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The Canin Bulletin
_ Singapore
VOL. 56 (Parts 1 & 2) September 2004 | ia ISSN 0374-7859
NATIONAL PARKS BOARD Ho ee at
Singapore Botanic Gardens Cluny Road Singapore 259569 Tel: 64741 165 Telefax: 64754295
THE GARDENS’ BULLETIN SINGAPORE
The Gardens’ Bulletin Singapore publishes papers on plant taxonomy (including
revisions), horticulture, phytogeography, floristics, morphology, anatomy and
related fields with emphasis on plants in the West Malesian region.
EDITORIAL BOARD
Dr R. Kiew Dr R.T. Corlett
(Editor) University of Hong Kong
Singapore Botanic Gardens Hong Kong
Dr P.Y. Tan Dr M.C. Roos
(Assist. Editor)
Singapore Botanic Gardens
Dr S.C. Chin
Singapore Botanic Gardens
Dr. M.J.E. Coode
Royal Botanic Gardens,
Kew, U.K.
National Herbarium of the Netherlands
Leiden University,
Leiden, The Netherlands
Dr E. Soepadmo
Forest Research Institute Malaysia
Kepong, Malaysia
Dr W.K. Tan
National Parks Board, Singapore
The Gardens’ Bulletin is published twice yearly by the National Parks Board,
Singapore. Neither the National Parks Board nor the Editorial Board is responsible
for the opinions or conclusion expressed by the contributing authors.
The annual subscription for the Gardens’ Bulletin is Singapore $100.00
including postage. Overseas subscribers are required to make payment in the form
of bank drafts or international money orders in Singapore currency payable to
National Parks Board, Singapore.
Instructions for contributing authors are found on the inside back cover.
The Gukdens’ Bulletin
Singapore
Vol. 56 (Parts 1 & 2) September 2004 ISSN 0374-7859
CONTENTS
Zumbroich, T.J. and Chay Y.W.
Singapore’s first spice plantation and botanic garden 1819-1859 ...... l
Duistermaat, H.
New grass (Poaceae) records for Singapore,
MS Fe ci LAN MEW TOP ASIA wi... sectecscereoeecansecscsetersensenseeseseccocsnssonves 29
Skornickova, J., M. Sabu and M.G. Prasanthkumar
Curcuma mutabilis (Zingiberaceae): a new species from South India ............ 43
Zhu, H.
A tropical seasonal rain forest at its altitudinal and
Pitan Wits i southern Yunnan, SW CAIN «0.2.20. scccc cccccccoesseceesedsareessnees 35
Kiew, R. and Tan J.H.
Begonia sabahensis Kiew & J.H.Tan (Begoniaceae), a new
CMT Witt COTE LIN TSGIICO: «cance scnnsecncpsceten-ntacessqanesnsatcnectunsenancenrescess a3
Argent, G.C.G.
New species and comments on Rhododendron (Ericaceae)
POTN Rat ASLO ST PAV ARO OTD, FNS a caiporamessgninnss<sezuanerananeéansepurannasanoctnnaats fie,
Kiew, R.
Begonia sizemoreae Kiew (Begoniaceae),
tea i! PROT STOR VACANT oo. a nvonnennncenannsiorsrneannceconnensnreverosens ome
Pearce, K.G.
The vegetation and plants of Niah National Park. Borneo ................eeeees 101
Craven, Lyn A. and Siti Sunarti
A new Species of, and Reinstatements in, Octamyrtus (Myrtaceae) ............. 147
Ipor I.P., Tawan C.S. and Boyce P.C.
A new species of Amorphophallus (Araceae: Thomsonieae)
Spt bee ete 7 eh i on oa ee oh ee a 153
Poulsen, A.D. and Clausager K.E.
A new species of Stachyphrynium (Marantacaea) from Borneo.................++ 161
Date of publication : 25th October 2004
Published by
National Parks Board
Singapore Botanic Gardens
Cluny Road, Singapore 259569
Printed by KaroCraft
Digitized by the Internet Archive
in 2014
http://archive.org/details/gardensbulletins456unse
Gardens’ Bulletin Singapore 56 ( 2004 ) 1-28. I
Singapore’s First Spice Plantation and Botanic Garden
1819-1859
THOMAS J. ZUMBROICH
89 East ‘E’ Street, Encinitas, CA 92024, USA
AND
CHAY YUE WAH
Nanyang Technological University,
Nanyang Avenue, Singapore 639798
Abstract
The history of the spice plantation and Botanic Garden conceived by Sir Stamford Raffles
(1781-1826) for Government Hill (now Fort Canning Hill) in Singapore is documented from
its inception in 1819 using archival sources, contemporary accounts, as well as previously
published and unpublished maps. After Raffles’s departure from Singapore in 1823 and under
pressure for retrenchment, the patronage of economic botany shifted from the government to
entrepeneurs who revived Raffles’s original nutmeg (Myristica fragrans Houtt.) plantation in
the 1830’s, subsequently leading to frenzied nutmeg cultivation around the settlement. A
combination of competitive pressure, inadequate husbandry and an insect pest brought nutmeg
cultivation on Government Hill and elsewhere in Singapore to an end in 1859. Not only was
the spice plantation envisioned by Raffles in existence longer and at times more extensive
than previously thought, it also exerted a significant impact on land development in and around
the settlement. A map of Government Hill and the surrounding area is shown to illustrate the
extent of the Botanic Garden as originally planned, its approximate location in 1827 and the
gradual encroachment of other buildings and facilities over time.
Introduction
The powerful combination of George III (1738-1820) and Sir Joseph Banks
(1743-1820) created in late eighteenth-century Britain a spirit of agricultural
‘improvement’ that extended its reaches well beyond domestic agrarian reform.
The West Indies, after 1783 deprived of their connection to the American
mainland, saw the revival (e.g. St. Vincent) or founding (e.g. Grenada) of
Botanic Gardens to prepare for the acclimatization of new Asian plants as
2 Gardens’ Bulletin Singapore 56 (2004)
much needed staple foods or crops. In Asia itself, the East India Company
was faced with millions of pounds of debt and needed to seek new measures to
finance its China trade. Economic botany, in particular the cultivation of
commodities of value in Europe or China, it was hoped, would come to the
rescue of the East India Company’s trade balance (Drayton, 2000, pp. 85-
128).
When in 1786 Lieutenant-Colonel Robert Kyd (1746-1793) first
approached the Board of Directors with the suggestion for a Company
plantantion in Calcutta in order to transfer novel food plants to Bengal as a
measure against a future famine, his arguments were of a moral and political
nature (Kyd to Board of Dir., 15 April 1786, in Biswas, 1950). Subsequently,
he withdrew to more traditional mercantile reasoning and detailed the financial
benefits the Company could derive from undermining the profitable Dutch
monopoly for spices. He envisioned a Botanic Garden ‘not for the purpose of
collecting rare plants ... as things of mere curiosity’, but rather to provide a
depository for plants that would ‘tend to the extension of the national Commerce
and Riches’ (Kyd to Board of Dir., 1 June 1786, in Biswas, 1950; Chatterjee,
1948, p. 362; Shukla, 1994, pp. 28-30). As the cultivation of the “finer spices’,
nutmeg (Myristica fragrans Houtt.) and its derivative mace, cloves (Syzygium
aromaticum (L.) Merr. & Perry) and cinnamon (Cinnamomum verum Presl),
had so far and contrary to French successes, eluded the East India Company,
the Court of Directors seized upon this opportunity by endorsing Kyd’s plan
in July 1787. On a site of 310 acres along the Hooghly river, Kyd then
introduced about 300 species of commercially significant plants into the new
Calcutta Botanic Garden.
Early Spice Cultivation in Penang and Bencoolen -
The British occupation of the Moluccas from 1796 to 1803 for the first time
allowed unhindered access to the spice gardens of the Banda islands and Ambon
(Wright, 1958, p. 27-47), and Kyd’s successor, Dr William Roxburgh (1751-—
1815), immediately sent the botanist Christopher Smith (d. 1807) to organise
the transportation of large amounts of nutmeg and clove plants to areas under
the Company’s control. In the years 1800-1801 over 30,000 nutmeg seedlings
and plants and 31,000 clove seeds and plants were dispatched to Penang. The
larger spice plants, some as tall as 14 ft and over 15 years old, typically perished
during the journey, but not so the younger plant material (Leith to Government
(Gov.), 16 July 1800, in Braddell, 1851; Penang to Calcutta, 21 April 1802, in
Braddell, 1851; Hunter to Sec. to Gov., 1 July 1802,.in Braddell, 1851;
Warburg, 1897a, pp. 244-246). The spice plants were cultivated in an extensive
government-run spice plantation at Ayer Hitam, where, however, due to the
Singapore's First Spice Plantation and Botanic Garden 3
excessive inventory and ill-guided care the survival rate was low. When in
1805 the annual expenses for the plantations in Penang reached $11,909, the
Calcutta government ordered that they be disposed of, and Lieutenant-Governor
Robert Townsend Farquhar (1776-1830) did so by selling the surviving 7,800
spice plants to European individuals for a total of $9,656 (Farquhar to incoming
Gov., 1805, in Braddell, 1851; Penang to Gov., 12 November 1805, in Braddell,
1852). Spanish dollars ($) were worth about 5 shillings sterling in the early
19" century. Except for a feeble and ultimately abortive attempt at establishing
another Company Garden between 1822 and 1834, the further development of
the spice industry of Penang was left entirely to private initiative (Low, 1836,
p. 20; Warburg, 1897a, p. 246; Jackson, 1968, table 7, p. 105).
In Bencoolen, the British settlement in West Sumatra, fruitless attempts
had been made since the early 1770’s to procure spice plants, until in 1798 a
first shipment of about 850 nutmeg and 70 clove plants arrived (Warburg,
1897a, pp. 226-228; Bastin, 1965, pp. xxxii—xxxii1). Since the initial progress
of the nutmeg plants was encouraging, a much larger shipment from the
Moluccas followed in 1803, and, despite the Court of Directors’ fiscal concerns,
the local authorities at Fort Marlborough in Bencoolen decided to cultivate
spices in a significant Company-owned nursery, where by 1809 about 3,200
nutmeg trees and 200 clove trees thrived (Fort Marlborough to Court of Dir., 7
March 1804, in Bastin, 1965). However, the second British occupation of the
Moluccas from 1810-1817 and the ensuing excessive imports brought about a
glut of spices on the London market, and when Sir Stamford Raffles (1781-
1826) was dispatched to his post as Lieutenant-Governor of Fort Marlborough
in October 1817, he had clear instructions to dispose of the Company’s spice
plantation there. Unlike Robert Farquhar, who had been critized for his fire
sale of the government plantation in Penang, Raffles resisted the immediate
execution of his order and only gradually sold off the stock in the government
plantation to locals planters whose spice cultivation he steadfastly encouraged
(Remarks on Farquhar’s Report by Governor in Council, 20 February 1806, in
Braddell, 1851; Raffles to Court of Dir., 18 April 1818, in Bastin, 1965).
In late 1818, Raffles stayed in Calcutta in order to lobby for his plan of
permanently securing the British trade route through the Straits of Malacca
and thus had the opportunity to explore the Calcutta Botanic Garden where he
became acquainted with the Danish surgeon and botanist Dr Nathaniel Wallich
(1786-1854). In 1809, Roxburgh’s support had helped Wallich to be employed
by the East India Company as a collector ‘to travel about in search of the
unknown objects of natural history’ (Roxburgh to Gov., 24 February 1809, in
Bastin, 1981), and in 1817 Wallich was appointed permanent Superintendent
of the Calcutta Garden. Wallich introduced Raffles to Dr William Jack (1795—
1822), a physician and outstanding systematic botanist, staying at the Garden
to pursue his botanical interests. Raffles immediately engaged Jack as his
4 ' Gardens’ Bulletin Singapore 56 (2004)
naturalist. Jack was to replace Raffles’s previous naturalist Dr Joseph Arnold
(1782-1818) who had died in July 1818, shortly after discovering with Raffles
and Lady Raffles the parasitic plant that would be named Rafflesia arnoldii
R.Br. (Burkill, 1916, p. 149; van Steenis-Kruseman, 1950, pp. 23-24; Bastin,
1990, p. 12).
First Spice Plantation in Singapore, 1819-1822
When in January 1819, Raffles and Major William Farquhar (1770-1839)
settled on Singapore island for a new British trading post, they found a Malay
fishing village surrounded by lush primary forest with ample fresh water
supplies. Even during those first days spent on Singapore island, Raffles took
time to study the vegetation and engage in some botanising (Raffles to Marsden,
31 January 1819, in Raffles, 1830). Raffles returned to Singapore a second
time in May 1819 for about three weeks to instruct the new Resident Farquhar
on numerous administrative issues, and this time he brought along a group of
naturalists comprised of Jack and two Frenchmen, Alfred Duvaucel (1793-—
1863) and Pierre Diard (1795-1863) who later became Director of the Botanic
Garden at Buitenzorg (now Bogor) in Java (Treub, 1893, p. 34; van Steenis-
Kruseman, 1950, p. 136). After exploring the island, Jack was overwhelmed
with the abundance of the vegetation, reporting to Wallich:
‘Flora here luxuriates in endless varieties, where she finds soil, climate
and everything congenial. I find many, or most of my Penang acquaintances
with others surpassing them in magnificence’ (Jack to Wallich, 8 June 1819,
in Burkill, 1916).
If the luxuriant growth of tropical vegetation was not enough to raise
Raffles’s hopes for successful plantations on the island, there was also evidence
of past and ongoing agriculture around what was known to the local Malays as
Bukit Larangan (Forbidden Hill). This cone-shaped eminence, located between
the Singapore River and a fresh water stream (Bras Basah River), in March
1819 had undergone some clearing (Hill, 1970, p. 146; ibid., p. 168) whereby
workers came upon an abandoned orchard on the apparently fertile hillside. A
few years later this orchard was still being recognized as a remnant of
Singapore’s past:
‘It is remarkable, that many of the fruit-trees cultivated by the ancient
inhabitants of Singapore are still existing, on the eastern side of the hill,
after a supposed lapse of near six hundred years. Here we find the durian,
the rambutan, the duku, the shaddock, and other fruit-trees of great size’ (4
February 1822; Crawfurd, 1828, p. 47).
Lacking other historical or physical evidence, the ctl age and origin
of the trees cannot now be ascertained, but the presence of fruit trees was
rightly taken as evidence of previous arboriculture on the hillside. Early British
Singapore’s First Spice Plantation and Botanic Garden 5
observers also found established plantations in the hills surrounding the mouth
of the Singapore River where, in exchange for rent, Temenggong Abdul Rahman
(d. 1825) had allowed various Malays and Chinese to establish about 20 gambier
(Uncaria gambier Roxb.) plantations prior to the arrival of the British (Jack to
Wallich, 8 June 1819, in Burkill, 1916; Singapore Straits Records (SSR, National
Archives, Singapore) L11, pp. 139-141, Farquhar to Sec. to Lieutenant
Governor, 23 December 1822; Bartley, 1933, p.117; Jackson, 1968, pp. 7-8).
This was recognized in the treaty of 26 June 1819 between the British and the
Temenggong, in which he, as the proprietor of the soil, was given continuing
and exclusive control over the disposal of land for gardens and plantations,
while the boundaries of the land under British control were restricted to a
stretch of sea front of about six miles long and landwards to the range of a
cannon shot (SSR, L17, p. 57A, Sec. to Lieutenant Governor to Farquhar, 4
February 1823; Buckley, 1902, pp. 58-59; Wake, 1975, pp. 60-61). Before
his departure, Raffles decreed as part of his instructions for the town planning
that the whole of the hill, subsequently known as ‘Government Hill’, should
be reserved for the ‘exclusive accomodation of the Chief Authority’ and that a
bungalow later be added on top as a residence (SSR, L10, pp. 71-75, Raffles to
Farquhar, 25 June 1819). No written orders to Farquhar were included for the
location of the planned spice plantation, though he would have received at
least oral directions from Raffles. Cross-referencing later documents allows
us to place the first spice plantation on the more gradual lower eastern slopes
of the hill, close to the later Government House, and not too far from the site
of previous awricultural activities. This particular location would prove well-
suited to the cultivation of nutmegs (Wallich to Raffles, 21 November 1822, in
Hanitsch, 1913; Pearson, 1953, pp. 200-204).
After 1818, the nurseries of the private and public spice plantations near
Fort Marlborough in Bencoolen contained such a tremendous supply that many
plants perished from neglect (Lumsdaine, 1821). Upon returning to Bencoolen
in July 1819, Raffles was therefore quickly able to assemble a collection of
1000 nutmeg and 350 clove seeds as well as 100 seedlings and 25 large plant
specimens each of both species, and on 18 August this shipment was dispatched
to Singapore on the Jndiana (Captain J. Pearl). Raffles explained to his superiors
that the purpose of the endeavour was ‘to extend the cultivation of the nutmeg
and clove tree as much as possible’ advising Farquhar ‘to exert [his] utmost
endeavours to establish the cultivation under [his] immediate authority’ (SSR,
L10, p. 125, Raffles to Farquhar, 18 August 1819; Raffles to Gov., 26 August
1819, in Bastin, 1965). The plant material arrived in Singapore on 27 September
1819 in the care of M.W. Dunn. The latter was apparently not so much a
gardener, but a long-time employee of the Company in Batavia (today’s Jakarta)
and Bangka sent by Raffles for general assistance to the administration in
Singapore where Dunn can be traced till July 1821 as an occupant of land
6 Gardens’ Bulletin Singapore 56 (2004)
(SSR, L10, p. 179, Farquhar to Raffles, 27 October 1819; Braddell, 1855, p.
451; Kathirithamby-Wells, 1969, p. 51).
Farquhar himself was known to have a long-standing interest in botany.
During his tenure as Resident of Malacca, 1803-1818, he collected plants on
Mount Ophir and had Chinese artists prepare a large collection of botanical
drawings (Jack to Wallich, 14 January 1819, in Hanitsch, 1916; Goh, 1999).
Farquhar soon reported back from Singapore that the larger plants had been
established in their permanent location and that seeds and seedlings were in a
nursery, all ‘in a thriving state’ ’(SSR, L10, pp. 182-183, Farquhar to Raffles,
28 October 1819). Farquhar’s initiative to employ Mr Brooks, a European
gardener, at $40 a month as Botanist and Superintendent for the spice plantation
on Government Hill, drew an immediate rebuke from Raffles. Raffles cited
orders of the Supreme Government to prune expenses in Singapore and
abolished the post of Brooks in December 1819 (SSR, L10, pp. 239-243, Raffles
to Farquhar, 29 December 1819, enclosing letter from Governor General, 15
October 1819; Kathirithamby-Wells, 1969, p. 51). The spice plantation was
thereafter under the direction of Farquhar with no further guidance from Raffles
until he eventually returned to Singapore.
Raffles’s extreme disdain for the Dutch helped motivate his own zealous
pursuit of spices. He resented the return of the ‘fertile and important Islands
of Java and the Moluccas’ to the Dutch, and one of his declared objectives in
founding Singapore was to ‘eventually destroy the spell of Dutch monopoly’
(Raffles to Col. Addenbrooke, 10 June 1819, in Willians, 1878). Fostering the
cultivation of spices to this end was, however, at least a ten-year project and
reveals Raffles’s long-term hopes for the British foothold at a time when the
future of Singapore was still caught up in a ‘paper war’ (Wurtzburg, 1954, pp.
525-542). Besides sending spice plants to Singapore, Raffles continued to
entice planters in Bencoolen to increase nutmeg production. In Penang in
1818, 79,000 nutmeg and 104,000 clove trees had been counted, and harvests
were beginning to swell during the 1820’s (Penang Consultations, 7 October
1818, in Jackson, 1968). Raffles grossly overestimated both the future demand
for spices in Great Britain and the price they would command. In 1822, the
Company’s London warehouses had about 1,000,000 Ibs of nutmeg and 200,000
Ibs of mace in stock, and further imports of spices from Bencoolen could only
be sold, if at all, at great losses (Bastin, 1960, p. xxxvii). Hence, Raffles
agricultural endeavours in Bencoolen proved financially disastrous for the
Company and did little to inspire official support for Raffles’s new agricultural
experiments in Singapore.
In 1819, Raffles himself also imported to Singapore cotton (Gossypium
sp.) seeds of the pernambuco variety, probably from Penang, and the plants
were successfully raised on Government Hill. While claims of cacao trees
(Theobroma cacao L.) and cultivated orchids thriving nearby are apocryphal
Singapore’s First Spice Plantation and Botanic Garden 7
(Ridley, 1905, p. 296; ibid., p. 303; Knecht, 2000, p. 194), there is evidence
that tea (Camellia sinensis (L.) Kuntze) was flourishing in the Garden by 1822,
though nutmeg and clove trees would remain its dominant features (Wallich to
Raffles, 2 November 1822, in Hanitsch, 1913; Jack to Wallich, 15 March
1819, in Burkill, 1916).
By May 1821, access to the spice plantation was improved with small
roads over the top and around Government Hill, and an additional $80 had
been spent on a bungalow, presumably a small structure to house a caretaker
and tools (Buckley, 1902, pp. 68-69; Bastin, 1960, pp. 146-147). During the
same time period, Chinese gambier and pepper (Piper nigrum L.) plantations
continued to slowly expand inland (Finlayson, 1826, pp. 61-62), but the unclear
future of Singapore and uncertain land rights caused most of the 22 Europeans
that occupied land in 1821 to abstain from engaging in agricultural ventures
(Braddell, 1855, p. 451; Miller, 1941, p. 194). A notable exception and case
in point regarding the issue of land tenure was Farquhar himself who started a
‘Garden & Plantation’ at his own expense on a hill north of the settlement.
Despite his insistence that he had received sanction from the Temenggong
according to the treaty of June 1819, his claim on the land was later denied by
Raffles (SSR, L11, pp. 139-141, Farquhar to Sec. to Lieutenant Governor, 23
December 1822).
Plans for a Botanic and Experimental Garden, 1822-1823
Raffles returned from Bencoolen to Singapore on 10 October 1822 and, out of
concern about the haphazard growth of the settlement, he mounted a final
effort to shape its design and character to match his vision for a ‘civilized’
society. By coincidence, in early September 1822 Wallich had arrived in
Singapore from Calcutta on his way to China to recover from malaria. He
decided to remain on the island for the rest of his leave, ultimately taking with
him ‘a curious and extensive botanical collection’ as the result of his botanising
in Singapore (Crawfurd, 1828, p. 297; Wallich to Secretary to Gov., 19 July
1822, in Hanitsch, 1913).
Wallich’s friendship with Raffles and Lady Sophia Raffles (1786-1858)
led to frequent social meetings and shared excursions over the course of six
weeks. They agreed to significantly expand the existing ‘Government Garden’
into a larger ‘Botanic and Experimental Garden’. The timing of their plan was
not accidental: Under the guidance of Raffles, a “Land Allotment Committee’,
in which Wallich participated, and a “Town Planning Committee’ were formed
to advise on developing a detailed masterplan for the settlement. Dr John
Lumsdaine, Superintendent of the Company’s Spice Plantation at Fort
Marlborough and an expert in the cultivation of spices, had accompanied Raffles
to Singapore and joined Wallich on the first committee. The actual map of the
8 Gardens’ Bulletin Singapore 56 (2004)
town was to be drawn up by Lieutenant Philip Jackson (1802-1879), the
Assistant Settlement Engineer (Report of the Committee to Raffles, 23 October
1822, in Hanitsch, 1913; Pearson, 1953, pp. 200-204; Wurtzburg, 1954, pp.
608-610; Bastin, 1981, p. 49). In the course of this town planning process,
and no doubt with much input from Raffles himself, on 2 November 1822
Wallich formally suggested to Raffles to include an extended garden into the
redesigned town plan:
‘... I beg leave to recommend that a suitable piece of ground may be
appropriated in the neighbourhood of the European town for the purposes
of a botanic garden and for the experimental cultivation of the indigenous
plants of Singapore and the adjacent Islands, as well as of such others of
foreign growth, as it might be desirable to submit to a skilful trial, previous
to encouraging their general introduction’.
In Wallich’s opinion Singapore’s location was ideal for such an institution:
‘... the most favourable for indigenous as well as foreign vegetation and
forming part of the richest archipelago in the world - its soil yielding to
none in fertility, its climate not exceeded by any in uniformity, mildness
and salubrity’ (Wallich to Raffles, 2 November 1822, in Hanitsch, 1913;
SSR, LY, pp. 91-100, 1822).
As an indication of the commercial promise of Singapore’s agriculture,
Wallich pointed to a number of wild nutmeg species found in the surrounding
forests, Farquhar’s successful experiments with spices and pernambuco cotton
in the existing garden, as well as the local pepper and gambier production. He
closed by expressing his hopes of becoming the Superintendent of this Garden,
which he planned to staff with a number of apprentices and experienced
gardeners from Calcutta. Raffles’s and Wallich’s letter was carefully designed
to aid their bid for financial support from the East India Company, hence it
overstated the bright prospects of economic botany in Singapore. Wallich
had, indeed, collected at least six species of wild nutmegs on the island, one of
which he named in Farquhar’s honour Myristica farquhariana (now known as
Gymnacranthera farquhariana (Wall. ex Hook. f. & Thomson) Warb. var.
farquhariana; Warburg 1897b: p. 367; de Wilde, 2000, pp. 4446), but none
of those species had the slightest aromatic quality. Over the years, some other
of Wallich’s supporting arguments also proved to be spurious, for example,
the lack of seasonality had a deleterious effect on some crops, like cotton, and
made the cultivation of others, like coffee (Coffea arabica L.), costly by
extending the harvest over a long period (Thomson, 1850, p. 141; Crane,
1851, p. 122).
The planning for the Botanic Garden was cut short when suddenly an
opportunity arose for Wallich to catch a return passage to Calcutta (Wurtzburg,
1954, p. 614). On 21 November, one day before his embarkation, Wallich
outlined to Raffles a plot for the Garden, which was soon after surveyed by
Jackson, the planner. An area of 48 acres on the northeastern side of
Singapore's First Spice Plantation and Botanic Garden 9
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Figure 1. This ‘Plan of the Town of Singapore by Lieut. Jackson’ represents the
proposed lay-out of the town as of December 1822 (Crawfurd, 1828, opposite
p. 529; Pearson, 1953, pp. 200-204) with a revised scale added.
The road along the ‘Fresh water Rivulet’ (Bras Basah River) corresponds
to Hospital Street, now Stamford Road, and Selegy Road was soon renamed
Brass Bassa Road (now Bras Basah Road). The area east of Hill Street, originally
reserved for church, government offices and court house as shown in this lay-
out, accomodated the convict lines after 1825. The total area of the planned
‘Botanical and Experimental Garden’ within the borders of the roads as shown
here is 48 acres. The dotted area of ei ght acres marks the approximate extent of
the Government Garden under Montgomerie’s superintendency in 1827 and
includes the original plantings by Farquhar. The calculation of the planted area
is conservatively based on a distance of thirty feet between trees, equivalent to
fifty trees per acre (Low, 1836, pp. 28-30: Ridley, 1912, p. 445).
(A) The cemetery opened in 1822 and was extended across the road from town
in the late 1840’s.
(B) The Armenian Church was erected in 1835 in proximity to the location of
the first home of the merchant A. L. Johnston. By 1840 Armenian Street
ran parallel to Hill Street on the lower slopes of Government Hill with
houses encroaching on the spice plantation.
(C) After 1830 the area (between the current Bencoolen and Waterloo Streets)
was used for different hospitals, hence the early name Hospital Street for
Stamford Road.
(D) A new convict jail was erected by convicts in 1841 and expanded considerably
between Hospital and Brass Bassa Road over the next fifteen years.
(E) The location of the Catholic Church of the Good Shepherd since 1845.
10 Gardens’ Bulletin Singapore 56 (2004)
Government Hill including the Fresh Water Stream and some low ground were
to be enclosed appropriately as the new ‘Singapore botanic and experimental
institution’ (Fig. 1). Furthermore, Wallich sweepingly proposed that all of
Government Hill should be landscaped with indigenous and non-indigenous
ornamental trees and shrubs by the Superintendent of the Garden so as to
complement the Botanic Garden (Wallich to Raffles, 21 November 1822, in
Hanitsch, 1913). Raffles, not one to let financial considerations stand between
him and his wishes (Hill, 1970, p. 78), donated $1,000 for the layout and
enclosure of the Garden. He also ordered an initial public subsidy of $60 a
month, less than 2% of the total budget for Singapore at the time and just
sufficient for about ten workers and some tools and materials (Braddell, 1853,
p. 347; Raffles to Wallich, 15 November 1822, in Bastin, 1981). In addition,
Raffles sent a formal letter, complete with supporting enclosures by Wallich,
to the Supreme Government in Calcutta to garner recognition and funding for
his project (Raffles to Wallich, November 1822, in Bastin, 1981; Raffles to
Wallich, 17 April 1823, in Bastin, 1981).
By early January 1823, Raffles and his wife had moved into a new
bungalow at the top of Government Hill overlooking the settlement. Despite
his serious and recurrent health problems, he was now anxious to begin the
design of his ‘future Superstructure’ by expanding the existing Botanic Garden
and turning the remainder of the Hill into a ‘very pretty Park’. He even planned
for an enclosure with 200 Spotted Deer (Axis axis Erxleben), though he soon
scaled back and abandoned the idea, recognizing that his successor would not
look favourably upon it (Raffles to Wallich, 5 January 1823, in Bastin, 1981;
Raffles to Marsden, 21 January 1823, in Raffles, 1830). Without official
approval from Calcutta for the project, Raffles prepared for the start of
construction: He gave orders to evict locals residing within the area of the
planned Botanic Garden and to compensate them for their dwellings and
surrounding cultivated areas (SSR, L13, p. 14, Farquhar to Asst. to Police Dept.
Bernard, 11 January 1823). Engineer Jackson began building a wall made
from a combination of brick and wood around the part of the Botanic Garden
facing the town, and workers began cutting a number of ‘noble’ walks through
the grounds (SSR, L13, p. 3, Farquhar to Jackson, 6 January 1823; SSR, N1, p.
103, Montgomerie to Prince, 8 February 1827). Raffles’s grand vision of
Government House overlooking a magnificently landscaped park and Botanic
Garden appears modelled after the private estates of the influential and wealthy
members of Penang’s British community, such as Governor William Edward
Phillips’s (b. 1769) Suffolk House or David Brown’s (d. 1825) Glugor House
(Crawfurd, 1828, p. 10; Stevens, 1929, pp. 405-411). For Singapore, Raffles’s
plan for his Garden was way out of proportion with the simple Government
Residence it was to surround, and, more importantly, would have required
substantial public financing.
Singapore’s First Spice Plantation and Botanic Garden 11
Montgomerie’s Superintendency of the Garden, 1823-1827
Initially, Wallich had planned to return to Singapore soon and was anxious to
secure a land grant from Raffles for a future home on what would be known as
Mount Wallich (Raffles to Wallich, 17 April 1823, in Bastin, 1981). But before
committing any resources to the development of the Garden, Wallich awaited
the Supreme Government’s decision on the project including his confirmation
as Superintendent in Singapore. As the endorsement failed to materialise, he
completely withdrew his support and, despite Raffles’s pleas for help (‘- pray
send down something like a Gardener or Head Man’), no personnel from
Calcutta ever arrived (Raffles to Wallich, 8 March 1823, in Bastin, 1981).
Thus, when Raffles made his final departure from Singapore in June 1823, the
nascent Garden lost its most ardent and, by then, only supporter.
The incoming Resident, John Crawfurd (1783-1868), drew his own
conclusions on Singapore’s agricultural capabilities from his study of
‘agricultural geology’ (Khoo, 1996, pp. 61—70). The absence of ‘alluvial plains’
and sufficient ‘rich black mold’ on the one hand and the presence of the “poor
red soil of the hills’ on the other hand made Singapore ill-suited for raising
most tropical crops of commercial importance (Crawfurd, 1828, p. 534;
Crawfurd, 1849, pp. 508-509). The relatively poor soil of Singapore island
had been noticed as early as the 1330’s by the travelling Chinese trader Wang
Ta Yuan in his Tao-i Chih-lioh who remarked about the settlement surrounding
Government Hill: “The soil is poor and grain scarce’ (Wheatley, 1961, p. 83).
Crawfurd’s dire predictions for the cultivation of the finer spices were
based on his calculations of the inherent value versus the actual price obtained
for spices. He became convinced that competitive pressure from the Moluccas
would eventually erode profits for the British planters if ever the Dutch restored
free culture and the artificially high prices were to disappear (Crawfurd, 1820,
vol. III, p. 409). Declining prices did, indeed, become a challenging issues for
the spice planters around the middle of the nineteenth century, and Crawfurd’s
cautious assessment of the island’s agricultural potential would turn out to be
more accurate than Raffles’s and Wallich’s glowing endorsements. To make
matters worse for the future of the Garden, in February 1823, Crawfurd had
bitterly fallen out with Wallich while both were in Calcutta, at the time when
Raffles was lobbying the authorities for support of the project (Raffles to
Wallich, 8 March 1823, in Bastin, 1981). It was no surprise then that Crawfurd
gave no further support to the Garden than the stipulated monthly allowance
and polite appreciation for the work of the new Superintendent of the Gardens,
Dr William Montgomerie (1797-1856; Crawfurd, 1828, p. 534).
Montgomerie had been in Singapore since May 1819 as Assistant Surgeon
in general medical charge of the troops and from January 1823, after the
dismissal of his colleague Dr Thomas Prendergast, was alone in medical charge
; of the settlement (Makepeace et al., 1921, vol. I, p. 517; Pearson, 1955, pp.
12 Gardens’ Bulletin Singapore 56 (2004)
41-46). In the course of his own rushed departure, Wallich had suggested that
Montgomerie, known for his interest in natural history, should temporarily
oversee the Garden - though without any addition in pay ‘as the thing was
upon a small scale’ (Braddell, 1855, p. 62). Montgomerie, initially reluctant,
was put to work willy nilly ‘nolens volens’ under Raffles’s direct supervision,
but after Raffles’s departure, Montgomerie found himself left entirely to his
own devices since Wallich never contacted him again nor sent the promised
botanical expert (Raffles to Wallich, 5 January 1823, in Bastin, 1981).
This mirrors the experience of the botanist George Porter (d. 1833) who
in 1822 on Wallich’s recommendation was put in charge of the new Government
Botanic Garden in Penang, but he, too, waited in vain for the expected support
from Calcutta (SSR, A19, p. 386, Establishment of a Botanic Garden at Penang,
1823; Bastin, 1981, pp. 43-44). In fact, during the period between Wallich’s
departure from Singapore and 1830, not a single botanical collector brought
his expertise to the settlement (van Steenis-Kruseman, 1950, p. LXXXIV).
Hence Montgomerie lacked the inspiration and means to establish a botanical
collection, but instead concentrated on the development of the experimental
spice plantation, marking the start of his enthusiastic involvement in plantation
agriculture.
Montgomerie completed the enclosure of this plantation with a bamboo
fence on the hill side and improved the roads in the area. He also attempted in
vain to drain some of the low ground along the Bras Basah River which was
prone to flooding and remained unsuitable for the cultivation of spices. As
late as 1841, when a permanent convict jail was erected on the northern side of
the Bras Basah River, the marsh had to be reclaimed by raising the site by 2—
4 ft with soil from Government Hill (McNair, 1899, pp. 147-148). In addition
to the nutmeg and clove trees already planted by Farquhar, he continued to
procure seeds, raise them in a nursery and set out seedlings on 18 foot-wide
terraces on the lower eastern slopes of Government Hill. When in April 1825,
the first 200 Indian convicts were transferred from Bencoolen to Singapore
and accomodated in sheds built between Hill Street and North Bridge Road,
1.e., just south-east of the spice plantation, Montgomerie was able to supplement
his small work force with a few extra convict labourers, some even with working
experience from the spice plantations of Bencoolen (Jannings et al., 1821;
Raffles, 1830, fold-in plate, facing p. 525; McNair, 1899, pp. 38-39). .
The new Resident Councillor Prince, in charge of the town from 1826 to
1828, was far more sympathetic towards the cause of botany than Crawfurd:
Prince himself made botanical collections and re-established communications
with Wallich by sending him living specimens to Calcutta (van Steenis-
Kruseman, 1950, pp. 416-417; Wurtzburg, 1950, p. 141). In February 1827,
Montgomerie reported to Prince that a total of about 300 nutmeg and 100
clove trees were growing on the slope of Government Hill. The first small
crop of nutmegs had been harvested in 1826 and from selected fruits 200
Singapore's First Spice Plantation and Botanic Garden 13
seedlings were being propagated in a nursery that held a total of 300 nutmeg
and 200 clove plantlets (SSR, N1, pp. 101-106, Montgomerie to Prince, Report
upon the present State of the Honorable Company’s Botanical and Experimental
Garden at Singapore, 8 February 1827). An estimated eight acres plus the
nursery, i.e. a fraction of the plot originally allocated in 1822, was being
cultivated with nutmegs and cloves (Fig. 1, dotted area). Despite
Montgomerie’s claim that the spices ‘appeared to thrive uncommonly well’
(SSR, N1, p. 102, Montgomerie to Prince, 8 February 1827), this was only
true for the nutmeg trees, since the clove trees, some in their ninth year, still
showed no sign of bearing (Ridley, 1912, p. 176; Bastin, 1960, p. 192). Stull,
Montgomerie suggested that the government would eventually profit if it
developed spice plantations with convict labour and leased them out once
they approached bearing - a plan that was both ill-timed and unrealistic.
The Anglo-Dutch treaty of London of March 1824 that had ceded
Bencoolen and upheld the Dutch trade monopoly in the Moluccas, raised largely
unwarranted fears that the Dutch hold on the spice trade would tighten again.
While the treaty caused ruin to the British nutmeg farmers in Bencoolen, giving
up the costly dependency was a financial blessing for the Company. In
parliament, Foreign Secretary George Canning (1770-1827) used the news
about successful experiments with spices in Singapore to assure critics of the
treaty that Britain’s spice supply would not be endangered in the future (Wright,
1958, p. 101; Newbold, 1839, p. 271). This promise, however, had no bearing
on any further investment into spice cultivation by the Company whose prime
concern for the United Presidency Administration of Penang, Malakka and
Singapore was retrenchment (Kathirithamby-Wells, 1969, p. 53). Lacking
sufficient funding, Resident Councillor Prince could do little more to encourage
private involvement in spice cultivation than to offer the limited supply of
nutmeg and clove seedlings available in Singapore. In 1827 he directed
Montgomerie to increase production of seedlings in the nursery and also
approached the Governor in Penang with a request for the shipment of young
plants or even just germinating seeds since ‘many of the European Land holders
... [were] now preparing their Lands for the reception of spices’ (SSR, N1, p.
107, Resident’s diary, 9/10 February 1827; Prince to Sec. Gov. Penang, 15
February 1827, in Cowan, 1950).
During the same year a period of decline began for the Garden as
Montgomerie left Singapore with the Bengal troops for India and financial
support of the Garden was limited to a staff allowance of 20 rupees (Makepeace
et al., 1921, vol. I, p. 490; ibid, vol. I, p. 517). ASicca rupee was worth about
2 shillings sterling. The supervision was first put into the hands of a professional
English gardener turned artilleryman with the newly arrived Madras troops,
later taken on by a Mr Caswall in charge of Medical Garrison Staff (SSR, V2,
pp. 307-308, Fullerton and Prince to Court of Dir., June 1828). Upon receiving
a report that the Garden was “by no means in good order and very confined’
-
14 . Gardens’ Bulletin Singapore 56 (2004)
the Governor-General ordered that the establishment be discontinued. Effective
30 June 1829, all financial support was cut, and the grounds from then on left
under the exclusive care of ten convicts (SSR, N6, p. 160, Act. Resident
Presgrave to Supt. of Convicts Bonham, 30 June 1829).
Despite its limited scale, the government plantation had in principle
demonstrated the possibility of growing nutmegs on the island by bringing the
first nutmeg trees to fruiting, and thus inspired other residents to slowly follow
suit with nutmeg cultivation. However, soil analyses performed by Major
James Low (1791-1852) during his time as Assistant Resident in 1840-1841,
revealed that the soil on the slopes of Government Hill was particularly fertile
and suited to the cultivation of nutmegs (Singapore Free Press, 25 November
1841). Therefore the initial success had exaggerated the commercial
opportunities for nutmegs growing in other locations in Singapore on inferior
soils.
The government plantation also presented a ready source for young
plantlets. As the viability of nutmeg seeds falls rapidly within a week, but
they self-seed easily, young plants could be dug up around the existing trees
and transplanted from there. In this respect, the plantation fulfilled a continuing
role by providing propagation material, at times supplemented from Penang,
for most of the other planters in Singapore (Buckley, 1902, p. 198). Even the
less successful cloves in the government plantation continued to attract attention
until about 1840, when it was eventually recognized that their best use in
Singapore was as an ornamental tree (Belcher, 1848, vol. II, p. 381; Thomson,
1850, pp. 102-103).
Montgomerie had started out as an amateur, unfamiliar with agriculture
under equatorial conditions and drew on the experience of the British spice
planters from Bencoolen and Penang. He had at his disposal Lumsdaine’s
often reprinted “Report on the Cultivation of Spices at Bencoolen, 1819-20’
that became the standard manual for British nutmeg planters for years to come
(Lumsdaine, 1821; Singapore Chronicle 15 May 1834; Low, 1836, pp. 28—
30; Lumsdaine, 1851, pp. 78-84; Ferguson, 1889, pp. 102-108). Montgomerie
also had the opportunity to receive direct advice from Lumsdaine when the
latter visited Singapore with Raffles in 1822-1823. Consequently,
Montgomerie adopted in Singapore Lumsdaine’s practice of growing nutmeg
trees without the permanent shelter of shade trees. While on the Banda Islands
trees were grown in alleys beneath tall kanari trees (Canarium vulgare Leenh.),
in the British plantations only newly-planted trees were protected with
individual shade structures made of atap (fronds of the mangrove palm, Nypa
fruticans Wurmb.). The slow growth of the shade trees, the terraced hill sides
of the plantations and the scarcity of fertile soil prevented protective shade
trees from being considered a useful practice in the British plantations. In the
long-run this decision had crucial consequences for Singapore’s nutmeg
cultivation: Not only did the open design of plantations promote excessive
Singapore’s First Spice Plantation and Botanic Garden 5
growth of invasive lalang grass (Imperata cylindrica (L.) Raensch.), thus
requiring frequent expensive weeding, it also made the trees more susceptible
to the insect pest that later devastated the spice plantations.
Land Development around the Botanic Garden, 1830s to 1840s
Parts of the large area that had originally been designated for the Botanic
Garden gradually began to be utilized for other purposes. A new Christian
cemetery had been opened in 1822 on the upper slopes of Government Hill to
take the place of the very first cemetery that was located inappropriately close
to Government House. This new cemetery gradually expanded downhill
towards the Spice Garden, but did not approach planted areas until the late
1840s (Fig. 1, A; Stallwood, 1912, plate I). In the direction of town, the spice
plantation early on bordered the gardens of the houses along Hill Street on the
lower slopes of Government Hill (Pearson, 1955, frontispiece; ibid., p. 67). In
April 1833, the Armenian community petitioned the Resident Councillor for a
grant of land from the Botanic Garden facing Hill Street to build a church
(Fig. 1, B). A subsequent enquiry with the government in Calcutta led to
Raffles’s orrginal land grant for the Botanic Garden being cancelled in July
1834. The desired parcel was then assigned to the Armenian community which
completed the Armenian Church in January 1835 and had it consecrated in
March 1836 (SSR, Z8, pp. 175-176, Act. Governor to Resident Councillor,
Aug. 1834, with enclosing letter to Sec. Gov. to Act. Govenor, 28 July 1834).
Records for the early land grants in the area of the original Botanic
Garden failed to turn up in a search at the Singapore Land Authority. Some
had apparently been reissued only in recent years. A school operated by the
Rev Darrah from August 1834 to December 1837 at the bottom of Government
Hill opposite the top of High Street, apparently was close to, but did not
encroach on the original Garden (Buckley, 1902, p. 128). On the north side of
the Bras Basah River, a simple Pauper Hospital was built between Bencoelen
(now Bencoolen) and Church (now Waterloo) Street in 1830 (Fig. 1, C). Three
years later a larger brick building was erected nearby as a permanent hospital
for convicts, paupers and European seamen. This building was located
approximately at the corner of Hospital Street (later Stamford Road) and
Bencoolen Street, i.e. across the street from where the Singapore History
Museum now is (Makepeace et al., 1921, vol. I, pp. 493-494). Beginning in
1841 a convict jail and other convict facilities began spreading out in the area
to the east of the hospital (Fig. 1, D; McNair, 1899, 54; ibid., plate X, facing
p. 77), and, yet further towards the sea, the new Roman Catholic Church
(Church of the Good Shepherd) was erected between 1843 and 1846 (Fig. 1,
E; Buckley, 1902, p. 248).
16 Gardens’ Bulletin Singapore 56 (2004)
Through the early 1830’s parts of the original Garden survived
insufficient maintenance, weeds and depredation by cattle (Bennett, 1834, vol.
I, p. 174; Earl, 1837, p. 350). A drive to the top of Government Hill, past
what was still known as the Botanic and Experimental Garden, became a
popular evening outing:
‘The drive up to this spot is exceedingly romantic - A spiral carriage road
winds up the hill, and, at each progressive step, fresh beauties attract the
eye. Eminences, undulating above each other, display broad patches either
cleared for cultivation, or shining in the bright green livery of clove
plantations’ (Begbie, 1834, p. 353).
The area provided the closest to a park-like ‘recreational’ space Singapore
had to offer and a respite from what was perceived as ‘the gloomy mass of
forest scenery’ (Bennett, 1834, vol. II, p. 139). Within the confines of the
former Garden visitors encountered a variety of ornamental and fragrant trees,
some indigenous like Kedah Gardenia (Gardenia carinata Wall.), others
imported such as Yellow Bauhinia (Bauhinia tomentosa L.) and Champaca
(Michelia champaca L.). There were also common medicinal plants, Purging
Croton (Croton tiglium L.) and the native gelam tree (Melaleuca cajuputi
Powell) that lent its name to Muslim quarter Kampong Glam, as well as the
introduced Borneo Camphor tree, Dryobalanops aromatica C.F.Gaertn.
(Burkill, 1935, vol. I, pp. 862-863). Many of the older nutmeg and clove trees
continued to produce fruit, and even some of the cotton plants from Raffles’s
days survived so that in 1836 their seeds could be collected to conduct a trial
for larger scale, private cotton plantations (Bennett, 1834, vol. II, pp. 174—
176; Thomson, 1850, p. 142; Crane, 1851, p. 121). This was part of broader
surge in agricultural interest that began to accelerate during the 1830s.
Singapore Agricultural and Horticultural Society, 1836 to c. 1845
Despite some initial interest shown by Europeans, development of plantation
agriculture had been hampered by a number of issues. During the 1820s access
to cultivatable areas further away from town was difficult, as well as dangerous,
so that the interior of the island remained almost unknown to Europeans (Earl,
1837, p. 353). The reluctance of the government to sell land in perpetuity
combined with the long maturation period of spice trees also made the shorter
term leases offered unattractive to planters (Thomson, 1850, pp. 210-219;
Makepeace et al., 1921, vol. I, pp. 301-311). Until about 1835 the cultivation
of spices was therefore restricted to about a dozen merchants with ‘small
amateur plantations of spice-trees near their residences’, close to town
(Singapore Chronicle, 15 May 1834; Earl, 1837, p. 410). Gradually some
long-term residents of Singapore accumulated enough free capital to explore
local investment options besides trade, and the relative success of the existing
Singapore’s First Spice Plantation and Botanic Garden 17
nutmeg plantations and other potential crops caught their attention (Little, 1849,
p. 678; Cameron, 1865, p. 168). Renewed interest in spices was helped when
locally the price of nutmegs spiked after 1835, aided by the rapidly declining
output of Bencoolen’s neglected plantations and diminished production of the
Banda spice gardens that were recovering from the impact of volcanic eruptions
in the 1820s (Warburg, 1897a, pp. 155-157; ibid., pp. 262—263; Jackson,
1968, table 11, p. 123).
Montgomerie, who had returned to Singapore in December 1834 as
Senior Surgeon, was one of the Europeans who rapidly developed spice
plantations. In 1836, he purchased Ryan’s Hill, a ten-year-old 16 acre nutmeg
plantation and developed it in conjunction with the neighbouring Craig Hill
into a large estate (Makepeace et al., 1921, vol. I, p. 517; Gibson-Hill, 1958,
pp. 147-148). His medical colleague and later successor as Senior Surgeon,
Dr Thomas Oxley (1805-1886), had an equally strong interest in economic
plants and botany. He dispatched plant specimens to Calcutta and Kew as
well as contributing “a large number of peculiar plants’ to the Buitenzorg Garden
(now Kebun Raya, Bogor) in Java (Treub, 1893, p. 48; Burkill, 1927, p. 129).
His nutmeg plantation on the slopes of “Oxley Hill’ became renowned for its
beauty, and his residence on top of the hill was surrounded by a considerable
collection of rare ornamental plants (Jagor, 1866, pp. 18-20; Buckley, 1902,
p. 405). Montgomerie’s and Oxley’s enthusiasm for nutmeg cultivation
interfered sufficiently with their medical duties that after 1857 medical officers
were forbidden to engage in any agricultural or commercial pursuits (Turnbull,
1972; B..217):
In May 1836, a number of influential European residents, Montgomerie
and Oxley amongst the leadership, organized the Singapore Agricultural and
Horticultural Society to represent their interest as planters. Their ultimate
goal was that ‘this island should all be cleared and cultivated; in fact become a
large Garden’ (Singapore Free Press, 9 June 1836; Singapore Free Press, 6
April 1837; Earl, 1837, p. 409; Buckley, 1902, pp. 305-306). The members
soon addressed their most pressing issue by petitioning the Governor-General
to review the restrictive land rights, although it took until 1845 that, with the
approval of the Court of Directors, large areas of land could be sold in perpetuity
(Singapore Chronicle, 15 October 1836; Makepeace et al., 1921, vol. I, pp.
310-311). As a means of funding its activities, on 19 November 1836 the
Agricultural and Horticultural Society received a land grant for seven acres on
Government Hill and access to convict labour so that it could resuscitate the
neglected nutmeg plantation (Makepeace et al., 1921, vol. I, p. 70). Fortunately
a significant number, by one estimate in excess of 200 nutmeg trees, had
survived in the original Garden from the early 1820 (Singapore Free Press, 25
November 1841) and in 1837 the Society was already able to earn $270 from
the sale of about 325 Ib of nutmegs. Among their other activities was the mutual
exchange of seeds with similar Societies in Calcutta and Bombay. It also tried
18 Gardens’ Bulletin Singapore 56 (2004)
encouraging the Chinese population to cultivate a broader range of crops and
vegetables by distributing pamphlets and seeds, though apparently with little
success (Singapore Free Press, 6 April 1837; Singapore Free Press, 6 July
1837; Buckley, 1902, p. 361). The Society continued to formally operate for
about ten years but with increasingly less visibility, and, as its members became
absorbed with their own agricultural endeavours, the nutmeg plantation on
Government Hill reverted back to the government.
‘Nutmeg Mania’
For this was the time when first many European, then Chinese and Malay
inhabitants, were swept away in ‘nutmeg mania’, frantically buying up land
and expanding their plantations (Belcher, 1848, vol. I, p. 381; Cameron,
1865, p. 168). Consequently the town’s surroundings to a distance of about
five miles underwent a dramatic transformation during the 1840s:
‘A few years ago it was a dense jungle: On every hill may now be seen the
residence of some hospitable merchant, surrounded by plantations of
nutmeg or other spice trees’ (Keppel, 1853, vol. I, p. 409).
Corresponding developments took place around Government Hill and
can be traced with early maps of Singapore. The first map to show land use
patterns was prepared in 1835 by George Drumgoole Coleman (1795-1844)
based on his surveys of the early 1830’s. At the time only a few scattered
spice plantations existed around Singapore and therefore, unlike other crops,
nutmeg and cloves were not yet specifically identified on his map. A strip
along Stamford Road, across from the Chinese Hospital, appears to have some
agricultural use, but it is impossible to clearly identify the remnants of the
government spice plantation as such on Coleman’s map (Tassin, 1836; Tassin,
1839; Goh, 1990, pp. 60-64).
In 1841, John Turnbull Thomson (1821-1884) was appointed
Government Surveyor of Singapore to help clarify the issue of land rights.
His Plan of Singapore Town and Adjoining Districts, based on surveys
conducted in 1842, represented the first map of Singapore to establish accurate
property boundaries. His maps clearly delineated a significant nutmeg
plantation in an area labelled ‘Botanical Garden’ on the north-eastern side of
Government Hill, indicating the renewed agricultural use after the Agricultural
and Horticultural Society had taken over the management of the site (Thomson,
1846; Thomson, 1854). By the end of 1848 this plantation contained 778
nutmeg trees, of which less than 50 were ‘of the old stock, most having been
planted since 1836’, and it had expanded well beyond the boundaries of
Montgomerie’s previous plantation to over 13 acres (Oxley, 1848, p. 657). A
total of 350 trees were bearing in 1848, the rest being too young or unproductive,
and yielded the government an income of $1134, while expenses were kept
Singapore's First Spice Plantation and Botanic Garden 19
low due to the use of free convict labour (Oxley, 1848, p. 659; Thomson,
1850, p. 31). This was about the height of Singapore’s nutmeg craze when a
total of 1,190 acres around town were planted with over 71,000 nutmeg trees
_ (Thomson, 1850, fold-out tables, facing p. 219).
Demise of Nutmeg Cultivation, 1859
As nutmeg production in the British and Dutch possessions gradually expanded,
demand proved to be inelastic, and prices on the saturated world markets
continued their long-term downward trend between 1840 and 1860 (Warburg,
1897a, pp. 518-519; Jackson, 1968, table 11, p. 123). Nutmeg planters in
Singapore and Penang incurred significantly higher expenses for labour and
fertilizer than their competitors in the Moluccas, and hence found it more and
more difficult to compete. What promised to be a profitable undertaking when
the nutmeg trees were planted, no longer was so when the first harvests were
reaped five to ten years later.
British planters were put at a further disadvantage when fungal and insect-
born diseases began to spread through their plantations in the late 1840’s. The
most significant disease was’ ‘nutmeg canker’, which caused premature
dehiscence and became quite prevalent in Penang, but also took effect in
Singapore (Little, 1849, pp. 678-681; Singapore Free Press, 12 July 1855;
Ridley, 1912, pp. 135-138; Flach & Tjeenk Willink, 1999, p. 147). Over the
course of the 1850s the first European planters began abandoning their ventures,
but the final demise of nutmeg cultivation in Singapore began in 1859 when a
mysterious disease ravaged the plantations (Singapore Free Press, 7 June 1860;
Cameron, 1865, pp. 168-170; Jagor, 1866, pp. 21-22), sending trees into
rapid decay:
‘In the night a tree would be attacked, and the morning light would show its
topmost branches withered; the leaves fell off; the disease slowly spread
downwards, chiefly on one side of the tree; and, in spite of every attempt
to check it (the lower portion often being for a long time green and bushy),
the tree became an unsightly mass of bare and whitened twigs’
(Collingwood, 1867, p. 47).
The pest was later identified as a small bark boring scolytid beetle
(Hyledius cribratus Blandf., previously Phloeosinus cribratus Blandf.), which
killed off the trees by attacking the cambium layer (Ridley, 1896, p. 92; Ridley,
1912, pp. 125-130; Flach & Tjeenk Willink, 1999, p. 147). By 1864 all but
one nutmeg plantation in Singapore and most in Penang had been destroyed.
Nutmeg mania in Singapore ended with large tracts of abandoned land
surrounding the town overgrown by secondary vegetation and financial
devastation for private planters who had speculated on their crops as much as
on real estate values.
20 Gardens’ Bulletin Singapore 56 (2004)
In 1851, the Kew-trained botanical collector Berthold Carl Seemann
(1825-1871), stopping over in Singapore on a voyage around the world still
had cause to admire the slopes of Government Hill as ‘clothed with numerous
Nutmeg-Trees, and a turf of brilliant green’ (Seeman, 1852, p. 82).
Even an updated map of 1857 shows a significant nutmeg plantation
between Armenian Street and the by-now expanded cemetary on the Hill, though
it is unclear to what degree the trees were actively maintained (Narayanan,
1857). In November 1859 some ‘enterprising citizens’ formed a second
Singapore Agri-Horticultural Society to stimulate the culture of flowers and
vegetables with the ultimate goal of creating a pleasure garden (Straits Times,
12 November 1859; Burkill, 1918, p. 55). The government was quick to offer
the defunct plantation on Government Hill as well as the labour of convicts,
but the Society wisely declined the land and settled a few weeks later for a
larger plot of 56 acres at Tanglin, which has remained the site of the current
Botanic Gardens (Straits Times, 24 December 1859; Makepeace et al., 1921,
vol. II, p. 73). As no further agricultural or horticultural use could be found
for the eastern side of Government Hill, now overlooked by the newly-built
Fort Canning, the land was gradually allocated to a handful of ecclesiastical
and public buildings that sparsely filled some of the area originally reserved
for Singapore’s first Botanic Garden.
Although there are now no remnants of the original Garden, Raffles’s
overall design for Government Hill left an enduring legacy: Initial exclusion
of the Garden area from commercial and residential use allowed its partial
preservation and enhancement as a green space after the Botanic Garden was
discontinued. Despite continued encroachment over the past 180 years, a
significant “green island’ on Fort Canning Hill has survived as a consequence
of Raffles’s original plan.
The Failure of Raffles’s Programme of ‘Cultivation’
Taking a broader perspective, it becomes apparent that Raffles’s agenda as an
advocate for a Botanic Garden extended well beyond his passion for natural
history or the hope for the ultimately elusive financial rewards of spices. True
to the Georgian ideology that the advance of agriculture lay at the heart of
creating an ‘improved’ society, Raffles had previously proclaimed for
Bencoolen: ‘Iam endeavouring to cultivate the soil, and to civilize the people’
(Raffles to ?, 14 July 1820, in Raffles, 1830). When he, through Wallich,
pleaded with the government to support the Garden in Singapore, he not only
promised rewards of a commercial nature but also an ameliorating influence
on the public at large (Wallich to Raffles, 2 November 1822, in Bastin, 1981).
In the spirit of enlightened imperialism, Raffles considered the Botanic Garden
an instrument of development:
Singapore’s First Spice Plantation and Botanic Garden 21
‘I am laying out a botanic and experimental garden, and it would delight
you to see how rapidly the whole country is coming under cultivation’
(Raffles to Marsden, 21 January 1823, in Raffles, 1830).
His broader programme of ‘cultivation’ was to be complemented by
another grandiose idea, the Singapore Institution, combining a new Malay
College and the Anglo-Chinese College of Malacca to create ‘the means of
civilizing and bettering the condition of millions’ (Raffles to Wallich, 17 April
1823, in Bastin, 1981). The idea of moving the Malacca College to Singapore
arose in January 1823 during a visit to Singapore of Rev Dr Robert Morrison
(1782-1834), the founder of the Anglo-Chinese College in Malacca. Morrison
was a ‘devoted friend’ to botany (Broomhall, 1924, pp. 105-107; Harrison,
1979, figure 5; ibid., p. 60) and therefore shared another vision with Raffles,
namely to establish a botany department and extend the garden attached to his
Malacca College:
‘A botanical garden was originally projected, but has failed, from
mismanagement. The object of it was to collect, under one view, all the
tropical plants of the Eastern Archipelago’ (Newbold, 1839, p. 183).
Despite his initial enthusiasm, in 1826 Morrison finally had to abandon
the plan of moving his Anglo-Chinese College to Singapore and blamed
Crawfurd, ‘the infidel doctor-civilian’, for it (Harrison, 1979, p. 75). Raffles’s
Singapore Institution was formally founded in April 1823 and amongst the
numerous provisional appointments was Montgomerie as Professor of Natural
Philosophy. But upon Raffles’s departure this project did not fare much better
than the Botanic Garden: By the early 1830s the poorly constructed building
of the Institution had turned into a landmark eye-sore (Earl, 1837, p. 351).
Raffles’s aspirations for a botanical enterprise collided with a broader
shift in British policy triggered by the death of both George III and Banks in
1820. The loss of the great sponsor and defender of the usefulness of science
and, in particular, botany, eventually sent many of the Banksian public initiatives
into retreat. As early as 1821 the War Office cut funding for the Botanic Garden
in St. Vincent (Guilding, 1825, pp. 22—23), while Kew Garden’s plant collectors
began to be recalled from abroad. This official pressure for retrenchment in
the pursuit of natural history increasingly also pervaded the East India Company
where the practical value of the existing Botanic Gardens was being questioned
(Drayton, 2000, pp. 130-131). Raffles’s plans for his Botanic and Experimental
Garden with a focus on spice cultivation not only violated basic economic
considerations, when London’s storehouses were still overflowing with spices,
but were also based on an outdated paradigm of public funding for his ventures.
In the age of retrenchment, the initiative and burden was shifting to private
support by local amateurs interested in botanical pursuits, often with an eye to
investment opportunities. This is exemplified by Montgomerie, a member of
the East India Company’s medical service, whose passion for economic botany
eventually eclipsed his dedication to medicine. Not only did he emerge in
22 Gardens’ Bulletin Singapore 56 (2004)
Singapore as a key supporter of private spice growers, he was also one of the
first two Europeans to attempt large scale sugar cultivation and claimed priority
for introducing the uses of Gutta Percha (Palaquium gutta (Hook.) Baill.) to
Europeans (Oxley, 1847, p. 22; Thomson, 1848, pp. 138-140).
After 1820, both domestically and abroad, Agri-Horticultural Societies
organized efforts to substitute government patronage of Botanic Gardens and
botanical exploration in general (McCracken, 1997, pp. 7-8; Drayton, 2000,
pp. 132-133). In Singapore, the first local Society operated Raffles’s Botanic
Garden primarily to fund its main objective of supporting the planters’ interests.
On the botanical front, their efforts fell short, since amateurish enthusiasm
could not make up for proper training or systematic experimentation. H.N.
Ridley, the first Director of the current Singapore Botanic Gardens, later
claimed that the ‘apathy of Government’ caused the disastrous decline of spice
cultivation in Singapore and Penang, because nutmegs were grown ‘only
empirically and without the aid of any scientific botanist’ (Ridley, 1905, p.
297; ’Ridley, 1910, p. 103), but this criticism would apply to both public and
private efforts. Indeed, pursuit of a more rigorous ‘Experimental Garden’ as
originally envisaged by Raffles might have revealed much sooner that nutmeg
culture on a larger scale was not viable in Singapore.
Conclusions
Raffles’s vision of spice cultivation in Singapore was embedded in the Georgian
philosophy of improvement, but was also based on flawed economic and
agronomic premises, and as such was doomed to failure from the onset.
Nonetheless, Raffles’s Botanic and Experimental Garden was far more than a
‘false start’ for the current Botanic Garden (McCracken, 1997, p. 8; Tinsley,
1989, pp. 14-17). Characteristic for the changing patronage for botany at the
time, it spawned a group of ‘botanic entrepeneurs’ who for a period of time
organized an Agricultural and Horticultural Society. In their pursuit of nutmegs
and other crops, they brought large areas of forest around Singapore town
under cultivation, thus leaving an indelible imprint on the environment. The
‘nutmeg mania’ that ultimately ensued was not only a speculation on future
nutmeg prices but also bore the signs of an ‘asset bubble’ in real estate that
quickly collapsed when nutmeg cultivation finally failed. The nutmeg
plantation on Government Hill survived in its original location until the late
1850s, much longer than previously thought, reaching its greatest extent around
1848 during the peak of frantic spice cultivation in Singapore. It exerted a
significant influence on land development in the vicinity of the settlement, as
well as helped to preserve ‘green space’ near the centre of town.
Singapore’s First Spice Plantation and Botanic Garden 23
Acknowledgements
The authors thank John Bastin for his continued encouragement during this
study and gratefully acknowledge the support of Ms Julie Yeo of Antiques of
the Orient, Singapore, and Mr Leong Foke-Meng, Singapore, who provided
previously published and unpublished maps. Ron Ranson, University of
California San Diego, greatly assisted in accessing literature.
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Gardens’ Bulletin Singapore 56 (2004) 29-42. 29
New Grass (Poaceae) Records for Singapore,
Including Panicum laxum New for Asia
HELENA DUISTERMAAT
The Herbarium, Singapore Botanic Gardens,
1 Cluny Road, Singapore 259569
Abstract
Working on a field guide to the grasses of Singapore, 15 records not included in The Concise
Flora of Singapore were discovered. Eight species and a variety are either recorded for the
first time for Singapore, or their presence, which was doubted before, is confirmed. They are:
Cyrtococcum patens (L.) A.Camus, Dichanthium annulatum (Forssk.) Stapf, Eragrostis
cilianensis (Bellardi) Vignolo ex Janch., Panicum paludosum Roxb., Panicum laxum Sw.,
Paspalum plicatulum Michx., Rottboellia cochinchinensis (Lour.) W.D. Clayton, and
Sporobolus indicus (L.) R.Br var. pyramidalis (Beauv.) Veldt. and S. tenuissimus (Schrank)
Kuntze. Panicum laxum is a new record for Asia. The occurrence of six more species already
known from Singapore but not reported in local floras is noted: Cenchrus echinatus L.,
Digitaria bicornis (Lam.) Roem. & Schult., D. mollicoma (Kunth) Henrard, Eragrostis brownii
(Kunth.) Nees (including E. cumingii Steud. var. cumingii), Panicum trichocladum Hack. ex
K. Schum., and Urochloa piligera (F. Muell. ex Benth.) R.D. Webster. For the sake of
completeness, they are included with a short discussion.
Introduction
Ridley (1907) provided the first account of the grass family (Poaceae) for
Malaya (including Singapore), followed much later by Gilliland (1971).
Recently, Keng et al. (1998) summarised the family for Singapore. The keys
provided in all works (in the latter unfortunately not to the species level) are
very technical. Also, some quite common species rarely seem to flower in
Singapore. They cannot be identified with the available literature. To tackle
the diverse identification problems that exist, 1am working on a field guide to
the grasses of Singapore (excluding the woody bamboos). During the course
of this project, I came across 15 new records, not mentioned or accepted in
Keng et al. (1998). The species belong to the genera Cenchrus, Cyrtococcum,
Dichanthium, Digitaria, Eragrostis, Panicum, Paspalum, Rottboellia,
Sporobolus and Urochloa. The presence in Singapore of some of them had
already been mentioned by others, but for the sake of completeness they are
included here. The species are discussed in alphabetical order.
Cenchrus
In Keng et al. (1998), Cenchrus brownii Roem. & Schult. was considered in
the wider sense, including C. echinatus L. Veldkamp (1999) accepts both
30 Gardens’ Bulletin Singapore 56 (2004)
species, and confirms the latter for Singapore. Both species are weeds of ruderal
places, native to America. Although Cenchrus brownii was first recorded for
Singapore (Sinclair SFN 38881, 1950), it is now more rare than the more
recently introduced C. echinatus (first collected in 1994). Both have spikelets
in groups (or burs) surrounded by a dentate involucre, with the burs arranged
on a single common axis. —
Key to the Cenchrus species of Singapore
la. Burs 1-2 mm apart, spines at base slender, 0.1—0.2 mm wide, erect, 24
(—7) mm long, apical lobes at margin with hairs up to 0.5 mm long. Spikelet
4—5 mm long. C. brownii
1b. Burs 1.5-4.0 mm apart, spines at base stout, 0.3-0.5 mm wide, recurved,
up to 4 mm long, apical lobes at margin with at least few hairs 1.0—1.5 mm
long. Spikelet 5—6.2 mm long. C. echinatus
Cenchrus echinatus L.
Singapore collections (in SING unless indicated otherwise): Ali Ibrahim &
Chin AI223 (1994) Armenian Street; Ali Ibrahim AI224 (1994) Bedok Road,
Bedok Corner; Tan et al. 2043a (1996) Pulau Tekong (SINU). Teo TM11 (2000)
Tanah Merah (SINU). Samsuri, A. et al. 290 (2002) Pulau Tekong; Duistermaat
$120 (2003) Pulau Ubin, near Jetty, ruderal place on sandy soil, unshaded;
$149 (2003) Orchard Road, next to Specialist Shopping Centre, ruderal open
place, shaded; S224 (2003) Cluny Road, recently replanted flowerbed with
fertile, humus-rich soil, partly shaded; S230 (2003) Labrador Villa Road, edge
of field, unshaded.
Cyrtococcum
Keng et al. (1998) mentioned two species of Cyrtococcum for Singapore, C.
accrescens (Trin.) Stapf and C. oxyphyllum (Steud.) Stapf. A third species, C.
patens (L.) A.Camus, was mentioned for Malaysia, in shady places, but not for
Singapore (Gilliland, 1971). I found collections from 1993 onwards of C. patens
in the herbaria of both SING and SINU, and collected it in secondary to primary
forest in Singapore.
Key to the Cyrtococcum species of Singapore
(Veldkamp, pers. comm.).
la. Longest pedicel of the pair shorter than spikelet. Inflorescence contracted.
C. oxyphyllum
1b. Longest pedicel of the pair longer than spikelet. Inflorescence more open.
2
New Grass (Poaceae) Records for Singapore, Including Panicum laxum New for Asia a1
2a. Erect part of culms 30-100 cm long. Sheath usually pilose all over. Panicle
20-50 by 6-30 cm. Spikelets usually 1.35—1.5 mm long.
C. accrescens
2b. Erect part of culms 10-30 cm long. Sheath usually pilose along margins
only. Panicle 3-18 by 0.8—2.3 cm. Spikelets usually 1.5—1.8 mm long.
C. patens
Cyrtococcum patens (L.) A.Camus
Culm with erect part 10-30 cm tall. Nodes hairy on one side. Sheath with
hairy margins only. Ligule 0.4—0.8 mm long, membranous. Blade 4—7 cm long,
5—8 mm wide, base at margin with up to 4 mm-long bulbous-based hairs.
Mature inflorescence 4—9 by 1—2.3 cm. Longest pedicel of the pair 2.5-4.0
mm long. Spikelet 1.5—1.8 mm long, 0.9-1.0 mm wide.
Singapore collections: Chua et al. H135 (1993) Pulau Hantu (SINU). Lai &
Saifudin LJ549 (1999) Loyang Avenue, back mangrove (SINU). Duistermaat
$176 (2003) Pulau Ubin, E of Kampong Melayu, abundant along paths and
open patches in old rubber plantation with remnants of old forest on top of the
hill; S219 (2003) Bukit Timah Nature Reserve, Senapang Road, shaded roadside
in secondary forest; $226 (2003) Pulau Ubin, Jalan Ubin, secondary forest
(SING).
Another specimen, Wong PW. s.n. (1959; SINU) is intermediate
between C. accrescens (sheath sparsely hairy all over, panicle 20—35 cm long)
and C. patens (spikelet 1.6—1.7 mm long). It could either be a hybrid, or a form
of C. accrescens with unusually small spikelets.
Dichanthium
Although Gilliland (1971) mentioned Dichanthium annulatum (Forssk.) Stapf
for Singapore, Keng et al. (1998) did not include it, because they had not seen
any Singapore material. The species is different from both D. caricosum (L.)
A. Camus and D. mucronulatum Jansen in having an inflorescence with 3—5
racemes (only | raceme in the other two). Superficially, it resembles a poorly
developed Bothriochloa bladhii, the inflorescence of which as a rule has more
than 10 racemes. Moreover, the racemes and pedicels are furrowed in the latter,
not so in D. annulatum. I was able to identify collections as D. annulatum both
in SING and SINU, and made three collections (all from one area) in Singapore.
This species of Paleotropic origin is mainly a roadside ruderal. Because
of its tufted habit with rather long and tough decumbent culms, it can be a
nasty weed in lawns.
32 Gardens’ Bulletin Singapore 56 (2004)
Dichanthium annulatum (Forssk.) Stapf
50-200 cm tall. Tufted, decumbent. Nodes hairy. Sheath 3—7 cm long, glabrous,
slightly compressed, not keeled, margin glabrous. Ligule 1.5 mm long, hairy,
membranous. Blade 7-17 cm long, 3—6 mm wide, hairy at least on upperside,
base rounded, with a few up to 5 mm-long hairs. Inflorescence with 3-5 simple
racemes, 3.5—7 cm long, axils hairy. Spikelets in pairs, one sessile, one
pedicelled. Pedicel 1.8—2.0 mm long, somewhat flattened but not furrowed,
hairy on one margin. Sessile spikelet 3.5-3.9 mm long, 1.0—1.2 mm wide,
dorsiventrally flattened, 2-flowered, awned; both lowest bracts (glumes) as
long as the spikelet. Awn 2—2.5 cm long. Pedicelled spikelet reduced to 2
glumes.
Singapore collections: Teruya 2012 (1932) Newton (SING). Sinclair SFN
39429 (1951) Tanjong Pagar Docks (SING). Wong P.W. s.n. (1959) General
Hospital grounds (SINU). Sinclair SFN 10784 (1965) Tanjong Pagar, Nelson
Road (SING). Ali Ibrahim AI337 (1998) Upper East Coast Road (SING). Lai
LJ498 (1999) Tanglin, Dempsey Road, Civil Service Club (SINU).
Duistermaat S16 (2002) Mount Sinai Drive, shaded grassy roadside; S25
(2002) Mount Sinai Drive, grassy roadside, partly shaded, frequently mown;
$118 (2003) Mount Sinai Drive, on 300 cm high stone wall in backyard,
unshaded (SING).
Digitaria
The genus Digitaria comprises worldwide c. 170 species, the delimitation of
which has long been a problem. For the 27 Malesian species, there is the
revision by Veldkamp (1973). The treatment of the genus in Keng et al. (1998)
obviously follows this revision. However, they did not include D. bicornis
(Lamk.) Roem. & Schult. and D. mollicoma (Kunth) Henr., although in the
revision the first was recorded as ‘throughout Malesia’ and the latter as
‘Singapore [among others]’. Material of both species is present in both SING
and/or SINU, and I collected them recently as well. Veldkamp (pers. comm.)
provided a separately distributed identification list of the specimens he had
examined, including those from SING. An updated list is given here.
Digitaria bicornis is a species from waste places, often on sand, also along
the shore. Its recorded presence on the island dates back to the 19" century.
The paired spikelets are heteromorphous, i.e. sterile lemma of the sessile
spikelet glabrous with nerves equidistant to slightly pubescent with nerves
not equidistant, that of the pedicelled spikelet always more pubescent to
bristled, nerves not equidistant; if basal spikelets homomorphous, then
completely glabrous. In Singapore there is only one other taxon with
heteromorphous spikelets, D. setigera Roem. & Schult. var. calliblepharata
New Grass (Poaceae) Records for Singapore, Including Panicum laxum New for Asia 33
(Henr.) Veldk. The two can be separated by the absolute and relative length of
the upper glume: 1—2.75 mm long and 0.35-0.8 times as long as the spikelet
in—D. bicornis, versus 0.3—1 mm long and 0.15—0.3 times as long as the spikelet
in—D. setigera var. calliblepharata (Veldkamp, 1973, pers. comm.). Singapore
collections (all in SING): Ridley s.n. (1890) Changi. Burkill 4669 (1919) Tanah
Merah Besar. Teruya 2221 (1932) Newton. Corner s.n. (1941) Yio Chu Kang
(Veldkamp det. as D. nuda). Duistermaat S142 (2003) Gim Moh, raised
flowerbed, unshaded; $184 (2003) Pulau Ubin, Chek Jawa, house no 1, ‘The
English House’, on rock facing the sea; S217 (2003) East Coast Park; $253
(2003) Sarimbun, Jalan Bahtera, grassy roadside, under trees, in shade.
Digitaria mollicoma, like D. longiflora (Retz.) Pers., is a stoloniferous, mat-
forming grass with spikelets in groups of three. Both grow on open, shaded to
unshaded, humid, sandy to rocky soil. Both species seem to be fairly recent
introductions to Singapore, the oldest record is from 1955 for the first, 1958
for the latter (H.B. Gilliland 1782). The two differ in the length of the spikelets,
relative length of the upper glume, and the positioning of the veins on the
sterile lemma (Veldkamp, 1973, pers. comm.):
la. Spikelets 1.8—2.5 mm long. Upper glume 0.7—1 times as long as the spikelet.
Sterile lemma with 7 more or less equidistant veins. D. mollicoma
lb. Spikelets 1.3-1.7(-1.9) mm long. Upper glume as long as the spikelet.
Sterile lemma with 5—7 more or less inequidistant veins. _D. longiflora
Singapore collections: Abu Kassim s.n. (1955) Bedok roadside (SINU); 1754
(1958) University grounds. Jumali K994 (1964) Botanic Gardens, near the
pond (SINU). Chan s.n. (1984) NUS, Kent Ridge Campus, waste ground
(SINU). Lai LJ493 (1999) junction of Sembawang Road and Gambas Ave,
open waste ground (SINU). Duistermaat S31 (2002) Holland Road, inclined
verge with grass vegetation, shaded; S160 (2003) Upper Pierce Reservoir, dirt
road through forest, in low vegetation on wet soil along free running streamlet;
$240 (2003) Pulau Subar Laut (Big Sisters Island), sandy soil just above high
water-line, underneath coconut trees (SING).
Eragrostis
The delimitation of the species of Eragrostis (c. 350 species worldwide) is
troublesome. However, for the Malesian region they were recently revised by
Veldkamp (2002), who clarified many problems. Most of the names he mentions
for Singapore are included in Keng ef al. (1998), although the delimitation is
different in a few cases. Most dramatic is the change in the E. atrovirens-
elongata complex. The name E. elongata (Willd.) J.Jacq., now to be regarded
as a synonym for the E. Malesian-Australian EF. diandra (R.Br.) Steud., appears
34 Gardens’ Bulletin Singapore 56 (2004)
to have been widely used for other taxa in Singapore including E. atrovirens
(Desf.) Trin. ex Steud., E. brownii (Kunth.) Nees and E. cumingii Steud. var.
cumingii. The last two, not mentioned in Keng et al. (1990), are not easily
distinguished. The discriminating character is in the length and shape of the
anthers, and to a lesser extent also in the colour of the pericarp (Veldkamp,
2002): anthers 0.1—0.2 mm long, globose, and pericarp cinnamon-coloured in
E. cumingii var. cumingti, versus anthers 0.3-0.4 mm long, ellipsoid, and
pericarp dark tea-coloured, rarely cinnamon-coloured as in E. brownii.
However, in the Singapore material the anthers are never globose (1.6 to 3.0
times as long as wide), the length of the anthers ranges continuously between
0.15 and 0.34 mm, and differences in the colour of the pericarp do not
correspond with this. I have therefore decided to treat the Singapore material
as one species, E. brownii in the wider sense (s.l.). It is distinguished from the
other Singapore species by the persistent paleas and the jointed rachilla that
will ultimately break up from above downward. It was collected for the first
time in 1930, and is today rather common, also in the urban environment.
Eragrostis brownii s.l.
Singapore collections (not already cited in Veldkamp, 2002; herbarium only
mentioned if not SING): Teruya 1299 (1930) Geylang, Teruya 2023 (1932)
Newton. Jeo TM30 (2000) Tanah Merah (SINU). Samsuri et al. 44 (2001)
Pulau Tekong. Duistermaat S4 (2002) Mount Sinai Drive, road side and between
pavement; S24 (2002) Holland Road area, Mount Sinai Rise, bank of terrace
house, with grass vegetation; S53 (2002) Sungei Kadut area, Mandai Quarry,
off Woodlands Road, waterlogged soil; S68 (2002) Sungei Buloh, on gravel
path, unshaded, low density vegetation; S145 (2003) Holland Road area, Mount
Sinai Drive, bank of terrace house, with grass vegetation, unshaded; S185
(2003) Pulau Ubin, Chek Jawa, house no 1, ‘The English House’, between
cracks of staircase leading to private jetty; S213 (2003), N-side of Sultan’s
Land, off Gallop Road, steep slope with seepage water, low and dense
vegetation, unshaded; S256 (2003) Sarimbun, Bahtera Track, sunny roadside
on sandy clay.
Furthermore, the collections in SING revealed, rather surprisingly, an
old collection of E. cilianensis (Bellardi) Vignolo ex Janch. (Corner s.n., 1941,
Yio Chu Kang; Veldkamp det. as E. unioloides, but lemmas lack the granular
structure that is typical for this species). The species is recognised by its
persistent paleas and the presence of obvious glands on leaf sheath and blade.
In Malesia, it is recorded as a weed of waste places, especially in regions with
a distinct dry season, from the Philippines, Java, Lesser Sunda Islands,
Moluccas, and introduced in New Guinea (Veldkamp, 2002). However, the
species does not seem to be part of the present day flora of Singapore.
New Grass (Poaceae) Records for Singapore, Including Panicum laxum New for Asia 35
Panicum
With 450 species worldwide, the genus Panicum is one of the largest within
the grass family (Poaceae). Only 26 species occur in the Malesian region
(Veldkamp, 1996, 1999). According to Aliscioni et al. (2003, and references
therein), Panicum is a polyphyletic group. They propose a new generic
delimitation and divide Panicum into various genera (with a number of sections
still as incertae sedis). Because many of the Malesian species were not included
in the analysis, I prefer to consider their results as preliminary. I therefore
retain Panicum with its traditional delimitation (Aliscioni et al., 2003). The
genus is recognized by a lax to dense panicle with rounded to triquetrous
branches. The spikelets are 2-flowered and unawned with the upper glume as
long as the spikelet.
Six species of Panicum were confirmed by Keng et al. (1998) based on
Veldkamp (1996). Panicum trichocladum Hack. ex K.Schum. was confirmed
by Veldkamp (1999), whereas material of one more Panicum species was found
in the herbarium, and a third species was collected in the field. Panicum is
now represented by nine species in Singapore.
Panicum trichocladum, when present, is often abundant, scrambling
through shrubs up to 3 m high. Its diagnostic features include ligule
membranous, ciliolate, panicle branches at base without spikelets, lower glume
0.3-0.6 mm long and 0.1-0.2 times as long as the spikelet, and 2"4 lemma
apiculate and incurved. It has been collected from three localities in 1998
(specimens cited in Veldkamp, 1999), and more recently in the secondary forest
between Holland Road and Tyersall Avenue (Duistermaat S200, 2003; SING),
and inside Bukit Timah Nature Reserve, Senapang Link (Duistermaat S220,
2003; SING).
In the collections of SINU, I found two specimens of Panicum paludosum
Roxb. This species is similar to P. repens L. in having a collar-shaped lower
glume, but differs in having an entirely glabrous sheath and blade, and the
sterile lower floret without a palea or, when present, up to 0.7 times as long as
the lemma, while P. repens has a male lower flower with the palea more than
0.8 times as long as the lower lemma. It also has larger spikelets (3.3-4.5 mm
long against 2.6-3.25 mm; see Veldkamp, 1996). Panicum paludosum is an
aquatic species, favouring shallow waters, whereas P. repens usually grows in
terrestrial conditions and is only able to survive inundation temporarily
(Gilliland, 1971). Its presence in Singapore is a little surprising, as in Malaysia
P. paludosum is known only from the northern and central states of Kedah
(including Langkawi), Pahang and Penang. In Indonesia, the nearest localities
are in Sumatra and Borneo. The oldest Singapore collection is from 1959 (Wong
P.W. s.n., Tampines, floating in water). The second and newest collection is
from 1965 (Keng et al. s.n., unfortunately without locality). Although I have
not seen the species on my collecting trips, it could still be present in shallow
waters.
36 Gardens’ Bulletin Singapore 56 (2004)
In 2002, I found a delicate, c. 20 cm high grass with minute spikelets
c.1.3 mm long. It grew on a shaded roadside with rather dense, lawn-like
vegetation. At the time I could only identify it to Panicum. Later, on a field
trip with Tan Kai Xin (NUS) on Coney Island (Pulau Serangoon), I found a
similar plant growing near the coast on the reclaimed part of the island, on
coarse sand in rather open, unshaded vegetation. Finally, I discovered along
Clementi Road a third population of over a hundred plants in an open field
with clayey, waterlogged soil, and a fourth of some tens of plants on the shaded
roadside along Seletar West Farmway 1 (off Jalan Kayu). Bor (1960) and
Veldkamp (1996, 1999) do not list the species, and even the World Grass Species ~
Database at the Kew website (http://www.rbgkew.org.uk/data/grasses/
grasses.ink) yielded only a single, but not identical species. Finally, Hitchcock
& Chase (1910, 1915) and Hepper (1972) lead beyond doubt to P. laxum Sw.
A recent revision of section Laxa (Zuloaga et al., 1992) resulted in the same
positive identification.
The species is widely distributed:in America from Mexico to Argentina,
where it is common in wet and open, disturbed places, margins of roads, swamps
and rivers, at altitudes of 0-1500 m. During the early 20" century, the species
was introduced to West Africa (first collected in 1927) and is there now
naturalized on roadsides and in clearings, particularly on damp soil. Later, it
was also found in Australia (Simon, 1992, as Cliffordiochloa parvispiculata
and Simon, 2003, as Steinchisma laxa), confined to water channels
(Queensland). As far as I know, this is the first time Panicum laxum is recorded
for Asia. Its habitat in Singapore is comparable to where it is found in Africa,
but the Singapore plants seem to be smaller. Duistermaat S37 is different from
the other two collections in being single and prostrate, rather than tufted and
geniculate. It could be the cv. ‘Shadegro’, a form with a potential as a turf
grass for shaded conditions, which is established in the Brisbane Botanic
Gardens (Simon, 2003).
Panicum laxum is more closely related to the Asian P. auritum (also
occurring in Singapore, and the only two species of section Laxa occurring in
Malesia) than to any of the other species occurring in Malesia. The two can be
separated as follows:
la. Plant 80-120 cm tall. Leaf sheath and blade without transverse veinlets.
Spikelet 2.0-3.0 mm long. Upper glume and 1* lemma with (faint)
transverse veinlets. 1*t palea up to 0.75 times as long as 1* lemma.
P. auritum
1b. Plant 20-45 cm tall. Leaf sheath and blade with minute transverse veinlets.
Spikelet 1.3—1.4 mm long. Upper glume and 1* lemma without transverse
veinlets. 1* palea as long as 1* lemma. P. laxum
New Grass (Poaceae) Records for Singapore, Including Panicum laxum New for Asia 37
Panicum laxum Sw.
(Steinchisma laxa (Sw.) Zuloaga)
Culms 20-45 cm tall. Single or tufted, geniculate to prostrate. Nodes glabrous.
Sheath glabrous, with minute transverse veins, margin hairy at apex or upper
half only. Ligule 0.3-0.4 mm long, membranous, fimbriate. Blade 4-7 cm
long, 2-5 mm wide, glabrous. Inflorescence 5-13 cm long, 3-10 branches
scattered along rachis, patent, longest branch 2.5-6.0 cm long, branchlets
appressed to 1“ order branch. Pedicel 0.5—1.0 mm long. Spikelet 1.3—1.4 mm
long, 0.7 mm wide, obtuse. Lower glume 0.5—0.6 mm long, 0.5 times as long
as the spikelet, glabrous, 3 veins. Upper glume 1.2 mm long, glabrous, 5 veins.
1 lemma as upper glume, 1.2—1.3 mm long, 3 veins, obtuse to acute. 1“ palea
1.2-1.3 mm long.
Habitat in Singapore: Open to shaded roadside or field, on clayey or sandy
soil.
Singapore specimens: Duistermaat S37 (2002) Holland Grove Road, Henry
Park Apartments, roadside lawn, shaded (SING, L). Duistermaat NUS 1269
(2003) Coney Island, raised bank of reclaimed area near the coast, on coarse
sand in rather open and unshaded vegetation. (SINU, L). Duistermaat S285
(2004) Clementi Road, at junction with Dover Road, open field on waterlogged,
clayey soil, more than 100 plants; S287 (2004) Seletar West Farmway 1, on
shaded roadside on clayey soil, some tens of plants.
Paspalum
The collections in SINU include one specimen of Paspalum plicatulum Michx.
(Teo L3 (2000) Lazarus Island). The species is readily recognised by a
conspicuous dark centre on the oblong to (ob-)ovate, paired spikelets. It is
native to America, and has been found several times in northern Australia
(Mallett & Orchard, 2002: p. 348). It was found in Malesia only once, in a
cultivation plot in Papua New Guinea (De Koning & Sosef, 1985: p. 313, and
included in their key to the Malesian species). Plants are cultivated for their
high palatabilty to cattle in (sub-)tropical America (known as Brownseed
Paspalum), the Chinese province of Gansu (Chen & Phillips, undated), and
Papua New Guinea (De Koning & Sosef, 1985). It remains to be seen if this
species will establish itself in Singapore.
Rottboellia
In the collections of SINU, I found a specimen (Chua et al. 1074) that, with its
80 cm-long culms growing in tufts and with several spike-like racemes from a
single culm, superficially resembles Mnesithea glandulosa. With this species
it also shares the sessile spikelets with winged lower glumes, and the sterile
pedicelled spikelets (Veldkamp et al., 1986), but the gland-like warts on the
margins of the lower glumes of the sessile spikelets, which are characteristic
for this species, are lacking. Moreover, the male 1“ floret of the sessile spikelet,
38 Gardens’ Bulletin Singapore 56 (2004)
and the sheaths with bulbous-based hairs speak more for Rottboellia
cochinchinensis (Lour.) W.D. Clayton (Veldkamp et al., 1986). Thus, the
specimen is intermediate between the genus Mnesithea and R. cochinchinensis.
In SING, I found a second intermediate specimen (Samsuri et al. 315), but this
one has unwinged lower glumes, a less inflated rachis, and sterile pedicelled
spikelets of only 2 glumes. Earlier, an intermediate specimen from the
Philippines was described as Rottboellia paradoxa Koning & Sosef (Veldkamp
et al., 1986), but here the pedicels are only partly fused with the rachis; they
are completely fused in the Singapore material. All intermediate specimens
are different from each other. Study of the Malesian collections of R.
cochinchinensis in SING revealed that this species is probably much more
variable than earlier thought. I therefore keep both Singapore specimens under
this name. Because both were collected on recently reclaimed land, it remains
to be seen whether the species will become established in Singapore.
Rottboellia cochinchinensis (Lour.) W.D. Clayton
R. exaltata L.f. (see Clayton, 1981)
Culms up to 80 cm tall, tufted. Nodes glabrous. Sheath 6-7 cm long, with
bulbous-based hairs, rounded, margin hairy. Ligule 1.5 mm long. Blade 21-45
cm long, 11 mm wide. Inflorescence spike-like raceme, 5—7 cm long, several
from | stem, basal part internode flattened, 2—3.8 mm wide, upper part rounded,
24.5 mm wide. Spikelets in pairs, 1 sessile, 1 pedicelled; pedicel glabrous,
one side completely fused with rachis; sessile spikelet 4.5—5.0 mm long, 1.7-
4 mm wide, 1“ floret male. Lower glume granulate, margin smooth, apex
unwinged or with indurated wings. Pedicelled spikelet c. 4.5 mm long, of two
reduced, herbaceous, flat glumes, 0—2 smaller hyaline bracts inside.
Habitat in Singapore: Open wasteland on sand near the coast, waterlogged,
unshaded.
Singapore specimens: Chua et al. 1074 (1995) Marina East, reclaimed land
(SINU). Samsuri et al. 315 (2002) Pulau Tekong, reclaimed land (SING).
Sporobolus
In mid-2003, a small-tufted grass with lax panicles attracted my attention. I
have found it several times in Singapore, on roadsides, in flower beds and
flower pots The species is not mentioned in Gilliland (1971) or Keng et al.
(1998). The keys in Bor (1960), however, lead to Sporobolus tenuissimus, a
species native to tropical Africa, America, and India. Introduction of the species
with potting mix or garden material is obvious.
Baaijens and Veldkamp (1991) already noticed that this species had been
introduced into other regions in Asia (Vietnam, Indonesia: Java). It seems to
be spreading (rapidly) in India (Sreekumar, 1994), Indonesia (Veldkamp, 1997)
and Thailand (Veldkamp, 2003). We should therefore expect to see more of
this species in Singapore and Malaysia; it could even spread to everyone’s
highrise garden!
New Grass (Poaceae) Records for Singapore, Including Panicum laxum New for Asia 39
The plant superficially resembles Eragrostis amabilis (L.) Wight & Arn.
ex Nees, with which it was found growing, in the panicle shape and in the
hyaline parts of the spikelet. It immediately stood out because of the strictly
erect stems (prostrate to ascending in E. amabilis) and the 1-flowered spikelets
(vs. 3—6-flowered).
During fieldwork with H.T.W. Tan et al., I found a few plants of
Sporobolus indicus (L.) R.Br. that seemed particularly tall. For that reason I
collected them, and checking with Baaijens and Veldkamp (1991) they appeared
to be S. indicus var. pyramidalis (Beauv.) Veldk., a taxon from Africa and S.
America. Because it had been reported for Australia and the Pacific they
suggested it might turn up in Malesia. Veldkamp (pers. comm.) was able to
confirm this identification and that it is the first record of the variety for Malesia.
Key to the Sporobolus species and varieties in Singapore
(after Baaijens and Veldkamp, 1991)
la. Stoloniferous plants. Lower glume 0.55-0.9 times as long as spikelet, upper
glume 0.75-—1 times as long. S. virginicus
1b. Tufted plants. Lower glume 0.2—0:5 times as long as spikelet, upper glume
up to 0.67 times as long. 2
2a. Spikelet 0.9—1.25 mm long; pedicel 2-6 mm long. Lemma and palea hyaline.
S. tenuissimus
2b. Spikelet longer than 1.25 mm; pedicel less than 1 mm long. Lemma and
palea herbaceous. 3
3a. Upper glume truncate, less than half as long as the spikelet, slightly longer
than the lower glume. (Spikelets 1.7—2.2 mm long. Anthers 3).
S. indicus var. pyramidalis
3b. Upper glume more or less acute, 0.4—0.67 times as long as spikelet,
distinctly longer than the lower glume. . 4
4a. Panicle usually somewhat lax and branches with loose spikelets. Spikelets
usually 1.4—1.6 mm long. Anthers usually 2, 0.5—0.8 mm long. Seed 0.6—
0.9 mm long. S. indicus var. flaccidus
4b. Panicle usually contracted and branches with dense spikelets. Spikelets
usually 1.8—1.9 mm long. Anthers usually 3, 0.7—1 mm long. Seed 0.9-1.1
mm long. S. indicus var. major
Sporobolus indicus (L.) R.Br. var. pyramidalis (Beauv.) Veldk.
Plant 1.55 m high, tufted. Culm erect. Spikelet 1.9 mm long. Lower glume 0.3
mm long, truncate. Upper glume 0.6 mm long, truncate. Anthers 3, 0.9 mm long.
Habitat in Singapore: Unshaded, open vegetation, sandy soil.
Singapore specimens: Duistermaat and Tan et al. S304 (2004) 38x39 Neo
Tiew Crescent, open field of private company (SING, duplicated in L, living
collection in SINU).
40 Gardens’ Bulletin Singapore 56 (2004)
Sporobolus tenuissimus (Schrank) Kuntze |
Plant 20-70 cm high, single or in small tufts. Culm erect. Nodes glabrous.
Sheath and blade glabrous. Ligule 0.2 mm long, fimbriate. Blade up to 14 cm
long, 2 mm wide. Inflorescence a lax panicle, 15-36 by 4—6 cm, longest
branches 4 cm long, glabrous, very thin, with secondary branchlets. Spikelets
well-spaced, pedicelled, single, 1-flowered, 1.3 mm long, hyaline. Both glumes
much shorter than spikelet. Anthers 0.3—-0.4 mm long.
Habitat in Singapore: In flowerbeds and on roadsides, open vegetation,
unshaded. Seems to be associated with Eragrostis amabilis.
Singapore specimens (all in SING, duplicated in L): Duistermaat S131 (2003)
as a weed in potted plant at 56 Mt. Sinai Drive; $223 (2003) Bukit Timah
campus of Singapore Management University, flowerbed with fertile humus-
rich soil, unshaded; $259 (2003) Waterloo Street, near the church of St Peter
& St Paul, on sandy roadside with cement and brick debris, unshaded; S260
(2004) Cluny Road, in unshaded flowerbed with clayey sand.
Urochloa
Veldkamp (1999) noticed that besides the three Urochloa species mentioned
in Keng et al. (1998), a fourth, U. piligera (F. Muell. ex Benth.) R.D. Webster,
is present in Singapore. Like U. subquadripara (Trin.) R.D. Webster, spikelets
are solitary, and the lower glume has overlapping margins (Veldkamp, 1996).
It is immediately recognised as different, however, because of the hairy upper
glumes (glabrous in the latter). It is native to Australia and east Malesia
(Sulawesi to New Guinea), where it grows in open sandy places near the
seashore, and in dry rice fields. The first records for Singapore (1959, specimens
in SINU, cited in Veldkamp, 1999), were found inland, in swampy places, on
the bank of a canal and on a roadside. It was not collected again, until in 2002
I found the species in Sungei Buloh (Duistermaat S72), and in 2003 in Sarimbun
(Duistermaat S243; collections in SING). In both places it grew on dry to
damp roadsides on sandy clay in coastal areas on the northwest of the island,
a habitat comparable to its native one.
Acknowledgements
I thank Dr J.F. Veldkamp for his enthusiastic support of the project, his updated
identification keys for Flora Malesiana, and for the confirmation of the
identification of Panicum laxum and Sporobolus indicus var. pyramidalis and
S. tenuissimus. Singapore Botanic Gardens (Dr S.C. Chin), SING (Dr R. Kiew
and staff), and SINU (especially Prof H.T.W.Tan, Ms Morgany, and Mr Chua)
are gratefully acknowledged for their hospitality. It was a great pleasure to be
in the field with Tan Kai Xin, who also helped me in the library. Joseph Lai
and Keith Hillier accompanied me in Sungei Buloh. The staff of the SING
New Grass (Poaceae) Records for Singapore, Including Panicum laxum New for Asia 41
herbarium (Serena Lee, Paul K.F. Leong, Samsuri Ahmad) and Ali Ibrahim
(NParks) allowed me to join them on their trip to Pulau Ubin. All these people
have motivated me to continue the project on the grasses of Singapore, for
which I am grateful.
References
Aliscioni, S.S., L.M. Giussani, F.O. Zuloaga and E.A. Kellogg. 2003. A molecular phylogeny
of Panicum (Poaceae: Paniceae): tests of monophyly and phylogenetic placement within
the Panicoideae. American Journal of Botany. 90: 796-821.
Baaijens, G.J. and J.F. Veldkamp. 1991. Sporobolus (Gramineae) in Malesia. Blumea. 35:
393-458.
Bor, N.L. 1960. The Grasses of Burma, Ceylon, India and Pakistan. Pergamon Press, Oxford,
London, U.K.
Chen, S. and S.M. Phillips. Undated. http://www.fna.org/china/mss/volume22/
htm 21 Paniceae.htm#tribe
Clayton, W.D. 1981. Notes on the tribe Andropogoneae (Gramineae). Kew Bulletin. 35: 813-
818.
Koning, R. de and M.S.M. Sosef. 1985. The Malesian species of Paspalum L. (Gramineae).
Blumea. 30: 279-318.
Gilliland, H.B. 1971. Revised Flora of Malaya. 3: Grasses of Malaya. Government Printing
Office, Singapore.
Hepper, F.N. 1972. Panicum Linn. Flora of West Tropical Africa. 3: 426-435.
Hitchcock, A.S. and A. Chase. 1910. The North American species of Panicum. Contributions
from the United States National Herbarium. 15: 1-396.
Hitchcock, A.S. and A. Chase. 1915. Tropical North American species of Panicum.
Contributions from the United States National Herbarium. 17: 459-539.
Keng, H., S.C. Chin and H.T.W. Tan. 1998. Gramineae. The Concise Flora of Singapore,
Monocotyledons. 2: 145-188.
Mallett, K. and A.E. Orchard (eds). 2002. Poaceae 1: Introduction and Atlas. Flora of Australia.
43: 1-319.
Ridley, H.N. 1907. Gramineae. Materials for a Flora of the Malay Peninsula. 5: 118-197.
Simon, B.K. 1992. Studies in Australian grasses 6. Alexfloydia, Cliffordiochloa and
Dallwatsonia, three new panicoid grass genera from eastern Australia. Austrobaileya. 3:
669-68 1.
42 Gardens’ Bulletin Singapore 56 (2004)
Simon, B.K. 2003. Steinchisma laxa (Sw.) Zuloaga, the correct name for Cliffordiochloa
parvispiculata. Austrobaileya. 6: 561-562.
Sreekumar, P.V. 1994. New plant records for Bay Islands. Journal of Economic and Taxonomic
Botany. 18: 185-187.
Veldkamp, J.F. 1973. A revision of Digitaria Haller (Gramineae) in Malesia. Blumea. 21: 1-80.
Veldkamp, J.F. 1996. Revision of Panicum and Whiteochloa in Malesia (Gramineae-Paniceae).
Blumea. 41: 181-216.
Veldkamp, J.F. 1996. Brachiaria, Urochloa (Gramineae—Paniceae) in Malesia. Blumea. 41:
413-437.
Veldkamp, J.F. 1997. IX. Overlooked genera and species in the Malesian flora: the case of
Crocosmia (Iridaceae) and some others. Flora Malesiana Bulletin. 11: 512.
Veldkamp, J.F. 1999. New records of Gramineae for Malesia. Flora Malesiana Bulletin. 12,
5: 231-239.
Veldkamp, J.F. 2002. Revision of Eragrostis (Gramineae, Chloridoideae) in Malesia. Blumea.
47: 157-204.
Veldkamp, J.F. 2003. Miscellaneous notes on Thai Gramineae. Blumea. 48: 495-501.
Veldkamp, J.F., R. de Koning and M.S.M. Sosef. 1986. Generic delimitation of Rottboellia
and related genera (Gramineae). Blumea. 31: 281-307.
Zuloaga, F.O., R.P. Ellis and O. Morrone. 1992. Arevision of Panicum subgenus Phanopyrum
section Laxa (Poaceae: Panicoideae: Paniceae). Annals of the Missouri Botanical Garden.
79: 770-818.
Added in press: Three more new records need to be added.
Neyraudia 3
Veldkamp (1999) reported Neyraudia arundinacea (L.) Henr. var. zollinger (Buse) Henr. as a
new record for Singapore (Wong W.P. s.n., August 1959, Bartley Road, SINU). It has not been
found since. This grass with a reed-like habit and silky-hairy panicles is distinguished from
Phragmites by the hairy lemmas. It also grows in drier places than the latter (Gilliland 1971).
Oplismenus
Oplismenus is characterized by awned glumes and ovate to lanceolate leaf blades (Gilliland
1971). The presence in Singapore of O. burmanni (Retz.) P. Beauv. and O. compositus (L.) P.
Beauv. was mentioned by Veldkamp (1999). Besides the specimens cited by him, I have seen
the first in Nee Soon forest (Duistermaat S166, 2003, SING). The two are best distinguished
by the type of awn: antrorsely scaberulous and filiform in O. burmanni vs. smooth and rather
thick in O. compositus.
Gardens’ Bulletin Singapore 56 (2004) 43-54. 43
Curcuma mutabilis (Zingiberaceae): a New Species from
South India
JANA SKORNICKOVA, M. SABU & M.G. PRASANTHKUMAR
Department of Botany, Calicut University, 673 635 Kerala, India
Abstract
Curcuma mutabilis Skornickova, M.Sabu & M.G.Prasanthkumar, sp. nov. is described from
Northern Kerala, South India, including illustrations and notes.
Introduction
The genus Curcuma L. is of great economic and ornamental importance and
several species such as C. longa L., C. aromatica Salisb., C. zanthorrhiza
Roxb., C. alismatifolia Gagnep., etc., can be found all over the tropics
introduced, cultivated and sometimes naturalized. The genus is distributed
mostly in tropical Asia with a few species extending to Australia and the South
Pacific. Larsen et al. (1998) stated there are about 80 species, Sirirugsa (1996)
estimated the number at around 100. Presently there are 103 validly published
names (excluding those that have been transferred to other genera) and another
6 species were published only as nomen nudum. Without a doubt, a number of
names will turn out to be synonyms, but as recent field exploration in India
and SE Asia proceeds, the number of species will probably reach 120. Three
new species were recently described from SE Asia (Sirirugsa & Newman,
2000; Mood & Larsen, 2001). In India, the genus is represented by 29 species
(Karthiykeyan et al., 1989; Jain and Prakash, 1995) and has been revised for
South India by Mangaly and Sabu (1993). Since then exploration of remote
areas in India has brought to light another three new species (Tripathi, 2001;
Skornickova et al., 2003a, b).
While working for the project ‘Revision of Indian Zingiberaceae’, the
authors encountered an interesting seed-setting Curcuma species from
Nilambur, North Kerala. At the beginning of the monsoon season, the plant
resembled C. oligantha Trimen in lacking an obvious coma, by the inflorescence
appearing before the first leaf expands, the flowers being exerted from the
bracts, and in rhizome shape. However, critical studies of the plants observed
at different times in relation to the monsoon season revealed that the species is
different from C. oligantha, especially in its general habit and size, shape and
coloration of the bracts, and size and shape of flower parts.
Subsequently, we discovered that Velayudhan et al. (1999) had described
this species as C. nilamburensis based on a collection from the same locality.
44 Gardens’ Bulletin Singapore 56 (2004)
Unfortunately, this name is not validly published according to the St Louis
Code (Greuter et al., 2000) because the description lacked a Latin diagnosis
and a type was not designated. In addition, the publication by Velayudhan et
al. (1999) is of limited circulation, consequently, the species is described
appropriately below.
Curcuma mutabilis Skornickova, M. Sabu & M.G. Prasanthkumar sp. nov.
(Fig. 1, Plate 1.)
C. nilamburensis in Velayudhan K.C., V.K. Muralidharan, V.A. Amalraj, P.L.
Gautham, S. Mandal & D. Kumar, Curcuma Genetic Resources. 1999. p. 42.
nom. illegit.
C. neilgherrensis auct. non Wt.: Sivar. & P. Matthew, Fl. Nilambur. 1996. p.
708.
Curcumae oliganthae Trimen in coma inconspicua rhizomatis figura magnitudo
colorque similis, habitu maiore robustiore (60 cm longo), bracteis fertilibus
ovato-ellipticis in dimidio inferiore connatis apice rodundato macula fusce
brunneo-violacea proviso, floribus bracteis minus exsertis, labello minore (14-
16 mm longo), staminodiis lateralibus brevioribus (13-16 mm longis) corollae
lobo dorsali non multo excedentibus, antherarum calcaribus sursum versis
differt. Typus: India, Kerala, Malappuram District, Nilambur, Skornickova &
Prasanthkumar 84145 (holo MH; iso K, CALI, SING).
Rhizomatous herb, 10—60 cm tall. Rhizome ovoid without branches, cylindrical
to conical, up to 5 x 2.5 cm, deeply buried in the ground, light brown externally,
glabrous, sheathed by papery brown scales and bases of the leaf sheaths, which
leave vertical scars after decaying, creamy yellowish internally, faintly aromatic.
Roots fleshy ending in ovoid root tubers 2-4 x 1—2 cm distanced 2-15 cm
from the main rhizome, externally light brown, glabrous, pearly white inside,
non-aromatic. Pseudostem 5—30 cm long, deeply buried in the soil, formed by
leaf sheaths and 34 sheathing bracts, green or with ared tinge, drying towards
the end of season and becoming brown and papery, ligule 3-4 mm, bilobed,
translucent greenish, hairy outside and on the margin, hairs 0.2—0.3 mm; leafy
shoot 15-60 cm long. Leaves at the beginning of the season 1—2 with a short
petiole or even almost sessile, later with up to 7 leaves, gradually with longer
petioles up to 20 cm long, petiole green or with a red tinge, glabrous or shortly
Plate 1. Curcuma mutabilis
1. Inflorescence of a yellow-flowered plant (seen from above); 2. Detail of arillate seeds;
3. Inflorescence of a white-flowered plant; 4. Variability of inflorescences; 5. Habit at type
locality; 6. The whole plant including rhizome. All photographs are of the type material
Skornickova & Prasanthkumar 84145 with lateral inflorescences at the beginning of season.
Photos J. Skornickova.
Curcuma mutabilis (Zingiberaceae): a New Species from South India
46 Gardens’ Bulletin Singapore 56 (2004)
hairy; lamina ovate-elliptic, 14-35 x 7-11 cm, adaxially deep green,
prominently veined, prominent veins quite closely arranged c. 5 mm apart,
sulcate in between, hairy on the prominent raised veins and towards the margin
especially in the upper half of the lamina, hairs c. 0.3 mm long (rarely almost
glabrous on the upper surface), abaxially paler green, usually glabrous, rarely
densely velvety pubescent; margin hyaline, translucent white, c. 0.2 mm wide,
hairy in the apical part and glabrous in the distal part of lamina, tip c. 0.5 cm
long, acuminate, densely hairy, base attenuate to slightly cordate, oblique,
midrib green, glabrous. /nflorescence lateral at the start of the season, central
later in the season. Peduncle 4—20 cm long, 2.5—8 mm diam., glabrous, whitish,
light green or with a red tinge, peduncle of the vernal inflorescence sheathed
by bracts, central one hidden within the pseudostem. Spike 3.5—15 x 2.5—5 cm,
consisting of 6—70 bracts. Coma inconspicuous, usually only the uppermost 2
or 3 bracts are sterile and more linear than the fertile ones, 2.5—3.5 x 0.7—1.5
cm, light green or with a red tinge (sometimes deep red), upper side sparsely
hairy, lower side glabrous, tips rounded with a deep violet brown patch. Fertile
bracts ovate-elliptic with visible parallel veinlets, tip obtuse or slightly
acuminate, both sides glabrous, connate in the lower half, 2.5-3.5 x 1.5—2.7
cm, whitish, light green, green or with a red tinge verging to brown red, but all
bracts (including uppermost sterile ones) always with a dark violet mauve tip,
which is usually larger in the upper bracts and less conspicuous in the lower
ones. Cincinni with 2-4 flowers. Bracteoles one per flower, 5—7 x 24 mm,
hyaline, translucent white or with a pink or a red tinge, almost glabrous or
puberulus with a few hairs 0.2—0.3 mm long on the tip. Flowers 4.5—5 cm
long, exserted from bracts. Calyx 8—11 mm long, 3-toothed, unilaterally split
3—4.5 mm deep, translucent white or tinged with pink or dark violet, quite
glabrous, but with sparsely hairy teeth and sometimes along the upper part of
the vein leading from teeth to the base. Corolla tube c. 2.5—3 cm, at the base
white, yellowish or yellow, towards the lobes sometimes tinged with pink, red
or violet, glabrous; dorsal corolla lobe c. 1.4—1.8 x 1—1.4 cm, triangular-ovate,
concave, apex mucronate, mucro 1—2 mm long, glabrous, varying from white,
yellowish or yellow, sometimes tinged pink red or deep violet-bluish, lateral
corolla lobes \.4—1.7 x 0.6—-0.8 cm, triangular with rounded, slightly concave
tip, glabrous, with the same coloration as the dorsal lobe but less intensive,
usually overlapping in the tip portion on the ventral side of the flower. Lateral
staminodes obovate-rhomboid, 1.3—1.6 x 1—1.2 cm, usually yellow or yellow
with a reddish base, less frequently white or white with a yellow or reddish
base, glandular hairs present on the raised middle portion. Labellum 1.4—1.6 x
1.5—1.9 cm, emarginate, split 3-6 mm long (opening wider as the flowers age
and wilt), yellow or less frequently white, centre deep yellow or yellow, rarely
also white, base of labellum usually yellowish, rarely with a deep red patch or
tinge. Anther spurred, glandular hairs present on the sides and back part, anther
Curcuma mutabilis (Zingiberaceae): a New Species from South India 47
Simm
2mm
Simm
2mm
\ , 1mm
3mm
~ ae
— a
=.=
Figure 1. Curcuma mutabilis
a. Habit; b. Flower (side view); c. Bracteoles; d. Dorsal corolla lobe; e. Lateral corolla
lobe; f. Labellum; g. Lateral staminode; h. Anther (front); 1. Anther (side); j. Calyx; k.
Ovary and epigynous glands; I. Ovary (cross section); m. Seed. Based on the type
material Skornickova & Prasanthkumar 84145. Del. J. Skornickova.
48 Gardens’ Bulletin Singapore 56 (2004)
thecae whitish, 3.5—4 mm long; filament 3-4 mm, light yellow, yellow or with
deep red tinge, constricted, 3 mm broad at base, 2 mm broad at upper part.
Anther spurs 2—2.5 mm long, white or creamy yellowish, pointing upwards.
Anther crest present, roundish to slightly truncate, c. 1.5 mm broad and 0.5
mm long. Ovary trilocular, 3—3.5 x 2.5-3 mm, hairy, white pubescent, hairs
0.3-0.4 mm long, ovules many. Stigma 1-1.4 x 0.9-1.1 mm, white, ciliate,
exserted 1—2.5 mm. Epigynous glands 2, creamy yellow, 3—5 mm long, 0.5—
0.6 mm diam. Fruit a dehiscent capsule, spherical, c. 1 x 1 cm. Seeds 3 x 2
mm, light brown, shiny glabrous, non-aromatic, aril translucent white, laciniate,
lobes up to 6 x 0.7—1.5 mm.
Distribution: India: Nilambur in Kerala, so far known only from the type locality
and adjacent areas.
Habitat: In lowland areas c. 50-100 m asl, in the undergrowth of teak
plantations, secondary forests and shrubby vegetation on dry lateritic soil.
Flowering and fruiting: Lateral inflorescences appear in May to June just before
or simultaneously with the leaves. Terminal (central) spikes are produced in
September provided the monsoon was sufficient. Fruiting occurs about 3-4
weeks after flowering.
Etymology: The specific epithet refers to the extremely variable colour of
floral parts (corolla lobes, labellum and lateral staminodes) and size of the
plants.
Notes: Curcuma mutabilis appears to be closely related to C. oligantha Trimen
in having an inconspicuous coma and in its ovoid rhizome internally creamy
yellow and without branches, but it differs in the following salient characters.
Most remarkable is the more robust habit (up to 60 cm tall), the ovate-elliptic
lamina and the size of inflorescence that consists of 6—70 bracts. The ovate-
elliptic bracts and prominent dark violet brownish patch at their rounded tips
as well as the fact they are connate in the lower half make this species readily
recognizable from C. oligantha, which rarely exceeds 15—20 cm in height, has
ovate-lanceolate laminas and its inflorescences consist of 5—10 (rarely up to
15) lanceolate bracts, which are connate in the lower quarter or even less and
have no prominent coloration at the tips that are acuminate. Even though C.
mutabilis is more robust in its vegetative parts, the flowers are overall smaller
(up to 5 cm long with the calyx 8—11 mm long, lateral staminodes up to 1.6 x
1.2 cm and which do not protrude much beyond dorsal corolla lobe) compared
with those of C. oligantha (5-7 cm long, calyx 15-21 mm long, lateral
staminodes up to 3 x 1.4 cm prominently protruding beyond the dorsal corolla
Curcuma mutabilis (Zingiberaceae): a New Species from South India 49
lobe, which make flowers of C. oligantha much more exserted out of the fertile
bracts than those of C. mutabilis). Notable also is the difference in anther
spurs, which point upwards in C. mutabilis, but downwards more or less
following the anther thecae direction in C. oligantha. Further details in
morphological characters and measurements of C. mutabilis and C. oligantha
are compared in detail in Table 1. More details on C. oligantha were published,
for example, by Trimen (1885, 1898a), Burtt and Smith (1983) and Bhat (1987).
There is a colour plate of C. oligantha in Trimen’s Handbook to the Flora of
Ceylon (1898b).
Great variability in several Curcuma species have already been observed
by others, for example, C. pseudomontana Graham (Santapau 1945, 1952), C.
aurantiaca Zijp (Valeton, 1918), and during the past several years we have
observed the same feature in several other species (C. oligantha Trimen, C.
inodora Blatt., C. neilgherrensis Wt.). All these species share in common
propagation by seed. Most of such species have ovoid rhizomes without
branches (usually referred as sessile or palmate tubers), and thus there is no
provision for vegetative propagation.
Many Curcuma species do not set seed and reproduce vegetatively by
the rhizome branches (e.g., C. aeruginosa Roxb., C. amada Roxb., C. longa
L., C. zanthorrhiza Roxb.). Very rarely in some of these species formation of
fruit can be observed and, even more rarely, the presence of developed seeds.
These species are morphologically quite uniform not only within population,
but also between remote populations. Root tubers, which are present in both
seed-setting and non-seed-setting Curcuma species, are not capable of sprouting
and functioning to sustain the plant during periods without rain, when the
leafy shoot dries up.
Like other seed-setting species, C. mutabilis possesses high variability
within populations as already mentioned by Sivarajan and Matthew (1996). In
fact, C. mutabilis is one of the most variable species we have encountered in
the genus. Most remarkable is the difference in flower colour. Corolla lobes
can vary from whitish pink, pink-red, reddish orange, dark pink to dark violet;
labellum and lateral staminodes can be pure white, white with a yellow or
reddish tinge in the throat of the labellum or base of the lateral staminodes or
different shades ranging from creamy, light yellow to deep yellow. The colour
of the corolla and staminodes varies independently and in fact hardly any two
individuals look identical. The leaves are also highly variable regarding
pubescence. All these differences can be observed within a few square metres.
Also remarkable is the difference in size of the plants, which ranges from a
few centimetres to about 60 cm tall, as well as in the number of the bracts,
which can vary from 6—70. Size and number of bracts seems to be correlated
with the age of the plant, which is a feature we have observed in some other
seed-setting Curcuma species.
50 Gardens’ Bulletin Singapore 56 (2004)
This great variability might suggest that hybridisation has occurred.
However, it can be ruled out in the case of C. mutabilis because the pollen is
fully formed (although it has not been tested yet for viability), seed is viable
and readily germinates, and many seedlings are observed in wild populations,
the range of variation is similar in the separate populations and there are no
species within this area that could be putative parents.
The species that appeared under the name Curcuma neilgherrensis Wt.
in the Flora of Nilambur (Sivarajan & Matthew, 1996) is C. mutabilis and not
C. neilgherrensis, which is a completely different species that grows in high
altitude grasslands (700-1200 m) and which is common on southwest slopes
of the southern range of Neilgherries. Even though C. neilgherrensis is also
seed-setting and quite a variable species, especially in size of the plant and
inflorescence, coma colour (pinkish to deep pink) and indumentum of leaves,
it can easily be distinguished from C. mutabilis by its lanceolate and firm
leaves, spikes with a prominent coma, green fertile bracts with no dark violet
patch and yellow flowers, which are not exserted out of the fertile bracts. In
contrast, C. mutabilis is lowland species (growing at 50-100 m altitude) found
in the undergrowth in shady places and it has rather thin, ovate-elliptic leaves,
with prominent sulcate venation, an inconspicuous coma, bracts tipped by dark
violet brown patch and the flowers (highly variable in colour) are exserted out
of the fertile bracts. We have so far not observed such colour variability of
flower parts in the C. neilgherrensis.
Table 1. Comparison of morphological characters of Curcuma mutabilis and
its closest ally, C. oligantha Trim. Diagnostic characters are in bold.
Rhizome
Leafy shoot
Main rhizome ovoid,
unbranched, up to 5 x 2.5 cm.
Inwardly creamy-yellowish
colour, faintly aromatic.
Root tubers 2-4 x 1—2 cm,
pearly white inside.
Leafy shoot to 60 cm tall
with 2-7 leaves.
Pseudostem and peduncle
sheathed by green or reddish
green bracts.
Main rhizome ovate-conical,
unbranched, up to 5 x 1.5 cm.
Inwardly creamy white-
yellowish colour, no obvious
aroma.
Root tubers 1.5—4 x 0.8-1.5
cm, pearly white inside. .
Leafy shoot to 20 cm tall
with 2-6 leaves.
Pseudostem and peduncle
sheathed by green or reddish
green bracts.
Curcuma mutabilis (Zingiberaceae): a New Species from South India
Table 1. Continued:-
Leafy shoot
Lamina
Inflorescences
Flowers
Ligule 34 mm, 2-lobed,
hairy along the margin,
hairs 0.2-0.3 mm long.
Lamina ovate-elliptic, 14—35
x 7-11 cm, base attenuate-
slightly cordate, oblique.
Adaxially deep green,
prominently sulcate, hairy
along the main veins
especially at the distal half of
the lamina and near margins
(rarely some individuals
almost glabrous), abaxially
pale green, usually glabrous,
rarely individuals densely
velvety hairy.
Vernal, lateral, later in season
central.
Peduncle 4—20 cm, spike c.
3.5-15 x 2.5-—5 cm,
comprised of 6—70 bracts.
Coma inconspicuous, few
uppermost bracts sterile and
more linear.
Bracts ovate-elliptic with
rounded tip, light green ,
green or with red tinge with
dark brownish-violet patch at
the taps) c:2-5-—3.5 x 1.5—2.7
cm, connate in lower half.
Cincinni with 2-4 flowers.
Flower 4.5—5 cm long,
exserted from the fertile
bracts.
Bracteoles 5—7 x 2-4 mm,
hyaline, translucent white or
pinkish with red tinge,
almost glabrous.
51
Ligule 1-1.5 mm, obscurely
2-lobed, glabrous.
Lamina ovate-lanceolate, 7—
19 x 3.2—7 cm, base
attenuate-slightly cordate,
oblique. Adaxially green,
glabrous or sparsely hairy,
always hairy along the main
veins especially at the distal
half of the lamina and near
margins, abaxially pale green,
glabrous.
Vernal, lateral, later in season
central.
Peduncle 4—11 cm, spike c.
3—6 x 2-4 cm, comprised of
4-10 (rarely 15) bracts.
Coma inconspicuous.
Bracts lanceolate, erect with
acuminate tip, light green or
| green, rarely with reddish
tinge, no obvious patch at the
tips, c.2.5—-4 x 1-2 cm,
connate in lower quarter or
even less. Cincinni with 2-3
flowers.
Flower 5-7 cm long, highly
exserted from the fertile
bracts.
Bracteoles 5—9 x 3.5—4 mm,
hyaline, whitish translucent,
usually hairy at the tip.
52
Table 1. Continued:-
Flowers
Calyx 8-11 mm long, 3-
toothed, unilaterally split
3—4.5 mm, translucent white
or tinged with pink or tinged
with dark violet, sparsely
hairy.
Corolla tube 2.5—3 cm,
glabrous, white or yellow at
base, towards lobes
sometimes tinged with red.
Labellum 1.4—1.6 x 1.5-1.9
cm, emarginate, yellow.
white with yellow center or
white, with or without red
tinge at the base.
Lateral staminodes 1.3-1.6 x
1-1.2 cm, yellow, white with
yellow or white with or
without red tinge at the base.
Anther thecae 3.5-—4 mm
long, anther spurs 2-2.5
mm white or creamy,
pointing upwards.
Ovary 3-3.5 x 2.5—3 mm,
hairy.
Seeds brown, glossy with
laciniate aril.
Epigynous glands, c. 3-5 mm
long, 0.5—0.6 mm diam.,
creamy yellow
Gardens’ Bulletin Singapore 56 (2004)
Calyx 15-21 mm long, 3-
toothed, unilaterally split
6-10 mm, translucent
greenish-white or tinged
with pink, shortly hairy.
Corolla tube 1.7—3.5 cm,
shortly hairy, hairs 0.1—0.2
mm, appressed, yellow or
white, with or without red
tinge.
Labellum 1.8-1.2 x 1.5-2.2
cm, emarginate, yellow.
white with yellow center or
pure white.
Lateral staminodes 2.2-3 x
1-1.4 cm, yellow or white
greatly exceeding beyond
dorsal corolla lobe.
Anther thecae 4-5 mm
long, anther spurs c. 3mm
white or yellow, pointing
downwards following
more or less anther thecae
direction.
Ovary 3—3.5 x 2—3 mm,
hairy.
Seeds brown, glossy with
laciniate aril.
Epigynous glands, c. 2.54
mm long, c. 0.4 in diam.,
yellow or creamy.
Acknowledgements
The authors thank the Department of Science and Technology, Govt. of India
for financial support (Order No. SP/SO/A-20/99 dt. 09.11.2001). We are also
indebted to authorities of Ministry of Environment and Forests for granting
permission for collecting. The first author thanks the Indian Council for Cultural
Relations, New Delhi, India, and Ministry of Education of the Czech Republic
for awarding a research fellowship; to Grant Agency of the Academy of
Curcuma mutabilis (Zingiberaceae): a New Species from South India 53
Sciences of the Czech Republic (Grant No. B6407401) and International
Association of Plant Taxonomy for financial support; to the Singapore Botanic
Gardens for providing facilities during her repeated visits; to Dr. J.J. Vermeulen
(SING) for his help and comments on drawing, to Dr. R. Kiew (SING) and to
Dr. Axel D. Poulsen (E) for comments on the manuscript. We also thank Dr. J.
F. Veldkamp, National Herbarium of Netherlands, Leiden University Branch,
for the Latin diagnosis.
References
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Journal of Forestry. 10: 66-68.
Burtt, B.L. and R.M. Smith. 1983. Zingiberaceae. In: Dassanyake, M.D. and F.R.Fossberg
(Eds.). A Revised Handbook to the Flora of Ceylon. Amerind. Pub. Co. New Delhi, India.
4: 488-5372.
Greuter, W., F.R. Barrie, H.M. Burdet, W.G. Chaloner, V. Demoulin, T.S. Filgueiras, D.H.
Nicholson, P.C. Silva, J.E. Skog, P. Trehane, N.J. Turland and D.L. Hawksworth. 2000.
International Code of Botanical Nomenclature (St Louis Code) adopted by the Sixteenth
International Botanical Congress St Louis, Missouri, July-August 1999. Regnum Vegetabile.
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Jain, S.K. and V. Prakash. 1995. Zingiberaceae in India: Phytogeography and Endemism.
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Karthikeyan, S., S.K. Jain, M.P. Nayar and M. Sanjappa. 1989. Florae Indicae Enumeratio
Monocotyledoneae. Botanical Survey of India, Calcutta, India. pp. 289-299.
Larsen, K., J.M. Lock, H. Maas and P.J.M. Maas. 1998. Zingiberaceae. In: K. Kubitzki (Ed.)
The Families and Genera of Vascular Plants IV. Springer Verlag, Berlin, Heidelberg,
Germany. pp. 474-495.
Mangaly, J.K. and M. Sabu. 1993. A taxonomic revision of the South Indian species of Curcuma
L. (Zingiberaceae). Rheedea. 3: 139-171. ;
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Santapau, H. 1952.Onacommon species of Curcuma of Bombay and Salsette Islands. Journal
of the Bombay Natural History Society. 51: 135-139.
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Zingiberaceae. Zhongshan University Press. Guangzhou, China. pp. 39-46.
Sirirugsa, P. and M. Newman. 2000. A new species of Curcuma L. (Zingiberaceae) from S.E.
Asia. The New Plantsman. 6: 196-197.
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Sivarajan, V.V. and P. Matthew. 1996. Zingiberaceae. Flora of Nilambur (Western Ghats,
Kerala). Bishen Singh Mahendra Pal Singh, Dehra Dun, India. pp. 702-716.
Skornickova, J. and M. Sabu. 2002. The genus Curcuma L. in India: Resume and Future
Prospects. In: A.P. Das (Ed.) Perspectives of Biology, Bishen Singh Mahendra Pal Singh,
Dehra Dun, India. pp. 45-51.
Skornickova, J., M. Sabu and M.G. Prasanthkumar. 2003a. New species of Curcuma from
Mizoram. Gardens’ Bulletin Singapore. 55: 89-95.
Skornickova, J., M. Sabu and M.G. Prasanthkumar. 2003b. Curcuma codonantha
(Zingiberaceae)—A new species from Andaman Islands, India. Gardens’ Bulletin Singapore.
55: 219-228.
Trimen, H. 1885. Curcuma oligantha. Notes on the flora of Ceylon. Journal of Botany. 23:
245.
Trimen, H. 1898a. Scitamineae. A handbook to the Flora of Ceylon. 4: 238-265.
Trimen, H. 1898b. Curcuma oligantha. A handbook to the Flora of Ceylon—Atlas of Plates.
Pi. XE:
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Gardens’ Bulletin Singapore 56 (2004) 55-72. 55
A Tropical Seasonal Rain Forest at its Altitudinal and
Latitudinal Limits in Southern Yunnan, SW China
H. ZHU
Xishuangbanna Tropical Botanical Garden,
The Chinese Academy of Sciences,
Xue Fu Road 88,
Kunming 650223, Yunnan, P. R. China
Abstract
The tropical seasonal rain forest (semi-evergreen rain forest) in Caiyanghe Nature Reserve,
Simao, Yunnan, China, is at the northern and altitudinal limits of tropical rain forest in Asia.
It shares the main physiognomic characteristics of tropical rain forest. Compared with lowland
rain forests in China and equatorial regions in Asia, it has lower species diversity with more
micro- and nano-phanerophytes and herbaceous phanerophytes in the life form spectra, more
microphyllous plants and more plants with non-entire leaves. In floristic composition, it is
dominated by tropical Asian elements and characterized by the Indo-Malayan tropical flora.
It is suggested that the occurrence of tropical rain forest in southern Yunnan is more influenced
by topography and local habitats than the regional climate.
Introduction
Tropical rain forests occur mainly in those wet tropical areas between the
Tropics of Cancer and Capricorn with a mean annual temperature above 24°C
and the mean monthly temperature above 18°C in the coldest month; and an
annual rainfall of at least 1700 mm and usually above 2000 mm (Richards
1996). However, the northern and southern borders of tropical rain forests do
not coincide exactly with any latitudinal limits and in some places they extend
somewhat beyond the Tropics of Cancer and Capricorn. For example, the Indo-
Malayan rain forests extend beyond the Tropic of Cancer at 27° 31°N in
northeastern India (Proctor et al., 1998), Myanmar (Kingdon- Ward 1945) and
southern Yunnan, China (Zhu 1997), where they occur as small patches forming
mosaics within the monsoon forests (Whitmore 1990; Richards 1996; Morley
2000). These Indo-Malayan tropical rain forests north of the Tropic of Cancer
are little known apart from some descriptive accounts (Kingdon-Ward 1945;
Proctor et al. 1998).
Southernmost Yunnan, i.e. Xishuangbanna, which borders Myanmar and
Laos, is a mountainous area on the northern margin of tropical Southeast Asia.
Tropical rain forest in this region was first mentioned by Wang (1939). Between
1955 and 1960, these forests in southern Yunnan were investigated in detail by
56 Gardens’ Bulletin Singapore 56 (2004)
Sino-Russian expeditions and Yunnan University. It was then suggested that
biogeographically tropical rain forests existed in southern Yunnan, but these
were considered to be a type different from the ones in Indo-Malaysia because
of the lack of representatives of Dipterocarpaceae, which dominates the rain
forests of tropical SE Asia (Fedorov 1958; Qu 1960; Wang 1961). Since the
tropical rain forest in Xishuangbanna occurs at the climatic limits of rain forest
(lower mean annual temperature and annual precipitation than usual) and has
a clear change of physiognomy between different seasons, Chinese botanists
prefer the term ‘tropical seasonal rain forest’ (Wu 1987; Zhu 1992; Cao 1996;
Jin 1997), although it conforms to the semi-evergreen rain forests of SE Asia
as defined by Whitmore (1984, 1990) and to a lowland rain forest (Zhu 1997).
Although several studies on the tropical rain forests in southern Yunnan
have been carried out (Wu 1987; Xu et al. 1987; Jin 1997; Zhu et al. 1998a,
2002), little has been published in English (Cao & Zhang 1997; Zhu 1997;
Zhu et al. 1998b) and most of these focused on the tropical seasonal rain
forest in Xishuangbanna where they occur only in areas below 900 m asl.
Recently, tropical seasonal rain forest was found in the Caiyanghe Nature
Reserve, Simao County, just north of Xishuangbanna at an altitude of 1300 m
and for which Zhu et al. (2000) provided a descriptive report. However, the
occurrence of tropical seasonal rain forest here is unusual more for its high
altitude than its northerly latitude. Its floristic composition and physiognomy
are presented in this paper.
The Caiyanghe Nature Reserve
Location and topography |
The Nature Reserve is in southeast Simao County, Yunnan, 101°7°~ 101°15° E
22°30°~22°38'N (Fig. 1). It has a mountainous topography with its main range
running east to west with the highest elevation (1698 m) in the north and the
lowest elevation (980 m) on the Caiyanghe River on the southwest slope of
the mountain.
Climate and Soil
There are no meteorological records for Caiyanghe, but from the records from
Simao county climate station c. 20 km away, the region has a monsoon climate
with a mean annual temperature of 17.7°C, a mean for the coldest month of
11.4°C, annual temperature accumulation (the sum of daily temperature means
of > 10°C) of 6253°C and mean annual precipitation of 1547.6 mm, of which
more than 80% falls during the rainy season between May and the end of
October.
Tropical Seasonal Rainforest From Southern Yunan 57
96° 97° 98° 5oF 100° 101° 102° 103° 104° 105°
aSt
rh
24°
eff
aoe
Myanmar
gees
Vietnam
Kilometers
pare,
21°
alZ
a0Z
19°
eG
18°
3
96° 97° 98° 99° 100? 101° 102? 103? 104? 105°
Figure 1. The location of Caiyanghe Nature Reserve in Simao, southern
Yunnan, Southwestern China
The soil is oxisol with a deep solum but a thin humus horizon. The oxisol
was derived from sandstone (above 1200 m asl) and from muddy shale (below
1300 m asl).
Vegetation types
The main vegetation formation at Caiyanghe is a montane evergreen broad-
leaved forest_in terms of physiognomy and habitat, which was called monsoon
evergreen broad-leaved forest in Chinese literature (Wu 1987). It occurs on
mountain slopes and summits above 1200 m asl and in valleys at 1300~1500
m altitude, and makes up 92% of the natural forest cover in the Nature Reserve
(Cao, 2003).
The focus of this study was on the tropical seasonal rain forest in
Caiyanghe, which makes up less than 2% of the natural forest cover in the
Nature Reserve and which is dominated by Pometia tomentosa (Sapindaceae)
and Garuga floribunda var. gamblei (Burseraceae). It occurs only in valleys
below 1300 m altitude and usually on south-facing slopes. The tropical seasonal
rain forest in Caiyanghe is an extension of the ones in Xishuangbanna.
58 Gardens’ Bulletin Singapore 56 (2004)
Methods
Several small patches of tropical seasonal rain forest occur in the Nature
Reserve. A representative stand in a valley was selected for a plot study. The
plot size was 25 m x 100 m (with the plot length along the valley). All trees in
the plot were identified and their dbh (minimum 5 cm), height and crown cover
were measured. The plot was divided into five subplots (25 m x 20 m each) so
that frequency of tree species could be calculated. The forest profile was drawn
from a strip stand of 50 m x 10 min the plot. Five small plots of 5m x 5m
each were set up for a floristic survey of the understorey. In these small plots,
saplings, shrubs and herbaceous plants were counted. Epiphytes and lianas in
the plot were identified and their abundance estimated visually. Importance
value indices (IVI) for tree species were calculated (Curtis & McIntosh 1951).
A rank/abundance diagram (Magurran 1988) of species in the plot was plotted.
Shannon-Wiener’s indexes for species diversity (Shannon and Wiener 1949)
were calculated from the plot data. Physiognomy (life forms and leaf size) was
analyzed using Raunkiaer’s criteria (1934). Specimens were collected and
identified. Species authorities follow Flora Reipublicae Popularis Sinicae (Flora
of China). Specimens are deposited in the herbarium of Xishuangbanna Tropical
Botanical Garden, Chinese Academy of Science. 3
Results
Species composition
The tropical seasonal rain forest in the Nature Reserve can be recognized as a
Garuga floribunda var. gamblei—Pometia tomentosa formation based on the
dominant species. It occurs in the valleys below 1300 m altitude. It is c. 35 m
tall with three tree layers (Fig. 2). The top tree layer, composed of emergent
trees with buttresses, is 25~35 m tall and has a crown cover of 40~50%. The
most dominant tree species in this layer is Pometia tomentosa. The second tree
layer is 10~25 m tall with a crown cover of 50-60%. The most dominant tree is
Alphonsea monogyna (Annonaceae). The third tree layer is 3~10 m tall with a
crown cover of 50% and Cleidion spiciflorum (Euphorbiaceae) is the most
dominant species. Some species in the tree layer are cauliflorous and many
have leaves with a drip tip. The sapling-shrub layer is 1~3 m tall with a cover
of 30~40 %. It is composed mainly of juvenile trees. The commonest shrub
species is Mycetia glandulosa (Rubiaceae). The herb layer is 1 m tall with a
cover of c. 25%. The woody lianas are abundant, but there are relatively few
epiphytes. |
Evergreen trees both in species and individuals dominate the forest. Only
two deciduous species were recorded from the forest plot. Garuga floribunda
Tropical Seasonal Rainforest From Southern Yunan 59
Figure 2. Profile diagram of the tropical seasonal rain forest in Caiyanghe
] Pometia tomentosa, 2 Cleidion spiciflorum, 3 Alphonsea monogyna,
4 Cinnamomun bejolghota, 5 Phoebe lanceolata, 6 Elaeocarpus sikkimensis,
7 Dead tree, 8 Garuga floribunda var. gamblei, 9 Saurauia macrotricha,
10 Baccaurea ramiflora, 11 Macropanax dispermus, 12 Flacourtia rukam,
13 Duabanga grandiflora, 14 Phoebe puwenensis
var. gamblei, although not dominant in individuals, is the largest deciduous
tree in canopy layer, and Radermachera igneum (Bignoniaceae) in the second
tree layer. 7
The species composition and their phytosociological importance are
given in the Appendix.
Physiognomy
The forest in Caiyanghe is characterized by phanerophytes, of which woody
phanerophytes contribute the most (Table 1). Among 52 woody plant species
from the plot, ones with mesophyllous leaves contribute 69.2% of the total,
ones with microphyllous leaves 17.3% and ones with macrophyllous leaves
13.5%.
Compared with tropical seasonal rain forests at lower elevations in
Mengla and Mengyang of Xishuangbanna (Zhu et al. 1998b), the forest in
60 Gardens’ Bulletin Singapore 56 (2004)
Table 1. Life form spectrum of the plants from the tropical season rain
forest in Caiyanghe.
Life form Epiphyte Liana Woody plants Herb plants Total
Megaph. Mesoph. Microph. Nanoph. Hph. Ch
No. of species 2 7 9 20 18 5 = ae. 81
% 2.5 8.6 11.1 24.7 teed 6.2 18.5 6.2 100
Megaph.: Megaphanerophytes; Mesoph.: Mesophanerophytes; Microph.: Microphanerophytes;
Nanoph.: Nanophanerophytes; Hph.: Herbaceous phanerophytes; Ch: Chamaephytes
Caiyanghe has fewer liana species, but more micro- and nano-phanerophytes,
and herbaceous phanerophytes (Fig. 3), Caiyanghe rain forest has a similar
leaf size spectrum to seasonal rain forests in Xishuangbanna and lowland
evergreen rain forest in northeastern India, which show a higher percentage of
microphyllous plants than those of equatorial lowland rain forests in Borneo
and the Philippines (Fig. 4). Like the lowland evergreen rain forest in
northeastern India, the rain forest in Caiyanghe also has a higher percentage of
species with simple leaves and non-entire leaves (Table 2). A relatively high
proportion of plants with non-entire leaf margins could be a character of the
rain forest either at latitudinal or altitudinal limits because a higher proportion
of non-entire leaved species is usually present in subtropical evergreen broad-
leaved forest in eastern Asia and in tropical montane forest in southeastern Asia.
Tropical Seasonal Rainforest From Southern Yunan 61
S&S Ep
(41 Ph
# Ch
HG
Species %
LHSR RSR SRC
Forest type
a. For all plant species
= Hph
40
# Lph
gf. (30 "| Mega-Mesoph
o 1! Micro-Nanoph
3 20
op)
LHSR RSR SRC
Forest type
b. For phanerophytes
Figure 3. Comparison of life form spectra from the tropical seasonal rain forest in
Caiyanghe and the tropical seasonal rain forests at lower altitude in southern Yunnan
In Caiyanghe - SRC: Seasonal rain forest. In southern Yunnan (Zhu et al., 1998b)
- LHSR: Lower hill seasonal rain forest; RSR: Ravine seasonal rain forest.
Ep=Epiphyte; Ph=Phanerophyte; Ch=Chamaephytes; G=Geophyte; Lph=Liana-
phanerophyte
Hph=Herbaceous phanerophyte; Mega-Mesoph=Megaphanerophyte +
Mesophanerophyte; Micro-Nanoph=Microphanerophyte + Nanophanerophyte
62 Gardens’ Bulletin Singapore 56 (2004)
I
UT
{Ui
ie s ia
* | | | i Nano-Micro.
“ | | | = Meso.
a | | | = Macro.
| a os Gigan.
|
wa _—
FB
WN
v2)
‘o)
eS
v2)
—
og
v2)
Ty
e
v2)
Forest type
Figure 4. Comparison of leaf size spectra from tropical rain forests in
Caiyanghe, southern Yunnan, India and equatorial lowlands
Caiyanghe - SRC; Southern Yunnan - LHSR: Lower hill seasonal rain forest
(Zhu , 2000); RSR: Ravine seasonal rain forest (Zhu et al., 1998b); India -
LRI: Lowland evergreen rain forest (Proctor et al., 1998); Equatorial lowlands
- LRF: Lowland rain forest in the Philippines (Brown,1919); LRFB: Lowland
rain forest in Borneo (Vareschi, 1980).
Nano-Micro.: Nanophyll + Microphyll; Meso.: Mesophyll; Macro.:
Macrophyll; Gigan.: Gigantophyll
Table 2. Comparison of leaf type and leaf margin spectra for phanerophytes
from the tropical rain forest in Caiyanghe, the rain forest in southern Yunnan,
the rain forest in northeastern India and the equatorial tropical rain forest in
New Guinea.
Forest type Leaf type Leaf margin
Simple | Compound Entire Non-entire
Tropical seasonal rain forest in Caiyanghe 78.9 21.1 76.9 23.1
Tropical seasonal rain forest in Xishuangbanna, 78.6 21.4 80 20
southern Yunnan?
Lowland evergreen rain forest in northeastern 80.5 19.5 | 722 27.1
India'
Tropical rain forest in New Guinea’ + i a 85 15
'Proctor et al. (1998); 7Paijmans (1970); *Zhu (1997).
Tropical Seasonal Rainforest From Southern Yunan 63
Table 3. Comparison of species diversity among different tropical rain forest
in southern Yunnan.
Forest type Plot Area | Alt f-piope .) NS. | NL: HT
code far) | tm) 2 ©)
Tropical seasonal rain forest in 2500 | 1200 2.528
Caiyanghe
Ravine seasonal rain forest! in 940102 5-10
Menglun, Xishuangbanna 940103 30
940101 25 48
Lower hill seasonal rain forest! in 931206 10 52
Menglun, Xishuangbanna 9201 30 46
Seasonal rain forest dominated 2500 20 =
by dipterocarp” in Mengla, 2500 30 56
Xishuangbanna
H’: Shannon-Wiener’s diversity indexes; Base:2.718283; N.S.: Number of species; N.I.: Number of individuals.
'data from Zhu et al. (1998b) *data from Zhu (2000)
Tree population patterns and diversity
A comparison of the rank/abundance diagram of species based on 0.25 ha plot
each between the tropical rain forest in Catyanghe and other tropical rain forest
types in southern Yunnan was made to show tree population patterns (Fig. 5).
The graph for the rain forest in Caiyanghe has a shorter tail in the species
sequence curve, which indicates fewer species with only one or two individuals
than for the other rain forests.
A total of 29 tree species > 5 cm d.b.h. was obtained from a plot of 0.25
ha in tropical seasonal rain forest at Caiyanghe. Comparable sites in seasonal
rain forests in valleys and on lower hills, and the seasonal rain forest dominated
by dipterocarps at lower altitudes in Xishuangbanna yielded 51, 49 and 56 tree
species respectively (Table 3). The Shannon-Wiener’s diversity index is lower
for the forest in Caiyanghe. This indicates that the tropical rain forest in
Caiyanghe at both of the altitudinal and latitudinal limits has lower species
diversity than the other rain forests at the latitudinal limit in southern Yunnan.
64 Gardens’ Bulletin Singapore 56 (2004)
Rain forest in Caiyanghe LHSR
o ~
100 =
v ©
: :
nm
i) 10 e
e “
o 1 E
oe) o
¢ a
0
0 20 40 60 80
Species sequence Species sequence
Diptorecarp rain forest RSR
= o
= = 100
4 iS)
n a
m
no
E te
ae
€ F 4
2 h eee
n n '
CARES DER EA,
0
0 20 40 60 80
Species sequence Species sequence
Figure 5. Comparison of rank/abundance diagram of tree species between the
tropical seasonal rain forest in Caiyanghe and other tropical rain forest types in
southern Yunnan
LHSR*: Lower hill seasonal rain forest; RSR*: Ravine seasonal rain forest;
Dipterocarp rain forest*: Seasonal rain forest dominated by dipterocarps.
* Zhu (2000).
Discussion
The tropical rain forest in southern Yunnan is considered to be a type of Indo-
Malayan rain forest at the northern margin of tropical Asia as judged by its
similarity to the Indo-Malayan forests in forest profile, physiognomy and
floristic composition (Zhu 1997). Occurring at the area with a seasonal dry
period, the tropical rain forest in southern Yunnan is a semi-evergreen rain
forest with deciduous trees making up 5 — 30% of the number of canopy species
or individuals (Zhu et al. 1998a). Interestingly, tropical seasonal rain forests
in Yunnan have a profile similar to those of lowland rain forests although it
occurs in mountain habitats with higher altitude (Zhu, 1997). In terms of
physiognomy, the tropical rain forest in Yunnan is similar to the lowland semi-
65
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Tropical Seasonal Rainforest From Southern Yunan
66 Gardens’ Bulletin Singapore 56 (2004)
evergreen rain forests in SE Asia as defined by Whitmore (1984, 1990). The
rain forest in Caiyanghe, although occurring further north and at higher
elevations with annual mean temperatures and annual temperature accumulation
lower than those Xishuangbanna of southern Yunnan (Table 4) is up to 35 — 40
m tall with three tree layers. It has the main physiognomic characteristics of
tropical rain forest, such as the upper layer trees with huge buttresses, the
lower layer trees with cauliflory, leaves with drip tips, and the relative
abundance of woody lianas and some epiphytes. Like the seasonal rain forest
in Xishuangbanna, the rain forest in Caiyanghe is also semi-evergreen due to
some deciduous trees in the canopy layer. The rain forest in Caiyanghe is the
extension of seasonal rain forest in Xishuangbanna. It is not only one of the
northern-most types of the Indo-Malayan rain forests, but it is also the northern-
most and the highest altitudinal type of tropical seasonal rain forest in Yunnan.
Beyond the northern watershed line of the mountain of Caiyanghe, tropical
rain forest is not seen, although there are still some tropical floristic elements.
It is possible that the tropical rain forest in southern Yunnan never extended
beyond the mountains of Caiyanghe in the north.
The tropical seasonal forest at Catyanghe has also lower species diversity
than the seasonal rain forests at lower altitude in Xishuangbanna of southern
Yunnan. However, the differences in physiognomy and species diversity
observed from the seasonal rain forest in Caiyanghe do not surpass the
differences observed in other locations in the climatic and geographic zone
categorized as tropical.
In floristic composition, the seasonal rain forest in Caiyanghe is
dominated by tropical Asian elements that characterize the Indo-Malayan flora,
as do the other tropical rain forests in southern Yunnan. However, the dominant
tree species, such as Pometia tomentosa, Alphonsea monogyna, Duabanga
grandiflora (Sonneratiaceae), Garuga floribunda var. gamblei and Terminalia
myriocarpa (Combretaceae), are all near the northern limits of their tropical
Asian distributions. Some strictly tropical Asian species, such as Barringtonia
macrostachya (Lecythidaceae), Myristica yunnanensis and Knema furfuracea
(Myristicaceae), and Pouteria grandiflora (Sapotaceae), which are the climax
species in the tropical seasonal rain forest in the lowlands of southern Yunnan,
are not seen in the rain forest in Caiyanghe. This shows that the tropical seasonal
rain forest in Caiyanghe is really at the altitudinal and latitudinal limits in
southern Yunnan.
Compared with equatorial lowland rain forest in southeast Asia, the
seasonal rain forest in Caiyanghe has more microphyllous plants and more
plants with non-entire leaves, which may be a result of the fact that it occurs at
both latitudinal and altitudinal limits and shows some similarities with
subtropical evergreen broad-leaved forest in eastern Asia and montane rain
forests in southeastern Asia.
Tropical Seasonal Rainforest From Southern Yunan 67
Although the tropical rain forest in Caiyanghe is composed mainly of
tropical floristic elements, the ones of strictly tropical distribution are still
under-represented compared with the Malesian flora. Many Malesian elements
reach to their northern limits here. This indicates that the tropical rain forest is
floristically transitional to the subtropical forests of SW China.
According to the climatic classification for tropical rain forest regions
(Richards 1996), the Caiyanghe Nature Reserve falls outside it because it has
a mean annual temperature of just 17.7°C and a coldest monthly mean of 11.4°C.
The tropical seasonal rain forest in Caiyanghe therefore occurs at the climatic
limits of tropical rain forest. Its occurrence is unusual more for its high altitude
than its northerly latitude.
The tropical seasonal rain forest in Catyanghe occurs quite locally in
limited habitats in valleys. Its occurrence therefore seems to correspond more
with topography than with regional climate. The tropical seasonal rain forest,
which occurs locally in wet valleys and on lower slopes, forms a mosaic with
montane evergreen broad-leaved forests and monsoon forests in southern
Yunnan. This implies that the distribution of tropical rain forest in southern
Yunnan is more influenced by local habitats and microclimates than regional
climate.
Acknowledgements
This project was funded by The National Natural Science Foundation of China
(40271048) and the Yunnan Natural Science Foundation (2002C0067M). Mr.
Li Bao-gui is thanked for help with fieldwork and Mr. Shi Ji-pu and Ms. Cai
Lin for help with the plot data analysis. I also thank Dr. Ruth Kiew and the
anonymous reviewers for their constructive comments on this article.
References
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Manila, the Philippines.
Cao M. 1996. Tree species composition of a seasonal rain forest in Xishuangbanna, Southwest
China. Tropical Ecology. 37: 18-192.
Cao M. and Zhang J.H. 1997. Tree species diversity of tropical forest vegetation in
Xishuangbanna, SW China. Biodiversity and Conservation. 6: 995—1006.
Cao S.S. (ed.) 2003. Caiyanghe Nature Reserve. Yunnan Sci. & Tech. Press, Kunming, China.
p. 17.
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Curtis, J.T. and R.P. MacIntosh. 1951. An upland forest continuum in the prairie-forest border
region of Wisconsin. Ecology. 32: 467-496.
Fedorov, A. A. 1958. The tropical rain forest of China (in Russia with English summary).
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Jin Z.Z. 1997. The diversity features of the floristic elements in the tropical rain forest of
Xishuangbanna, Yunnan. Acta Botanica Yunnanica. Suppl. 9: 129-148.
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Magurran, A.E. 1988. Ecological Diversity and its Measurement. Croon Helm Limited, London,
UK.
Morley, J.R. 2000. Origin and Evolution of the Tropical Rain Forests. John Wiley. England:,
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Paijmans, K. 1970. An analysis of four tropical rain forest sites in New Guinea. Journal of
Ecology. 58: 77-101.
Proctor, J., K. Haridasan, and G.W Smith. 1998. How far north does lowland evergreen tropical
rain forest go? Global Ecology and Biogeography Letters. 7: 141-146.
Qu, Z.X. 1960. Nature reserves in Yunnan. (in Chinese). Journal Yunnan University (Natural
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Raunkiaer, C. 1934. The Life Forms of Plants and Statistical Plant Geography. Oxford
University Press, Oxford, UK.
Richards, P.W. 1996. The Tropical Rain Forest: an Ecological Study. Second edition, Cambridge
Univ. Press, Cambridge, UK. pp.1—19, 159-205.
Shannon, C.E. and W. Wiener. 1949. The Mathematical Theory of Communication. Univ.
Illinois Press, Urbana, USA.
Vareschi, V. 1980. Vegetationsoekologie der Tropen. Stuttgart, Eugen. Ulmer, Germany. pp.
141-148.
Wang C.W. 1939. A Preliminary Study of the Vegetation of Yunnan. Bull. Fan. Mem. Inst.
Bot. IX
Wang C.W. 1961. The forests of China with a survey of grassland and desert vegetation.
Maria Moors Cabot Foundation Publication No. 5. Harvard Univ. Cambridge
Massachusetts., pp. 155-164
Whitmore, T.C. 1984. Tropical Rain Forests of Far East. (Second edition). Clarendon Press,
Oxford, UK.
Tropical Seasonal Rainforest From Southern Yunan 69
Whitmore, T.C. 1990. An Introduction to Tropical Rain Forests. Clarendon Press, Oxford,
U.K. pp. 9-36.
Wu C.Y. 1987. Vegetation of Yunnan. Science Press, Beijing, pp. 97-143.
Xu Y.C., Jiang H.Q. and Quan F. 1987. Reports on the Nature Reserve of Xishuangbanna.
Yunnan Sci. & Techn. Press, Kunming. (in Chinese)
Zhu H. 1992. The tropical rainforest vegetation in Xishuangbanna. Chinese Geographical
Science. 2 (1): 64-73
Zhu H. 1997. Ecological and biogeographical studies on the tropical rain forest of south Yunnan,
SW China with a special reference to its relation with rain forests of tropical Asia. Journal
of Biogeography. 24: 647-662.
Zhu H., Wang H. and Li B.G. 1998a. Research on the tropical seasonal rainforest of
Xishuangbanna, South Yunnan. Guihaia. 18 (4): 371-384 (in Chinese with English abstract).
Zhu H., Wang H. and Li B.G. 1998b. The structure, species composition and diversity of the
limestone vegetation in Xishuangbanna, SW China. Gardens’ Bulletin Singapore. 50: S—
35;
Zhu H., LiB.G., Deng S.C., Cao Y.G. and Zhang X.B. 2000. Tropical rain forest of Caryanghe
Nature Reserve,-Simao, Yunnan and its biogeographical significance. Journal of Forestry
University NE China. 28 (5): 87-93 (in Chinese with English abstract).
Zhu H. 2000. Ecology and Biogeography of the Tropical Dipterocarp Rain Forest in
Xishuangbanna. Yunnan Science & Technology Press, Kunming.
Zhu H. and Zhou H.X. 2002. A comparative study on the tropical rain forests in Xishuangbanna
and Hainan. Acta Botanica Yunnanica. 24: 1-13.
70 Gardens’ Bulletin Singapore 56 (2004)
Appendix. Plot table of the tropical rain forest in Caiyanghe Nature Reserve,
Simao, Yunnan
Plot information:
Plot size: 25 x 100 m Altitude:1200 m Slope aspect:SW
Height of forest: 40 m Coverage: >95% Slope: 25°
Trees
Name of trees Stem% Base area% Frequency% IVI!
Pometia tomentosa 12.34 28.10 8.47 48.91
Alphonsea monogyna pay po a poke yd 8.47 41.27
Duabanga grandiflora 5.19 22.47 6 Be 31.05
Cleidion spiciflorum 18.83 2:09 8.47 29.36
*Garuga floribunda var. gamblei 3.25 13.52 6.78 Pa Fe
Terminalia myriocarpa 0.65 14.82 1.69 17.16
Homalium laoticum 3.90 6.34 6.78 17.02
Garcinia cowa 3.29 1.09 5.08 9.42
Ostodes paniculata Was) 0.43 5.08 8.76
Phoebe puwenensis pe a 0.38 5.08 8.72
Cinnamomum bejolgota 2.60 1.05 o:39 7.04
Baccaurea ramiflora 1:95 0.24 3.39 5.58
Syzygium szemaoense 1.30 0.48 3.39) 5.16
Mitrephora maingayi 1.30 0.38 339 Sad,
Randia wallichii 1.30 0.08 5.39 4.76
Flacourtia rukam 1295 0.97 1.69 4.61
Elaeocarpus sikkimensis 0.65 0.82 1.69 3.16
Amoora tetrapetala 0.65 0.53 1.69 2:87
Horsfieldia kingii 0.65 0.21 1.69 2
*Radermachera igneum 0.65 ey ae 1.69 2.51
Caryota monostachys 0.65 (OA 1.69 2.46
Phoebe lanceolata 0.65 0.12 $69 2.46
Drypetes indica 0.65 0.03 1.69 238
Saurauia macrotricha 0.65 0.03 1.69 pre |
Dichapetalum gelonioides 0.65 0.02 1.69 2.36
Syzygium cathayensis 0.65 0.02 1.69 2.30
Magnolia henryi 0.65 0.01 1.69 2.36
Trigonostemon thyrsoideum 0.65 0.01 169. 2.36
Glycosmis pentaphylla 0.65 0.01 1.69 2.36
Total: 29 species 100 100 100 300
‘Importance Value Indices; *Deciduous trees
Tropical Seasonal Rainforest From Southern Yunan
Sapling and shrub (frem 5 subplots of 5 x 5 m )
Species
Mycetia glandulosa
Cleidion spiciflorum
Alphonsea monogyna
Trigonostemon thyrsoideum
Garuga floribunda var. gamblei
Pandanus furcatus
Magnolia henryi
Desmos dumosus
Baccaurea ramiflora
Leea compactiflora
Mitrephora maingayi
Litsea monopetala
Mallotus philippinensis
Semecarpus reticulatus
Ficus tinctoria ssp.gibbosa
Caryota urens
Flacourtia rukam
Clausena dunniana
Horsfieldia amygdalia
Ardisia arborescens
Sarcosperma kachinensis
Pometia tomentosa
Nephelium lappaceum var. pallens
Listea pierrei var. szemaois
Phoebe puwenensis
Glycosmis pentaphylla
Reevesia pubescens
Ficus collosa
Sapium baccatum
Ficus cyrtophylla
Maesa permollis
Litsea dilleniifolia
Bischoffia javanica
Calophyllum polyanthum
Randia wallichii
Total: 35 species
Individual
—
on)
Pm a a CO ONO ONO ON ON DH WH WH fh ~)
Frequency %
60
40
40
40
20
40
40
20
40
40
20
40
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
Life form
shrub
sapling
sapling
sapling
sapling
shrub
sapling
shrub
sapling
shrub
sapling
sapling
sapling
sapling
sapling
sapling
sapling
sapling
sapling
sapling
sapling
sapling
sapling
sapling
sapling
sapling
sapling
sapling
sapling
sapling
shrub
sapling
sapling
sapling
sapling
ed Gardens’ Bulletin Singapore 56 (2004)
Herbaceous plants (from 5 subplots of 5 x 5 m )
Species Individual Frequency %
Costus speciosus 2 20
Alocasia macrorrhiza ] 20
Pseudoranthemum malaccense 2 20
Aglaonema pierreanum 2 | 20
Alpinia bracteata z 20
Piper bavinum 2 20
Mananthus patentiflora 5 20
Phrynium capitatum 5 20
Rhynchotechum obovatum he 40
Chloranthus spicatus 2 20
Elatostema megacephalum 8 20
Steudnera colocasiaefolia 3 20
Musa acuminata 2 20
Elatostema macintyrei 3 20
Ophiopogon griffithii 2 20
Ophiopogon sarmentosus 10 20
Senecio densiflorum 2 20
Amischotolype hookeri 3 20
Globba racemosa 1 20
Boehmeria siamensis 1 20
Total: 20 species
Lianas and epiphytes
Species Dominance
Combretum latifolium rare liana
Milletia dorwardii rare liana
Heterostemma wallichii rare liana
Gymnostemma pentaphylla rare liana
Tetrastigma crucistum frequent liana
Byttneria grandifolia frequent liana
Toxocarpus villosus frequent liana
Piper flaviflorum frequent _ epiphyte
Rhaphidophora hongkongensis abundant epiphyte
Total: 9 species
Gardens’ Bulletin Singapore 56 (2004) 73-77. 73
Begonia sabahensis Kiew & J.H.Tan (Begoniaceae), a New
Yellow-flowered Begonia from Borneo
RUTH KIEW
Singapore Botanic Gardens,
1 Cluny Road, Singapore 259569
AND
TAN JIEW-HOE
37 Beach Road #03-01, Singapore 189678
Abstract
Begonia sabahensis Kiew & J.H.Tan, (Begoniaceae) is described from Sabah, Malaysia. It
belongs to sect. Diploclinium and is most remarkable for its pure yellow flowers.
Introduction
In 2000, Recin Sapau collected a yellow-flowered begonia from the hills behind
Kampung Katabelangan, about 40 km south of Tenom, Sabah. It was planted
in the Tenom Agricultural Park but never flowered so its flower colour could
not be confirmed. As far as we know, this is the first record of a yellow-flowered
begonia in Borneo. Most Bornean begonias have white or pink or, occasionally,
orange to red flowers, the closest they come to yellow is a greenish cream
colour.
To re-find this species and verify its flower colour, one of us (TJH)
returned to the original site with Recin Sapau and there found a few plants.
The fact that it is known only from one small population in just one locality
suggests it is not a common begonia.
Begonia sabahensis Kiew & J.H. Tan, sp. nov.
A Begonia calcarea Ridl. floriis canarinis (nec tangerinis vel rubis) et majoribus
differt. Typus: Recin Sapau & Tan JH. AL 727/2004 11 Feb 2004 Sungai
Telekoson, Tenom, Sabah, Malaysia (holo SAN).
Plant rhizomatous, hairy, hairs straight, unbranched; c. 2-3 mm long and dense
on the young stems, stipules, petioles and veins on the lower lamina surface;
on the lamina c. 1 mm long, very scattered on the upper surface, sparse beneath;
c. 1 mm long and dense on the veins on the upper lamina surface; brown on the
74 Gardens’ Bulletin Singapore 56 (2004)
stem; red on the young petiole and lamina, transparent on older leaves. Stem
creeping and rooting at nodes, up to 30 cm long, c. 5 mm thick when dried,
green, succulent, unbranched, nodes slightly swollen; without a tuber. Stipules
ovate, 10-15 x c. 5 mm, margin entire, apex acuminate, soon falling. Leaves
distant, 3—7 cm apart; petiole pink, 8—17 cm long, succulent and up to 9 mm
thick in life, c. 3 mm thick when dried; lamina oblique, plain dark green,
glossy above, succulent in life, thinly leathery when dried, broadly ovate,
strongly asymmetric, 9—-10.5 x (9—)10—12 cm, broad side 5.7-6.7 cm wide,
base cordate, slightly unequal, basal lobes rounded, 24 cm long, margin entire,
fringed with hairs 1-3 mm long, apex acuminate 5—8 mm long; veins palmate-
pinnate, 2—3 pairs of secondary veins at the base with another 1-2 pairs along
the midrib branching c. halfway to the margin, and 3 veins in each of the
basal lobes, slightly impressed above, beneath prominent and green.
Inflorescences axillary, pink, erect, shorter than the leaves, male and female
flowers on separate inflorescences. Male inflorescences c. 3.3 cm long,
umbellate with 2 or 3 flowers; bracts pink, ovate, c. 4 x 3 mm, margin entire,
apex setose, persistent. Bracteoles white and transparent, ovate, 2-3 mm long.
Female inflorescences c. 1.5 cm long with a single flower; bracts pink, ovate,
2—3 mm long, margin entire, persistent; bracteoles white and transparent, ovate,
c. 1.5 mm long. Male flowers with a white pedicel 15—25 mm long; tepals 4,
isomorphic, canary yellow, glabrous, broadly oval, slightly narrowed to the
base, margin entire, apex rounded, outer two tepals 12—14 x 9-11 mm, inner
two tepals slightly smaller, 11-13 x 8-13 cm; stamens 20-23, in a lax, sessile
cluster, filaments 1.5—2 mm long, anthers deep yellow, narrowly oblong, 1.5-
2 mm long, apex rounded, opening by lateral slits. Female flowers with a pink
pedicel 2—3 mm long; ovary white becoming pale yellow, 12-13 x 10-15
mm, turbinate, wings 3, equal, rounded c. 7 mm wide, locules 3, placentas
bifid; tepals 5, canary yellow, glabrous, isomorphic, oval slightly narrowed to
base, margin entire, apex slightly acute, 12-13 x 8-11 mm; styles 3, free,
styles and stigma deep yellow, 6-8 mm long, stigmas forming a papillose
spiral band. Fruits not known. —
Distribution: Endemic in Sabah, Malaysia, known only from the type locality
near the Sungai Telekoson, Tenom.
Habitat: Growing in deep shade on thick leaf litter on a steep hill slope in
forest in the transition zone between hill forest and lower montane forest at
1,115 m altitude.
Notes: This new species is very distinctive, not only in its pure yellow flowers,
but also in producing male and female flowers on separate, short inflorescences
from the prostrate rhizome, often from the old stem from which the leaves
have already fallen. The male inflorescences are umbellate, while the female
Begonia sabahensis from Borneo
Plate 1. Begonia sabahensis Kiew & J.H.Tan and Begonia calcarea Ridl.
Begonia sabahensis. A. The plant showing the red-haired young petiole and
the pure yellow flowers produced from the old stem. B. The female flowers
have five tepals and the male four isomorphic tepals. Tan J.H.
Begonia calcarea. C. The plant showing the tufted habit, red veins on the
young leaves and bristles on the upper leaf surface. D. The peachy orangey-
red flowers. R. Kiew
76 Gardens’ Bulletin Singapore 56 (2004)
inflorescences apparently produce a single flower. The leaves are broadly
rounded and wider than long. The succulent petioles with dense, long red hairs
are also striking. The male flowers have four isomorphic tepals. In all these
characters it resembles Begonia calcarea Ridl., which grows on a few limestone
hills around Bau, Sarawak (Ridley, 1906; Kiew and Geri, 2003). In addition,
they both have brightly coloured flowers, rather than the usual white or pale
pink ones.
However, Begonia sabahensis is distinct from B. calcarea in a number
of characters (Table 1), the most conspicuous being flower colour. The flowers
of B. calcarea are orangey red, in male flowers becoming deeper red toward
the margin. The male and female flowers of B. sabahensis are pure yellow
without a tinge of red, a unique feature among Bornean begonias.
Both Begonia calcarea and B. sabahensis belong to section Diploclinium
in possessing rhizomes, male flowers with 4 tepals and ovaries with three
locules each with a bifid placenta.
This fine begonia is so named because it is endemic in Sabah, Malaysia.
It is apparently a rare and local begonia and, because the area is currently
being logged, there is concern for its long-term survival.
Table 1. Differences between Begonia sabahensis and B. calcarea
Character Begonia sabahensis_ | Begonia calcarea
Leaves distant on the stem tufted
Lamina width (cm) 9-12 cals
Lamina not bristly above bristly above
Vein colour on the lower
surface of young leaves green red
Flower colour pure yellow orangey red
Male tepals (mm) 11-14 x 8-13 7-11 x4-9
Stamen number 20-23 25-35
Female tepals (mm) 12-13 x 8-11 COT
Style length (mm) 6-8 4
Begonia sabahensis from Borneo VT
Acknowledgements
We thank Recin Sapau for guiding TJH to the site, the Manager and Botanist
of the Sabah Agricultural Park for permission for Aninguh (Andy) Surat to
accompany TJH in the field, Anthony Lamb for detailed colour notes of the
living plant, and the Curator of the Sabah Forest Department Herbarium (SAN)
for permission to borrow the specimen.
References
Kiew, R. and C. Geri. 2003. Begonias from the Bau limestone, Borneo, including a new
species. Gardens’ Bulletin Singapore. 55: 113-123.
Ridley, H.N. 1906. Begonias of Borneo. Journal of the Straits Branch of the Royal Asiatic
Society. 46: 246-261.
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Gardens’ Bulletin Singapore 56 (2004) 79-94. 79
New Species and Comments on Rhododendron (Ericaceae)
from the Island of Palawan, Philippines
G.C.G. ARGENT
Royal Botanic Garden Edinburgh, Edinburgh EH3 5LR, U.K.
Abstract
Three new species of Rhododendron section Vireya are described: R. mendumiae; R reynosoi
and R. wilkiei, together with a new subspecies of R. javanicum ssp. palawanense and new
records of R. edanoi, which is shown to be the same species as R. pneumonanthum of Borneo
although differing at the sub-specific level. Keys are given to all the Rhododendron species
currently known from the island of Palawan.
Introduction
The island of Palawan (Fig. 1) has high biodiversity (Anon., 1997). It has a
distinctive Bornean element in its flora resulting from the fact that it is the
only part of the Philippines lying on the Sunda Shelf (Whitmore, 1975) so that
it has been connected with land bridges to the west in recent periods of low sea
level. It has been, and still is, in many respects poorly investigated and Sleumer
(1966) in his Flora Malesiana account of Rhododendron only reported three
species specifically from the island, two endemics (R. acrophilum Merr. &
Quisumb. and R. edanoi Merr. & Quisumb.) and one more widespread species
(R. javanicum (Blume) Benn. var. schadenbergii (Warb.) Sleumer). Both
endemics were known only from single collections at the time of his publication.
Joint National Museum (Manila) and Royal Botanic Garden Edinburgh
expeditions in 1992, 1997, 1998 and 1999 have added significantly to our
knowledge of the flora of the island. R. madulidii Argent was described as a
new endemic (Argent 1998) and R. acrophilum was rediscovered and the
description amended after being introduced to cultivation and successfully
flowered (Argent & Madulid, 1995). In this account, three new species are
described from Cleopatra Needle and Thumb Peak (Fig. 1) together with new
subspecies of R. javanicum (Blume) Benn. and R. edanoi. Of these subspecies,
R. javanicum shows great similarity to the Mt. Kinabalu populations of this
species and R. edanoi is shown to be insufficiently distinct from R.
pneumonanthum Sleumer of Borneo to maintain this as a separate species.
Rhododendron bagobonum H.F.Copel. was recorded from Mt Mantalingjahan
in the south of the island (Argent RBGE Acc.No. 19922777). Originally
described from the island of Mindanao in the Philippines, this record is an
80
Gardens’ Bulletin Singapore 56 (2004)
extension of its known range in the Philippines and is yet another link to the
Bornean flora where this species is widespread (Sleumer 1966, Argent 2003).
—_—
N
—~
Vegetative Key to Rhododendron from Palawan
Largest leaves more than 5 cm wide,
petioles more than 2 cm long jJavanicum
Largest leaves up to 3 cm wide, petioles up to 1.5 cm long 2
Leaves less than 1 cm wide bagobonum
Leaves more than 1 cm wide 3
Stems minutely puberulent (and with scales),
leaves narrowly but distinctly revolute madulidii
Stems with scales only, leaf margin flat
(or only very indistinctly revolute in R. mendumiae) 4
Leaf apex rounded (except sometimes with a mucronate point) edanoi
Leaf apex obtusely or acutely pointed 5
Leaves mostly, or at least commonly, broadest in the upper half 6
Leaves broadest in the middle, only rarely broader in the upper half 7
Leaves in tight pseudowhorls spread over c. 6 mm of stem,
leaf blades up to 4 cm long acrophilum
Leaves in loose pseudowhorls spread over c. 20 mm of stem,
leaf blades often more than 5 cm long mendumiae
Largest leaf blades more than.5 cm long,
lateral veins more than 6 per side reynosoi
Largest leaf blades up to 4 cm long,
lateral veins 3 or 4 per side wilkiei
Rhododendron from Palawan, Philippines 81
PALAWAN
11°
i |
TAYTAY
Cleopatra's ‘a ‘
10°
Thumb Peak.
PUERTO PRINCESA
QUEZON
9°
«Mt. Manfalingajan
Figure 1. The island of Palawan, Philippines.
82 Gardens’ Bulletin Singapore 56 (2004)
Key to flowering specimens of Rhododendron from Palawan
1 Flowers white or very pale pink 2
Flowers coloured, red orange or yellow and orange 4
2 Flowers hypocrateraeform, the tube up to 6 mm in diameter edanoi
Flowers funnel-shaped, the tube more than 15 mm in diameter 3
3 Flowers up to 3 per umbel, bract tips reflexed,
stems and pedicels without a fine puberulence mendumiae
Flowers more than 3 per umbel, bract tips erect,
stems and pedicels with a fine puberulence madulidii
4 Flowers longer than broad, up to 1 cm wide bagobonum
Flowers broader than long or more than 2 cm wide 5
5 Flowers more than 6 cm long javanicum ssp. palawanense
Flowers less than 4 cm long 6
6 Filaments and inside of the corolla tube glabrous
corolla lobes red wilkiei
Filaments and inside of the corolla tube hairy, corolla lobes orange 7
7 More than 5 flowers in each umbel,
the flowers bright orange with a red star in the throat reynosot
Less than 4 flowers in each umbel,
the flower bicoloured with a yellow tube and orange lobes acrophilum
1. Rhododendron mendumiae Argent sp.nov.
Named in memory of Mary Mendum [1945-2004] who was on the expedition
which collected this species. A tireless enthusiast for SE Asian botany and
respected research worker.
R. madulidii simillima sed floribus majoribus fragrantibus per inflorescentiam
uno duobus tantum, caulibus pedicellisque sine pilis simplicibus, et disco glabro
differt.
Typus: Philippines, Palawan, Cleopatra Needle. 1600 m. Argent, Chavez, Cronk,
Fuentes, Mendum, Middleton & Wilkie 53. Original collection 22 January 1998.
Cultivated collection: 20 Sept. 2003. Ac. No. 19981815, (holo PNH; iso E).
Rhododendron from Palawan, Philippines 83
Figure 2
Shrub to c. 1 m. Stems pale green, terete, c. 3 mm diam., moderately densely
covered in brown stellate scales. Leaves in rather loose pseudowhorls 3—8 cm
apart. Petiole 4—7 x 2-3 mm, faintly grooved above, pale green or pale pink
with brown scales. Leaf blade narrowly oblanceolate, occasionally elliptic,
40-75 x 17-30 mm, at first scaly with silvery or brown scales above, quickly
glabrescent, below brown scaly, the scales rounded to weakly lobed with darker
centres which are often as broad in diameter as the flanges, distributed 1—2
diameters apart in the mature leaves. Leaf base cuneate, often slightly decurrent
into the petiole; the margin entire, flat or slightly and narrowly recurved; the
apex obtuse, with a somewhat obscure, white, non-protruding, gland-like
structure at the point, midrib narrowly impressed above, distinctly prominent
in the proximal half to two-thirds beneath, pink in colour especially towards
the base, lateral veins plane, rather indistinct, 5—8 per side, the basal arising at
an acute angle, the upper ones wide-spreading, all disappearing before the
edge of the leaf.
Flower buds green, up to 40 x 22 mm, conical but contracted near the base,
imbricate with the tips of the bracts reflexed. Bracts minutely hairy and with
a few small, scattered scales outside and scales along the margins, inner ones
often shortly emarginate. Pedicels 18-22 x 2—3 mm, green, laxly scaly but
without simple hairs. Flowers one or two together, terminal, held more or less
horizontally, white with a cream throat, very strongly and sweetly scented,
52-60 x 102-105 mm; calyx a lobed disk, densely scaly below, less so near the
perimeter; corolla tube 26-30 x c.15 x 17-24 mm, glabrous outside but with
retrorse white hairs in the basal half inside, the lobes c. 46 x 35-46 mm,
spreading more or less horizontally, overlapping to about half their length,
with a rather irregular ‘frilled’ margin.’ Stamens clustered on the lower side of
the flower, exserted 20-25 mm from the mouth, the filaments white, slightly
dimorphic, 38-43 x 0.2 mm wide expanded suddenly for the basal 2.5 mm to
c. 1.4mm wide, glabrous but densely white hairy in the basal expanded portion,
anthers brown, c. 7 mm long, shortly but distinctly apiculate at the base. Disc
green with dense short. white, erect hairs. Ovary c. 6 x 5 mm, broadly trapezoid
in outline, densely silvery scaly and with rather sparse short patent hairs mostly
towards the top. Style cream, lying on the lower side of the corolla tube but
curving upwards when the stigma becomes receptive, c. 50 mm, rather densely
patent-hairy and sparsely scaly in the basal third, the scales arising on low
papillae. Stigma white, 4-7 mm in diam., somewhat crown-shaped, exserted
up to 15 mm beyond the anthers. Fruit 25 x 12 mm, pale green with a covering
of dense brown scales, the pericarp only weakly separating from the valves
which curve outwards on opening without twisting, the placentae separating
from the central column. Seeds c. 2.3 mm long, without tails 1.0 mm, the
longest tail 0.8 mm.
84 Gardens’ Bulletin Singapore 56 (2004)
Notes: Rhododendron mendumiae is somewhat reminiscent of R. madulidii
from Mt Mantalingajan near the southern tip of Palawan but that species has
many more flowers in the inflorescence, puberulous stems and pedicels, and
very different flower bud morphology with erect not reflexed tips to the bracts.
Vegetatively, it looks very similar to R. jasminiflorum Hook. var. copelandii
(Merr.) Sleumer, but the massive flowers of this new species are totally different
to the slender flowers with very small lobes of that species and the bud
morphology again is also very different.
This very distinctive species should be associated with Sleumer’s (1966)
subsection Euvireya on the basis of its short broad corolla tube relative to the
enormous lobes and the well spaced, stellate scales on the leaves. It is slightly
anomalous in that, from the single observation of fruit dehiscence, the pericarp
remains closely attached to the valves, only weakly separating near the tips
but it clearly does not belong in the groups with adherent pericarps from the
scale morphology. On the basis of leaf size it could be placed in either series
Buxifolia or series Javanica. In series Buxifolia, it clearly does not key out
having a very distinct combination of characters. In series Javanica, it would
key out with the Indonesian species: R. lompohense J.J.Sm., R. bloemberginii
Sleumer, from Sulawesi or R. buruense J.J.Sm. from Maluku, all rather unlikely
geographical alliances. All three of these species have many more flowers in
the inflorescence, much smaller corolla lobes and ovaries that are much longer
than broad. When this species flowered in cultivation for the first time in 2002
with a single flower, it was thought this must be an anomalous peloric form
but the following year two-flowered inflorescences were produced on several
plants and it was concluded that these large flowers are normal. The seeds are
rather unusual in having such short tails, these are characteristic of vireyas
which occur in open situations on mountain peaks. R. retusum (Blume) Benn.,
R. adinophyllum Merr. and R. abietifolium Sleumer have similar seeds with
very reduced tails. Since this species was collected near the summit of Cleopatra
Needle where the vegetation was dense, mossy forest, this 1s a surprising
character. Plants have been raised from both seed and cuttings collected in the
wild and from the isolation of the peak it seems virtually impossible that these
are hybrids. Given the isolation of the habitat and the very restricted area of
mossy forest on the mountain, the population of this species is very small and
must be vulnerable to any kind of habitat disturbance.
Additional specimens: All from the type locality with the same collecting
details as the type: 19981798; 19981800; and 20031269.
2. Rhododendron reynosoi Argent sp. nov.
Named in memory of Ernesto Reynoso who was a wonderful companion on
several expeditions and made a considerable contribution to Philippine botany
working for the National Museum, Manila.
Rhododendron from Palawan, Philippines 85
Figure 2. Rhododendron mendumiae
a Branch with inflorescence; b bracteole; c, d, f bracts; e enlargement of
indumentum on abaxial surface of bract; g enlargement to show scales on bract
margins; h part flower, opened; i anthers; 7 enlargement to show ovary
indumentum; k pistil.
86 Gardens’ Bulletin Singapore 56 (2004)
Umbella erecta florum clare aurantiacorum, foliis comparate parvis et foliis
foliaceis bracteas sensim transientibus distincta. Typus: Philippines, Palawan,
Cleopatra Needle, 22 January 1998, just below summit, moss forest, 1600m.
Argent, Chavez, Cronk, Fuentes, Mendum, Middleton & Wilkie 44A. Original
collection 22 Jan. 1998. Cultivated collection: 29 August 2002, Ac. No.
19981806 (holo PNH; iso BE).
Figure 3
Shrub to c. 80 cm. Stems green, terete, c. 3 mm diam., moderately densely
covered in brown stellate scales. Leaves in close pseudowhorls 1—2 cm apart,
consisting mostly of 6—7 leaves. Petiole 6-10 x 1.5—2 mm, not grooved above,
pale green with brown stellate scales. Leaf blade elliptic to slightly oblanceolate
50-80 x 15-30 mm, with some sparse silvery, sub-circular scales at first above
but quickly glabrescent here, below moderately brown scaly, the scales rounded
or weakly lobed slightly impressed, the centres small 1—3 diameters apart in
the mature leaves. Leaf base broadly cuneate; the margin entire, flat, narrowly
cartilaginous; the apex acute to broadly acute, sometimes shortly acuminate or
apiculate, midrib narrowly impressed above although slightly prominent just
above the petiole where it is also faintly grooved; beneath broadly raised for
about three-quarters of the length proximally, lateral veins plane, 6—10 per
side, spreading broadly but then curving upwards to link with the vein above
before the margin, distinct above but obscure below.
Flower buds green, up to 40 x 20 mm, ovoid, sharply pointed with a gradual
transition from foliage leaves to bracts these transitional forms having a smaller
but distinct foliar blade with broad translucent decurrent margins instead of a
petiole, the inner bracts narrowly lanceolate to subulate with spreading points.
Bracts glabrous or with a few, scattered scales outside and along the margins
but no simple hairs. Pedicels 9-15 x 1-2 mm, orange, moderately scaly but
without simple hairs. Flowers terminal, 6-10 in an umbel, held erect or semi-
erect, bright orange with a red star in the throat the points extended as lines
along the lobes, without scent, 32-40 x 30-40 mm; calyx a low scaly disc.
Corolla tube c. 15 x 6 x 13 mm, glabrous outside, white hairy in the proximal
half inside, the lobes 15-30 x 12—22 mm, semi-erect, overlapping half to three
quarters of their length, rounded or weakly emarginate at the apex. Stamens
regularly arranged around the mouth of the flower, exserted to c. 5 mm,
distinctly dimorphic, filaments pale orange, with white shaggy hairs in the
basal third glabrous above, the anthers dark purple c. 3 x 1.5 mm. Disc green
almost glabrous but with a few short hairs on the upper margin. Ovary c. 6.5 x
3 mm, ellipsoid, silvery-scaly and densely white hairy. Style orange, at first
lying on the lower side of the corolla tube, rising to a central position as the
flower ages, c.14 x 0.8 mm, glabrous. Stigma purplish-red, hardly expanded
from the style, c. 3 mm in diam. Fruits sub-cylindrical, tapering distally to the
Rhododendron from Palawan, Philippines 87
Figure 3. Rhododendron reynosoi
a branch with inflorescence; b bracteole; c bract with enlargement to show scales
on the margin; d flower cut open; e stamen with anther back view; f pistil with
enlargement of ovary indumentum.
8& Gardens’ Bulletin Singapore 56 (2004)
style, grooved longitudinally in the proximal two thirds, 20-25 x c. 6 mm, the
calyx slightly accrescent. On splitting the outer layer peeling back irregularly,
the valves spreading to c. 45°, the placentae remaining more or less adherent
to the central column. Seeds c. 3.7 mm long, with tails c. 0.8 mm, the longest
tail 1.5 mm.
Notes: This species keys out in Sleumer (1966) to Rhodendron leytense Merr.
differing, however, not only in the colour of the flowers, which is recorded as
yellow in R. leytense but in the non- puberulous pedicels and it has more flowers
in the inflorescence than that species. A unique feature, at least amongst the
Philippine rhododendrons, is the gradual transition from foliage leaves to bracts,
the intermediates are longer than the bracts, have broad sheathing bases but
progressively smaller blades than the foliage leaves.
This species is very distinct with its bright vibrant orange flowers in a
tight, erect umbel. This alone distinguishes it from all other Philippine species
as those with several flowered umbels either have very differently sized flowers
or have a totally different disposition. It is much smaller leaved than R.
javanicum (Blume) Benn. var. schadenbergii (Warb.) Sleumer as conceived
by Sleumer (1966).
3. Rhododendron wilkiei Argent sp. nov.
Named after Peter Wilkie, botanist and explorer in SE. Asia, who nearly died
of a malarial attack helping to collect this species.
R. acrophilo similis a qua flore intense rubro haud bicolori et filamentis ac
corollae parte interiore glabris (haud pilosis) differt. Typus: Philippines,
Palawan, Thumb Peak, Argent, Chavez, Cronk, Mendum, Middleton & Wilkie
48. Original collection 30 January 1998; Cultivated collection Ac.No.19981810
11 July 2003. (holo PNH; iso E).
Figure 4
Weak shrub to c. 30 cm. Stems pale green, terete, c.1.5 mm diam., moderately
densely covered in brown stellate scales. Leaves in pseudowhorls 1—2 cm apart,
consisting of 34 larger and 2—3 smaller leaves, mostly only possessing leaves
at the terminal pseudowhorl. Petiole 2-3 x c.1.5 mm, not distinctly grooved
above but with a central line, pale green with brown stellate scales. Leaf blade
elliptic 25—40 x 10-15 mm, with some sparse silvery, stellate scales at first but
quickly glabrescent above, below moderately brown scaly, the scales rounded,
lobed to sub-stellate, impressed into small pits the centres small and indistinct
distributed 24 diameters apart in the mature leaves. Leaf base broadly cuneate;
the margin entire, flat; the apex broadly acute to obtuse, sometimes with an
Rhododendron from Palawan, Philippines 89
| 2mm
Figure 4. Rhododendron wilkei
a branch with inflorescence and opening bud; b anthers face and back view;
c flower cut open; d inner bract with enlargement to show marginal scales;
e bract. |
obscure, white, non-protruding, gland-like structure at the point, midrib
narrowly impressed above, slightly prominent just above the petiole but then
somewhat impressed distally beneath, translucent green especially towards
the base, lateral veins plane, 2-4 per side, spreading broadly but disappearing
before the margin, distinct above but obscure below.
Flower buds green, up to 18 x 8 mm, narrowly conical but contracted near the
base, sharply pointed with the basal bracts spreading, the upper (inner) bracts
mostly appressed. Bracts glabrous or with a few small, scattered scales outside
and with scales along the margins. Pedicels 10-15 x 1-2 mm, red, moderately
90 Gardens’ Bulletin Singapore 56 (2004)
scaly but without simple hairs. Flowers terminal, solitary or paired, held half-
hanging, red, without scent, c. 27 x 44 mm; calyx a low scaly disc or with two
irregular longer lobes up to 2 mm long. Corolla tube c. 15 x 4x 10 mm, glabrous
outside and inside, the lobes c. 19 x 21 mm, spreading horizontally, overlapping
to about half to two thirds of their length, emarginate at the apex. Stamens
regularly arranged around the mouth of the flower, filaments pale pink, glabrous,
the anthers dark purple 2—2.5 x c.1.25 mm. Disc green glabrous. Ovary c. 6 x
2.5 mm, ellipsoid, rather sparsely brown-scaly, without simple hairs. Style pink,
at first lying on the lower side of the corolla tube, rising to a central position as
the flower ages, c. 20 mm, glabrous. Stigma purplish-red, hardly expanded
from the style, c. 1.25 mm diam. Fruits not seen.
Notes: This keys out in Sleumer (1966) to Rhododendron porphyranthes
Sleumer, known from only a single collection from the Arfak Mountains in
New Guinea. Apart from being a most unlikely distribution pattern, R.
porphyranthes differs in its rounded to emarginate leaves, in having much
smaller corolla lobes, a smaller ovary and a style only half as long as in this
new species. It is probably most similar to R. acrophilum, which is at least
from the same island, but differs in the flower colour, which is bicoloured with
a yellow tube and orange lobes in that species (Argent & Madulid, 1995) and
that the filaments and the inside of the corolla are glabrous in R. wilkiei, not
hairy. This new species also lacks the white waxy substance associated with
the flower buds of R. acrophilum and is much less vigorous in its growth habit.
Additional specimen with the same collecting details as the type: 55,
Ac.No.19981817 (PNH, E).
Rhododendron javanicum (Blume) Benn. ssp. palawanense Argent ssp. nov.
Named from the island of Palawan from where it was collected.
Folia spiraliter disposita laevia. Gemmae anguste ovoideae apice acuto
bracteisque adpressis. Flores c. 10 cm longi, aurantiaci fauce luteo, lobis circiter
duplo longioribus quam latioribus. Typus: Philippines, Palawan, Mt.
Mantalingajan. Argent & Romero Ac. 19922770 Cultivated specimen collected
16 January 1998. (holo PNH, iso E).
Shrub to 1.5 m. Leaves spirally arranged, smooth. Flower buds green, narrowly
ovoid with an acute apex and appressed bracts. Pedicels glabrous or sparsely
scaly. Flowers c. 10 x 9 cm, bicoloured, with a yellow tube, hairy inside near
the base and forming a yellow ‘star’ in the mouth, lobes orange, c. 5 x 2.5 cm.
Stamens with hairs in the basal quarter of the filaments, anthers c. 5 mm with
grey pollen. Ovary glabrous, (in the type), or variably hairy and scaly.
Rhododendron from Palawan, Philippines 91
Notes: Following Argent et al. (1988) and Argent (2003), Rhododendron
javanicum is presently treated in a broad sense including R. brookeanum H.Low
ex Lindl. and R. moultonii Rid|. This complex of mainly low altitude forest
epiphytes is characterised by large, more or less elliptic, acutely pointed leaves
and large funnel-shaped flowers in a range of colours. As conceived here, it is
widespread occurring through Sumatra, Peninsular Malaysia, the Philippines,
Borneo, Java, Bali to Sulawesi. It is highly variable in its leaf arrangement and
ovary indumentum even within some populations and is difficult to deal with
satisfactorily although various populations are distinct. This new subspecies is
very similar to the Mt. Kinabalu populations: R. javanicum ssp. brookeanum
(H.Low ex Lindl.) Argent & Phillipps var. kinabaluense Argent, A.L.Lamb &
Phillipps, which is similarly a high altitude form with large, conspicuously
bicoloured flowers, broad flower buds with appressed bracts that are often
emarginate and rugose leaves arranged in regular spirals. The Palawan plants
differ, however, in having smooth leaves, more slender flower buds with acutely
pointed bracts and longer flowers, with longer and narrower lobes to the corolla,
and grey pollen. The significance of pollen colour is not yet known but it was
considered important as a specific character in Aeschynanthus (Gesneriaceae)
by Mary Mendum (pers. comm.). Within the Philippines, all plants of
Rhododendron javanicum were included in var. schadenbergii (Warb.) Sleumer
(Sleumer, 1966). However, these recent collections from Mt. Mantalingajan
are very different from the majority of Philippine collections, mostly from the
island of Mindanao. These have much shorter, uniformly red flowers and cream
pollen. The ovary indumetum in ssp. palawanense varies from completely
glabrous to hairy and scaly in the proximal two thirds. Thus this Palawan
population appears to be similar but not identical to the Kinabalu one and very
different to at least that on Mindanao. It seems best to give these plants
subspecies status at present which draws attention to their unique character
and distribution. Further collections of this species complex from other islands
in the Philippines are badly needed to satisfactorily understand the variation.
Additional collections that are very similar are from Cleopatra Needle, Ac.
No. 19981796, and from Thumb Peak, Ac. Nos. 19981802, 19981812. A
vegetative herbarium specimen in PNH from Victoria Peak is probably the
same taxon.
New Records of Rhododendron edanoi Merr. & Quisumb.
Philippines, Palawan, Thumb Peak, near summit c.1200 m Argent, Chavez,
Cronk, Mendum, Middleton & Wilkie 46, Ac. 19981808 Original collection 30
Jan.1998; Philippines, Palawan, Cleopatra Needle, just below summit c.1500
m, in mossy forest Argent et al. Ac.19981819. Original collection 15 January
1998. (All preserved in PNH. and E).
92 Gardens’ Bulletin Singapore 56 (2004)
Notes: This species was previously only known from the type collection made
on Mt. Mantalingajan near the southern tip of the island of Palawan on 13"
May 1947 by G.E. Edanio. The materials collected in 1998 on the tiny isolated
area of mossy forest on the summit of Cleopatra Needle and from Thumb
Peak, a similar isolated area of mossy forest a little further to the south, were
both seed and cuttings of what was designated in the field as the ‘round leafed’
rhododendron. None of these rhododendrons was in flower at the time of
collection and identification had to await flowering in cultivation. This occurred
first for the cutting material in March 2000 and for the seed in 2003. The
interest in these collections is not just that this.extends the range of this otherwise
point endemic to most of the length of the island of Palawan but that it shows
this species to be conspecific with that of Rhododendron pneumonanthum
Sleumer of Borneo. Sleumer (1966) distinguished these two species at couplet
‘31’ on the characters “Corolla tube more or less manifestly and gradually
narrowed from the base upwards. Leaves sub-sessile (R. pneumonanthum) vs.
Corolla tube equally wide all over or slightly and gradually widened from the
base upwards. Leaves distinctly petioled. (R. edanoi).
From examination of isotypes of the original collection of Rhododendron
edanoi from BM, K and L, the first character is not a valid difference. Several
corolla tubes show a quite definite narrowing from the proximal to the distal
ends. The corollas from cultivated plants from both Palawan and Borneo, in
the fresh state all quite clearly taper from base to mouth. After pressing, the
tube can appear to at least look parallel-sided. There does appear to be a
difference in petiole length between Bornean and Philippine materials but again
there is a lot of variation even on one plant and this alone would not support
maintaining these plants as different species. There is a very small difference
in the scales on the leaves from the two islands, those from Bornean materials
are darker and more variable in size but both have essentially the same sub-
stellate shape, are rather tall and set on pronounced epidermal tubercles. The
vegetative habit of the plants in cultivation from the different islands 1s certainly
very different. Both Philippine collections are slow and low growing compared
to the tall ‘leggy’ and much more vigorous Bornean plants. It is tempting to
imply that the Philippine plants growing in such small isolated populations are
suffering from inbreeding depression compared with the much larger
populations of R. pneumonanthum in northern Sarawak where it is reported to
be common over large areas close to Bario on the slopes of Mt. Murud (A.
Lamb pers. comm.). Differences in habit between different populations of the
same species are known elsewhere in section Vireya. R. burttii P.Woods varies
on different mountain ranges (Argent et al., 1988) and R. leptanthum F.Muell.
(pers. obs.). The clinching factor in regarding the Bornean R. pneumonanthum
to be, at best, a subspecies and not specifically distinct from R. edanoi is the
fact that the flower bud morphology is identical and very distinctive from both
Rhododendron from Palawan, Philippines 93
island populations. The flower buds are broadly oval with reflexed points to
the tips of the bracts (perulae of Sleumer, 1966).
Thus the Bornean populations are reduced to the status of subspecies
which reflects the relationship between Philippine and Bornean plants more
accurately:
R. edanoi ssp. pneumonanthum (Sleumer) Argent comb. nov.
Basionym: R. pneumonanthum Sleumer, Reinwardtia 5 (1960) 132.
Key to the subspecies
Small spreading plants, mostly less than 30 cm,
petioles mostly more than 5 mm long,
distinctly longer than wide, scales pale brown ssp. edanoi
Tall erect plants, often more than | m, petioles up to
3 mm long, hardly longer than wide, scales
blackish-brown when mature ssp. pneumonanthum
Acknowledgements
The author is grateful to the Royal Botanic Garden Edinburgh for sponsoring
the expeditions that collected the materials on which this paper is based and to
SEERAD for funding the research. The National Museum, Manila, Philippines
supported and partnered the expeditions and I am especially indebted to Dr.
Domingo Madulid, Chief of the Division of Botany who provided support staff
and helped us in many ways and to the officials of the Palawan Tropical Forestry
Protection Program and the Palawan Council for Sustainable Development.
Curators of herbaria at BM, K, L and PNH are thanked for providing access to
relevant herbarium collections. Dr Robert Mill translated the Latin diagnoses
and made many helpful comments and Eve Bennett provided the elegant
illustrations.
References
Anon. 1997. Philippine Biodiversity: an Assessment and Plan of Action. Philippines,
Department of Environment and Natural Resources. Makati City: Bookmark, 1-298.
Argent, G. 1998. Rhododendron madulidii (Ericaceae): a new Philippine species from Palawan.
The New Plantsman 5: 204-211.
94 Gardens’ Bulletin Singapore 56 (2004)
Argent, G. 2003. Species patterns in Rhododendron section Vireya from sea level to the snow
line in New Guinea, 160-170. In: G. Argent and M. McFarlane. (eds.) Rhododendrons in
Horticulture and Science. Royal Botanic Garden Edinburgh, U.K.
Argent, G., A. Lamb, A. Phillipps and S. Collenette. 1988. Rhododendrons of Sabah. Sabah
Parks Publication No. 8. 1-146.
Argent, G. & Madulid, D. 1995. Rediscovery of Rhododendron acrophilum. Rhododendrons
with Camelias and Magnolias. 47: 39-40 & fig.14.
Sleumer, H. 1966-67. Ericaceae. Flora Malesiana. 6: 469-914.
Whitmore T.C. 1975. Tropical Rain Forests of the Far East. Clarendon Press, Oxford, U.K.
1-282.
Gardens’ Bulletin Singapore 56 (2004) 95-100. 95
Begonia sizemoreae Kiew (Begoniaceae), a Handsome New
Begonia from Vietnam
RUTH KIEW
Singapore Botanic Gardens, Singapore 259569
Abstract
Begonia sizemoreae Kiew (sect. Platycentrum) is described from the Ba Vi National Park in
North Vietnam. Closely related to B. rex Putzeys and with similarly fine variegated leaves, it
is striking for the crimson reticulation of veins in the central and marginal parts of the leaves.
Introduction
This attractive begonia was first discovered by Mary Sizemore on 6" November
1996 in Ba Vi National Park. c. 80 km west of Hanoi, Vietnam. It was listed in
the American Begonia Society’s Unidentified Species List as Begonia U388
and, because of its beautiful leaves, was quite widely circulated among the
Society’s members (Keepin, 2003). Naming such a striking plant, which is a
significant addition to the foliage begonias already in cultivation, is long
overdue. It is here described and illustrated and is named in honour of Mary
Sizemore.
Begonia sizemoreae Kiew, sp. nov.
A Begonia rege Putzeys petiolis quam laminis subduplo longioribus, venis
tertiariis foliorum rubris et cymis 3-floris statim dignoscenda. Typus: Vietnam,
Ba Vi National Park, Ha Tay Province, c. 80 km west of Hanoi, Accession No.
20020399 cult. in Singapore Botanic Gardens ex Palm Hammock Orchid Estate,
Miami, U.S.A. R. Kiew 5304 (holo SING, iso HN).
Plate 1
Begonia rhizomatous. Stem creeping, not branched, succulent, c. 2.5 cm long
and c. 5 mm thick; reddish brown or purplish; without a tuber. Indumentum of
white, c. 10 mm long, uniseriate hairs, scattered on the upper lamina surface
(not on the veins), these c. 5 mm long and dense on the stem, stipules and
petiole, 2-4 mm long and dense on the lamina margin and on the undersurface
96 Gardens’ Bulletin Singapore 56 (2004)
of the midrib, secondary and tertiary veins. Stipules persistent, broadly
triangular tapering to a setose point, 8-12 x 4-9 mm, reddish when young,
becoming pale yellowish green with a reddish band along the midrib, margin
entire. Leaves alternate, tufted; petiole terete, (4.5—)10.5—19 cm long, pale red
or purplish; lamina oblique, asymmetric, broadly ovate, 6.5—11.5 x 5.5—10.5
cm, broad side 4—6.5 cm wide, base unequally cordate, larger basal lobe 2.25—
3.5 cm long, margin entire, slightly undulate, apex slightly acute, sometimes
rounded, lamina slightly velvety above and thinly succulent in life, papery
when dried, variegated on the upper surface with the basal part (less than half
the lamina width) and a band c. 10-15 mm wide around the margin jade green
sometimes with a blackish hue and with the middle part pale silvery grey-
green, secondary veins green in the basal part, white in the middle part and
deep crimson in the marginal band, tertiary veins deep crimson in the basal
and marginal parts and white in the middle part, on the lower lamina surface
green with secondary veins greenish brown, tertiary veins deep crimson;
venation palmate-pinnate with 2 pairs of secondary veins at the base and 2
pairs along the midrib with another vein in the basal lobe, secondary veins
branching c. halfway to the margin, impressed above, prominent beneath.
Inflorescences axillary, monochasial cymes, protandrous with two male flowers
and one female flower, glabrous, 7.25—19.5 cm tall with two branches 2-3
mm long, rosy red, greenish distally. Bracts narrowly ovate, 11-15 x 4-5 mm,
glabrous, pale green, almost transparent, reddish along the midrib and towards
the tip, apex acutely pointed, margin entire, caducous. Male flower: pedicel
18-20 mm long, pale pink or white; tepals 4, glabrous, margin entire, apex
rounded, outer two tepals ovate, 18—25 x 13-14 mm, deep pink paler towards
base, longitudinal veins slightly impressed; inner two tepals oval, 16—25 x 9—
11 mm, pale pink, veins not impressed; stamens many, joined into a more or
less globose cluster, 5—7 x 5 mm; filaments joined into a column for 1—3.5 mm
and free for c. 1.25 mm, white; anthers narrowly oblong, c. 2.5 mm long, deep
yellow, apex strongly apiculate, c. 1 mm long, thecae dehiscing through lateral
slits. Female flower: pedicel 15—25 mm long, slightly reddish; ovary pale green,
7-8 x 20-23 mm, wings 3, unequal, long wing 12-17 mm wide, two shorter
wings 3—5 mm wide, locules 2, placentas bifid; tepals 5, obovate tapered to an
acute apex, more or less isomorphic, 19-21 x 10-12 mm, deep pink at the tip,
paler at the base, margin entire and undulate; styles and stigma golden yellow
or yellowish green, styles 2, 8-9 mm long, joined for c. 2 mm at the base,
bifurcating in the upper third; stigmas a spiral papillose band. Fruit and seed
not known.
Distribution: Endemic in N. Vietnam (Ba Vi National Park).
Habitat: Locally common on earth banks beside the road in medium shade at
c. 800 m altitude.
Begonia sizemoreae from Vietnam 97
Plate 1. Begonia sizemoreae Kiew.
A. The plant. B. Monochasial cyme with two male flowers and the developing
female flower. C, D. Male flower. E, F. Female flower. Serena Lee
9§ Gardens’ Bulletin Singapore 56 (2004)
Other specimen examined: U.S.A., Palm Hammock Orchid Estate, Miami,
Kiew s.n. 10 Feb 2002 (SING).
Notes: Begonia sizemoreae belongs to sect. Platycentrum in being rhizomatous,
protandrous, and having a male flower with four tepals and a female flower
with an ovary with one wing much longer than the other two, two styles, and
two locules each with a bifid placenta.
This new species is very similar to Begonia rex Putzeys from Assam,
India, in its leaf shape, the silvery band around the middle of the leaf and in
flower structure. However, B. sizemoreae is distinct in its petioles being much
longer (over 1.5 times longer) than the lamina and densely hairy, whereas in B.
rex the petioles are shorter than the lamina and are less hairy. The lamina of B.
rex 1s distinctly bullate, while that of B. sizemoreae is not. The details of the
variegation are significantly different: in B. sizemoreae the basal and marginal
parts of the lamina are jade green with striking deep crimson venation, while
in B. rex the basal and marginal parts are dark green to bronzy green and the
venation is concolorous. In addition, the leaf base is never overlapping, as is
seen in wild collections of B. rex (e.g. Griffith 2588, K). In B. sizemoreae, the
inflorescence is a three-flowered, monochasial cyme with a single female
flower, while in B. rex it is a dichasial cyme with four female flowers.
The introduction of Begonia rex is well documented-it was inadvertently
introduced from Assam together with a consignment of orchids and was bought
by J. Linden for 10,000 francs (Thompson & Thompson, 1981). The original
French publication described it as “ce merveille Begonia’ (Hooker, 1859). Since
then it has been hybridized with a wide range of species, both closely and
distantly related, to produce an amazing variety of leaf shapes (some even
spiral!), colours, patterns and leaf textures but most with the characteristic
silvery sheen. It is doubtful now whether any ‘pure’B. rex plants exist in
cultivation considering the ease with which it hybridizes. As Krempin (1993)
records, the hybrids of B. rex are ‘never ending’ and there are probably more
than a thousand named and unnamed. Begonia rex is not recorded wild from
Vietnam. H6 (1991, 1999) included it in his account of Vietnamese begonias
as a cultivated species.
Golding (2004) recently identified Begonia U388, which is here described
as Begonia sizemoreae, as B. longiciliata C.Y. Wu, a begonia from Guizhou,
China. However, B. sizemoreae is different from this species in its variegated
leaves, the larger basal lobe (one third as long as the midrib) and the entire
margin; the inflorescence is a monochasial cyme with two much larger male
flowers. In contrast, the lamina of B. longiciliata is plain deep green above
and green beneath, the larger basal lobe is only one fifth to one sixth as long
as the midrib, and the margins are denticulate; the outer tepals of the male
flowers are 13 by 8 mm and the inner ones 9 by 3.5 mm. Although Wu in Wu
Begonia sizemoreae from Vietnam 99
& Ku (1995) described the inflorescence as monochasial with two flowers,
the illustration shows it to be dichasial with potentially six male flower and
four fruits. In this latter character, it resembles B. rex.
Apart from its stunning foliage, Begonia sizemoreae has the advantage
that it is easy to grow in cultivation unlike many other wild species in Asia. In
cultivation, its leaves can grow up to 26 by 16.5 cm (M. Sizemore, pers. comm.).
It flowers freely and, in Singapore, it is not seasonal. It propagates easily from
the leaves. Keepin (2003) reports that it is self compatible and produces viable
seed. This new species is therefore an important addition to the stable of
cultivated begonias. Its charm lies in the deep crimson tracery of the fine
network of veins—a feature not seen in B. rex. No doubt, it too will prove easy
to hybridise, but it is important to keep a pure line so that its identity is not
lost.
Acknowledgements
I am grateful to Mary Sizemore who shared her discovery with other begonia
enthusiasts and who provided habitat data; to the Institute of Ecology and
Biological Research (NCST), Hanoi, Vietnam for giving M. Sizemore
permission to collect plants and to Nguyen Van Du, who accompanied her; to
Tim Anderson, Palm Hammock Orchid Estate, who drew my attention to this
species and provided me with material; to Norhayati Mohd Din, for her care of
the Singapore Botanic Gardens begonia collection; to Serena Lee for the
photographs; to M.J.E. Coode for correcting the botanical Latin; and to the
Keeper of the Herbarium, Royal Botanic Gardens Kew, for permission to
examine specimens in his care.
References
Golding, J. 2004. Begonia U 388 is Begonia longiciliata C.Y.Wu. The Begonian. 71: 154—
156.
Ho, P. H. 1991. Figure 2053 Begonia rex. Cay Co Viét Nam. 1: 739.
Ho, P. H. 1999. Figure 2339. Begonia rex. Cay Cé Viét Nam. 1: 585.
Hooker, W. J. 1859. Begonia rex, Royal Begonia. Curtis’s Botanical Magazine. 85: Plate
5101.
Keepin, T. 2003. Begonia U388 out of a terrarium in Houston. The Begonian. pg. 128.
Krempin, J. 1993. Know Your Begonias. Krempin Book, Australia.
100 Gardens’ Bulletin Singapore 56 (2004)
Thompson, M.L. and E.J. Thompson. 1981. Begonia—The Complete Reference Guide. Times
Books, U.S.A.
Wu, C.Y. and T.C. Ku. 1995. New taxa of the Begonia L. (Begoniaceae) from China. Acta
Phytotaxonica Sinica. 33: 251-280.
Note added in press: A begonia called ‘Vietnamese hairy Begonia’ was
illustrated in The Garden (2004) Volume 129 (7): 515, after it had been exhibited
at the Chelsea Flower Show. From the photograph, it looks very like Begonia
sizemoreae, which suggests that this species is beginning to enter the
horticultural trade.
Gardens’ Bulletin Singapore 56 (2004) 101-145. 101
The Vegetation and Plants of Niah National Park, Borneo
KATHARINE GEORGINA PEARCE
32 Lorong Kumpang 4
93200 Kuching, Sarawak, Malaysia
Abstract
Based on survey work and specimen collection carried out during 2001 to 2003, vegetation
types that occur in Niah National Park (the various facies of limestone vegetation—the Subis
limestone is the most prominent feature of the Park—mixed dipterocarp forest, seasonal swamp
forest on clayey marls and on peat soils, riparian forest, and regenerating forest developing
after shifting cultivation) are described. A provisional checklist of species includes more than
300 species. Ferns, Acanthaceae, Annonaceae, Araceae, Begoniaceae, Euphorbiaceae,
Gesneriaceae, Moraceae, Rubiaceae and Urticaceae are well represented, while orchids and
palms appear to be under-collected. A number of fruit tree species and other useful plants
occur in the Park. The Park is home to at least 12 endemics found nowhere else, and 5 possible
new species.
Introduction
The following account of the plants of Niah National Park (Niah NP) grew out
of work done to map and describe the vegetation types at Niah NP under the
Sarawak Forest Department/DANIDA Project Support to Wild Life Master
Plan Implementation through the Improved Management of Totally Protected
Areas, Sarawak, Malaysia (SWMPI) completed in 2003. An understanding of
the Park’s vegetation and flora is, of course, a sine qua non for effective park
management.
Niah NP lies close to the north coast of Sarawak in the lower reaches of
Sungai (Sg) Niah catchment. The Park’s most prominent feature, located almost
entirely within the Park, is the striking Subis limestone massif (the remains of
an ancient coral reef), which rises vertically out of the Sg Niah floodplain. The
Park area has a unique history of human interaction dating from prehistoric
times to the present. Constituted in 1974 (with the inclusion of the Niah Great
Cave and Historical Monument in 1994), Niah NP covers 3,139 ha. Local
people formerly farmed and planted fruit trees in the area and some local
inhabitants have been accorded a range of gazetted rights within the Park,
such as the right to collect fruit and forest products.
Although the first plant collections were made at Subis in 1894, the
Gunung Subis limestone outcrop, until this study, was somewhat neglected by
plant collectors (Pearce, 2003). Particularly for some of the earlier collections
102 Gardens’ Bulletin Singapore 56 (2004)
from non-limestone habitats that give the locality as “Niah’, it cannot be
ascertained whether the collections were made within what is now the Niah
NP area.
Methodology
This account is based on fieldwork at Niah NP during the period 2001 to 2003
and an examination of available literature. Field work included observations,
an investigation of the use of gazetted rights and forest products and local
botanical knowledge at Niah NP and recording plants in verification plots in
seven vegetation types. The botanical expedition teams collected 303 fertile
specimens of trees, shrubs and herbs and covered most of the vegetation types
represented in the Park including some less well-collected areas, such as the
summit of Gunung Subis, and areas to the Park’s east.
The specimens were subsequently identified by reference to the literature
and comparison with specimens 1n the Herbarium, Sarawak Forest Department
(SAR) and, for the Araceae, Macaranga species, Orchidaceae, Sterculiaceae
and Zingiberaceae, with the help of experts. The identification of species of
Piper, Syzygium, Lauraceae and Urticaceae posed problems as up-to-date
revisions and/or well-annotated SAR herbarium specimens were not available.
The checklist includes specimens collected, recorded or observed during
the DANIDA/SWMPI Project period, some mentioned in the literature and
some SAR collections reliably identified by specialists. Fern classification is
according to Parris and Latiff (1997).
Vegetation Types in the Park
Though small, the Park includes six distinct vegetation types, namely:
¢ Limestone vegetation on karst (rough limestone country with
underground drainage) covers approximately 16 km? or c. 40% of
the Park’s area). It includes both the massif and surrounding scattered
limestone fragments, large and small, which provide habitats for a
variety of limestone plants; Sg Subis drains forest west of the
limestone massif and flows over limestone bedrock in places
¢ Mixed Dipterocarp Forest (MDF) covering c. 16% of the Park
chiefly on gentle to steep slopes to the Park’s southwest where
weathering has exposed the sandstone that underlies the limestone
(Sg Trusan drains part of this MDF) and also a patch traversed by
the Plank Walk from the Park Head Quarters to the Great Cave and
also, south of this, a patch of MDF-Kerangas forest traversed by the
Jalan Madu Trail
Plants of Niah National Park, Borneo 103
113°A0' 113°45' 113°50' 113°55' 114°00
Elevationinfeet [ ) Catchment Boundary
[—] TPA Boundary
4 High Point
JV Rivers
/\/ Permanent Streams
*.” Seasonal Streams
Infrastructure
© Communities
/ 1 Main Road
‘ Tracks
4 Trails
GB 950 - 1050
- 1050 - 1480 /V/ Proposed pat Road
1 0 1 2 3 4 Kilometres aK
a ee | x E
s
The National Park boundary marked on the map is approximate
only and that demarcated on the ground shall be considered
correct
Prepared by the DANIDA/SWMPI Project (August 2003)
113°40’ 113°45' 113°50' 113°S5" 114°00'
Map 1. Topography of Niah National Park and Surrounding Areas.
(reproduced with permission of the Sarawak Forest Department).
104 Gardens’ Bulletin Singapore 56 (2004)
¢ Seasonal Swamp Forest on clayey marl soils covers c. 10% of the
Park on the Sg Niah and Sg Tangap floodplains, which are
periodically inundated, sometimes to a depth of 3 m (the height of
stranded debris), when, after intense rainfall, the rivers flood their
banks and rainwater is slow to drain
¢ Seasonal Swamp Forest on peat soils covers c. 5% of the Park and
is found in a patch of low-lying and periodically inundated land that
lies between the levees of Sg Niah and Sg Tangap at the north of the
Park and which has a different forest type from seasonal swamp
forest on clayey marl soils
¢ Riparian Forest fringes the tidal and navigable Sg Niah and its
slow-moving tributaries, Sg Tangap and Sg Sekaloh, the three rivers
that bound the Park to the west, northeast and south, respectively
¢ Regenerating Forest covers c. 29% of the Park in areas that have
been disturbed.
Limestone vegetation on karst
Karst hills provide a variety of microhabitats (Kiew 1991). Niah National Park’s
karst landscape includes summits, steep to vertical cliffs rising to more than
100 m, narrow gorges, broad valleys, complexes of extensive and spectacular
caves and a pool, “Telaga Kumang’, located at the base of a karst wall.
The Subis massif summits are of jagged karst rocks with pockets of
humus and soil in crevices (acidic where litter has accumulated) and surrounding
skeletal (< 25 cm deep) mineral soils with rock particles. The massif’s high
point, Gunung Subis at 394 m, does not match altitudes reached by karst at
Mulu and few collections mention the existence of a summit peat or ‘mor’
layer such as occurs on even quite low hills in the Kuching Division (Kiew et
al. 2004).
Summit vegetation is characterized by a few species of trees.
Gymnostoma nobile (ru ronang), with its characteristic rounded, soft-looking
crowns of fine needle-twigs, is typical and abundant on undisturbed summits
and there are also shrubs, numerous herbs and epiphytes. Orchids and ferns
are well represented and Paraboea speluncarum*, with its distinctive rosettes
of pale leaves, is abundant.
Vegetation may differ from summit to summit. Exposed karst rocks and
surrounding humus soil at the summit of Gunung Subis (partly disturbed by
the setting up of a camp) support Ficus deltoidea and F: sumatrana; Schefflera
burkillii and Schefflera species, Medinilla botryocarpa*, a species each of
E
Endemic to Borneo
Plants of Niah National Park, Borneo 105
Cayratia, Mussaenda and Fagraea and orchid species while just below the
summit were Pandanus calcinactus®, Podocarpus confertus, the jewel orchid
Dossinia marmorata™, species of Hemigraphis and Justicia and the fern
Paragramma longifolia. These species were not noted on the only accessible
portion of Bukit Kasut summit (220 m, burnt in the past and suffering high
visitor pressure). Altitude may be a factor restricting the distribution of
Pandanus calcinactus and Schefflera burkillii to the summit of Gunung Subis
since SAR specimens of the former species have (with one exception) only
been collected from an altitude of at least 400 m and specimens of the latter
from at least 780 m. However, SAR specimens of Podocarpus confertus have
been collected from as low as 100-133 m altitude.
Species found at the summit of Bukit Kasut included Macaranga
endertii®, the ferns Antrophyum parvulum, Drynaria quercifolia, Phymatosorus
scolopendria, and species of Asplenium, Pyrrosia and Loxogramme; as well as
Dischidia albida (Asclepiadaceae) and the legume Ormosia sumatrana, none
of which were noted on the summit of Gunung Subis.
Burning has clearly modified summit vegetation. Distinct differences
between the vegetation of burnt and un-burnt areas are apparent on Bukit Kasut,
said to have been burnt c. 1997, and on summits opposite the West Mouth of
the Great Cave, burnt in 1988 due to a lightning strike. There is much exposed
limestone both at the summit and on slopes of the burnt area of Bukit Kasut
and vegetation comprises scattered trees and shrubs to c. 4 m tall, with no
Gymnostoma nobile, and a herb layer that includes drought resistant plants,
such as Paraboea speluncarum and Phymatosorus sp. By comparison, a peak
immediately to the southeast, separated by a ravine and thus inaccessible, is
dominated by well-developed specimens of G. nobile to at least 10 m tall, with
an understorey of shorter trees and bare limestone visible only on sheer vertical
slopes. Similarly, the skyline of the cliff facing the Great Cave West Mouth,
viewed from the West Mouth, has sparse G. nobile to the left (south) but a
well-developed canopy of G. nobile and other species to the right (north).
Visitor pressure on steep, open parts of the trail leading to Bukit Kasut
(the only trail to a limestone summit easily accessible to park visitors and also
the habitat of the endemic Begonia kasutensis) has loosened the friable soil in
places.
Exposed limestone faces, including cliffs and the sides of shady sinkholes,
are typically dry and support little growth besides ferns and scattered clinging
shrubs and herbs. On the other hand, limestone slopes can be comparatively
well vegetated with scattered trees, some to 30 m tall, and an open canopy.
This habitat supports some large dipterocarps including species of Shorea (lun
or damar hitam and selangan batu), Dipterocarpus and Dryobalanops including
D. lanceolata (kapur paji) and D. beccarii (kapur bukit), Koompassia excelsa
(tapang), Hopea andersonii (luis somit), Artocarpus sp., Popowia spp., Ficus
spp. and Eusideroxylon zwageri (belian) are present. Herbs include aroids and
ferns.
106 Gardens’ Bulletin Singapore 56 (2004)
Exposed limestone hills and cliffs may support scattered large trees and
smaller trees, such as Mallotus miquelianus, Streblus macrophyllus® and
Hydnocarpus gracilis. Herbs include gingers, Costus, aroids and ferns, such
as Asplenium nidus (bird’s nest fern) that may reach a very large size, A.
salignum and Antrophyum parvulum.
Shaded limestone outcrops occur in the seasonal swamp forest. They
vary in size and shape and provide a range of niches differing in exposure and
thus light, humidity and moisture availability, surface features (presence or
absence and tilt of eroded bedding planes, which give the appearance of worn
brickwork), aspect, inclination and presence of soil pockets in crevices and
associated sinkholes and small ravines. Large strangling fig trees rooting into
these outcrops are conspicuous and abundant. Lithophytes include aroids,
orchids, ferns (e.g. Cyclopeltis presliana), begonias and the limestone specialists
Elatostema acuminata, Epithema sarawakensis, and Monophyllaea merrilliana.
Caves at Niah provide a continuum of plant microhabitats differing in substrate
(combinations of bare rock/guano/soil), light and moisture availability (rock
surfaces may be dry, damp or wet), inclination, aspect, altitude, etc. In the
deepest shade algae and patches of liverwort cover the rocks. Ferns (Tectaria
devexa, Tectaria sp., Asplenium spp.) occupy shaded areas, as do Cyrtandra
spelaea®, Clerodendrum sp. and Elatostema acuminata.
At the Great Cave entrance, karst specialist species include the ferns
Heterogonium pinnatum and Cyclopeltis presliana, Epithema sarawakensis
and two Begonia species. On exposed karst rocks and dry, soil-covered slopes
in the same area six Ficus (fig) species, Alstonia scholaris (pelai) and Piper
(wild pepper) occur. On slopes just outside the mouth of the Great Cave,
cultivated/exotic species that have been planted or have escaped include
Sauropus androgynus (cankok manis - a local vegetable, also recorded from
Gua Pangomah in the Park), Capsicum sp. (chilli), Areca catechu (pinang),
Alocasia macrorrhizos (keladi), Carica papaya (papaya) and an ornamental
Codiaeum. Weeds (Mikania cordata, Ageratum conyzoides and Oxalis
corniculata) also occur in this habitat.
Aroid species in limestone habitats at Niah NP included Amorphophallus
brachyphyllus, A. hewittii, Arisaema filiforme, Homalomena humilis,
Raphidophora lobbii and Schizmatoglottis elegans, S. niahensis and
Schizmatoglottis sp. (the last is possibly a new species - A. Hay, pers. comm.).
Many limestone plants have very exacting niche requirements. For
example, four of Niah’s five Begonia species and Monophyllaea merrilliana
each occupied highly localized limestone microhabitats.
Mixed Dipterocarp Forest
The mixed dipterocarp forest (MDF) in the Niah comprises impressive, mature
trees, including emergents. Trees exceeding 100 cm dbh (exceptionally 200
Plants of Niah National Park, Borneo 107
cm dbh) were observed. The understorey comprised numerous saplings and
seedlings, woody climbers and a few herbs. The MDF is likely to be of the
humult ultisol type typically found on sandy soils with a surface horizon of
loosely matted and densely rooted raw humus, low nutrient status and low
water retention capacity (Ashton 1995). Dryobalanops lanceolata, Shorea
superba and Dipterocarpus caudiferus, which can indicate this type of MDF,
occur at Niah (D. lanceolata was observed in the Park). According to Ashton,
humult ultisol MDF ‘appears to be the richest in species of all forest types in
Malaysia, because of its remarkable endemism’.
Large trees north of Sg Sekaloh and along Sg Trusan include Shorea
spp. (meranti, selangan batu), Dipterocarpus spp. (keruing), Sindora leiocarpa
(tampar hantu), Anacardiaceae sp. (rengas) and Chrysobalanceae sp. (merbatu).
The MDF on the sandstone slopes at the Park’s southwest included
Dryobalanops spp. (kapur), Dipterocarpus confertus (keruing kobis) and
Shorea quadrinervis (meranti sudu). In this area, belian has been extracted
and Antiaris toxicaria (ipoh) tapped. The ground flora includes aroids.
Seasonal Swamp Forest on Clayey Marls
Seasonal swamp forest in Niah NP occurs on somewhat acid clayey marls, has
a broken canopy, no emergents (some trees reach 20 m but the majority are
shorter), and a dense understorey with herbs chiefly represented by climbers.
There is low tree species diversity but high levels of gregariousness. Octomeles
sumatrana (benuang), Pterospermum subpeltatum (bayur) and Pandanus
basilocularis (a stilted screw-pine that reaches 10 m and is abundant beside
the plank walk to the Painted Cave) are gregarious in this forest. Dracontomelum
dao (sengkuang), Pometia pinnata (kasai) and Anthocephalus cadamba
(kelampayan) also occur in this habitat. Much of the seasonal swamp forest at
Niah has been disturbed. There are large belian stumps and the canopy is
generally broken.
In seasonal swamp forest below the West Mouth of the Great Cave,
belian, Diospyros euphlebia’, Ficus spp. and Saurauia sp. (mata ikan)
predominate. Aroids carpet the ground and some, e.g. Pothos scandens, climb.
Forest transitional between riverine and seasonal swamp forest occurs
between Kuala Sg Subis and the foot of Bukit Kasut. Here are large fig trees,
Mallotus floribundus (buantik), Macaranga (benuang), sengkuang, gregarious
Schoutenia glomerata and Lithocarpus blumeanus (empili). Rattans (Calamus
sp., Daemonorops sparsiflora, Korthalsia sp.) and Caryota sp., erect and
creeping aroids, Nepenthes (pitcher plants), wild Piper species (including the
locally abundant P. sarmentosum), Costus speciosus, gingers (Boesenbergia
sp. and Globba sp.) and urticaceous herbs, as well as a number of woody and
herbaceous climbers, occur here. At the base of Bukit Kasut and on the way to
Telaga Kumang, mosses grow on rocks beside small channels.
108 Gardens’ Bulletin Singapore 56 (2004)
Seasonal Swamp Forest on Peat Soils
At the north of the Park, a small area between the levees of Sg Niah and Sg
Tangap had, where sampled, an organic horizon >150 cm deep and peat soil
with 76% loss on ignition (peat soil has >65% loss on ignition). Small channels
and pools of standing water occur. This forest had a broken canopy (extraction
was reported to have occurred in the past), some emergents to 25 m and shorter
trees to 20 m or less, a dense understorey with abundant saplings and seedlings
of primary forest species, herbs, woody climbers and abundant bryophytes on
tree roots.
Peat swamp timber species that occur in this forest include Copaifera
palustris (sepetir paya), Dactyloclados stenostachys (jongkong), Gonystylus
sp. (ramin), Shorea sp. (meranti), Dryobalanops rappa (kapur paya) and
Combretocarpus rotundatus (keruntum) as well as large trees of Parishia sp.
(upi). There are also seedlings of Parishia maingayi (upi paya) and Vatica
mangachapoi (resak paya).
Other trees include the riverine kasai, Elaeocarpus stipularis (sengkurat)
and Vatica umbonata, the swamp species Quassia indica, and other trees
including figs, Stemonurus secundiflorus var. lanceolatus (semburok),
Blumeodendron tokbrai (merbulan), Diospyros pilosanthera (kayu malam),
Syzygium spp. (ubah), Memecylon sp. (nipis kulit) and Santiria laevigata. The
swamp palm Eleiodoxa conferta (asam paya) and the aroids Raphidophora
korthalsii and R. lobbii occur here.
Riparian Forest
The banks of Sg Niah, where it bounds the Park, are occasionally flooded and
are occasionally under saline influence as the river is tidal at this point. A well-
developed riparian fringe includes abundant Lithocarpus blumeanus, frequent
sengkurat and buantik and frequent, large specimens of sengkuang. The climber
Willughbeia sp. (akar kubal) is moderately frequent and kasai scattered.
Dipterocarpus oblongifolius (ensurai) was recorded. Fruit trees that have been
planted behind the riverbanks include Durio zibethinus (the cultivated durian),
two wild Durio species with edible fruits (nyakak and isu), Mangifera payang
(embang) and Artocarpus odoratissimus (terap).
Sungai Sekaloh appears to have lost most of its original riparian
vegetation. Sungai Subis, a muddy, slow flowing river with muddy banks,
flows through seasonal swamp forest at the north and west of the Subis massif.
No macrophytes were noted either in the water or on the banks 1n this moderately
to deeply shaded habitat. To the west of the Subis massif, towards Kuala Subis,
the stream is bigger and the riverside trees sengkuang, kasai and benuang are
common while fig trees are abundant.
Plants of Niah National Park, Borneo 109
Regenerating Forest
About 29% of the Park is now in various stages of regeneration after having
been disturbed. Some of the area was formerly used for shifting cultivation.
Some timber extraction has taken place. Regeneration, occurring over different
time intervals, on different soil types and presumably after different types and
degrees of disturbance, is by no means uniform. A plot set in old secondary
forest, where a charcoal layer in the clay loam to sandy loam soil suggested
past clearance for agriculture, had emergents reaching 30 m and 110 cm dbh,
though most trees were 25 m or less. In this 20 x 20 m plot there was a dense
understorey of numerous saplings of primary forest species, a few herbs, some
woody climbers and almost no ferns or palms. The mix of primary forest species
in this plot included only one individual belonging to the Dipterocarpaceae.
The Park’s Flora
The flora of Niah NP is as yet incompletely known. To date, more than 800
plant specimens from Niah NP are deposited at SAR. This includes more than
300 specimens collected during the SWMPI project botanical field
investigations. Orchids and palms appear to be under-collected. New species
of aroid (A. Hay, P. Boyce, pers. comm.) Paraboea, Schumannianthus and
Boesenbergia (A. Poulsen, pers. comm.) are still to be described from specimens
collected in the park.
Orchid species reported with certainty from the Park to date are
surprisingly few. Beaman ef al. (2001) list nine genera and species
(Agrostophyllum sp., Bulbophyllum minutulum, B. vaginatum, Calanthe
triplicata, Coelogyne verrucosa, C. pandurata, Didymoplexiella borneensis€,
Dossinia marmorata* and Plocoglottis borneensis). However, they mention
31 orchid species as originating from ‘Niah’ (six terrestrial, one lithophytic
and two saprophytic) and at least some of these are likely to be found in the
Park as all but four are noted as occurring at least occasionally on karst. The
SWMPI study has added Agrostophyllum glumaceum, Liparis elegans and
Thecostele alata while Anoectochilus sp. was photographed and Claderia
viridiflora observed just off park trails.
Dossinia marmorata 1s an attractive jewel orchid of karst found also at
Bau and Mulu. Adenoncos triloba®, an epiphytic orchid of lowland karst hills
described in 1935, is known only from the type specimen from Niah. Beaman
et al. (2001), applying IUCN Red List Categories to orchid taxa known only
from Sarawak, consider Adenoncos triloba to be Critically Endangered and
both Malaxis andersonii (a terrestrial karst species) and Didymoplexiella
borneensis (a saprophyte of hill forest sometimes found on karst) as
_ Endangered. There are unconfirmed reports that a slipper orchid
(Paphiopedilum sp.) occurs near the karst summits at Niah.
1 10 Gardens’ Bulletin Singapore 56 (2004)
Orchids not associated with karst include two attractive terrestrial species,
Plocoglottis borneensis and Calanthe triplicata. |
Palms observed, reported or recorded from in and around the Park
include Caryota no, Areca andersonii", which is known only from the type
specimen from Niah (Dransfield 1984) and from Brunei , Licuala borneensis*
(previously collected in 1867 in Tubau but no longer there (Saw Leng Guan
pers. comm.) and at Niah collected in secondary forest), and rattans including
Calamus myriacanthus®, C. caesius and C. scipionum, Daemonorops
microstachys® and D. sparsiflora*.
Well-represented groups at the Park include:
ferns and fern allies - 31 species including Taenitis trilobata (Holttum 1968)
described from Niah and known from only two localities in Sarawak; possibly
two new records for Sarawak (Adiantum cf. malesianum and Asplenium
subnormale), Mesophlebion oligodictyon (originally described as Dryopteris
oligodyctia from Niah) and five species (Syngramma wallichii, Lindsaea
borneensis, Tectaria aurita, Leptochilus macrophyllus and Monogramma
paradoxa) collected by H. Schneider (pers. comm.),
Acanthaceae with 9 species including Cosmianthemum dido, described from
within the Park,
Annonaceae with 15 species,
Araceae with 19 species includes Schizmatoglottis niahensis and Alocasia
venusta, which are both endemic to Niah NP,
Begoniaceae with five Begonia species, all new to science and endemic to
Gunung Subis (Pearce 2003),
Euphorbiaceae with 13 species,
Gesneriaceae includes Cyrtandra cretacea® (Burtt 1970) and C spelaea® (Burtt
1978), both described from Niah and the latter endemic to the Park;
Monophyllaea merrilliana (Monophyllaea had not previously been reported
from Niah), and Paraboea speluncarum and Paraboea sp. nov. endemic to
the Park,
Moraceae - Ficus is especially diverse with eight species and Streblus
macrophyllus is a new record for Sarawak,
Rubiaceae with 14 species,
Urticaceae is represented by 11 species, and
Vitaceae includes Ampelocissus capillaris and a Cayratia from the summit of
Gunung Subis that could not be matched at SAR.
This array of plant groups largely mirrors those Anderson (1965) reported
as being well represented on limestone in Sarawak (Annonaceae, Begoniaceae,
Euphorbiaceae, Gesneriaceae and Moraceae).
The Park is reported to be rich in both wild and cultivated fruit tree
species. Informants interviewed regarding gazetted rights to collect fruits, etc.,
in the Park mentioned a total of more than 30 fruit tree species. These include
species of Mangifera (Anacardiaceae), Willughbeia (Apocynaceae), Durio
Plants of Niah National Park, Borneo 111
(Bombacaceae), Lansium (Meliaceae), Artocarpus (Moraceae) and Nephelium
(Sapindaceae). Other species include Gnetum gnemon (meninjau), Pangium
edule (kepayang) and Dialium indum (keranj1).
Useful species in the Park include Eurycoma longifolia (tongkat alt),
locally well-known owing to its reputed aphrodisiac properties; Dyera
polyphylla (jelutong paya) formerly tapped for latex for export to make chewing
gum (a tree showing tapping scars is just beside the plank walk to the Great
Cave and jelutong is said to have been harvested for timber in the Park);
Goniothalamus velutinus (kayu hujan panas, lim panas)—a treelet common on
sandy soil and kerangas forest that is locally highly valued based on its supposed
magical powers to protect against evil forces; Antiaris toxicaria (ipoh) - a tree
tapped for its cardiac-glycoside-containing latex, an ingredient of blowpipe
dart poison; palms with edible cabbages; rattan canes (Calamus caesius (sega)
and other species) for tying scaffolding used to collect edible birds’ nests in
caves, for handicrafts and for weaving; and Pandanus andersonii (kerupok)
and P. ?vinaceus - (akas or akah) whose leaves are collected for making mats.
Endemicity
Little information on endemicity is available for most plant groups in Sarawak.
Wong (1998) listed endemic species of selected taxa in Borneo, which indicated
endemicities ranging from 8% (Aglaia) to 92% (Vaccinium).
Limestone forest in Borneo is rich in endemics particularly among
herbaceous species (Ashton 1995). Wong (1998) noted that nearly all of the 15
Paraboea species in Borneo are locally endemic to karst and 16 of the 17
species of Monophyllaea are specifically confined to it. Wong mentioned
Cyrtandra species and Pandanus calcinactus as Sarawak karst plants and notes
that the ‘inselberg’ (island mountain) nature of the majority of the karst outcrops
in Borneo appears to have promoted speciation among such groups as the
gesneriads, balsams and begonias. Likewise, Kiew (1991), with reference to
the limestone flora in Peninsular Malaysia, states that for some herbaceous
groups such as the Paraboeas, recent speciation appears to have occurred as a
result of isolation, so that many species are confined to a particular hill or
group of hills.
The Subis massif is indeed well isolated from other Sarawak karst
formations (Bau and Padawan-Serian in the southwest; Bukit Sarang southeast
of Bintulu; Mulu to the northeast, and the Upper Baram limestone in the
interior). So far, the following endemics from the Subis massif have been
reported: Alocasia venusta and Schismatoglottis niahensis (Araceae), Begonia
kachak, B. kasutensis, B. niahensis, B. stichochaete and B. subisensis
(Begoniaceae), Tectaria andersonii and T. subdigitata (Dryopteridaceae)
(Holttum 1991) both known only from the type specimens, Cyrtandra spelaea
112
Gardens’ Bulletin Singapore 56 (2004)
Table 1. List of species endemic to Borneo reported from Niah National Park
*endemic to the Niah National Park, excluding new species
Adiantaceae
Annonaceae
Araceae
Arecaceae
Aspleniaceae
Begoniaceae
Dryopteridaceae
Ebenaceae
Euphorbiaceae
Gesneriaceae
Melastomataceae
Moraceae
Orchidaceae
Pandanaceae
Adiantum malesianum Ghatak
Goniothalamus calcareus Mat Salleh
Goniothalamus velutinus Airy Shaw
Orophea sarawakensis Kessler
*Alocasia venusta A. Hay
*Schismatoglottis niahensis A. Hay
Areca andersonii J.Dransf.
Calamus myriacanthus Becc.
Daemonorops microstachys Becc.
Licuala borneensis Becc.
Asplenium subnormale Copel.
*Begonia kachak K. G. Pearce
*B. kasutensis K. G. Pearce
*B. niahensis K. G. Pearce
*B. stichochaete K. G. Pearce
*B. subisensis K. G. Pearce
*Tectaria andersonii Holttum
*T. subdigitata (Baker) Copel.
Diospyros euphlebia Merr.
Macaranga endertii Whitmore
Cyrtandra cretacea Kraenzlin
*C. spelaea B. L. Burtt
*Paraboea speluncarum (Burtt) Burtt
Medinilla botryocarpa Regalado
Ficus hemselyana King
Streblus macrophyllus Blume
*Adenoncos triloba Carr
Didymoplexiella borneensis (Schltr.) Garay
Dossinia marmorata E.Morren
Pandanus calcinactus St John ex Stone
Plants of Niah National Park, Borneo 113
and Paraboea speluncarum (Gesneriaceae) and Adenoncos triloba
(Orchidaceae), known only from the type specimen. In addition, five species
are probably new to science but have yet to be described: Schefflera ‘niah’
(Araliaceae), which grows on mudstone outcropping from riverbank (D. Frodin
pers. comm.), Schismatoglottis (Araceae) from the foot of the limestone hill
(A. Hay, pers. comm.), an Anadendrum (P.C. Boyce, pers. comm.), a Paraboea
(Gesneriaceae), a Schumannianthus (Marantaceae, A. Poulsen pers. comm.),
and a Boesenbergia sp. collected by A. Poulsen (A. Poulsen pers. comm..).
Conclusion
Niah NP is remarkable on account of the variety of vegetation types represented,
including a variety of limestone habitats, as well as its history of human
interaction. That the Park should support a rich and varied flora is not
unexpected in view of the diversity of substrates, relief (especially in karst
areas) and processes (flooding, fire and human disturbance, including shifting
cultivation and extraction of timber and forest products). While the limestone
is relatively dry and mostly without peat formation at summits, it still supports
a number of plants that have yet to be described or could not be identified by
reference to the SAR collection, as well as new records for Sarawak and Niah.
Inaccessible parts of the limestone are still to be explored botanically.
Meanwhile, a range of stresses impacts the Park. These include its isolation by
surrounding development, human disturbance by bird nest and guano collectors,
visitors and, just outside the Park boundary, quarrying, and the depletion of
Park resources (birds’ nests, guano, timber, forest products, ornamentals, hunted
wildlife, etc). An initiative to fully explore, interpret and conserve this special
area is urgently needed.
Acknowledgements
This paper is an outcome of botanical field investigations at Niah National
Park carried out under the Sarawak Forest Department/DANIDA Project
Support to Wild Life Master Plan Implementation through the Improved
Management of Totally Protected Areas, Sarawak, Malaysia (SWMPI). The
author is grateful to Julian T. Inglis, Chief Technical Advisor of the SWMPI
Project, for the opportunity to undertake a botanical expedition to, and a study
of the use of gazetted rights at Niah National Park under the Project and to the
Sarawak Forest Department for permission to reproduce the map. Ruth Kiew,
expert in limestone floras of the region, provided both the encouragement for
this paper to be started and completed and comments on the manuscript. The
author is most grateful for identifications of the Niah material kindly provided
114 Gardens’ Bulletin Singapore 56 (2004)
by Peter Boyce and Alistair Hay (Araceae), Stuart Davies (Macaranga),
Anthony Lamb (Orchidaceae), Axel Poulsen (Zingiberaceae) and Peter Wilkie
(Sterculiaceae). The author is also most grateful to the staff of the Botany
Unit, Forest Research Centre and of the Niah National Park, National Parks
and Wildlife Department, Sarawak Forest Department and would like to thank
members of the local communities at Niah who participated in the survey of
gazetted rights and/or assisted during collecting trips.
References
Anderson, J.A.R. 1965. Limestone habitat in Sarawak. Ecological Research in the Humid
Tropics Vegetation. Kuching, Government of Sarawak & UNESCO: 49-57.
Ashton, P.S. 1995. Biogeography and ecology. Tree Flora of Sabah and Sarawak. 1: xliti-li.
Beaman, T.E., J.J. Wood, R.S. Beaman and J.H. Beaman. 2001. Orchids of Sarawak. Natural
History Publications (Borneo), Kota Kinabalu, Sabah, Malaysia.
Burtt, B.L. 1970. Studies in the Gesneriaceae of the Old World XXXII: Some species of
Cyrtandra, chiefly Bornean. Notes Royal Botanic Gardens Edinburgh. 30: 23-42.
Burtt, B.L. 1978. Studies in the Gesneriaceae of the Old World XIV: New and little known
species of Cyrtandra, chiefly from Sarawak. Notes Royal Botanic Garden Edinburgh. 36:
157-179.
Dransfield, J. 1984. The genus Areca (Palmae: Arecoideae) in Borneo. Kew Bulletin. 39: 1-22.
Hay, A. 1998. The genus Alocasia (Araceae-Colocasieae) in West Malesia and Sulawesi.
Gardens Bulletin Singapore. 50: 221-334.
Holttum, R.E. 1968. A re-definition of the fern-genus Taenitis Willd. Blumea. 16: 87-95.
Holttum, R.E. 1991. Tectaria group, Flora Malesiana Series 2, 2: 1-132.
Kiew, R. 1991. The Limestone Flora. In: R. Kiew (ed.) The State of Nature Conservation in
Malaysia. Malayan Nature Society, Kuala Lumpur, Malaysia. Pp. 42—50.
Kiew, R., C. Geri, S. Julia, P.-C. Boyce. 2004. Understorey flora. In: Sarawak Bau Limestone
Biodiversity. H.S. Yong, F.S.P. Ng and E.E.L. Yen (eds.). Sarawak Museum Journal. 59
(No. 80, New Series), Special Issue No. 6: 105-146.
Parris, B.S. and A. Latiff. 1997. Towards a pteridophyte flora of Malaysia: A provisional
checklist of taxa. Malaysian Nature Journal. 50: 235-280.
Pearce, K.G. 2003. Five new Begonia species (Begoniaceae) at the Niah National Park, Sarawak,
Malaysia. Gardens Bulletin Singapore. 55: 73-88.
Wong, K.M. 1998. Patterns of plant endemism and rarity in Borneo and the Malay Peninsula.
Academia Sinica Monograph Series. 16: 139-169.
Plants of Niah National Park, Borneo £15
Appendix: Preliminary Checklist of Plants Collected and
Reported from Niah National Park
Note: Most descriptions of plants and habitats are taken directly from field notes. The
specimens listed are deposited in the Herbarium, Sarawak Forestry Department (SAR).
emperan is the Sarawak term for alluvial forest
kerangas is the Iban word for forest that when cleared is not suitable for growing hill padi
* species associated with limestone at Niah National Park
E species endemic to Borneo
G Gunung (mountain)
MDF mixed dipterocarp forest
NP National Park
Sg Sungai (river or stream)
Tj Tanjung (headland)
ACANTHACEAE
Cosmianthemum dido Burtt & Smith
Pathside in primary forest. Pengkalong Lobang to Niah Cave B.L. Burtt & P.J.B. Woods B
1998.
Cosmianthemum magnifolium Brem.
Herb 25—40 cm tall. Disturbed peat swamp forest, open area by trail, peat soil. Along N Park
boundary from Tj Belipat Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89235;
MDF, brown clay soil. Sekaloh Jemree Sablee et al. S 89017.
*Hemigraphis sp.
Woody shrub 60 cm tall. Exposed limestone rocks and humus soil, open, canopy 5 m tall.
Just below summit of G Subis. Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S
89429.
*Justicia sp.
Woody shrub c. | m tall. Exposed limestone rocks and humus soil, open, canopy 5 m tall.
Just below summit of G Subis Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S
89430.
*Linariantha bicolor Burtt & Smith
Creeping herb. Limestone forest at foot of limestone hill, light shade, in soil at base of
limestone rocks. On path to Telaga Kumang Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri
Johari S 89410.
?Polytrema sp.
Herb 25 cm tall. On trail, open area, clay soil. Near Sg Subis Kit Pearce, Julaihi Abdullah &
Serukit Dubod S 89457.
*) Pseuderanthemum sp.
Shrub 40 cm tall. Foot of limestone hill. Ulu Sg Tangap Jemree Sablee et al. S 89072.
116 Gardens’ Bulletin Singapore 56 (2004)
*Pryssiglottis sp. }
Semi-woody plant c. 60-200 cm tall. Forest at base of limestone crags, light shade, humus
soil also high ground, light shade by path, loamy soil (locally common) Kit Pearce, Yahud
Wat, Serukit Dubod & Jeffri Johari S 89272; Along N Park boundary from Tj Belipat. Kit
Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89236.
Acanthaceae sp.
Creeping herb. On trail in open, clay soil. Near Sg Subis Kit Pearce, Julaihi Abdullah &
Serukit Dubod S 89456.
ADIANTACEAE
*Adiantum cf. malesianum © Ghatak
Creeping herb c.15 cm tall. Light shade in forest, humus soil among limestone outcrops,
under wall of limestone, below summit of G Subis Kit Pearce, Yahud Wat, Serukit Dubod &
Jeffri Johari S 89438.
Syngramma alismifolia (C. Presl) J.Sm.
Herb 30 cm tall. Disturbed peat swamp forest, open area by trail, peat soil. Along N Park
boundary from Tj Belipat Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89219.
Syngramma wallichii Hook.
Collected by H. Schneider (pers. comm.).
Taenitis trilobata Holttum
Niah. C. Hose s.n., 1890. (in Holttum, 1968).
ANACARDIACEAE
*Dracontomelum dao (Blanco) Merr. & Rolfe
Tree c. 1 m girth, 24 m high. S side of G Subis, in Sg Sekaloh, Niah. J.A.R. Anderson, Sonny
Tan & E. Wright S 27577.
Mangifera pajang Kosterm.
Observed behind the banks of Sg Niah.
Semecarpus rufovelutinus Ridley
Tree 8 m tall, c. 5 cm diam. Disturbed peat swamp forest, open area by trail, peat soil with
pools of standing water. Along N Park boundary from Tj Belipat Kit Pearce, Yahud Wat,
Serukit Dubod & Jeffri Johari S 89216.
ANISOPHYLLACEAE
Combretocarpus rotundatus (Miq.) Danser
Observed in seasonal swamp forest on peat soils in the N of Park.
ANNONACEAE
Cyathocalyx cf. havilandii Boerl.
Treelet 2 m tall. Disturbed forest, moderate shade. Tj Belipat, along boundary from Sg Niah,
100 m from river Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89203.
Ellipeia cuneifolia J.D.Hook. & Thompson
Woody climber 12 m tall, base of stem 6 cm diam. Seasonal swamp forest (disturbed), light
shade, clay soil. Trail into park from Kuala Sg Subis Kit Pearce, Yahud Wat, Serukit Dubod
& Jeffri Johari § 89284.
Plants of Niah National Park, Borneo V7
*Enicosanthum macranthum (King) J. Sinc.
Tree 10 m tall, 16 cm diam. Alluvial limestone lowland forest at the foot of limestone hills
from Rumah Chang, Sg Tabau Julaihi Abdullah, Kit Pearce et al. S 89321.
Goniothalamus cf. calcareus® Mat Salleh
Treelet 1.5—2 m tall, 1 cm diam. MDF, brown clay soil also secondary forest, moderate shade,
loam soil. Sekaloh. Jemree Sablee et al. S 89008; Close to Sg Niah Kit Pearce, Yahud Wat,
Serukit Dubod & Jeffri Johari S 89251.
*Goniothalamus cf. uvarioides King
Tree 7 m tall, 6 cm diam. Limestone forest, alluvial, medium shade. Gua Sibau, trail from
Rumah Chang, NE of the Park. Julaihi Abdullah, Kit Pearce et al. S 89318.
Goniothalamus velutinus © Airy Shaw
Treelet 2 m tall, 2 cm diam. MDF, brown clay soil. Sekaloh Jemree Sablee et al. S 89018.
Orophea kostermansia Kessler
Tree to 6 m tall, 20 cm diam. Shady lowland alluvial forest. Along plank walk to Niah Caves
Julaihi Abdullah, Kit Pearce et al. S 89305.
*Orophea sarawakensis © Kessler
Tree 7 m tall, 5 cm diam. S boundary of park, E of Quarry Kit Pearce, Julaihi Abdullah &
Serukit Dubod S 89495.
Polyalthia cauliflora J.D.Hook. & Thompson var. beccarii (King) Sinclair
Tree 6 m tall, 7 cm diam. MDF, Sg Sekaloh Jemree Sablee et al. S 89031.
Polyalthia motleyana (Hk.f.) Airy Shaw var. motleyana
Tree 4 m tall, 7 cm diam. MDF, Sg Sekaloh Jemree Sablee et al. S 89036.
Popowia pisocarpa (Blume) Endl.
Tree c 7 m tall, 5 cm diam. Secondary forest, moderate shade, loam soil. Close to Sg Niah Kit
Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89250.
*” Popowia
Small tree 3 m tall, 6 cm diam. Limestone forest. G Subis Jemree Sablee et al. S 89046.
*Uvaria grandiflora Roxb. ex Homem.
Woody climber 17 m tall, 2 cm diam. Limestone forest, light shade, humus soil among limestone
rocks at the top of limestone summit Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S
89404.
Uvaria sp.
Woody climber c. 15 m tall, stem 3 cm diam. Disturbed peat swamp forest, open area by trail,
peat soil. Along N Park boundary from Tj Belipat Kit Pearce, Yahud Wat, Serukit Dubod &
Jeffri Johari S 89225.
Woodiellantha sympetala (Merr.) Rauschert
Small tree 8 m tall, 7 cm diam. MDF. Sg Sekaloh Jemree Sablee et al. S 89024.
118 Gardens’ Bulletin Singapore 56 (2004)
APOCYNACEAE
*Alstonia scholaris R. Br.
Observed at the mouth of the Great Cave.
Dyera polyphylla (Miq.) Ashton
Observed along the plank walk to the Great Cave.
Tabernaemontana macrocarpa Jack
Tree 7 m tall, 6.5 cm diam. Disturbed forest, grey clay under 25 cm peat, moderate shade.
Close to Sg Niah Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S &9257.
Tabernaemontana pauciflora Blume
Shrub | m tall. Seasonal swamp forest (disturbed), open area. Beside plank walk to Great
Cave Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89469.
AQUIFOLIACEAE
*Tlex sp.
Tree to 3 m tall, 10 cm. diam. Shady lowland alluvial forest. Limestone hill next to Quarry
Julaihi Abdullah, Kit Pearce et al. S 89313.
ARACEAE
Aglaeonema nebulosum N.E.Br.
Herb c.15—20 cm tall. Tj Belipat, along boundary from Sg Niah Kit Pearce, Yahud Wat, Serukit
Dubod & Jeffri Johari S 89208; On path to Telaga Kumang Kit Pearce, Yahud Wat, Serukit
Dubod & Jeffri Johari S 89409.
Aglaeonema nitidum (Jack) Kunth
Herb 60-100 cm tall. Just beside inundated area with standing water, clay soil, moderate
shade, also seasonal swamp forest (disturbed), light shade and secondary forest, moderate
shade, loam soil. Tj Belipat, along boundary from Sg Niah Kit Pearce, Yahud Wat, Serukit
Dubod & Jeffri Johari S 89211; Beside plank walk to Great Cave Kit Pearce, Julaihi Abdullah
& Serukit Dubod S 89464; Close to Sg Niah Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri
Johari S 89247.
*Alocasia venusta A. Hay
Herb to 45 cm tall. Lithophytic on limestone in swamp forest c. 200 m alt. Path between Niah
town and Niah caves Hay et al. 9346.
*Amorphophallus brachyphyllus Hett.
Lithophytic herb c. 1.5 m tall, stem c. 60 cm, 2.5 cm diam. Disturbed forest among limestone
rocks, light shade also shady lowland alluvial forest. Below W mouth of Great Cave Kit
Pearce, Julaihi Abdullah & Serukit Dubod S 89487, Along plank walk to Niah Caves Julathi
Abdullah, Kit Pearce et al. S 89309.
*Amorphophallus hewittii Alderw.
Herb | m tall. Limestone forest, light shade, humus soil amongst limestone rocks. Top of
limestone summit Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89408.
Anadendrum microstachyum (deVr. & Miq.) Baker & Alderw.
Herbaceous climber 1.5 m tall. Emperan forest, dark brown clay soil, also MDF, light shade,
Plants of Niah National Park, Borneo 1 19
loam soil Jemree Sablee et al. S 89006; Across river from Park HQ Kit Pearce, Yahud Wat,
Serukit Dubod & Jeffri Johari S 89444.
Anadendrum sp. nov.
Climbing herb 1 m tall. Cut petiole without raphides, immature fruits yellow-green. Disturbed
swamp forest in standing water, moderate shade, grey clay with wet humus layer. Close to Sg
Niah Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89264.
*Arisaema filiforme Blume
Herb 30 cm tall. Forest at base of limestone crags, light shade, humus soil Kit Pearce, Yahud
Wat, Serukit Dubod & Jeffri Johari S 89273.
Cryptocoryne zonata de Wit
Aquatic herb with leaves slightly above water surface. In water in dry regenerating peat swamp
forest also in stagnant pool, light shade, disturbed forest. Jemree Sablee et al. S 89041; Across
river from Park HQ Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89440.
Homalomena cf. hostifolia Engl.
Herb 20 cm tall. Undisturbed MDF, medium shade, loamy soil Kit Pearce, Yahud Wat, Serukit
Dubod & Jeffri Johari S 89268.
*Homalomena humilis (Jack) Hk.f.
Herb15 cm tall. Limestone forest on stone, also on limestone rocks in forest at base of limestone
crags in light shade, stony mineral soil and on limestone rocks in pockets of mineral soil,
medium shade. G Subis Jemree Sablee et al. S 89042; Kit Pearce, Yahud Wat, Serukit Dubod
& Jeffri Johari S 89275; Near Sg Subis Kit Pearce, Julaihi Abdullah & Serukit Dubod S
89459.
Homalomena sp.
Herb 70 cm tall. MDF, light shade, loamy soil. At base of limestone crags Kit Pearce, Yahud
Wat, Serukit Dubod & Jeffri Johari S 89279.
Lasia spinosa (L.) Thwaites
Herb c.1.3 m tall. Just beside inundated area with standing water, clay soil, moderate shade,
among litter. Tj Belipat, along boundary from Sg Niah Kit Pearce, Yahud Wat, Serukit Dubod
& Jeffri Johari S 89209.
Raphidophora korthalsii (tenuis facies) Schott.
Herb c. 50 cm tall. Disturbed peat swamp forest, open area by trail, peat soil. Along N Park
Boundary from Tj Belipat Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89234.
*Raphidophora lobbii Schott.
Climber 10-350 cm tall. Disturbed peat swamp forest among limestone rocks, also open area
by trail, peat soil with pools of standing water, light shade. Below W mouth of Great Cave Kit
Pearce, Julaihi Abdullah & Serukit Dubod S §9486; Along N Park boundary from Tj. Belipat
Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89217; Kit Pearce, Yahud Wat, Serukit
Dubod & Jeffri Johari S 89222.
*Schismatoglottis elegans A. Hay
Lithophytic herb c. 50 cm tall. Limestone forest, alluvial, medium shade, on limestone outcrops.
Gua Sibau, trail from Rumah Chang, NE of Park Julaihi Abdullah, Kit Pearce et al. S 89319.
120 Gardens’ Bulletin Singapore 56 (2004)
*Schismatoglottis niahensis A. Hay
Clustering herb 0.6 m tall. Limestone forest, medium shade, peaty soil at base of limestone
rocks. Near Sg Subis Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89454
*Schismatoglottis puberulipes Alderw.
Limestone. Hay et al. 9369.
*Schismatoglottis sp.nov. (P.C. Boyce, pers. comm.)
Herb 25 cm tall. Leaf upper surface lustrous, dark green, pale beneath, spathe pale green
with brownish purple bracts, ovaries green, stamens (above) cream. Limestone forest at foot
of limestone hill, light shade, in soil at base of limestone rocks. On path to Telaga Kumang Kit
Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89411.
ARALIACEAE
*Schefflera burkillii Merr.
Herbaceous climber 1.5 m tall. Open summit of G Subis Kit Pearce, Yahud Wat, Serukit
Dubod & Jeffri Johari S 89413.
*Schefflera petiolosa (Miq.) Harm.
Lithophytic shrub 1.5—5 m tall. Limestone rock in secondary seasonal swamp forest also
disturbed seasonal swamp forest, light shade, by stream. Trail into park from Kuala Sg Subis
Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89290; Beside Plank Walk to Great
Cave Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89467.
Schefflera sp. nov. (D. Frodin, pers. comm.)
Epiphyte. Greenish orange fruits. Ulu Sg Sekaloh, Sg Niah J.A.R. Anderson S 27296.
*Schefflera sp.
Herb | m tall, climbing on rocks. Open summit of G Subis. Kit Pearce, Yahud Wat, Serukit
Dubod & Jeffri Johari S 89412.
ARECACEAE
*Areca andersonii © J.Dransf.
Palm 1-2. 6 m tall, 1.5 cm diam. MDF, Sekaloh Jemree Sablee et al. S 89019; Sg Sekaloh, G
Subis, Niah J.A.R. Anderson S 31937.
Areca minuta Scheff.
Palm 2.5 m tall, 1 cm diam at base. MDF/Kerangas forest, moderate shade, sandy loam soil.
Jalan Madu Trail Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89447.
Calamus myriacanthus © Becc.
Rattan 2—7 m tall. MDF, light shade, loam soil. Across river from Park HQ Kit Pearce, Julathi
Abdullah & Serukit Dubod S 89445 & S §9446.
Calamus myriacanthus © Becc.
(All Borneo Rattan Database, J. Dransfield, 2000)
Caryota no Becc.
Observed in the valley outside the West Mouth of the Great Cave.
Plants of Niah National Park, Borneo 121
Daemonorops korthalsii Blume
Rattan c. 10 m tall. Disturbed secondary forest, moderate shade, clay soil. Along N Park
boundary from Tj Belipat. Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89212.
*Daemonorops longipes (Griff.) Mart.
Rattan 6 m long. Foot of limestone, 17 m asl. G Brangin, Ulu Sg Subis, Niah NP Yii Puan
Ching S 40174.
Daemonorops sparsiflora Becc.
Rattan 10 m long. Riverine forest, along Sg Niah, Niah NP Yii Puan Ching S 40147.
Iguanura curvata Kiew
Palm c. 1 m tall. In disturbed alluvial forest with silty clay soil (not very common). Niah NP
Paul Chai S 40056.
Licuala borneensis ® Becc.
Clustered palm 80 cm tall. Secondary forest, moderate shade, loam soil. Close to Sg Niah.
Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89248.
Licuala sp. 1
Solitary to 1 m tall. Shady lowland alluvial forest. Along plank walk to Niah Caves Julaihi
Abdullah, Kit Pearce et al. S 89301.
Licuala sp. 2
Clustered palm | m tall. Disturbed swamp forest, in standing water, moderate shade, grey
clay with wet humus layer. Close to Sg Niah. Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri
Johari § 89263
*Pinanga albescens Becc.
Palm in shade on limestone rock at base of limestone hill /.A.R.Anderson S 16028.
ASCLEPIADACEAE
*Dischidia albida
Vegetation verification plot specimen at summit of Bukit Kasut.
*Dischidia hirsuta (Blume) Decne.
Small climbing epiphyte. Lower slopes of limestone hill, large limestone boulders. Sg
Sekaloh, G Subis, Niah J.A.R. Anderson S 31924.
*Dischidia cf. tubulifera King & Gamble
Small climber. Summit ridge of limestone, 270 m altitude, limestone strongly dissected and
covered by 15-22 cm peat layer. Sg Sekaloh, G Subis, Niah J.A.R. Anderson S 31930
ASPLENIACEAE
* Asplenium macrophyllum Sw.
Terrestrial fern. Rare on limestone rocks in semi-exposed position, slope between caves,
limestone rocks and residual soil, trees mainly felled, habitat semi-exposed. also inside Gua
Tulang on rock. Great Cave, G Subis, Niah J.A.R. Anderson S$ 31916; Mohidin S 21691;
Niah Ahmad 104.
122 Gardens’ Bulletin Singapore 56 (2004)
*Asplenium nidus L.
On tree in old forest. G Subis Ahmad 102.
*Asplenium nitidum Sw.
Epiphyte on rock. Shady lowland alluvial forest. Along plank walk to Niah Caves Julaihi
Abdullah, Kit Pearce et al. S 89307.
*Asplenium salignum Blume
Herb (may be epiphytic) 60 cm tall. Disturbed limestone forest on and among rocks, light
shade. Below W mouth of Great Cave Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89485;
Ulu Sg Sekaloh, near Quarry Jemree Sablee et al. S 89069.
*Asplenium scandens J.Sm. ex Mett.
Herb c. 50 cm tall. On limestone rocks, light shade, almost no soil. Near Sg Subis Kit Pearce,
Julaihi Abdullah & Serukit Dubod S 89461.
*Asplenium subnormale © Copel.
Creeping herb. On limestone rocks in pockets of mineral soil, medium shade also shady lowland
alluvial forest. Near Sg Subis Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89453; Along
plank walk to Niah Caves Julaihi Abdullah, Kit Pearce et al. S 89304.
*Asplenium vittaeforme Cav.
Epiphytic fern on fallen branch. Foot of limestone forest, sandy soil. Ulu Sg Trusan Jemree
Sablee et al. S 89056.
BEGONIACEAE
*Begonia kachak K.G.Pearce
Creeping herb, 11—25 cm tall. Foot of limestone hill, on rocks, light shade, almost no soil also
shade at base of limestone cave wall. Great Cave Jemree Sablee et al. (S 89062), Kit Pearce,
Bibian Diway, Saupel Atot and Dami Jude S 78536; Near Sg Subis Kit Pearce, Julaihi Abdullah
& Serukit Dubod S 89463.
*Begonia kasutensis K.G.Pearce
Creeping fleshy herb c. 10 cm tall. Limestone forest also on steep path in soil between limestone
rocks where canopy cover starts to thin out. G Subis Jemree Sablee et al. S 89049; On route to
Bukit Kasut Kit Pearce & Narawi Johari S 78596.
*Begonia niahensis K.G.Pearce
Sprawling herb c. 40 cm long. On limestone rocks, light shade, almost no soil also exposed
steeply sloping limestone rocks, rooted in a pocket of soil in the rocks. Near Sg Subis Kit
Pearce, Julaihi Abdullah & Serukit Dubod S 89460; Great Cave Kit Pearce, Bibian Diway,
Saupel Atot and Dami Jude S 78537.
*Begonia stichochaete K.G.Pearce
Erect herb 15—25 cm tall. Undisturbed MDF, medium shade, loamy and brown clay soil, also
disturbed emperan forest, deep alluvial soil, light shade also on low limestone outcrops and in
soil surrounding limestone outcrops in deep shade and damp conditions. Sekaloh Jemree Sablee
et al. § 89014; Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89267; Banks of Sg
Niah near confluence with Sg Subis Kit Pearce, Bibian Diway, Saupel Atot and Dami Jude S
78539; On route to Bukit Kasut Kit Pearce & Narawi Johari S 78597.
Plants of Niah National Park, Borneo 123
*Begonia subisensis K.G.Pearce
Herb c. 45 cm tall. Near summit on limestone rooted in crevices in soil pockets, partial shade
also on steep slopes between limestone rocks where canopy cover starts to thin out (locally
frequent). Bukit Kasut Kit Pearce, Bibian Diway, Saupel Atot and Dami Jude S 78538; On
trail to Bukit Kasut Kit Pearce & Narawi Johari S 78598.
BIGNONIACEAE
*?Tecomanthe sp.
Woody climber 5 m tall. In forest with canopy c. 20 m tall, light shade, humus soil among
limestone outcrops. Below summit of G Subis Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri
Johari S 89436.
CASUARINACEAE
Gymnostoma nobile (Whitmore) L.A.S. Johnson
Observed on summits. Abundant.
CELASTRACEAE
Microtropis sp.
Treelet 1.5 m tall, 2 cm diam. MDF. Sg Sekaloh Jemree Sablee et al. S 89034.
CHLORANTHACEAE
*Chloranthus erectus (Buch.-Ham.) Verdc.
Herb c. | m tall. Limestone forest, alluvial, medium shade. Gua Sibau, trail from Rumah
Chang, NE of park Julaihi Abdullah, Kit Pearce et al. S 89324.
CHRYSOBALANACEAE
Parinari sp.
Tree c. 40 m tall, 90 cm diam above 6 m tall buttresses. Secondary forest, moderate shade,
loam soil. Close to Sg Niah Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89252.
CLUSIACEAE
*?Mesua/Mammea/Kayea
Tree 2.2 m tall, 1.5 cm diam. Forest at base of limestone crags, light shade, stony mineral soil
Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89269.
CONVALLARIACEAE
*Peliosanthes teta Andr. ssp. humilis (Andr.) Jessop
Erect herb c. 35 cm tall. Limestone forest, medium shade, in litter at base of limestone rock.
Abundant in this seasonally wet location. Beside plank walk to Great Cave Kit Pearce, Julaihi
Abdullah & Serukit Dubod S 89472.
COSTACEAE
Costus speciosus (Koenig) R.M. Smith
CRY PTERONIACEAE
Dactyloclados stenostachys Oliv.
Observed in seasonal swamp forest on peat soils in the N of the Park.
CYPERACEAE
Hypolytrum nemorum (Vahl) Spreng
Herb c. 80 cm tall. Along N Park boundary from Tj Belipat Kit Pearce, Yahud Wat, Serukit
Dubod & Jeffri Johari S 89241.
124 Gardens’ Bulletin Singapore 56 (2004)
DATISCACEAE
Octomeles sumatrana Miq.
Medium sized tree 23 m tall, 1 m girth. River bank on sandy soil. Ulu Sg Sekaloh, Sg Niah
J.A.R. Anderson S 27298.
DENNSTAEDTIACEAE
Lindsaea borneensis
Collected by H. Schneider (pers. comm.).
Lindsaea doryphora Kramer
Terrestrial herb c. 25 cm tall. Disturbed peat swamp forest, moderate shade, peat soil. S of
trail from Tj Belipat along N Park boundary Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri
Johari S$89233.
DIPTEROCARPACEAE
Dipterocarpus confertus Slooten
Observed in MDF in the SW of the Park.
Dipterocarpus oblongifolius Blume
Large tree 3 m girth, 17 m tall. On riverbank, sandy soil. About 1 km upstream from Kuala
Sekaloh of Sg Niah. Banyeng & Adenan S 22571.
Dryobalanops beccari Dyer
Observed in MDF on route to G Subis.
Dryobalanops lanceolata Burck
Observed in MDF.
Dryobalanops rappa Becc.
Observed in seasonal swamp forest on peat soils in the N of the Park.
*Hopea andersonii Ashton ssp. andersonii
Large tree 2 m girth, 33 m tall. On limestone rock near river on S slope of G Subis in Sg
Sekaloh, Niah. Sibat & Adenan S 22566; G. Subis, Niah, S side near Sg Sekaloh J.A.R. Anderson
S 31645.
*Hopea dryobalanoides Miq.
Tree 2 m girth, 27 m tall. On limestone rock near river on S slope of G Subis in Sg Sekaloh,
Niah. Sibat & Adenan S 22563.
*Hopea pachycarpa (Heim.) Sym.
Tree 2 m girth, 27-37 m tall. On limestone rock near river, also on lower slopes of Cana
hill and on limestone boulders on southern slope of G Subis in Sg Sekaloh, Niah. Sibat &
Adenan § 22565; Sonny Tan & E. Wright S 27262.
Parashorea macrophylla Wyatt-Smith
Observed along Jalan Madu Trail, MDF/Kerangas forest.
Shorea agami Wood. ssp. agami
Tree. Lowland poorly drained site. Niah NP. S.H. Rogstad 725.
Plants of Niah National Park, Borneo 125
*Shorea falciferoides Foxw. ssp. glaucescens Meijer
G. Subis area, Niah. Mohidin S 21681.
*Shorea guiso (Blanco) Blume
Large tree 2.7 m girth, 37 m tall. Lower slopes of limestone hill, numerous limestone boulders.
Southern slopes of G Subis, near Sg Sekaloh Sonny Tan & E. Wright S 27265.
*Shorea havilandii Brandis
Medium-sized tree, 75-120 cm girth, 13-23 m tall. Lower slopes of steep limestone hill,
numerous limestone boulders. South side of G Subis, in Sg Sekaloh, Niah J/.A.R. Anderson,
Sonny Tan & E. Wright S 27555; Sibat & Adenan S 22570.
*Shorea parvifolia Dyer
Large tree about 40 m high, 100 cm diam. On relatively steep slope of small river valley in
rather open Dipterocarpus forest. Limestone area of Bukit Subis to right of Sg Sekalau, to
right of footpath from Kuala Sekalau to Bukit Drusau, 120 m asl. H.P. Fuchs 21275.
*Shorea patoiensis Ashton
Tree 15-17 cm girth and 23 m tall. On limestone rock near river, south side of G Subis, in Sg
Sekaloh, Niah Sibat & Adenan S 22567.
Shorea quadrinervis Slooten.
Observed in MDF in the SW of the Park.
*Vatica chartacea (Ashton) Kost.
Tree 15 m tall, 45 cm girth. Foot of limestone 17 m asl, G Branging, Ulu Sg Subis, Niah NP.
Yii Puan Ching S 40177.
*Vatica rynchocarpa Ashton
Small tree 87 cm girth, 17 m tall. On riverbank, sandy soil. About '/, mile upstream from
Kuala Sekaloh of Sg Niah Banyeng & Adenan S 22572.
Vatica mangachapoi Blanco
Observed in seasonal swamp forest on peat soils in the N of the Park.
DRY OPTERIDACEAE
*Cyclopeltis presliana J.Sm.
Lithophytic or terrestrial herb 50 cm high. Foot of limestone forest, sandy soil, on limestone
cliff also rocks in pockets of mineral soil, medium shade. Ulu Sg Trusan Jemree Sablee et al.
S 89058; Near Sg Subis Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89458.
*Heterogonium pinnatum (Copel.) Holttum
Herb 50 cm tall. Limestone forest on soil on rock on hillside, moderate shade, loamy soil with
humus. SW of Telaga Kumang Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89297.
*Tectaria andersonii Holttum
Terrestrial fern in shaded localities on limestone outcrop. Upper slopes of limestone hill, 12.5
m below summit ridge, limestone rocks outcropping and deep ‘mor’ soil. Sg Sekaloh, G
Subis, Niah J.A.R. Anderson S 31963
Tectaria aurita (Sw.) S. Chandra
Collected by H. Schneider (pers. comm.).
126 Gardens’ Bulletin Singapore 56 (2004)
*Tectaria devexa (Kunze) Copel.
Lithophytic herb c 25 cm tall. On limestone rocks in pockets of soil between rocks. Medium
shade. Beside plank walk to Great Cave Kit Pearce, Julaihi Abdullah & Serukit Dubod S
89478.
*Tectaria labrusca (Hook.) Copel.
Herb c. 60 cm tall on rocks. Forest canopy c. 20 m tall, light shade, humus soil among limestone
outcrops. Below summit of G Subis Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S
89437.
Tectaria subdigitata (Baker) Copel.
Rheophyte. Niah C. Hose 196 (from Holttum, 1991).
*Tectaria ternata (Baker) Copel.
Terrestrial. Foot of limestone forest, sandy soil. Ulu Sg Trusan Jemree Sablee et al. S 89052.
EBENACEAE
*Diospyros euphlebia © Merr.
Tree 34 m tall, 3-4 cm diam. Disturbed forest among limestone rocks also foot of limestone
hill, light shade. Below W mouth of Great Cave Kit Pearce, Julaihi Abdullah & Serukit Dubod
S 89481; Ulu Sg Tangap Jemree Sablee et al. S 89073.
*Diospyros pilosanthera (Blanco) var. pilosanthella
Large tree about 30 m high, 80 cm diam. On steep slope of small river-valley in rather open
Dipterocarpus-forest. Limestone area of G Subis to right of Sg Sekalau, to right of footpath
from Kuala Sekalau to Bukit Drusau, 120 m asl. H.P. Fuchs 21275.
Diospyros pyrrhocarpa Miq.
Tree c. 20 m tall, c. 15 cm diam. By path, moderate shade, wet clay soil. Along N Park
boundary from Tj Belipat Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89238.
*Diospyros sumatrana Miq.
Small tree 3.5—5 m tall, 2-8 cm diam. Dry regenerating peat swamp forest also limestone
forest, alluvial, medium shade. Jemree Sablee et al. S 8904; Gua Sibau, trail from Rumah
Chang, NE of park Julaihi Abdullah, Kit Pearce et al. S 89322.
*Diospyros toposia Ham. var. toposioides (King & Gamble) Phengklai
Tree 10 m tall, 5 cm diam. Limestone forest, light shade, humus soil among limestone rocks.
Top of limestone summit. Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89405.
? Diospyros sp.
Woody climber [sic] c. 7 m tall, 5 cm diam. Just beside inundated area with standing water,
clay soil, moderate shade. Tj Belipat, along boundary 50 m from Sg Niah Kit Pearce, Yahud
Wat, Serukit Dubod & Jeffri Johari S 89207.
ELAEOCARPACEAE
Elaeocarpus stipularis Blume
Shrub about 6 m tall. Riverine forest about 17 m asl, soil loamy. Kuala Sg. Sekaloh, Niah NP
Yii Puan Ching S 40105.
Plants of Niah National Park, Borneo 407
EUPHORBIACEAE
*Antidesma montanum Blume
Tree 2—5 m tall, 3 cm diam. Alluvial limestone lowland forest, on limestone cliff also disturbed
peat swamp forest, open area by trail, peat soil. Foot of limestone hills from Rumah Chang,
Sg Tabau Julaihi Abdullah, Kit Pearce et al. S 89326; Along N Park boundary from Tj. Belipat
Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89223.
Antidesma neurocarpum Miq.
Tree 5 m tall, 4 cm diam. Undisturbed MDF, medium shade, loamy soil Kit Pearce, Yahud
Wat, Serukit Dubod & Jeffri Johari S 89271.
Antidesma tomentosum Blume var. tomentosum
Small tree 3—7 m tall, 3-10 cm diam. Emperan forest, dark brown clay soil also MDF/kerangas
forest, moderate shade, sandy loam soil. Jemree Sablee et al. S 89005; Jalan Madu Trail Kit
Pearce, Julaihi Abdullah & Serukit Dubod S 89450.
Baccaurea sp.
Tree 9 m tall, 18 cm diam. MDF, brown clay soil. Sekaloh Jemree Sablee et al. S 89012.
Blumeodendron tokbrai (Blume) Kurz.
Tree 20 m tall, 75 cm girth. Peat swamp forest. Niah NP. Bernard Lee M.H. S 40065.
?Claoxylon
Treelet 1.5 m tall, 2 cm diam. MDF, Sg Sekaloh Jemree Sablee et al. S 89035.
Cleidion javanicum Blume
Tree 7-10 m tall, 15—20 cm diam. Seasonal swamp forest (disturbed) also shady lowland
alluvial forest, open area. Beside plank walk to Great Cave Julaihi Abdullah, Kit Pearce et al.
S 89308; Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89470.
*Cleistanthus megacarpus C.B.Rob.
Tree 6-8 m tall, 4-10 cm diam. Alluvial limestone lowland forest also by path, moderate
shade, wet clay soil. Foot of limestone hills from Rumah Chang, Sg Tabau Julaihi Abdullah,
Kit Pearce et al. S 89325; Along N Park Boundary from Tj. Belipat Kit Pearce, Yahud Wat,
Serukit Dubod & Jeffri Johari S 89237.
Croton oblongus Burm.f. .
Treelet c. 9 m tall, 18 cm diam. Just beside inundated area with standing water, clay soil,
moderate shade. Tj Belipat, along boundary from Sg Niah (200 m from river) Kit Pearce,
Yahud Wat, Serukit Dubod & Jeffri Johari S 89204.
Drypetes microphylla (Merr.) Pax et Hoffm.
Tree 8 m tall, 8 cm diam. MDF/kerangas forest, moderate shade, sandy loam soil. Jalan Madu
Trail Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89449.
*? Frismanthus obliquus Wall ex Muell.-Arg
Small tree 4—7 m tall, 7-10 cm diam. Limestone forest, light shade, humus soil amongst
limestone rocks. G Subis Jemree Sablee et al. S 89043; Top of limestone summit Kit Pearce,
Yahud Wat, Serukit Dubod & Jeffri Johari S 89401.
128 Gardens’ Bulletin Singapore 56 (2004)
*Macaranga endertii Whitmore |
Tree 35 cm girth, 15 m high. Lower slopes of limestone hill, numerous limestone boulders. S
slopes of G Subis, near Sg Sekaloh. Sonny Tan & E. Wright, S 27275.
Mallotus floribundus (Blume) Muell.-Arg.
Tree 6-10 m tall, 3-17 cm diam. Just beside inundated area with standing water, clay soil,
moderate shade also seasonal swamp forest (disturbed), light shade, clay soil, beside water
channel. Tj Belipat, along boundary from Sg Niah Kit Pearce, Yahud Wat, Serukit Dubod &
Jeffri Johari S §9206; Trail into park from Kuala Sg Subis Kit Pearce, Yahud Wat, Serukit
Dubod & Jeffri Johari S 89289.
*Mallotus miquelianus (Scheff.) Boerl.
Treelet 2 m tall, 1.5 cm diam. Limestone forest among rocks on hillside, moderate shade,
loamy soil with humus. SW of Telaga Kumang Kit Pearce, Yahud Wat, Serukit Dubod &
Jeffri Johari S 89295.
*Wetria insignis (Steud.) Airy Shaw
Tree 20 m tall, 4 cm diam. Limestone forest, light shade, humus soil amongst limestone
rocks. Top of limestone summit. Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari §
89403 :
*Euphorbiaceae sp.
Tree 4 m tall, 2 cm diam. On humus soil among limestone outcrops. Beiow summit of
G Subis Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89434.
FABACEAE (PAPILIONOIDEAE)
Canavalia cathartica Thouars
Semi-woody plant to 0.5 m tall. Shady lowland alluvial forest. Along plank walk to Niah
Caves Julaihi Abdullah, Rit Pearce et al. S 89310.
Copaeifera palustris (Sym.) de Wit
Observed in seasonal swamp forest on peat soils in the N of the Park.
Koompassia excelsa (Becc.) Taubert
Observed in MDF on route to G Subis.
Ormosia sumatrana (Miq.) Prain
Observed on the summit of Bukit Kasut.
FAGACEAE
Lithocarpus blumeanus (Korth.) Rehder
Observed in forest transitional between riverine and seasonal swamp forest.
Lithocarpus gracilis (Korth.) Soepadmo
Tree c. 25 m tall, 45 cm diam. Seasonal swamp forest (disturbed), light shade, clay soil. Trail
into park from Kuala Sg Subis Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89288.
Lithocarpus sp.
Tree c. 5 m tall, 15 cm diam. Disturbed peat swamp forest, moderate shade, peat soil. S of
trail from Tj Belipat along the N boundary of Niah NP. Kit Pearce, Yahud Wat, Serukit Dubod
& Jeffri Johari S 89228.
Plants of Niah National Park, Borneo 129
FLACOURTIACEAE
Casearia grewiifolia Vent.var. gelonioides (Bl.) Sleumer
Treelet 2 m tall, 1.5 cm diam. Disturbed peat swamp forest, open area by trail, peat soil.
Along N Park Boundary from Tj Belipat Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri
Johari S 89218.
Hydnocarpus gracilis (Sloot.) Sleum.
Tree 20 m tall, 45 cm diam. MDF, Sg Sekaloh Jemree Sablee et al. S 89029.
GESNERIACEAE
*Aeschynanthus tricolor Hook.
Epiphyte. On trees on swampy alluvial soils at base of limestone hill, subject to periodic
inundation. On southern slopes of G Subis in Sg Sekaloh, Niah J.A.R. Anderson, Sonny Tan &
E. Wright S 26086.
*Aeschynanthus cf. tricolor Hook.
Herbaceous climber c. 10 m up on trunk of tree. Limestone forest (disturbed including by
logging), on exposed rocks, shady area, soil peaty humus between rocks. S boundary of Park,
E of Quarry Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89494.
*Aeschynanthus parvifolius R.Br.
Epiphytic on tree. Foot of limestone hill, Great Cave Jemree Sablee et al. S 89065.
*Cyrtandra bracheia B.L.Burtt
Shrublet 45—60 sm high. Broad sloping ravine between two vertical limestone hills, numerous
limestone boulders, some residual soil (very localized). On southern slopes of G Subis in the
Sg Sekaloh, Niah J.A.R. Anderson, Sonny Tan & E. Wright S 26092.
*Cyrtandra cretacea® Kraenzl.
On rough limestone rocks just before reaching caves. Niah Caves B.L. Burtt & P.J.B. Woods
B 1991; Haviland & Hose 3530.
Cyrtandra erectipila B.L.Burtt
Herb 30 cm tall, 1 cm diam. MDF. Sg Sekaloh Jemree Sablee et al. S 89033.
*Cyrtandra incrustata® B.L.Burtt
Solitary herb or small shrublet 15—22 cm tall. In limestone forest on exposed rocks, medium
shade also base of limestone cliff, somewhat dry and shaded at 100 m altitude. S boundary of
the Park, E of Quarry Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89496; South side of G
Subis, in Sg Sekaloh, Niah J.A.R. Anderson, Sonny Tan & E. Wright S 27575.
*Cyrtandra oblongifolia (Blume) C.B.Clarke
Epiphytic shrub. In limestone forest on rocks, medium shade, also on limestone rock at 50 m
altitude. S Boundary of park, E of Quarry Kit Pearce, Julaihi Abdullah & Serukit Dubod S
89499; Valley in front of Niah Caves B.L. Burtt & P.J.B. Woods B 2037; Ulu Sg Subis, Niah
NP Bernard Lee S 40083.
*Cyrtandra spelaea*B.L.Burtt
Prostrate herb or shrublet 60-90 cm tall. Back of shallow cave, base of cave wall, somewhat
dry and completely shaded, slopes of limestone hill immediately below cliff c 130 m altitude,
130 Gardens’ Bulletin Singapore 56 (2004)
litter layer overlying limestone boulders, abundant on vertical limestone rock face. Niah Cave
B.L. Burtt & P.J.B. Woods B 2018 & B 3020 and Alphonso & Samsuri A 239; Sg Sekaloh, G
Subis J.A.R. Anderson § 31958; Great Cave J.A.R. Anderson S 31918; Inside Niah Cave on
rock Ahmad 11; Niah Haviland & Hose 3531; Synge 615.
*Cyrtandra trisepala C.B.Clarke
Herb or shrublet 85 cm tall. On limestone rocks and pavement, gully between limestone hill,
heavily dissected, mineral soil very sparse. G Subis J.A.R. Anderson S 31671.
*Cyrtandra tubiflora Kraenzl.
On bare limestone, shaded. Valley below main Niah Cave B.L. Burtt & P.J.B. Woods B 2015.
*Epithema sarawakensis Hill & B.L.Burtt
Herb. Limestone forest on stone, G Subis Jemree Sablee et al. S 89048.
Henckelia cf violoides (C.B.Clarke) B.L.Burtt
Niah. Haviland & Hose 3534.
*Monophyllaea merrilliana Kraenzl.
Herb. Infrequent on damp limestone rock in deep shade Kit Pearce & Narawi Johari S 78595.
*Paraboea speluncarum (B.L.Burtt) B.L.Burtt
Herb or shrublet. Cave and large overhang 100 m up on steep limestone hill, light dry soil,
partly composed on guano, also forest. Gua Pangomah, G. Subis, Niah J.A.R.Anderson S
31675; Niah Caves Alphonso & Samsuri 241.
Paraboea sp. nov.
Large herb 75-120 cm tall. On bare, exposed limestone rock near summit of limestone pinnacles
or crest of limestone hill, ‘mor’ soil, 130 m asl, Niah J.A.R. Anderson S 16045; S slope of G
Subis near Sg Sekaloh Sonny Tan & E. Wright S 27279.
GNETACEAE
Gnetum gnemon L.
Tree 5 m tall, 2 cm diam. Disturbed secondary forest, moderate shade, clay soil. Along N Park
boundary from Tj. Belipat Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89215.
*Gnetum leptostachyum Blume
Woody climber 10 m tall, 2.5 cm diam. Limestone forest among rocks on hillside, moderate
shade, loamy soil with humus. SW of Telaga Kumang Kit Pearce, Yahud Wat, Serukit Dubod
& Jeffri Johari § 9298.
Gnetum neglectum Blume
Woody climber or liana at least 10 m tall, 2 cm diam. Secondary forest, moderate shade, loam
soil under 15 cm humus, moderate shade. Close to Sg Niah Kit Pearce, Yahud Wat, Serukit
Dubod & Jeffri Johari S 89259 & S 89246.
HANGUANACEAE
Hanguana malayana (Jack) Merr.
Herb | m tall. Disturbed swamp forest, in standing water, moderate shade, grey clay with wet
humus layer. Close to Sg Niah Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89242.
Plants of Niah National Park, Borneo 131
ICACINACEAE
Gonocaryum macrophyllum (Blume) Sleumer
Treelet 4 m tall, 2.5 cm diam. Secondary forest, moderate shade, loam soil. Close to Sg Niah
Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89245.
Stemonurus secundiflorus var. lanceolatus
Tree 15 m tall, 10 cm diam. Disturbed peat swamp forest, open area by trail, peat soil. Along
N Park boundary from Tj Belipat Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S
89226.
LAURACEAE
*Dehaasia brachybotrys (Merr.) Kosterm.
Tree 5 m tall, 5 cm diam. Foot of limestone hill. Ulu Sg Tangap Jemree Sablee et al. S 89076.
Eusideroxylon zwageri Teijsm. & Binn.
Cut stumps observed in alluvial habitats, on hills and in damp valley.
Litsea ?oppositifolia (Blume) Vill.
Tree c. 6 m tall. Just beside inundated area with standing water, clay soil, moderate shade. T}
Belipat, along boundary from Sg Niah Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari
S 89210.
Litsea sp: 1
Treelet 2 m tall, < 1 cm diam. Disturbed peat swamp forest, open area by trail, peat soil.
Along N Park boundary from Tj Belipat Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari
S 89220.
*Litsea sp. 2
Tree 3 m tall, 1.5 cm diam. Limestone forest, alluvial, medium shade. Gua Sibau, trail from
Rumah Chang, NE of Park. Julaihi Abdullah, Kit Pearce et al. S 89317.
Lauraceae sp. 1
Treelet 1.5 m tall, 1 cm diam. Dry regenerating peat swamp forest Jemree Sablee et al. S 89039.
Lauraceae sp. 2 .
Small tree 3 m tall, 4 cm diam. MDF, brown clay soil. Sekaloh Jemree Sablee et al. S 89013.
Lauraceae sp. 3 .
Small tree 5 m tall, 7 cm diam. MDF, brown clay soil. Sekaloh Jemree Sablee et al. S 89016.
LOGANIACEAE
*Fagraea sp.
Tree 2 m tall, 3 cm diam. Exposed limestone rocks with humus soil around, summit of G
Subis Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89420.
LOMARIOPSIDACEAE
Bolbitis heteroclita (C. Presl) Ching
Herb 30 cm tall. Disturbed seasonal swamp forest, light shade, grey clay soil. Sg Subis Kit
Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89294
l 3 ¥ 4 Gardens’ Bulletin Singapore 56 (2004)
LYCOPODIACEAE
Huperzia phlegmaria (L.) Rothm.
Herbaceous epiphyte c. 1.5 m long. Seasonal swamp forest (disturbed), light shade, clay soil.
Trail into Park from Kuala Sg Subis Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S
89283.
MARANTACEAE
Schumannianthus sp. nov.
Clustering herb | m tall. MDF, light shade, loam soil. Across river from Park HQ Kit Pearce,
Julaihi Abdullah & Serukit Dubod S 89442.
MELASTOMATACEAE
*Medinilla botryocarpa* Regalado
Woody shrub | m tall. Exposed limestone rocks with humus soil, summit of G Subis Kit
Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89419.
Medinilla crassifolia (Reinw. ex Blume) Blume
Semi-woody epiphyte 3.5 m up tree. MDF/kerangas forest, moderate shade, sandy loam soil.
Jalan Madu Trail Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89448.
Memecylon paniculatum Jack
Tree 6 m tall, c. 3 cm diam. Disturbed secondary forest, light shade, clay soil. Along N Park
Boundary from Tj Belipat Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89214.
Pternandra cf. crassicalyx Maxwell
Tree 6 m tall, c. 4 cm diam. Secondary forest, moderate shade, loam soil. Close to Sg Niah Kit
Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89253.
MELIACEAE
* Aglaia korthalsii Miq.
Tree 10 m tall, 10 cm diam. Forest on humus soil among limestone outcrops. Below summit
of G Subis Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89435.
*A glaia simplicifolia (Bedd.) Harms
Small tree 4 m tall, 5 cm diam. Foot of limestone forest, sandy soil. Ulu Sg Trusan Jemree
Sablee et al. S 89051.
Chisocheton polyandrus Merr.
Small tree 5 m tall, 4 cm diam. MDF, brown clay soil. Sekaloh Jemree Sablee et al. S 89021.
Dysoxylum excelsum Blume
Tree 10 m tall, 13 cm diam. Emperan forest, dark brown clay soil Jemree Sablee et al. S
89001.
Dysoxylum oppositifolium F.Muell.
Small tree 2 m tall, 6 cm diam. MDF, Sg Sekaloh Jemree Sablee et al. S 89027.
MENISPERMACEAE
?Parabaena hirsuta (Becc.) Diels
Herbaceous climber, twining to 6 m tall. Seasonal swamp forest (disturbed), light shade, clay
soil. Trail into the Park from Kuala Sg Subis Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri
Johari S 89286.
Plants of Niah National Park, Borneo 133
MONIMIACEAE
*Matthaea sancta Blume
Small tree 4 m tall, 7 cm diam. Foot of limestone forest, sandy soil. Ulu Sg Trusan Jemree
Sablee et al. S 89054.
MORACEAE
Antiaris toxicaria Lesch.
Photographed in MDF.
Artocarpus odoratissimus Blanco
Observed behind the banks of Sg Niah.
*Ficus deltoidea Jack var. deltoidea
Woody shrub 70 cm tall or epiphyte. Dry limestone forest, open, with litter and exposed
limestone rocks with humus soil around, open. Limestone hill next to Quarry Julaihi Abdullah,
Kit Pearce etal. S 89311; Summit of G Subis Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri
Johari S 89416.
*Ficus depressa Blume
Epiphyte or huge strangler to 4 m tall, 20 cm diam. Limestone forest or shady lowland
alluvial forest. Ulu Sg Sekaloh, near Quarry Jemree Sablee et al. S 89070; Limestone hill
next to Quarry Julaihi Abdullah, Kit Pearce et al. S 89316.
*Ficus hemselyana © King
Lithophytic shrub c. 2 m tall, 1.5 cm diam at base. Disturbed forest among limestone rocks,
light shade also in limestone forest on exposed rocks, medium shade. Below W mouth of
Great Cave Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89488; S Boundary of Park, E
of Quarry Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89497.
Ficus obscura Blume var. lanata Kochummen
Epiphytic tree c. 4 m tall, 3 cm diam. about 4 m from ground on large tree. Seasonal swamp
forest (disturbed), light shade. Beside plank walk to Great Cave Kit Pearce, Julaihi Abdullah
& Serukit Dubod S 89465.
*Ficus subulata Blume .
Shrub sprawling to c. 4m or much branched lithophytic climber at least 10 m tall, c. 5 cm
diam at base. On limestone rocks in pockets of mineral soil, medium shade. Near Sg Subis
Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89455; Beside plank walk to Great Cave
Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89477.
*Ficus sumatrana (Miq.) Miq.
Woody shrub | m tall. Exposed limestone rocks with humus soil, summit of G Subis
Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89415.
*Ficus tinctoria G.Forst. var. gibbosa (Blume) Corner
Shrubby epiphyte to 1 m long. Shady lowland alluvial forest, limestone hill next to Quarry
Julaihi Abdullah, Kit Pearce et al. S 89312.
Ficus sp.
Epiphyte. Dry regenerating peat swamp forest Jemree Sablee et al. S 89038.
134 Gardens’ Bulletin Singapore 56 (2004)
*Streblus macrophyllus® Blume
Small tree 1.5—2.5 m tall, 1-5 cm diam. Foot of limestone hill also on limestone rocks in
forest at base of limestone crags, light shade, stony mineral soil. Ulu Sg Tangap. Jemree
Sablee et al. (S 89075); At base of limestone crags Kit Pearce, Yahud Wat, Serukit Dubod &
Jeffri Johari S 89274.
MYRISTICACEAE
Knema glauca (Blume) Peterman
Tree 12 m tall, 20 cm diam. MDF, brown clay soil. Sekaloh Jemree Sablee et al. S 89023.
*Myristica guatteriifolia A.DC.
Tree 20 m tall, 30 cm diam. Summit of limestone hill, light shade, with loose limestone
rocks and humus soil. SW of Telaga Kumang Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri
Johari S 89299.
MYRSINACEAE
Ardisia sanguinolenta Blume
Small tree 5—7 m tall, 4-12 cm diam. Emperan forest, dark brown clay soil Jemree Sablee et
al. S 89003 & S 89007.
Labisia pumila Benth. & Hook.f.
Herb 35 cm tall. Disturbed peat swamp forest, open area by trail, peat soil. Along N Park
boundary from Tj Belipat Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89227.
*Systellantha brookeae Stone
Treelet 1—2.5 m tall, 1-1.5 cm diam or shrub c. 40 cm tall. By path, moderate shade, wet clay
soil also foot of limestone hill, also limestone forest (disturbed, including by logging) on
exposed rocks, light shade, soil peaty humus between rocks. Along N Park boundary from
Tj Belipat Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89239; Ulu Sg Tangap.
Jemree Sablee et al. S 89071; S Boundary of park, E of Quarry Kit Pearce, Julaihi Abdullah
& Serukit Dubod S 89493.
MYRTACEAE
*Syzygium hirtum Korth.
Tree 14 m tall, 20 cm diam. Shady lowland alluvial forest. Limestone hill next to Quarry
Julaihi Abdullah, Kit Pearce et al. S 89314.
Syzygium sp. 1
Tree c. 10 m tall, 20 cm diam. Disturbed peat swamp forest, moderate shade, peat soil. S of
trail from Tj Belipat along N Park boundary. Kit Pearce, Yahud Wat, Serukit Dubod & & Jeff
Johari S 89229.
*Syzygium sp. 2
Tree 3 m tall, 3 cm diam. Exposed limestone rocks with humus soil around, summit of G
Subis Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89417.
*Syzygium sp. 3
Small tree 2 m tall, 5 cm diam. Foot of limestone forest, sandy soil. Ulu Sg Trusan Jemree
Sablee et al. S 89055.
Plants of Niah National Park, Borneo £35
*Syzygium sp. 4
Small tree 3 m tall, 3 cm diam. Foot of limestone hill. Great Cave Jemree Sablee et al. S 89066.
*Syzygium sp. 5
Tree 6 m tall, 7 cm diam. Foot of limestone hill. Ulu Sg Tangap Jemree Sablee et al. S
89074.
*Syzygium sp. 6
Tree 7 m tall, 6 cm diam. Forest on humus soil among limestone outcrops, below summit of
G Subis Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89433.
Syzygium sp. 7
Tree 12 m tall, 80 cm diam. Secondary forest, loam soil under 15 cm humus, moderate
shade. Close to Sg Niah Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89261.
NEPENTHACEAE
Nepenthes mirabilis (Lour.) Druce
Climber 4 m long. Disturbed forest, grey clay under 25 cm peat, light shade. Close to Sg
Niah Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89254.
OPHIOGLOSSACEAE
Helminthostachys zeylanica (L.) Hook.
Terrestrial fern, 40 cm tall. On rock near river. Sg Sekaloh Jemree Sablee et al. S 89037.
ORCHIDACEAE
*Adenoncos triloba *Carr
Epiphyte. Lowland limestone hills at 300 m altitude. Niah Synge S 603. (from Beaman et
al., 2001).
*A grostophyllum glumaceum Hook.f.
Epiphytic herb c. 2.5 cm tall, c. 2.5 cm up woody liana. Limestone forest, alluvial, medium
shade. Gua Sibau, trail from Rumah Chang, NE of Park Julaihi Abdullah, Kit Pearce et al.
S 89323.
*A grostophyllum sp.
Epiphyte. Exposed limestone rocks and humus soil, just below summit of G Subis Kit Pearce,
Yahud Wat, Serukit Dubod & Jeffri Johari S 89432.
*Anoectochilus sp.
Terrestrial jewel orchid photographed by trail to Bukit Kasut.
*Appendicula cf. anceps Blume
Herb 30 cm tall. Foot of limestone forest, sandy soil. Ulu Sg Trusan Jemree Sablee et al. S
89060.
*Appendicula cf. uncata Ridley ssp. sarawakensis J.J. Wood
Herb 25 cm tall or epiphytic. Disturbed peat swamp forest, moderate shade, peat soil also
exposed limestone rocks with humus soil around, open. S of trail from Tj Belipat along N
Park boundary. Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89232: Summit of
G Subis Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89422.
136 Gardens’ Bulletin Singapore 56 (2004)
*Appendicula sp.
Epiphyte. Disturbed forest among limestone rocks, light shade. Below W mouth of Great
Cave Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89489.
*Bulbophyllum cf. acutum J.J.Sm.
Epiphyte. Limestone forest, alluvial, medium shade. Gua Sibau, trail from Rumah Chang,
NE of Park Julaihi Abdullah, Kit Pearce et al. S 89320.
*Bulbophyllum minutulum Ridley
Epiphyte. On limestone. Niah Vermeulen LC 26546 (from Beaman et al. 2001).
*Bulbophyllum vaginatum (Lindl.) Rchb.f.
Epiphyte. G Subis, 50 m Vermeulen 117] (from Beaman et al. 2001).
Calanthe triplicata (Willemet) Ames
Terrestrial. G Subis J.A.R. Anderson S 31668; Niah, 300 m Burtt & Woods B 2031; Niah.
Synge, S. 577. (from Beaman et al. 2001).
*Calanthe cf. triplicata (Willemet) Ames
Lithophytic herb 80 cm tall. On limestone rocks in forest at base of limestone crags, light
shade, stony mineral soil Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89270.
*Calanthe sp.
Herb 50 cm tall. Limestone forest. G Subis Jemree Sablee et al. S 89050.
Claderia viridiflora Hook.f.
Terrestrial. Observed by path from Pengkalan Lobang to Great Caves.
*Cleisostoma or Pomatocalpa sp.
Herb creeping on rock. Exposed limestone rocks with humus soil, summit of G Subis
Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89414.
Coelogyne verrucosa S.E.C. Sierra
Epiphyte. Niah 10-300m Vermeulen LC 26555 (from Beaman et al. 2001).
Coelogyne pandurata Lindl. .
Epiphyte. G Subis Mohidin S 21682 (from Beaman et al. 2001).
Didymoplexiella borneensis © (Schltr.) Garay
Saprophyte. G Subis Sonny Tan & Wright S 27273 (from Beaman et al. 2001).
*Dossinia marmorata © E.Morren
Small terrestrial orchid. On moss on exposed limestone rocks and humus soil, also in shade
near base of limestone hill. Just below summit of G Subis Kit Pearce, Yahud Wat, Serukit
Dubod & Jeffri Johari S 89425; Niah J.A.R. Anderson S 16025.
*Eria (Sect. Cylindrolobus) aff. leptocarpa Hook.f.
Epiphyte. Disturbed forest among limestone rocks, light shade. pkI6w W mouth of Great
Cave Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89480.
Plants of Niah National Park, Borneo 137
*Eria sp.
Epiphyte. MDF, below limestone, moderate shade, loamy soil. Abundant. At base of limestone
crags Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89278.
*Liparis cf. gibbosa Finet
Epiphyte. Exposed limestone rocks and humus soil. Just below summit of G Subis Kit Pearce,
Yahud Wat, Serukit Dubod & Jeffri Johari S 89426.
*Liparis elegans Lindl.
Epiphytic herb c. 20 cm tall. Exposed limestone rocks with humus soil. Summit of G Subis
Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89424.
*Liparis cf. grandiflora Ridley
Epiphyte. Disturbed forest among limestone rocks, light shade. Below W mouth of Great
Cave Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89483.
Liparis wrayi Hook.f.
Terrestrial herb 20 cm tall. By path, moderate shade, wet clay soil. Along N Park boundary
from Tj Belipat Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89240.
Malaxis andersonii (Ridl.) Ames
Terrestrial. Niah Synge S 599 (from Beaman et al. 2001).
Plocoglottis borneensis Rid.
Terrestrial orchid in forest. Niah Ahmad 45. G Subis Mohidin S 21678 (from Beaman et al.
2001).
Plocoglottis cf. borneensis Ridl.
Terrestrial herb 40 cm tall. Disturbed swamp forest, in standing water, moderate shade, grey
clay with wet humus layer. Close to Sg Niah Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri
Johari § 89265.
*Podochilus cf. serpillifolius (Blume) Lindl.
Epiphyte. Exposed limestone rocks with humus soil, summit of G Subis Kit Pearce, Yahud
Wat, Serukit Dubod & Jeffri Johari S 89423..
Thecostele alata (Roxb.) C.S.P.Parish
Herbaceous epiphyte. MDF/kerangas forest, moderate shade, sandy loam soil. By Sg Subis,
Jalan Madu Trail Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89451.
OXALIDACEAE
*Oxalis corniculata L.
Lithophytic herb creeping on limestone rock. Limestone forest, moderate shade, beside plank
walk to Great Cave Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89475.
PANDANACEAE
*Freycinetia rigidifolia Hemsley
Shrub | m tall. On ‘mor’ layer on summit of sharp limestone hill. G Subis, Niah 130 m.
J.A.R. Anderson S 16426.
l 3 § Gardens’ Bulletin Singapore 56 (2004)
Pandanus basilocularis Martelli
Observed along plank walk to Painted Cave.
Pandanus calcinactus St John ex Stone
Observed just below the summit of G Subis.
PASSIFLORACEAE
Adenia macrophylla (Bl1.) Koord. var. smilacina (Hall.f.) deWilde
Woody climber 7 m tall, c. 1 cm diam. Disturbed emperan forest, by path in open area, clay
soil. Tj Belipat, along boundary from Sg Niah (50 m from river) Kit Pearce, Yahud Wat,
Serukit Dubod & Jeffri Johari S 89201.
PIPERACEAE
Piper caninum Blume
Semi-woody climber 1.5 m tall. Disturbed peat swamp forest, open area by trail, peat soil.
Along N Park boundary from Tj. Belipat Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri
Johari S 89221].
Piper muricatum Blume
Herb c. 30 cm tall. Well-developed secondary forest by stream, moderate shade, alluvial soil.
Close to Sg Niah Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89262.
Piper sarmentosum Roxb. ex Homem.
Sprawling herb c. 60 cm tall. Disturbed seasonal swamp forest, light shade, grey clay soil. Sg
Subis Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89292.
*Piper ?stylosum Miq.
Climber up to 1.5 m high on small tree. Foot of limestone forest, sandy soil. Ulu Sg Trusan
Jemree Sablee et al. § 89053.
*Piper aff. vestitum C.DC.
Erect herb 25—50 cm tall. Limestone forest, light shade, humus soil amongst limestone rocks,
also on limestone rocks in forest at base of limestone crags, light shade, stony mineral soil
and top of limestone summit Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89407;
At base of limestone crags Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89276.
Piper sp. 1
Herbaceous climber 1.2 m tall. Disturbed forest, grey clay under 25 cm peat, moderate shade.
Close to Sg Niah Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89255.
Piper sp. 2
Herbaceous climber, c. 2 m tall. Disturbed seasonal swamp forest, light shade, grey clay soil.
Sg Subis Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89293.
Piper sp. 3
Climber on tree. MDF. Sg Sekaloh Jemree Sablee et al. S 89028.
Piper sp. 4
Shrub 1.5 m tall. Secondary forest, moderate shade, loam soil. Close to Sg Niah Kit Pearce,
Yahud Wat, Serukit Dubod & Jeffri Johari S 89243.
Plants of Niah National Park, Borneo 139
POACEAE
Schizostachyum or Bambusa
Bamboo to 3 m tall, 3 cm diam. Shady lowland alluvial forest. Along plank walk to Niah
Caves. Julaihi Abdullah, Kit Pearce et al S 89303.
PODOCARPACEAE
*Podocarpus confertus deLaubenfels
Tree 10 m tall, 12 cmdbh. Exposed limestone rocks and humus soil, just below summit of
G Subis Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89428.
POLY PODIACEAE
Microsorium linguiforme (Mett.) Copel.
Epiphyte. Shady lowland alluvial forest. Along plank walk to Niah Caves Julaihi Abdullah,
Kit Pearce et al. S 89302.
*Paragramma longifolia (Blume) T. Moore
Epiphyte c. 40 cm tall. Exposed limestone rocks and humus soil, just below summit of G
Subis Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89431.
*Phymatosorus nigrescens (Blume) Pic.Serm.
Lithophytic herb 1 m tall: Limestone forest, moderate shade, on limestone rock. Beside
plank walk to Great Cave Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89473.
*Pyrrosia lanceolata (L.) Farw.
Epiphyte. Foot of limestone forest, sandy soil. Ulu Sg Trusan Jemree Sablee et al. S 89057.
RUBIACEAE
*? Timonius
Shrub | m tall. Limestone forest (disturbed, including by logging) on exposed rocks, shady
area, soil peaty humus between rocks. S Boundary of Park, E of Quarry Kit Pearce, Julaihi
Abdullah & Serukit Dubod S 89500.
Acranthera cf. involucrata Val.
Herb 30-150 cm tall, 1 cm diam. MDF, also secondary forest, moderate shade, loam soil.
Sg Sekaloh Jemree Sablee et al. S 89032; Close to Sg Niah Kit Pearce, Yahud Wat, Serukit
Dubod & Jeffri Johari S 89244.
Anthocephalus cadamba (Roxb.) Miq.
Vegetation verification plot specimen in seasonal swamp forest on gley soils.
Cephaelis stipulacea Blume
Treelet 1 miall, 0.8 cm diam. MDF, brown clay soil. Sekaloh Jemree Sablee et al. S 89009.
Geophila repens (L.) Johnston
Creeping herb. Seasonal swamp forest (disturbed), by path, light shade, clay soil.
Along Jalan Madu Trail Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89490.
Lasianthus constrictus Wight
Treelet 1 m tall. MDF, brown clay soil. Sekaloh Jemree Sablee et al. S 89015.
140 Gardens’ Bulletin Singapore 56 (2004)
Lasianthus stipularis Blume }
Treelet 2.5 m tall, 1.5 cm diam. Seasonal swamp forest (disturbed), light shade, clay soil.
Trail into park from Kuala Sg Subis Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari
S 89287.
*Mussaenda sp.
Herbaceous climber 3 m long. Exposed limestone rocks with humus soil, summit of G
Subis Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 8942].
*Ophiorrhiza sp.
Herb to 25 cm tall. Dry limestone forest, open, with litter. Limestone hill next to Quarry
Julaihi Abdullah, Kit Pearce et al. § 89315.
Pavetta indica L.
Treelet 2-4 m tall, < 1—-2.5 cm diam. Disturbed forest, grey clay under 25 cm peat, moderate
shade also disturbed, regenerating emperan forest, light shade, alluvial soil. Close to Sg
Niah Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89256; Kuala Sg Subis Kit
Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89281.
Psychotria cf. aurantiaca Blume
Shrub or treelet 70-100 cm tall. Emperan forest, dark brown clay soil also just beside
inundated area with standing water, clay soil, moderate shade. Jemree Sablee et al. S 89004;
Tj Belipat, along boundary 50 m from Sg Niah Kit Pearce, Yahud Wat, Serukit Dubod &
Jeffri Johari § 89205.
*Psychotria cf. obovata Wall.
Herbaceous climber to | m tall. Exposed limestone rocks and humus soil, just below summit
of G Subis. Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89427.
Saprosma arborea Blume
Treelet 4 cm diam. Disturbed secondary forest, moderate shade, clay soil. Along N Park
Boundary from Tj Belipat Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89213.
Tarenna aff. costata Val.
Treelet 1.5 m tall. MDF, brown clay soil. Sekaloh Jemree Sablee et al. S 89022.
Urophyllum hirsutum Wight
Tree 3 m tall or shrub 1.5 m tall, 1.5 cm diam. Disturbed MDF, light shade, grey clay soil
ith humus 5 cm deep (locally abundant), also emperan forest, dark brown clay soil. Close
(0 Sg Niah Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89249; Jemree Sablee et
al. S 89002.
RUTACEAE
*Micromelon minutum (G. Forst) Wight & Arn.
Tree c. 6 m tall, c. 5 cm diam. Disturbed forest among limestone rocks, light shade. Below
W mouth of Great Cave Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89484.
SAPINDACEAE
Allophyllus cobbe (L.) Raeuschel
Leaning treelet 4 m tall, 3 cm diam. Seasonal swamp forest dunce light shade, clay
soil. Trail into Park from Kuala Sg Subis Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri
Johari § 892835.
Plants of Niah National Park, Borneo 141
*Lepisanthes falcata (Rad\lk.) Leenh.
Treelet 7 m tall, 5 cm diam. Limestone forest among rocks on hillside, moderate shade,
loamy soil with humus. SW of Telaga Kumang Kit Pearce, Yahud Wat, Serukit Dubod &
Jeffri Johari S 89300.
*Lepisanthes tetraphylla (Vahl) Radlk.
Tree 2 m tall, 2 cm diam. In limestone forest at summit on exposed rocks, medium shade. S
boundary of Park, E of Quarry Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89498.
Nephelium sp.
Tree 7 m tall, 1.5 cm diam. MDF, brown clay soil. Sekaloh Jemree Sablee et al. S 89011.
Pometia pinnata J.R. Forst. & G.Forst.
Observed along riverbanks.
?Sapindaceae
Small tree 4 m tall, 6 cm diam. MDF Sg Sekaloh Jemree Sablee et al. S &9030.
SAURAUIACEAE
Saurauia sp.
Tree to 2 m tall, 4 cm. diam. Shady lowland alluvial forest. Along plank walk to Niah Caves
Julaihi Abdullah, Kit Pearce et al. S 89306.
STERCULIACEAE
Leptonychia caudata (Wall. ex G.Don) Burret
Small tree 3 m tall, 4 cm diam. MDF, brown clay soil. Sekaloh Jemree Sablee et al. S 89010.
Pterospermum subpeltatum C.B. Rob.
Vegetation verification plot specimen in seasonal swamp forest on gley soils.
Sterculia aff. coccinea Jack
Small tree. Seasonal swamp forest (disturbed), open area. Beside plank walk to Great Cave
Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89471.
Sterculia rubiginosa Vent.
Tree c. 11 m tall, 25 cm diam. Disturbed, regenerating emperan, light shade, alluvial soil.
Kuala Sg Subis Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89280.
Sterculia stipulata Korth.
Tree 4 m tall, c. 3 cm diam. Seasonal swamp forest (disturbed), light shade by stream.
Beside plank walk to Great Cave Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89466.
*Sterculia rubiginosa or stipulata
Treelet 6 m tall, 5 cm diam. Disturbed forest among limestone rocks, light shade. Below W
mouth of Great Cave Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89482.
?Sterculia
Small tree 7 m tall, 5 cm diam. MDF, Sg Sekaloh Jemree Sablee et al. S 89026.
142 Gardens’ Bulletin Singapore 56 (2004)
THELY PTERIDACEAE
Mesophlebion oligodictyon (Baker) Holltum
Niah C. Hose 1341.
TILIACEAE
Grewia aff. pearsonii (Merr.) Burret
Tree 2.5 m tall, 1.5 cm diam. MDF, below limestone, loamy soil. At base of limestone crags
Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89277.
Shoutenia glomerata King
Observed growing gregariously along Sg Subis along Jalan Madu Trail.
URTICACEAE
*Flatostema acuminata (Poir.) Brongn.
Creeping terrestrial herb c. 20 cm tall. Limestone forest on stones and humus on top of rock
in secondary seasonal swamp forest. G Subis Jemree Sablee et al. S 89047; Trail into park
from Kuala Sg Subis Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89291
Elatostema sp.
Herb 25 cm tall. Secondary forest, loam soil under 15 cm humus, moderate shade. Close to
Sg Niah Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89260.
*Oreocnide trinervis (Wedd.) Miq.
Small tree 4-5 m tall, 4-5 cm diam. Limestone forest, light shade, humus soil amongst
limestone rocks. Ulu Sg Sekaloh, near Quarry Jemree Sablee et al. S 89067; Top of limestone
summit Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89402.
?Pilea sp. 1
Sprawling herb c. 30 cm tall. Disturbed peat swamp forest, moderate shade, peat soil. S of
trail from Tj Belipat along N boundary of Niah N.P. Kit Pearce, Yahud Wat, Serukit Dubod
& Jeffri Johari § 89231.
?Pilea sp. 2
Semi-woody plant c. 20 cm high. Disturbed peat swamp forest, open area by trail, peat soil.
Along N Park boundary from Tj Belipat Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri
Johari § 89224.
*Urticaceae sp. |
Creeping herb. Foot of limestone hill, on rock. Great Cave Jemree Sablee et al. S 89063.
*Urticaceae sp. 2
Creeping herb. Forest canopy c. 20 m tall, light shade, humus soil among limestone NNT,
on loose soil below wall of limestone. Below summit of G Subis Kit Pearce, Yahud Wat,
Serukit Dubod & Jeffri Johari S 89439.
*Urticaceae sp. 3
Terrestrial herb. Limestone forest. G Subis Jemree Sablee et al. S 89044.
*Urticaceae sp. 4
Lithophytic herb. Limestone forest (disturbed, including by logging) on exposed rocks,
Plants of Niah National Park, Borneo 143
shady area, soil peaty humus between rocks. S boundary of park, E of Quarry Kit Pearce,
Julaihi Abdullah & Serukit Dubod S 89492.
*Urticaceae sp. 5
Herb 35 cm tall. MDF, Sg Sekaloh Jemree Sablee et al. S &9025.
*Urticaceae sp. 6 .
Herb c. 30 cm tall. On limestone rocks, light shade, almost no soil. Near Sg Subis Kit
Pearce, Julaihi Abdullah & Serukit Dubod S 89462.
VERBENACEAE
*Clerodendrum sp.
Semi-woody plant 25 cm tall. On limestone rocks in pockets of soil between rocks, medium
shade. Beside plank walk to Great Cave Kit Pearce, Julaihi Abdullah & Serukit Dubod S
89479.
*Verbenaceae sp.
Woody herb 30 cm tall. Foot of limestone hill. Great Cave Jemree Sablee et al. S 89064.
VIOLACEAE
*Rinorea bengalensis (Wall.) O.K.
Treelet 2.5 m tall, 2 cm diam. Limestone forest among rocks on hillside, moderate shade,
loamy soil with humus. SW of Telaga Kumang Kit Pearce, Yahud Wat, Serukit Dubod &
Jeffri Johari S 89296.
*Rinorea horneri (Korth.) O.K.
Small tree 1.5—5 m tall, 3-8 cm diam. Limestone forest. G Subis Jemree Sablee et al. S
89045; Ulu Sg Sekaloh, near Quarry Jemree Sablee et al. S 89068.
VISCACEAE
Ginalloa falcata Dans.
Hemi-parasitic shrub on branch. Seasonal swamp forest (disturbed), open area. Beside plank
walk to Great Cave Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89468.
VITACEAE
*?Cayratia sp. 1
Herbaceous creeping scrambler 2 m long. Exposed limestone rocks with humus soil, summit
of G Subis Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89418.
*?Cayratia sp. 2
Semi-woody climber c. 10 m tall, 1 cm diam. Limestone forest, light and moderate shade,
humus soil amongst and climbing over limestone rocks. Top of limestone summit of G
Subis Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89406; Beside plank walk to
Great Cave Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89476.
*?Cayratia sp. 3
Creeping herb. Limestone forest. Moderate shade on limestone rock. Beside plank walk to
Great Cave Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89474.
144 Gardens’ Bulletin Singapore 56 (2004)
Cissus cf. angustata Ridley
Semi-woody climber c. 5 m tall. MDF, light shade, loam soil. Across river from Park HQ Kit
Pearce, Julaihi Abdullah & Serukit Dubod S 89443.
Pterisanthes polita (Miq.) Lawson
Herbaceous climber 1.5 m long. MDF, light shade, loam soil. NE of Kuala Sg Sekaloh Kit
Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89266.
Tetrastigma cf. diepenhorstii (Miq.) Latiff
Woody climber c. 12 m long, c. 7 cm diam at base. Disturbed forest, by path, moderate
shade. Tj Belipat, along boundary from Sg Niah (100 m from river) Kit Pearce, Yahud Wat,
Serukit Dubod & Jeffri Johari S 89202.
VITTARIACEAE
*Antrophyum parvulum Blume
Vegetation verification plot specimen near summit of Bukit Kasut.
Monogramma paradoxa Bedd.
Collected by H. Schneider (pers. comm.).
ZINGIBERACEAE
*Alpinia ligulata K.Schum.
Clustering herb 2.5 m tall. Foot of limestone hill. Great Cave Jemree Sablee et al. S 89061.
Amomum borneense K.Schum.
Epiphytic herb. Locally abundant. MDF, light shade. Across river from Park HQ Kit Pearce,
Julaihi Abdullah & Serukit Dubod S 8944].
*Boesenbergia sp.
Creeping herb. At base of limestone rocks, medium shade, damp. Along Jalan Madu Trail
Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89491.
This species belongs to the B. aurantiaca/belalongensis/ornata group.
*Boesenbergia sp. nov. (A. Poulsen, pers. comm.)
Yellow-flowered herb collected by A. Poulsen.
Globba atrosanguinea Teijsmann & Binnend.
Herb 60 cm tall. Disturbed forest, grey clay under 25 cm peat, moderate shade. Close to Sg
Niah Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89258.
*Globba ?propinqua
Herb 50 cm tall. Limestone forest, medium shade, peaty soil at base of limestone _—
Near Sg Subis. Kit Pearce, Julaihi Abdullah & Serukit Dubod S 89452.
Globba tricolor Ridley var. gibbsiae (Ridley) R.M.Smith
Solitary herb 40 cm tall. MDF, brown clay soil. Sekaloh Jemree Sablee et al. S 89020.
Zingiber cf. albiflorum (Ridley) R.M.Smith
Clustering herb 1.5 m tall. Disturbed peat swamp forest, moderate shade, peat soil. S of trail
from Tj Belipat along N Park boundary. Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri
Johari S 89230.
Plants of Niah National Park, Borneo 145
Zingiber cf. martinii R.M.Smith
Erect, clustering herb 2.4 m tall. Disturbed, regenerating emperan forest, light shade, alluvial
soil. Kuala Sg Subis Kit Pearce, Yahud Wat, Serukit Dubod & Jeffri Johari S 89282.
*Family unknown
Climber on tree. Limestone forest, sandy soil. Ulu Sg Trusan Jemree Sablee et al. S 89059.
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Gardens’ Bulletin Singapore 56 (2004) 147-152. 147
A New Species of, and Reinstatements in, Octamyrtus
(Myrtaceae)
LYN A. CRAVEN
Australian National Herbarium, CPBR, CSIRO Plant Industry,
GPO Box 1600, Canberra, ACT 2601, Australia
AND
SITI SUNARTI
Herbarium Bogoriense, Bidang Botani, PPB-LIPI,
Jalan Raya Juanda 22, Bogor 16122, Indonesia
Abstract
The taxonomy of Octamyrtus Diels is reviewed and six species are recognised. Octamyrtus
halmaherensis Craven & Sunarti is newly described and O. arfakensis Kaneh. & Hatus. ex
C.T. White and O. glomerata Kaneh. & Hatus. ex C.T. White are reinstated. Neotypes are
designated for O. behrmannii Diels and O. insignis Diels. A key to the species is provided.
Introduction
Due to its largish, multipetalled flowers that commonly are red or pink,
Octamyrtus Diels is an intriguing genus of Papuasian Myrtaceae. The syndrome
of floral features, i.e. long, more or less tubular, often brightly coloured flowers,
suggests that its several species may be pollinated by birds. The genus was
revised in 1978 by Scott who recognised three species, O. behrmannii Diels,
O. insignis Diels and O. pleiopetala Diels, with the last-named species
containing two varieties, O. pleiopetala var. arfakensis (Kaneh. & Hatus. ex
C.T. White) A.J. Scott and the typical morph (Scott 1978a).
The first author’s long-standing interest in this attractive genus was re-
kindled by an inquiry from Marcel Polak, Leiden, as to the identity of collections
he had made on the Vogelkop, Indonesian New Guinea, in 1995 and 1996.
Polak’s specimens had extremely large leaves with the lamina decurrent to the
petiole and yellowish-coloured flowers; they keyed to O. insignis in Scott’s
key (1978a). Upon comparison with collections lodged in herb. CANB, and
those cited by Scott (1978a) in his revision as being’O. insignis, the Polak
specimens proved to be significantly different. The CANB specimens of O.
insignis had distinctly petiolate, non-decurrent leaves and pink to red flowers.
During collaborative investigations of Octamyrtus specimens in BO, the first
author noted that the Polak specimens had much in common with an isotype
148 Gardens’ Bulletin Singapore 56 (2004)
specimen of O. glomerata Kaneh. & Hatus. ex C.T. White, a species placed in
synonymy with O. insignis by Scott (1978a).
The investigations were continued during a study visit by the second
author to CANB in 2003 with the conclusion that the Polak specimens were
conspecific with the type collection of O. glomerata and that this species was
distinct from O. insignis. It was further concluded that Scott had inappropriately
combined O. arfakensis Kaneh. & Hatus. ex C.T. White with O. pleiopetala
although he did give the former taxon status at the rank of variety. An anomalous
collection from Maluku (de Vogel 3140), differs from all other collections of
the genus that we have seen in possessing sessile, cordate leaves and we
conclude that this represents a new species, described below as O.
halmaherensis Craven & Sunarti.
Octamyrtus has much in common with Rhodomyrtus (DC.) Rchb., but
differs from this genus in having a greater number of petals, 6, 8 or 12 as
against 4 or 5 in Rhodomyrtus (Scott 1978a). The petals in Octamyrtus are
often red or pink and may be up to c. 6cm long. In most species of Malesian
rainforest Myrtaceae, the petals are relatively insignificant with the stamens
providing the primary visual signal for pollinators; in this respect Octamyrtus
appears to be unique. Nonetheless, in features of the ovules and placentation,
and seeds and fruit morphology the species of Octamyrtus are so similar to
Papuasian Rhodomyrtus that they may in future be found to be better placed in
that genus.
Taxonomy
Octamyrtus Diels
Octamyrtus Diels, Bot. Jahrb. Syst. 57 (1922) 373. —Type: Octamyrtus insignis
Diels (lecto, vide McVaugh, Taxon 5 (1956) 144).
1. Octamyrtus arfakensis Kaneh. & Hatus. ex C.T. White
Octamyrtus arfakensis Kaneh. & Hatus. ex C.T. White, J. Arnold Arb. 32
(1951) 144. Octamyrtus pleiopetala var. arfakensis (Kaneh. & Hatus.
ex C.T. White) A.J. Scott, Kew Bull. 33 (1978) 305. —Type: Indonesian
New Guinea, Arfak Mountains, Lake Gita, 1900 m, Kanehira & Hatusima
14028 (lecto A, not seen (image seen); isolecto BO, BRI not seen).
Notes: Although White (1951) cited the Kanehira and Hatusima collection as
being the type when he described O. arfakensis, he did not. designate a holotype.
Scott (1978a) designated the specimen lodged in A as holotype and thus
effectively has selected the lectotype. The BRI material cited by Scott as an
isotype is fragmentary, consisting of a single leaf only (A.R. Bean, pers. comm.)
and Scott’s choice of the A specimen as lectotype is appropriate.
A new species of, and reinstatements in, Octamyrtus (Myrtaceae) 149
2. Octamyrtus pleiopetala Diels
Octamyrtus pleiopetala Diels, Bot. Jahrb. 57 (1922) 373. Eugenia pleiopetala
F. Muell., Descr. Notes Papuan Pl. 1 (1877) 106 sub Myrtella hirsutula,
nom. inval. (nom. prov.). —Type: Indonesia, Moluccas, Aru Islands, Beccari
s.n. (2lecto MEL, not seen; isolecto K, L, not seen (images seen)).
Octamyrtus lanceolata C.T. White, J. Arnold Arb. 32 (1951) 145. Type: Papua
New Guinea, Western Province, Brass 770] (lecto A, not seen (image seen);
isolecto BO; BM, BRI, K, L, not seen (image seen from L)).
Notes: Scott (1978a) followed Diels (1922) in treating the name O. pleiopetala
as being acombination resulting from the transfer of Mueller’s (1877) Eugenia
pleiopetala to Octamyrtus. Scott, however, appears to have overlooked the
long-standing provision of the various editions of the International Code of
Botanical Nomenclature, including the most recent version (the Saint Louis
Code), that pertains to names not accepted by their author(s) (Art. 34.1; Greuter
et al. 2000). Mueller’s (1877) name is invalidly published as he was tentative
as to bestowing the epithet pleiopetala upon it (i.e. “to which the name E.
pleiopetala might be given’, p. 106); that he was also unsure as to which
genus the new plant belonged is immaterial. Diels (1922) did not validate
Mueller’s name in Eugenia as he was recognising the species as belonging to
his new genus, Octamyrtus, and treated the species accordingly. Thus Diels
apparently has provided the first validly published name for the plant and, as
he was treating it under a different genus to Mueller, the name Octamyrtus
pleiopetala should not be treated as a new combination, even though Diels’
name is based on the species concept and descriptive text Mueller derived
from Beccari’s specimens and is to be typified by those specimens.
Scott (1978a) cited specimens of O. pleiopetala in K and L as isotypes,
presumably regarding the material assumed to be at MEL as the primary type,
in this case lectotype as Mueller did not designate a holotype. Type material
of O. pleiopetala has not been located in MEL (J.H. Ross, pers. comm.) and
may no longer be extant in which case a lectotype will need to be selected
from the other sheets of Beccari’s collection that are available.
As with O. arfakensis, White (1951) did not designate a holotype when
describing O. lanceolata and Scott’s (1978a) designation of the material at A
as holotype has effected lectotypification. The BRI isolectotype material fide
A.R. Bean (pers. comm.) consists of an adequate flowering specimen.
3. Octamyrtus behrmannii Diels
Octamyrtus behrmannii Diels, Bot. Jahrb. 57 (1922) 376. —Neotype (here
designated): Papua New Guinea, Morobe Province, Aseki Patrol Area, near
Haumga, 6 April 1966, Craven & Schodde 1195 (neo CANB; isoneo A,
BRI, CHR, K, L, LAE, TNS).
150 Gardens’ Bulletin Singapore 56 (2004)
Notes: With the loss of the type designated by Diels (1922), i.e. Behrmann in
Ledermann 6969, during the Second World War, a neotype should be selected.
Scott (1978a) apparently did not locate any of the original material and a neotype
that conforms with Diels’ circumscription has been designated above.
4. Octamyrtus insignis Diels
Octamyrtus insignis Diels, Bot. Jarhb. 57 (1922) 374. —Neotype (here
designated): Papua New Guinea, Morobe Province, foothills NW of the
Waria River, alluvial terraces along Pao streambed by Yai Village, 4 June
1999, Takeuchi et al. 13126 (neo CANB; isoneo LAE not seen).
Notes: With the loss of the type designated by Diels (1922), 1.e. Schlechter
17428, during the Second World War, a neotype should be selected. Scott
(1978a) apparently did not locate any of the original material and a neotype
that conforms with Diels’s circumscription has been designated above.
5. Octamyrtus halmaherensis Craven & Sunarti, sp. nov.
Octamyrtus halmaherensis Craven & Sunarti, a congeneribus foliis sessilibus
et lamina cordata differt. -—Typus: Indonesia, Maluku, Halmahera, Ekor, G.
Panjang, 27 September 1974, de Vogel 3140 (holo BO; iso CANB, L not
seen).
Tree 10 m tall, clear bole to 5 m, dbh 10 cm, rather gnarled, without buttresses
or knots; bark red-brown, not fissured, strongly peeling. Branchlets finely
pubescent, 5—7 mm wide. Leaves sessile, obovate, finely pubescent with the
hairs weathering away on the adaxial surface, with oil dots (the white material
present on the only specimens seen apparently is an artefact of preservation),
22—31.4 x 9.2-12.5 cm; base cordate; apex acute to short acuminate; midrib
strongly prominent and rounded abaxially and slightly impressed to flat
adaxially; primary veins strongly prominent abaxially and slightly impressed
adaxially, arching, confluent at their apex with the next vein and an intramarginal
vein not developed; secondary veins not or poorly developed; tertiary veins
prominent abaxially, linking the primary veins in a more or less ladder-like
pattern. Flowers 1-2 on a distal-axillary spicate inflorescence, 3 x 1 cm,
inflorescence axis up to c. | cm long, the axis bracts, bracteoles, hypanthium
and sepals densely and finely pubescent; bracteoles persistent in fruit, narrowly
ovate, 10 mm long, pedicel 3-4.5 mm long. Hypanthium funnel-shaped, very
slightly stipitate or estipitate, 5.5—7 x 5—5.5 mm. Sepals 4, green, persistent in
fruit, ovate to subtriangular, 10.5—13.5 x 6—6.5 mm. Petals 8, bright red, blade
glabrous, margin pubescent, narrowly elliptic to elliptic, 20-28 mm long, oil
glands present. Stamens 35—37 mm long, filaments violet, anthers yellow, 2.4—
2.8 mm long. Style at least 30 mm long (fully mature style not seen), stigma
A new species of, and reinstatements in, Octamyrtus (Myrtaceae) 15 1
green, peltate. Ovary 4-locular, with 2 rows of ovules in each locule. Immature
fruit ellipsoid, 18 mm long, seeds not seen but horizontal and vertical septa
present.
Habitat & ecology: Found in rather dense primary forest 20 m tall, with little
undergrowth, on arather steep hill side on deep clayey soil, 40 m alt, gregarious.
Distribution: Known only from the type collection.
Notes: Octamyrtus halmaherensis is readily distinguished from all other species
of the genus in its leaves, which are sessile and cordate. Although known from
limited material only, its diagnostic features are unique in the genus.
6. Octamyrtus glomerata Kaneh. & Hatus. ex C.T. White
Octamyrtus glomerata Kaneh. & Hatus. ex C.T. White, J. Arnold Arb. 32 (1951)
145. —Type: Indonesian New Guinea, Kanehira & Hatusima 14126 (lecto
A, not seen (image seen); isolecto BO, BRI fragment not seen).
Notes: As with the other Octamyrtus species described by White (1951), a
holotype was not designated. Scott’s (1978a) designation of the A specimen
as holotype effectively has nominated the lectotype. The BRI material cited
by Scott as an isotype is fragmentary, consisting of some pieces of material in
a packet (A.R. Bean, pers. comm.) and Scott’s choice of the A specimen as
lectotype is appropriate.
Key to the Species of Octamyrtus
la. Leaves distinctly petiolate, the lamina not decurrent
2a. Sepals up to 10 mm long
3a. Leaves with a distinct intramarginal vein; petals white, lemon-
white, greenish white or light yellow ............. 1. O. arfakensis
3b. Leaves without an intramarginal vein; petals pink or red, rarely
yellowish white
4a. Flowers inserted in the distal leaf axils .....2.0. pleiopetala
4b. Flowers usually inserted on the trunk or on branchlets below
the leaves, rarely in leaf axils ............... 3. O. behrmannii
Zoe Sepals 1292. more mim, lomg).<).yaieccti ho bednalas de. 4. O. insignis
1b. Leaves sessile or the lamina strongly decurrent to the petiole
Jan caves sessnc the lanimacordate 22 ao. 5. OQ. halmaherensis
5b. Leaf lamina shortly petiolate, the lamina strongly decurrent to the
I i ah ae te Rit ag Ai lbenno de 6. O. glomerata
152 Gardens’ Bulletin Singapore 56 (2004)
Excluded species
Octamyrtus elegans (Bl.) Steen., Blumea 19 (1971) 146; Macropsidium elegans
BI., Mus. Bot. Lugd. Bat. 1 (1849) 85; Psidium elegans (B1.) Migq., Fl. Ind.
Bat. 1, 1 (1855); Rhodomyrtus elegans (Bl.) A.J. Scott, Kew Bull. 33 (1978)
320. —Type: Indonesia, Maluku, Halmahera, in forests near Singanoli, Gilolo,
July 1861, Forsten s.n. (holo L, not seen (image seen)).
Scott (1978a) considered this species to be conspecific with Rhodomyrtus
calophlebia C.T. White and in a later paper (Scott 1978b) published the
combination Rhodomyrtus elegans (Bl.) A.J. Scott.
Acknowledgements
The Directors and/or Curators of the following herbaria are thanked for the
opportunity to study specimens in their care: BO, CANB and L. Particular
thanks are due to Marcel Polak (L), for sending some stimulating materials
from the Vogelkop Peninsula to the first author; these specimens rekindled his
interest in this fascinating group, culminating in the present paper. Emily
Wood and Walter Kittredge (A) and Eve Lucas (K) are thanked for making
available digital images of Octamyrtus types. Tony Bean (BRI) and Jim Ross
(MEL) kindly reported upon the status of type materials at their respective
institutions. Siti Sunarti was the recipient of a CSIRO-LIPI Award that made
possible her study visit to Canberra.
References
Diels, L. 1922. Die Myrtaceen von Papuasien. Botanische Jarhbiicher fiir Systematik,
Pflanzengeschichte und Pflanzengeographie 57: 356-426.
Greuter, W., J. McNeill, FR. Barrie, H.M. Burdet, V. Demoulin, T.S. Filgueiras, D. Nicolson,
P.C. Silva, J.E. Skog, P. Trehane, J.N. Turland and P.M. Hawksworth. 2000. /nternational
code of botanical nomenclature (Saint Louis Code) adopted by the sixteenth International
Botanical Congress, St Louis, Missouri, July-August 1999. Koeltz, Konigstein. (Regnum
Vegetabile 138). :
Mueller, F. 1877. Myrtella hirsutula. In, Descriptive notes on Papuan plants. 106—107. George
Skinner, Melbourne.
Scott, A.J. 1978a. A revision of Octamyrtus (Myrtaceae). Kew Bulletin 33: 303-309.
Scott, A.J. 1978b. A revision of Rhodomyrtus (Myrtaceae). Kew Bulletin 33: 311-329.
White, C.T. 1951. Some noteworthy Myrtaceae from the Moluccas, New Guinea, and the
Solomon Islands. Journal of the Arnold Arboretum 32: 139-149.
Gardens’ Bulletin Singapore 56 (2004) 153-159. 153
A new species of Amorphophallus (Araceae: Thomsonieae)
from Sarawak, Borneo
I.B. IPOR', C.S. TAWAN! AND P.C. BOYCE?
'Faculty of Resource Science & Technology, Universiti Malaysia Sarawak,
94300 Kota Samarahan, Sarawak
? Malesiana Tropicals, Suite 9-04, Tun Jugah Tower, No. 18,
Jalan Tunku Abdul Rahman
93100 Kuching, Sarawak, Malaysia
Abstract
Amorphophallus julaihii Ipor, Tawan & P.C. Boyce a new species from forested limestone in
Mulu National Park, Sarawak, Borneo is described and illustrated.
Introduction
Recognition of Amorphophallus julaihii as a new species takes to fifteen the
number of indigenous species of Amorphophallus recorded from Borneo
(Bogner 1989; Hetterscheid 1994, 2001). Including A. julaihii there are eight
species in Sarawak, viz: A. angulatus Hett. & A.Vogel, A. brachyphyllus Hett.,
A. eburneus Bogner, A. hewittii Alderw., A. hottae Bogner & Hett., A.
infundibuliformis Hett., A. Dearden & A. Vogel, A. pendulus Bogner & Mayo.
Five species have been recorded from Sabah: A. hottae, A. lambii Mayo &
Widjaja, A. rugosus Hett. & A.L. Lamb., A. tinekeae Hett. & A. Vogel and A.
venustus Hett., A. Hay & J. Mood. Eight species are recorded from Kalimantan:
A. borneensis (Engl.) Engl. & Gehrm., A. costatus Hett., A. hewittii, A.
infundibuliformis, A. lambii, A. linguiformis Hett., A. pendulus and A. prainii
Hook.f. (the last perhaps based on a mis-labelled specimen; A. prainii is
otherwise known only from Peninsular Malaysia and Sumatera). With the
exception of A. prainii all Bornean Amorphophallus are endemic to Borneo.
The most remarkable aspect of these data is that 13 of these 15 species have
been described within the past 25 years. This extraordinary increase in
recognized species is being repeated throughout the range of the genus such
that Amorphophallus now numbers over 200 species, of which in excess of
one third are novel taxa described since 1980. Recent and on-going fieldwork
indicates that there are more novel Bornean Amorphophallus awaiting
description.
154 Gardens’ Bulletin Singapore 56 (2004)
Ecology of Amorphophallus in Sarawak
Two of the authors (1.1. & P.C.B.) have been observing Amorphophallus
populations in North Borneo for many years and based on these observed data
the following summary of Amorphophallus ecology in Sarawak may be drawn.
Five of the eight Sarawak Amorphophallus species occur in limestone
forest with three, A. brachyphyllus, A. eburneus and A. julathii, seemingly
restricted to this habitat. Amorphophallus brachyphyllus and A. eburneus occur
sporadically and allopatrically on both the Bau and Padawan limestones. There
are also records of A. brachyphyllus from the Mulu and Niah limestones,
although these records have yet to be verified and it is quite possible that the
Mulu and Niah plants represent one or more additional undescribed vicariant
taxa in the species-rich Manta Group (Hetterscheid, in prep.). The single record
from Mulu for the otherwise Sabahan A. hottae also requires verification not
least because in Sabah A. hottae is never associated with limestone.
Amorphophallus hewittii as currently circumscribed is frequently found
in association with limestone, occurring commonly on both the Bau and
Padawan limestones where the flowering of large specimens occasionally
receives coverage in the local press. However, A. hewittii is not restricted to
calcareous rocks, and is also found on the hard sandstones of the Penrissen
Range and occurs in several widely scattered, mostly sandstone, locations
throughout Sarawak. It should be noted that this apparent ecological diversity
may be an artifact of imperfect taxonomy. The large size of this plant, both
florally and vegetatively, makes it an unpopular subject for herbarium collection
and our knowledge of its morphology is based on a decidedly meagre collection
of mediocre specimens.
All other Sarawak species appear to be sandstone or shale. associated.
Amorphophallus pendulus, described from Gn. Matang (Bogner et al. 1985),
is widespread and frequently locally common on moist sandstones at least as
far east as Kapit (Belaga) (Boyce, pers. obs.). Amorphophallus
infundibuliformis is widespread but scattered and seldom locally abundant in
Kuching and Sri Aman Divisions, with collections known from wet but well-
drained sandstone sites between 60-870 m asl in Lundu, Padawan, Bau & Ulu
Batang Ai. Amorphophallus angulatus was described from the sandstones of
Gunung. Selantik (Sri Aman), is also recorded from Gunung. Ampungan
(Samarahan) and has recently been discovered at Nanga Gaat (Kapit) where it
occurs on hard shales exposed by stream action (Boyce, pers. obs.).
Relationships of Amorphophallus julaihu
Amorphophallus julaihii fits uncontroversially into Hetterscheid’s “Manta
Group’ (Hetterscheid, in prep), into which also belong all the other Sarawak
A new species of Amorphophallus (Araceae: Thomsonieae) from Sarawak, Borneo 15
(|=
Figure 1. Amorphophallus julaihii \por, Tawan & P.C. Boyce.A. seedling, B.
leaflet of seedling, C. mature plant, D. tuber with adventitious roots, E. tuber
from inflorescence, F. leaflets of mature plant, G. inflorescence with cataphylls,
H. LS. of the inflorescence, I. male and female zones, J. stamens, K. pistils, L
infrutescence with berries, M. berry and in L.S..
A,B, C & D, E, FE G, H, I, J, K from CST 2527, L & M from CST 2530 —
drawings from fresh samples.
156 Gardens’ Bulletin Singapore 56 (2004)
species to which A. julaihii is most similar. The Manta Group is defined as:
Small to medium-sized herbs with tubers depressed-globose or subglobose
that are not offsetting; petioles sometimes forming an intercalary bulbil at the
junction with the lamina (A. angulatus). Seedling leaves often flushed with
dark red or lilac-red (A. angulatus, A. pendulus, - Boyce, pers. obs.) Spathe
linguiform, narrow or broad. Spadix shorter than or only slightly longer than
spathe. Male flowers often longitudinally elongate and fused into rows.
Amorphophallus julathii \por, Tawan et P.C. Boyce sp. nov. Ab omnibus
speciebus in habitu calcicola lithophytica Borneensibus borealis spatha parva
in toto atropurpureis differt. Typus. Sarawak, Miri Division, Mulu National
Park, forest on limestone, C.S. Tawan & I.B. Ipor CST 2527, 14 April 2004
(Holotypus: HUMS (Herbarium Universiti Malaysia Sarawak) inflorescence
and tubers in spirit; vegetative parts (petiole with leaflets, seedlings) - herbarium
specimens.)
Small to medium-sized herb, 50—100 cm tall. Tuber depressed sub-cylindrical
to globose, with irregular raised areas, up to 5.8 cm diam., 6.0 cm high, surface
dull brown greyish, inner part fleshy, whitish. Adventitious root scars present
on top portion of tuber at flowering stage, new roots developing during
vegetative stage. Petiole up to 44 cm long, c. 19 mm diameter at base, turgid,
cylindrical, smooth, bright green, enveloped basally by dried decaying brownish
cataphylls; lamina highly dissected, rachises naked, narrowly channelled,
yellowish green; leaflets elliptic-lanceolate, 11.0—-13.0 x 4.0-+4.4 cm, some
petiolulate, petiolule 1.5—3.0 cm long, slightly channelled adaxially, ultimate
leaflet sessile, leaflets asymmetrical, apex apiculate 1.5—2.0 cm long; base
unequal; margin slightly wavy and sparsely fine-toothed; adaxial surface bright
green, thin slightly leathery, abaxial surface pale green; 6 —16 pairs of secondary
veins with intermediate veins per leaflet, these adaxially channelled and
adaxially raised; venation forming distinct submarginal veins; tertiary veins
reticulate; lamina texture leathery when fresh, chartaceous when dry.
Inflorescence solitary, flowering without foliage leaves; peduncle cylindrical,
up to 17.5 cm long, 8-9 mm diam. at base, yellowish to creamy. Cataphylls 6;
first 1-4 ovate to linear, 3—13 cm x 1.5—2.0 cm, brownish to dark brown, thin,
soon withering and decaying; the next 5—6 linear-oblong, 15—21 x 2.4—3.5 cm,
light purplish to whitish and slightly purplish stained. Spathe elongate-oblong,
limb erect at anthesis, later recurved and twisted towards the base, up to 12 cm
long, width at base 4.0 cm, at middle 4.0 cm, at apex 4.5 cm, lower spathe
strongly convolute and forming a short tube 3.5-4.0 cm long; inner surface of
upper spathe limb ribbed with distinct venation, purplish, lower part rough
and verrucose, deep purple; outer surface of upper spathe limb with distinct
longitudinal venation, margin thinly undulate, apex dentate; upper part of spathe
A new species of Amorphophallus (Araceae: Thomsonieae) from Sarawak, Borneo 57
a. tuber, 6. inflorescence with tuber, Foliage (Seale bar — 6 cm)
C. leaflet (Scale bars — 3 cm)
ayia *
ro
Inflorescence (scale bars — 6 c
F, Re
m) Spathe (Scale bars — 4 cm)
ae : a a=
aS fi
5 i = &
' s
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e -
4
oO i;
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. .
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a . _
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a. Spathe and spadix, b. Infrutescence. Male and female zones of the spadix
(Scale bars-2 cm) (Scale bar—2 cm)
Figure 2. Amorphophallus julaihii Ipor, Tawan & P.C. Boyce
158 Gardens’ Bulletin Singapore 56 (2004)
limb rich purple-scarlet, middle part and margins darker. Spadix exceeding
spathe, 18.5—19.0 cm, short stipe, 1.0-1.5 cm long. Appendix up to 14 cm
long, cylindrical, spongy, purplish, 9—9.2 mm diam. at base; 7.5 mm diam. at
middle, 2 mm diam. at apex, surface verrucose with longitudinal lines, appendix
producing an unpleasant odour similar to that of rotten fish. Flowers unisexual;
male zone cylindrical, up to 3 cm long, 9 mm diam. at middle, whitish to
creamy; stamens connate and short, c. 1 mm long, c. 0.5 mm broad across,
fused with adjacent stamens, filaments c. 5 mm, truncate, pores apical, rounded
or variously elongate. yellowish or creamy, pollen brownish. Female zone
cylindrical, separated by irregular zone of sterile stamens from the male zone,
1.5—2 cm long, 10 mm diameter; pistils in irregularly rows, rather lax,
sometimes almost in V-shaped rows; ovaries ovoid-subglobose, 2.5—3.0 mm
in long, 1.8—2.0 mm broad at base, dark purple, unilocular rarely bilocular;
stigma sessile, two to three lobed or sometimes irregularly lobed,
Infructescence with up to 35 berries, pedunculate up to 22 cm long, 6 mm
diam. at base, 9 mm diam. at apex, with blackish-dark brown V-shaped scar
from the marcescent spathe, basally with remains of the cataphylls. Berries
when ripe deep orange, ellipsoid 15—16 mm x 8-10 mm, apex rounded, with
blackish stigma remnants, one seeded. Seeds ellipsoid, 12-14 mm x 7-8.2
mm wide, testa smooth, thin, yellowish green, seed copiously starchy, embryo
small.
Distribution: Endemic in Sarawak, so far recorded only from Mulu National
Park, Miri Division.
Ecology: Limestone forest, growing in shady areas, in humus-filled fissures
and holes in limestone. Flowering recorded in April.
Notes: Amorphophallus julathii is most similar to A. angulatus, A.
brachyphyllus and A. eburneus, (all Sarawak) and A. costatus (Kalimantan).
From A. angulatus (with which A. julaihii shares a purple spathe), it differs by
the spadix appendix exceeding the spathe and by the smaller, not diamond-
shaped male flowers. From A. eburneus and A. brachyphyllus (also both
restricted to limestone), A. julaihii is immediately separable by the purple ribbed
spathe. The resemblance of A. julaihii to A. costatus is in having an erect
elongate triangular spathe with the base strongly convolute forming a narrow
tube. In both species, the spadix exceeds the spathe but in A. julaihii the spadix
is shortly stipitate while it is sessile in A. costatus; the male zones in both
species are similar - cylindrical, flowers slightly distinct, irregularly arranged
or in longitudinal oblique, interrupted rows. The matt-purple spathe of A. julaihii
readily separates it from A. costatus in which the spathe interior is glossy
maroon. To date A. costatus is recorded only from southern Kalimantan.
A new species of Amorphophallus (Araceae: Thomsonieae) from Sarawak, Borneo 159
Etymology: This species is named after Mr. Julaihi Abdullah, Deputy Research
Manager of Sarawak Forestry Corporation, who first showed the first two
authors this species in the Mulu National Park, Sarawak. He was formerly
Botanist of the Sarawak Forest Department, Kuching.
Other specimens seen: Type locality, inflorescence, C.S. Tawan & I.B. Ipor
CST 2528, 14 April 2004 (HUMS); Type locality, inflorescence, C.S. Tawan
& I.B. Ipor CST 2529, 14 April 2004 (SAR); Type locality, infructescence in
spirit C.S. Tawan & I.B. Ipor CST 2530, 14 April 2004 (HUMS); Sarawak,
Miri Division, Mulu National Park, unspecified locality, J.Brodie AM-39, 12
May 1999 (SAR).
Acknowledgements
The first two authors thank Universiti Malaysia Sarawak for the grant to
undertake this project. All authors would also like to thank the Director of
Sarawak Forest Department for permission to collect the specimens, and also
to Julaihi Abdullah, Sulaiman Jamhari, Malcom Demies, Meekiong Kalu, James
Abai, Mohd. Rizan Abdullah, Hidir Marzuki and Sekudan Tedong for their
help. Our appreciation also goes to Joseph Pao for the botanical illustrations.
References
Bogner, J. 1989. Anew Amorphophallus (Araceae) from Sarawak. Willdenowia 18: 441-443.
Bogner, J., S. Mayo and M. Sivadasan, M. 1985. New species and changing concepts in
Amorphophallus. Aroideana 8: 14—25.
Hetterscheid, W.L.A. 1994. Notes on the genus Amorphophallus (Araceae) — 2. New species
from tropical Asia. Blumea 39: 237-281.
Hetterscheid, W.L.A. 1994. Notes on the genus Amorphophallus (Araceae) — 11. New and
obsolete species from East Malaysia and continental Southeast Asia.—-Blumea 46: 253-
2e2..
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Gardens’ Bulletin Singapore 56 (2004) 161-166. 161
A New Species of Stachyphrynium (Marantaceae) from
Borneo
A.D. POULSEN
Royal Botanic Garden Edinburgh, Scotland, U.K.
AND
K.E. CLAUSAGER
Doktorvejen 15, DK-9940 Lesg, Denmark
Abstract
Stachyphrynium calcicola, anew species of Marantaceae, was recently discovered in the Bau
limestone area, Sarawak. It also occurs in Kalimantan. The new species is described and
illustrated and the uses of Bornean Marantaceae are given.
Introduction
During a recent inventory of the flora of the limestone hills at Bau, Sarawak, a
species was discovered (Poulsen et al., 2004), which matched other collections
in Borneo but none of the known types of species of Marantaceae. The new
taxon is clearly a Stachyprynium, because the sepals are very short relative to
the corolla tube, whereas in Phacelophrynium K.Schum. and Phrynium Willd.,
the sepals are half the length of the corolla tube or longer.
In Borneo, the most recent revision of Marantaceae included only those
from Sabah (Clausager & Borchsenius, 2003). The earliest account of the
Bornean species by Ridley (1903) included two species, Phrynium parviflorum
Roxb. (which is synonymous with Stachyphrynium placentarium (Lout.)
Clausager & Borchs.) and Phrynium fissifolia Ridl. Existing accounts for nearby
areas, include those for the Malay Peninsula (Ridley, 1899; Holttum, 1951),
and the Philippines (Ridley, 1909). The largest part of Borneo, Kalimantan, is
also the least studied and an intensified inventory on Marantaceae in Kalimantan
with a subsequent flora account for Borneo is therefore much needed, not
least because several species are useful to the indigenous communities
(Christensen, 2002). As none of 13 species of Marantaceae recognized in the
account for Sabah (Clausager & Borchsenius, 2003) are endemic to Sabah,
this account is meanwhile a useful tool for the identification of species
elsewhere in Borneo.
162 Gardens’ Bulletin Singapore 56 (2004)
In Sabah, three species of Stachyphrynium are presently recognized
(Clausager & Borchsenius, 2003): S. borneense Ridl. is widespread in Borneo
and may be synonymous with S. sumatranum (Migq.) K.Schum.; S. latifolium
(Blume) K.Schum. is widespread in the Malesian region (in Sabah it is found
east of the Crocker Range, and a recent collection (Poulsen 2301 from Mulu)
confirms it from Sarawak west of the main range); and S. placentarium is
widespread in SE Asia.
A fourth species, Stachyphrynium lancifolium Suksathan & Borchs., was
recently described from Brunei (Suksathan & Borchsenius, 2003) and this
species also occurs in Sarawak. The new species described below is
consequently the fifth Bornean species of Stachyphrynium K. Schum.
In the description, the terminology of the inflorescence is taken from
Clausager and Borchsenius (2003), as there still remains uncertainty as to the
derivation of the inflorescence and its structure (see Andersson, 1976, 1981,
1998; Kunze, 1985, 1989).
Stachyphrynium calcicola A. D. Poulsen & Clausager sp. nov.
Stachyphrynio borneensi similis in foliorum statura formaque, inflorescentia
elongata differt. Praeterea S. lancifolio similis ac habitu generali ac
inflorescentia elongata necnon corollae tubo longissimo, sed foliorum forma
oblongo-elliptica nec lanceolata, inflorescentia multo maiore ramosa laxa,
corollae lobis multo latioribus, staminodiis exterioribus maioribus differt.
Typus: Malaysia, Borneo, Sarawak, Bau area, Kampung Skiat (1°23°N; 110°
12°E), 30 June 2003. Poulsen, Jugah, Jais & Clausager 2026 (holo SAR; iso
AAU, K, L, Sarawak Biodiversity Centre Flora Depository).
Figure 1.
Rhizomatous ground herb to 50 cm tall. Leaves basal, 2—5 per shoot, distichous;
sheath to 16 cm long, greenish; petiole 8-18 cm long, greenish, glabrous;
pulvinus 10-15 mm long, glabrous; lamina oblong-elliptic with entire margin,
apex acuminate, 26.5 by 9 cm, glabrous, green. /nflorescence interfoliar, erect
to 37 cm long; peduncle 1-17 cm long, glabrous, pale yellowish green;
synflorescence lax, spiciform, simple or branched, to 20 cm; individual bracts
elliptic to oblanceolate, to 3 by 0.8 cm, light green; bracts sub-distichously
arranged, each covering a special paraclade, these with 1—2 flower pairs, each
associated with a 2-keeled prophyll, oblong-lanceolate, 12—19 by 3.5—5.5 mm,
and a lanceolate interphyll 8-15 by and 2.5-4 mm. Flowers c. 30-36 mm
long, cream white except for a yellow touch on the mouth of the corolla tube.
Sepals 3, triangular, free, sparsely hairy at base, 4 by 1 mm. Petal lobes subulate,
A New Species of Stachyphrynium (Marantaceae) from Borneo 163
Figure 1. Stachyphrynium calcicola A.D.Poulsen & Clausager
A. habit; B. sepal; C. flower; D. fruit; E. seed with bilobed aril. Drawn by
Piyakaset Suksathan from the type (Poulsen et al. 2026).
164 Gardens’ Bulletin Singapore 56 (2004)
c. 5 by 3 mm; corolla tube 25—26 mm long, yellow on the inside. Staminodes
and stamen basally adnate to the corolla tube; outer staminodes 2, subequal,
obovate, with petaloid lobe 5—6 by 4-5 mm; cucullate staminode with the free
part 3 by 1.5 mm; callose staminode truncate with the free part 2.5 by 2.5 mm;
fertile anther 2.5 mm long with petaloid cucullate appendage 3 by 1.5 mm;
style hooked with the free part 2.5 mm long; ovary 2.5 mm long. Fruits 10 by
4 mm, with one seed and persistent calyx, 3.5 mm. Seeds 8 by 3.8 mm, aril 2-
appendiculate.
Distribution: Endemic in Borneo in lowland forest. Southern Sarawak and
West and Central Kalimantan.
Ecology and etymology: The type collection was found in open forest with
bamboo, growing near limestone outcrops and a small stream. Labels on most
other collections mention presence of limestone in the habitat, which is the
reason for the choice of epithet.
Notes: Stachyphrynium calcicola is distinguished from all other
Stachyphrynium species from Borneo by the elongated and lax inflorescence.
At the type locality this is at the most with two well-developed branches but
some of the Kalimantan material (Burley et al. 3239) has at least six well-
developed branches. In Borneo, the inflorescence of S. calcicola bears most
resemblance to S. lancifolium, but can be distinguished simply by its much
broader leaves. The leaves of S. calcicola resemble the widespread
Stachyphrynium borneense Ridl., but in this case the difference between
inflorescences is pronounced as S. borneense has compact inflorescences with
red bracts, and S. calcicola has elongate, lax inflorescences with light green
bracts. S. latifolium and S. placentarium are much larger plants than S. calcicola;
the former also differs in the separate and simple inflorescence with water-
containing bracts; the latter by its capitate inflorescence with pointed and stiff
bracts.
In Sarawak, the leaves of most Marantaceae species are used by local
people for wrapping food and for roofing (Christensen, 2002). One collection
of S. calcicola from Sarawak (Jamree et al. §.73204) gives the vernacular
name rirek. Rirek or ririk is a generic name in some Iban dialects for different
species of Marantaceae (H. Christensen, pers. comm.). Some species that are
used as a source of starch (Maranta arundinacea L.) are called mulung ririk,
others where the leaves are used are called daun (meaning leaf) ririk
(Christensen, 2002). The most commonly used big-leaved species,
Phacelophrynium maximum (Blume) K. Schum, is called daun lung besai in
one Iban dialect (Christensen, 2002). Another small-leaved species,
Schumannianthus acaulis Suksathan, Borchs. & A.D.Poulsen ined., is also
called daun ririk in Iban (Poulsen 1943 from Niah, Sarawak). It is perhaps
A New Species of Stachyphrynium (Marantaceae) from Borneo 165
surprising that species with small leaves are used. Possibly they substitute
when the large-leaved species are not available.
Other specimens examined: SARAWAK. SE slope of Gunung Badug, Bau,
lower slope of limestone hill, 300 ft, 1 Apr 1966, Anderson S.19683 (SAR, K);
Gunung Selabor, Upper Sadong Distr., wet place among fallen leaves at foot
of limestone hill, 28 May 1975, Burtt B8210 (E, SAR); Bukit Terebat, Mongkos,
Serian, near river bank, sandy soil, 18 Jun 1996, Jamree, Banyeng & Enjah
S.73204 (SAR, KEP n.v., K n.v., Ln.v.); Bukit Jebong, Bau, at base of mountain
in damp dyke area, forest, 350 ft, 7 Aug 1970, Lehmann S.29433 (SAR); Gunung
Doya, Bau, limestone, 160 m, 26 Feb 2002, Meekiong Kalu et al. SBC 2234
(Sarawak Biodiversity Centre Floral Depository); Gunung Kawa, Bau,
limestone foothill, 200 m, 7 May 2002, Meekiong Kalu & Stephen J.J. SBC
3128 (Sarawak Biodiversity Centre Floral Depository); Gunung Lanyang, Bau,
limestone, rock surface at boulders at foothill on the trail to G. Lanyang, 11
Apr 2002, Julia Sang et al. SBC 2681 (Sarawak Biodiversity Centre Floral
Depository); Gunung Tabai, Bau, limestone, flat area at foothill, 13 Mar 2002,
Julia Sang et al. SBC 2546 (Sarawak Biodiversity Centre Floral Depository);
Gunung Berloban, 10 km Tebakang—Tebedu road, limestone cliff, 270 m, 24
Jun 1983, Yii & Othman S.46243 (K, SAR).
KALIMANTAN. Headwaters of Sungai Kahayan, 5 km NW of Tumbang Sian
logging camp, Sikatan Wana Raya logging concession, primary lowland forest,
150 m, 24 Apr 1988, Burley et al. 798 (A, SAR); West Kalimantan, 5—10 km
N of Masa village, 150 km NE of Pontianak. Ridge SW of Gunung Bentuang,
mixed dipterocarp forest, sloped area, 700 m, 28 Jun — 6 Jul 1989, Burley et al.
3239 (A).
Acknowledgements
The study of Marantaceae from Bau was requested by the Sarawak Biodiversity
Centre (SBC), who also provided permits for research on Zingiberales in
Sarawak for A. D. Poulsen. His fieldwork was kindly supported by HRH Crown
Prince Frederik’s Foundation. We should also like to thank the keepers of the
herbaria SAR, K, and AAU, staff at SBC and Forestry Research Centre for
logistic and other help, J. KF. Veldkamp for Latinizing the diagnosis, and
particularly Piyakaset Suksathan for drawing and providing constructive
suggestions to the manuscript. Finally, we would like to thank F. Borchsenius
and an anonymous reviewer for constructive comments on the manuscript.
References
Andersson, L. 1976. The synflorescence of the Marantaceae. Organization and descriptive
terminology. Botaniske Notiser. 129: 39-48.
166 Gardens’ Bulletin Singapore 56 (2004)
Andersson, L. 1981. The neotropical genera of Marantaceae. Circumscription and relationships.
Nordic Journal of Botany. 1: 218-245.
Andersson, L. 1998. Marantaceae. In: Kubitzki (ed.), The Families and Genera of Vascular
Plants. pp. 278-293. Springer Verlag, Berlin.
Christensen, H. 2002. The Ethnobotany of the Iban and the Kelabit. A joint publication by
Forest Department Sarawak, Malaysia, Nepcon, Denmark & The University of Aarhus,
Denmark. 381 pp.
Clausager, K. and F. Borchsenius. 2003. The Marantaceae of Sabah, northern Borneo. Kew
Bulletin. 58: 647-678.
Holttum, R.E. 1951. The Marantaceae of Malaya. The Gardens’ Bulletin Singapore. 13: 254—
269.
Kunze, H. 1985. Die Infloreszenzen der Marantaceen und ihr Zusammenhang mit dem Typus
der Zingiberales-Synfloreszenz. Beitrdge zur Biologie der Pflanzen. 60: 93-140.
Kunze, H. 1989. Probleme der Infloreszenztypologie von W. Troll. Plant Systematics and
Evolution. 163: 187-199.
Poulsen, A.D., Geri, C., Kalu, M. and Roos, M. 2004. Zingiberaceae and Marantaceae. In:
H.S. Yong, F.S. P. Ng & E.E.L. Yen, (eds). Sarawak Bau Limestone Biodiversity. The
Sarawak Museum Journal 54, No. 80 (New Series), Special Issue 6. Pp. 163-169.
Ridley, H.N. 1899. The Scitamineae of the Malay Peninsula. Journal of the Straits Branch of
the Royal Asiatic Society. 32: 175-182.
Ridley, H.N. 1906. Scitamineae of Borneo. Journal of the Straits Branch of the Royal Asiatic
Society. 46: 229-246.
Ridley, H.N. 1909. The Scitamineae of the Philippine Islands. Philippine Journal of Science,
C. 45: 155-199.
Suksathan, P. and Borchsenius, F. 2003: Two new species of Stachyphrynium (Marantaceae)
from SE Asia. Willdenowia. 33: 403-408.
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