THE GARDENS’ BULLETIN

EDITORIAL COMMITTEE

Chairman: Bernard T.G. Tan, B.Sc.Hons. (Singapore); D. Phil. (Oxon.)

Members: Syed Yusoff Alsagoff

C.J. Goh, B.Sc.Hons. (Singapore); Ph.D (Newcastle upon Tyne)

Leo W.H. Tan, B.Sc., Ph.D. (Singapore)

W.K. Tan, B.A. (Wms Coll., Mass.); M.Sc. (M.S.U., Mich.);

Ph.D. (U.M., FI.)

Lawrence C.C. Leong, B.Sc., M.Sc., Ph.D. (Malaya)

Jennifer Ng-Lim Cheo Tee, B.Sc., M.A. (Dublin)

Editors : Chin See Chung, B.Sc.Hons (Singapore); M.Sc. (Malaya); M.S.,

M. Phil., Ph.D. (Yale)

S. Y. Geh, B.Sc.Hons., M.Sc. (Singapore); Dip.Hort.Sc. (Massey)

Foong Thai Wu, B.Sc. Hons (Cant.); Ph.D. (Cant.); MBA (UK)

Reviewer: Chin, S.C., Botanic Gardens, Singapore

Dransfield, J., Royal Botanic Gardens, Kew, England

Lim, A.L., University of Malaya, Kuala Lumpur, Malaysia

Paris, B.S., Takapuna, Auckland, New Zealand

Prakash, N., University of New England, Armidale, Australia

Primack, R.B., Boston University, Boston, Massachusetts, USA

Soepadmo, E., Forest Research Institute of Malaysia, Kepong, Malaysia

Tattar, T.A., University of Massachusetts, Amherst, Massachusetts, USA

Whitmore, T.C., University of Cambridge, England

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Singapore Botanic Gardens

NATIONAL PARKS BOARD

The GARDENS’ BULLETIN Singapore

Volume 48 (Part 1 & Part 2) June & Dec 1996

Pages 239 and 245 are defective.

We apologise. Replacement copies for these pages are mailed to you

together with this copy of the Garden’s Bulletin.

Thank you.

1.3 ARNOLD

WIN in ale + mAAE r

JUN 15 1898 2730RETUN

The Gardens' Bulletin

Singapore

VOL. 48 (Part 1 & Part 2) June & Dec 1996 ISSN 0374-7859

CONTENTS

Lim, C.K.

Unravelling Jguwanura BL. (Palmae) in Peninsular Malaysia ..............:1cseeseseeeeeereeee 1 — 64

i Lim, C.K.

Palms in the Farquhar collection of natural history drawings .............:.::::seeeee 65 — 74

| LaFrankie, J.V.

Distribution and abundance of Malayan trees:

significance of family characteristics for CONSETVAtION ............ceceeseeseeeeseeeseeeeeeeeeeeerees 75 — 87

Sanderson, F.R., Fong, Y.K., Saiful Anuar, M.S., Yik, C.P., and Ong, K.H.

A Fusarium wilt (Fusarium oxysporum) of Angsana (Pterocarpus indicus)

PEM SEEL EAA FYOL oe see gece es cee nas ve nceats rae ce sucesso ceva stra oudos icbasceltonsbsutadeseqevenoeatbsbszosucouescesdvosnsectase 89 — 127

Turner, I.M., Boo, C.M., Wong, Y.K., Chew, P.T. and Ali Ibrahim

Freshwater swamp forest in Singapore, with particular reference to that

FO MEICATO LEG CHE INCE, SOOM EIN D RANGES -2.-cococerczneconeesscsesesecscesecevanesucesssssesasednces 129 — 157

Zulkifli, M., Latiff, A., Bidin, A.A. and Jaman, R.

A preliminary survey of ferns and fern-allies of Gunung Kajang area,

IESE APB Md OETA AEN cre cote ec cree aaa Beate och a pied sn daenn enn see nonce tocuki<ivuceeseutsoaenetanadceetene suche: 159 — 188

Faridah-Hanum, I.

Morphological variation of Pangium edule Reinw. fruits in Malaysia ................. 189 — 194

Latiff, A., Natrah Mohamad and Zainudin Ibrahim, A.

Ternstroemia magnifica Stapf ex Ridley (Theaceae) and

Kibatalia macrophylla (Pierre) Woodson (Apocynaceae),

EVORSPECIES LEW EO RCMEISU LAL IVI ALAY SIG onc cczscseccnactdedneccrorsecesaseucennccswssevevevcucceveceesesss 195 — 200

Chantaranothai, P.

A new species of Barringtonia (Lecythidaceae) from Peninsular Malaysia........ 201 — 202

Khatoon, S.

Ontogenetic basis of polyad symmetry in Samanea saman (Jacq.) Merr. ........... 203 — 206

Theilade, I.

Revision of the genus Zingiber in Peninsular Malaysia ................2.:2:e:ceseeseeeeees 207 — 236

Saunders, R.M.K.

he Anpiosperm: Plora/ Of Singapore Part «5, 2..0-c-...50.ceosecsaetscnssansusvesossuvavesseassense 237 — 240

Saunders, R.M.K.

eT AMPTOSPELEU MLOLAlGb SIP APOLS: PALt 3) ..-.sr-ccsvccnsedenenrsesivancsscsuseseceseosecoeegeare=ss2 241 — 251

nN ep RSMO ENED oe eto Sateen een chen cnkonctexopexcvevecnsninensecniedennattvestetantiase ZOO —— 2O4

Date of Publication: 28 Feb 1998

Published by

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Singh, H. (1976). Sclereids in Fagraea. Gard. Bull. Sing. 22, 193-212.

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Unravelling [guanura BI. (Palmae) in Peninsular Malaysia

Lim Chong Keat

Palm Search Malaysia Project

215 Macalister Road, 10450 Penang, Malaysia

Abstract

Based on over four years of field studies, a revision of the palm genus Jguanura Bl. in

Peninsular Malaysia is presented, listing 16 taxa including seven new species and one new variety.

Preamble: My interest in palms began with the realisation that I knew little

about them, whether they were introduced exotics or were local and indigenous or

endemic. I decided to learn and investigate firsthand those found within Peninsular

_ Malaysia (initially) and to study them in their natural habitat wherever possible,

and also to research on available information in the herbaria with emphasis on the

genera /guanura, Pinanga and Areca. The rediscovery of historical or forgotten

species has been both rewarding and exciting, as was the encounter with those

that were evidently new. With other team members of the privately sponsored

project called Palm Search Malaysia, we were encouraged at an early stage by the

fortuitous discovery in N. Perak of a new species, Areca tunku, which I published

jointly with John Dransfield who had also found the new taxon in Sumatra and in

Peninsular Malaysia (see Principes 36 (2), 1992: 79-83). The result of the field

work begun in 1989 has provided the essential basis for the revision of /guwanura

within Peninsular Malaysia. As the genus also occurs in other parts of S.E. Asia,

the revision will undoubtedly be interim, and will need to be followed up by

broader regional studies and comparisons. Nevertheless existing documentation

and taxonomic accounts of the Malayan flora require correction, unscrambling or

unravelling, in the light of fresh data and of new species and varieties.

Chronology

The genus /guanura was first described by Blume in 1838 based on the

specimen he named /. /eucocarpa which had been collected in Sumatra by Korthals

and others. The earliest species found in Peninsular Malaysia was from Penang

Island, collected by Porter in 1822 which featured in Wallich’s catalogue of 1828,

and was published by Martius as Areca wallichiana in 1837; it was subsequently

reidentified much later in 1883 by J.D. Hooker as /. wallichiana. Among other

early palms collected in the Malacca area around 1841 was Slackia geonomaeformis

which Griffith published in 1844; his material had also been seen by Martius who

described it as J. geonomaeformis Griff. ex Mart. in 1849, placing it under Blume’s

new genus.

In 1886, Beccari began his monumental publication on Asiatic Palms (Malesia

III) describing his Bornean discoveries, also adding notes on J. wallichiana, using

Wallich’s specimens in the Munich Herbarium. He also claimed a new species, /.

malaccensis, based on Kehding’s collection of 1878 from Klang; which is arguably

synonymous with /. geonomiformis (as per the revised spelling).

Within the section of Malesia II, dated August 1889, Beccari provided

further important accounts based on material supplied by Dr King of Calcutta

which had been collected by Kunstler in Perak, and by Father Scortechini, who

had died of dysentery in Calcutta in November 1886. The new species published

were I. corniculata (collected by Kunstler from Selama), I. bicornis (by Scortechini

from Gunung |juk), 7. polymorpha and its variety canina (also by Scortechini,

from Perak), as well as numerous taxa from other genera.

In 1892, J.D. Hooker published “Flora of British India”, describing Beccari

as the precedent co-author for the section on Palmae, incorporating his manuscript

data on new /guanura taxa, including J. wallichiana v. major , I. wallichiana v.

minor, I. diffusa (collected by Scortechini from “Gunong Tjok”) and J. parvula

(collected by Scortechini from Perak), hinting however that the last might be a

“very small form of J. polymorpha”.

In 1903, H.N. Ridley cited two new species, J. ferruginea and I. spectabilis,

the latter being the spectacular large entire-leafed form found in the G. Bubu

area, which had apparently been preceded by Dr Masters who had exhibited and

published it in 1898 as a botanical exotic called Geonoma Pynaertiana. However,

the syntypes for /. wallichiana var. major are clearly similar to this, and Masters’

species and /. spectabilis are consequently reduced to synonyms of the latter. In

“Materials for a Flora of the Malay Peninsula” published in 1907, Ridley listed

other historical taxa of [guanura, but included J. diffusa and I. parvula as synonyms

of I. wallichiana, and I. brevipes under I. polymorpha. Perplexed by fresh collections

of I. geonomiformis and the variations in leaf division and inflorescence rachilla,

he added to I. geonomiformis var. typica, I. geonomiformis sub. var. ramosa and I.

geonomiformis var. malaccensis (which he had cited in 1903). Within this revision

these will be treated as forms of I. geonomiformis.

The sequel to “Materials” was Ridley’s “Flora of the Malay Peninsula”,

published in 1925, which contained several editorial and textual inconsistencies

within the Palmae section, including the description of Licuala ferruginea in place

of its /gwanura namesake (correct in the 1907 publication), as first pointed out by

Whitmore (1973 endnote 78).

The syntypes of J. ferruginea came from different locations, displaying

variations in inflorescence, but Ridley’s citation positively identifies characters

that are found in J. polymorpha Becc. and its var. canina in particular. Furriness

on which it was distinguished is in fact quite common within the genus and is not

necessarily a reliable differentiation for taxonomic purposes.

@eo-

Plate 1 I. wallichiana var. major: Watercolour by Charles De Alwis titled “/guanura spectabilis

Ridley” (by courtesy of National Parks Board and Singapore Botanic Gardens)

(ee =A oe phts ros ee = —_ — a = = es eS

Plate 2 I. geonomiformis: Watercolour by Charles De Alwis titled “/guwanura geonomiformis”

(by courtesy of National Parks Board and Singapore Botanic Gardens)

Ridley had also collected a specimen from Ara Kuda in Province Wellesley,

Penang, which he labelled “Pinanga canina Becc.”, probably on account of the

fruit shape, although none is on the herbarium sample itself. In 1934, Furtado

published this as a new species, /. arakudensis, characterised by the broad apical

leaflets. Earlier taxonomic attempts to use variations in leaf shape and form as

determinants were then undoubtedly part of a learning curve. As the type location

has since been turned into rubber and oil palm plantations, the species was deemed

extinct (Whitmore 1973), indeed our own field searches have proved abortive.

However, other equivalent low-lying terrain in north Malaysia has yielded enough

specimens to indicate that Furtado’s taxon with its three-branched inflorescence

is, like 1. ferruginea, a form of I. polymorpha.

In 1973 Whitmore published “Palms of Malaya”, intending it to be a popular

guide to the flora, then much misunderstood or unknown. It was a remarkable

work, produced apparently in between more serious work and forestry duties. It

is still the key introduction and reference for many, and provides a clear overview

of the /gwanura (and other) taxa. He revives the identity of /. diffusa, I. parvula,

I. brevipes and I. arakudensis (which he has suggested might be extinct), and

reaffirms /. wallichiana, I.geonomiformis, I.corniculata, I.bicornis and I.polymorpha.

In his earlier “Taxonomic Notes” (Principes 1970, Vol. 14: 124) he has contradicted

Ridley and held that I. brevipes was distinct from /. polymorpha on the basis of

the crownshaft and its interfoliar as against the infrafoliar inflorescence of the

latter. Whitmore however realised and suggested more field specimens were

needed for fuller determination of the 1. polymorpha complex. J.D. Hooker’s

type for 1. brevipes collected from Bukit Larut from the same vicinity as /. bicornis

had no fruit, and its habit, whether solitary or clustering, was not described.

Subsequent herbarium specimens described as /. brevipes (by Furtado, Whitmore

and others) are in my view not at all certain, and are more likely to be 1. polymorpha

or I. bicornis.

Whitmore himself and his colleagues collected strenuously and made

substantial deposits of palm specimens at the Kepong herbarium; his field coverage

and notes have significantly extended postwar botanical investigations. He

rediscovered J. diffusa in Taman Negara; although it has not been seen in the

original type location in Perak, the Pahang form appears to fit the description by

Beccari, per J.D. Hooker.

In 1976, Ruth Kiew (née Evans) published the first major revision of the

genus, covering the entire known range, including Thailand, Sumatra, Malaya and

Borneo (Gardens Bulletin Vol. 28: 191-230). As an extension of her doctoral

interest in the genus, the revision itself “was based on one year’s field work in

Malaya and one month’s field work in Sarawak, in addition to the study of

herbarium specimens”. She lists for the region a total of 16 species including four

new Bornean ones that she has determined, with further taxa at the rank of

subspecies and variety. Later, she added two further new species described from

Sarawak (Kew Bulletin 1978 Vol. 34: 143, 144) with supplementary notes on

Malayan taxa also found in Thailand. For Peninsular Malaysia, she reduces the

spectrum to four species: J. bicornis, I. corniculata, I. polymorpha and I. wallichiana,

with two subspecies for the last: ssp. wallichiana and ssp. malaccensis, each with

two or three varieties respectively, totalling eight taxa at various ranks, including

two new varieties.

Kiew’s revision separates the Malayan taxa on the basis of leaf abscission

or marcescence, and of leaflet shape; her “trapezoidal” section lists the first three

species mentioned above. She errs however by including /. arakudensis within I.

bicornis, whereas it should be regarded as a form of J. polymorpha, under which

she includes the var. canina, I. brevipes, I. ferruginea and also, erroneously, I.

parvula. In summary, she places all forms with parallel-sided leaflets, or with

entire leaves, within the /. wallichiana complex, creating two subspecies, with

synonymy as follows:-

1.1 J. wallichiana (Wall. ex. Mart.) J.D. Hooker ssp. wallichiana

var.wallichiana

(synon.: Areca wallichiana Wall. ex. Mart., I. diffusa Becc., I. wallichiana

var. minor Becc. in. J.D. Hooker)

1.2 I. wallichiana ssp. wallichiana var. major (Becc. in J.D. Hooker ) Kiew

(synon.: I. wallichiana var. major Becc. in J.D. Hooker, Geonoma

pynaertiana Masters, I. spectabilis Ridley)

2.1 J. wallichiana ssp. malaccensis (Becc.) Kiew var. malaccensis

(synon.: Slackia geonomiformis Griff., I. geonomiformis Griff. ex. Mart.,

I. malaccensis Becc., I. geonomiformis var. malaccensis (Becc.) Ridley,

I. geonomiformis sub.-var. ramosa Ridley)

2.2 I. wallichiana ssp. malaccensis var. humilis Kiew

2.3. I. wallichiana ssp. malaccensis var. elatior Kiew.

Kiew attempted to localise her J. wallichiana taxa into an elegant distribution

pattern within the map of the Peninsula, which cannot but be outdated by evidence

from more collections and further field samplings. For example her ssp. malaccensis

can be found more northerly well into Perak and Kelantan.

The basis for Kiew’s preference for Beccari’s malaccensis as against the

Griffith-Martius geonomiformis was that she believed Beccari was first to correctly

identify the characteristic spicate inflorescence, whereas Griffith and Martius

included “branched” inflorescence - which rendered their earlier accounts

synonymous with Martius’ own Areca wallichiana. However, the illustrations of /.

geonomiformis within Martius and Griffith (q.v.) are quite indicative and show bi-

furcating and tri-furcating inflorescences, which are usual variations of the spicate

form commonly seen. These forms are quite unlike the divaricate, paniculate

branching of /. wallichiana and its allies.

In addition to herbarium reference and historical research, the field

observations and new collections within the Palm Search programme lead to a

rather different scenario from Kiew’s overview of the /. wallichiana complex.

Working quite independently, but greatly benefitting from her published work

and friendly communications, I propose to revise and update her listing as follows:-

1.1 LF. wallichiana (Wall. ex. Mart.) J.D. Hooker var. wallichiana

(synon.: Areca wallichiana Wall. ex Mart. I. wallichiana var. minor

Becc. in J.D. Hooker, J. wallichiana (Wall. ex. Mart.) J.D. Hooker ssp.

wallichiana vat. wallichiana);

1.2. L. wallichiana var. major Becc. in J.D. Hooker

(synon.: Geonoma pynaertiana Masters, I. spectabilis Ridley,

I. wallichiana ssp. wallichiana var. major (Becc. in J.D. Hooker) Kiew);

1.3. I wallichiana var. rosea C.K. Lim var. nov.

1.4 I. diffusa Becc. ex. J.D. Hooker

1.5 J. asli C.K. Lim sp. nov.

1.6 JL. kelantanensis C.K. Lim sp. nov.

1.7 I. piahensis C.K. Lim sp. nov.

2.1 I. geonomiformis (Griff.) Mart.

(synon.: Slackia geonomiformis Griff., J. malaccensis Becc., I.

wallichiana ssp. malaccensis var. malaccensis (Becc.) Kiew, I. wallichiana

ssp. malaccensis var. elatior Kiew)

2.2 I. humilis (Kiew) C.K. Lim stat. nov.

(synon.: I.wallichiana ssp. malaccensis var. humilis Kiew)

It might be mentioned at this juncture that the range of variations and

forms within the wallichiana and geonomiformis groups gives scope for further

field investigations. Kiew’s var. elatior remains tantalising, but seems to be similar

to certain other widely distributed forms of 1. geonomiformis. There are indeed

other robust forms of the taxon to be seen in G. Bintang and in Negeri Sembilan;

the latter might well be the epicentre on the basis of size. I. humilis deserves its

new status, as was indeed suggested by Furtado on Corner’s specimen; and although

other forms of J. geonomiformis can have precocious inflorescences at the

acaulescent stage, they are usually clustering, whereas the new taxon is solitary

with a short stem. Our field observations reveal certain other “grey” areas between

the two groups or within each, but this has been seen mainly in particular locations

(Pahang, Kelantan and Terengganu). In the more well-known collecting areas the

distinctions seem evident enough to justify maintaining the historical (and

convenient) separation of J. wallichiana from I. geonomiformis.

The field strategy has obviously been to check on or to get close to type

locations, and to corroborate against type specimens. J. wallichiana (var.

wallichiana) can still be found on Penang Hill, but in diminishing stands. I.

geonomiformis is probably extinct at Ayer Panas, but is not uncommon within the

Malacca area. I. diffusa has yet to be found in the G. Ijok (or G. Hijau?) vicinity,

but the Taman Negara form (or its look-alike) is quite accessible and distinctive.

Within the . wallichiana complex, Ridley’s I. spectabilis would have been a

wonderful name, which would have been subsumed into “/. pynaertiana”. However,

as mentioned above, they are both synonymous with Beccari’s J. wallichiana var.

major, the distinctive taxon with erect inflorescence and large entire leaves, as

seen on G. Bubu. Other forms (eg. J. wallichiana var. minor) can be readily

differentiated from this variety in the field, and are included under J. wallichiana

var. wallichiana.

I propose to recognise a new variety, /. wallichiana var. rosea, firmly on its

distinctive fruit colour, which is pink from the start, ripening red, as found in two

particular locations in Perak and Kelantan. Fruit colour difference as a positive

character is also found in the new species to be named I. asli, which is quite

widespread within [. geonomiformis (Kiew’s var. elatior) territory, from eastern

Johor to Pahang and Terengganu, which similarly has immature pink fruit turning

candy pink (rather than cerise red), somewhat rivalling adjacent populations of

Licuala ferruginea in colour and splendour. The inflorescence is interfoliar, with

curled rachillae, and short peduncles within the sheaths.

L.kelantanensis is likely to be more widespread but has so far been seen

only in its type locality within the named state. It is acaulescent, and solitary, with

variable branching inflorescence, usually having short peduncles.

Another new species, which has so far been seen only in a particular area

and is thereby named /.piahensis, has a particular distinguishing characteristic that

may intrigue the student of dried leaves (as on herbarium sheets) - its fine texture,

papery and silky to the touch. Furthermore, the leaf shapes are distinguishable

from others of the wallichiana/geonomiformis complex; the entire forms are

unusually long and parallel-sided. The lobed forms can also be identified, with

some comparative practice. Its inflorescence is branching, usually fine and modest,

similar to some collections of “/. wallichiana var. minor”, which however has

standard wallichiana type leaves.

The rediscovery of Beccari’s diminutive /. parvula was indeed the

providential highlight of our field journeys. Apart from Whitmore’s steadfast

belief, it had been regarded either as an aberrant or immature aspect of J.

polymorpha or of I. wallichiana (per Ridley). Some strange magnetism led us to

stray into its remnant (and threatened) habitat in Kedah, in rather unlikely

circumstances, to find among logging debris this highly distinctive clustering palmlet,

then “new” to us. A month earlier, I had made a series of photographs of Beccari’s

type specimen without attaching importance to its species status, and was therefore

able to make a clear identification. Beccari was indeed so right in his descriptions.

Although the type location was elsewhere in neighbouring Perak, it does not seem

to have been collected there since. However, I subsequently located a fine specimen

in the Singapore Herbarium collected also from Kedah by Haniff (SFN 21104) in

1928, which had been filed under /. polymorpha despite Furtado’s correct

determination.

The other two unmistakable Beccari species are of course J. bicornis and I.

corniculata, which, as might be said, require no introduction - after the full

description in Malesia III. The former is relatively common, but has so far not

been found other than in Kedah and Perak, and usually on high ground above

500m. Our field observations show a range in size and stature from small clumps

not taller than 1m to robust ones reaching over 3m. The largest of the fully ripe

bigibbous fruit that we have encountered measure 2cm across, and are luscious

and sweet, as all Jguanura drupes should be. /. corniculata is a different proposition

in terms of its survival. Because of the fine type specimens made by King’s

collector, Kunstler, from Selama, Perak in 1881, it had remained listed as hopefully

extant but probably extinct, and as mentioned by Kiew and others, it had not been

seen or collected since, ie. within the last 100 years or so. I would be inclined to

treat as suspect the specimen collected by Henderson from near Raub, which he

himself considered to be J. ferruginea. Several forays into the Selama localities

have been abortive, and the prospect of discovery there seems dismal as logging

and plantation expansion is extensive, especially in the altitude range of 100-

150m. I finally found the elusive taxon during the most recent stages of field work

- not in Perak, but in its adjacent state - providentially indeed to rediscover I.

corniculata within Penang boundaries, after failing to resurrect J. arakudensis!

Beccari’s type specimen of /. polymorpha and its var. canina were apparently

collected on Bukit Larut, which was also the type locality for 1. bicornis and for

J.D. Hooker’s I. brevipes - all from altitudes above 1000m. Ridley’s /. ferruginea

syntypes. were from two locations. Later collections by Ridley and others came

from various places, Perak to Kedah and Kelantan; several were from low altitude

areas, for example, Furtado’s 1. arakudensis, from Province Wellesley. The cited

lectotype for 1. ferruginea (Fox 10684) had a stout 10-branched inflorescence and

is clearly synonymous with I. polymorpha. The other syntype seemed different,

with 2 rachillae but with the characteristic curved fruit. The taxon is variable in its

rachillae, often trident, sometimes with 1 or 2 branches, (suggesting a link with /.

corniculata). From our own field collections and other herbarium specimens we

have noticed that sterile specimens of I. bicornis and I. polymorpha have sometimes

been confused. Indeed /. brevipes might have been based on a bicornis specimen.

With regard to the inter/intrafoliar distinctions set out by Hooker (and later

espoused by Whitmore), Kiew has elucidated on this, pointing out the consequences

of variable abscission conditions. Our own observations are on similar lines, as all

taxa in the “trapezoidal” group can be seen with inter and/or infrafoliar

inflorescence with the obvious conclusion that the peduncle was able to hold

laden fruit before or after leaf fall, even if some heavier ones would be more

“comfortable” while being interfoliar. Besides, the conditions for abscission would

be different for clustering, as from solitary forms of growth.

Returning to the 1. polymorpha complex, the scenario initially seemed to

suggest the existence of 4 taxa: 1. polymorpha (with ovoid, not curved fruit), /.

polymorpha var. canina (with distinctly curved fruit), and /. ferruginea/arakudensis

(also with curved fruit) for lower altitude forms (all clustering and not solitary).

Although we have not yet seen live populations of 1. polymorpha or its allies on

Bukit Larut, we have observed interesting counterparts on Cameron Highlands,

where a range of variation in leaf divisions can be seen. There, within particular

colonies, the fruit shapes vary from globose to ovoid and curved, with seeds

thicker than for I. ferruginea forms. The lowland variant is found all over Perak,

Kelantan, Terengganu and Pahang, and is the most commonly encountered form.

Our own relatively recent collections from upper Perak, in the vicinity of

Terunok (according to Ridley and Whitmore the local word for /guwanura, and

perhaps for this particular taxon itself), have broadened our concept of I.

polymorpha. As against the 2-2.5m clumps that we had seen elsewhere, we were

astonished by the robust 3-3.5m stems, with leaves twice as large (as elsewhere)

and an amazing variety of inflorescence form, from two to nine branches, the last

sub-branching to 16 or more. Some rachillae were over 75cm in length, inclusive

of the broad (1.5cm) peduncles, borne infrafoliar or interfoliar. On some rachillae

we counted over 95 buds; the immature fruit were typically curved and irregular

in its process of distention. The altitude was c.300m., with more colonies likely

higher up or lower (now under water level because of damming). The evidence

seems incontrovertible that we are faced with a single main taxon of J. polymorpha.

I. polymorpha var. canina, I. brevipes, I. ferruginea, and I. arakudensis should thus

be regarded as synonyms. Beccari’s taxon is indeed polymorphic.

Again within Perak, we have found two new members of the “trapezoidal”

complex, the first solitary in habit identified in 1989, which we then decided to

name I. belumensis, after its location. Later we came across herbarium specimens

of the same or a similar taxon, originating from Bujang Melaka and G. Bubu,

labelled as 1. polymorpha. The other new species is exceedingly rare and highly

endangered; over many trips we have yet to find more than ten plants in its only

known location. It is a 4m tall, solitary, (occasionally with basal branches) giant

Iguanura, with two forms of leaf divisions, twice as large in size as the sympatric J.

bicornis neighbours. It is to be called 1. perdana.

The underlying paradigm behind the Palm Search project has been to seek

in the field as widely and repeatedly as possible, in parallel with herbarium research

work. Our original purpose was to learn about existing, known taxa and their

conditions of endangerment. To alight upon a new species is surely an act of

providence, especially in unsuspecting circumstances, but it has undoubtedly given

fuller impetus to persistent botanical effort.

The ultimate new taxon among Peninsular /guanura was stumbled upon in

a chance sidetrip from a main botanical field expedition. In a sense it is not

“new ’’, in that countless locals and foresters would have known it, and as it happens,

a juvenile specimen had been collected and emplaced within the Kew Herbarium,

awaiting determination, as we subsequently learnt. The new species to be called /.

mirabilis is caespitose and has strongly plicate entire leaves and curved fruit, and

although unlike the “trapezoidal” members, it also falls within the abscissing group.

Affinities with the Sarawak taxa, especially /. elegans and I. sanderiana, are

suggested, but the new taxon is clearly on its own within the genus.

With regard to local names for /gwanura, Ridley (1903) had recorded that

the striking palm he called /. spectabilis was known to the (local) Malays as

“Teruno” (Tronoh, Terunoh, Terunok), a name now unfamiliar to most. Griffith

earlier noted that the 1. geonomiformis was called “Pinang Rambeh”, to which

Ridley added “Pinang Sapadan”, further noting that “Pinang Pachat”, “Pinang

Burong Tikus” and “Pinang K’lasak”, were used for /. wallichiana. According to

Kiew, the latter is called “Terunok” in Kelantan. In north Perak, along the river,

there is a place called Terunoh, probably a village now obscure or vanished under

water. In that vicinity, robust forms of J. polymorpha abound and it might be

conjectured that the word, clearly botanical, applied historically to this particular

taxon. Burkill (1935) quotes and extends Ridley’s list, and noted that 1. corniculata

was called “Pinang Angin”. In his reference however, “Terunok” was misprinted

as “Termoh”. Many loca! names might have been ad hoc concoctions. Generally

there has been a decline in specificity in knowledge of traditional names for

plants. Most understorey palms including the /gwanura and the ubiquitous Pinanga

malaiana (known as “Legong”) are loosely now referred to as “Pinang”. The

Temiar (Senoi) folk are however still in command of their botanical names, also

for the palms (including rattans) within their traditional domain. They call the

Iguanura wallichiana alliance “Bog”, but do not use this for the polymorpha

forms not known to them. With more anthropological research, other aboriginal

names will be reidentified, as used by the Semai and others.

Many of the /guanura taxa may be considered endemic to the Peninsula,

although territorial boundaries are in fact arbitrary from a botanical point of view;

South Thailand, for example, is indeed part of the same bio-domain, unlike Sarawak

and Sabah, which are placed in a separate botanical division, although within the

same nation. Knowledge of the confines of a taxon depends on the extent of the

collections and will be modified by further field surveys; for example, J. parvula

and the new species [. belumensis may well be found in South Thailand, where J.

polymorpha and I. wallichiana abound.

Our unravelling of /guanura in Peninsular Malaysia is intended to provide a

useful framework for a fuller regional taxonomy of the whole genus. Further field

work will undoubtedly yield fresh data and refinements or shifts in perceptions,

but the next priorities lie in widening the territorial scope to include Thailand and

Sumatra, our immediate floristic bio-domain, and then Borneo, in particular

Sarawak, where other exciting species of Jguanura are to be found, surprisingly

different from the Peninsular forms. Contrary to earlier suggestions (Kiew et

alia), preliminary observations already suggest that none of the Malayan taxa are

to be found in Sarawak.

Iguanura Bl. (Palmae)

Bulletin des Sciences Physiques et Naturelles en NEerlande 1: 66. 1838.

Type: I. leucocarpa Blume; Kiew, Gdns. Bull. 28: 191-230; Uhl and Dransfield,

Genera Palmarum: 421-423. 1987.

Blume first described the genus in 1838 with 1. leucocarpa collected from

Sumatra as his type species. Slackia of Griffith, exemplified by S. geonomiformis

published in 1844 is thus a synonym of the genus. Full citations have been listed

in Kiew (1976), and most recently by Uhl and Dransfield (Genera Palmarum: 421-

423, fig. 133. 1987). These include similar taxonomic accounts, and Uhl and

Dransfield have classified the genus within the tribe /guanurinae J.D. Hooker in

Bentham and J.D. Hooker, under the sub-family of Arecoideae.

The present revision is confined to taxa found in Malaya, and will contribute

fresh data on new species and varieties, with more emphasis on field characters

and information, but less on herbarium research requiring laboratory and

microscope work, eg. on floral, pollen or DNA examinations, which require fuller

samplings and remain largely unstudied, giving scope for further inputs.

Blume’s type species, I. leucocarpa, is of the form with trapezoidal leaflets,

ie. similar to the polymorpha complex, with cleanly abscissing leaves, but the

genus now includes other entire-leafed, marcescent forms. It is not certain whether

or not J. leucocarpa was solitary in habit, but Kiew’s account confirms that the

fruit was olive-shaped, the inflorescence was spicate, seeds “not ribbed or ridged”,

the drupes were white in colour (per the epithet, when unripe). So far no Peninsular

taxa have these seed features. Presumably from examination of the type specimens

at Leiden and Bogor, she noted that the anthers were lobed, a character that she

has recognised and confirmed also for certain Sarawak species (J. sanderiana, I.

ambigua and I. remotiflora), and by implication suggesting that all other species

had anthers not lobed. This may need to be rechecked with more samplings. It

might be suggested however, that anther lobes may tend to crenulate in drying,

and therefore examination of fresh material may be essential.

For purposes of this revision I have examined the /guanura collections in

Calcutta, Florence, Kepong, Kew and Singapore, and also at UPM, Kuching and

Sandakan. Unless otherwise indicated, all specimens quoted have been viewed.

This has complemented the substantial field data and our own collections, providing

the basis for determinations and for recording distribution of the taxa. Many of

the herbarium specimens, both historical and recent, need to be verified; indeed, a

major updated checklist correlating specimens in Calcutta, Florence, Kew and

Singapore needs to be made, which will also be interesting for botanical history.

Pending this, and to avoid perpetuating existing discrepancies, I allude mainly to

types and syntypes for the revised taxa and their synonyms, and to limited selected

specimens, listed under Reference Collections, including those in the Palm Search

Malaysia herbarium (designated here as PSM).

Within its natural habitat, /gwanura is arguably the most attractive of

understorey palms and is easily recognisable by its leaf shapes and forms, with

their distinctive praemorse margins, and the red or bronze flush seen in new

leaves. At bifid seedling stages, all taxa are practically identical. With some

experience, however, the eophylls and juvenile leaves of the two main groups - the

marcescent wallichiana-geonomiformis complex and the abscissing trapezoidal

alliance - can be differentiated, the latter being darker green and more oblong. J.

mirabilis is particularly distinct in its discernibly plicate leaves. Historically,

botanists have dwelt on leaf divisions, shapes and sizes, petiole lengths etc. as

determinants. For most taxa, we must now recognise a wider range of variations.

Leaf forms may be entire or divided; the division may conveniently be described

as lobed or pinnate. Certain species when mature continue to have undivided

bifid leaves, which vary in shape from cuneate to oval and oblong, all with the

characteristic apical cleft. They also vary in size from the diminutive 20cm long /.

parvula to the 2m or longer gigantic blades of /. wallichiana var. major. Large

bifid leaves tend to have short petioles for structural reasons.

Leaf divisions described as lobed, are those split into broad leaflets, with

two to five pairs (or more), opposite, alternate or irregular; the leaflets may be

spaced closely (with short rachial lengths between) or more distantly. The more

common leaf division is the pinnate form with many pairs of narrow parallel-sided

leaflets, opposite or alternate, but usually with broader apical leaflets. An extreme

form of division can be seen in /. diffusa, where unicostate leaflets (including

apical ones) have been found. Both for /. wallichiana and for I. geonomiformis

and their allies, the lamina can vary from entire to lobed and pinnate, within the

same colony, and sometimes on the same plant.

Leaf sheaths are either fibrous and persistent (i.e. marcescent), or self-

peeling and abscissing; this differentiation is important, as was well appreciated by

Kiew; and obviously has a bearing on the nodal scars on the stems and the

internodal dimension. Apart from the entire-leafed 1 parvula and I. mirabilis,

others with abscissing leaves are those that divide (after the juvenile stage) into

trapezoidal leaflets, closely or more widely spaced along the rachis, opposite or

alternate, sometimes quite irregular in leaflet widths (as in 1. polymorpha). The

leaflets which flare distally are like butterflies in shape, but there are great

variations, sometimes not rhomboid but almost triangular. Again, the leaf divisions

may be lobed or pinnate, with two to twelve pairs of leaflets. The apical pair are

usually broad and multi-costate; the tip angles can vary from acute to right-angled

or obtuse.

Herbarium specimens, being desiccated, usually fail to convey the texture

and relative thickness of lamina and other parts observed in the forest. Although

there are broad variations, especially between juvenile and older leaves, certain

taxa have a distinctive sheen and tactile quality on their leaf surfaces, as with I.

parvula and I. piahensis, where the silky texture can be relied upon for field

identification. Prominence of nerves and the leaf fold is also characteristic in the

case of I. mirabilis, which has other obvious features. Colour is also lost on

herbarium sheets, though botanists have tried to describe the hue after drying.

Tomentosity and glaucosity are often obscured. Indumentum is usually well

preserved, but this character is often variable and found in most taxa, on the

sheaths and rachis, leaf nerves and inflorescence.

All [guanura are stilt-rooted. Some are caespitose and clustering, others

are solitary, but occasionally with basal branches. Stems vary from the arudinaceous

to the robust, eg. 2. parvula (Smm in diameter, 1m in height) and /. wallichiana

var. major (2cm in diameter, up to 4m or more in height), respectively. Basal

branches may produce precocious fruiting at low level, especially when the main

stems have been destroyed. Caespitose taxa may also appear to be solitary before

basal branching develops (as is often seen in 1. geonomiformis).

Inflorescences spring from within the leaf sheaths, usually interfoliar, but

for abscissing forms, they can also develop in an infrafoliar position. The peduncles

can be long and extended (sometimes erect, as with /. wallichiana var. major), or

short and enclosed within the sheaths. The inflorescence may be spicate or

divaricately paniculate, (varying in robustness from stout and “succulent” to fine

and “twiggy”), the latter sometimes branching to second or third orders. The

spicate forms (J. geonomiformis complex) may split or “furcate” into two to seven

parallel tails (sometimes sub-branching), but unlike the paniculate forms, the

rachillae are not divaricate (as in J. wallichiana and its allies). As a convenient

gauge of relative inflorescence size, that of the ubiquitous I. wallichiana is often

90cm long (including a 65cm peduncle); and of 1. geonomiformis, 80cm long.

Floral characteristics include the deeply sunken triads within the rachilla,

spirally disposed. Kiew (1976) gives a full account of the details, indicating that

“the male and female flowers are uniform throughout the genus, except for the

lobing of the anther” (ibid. 199). There are of course variations in sizes and

colours of flowers, but these have not been examined within this revision, in the

absence of adequate samplings, and also because of our field-based emphasis.

Anther lobes have not been observed in Peninsular Malaysian taxa, and is a

character more pertinent to those in Borneo, as also the ridged or ribbed seed

forms found there. We have also not examined systematically the seed endosperm

for the variations in homogeneity, whether ruminate or equable; this can be the

subject for further study. For present purposes, fruit shape is adequately indicative,

and the basal stylar remains on /guanura fruit is indeed the unmistakable feature

of the genus. The fruit develops eccentrically, and may be globose, ovoid, olive-

shaped, oblong and bigibbous (J. bicornis), or “canine”, curved (J. polymorpha),

and hooked (J. corniculata). Fruit sizes vary; those described as large would be

near 2cm in diameter, the average being 1.2cm and the small, 8mm.

The pericarp forms like a mould within which the endosperm solidifies

relatively later after the drupe assumes its shape. The endocarp then swells to

sweet, luscious fullness. It is important to note the fruit colour. Commonly,

young fruit is white turning cream or orange, then ripening cerise to crimson red

(eventually black). In some taxa the drupes are light green turning yellow to

orange and red, as in /. parvula and I. piahensis. There also appear to be distinct

forms that have pink (not white) fruit turning bright candy pink and dark red,

consistently seen in particular populations, a determining character which I propose

to recognise.

The revised list of 16 taxa has been arranged not alphabetically or

chronologically, but by similarity, which I believe to be convenient for ease of

comprehension. Its accompanying key is based on simple field characters that

facilitate identification, without laboratory equipment. Generally, descriptions,

notes on colours, textures etc., are based on live and not on dried specimens.

From field surveys of extant /guwanura populations, I have also given weight to

location and distinctive characters (eg. habit and fruit colour) which are not seen

as intermediary forms in other areas. Accurate collection notes on these would be

essential for future herbarium identification, and adequate samplings from

populations would be desirable.

In summary, the key adopts the following main features and differentiations:

leaves marcescent or abscissing; habit caespitose or solitary; leaves entire or divided

(lobed or pinnate), leaflets parallel-sided or trapezoidal; texture of lamina;

inflorescence spicate (splitting or “furcating”) or paniculate (branching divaricately,

sub-branching); fruit shape globose, ovoid, curved, hooked, or oblong and

bigibbous; fruit colour (within location confines).

Revised List of Jguanura in Peninsular Malaysia

16 taxa, including seven new species and one new variety, and their synonyms

1. [guanura wallichiana (Wall. ex Mart.) J.D. Hooker var. wallichiana

synonyms: Areca wallichiana Wall. ex. Mart.

I. wallichiana var. minor Becc. in J.D. Hooker

I. wallichiana (Wall.ex.Mart.)J.D.Hooker ssp. wallichiana. var.

wallichiana Kiew

2. Iguanura wallichiana var. major Becc. in J.D. Hooker

synonyms: Geonoma pynaertiana Masters

I. spectabilis Ridley

I. wallichiana ssp. wallichiana. var. major (Becc. in J.D. Hooker)

Kiew synon. nov.

Iguanura wallichiana var. rosea C.K. Lim yar. nov.

Iguanura diffusa Becc. in J.D. Hooker

Iguanura asli C.K. Lim sp. nov.

Iguanura kelantanensis C.K. Lim sp. nov.

Iguanura piahensis C.K. Lim sp. nov.

ea Sy ae oe

Iguanura geonomiformis (Griff.) Mart.

synonyms: Slackia geonomiformis Griff.

I. malaccensis Becc.

I. geonomiformis var. malaccensis (Becc.) Ridley

I. geonomiformis sub. var. ramosa Ridley

I. wallichiana ssp. malaccensis var. malaccensis (Becc) Kiew

synon. nov.

I. wallichiana ssp. malaccensis var. elatior Kiew synon. nov.

9. Iguanura humilis (Kiew) C.K. Lim stat. nov.

synonym: J. wallichiana ssp. malaccensis var. humilis Kiew synon. nov.

10. Iguanura parvula Becc. in J.D. Hooker

11. Iguanura bicornis Becc.

12. Iguanura corniculata Becc.

13. Iguanura polymorpha Becc.

synonyms: J. polymorpha var. canina Becc.

I. brevipes J.D. Hooker

I. ferruginea Ridley

I. arakudensis Furtado

14. Iguanura belumensis C.K. Lim sp. nov.

15. Iguanura perdana C.K. Lim sp. nov.

16. Iguanura mirabilis C.K. Lim sp. nov.

Key to identification of Peninsular Malaysia taxa of Iguanura

Leaves persistent (marcescent), leaf sheath fibrous «0.0.0.0... eeeeeeseeeeeeeeees 2

Leaves abscissing, leaf sheath smooth inside ...............eseseeseeseeeeeeeeeeeeeeeeeeeees 10

Inflorescence spicate, or furcating into tWO tO SEVEN ......... ee eeeeeeeeeeseeeeteeeeesees 3

Inflorescence paniculate, with divaricate branches sometimes to second and

LEE EP a I Ee nO earn Ree ee ee ~

Caespitose (sometimes solitary), stems to 4m; leaves entire, lobed or pinnate

ee Eee ee ee ace ahccasts alba deaatcnucadponseatucsanth sotnonl 8. I. geonomiformis

Solitary, stem to 50cm, often covered with roots; leaves entire or pinnate

Bee aaa e ce eects tengo ans tene xe eot ens llawadvendacostsuscsoesdbccenesd socesdactase 9. I. humilis

LEE a heise Seas LR dks TR Roy na oe 2 a 5

RSs ALO tape san tesco ac ban donc cs otc scsse esas eoncvacuasseseennssrvavedteabeesensvuatedsiesties 6

Acaulescent; leaves pinnate; inflorescence variable, peduncle often within

BRA aI eset ene eae eae a anet ne ava coassshadbsvl actecusttaaizenee 6. I. kelantanensis

Stem to 1m; leaves pinnate; leaflets often unicostate; inflorescence branching

Rs A GION PEIEEE, CORA Ta ood 2 2ecaSacestee ne enciin a fenssueoswdongonctatcesecscanestevatucs 4.1. diffusa

Fruit globose, pink (not white) ripening deep pink or red ........ eee qf

Fruit globose, white, light green, yellow ripening red ..............eseseeeeeeeereeeee 8

Peduncle short to 15cm, within sheath, rachillae curled .................. 5. I. asli

Peduncle long to 50cm or more, rachillae divaricate; uncommon ...............++.

Pa Pe ea Fe Bch cone eet nga his ieudyennh svestncectvadcbcces 3. I. wallichiana var. rosea

Fruit small to 8mm diameter, light green (not white), cream, orange ripening

to red; inflorescence short, rachillae slender; leaves oblong, entire, or lobed,

PRNRIy SACRE sored as teen eae PE oh EAS s Pea vt rasenndcastbese ce ncedonavsnnveeatve 7. I. piahensis

Bat Memmi to lake, Nem Giamerer OF MOLE \..2....22sSisse eeecaee a cae ce tess -cecacee 9

fale

13:

14.

iS.

Leaves pinnate or entire to 1.3m; inflorescence to 90cm, axillary, spreading;

COMMON, 3-4 ees vecnaserece eee 1. I. wallichiana var. wallichiana

Leaves usually entire, very large to 2m or more; inflorescence to more than

{00cm; eréct-and robust S22... eee 2. I. wallichiana var. major

Te aves @ntare: fai iagececccoc te easkenteeee Pere Ora cee ee oe Soe Eo 11

Leaves lobed or pinnate, leaflets flaring distally (trapezoidal) ................. AD,

Leaves small to 30cm, oblong; stems caespitose, arudinaceous, short to 1.2m;

peduncle imsheath: trum sSlohose sere re eee 10. I. parvula

Leaves large to 85cm, cuneate, often truncated at apex, strongly plicate;

stems caespitose, to 2cm diam., to 3m; inflorescence three to nine branched

with purple indumentum; fruit cumveds 125 essences 16. I. mirabilis

SOMMATY. cccicengetiodes esd ose tite dec tecae ccc cee cette ee tee eee eee Oe nec 13

CaespitOse -...0..5is0s..cecesasdelnegeodeaveedsnanedstopscedecn cater scope teae sens caeme ee dees toe eee eee 14

Fruit ovoid or curved; stem 1.5cm diameter, to 3m; leaves to 60cm;

inflorescence interfoliar or infrafoliar, paniculate with twiggy branches

Saba betiisacan didn esc aE ON EN oA OE EEE 14. I. belumensis

Fruit eccentric with flat top; stem (sometimes with basal branches) 2cm

diameter to 4m; leaves to 100cm; inflorescence usually interfoliar, paniculate

with succulent branches; very mane ats: c ence ree rece aces 15. I. perdana

Inflorescence spicate, to 20cm or more; fruit curved, sometimes with a

pronounced HOOK wer vcceeti sehen sess tere aoe ene eer 2. I. corniculata

Inflorescence branched (rarely spicate, if so less than 10cm); stems to 3m

Fruit bigibbous, inflorescence with five to seven rachillae; stem to lcm

diameter, chestnut colour; habitat montane .............::ceeee 11. I. bicornis

Fruit ovoid or curved (not hooked); inflorescence variable, three (rarely

two or one) to nine rachillae, sometimes sub-branching; stems arundinaceous

or robust, not chestnut colour; habitat sea level to montane, distribution

WIGESPTE A» ...cactstssvcrss vncens oars sures coopera ences ae eee ene ete eared 13. I. polymorpha

Plate 4 I. wallichiana var. wallichiana: Characteristic inflorescence, paniculate, to 90cm.

1. Iguanura wallichiana (Wall. ex Mart.) J.D. Hooker var. wallichiana. Areca

wallichiana Wallich’s Catalogue no. 8600 (1828); Martius, Hist. Nat. Palm. 3: 178

(1837); Beccari, Malesia. 3: 100 (1886): J.D. Hooker, Fl. Brit. Ind. 6: 416 (1892);

Ridley, Mat. Fl. Mal. Pen. (Monoc) 2: 151 (1907); Fl. Mal. Pen. 5: 14 (1925);

Whitmore, Palms of Malaya: 65 (1973); Kiew, Gdns. Bull. 28: 216-224 (1976).

Synonyms: [. wallichiana var. minor Becc. in J.D. Hooker, FI. Brit. Ind. 6:

416 (1892); I. wallichiana (Wall. ex. Mart.) J.D. Hooker ssp. wallichiana var.

wallichiana Kiew synon. noy., Gdns Bull 28: 221 (1976).

This earliest of guanura taxa collected in Malaya, by Porter in 1822, came

from Penang, and from examination of remnant populations on Penang Hill at c.

600m, a good idea of the characteristics of the topotype can be obtained. It grows

on hill slopes, along or within rocky streams, and is caespitose (not solitary),

clustering in a rather untidy way, with stems up to 2m, and persisting pinnate

leaves. The inflorescences extend out from among the leaves, with long peduncles,

branching with several divaricate rachillae (usually robust, but sometimes finer).

The species is to be differentiated from 1. geonomiformis mainly by the inflorescence

configuration. Although in the latter it is often spicate, forms that fork into many

parallel rachillae are also common. The fruit are characteristic of the genus,

globular with basal stylar remains, and ripening in colour stages from ivory white

or light green to cream, then cerise or crimson, eventually black. Eophyll and

juvenile leaves are easily identified as of all within the genus, always entire, cuneate

or oval, and new leaves which may be entire or lobed are often flushed, reddish or

bronze.

From viewing the syntypes of var. minor (King’s Collector 454, 7941, 7999),

I concur with Kiew that they are within the range of forms of /. wallichiana var.

wallichiana. As described in J.D. Hooker, with stems up to 30 inches, and leaves

simple or pinnatisect, and finer inflorescence and fruit, they represent the smaller

variants, with entire or divided leaf forms, which Beccari called “flabelliformis” or

“pinnatisect”.

The main taxon is variable in stature, with forms that are taller, up to 4m in

height, often with dominant stems and basal branches, and thrives at altitudes

from lowland to mountain slopes at 1000m. Variations in leaf divisions and leaflet

widths can also be observed; entire or lobed forms are not uncommon, but the

pinnate with many leaflets are also ubiquitous. The northern populations could

be considered typical of the species. Those in Perlis thriving in riverine swamps

near limestone outcrops, point to a larger common domain that would include

south Thailand.

Distribution: Kedah, Penang, Perak, Perlis.

Type: Penang, 1822, Porter 8600, (holotype K)

Reference Collections:-

Kedah, Weng, Furtado 33070, 33071 K, SING; Cherok Bokbak FR, Meh

10100 K;G. Bongsu C.K. Lim H 1088 PSM Collection; Mahang, C.K. Lim H 11, H

81, H 955, H 976, H 1019, H 1042, H 1046, H 1086, H 1203 PSM Collection;

Sintok, C.K. Lim H 29 PSM Collection; G. Inas, C.K. Lim H 99 PSM Collection;

Penang, Penang Hill, 1879, King’s Collector 6002 CAL; Ridley 7101, 10344 SING:

Bukit Mertajam, C.K. Lim H 24 PSM Collection; Bukit Panchor, C.K. Lim H

1567 PSM Collection; Perak, Gopeng, 1880, King’s Collector 454 (Type for I.

wallichiana var. minor) FI, K; Batang Padang district, 1885, Kunstler 7941 FI, 7941

K, 7999 FI, K; Selama, King’s Collector 3127 Fl, K; Chior FR, F.S.P. Ng FRI 5792

KEP, K; Pos Legap, Gianno E 326 K; G. Kerbau (Korbu) Robinson s.n. K, C.K.

Lim H 1137, H 1249 PSM Collection; Kg. Sukam, Avé 705 K; Bintang Hijau FR,

Everett FRI 14529 KEP, K; Kenayat, C.K. Lim H 44, H 109, H 619 PSM Collection;

Banding, C.K. Lim H 128 PSM Collection; Sg. Piah, C.K. Lim H 396, H 455, H

549, H 1100, H 1108, H 1591 PSM Collection; Pos Salim, C.K. Lim H 413, H 418

- PSM Collection; Kg. Ijok, C.K. Lim H 942, H 1377 PSM Collection; Sg. Bayor,

C.K. Lim H 956 PSM Collection; Krokh, Furtado 33008 K, SING; Perlis, Kaki

Bukit, C.K. Lim H 1607 PSM Collection.

2. Iguanura wallichiana var. major Becc. in J.D. Hooker, Fl. Brit. Ind. 6: 416

(1892); Ridley, Mat. Fl. Mal. Pen. (Monoc) 2: 152 (1907); Fl. Mal. Pen. 5: 14

(1925); Whitmore, Palms of Malaya: 65 (1973); Kiew, Gdns. Bull. 28: 221, 222

(1976).

Synonyms: Geonoma pynaertiana Masters, Gdnrs’ Chronicle XXIII, 1098:

258 f. 90 (1898); I. spectabilis Ridley, J. Straits Branch Roy. Asiat. Soc. 47: 40

(1903); Whitmore, Palms of Malaya. 65 (1973); J. wallichiana ssp. wallichiana var.

major (Becc.) Kiew synon. noy., Gdns. Bull. 28: 221, 222 (1976).

The variety was first recognised by Beccari as being more robust and having

larger entire leaves. The syntypes cited by J.D. Hooker, basing on Beccari’s

manuscript, were collected between 1880 and 1888, from the Gopeng area: (King’s

Collector Nos. 431, 8227). The collection notes indicated that the fronds were “4-

6ft long, sometimes longer”. The similarity with /. spectabilis is unmistakable, and

indeed Ridley himself stated that “Some specimens of King’s Collection, viz. 431

and 8227, described as the var. major Becc. in the Flora of British India are

apparently either large forms of I. geonomaeformis (sic.) or I. spectabilis Ridley”

(Materials: 152, 1907). By taxonomic precedence, both Geonoma pynaertiana and

I. spectabilis are thus synonyms. Kiew’s I. wallichiana ssp. wallichiana var. major

also becomes a new synonym within this revision, which dispenses with the

subspecies rank by reinstating /. wallichiana and I. geonomiformis. Her description

of the variant is conservative in dimensions, and perhaps Ridley’s notes on /.

spectabilis provides a more impressive account. Ridley first saw the “superb palm

I. wallichiana var. major: Leaf large and entire, with erect robust inflorescence.

Plate

known to Malays as the Teruno” at Bruas where it had been collected by Curtis,

who sent it to Dr Masters. He later saw even more robust specimens on Hermitage

Hill, a foothill of G. Bubu from where the early ascents by Swettenham and

others were made.

In 1937, Furtado collected specimens from the Gopeng area, which had

divided leaves, “5-7ft. long”. The variety is similar to 1. wallichiana var. wallichiana,

but has robust stems to 4m, 3cm or more in diameter; the lamina is usually entire

(rarely divided), cuneate or oblong, strongly ribbed, to 2m or more in length;

characteristically, the inflorescence is ascending erect, thick, paniculate to 8 or

more branches; fruit is globose or ovoid, larger (2cm or more) than for I. wallichiana

var. wallichiana, colour in ripening similar; flowers also similar. Common in G.

Bubu FR, where according to L.G. Saw, an unusual pinnate form has also been

seen.

Distribution: Kedah, Perak.

Type: Perak; Gopeng, King’s Collector 431 (holotype K, isotype FI)

Reference Collections:-

Kedah, G. Bongsu FR, 1940, Corner 35837 SING; Perak, Gopeng, 1937,

Furtado 33089 SING; Bruas, Ridley 8403 SING; C.K. Lim H 1380 KEP; Hermitage

Hill, 1892, Ridley s.n. (Type for I. spectabilis) SING; G. Bubu, 1966, Whitmore

FRI 626 KEP, SING; Keramat Pulai, C.K. Lim H 1268 KEP; C.K. Lim H 424, H

1102, H 1141, H 1142, H 1270 PSM Collection.

3. TIguanura wallichiana var. rosea C.K. Lim var. nov.

A varietate typica fructibus immaturis roseis bene distincta. Typus: Perak:

Belukar Semang, 1992, C.K. Lim H1126 (holotypus KEP)

Caespitose, stem and leaves similar in size and form to /. wallichiana var.

wallichiana, inflorescence also similar, but differs and is distinct in its fruit colour,

pink when immature ripening dark pink or red. Two specific colonies, in Perak

and Kelantan, were found to display the consistent fruit colour distinction. The

drupes are globose or ovoid and similar in size to J. wallichiana, and also in the

carriage and branching of the inflorescence. Both populations studied over a two-

year period are locally abundant, relatively robust, some with stems to 4m in

height, with large leaves to 1.5m, pinnate with broad or narrower leaflets. We

have so far not encountered intermediaries in colour in other locations, and believe

that this character can be used for varietal differentiation.

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Plate 6 I. wallichiana var. rosea: Type specimen: Perak: Belukar Semang, 1992, C.K.Lim

H1126 (KEP)

Plate 7 JI. wallichiana var. rosea: Inflorescence similar to var. wallichiana but with distinctly

pink fruit.

Plate 8 J.asli: Inflorescence with peduncle within sheath, and curved rachillae, and fruit

characteristically candy pink.

Plate 9 JI. diffusa: Type specimen: Perak: G. Tjok, 1884, Scortechini 1189 (FI) (by courtesy

of Erbario Centrale, Florence)

Plate 10 I. diffusa: Specimen from Taman Negara: stem solitary, leaves pinnate with narrow

leaflets; inflorescence interfoliar

The fruit colour distinction has also been displayed by another new species,

I. asli. Although the variation had been considered as “minor local differences”

within /. wallichiana and I. geonomiformis by Kiew (1976: 221), the forms she

mentioned as seen in Taman Negara and elsewhere with pink immature fruit,

might well have been that new species, which has other distinctions in its

inflorescence. I believe that further surveys will locate other consistent populations

of this taxon.

Distribution: Kelantan, Perak. Habitat: hill forest slopes, at. c. 500m.

Type: Perak: Belukar Semang, 1992, C.K. Lim H 1126 (holotype KEP)

Reference Collections:-

Kelantan, Sg. Pergau, C.K. Lim H 1173 PSM Collection; Perak, Belukar

Semang, C.K. Lim H 1192, H 1253, H 1413, H 1437, H 1573, H 1574 PSM Collection.

4. Iguanura diffusa Becc. in J.D. Hooker, Fl. Brit. Ind. 6: 416 (1892); Ridley,

Mat. FI. Mal. Pen. (Monoc) 2: 152 (1907); Whitmore, Palms of Malaya: 65 (1973);

Kiew, Gdns Bull. 28: 221 (1976)

J.D. Hooker’s description (Flora of Br. India, 1892) based on Beccari’s

manuscript, noted that this was “one of the largest of the genus, leaves about 3-

4ft, very numerous leaflets, narrow, one to one-and-a-half inches, inflorescence

filiform, branches eight to ten, branching to second or third orders”. He further

admitted that the specimen was imperfect and “may be a luxuriant state of /.

polymorpha” In fact, the type included a good inflorescence, but too few leaflets,

and had an inscription by Scortechini: “/. polymorpha var.?” Also in Florence, is

another specimen under this name, (presumably accepted by Beccari), collected

by: Ridley in 1895 from Ara Kuda, which consists of one juvenile leaf and is rather

unindicative. In 1907 Ridley himself mentioned the taxon as a variant of /.

wallichiana (Materials: 152). Since then, many others have tried to match collections

to the name, but no specimens have yet been found in or near the type location

which may be G. Hijau (Bukit Larut) rather than G. Ijok near Selama. In Kelantan,

Henderson found a short-stemmed form which he thought to be /. diffusa, which

had a six-branched inflorescence, further divided into second and third order, with

peduncles shorter than in the type specimen.

In 1968, John Dransfield collected a specimen from Taman Negara (686

KEP), noting that the leaves were 4ft in length, with leaflets very narrow, some

unicostate, with short stem to 2ft, with “compound” inflorescence. Whitmore

subsequently determined this as /guwanura diffusa, and indeed our own field

collections in the same area have yielded equivalent samples (eg. H 1542 SING)

which have a range of variations in leaflet widths, some totally unicostate, including

(unusually) the apical pair. Furthermore, within the locally common population

we saw only solitary plants with stems to 1m, and would put this up as a character

in habit, that would further differentiate it from /. wallichiana. The inflorescence

was interfoliar, some shorter in peduncle than others., finely paniculate, with six

to eleven branches, often sub-branching, with fruit similar to 1. wallichiana in

colour and size. The Taman Negara collections appear to fit the original citations,

but because of its geographical disjunction from the type location, an element of

doubt persists for the field-biased taxonomist, and it remains desirable to find it

nearer the type locality.

Distribution: Kelantan, Pahang, Perak. Habitat: low hill forests, near river,

locally common.

Type: Perak: “G. Tjok”, 1884, Scortechini 1189 (holotype FI)

Reference Collections:-

Kelantan, Bukit Batu Papan, 1935, Henderson 29501 SING; Pahang, Kuala

Kenyam, 1968, J. Dransf 686 KEP; 1993, C.K. Lim H 1542 SING, PSM Collection.

Sh Iguanura asli C.K. Lim sp. nov.

Fructibus immaturis roseis; a I. wallichiana et geonomiformi inflorescentiae

interfoliaceae pedunculo brevi vagina folii occulto, rachillis arcuatis differt. Typus:

Pahang: Berkelah FR, 1993, C.K. Lim H 1539 (holotypus SING)

Caespitose, with dominant stems from 1.5-2.5m, leaves marcescent, divided,

similar to /. geonomiformis, variable in numbers and leaflet widths; inflorescence

branching two to nine, rarely sub-branching, often curled interfoliar, with short

tomentose peduncle enclosed in the leaf sheath; fruit globose, 4-8mm, immature

pink ripening brilliant candy pink; flowers and seed not examined.

This is indeed a widespread species found from Terengganu to Johor where

it might have been taken to be a form of /. wallichiana although it is usually within

I. geonomiformis territory. Indeed the habit and leaf variations are visually more

similar to the latter. We later recalled that we had found and photographed

examples in other locations, mainly within Johor, which we had assumed to be J.

wallichiana. There have indeed been earlier collections, and within the Singapore

Herbarium, Holttum’s specimen (9476) is an example. Kiew’s observations on

fruit colour variations in her ssp. malaccensis, (Kiew 1976: 221) is obviously of this

new species. In the Rompin area, the brilliant candy pink fruit often rivals adjacent

Licuala ferruginea not only in colour, but sometimes in size and robustness of the

rachillae and fruit. The inflorescence, although variable, is apparently distinct

Plate 12 I. kelantanensis: Type specimen: Plate 13 I. kelantanensis: Stemless and

Pahang: Gua Musang, 1992, solitary, with interfoliar inflorescence.

L.G.Saw FR137607 (KEP)

from the other taxa, usually curled among the leaf petioles, with the peduncles

mainly enclosed within the leaf sheath.

The inspiration for the name arose out of encountering the aboriginal groups

living near to the first fruiting populations that we saw; it was also determined

within the Year of the Indigenous (Orang Asli) Peoples.

Distribution: Johor, Pahang, Terengganu; not rare, in low hill forests, riverine

swamp.

Type: Pahang: Berkelah FR, 1993, C.K. Lim, H 1539 (holotype SING)

Reference Collections:-

Johor, Kluang, 1922, Holttum 9476 SING; Kahang, C.K. Lim H 1523 PSM

Collection; Jamari, C.K. Lim H 1401 PSM Collection; H 1524 SING; Pahang,

Kedaik, 1991, C.K. Lim H 1000 PSM Collection; H 1041 KEP; Lesong FR, Samsuri

433 SING; Balok 1994, C.K. Lim H 1599 PSM Collection; Gambang, 1992, C.K.

‘Lim H 1229 PSM Collection.

6. Iguanura kelantanensis C.K. Lim sp. nov.

I. wallichianae affinis sed habitu solitario acaulescenti differt. Typus:

Kelantan: Gua Musang, 1992, L.G. Saw FRI 37607 (holotypus KEP)

Differing from caespitose J. wallichiana by being solitary, acaulescent

(sometimes with short stems to 10cm), leaves marcescent, pinnate to 20 pairs of

leaflets, often bicostate, inflorescence branching among petioles, peduncles short

or longer (varying by 20cm or more), rachillae six to nine, curled or not; often

branching to second order, fruit as for 1. wallichiana, white ripening red.

To the east of the Main Range, in Pahang and Kelantan, there appear to be

considerable variation in the forms within the /. wallichiana/geonomiformis complex,

especially in the branching of the inflorescence. While surveying this area, we

first encountered this stemless, solitary /guanura, which had also been collected

by L.G. Saw of FRIM, whose specimen we have chosen as Type. Subsequent

collections display wide variability in leaflet width and divisions, and also in size

and form of rachillae. I would like to credit Saw with the perception of acaulescence

as a distinguishing character, until then not observed in the genus, and quite

unlike the short-stemmed forms of /. geonomiformis. So far the taxon has only

been found in one area, and thus it will be named after the state of Kelantan. In

an adjacent forest, Henderson had collected a specimen (2030/) identified as J.

diffusa (q.v.), which has similar inflorescence, and there may be a relationship

between the two taxa.

Distribution: Kelantan; habitat: hill forests c. 300m

Type: Kelantan: Gua Musang, 1992, L.G. Saw FRI 37607 (holotype KEP)

Reference Collections:-

Kelantan, Gua Musang, 1991, C.K. Lim H 984, 1992, H 1261 PSM Collection.

x Iguanura piahensis C.K. Lim sp. nov.

I. wallichianae affinis sed lamina oblongata integra vel lobata, glabra,

papyracea, laeve sericea differt. Typus: Perak, Piah FR, 1992, C.K. Lim H 1266

(holotypus KEP).

Caespitose, clustering, stems to 2m, leaves marcescent, sometimes entire,

usually divided into two or more pairs of lobes but closely spaced along rachis,

lamina oblong or elliptical, glabrous, texture papery, smooth and silky to the

touch; inflorescence fine and sparsely paniculate four to seven branches or more,

with fruit slightly smaller (c. 6mm in diameter) than /. wallichiana, usually light

green, turning cream to yellow, ripening red.

This is yet another species found so far only in a limited location, where

there are contiguous populations of the more gregarious and common /. wallichiana.

It is an elegant taxon that can quickly be identified once one “gets one’s eye in”,

especially when the oblong entire forms are encountered. Another useful field

indicator is the feel of the relatively thin lamina; the nerves are fine and unobtrusive

and give rise to the smooth and silky glabrous sheen. Young or juvenile leaves of

other taxa can be similar, especially within the 1 polymorpha alliance, and

conversely some old leaves of the new taxon could be coarser. Fruit colours, in

the more evidently yellow-cream (not white) stage are reminiscent of /. parvula

Becc. (q.v.)

Certain herbarium specimens labelled /. wallichiana var. minor display

similar inflorescence, but differ in leaf shape. The specimen collected in 1889 by

Wray (3628, K) from an unknown location in Perak, is an example that I consider

to be this new taxon. Its name is based on the type location which is one of the

home districts of the Temiar tribe.

Distribution: Perak. Habitat: Hill forest 300m alt. Locally not rare.

Type: Perak: Piah FR., 1992, C_K. Lim H.1266 (holotype KEP)

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Plate 14

Lpiahensis: Type specimen: Perak: Piah FR, 1992, C.K.Lim H1266 (KEP)

Plate 16 J. piahensis: Inflorescence with seven rachillae and fruit to 8mm diameter.

Reference Collections:-

Perak, 1889, Wray 3628 K; Piah FR, 1992; C.K. Lim H 454, 1991, H 587, H

1099, 1992, H 1172, H 1202, H 1266, 1993, H 1357 PSM Collection.

8. Iguanura geonomiformis (Griff.) Mart., Hist. Nat. Palm. 3: 229 (1837-1850)

and pl. 178 (1894); Slackia geonomaeformis Griffith, Calcutta J. Nat. Hist. 5: 469

(1845); Palms of British East India, (1850); Miquel Fl. Ind. Bat. 3: 44 (1855); J.D.

Hooker FI. Brit. Ind., 6: 415 (1892); Ridley, Mat. Fl. Pen. (Monoc) 2: 150 (1907);

Fl. Mal. Pen. 5: 13 (1925); Martelli, Nuovo G. bot. ital. 42: 52 (1935); Whitmore,

Palms of Malaya: 63 (1973); ; Kiew, Principes /6: 3-10 (1972); Gdns Bull. 28: 216

(1976).

Synon.: I. malaccensis Becc., Malesia 3: 102 (1886); J.D. Hooker, FI. Brit.

Ind. 6: 416 (1892); Whitmore, Palms of Malaya: 114 (1973); I. geonomiformis var.

malaccensis (Becc.) Ridley, Mat. Fl. Mal. Pen. (Monoc) 2: 150 (1907), Fl. Mal.

Pen. 5: 14 (1925); 1. geonomiformis sub. var. ramosa Ridley, Mat. Fl. Mal. Pen. 2:

151 (1907); I. wallichiana ssp. malaccensis var. malaccensis. (Becc.) Kiew syn. nov.,

Gdns Bull. 28: 220 (1976); I. wallichiana ssp. malaccensis var. elatior Kiew syn.

nov., Gdns Bull. 28: 224 (1976).

The reasons for reinstating 1. geonomiformis have been explained (which

includes the /. wallichiana ssp. malaccensis propositions of Kiew). Griffith’s

description and illustration (Palms of Brit. Ind., 1850: 162, tab. 234) bear out the

characteristics as found in Ayer Panas, Malacca, the bifurcating spadix being

common. Martius using the same collection contributed to some confusion by his

citation: “... spadice saepe simpliciter ramoso”. Incidentally, he also clouded his

description of Areca wallichiana, by noting: “spadicibus simpliciter valde ramosis”,

but described it elsewhere as having 10-15 branching inflorescences. As with J.

wallichiana, which is still extant in its type location, the populations of J.

geonomiformis in and around Malacca can still be referred to.

I. geonomiformis has very much the habit of /. wallichiana - caespitose,

clustering often with dominant stems up to 4m, leaves marcescent entire or pinnate,

some with narrow leaflets, sometimes divaricate; the long, stout and tomentose

inflorescence is often spicate but may fork into parallel tails, sometimes varying in

numbers on the same plant. The fruit is similar to its relative, although in colour

it is often white rather than green, before maturing red. The individuals in Negri

Sembilan are particularly large in stature and fruit size. The taxon has also often

been seen in a solitary form, where basal branching had not yet developed.

J.D. Hooker had listed the synonymous /. malaccensis Becc. as a “closely

allied” species; its type was collected by Kehding in Klang, where the main taxon

is common. Likewise most of the collections cited by Ridley for his varieties came

from the southern part of the Peninsula, including Singapore. The species is by no

a hg

LUUANE RA eeonomartorins

Plate 17 I. geonomiformis: Drawing showing furcating inflorescence; (Tab. 178, Martius,

Historia Naturalis Palmarum Vol. 3. 1837-1850).

means confined to the south, and has been collected on Larut and Bujang Melaka

in Perak and on the eastern side of the Main Range, north to Kelantan, where

curious forms present scope for further study.

The vast collection in the Singapore herbarium displays the wide range of

inflorescence divisions, from the thick robust spikes, often forking up to two or

three, to the “ramose” form, with seven or more tails, sometimes forking again to

second order. Ridley and others devised new appellations to cope, including var.

malaccensis, sub. var. ramosa, and “var. spectabilis” (unpublished). The lamina of

I. geonomiformis varies from entire to lobed and narrowly pinnate, as with /.

wallichiana. Certain historical specimens of /. wallichiana var. minor, eg. King’s

Collector 7996 from Batang Padang, Perak, might well be similar to the form of J.

geonomiformis as found on Bujang Melaka.

To describe the larger entire leaf form as found around Belumut, Kiew

published her var. e/atior, which I include within the main taxon. In the same

vicinity, pinnate forms are also found. Similar variations with entire leaves can be

seen on Bujang Melaka in Perak, and near Jerantut, Pahang. In other locations,

eg. at G. Panti, precocious flowering acaulescent individuals have been seen, but

these are usually caespitose, and may well be basal branches of a mature plant

where the dominant stems have been destroyed. So far we have not encountered

variants of 1. geonomiformis with pink fruit, such as those of I. asli. There may

well be other variants to be described especially from Kelantan, Pahang and

elsewhere, which will require more field collections and samplings of populations.

Distribution: Johor, Kelantan, Malacca, Negeri Sembilan, Pahang, Perak,

Selangor.

Type: Malacca, Ayer Panas, Griffith 6406, 6407 (holotype K)

Reference Collections:-

Johor, Mawai, 1934, Corner 29001, K, SING; G. Blumut, Holttum 10299

(Type for I. wallichiana ssp. malaccensis var. elatior) SING; F.S.P. Ng 98021, KEP;

Gemas, 1966, Palmer 814 SING; Mt. Ophir, 1888, Hullett 751 SING; Ridley 3141

SING; G. Panti, 1892, Ridley s.n. SING; C.K. Lim H 259, H 260, H 914, H1198

PSM Collection; Linggiu, C.K. Lim H 923, H 1023, H 1024, H 1342, H 1345, H

1528 PSM Collection; Ulu Sedili, C.-K. Lim H 925; Labis, C.K. Lim H 1520 PSM

Collection; Lenggor, C.K. Lim H 931, H 1073, H 1590 PSM Collection; Malacca,

Ayer Panas, 1891, Hervey s.n. SING; 1893, Ridley 1583, 1618 SING; Lubok

Kedondong, 1892, Ridley s.n. SING; Negri Sembilan, Perhentian Tinggi, 1890,

Ridley s.n. (Type for I. geonomiformis sub. var. ramosa) SING; Kuala Klawang,

C.K. Lim H 468, H 492, H 1334 PSM Collection; G. Angsi, C.K. Lim H 998 PSM

Collection; Bukit Tampin, 1894, Ridley 189] FI, SING; G. Angsi, 1904, Ridley

11968 FI, SING; Pahang, Tahan Woods, 1891, Ridley s.n. SING; Temerloh, 1913,

Murdoch 174 SING; Kuala Lipis, 1924, Burkill & Haniff 17188 SING; Kuala

Tembeling, 1928, Holttum 20538 SING; Sg. Tahan, 1967, Palmer 12, 13 SING;

Tembeling, 1929, Henderson 21874 SING; Fraser’s Hill, 1929, Holttum 21535 SING;

Sg. Yu, Hardial & Noor 84 K, SING; Taman Negara, Loy FRI 17254 KEP; K,

Temerloh, Hamid 10596 K; Kuala Tenok, Ridley 3138 Fl, SING; Pulau Tawar,

Ridley 3514 FI; Aur Guar, C.K. Lim H 982, H 983, H 1258, H 1479 PSM Collection;

Stong C.K. Lim H 986 PSM Collection; Taman Negara C.K. Lim H 154] PSM

Collection: Jerantut, C.K. Lim H 1256, H 1477 PSM Collection; Perak, Larut,

1881, King’s Collector 1869 CAL, K; 1882, King’s Collector 2994 CAL, FI, K;

1949, Sinclair & Kiah 38783 SING; Ulu Bubong, 1886, King’s Collector 10270,CAL,

K; Scortechini 502 FI; Bujang Melaka, C.K. Lim H 1423 PSM Collection; Selangor,

Klang, 1878, Kehding 108 (Type for 1. malaccensis Becc.) Fl; Kuala Lumpur, 1889,

Ridley s.n. SING; Petaling, 1889, Ridley s.n. SING; Rawang, 1894, Ridley s.n.

SING; Ginting Peras, 1896, Ridley 7887 SING; Semangko Pass, Ridley s.n. SING;

Bukit Kutu, 1896, Ridley 7889 SING; Ulu Langat, 1912, Kloss s.n. K; Genting,

1917, Ridley et alia s.n. K; Gombak, Croat 53311 K; Sg. Lallang, C.K. Lim H 1373

PSM Collection; Awana C.K. Lim H 1535, H 1550 PSM Collection; Singapore,

1890, Ridley 3140 FI, SING; 1891, Ridley 1664 SING; 3139 FI, SING.

9. TIguanura humilis (Kiew) C.K. Lim stat. nov.

Synon.: [. wallichiana ssp. malaccensis var. humilis Kiew syn. novy.:Gdns

Bull. 28: 223, 224 (1976)

I propose to elevate in rank this solitary, short-stemmed relative of /.

geonomiformis with the upward growing roots, as well described by Kiew who

provided a full account of her new variety. The leaves are however, more variable

than as identified and illustrated; other specimens from the type location also

include those that are pinnate. The inflorescence is spicate, and erect;

characteristically, the fruit is globose, white in colour, ripening red. It is interesting

that Furtado, who viewed Corner’s specimen, had also noted that it might be a

new species. It is common and widespread in Terengganu and is possibly also

found in bordering Pahang. It could however be confused with short or stunted

forms of /. geonomiformis which have been seen to flower in acaulescence, as in

G. Panti which are usually caespitose, whereas J. humilis is distinctly solitary in

habit, and mostly not taller than 75cm. In parts of Taman Negara, and in northwest

Pahang, however, solitary forms of J. geonomiformis may be seen, but these may

well be intermediate variants that may relate with /. humilis. Although undivided

leaves are usual especially in the Ulu Setiu area, /. humilis is also often seen with

broad pinnate leaves. The ascending growth of the roots may be correlated with

the swampy habitat, and there may well be upland individuals without this feature.

Distribution: Kelantan, Pahang, Terengganu.

Type: Terengganu, Ulu Bendong, 1935, Corner 30095, (holotype SING,

isotype K)

CARD-INOEY

fe ale a ala =) a 8

CARD-INOEX

ENTERED IN

store

MALAY POmnsuce

Nee oF koakwae

30095

Plate 18 I. humilis: Type specimen: Terengganu: Ulu Bendong, 1935, Corner 30095 (SING)

“ge ra : >... ~

, = oS -_ =

AS ie § GSO ae

aT hee Say SN

Plate 19 I. humilis: Short-stemmed, with entire (or pinnate) leaves, and spicate inflorescence.

Reference Collections:-

Kelantan, Gua Panjang, 1927, Henderson 19502 SING; Terengganu, Ulu

Ayam, Corner 30258 SING; 30259 SING; 30260 SING; Bukit Kajang Corner 30237

SING; Ulu Setiu 1977, J. Drans 5156 K; C.K. Lim H1451 KEP; H 1518 SING;

C.K. Lim H 1299, H 1325, H 1369, H 1542 PSM Collection; Sg. Kemaman, C.K.

Lim H 1491 PSM Collection.

10. Iguanura parvula Becc. in J.D. Hooker, Flora of British India 6: 417 (1892);

Ridley Mat. Fl. Mal. Pen (Monoc) 2: 152 (1907); Whitmore, Palms of Malaya: 63

(1973); Kiew, Gdns Bull. 28: 212 (1976)

From Beccari’s manuscript, J.D. Hooker’s description is pertinent: “Leaves

small, oblong, tip forked, margin undulate; spadix filiform, sparingly branched,

peduncle about as long as the petiole more than half embraced by the spathes”.

He further noted that the leaves were 8"-10" by 3", which is typical, although the

undulate margin might be the result of drying and is not as evident in the live

-plants. He also added that it might be “possibly a very small form of J. polymorpha”

(as annotated by Scortechini), leading others to this assumption. Ridley, however,

erroneously considered it (and also /. diffusa) a form of I. wallichiana (Materials:

i52, 1907)

The taxon is the most diminutive of the genus, caespitose, with

arundinaceous, sinuous stems which often sprout aerial branches. The leaves are

entire (so far no divided forms have been seen) and could be mistaken for young

entire leaves of the other taxa in the /. polymorpha complex, but they are held

stiffly at right angles to the sheath and distinctive in texture. The inflorescence

has been accurately described by Beccari. The peduncles are indeed mainly

enclosed, but are frequently exposed by early abscission, and become infrafoliar.

The fruit is globose, smaller than /. wallichiana, ripening from light green, cream,

yellow (not white) to red.

The aptly named species has been ignored or misunderstood partly because

there was only one historical specimen (in Florence); the evidence in Kew consisted

only of a drawing of the holotype, which is nevertheless a fine sketch.

The type location in Perak is not known, but Furtado recognised the species

as distinct in 1933, when he correctly determined a fine specimen in the Singapore

Herbarium (SFN 21104) collected from the Kedah-Perak border by Haniff in

1928. No other collections appear to have been made until our recent finds in

Kedah, now deposited at KEP, K and SING, which are from one specific area on

forested hill slopes at 200m unfortunately threatened by re-logging activities. We

have also encountered rare individuals of this species at 1000m alt. in a logged

remnant forest in the G. Bintang Forest Reserve. These then are its residual

known distributions.

Plate 20 I. parvula: Type specimen: Perak: c1886,Scortechini s.n.(FI) (by courtesy of

Erbario Centrale Florence)

Distribution: Perak, Kedah.

Type: Perak: c. 1886, Scortechini s.n. (holotype FI).

Reference Collections:-

Kedah, Kuala Ketang, 1928, Haniff 21104 SING; G. Inas, 1991, C.K. Lim H

962 SING; H 975, H 1016, H 1077, 1992, C.K. Lim H 1123 PSM Collection.

11. Iguanura bicornis Becc., Malesia III: 188 (1889), J.D. Hooker, Fl. Brit.

India 4: 417 (1892); Ridley, Mat. Fl. Mal. Pen. (Monoc) 2: 153 (1907); FI. Mal. Pen.

5: 16 (1925); Whitmore, Palms of Malaya: 63, (1973); Kiew, Gdns Bull 28: 204

(1976)

In 1889, Beccari was stimulated by fresh collections sent to him by Dr King

from Calcutta, made by Kunstler and by the late Father Scortechini. The species

named I. bicornis was one of the New Asiatic Palms he listed, citing two specimens:

_the one Scortechini collected at c. 1300m from G. Ijok (more correctly G. Hijau

which is adjacent to Bukit Larut), and the other by Kunstler (6375) also from

Bukit Larut. His description highlights the unique fruit tipped by “two obtuse

unequal bosses” (per J.D. Hooker), which is also described as bigibbous. The

fruit shape varies from oblong to broad and square; ripening from light green to

yellow and brilliant cherry red.

The palmlet is caespitose, clustering often with numerous basal branches,

stems to 2-5m and up to 3-5m (in recently observed robust populations), 6-8mm in

diameter, usually chestnut brown in colour. The leaf sheath is green, abscissing,

leaves entire when young, pinnate with two to eight pairs of trapezoidal leaflets,

flaring distally, varying in the angle of the apical tips, from less than 30 degrees to

more than 90 degrees, typical and similar within the polymorpha complex. The

leaves appear to abscise more readily than others in the complex, giving rise to the

smooth stems, and to the infrafoliar position of the inflorescence, which nevertheless

usually forms within the sheath, as characteristic in that genus.

The taxon in fruit is especially distinctive, and appears to thrive and is quite

abundant in mountain forests usually above 400m to 1500m alt. (it has not been

seen in lowlands), from Bukit Larut northwards to Belum Forest Reserve, Gerik,

also common in Kedah; Weng, and in S. Thailand. Kiew incorrectly included

within this taxon the lowland /. arakudensis Furtado (which was based on a

specimen misidentified by Ridley as Pinanga canina Becc.), whereas it is indubitably

a form of I. polymorpha.

Distribution: Kedah, Perak.

Type: Perak: G. Ijuk, 1886, Scortechini 1188, (holotype FI)

Plate 21 I. bicornis: Inflorescence with

bigibbous fruit, trapezoidal leaflets

a Ss

Plate 22 JI. bicornis: Variations

in leaf form

Plate 23 I. bicornis (top), I. polymorpha (left) and I. mirabilis (right): Fruit and seeds.

Reference Collections:-

Kedah, G. Lang, Kiah 35048 K, SING; G. Inas, Whitmore 4650 KEP; Bukit

Palong, C.K. Lim H 1505, H 1558 PSM Collection; Perak, Larut, King’s Collector

6375 CAL, FI, K, SING; Wray 695 FI; Scortechini s.n. Fl; C.K. Lim H 1391, H

1096 PSM Collection; Kenayat, C.K. Lim H 53, H 58, H 110, H 115, H 117, H 120,

H 617, H 792 PSM Collection; Belum FR, C.K. Lim H 610, H 612, H 1482 PSM

Collection; Bintang Hijau, C.K. Lim H 719, H 754, H 779, H 1043, H 1387 PSM

Collection; Belukar Semang, C.K. Lim H 1124, H 1127, H 1252, H 1365 PSM

Collection.

12. Iguanura corniculata Becc., Malesia III: 187 (1889), J.D. Hooker FI. Brit.

India 6: 417 (1892); Ridley, Mat. Fl. Mal. Pen. (Monoc.) 2: 152 (1907); Fl. Mal.

Pen. 5: 15 (1925). Whitmore, Palms of Malaya: 63 (1973); Kiew, Gdns Bull 28: 204

(1976)

The original specimens from the Selama area are so distinctive that this, the

_ first of the new /guanura Beccari described in 1889, has retained pride of place in

all taxonomic accounts although it had not been collected until recently, and had

been deemed extinct, especially as its type location (at 100-170m alt.) has largely

been cleared for plantations. Fortuitously, our recent field trips have yielded a

newly discovered population, not in Perak but in adjacent Penang, at low altitudes.

As inferred by Whitmore (1973) and Kiew (1976), it is very much a member

of the J. polymorpha alliance. It is caespitose, and is distinguished by its long

spicate inflorescences (22cm in the type specimen), and fruit more pronouncedly

hooked than the narrowest forms in J. polymorpha. The recent collections from

Penang, however, have fruit more similar to /. polymorpha, where the hook is less

pronounced. The plant is smaller, up to 2m in height, with finer trapezoidal

leaflets. It should be noted that some forms of [. polymorpha may be similarly

diminutive, as at Merapoh, where individuals with spicate but shorter inflorescence

can occasionally be found, clearly a variation among those predominantly with

two or three branches. The flowers of the two taxa are observably different in

size, those of I. corniculata being half to two-thirds as large, and having sepals

tinged red at the tips.

So far, the known distribution seems limited to near the Perak-Penang

boundary. The collection by Henderson from Pahang, which he labelled as J.

ferruginea, may need to be corroborated, especially as 1. polymorpha forms with

one to three rachillae have been seen in that state and also elsewhere. Furtado

had determined it to be /. corniculata on the basis of leaflet shape, and may have

introduced (from Kew?) the hooked fruit and leaflet cuttings now in the sachets

on the specimen. Yet another fresh lead might be provided by a recent collection

in 1990 by Kiew and Anthonysamy (RK 2908, UPM) from the Nenggiri area in

Kelantan; the specimen, classified as I. polymorpha, has spicate inflorescence, but

7a 1th

York, Hert, Beh Orkrationnin

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. 4 1LL de Kings Cthetor

Plate 24 I. corniculata: Type specimen: Perak: Selama, Kunstler (King’s Collector)3131 (FI)(by

courtesy of Erbario Centrale, Florence).

Plate 26 I. corniculata; Dwarf clustering palm with long spicate inflorescence; specimen

from Penang.

unfortunately, no fruit were collected or described. [. corniculata remains one of

the rarest endemic species, requiring the fullest conservation measures.

Distribution: Pahang, Perak, Penang.

Type: Perak: Selama, 1882, Kunstler (King’s Collector) 3131, Holotype FI,

isotypes CAL, K.

Reference Collections:-

Pahang, Raub, 1931, Henderson 2509 SING; Penang, Seberang Prai, C.K.

Lim H 1626 PSM Collection; H 1672 KEP.

13. Iguanura polymorpha Becc., Malesia III: 189 (1889), J.D. Hooker, Fl. Brit.

India 6: 417 (1892); Ridley, Mat. Fl. Mal. Pen. (Monoc.) 2: 152 (1907); Fl. Mal.

Pen. 5: 15 (1925); Whitmore, Principes /4: 124 (1970), Palms of Malaya: 64 (1973);

Kiew, Gdns Bull 28: 212 (1976)

Synonyms: /. polymorpha var. canina Beccari, Malesia 3: 190 (1886); 1.

brevipes J.D. Hooker, Fl. Brit. India 6: 416 (1892); 1. ferruginea Ridley, J. Straits

Branch Roy. Asiat. Soc. 47: 40 (1903); Mat. Fl. Mal. Pen. (Monoc) 2: 152 (1907); 7.

arakudensis Furtado, Reprium nov. Spec. Regni veg. 35: 273 (1934).

Like the others with abscissing, pinnate, leaves and trapezoidal leaflets, this

caespitose taxon is indeed appropriately named, with a range of forms that has

given rise to three other proposed species and one variety, now reduced to

synonymy. From extensive field observations, the variations in leaf division, forms

and apical tip angles, are typical of the whole complex (including J/. bicornis etc.)

Beccari’s description of the new species and its variety canina (Scortechini

sn., FI) were based on specimens collected by Scortechini, but the type locations

in Perak are not certain. We would infer that they came from relatively high

altitudes (c. 1200+m), probably from Bukit Larut. Our reexamination of Beccari’s

concept of the taxon has been based on equivalent populations from Cameron

Highlands, some with sinuous stems and leaves irregularly pinnate with broad and

narrow leaflets on the same leaf, a feature which initially seemed distinctive, not

often seen in herbarium samples. Contiguous and intermixed forms can also be

observed with a range of fruit shapes - ovoid, elliptic (but not curved) to the

lightly curved “tooth” (canine) form. The fruit ripens from white to pink and red.

The rachillae vary in number from three to nine, within both forms and are not as

clearly different as suggested by Beccari and others

The /guanura brevipes of J.D. Hooker was also collected from above 1200m

at Bukit Larut, where /. bicornis and I. polymorpha are also found. The holotype

4008 :

bh neena.

io a Pa,

ve (AKIUM MUM PRRAK

Ky Pas tps 3

/ /

f aise ghadatmpad (016

j /

ees f* rEnTA bd i

Ubehi tebe FO Ad! hoy a ee

F bes Lh ftifpte. tie = Diaipiek GE Powel + Malay Pontes

J Aide ? inceel Aiea t a

2 ; Caticonad tig Reed, Pie jeerienhtad GN

ate i

Plate 27 I. polymorpha: Type specimen: Perak: c1886, Scortechini 318b (FI) (by

courtesy of Erbario Centrale, Florence)

Pas

é.

'¥

Plate 28 I. polymorpha: Inflorescence with ripe fruit.

(Kunstler 2029, K) had no fruit, but had flowers which, as described, are

indistinguishable from the others in the complex. The spadix, with three to four

branches, were indeed interfoliar, but it is not certain if any remnant infrafoliar

ones had fallen off. The leaf sheath forming the crownshaft were certainly of the

abscissing type, and although Hooker referred to the slender stem, he did not

indicate whether the taxon was solitary or caespitose, but noted that it was 2-S5m

in height. Amongst abscissing /guanura taxa, the inflorescences can be within or

below the sheaths, sometimes both; leaf fall varies with moisture conditions and

persistence of the nodal grip of the peduncle, and the size and weight of the

inflorescence would also have a bearing. The mixed conditions can often be seen

in the field. Practically all the subsequent herbarium specimens we have seen

labelled as /. brevipes could fit within 1. polymorpha - and some within J. bicornis

(one actually had its typical fruit!).

By the time Ridley published Flora of the Malay Peninsula in 1925, he had

overlooked many editorial errors and credited his /. ferruginea to Beccari, also

providing it with the description of its Licuala namesake in the same volume!

When he first published the new taxon in 1903, he noted that it had the habit of /.

_ polymorpha, and distinguished it on the basis of its “stouter inflorescence on a

longer peduncle and covered with red wool, the larger flowers, and curved cylindric

fruit”. His syntypes were from “Thaiping Hills common from 2000ft upwards”

(Fox 10684, and Ridley s.n.) and from Bujang Melaka (Curtis 3164). In “Materials”

(1907) he characterised the inflorescence as being stout and having nine to eleven

branches, sometimes branched again, and re-emphasised the furry indumentum.

In effect by citing the characteristics of his 1891 specimen, he rendered it

synonymous within /. polymorpha. indumentum on rachillae and leaf parts is,

however, not uncommon for all /guanura taxa.

The other syntypes displayed inflorescences with two or three rachillae, and

curved seeds. This is a form that is common and widespread, and is found in

many parts of Perak, Kedah, Kelantan, Terengganu and Pahang, mainly at lower

altitudes. /. arakudensis Furtado is almost certainly of this form; its type was

based on Ridley’s misidentification of “Pinanga canina” (Ridley 7027), which had

broad apical leaflets and immature three-branched inflorescence. It would have

been tempting to erect a distinct taxon, at least at varietal rank, for /. ferruginea

based on the trident inflorescence and the curved seeds which are more slender

than in var. canina samples. It is now evident that curved and ovoid fruit can be

found within the same taxon, with the “canine” shapes more common. As observed

by Beccari and others, the drupes form eccentrically from the basal stylar remains

which can be on the concave or the convex side. Inflorescence branching and size

are also variable. This was further resolved by the collection of fresh specimens

(from north. Perak) from taller, clustering, robust forms with stems up to 3.5m,

larger leaves (72cm x 40cm), and gigantic inflorescences, variable from three or

four to nine branches, sub-branching sometimes to 16. The longest rachillae

measured over 78cm, including a peduncle of 33cm (vide. C.K. Lim H 1671).

Plate 29 I. polymorpha: Robust form with two to nine branched inflorescence:

specimen from Belum, Upper Perak.

Plate 30 I. polymorpha: Leaflets variable, inflorescence with ovoid or curved fruit.

In summary, the concept of the aptly named /. polymorpha would include a

wide range of variations. Its habitat ranges from sea level to 1500m or more. In

habit it is always caespitose, stilt rooted, with fine or stout stems, 1m to 3.5m in

height, leaves always abscissing, trapezoidal, variable in divisions, leaflet widths

and apical tip angles, with short or long peduncles and inflorescences fine or stout,

with or without ferrugineous indumentum, with three (rarely one or two) to nine

branches, sometimes branching to second order with up to 18 or more branches,

with fruit ovoid to curved, ripening from white to pink and red.

Distribution: Kelantan, Pahang, Perak, Terengganu:

Type: Perak, c. 1886, Scortechini 318b, (holotype FI)

Reference Collections:-

Kelantan, Sg. Bring, 1990, Kiew 2808 UPM; Kota Bharu, Gwynne-Vaughan

560 CAL, K; Sg. Kerteh, Nur 12052 K, SING; Jeli FR, Chelliah FRI 6528 KEP, K;

1911, Anderson 165, 173 SING; 1899, Fox 162, 10684 SING; Burkill & Haniff

12715 SING; Barnard CF 29 SING; Ulu Sat, C.K. Lim H 1164, 1509 PSM

- Collection; Sg. Mekong C.K. Lim H 1302 KEP; Loyjing, C.K. Lim H 1247, H 1264,

H 1347, H 1630 PSM Collection; Pahang, Sg. Gasoh, Jaamar 28264 SING; Merapoh,

C.K. Lim H 1543 PSM, H 1576 PSM Collection; Penang, Ara Kuda, Ridley 7027

(Type for 1. arakudensis) SING; Perak, Scortechini s.n. (Type for I. polymorpha

var. canina), FI; Taiping Hills, 1881, King’s Collector 2029 (Type for I. brevipes)

K; 1891, Ridley s.n. (Type for I. ferruginea) SING; Larut, 1899, Fox 10684 SING;

Ridley 3157 Fl, SING; Wray 714 FI, Ridley 11405 CAL, SING; Curtis 2078 SING;

G. Batu Puteh, Wray 396 SING; Bujang Melaka, Curtis 3164 (Syntype I. ferruginea)

SING; Ridley 9803 FI, SING; Tapah, Avé 132 K; Tambun, Burkill 6299 SING;

Padang Rengas, Burkill & Haniff 13577 SING; Kati, C.K. Lim H 1109, H 1202

PSM Collection; Nenering C.K. Lim H 1411 PSM Collection; Belum FR, C.K.

Lim H 1132 PSM Collection; C.K. Lim H 1671 KEP, K; Kroh, Furtado s.n. SING;

Terengganu, Kg. Tok Dor C.K. Lim H 1156, H 1440 PSM Collection; Dungun,

C.K. Lim H 1169 PSM Collection; G. Tebu FR, Sinclair & Kiah 40821 K, SING;

C.K. Lim H 1416 PSM Collection; Besut, J. Drans. 6504 K; Sg. Tong FR, Meier &

Yong, KEP 94780 KEP, K,

14. _Iguanura belumensis C.K. Lim sp. nov.

I. polymorphae affinis sed habitu solitario, inflorescentiis rachillis 10-14,

distincta. Typus: Perak, Belum FR, 1992, C.K. Lim H 1281 (holotypus KEP).

Solitary, stilt-rooted, stem grey or brown, erect to 4m or more, to 1.5cm

diam., leaves abscissing, nine or more in crown, pinnate three to five pairs leaflets,

trapezoidal (size and variations similar to J. polymorpha), leaf sheath brown,

internode 1-1.4cm, inflorescence often numerous, inter or infrafoliar, finely

4 i ‘ s

fale .

test

’

—

Plate 32 I. belumensis: Type specimen: Perak: Belum FR, 1992, C.K.Lim H1281 (KEP)

Plate 33 I. perdana: Inflorescence interfoliar, with rachillae thick and long.

paniculate with many branches, 10-14, often sub-branching, usually profuse in

flower or fruit; fruit ovoid (also known to be curved), white, pink tinged, ripening

red; seed not ridged.

Found on hill forest slopes at alt. c. 800m, not seen on lower locations, its

solitary habit and more robust erect stem sets it apart, as does the profuse

inflorescence (though sometimes sparse in poor soil conditions), flowering from

lm to 4m or more. Not uncommon but endangered by logging along the E.W.

Highway, it has so far been encountered only in isolation from other /guanura

taxa, although /. bicornis, I. wallichiana and I. geonomiformis have been seen at

similar altitude. In the Belum FR, other sympatric palm species include Pinanga

subintegra var., P. simplicifrons, P. perakensis, P. malaiana, Nenga macrocarpa

and Johannesteijsmannia altifrons. In that area, visited over four years, I.

polymorpha has not been found at this altitude, although to the east and west, at

lower levels, that taxon abounds, and also to the north (probably extending into

Thailand), where robust colonies of the clustering relative are dominant.

On G. Bubu, a specimen collected in 1972 at 1000m (Evans 722, UPM)

displayed curved fruit, but in another (FSP Ng FRI 6134) they were ovoid. Like

others, the fine specimen with robust stems from Bujang Melaka (Shah 3369) was

thought to be /. polymorpha. The epithet belumensis was chosen for the locality

in which we found it in 1989, and used (ined.), for purposes of herbarium

depositions; the choice becomes especially appropriate in the light of more recent

national conservation efforts in the type area.

Distribution: Perak.

Type: Perak: Belum FR, 1992, C.K. Lim H 1281 (holotype KEP)

Reference Collections:-

Perak, Bujang Melaka, 1975, Mohd. Shah 3369 SING; G. Bubu, 1966,

Whitmore FRI 633 KEP, SING; F.S.P. Ng FRI 6134 KEP; Evans 722 UPM; Belum

FR; CK. Lim H 064, A 122, H 531, H 606, H 621, H 787, H 800, H 991, H 1128, H

1153, H 1163, H 1195, H 1243, H 1277, H 1306, H 1438, H 1466, H 1468, H 1472

PSM Collection.

15. Iguanura perdana C.K. Lim sp. nov.

Affinis I. bicorni sed habitu solitario raro caespitoso; elatior, erecta ad 4.5m,

laminis majoribus inflorescentiis robustis fructa non bicorni, excentrico apico plano

bene distincta. Typus: Perak: Kroh, 1992, C.K. Lim, H 1125 (holotypus KEP)

Plate 34 I. perdana: Type specimen: Perak: Kroh,1992, C.K.Lim H1125 (KEP)

Plate 35 I. perdana: Leaves large to 100cm, pinnate forms with trapezoidal or parallel-sided

leaflets.

Usually solitary, rarely with basal branches, stilt-rooted, stem greyish brown,

erect, 1.8cm diam., robust, erect, to 4.5m; internodes 4-5cm, leaves abscissing,

large 100cm x 50cm, pinnate with two forms of leaflets, few (six pairs) broadly

flaring distally or numerous (15-17 pairs) narrowly parallel-sided, petiole 20cm,

leaf sheath 29cm, green with brown indumentum; often persisting in inflorescence

thereby remaining interfoliar, rachillae stout, succulent, nine to ten branches, to

45cm long with 30cm peduncle; flowers widely spaced, relatively large; fruit (only

one seen to date), unripe yellow, pink, like bicornis but eccentric, with flat top,

not two-lobed.

This tantalising taxon needs to be described and recorded even if somewhat

incompletely, in view of its rarity; we have only found less than ten plants in one

threatened locality. Although it is within a Forest Reserve, adjacent felling and

land schemes may affect its survival. It grows sympatrically, with the common /.

bicornis and the new I. wallichiana var. rosea. The very large leaves are a third to

twice the size of the adjacent /. bicornis, usually trapezoidal, but another pinnate

form has been seen with parallel-sided leaflets. The inflorescences are signally

different, lingering (perhaps because of size) among the leaf sheaths, which are of

the abscissing type, leaving clean scars, but are more fibrous, and appear to persist

in support of the stout but gangling rachillae. The unique fruit collected was

somewhat like a dirty tooth, irregular but not bigibbous. This sampling is

incomplete, but other characteristics set this taxon apart not only from J. bicornis

and J. polymorpha but also from J. belumensis.

The population is certainly not sterile, as seedlings and the one

tantalisingly solitary fruit collected prove. It is indeed an imposing erect taxon,

and the epithet, which means prime (in Malay), alludes at attributes which a

leader primus inter pares, should have.

Distribution: Perak; damp hill slopes, alt. 500m.

Type: Perak: Kroh, 1992, C.K. Lim H 1125, (holotype KEP)

Reference Collections:-

Perak, Kroh, H 1507 SING; C.K. Lim H 1254, H 1366, H 1412, H 1572 PSM

Collection.

16. Iguanura mirabilis C.K. Lim sp. nov.

A ceteris speciebus malaysianis foliis integris valde plicatis, pedunculo brevi,

fructibus curvatis differt. Typus: Terengganu: Ulu Setiu, 1993, C.K. Lim H 1448

(holotypus KEP).

Plate 37 I. mirabilis: Rachillae (three to nine branched) with purple indumentum; fruit curved.

Caespitose, stilt-rooted, with basal branches, dominant stems to 3m, 1-2cm

diam., internodes 2-3cm, leaves eight to ten at crown, held stiffly with short petioles

3cm or less, abscissing, entire, lamina strongly plicate, 85cm x 28cm, with deep

(23cm) apical cleft, often with blunt truncated apical edges sometimes attenuated;

leaf sheath 14cm; inflorescence among or below sheath, stout, horizontal or erect,

with purple indumentum, 5 or more branches sometimes sub-branching, rachillae

to 15cm or more, 4-6mm thick peduncle up to 10cm, 10mm x 5mm, often within

sheath; buds prominent pink, with red indumentum, male flowers large 5mm wide

when open, with pinkish sepals not examined in detail. Fruit curved (like /.

polymorpha, also in size), white, ripening cherry red; seed not ridged.

This is indeed a spectacular new species, quite unlike any others within the

genus in Peninsular Malaysia. The plicate leaves are stiff, with deep apical clefts,

and often blunt at the top edges (oar-shaped), sometimes with extended tips. The

leaves abciss cleanly and are entire. The taxon is clearly distinct from the

polymorpha complex, although the curved fruit resemble those of /. polymorpha,

and has been placed (together with /. parvula) for convenience with the trapezoidal

leaflet members, under the abscissing group.

A small specimen with inflorescence had been collected earlier from the

same area but remained unidentified at Kew (J.Dransfield 5145); but no other

collections had previously been deposited in KEP or SING. In Sarawak, two

species (J. sanderiana and I. elegans) may seem to have affinities, their leaves also

being plicate and entire (although lobed forms have recently been found with

variable apical clefts); however, these are solitary in habit, and the inflorescence

and fruit are quite different, globose, with seeds that are ribbed or ridged, features

not observed in Peninsular taxa of Jguanura. The new species has been a wonderful

culmination for this stage of our palm studies, and the name /. mirabilis is surely

appropriate.

Distribution: Terengganu, where it is limited but not uncommon in particular

flood-prone swamp forests, sharing the habitat with /. humilis.

Type: Terengganu: Ulu Setiu, 1993, C.K. Lim H 1448 (holotype KEP)

Reference Collections:-

Terengganu, Ulu Setiu, J. Dransfield 5145 K; H 1518 SING; C.K. Lim H

1300, H 1379 KEP; H 1326, H 1418, H 1474, H 1475 PSM Collection.

Plate 38 I. mirabilis: Caespitose, leaves entire, strongly plicate.

Index of Illustrations and photographs

All photographs are the copyright of the Singapore Gardens’ Bulletin and may not be reproduced

without written permission.

nn & W

Page

I. wallichiana yar. major: Watercolour by Charles De Alwis titled “Jguanura spectabilis

Ridley” (by courtesy of National Parks Board and Singapore Botanic Gardens) ...............:.:00++ 3)

I. geonomiformis: Watercolour by Charles De Alwis titled “/guanura geonomiformis” (by

courtesy of National Parks Board and Singapore Botanic GardeMs) ..00.........c.ccccscceceeseseseeseeeeseseeeees 4

I. wallichiana var. wallichiana: Leaves in lobed and pinnate forms. ..............c.cecceseseseseseeeeseeeeeee 19

I. wallichiana var. wallichiana: Characteristic inflorescence, paniculate, to 90cm. 0.0.0.0... 19

I. wallichiana var. major: Leaf \arge and entire, with erect robust inflorescence. ...0...........0.0.- 22

I. wallichiana var. rosea: Type specimen: Perak: Belukar Semang, 1992, C.K.Lim H1126

SIETP)) seconde -esidenace Sone = Bocce heeee Bac Oc c-Si REE Dace ey nee SE ne 24

I. wallichiana var. rosea: Inflorescence similar to var. wallichiana but with distinctly pink

JETER. cooeordseccebendetine eeteheckcenat eet cule cee ct aap OSE RaR EEE Sees a ie eae ne 25

T.asli: Inflorescence with peduncle within sheath, and curved rachillae, and fruit

“el hie FEI SEIS COZ TT GTEC Ny OT ea a ee et oe 25

I. diffusa: Type specimen: Perak: G. Tjok, 1884, Scortechini 1189 (FI) (by courtesy of Erbario

ORE ere TAN em ELT INCES) pm eee eee crete nace ee na ceve exer ot rac etes oot eat earaacossceceassereceseus cons sevsey suesaveccoussveuivonsee 26

I. diffusa: Specimen from Taman Negara: stem solitary, leaves pinnate with narrow leaflets;

METH Re SCONCE HITILE Hi OMA Nate ee sete te eas coon sc sak Ge Stas eacag chasusane /ecvet Pact sesestectsciusuasneecensursacUesdedvecesenevetesvéscneders 26

I. asli: Type specimen: Pahang: Berkelah FR, 1993, C.K.Lim H1539 (SING). .......cesseeseseseeeees 29

I. kelantanensis: Type specimen: Pahang: Gua Musang, 1992, L.G.Saw FR137607 (KEP) ........ 29

I. kelantanensis: Stemless and solitary, with interfoliar inflorescence. «........-.-::sccscsesceseseseseseeeeeees 29

I piahensis: Type specimen: Perak: Piah FR, 1992, C.K.Lim H1266 (KEP) ........:.ccceseeseseseeeeeeee 32

I piahensis: Leaves silky textured, sometimes undivided, ObIONG. ...............cscscssseseseseeseeeeeeeeeeees 33

I. piahensis: Inflorescence with seven rachillae and fruit to 8mm diameter. ..0....0........eeceeeeeeee 33

I. geonomiformis: Drawing showing furcating inflorescence; (Tab. 178, Martius, Historia

AEE AM SS AMMEATEIENINV Ol SEAN O Sl ODO) bye oc coc igtecstaccoscoassteescscsceucoateonvatevscennecencecvsasdevcsevqereseruvansenveeen, BO)

I. humilis: Type specimen: Terengganu: Ulu Bendong, 1935, Corner 30095 (SING)...........000+- 38

I. humilis: Short-stemmed, with entire (or pinnate) leaves, and spicate inflorescence. .............. 39

I. parvula: Type specimen: Perak: c1886,Scortechini s.n.(FI) (by courtesy of Erbario Centrale

EL rerentytos ong VERCec PEN RCE ect ee ee ee 41

I. bicornis: Inflorescence with bigibbous fruit, trapezoidal leaflets 0c ccecececeseceeeeees 43

Rae Manas aa eA NUT SCM) NE OTN ces nucnsscasdacseentnescorbosneninscasncanccastassdvteanvessocincTeendti seni 43

I. bicornis (top), I. polymorpha (left) and I. mirabilis (right): Fruit and seeds... 43

I. corniculata: Type specimen: Perak: Selama, Kunstler (King’s Collector)3131 (FI)(by

CANMes yal ELD AOMC EN tale. BIOLETMCEN 9. .0.¢cvctscsvecaarvsedtsctoeavscestoecr sossessasdessnsconstdesstsevavesseorsosoosietevs 45

[he ODETTE PLENTE VENRUNL GVETG LS eYe7 0 ee eh ae, En ere 46

I. corniculata: Dwarf clustering palm with long spicate inflorescence; specimen from Penang. .... 46

I. polymorpha: Type specimen: Perak: c1886, Scortechini 318b (FI) (by courtesy of Erbario

CUD TENS, EMO HESAGS) ssirecesnenn sacar Sa ce be rer See EPC en Sr 48

RPO MOTPUATNTOLESCENCE WIE TAP © LUUL be <5 2.ch<n-n<-nervsarsrevseacovnsosszatasasaessvsessussvosususGessccesatsescrsrsedner 49

I. polymorpha: Robust form with two to nine branched inflorescence; specimen from Belum,

De aa iA eT ee See ec Sree nsf snopes chine satan engacab inate vn ged ab daccenensstecshcovis 51

I. polymorpha: Leaflets variable, inflorescence with ovoid or curved fruit. ooo... cceeeereeeeeeee DL

I. belumensis: Solitary, with infrafoliar, multi-branched inflorescence. .............ccccceeseseseeeeeeeeeeee 53

I. belumensis: Type specimen: Perak: Belum FR, 1992, C.K.Lim H1281 (KEP) oc 53

I. perdana: Inflorescence interfoliar, with rachillae thick and long. ..............:eccseseseseeeeeeeeeeeeeeeees 54

I. perdana: Type specimen: Perak: Kroh,1992, C.K.Lim H1125 (KEP), «........cesccsesesssesesesssessssseeees 56

I. perdana: Leaves large to 100cm, pinnate forms with trapezoidal or parallel-sided leaflets. .... 56

Lmirabilis: Type specimen: Terengganu: Ulu Setiu. 1993, C.K.Lim H1448 (KEP). 58

I. mirabilis: Rachillae (three to nine branched) with purple indumentum; fruit curved. ........... 58

PS MILADIS-\@aespitOsenleaves Clie, StLOMP LY PICALC arancsecenecysetccenesesacceaecees seen ate nce ceseeceeeceore 60

Acknowledgements

I would like to mention Dr Ivan Polunin and Dr Francis S.P. Ng for their

initial stimulus in encouraging me to diminish my ignorance of Malaysian flora,

and of Palms in particular. In pursuing this study of Jguanura, I am particularly

grateful to Dr John Dransfield, who not only supplied the Latin for the diagnoses

but also gave unstinted time in advising on the early draft; to Dr Ruth Kiew, Dr

Saw Leng Guan (and his colleagues at KEP) and to Dr Tim Whitmore, who has

been a long-standing, friendly mentor.

Acknowledgements are also due to the herbaria with significant Malaysian

collections: Calcutta, Florence, Kepong, Kew, Singapore, and also Kuching,

Sandakan and Universiti Pertanian Malaysia (UPM), and to their staff: Dr S.C.

Chin, Dr P. Cuccuini, Dr C. Nepi, Mr Tay Eng Pin, Dr Vasu, Dr Wong Khoon

Meng; also to Abang Mohamed Mohtar bin Abang Pawozan, Ali Ibrahim,

S.Anthonysamy, Baya Busu, Kamarudin bin Saleh, Mat Asri bin Ngah Sanah and

Mohamad Shah bin Mohamad Noor.

I would like to extend special thanks to the Minister for Primary Industries

Malaysia, Datuk Seri Dr Lim Keng Yaik, who gave his personal encouragement

to the Palm Search Malaysia project; to Dato’ Mohamad Darus bin Haji Mahmud

the former Director General of the Forestry Department, and his successor Dato’

Ismail bin Awang, and all the many state Forestry Directors, their officers and

rangers who have been helpful, and also to the Wildlife Department and its Director

General, Mohamed Khan bin Momin Khan, and his successor, Musa bin Nordin.

The project benefitted by the collaboration of the Penang Botanical Gardens at

its initial stages, and thanks are due to Mr S. Nadarajah, Ahmad Ismail and

Mohamad Noor Jamalulail. The PSM crew has included Abu Alang, Adam bin

Abdul Rahman, Adnan bin Yusuf, Ahmadi bin Tohari, Chew Sin Huat, Hamid

Busu, Ishak bin Mohamad, Johari Alang, Kasiman bin Rasda, Mohamad bin Sikin,

Mohamed Noh bin Muhamad, Mohamed Yassin bin Mohamad, Othman bin Yusuf,

Razali bin Yusoff, Sani Busu, Sahabuddin bin Ishak, Sari Long, Yusuhairi bin

Yusuff, Zanirros Uda and Zawawi Mat.

For research and secretarial contributions, I would like to thank Majella

Gomes, Betty Teoh, T. Puventhiran and Sani Hussain.

The Palm Search Malaysia project has been privately funded by Heritage

Research Sdn. Bhd., Penang, which will continue to support its work on palms and

other endangered flora, and to maintain and extend its botanical collection.

References

Beccari, O.D. (1886). Nuovi studi sulle palme asiatiche. Malesia 3: 58-149.

Beccari, O.D. (1889). Nuove palme asiatiche. Malesia 3: 187-191.

Blume, C.L. (1838). Bulletin des Sciences Physiques et Naturelles en Néerlande

1: 61-67.

Blume, C.L. (1838-39). Rumphia 2, pt. 18: 105.

Burkill, I.H. (1935). A Dictionary of the Economic Products of the Malay Peninsula

Vol. II (I-Z): 1222. Crown Agents for the Colonies, London.

Dransfield, J. (1969). Palms in the Malayan Forest. Malay. Nat. J. 22: 144-51

Dransfield, J. & T.C. Whitmore. (1969). Palm Hunting in Malaya’s National Park.

Principes 13: 83-98.

Furtado, C.X. (1934). Palmae Malesicae. Fedde. Reprium nov. Spec. Regni vag.

35: 273-83.

Griffith, W. (1844). Calcutta J. Nat. Hist. 5: 445-491.

Griffith, W. (ed. M’Clelland). (1850). Palms of British East India.

Hooker, J.D. (1883). Palmae in Bentham, G. and Hooker, J.D. Genera plantarum

3: 870-948.

Hooker, J.D. (1892). Flora of British India vol. 6: Reeve, London.

Kiew, R. (1972). The Natural History of [guwanura geonomaeformis Martius:

A Malayan undergrowth palm. Principes. 16: 3-10.

Kiew, R. (1976). The Genus /guanura Bl. (Palmae). Gardens’ Bulletin, Singapore.

Vol. 28: 191-230.

Kiew, R. (1978). New species and records of /gwanura (Palmae) from Sarawak

and Thailand. Kew Bulletin Vol. 34: 143-145

Kiew, R. (1989). Utilization of Palms in Peninsular Malaysia. Malay. Nat. J. 42

(1/2): 43-67

Martius, C.F.P. (1837-1850). Historia Naturalis Palmarum Vol. 3.

Ridley, H.N. (1903). New Malayan Palms. J. Straits Branch Roy. Asiat. Soc. 41:

31-51.

Ridley, H.N. (1907). Materials for a Flora of the Malay Peninsula (Monocotyledons)

II: 133-221. Methodist Publishing House, Singapore.

Ridley, H.N. (1925). Flora of the Malay Peninsula. Vol. 5. Reeve, London.

Uhl, N.W. and J. Dransfield. (1987). Genera Palmarum. Allen Press. Kansas,

USA.

Whitmore, T.C. (1970). Taxonomic notes on some Malayan palms. Principes 14:

123-127.

Whitmore, T.C. (1973). Palms of Malaya. Oxford University Press. Kuala Lumpur,

Singapore, London.

Palms in the Farquhar Collection

of Natural History Drawings

Lim Chong Keat

Heritage Research Sdn Bhd

215 Macalister Road, 10450 Penang, Malaysia

Over the period of his stay in Malacca from 1795 to 1818, Major-General

William Farquhar accumulated an extensive collection of drawings of flora and

fauna which he presented to the Royal Asiatic Society in 1827. It was sold in

auction at Sotheby’s in 1994 by the Society, and was subsequently acquired and

magnanimously donated in 1996 to the Singapore Heritage Museum where the

drawings have been given appropriate pride of place. The works by local artists -

said to be Chinese - were not the only items collected or kept by Farquhar; there

were others sent by him to Marquis Wellesley, then Governor-General of India

from 1798-1805. The Wellesley Collection might indeed have offered interesting

comparisons. It consisted of 2660 folios of natural history subjects, including 15

volumes on plants, kept at the India Office Library in London. Other East India

Company botanical drawings have been deposited at Kew.

From my initial scan of the material pertaining to flora, viewing the

marvellous Raffles items and others within the India Office Library, there appear

to be no duplicates or similar drawings. Indeed, there were no records there

pertaining to Farquhar as a collector of natural history drawings. There are believed

to be other items in private ownership from the same artists and vintage; some of

the Farquhar ones have watermarks dated 1796. At the Kew Library, I eventually

came across two palm drawings which were exact and undoubtedly

contemporaneous copies, within the collection of George Findlayson, who had

accompanied Crawfurd to Siam in 1821-1822, and was likely to have had access to

the same artists in Malaya. It can thus be conjectured that Farquhar’s items were

not necessarily exclusive, and that the Chinese artists had a wider clientele; indeed

they would have been practising their trade for quite some time, catering for

European interest in exotic natural history and other subjects.

In China, John Reeves had begun to commission local painters since

1812, and he amassed a collection of some 2000 items now at the British Natural

History Museum. He had supplied the Horticultural Society of London with

drawings by artists whose names have been recorded as “Akut, Akam, Akew and

Asung”. The employment of artists from Canton and Macao went back even

earlier; J. Cunninghame probably used them for the 800 drawings he sent home

between 1698-1703. British interest in botany and the commercial potential of

Asian plants had been concomitant with the activities of the East India Company

since 1698, in common with the Dutch whose botanists since Rumphius had also

been energetic in the region, followed by Blume. Joseph Banks and Daniel Carl

Solander made extensive collections during the Cook voyages.

In India, the work of Roxburgh, Hunter, Wallich, and Jack laid the

foundation for others such as Griffith; within the first half of the 19th century they

provided the earliest taxonomic coverage of the flora of Penang and Malacca.

Indeed Farquhar showed the collection of drawings to Dr. William Jack (who

accompanied Raffles to Penang) in 1818, who was rather critical about their

botanical adequacy. It is more than likely that Jack would have been familiar with

the excellent and meticulous herbarium drawings being amassed by Roxburgh at

the Calcutta Botanic Gardens in India, including the superb watercolours

commissioned by William Hunter in 1802 to represent the flora of Prince of

Wales’s Island, which gave an indication of the high quality expected in botanical

illustration. Indian artists were employed in the main for botanical and zoological

renderings, but it is thought that the Hunter items were by Chinese, probably in

Penang. The botanists and amateurs of the day would have supervised the

illustrations, and often supplied the paper stock and the specimens; they were

often already acquainted with the scientific names given to plant names by Linnaeus

and others.

The Farquhar collection is nevertheless important and significant

historically, and includes many items in the region then not yet scientifically

recorded; some collected and represented from Mount Ophir are believed to be

the original holotypes. There are indeed many illustrations that are artistically

charming and botanically informative - although some are naive and not quite

accurate.They nevertheless provide interesting challenges for the identification of

the species observed and recorded, and also for unravelling their actual location

and extance. With regard to flora it might appear that the items were not all from

Malacca or from Mount Ophir - or from the Malay Peninsula - as the drawing of

the Double Coconut of the Seychelles would infer. This and the other eleven

palm items were not included in the two bound volumes then owned by the Royal

Asiatic Society which were perused by I.H. Burkill and described by him in his

articles in the Gardens’ Bulletin (Vol.XII,1949:404-407; Vol. XIV,1955:530-533).

Much of the flora had remained unidentified until then, but some of his

determinations are due for correction and updating, and have stimulated this

paper - which will be confined mainly to the palm taxa, sixteen within the two

volumes, and a total of 28 in the whole collection.

The 472 drawings were catalogued in 1991 within the total collection of

the Royal Asiatic Society by the indefatigable Raymond Head, who recorded that

the Farquhar items were by then no longer in their original bound volumes. He

provided an index for the set, numbered 016.001-016.472, and deciphered their

inscriptions as best he could, unfortunately without the help of botanists or

zoologists - or someone familiar with Jawi or Romanised Malay. There are therefore

consequential misrenderings that need to be corrected; for example: item 016.469

is listed as “Caryota wiens”, which is clearly Caryota mitis. There are indeed

scribbles difficult to make out, and Jawi does lend itself to various transliterations,

calling for familiarity with the local names of the locality and vintage.

8 eee eee aaaSSaeaQQQS_ aE... eee...

Mild. Fyrhel Rat

Plate 1 Pinanga malaiana: “Wild Betel Nut”; (RAS: 016.048).

In the first volume (016.001-016.054 in Head) inscribed “Medicinal plants,

etc., of Malacca” on the flyleaf, apparently in Farquhar’s hand, Burkill noted 55

items, of which only one was a palm (No. 49; 016.048 in Head), which he identified

not incorrectly as a Pinanga sp. That drawing carried the inscriptions in English

“Wild Betel Nut”, and in Jawi “Pinang Utan”; (there was also an irrelevant

superscription “Calamus Ketang” by pen, and a pencil note: “Rotang”, on the

sheet). It becomes part of the sleuthing exercise to figure out the dates of the

annotations, in English and in Jawi - as to which were original - and their authority;

Burkill suggests that Linley had contributed a note probably in London, and it is

not unlikely that Jack might also have added notes. Local names in Jawi, and the

Rumi transliterations have provided an important basis for botanical identification;

they certainly guided Burkill’s taxonomic determinations. None of the palm species

included in the two volumes had been named by 1827, but some of the other

twelve elsewhere in the Farquhar collection already had established scientific names,

some attributed to Linnaeus, and were so inscribed, and these will be mentioned

later. It is open to conjecture if Farquhar was himself versed in these names, and

would have been responsible for the titles. Although he employed artists who

were Chinese, according to Wallich, the Jawi vernacular names for distinctive

_local flora were obviously by Malay scribes of the vintage and literacy of Munshi

Abdullah. Many items in the vast Indian natural history collections also had Jawi

superscriptions.

In Burkill’s notes on the first bound volume and with reference to the

“Wild Betel Nut”, it is surprising that he did not offer the obvious determination

as the ubiquitous Pinanga malaiana (Mart.) Scheff., published in 1838 (see Plate

1), which has been collected both in Penang and Malacca (and elsewhere throughout

the Peninsula and in Sumatra), and well represented in the Singapore Herbarium.

He had however 54 other non-palm items to deal with.

In the second book (016.055-016.096 in Head), which had originally been

bound in a volume measuring 37 x 42 cm, inscribed “Drawings of Plants from

Malacca Presented by Col. Farquhar Vol. 2”, there are 42 drawings. Burkill notes

that items 17 to 21 were from Mount Ophir. It is not known where the first sixteen

items came from; they were all labelled rattans in Jawi, but as Burkill observed,

item 16(016.070), “rotan sega” is not a rattan or at all a palm but is Flagellaria

indica Linn., and that the term “rotan dini” is sometimes applied to this type of

plant. Apart from item 14 the others are indeed rattans likely to have been found

in the Malacca area, but not exclusively so. The only inscriptions on the pictures

are in Jawi. Drawing from more recent taxonomic information, notably by John

Dransfield (“A Manual of the Rattans of the Malay Peninsula”), an update on

Burkill’s identifications is presented herewith, with corrections in italics:-

1 (016.055). “Rotan perachit” : Daemonorops angustifolia (Griff.) Mart. (1850);

now known as rotan getah.

2 (016.056). “Rotan batu” : Calamus insignis var. insignis Griff. (1844); the local

name is still used.

Plate 2 Calamus speciosissimus:

“Rotan sega badak”; (RAS: 016.058).

Plate 3 Myrialepis paradoxa:*Rotan kertang”; Plate 4 Korthalsia rostrata: “Rotan semut”;

(RAS: 016.059). (RAS: 016.060).

3 (016.057). “Rotan jerenang” : Daemonorops propinqua Becc.in J.D.Hooker

(1893); the local name is also used for three other rattan taxa.

4 (016.058).”Rotan sega badak” : Calamus speciosissimus Furt. (1956) (see Plate

D):

5 (016.059). “Rotan kertang” : Myrialepis paradoxa (Kurz) J. Drans. (synon: M.

scortechinii Becc. 1893); the local name also spelt “kertong” (see Plate 3).

6 (016.060). “Rotan semut” : Korthalsia rostrata Bl. (synon: K. scaphigera Griff.

ex Mart. 1849); the local name applies also to other species of Korthalsia, as

noted by Burkill (see Plate 3).

7 (016.061). “Rotan tunggul” : Plectocomiopsis geminiflora (Griff.) Becc. 1893;

currently called rotan rilang; rotan tunngal is Calamus laevigatus.

8 (016.062). “Rotan manau” : Plectocomia elongata Mart. ex Bl. (synon: P.

griffithii Becc. 1893); usually called rotan mantang; rotan manau is Calamus

manan.

9 (016.063). “Rotan kemandong” : Calamus sp., probably C. speciosissimus, see

item 4; the local name is not known currently.

10 (016,064).”Rotan dahanan” : Korthalsia rigida Bl. (1843); rotan dahan also

applies to other species of Korthalsia, sometimes also called rotan semut.

11 (016.065).”Rotan semambu” : not identifiable from the drawing; probably

Calamus scipionum Lour. 1790, to which the local name refers.

12 (016.066).”Rotan sisir” : not identifiable from the drawing; “Calamus

griffithianus Mart.” as mentioned by Burkill is not known in current checklists;

the local name is also not known, and could have been applied to any finely

pinnated taxon e.g. Calamus exilis Griff..

13 (016.067).”Rotan gelam” : not identifiable from the drawing; not Daemonorops

verticillaris (Griff.) Mart.

14 (016.068). to be discussed below.

15 (016.069).”Rotan getah” : a Daemonorops sp., probably D. angustifolia, from

the local name.

Item 14 is labelled “Rotan pinang-pinang”, but it is obviously not a

rattan, and Burkill suggests that its stem would have been used as a walking-stick

or ‘rotan’ cane. He misidentifies it as Pinanga disticha Bl., which is quite surprising

as that taxon is easily recognisable by leaf and inflorescence and is again well

represented in herbarium collections. It is in fact not a Pinanga but rather an

Iguanura sp. (see Plate 5); the genus was only described in 1838 by Blume based

on a Sumatran type specimen. This particular species with pinnate trapezoidal

leaflets has quite positively not been found in Malacca, or south of Perak - or on

Penang Island, another well-known botanical collection site. Thus unless it proves

Plate 5 Iguanura sp.: “Rotan pinang-pinang”; (RAS: 016.068).

to be an utterly extinct or undiscovered species, it suggests that Farquhar’s artist

had access to collections from areas beyond the British settlements. From recent

field research a new species of single-stemmed Jguanura from the Belum area

(also found in Gunung Bubu, and Bujang Melaka) has similar features.

There are twelve other palm drawings which are not part of the two

volumes described by Burkill; their place in the Farquhar collection can be traced

from the Head index number (shown in brackets). They were apparently bound in

two albums, each with 77 items, and similarly inscribed “Drawings of Plants from

Malacca”, although we are quite certain that not all the plants hail from Malacca.

Six of the palm with known botanical names were annotated accordingly: “Coco

Nut Tree: Cocos nucifera” (016.441), “Pinang Betle Nut Tree - Beetle Tree: Areca

catechu”’ (016.446), “Buah Lontar - Palmyra : Borassus flabelliformis (Lin.)” -

correctly B. flabellifer L.,synon: B. flabelliformis J.A.Murray (016.447),“Tookas:

Caryota mitis”(016.469), “Nipah: Nipa fruticans”(016.422), and “Corma -Dabe,

Elate sylvestris” - which was the Linnaeus name, now Phoenix sylvestris (L.) Roxb.

(probably 016.443, listed uninscribed), the last-mentioned being a species then

already introduced to Malacca. The first four items had no Jawi inscriptions -

suggesting that their naming required no local assistance, whereas the Caryota

’ had a faintly pencilled Jawi corroborative note, and also the Nipah, which was

annotated within the picture as “Pokok Nipah”. Another charming drawing is of

“Sago”: Metroxylon sagu Rottl. (016.470), which had no Jawi but indecipherable

English annotations. There is another labelled only in Jawi with no Rumi

transliteration : a naive but elegant rendering of Arenga pinnata (Wurmb) Merr.

(016.452, uninscribed), yet another introduced species, also commonly found in

kampongs. This is one of the two duplicated in Findlaysonis collection, which is

labelled “Arenga saccherifera Bl.”.

Two further drawings are of “Poko Nibong” : Oncosperma tigillarium

(Jack) Ridl. (016.433), “Sala - The Sala(?) Fruit”: Salacca sp.(016.454) (see Plate

6). The items mentioned so far were from the album dated c.1805-1818, but there

is yet another interesting palm within the other earlier album, the “Palas batoo”:

Licuala longipes Griff. (016.346) (see Plate 7), with inconspicuous Jawi notes. This

is indeed a taxon common in Malacca and on Mount Ophir, where it was later

described by Griffith in 1845.

The final palm to be discussed is from the album containing ten others

mentioned above. It is that famous endemic of the Seychelles - the Coco-de-Mer,

Lodoicea maldivica (Gmel.) Pers. (016.434) (see Plate 8). The drawing bears a

pencilled inscription “Lodoicea Sechellarum”, a name geographically correct,

unfortunately rendered invalid by taxonomic precedence. It would have seemed

most unlikely that there would have been fully grown specimens in Malacca at

that time, and that Farquhar’s artists have drawn it at other locations or even from

illustrations, as it would have been a noted exotic from the sea route to India and

Malaya. A copy of the. same drawing is also in the Findlayson collection at Kew,

and it is therefore not unique as evidence of flora from a particular locality. The

mystery dispelled by Burkill (1935) citing an amazing record by Koenig (see JRAS

Plate 7 Licuala longipes: “Palas batoo”;

(RAS: 016.346).

Plate 8 Lodoicea maldivica:

“Lodoicea Sechellarum”; (RAS: 016.434).

Str. Br. 26, 1894: 104) that a three-year old palm had been seen growing in the

garden of a rich man, Bartolomei de Vents by name, in 1778; this indeed gives us

positive evidence that the Farquhar drawing if made c.1818 of the same palm,

would have been 43 years old, and thus quite fecund - also implying that suitable

floristic mates or progeny were also growing nearby.

Farquhar’s collection is certainly amply fascinating even from viewing a

narrow botanical sector such as palms; the total rich trove of the other natural

history illustrations will undoubtedly yield a wealth of early perceptions in the

depiction of Malesian flora and fauna at the commencement of the 19th century.

Acknowledgments

Yu-Chee Chong, Dr John Dransfield, Dr Ivan Polunin, Singapore Botanic

Gardens, Singapore Heritage Museum, India Office Library, British Museum

(Natural History), Kew Library.

References

Archer, M. (1969). Natural History Drawings in the India Office Library. London.

Burkill, I.H. (1935). A Dictionary of the Economic Products of the Malay Peninsula.

2 vols. London: 1383-1385.

Burkill, IH. (1949). William Farquhar’s Drawings of Malacca Plants, Gardens’

Bulletin XII: 404-407.

Burkill, 1.H. (1955). William Farquhar’s Second Book of Drawings of Malacca

Plants, Gardens Bulletin XIV: 530-533.

Chong, Y.C. (1987). Exploring the East Indies. London.

Dransfield, J. (1979). A Manual of the Rattans of the Malay Peninsula. Kuala

Lumpur.

Gallop, A.T. (1995). Early Views of Indonesia. London.

Griffith, W. (ed. John McClelland). (1850). Palms of British India. Calcutta.

Head, R. (1991). Catalogue of Paintings, Drawings, Engravings and Busts: The

Collection of the Royal Asiatic Society. London.

Hunter, W. (1909). Plants of Prince of Wales Island. Journal Straits Branch R.A.S.

53:52-127.

Jack, W. (1916). Letters to Wallich. Journal Straits Branch R.A.S. 73: 151.

Whitehead, P.J.P. & P.I. Edwards. Chinese Natural History Drawings selected

from the Reeves Collection, British Museum (Natural

History).

Distribution and Abundance of Malayan Trees: Significance

of Family Characteristics for Conservation

James V. LaFrankie

Center for Tropical Forest Science

National Institute of Education

469 Bukit Timah Road

Singapore 1025

Abstract

Taxonomic families of plants that characterize the lowland Malayan rain forest differ from one another nearly

ten-fold in quantitative measures of distribution and abundance. A 50-ha sample of 300,000 trees includes 814

species or fully one-fourth of the Malayan tree-flora. The median adult population size for trees and shrubs is a

linear function of area. From the Pasoh equations, we can calculate the area needed to capture an adult population

of a specific size for a particular fraction of the flora, i.e., for 90% of the Pasoh tree flora to be represented by

more than 200 adults per species will require about 3000 ha of forest. These equations indicate how many species

will have a specific population size within a forest, but not which species. I test the alternative hypotheses that

the large characteristic families of the Malayan forest either do or do not differ more than 10-fold in median

species abundance and species representation. The Pasoh data reject the latter hypothesis. The characteristic

taxonomic families of the lowland forest, e.g., Dipterocarpaceae, Sapotaceae and Burseraceae vary in

representation from 10% of regional species to 60%, the power functions of species- area curves vary nearly

10-fold, and median abundances vary from less than 1 to more than 10 individuals per ha. These findings are

confirmed in part from an analysis of the flora of Singapore which, with regard to representation, illustrates

patterns identical to those at Pasoh. The consequences for conservation are two-fold: (1) general conservation

strategies should not be based on studies of focal families; (2) different taxonomic families of trees and shrubs

will require very different strategies of reserve design for their conservation.

Introduction

Do taxonomic families of plants differ from one another in quantitative

measures of abundance and distribution? Of course, taxonomic families differ

markedly in distribution at the broadest scales of geography: Dipterocarpaceae

are found in Asia, bromeliads in the Neotropics, Proteaceae are predominantly in

Australia and Africa. But within a geographic province, and within a single habitat

such as the Malayan lowland rain forest, do the plant families that characterize

that habitat, such as the Dipterocarpaceae, Myristicaceae, Sapotaceae and

Lauraceae, differ in quantitative features of distribution and abundance?

Plant collectors and field botanists know in a qualitative fashion that some

tropical families tend to show up in characteristic ways. For example, Symington

(1943, pp xii) wrote of the Dipterocarpaceae of the Malay Peninsula: “In these

lowland Dipterocarp forests about 130 species of dipterocarps, representing all

the main groups, occur, but in any one district it is doubtful whether there are

more than 60 species, and the representation of groups may be incomplete. In any

one reserve there may, exceptionally, be as many as 40 species, but between 10

and 30 is the more usual number.” We could add that those few species of the

Dipterocarpaceae that do occur tend be among the most abundant trees in that

forest. Nutmegs are quite different, and most of the 53 Malayan species can be

collected without traveling far.

A quantitative comparison of families along these lines could influence our

ideas about conservation biology in two ways. First, large differences in local

representation and abundance among taxonomic families would indicate that the

numerical loss of species with reducing land area would not be a random loss

from the flora, but would rather be concentrated in certain families. Second, if

large inter-familial differences are found, then efforts either to study biodiversity

or to develop conservation policies, should avoid the use of one taxonomic group

as a proxy representative of overall biodiversity.

This paper compiles the results of a large-scale inventory of the lowland forest

of Malaya (50 ha, 320,000 trees, 814 species) and quantitatively compares those

results with the entire known tree flora of Malaya (13 million ha, 3500 species),

thereby testing the alternative hypotheses that the large characteristic families of

trees either do or do not differ more than 10-fold in median species abundance

and species representation. The findings are then qualitatively compared with the

‘findings of floristic inventory from Singapore.

Methods and study site

The study was made at Pasoh Forest Reserve, Malaysia, which is located at

2° 59' N latitude and 102° 18' W longitude, or about 140 km south-east of Kuala

Lumpur, in the interior portion of the state of Negeri Sembilan amidst a broad

expanse of flat lands and gently rolling ridges that abut the westward side of the

Main Range (Fig. 1). Prior to 1900, this South-Central portion of the Malay

Peninsula comprised nearly 100,000 ha of relatively unbroken forest. The plot is

situated in the last remnant of that forest.

The 50-ha plot is situated in the center of the primary forest reserve, and

comprises 1250 contiguous quadrates each 20 m on a side. Over the course of

three years, a team of roughly 20 people enumerated all the free-standing woody

plants that exceeded | cm diameter at a height of 1.3 meters. The identifications

were documented through collection of about 3000 herbarium specimens that

include either flower or fruit, and through another 4000 specimens of sterile

material. Specimens are stored at KEP and cited in Manokaran et al. (1990),

together with a complete list of species. Further details of the plot survey methods,

including the methods of measuring and identifying trees, are also described in

Manokaran et al. (1990). The base data set has been published in Manokaran et

al. (1993).

The present tabulation of the Malayan tree and shrub flora is based upon a

1990 analysis of the published Tree Flora of Malaya (Whitmore 1973, 1974; Ng

1978, 1989), and further details can be found in Kochummen et al. (1990). In

tabulating the flora, we used the same definition of a tree that was used in the 50-

ha plot, creating a slight discordance with the flora as originally published: we

excluded climbers, epiphytes and stranglers, and scrambling shrubs, and also a

few of the smaller plants in the Rubiaceae. On the other hand, we added a few

shrubs that were not included in the treatment of the Melastomataceae. For the

Dipterocarpaceae, we followed Ashton (1983), and in a few other minor ways our

tabulation departed from the Tree Flora of Malaya series (e.g., we include the tiny

families Dichapetalaceae and Goodeniaceae which were overlooked in the Flora),

and for these reasons, our tabulation differs from those in Whitmore (1973) and

Ng (1988).

Those species in the plot that were new records for the Malay Peninsula (about

2% of the plot flora) were also added to the total Malayan flora. Monocotyledons,

which represent 1-2% of the plot flora, and less than 1% of the trees, were excluded

from the analysis because the regional floristic tally has yet to be completed. A

brief discussion of the distribution of individual trees species based on an earlier

version of this tabulation was presented in Kochummen et al. (1992).

For the purposes of the present analysis, the 50-ha plot is treated as one very

large and robust sample of over 320,000 trees that is representative of the South-

Central lowland forest type, and the results are used as ecological descriptions of

the constituent families. Tests of statistical significance for differences among

families cannot be derived from the plot itself because it is being considered as a

single sample. It would be improper to use the fifty contiguous one-ha plots as

independent samples because they are spatially contiguous, and because the plot

was specifically chosen for its homogeneity.

Percent representation for each family is the number of species represented

by at least one tree in the 50-ha plot divided by the number of species in the Tree

Flora of Malaya. To find the median abundance for each family, we first tally the

number of individuals for each species, then sort the species by family. For each

family the median abundance is the abundance of the species closest to the 50th

percentile. For a family represented in the 50-ha plot by 11 spp, this would be the

6th most abundant species, with 5 species more abundant and 5 species less

abundant. The median is preferable to the mean in that the latter is greatly

influenced by the magnitude of the most abundant species.

Species area curves were prepared for selected families using nested cumulative

plots of 0.1, 1, 10, 30 and SO ha and the entire Malayan flora. An estimate was

made for the each family at the scale of 100,000 ha in the South-Central Peninsula

based on the institutional records of the herbarium in Kepong. This estimate is

distinguished from the actual plot data.

To determine the number of adults of each species per unit area, I estimated

the diameter at first reproduction based on published reports, advice of local

residents, and personal field experience. These were liberal estimates erring toward

smaller diameters. Then I tallied the number of adults for each species within areas

of 1, 2, 4, 8, 16, 32 and 50 ha, and then determined for each area sample the 50th,

75th, and 90th percentile ranking, that is, the adult population that 50%, 75% and

90% of the species exceeded. The samples of 1, 2, 4, 8, and 16 ha were independent

of one another, while the 32 and 50 ha samples necessarily included the smaller

samples. Thus, the samples are, in part, autocorrelated, which precludes a strict

determination of significance for the r* values.

Results and Discussion

1. The sample, covering only 0.0004% of land area of Malaya, includes at

least one individual of 814 species or 25% of the Malayan tree flora. This is much

higher than anticipated and suggests that the usefulness of small reserves for

conservation of trees may have been previously underestimated. Several important

caveats are rather obvious: one individual does not constitute a conserved

_population, and no population is secure if a obligate pollinator has been lost from

the system. On the other hand, a single self-fertile canopy tree could live for a

hundred years or more in relative isolation, and produce thousands of offspring

in a single season.

2. The median adult population size for trees and shrubs is a linear function

of area. The relationship 1s the outcome of three constituent functions: the number

of trees per area, which is linear; the number of species per area, which is a power

function; and the number of individuals per species, which is close to a negative

binomial. That the resulting function is very nearly linear is empirically interesting,

and can be useful for calculating the numbers of adult trees in this lowland rain

forest. The relationship between median adult number (N) and area (A) in ha

follows the equation N = 2.02 + 0.56 *A. The relationship is non-linear over the

first hectare, as species rapidly accumulate, but thereafter conforms closely to the

equation, with an r? value of 0.99 (Fig. 2). The relationship between the 75th

percentile population and area is much less steep, and is fitted to the equation N

= 0.96 + 0.16*A, and again has an r? value of 0.99. The 90th percentile was only

three adults per species at the 50 ha sample, i.e., 10% of the species were

represented by fewer than three adults in 50 ha. The relationship follows the

equation N = —.19 + 0.07 * A, and the r? value is 0.98.

If we choose 200 adults as a safe population level, then this level would be

achieved for half of the species in the Pasoh tree flora in 357 ha; for 75% of the

tree flora in 1250 ha; and for 90% of the tree flora in 2856 ha.

3. These equations indicate how many species might be considered ‘safe’

within a forest, but not which species. Individual taxonomic families differ

markedly in representation of species (Table 1). First, restricting ourselves to

families with more than 5 species in Malaya, representation in the 50- ha plot

ranged from 66% for the Hypericaceae (consisting solely of the genus

Cratoxylum) to 0% for six families. Among the larger families, representation

ranged from 32 out of 53 spp., about 60 % for the Myrsisticaceae to 5 out of 88

spp. or 5% for the Myrsinaceae. The large and characteristic families of the lowland

forest differed in the best-fit power functions from 0.06, for the Anacardiaceae

and Burseraceae, to 0.15 for the Dipterocarpaceae and Rubiaceae. These different

species-area curves are graphically illustrated in Figure 3.

4. Furthermore, median abundance of species varies ten-fold with taxonomic

family. The overall median abundance of individual trees greater than 1 cm dbh

is about 2 per hectare. Thus, about 400 species are represented by more than 2

individuals per hectare and 400 by less. Median species abundances varied among

the larger characteristic rain forest families by nearly 100-fold (Table 2).

Considering first those families with 5 or more species within the plot, the

abundances range from a high of 14 individuals per ha for the Ulmaceae, and 11

individuals per ha for the Dipterocarpaceae, to lows of 0.4 and 0.1 per ha for the

Symplocaceae and Combretaceae respectively.

5. Characteristic taxonomic families of the lowland forest, Dipterocarpaceae,

Rubiaceae, Sapotaceae and Burseraceae will have varying levels of species

representation and numbers of individuals at different spatial scales. Tables 1 and

2 can be usefully combined by taking all Malayan families represented by more

than five species in the 50-ha plot, and ranking them in thirds by percent

representation and by abundance, then cross-tabulating the families into a nine-

cell (Table 3). The families in the upper left are those that are relatively well-

represented in a single forest and also have a relatively high median abundance

in that forest. These families comprise what are widespread and abundant species,

e.g., Burseraceae. Those in the lower left, e.g., Rubiaceae, are represented by

relatively few of the regionally available species, but the species that do occur in

a particular forest are very abundant.

The most obvious question to ask at this point is whether or not these patterns

are truly recurrent and characteristic features of the taxonomic families, or are

they peculiar to Pasoh Forest Reserve. Are there not lowland forests in Malaya

where the patterns are reversed for different families, where the Rubiaceae are

represented by 50% of their species and the Myristicads by 15%?

A natural, but misleading, approach to that question would be to statistically

compare each of the 50 individual one-hectare samples of roughly 6,000 trees each.

Doing this, one finds nearly complete concordance among the fifty samples.

However, this does not answer the question, but rather only tells us that the one-

ha plots, being spatially proximate, are autocorrelated in compositional features,

and represent a relatively homogenous forest, something that we already knew

having specifically chosen this particular forest for its homogeneity. What we want

to know is whether or not these patterns are found recurrently in further samples

around Malaya and Borneo.

There are no other plots comparable to the Pasoh plot where these hypothesis

can be tested. The many line, quadrat, and growth and yield samples made in the

Forest Reserves of Malaya over the past hundred years do not reject the hypothesis

or contradict the findings, but they too heterogenous in method to be used for

quantitative comparisons.

An alternative test is to use verified lists of species compiled for local floras,

and while these offer no insight on abundances, they do represent hard evidence

of a species occurrence. Singapore is one of the better collected tropical locales,

and it’s well-documented list of species describes the original forest cover at a

time long before the great changes brought to the island by economic growth

(Turner et al. 1990). A compilation of percent representation of the characteristic

lowland families represented in on the island (corrected to include only shrubs

and trees) would reject the hypothesis that families display haphazard quantitative

patterns of occurrence. Ulmaceae issrepresented by 7/8 of the Malayan flora,

Myristicaceae by 34/53 or 64%; at the other end, the Dipterocarps are represented

~ in Singapore by 25/156 species or 16%; the Lauraceae by 45/213 or 18%. No

family is represented by more species than might be predicted from Figure 3

for an area of 60,000 ha. For those families of small shrubs that bloom frequently

and which, therefore, should be expected to be thoroughly collected, such as

the Rubiaceae, the representation is roughly as predicted : 89/296 or 30%, and

the Myrsinaceae: 14/88 or 15.9%. This analysis shows that the data from an

ecological sample from Pasoh can successfully predict the percent representation

of taxonomic families in Singapore, something that was not at all intuitively

obvious.

The description of these patterns and the magnitude of the differences among

the families leads to two question: First, what is the ecological basis for these

patterns, and second, what significance do these patterns hold for conservation

policy?

Why should taxonomic families differ from one another in their patterns of

distribution and abundance? Why should species of laurels be patchy in

geographic distribution and sparse in local density, while species of dipterocarps

are patchy in geographic distribution but high in local density, and the

kedendongs (Burseraceae) are both widely distributed and relatively high in local

density?

One of the more obvious factors regulating geographic distribution is the

capacity for wide dispersal. The big trees with low representation, the

Dipterocarps and very big Apocynads, seem to be invariably wind-dispersed.

By contrast, the highest representations are found in trees such as the Anacards,

nutmegs, and kedendongs, which bear large, often arilate seeds in fleshy fruits,

many of which are routinely dispersed by mammals and strong flying birds such

as the hornbills. But we can see that dispersal is not a completely adequate

explanation, for among the species with low representation, along side the

Dipterocarps and Apocynads, are the berry-fruited species of the Rubiaceae and

Myrsinaceae. Those latter two families are comprised chiefly of shrubs and

smaller trees, in complete contrast with the stature of the towering Dipterocarps.

Stature is an interesting factor, in that the highest levels of very local endemism

among Malayan plants seem to invariably occur among small herbaceous genera.

According to Kiew (1990), 50 percent of the species of Malayan Begonia

(Begoniaceae) and 75 percent of 80 species of Malayan Didymocarpus

(Gesneriaceae) are known from single localities. There are no genera of large

trees that have distribution patterns anything like that. The only genera that might

come close are the smaller shrubs in the Rubiaceae and Myrsinaceae. More

complete data on reproductive ecology of these trees and a more thorough

statistical approach may reveal further patterns and correlations.

The consequences of these patterns for conservation are two-fold. First of all,

with regard to the methods of policy formulation, it is clear that the focal-family

approach should be avoided. Choosing one taxon as a proxy representative for

biodiversity is sometimes done explicitly, but more often it is only tacitly done

through the limitations inherent in personal knowledge. Consultants who are

engaged to aid the development of conservation plans often have, as a necessity

of modern research, an intimate taxonomic and ecological knowledge of one or

two families, and they perhaps inevitably draw upon their knowledge in forming

opinions. The results from Pasoh show that this practice is very dangerous. In

deciding upon the size and number of nature reserves, we would reach very

different recommen-dations if we base our study on the Dipterocarpaceae rather

than on, e.g., Myristicaceae.

Second, with regard to a national strategy for conservation, reserves of different

size and number will have a varying success in securing safe populations of species

representative of different families. One large preserve, such as Taman Negara in

the center of the Peninsula will include the overwhelming majority of species in

the Anacardiaceae, Burseraceae, and Myristicaceae. But it will certainly not include

the major portion of Dipterocarps, Rubiaceae or Myrsinaceae. Where the represen-

tative species of these latter families occur, they are extremely abundant and a

small park would include a large population. For these, a greater number of small

parks in more scattered locations about the Peninsula would better serve their

preservation. For the Lauraceae and Sapotaceae, trees with very low abundance

and low representation, it may not be possible preserve their diversity in the wild

through any means short of very large-scale conservation of the landscape. If only

10% of the Malayan Peninsula is retained under mixed forest cover, then for these

families, ex situ conservation may be a priority.

Table 1 Representation of families of Malayan trees in 50-ha plot at Pasoh Forest Reserve, arranged in descending

order of representation; families with more than five species appear before families with less than five

species.

Total Pasoh Total Pasoh

Family Malayan spp. Plot spp. Percent Family Malayan spp. Plot spp. Percent

Hypericaceae 6 4 66.7 Aquifoliaceae 19 2 10.5

Ulmaceae 8 5 62.5 Podocarpaceae 10 1 10.0

Burseraceae 36 22 61.1 Araliaceae 21 2 95

Myristicaceae 53 32 60.4 Saxifragaceae 17 1 5.9

Alangiaceae fl 4 Sel Myrsinaceae 88 5 5)H/

Olacaceae 9 5 55.6 Ericaceae 34 0 0.0

Combretaceae 10 5) 50.0 Actinidiaceae 10 0 0.0

Polygalaceae 21 10 Lythraceae 7 0 0.0

47.6Meliaceae 98 42 42.9 Hamamelidaceae 6 0 0.0

Anacardiaceae 78 32 41.0 Urticaceae 6 0 0.0

Sapindaceae 52 20 38.5

Rosaceae 27 10 37.0 —Families with less than Five Species—

Icacinaceae 19, th 36.8

Ebenaceae 65 23 35.4 Ixonanthaceae 2 2 100.0

Moraceae 72 25 34.7 Asteraceae 1 1 100.0

Violaceae 9 3 3333 Convolvulaceae 1 1 100.0

Annonaceae 133 43 B25 Dichaepetalaceae l 1 100.0

Flacourtiaceae 48 15 31.6 Gnetaceae 1 1 100.0

Dilleniaceae 10 3 30.0 Irvingiaceae 1 1 100.0

Melastomataceae 56 16 28.6 Linaceae 1 100.0

Elaeocarpaceae 28 8 28.6 Santalaceae 1 1 100.0

Leguminosae 102 28 PA fe) Trigoniaceae l 1 100.0

Sterculiaceae 48 13 27.1 Cornaceae 3 2 66.7

Euphorbiaceae 324 87 26.9 Ochnaceae 4 2 50.0

Guttiferae 122 32 26.2 Erythroxylaceae ?) 1 50.0

Boraginaceae 8 2 25.0 Monimiaceae 2 1 50.0

Fagaceae 64 16 25.0 Oxalidaceae >) 2 40.0

Symplocaceae 21 >) 23.8 Capparidaceae 3 1 Ses)

Lecythidaceae 17 4 2359 Crypteroniaceae 3 1 33.3

Myrtaceae 209 49 23.4 Juglandaceae 4 1 25.0

Bombacaceae 22 5 22rT Opiliaceae 4 1 25.0

Lauraceae 213 48 22.5 Staphyleaceae 4 l 25.0

Oleaceae 18 4 22.2 Styracaceae 4 1 25.0

Celastraceae 33 7 21.2 Aristolochiaceae 5 1 20.0

Bignoniaceae 10 2 20.0 Sonneratiaceae 5 0 0

Dipterocarpaceae 156 30 19.2 Caprifoliaceae 4 0 0

Sapotaceae 73 14 19.2 Malvaceae 4 0 0

Tiliaceae 45 8 17.8 Rhamnaceae 4 0 0

Verbenaceae 45 8 17.8 Illiciaceae 3 0 0

Apocynaceae 36 6 16.7 Nyctaginaceae 3 0 10)

Proteaceae 12 2 16.7 Araucariaceae z 0 0

Sabiaceae 6 l 16.7 Clethraceae 2 0 0

Thymelaeaceae 18 3 16.6 Daphniphyllaceae 2 0 0

Rubiaceae 296 46 Il)-5) Hernandiaceae 2 0 0

Rutaceae 46 6 13.0 Pittosporaceae 2 0 0

Loganiaceae 8 | 12.5 Sarcospermataceae 2 0 0

Vitaceaeae 8 1 125 Aceraceae i 0 0

Rhizophoraceae 26 3 LS Casuarinaceae i 0 0

Simaroubaceae 9 1 11.1 Connaraceae 1 0 0

Theaceae 36 4 Nit! Cunoniaceae 10 0

Magnoliaceae 18 2 11.1 Datiscaceae 1 0 0

Pasoh Median Pasoh Median

Family spp. population Family spp. population

Epacridaceae 1 0 0 Nyssaceae 1 0 0

Goodeniaceae 1 0 0 Pentaphylaceae 1 0 0

Myricaceae 1 0 0 Tetrameristaceae 10 0

Table 2 Median population size of species in the Pasoh 50-ha plot. arranged by family. Median population is in

individuals larger than 1 cm dbh per ha.

Pasoh Median Pasoh Median

Family spp. population Family spp. population

Ulmaceae 5 13.70 Opiliaceae 1 11.21

Dipterocarpaceae 30 10.96 Trigoniastraceae 1 7.34

Ebenaceae 23 7.58 Rhizophoraceae 3 7.08

Myrsinaceae 5 7.30 Santalaceae l 6.26

Burseraceae 22 6.70 Convolvulaceae ] 6.22

Melastomataceae 16 6.66 Proteacaceae 2 Spl

Tiliaceae 8 6.30 Violaceae 3 4.90

Polygalaceae 10 3.63 Irvingiaceae 1 4.62

Annonaceae 43 3.58 Alangiaceae 4 3.90

Rubiaceae 46 3:53 Oxalidaceae 2 3.86

Apocynaceae 6 3.46 Linaceae 1 3.62

Euphorbiaceae 87 3.16 Lecythidaceae 4 2.67

Celastraceae 7 3.06 Asteraceae l 22,

Myrtaceae 49 2.92 Thymeliaceae 3 2.50

Anacardiaceae 32 2.76 Ochnaceae 2 2.40

Rutaceae 6 2.74 Dichaepetalaceae 1 2.06

Sterculiaceae 13 2.68 Aquifoliaceae 2 1.99

Rosaceae 10 2.67 Loganiaceae 1 1.92

Guttiferae 32 2S Oleaceae 4 1.39

Leguminosae 28 2.44 Erythroxalaceae 1 1.10

Fagaceae ‘16 2.40 Dilleniaceae 3 0.80

Icacinaceae 7 2.20 Magnoliaceae 2. 0.80

Meliaceae 42 1.97 Theaceae 0.78

Bombacaceae 5 1.82 Crypteroniaceae 1 0.64

Myristicaceae 32 1.82 Hypericaceae a 0.56

Elaeocarpaceae 8 1.70 Podocarpaceae | 0.54

Flacourtiaceae 15 1.62 Aristilocaceae 1 0.50

Sapotaceae 14 iad! Juglandaceae 1 0.40

Sapindaceae 20 1.14 Monimiaceae 1 0.36

Lauraceae 48 0.96 Boraginaceae 2 0.36

Olacaceae 5 0.90 Cornaceae 2 0.28

Verbenaceae 8 0.84 Araliaceae 2 0.16

Moraceae2 5 0.72 Capparidaceae 1 0.04

Symplocaceae 5 0.40 Vitaceae 1 0.04

Combretaceae 5 0.10 Bignoniaceae 2 0.02

Loganiaceae 1 0.02

—Families with less than five species— Sabiaceae 1 0.02

Staphyleaceae 1 0.02

Ixonanthaceae 2 32.01 Styracaceae 1 0.02

Simaroubaceae 1 17.14 Saxifragaceae 1 0.02

Gnetaceae 1 11.60

ABUNDANCE

Table 3. The major taxonomic families characteristic of the lowland Malaysian rain forest divided by thirds with

regard to their abundance with representationin the 50-ha plot in Pasoh Forest Reserve.

HIGH MEDIUM LOW

Burseraceae (22 spp.) (61.1 %) (6.70 ind.) | Meliaceae (42 spp.) (42.9 %) (1.97 ind.) | Sapindaceae (20 spp.) (38.5 %) (1.14 ind.)

Polygalaceae (10 spp.) (47.6 %) (3.63 ind.) | Anacardiaceae (32 spp.) (41.0 %) (2.76 ind.) | Combretaceae (5 spp.) (50.0 %) (0.10 ind.)

Ulmaceae (5 spp.) (62.5 %) (13.70 ind.) | Myristicaceae (32 spp.) (60.4 %) (1.82 ind.) | Olacaceae (5 spp.) (55.6 %) (0.90 ind.)

Rosaceae (10 spp.) (37.0 %) (2.67 ind.)

Icacinaceae (7 spp.) (36.8 %) (2.20 ind.)

HIGH

Ebenaceae (23 spp.) (35.4 %) (7.58 ind.) | Euphorbiaceae (87 spp.) (26.9 %) (3.16 ind.) | Symplocaceae (5 spp.) (23.8 %) (0.40 ind.)

Annonaceae (43 spp.) (32.3 %) (3.58 ind.) | Myrtaceae (49 spp.) (23.4 %) (2.92 ind.) | Flacourtiaceae (15 spp.) (31.6 %) (1.62 ind.)

Melastomataceae (16 spp.) (28.6 %) (6.66 ind.) | Leguminosae (28 spp.) (27.5 %) (2.44 ind.) | Elaeocarpaceae (8 spp.) (28.6 %) (1.70 ind.)

Fagaceae (16 spp.) (25.0 %) (2.40 ind.) | Moraceae (25 spp.) (34.7 %) (0.72 ind.)

Sterculiaceae (13 spp.) (27.1 %) (2.68 ind.)

Guttiferae (32 spp.) (26.2 %) (2.53 ind.)

REPRESENTATION

MEDIUM

Rubiaceae (46 spp.) (15.5 %) (3.53 ind.) | Celastraceae (7 spp.) (21.2 %) (3.06 ind.) | Sapotaceae —_ (14 spp.) (19.2 %) (1.21 ind.)

Dipterocarpaceae (30 spp.) (19.2 %) (10.96 ind.) | Rutaceae (6 spp.) (13.0 %) (2.74 ind.) | Verbenaceae _—(8 spp.) (17.8 %) (0.84 ind.)

Myrsinaceae (5 spp.) (5.7 %) (7.30 ind.) Bombaceae (5 spp.) (22.7 %) (1.82 ind.)

Tiliaceae (8 spp.) (17.8 %) (6.30 ind.) Lauraceae (48 spp.) (22.5 %) (0.96 ind.)

Apocynaceae (6 spp.) (16.7 %) (3.46 ind.)

LOW

Figure 1 Map of Pasoh and Malaya depicting the number of trees and shrubs with documented occurrence.

(These figures exclude monocotyledons, and differ from the tallies in the Tree Flora of Malaya and the

Pasoh data set in a few other minor respects which are described in the methods.)

Bangkok

Gates EA

m Penhea

Ho Chi Min

Malay Peninsula

13,158,700 ha

3197 spp

Pasoh Forest

50 ha

802 spp

Figure 2 The 50th (median), 75th and 90th percentile of adult trees per species related to area at Pasoh Forest

Reserve, Malaysia.

Adult Population

30 7?

1 va

20— @

4 A

| ev

10-4 an

| x of +

ae Qs ey

Ame

0 == eee a = ae T T T

0 10 20 30 40 50

Area ha

Figure 3 For six characteristic families of the lowland tropical rain forest, the number of species per area, all trees

over | cm dbh, starting at Pasoh Forest Reserve Malaysia. Data points for 100,000 ha is estimated from

records of plant collectors.

Percent of Malayan Species

100 aoe ——

Anacardiaceae

Euphorbiaceae

Dipterocarpaceae

Rubiaceae

| | |

3 4 5 6

Log Area ha

Acknowledgments

This study was presented at the International Symposium on Ecological

Perspectives on Biodiversity at Kyoto University in November 1993, and submitted

as a report to the Director-General of the Forest Research Institute of Malaysia.

The large-scale forest plot at Pasoh Forest Reserve is an ongoing project of

the Malaysian Government, directed by the Forest Research Institute Malaysia

through its Director-General, Datuk Dr. Salleh Mohd. Nor, under the leadership

of N. Manokaran in cooperation with the Center for Tropical Forest Science, Peter

S. Ashton of Harvard University and Stephen P. Hubbell of Princeton University.

The identification of trees in the 50-ha plot was supervised by Mr. K.M.

Kochummen through the support of a Smithsonian Senior Fellowship.

Supplemental funds are very gratefully acknowledged from the following sources:

National Science Foundation (USA), Conservation, Food and Health Foundation,

Inc. (USA); UNESCO-MAB and the continuing support of the Smithsonian

Tropical Research Institute (USA), Barro Colorado Island, Panama.

References

Ashton, P.S. 1983. Dipterocarpaceae. in Flora Malesiana Series I. Vol. 9.

Kiew, R. 1990. Conservation of plants in Malaya. 313-323 in Plant Diversity of

Malesia. (Baas, P., K. Kalkman & R. Geesink, eds.) Kluwer Academic

Publishers. London.

Kochummen, K.M. J.V. LaFrankie and Manokaran, N. 1991. Floristic composition

of Pasoh Forest Reserve, a lowland rain forest in Peninsular Malaysia. Journal

of Tropical Forest Science. 3(1):1-13.

Kochummen, K.M., J.V. LaFrankie and Manokaran, N. 1992. Representation of

Malayan trees and shrubs at Pasoh Forest Reserve. pp: 545-554 in Yap., S. K.

& S. W. Lee (eds.) In Harmony With Nature. Proceedings of an inter-national

symposium on the conservation of biodiversity. The Malayan Nature Society.

Manokaran, N., J.V. LaFrankie, K.M. Kochummen, E.S. Quah, J.E. Klahn, P.S.

Ashton, and S.P. Hubbell. 1990. Methodology for the 50-ha plot at Pasoh Forest

Reserve. Forest Research Institute Malaysia, Research Pamphlet No.104. 69

Pp.

Manokaran, N., J.V. LaFrankie, K.M. Kochummen, E.S. Quah, J.E. Klahn, P.S.

Ashton, and S.P. Hubbell. 1993. Maps and stand tables for the 50-ha research

plot at Pasoh Forest Reserve. Forest Research Institute Malaysia, Research

Data Series, No. 1. 454 pp.

Ng, ES.P. (ed.) 1978. Tree Flora of Malaya. Vol 3. Longmans, London.

— (ed.) 1988. Forest tree biology. pp 102-125 in Key Environments: Malaysia

(Cranbrook, ed.). Pergamon Press, Oxford.

— (ed.) 1989. Tree Flora of Malaya. Vol 4. Longmans, London.

Soepadmo, E. & T. Kira. 1979. Contribution of the IBP-PT Research Project to

the understanding of Malaysian forest ecology. in A New Era in Malaysian

Forestry (S. Manap, ed.)

Symington, C.F. 1943. Foresters’ Manual of Dipterocarps. Malayan Forest

Records 16.

Turner, I.M., K.S. Chua and H.T. Tan 1990. A checklist of the native and

naturalized plants of the Republic of Singapore. Journal of the Singapore

National Academy of Science. 18:58-88.

Whitmore, T. (ed.) 1971. Tree Flora of Malaya. Vol. 1. Longmans, London.

— (ed.) 1973. Tree Flora of Malaya. Vol. 2. Longmans, London.

A Fusarium Wilt (Fusarium oxysporum) of Angsana

(Pterocarpus indicus) in Singapore.

F. R. Sanderson!*, Fong Yok King!, Saiful Anuar', Yik Choi Pheng? and

Ong Keng Ho?

Singapore Botanic Gardens, National Parks Board, Cluny Road, Singapore 259569.

Sembawang Field Experimental Station, Primary Production Department, 16 km Sembawang Road, Singapore

769194.

* Present Address: PNG Oil Palm Research Association, P O Box 36 Alotau, Milne Bay Province, Papua New

Guinea.

Abstract

The Angsana wilt disease affecting Pterocarpus indicus in Singapore and caused by Fusarium oxysporum was

first reported in Malacca in 1870. Following several outbreaks in various parts of the Peninsular Malaysia the disease

was recorded in Singapore in 1914 and by 1919 many of Singapore’s Angsanas had either been killed by the disease

or removed to prevent iis further spread. Sporadic occurrences of the disease occurred around Singapore between

1970 and 1982 resulting in a rapid investigation of the disease and the implementation of control measures. Between

1980 and 1992, more than 800 Angsanas were removed as a consequence of the disease.

Although both F. oxysporum, and F; solani were consistently isolated from infected trees only F. oxysporum

proved to be pathogenic in inoculation experiments.

During a 10 month period, 170 Angsana trees were inspected because they had symptoms similar to the Angsana

wilt disease. Of the 170 trees, 86% (147) were infected with F’ oxysporum, while the other 14% were the result of

lightning strikes. Of the 147 infected trees, 90% had also been struck by lightning and 87% had both lightning and

ambrosia beetle infestations. The remaining 15 trees (10%) which were not struck by lightning were at secondary

infection sites where an adjacent Angsana had already been removed because it was infected with F. oxysporum.

The hypothesis presented here for the life cycle of the Angsana wilt disease is that lightning damage to an

Angsana, provides the stress which attracts the ambrosia beetles. If these beetles are contaminated with F oxysporum

spores, then infection is likely to follow. The secondary spread away from this primary infection site, is by

F. oxysporum which has entered the soil from the infected tree.

Short term control strategies are discussed which include the rapid removal of all lightning damaged trees and

the use of insecticides and fungicides either sprayed or injected to prevent the establishment of new infection sites.

Long term control is anticipated following screening of Angsanas collected from a wide geographical area, and

selection for resistance to F. oxysporum.

Key Words: Ambrosia beetles; Angsana; Angsana wilt; Fusarium oxysporum; Fusarium wilt; injection; lightning;

Platypus parallelus; Pterocarpus indicus; resistance.

1 INTRODUCTION

The wilt disease of Angsana (Pferocarpus indicus) caused by Fusarium

oxysporum was responsible for killing on average about 28 trees a month in

Singapore between 1989 and 1995. The first indication of infection is the yellowing

of leaves on one branch followed by their death. This is followed by the yellowing

of leaves on subsequent branches until the tree is completely killed (Plates 1.1,

1.2), a process which can take either a few weeks or many months. Infection is

confirmed by the presence of darkened xylem vessels within the primary xylem

and the isolation of F oxysporum.

Plate 1.1

An Angsana infected with the Angsana wilt disease. The disease has already killed one of the branches, and a

second branch is starting to show the early signs of infection. The leaves turn bright yellow before turning brown

and falling. This magnificent tree was the last in a avenue of about 10 trees.

Plate 1.2

The same tree five weeks later showing the rapid progress of the disease.

The disease was first reported in Malacca, on the South-west coast of Malaysia,

where between 1870-1880 the disease practically wiped out a magnificent avenue of

trees which adorned the sea shore (Fox, 1910).

Thirty years later the disease started to appear in other areas of Malaysia, where,

between 1906-1910, around 100 trees were killed by the disease on the island of

Penang and at about the same time some Angsanas in Tapah (Perak) were also

killed. Soon after, many trees died in Kuala Kubu , Kuala Lumpur (Selangor) and

in Taiping (Perak) (Bancroft, 1912) .

The disease first appeared on Pulau Brani, an island in the port of Singapore,

during 1914. From there it jumped to Connaught Drive on the waterfront, then a

kilometre inland to Dhoby Ghaut. It subsequently appeared in the grounds of the

Istana where the disease developed in an avenue of Angsanas near the gate to

Orchard Road. The avenue was immediately cut in the hope of restricting the

spread of the disease (Burkill, 1918). A localised strain of this disease is still

endemic in these grounds today (Crowhurst et. al., 1995).

The disease continued inland during early 1919 and by May some trees at the

end of an avenue at Tanglin Barracks, 5 kilometres inland showed sign of Angsana

wilt. Although the disease spread to many other parts of Singapore, the epidemic

was held in check by the rapid removal of any tree showing signs of the disease

(Furtado, 1935b).

Sporadic occurrences of the Angsana wilt disease occurred around Singapore

between the late 1970s and 1982, however, an outbreak which affected 28 trees

in an avenue of Angsanas along Tampines Road, resulted in a rapid investigation

into the disease. This investigation culminated in a report written by John Harden

(1982) who formulated a number of control measures which were implemented.

These included the removal of trees once they were confirmed to be infected, with

the trunks and branches being burnt. The areas around infected trees, and around

trees suspected of being infected, were drenched with fungicides. Field officers

were advised to closely monitor any further spread of the disease and strict

horticultural sanitation was introduced.

With the removal of the infected trees and the intensive soil drenching with

fungicides, only 8 further infections were encountered during the next year and

the spread of the disease again appeared to be under control. The next report of

trees being removed because of Angsana wilt was in October 1988, when two

Angsanas along Collyer Quay were found to be diseased. Since 1988 this latest

epidemic has gathered momentum and over 800 trees have been removed as a

consequence of the disease, by the Parks and Recreation Department’s

Maintenance Division.

During 1990, the Parks and Recreation Department, because of their concern

regarding this disease, organised a seminar and a field trip to familiarise field

officers with this disease. Later in that year a joint application was made by

National Parks Board, Primary Production Department and the Parks and

Recreation Department to the National Science and Technology Board and to SGS

SINGAPORE Pte. Ltd, for funding to study this disease.

2 THE CAUSAL ORGANISM

During the 1982 investigation, the causal organism was tentatively identified

by Professor Gloria Lim and Fong Yok King, and confirmed by the Commonwealth

Mycological Institute, as Fusarium oxysporum. Similar wilt diseases occur on

Pterocarpus angloensis in Africa (Piearce, 1979), and on Albizzia julibrissin

(Pirone, 1988) in the United States. No successful pathogenicity test was conducted

with the fungus at that time.

Fusarium oxysporum is a common pathogen of crop plants causing considerable

economic losses in peas, beans, tomatoes, cotton and bananas. There are also

- examples of F oxysporum being pathogenic to palms where it is of commercial

importance on both oil and date palms (Turner, 1981; Louvet and Toutain, 1973).

It has also been recorded on ornamental palms in Singapore on a number of

occasions by staff of the Plant Health Diagnostic Branch of the Primary Production

Department in Singapore (PPD Disease Records).

2.1 Isolation

During the preliminary investigation a number of Angsanas showing

symptoms of Angsana wilt were sampled and fusaria isolated. The isolates

consisted of what were considered typical Fusarium oxysporum with short

phialides and typical Fusarium solani with long branched phialides (Toussoun

and Nelson, 1968; Booth, 1977; and Burgess et. al., 1988). There was also,

however, a wide range of fusaria covering the entire range from short oxysporum-

like, to long solani-like phialides.

During November and December 1991 thirteen sites consisting of 61 trees

showing symptoms of Angsana wilt were sampled. The dead areas of the bark

were identified by removing the outer bark (Plates 2.1, 2.2, 2.3) using a sharp, 3

cm wood chisel and hammer. The inner bark, together with a thin portion of

underlying wood was removed and taken as the sample. Between 3 and 5 samples

were taken per tree. The first sample was taken at the vertical boundary between

healthy and diseased tissue, with subsequent sampling across the dead tissue to

the opposite boundary. The samples were placed in a plastic bag which was then

labelled.

Plate 2.1

Chopping back the bark

reveals the demarcation

between the healthy beige

tissue and the brown to very

dark brown F. solani infected

dead bark. Between the bark

and the wood are many

pockets of F oxysporum. The

vascular staining of the F

oxysporum infection can be

seen in the small area of

exposed wood at the bottom of

the plate. The holes made by

the ambrosia beetles are

frequently to be found

associated with the infected

tissues.

In the laboratory, the samples were first washed in clean tap water and then for

2 minutes in a 20% sodium hypochlorite solution (1:5 Chlorox® and water), rinsed

in sterile distilled water to remove any excess chlorine then transferred to a 15 cm

glass petri dish containing sterilised moist filter paper, for incubation. The samples

were checked daily for developing fungal growth and spores were transferred to

PDA. (Potato Dextrose Agar).

From the 61 trees sampled, Fusarium oxysporum was isolated from 56, and

Fusarium solani from 45. From 40 of the 56 (71%) trees from which F oxysporum

was isolated, F. solani was also isolated, suggesting that this is a disease complex

involving both fusaria.

For all samples collected later in the project, the inner bark and a thin layer of

the outer wood was removed and discarded. This exposed the primary xylem, which

if infected with F oxysporum, would show the characteristic vascular staining

(Plates 2.4). Where possible samples were collected from areas invaded by the

ambrosia beetles. Such samples of primary xylem usually resulted in pure cultures

of F. oxysporum (Plate 2.5, 2.6).

Plate 2.2

A small Angsana transplated

into contaminated soil has

become infected with the

Angsana wilt disease (a).

Removing the bark from the

buttress and lower trunk has

exposed the discoloration of

the tissues underneath. The

discoloured tissues are

invaded by both F. solani and

F oxysporum (b).

Plate 2.3

Infection starting from the

roots and spreading upwards

into the trunk is a good

indication of secondary

spread.

Plate 2.4

Woody tissue of a diseased

Angsana showing the brown

streaks of the infected xylem

vessels, the presence of

ambrosia beetle holes and

white powdery frass.

2.2 Identification

Both Fusarium oxysporum and Fusarium solani were tentatively identified by

the project team, using the criteria of phialide characteristics, macro-spore size,

colony edge morphology, and culture colour (Table 2.1, Plates 2.7, 2.8), as set out

in the keys of Toussoun and Nelson (1968), Booth (1977), and Burgess et. al. (1988).

Table 2.1 Culture characteristics of Fusarium oxysporum and Fusarium solani

FE. oxysporum

E solani

Culture pigmentation

Top white and fluffy white range of colours from light

brown through yellow to orange.

Bottom turn peach to salmon pink amber to red turning dark brown or

black.

Phialide characteristics

short and simple

long and branch

Macro-conidia size

20 —32u x 3 — 5u

40 — 75u x 4.5 — Ju

Colony edge morphology

irregular with micro-conidia

produced within the agar

smooth uniform hyphae

Plate 2.5

Close up view of the same colonised vessel. Bunches of conidia of F oxysporum on short phialides characteristics

of this fungus, can be seen on the mycelium.

Plate 2.6

An ambrosia beetle coming out from its hole in surface sterilized diseased wood. The white cottony mycelium of

F. oxysporum can be seen growing from the frass thrown out by the insect.

Plate 2.7

Slide made from F oxysporum growing on surface-sterilised diseased wood tissue collected in Singapore. (a)

Chlamydospores, (b) mycelium with short simple phialides.

Plate 2.8

E oxysporum isolated from diseased Angsana tissue collected in Singapore. Sizes of macro-conida range from 18-

28 (a), (b) and micro-conidia 4-144 (c).

For confirmation of their identity, three isolates of each species were sent to

the International Mycological Institute, UK, Sydney University’s Fusarium

Research Laboratory, Australia and the Crown Research Institute, Auckland, New

Zealand during the course of the investigation. The three isolates of F oxysporum

and F. solani represented the range within the population. All three institutes to

which the isolates of F. oxysporum were sent, confirmed their identity, however,

with the F. solani isolates, there was no consensus as to their identity. In two

instances different names were given to the different isolates. This is not surprising

because of the variation between the F. solani isolates. Some isolates of F. solani

were impossible to differentiate from FE oxysporum on phialide characteristics

when viewed on wood, the difference only became obvious because of colony

morphology when the isolates were growing on PDA. A complete range of phialide

length along with a wide range of colony colour was found within the population

of F. solani. It is suggested that when isolates were sent for identification they

were seen as individuals from separate populations, rather than the natural variation

within a sexually active and therefore a segregating population.

As we are not in a position to judicate on the taxonomy of the F’ solani, and to

‘ prevent further confusion in the literature, F solani is retained for this publication.

Fusarium solani was consistently isolated from the dead bark outside the F

oxysporum infected primary wood tissue. It was also consistently isolated from

ambrosia beetles sampled. The status of F so/ani in this disease complex, therefore

warrants further investigation.

2.3 Pathogenicity Test

A root inoculation test was conducted to test the pathogenicity of the two fusaria

isolated. The two fusaria were consistently isolated from the wood material collected

from infected trees.

The method used was that described by Burgess et. al. (1988 ) in the Laboratory

manual for Fusarium Research, and by Piearce (1979).

One hundred and fifteen, well rooted hardwood cuttings of Angsana were removed

from their pots and the soil thoroughly washed from the roots. The roots were then

trimmed to 50% of their original length before the cuttings were placed in a spore

suspension and left in the sun for four hours. The spore suspension was made by

macerating three plates of one-week old fungal lawns in 500 ml of sterile water.

Sixty cuttings were inoculated with Fusarium oxysporum, 45 with F. solani and

10 inoculated with macerated PDA as controls. After four hours the cuttings were

re-potted using the original soil and watered with the fungal lawn suspension. No

watering was done for the next three days, and subsequently every three days.

The first cutting started to wilt after 19 days with the majority of the cuttings

wilting between days 21 and 28. The trial was terminated after 35 days.

As can be seen from the results of the inoculation test (Table 2.2, Plate 2.9) the

only deaths occurred with those hardwood cuttings inoculated with F oxysporum

where 96% of the hardwood cuttings died. The identity of the causal organism was

therefore confirmed as F oxysporum.

Table 2.2 Pathogenicity test of the two fusaria isolated from diseased Angsanas

Number Number Number

inoculated died survived

Plate 2.9

Well rooted hardwood cuttings of Angsana inoculated with Fusarium oxysporum and Fusarium solani. Five

weeks after inoculation, all except 3 cuttings inoculated with F oxysporum showed symptoms of Angsana Wilt

Disease and died eventually (a). Those cuttings which were inoculated with F. solani (b) and those uninoculated

plants (control) (c) remained healthy.

3 EPIDEMIOLOGY

3.1 Lightning

Damage to trees caused by lightning has been the subject of periodic

publications in Singapore (Thomas, 1900; Fox, 1910; Futardo, 1935a). Osmaston

100

(1920: page 120) made the statement “From general observations which are not,

however, based on definite countings, I believe that under existing conditions (in

Asia) about 50% of trees struck (by lightning) survive, and the death of the

remaining 50% is, I believe, mainly brought about by insect and perhaps also

fungal attack. On more than one occasion I have noticed how Platypus biformis

and bark beetles may at once attack a struck tree commencing at first on either

side of the rift in the bark and thence gradually extend their operations completely

round the stem.” Sharples (1933) suggested, that in the case of rubber trees (Hevea

brasiliensis), lightning discharges rendered these trees susceptible to the attack

of parasitic insects or fungi.

The relationship between lightning and insect attack has received considerable

attention (Miller and Keen, 1969; McMullan and Atkins, 1962; Anderson and

Anderson, 1968; Anderson and Hoffard, 1978 and Schmitz and Taylor, 1969).

Taylor (1974 : page 843) comments that “several genera of forest insects

apparently respond to the olfactory attraction of volatile extractives released by

a tree newly ruptured by lightning, and the few initially attacking insects may

create sexual stimuli that trigger a mass attack on the struck tree and its

-neighbours.”

Harden (1982) in his report to the Parks and Recreation Department, suggested

the possible involvement of the ambrosia beetles in the disease cycle of the

Angsana wilt disease. However, such a suggestion posed the enigma that ambrosia

beetles only infest weakened trees. For this reason it was generally assumed that

the ambrosia beetles infested the Angsanas only after they had been weakened by

Angsana wilt.

The puzzle which confronted the project team at the beginning of their

investigation was, if F oxysporum is a soil borne organism, how do new infection

sites develop several kilometres away from an old infection site, and are the

ambrosia beetles, as suggested by Harden, and which are nearly always associated

with the infected trees, part of the disease process.

The Housing Development Board’s tree nursery along University Road, gave

us our first indication that lightning might be associated with the epidemiology

of the disease. This occurred when a tree growing adjacent to four trees which

had been unsuccessfully stem inoculated with F oxysporum, attracted our attention

because of a line of frass which extended from the ground to the tip of a main

branch (Plates 3.1, 3.2). On close inspection the tree exhibited all the symptoms

of Angsana wilt, and F: oxysporum was isolated from the tree. This situation caused

considerable confusion and debate as to why an adjacent tree, and not one of the

inoculated trees, should develop the symptoms of the disease. During the following

weeks, however, obvious symptoms of what we perceived to be lightning damage

began to appear. After three months a 5 - 10 cm strip of bark had peeled off,

revealing the wood from just above the ground level to the top of the main branch.

101

Plate 3.1 & 3.2

White powdery frass on the

Angsana trunk and root flare

indicating the presence of the

ambrosia beetles actively

invading the tree. When this

photograph was taken the

leaves on the affected

branches were drooping and

dull green, suggesting that it

had only been a few days

since the lightning strike. The

tree was the third in a row to

have been affected.

102

This was the area which had originally been the site of the ambrosia beetle

invasion. The disease subsequently spread to two neighbouring trees.

Of the 21 trees investigated during the following five weeks from which F

oxysporum was isolated, 17 of the trees also exhibited symptoms of what we

suspected were caused by a lightning strike. These symptoms included, the rapid

defoliation of one or several branches of the tree, areas of thin outer bark which

had been lifted off in large sheets, and a line of cracked bark down the tree,

defining the path of the lightning. The bark along the cracks often appeared to

have been burnt or finely fragmented. Fifteen of the 17 lightning trees were also

colonised by ambrosia beetles within days of the lightning strikes, with numbers

of 500 - 2,000 beetles/m? being common.

3.1.1 Water Solubility Of Resin

Because there appeared to be an association between lightning damage and

trees infected with Angsana wilt, it was important to determine with certainty,

whether or not a tree had been struck by lighting. To this end we started looking

at ways to confirm lightning strikes on Angsana.

An observation that the red stains on clothes, caused by the resin from

Angsanas, could be washed out by vigorous scrubbing in cold water, but if the

clothes were washed and subsequently ironed, then the stains became permanent

and impossible to remove, led us to investigate the solubility of the Angsana resin.

It was found that the resin from both the healthy Angsana, and those colonised

by F oxysporum and F. solani, although not soluble in organic solvents, was

soluble in water. Conversely, resin from the bark samples suspected of having

lightning damage, were not water soluble.

To determine the temperature at which the resin became insoluble in water,

1.5 g samples of air dried, healthy bark were placed into open glass petri dishes,

and heated in an oven. One petri dish sample of bark was removed at every 10°C

rise in temperature between 70°C and 220°C. A glass petri dish lid was placed

over the petri dish as it was removed from the oven. When these samples were

placed in water, it was found that the resin was water soluble for those samples

which had been removed at a temperatures of 150°C and below, whereas the resin

was no longer water soluble for those samples heated to 160°C and above.

Assuming that the only natural way for the bark to be heated to temperatures

above 160°C is the passage of a very high electric current through the bark, 1.e.

lightning. We had developed a very simple test to determine whether or not an

Angsana has been struck by lightning. The test results can be obtained within five

minutes, with complete repeatability, and thus a very high degree of accuracy.

103

3.1.2 Tree Survey: November 1993 - August 1994

Each week, from November 1993 until August 1994, trees showing symptoms

of Angsana wilt (Plates 1.1, 1.2) were inspected and detailed records made of

lightning symptoms, location, ambrosia beetle infestation, percentage of the canopy

and bark affected, size of tree, and the number of nearby Angsana. Bark samples

were collected from both the suspected lightning damage and adjacent trees. These

were tested in the laboratory for the water solubility of the resin, to determine

whether a tree had or had not been struck by lightning.

During the 10 month period, 170 Angsanas were inspected because they had

symptoms similar to Angsana wilt. Of the 170, 147 (86%), were infected by

F. oxysporum, the remaining 23 (14%) had been struck by lightning but no

infection was detected. Of the 147 infected trees, 132 (90%) had been struck by

lightning and 128 (87%) had both lightning and ambrosia beetle attack.

All the 15 (10%) trees where lightning damage was not recorded, were

secondary infection sites being adjacent to a site where a tree had already been

removed because of the Angsana wilt disease. The time taken for the secondary

infection to move from an infected tree to the adjacent tree varied from 22 days

in the case of two trees in Holland Avenue, to 131 days between two adjacent

trees at the Pioneer Rd/International Rd corner. The mean time taken for the

secondary infection to move between two trees was 73 days (15 observations).

3.1.3 Symptoms Of Lightning Damage

The first symptoms occur at the time of the lightning strike and these are the

physical damages that are associated with the expansion of gases as the liquids in

the bark are heated past their boiling points. In the majority of cases, sheets of

outer bark, usually in the order of 50 x 20 cm, are either lifted off together with

the thicker inner bark or they are completely blown off the tree (Plate 3.4). Small

areas (1 x 5 cm) of inner bark may also be completely disrupted producing an

area of loosely held fibres (Plates 3.5, 3.6).

During the days following the strike it becomes apparent that the tree has

sustained a lightning strike as the leaves dry up, turn a dull green then brown before

falling (Plate 3.7). If the tree is left standing, the affected branches will flush with

new leaves, which may also wilt before reaching full size. In the absence of the

F. oxysporum fungus, death or survival of a lightning struck tree depends on many

factors including the size of the tree and the extent of the lightning strike.

During this period the bark along the path of the strike will start to dry and

crack. It is along the edge of these cracks that the resin appears to have been baked

and it is from this area that samples are collected for testing the water solubility

of the resin. The drying and cracking of the bark continues over the next few

104

Plate 3.4

In the majority of severe

lightning strikes, large sheets of

outer bark get lifted off from

the trunk.

months until the bark falls to the ground and the area of damage is outlined by

the absence of bark (Plate 3.8).

It was common to record more than one tree which had apparently been struck

during the same storm. In one instance a group of five neighbouring trees all

showed similar symptoms (Plate 3.9). It is also suspected that trees which show

symptoms within a few weeks of the first tree dying, might also have been struck

at the same time. This delay has been attributed to the shock being transferred

through the root system (Futardo, 1935a).

3.1.4 Ambrosia Beetles

Dutch elm disease is the best known example of a disease - insect relationships

because of the widespread death of elms in both Europe and the United States.

The insect-pathogen relationship is a passive one, with the fungal spores sticking

105

Plate 3.5

Below the sheets of thin outer

bark which have been separated

from the inner bark, are the

vertical cracks and often small

areas of completely disrupted

bark producing areas of loosely

held fibres.

Plate 3.6

Large sheets of outer bark are

separated from the inner bark

revealing the vertical cracks

and areas of disrupted fibres.

106

Plate 3.8

The area of exposed wood

showing the old ambrosia

tunnels, which three months

earlier had delimited the area of

damage and from which

F. oxysproum had been isolated.

Plate 3.7

An Angsana along Adam Road

showing what appeared to be

the symptoms of Angsana wilt

disease. On close inspection the

trunk showed damage due to a

probable lightning strike. The

leaves turned dull green then

brown before falling. This

process took about 7 days.

107

Plate 3.9

Two of a group of five

Angsanas at a vacant plot of

land on Adam Road, all

showed similar lightning

damage, suggesting that all

five had been struck

simultaneously.

to the insects before the adults leave the diseased tree, then being physically

brushed off in the new feeding tunnels.

Ambrosia larvae on the other hand feed on fungi which are cultivated by the

adults, and the spores of these fungi are actively carried from one tree to another

in special pouches, mycangia, on the thorax of adults. The ambrosia beetles

therefore actively inoculate newly colonised trees with their ambrosia fungi. It is

not surprising therefore that when a plant pathogen becomes involved as part of

this fungal-insect symbiosis that a fungal-insect-disease situation arises. Current

research in New Zealand suggests that an ambrosia beetle (Platypus spp.) might

also be involved as a vector of Dutch Elm Disease in that country (Scott C. pers.

comm. ).

In 1973 Zimmermann, found Fusarium javanicum to be among a range of fungi

colonising the galleries produced by the ambrosia beetle, Xy/ebrus. Appreciating

the significance that some of these fungi might be plant pathogens, he tested them

in the laboratory and found that F javanicum was indeed a pathogen, which caused

a canker on tomato shoots.

The first link between an ambrosia beetle being a vector of a plant disease was

when Kessler (1974) suggested that the apparent symbiosis between a Fusarium

species and ambrosia beetle (Xylosandrus remanus) causes black walnut canker,

108

a link that was confirmed by Weber (1979, 1985). Hara and Beardlsey (1976)

studied the biology of the black twig borer, a severe pest of shrubs and trees,

causing extensive economic damage to coffee and cacao in tropical Africa,

Indonesia and Southern India and to tea in Japan, and found that the eggs were

laid on the ambrosia fungus, F: solani. The symptoms of the attack were necrosis

of leaves and stems, which extended from the entry hole to the terminal shoots of

the branch. These are leaf and stem symptoms characteristic of Fusarium attack

on crop plants such as peas and beans.

In 1978 Anderson and Hoffard demonstrated the link between Fusarium canker

(F: solani), and the ambrosia beetle Xylosandrus germanus and Xylebrus sayi on

tulip poplar trees in Ohio.

There are three reported examples of ambrosia beetles being associated with

diseases of pines. Frederick (1976) provided data which suggested the scolytid

beetles may be important vectors in the transmission of Scieroderris lagerbergii

under specific conditions. Wingfield and Marasas (1980) suggested that the fungus-

insect relationship of Ceratocystis ips and Orthotomicus erosus is an important

- part of a disease complex that results in significant losses in pine plantations. In

1983, the same authors demonstrated the pathogenicity of the root pathogens

Verticicladiella procera and 2 new species and showed that they were carried by

scolytid bark beetles.

Other examples of ambrosia beetles being linked as the vectors of plant

pathogens are: the mortality of the Red Beech (Nothofagus fusca) which is a direct

result of the invasion of the sapwood by a fungal pathogen Sporothix sp. introduced

by the ambrosia beetle Platypus sp. (Faulds ,1977); the sudden death syndrome

of cocoa, caused by Phytophthora palmivora in Papua New Guinea, which Prior,

(1986) linked to the scolytid bark beetles and the ambrosia Platypus sp.. In this

study Prior found that F: solani was also a part of this disease-insect syndrome;

and in 1991 Hijii, et. al. suggested that the mass mortality of oak trees in Japan

was the result of a range of pathogens introduced by the ambrosia Platypus

quercivorus and P. clamus.

In our current study F. solani was consistently isolated from ambrosia beetles

sampled. It was also consistently isolated from the dead discoloured bark outside

the primary xylem infected with F oxysporum. This bark is characteristically wet,

and smells of decay, and is the marker used for locating the F oxysporum infection.

This association of ambrosia beetle, F’ solani and the F oxysporum warrants further

investigation.

Of the 132 infected and lightning damaged trees inspected during the last 10

months of the project, 128 (97%) were infested with ambrosia beetle at the time

of inspection (Table 3.1).

109

Table 3.1

Classification of the 170 = os

r [= = ®

unhealthy trees inspected ol vas| Set Ae Sie eine Tella 1G

F : Sy) 9S) | Bho aee eS lee eels Baka mee co

during the 10 month period = eee ba eee eae lSe waren Se Sultae |S

= ® = 2 =| = ec =]

November 1993 — August 1994. Te SG este pee oe ences Veg. ios So

> cot | Bea, ct ys Selpery e @ | ¥

5 Zi €z oO re || s % | & a

<= = = § 3 — =) 2 “< — 3 = =

aS Sree ny | eee ens ae om [ieee

rion eee ei) Mr ieee) Sirsa (seek oda

ee aka satis erm ee ie P| a a

a é

Lightning damage Secondary infection site 100 | 103 103

No infection

Seconday infection

No infection

Totals

Lightning damage

No lightning damage

Percentage

Because of this apparent close relationship between lightning damage, the

presence of ambrosia beetles (Plate 3.10), and the subsequent infection by

F. oxysporum, and the demonstrated link of infection by F solani and ambrosia

Plate 3.10

Ambrosia beetles actively

invading an Angsana trunk

which had recently been struck

by lighting.

i Sy

110

beetles in the literature, it was important to determine, firstly the identification of

the ambrosia beetles and secondly, whether these beetles were capable of carrying

F. oxysporum as part of their fungal flora.

Collection of ambrosia beetles were carried out between April 1992 and

September 1994 at University Road, Holland Road Fringe car park, Pioneer Road

and Sembawang Field Experimental Station. The beetles were trapped using

Window Flight Traps (Beaver & Loyttyniemi, 1991; Martin, 1977), funnel traps

and also collected individually using forcepts.

Whole beetles and severed parts, such as head, thorax and abdomen were

analysed for the presence of micro-organisms. The selective media used was a

Peptone PCNB agar (PPA) as described by Burgess et. al. (1988). This media was

later modified with the addition of | g per litre of carbofuran (Furadan® 5G)

granules to control the nematodes carried by the beetles, which quickly resulted

in bacterial contamination swamping the plates.

3.2.1 Identification.

The ambrosia beetle which predominated in all the collections was identified

as Platypus parallelus (Fabricus) (= P. linearis Stephens), Family :Platypodidae.

(Plate 3.11). This was confirmed by Dr. Mick Cox of the International Institute

of Entomology, CAB International, and also by Prof. D. H. Murphy from the

National University of Singapore.

The beetle is polyphagous and is found throughout the tropical and subtropical

regions of the New World, and in Africa and Malaysia. Dr. Cox (pers. comm.)

communicated that the beetle could transmit Fusarium fungus since the platypodid

Cylindropalpus auricimans has this ability in West Africa.

The other species of ambrosia beetle, infrequently caught in the collections,

was identified as Platypus geminatus by Prof. D. H. Murphy from the National

University of Singapore .

3.2.2 Isolation of Fusarium oxysporum from ambrosia beetle

F. oxysporum, F. solani, Penicillium sp., Gliocladium sp., and Trichoderma sp.,

were isolated from larvae taken from the galleries. Saprophytic bacteria were also

isolated in some cases.

Similarly, the adult insects (male and female) also yielded F oxysporum,

F: solani, Penicillium sp., and Gliocladium sp., and also Verticillium sp.

In an effort to localise these micro-organisms, parts of the adult insects were

plated and analysed for the presence of fungal growth. F. solani was found on the

head, thorax and abdomen of the insects, whereas F oxysporum was only detected

111

Plate 3.11

The ambrosia beetles (Platypus parallelus, family: Platypodidae) trapped from the diseased Angsana. Female

(a); Male (b).

on the abdominal parts. Cladosporium sp. and Nigrospora sp. were also detected

in these experiments.

Besides studies on mycangia, other workers were cited by Browne, (1961) who

reported that spores could be carried by the beetle on brushes or hairs on the head,

under the elytra, and on ventral abdominal hairs. It is also generally agreed that

mycelium is utilised as food, but spores usually pass out undigested.

Out of the 200 beetles collected at University Road, only 6% carried

F. oxysporum, while beetles taken from funnel traps showed 33% had F. oxysporum

on them. Collections of beetles taken at Holland Road fringe car park, pooled

collection of 50 - 100 beetles each, revealed that 30 - 40% of them carried

F. oxysporum. Of the twenty-six beetles which were either drowned in the resin

or captured alive while initiating attack on the bark, 15% were found to carry

F. oxysporum.

These results demonstrated that where inoculum was high as a result of

increased numbers of infected trees, it followed that a larger percentage of the

beetles emerging from the area would carry the pathogen. At the university Road

. there was only one infected tree at the time of trapping, whereas the Holland Road

fringe car park area was in the midst of an outbreak. Several trees were diseased

or became infected over the trapping period.

3.3 Discussion

The link between ambrosia beetles and Fusarium solani as an ambrosia fungus

is well documented (Kok, 1972; Zimmermann, 1973; and Hara and Beardsley,

1976). That F solani, an aggressive plant pathogen, was also linked with the

ambrosia beetles as a vector of the disease is therefore to be expected. (Kessler,

1974: Anderson and Hoffard, 1978 and Weber, 1979; 1985)

Fusarium oxysporum is very similar to F. solani in respect to the production

of micro-conidia, so that any F oxysporum which had colonised the ambrosia

tunnels, would be picked up by the adult ambrosia beetles leaving the tunnels and

transported to, and inoculated into, any new susceptible trees colonised by the

beetles.

Murphy (1994 pers. comm.) stated that “the biology of the ambrosia beetles

is centred around the transmission of their fungal symbionts, for which they have

specialised anatomical devices such as hair tufts or chitonous pockets designed

to carry the spores. If a tree became infected with a fungal pathogen such as has

happened in the Angsana wilt disease, it would be almost impossible for ambrosia

beetles not to pick it up on emergence, and almost inevitable that they would infect

any new tree that they successfully infested themselves.”

Throughout the project it was common when isolating F’ oxysporum, to find

the mycelium to be associated with the ambrosia holes in the wood (Plate 3.12),

113

Plate 3.12

White cottony mycelium F. oxysporum growing out from Ambrosia beetle holes on surface sterilized diseased

wood tissue placed in petri dish. The only mycelial growth to be seen on the wood is associated with the ambrosia

tunnel.

however, it was only towards the end of the project that we demonstrated that

ambrosia beetles do carry the F oxysporum. In one sample 40% of those beetles

sampled were carrying F oxysporum.

Of the 170 unhealthy trees inspected, 147 (86%) were infected with F

oxysporum, and of these, 132 (90%) were also struck by lightning and 128 (87%)

had both lightning and ambrosia beetles. Only 15 (10%) were due to secondary

spread and not related to lightning (Table 3.1).

Of the remaining 23 (14%) trees, all were struck by lightning but no F

oxysporum was isolated.

These figures demonstrated dramatically the importance of lightning in the

epidemiology of the disease, not only in initiating the primary infection sites (97%)

but also in the secondary infection sites (86%).

Of the 170 trees examined in the last 10 months of the project, only 4 (3%) of

those which had been struck by lightning had no ambrosia beetles associated with

the damage at the time of inspection (Table 3.1).

The hypothesis presented here for the life cycle of the Angsana wilt disease,

is that lightning damage to an Angsana provides the stress which attracts the

114

ambrosia beetles. If these beetles are contaminated with F oxysporum spores, then

infection of the already stressed tree is likely to follow. The secondary spread away

from this primary infection site, is by F oxysporum which has entered the soil

from the infected tree.

The present epidemic of the Angsana wilt disease is a result of a large build

up in the number of infectious ambrosia beetles, and it is this population which

should be targeted as the weak link in the disease cycle where, with a combined

effort of the various government organisations involved, control of the disease

can be achieved.

4. THE CONTROL STRATEGY

Sporadic occurrences of Angsana wilt disease occurred around Singapore

between the late 1970s and 1982. This culminated in the visit of Harden (1982)

and the following control measures being implemented by Parks and Recreation

Department:

* Drenching of the surrounding soil with fungicide.

¢ Horticultural sanitation.

@ Monitoring.

With the removal of the infected trees and the intensive soil drenching with

fungicides, no further infections were encountered and the spread of the disease

appeared to be under control. In October 1983 the Plant Protection Unit of Parks

and Recreation Department was disbanded, and no cases of Angsana wilt were

recorded between October 1983 and October 1988. In late 1988 two Angsanas

along Collyer Quay were found to be diseased. Between 1988 and 1991 over 800

trees died in Singapore because of the disease.

As the implementation of these stringent control measures failed to eradicate

the disease, it became one of the primary objectives of the Angsana wilt project

to establish possible reasons for the failure, and to formulate a modified disease

control strategy, based on the findings of the epidemiological studies.

The control strategy developed in 1982 was based on the assumption that the

disease was solely caused by the soil-borne fungus F. oxysporum, and therefore

as it was assumed that the disease originated from the soil, there was no urgency

to remove infected trees. It was also postulated that drenching the soil with

fungicides would eradicate the pathogen from the soil, and also, because the

chemicals used were systemic, they would be translocated and therefore have

activity against the pathogen within the tree.

The current research, however, has empirically demonstrated that although the

disease is of soil-borne origin, the initial spread of the fungus to new sites is by

115

an insect vector. Controlling the disease once the pathogen has reached the soil

in a new site, is therefore to implement control measures at the end of the primary

infection cycle. Control, to be successful must be implemented at the beginning

of the primary infection cycle.

4.1 Development Of The Control Strategy

The modified control strategy has two objectives. Firstly to prevent the disease

from becoming established in new sites, and secondly to slow down the

progression of the disease in those areas where secondary infection is already

established.

No control strategy will stop an epidemic instantly. Success depends on slowing

down the progress of the epidemic, turning the positive multiplication rate of an

active epidemic, into a negative multiplication rate which eventually results in the

total control of the disease as the pool of inoculum is gradually reduced to one

below epidemic thresholds. Sporadic incidences could then be mopped up.

4.2 Overview.

There are two main methods of spread of the Angsana wilt disease. Firstly by

the F oxysporum infested ambrosia beetles which invade a damaged tree following

a lightning strike, and secondly by the traditionally accepted method of soil-borne

hyphae and spores, and by root-to-root contact.

The lightning - ambrosia beetle - F oxysporum complex explained 97% of the

primary infection sites inspected during the 1993-94 survey. This complex also

played a significant role, being associated with 87% of the secondary infection sites

(Table 3.1).

Most of the pockets of secondary infection sites scattered around Singapore

are well established, and the pattern of diseased trees within these pockets suggests

that the spread of the disease is predominantly localised. The immediate neighbours

of an infected tree are highly at risk with a more than an 80% chance of becoming

infected . All Angsanas within 100 m of an infected tree are also at risk, although

this distance will vary depending on the terrain between the trees and the assistance

this provides for the spread of the fungus.

As a result of the trials on injecting infected trees to stop the development of

the disease, it was apparent that once a tree was infected, and subsequently when

the symptoms of the disease appeared, it was already too late to control the disease

within the tree by chemical treatment. This is understandable as F oxysporum is

a vascular pathogen attacking and destroying the vessels which move the

fungicides around the tree. Once these vessels are destroyed systemic action of

the fungicide stops. Although it was demonstrated that chemical treatment will

116

delay the onset of the disease symptoms for between 4 - 6 months, it will not

prevent the inevitable death of the tree. The fungus will again advance as the

fungicide within the healthy vessels becomes diluted or is broken down. Secondly

once infection is established, ambrosia beetles rapidly help spread the pathogen

within the tree.

The time requirements for the removal of the lightning damaged trees is

therefore based on the life cycle of the beetles. This time period, from the first

invasion of the tree to the emergence of the second generation of infected beetles

can be predicted within narrow limits.

As the ambrosia beetles are the weak link in the disease cycle, and a major

contributing factor to the current epidemic, it was pertinent to develop the control

strategy around the life cycle of the beetles.

4.3 The Control Strategy

The control strategy is aimed at breaking the disease cycle by controlling the

ambrosia beetles, and thus preventing F oxysporum from infecting the lightning

- damaged tree and becoming established in the soil. The strategy is based on the

rapid removal of these lightning damaged trees, chemical treatments in the form

of sprays to prevent insect invasion of the damaged trees and sprays and injections

to protect the surrounding trees from invasion and infection.

# Monitoring. The success or failure of the control strategy will depend on the

efficiency of the monitoring team in detecting the lightning damaged trees early

and co-ordinating the treatment programme.

* Removal of lightning damaged trees. It is paramount to the success of the

control strategy that: 1) insecticide sprays be applied to the damaged tree within

days of the first symptoms of wilting being noted to prevent the colonizarion

by ambrosia beetles, and 2) the tree and as much of the root ball as is possible

be removed within three weeks of the lightning strike.

¢ Treatment of High Risk trees by injection. All Angsanas within 100 m of

an infected tree are considered to be highly at risk and therefore likely to

become infected in the future. All high risk trees should be sprayed to run-off

as far up the trunk as is practicable, using a conventional horticultural high

volume sprayer with a mixture of fungicide and insecticide and injected with

a mixture of fungicide and insecticide within three days of the lightning

damaged tree being noted. Addition of a sticker to the chemical spray is

recommended.

@ Second injection. Trees should be re-injected 4 months later.

¢ Trenching. Where possible trenching should be carried out between damaged

Ly,

and undamaged trees to minimize the likelihood of tree to tree spread via root

contact.

@ Hygiene. As with all infectious diseases, hygiene is of paramount importance.

All tools which are used on or around infected trees should be treated with

95% alcohol using a hand-held plastic sprayer. Extreme care must be taken to

ensure that ambrosia beetles do not get inside vehicles and transported to new

areas. Felled trees should be burned to rid them as a source of beetle dispersal.

¢ Replanting with Angsanas. As long as the lighting damaged tree is removed

before infection is established within the damaged tree, then there should be

no danger in replanting with a second Angsana. However, once infection is

established in the damaged tree, then it must be assumed that the fungus has

reached the soil and measures taken accordingly.

¢ Trees outside the jurisdiction of Government Departments. Although the

success of this control strategy can proceed independently of the removal of

lightning damaged trees within private property, owners should be advised that

these trees are not only a physical danger to property, but also a health hazard

to surrounding Angsanas, and that it is important that these trees be rapidly

removed. It would be frustrating to see the control efforts wasted because of

one or two small hot-spots of infection remaining within privately controlled

land. Hot-spots which would act as the source of infection for subsequent

outbreaks of the disease.

4.4 Discussion

Currently 87% of new infections of Angsana wilt disease are ambrosia beetle

related and it is this population of infectious beetles which is the initial target of

the control strategy. To control this population is to control the epidemic. This

can only be done by the rapid removal of all lightning damaged trees, the potential

sites of infection. Initially, this will therefore mean the removal of 10% of trees

which would not have become infected, however, this is a small price to pay for

the ultimate success of the control strategy. As the control strategy takes effect

and the population of infectious ambrosia beetles has reduced then the number of

trees likely to remain healthy will rise to an ultimate aim of 100%. As the

population of infectious ambrosia beetles drops, decisions will have to be made

as each new lighting damaged tree is detected, as to whether or not the tree should

be removed on the basis of the likelihood of it becoming infected.

Following the implementation of the control strategy, there will be a period of

several months before there is a reduction in the number of trees being removed

due to Angsana wilt each month. This lag period will correspond to the time that

it takes for all those trees which are already infected at the time of treatment, to

develop symptoms and be removed. Once this reservoir is removed and the

118

population of infectious ambrosia beetles is reduced, rapid progress should be made

in controlling this insidious disease.

5 SCREENING OF ANGSANA (Pterocarpus indicus) FOR

RESISTANCE TO Fusarium oxysporum

Although the short term aim of the project was to develop a control strategy

based on chemical control, the long term aim was to identify lines of P. indicus

resistant to FE. oxysporum.

5.1 Collection of planting material

Planting material was collected from four localities in West Malaysia, ten

localities in the Philippines, three in Myanmar, five in the Solomon Islands, three

in Papua New Guinea, three in Java, eight in the Sulawesi Islands and six localities

in the Maluku Islands. In total, 610 trees were sampled for hardwood cuttings and

188 samples of seeds were collected (Fig. 1).

Pterocarpus indicus is a natural component of the coastal low-land flora of

South East Asia, however, because it is prized as a timber for both building and for

furniture, it is very rare to see mature trees in the jungle accessible to the local

population. Conversely, because it is so easily propagated and provides excellent

shade, Angsanas were commonly found growing in villages as either single trees

or as avenues for shade trees, and as rows of trees for fencing. It was also found as

isolated shade trees in cleared jungle being used for agriculture.

Planting material was also collected from the Angsanas growing around

Singapore on a weekly basis between February and November 1993. During which

time 270 samples of cuttings and 58 collections of seed were made.

Where possible 100 seeds per tree were collected (Plate 5.1) as, not only were

they physically easier to collect and gave a greater establishment rate, but it also

meant that we were selecting within a segregating population where each seedling

was genetically unique.

Where seeds were not available, 3 or 4 hardwood cuttings, 20 - 25 cm long and

between 2 - 3 cm in diameter were cut. Hardwood cuttings were collected by

removing a 2 - 3 cm diameter branch, using either a pruning saw or a tree pruner

which was extendible to 5 m. The branch was then cut into 3 - 4, 20 cm lengths.

These were placed into a zip-lock® plastic bag for transport. Data were recorded

on tree locality and physical characteristics, such as tree form, leaf shape and size

etc. Cuttings were soaked in water overnight, then soaked for 30 min in a 1:10

sodium hypochlorite solution (Chlorox®), wrapped in paper towels treated with

Benomyl (Benlate® S5OWP), and returned to the zip-lock® plastic bags for transport.

Every effort was made to ensure that all hardwood cuttings were planted into soil

within 14 days from collection.

119

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120

Plate 5.1

Angsana seeds collected from both Singapore and overseas exhibit a large variation in size across the wing,

from 3.5 cm to 7 cm.

In instances where neither seeds nor hardwood cuttings were available, then

softwood cuttings were taken.

The wings of the seeds of Pterocarpus indicus were cut to expose the seeds

inside the pods. Seed pods were soaked in water overnight before being sown in

washed coarse sand in 450 x 345 x 70 mm plastic trays. Fresh seeds usually

germinated within three days. However, this period is extremely variable with some

seeds germinated overnight while a few exceptions took over a year to germinate.

Vigorous seedlings were pricked out into 75 mm plastic pots containing a general

potting mix. Seedlings were allowed to grow until they were between 10 - 15 cm

tall before being screened for resistance.

Hardwood cuttings when they arrived from overseas, were thoroughly cleaned

and soaked in a dilute fungicide solution for 30 min. before being potted in 75 or

120 mm plastic pots in either sand, soil or a general potting mix . The cuttings

were allowed to grow until they were well rooted and growing vigorously before

being put through the screening process. The establishment of hardwood cuttings

depended on the time from cutting to planting. The establishment rate for the

Singapore cuttings was 62% while the establishment of those collected overseas

ranged from 13% to 19%.

121

5.2 Screening for Resistance

A weekly cycle was developed to enable an efficient screening of the large

number of established hardwood cuttings and seedlings established in the nursery.

Five different isolates of F oxysporum were grown on PDA (Potato Dextrose

Agar) at 28°C for 5 days. Five plugs were removed using a sterilised 5 mm cork

borer and ground in a 10 ml glass macerator to form a thick suspension. This

suspension was added to 500 ml of PDA at 45 - 50°C just prior to pouring. The

petri-dishes were left on a bench in natural light to stimulate the production of

micro-conidia. After seven days the fungal lawns were blended together in sterile

water. The final concentration used was three plates to 500 ml of sterile water.

Cuttings and seedlings were removed from their pots, labelled, and their roots

washed clean and trimmed to about 1.0 - 1.5 cm in length. They were then placed

into plastic jars containing the inoculum. They were left in the inoculum for four

hours under full sunlight, before being re-potted using the original potting mix.

The pots were then watered with the inoculum from the jars. The inoculated

cuttings or seedlings were only watered when the soil became dry in order to

maintain them under mild stress. Inoculated plants were monitored closely for five

weeks, and plant deaths were recorded accordingly.

5.3 Results

Inoculated plants usually started to show signs of wilting (yellowing and

dropping of leaves) towards the end of the third week. Scraping the stem to reveal

some discoloration and vascular staining would confirm that the material was

infected. Some plants died during the third and fourth weeks, however, the majority

of plants died during the fifth week, with a few taking more than five weeks to

wilt and die.

At the end of five weeks, those plants that had survived were checked, and

healthy plants were re-labelled and returned to the nursery area to await subsequent

screening.

From November 1992 to Nov. 1995, 35 batches of P. indicus, consisting of

over 2,600 seedlings and 650 cuttings were subjected to one screening, and from

those, 1,214 seedlings and 69 cuttings have since been inoculated the second time,

and 652 seedlings and 47 cuttings were inoculated for the third time.

All plants will now be planted into natural soil known to contain & oxysporum,

for a final confirmation of resistance.

5.4 Discussion

During the course of the project over 650 hardwood cuttings and 2,600

122

seedlings which had been collected from 7 countries were tested. Because we are

looking for a single gene resistance, which is expressed as an all-or-nothing

reaction, resistant plants survive and susceptible plants die.

All the material had been subjected to three screenings. All surviving plants

will now be planted into natural soil which is known to contain F oxysporum before

we can confidently describe this material as resistant to F oxysporum.

From Table 5.1 it can be seen that there is a wide range in the percentage survival

of the hardwood cuttings. This is understandable as the resistance/susceptibility of

the hardwood cuttings is that of the parent tree. The low incidence of survival of

hardwood cuttings from Singapore (25%) would suggest a low incidence of

resistance within the Singapore Angsana population. This could go some way to

explain the development of the disease here rather than in other regions.

Table 5.1 Screening of Angsana seedlings and hardwood cuttings for resistance to Fusarium oxysporum.

Cn Ee ee eee eee Eee

ee ee ee are Pe

on Ree Roe neo ee ee)

123

The ratio of resistance to susceptible plants in the seedling population must,

however, reflect the genetic composition of the population. That the overall

percentage of survival was close to 50% (535) is encouraging, and further genetical

studies will be carried out.

That the survival of the seedlings increased from 54% in the first inoculation to

82% in the second and to 95% in the third inoculation is an encouraging indication

that resistance has been identified. On the other hand, at no time did the results

obtained with the hardwood cuttings parallel the success obtained with seedlings.

Not only was the initial establishment of the hardwood cuttings very low (less than

20% for overseas cuttings) but the attrition rate while growing the plants in small

plastic bags under the direct tropical sun was very high.

This work will continue and it will take an estimated further two years to have

resistant planting material available for multiplication.

ACKNOWLEDGEMENTS

It is with gratitude that we wish to thank the National Science and technology

Board and Mr. Jim Baker from SGS Singapore Pte. Ltd., for their foresight and

financial support that made this project possible, and Mr. Robert Lee of SGS AIl-

Pest Pte. Ltd. for his support throughout this project.

We would also like to thank Professor Gloria Lim; Dr. D. Brayford of

International Mycological Institute, England; Professor L. Burgess of Sydney

University; Dr. B. Summerell of Royal Botanical Gardens, Sydney, Australia; Dr

B. Hawthorn of Food and Hort Research, CRI of New Zealand; and Dr. H. Nirenberg

of Federal Biological Research Centre for Agriculture and Forestry, Germany for

their assistance in the identification of the Fusarium isolates.

Also to the following, without whom, the collection of the Angsana material

from such a wide geographical area would not have been feasible : Mr. U. Aung

Than and staff of the Forest Department, National Parks and Wildlife Conservation

Division, Myanmar; Dr. Dimingo A. Madulid, Mr. Toy Sageal, and Mr. Efran Romeo

of the Botany Division, National Museum, Manila, Philippines; Mr. Myknee Qusa

and Mr. Patterson of the Forestry Division, Ministry of National Resources, Honiara,

and Mr. Geoff F.C. Dennis of Solomon Islands; Mr. Ian Orrell of the Oil Palm

Research Association, and Mr. Jim Bell of Papua New Guinea; and finally Ms.

Sandra Upston and Mr. Julian Sanderson of SGS Singapore Pte. Ltd. who made the

collection in the Sulawesi and the Maluku Islands possible.

The assistance of Professor Lee Seng Kong and Mr. Ho A.C. of Nanyang

Technological University in the development of the aeroponic system is fully

acknowledged. In addition the assistance of Professor D. H. Murphy of National

University of Singapore is also acknowledged.

124

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Freshwater Swamp Forest in Singapore, with Particular

Reference to That Found Around the Nee Soon Firing Ranges

I.M. Turner’, Boo Chih Min', Wong Yew Kwan’, Chew Ping Ting? and

Ali bin Ibrahim?

"School of Biological Sciences, National University of Singapore, Singapore 119260

289 Soo Chow Garden Road, Singapore 575526

3. National Parks Board, Singapore Botanic Gardens, Singapore 259569

Abstract

The freshwater swamp forest found around the firing ranges at Nee Soon is the last remaining area of this

forest formation in Singapore. The vascular plant flora of freshwater swamp forest in Singapore is reviewed, with

particular reference to the Nee Soon forest. Some preliminary soil, water and foliar analyses indicate that the

swamp forest at Nee Soon may have considerably higher amounts of phosphorus available to the vegetation than

the dryland forest of Bukit Timah Nature Reserve. Nitrogen and potassium availabilities appear to be similar at

the two sites. The Nee Soon swamp forest is an extremely important site for Singapore’s native biota and should

receive the highest priority for conservation.

Introduction

One of the least well-known of the forest formations of West Malaysia is

freshwater swamp forest. So named to distinguish it from the saltwater mangrove

swamps, it is a forest type of wet, often seasonally flooded, lowland areas (Whitmore

1975). The only work of any note specifically concerning freshwater swamp in this

region is Corner’s (1978) account of his studies, which were mostly floristic in

nature, of these forests in southeast Johore and Singapore. Corlett (1991) has

estimated that approximately 5% of Singapore was covered in freshwater swamp

before the major forest clearance of the nineteenth century began. The only

remaining area of swamp forest in Singapore today is to be found around the Nee

Soon Firing Ranges in the Central Catchment Nature Reserve (Fig. 1). The fauna

of the Nee Soon forest and surrounding areas has been outlined by Ng and Lim

(1992). Botanically, this area has never been studied closely, and it seems timely to

consolidate our current knowledge of the forest into one publication.

The Nee Soon Swamp Forest

Swamp forest occurs in low-lying areas where slow-flowing streams drain

shallow valleys. The ground becomes saturated with the water table close to the

soil surface and there are often periods of flooding. Such swamps probably occurred

historically in the upper reaches of all the rivers in Singapore (Corlett 1991). Corner

(1978) made detailed studies of the flora and forest structure of the swamp forest

near Mandai Road, which is now submerged beneath Upper Seletar Reservoir, and

in Jurong. Industrial and housing development have totally destroyed the Jurong

swamps. This leaves the Nee Soon swamp as the last remnant of the forest formation

in Singapore. It has probably survived because of the Firing Ranges, which lack

130

baffling walls or banks, making the surrounding area dangerous to the public and

unsuitable for any form of development. The approximate extent of the swamp

forest is shown in Figure 1. This has been delineated from data collected by a

number of field researchers, particularly D.H. Murphy who has mapped many of

the main streams of the area. It is estimated that the swamp forest covers about 87

ha, though much of this has been disturbed quite heavily at various times in the

past. There is concern that changes in the drainage regime of the forest may be

causing excessive soil erosion along some of the streams and causing the death of

the larger trees in certain areas (D.H. Murphy pers. comm.).

G + rst WS

e, UPPER XQ

eins) \

a SELETAR cure ye

RESERVOIR \ ery

7 N

Figure 1. Map showing the location of the Nee Soon Freshwater Swamp Forest, Singapore. The stippled area

indicates the approximate extent of the swamp forest.

Floristics

Appendix | gives a list of vascular plant species recorded from swamp forest in

Singapore. It is compiled from the list given by Corner (1978) for his plots at

Jurong and Mandai Road, specimens in the herbarium of the Singapore Botanic

Gardens (SING) where Nee Soon or Chan Chu Kang are given as collecting

localities, recent collections from the Nature Reserves mostly as outlined by Turner

et al. (1994) and the results of the forest survey of Wong et al. (1994). Clearly, the

flora of the swamp forest is rich, with more than 700 species having been recorded

from this vegetation type in Singapore.

131

Vegetation Structure and Physiognomy

The forest at Nee Soon has an appearance similar to that outlined for freshwater

swamp forestin other localities by Corner (1978). The ground is saturated with frequent

small streams and pools of water. Many of the trees exhibit the prolific development

of large stilt roots and/or abundant pneumatophores of various descriptions, that

Corner illustrates profusely in his book. In apparently primary areas, the trees can

reach large size, with heights of 40 m or more being recorded (Hill 1977).

During a survey of the forest communities of the Central Catchment Nature

Reserve (Wong et al. 1994) three 0.2 ha plots were enumerated in the swamp forest.

The location of plots 26, 27 and 28 are given in Figure |. The floristic composition

for trees of greater than 30 cm girth at breast height (1.3 m) in the three plots is

given in Table 1. The composition of the three plots was different from the plots

surveyed in the primary forest on dryland sites (Turner et al. 1996b). Important

species in terms of basal area were Pometia pinnata, Palaquium xanthochymum,

Mangifera griffithii, Gluta wallichii and Strombosia ceylanica. Corner (1978)

reported these to be abundant in his Mandai Road plots, indicating a strong similarity

in the forest between the two sites, which is not surprising given their propinquity.

Table 1

Composition of the forest community (> 30 cm gbh) in three 0.2 ha plots in the Nee Soon swamp

forest. n = number of individuals of each species, ba = total basal area (cm2). Species are in descending

order of total basal area summed for the three plots. Some individuals could not be identified and are

referred to by code numbers.

plot

26 27 28

species n ba n ba n ba

Pometia pinnata 6 10599 5 10357 2 6531

Palaquium xanthochymum 5 17983 2 1866 D, 7128

Mangifera griffithii 5 8853 7 4649 6 6160

Gluta wallichii 5 6466 3 3082 5 6362

Strombosia ceylanica 10 8104 3 1017 11 5295

Z9 1 13440

Pentace triptera 5 7322

Z 03 I 6237

Palaquium rostratum yy 3107 1 2114

Santiria rubiginosa 5 4955

Mussaendopsis beccariana 4 1406 5 1514 4 1912

Gonystylus maingayi 2 4494

Polyalthia glauca 5 1635 6 2521

Campnosperma auriculatum l 3677

Bhesa paniculata 2 1571 1 1559 4 484

Lophopetalum multinervium 1 232 4 2982

Calophyllum teysmannii 2 3199

Ull 1 3182

Xanthophyllum ellipticum 2 2750

Gymnacranthera farquhariana 4 1656 1 447 2 604

132

Vatica ridleyana

Ganua motleyana

Ochanostachys amentacea

Sandoricum beccarianum

Myristica cinnamomea

Myristica elliptica

Xanthophyllum affine

Carallia brachiata

Macaranga conifera

Alstonia angustiloba

Horsfieldia polyspherula

Triomma malaccensis

Scaphium macropodum

Artocarpus kemando

Xylopiamalayana

Eugenia cerina

Diospyros maingayi

Jackiopsis ornata

Lithocarpus bennettii

Sterculia macrophylla

Alseodaphne bancana

Sterculia cordata

Magnolia candollii

Z09

Campnosperma squamatum

Aquilaria malaccensis

Tristaniopsis merguensis

Archidendron ellipticum

Horsfieldia sucosa

Aglaia rubiginosa

7, 17]

Blumeodendron tokbrai

Pellacalyx axillaris

Horsfieldia crassifolia

Xylopia fusca

Alangium nobile

Xanthophyllum obscurum

Knema communis

Osmelia philippinensis

Litsea erectinervia

Memecylon edule

Knema curtisii

Lophopetalum wightianum

Garcinia parvifolia

Kibatalia maingayi

Knema conferta

Mastixia trichotoma

Cryptocarya ferrea

MYRISTICACEAE 1

Beilschmiedia kunstleri

Castanopsis nephelioides

735

2060

758

1422

558

733

1450

718

828

241

1146

1052

509

471

140

575

Di,

Ke wm he

plot

215

2192

588

207

1790

1209

376

1559

1384

1263

740

263

998

993

985

409

115

176

127

320

602

454

535

509

NK NDN We

Ss JS eS) FS)

— es OD

1758

2166

2027

1353

1174

199

1811

1283

1515

736

692

1386

382

119]

194

832

1016

926

1165

447

796

616

688

328

644

2982

366

460

447

436

a ae

277

115

bo bv

159

361

183

412

14)

241

215

191

176

109

134

Buchanania sessilifolia

Z 16

Elaeocarpus petiolatus

Lithocarpus sundaicus

Eurya acuminata

Sterculia rubiginosa

Gymnacranthera bancana

Microcos blattifolia

Bhesa robusta

Stemonoporus scorpioides

Memecylon lilacinum

Diplospora malaccensis

Dysoxylum excelsum

Ardisia colorata

Symplocos fasciculatus

Xanthophyllum vitellinum

TOTAL (bain m’)

26 Dif

1 97

]

1 72

108 9.18 107

plot

6.64

f2,

1 97

1 81

121 8.20

The terrestrial herbs have been surveyed also in these three plots (Turner et al.

1996a) . Twenty species were recorded (Table 2), with only Aglaonema nebulosum

and Labisia pumila being found in all three plots. The Araceae proved to be

particularly diverse.

Table 2

Terrestrial herbaceous species recorded from the three 0.2 ha plots enumerated in Nee Soon swamp

forest. + indicates presence.

species

Aglaonemanebulosum

Aglaonema nitidum

Alocasia denudata

Amischotolype gracilis

Cryptocoryne griffithii

Cyrtosperma merkusii

Homalomena sp. 1

Homalomena sp. 2

Labisia pumila

Lecanorchis malaccensis

Leptaspis urceolata

Lindsaea doryophora

Mapania cuspidata

Mapania squamata

Ophiorrhiza singaporensis

Peliosanthes teta ssp. humilis

Plocoglottis javanica

Schismatoglottis wallichii

Syngramma alismifolia

Trichomanes obscurum

Total number of species

plot

plo

t++teteteeet

plot

135

Soil

The soil in freshwater swamp forest is generally rich in organic matter,

presumably because the anaerobic nature of the waterlogged conditions reduces

the rate of decomposition. However, this process has not proceeded far enough

to develop soils of high enough organic matter content to be considered peats

(generally >90% loss-on-ignition by mass). Some preliminary analyses for soil

samples from Nee Soon (Table 3) showed the soil surface to have high loss-on-

ignition, averaging nearly 80% and approaching 90% in places, though the values

dropped rapidly below 5 cm depth to generally less than 50% loss-on-ignition.

Some pH measurements showed that stream water and soil water lay in the range

4.6-5.5, with soil pH at 5 cm depth to be lower, 4.0-4.5. The concentrations of

the major plant nutrients, nitrogen, phosphorus and potassium, were measured in

the litter, soil and some foliar samples (shade leaves) gathered at Nee Soon (Table

4 and 5). Surprisingly, the litter generally had lower concentrations of the important

elements than the top 5 cm layer of the soil. Possibly this represents the flushing

of nutrients down through the soil.

The nutrient concentration data can be compared with values obtained for the

coastal hill dipterocarp forest at Bukit Timah (Grubb et al. 1994). The comparison

of the soil values between the two sites is difficult given the very different soil

bulk densities because of the high organic matter content of the soils at Nee Soon.

Bukit Timah has more acidic soil (3.7-4.0 versus 4.0-4.5) and slightly higher litter

concentrations of N (12.9-14.5 versus 9.4 mg g"'), lower concentrations of P

(0.21-0.23 versus 0.47 mg g'') and higher concentrations of K (2.0-5.9 versus 0.48

mg g'). The foliage samples show a similar pattern with higher mean P in the

leaves at Nee Soon (1.72 versus 0.68 mg g''), but lower N (13.2 versus 17.1 mg

g') and K (8.3 versus 10.5 mg g') concentrations. It would appear that Nee Soon

probably has more phosphorus available for plant growth than Bukit Timah, but

possibly slightly less nitrogen and potassium. Phosphorus appears to be the nutrient

limiting the growth of non-mycorrhizal plants on Bukit Timah soil (Burslem et

al. 1994), though mycorrhizal plants seem not to be responsive to P fertilization

(Burslem et al. 1995). Nee Soon may, therefore, present a potentially more fertile

site for plant growth, but of course the water-logged conditions are highly

unfavourable to many species. This relatively high nutrient availability may be

what prevents peat formation from occurring.

136

Table 3 Table 4

Analyses of soil and leaf litter from Nee Soon Swamp Foliar analyses for shade leaves from trees at Nee Soon

Forest, Singapore. Swamp Forest, Singapore.

loss-on- total N totalP total K species total N totalP total K

ignition (mg g') (mg g') (mg g") (mg g') (mg g') (mgg")

(%) Archidendron sp. 18.9 1.28 8.9

leaf litter Calophyllum sp. Ici 0.55 >is

sample | - 9.88 0.52 0.52 Desmos dasymaschala_ 16.2 2.86 11.0

sample 2 - 8.50 0.42 0.48 Diospyros sp. 10.6 1.88 7.5

sample 3 - 9.83 0.49 0.43 Litsea ridleyi O77, 2.41 12.3

mean - 9.40 0.47 0.48 Nephelium sp. 14.2 1.95 7.1

Pternandraechinata 12.4 1.22 5.8

soil (0-5 cm) Strombosiaceylanica 14.3 1.16 8.3

sample | 88.7 22.2 0.87 0.51 sp. indet. 14.8 Dale) Wes

sample 2 60.8 12.8 0.70 0.52 mean 13.2 1.72 8.3

sample 3 89.6 17.5 0.87 0.64

mean TE 17.5 0.81 0.56

soil (5-10 cm)

sample | 54.8 9.53 0.84 0.72

sample 2 34.8 6.25 0.45 0.69

sample 3 47.5 7.41 0.54 1.16

mean 45.7 7.73 0.61 0.86

Conservation

Nee Soon is now Singapore’s last fragment of freshwater swamp remaining.

Many species of plants and animals are now restricted to this area, hence making

Nee Soon of great importance in terms of the biodiversity of Singapore. The plant

list includes Aeschynanthus wallichii, Bulbophyllum macranthum, Cystorchis

variegata, Kopsia singapurensis, Luvunga crassifolia, Neoscortechinia kingii and

Xylopia fusca. Ng and Lim (1992) list many animal species restricted, in Singapore,

to Nee Soon. Species found at Nee Soon are generally not found at Bukit Timah,

and vice versa, so the two are the twin pillars of biodiversity in Singapore and

require strict protection.

Acknowledgements

We are grateful to D.H. Murphy, H.T.W. Tan, K.S. Chua and Haji Sidek for

sharing their expertise and data concerning Nee Soon swamp forest. Jeanette Ong

provided valuable technical assistance in the laboratory, and Jean Yong helped in

preparing the figure.

References

Burslem, D.E.R.P., Turner, I.M. & Grubb, P.J. (1994). Mineral nutrient status of

coastal hill dipterocarp forest and adinandra belukar in Singapore: bioassays of

nutrient limitation. J. Trop. Ecol. 10:579-599.

137

Burslem, D.F.R.P.. Turner, I.M. & Grubb, P.J. (1994). Responses to elevated nutrient

supply and simulated drought among shade-tolerant tree seedlings of lowland

tropical forest in Singapore. J. Ecol. 83:123-132.

Corlett, R.T. (1991). Vegetation. In: Chia, L.S., A. Rahman & D.B.H. Tay (eds.),

The Biophysical Environment of Singapore. Singapore University Press, 134-

154.

Comer, E.J.H. (1978). The freshwater swamp-forest of South Johore and Singapore.

Gdns’ Bull., Sing., suppl. 1:1-266.

Grubb, P.J.. Turner, I.M. & Burslem, D-.F.R-P. (1994). Mineral nutrient status of

coastal hill dipterocarp forest and adinandra belukar in Singapore: analysis of

soil, leaves and litter. J. Trop. Ecol. 10:559-577.

Hill, R.D. (1977). The vegetation map of Singapore: A first approximation. J. Trop.

Geogr. 45:26-33.

Kostermans, A.J.G.H., & Bompard, J.-M. (1993). The Mangoes. Academic Press,

London.

Ng, P.K-L.. & Lim, K.K-P. (1992). The conservation status of the Nee Soon

freshwater swamp forest of Singapore. Aquatic Conserv. 2:255-266.

Turner, I.M. (1994). The taxonomy and ecology of the vascular plant flora of

Singapore: a statistical analysis. Bot. J. Linn. Soc. 114: 215-227.

Turner, I.M., Tan, H.T.W., Chua, K.S., & Metcalfe, D.J. (1994). Recent botanical

collections from the Nature Reserves of Singapore. Gdns’ Bull., Sing. 46(2):1-

36.

Turner, I.M., Tan, H.T.W., & Chua, K.S. (1996a). Relationships between herb layer

and canopy composition in a tropical rain forest successional mosaic in

Singapore. J. Trop. Ecol. 12: 843-851.

Turner, I.M., Wong, Y.K., Chew, P.T. & Ali bin Ibrahim (1996b). Rapid assessment

of tropical rain forest successional status using aerial photographs. Biol. Conserv.

77: 177-183.

Whitmore, T.C. (1975) Tropical Rain Forests of the Far East. Clarendon Press,

Oxford.

Wong, Y.K., Chew, P.T. & Ali bin Ibrahim (1994). The tree communities of the

Central Catchment Nature Reserve, Singapore. Gdns’ Bull., Sing. 46(2):37-78.

138

APPENDIX 1

SPECIES LIST OF VASCULAR PLANTS FOR FRESHWATER SWAMP FOREST IN SINGAPORE

Species marked with an asterisk are those recorded by Corner (1978) from freshwater swamp

forest in Singapore. Where taxonomic revisions have necessitated name changes, or specimens were

misidentified, the name used by Corner is given in parentheses. Other species are those collected

recently or known from herbarium specimens where Nee Soon swamp forest or Chan Chu Kang are

given as the collecting locality. For more information regarding NRS series collections see Turner et

al. (1994). Species recorded from the three 0.2 ha plots at Nee Soon (Wong et al. 1994) are marked

with a dagger (7+). Only forest species are included. The determination of Mangifera species follows

Kostermans & Bompard (1993).

LYCOPODOPHYTA

Lycopodiaceae

Huperzia nummulariifolia (Blume) Jermy - Ridley s.n. (Oct 1889)

Huperzia phlegmaria (L.) Rothm. (Lycopodium phlegmaria) - Ridley 2420 (1884)

* Huperzia squarrosa (G. Forst.) Trevisan (Lycopodium squarrosum)

Selaginellaceae

Selaginella argentea (Wall.) Spr. - NRS1972

FILICINOPHYTA

Adiantaceae

+ Syngramma alismifolia (C. Presl) J. Sm. - NRS1610

* Taenitis blechnoides (Willd.) Sw.

Aspleniaceae

Asplenium batuense Alderw. - NRS1185

Asplenium macrophyllum Sw. - Goodenough s.n. (1890)

* — Asplenium nidus L.

* Asplenium nitidum Sw. (Asplenium glaucophyllum)

Blechnaceae

Blechnum finlaysonianum Hook. & Grev. - Ridley 6121 (1894)

* — Blechnum orientale L.

* — Stenochlaena palustris (Burm.) Bedd. - NRS1589

Cyatheaceae

Cyathea glabra (Blume) Copel. - NRS1526

* Cyathea latebrosa (Wall. ex Hook.) Copel. - NRS1508

Davalliaceae

Davallia angustata Wall. ex Hook. & Grev. - Ridley 3599a (189?)

Dennstaedtiaceae

* Histiopteris incisa (Thunb.) J.Sm.

N59

Lindsaea doryophora Kramer - NRS1519

* Lindsaea parasitica (Roxb. ex Griff.) Hieron. (L. scandens) - NRS0158

Dryopteridaceae

Tectaria barberi (Hook.) Copel. - Maxwell 82-71 (18 Mar 1982)

Tectaria singaporeana (Wall. ex Hook. & Grev.) Copel.

Hymenophyllaceae

Trichomanes christii Copel. - NRS0867

* — Trichomanes motleyi (Bosch) Bosch

*} Trichomanes obscurum Blume - Ridley 6119 (1894)

Lomariopsidaceae

Teratophyllum aculeatum (Blume) Mett. ex Kuhn - NRS0051

* Teratophyllum ludens (Fée) Holttum - NRS0132

Polypodiaceae

* ~Drynaria quercifolia (L.) Sm.

Lecanopteris sinuosa (Wall. ex Hook.) Copel. - NRSO157

Microsorum punctatum (L.) Copel. - Ridley s.n. 1896

Platycerium coronarium (D. Koenig ex O.F. Miill.) Desv. - NRS1603

Pyrrosia angustata (Sw.) Ching

Pyrrosia longifolia (Burm.) Morton - NRS0131

Schizaeaceae

Lygodium longifolium (Willd.) Sw. - Ridley 4229 (Dec 1892)

Schizaea digitata (L.) Sw. - NRSO171

Thelypteridaceae

Christella parasitica (L.) Lév. - NRS0145

* Mesophlebion chlamydophorum (Rosenst. ex C. Chr.) Holttum (Thelypteris chlamydophora)

Sphaerostephanos heterocarpus (Blume) Holttum - Ridley 6120 (1894)

Vittariaceae

* Vittaria ensiformis Sw.

Woodsiaceae

Diplazium crennatoserratum (Blume) Moore - Ridley 4399 (1892)

GNETOPHYTA

Gnetaceae

+ Gnetum gnemon L.

* Gnetum gnemonoides Brongn. - NRSO0781

Gnetum macrostachyum Hook.f. - NRSO196

* Gnetum microcarpum Blume - Maxwell 82-198 (29 Jul 1982)

140

CONIFEROPHYTA

Podocarpaceae

* Nageia wallichiana (Presl) Kuntze

ANGIOSPERMOPHYTA

Acanthaceae

Hygrophila ringens (L.) R.Br. ex Steud. - NRS2438

Justicia vasculosa Wall. - NRS1388

Staurogyne setigera Kuntze - NRS0166

Alangiaceae

* Alangium ebenaceum (C.B. Clarke) Harms var. tutela (Ridl.) Kochummen

+ Alangium nobile (C.B. Clarke) Harms

Anacardiaceae

+ Buchanania sessifolia Blume

*+ Campnosperma auriculatum (Blume) Hook.f.

*+ Campnosperma squamatum Ridl. - NRS101

*+ Gluta wallichii (Hook.f.) Ding Hou (Melanorrhoea wallichii) - Hullett 601 (8 Feb 1887)

+ Mangifera foetida Lour.

+ Mangifera griffithii Hook.f.

* Mangifera magnifica Kochummen (Mangifera quadrifida)

* Mangifera paludosa Kosterm. (Mangifera aff. parvifolia)

* ~Melanochyla auriculata Hook.f. - Ridley 3975 (1892)

*+ Melanochyla caesia (Blume) Ding Hou (Melanochyla kunstleri)

+ Parishia maingayi Hook.f.

Ancistrocladaceae

Ancistrocladus tectorius (Lour.) Merr. - NRS1523

Anisophylleaceae

* Anisophyllea disticha (Jack) Baill. - Maxwell 82-101 (7 Apr 1982)

Annonaceae

Artabotrys costatus King - NRS1358

Artabotrys suaveolens (Blume) Blume - Maxwell 82-70 (18 Mar 1982)

*+ Cyathocalyx ramuliflorus (Maingay ex Hook.f. & Thomson) Scheff.

* ~Cyathostemma viridiflorum Griff. - Maxwell 80-211 (28 Nov 1980)

+ Desmos dasymaschala (Blume) Saff.

* Desmos dumosus (Roxb.) Saff. - Maxwell 81-57 (1 Apr 1980)

* Friesodielsia biglandulosa (Blume) Steenis (Oxymitra biglandulosa) - Sinclair SFN40271 (1

May 1954)

* Friesodielsia borneensis (Miq.) Steenis (Oxymitra borneensis)

* Friesodielsia glauca (Hook.f. & Thomson) Steenis (Oxymitra glauca)

Friesodielsia latifolia (Hook.f. & Thomson) Steenis - Goodenough s.n. (189?)

Goniothalamus macrophyllus (Blume) Hook.f. & Thomson - NRS1171

* Goniothalamus ridleyi King - NRSO160

141

Mezzettia parviflora Becc. (Mezzettia leptopoda)

Mitrella kentii (Blume) Mig. - NRSO119

* Polyalthia angustissima Ridl.

*+ Polyalthia glauca (Hassk.) Muell.

Polyalthia hypoleuca Hook.f. & Thomson

Polyalthia lateriflora (Blume) King

Polyalthia sclerophylla Hook.f. & Thomson

Popowia tomentosa Maingay ex Hook.f. & Thomson (Popowia hirta)

Pyramidanthe prismatica (Hook.f. & Thomson) Merr.

Uvaria cordata (Dunal) Alston - NRSO120

* Uvaria leptopoda (king) R.E. Fries

* Xylopia caudata Hook.f. & Thomson

*+ Xylopia fusca Maingay ex Hook.f. & Thomson

*+ Xylopia malayana Hook.f. & Thomson - DJM100

* * * &

Apocynaceae

Alstonia angustifolia Wall. ex A.DC.

Alstonia angustiloba Miq.

Alstonia spatulata Blume

Anodendron candolleanum Wight

Kibatalia maingayi (Hook.f.) Woodson

Kopsia singapurensis Ridl. - NRSO0126

Leuconotis griffithii Hook.f. - Ridley s.n. (1894)

Leuconotis maingayi Dyer ex Hook.f. - NRS0169

* Parameria polyneura Hook.f.

Parsonsia stenocarpa King & Gamble - Maxwell 80-208 (28 Nov 1980)

*#—- * # —r

* Tabernaemontana corymbosa Roxb.

* Urceola torulosa Hook.f.

* Urnularia flavescens (Dyer ex Hook.f.) Stapf

* Willughbeia coriacea Wall. - NRS0048

Aquifoliaceae

* Ilex cymosa Blume - Ridley s.n. (23 Sep 1890)

* [lex macrophylla Hook.f.

Araceae

+ Aglaonema nebulosum N.E.Br. - NRSO161

*+ Aglaonema nitidum (Jack) Kunth - NRS0766

Aglaonema simplex Blume - NRS0868

*+ Alocasia denudata Engl. - Ridley s.n. (30 Oct 1889)

* Anadendrum montanum (Blume) Schott - NRS2418

+ Cryptocoryne griffithii Schott - NRS1525

¥ Cyrtosperma merkusii (Hassk.) Schott - NRSO889

Homalomena griffithii (Schott) Hook.f. - Ridley s.n. (May 1889)

Homalomena sagittifolia Jungh. ex Schott - NRS2419

* Pothos latifolius Hook.f.

Rhaphidophora korthalsii Schott - NRS1591

Rhaphidophora sylvestris (Blume) Engl. var. montana (Blume) Nicols. - NRS0992

Schismatoglottis wallichii (Roxb.) Hook.f. - NRS1604

Scindapsus hederaceus (Zoll. & Moritzi) Mig. - NRS1509

142

Araliaceae

Arthrophyllum diversifolium Blume (Arthrophyllum ovalifolium)

* Schefflera cephalotes (C.B. Clarke) Harms

Asclepiadaceae

Dischidia albiflora Griff. (Dischidia collyris)

Dischidia complex Griff.

Dischidia hirsuta (Blume) Decne.

Dischidia major (Vahl) Merr. (Dischidia rafflesiana)

Dischidia nummularia R.Br.

Toxocarpus griffithii Decne. (Toxocarpus glabrescens)

Fe 9h OE

Bignoniaceae

* — Deplanchea bancana (Scheff.) Steenis

Bombacaceae

* Coelostegia griffithii Benth.

* Durio singaporensis Ridl.

* — Neesia malayana Bakh. - Ridley 3770 (1892)

Neesia synandra Mast. - NRS0996

Burseraceae

Canarium littorale Blume - Ridley 44 (Nov 1889)

* ~Canarium odontophyllum Mig. - Corner’s specimen (s.n. 25 Apr 1934) taken from a 12 foot

sapling is the only record of this species from the Malay Peninsula.

* ~Canarium pilosum Benn.

* ~Dacryodes incrvata (Engl.) H.J. Lam

* Dacryodes rostrata (Blume) Lam

* — Santiria apiculata Benn.

* Santiria griffithii (Hook.f.) Engl.

* Santiria laevigata Blume

*+ Santiria rubiginosa Blume

+ Triomma malaccensis Hook.f.

Caprifoliaceae

Viburnum sambucinum Blume - Ridley 6829

Cecropiaceae

Poikilospermum suaveolens (Blume) Merr. - NRS1020 CCA 18

Celastraceae

*+ Bhesa paniculata Arn. - NRSO170

+ Bhesa robusta (Roxb.) Ding Hou

* Euonymus javanicus Blume

*+ Lophopetalum multinervium Ridl.

+ Lophopetalum wightianum Arn.

Salacia grandiflora Kurz - NRSO128

143

Chrysobalanaceae

Auna racemosa Rafin. - Ridley 1898 (7 Dec 1890)

Liania splendens (Korth.) Prance - Ridley 4732 (15 Feb 1893)

* Parastemon urophyllus (Wall. ex A. DC.) A. DC.

* Parinari costata (Korth.) Blume

* Parinari oblongifolia Hook.f.

Combretaceae

Combretum sundaicum Mia. - Mat 6783 (1985)

Quisqualis indica L. - Ridley 8053 (1896)

* Terminalia phellocarpa King

Commelinaceae

+ Amischotolype gracilis (Ridl.) I.M. Turner - NRSO0179

Compositae

Vernonia arborea Buch.-Ham. - Goodenough 2739 (1891)

Connaraceae

* Agelaea borneensis (Hook.f.) Merr. - Maxwell 82-107 (28 May 1981)

* Cnestis palala (Lour.) Merr.

Connarus ferrugineus Jack - Maxwell 83-26 (13 Mar 1983)

Connarus grandis Jack - J. Sinclair SFN40321 (3 Jul 1954)

* Connarus monocarpus L. ssp. malayanus Leenhouts

* Connarus semidecandrus Jack

Rourea fulgens Planch. - NRSO0860

* Rourea minor (Gaertn.) Leenhouts

Convallariaceae

+ Peliosanthes teta Andr. ssp. humilis (Andr.) Jessp. - NRS1169

Convolvulaceae

* Argyreia ridleyi Prain ex Ooststr. - NRSOOO1 CCA 13

* Erycibe griffithii C.B. Clarke

Erycibe tomentosa Blume var. tomentosa - NRSO181

Cornaceae

+ Mastixia trichotoma Blume - Maxwell 82-258 (23 Jun 1982)

Cucurbitaceae

Trichosanthes celebica Cogn. - Maxwell 80-176 (31 July 1980)

Cyperaceae

Hypolytrum nemorum (Vahl) Spreng. - NRS1176

Mapania cuspidata (Miq.) Uittien - NRS1620

Mapania enodis (Miq.) C.B. Clarke

Mapania lorea Uittien

Mapania squamata (Kurz) C.B. Clarke - NRS1505

Thoracostachyum bancanum (Miq.) Kurz

Cad) ae ae a

144

Dilleniaceae

* Acrotrema costata Jack - no herbarium material of this species collected in Singapore can be

found.

Dillenia grandifolia Wall. ex Hook.f & Thomson

Dillenia pulchella (Jack) Gilg

Tetracera akara (Burm.f.) Merr.

Tetracera arborescens Jack

Tetracera indica (Christm. & Panz.) Merr. - NRS0184

* * +

Dioscoreaceae

Dioscorea laurifolia Wall. ex Hook.f. - NRS0042

Dioscorea polyclades Hook.f. - Mat s.n. (15 May ?)

Dipterocarpaceae

* Hopea mengerawan Miq. - Goodenough 5094 (8 Apr 1893)

* Shorea gratissima Dyer

*+ Shorea macroptera Dyer

+ Shorea ovalis Blume (S. ?eximia)

*+ Shorea platycarpa Heim

* — Vatica pauciflora Blume (Vatica wallichii) - Hill & Samsuri H3001 (22 Oct 1971)

+ Vatica ridleyana Brandis

Dracaenaceae

Dracaena aurantiaca Wall. - NRS0139

Dracaena elliptica Thunb. - Ridley s.n. (1894)

Dracaena singapurensis Ridl. - Ridley 6235 (1894)

Dracaena umbratica Ridl. - NRS1167

Ebenaceae

* Diospyros coriacea Hiern

*+ Diospyros lanceifolia Roxb. - Ridley 6695 (1894)

*+ Diospyros maingayi (Hiern) Bakh.

*+ Diospyros pilosanthera Blanco var. oblonga (Wall. ex G. Don) Ng (Diospyros oblonga) - Hill

& Samsuri H3025 (22 Oct 1971)

* Diospyros siamang Bakh.

Diospyros sumatrana Miq. - Maxwell 82-73 (18 Mar 1982)

Elaeocarpaceae

Elaeocarpus acmosepalus Rid. - Maxwell 83-25 (13 Mar 1983)

Elaeocarpus ferrugineus (Jack) Steud. - Goodenough 4948 (24 Feb 1890)

Elaeocarpus floribundus Blume - Sidek 605 (18 Mar 1982)

* Elaeocarpus griffithii (Wight) A. Gray - Ridley 418 (Nov 1889)

* Elaeocarpus mastersii King - Ridley 274 (Nov 1889)

Elaeocarpus nitidus Jack var. salicifolius (King) Ng - Samsuri Ahmad 1522 (17 Sept 1981)

+ Elaeocarpus petiolatus (Jack) Wall.

* Elaeocarpus polystachyus Wall. - Maxwell 81-214 (24 Sept 1981)

Elaeocarpus stipularis Blume - Maxwell 82-69 (18 Mar 1982)

Euphorbiaceae

* Agrostistachys longifolia (Wight) Benth. ex Hook.f. (Agrostistachys sessilifolia) - NRSO18

145

Alchornea villosa Miill.Arg.

Antidesma coriaceum Tul. - Maxwell 81-60 (1 Apr 1981)

Antidesma cuspidatum Miill.Arg. - NRS0856

Antidesma neurocarpum Mig. - NRSO116

Aporusa confusa Gage - NRS1359

Aporusa frutescens Blume - NRSO193

+ Aporusa penangensis (Ridl.) Airy Shaw

Aporusa symplocoides (Hook.f.) Gage - NRS0876

* Austrobuxus nitidus Miq.

+ Baccaurea hookeri Gage

* Baccaurea kunstleri King ex Gage

+ Baccaurea minor Hook.f.

+ Baccaurea parviflora (Miill.Arg.) Miill.Arg.

+ Baccaurea racemosa (Reinw.) Miill.Arg.

*+ Blumeodendron tokbrai (Blume) Kurz

* Bridelia griffithii Hook.f.

* Croton laevifolius Blume - NRSO872

* Drypetes pendula Rid].

* Endospermum diadenum (Migq.) Airy Shaw

* Glochidion hypoleucum (Migq.) Boerl. - Maxwell 82-181 (1-Jul 1982)

* Glochidion rubrum Blume

* Glochidion sericeum (Blume) Zoll. & Moritzi - Maxwell 82-53 (26 Feb 1982)

Glochidion superbum Baill. - NRSO007

* Hymenocardia sp. [the correct identity of this species has still not been determined]

*+ Macaranga conifera (Zoll.) Miill.Arg.

* Macaranga gigantea (Rchb. f. & Zoll.) Miill.Arg.

* Macaranga griffithiana Miill.Arg.

* Macaranga hypoleuca (Rchb.f. & Zoll.) Miill.Arg.

* ~Macaranga punticulata Gage

* Macaranga recurvata Gage

* Macaranga trichocarpa (Rchb.f. & Zoll.) Miill.Arg. - Maxwell 81-81 (29 Apr 1981)

*+ Macaranga triloba (Reinw.) Miill.Arg. - NRS0037

+ Neoscortechinia kingii (Hook.f.) Pax & K. Hoffm. - NRS0771

*+ Pimelodendron griffithianum (Miill.Arg.) Benth.

* Sapium discolor (Champ. ex Benth.) Miill.Arg.

Fagaceae

* Castanopsis inermis (Linfl. ex Wall.) Benth. & Hook.f.

+ Castanopsis nephelioides King ex Hook.f.

Castanopsis wallichii King ex Hook.f. - Ridley 3842 (Jun 1892)

*+ Lithocarpus bennettii (Miq.) Rehder - E. Tang & Sidek 265 (12 Jan 1995)

* Lithocarpus elegans (Blume) Hatus. ex Soepadmo

7 Lithocarpus sundaicus (Blume) Rehder

Flacourtiaceae

Casearia capitellata Blume - Ridley s.n. (1892)

Casearia lobbiana Turcz. - Ridley s.n. (1894)

Casearia sp. - the specimen cited by Corner (SFN32517) is Campnosperma squamatum,

presumably the wrong number was referred to.

Flacourtia rukam Zoll. & Moritzi - NRS1015

146

* Osmelia maingayi King

{+ Osmelia philippina (Turcz.) Benth. - Maxwell 83-18 (17 Feb 1983)

Flagellariaceae

* Flagellaria indica L. - NRSO015

Gesneriaceae

Aeschynanthus albidus (Blume) Steud. (= A. purpurascens Hassk.) - NRS1012

Aeschynanthus parvifolius R.Br.

* — Aeschynanthus wallichii R.Br.

Gonystylaceae

* Gonystylus confusus Airy Shaw

+ Gonystylus maingayi Hook.f.

Gramineae

* — Leptaspis urceolata (Roxb.) R.Br. - NRS0039

Lophatherum gracile Brongn. - Goodenough 1701 (8 Feb 1890)

Guttiferae

+ Calophyllum dispar P.F. Stevens

* Calophyllum ferrugineum Ridl. (Calophyllum retusum)

* ~Calophyllum rubiginosum M.R. Hend. & Wyatt-Sm. (C. wallichianum)

*+ Calophyllum tetrapterum Mig. (Calophyllum floribundum)

*+ Calophyllum teysmannii Mig. (Calophyllum inophylloide var. singapurense)

* ~Calophyllum wallichianum Planch. & Triana var. incrassatum (M.R. Hend. & Wyatt-Sm.) P.F.

Stevens (Calophyllum incrassatum)

* ~Cratoxylum arborescens (Vahl) Blume

* ~~ Cratoxylum cochinchinense (Lour.) Blume (Cratoxylum ligustrinum) - NRSO172

* Cratoxylum formosum (Jack) Dyer

+ Garcinia eugeniifolia Wall. ex T. Anderson - Ridley s.n. (1893)

*+ Garcinia forbesii King - NRS0887

Garcinia griffithii T. Anderson - DJM101

2? Garcinia hombroniana Pierre - Ridley s.n. (1893)

Garcinia mangostana L. - Ridley s.n. (1892)

+ Garcinia nervosa Mid. - J. Sinclair SFN10915 (6 Sep 1966)

* Garcinia nigrolineata Planch. ex T. Anderson

*+ Garcinia parvifolia (Mig.) Mig. - Ridley s.n. (Nov 1909)

* Garcinia scortechinii King (Garcinia gaudichaudii)

Hanguanaceae

* —Hanguana malayana (Jack) Merr. - NRSO035

Hypoxidaceae

Curculigo latifolia Dryand. - NRS0025 CCA 13

Icacinaceae

* Cantleya corniculata (Becc.) Howard

Gomphandra quadrifida (Blume) Sleumer var. ovalifolia (Ridl.) Sleumer - NRSO777

Todes cirrhosa Turcz. - NRS0026

147

Iodes ovalis Blume - NRS0004

* Phytocrene bracteata Wall.

* Phytocrene latifolia Blume

*7 Stemonurus scorpioides Becc. - Ridley s.n. (1892)

ervaars

* Irvingia malayana Oliv.

Ixonanthaceae

Ixonanthes icosandra Jack - Ridley 1937 (1890)

Ixonanthes reticulata Jack - NRSO0783

Lauraceae

* Actinodaphne macrophylla (Blume) Nees

+ Alseodaphne bancana Mia. - Samsuri 1401 (9 Feb 1977)

*+ Beilschmiedia kunstleri Gamble

Cryptocarya ferrea Blume - NRS1005

Cryptocarya griffithiana Wight

Endiandra maingayi Hook.f. (Hexapora ? curtisii)

Lindera lucida (Blume) Boerl. (Lindera malaccensis) - Sinclair SFN40520 (22 Dec 1954)

Litsea elliptica Blume - Samsuri 1446 (9 Feb 1977)

Litsea erectinervia Kosterm.

Litsea ferruginea Blume

Litsea firma Hook.

Litsea gracilipes Hook.f. - Sinclair SFN40274 (1 May 1954)

Litsea grandis Hook.f. - Ridley 3962 (1892)

Litsea lancifolia (Roxb.) Hook.f. - Maxwell 82-78 (19 Mar 1982)

Litsea ridleyi Gamble - NRS1183

7 Litsea robusta Blume

7 Phoebe grandis Mert.

+ eH ml

* # # =

Leeaceae

Leea indica (Burm.f.) Mert. - NRS0046

Leguminosae

* Albizia splendens Mia. (Pithecellobium confertum)

Archidendron clypearia (Jack) 1.C. Nielsen - NRS0421

*+ Archidendron ellipticum (Blume) LC. Nielsen (Pithecellobium ellipticum) - Maxwell 81-146

(18 Jun 1981)

Archidendron microcarpum (Benth.) I.C. Nielsen - Samsuri Ahmad 1399 (9 Feb 1977)

Bauhinia semibifida Roxb. - NRS0990

Crudia caudata Prain [we have not been able to trace the specimen referred to by Comer]

* Dalbergia havilandii Prain

Dalbergia pseudosissoo Mia. - J. Sinclair SFN39999 (2 Aug 1953)

Derris amoena Benth. var. maingayana (Baker) Prain - NRSO118 CCA 18

Derris trifoliata Lour. - Goodenough s.n. (30 Oct 1889)

* Dialium patens Baker

Entada spiralis Ridl. - Samsuri Ahmad 1451 (8 Mar 1977)

Koompassia malaccensis Maingay ex Benth.

Kunstleria ridleyi Prain

148

Milletia eriantha Benth. (Adinobotrys eriantha)

Ormosia macrodisca Baker

Parkia speciosa Hassk. - Ridley 419 (Nov 1889)

Saraca thaipingensis Cantley - NRS0779

Sindora wallichii Grah. ex Benth.

Spatholobus ridleyi Prain ex King - Mat 6795 (1894)

Linaceae

Indorouchera griffithiana (Planch.) Hallier f. - NRSO155

Loganiaceae

Fagraea auriculata Jack

Fagraea racemosa Jack ex Wall. - NRS1027

Loranthaceae

Macrosolen cochinchinensis (Lour.) Tiegh. - Maxwell 82-257 (23 Sept 1982)

Magnoliaceae

Magnolia candollii (Blume) H. Keng var. singapurensis (Ridl.) Nooteboom (Talauma

singapurensis, T: lanuginosa) - Maxwell 81-145 (18 Jun 1981)

Malpighiaceae

Aspidopteris concava (Wall.) Juss. - Maxwell 82-46 (17 Feb 1982)

Melastomataceae

Clidemia hirta (L.) D. Don. - NRS0043

Diplectria viminalis (Jack) Kuntze (Anplectrum viminale)

Macrolenes echinulata (Naud.) Bakh.f. (Marumia rhodocarpa)

Medinilla crassifolia (Reinw. ex Blume) Blume (?Medinilla rubicunda var. hasseltii) - Ridley

273 (Oct 1889)

Memecylon edule Roxb.

Memecylon floridum Ridl. (Memecylon sp.)

Memecylon lilacinum Zoll. & Moritzi

Pachycentria constricta (Blume) Blume (Pachycentria tuberculata)

Pternandra coerulescens Jack - NRSO754

Pternandra echinata Jack - NRSO192

Sonerila heterostemon Naud. - NRSO0778

Meliaceae

Aglaia odoratissima Blume - Maxwell 82-282 (4 Nov 1982)

Aglaia oligophylla Mia. - Ridley 3919 (1892)

Aglaia rubiginosa (Hiern) Pannell - Ridley 3790 (1892)

Aphanamixis polystachya (Wall.) Parker - Maxwell 82-282 (2 Jul 1981)

Dysoxylum cauliflorum Hiern. - Goodenough 5080 (1893)

Dysoxylum cyrtobotryum Migq. - Goodenough s.n. (Mar 1890)

Dysoxylum densiflorum (Blume) Miq.

Dysoxylum excelsum Blume - Goodenough s.n. (15 Mar 1890)

Dysoxylum flavescens Hiern - J. Sinclair SFN40262 (23 Apr 1954)

Lansium domesticum Correa

Pseudoclausena chrysogyne (Miq.) T.P. Clark - Maxwell 81-199 (13 Aug 1981)

149

*+ Sandoricum beccarianum Baill. (Sandoricum emarginatum)

Menispermaceae

Fibraurea tinctoria Lour. - NRS0143

Limacia scandens Lour. - NRSO147

Stephania capitata (Blume) Spreng. - NRSO159

Tinospora macrocarpa Diels - NRS0849

Monimiaceae

+ Kibara coriacea (Blume) Tul.

Matthaea sancta Blume - NRS1003

Moraceae

* Artocarpus elasticus Reinw. ex Blume - Chew CWL34 (5 Mar 1957)

Artocarpus fulvicortex Jarrett - Ridley 4129 (1892)

*+ Artocarpus kemando Miq. - Sinclair SFN40322 (3 Jul 1954)

Artocarpus nitidus Tréc. ssp. griffithii (King) Jarrett

Artocarpus scortechinii King

Ficus albipila (Mig.) King

Ficus apiocarpa Miq. - NRS2128

Ficus binnendykii Mia. var. coriacea Corner

Ficus bracteata Wall. ex Miq.

Ficus consociata Blume var. murtoni King - E. Tang & Sidek 785 (10 Jul 1995)

Ficus excavata King

Ficus fistulosa Reinw. ex Blume var. fistulosa - NRSO010

Ficus globosa Blume - NRS0183

Ficus heteropleura Blume - NRS1180

Ficus microcarpa L..

Ficus microsyce Ridl.

Ficus obscura Blume var. borneensis (Miq.) Corner

Ficus pellucido-punctata Griff.

Ficus pisocarpa Blume - DJM318

Ficus recurva Blume var. ribesioides King

Ficus retusa L.

Ficus ruginervia L.

Ficus sagittata Vahl -

Ficus scortechinii King - NRS1174

Ficus sinuata Thunb. - NRSO186

Ficus sundaica Blume - NRS0014

Ficus trichocarpa Blume - DJM171

Ficus variegata Blume - Ridley s.n. (189?)

* Ficus villosa Blume - NRS1618

Ficus virens Ait. var. glabella (Blume) Corner - Ridley 1602 (30 Oct 1889)

* Ficus xylophylla Wall. ex Miq.

Parartocarpus bracteatus (King) Becc. - Ridley 4128 (1892)

Parartocarpus venenosus (Zoli. & Moritzi) Becc. ssp. forbesii (King) Jarrett

Streblus elongatus (Miq.) Corner

* + © © 4H HH HH & &

* * & +

Myristicaceae

* Endocomia canarioides (King) W.J. de Wilde (Horsfieldia macrocoma vat. canarioides)

150

*+ Gymnacranthera bancana (Miq.) J. Sinclair

*+ Gymnacranthera farquhariana (Hook.f. & Thomson) Warb. (Gymnancranthera eugeniifolia var.

griffithii) - Ridley 1834 (9 Dec 1891)

Gymnacranthera forbesii (King) Warb. - NRS1190

*+ Horsfieldia crassifolia (Hook.f. & Thomson) Warb. - Ridley 8040 (Feb 1896)

* Horsfieldia grandis (Hook.f.) Warb.

*+ Horsfieldia polyspherula (Hook.f.) J. Sinclair (Horsfieldia subglobosa)

*+ Horsfieldia sucosa (King) Warb. (Horsfieldia bracteosa)

* Horsfieldia superba (Hook.f. & Thomson) Warb.

+ Horsfieldia wallichii (Hook.f. & Thomson) Warb. - Ridley 5061 (1893)

+ Knema communis J. Sinclair - Ridley 1833

*+ Knema conferta (King) Warb.

*+ Knema curtisii (King) Warb. var. paludosa J. Sinclair

* Knema glaucescens Jack

+ Knema hookeriana (Wall. ex Hook.f. & Thomson) Warb.

* Knema intermedia (Blume) Warb.

+ Knema latericia Elm. - NRSO869

+ Knema laurina (Blume) Warb.

*+ Knema malayana Warb. - Maxwell 81-87 (29 Apr 1981)

*+ Myristica cinnamomea King - Sinclair SFN40374 (25 Sep 1954)

*+ Myristica elliptica Wall. ex Hook. f. & Thomson - NRS0049

*t Myristica iners Blume

*+ Myristica lowiana King

Myristica maxima Watb. - Sinclair SFN 40304 (22 May 1954)

Myrsinaceae

+ Ardisia colorata Roxb. - NRSO018

Ardisia miqueliana Scheff. - NRSO767

Ardisia sessilis Scheff. - Ridley 3844 (Jun 1892)

Ardisia teysmanniana Scheff. - Maxwell 82-147 (23 Apr 1982)

* Ardisia tuberculata Wall. ex A.DC.

Ardisia villosa Roxb. - NRS0883

Embelia canescens Jack - NRSO0947

* Embelia coriacea Wall. ex A. DC.

* Embelia fulva Mez (E. ?amentacea)

Labisia pumila (Blume) Fern.-Vill. - NRS1186

* — Maesa ramentacea Wall. ex Roxb. - Maxwell 81-240 (3 Dec 1981)

Myrtaceae

* Aphanomyrtus rostrata Miq. (Pseudoeugenia singaporensis) - NRSO878

* Decaspermum fruticosum J.R. Forst. & G. Forst - DJM174

*+ Eugenia cerina M.R. Hend. - Samsuri Ahmad 1396 (9 Feb 1977)

Eugenia chlorantha Duthie - Ridley 3910

* Eugenia cumingiana Vidal

Eugenia filiformis Duthie var. clavimyrtus (Koord. & Valeton) M.R. Hend.

Eugenia grandis Wight - Maxwell 81-103 (28 May 1981)

* Eugenia kunstleri King

Eugenia leptostemon (Korth.) Miq. - Mat s.n. (5 May 1895)

* Eugenia longiflora (Presl) Fern.-Vill.

* Eugenia microcalyx Duthie

151

Eugenia muelleri Miq.

Eugenia nemestrina M.R. Hend. - Corner SFN37396 (28 Dec 1940)

Eugenia oblongifolia Duthie - Ridley 360 (Nov 1889)

Eugenia oleina Wight

Eugenia pachyphylla Kurz - NRS0138

Eugenia papillosa Duthie

Eugenia polyantha Wight - NRS0562

Eugenia pseudocrenulata M.R. Hend.

Eugenia pseudosubtilis King - Goodenough 4487 (21 Dec 1889)

Eugenia pustulata Duthie - Mat 6806 (15 May 1895)

Eugenia ridleyi King - Mat 6805 (15 May 1895)

Eugenia rugosa (Korth.) Merr.

Eugenia tumida Duthie - Maxwell 81-210 (24 Sep 1981)

Rhodamnia cinerea Jack - Goodenough s.n. (10 Feb 1890)

Tristaniopsis merguensis (Griff.) Peter G. Wilson & J.T. Waterh.

Najadaceae

Najas indica (Willd.) Cham. - NRS0838

Nepenthaceae

Nepenthes ampullaria Jack

Nepenthes gracilis Korth.

Nepenthes rafflesiana Jack

Nymphaeaceae

Barclaya motleyi Hook.f. - Sinclair SFN40337 (17 July 1954)

Ochnaceae

Brackenridgea hookeri (Planch.) A. Gray

Brackenridgea palustris Bartell. - Ridley 5896 (1894)

Olacaceae

;

Erythropalum scandens Blume - NRS0845

Ochanostachys amentacea Mast.

Strombosia ceylanica Gardn. - NRS1946

Oleaceae

Chionanthus ramiflorus Roxb. (Linociera pauciflora)

Jasminum aemulum R.Br. - Mohd. Shah & Ali MS4153 (3 Dec 1981)

Orchidaceae

* + & & &

Agrostophyllum bicuspidatum J.J.Sm.

Anoectochilus geniculatus Ridl. - Ridley 5896 (1894)

Appendicula lucida Ridl.

Biermannia laciniata (Carr) Garay (Chamaeanthus laciniatus)

Bulbophyllum acuminatum (Ridl.) Ridl.

Bulbophyllum apodum Hook.f. - Ridley 3933 (1892)

Bulbophyllum blumei (Lindl.) J.J.Sm. - Mat s.n. (1894)

Bulbophyllum botryophorum Ridl.

Bulbophyllum gusdorfii J.J.Sm.- NRS1683

Bulbophyllum macranthum Lindl. - NRS1320

Bulbophyllum medusae (Lindl.) Rchb.f. - Goodenough s.n. (1891)

Bulbophyllum ovalifolium (Blume) Lindl. - Mat s.n. (1894)

* Bulbophyllum pileatum Lindl.

* Bulbophyllum pulchellum Ridl. - Ridley s.n. (19 Dec 1890)

Bulbophyllum purpurascens Teijsm. & Binnend.

Bulbophyllum sessile (Koenig) J.J.Sm. - Ridley s.n. (1892)

Bulbophyllum vaginatum (Lindl.) Rchb.f.

Calanthe pulchra (Blume) Lindl. - Chew & Whitmore s.n. (5 Mar 1957)

Ceratostylis subulata Blume (Camerotis adnata)

Claderia viridiflora Hook.f. - NRSO002

Coelogyne mayeriana Rchbf. - Ridley s.n. (1891)

Corymborkis veratrifolia (Rienw.) Blume - Mat 6756 (1894)

Cymbidium finlaysonianum Lidl. - Ridley s.n. (1891)

Cystorchis variegata Blume - Sinclair SFN40277 (8 May 1954)

* Dendrobium aloifolium (Blume) Rchb.f.

* Dendrobium concinnum Migq. (Dendrobium carnosum) - Ridley s.n. (1892)

* Dendrobium crumenatum Sw.

Dendrobium indivisum (Blume) Mia. - Ridley s.n. (1892)

* Dendrobium leonis (Lindl.) Rchb.f. - Ridley s.n. (30 Oct 1889)

Dendrobium planibulbe Lindl. - Goodenough s.n. (29 Aug 1890)

* Dendrobium prostratum Ridl.

Dendrobium pulchellum Roxb. ex Lindl. - Ridley s.n. (May/Jun 1890)

Dendrobium setifolium Ridl. - Goodenough s.n. (21 Dec 1889)

Dendrobium spegidoglossum Rchb.f. - Ridley s.n. (1894)

* Dendrobium spurium (Blume) J.J.Sm.

* Dendrochilum longifolium Rchb.f.

Didymoplexis pallens Griff. - Ridley s.n. (1889)

Eria bractescens Lindl. - Ridley s.n. (1890)

Eria floribunda Lindl. - Goodenough s.n. (13 Jan 1892)

Eria neglecta Ridl.

Eria nutans Lindl.

Eria pannea Lindl. - Goodenough s.n. (1891)

Eria pulchella Lindl. - Ridley s.n. (1892)

Eulophia spectabilis (Dennst.) Suresh (Eulophia squalida) - Ridley s.n. (1892)

Flickingeria comata (Blume) Hawkes (Dendrobium comatum)

Flickingeria fimbriata (Blume) Hawkes (Dendrobium plicatile)

Galeola nudifolia Lour. (Galeola kuhlii) - Ridley 4037 (28 Oct 1892)

Hylophila mollis Lindl. - Ridley s.n. (Oct 1889)

Lecanorchis malaccensis Ridl. - NRS1533

Lipparis gibbosa Finet - Goodenough s.n. (1891)

Lipparis tricallosa Rchb.f. - Goodenough s.n. (13 Jan 1892)

* Lipparis wrayi Hook.f.

Malaxis latifolia J.E.Sm. - Ridley 3922 (1892)

Malaxis micrantha (Hook.f.) Kuntze - Ridley s.n.

Microsaccus javensis Blume - Ridley s.n. (1894)

Nephelaphyllum pulchrum Blume - NRSO775

* Nervilia punctata (Blume) Makino

Oberonia stenophylla Ridl. - Corner 393 (Dec 1930)

Plocoglottis gigantea (Hook.f.) J.J.Sm. - NRSO122

*+ Plocoglottis javanica Blume - NRS0999

Ee Sey Oe OE:

ec, a a

e—- e KK He &

153

Podochilus microphyllus Lindl. - Goodenough s.n. (15 Oct 1889)

Pteroceras pallidum (Ridl.) Holttum - NRS1935

* Schoenorchis secundiflora (Ridl.) J.J.Sm.

* Taeniophyllum filiforme J.J.Sm.

* Taeniophyllum obtusum Blume - NRS1318

* Thelasis micrantha (Brongn.) J.J.Sm.

* Thelasis pygmaea (Griff.) Lindl. (Thelasis triptera)

Thrixspermum trichoglottis (Hook.f.) Kuntze - NRS1319

* Trichotosia velutina (Lodd. ex Lindl.) Kraenzl. (Eria velutina)

Vrydagzynea lancifolia Ridl. - Ridley s.n. (Jan 1890)

Oxalidaceae

* Dapania racemosa Korth.

+ Sarcotheca griffithii (Planch. ex Hook.f.) Hallier f.

Palmae

Calamus ridleyanus Becc. - Hill & Samsuri 3108 (22 Oct 1971)

* Cyrtostachys renda Blume (Cyrtostachys lakka)

Daemonorops angustifolia (Griff.) Mart.- NRS0031

Daemonorops didymophylla Becc. - Ridley 3476 (1892)

Daemonorops hystrix (Griff.) Mart. - Ridley s.n.

Daemonorops kunstleri Becc. - Goodenough 1667 (21 Jun 1890)

* Daemonorops leptopus (Griff.) Mart.

Daemonorops longipes (Griff.) Mart. - NRS2424

Daemonorops sabut Becc. - Sinclair SFN40303 (22 May 1954)

Eleiodoxa conferta (Griff.) Burret (Salacca conferta) - Ridley 3502 (1892)

Iguanura wallichiana (Wall. ex Mart.) Hook.f. - NRSO177

Korthalsia echinometra Becc. - NRS1175

Licuala ferruginea Becc. - NRS1177

Myrialepis paradoxa (Kurz) J. Dransf. - Ridley 3503 (1892)

* Nenga pumila (Mart.) Wendl. - Ridley s.n. (1900)

* Onchosperma horridum (Griff.) Scheff. - Ridley s.n. (1891)

Orania sylvicola (Griff.) H.E. Moore - Ridley 3144 (1891)

Pinanga malaiana (Mart.) Scheff. - NRSO135

Rhopaloblaste singaporensis (Becc.) Hook.f. - Ridley 2134 (1891)

Pandaceae

* Galearia fulva (Tul.) Miq. - NRSO881

Pandanaceae

* Freycinetia angustifolia Blume - Samsuri 1451 (8 Mar 1977)

* Freycinetia confusa Ridl.

* Freycinetia corneri B.C. Stone

* Frecinetia javanica Blume - NRS1527

* Pandanus atrocarpus Griff.

* Pandanus kamiae B.C. Stone

Passifloraceae

Adenia macrophylla (Blume) Koord. var. singaporeanna (Wall. ex G. Don) W.J. de Wilde

-NRS1028 CCA 18

154

Phormiaceae

* Dianella ensifolia (L.) DC. - NRS0281 CCA 18

Piperaceae

* Piper flavimarginatum C. DC.

* Piper miniatum Blume

Piper muricatum Blume - NRS1187

Polygalaceae

*+ Xanthophyllum affine Korth. - NRSO0995

+ Xanthophyllum ellipticum Korth.

+ Xanthophyllum obscurum Benn.

+ Xanthophyllum vitellinum (Blume) Dietr.

Rhamnaceae

Ventiiago malaccensis (Lour.) Merr. - Maxwell 81-64 (1 Apr 1981)

* Ziziphus elegans Wall.

Rhizophoraceae

*+ Carallia brachiata (Lour.) Merr. - Maxwell 81-102 (28 May 1981)

*t+ Gynotroches axillaris Blume - Sidek & Ali SK620 (9 Dec 1982)

*+ Pellacalyx axillaris Korth. - NRS0146

Rosaceae

* Prunus arborea (Blume) Kalkman

* Prunus grisea (C. Muell.) Kalkman

+ Prunus polystachya (Hook f.) Kalkman - NRSO191

Rubiaceae

* Aidia corymbosa (Blume) K.M. Wong (Randia auriculata)

Aidia wallichiana Tirveng. - Ridley 5662 (1892)

* Caelospermum truncatum (Wall.) Baill. ex K. Schum.

Canthium horridum Blume - NRS0044

Chasalia chartacea Craib - NRS1025

Coptosapelta griffithii Hook.f. - Mat 6891 (1895)

+ Diplospora malaccensis Hook.f. - NRSO763

Gardenia griffithii Hook.f. - Hullett 611 (21 Aug 1888)

Gynochthodes sublanceolata Mig. - Samsuri Ahmad 1521 (17 Sep 1981)

Hedyotis capitellata Wall. ex G. Don - DIM166

Hedyotis congesta R.Br. ex G. Don - DJM139

* Hydnophytum formicarum Jack

Ixora coccinea Hook.f. - Mat 6714 (1894)

* Ixora congesta Roxb.

Ixora lobbii King & Gamble - NRS2410

Ixora pendula Sack - Ridley 5676

*+ Jackiopsis ornata (Wall.) Ridsdale (Jackia ornata)

Lasianthus appressus Hook.f. or L. attenuatus Jack - NRSO998

Lasianthus attenuatus Jack - NRSO023

Lasianthus constrictus Wight - NRS0034

Lasianthus densifolius Mig. - NRSO756

155

* Lasianthus griffithii Wight - NRS1949

Lasianthus ridleyi King & Gamble - Maxwell 82-100 (7 Apr 1982)

* Lecananthus erubescens Jack

Lucinaea membranaea King - NRS0862

Morinda citrifolia L. - NRSO167

Morinda ridleyi (King & Gamble) Rid]. - Maxwell 81-235 (3 Dec 1981)

Morinda rigida Miq.

Morinda umbellata L. - NRS2411

Mussaenda glabra Vahl - NRS0047

*+ Mussaendopsis beccariana Baill. - NRS0991

Myrmecodia tuberosa Jack

Ophiorrhiza singaporensis Ridl. - NRSO165

Oxyceros scandens (Blume) Tirveng. (Randia clarkei)

Pavetta wallichiana Steud. (Pavetta indica) - NRS0759

Porterandia anisophylla (Jack ex Roxb.) Ridl. - NRSO0852

Prismatomeris glabra (Korth.) Valeton - Ridley 6150 (Apr 1894)

Psychotria sp. 9 - Hullett s.n. (1896)

* Psychotria maingayi Hook.f.

* Psychotria obovata Wall.

Psychotria penangensis Hook.f. - NRSO770

Psychotria sarmentosa Blume - NRSO173

* Rothmannia macrophylla (R.Br. ex Hook.f.) Bremek. (Randia macrophylla)

* Tarenna adpressa (King) Merr.

Tarenna odorata (Roxb.) B.L. Rob. - NRS0871

* Timonius flavescens (Jack) Baker - Maxwell 81-82 (29 Apr 1981)

* Timonius wallichianus (Korth.) Valeton - NRSO857

* Uncaria acida (Hunter) Roxb. (Uncaria ovalifolia)

Uncaria cordata (Lour.) Merr. - DJM172

Uncaria gambir (Hunter) Roxb. - Maxwell 82-50 (17 Feb 1982)

Uncaria lanosa Wall. var. glabrata (Blume) Ridsdale

Uncaria longiflora (Poir.) Merr. var. pteropoda (Miq.) Ridsdale - Ridley s.n.

Urophyllum blumeanum (Wight) Hook.f. - NRS0752

* Urophyllum glabrum Wall. - NRSO162

Urophyllum sp. 2 of Wong (Tree Flora of Malaya Vol. 4) - NRSO176

* Urophyllum streptopodium Wall. ex Hook.f.

+ —- +

Rutaceae

Glycosmis chlorosperma (Blume) Spreng. - Ridley 3912 (1892)

Luvunga crassifolia Tanaka - NRS0635

Maclurodendron porteri (Hook.f.) T.G. Hartley - Hill & Samsuri H3017 (22 Oct 1971)

Sapindaceae

* Lepisanthes fruticosa (Roxb.) Leenh.

Nephelium cuspidatum Blume var. eriopetalum (Miq.) Leenh.

* Nephelium lappaceum L. (N. glabrum) - NRSO006

* Nephelium laurinum Blume - Ridley 6211 (Apr 1894)

*+ Pometia pinnata Forst. - Sinclair SFN40519 (22 Dec 1954)

Xerospermum noronhianum Blume - Ridley s.n. (1894)

156

Sapotaceae

Ganua motleyana (de Vr.) Pierre ex Dubard

Madhuca malaccensis (C.B. Clarke) Lam - Mat 6500 (1894)

Palaquium hexandrum (Griff.) Baill.

Palaquium gutta (Hook.f.) Baill. - Hill & Samsuri H3038 (22 Oct 1971)

*+ Palaquium rostratum (Miq.) Burck

*+ Palaquium xanthochymum (de Vr.) Pierre - Ridley 3774 (1892)

* Planchonella maingayi (C.B. Clarke) van Royen

Smilacaceae

Smilax setosa Miq. - NRSO129

Smilax calophylla Wall. - NRS0997

Sterculiaceae

Byttneria maingayi Mast. - Maxwell 82-59 (26 Feb 1982)

Commersonia bartramia (L.) Merr. - NRS1018

* Heritiera elata Ridl.

Scaphium linearicarpum (Mast.) Pierre - Himat (?) s.n. (11 Oct 1956)

+ Scaphium macropodum (Miq.) Beumee ex Heyne

* Sterculia coccinea Jack - NRSO121

+ Sterculia cordata Blume

* Sterculia gilva Miq. (Stercula bicolor)

*+ Sterculia macrophylla Vent. - Samsuri Ahmad 1455 (8 Mar 1977)

*+ Sterculia rubiginosa Vent. - Samsuri Ahmad 1395 (9 Feb 1977)

Symplocaceae

* ~Symplocos barringtoniifolia Brand - Maxwell 82-180 (1 July 1982)

+ Symplocos fasciculata Zoll.

Taccaceae

* — Tacca integrifolia Ker-Gawl. - Goodenough s.n. (14 Apr 1890)

Theaceae

Adinandra acuminata Korth. - Ridley s.n. (1894)

Adinandra dumosa Jack - NRSO198

+ Eurya acuminata DC. - NRSOOS2

Gordonia penangensis Ridl. - Ridley 3913 (1892)

+ Pyrenaria acuminata Planch. ex Choisy - NRS1016

Thymelaeaceae

*+ Aquilaria malaccensis Lam.

Enkleia malaccensis Griff. - DJM153

Tiliaceae

* Grewia acuminata Juss. (Grewia umbellata) - NRSO185

*+ Microcos blattifolia (Corner) Rao (Grewia blattifolia)

*+ Pentace triptera Mast.

Ulmaceae

Gironniera parvifolia Planch.

157

Verbenaceae

Clerodendrum deflexum Wall. - NRS1009

Clerodendrum laevifolium Blume - NRS0758

Clerodendrum paniculatum L. - NRS0846

Clerodendrum villosum Blume - NRS0005

Vitex pinnata L. - NRS0136

Viscaceae

Viscum articulatum Burm.f. - Ridley 8054 (1896)

Viscum ovalifolium Wall. ex DC. - Ridley s.n. (1896)

Vitaceae

Ampelocissus elegans (Kurz) Gagnep. - NRS0011

Ampelocissus gracilis (Wall.) Planch. - NRS1170

Cayratia mollissima (Wall.) Gagnep. (Vitis mollissima) - DJIM1696

Cissus hastata (Miq.) Planch. (Vitis hastata) - NRSO187

Cissus nodosa Blume (Vitis glaberrima) - Ridley 6789 (7 Aug 1895)

-Cissus repens Lam. - NRS0884

Pterisanthes eriopoda (Migq.) Planch. - Ridley 425 (Nov 1889)

Pterisanthes polita (Miq.) Laws. - NRS0127

Zingiberaceae

Amomum hastilabium Ridl.

Amomum xanthophlebium Baker - Ridley s.n. (1899)

Etlingera punicea (Roxb.) R.M. Sm. - Maxwell 81-101 (28 May 1981)

Globba leucantha Mia. - Betty Allen s.n. (28 May 1949)

Hornstedtia leonurus (Koenig) Retz. - Ridley s.n. (1892)

Hornstedtia scyphifera (Koenig) Steud. -

Plagiostachys lateralis (Rid\.) Ridl. - Ridley s.n. (1892)

Zingiber griffithii Baker - NRSO787

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<4) ;

A Preliminary Survey of Ferns and Fern-Allies of

Gunung Kajang Area, Pulau Tioman

M. Zulkifli, A. Latiff, A.A. Bidin and R. Jaman

Department of Botany,

Faculty of Life Sciences,

Universiti Kebangsaan Malaysia,

43600 UKM Bangi, Selangor, Malaysia.

Abstract

A total of 149 specimens of ferns and fern-allies were collected during two trips to Gunung Kajang, Pulau

Tioman. They were identified to 24 families, 59 genera, 95 species, 1 subspecies and 11 varieties, excluding 2

species whose identity were uncertain. Asplenium contributes the largest number among genera, with 9 species,

followed by Selaginella (6 species) and Selliguea (4 species), and 39 species were identified as new records to

Pulau Tioman fern flora.

Introduction

Gunung Kajang (1030 m.) which is situated in the south-eastern part of Pulau

Tioman is the highest peak of the island (Figure 1). There is a trail from Kampung

Paya, inland of Tanjung Pauh on the south-west coast cutting several small streams

to the summit of Gunung Kajang. The trail runs upwards along ridges and valleys

in a generally west by south-east direction. The small ridge known as Bukit

Permatang runs from east to west and one small cave found on the way to the

summit at 950 m. elevation is known as Gua Tahi Angin. Another trail to the summit

of Gunung Kajang runs from Kampung Juara at Teluk Juara on the other side of the

island has been demolished because of the dangerous slopes on the north and south

banks.

The geology of the Gunung Kajang area is dominated by granitoid rocks which

give the rocky structure such as exposed granite rock and thin layer of soil. Exposed

rock is found near the sea at Kampung Paya and at several places along the trail to

the summit of Gunung Kajang.

Two trips of survey have been made to collect and identify the ferns and fern-

allies from Kampung Paya to the summit of Gunung Kajang and several meters

down from the summit to Teluk Juara. Most of the samples were collected in the

form of herbarium specimens except several of them were only observed and noted.

Voucher specimens were kept in the Herbarium, Universiti Kebangsaan Malaysia

(UKMB).

Survey of Pulau Tioman Ferns

Several collections of plants of Pulau Tioman were made by various collectors.

In August 1889, Ridley collected a little on the west coast at the place he called

Nypa Bay (Henderson, 1930). In June 1915, Burkill and Robinson collected at

160

Juara Bay and Tanjung Duata (Burkill, 1927) while Henderson (1930) collected

mainly on higher altitude especially on Gunung Kajang and Gunung Rokam between

May 10th and May 29th, 1927. It was until 1977 when Lee and members of Universiti

Malaya made a list of ferns and fern-allies of Pulau Tioman. A total of 112 species

from 23 families and 62 genera have been recorded from Pulau Tioman and Pulau

Tulai from their collections and previous records of herbarium specimens. In this

preliminary survey, a total of 149 specimens were collected and classified into 24

families, 59 genera and 95 species which include 39 new records for Pulau Tioman

(Table 1).

Table 1 List of ferns and fern-allies collected from Gunung Kajang, including

previous records

et all. Records

LYCOPODIACEAE

Selaginella alutacia

Selaginella wallichi a -

Selaginella willdenovii mi

PSILOTACEAE ee

Psilotum complanatum +

ADIANTACEAE

Acrostichum aureum

Acrostichum speciosum

Adiantum caudatum

Adiantum stenochlamys

Pityrogramma calomelanos

Lee

et al.

Taenitis blechnoides

Taenitis dimorpha

ASPLENIACEAE

161

Zulkifli Lee New

et al. et all. Records

FAMILY / Species

Asplenium affine

Asplenium batuense

Asplenium borneense

Asplenium macrophyllum

Asplenium montanum

Asplenium nidus

Asplenium nitidum

Asplenium normale

Asplenium robustum

Asplenium salignum

Asplenium scolopendrioides

Asplenium sublaserpitiifolium

Asplenium tenerum

Asplenium unilaterale

BLECHNACEAE

| Blechnumfinlaysonianum | | |

Es es

| Stenochleanapalustris | + | |

CHEIROPLEURIACEAE

Cheiropleuria bicuspis ~

CYATHEACEAE

[Cyatheacriophora—SSSS*dC

F Cyatheahymenodes———SSS—«d

[Cyathea obscura ——SSCSC*~dSCSt

[Araiostegiahymenophylloides |

[Davalladivaricaa ———SC~d

162

et al.

MEE MORad Di. 1

Histiopterisincisa Sid

[Lindsaeadoryphora it

Tapeiniivon luzonicur var luzonicum [4

“Tapeinidiumpinnaum Sd

Wet ees

Dipterisconjugaa ———SSSSS~dC

ann en

Ditdymochlaenatruncaudla =

Pleocnemiaconjugaa———SSS=dS

[Polystichum prolficans Si

[Preridryssyrmatica——SSSC~dC

[“Tectaria grandidentata———SSS~dSC

Mane aan

[Dicranopteriscurranii SSCs

New

Records

Lee

et al.

163

FAMILY / Species Zulkifli Lee

et all. :

Dicranopteris linearis var. subpectinata

Diplopterygium longissimum

Gleichenia microphylla var. semivestita

Sticherus truncatus var. truncatus

GRAMMITIDACEAE

Ctenopteris blechnoides

et al

Grammitis adspersa

Prosaptia celebica

Prosaptia leysti

Scleroglossum debile

+/+ ]+

Tee

=e ew

[ Microgonium bimarginauum ——~(| ++i SiC

Nesopterissuperba——=SCS*~‘dSSC~‘“dtCSCC*d

Faaatae ees

te 5 1 ae

Vandenboschia maxima

164

Zulkifli

et al.

FAMILY / Species Li New

et al. Records

ia’)

MARATTIACEAE

Angiopteris angustifolia

Angiopteris evecta

NEPHROLEPIDACEAE

Nephrolepis auriculata

Nephrolepis davallioides

Nephrolepis falcata

Nephrolepis hirsutula

OLEANDRACEAE

Oleandra pistillaris - -

[ee a

| OPHIOGLOSSACEAE 5)

| Helminthostachyszeylanica | + | + |

| POLYPODIACEAE | | |) ee

Colysis macrophylla +

Colysis pedunculata

Colysis pteropus

Drynaria sparsisora

Goniophlebium korthalsii

Goniophlebium sp.

Loxogramme avenia

Loxogramme subecostata

Microsorum heterocarpum

Phymatosorus nigrescens

Pyrrosia lanceolata

Pyrrosia rasamalae

Selliguea enervis

Selliguea heterocarpa

Selliguea taeniata var. palmata

Selliguea triloba

PTERIDACEAE

Pteris ensiformis

Pteris grevilleana

FAMILY / Species

Pteris mertensioides

Pteris oppositipinna

Pteris tripartita

Pteris vittata

SCHIZAEACEAE

Lygodium circinatum

Lygodium flexuosum

Lygodium microphyllum

Schizaea dichotoma

Schizaea digitata

THELYPTERIDACEAE | |

Amphineuron interruptum

Pronephrium repandum + ee:

Pronephrium rubicundum | - |

Sphaerostephanos heterocarpus -

VITTARIACEAE

Antrophyum callifolium - ~

Monogramma dareicarpa -

Vittaria angustifolia - ~

Vittaria scolopendrina - ~

WOODSIACEAE

Diplazium sorzogonense |

Diplazium tomentosum ~ ~

Total number of species 95 112 39

165

New

Records

On the coastal area at Kampung Paya in Tanjung Pauh, Nephrolepis auriculata,

Stenochlaena palustris and Pteris ensiformis dominate the rocks in open places.

There are several others like Asplenium nidus, A. macrophyllum and Pyrrosia

166

lanceolata which were found on rocks by the beach, whereas Lygodium flexuosum

can be easily seen scrambling over the bushes nearby. A single individual of

Helminthostachys zeylanica was observed in open ground.

The trail from Kampung Paya runs across several small streams under primary

forest of Gunung Kajang foothill. One small stream known as Sungai Ayer Pertama

is dominated with a species, Bolbitis sinuata, on rock and terrestrial in flood area

of the stream. Along streams, Bolbitis appendiculata ssp. appendiculata grows in

similar habitat with the above species. Other rock ferns such as Cephalomanes

Javanicum grow on expose rock surface in the stream.

Most lowland ferns grow on the forest floor similar to that of the lowland ferns

in similar habitat in the mainland. The forest floor is dominated by Selaginella

wallichi, Selaginella wildenovii and Colysis pedunculata among others. Another

small stream called Ayer Suring shows the same fern flora but no Bolbitis sinuata

found. Microsorum heterocarpum grows on rock in shady area. Holttum (1968)

recorded that this species has the south limit not further than the Main Range.

Riparian ferns such as Pleocnemia conjugata_and Pteridrys syrmatica are found

along streams but not in large groups, were new records for Pulau Tioman. Holttum

(1968), noted that Pleocnemia conjugata was less common in Peninsular Malaysia,

probably correlated with its lowland habitat and preference for lightly shaded places

which do not occur in normal fully shaded primitive forest.

Another notable place called Ayer Orang Puteh was furnished with naked surface

of rocks with a small stream that flows underneath, giving a wet and moist habitat

which is a suitable place for rock-loving ferns especially members of Hymeno-

phyllaceae such as Cephalomanes javanicum, Meringium acanthoides, Microgo-

nium bimarginatum and Selenodesmium obscurum. Other lithophytic ferns found

were Asplenium affine, A. borneense, A. scolopendrioides, Antrophyum callifolium,

Colysis macrophylla and Loxogramme avenia. Ayer Orang Puteh at the altitude of

650 m also the lowest limit of Cyatheaceae found in Gunung Kajang area; Cyathea

eriophora was found on the bank near rocky part of Ayer Orang Puteh.

The area upwards of Air Orang Puteh shows more diversity in fern flora. Out of

95 species of ferns and fern-allies collected from Gunung Kajang area, 49 species

(52 %) were collected in the very limited area above 700 m. Bukit Permatang (850

m.) was familiar with ferns of Cyatheaceae and Dennsteadtiaceae. Two species of

Cyathea i.e. Cyathea hymenodes and Cyathea obscura were new records to Pulau

Tioman. Histiopteris incisa grows on rock with a thin layer of humus. Asplenium

robustum which grows on high branches of trees in Main Range, especially at

Cameron Highlands, was found growing on rock at Bukit Permatang. This species

is now rare (Piggot, 1988).

Along the trail reaching the summit, ferns of Gleicheniaceae were found growing

in several places. Two species of Dicranopteris and one species each from Diplo-

pterygium, Gleichenia and Sticherus were found in this area but not found elsewhere

at the lower altitude. The summit of Gunung Kajang (1030 m.) is characterised

with the present of Cheiropleuria bicuspis which only grows in lower montane

167

forest. Holttum (1968), noted that specimen from Pulau Tioman has a special form

with four-lobed sterile fronds, while our collection showed the form with two-

lobed sterile fronds which is the common form of this species. Ferns of family

Grammitidaceae which grow on mossy branch of trees and rocks also characterised

the summit of Gunung Kajang. Among them were Prosaptia celebica and Prosaptia

leysii which were new records of Pulau Tioman ferns. Another notable fern species

at the summit was Histiopteris incisa which colonised the landslide area on the

north banks of the summit.

Checklist of Ferns and Fern-Allies of the Gunung Kajang Area

FERN-ALLIES

1. LYCOPODIACEAE

1.1 Huperzia Bernh.

One species found in Gunung Kajang area.

1.1.1 Huperzia phlegmaria (L.) Rothm.

Lycopodium phlegmaria L.

Epiphyte. Lower montane forest. Elevation: 1000-1030 m.

Material: SUMMIT TRAIL: Elevation 1000 m, Zulkifli & R. Jaman, RJ 2921 (UKMB): SUMMIT:

Elevation 1030 m, Zu/kifli & R. Jaman, RJ 2936 (UKMB).

1.2 Lycopodiella Holub

One species found in Gunung Kajang area, restricted mostly near the summit of the

hill.

1.2.1 Lycopodiella cernua (L.) Pic. Serm.

Lycopodium cernuum Linn.

Terrestrial. Upper hill dipterocarp to lower montane forest. Elevation: 950-1000 m.

Material: SUMMIT TRAIL: Elevation ca. 950 m, Zulkifli & R. Jaman, RJ 2888 (UKMB); ca.1000 m

elevation, RJ 2939 (UKMB).

2. SELAGINELLACEAE

2.1. Selaginella Beauv.

A total of 8 species recorded for Pulau Tioman and 6 from the Gunung Kajang area,

mostly in shaded area.

168

Kx... Gua Tahi Angin

G Kajor

ons

Figure 1. Map of Pulau Tioman, showing collection trail (....) to the summit of

Gunung Kajang and several previous collecting localities

169

2.1.1 Selaginella alutacia Spring

Lithophyte. Upper hill dipterocarp forest. Elevation: 780-940 m.

Material: GUA TAHI ANGIN TRAIL: Elevation ca. 780 m, Zulkifli & R. Jaman, RJ 2899 (UKMB); Elevation

ca. 940 m, Zulkifli & R. Jaman, RJ 2966 (UKMB).

2.1.2 Selaginella delicatula (Desv.) Alston

Terrestrial. Upper hill dipterocarp forest.

Material: BUKIT PERMATANG: Elevation ca. 780 m, Zulkifli & R. Jaman, RJ 2918 (UKMB).

2.1.3 Selaginella intermedia (B1.) Spring var. intermedia

Selaginella atroviridis Spring

Terrestrial. Upper hill dipterocarp forest.

Material: SUMMIT TRAIL: Elevation ca. 920 m, Zulkifli & R. Jaman, RJ 2924 (UKMB).

2.1.4 Selaginella selangorensis Bedd. ex Ridl.

Terrestrial. Upper hill dipterocarp forest to lower montane forest. Elevation 950-

1000 m.

Material: SUMMIT TRAIL: Elevation ca. 1000 m, Zulkifli & R. Jaman, RJ 2898 (UKMB); Elevation ca.

950 m, Zulkifli & R. Jaman, RJ 2935 (UKMB).

2.1.5 Selaginella wallichii (Hook. & Grev.) Spring

Terrestrial. Hill dipterocarp forest. Elevation 350-880 m.

Material:SUNGAI AIR PERTAMA TRAIL: Elevation ca. 380 m, Zulkifli & R. Jaman, RJ 2829 (UKMB);

Elevation ca. 350 m, Zulkifli & R. Jaman, RJ 2832 (UKMB); BUKIT PERMATANG: Elevation ca. 880 m,

Zulkifli & R. Jaman, RJ 2877 (UKMB).

2.1.6 Selaginella willdenovii( Desv. ) Baker

Terrestrial, forming thicket. Hill dipterocarp forest.

Material: SUNGAI AIR PERTAMA TRAIL: Elevation ca. 380 m, Zulkifli & R. Jaman, RJ 2830 (UKMB).

FERNS

3. ADIANTACEAE

3.1 Taenitis Willd

This genus has never been recorded from Pulau Tioman before. Two species were

found in Gunung Kajang area.

170

3.1.1 Taenitis blechnoides (Willd.) Sw.

Terrestrial. Upper hill dipterocarp forest to lower montane forest. Elevation:

940- 1030 m.

Material: SUMMIT TRAIL: Elevation ca. 940 m, Zulkifli & R. Jaman, RJ 2969 (UKMB); SUMMIT: Elevation

ca. 1030 m, Zulkifli & R. Jaman, RJ 2882 (UKMB).

3.1.2 Taenitis dimorpha Holttum

Terrestrial in semi-shade. Lower montane forest. Elevation: 1020-1030 m.

Material: SUMMIT: Elevation ca. 1030 m, Zulkifli & R. Jaman, RJ 2886 (UKMB); Elevation ca. 1020 m,

Zulkifli & R. Jaman, RJ 2960 (UKMB).

4. ASPLENIACEAE

4.1 Asplenium L.

This genus contributed the largest number of species among the fern families of

Gunung Kajang. A total of nine species were identified with two of them, A. nidus and

A. robustum being new records for Pulau Tioman.

4.1.1 Asplenium affine Sw.

Asplenium spathulinum J. Sm.

On semi-shaded rock. Hill dipterocarp forest.

Material: AYER ORANG PUTEH: Elevation ca. 600 m, Zulkifli & R. Jaman, RJ 2861 (UKMB).

4.1.2 Asplenium borneense Hook.

On semi-shaded rock. Hill dipterocarp forest.

Material: AYER ORANG PUTEH: Elevation ca. 600 m, Zulkifli & R. Jaman, RJ 2856 (UKMB).

4.1.3 Asplenium macrophyllum Sw.

On exposed rock. Coastal vegetation.

Material: KAMPONG PAYA: Elevation ca. 1 m, Zulkifli & R. Jaman, RJ 2817 (UKMB).

4.1.4 Asplenium nidus L.

On exposed rock. Coastal vegetation.

Material: KAMPONG PAYA: Elevation ca.1 m, Zulkifli & R. Jaman, RJ 2818 (UKMB).

4.1.5 Asplenium normale D.Don

On shaded rock. Hill dipterocarp forest.

Material: SUMMIT TRAIL: Elevation ca. 700 m, Zulkifli & R. Jaman, RJ 2920 (UKMB).

171

4.1.6 Asplenium robustum Bl.

On shaded rock. Upper hill dipterocarp forest.

Material: BUKIT PERMATANG: Elevation ca. 850 m, Zulkifli & R. Jaman, RJ 2864 (UKMB).

Note: Piggot (1988), noted that this species was commonly observed as epiphytes

on higher branches of trees in Cameron Highlands but is now difficult to find.

4.1.7 Asplenium scolopendrioides J. Sm. ex Hook.

On semi-shaded rock. Hill dipterocarp forest. Elevation: 460-600 m.

Material: AYER SURING TRAIL: Elevation ca. 460 m, Zulkifli & R. Jaman, RJ 2834 (UKMB); AYER

SURING: Elevation ca. 480 m, Zulkifli & R. Jaman, RJ 2839 (UKMB); AYER ORANG PUTEH: Elevation

ca. 600 m, Zulkifli & R. Jaman, RJ 2857 (UKMB).

4.1.8 Asplenium sublaserpitiifolium Ching

On semi-shaded rock. Hill dipterocarp forest. Elevation: 520 m.

Material: AYER ORANG PUTEH TRAIL: Elevation ca. 520 m, Zulkifli & R. Jaman, RJ 2845 (UKMB).

Note: This species probably is the second collection in Malaysia after a single

specimen collected by Burkill in June 1915 from Teluk Juara, Pulau Tioman, which

was described in Holttum (1968). The occurence of the species on Pulau Tioman is

thus an interesting extension of range as the species described by Ching were

collected from the neighbouring parts of Southern China.

4.1.9 Asplenium unilaterale Lam.

On semi-shaded rock. Hill dipterocarp forest.

Material: AYER SURING: Elevation ca. 480 m, Zulkifli & R. Jaman, RJ 2848 (UKMB); Zulkifli & R.

Jaman, RJ 2852 (UKMB).

5. BLECHNACEAE

5.1 Blechnum L.

A genus of six species in Peninsular Malaysia. This genus was not previously recorded

from Pulau Tioman. The shade loving-fern, Blechnum finlaysonianum was found on

forest floor in hill dipterocarp forest.

5.1.1 Blechnum finlaysonianum Hook. & Grev.

Terrestrial in shade. Hill dipterocarp forest.

Material: SUNGAI AYER PERTAMA TRAIL: Elevation ca. 350 m, Zulkifli & R. Jaman, RJ 2914 (UKMB).

5.2 Stenochlaena J. Sm.

A genus with only one species recorded from Malaysia and has not been recorded

before from Pulau Tioman.

5.2.1 Stenochlaena palustris (Burm. f.) Bedd.

Climber in open situations. Coastal vegetation.

Material: KAMPONG PAYA: Elevation ca.1 m, Zulkifli & R. Jaman, RJ 2821 (UKMB).

6. CHEIROPLEURIACEAE

6.1 Cheiropleuria C. Pres|

The only species in the genus recorded in Malaysia. It is a species found near the

summit of Gunung Kajang.

6.1.1 Cheiropleuria bicuspis (B1.) C. Presl

Terrestrial in shade. Lower montane forest.

Material: SUMMIT TRAIL: Elevation ca. 1020 m, Zulkifli & R. Jaman, RJ 2961 (UKMB).

7. CYATHEACEAE

7.1 Cyathea Sm.

A total of 7 species recorded for Pulau Tioman, 3 were new records, found in Gunung

Kajang area.

7.1.1 Cyathea eriophora Holttum

Terrestrial in semi-shade. Hill dipterocarp forest.

Material: AYER ORANG PUTEH: Elevation ca. 650 m, Zulkifli & R. Jaman, RJ 2879 (UKMB).

7.1.2 Cyathea hymenodes Mett.

Cyathea latebrosa (Wall.)Copel. var. indusiata Holttum

Terrestrial in light shade. Upper hill dipterocarp forest. Elevation: 850-950 m.

Material: BUKIT PERMATANG: Elevation ca. 850 m, Zulkifli & R. Jaman, RJ 2946 (UKMB), Elevation

ca. 880 m, Zulkifli & R. Jaman, RJ 2872 (UKMB); SUMMIT TRAIL TO TELUK JUARA: Elevation ca. 950

m, Zulkifli & R. Jaman, RJ 2933 (UKMB).

7.1.3 Cyathea obscura (Scort.) Copel.

Terrestrial in open situations. Upper hill dipterocarp forest to lower montane

forest. Elevation: 850-1020 m.

Material: BUKIT PERMATANG: Elevation ca. 850 m, Zulkifli & R. Jaman, RJ 2927 (UKMB); SUMMIT

TRAIL: Elevation ca. 1020 m, Zulkifli & R. Jaman, RJ 2959 (UKMB).

8. DAVALLIACEAE

8.1 Araiostegia Copel.

Only one species recorded from Malaysia. On Pulau Tioman this species was found at

Bukit Permatang.

8.1.1 Araiostegia hymenophylloides (B\.) Copel.

On rock in semi-shade. Upper hill dipterocarp forest.

Material: BUKIT PERMATANG: Elevation ca. 800 m, Zulkifli & R. Jaman, RJ 2863 (UKMB).

8.2 Davallia Sm.

One species found from three species recorded from Pulau Tioman.

8.2.1 Davallia divaricata Bl.

On rock in light shade. Upper hill dipterocarp forest.

Material: BUKIT PERMATANG: Elevation ca.850 m, Zulkifli & R. Jaman, RJ 2865 (UKMB).

8.3 Humata Cay.

Two species were found in Gunung Kajang area.

8.3.1 Humata angustata (Wall. ex Hook. & Grev.) J. Sm.

Terrestrial and climber, lithophyte and on old stump in semi-shade. Upper hill

dipterocarp forest to lower montane forest. Elevation: 780-1020 m.

Material: SUMMIT TRAIL: Elevation ca. 1000 m, Zu/kifli & R. Jaman, RJ 2887 (UKMB), Elevation 1000

m, RJ 2890 (UKMB), Elevation ca. 1020 m, Zulkifli & R. Jaman, RJ 2963 (UKMB); BUKIT PERMATANG

TRAIL: Elevation ca. 780 m, Zulkifli & R. Jaman, RJ 2919 (UKMB).

8.3.2 Humata repens (L. f.) Diels

Epiphyte in light expose. Upper hill dipterocarp forest to lower montane forest.

Elevation: 820-1030 m.

Material: BUKIT PERMATANG: Elevation ca. 820 m, Zulkifli & R. Jaman, RJ 2868 (UKMB); SUMMIT

TRAIL: Elevation ca. 1000 m, Zulkifli & R. Jaman, RJ 2889 (UKMB); SUMMIT: Elevation ca.1030 m,

Zulkifli & R. Jaman, RJ 2929 (UKMB).

9 DENNSTEADTIACEAE

9.1 Microlepia C. Pres

One species found in Gunung Kajang area.

174

9.1.1 Microlepia speluncae (L.) T. Moore var. villosissima C. Chr.

Terrestrial in light shade. Hill dipterocarp forest.

Material: SUNGAI AYER PERTAMA TRAIL: Elevation ca. 350 m, Zulkifli & R. Jaman, RJ 2833 (UKMB).

9.2 Histiopteris (Agardh) J. Sm.

One species found from two different habitats, i.e. terrestrial and rock.

9.2.1 Histiopteris incisa (Thunb.) J. Sm.

Terrestrial and lithophyte in open situations. Upper hill dipterocarp forest to

lower montane forest. Elevation: 840-1030 m.

Material: SUMMIT: Elevation ca. 1030 m, Zulkifli & R. Jaman, RJ 2883 (UKMB); BUKIT PERMATANG:

Elevation ca. 840 m, Zulkifli & R. Jaman, RJ 2956 (UKMB).

9.3. Lindsaea Dryander

Three species were found in Gunung Kajang area from five species recorded for Pulau

Tioman. One sterile specimen cannot be identified.

9.3.1 Lindsaea doryphora Kramer

Lindsaea scandens Hooker var. terrestris Holttum

Terrestrial in light shade. Hill dipterocarp forest to upper hill dipterocarp forest.

Elevation: 580-880 m.

Material: AYER ORANG PUTEH TRAIL: Elevation ca. 580 m, Zulkifli & R. Jaman, RJ 2847 (UKMB);

BUKIT PERMATANG: Elevation ca. 820 m, Zulkifli & R. Jaman, RJ 2875 (UKMB), Elevation ca. 880 m,

Zulkifli & R. Jaman, RJ 2878 (UKMB).

9.3.2 Lindsaea parasitica (Roxb. ex Griff.) Hieron.

Lindsaea scandens Hooker

Climber in shade. Upper hill dipterocarp forest. Elevation: 750-850 m.

Material: BUKIT PERMATANG TRAIL: Elevation ca. 750 m, Zulkifli & R. Jaman, RJ 2869 (UKMB);

BUKIT PERMATANG: Elevation ca. 850 m, Zulkifli & R. Jaman, RJ 2876 (UKMB).

9.3.3. Lindsaea repens (Bory) Thwaites var. pectinata (B1.) Mett. ex Kuhn

Epiphyte in semi-shade. Lower montane forest.

Material: SUMMIT TRAIL: Elevation ca. 1000 m, Zulkifli & R. Jaman, RJ 2942 (UKMB).

9.3.4 Lindsaea sp.

On rock and scrambling over small bushes in shade. Hill dipterocarp forest.

Sterile.

Material: AYER ORANG PUTEH TRAIL: Elevation ca. 550 m, Zulkifli & R. Jaman, RJ 2851 (UKMB).

175

9.4 Tapeinidium (C. Presl) C. Chr.

Two species are identified and 7: Juzonicum var. luzonicum is a new record for Pulau

Tioman.

9.4.1 Tapeinidium luzonicum (Hook.) Kramer var. luzonicum

Terrestrial in shade. Upper hill dipterocarp forest.

Material: SUMMIT TRAIL: Elevation ca. 960 m, Zulkifli & R. Jaman, RJ 2964 (UKMB).

9.4.2 Tapeinidium pinnatum (Cav.) C. Chr.

Terrestrial in shade. Upper hill dipterocarp forest.

Material: BUKIT PERMATANG: Elevation ca. 880 m, Zulkifli & R. Jaman, RJ 2874 (UKMB).

10. DIPTERIDACEAE

10.1 Dipteris Reinw.

One species found on summit of Gunung Kajang.

10.1.1 Dipteris conjugata Reinw.

Terrestrial in open situations. Lower montane forest.

Material: SUMMIT: Elevation ca.1030 m, Zulkifli & R. Jaman, RJ 2885 (UKMB).

11. DRYOPTERIDACEAE

11.1 Didymochlaena Desv.

Only one species occurs in Gunung Kajang area.

11.1.1. Didymochlaena truncatula (Sw.) Sw.

Terrestrial in shade. Hill dipterocarp forest.

Material: AYER ORANG PUTEH TRAIL: Elevation ca. 520 m, Zulkifli & R. Jaman, RJ 2842 (UKMB).

11.2 Pleocnemia C. Presl

One species is identified from Gunung Kajang area and is a new record for Pulau

Tioman.

11.2.1 Pleocnemia conjugata (B1.) C. Pres

Terrestrial in semi-shade. Lowland dipterocarp forest to hill dipterocarp forest.

Elevation: 210-560 m.

176

Material: SUNGAI AYER PERTAMA TRAIL: Elevation ca. 210 m, Zulkifli & R. Jaman, RJ 2827 (UKMB);

AYER SURING TRAIL: Elevation ca. 560 m, Zulkifli & R. Jaman, RJ 2906 (UKMB).

Note: Holttum (1968), noted that this species was collected twice in Peninsular

Malaysia, once near Melaka and once at Baling, Kedah.

11.3 Polystichum Roth.

This genus has never been recorded previously from Pulau Tioman.

11.3.1 Polystichum prolificans Alderw.

Terrestrial and lithophyte in light shade. Upper hill dipterocarp forest. Elevation:

800-880 m.

Material: BUKIT PERMATANG: Elevation ca. 800 m, Zu/kifli & R. Jaman, RJ 2862 (UKMB); SUMMIT

TRAIL: Elevation ca. 880 m, Zulkifli & R. Jaman, RJ 2871(UKMB).

11.4 Pteridrys C. Chr & Ching

One species found in Gunung Kajang area.

11.4.1 Pteridrys syrmatica (Willd.) C. Chr. & Ching

Terrestrial and lithophyte in light shade. Lowland dipterocarp forest. Elevation:

160-300 m.

Material: SUNGAI AYER PERTAMA TRAIL: Elevation ca. 160 m, Zulkifli & R. Jaman, RJ 2833 (UKMB);

Elevation ca. 210 m, Zulkifli & R. Jaman, RJ 2826 (UKMB); SUNGAI AYER PERTAMA: Elevation ca. 300

m, Zulkifli & R. Jaman, RJ 2909 (UKMB).

11.5 Tectaria Cav.

One species found out of two species recorded on Pulau Tioman.

11.5.1 Tectaria grandidentata (Cesati) Holttum

Terrestrial in shade. Hill dipterocarp forest.

Material: AYER ORANG PUTEH TRAIL: Elevation ca. 550 m, Zulkifli & R. Jaman, RJ 2846 (UKMB).

12; GLEICHENIACEAE

12.1 Dicranopteris Bernh.

Two species were found in Gunung Kajang. Dicranopteris curranii is anew record for

Pulau Tioman.

177

12.1.1 Dicranopteris curranii Copel.

Terrestrial and scrambling in exposed lower montane forest.

Material: SUMMIT: Elevation ca. 1030 m, Zulkifli & R. Jaman, RJ 2912 (UKMB).

12.1.2 Dicranopteris linearis (Burm. f.) Underw. var. montana Holttum

Terrestrial and scrambling in light-shaded lower montane forest.

Material: SUMMIT TRAIL: Elevation ca. 1000 m, Zulkifli & R. Jaman, RJ 2934 (UKMB).

12.1.3 Dicranopteris linearis (Burm. f.) Underw. var. subpectinata (H. Christ)

Holttum

Terrestrial in expose. Upper hill dipterocarp forest to lower montane forest.

Elevation: 950-1000 m.

Material: SUMMIT TRAIL: Elevation ca. 950 m, Zulkifli & R. Jaman, RJ 2931 (UKMB); Elevation ca.1000

m, Zulkifli & R. Jaman, RJ 2940 (UKMB).

12.2 Diplopterygium (Diels) Nakai

12.2.1 Diplopterygium longissimum (B1.) Nakai

Gleichenia longissima Bl.

Scrambling in expose. Lower montane forest.

Material: SUMMIT TRAIL: Elevation ca.1000 m, Zulkifli & R. Jaman, RJ 2881 (UKMB).

12.3 Gleichenia Sm.

One species was identified from Gunung Kajang.

12.3.1 Gleichenia microphylla R. Br. var. semivestita (Labill.) v.A.v.R.

Terrestrial in open situationse. Lower montane forest.

Material: SUMMIT TRAIL: Elevation ca. 1000 m, Zulkifli & R. Jaman, RJ 2932 (UKMB).

12.4 Sticherus Pres|

12.4.1 Sticherus truncatus (Willd.) Nakai var. truncatus

Gleichenia truncata (Willd.) Spring var. truncata; Gleichenia laevigata

(Willd.) Hk.

Terrestrial in light-shaded lower montane forest.

Material: SUMMIT: Elevation ca.1030 m, Zulkifli & R. Jaman, RJ 2938 (UKMB).

178

13. GRAMMITIDACEAE

13.1 Ctenopteris Bl. ex Kunze

One species was identified from Gunung Kajang.

13.1.1 Ctenopteris blechnoides (Grev.) W.H. Wagner & Grether

Ctenopteris moultonii (Copel.) C. Chr. & Tard.

Epiphyte and lithophyte in light shade. Upper hill dipterocarp forest to lower

montane forest. Elevation: 820-1030 m.

Material: SUMMIT TRAIL: Elevation ca. 820 m, Zulkifli & R. Jaman,RJ 2867 (UKMB); Elevation ca. 1000

m, Zulkifli & R. Jaman, RJ 2896 (UKMB); SUMMIT: Elevation ca. 1030 m, Zulkifli & R. Jaman, RJ 2930

(UKMB); BUKIT PERMATANG: Elevation ca. 860 m.(Trail to Gunung Kajang), Zulkifli & R. Jaman, RJ

2953 (UKMB).

13.2 Grammitis Sw.

A single species of Grammitis was found in Gunung Kajang area.

13.2.1 Grammitis adspersa Bl.

Epiphyte and lithophyte in shade. Hill dipterocarp forest to lower montane forest.

Elevation: 580-1030 m.

Material: SUMMIT: Elevation ca. 1030 m, Zulkifli & R. Jaman, RJ 2893 (UKMB); AYER ORANG PUTEH:

Elevation ca. 580 m, Zulkifli & R. Jaman,RJ 2943 (UKMB).

13.3. Prosaptia Pres|

Two species are found in Gunung Kajang and Prosaptia leysii is a new record for Pulau

Tioman.

13.3.1 Prosaptia celebica (B\.) Tagawa & K. Iwatsuki

Ctenopteris celebica (Bl1.) Copel.

Epiphyte in semi-shade. Lower montane forest.

Material: SUMMIT: Elevation ca. 1030 m, Zulkifli & R. Jaman, RJ 2892 (UKMB).

Note: Holttum (1968), stated that this species has been found twice in mainland

Peninsular Malaysia (once on Gunung Tahan (1830 m) and the other at an unrecorded

place in Perak.)

13.3.2 Prosaptia leysii (Baker) Ching

Ctenopteris leysii (Bak.) Holtt.

Epiphyte in semi-shade. Upper hill dipterocarp forest to lower montane forest.

Elevation: 860-1000 m.

179

Material: SUMMIT TRAIL: Elevation ca. 1000 m, Zulkifli & R. Jaman, RJ 2901 (UKMB); BUKIT

PERMATANG: Elevation ca. 860 m, Zu/kifli & R. Jaman, RJ 2952 (UKMB).

Note: Holttum (1968), noted that the southern limit of distribution of this species

in Peninsular Malaysia is Gunung Ledang (Mount Ophir) at elevations varying

from 150 m to 760 m.

13.4 Scleroglossum v.A.v.R.

Only one species occurs in Gunung Kajang area.

13.4.1 Scleroglossum debile (Kuhn) Alderw.

Epiphyte in shade. Upper hill dipterocarp forest.

Material: SUMMIT TRAIL: Elevation ca. 900 m, Zulkifli & R. Jaman, RJ 2894 (UKMB).

14. HYMENOPHYLLACEAE

14.1 Cephalomanes C. Pres

14.1.1 Cephalomanes javanicum (B1.) Bosch

Trichomanes jJavanicum Blume

On semi-exposed rock. Lowland dipterocarp forest to lower montane forest.

Elevation: 200-1030 m.

Material: SUNGAI AYER PERTAMA TRAIL: Elevation ca. 200 m, Zulkifli & R. Jaman, RJ 2825 (UKMB);

AYER SURING: Elevation ca. 480 m, Zulkifli & R. Jaman, RJ 2840 (UKMB); AYER ORANG PUTEH:

Elevation ca. 600 m, Zulkifli & R. Jaman, RJ 2858 (UKMB); SUMMIT: Elevation ca. 1030 m, Zulkifli & R.

Jaman, RJ 2937 (UKMB).

14.2 Meringium C. Presl

14.2.1 Meringium acanthoides ( Bosch ) Copel.

Hymenophyllum acanthoides Bosch

On shaded rock. Hill dipterocarp forest.

Material: AYER ORANG PUTEH: Elevation ca. 600 m, Zulkifli & R. Jaman, RJ 2970 (UKMB).

14.3. Microgonium C. Presl

14.3.1 Microgonium bimarginatum Bosch

Trichomanes bimarginatum Bosch

On shaded rock. Hill dipterocarp forest. Elevation: 650 m.

Material: AYER ORANG PUTEH: Elevation ca. 650 m, Zulkifli & R. Jaman, RJ 2970 (UKMB).

180

14.4 Selenodesmium (Prantl) Copel.

14.4.1 Selenodesmium obscurum (B1.) Copel.

Trichomanes obscurum Bl.

Terrestrial and lithophytic in shade. Hill dipterocarp forest to upper hill

dipterocarp forest. Elevation: 600 - 780 m.

Material: GUA TAHI ANGIN TRAIL: Elevation ca. 780 m, Zulkifli & R. Jaman, RJ 2900 (UKMB); AYER

ORANG PUTEH: Elevation ca. 600 m, Zulkifli & R. Jaman, RJ2900A (UKMB).

14.5 Vandenboschia Copel.

14.5.1 Vandenboschia birmanica (Bedd.) Ching

Trichomanes radicans Sw. auct.

On shaded rock. Hill dipterocarp forest.

Material: AYER SURING: Elevation ca. 489 m, Zulkifli & R. Jaman, RJ 2836 (UKMB).

_ Note: Holttum (1968), recorded that only one small specimen has been collected at

Cameron Highland and postulated that this highland is the southern limit of this

species in South-east Asia.

15. LOMARIOPSIDACEAE

15.1 Bolbitis Schott

Three species are found in Gunung Kajang and B. appendiculata ssp. appendiculata is

a new record for Pulau Tioman.

15.1.1 Bolbitis appendiculata (Willd.) K. Iwatsuki ssp. appendiculata

Egenolfia appendiculata (Willd.) J. Sm. ssp. appendiculata

On shaded rock. Lowland dipterocarp forest.

Material: SUNGAI AYER PERTAMA TRAIL: Elevation ca. 200 m, Zulkifli & R. Jaman, RJ 2835 (UKMB).

15.1.2 Bolbitis heteroclita (C. Presl) Ching

Terrestrial in shade. Hill dipterocarp forest.

Material: AYER SURING: Elevation ca. 480 m, Zulkifli & R. Jaman, RJ 2837 (UKMB).

15.1.3 Bolbitis sinuata (C. Presl) Hennipman

Bolbitis malaccensis (C. Chr.) Ching

On shaded rock. Lowland dipterocarp forest.

Material: KAMPONG PAYA: Elevation ca. 50 m, Zulkifli & R. Jaman, RJ 2910 (UKMB).

181

15.2 Elaphoglossum Schott

A genus of six species recorded from Peninsular Malaysia, but only one species occurs

in Gunung Kajang area.

15.2.1 Elaphoglossum melanostictum (B1.) T. Moore

On shaded rock. Hill dipterocarp forest to upper hill dipterocarp forest. Elevation:

580-880 m.

Material: BUKIT PERMATANG: Elevation ca. 880 m, Zulkifli & R. Jaman, RJ 2891 (UKMB); AYER

ORANG PUTEH TRAIL: Elevation ca. 580 m, Zulkifli & R. Jaman, RJ 2904 (UKMB); SUMMIT TRAIL:

Elevation ca. 840 m, Zulkifli & R. Jaman, RJ 2947 (UKMB).

Note: Holttum (1968) noted that this species was found in Penang, but nowhere

else in Peninsular Malaysia.

15.3. Teratophyllum Mett. ex Kuhn

One species found in Gunung Kajang area.

15.3.1 Teratophyllum aculeatum (B1.) Mett. var. montanum Holttum

Climber in light shade. Hill dipterocarp forest to upper hill dipterocarp forest.

Elevation: 550-920 m.

Material: SUMMIT-TELUK JUARA TRAIL: Elevation ca. 920 m, Zulkifli & R. Jaman, RJ 2965 (UKMB);

AYER ORANG PUTEH TRAIL: Elevation ca. 550 m, Zulkifli & R. Jaman, RJ 2851 (UKMB).

16. MARATTIACEAE

16.1 Angiopteris Hoffm.

Only one species occurs in Gunung Kajang.

16.1.1 Angiopteris evecta ( G. Forst. ) Hoffm.

Terrestrial. Hill dipterocarp forest.

Material: AYER ORANG PUTEH: Elevation ca. 610 m, Zulkifli & R. Jaman, RJ 2917 (UKMB).

17. NEPHROLEPIDACEAE

17.1 Nephrolepis Schott

Three species were identified from Gunung Kajang area and WN. hirsutula is a new

record for Pulau Tioman.

182

17.1.1 Nephrolepis auriculata (L.) Trimen

Nephrolepis biserrata (Sw.) Schott

Terrestrial and climbing, and lithophyte in shade and expose. Coastal vegetation

to upper hill dipterocarp forest. Elevation: 1-950 m.

Material: KAMPONG PAYA: Elevation ca. | m, Zulkifli & R. Jaman, RJ 2816 (UKMB); AYER SURING:

Elevation ca. 210 m, Zulkifli & R. Jaman, RJ 2843 (UKMB); BUKIT PERMATANG: Elevation ca. 820 m,

Zulkifli & R. Jaman, RJ 2922 (UKMB); SUMMIT-TELUK JUARA TRAIL: Elevation ca. 950 m, Zulkifli &

R. Jaman, RJ 2968 (UKMB).

17.1.2 Nephrolepis davallioides (Sw.) Kunze

Nephrolepis acuminata (Houtt.) Kuhn

Terrestrial and epiphyte in light shade. Lower montane forest.

Material: SUMMIT: Elevation ca. 1030 m, Zulkifli & R. Jaman, RJ 2884 (UKMB).

17.1.3 Nephrolepis hirsutula (G. Forst.) C. Presl

Terrestrial in expose. Lowland dipterocarp forest. Elevation: 70 m.

Material: AYER SURING TRAIL: Elevation ca. 70 m, Zulkifli & R. Jaman, RJ 2911 (UKMB).

18. OLEANDRACEAE

18.1 Oleandra Cay.

Only one species found from two species recorded from Peninsular Malaysia.

18.1.1 Oleandra pistillaris (Sw.) Cav.

Terrestrial in light shade. Lower montane forest.

Material: Gunung Kajang, near summit. Observed.

19. OPHIOGLOSSACEAE

19.1 Helminthostachys Kaulf.

Only one individual was observed during the survey.

19.1.1 Helminthostachys zeylanica (L. ) Hook.

Terrestrial. Alluvial vegetation.

Material: KAMPONG PAYA: Elevation ca. 2 m. Observed.

20. POLYPODIACEAE

20.1 Colysis C. Pres]

Two species are found from the Gunung Kajang area.

20.1.1 Colysis macrophylla (B1.) C. Presl

On rock in semi-shade. Hill dipterocarp forest.

Material: AYER ORANG PUTEH: Elevation ca. 600 m, Zulkifli & R. Jaman, RJ 2859 (UKMB).

20.1.2 Colysis pedunculata (Hook. & Grey.) Ching

On rock in shade. Lowland dipterocarp forest to upper hill dipterocarp forest.

Elevation: 180-850 m.

Material: SUNGAI AYER PERTAMA TRAIL: Elevation ca. 180 m, Zulkifli & R. Jaman, RJ 2824 (UKMB);

AYER SURING: Elevation ca. 480 m, Zulkifli & R. Jaman, RJ 2841 (UKMB); Elevation ca. 480 m, Zulkifli

& R. Jaman, RJ 2853 (UKMB); BUKIT PERMATANG: Elevation ca. 850 m, Zulkifli & R. Jaman, RJ 2895

(UKMB).

20.2 Drynaria J.Sm.

This genus was never recorded from Pulau Tioman before this survey. One species

occurs in Gunung Kajang area.

20.2.1 Drynaria sparsisora (Desv.) T. Moore

Epiphyte in semi-shade. Hill dipterocarp forest.

Material: SUNGAI AYER PERTAMA: Elevation ca. 460 m, Zulkifli & R. Jaman, RJ 2908 (UKMB).

20.3 Goniophlebium (B1.) C. Pres

Two specimens were collected from Gunung Kajang area. One sterile specimen needs

further determination. Four other species occur in Peninsular Malaysia, viz. G.

percussum, G. persicifolium, G. prainii and G. subauriculatum.

20.3.1 Goniophlebium korthalsii (Mett.) Bedd.

Epiphyte in light shade. Upper hill dipterocarp forest.

Material: SUMMIT TRAIL FROM TELUK JUARA: Elevation ca. 950 m, Zulkifli & R. Jaman, RJ 2967

(UKMB).

Note: Holttum (1968), noted that this species was only once collected in Peninsular

Malaysia by H. Kunstler (King’s Collector) in Larut, Perak.

20.3.2 Goniophlebium sp.

Terrestrial in semi-shade. Lowland dipterocarp forest. Sterile.

Material: AYER SURING TRAIL: Elevation ca. 210 m, Zulkifli & R. Jaman, RJ 2843. (UKMB).

184

20.4 Loxogramme C. Presl

Only one species found in Gunung Kajang area from two species recorded for Pulau

Tioman.

20.4.1 Loxogramme avenia (B1.) C. Presl

On exposed rock. Hill dipterocarp forest.

Material: AYER ORANG PUTEH: Elevation ca. 600 m, Zulkifli & R. Jaman, RJ 2860 (UKMB).

20.5 Microsorum Link.

One species was identified from Gunung Kajang area and is a new record for Pulau

Tioman.

20.5.1 Microsorum heterocarpum (B1.) Ching

On shaded rock. Hill dipterocarp forest to upper hill dipterocarp forest. Elevation:

480-800 m.

Material: AYER SURING: Elevation ca. 480 m, Zulkifli & R. Jaman, RJ 2838 (UKMB); SUMMIT TRAIL:

Elevation ca. 800 m, Zulkifli & R. Jaman, RJ 2926 (UKMB).

20.6 Phymatosorus Pic. Serm.

20.6.1 Phymatosorus nigrescens (Blume) Pic. Serm.

Phymatodes nigrescens (B1.) J. Sm.

On shaded rock. Hill dipterocarp forest.

Material: AYER ORANG PUTEH: Elevation ca. 580 m, Zulkifli & R. Jaman, RJ 2950 (UKMB).

20.7 Pyrrosia Mirbel

Only one species found during the survey.

20.7.1 Pyrrosia lanceolata (Linn.) Farwell

Pyrrosia adnascens (Sw.) Ching

On exposed rock. Coastal vegetation.

Material: KAMPUNG PAYA: Elevation ca. | m, Zulkifli & R. Jaman, RJ 2819 (UKMB).

20.8 Selliguea Bory

Four species found in Gunung Kajang area.

20.8.1 Selliguea enervis (Cav.) Ching

Cypsinus enervis (Cav.) Copel.

185

Epiphyte in light shade. Upper hill dipterocarp forest.

Material: SUMMIT TRAIL: Elevation ca. 950 m, Zulkifli & R. Jaman, RJ 2887 (UKMB).

20.8.2 Selligua heterocarpa (B1.) Bl.

On rock in light shade. Upper hill dipterocarp forest.

Material: SUMMIT TRAIL: Elevation ca. 750 m, Zulkifli & R. Jaman, RJ 2925 (UKMB).

20.8.3 Selliguea taeniata (Sw.) Parris

Crypsinus taeniatus (Sw.) Copel..

On exposed rock. Upper hill dipterocarp forest.

Material: BUKIT PERMATANG: Elevation ca. 820 m, Zulkifli & R. Jaman, RJ 2866 (UKMB).

20.7.4 Selliguea triloba (Houtt.) M.G. Price

Crypsinus trilobus (Houtt.) Copel.

Epiphyte in light-shade. Lower montane forest.

Material: SUMMIT: Elevation ca. 1030 m, Zulkifli & R. Jaman, RJ 2903, RJ 2905 (UKMB).

21. PTERIDACEAE

21.1 Pteris L.

Six species are recorded from Pulau Tioman. Two species are found in Gunung Kajang

area, including P. oppositipinna which is a new record for Pulau Tioman.

21.1.1 Pteris ensiformis Burm. f.

Terrestrial in open situations. Coastal vegetation.

Material: KAMPONG PAYA: Elevation ca. 3 m, Zu/kifli & R. Jaman, RJ 2822 (UKMB).

21.1.2 Pteris oppositipinna Fée

Pteris asperula J. Sm. ex Hieron.

Terrestrial in light-shade. Upper hill dipterocarp forest.

Material: BUKIT PERMATANG: Elevation ca. 840 m, Zulkifli & R. Jaman, RJ 2928 (UKMB).

Note: Holttum (1968), noted that this species was only recorded in Selangor, Pulau

Pinang and Negeri Sembilan.

22. SCHIZAEACEAE

22.1 Lygodium Sw.

Three species are recorded from Pulau Tioman but only one species found during the

survey in Gunung Kajang and surrounding area.

186

22.1.1 Lygodium flexuosum (L.) Sw.

Terrestrial and scrambling over bushes. Coastal vegetation.

Material: KAMPONG PAYA: Elevation ca. 1 m, Zulkifli & R. Jaman, RJ 2820 (UKMB).

22.2 Schizaea Smith

Only one species found, mostly near summit of Gunung Kajang.

22.2.1 Schizaea digitata ( Linn. ) Sw.

Terrestrial. Upper hill dipterocarp to lower montane forest. Elevation 950-

1020 m.

Material: SUMMIT TRAIL: Elevation ca. 1000 m, Zulkifli & R. Jaman, RJ 2897 (UKMB); Elevation ca.

950 m, Zulkifli & R. Jaman, RJ 2941 (UKMB); Elevation ca. 1020 m, Zulkifli & R. Jaman, RJ 2962

(UKMB).

23. THELYPTERIDACEAE

23.1 Pronephrium C. Presl

Two species are found in Gunung Kajang area and P. repandum is a new record for

Pulau Tioman.

23.1.1 Pronephrium repandum (Fée) Holttum

Terrestrial in light shade. Hill dipterocarp forest. Elevation: 380-580 m.

Material: SUNGAI AYER PERTAMA TRAIL: Elevation ca. 380 m, Zulkifli & R. Jaman, RJ 2828 (UKMB);

AYER ORANG PUTEH: Elevation ca. 580 m, Zulkifli & R. Jaman, RJ 2850 (UKMB).

23.1.2 Pronephrium rubicundum (Alderw.) Holttum

Terrestrial in shade. Hill dipterocarp forest. Elevation: 510-750 m.

Material: AYER ORANG PUTEH TRAIL: Elevation ca. 510 m, Zulkifli & R. Jaman, RJ 2844 (UKMB);

Elevation ca. 510 m, Zulkifli & R. Jaman, RJ 2849 (UKMB); BUKIT PERMATANG TRAIL: Elevation 750

m, Zulkifli & R. Jaman, RJ 2870 (UKMB).

23.2 Sphaerostephanos J. Smith

One species found and is a new record for Pulau Tioman.

23.2.1 Sphaerostephanos heterocarpus (B1.) Holttum

Cyclosorus heterocarpus (Bl1.) Ching

Terrestrial in light shade. Hill dipterocarp forest. Elevation: 360-550 m.

Material: AYER SURING TRAIL: Elevation ca. 360 m, Zulkifli & R. Jaman, RJ 2831 (UKMB); Elevation

ca.550 m, Zulkifli & R. Jaman, RJ 2907 (UKMB).

187

24. VITTARIACEAE

24.1 Antrophyum Kaulf.

Only one species found in Gunung Kajang area.

24.1.1 Antrophyum callifolium B.

On shaded rock. Hill dipterocarp forest.

Material: AYER ORANG PUTEH: Elevation ca. 600 m, Zulkifli & R. Jaman, RJ 2855 (UKMB).

24.2 Vaginularia Fée

24.2.1 Vaginularia trichoidea Fée

Monogramma trichoidea J. Sm.

Epiphyte in shade. Lower montane forest.

Material: SUMMIT: Elevation ca. 1030 m, Zulkifli & R. Jaman, RJ 2948 (UKMB).

24.3 Vittaria Sm.

Two species are found in the Gunung Kajang area.

24.2.1 Vittaria angustifolia BI.

On shaded rock. Upper hill dipterocarp forest. Elevation: 840-850 m.

Material: BUKIT PERMATANG: Elevation ca. 850 m, Zulkifli & R. Jaman, RJ 2902 (UKMB); Elevation

ca. 840 m, Zulkifli & R. Jaman, RJ 2951 (UKMB).

24.3.2 Vittaria scolopendrina (Bory) Thw.

On shaded rock. Upper hill dipterocarp forest.

Material: BUKIT PERMATANG: Elevation ca. 900 m, Zulkifli & R. Jaman, RJ 2902 (UKMB).

25. WOODSIACEAE

25.1 Diplazium Sw.

This genus was never recorded previously from Pulau Tioman. Two species are found

in Gunung Kajang area.

25.1.1 Diplazium sorzogonense (C. Presl) C. Presl

Athyrium sorzogonense (C. Presl) Milde

Terrestrial in open situations. Upper hill dipterocarp forest.

Material: BUKIT PERMATANG TRAIL: Elevation ca. 780 m, Zulkifli & R. Jaman, RJ 2873 (UKMB).

188

25.1.2 Diplazium tomentosum Bl.

On shaded rock. Upper hill dipterocarp forest.

Material: GUA TAHI ANGIN: Elevation ca. 950 m, Zulkifli & R. Jaman, RJ 2923 (UKMB).

Acknowledgements

We would like to thank the Curators of the Herbarium, Singapore Botanical Garden

(SING), especially Mr. Mohd Shah Mohd Noor and Mr. Ali, the Herbarium, Forest

Research Institute of Malaysia (KEP) for their cooperation during our visits. We

also thank Mr. Sani Miran and Mr. Idris Hj. Sharif for their company during the

fieldworks. This work was financially supported by the Department of Botany,

Universiti Kebangsaan Malaysia, IRPA 04-07-03-007 and IRPA 08-02-0009.

References

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Ferns and fern-allies. Royal Botanic Gardens, Kew. 165 pp.

Parris, B.S. and A. Latiff. 1997. Towards a pteridophyte flora of Malaysia: A

provisional checklist of taxa. Mal. Nat. J. 50: 235-280.

Piggot, A.G. 1988. Ferns of Malaysia in colour. Tropical Press Sdn. Bhd., Kuala

Lumpur. 458 pp.

Morphological Variation of Pangium edule Reinw. Fruits in

Malaysia

I. Faridah-Hanum

Faculty of Forestry,

Universiti Pertanian Malaysia

43400 Serdang, Malaysia

Abstract

Morphological variation of fruits and seeds occurring in Pangium edule is described. The fruits of typical

form, “kepayang lenga” are oblong with blunt apices , those of “kepayang papan” are subglobular and the apices

are sharp while those of “kepayang bubur” are ellipsoidal and the apices are blunt. Seeds differ in their sizes too.

Introduction

Pangium edule or commonly known as “kepayang”, is monotypic and belongs

to the family Flacourtiaceae. The trees are either found wild or more commonly,

cultivated on a small scale throughout Southeast Asia, Melanesia and Micronesia.

They reach the height of 25 m and frequently as tall as c. 40 m. As the boles are

straight and often reach the extreme diameter of 1 m, they offer good timber source

to the locals. The seeds contain high quality oil which has been used for cooking

for hundreds of years, moreover the roasted seeds have long been used for the

preservation of meat and fishes. Many more uses are documented by Sleumer (1954)

and Burkill (1966). In Indonesia and elsewhere the use of the fruits as food is very

widespread. In Sarawak the oil is used for cooking and for preserving fish and

meat.

Prior to this, the intraspecific variation especially that of the fruits and seeds has

not been described. During the trips made by the author to Sarawak to collect

samples of the species, observations were made on the occurrence of the variations

in this species and herein described.

Materials and Methods

The morphological observations were made on the materials either collected

personally by the author in the field or those deposited in various herbaria in the

country. Those of the former were deposited in the Herbarium, Forestry faculty,

Universiti Pertanian Malaysia (tentatively designated UPMF).

The fruits and seeds collected were observed on the following characters;

shape, size, shape and length of the apex. They were then opened to observe the

seeds; observation on the seeds were made for the characters of overall shape

and size.

190

Observations

General morphology

Tree branching at about 5 m, buttress small to spreading, up to 1.5 m high. Inner

bark cream-yellowish with orange tints. Sapwood yellowish to orangish, slash giving

a colourless exudate with a distinct smell of malt. Leaves are ovate and hetero-

phyllous, often on the same branch or tree of any age, up to 75 cm long. Flowers

rather fragrant, flowering starts immediately after every new flush of leaves. Fruits

1-(-4) per infructescence of varying sizes (16-24 x 10-16 cm), oblong to ellipsoidal

in shape with bluntly to distinctly sharp tips, rather heavy, each weighing up to 2.5

kg, fruiting 1-4 times per year, fruits smaller with massive fruiting, no indication of

maturity of fruits from the trees until they fall and rot, fruit stalk up to 60 cm long.

Seeds interlocked, embedded in a pungent mesocarp when ripe, almost triangular

in shape, 13-40 in number.

Fruit and seeds morphology

Ridley (1922) described the fruits of Pangium edule as 17-30 cm long and 9-

10. cm across and seeds were recorded as about 5 cm across and triangular in outline.

Sleumer (1954) gave a more extensive description of the fruits as being oblong-

ovoid, variable in size with blunt tips and containing about 20 irregularly shaped

seeds (3-)4-6 x 2-3(-4) cm in size. In this context he had mentioned that the fruits

are variable in size, and shape and the seeds are also variable in size. All these

descriptions seem to refer to the typical form or “kepayang lenga” (as used in this

paper).

Whitmore (1973) on the other hand, described the fruits of Pangium edule as

pear-shaped with blunt tips and 15-25 by 7-15 cm in size with seeds about 4 cm

long. This description seems to refer to “kepayang lenga” too.

While collecting specimens in Sarawak, the author recognised two distint variants

of the Pangium edule with respect to fruits and seeds morphology and they are

described below:

Typical form of Pangium edule

3. 16¢

Vernacular names: “kepayang”, “payang”, “buah keluak”, “kepayang lenga”,

“kepayang keluwak” or “pangi”.

Description: Fruits, oblong to oblong-oval, 18-23 x 10-14 cm, tips almost

rounded, blunt, ca. 0.5 cm long; seeds 3.5-4.0(-6.0) cm, seeds per fruit 25-30 (Fig.

| a-b).

Specimens studied: SARAWAK, Serian, Ilias Paie $.28040 (SAR); Miri, Jack

Liam s.n. (SAR); Miri, Anderson S.16428 (SAR); Niah, Anon. KFN 98660 (SAR);

Kuching, Faridah-Hanum FHI 398 (UPMF); Samarahan, Faridah-Hanum FHI 401,

19]

FHI 403, FHI 404, FHI 405, FHI 406, FHI 407, FHI 408, FHI 411 (UPMF); 7th

Division, Ulu Sg. Melinau, Hose Mt., P. Chai $.37224 (SAR); Kuching, Pedawan

Rd., P. Chai & Erwin S. 27525 (SAR); Niah, G. Subis, Anderson S. 16428 (SAR):

Belaga, B. Lee S. 39813 (SAR): SABAH, Inanam, Burgess SAN 28421 (SAN):

Beluran, Amin G. et al. SAN 93886 (SAN); Beaufort, Mikil SAN 34593 (SAN):

Keningau, Amin G. SAN 95499 (SAN): Tongod, Aban & Clement SAN 69926

(SAN). PAHANG, Ulu Dong, Faridah-Hanum FHI 360, FHI 361 (UPMEF):

Temerloh, Faridah-Hanum FHI 359 (UPMF): Sg. Triang. Kassim 40368 (KEP):

PERAK, Temenggor; King’s Coll. 10065 (K); Kg. Temenggor, Chelliah FRI 98660

(KEP); SELANGOR; s.]., Tachun bin Baba 57086 (KEP); Ulu Langat, Gadoh anak

Umbai KL 1871 (KEP): Kepong, Kochummen FRI 19098 (KEP); KELANTAN,

Gua Musang, Husmady et al. FRI 39709 (KEP).

The two variants observed are as follows:

_ Pangium edule “kepayang papan”.

Description: Fruits subglobular to ellipsoidal, 16-24 x 11-16 cm, tips oblong-

triangular, sharp, 1.5-2.0 cm long: seeds 3.5-5.5. cm, seeds per fruit (13-) 20-24 (-

29). (Fig. 1 c-d)

Specimens studied: SARAWAK, Seri Aman, Faridah-Hanum FHI 400 (UPMF):

Samarahan, Tarat, Faridah-Hanum FHI 410, FHI 412, FHI 413: Seri Aman Division,

Faridah-Hanum FHI 399 (UPMF);: Serian, Ilias Paie S. 28040 (SAR); Ulu Kapit, P.

Chai et al. S. 33309 (SAR).

Pangium edule “bubur’”.

Description: Fruits ellipsoid to oval, 18-20 x 12-14 cm, tips almost rounded,

blunt, ca. 0.5 cm long; seeds 2.5-3.0(-4.0) cm, seeds per fruit 25-30(-40).

(Fig. 1 e-f).

Specimen studied: SARAWAK, Seri Aman, Faridah-Hanum FHI 402 (UPMF):

Samarahan, Tarat, Faridah-Hanum FHI 409, FHI 414, FHI 415 (UPMF).

Distribution: Throughout Malesia, extending to Melanesia and Micronesia,

apparently there is some variation in Borneo too; in Malaysia throughout (Fig. 2).

Burkill (1966) cited the following vernacular names for Peninsular Malaysia,

viz. kepayang, payang and buah keluak. The typical Pangium edule or kepayang

lenga, kepayang, payang is distributed widely in Sabah, Sarawak and Peninsular

Malaysia but its variants, Kepayang papan and kepayang bubur so far have only

been in found Sarawak, Borneo.

Discussion

Except for the differences in the fruit and seed characters, other morphological

characters remain very similar. Recent observations and collections made by the

192

author showed that description by Corner (1940), Sleumer (1954) and Whitmore

(1973) all could be referred to “kepayang lenga’’, the typical P. edule. The other

two types observed are atypical of the “kepayang”.

While the fruits of kepayang papan are almost globular the seeds are similar to

those of kepayang lenga in both their shape and size. However, the fruits of kepayang

bubur are smaller and ellipsoidal and the seeds are relatively smaller too.

Fig. 1. Morphology of fruits and seeds of Pangium edule. (a-b, kepayang lenga; c-d, kepayang papan; e-f,

kepayang bubur)

Distribution of Pangium edule in Malaysia. ( @ kepayang lenga, kepayang papan, and O kepayang bubur).

Fig, 2.

194

Acknowledgements

The author would like to thank the International Foundation of Science (IFS),

Sweden for the financial support given through Grant No. D/2255-1. I am grateful

to the Director of the Agriculture Department, Sarawak and his staff for support

while I was in Sarawak. Thanks are also due to Mr. Chai Chen Chong of the

Agriculture Research Centre, Semonggok, Sarawak for assisting me in the field.

References

Burkill, I. H. (1966). A dictionary of the economic products of the Malay Peninsula

2 : 1680-1682. Ministry of Agriculture and Cooperatives, Kuala Lumpur,

Malaysia.

Corner, E.J.H. (1940). Wayside Trees of Malaya. 3rd Ed. 1 : 348 Malayan Nature

Society.

Ridley, H. N. 1922. Flacourtiaceae. Flora of the Malay Peninsula | : 163, Reeves

& Co., London

Sleumer, H. (1954). Flacourtiaceae. In : Steenis, van C.G.G.J. (Ed.). Flora Malesiana

Ser. 1, Vol.5 > 35-39

Whitmore, T. C. 1973. Flacourtiaceae. In : T. C. Whitmore (Ed.). Tree Flora of

Malaya 2 : 155, Longmans Malaysia

Ternstroemia magnifica Stapf ex Ridley (Theaceae) and

Kibatalia macrophylla (Pierre) Woodson (Apocynaceae),

Two Species New to Peninsular Malaysia.

A. Latiff, Natrah Mohamad & A. Zainudin Ibrahim

Department of Botany, Universiti Kebangsaan Malaysia,

43600 Bangi, Malaysia.

Abstract

Recent collections of Ternstroemia magnifica Stapf ex Ridley (Theaceae) and Kibatalia macrophylla (Pierre)

Woodson (Apocynaceae) were made in Bangi Permanent Forest Reserve, Selangor, and Langkawi Islands, Kedah,

respectively, representing new records for the flora of Peninsular Malaysia. Diagnostic descriptions and keys to

species are presented with some morphological notes.

Introduction

The pantropical genus Jernstroemia Mutis ex L. f. is represented by shrubs and

trees in Peninsular Malaysia. Keng (1978) listed a total of seven species as occurring

in the peninsula, viz. 7: bancana Miq., T: corneri H. Keng, T: evenia (King) A.C.

Smith, 7: maclellandiana Ridl., T. montana Ridl., T: penangiana Choisy and T.-

wallichiana (Griff.) Engler. In the course of our preparation of a checklist of the

flowering plants of Bangi Permanent Forest Reserve, Selangor, one theaceous

specimen belonging to Ternstroemia magnifica was collected. The discovery of

this species in Peninsular Malaysia is both suprising and phytogeographically

noteworthy. In a similar exercise of collecting plants in the Langkawi Archipelago,

Kedah, for the preparation of the Checklist of Langkawi flowering plants, two

specimens of an interesting apocynaceous tree species were collected. It was

subsequently identified as Kibatalia macrophylla which proved to be a new record

for both Peninsular Malaysia and the Malaysian flora. A diagnostic description and

a key to the species is herein given.

Ternstroemia magnifica Stapf ex Ridl., Kew Bull. (1938) 175

Tree c. 20 m tall, no buttress. Bole straight, bark profusely lenticellate becoming

large cracks; inner bark reddish, sapwood cream; branches terete, greyish, the

ultimate ones c. 5 mm in diameter; branching terminalian-type. Leaves spirally

arranged, crowded at the shoot, very coriaceous, 16.4-18.7 « 7.0-9.2 cm, oblanceolate

to obovate, apex abruptly acuminate, base cuneate, margin entire, lateral nerves c.

12 on each side, very inconspicuous; petioles c. 2 cm long. Flowers not observed.

Fruits ellipsoid, brown when dry, smooth, 7.6 = 6.4 cm, calyx-lobes thick, persistent,

verrucose, strongly attached to the base, the stalk c. 2.4 cm long. Seeds oblong,

rounded at both ends, c. 6 x 2.4 cm attached to the central axis by funiculus, the

funiculus flat, filiform.

196

Specimen examined: Peninsular Malaysia, Selangor, Bangi Permanent Forest

Reserve; 22.1.1994, A. Zainudin et al. AZ 4751 (UKMB).

Representative Specimens from Borneo examined: Sarawak, P. Chai S. 19717

(K, SAR), Ilias Paie S. 16973 (K, SAR); Sabah, Aban & Saikeh SAN 71858 (SAN,

K), Chew & Corner RSNB 4038 (SAN, K), Jacobs 5702 (L, K, SAR), Mikil 33943

(SAN), Mujin SAN 33772 (K, SAN).

Distribution: Borneo and Malay Peninsula.

Ecology: In Borneo the species is found in the lowland dipterocarp forest of

Sabah, Sarawak and Kalimatan. The new record in the Malay Peninsula was

collected from a twice logged-over lowland dipterocarp forest of the Bangi

Permanent Forest Reserve, Selangor. The occurrence of the species in Peninsular

Malaysia is therefore very interesting as it extends its present geographical range

of distribution westwards from Borneo island. The close affinity of the Bornean

flora to that of the southeast coast of Peninsular Malaysia is quite well accepted

and known. However, the new locality for the species at the Bangi Permanent

Forest Reserve, Selangor is about 300 km inland from the east coast of Peninsular

- Malaysia. By extrapolation, it is quite probable that the species is also found in the

south of the peninsula, notably in the Endau-Rompin area.

T. magnifica is related to T. evenia because of the character of inconspicuous

leaf venation on the lower leaf surfaces but differs in fruit morphology and also to

T. philipinensis Merr. but differs from it in the character of the fruits and leaves. In

Peninsular Malaysia it is perhaps more closely related to 7? corneri, from which it

can be distinguished as follows:-

Leaves 20-28 cm long; petioles c. | cm long; fruits 4.5-5 x 2-2.5 cm, apex

briefly forked; calyx smooth, margin free, deflexed; swamp forest........ T. corneri

Leaves 16-19 cm long; petioles c. 2 cm long; fruits 7.6 x 6.4 cm, apex not

forked; calyx verrucose, margin firmly attached; dry lowland

FOTOS... S6s.ssad spake be cal cus Texaee cule cae ooo ee NUE aa te Re cee a aera T. magnifica

Morphological notes. The type specimen (Haviland 1984, K!) was collected

in the vicinity of Kuching, Sarawak, and the species is now known to be widely

distributed in Borneo. Generally the fruits of the type specimen are not as large as

that of the Bangi specimen and the persistent calyx are always more or less deflexed

or at least the margin is free from adnation to the mature fruit. Dr. S. C. Chin

(SING) confirmed the occurrence of leaf sclereids, a characteristic feature of the

Theaceae. However, the character of the seeds bothered Dr. Hsuan Keng (SING) a

little. The ovules (and later seeds) are generally attached at the central axis of the

ovary in almost all the theaceous plants known to him but in the Bangi specimen

the seeds are attached to the funiculus which arise from the top of the central column.

This appears to be a discrepancy, axile versus basal placentation in the Theaceae.

According to Dr. M. van Balgooy (L. pers. comm.) this is also true for the Kalimantan

specimens which he observed. It appears that this character is common for the

species in its range of distribution. When describing the species, Ridley (1938)

stated that it is unlike any other species in the genus in its very large staminate

flowers and glaucous leaves. The above variations prompted us to suggest that the

position of this species within the genus should be assessed. After all the position

of 7ernstroemia itself within the Theaceae has been assessed many times in its

taxonomic history.

Notes on other Ternstroemia species in Peninsular Malaysia.

T. bancana Miq,., Fl. Ind. Bat. suppl. (1861) 477. Quite widespread, more often

near the sea.

T. corneri H. Keng, Gdns’. Bull. Sing. 29 (1977) 143-144. Endemic to the swamp

forests of South Johor.

T. evenia (King) A.C. Smith, Sargentia 7 (1947) 78. Endemic to Peninsular Malaysia,

recorded from lowland to montane forests in Perak, Kelantan, Pahang and Selangor.

T. maclellandiana Ridl., J. Fed. Mal. Stat. Mus. 6 (1915) 140. Widespread in

montane forests.

T. montana Ridley, J. Str. Br. Roy. As. Soc. 73 (1916) 141. Restricted to the montane

forests of Gunung Jerai (Kedah), Gunung Korbu (Perak) and Cameron Highlands

(Pahang).

T. penangiana Choisy, Mem. Soc. Phys. Hist. Nat. Geneve | (1822) 118. Widespread.

T. wallichiana (Griff.) Engler, Pflanzenfam. Nachtr. 1 (1897) 246. Widespread in

the lowland forests.

Kibatalia macrophylla (Pierre) Woodson, Philipp. Journ. Sci. 60 (1936) 214;

Rudjiman, Agric. Univ. Wagenigen Papers 86(5) (1986) 38 (as K. anceps), fig. 10.

Tree c. 12 m high, probably deciduous. Trunk c. 15 cm in diameter. Leaves 12-

14 by 2.3-4.1 cm, elliptic; apex acuminate; base cuneate to obtuse; margin entire,

glossy, glabrous above, beneath sparsely pubescent, coriaceous, with 18 secondary

veins on each side; tertiary veins inconspicuous; petioles 0.5 cm, glabrous.

Infrutescence lax; mericarps 9-12 = 0.5 cm, narrowly ellipsoid; pedicels 1.5 cm

long; peduncle 0.2 cm long.

Ecology: A component of beach forests.

Specimens studied: Malaysia, Kedah, Pulau Langkawi, Pulau Beras Basah, A.

Latiff, A. Zainudin & Hamid Salleh ALM 3568, 21.11.90 (UKMB); Pulau Singa

Besar, A. Zainudin et al. AZ 4394, 19.11.92 (UKMB).

Other specimens examined: Thailand, Winit 1246 (K); Vietnam, Tonkin, Balansa

2103 (K), Poillane 17/2 (K)

198

Distribution: China (Yunan), Peninsular Burma, Thailand, Indochina and

Peninsular Malaysia (Langkawi island).

Rudjiman (1986) recognised 15 species of Kibatalia in the world and five of

them occur in Malaysia, viz. K. arborea (Bl.) G. Don, K. borneensis (Stapf) Merr.,

K. laurifolia (Ridl.) Woodson, K. maingayi (Hook. f.) Woodson and K. villosa

Rudjiman. K. borneensis is endemic to Sarawak. Earlier, Whitmore (1973) gave an

account for Peninsular Malaysia and listed two species only, namely K. maingayi

and K. arborea. Ridley (1923) considered the taxa under Vallaris. With the discovery

of this new record for Peninsular Malaysia the number of species for the peninsula

is now five and for Malaysia six. The occurrence of the species in the Langkawi

archipelago represents the current southermost limit for the species as before this

the southern limit known is Tennaserim in Burma. Similar pattern of distribution

was shown by K. laurifolia, another Asiatic species.

Key to the Kibatalia species in Malaysia (adapted from Rudjiman, 1986)

L., Stamens exserted: for 05-4 amin 2, 62:5, osaeeee eyed suedeecee eed 2

Stamens included for 3-30 mm .3:...0:.:c.acccscevenesosateeeressaeea cee meen 2)

2. Leaves 12-34 cm long; costa hairy beneath; petioles 6- 15 cm long, sparsely

PUDESCEME 2.60. /ossseva ecnpnteasaanecunoetis eel acceeateaes aera nee ee eee eae K. macrophylla

Leaves 4-19 cm long; costa glabrous; petioles 2-10 cm long, glabrous............ 3

3. Leaves 4-14 x 0.8-5.5 cm; secondary veins 4-8 on each side; corolla tube 5-10

101 008 (0) || eee eae nn ae een Alin ARI Lt es nner Ne ENG BM sion. tho yh K. maingayi

Leaves 8-19 x 2-7 cm; secondary veins 7-14 on each side; corolla tube 11-14

OI LOWMP (i. eseidleceascnecties ace esdeden bebe meee eee ne aaa Un seamen he Rei aan Eee ee 4

4. Corolla mouth glabrous; inflorescence |- to 6-flowered; axis with many bracts;

petioles 2-6 mama Lome. ois. cessed eteceae ee hanec nes eee en ee se K. laurifolia

Corolla mouth hairy; inflorescence 8- to 25-flowered; axis without bracts;

petioles 5-15 ‘mum LOM .5..3.- cteteecace eee seeeee ease sexton ees eee ee K. villosa

5. Leaves 16-26 x 8-13 cm; leaf apex acuminate; veins 11-18 on each side; corolla

tube glabrous; pedicels 4-6 cm long; peduncles 2-6 mm long ........... K. arborea

Leaves 6-20 x 1-6.5 cm; leaf apex caudate; veins 8-11 on each side; corolla tube

hairy; pedicels 7-10 mm long; peduncles 12-23 mm long ............ K. borneensis

Notes on other Kitabalia species in Malaysia.

K. arborea (Bl.) G. Don, Gen. Syst. Bot. 4 (1837) 86; Whitmore, Tree Fl. Mal. 2

(1973) 18. Quite rare. Peninsular Malaysia (Pahang and Selangor), and Sabah.

K. borneensis (Stapf) Merrill, Philip. J. Sc. 17 (1920) 309. A species endemic to

Sarawak, in the swamp or heath forests.

K. laurifolia (Ridl.) Woodson, Philip. J. Sc. 60 (1936) 212. Rudjiman (1986)

199

recorded this species for Peninsular Malaysia. Known from one collection from

Perak.

K. maingayi (Hook. f.) Woodson, Philip. J. Sc. 60 (1936) 213; Whitmore, Tree FI.

Mal. 2 (1973) 1973; Vallaris maingayi Hook. f. in Ridley, Fl. Mal. Penin. 2 (1923)

351.

Common throughout Peninsular Malaysia and in Sarawak known only in

Semenggoh Forest Reserve.

K. villosa Rudjiman, Agric. Univ. Wagenigen Papers 86-5 (1986) 82. Rudjiman

(1986) recorded this species for Malaysia. Apparently very rare, known only from

Rengam FR. (Johor) and a few localities in Sarawak.

Fig. 1. Ternstroemia magnifica. A. fruiting branch; note the verrucose calyx

which are tightly attached to the base. B. seeds with funiculus attached.

200

Acknowledgements

We are grateful to Prof. E. Soepadmo, Mr. K. M. Kochummen and Dr. L. G.

Saw (all from KEP) for their initial assistance in identifying Ternstroemia, Dr.

Hsuan Keng (SING) who confirmed the identity of the species, Dr. S. C. Chin

(SING) who confirmed the presence of leaf sclereids and Dr. M. van Balgooy (L)

for highlighting the character of the ovules, and Dr. A.J.M. Leeuwenberg (WAU)

for alerting the correct name of the species, Kibatalia macrophylla. Lastly to the

Directors or Curators of the following herbaria, K, L, SING, KEP for allowing us

to study their materials. This study and publication is supported by IRPA Grant 04-

07-03-007 for which the authors are grateful.

Reference

Keng, H. 1978. Theaceae. In Ng. F. S. P. (Ed.). Tree Flora of Malaya 3 : 293-295.

Longman Malaysia Sdn. Bhd.

Ridley, H. N. 1922. Theaceae. Flora of the Malay Peninsula I : 192-209. Reeves &

Sons, London.

Ridley, H. N. 1923. Apocynaceae. Flora of the Malay Peninsula 2 : 351-352. Reeve

& Sons, London.

Ridley, H. N. 1938. Additions to the flora of Borneo and other Malay islands: VII.

Kew Bull. No. 5 : 175

Rudjiman. 1986. A revision of Beaumontia Wallich, Kibatalia G. Don and

Vallariopsis Woodson (Apocynaceae). Agricultural University Wagenigen

Papers 86(5) : 36-89

Whitmore, T.C. 1973. Apocynaceae. In T.C. Whitmore (Ed.). Tree Flora of Malaya

3: 16-18. Longman Malaysia Sdn. Bhd.

A New Species of Barringtonia (Lecythidaceae) From

Peninsular Malaysia

P. Chantaranothai

Department of Biology, Faculty of Science, Khon Kaen University, Khon Kaen, Thailand

Abstract

A new species, of Barringtonia, B. terengganuensis P. Chantaranothai, is described and illustrated from

Terengganu, Peninsular Malaysia.

Introduction

The genus Barringtonia occurs from eastern Africa to northern Australia and

comprises some 56 species. In preparation for publication of the Lecythidaceae for

the Flora of Thailand, I also examined material from other regions for comparison.

It has revealed the existence of a previously undescribed species of Barringtonia.

Barringtonia terengganuensis P. Chantaranothai sp. nov.

Differt a B. pendula hypanthio minore, petiolis brevioribus, foliis basi cordatis vel rotundatis. Typus:

Peninsular Malaysia, Terengganu, P F Cockburn FRI 8363 (holotypus K).

Small tree up to 6 m tall. Rachis and hypanthium pubescent. Leaves with petioles

4-5 mm long; lamina 26-28.5 x 7-7.5 cm, chartaceous, elliptic, apex acuminate,

base slightly cordate or rounded, margin slightly serrate-crenulate; secondary veins

22-23 pairs, 1.4-2 cm apart. Inflorescence a terminal spike, c. 25 cm long, densely

flowered, with c. 50 flowers; rachis base 2 mm in diam. tapering to 0.8 mm in

diam. near apex; cataphylls 10-12 x 3-5 mm, triangular; bracteoles 0.4-1 « 0.2 mm,

triangular, caducous. Flowers in bud pink; hypanthium without pedicel, funnel-

shaped, quadrangular, with 4 grooved at the corner, pubescent; sepal 4, orbicular or

suborbicular; petal 4, orbicular; ovary 2-locular, ovules 3-6 per locule. Fruits

unknown. Fig. 1.

Peninsular Malaysia. Terengganu: Ulu Sungai Terengganu near Jaram

Galong, primary forest, hillsides, alt. 600 m, 31 May 1968, RF Cockburn FRI

8363 (Holotype K).

Barringtonia terengganuensis is a distinct species which is closest to B. pendula

(Griff.) Kurz. Both species have grooves at the corners of the hypanthium. It differs

from B. pendula mainly in its smaller hypanthium, shorter petiole and slightly

cordate or rounded leaf base. P F Cockburn FRI 8363 (K) was originally determined

as B. fusiformis King. The leaves of B. terengganuensis and B. fusiformis are

similar but the two species differ in the former having a grooved hypanthium and

no pedicel.

202

ntebis NY

Fig. 1. Barringtonia terengganuensis. A. habit; B. floral bud; C. floral bud in

longitudinal section. All from Cockburn FRI 8363. Drawn by M. Tebbs

Acknowledgements

I wish to thank Khon Kaen University for financing my work at the Royal Botanic

Gardens, Kew (K); Assist. Prof. B.Watanakij, Assoc. Prof. Dr. L. Sanoa-muang

and Dr. S. Santawananpas for encouragement; Keeper of the Kew Herbarium for

the use of specimens; the late Prof. D. A.Webb and Alan Radcliffe-Smith for the

Latin diagnosis; M. Tebbs for the illustration and the Programme for Biodiversity

Research and Training, Thailand (TRF - NSTDA / BIOTEC) for the illustration

cost.

Ontogenetic Basis of Polyad Symmetry in

Samanea saman (Jacq.) Merr.

Surayya Khatoon

Department of Botany,

University of Karachi, Karachi-75270, Pakistan.

Abstract

Samanea saman (Jacq.) Mert. bears more or less radially symmetric polyad. The polyad is formed by eight

equal sized, decussate tetrads, thus consisting of 32 pollen grains in total. In the present work, the ontogeny of

the polyad is studied and the results show that the symmetry of the polyad is effected by the fact that the premeiotic

nuclear divisions in the sporogenous cell precede cell-wall formation.

Introduction

-Compound pollen grains, i.e. tetras and polyads, occur in almost all tribes of

Mimosaceae (Leguminosae-Mimosoideae) with high frequency (Guinet 1981). The

number of pollen grains per polyad ranges from eight to 64 in various members

of the family (Maheshwari 1950). Dnyansagar (1951) has described and illustrated

the morphology of the mature polyad of Samanea saman (as Pithecolobium saman

Benth.) He also described the microsporogenesis, but did not study the premeiotic

events. Kenrick and Knox (1982) for the first time suggested that the pollen mother

cells which give rise to a polyad, descend froin single sporogenous cell in Acacia.

On the basis of their studies on Australian species of Acacia, Knox and Kenrick

(1983) concluded that the number of premeiotic mitoses determined the number

of pollen grains per polyad. The mature polyad is generally highly symmetric,

whatever the grain number may be (Guinet 1983). According to him, this symmetry

is related to the ontogenctic sequence which, however, has not been worked out.

In the present work, premeiotic events are investigated in Samanea saman with a

view to determine the ontogenetic sequence that leads to the symmetry of polyad.

Materials and Methods

Young floral heads of Samanea saman were fixed in Carnoy’s solution (ethanol-

glacial acetic acid in a ratio of 3:1) and stored, at 5°C. The slides were prepared

by squashing the anthers in 1% propionic carmine. Photographs were taken from

temporary mounts.

Results and Discussion

The study of polyad ontogeny in Samanea saman revealed that three rounds

of premeiotic mitosis occur in the sporogenous cell, giving rise to eight nuclei.

However, these nuclear divisions are not immediately followed by cytokinesis and

well-wall formation, thus all these nuclei remain within the same cell. The eight

204

nuclei are arranged in a circle near the periphery of the cell at almost equal

distances from one another (Fig. 1A); then radial walls are laid down and a group

of eight pollen mother cells is formed (Fig. 1B). This group of radially arranged

pollen mother cells undergoes meiosis and gives rise to the polyad, which consists

of eight radially arranged decussate tetrads (Fig. 2A & 2B). Dnyansagar (1951)

has described these tetrads as tetrahedral, but according to the terminology of

Maheshwari (1950), these tetrads are decussate rather than tetrahedral since each

tetrad has two pollen grains arranged in one plane and the other two are in the

plane perpendicular to the first.

Knox and Kenrick, (1983) suggested that the sporogenous cell in Acacia

divides once to form two pollen mother cells in the case of the polyad comprising

eight pollen grains, and that there are two rounds of premeiotic mitosis in the case

of the polyad comprising 12 or 16 pollen grains. All the polyad comprising 8, 12

or 16 pollen grains were symmetric, containing equal sized tetrads. However, Knox

and Kenrick (1983) did not study how the polyad composed of 12 pollen grains

was formed. Guinet (1983) discussed that the symmetry of the polyad implies the

symmetric arrangement of pollen mother cells which form the polyad. However,

’ it has not been explained how the pollen mother cells achieve this symmetry. Their

symmetry seems logical in cases where the pollen mother cells are in even number,

such as 2, 4 or 8; but in those where the pollen mother cells are in odd numbers

(such as 3 or 7, i.e. in the instances of polyad composed of 12 or 28 pollen grains),

it is obvious that one of the daughter cells of the original sporogenous cell remains

undivided, and is therefore larger than other cells. This may result in unequal sizes

of the pollen grains thus disturbing the symmetry of the polyad (cf. Knox and

Kenrik 1983, Fig. lc). The ontogenetic sequence described here for Samanea

saman ensures the formation of equal sized pollen mother cells, as the nuclear

divisions without cell-wall formation allow the nuclei to be arranged at equal

distances from one another, followed by the formation of cell-wall. This

mechanism would also allow the formation of equal sized pollen mother cells in

those instances where the pollen mother cells are in odd numbers. Further studies

in other polyad bearing members of Mimosaceae would reveal if such a mechanism

exists in them also.

Fig. 1. Premeiotic developmental stages of the polyad of Samanea saman

(Jacq.) Merr.:

A. Sprogenous cell with eight nuclei arranged near periphery.

B. Group of eight young pollen mother cells formed by laying down of

radial walls in the 8-nucleate sprogenous cell.

Fig. 2. A. Mature of polyad of Samanea saman (Jacq.) Merr. (unacetolysed).

B. One of the component tetrads drawn separately to show the decussate

arrangement of the pollen grains.

206

Acknowledgements

I am grateful to Prof. Dr. S.I. Ali for developing my interest in the reproductive

biology of Mimosaceae.

References

Dnyansagar, V.R. (1951) Embryological studies in the Leguminosae III. A

contribution to the embryology of Pithecolobium saman Benth. Proc. Indian

Acad. Sci. B. 34: 188-198.

Guinet, Ph. (1981) - Comparalive account of pollen characters in the Leguminosae.

In: Polhill, R.M. and Raven, P.H (eds.) Advances in Legume Systematics, Part

2. Royal Botanic Gardens, Kew, England, 789-799 pp.

Guinet, Ph. (1983). Distribution of the pollen characters in some hybrids and in

some suspected hybrids in the genus Acacia. Bull. Groupe Int. Etude, Mimos.

II: 25-28.

Kenrick, J. and Knox, R.B. (1982). Function of the polyad in reproduction of

Acacia. Ann. Bot. 50: 721-727.

Knox, R.B. and Kenrick, J. (1983). Polyad function in relation to the breeding

system of Acacia, In: Mulcahy, D.L. and Ottaviano, E. (eds.) Pollen: Biology

and Implications for Plant Breeding. Elsevier Science PublistingCo., Inc.; 411-

417 pp.

Maheshwari, P. (1950). An introduction to the Embryology of Angiosperms.

McGraw-Hill International Book Co., Singapore.

Revision of the Genus Zingiber in Peninsular Malaysia

Ida Theilade, Department of Systematic Botany, University of Aarhus, Nordlandsvej 68,

DK-8240 Risskoy, Denmark.

Abstract

Zingiber (Boehm.) comprises nineteen species in Peninsular Malaysia. Seventeen of these are included in the

section Zingiber while two belong to the section Cryptanthium Horan. A new species, Z. fraseri, from Fraser’s

Hill in Pahang, and a new variety, Zingiber officinale var. rubrum, are described. The latter is widely used in

malay traditional medicine. A new combination Z. montanum (Koenig) Theil. comb. nov. is proposed based on

the rediscovery of some of Koenig’s collections from Phuket. Z. griffithii var. citrinum Holtt., and the four varieties

of Z. gracile Jack recognized by Holttum have been ranked as species. Key to the species and varieties are

provided, as well as species descriptions, distribution and specimen citations. The taxa have as far as possible

been typified.

Contents

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Relationships within the Malayan species of Zingiber ...............cceseeeeeeeeeeeeee 154

Origin and evolution .............. Pe ae eae Pe re ian bith Sta S aa. wars icc aeceta ened 155

Key to the genus Zingiber in Peninsular Malaysia.................::ccssccessccceseeeeeeeees 156

NNN Ne 5 2a se yh osc oe 7s 5 cp sos ane sp vtichessidavecvcanésvedaesvdscetenecedive 176

Introduction

The genus Zingiber is distributed throughout tropical Asia with the center of

diversity in Southeast Asia. It is a large and complicated genus comprising of about

100 species (Burtt 1972, Larsen 1980). Within the last few years several undescribed

taxa have been found during field work in China, Thailand and Malaysia, indicating

an even greater species diversity.

Zingiber is a constituent of the undergrowth in the tropical forests. The plants

are perennial and grow mainly in damp places, but they are also frequent in secondary

forest and disturbed sites. In Peninsular Malaysia they are most common in lowland

and mid-mountain forest, but a few species grow on high mountain ridges (Holttum

1950).

The genus is characterized by the long, curved anther appendage enfolding the

style, the three-lobed lip, and the relatively large bracts, each subtending a non-

tubular bracteole and a single flower. At the time of flowering the bracts are usually

orange or red, and often change to a darker colour as they grow older. The lip is

cream or white in some species, and purple mottled with cream in others.

208

The existing classification of Zingiber is based on the eminent works of

Schumann (1904) and Valeton (1918), which are still veryuseful in revising the

genus. Ridley described many new Malayan species in his Flora of the Malay

Peninsula (1924). Similarly the later revision of the genus by Holttum (1950) deals

with the species in the Malay Peninsula. The key presented here largely follows

that of Holttum. Two species, Z. sulphureum Burkill ex Theilade, Z. fraserii sp.

nov., and one variety Z. officinale var. rubrum var. nov. have been added. Z. griffithii

var. citrinum and the four varieties of Z. gracile recognized by Holttum (1950)

have been ranked as separate species. Apart from that only minor additions have

been made. The author is working on a similar revision for Zingiber in Thailand

and in Sabah, Malaysia.

Materials and Methods

The present revision is based primarily on the study of collections of Zingiber

from the following herbaria: AAU, C, E, K, KEP, L, P, PSU, SING, and UKM. Z.

gracile, Z. elatior, Z. auranticum, Z. griffithii , Z. puberulum, Z. montanum and Z.

officinale var. officinale and var. rubrum were studied in the field as well. All

" measurements refer to dried material, except those of floral parts. Flowers were

boiled in spirit prior to measuring. Most of the type collections have been examined.

Notes on the Key

A major difficulty in working with the genus Zingiber is that many of the

collections are incomplete. Most of them include an inflorescence, but often no

flowers. Even when present at the time of collection, the delicate flowers often

deteriorate during the drying procedure. Therefore, vegetative characters and the

structure of the inflorescence have been used in the key whenever possible. Hairiness

is a variable character and has been added only when it is very conspicuous. Colours

of the labellum have been included as supplementary characters. An attempt has

been made to offer a simple and workable key for the genus as it is found in

Peninsular Malaysia, but it does not necessarily reflect the true relationship within

the genus. It should allow identification of species whenever an inflorescence is

available regardless of the presence of flowers. In many cases vegetative characters

are sufficient for a proper identification though some species of Zingiber are difficult

to distinguish based on these characters alone.

It should be kept in mind that the large, juicy inflorescences shrink considerably

when dried. A key with measurements based on live specimens for identification in

the field is desirable, but this will need more field study.

Taxonomic Characters

Measurements of leaves and ligula are taken from the middle of the stem.

Dimensions of inflorescences are given at anthesis. The spikes alter shape according

209

to age, older and fruiting inflorescences are much wider and the bracts are loosely

imbricate. Finally the bracts spread irregularly as the fruits dehisce within them

giving the spike a shaggy appearance. The measurements of the bracts are given

for the lower to middle ones. The lowest bracts are much bigger than the others.

The colour of the bracts changes as they grow older, that at anthesis is indicated in

the descriptions. The length of the calyx, corolla and labellum includes the ovary.

The width of the corolla lobes is measured at the base. For the labellum, the length

of the midlobe is from the junction with the sidelobes to the apex, and the length of

the sidelobes is from the junction with the midlobe to the apex (Fig. 1). The characters

of the ovary, stylodes and stigma are not distinctive and have been omitted in the

descriptions.

Floral Biology and Seed Dispersal

For each species flowering time is given following the description, whenever

data are available. Most species flower mainly between July and September and

fruiting occurs from November to January.

Fig. 1. Labellum outline showing the measurements of the length and width of

the midlobe (q, x) and sidelobes (y, z).

210

The flowers are tubular in their lower part and contain nectar. The yellowish

colour of many species of Zingiber is common of flowers with entomogamy (Jones

1983) and is thought to be important in attracting pollinators. The broad labellum

provides an excellent landing place for insects. The contrasting deep red-purple

markings on the labellum of some species presumably act as nectar guidelines

helping insects to find the entrance to the nectar and pollen store.

The flower is constructed so that the long anther at the back of the flower is bent

forward in the flower mouth with the long, curved, hornlike anther appendage

extending in front of it. The downbent stigma just emerges at the top of the appendage

and occupies the entrance of the mouth a little above the lip. The pollen sacs and

the stigma are oriented towards the labellum. Bees visiting the flower must force

their way in between the anther and lip. Thus the visitor will be covered by pollen,

which at anthesis hangs in loose clusters under the anther. When a visitor leaves

the flower some of the pollen may be rubbed off against the stigma (Valeton 1918).

Whether this results in self-pollination, or whether some kind of self-incompatibility

system has developed, is not known.

Few observations have been made on the pollination of Zingiber, and no details

on the pollination or breeding system of Zingiber have been published. Flowers of

Z. aromaticum open before noon and are regularly visited by swarms of Apis indica

and stray individuals of Anthophora zonata (Valeton 1918). Z. zerumbet is regularly

visited by small, solitary bees collecting pollen (personal observation).

The fruits are dehiscent loculicidally within the persistent bracts. At the time of

fruiting the bracts bend obliquely outwards presenting the black seeds surrounded

by the fleshy, white arils. The conspicuous red bracts together with the fruit valves,

which are bright red inside, and the fleshy, white or yellow aril are suggested to be

an adaptation to bird dispersal. However, so far no observations have been made to

confirm this. Further field studies are necessary to elucidate the pollination and

seed dispersal in the genus.

Relationships Within the Malayan Species of Zingiber

Of the nineteen species of Zingiber described from Peninsular Malaysia sixteen

are thought to be indigenous while the widely cultivated Z. officinale, Z. zerumbet

and Z. montanum are regarded as introduced.

Seventeen of the nineteen Peninsular Malaysian species have the inflorescence

on a radical, erect peduncle and are included in the section Zingiber. Only two

species, Z. wrayi and Z. fraseri, have a radical procumbent or very short peduncle

and belong to the section Cryptanthium.

Within the section Zingiber the species may be divided into two groups. One

group have bracts which are incurved or curved outwards and the labellum mottled

purplish or red-brown and cream. The other group is characterized by closely

overlapping bracts with no curvature, and the labellum cream or white without

211

mottling. Within the first group Z. kunstleri and Z. wrayi have apices of the bracts

pointed and curved outwards. The remaining species in the group have rounded

bracts with incurved margins. Of these, the smaller species Z. officinale, Z. curtisti

and Z. chrysostachys are similar in having light green to yellow bracts, while the

larger species Z. spectabile, Z. ottensii and Z. multibracteatum have yellow-orange,

red or dull purple bracts. Thus, it seems that the character of the bracts together

with the colour of the labellum make up the most important features for classifying

the genus as it occurs in Malaysia. The size of the species and the hairiness are

more variable characters.

The relationship within the group with closely overlapping bracts and cream

labellum is more difficult to outline. The complex of species comprizing Z. griffithii,

Z. puberulum and Z. gracile has long caused much confusion. Holttum (1950)

reported that there seemed to be no clear line of distinction between Z. griffithii

and Z. gracile and possibly hybridization could take place. However, the description

of Z. sulphureum cleared up some of the confusion (Cowley & Theilade 1995). Z.

sulphureum is a small species with ovate leaves like Z. griffithii but an inflorescence

like Z. gracile, and for long it blurred the distinction between the two species.

Thus, it seems that it was an undescribed species rather than a hybridization, which

caused identification problems.

Holttum (1950) recognized four varieties of Z. gracile: var. gracile, elatior,

aurantiacum, and petiolatum although they vary greatly in size and morphology,

and he found it likely that all four varieties would later rank as distinct species. In

fact, Z. elatior was described as a species by Ridley (1899). Z. gracile is a small

plant with a long, thin ligule, while Z. elatior, Z. aurantiacum and Z. petiolatum are

much larger plants, Z. elatior and Z. aurantiacum with short ligules. Z. elatior is

distinguished by its linear leaves, short petiole and pubescent bracts, and as such it

has affinity to Z. montanum. Z. aurantiacum is distinguished by its wider, lanceolate

leaves but otherwise it is much like Z. elatior. Z. petiolatum is a much larger species

distinguished by the tall, leafy stems, the long petiole, large leaves, the extremely

long scape and spike and the tough pink bracts. Z. petiolatum is most similar to Z.

puberulum in the large leaves and tough pink bracts. In this treatment all varieties

have been ranked as separate species.

Origin and Evolution

The Zingiberaceae is a pantropical family. The largest concentration of genera

and species is in Southeast Asia, and judging by the existing distribution of the

family the place of origin was within the Indo-Malayan region (Holttum 1950).

Through geological time, the landmass of the Malayan Peninsula was unaffected

by glaciation, drastic climatic changes or sea flooding, permitting a continuous

evolutionary history of about 140 million years (Flenley 1979, Whitmore 1984).

Presumably Zingiber, like many genera of the family, evolved in the stable climate

in the landmass of the Malay Peninsula and later spread to its present distribution.

Seventeen of the nineteen species of Zingiber found in Peninsular Malaysia

belong to section Zingiber. In the neighbouring territories of Thailand and Borneo

the genus has proliferated not only within section Zingiber but also within section

Cryptanthium. If the ancient rain forest of Peninsular Malaysia is assumed to be

the center of origin of the genus, section Zingiber may be regarded as primitive

while section Cryptanthium is derived or advanced. Thailand and Borneo would

then be secondary centers of diversity.

Most of the species of the Malay Peninsula are large plants with fairly large,

lanceolate or oblong leaves and a relatively tall, erect inflorescence with tough

bracts. It is suggested that these are primitive characters, which are retained in

species like Z. spectabile, Z. multibracteatum, Z. puberulum and Z. ottensii.

However, for a better understanding of the evolution within the genus a proper

cladistic analysis, preferably including chloroplast DNA analysis, would be

desirable.

Key to the genus Zingiber in Peninsular Malaysia

la. ‘Inflorescence: radicalS procuimbenttye rec cnceccceneeccerersceeteeecente care sereesneceaee rere ene eee sect. Cryptanthium 2

Ib: Inflorescence radical) erect, .....cc..cescsecnerenesacoes sveseeoreveascesencctbexsensent ste teaser cdeecestecesneenter festa sect. Zingiber 3

2a. Leafy shoots 3-4 m long, scrambling, leaves lanceolate, inflorescence ovate tapering to a

pointed apex, bracticlosely OVeEml ap pun Py sc. .esacrenc scenes mene see eac ones cases are cra eer cee een eee nena 1. Z. fraseri

2b. Leafy shoots to 2 m long, leaves elliptic, inflorescence cylindric, bracts deflexed

3a. _Bracts with their apical margins incurved; or with their apices curved outwards and free,

not closely imbricating, labellum mottled purplish or red brown and cream ...........2::cc:cseccesseeeescereeneeeeees 4

3b. Inflorescence ovoid to fusiform or cylindric, the bracts closely overlapping their apices not

curved, labellum cream) or whitetwithout mottlin gy cor .ssccccesceessereensceeneseces hes eeenseseeer ences tent eeeeneeanc ee ere 10

4a. Apices of bracts narrowed to a blunt point and curved outwards

4b. Apices of bracts rounded and slightly incurved or with incurved margins .............:cc:ceseeseeseeseseeseeseeeceneneess 5

5a. Leafy shoots'60-1100 cmitall largest leaves}toi20/ crm Lome crccecenessescescsucseeecsnseueesceenerereperce sees tcaeeeeee earner 6

5b. Leafy shoots 150-300(-350) cm tall, leaves commonly 30 cm long OF MOTE ............:::cesceceesesceseeseeeeeeeesesees 8

6a; | Leavesilinear, 15223! Dyslisd=2.5) Crm aie. gence cee re acca see atn cat on cuot pe aces aveeb aie asec rk aentarratanterstesecereoee 4. Z. officinale

6b; Leaves lanceolate or ovate 12-17 Dyi4= SPC ri eeree wcrc cee Cres eccmeneeocerreecessnen cre enasunstlce spent snescber taster teen atest

7a. Labellum closely blotched with purple throughout, including the sidelobes, bracts about

BaBiS DY LS Cm sis ctiastveteces eecentenescencesseectereshcssene teaarbwereenwa sivacesecartatestateestanene seueenencacespstbcteesene 5. Z. curtisti

7b. Labellum with almost entirely crimson midlobe and white sidelobes, bracts about

DeT Wy DA CMM vices Leseeskeoee Aries atabia sciatic aven vane oa stent cons beconumtoadovse ate gud teectcetesrs negtessavendooesnsasees 6. Z. chrysostachys

8a. Inflorescence 12-30 cm tall, cylindric, bracts with their apices free forming open

POUCHES, ox co vccisesececuaneces ove oeigateneeoes suave cnumcere thee te sncsseecht nt cask cette cna scueetek nents Staceacah ce Re eee eens 7. Z. spectabile

8b. Inflorescence usually less than 12 cm long, ovoid to ellipsoid, bracts not forming open pouches............. £

9a. Leaf sheaths and ligule sparsely hairy, labellum pale yellow with faint red-brown

markings, village pla tis tye srecesratcpecetecgaee se ate cte neve ek eb vaca seame sateen thi on Ceneercetyes stetren cess ireeterare 8. Z. ottensii

9b. Leaf sheaths and ligule velutinus, labellum dark purple spotted with cream,

MOUMAIN Plats oso. ccesseedeiccseze-aneesconavesws cers taeedosr vavatoancts seu coe usentenevel esa enenancey esate 9. Z. multibracteatum

10a; Leaves linear; about! 20-30) Dy 2-SiCmy (tock. snes cicenserenal or teteas crveteccetsetnece trans ncateney adres Sonatas oo ttane Nt eRe enEn 11

10b. Leaves lanceolate, 4 cm or more wide, proportionately wider than above ............:...:cssssesceseseeeeseeeseseneene 12

lla. Inflorescence fusiforme or cylindric-ovate, 3-3.5 cm wide, bracts brownish with a

preenimarginy Village plants c.2.0.:2c.cceupsavtectesenecvesvaredes casuteuesbesteetesnvstunctse sWestteas bens vanaingasees 10. Z. montanum

11b. Inflorescence slender fusiform, 2-2.5 cm wide, bracts orange turning red, forest plant .......... 11. Z. elatior

12a. Bracts green when young turning red, slightly convex, ligules 1.5-2.5 cm long, papery,

RTL RESET GE CTI WLC asco ess oee Mechelen rece wn Poms See sae acc Se Tes nas cat Neyo ve bees bestaseeis 12. Z. zerumbet

12b. Bracts pink, yellow or orange sometimes turning red, ligule short or ligule1.5-1.8 cm and

Bs Meg SACEN ATI a IMME WAAC ceeserresca tae oncetaucaees treat cnet verQes ane casneessccrescveestieeeteat ch setecssacdecseert ete oSeensoeree ones atteeaes 13

13a. Leaves lanceolate-ovate 15-30 by 5-10 cm or 12-14 by 4-4.5 CM .......0...ccccccessessesssseesceeescsessecsseecesonsonevess 14

Mnmelvedves lanceolate proportionately MAMLOWEL -:./..c<c-c:r-scesecscesessosursseces deve saceeesssoceoe dussndvococtbesSedebecuccustesevese 16

14a. Leaves 15-20(-25) by 5-8 cm with fine silky hairs below, bracts pink to red .................000000-- 3. Z. griffithii

14b. Leaves 24-30 by 8-10 or 12-14 by 4-4.5 cm, bracts yellow sometimes turning red .............:cceeceseseeeeeees 15

15a. Leaves 24-30 by 8-10 cm, inflorescence cylindrical, 12-15 by 4-5 cm, bracts lemon yellow 14. Z. citrinum

15b. Leaves 12-14 by 4-4.5 cm, inflorescence fusiform, slender, 8-10 by 1.5 cm, bract

VRAD RUNTGRY CAN OW descreettne eerste ae cease erento nen eect cus ons Sonsnvaebaspescaswacdecuedrsistesesvesscuvisissseveces 15. Z. sulphureum

Mies ANCA ESHOOIS eesMtAl LEAVES ILO 2 TNO YA. Ole: cx. se sads-ccserscx-vessncceetentesessstescnaetdndlstasieaducssibasetexstenterdettavosectese 17

Spec ALVES HOOLS 2 -SINitAll) LeAVESILO4O)DV/S \CLIN <<-cs-c0rcceesesesveneaesdcns Senecvectsadeissonci suvacessdosoutaceosssancseasdeceseccsec vee 18

17a. Leafy shoots to 1 m tall, leaves to about 18 by 4 cm, ligule 1.5-1.8 cm long ...............000:000+- 16. Z. gracile

17b. Leafy shoots 1.5-2.0 m tall, leaves 20-27 by 3.5 cm, ligule 0.5 cm long ...............::0000 17. Z. aurantiaum

18a. Leaf sheaths glabrous, inflorescence fusiform, slender, 30-45 cm long, apices of

SEA EST AGEN eset Barren ceases ae Ben arse hecho Scere <sebhren cnet se sckddduc nopspadesusuarsoraaeebosttnasteazest 18. Z. petiolatum

18b. Leaf sheaths hairy to velutinous, inflorescence ovoid, to 15 cm long, apices of bracts

RVEDURES Careers tec nesae sh ctacies svhica bse a dncevsedakucevcesceceasesacsueaicestsdiut costes aceqssacesceessbcvecuprereedesestasseveres 19. Z. puberulum

Zingiber Boehmer in Ludwig, Defin. Pl. (1760) 89; vide Dandy, Ind. Gen. Vasc.

Pl. (1967) 1753-74 (Regn. Veg. 51) 91. - nom. cons.

Adans., Fam. Pl. 2 (1763) 66; Roscoe, Monandr. Pl. Scitam. (1828) 33, t. 83; Horan.,

Prodr. Monogr. Scitam. (1862) 27; Benth. & Hook. f., Gen. Pl. 3 (1883) 646; Baker

in Hook. f., Fl. Br. India. 6 (1892) 243; Schumann in Pflanzenr. Zing. (1904) 163-

64; Holttum, Gdns’. Bull. Singapore. 13 (1950) 48-49. R. Dahlgren et al., Fam.

Monocot. (1985) 364; — Lampujang and Thumung Konig in Retz. Observ. 3 (1783)

62. — Jaegera and Dietrichia Giseke, Prael. ord. nat. pl. (1792) 203, 208. —

Cassumunar Colla, Novi Scitaminearum generis (1830) 1 tab. 1. — Zerumbet Lestib.

in Ann. Sci. Nat. 2. Ser. 15 (1841) 329. —Dymczewiczia Horan., Prodr. Monogr.

Scitam. (1862) 26.

Rhizome horizontal, at or near surface of ground, tuberous, aromatic. Leafy shoots

0.5 - 3.5 m tall, plane of leaves parallel with the rhizome. Leaves distichous, linear

or oblong-lanceolate, sessile or with short petioles, ligules short to long, entire or

deeply bilobed. Inflorescence a spike, usually radical, rarely terminal on the leafy

stem; scape erect, procumbent or very short at base of leafy stem; spike compact,

fusiform or ovoid to cylindrical. Bracts persistent, closely imbricating or with apices

free, single flowered, at first green, yellow or reddish turning bright yellow or red.

Bracteoles one to each flower facing the bract, narrower than the bract, usually

persisting and enclosing the fruit. Calyx hyaline, tubular-spathaceous, usually shorter

than the bracteole, tripartite. Corolla tube slender; dorsal lobe broader than lateral

lobes, concave; lateral lobes below the lip, usually joined partly together by adjacent

sides and to the lip. Labellum 3-lobed, cream to white with or without purple

mottling; midlobe oblong-obovate, apex retuse or cleft; lateral staminodes oval or

214

acute, adnate to the midlobe. Filament short; anther 1-1.5 cm long, narrow;

connective prolonged into a slender, curved beak embracing the style. Stigma

protruding just below the apex of the appendage, with a circular apical aperture

surrounded by stiff hairs. Ovary trilocular with two, slender epigyneous glands.

Fruit a capsule, oblong with a fleshy wall when fresh, more or less leathery when

dry, trilocular with axil placentation, dehiscent loculicidally within the persistant

bracts. Seeds numerous, ellipsoid, black or maroon, covered by a saccate, fleshy,

white or yellow, lacerate aril.

Type: Zingiber officinale Roscoe, Trans Linn. Soc. London 8 (1807) 358;

Monandr. Pl. Scitam. (1824-1828) t. 83.

Distribution: Throughout tropical Asia and to tropical Australia and Japan.

Ecology: Tropical evergreen and monsoon forests, in moist, humus rich soils in

shady habitats. Secondary forests, open habitats at the edges of the forests, disturbed

places, and bamboo thickets on rocky ground, up to 3000 m.

Uses: The only species extensively used as a flavouring in food is the true ginger

_ Z. officinale. Several species of the genus are known to be used in local medicine

throughout Asia. Some species are grown as ornamentals.

1. Zingiber fraseri Theilade sp. nov. (Figs. 2 & 3)

Type: Peninsular Malaysia, Bukit Fraser, trail 4, behind the golf course, Sep.

1993, Theilade 12 (AAU holotype!, K isotype!).

Species Z. griffithii affinis a qua differt surculis longis scadentibus, foliis

angustioribus lanceolatis glabris, scapo brevi procumbente, inflorescentia ovoidea

in apicem acuto angustata, bracteis et bracteolis longioribus

Leafy shoots slender, to 4 m long with a scrambling habit, leaf sheaths finely

hairy. Ligula 2 mm, hairy. Petiole 2 mm, hairy. Leaves lanceolate, 14-20 by 3.5-5.0

cm, glabrous. Inflorescence radical, procumbent. Scape to 8 cm, whitish. Spike

ovate to fusiform, 12 by 3.0 cm, tapering to a pointed apex. Bracts lanceolate, 4.0-

5.0 by 1.5 cm, bright red, hairy, apex acuminate. Bracteole 3.5 by 0.7 cm, whitish.

Calyx 3.0 cm long. Flowers unknown.

Z. fraseriis related to Z. griffithii but differs in the long, scrambling leafy stems,

the narrower, lanceolate, glabrous leaves, the short, procumbent scape, the ovoid

inflorescence tapering to a pointed apex and the longer bracts and bracteoles.

Vegetatively Z. fraseri is easily recognized by the long, scrambling leafy stems,

which is unlike any Zingiber described so far.

Collections other than type: Bukit Fraser, trail 5 behind Rompin’s House, Nov.

1995, Theilade 68 (AAU). In cultivation at Waimea Botanical Gardens, Hawaii.

Ecology: Montane forest at 1300 m altitude. Flowering in August-September,

fruiting in October. Probably endemic to Fraser’s Hill.

5cm

Fig. 3. Zingiber fraseri Theilade sp. nov. Leafy shoot and inflorescence.

216

2. Zingiber wrayi Ridley, J. Straits Brch. R. Asiat. Soc. 41 (1904) 31; Mat. FI.

Malay. Penins. 2 (1907) 27; Fl. Mal. Pen. 4 (1924) 259; Holttum, Gdns’. Bull.

Singapore 13 (1950) 53.

Leafy shoots to 1.80 m, lower sheaths flushed with purple. Ligula bilobed, lobes

broadly rounded, to 5 mm long, thin, glabrous to sparsely pilose. Petiole 2 mm.

Leaves lanceolate to evenly elliptic, 30-40 by 7-10 cm, glabrous, apex acute.

Scape radical, procumbent, 10-30 cm; sheaths 5—8 cm long, slightly hairy at tips.

Spike cylindric—ellipsoid, later almost globose, 6-13 by 5-10 cm. Bracts oblong

to evenly elliptic, 4-5 by 2-3 cm, red, slightly hairy, apex shortly pointed, with

slightly inflexed edges. Bracteole lanceolate, 3-4 by | cm, tinged with pink,

sparsely pilose, apex acute Calyx 2.8 cm long, sparsely pilose. Corolla 6.5 cm

long; dorsal lobe 2.0 by 1.2 cm; lateral lobes more narrow, lobes pale yellow.

Labellum 6.0 cm long, pale yellow mottled with purple; midlobe ovate, 1.2 cm

long, apex slightly retuse margin crenate; sidelobes oblong, 0.8 cm long. Anther

appendage dark purple.

Type: Peninsular Malaysia, Upper Perak, Wray 3735 (SING holotype!, K

isotype!).

Collections: Corner 37054; A. Latiff 88; Md. Nur 18569; Md. Shah & Md.

Nur 1990; Moysey & Kiah 33712; Sinclair 6181.

Distribution: Peninsular Malaysia and Peninsular Thailand.

Malaysia: Johor, Pahang, Perak, Terengganu.

Ecology: At streams, to 800 m. Flowering in July to September.

Note: Easily recognized by the deflexed bracts and the broad leaves. The

inflorescence of Z. wrayii has affinity to Z. kunstleri but Z. wrayi is a smaller plant

and the leaves are broader.

3. Zingiber kunstleri Ridley, J. Straits Brch. R. Asiat. Soc. 32 (1899) 127-128;

Mat. Fl. Malay. Penins. 2 (1907) 27; Fl. Mal. Pen. 4 (1924) 259; Schumann in

Pflanzenr. Zing. (1904) 180; Holttum, Gdns’. Bull. Singapore 13 (1950) 52.

Rhizome internally purplish—lilac. Leafy shoots to 2 m tall, slightly swollen at

base with a pale lilac hue, internally purplish—lilac. Leaf sheath glabrous. Ligula

bilobed, lobes broadly rounded, 1—2 mm long, glabrous. Petiole 4—5 mm. Leaves

linear-lanceolate, 45 by 6 cm, glabrous, apex acuminate. Scape radical, erect, 30

cm; sheaths 7.5 cm long, slightly hairy at the tips. Spike ovate to cylindric, later

globose, to about 14 by 9.0 cm. Bracts densely imbricate, linear to slightly

lanceolate, about 6 by 2-3 cm, pink to red, glabrous, apex narrowed, bluntly

pointed, curved outwards or deflexed. Bracteole nearly 5 cm long. Corolla shorter

than bracts, pale white. Labellum narrow, lanceolate, shorter than corolla lobes,

reddish-brown, apex acute; side-lobes hardly distinct.

217

Type: Peninsular Malaysia, Taiping Hills, Ridley 11449 (SING lectotype!, K

isotype!).

Collections: Corner 30588; Kunstler 2219 (drawing only); Kiah s.n. July 1936;

Sinclair & Kiah 38784; Ridley 2401.

Malaysia: Pahang, Perak, Terengganu.

Ecology: In swamps and in hills, 150-950 m, flowering from late July until

August, fruiting in November.

Note: Z. kunstleri is easily recognized by the deflexed bracts and the linear

leaves.

Affinities: In the deflexed bracts Z. kunstleri is similar to Z. wrayi, but the leaves

are narrower and the inflorescence is larger.

4. Zingiber officinale Roscoe, Trans. Linn. Soc. 8 (1807) 348; Valeton, Bull. Jard.

bot. Buitenz. 27 (1918) 128; Holttum, Gdns’ Bull. Singapore. 13 (1950) 54; Backer,

Fl. Java. 3 (1968) 45-46. — Amomum zingiber Linné, Sp. Pl. 1 (1753) 1. — Curcuma

longifolia WALL. (1828), Cat. No. 6612.

Key to varieties:

la. Rhizomes yellow externally, leafy shoots green basally, labellum dark purple mottled

RPMI RIMS ERAN TMs ee en cease nec one a ee coun ccd cae ace sc aasa ss renues aes seas cucu. gaucstcds seceUeweaseeusoycesnesvs cab var. officinale

lb. Rhizomes reddish externally, leafy shoots red basally, labellum scarlet red mottled with

MEM AMA eer eee oa ren eas acess savudue recs spus cece teusnverrsansdacudvtspvbataccssicaessacdbecsosscsdtucandssesteecaessascsteesdesvexeescade var. rubrum

var. officinale

Rhizome yellow inside. Leafy shoots 0.5-1.0 m, glabrous except for short hairs

near base of each leaf-blade. Petiole 2 mm long. Ligula 3-5 mm, slightly bilobed,

glabrous. Leaves linear, 15-23 cm by 1.5-1.8 cm, narrowed to a slender tip. Scape

radical, erect, slender 15-20(-30cm) tall; sheaths tightly appressed, 4-5 cm long,

the upper sometimes enlarged and leaf-like. Spike elliptic or oblong, 4.0-5.0 by

1.5-2.0 cm. Bracts obovate, 2.0-3.0 by 1.5-2.0 cm, light green turning yellow,

glabrous with a thin, slightly incurved, margin. Bracteole elliptic, 2.5-3.0 cm long,

often longer than the bract. Calyx 1.2 cm long. Corolla 4.5 cm long, lobes yellow;

dorsal lobe 1.8 by 0.8 cm; lateral lobes 1.6 by 0.6 mm. Labellum dull purple mottled

with cream; midlobe circular, entire, 1.2 cm long; side-lobes rather narrow, 0.6 by

0.4 cm, acute. Anther cream. Appendage dark purple. Capsule red. Pollen globose

with cerebroid sculpturing.

Type: Drawing in Roscoe Monandr. PI. Scitam. t. 83 (1828). Lectotype here

selected. No specimen at LINN or in Herb. Cliff (BM) (Burtt 1972).

Distribution: Cultivated in tropical Asia from ancient times and now through-

out the tropics. Its country of origin is unknown. It is considered to be indigenous

218

to Eastern India (Dahlgren et al., 1985). On the other hand it is suggested that the

area of origin could be somewhere between the Yangtze and Yellow Rivers, and

that due to great changes in natural environments wild ginger disappeared from

its original range (Wu, 1985). Z. officinale grows well in Thailand and Malaysia.

Ecology: Occurs as naturalized on wasteground, in deciduous forests and dry

rocky bamboo thickets.

Uses: Medicinal plant and spice throughout Southeast Asia. Various races are

grown in Malaysia and Thailand where it is a common village plant. The aromatic

rhizomes are widely used in cooking or made into marmelade. Occasionally the

young rhizomes and parts of the stem are eaten raw. It is also boiled, sun-dried and

made into powder, and then used as a condiment to flavour cakes and ginger-beer.

Commercially, Jamaica, India, Sri Lanka and China are the most important producer

countries.

Medicinally, the rhizome is considered to have diaphoretic, stomachic,

carminative and diuretic properties. It is prescribed for heavy colds, coughs and

congestion of the chest. It is also used as a remedy for diarrhoea and dysentery. The

“juice of the rhizome is used to treat migraine and it relieves menstrual cramps.

Various lotions, decoctions or poultices may be rubbed on the body after childbirth,

applied to swellings, contusions, for rheumatism and further as a bath to lower

fever. The crushed rhizome is often smeared on the head for headaches. In China it

is used as an antidote to fish and crab poison.

Malay name: “Halia betul” or “real ginger”.

Notes: Z. officinale 1s recognized by the linear leaves and glabrous ligula; the flowers

are characterized by the dull purple labellum which is spotted with yellow.

Affinities: In its narrow leaves Z. officinale resembles Z. montanum Roxb. but

the latter has much taller leafy stems and may be distinguished by its hairy ligules

and ferruginous, pubescent bracts.

var. rubrum Theilade var. nov. —Haliya padi Valeton, Bull. Jard. bot. Buitenz. 27

(1918) 128; —Halia bara or padi and Halia udang Holttum, Gdns’ Bull. Singapore

13 (1950) 54. (Figs. 4a & 4b)

A varietati officinale differt rhizomatibus minoribus externe rubellis, surculis

foliosis ad basin rubris, petiolis rubellis, labello scarlatino cremicolore maculato.

Rhizome reddish outside, yellow inside, pungent. Leafy shoots 0.5-1.25 m, basally

dark red, glabrous except for short hairs near base of each leaf-blade. Petiole 2 mm

long, more or less reddish. Ligula 1-5 mm, slightly bilobed, glabrous. Leaves linear,

25-32 by 1.5-1.8 cm, narrowed to a slender tip. Scape radical, erect, slender 20-30

cm tall; sheaths tightly appressed, 4-5 cm long. Spike elliptic or oblong, 4.0-5.0 by

1.5-2.0 cm. Bracts obovate, 2.0-3.0 by 1.5-2.0 cm, green with a thin, slightly involute

Fig. 4b. Zingiber officinale var. rubrum Theilade var. nov. Leafy shoot showing

reddish petioles.

220

margin. Bracteole elliptic, 2.5-3.0 cm long, often longer than the bract. Calyx 1.2

cm long. Corolla 5.0 cm long, yellow. Labellum scarlet red mottled with cream;

midlobe circular, 1.5 cm long, apex retuse; side-lobes oblong, 6 by 4 mm. Anther

cream. Appendage dark purple. Capsule red. Pollen globose, sculpturing cerebroid.

Type: Malaysia, Sabah, Kota Kinabalu, Central Market, Theilade 66 (SAN

holotype! incl. spirit coll. and photos, AAU, SING isotypes). In cultivation at AAU.

Collections other than type: Weber s.n. 1988; Theilade 60 in cultivation at UKM.

Distribution: Cultivated in Southeast Asia for medicinal purposes and as a spice.

Frequently sold at markets in Malaysia.

Ecology: Z. officinale var. rubrum grows well in Malaysia, but rarely flowers.

Uses: Var. rubrum is grown on a small scale for medicinal use and as a spice.

Various lotions, decoctions or poultices may be rubbed on the body after childbirth.

The juice of the rhizome is used to relieve menstrual cramps. See Burkill (1966)

for a more extensive account on medicinal uses.

Malay names: The Malays recognize more kinds of the red variety; “halia merah”

‘(merah means red), “halia padi” (small), “haliya bara” (hot coals) and “halia

hudang” (prawn). Whether these differ is not certain, but they are all used

medicinally like “halia padi’. Ridley (1912) regards “halia bara” as synonymous

with “halia padi’.

Notes: Z. officinale var. rubrum differs vegetatively from var. officinale by the

smaller, red colored rhizomes which have a stronger, more pungent smell, the red

coloring of the basal parts of the leafy stems and petioles and larger leaves.

Furthermore, the labellum is larger and scarlet red mottled with cream.

According to Burkill (1966) Rumphius in his Herbarium Amboinense (1747)

first described two varieties of Z. officinale as Z. majus— the larger plant known in

Malay as “halia” and Z. minus— the smaller plant, or “halia padi’. Of the latter

Rumphius distinguished two forms. Valeton (1918) stated that the Javanese “haliya

padi” or “sunti” differs from normal Z. officinale in the rhizome and the ovate

staminodes with a rounded base. This study has shown that the size of var. rubrum

varies greatly with growing conditions. Grown in the villages the plant is usually

smaller than var. officinale, but a rhizome brought back to Denmark and cultivated

at AAU in rich soil grew 1.25 m tall and has larger leaves. The red coloring of the

petioles was most distinct at the time of flowering. It remains to ascertain how

many races enter into var. rubrum and whether they genetically differ in size.

5. Zingiber curtisii Holttum, Gdns’ Bull. Singapore 13 (1950) 54-55. (Figs. 5)

Leafy shoot 0.6-1.0 m, slender, the lower sheaths flushed with purple. Ligula 4-

5 mm long, thin, hairy towards the base. Leaves ovate, 12-17 by 4-5.5 cm, base

broadly cuneate, apex acute, glabrous except for the hairy base of the lower surface

221

of the midrib. Scape aboutl0 cm long, sheaths purple. Spike to 14 by 3 cm,

cylindrical, apex obtuse. Bracts elliptic, 3-3.5 by 1.5 cm, pale yellow-green,

glabrous, apex obtuse and slightly inflexed. Bracteole slightly shorter than bract.

Calyx 2.0 cm long. Corolla 5.5 cm long, lobes about 2.0 cm long, white. Labellum

same length as corolla lobes, with deep purple markings throughout including the

sidelobes. Anther appendage deep purple.

Type: Peninsular Malaysia, Bujong Malacca, August 1898, Curtis s.n. (SING

holotype!).

Notes: This species is not distinguishable vegetatively from Z. chrysostachys.

However, the inflorescence is longer and more slender, the bracts are longer and

narrower, pale yellow green, only slightly inflexed at the tips and forming a closer

spike. The lip has deep purple markings throughout including the sidelobes and the

anther appendage is deep purple. The type was cultivated in The Botanical Garden,

Penang where it flowered in June 1902. Known only from the type collection and a

coloured drawing by Hussein done when the species flowered in Penang and now

deposited in the Singapore Herbarium.

6. Zingiber chrysostachys Ridley, J. Straits Brch. R. Asiat. Soc. 32 (1899) 129; Fl.

Fig. 5. Zingiber curtisii Holttum. Fig. 6. Zingiber chrysostachys Ridley.

Drawing by Hussein. Drawing by De Alwis.

222

Mal. Pen. 4 (1924) 260; Holttum, Gdns’. Bull. Singapore 13 (1950) 55. (Fig. 6)

Leafy shoots 0.6 m tall. Leaf sheaths the lower sheaths flushed with purple.

Ligula 5 mm, glabrous or hairy, bilobed; lobes green with scarious margin. Leaves

lanceolate, 12-17 by 4.0-5.5 cm, glabrous except for the hairy base of the midrib

below, base broadly cuneate, apex acuminate. Scape radical, erect, 7-10 cm long,

often two or more in succession from the same rhizome; sheaths flushed with purple.

Spike oblong, to 10 cm long and 4 cm wide, apex rounded. Bracts loosely

imbricating, obovate-suborbicular, 2.5-3.0 by 2.3-2.5 cm, bright yellow, sparsely

hairy, convex with inflexed upper margin. Bracteoles 2.5 cm long, hairy at base.

Calyx 1.5 cm long. Corolla about 4.5 cm long, pale yellowish. Labellum as long as

corolla lobes; midlobe obovate, 1.2 cm long, almost entirely crimson with irregular

white margins, apex broadly rounded; side-lobes ovate, 0.8 by 0.5 cm, white,

spreading to a width of 1.5 cm when flattened. Anther appendage mottled pink to

red. Pollen globose with cerebroid sculpturing.

Type: Peninsular Malaysia, Bukit Stau, 1891, Ridley s.n. (SING lectotype!, K

_ isotype!). Lectotype here selected.

Collections: Burkill & Haniff 13830; Curtis 2716; Hervey s.n. 1889, Kiah S.341;

Ridley 5199; SEN 35834; Wray 3549, 3110; Zainudin 4471.

Malaysia: Kedah, Perak.

Ecology: Evergreen forest and dry bamboo forest on limestone hills, 200-

1400 m.

Uses: In 1924 a bomoh (Malay traditional healer) of Grik in Upper Perak brought

leaves of what appeared to be this species to Burkill and Haniff stating that in fever

a decoction is administered. He called it “/empui’, a form of “/ampoyang” (Burkill

1966).

Affinities: Z. chrysostachys is most closely related to Z. curtisii but the latter

has the labellum coloured purple throughout including the side-lobes. No other

small species has yellow bracts of this character. In the inflexed bracts and red-

marked lip Z. chrysostachys appears to be related to Z. ottensii and Z. spectabile

but it is a very much smaller species than either.

7. Zingiber spectabile Griffith, Not. pl. asiat. 3 (1853) 413; Baker in Hook. f, FI.

Br. India 6 (1892) 247-248; Ridley, Mat. Fl. Malay. Penins. 2 (1907) 26-27; FI.

Mal. Pen. 4 (1924) 258; Holttum, Gdns’. Bull. Singapore 13 (1950) 56.

Leafy shoots 2.0—3.5 m, basal leafless part to | m tall, swollen at base. Leaf

sheaths sparsely pilose, margin scarious. Ligula deeply bilobed, the lobes to 1.5

cm long, broad, pale green. Leaves lanceolate, 30-50 by 6-10 cm, glabrous or slightly

hairy at the base below. Scape radical, erect, 20-40 cm; sheaths 5 cm long, green to

reddish. Spike cylindric, 10-30 by 6-7 cm, apex rounded. Bracts obovate, 4.5 cm

223

long, at first yellow turning orange, when old entirely red, fleshy, curved outwards

with the broadly rounded edge incurved forming open pouches. Bracteole linear,

to 4 cm. Calyx to 35 mm long, cream—pinkish. Corolla 70 mm long, yellow; dorsal

lobe to 30 by 17 mm; lateral lobes 18 by 6 mm. Labellum 40-60 mm, dark purple

with yellow spots; midlobe ovate, 16 by 14 mm, shorter than or of equal length

with the lateral corolla lobes, apex cleft; side-lobes broadly rounded, 10 by 10 mm.

Anther yellow. Anther appendage purple. Capsule ovoid, 30 by 10 mm, sparsely

pilose. Pollen spherical with cerebroid sculpturing.

Type: Peninsular Malaysia, Malacca, Griffith 5762 (K lectotype!).

Collections: Burkill 1985; Corner 1587; Curtis 1978, 2161; Henderson 21851;

Dr. King’s Collector s.n., 3205; Maingay 1567; Nur 34215; Ridley, s.n. (1895);

Sinclair 7856; Wray 3578.

Distribution: Peninsular Thailand and Peninsular Malaysia

- Malaysia: Penang, Perak, Terengganu, Pahang, Selangor, Negri Sembilan.

Ecology: In evergreen forests, along trails, roadsides, streams and edges of the

forest, on hillsides, disturbed sites, up to 1000 m. Flowering July till September.

Fruiting in November.

Uses: Its leaves may be pounded and used for poulticing swellings. Used by

Orang Asli against headaches and backaches. Sometimes used by the Malays as a

flavouring (Burkill 1966). The tall, colourful inflorescences are sometimes cut and

used as ornamentals.

Affinities: Z. spectabile has affinity to Z. ottensii, but it is a much larger species

easily recognized by the large, orange inflorescence with incurved bracts forming

open pouches. It has the largest inflorescence of any Malayan species.

8. Zingiber ottensii Valeton, Bull. Jard. Bot. Buitenz. 27 (1918) 136, t. 19. Ridley,

Fl. Mal. Pen. 4 (1924) 259.

Rhizome pale grey-purple within having a pungent smell. Leafy shoots to 1.5 m

tall. Leaf sheaths broad, slightly hairy near base and apex. Ligula broad, thin, entire,

to about 1.2 cm long, hairy towards the base. Petiole 5 mm, finely hairy. Leaves

elliptic or widest above the middle, 35(-40) by 6(-8) cm, lower surface slightly

hairy towards the base, apex acuminate. Scape radical, erect, 25-40 cm. Spike evenly

ellipsoid to cylindrical with a broad apex, 10-12 by 4 cm. Bracts obovate, 4 cm

long and almost as wide, convex with incurved tips, dull red to bright red when

old. Bracteole linear to lanceolate, 3.2 cm long. Calyx 2.3 cm long, white. Corolla

5.7 cm long, cream to yellow; dorsal lobe 2.2 by 1.1 cm; lateral lobes 2.0 by 0.6

mm. Labellum 5.5 cm long, pale yellow with faint red-brownish markings; midlobe

oblong almost round, 2.0 by 1.5 cm, apex rounded and slightly cleft; side-lobes

ovate, 1.5 by 0.9 cm. Anther 1.2 cm long, pale yellow. Capsule oblong, red.

224

Type: Java, Bogor, Ottens 676 (L lectotype ex Hort. Bot. Bog.!, K isotype).

Lectotype here selected.

Collections: Curtis Apr. 1900; Birch Oct. 1901; Burkill July 1914; Henderson

20200; Holttum 17671; Ridley 7799, s.n. Jun 1893.

Distribution: Thailand, Malaysia, Java and Sumatra.

Malaysia: Penang, Kedah, Selangor, Terengganu.

Uses: Z. ottensii is cultivated and used in traditional medicine. The rhizomes

are pounded into a poultice used after childbirth.

Malay name: “Lempoyang hitam” or “bonglai hitam” referring to the grey-purple

colour of the rhizome.

Affinities: Z. ottensii is closely allied to Z. zerumbet but differs in the convex

bracts with incurved tips and the pale yellow flowers with faint red or brownish

markings. Furthermore, Z.oftensiii can be distinguished by the rhizome, which is

dark purple inside in contrast to the yellow rhizome of Z. zerumbet and Z.

- cassumunar.

9. Zingiber multibracteatum Holttum, Gdns’ Bull. Singapore. 13 (1950) 57.

Key to varieties:

la. Leaves 6.0-9.5 cm wide, spike ovoid to oblong, bracts dull purple ...............:.:e:ceeee var. multibracteatum

lb. Leaves to 5.0 cm wide, spike cylindrical, bracts light green ................:.:csscsseesceseeceeeceeescneceseeeeneens var. viride

var. multibracteatum

Leafy shoots to 3 m tall. Leaf sheaths brownish velutinus. Petiole 2-3 mm long,

velutinus with brown-yellowish hairs. Ligula 3-5 mm long, brownish velutinus.

Leaves lanceolate or obovate, 35-40 by 6.0-9.5 cm, lower surface sericeus, midrib

pubescent or densely so, edge hairy in the young leaves, apex acute. Scape radical,

erect, 25-85 cm; sheaths pubescent. Spike ovoid to oblong, 10-12 by 4.5-7.5 cm,

apex obtuse. Bracts obovate, convex, 3.5-4.0 by 3.2 cm, dull purple, pubescent,

apex broadly rounded, incurved, margin 1.5 mm wide, thin. Bracteoles lanceolate,

3.0-3.5 by 1.5 cm. Calyx tubular, 3.0 cm long. Corolla 8.0 cm long, lobes 3.0 cm

long, reddish. Labellum purple, spotted cream; midlobe obtuse, 2.5-3.0 cm long,

apex retuse; side-lobes oblong, 0.8 cm long. Capsule 2.0 cm long. Seeds maroon.

Type: Peninsular Malaysia, Pahang, Fraser’s Hill, 1300 m. Corner S.R.N. 33174

(SING holotype!, K, L isotypes).

Collections: Burkill & Haniff 12765; Mohd. Shah et al. 1059; Nur S.E.N. 32869;

Sinclair 6083, 38693; Theilade 17, 18.

Malaysia: Kelatan, Pahang, Perak.

Ecology: Common in the Cameron Highlands, 1100-1300 m.

223

Notes: Z. multibracteatum is a large plant only found at high altitudes. It is

characterized by the velutinus petiole and ligule, the very broad ovoid inflorescence

of convex, firm, dull purple bracts with thin edges, and the large flowers with

purple cream-spotted lip.

Affinities: Vegetatively Z. multibracteatum is similar to Z. puberulum Ridl., but

in the inflorescence and colour of labellum it is clearly allied to Z. spectabile and Z.

ottensii.

var. viride Holtum (1950), l.c.

Z. multibracteatum var. viride differs from the typical form in having leaves to

5.0 cm wide, a cylindrical inflorescence and bracts to 5 cm wide, light green.

Type: Peninsular Malaysia, Cameron Highlands, Tanah Rata, Holttum s.n. Aug.

1946 (SING holotype).

- Collections: Lewis 166.

Malaysia: Pahang.

Ecology: Upper montane forest in Cameron Highlands, 1800-2000 m.

10. Zingiber montanum (Koenig) Theilade comb. nov.— Basionym Amomum

montanum Koenig in Retz. Observ. 3 (1783) 51. — Z. purpureum Roscoe, Trans

Linn. Soc. 8 (1807) 348; Scitaminae (1828) t. 85. — Zingiber cassumunar Roxb.,

Asiat. Res. (1810) 347, t.5; Fl. Ind. 1 (1820) 49. Bot. Mag. t. 1426. Schumann in

Pflanzenr. Zing. (1904) 179. Valeton, Bull. Jard. bot. Buitenz. 27 (1918) 138, t.15

f.13, t.20 f.14,15. Ridley, Fl. Mal. Pen. 4 (1924) 259. (Fig. 7)

Rhizome pale carrot colour internally, strongly aromatic. Leafy stems 1.2-1.8 m,

sheaths glabrous or hairy near edges. Ligula bilobed, about 2 mm, hairy. Leaves

linear, 20-35 by 2-4 cm, pubescent beneath, evenly narrowed to the tip. Scape

erect, 20-25 cm long. Inflorescence fusiforme or cylindric-ovate, 10-16 by 3.0-3.5

cm, apex acute. Bracts ovate, 3-3.5 cm long, brownish with green, papery edges,

pubescent. Bracteole 1-1.5 cm. Calyx 1.2 cm. Corolla 6.0 cm long, pale yellow.

Labellum 6.0 cm long, pale yellow; midlobe broadly rounded, 2.0 cm, apex bilobed,

deeply split when old; side-lobes oblong.

Type: Thailand, Phuket, Koenig s.n. (C holotype!).

Collections: Burkill & Haniff 14066, 16482, 17551; Gianno 416.

Distribution: This species is probably native to India. In Sanskrit it is called

“yanaardraka’’, in Hindi “banada” and in Kannada it is called “kadu shunti”’. All

the names signify that it is a “wild ginger’, i.e. a forest ginger in contrast to the

cultivated Z. officinale. It occurs widely in Southeast Asia as a village plant.

According to Holttum (1950) it is not very common in Malaysia.

Pa ww

Pe Sra

Fig. 7. Zingiber montanum (Koenig) Theilade comb. nov. Inflorescence with

opened flowers.

Malaysia: Pahang, Perak.

Uses: Z. montanum is used in traditional medicine all over tropical Asia. It is

primarily a carminative and a stimulant for the stomach used in cases of diarrhoea

and colic. In Malay medicine the rhizome is rubbed on the body to reduce fever

and to heal a person infiltrated by spirits. Other uses of the plant are very similar to

those of Z. officinale and it can be regarded as a substitute for this species.

Vernacular names: In Ayurvedic medicine Z. montanum is sometimes called

“camphor ginger” to distinguish it from Z. zerumbet called “stone ginger”. The

latter has some bitterness in the taste while Z. montanum has not. In Malaysia Z.

montanum is called “bunglai” or “bolat’.

Notes: Koenig’s specimens from Phuket have long been considered lost at sea and

it was impossible to verify the species he named from his rather short descriptions.

Recently some of his collections were rediscovered in the herbarium in Copenhagen

including a collection of this species to which Koenig then gave the name Amomum

montanum providing the first valid epithet. Not withstanding the above findings

the species has for long wrongly been named Z. cassumunar. As Roscoe described

the species in 1807 as Z. purpureum this provides an earlier epithet than cassumunar

used by Roxburgh three years later in 1810.

Affinity: This species is related to Z. zerumbet but can be easily distinguished

227

by the linear leaves and very short ligules as well as the brown bracts.

11. Zingiber elatior Ridley (1899), J. Straits Brch. R. Asiat. Soc. 32: 130; (1924)

Fl. Mal. Pen. 4: 260. — Z. gracile var. elatior Holttum (1950), Gdns’ Bull. Singapore

13: 64.

Leafy shoots to 2 m tall. Ligula bilobed, 4 mm, pubescent. Leaves linear, 20-30

by 2.0-2.8 cm, lower surface and the midrib towards the base pubescent. Scape to

30(-40) cm long. Spike slender fusiform, 10-20(-25) cm long and 2.0-2.5 cm wide.

Bracts orange turning red, pubescent along margin. Bracteoles 2.5-3.0 cm Calyx

about 2.5 cm long. Corolla 6.0 cm cream. Labellum deeply bifid, yellow.

Type: Peninsular Malaysia, Penang, Apr. 1896, Ridley 9340 (SING lectotype!,

K isotype!). Lectotype here selected.

Collections: Burkill 1529, 3312; Burkill & Haniff 12712; Dr. King 7954; W.

Fox 61; Mohd. Shah & Sanusi 2166; Ridley 7954, Aug. 1904, Dec. 1905.

Affinities: Vegetatively Z. elatior resembles Z. montanum, a species native to

India, in the linear leaves. Z. elatior is recognized by the somewhat narrower leaves,

and the slender fusiform inflorescence with orange to red bracts.

Malaysia: Johore, Penang, Perak, Selangor.

Ecology: Up to 1150 m.

Notes: The plants here represent a distinct species as proposed by Ridley. Whether

the lip has always the small red and black lines reported only by Burkill for SFN.

3312 is unknown.

12. Zingiber zerumbet Smith, Exot. bot. 2 (1805) 105 t. 112; Roscoe, Trans. Linn.

Soc. Lond. 8 (1817) 348; Monandr. pl. Scitam. (1828) 35; Blume, Enum. pl. Javae

(1827) 42; Roxb., Fl. ind. (1832) 47; Baker in Hook. f., Fl. Brit. India 6 (1892) 247;

Hook. f. in Trimen, Handb. fl. Ceylon 4 (1898) 259; Schumann in Pflanzenr. Zing.

(1904) 172-173; Ripvey, Mat. Fl. Malay. Penins. 2 (1907) 27-28; Gagnepain in M.

Lecomte, Fl. Gen. Indo-Chine 6 (1934) 84-85: Valeton, Bull. Jard. bot. Buitenz.

27 (1918) 129-131; Holttum, Gdns’. Bull. Singapore 13 (1950) 59-60; Backer &

Bakh. f., Fl. Java 3 (1968) 45. — Ammomum zerumbet Linn., Sp. pl. 1 (1753) 1. —

A. spurium J. Gmelin, Syst. nat. 1 (1792) 6. — Z. amaricans Blume, Enum. pl.

Javae 1 (1827) 43. — Z. aromaticum Valeton, Bull. Jard. bot. Buitenz. 27 (1918)

131-133 — Z. ovoideum Noronha, Verh. Batav. Genootsch. Kunst. Wet. ed. 1 5(4)

(1790) 28; Blume (1827), l.c. — Z. spectabile Griffith, Not. pl. asiat. 3 (1851) 412,

Icon. (1847) t. 351. — Z. truncatum Stokes, Bot. mat. med. | (1848) 68. — Z.

blancoi Hassk. in Flora Philip. 47 (1864) 20. — Z. aromaticum Valeton, Bull. Jard.

bot. Buitenz. 27 (1918) 131; Ridley, Fl. Mal. Pen. 4 (1924) 259. — A. silvestre

Poiret, Encycl. met. bot. suppl. 5 (1817) 548. — A. zingiber Blanco, FI. Filipp. ed.

3 (1837) 2-3, non L. — Zerumbet zingiber Lestib. f. in Ann. Sci. nat. 2. ser. 15

228

(1841) 329.

Rhizome tuberous, internally pale to brighter yellow, aromatic. Leafy shoots 1.25-

1.75 m. Leaf sheaths sparsely hairy. Ligula papery, 1.5—2.5 cm long, scarious. Petiole

finely hairy. Leaves broadly lanceolate, 25-40 by 5-8 cm, apex acuminate. Scape

radical, erect, 10-30 cm; sheaths green. Spike cylindric to ovate, 6-14 by 4-5 cm,

apex obtuse. Bracts obovate, 3.0-4.0 by 2.5 cm, green when young, red when old,

convex near upper edge, apex broadly rounded with a thin, papery margin. Bracteoles

linear to lanceolate, 2.5—3.5 long.Calyx 2.5 cm long, shorter than bracteole,

white.Corolla 5.5 cm long, lemon yellow; dorsal lobe 2.5 by 2.0 cm; lateral lobes

1.6 by 0.7 cm. Labellum 5.5 cm long, lemon yellow, margin crenate; midlobe oblong

almost round, 1.5 cm long, apex cleft; side-lobes ovate, 0.8 cm long. Anther pale

yellow. Capsule oblong, 1.5 cm long, red.

Type: Burma, Pegu, 1826, C.W. s.n. (K lectotype!). Lectotype here selected. No

specimen in LINN or Herb. Cliff (BM).

Collections: Burkill & Haniff 13712, 13421; Collins 1053, 1611; Corner 31559;

Curtis 1924; Zainudin 3826, 4466.

Distribution: Cultivated in India, China and throughout Southeast Asia. It is

probably indigenous to India.

Malaysia: Kedah, Perak, Selangor, Johor.

Ecology: Cultivated or naturalized on margins of forests or waste ground near

villages. Grown up to 1200 m alt, flowering June to September, fruiting October

till January.

Uses: The rhizome is used as a spice as well as for medical purposes.

Vernacular names: India and Malaysia: “Lampoyang”. In Ayurvedic medicine

Z. zerumbet is called called “stone ginger” in order to distinguish it from Z.

montanum called “camphor ginger’. Z. zerumbet has some bitterness in the taste

which Z. montanum lacks.

Notes: Valeton (1918) described three species from Java, Z. amaricans Z.

aromaticum and Z. littorale, as closely allied to Z. zerumbet. It might be possible to

distinguish Z. aromaticum by its fleshy white rhizome and stronger taste as described

by Valeton. Unfortunately both characters require live material, which was not

available for this study. In this treatment Z. amaricans Z. aromaticum and Z. littorale

are regarded as varieties of Z. zerumbet.

Affinities: Z. zerumbet is allied to Z. ottensii but the bracts are green at anthesis

and the labellum is lemon yellow without markings. Furthermore, Z. oftensii can

be distinguished by the rhizome, which has a purplish color inside.

cultivar darceyi Veitch Burtt, Notes Roy. Bot. Gard. Edinb. 31 (1972) 315-316. —

Zingiber darceyi hort. syn. Veitch, Veitch Cat. (1890) 13; Wien Ill. Gart. Zeit. (1890)

229

398; Garden (1890) 43; Kew Bull. app. 2 (1891) 54; Roy. Hort. Soc. Dict. 4 (1951)

2310.

This cultivar is similar to the typical form except for the variegated leaves which

are bright green with a broad white margin and oblique stripes of the same colour.

Futhermore, the bracts of the inflorescence have whitish stripes near the margin.

Note: This cultivar is the only known Zingiber with variegated leaves and has great

horticultural potential. Z. zerumbet cultivar darceyi was introduced to Kew in 1890

from Sydney Botanic Garden and is said to come from the South Sea Islands (Burtt

1972).

13. Zingiber griffithii Baker in Hook. f., Fl. Brit. India. 6 (1892) 246; Ridley,

Asiat. Soc. Beng. (1899) 131; Schumann in Pflanzenr. Zing. (1904) 177; Ridley,

Fl. Mal. Pen. 4 (1924) 260. Holttum, Gdns’. Bull. Singapore 8 (1950) 60-61.

Leafy shoots 0.5-1.2 m tall. Leaf sheaths pubescent, indumentum increasing

towards the petiole. Petiole 2-3 mm long, pubescent. Ligula bilobed, lobes broadly

rounded, 3 mm long, pubescent, margin scarious, dotted. Leaves broadly lanceolate

or ovate, 15 by 5 cm to 20(-25) by 8 cm, upper surface smooth, lower surface and

midrib sericeus, finely dotted throughout, base broadly to narrowly cuneate, apex

mucronate, sidenerves finely raised in dotted lines when dried giving a finely ribbed

appearance. Scape radical, more or less erect, 4-10(-15) cm. Spike 10-15 by 1.5-

3.5 cm, fusiform when young, wider and nearly evenly cylindric when in fruit,

apex acuminate. Bracts elliptic, 2.5-4.0 by 1.5-2.5 cm, thin, pink turning red, finely

hairy. Bracteoles absent. Calyx 2.5 cm long, longer than the bracteole. Corolla 5.0

cm long, white to cream; dorsal lobe 2.0 by 1.0 cm; lateral lobes joined together for

nearly half their length below the lip. Labellum white to cream; midlobe triangular,

1.7 by 0.6 cm, apex acute or sometimes cleft; lateral staminodes ovate, 0.8 by 0.4

mm, tips rounded. Capsule glabrous, flowers persisting on top of the fruit. Seeds

maroon. Pollen globose, sculpturing cerebroid.

Type: Peninsular Malaysia, Malacca. Griffith, Kew Distrib. 5731 (K lectotype).

Collections: Burkill 1170, 4485; Corporal s.n. Feb. 1890; Goodenough 1434,

s.n. (12 Feb. 1890); Holttum 9393, 10926; s.n. 10 Apr. 1890; Lake & Kelsall s.n.

22 Oct 1892. Latiff 856, 3151; Mohd. Shah et al. S.1700, 3633; Ridley s.n., s.n. (20

Aug.1892), 6011, 9188; Sinclair 5105, s.n. 25 Feb. 1950, 10616; Smith 11;

Symington 22750, 25669.

Distribution: Peninsular Thailand, Malaysia and Singapore

Malaysia: Johore, Melaka, Negri Sembilan, Pahang, Selangor, Terengganu.

Ecology: Lowland evergreen forest or secondary forest in damp, shady places

in humus rich soil. Common in lowland forests in the southern half the Malay

Peninsula.

230

Uses: The plant may be used for poulticing.

Vernacular names: “Jepus merah” (red gingerwort), “Tepus kechil’ (little

gingerwort), “Zepus huma” (uplands clearing gingerwort).

Notes: Z. griffithii is well characterized by its broad leaves with silky hairs on the

lower surface and finely raised veins when dried. Holttum regarded Z. griffithii var.

major as a synonym to Z. puberulum and his view has been followed in this

treatment.

Affinities: Z. griffithii is closely related to Z. puberulum and Z. gracile, but the

leaves are broader and the inflorescence is more cylindric than either of these.

Furthermore, the bracts in Z. griffithii are much less tough than in Z. puberulum

and Z. gracile.

14. Zingiber citrinum Ridl. (1899), J. Straits Brch. R. Asiat. Soc. 32: 129; (1907)

Mat. Fl. Malay. Penins. 2: 28-29; (1924) Fl. Malay. Penins. 4: 260; Schumann in

Pflanzenr. Zing. (1904) 174. — Zingiber griffithii var. citrinum Holttum, Gdns’

Bull. Singapore 13 (1950) 61.

Leafy stem 0.5—0.6 m, flushed with purple. Leaf sheath usually glabrous, margin

scarious. Ligula entire, 24 mm long, glabrous. Petiole 2 mm, sparsely hairy. Leaves

lanceolate to ovate, 24-30 by 8-10 cm, sparsely sericeus, base cuneate, apex

acuminate. Scape radical, erect, 7.0 cm long. Spike cylindric, 12-15 by 4-5 cm,

apex acute. Bracts densely imbricate, elliptic to ovate, 3.5-4.0 by 2—3 cm, lemon

yellow turning pink to red in fruit, glabrous or sparsely pilose, margin scarious,

apex acute. Bracteoles absent. Calyx 2.5 cm long, translucent—yellow. Corolla 5.6

cm long, white to yellow; dorsal lobe 2.4 by 0.8 cm. Labellum 5.4 cm, white to

yellow; midlobe elliptic, 1.0 by 0.6 cm, apex bifid; lateral staminodes 0.5 by 0.4

cm.

Type: Peninsular Malaysia, Genting Bidai, May 1896, Ridley 7797a (K

lectotype!). Lectotype here selected.

Collections: Burkill & Haniff 16444; Foxworthy & Burkill 1921; Holttum 9621;

Dr Kings coll. 10263; Lake & Kelsall 1892; Ridley s.n. (1900), s.n. 1899.

Malaysia: Johor, Negri Sembilan, Perak, Selangor,.

Ecology: On rich soil, 120-180 m, flowering in May-June.

Vernacular name: “TJepus tenok” meaning tapir’s gingerwort (Ridley 1924).

Notes: Easily recognized by the lemon yellow bracts. Ridley’s drawing of the plant

brought from Dusun Tua is found at Kew. Holttum (1950) considered it doubtful

whether the large leaves were always associated with the lemon colour of the bracts

and included Z. citrinum in Z. griffithii.

Affinities: Z. citrinum is has affinity to Z. griffithii. It is distinguished by the

larger leaves, the broader inflorescence and the lemon yellow bracts.

Zoi

15. Zingiber sulphureum Burkill ex Theilade, Curtis’s Bot. Mag. 12 (1995) 7377.

Leafy shoots to 0.7 m tall. Petiole 3-4 mm, yellowish pubescent. Ligula 4-6

mm, biparted, hairy when young. Leaves ovate, 12-14 by 4.0-4.5 cm, glabrous

above, hairy below. Scape 5-10(-15) cm long. Spike fusiform, slender, 8-10 by 1.5

cm, apex acuminate. Bracts ovate, 3.5 by 1.5 cm, sulphur yellow later turning

pink, pubescent, apex acute, finely dotted along the margin when dried. Bracteole

absent. Calyx 2.3 cm long, pale yellow. Corolla 5.5 cm long, pale yellow; dorsal

corolla lobe 2.3 by 0.4 cm; lateral corolla lobes 1.8 by 0.3 cm Labellum 5.0 cm

long, pale yellow; midlobe oblong, 1.3 by 0.4 cm, retuse; side-lobes 0.5 by 0.4 cm,

oblong, total width when flattened 1.5 cm. Anther pale yellow. Pollen globose with

cerebroid sculpturing.

Type: Peninsular Malaysia, Pahang, Gunung Tahan, 1150 m, 18 June 1922,

Haniff & Nur SNF 8016 (K holotype!, SING isotype!).

Collections: D. Cooke 23. Evans s.n. (June 1917). Willis R. Littke 421.

Distribution: Malaysia in Pahang state. Localities: Taman Negara, Gunung

Senyum, Fraser’s Hill.

Ecology: Evergreen rain forest, on granite or limestone, 50-1200 m. Flowering

June to August.

Notes: Zingiber sulphureum is closely related to Zingiber griffithii and Zingiber

aurantiacum but it is a smaller plant than either of these. Zingiber sulphureum is

recognized by the small ovate leaves, the short, narrow spike with sulphur yellow

bracts and the short calyx. Burkill used the epithet su/phureum on herbarium

specimens SNF 8016.

16. Zingiber gracile Jack, Malayan Misc. | (1820) 1; Baker in Hook. f,, Fl. Brit.

India. 6 (1892) 246; Ridley, J. Straits Brch. R. Asiat. Soc. 32 (1899) 130; Fl. Mal.

Pen. 4 (1924) 260. Holttum, Gdns’ Bull. Singapore 13 (1950) 63-65.

Leafy shoots to 1 m tall. Ligula about 1.8 cm, bilobed, scarious, with black dots.

Leaves lanceolate, to 18 by 4.0 cm, upper surface smooth, lower surface and midrib

pubescent or sericeus, base cuneate, apex acuminate. Scape to 10-20 cm.

Inflorescence cylindrical about 15 by 1.5 cm. Bracts ovate, 3.5-5.0 by 2.0 cm, pink

or orange when young later turning red, pubescent, apex acute. Bracteole 2.0-2.5

cm shorter than calyx. Calyx 2.5-3.0 cm long. Corolla 6.0 cm; lobes 1.5-2.0 cm

long, cream. Labellum to 6.0 cm, cream; midlobe oblong, 2.2 cm long, apex retuse

or bifid; side-lobes 0.2 cm long, oval. Capsule 2.5 cm long, glabrous; seeds maroon.

Pollen globose, sculpturing cerebroid.

Type: Peninsular Malaysia, Penang, Waterfall, Apr. 1890, Curtis (SING

lectotype!). Lectotype here selected. No authentic specimens of Jack’s plants exist.

Collections: Burkill 16141; Fox 12708; Foxworthy & Burkill Nov. 1921; Mohd.

Shah 2460; Ridley 1640, 7235, 12700.

232

Malaysia: Johore, Malacca, Pahang, Penang, Perak, Selangor.

Uses: The use of the name “mempoyang” for “lempoyang” suggests that it may

be a substitute for other gingers. The name was obtained by Ridley and Goodenough

in Malacca (Burkill 1966).

Note: This is the species that corresponds to Jack’s original description. Holttum

recognized in addition to the typical form three varieties of Z. gracile. In this

treatment they have been ranked as separate species. Z. gracile is distinct by the

long thin ligule; the lanceolate leaves and the short scape and spike. It is probably

closely related to Z. griffithii.

King’s 7954 and Hullett’s 854, cited by Baker (1892) as probably Z. gracile,

have been identified as Z. elatior and Z. aurantiacum.

17. Zingiber aurantiacum Theilade stat. nov. — Zingiber gracile Jack var.

aurantiacum Holttum, Gdns’. Bull. Singapore. 13 (1950) 63.

Leafy shoots 1.5 to 2 m tall. Ligula 5 mm long. Leaves narrowly lanceolate, to

_ 27 by 3.5 cm. Scape 15 to 35 cm. Spike slender, 15-20 cm long and 2.5 cm wide.

Bracts bright orange turning red when fruiting. Bracteole 2.5 cm long. Calyx 2.5-

3.0 cm long. Corolla 6.0 cm; lobes 2.0 cm long, cream. Labellum to 6.0 cm, cream;

midlobe oblong, 2.2 cm long, apex retuse or bifid; side-lobes oval. Capsule to 2.5

cm long, glabrous; seeds maroon. Pollen globose, sculpturing cerebroid.

Type: West Malaysia, Pahang, Fraser’s Hill, Burkill & Holttum SEN. 8806 (SING

lectotype!). Lectotype here selected.

Collections: Burkill 2337, 3162, 5988; Burkill & Holttum 8633; Corner Nov.

1935, Aug. 1937, 33191; Henderson 11416; Hullett Apr. 1888; Kelsall 1978; Mohd.

Shah 2765; Ridley Jun. 1889, Dec. 1920; Wray 5365.

Malaysia: Johore, Malacca, Negri Sembilan, Pahang, Selangor.

Ecology: Common at moderate elevations on the southern part of the Main

Range in Cameron Highlands (Holttum, 1950).

Note: This species is distinguished by the long, slender inflorescence, and the orange

bracts later turning red. It has affinity to Z. gracile but differs in being a larger plant

with a short ligule. The long and slender inflorescence approaches Z. petiolatum

but the leaves are much smaller.

18. Zingiber petiolatum Theilade stat. nov. — Zingiber gracile Jack var. petiolatum

Holttum, Gdns’. Bull. Singapore. 13 (1950) 63.

Leafy shoots to 2.5 m tall. Petiole 10-15 mm. Ligula bilobed, 3-4 mm. Leaves

lanceolate, to 40 by 8 cm, lower surface glabrous. Scape to 75 cm long. Spike 30-

45 cm long, fusiform. Bracts lanceolate, 7.0 by 3.0 cm, tough, rose-pink, apex

acute. Bracteole to 3.0 cm long. Corolla 6.0 cm; lobes 2.0 cm long, cream. Labellum

233

to 6.0 cm, cream; midlobe oblong, 2.5 cm long, apex retuse or bifid; side-lobes 0.3

cm long, oval.

Type: Peninsular Malaysia, Kedah, Pass from Kroh to Baling, 350 m. Corner

S.F.N. 31570 (SING holotype!, E!, K! and L isotypes).

Collections: Henderson 21794; Kerr 7353; Md. Nur 34284; Ridley s.n. Aug.

1891, s.n. 18 Mar. 1893.

Distribution: Peninsular Thailand and Peninsular Malaysia.

Malaysia: Kedah, Pahang.

Ecology: Evergreen forest at low elevations.

Note: Z. petiolatum is distinct by the large leaves, the long petiole, the extremely

long scape and the long, slender spike with large tough bracts. In the large leaves

and tough bracts Z. gracile var. petiolatum approaches Z. puberulum.

19. Zingiber puberulum Ridley, J. Straits Brch. R. Asiat. Soc. 32 (1899) 131; Fl.

Mal. Pen. 4 (1924) 261; Holttum, Gdns’ Bull. Singapore. 13 (1950) 61-62. — Z.

griffithii var. major Ridley, J. Straits Brch. R. Asiat. Soc. 32 (1899) 132.

Key to varieties:

Ren PEEL OLES GENCE OVEIG APE R: ROW GEM rey .c ees nacsctacaseasnshsssersodcosscecescuaseuscsevounescovevseaveseoseaascessescceles var. ovoideum

lb. Inflorescence fusiform to ovoid tapering to a pointed aPeX .............sceceeeececeeseceseeeeseeeeeesceceececeecuceeeeeseeeneeees 2

2a. Leaf sheaths velutinous with brownish hairs, bracts pink ..............:.:c:escesseeseseseeeeeeneeeeeeeesees var. puberulum

PP ledies heats) Su Pty hairy, IDIACtS AV ELLOW .<---ccecx-secen-acsurueaceeree<ovecvoverssentiveseieonsutdeeecessesiceacetenes var. chryseum

var. puberulum

Leafy shoots 2-3 m tall. Leaf sheaths brownish velutinus. Ligula entire, to 5 mm

long, yellowish hairy or velutinus. Leaves lanceolate or widest above the middle,

25-55 by 6-10 cm, upper surface smooth, rather grey-green, lower surface sometimes

pubescent throughout, densely so on midrib towards the base, hairs yellowish-

brown, margin hairy, apex acuminate. Scape radical, erect, 10-25 cm long. Spike

10-15 by 2.5-3.5 cm, fusiform to ovoid tapering upwards to a pointed apex. Bracts

ovate, 3.5-5.0 by 2.0-3.0 cm, texture firm, pink, pubescent, apex obtuse, margin

scarious, usually conspiciously hairy. Bracteole ensiforme, 1.5-2.0 by 2.4-2.8 cm.

Calyx 2.5-3.3 cm long. Corolla to 6.5 cm, white to cream; dorsal lobe 2.5 by 0.7

cm wide; lateral lobes 2.0 by 0.4 cm. Labellum nearly as long as the corolla lobes,

cream; midlobe about 1.2 by 0.7 mm, apex obtuse with a crenate margin; side-

lobes oblong, reaching nearly half the total length of lip, spreading to a total width

of 1.6 cm when flattened. Anther yellow. Capsule 2.5 cm long, pubescent. Pollen

globose, sculpturing cerebroid.

Type: Singapore, Bukit Timah, Ridley s.n. 1894, (K lectotype!, SING isotype).

Lectotype here selected.

234

Collections: Burkill 2670, 5990; Corner 30112, 30266; 30969, 31477; Curtis

3037; Dr Kings Coll 2163; Henderson 22380, 25003, 36608; Holttum 10842;

Maxwell 81-96, 81-133; Md. Nur sn. 1937; Ridley s.n. Bukit Timah 1892, s.n.

Bukit Tangga 1920, 4613; Sinclair 10616, 10622; Stone 10877.

Distribution: Peninsular Thailand, Peninsular Malaysia, and Singapore.

Malaysia: Penang, Perak, Terengganu, Pahang, Selangor, Johore.

Ecology: Evergreen forest in damp places, peaty soil in fresh-water swamp forest,

up to 600 m.

Vernacular name: “Lempoyang anjing” or dog’s “lempoyang” (Burkill 1966).

Notes: The indumentum of var. puberulum is very variable, but it is always hairy

on the ligule and leaf sheaths. According to Ridley (1899) Z. griffithii var. major

differs from var. puberulum in the glabrous leaf sheaths. In Ridley’s type specimen

of Z. griffithii var. major (s.n. July 1891, Kuala Sembiling, Pahang) the leaf sheaths

and petioles was found to be hairy though the indumentum was missing in some

parts of the leaf sheaths. Holttum (1950) considered Z. griffithii var. major Ridl. to

be a synonym to Z. puberulum and his view is followed here.

Affinities: var. puberulum is closely allied to Z. griffithii and Z. petiolatum. but

has larger leaves than the former, smaller inflorescence than the latter, and is easily

recognized by the velutinus leaf sheaths and ligules.

var. ovoideum (Ridl.) Holttum, Gdns’ Bull. Singapore 8 (1950) 63.

This variety is rarely, if ever, as hairy as the typical form, but otherwise it is

indistinguishable vegetatively. It differs from the typical form in the following:

Scape 5-10 cm long. Spike ovoid, about 8 cm long, apex rounded.

Type: Peninsular Malaysia, Pahang, river Tahan, Ridley s.n. Aug. 1891 (SING

lectotype!). Lectotype here selected.

Henderson S.F.N. 21781, 21857, 22554, 19566; Moysey & Kiah 33732; Ridley

Sn, 1892.

Malaysia: Kelantan, Perak, Pahang.

Notes: This variety has been collected mainly in Pahang. The less hairy character,

the short scape and the short, ovoid inflorescence with rounded apex seems to be

constant.

var. chryseum (Ridl.) Holttum, Gdns’ Bull. Singapore 8 (1950) 62. —Zingiber

chryseum Ridley, J. Straits Brch. Asiat. Soc. 50 (1908) 149. Fl. Mal. Pen. 4 (1924)

260.

Differs from the typical form in having pale yellowish bracts and in the whole

plant being almost glabrous.

235

Type: Singapore, Stagmount, Ridley 13330, 1908 (SING holotype!, K isotype!)

Collections: Corner 30658 Sungai Panti, Pahang.

Distribution: Pahang and Singapore. The type specimen was collected in a wood

at Stagmount in 1908 but the habitat was cleared and burnt a year or so later (Ridley,

1924).

Ecology: lowland forest, up to 200 m.

Notes: In size and shape of the leaves and inflorescence it does not differ in any

way from var. puberulum, and the flowers are described in same terms by Ridley.

Var. chryseum differs in the pale yellowish bracts and in the whole plant being

almost glabrous. Whether the yellow bracts and general glabrous character is always

associated is not known.

References

Burkill, I. H. (1966). A dictionary of the economic products of the Malay Peninsula.

Art Printing Works, Kuala Lumpur: 2335-2346.

Burtt, B.L. (1972). General Introduction to papers on Zingiberaceae. Notes Roy.

Bot. Gard. Edinb. 31: 155-228.

Cowley, J. & Theilade, I. (1995). Zingiber sulphureum (Zingiberaceae). Curtis's

Bot. Mag. 12: 73-77.

Dahlgren, R.M.T., Clifford, H.T. & Yeo, P.F. (1985). The families of the

Monocotyledons. Springer Verlag, Berlin: 350-352, 360-367.

Flenley, J. (1979). The equatorial rain forest: a geological history. Butterworths,

London: 77-98.

Holttum, R.E. (1950). Zingiberaceae. Gdn’s. Bull. Singapore 13: 38-65.

Jones, C.E. (1983). Handbook of experimental pollination biology, eds. C.E. Jones

& R.J. Little. Scientific & Academic editions, New York: 3-30.

Larsen, K. (1980). Annotated key to the genera of Zingiberaceae of Thailand. Nat.

Hist. Bull. Siam Soc. 28: 151-169.

Ridley, H.N. (1899). Zingiber elatior. J. Straits Brch. R. Asiat. Soc. 32: 130.

Ridley, H.N. (1912). Spices. MacMillan, London: 390.

Ridley, H.N. (1924). Zingiber. The Flora of the Malay Peninsula 4: 257-261. Reeve

& Co., London.

Rumphius, G.E. (1747). Herb. Amboin. 5. Utrecht. Not seen.

Schumann, K. (1904). Zingiber. Das Pflanzenreich. Wilhelm Engelmann, Leipzig:

163187.

Valeton, T. (1918). New notes on the Zingiberaceae of Java and Malaya. Bull.

Jard. Bot. Buitenz. 27: 1-176.

Whitmore, T.C. (1984). Tropical rain forests of the Far East, ed. 2; Oxford University

Press, New York: 12-14, 238-239.

Wu, T.-L. (1985). The origin of Zingiber officinale. Agricultural Archeology 2:

247-250.

236

Acknowledgements

Professor Kai Larsen, Department of Systematic Botany, University of Aarhus,

is gratefully acknowledged for valuable discussions and advice during the work. I

am indebted to the curators of BM, E, K, KEP, L, P, SING, UKMS, UM for placing

their material at my disposal. I also like to thank Jorgen Theilade and Anthea Phillips

for usefull comments on the script. Benjamin @llgaard kindly latinized the diagnoses

and Bgrge Madsen made the drawings for Fig. 1 & 2.

The drawings of Zingiber curtisii (Fig. 4) and Z. chrysostachys (Fig. 5) are in the

collection of the Singapore Botanic Gardens and are used here with permission.

This study was financed by a scholarship from Danida. Sibbald Trust Fund supported

my stay at The Royal Botanic Gardens Edinburgh.

The Angiosperm Flora of Singapore Part 4

SCHISANDRACEAE

R.M.K. SAUNDERS

Department of Ecology & Biodiversity,

The University of Hong Kong,

Pokfulam Road,

Hong Kong

Kadsura Juss.

Ann. Mus. Hist. nat. 16 (1810) 340; Ridl., Fl. Malay Penins. 1 (1922) 20;

A.C.Sm., Sargentia 7 (1947) 156; Backer & Bakh.f., Fl. Java 1(1963) 99; R.M.K.

Saunders, Fl. Males. 1:13 (1997) 190; Syst. Bot. Monogr. (in press).

Sarcocarpon Blume

Woody, monoecious lianes. Leaves simple, alternate; exstipulate; lamina elliptic

to ovate, papyraceous to coriaceous, apex acute or acuminate, margins denticulate

to entire, base cuneate (especially when young), obtuse or truncate; petioles grooved

adaxially. Flowers unisexual; in axils of leaves or fugaceous bracts, generally

solitary, occasionally with a secondary flower growing in the axil of the prophyll,

or in clusters of 2-4 forming glomerules, occasionally cauliflorous; tepals 7-24,

imbricate at anthesis, suborbicular, elliptic or ovate, rarely obovate, outermost and

innermost tepals + reduced, inner and middle tepals white, cream, yellow, pink or

red, outer tepals often green; stamens 13-80, either essentially free but connate at

the base of the filaments (occasionally with subulate appendages at the distal apex

of the receptacle), or else stamens aggregated into a compact subglobose head with

very broad connectives; pollen hexacolpate, distally syncolpate; carpels 17-300,

free; ovaries with 2-5(-11) pendulous or ventrally attached ovules. Fruit a subglobose

aggregate of berries attached to an ellipsoid or clavate receptacle; berries ripening

red or yellow. Seeds 1-4(-11) per berry, smooth, hilum lateral or apical.

Distribution - There are 16 species in Kadsura, with a southern Chinese and

Indo-Chinese centre of distribution, extending from southern Japan in the north-

east, to Sulawesi and Java in the south-east, and Sri Lanka in the west (Smith,

1947; Saunders, in press). Only K. scandens is found in Singapore.

Ecology - Scrambling and twining woody vines of warm and subtropical broad-

leaved forests, with some species extending into humid montane forests of up to

2400 m altitude.

Uses - See under K. scandens.

Notes - The most recent monograph of Kadsura is by Saunders (in press). A

detailed review of the palynology of the family has been published by Praglowski

(1976).

>» > ° fi ; 4 iy

% I et ae ee ; Cs Fay, 5 ips

ae i ee 5 i . is

a ae tty ar ht

7 ae aee 4 ri

* i

ney st Sf

‘ae

239

Fig. 1. Kadsura scandens (Blume) Blume. a. Flowering branch. b. Female flower

with perianth removed, showing gynoecium. c. Isolated carpel (lateral

view). d. Male flower with perianth removed, showing androecium. e.

Isolated stamen (lateral view). f. Fruit. g. Seed. [a. H.N. Ridley 6354

(SING); b.-c. M. Nur 26103 (SING); d.-e. P.W. Korthals s.n. (L); f. redrawn

from van Steenis (1972: fig. 29.3); g. N. Wirawan 134 (L)]. Del. H.L.

Wilks. (Reproduced with permission from Flora Malesiana 13).

240

Acknowledgements

I would like to thank the Directors of L and SING for the loan of herbarium

specimens, Mr Ali bin Ibrahim for the current location of Kadsura scandens

in Singapore, and Hazel Wilks for drawing Fig. 1.

References

Burkill, I.H. (1966). A Dictionary of the Economic Products of the Malay

Peninsula. Vol. 2. Ministry of Agriculture and Co-operatives; Kuala Lumpur;

1296.

Perry, L.M. (1980). Medicinal Plants of East and Southeast Asia: Attributed

Properties and Uses. MIT Press; Cambridge, Massachusetts; 382.

Praglowski, J. (1976). Schisandraceae Bl. World Pollen and Spore Flora 5, \-32.

Rao, H.S. (1939). Cuticular studies of Magnoliales. Proc. Indian Acad. Sci. B:9,

99-116, pl. 4-21.

Saunders, R.M.K. (In press). A monograph of Kadsura (Schisandraceae). Syst. Bot.

Monogr.

Smith, A.C. (1947). The families Illiciaceae and Schisandraceae. Sargentia 7, 1-

224.

Steenis, C.G.G.J. van (1972). The Mountain Flora of Java. Brill; Leiden.

Wodehouse, R.P. (1959). Pollen Grains: Their Identification and Significance

in Science and Medicine. Hafner; New York; 340.

The Angiosperm Flora of Singapore Part 5

BURMANNIACEAE

R.M.K. SAUNDERS

Department of Ecology & Biodiversity,

The University of Hong Kong,

Pokfulam Road,

Hong Kong

Major references: Jonker, Meded. bot. Mus. Rijks-Univ. Utrecht 51 (1938) 1-

279; Fl. Males. 1:4 (1948) 13-26; R. Dahlgren et al., Fam. Monocot. (1985) 216-

219; Maas et al., Fl. Neotropica, Monogr. 42, Burmanniaceae (1986) 1-189; T.

Riibsamen, Diss. Bot. 92 (1986) 1-310, pl. 1-98.

Small saprophytic or autotrophic, annual or perennial herbs, often growing from

rhizomes or tubers. Leaves simple, exstipulate, usually alternate, lamina entire;

autotrophic spp. with basal rosette of linear or lanceolate green leaves; saprophytic

spp. with colourless to yellowish or reddish, scale-like leaves. Flowers bisexual,

usually actinomorphic, solitary or in terminal cymes or racemes; perianth (3-)6-

lobed, corolline, tubular or campanulate, tube occasionally 3-angled or 3-winged,

outer parts (sepals) valvate, inner parts (petals) generally smaller, induplicate-

valvate, occasionally absent, 3 or all 6 of the lobes often each with an elongate,

slender, terminal appendage; stamens 3 or 6, if 3, then subsessile in the perianth

throat, with anther dehiscence by horizontal slits if 6, then pendant in perianth

tube, with anther dehiscence by longitudinal slits; connectives large, often

appendiculate; ovary inferior, 1-loculate with parietal placentation, or 3-loculate

with axile placentation, ovules numerous, anatropous, bitegmic; style filiform,

shortly cylindrical or conical; stigmas 3, sometimes connate. Fruit capsular,

occasionally fleshy, with either persistent perianth tube and style or persistent basal

ring of perianth only, dehiscence irregular or by transverse ventral slits. Seeds small,

subglobose to linear, numerous, sometimes with a loose reticulate testa; endosperm

present. x = 6 or 8 (Riibsamen, 1986).

Distribution - C. 20 genera and 130 spp., with an essentially pantropical

distribution but also extending into temperate regions (Jonker, 1938).

Ecology - The Burmanniaceae consist of both saprophytes and autotrophs (semi-

saprophytic). The autotrophic spp. generally occur as sparse individuals in open

fields and savannahs, whereas the saprophytes occur almost exclusively on decaying

leaves, wood and roots in the deep shade of wet tropical forests (Jonker, 1938). The

seeds are tiny, with little food reserve, but are produced in very large quantities.

Spp. in open grassland habitats therefore presumably disperse their seeds by

wind,whereas water may be the primary means of dispersal in spp. of forested

areas (Maas et al., 1986).

Uses - The Burmanniaceae are of no known economic importance.

242

Notes - The phylogenetic relationships of the family are rather unclear, due in

part to convergence in evolution of achlorophyllous plants. The Burmanniaceae

are clearly isolated, but possess greatest similarities with the Orchidaceae, although

not with the subfamily Apostasioideae (Riibsamen, 1986), as previously suggested.

The Burmanniaceae have been classified into 3 tribes (Burmannieae, Thismieae

and Corsieae) based on the inflorescence type, and stamen number and attachment,

although the Corsieae are generally elevated to the segregate family Corsiaceae

(e.g., Jonker, 1938; Riibsamen, 1986). The Thismieae have also been isolated as a

distinct family (e.g., Dahlgren et al., 1985), although the present account follows

most other recent treatments in describing the family Burmanniaceae sensu lato.

Key to the Genera

la. Perianth tube cylindrical or trigonous. Style of + equal length as the tube.

StAMIONS) 3 id sascsusat telsheed. Renews cae bashe rere caeslenms once ek ee eee 2

lb. Perianth tube urceolate, circumscissile. Style very short.

StdMENS 6c 58onsacaet che eateds otaca ee inet! Ee eee ee ee Thismia

2a. Perianth persistent after anthesis, together with ovary often prominently 3-

winged, sometimes 3-costate or wingless. Ovary and capsule 3-loculate with

axile placemtato ir. 364.1 ci0.2 docsde seb! Gobaceusctec tere eee ee eee Burmannia

2b. Perianth caducous after anthesis, together with ovary wingless. Ovary and

capsule 1-loculate with parietal placentation. .................00 Gymnosiphon

Burmannia L.

Sp. pl., ed. 1 (1753) 287; Ridl., Mat. fl. Malay. Penins. 2 (1907) 69; Fl. Malay

Penins. 4 (1924) 303; Jonker, Meded. bot. Mus. Rijks-Univ. Utrecht 51 (1938) 18,

57; Fl. Males. 1:4 (1948) 15; M.R. Hend., Malay. Wild Flowers, Monocot. (1954)

170; Backer & Bahk. f., Fl. Java 3 (1968) 213.

Achlorophyllous saprophytic or green autotrophic, annual or perennial herbs;

autotrophs with basal rosette of green leaves, and smaller green leaves appressed

to the stem; saprophytes with only small bract-like, achlorophyllous, stem leaves.

Flowers solitary or in terminal cymes; perianth persistent after anthesis, often

3-winged, occasionally wingless, tube generally 6-lobed, cylindrical to trigonous,

outer segments relatively large, inner segments often minute, occasionally lacking;

stamens 3, sessile or subsessile on perianth tube, connective occasionally with

(1-) 2 apical glandular crests and/or a central, pendant, basal spur; ovary trigonous,

3-loculate, placentation axile; styles fused, branched towards the stigmas; stigmas

3, occasionally sessile, funnel-shaped. Capsule with persistent perianth, dehiscence

generally irregular. Seeds numerous, oblong or ellipsoidal.

Distribution - C. 60 spp., throughout the range of the whole family (Jonker,

1938) although with an Asian centre of diversity. Only B. championii, B. coelestis

and B. wallichii have been recorded in Singapore.

243

Ecology - As with the family, most spp. of Burmannia are either autotrophs of

wet areas of open savannahs, or saprophytes of wet tropical forests (Jonker, 1938),

although | neotropical sp. is known to be epiphytic (Maas et al., 1986). Although

the brightly coloured perianth, septal nectaries and glandular staminal crests are

indicative of entomophily, no such system has ever been observed. Protandry has

been demonstrated for some Burmannia spp. (Malme, 1896), and it has been

suggested that the lower lip of the stigmatic lobes may be a means of avoiding

autogamy. The stigmas and anthers are, nonetheless, located at the same height in

the perianth tube, often almost occluding the throat of the tube, and Schoh (1920)

believed that Burmannia must be autogamous since the stigmatic lobes prevent the

dispersal of pollen from the thecae.

Notes - The distinction between autotrophic and saprophytic habits was

previously believed to be of some taxonomic significance, but is now considered

of little value due to the occurrence of pairs of closely related saprophytic and

autotrophic spp. (Jonker, 1938), suggesting that saprophytism is polyphyletic in

the genus. The pollen is mono- or biporate (Chakrapani & Raj, 1971).

Key to the Species

Remiee aver Ml a DASALTOSEIC, PTCOM «....22..-.2ic ene ecensekeruscieseavessscesecatene B. coelestis

ih Leaves not ina basal rosette, achlorophyllous ............0......c..sseceteesceosebneeseee f

2a. Inflorescences (2-)3-9(-14)-flowered. Perianth tube without lateral

Mareen ane Pere NE yes Eh ein 5 Siig avn deteh Ske stb kes B. championii

2b. Flowers generally solitary. Perianth tube with 3 lateral wings.... B. wallichii

1. B.championii Thwaites

Enum. pl. Zeyl. 5 (1864) 325; Ridl., Mat. fl. Malay. Penins. 2 (1907) 72; A.

Ernst & C. Bernard, Annls Jard. bot. Buitenz. 24 (1911) 81-82; Ridl., Fl. Malay

Penins. 4 (1924) 306; Jonker, Meded. bot. Mus. Rijks-Univ. Utrecht 51 (1938)

138; Fl. Males., 1:4 (1948) 17; M.R. Hend., Malay. Wild Flowers Monocot. (1954)

172; Backer & Bakh. f., Fl. Java 3 (1968) 213; H. Keng, Gdns’ Bull., Singapore 40

(1987) 123; 1.M. Turner, K.S. Chua & H.T.W. Tan, J. Singapore natn. Acad. Sci. 18

& 19 (1990) 63; I.M. Turner, Gdns’ Bull., Singapore 45 (1993) 47.

B. capitata Makino; B. chionantha Schltr.; B. dalzieli Rendle; B. japonica

Maxim. ex Makino; B. tuberosa Becc.

Saprophytic annual herbs, flowering at 9-27 cm tall; stem unbranched, filiform,

growing from a tuberous rhizome, covered with hair-like roots, giving rise to small

adventitious tubers. Stem /eaves appressed, achlorophyllous, bract-like, linear to

subulate, 2-8 mm long. Inflorescence of (2-)3-9(-14) flowers: bracts lanceolate,

2-6 mm long, apex acute. Flowers 3-11 mm long; perianth yellow-white to bright

yellow, tube 3-costate, without lateral wings, outer lobes triangular, c. 1-2.5 mm

244

long, apex acute involute, inner lobes spathulate, to c. 0.75 mm long, apex rounded;

staminal connective with single, central crest at apex, basal spur absent; gynoecium

c. 3 mm long; ovary c. 2-3 mm long; flowers pedicellate or subsessile. Capsule

dehiscing by fragmentation of the wall between costae. n = 6, 2n = 12 (Ernst and

Bernard, 1912).

Distribution - Japan, southern China, Sri Lanka and Malesia (Malay Peninsula,

Bangka, Java, Borneo and Papua New Guinea) (Jonker, 1938).

Ecology - Grows in wet lowland forests.

Notes - The pollen is monoporate (Chakrapani and Raj, 1971).

2. B. coelestisD.Don

Prodr. fl. Nepal. (1825) 44; Ridl., J. Straits Brch R. Asiat. Soc. 33 (1900) 152;

Mat. fl. Malay. Penins. 2 (1907) 71; Fl. Malay Penins. 4 (1924) 304; Jonker, Meded.

bot. Mus. Rijks-Univ. Utrecht 51 (1938) 120; Fl. Males., 1:4 (1948) 17; M.R. Hend.,

Malay. Wild Flowers Monocot. (1954) 171; Backer & Bakh. f., Fl. Java 3 (1968)

_ 213; H. Keng, Gdns’ Bull., Singapore 40 (1987) 123; I.M. Turner, K.S. Chua &

H.T.W. Tan, J. Singapore natn. Acad. Sci. 18 & 19 (1990) 63; I.M. Turner, Gdns’

Bull., Singapore 45 (1993) 47.

B. azurea Griff; B. borneensis Gand.; B. javanica Blume; B. malaccensis Gand;

B. selebica Becc.; B. triflora Roxb.

Autotrophic annual herbs, flowering at 7-40 cm tall; stem usually unbranched,

robust. Leaves with basal rosette and stem forms; basal rosette leaves linear or

lanceolate, 3-26 x 0.5-3.5 mm; stem leaves appressed, sometimes imbricate towards

the base, linear to subulate, 3-30 mm long, apex acute. Inflorescence a solitary flower

or cluster of 2-4(-6) flowers; bracts linear to subulate, 1-9 mm long, apex acute.

Flowers 6-17 mm long; perianth blue, purple or white, often with yellow lobes, tube

cylindrical-trigonous with lateral wings, c. 5 mm long, lateral wings half-elliptical

to half-obovate, c. 10 x 2.5 mm, outer lobes ovate, to c. 1.5 mm long, double

margined, apex apiculate, inner lobes lanceolate, to c. 0.5 mm long, double margined,

apex apiculate; staminal connectives with 2 apical crests and a basal pendant spur;

gynoecium c. 4mm long; ovary c. 5 mm long; flowers subsessile. Capsule dehiscing

+ irregularly, by transverse splits. n = c. 16 (Sarkar et al., 1973). - Fig. 1.

Distribution - Singapore: currently in MacRitchie Reservoir, Poyan Reservoir,

Kent Ridge Park, etc. India, Nepal, southern China, Burma, Thailand, Indochina,

and throughout Malesia (Jonker, 1938).

Ecology - Grows in wet areas of open grassland.

Notes - The pollen is monoporate (Chakrapani and Raj, 1971). Larsen (1963)

reported a chromosome count of 2n = c.32 for ‘B. coelestis’; examination of the

voucher specimen resulted in its redetermination as B. chinensis Gand.

245

vem 0.5mm

Fig. 1. Burmannia coelestis D. Don. a. Habit. b. Complete flower. c. Half flower.

d. Stamen, with lateral thecae and connective consisting of 2 apical crests

and a pendant basal spur. e. 3 stigmas with united style. f. Ovary (TS),

showing axile placentation. g. Seeds, with spirally striate testa. [a.-f. K.S.

Chua & H.T.W. Tan 281 (Herbarium, The University of Hong Kong); g. Z.

Teruya 2210 (SING)]. Del. R.J. Nicholls.

246

3. B. wallichii (Miers) Hook. f.

Fl. Brit. India 5 (1888) 666; Ridl., Mat. fl. Malay. Penins. 2 (1907) 71; FI.

Malay Penins. 4 (1924) 305; Jonker, Meded. bot. Mus. Rijks-Univ. Utrecht 51

(1938) 145; R.M.K. Saunders, Blumea 41 (1996) 336.

Saprophytic annual herbs, flowering at 4.5-11 cm tall; stem unbranched, filiform.

Stem /eaves + appressed, achlorophyllous, bract-like, subulate-triangular, sometimes

keeled, 1-2.8 mm long, apex acute. Inflorescence a solitary flower or occasionally

a pair; bracts lanceolate, c. 2.5-4 mm long. Flowers 6-9.5 mm long; perianth white

or bluish; tube cylindrical, c. 3 mm long, lateral wings narrow, + linear, c. 4.5 x 0.5

mm; outer lobes obtuse-triangular, to c. | mm long, inner lobes orbiculate, apex

rounded; staminal connectives without apical crests, but with a short basal spur;

gynoecium c. 3 mm long, ovary c. 2.5 mm long; flowers subsessile. Capsule

dehiscing + irregularly by transverse splits. n = 16 (Larsen, 1963).

Distribution - India, Burma, Thailand, Indo-China, southern China and the Malay

Peninsula (Jonker, 1938).

- Ecology - B. wallichii is clearly saprophytic; otherwise little is known of its

ecology.

Notes - B. wallichii has only been recorded from Singapore once, last century

(Ridley’s collector s.n., Kranji [SING 074436]); as it has not been collected since,

it is now probably extinct in Singapore.

Gymnosiphon Blume

Enum. pl. Javae 1 (1827) 29; Ridl., Mat. fl. Malay. Penins. 2 (1907) 73; FI.

Malay Penins. 4 (1924) 306; Jonker, Meded. bot. Mus. Rijks-Univ. Utrecht 51

(1938) 27, 168; Fl. Males., 1:4 (1948) 20; Backer & Bakh. f., Fl. Java 3 (1968) 214.

Saprophytic annual herbs. Leaves achlorophyllous, bract-like, small. Flowers

borne in 3 to many-flowered racemes, rarely solitary; perianth tube 6-lobed, without

lateral wings, tube from below point of insertion of stamens deciduous after anthesis,

lower part persistent, outer lobes larger than the inner ones, slightly 3-lobed; anthers

3, sessile in the tube throat, dehiscing horizontally, connectives rather broad,

inappendiculate or mucronulate at the top; ovary ovoid to globose, with 3 parietal

placentas, placentas each with a large globose gland at both sides of the apex; style

filiform, branching; stigmas 3, often appendiculate. Capsule with persistent perianth

tube base, dehiscence generally irregular or by longitudinal slits. Seeds ovoid to

globose, with a reticulate testa.

Distribution - C. 25 spp., occurring in the neotropics, tropical Africa and

Madagascar and South-east Asia (Jonker, 1938). Only G. aphyllus has been recorded

in Singapore.

Ecology - All spp. of Gymnosiphon are saprophytes of wet tropical forests.

247

Although cross-pollination has never been demonstrated for the genus, several floral

structures are indicative of entomophily: coloured perianths; septal nectaries (Maas

et al., 1986); stigmatic appendages that exceed the floral tube; putative protandry

(Riibsamen, 1980); and reports of scented flowers in 2 spp. (Bentham, 1855; Vogel

1962, cited in Maas et al., 1986). Riibsamen (1986) has furthermore observed insect

larvae within Gymnosiphon flowers. Autogamy is also possible, however, as the

stigmas and anthers are located at the same height in the perianth tube, and are

often intimately associated.

Notes - The upper region of the perianth tube (including the sessile anthers) is

deciduous after anthesis, and so the identification of post-anthesis and fruiting

specimens is therefore often impossible.

1. G. aphyllus Blume

Enum. pl. Javae 1 (1827) 29 (‘G. aphyllum’); Ridl., Mat. fl. Malay. Penins. 2

(1907) 73; Fl. Malay Penins. 4 (1924) 306; Jonker, Meded. bot. Mus. Rijks-Univ.

Utrecht 51 (1938) 30, 170; Fl. Males., 1:4 (1948) 20; Backer & Bakh. f., Fl. Java 3

(1968) 214; H. Keng, Gdns’ Bull., Singapore 40 (1987) 123; I.M. Turner, K.S.

Chua & H.T.W. Tan, J. Singapore natn. Acad. Sci. 18 & 19 (1990) 63; I.M. Turner,

Gdns’ Bull., Singapore 45 (1993) 47.

G. borneense Becc.; G. pedicellatum Schltr.

Herbs, flowering at 8-17 cm tall; stem often branched. Leaves appressed, bract-

like, narrowly ovate-triangular, 1-2.8(-4.5) mm long, apex acute. Jnflorescence

racemose, branched or not, many-flowered; bracts appressed, ovate-triangular, 1.2-

3.5(-5) mm long, apex acute. Flowers with white to violet perianth, tube to 4 mm

long, outer lobes ovate, c. 2-2.5 mm long, apex obtuse, with narrow, crenate lateral

lobes, inner lobes linear-lanceolate, minute; anthers inserted immediately below

the inner perianth lobes; stigmas curved, funnel-shaped, inappendiculate; pedicel

1.5-4.0 mm long. Capsule c. 3 mm long, dehiscing by perforation of the wall between

costae. Seeds ovoid, with striate testa.

Distribution - Singapore: formerly collected from Bukit Timah, but now probably

extinct in Singapore (Keng, 1987). Thailand and Malesia (Malay Peninsula, Java,

Borneo, Sulawesi and New Guinea) (Jonker, 1938).

Ecology - Grows on decaying vegetation and humus, in wet shaded forests below

1500 m in its range.

Thismia Griff.

Proc. Linn. Soc. Lond. | (1844) 221; Ridl., Mat. fl. Malay. Penins. 2 (1907) 73;

Fl. Malay Penins. 4 (1924) 307; Jonker, Meded. bot. Mus. Rijks-Univ. Utrecht 51

(1938) 42, 227; Fl. Males., 1:4 (1948) 21; Backer & Bakh. f., Fl. Java 3 (1968) 214.

248

Saprophytic, fleshy, annual herbs with generally tuberous, coralliform or

vermiform and creeping subterranean parts, stem usually short, rarely branched.

Leaves achlorophyllous, bract-like, small. Flowers actinomorphic (occasionally

zygomorphic); subtending bracts sometimes forming an involucre; perianth 6-

merous, segments either free and of equal length, or else with very small outer

lobes and larger inner lobes connivent or connate at the apex, forming an erect

mitre with 3 holes; tube urceolate to campanulate, longitudinally striate, with a

prominent annulus at the tube mouth; stamens 6, generally laterally connate, forming

an anther tube, hanging from the annulus, occasionally free, filaments filiform or

taeniform, short; ovary 1-loculate, with 3 parietal placentas, obconical or obovoid;

style cylindrical or conical, short, thick; stigmas 3, simple or bilabiate. Fruit fleshy,

cupulate, crowned by the persistent, fleshy, basal ring of the perianth tube and the

style and stigmas, dehiscing by abscission of the apical lid. Seeds numerous, oblong,

with reticulate testa.

Distribution - C. 30 spp., distributed in tropical regions of America and Indo-

Malesia and temperate Australasia (Tasmania and New Zealand) (Jonker, 1938); 1

further sp., now extinct, was endemic to Illinois, U.S.A. Only 7: aseroe and T:

fumida have been recorded in Singapore.

Ecology - All Thismia spp. are saprophytic, predominantly occurring among

leaf litter of the tropical forest floor. As with other genera in the Burmanniaceae,

Thismia possesses several floral characteristics typical of entomophily: some spp.

have a brightly coloured annulus with nectar guides directed towards the perianth

tube; others have tentacle-like extensions of the perianth, which may act as access

routes for insects of the forest floor; the stamens often form a guide, leading the

pollinator to the stigma; many have specialized or restricted exit routes for

pollinators, ensuring that they brush past the anthers; and some spp. have glandular

swellings at the tip of the perianth segments, presumably acting as osmophores, or

glands at the base of the perianth (Vogel, 1962, cited in Maas et al., 1986). Stone

(1980) suggested that pollination of some spp. may be effected by flies, and cited

the mitriform perianth apex with its openings as evidence of the myophilous

syndrome. Self-pollination has also been demonstrated in many spp. (Maas et al.,

1986). Abscission of the perianth tube following anthesis and the fragility of the

membrane covering the ovarian chambers are indicative of seed dispersal by rain-

splash (Stone, 1980).

Key to the Species

la. Perianth tube brown to grey-yellow below and bright orange-yellow above,

with bright orange-yellow lobes which are red basally; short extensions present

between the LODES. \o5 os. -...5..0g</us veuneasscentcemtonretemapavegtecasete Ms atavtecstets T. aseroe

lb. Perianth tube white with pink stripes, and grey-green lobes; extensions between

lobes absent ssc. ...20./5..3 cise tose pateeexes rae aoe nce ay ee T. fumida

249

1. T. aseroe Becc.

Malesia | (1878) 252: Ridl., J. Straits Brch R. Asiat. Soc. 22 (1890) 336; J.

Straits Brch R. Asiat. Soc. 33 (1900) 153; Mat. fl. Malay. Penins. 2 (1907) 74; FI.

Malay Penins. 4 (1924) 308; Jonker, Meded. bot. Mus. Rijks-Univ. Utrecht 51

(1938) 240; Fl. Males., 1:4 (1948) 23; H. Keng, Gdns’ Bull., Singapore 40 (1987)

123; I.M. Turner, K.S. Chua & H.T.W. Tan, J. Singapore natn. Acad. Sci. 18 & 19

(1990) 63; I.M. Turner, Gdns’ Bull., Singapore 45 (1993) 47.

Herbs, flowering at up to c. 9 cm tall; stem usually unbranched, + succulent,

growing from the creeping white rhizome. Leaves appressed, lanceolate, c. 4 mm

long, apex obtuse. Flowers solitary, terminal on branches; bracts lanceolate, forming

an involucre at the flower base, apex acute; perianth tube brown to grey-yellow

below bright orange-yellow above, obconic-campanulate, flaring into a narrow rim,

c. 10-12 mm long, with a prominent raised annulus at the mouth, basal 5 mm of

tube with transverse bars internally; perianth lobes bright orange-yellow with bright

_ yellow or orange tentacles, red at base, triangular, equal-sized, c. 10 mm long, with

short extensions of the tube present between the lobes; anthers pendulous from the

annulus, with filaments near the top, laterally connate, forming a staminal tube,

‘ connective extensions forming broad, dorsal quadrangular wings on the outer

_ Margin, with 2 nectaries located in the furrows between extensions, the staminal

tube base with several pendulous lobes; ovary obovoid, c. 3 mm long; style short;

stigmas 3. Capsule c. 5 mm long, ribbed, with persistent style, pedicel lengthening

c. 5-7 mm above the involucre. Seeds ellipsoid.

Distribution - Singapore: currently there is only 1 known population, in Fern

Valley at the Bukit Timah Nature Reserve. Only known from the Malay Peninsula

(Singapore and Perak) (Jonker, 1938).

Ecology - Grows in the leaf litter of dense forests.

Notes - The complex floral anatomy is described in detail and illustrated by

Groom (1895).

2. T. fumida Ridl.

J. Straits Brch R. Asiat. Soc. 22 (1890) 338; J. Straits Brch R. Asiat. Soc. 33

(1900) 153; Mat. fl. Malay. Penins. 2 (1907) 74; Fl. Malay Penins. 4 (1924) 307;

Jonker, Meded. bot. Mus. Rijks-Univ. Utrecht 51 (1938) 240; Fl. Males., 1:4 (1948)

23; H. Keng, Gdns’ Bull., Singapore 40 (1987) 123; I.M. Turner, K.S. Chua &

H.T.W. Tan, J. Singapore natn. Acad. Sci. 18 & 19 (1990) 63; I.M. Turner, Gdns’

Bull., Singapore 45 (1993) 47.

Herbs, flowering at up to c. 10 tall; stem usually unbranched, + succulent,

growing from a brownish rhizome. Leaves appressed, lanceolate, apex acute to

acuminate. Flowers solitary, terminal on branches, c. 5-10 mm long, bracts not

250

observed; perianth tube white with pink stripes, + cylindrical, constricted above

the ovary and broadened below the limb, without a noticeable apical ring, annulus

prominent; perianth lobes grey-green, spreading, lanceolate, apex acute, extensions

between lobes absent; anthers not observed; ovary obconical; style very short;

stigmas 3, recurved. Capsules cupulate, ribbed, scabrid, crowned by the crenulate

basal ring of the perianth. Seeds not observed.

Distribution - 7. fumida has only ever been recorded from Petaling in Selangor,

Malaysia and Chan Chu Kang in Singapore (Saunders, 1996a).

Ecology - Grows on rotting logs in dense forests.

Notes - The scarcity of collections and their poor quality prevents descriptions

of some structures, including the bracts, anthers and seeds. The above description

is consequently largely based on that of Ridley (1890) and his drawings, now held

in the library of the Royal Botanic Gardens at Kew (published in Saunders, 1996a).

As both collections were made in the last century, the sp. is now believed to be

extinct in Singapore (Keng, 1987).

Acknowledgements

I would like to thank the Directors and Curators of B, BM, C, G, K, L, SING

and SINU for the loan of herbarium specimens and drawings, Dr D.J. Metcalfe for

the current location of Thismia aseroe in Singapore, and Rosemary Nicholls for

drawing Fig. 1.

References

Bentham, G. (1855). On the South American Triurideae and leafless Burmanniaceae

from the collections of Mr Spruce. Hooker’s J. bot. Kew Gard. Misc. 7, 8-17.

Chakrapani, P. and Raj, B. (1971). Pollen morphological studies in the

Burmanniaceae. Grana palynol. 11, 164-179.

Dahlgren, R.M.T., Clifford, H.T. and Yeo, P.F. (1985). The Families of the

Monocotyledons: Structure, Evolution, and Taxonomy. Springer-Verlag; Berlin;

215-220.

Ernst, A. and Bernard, C. (1912). Entwicklungsgeschichte des Embryosackes und

des Embryos von Burmannia candida Engl. und B. championii Thw. Annls Jard.

bot. Buitenz. 10, 161-188.

Groom, P. (1895). On Thismia aseroe (Beccar1) and its mycorrhiza. Ann. Bot. 34,

327-361, pl. 13-14.

Jonker, F.P. (1938). A monograph of the Burmanniaceae. Meded. bot. Mus. Rijks-

Univ. Utrecht 51, 1-279.

251

Keng, H. (1987). Annotated list of seed plants in Singapore (XI). Gdns’ Bull.,

Singapore 40, 113-132.

Larsen, K. (1963). Studies in the flora of Thailand. 14. Cytological studies in vascular

plants of Thailand. Dansk bot. Ark. 20, 210-275.

Maas, P.J.M., Maas-van de Kamer, H., Benthem, J. van, Snelders, H.C.M., and

Riibsamen, T. (1986). Flora Neotropica Monograph 42. Burmanniaceae. New

York Botanical Garden; New York; 1-189.

Malme, G.O.A. (1896). Die Burmannien der ersten Regnel’schen Expedition. Bih.

K. Svenska Vetensk.-Akad. Handl. 22, 1-32.

Ridley, H.N. (1890). The Burmanniaceae of the Malay Peninsula. J. Straits Brch R.

Asiat. Soc. 22, 331-339.

Riibsamen, T. (1980). Beitrage zur Mikromorphologie der Testa und zur

Embryologie amerikanischer Burmanniaceen und Triuridaceen. Diplomarbeit,

Ruhr-Universitat Bochum 1, 1-355; 2, 356-528.

— (1986). Morphologische, embryologische und systematische Untersuchungen

an Burmanniaceae und Corsiaceae (Mit Ausblick auf die Orchidaceae-

Apostasioideae). Diss. Bot. 92, 1-310.

Sarkar, A.K., Datta, N., Chatterjee, U. and Datta, R. (1973). IOPB chromosome

number reports XLII. Taxon 22, 647-654.

Saunders, R.M.K. (1996a). Typification of the name Thismia fumida Ridl.

(Burmanniaceae). Jaxon 45, 107-109.

— (1996b). The occurrence and taxonomic relationships of Burmannia wallichii

(Burmanniaceae) in Malesia. Bluemea 41, 333-337.

Schoh, M. (1920). Entwicklungsgescichtlich-cytologische Untersuchungen itiber

die Pollenbildung und Bestaéubung bei einigen Burmannia-Arten. Arb. Inst. Allg.

Bot. Univ. Ziirich 24. Diss. Ziirich.

Stone, B.C. (1980). Rediscovery of Thismia clavigera (Becc.) F. v. M.

(Burmanniaceae). Blumea 26, 419-425.

Vogel, S. (1962). Duftdriisen im Dienste der Bestéubung. Akad. Wiss. Abh. Math.-

Naurwiss. Kl. 10, 601-763.

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OBITUARY

Edred John Henry Corner

12 Jan 1906 - 14 Sep 1996

E.J.H. Corner came out East as a young man in 1929 and served as Assistant Director

of the Singapore Botanic Gardens under Professor R.E. Holttum till 1945. His

research duties were mainly concerned with the fungi, a great diversity of which

are found in the tropics. Through his wanderings in the forests of Singapore and

the Malay Peninsula he amassed a great collection of fungi and he also gradually

build up an extensive knowledge of trees and palms of the region.

In 1940 Corner published his first book, Wayside Trees of Malaya, in two volumes.

This work written in his lucid and friendly style is full of detail that reveal a personal

familiarity with the subject matter. Now in its third edition, this is still the best

book on the common (and many not so common) trees of the area covered. To

solve the problem of collecting botanical specimens from tall trees Corner developed

an idea inspired by watching monkeys trained to pick coconuts. Acquiring some

of these animals, he had them retrained as botanical specimen collectors - the first

apes in the civil service.

During the Japanese occupation Corner remained in the Gardens where he was

allowed to continue with his research. He left the Gardens in 1946, and after a

short interlude with UNESCO in Brazil, took up a teaching position at Cambridge

University where he remained until he retired in 1973 as Professor of Tropical

Botany. Reputed to be an eloquent and persuasive teacher, he attracted to tropical