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57 (Part 1) September 2005

Singapore

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THE GARDENS’ BULLETIN SINGAPORE

The Gardens’ Bulletin Singapore publishes papers on plant taxonomy (including revisions),

horticulture, phytogeography, floristics, morphology, anatomy and related fields with

emphasis on plants in the West Malesian region.

EDITORIAL BOARD

Dr R. Kiew

(Editor)

Singapore Botanic Gardens

Dr P.Y. Tan

(Assist. Editor)

Singapore Botanic Gardens

Dr $.C. Chin

Singapore Botanic Gardens

Dr M.J.E. Coode

Royal Botanic Gardens

Kew, U.K.

Dr R.T. Corlett

University of Hong Kong

Hong Kong

Dr M.C. Roos

National Herbarium of the Netherlands

Leiden University

Leiden, The Netherlands

Dr E. Soepadmo

Forest Research Institute Malaysia

Kepong, Malaysia

Dr W.K. Tan

National Parks Board, Singapore

The Gardens’ Bulletin is published twice yearly by the National Parks Board, Singapore.

Neither the National Parks Board nor the Editorial Board is responsible for the opinions or

conclusions expressed by the contributing authors.

The annual subscription for the Gardens’ Bulletin is Singapore $100.00 including postage.

Overseas subscribers are required to make payment in the form of bank drafts or

international money orders in Singapore currency payable to National Parks Board,

Singapore.

Instructions for contributing authors are found on the inside back cover.

yet ‘Ss ) a

3 eo ARBORETUM

The Gardens' Bulletin

Singapore

VOL. 57 (Part 1) August 2005 ISSN 0374-7859

CONTENTS

Ipor, I.B., C.S. Tawan and N. Jacobsen

Bonew Species ol Cryplocon ne (Ataceac) TOM! BOTMEOs:c.,.20.:.:scscscseesndesescccesentonesenctatense 1

Baharuddin Sulaiman and P.C. Boyce

A remarkable new species of Homalomena (Araceae) from Peninsular Malaysia ..... i

Boyce, P.C., Jelend Kisai and Jepom Tisai

Hapaine appendiculata (Araceae: Caladicae) tedisCOVered ...:.......2c.00c..2-s-.0cencaessasseceee 13

Truong Q.T., R. Kiew and J.J. Vermeulen

Begonia bataiensis Kiew, a new species in section Leprosae (Begoniaceae) from

Ace ain re ee Ra ee oR occ et reds oss Et aac ra dvCd Gnceapbaveaslnocead Mtaetbanuseetiramecoas¥instdhane’ ie

Turner, ILM. and K.N. Gandhi

PRCOmectOm im CleMsTOcalyx (NINTLACCAC)) <aisdccnssecesecccerednsstecasaeesenseed-seedecasucnsensavatsdeereedeneas 25

Poulsen, A.D. and R.J. Searle

Scaphochlamys calcicola (Zingiberaceae): a new and unusual species from Borneo... 29

Skornickova, J. and M. Sabu

The recircumscription of Curcuma L. (Zingiberaceae) to include the genus

Hag IEEE (EVER EAY CURRY ts SUI eet eran SE gee ass Sc nad y cee zac euiss scusdatacec lewenauaeiisar (tunica tes¥enainestanaceTaieanedsvenedaves 3]

Wilde, W.J.J.O. de and B.E.E. Duyfijes

New taxa and taxonomic status in Xanthophyllum Roxb. (Polygalaceae) from

EXCUSES) wsctewedentloshadeoga ss ¢ec6ck) 00 HEREC ere ac unENEEce OHT eEe 47

Ng, F.S.P.

Vaxonomic notes on Boriean Crypiocarya RBI; Gauraceae) ..................c.ceceecsssesnjeeee 63

Julia, S.

A synopsis of the genus Actinodaphne Nees (Lauraceae) in Sabah and Sarawak,

yeaa SUN aoe ee ce INC ERED OMe MOD Sea SMe ee cc sada ac eunsosnceanectarsussenovuieanad¥oetehnnes 69

Chung, R.C.K., E. Soepadmo and A.L. Lim

Pusymopsis on the Bomean species ol Microcos Li (Tiliacéae) 2.2.6.0icecccde.tedecsctdienss- 101

Kiew, R.

A correction: Begonia nubicola Kiew (Begoniaceae) is renamed B. oreophila Kiew

Vermeulen, J.J. and P. O'Byrne

Two new species resembling Bulbophyllum plumatum Ames and B. mirum J.J.Sm.

(Orchidaceae). .2.vidsdecec ccd de Gastnahensts cede nee eee ee

Book reviews:

Orchid Hybrids of Singapore 1893-2003 by J. Elliott Henry Oakeley

Field Guide to the Grasses of Singapore (Excluding the Bamboos)

by H. Duistermaat J.F. Veldkamp

Epiphyllous Liverworts of China by Zhu R.-L. and So M.-L. B.C. Tan

Date of publication: 30 September 2005

Published by

National Parks Board

Singapore Botanic Gardens

Cluny Road

Singapore 259569

Printed by Oxford Graphic Printers Pte Ltd

Gardens’ Bulletin Singapore 57 (2005) 1—6 |

A New Species of Cryptocoryne (Araceae)

from Borneo

I.B. IPOR, C.S. TAWAN

'Faculty of Resource Science & Technology, Universiti Malaysia Sarawak,

94300 Kota Samarahan, Sarawak

AND

N. JACOBSEN

Botanical Section, Department of Ecology, The Royal Veterinary and Agricultural

University, Rolighedsvej 21, DK-1958 Frederiksberg C, Denmark

Abstract

Cryptocoryne zaidiana Ipor & Tawan, a noteworthy new species from Long Tran, Tinjar,

Miri Division, Sarawak, is described and illustrated. This amphibious species grows on

mudflats in sandy clay soil of the intertidal freshwater zone of Sungai Mering. Its leaf shape

and texture are similar to those of Cryptocoryne lingua Engler but differ especially in its

spathe characteristics. It has a strongly recurved ovate limb, the upper surface is distinctly

covered with purplish and creamy protuberances and it has a distinct creamy collar.

Introduction

In Sarawak Cryptocoryne species occur in the inner tidal zone in rivers and

streams, on riverbanks of slow and fast flowing rivers or streams and in

ditches of swampy and flooded areas. In most places they are found in

shaded locations. They are known as kalakatai (Iban - Sarawak); kiambang

batu (Malay-Sarawak and Peninsular Malaysia) and tropong air

(Banjarmasin-Kalimantan).

The discovery of the new species Cryptocoryne zaidiana raises the

current total number of Cryptocoryne species in Sarawak to fourteen. The

eleven species reported by Jacobsen (1985) are C. ciliata (Roxburgh) Schott,

C. auriculata Engler, C. bullosa Engler, C. ferruginea Engler, C. keei N.

Jacobsen, C. lingua Engler, C. longicauda Engler, C. pallidinervia Engler,

C. striolata Engler, C. zonata De Wit and C. grabowskii Engler. Jacobsen

(2002) regards C. zonata as C. cordata Griffith var. zonata (De Wit)

N.Jacobsen and C. grabowskii as C. cordata Griffith var. grabowskii (Engler)

N.Jacobsen. Recently two other species have been described, viz. C. uenoi

Y.Sasaki (2002) and C. yujii Bastmeijer (2002), which were collected from

Gard. Bull. Singapore 57 (2005)

Sabal Keruing, Sri Aman Division and Sungai Nibong, Durin, Sibu Division,

respectively (Bastmeijer, 2005). It is essential to conduct more field work

to obtain extensive records of the existence and occurrence, as well as

understanding of the natural diverse habitats of Cryptocoryne species in

Sarawak.

Description of the Species

Cryptocoryne zaidiana Ipor & Tawan sp. nov.

Differt a C. lingua folia majore, limbus ovatus, valde recurvatus,

protuberance supra sed. Typus - Sarawak: Miri Divison, Tinjar, Long

Tran, Sungai Mering, whole plant with inflorescence and young syncarp, 2

July 2004, C.S. Tawan, I.B. Ipor & A. Mohd Rizan CST 2548, (holotypus,

HUMS (Herbarium Universiti Malaysia Sarawak) - in spirit).

Rhizome stout, internode length 1.0—3.0 cm, 4.5—6.8 mm diam., whitish,

smooth, adventitious roots contracted at the nodes, new roots whitish and

later becoming brownish due to mud stains, 1—14 cm long, Cataphylls 1—2

keeled adaxial to the flowering shoot, ovate, 5 cm long, 1.5 cm across at

the middle, convolute, base whitish, light greenish upper portion, slightly

hyaline near the margin, smooth on both surfaces, base truncate, apex

acute. Leaves with upper surface green, smooth and a shiny, lower surface

smooth and pale green or sometimes slightly purplish; blade ovate, 9.5—

13.5 x 4.0-6.0 cm, base truncate-cordate, apex acute, margin entire

sometimes slightly wavy near the base; midrib flattened on both surfaces,

secondary veins 4 pairs, not prominent, slightly channeled on both surfaces;

petiole elongate, 19—23 cm long, 3.6—5.6 mm diam. in middle, leaf sheath

c. 3.5—4.5 cm long with hyaline margins, middle portion rounded or

sometimes angled, upper part slightly flattened, green, lower part usually

brownish due to mud stains. Spathe 9—14 cm long, pedunculate 1.5—2.5

cm long, 6 mm diam., whitish, smooth; kettle 2—2.8 cm long, constricted

about 1/3 from the top; outer surface with a whitish background with faint

purplish longitudinal lines, inner surface 1/3 of the upper portion, whitish,

2/3 lower portion sprayed with fine purple spots; tube 4.5—7 cm long,

narrow, cylindrical at base 5 mm diam., broader in the upper part, 7.5 mm

diam., outer surface faintly purplish, more intense in the upper part; inner

surface whitish, slightly twisted, tube fusion line distinct, purple; limb

strongly recurved, 3.0—4.0 cm long, 1.5—2 cm across (near the collar), ovate,

apex apiculate, 3—4 mm long, dark purple, upper surface with protuberances

or warty, creamy-whitish, margin and surface towards the apex covered

Cryptocoryne zaidiana in Borneo

1.5 cm

2.5 mm

2cm

Figure 1. Cryptocoryne zaidiana \por & Tawan.

A Plant with inflorescence and fruit; B inner surface of the kettle; C limb surface and collar;

D spadix showing the male and female flowers; E thecae; F stamen; G syncarp with verrucose

surface; H seed. (A-F from the holotype CST 2548; G-F from the CST 2549 - drawn from the

fresh material).

4 Gard. Bull. Singapore 57 (2005)

with distinct dark purple protuberances, lower surface whitish or faintly

purplish; collar distinct, creamy; throat surface deep purplish or sprayed

with purplish spots, surface smooth. Female flowers c. 6, stigma ovate-

elliptic, purplish; ovary whitish elongate, 3 mm long, 1 mm wide, Male

flowers c. 50, smooth, creamy or light yellow; naked axis spadix 9-10 mm

long, purplish; sterile appendix c. 1.5 x 1.0 mm, ovate, deep purple; olfactory

bodies dark purple; flap ovate, 4-5 x 3—3.5 mm, whitish. Fruit peduncle

3.5—6.0 cm long, 3—4.5 mm diam., whitish or sometimes speckled with dark

purple spots. Syncarp broadly ovoid, 1.5—1.7 cm long, 12—16.5 mm diam.,

dull green speckled with dark purple, slightly verrucose surface, apex

distinctly apiculate, dark green-purple. Seeds elongate 6—7 mm long, 1.8—

2.0 mm broad at base, dark purple in the upper portion, lower part whitish,

surface slightly striated. Embryo with three long, green plumulary processes.

Other specimens studied: Type locality, whole plant with inflorecence, 16

June 2004, C.S. Tawan , I.B. Ipor & A. Mohd Rizan CST 2545 (HUMS) -

in spirit from cultivated plant collected from the type locality; herbarium

specimens from the type locality, whole plant with mature syncarp, 2 July

2004, C.S. Tawan , I.B. Ipor & A. Mohd Rizan CST 2550 (HUMS); whole

plant with inflorescence and young syncarp, 2 July 2004, C.S. Tawan, I.B.

Ipor & A. Mohd Rizan CST 2551 (HUMS); whole plant with inflorecence,

2 July 2004, C.S. Tawan, I.B. Ipor & A. Mohd Rizan CST 2552 (SAR).

Distribution: Endemic in Sarawak, as yet known only from Sungai Mering,

Tinjar, Miri Division.

Habitat: Cryptocoryne zaidiana occurs in small patches on muddy ground

(sandy clay soil) with a litter of leaves and twigs. The river is approximately

5—7 m wide and flooded with a considerable slow current after a period of

heavy rain. The riverbank is established with secondary riverine forest

(15-18 years after padi planting according to the local people). The forest

undergrowth is mainly dominated by bemban, Donax grandis

(Marantaceae). The river normally becomes shallow or sometimes dries

up after dry periods. At this time, this river is a popular place for the local

people to ‘mansai’ (to fish using round-shaped nets to scoop in the shallow

water to catch small fish). This regular activity appears to disturb the

habitat of C. zaidiana. More severe disturbance is imminent as the area is

earmarked for oil palm plantations.

Notes: Cryptocoryne zaidiana shows in its habitat and morphological

characteristics certain similarity with C. lingua. These two species thrive

well on mudflats of riverine clay soil along the fringes of riversides and

Cryptocoryne zaidiana in Borneo 5)

A Scale bar-6 cm H cale bar-1 cm

Plate 1. Cryptocoryne zaidiana Ipor & Tawan.

A Plant with unopened inflorescence; B, C limb surface and collar; D syncarp with verrucose

surface; E seeds; F the male and the female zones.

6 Gard. Bull. Singapore 57 (2005)

ditches. C. lingua grows well in a certain level of salinity during short

drought spells in the Sungai Sarawak (Ipor, pers. obs.). To date, C. zaidiiana

has only been found in freshwater within the intertidal zone. Both species

have spongy leaves but C. lingua has an ovate to oblong blade with a

truncate base whereas C. zaidiana has ovate blades with a truncate to

cordate base. This species is different from C. /ingua in having an ovate,

strongly recurved limb with protuberances on the upper surface. C. lingua

has a caudate, smooth limb without protuberances on the upper surface.

Etymology: This species is named in honour of the late Tun Ahmad Zaidi

Adruce, who was the first Chancellor of Universiti Malaysia Sarawak

(UNIMAS) and the fifth Governor of Sarawak, in recognition of his great

contribution to the state of Sarawak and in particular to research

development at UNIMAS during his lifetime.

Acknowlegements

We thank Universiti Malaysia Sarawak for the short-term grant to undertake

this project. Our gratitude also goes to the Sarawak Forestry Department

for permission to collect the specimens. We should also like to thank

Mohd Rizan Abdullah, James Abai and Hidir Marzuki for their help. Our

appreciation also goes to Joseph Pao for the botanical illustration.

References

Bastmeijer, J.D. 2002. Cryptocoryne yujii Bastmeijer (Araceae), eine neue

Art aus Sarawak. Aqua Planta. 27: 145—146.

Bastmeyjer, J.D. 2005. The Crypts pages http://132.229.93.11/Cryptocoryne/

index.htm]

Jacobsen, N. 1985. The Oe: (Araceae) of Borneo. Nordic Journal

of Botany. 5: 31—S0.

Jacobsen, N. 2002. Der Cryptocoryne cordata Griffith — complex (Araceae)

in Malesian. Aqua Planta. 27: 150-151.

Sasaki, Y. 2002. Cryptocoryne uenoi Y. Sasaki (Araceae), eine neue Art

aus Sarawak. Aqua Planta. 27: 147-148.

Gardens’ Bulletin Singapore 57 (2005) 7—11 ‘|

A Remarkable New Species of Homalomena

(Araceae: Homalomeneae)

from Peninsular Malaysia

BAHARUDDIN SULAIMAN

School of Biological Sciences, Universiti Sains Malaysia,

11800 Minden, Pulau Pinang, Malaysia

AND

PETER-C. BOYCE

Malesiana Tropicals, Suite 4, Level 9, Tun Jugah Tower,

Jalan Abdul Rahman, 93000 Kuching, Malaysia

Abstract

Homalomena pineodora Sulaiman & P.C.Boyce is described as a new species from Peninsular

Malaysia. Its similarities to H. bellula Schott (Java), H. elegantula A.Hay & Herscovitch

and H. hastata M.Hotta (both Sumatera) and an as yet undescribed Homalomena from

North Vietnam are discussed.

Introduction

Homalomena is a genus of more than 120 species of terrestrial or lithophytic,

seldom rheophytic, clumping, rarely rhizomatous, very rarely climbing,

mostly aromatic herbs distributed in the Neo- and Asian tropics. As noted

by Hay & Herscovitch (2002), the genus is overwhelmingly Asian in

distribution with the greatest number of species occurring in archipelagic

Malesia. The taxonomy of Homalomena is presently in disarray, the problem

compounded by the poor condition of many of the historical types and

inadequate understanding of interspecific variability.

There is a tendency in herbaria to apply the names of continental

Asian, Sumateran, Javan and Bornean species somewhat indiscriminately

while ignoring the high levels of endemism known to exist for other

terrestrial aroid genera (e.g., Alocasia, Schismatoglottis) in these regions.

With more fieldwork it is becoming apparent that for these land masses

the level of species endemism averages 80% (Boyce, unpublished data)

and that species novelty for most terrestrial aroid genera is in excess of

75% (Boyce, unpublished data); there is no evidence to suggest that

Homalomena is any different: Sarawak alone appears to have at least 80

8 Gard. Bull. Singapore 57 (2005)

species of Homalomena of which no fewer than 65 are as yet undescribed.

Until such time as a full revision of Homalomena is attempted, there is still

a need to be able to identify the more distinctive taxa. The only practical

approach is to describe new taxa as they become apparent with the caveat

that at a future date some of these taxa may prove to be unsustainable in

light of a more comprehensive and thorough revision.

Despite Furtado’s attempts to unravel Homalomena in Peninsular

Malaysia (Furtado, 1939) and accepting the evidence that Homalomena is

far more species-rich in archipelagic rather than in continental Asia, much

remains to be done in Peninsular Malaysia before it can be said that

Homalomena is properly understood. That much remains to be discovered

was evidenced during fieldwork by the first author at Pondok Tanjung,

Selama, Perak, where a collection was made that at first appeared to be a

species of Piptospatha in that it was growing in a rheophytic habitat and

carried nodding inflorescences. However, further examination revealed that

the petiolar sheaths were fully adnate to the petiole (adnate basally but

otherwise free and long ligulate in Piptospatha), the spathe showed no

abscission layer between the lower part and the limb (all west Malesian

Piptospatha shed the upper spathe at anthesis), and that the leaf tissue

when crushed emitted a very strong resinous smell (tissue odourless in

Piptospatha). Examination of fresh inflorescences revealed the pistils were

each associated with a staminode confirming, together with the above listed

characters, that the plant belongs to the genus Homalomena. It is here

described.

Homalomena pineodora Sulaiman & P.C.Boyce, sp. nov.

Ab alii Homalomena Malesiarum caudice repenti hypogeo pleionathico et

inflorescentia cernuua differt. Typus: Malaysia, Perak, Pondok Tanjung,

Selama, 5° 0'N; 100° 45'E, B. Sulaiman BS 1259 (holo Universiti Sains

Malaysia 10811; iso KEP, SING).

Dwarf creeping evergreen strongly aromatic (terpenoids - reminiscent of

Pinus) when cut or crushed herb to 25 cm tall; cut surfaces producing

copious watery, later sticky, aromatic sap. Stem rhizomatous, creeping, c.

8-10 mm diam., rooting along its length over the entire surface, oldest

parts of stems clothed with papery to fibrous leaf base remains, roots

penetrating this fibrous layer, roots also penetrating the leaf bases in the

lower active portion; distal active shoot erect to semi-procumbent; active

part of shoots densely clothed with spiro-distichous leaves; shoots

pleionanthic. Leaves clustered towards shoot tips; petiole partially clasping

the rhizome and then expanding into a persistent petiolar sheath, petiole

8.5—9 cm long, non-sheathing portion c. 2.5 mm diam., weakly D-shaped in

Homalomena pineodora from Peninsular Malaysia 9

Beer Mears!

Figure 1. Type plant of Homalomena pineodora Sulaiman & P.C. Boyce showing the distinctive

nodding inflorescences.

cross-section and dorsally shallowly grooved, the groove extending to the

insertion of the lamina; sheath extending to c. °/s petiole length, margins

hyaline, long-persistent; /amina leathery, elliptic to weakly oblongo-elliptic,

9—12 x 4—5 cm, base cuneate to ovate, apex acute with a prominent 3.5—4

mm long tubule, margins minutely hyaline (margins c. 0.5 mm wide) and

tending to crispulate at the petiole insertion, abaxial surface pale green

with up to 5 (mostly 3—4) weakly prominent and slightly darker primary

lateral veins on each side of the mid-vein; interprimaries much less

prominent and not at all darker than the abaxial lamina; adaxial lamina

surface slightly lustrous dark green, primary lateral veins slightly impressed,

interprimary veins hardly discernible. [Inflorescences nodding, two together,

maturing sequentially, each subtended by a two-keeled prophyll. Peduncle

c.5 cm long, 2.5 mm diam., weakly D-shaped in cross-section and expanding

distally at the insertion of the spathe. Spathe unconstricted, ovate-ellipsoid,

inflated and gaping at anthesis, c. 3 x 1.8 cm (rolled) (5.5 cm wide when

flattened) ovato-triangular, apex acuminate; spathe exterior glossy bright

green, the upper half deeper green, the lower half somewhat yellow-white,

at anthesis spathe margins recurved, hyaline and slightly brownish along

10 Gard. Bull. Singapore 57 (2005)

the edge; spathe interior glossy greenish white. Spadix 20 x 8 mm, stipitate,

stipe c. 1 mm. Pistillate flower zone 6 x 6 mm with a zone of very slender-

stalked globose-claviform 0.1 x 0.2 mm glossy white staminodes basally.

Ovaries compressed-globo-cylindrical, 0.2 x 0.4 mm, stigma sessile, circular

and slightly umbonate, overhanging the ovary, 0.7 x 0.4 mm, ovaries pale

green, stigma translucent greenish white. Male flower zone ovoid, 10 x 7

mm, fertile to the apex, flowers comprised of two stamens, rarely one

stamen aborted, 0.4 x 0.5 mm, connective barely prominent, male flowers

pale cream. /nfructescence and seeds not observed.

Distribution: Peninsular Malaysia (NW Perak), known only from the type

collection.

Habitat: Riverside, growing in clumps on an inundated site under forest

canopy. Sea level.

Notes: The rhizomatous stems, pleionanthic shoots and inflorescences

nodding at anthesis immediately distinguish Homalomena pineodora from

any Homalomena hitherto described from Peninsular Malaysia and adjacent

peninsular Thailand. Rhizomatous creeping stems are a feature of two

species described from Sumatera (H. elegantula A.Hay & Herscovitch, H.

hastata M.Hotta,) and one from Java (H. bellula Schott). Among characters

that distinguish these species from H. pineodora are hapaxanthic shoots

(all), inflorescences erect at anthesis (all), hastate leaves (H. hastata),

apically geniculate petioles (H. elegantula) and smaller (1 cm long) spathes

(H. elegantula). An as yet undescribed Homalomena from Ba Vi, north

Vietnam, has rhizomatous creeping stems, pleionanthic shoots and nodding

inflorescences but differs from H. pineodora by the longer petioles with

the petiolar sheath c. 1/4 the length of the petiole, broader ovato-cordate

leaf laminae, a longer, narrower spadix and a markedly different smell

produced by the crushed tissue (the Ba Vi Homalomena has a smell

resembling Juniperus — H. pineodora smells of Pinus). The odours produced

by the damaged tissue of H. elegantula, H. hastata and H. bellula are not

recorded.

The specific epithet is from the Latin pinea and odora meaning

‘pine-scented’ in allusion to the very strong smell of Pinus produced by

cutting or crushing the plant tissue.

Homalomena pineodora from Peninsular Malaysia a

References

Furtado, C.X. 1939. Araceae Malesicae II. Notes on some Indo-Malaysian

Homalomena species. Gardens’ Bulletin Straits Settlements. 10: 183-238.

Hay, A. and C. Herscovitch. 2002. Two remarkable new West Malesian

Homalomena (Araceae) Species. Gardens’ Bulletin Singapore. 54: 171—

178.

she ; hie (of we

ui Lobat ey un

c it i wales een Bt

i r we | aa ina eT ah

“s

Gardens’ Bulletin Singapore 57 (2005) 13—18 IS

Hapaline appendiculata (Araceae: Caladieae)

Rediscovered

PETER C. BOYCE, JELEND ak KISAI AND JEPOM ak TISAI

Malesiana Tropicals, Suite 9-04, Tun Jugah Tower,

No. 18, Jalan Tunku Abdul Rahman,

93100 Kuching, Sarawak, Malaysia

Abstract

Hapaline appendiculata Ridl., a species endemic to Sarawak, Borneo, and hitherto known

from only three collections, has recently been recollected. An expanded description for the

species, first ever published field photographs and a key to Hapaline in Borneo is presented.

Introduction

Recent and on-going fieldwork in Sarawak continues to reveal a wealth of

novel aroid species and has led to the rediscovery of numerous others

known from fewer than five collections. One such recent discovery was

that of Hapaline appendiculata Ridl., an aroid known from just three

previous collections, all fragmentary, and thus seemingly representing one

of the rarest aroids in Borneo.

Hapaline

Hapaline is a genus of seven published species occurring from Myanmar to

China (Yunnan) and south to western Sarawak. All species so far discovered

are diminutive to moderate-sized, slender, tuberous or stoloniferous, clump-

forming, terrestrial and seasonally dormant or (rarely) evergreen herbs.

See Boyce (1996) for an extensive commentary and generic description.

Hapaline is superficially similar to species of the genus Typhonium

Schott (tribe Areae), most notably in the sagittate leaves with prominently

reticulate venation. In Borneo there are only three 7yphonium species, all

introduced and all associated exclusively with ruderal habitats whereas

Hapaline is only found in natural forest. Elsewhere in the range of Hapaline

(e.g., Thailand) there are strikingly similar species of Typhonium in natural

habitats and confusion is possible between these and Hapaline. In these

instances the floral characters to distinguish Typhonium include the naked

spadix appendix (spadix appendix covered in synandrodes: Hapaline), the

inflated, convolute lower spathe (lower spathe clasping the female flower

14 Gard. Bull. Singapore 57 (2005)

zone: Hapaline), free male flowers (stamens fused into synandria: Hapaline)

and the female flower zone free (fused to the abaxial lower spathe surface:

Hapaline).

Hapaline appendiculata Ridl.

Hapaline appendiculata Ridl., J. Straits Branch Roy. Asiat. Soc. 49 (1908

(‘1907°)) 47; Bogner, Pl. Syst. Evol. 144 (1984): 62 (1984); Boyce, Kew

Bull. 51(1) (1996) 63—82 (1996). Type: Sarawak, Kuching Division, Puak,

Sept. 1890, Ridley 12411 (holotype SING!; isotype K!).

Slender, tuberous, seasonally dormant perennial herb up to 25 cm tall.

Stem: tuberous, + cylindric, c. 1.5 x 1 cm; plants frequently producing two

to five slender (c. 3 mm diam.) stolons to 30 cm long, these rooting

terminally and forming additional tubers. Roots c. 0.25—0.33 mm in diam.,

mostly spreading through the leaf litter-soil interface. Leaf prophyll linear,

up to 9x c. 4 mm, acute; cataphylls oblong-lanceolate to linear triangular

or triangular, up to 10 cm x 5 mm, attenuate to acute, prophyll and cataphylls

at first membranous and pale green, soon darkening and drying papery;

petiole 2.5—18 cm x 1—2 mm; leaf blade ovate to hastate or subsagittate,

10-21 x 3.5-8.5 cm, thinly coriaceous or coriaceous, even on the same

plant, pale to dark green, occasionally with various greyish to pale green

blotchy or/or cloudy markings adaxially, abaxial surface much paler,

sometimes suffused reddish purple in which case primary mid-vein and

primary lateral veins on abaxial surface purple-red, margins smooth, apex

acute to acuminate, posterior lobes rounded to subacute, divergent to almost

parallel, sometimes + absent. Inflorescence (1)2—4 together held level with

or below the leaves; peduncle 4—25 cm x 0.25-0.5 mm; spathe 2—7 cm

long; spathe limb elliptic, 1.6—2.6 cm x 5—7 mm, apex acute to briefly

attenuate, base decurrent into lower spathe; lower spathe margins clasping

the ovaries, 4—8 x c. 1.5 mm; spadix 2.5—3.5 cm x 0.25—1 mm, free portion

cylindric, up to 14 mm long, tapering apically into a greatly attenuated

appendix to 9 mm long composed of connate synandrodes. Flowers

synandria irregularly elongate in plan view, 2—3 x 0.5—1 mm; ovaries bottle-

shaped, 1.5 x 0.7 mm, two or three in a single row aligned longitudinally

along the spadix axis; stigma capitate, c. 0.2 mm in diam., papillose; style

very short. Infructescence carried on declinate to reflexed peduncle,

enclosed by the persistent lower spathe, 2 cm x 4 mm, few-berried; berries

more or less globular, ripening pale green, c. 4 mm in diam, stigmatic

remains persistent, not prominent. Seed ellipsoid, c. 3 x 2 mm, glossy pale

brown with a conspicuous white oily raphe.

Distribution: Endemic to Sarawak with four collections to date, these only

Hapaline appendiculata from Borneo

Plate 1. Hapaline appendiculata Ridl. 1. A typical flowering individual at Tringgus (Boyce,

Kisai & Tisai AR-1017). Note the greatly elongated appendix comprised of fused synandrodes;

2. A form with little or no posterior leaf lobe development; 3. An example of the leaf

variegation present at Tringgus.

16 Gard. Bull. Singapore 57 (2005)

from Kuching Division (three collections) and Kapit Division (one

collection).

Other specimens seen: SARAWAK. Kuching Division: Krokong, Kampung

Tringgus, Sungai Bong, 01° 15' 22.8"; 110° 05' 53.9", 4 March 2005, P. Boyce,

Jelend & Jepom AR-1017 (SAR, UNIMAS); Bau, Bidi, 6 Dec. 1905, Hewitt

476 (SING!). Kapit Division: Belaga, Long Linau, near Punan Lusong to

Long Jakah, 8 Sept. 1978, Burtt & Woods 11477 (E!).

Ecology: Lowland evergreen moist valley forest on shales and basalt. Only

recorded altitude c. 150 m asl (Tringgus). Both Bornean Hapaline are

associated with shale and basalt whereas all other Hapaline species (all

extra-Bornean) are restricted to karst limestone. Hewitt 476 is purportedly

from a limestone habitat (Bidi). However, repeated visits to the site by the

authors have failed to refind Hapaline and it is possible that the locality

data are in error.

Notes: Based on the paucity of collections, Hapaline appendiculata appears

to be one of the rarest aroids in Borneo. However, considering its diminutive

size, the periodically dormant nature of the plant and the fact that aroids

have received scant attention from field workers since Ridley’s time, it is

equally possible that it is simply overlooked during fieldwork. It is worth

noting that the Kuching and Kapit collections of H. appendiculata are 340

km apart with much of the intervening forest botanically unexplored.

Hapaline appendiculata was not visible during several previous visits

by the authors to Tringgus although on the trip that lead to the collection

discussed here it was abundant and flowering gregariously.

The method of seed dispersal is unknown although the oily raphe

suggests ant dispersal, as occurs in other aroid genera with similar seed,

e.g., Biarum Schott and Eminium Schott (both Mediterranean). The first

author has observed morphologically similar seed of Zingiber

pseudopungens R.M.Sm. and Z. pachysiphon B.L.Burtt & R.M.Sm.

(Zingiberaceae) being carried by solitary foraging ants of the genus

Campanotus.

Hapaline appendiculata differs from H. celatrix P.C.Boyce (Brunei)

by its seasonally dormant habit, thinly coriaceous leaves and greatly

elongated sterile appendix. The habitat of H. celatrix is briefly seasonally-

dry riverine forest on shale. Both known localites of H. celatrix are more

exposed (higher light levels) and presumably less humid for at least part of

each day than the habitats of H. appendiculata. In neither locality is H.

celatrix abundant.

Hapaline appendiculata from Borneo

Plate 2. Hapaline appendiculata Ridl. 4. A form with hastate concolorous leaves: 5. Showing

the range of variation in leaf shape and markings present at Tringgus. These individuals were

collected within 20 metres of one another.

18 Gard. Bull. Singapore 57 (2005)

Key to the Two Species of Hapaline in Borneo

la. Spadix sterile apex greatly elongated, exceeding spathe limb; foliage

thinly Ccomacéous. 4225 620. - eee H. appendiculata

1b. Spadix sterile apex not greatly elongated, not exceeding spathe limb;

foliage thickly comaceous:......4.72., 099s 2o eee ee H. celatrix

Acknowledgements

The collaboration and support of the Sarawak Forestry Department, the

Sarawak Biodiversity Centre, in particular Datin Eileen Yen Ee Lee, the

Forest Research Centre (Kuching), notably L.C.J. Julaihi, and UNIMAS,

in particular Assoc. Prof. Isa Ipor, is gratefully acknowledged. Thanks are

due to Datuk Amar (Dr) Leonard Linggi Tun Jugah, Graeme Brown and

Dr Timothy Hatch of Malesiana Tropicals Sdn Bhd for their support and

funding of fieldwork in Sarawak.

References

Bogner, J. 1984. On Hapaline appendiculata and Phymatarum borneense,

two rare Araceae from Borneo. Plant Systematics and Evolution. 44: 59—

66.

Boyce, P.C. 1996. Hapaline (Araceae: Aroideae: Caladieae). Kew Bulletin.

51: 63-82.

Gardens’ Bulletin Singapore 57 (2005) 19—23 19

Begonia bataiensis Kiew, a New Species in

Section Leprosae (Begoniaceae) from Vietnam

TRUONG QUANG TAM

Institute of Tropical Biology

85 Tran Quoc Toan, Quan 3

Hochiminh City, Vietnam

RUTH KIEW

AND

J.J. VERMEULEN

The Singapore Botanic Gardens

1 Cluny Road, Singapore 259569

Abstract

Begonia bataiensis Kiew with long, cylindric, fleshy fruits is the fourth species to be described

in the largely Chinese section Leprosae.

Introduction

The great majority of Begonia species have dry capsular fruits that are

conspicuously winged and that dehisce to release the seeds. The only Asian

sections with berries (fleshy fruits that do not dehisce) are sect. Sphenanthera

and sect. Leprosae, which differ primarily in fruit shape - turbinate in sect.

Sphenanthera and narrowly cylindric in sect. Leprosae. This new species,

Begonia bataiensis Kiew, with its cylindric, fleshy, indehiscent fruit therefore

falls within sect. Leprosae.

Begonia sect. Leprosae accommodates, besides this new species,

three others, B. leprosa Hance, B. longicarpa K.Y.Guan & D.K.Tian and

B. cylindrica D.R.Liang & X.X.Chen (Shui et al., 2002). Apart from B.

longicarpa, which has been collected from the Lao Cai Province, in the

extreme north of Vietnam, just across the border from China, the other

two are confined to southern China (Shui et al., 2002). This new species

therefore extends the distribution of this section much further south to

Kien Giang Province in southwest Vietnam.

While sharing the character of the fleshy, narrowly cylindric fruit,

it is quite unlike the other three species in habit - the other three attain a

20 Gard. Bull. Singapore 57 (2005)

much larger size and are plants with rhizomes, leaves with a ferrugineous

indumentum and dichasial inflorescences that are shorter than the leaves

as compared with B. bataiensis, which is stemless, has a basal tuber that

produces just one or two or up to five small leaves with scattered translucent

hairs, and racemose inflorescences longer than the leaves (Plate 1A). In

addition, its fruit is not clavate as are those of B. /eprosa and B. longicarpa,

but instead are narrowly tapered distally.

It is probable that this section is artificial and that this fruit type

has evolved more than once. Indeed, in its habit (stemless with a tuber), its

almost symmetric leaves and its racemose inflorescence, this new species

more closely resembles species in sect. Diploclinium III of Doorenbos et

al. (1998) than other species in sect. Leprosae.

Begonia bataiensis Kiew, sp. nov.

Differt a Begonia leprosa tuberiformis (nec rhizomatis), foliis 5-6 cm

longis (nec (4—)7.5—15 cm longis) et inflorescentia racemosa (nec dichasia).

Typus: Vietnam, Ba Tai Hill, Kien Giang Province. J.J. Vermeulen 2586

(spirit collection 26/0) October 2004 (holo HCMC, iso HN, SING).

Tuberous begonia without a stem. Tuber turbinate, mature ones smooth,

brown and glabrous, c. 10 x 12 mm; young ones white, ellipsoid, c. 6 x 12

mm, shoot apex covered in many erect, fleshy stipules, narrowly triangular

3—4 x 1 mm. Leaves 1 or 2 or up to 5 per tuber. Petiole pale red, glabrous,

1.5—3.5 mm long, terete. Lamina not oblique, held horizontally; upper

surface plain dark green or pale green and flecked by the many raised

white hair bases, darker around the margin with white teeth and with a red

patch at the base above the junction with the petiole (Plate 1B), the lower

surface paler and scintillating or the upper surface brownish red and the

lower surface pale rosy purple, in life thin and slightly fleshy, hairs erect on

a raised hair base, translucent, uniseriate, c. 0.25 mm long, glabrous beneath,

ovate, symmetric except that the basal lobes are sometimes unequal, 4.5—6

x 5—6.5 cm, base deeply cordate, basal lobes equal or unequal, not

overlapping, 0.5—2 cm long, margin doubly crenate, each tooth tipped by a

hair c. 0. 5 mm long, apex acute to acuminate; venation basically palmate

with 1—2 pairs of veins at the base and c. 2—3 pairs of fine veins along the

midrib, and 1—2 veins in the basal lobes, veins branching c. half-way to the

margin, slightly impressed above, slightly prominent beneath, glabrous on

both surfaces.

Inflorescences axillary, reddish purple, glabrous, erect and longer than the

leaves; racemes 6—12 cm long with peduncles 5.5—9 cm long, cymules spaced

1—1.5 cm apart, lower two cymules comprising one male and one female

Begonia bataiensis from Vietnam

Plate 1. Begonia bataiensis Kiew. A. Plants in nature with male flower and fleshy cylindric

purple fruits, B. Protandrous raceme with the first male flower open, C. Female flower. J/./.

Vermeulen

22 Gard. Bull. Singapore 57 (2005)

flower or a short branch c. 2 mm long with three flowers (one female and

two male flowers), upper cymules with male flowers only, protandrous.

Bract pair broadly ovate, recurved, c. 2.5 x 2 mm; bracteoles ovate, c. 4 x 1

mm or linear, c. 1 mm long. Male flowers with pale reddish purple pedicels

up to 10 mm long; tepals 4, in bud greenish white and minutely hairy

outside, at anthesis glabrous and pure white, margin entire, apex rounded

to slightly acute, outer two broadly ovate to circular, 5—6 x 5.5—7 mm,

inner two elliptic to slightly obovate, 4.5—5 x 1.75—2.5 mm; stamens c. 30—

40, cluster hemispherical 2.5—3 mm diam., joined at base for c. 0.5 mm,

filament 0.5—1 mm, anther pale yellow, obovoid, c. 1 mm long, apex

emarginate, opening by lateral slits. Female flowers with reddish purple

pedicel 1.5—2 mm long; ovary deep reddish purple, narrowly cylindric, 7—

16 x 1.5 mm, glabrous, locules 3, placentas fleshy and bifid; tepals (4 or) 5,

pure white, margin entire, glabrous, apex rounded or slightly acute, outer

two broadly ovate to circular, 4-8 x 3—5 mm, inner (two or) three narrowly

obovate, 3.5—7 x 1.75—4 mm; styles and stigmas yellow, 2—3.5 mm long,

styles 3, almost free to the base, bifid and webbed, stigmas horizontal and

sigma-shaped. Fruit a fleshy pendent berry, pedicel 1.5—2 mm, stout; berry

slender, smooth and cylindric, tapered at both ends, 25—30 x 2—2.5 mm,

glabrous, without wings or ribs, indehiscent. Seeds truncate, brown and

sculptured, c. 0.3 mm long, collar cells c. half the seed length.

Distribution: Endemic to Ba Tai Hill and Bai Voi (Mo So) Hill, 3 km

NNW of Hon Chong, Kien Giang Province, SW Vietnam.

Habitat: Rooting in soil pockets in clefts in limestone rock, well shaded, at

c. 50 m altitude.

Specimens examined: Ba Tai Hill J.J. Vermeulen 2587 (spirit collection

2611) (SING); Bai Voi (Mo So) Hill Truong Quang Tam MS 056 (HCMC).

Notes: This species was discovered by the first author in October 2000 on

the Ba Tai and Bai Voi Hills. It takes its name from Ba Tai Hill. Both are

limestone karst hills and Bai Voi Hill is scheduled for quarrying with

about two thirds of the hill to be exploited for cement production. Begonia

bataiensis is therefore highly endangered, because its narrow distribution

and small population size make it vulnerable to extinction.

The racemose inflorescence produces female flowers only in the

two lowermost cymules. In these cymules, the male flowers open first and

have fallen before the female flower opens. Thereafter, a series of cymules

is produced that are comprised of only male flowers.

Begonia bataiensis from Vietnam 23

Acknowledgements

J.J. Vermeulen thanks Holcim, the parent company of the Swiss quarrying

company, Morning Star, for sponsoring the biodiversity survey of the

limestone hills.

References

Doorenbos, J., M.S.M. Sosef and J.J.F.E. de Wilde. 1998. The sections of

Begonia including descriptions, keys and species lists. Wageningen

Agricultural University Papers. 98-2: 1—266.

Shui, Y.M., C.I.- Peng and C.Y. Wu. 2002. Synopsis of the Chinese species

of Begonia (Begoniaceae), with a reappraisal of sectional delimitation.

Botanical Bulletin of Academia Sinica. 43: 313—327.

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Gardens’ Bulletin Singapore 57 (2005) 25—27 25

A Correction in Cleistocalyx (Myrtaceae)

I. M. TURNER

P.O. Box 20, Winchelsea, East Sussex TN36 4WA, UK

AND

K. N. GANDHI

Harvard University Herbaria, Divinity Avenue, Cambridge, Massachusetts

02138, USA

Abstract

Cleistocalyx nervosus (DC.) Kosterm. is an illegitimate later homonym of Cleistocalyx

nervosus (Lour.) Blume. Therefore, the new combinations Cleistocalyx cerasoides (Roxb.)

I.M. Turner and C. cerasoides var. paniala (Roxb.) I.M. Turner are proposed. Eugenia

cerasoides Roxb. is lectotypified.

Cleistocalyx nervosus

When Blume described the genus Cleistocalyx in 1850 he included two

species. These were Cleistocalyx nitidus (Korth.) Blume, based on Jambosa

nitida Korth., and Cleistocalyx nervosus (Lour.) Blume, based on Eugenia

nervosa Lour. Merrill and Perry (1937) resurrected Blume’s genus from

obscurity, including 21 species in their revision. One of these was

Cleistocalyx operculatus (Roxb.) Merr. & L.M. Perry based on Eugenia

operculata Roxb. Roxburgh first used this name in his Hortus Bengalensis

published in 1814, but lacking a description the name was not properly

validated until the posthumous publication of the second edition of Flora

Indica that appeared in 1832. However, before that in 1828, De Candolle

had described the species as Syzygium nervosum, referring to Roxburgh’s

nomen nudum Eugenia operculata. Thus the oldest valid name for this

species is Syzygium nervosum DC. Panigrahi and Mishra (1985) argued

that the 1832 publication of Eugenia operculata Roxb. represents an

illegitimate renaming of Syzygium nervosum DC.

Recognising Syzygium nervosum DC. as the oldest name available,

Kostermans (1987) made the transfer of the epithet to Cleistocalyx.

However, Cleistocalyx nervosus (DC.) Kosterm. is an illegitimate later

26 Gard. Bull. Singapore 57 (2005)

homonym of Blume’s second species of the genus Cleistocalyx nervosus

(Lour.) Blume. Unfortunately, Kostermans’ name has, incorrectly, been

taken up by several workers (Turner 1995, Chantaranothai & Parnell 2002,

Kress et al. 2003). As this is a wide-ranging (India to Australia) and quite

common species, it is important to clarify and correct its nomenclature.

While Merrill and Perry intended to transfer Eugenia operculata Roxb. to

Cleistocalyx, their combination, Cleistocalyx operculatus Merr. & L.M.Perry,

can be considered as a nomen novum for Syzygium nervosum DC. as they

cited this name in the synonymy. However, there are a number of synonyms

of Syzygium nervosum DC. that have priority over Cleistocalyx operculatus

Merr. & L.M.Perry. The oldest available appears to be Eugenia cerasoides

Roxb. and this is formally transferred to Cleistocalyx below. The correct

name for the species in Syzygium remains as Syzygium nervosum DC. For

those who maintain Eugenia in the widest sense, the species is correctly

Eugenia cerasoides Roxb.

A new combination is also made for the variety of the species

based on Eugenia paniala Roxb. Valid combinations for variety paniala

have yet to be made in either Syzygium or Eugenia.

Cleistocalyx cerasoides (Roxb.) I.M. Turner, comb. nov.

Basionym: Eugenia cerasoides Roxb., Fl. Ind. ed. 1832, 2 (1832) 488.

Syzygium cerasoides (Roxb.) Raizada, Ind. Forester 84 (1958) 478.

Type: Ic. Roxb. no. 2256 (lecto, designated here, K) (reproduced as Wight

Ie;615),

Syzygium nervosum DC., Prod. 3 (1828) 260. Cleistocalyx nervosus (DC.)

Kosterm., Bull. Bot. Surv. India 29 (1989) 17 sphalm. nervosum, comb.

illegit., non Cleistocalyx nervosus (Lour.) Blume, Mus. Bot. 1 (1850) 85.

Cleistocalyx operculatus Merr. & L.M. Perry, J. Arn. Arb. 18 (1937) 337,

nom. nov. pro 8. nervosum DC.

Type: Roxburgh s.n., herb. Lambert 1816 (holo, G-DC).

Eugenia operculata Roxb., Fl. Ind. ed. 1832, 2 (1832) 486, nom. illegit.

Syzygium operculatum Nied., Nat. Pflanzenfam. 3(7) (1893) 85, nom. illegit.

Type: Roxburgh s.n., herb. Lambert 1816 (lecto, designated by Panigrahi

& Mishra 1985, G-DC).

var. paniala (Roxb.) I.M. Turner, comb. nov.

Basionym: Eugenia paniala Roxb., Fl. Ind. ed. 1832 (1832) 489. Eugenia

operculata Roxb. var. paniala (Roxb.) Duthie, Fl. British India 2 (1879)

498, comb. illegit. Cleistocalyx operculatus Merr. & L.M. Perry var. paniala

(Roxb.) P. Chantaranothai & J. Parn., Kew Bull. 48 (1993) 591. Cleistocalyx

nervosus (DC.) Kosterm. [non (Lour.) Blume] var. paniala (Roxb.) P.

A Correction in Cleistocalyx 2a

Chantaranothai & J. Parn., Novon 6 (1996) 201, comb. illegit.

Type: Ic. Roxb. no. 2255 (lecto, designated by Chantaranothai & Parnell

(1993), K).

Acknowledgements

Dr Martin Cheek is thanked for assistance at the Herbarium, Royal Botanic

Gardens, Kew.

References

Chantaranothai, P. and J. Parnell 1993. New taxa and combinations in

Cleistocalyx and Syzygium (Myrtaceae) in Thailand. Kew Bulletin. 48:

589—610.

Chantaranothai, P. and J. Parnell 2002. Myrtaceae. Flora of Thailand. 7(4):

778-914.

Kostermans, A. J. G. H. '1987' (1989). Some taxonomic and nomenclatural

notes. Bulletin of the Botanical Survey of India. 29: 17—18.

Kress, W. J., R. A. DeFilipps, E. Farr and Yin Yin Kyi 2003. A checklist of

the trees, shrubs, herbs, and climbers of Myanmar. Contributions from

the United States National Herbarium. 45: 1—590.

Merrill, E. D. and L. M. Perry 1937. Reinstatement and revision of

Cleistocalyx Blume (including Acicalyptus A. Gray), a valid genus of the

Myrtaceae. Journal of the Arnold Arboretum. 18: 322-343.

Panigrahi, G. and S. C. Mishra 1985. Syzygium nervosum DC., correct

name for S.operculatum Niedenzu (Myrtaceae). Taxon. 34: 298—299.

Turner, I. M. ‘1995’ (1997). A catalogue of the vascular plants of Malaya.

Gardens’ Bulletin, Singapore. 47: 1—757.

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Gardens’ Bulletin Singapore 57 (2005) 29—35 29

Scaphochlamys calcicola (Zingiberaceae): a

New and Unusual Species from Borneo

A.D. POULSEN

Royal Botanic Garden, 20A Inverleith Row, Edinburgh EH3 5LR,

Scotland, U.K.

AND

R.J. SEARLE

48 Bartons Drive, Yateley, Hampshire, GU46 6DP, UK.

Abstract

Scaphochlamys calcicola A.D.Poulsen & R.J.Searle is described from the Bau limestone

area in Sarawak, Borneo. Its placement within Scaphochlamys and relationship with

Boesenbergia and Distichochlamys are discussed.

Introduction

The genus Scaphochlamys Baker comprises 27 described species distributed

from the southernmost provinces of Thailand, through Peninsular Malaysia,

Singapore, and the island of Borneo. The most comprehensive review of

the genus is by Holttum (1950) as part of a treatment of the Zingiberaceae

in Peninsular Malaysia, which includes 20 species of Scaphochlamys. The

four species from Borneo were reviewed by Smith (1987), and since then

three new species have been described from Thailand (Sirirugsa & Larsen,

1991; Jenjittikul & Larsen, 2002).

The current species was discovered during a botanical inventory of

the Bau limestone area conducted by staff of the Sarawak Biodiversity

Center in 2001—2002. This project subsequently involved the first author in

assisting with the identification of Zingiberaceae material and compiling a

preliminary checklist of all species. This list (Poulsen et al., 2004) included

a species of Scaphochlamys which follow-up fieldwork, including the first

author in 2002—2003, has confirmed to be new. During this fieldwork,

several flowering individuals were found, photographed and preserved in

spirit for further studies essential for making the following description

which forms part of a revision of the genus due for publication by the

second author.

30 Gard. Bull. Singapore 57 (2005)

Scaphochlamys calcicola A.D.Poulsen & R.J.Searle, sp. nov.

Scaphochlamydi sylvestri, S. occulatae et S. breviscapaea similis. Ab eis

inflorescentiis compactis spiraliter dispositis differt.

Typus: Malaysia, Borneo, Sarawak, Kuching Division, Bau area, Gunung

Tai Ton (1°24’N; 110°8’E), altitude 50 m, in a recently deforested limestone

gorge, 20 June 2003. Poulsen, Jugah & Clausager 2022 (holo SAR; iso

AAU, E, L ,K).

Terrestrial herb 30-60 cm tall. Rhizome horizontally creeping above the

ground, robust, about 7 mm diam. when dried, sometimes forking, with

well-developed anchoring roots (not stilted). Leafy shoots 2—20 cm apart,

distinctly unifoliate; Jaminate leaf tightly enclosed by 3(—5) leafless sheaths,

the longest to 18 cm, cream at base, flushed green towards apex, becoming

brown and papery with age, shredding; these completely hide the much

shorter sheath and ligule of the leaf; sheath to 3 cm, margin thin and

densely ciliate terminating in a + obscure, bilobed, to 5 mm (when fresh),

membranous ligule; petiole (measured from the base of the shoot because

the ligule is often hard to establish)10—27(—39)cm long, canaliculate; lamina

15—37 x 9-18 cm, broadly elliptic to lanceolate, slightly asymmetric, plicate,

upper surface plain green, lower surface very pale green and villose (up to

1 mm long, appressed, white hairs, scattered throughout the lower surface

but most dense near and on the sides of the midrib); base rounded to

slightly cordate and attenuate; apex distinctly acuminate to c. 1 cm.

Peduncle to 1.5 cm, slender, glabrous, hidden at base of the leaf;

inflorescence derived from near the leaf base inside the leafless sheaths,

flowering from base to apex; inflorescence head 7—10 cm long, bilaterally

flattened: 1.5—2:0 wide and 0.5—0.6 cm deep, tapering towards a pointed

apex, with 8—13 bracts arranged distichously and 0.3—0.7 cm apart on an

elongated spike; bracts pale green, 2.5—3.2 cm long and 0.9 cm wide near

the base, held stiffly upright, boat-shaped, outer surface glabrous or covered

in short spiky hairs; apex acute, edges involute and overlapping; bracts

subtending (2—)3 flowers; bracteole 2.0—3.3 cm (first; second and third

decreasing in size), 2-keeled, generally longer and opposite to the bract,

wrapped tightly around the flower(s), slit to the base, edges overlapping,

covered in short spiky hairs; calyx with ovary 12—13 mm long, fissure c. 4

mm, with rounded to acute, irregularly tridentate apex with a scattering of

short spiky hairs, otherwise glabrous; corolla tube 2.4—4 cm long, glabrous,

corolla lobes white, linear, 1.5 x 0.5 cm, apex mucronate and hooded;

staminodes oblong, 1.2 x 0.3—0.4 cm wide, papillose, apex obtuse; /abellum

1.4-1.7 x 0.9-1.1 cm, spathulate, apex bilobed (indented 3—4 mm), lobes

overlapping, white with pale yellow-green centre; stamen 11—12.mm;

filament 5—7 x 2 mm, anther thecae 4—5 x 1.5 mm, not spurred, dehiscing

Scaphochlamys calcicola from Borneo

’

Figure 1. Scaphochlamys calcicola A.D. Poulsen & R.J. Searle. A. Habit, leaf and inflorescence

with the sheaths removed; B & C. Flowers (manipulated). Bract and bracteole removed on

flower to the right exposing calyx and second bracteole; D. Ovary and calyx (corolla tube

removed); E. Ovary with epigynal gland and base of style; F Stamen, style and stigma, ventral

view; G. Stamen, style and stigma, side view. Drawn by A. D. Poulsen from the type.

32 Gard. Bull. Singapore 57 (2005)

Table 1. A comparison of key characters between the new species,

Scaphochlamys, Distichochlamys and Boesenbergia. Traditionally six

characters have been used (Smith, 1987; Newman, 1995) to which we have

added the splitting of the bracteole.

Scaphochlamys | Scaphochlamys | Distichochlamys | Boesenbergia

calcicola |

Bract spiral distichous distichous distichous

arrangement

Flowering base to apex base to apex base to apex apex to base

mode

Flower cincinni cincinni of 2—3 | cincinni single

arrangement flowers

First bracteole more or less 2-keeled, 2-keeled, boat-shaped,

(shape and keeled, arising | opposite to opposite to arising at right

position) opposite to bract| bract bract, tubular angle to bract

Bracteole split to base split to base Rt tubular, split 2/3 | split to base

splitting

Labellum bilobed or bilobed, flat, bilobed, not usually saccate,

entire, lobes lobes saccate lobes entire, rarely

overlapping overlapping not overlapping | emarginate,

margin recurved

Base of thecae | with very short | no spurs no spurs no spurs

free basal spurs

for their entire length; crest rounded, to 2 mm; stigma 1 mm across, club-

shaped with two dorsal knobs, ostiole ciliate, forward-facing; ovary 2 mm,

glabrous, epigynous glands 4—5.5 mm, bilobed, needle-shaped. Fruit not

seen.

Distribution: Endemic to Borneo in lowland forest. So far, only documented

from near Bau, southwest Sarawak.

Ecology and etymology: From information on labels, this species seems to

be found exclusively on limestone, which is the reason for the choice of

epithet. The species occurs at the base or shoulder of limestone hills, on

boulders near streams.

Notes: The 2-keeled bracteole is an unusual character for Scaphochlamys.

This, however, is characteristic for the genus Distichochlamys M.F.Newman,

a genus recently described from Vietnam with three species (Newman,

1995; Larsen & Newman, 2001; Rehse & Kress, 2003). Comparing the new

taxon to the most closely related genera (Table 1), one could be tempted

to think it is actually a species of Distichochlamys. However, it does not

have the distinctly tubular bracteole of Distichochlamys.

Scaphochlamys calcicola from Borneo 33

Figure 2. Scaphochlamys calcicola A.D. Poulsen & R.J. Searle. Photographs of Connie Geri

et al. SBC 3849; same population as the type. Photos: A. D. Poulsen.

Scaphochlamys calcicola also shares the distichous inflorescence

and lack of spurs with Boesenbergia Kuntze, from which it differs in four

other key characters (Table 1).

Preliminary findings from cladistic analysis of the Internal

Transcribed Spacer (ITS) region of the nuclear NDA (unpublished data)

support the inclusion of the new species within the genus Scaphochlamys,

and it appears to fall within a subclade consisting of other Bornean species

of this genus.

In Sarawak, the material from Bau at first seemed similar to another

undescribed limestone species in southwest Sarawak, which Smith (1987)

mentioned in her account as S. sp. aff. breviscapa Holttum (Burtt 8798; E},

K!). Both species have shoots with a single, erect, long-petiolate (29 cm

long), large leaf (24 x 12 cm). However, the Burtt collection differs by the

34 Gard. Bull. Singapore 57 (2005)

leaf being ovate with a bluntly pointed apex, the peduncle being longer (c.

5 cm) and the spike being compact with spirally arranged bracts. Also, the

labellum has purple around the central line of the labellum (similar to at

least two other Sarawakean species — S. petiolata (K. Schum.) R.M. Sm.

and S. reticosa (Ridl.) R.M.Sm.). The sterile characters of the Burtt

collection bear more similarity to S. oculata (Ridl.) Holttum than to S.

breviscapa.

Of the other species of Scaphochlamys mentioned for Sarawak by

Smith (1987), none has as large leaves as S. calcicola; the most similar

being S. petiolata in which the lamina is up to 14 x 4 cm — much smaller

than S. calcicola. Also, the inflorescence of S. petiolata is much reduced in

comparison. A recent collection, P.Boyce, J.Kisai & S.Kutuh ZI-658 (E,

SAR) from near Serian also has a distichous inflorescence like those of S.

calcicola but differs in the much smaller, cordate leaves, and the smaller

flowers with a lilac labellum. The collection probably represents another

new species and indicates that a distichous inflorescence in Scaphochlamys

is not unique to S. calcicola. This collection and others recently made by

Boyce illustrate that several new species are likely to be discovered and

described from Sarawak in the future.

Considering the unifoliate species in Peninsular Malaysia, the leaf

of S. calcicola is similar in size and shape to S. sylvestris (Ridl.) Holttum, S.

oculata and S. breviscapa. All of these differ by having compact, spirally

arranged inflorescences. In addition, S. sylvestris has a much longer peduncle

(15 cm); S. oculata has a slightly longer peduncle (2—5 cm), whereas that of

S. calcicola is within the range of S. breviscapa (1—3.5 cm).

The collection from the type locality shows some variation within

the population, e.g. length of corolla and anther crest but the flowers are

consistently white with a yellow-green centre. It seems that the colour of

the labellum in S. kunstleri (Baker) Holttum in Peninsular Malaysia can be

quite variable from almost pure white to dark red (Lim, 2001). Future

studies on Bornean Scaphochlamys should investigate if species with a

pale lilac labellum are consistently so or whether they can occasionally be

white like S. calcicola. Clarification of this issue will be of importance for

using this diagnostic character in species identification.

Other specimens examined: SARAWAK. Kuching Division, Bau area:

Gunung Apin, 6 Nov 2001 Malcom Demies et al. SBC 1123 (SAR); Gunung

Krian, 27 Nov 2001 Meekiong Kalu et al. SBC 1613 (SAR); Gunung

Lanyang, 10 Apr 2002 Julia Sang SBC 2919 (SAR); Gunung Poing, 23 Sep

2001 Julia Sang et al. SBC 343 (SAR), 13 May 2002 Malcom Demies et al.

SBC 1547 (SAR), 11 July 2002 Meekiong Kalu et al. 3246 (SAR); Gunung

Tai Ton, 27 Mar 2002 Julia Sang et al. SBC 2654 (SAR), 30 Oct 2002

Connie Geri et al. SBC 3749 (SAR, SING).

Scaphochlamys calcicola from Borneo 35

Acknowledgements

The Sarawak Biodiversity Centre (SBC) requested the first author to assist

in the study of the Zingiberaceae from the Bau limestone area. SBC also

provided permits for his research on Zingiberales elsewhere in Sarawak

and his fieldwork was kindly supported by HRH Crown Prince Frederik’s

Foundation. We should also like to thank staff at SBC and the Forestry

Research Centre for logistic and other help, the keepers of the herbaria

SAR, K, and AAU, and J. F. Veldkamp for Latinizing the diagnosis. Finally,

we should like to thank two anonymous reviewers for constructive

comments on the manuscript.

References

Holttum, R.E. 1950. The Zingiberaceae of the Malay Peninsula. Gardens’

Bulletin Singapore. 13: 1—249.

Jenjittikul, T. and K. Larsen. 2002. Two new species of Scaphochlamys

(Zingiberaceae) from Thailand. Nordic Journal of Botany. 22: 35-38.

Larsen, K, and F.M. Newman. 2001. A new species of Distichochlamys

from Vietnam and some observations on generic limits in Hedychieae

(Zingiberaceae). Natural History Bulletin of the Siam Society. 49: 77—80.

Lim, C.K. 2001. Notes on Scaphochlamys kunstleri (Zingiberaceae) and its

varieties. Folia Malaysiana 2: 117-126.

Newman, F.M. 1995. Distichochlamys, a new genus from Vietnam.

Edinburgh Journal of Botany. 52: 65—69.

Poulsen, A.D., C. Geri, M. Kalu, and M. Roos. 2004. Zingiberaceae and

Marantaceae. In: H.S. Yong, F.S.P. Ng & E.E.L. Yen, (eds). Sarawak

Bau Limestone Biodiversity. The Sarawak Museum Journal. 54 (No. 80

(New Series), Special Issue 6): 163-169.

Rehse, T. & W.J. Kress. 2003. Distichochlamys rubrostriata (Zingiberaceae),

a new species from north Vietnam. Brittonia. 55: 205—208.

Sirirugsa, P. and K. Larsen. 1991. A new species of Scaphochlamys

(Zingiberaceae) from Thailand. Nordic Journal of Botany. 11: 93—95.

Smith, R.M. 1987. A review of Bornean Zingiberaceae: 3 (Hedychieae).

Notes Royal Botanic Garden Edinburgh. 44: 203-232.

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Gardens’ Bulletin Singapore 57 (2005) 37—46 ay

The Recircumscription of Curcuma L. to

Include the Genus Paracautleya R.M.Sm.

J. SKORNICKOVA"2 AND M. SABU!

‘Department of Botany, University of Calicut, 673635 Kerala, India.

"Department of Botany, Charles University, Bendatska 2, Prague, 12802, Czech

Republic.

Abstract

Based on recent living material from the type locality, the monotypic genus Paracautleya is

reduced to synonymy with Curcuma and a new combination is made for C. bhatii (R.M.Sm.)

Skornick. and M.Sabu. A detailed description of the species, including a colour plate, is

given.

Introduction

The monotypic and narrowly endemic genus Paracautleya was established

by R.M. Smith in 1977. Its type species, Paracautleya bhatii R.M.Sm., was

described based on a collection by K.G. Bhat made near Udupi, South

India. In relation to the description, Smith discussed possible affinities of

Paracautleya with four other genera (Camptandra Ridl., Cautleya Hook.f,

Curcuma L. and Roscoea Sm.) in the tribe Hedychieae, which all have a

truly versatile anther like P. bhatii. Several characters were used to clarify

the affinity of Paracautleya but unfortunately, many of them were not

known satisfactory at the time (Table 1), such as the habit and position of

inflorescence, shape of the fertile bract, number of flowers per bract, absence

of bracteole, and ovary division.

Kress et al. (2002) provided a new classification of the Zingiberaceae

based on DNA sequences of the nuclear internal transcribed spacer (ITS)

and plastid matK region. However, they lacked samples of a number of

rare monospecific genera, Paracautleya among them. Based on morphology

alone, they tentatively placed the genus Paracautleya in the Zingibereae

tribe. This was recently confirmed by Ngamriabsakul et al. (2004), who

elaborated the phylogeny of the tribe Zingibereae using the same nuclear

gene as Kress et al. (2002) but a different chloroplast gene, trnL-F. Their

phylogenetic analysis of the combined data set showed that Paracautleya

grouped with Curcuma subg. Curcuma whereas the other genera mentioned

by Smith (1997) were placed in distant clades. Ngamriabsakul et al. (2004)

pointed out that all genera recognized in the Curcuma complex may be

Gard. Bull. Singapore 57 (2005)

Table 1. A re-examination of characters in Curcuma and Paracautleya.

Characters and character states stressed by Smith (1997) in bold.

absent

white, laciniate

i _

Cautleya Curcuma Paracautleya Roscoea

Rhizome Swollen fleshy roots | Rhizome and tubers} Rhizome and tubers} Swollen fleshy roots

Habit Leafy stems Leaf tuft (forming | Leaf tuft (stemless) | Leafy stems

pseudostem or

stemless)

Leaf sheath Closed Open Open Closed

Inflorescence Terminal Terminal or radical | Terminal Terminal

or both

Bract Free Pouch-like or free | Free or pouch-like | Free

Bracteole Absent Present or absent | Present or absent | Absent

Flowers Single Cincinni or single | Single Single

Anther Spurred, L-shaped, | Spurred or not, Spurred, vestige of | Spurred, L-shaped,

crested crested or a vestige | crest present crested

present or absent

—|

Epigynous gland Sharp Blunt Blunt Sharp

apex

Ovary Trilocular Trilocular, rarely Imperfectly Trilocular

imperfectly trilocular

trilocular

Placentation Axile, ovules Axile, many ovules | Basal Axile, ovules

throughout the axis | towards the base, throughout the axis

less towards the top

2 a —4

Capsule Many-seeded. Few to many- Few-seeded Many-seeded

seeded (10 — 25 (5-12 seeds)

seeds)

Seed Black to brown, Ovoid, light brown | Ovoid, light brown, | Ovoid, arillate

grey or red, angled, | to dark brown, shiny, arillate, aril

aril present or shiny, arillate, aril white, laciniate,

usually positioned

towards one side

—|

regarded as a single genus, though there are some morphological characters

supporting the separation of each taxon. These characters they suggest

are, however, autapomorphic. Distinguishing characters of the genera in

the Curcuma complex, namely Curcuma and Paracautleya, as tabulated by

Ngamriabsakul et al. (2004) included the following: in Curcuma adnate

bracts, several flowers per bracts, bracteoles present, and labellum

emarginate and rarely split versus in Paracautleya free bracts, one flower

per bract, bracteoles absent and labellum split.

Curcuma bhatii from India 39

The purpose of the present paper is to evaluate whether the

morphological evidence based on newly available material still justifies

keeping Paracautleya separate from Curcuma.

Material and Methods

In June 2004, Paracautleya bhatii was observed flowering in its type locality.

Plants were collected and transplanted to Calicut University Botanic Garden

for further observation. During the past four years we have also extensively

studied Curcuma species throughout the Indian subcontinent both from

living flowering material as well as herbarium specimens including type

material (BM, BSI, CAL, CALI, E, G, K, L, MH, PDA, PR, SING) paying

particular attention to the extreme variability of vegetative and floral

characters.

Paracautleya R.M.Sm. versus Curcuma L.

Rhizome structure

The genus Curcuma has a conspicuous rhizome structure. The main rhizome

can be simple or branched and the roots end in round, ovoid root tubers,

which may be almost sessile or distant 20-30 cm from the main rhizome or

branches. Root tubers are present invariably in both seed-setting and non

seed-setting species. They are not capable of sprouting and their exclusive

function is to sustain the plant during dry periods when the leafy shoots

dries up. Our recent studies reveal that Paracautleya bhatii has an ovoid

main rhizome usually without branches or with one branch and root tubers,

typical for the genus Curcuma.

Bract

As described by Smith (1977), unlike Paracautleya with singly borne flowers

and bracts that do not form pouches, the flowers of Curcuma commonly

arise in cincinni held within pouches formed by adnate bracts. This

traditional diagnostic character of the genus Curcuma is based mainly on

early descriptions made by Roxburgh (1820) and subsequent workers when

only a few species of Curcuma were known. However, species lacking this

bract formation were described from Sri Lanka (C. albiflora Thwaites, and

to some extent also C. oligantha Trim.). Kress et al. (2002) have pointed

out that pouched inflorescences are neither unique nor universal in the

genus Curcuma.

As a matter of terminology, the term ‘adnate’ is used when two

non-homologous entities are fused, while the term ‘connate’ is appropriate

when two homologous entities are fused, e.g. two bracts. Thus, the pouches

40 Gard. Bull. Singapore 57 (2005)

in Curcuma are, in most species, formed by both means on the same

inflorescence. It is possible to say that pouches in all Curcuma species are

formed at their base by being adnate to the inflorescence axis and in most

species their sides further up are connate to the basal part of the bracts

positioned above them. The degree of connation varies considerably within

the genus and may in some species be negligible.

Smith (1977) mentioned that in Curcuma albiflora a few free bracts

may occur at the base of the inflorescence. We have studied C. albiflora

from herbarium material (BM, E, G, K, PDA). For most sheets, including

type specimens, there is no connation of bracts at all; the bracts form

shallow pouches by the basal part being adnate only to the inflorescence

axis. Bracts of C. albiflora are rarely connate to each other and if so, then

only in the uppermost part of inflorescence. Connation of bracts is also

very much reduced in the case of C. oligantha, where the situation is less

obvious than in C. albiflora. In C. oligantha, one or two free fertile bracts

may occur in the basal part of the inflorescence, where the lowest bracts

may be joined only to the inflorescence axis and the upper bracts are

connate just in its lowermost, c. 1-6 mm, portion.

During our fieldwork, we have examined over 30 inflorescences

from different populations from the type locality and found that bracts in

Paracautleya bhatii are mostly free but do form pouches by being adnate

at its base to the inflorescence axis. Such a pouch can be almost a third of

the bract length, but individuals with their bracts connate for 1—2 mm can

also be found and if such are present, they are usually positioned in the

upper part of the inflorescence. There is no doubt about the generic identity

of C. albiflora and C. oligantha and therefore the pouched bract character

alone is not sufficient to maintain Paracautleya bhatii separate from

Curcuma. |

Number of flowers per bract

In the genus Curcuma, there are usually two or more flowers subtended by

each bract forming a cincinnus while Paracautleya has its flowers borne

singly. Yet, there is in India a complex of four species, namely C. reclinata

Roxb., C. decipiens Dalzell, C. inodora Blatt. and C. sulcata Haines, which

share in common strongly reduced or missing bracteoles and flowers that

are born singly or with a maximum of two per bract.

We have re-collected all these species from their type or near to their

type localities. Curcuma decipiens (Skornickova 73443 and 73445; CAL,

CALI, K, MH, PR, SING) has 1—2 flowers per bract and the number of

flowers may not be consistent throughout the spike. In some plants every

bract subtends just one flower, in others bracts subtend two flowers in the

basal part of inflorescence and one flower in its upper part. C. sulcata

Curcuma bhatii from India 41

(Skornickova 73467, CALI, MH, K, PR, SING), C. reclinata (Skornickova

73477, CAL, CALI, MH, K, PR, SING) and C. inodora 73403 (Skornickova

73403, CALI, SING) have only one flower per bract and we did not

encounter a single plant that had two or more flowers per bract.

Bracteole

Paracautleya has been distinguished from Curcuma on the grounds of its

lacking bracteoles (Smith, 1977). Based on earlier generic and specific

descriptions of Curcuma, there is one boat-shaped bracteole per flower.

From our field observations, there are great differences in bracteole size

across the genus Curcuma. The non-seed setting species usually possess

large bracteoles that can reach over 3 cm long while the bracteoles of some

seed-setting species can either be well developed (e.g., C. montana Roxb.)

or more or less reduced. As already mentioned above, C. decipiens, C.

inodora, C. reclinata and C. sulcata bracteoles are reduced to 1-2 mm or

are often absent. This feature may not be constant within the species or

whole population (e.g., the same population includes plants with strongly

reduced bracteoles as well as plants where the bracteoles are completely

lacking), but it is usually constant within a single plant. In the type locality

of P. bhatii, we found plants without bracteoles (more common) as well as

plants with bracteoles up to 2.5 mm long.

Labellum

An emarginate labellum is commonly found in a number of Indian seed-

setting species distributed in Western Ghats area, e.g. C. oligantha, C.

mutabilis Skornick. et al., C. pseudomontana J.Graham, where the split

often progresses as the flower ages towards the end of the day and wilts.

This is thus not unique to Paracautleya contrary to what was tabulated by

Ngamriabsakul et al. (2004).

Anther structure

Anther structure of Paracautleya is similar in structure to several seed-

setting Curcuma species being versatile, having two anther spurs at its base

and, in the adaxial part of the anther, a vestige of a reduced anther crest.

The whole morphology of the anther closely resembles, for example, anthers

of C. neilgherrensis Wight, C. decipiens or C. oligantha.

Ovary structure

As far as we know, there is no specific work dealing in detail with ovary

structure in the genus Curcuma. The ovary in Zingiberaceae is tricarpellate

(except the tribe Globbae and genus Tamijia, in which the ovary is unilocular

with parietal placentation), and is generally described as trilocular with

42 Gard. Bull. Singapore 57 (2005)

axile placentation. Hamza (1989) observed that many Zingiberaceae species

described with trilocular ovaries and axile placentation, have incomplete

septation at the top and are thus unilocular in this region. He did not

include any representatives of the genus Curcuma in his study, but we

have seen similar ovaries when observing transverse sections from different

parts of ovaries of different species of Curcuma e.g., C. decipiens, C.

oligantha, C. neilgherrensis, C. karnatakensis Amalraj et al. Sections near

the base are always trilocular with a large number of ovules, but sections

towards the top have a much reduced ovule number or are empty and the

septa are not always joined. In Paracautleya incomplete septa are clearly

visible.

Aril and seed shape

For the genus Curcuma, there are few descriptions or drawings of fruits

and seeds but we have been able to observe fruits and seeds of more than

15 species. Seeds are ovoid, light brown to dark brown and shiny. The aril

is white, laciniate and is rather uniform throughout the genus, although the

size of the aril may vary among the species. The only Curcuma species

with seeds described as exarillate appears in the original description of C.

oligantha (Trimen, 1885). We have examined the type specimen of C.

oligantha (PDA) and confirmed the presence of an aril on the seeds from

an unopened fruit. Seeds of Paracautleya bhatii agree in shape and colour

and the presence of a laciniate aril with those observed in the genus

Curcuma. The laciniate aril in Paracautleya is usually positioned towards

- one side and its size and shape vary slightly within a population. The aril in

unripe seeds is smaller than in fully ripe seeds.

Other observations

The inflorescence in Paracautleya bhatii is invariably terminal on a long

naked peduncle and its leaves are rather narrow. In its general habit it

most resembles some of the SE Asian Curcuma species, e.g. C. gracillima

Gagnep., which also has this type of inflorescence and narrow leaves. In

the Indian region, the most similar species are perhaps C. oligantha (India

and Sri Lanka) and C. albiflora (Sri Lanka).

While dealing with the ovary and fruit of Paracautleya bhatii, we

also studied the shape of the epigynous glands. They agree well with those

found in the genus Curcuma in being linear with a blunt apex. The shape

of the stigma and presence of cilia at the ostiole of Paracautleya bhati also

agrees well with Curcuma.

Curcuma bhatii from India 43

Conclusion

The conclusion is that the generic delimitation of Paracautleya as defined

by Smith (1977) is not strong enough to warrant keeping it as a separate

genus. As well as similarity in habit, we found that the generic diagnostic

characters including floral morphology described for Paracautleya can be

observed in at least some members of the genus Curcuma. Thus, its generic

status cannot be justified. This necessitates a new combination for the

Paracautleya species.

Curcuma bhatii (R.M.Sm.) Skornick. and M.Sabu comb. nov.

Basionym: Paracautleya bhatii R.M.Sm., Notes Roy. Bot. Gard. Edinburgh.

35 (1977) 367; K.G. Bhat, Fl, Udupi. (2003) 634.

Type: India, Karnataka, South Kanara, Manipal, 1.VI. 1975 Bhat 204 (holo

E!; iso BSI!, C, CAL!).

Small rhizomatous herb, 5—20 cm tall. Rhizome ovoid mostly unbranched,

5—10 x 4-5 mm, rarely with one branch, which gives rise to a new plant,

light brown externally, sheathed by papery remains of leaf sheath bases,

creamy yellowish internally, non-aromatic. Roots fleshy, root tubers up to

1—2.5 x 0.5 cm, almost sessile to the main rhizome, externally whitish (when

young) to brown (when older), internally pure white, non-aromatic.

Pseudostem 2—5 cm long, formed by leaf sheaths and 1—2 sheathing bracts,

whitish, drying soon and becoming light brown, thin and papery, ligule 1.5

mm, bilobed, translucent greenish-white, glabrous. Leaves 2—5 (—7), sessile

or with a very short petiole c. 1 cm long (gradually changing into a narrowly

attenuate lamina base); /amina lanceolate, 5-12 x 0.7—1.5 cm, adaxially

green, glabrous, abaxially lighter green, glabrous; margin hyaline, translucent

white, c. 0.1 mm wide, tip acute, base attenuate, midrib green, glabrous.

Inflorescence invariably central. Peduncle 3.5—13 cm, c. 1—1.5 mm diam.,

green, glabrous, partly hidden within the pseudostem. Spike 2—5 x 1-—2.5

cm, consisting of 5—23 green bracts. Coma inconspicuous, usually only the

uppermost 2-3 bracts are sterile and smaller in size than the fertile ones.

Fertile bracts ovate, 1—1.4 x 0.7—1 cm, tip acute, both sides glabrous, green,

usually free especially at the base of inflorescence, but occasionally connate

by the lower fifth of bracts (c. 1.5—2.5 mm), 1 flower per bract. Bracteoles

usually absent (reduced), rarely present, one per flower, 0.5—2.5 x 0.5—1

mm, translucent white, glabrous. Flowers 1.8—2 cm long, exserted from

bracts. Calyx 3.5 mm long, obscurely 3-toothed, translucent white with

greenish tinge, glabrous. Corolla tube c. 9 mm, deep yellow, glabrous;

dorsal corolla lobe c. 6—7 x 7 mm, triangular-ovate, concave, apex with

obscure blunt mucro 0.2 mm, translucent yellow, glabrous, lateral corolla

44 Gard. Bull. Singapore 57 (2005)

lobes ovate, slightly concave at tip, translucent yellow, glabrous. Lateral

staminodes c. 7 x 4.5 mm, deep yellow, glandular hairs present on the

slightly raised middle portion. Labellum c. 9 x 9 mm, emarginate, split 3—4

mm long (opening deeper and wider as the flowers age and wilt), deep

yellow. Anther versatile, spurred, deep yellow, short glandular hairs present

on the sides and adaxially, anther thecae whitish, 1.5 x 0.4 mm; filament 1

mm long, deep yellow, constricted, 2.5 mm broad at base, 1 mm broad at

upper part. Anther spurs 0.9—1 mm long, yellow. Anther crest small, reduced

c. 0.5 x 0.4 mm, yellow. Ovary imperfectly trilocular (trilocular at basal

part, septa incomplete in the upper part), c. 1.8 x 2 mm, white, glabrous,

ovules c. 3—8. Stigma c. 0.7—0.9 x 0.7—0.9 mm, creamy white, ciliate, not

exserted. Epigynous glands 2, creamy yellowish, c. 1—1.2 mm long, 0.2—0.3

mm diam. Fruit a dehiscent capsule, spherical, 5—6 x 5—6 mm light green to

whitish, glabrous, calyx persistent. Seeds 3.5—4 x 2 mm, greenish creamy

(unripe) to light brown (ripe), shiny glabrous, non-aromatic, aril translucent

white, laciniate, lobes up to 4 x 0.5—1 mm (smaller in unripe seeds), arranged

towards one side.

Other specimens examined: India, Karnataka, Udupi Dt., Manipal,

Skornickova: 73446 (CAL, CALI, K, MH, SING, PR); Manipal, Sabu s.n.

(CALI); Alevoor, Bhat 11349 (E!).

Distribution, habitat and phenology: This species is very rare and so far is

known to occur only in Karnataka, Udupi District, at its type locality and

adjacent territory. Curcuma bhatii grows on lateritic slopes in rock crevices

in areas that are rich in monsoon rains. Its small size may be due to its

habitat, which is unique compared with other Curcuma species. It flowers

from June to August. This species is highly endangered due to rapid loss of

its habitat.

It is notable that maximum diversity of seed-setting Curcuma species

is along the western side of the Western Ghats, South India, where, for

example, C. oligantha, C. vamana Sabu et Mangaly; C. karnatakensis

Amalraj et al.; C. pseudomontana Graham, as well as C. bhatii, occur. This

region is known as one of the world’s biodiversity ‘hot spots’.

Plate 1. Curcuma bhatii (R.M.Sm.) Skornick. and M.Sabu

1. Anther (front view); 2. Anther (lateral view); 3. Anther (back view); 4. Inflorescence with

flower in side view (half bract dissected); 5. Whole plant with detail of the flower in front

view; 6. Epigynous glands; 7. Detail of the arillate seed; 8. Rhizome structure — main rhizome

on the right, root tuber on the left; 9. Rhizome structure dissected — main rhizome on the

right, root tuber on the left; 10. Curcuma bhatii on its type locality. es 73446).

Photo J. Skornickova.

Curcuma bhatii from India

46 Gard. Bull. Singapore 57 (2005)

Acknowledgements

The authors thank the Department of Science and Technology, Government

of India (Order No. SP/SO/A-20/99 dt. 09.11.2001), the Grant Agency of

the Academy of Sciences of the Czech Republic (Grant No. B.6407401)

and the International Association of Plant Taxonomy for financial support.

The first author thanks curators of BM, BSI, CAL, CALI, E, G, K, L, MH,

PDA, PR, SING for letting her examine specimens in their care, to

Singapore Botanic Gardens for awarding a Singapore Botanic Gardens

Research Fellowship and providing facilities during her repeated visits, Dr.

Axel Dalberg Poulsen (E), Dr. Mark F. Newman (E) and Dr. Ruth Kiew

(SING) for comments on the manuscript.

References

Baker, J. G. 1890. Scitamineae. Flora of British India. 4:198—264.

Hamza, P.V. 1989. A Study on Placentation in Zingiberaceae. M. Phil.

Thesis, Calicut University, India (Unpublished).

Kress, J. W., L.M. Prince and K.J. Williams. 2002. The phylogeny and a

new classification of the gingers (Zingiberaceae): evidence from molecular

data. American Journal of Botany. 89: 1682—1696.

Ngamriabsakul, C., M.F. Newman and Q.C.B. Cronk. 2004. The phylogeny

of the tribe Zingibereae (Zingiberaceae) based on ITS (nrDNA) and

trnL-F (cpDNA) sequences. Edinburgh Journal of Botany. 60: 483—507.

Roxburgh, W. 1820. Monandria Monogynia. Flora Indica. Mission Press,

Serampore, India. Pp. 1—84.

Smith, R.M. 1977. A new genus of Zingiberaceae from S. India. Notes

from Royal Botanic Garden Edinburgh. 35: 365—368.

Trimen, H. 1885. Notes on the Flora of Ceylon. Journal of Botany. 23:

238—245.

Gardens’ Bulletin Singapore 57 (2005) 47—61 47

New Taxa and Taxonomic Status in

Xanthophyllum Roxb. (Polygalaceae)

from Borneo

W.J.J.O0. DE WILDE AND BRIGITTA E.E. DUYFJES

National Herbarium of the Netherlands, Leiden Branch

P.O. Box 9514, 2300 RA Leiden, The Netherlands

Abstract

Thirteen new taxa or taxa with a new status in Xanthophyllum (Polygalaceae) from Borneo

are described. The ten new species described in this paper are: X. bicolor W.J. de Wilde &

Duyfjes, X. brachystachyum W.J. de Wilde & Duyfjes, X. crassum W.J. de Wilde & Duyfjes,

X. inflatum W.J. de Wilde & Duyfjes, X. ionanthum W.J. de Wilde & Duyfjes, X. longum

W.J. de Wilde & Duyfjes, X. nitidum W.J. de Wilde & Duyfjes, X. pachycarpon W.J. de

Wilde & Duyfjes, X. rectum W.J. de Wilde & Duyfjes and X. rheophilum W.J. de Wilde &

Duyfjes, and the new variety is X. griffithii A.W. Benn var. papillosum W.J. de Wilde &

Duyfjes. New taxonomic status has been accorded to X. adenotus Miq. var. arsatii (C.E.C.

Fisch.) W.J. de Wilde & Duyfjes and X. lineare (Meijden) W.J. de Wilde & Duyfjes.

Introduction

During the study of Xanthophyllum carried out in the BO, KEP, L, SAN,

SAR and SING herbaria for the account of Polygalaceae in the Tree Flora

of Sabah and Sarawak, several new taxa were defined. Their taxonomic

position within the more than 50 species of Xanthophyllum recognised in

Sabah and Sarawak will be clarified in the treatment of the family in the

forthcoming volume of the Tree Flora of Sabah and Sarawak series. In

addition, three new species from Brunei and Kalimantan are also described

below.

1. Xanthophyllum adenotus Miq. var. arsatii (C.E.C. Fisch.) W.J. de Wilde

& Duyfjes, stat. nov.

Basionym: Xanthophyllum arsatii C.E.C. Fisch., Bull. Misc. Inform., Kew

(1932) 176. Type: Arsat BNBFD 1213, Borneo, Sabah, Lokan [Lukan]|

(holo K).

Distribution: Endemic to Borneo (Sabah, Sarawak, Brunei, Kalimantan).

Ecology: In mixed dipterocarp forest, along streams, on flat land or on

ridges, on sandy soil, sandstone, shale or silty clay, at altitudes to 400 m.

48 Gard. Bull. Singapore 57 (2005)

Notes: We recognise two varieties of Xanthophyllum adenotus: var. adenotus

and var. arsatii. The latter differs from the typical variety in its leaves that

have a cuneate or rounded base (not rounded-cordate) with flat (not

upturned) margins on both sides of the petiole.

Selection of specimens examined: SABAH - Kinabatangan District,

Tangkulap Forest Reserve (e.g SAN 124676); Labuk Sugut District, Bidu

Bidu Forest Reserve (e.g., SAN 130719); Pensiangan District, Sogo Sogo

(e.g., SAN 91/17); Ranau District, Kilimu Forest (e.g., SAN 6511/1);

Sandakan District, Kebon China Forest Reserve (e.g., SAN 37456), Sipitang

District (e.g., SAN 133578). SARAWAK - Kapit District, Sungai Mengiong

(e.g., § 29755 and S 41217); Kuching District (e.g., S 20887 and S$ 32692);

Lubok Antu District (e.g., S 33564); Lundu District, Sungai Sebuloh (e.g.,

S 49914 and S 59969), Gunung Pueh (e.g., S 34490); Miri District, Lambir

Hill National Park (e.g., S 46424). BRUNEI - Belait District, Sungai Liang

Arboretum (e.g., Niga Nangkat NN 94), Brunei Town (e.g., Hotta 13182

and Sands 5673); Tutong District (e.g., BRUN 15060 and BRUN 15370).

KALIMANTAN - WEST KALIMANTAN, Bukit Baka National Park

(e.g., Church 159), Gunung Bentuang (e.g., Yahud ITTO/BA 792), Gunung

Senuyah (e.g., Suzuki 9675); CENTRAL KALIMANTAN, Sambas (e.g.,

Jarvie 5668), Sungai Katingan (e.g., Wiriadinata 3566), Tumbong Riang

(e.g., Mogea 3626); EAST KALIMANTAN, Bilatalang (e.g., Kostermans

12707), Djenean (e.g., Endert 4958), Nunukan Island (e.g., Meijer 2424).

2. Xanthophyllum bicolor W.J. de Wilde & Duyfjes, sp. nov.

Iuxta Xanthophyllum penibukanensem Heine, foliis longioribus 14—24 cm

longis, basi in petiolo longo attenuata 20-30 mm longo, fructu 2 cm diam.

distinctum. Typus: Niga Nangkat NN 182, Borneo, Brunei, Belait District,

Sungai Mau (holo KEP; iso BRUN, L, SAN, SAR, SING).

Tree to 30 m tall and 32 cm diam. Twigs grey. Bark dark brown, smooth,

inner bark pale yellow. Axillary buds 1—3, conical, 2—4 mm long, glabrous,

with corky thickenings at base. Leaves glabrous, green and shiny above,

contrastingly pale cinnamon and papillose beneath; blade oblong, 14—35 x

4.5—10 cm, base long-attenuate and forming part of the petiole, apex rounded

to bluntly acute; midrib somewhat raised above; lateral veins 8—12 pairs,

indistinct, basal veins hardly reaching the middle of the leaf, intramarginal

vein faint; intercostal venation coarsely reticulate, faint; glands inconspicuous;

petiole 20-30 mm long, consisting of the basal half grey-brown, transversely

or longitudinally furrowed or wrinkled, and the distal part smooth yellowish-

coloured like the midrib. Inflorescences nearly half as long as the leaves, 6—

11 cm long, unbranched; axis minutely sparsely appressed hairy, 15—25—

Xanthophyllum in Borneo 49

flowered. Flowers: 1—3 per bract, bracts minute; pedicel 5-10 mm long,

appressed fine-hairy or subglabrous; sepals c. 4 mm long, subglabrous;

petals glabrous, yellowish orange when fresh, brown-orange on drying;

ovary ovoid, c. 4 mm diam., sparsely minutely appressed hairy, c. 2 mm

stipitate, style caducous, ovules 8 (?). Fruits globose, 1.5—2 cm diam., 2 mm

stipitate, blue when ripe, drying light brown, glabrous; pericarp thin; fruiting

pedicel c. 10 mm long.

Distribution: Endemic to Borneo (Sabah, Sarawak and Brunei).

Ecology: Mixed dipterocarp, riverbank and hill forest, on peaty or (yellow)

sandy clay soil, at low altitudes.

Notes: Xanthophyllum bicolor belongs to a group of species with 8-16

ovules per ovary and a pale, whitish, papillose leaf lower surface, to which

also belong X. discolor, X. penibukanense and X. pulchrum. It differs from

X. discolor and X. penibukanense, by its long petiole, which is two-coloured

as it actually consists of a brown basal part that appears to be the ‘true’

petiole and a more greenish part which can be regarded as derived from

the base of the leaf blade narrowed to the same width as the ‘true’ petiole.

This phenomenon is less conspicuously seen in X. penibukanense, and

even more so in the unrelated X. Jongum, described below. The lower leaf

surface is of a distinctive pale cinnamon, much contrasting with the dark

green upper surface. Although known only from few specimens, X. bicolor

seems to be confined to forest on poor wet soils or peaty riverbank swamp

forest, whereas X. penibukanense is found mainly in forest on well-drained

soils.

Specimens examined: SABAH — Beaufort District (SAN 43595, SAN 77790

and SAN 86129); Tenom District (SAN 64344); SARAWAK — Lubok

Antu District, Ulu Sungai Engkari (S$ 34723). BRUNEI — Mostly from

Belait District (BRUN 17868, BRUN 17936, Joffre et al. JAA 10, Haslani

Abdullah HA 80, Niga Nankat NN 182 [type] and Niga Nankat NN 369).

3. Xanthophyllum brachystachyum W.J. de Wilde & Duyfijes, sp. nov.

Tuxta Xanthophyllum pedicellatum Meijden, racemis brevioribus paucifloris

c. 5 mm longis, pedicellis 2—4 mm longis differt. Typus: Normaya & Sirukit

S 91521, Borneo, Sarawak, Marudi District, Sungai Silat Basin, Bukit

Palutan (holo SAR; iso KEP, L, SAN, SING).

Tree to 12 m tall and 9 cm diam. Bark smooth, grey-green or whitish, slash

bark yellow, tough. Sapwood yellow, hard. Twigs 0.5—1 mm diam., brown

50 Gard. Bull. Singapore 57 (2005)

patently hairy, hairs c. 0.5 mm long, later glabrescent. Axillary buds less than

1 mm long, hairy. Leaves green, hairy mainly on midrib, papillose beneath;

blade narrowly oblong, 3—7(—9.5) x 0.5-1(—2) cm, base narrowly rounded or

cuneate, apex long-acute; lateral veins 4~7 pairs, forming a faint intramarginal

vein; intercostal venation reticulate; glands numerous, minute, c. 0.1 mm diam.,

scattered; petiole c. 2 mm long, hairy. Inflorescences much shorter than the

leaves, c. 0.5 cm long, unbranched, with 2-4 flowers; axis glabrescent. Flowers:

pedicel 2-4 mm long, subglabrous; sepals sparsely appressed hairy or

subglabrous, outer sepals 1.5—2 mm long, inner sepals c. 2.5 mm long; petals

pale purplish, drying orange brown, (sub)glabrous, c. 12 mm long; filaments

subglabrous, anthers c. 0.5 mm long, with some hairs at base; ovary c. 1 mm

stipitate, densely light brown half-patently hairy, hairs 0.5—1 mm long, style

glabrous in apical part, ovules presumably 8 or more. Fruits globose, 1—1.5

cm diam., pale brown, hairy; fruiting pedicel c. 5 mm long.

Distribution: Endemic to Borneo (Sabah and Sarawak).

Ecology: Hill mixed dipterocarp forest, at altitudes between 500 and 900 m.

Notes: This species resembles Xanthophyllum purpureum Ridl. because its

midrib is densely hairy beneath. X. brachystachyum is easily distinguished

by its strikingly delicate branches (1 mm thick or less) and small leaves, by

which it is reminiscent of X. pedicellatum. The latter, however, differs by

its longer inflorescences with more flowers with longer pedicels, 10—15

mm long (pedicels 2—4 mm long in X. brachystachyum). Both X.

pedicellatum and X. brachystachyum can possibly be regarded as derived

from delicate ecotypic forms, possibly confined to ridge forest on poor

soils, of the very variable X. purpureum.

Specimens examined: SABAH—Beaufort District, Mt. Sunggau (SAN

77425). SARAWAK-Marudi District, Bukit Palutan (S 9/52/ [the type]

and S$ 9/825); Miri District (S 3735).

4. Xanthophyllum crassum W.J. de Wilde & Duyfjes, sp. nov.

Tuxta Xanthophyllum vitellinum (Blume) D. Dietr., foliis crassis coriaceis

nervatura inconspicua, inflorescentiis validissimis eramosis differt. Typus:

Sugau SAN 134307, Borneo, Sabah, Kinabatangan District, Bukit Tawai

(holo SAN).

Figure 1

Shrub or treelet c. 2 m tall and c. 3 cm diam. Bark blackish brown. Twigs

black, glabrous, 4-5 mm thick. Axillary buds long-conical, (2—)4—5 mm

Xanthophyllum in Borneo

Bw mar

Figurel. Xanthophyllum crassum W.J. de Wilde & Duyfjes

A. Flowering leafy twig; B. Part of inflorescence. (from SAN 134307)

52 Gard. Bull. Singapore 57 (2005)

long, glabrous. Leaves thickly coriaceous, glabrous, on drying dark brown

above; concolorous and not papillose beneath; blade (shortly) (ovate-)

elliptic, 6—10(—12) x 6(—7) cm, base broadly rounded, apex rounded with

short, broad, blunt tip; midrib slightly raised above; lateral veins 4—7 pairs,

not forming an intramarginal vein; intercostal venation reticulate, flat and

indistinct beneath; glands few, scattered, less than 0.5 mm diam.; petiole 8—

10 mm long, stout, black on drying, without glands. Inflorescences stout,

erect, (almost) unbranched, finely yellow-brown appressed hairy; axis 5—7

cm long, 3—4 mm thick, 10—15—flowered. Flowers solitary or 2—3 together;

pedicel short, stout, 4-5 mm long; sepals 5—6 mm long, densely appressed

yellow-brown hairy; petals 14-15 mm long, purple-black on drying, partly

grey-yellow appressed hairy; stamens and pistil not seen; ovary subglobose,

c. 2.5 mm diam., densely yellow-brown subpatently hairy, hairs c. 0.5 mm

long, ovules 4. Fruits not seen.

Distribution: Endemic to Sabah, where it is known only from the type

locality in the Bukit Tawai Forest Reserve, Kinabatangan District.

Ecology: Stunted montane forest over ultrabasic substrate, at

c. 1250 mraltitude.

Notes: Xanthophyllum crassum is obviously close to X. vitellinum, which

has membranous leaves and branched inflorescences. It differs, however,

from X. vitellinum in its stout habit, (ovate-)elliptic, very coriaceous leaves

with faint venation, and stout, erect, almost unbranched inflorescences. It

~ is also similar to _X. rectum from lowland kerangas in west Sarawak, which

differs in its prominent and distinct coarsely reticulate leaf venation.

5. Xanthophyllum griffithii A.W. Benn. var. papillosum W.J. de Wilde &

Duyfjes, var. nov.

Tuxta Xanthophyllum griffithii A.W. Benn. var. angustifolium Ng, ramulis

2mm diam., gemmis axillaribus 5-10 mm longis pubescentibus, foliis infra

distincte papillosis differt. Typus: Abang Mohtar S 54289, Borneo, Sarawak,

Lundu District (holo SAR; iso K, KEP, L, MO, SAN). |

Synonym: Xanthophyllum griffithii A.W. Benn. subsp. angustifolium (Ng)

Meijden, Leiden Bot. Ser. 7 (1982) 94, p.p., excluding the collections cited

for Peninsular Malaysia.

Twigs hairy, towards apex 1—2 mm thick. Axillary buds long-triangular or

oblong (-linear), S—9(—10) mm long, hairy. Leaves (ovate or) elliptic-oblong,

S—10(—13) x 2—5 cm, apex acute-acuminate, dull, pale (grey-glaucous) and

Xanthophyllum in Borneo 53

distinctly papillose beneath; petiole 6—12 mm long, hairy. Branches of

inflorescences 1—2 mm thick, densely brown hairy. Fruits densely patently

hairy, sometimes glabrescent; fruiting pedicel c. 3 mm long, densely fine-

hairy.

Distribution: Endemic to Borneo (Sarawak and Kalimantan).

Ecology: In mixed lowland dipterocarp and heath forest, also close to the

sea.

Notes: Two varieties of Xanthophyllum griffithii, var. angustifolium and

var. papillosum, can be found in the same area, e.g. at Gunung Santubong,

Sarawak, where the former seems to grow at somewhat higher altitudes.

The type of X. griffithii var. angustifolium from Peninsular Malaysia (Ng,

1970, 1972) has axillary buds that are somewhat longer (S—7 mm long), the

branches slightly thicker, and the flowers more hairy, as compared to the

Bornean material. Xanthophyllum griffithii var. papillosum comes close to

X. pseudostipulaceum Merr. from Luzon, the Philippines, but the latter

differs in its larger and glabrous axillary buds, and glabrous twigs.

Specimens examined: SARAWAK - Bintulu District (S$ 54289); Kuching

district, Gunung Santubong (S 47097, S 60111, S 91154 and S 91159), Bako

National Park (S 6730, S 7006 and S 54289 |the type], Gunung Matang (S

81803); Lundu District, Samunsam Wildlife Sanctuary (S 40567). EAST

KALIMANTAN, Berau (Ambriansyah et al. 942, Arifin et al. 1108 and

Kessler et al. 651).

6. Xanthophyllum inflatum W.J. de Wilde & Duyfjes, sp. nov.

Xanthophyllo flavescenti Roxb. similis, fructu maiore c. 4 cm diam.,

infructescentia racemosa eramosa differt. Typus: Ambriansyah AA 2772,

Borneo, Central Kalimantan, Barito River, Batampang Village area, 2° 01

2S 114° 38 2 E (holo L; iso WAN).

Small tree to 5 m tall and 7 cm diam. Bark greyish. Axillary buds minute,

less than 1 mm long. Leaves coriaceous, yellow on drying, glabrous, not

papillose beneath; blade oblong, 12—20 x 3—5.5 cm, base cuneate, apex

acute-acuminate; midrib prominent beneath; intercostal venation (in parts

of the leaf) scalariform; petiole 10-15 mm long. Inflorescences

(infructescences) terminal, /ong-racemose, unbranched, 15—20 cm long, 20—

25—flowered (flower-scars). Flowers not known. Fruit developed near the

apex of the infructescence, single, globose, large, 3.5—4 cm diam. (c. 7 cm

diam. and orange when fresh), glabrous; pericarp c. 5 mm thick, somewhat

54 Gard. Bull. Singapore 57 (2005)

wrinkled, (yellow-)brown on drying; sepals not persistent; fruiting pedicel

stout, 4-5 mm long. Seed single, globose, c. 2 cm diam.

Distribution: Endemic to Borneo; confined to Kalimantan where it is known

only from the type.

Ecology: Peat swamp forest, at c. 25 m altitude.

Notes: This species from peat swamp forest is, in its vegetative characters,

closely related to the large, variable species Xanthophyllum flavescens,

which differs in having branched inflorescences and smaller fruits (1—2 cm

diam.) with a thinner pericarp.

7. Xanthophyllum ionanthum W.J. de Wilde & Duyfjes, sp. nov.

Xanthophyllo pedicellato Meijden et X. neglecto Meijden similis, foliis infra

pubescentibus nec papillosis, basi anguste cuneata, inflorescentia brevi c. 0.5

cm longa pauciflora, pedicellis 6—8 mm longis, ovulis 4 differt. Typus: Susanto

& Peters 1177, Borneo, West Kalimantan (holo L; iso BO, NY).

Treelet, stem 3—6 cm diam. Axillary buds long-triangular, c. 3 mm long,

hairy. Leaves green on drying, (sparsely) patently hairy and not papillose

beneath; blade oblong, 8—13 x 2—4 cm, base long-cuneate, apex acute-

acuminate; intercostal venation reticulate; petiole 5—7 mm long, short-hairy.

Inflorescences | or 2 on the node or on the older wood, unbranched, short,

0.5(—1) cm long, pubescent, 3—5-flowered; bracts minute; pedicel 6-8 mm

long, hairy; flowers yellow, sepals hairy; ovary ovoid, stipitate, hairy, ovules

4. Fruits (immature) subglobose, c. 6 x 5 mm, densely hairy; fruiting pedicel

8—10 mm long.

Distribution: Endemic to Borneo: confined to West Kalimantan.

Ecology: Mixed dipterocarp forest, on rocky soil, at 100—300 m altitude.

Notes: The long-pedicelled flowers of the (sometimes paired) short

inflorescences give the impression that the flowers are more or less fascicled.

Xanthophyllum ionanthum resembles X. beccariana or X. pedicellatum in

its similar hairy leaves, but the last two species have a much broader leaf

base, purplish flowers and 8—12 ovules per ovary. It is also superficially

reminiscent of X. neglectum, a largely glabrous species with 4 ovules per

ovary.

Specimens examined: WEST KALIMANTAN (Susanto & Peters 1177 |the

Xanthophyllum in Borneo DS

type], Suzuki K9720 and Suzuki K10071).

8. Xanthophyllum lineare (Meijden) W.J. de Wilde & Duyfjes, stat. nov.

Basionym: Xanthophyllum adenotus Mig. var. lineare Meijden, Leiden Bot.

Ser. 7 (1982) 101; Fl. Males. 1, 10 (1988) 516. Type: Sinanggul SAN 57294,

Borneo, Sabah, Lahad Datu District, Bukit Silam (holo K; iso SAN).

Notes: Additional collections made since its original description as a variety

of Xanthophyllum adenotus indicate that this taxon can best be regarded

as a distinct species. It is possibly derived from the widespread X. adenotus,

but it is different especially in its coriaceous linear leaves. It seems restricted

to an environment with divergent ecology, namely, in stunted forest on

ultrabasic bedrock on Bukit Silam. Some other local endemic species, like

X. crassum, X. petiolatum Meiden and X. rectum, may similarly be regarded

as derived from the widespread_X. vitellinum, induced by special edaphically

defined environments.

Specimens examined: SABAH — Lahad Datau District, Bukit Silam

(Beaman et al. 10030, Rimi Repin et al. SP 6225, SAN 29652, SAN 57294

[the type], SAN 95535, SAN 109836, SAN 144533 and Wood SAN A 4182).

9. Xanthophyllum longum W.J. de Wilde & Duyfjes, sp. nov.

Iuxta Xanthophyllum vitellinum (Blume) D. Dietr., foliis tenuibus

concoloribus petiolis gracilibus 35—40 mm longis differt. Typus: Sigin, Lideh

& Patrick SAN 107165, Borneo, Sabah, Kinabatangan District, Ulu Sungai

Pinangah (holo SAN; iso KEP, L, SAR).

Figure 2

Tree to 5 m tall and 10 cm diam. Bark pale greenish or blackish. Sapwood

white. Axillary buds | (or 2), less than 1 mm long, minutely hairy. Leaves

green on both surfaces, glabrous, except for minutely patently hairy midrib

and petiole, not papillose beneath; blade (narrowly) oblong, 15-20 x 6—7

cm, base rounded to short-attenuate, apex acute-acuminate; midrib raised

above; lateral veins 8—10 pairs, forming an intramarginal vein; intercostal

venation reticulate, glands inconspicuous; petiole (20-)30—40 mm long, 1—2

mm thick, the basal portion of c. 15 mm brown, the rest slightly narrower

and drying as green-yellow as the midrib, glands absent. Inflorescences

about half as long as the leaves, subapical, branched, axes minutely patently

hairy. Flowers (after anthesis) single; pedicel 1-2 mm long; perianth, stamens

and pistil unknown; ovary globose, densely grey(-brown) patently hairy

(hairs c. 0.5 mm long), style caducous, ovules 4. Fruits hairy, brownish,

globose, 1.5—1.7 cm diam.; pericarp thin; fruiting pedicel c. 4 mm long.

56 Gard. Bull. Singapore 57 (2005)

Distribution: Endemic to Borneo (Sabah and Sarawak).

Ecology: Lowland forest, along streams, on undulating land and hillsides.

Notes: Xanthophyllum longum resembles X. vitellinum but differs, in general

aspect, from the latter (and from all other known species in Sabah and

Sarawak), by its thin, bright green leaves and extremely long and slender

petioles, (20—)30—40 mm long (versus 8—14(—16) mm long in X. vitellinum).

The upper two-thirds of the petiole is obviously derived from the leaf

blade as it has the same green-yellow colouring as the midrib; the basal

one-third dries brown as is the case in most species of Xanthophyllum.

Specimens examined: SABAH - Kinabatangan District, Ulu Sungai Pinangah

(SAN 81178, SAN 95970, SAN 107165 |type], SAN 107277 and SAN 107314);

Tawau district (SAN 95970). SARAWAK - Limbang District (Brunig 48,

specimen slightly deviating).

10. Xanthophyllum nitidum W.J. de Wilde & Duyfjes, sp. nov.

Tuxta Xanthophyllum vitellinum (Blume) D. Dietr., partibus omnibus

flavovirentibus, venatione intercostali tenuissima et acuta in folli pagina

inferiore nitidi, areolis consimilibus c. 0.5 mm diam. differt. Typus: Dewol,

Tuyok & Langkap SAN 108778, Borneo, Sabah, Kinabatangan District,

Bukit Tawai (holo SAN; iso A, BO, K, KEP, L, SAR, SING).

Tree to 20(—30) m tall and 20(—35) cm diam. Twigs yellow. Bark smooth,

black, inner bark yellowish. Sapwood white. Axillary buds c. 1 mm long,

minutely hairy. Leaves green-yellow, shiny above, concolorous, not papillose

beneath; blade oblong-lanceolate, 7—11 x 2—4 cm, base cuneate, apex acute-

acuminate; midrib above flat; lateral veins 4 or 5 pairs, intramarginal vein

indistinct; intercostal venation finely reticulate on both surfaces, areoles all

of about the same size, small, c. 0.5 mm diam., glands inconspicuous; petiole

8—12 mm long, transversely wrinkled. Inflorescences 5—10 cm long,

branched, minutely light brown hairy. Flowers 2 together at base, other

flowers solitary: pedicel 2-3 mm long, hairy; perianth, stamens and pistil

not known; developing ovaries and immature fruit sessile, globose, 4—6

mm diam., (not densely) hairy, light green, ovules 4. Fruit not known.

Distribution: Endemic to Borneo (Sabah and Kalimantan).

Ecology: Lowland forest, on brown soil over ultrabasic bedrock, at 100—400

m altitude.

Xanthophyllum in Borneo a7

SS we Mar

Figure 2. Xanthophyllum longum W.J. de Wilde & Duyfjes

A. Fruiting leafy twig; B. Part of inflorescence (post-anthesis) with developing ovaries

(young fruits). (A from SAN 107277, B from SAN 107165).

58 Gard. Bull. Singapore 57 (2005)

Notes: Xanthophyllum nitidum resembles superficially the variable and

widespread X. flavescens in its overall yellow-green colour of the dried

leaves, its yellow twigs, and its widely branched inflorescences. However,

it differs from the latter by its ovary, which is hairy all-over (glabrous in_X.

flavescens), shiny aspect of the leaves (not shiny in X. flavescens) and

finely and regularly areolate-reticulate venation (scalariform in X.

flavescens). It is also close to X. vitellinum, a species with the leaves dull

and mostly green-brown on drying and with coarser, irregularly sized

areoles. The collections from Sabah are from ultrabasic localities.

Specimens examined: SABAH - Kinabatangan District, Bukit Tawai (SAN

108778, the type); Sandakan District, Bukit Tangkunan (SAN 7/499), Bukit

Malawali (SAN 46624). EAST KALIMANTAN, Kutai (Arifin AA 968

and Sidiyasa 1118).

11. Xanthophyllum pachycarpon W.J. de Wilde & Duyfjes, sp. nov.

Iuxta Xanthophyllum tenuem Chodat, fructu maiore 2—3 cm diam.,

pericarpio crassiore 5—10 mm crasso differt. Typus: Lai et al. S 69651,

Borneo, Sarawak, Lubok Antu district, Nanga Segara, Sungai Engkari (holo

SAR; iso K, KEP, L, SAN, MO).

Tree 12-30 m tall and to 30 cm diam. Bark greyish or darkish green,

smooth or with large warty lenticels. Sapwood orange-yellow. Twigs

glabrous, smooth, yellowish. Axillary buds long-triangular, 1—1.5 mm long,

glabrous. Leaves glabrous, green-brown or light brown above, concolorous,

not papillose beneath; blade oblong, 8—16 x 2.5—6 cm, base cuneate, apex

acute-acuminate; midrib scarcely raised above, prominent beneath; lateral

veins 4-6 pairs, forming an irregular intramarginal vein; intercostal venation

finely and sharply reticulate on both surfaces; glands inconspicuous; petiole

6-12 mm long, longitudinally and transversely wrinkled, glabrous, glands

absent. Inflorescences much shorter than the leaves, unbranched, with 7-10

flowers (flower-scars), axis short hairy, glabrescent. Flowers not known.

Fruits glabrous, globose, 2-3 cm diam, light brown, coarsely wrinkled on

drying; pericarp 5-10 mm thick, solid or spongy due to irregularly-sized

scattered hollows; fruiting pedicel 3-4 mm long. Seed 1.

Distribution: Endemic to Borneo (Sabah, Sarawak and W Kalimantan).

Ecology: Mixed dipterocarp forest, on hill ridges and slopes, at altitudes to

650 m, also in forest on brown soil over ultrabasic bedrock.

Xanthophyllum in Borneo 59

Notes: This species, for which flowers are unknown, can be readily

recognized by its diagnostic large globose fruits, 2—3 cm in diam., which

dry light brown with a coarsely, wrinkled surface. The dry pericarp is

somewhat woody, 5—10 mm thick, solid or with scattered hollows of various

sizes. The venation of the glabrous concolourous leaves is fine and distinct

on both surfaces, by which the plant is reminiscent of Xanthophyllum

nitidum, but in the latter species the distinct venation 1s still finer, the

inflorescences branched and the young fruits densely hairy. In X.

pachycarpon, the fruits (and presumebly the ovaries) are glabrous.

Specimens examined: SABAH - Keningau District (SAN 78275 and SAN

78258); Kinabatangan District (SAN 52086, SAN 52099, SAN 53303 and

SAN 54072); Labuk Sugut District (SAN 82833); Ranau District (SAN

76778 and SAN 97680); Sandakan District (SAN 51297); Tenom District

(Aban et al. 65263). SARAWAK - Belaga District, Sungai Kenaban (S$

3525 and § 3541); Lubok Antu District (S 69651, the type); Serian District,

Sungai Engkabang (S$ 27397). WEST KALIMANTAN - Senawai (Church

et al. 1524).

12. Xanthophyllum rectum W.J. de Wilde & Duyfjes, sp. nov.

Tuxta Xanthophyllum vitellinum (Blume) D. Dietr., inflorescentiis crassis et

validis erectis et eramosis differt. Typus: Ilias Paie S 17903, Borneo, Sarawak,

Kuching District, Bako National Park, Lintang path (holo SAR; iso A,

BO, K, KEP, L, MEL, SAN, SING).

Tree 5—12 m tall and to c. 20 cm diam.. Bark pale grey, smooth or fine-

fissured. Inner bark yellow-brown. Twigs black, glabrous, c. 5 mm thick.

Axillary buds less than 0.5 mm long, subglabrous. Leaves coriaceous, dark

grey(-green) brown above, chocolate-brown and not papillose beneath;

blade elliptic-oblong, 7—13 x 3—7.5 cm, base rounded or short-cuneate,

apex acute-acuminate (very tip bluntish); midrib above flat or slightly raised;

lateral veins 4—7 pairs, not forming an intramarginal vein; intercostal

venation reticulate, prominent and distinct beneath; glands absent or

inconspicuous; petiole 7-12 mm long, without glands. Inflorescences stout,

straight, erect, unbranched, axis 15—20-flowered, 8—13 cm long, 2—8 mm

thick, sparsely minutely appressed brown hairy. Flowers solitary: pedicel c.

1 mm long; sepals c. 4 mm long, drying blackish; petals not seen; stamens

and mature pistil not seen; ovary sessile, (densely) hairy, ovules 4. Fruit

globose, c. 1.8 cm diam., smooth, + shiny, dark-brown, + sparingly hairy,

glabrescent; pericarp thin, seed 1; fruiting pedicel c. 2.5 mm long, 3—4 mm

thick.

60 Gard. Bull. Singapore 57 (2005)

Distribution: Endemic to Sarawak.

Ecology: Lowland heath forest, ridge of dipterocarp forest, and rocky

padang.

Notes: Earlier collected specimens of this species were left unnnamed by

Ng (in herb. KEP, 1968), but determined by van der Meijden (1982) as

Xanthophyllum vitellinum, which it resembles. Xanthophyllum vitellinum

normally has branched inflorescences, but occasionally has unbranched

ones, and it has membranous leaves. Xanthophyllum rectum 1s conspicuously

stout with thick twigs and coriaceous leaves with prominent venation

beneath, the whole plant dries dark brown, with stout, erect and unbranched

inflorescences. Apparently, it is confined to low heath forest of the Sampadi

Forest Reserve and Bako National Park, at low altitudes, in west Sarawak.

Specimens examined: SARAWAK - Kuching District, Bako National Park,

Lintang path (S 4446 and S 17903, the type), Sampadi Forest Reserve (S

66789), Selang Forest Reserve (Brunig 7343); Lundu District, Bedaun,

Sematan (S$ 65493).

13. Xanthophyllum rheophilum W.J. de Wilde & Duyfjes, sp. nov.

Xanthophyllo flavescenti Roxb. differt in foliis lineare-lanceolatis, petiolis

3—4 mm longis, fructu c. 1 cm diam. Typus: Ridsdale PBU 97, Borneo,

Central Kalimantan, Barito Ulu, 0.02' S 114° 06' E, (holo L; iso BO).

Shrub c. 3 m tall. Axillary bud minute. Leaves glabrous, yellow on drying,

lower surface concolorous, not papillose; blade lanceolate-linear, 6—/2 x

0.5—1 cm, base attenuate, apex long-acuminate; intercostal venation

reticulate; petiole 3-4 mm long, glabrous. Inflorescences racemose or few-

branched, c. 7 cm long, (S—)10—15-flowered. Flowers: pedicel c. 5 mm long;

petals cream; ovary subglobose, glabrous, except for a few hairs at apex,

ovules 12. Fruits globose, 0.8—1 cm diam., glabrous, blackish on drying.

Distribution: Endemic to Central Kalimantan; known only from the type.

Ecology: Riverine vegetation, at low altitude.

Notes: Xanthophyllum rheophilum is close to the broadly defined xX.

flavescens, a species usually with the intercostal venation scalariform and

with 5-15 mm-long petioles. The lanceolate-linear leaves with a short petiole

make X. rheophytum distinctive.

Xanthophyllum in Borneo 61

Acknowledgements

We thank the Director General of Forest Research Institute Malaysia, the

Director of the Sabah Forestry Department, the Director of the Sarawak

Forestry Department and Engkik Soepadmo (KEP) who gave us the

opportunity of working for the Tree Flora of Sabah and Sarawak Project.

We thank the Curators of the herbaria BO, KEP, L, SAN, SAR, and

SING for their hospitality and permission to study their collections. We

are especially grateful to Normaya Nordin and Soh Wuu Kuang (KEP),

Julia Sang (SAR) and Joan T. Pereira (SAN) for their assistance and

friendship. Ubaldus Majawal (SAN) made the beautiful drawings, and Jan

Frits Veldkamp (L) translated the diagnoses into Latin.

References

Mejjden, R. van der. 1982. Systematics and evolution of Xanthophyllum

(Polygalaceae). Leiden Botanical Series. 7: 1-159.

Meijden, R. van der. 1988. Polygalaceae. Flora Malesiana. 1, 10: 455—539.

Ng, F.S.P. ‘1968’ (1970). X. Xanthophyllum (Polygalaceae). In: K.M.

Kochummen, F.S.P. Ng and T.C. Whitmore, Notes on the systematy of

Malayan Phanerogams: VI—X, Federation Museums Journal (new series).

AS: 157.

Ng, F.S.P. 1972. Polygalaceae. Tree Flora of Malaya. 1: 351—366.

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as 96r5) Glan Cea ert lide =.

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Gardens’ Bulletin Singapore 57 (2005) 63—68& 63

Taxonomic Notes on Bornean Cryptocarya R.Br.

(Lauraceae)

F.S.P.. NG

Forest Research Institute Malaysia

52109 Kepong, Selangor, Malaysia

Abstract

Cryptocarya is sharply defined from other genera in the Lauraceae by floral morphology

and fruit development. The ovary is superior and seated within a deep and distinct

hypanthium. The developing fruit remains free from the hypanthium but is completely

enclosed by it. The resulting inferior fruit has two walls - an outer wall developed from the

hypanthium, and an inner wall derived from the ovary wall. The only other tree genera of

Lauraceae in SE Asia with inferior fruits are Eusideroxylon and Potoxylon, but in these

two genera, the ovaries are semi-inferior and the pericarp develops from the ‘fusion’ of the

receptacle and the ovary wall resulting in a single fruit wall.

In Borneo, 15 species of Cryptocarya are recognized. Seven species are reduced to synonymy.

Introduction

Cryptocarya is a genus of monoecious trees with leaves alternate or spiral,

mostly penniveined but in a few species triveined at the base. The

inflorescences are paniculate with first, second, third or, rarely, fourth

order branching, are terminal and axillary, and irregularly bracteate

(sometimes with a bract subtending every flower, sometimes not). The

flowers are bisexual, occupying terminal and lateral positions on the

panicles, small in all species (2-3 mm long and 1—2 mm wide in dried

specimens), bearing 6 perianth lobes (three outer and three inner), a ring

of 6 stamens alternating with 6 glands, and an inner ring of 3 stamens

alternating with 3 staminodes, at the rim of a distinct and deep hypanthium

(Fig. 1.Aa). The anthers are 2-locular. The ovary is completely surrounded

by the wall of the hypanthium but is free from it, and it remains free

throughout the development of the fruit (Fig. 1.Ab).

As the fruit develops, the perianth lobes, anthers and staminodes

absciss and the hypanthium wall becomes the outer fruit wall. This wall is

zoned into a thin skin, a fleshy middle layer and a hard inner layer. Within

this, the seed develops within the pericarp, which is dry at maturity. The

fruit of Cryptocarya, based on the character of the hypanthium wall, can

be considered as drupaceous.

64 Gard. Bull. Singapore 57 (2005)

Two other tree genera of Lauraceae in Borneo, Eusideroxylon and

Potoxylon, also have drupaceous inferior fruits. Kostermans (1957)

described the fruit development of Eusideroxylon as follows: “In

Eusideroxylon the flower tube is shallow, although it completely envelopes

the mature fruit”. Later, in describing the new genus Potoxylon, Kostermans

(1978) wrote: “Ovary base immersed in the perianth tube and adnate with

it...”. Kostermans’ comments are based on the definition of the inferior

fruit being the result of ‘fusion’ or ‘adnation’ of the ovary with a flower

tube or perianth tube.

There is no structure in Eusideroxylon and Potoxylon comparable

with the hypanthium of Cryptocarya. What we actually see in the flower of

Eusideroxlon and Potoxylon is a semi-inferior ovary, in which about half

of the ovule is embedded in the receptacle and half is above it (Fig. 1.Ba,

Ca). The fruit is the result of massive enlargement of the receptacle and

ovary wall resulting in the perianth lobes, stamens and staminodes persisting

as vestiges at the apex of the fruit (Fig. 1.Bb, Cb, Cc). The fruit is a drupe,

with its pericarp zoned into a skin-like epicarp, fleshy mesocarp and hard

endocarp. The fruits of Eusideroxylon and Potoxylon are similar in their

morphology but differ from the drupaceous fruit of Cryptocarya in the

receptacle and ovary wall being ‘fused’ to form the pericarp (Fig. 1.Bb, Cb,

Cc), whereas in the fruit of Cryptocarya the hypanthium wall remains free

from the pericarp (Fig. 1.Ab).

There are 15 species of Cryptocarya in Borneo, of which the most

common and variable one is C. ferrea.

Taxonomic Changes

1. Cryptocarya ferrea Blume

Bidr. Fl. Ned. Ind. (1826) 557. Type: Blume 1559, Java (holotype L).

This species is probably the commonest and most widely distributed in the

genus. It occurs throughout SE Asia and is highly variable in leaf size, leaf

shape and angle of inclination of the lateral veins. In this treatment, three

varieties, var. erectinervia, var. ferrea and var. scortechinii, are recognised

in Borneo, into which most specimens can be placed but many specimens

are intermediate and their placement only approximate.

Var. erectinervia (Kosterm.) Ng, stat. nov.

Basionym: C. erectinervia Kosterm., Reinwardtia 7 (1968) 307. Type:

Kostermans 9908, Kalimantan, Samarinda (holo L; iso BO, KEP, SAN).

This variety is characterized by lateral veins acutely inclined and almost

Cryptocarya in Borneo 65

Cc Cb

Figure 1. Longitudinal sections of the flowers and fruits of Cryptocarya, Eusideroxylon and

Potoxylon

Aa. flower of Cryptocarya ferrea var. ferrea (KMS 3299); Ab. young fruit of Cryptocarya ferrea

var. erectinervia (SAN 92730); Ba. flower of Potoxylon melagangai (S 57892); Bb. young fruit

of P melagangai (SAN 73320); Ca. flower of Eusideroxylon zwageri (KEP 98302); Cb and Cc:

young and mature fruits of E. zwageri (FSP Ng s.n., 14 May 05, freshly fallen, in FRIM

plantations.

ov ovule; ow ovary wall; h hypanthium; t tepal; s stamen or staminode; tes testa; en endocarp;

m mesocarp; ex exocarp. Scale indicates 1 mm.

66 Gard. Bull. Singapore 57 (2005)

straight until they approach the leaf margin; the leaf blades are longer than

15 cm and oblong in shape.

Distribution: Peninsular Malaysia and Borneo. In Borneo: Sabah - Kota

Kinabalu, Keningau, Ranau, Sipitang, Tawau (e.g. SAN 16485, SAN 16761,

SAN 38503 and SAN 90730); Sarawak - Sungai Jelalong, Kakus, Mt.

Mersing, Tubau (e.g. S 4947, S 21828, S 22490 and S 48984); Kalimantan -

E. Kutei and Samarinda (e.g. Kostermans 5566, 5908, 9931, 9908).

Var. scortechinii (Gamble) Ng, stat. nov.

Basionym: C. scortechinii Gamble, Bull. Misc. Inform. Kew (1910) 143.

Type: King’s Collector 6297 (lectotype L = Sheet No. 003624880761, here

designated).

This variety is characterized by leaves broadly oblong or broadly elliptic,

with lateral veins distantly spaced.

Distribution: Peninsular Malaysia and Borneo. In Borneo: Sabah - Lamag,

Mostyn, Pulau Sapanggar (e.g. SAN 23858, SAN 35293 and SAN 57269);

Sarawak - Belaga, Lambir, Lubok Antu, Selampit, Semengoh (e.g. S 24109,

S 26987, S 44076 and S 69647); Brunei — Andulau, Belait, Bukit Puan (e.g.

Wong WKM 950, BRUN 578 and BRUN 2635).

Var. ferrea

New synonyms:

C. tomentosa Blume, Mus. Bot. Lugd. Bat. 1 (1851) 335. Type: Blume s.n.,

Java (holo L = Sheet No. 8983353).

C. kurzii Hook. f;, Fl. Brit. Ind. 5 (1886) 119. Type: Gnjjith 1742 (= Kew

Distr. 4274), Burma, Mergui (holo K).

C. bicolor Merr., Phil. J. Bot. 4 (1909) 255. Type: Hutchinson For. Bur.

6548, Philippines, Mindanao (holo not seen; iso SING).

C. argentea Gamble, Bull. Misc. Inform. Kew (1910) 144; C. kurzii Hook. f.

var. argentea (Gamble) Airy Shaw, Bull. Misc. Inform. Kew (1939)

535. Type: King’s Collector 7966, Perak (holo K; iso SING).

C. tawaensis Merr., Plantae Elmerianae Borneenses. (1929) 89. Type: E/mer

21418, Sabah, Tawao (holo not seen; iso L, U). (Elmer spelled

Tawau as “Tawao’ on the collecting label).

C. kurzii Hook. f. var. subsericea Airy Shaw, Bull. Misc. Inform. Kew

(1939) 535. Type: Richards 2443 Sarawak, Gunung Balapau (holo

K; iso SING).

C. borneensis Kosterm., Reinwardtia 7 (1968) 302. Type: Wood SAN 16257,

Sabah, Sipitang (holo L).

Cryptocarya in Borneo 67

This variety 1s characterized by leaves narrowly elliptic, ovate or obovate,

with lateral veins closely spaced.

Distribution: Myanmar, Peninsular Malaysia, Borneo, the Philippines, Java

and Sulawesi; probably throughout SE Asia. In Borneo, common

throughout Sabah (e.g. SAN 27336, SAN 46277 and SAN 76444); Sarawak

at Bako, Batang Tinjau, Belaga, Sungai Temulan and Tubau (e.g. S 39990,

S$ 41010 and SFN 35720); Brunei at Andulau Forest Reserve (e.g. BRUN

16914); Kalimantan at Balikpapan, Gunung Bentuang, Nunukan,

Samarinda, Sankulirang (e.g. Kostermans 5953, Kostermans 8937 and

Kostermans 9946).

2. Cryptocarya griffithiana Wight

Icon. 5 (1852) 12, t. 1830. Type: Griffith s.n., Malacca (not located).

A widespread species ranging from Myanmar to Sumatra, Peninsular

Malaysia, Borneo and the Philippines. Two varieties are recognised,

differing consistently only in the length of the bracts that subtend the

flowers.

Var. crassinervia (Miq.) Ng, stat. nov.

Basionym: Cryptocarya crassinervia Miq., Fl. Ind. Bat. 1, 1 (1858) 924.

Type: Teijsmann s.n., Sumatra, Fort de Kock (holo L = Sheet No.

905229445).

This differs from the typical variety in the floral bracts shorter than the

flowers they subtend.

Distribution: Sumatra, Peninsular Malaysia, Borneo and the Philippines.

In Borneo: Sabah - Beaufort, Kalabakan, Kota Belud, Sandakan, Semporna,

Sipitang, Tawau, (e.g. SAN 86242, SAN 88178, SAN 102650 and SAN

121972); Sarawak - Belaga, Kuching, Limbang (Gunung Pagon), Kelabit

Highlands, Lambir, Lundu (Bukit Berumput), Simunjan (e.g. Haviland &

Hose 3295, S 38465, S 46345 and S 70823); Brunei - Bukit Apoi and Belait

(e.g., Coode 7921, Forman 866 and Wong WKM 1281) and Kalimantan -

Tumbang Tapi and Bukit Raya (e.g. Veldkamp 8252 and Veldkamp 8549).

Var. griffithiana

The typical variety has floral bracts longer than the flowers they subtend.

It is rare or possibly extinct in Borneo, being known only by one specimen,

Haviland 870, from Lundu, Sarawak. The type specimen of this variety

could not be found at Kew, but this taxon is well-known in the Malay

68 Gard. Bull. Singapore 57 (2005)

Peninsula and there is no doubt about its identity.

3. Cryptocarya wrayi Gamble

Bull. Misc. Inform. Kew (1910) 142. Type: Wray 3853, Peninsular Malaysia,

Gunung Bubu, (holotype K).

New synonym: C. tuanku-bujangii Kosterm., Reinwardtia 8 (1970) 79. Type:

Paie S 26404, Sarawak, Lawas, path to Gunung Murut (second summit).

(holo SAR; iso L, SING).

This is a small tree to 4 m tall, on the summits of mountains. The plants in

the Matay Peninsula named Cryptocarya wrayi and those in Borneo named

C. tuanku-bujangii both occupy mountain summits and both have triveined,

ovate, coriaceous leaves. I think they are at most only minor geographic

variants of the same species.

Distribution: Peninsular Malaysia and Borneo. In Borneo: Brunei - Gunung

Pagon Periok (e.g. Ashton BRUN 2388) and Sarawak - Batu Lawi, Gunung

Murud and Gunung Mulu, (e.g. S 26404, S$ 38839 and S 50869).

Acknowledgements

I am grateful to the curators of the herbaria at SAN, SAR and SING for

the loan of specimens examined for this study, to the Tree Flora of Sabah

and Sarawak Project for funding the study and to Prof. Jens Rohwer and

Dr Henk van der Werff for kindly reviewing the manuscript.

References

Kostermans A.J.G.H. 1957. Lauraceae. Communications of the Forest

Research Institute (Indonesia). 57: 1—64.

Kostermans, A.J.G.H. 1978. Potoxylon, a new Bornean genus of Lauraceae.

Malayan Nature Journal. 32: 143-148.

Gardens’ Bulletin Singapore 57 (2005) 69—100 69

A Synopsis of the Genus Actinodaphne Nees

(Lauraceae) in Sabah and Sarawak, Malaysia

S. JULIA

Forest Research Centre

KM10, Jalan Datuk Amar Kalong Ningkan,

93250 Kuching, Sarawak, Malaysia

Abstract

In Sabah and Sarawak, a total of 20 species of Actinodaphne are recognized including one

imperfectly known species. Of these, eight (Actinodaphne kostermansii S. Julia, Actinodaphne

percoriacea S. Julia, Actinodaphne robusta S. Julia, Actinodaphne semengohensis S. Julia,

Actinodaphne soepadmot S. Julia, Actinodaphne spathulifolia S. Julia, Actinodaphne sulcata

S. Julia and Actinodaphne venosa S. Julia) are new to science. In addition, two varieties,

Actinodaphne kostermansii var. glabrescens S. Julia and Actinodaphne sulcata var.

longipetiolata S. Julia are also described as new. Relevant references, basionyms, type

specimens (if known), synonyms, distribution, ecology and notes for each species occurring

in Sabah and Sarawak are provided. An identification list for all specimens examined is

given.

Introduction

The genus Actinodaphne Nees was established by C. G. Nees von Esenbeck

in 1831 based on A. pruinosa Nees from Peninsular Malaysia. Since then, a

total of 150 binomials have been published by various authors (IPNI, 2005).

Of these, 14 binomials were attributed to species from Borneo of which 11

occur in Sabah and Sarawak (Blume, 1851; Miquel, 1858; Merrill, 1921,

1929; Masamune, 1942; Burgess, 1966; Anderson, 1980; Kochummen, 1989;

Coode et al., 1996; Argent et al., 1997; and Beaman et al., 2001).

The present study, based mainly on available specimens collected

from Borneo and preserved at the BO, KEP, PNH, Kinabalu Park, SAN,

SAR and SING herbaria and supplemented with images of type specimens

from the data bases/websites of the National Herbarium of the Netherlands,

University of Leiden Branch (L), New York Botanic Garden (NY) and

Royal Botanic Gardens, Kew (K) resulted in 20 species (including one

incompletely known species) being recognized. Of these, eight are new to

science and one is a new record for Borneo (A. johorensis Gamble). Of

the eight new species, four (A. percoriacea, A. semengohensis, A.

spathulifolia and A. sulcata) are endemic in Sabah and Sarawak, two in

Borneo (also occurring in Brunei and/or Kalimantan; A. kostermansii and

A. venosa) and two also occur outside Borneo (A. robusta, Peninsular

Malaysia and the Philippines and A. soepadmoi, Peninsular Malaysia).

70 Gard. Bull. Singapore 57 (2005)

The genus Actinodaphne comprises about 100 recognized species

(Kostermans, 1957; Rohwer, 1993; van der Werff, 2001), distributed from

India and Sri Lanka to Myanmar, Thailand, Indo-China, China, Korea,

Japan, Malesia and the Solomon Islands.

In Borneo, species of Actinodaphne occur in various forest types

on different soils, including mixed dipterocarp forest, peat swamp forest,

kerangas forest, riparian forest and forest on limestone and ultrabasic soils,

at altitudes from sea level to 2400 m.

Actinodaphne Nees

Actinodaphne Nees in Wallich, Pl. As. Rar. 2 (1831) 61, 68; Gamble, J. As.

Soc. Beng. 75, 1 (1912) 112; Mernill, J- Sir Br. Koy As: Soc. Spee

Number (1921) 274, Univ. Calif. Publ. Bot. 15 (1929) 78; Ridley, FI.

Malay Penins. 3 (1924) 107; Masamune, En. Phan. Born. (1942) 306;

Kostermans, Comm. For. Res. Inst. Bogor (1957) 42; Backer &

Bakhuizen f,, Fl. Java 1 (1964) 124; Burgess, Sabah For. Record 6 (1966)

330; Anderson, Checkl. Trees Sarawak (1980) 220; Corner, Wayside

Trees of Malaya a edition, 1 (1988) 382; Kochummen, Tree FI. Malaya

4 (1989) 102; Rohwer in Kubitzki et al. (eds.), Fam. Gen. Vas. Pl. 2

(1994) 366; Turner, Gard. Bull. Sing. 47 (1995) 273; Coode et al. (eds.),

Checkl. Flow. Pl. Gymno. Brunei (1996) 149; Argent et al. (eds.), Man.

Non-Dipt. Trees Centr. Kalimantan 1 (1997) 301; Beaman et al., Pl. Mt.

Kinabalu (2001) 393; van der Werff, Blumea 46 (2001) 125.

Type species: Actinodaphne pruinosa Nees.

Terminal (vegetative) buds perulate; scales imbricate, often leaf-like, on

falling leaving distinct scars just above the whorls of leaves. Leaves pinnately

veined, usually in whorls of 3—9 or rarely not strictly in whorls; upper

surface plane or bullate, often shining, lower surface often glaucous; lateral

veins disappearing towards the margin or occasionally joining near the

margin at the upper half of the leaf blade; tertiary veins scalariform,

subscalariform or reticulate. Inflorescences axillary or extra-axillary, usually

condensed umbels, pseudo-umbels or glomerules (or the males rarely

racemes as in A. johorensis Gamble and A. semengohensis, or panicles as

in A. montana Gamble) of sessile or pedicelled flowers borne on short

lateral shoots with or without terminal vegetative buds, surrounded by

sessile involucral, imbricate bracts which on falling leave distinct scars at

the base of the inflorescence. Flowers trimerous, unisexual; perianth lobes

6, outer lobes slightly larger than the inner ones; fertile stamens in male

flowers and staminodes in female flowers usually 9, arranged in 3 whorls,

those of the first and second whorls non-glandular, that of the third whorl

Actinodaphne in Borneo 71

with stalked glands on each side at the base; filaments longer or shorter

than anthers, anthers 4-locular, the /ocules all introse, arranged in two

pairs above each others; pistillode in male flower rudimentary and minute

or absent; ovary in female flowers superior, narrowed towards the style,

stigma peltate, dilated or discoid. Infructescences each bearing 1—18 fruits.

Fruits drupaceous, seated on a flat or shallowly saucer-shaped or deeply

cup-shaped, accrescent or non-accrescent cupule with or without a remnant

of perianth lobes.

Enumeration of Species Occurring in Sabah and Sarawak

1. Actinodaphne borneensis Meisn.

Actinodaphne borneensis Meisn. in A.DC., Prodr. 15, 1 (1864) 213; Merrill,

J. Str. Br. Roy. As. Soc., Special Number (1921) 274; Masamune, En.

Phan. Born. (1942) 306; Burgess, Sabah For. Record 6 (1966) 330;

Anderson, Checkl. Trees Sarawak (1980) 220; Coode et al. (eds.), Checkl.

Flow. Pl. Gymno. Brunei (1996) 149. Type: Lobb s.n., Borneo (holo

n.v.; iso K [K000009921]).

Distribution: Endemic in Borneo (Brunei, Kalimantan, Sabah and Sarawak).

Ecology: In mixed dipterocarp, beach, peat swamp, kerangas and

submontane forests (including forest on ultrabasic soil) at altitudes from

sea level to 1500 m.

Notes: A widespread species in Borneo, characterized by its well-spaced

lateral veins of 3—7 pairs, tertiary veins obscure on both surfaces or obscure

above, prominulous below, pseudo-umbellate inflorescences each bearing

up to 7 flowers, and fruit, which is seated on a shallowly saucer-shaped or

deep cup-shaped cupule. The leaves are highly variable in size, ranging

from 3.5 x 2 cm to 25 x 8.5 cm.

The species resembles Actinodaphne pruinosa Nees and A. oleifolia

Gamble but can be differentiated from the former by its lax or well-spaced

(vs. dense) lateral veins and from the latter by its scalariform (vs. pitted or

reticulate) and obscure or prominulous tertiary veins underneath (vs.

distinctly prominent).

Kochummen (1989) described specimens from Peninsular Malaysia

(Corner SFN 21345, Kiah SFN 32386 and FRI 13451) as A. borneensis.

However, after re-examining the specimens and comparing them with that

of the type as well as other specimens of A. borneensis collected from

Borneo, I am of the opinion that the three specimens from Peninsular

72 Gard. Bull. Singapore 57 (2005)

Malaysia belong to A. malaccensis Hook.f., which does not occur in Sabah

and Sarawak.

2. Actinodaphne diversifolia Merr.

Actinodaphne diversifolia Merr., J. Str. Br. Roy. As. Soc. 85 (1922) 191;

Masamune, En. Phan. Born. (1942) 306; Burgess, Sabah For. Record 6

(1966) 330; Coode et al. (eds.), Checkl. Flow. Pl. Gymno. Brunei (1996)

149; Beaman et al., Pl. Mt. Kinabalu (2001) 393. Type: Ramos 1838,

Borneo, Sabah, Sandakan District, Sebuga (holo K; iso, n.v.: A, BO).

Distribution: Endemic in Borneo (Brunei, Kalimantan, Sabah and Sarawak).

Ecology: In beach, peat swamp, kerangas, mixed dipterocarp and

submontane forests at altitudes to 1200 m.

Notes: Common and widespread in Sabah and well represented in Gunung

Kinabalu National Park, Crocker Range National Park and Maliau Basin

Conservation Area but it is very rare in Sarawak. Its narrowly elliptic

leaves that are hairy on the lower surface readily distinguish the species.

The species resembles Actinodaphne kostermansii var. kostermansii

but can be differentiated by its linear (vs. lanceolate) bud-scales, narrow

elliptic, unequal-sided leaves (vs. broadly elliptic; equal-sided) with sharply

acute (vs. rounded or broadly acute) base, slightly revolute (vs. strongly

revolute) margins, and closely scalariform (vs. distantly scalariform) tertiary

veins.

3. Actinodaphne fuliginosa Airy Shaw

Actinodaphne fuliginosa Airy Shaw, Bull. Misc. Inform., Kew (1939) 535;

Masamune, En. Phan. Born. (1942) 306. Type: Synge 1893, Borneo,

Sarawak, Dulit Range (holo K; iso SING).

Distribution: Endemic in Borneo (Sarawak). Very rare and known only

from the type specimen from Dulit Range in Sarawak.

Ecology: In open mossy forest on exposed peak at c. 1400 m altitude.

Notes: This distinctive species is characterized by its small, 2.5—4.5 x 1.5—

2.5 cm, obovate leaves with a rounded apex.

Actinodaphne in Borneo 13

4. Actinodaphne glabra Blume

Actinodaphne glabra Blume, Mus. Bot. Lugd-Bat. I (1851) 344; Backer &

Bakhuizen f., Fl. Java 1 (1964) 125; Beaman et al., Pl. Mt. Kinabalu

(2001) 393. Type: Blume s.n., Java (lecto L[NHN-L Acc. No. 905220121],

designated here).

Distribution: Peninsular Malaysia, Java and Borneo (Brunei, Kalimantan,

Sabah, Sarawak).

Ecology: In mixed dipterocarp and riparian forests on alluvial and ultrabasic

soils at altitudes to 600 m.

Notes: The species is characterized by its long and narrowly elliptic or

oblanceolate leaves with long acuminate or pointed leaf apex. In Borneo,

however, there are specimens with rounded, obtuse or shortly acuminate

leaf apex (e.g. S S151, SAN 25354, SAN 37438, SAN 55839 and SAN

132242). Images of type specimens of two other species (Actinodaphne

pubescens Blume from Java and A. rumphii Blume from the Moluccas)

strongly suggest that these two species may be conspecific with A. glabra.

More specimens from the relevant areas for detailed study are, however,

needed to elucidate the taxonomic status of the two taxa.

5. Actinodaphne glomerata (Blume) Nees

Actinodaphne glomerata (Blume) Nees, Syst. Laur. (1836) 597; Blume,

Mus. Bot. Ludg.- Bat. 1 (1851) 343; Miquel, Fl. Ind. Bat. 1(1858) 968;

Gamble, J. As. Soc. Beng. 75, 1 (1912) 116; Ridley, Fl. Malay Penins. 3

(1924) 108; Backer & Bakhuizen f, Fl. Java 1 (1964) 124; Burgess,

Sabah For. Record 6 (1966) 330; Anderson, Checkl. Trees Sarawak

(1980) 220; Corner, Wayside Trees of Malaya 3 edition, 1 (1988) 220;

Kochummen, Tree Fl. Malaya 4 (1989) 104; Turner, Gard. Bull. Sing. 47

(1995) 273; Coode et al. (eds.), Checkl. Flow. Pl. Gymno. Brunei (1996)

149; Argent et al. (eds.), Man. Non-Dipt. Trees Centr. Kalimantan 1

(1997) 303; Beaman et al., Pl. Mt. Kinabalu (2001) 393. — Litsea

glomerata Blume, Bijdr. Fl. Ned. Ind. (1826) 566. Type: Blume s.n., W

Java, Gunung Salak, (holo n.v.; iso NY [00355279]).

Syn. nov.: Actinodaphne maingayi Hook.f. var. macrocarpa Gamble, J. As.

Soc. Beng. 75, 1 (1912) 115; Actinodaphne sesquipedalis Hook.f. &

Thoms. var. macrocarpa (Gamble) Ridl., Fl. Malay Penins. 3 (1924)

108. Type: Ridley 11675, Borneo, Sarawak, Matang (/ecto SING,

designated here)

74 Gard. Bull. Singapore 57 (2005)

Distribution: Sumatra, Java, Peninsular Malaysia, Singapore, Borneo

(Brunei, Kalimantan, Sabah and Sarawak) and Sulawesi.

Ecology: A very common and widespread species occurring in mixed

dipterocarp, riparian and submontane forest at altitudes to 1500 m, on

various soil types.

Notes: In Borneo, the leaves of this species are very variable in shape,

ranging from obovate to broadly elliptic or suborbicular and in size from

11—41 cm long and 5—23 cm wide, with a length:width ratio of 2:1 to 1:1

depending on their position on the branch and the habitats where the trees

grow. However, its thinly coriaceous leaves with a glaucous or brownish

undersurface and many-flowered, condensed, glomerulate inflorescences

easily distinguish the species. The specimen (Mikil SAN 38734) from Mount

Kinabalu cited by Beaman et al. (2001) as Actinodaphne sesquipedalis

Hook.f. & Thoms. belongs to A. glomerata.

6. Actinodaphne johorensis Gamble

Actinodaphne johorensis Gamble, Bull. Misc. Inform., Kew (1910) 313;

Gamble, J. As. Soc. Beng. 75, 1 (1912) 117; Ridley, Fl. Malay Penins. 3

(1924) 109; Kochummen, Tree Fl. Malaya 4 (1989) 104; Turner, Gard.

Bull. Sing. 47 (1995) 273. Type: Ridley 4419, Johor, Gunung Panti (holo

SING; iso SING).

Distribution: Peninsular Malaysia and Borneo (Sarawak).

Ecology: Confined to kerangas forest at altitudes to 400 m.

Notes: The species was previously known as endemic in Peninsular Malaysia

(Johor and SW Pahang). This is a new record for Borneo and is known

only from the Kuching, Sri Aman and Lundu Districts, Sarawak.

Actinodaphne johorensis can easily be recognized by its thickly

coriaceous, glabrous leaves (except on the midrib and lateral veins), usually

glaucous underneath and slightly prominent lateral veins, almost obscure

tertiary veins and racemose male inflorescence.

7. Actinodaphne kinabaluensis Kosterm.

Actinodaphne kinabaluensis Kosterm., Reinwardtia 7 (1969) 452; Beaman

et al., Pl. Mt. Kinabalu (2001) 394. Type: Chew et al. RSNB 196, Borneo,

Sabah, Gunung Kinabalu (holo BO; iso K, L, SING).

Actinodaphne in Borneo TO

Distribution: Endemic in Borneo (Sabah); rare and known only from the

type.

Ecology: Montane forest at 2400 m altitude.

Notes: Even though this species is known only from one fruiting specimen,

it definitely belongs to Actinodaphne as shown by its vegetative and fruit

characters that conform to those of the genus.

The species can easily be recognized by its thickly coriaceous,

densely hairy, bullate leaves and ellipsoid fruits. It is the only Actinodaphne

species in Sabah and Sarawak with ellipsoid fruit.

8. Actinodaphne kostermansii S.Julia, sp. nov.

(André Joseph Guillaume Henri Kostermans, 1906—1981, prominent

botanist at the Herbarium Bogoriense)

Actinodaphne diversifoliae similis, perulis lanceolatis (vs. linearibus), foliis

late (vs. anguste) ellipticis aequilateris (vs. inaequilateris), basi acuta ad

late (vs. argute) acuta, apice acuto ad breviter acuminato (acumine, 0.5—1

cm vs. 1.5—2 cm longo), nervis tertiariis distantiter (vs. arte) scaliformibus

differt. Typus: Clemens 50386, Borneo, Sabah, Kota Kinabalu District,

Gunung Kinabalu, Penibukan (holo L).

Tree 6—20 m tall, 7—30 cm diam.; bole straight, c. 6 m tall; buttresses

absent. Bark brownish green or dark grey, smooth; inner bark yellowish,

fibrous. Sapwood yellowish. Twigs drying brown, glabrescent, smooth.

Terminal (vegetative) buds: scales lanceolate, 10—30 x 4-8 mm, densely

hairy. Leaves in whorls of 3—5, thickly coriaceous, glabrous or very sparsely

hairy above, glabrous or densely hairy below, drying brown or greenish

brown above, greyish or brownish below; blade broadly elliptic, (15.5—)

21—31(—39) x 7—15.5 cm, base broadly acute, margin strongly revolute,

apex acute or acuminate, acumen 0.5—1 cm long; midrib raised on both

surfaces, stronger below, glabrous or sparsely hairy above, glabrous or

densely hairy below; lateral veins 7—11 pairs, lax, at an angle of c. 30° from

the midrib, flat or sunken above, strongly raised below, joining towards

margin at the upper half of leaf; tertiary venation slightly distinct or obscure

above, distinct below, distantly scalariform; petiole 1.5—3.5 cm long, sparsely

or densely hairy, drying black or dark brown. Inflorescences and flowers

unknown. Infructescences each bearing 1—7 fruits (in glomerulate

arrangement); vegetative terminal buds present. Fruits globose, 1.3—1.5 cm

diam., drying black; cupule sometimes accrescent, deeply or shallowly

saucer-shaped, 1—1.2 cm across, remnant of perianth lobes absent or present;

76 Gard. Bull. Singapore 57 (2005)

pedicels 5—7 mm long. Seeds globose, 0.8—1 cm diam., drying dark brown.

Distribution: Endemic in Borneo (confined to Brunei and Sabah).

Ecology: Uncommon in mixed dipterocarp, submontane, riparian forests

and forest on ultrabasic soil at altitudes to 1400 m.

Notes: Even though inflorescences and flowers are not available, this species

is placed in Actinodaphne based on its vegetative (e.g. perulate terminal

bud, imbricate bud scales, pinnately veined leaves arranged in whorls of 3—

5 and distinct bud-scale scars above the whorls of leaves) and fruiting

characters.

The species is reminiscent of A. diversifolia but differs by its

lanceolate (vs. linear) bud-scales, broadly elliptic, equal-sided leaves (vs.

narrow elliptic, unequal-sided) with rounded or broadly acute (vs. sharply

acute) base, strongly revolute (vs. slightly revolute) margins, and distantly

scalariform (vs. closely scalariform) tertiary veins.

In 1977, Kostermans annotated specimen Clemens 50386 as A.

clemensii, a new species but he never validated the name of the new taxon.

This new species is renamed in honour of Dr. A.J.G.H. Kostermans who

made an enormous contribution toward the advancement of our knowledge

on the Lauraceae of the Malesian region.

Beaman et al. (2001) incorrectly identified the same specimen

(Clemens 50386) as A. sesquipedalis Hook.f. & Thoms., a species only

known from Myanmar and Peninsular Malaysia and differing from A.

kostermansii by its oblanceolate, obovate or elliptic-oblong (vs. broadly

elliptic) leaves arranged in whorls of 5—13 (vs. in whorls of 3—5) with

strongly prominent (vs. obscure or prominulous) tertiary veins.

In Sabah and Sarawak, two varieties are recognized, viz. var.

kostermansii and var. glabrescens.

var. kostermansii

Distribution: Endemic in Borneo (Sabah and Brunei). Found in montane,

riparian and mixed dipterocarp forests, at altitudes to 1400 m.

Notes: Leaves and midrib very sparsely hairy above, densely hairy below.

Other specimens examined: BORNEO —- BRUNEI, Temburong District,

Amo, Ulu Temburong, Coode et al. MC 7869 (KEP, SAR); SABAH, Ranau

District, Sosopodon Forest Reserve, Aban SAN 64101 (SAN).

Actinodaphne in Borneo Ta

var. glabrescens S.Julia, var. nov.

(Latin, glabrescens=becoming glabrous; referring to the leaves)

A var. typica folis glabris (vs. infra dense pubescentibus) differt. Typus:

Proctor SAN 98112, Borneo, Sabah, Lahad Datu District, Gunung Silam

(holo SAN).

Distribution: Endemic in Sabah and found in mixed dipterocarp forests

and forest on ultrabasic soil, at altitudes to 800 m.

Notes: This new variety differs from var. kostermansii in having leaves that

are glabrous (on both surfaces) or very sparsely hairy (particularly on the

midrib and lateral veins below).

Other specimens examined: BORNEO — SABAH, Kota Belud District,

Melangkap Tomis, Lorence Lugas 1922 (BO, KEP, KNP, SAN, SAR),

Lahad Datu District, Gunung Silam, Mujin SAN 37821 (SAN), Proctor

SAN 98060 (SAN), Proctor SAN 98118 (SAN), Proctor SAN 100721 (SAN),

Proctor SAN 100729 (SAN), Proctor SAN 101957 (SAN), Rimi et al. SP

6276 (KNP), Ranau District, Kinabalu National Park, Kokawa & Hotta

5603 (SAN).

9. Actinodaphne macrophylla (Blume) Nees

Actinodaphne macrophylla (Blume) Nees, Syst. Laur. (1836) 598; Blume,

Mus. Bot. Ludg.-Bat. 1 (1851) 341; Miquel, Fl. Ind. Bat. 1,1 (1858) 965;

Backer & Bakhuizen f., Fl. Java 1 (1964) 125; Kochummen, Tree FI.

Malaya 4 (1989) 105; Turner, Gard. Bull. Sing. 47 (1995) 273; Coode et

al. (eds.), Checkl. Flow. Pl. Gymno. Brunei (1996) 149; Argent et al.

(eds.), Man. Non-Dipt. Trees Centr. Kalimantan 1 (1997) 303. — Litsea

macrophylla Blume, Biydr. Fl. Ned. Ind. (1826) 567. Type: Blume s.n.,

Java, Nusa Kambangan (7.v.).

Actinodaphne maingayi Hook.f., Fl. Brit. Ind. 5 (1886) 151; Gamble, J. As.

Soc. Beng. 75, 1 (1912) 114; Merrill, J. Str. Br. Roy. As. Soc., Special

Number (1921) 274; Ridley, Fl. Malay Penins. 3 (1924) 108; Masamune,

En. Phan. Born. (1942) 306; Corner, Wayside Trees of Malaya aa edition,

1 (1988) 345. Type: Maingay 1258, Peninsular Malaysia, Malacca (lecto

K, designated here; iso L).

Distribution: Peninsular Malaysia, Singapore, Java and Borneo (Brunei,

Kalimantan, Sabah and Sarawak).

78 Gard. Bull. Singapore 57 (2005)

Ecology: In mixed dipterocarp, riparian, kerangas and swamp forests, at c.

890 m altitude.

Notes: The species is characterized by its thickly coriaceous, broadly elliptic-

oblong or oblanceolate leaves, which are densely hairy underneath with

very strongly prominent lateral veins and sturdy petiole. Actinodaphne

macrophylla is very close to A. sesquipedalis from Myanmar and Peninsular

Malaysia but can be differentiated by its very densely hairy (vs. glabrous)

twig, leaves densely hairy underneath (vs. sparsely hairy or glabrous) with

very distinctly prominent (vs. almost obscure or slightly prominent) tertiary

veins.

10. Actinodaphne myriantha Merr.

Actinodaphne myriantha Merr., Univ. Calif. Publ. Bot. 15 (1929) 78;

Masamune, En. Phan. Born. (1929) 306; Burgess, Sabah For. Record 6

(1966) 330; Anderson, Checkl. Trees Sarawak (1980) 220. Type: Elmer

21335, Borneo, Sabah, near Tawau (holo PNH 7; iso BO, L, MO, SING).

Distribution: Endemic in Borneo (Brunei, Kalimantan, Sabah and Sarawak).

Ecology: In mixed dipterocarp, peat swamp, and riparian forests at altitudes

to 700 m.

Notes: This species resembles Actinodaphne glomerata but differs by its

narrowly obovate or narrowly elliptic (vs. broadly obovate or broadly

elliptic), much longer and narrower leaves with a length:width ratio of 3:1

(vs. length:width ratio of 2:1 or 1:1).

11. Actinodaphne oleifolia Gamble

Actinodaphne oleifolia Gamble, Bull. Misc. Inform., Kew (1910) 313;

Gamble, J. As. Soc. Beng. 75, 1 (1912) 121; Merrill, J. Str. Br. Roy. As.

Soc., Special Number (1921) 274; Ridley, Fl. Malay Penins. 3 (1924)

111; Masamune, En. Phan. Born. (1942) 306; Burgess, Sabah For. Record

6 (1966) 330; Kochummen, Tree FI. Malaya 4 (1989) 106; Turner, Gard.

Bull. Sing. 47 (1995) 274; Coode et al. (eds.), Checkl. Flow. Pl. Gymno.

Brunei (1996) 149. Type: Ridley 13728, Peninsular Malaysia, Pahang,

Gunung Berembun (lecto K, designated here; iso SING).

Actinodaphne gelonioides Ridl., Fl. Malay Penins. 3 (1924) 111. Type:

Robinson s.n., Peninsular Malaysia, Perak, Gunung Kerbau (holo K).

Syn. nov.: Actinodaphne foxworthyana Gibbs, J. Linn. Soc. Bot. 42 (1914)

Actinodaphne in Borneo 719

129; Merrill, J. Str. Br. Roy. As. Soc., Special Number (1921) 274;

Masamune, En. Phan. Born. (1942) 306; Burgess, Sabah For. Record 6

(1966) 330. Type: Gibbs 3135, Borneo, Sabah, Kota Belud District,

Gunung Kinabalu, Paka Cave (holo BM; iso K).

Distribution: Peninsular Malaysia and Borneo (Brunei, Kalimantan, Sabah

and Sarawak).

Ecology: In mixed dipterocarp, submontane to upper montane, limestone,

kerangas and peat swamp forests and forest on ultrabasic soil at altitudes

to 1800 m.

Notes: The species is rather similar to Actinodaphne pruinosa in its smallish

leaves but it is distinct in having obtuse or shortly acuminate leaves, with

an acumen 1—1.5 cm long (vs. with an acumen 1.5—3 cm long) and pitted

or reticulate (vs. scalariform) tertiary veins that are distinctly prominent

on both surfaces (vs. obscure or prominulous above, prominent below).

From the small-leaved A. borneensis, the species differs by its dense

(vs. lax or well-spaced) lateral veins and reticulate (vs. scalariform) and

distinctly prominent (vs. obscure or slightly prominent) tertiary veins.

Specimens that were previously recognized by Kochummen (1989)

as A. oleifolia and/or A. gelonoides (FRI 10111, Holttum SFN 20725, Megsay

& Kiah SFN 31832, Mohd. Shah MS 1461, FMS 43076, Symington & Kiah

SFN 28831, Seimund 341) from Peninsular Malaysia may respresent an

undescribed new species differing from A. oleifolia by its almost rounded

and broader thickly coriaceous leaves.

12. Actinodaphne percoriacea S. Julia, sp. nov.

(Latin, per=very; coriaceous=leathery; referring to the very thick leaves)

Actinodaphne borneensis foliis magnis revocans, foliis glabris (vs. infra

sparse pubescentibus), basi rotundata ad truncata (vs. acuta), venis

lateralibus densis (vs. laxis ad clare dispositis) facile distinguenda. Typus:

Clemens 31493, Borneo, Sabah, Kota Kinabalu District, Gunung Kinabalu,

Penibukan (holo BO).

Small tree or treelet to 6 m tall. Bark pale brown, smooth; inner bark

yellowish. Sapwood yellowish. Twigs drying greyish brown, densely hairy

when young, sparsely hairy when older, smooth. Terminal buds: scales

ovate, 3—9 x 3—5 mm, densely hairy. Leaves in whorls of 3—5, coriaceous to

thickly coriaceous, shiny above, glabrous on both surfaces or sparsely hairy

below, drying red-brown on both surfaces, darker above, sometimes

80 Gard. Bull. Singapore 57 (2005)

glaucous below; blade usually elliptic or ovate, rarely obovate, 5.5—16 x

2.5—8.5 cm, base acute to broadly acute, margin revolute, apex acuminate,

acumen 0.5—1.5 cm long; midrib raised on both surfaces, stronger below,

glabrous or sparsely hairy below; lateral veins 4—7(—10) pairs, dense, at an

angle of 45°—50° from the midrib, flat above, slightly raised below,

disappearing towards the margin, sometimes joining near the margin on

the upper half of leaf; tertiary venation slightly or distinctly prominent

above, slightly prominent below, reticulate or pitted; petiole (0.7—)1—2.5

cm long, drying dark brown. Inflorescences umbellate, axillary or borne

along twigs between the whorls of leaves, sessile; vegetative terminal bud

absent; bracts rounded, ovate or elliptic, 3—4 x 3 mm, densely hairy outside,

glabrous inside. Male flowers: pedicels 2-4 mm long; perianth lobes ovate

or elliptic, 2—3 x 2—2.5 mm, outer lobes slightly larger than the inner ones,

thin, densely hairy outside, glabrous inside; stamens sparsely hairy at base,

anthers 1-—1.2 mm long, filaments 0.8—1 mm long; pistillode c. 1.5 mm long.

Female flowers pedicels c. 2.5 mm long, perianth lobes obovate or rounded,

2—3 mm across, outer lobes slightly larger than the inner ones, thin, densely

hairy outside, glabrous inside; ovary ovoid, c. 0.5 x 0.2 mm, style thick,

stigma 0.5—1 mm across, densely hairy; staminodes spathulate, 1.5—2 mm

long. Infructescences each bearing 3—5 fruits; vegetative terminal buds

absent. Fruits globose, 0.5—0.8 cm diam., fleshy, drying black; cupule saucer-

shaped, 0.3—0.5 cm across, remnant of perianth lobes absent; pedicels to 5

mm long. Seeds globose, 0.3—0.4 cm diam., drying black.

Distribution: Endemic in Borneo (Sabah and Sarawak).

Ecology: In montane and limestone forests and also forest on ultrabasic

soil at altitudes to 2300 m.

Notes: The species resembles the large-leaved Actinodaphne borneensis

but can be differentiated by its glabrous (vs. sparsely hairy below) leaves

with a broadly acute or truncate (vs. acute) base, dense (vs. lax or well-

spaced) lateral veins and reticulate (vs. scalariform) tertiary veins. The

leaves of this species are highly variable in size and shape: the larger leaves

usually ovate, the medium-sized ones either broadly elliptic or almost

rounded and the small-sized leaves elliptic, ovate or slightly obovate. In

1968, Kostermans annotated four specimens (Clemens 20620, Clemens

31388, Clemens 40072, Clemens s.n.) as a new species, A. percoriacea, but

he never validated the name of the new taxon.

Other specimens examined: BORNEO - SABAH, Labuk Sugut District,

Gunung Tawai, Sugau SAN 138834 (SAN), Ranau District, Colombon

Actinodaphne in Borneo 81

Basin, Clemens 40072 (BO), Clemens 28933 (BO), Clemens 30226 (BO),

Kamborongo, Henry SAN 38309 (SAN), Kinabalu National Park, Carr

SFN 27725 (SING), Panar Laban, Sato 1099 (SAN), Kostermans SAN

38469 (SAN), Meijer SAN 46520 (KEP, SAN, SAR), Sinclair & Kadim

9045 (BO), Penibukan, Clemens 31388 (BO), Clemens s.n. (BO), Rao 134

(SING), Wood & Wyatt-Smith SAN A 4491 (KEP, SING), Pig Hill,

Barkman 107 (KNP), Tenom District, Mount Tomanis, Dolois et al. SP

15067 (KNP); SARA WAK, Bau District, Bidi Cave, Clemens 20620 (SAR),

Ridley s.n. (SING), Lawas District, Gunung Murud, Julaihi et al. S 80034

(KEP, SAR), S 80093 (KEP, SAR).

13. Actinodaphne pruinosa Nees

Actinodaphne pruinosa Nees in Wall. Pl. As. Rar. 2 (1831) 68; Gamble, J.

As. Soc. Beng. 75, 1 (1912) 119; Ridley, Fl. Malay Penins. 3 (1924) 110;

Anderson, Checkl. Trees Sarawak (1980) 220; Kochummen, Tree FI.

Malaya 4 (1989) 106; Turner, Gard. Bull. Sing. 47 (1995) 274; Coode et

al. (eds.), Checkl. Flow. Pl. Gymno. Brunei (1996) 149; Beaman et al.,

Pl. Mt. Kinabalu (2001) 394 (p.p., excl. syn. Actinodaphne foxworthyana

Gibbs). Type: Wallich Cat. 2584b, Peninsular Malaysia, Penang (holo

K; iso BO, L).

Actinodaphne pruinosa var. kunstleri Gamble, J. As. Soc. Beng. 75, 1 (1912)

120; Ridley, Fl. Malay Penins. 3 (1924) 110; Burgess, Sabah For. Record

6 (1966) 330; Turner, Gard. Bull. Sing. 47 (1995) 274. Type: King’s

collector 6063, Peninsular Malaysia, Perak, Larut (holo K).

Syn. noyv.: Actinodaphne concinna Ridl., J. Fed. Mal. States Mus. 5 (1914)

44, Fl. Mal. Pen. 3 (1924) 110; Kochummen, Tree Fl. Malaya 4 (1989)

104; Turner, Gard. Bull. Sing. 47 (1995) 273. Type: Robinson s.n.,

Peninsular Malaysia, Selangor, Gunung Mengkuang Lebah (holo K; iso

SING)

Distribution: Peninsular Malaysia, Singapore and Borneo (Brunel,

Kalimantan, Sabah and Sarawak).

Ecology: In peat swamp, kerangas, mixed dipterocarp, submontane to upper

montane forests at altitudes to 4000 m.

Notes: The species closely resembles Actinodaphne oleifolia and A.

borneensis but differs from both in having characters discussed under the

notes for A. oleifolia and A. borneensis.

82 Gard. Bull. Singapore 57 (2005)

14. Actinodaphne robusta S Julia, sp. nov.

(Latin, robustus=robust; referring to the leaves)

Actinodaphne macrophyllae in habitu similis, foliis obovatis ad late ellipticis

(vs oblanceolatis ad elliptice oblongis), venis lateralibus valde ascendentibus

sed paucioribus differt. Typus: /lias S$ 41159, Borneo, Sarawak, Kapit

District, Ulu Sampurau, Bukit Sampandai (holo SAR; iso KEP, SAN).

Tree 12—33 m tall, 30—35 cm diam.; bole straight, c. 7.5 m tall. Bark brown,

scaly; inner bark red-brown, granular. Sapwood whitish. Twigs drying dark

brown or greyish, sparsely or densely hairy, smooth. Terminal buds: scales

elliptic or ovate, 15—20 x 10—15 mm, densely hairy. Leaves in whorls of 5-

7, coriaceous to thickly coriaceous, glabrous above, densely hairy below,

drying dark brown on both surfaces or greenish brown below; blade usually

broadly obovate or rarely elliptic, (16.5—)22—40(—49) x 7.5-14(—18) cm,

base acute to attenuate, margin strongly to slightly revolute, apex acuminate,

acumen ().5—1 cm long; midrib raised on both surfaces, stronger below,

glabrous or sparsely hairy above, densely hairy below; lateral veins 7—15

pairs, lax, at an angle of 45°—50° from the midrib, flat or sunken above,

strongly raised below, joining near the margin on the upper half of leaf;

tertiary venation obscure or slightly distinct and impressed above, distinct

below, distantly scalariform; petiole 2.5—6 cm long, drying brownish or

dark brown, densely hairy. Inflorescences glomerulate, borne along twigs

between whorls of leaves; peduncle c. 3 mm long; vegetative terminal bud

absent; bracts ovate or elliptic, c. 4x 3 mm, densely hairy outside, glabrous

inside. Male flowers unknown. Female flowers: pedicels 2—3 mm long;

perianth lobes ovate-rounded, 2—2.5 mm across, outer lobes slightly larger

than the inner ones, thin, densely hairy outside, glabrous inside; ovary

ovoid, 2-3 mm, style thick, stigma 1—2 mm across, densely hairy; staminodes

spathulate, 1.5—2.5 mm long. Infructescences each bearing 2—8 fruits;

vegetative terminal bud present. Fruits globose, 1—1.8 cm diam., fleshy,

yellowish when mature, drying black or dark brown; cupule saucer-shaped,

0.7—1.5 cm across, remnant of perianth lobes absent or sometimes present;

pedicels 4-5 mm long. Seeds globose, 1.2—1.5 cm diam., drying black.

Distribution: Peninsular Malaysia, Borneo (Kalimantan and Sarawak) and

the Philippines.

Ecology: In mixed dipterocarp forest, at altitudes to 1100 m.

Notes: The new species resembles Actinodaphne macrophylla but differs in

having obovate or broadly elliptic leaves (vs. oblanceolate or elliptic-oblong)

Actinodaphne in Borneo 83

and markedly ascending but fewer lateral veins.

Other specimens examined: PENINSULAR MALAYSIA -— Johore, Labis

Forest Reserve, Saw FRI 36355 (KEP), Mersing, Teo & Din KL 4940

(KEP), Pahang, Jerantut, Taman Negara, Lata Berkoh, Ang FRI 23426

(KEP); BORNEO — SARAWAK, Miri District, Bakong, Ulu Mamut, J//ias

S 24362 (KEP, SAN, SING); EAST KALIMANTAN, Gunung Ilas

Bungaan, Kostermans 13741 (BO).

15. Actinodaphne semengohensis S.Julia, sp. nov.

(of Semengoh Forest Reserve, Sarawak)

Actinodaphne pruinosae similis, foliis oblanceolatis ad anguste ellipticis

(vs. ellipticis ad obovatis) in verticillis non strictis (vs. 3—6 in verticillis

strictis) dispositis, inflorescentiis masculis umbellatis ad glomerulatis (vs.

racemosis) facile distinguenda. Typus: Anderson & Asah § 12724, Borneo,

Sarawak, Kuching District, Semengoh Forest Reserve (holo SAR; iso BO,

SAN, SING).

Figure 1

Tree 15—24 m tall, 18—50 cm diam.; bole straight, c. 20 m tall; buttresses

absent. Bark brown, smooth; inner bark ochre or brownish, fibrous.

Sapwood yellowish or whitish. Twigs drying greyish brown or black, sparsely

hairy, smooth. Terminal buds: scales ovate, 3—4 x 2—3 mm, densely hairy.

Leaves not strictly in whorls, coriaceous, glabrous above, glabrous or

sparsely hairy below, drying dark brown above, glaucous below; blade

oblanceolate or narrowly elliptic, (S—)7.5-9.5(—12) x (1.8—)2-2.5(—3) cm,

base sharply acute, margin flat, apex acute or acuminate, acumen 0.5—1.5

cm long; midrib raised on both surfaces, stronger below, glabrous above,

sparsely hairy below; lateral veins 4—6 pairs, lax, at an angle of 30°—40°

from the midrib, slightly raised on both surfaces, inconspicuously joining

towards the margin; tertiary venation obscure on both surfaces; petiole 1—

2 cm long, drying dark brown. Male inflorescences racemose, axillary or

borne along twigs between the whorls of leaves; peduncle 3—5 mm long;

vegetative terminal bud absent; bracts ovate-rounded, 3—4.5 x 2—2.5 mm,

densely hairy outside, glabrous inside. Male flowers: pedicels 1—1.5 mm

long; perianth lobes elliptic or ovate-rounded, 1.5—2 x 1—1.8 mm, outer

lobes slightly larger than the inner ones, thin, densely hairy outside, glabrous

inside; stamens sparsely hairy at base, anthers 1—1.5 mm long, filaments c.

0.5 mm long; pistillode absent. Female inflorescences, flowers and fruits

unknown.

$4 Gard. Bull. Singapore 57 (2005)

Distribution: Endemic in Borneo (Sarawak and Sabah).

Ecology: In primary mixed dipterocarp and submontane forests at altitudes

to 1500 m.

Notes. The species resembles Actinodaphne pruinosa but can be

distinguished by its oblanceolate or narrowly elliptic (vs. elliptic or obovate)

leaves arranged in pseudo-whorls (vs. in whorls) and racemose (vs.

umbellate or glomerulate) male inflorescences. (Although the original

description of A. pruinosa described the inflorescence as racemose, at least

in the male inflorescence, racemose inforescences have not been observed

in Bornean specimens, which are consistently umbellate). It is also

reminiscent of A. oleifolia but differs by its pseudo-whorled (vs. whorled)

leaves with obscure (vs. distinctly pitted or reticulate) tertiary veins and

racemose (vs. umbellate) male inflorescences.

Other specimens examined: BORNEO — SABAH, Keningau District, Trus

Madi, Meijer SAN 122587 (SAN), Ranau District, Kinabalu National Park,

Chow & Madani SAN 74527 (SING), Meijer SAN 57510 (SAN);

SARA WAK, Kuching District, Semengoh Forest Reserve, Bojeng S 14620

(BO, SAN, SAR, SING), Rosli S 16425 (BO, SAN, SAR, SING), Rosli S

16453 (BO, SAN, SAR, SING), Othman Ismawi S 57220 (SAN, SAR).

16. Actinodaphne soepadmoi S.Julia, sp. nov.

(Engkik Soepadmo, 1993—present, Coordinator/Chief Editor of the Tree

Flora of Sabah and Sarawak Project)

Actinodaphne glabram approximata, foliis late obovatis (vs. ellipticis vel

elliptice oblongis vel oblanceolatis) indice longitudinis/latitudinis 2:1 (vs.

3—4:1) facile distinguenda. Typus: Julia & Sirukit S 91375, Borneo, Sarawak,

Kuching District, Semengoh Forest Reserve (holo SAR; iso KEP).

Figure 1

Tree 20-30 m tall, 15—40 cm diam.; bole straight, c. 15 m tall; buttresses

0.5—0.6 m tall, c. 6 cm wide, not spreading. Bark blackish or reddish brown,

smooth or with sparse rings; inner bark brownish, granular. Sapwood

yellowish. Twigs drying blackish, sparsely hairy, sparsely lenticellate.

Terminal buds: scales lanceolate, 10-15 x 4-6 mm, densely hairy. Leaves

in whorls of 5—6, thickly coriaceous, shiny above, glabrous on both surfaces

or sparsely hairy below, drying brown on both surfaces or greenish brown

above, glaucous below; blade plane, obovate, 14.5—20.5 x 4.5—9 cm, base

acute, margin strongly revolute, apex rounded or bluntly acute; midrib

Actinodaphne in Borneo 85

Figure 1. Actinodaphne semengohensis. A, leafy twig; B, male inflorescence; C, male flower;

D, male flower showing stamens; E, longitudinal section of male flower; F, bract; G & H,

perianth lobes; I, stamen with glands; J, stamen without gland (all from S 12724)

86 Gard. Bull. Singapore 57 (2005)

sunken above, strongly raised below; lateral veins 6—8 pairs, lax, at an

angle of 40°—45° from the midrib, slightly sunken above, strongly raised

below, disappearing towards the margin, sparsely hairy on both surfaces;

tertiary venation distinct on both surfaces, closely scalariform; petiole 2—6

mm long, drying dark brown, sparsely hairy. Male inflorescences and flowers

unknown. Female inflorescences glomerulate, borne along twigs between

the whorls of leaves; peduncle 5—14 mm long; vegetative terminal bud

present; bracts ovate or rounded, 3.5—5 x 5—6 mm, sparsely hairy outside,

glabrous inside. Female flowers: pedicels 2—3 mm long; perianth lobes

ovate, 2—4 x 1.5—4 mm, outer lobes slightly larger than the inner ones,

thin, densely hairy outside, glabrous inside; ovary ovoid, c. 1.5 x 0.5 mm,

style thick, stigma 0.8—1.5 mm across, densely hairy; staminodes spathulate,

1—1.5 mm long. Infructescences each bearing 1—4 fruits; vegetative terminal

bud present. Fruits globose, 0.7—1.5 cm diam., fleshy, yellow turning to red

when mature, drying black; cupule sometimes swollen, saucer-shaped, 0.6—

1.3 cm across, remnant of perianth lobes absent; pedicel 5-12 mm long.

Seeds globose, 0.8—1 cm diam., drying dark brown.

Distribution: Peninsular Malaysia and Borneo (Sarawak).

Ecology: In mixed dipterocarp forest at altitudes to 300 m.

Notes: Even though the male inflorescence and flower are not available,

this species is placed in Actinodaphne based on the combination of its

vegetative characters (perulate buds, bud scales leaving scars above the

whorled leaves) and the characters of its female inflorescence and trimerous

flower. |

Actinodaphne soepadmoi resembles A. glabra but differs in having

obovate (vs. oblanceolate or narrowly elliptic) leaves. The new species is

also reminiscent of Actinodaphne myriantha but can be distinguished by its

obovate (vs. obovate-elliptic or narrowly elliptic) leaves having an obtuse

or almost rounded (vs. acute or acuminate) apex.

Kochummen (1989) identified and described a single collection

(Ridley 16125) from Gunung Tahan, Peninsular Malaysia as A. obovata. In

comparing the specimen with the type image and description of A. obovata

(Wall. ex Nees) Blume, it is concluded that the specimen does not belong

to A. obovata but to A. soepadmoi.

This new species is named in honour of Dr. E. Soepadmo for his

dedication and contribution to the knowledge of the Tree Flora of Sabah

and Sarawak.

Other specimens examined: PENINSULAR MALAYSIA - Pahang,

Actinodaphne in Borneo 87

Figure 2. Actinodaphne soepadmoi. A, fruiting leafy twig; B, female inflorescence; C, female

flower; D, longitudinal section of female flower showing pistil and staminodes: E, longitudinal

section of female flower; F-G, perianth lobes; H, staminodes: I, pistil: J, fruit: K, longitudinal

section of fruit (A from § 37704, B-I from § 91375, J & K from S 45548)

88 Gard. Bull. Singapore 57 (2005)

Gunung Tahan, Ridley 16125 (SING). BORNEO - SARAWAK, Kuching

District, Semengoh Forest Reserve, lias § 37704 (KEP, SAN, SAR), Belaga

District, Sungai Iban, Bernard S 45548 (KEP, SAN, SAR).

17. Actinodaphne spathulifolia S Julia, sp. nov.

(Latin, spathulatus=spathula-shaped, folium=leaf)

Actinodaphne fuliginosam revocans, foliis oblanceolatis (vs. obovatis)

maioribus (5—7.5 cm longis, 2.5—4 cm latis, vs. 2.5—4.5 cm longis, 1.5—2.5

cm latis), costa venis lateralibus tertiariisque conspicuis (vs. obscuris) differt.

Typus: Latiff et al. ALM 4178, Borneo, Sarawak, Marudi District, Kelabit

Highlands, Bario (holo SAR).

Tree or treelet 4-33 m tall. Twigs drying black, densely hairy when young,

sparsely hairy when older, smooth. Leaves in whorls of 4—5, thickly

coriaceous, glabrous on both surfaces, drying shining and greenish brown

above, greyish brown below; blade oblanceolate, 5—7.5 x 2.5—4 cm, base

cuneate or sharply acute, margin strongly revolute, apex rounded; midrib

sunken above, raised below, sparsely hairy below; lateral veins 5—6 pairs,

lax, at an angle of c. 40° from the midrib, sunken above, raised below,

disappearing towards the margin; tertiary venation obscure on both surfaces;

petiole 1.2—1.5 cm long, drying black. Male inflorescences and flowers

unknown. Female inflorescences glomerulate, axillary or borne along twigs

between the whorls of leaves; vegetative terminal bud absent; bracts elliptic,

2.5—3 x 2—2.2 mm, sparsely hairy outside, glabrous inside. Female flowers

(young): pedicels 1.8—2 mm long; perianth lobes ovate-rounded, 1—1.5 x

1—1.2 mm, outer lobes slightly larger than the inner ones, thin, densely

hairy outside, glabrous inside; ovary ovoid, c. 0.8 x 0.4 mm, style c. 0.4 mm

long, stigma c. 0.3 mm across, densely hairy; staminodes spathulate, 0.8—1

mm long. Infructescences and fruits unknown.

Distribution: Endemic in Borneo (Sabah and Sarawak).

Ecology: In kerangas and submontane forests at altitudes to 1800 m.

Notes: Even though the male inflorescence and flower are not available, this

species is placed in Actinodaphne based on the combination of its vegetative

characters (perulate buds, bud scales leaving scars above the whorled leaves)

and the characters of its female inflorescence and trimerous flowers.

The species resembles A. fuliginosa but differs by its oblanceolate

(vs. obovate) and broader, 2.5—4 cm wide (vs. 1.5—2.5 cm) leaves with

prominent (vs. obscure) midrib, lateral veins and tertiary veins.

Actinodaphne in Borneo 89

Other specimens examined: BORNEO - SABAH, Ranau District, Kinabalu

National Park, Kitayama 4480 (KNP); SARA WAK, Marudi District, Kelabit

Highlands, Bario, Nooteboom & Chai 2150 (KEP, SAR).

18. Actinodaphne sulcata S. Julia, sp. nov.

(Latin, su/catus=grooved; referring to the midrib and lateral veins on upper

leaf surface)

Actinodaphne kinabaluensi foliis bullatis similis, foliis obovatis (vs. ellipticis),

fructibus globosis (vs. ellipsoideis) differt. Typus: Nooteboom & Chai 2245,

Borneo, Sarawak, Marudi District, Kelabit Highlands (holo SAR; iso KEP).

Tree or treelet 3—10 m tall, 1.5—17 cm diam.; bole straight, c. 4 m tall. Bark

creamy or brownish, smooth; inner bark greenish, granular. Sapwood

yellowish or whitish. Twigs drying greyish or dark brown, sparsely or densely

hairy when young, sparsely hairy when older, smooth or sparsely large-

lenticellate. Terminal buds: scales ovate, 5—10 x 3—5 mm, densely hairy.

Leaves in whorls of 3—5, thickly coriaceous, glossy and glabrous above,

sparsely hairy below, drying red-brown or greenish brown above, brown

below; blade bullate, obovate, 10—15(—19) x 4~7(—8.5) cm, base sharply

acute, margin revolute, apex acuminate, acumen ().5—1.5 cm long; midrib

sunken or flat above, strongly raised below, glabrous or sparsely hairy

below; lateral veins 7—10 pairs, lax, at an angle of 40°—45° from the midrib,

strongly sunken above, strongly raised below, disappearing towards the

margin or joining near the margin in the upper half of leaf; tertiary venation

distinct on both surfaces or sunken above, distantly scalariform; petiole

0.6—1(—5) cm long, sparsely or densely hairy, drying dark brown or black.

Inflorescences umbellate (female) or racemose (male), axillary or borne

along twigs between the whorls of leaves, subsessile; vegetative terminal

bud absent; bracts ovate-rounded, 3—4 x 3.5—4 mm, sparsely hairy outside,

glabrous inside. Male flowers: pedicels c. 5 mm long; perianth lobes elliptic

or elliptic-oblong, 3—3.5 x 1.2—2.2 mm, outer lobes slightly larger than the

inner ones, thin, densely hairy outside, glabrous inside; stamens sparsely

hairy at base, anthers 0.8—1.2 mm long, filaments 0.8—1.2 mm long; pistillode

0.6—0.8 mm long. Female flowers: pedicels c. 5 mm long, perianth lobes

elliptic or oblong, 3—3.5 x 1.5—2.2 mm, outer lobes slightly larger than the

inner ones, densely hairy outside, glabrous inside; ovary ovoid, c. 2 x 1.5

mm, style thick, stigma 0.8—1 x 0.8 mm, densely hairy; staminodes spathulate,

1.5—2.5 mm long. Infructescences each bearing 1—5 fruits; vegetative

terminal bud absent. Fruits globose, 0.6—0.8 cm diam., drying black; cupule

swollen, saucer-shaped, 0.5—0.8 cm across, remnant of perianth lobes absent;

pedicel 5—8 mm long. Seeds globose, 0.3—0.5 cm diam., drying dark brown.

90 Gard. Bull. Singapore 57 (2005)

Distribution: Endemic in Borneo (Sabah and Sarawak).

Ecology: In mixed dipterocarp and submontane forests at altitudes to 1400 m.

Notes: The species resembles Actinodaphne kinabaluensis in its bullate

leaves but differs in having globose fruits (vs. ellipsoid) and obovate leaves

(vs. elliptic leaves).

In Sabah and Sarawak two varieties, var. sulcata and var. longipetiolata,

are known.

var. sulcata

Distribution: Endemic in Borneo (Sabah and Sarawak). In mixed

dipterocarp and submontane forests, at altitudes 1200-1400 m.

Other specimens examined: BORNEO — SABAH, Sipitang District, Bukit

Rimau, Pius & Ubaldos SAN 143457 (KEP, SAN, SAR, SING), Long

Miau, Meligan, Dewol & Kambira SAN 141813 (SING); SARA WAK, Lawas

District, Ulu Sungai Belaban, Gunung Murut, Ilias S 26424 (SAR).

var. longipetiolata S Julia, var. nov.

(Latin, /ongi=long, petiolatus=petiole)

A var. typica foliis petiolisque longioribus, 3—5 cm longis (vs. 0.6—1 cm

longis) differt. Typus: Julius et al. SAN 132805, Borneo, Sabah, Sipitang

District, Maligan Forest Reserve (holo SAN; iso KEP, SAR).

Distribution: Endemic in Borneo (Sabah) and known only by the type

specimen. Found in mixed dipterocarp forest, at c. 1400 m altitude.

Notes: This variety can be distinguished from var. sulcata by its longer (3—

5 cm vs. 0.6—1 cm), sparsely hairy (vs. densely hairy) petiole.

19. Actinodaphne venosa S.Julia, sp. nov.

(Latin, venosus=conspicuously veined; the leaves)

Actinodaphne malaccensi e Malaysia peninsulari similis, ramulis glabris

(vs. dense ferrugineo-tomentosis), foliis ellipticis ad oblanceolatis (vs.

elliptice oblongis ad lanceolatis) venis tertiariis conspicuis (vs. obscuris ad

inconspicuissimis) facile distinguenda. Typus: Ampuria SAN 32859, Borneo,

Sabah, Beluran District, near Tidog Camp (holo SAN; iso BO, KEP, SING).

Actinodaphne in Borneo 9]

Tree or treelet, (2—)4—35 m tall, (5—)15—45 cm diam.; bole straight, 2—9 m

tall; buttresses absent. Bark brownish, greenish or greyish, smooth; inner

bark yellowish, fibrous. Sapwood yellowish. Twigs drying dark brown or

greyish brown, densely hairy when young, sparsely hairy when older, smooth

or sometimes fissured or sparsely large-lenticellate. Terminal buds: scales

ovate, lanceolate or linear, 3—5 x 1—2 mm, densely or sparsely hairy. Leaves

in whorls of 3—5, thinly coriaceous, sometimes shiny above, glabrous on

both surfaces or sparsely hairy below, drying brownish, greenish brown or

reddish brown on both surfaces or glaucous below; blade obovate or elliptic,

9.5—16.5(—18) x 3—6(—9) cm, base acute, margin flat or slightly revolute,

apex acuminate or shortly cuspidate, acumen c. | cm long; midrib sunken

or flat above, strongly raised and sparsely hairy below; lateral veins 4—7(—

9) pairs, dense, at an angle of 30°—40° from the midrib, slightly or strongly

sunken above, strongly raised below, joining towards the margin; tertiary

venation distinct on both surfaces, sunken above, closely scalariform; petiole

1.2—2(—2.5) cm long, drying brown or black, sparsely hairy. Inflorescences

umbellate, borne along twigs between the whorls of leaves, sessile;

vegetative terminal bud absent; bracts rounded or ovate, 1.5—3 x 1—2 mm,

sparsely to densely hairy outside, glabrous inside. Male flowers: pedicels

2—4 mm long; perianth lobes ovate or elliptic, 1—2 x 0.8—1 mm, outer lobes

slightly larger than the inner ones, thin, densely hairy outside, glabrous

inside; stamens sparsely hairy at base, anthers c. 1.5 mm long, filaments c.

1.5 mm long; pistillode c. 1.5 mm long. Female flowers: pedicels 1-2 mm

long, perianth lobes ovate or elliptic, 1—2 x 0.5—1 mm, outer lobes slightly

larger than the inner ones, thin, densely hairy outside, glabrous inside;

ovary ovoid, c. 1 x 0.5 mm, style thick, stigma c. 0.5 mm across, densely

hairy; staminodes spathulate, c. 1 mm long. Infructescences each bearing

1—4 fruits; vegetative terminal bud absent. Fruits globose, 0.6—1.5 cm diam.,

fleshy, drying black; cupule saucer-shaped, 0.5—0.8 cm across, remnant of

perianth lobes present or absent; pedicel 3—7 mm long. Seeds globose,

0.5—1 cm diam., drying black.

Distribution: Endemic in Borneo (Brunei, Kalimantan, Sabah and Sarawak).

Ecology: In mixed dipterocarp, submontane, riparian forests and forest on

ultrabasic soil at altitudes to 1300 m.

Notes: The new species resembles Actinodaphne malaccensis Hook.f. from

Peninsular Malaysia but can readily be distinguished by its glabrous (vs.

densely rusty-tomentose) twig, elliptic or oblanceolate (vs. elliptic-oblong

or lanceolate) leaves with distinctly prominent (vs. obscure or very faintly

visible) tertiary veins. The species also resembles A. borneensis and A.

92 Gard. Bull. Singapore 57 (2005)

pruinosa but differs from the former by its thinly coriaceous (vs. coriaceous

or thickly coriaceous) leaves, dense (vs. lax or well-spaced) lateral veins

and distinct (vs. obscure) tertiary veins, and from the latter in having

leaves with a length:width ratio of 2:1 (vs. length: width ratio of 3:1) and

sunken lateral veins (vs. flat) above. In vegetative characters the species is

also close to A. montana Gamble from Peninsular Malaysia but differs by

its strongly impressed (vs. raised) lateral veins on upper surface of leaves

and umbellate (vs. paniculate or racemose) inflorescence.

Other specimens examined: BORNEO - SABAH, Beaufort District, Klias,

Meijer SAN 31411 (SAN), Lumat, Daud & Karim SAN 77929 (SAN),

Kinabatangan District, Maliau Basin, Ming & Sidkan, MB &14 (SAN), Ulu

Sungai Pingas-pingas, Sumbing SAN 110950 (SAN), Kuala Penyu District,

Klias, Meijer SAN 31411 (SAN), Labuk Sugut District, Beluran, Ampuria

SAN 32859 (BO, KEP, SAN, SING), Lungmanis, Wood A 3994 (KEP,

SING), Lahad Datu District, Binuang, Nordin SAN 54576 (SAN), Danum

Valley, Dewol SAN 129455 (SAN), Dewol SAN 134978 (SAN), Campbell

214 (SAN), Gunung Silam, Agam, SAN 37165 (SING), Kalumpang Forest

Reserve, George et al. SAN 123904 (SAN), Agam SAN 40863 (BO, SAN),

SAN 37165 (SAN, SING), Madai Forest Reserve, Nordin SAN 47763

(SAN), Segama, Bukit Belachan, Chai SAN 31722 (KEP, SAN, SAR,

SING), Ulu Sungai Tabin, Dewol SAN 129528 (SAN), James SAN 35358

(SAN), Ranau District, Gunung Kinabalu, Clemens 31728 (BO), Clemens

32292 (BO), Semporna District, Mount Pock Forest Reserve, Nordin SAN

54463 (SAN), Tambunan District, Trus Madi, Kamaruddin & Latiff KMS

1782 (SAN), Tawau District, Elmer 20947 (BO, PNH, SING), Betoton,

Orolfo BNB 3221 (PNH); SARAWAK, Bau District, Mamit S 29592 (BO,

SAR), Kapit District, Bukit Kumbong, Runi et al. S 60068 (SAN, SAR),

Belaga District, Upper Rejang, Clemens 21861 (BO, SAR), Kapit District,

Ulu Sungai Melatai, Yii S 48364 (SAR), Kuching District, Padawan, Gunung

Siruruh, Yii S$ 55270 (SAN, SAR), Bako National Park, Telok Asam,

Purseglove P 4973 (SAR), Limbang District, Sungai Ensungei, Rena et al.

S 42875 (KEP, SAN, SAR), Rena et al. S 42934 (KEP, SAN, SAR), Lubok

Antu District, Nanga Sumpa, Christensen 1493 (SAR), Marudi District,

Gunung Mulu National Park, Chai S 39750 (KEP, SAN, SAR), Chai S$

39667 (KEP, SAN, SAR), Yii & Talib S 58659 (KEP, SAR), Sri Aman

District, Lingga, G. Lesung, Hansen 1019 (KEP, SAN, SAR); BRUNET,

Belait District, Melilas, Atkins et al. SA 539 (KEP, SAN, SING);

KALIMANTAN, Without locality, Kostermans 8977 (BO) and Kostermans

s.n. (BO), Kutai, Belajan River, Endert 1927 (BO) and Sungai Wain,

Kostermans 4328 (BO).

Actinodaphne in Borneo 93

Incompletely Known Species

20. Actinodaphne sp. 1

Twigs drying black, densely hairy when young, sparsely hairy when older,

smooth. Terminal buds: scales ovate, c. 3 x 2 mm, sparsely hairy. Leaves in

whorls of 3—5, thickly coriaceous, shiny and glabrous above except on the

midrib and lateral veins, sparsely hairy below, drying greyish brown above,

brown below; blade elliptic, 7.5—10 x 3—4 cm, base acute, margin strongly

revolute, apex acute; midrib raised on both surfaces, stronger below; lateral

veins 4—5 pairs, lax, at an angle of c. 45° from the midrib, flat or sunken

above, raised below, disappearing towards the margin; tertiary venation

distinct on both surfaces, sunken above, distantly scalariform; petiole 0.8—

1 cm long, drying black, densely hairy, glabrescent. Male inflorescences

and flowers unknown. Female inflorescences racemose, borne along twigs

between the whorls of leaves; peduncle 3—5 mm long, densely hairy;

vegetative terminal bud present; bracts ovate, c. 0.3 x 0.2 mm, densely

hairy outside, glabrous inside. Female flowers: pedicels 2—3 mm long;

perianth lobes elliptic, c. 1.5 x 1 mm, outer lobes slightly larger than the

inner ones, thin, densely hairy outside, glabrous inside; ovary ovoid, c. 0.5

x 0.3 mm, style thick, c. 1 mm long, stigma c. 0.4 mm across, densely hairy;

staminodes spathulate, c. | mm long. Infructescences and fruits unknown.

Distribution: Known only from a single collection (Chew et al. RSNB 1840)

from Gunung Kinabalu, Sabah.

Notes: This taxon resembles Actinodaphne kinabaluensis but differs in

having plane (vs. bullate) leaves, shorter petiole and yellowish (vs. brownish)

hairs on the leaf undersurface. The species is placed in Actinodaphne based

on the combinations of its vegetative (perulate buds, bud scales leaving

scars above the whorled leaves) and reproductive characters (trimerous

flowers). However, to formally describe the specimen as a new species

more specimens are needed to elucidate its taxonomic status, as there is a

possibility that this incompletely known species may represent a variety of

the already known species.

Acknowledgements

This synopsis represents a precursory paper of the revision of Actinodaphne

for the Tree Flora of Sabah and Sarawak Project funded by the Malaysian

Ministry of Science, Technology and Innovation (MOSTI) under the IRPA

94 Gard. Bull. Singapore 57 (2005)

Research Grant No. 08-04-01-0165. I wish to extend my gratitude for the

support and encouragement given by the Director-General of the Forest

Research Institute Malaysia and the Directors of the Forestry Departments

of Sabah and Sarawak; to the Curators of the herbaria BO, K, KEP, L,

PNH, SAR, SAN, SING and the Kinabalu National Park, Sabah, for the

loan of specimens and images of type specimens and also for their

cooperation and hospitality during my visits to their institutions. I am

grateful to Dr. E. Soepadmo (KEP) and Dr. Ruth Kiew (SING) for their

constant guidance, critical comments and valuable advice during the

preparation of this paper. Dr. J.F. Velkamp (L) is thanked for preparing

the Latin diagnoses of the new taxa and Joseph Pao (SAR) for preparing

the botanical illustrations.

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Identification list of specimens examined

The number after the collector and/or serial number refers to the following

Actinodaphne taxa.

1 = A. borneensis Meisn. 11 =A. oleifolia Gamble

2 =A. diversifolia Merr. 12 = A. percoriacea S. Julia

3 =A. fuliginosa Airy Shaw 13 = A. pruinosa Nees

4= A. glabra Blume 14 = A. robusta S. Julia

5 = A. glomerata (Blume) Nees 15 = A. semengohensis S. Julia

6 = A. johorensis Gamble 16 = A. soepadmoi S. Julia

7 = A. kinabaluensis Kosterm. 17 = A. spathulifolia S. Julia

8a = A. kostermansii S. Julia var. kostermansii 18a = A. sulcata S. Julia var. longipetiolata S. Julia

8b = A. kostermansii S. Julia var. glabrescens S. Julia 18b = A. sulcata S. Julia var. sulcata

9 = A. macrophylla (Blume) Nees 19 = A. venosa S. Julia

10 = A. myriantha Merr. 20:= AL spd

Achmad 1356: 5 — Anderson s.n.: 1; Anderson s.n.: 4 — Ando et al. AKK

30: 9; AKK 42: 9 — Argent & Amiril 9320: 11 — Ariffin ARK 35: 5 —

Ariffin & Arbainsyah AA 977: 10 — Atkins et al. SA 439: 5; SA 450: 1; SA

539519:

Balajadia BNB 3061: 13; 7068: 5 — Barkman 107: 12 — Benedict SP 5939:

13 — BRUN 565: 2; BRUN 1465: 1; BRUN 2412: 5; BRUN 10384: 13;

BRUN 15026: 1; BRUN 15055: 5; BRUN 16933: 5; BRUN 16634: 5; BRUN

96 Gard. Bull. Singapore 57 (2005)

16209: 1; BRUN 18061: 13; BRUN 18285: 9.

Campbell EG 199: 5; EG 214: 19; 271:12 — Chew CWL 452: 4; CWL 507:

5; CWL 1212: 4 — Christensen 1493: 19 — Christensen & Apu 486: 13; 582:

13 — Church 424: 5; 473: 5 — Church & Mahyar 1879: 10 — Clemens 111:

4; 20042: 13; 20157: 13; 21859: 4; 21860; 4 21861: 1: 2666825227 eee

28814: 5; 29408:.5: 20620: 12; 28933: 12-.29116; ite 30008; 133-40776. 4

31159: 2; 31278: 1; 31364: 2;°31388: 12; 31493-1231 Za ee

33629: 11; 33798: 113351132 1k 30467: 13: 324357-13. 32490: 2 S26 ee

34340: 13; 37707: 13; 40072: 12: 50386: 8a; Clemens:s.n.: 12 — Coode ef ak

MC 7590: 11; MC 7869: 8a — Cousens 69770: 9 — Curtis 1020: 13.

Daim 302: 2; 635: 13; 639: 2; 643: 5; 673: 5; 6922 13; 966: 13'—= DeVore

1702: 5 — Dewanie 51: 2 — Dolois et al. SP 15067: 12 — Dransfield JD

7467: 2 — Dransfield et al. JD 7096: 5; JD 7171: 13.

Elmer 20947: 19; 21238: 10; 21335: 10; 21802: 10; 21809: 10 — Enchai 10382:

5 — Endert 1927: 19; 2249: 5; 3330: 9; 4733: 5; 5149: 10 — Enggoh 10421: 5.

Forman & Blewett LLF 805: 9 — FMS 17513: 13; FMS 20441: 11; FMS

34480: 4; FMS 36677: 5; FMS 37221: 4; FMS 37392: 9; FMS 37393: 13; FMS

41203: 5; FMS 41223: 5; FMS 42900: 5; FMS 48728: 5; FMS 49161: 5 — FRI

2036: 9; FRI 2073: 13; FRI 3237: 11; FRIG135..11 FRISD50: G PRIA

11; FRI 13229: 13; FRI 13725: 13; FRI 13889: 13; FRI 16998: 11; FRI 19870:

11; FRI 20395: 13; FRI 20684: 11; FRI 21864: 13; FRI 23426: 14; FRI 36355:

14; FRI 41338: 1; FRI 124124: 9.

Gadoh KL 837: 5 — Geh GSY 802: 5 — Geofarry et al. SP 17178: 2 —

Gibbs 3135: 2 — Gusili SP 9937: 13.

Hallier 1032: 5; 1059: 5; 1713: 13; B°1859: 13 — Hansen 801: 5; 1019: 19 —

Haviland 629: 5; Haviland 737: 5; Haviland 1756: 10; Haviland 1944: 13;

Haviland 2490: 13; Haviland 3328: 10 — Haviland ... Hose 3647A: 6 —

Henry & Sidkan MB 96: 11; MB 116: 11 — Hotta 13948: 5 — Hou 415: 10.

Jacobs 5086: 5 — John Sugau SIP-B 69: 4 — Junghuhns.n.: 5.

Kamaruddin KA 30: 2 — Kamaruddin & Latiff KMS 1782: 19 — Keith

7088: 5 — KEP: 10770: 2; KEP 20503: 13;-KEP. 51761213; KEP 55426...

KEP 64756: 13; KEP 66577: 11; KEP 78898: 13; KEP 84780: 9; KEP 85040:

13; KEP 85056: 13; KEP 94469: 13; KEP 94823: 13; KEP 99378: 13; KEP

104628: 13 — Kessler PK 917: 10 — King’s Collector 6172: 9; 6435: 5; 7349:

11 — Kinsun 469: 4 — Kirkup et al. DK 438: 1; DK 749: 1 — Kitayama

4480: 17 — Kokawa & Hotta 5603: 8b — Koorders 2788: 5; 3264: 5; 3619: 5;

3707: 5: 3718: 5: 5146: 5: 10960: 5: 1096668:;.5: 158078: 5S: 13924 s) tae

26772: 5: 30515: 5: 137814: 5 — Kostermans 3208: 9::4328:- 419: 4412-4:

4558: 10; 4696: 2: 4888: 12; 4957: 5: 5146: 5: 5285: 5: 3352242 5479: 3S ioo:

5859: 5; 5986: 9; 6007: 4; 6815: 4; 6954: 2; 7333: 13; 7361: 9; 7544: 13; 7560: 9;

7615: 13; 8116: 1; 8953: 4; 8789: 5; 8977: 19: 8942: 5; 9064: 13; 9069: 13; 9106:

10; 9242: 1; 9929: 5; 10054: 13; 1060825; 10676: 1: 12660: 3: 12S 75° ES. Pe:

Actinodaphne in Borneo 97

PieM3UGS: 132 15730: 5; 1347975; 13741: 1421013: 5; 21402: 5; 21640: 5;

23020: 5; 23932: 5; s.n.: 19 — Kuswata 964: 5; 1021: 1.

Latiff et al. ALM 4178: 17 — Leighton 13: 10; 133: 13 — Lim et al. LSP 1184: 4

- Lobb s.n.: 1 — Lomudin 510: 1 — Lorence Lugas 876: 4; 970: 13; 1922: 8b;

2387: 4.

Mahyar et al. 1172: 13 — Maingayi 1258: 9; 1275: 9 — McDonald & Ismail

4050: 4 — Ming & Sidkan, MB 814: 19 — Ming & Sidkan MB 814: 19 —

Mogea & de Wilde 3823: 13 — Mohd Nur 11232: 11 — Mohd. Shah 3502:

13 — Mohd Shah & Ahmad Shuker MS 3253: 4 — Mohizah & Yahud

ITTO/BC 69: 4 — Murata B-2600: 11.

Native Collector 1690: 13; 2174: 6 — Norazami AZ 17: 2 — Nooteboom &

Chai 1895: 4; 2150: 17; 2245: 18b — Orolfo BNB 3221: 19.

Pereira JTP 327: 2; JTP 372: 11 — Phillipps SNP 1926: 13; SNP 3029: 11 —

Podzorski SMHI 757: 14 — Poore H 308: 13 — Purseglove P 4973: 1.

Ramlanto & Fanani 678: 5 — Ramos 1159: 1; 1216: 5; 1838: 2 — Rao 134: 12

— Reksodihajo 750: 5 — Reza RA 129: 5 — Ridley 4419: 7; Ridley 6296: 5;

Ridley 6741: 5; 11675: 5; 13728: 11; 13783: 5; 16125: 16; Ridley s.7,: 12; s.n.:

5 — Ridsdale SMHI 21: 4; SMHI 1514: 14 — Rimi & Geofarry SP 8112: 11

— Rimi ef al. SP 6276: 8b; SP 6284: 11; SP 6311: 11; 7374: 13; SP 9289: 11;

TS 6573: 1 — Robinson s.n.: 11 — RSNB 13: 2; RSNB 137: 13; RSNB 196:

7, RSNB 1251: 2; RSNB 1840: 20; RSNB 4113: 5; RSNB 4383: 11; RSNB

4567: 5; RSNB 4660: 11; RSNB 4754: 11; RSNB 4832: 11.

» 97-10: S448: 10: S 145574: S 1583: 10; S 2179: 1:'S 4437: 6: S 4607: 11:8

Ay es Sats lo -.4- 8.6294" FS 9160713: S-12774: 15,9 1313F- 6; S 14620: 15:

164252152 Ss 10453: 15: S.16988: £3:-S 17624: 5; 818227: 4; 8S. 1872771; 8

1DOTo EE SH19114: 12S. 208742 105822104: 128'22182: 4: S 22783:-1;'S8

24362: 14: S 25437: 13; S 26424: 18b; S 27391: 10; S 28414: 9; S 28492: 5; 8

29597: 19S 35554: °5: S 33793: -13; S 34520: 5: S 34665: 10; S-34763: 5; S

35668: 1; S 37704: 16; S 38318: 1; S 38403: 5; S 39667: 19; S 39750: 19; S

39796: AS 42 ie 4: S 40159: 14:°S: 41423: 4: S 41895: 45 842735: 13; S

42875: 19; S 42934: 19; S 43657: 13; S 44704: 5; S 45548: 16; S 46397: 1; S

#0625. ); S 4/040: tt S°47657- 2: S 47783? 13;:S 48364: 19: 'S 49318: 5; S

Sol 1s SUE 428.505 90: 1 eS. 50767: 112 S 509038: 11: 8$-51430: 35; S

Se 78 552 702 19: SS 55604: 1h: S'55966: 11:-S 55968: 11; S 56471: 5; S

Doves dle S57 2202 19. S:974932 98:8 574412 10: S 57789: 5; S. 5786327 4;'S

58659: 19; S 59021: 1; S 60068: 19; S 61440: 5; S 62161: 4; S 64289: 5; S

G76: 42.5 65412-.6: S 66173: 5; S°66829: 5: S 68074: 13: S 71953: 11; S

Ponda Ss. 0 4442-> 5: 8S 74627421 S 7032429; S 76964: 13; S 78301: 9; S

80034: 12; S 80093: 12; S 80192: 11; S 80210: 13; S 81713: 5; S 81888: 13; S

87548: 6; S 88004: 14; S 88190: 13; S 91375: 16; S 94780: 5 — S A 906

(Egon): 13 — SAN A 470: 4; SAN A 1147 (Cuadra): 5; SAN A 1309

(Cuadra): 5; SAN A 2264: 5; SAN A 2797(Kadir): 5; SAN A 2892: 5; SAN

98 Gard. Bull. Singapore 57 (2005)

A 3302 (Kadir & Jiran): 5; SAN A 3568 (Kadir): 5; SAN A 3874: 1; SAN

A 3994: 19; SAN A 4491: 12; SAN A 4765: 10; SAN A 10512 (Enggoh): 5

— SAN 16576: 4; SAN 16856: 5; SAN 17570: 9; SAN 19213a: 5; SAN

20624: 2; SAN 21233: 13; SAN 21279: 5; SAN 21817: 1; SAN 22264: 5; SAN

22827: 9; SAN 23284: 1; SAN 23340: 5; SAN 25343: 1; SAN 25354: 4; SAN

26348: 5; SAN 26484: 4; SAN 27158: 5; SAN 27438: 4; SAN 28365: 5; SAN

29110: 13; SAN 30304: 4; SAN 30650: 2; SAN 31059: 5; SAN 31411: 19;

SAN 31614: 5; SAN 31722: 19; SAN 32029: 5; SAN 32859: 19; SAN 33160:

5; SAN 34426: 5; SAN 35088: 5; SAN 35358: 19; SAN 35650: 2; SAN 36546:

5; SAN 36583: 5; SAN 37056: 1; SAN 37165: 19; SAN 37683: 13; SAN

37821: 8b; SAN 38309: 12; SAN 38469: 12; SAN 38508: 13; SAN 38734: 5;

SAN 38823: 1; SAN 39269: 2; SAN 40863: 19; SAN 42206: 2; SAN 42400: 5;

SAN 43164: 4; SAN 43345: 5; SAN 44718: 5; SAN 46313: 5; SAN 46520: 12;

SAN 47327: 1; SAN 47402: 10; SAN 47763: 19; SAN 48181: 1; SAN 48287:

1; SAN 48992: 5; SAN 49248: 10; SAN 50065: 5; SAN 50564: 5; SAN 51016:

1; SAN 52516: 4; SAN 52789: 10; SAN 53552: 5; SAN 53921: 5; SAN 54463:

19; SAN 54576: 19; SAN 55793: 5; SAN 55839: 4; SAN 57104: 2; SAN

57143: 10; SAN 57510: 15; SAN 58090: 5; SAN 58182: 5; SAN 60576: 5;

SAN 61467: 4; SAN 62093: 1; SAN 62541: 1; SAN 64101: 8a; SAN 67566:

13; SAN 70344: 13; SAN 71505: 2; SAN 71760: 5; SAN 72002: 2; SAN

72064: 2; SAN 72067: 11; SAN 72771: 5; SAN 74278: 1; SAN 74279: 1; SAN

74338: 13; SAN 74527: 15; SAN 75940: 10; SAN 76177: 12; SAN 76181: 13;

SAN 76235: 4; SAN 77929: 19; SAN 78186: 2; SAN 78482: 2; SAN 79487: 2;

SAN 79943: 5; SAN 80030: 1; SAN 80201: 2; SAN 80226: 5; SAN 81172: 5;

SAN 81642: 5; SAN 81966: 1; SAN 82057: 5; SAN 82412: 11; SAN 84457: 5;

SAN 84766: 5; SAN 86368: 1; SAN 86369: 1; SAN 86759: 4; SAN 87194: 11;

SAN 87905: 10; SAN 87977: 10; SAN 89151: 5; SAN 90022: 13; SAN 90937:

5; SAN 92438: 5; SAN 93077: 5; SAN 93277: 2; SAN 98060: 8b; SAN 98112:

8b; SAN 98118: 8b; SAN 99879: 2; SAN 100721: 8b; SAN 100729: 8b; SAN

101957: 8b; SAN 102413: 2; SAN 103000: 1; SAN 105188: 1; SAN 106127:

5; SAN 106867: 11; SAN 108848: 2; SAN 110950: 19; SAN 111238: 2; SAN

113851: 13; SAN 114046: 2; SAN 114125: 13; SAN 114150: 2; SAN 114274:

2; SAN 114308: 5; SAN 116415: 13; SAN 116423: 13; SAN 117957: 5; SAN

118442: 13; SAN 118833: 13; SAN 120110: 2; SAN 120336: 11; SAN 122207:

13; SAN 122587: 15; SAN 122626: 11; SAN) 1228515 25> SANA23433243;

SAN 123918: 5; SAN 123904: 19; SAN 123986: 11; SAN 124769: 2; SAN

125360: 2; SAN 126094: 13; SAN 126440: 1; SAN 128051: 2; SAN 128635:

13; SAN 128848: 5; SAN 129455: 19; SAN 129528: 19; SAN 130104: 13;

SAN 130202: 2; SAN 132242: 4; SAN 132805: 18a; SAN 134978: 19; SAN

135198: 2; SAN 136926: 13; SAN 137056: 1; SAN 138834: 12; SAN 138879:

11; SAN 139425: 11; SAN 139568: 13; SAN 141813: 18b; SAN 143457: 18b

— Sands et al. 5328: 11 — Sathvinderjit SKK 8: 13 — Sato 1099: 12 — Sato

Actinodaphne in Borneo 99

ep a) 237 13 — SEN 2/137: 5; SPN27291:,4; SEN 27686: 13; SFN 27725: 12;

SFN 36060: 4; SEN 40926: 5; SEN 11314: 9 — Schultze 35a: 2 — Shashi et al.

DSD 28: 2 — Sidiyasa et al. 1195: 10 — Sinclair & Kadim 9045: 13; 10522: 10

— Slooten 2274: 10 — Soejarto 69: 5 — Soepadmo ES 964: 11 — Soepadmo

& Mahmud ES 1049: 11 — Steenis 17393: 5 — Stevens PSF 393: 1; PSF 201:

1 — Stevens et al. PSF 482: 13 — Synge 1893: 3.

Tagawa et al. 321: 5 — Tandom BNB 4822: 1 — Teo & Din KL 4950: 14,

KL 4940: 5 — Teo & Remy KL 4086: 5 — Teysmann s.n.: 1.

Van Balgooy 4022: 5.

Wallich Cat. 2584b: 13 — Webb 715: 1 — Webb et al. MB 159: 13 — Wong

WKM 20: 1; WKM 36: 1; WKM 68: 5; WKM 144: 1; WKM 163: 1; WKM

835: 5; WKM 1116: 1; WKM 1368: 11 — Wong & Dransfield WKM 499: 9

— Wood 761: 1 — Wong & Wyatt-Smith W 81: 11 — Wray 280: 13.

Yabainus et al. SP 16287: 13.

Zainuddin AZ 17: 2; AZ 4918: 2; AZ 4984: 1 — Zainuddin et al. AZ 5674:

10 — Zazmee ZASZ 3: 2.

List of basionyms, synonyms and accepted names of Actinodaphne

species in Sabah and Sarawak

Basionyms and synonyms are given in italic and the accepted names in

bold and italic.

A. borneensis Meisn.

A. concinna Ridl. = A. pruinosa Nees

A. diversifolia Merr.

A. foxworthyana Gibbs = A. oleifolia Gamble

A. fuliginosa Airy Shaw

A. gelonioides Ridl. = A. oleifolia Gamble

A. glabra Blume

A. glomerata (Blume) Nees

A. johorensis Gamble

kinabaluensis Kosterm.

kostermansii var. glabrescens S. Julia

kostermansii var. kostermansii

macrophylla (Blume) Nees

maingayi Hook.f. = A. macrophylla (Blume) Nees

maingayi Hook.f. var. macrocarpa Gamble = A. glomerata (Blume) Nees

myriantha Mert.

oleifolia Gamble

percoriacea 8S. Julia

pruinosa Nees

pruinosa Nees var. kunstleri Gamble = A. pruinosa Nees

robusta S. Julia

semengohensis S. Julia

sesquipedalis Hook.f. & Thoms. var. macrocarpa (Gamble) Ridl. = A. glomerata (Blume)

~PDR PPP PR RD DDD

100

. soepadmoi S. Julia

. spathulifolia S. Julia

. sulcata S. Julia var. longipetiolata S. Julia

. sulcata S. Julia var. sulcata

. venosa S. Julia

Litsea glomerata Blume = A. glomerata (Blume) Nees

Litsea macrophylla Blume = A. macrophylla (Blume) Nees

Gard. Bull. Singapore 57 (2005)

Gardens’ Bulletin Singapore 57 (2005) 101—130 101

A Synopsis of the Bornean Species of

Microcos L. (Tiliaceae)

R. C. K. CHUNG, E. SOEPADMO

Forest Research Institute Malaysia

52109 Kepong, Selangor, Malaysia

AND

Axl. 151M.

Institute of Biological Sciences, University of Malaya

50603 Kuala Lumpur, Malaysia

Abstract

In preparing the treatment of the family Tiliaceae for the Tree Flora of Sabah and Sarawak

Project, a revision of the genus Microcos in Borneo was conducted resulting in 27 species

being recognised for Borneo, of which 25 species occur in Sabah and Sarawak; two (M.

laurifolia and M. tomentosa) are newly recorded, three (M. creaghii, M. elmeri and M.

ovato-lanceolata) are reduced to synonymy and one (M. longipetiolata) is excluded from

the genus. Fifteen species are endemic to Borneo. A complete list of exsiccatae,

nomenclatural (typification and synonymy) and taxonomic notes and distribution of the

recognised species are provided.

Introduction

The genus Microcos L. was founded by Linnaeus in 1753 based on the Sri

Lankan species, M. paniculata L. In 1767, Linnaeus reduced Microcos to

the synonymy of Grewia L. In the past few decades, the delimitation and

taxonomic status of Microcos and Grewia have been the subject of

controversy. Recently, Bayer et al. (1999), Chung (2002, 2003), Bayer &

Kubitzki (2003) and Chung et al. (2003, 2005) recognised Microcos as

distinct from Grewia based on morphological, wood anatomical, leaf

epidermal, pollen morphological characters and combined analyses of

plastid atpB and rbcL DNA sequences.

Microcos species are found mainly in the Malesian region, with

their centre of distribution probably in Borneo. Burret (1926) using

morphological characters of the flower (such as the pedicel length, stamen

number, ovary shape, ovule number per locule, and style length) recognised

two subgenera within Microcos: subgen. Microcos Burret (species from

Africa, Asia and West Malesia) and subgen. Eumeriandra Burret (species

102 Gard. Bull. Singapore 57 (2005)

from the Moluccas and New Guinea).

Up to 2005, a total of 97 binomials for Microcos had been published,

representing taxa from tropical Africa to Indo-Malesia (The International

Plant Names Index, 2005). Twenty-one binomials have been published for

plants occurring in Borneo (Merrill, 1921; Masamune, 1942; Anderson,

1980; Ashton, 1988; Whitmore et al., 1990; Kessler et al., 1992: Kessler &

Sidiyasa, 1994; Coode et al., 1996; Argent et al., 1997). In this paper, we

recognise 27 species as occurring in Borneo.

Microcos L.

Microcos L., Sp. Pl. 1 (1753) 514, Gen. Pl. ed. 5 (1754) 230; Burret, Notizbl.

Bot. Gart. Berl.-Dahl. 9 (1926) 756; Merrill, Univ. Calif. Publ. Bot. 15

(1929) 185; Masamune, En. Phan. Born. (1942) 449; Backer & Bakhuizen

f., Fl. Java 1 (1964) 393; Meijer, Bot. News Bull. Sandakan 7 (1967) 107;

Whitmore & Tantra, Tree Fl. Indon., Checkl. Sumatra (1986) 241;

Phengklai, Thai For. Bull., Bot. 16 (1986) 15, Fl. Thailand 6, 1 (1993)

33; Whitmore et al., Tree Fl. Indon., Checkl. Kalimantan 2, 1 (1990)

357; Kessler et al., Checkl. Tree Fl. Balikpapan-Samarinda Area (1992)

66; Kessler & Sidiyasa, Trees Balikpapan-Samarinda Area, Tropenbos

ser. 7 (1994) 228; Turner, Gard. Bull. Sing. 47 (1995) 487; Coode et al.,

Checkl. Flow. Pl. Gymno. Brunei (1996) 323; Boer & Sosef in Sosef et

al., Pl. Resources of South-East Asia 5, 3 (1998) 378. - Grewia L.

subgen. Microcos (L.) J.R.Drumm. in Gamble, Fl. Madras 1, 1 (1915)

114..- Grewia L. sect. Microcos (L.) Wight & Arn., Prodr. Fl. Ind.

Orient. 1 (1834) 81; King, J. Roy. As. Soc. Beng. 60, 2 (1891) 109. Type

species: Microcos paniculata L.

Arsis Lour., Fl. Cochinch. ed. 1, 1 (1790) 335. Type species: Arsis rugosa

Lour., Indo-China (= Microcos paniculata L.).

Fallopia Lour., Fl. Cochinch. ed. 1, 1 (1790) 335. Type species: Fallopia

nervosa Lour., China, Canton [= Microcos nervosa (Lour.) S.Y.Hu].

Omphacarpus Korth. in Temminck, Verh. Nat. Gesch. Ned. Bezitt., Bot. 6

(1842) 192, Verh. Nat. Gesch. Ned. Bezitt., Bot. 5 (1842) t. 42. - Grewia

L. sect. Omphacarpus (Korth.) Migq., Fl. Ind. Bat. 1, 2, 2 (1859) 204;

King, J. Roy. As. Soc. Beng. 60, 2 (1891) 109. Type species: Omphacarpus

opacus Korth., Borneo, Kalimantan [= Microcos opaca (Korth.) Burret].

Inodaphnis Miq., Fl. Ind. Bat., Suppl. 3 (1861) 357. Type species: Inodaphnis

lanceolata Mig., Sumatra [= Microcos lanceolata (Miq.) Burret).

Grewia L. p.p.: King, J. Roy. As. Soc. Beng. 60, 2 (1891) 108; p.p. (excl. G.

umbellata Roxb.); Ridley, Fl. Malay Penins. 1 (1922) 299, p.p. (excl. G.

umbellata Roxb.); Kochummen in Whitmore, Tree Fl. Malaya 2 (1973)

Microcos in Borneo 103

396, p.p. (excl. G. acuminata Juss., G. viminea Wall. ex Burret et G.

sclerophylla Roxb. ex Don); Corner, Wayside Trees Malaya 3rd edition,

2 (1988) 732, p.p. (excl. G. umbellata Roxb.).

Distribution: The genus comprises about 80 species occurring in tropical

Africa (not Madagascar), India, Sri Lanka, Myanmar, Indo-China, S China,

Hainan, Thailand, and throughout Malesia (except Lesser Sunda Islands).

In Malesia, some 42 species are known with two centres of diversity: 36

species in the West Malesia and 16 species in the Moluccas and New

Guinea. In Borneo there are 27 species with 15 endemics; Sarawak has 22

species with one endemic; Sabah 18 species with one endemic; Brunei 16

species (none endemic) and Kalimantan 16 species (none endemic).

Notes: Two basic inflorescence types are seen in Microcos (Chung 2001,

2003): Type A panicle where only first-order branches are conspicuous,

and Type B panicle where at least two orders of branching are conspicuous.

Two domatia types, pocket- or sac-type, are found on the abaxial

leaf surface either in the axils of basal pair of secondary veins or of other

secondary veins in some species of Microcos.

Four main types of non-glandular trichomes were observed in

Microcos (Chung, 2002), namely, simple, tufted (branched from the base

upwards), stellate (star-shaped) and cushioned stellate trichomes.

1. Microcos antidesmifolia (King) Burret

Microcos antidesmifolia (King) Burret, Notizbl. Bot. Gart. Berl.-Dahl. 9

(1926) 780; Turner, Gard. Bull. Sing. 47 (1995) 487; Coode et al. (eds.),

Checkl. Flow. Pl. Gymno. Brunei (1996) 323. — Grewia antidesmifolia

King, J. As. Soc. Beng. 60, 2 (1891) 113; Ridley, Fl. Malay Penins. 1

(1922) 302; Kochummen in Whitmore, Tree Fl. Malaya 2 (1973) 397,

p.p. (excl. syn. G. antidesmifolia King var. hirsuta King); Corner, Wayside

Trees Malaya 3rd edition, 2 (1988) 733; Anderson, Checkl. Trees Sarawak

(1980) 338; Ashton, Man. Non-Dipt. Trees Sarawak 2 (1988) 441, p.p.

(excl. syn. G. antidesmifolia King var. hirsuta King and M. subepetala

Stapf ex Ridl.); Whitmore ef al. (eds.), Tree Fl. Indon., Checkl.

Kalimantan 2, 1 (1990) 355; Kochummen, Tree Fl. Pasoh (1997) 428.

Type: King’s Collector 4029, Peninsular Malaysia, Perak, Larut (lecto

K., here designated).

Distribution: Peninsular Malaysia and Borneo.

Notes: In leaf shape, distance of the basal pair of secondary veins, and

104 Gard. Bull. Singapore 57 (2005)

fruit, Microcos antidesmifolia is closely related to M. triflora but differs in

the leaf texture (chartaceous vs. subcoriaceous), basal pair of secondary

veins running parallel to midrib at the base (absent vs. present), domatia

type in the axil of basal pair (pocket-type vs. sac-type), petiole (not swollen

vs. swollen), and the floral characters.

A species with two recognised varieties, in Borneo only var. hirsuta

is known. The other, var. antidesmifolia, occurs in Peninsular Malaysia.

var. hirsuta (King) Burret

Microcos antidesmifolia var. hirsuta (King) Burret, Notizbl. Bot. Gart. Berl.-

Dahl. 9 (1926) 780. — Grewia antidesmifolia King var. hirsuta King, J.

As. Soc. Beng. 60, 2 (1891) 113; Ridley, Bull. Misc. Inform., Kew (1933

489. Type: King’s Collector 10185, Peninsular Malaysia, Perak (lecto K,

here designated).

Microcos elmeri Merr., Univ. Calif. Publ. Bot. 15 (1929) 186; Masamune,

En. Phan. Born. (1942) 450; Coode et al. (eds.), Checkl. Flow. Pl. Gymno.

Brunei (1996) 323, p.p.. syn. nov. — Grewia elmeri (Mertr.) P.S.Ashton,

Man. Non-Dipt. Trees Sarawak 2 (1988) 443; Anderson, Checkl. Trees

Sarawak (1980) 338; Whitmore er al. (eds.), Tree Fl. Indon., Checkl.

Kalimantan 2, 1 (1990) 356. Type: Elmer 20911, Borneo, Sabah, Tawau

(A, BO, GH, K, L, NY, SING, UC).

Microcos creaghii Ridl., Bull. Misc. Inform., Kew (1933) 490, syn. nov.;:

Masamune, En. Phan. Born. (1942) 449. Type: Creagh s.n., Borneo,

Sabah, Sandakan (K).

Distribution: Peninsular Malaysia (confined to Perak) and Borneo (Sarawak,

Sabah, Brunei, and Kalimantan).

2. Microcos borneensis Burret

Microcos borneensis Burret, Notizbl. Bot. Gart. Berl.-Dahl. 9 (1926) 772:

Whitmore ef al. (eds.), Tree Fl. Indon., Checkl. Kalimantan 2, 1 (1990)

357: Coode et al. (eds.), Checkl. Flow. Pl. Gymno. Brunei (1996) 323. —

Grewia borneensis (Burret) P.S.Ashton, Man. Non-Dipt. Trees Sarawak

2 (1988) 442: Anderson, Checkl. Trees Sarawak (1980) 338. Type:

Haviland 2837, Borneo, Sarawak, Kuching (holo K; iso BO [3X], K,

SAR, SING).

Distribution: Endemic in Borneo (confined to Sarawak and Brune’).

Notes: Burret (1926) erroneously cited Hose 2837 as the type. However,

Microcos in Borneo 105

the correct type specimen for this species is Haviland 2837 (see Steenis,

1954) and the collection date of this specimen matches well with that cited

by Burret (1926). Two sheets of this collection (at flower-bud stage) were

examined at K. The sheet with the notes “flowers yellow” is the holotype,

while the sheet with the notes “near M. florida of Sumatra, venation of

leaves is different. It is not Motleys 1216 or 1260” is the isotype.

Microcos borneensis is closely related to M. riparia but the latter

differs from the former by its larger and thicker leaves with a greater

number of secondary veins (5—7 pairs), and longer inflorescences (3—8 cm

long), lower part of the androgynophore is longer (1.5—2 mm long) and

concave in side view, sparsely stellate-hairy style to between ‘/: and ‘/2 its

length, longer fruit (2.5—2.7 mm long) with 5—7 mm long pseudostalk,

thicker mesocarp (5—6 mm thick), thinner endocarp (c. 0.5 mm thick),

always with one fertile pyrene, and inconspicuous sterile pyrene.

3. Microcos cinnamomifolia Burret

Microcos cinnamomifolia Burret, Notizbl. Bot. Gart. Berl.-Dahl. 9 (1926)

770; Merrill, Univ. Calif. Publ. Bot. 15 (1929) 187; Masamune, En. Phan.

Born. (1942) 449; Whitmore ef al. (eds.), Tree Fl. Indon., Checkl.

Kalimantan 2, 1 (1990) 357; Kessler & Sidiyasa, Trees Balikpapan-

Samarinda Area, Tropenbos ser. 7 (1994) 228; Coode et al. (eds.), Checkl.

Flow. Pl. Gymno. Brunei (1996) 323; Boer & Sosef in Sosef et al., Pl.

Resources of South-East Asia 5, 3 (1998) 380. — Grewia cinnamomifolia

(Burret) P.S.Ashton, Man. Non-Dipt. Trees Sarawak 2 (1988) 443;

Anderson, Check]. Trees Sarawak (1980) 338; Argent et al. (eds.), Man.

Non-Dipt. Trees Centr. Kalimantan 2 (1997) 641. Type: Beccari PB

1617, Borneo, Sarawak (lecto K, here designated; isolecto BO, L).

Distribution: Endemic in Borneo (Sarawak, Sabah, Brunei, and

Kalimantan).

Notes: A very distinct species, it is the only Microcos with only a basal pair

of secondary veins that reach almost to the apex of the blade. Nevertheless.

it appears to be allied to M. Jaurifolia, with which it shares almost the same

leaf shape (narrowly elliptic to elliptic or lanceolate to ovate).

4. Microcos crassifolia Burret

Microcos crassifolia Burret, Notizbl. Bot. Gart. Berl. —Dahl. 9 (1926) 780,

p.p.. Merrill, Pl. Emer. Born. (1929) 186, p.p.: Masamune, En. Phan.

Born. (1942) 449: Whitmore ef al. (eds.), Tree Fl. Indon., Checkl.

106 Gard. Bull. Singapore 57 (2005)

Kalimantan 2, 1 (1990) 357; Kessler & Sidiyasa, Trees Balikpapan-

Samarinda Area, Tropenbos ser. 7 (1994) 228. — Grewia pyriformis

Merr., J. Str. Br. R. As. Soc. 86 (1922) 327, p.p., nom. illegit., non G.

pyriformis Elmer, Leafl. Philip. Bot. 8 (1915) 2841. Type: Ramos 1704,

Borneo, Sabah, Sandakan (holo K; iso L, US).

Distribution: Endemic in Borneo (confined to Sarawak, Sabah and Brune’).

Notes: In overall appearance, this species quite closely resembles Microcos

pachyphylla trom Borneo. However, it is sufficiently different to be

maintained as a separate species (see note in M. pachyphylla). Oblong,

ellipsoid or ovoid fruit galls, measuring 17—20 x 9-15 mm are sometimes

found in this species.

5. Microcos dulitensis Airy Shaw

Microcos dulitensis Airy Shaw, Kew Bull. (1949) 159. — Grewia dulitensis

(Airy Shaw) P.S.Ashton, Man. Non-Dipt. Trees Sarawak 2 (1988) 443;

Anderson, Checkl. Trees Sarawak (1980) 338; Whitmore et al. (eds.),

Tree Fl. Indon., Checkl. Kalimantan 2, 1 (1990) 355. Type: Richards

1767, Borneo, Sarawak, Mt. Dulit (holo K; iso BO, SING).

Distribution: Endemic in Borneo (Sarawak). Reported only from Mt. Dulit

and Usun Apau plateau.

6. Microcos fibrocarpa (Mast.) Burret

Microcos fibrocarpa (Mast.) Burret, Notizbl. Bot. Gart. Berl.-Dahl. 9 (1926)

782; Phengklai, Thai For. Bull., Bot. 16 (1986) 54, f. 24; Phengklai, Fl.

Thailand 6, 1 (1993) 38, f. 24; Turner, Gard. Bull. Sing. 47 (1995) 487;

Coode et al. (eds.), Checkl. Flow. Pl. Gymno. Brunei (1996) 323, p.p.;

Boer & Sosef in Sosef et al., Pl. Resources of South-East Asia 5, 3

(1998) 380, p.p. (excl. syn. Microcos reticulata Ridl.). — Grewia fibrocarpa

Mast. in Hooker f., Fl. Brit. India 1, 2 (1874) 391; King, J. As. Soc.

Beng. 60, 2 (1891) 111; Ridley, Fl. Malay Penins. 1 (1922) 301;

Kochummen in Whitmore, Tree Fl. Malaya 2 (1973) 397; Anderson,

Checkl. Trees Sarawak (1980) 338; Ashton, Man. Non-Dipt. Trees

Sarawak 2 (1988) 444, p.p. (excl. syn. M. reticulata); Corner, Wayside

Trees Malaya 3rd edition, 2 (1988) 733; Whitmore et al. (eds.), Tree FI.

Indon., Checkl. Kalimantan 2, 1 (1990) 356. Type: Maingay 1080,

Peninsular Malaysia, Malacca (lecto K [with flower buds and fruits],

Microcos in Borneo 107

here designated; isolecto K [with flower buds], K [with fruits]).

Distribution: India (fide Phengklai 1993), peninsular Thailand, Peninsular

Malaysia, and Borneo (Sarawak, Sabah and Kalimantan).

Notes: This species is easily confused with Microcos reticulata, but can be

distinguished by the distantly and obscurely serrulate leaf margin, impressed

midrib and secondary veins above, 1—1.5 mm long and glabrous lower part

of the androgynophore, broadly ovoid or globose ovary, which is broadly

ovate in cross section. It has soft indumentum on the twigs, lower leaf

surface and infructescences.

7. Microcos gracilis Stapf ex Rid.

Microcos gracilis Stapf ex Ridl., Bull. Misc. Inform., Kew (1938) 229;

Masamune, En. Phan. Born. (1942) 450; Coode et al. (eds.), Checkl.

Flow. Pl. Gymno. Brunei (1996) 323, p.p. — Grewia gracilis (Stapf ex

Ridl.) P.S.Ashton, Man. Non-Dipt. Trees Sarawak 2 (1988) 444;

Anderson, Checkl. Trees Sarawak (1980) 338; Whitmore et al. (eds.),

Tree Fl. Indon., Checkl. Kalimantan 2, 1 (1990) 356. Type: Haviland

1509, Borneo, Sarawak, Kuching (holo K; iso A, SING).

Distribution: Endemic in Borneo (Sarawak, Sabah, Brunei, and

Kalimantan).

Notes: Microcos gracilis is closely related to M. sumatrana in having

exclusively Type A panicles, glabrous lower part of the androgynophore,

globose ovary, and densely stellate-hairy obovoid fruits. It is different in its

basal pair of secondary veins reaching between !/1 and '/ the length of the

blade (vs. up to c. '/4 in M. sumatrana), petiole 0.5—1 mm long and not

swollen at the distal end (vs. 1—1.5 mm, and slightly swollen at the distal

end), narrowly elliptic and 0.5—1 mm wide involucral bracts of the inner

whorl (vs. oblanceolate, 1-1.5 mm wide), oblong sepals (vs. linear or

oblanceolate), oblong or obovate petals (vs. lanceolate), and 1n cross section

the ovary is elliptic with three shallow ridges (vs. circular).

8. Microcos henrici (Baker f) Burret

Microcos henrici (Baker f.) Burret, Notizbl. Bot. Gart. Berl.-Dahl., 9 (1926)

781, p.p.; Whitmore & Tantra, Tree Fl. Indon., Checkl. Sumatra (1986)

241; Coode et al. (eds.), Checkl. Flow. Pl. Gymno. Brunei (1996) 323,

p.p. — Grewia henrici Baker f., J. Bot. 62 (1924) 13. Type: Forbes 3014,

108 Gard. Bull. Singapore 57 (2005)

Sumatra, Palembang, Rupit River, Bingin (holo L [no. 908.144.547]; iso

BO, GH, L [nos. 908.144.548, 908.140.1977, 409613], SING).

Distribution: Sumatra (Jambi and Palembang) and Borneo.

Notes: Two subspecies are recognised: subsp. henrici (represented by Forbes

3014 and Posthumus 1036), which is endemic in Sumatra, and subsp. acuta

R.C.K.Chung from Borneo.

subsp. acuta R.C.K.Chung

Microcos henrici (Baker f:) Burret subsp. acuta R.C.K.Chung, Kew Bull.

58 (2003) 242, fig. 5. Type: Kostermans 12681, Borneo, Kalimantan,

Kutei, Belajan River, Tabang (holo L [no. 958.349.186]; iso L [no.

960.24.012], SING).

Grewia sp. 1, Ashton, Man. Non-Dipt. Trees Sarawak 2 (1988) 447.

Distribution: Endemic in Borneo (confined to Sarawak, Sabah and

Kalimantan).

9. Microcos hirsuta (Korth.) Burret

Microcos hirsuta (Korth.) Burret, Notizbl. Bot. Gart. Berl.-Dahl. 9 (1926)

782; Merrill, Univ. Calif. Publ. Bot. 15 (1929) 185; Masamune, En. Phan.

Born. (1942) 450; Whitmore & Tantra, Tree Fl. Indon., Checkl. Sumatra

(1986) 241; Whitmore et al. (eds.), Tree Fl. Indon., Checkl. Kalimantan

2, 1 (1990) 357; Kessler et al., Checkl. Tree Fl. Balikpapan-Samarinda

Area (1992) 66; Cheek & Turner, Kew Bull. 50 (1995) 129; Turner,

Gard. Bull. Sing. 45 (1993) 221, Gard. Bull. Sing. 47 (1995) 487; Coode

et al. (eds.), Checkl. Flow. Pl. Gymno. Brunei (1996) 324, p.p. —

Omphacarpus hirsutus Korth. in Temminck, Verh. Nat. Gesch. Ned.

Bezitt., Bot. 6 (1842) 193, Verh. Nat. Gesch. Ned. Bezitt., Bot. 5 (1842)

t. 42. — Grewia omphacarpa Migq., Fl. Ind. Bat. 1, 2, 2 (1859) 204; Merrill,

J. Str. Br. Roy. As. Soc., Spec. No. (1921) 373; Ridley, Fl. Malay Penins.

1 (1922) 301; Anderson, Checkl. Trees Sarawak (1980) 338; Ashton,

Man. Non-Dipt. Trees Sarawak 2 (1988) 445, p.p.; Whitmore et al. (eds.),

Tree Fl. Indon., Check]. Kalimantan 2, 1 (1990) 356. — Grewia hirsuta

(Korth.) Kochummen, in Whitmore, Tree Fl. Malaya 2 (1973) 39, nom.

illegit. et nom. superfl., non G. hirsuta Vahl, Symb., Bot. 1 (1790) 34;

Whitmore & Tantra, Tree FI. Indon., Check]. Sumatra (1986) 240. Type:

Korthals s.n., Borneo, Kalimantan, Doesoen River (lecto L [no.

908.253.200], here designated; isolecto L [nos. 944.56.122-124,

Microcos in Borneo 109

908.253.353, 908.253.341—342]).

Grewia palembanica Migq., Fl. Ind. Bat., Suppl. 3 (1861) 405. Type: Teijsmann

HB 3658, Sumatra, Palembang, Muara Enim (BO, L [no. 908.253.799],

Libgey.):

Distribution: Sumatra, Peninsular Malaysia, Singapore, and Borneo

(Sarawak, Sabah, Brunei, and Kalimantan).

Notes: Microcos hirsuta is closely related to M. phaneroneura (see note

under M. phaneroneura).

10. Microcos kinabaluensis R.C.K.Chung

Microcos kinabaluensis R.C.K.Chung, Kew Bull. 58 (2003) 330, fig. 1. Type:

Chew & Corner RSNB 4994, Borneo, Sabah, Mt. Kinabalu, Mesilau

River (holo SING; iso SAN, US).

Distribution: Endemic to Borneo (Mt. Kinabalu, Sabah).

11. Microcos latifolia Burret

Microcos latifolia Burret, Notizbl. Bot. Gart. Berl.-Dahl. 9 (1926) 781. —

Grewia latifolia Mast. in Hooker f., Fl. Brit. India 1, 2 (1874) 392, non

G. latifolia F.Muell. ex Benth. in Bentham & Mueller, Fl. Austral. 1

(1863) 271; King, J. As. Soc. Beng. 60, 2 (1891) 112; Gagnepain in

Lecomte, Not. Syst. 1 (1909) 132: Ridley, Fl. Malay Penins. 1 (1922)

300. — Grewia blattaefolia Corner, Gard. Bull. S.S. 10 (1939) 262:

Kochummen im Whitmore, Tree Fl. Malaya 2 (1973) 399; Anderson,

Checkl. Trees Sarawak (1980) 338; Ashton, Man. Non-Dipt. Trees

Sarawak 2 (1988) 442, p.p.; Corner, Wayside Trees Malaya 3rd edition,

2 (1988) 733: Whitmore et al. (eds.), Tree Fl. Indon., Checkl. Kalimantan

2, 1 (1990) 355; Kochummen, Tree Fl. Pasoh (1997) 429. — Microcos

blattaefolia (Corner) R.S.Rao, J. Bomb. Nat. Hist. Soc. 48 (1949) 300,

p.p.. Turner, Gard. Bull. Sing. 45 (1993) 221, Gard. Bull. Sing. 47 (1995)

487: Boer & Sosef in Sosef et al., Pl. Resources of South-East Asia 5, 3

(1998) 380. Type: Maingay 3150 (= Kew Distr. No. 249), Peninsular

Malaysia, Malacca (lecto K, here designated).

Distribution: Peninsular Malaysia, Singapore and Borneo (confined to

Sarawak and Brunei).

110 Gard. Bull. Singapore 57 (2005)

Notes: Microcos latifolia, a highly variable species with respect to vegetative

characters, is sometimes confused with M. globulifera from Peninsular

Malaysia. Two rather distinct entities can be recognised. The first,

represented by specimens collected from Peninsular Malaysia and Singapore

are characterised by elliptic to broadly elliptic leaves with an acute or

obtuse apex and impressed hairy midrib and secondary veins above; and

the second by specimens from Sarawak and Brunei which have narrowly

elliptic leaves with acuminate apex, and raised glabrous midrib and

secondary veins above.

The name Microcos latifolia (Mast.) Burret (Burret, 1926: 781) is

an illegitimate name because it was based on an illegitimate basionym

(Grewia latifolia Mast., 1874, non G. latifolia Benth. & F.Muell, 1863). In

accordance with the International Code of Botanical Nomenclature 2000

(St. Louis Code) Art. 58.3 with Ex. 2 (Greuter et al., 2000), the correct

name for the species is Microcos latifolia Burret with no parenthetic author

citation.

The epithet of “blattaefolia” is not to be changed to “blattifolia”

since Corner (1939) avowedly based it on Blatta, the Latin name for the

cockroach used by Pliny, Vergil, Horace, etc., and has nothing to do with

Blatti Adanson, a listed rejected name in the synonymy of Sonneratia nom.

cons.

12. Microcos latistipulata (Ridl.) Burret

Microcos latistipulata (Ridl.) Burret, Notizbl. Bot. Gart. Berl.-Dahl. 9 (1926)

795; Whitmore & Tantra, Tree Fl. Indon., Checkl. Sumatra (1986) 241:

Turner, Gard. Bull. Sing. 47 (1995) 487; Coode et al. (eds.), Checkl.

Flow. Pl. Gymno. Brunei (1996) 324, p.p. — Grewia latistipulata Ridl.,

Bull. Misc. Inform., Kew (1924) 262, Fl. Malay Penins. 5 (1925) 293;

Anderson, Checkl. Trees Sarawak (1980) 338; Ashton, Man. Non-Dipt.

Trees Sarawak 2 (1988) 444, p.p. (excl. syn. M. crassifolia Burret et M.

pachyphylla Merr.); Whitmore et al. (eds.), Tree Fl. Indon., Check.

Kalimantan 2, 1 (1990) 356. Type: Burkill SFN 7526, Peninsular Malaysia,

Selangor, Klang (holo K; iso SING).

Distribution: Sumatra, Peninsular Malaysia and Borneo.

Notes: A species with two recognised varieties: var. latistipulata is known

only from Peninsular Malaysia and Borneo and var. lanceolata (Ridl.) Burret

from Sumatra.

var. latistipulata

Microcos in Borneo at

Distribution: Peninsular Malaysia and Borneo (confined to the east coast

of Sabah).

Notes: Microcos latistipulata is characterised by the coriaceous and glabrous

leaves, as well as by the distinctly sharply prominent midrib and secondary

veins beneath, and glabrous pyriform fruits with pseudostalk. Specimens

of var. /atistipulata from Borneo differ from the Peninsular Malaysian ones

by their glabrous lower part of the androgynophore (vs. stellate-hairy) and

glabrous ovary (vs. stellate-hairy). However, the presence of a number of

specimens with intermediate characters means that the populations in

Peninsular Malaysia and Borneo should be grouped under var. Jatistipulata.

Some of these intermediate specimens are: Charington SAN 24716, Elmer

20310 and Meijer SAN 25133.

13. Microcos laurifolia (Hook.f. ex Mast.) Burret

Microcos laurifolia (Hook.f. ex Mast.) Burret, Notizbl. Bot. Gart. Berl.-

Dahl. 9 (1926) 771; Phengklai, Thai For. Bull., Bot. 16 (1986) 56, f. 25,

Fl. Thailand 6, 1 (1993) 39, f. 25; Turner, Gard. Bull. Sing. 47 (1995)

487; Boer & Sosef in Sosef et al., Pl. Resources of South-East Asia 5, 3

(1998) 381. — Grewia laurifolia Hook. f. ex Mast. in Hooker f, Fl. Brit.

India 1, 2 (1874) 392; King, J. As. Soc. Beng. 60, 2 (1891) 114; Baker f,

J. Bot. 62 (1924) 13; Kochummen in Whitmore, Tree Fl. Malaya 2

(1973) 399; Corner, Wayside Trees Malaya 3rd edition, 2 (1988) 733;

Kochummen, Tree Fl. Pasoh (1997) 429. Type: Maingay 1647, Peninsular

Malaysia, Malacca (lecto K [with flower buds], here designated; isolecto

K [with fruits]).

Distribution: India (fide Phengklai 1993), peninsular Thailand (fide Phengklai

1993), Sumatra, Peninsular Malaysia, and Borneo (confined to Brunei).

Notes: The specimen from Brunei (Dransfield et al. JD 6825) represents a

new record of the species for Borneo. The young twigs, petioles, midrib

and secondary veins of this specimen are covered with simple hairs (vs.

glabrous, except for the sparsely minutely stellate-hairy petioles for the

Peninsular Malaysian specimens).

Morphologically, M. laurifolia is closely allied to M. florida (Miq.)

Burret and M. loerzingii Burret (from Sumatra), M. pyriformis (Elmer)

Burret (from the Philippines) and M. kinabaluensis (from Borneo; see

Chung (2003) for further details).

Microcos florida differs from M. laurifolia by its hairy sac-type

domatia (vs. glabrous pocket-type), densely stellate-hairy ovary (vs. sparsely

112 Gard. Bull. Singapore 57 (2005)

covered with glandular trichomes in buds and less in flowers), fruits with a

short remnant of style (vs. without a style remnant), endocarp of less than

0.5 mm thick (vs. 0.5—1 mm thick), and consistently 1 fertile pyrene (vs. 1—

2).

Microcos laurifolia can be distinguished from M. loerzingii by its

equilateral leaf base (vs. inequilateral, i.e. with halves or sides unequal in

shape and size), thinner (1—1.5 mm) petioles (vs. c. 2 mm thick), sparsely

simple- or minutely stellate-hairy petioles (vs. glabrous), ovary sparsely

covered with glandular trichomes (vs. densely stellate-hairy), and glabrous

style (vs. sparsely stellate-hairy).

From M. pyriformis, M. laurifolia differs by the concolourous

subcoriaceous leaves (vs. discolourous chartaceous leaves), pyriform fruits

with distinct pseudostalk (vs. obovoid without pseudostalk), smaller fruits

1.5—2 x 0.8—1.3 cm (vs. 2.5—3 x 1.8—2 cm), thinner (1—1.5 mm) mesocarp

(vs. 4-6 mm thick), thinner (0.5—1 mm) endocarp (vs. 1-2 mm thick),

consistently with 1—2 fertile pyrenes (vs. 1), and shorter (4—6 mm) fertile

pyrenes (vs. 8-10 mm long).

Baker f. (1924), Ashton (1988) and Kessler et al. (1992) reported

that M. laurifolia occurs in Palembang, Sumatra (specimens Baker f. 3005

and 31/46), Sarawak (Ridley s.n.) and Kalimantan (S 292) respectively.

However, during this study we could not locate these specimens in order

to confirm their identity.

Ridley (1938) identified one collection from Sarawak (Beccari PB

3473) as that of M. laurifolia. Careful study of the specimen revealed that

it belongs to M. riparia.

14. Microcos membranifolia R.C.K.Chung

Microcos membranifolia R.C.K.Chung, Kew Bull. 58 (2003) 336, fig. 3.

Type: Singh SAN 30672, Borneo, Sabah, Sandakan, Sungai Binuang

(holo KEP; iso K, SAN, SAR, SING).

Distribution: Endemic in Borneo (confined to Sarawak and Sabah).

15. Microcos opaca (Korth.) Burret

Microcos opaca (Korth.) Burret, Notizbl. Bot. Gart. Berl.-Dahl. 9 (1926)

781; Whitmore ef al. (eds.), Tree Fl. Indon., Checkl. Kalimantan 2, 1

(1990) 357; Kessler et al., Checkl. Tree Fl. Balikpapan-Samarinda Area

(1992) 66. — Omphacarpus opacus Korth. in Temminck, Verh. Nat.

Gesch. Ned. Bezitt., Bot. 6 (1842) 193. - Grewia opaca (Korth.) Migq.,

Fl. Ind. Bat. 1, 2, 2 (1859) 204; Merrill, J. Str. Br. Roy. As. Soc., Spec.

Microcos in Borneo ] 13

No. (1921) 373; Masamune, En. Phan. Born. (1942) 449; Anderson,

Checkl. Trees Sarawak (1980) 338; Ashton, Man. Non-Dipt. Trees

Sarawak 2 (1988) 446. Type: Korthals s.n., Borneo, Kalimantan, Gunung

Sakoembang (lecto L [no. 908.253.752], here designated; isolecto L [no.

908.253.767]).

Distribution: Endemic in Borneo (confined to Sarawak, Sabah and

Kalimantan).

Notes: This species is sometimes confused with M. henrici subsp. acuta

because of its leaf shape, inequilateral leaf blade, and flattened midrib

above. Microcos opaca is sufficiently different in its inflorescence type,

filament and style indumentum, ovary shape, and fruit shape. Additionally,

it has acuminate leaves and the acumen with blunt tip.

16. Microcos ossea Burret

Microcos ossea Burret, Notizbl. Bot. Gart. Berl.-Dahl. 9 (1926) 779;

Whitmore et al. (eds.), Tree Fl. Indon., Check]. Kalimantan 2, 1 (1990)

357; Coode et al. (eds.), Checkl. Flow. Pl. Gymno. Brunei (1996) 324,

p.p. — Grewia ossea (Burret) P.S.Ashton, Man. Non-Dipt. Trees Sarawak

2 (1988) 446, p.p.; Anderson, Checkl. Trees Sarawak (1980) 338. Type:

Haviland 42, Borneo, Sarawak, Kuching (lecto K [with flower buds and

flowers], here designated; iso BO, K [with young and mature fruits], L

[no. 908.254.267], SING).

Microcos paucicostata Burret, Notizbl. Bot. Gart. Berl.-Dahl. 12 (1935)

602; Whitmore ef al. (eds.), Tree Fl. Indon., Checkl. Kalimantan 2, 1

(1990) 358. Type: Clemens 21071, Borneo, Sarawak, Kapit (A, K, NY,

SAR).

Distribution: Endemic in Borneo (Sarawak, Sabah, Brunei, and

Kalimantan).

Notes: In his original publication for the species, Burret (1926: 779) did not

specify the collector and collection number of the type. However, under

the distribution note, he stated that the species was described based on a

specimen on loan from K collected from a small tree near Kuching on 2

March 1893. We have examined specimens deposited at BO, K, L, and

SING and found that Haviland 42 bears the same notes on the habit,

locality and date of collection. Based on this, we concluded that Haviland

42 is the type of Microcos ossea Burret and here designate it as such.

114 i Gard. Bull. Singapore 57 (2005)

17. Microcos pachyphylla Merr.

Microcos pachyphylla Merr., Univ. Cahf. Publ. Bot. 15 (1929) 187;

Masamune, En. Phan. Born. (1942) 450. Type: Elmer 21880, Borneo,

Sabah, Tawau (holo US; iso A, BO, K, L, NY, SING).

Distribution: Endemic in Borneo (confined to Sabah, Brunei and E

Kalimantan). This species is common in Sabah and Kalimantan but rare in

Brunei. In Sabah, it is found mainly in the east coast districts (Sandakan

and Tawau), with one collection from Beaufort on the west coast.

Notes: Microcos pachyphylla is closely refated to M. crassifolia. In both

species the leaf is coriaceous and glabrous, the mesocarp is dorsilaterally

ribbed, and the specimens of fruits are often infested with galls. The former,

however, differs from the latter in having leaves shiny on both surfaces (vs.

not shiny), short petioles of less than 12 mm long (vs. 7-18 mm long), and

globose fruits (vs. ellipsoid or obovoid). Fruit galls, ovoid to broadly ovoid

or occasionally ellipsoid to oblong and measuring (9—)12—14 x 9-12 mm,

are often found in this species.

18. Microcos pearsonii (Merr.) Burret

Microcos pearsonii (Mertr.) Burret, Notizbl. Bot. Gart. Berl-Dahl. 9 (1927)

1171; Merrill, Univ. Calif. Publ. Bot. 15 (1929) 185; Masamune, En.

Phan. Born. (1942) 450; Whitmore et al. (eds.), Tree Fl. Indon., Checkl.

Kalimantan 2, 1 (1990) 358; Coode et al. (eds.), Checkl. Flow. Pl. Gymno.

Brunei (1996) 324. — Grewia pearsonii Merr., Philipp. J. Sci. 30 (1926)

Bot. 83; Masamune, En. Phan. Born. (1942) 449; Anderson, Checkl.

Trees Sarawak (1980) 338; Ashton, Man. Non-Dipt. Trees Sarawak 2

(1988) 447. Type: Wood 1216, Borneo, Sabah, Kudat, Lingkongan River

(holo UC [barcode UC 232336]).

Distribution: Endemic in Borneo (confined to Sarawak, Sabah, and Brunei).

Notes: The collections from Sarawak and Brunei have longer (9-15 mm

long) and sparsely stellate-hairy petioles and differ from the “typical”

collections from Sabah which have shorter petioles (2-7 mm long) and are

densely stellate-hairy. A number of collections (e.g., Dewol & Donggap

SAN 129474 from Sabah and Nielsen & Balslev 1109 from Brunei) have

the secondary veins and tertiary venation impressed above which differ

from those of the “typical” collections.

Microcos in Borneo : it 15

Microcos pearsonii, with its cordate leaf base, is distinctive among the

Bornean species of Microcos. It is closely allied to M. erythrocarpa and M.

malayana from Peninsular Malaysia, but differs from both by its cordate

leaf base, type of indumentum of the twigs, veins, petioles, inflorescences

and infructescences, and by its floral characters.

19. Microcos phaneroneura Burret

Microcos phaneroneura Burret, Notizbl. Bot. Gart. Berl.-Dahl. 12 (1934)

163; Whitmore et al. (eds.), Tree Fl. Indon., Checkl. Kalimantan 2, 1

(1990) 358; Kessler et al., Checkl. Tree Fl. Balikpapan-Samarinda Area

(1992) 66. Type: Hallier 2868, Borneo, Lianggagang (holo BO [with

white specimen tag.; iso BO [without white specimen tag], SING).

Distribution: Endemic in Borneo (confined to Sarawak, Brunei and

Kalimantan).

Notes: Microcos phaneroneura is closely related to M. hirsuta in the smooth

~ twigs, basal pair of secondary veins reaching between '/; and ‘/ the length

of the blade and forming an angle of less than 45° from midrib, lanceolate

petals, and smooth in the lower part of the androgynophore. However, the

former differs from the latter by its chartaceous leaf (vs. subcoriaceous),

sparsely simple- and stellate-hairy leaf above (vs. glabrous), impressed

secondary veins and tertiary venation (vs. flattened), exclusively Type A

panicles (vs. mixture), lanceolate involucral bract lobes of the outer whorl

(vs. ovate), narrowly elliptic involucral bracts of the inner whorl (vs.

oblanceolate), acuminate petals (vs. with shallow 2-3 teeth), and sparsely

stellate-hairy lower part of the androgynophore with cylindrical side view

(vs. glabrous with concave side view).

20. Microcos reticulata Ridl.

Microcos reticulata Ridl., Bull. Misc. Inform., Kew (1933) 490, p.p.;

Masamune, En. Phan. Born. (1942) 450; Coode er al. (eds.), Checkl.

Flow. Pl. Gymno. Brunei (1996) 324. Type: Creagh s.n., Borneo, Sabah

(holo K; iso K).

Distribution: Sumatra and Borneo (Sarawak, Sabah, Brunei, and

Kalimantan). Very common in Borneo.

Notes: Two sheets of Creagh s.n. collection (at flower-bud stage) were

examined at K. The sheet with a white envelop and a determination slip by

116 Gard. Bull. Singapore 57 (2005)

J.R. Drummond is considered the holotype, while the other sheet without

the envelop and Drummond’s determination slip is the isotype.

Ridley (1933) cited specimen Anderson 160 from Sarawak as

belonging to M. reticulata. However, this specimen has elliptic leaves with

an unequal base and pyriform fruits with narrowed pseudostalk, and thus

matches well with Haviland 1685 belonging to M. stylocarpoides Burret, a

species restricted to Sarawak and Kalimantan.

21. Microcos riparia (Boerl. & Koord.) Burret

Microcos riparia (Boerl. & Koord.) Burret, Notizbl. Bot. Gart. Berl.-Dahl.

9 (1926) 795; Whitmore & Tantra, Tree Fl. Indon., Checkl. Sumatra

(1986) 241. — Grewia riparia Boerl. & Koord. in Koorders-Schumacher,

Syst. Verz. 2 (1911) 35. Type: Koorders 10450b, Sumatra (holo BO).

Microcos ovato-lanceolata Burret, Notizbl. Bot. Gart. Berl.-Dahl. 12 (1934)

163, syn. nov.; Whitmore et al. (eds.), Tree Fl. Indon., Check]. Kalimantan

2, 1 (1990) 358; Kessler et al., Checkl. Tree Fl. Balikpapan-Samarinda

Area (1992) 66; Argent er al. (eds.), Man. Non-Dipt. Trees Centr.

Kalimantan 2 (1997) 641. Type: Hallier 1314, Borneo, Kalimantan, Sungai

Keniboeng (BO [3X]).

Distribution: Sumatra, Peninsular Malaysia (only recorded in Rompin FR,

Pahang) and Borneo (confined to Sarawak and E Kalimantan, recorded in

the area around Sungai Keniboeng, Sungai Kenepai, and Wanariset

Research Institute).

Notes: Burret (1934) distinguished Microcos ovato-lanceolata from M. riparia

solely by its lanceolate to ovate leaves with obtuse to rounded base (vs.

broadly ovate with truncate base in M. riparia). Detailed study on the

available specimens of both species showed that these distinguishing

characters of the leaves intergrade. In addition, the fruits of the two species

also possess many similar characters. In conclusion, M. ovato-lanceolata is

here reduced to the synonymy of M. riparia.

Sterile specimens of M. riparia, especially those with small leaves,

can be easily confused with M. borneensis. However, the latter can be

distinguished by its glabrous older twigs and leaves, chartaceous leaves,

and brown leaf colour when dried (see note under M. borneensis).

22. Microcos stylocarpoides Burret

Microcos stylocarpoides Burret, Notizbl. Bot. Gart. Berl.-Dahl. 12 (1934)

162; Whitmore et al. (eds.), Tree Fl. Indon., Checkl. Kalimantan 2, 1

Microcos in Borneo 17

(1990) 358; Kessler et al., Checkl. Tree Fl. Balikpapan-Samarinda Area

(1992) 66; Argent et al. (eds.), Man. Non-Dipt. Trees Centr. Kalimantan

2 (1997) 641 [cited as (Grewia?) Microcos stylocarpoides Burret]. Type:

Hallier 756, Borneo, Kalimantan, Soemedoene (holo BO).

Distribution: Endemic in Borneo (confined to Sarawak and Kalimantan).

Notes: Microcos stylocarpoides is closely related to M. henrici subsp. acuta.

In both taxa the leaves are subcoriaceous, elliptic or obovate, and with

flattened midrib and secondary veins above. In addition, the side axes of

the inflorescences bear bracteole and peduncle scars, the lower part of the

androgynophore is obovate in side view, and the style is sparsely stellate-

hairy for at least a quarter of its length. Microcos stylocarpoides differs

from M. henrici subsp. acuta mainly in having obovoid flower buds (vs.

narrowly oblong), linear sepals (vs. oblong), subglobose or broadly ellipsoid

ovary without ridges (vs. ellipsoid with 6 distinct and sharp ridges), and

pyriform fruits without ridges (vs. ellipsoid or obovoid with 6 distinct ridges).

23. Microcos subcordifolia R.C.K.Chung

Microcos subcordifolia R.C.K.Chung, Kew Bull. 58 (2003) 339, fig. 4. Type:

Kirkup & Thomas DK 727, Borneo, Brunei, Tutong, Lamunin, Kampung

Menangah (holo KEP [barcode 77837]; iso BRUN, KEP [barcodes

77838—77840]).

Distribution: Endemic in Borneo (confined to Sarawak and Brune’).

24. Microcos subepetala Stapf ex Rid.

Microcos subepetala Stapf ex Ridl., Bull. Misc. Inform., Kew (1938) 228;

Masamune, En. Phan. Born. (1942) 450. Type: Haviland 1885, Borneo,

Sarawak, Kuching (holo K).

Distribution: Endemic in Borneo (confined to Sarawak and Sabah).

Notes: The species is characterised by its glabrous twigs, conspicuously

small elliptic or ovate leaves, glabrous and subcoriaceous leaves with hardly

prominent venation, 4—6(—8) secondary veins, dense Type A or Type B

panicles, and glabrous ovary. Specimens from Sarawak consistently have

four pairs of secondary veins and pocket-type domatia, while those from

Sabah have 4—8 pairs of secondary veins and domatia are absent.

The species is closely related to M. antidesmifolia var. hirsuta in

118 Gard. Bull. Singapore 57 (2005)

leaf shape but can be distinguished by its subcoriaceous leaves with a

pointed tip, densely flowered inflorescences, broadly obovoid flower buds,

cup-shaped lower part of the androgynophore, and glabrous ovary.

25. Microcos sumatrana (Baker f.) Burret

Microcos sumatrana (Baker f.) Burret, Notizbl. Bot. Gart. Berl.-Dahl. 9

(1926) 783; Whitmore & Tantra, Tree Fl. Indon., Checkl. Sumatra (1986)

241; Whitmore et al. (eds.), Tree Fl. Indon., Checkl. Kalimantan 2, 1

(1990) 358; Kessler et al., Checkl. Tree Fl. Balikpapan-Samarinda Area

(1992) 66. — Grewia sumatrana Baker f., J. Bot. 62 (1924) 13. Type:

Forbes 2684, S Sumatra, Palembang, Batoe Pantjeh, Moesi River (lecto

L [no. 908.140.744], here designated; isolecto L [nos. 908.140.1824,

409657]).

Distribution: Sumatra and Borneo (Sarawak, Sabah, Brunei, and

Kalimantan).

Notes: See note under M. gracilis.

26. Microcos tomentosa Sm.

Microcos tomentosa Sm. in Rees, Cycl. 23, 2, 46 (1813) 2; Jack in Hooker,

Bot. Misc. 1, 3 (1830) 281, t. 60; Don, Gen. Hist. 1 (1831) 551; Backer &

_Bakhuizen f., Fl. Java 1 (1964) 393; Whitmore & Tantra, Tree Fl. Indon.,

Checkl. Sumatra (1986) 241; Phengklai, Thai For. Bull., Bot. 16 (1986)

" 92, £. 23; Phengklai, Fl. Thailand 6;°1°(1993) 37; 23) Taran eee

Bull. Sing. 47 (1995) 487. — Grewia paniculata Roxb. ex DC., Prodr. 1

(1824) 510; Blume, Bijdr. Fl. Ned. Ind 3 (1825) 115; Roxburgh, FI. Ind.

ed. 1832, 2 (1832) 591; Miquel, Fl. Ind. Bat. 1, 2, 2 (1859) 203; Masters

in Hooker f., Fl. Brit. India 1, 2 (1874) 393 (excl. Ins. Philipp.); King, J.

As. Soc. Beng. 60, 2 (1891) 110; Pierre, Fl. Forest. Cochinch. 11 (1888) ¢.

153; Gagnepain in Lecomte, Not. Syst. 1 (1909) 131 (excl. Ins. Philipp.),

in M.H.Lecomte, Fl. Indo-Chine 1, 5 (1911) 544; Koorders, Exkurs.-Fl.

Java 2 (1912) 577; Koorders & Valeton, Bijdr. Boomsoort. Java 1 (1894)

225, Atlas Baumart. Java 2, 8 (1914) figs. 393 & 394; Ridley, Fl. Malay

Penins. 1 (1922) 300; Kochummen in Whitmore, Tree Fl. Malaya 2

(1973) 397; Whitmore & Tantra, Tree Fl. Indon., Check]. Sumatra (1986)

241; Corner, Wayside Trees Malaya 3rd edition, 2 (1988) 734. Type:

Roxburgh s.n. in Herb. EIC 1097B, buds & fl., Peninsular Malaysia,

Penang (holo K-W, photo; iso BR [barcode BR-S.P. 817069], photocopy).

Grewia blumei Hassk. in Hoeven & de Vriese, Tijdschr. Nat. Geschied. 12

Microcos in Borneo 1 19

(1845) 130; Miquel, Fl. Ind. Bat. 1, 2, 2 (1859) 203. Type: Teijsmann s.n.,

Java (n.v.).

Grewia cumingiana Turcz., Bull. Soc. Natural. Moscou 1854, 31, 1 (1858)

231. Type: Cuming s.n., Peninsular Malaysia (B*).

Distribution: Myanmar (fide Phengklai 1993), S China (fide Phengklai 1993),

Indo-China (fide Phengklai 1993), Thailand, Sumatra, Peninsular Malaysia,

Singapore, Java, Borneo, and the Philippines (fide Phengklai 1993). In

Peninsular Malaysia, this species is common throughout except in the

southern states.

Notes: Microcos tomentosa is a new record for Borneo with single collection

from Kalimantan (de Vriese 111 = L sheet nos. 899.300.63 & 899.300.64).

It is closely related to M. paniculata in having peduncles, which arch out

and droop, and in its curved-striate exocarp. However, the two species can

be distinguished by characters of their leaves, flowers and fruits.

27. Microcos triflora (Blanco) R.C.K.Chung

Microcos triflora (Blanco) R.C.K.Chung, Kew Bull. 58 (2003) 346. —

Helianthemum triflorum Blanco, FI. Filip., ed. 2 (1845) 309, FI. Filip.,

ed. 3, 2, 12/13 (1878) 208, non Grewia triflora (Bojer) Walp., Repert.

Bot. Syst. 5, 1 (1845) 119; Merrill, Sp. Blancoan. (1918) 250. Type: Sp.

Blancoan. 864, the Philippines, Luzon Laguna, Mt. Maquiling (neo US

[barcode US 904559], “illustrative specimen” of Merr., Sp. Blancoan.

(1918) 250, designated by Chung, 2003; isoneo L [no. 921.22.116], PNH*].

Grewia stylocarpa Warb. in Perkins, Fragm. Fl. Philipp. 1 (1904) 104;

Gagnepain in Lecomte, Not. Syst. 1 (1909) 131; Merrill, Philipp. J. Sci. 1

(1906) Suppl. 90, Sp. Blancoan. (1918) 250, Enum. Philipp. FI. Pl. 3, 1

(1923) 27; Anderson, Checkl. Trees Sarawak (1980) 338; Ashton, Man.

Non-Dipt. Trees Sarawak 2 (1988) 447. — Microcos stylocarpa (Warb.)

Burret, Notizbl. Bot. Gart. Berl.-Dahl. 9 (1926) 780; Whitmore et al.

(eds.), Tree Fl. Indon., Checkl. Kalimantan 2, 1 (1990) 358; Coode et al.

(eds.), Checkl. Flow. Pl. Gymno. Brunei (1996) 324; Boer & Sosef in

Sosef et al., Pl. Resources of South-East Asia 5, 3 (1998) 381. Syntypes:

Warburg 11870, the Philippines, N Luzon, Malunu (B+, PNH7); Warburg

13072, the Philippines, C Luzon, Tayabas, Sampaloc (B+, PNHT7).

Distribution: Borneo, the Philippines and Sulawesi.

Notes: Two varieties are recognised in Borneo.

120 Gard. Bull. Singapore 57 (2005)

a. var. triflora

Distribution: Borneo (Sarawak, Sabah, Brunei, and Kalimantan), the

Philippines and Sulawesi.

b. var. longipetiolata (Merr.) R.C.K.Chung

Microcos triflora (Blanco) R.C.K.Chung var. longipetiolata (Merr.)

R.C.K.Chung, Kew Bull. 58 (2003) 347. — Grewia stylocarpa Warb. var.

longipetiolata Merr., J. Str. Br. R. As. Soc. 76 (1917) 97; Merrill, J. Str.

Br. Roy. As. Soc., Spec. No. (1921) 373; Ashton, Man. Non-Dipt. Trees

Sarawak 2 (1988) 447. — Microcos stylocarpa (Warb.) Burret var.

longipetiolata (Merr.) Burret, Notizbl. Bot. Gart. Berl.-Dahl. 9 (1926)

780; Merrill, Univ. Calif. Publ. Bot. 15 (1929) 186; Masamune, En. Phan.

Born. (1942) 450. Type: Villamil 243, Borneo, Sabah, Kalabakan

watershed (holo K; iso BO, SING, US).

Microcos havilandii Ridl., Bull. Misc. Inform., Kew (1938) 228; Masamune,

En. Phan. Born. (1942) 450. Type: Haviland 2332, Borneo, Sarawak,

Kapit, Rejang (A, K, SAR, SING).

Distribution: Endemic in Borneo (Sarawak, Sabah and Brune’).

Excluded species

Microcos longipetiolata Kosterm., Reinwardtia 6, 3 (1962) 301. Type: Meyer

SAN 19494 (holo K; iso SAN), Borneo, Sabah, Tawau, Tawau FR =

Scaphium longipetiolatum (Kosterm.) Kosterm. (Sterculiaceae). See:

Kostermans, Bull. Bot. Surv. India 7 (1965) 128.

Acknowledgements

We acknowledge the generosity of the directors, keepers and curators of

herbaria (A, B, BM, BO, BOL, BR, BRUN, C, CAL, FL, G, GRA, K,

KEP,.KLU, L; MEL, NYsP, PNE.PREsS. SAN. SARs SINGa Lia

UKMB, US, Z and the Kinabalu National Park, Sabah) for the loan of

specimens and facilities rendered. We are also grateful to Dr Dan H.

Nicholson (United States National Herbarium, Washington DC) and Dr J.

F. Veldkamp (Nationaal Herbarium Nederland, Leiden) who helped clarify

some nomenclature problems, Dr Peter C. van Welzen, Dr Ding Hou

(Nationaal Herbarium Nederland, Leiden) and Dr Martin Cheek (Royal

Botanic Gardens, Kew) for their help in sourcing references and materials.

Microcos in Borneo 121

This study was supported financially by IRPA grants 01-04-01-024 (RMKe-

7), 09-04-01-0073-EA001 (RMKe-8) and University of Malaya postgraduate

research grants 489/97, 225/98, and 209/99.

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Identification list

The number after the collector numbers refers to the following Microcos

taxa. When the number of the collection is not available or unknown then

the dates or sheet numbers are mentioned between brackets.

1 =M. antidesmifolia var. hirsuta 15 = M. opaca

2 = M. borneensis 16 = M. ossea

3. =M. cinnamomifolia 17 = M. pachyphylla

4 =M. crassifolia 18 = M. pearsonii

5 =M. dulitensis 19 = M. phaneroneura

6 =M. fibrocarpa 20 = M. reticulata

7 =M.- gracilis 21 = M. riparia

8 =M. henrici subsp. acuta 22 = M. stylocarpoides

9 =M. hirsuta 23 = M. subcordifolia

10 = M. kinabaluensis 24 = M. subepetala

11 = M. latifolia 25 = M. sumatrana

12 = M. latistipulata var. latistipulata 26 = M. tomentosa

13 = M. laurifolia 27a= M. triflora var. triflora

14 = M. membranifolia 27b= M. triflora var. longipetiolata

124 Gard. Bull. Singapore 57 (2005)

Aban SAN 29557: 9; SAN 30365: 3; SAN 30400: 3; SAN 30437: 16; SAN 30444: 17; SAN

30445: 16; SAN 30468: 18; SAN 30494: 9; SAN 30536: 27a; SAN 30545: 3; SAN 30547:

17; SAN 31186: 17; SAN 31201: 8; SAN 31231: 27a; SAN 31309: 17; SAN 32411: 9; SAN

32433: 3; SAN 32527: 1; SAN 33028: 18; SAN 33039: 20; SAN 34126: 1; SAN 35880: 17;

SAN 35959: 3; SAN 55485: 14; SAN 66719: 20; SAN 66758: 27a; SAN 66935: 9: SAN

73775: 27a; SAN 76683: 12; SAN 92479: 20; SAN 94313: 14; SAN 100338: 20 — Aban &

Dewol SAN 91619: 27a — Aban & Minjulu SAN 79677: 9; SAN 79825: 18; SAN 79830: 9

— Aban & Kodoh SAN 81001: 1; SAN 81033: 1 — Aban & Nordin SAN 47739: 20 —

Aban & Patrick SAN 73669: 27a; SAN 74285: 17 — Aban & Saikeh SAN 79374: 9 —

Aban & Soinin SAN 66340: 4; SAN 66898: 27b — Aban et al. SAN 66886: 20 — Abang

Daud S 433: 7 — Abang Mohtar S 53957: 18 — Abang Mohtar & Yii S 66193: 19; S 66199:

4; S 66238: 19 — Abang Mohtar et al. S 49273: 19; S 47121: 20; S 49546: 8; S 49589: 16; S

51418: 1 — Abdul Karim SAN 80225: 1 — Abdul Latiff ALM 2911: 8 — Abdul Rahim

SAN 92971: 9 — Agam SAN 37215: 12 — Agam & Aban SAN 40859: 8; SAN 40875: 9;

SAN 40876: 18; SAN 41604: 1 — Agama 555: 17 — Agama & Kadir SAN A 3505: 9 —

Ahmad SAN 50547: 20; SAN 52763: 9; SAN 54858: 27a; SAN 54951: 27a; SAN 55004: 7:

SAN 68343: 27a; SAN 70522: 12; SAN 70547: 14 — Ahmad & Latip SAN 52523: 27a:

SAN 52525: 14; SAN 52532: 27a; SAN 52540: 27a; SAN 52545: 27a; SAN 52570: 14;

SAN 52574: 27a — Ahmad & Nordin SAN 48967: 1 — Ahmad & Sairin SAN 59520: 17 -

Ahmad & Termiji SAN 67973: 8; SAN 68352: 20 — Allen 630: 1 — Ahmad Zainuddin et

al. AZ 4861: 25 — Aloysius SAN 78567: 14 — Ambri & Arifin W 382: 21; W 524: 21 —

Amin SAN 105194: 16; SAN 106128: 1; SAN 111510: 20; SAN 126416: 3; SAN 126618:

3; SAN 127039: 16; SAN 132040: 27a — Amin & Jarius SAN 116411: 20 — Amin &

Lideh SAN 70341: 4; SAN 110480: 3 — Amin & Matin SAN 108057: 9 — Amin & Sigin

SAN 111693: 12 — Amin & Zainuddin SAN 117380: 27a — Amin, Ismail & Hassan SAN

113604: 9 — Amin et al. SAN 59499: 1; SAN 67317: 4; SAN 69495: 14; SAN 69989: 9:

SAN 107147: 9 — Amir & Jafar SAN 40989: 9 — Ampuria SAN 19092: 4; SAN 32671A:

4: SAN 32764: 9; SAN 32855: 9; SAN 32873: 8; SAN 35309: 9; SAN 36501: 20; SAN

41262: 6; SAN 41274: 20 — Anak SAN 24870: 1 — Anderson 160: 22; S 4027: 7; S 4067: 8;

S 14709: 18; S 15956: 2; S 19135: 22; S 25147: 3 — Anderson & Ilias S 28502: 8 —-

Anderson & Keng K 70: 8 — Argent & Amiril 9380: 1 — Argent et al. 93195: 15 — Ariffin

et al. ARK 7: 9; BRUN 16876: 27a; BRUN 17460: 11; BRUN 17468: 3; BRUN 17720:

27b; BRUN 18257: 3 — Asah BRUN 3132: 3 — Ashton BRUN 31: 17; BRUN 62: 3;

BRUN 155: 9; BRUN 248: 25; BRUN 476: 16; BRUN 5505: 3; S 17622: 9; S 17800: 20; S

17946: 11; S 17956: 2; S 18198: 27b; S 18416: 27b; S 19788: 19; S 5936: 3 — Asik SAN

104276: 9; SAN 115802: 9 — Atkins et al. SA 433: 9; SA 460: 20; SA 504: 9 — Au S 17270:

11 - Awang Ahmad SAN 133138: 27a — Awang Enjah S 81830: 20 —- Awang Safri S

9648: 16.

Bakar SAN; 17312: 18; SAN 24963: 27a - Banang SAN 36915: 9; SAN 36925: 20; SAN

49356: 3; SAN 52059: 12; SAN 54076: 8 — Banyeng S 22029: 27a - Banyeng & Dami S

43677: 15 —- Banyeng & Jugah S 26703: 7 - Barttett s.n. (23/05/1893): 7 — Beccari PB

1617:.3; PB 1804: 3;.PB 1993:/22;°PB. 3217: 3; PB. 3473:21,= Benang ss 2alli. 464 —

Bernstein JHB 307: 3 - Binson SAN 62956: 27a — Binson & Lasan SAN 53179: 3 -

Blicher & Rantai S 57925: 8 - Bongsu SAN 62905: 27a- Bousi et al. SAN 123779: 27b;

SAN 123787: 27b - Brain S 14480: 11 - Brand SAN 21466: 17; SAN 30984: 4; SAN

30991: 12 - Brunig S 7440: 2 - Bujang S 12988: 16; S 20851: 2; S 20870: 2 - Bukah SAN

A 3416: 14 - Buntar SAN 25864: 9; SAN 27286: 16 - Burgess & Charington SAN

19110: 14 - Burley et al. 2413: 9; 2935: 9; NGS 407: 15; NGS 428: 1; NGS 484: 9; NGS

2307: 9; NGS 2356: 16; NGS 2392: 16; NGS 2824: 15; NGS 2900: 22: NGS 3018: 15; NGS

3243: 22; NGS 3248: 20 — Burkill SFN 7826: 12.

Microcos in Borneo 5

Campbell SAN 112325: 20 — Campbell EG 323: 20 — Carr SFN 26541: 1 — Castro 3792: 15;

Castro SAN A 819: 1 — Chai S 19274: 1; S 34080: 8; S 34667: 1; S 34769: 8; S 36100: 19; S

36174: 8; S 37226: 18; S 39657: 1; SAN 21682: 27a; SAN 25958: 8; SAN 26066: 7; SAN

26106: 20; SAN 26107: 18; SAN 26983: 27a; SAN 29304: 9; SAN 29660: 27a; SAN 29712:

27a; SAN 29741: 12; SAN 29824: 27b; SAN 31668: 9; SAN 33360: 18 — Chai & Ilias S

31118: 27b; S 31504: 4; S 31534: 9 — Chai et al. S 68827: 1 — Charington SAN 24680: 1;

SAN 24716: 12; SAN 26354: 27b; SAN 29852: 3 — Chew & Corner RSNB 4216: 10;

RSNB 4994: 10 — Chew et al. 33: 9; 100: 27a; 534: 9; 1468: 27b — Chin CSC 2688: 20;

KLU 2536: 19 — Chow SAN 75346: 8; SAN 75663: 8 — Chow & Leopold SAN 76407: 10

— Chung FRI 40317: 7; FRI 40318: 3; FRI 40338: 12 — Chung & Banyeng FRI 40315: 16;

FRI 40316: 16; FRI 40319: 16; FRI 40321: 4; FRI 40324: 16; FRI 40325: 16; FRI 40329:

16; FRI 40330: 16; FRI 40332: 4; FRI 40333: 16 — Chung & Dewol FRI 40335: 4; FRI

40339: 9; FRI 40340: 27a; FRI 40341: 1; FRI 40343: 4 — Church et al. 214: 1; NSF 13: 1;

NSF 21: 16; NSF 26: 8: NSF 88: 16; NSF 299: 1; NSF 318: 8; NSF 495: 16 — Clemens

20154: 20; 21071: 16; 27284: 9; 28828: 10; 31148: 27a — Cockburn SAN 68472: 12; SAN

70041: 14; SAN 83330: 27b; SAN 85050: 12; SAN 85115: 20 — Coode MC 7716: 3 —

Coode et al. MC 7923: 4; MC 6776: 7; MC 6914: 11; MC 7110: 9; MC 7865: 1 - Creagh

s.n (8/1895): 20; s.n. (5/1895): 1 — Cuadra SAN A 2278: 12; SAN A 2290: 14.

Dagang SAN A 2751: 27a — Daim DA 507: 18; DA 754: 18 — Dan S 16505: 4 — Daud &

Tachun SFN 35631: 16; SFN 35719: 22 — David S 10109: 20 - Dayang Awa & Ilias S

45267: 20; S 45291: 22; S 45589: 25; S 45620: 20; S 45689: 16; S 48783: 19 — Dayang Awa

& Lee S 47855: 27a; S 50065: 16 — Dayang Awa & Othman S 47035: 11 — De Vogel 856:

15; 885: 15 — De Vriese 111: 26 — Dewol SAN 55979: 12; SAN 71503: 9; SAN 78341: 14;

SAN 78492: 27a; SAN 80920: 27b; SAN 83791: 20; SAN 87969: 9; SAN 90341: 9; SAN

90450: 9; SAN 92192: 25; SAN 99477: 3; SAN 129459: 20; SAN 129540: 8 — Dewol &

Alexius SAN 88397: 20 — Dewol & Donggop SAN 109282: 1; SAN 129474: 18 — Dewol

& Karim SAN 80256: 1 — Dewol & Kodoh SAN 89316: 25 — Dewol & Petrus SAN

89581: 3 — Dewol & Termiji SAN 78247: 3 — Dewol et al. SAN 69909: 14; SAN 71196:

20; SAN 74951: 4; SAN 109631: 9; SAN 124624: 8 — Dransfield et al. JD 6825: 13; JD

W22E. Js. WIT 2AS: 29) ID 252;-35: YD 7255: 20; ID 7376: 3.

Boon F591: 16 — Bimer 20135: 12-2031 0: 12. 20911; 1; 21043: 3; 21094: 3; 21315: 9: 21365: 3;

21463: 18; 21591: 27b; 21704: 17; 21880: 17 — Enchai FMS 49060: 27a — Endert 1844: 7 —

Enggoh FMS 48988: 1; SAN 10541: 9 — Evangelista 761: 12 — Everett KEP 104926: 11.

Fabia SAN A 4002: 27a — Fidilis SAN 68758: 25; SAN 82152: 1; SAN 87322: 9; SAN 89811:

9; SAN 94782: 9; SAN 94800: 3; SAN 94856: 20; SAN 94895: 27a; SAN 95781: 25; SAN

95849: 9; SAN 95989: 9; SAN 96048: 9; SAN 103490: 9; SAN 110238: 20; SAN 116748:

27b; SAN 118738: 1; SAN 118812: 1; SAN 129982: 1; SAN 131539: 3; SAN 135832: 9;

SAN 135972: 20; SAN 136883: 8 — Fidilis & Asik SAN 96375: 1; SAN 110139: 16 —

Fidilis & Awang SAN 79236: 16 — Fidilis & Soludi SAN 105336: 16 — Fidilis & Sumbing

SAN 68804: 9; SAN 88219: 18; SAN 88232: 1; SAN 88333: 14; SAN 88493: 27a; SAN

89710: 14; SAN 91405: 9; SAN 91491: 3; SAN 91758: 9; SAN 91832: 27a; SAN 101394: 9;

SAN 114013: 20; SAN 125445: 16 — Forbes 2684: 25 — Forman & Blewett LLF 1053: 23

— Francis SAN 42870: 10.

Galau S 15645: 3 — Gambio SAN 117552: 12 — Gambio et al. SAN 133700: 7 — Geh &

Samsuri GSY 402: 20 — George SAN 121786: 9; SAN 128807: 9 — George & Joseph

SAN 130782: 9 — George et al. SAN 120513: 14; SAN 120978: 25; SAN 138330: 9 —

126 Gard. Bull. Singapore 57 (2005)

Good SAN 118088: 27a.

Hallier 756: 22; 1314: 21; 1833: 20; 1965: 21; 2208: 21; 2868: 19; 3020: 8 — Harvey & Melegrito

SH 10129 (= FMS 48903): 8 — Hashim SAN 33463: 3 — Hassan SAN A 3702: 27a —

Haviland c.c.m.a. (= 42): 16; c.L.v.n. (= 43): 11; s.1.: 11; s.n. (L sheet no. 908.254.234): 16;

s.n. (10/02/1894): 16; s.n. (10/09/1894): 7; s.n. (18/09/1894): 7; 42 bis: 16; 1471: 9; 1509: 7;

1683: 7; 1685: 22; 1742: 7; 1885: 24; 2256: 3; 2332: 27b; 2837: 2 —- Hemingway et al. CAH

80: 27b — Hose 626: 8.

Idris et al. BRUN 15652: 11 — Ignasius SAN 138762: 17 — Ilias S 13349: 19; S 15570: 4; S

15890: 16; S.16342: 15;.S 22901: 16; S 25762: 16;.S. 26022: 1; S 26185: 22; S 76004. 35

28128: 19: .S 28428: 19:-S 36373: 23: $:30520: 8:'S 39118: 20S 39139: 15: S 3uiaae

39248: 7; S 41035: 8; S 42089: 25; S 42480: 20; S 43423: 19; S 44192: 19; S 45121: 8 — Ilias

& Azahari S 35654: 4; S 35685: 1 — Ilias & Dami S 45004: 19 — Ilias & Yeo S 38372: 11 —

Imbungan & Patrick SAN 74182: 14.

Jacobs 5120: 16; 5122: 9 — Jahudin SAN 70467: 17; SAN 70476: 17; SAN 70478: 17; SAN

71653: 20; SAN 71720: 8; SAN 71785: 14 — James SAN 19045: 12; SAN 34937: 12: SAN

34965: 9; SAN 35490: 27a; SAN 40057: 17; SAN 40185: 1 — James et al. S 33746: 16; S

34506: 16; S 35089: 22 — Joffre et al. BRUN 16539: 11; BRUN 16554: 18; BRUN 17508:

18; BRUN 17533: 20; BRUN 17872: 9; BRUN 17876: 16 — Johari SAN 59641: 27a —

Joseph SAN 116938: 20 — Joseph & Lideh SAN 116883: 20; SAN 133008: 27a — Joseph

& Tuyuk SAN 119049: 9 — Joseph et al. SAN 113585: 1; SAN 123817: 12 — Jugah S

23857: 19; S 27756: 20 — Julius SAN 89414: 27a; SAN 119348: 27a; SAN 124172: 9; SAN

131043: 9; SAN 132992: 9 — Julius et al. SAN 124169: 20 — Jusimin JD 376: 8.

Kadir SAN A 2893: 1 — Kadir & Jiran SAN A 3367: 14 — Kamarudin et al. RC 97: 6 —

Kamideh SAN 66590: 27a — Kanis & Ding SAN 49318: 27a — Kanis & Sinanggul SAN

49337: 27a — Kapis SAN 46952: 3; SAN 46966: 3; SAN 46969: 3; SAN 46970: 3; SAN

46975: 3; SAN 46978: 3; SAN 46979: 3 — Kartawinata 696: 8; 815: 1; 963: 15 — Kessler

SAN 118158: 9 — King’s Collector 10185: 1 — Kinted SAN 15873: 27a — Kirkup &

Thomas DK 727: 23 — Kirkup et al. 323: 27a; DK 259: 3; DK 314: 16; DK 427: 3; DK

855: 9; DK 935: 9 — Kloss 19086: 20 — Kodoh SAN 87736: 12; SAN 87777: 20 — Kodoh

& Aban SAN 81928: 20 — Kokawa & Hotta 480: 20; 696: 14 — Koorders 10450b: 21 —

Korthals 489 (L sheet nos. 899.216.268, 899.216.269): 15; s.n. (L sheet nos. 899.310.11,

899.310.12, 908.253.178, 908.253.200, 908.253.341, 908.253.342, 908.253.353, 908.253.357,

908.253.358, 944.56.122, 944.56.123, 944.56.124): 9; s.n. (L sheet nos. 908.253.752,

908.253.767): 15 — Kostermans 4243: 20; 5095: 15; 5797: 3; 6004: 27a; 6488: 1; 6918: 27a;

7075: 3: 7280: 1: 7574: 8; 8715: 8: 8792: 16: 9635: 16; 9643: 20: 9729: 8: 10076: 17; 101s:

17; 10184: 7; 10424: 8; 12681: 8; 13466: 15; 13593: 3; 13603: 20; 21765: 8 — Kram S 1207: 4

— Kumin SAN 43119: 12; SAN 70570: 20; Kuripin SAN 25776: 6 — Kuzza s.n. (04/1976):

25.

Lai et al. S 68522: 3; S 74043: 8; S 72424: 24; S 75359: 8 — Lajangah SAN 28767: 27a; SAN

32261: 27b; SAN 47796: 18 — Lakasing SAN 70167: 8 — Lee S 39337: 3; S 39406: 11; S

41852: 20; S 43227: 15; S 52308: 1; S 54670: 19; SAN A 3107: 4 — Leopold SAN 33167:

12; SAN 33236: 9; SAN 36755: 17; SAN 47165: 27a; SAN 53133: 16; SAN 63511: 17;

SAN 63533: 20; SAN 73622: 14; SAN 76003: 4; SAN 78703: 20; SAN 81477: 12; SAN.

81483: 8; SAN 81584: 9; SAN 83567: 27a; SAN 89505: 1; SAN 91142: 9; SAN 92040: 25;

SAN 102192: 18; SAN 107940: 9; SAN 112940: 12; SAN 124404: 24; SAN 133490: 9;

Microcos in Borneo 7

SAN 133902: 9 — Leopold & Amin SAN 75368: 1; SAN 75372: 25 — Leopold & Dewol

SAN 60146: 20; SAN 60220: 1; SAN 74343: 20 — Leopold & Ismail SAN 108606: 18;

SAN 108636: 9; SAN 108678: 12 — Leopold & Kodoh SAN 81359: 9; SAN 81426: 9;

SAN 81557: 20; SAN 81567: 9 — Leopold & Patrick SAN 72589: 27a; SAN 72636: 20;

SAN 72650: 12; SAN 72680: 8 — Leopold & Pitty SAN 58956: 27a — Leopold & Saikeh

SAN 82587: 20 — Leopold & Sigin SAN 111472: 18 — Leopold et al. SAN 73816: 12;

SAN 73840: 12; SAN 73847: 20; SAN 114525: 20; SAN 133907: 9 — Lorence LL 540: 27a;

LL 714: 14.

Mahyar et al. 966: 6; 1123: 1; 1190: 15; NSF 786: 9 — Maikin & Tuyuk SAN 117531: 9 —

Maikin et al. SAN 132177: 20; SAN 132950: 27a — Maingay 1080: 6; 1647: 13; 3150: 11 —

Manis & Salang S 41592: 19 —- Mansus & Aban SAN 69232: 20 — Mansus & Dewol SAN

109240: 14 — Mansus et al. SAN 122293: 1 — Matin SAN 103515: 1 — Masirom SAN

43174: 14 — Meijer 2116: 1; 2146: 20; 2181: 16; 2228: 16; 2264: 20; 2525: 8; SAN 19169: 1;

SAN 19539: 8; SAN 20042: 3; SAN 21216: 27a; SAN 21235: 9; SAN 25133: 12; SAN

25377: 1; SAN 29500: 9; SAN 42493: 27a; SAN 44072: 27a; SAN 48135: 27a; SAN 51184:

3; SAN 53551: 14 — Meijer & Brand SAN 20068: 27a — Meijer & Patrick SAN 57476: 12

— Meijer & Pereira SAN 47225: 18 — Meijer & Singh SAN 24791: 17 — Meijer et al.

SAN 122556: 9 — Mendoza FMS 36793: 9 — Mikil SAN 30229: 27a; SAN 30339: 1; SAN

31390: 12; SAN 42061: 20; SAN 44334: 27b; SAN 46721: 27a — Minjulu SAN 77092: 16 —

Mogea & de Wilde 4347: 16 — Mohd. Haslani HA 26: 20; HA 36: 16 — Motley 1260: 27a

— Muas § 13354: 16; S 13388: 3 — Mukmin SAN 35601: 9; SAN 35613: 9; SAN 35614:

27a.

Nahar S 12662: 15 — Native 874: 9 — Native Collector 140: 2; 181: 7; 237: 7; 873: 7; 2505: 15

— Nedi 721: 17 — Neth. Ind. For. Service bb. 16252: 17; bb. 28325: 17 — Nicholson &

Patrick SAN 17045: 3; SAN 17707: 17 — Nielsen & Balslev 1006: 16; 1109: 18 — Niga

BRUN 15479: 3; NN 118: 23 — Niga et al. BRUN 15212: 11; BRUN 15448: 11; BRUN

15599: 9; BRUN 16779: 25 — Norasmah SAN 116487: 9 — Nordin SAN 46189: 27b; SAN

46256: 8; SAN 46317: 9; SAN 46332: 18; SAN 54500: 27b; SAN 54589: 9; SAN 85671: 8;

SAN 85786: 1 — Nordin & Ali SAN 54447: 9.

Ong OHC 716: 19 — Othman S 19944: 19; S 21354: 8; S 37444: 22 — Othman & Rantai S

57464: 16 — Othman et al. S 41351: 1; S 41483: 16; S 43361: 9; S 49042: 20; S 57187: 20; S

57801: 16; S 65469: 2.

Patrick SAN 20608: 17; SAN 21442: 16; SAN 23409: 12; SAN 39474: 3; SAN 65608: 12;

SAN 92077: 27a — Patrick & Leopold SAN 53031: 14; SAN 53038: 27a — Patrick &

Kadir SAN A 1852: 12 — Patrick & Matin SAN 116781: 14 — Patrick & Nicholson SAN

17701: 3 — Pereira SAN 43432: 27a — Pereira et al. JTP 248: 20; JTP 421: 9 — Pickles S

3680: 5; S 3689: 5 — Pitty SAN 55164: 12; SAN 55748: 1; SAN 58975: 27a; SAN 59795: 9

— Pitty & Binson SAN 61723: 12 — Pitty et al. SAN 61620: 17 — Pius SAN 108939: 9;

SAN 117717: 14 — Pius & Suali SAN 117725: 9 — Primack S 42409: 16 — Puasa A 3991:

12; CF 4627: 20; SH 10104 (= FMS 48878): 27b; SAN 36769: 12 — Prot SAN 32310: 27a;

SAN 32345: 27a; SAN 32376: 27a — Purseglove & Mohd. Shah P 4503: 11; P 4577: 24.

Ramos 1438: 12: 1704: 4 — Rantai et al. S 66605: 16; S 66622: 16 — Richards 1129: 9: 1767: 5

— Ridsdale 1926: 27a — Robinson SAN A 1: 4 — Roxburgh s.n. (in Herb. EIC 1097B):

26.

128 Gard. Bull. Singapore 57 (2005)

Sabana 5: 20; SAN 49708: 10 — Saigol SAN 93053: 9; SAN 93093: 25 — Saikeh SAN

64759: 1; SAN 69276: 27a; SAN 69284: 20; SAN 69287: 3; SAN 72053: 20; SAN

72193: 1; SAN 75493: 12; SAN 82369: 9; SAN 82384: 20; SAN 83183: 27a; SAN

87805: 14; SAN 87867: 14 — Saikeh & Leopold SAN 74478: 27a — Salleh et al.

BRUN 15315: 17; BRUN 16834: 9; BRUN 16853: 16 - Sani SS 629: 3; SS 664: 27a —

Sarawak Museum Series 165 (= Sarawak Museum T, 2, 3): 11; 171: 7; 763: 7 —

Sauveur 90: 20 — Sawan SAN 120370: 1; SAN 128068: 9 - Sayu SAN 35600: 9 —

Shea SAN 75215: 27a; SAN 75223: 27a; SAN 75615: 9 - Shea & Chow SAN 75703:

27a; SAN 75717: 8 = Sibat S 21727: 8; S 21790: 16; S 219796: 1; S 22544-°3: S 27622

27b; S 23033: 18; S 24381: 18; S 24508: 11; S 24565: 16; S 25179: 1 — Sidiyasa et al.

BRF 1790: 16; PBU 620: 8 — Sie S 35468: 9 - Sigin SAN 95034: 9 — Sigin & Amin

SAN 67559: 20 - Sigin & Joseph SAN 97151: 8 — Sigin et al. SAN 66534: 9; SAN

107409: 20; SAN 107703: 25 — Sinanggul SAN 51308: 20; SAN 54546: 12; SAN

56183: 20; SAN 56975: 16; SAN 57061: 27a; SAN 57105: 9; SAN 57303: 9; SAN

58036: 12; SAN 58042: 27a; SAN 58127: 27a; SAN 58128: 27a; SAN 58142: 27a;

SAN 58160: 27a — Sinclair & Kadim S 10287: 20 — Singh SAN 22765: 17; SAN

22801: 17; SAN 24003: 27a; SAN 24232: 20; SAN 29960: 17; SAN 29976: 27a; SAN

30100: 17; SAN 30672: 14; SAN 30685: 9; SAN 30716: 27b; SAN 30724: 1; SAN

30735: 27b; SAN 30738: 27b; SAN 31096: 12; SAN 34729: 9; SAN 34745: 27a; SAN

53452: 27b; SAN 58758: 20 - Singh & Aban SAN 29970: 17; SAN 29989: 18; SAN

29990: 1; SAN 30027: 1; SAN 30135: 18 — Singh & Hui SAN 30656: 27a - Singh &

Noordin SAN 48402: 9 — Singh et al. SAN 48870: 20 — Smythies S 12632: 25 —

Smythies et al. S 5877: 2 — Soegang 744: 17; 751: 17 — Soepadmo & Chai S 27611: 20

- Sow KEP 80180: 1 - Stevens et al. 402: 12 — Stone SAN 85179: 12 — Students s.n.

(08/1971): 26 — Suah SAN 36318: 1; SAN 37418: 9 - Suali SAN 117571: 9 — Sugau

JBS 242: 9; SAN 145223: 18 — Suhaili et al. BRUN 17437: 7; BRUN 18028: 3 — Suib

S 23458: 27b — Sumbing SAN 101452: 9; SAN 103566: 9; SAN 106907: 8; SAN

107041: 9; SAN 115844: 1; SAN 118376: 20; SAN 118533: 3; SAN 118684: 20; SAN

122040: 20; SAN 125305: 9; SAN 128267: 8; SAN 128519: 9; SAN 129924: 3; SAN

135695: 27b — Sumbing & Asik SAN 110245: 3; SAN 110869: 8; SAN 112958: 8.

Taib SAN 45165: 12 - Taleon 144: 8; KEP 55316: 8 — Talib SAN 84566: 1; SAN 84817: 3 —

Talip & Ejan SAN 85704: 14 — Tan S 37480: 3 - Tay & Mohd. Shah 176: 27b — Tay &

Othman 93-1113: 8 — Tay et al. 91-7: 6 — Teijsmann 10911: 9; 10914: 9; HB 3658: 9 —

Termiji SAN 72872: 9; SAN 76378: 12; SAN 81296: 27a; SAN 81538: 12; SAN 82860: 8;

SAN 85473: 24; SAN 90878: 12 — Termiji & Ali SAN 82931: 8; SAN 82937: 1 — Termiji

& Bongsu SAN 75885: 10; SAN 75820: 4 — Termiji & Dewol SAN 80036: 3 — Termiji &

Paul SAN 87643: 1 — Tiggi S 3311: 2 — Tingguan SAN 18791: 1 — Tong S 33866: 16; S

34324: 16; S 34916: 3; S 35005: 20; S 35035: 18; S 36536: 11; S 36584: 27b — Tong et al. S

34310: 15 — Tuah & Chai SAN 58709: 3 — Tuke P1-1005: 8.

Unknown FMS 25500: 6; FMS 34444: 20; FMS 34460: 3; SAN 44881: 12.

Veldkamp 8139: 9; 8292: 7; 8530: 8 — Villamil 243: 27b.

Wilkie 9544: 9 — Winkler 735: 8 — Wong SAN 68153: 12; SAN 68160: 4; WKM 379: 16:

WKM 704: 19; WKM 1244: 27b; WKM 1290: 16; WKM 1544: 9; WKM 1545: 11; WKM

1564: 7; WKM 1947: 1; s.n. (03/10/1988): 3 — Wood 1216: 18; 1281: 18; 1989: 17; 2273:

27a; SAN 15063: 3; SAN 15174: 3; SAN 15278: 17; SAN 16126: 1; SAN A 1980: 1; SAN

A 1992: 3; SAN A 2913: 9; SAN A 3682: 1; SAN A 3963: 27a; SAN A 4047: 27a; SAN A

4709: 14; SAN A 4723: 20; SAN A 4730: 12; SAN A 4788: 20; SAN A 4840: 1 — Wood

Microcos in Borneo 129

1216: 18 — Wood & Charington SAN 16511: 27a — Wood & Kadir SAN 17212: 27a —

Wood & Kapis SAN 16928: 3 — Wood & Wyatt-Smith SAN A 4290: 12 — Wright S

23589: 197 S72 7985-6; 5) 29013: 18.

Yahud et al. S 61960: 1; S 82558: 2 — Yap SAN A 716: 12 — Yui S 37887: 9; S 41139: 15; S

42004: 16; S 45938: 16; S 55824: 27a; S 61492: 14 — Yu & Alex S 66403: 16 — Yii & Shalih

S 64337: 16 — Yui et al. S 37752: 7; S 49164: 27a.

Zain SAN 61339: 27a; SAN 64703: 27a.

130 Gard. Bull. Singapore 57 (2005)

Index

The accepted names are in roman type, the synonymy and excluded names in italics. The

numbers after the names refer to the numbered species in this synopsis; (gen) = taxa listed

under the generic synonyms; (ag) = accepted taxa listed under the genus; (x) = excluded

taxa.

cinnamomifolia Burret 3

crassifolia Burret 4

Arsis Lour. (gen)

rugosa Lour. (gen)

Fallopia Lour. (gen) creaghii Ridl. 1

nervosa Lour. (gen) dulitensis Airy Shaw 5

Grewia L. elmeri Merr. 1

subgen. Microcos (L.) J.R.Drumm. (gen) fibrocarpa (Mast.) Burret 6

sect. Microcos (L.) Wight & Arn. (gen) gracilis Stapf ex Ridl. 7

sect. Omphacarpus (Korth.) Mig. (gen) havilandii Ridl. 27

antidesmifolia King 1

var. hirsuta King 1

blattaefolia Corner 11

blumei Hassk. 26

borneensis (Burret) P.S.Ashton 2

cinnamomifolia (Burret) P.S.Ashton 3

cumingiana Turcz. 26

dulitensis (Airy Shaw) P.S.Ashton 5

elmeri (Merr.) P.S.Ashton 1

fibrocarpa Mast. 6

gracilis (Stapf ex Ridl.) P.S.Ashton 7

henrici Baker f. 8

hirsuta (Korth.) Kochummen 9

latistipulata Ridl. 12

latifolia Mast. 11

laurifolia Hook. ex Mast. 13

omphacarpa Mia. 9

opaca (Korth.) Mig. 15

ossea (Burret) P.S.Ashton 16

palembanica Mig. 9

paniculata Roxb. ex DC. 26

pearsonii Merr. 18

pyriformis Merr. 4

riparia Boerl. & Koord. 21

stylocarpa Warb. 27

var. longipetiolata Merr. 27

sumatrana Baker f. 25

sp. 18

Helianthemum Mill.

triflorum Blanco 27

Inodaphnis Mig. (gen)

lanceolata Miq. (gen)

Microcos L.

antidesmifolia (King) Burret 1

var. antidesmifolia 1

var. hirsuta (King) Burret 1

blattaefolia (Corner) R.S.Rao 11

borneensis Burret 2

henrici (Baker f.) Burret 8

subsp. henrici 8

subsp. acuta R.C.K.Chung 8

hirsuta (Korth.) Burret 9

kinabaluensis R.C.K.Chung 10

latifolia Burret 11

latistipulata (Ridl.) Burret 12

var. latistipulata 12

var. lanceolata (Ridl.) Burret 12

lanceolata (Miq.) Burret (ag)

laurifolia (Hook. ex Mast.) Burret 13

longipetiolata Kosterm. (x)

membranifolia R.C.K.Chung 14

nervosa (Lour.) S.Y.Hu (ag)

opaca (Korth.) Burret (ag), 15

ossea Burret 16

ovato-lanceolata Burret 21

pachyphylla Merr. 17

paniculata L. (ag)

paucicostata Burret 16

pearsonii (Merr.) Burret 18

phaneroneura Burret 19

reticulata Ridl. 20

riparia (Boerl. & Koord.) Burret 21

stylocarpa (Warb.) Burret 27

var. longipetiolata (Merr.) Burret 27

stylocarpoides Burret 22

subcordifolia R.C.K.Chung 23

subepetala Stapf ex Ridl. 24

sumatrana (Baker f.) Burret 25

tomentosa Sm. 26

triflora (Blanco) R.C.K.Chung 27

var. triflora 27

var. longipetiolata (Merr.) R.C.K.Chung 27

Omphacarpus Korth. (gen)

hirsutus Korth. 9

opacus Korth. (gen), 15

Gardens’ Bulletin Singapore 57 (2005) 131 131

A Correction: Begonia nubicola Kiew

(Begoniaceae) is renamed B. oreophila Kiew

RUTH KIEW

The Singapore Botanic Gardens,

Singapore 259569

The Cloud Begonia, Begonia nubicola Kiew, is a unique begonia in

Peninsular Malaysia in that it lives on a dry coastal hill on boulders at the

headwaters of intermittent streams at the exact elevation where the cloud

cap forms on most mornings. Hence, ‘nubicola’ (cloud-dwelling) is a most

apposite name. Unfortunately, this name was already used for a Venezuelan

begonia (L.B. Smith & B.G. Schubert, Mem. New York Bot. Gard., 9

(1957) 354) so that B. nubicola Kiew is a later homonym. It is here proposed

to rename this species, B. oreophila Kiew as follows:

Begonia oreophila Kiew, nom. nov.

Begonia nubicola Kiew, Begonias of Peninsular Malaysia. (2005) 275 non

L.B. Smith & B.G. Schubert (1957).

Type: Kiew RK 5091, 31 Aug 2000, Bukit Labohan, Trengganu (holo SING;

iso K, KEP).

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Gardens’ Bulletin Singapore 57 (2005) 133—137 153

Two New Species Resembling Bulbophyllum plumatum Ames

and B. mirum J.J.Sm. (Orchidaceae)

JJ. VERMEULEN

Singapore Botanic Gardens

Singapore 259569

AND

P-O7BYRNE

14 Clementi Crescent

Singapore 599517

Abstract

Two new species of Bulbophyllum (Orchidaceae) are described, both similar to B. plumatum

Ames and B. mirum J.J. Sm. Bulbophyllum scotinochiton J.J.Verm. & P. O’Byrne is from

Sumatra, Indonesia; B. thiurum J.J.Verm. & P. O’Byrne is from Johore, Peninsular Malaysia.

Pictures of all four species are given.

Introduction

Bulbophyllum scotinochiton, described below, has been available in trade

for several years now, usually under the name Bulbophyllum mirum J.J.

Sm. Recently, the provenance of B. scotinochiton has become known, adding

to our knowledge about this species. It can now be formally described. The

Sumatran forests that appear to have yielded B. scotinochiton are quickly

disappearing. Besides illegal logging, the thick peat on the forest floor is

collected on a large scale to be used as garden mulch in nearby towns. The

collectors take care to hide their activities by leaving untouched a strip of

forest bordering the road. Elsewhere, large tracts of forest are slowly dying

because the canopy trees have lost their foothold.

The second species, B. thiurum, is said to occur in the Endau

Rompin State Park in the Johore lowlands, Malaysia. Locally, it has been

traded as ‘the yellow B. plumatum’, but very recently plants have also

appeared on the European market. Rumours suggest that, regrettably, the

population is already depleted.

Bulbophyllum scotinochiton J.J. Verm. & P. O’Byrne, sp. nov. - Plate 1.2

Bulbophyllum scotinochiton J.J. Verm. & P. O’Byrne, a Bulbophyllo miro

inflorescentia uniflora, floribus c. duplo majore (c. 5 cm longis) differt. -

134 Gard. Bull. Singapore 57 (2005)

TYPE: Indonesia, Sumatra, Lake Toba area, SBG-O 092] (SING, holo.).

Roots below the pseudobulbs. Rhizome creeping, c. 2.5 mm diam., sections

between pseudobulbs 2.5—3 cm long, bracts soon rotting away. Pseudobulbs

distant, ovoid, 1.3—1.5 x 0.7—1.1 cm, distinctly and rather sharply 4-angled.

Petiole 0.3—0.4 cm long. Leaf blade elliptic to ovate, 3.8—4.5 x 2.1—2.4 cm,

index (length/width) 1.5—2.2; subacute. Inflorescence porrect to patent, c.

10 cm long, 1-flowered. Peduncle c. 5 cm, bracts c. 4, the longest c. 10 mm

long. Floral bracts c. 5 mm long, acute. Flowers not fully opening. Pedicel

and ovary c. 10 mm long, basal node + flush with the surface of the rhachis.

Median sepal + porrect, ovate, c. 19 x 12 mm, index 1.5—1.6; top acute,

margins entire, base narrowly attached, rather thin, glabrous. Lateral sepals

adnate along the upper margins in the top half of the sepal, each oblique,

more or less elliptic, c. 50 x 10 mm, index c. 5, top obtuse, base narrowly

attached; adaxially glabrous, abaxially finely papillose; otherwise as the

median sepal. Petals porrect, transversely elliptic, c. 1.3 x 2.7 mm, index

0.4—0.5; rounded, top margin reflexed, entire, with c. 12 appendages, base

broadly attached; very thick, surface glabrous; appendages mobile, more

or less elliptic, up to 5.5 x 1 mm, subacute, margins slightly erose, base very

narrowly attached; thin, glabrous. Lip recurved about half way, general

outline subtriangular, c. 3.3 x 2.2 mm, index c. 1.5 (all without artificial

spreading); obtuse, margins entire; very thick, glabrous; adaxially more or

less flat near the base, with 2 more or less parallel, inconspicuous, low,

rounded ridges about half-way up the length of the lip, surface convex

towards the tip; abaxially with a distinct, retuse ridge over most of the

length of the lip. Column c. 3.2 mm long, stigma without ridges inside, with

2 distinct, longitudinal, short, rounded teeth at its base, column foot without

teeth. Stelidia c. 1 mm long, triangular, acute, with a small, triangular,

obtuse tooth along the upper margin and a distinct, deltoid, obtuse wing

along the lower margin. Anther abaxially with a distinct crest near the

attachment and a distinct, transverse crest overtopping the front margin,

surface somewhat papillose, front margin drawn out, truncate, entire.

Pollinia 4, the inner + as long as the outer, drop-shaped, the outer ellipsoid-

ovoid; no appendages present.

Colour: Median sepal white, slightly greenish in centre, with large

blackish purple spots and blackish purple margins. Lateral sepals white

near the base, but suffused with purple towards the tip, tip itself blackish

purple; lateral sepals entirely covered with small.purple spots. Petal

appendages pinkish purple towards the base, white at the tip. Lip pale

greenish, purple near the base. Column pale greenish.

Habitat: Understorey epiphyte in mossy montane forest dominated by oaks.

Altitude c. 1500 m.

Bulbophyllum scotinochiton and B thiurom

Plate 1. Four Bulbophyllum species.

la, b. Bulbophyllum mirum J.J.Sm.(SBG-O 4762); 2. Bulbophyllum scotinochiton J.J.

Verm. & P.O’Byrne (SBG-O 0921);3. Bulbophyllum plumatum Ames (SBG-O 5395);

4. Bulbophyllum thiurum J.J. Verm. & P.O’Byrne (SBG-O 5395). Photos : J.J. Vermeulen

136 Gard. Bull. Singapore 57 (2005)

Distribution: INDONESIA: Sumatra, Lake Toba area (2 specimens

seen).

Notes: Bulbophyllum scotinochiton is most similar to B. mirum

J.J.Sm. It differs in always having 1-flowered inflorescences and distinctly

larger flowers: median sepal c. 19 mm long (6—8 mm in B. mirum); lateral

sepals c. 50 mm long (20—26 mm in B. mirum). The petals and the lip,

however, are about the same size in both species. The information on B.

mirum was compiled from specimens from Indonesia (Java), and Malaysia

(Peninsula and Sabah); B. scotinochiton clearly falls outside the size range

of B. mirum. Additional differences between the two may be found in the

more distinct, deltoid wing along the lower margin of the stelidia, as well

as the somewhat wider appendages on the petals in B. scotinochiton (c. 1

mm wide; c. 0.5 mm wide in B. mirum).

Bulbophyllum thiurum J.J. Verm. & P. O’Byrne, sp. nov. — Plate 1.4

Bulbophyllum thiurum J.J. Verm. & P. O’Byrne, a Bulbophyllo plumato

sepalis lateralibus basi ovatis apicibus longe attenuates differt. - TYPE:

Malaysia, Johore, SBG-O 5395 (SING, holo.).

Roots below the pseudobulbs. Rhizome creeping, c. 1.5 mm diam., sections

between pseudobulbs 1.2—2.3 cm long, bracts little persistent. Pseudobulbs

distant, ovoid, 1.6—2.3 x 0.7-0.9 cm, obtusely 3—4-angled. Petiole 0.3—0.7

cm long. Leaf blade elliptic to ovate, 5.2—8.5 x 2.1—2.6 cm, index (length/

width) 2.5—3.4; obtuse. Inflorescence a subumbellate raceme, c. 20 cm long,

3—4-flowered. Peduncle c. 10 cm, bracts c. 5, the longest c. 9.5 mm long.

Rhachis nodding, somewhat thickened, c. 1.2 mm long. Floral bracts c. 7—8

mm long, acute. Flowers not fully opening. Pedicel and ovary c. 5.5 mm

long, basal node on a c. 0.5 mm-long thickened stump. Median sepal +

porrect, more or less orbicular, c. 6 x 6 mm, index c. 1; top shortly acuminate,

margins entire, base narrowly attached, rather thick, glabrous. Lateral sepals

proximally adnate along the upper (for 1 cm starting 0.5 cm from its base)

and lower (for 1 cm, starting at the base) margin, each oblique, ovate at

the base, with a long drawn-out top part, c. 94 x 6.5 mm, index 14—15, tip

acute, base narrowly clawed; thin, glabrous; otherwise as the median sepal.

Petals porrect, more or less elliptic, c. 1.7 x 1.3 mm, index 1.3—1.4; rounded,

entire, with c. 36 appendages, base rather broadly attached; thick, surface

glabrous; appendages somewhat mobile, ellipsoid-obovoid, up to c. | x

0.2—0.3 mm, tip obtuse, base tapering into a stalk of 1—1.5 mm, surface

consisting of large vesicles which become more elongate, almost hair-like,

towards the tip of the appendage. Lip recurved about half way, general

outline subtriangular, c. 1.7 x 1.2 mm, index c. 1.4 (all without artificial

spreading); obtuse, margins entire; thick; adaxially about flat near the base,

Bulbophyllum scotinochiton and B thiurom 137

with 2 about parallel, inconspicuous, low, rounded ridges about half-way

up the length of the lip, surface convex towards the tip, adaxial surface

finely and shortly pubescent; abaxially with a rather distinct, retuse ridge

over most of the length of the lip, surface glabrous. Column c. 1.7 mm

long, stigma without ridges inside, with 2 starting along its sides, converging

towards its base and continuing on to the column foot, column foot without

teeth. Stelidia c. 0.5 mm long, triangular, acute, upper margin erose, lower

margin with a distinct, deltoid, obtuse wing. Anther abaxially with an

inconspicuous crest, surface somewhat papillose, front margin drawn out,

truncate, entire. Pollinia 4, the inner slightly shorter than the outer, all

drop-shaped; no appendages present.

Colour: Flowers lemon yellow; median sepal with reddish spots,

lateral sepals with a patch of fine specks near the base, petals with pinkish

appendages. No scent.

Habitat: Epiphyte in lowland swamp forest.

Distribution: MALAYSIA: Johore (1 specimen seen); according to

local nurserymen a population occurs in the Endau Rompin State Park

area.

Notes: Bulbophyllum plumatum Ames is most similar. B. thiurum

differs in having the lateral sepals fused close to their base only, and along

the upper as well as the lower margins. In B. plumatum, these are fused

over a considerable part of their length, but along the upper margin only.

The shape of the lateral sepals is also different: wide and ovate in B.

thiurum, tapering into a caudate top part; but strap-shaped and barely

widened near their base in B. plumatum. Besides the appendages on the

petals, B. thiurum has almost hair-like vesicles near the top; in B. plumatum

all vesicles, up to the tip of the appendage, are rounded.

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Gardens’ Bulletin Singapore 57 (2005) 139-140 139

Book review: John Elliott. 2005 Orchid Hybrids of Singapore 1893-2003.

The Orchid Society of South East Asia, Singapore. ISBN 981 05 1752 1.

Price: Singapore dollars $120 (about £40 British pounds)

Orders to ossea@pacific.net.sg

Three hundred and four pages in hardback with stunning, glossy, pictorial

dust jacket, containing text written with eloquence and flair; 750 beautifully

rendered colour photographs with brief and elegant annotated descriptions

of 2,100 orchid hybrids raised in Singapore. Wow! This is a landmark

book, not only in tracing the history of orchid breeding in Singapore, the

history of the Orchid Society of South East Asia through 77 often turbulent

years and the rise of the orchid cut-flower trade, but also (and especially)

for recording and documenting these hybrids with the fascinating minutiae

of their disputed parentages, their raisers and whom they were named

after.

If I may begin at the end; the genius is in the index where the

orchids are listed in alphabetical order of the hybrid (grex) name. This a

real bonus at a time when the names of genera and hybrid genera are in

the process of being changed more frequently than a butterfly flaps its

wings. The new names given to the hybrid genera containing

Paraphalaenopsis have not been accepted, and the old names that

commemorate famous names in the history of Singapore’s orchid history

have been retained. John Elliott cogently argues the case for a reversal of

the International Orchid Registrar’s recent name changes which may

become even more confusing to ordinary growers when the status of

Euanthe, Papilionanthe, Ascocenda etc., are revised in the next edition of

Genera Orchidacearum.

The glory is in the main text, where the hybrids are listed, illustrated

and described by genus, in chronological order of registration so one may

the more easily see the changes in breeding over the years since Vanda

Miss Joaquim was registered in 1893. Information, information, information

is the joy of the reviewer. While it is a magnificent ‘coffee-table book’ it is

so full of information, from the origins of Vanda Miss Joaquim (was it an

artificial or a natural hybrid?) to the recording of great dedication (Christiera

Ramiah commemorates a gardener, Muthiya Pillai Ramiah who worked

for 50 years in the Singapore Botanic Gardens’ nursery) that every page is

worth reading as well as admiring.

There are biographical vignettes of the great Singapore orchidists

of the past - Holttum, Galistan and Laycock and we are introduced to the

orchidists of today: Dr Tan Wee Kiat VMH (Victoria Medal of Honour of

the Royal Horticultural Society, Gold Medallist of the American Orchid

Society); Tan Jiew Hoe (generous sponsor of this book, son of Tan Hoon

140 Gard. Bull. Singapore 57 (2005)

Siang past-President of OSSEA and raiser of Vanda Tan Chay Yan ‘Pride

of Singapore’ FCC/RHS); Peggy Tan (Vice-President of OSSSEA) who

designed it and who with Wendy Chew compiled pictures and information;

Syed Yusof Alsagoff (past President of OSSEA; raiser of 154 hybrids

including the only FCC ever awarded by OSSEA, and the inspiration

behind this book); Dr Yam Tim Wing (Senior Researcher, Orchid Breeding)

Singapore Botanic Gardens, who carries on the Gardens’ long tradition of

orchid breeding) - but one can, as the author notes, only list a few.

What a galaxy of talent must exist in Singapore to produce such

beautiful hybrids, and such a book! I loved it. Buy it. Treasure it. Imitate it

(if you can).

Henry Oakeley

Chairman

Royal Horticultural Orchid Committee

Wisley, U.K.

Gardens’ Bulletin Singapore 57 (2005) 141—142 141

Book review: Duistermaat, H. 2005. Field Guide to the Grasses of

Singapore (Excluding the Bamboos). Gardens’ Bulletin Singapore, Suppl.

57: 1-177, illus. ISSN 0374-7859, SING$ 30 (inclusive of p&p) from The

Library, Singapore Botanic Gardens, | Cluny Road, Singapore 259569.

The grasses are economically the most important family of flowering plants

in the world. It is therefore a pity that recent Malesian taxonomists take

so little interest in them (except for some students of bamboos). The

publication of this Guide is therefore a great occasion for the area. About

700 herbaceous grasses have been reported for Malesia, so the 134 treated

here may seem comparatively few, but because many of these have a very

wide distribution, the Guide therefore has a much wider application than

to the tiny island of Singapore alone.

The introduction is well-illustrated and the keys seem to work

well. Turf grasses generally do not get the chance to flower, so identification

with the usual keys is impossible. A novelty here is a key based on

vegetative characters. Another more extensive key to all species in the

Guide based on vegetative characters given at the end is also something

new. The argument for these two keys is that neglected lawns are invaded

by other species.

The genera and species are arranged alphabetically by their scientific

names. Usually a very brief synonymy is given, but the descriptions are

extensive and analytical. The first occurrence is noted. It is remarkable

that, although collecting in Singapore started in the early 19th century,

many ‘firsts’ are from the end of that period or early 20th century. The

difference in time may be that the older collections are not present in The

Singapore Herbarium (SING) and the Herbarium of the National

University of Singapore (SINU) on which this treatment was mainly based.

This reminds me of a report (perhaps a reader can tell me who published

it and where?) that the 19th century British colonial army had no hay and

straw for their horses so it was imported from Burma, whereby most of

the species now widespread in the Malay Peninsula were introduced at

that time.

Of great advantage are the numerous figures of the spikelets, where

the glumes have been shaded for better contrast. They were prepared by

J.J. Vermeulen, otherwise known for his taxonomic studies and marvellous

drawings in orchidology and malacology. He also made some of the

excellent colour photographs.

These, together with the careful introduction and the various keys,

make the excuse that grasses are ‘difficult’ a rather lame one.

The common weed Eragrostis cumingii has been reduced to E.

brownii. I can well understand that, for in the herbarium the differences

142 Gard. Bull. Singapore 57 (2005)

are slight and cumbersome: mainly in the basal branching system, which

often is lacking, size of the anthers, and the colour of the caryopsis.

Apparently in the field there are no supporting habitual characters. Current

studies indicate that the correct name for what I and others previously

have called EF. amabilis is E. tenella after all (Veldkamp, unpublished).

Figures 60 and 63 of E. atrovirens and E. gangetica are misleading, as only

one or no glumes at all are depicted. In Figure 66 the fine sculpture of the

lemmas of FE. unioloides should have been accentuated whereby the species

is immediately distinct from all others and especially from E. montana

(which, as is shown, subtly differs by the shape of the glumes and lemmas,

too, for the cognoscendi).

There is a challenge to experimenters with the diaspores of

Mnesithea glandulosa and Rottboellia cochinchinensis: the joints of the

fragile racemes have a knob at the base, which is an elaiosome and attracts

ants: do they carry them off? Contrary to the remark here (p. 90), Beumee

[Handl. 4e Congr. Ned.-Indie Nat. Wet. (1927) 419] already reported it for

both.

Other ant plants may be the bottle-brush species of Setaria.

Harvester ants (Camponotus spp.) cut up the inflorescences and take the

spikelets to their lairs as I observed with S. italica here in Europe. Collect

both ants and grass!

It might be noted that Panicum maximum, according to recent

studies, is actually a Urochloa with the first record for Singapore by Hullett,

27 Jan 1894.

J.F. Veldkamp

National Herbarium of the Netherlands

University of Leiden Branch

The Netherlands

Gardens’ Bulletin Singapore 57 (2005) 143—144 143

Book review: Rui-Liang Zhu and May-Ling So. 2001. Epiphyllous

Liverworts of China. Nova Hedwigia Beiheft 121. 2001. 418 pp. + 140

figures and 5 tables. J. Cramer. Berlin/Stuttgart. ISBN 3-443-51043-4

(Paper). |

This is a very well written and fully illustrated flora of epiphyllous liverworts

found in China. A total of 168 epiphyllous species belonging to 28 genera

in 10 families are recognized, described, and illustrated. Of these, 43

species are obligate epiphyllous liverworts, with the rest either as accidental

or facultative epiphyllous taxa. For each species, the distribution inside

China and their worldwide range are enumerated. Dichotomous keys to

the families, genera and species are presented. Two new species and one

new variety, and full synonymy for each taxon, of which 10 are new

synonyms, are proposed. The volume also contains several nomenclatural

changes and new lectotypifications. A glossary and index to scientific

names help a great deal in understanding the technical terms and in

accessing the rich information provided by this well-done revision.

Undoubtedly, the presence of epiphyllous liverworts forms one

interesting and unique visual phenomenon of the wet tropical rain forest.

Biologically speaking, this group of tiny plants and their adaptations to

grow on the wet surface of leaves of numerous plants has been a challenging

area of investigation for more than a century. Although many of the

species of hepatic epiphyllae are not a natural group from the phylogenetic

point of view, nevertheless, their repeated occurrence across the wide

tropical rainforest belt represent a remarkable convergent pattern of

evolution. As such, the reader is recommended to be educated first on

this topic by reading carefully the introductory discussion of the

morphological characteristics of epiphyllous liverworts and their adaptations

to the phyllosphere, as well as the summary on the ecology and distribution

of epiphyllous liverworts, found on pages 9—30, before attempting to use

the volume for any taxonomic identification.

Although the epiphyllous liverwort flora described in this volume

is based on the Chinese materials, more than a hundred of the species

recognized are, however, widespread in E and SE Asia. As such, the flora

is an up-to-date reference and useful guide to the study of epiphyllous

liverworts in Malesia, including Singapore.

In using the volume to key out some of my epiphyllous liverworts

collected from Nee Soon Swamp in Singapore, I have found the species

description too lengthy and the many measurements of leaf sizes and leaf

cells too detailed to be of great help to the reader who often needs to

decide on a species determination under time pressure. In the end, I have

resorted many times to base my conclusion about the species identity on

144 Gard. Bull. Singapore 57 (2005)

the short paragraphs of taxonomic synopsis and the diagnostic comments

that are concisely and sufficiently written for each taxon. Having said this,

the excellent and accurately executed line drawings of the species are very

useful in illustrating the species concept, especially for beginners.

My other enquiry pertains to the five centres of occurrence of

epiphyllous liverworts in China, presumably, these are the local areas with

high species number and diversity of epiphyllous liverworts. They are

identified as (1) Hainan, (2) Taiwan, (3) southwestern Zhejiang and

northern Fujian, (4) southern Yunnan, and (5) northwestern Yunnan and

southeastern Xizang. An oversight in this part of the discussion lies in the

failure of the authors to indicate the exact number of epiphyllous taxa

reported from each of the five centres. Instead, the richness of the taxa for

the five centres is represented in the map (Fig. 10) by the proportional

length of bars. It becomes difficult to estimate, by visual judgment of the

comparative length of the five bars, the differences in the number of taxa,

say between Centre 1 & 2 or Centre 2 & 3. Additionally, my own limited

experience with the study of tropical mosses in China has called my attention

to the richness of tropical bryophytes in Guangdong Province. Does the

omission of a Guangdong Centre represent a reality or an artifact of under-

collecting of epiphyllous liverworts in this province?

In collaboration with outsiders, resident Chinese plant taxonomists

have in recent years published many excellent floristic revisions and modern

systematic monographs that have shown the world the great diversity and

richness of their indigenous flora. This volume adds more evidence to this

fact by revealing convincingly the little known and yet very diverse world

of epiphyllous liverworts that exists in the wet tropical forests in China.

Workers in E and SE Asia can only thank the two authors for rendering a

great service in producing such a magnificent piece of summary work based

on their many years of hard work in the field and careful observation of

minute taxonomic characters under the microscope for each of the species

described. A copy of this supplement volume of the journal Nova Hedwigia

is a must for libraries specializing on tropical botanical literature.

Benito C. Tan

Department of Biological Sciences

National University of Singapore

Singapore

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Medinilla alternifolia Blume, Mus. Bot. Ludg.-Bat. I:1 (1849) 19.

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VOL. 57 (Part 2) December 2005 . oe ISSN 0374-7859

| \ NATIONAL PARKS BOARD |...

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THE GARDENS’ BULLETIN SINGAPORE

The Gardens’ Bulletin Singapore publishes papers on plant taxonomy (including revisions),

horticulture, phytogeography, floristics, morphology, anatomy and related fields with

emphasis on plants in the West Malesian region.

EDITORIAL BOARD

Dr R. Kiew Dr R.T, Corlett

(Editor) University of Hong Kong

Singapore Botanic Gardens Hong Kong

De PY. Yan Dr M.C. Roos

(Assist. Editor)

Singapore Botanic Gardens

Drs.€, Chin

Singapore Botanic Gardens

Dr M.J.E. Coode

Royal Botanic Gardens

Kew, U.K.

National Herbarium of the Netherlands

Leiden University

Leiden, The Netherlands

Dr E. Soepadmo

Forest Research Institute Malaysia

Kepong, Malaysia

Dr W.K. Tan

National Parks Board, Singapore

The Gardens’ Bulletin is published twice yearly by the National Parks Board, Singapore.

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nike, APR 0 6 2006 ( ARNOLD ARBORETUM

| UBRARY

The Gardens’ Bulletin

Singapore

VOL. 57 (Part 2) December 2005 ISSN 0374-7859

CONTENTS

Pearce, K. G.

A new Bornean Paraboea (Gesneriaceae) species endemic to Niah National

Patk sarawak, Malaysia. and further plant records from Niall. «..-...2....:cssc--ocseuseesesonncbesseoncesne 145

Pannell, C. M.

Aglaia scepadmot (Meliaceac), a. new Species for BOLNEO 2 c.ccseuscsesescisecsanebesseratencdenrees 183

Skornickova, J. and M. Sabu

Curcima roscoeana Wall(Zineiberaceae) 1 WII a’ 2... .e.ctecenvesecaacnconeatactavesndeateanbenetend 187

Skornickova, J. and M. Sabu

The identity and distribution of Curcuma zanthorrhiza Roxb.(Zingiberaceae) ........ 199

Boyce, P. C.

A new species of Rhaphidophora Hassk. (Araceae-Monstereae) from Borneo) ...... a)

Ng, F.S.P.

Taxonomic notes on Bornean Litsea, Lindera, Neolitsea and Iteadaphne

(LEVIES CEES) Ao Do pag ee EL si (Ger Re eee Pen er oe ee ee ene ee ee ZAG

Nguyen Q. H. and M. C. Tebbitt

An unusual new species of Begonia (Begoniaceae) from Vietnam ........... cece 247

Sam Y.Y. and L.G. Saw

Three new species of Scaphochlamys (Zingiberaceae) from Peninsular Malaysia ..... 253

Jutta, M. and Q.Z. Faridah

Malaxis inexpectata and Habenaria paradiseoides (Orchidaceae), new records for

Re AE A reac SU Aion gaceseeseran sc SRO eee MP a otis Scares acy sce avons ciomafenu'vasbnaaineieswpcodscaupewuiiis'ees 263

Ashton, P.S.

New Tristaniopsis Peter G.Wilson & J.T.Waterh. (Myrtaceae) from Borneo. ........... 269

Geri, C and F.S.P. Ng

Musa lokok (Musaceae), a new species of banana from Bario, Borneo ...................++ 279

S. Lee and Ghani, A. A.

Dipterocarpus tempehes (Dipterocarpaceae) - a new record for Singapore ............. 285

BOOK REVIEW

Rubiaceae of Thailand. A Pictorial Guide to Indigenous and Cultivated Genera

Kale Seak cides onus nnsinssivsonjnoleeustdema naar UabeernaSgessnce-bieueascnn Ce REee eee a eee a K.M Wong 289

Obituary

Arthur George Alphonso: ..2.5..66.2:6:0:..525 eee SY. Geile 21

Date of publication: 30 December 2005

Published by

National Parks Board

Singapore Botanic Gardens

Cluny Road

Singapore 259569

Printed by Oxford Graphic Printers Pte Ltd

Gardens’ Bulletin Singapore 57 (2005) 145—182 145

A New Bornean Paraboea (Gesneriaceae)

Species Endemic to Niah National Park,

Sarawak and Further Plant Records from Niah

KATHARINE G. PEARCE

32 Lorong Kumpang 4

93200 Kuching, Sarawak, Malaysia

Abstract

Paraboea culminicola K.G. Pearce, endemic to the Subis limestone at Niah, was first collected

in 1932 and subsequently in 1961, 1962 and 2002. The species is described and illustrated.

In addition, a list is provided of 357 taxa (356 species) from Niah represented by specimens

in the herbarium of the Sarawak Forestry Corporation (SAR), bringing the total number of

taxa recorded for Niah National Park to c. 550 and the total number for the Park and its

vicinity to c. 700; the total number of taxa associated with limestone at Niah to 281; and the

total of Borneo endemics found in the Park to 19 taxa.

Introduction

Paraboea in Borneo is currently represented by 14 described species. All

but one (P. paraboeoides (Kranzlin) B.L.Burtt) are found on limestone

and typically the species have restricted distributions, are described from a

single or few collections and from remote locations (Burtt, 1984; Xu &

Burtt, 1991).

In limestone locations in Sarawak for which data are available,

more than one Paraboea species co-exist on the same hill (west Sarawak

with P. clarke: B.L.Burtt and P. havilandii (Ridl.) B.L.Burtt (Kiew et al.

2004) and Mulu with P. banyengiana B.L.Burtt, P. candidissima B.L.Burtt,

P. effusa B.L.Burtt and P. meiophylla B.L.Burtt (Burtt, 1982) and now on

the Niah limestone with P. speluncarum B.L.Burtt (Burtt, 1971) and the

new species, P. culminicola K.G.Pearce, here described.

In preparation of the checklist of plants from the Niah National

Park, (Pearce, 2004), based on botanical field investigations and examination

of herbarium specimens carried out as part of the Sarawak Forest

Department/DANIDA Project ‘Support to Wild Life Master Plan

Implementation through the Improved Management of Totally Protected

Areas in Sarawak, Malaysia’, a re-examination of the Paraboea specimens

showed that one specimen from near the summit of Bukit Kasut did not

conform to any species yet described from Sarawak. It was neither P.

speluncarum, which is endemic to the Park, nor did it fit the description

146 Gard. Bull. Singapore 57 (2005)

for P. treubii H.O. Forbes as suggested by Burtt (1971). This prompted a

further look at other Niah Paraboea specimens in SAR with the conclusion

that it is a distinct new species, here described as P. culminicola.

The Niah National Park includes the striking Gunung Subis

limestone massif that reaches 394 m elevation and Bukit Kasut, a smaller

limestone hill 220 m high. The Great Cave, world famous for its edible

birds nests and prehistoric remains, and sometimes called ‘Niah Caves’ on

herbarium labels, is located in the northern part of the Gunung Subis

massif.

This new species was first collected in 1932 (Synge 589) and

subsequently in 1961 (Anderson S16045) and 1962 (Burtt & Woods B2023).

In 1971, Burtt described Paraboea speluncarum (then placed in the genus

Boea) from Niah Caves. In the same paper he mentioned Paraboea treubii

(H.O.Forbes) B.L.Burtt originally described from Karangnata, Sumatra

(Forbes, 1882) ascribing it to the three specimens from ‘Niah Caves’ and

Gunong Subis, together with Henderson SFN 19459, a specimen from Gua

Tipus, Pahang, Peninsular Malaysia. At the time, Burtt considered that

Boea havilandii (a monocarpic species from west Sarawak) ‘may well be’ a

synonym of B. treubii as he then understood it, although he noted that B.

havilandii differed in ‘detail of indumentum and nervature on the underside

of the leaf’.

In 1982, Burtt described four new species of Paraboea from Gunung

Mulu National Park, Sarawak (P. banyengiana, P. candidissima, P. effusa

and P. meiophylla). Burtt noted that P. effusa is ‘similar to P. treubii

(H.O. Forbes) B.L.Burtt but differs from this in its habit, which is not

monocarpic, and its glabrous sepals’. In 1984, Burtt published revised generic

concepts for Boea Lam and its allies, citing additional specimens from Gua

Musang, Kelantan for P treubii.

In 1991, Xu and Burtt described a new species, P. nervosissima,

from Peninsular Malaysia, based on specimens from Gua Tipus and Gua

Musang. Kelantan. However, they made no mention of the Niah specimens.

This left the three Niah specimens (Synge 589, Anderson S 16045 and Burtt

& Woods B 2023) doubtfully as P. treubii. A further Niah specimen

deposited at SAR (Sonny Tan & E. Wright S 27279) and a DANIDA

SWMPI Project plot specimen from the summit of Bukit Kasut, Niah

National Park, also represent this taxon.

This new species differs from Paraboea havilandii in the felty rather

than woolly tomentum covering the leaf undersurface, the shorter petiole

(3-6 mm long not 7-10 mm long), the greater number of pairs of leaf

veins (> 30 not 15-16 pairs) and more elongate calyx (> 3 times rather

than 2 times as long as wide at the midpoint) and shorter fruit (inclusive of

persistent style 1.5—2 cm, not 3.5 cm long). The new species differs from P.

Paraboea culminicola and plants from Niah, Borneo 147

effusa in its greater number of veins (> 30 pairs rather than 12-21 pairs),

relatively narrower calyx lobes (3-5 times rather than up to 2.5 times as

long as wide at the midpoint) and shorter fruit (1.5-2 cm, not 2.8 cm long).

It is therefore a distinct new species and is described below.

Paraboea culminicola K.G.Pearce, sp. nov.

Paraboea culminicola K.G.Pearce, sp. nov. a P. havilandii (Ridl.) B.L.Burtt

et P. effusa B.L.Burtt nervis foliorum magis numerosis (>30-jugis non 12-

21) et fructibus brevioribus (1.5-2 cm non 2.8-3.5 cm) distincta et etiam a

P. culminicola petiolis brevioribus (3-6 mm longis non 7-10 mm) differt.

Typus: Southern slopes of Gunong Subis, near Sekaloh river; Miri District;

Fourth Division Sonny Tan & E. Wright S 27279 (holo SAR).

Unbranched herb, 75-120 cm tall. Stem woody, with or without shrivelled

remains of dead leaves persisting below, with close, felty indumentum of

matted hairs. Young parts with a network of fine, cobwebby white hairs,

eventually lost. Leaves in whorls of 4; petiole winged, to 7-10 mm long;

blade discolorous, adaxial surface reddish grey-brown, with fine, cobwebby

white hairs eventually lost (barely visible at 10x magnification, thickly

scattered globose, glistening (apparently) glandular hairs), abaxial surface

with a pale cinnamon-brown, close, thin felty, persistent indumentum,

oblanceolate, somewhat falcate, 25-30 x 5.5-6 cm, base with wings

decurrent on the petiole, margin entire to obscurely dentate, veins c. 35

pairs, prominent below. Inflorescence a terminal panicle, its branches

subtended by more or less sessile foliose bracts, axes tomentose as the

stem, lowest branches 20-40 cm long, flowers with pedicel to 2.2 cm long,

calyx 2-5.5 mm long, split almost to the base, lobes narrowly triangular,

3-5 or more times as long as the width at the midpoint, apices incurved,

corolla in life pinkish cream or very pale blue, tube c. 2 mm long, the two

posterior lobes c. 8 x 6 mm divided by a 2 mm-long sinus, the anterior lip c.

1.1 cm long, trilobed, median lobe 6 x 6 mm; stamens arising at the mouth

of the tube, filaments c. 3 x 0.5 mm, flat, anthers transverse, cohering face

to face, 4 x 2.5 mm, beaked; ovary c. 1.5 mm, glabrous, narrowed to a

glabrous style 3.5-6 mm long. Fruit a capsule to 2 cm long, glabrous,

spirally twisted.

Distribution: Borneo - SARAWAK: endemic to the Subis limestone

Habitat: On bare, exposed limestone rocks or on mor soil at or near the

summit of limestone hills or pinnacles at 120-220 m altitude.

148 Gard. Bull. Singapore 57 (2005)

Notes: Paraboea culminicola is named for its habitat (at or near summits)

and to contrast its habitat as one of a pair of Paraboea species endemic to

Niah, the other being P. speluncarum. It would not be confused with P.

speluncarum, which has leaves with c. 20 pairs of veins, which tend to be

obscured by a thick woolly indumentum on the abaxial leaf surface.

Other specimens examined:

SARAWAK: Miri District, Niah National Park - Anderson S16045 (SAR),

southern slopes of Gunung Subis, near Sekaloh river; Bukit Kasut

DANIDA/SWMPI Plot Specimen Plot 20 No. 1 (SAR).

Further Plant Records from Niah

A checklist of plant species occurring at or in the vicinity of Niah National

Park, Sarawak, (Pearce, 2004), included c. 343 taxa (not all identified to

species), represented by >300 collections made during recent investigations

in the Park as part of the DANIDA/SWMPI Project Support to Wild Life

Master Plan Implementation through the Improved Management of Totally

Protected Areas in Sarawak, Malaysia (2000 — 2003) and >100 other records,

chiefly earlier SAR collections.

A second checklist presented here includes an additional 357 taxa

not listed in the earlier checklist, represented by >450 specimens in the

SAR collection. The majority of these have been collected by the Sarawak

Forest Department. Of these, 115 taxa are noted as being associated with

limestone. This brings the total number of limestone-associated taxa at

Niah to c. 281 (Pearce, 2004 reported 167 taxa associated with limestone

for Niah, now 166 with the confirmation of S 89270 as Calanthe triplicata).

Not all the specimens here cited were collected from what is now

Niah N.P. About 180 specimens, mostly collected from Ulu Sg. Sekaloh,

may have been located up to 15 km from the Park, the distance Sg. Sekaloh

extends beyond the Park boundary. These specimens are mostly trees,

collected in the 1960s, some from logging areas, and include many

dipterocarp species. They indicate the richness of dipterocarp forest and

some dimensions reached by trees in this area. A portion of this mixed

dipterocarp forest is still protected in the Park’s south west. Approximately

144 of the species in this checklist are represented only by such ‘park

vicinity’ specimens.

The c. 90 tree and c. 21 epiphyte species from the Park listed here

serve to provide a more complete picture of the flora of Niah N.P., the

herb and shrub flora having been well-collected through the efforts of

G.D. Haviland, C. Hose, Ahmad, W.M.A. Brooke, B.L. Burtt, the

DANIDA/SWMPI Project and others.

Paraboea culminicola and plants from Niah, Borneo 149

Figure 1. Paraboea culminicola K.G.Pearce. A. Shoot tip with inflorescence; B. leaf; C.

indumentum of the lower leaf surface; D. flower, E. front view of flower, F. L.S. flower; G.

flower after the corolla has fallen; H. stamens; I. fruit after dehiscence. (A-H DANIDA/SWMPI

Plot Specimen Plot 20 No. 1; 1.827279).

150 Gard. Bull. Singapore 57 (2005)

However, species identifications of Psychotria, Gnetum and Saurauia

remain uncertain.

At least 35 taxa here listed are Borneo endemics. Of these, seven

(Endocomia virella W.J. de Wilde, Gonystylus calophyllus Gilg, Paraboea

culminicola Pearce, Shorea calcicola P.S.Ashton, S. seminis (de Vr.) Slooten,

Vatica badiifolia P.S.Ashton, Voacanga havilandii Ridley and Zizyphus

borneensis Metr.) occur within the Park, bringing the number of Borneo

endemics found within the Park to 19 (Pearce, 2004).

This second checklist of taxa is not the last word on species that

occur in Niah N.P. and in its vicinity. Some specimens collected from Niah

are not represented at SAR (e.g. those collected by P.M. Synge in 1932

(Pearce, 2003)). Furthermore, some specimens recorded in the SAR card

index file representing species not already included in this or the earlier

(Pearce, 2004) checklist could not be located on the shelves.

Acknowledgements

The author is most grateful to Ruth Kiew, whose knowledge of the

limestone flora of the region in general and the Gesneriaceae in particular

prompted the careful examination of specimens of Paraboea from Niah

National Park and provided the stimulus for the recording of further species

known from Niah National Park, and for comments on the manuscript.

The author would also like to thank Mark J.E. Coode for translating the

diagnosis into Latin, and the curator of the SAR herbarium for permission

to examine specimens and use the Botany Unit Library at Sarawak’s Forest

Research Centre, to P.S. Ashton for making available his checklist of

Bornean Syzygium, and Joseph Pao for his excellent botanical illustrations.

References

Burtt, B. L. 1971. Studies in the Gesneriaceae of the Old World XXXIV:

A miscellany from South Eastern Asia. Notes Royal Botanic Garden

Edinburgh. 31: 36.

Burtt, B. L. 1982. New species and combinations in Gesneriaceae. In

Jermy, A.C. and Kavanagh, K.P. (eds.) Notulae et novitiates Muluenses

No. 2. Botanical Journal Linnean Society. 85:17—30.

Burtt, B. L. 1984. Studies in the Gesneriaceae of the Old World: XLVII.

Revised generic concepts for Boea and its allies. Notes Royal Botanic

Garden Edinburgh. 41: 401-452.

Paraboea culminicola and plants from Niah, Borneo 151

Forbes, H.O. 1882. On two new, and one wrongly referred, Cyrtandreae.

Journal Linnean Society Botany. 19: 279.

Kiew, R., C. Geri, S. Julia, P.C. Boyce. 2004. Understorey flora. In: Sarawak

Bau Limestone Biodiversity. H.S. Yong, F.S.P. Ng and E.E.L. Yen (eds.).

Sarawak Museum Journal. 59 (No. 80, New Series), Special Issue No. 6:

105—146.

Pearce, K.G. 2003. Five new Begonia species (Begoniaceae) at the Niah

National Park, Sarawak, Malaysia. Gardens Bulletin Singapore. 55: 73—

88.

Pearce, K.G. 2004. The Vegetation and Plants of Niah National Park,

Borneo. Gardens’ Bulletin Singapore. 56: 101-145.

Ridley, H.N. 1905. The Gesneriaceae of the Malay Peninsula. Journal Straits

Branch of the Royal Asiatic Society. 44: 73.

Xu, Z. and B.L. Burtt. 1991. Towards a revision of Paraboea (Gesneriaceae):

I. Notes Royal Botanic Garden Edinburgh. 48: 1-18.

Appendix — Checklist of Plants Collected from Niah National Park

Note: descriptions of plants and habitats from field notes have been merged

when there are more than one specimen.

MDF = Mixed Dipterocarp Forest; P.F. = Permanent Forest

G. = Gunung; Sg. = Sungai; Kg. = Kampung

3 species associated with limestone at Niah National Park

species endemic to Borneo

ACANTHACEAE

*Borneacanthus grandifolius Bremek.

Shrub 1 m tall. At base of limestone hill near river, alluvial soil, also limestone

scree below cave, mineral soil. Southern slopes of G. Subis near Sekaloh River

Sonny Tan & E. Wright S 27261; Sekaloh, G. Subis, Niah J.A.R. Anderson S 31960.

HAygrophila angustifolia R.Br.

Flat, wet ground at foot of hill. G. Subis, Niah B.L. Burtt & P.J.B. Woods B 2022.

Hygrophila salicifolia (Vahl) Nees

Niah Haviland 3513A.

Pseuderanthemum crenulatum Radlk.

G. Subis area, Niah Mohidin S 21676.

*Ptyssiglottis caudata (Stapf) B. Hansen

Limestone. Valley in front of Niah Caves B.L. Burtt & P.J.B. Woods B 2013.

152 Gard. Bull. Singapore 57 (2005)

*Ptyssiglottis frutescens Hallier

Shrublet 60-100 cm tall. Lower slopes of limestone hill, numerous limestone

boulders. G. Subis area, Niah Mohidin S 21679; Southern slopes of G. Subis near

Sekaloh River Sonny Tan & E. Wright S 27271.

Ptyssiglottis psychotriifolia (Stapf) B. Hansen

G. Subis area Niah Mohidin S 21667.

Rungia salaccensis Valeton

1-1.3 m tall. In jungle on wet ground. Kg. Tangap, Niah Ahmad 26; G. Subis area,

Niah Mohidin § 21655.

Staurogyne sp.

G. Subis area, Niah Mohidin S 21654.

ADIANTACEAE

* Adiantum caudatum L.

Limestone. Niah C. Hose s.n.; G. Subis area, Niah Mohidin S 21695.

*Adiantum hosei Baker

Small terrestrial fern. In damp, shady crevices of limestone rockface at 67 m a.s.l.

Great Cave, G. Subis, Niah J/.A.R. Anderson S 31938.

*Taenitis cordata (Gaudich.) Holttum

Small terrestrial fern. In shaded, dry rock crevices on damp, shaded limestone

rockface at 67 m a.s.l., Great Cave, G. Subis, Niah J.A.R. Anderson S 31945.

AGAVACEAE

Dracaena elliptica Thunb.

G. Subis area, Niah Mohidin S 21675.

ALANGIACEAE

Alangium javanicum (Blume) Wangerin var. javanicum

Tree 18 m tall, 45 cm girth. Lowland MDF 150 m a.s.l., soil sandy clay. Sg. Sekaloh,

Niah N.P. Yii Puan Ching S 40127.

ANACARDIACEAE

*Buchanania arborescens (Blume) Blume

Tree 3 m tall, 20 cm girth. Limestone hill summit 110 m a.s.l. Yii Puan Ching S$

40184.

Parishia maingayi Hook.f.

Tree 31 m tall, 75 cm girth. G. Subis P.F., Niah J.A.R. Anderson & Chew Wee Lek

S 16022.

Semecarpus glaucus Engl.

Tree 5.3 m tall, 12.5 cm girth. Primary lowland dipterocarp forest on clay loam

soil. Ulu Sg. Sekaloh, Niah river Sibat ak Luang S 26136.

ANNONACEAE

Alphonsea ?johorensis J. Sinclair

Paraboea culminicola and plants from Niah, Borneo 1535

Tree 17 m tall, 37 cm girth. MDF with reddish clay loam soil. Ulu Sg. Sekaloh,

Niah Erwin Wright S 29271.

Cyathocalyx magnifica Diels

Tree 8.7 m tall, 17.5 cm girth. Lowland dipterocarp forest, alluvial soil beside

stream. Ulu Sg. Sekaloh, Niah river Sibat ak Luang S 27877.

Dasymaschalon blumei Finet & Gagnep.

Treelet 4 m tall. Primary lowland dipterocarp forest, clay sandy soil beside river.

Ulu Sg. Sekaloh, Niah river Benang ak Bubong S 27873.

Desmos teijsmannii Merr.

Climber on tree. Riverine forest also MDF near stream on steep slope. Along Sg.

Niah, Niah N.P. Paul Chai S 40057, Yii Puan Ching S 40121.

*Enicosanthum coriaceum (Hook.f. & Th.) Airy Shaw

Tree 8 m tall, 30 cm girth. Primary forest, lower flanks of limestone mountain. G.

Subis, Niah J.A.R. Anderson S 31646.

Goniothalamus uvarioides Ridl.

Shrub 2.3 m tall. Primary lowland dipterocarp forest on clay loam soil. Ulu Sg

Sekaloh, Niah river Sibat ak Luang S26121.

Monocarpia borneensis J.B. Mols & P.J.A. Kessler

Tree 22 m tall, 45 cm girth. MDF with reddish clay loam soil. Ulu Sg. Sekaloh,

Niah Erwin Wright S 29268.

Polyalthia sumatrana (Miq.) Kurz

Tree 23 m tall, 90 cm girth. Primary lowland dipterocarp forest on clay loam soil.

G. Subis area, Niah Mohidin S 21644; Ulu Sg. Sekaloh, Niah river Benang ak

Bubong S 26104, Erwin Wright S 29120.

Pseuduvaria pamattonis (Miq.) Su & Saunders

Tree 18 m tall, 39 cm girth. Lowland MDF 150 mass.1., soil sandy clay. Sg. Sekaloh,

Niah N.P. Yii Puan Ching S 40126.

*Stelechocarpus cauliflorus (Scheff.) R.E. Fr.

Treelet 5 m tall. On steep slope of limestone hill at 80 m a.s.l. Niah N.P. Bernard

Lee § 40078.

Uvaria littoralis Blume

Climber on tree. On mudstone outcropping from riverbank also riverine forest.

Nangau Sekalan, Ulu Sungei Sekaloh, Niah river J.A.R. Anderson S 27599; along

Sg. Niah, Niah N.P. Yii Puan Ching S 40114.

APOCYNACEAE

*Kopsia arborea Blume

6 m tall, 27 cm girth. In exposed situation on limestone cliff at 117 m alt. G. Subis,

Niah J.A.R. Anderson S 16033.

aoheae

Voacanga havilandii Ridley

Tree 17 m tall, 45 cm diam. On wet podsolic soil. Niah N.P. Rena George S 39934.

154 Gard. Bull. Singapore 57 (2005)

AQUIFOLIACEAE

Ilex cymosa Blume

Tree 3 m tall, 25 cm girth. Riverine forest. Along Niah river, Niah Yii Puan Ching

S 40146.

*Ilex malaccensis Loes.

Small tree or shrub. Summit ridge of limestone 267 m alt., limestone strongly

dissected and covered by 15-23 cm ‘mor’ layer. Sg. Sekaloh, G. Subis, Niah J.A.R.

Anderson § 31929.

ARACEAE

Homalomena elliptica Hook.f.

G. Subis area, Niah Mohidin S 21630.

*Homalomena griffithii Hook.f. forma acuminata (Ridl.) Furtado

Herb on limestone rock. Foot of limestone at 17 m a.s.l. G. Brangin, Ulu Sg.

Subis, Niah N.P. Yii Puan Ching S 40176.

Homalomena rostratum Griff.

Boggy ground at edge of primary forest. Pengkalan Lobang to Niah Caves B.L.

Burtt & P.J.B. Woods B 1999.

*Scindapsus coriaceus Engl.

Epiphytic shrub 60—100 cm tall. Limestone hill, lower summit ridge, limestone

pinnacles and ‘mor’ soil. South side of G. Subis in Sekaloh river, Niah J.A.R.

Anderson, Sonny Tan & E. Wright S 27563.

ARECACEAE

Pinanga mooreana J. Dransf.

Palm 7 m tall. Ulu Sg. Sekaloh, Niah river Erwin Wright S 29126.

ASPLENIACEAE

*Asplenium filicipes Copel.

Small terrestrial fern. On limestone rocks, ravine between two limestone hills,

limestone pavement and small heavily eroded limestone hillocks. G. Subis area,

Niah Mohidin § 21620; Outside Great Cave, G. Subis, Niah J.A.R. Anderson

31906.

*Asplenium phyllitidis D. Don.

In side (sic) Gua Tulang on rock Ahmad 103.

*Asplenium squamulatum Blume

Epiphytic fern on small tree. Damp and shaded at base of limestone rock face at

67 ma.s.l. Great Cave, G. Subis, Niah J.A.R. Anderson S 31943.

*Asplenium thunbergii Kunze

Small terrestrial fern. On rocks, also epiphytic at base of small trees, ravine between

two limestone hills, limestone pavement and small heavily eroded limestone hillocks.

Outside Great Cave, G. Subis, Niah J.A.R. Anderson 31904.

*Asplenium aff. unilaterale Lam.

Small terrestrial fern. Slope between caves, limestone rocks, on residual soil not

Paraboea culminicola and plants from Niah, Borneo 155

on limestone, in semi-shaded locality, habitat semi-exposed, trees mainly felled.

Great Cave, G. Subis, Niah J.A.R. Anderson 31914.

ASTERACEAE

Blumea balsamifera (L.) DC.

2 m tall. In jungle on wet ground. Kg. Tangap, Niah Ahmad 18.

Sparganophorus vailantii Crantz

Wet, flat ground at foot of hill. G. Subis, Niah B.L. Burtt & P.J.B. Woods B 2023.

Synedrella nodiflora Gaertn.

At path side. Niah Caves B.L. Burtt & P.J.B. Woods B 2011.

BIGNONIACEAE

Deplanchea bancana (Scheff.) Steenis

Tree 40 m tall, 200 cm girth. Primary forest, alluvial soils, sandy, approaching

kerangas type. Path from Pangkalan to Great Cave, Niah J.A.R. Anderson 16427.

BORAGINACEAE

*Tournefortia sp.

Small climber or scrambler. Hanging on limestone cliff face. Outside Great Cave,

G. Subis, Niah J.A.R. Anderson S 31917.

BURMANNIACEAE

Burmannia sp.

G. Subis area, Niah Mohidin S 21638. (Mixed collection)

Gymnosiphon aphyllum Blume

G. Subis area, Niah Mohidin S 21638 (Mixed collection).

BURSERACEAE

*Canarium odontophyllum Miq.

Tree 20 m tall, 65 cm girth. On steep slope of limestone hill at 80 m a.s.l. Niah N.P.

Bernard Lee S 40073.

Santiria apiculata A.W. Benn. var. apiculata

Shrub 3 m tall. Primary lowland dipterocarp forest on clay loam soil. Ulu Sg.

Sekaloh, Niah river Sibat ak Luang S 26133.

Santiria mollis” Engl.

Tree 23 m tall, 120 cm girth. MDF with reddish clay loam soil. Ulu Sg. Sekaloh,

Niah Erwin Wright S 29272.

CAPPARACEAE

*Crateva magna (Lour.) DC.

Tree 10 m tall, 53 cm girth. On steep slope of limestone hill at 80 m a.s.1. Niah N.P.

Bernard Lee S 40074.

CECROPIACEAE

Poikilospermum cordifolium (Barg.-Petr.) Merr.

Large climbing epiphyte to 10 m. Disturbed belian forest, alluvial soils at base of

156 Gard. Bull. Singapore 57 (2005)

limestone hill. Between mouth of Sg. Sekaloh and G. Subis, Niah J.A.R. Anderson

S 27582.

*Poikilospermum scabrinervium (Barg.-Petr.) Merr.

Woody herb to 2 m tall. On limestone rock c. 50 m a.s.l. Niah N.P. Bernard Lee S

40087.

Poikilospermum suaveolens (Blume) Merr.

Climber 15-27 m tall entwined round tree. MDF with reddish clay loam soil. Ulu

Sg. Sekaloh, Niah Benang ak Bubong S 27885, Erwin Wright S 29275.

CELASTRACEAE

Bhesa paniculata Arn.

Tree 20 m tall, 60 cm girth. MDF on clay loam soil, 100 m alt. Ulu Sg. Sekaloh,

Niah river J.A.R. Anderson, Sonny Tan & E. Wright S 26068.

Kokoona sp.

Tree 25 m tall, 100 cm girth. Lowland MDF 80 m az:s.l., soil sandy loam. Sg.

Sekaloh, Niah N.P. Yii Puan Ching S 40130.

Lophopetalum glabrum- Ding Hou

Tree 15 m tall, 40 cm girth. Primary MDF on gentle slope, clay loam soil. Ulu Sg.

Sekaloh, Niah river J.A.R. Anderson, Sonny Tan & E. Wright S 26124; Niah N.P.

Bernard Lee S 40072.

*Reissantia grahamii (Wight) Ding Hou

Creeper. Limestone hill summit 110 m a.s.]. G. Brangin, Niah N.P. Yii Puan Ching

S 40185.

*Siphonodon celastrineus Griff.

Tree 10 m tall, 40 cm girth. On near vertical slope of limestone hill, dry and in

partial shade, 67 m alt. G. Subis, Niah J.A.R. Anderson S 16034.

CHRYSOBALANACEAE

Maranthes corymbosa Blume

Tree 18 m tall, 90 cm girth. MDF remnant forest. Niah N.P. Bernard Lee S 40067.

CLUSIACEAE

Garcinia parvifolia Miq.

Tree 17 m tall, 60 cm girth. On mudstone outcropping from river bank. Nangau

Sekalan, Ulu Sg. Sekaloh, Niah river J.A.R. Anderson S 27288, Erwin Wright S

29264.

COMBRETACEAE

Combretum tetralophum C.B.Clarke

Climber on tree. Riverine forest. Along Sg. Niah, Niah N.P. Yii Puan Ching S

40115.

CONNARACEAE

Agelaea borneensis Mert.

Small climber. MDF, clay loam soil, sedimentary rocks, 100 m alt. Ulu Sg. Sekaloh,

Paraboea culminicola and plants from Niah, Borneo 57

Niah river E. Wright S 27251.

Connarus grandis Jack

Climber 23 m tall. MDF. Ulu Sg. Sekaloh, Niah Erwin Wright S 29273.

Ellipanthus beccarii Pierre var. peltatus’ (Schellenb.) Leenh.

Tree 7 m tall, 12.5 cm girth. Primary lowland dipterocarp forest on clay loam soil.

Ulu Sg. Sekaloh, Niah river Benang ak Bubong S 26109.

Rourea mimosoides Planch. f. obtusifolia Leenh.

Climber on tree. Riverine forest and river bank on sandy clay soil. Ulu Sg. Sekaloh,

Niah river J.A.R. Anderson 27295; Along Niah River, Niah N.P. Yii Puan Ching S

40148.

COMMELINACEAE

Amischotolype marginata Hassk.

Large herbaceous creeper. Alluvial soil at base of limestone hill near river. Southern

slopes of G. Subis, near Sekaloh River Sonny Tan & E. Wright S 27258.

CONVOLVULACEAE

Erycibe bullata Ridl. ex Hoogl.

Tree 5.7 m tall, 10 cm girth. Primary lowland dipterocarp forest on clay loam soil.

Ulu Sg. Sekaloh, Niah river Sibat ak Luang S 26119.

Erycibe glomerata Blume

Treelet 2.4 m tall. Primary lowland dipterocarp forest on clay loam soil. Ulu Sg.

Sekaloh, Niah river Erwin Wright S 29107.

CUCURBITACEAE

*Gynostemma pentaphyllum (Thunb.) Makino

Small climber climbing by tendrils to 7 m. Swampy area near stream, alluvium

with limestone influence, 33 m a.s.]. Path from Pangkalan Lobang to Great Cave,

Niah J.A.R. Anderson, Sonny Tan & E. Wright S 26073.

DAVALLIACEAE

Davalia solida (G. Forst.) Sw.

Riverine forest. Along Sg. Niah, Niah N.P. Yii Puan Ching S 40155.

DENNSTAEDTIACEAE

Lindsaea scandens Hook. var. terrestris Holttum

Fronds dark shining green above, paler below. In loose litter on forest floor.

Pengkalan Lobang to Niah Caves B.L. Burtt & P.J.B. Woods B 2002.

DICHAPETALACEAE

Dichapetalum setosum Leenh.

Climber to 17 m tall. Primary lowland dipterocarp forest on clay loam soil. Ulu Sg.

Sekaloh, Niah river Benang ak Bubong S 26106.

DILLENIACEAE

Dillenia excelsa Martelli (Jack) Gilg var. pubescens (Corner) Corner ex Masamune

Tree 17 m tall, 25 cm girth. Primary lowland dipterocarp forest on clay loam soil.

158 Gard. Bull. Singapore 57 (2005)

Ulu Sg. Sekaloh, Niah river Sibat ak Luang S 26147.

Tetracera korthalsii Miq. var. subrotunda (Elmer) Hoogl.

Climber on tree. Along river bank, riverine forest. Along Niah river, Niah N.P.

Yiu Puan Ching S 40141.

DIPTEROCARPACEAE

Anisoptera costata Korth.

Tree 17 m tall, 210 cm girth. Primary lowland dipterocarp forest on clay loam soil.

Ulu Sg. Sekaloh, Niah river Edwin Wright S 29127

Anisoptera grossivenia * Slooten

Tree to 27 m tall, 230 cm girth. Primary lowland dipterocarp forest on clay loam

soil, 100 m a.s.l. Ulu Sg. Sekaloh, Niah river Sibat, Banyeng & Adenan S 26164, S

26167.

Dipterocarpus acutangulus Vesque

Tree 32 m tall, to 203 cm girth. Primary lowland dipterocarp forest on clay loam

soil, 100 m a.s.]. Ulu Sg. Sekaloh, Niah river Sibat, Banyeng & Adenan S 26168, S

26170.

Dipterocarpus applanatus * Slooten

Tree 30 m tall, 273 cm girth. Primary lowland dipterocarp forest on clay loam soil,

100 m a.s.l. Ulu Sg. Sekaloh, Niah river Sibat & Adenan § 22553, Benang &

Adenan S§ 22594.

Dipterocarpus caudatus Foxw. ssp. penangianus (Foxw.) P.S.Ashton

Tree 32 m tall, 85 cm girth. Primary lowland dipterocarp forest on clay loam soil,

100 m alt. Ulu Sg. Sekaloh, Niah river Sibat, Banyeng & Adenan S§ 26173.

Dipterocarpus caudiferus Merr.

Tree to 33 m tall, 315 cm girth. Primary lowland dipterocarp forest on clay loam

soil, 100 m a.s.l. Ulu Sg. Sekaloh, Niah river Sibat & Adenan S 22590, § 22591, $

22592.

Dipterocarpus elongatus Korth.

Tree to 30 m tall, 133 cm girth. Primary lowland dipterocarp forest on clay loam

soil, 100 m a.s.l. Ulu Sg. Sekaloh, Niah river Banyeng & Adenan S 22573.

Dipterocarpus verrucosus Foxw. ex Slooten

Tree to 30 m tall, 225 cm girth. Lowland MDF on clay loam soil, 100-150 m a.s.l.

Ulu Sg. Sekaloh, Niah river Sibat & Adenan S 22559, § 22560, Adenan bin Dillah

& Banyeng ak Nudong S 26151; Sg Sekaloh, Niah N.P. Yii Puan Ching S 40125.

*Hopea plagata (Blanco) Vidal

Tree 23 m tall, 148 cm girth. On limestone. South side of G. Subis in Sekaloh

River, Niah Sibat & Adenan § 22568.

Parashorea cf. lucida (Miq.) Kurz

Tree 23 m tall, 188 cm girth. Primary lowland dipterocarp forest on clay loam soil,

100 m alt. Ulu Sg. Sekaloh, Niah river Sibat & Adenan S 22588.

Paraboea culminicola and plants from Niah, Borneo 159

Shorea almon Foxw.

Tree 30 m tall, 308 cm girth. Primary lowland dipterocarp forest on clay loam soil,

100 m a.s.l. Ulu Sg. Sekaloh, Niah riverAdenan bin Dillah & Banyeng ak Nudong

S 26153, S 26163, Banyeng & Adenan S 22576.

Shorea argentifolia Z Symington

Tree 33 m tall, 285 cm girth. Primary lowland dipterocarp forest on clay loam soil,

100 m a.s.l. Ulu Sg. Sekaloh, Niah river Banyeng & Adenan S§ 22578, Sibat &

Adenan § 26175.

Shorea cf. atrinervosa Symington

Tree 33 m tall, 193 cm girth. Primary lowland dipterocarp forest on clay loam soil,

100 m a.s.l. Ulu Sg. Sekaloh, Niah river Sibat, Banyeng & Adenan S 26171, Sibat &

Adenan S 26176.

Shorea balanocarpoides Symington

Tree 23 m tall, 88 cm girth. Primary lowland dipterocarp forest on clay loam soil,

100 m a.s.l. Ulu Sg. Sekaloh, Niah river Sibat & Adenan S 26174.

Shorea beccariana’ Burck

Tree 31 m tall, 323 cm girth. Primary lowland dipterocarp forest on clay loam soil,

100 m a.s.1. Ulu Sg. Sekaloh, Niah river Sibat & Adenan § 22554, Sibat, Banyeng &

Adenan S 26166.

*Shorea calcicola’ P.S. Ashton

G. Subis, Miri District, Sg. Sekaloh. Sylvester S 27282.

Shorea confusa * PS. Ashton

Tree 27 m tall, 330 cm girth. Primary lowland dipterocarp forest on clay loam soil,

100 m a.s.l. Ulu Sg. Sekaloh, Niah river Banyeng & Adenan S 22575.

Shorea domatiosa’ P.S. Ashton

Tree 33 m tall, 318 cm girth. Primary lowland dipterocarp forest on clay loam soil,

100 m as.l.. Ulu Sg. Sekaloh, Niah river Sibat & Adenan S 22593, Benang &

Adenan S 22597, S 22598.

Shorea faguetiana F.Heim

Tree 27 m tall, 208 cm girth. Primary lowland dipterocarp forest on clay loam soil,

100 m a.s.l. Ulu Sg. Sekaloh, Niah river Sibat & Adenan S 22587.

Shorea faguetioides~ P.S. Ashton

Tree 30 m tall, 208 cm girth. Primary lowland dipterocarp forest on clay loam soil,

100 m asl. Ulu Sg. Sekaloh, Niah river Banyeng & Adenan S$ 22579, S 22580,

Adenan bin Dillah & Benang ak Bubong S 26161.

Bn,

Shorea fallax Meijer

Tree 23 m tall, 128 cm girth. Primary lowland dipterocarp forest on clay loam soil,

100 m a.s.]. Ulu Sg. Sekaloh, Niah river Adenan bin Dillah & Benang ak Bubong S

26160.

2 E :

Shorea ferruginea Dyer ex Brandis

Tree 30 m tall, 88 cm girth. Primary lowland dipterocarp forest on clay loam soil,

160

Gard. Bull. Singapore 57 (2005)

100 m alt. Ulu Sungei Sekaloh, Niah river Adenan bin Dillah & Banyeng ak

Nudong S 26154.

Shorea macroptera Dyer

Tree 30 m tall, 233 cm girth. Primary lowland dipterocarp forest on clay loam soil,

100 m a.s.l. Ulu Sg. Sekaloh, Niah river Benang & Adenan S 22595, Adenan bin

Dillah & Banyeng ak Nudong S 26156.

Shorea macroptera Dyer ssp. macropterifolia * PS. Ashton

Tree 30 m tall, 140 cm girth. Primary lowland dipterocarp forest on clay loam soil,

100 m a.s.]. Ulu Sg. Sekaloh, Niah river Adenan bin Dillah & Benang ak Bubong S

26157, S 26159.

Shorea mecistopteryx * Ridl.

Tree 33 m tall, to 350 cm girth. Primary lowland dipterocarp forest or MDF on

reddish clay loam soil, 100 m alt. Ulu Sg. Sekaloh, Niah river Sibat & Adenan S

22557, S 22589, Benang & Adenan § 22585, Adenan bin Dillah & Banyeng ak

Nudong S 26152, E. Wright S 29278.

Shorea ochracea™ Symington

Tree 33 m tall, to 488 cm girth. Primary lowland dipterocarp forest or MDF on

reddish clay loam soil, 100 m a.s.l. Ulu Sg. Sekaloh, Niah river Benang & Adenan

22600, Adenan bin Dillah & Benang ak Bubong § 26158.

Shorea parvistipulata * F.Heim ssp. albifolia P.S. Ashton

Tree 30 m tall, to 320 cm girth. Primary lowland dipterocarp forest or MDF on

clay loam soil, 100 m a.s.]. Ulu Sg. Sekaloh, Niah river Sibat & Adenan S$ 22561,

Adenan bin Dillah & Benang ak Bubong S 26162.

Shorea pauciflora King

Tree 27 m tall, to 293 cm girth. Primary lowland dipterocarp forest or MDF on

clay loam soil, 100 m a.s.l. Ulu Sg. Sekaloh, Niah Benang & Adenan § 22599.

‘ E

Shorea saggitata P.S. Ashton

Tree 23 m tall, to 233 cm girth. Primary lowland dipterocarp forest or MDF on

clay loam soil, 100 m a.s.l. Ulu Sg. Sekaloh, Niah Sibat & Adenan S§ 22558.

*Shorea seminis (de Vr.) Slooten

Tree 27 m tall, 2 m girth. Alluvial soils by Sg. Sekaloh at base of limestone cliff.

On southern slope of G. Subis in Sekaloh river, Niah Sibat & Adenan S 22567.

Shorea smithiana’ Symington

Tree 33 m tall, to 323 cm girth. Primary lowland dipterocarp forest or MDF on

clay loam soil, 100 m a.s.l. Ulu Sg. Sekaloh, Niah Banyeng & Adenan S$ 22577, S

22581, S 22586.

Shorea superba Symington

Tree 33 m tall, to 325 cm girth. Primary lowland dipterocarp forest or MDF on

clay loam soil, 100 m a.s.]. Ulu Sg. Sekaloh, Niah Sibat & Adenan S 22556, Adenan

bin Dillah & Banyeng ak Nudong S 26155.

Vatica badiifolia’ P.S. Ashton

Paraboea culminicola and plants from Niah, Borneo 161

Tree 31 m tall, to 298 cm girth. Primary lowland dipterocarp forest or MDF on

clay loam soil, 100 m a.s.l.. Ulu Sg. Sekaloh, Niah Sibat, Banyeng & Adenan S

26165.

*Vatica globosa * B.S. Ashton

Limestone. Niah G.D. Haviland & C. Hose Hav. 3159.

DRYOPTERIDACEAE

*Tectaria brooksii Copel.

Small terrestrial fern. On dry, shaded vertical bare limestone rock also cave and

large overhang 100 m up on steep limestone hill, light dry soil partly composed of

guano and damp also shaded limestone rockface at 67 m as.l. G. Subis, Niah

Mohidin § 21690 J.A.R. Anderson S 16024; Gua Pangomah, G. Subis, Niah J.A.R.

Anderson § 31687, Great Cave, G. Subis, Niah J.A.R. Anderson § 31949.

*Tectaria melanorachis (Baker) Copel.

Terrestrial fern. On damp and shaded limestone rockface at 67 m a.s.l. Great

Cave, G. Subis, Niah J.A.R. Anderson S 31944.

*Tectaria pleiosora (Alderw.) C. Chr.

Terrestrial fern 60-100 cm high. Markedly shaded localities on vertical limestone

rock, cave and large overhang 100 m up on steep limestone hill, light dry soil

partly composed of guano also on residual soils in semi-exposed position, slope

between caves, limestone rocks and residual soil, trees mainly felled. Gua

Pangomah, G. Subis, Niah J.A.R. Anderson S 31689; Great Cave, G. Subis; Niah

JLA.R. Anderson S 31915. i

*Tectaria sp.

Terrestrial fern. On limestone rock, foot of limestone at 17 m a.s.l. G. Brangin,

Ulu Sg. Subis, Niah N.P. Yii Puan Ching S 40180.

EBENACEAE

*Diospyros cauliflora Blume

Tree 13 m tall, 38 cm girth. On summit of dry limestone rock at base of limestone

hill. Near Great Cave, G. Subis, Niah J.A.R. Anderson S 16026.

Diospyros foxworthii Bakh.

Tree to 27 m tall, 67 cm girth. Primary MDF on clay loam soil. Ulu Sg. Sekaloh,

Niah river Sibat ak Luang S 27881, Edwin Wright S 29115.

Diospyros korthalsiana Hiern

Tree 20 m tall, 60 cm girth. Primary lowland dipterocarp forest on clay loam soil.

Ulu Sg. Sekaloh, Niah river Edwin Wright S 29102, S 29141.

*Diospyros venosa Wall. ex A.DC.

Tree 7 m tall, 30 cm girth. On ‘mor’ layer near summit of limestone hill. G. Subis,

Niah J.A.R. Anderson S 16050.

ELAEOCARPACEAE

Elaeocarpus obtusus Blume |

Tree 30 m tall, 150 cm girth. Ulu Sg. Sekaloh, Niah river Banyeng ak Nudong S

162 Gard. Bull. Singapore 57 (2005)

26144.

Elaeocarpus submonoceras Miq. ssp. lasionyx (Ridl.) R. Weibel

Tree 3 m tall, 20 cm girth. Riverine forest. Along Niah river, Niah N.P. Yii Puan

Ching S 40140.

ERICACEAE

*Vaccinium leptanthum Miq.

Epiphyte or straggling shrub. Near or on summit of limestone hill. G. Subis, Niah

J.A.R. Anderson S 16048; G. Brangin, Niah N.P. Yii Puan Ching S 40181.

EUPHORBIACEAE

Agrostistachys longifolia (Wight) Benth. ex Hook.f. var. leptostachya (Pax & K.

Hoffm.) Whitmore

Tree 7 m tall. Summit of small sandstone hill, primary forest on clay loam soil.

Between mouth of Sg. Sekaloh and G. Subis, Niah J.A.R. Anderson S 27592.

*Antidesma leucopodum Miq.

G. Subis area, Niah Mohidin S 21646.

Antidesma cf. pendulum Hook.f.

Tree 7 m tall, 10 cm girth. Primary lowland dipterocarp forest on clay loam soil.

Ulu Sg. Sekaloh, Niah river Benang ak Bubong S 27862.

*Aporosa grandistipulata Merr.

Tree c. 3 m tall, 5 cm diam. Limestone area, relatively open Dipterocarpus forest.

G. Subis, to right of Sg. Sekalau, at footpath from Kuala Sekalau to Bukit Drusau

ALP: Fuchs 21271.

Aporosa nigricans Hook.f.

Tree 7 m tall. Primary lowland dipterocarp forest on clay loam soil. Ulu Sg.

Sekaloh, Niah river Sonny Tan & E. Wright S 26101.

Aporosa nitida Merr.

Tree 8 m tall, 15 cm girth. Primary lowland dipterocarp forest on clay loam soil.

Ulu Sg. Sekaloh, Niah river Erwin Wright S 29118.

*A porosa prainiana King ex Gage

Tree c. 7 m tall, to 10 cm thick. Limestone area 100 m a.s.l., also clay loam soil,

sedimentary rocks relatively open MDF. Ulu Sg. Sekaloh and to right of Sg.

Sekalau at footpath from Kuala Sekalau to Bukit Drusau, G. Subis H.P. Fuchs

21264, 21287; J.A.R. Anderson, Sonny Tan & E. Wright S 26064, E. Wright S

LILDD:

Baccaurea lanceolata (Miq.) Muell.-Arg.

Tree 17 m tall, 55 cm girth. Alluvial soils by Sg. Sekaloh at base of limestone cliff.

S. side of G. Subis, Niah J.A.R. Anderson, Sonny Tan & E. Wright S 27552.

Baccaurea sarawakensis Pax & K. Hoffm.

Tree 12 m tall, 30 cm girth. Primary lowland dipterocarp forest on clay loam soil.

Ulu Sg. Sekaloh, Niah river Sibat ak Luang S 27859.

Paraboea culminicola and plants from Niah, Borneo 163

Baccaurea trunciflora Merr.

Tree 7 m tall, 20 cm girth. Primary lowland dipterocarp forest on clay loam soil.

Ulu Sg. Sekaloh, Niah river Banyeng ak Nudong S 26142.

Cleistanthus pyrrhocarpus Airy Shaw

Slender tree to 8 m tall. MDF between two steep slopes at 85 m a.s.l. Niah N.P. Yii

Puan Ching S 40070.

Glochidion rubrum Blume

Tree 8 m tall, 35 cm girth. Riverine forest c. 17 m a.s.l., loamy soil. Kuala Sg Sekaloh,

Niah N.P. Yii Puan Ching S 40102.

Drypetes crassipes Pax & K. Hoffm.

Tree 17 m tall, 100 cm girth. Primary lowland dipterocarp forest on clay loam soil.

Ulu Sg. Sekaloh, Niah river Sibat ak Luang S 26129, Erwin Wright S 29121.

Koilodepas longifolium Hook.f.

Tree 7 m tall, 15 cm girth. Primary lowland dipterocarp forest on clay loam soil.

Ulu Sg. Sekaloh, Niah river Erwin Wright S 29117.

Macaranga bancana (Miq.) Muell.-Arg.

Tree 7 m tall, 30 cm girth. Riverine forest c. 17 m a.s.l., soil loamy. Kuala Sg

Sekaloh, Niah N.P. Yii Puan Ching S 40108.

*Mallotus dispar (Blume) Muell.-Arg.

Bushy tree 7 m tall 25 cm girth. On limestone rocks, base of limestone hill also

ravine between two limestone hills, limestone pavement and small, heavily eroded

limestone hillocks. G. Subis, path to Great Cave J.A.R. Anderson S 31693; Outside

Great Cave, G. Subis, Niah J.A.R. Anderson S 31910.

*Mallotus korthalsii Muell.-Arg.

Tree c. 3 m tall, 8 cm diam. In small river valley between limestone boulders,

relatively open Dipterocarpus forest, 150 m a.s.l. G. Subis to right of Sg. Sekalau,

at footpath from Kuala Sekalau to Bukit Drusau H.P. Fuchs 21272, 21273.

Mallotus wrayii King ex. Hook.f.

Tree 7 m tall, 12.5 cm girth. MDF on clay loam soil, 100 m a.s.l. Ulu Sg. Sekaloh,

Niah river J.A.R. Anderson, Sonny Tan & E. Wright S 26063.

Margaritaria indica (Dalziel) Airy Shaw

Tree 10 m tall, 40 cm girth. Riverine forest. Along Sg. Niah, Niah N.P. Yii Puan

Ching S 40160.

*Neoscortechinia sumatrensis S. Moore

Tree 17 m tall, 15 cm diam. with stilt roots. On limestone. Along the plankwalk to

Niah Cave, Niah N.P. Rena George S 39936.

Pimelodendron griffithianum (Muell.-Arg.) Benth.

Tree 11.7 m tall, 18 cm girth. Primary lowland dipterocarp forest on clay loam soil.

Ulu Sg. Sekaloh, Niah river. Sibat ak Luang S 26135.

Ptychopyxis grandis Airy Shaw

164 Gard. Bull. Singapore 57 (2005)

Tree 8.3 m tall, 30 cm girth. Primary lowland or MDF on clay loam soil. Ulu Sg.

Sekaloh, Niah river. Benang ak Bubong S 26112, Erwin Wright S 29149.

*Sauropus androgynus (L.) Merr.

Shrub 1 m tall. Cave and large overhang 100 m up on steep limestone hill, light

dry soil, partly composed of guano. Gua Pangomah, G. Subis, Niah J.A.R. Anderson

S 31682.

Trigonostemon merrillianus Airy Shaw

Tree 3 m tall, 10 cm girth. Primary lowland dipterocarp forest on clay loam soil.

Ulu Sg. Sekaloh, Niah river Sibat ak Luang S 26123, Erwin Wright S 29123.

FABACEAE

*Bauhinia endertii K. & S.S. Larsen Creeper on limestone or climber on tree.

Riverine forest along Sg Niah also limestone hill summit 110 m a.s.l. Niah N.P. Yii

Puan Ching S 40157; G. Brangin, Niah N.P. Yii Puan Ching S 40182.

Bauhinia excelsa (Blume ex Miq.) Prain

Climber to 40 m tall on tree, 30 cm girth at base. Mudstone outcropping from

river bank of Nangau Sekalan also disturbed belian forest, alluvial soils at base of

limestone hill. Ulu Sg Sekaloh, Niah River J.A.R. Anderson S 27291; Between

mouth of Sg. Sekaloh and G. Subis, Niah J.A.R. Anderson S 27581.

Derris elegans Benth.

Climber. Riverine forest. Along Sg. Niah, Niah N.P. Yii Puan Ching S 40117.

*Dialium indum L.

Tree 25 m tall, 200 cm girth. Foot of limestone 17 m a.s.l. G. Brangin, Ulu Sg.

Subis, Niah N.P. Yii Puan Ching S 40173.

Parkia sumatrana Miq.

Niah G.D. Haviland & C. Hose Haviland 3273.

*Saraca declinata (Jack) Miq.

Tree to 20 m tall and 60 cm girth. Base of limestone hill on limestone rocks and

lower slopes of steep limestone hill, numerous limestone boulders also primary

lowland dipterocarp forest on clay loam soil. Ulu Sg. Sekaloh, Niah river Sibat ak

Luang S 26150; South side of G. Subis in Sekaloh river, Niah J.A.R. Anderson,

Sonny Tan & E. Wright S 27554; G. Subis, path to Great Cave J.A.R. Anderson §

31694.

*Whitfordiodendron niewenhuisii (J.J. Sm.) Dunn

Climbing shrub. Limestone area, relatively open Dipterocarpus forest, 150 m a.s.l.

G. Subis to right of Sg. Sekalau, at footpath from Kuala Sekalau to Bukit Drusau

EP. Fuchs 21262.

FAGACEAE

Lithocarpus meijeri Sopadmo

Tree 23 m tall, 240 cm girth. Primary forest on ridge at 133 m a.s.l. Niah Dan bin

Hj. Bakar S 16524.

FLACOURTIACEAE

Paraboea culminicola and plants from Niah, Borneo 165

Casearia rugulosa Blume

Tree 4 m tall, 7.5 cm girth. Primary lowland dipterocarp forest on clay loam soil.

Ulu Sg. Sekaloh, Niah river Erwin Wright S 29142.

Hydnocarpus subfalcata Merr.

Tree 13.3 m tall, 45 cm girth. Primary lowland dipterocarp forest on clay loam soil.

Ulu Sg. Sekaloh, Niah river Erwin Wright S 29110.

*Osmelia philippina (Turcz.) Benth.

Tree 10 m tall, 28 cm girth. On limestone rocks near river and alluvial soils at base

of limestone hill in disturbed belian forest. On southern slopes of G. Subis in the

Sekaloh river. Niah J.A.R. Anderson, Sonny Tan & E. Wright S 26083; between

mouth of Sg. Sekaloh and G. Subis, Niah J.A.R. Anderson § 27579.

Pangium edule Reinw.

Tree 33 m tall, 90 cm girth. On alluvial soil at base of limestone hill. Just beyond

Great Cave, G. Subis. Niah J.A.R. Anderson S 16428.

Ryparosa acuminata Merr.

Tree 7 m tall 13 cm girth. Primary lowland dipterocarp forest on clay loam soil.

Ulu Sg. Sekaloh, Niah river Sibat ak Luang S 26122.

Ryparosa kostermansii Sleumer

Tree 17 m tall, 75 cm girth. MDF on clay loam soil, 100 m a.s.]. Ulu Sg. Sekaloh,

Niah river J.A.R. Anderson, Sonny Tan & E. Wright S 26054.

FLAGELLARIACEAE

Flagellaria indica L.

Climber on tree. Riverine forest. Along Sg. Niah, Niah N.P. Yii Puan Ching S$

40113.

GESNERIACEAE 2

*Paraboea culminicola K.G. Pearce

Large herb 75-120 cm tall. On bare, exposed limestone rock near summit of

limestone pinnace or crest of limestone hill, ‘mor’ soil, 170 m a.s.l. Niah J.A.R.

Anderson § 16045; Southern slopes of G. Subis, near Sekaloh river Sonny Tan &

E. Wright S 27279.

Included in Pearce, 2004 as Paraboea sp.

GNETACEAE

Gnetum cuspidatum Blume

Woody climber c. 33 m long, 15 cm girth on large tree. Riverine forest c. 17 m

a.s.l., soil loamy also primary lowland dipterocarp forest on clay loam soil. Kuala

Sg. Sekaloh, Niah N.P. Yii Puan Ching S 40107; Ulu Sg. Sekaloh, Niah River

Erwin Wright S 29105.

*Gnetum diminutum Marker.

Climber to 27 m. Slopes of limestone hill, immediately below cliff, 133 m a.s.l.,

litter layer overlying limestone boulders also limestone hill, lower summit ridge,

limestone pinnacles and ‘mor’ soil. South side of G. Subis in Sekaloh river, Niah

J.A.R. Anderson, Sonny Tan & E. Wright S 27560; Sg. Sekaloh, G. Subis, Niah

166 Gard. Bull. Singapore 57 (2005)

J.A.R. Anderson 31954.

Gnetum gnemonoides Brongn.

Entwined round tree. Mudstone outcropping from river bank. Nangau Sekalan,

Ulu Sg. Sekaloh, Niah river J.A.R. Anderson S 27286.

*Gnetum klossii Merr. ex Marker.

Climber to 13 m. Lower slopes of steep limestone hill, numerous limestone boulders.

South side of G. Subis in Sekaloh river, Niah J.A.R. Anderson, Sonny Tan & E.

Wright S 27556

Gnetum loerzingii Markgr.

Scrambler to 13 m high. Remnant forest near river bank. Niah N.P. Bernard Lee

M.H. S 40066.

Gnetum macrostachyum Hook.f.

Climber 27 m high. Primary lowland dipterocarp forest on clay loam soil. Ulu Sg.

Sekaloh, Niah river Benang ak Bubong S 26116.

HYMENOPHYLLACEAE

Crepidomanes bipunctatum (Poir.) Copel.

G. Subis area, Niah Mohidin S 21625.

Crepidomanes christii (Copel.) Copel.

G. Subis area, Niah Mohidin S 21614.

HYPERICACEAE

Cratoxylon formosum (Jack) Dyer ssp. formosum

Tree c. 10 m tall, 10 cm girth. Riverine forest along Sg. Niah. Niah N.P. Yii Puan

Ching S 40152.

ICACINACEAE

Cantleya corniculata (Becc.) R.A. Howard

Tree 20 m tall, 80 cm girth. Lowland MDF, 80 m a.s.l., soil sandy loam. Sg.

Sekaloh, Niah N.P. Yii Puan Ching S 40131.

*Gomphandra cumingiana (Miers) F.-Vill.

Treelet 5 m tall, 8 cm girth. 17 m below summit ridge of limestone hill, limestone

outcropping and ‘mor’ soil present, also lower slopes of limestone hill, numerous

limestone boulders and primary lowland dipterocarp forest on clay loam soil.

Southern slopes of G. Subis near Sekaloh river, Niah Sonny Tan & E. Wright S$

27263, Ulu Sungei Sekaloh, Niah river Erwin Wright S 29104; J.A.R. Anderson S$

31934.

LAMIACEAE

Anisomeles indica O. Kuntze

In secondary growth on ground. Niah Ahmad 47.

Pogostemon auricularis (L.) Hassk.

Niah Haviland 3598.

LAURACEAE

Paraboea culminicola and plants from Niah, Borneo 167

Litsea lancifolia (Roxb. ex Wall.) Hook.f.

Tree 8 m tall, 25 cm girth. MDF with reddish clay loam soil. Ulu Sg. Sekaloh, Niah

Erwin Wright S 29262.

Nothaphoebe sarawacensis Gamble

Tree 4 m tall, 45 cm girth. Along Sg Niah, Niah N.P. Yii Puan Ching S 40119.

LECY THIDACEAE

Barringtonia lanceolata’ (Ridl.) Payens

Tree 12 m tall, 25 cm girth. MDF on clay loam soil. Ulu Sg. Sekaloh, Niah river

Erwin Wright S 29124.

Barringtonia macrostachys (Jack) Kurz

Tree 8 m tall, 38 cm girth. MDF with reddish clay loam soil. Ulu Sg. Sekaloh, Niah

Erwin Wright S 29269.

LEEACEAE

Leea indica (Burm.f.) Merr.

Treelet 2.7 m tall. Lowland dipterocarp forest on clay loam soil. Ulu Sg. Sekaloh,

Niah river Sibat ak Luang S 27879.

LORANTHACEAE

Lepeostegeres beccarii (King) Gamble

Epiphyte. Riverine forest. Along Niah river, Niah N.P. Yii Puan Ching S 40149.

*Macrosolen beccarii Tiegh.

Parasite on small tree. On limestone rocks near river. On southern slopes of G.

Subis in Sekaloh river, Niah J.A.R. Anderson, Sonny Tan & E. Wright S 26081.

Trithecanthera cf. xiphostachys Tiegh.

Large parasite in crown of Pometia pinnata at 20 m. Alluvial soils by Sg. Sekaloh

at base of limestone cliff. On southern slopes of G. Subis in Sekaloh river, Niah

J.A.R. Anderson, Sonny Tan & E. Wright S 26097.

MALVACEAE

Sida acuta Burm.f.

In secondary growth on ground. Rumah Pasang, Niah Ahmad 49.

Urena lobata L. ssp. lobata f. sinuata (L.) Borss.

120-150 cm tall. In jungle on wet ground. Rumah Pasang, Niah Ahmad 19.

MARANTACEAE

Phrynium capitatum Willd.

Herb 1 m tall. In jungle on ground, also alluvial soil at base of limestone hill near

river. Rumah Pasang, Niah Ahmad 41; Southern slopes of G. Subis near Sekaloh

river, Niah Sonny Tan & E. Wright S 27257.

Schumannianthus dichotomus (Roxb.) Gagnep.

Shrub c. 2 m high. Secondary shrubby vegetation along small creek to right of Sg.

Sekalau, to left of footpath from Kuala Sekalau to Bukit Drusau, near Kuala

Skaloh longhouse, limestone area, G.Subis H.P. Fuchs 21284.

168 Gard. Bull. Singapore 57 (2005)

MELASTOMATACEAE

Dissochaeta annulata Hook.f. ex Triana

G. Subis area. Niah Mohidin S 21636.

Dissochaeta hirsuta Hook.f. ex Triana

Climber on tree. Riverine forest. Along Sg. Niah, Niah N.P. Yii Puan Ching S

40124.

Dissochaeta rostrata Korth.

Climber entwined round tree. Lowland dipterocarp forest on clay loam soil. Ulu

Sg. Sekaloh, Niah river Sibat ak Luang S 27878.

Medinilla macrophylla Blume

Epiphyte 27 m high on tree. Primary lowland dipterocarp forest on clay loam soil.

Ulu Sg. Sekaloh, Niah river Benang ak Bubong S 26105.

*Memecylon paniculatum Jack

Tree c. 6 m tall, 10 cm thick. Limestone area, relatively open Dipterocarpus forest,

150 m a.s.l. G. Subis, to right of Sg. Sekalau, at footpath from Kuala Sekalau to

Bukit Drusau H.P. Fuchs 21263.

Memecylon scolopacinum~ Rid.

Shrub 2 m high. Primary lowland dipterocarp forest on clay loam soil. Ulu Sg.

Sekaloh, Niah river Sibat ak Luang S 26130.

*Pachycentria microsperma Becc.

Epiphyte on limestone hill summit. Limestone hill summit 110 m a.s.l. G. Brangin.

Niah N.P. Yii Puan Ching S 40183.

Plethiandra motleyi Hook.f.

Epiphyte 8 m high on Dipterocarpus oblongifolia. Mudstone outcropping from

river bank. Nanagu Sekalan, Ulu Sg. Sekaloh, Niah river J.A.R. Anderson S 27287.

*Plethiandra robusta (Cogn.) Nayar ex Bakh.

Epiphyte 23 m high on tree trunks, and on rocks. Limestone hill, lower summit

ridge, limestone pinnacles and ‘mor’ soil also primary lowland dipterocarp forest

on clay loam soil. South side of G. Subis in Sekaloh river, Niah J.A.R. Anderson,

Sonny Tan & E. Wright S 27562; Ulu Sg. Sekaloh, Niah river Erwin Wright S$

29109.

Pternandra crassicalyx J.F. Maxwell

Tree 8 m tall, 15 cm girth. Primary lowland dipterocarp forest on clay loam soil.

Ulu Sg. Sekaloh, Niah river Sibat ak Luang S 26131.

Pternandra multiflora Cogn.

Tree 15 m tall, 60 cm girth. Primary lowland dipterocarp forest on clay loam soil.

Ulu Sg. Sekaloh, Niah river Erwin Wright S 29135.

Pternandra rostrata (Cogn.) M.P. Nayar

Tree to 12 m tall, 28 cm girth. Primary lowland dipterocarp forest on clay loam

soil. Ulu Sg. Sekaloh, Niah river Sibat ak Luang S 27855, Erwin Wright S 29129.

Paraboea culminicola and plants from Niah, Borneo 169

MELIACEAE

* Aglaia elaegoinidea (A. Juss.) Benth.

Tree 7 m tall, 20 cm girth. Partially exposed situation on dry limestone cliff. Niah

J.A.R. Anderson S 16032.

Aglaia elliptica Blume

Tree. Riverbank on sandy clay soil. Ulu Sg. Sekaloh, Niah river J.A.R. Anderson S

27297.

*A glaia odoratissima Blume

Tree to 27 m tall, 90 cm girth. MDF with reddish clay loam soil, also lower slopes

of limestone hill with numerous limestone boulders, steep slopes of limestone hill

with deep litter layer and narrow limestone ridge at 130 m a.s.]. Ulu Sg. Sekaloh

Sibat ak Luang S 26149, Benang ak Bubong S 27861, Erwin Wright S 29274;

Southern slopes of G. Subis near Sekaloh river Sonny Tan & E. Wright S 27276,

J.A.R. Anderson & Sonny Tan § 27573; Niah N.P. Paul Chai S 40059.

*A glaia speciosa Blume

Tree 7 m tall, 25 cm girth. Knife-edge ridge near summit of limestone hill, limestone

pinnacles and deep (45 cm) ‘mor’ soil. South side of G. Subis in Sekaloh river,

Niah J.A.R. Anderson, Sonny Tan & E. Wright S 27569.

Aglaia tomentosa Teijsm. & Binn.

Tree 6 m tall, 8 cm girth. Primary lowland dipterocarp forest on clay loam soil.

Ulu Sg. Sekaloh, Niah river Banyeng ak Nudong S 26140.

Chisocheton ceramicus (Miq.) C.DC.

Tree 15 m tall, 50 cm girth. Riverine forest. Along Niah river, Niah N.P. Yii Puan

Ching S 40144.

Chisocheton sarawakanus (C.DC.) Harms

Treelet 5 m tall. Primary lowland dipterocarp forest on clay loam soil. Ulu Sg.

Sekaloh, Niah river Erwin Wright S 29148.

*Dysoxylum alliaceum (Blume) Blume

Tree to 33 m tall and 180 cm girth. Steep slopes of limestone, numerous limestone

boulders, deep litter layer also primary lowland dipterocarp forest on clay loam

soil. South side of G. Subis in Sekaloh river, Niah J.A.R. Anderson, Sonny Tan &

E. Wright S 27576; Ulu Sg. Sekaloh, Niah river Erwin Wright S 29114.

Dysoxylum brachybotrys Merr.

S. Subis area, Niah Mohidin S 21643.

Sandoricum borneense Mig.

Tree 18 m tall, 70 cm girth. Riverine forest. Along Niah river, Niah N.P. Yii Puan

Ching S 40145.

Walsura dehiscens T.P. Clark

Tree 13 m tall, 40 cm girth. Lowland MDF 80 m a.s.l., soil sandy loam. Sg Sekaloh,

Niah N.P. Yii Puan Ching S 40128.

MENISPERMACEAE

170 Gard. Bull. Singapore 57 (2005)

Fibraurea tinctoria Lour.

Climber to 5 m on Agrostistachys. Primary tropical heath forest, 33 m a.s.l. Path

from Pangkalan Lobang to Great Cave, Niah J.A.R. Anderson, Sonny Tan & E.

Wright S 26070.

MONIMIACEAE

*Kibara coriacea (Blume) Tulasne

Tree to 10 m tall, 25 cm girth or 25 cm diam. Limestone forest at 80 m a.s.l. and

along plankwalk to Great Cave. Niah N.P. Rena George S 39938, Yii Puan Ching

S 40200.

MORACEAE

Ficus aurata (Miq.) Miq.

Tree 5 m tall, 18 cm girth. Primary lowland dipterocarp forest on clay loam soil.

Ulu Sungei Sekaloh, Niah river Erwin Wright S 29128.

*Ficus aurita Blume var. auriculifera (Merr.) Corner

Bushy tree 6 m tall. Base of limestone hill on limestone rocks at 17 m a.s.l. G.

Subis area, Niah Mohidin S 21604; Path to Great Cave, G. Subis J.A.R. Anderson

S 31695; G. Brangin, Ulu Sg. Subis, Niah N.P. Yii Puan Ching S 40169.

Ficus beccarrii King

Tree 3 m tall. Disturbed belian forest, alluvial soils at base of limestone hill.

Between mouth of Sg. Sekaloh and G. Subis, Niah J.A.R. Anderson S 27589.

*Ficus benjamina L.

Tree 23 m tall, 90 cm girth or shrub. Vertical exposed limestone cliff, also swampy

alluvial soils at base of limestone hill subject to periodic inundation. Niah J.A.R.

Anderson § 16029; On southern slopes of G. Subis in the Sekaloh river, Niah

J.A.R. Anderson, Sonny Tan & E. Wright S 26087.

*Ficus binnendykii (Miq.) Miq. var. coriacea Corner

Tree 13 m tall, 28 cm girth. Primary lowland dipterocarp forest on rocks beside

waterfall. Ulu Sg. Sekaloh, Niah river Benang ak Bubong S 27871.

Ficus delosyce Corner

Epiphyte 3 m tall, 12 cm girth. Riverine forest. Along Sg. Niah, Niah N.P. Yii

Puan Ching S 40122.

Ficus fistulosa Reinw. ex Blume

Tree 5 m tall, 12 cm girth. River bank of Nangau Selakan on clay soil. Ulu Sg.

Sekaloh, Niah river J.A.R. Anderson S 27600.

Ficus laevis Blume var. tomentosa King

G. Subis area, Niah Mohidin S 21671.

Ficus lepicarpa Blume

Tree 8 m tall, 50 cm girth. Swampy alluvial soils at base of limestone hill subject to

periodic inundation, also at foot of limestone hill near stream 50 m a.s.]. Ulu Sg.

Subis, Niah N.P. Bernard Lee S 40086; On southern slopes of G. Subis in the

Sekaloh river, Niah Mohidin S 21609, J.A.R. Anderson, Sonny Tan & E. Wright S

26089.

Paraboea culminicola and plants from Niah, Borneo 7h

Ficus magnoliifolia Blume

Tree 23 m tall, 60 cm diameter. On sandy soil at starting point of path to Niah

Cave. Niah N.P. Rena George S 39932.

*Ficus pisocarpa Blume

Climber entwined round tree. Hillslope on limestone rocks, 167 m a.s.l. Ulu Sg.

Sekaloh, Niah river J.A.R. Anderson S 27570.

*Ficus sundaica Blume var. sundaica

Tree 17 m tall. Foot of limestone 17 m a.s.l. G. Brangin, Ulu Sg. Subis, Niah N.P.

Yui Puan Ching S 40175.

Ficus treubii_ King

Tree 10 m tall, 20 cm girth. On rocks beside swift flowing stream. Niah Banyeng

Bubong S 27869.

Ficus uniglandulosa Wall. ex Miq.

Climber 15 m high. Primary lowland dipterocarp forest on clay loam soil. Ulu Sg.

Sekaloh, Niah river Erwin Wright S 29132.

MYRISTICACEAE 1

Endocomia virella W.J. de Wilde

Tree 20 m tall, 50 cm diam. Along path to Niah Cave, Niah N.P. Rena George S

39933.

Gymnacranthera farquhariana (Hook.f. & Thom.) Warb. var. zippeliana (Miq.)

R.T.A.Schouten.

Tree 23 m tall, 50 cm girth. Primary lowland dipterocarp forest on clay loam soil.

Ulu Sg. Sekaloh, Niah river Sonny Tan & Benang ak Bubong S 26103.

Knema curtisii (King) Warb. var. curtisii

G. Subis area, Niah Mohidin S 21659.

Knema latifolia Warb.

Tree 19 m tall, 30 cm girth. Primary lowland dipterocarp forest on clay loam soil.

Ulu Sg. Sekaloh, Niah river Benang ak Bubong S 26107.

Knema tridactyla’ Airy Shaw ssp. tridactyla

Tree to 5 m tall, 8 cm girth. Primary lowland dipterocarp forest on clay loam soil.

Ulu Sg. Sekaloh, Niah river Sibat ak Luang S 26128, Erwin Wright S 29136.

Myristica iners Blume ;

Tree 22 m tall, 150 cm girth. MDF between two steep slopes at 85 m a.s.]. Niah

N.P. Normah Yusoff S 40069.

Myristica malaccensis Hook.f.

Tree 15 m tall, 40 cm girth. Primary lowland dipterocarp forest on clay loam soil.

Ulu Sg. Sekaloh, Niah river Erwin Wright S 29145.

MYRSINACEAE

* Ardisia breviramea Mertr.

Treelet to 2.7 m tall. Lower slopes of limestone hill, large limestone boulders. Sg.

172 Gard. Bull. Singapore 57 (2005)

Sekaloh, G. Subis, Niah J.A.R. Anderson § 31923.

Ardisia livida Mez

Treelet 6 m, 15 cm girth. MDF on clay loam soil. Ulu Sg. Sekaloh, Niah river

J.A.R. Anderson, Sonny Tan & E. Wright S 26055.

MYRTACEAE

*’ Eugenia aff. glanduligera Ridl.’

Tree 49 cm girth. Primary forest, lower slopes of limestone mountain. South side

of mountain near Sg. Sekaloh, G. Subis, Niah J.A.R. Anderson S 31641.

‘Eugenia prasiniflora Ridl.’

Niah Haviland & C. Hose, Haviland. 3127A.

*Syzygium aff. castaneum (Merr.) Merr. & Perry

Limestone. Niah Haviland & C. Hose Hav. 3215.

* Syzygium caudilimbum (Merr.) Merr. & Perry

Tree 10 m tall, 33 cm girth. Knife-edge ridge near summit of limestone hill,

limestone pinnacles and deep (45 cm) ‘mor’ soil, also lower slopes of limestone

hill. South side of G. Subis in Sekaloh river, Niah J.A.R. Anderson, Sonny Tan &

E. Wright S 27274, S 27572.

Syzyium foxworthyianum (Ridl.) Merr. & Perry

G. Subis area, Niah Mohidin S 21666.

*Syzygium nigricans (King) Merr. & Perry

Tree 10 m tall, 35 cm girth. Hill slope on limestone rocks, 167 m a.s.l. Ulu Sungei

Sekaloh, Niah river J.A.R. Anderson S 27559.

Syzygium pycnanthum Merr. & Perry

Tree 10 m tall, 45 cm girth. On mudstone outcropping from river bank and

riverbank on sandy clay soil. Nangau Sekaloh, Ulu Sg. Sekaloh, Niah river J.A.R.

Anderson S 27293, § 27596.

Syzygium remotiflorum (Ridl.) Merr. & Perry

Tree 17 m tall, 60 cm girth. On riverine alluvium. Niah Dan bin Hj. Bakar S 16526.

OCHNACEAE

Campylospermum serratum (Gaertn.) Bittrich & M.C.E. Amaral

Tree 3 m tall. Disturbed belian forest, alluvial soils at base of limestone hill.

Between mouth of Sg. Sekaloh and G. Subis, Niah J.A.R. Anderson S 27580.

PASSIFLORACEAE

Passiflora foetida L.

Climbing on small tree. In jungle on ground. Kg. Tangap, Niah Ahmad 25.

PIPERACEAE

*Piper arborescens Roxb.

Root climber to 5 m. Knife-edge ridge near summit of limestone hill, limestone

pinnacles and deep (45 cm) ‘mor’ soil, South side of G. Subis in Sekaloh river,

Niah Mohidin S 21629; J.A.R. Anderson, Sonny Tan & E. Wright S 27571.

Paraboea culminicola and plants from Niah, Borneo 173

*Piper vestitum C.DC.

Shrublet 60 cm high or creeper on slope of limestone hills also on limestone rocks

at 50 m a.s.l. and on residual soils. Great Cave, G. Subis. Niah J.A.R. Anderson S

31919, G. Brangin, Niah National Park Yii Puan Ching S 40085, S 40188.

PITTOSPORACEAE

*Pittosporum ferrugineum Aiton

Small tree or shrub. On exposed rockface immediately above large cave overhang

100 m up on steep limestone hill, light dry soil, partly composed of guano. Gua

Pangoma, G. Subis, Niah J.A.R. Anderson S 31680.

POACEAE

Phragmites karka (Retz.) Trin. ex Steud.

G. Subis area, Niah Mohidin S 21635.

POLYGALACEAE

Xanthophyllum flavescens Roxb.

Tree to 23 m tall, 210 cm girth. Primary lowland dipterocarp forest on clay loam

soil, also riverine forest along Sg. Niah. Ulu Sg. Sekaloh and Sg Niah Sibat ak

Luang S 27852, Benang ak Bubong S 27883, S 27884, Erwin Wright S 29131, S

29140, S 29150, Yii Puan Ching S 40151.

Xanthophyllum velutinum Chod.

Tree 16 m tall, 40 cm girth. Between two steep slopes, MDF at 85 m a.s.]. Niah

N.P. Bernard Lee S 40071.

PSILOTACEAE

*Psilotum complanatum Sw.

Epiphyte at low height on tree. Limestone hill, lower summit ridge; limestone

pinnacles and ‘mor’ soil. South side of G. Subis in Sekaloh river, Niah J.A.R.

Anderson, Sonny Tan & E. Wright S 27566.

PTERIDACEAE

Pteris longipinnula Wall. ex J. Agardh

G. Subis area, Niah Mohidin S 21610.

*Pteris vittata L.

On rock near entrance, Niah Cave. Niah Ahmad 56: G. Subis area. Niah Mohidin

S 21696.

RHAMNACEAE z

Zizyphus borneensis Mert.

Climber creeping on tree. Riverine forest c. 17 m a.s.l. Kuala Sg. Sekaloh, Niah

N.P. Yii Puan Ching & Bernard Lee S 40106.

Zizyphus horsfieldii Miq.

Climber entwined round tree. Mudstone outcropping from river bank of Nangau

Sekaloh. Ulu Sg. Sekaloh, Niah river J.A.R. Anderson § 27284.

RHIZOPHORACEAE

*Carallia brachiata (Lour.) Merr.

Tree 43 cm girth. Lower flanks of limestone mountain, primary forest. South side

174 Gard. Bull. Singapore 57 (2005)

of mountains near Sg. Sekaloh, G. Subis, Niah J.A.R. Anderson S 31642.

RUBIACEAE

*Acranthera multiflora Valeton

Succulent herb. On steep slope of limestone hill at 80 m a.s.]. Niah N.P. Bernard

Lee S 40076.

*Argostemma borragineum Blume

Herb 15-23 cm tall, stock woody. Crevices on vertical shady limestone rocks and

in gully between limestone hill, heavily dissected limestone rocks and pavement,

mineral soil very sparse. Pengkalan Lobang to Niah Caves B.L. Burtt & P.J.B.

Woods B 2005; Gunong Subis J.A.R. Anderson S 31672.

*Argostemma havilandii Rid.

Herb 7.5 cm tall. On limestone rock face at 67 m a.s.l., damp and shaded. Great

Cave, G. Subis, Niah J .A.R. Anderson S 31946.

*Canthium sp.

Tree 10 m tall, 60 cm girth. Narrow limestone ridge at 130 m a.s.l. Niah N.P. Yii

Puan Ching S 40058.

Cephaelis sp.

Shrublet 30-37 cm tall. Disturbed belian forest, alluvial soils at base of limestone

hill. Between mouth of Sg. Sekaloh and G. Subis J .A.R. Anderson S§ 27586.

Chasallia curviflora (Wall.) Thw.

Niah Haviland & C. Hose Haviland 3454A.

Coptosapelta sp.

Climber on large tree. Lowland MDF, 80 m a.s.l., soil sandy loam. Sg. Sekaloh,

Niah N.P. Yii Puan Ching S$ 40134.

*Diplospora singularis Korth.

Small tree 40 cm girth. Ravine between two limestone hills, limestone pavement

and small heavily eroded limestone hillocks. Outside Great Cave, G. Subis, Niah

J.A.R. Anderson § 31909.

Gaertnera vaginans (DC.) Merr. ssp. junghuhniana (Miq.) Beusek.

Tree 3.3 m tall. Primary MDF on reddish clay loam soil. Ulu Sungei Sekaloh, Niah

Banyeng ak Nudong S 26139, Erwin Wright S 29267.

Geophila aff. hirta ( L.) Pearson

Scrambling herb. MDF, yellow clay soil. Niah N.P. Paul Chai et al. S 40053.

Hedyotis congesta Wall. & G. Don

Niah Haviland & C. Hose Haviland 3459A.

*Hydnophytum formicarum Jack

Shrublet 67 cm tall. Crest of limestone hill, limestone pinnacles on ‘mor’ soil.

Southern slopes of G. Subis near Sekaloh river Haviland & C. Hose Haviland

3448, J.LA.R. Anderson § 16043, Sonny Tan & E. Wright S 27280.

Paraboea culminicola and plants from Niah, Borneo . MES

Ixora brevicaudata Bremek.

Treelet 2.3 m tall. Lowland dipterocarp forest on clay loam soil. Ulu Sg. Sekaloh,

Niah river Mohidin S 21642; Sibat ak Luang S 27880.

Ixora stenophylla (Korth.) Kuntze

Shrub 1 m tall. Primary lowland dipterocarp forest on clay loam soil. Ulu Sg.

Sekaloh, Niah river Erwin Wright S 29116.

Ixora woodii Bremek.

Treelet 6 m tall, 13 cm girth. Primary MDF on a ridge and on clay loam soil,

sedimentary rocks 80—100 m a.s.]. Ulu Sg. Sekaloh, Niah river Benang ak Bubong

S 26110; Erwin Wright S 27252.

Lasianthus cf. maingayi Hook.f.

G. Subis area, Niah Mohidin S 21641.

Lucinaea membranacea King

Treelet 3.3 m high, 8 cm girth. Primary lowland dipterocarp forest on clay loam

soil. Ulu Sg. Sekaloh, Niah river Erwin Wright S 29134.

*TLudekia borneensis C.E,Ridsdale

Tree 25 m tall, 200 cm girth. Foot of limestone, 17 m a.s.l. G. Brangin, Ulu Sg.

Subis, Niah N.P. Yii Puan Ching S 40171.

*Mitreola sphaerocarpa (Leenh.) Leenh.

Creeping herb. In shade at base of limestone cliff, 167 m alt. On southern slopes

of G. Subis in the Sekaloh river, Niah J.A.R. Anderson, Sonny Tan & E. Wright S

260906.

Motleyia borneensis J.T. Johansson

G. Subis area, Niah Mohidin S 21665.

Mussaendopsis beccariana Baill.

Tree 30 m tall, 100 cm girth. Primary lowland forest. Niah J.A.R. Anderson &

Chew Wee Lek S 16020.

*Mycetia javanica (Blume) Korth.

Fleshy herb or slender shrub to 2 m tall. On limestone at 17 m a.s.l., also in

secondary alluvial forest with heavy silty clay soil near riverbank also on broad

sloping ravine between two vertical limestone hills with numerous limestone

boulders and some residual soil, c. 167 m a.s.l.. G. Subis area, Niah Mohidin S$

21605; Niah Caves Chew Wee-Lek CWL 316; On southern slopes of G. Subis, in

the Sekaloh river. Niah J.A.R. Anderson, Sonny Tan & E. Wright S 26091; Niah

N.P. Paul Chai S 40052, G. Brangin, Ulu Sg. Subis Yii Puan Ching S 40170.

Myrmeconauclea strigosa (Korth.) Merr.

Shrub 1.3 m tall. Primary lowland dipterocarp forest on rocks beside swift running

stream. Ulu Sg. Sekaloh, Niah river Benang ak Bubong S 27868.

Nauclea parva (Havil) Merr.

Shrub about 3 m tall. Riverine forest. Along Sg. Niah, Niah N.P. Yii Puan Ching S

40116.

176

Gard. Bull. Singapore 57 (2005)

*Ophiorrhiza fibrillosa Ridl.

Herb c. 67 cm tall. On broad sloping ravine between two vertical limestone hills

with numerous limestone boulders and some residual soil. Niah Caves area B.L.

Burtt and P.J.B. Woods B 2029; Southern slopes of G. Subis in the Sekaloh river,

Niah J.A.R. Anderson, Sonny Tan & E. Wright S 26095.

Petunga coniocarpa Korth.

Tree 5 m tall, 10 cm girth. Primary lowland dipterocarp forest on clay loam soil.

Ulu Sg. Sekaloh, Niah river Sibat ak Luang S 26126.

Pleiocarpidia enneandra (Wight) K. Schum. & Bremek.

Tree 15 m tall, 48 cm girth. Primary lowland dipterocarp forest on clay loam soil.

Ulu Sg. Sekaloh, Niah river Banyeng ak Nudong S 26143.

Pleiocarpidia opaca Bremek.

Tree 6 m tall, 8 cm girth. Primary MDF on clay loam soil, sedimentary rock, 100 m

a.s.l. Ulu Sg. Sekaloh, Niah river Banyeng ak Nudong S 26138, E. Wright S 27253.

Pleiocarpidia sandahanica Bremek.

Tree 21 m tall, 75 cm girth. Primary lowland dipterocarp forest on clay loam soil.

Ulu Sg. Sekaloh, Niah river Benyang ak Bubong S 27867.

Porterandia anisophylla (Jack ex Roxb.) Rid.

Tree 13 m tall, 43 cm girth. MDF on clay loam soil, 100 m a.s.1. Ulu Sg. Sekaloh,

Niah river J.A.R. Anderson, Sonny Tan & E. Wright S 26051.

Prismatomeris beccariana (Baillon) J.T. Johansson.

Treelet 2 m tall. Primary lowland dipterocarp forest on clay loam soil. Ulu Sg.

Sekaloh, Niah river Erwin Wright S 29119.

*Psychotria elmeri Merr.

Tree 3 m tall, 10 cm girth. Lower crest of limestone hill, limestone pinnacles on

‘mor’ soil, 167 m a.s.l. Southern slopes of G. Subis near Sg. Sekaloh Niah Mohidin

S 21617, Sonny Tan & E. Wright S 27593.

*Psychotria cf. expansa Blume

Limestone. Niah Haviland & C. Hose, Haviland 3462A.

Psychotria laxiflora Blume

Climber. Primary lowland dipterocarp forest on clay loam soil. Ulu Sg. Sekaloh,

Niah river Benyang ak Bubong S 27882.

*Psychotria pachyphylla Ridl.

Epiphyte at 1.3 m near base of small tree. Summit ridge of limestone, 267 m a.s.L.,

limestone strongly dissected and covered by 15-23 cm ‘mor’ layer. Sg. Sekaloh, G.

Subis, Niah J.A.R. Anderson S 31932.

*Psychotria sarmentosa Blume

Creeping herb. Crest of limestone hill, limestone pinnacles and ‘mor’ soil. Southern

slopes of G. Subis near Sekaloh river, Niah Sonny Tan & E. Wright S 27281.

Psychotria woodii Merr.

Paraboea culminicola and plants from Niah, Borneo v7

Small tree. In forest on ground. Kg. Tangap, Niah Ahmad 30.

*Timonius matangensis Valeton

Tree 10 m tall, 23 cm girth. Limestone hill, lower summit ridge, limestone pinnacles

and ‘mor’ soil. South side of G. Subis in Sekaloh river, Niah J.A.R. Anderson,

Sonny Tan & E. Wright S 27561.

Uncaria cordata (Lour.) Merr. f. sundiaca C.E. Ridsdale

Climber entwined around tree. Primary lowland dipterocarp forest on clay loam

soil. Ulu Sg. Sekaloh, Niah river Sibat ak Luang S 26132.

Uncaria ferrea DC.

G. Subis area, Niah Mohidin S 21633.

Uncaria insignis DC.

Climber creeping on trees. Riverine forest c. 17 m a.s.l., soil loamy. Kuala Sg.

Sekaloh, Niah N.P. Yii Puan Ching S$ 40110.

Xanthophytum glabrum Axelius

G. Subis area, Niah Mohidin S 21651.

RUTACEAE

Clausena excavata Burm.f.

Tree 4 m tall, 35 cm girth. Flowers in bud light green, fruits green when young,

whitish when mature. Riverain forest. Along Niah River, Niah N.P. Yii Puan

Ching S 40143.

Glycosmis chlorosperma (Blume) Spreng. var. elmeri (Merr.) Tanaka

Treelet 3 m tall. Primary lowland dipterocarp forest on clay loam soil. Ulu Sg.

Sekaloh, Niah river Sibat ak Luang S 27856.

Glycosmis superba. B.C. Stone

Tree to 7 m tall, 12.5 cm girth. MDF on clay loam soil at 100 m. Ulu Sg. Sekaloh,

Niah river J.A.R. Anderson, Sonny Tan & E. Wright S 26059 (type specimen),

Benang ak Bubong S 26118.

*Tunasia amara Blanco

Slender shrub or treelet 2—3.3 m tall. On ‘mor’ soil, summit ridge of limestone

130—267 m a.s.l., limestone strongly dissected and covered by 15-23 cm ‘mor’

layer. Sg. Sekaloh, G. Subis, Niah J.A.R. Anderson S 31925; Niah N.P. Paul Chai S

40061.

SABIACEAE

Meliosma sumatrana (Jack) Walp.

G. Subis area, Niah Mohidin S 21674.

SAPINDACEAE

Dimocarpus longan Lour. ssp. malesianus Leenh.

Tree 6 m tall. Primary lowland dipterocarp forest on clay loam soil. Ulu Sg.

Sekaloh, Niah river Sibat ak Luang S 26148.

Guoia bijuga (Hiern) Radlk.

178 Gard. Bull. Singapore 57 (2005)

Tree 10 m tall, 40 cm girth. Riverine forest. Along Sg. Niah, Niah N.P. Yii Puan

Ching S 40153.

*Harpullia cupanioides Roxb.

Tree 13 m tall, 58 cm girth. Broad sloping ravine between two vertical limestone

hills with numerous limestone boulders and some residual soil. On southern slopes

of G. Subis, in the Sekaloh river. Niah J.A.R. Anderson, Sonny Tan & E. Wright S

26093.

Lepisanthes alata (Blume) Leenh.

Slender tree. On riverbank, riverine forest. Along Niah river, Niah N.P. Yii Puan

Ching S 40142.

*Lepisanthes fruticosa (Roxb.) Leenh.

Tree to 17 m tall and 28 cm girth. Primary MDF on clay loam soil at 100 m a.s.],

also on limestone rocks near river. Ulu Sg. Sekaloh, Niah river J.A.R. Anderson,

Sonny Tan & E. Wright S 26057, 26079, Benang ak Bubong S 26113, Sibat ak

Luang S 27854.

SAPOTACEAE

*Tsonandra lanceolata Wight var. lanceolata

Tree to 27 m tall and 133 cm girth. Crest of limestone hill, limestone pinnacles and

‘mor’ soil and narrow limestone ridge at 130 m a.s.l.. Southern slopes of G. Subis.

Near Sekaloh river Sonny Tan & E. Wright S 27278; Niah N.P. Paul Chai S 40060.

Madhuca ct. spectabilis P. Royen

Tree 14 m tall, 45 cm girth. MDF on clay loam soil at 100 m a.s.1. Ulu Sg. Sekaloh,

Niah River J.A.R. Anderson, Sonny Tan & E. Wright S 26058.

*Pouteria obovata (R.Br.) Baehni

Tree 10 m tall, 38 cm girth. Summit ridge of limestone 267 m az.s.l., limestone

strongly dissected and covered by 15-23 cm ‘mor’ layer. Sg. Sekaloh, G. Subis,

Niah J.A.R. Anderson S 16049, S 31928.

SAURAUIACEAE

*Saurauia acuminata Merr.

7 m tall with many arching stems. From limestone bank and also base of limestone

hill on limestone rocks. Niah Caves B.L. Burtt and P.J.B. Woods B 2038; Path to

Great Cave J.A.R. Anderson S 31692.

Saurauia cf. ferox Korth.

Treelet 2.7 m tall. Primary lowland dipterocarp forest on clay loam soil. Ulu Sg.

Sekaloh, Niah river Benang ak Bubong S 27863.

Saurauia reinwardtiana Blume

G. Subis area, Niah Mohidin S 21680.

SCROPHULARIACEAE

*Lindernia serrata (Roxb.) O. Kuntze

Limestone. Niah Haviland & C. Hose, Haviland 3511A.

Torenia polygonoides Benth.

Paraboea culminicola and plants from Niah, Borneo 179

Niah Haviland & C. Hose, Haviland 3508A.

Vandellia sp.

Niah Haviland & C. Hose, Haviland 3538A.

SCHIZAEACEAE

Lygodium borneense Alderw.

G. Subis area, Niah Mohidin S 21661.

Lygodium circinnatum (Burm.f.) Sw.

In Subis forest on ground, Niah Ahmad 87.

Schizaea dichotoma (L.) Sm.

In forest on ground. Rumah Pasang, Niah Ahmad 42.

SELAGINELLACEAE

*Selaginella alutacia Spring

Hanging on limestone rockface. Niah Lord Medway 8950.

*Selaginella hewittii Hieron.

On damp limestone rockface. Niah Mohiden 8960.

Selaginella paxii Hieron.

G. Subis area, Niah Mohidin S 21615.

Selaginella phanerotricha Baker agg.

G. Subis area, Niah Mohidin S 21693.

Selaginella ct. roxburghii (Hook. & Grev.) Spring

G. Subis area, Niah Mohidin S 21699.

SOLANACEAE

Nicotiana tabacum L.

Cultivated, Niah Haviland & C. Hose Haviland 3536.

STERCULIACEAE

*Sterculia coccinea Roxb.

Tree 3.3 m tall. Base of limestone hill on limestone rocks. Path to Great Cave, G.

Subis J.A.R. Anderson S 31696.

*Sterculia macrophylla Vent.

Tree 30 m tall, 120 cm girth. On limestone rocks amidst alluvium by stream at

base of limestone hill also on MDF with reddish clay loam soil. South side of G.

Subis in Sekaloh river, Niah J.A.R. Anderson, Sonny Tan & E. Wright S 27578;

Ulu Sg. Sekaloh, Niah Erwin Wright S 29263.

*Sterculia megistophylla Ridl.

Small tree. On limestone rocks or scree c. 100 m a.s.l.. Vicinity of Niah Caves B.C.

Stone 13729.

THEACEAE

Adinandra dumosa Jack

180 Gard. Bull. Singapore 57 (2005)

Tree 40 m tall, 300 cm girth. Lowland MDF 80 m a.s.l, soil sandy loam. Sg.

Sekaloh, Niah N.P. Yui Puan Ching S 40137.

THELYPTERIDACEAE

Christella arida (D. Don) Holttum

1.3 m long. In Subis forest on ground. Niah Ahmad 88.

Christella subpubescens (Blume) Holttum

G. Subis area, Niah Mohidin S 21621.

Cyclosorus sp.

G. Subis area, Niah Mohidin S 21684.

*Pronephrium simillimum (C. Chr.) Holttum

Terrestrial fern. In shaded or semi-shaded localities under large overhang 100 m

up on steep limestone hill, light dry soil, partly composed of guano. Gua Pangomah,

G. Subis, Niah J.A.R. Anderson S 31690.

Pronephrium sp.

On rock near entrance Niah Cave. Niah Ahmad 39.

THYMELAEACEAE :

Gonystylus calophyllus Gilg

Tree 5 m tall. Primary forest at base of sandstone hill, clay loam soil. Between

mouth of Sg. Sekaloh and G. Subis, Niah J.A.R. Anderson 27583.

*Wikstroemia indica (L.) C.A. Mey.

Shrub 1.7 m high. Crest of limestone hill, limestone pinnacles and ‘mor’ soil.

Southern slopes of G. Subis, near Sekaloh river Sonny Tan & E. Wright S 27277.

TILIACEAE

Brownlowia sp.

Tree 27 m tall. In poorly drained area with dense primary forest. Niah N.P. Rogstad

729,

TRIURIDACEAE

Sciaphila sp.

G. Subis area, Niah Mohidin S 21638. (Mixed collection)

URTICACEAE

*Flatostema palioneurum

Limestone. Niah Haviland & C. Hose Haviland 3318H

*Elatostema variolaminosum H. Schr. var. latum Schrooter

Herb. On limestone ridge at 130 m a.s.l. G. Branchin, Ulu Sg. Subis, Niah N.P. Yii

Puan Ching S 40165.

*Pipturus argenteus (G. Forst.) Wedd.

Shrub c. 5 m tall. On limestone c. 167 m, also at the foot of limestone hill near

stream at 20 m a.s.l. Niah Caves Chew Wee-Lek CWL 312: Ulu Sg. Subis, Niah

N.P. Bernard Lee S 40081.

Paraboea culminicola and plants from Niah, Borneo 181

Pouzolzia zeylanica (L.) Benn.

Niah Haviland & C. Hose Haviland 3319.

VERBENACEAE

Clerodendrum fistulosum Becc.

1.3 m tall. In forest on ground. Niah Ahmad 51.

*Clerodendrum laevifolium Blume

Herb or shrub 30—67 cm tall. Limestone forest at 80 m a.s.l. and on limestone in

shelter of cave mouth, damp and partially shaded. Northern entrance to Great

Cave and G. Subis area, Niah G.D. Haviland & C. Hose Haviland 3556, Ahmad

79, Mohidin S 21602, J.A.R. Anderson, Sonny Tan & E. Wright S 26071, Yii Puan

Ching S 40197.

Stachytarpheta dichotoma (L.) Vahl

1-1.3 m tall. In jungle on wet ground. Kuala Tangap, Niah Ahmad 20.

VITACEAE

Ampelocissus capillaris Merr.

G. Subis Area. Niah Mohidin S 21668.

*Tetrastigma diepenhorstii (Miq.) Latiff

Climber to 13 m tall. Primary lowland dipterocarp forest on clay soil and limestone

hill, lower summit of ridge, limestone pinnacles and ‘mor’ soil. South side of G.

Subis in Sekaloh river, Niah J.A.R. Anderson, Sonny Tan & E. Wright S 27565;

Ulu Sg. Sekaloh, Niah river Erwin Wright S 29139.

VITTARIACEAE

*Antrophyum callifolium Blume

Small terrestrial fern. In Subis forest on ground, also on limestone rocks, slope

between caves, limestone rocks and residual soil, and as an epiphytic at base of

trees, on trees mainly felled, habitat semi-exposed. Great Cave, G. Subis, Niah

J.A.R. Anderson 31920; Niah Ahmad 86.

Vittaria elongata Sw.

G. Subis area, Niah Mohidin S 21677.

WOODSIACEAE

*Diplazium crinitum (Baker) C.Chr.

Frond 1.3 m long. Below Niah cave on rock. Niah C. Hose 1230, Ahmad 75, G.

Subis area, Niah Mohidin S 21607.

ZINGIBERACEAE

Alpinia sp.

2.7 m tall. Flower buds bright red. Lowland dipterocarp forest, alluvial soil beside

stream. Ulu Sg. Sekaloh, Niah river Sibat ak Luang S 27875.

*Burbidgea stenantha Ridl.

Herb, stem bronze. On vertical limestone, rooting in crevices, gully between

limestone hill, limestone rocks and pavement, heavily dissected, mineral soil very

sparse also on limestone wooded rocks. G. Subis B.L. Burtt and P.J.B. Woods B

2007, J.A.R. Anderson 31669.

182 Gard. Bull. Singapore 57 (2005)

Globba affinis Rendle

Herb 25 cm tall. Clayey soil not far from stream. Ulu Sg. Sekaloh, Niah Erwin

Wright S$ 29270.

Plagiostachys strobilifera (Bak.) Ridl.

3 m tall. Flower buds bright red, fruits brown. Lowland dipterocarp forest, alluvial

soil beside stream. Ulu Sungei Sekaloh, Niah river Sibat ak Luang S 27874.

Erratum from ‘The Vegetation and Plants of Niah National Park, Borneo’

DRYOPTERIDACEAE

Tectaria andersonii Holttum. J.A.R.Anderson S 31936 (not S 31963).

ORCHIDACEAE

Appendicula undulata Blume var. longicalcatara (Rolfe) Ames

Pearce et al. S 89489

Calanthe triplicata (Willemet) Ames

Jamree et al. § 89050, Pearce et al. S 89270.

Gardens’ Bulletin Singapore 57 (2005) 183—185 183

Aglaia soepadmoi Pannell (Meliaceae), a New

Species for Borneo

C.M. PANNELL

Daubeny Herbarium (FHO), Department of Plant Sciences,

University of Oxford, South Parks Road, Oxford OX1 3RB, U.K.

More species of Aglaia Lour. occur in Borneo than anywhere else in the

range of the genus. Sixty species are known from the island, of which 12

are endemic. The species described here is a rare plant recorded from

Lundu District in west Sarawak and from Sumatra. It belongs to section

Aglaia.

Aglaia soepadmoi Pannell, sp. nov.

(Engkik Soepadmo, Coordinator and Chief Editor of the Tree Flora of

Sabah and Sarawak project)

Aglaia soepadmoi ab A. eximia foliolis paucioribus latioribusque siccitate

nervatura magis conspicua nervis lateralibus prominentibus et

brochidodromis, reti subprominenti, venatione in pagina superiore tota

impressa differt. Typus: Othman et al. S 59970, Borneo, Sarawak, Lundu

District, Sungai Sebuloh, fl 15 August 1990 (holotypus SAR!; isotypus

KEP!).

Small tree to 7 m. Twigs longitudinally channelled, densely covered with

dark brown or blackish brown stellate hairs, with pale stellate scales

interspersed. Leaves and inflorescences crowded near the apices of the

shoots. Leaves usually imparipinnate, rarely lacking the terminal leaflet, to

95 cm long, 44 cm wide; petiole 22—28 cm long; the petiole, rachis and

petiolules channelled and with hairs and scales like those on the twigs.

Leaflets 9-11, shiny green above, densely covered with white stellate scales

interspersed with dark brown hairs beneath; blades ovate or elliptical, the

terminal one often markedly obovate, 15.5—24 x. 6—9 cm, base cordate,

apex acuminate, acumen acute to 16 mm long, sometime obovate and apex

rounded; midrib and lateral veins impressed and reticulation visible above;

midrib prominent, lateral veins 11—17 pairs, ascending and markedly curved

upwards at the margin, nearly or quite anastomosing, subprominent and

reticulation slightly prominent beneath; petiolules 0.5 cm long. Inflorescences

to 17 x 9 cm, densely covered with indumentum like that on the twigs.

Flowers sessile, subglobose, c. 1.3 x 1.3 mm; calyx divided into 5 lobes,

184 Gard. Bull. Singapore 57 (2005)

densely covered with reddish-brown stellate hairs and scales outside; petals

5; staminal tube obovoid, with a shallowly lobed aperture c. 0.2 mm across;

anthers 5, almost as long as the staminal tube, inserted longitudinally and

visible through the aperture; ovary subglobose, stigma ovoid with two apical

lobes. Infructescences c. 18.5 cm long. Fruits indehiscent, subglobose, c. 2 x

2 cm, densely covered with dark brown stellate hairs with a few white hairs

interspersed.

Distribution: Known by two collections from Sumatra and five collections

from Sarawak (including the type).

Specimens examined: SUMATRA. South Sumatra, Lampong Prov., Mt

Tanggamus, fr. 1 May 1968, Jacobs 8190 (KEP, L); West Sumatra, Pasaman,

Mt Talamau (Ophir), fl., 8 Nov. 1984, Laumonier YL 6692 (FHO).

SARAWAK. Lundu District, Sungai Sebuloh, young fl 15 Aug. 1990,

Othman et al. 59970 (KEP, SAR), young fr 27 Nov. 1991, Othman et al. S

63857 (SAR); Datu Permanent Forest, fl., 17 May 1980, Bernard Lee S$

41873 (FHO, KEP, SAR); Gunung Gading, fl., 7 June 1991, Yahud et al. S$

54873 (SAR); Gunung Gading, Sekamal, 12 Dec 1995, S.T. Lai et al. S

54426 (KEP, SAR).

Ecology: Lowland dipterocarp forest, disturbed forest and rocky terrain in

beach forest, up to 1100 m altitude.

Notes: Aglaia soepadmoi differs from A. eximia in having fewer, broader,

leaflets, with more conspicuous venation when dry. The lateral veins are

prominent and looped at the margins, the reticulation is subprominent. All

venation is impressed on the upper surface.

Acknowledgements

The author thanks the following: Dr Engkik Soepadmo, who confirmed to

me that this species is distinguishable from Aglaia eximia Migq., to the

Directors, Curators and staff of the Forest Research Institute of Malaysia

and the Forestry Departments of Sabah and Sarawak for inviting me to

work in their herbaria, to Dr Robert Mill of Edinburgh Botanic Garden

for latinising the diagnosis and to Joseph Pao (SAR) for preparing the

illustration.

Aglaia soepadmoi from Borneo 185

1mm

Figure 1. Aglaia soepadmoi Pannell. A, leafy twig with young male inflorescences; B, detail

of lower leaflet surface showing indumentum; C, stellate hair; D, stellate scale; E, female flower;

F longitudinal section of female flower; G, longitudinal section of young male flower; H, young

infructescence; I, young fruit. (A, B, C, D and G from S$ 4/873; E and F from S 59970; H from

S 54426;1 from M. Jacobs 8190).

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Gardens’ Bulletin Singapore 57 (2005) 187—19& 187

Curcuma roscoeana Wall. (Zingiberaceae) in

India

J. SKORNICKOVA !2 AND M. SABU!

‘Department of Botany, University of Calicut, 673635 Kerala, India

"Department of Botany, Charles University, Benatska 2,

Prague, 12802, Czech Republic

Abstract

Collections from the Andaman Islands, formerly determined to Curcuma kurzii King ex

Baker or C. petiolata Roxb., match C. roscoeana Wall. A detailed description from living

material, a colour plate, as well as historical and nomenclatural details are provided and

lectotypes are designated for C. roscoeana and C. kurzii.

General Introduction

The genus Curcuma L., with an expected total of c. 120 species, occurs

throughout tropical ani sub-tropical Asia with a few species extending to

Australia and the Pacific region (Skornickova et al., 2004). It is of great

economic and ornamental importance and at the same time is one of the

genera within the family Zingiberaceae with polymorphic species, which

has caused confusion with species delimitation.

The first author is revising Curcuma in India. To date some 30 species

have been reported (Skornickova et al., 1989; Jain and Prakash, 1995;

Velayudhan et al., 1996; Skornickova and Sabu, 2002), including several new

taxa and new records (Skornickov4 Balachandran, 1983; Bhat, 1987;

Mangaly and Sabu, 1988; Mangaly and Sabu, 1993; Tripathi, 2001;

Skornickova et al., 2003a, b; Skornickova et al., 2004).

Our studies Sar that specific delimitation Hint the genus, synonymy

and even the identities of some of the commonly cultivated taxa are quite

confusing and that names are often misapplied. The main reasons include

the following.

1. Many taxa described at the beginning of 19° century have

scanty protologues and type specimens, if they were cited,

are either lost or have deteriorated.

2. The characters used to delimit species in Curcuma are not

obvious on herbarium sheets. This applies particularly to shape

188 Gard. Bull. Singapore 57 (2005)

and colour of the rhizome, position of the inflorescence, colour

of the bracts, and floral characters such as the shape of the

anther spurs, the colour and shape of floral parts.

3. Notes on herbarium labels made by people not familiar with

the floral structure of this genus can be quite misleading.

4. Many species are superficially very similar and, without a

detailed description based on living material, are difficult to

determine.

5. The huge area of the Indian subcontinent and the rather high

variability among seed-setting species result in some species

being described repeatedly under different names.

These points demonstrate clearly that fieldwork and observation of

flowering material, especially from the type locality or nearby, together

with a search for original historical material are key to an accurate

understanding of the genus Curcuma. During the course of our work we

intend to address and clarify individual problems connected with the

taxonomy and nomenclature of Indian representatives of the genus.

What is Curcuma kurzii King ex Baker?

Curcuma kurzii was described by Baker (1890) in The Flora of British

India, under the heading ‘imperfectly known species’. The description is

based on King’s manuscript and consist of two lines: ‘nearly allied to C.

petiolata, but leaves larger, petioles longer, scape longer, bracts more

imbricating, and their tips less spreading. — S. Andaman Islds’. From this

we presume that Baker was dealing with herbarium specimens and that he

had had no opportunity of seeing live material of C. kurzii. It is not

surprising that he found this plant closely allied to C. petiolata because in

general habit these two species are indeed very similar and the fertile

bracts of both species form rather deep pouches. Colours and delicate

flower parts are very rarely preserved in herbarium material and, as we

have confirmed, they are not present on the presumably original material

collected by S. Kurz from South Andaman Island at CAL and K.

Balakrishnan and Bhargava (1984), Srivastava (1998) and Tripathi and

Prakash (1998) treated C. kurzii as conspecific with C. petiolata Roxb.

During our fieldwork in the Andaman Islands, April-June 2002,

we collected sterile specimens of Curcuma, assuming that it would be C.

petiolata based on the earlier work of Balakrishnan and Bhargava (1984).

Rhizomes were successfully transplanted to Calicut University Botanical

Garden and, in late November 2002, one of them flowered, followed by

Curcuma roscoeana in India 189

others in the autumn of 2003 and in 2004. Though we found them identical

to the description given by Balakrishnan and Bhargava (1984), they neither

matched the protologue, nor the descriptions and colour drawings of C.

petiolata given by Roxburgh (1820) and others (Roscoe, 1827; Hooker,

1870).

In contrast to the original description of Curcuma petiolata, our

plants had no distinct coma, the whole inflorescence was bright orange-

red, the bracts gradually becoming yellow-green towards the base, and the

flowers were longer than the bracts and slightly exserted. Most strikingly,

the bracts were arranged in rows. These characters led us to determine our

material as the Burmese species C. roscoeana Wall. This was confirmed by

consulting researchers recently working on the genus Curcuma and

comparing our plant with photographs of C. roscoeana from Thailand and

Burma, where this species is found in wild and is widely cultivated.

As other seed-setting taxa, it shows a considerable degree of

variability leading to the opinion that there are no grounds for keeping C.

kurzii as a separate species or even as a distinct variety. Thus, C. kurzii is

treated here as conspecific with C. roscoeana Wall. and as such is an addition

to the Indian flora. A detailed description is given to include characters

observed from living plants (Skornickova and Prasanthkumar 73309, 73310),

which flowered at CUBG during September-November 2002, 2003 and

2004 (Plate 1.).

Curcuma roscoeana Wall. Pl. Asiat. Rar. I. (1829) 8, t. 9 — Hitchenia

roscoeana (Wall.) Benth. in Benth.& Hook.f. Gen. Pl. 3 (1883) 643. —

Hicheniopsis roscoeana (Wall.) Loes. Nat. Pflanzenfam. ed. 2, 15(A) (1930)

572. — Type: Wallich. Pl. Asiat. Rar. t. 9. (lectotype; designated here).

Curcuma coccinea Wall. ex Baker, in Hook. f.,, Fl. Brit. Ind. 6 (1890) 216,

nom. nudum (in syn.) C. kurzii King ex Baker, in Hook. f., Fl. Brit. Ind. 6

(1890) 216, syn. nov. —Type: Andaman Islands, South Andaman, Smith

Point, S. Kurz s.n. (K!, lectotype, designated here; CAL! [Acc. No. 467218],

isolectotype); Andaman Islands, South Andaman, without exact locality,

S. Kurz s.n. (K!, G!; putative isolectotypes).

C. petiolata auct. non. Roxb. Balakrishnan & Bhargava. J. Bombay Nat.

Hist. Soc. 81 (1984) 512; Srivastava S.K. Indian J. Forest. Add. Series X.

(1988)16; Tripathi & Prakash. J. Econ. Taxon. Bot. 22 (1998) 468.

Rhizomatous herb, to 90 cm tall. Rhizome ovoid without branches (rarely

with one branch, which later turns into another main rhizome), c. 3-4 x 1—

2 cm (increases with age), light brown externally, sheathed by bases of the

leaf sheaths, which leave vertical scars after decaying, creamy-yellowish

190 Gard. Bull. Singapore 57 (2005)

inside, not aromatic. Roots many, penetrating deeply into soil terminating

with small ovate tubers 0.5—2 x 0.5—1.5 cm, 2—15 cm from the main rhizome,

externally light brownish, glabrous, white internally. Leafy shoot 30—90

cm, pseudostem 15—25 cm long formed by leaf sheaths and a few sheathing

bracts (conspicuous especially at the beginning of the season, later drying

and decaying), softly pubescent, hairs 0.2 mm long, green with deep red-

violet tinge, /igule obscurely bilobed, 1.5—2.5 mm long, greenish translucent,

hairy on the margin, hairs 0.5 mm. Leaves 3—4 at the beginning of the

season, later to 8, petiole 7—25 cm long, shorter in the first leaves, gradually

longer in older leaves, deeply channelled, green or with reddish-violet

tinge, glabrous; /Jamina oblong-ovate, c. 16-35 x 5.5—13 cm (first leaves

smallest), glabrous on both sides, adaxially green, with prominent veins c.

0.7—1 cm apart, abaxially pale green; margin hyaline, white, 0.2 mm broad,

tip c. 1.5 cm long, acuminate, base rounded to cordate, slightly oblique,

midrib green, glabrous. Inflorescence always central. Peduncle 15—28 cm,

green, glabrous, most of it hidden within the pseudostem. Spike c. 8-15 x

4—7 cm, cylindrical, consisting of 15—45 bracts arranged in 3—5 serial rows,

base of spike attenuate formed by deep pouches of lowermost fertile bracts.

Size of the spike and number of bracts and rows increase with the age of

plants. Coma bracts similar in colour and size to fertile bracts, sterile c. S—

7, spirally arranged. Fertile bracts obovate-spatulate, c. 3.5—4.5 x 2.5—3 cm,

deep orange-red in upper part gradually becoming yellow—greenish at their

base (character which can be often seen in herbarium specimens as highly

glossy yellowish patch at the base of bracts, provided plants were processed

by the dry method) lower half forming deep funnel-shaped pouches

subtending cincinnus of 2—4 flowers, upper part spreading, glabrous or

shortly hairy, usually more hairy outside, margin shortly hairy, hairs 0.2—

0.3 mm long. Bracteoles one per flower, 5 x 3 mm, hyaline, creamy-white,

glabrous. Flowers 4.5—5.5 cm long, cream-white with yellow in the centre

of lip, slightly exserted from the bracts. Calyx 14-16 mm long, translucent

white, glabrous, obscurely 3-toothed, unilaterally split 4-5 mm. Corolla

tube 3—3.5 cm, creamy white, glabrous, dorsal corolla lobe 0.9-1.1 x 0.5—

0.7 cm, obtuse, creamy white, glabrous, lateral lobes 0.9—1 x 0.4—0.6 cm,

creamy white, glabrous. Labellum 1.4—1.6 x 1.7—2 cm, slightly emarginate,

split c. 1 mm, creamy white with deep yellow patch in the centre, yellow-

coloured area raised and swollen, forming a channel in the middle of

labellum (especially at the basal part of the labellum). Lateral staminodes

unequally rhomboid, 12—15 x 8—9 mm, creamy white, glabrous. Anther c. 5

Plate 1. Curcuma roscoeana Wall. from the Andaman Islands 1. Flower in bract (side view); 2.

Anther (lateral view); 3. Flower (front view); 4. Rhizome; 5. Inflorescence (young plant, c. 2—3

years old); 6. Flower (dissected). (Skornickova & Prasanthkumar 73309). Photo J. Skornickova.

—

ie)

is)

1S)

192 Gard. Bull. Singapore 57 (2005)

mm, yellowish white with yellow at apical part, anther thecae 3-4 mm

long, whitish, dehiscing by basal pore, filament 2—3 mm long, 3.5 mm

broad at base, 2.5 mm broad at upper part. Anther spurs absent. Anther

crest 1.5 mm long, hyaline, yellow, rounded, terminally recurved. Ovary

trilocular, 2—2.5 x 1.5—2 mm, white, glabrous, ovules many. Stigma 1.4 x 0.7

mm, translucent white, ciliate, cilia 0.3 mm, surrounded by anther crest,

not exserted. Epigynous glands 2, creamy-white, 4-5 mm long, 0.5 mm

diameter. Fruits not seen.

Specimens examined: INDIA, North Andamans: Sitanagar forest, 15.V.1982,

M.K. Vasudheva Rao 9032 (CAL, PBL); Laksmipur, 23.X1.1976, N.G. Nair

4881 (PBL, L-digital image seen ); Mayabunder, 6.XII.1992, B.K. Sinha

16263 (PBL); Mayabunder Dt., Chainpur, 19.V.2002, Skornickova &

Prasanthkumar 73310 (CALI, SING); Mayabunder Dt., Chainpur,

19.V.2002, Skornickova & Prasanthkumar 73309 (CALD: Middle

Andamans: Bakultala, 6.X1.1979, N. Bhargava 6406 (CAL, PBL, “i;

Mayabunder, 31.VII.1974, N. Bhargava 1941 (CAL, PBL); South Andaman:

Coatering Cave, S.’Kurz s.n., Acc. No. 467217 (CAL); Baboo ghat,

7.VIIL1894, King’s coll. s.n., 3 nos (CAL); Saroah Creek, Baratang,

25.X%.1979) DP: Basa 7551 (PBL): Andamans, sine loc: 22.VIII.1898, G.H.

Mann s.n. Acc. No. 467212 (CAL); sine loc., HBC s.n. Acc. No. 467207

(CAL); Bom. Lung. Jang., X.1915, Parkinson 678 (DD); Jharkhand:

Singbhum, IX. 1900, H.H. Haines 331 (K, CAL); Parasnath, 30. IX. 1873,

C.B.Clarke 20181 (CAL); Chota Nagpur, 4. IX. 1875, J.J.Wood s.n. (K);

Western Bengal & Behar: S. Kurz s.n., Acc. No. 467109 (CAL);

BANGLADESH: Sine loc., Griffith 5724 (K); BURMA (MYANMAR),

Repu Irawaddi ad Scandwya, Wallich 6597B (CAL); Upper Burma, 1897,

Veitch & sons s.n. (K); Rangoon District: Rangoon, sine dat., M. Sellan s.n.

(E); Hanthawaddy, 25.VIII.1932, Parkinson 14831 (DD); Prome Road,

24.VIII.1932, Parkinson 14831 (DD); Kamaynt, 24.VIII.1932, C.E.

Parkinson 14629 (DD); Pegu, McLeland s.n., 2 nos (K); Pegu, sine dat.,

sine coll., 2 nos (CAL); Minbu District: Dwe Chaung, 23.XI.1937, C.E.

Parkinson 79677 (DD); Tenasserim, Tavoy Dt., Paungdaw, IX. 1961, Keenan

et al. 1502 (E, K); South Tenasserim, Leikpok Chaung Mergui, 1925, Mr.

Braybon’s collector 223 (DD); Tenasserim and Andamans, Herb. Helfer

5711 (CAL); Toungoo Dt., Dongzayit, 4.VII.1911, J.H. Lace 5391 (CAL, E,

K); sine loc. 1880 D. Brandis s.n. (DD); THAILAND: Chiang Mai, 1965,

Johnston s.n. (BM); Chiang Mai, Queen Sirikit Bot. Garden, 10. VII. 1999,

C. Ngamriabsakul 41 (E); Hin Dat, Kanburi, 2. VII. 1926, Put 73 (BM, K);

Muang Ngao, Lampang, 15. VII. 1931, Put 3996 (K); Maehongson, Khon

Yuam, 5. IX. 1974, Larsen & Larsen 34161 (K); Wangka, 2. — 4. VII. 1946,

Kostermans 799 (K); sine loc.: VI. 1854, No. 26 (CAL), sine dat., sine coll.

Curcuma roscoeana in India 193

(BM); VIII. 1862, sine coll. (BM); 184547, Galathea Expedition, Herb.

Wallich 841 (C, KIEL).

Distribution: So far only known to occur wild in Myanmar, Thailand and

India, but widely cultivated elsewhere for its ornamental value and bred as

a cut flower and pot plant. The only confirmed field observations of

Curcuma roscoeana in India are from the Andaman Islands. It is interesting

to note, however, that we have found in CAL and K four herbarium

specimens of C. roscoeana collected from forests in Jharkhand and West

Bengal/Behar by different collectors between 1875 and 1900, which bear

the remark ‘bracts orange red’ or ‘spike uniform dark yellow orange’ and

the shape and arrangement of the bracts leaves no doubt as to their identity.

Since then, the species has not been recollected from these areas, even

though we made an effort to search near to these localities. Another

interesting find at K is a specimen of C. roscoeana (Griffith 5724) collected

in East Bengal (Bangladesh) as part of the Herbarium of the East India

Company and distributed by RBG Kew during the years 1863-64. As far as

we can ascertain, this is the only evidence of C. roscoeana from Bangladesh,

but it is not clear if it was collected from wild.

Lectotypification

Wallich described Curcuma roscoeana in Plantae Asiaticae Rariores (1829)

but did not designate type specimens nor cite any herbarium material. The

description of C. roscoeana is accompanied by a plate (No. 9), an excellent

colour icon, where the whole plant as well as the dissected flower are

depicted. The type locality is given as “Pegu et ad oram Tenasserim’.

Wallich’s specimens can nowadays be found in about 46 herbaria,

the main set being deposited at K-W (Stafleu and Cowan, 1988). His

‘Catalogue’ or more correctly ‘Numerical List’ (1829-1832 + supplements

1847-1849) lists all Curcuma species between numbers 6594-6613. They

were thus published in 1832 (Anon., 1913). There are two entries for C.

roscoeana, 6597A collected in Rangoon and 6597B marked as Repu

Irawaddi ad Scandwya (Burma), both collected in 1826 as per entry in the

Numerical List. Forman (1997), writing on typification of Roxburgh’s names,

explained that, for the purpose of interpreting the name of the species,

there are many instances where the illustration is far superior to the

corresponding specimen and therefore the drawing would be much

preferable as the type. While studying Wallich’s collections in various

herbaria, we have encountered the problem that some of Wallich’s

duplicates bearing the same number do not always belong to just one

194 Gard. Bull. Singapore 57 (2005)

taxon. Moreover, some of his sheets comprise mixed collections, which is,

for example, obvious in the case of the C. roscoeana specimen 6597B

deposited in CAL, where only one of the two plants on this sheet belongs

to this species, the other being an undeterminable Curcuma species with a

lateral inflorescence. Considering this confusion with specimens, we

therefore propose to designate Plate 9 published together with the original

description in Plantae Asiaticae Rariores as the lectotype of C. roscoeana.

After critical analysis of Wallich’s original description and drawing

of Curcuma roscoeana (Wallich, 1829), we cannot but admit that it fits our

Andaman plant including important floral details such as the shape of the

anther without basal spurs, but with a well-developed anther crest, which

is precisely depicted in Wallich’s drawing. Other drawings and descriptions

of C. roscoeana were published in Botanical Magazine No. 4667 (Hooker,

1852), in 1854 by Lemaire in Le Jardin fleuriste and a short description

without a drawing was published by Dammer (1890). It is interesting to

note that the type locality of C. roscoeana (Pegu, on the coast of Tenasserim)

is just opposite the Andaman Islands. The possibility of plants being

introduced into the Andamans cannot be ruled out.

Apparently all specimens of this taxon from the Andamans collected

by Bhargava and other workers between 1979-1992 and deposited in PBL

(all determined as C. petiolata) are identical with several sheets from the

Andamans deposited at CAL and K (determined as C. kurzii or C.

roscoeana). They also match all other sheets of C. roscoeana deposited at

BM, C, CAL, DD, E, K, KIEL and L. Some of the sheets designated as C.

kurzii were collected by S. Kurz from the Andaman Islands and presumably

represent the original material, which was accessible to King and subsequent

workers. One sheet at K bears the remark “C. kurzii sp. nov. King ms.”

Thus, we propose to designate this sheet as lectotype and the other Kurz

collections as isolectotypes of C. kurzii.

Other historical sheets of this taxon from the Andamans worth

mentioning are Dr. King’s collections in 1894 designated as C. kurzii (CAL)

and two other sheets collected by Mr. Mann and an anonymous collector

annotated as C. roscoeana (CAL).

Recently, C. petiolata was reported from Arunachal Pradesh in NE

India by Tripathi and Prakash (1998). Their description and illustration do

not match well with either C. petiolata or C. roscoeana, neither does the

specimen cited by them (Tripathi 21838, CDRI!). Thus, the presence of C.

petiolata in India has yet to be confirmed. The differences between C.

roscoeana and C. petiolata are shown in Table 1.

Acknowledgements

The authors thank the Department of Science and Technology, Government

Curcuma roscoeana in India

Table 1. Comparison of important morphological characters of Curcuma

roscoeana Wall., C. petiolata Roxb. and C. sp. ’petiolata’ sensu Tripathi and

Prakash. Main diagnostic characters are in bold.

C. roscoeana Wall.

C. petiolata Roxb.

C. sp. ‘petiolata’ sensu

Tripathi and Prakash

Main rhizome

Ovoid unbranched,

(rarely with one

branch), inwardly

creamy-yellowish.

Branched, branches

few and small,

inwardly pale yellow.

Branched, branches

few, inwardly pale

yellow in centre,

greyish towards

margins.

Inflorescence

Fertile bracts

orange-red, with

yellow at base,

arranged in

conspicuous 3-5

rows. Coma

inconspicuous,

uppermost sterile

bracts orange-red.

Fertile bracts light

green, bracts spirally

arranged. Coma

conspicuous, lilac

coloured.

Fertile bracts

yellowish, orange at

the tip, arrangement

not mentioned. Coma

conspicuous, bracts

pinkish-orange or

pinkish-purplish.

Flower

Longer than the

bracts, slightly

exserted from the

bracts.

Small, not exserted

from the bracts.

As long as or shorter

than the bracts.

Anther

Spurs absent.

Anther crest

present, prominent.

Spurs present.

Anther crest present,

small.

Spurs absent. Anther

crest short (as per

description, but missing

in illustration).

of India (Order No. SP/SO/A-20/99 dt. 09.11.2001), to Grant Agency of

the Academy of Sciences of the Czech Republic (Grant No. B.6407401)

and International Association of Plant Taxonomy for financial support.

We are indebted to the staff of the Forest Department of the Andaman

Islands for their help, to the authorities of Ministry of Environment and

Forests for granting permission for collecting specimens, and to the curators

or BMC; CAL, CALY DD, E, G, K, KIEL, L,. PBL.and SING for letting

us examine specimens in their care or providing us with digital images/

photocopies of herbarium sheets, in particular to the library at K for giving

access to Icones Roxburghianae. Prof. P. Sirirugsa (Prince of Songkla

University, Thailand), Miss T. Rehse (Duke University, U.S.A), Mr. P.

Suksathan (Aarhus University, Denmark) and Mr. Ch. Maknoi (Prince of

Songkla University, Thailand) kindly sent us their photographs of wild and

cultivated Curcuma roscoeana. The first author thanks the Singapore

196 Gard. Bull. Singapore 57 (2005)

Botanic Gardens for awarding a Singapore Botanic Gardens Research

Fellowship and facilities during her repeated visits and to Dr. M. F. Newman

(E), to Dr. A. D. Poulsen (E) and to Dr. R. Kiew (SING) for comments on

the manuscript.

References

Anon., 1913. XLIII.The Wallichian Herbarium. Bulletin of Miscellaneous

Information Kew. pp. 255—263.

Baker, J.G. 1890. Curcuma. The Flora of British India. 6: 209-216.

Balakrishnan, N.P. and N. Bhargava. 1984. The genus Curcuma L.

(Zingiberaceae) on Andaman and Nicobar Islands. Journal of the Bombay

Natural History Society. 81: 510—514.

Bhat, K.G. 1987. Curcuma oligantha Trimen (Zingiberaceae) — a new record

for India. Indian Journal of Forestry. 10: 66—68.

Dammer, U. 1890. Curcuma roscoeana Wall. The Gardeners’ Chronicle. p.

592.

Forman, L. L. 1997. Notes concerning the typification of names of William

Roxburgh’s species of Phanerogams. Kew Bulletin. 52: 513-534.

Hooker, J. D. 1852. Curcuma roscoeana. Botanical Magazine. 78: t. 4667.

Hooker, J. D. 1870. Curcuma petiolata. Botanical Magazine. 94: t. 5821.

Jain, S.K. and V. Prakash. 1995. Zingiberaceae in India: phytogeography

and endemism. Rheedea. 5: 154-169.

Karthikeyan, S.,S.K. Jain, M.P. Nayar and M. Sanjappa 1989. Florae Indicae

Enumeratio Monocotyledoneae. Botanical Survey of India, Calcutta. pp.

289-299.

Lemaire, C. 1854. Curcuma roscoeana. Le Jardin Fleuriste. 4: t. 327.

Mangaly, J.K. and M. Sabu. 1988. A new species of Curcuma from South

India. Notes from the Royal Botanic Garden Edinburgh. 45: 429-431.

Mangaly, J.K. and M. Sabu. 1993. A taxonomic revision of the South Indian

species of Curcuma L. (Zingiberaceae). Rheedea. 3: 139-171.

Roscoe, W. 1827. Curcuma petiolata. Monandrian Plants of the Order

Scitamineae. George Smith. Liverpool. England.

Curcuma roscoeana in India 197

Roxburgh, W. 1820. Monandria monogynia. Flora Indica. Serampore, India.

pp. 1-84.

Santapau, H. 1945. Miscellaneous Notes 19. Curcuma pseudomontana Grah.

Journal of the Bombay Natural History Society. 45: 618—624.

Sivarajan, V.V. and I. Balachandran. 1983. A new species of Curcuma

from Southern India. Notes from the Royal Botanic Gardens Edinburgh.

41; 321-3723.

Skornickova, J. and M. Sabu. 2002. The genus Curcuma L. in India: Resume

and Future Prospects. In: A.P. Das (Ed.) Perspectives of Biology, Bishen

. Singh, Mahendra Pal Singh, Dehra Dun. pp. 45—S1.

Skornickova, J., M. Sabu and M.G. Prasanthkumar. 2003a. New species of

Curcuma from Mizoram. Gardens’ Bulletin Singapore. 55: 89-95.

Skornickova, J.. M. Sabu & M.G. Prasanthkumar. 2003b. Curcuma

codonantha (Zingiberaceae) A new species from the Andaman Islands,

v India, Gardens’ Bulletin Singapore. 55: 219-228.

Skornickova, J.,M. Sabu and M.G. Prasanthkumar. 2004. Curcuma mutabilis

(Zingiberaceae): A new species from South India. Gardens’ Bulletin

» Singapore. 56: 43—54.

Srivastava, S.K. 1998. Zingiberaceae in Andaman and Nicobar Islands,

India. Higher plants of Indian sub-continent. Indian Journal of Forestry.

Additional Series X. 8: 1—33.

Stafleu, F.A. and R.S. Cowan. 1988. Wallich, Nathaniel. Taxonomic

Literature 7: 37-41, (2™ edition). _

Tripathi, S. 2001. Curcuma prakasha sp. nov. (Zingiberaceae) from North-

eastern India. Nordic Journal of Botany. 21: 549-550.

Tripathi, S. and V. Prakash. 1998. Studies on Zingiberaceae of N.E.India:.

A new distributional record of Curcuma petiolata Roxb. Journal of

Economic and Taxonomic Botany. 22: 468—470.

Valeton, T. 1918. New notes on the Zingiberaceae of Java and Malaya.

Bulletin du Jardin Botanique de Buitenzorg. 27: 1-166.

Velayudhan, K.C., V.A. Amalraj and V.K. Muralidharan. 1996. The

conspectus of the genus Curcuma in India. Journal of Economic and

Taxonomic Botany. 20: 375—382.

198 Gard. Bull. Singapore 57 (2005)

Wallich, N. 1829. Plantae Asiaticeae Rariores I: 8, t.9. London, England.

Wallich, N. 1829-1832. A Numerical List of Dried Specimens of Plants in

the East India Company’s Museum, Collected under the Superintendence

of Dr. Wallich of the Company’s Botanic Garden at Calcutta. London.

Gardens’ Bulletin Singapore 57 (2005) 199—210 199

The Identity and Distribution of Curcuma

zanthorrhiza Roxb. (Zingiberaceae)

J. SKORNICKOVA !2 AND M. SABU !

‘Department of Botany, University of Calicut, Kerala 673635, India

"Department of Botany, Charles University, Benatska 2,

Prague, 12802, Czech Republic

Abstract

Curcuma zanthorrhiza Roxb., a widely distributed and utilized Asian species, has been

misidentified in India for over 100 years. A description and colour plate of Curcuma

zanthorrhiza are provided, with additional notes on C. zedoaria (Christm.) Roscoe and C.

aromatica Salisb., which are two common misidentifications of C. zanthorrhiza in India. It

is postulated that it is of South Indian origin and has been widely distributed by the

Srivijaya civilisation that spread through SE Asia long before Western domination.

Introduction

Skornickova and Sabu (2005) provided a general introduction to the genus

Curcuma and pointed out that the identities of many Curcuma species

described in earlier taxonomic publications from India, including those

commonly used and cultivated, are still very often unclear. One such case

is discussed here in detail.

A large, handsome Curcuma species is widely grown in South India.

It has a red patch along the midrib of the leaves that penetrates to the

lower surface, a lateral inflorescence, a large branched rhizome and root

tubers, which are deep orange colour inside. Resembling true turmeric (C.

longa L.) in its orange rhizome, this is one of the most common Curcuma

species used by local women as a face rub, medicine and sometimes also as

a substitute for true turmeric.

This species is mostly referred to in both old literature and in recent

Indian floras as Curcuma zedoaria (Christm.) Roscoe or as C. aromatica

Salisb. Further investigation shows that descriptions of C. zedoaria and C.

aromatica in the Indian literature are often confusing and misleading. These

publications usually contain either a very short description based on earlier

works with not many details particularly of diagnostic characters, or else

they are obscure and do not match the original descriptions of either species.

Descriptions of these two species in Indian floras fit our plant, however, or

share a combination of characters appropriate for C. zedoaria and C.

200 Gard. Bull. Singapore 57 (2005)

aromatica. After critical examination of fresh flowering material, it was

clear that the plant we have collected several times from different and

quite distant localities was neither C. zedoaria (Christm.) Roscoe nor C.

aromatica Salisb., but C. zanthorrhiza Roxb. and that there is a need to

clarify the distinction between these three species.

Curcuma zanthorrhiza Roxb.

The name Curcuma zanthorrhiza first appeared as a nomen nudum in

Roxburgh’s Hortus Benghalensis (1814). A few years later Roxburgh

described it from cultivated material in the Calcutta Botanical Garden,

which had been donated in 1798 by C. Smith from Amboina (nowadays

called Ambon) and which flowered for the first time in April and May

1810 (Roxburgh, 1820). The main character that caught Roxburgh’s

attention was the deep yellow internal colour not only of the branched

rhizome, but particularly of the root tubers. Usually, root tubers in Curcuma

species are either pure white or creamy or yellowish and much lighter in

colour than the central part of the rhizome and branches. Another

prominent character mentioned in Roxburgh’s original description 1s ‘/eaves

broad lanceolar, and oblong; there is a narrow purple cloud down the middle

of them, which penetrates to the underside.’

Roxburgh (1820) in his original description did not cite any

herbarium material. As explained by Forman (1997), the main difficulty in

typification of Roxburgh’s species lies in locating original material.

Roxburgh apparently did not keep his own personal set and his collections

can be found in at least 16 herbaria. Moreover, in many cases original

labels have been discarded and replaced by newly written ones and, owing

to lack of direct evidence, such sheets are less desirable for typification.

Yet, for most of the species described by Roxburgh in Flora Indica there

are fine life-size colour drawings, which usually also depict details of

dissected flowers. Their importance has been emphasized by Sealy (1957)

and further elaborated by Forman (1997) who concluded that in many

cases the Roxburgh drawing is far superior to a corresponding specimen.

This is particularly true for gingers considering the difficulties in preparing

good and valuable specimens and the particular importance of the colours

of various parts of Curcuma species (e.g. colour of rhizome, flower parts),

which cannot be well preserved in herbarium specimens.

Recently, Newman et al. (2004) cited the type for C. zanthorrhiza

as ‘Icones Roxburghianae 2003 (CAL). However, there are two copies of

the drawing No. 2003 available, one at Kand one at CAL (Sealy, 1957;

Sanjappa, 1994) and Flora Indica drawings have never been officially

Curcuma zanthorrhiza 201

published. Thus, one of them should be selected as the lectotype. We have

been granted permission to access the [cones Roxburghianae at K only,

leaving us unable to verify the presence and identity of a presumably

identical CAL icon of C. zanthorrhiza.

Valeton (1918) and Holttum (1950) provided more detailed

descriptions of C. zanthorrhiza. As already pointed out by Valeton (1918),

Ridley in his earlier works (1899, 1907, 1909) also erred by associating C.

zedoaria with the description and vernacular name Temu Lawas, both of

which should undoubtedly be subsumed under C. zanthorrhiza.

Subsequently Ridley (1924) cited C. zanthorrhiza with its proper vernacular

name (Temu Lawas) as a cultivated species in the Malay Peninsula, but his

description of C. zedoaria remained confusing. Based on the quite detailed

descriptions of C. aromatica provided by Watt (1889) and Dymock et al.

(1893), there is no doubt that they also misidentified C. zanthorrhiza for C.

aromatica and that the English common names Wild Turmeric, Yellow

Zedoary and Cochin Turmeric quoted by both Watt (1889) and Dymock et

al. (1893) for C. aromatica are in fact common names of C. zanthorrhiza.

Curcuma zanthorrhiza Roxb., Fl. Ind. 1 (1820) 25. — Type: [cones

Roxburghianae 2003 (lectotype K!, designated here)

Curcuma xanthorrhiza Roxb., Syn. Pl. 1 (1839) 19., orth. var.

C. aromatica auctt. non Salisb.: G. Watt, Dict. Econ. Products India 2

(1889) 655-658; W. Dymock, Pharmacographia Indica 6 (1893) 396-398;

K.S.Manilal, Fl. Silent Valley. (1988) 311-312.

C. zedoaria auctt. non (Christm.) Rosc.: Ridley, J. Straits Branch Roy.

Asiat. Soc. 32 (1899) 119; Materials for a Fl. Malay Penins. (1907) 21;

Philipp. J. Sci. 4 (1909) 166; Mangaly & M. Sabu, Rheedea 3 (1993) 168.

Rhizomatous herb to 2 m high. Rhizome branched, central part oblong, c.

8—10 x 6—8 cm, brownish orange outside, deep bright orange to yellow-

orange inside, strongly aromatic with a carrot-like and camphoraceous

smell and taste, slightly bitter, rhizome branches 5—15 cm long, 1.5—4 cm in

diam., brownish-orange outside, deep yellow-orange to dark orange inside,

youngest branches lighter in colour. Root tubers present at the end of 1-3

mm thick roots at 5—20 cm from the main rhizome or branches, elliptic, 3—

8 x 1.5—3 cm, brown outside, usually with many small roots, deep yellow-

orange inside, aromatic (less so than the main rhizome), bitter in taste.

Pseudostem to 70 cm, green, composed of leaf sheaths and sheathed by 4—

5 green bracts, innermost bracts as long as the pseudostem, outer ones

gradually decreasing in length, /igu/e 2-3 mm, obscurely bilobed, hyaline,

translucent greenish white, glabrous, hairy on the margin, hairs c. 0.3—0.5

mm; leafy shoot to 2 m tall. Leaves at the beginning of the season 1-3, later

202 Gard. Bull. Singapore 57 (2005)

to 8; petiole 5—20 cm (first leaves almost sessile), winged on both sides,

glabrous; /amina oblong-lanceolate to elliptic-lanceolate, the very first leaves

more elliptic, c. 30-100 x 10—28 cm, glabrous on both surfaces, adaxially

deep green with red patch along both sides of midrib, particularly

conspicuous in young leaves, fading with age, abaxially lighter green, the

red patch is also lighter but usually visible; midrib glabrous, green to reddish

on the upper side, green below; margin translucent white, c. 0.7-1 mm

wide, glabrous; tip acuminate, 2—3 cm, slightly hairy; base attenuate,

decurrent. /nflorescence invariably lateral, arising together with the leaves

or shortly before. Peduncle 10—30 cm, 0.8—2 cm diam. (without scales),

green, glabrous, sheathed by 4—6 green, glabrous sterile bracts, innermost

bract longest, outer ones gradually smaller. Spike c. 15—25 (—30) x 8-14

cm. Coma present, forming upper third of inflorescence length, coma bracts

oblong-elliptic, c. 10-17, 6.5—8.5 x 2.5—4 cm, pink to deep reddish-pink,

shortly hairy on both surfaces, hairs 0.1—0.2 mm long, tip slightly mucronate,

hairy, lower coma bracts sometimes fertile, upper ones sterile. Fertile bracts

roundish-oblong, 5—6 x 4—5 cm, green, tips tinged with pink, almost glabrous,

connate to one another in the lower third. Cincinni with 5-7 flowers.

Bracteoles one per flower, ovate, boat-shaped, c. 3.5 x 2 cm to 1.5 x 0.7 cm

(outer one larger, inner ones are gradually smaller), translucent white,

glabrous, but the tip, upper part and margins sparsely hairy. Flowers 5—6

cm, as long as the bracts. Calyx 10-11 mm long, translucent white sometimes

with a slight pink tinge, sparsely hairy, unilaterally split for 4-5 mm, apex

with 3 teeth. Corolla tube 3—3.8 cm long, outside light yellow in the lower

part with a light pink tinge in the upper part, glabrous, inside yellow,

constricted c. 2.3 cm above the ovary, constriction densely hairy; dorsal

corolla lobe 1.5—2.1 x 1.3—2 cm, triangular ovate, concave, glabrous, light

pink or pink, apex mucronate, mucro 2—3 mm, lighter in colour, hairy,

hairs 0.2—0.3 mm; lateral corolla lobes 1.5—1.8 x 1.5—1.7 cm, triangular with

a rounded slightly concave tip, glabrous, light pink to pink, usually slightly

overlapping each other at the base. Lateral staminodes obovate, 1.3—1.5 x

0.9-1.1 cm, light yellow, glandular hairs present on the raised middle

portion. Labellum 1.7—2 x 1.8—2 cm, obscurely trilobed, lateral lobes folding

upwards, emarginate, split c. 3 mm, light yellow at the periphery, deep

yellow in the centre (golden median band). Anther spurred, glandular hairs

present on the sides and back part of the anther, anther thecae 3.5—4 mm

long, white; filament 4—5 mm long, light yellow, 3.54.5 mm at base, 2.5—3

Plate 1. Curcuma zanthorrhiza Roxb. 1. Midrib of leaf (adaxially); 2. Midrib (cross section); 3.

Midrib of leaf (abaxially); 4. Inflorescence; 5. Habit; 6. Flower in open fertile bract; 7. Dissected

flower; 8. Anther (front view); 9. Anther (side view); 10. Ovary and epigynous glands, cross

section of ovary in upper left corner. (‘kornikov- 73302 & 84107) Photo J. Skornickova.

Curcuma zanthorrhiza

ee! ee

6 7 8

204 Gard. Bull. Singapore 57 (2005)

mm at the top. Anther spurs 3—4 mm long. Stigma white, ciliate, exserted

from thecae by c. 1 mm. Epigynous glands two, light yellow, 4-5 mm long,

0.9-Imm diam. Ovary 4—5 x 34 mn, trilocular, hairy, hairs c. 0.3 mm.

Fruits not seen.

Specimens examined: AFRICA: Congo, Eala, 1930, Corbisier 980 (K);

MADAGASCAR: II. 1880, Hildebrant 3348 (BM); CHINA: Fokien, 7.

VII. 1909, Dunn 3547 (K); Wai-yeung, HI. 1932, Tsui 98 (K); INDIA:

Kerala: Kasargod District: Nileshwar, 12.V.1982, V.J.Nair 73856 (CAL,

MH); Cannanore District: Peria R.F., 17.1V.1966, J.L. Ellis 27113 (MH);

Waynad District: Sulthan Bathery, Beenachi Estate, 14.V.2003, Skornickova

84126 (CALI, SING); Pallakad District: Silent Valley, Sivarajan SV 10565

(CALI); Pallakad, 10.VI.1983, sine coll. (J.K. Mangaly?) CU 10364 (CALI);

Palai, 10.VI.1983, J.K. Mangaly CU 10365 (E); Idukki District: Kulamavu,

23.1X.2003, Skorni¢kova 84166 (CALI); Nadukani, 23.1X.2003, Skornickova

84172 (CALI); Kollam District: Dahli, 23.1V.2003, Skornickova 84107

(CALI, SING); Kottayam District, Vazhoor East, 8.1V.2002, Prasanthkumar

86111 (CALI); Pathanamthitta District, Sabarigiri, 25.1.1984, M. Sabu

CU37315 (CALI); Pamba, Sabarimala R.F., 24.1V.1984, Vajravelu 83580

(MH); Andaman Islands: Rangat District, Amkunj near Bakultala,

17.V.2002, Skornickova 73302 (CALI, SING); Jharkand: West Singbhum

District, Khutpani, 5.VII.2003, Skornickova 73420 (CALI, SING); sine loc:

Herb. Hort. Bot. Calcuttensis, sine dat., Acc. No. MH 72400 (MH); Hortus

Botanicus Calcuttensis, sine dat., Acc. No. CAL 467008 (CAL); sine col. et

dat. Acc. No. CAL 466954 (CAL); sine col.et dat., No. 1612 (DD); presented

1871, Wight s.n. (MH); sine dat., Wight s.n. (E); SRI LANKA: Gangaruwa,

22. VII. 1924, Siwa 167 (PDA); CAMBODIA: Koh Kong, 3. I. 2000, Meng

Monyrak 115 (K); MALAY PENINSULA: Johore, 21.V. 1954, Sinclair

8079 (E); Johore, 21.V. 1954, Sinclair 40295 (BM, E, K); SARAWAK:

Lubok Antu, 29. X. 1993, Christensen 1353 (K); Lundu, 19.IX. 1955,

Purseglove & Shah P 4584 (K, SING); JAVA: Buitenzorg, sine coll., Col.

No. 42 (BO); West Java, VII. 2000, M. Ardiyani 29 MA (E); SOUTH

KALIMANTAN: 8. XI. 1996, Kessler et al. PK1755 (K); PHILIPPINES:

Mindoro: Mansalay, IV. 1903, Merril 908 (K); Luzon: San Francisco del

Monte, Loher 682 (K); VI. 1904, Loher 7003; 3. V. 1908, Elmer 7003 (K);

V. 1905, Whitford 1267 (K); Manila, 17. V. 1890, Loher 683 (K); ORIGIN

UNKNOWN: 30. VII. 1999, M. Ardiyani 80 MA (E); 26. XI. 1978; HS.

McKee 36132 (E).

Flowering: In India, April and May.

Distribution and habitat: Common in South West India where it grows in

Curcuma zanthorrhiza 205

the edges of secondary forests or in the undergrowth. It is common in

semi-wild conditions e.g., teak plantations, coconut groves, along roadsides

and rarely also in high altitude grasslands (c. 1000 m a.s.1.), which suggests

it is native in South India.

We have collected it also in the Andaman Islands from the garden

of a Bengali family. It was certainly cultivated but its origin was obscure.

Either it was brought from Bengal by their ancestors or other settlers, who

came to the Andamans from South India, may have brought it. One recent

collection was from a garden in Jharkhand, Central India, where this

Curcuma plant is cultivated as a substitute for turmeric, C. Jonga. This is

probably a recent introduction.

It is interesting to note that in India up to 1845 C. zanthorrhiza

Roxb. was recorded only as cultivated in the East India Company’s

Botanical Garden in Calcutta from an introduction from Ambon (Roxburgh,

1814; Voigt, 1845). Thus, even though most probably native, widely

distributed, naturalized, cultivated and commonly used in South India for

centuries, C. zanthorrhiza is here recorded for the first time from India.

Since Roxburgh’s time, Curcuma zanthorrhiza has been reported

from Java (Valeton, 1918), Peninsular Malaysia (Holttum, 1950), Vietnam

(H6, 1993), Thailand (Larsen, 1996), the Philippines (Madulid, 1996) and

China (Wu and Larsen, 2000). It is therefore not surprising that a species

so widely distributed and frequently cultivated all over Asia occurs in

India.

In addition to literature records, we found specimens looking

identical to Curcuma zanthorrhiza from different parts of Africa,

Madagascar, Sri Lanka, Cambodia, and Borneo (Kalimantan). From

remarks on the sheets, it is obvious that most of the material was of

cultivated origin.

It is difficult to establish where Curcuma zanthorrhiza is native

because it is so widely distributed, cultivated and naturalized for centuries

all over Asia and can now be found as far away as West Africa. Yet, there

might be a historical explanation. During the first centuries A.D., the

Sriviaya Civilisation of the Hindu Kingdom in South India started

expanding through SE Asia. Based in eastern Sumatra, they dominated

the Malacca and Sunda straits, which were the two main sea routes between

the Indian Ocean, China Sea and Indonesia, and controlled the trade of

the region. They heavily influenced cultures in SE Asia until the 13 century

A.D., when Srivijaya had lost control. It is thus not surprising that the SE

Asian region was, prior to Western dominance, known as Greater India.

There is a possibility then that C. zanthorrhiza originated in the southern

part of the Indian subcontinent and was introduced to other countries in

SE Asia, perhaps as part of the spice and medicine trade. Dymock et al.

206 Gard. Bull. Singapore 57 (2005)

(1893) say that this plant is the Vana-haridra of Sanskrit writers (meaning

‘wild turmeric’) and also mention that it is the ‘turmeric-coloured zedoary

of Ainslie used by the Mahometans of Southern India as valuable medicine

in snake bite...” and say that it was well known to Rumphius (the Dutch

botanist working in Ambon in the 17" century), who called it Tommon

bezaar or Tommon primum. It was from Ambon that it was brought to

Calcutta where Roxburgh published it as a valid binomial. Roxburgh did

not work in South India so he did not realize it was native in there. And

today we know C. zanthorrhiza does not occur in Central and Northern

India, so he would have had no access to Indian material. This partly

explains the long history of misidentifying of C. zanthorrhiza in South

India.

Etymology & orthography: Greek: zanthos (yellow) rhizos (root or rhizome).

Curcuma zanthorrhiza versus C. aromatica and C. zedoaria

Curcuma. zanthorrhiza has for a long time been misidentified as C. zedoaria

and C. aromatica in India. They are all early flowering species with a

lateral inflorescence, which arises shortly before the leaves appear or

simultaneously with leaves, sharing in common the pink conspicuous coma

and yellow flowers. The main differences among these species are shown

in Table 1.

Acknowledgements

The authors thank the Department of Science and Technology, Government

of India (Order No. SP/SO/A-20/99 dt. 09.11.2001), the Grant Agency of

the Academy of Sciences of the Czech Republic (Grant No. B.6407401)

and the International Association of Plant Taxonomy for financial support.

We are indebted to authorities of Ministry of Environment and Forests for

granting permission for collecting. The first author thanks the curators of

BM, BO, CAL, CALI, DD, E, K, MH, PDA and SING for letting her

examine specimens in their care, and to the library of K for granting access

to [cones Roxburghianae, the Singapore Botanic Gardens for awarding a

Singapore Botanic Gardens Research Fellowship and facilities during her

repeated visits, to Dr. A. D. Poulsen (E), Dr. M. F. Newman (EB), Prof.

Emeritus K. Larsen (AAU), Ms. T. Rehse (Duke University), Dr. R.

Kiew (SING), for helpful comments on the manuscript.

Curcuma zanthorrhiza

207

Table 1. Comparison of important morphological characters of Curcuma

zanthorrhiza Roxb., C. aromatica Salisb. and C. zedoaria (Christm.) Roscoe.

Main diagnostic characters are in bold.

C. zanthorrhiza Roxb.

C. aromatica Salisb.

C. zedoaria

(Christm.) Roscoe

Main branched, inwardly branched, inwardly branched. inwardly

rhizome deep bright orange cream to pale brown cream to white-

to yellow-orange colour. yellowish colour.

colour.

Root deep yellow-orange white inside. white inside.

tubers inside.

Leafy to 2 (-2.5 m) tall. to 1.2 m tall. to 1 m tall.

shoot

Lamina oblong-lanceolate to oblong-lanceolate to oblong-lanceolate to

elliptic-lanceolate, elliptic- lanceolate, elliptic- lanceolate,

adaxially green with adaxially bright adaxially green with

red patch along the green, glabrous, red patch along the

sides of midrib, abaxially lighter sides of midrib,

particularly green, densely particularly

conspicuous in young shortly pubescent. conspicuous in young

leaves, fading with leaves, fading with

age, protruding age, glabrous,

underneath, glabrous, abaxially lighter

abaxially lighter green green, glabrous.

and visible red patch

(also lighter then on

adaxial side),

glabrous.

Flowers 53—6 cm long, as 5—6 cm long, longer 5—6 cm long, as

long as the bracts than the bracts long as the

or slightly slightly exserted. bracts or

exserted. slightly exserted.

Corolla __ conspicuously pink white, sometimes nearly white or

lobes to reddish. with slight pinkish with very slight

shade toward the

tips.

pinkish shade.

208 Gard. Bull. Singapore 57 (2005)

References

Burtt, B.L. 1977. Curcuma zedoaria. Gardens’ Bulletin Singapore. 30: 59—

62.

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Gardens’ Bulletin Singapore 57 (2005) 211—216 211

A New Species of Rhaphidophora Hassk.

(Araceae-Monstereae) from Borneo

Pe BOYCE

Malesiana Tropicals, Suite 9-04, Tun Jugah Tower,

No. 18, Jalan Tunku Abdul Rahman93100 Kuching, Sarawak, Malaysia

Abstract

Rhaphidophora typha P.C.Boyce, a new species from Borneo distinguished by its remarkably

long peduncle and leaves abaxially grey-glaucous, is described and illustrated.

Introduction

Revisionary work on Rhaphidophora (Boyce, 1999, 2000a, 2000b, 2001a,

2001b) has highlighted that although Borneo has a relatively few species

(13 species excluding the one described here), five (excluding that described

here) are endemic compared with 15/1 (total/endemic) in Peninsular

Malaysia and 15/4 in Sumatera. Thus Borneo, with the exception of the

Philippine Islands (11/7), has the richest level of Rhaphidophora endemism

in Sunda — although these data are insignificant compared with New Guinea

and tropical Australasia (30/29). Boyce (2001a) speculated that the paucity

of Rhaphidophora species recorded for Borneo was an artefact of

inadequate collecting.

Since the above publications, the author has had the opportunity

to spend an extended period undertaking fieldwork in Sarawak. While

astonishingly productive in locating novel terrestrial aroids, this fieldwork

has generally supported the impression that the species complement of

climbing aroids in Borneo is, indeed, meagre. Nonetheless, undoubtedly

further lianescent novelties still await discovery. One such novelty, a

herbarium specimen originating from Brunei, was located during a study

visit to the Singapore Herbarium (SING) in Janary 2004 and was

subsequently collected in Sarawak. It is described here.

Rhaphidophora typha P.C. Boyce, sp. nov.

Rhaphidophora typha a pluribus speciebus generis Borneensibus combinatio

foliorum subtus griseo-glauca, veneris laminorum valde reticulatis et

pedunculum infloresecentia alta demi petiolorum laminae differt.— TYPUS:

Brunei Darussalam, Temburong District. Sungai Temburong, just

Gard. Bull. Singapore 57 (2005)

downstream from Kuala Belalong. 19 July 1988, K.M. Wong WKM 242

(SING, holo!; BRUN!, K!, L!, iso).

Figure 1.

Slender liane to 3 m in height; seedling stage and pre-adult plants not

observed; adult shoot clinging and flowering terminally with inflorescence

soon displaced (well before infructescence maturity) and thus

physiognomically lateral; stems smooth, mid- to dark green, with long,

sparse petiolar sheath fibre on the most recently matured portions,

internodes up to 10.5 cm x c. 7 mm, separated on still-leafy portions by

deep brown clasping leaf base, this up to 6 mm long; flagellate foraging

stems not observed; clasping roots few, arising from the nodes, velvety;

feeding roots frequently stilt-like in terrestrial individuals, adhering to the

climbing surface and reaching the ground in climbing individuals, smooth;

leaves spiro-distichous, distally clustered on adult shoots; cataphylls and

prophylls membranous, soon drying and falling and only rarely persisting

as scattered long fibres; petiole narrowly canaliculate, 18—24 cm x c. 2 mm,

smooth, apical geniculum prominent in dried material, up to 3.5 cm long,

basal geniculum also prominent, up to 1.5 cm x 6 mm; petiolar sheath

prominent, extending to the apical geniculum, soon degrading into semi-

persistent long fibres; /amina entire, narrowly-oblanceolate, 34—35 x 7—7.5

cm, when fresh thinly leathery with adaxial surface matt pale olive-green

and abaxial surface obscurely grey glaucous, drying papery with adaxial

surface mid-orange-brown and abaxial surface slightly grey-glaucous, base

long-decurrent, apex acuminate-attenuate; midrib prominently raised

abaxially, level adaxially; primary venation pinnate, raised abaxially, slightly

impressed adaxially; interprimaries subparallel to primaries, slightly raised

abaxially, + flush adaxially, forming a weak reticulum distally; secondary

venation prominently reticulate, raised abaxially; tertiary venation a network

of broadly spaced tessellate veins arising at c. 90 from the midrib and

crossing the primaries and interprimaries; inflorescences solitary; peduncle

terete, up to 18 cm x 2 mm; spathe not observed but, based on observing

young infructescences, seemingly caducous; spadix cylindrical, sessile,

inserted level on peduncle, 6.5 cm x 6 mm, green in juvenile fruiting stage;

stylar region rhomb-hexagonal, c. 1.5 x 1.2 mm, truncate; stigma elliptic,

longitudinally orientated, up to 0.75 mm, prominent in dried material:

anthers not observed; mature infructescence not observed.

Figure 1. Rhaphidophora typha PC. Boyce Holotype specimen (SING) showing the diagnostic

long-pedunculate inflorescence.

Rhaphidophora typha from Borneo 213

fivegerngrng

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SINGAPORE BOTANIC GARDENS

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1 ~. 2 3 4 5 6 7 8 9 10 1 12 13 14 15 16 17 18 19 20 zt 22 23 24 25 26 j 28

i deep green. Spathe greenish. Spadix with green fruit.

Det. PC Bouce Detdate: 4 |

SINGAPORE BOTANIC GARDENS * U9 “0% K.M. Wong WKM 242 19 duly 1988

= ; Brunei. Temburong District. Sungai Temburong, Jus

: | Rhaphidephora Typh 4 eles Bouce domstrean from Kuala Belalong. Common herb. Foliage

i Rhap Lidopha ce wpe. nov

(/ Singapore. a Det. Date :

‘ Fiborycs zi /Z Jou ;

214 Gard. Bull. Singapore 57 (2005)

Distribution: BORNEO: Brunei and Sarawak.

Habitat: Although habitat is not recorded on the type, Kuala Belalong is

predominantly mixed moist lowland dipterocarp forest on Setap shales

with some gallery forest in the valley bottoms and kerangas elements on

the ridge tops. In Sarawak, R. typha occurs in riverine or gallery forest on

shales exposed by river action. Altitudes in Kuala Belalong range between

15 and 350 m. The Sarawak collection cited below is from 240 m.

Notes: 1. Rhaphidophora typha bears some resemblance to R. beccarii

(Engl.) Engl. but is readily separable by the much thinner abaxially grey-

glaucous leaf laminas, the slender, less thickly-rooted stems (roots arising

only at the nodes in R. typha, rooting along the internodes in R. beccarii),

the smooth, not scaly feeding roots and the markedly longer, thinner

peduncle. The prominently reticulate secondary and tessellate tertiary

venation is also noteworthy. As noted, the ecology of R. typha in Brunei is

not recorded. In Sarawak it grows as a low climber (rarely up to 3 m) on

slender trees, while R. beccarii is an obligate rheophyte.

2. There exist in Borneo three further lanescent aroids with glaucous

abaxial leaf surfaces with which confusion with R. typha might occur. They

are: Scindapsus longipes Engl., S. glaucescens (Engl. & K.Krause) Alderw.

and an as yet undescribed Anadendrum. Scindapsus longipes differs from

R. typha by the stiffly coriaceous ovate leaf lamina with the primary veins

hardly visible while S. glaucescens has oblong leaves up to 1 m long with

abaxially the primary lateral veins raised but the interprimary and secondary

veins hardly visible. Both Scindapsus are found only in hill forest on

sandstones. The Anadendrum has leaves similar in texture and venation to

those of R. typha but is readily distinguished by the strictly distichous leaf

arrangement.

3. The specific epithet is from the Greek typhe (cat-tail) and alludes to the

proportionately very long-pedunculate inflorescence, the peduncle reaching

over half the petiole length, which somewhat fancifully resembles the

inflorescence of Typha (Typhaceae - Poales). Such inflorescence

morphology is very unusual among long-petiolate Rhaphidophora and not

hitherto recorded for any Bornean species although occurring in a

morphologically very different species in the Philippines, R. monticola

K.Krause.

4. Rhaphidophora typha can be fitted into the key to Bornean

Rhaphidophora (Boyce 2001a) as follows:

Rhaphidophora typha from Borneo DAS

3a. Geniculum and abaxial surface of lamina pubescent..............ceeeeee 4

3b. Geniculum and abaxial surface of lamina glabrous or glaucous. ......... 5

4a. Plants flowering on clinging stems. Leaves of mature plants extensively

perforated, active shoot tips with black mucilage ........ 6 R. foraminifera

4b. Plants flowering on free lateral stems. Leaves of mature plants lacking

or with only with scattered, perforations; active shoot tips lacking black

TOA eee oR oe rd cecncee deg ea Renec anv ettutepioas- ab aptoes coco 12. R. puberula

5a. Leaves always shingling, even in flowering individuals. Flowering on

CLONES OIEES (C6 | Sa elite 6 os et oe en ee ee 8. R. latevaginata

5b. Leaves spreading in adult and flowering individuals. Flowering on

Hines aera (MD ONIN NOES ecsge fears ceees ac oe eee stand istened Utell llide deena 6

6a. Stems scabrid to asperous. Spathe exterior minutely puberulent ..........

rer ere ee tee ew SE, PEE oe acisedvisctdccistiesseedtanes 9. R. lobbii

6b. Stems smooth. Spathe (where known) exterior glabrous .................. 7

7a. Abaxial surface of lamina glaucous. Peduncle up to 18 cm long ............

Fe ee eat eee SANE Hat Seats Meta cocathcoaee Wen albidecbek Cbedueebivuvabentas R. typha

7b. Abaxial surface of lamina never glaucous. Peduncle not exceeding 10

SI OTR Scr Ne a8 ee Gin A OR ne ae | er 8

8a = 7a, etc., in Boyce (2001a)

Other specimen seen: SARAWAK. Kapit Division, Nanga Gaat, Rejang

Wood Concession, KM 55 road to Camp Gahada, 01° 44' 44.5"N; 113° 28'

32.3"E, 13 May 2004, P.Boyce, Jeland ak Kisai & Jipomak Tisai AR-374

(SAR).

References

Boyce, P.C. 1999. The genus Rhaphidophora Hassk. (Araceae-

Monsteroideae-Monstereae) in Peninsular Malaysia, and Singapore.

Gardens’ Bulletin Singapore. 51: 183-256.

Boyce, P.C. 2000a. The genus Rhaphidophora Hassk. (Araceae-

Monsteroideae-Monstereae) in the Southern and Western Indonesian

Archipeiago. Gardens’ Bulletin Singapore. 52: 101-183.

Boyce, P.C. 2000b. The genus Rhaphidophora Hassk. (Araceae-

Monsteroideae-Monstereae) in the Philippines. Gardens’ Bulletin

Singapore. 53: 213-256.

Boyce, P.C. 200la. The genus Rhaphidophora Hassk. (Araceae-

Monsteroideae-Monstereae) in Borneo. Gardens’ Bulletin Singapore. 53:

19-74.

216 Gard. Bull. Singapore 57 (2005)

Boyce, P.C. 2001b. The genus Rhaphidophora Hassk. (Araceae-

Monsteroideae-Monstereae) in New Guinea, Australia and the Tropical

Western Pacific. Gardens’ Bulletin Singapore. 53: 75—183.

Gardens’ Bulletin Singapore 57 (2005) 217—246 ZAG

Taxonomic Notes on Bornean Litsea, Lindera,

Neolitsea and Iteadaphne (Lauraceae)

F.S.P. NG

Forest Research Institute Malaysia

52109 Kepong, Selangor, Malaysia

Summary

Litsea Lam., Lindera Thunb., Neolitsea Merr. and Iteadaphne Blume share

a combination of many features that distinguish them collectively as a

well-defined natural group from the rest of the Lauraceae. However, within

the group, the genera are defined on features that are unsatisfactory whether

considered individually or in combination.

The main line of division is anther locule number: four versus two.

This feature separates the 4-locular genera Litsea and Neolitsea from

the 2-locular genera Lindera and Iteadaphne. However, there are

several species on one side of the division that are mirrored by species

on the other side, differing only by the anther locule number. In this

revision, four such sets of twins are united as four species with variable

anther locule number. One of these species is Litsea cubeba (Lout.)

Pers., in which 2-locular and 4-locular anthers can occur even within

the same flower.

Another line of division uses a combination of leaf venation and

floral merism, by which triveined dimerous Neolitsea has been

distinguished from penniveined trimerous Litsea. These two-feature

states are not fixed, and some authors have defined Neolitsea on

dimery with or without triveined leaves. However, dimery itself is a

variable feature and the type species, Neolitsea cassia, has been found

to have both dimerous and trimerous flowers.

Litsea species can be distinguished from Lindera if the fruit has a

prominent cupule, but there are a few Litsea species without cupule

development (e.g. Litsea elliptica and all the species with variable

anther locule number), and these have often been misidentified as

Lindera.

[teadaphne is distinguished from Lindera in its umbellules reduced

to a single floret. By default, Lindera consists of species with 2—20

florets per umbellule. However, the number of florets in the

umbellules is actually variable. The type species of [teadaphne has

itself been found to be part of a variable species, Litsea

subumbelliflora (Blume) Ng, with 2 or 4 anther locules and 1—6

218 Gard. Bull. Singapore 57 (2005)

florets per umbellule. Hence /teadaphne is reduced to Litsea.

e Litsea, Lindera and Neolitsea should be united as a single natural

genus, but their merger would result in a great nomenclatural

upheaval. The alternative, adopted here, is to retain them as artificial

genera, for nomenclatural stability.

e In writing the account for The Tree Flora of Sabah and Sarawak, 9

new taxa (species or varieties) are described, 47 species are reduced

to synonymy and one species is excluded from Lauraceae.

Introduction

Litsea Lam., Lindera Thunb., Neolitsea Merr. and Iteadaphne Blume are

collectively distinguished from the rest of Lauraceae by a suite of

reproductive characters. The species are nearly always dioecious, rarely

polygamous (Litsea cubeba (Lout.) Pers., Litsea lancifolia (Roxb. ex Wall.)

Hook. f.) The inflorescence is an axillary raceme terminating in a dormant

terminal bud. However, instead of flowers, the raceme bears ‘involucral

buds’, each involucral bud consisting of a perianth-like involucre of 4

(sometimes 3 or 5) decussate bracts enclosing 1—20 florets in an umbellule.

The stalk of the umbellule is here called a peduncle and the stalks of the

florets are called pedicels. At anthesis, each involucral bud behaves like a

unified flower, with all its florets opening together to expose a mass of

protruding stamens, staminodes and stigmas on an open involucre imitating

a perianth. In the female flower, the ovary sits within a hypanthium

(perianth tube), with the perianth lobes (small and flimsy), staminodes,

and glands arising from the mouth of the hypanthium. The hypanthium is

enlarged in fruit to form a cup-like or dish-like cupule in most species of

Litsea and to some extent in Neolitsea, but not in Lindera. The number of

perianth lobes varies from 3 to 9, with 6 as the most common number, but

aberrant specimens have 2 and even no perianth lobes. In the male flowers,

the staminodes are replaced by functional stamens and the pistil by a

pistillode. The stamens range in number from 3 to 12 and rarely 18, with 9

as the most common number. Glands are usually present, as swollen pads

of tissue between the stamens and staminodes, sometimes free standing

and sometimes attached to the sides of the filaments; they sometimes look

like sessile anthers with bulging but non-functional anther locules.

I recently found a population of Litsea cubeba with hermaphrodite

flowers in the Bario Highlands, Sarawak. About ten trees were all heavy

with fruit, but some also bore flowers toward the twig tips. The flowers

had functional stamens (with swollen anther locules opening by uplifted

flaps to release pollen) and ovaries containing well-formed ovules. I then

Litsea, Lindera and Neolitsea in Borneo 219

re-examined my previous drawings of dissected flowers and confirmed that

Litsea cubeba has three distinct kinds of flowers: male, female, and

hermaphrodite, and that hermaphrodite trees are not only in Sarawak, but

also in Ranau, Sabah (SAN 114/66). I also found a hermaprodite flower in

Litsea lancifolia var. lancifolia in a specimen from Ranau, Sabah (Matamin

Rumuto MR 27), which I had previously taken to be male.

Within the group, there is abundant evidence that the genera are artificial.

The major internal line of division is whether the anthers are 2- or 4-

locular. Lindera and Iteadaphne are 2-locular whereas Litsea and Neolitsea

are 4-locular. This dividing feature stands almost alone, partially supported

by the development of cupules in most species of Litsea and Neolitsea and

the non-development of cupules in Lindera and Iteadaphne. Division by

anther locule number has resulted in species of Litsea having ‘twin species’

in Lindera/Iteadaphne, differing only in the number of anther locules. In

such cases, only male specimens can be identified positively. Female and

sterile specimens can only be placed by looking for exact matches in leaves

or localities. Twin species could have arisen by parallel evolution in different

genera but are more likely to be the result of an arbitrary line of division

based on a character that is in fact variable. I have united these twins and

placed them in Litsea:

1. Litsea subumbelliflora (Blume) Ng includes Litsea lanceolata (Blume)

Kosterm. and Lindera subumbelliflora (Blume) Kosterm.

2. Litsea cubeba (Lour.) Pers. includes Lindera pipericarpa sensu auct. non

(Miq.) Boerl., Lindera oxyphylla (Nees) Hook.f. and Lindera reticulosa

Kosterm.

3. Litsea lancifolia (Roxb. ex Wall.) Hook.f. and Litsea sessiliflora Hook.f.

have both been found to include male specimens with 2-locular anthers

that technically would qualify these specimens to be new species of Lindera

or [teadaphne, but such ‘species’ would consist of male specimens only.

Further details are given in the respective species accounts below.

The well-developed cupule is associated with Litsea but a few species

of Litsea (e.g. Litsea elliptica Blume, and all the species with variable

anther locule number) have weakly developed or undeveloped cupules,

and fruiting specimens of these species are often misidentified as Lindera.

Neolitsea was included in Litsea, until raised to generic level by

Merrill (1906). Previously, it had been Litsea Lam. sect. Neolitsea Benth.

The section Neolitsea was described by Bentham (in Bentham and Hooker,

1880) as follows: Folia saepius triplinervia. Flores sepius 2-meri, perianthi

segmentis 4, staminibus perfectis 6 (Leaves often triplinerved. Flowers often

220 Gard. Bull. Singapore 57 (2005)

2-merous, perianth segments 4, perfect stamens 6). By default, Litsea

became a genus with penninerved leaves and trimerous flowers. However,

dimery is not always associated with triveined leaves, and some authors,

e.g. Kostermans (1957) have dropped the triveined character from the

definition of Neolitsea, leaving dimery as the key character. There are

indeed 4 perianth lobes in N. amabilis and N. villosa but I have found

florets with 3, 4 and 6 perianth lobes in N. cassia, the type species of

Neolitsea.

Lindera differs from Litsea and Neolitsea only by anther locule

number, associated with lack of cupule development in fruit. Lindera

parallels the situation in Litsea prior to Merrill in that it consists of two

subgroups of species, one with triveined leaves and the other with

penniveined leaves. Lindera also has dimerous as well as trimerous flowers.

Iteadaphne is distinguished from Lindera by its umbellules bearing

a single floret. By default, Lindera consists of species with 2—20 florets per

umbellule. The arbitrariness of this line of division is quite obvious. The

number of florets in the umbellule is variable in most, perhaps all species.

Neolitsea amabilis Airy Shaw and Lindera montanoides Kosterm.

have 1—3 florets per umbellule. Both species have triveined leaves but one

is a Neolitsea because of 4-locular anthers and the other a Lindera because

of 2-locular anthers. Lindera bibracteata (Nees) Boerl. is consistently

uniflorous and could have been placed in /teadaphne but was placed in

Lindera instead; it has triveined leaves.

The type species of [teadaphne is ‘Iteadaphne confusa Blume’ (=

Lindera subumbelliflora (Blume) Kosterm.), and this has been found to

consist of local populations with uniflorous umbellules, mirrored by Litsea

lanceolata (Blume) Kosterm. with 4 anther locules. The two combined

make the widespread Litsea subumbelliflora (Blume) Ng that has 2 or 4

anther locules and 1-6 florets per umbellule. This merger has the effect of

reducing /teadaphne to Litsea. There are two other species of Jteadaphne

to be accounted for: Jteadaphne caudata (Nees) H.W.Li and Iteadaphne

philippinensis Elmer. These should be examined carefully to see whether

they fit into Lindera, Litsea or Neolitsea. It is likely that they already have

‘twin species’ in those genera, with which they should be united. /teadaphne

caudata has triveined leaves, suggesting a relationship with Lindera

bibracteata.

Litsea, Lindera and Neolitsea could be merged into a single genus,

which would be a natural genus because it would share many traits all

supporting each other. Such a genus could then be re-sorted into sections

(I would expect five or more sections) that are better defined than the

current three genera. However, merger would result in considerable

nomenclatural upheaval because there are about 400 binomials in Litsea,

Litsea, Lindera and Neolitsea in Borneo a74

100 in Lindera and 100 in Neolitsea. Lindera is the oldest name. For the

Tree Flora of Sabah and Sarawak, for which this research has been carried

out, the preferred option is to maintain Litsea, Lindera and Neolitsea as

separate artificial genera.

The following definitions appear to me to be the most practical for

conserving existing species names.

Litsea: leaves penniveined; anther locules 4, but allowing for species

with anther locules variably 2 or 4; fruits with or without prominent

cupules.

Neolitsea: leaves triveined; anther locules 4; fruits with or without prominent

cupules.

Lindera: leaves penniveined or triveined; and anther locules 2; fruits without

prominent cupules.

New Taxa and Status Changes

Litsea Lam.

1. Litsea aban-gibotii Ng, sp. nov.

(Named for Aban Gibot, b. 1944, plant collector in the Sabah Forest

Department)

Folia infra nitida ob pilos tenues microscopicos (manifesti magnificatione

15x). Fructus in cupulis profundis poculiformibus sedentes. Typus: Aban

Gibot SAN 66744, Sabah, Papar District (holo KEP; iso SAN).

Figure 1

Small to medium-sized trees. Leaves alternate, undersurface with silky shee

n due to closely-packed microscopic hairs (visible at 15x magnification);

blades elliptic or obovate, 9—-13.5 x 44.8 cm, base cuneate to attenuate,

apex acuminate; midrib shallowly impressed above; lateral veins 9-10 pairs;

intercostal venation faint, closely scalariform (mostly closer than 0.2 cm);

petiole 1.5—2 cm long, up to 0.2 cm thick. Inflorescences borne on twigs up

to 1.2 cm diam.; raceme axis highly condensed, 0.2—0.4 cm long; florets 3 or

more per umbellule; peduncles 0.3—0.7 cm long. Fruits ovoid, c. 1.4 x 1.4

cm; hypanthium at first globose, finally forming a deep cup-shaped cupule

covering the basal half to two-thirds of the fruit; pedicel c. 0.4 cm long,

gradually narrowed downwards from the cupule.

Gard. Bull. Singapore 57 (2005)

Distribution: Endemic in Borneo: Sabah, Papar District; Sarawak, Bintulu

and Lundu Districts; Brunei, Belait District.

Specimens examined: Sabah, Papar (SAN 66744); Sarawak, Bintulu (S 18697)

and Lundu (S 76785); Brunei, Belait (Niga NN 242, Niga NN 244).

Notes: The silky sheen on the leaf undersurface, due to microscopic

appressed hairs, is highly characteristic of a small number of Litsea species,

which includes L. chewii Kosterm., L cuprea Merr. and L. staintonii

Kosterm. L. aban-gibotii difters from the others by its closely spaced (mostly

closer than 2 mm) scalariform intercostal veins.

2. Litsea accedens (Blume) Boerl.

Hand. Fl. Ned. Ind. 1, 3 (1900) 145.

Basionym: Tetranthera accedens Blume, Mus. Bot. Lugd. Bat. 1 (1851) 383.

Type: Korthals s.n., Borneo (holo not seen; iso U Acc. No. 188204B).

Notes: A highly variable species in which three varieties may be recognized:

i. var. accedens: intercostal venation reticulate; leaves variable in shape,

from elliptic to oblong-elliptic, obovate and lanceolate. Widespread.

ii. var. kinabaluensis (Kosterm.) Ng: intercostal venation scalariform; leaves

ovate or oblong-elliptic. Endemic to the Kinabalu region of Sabah.

iii. var. oblanceolata (Gamble) Ng: intercostal venation scalariform; leaves

oblanceolate. Widespread.

i. Var. accedens

New synonyms:

Litsea costata (Blume) Boerl., Handl. Fl. Ned. Ind. 3 (1900) 144. Basionym:

Cylicodaphne costata Blume, Mus. Bot. Lugd. Bat. 2 (1856) 13.

Type: Korthals s.n., Borneo (lectotype, here designated, L Acc.

No. 90523333).

Litsea insignis (Blume) Boerl., Handl. Fl. Ned. Ind. 3 (1900) 142. Basionym:

Tetranthera insignis Blume, Mus. Bot. Lugd. Bat. 1 (1851) 382.

Type: Korthals s.n., Sumatra (lectotype, here designated, L Acc.

No. 905233560).

Litsea ochracea (Blume) Boerl., Handl. Fl. Ned. Ind. 3 (1900) 144. Basionym:

Cylicodaphne ochracea Blume, Mus. Bot. Ludg. Bat. 2 (1856) 13.

Type: Korthals s.n., Sumatra (holo not seen; iso BO Acc. No.

1278346, U Acc. No. 38210).

Litsea, Lindera and Neolitsea in Borneo 223

Figure 1. Litsea aban-gibotii Ng. A, fruiting (young) leafy twig; B, detail of venation on lower

leaf surface; C, young fruit; D, older fruit. (A—B from SAN 66744, C from S 76785, D from

Niga NN 242.)

224 Gard. Bull. Singapore 57 (2005)

Litsea ridleyi Gamble, Bull. Misc. Inform. Kew (1910) 317. Type: Ridley

5101, Singapore (lectotype, here designated, SING).

Litsea wrayi Gamble, Bull. Misc. Inform. Kew (1910) 319. Type: Wray

4036, Perak, Waterfall Hill (lectotype, here designated, SING).

Litsea pentagona Mertr., Univ. California Publ. Bot. 15 (1929) 81. Type:

Elmer 20666, Sabah, Tawau (holo not seen; iso BO, NY, SING).

ii. Var. kinabaluensis (Kosterm.) Ng, stat. nov.

Basionym: Litsea kinabaluensis Kosterm., Reinwardtia 7 (1969) 506. Type:

Clemens 33053, Sabah, Gunung Kinabalu (holo BO).

ii. Var. oblanceolata (Gamble) Ng, stat. nov.

Basionym: Litsea oblanceolata Gamble, Bull. Misc. Inform. Kew (1910) 362.

Type: King’s Collector 2020, Perak, Larut (lectotype, here designated. K).

New synonyms:

Litsea ochracea (Blume) Boerl. var. oblanceolata (Gamble) Kochummen,

Tree Flora Malaya 4 (1989) 163.

Litsea perakensis Gamble, Bull. Misc. Inform. Kew (1910) 359. Type: King’s

Collector 5114, Perak, Larut (lectotype, here designated, K).

3. Litsea andreana Ng, sp. nov.

(Named for André Joseph Guillaume Henri Kostermans, 1907-1994, forest

botanist in Bogor, Indonesia)

Folia magna obovata ad oblanceolata infra tomentosa 29-50 cm longa 17-

19 cm lata basi subcordata. Racemi cauliflori 0.4—-1.2 cm longi. Typus: Ilias

Paie § 39032, Sarawak, Niah (holo SAR; iso KEP).

Figure 2

Small to medium-sized tree. Leaves alternate, densely tomentose below;

blades broadly obovate to oblanceolate, 29—50 x 17-19 cm, base subcordate,

apex rounded; midrib broad and raised to shallowly impressed above; lateral

veins 15—24 pairs; intercostal venation scalariform, prominent below; petiole

1—1.5 cm long, 0.6—0.9 cm thick. Inflorescences borne on the branches and

trunk; raceme axis 0.4—1.2 cm long, c. 0.5 cm thick; florets c. 7 per umbellule:

anthers 4locular (.$ 39032); peduncles c. 0.6 cm long. Fruit not seen.

Distribution: Endemic in Borneo: Sarawak, Semengoh FR; Kalimantan,

Gunung Bentuang and Sampit.

Specimens examined: Sarawak, Semengoh FR S 39302; Kalimantan, G.

Bentuang, Burley et al. 2858 and Sampit Kostermans 8057.

225

Litsea, Lindera and Neolitsea in Borneo

Beet {)

part ery

ayaa

+. \ wy,

Figure 2. Litsea andreana Ng var. andreana (A-F): A, leafy twig; B, detail of venation and

indumentum on lower leaf surface: C, cauliflorous inflorescence; D, involucral bud; E, umbellule

of florets within an involucral bud; F, stamen, staminode and glands; var. dewolii Ng: G, young

infructescence. (A-F from S 39032, G from SAN 142976.)

Gard. Bull. Singapore 57 (2005)

Var. dewolii Ng, var. nov.

(Named for Dewol Sundaling, b. 1950, plant collector in the Sabah Forest

Department)

A varietate typica foliis multo maioribus ad 100 cm longis 35 cm latis,

racemis longioribus c. 5.5 cm longis ramis majoribus insidentes differt. Typus:

Dewol SAN 142976, Sabah, Danum Valley (holo SAN; iso KEP).

Leaves to c. 100 x 35 cm. Inflorescences borne on large branches; the

raceme axis c. 5.5 cm long and c. 0.9 cm thick; umbellules and florets not

seen. Fruits with the basal three quarters enclosed in a cup-like cupule

when fresh, completely shrunken into the cupule when dried; cupule densely

tomentose, c. 2.5 cm wide and c. 2 cm deep, non-pedicellate when fresh,

basally narrowed into a 0.5 cm-long pedicel when dried.

Distribution: Endemic in Borneo: Sabah.

Notes: Known only from the type specimen.

4. Litsea cordata (Jack) Hook.f.

Fl. Brit. Ind. 5 (1886) 177. Basionym: Tetranthera cordata Jack, Malay.

Misc. ti (1822) 34. Type: unknown (Jack’s description).

New synonym: Litsea elmeri Merr., Univ. California Publ. Bot. 5 (1929) 82.

Type: Elmer 21221, Sabah, near Tawau (holo not seen; iso SING).

Notes: This is a well-known species and Jack’s name has been consistently

applied even though the type cannot be located. On the Singapore sheet of

Elmer 21221, (type of Litsea elmeri Merr.) there is a determination slip by

Kostermans re-determining the specimen as L. cordata. I agree with

Kostermans, but the reduction does not seem to have been published.

5. Litsea costalis (Nees) Kosterm.

Reinwardtia 7 (1969)501. Basionym: Alseodapne costalis Nees in Wallich,

Pl. As. Rar. 2 (1831) 72. Type: Wallich 2594B, Singapore (holo K; iso BO)

Var. nidularis (Gamble) Ng, stat. nov.

Basionym: L. nidularis Gamble, Bull. Misc. Inform. Kew (1910) 365. Type:

King’s Collector 6883, Perak, Larut (lectotype, here designated, K).

Notes: Litsea costalis var. nidularis differs from the typical variety in having

smaller, narrower leaves and less prominent venation, but there are

intermediates, and at least one specimen from Brunei (Dransfield JD 7249)

Litsea, Lindera and Neolitsea in Borneo 229

bears leaves of both types.

6. Litsea crassifolia (Blume) Boerl.

Hand. Fl. Ned. Ind. 3 (1900) 143.

Basionym: Tetranthera crassifolia Blume, Mus. Bot. Lugd. Bat. 1 (1851)

386. Type: sin. coll., s. n., Java (holo not seen, iso U Acc. No. 261746B).

New synonym: Litsea lithocarpoides Kosterm., Reinwardtia 7 (1968) 348.

Type: Clemens 33823, Sabah, Gunung Kinabalu (holo L; iso BO, K).

Notes: The isotype of Litsea crassifolia (U, viewed on the L database

website) consists of one detached leaf with no number and no locality.

However, the name has been consistently applied in the region and there

is no reason to dispute its application. Litsea lithocarpoides is merely a

high elevation form of L. crassifolia, which is already known to occupy a

broad elevational range, from sea level white sands and peat swamps to

high mountains where soil conditions are similarly sandy or peaty.

7. Litsea cubeba (Lour.) Pers.

Syn. Pl. 2 (1807) 4. Basionym: Laurus cubeba Lour., Fl. Cochin. (1790)

252. Type: not seen.

Synonym: Litsea citrata Blume, Bijdr. Fl. Ned. Ind. 2e (1826) 565 (reduced

by Kostermans in Reinwardtia 10 (1988) 465).

New synonyms:

Lindera pipericarpa auct non (Miq.) Boerl.: Gamble, J. As. Soc. Bengal 75,

2 (1912) 198; Ridley, Flora Malay Pen. 3 (1924) 135; Burkill, Dict.

Econ. Prod. Malay Pen. 2 (1936) 1371; Kochummen, Tree Flora of

Malaya 4 (1989) 147.

Lindera oxyphylla (Nees) Hook.f., Fl. Brit. Ind. (1886) 183. Basionym:

Daphniphyllum oxyphyllum Nees in Wallich, Pl. As. Rar. 2 (1831)

63. Type: Porter s.n., Penang (Wallich Cat. 2547, not seen).

Lindera reticulosa Kosterm., Reinwardtia 9 (1974) 102. Type: FRI 5016,

Pahang, Fraser’s Hill (holo K; iso K, KEP).

Figure 3

Notes: Litsea cubeba is geographically a very widespread species, ranging

from Nepal and Sikkim to Japan and Java.

While reviewing specimens from Borneo and the Malay Peninsula,

it struck me that the Bornean specimens identified as Litsea cubeba by

Kostermans resembled the Peninsular specimens named variously as

Lindera pipericarpa (Miq.) Boerl, Lindera oxyphylla and Lindera reticulosa.

Similarity in the leaves, fruits and female florets indicated that they are

one species in spite of specimens having been placed in two different

220 Gard. Bull. Singapore 57 (2005)

genera due to difference in anther locule number.

The mystery deepened when I found that Kostermans (1970) had

transferred Lindera pipericarpa to Litsea pipericarpa (Miq.) Kosterm.

Kostermans suggested that the earlier authors had not examined the anthers.

The lack of a cupule in the fruit would have suggested Lindera, but there

is another possibility, that the flowers examined by the earlier authors did

have two anther locules while those examined by Kostermans had four,

making Litsea pipericarpa a species with variably 2 and 4 anther lobes.

I examined the type of Lindera pipericarpa (now Litsea pipericarpa,

basionym Polyadenia pipericarpa Miq., Fl. Ind. Bat. 1, 1 (1858) 962) in

Bogor: Teysmann, HB 2214, Sumatra; (=BO Acc. No. 1277851) which bears

Kosterman’s annotations and found that Gamble (1912), Ridley (1924),

Burkill (1936) and Kochummen (1989) were all mistaken. Lindera

pipericarpa has leaves resembling Anisoptera (Dipterocarpaceae) in the

undersurface that is prominently reticulate-veined, finely tomentose and

tinged yellow. It is confined to Sumatra. Litsea cubeba, in contrast, has

leaves glabrous and glaucous below.

I have transferred all the Malay Peninsula specimens previously

identified as Lindera pipericarpa to Litsea cubeba on the grounds that the

anther locule number in Litsea cubeba is not a species-defining character,

but a variable character expressed differently in different parts of the

geographical range of the species. While the Peninsular plants all appear

to be 2-locular, the Bornean specimens may be 2-locular (e.g. SA 1284, S$

55827), 4-locular (e.g. SAN 114245) or have both conditions within the

same floret (e.g. SAN 114166 and TK 1122). In the population of Litsea

cubeba in Bario, Sarawak, that has hermaphrodite flowers, I found stamens

with 2 or 4 anther locules within the same flower, together with abnormal

stamens with 1 or 3 locules.

I thought that Litsea cubeba with 4-locular anthers might exist as a

rare condition in the Peninsula because of two specimens, Ridley 11390

(Taiping Hills, Perak), and Ridley 13780 (Telom, Pahang), placed by Gamble

(1912) in Litsea citrata Blume. These were reported to have ‘quadrate

anthers’. This reference was overlooked by Ridley (1924) but cited in

synonymy by Burkill (1936) under Lindera pipericarpa, implying that Burkill

disagreed with Gamble over the number of anther locules. Duplicates of

the two specimens were located in SING and examined by Julia Sang, who

found the anthers to be 2-locular. I nevertheless think that the specimens

examined by Gamble were 4-locular, otherwise he would not have said so.

In any case, Litsea citrata Blume has itself been reduced to Litsea cubeba

by Kostermans (1988).

In summary, both Litsea pipericarpa and Litsea cubeba have

synonyms in Lindera and the reason, I believe, is variation in the number

Litsea, Lindera and Neolitsea in Borneo 229

Figure 3. Litsea cubeba (Lout.) Pers. A, fruiting leafy twig; B, detail of venation and indumentum

on lower leaf surface; C, infructescence; D, female flower; E, hermaprodite flower with stamens

variably 2-locular and 4-locular. (A—C from SAN 60328, D from SAN 24022, E from SAN

1141606.)

230 Gard. Bull. Singapore 57 (2005)

of anther locules, which the authors had not been prepared for.

8. Litsea elliptica Blume

Bijdr. Fl. Ned. Ind. 1le (1826) 563. Type: s. coll., s. n., Java, Mt Salak (holo

L Acc. No. 905233282; iso BO Acc. No. 1274910).

New synonym:

Litsea pruriens Kosterm., Reinwardtia 8 (1970) 105. Type: Kostermans

12675, Kalimantan, W. Kutei, Belajan River (holo BO; iso SING).

Notes:

I can find no consistent difference between L. elliptica and L. pruriens.

Litsea elliptica has small or undeveloped cupules and fruiting specimens

are often mistaken for Lindera but the males consistently have 4-locular

anthers.

9. Litsea firma (Blume) Hook. f.

Fl. Brit. Ind. 5 (1886) 162. Basionym: Tetranthera firma Blume, Mus. Bot.

Lugd. Bat. 1 (1851) 381. Type: Korthals, s.n., Borneo (lectotype here

designcted BO Acc. No. 1245463).

New synonyms:

Litsea cylindrocarpa Gamble, Bull. Misc. Inform. Kew (1910) 318. Type:

King’s Collector 6673, Perak, Larut (lectotype, here designated, SING).

Litsea turfosa Kosterm., Reinwardtia 7 (1968) 353. Type: Lajanjah SAN

44550, Sabah, Beaufort (holo K; iso SAN, SAR, SING).

Notes: Litsea turfosa and L. cylindrocarpa are linked by intermediates to

L. firma (Blume) Hook. f., forming a single widespread and common

species.

10. Litsea fulva (Blume) F. Vill.

Nov. App. FI. Filip. (1880) 181. Basionym: Tetranthera fulva Blume, Mus.

Bot. Lugd. Bat. 1 (1851) 377. Synt Types: s. coll., s. n. Java and Sumatra

(according to Blume), not seen.)

Notes: The typical variety of this species ranges from Sumatra to Java,

Borneo and the Philippines, but the new variety described below is endemic

to the Kinabalu massif in Sabah.

Var. corneri Ng, var. nov.

(Named for E.J.H. Corner, 1906-1996)

A varietate typica foliis ovatis vel ellipticis aut subobovatis (vs. obovatorum)

Litsea, Lindera and Neolitsea in Borneo Dat

umbellulis fructibusque sessilibus (vs. pedunculatorum) differt. Typus: Chew

& Corner RSNB 4473, Sabah, Kinabalu, Bembangan River at c. 1700 m

(holo SING; iso SAN).

Leaves ovate, elliptic or sub-obovate,; 3-11 x 1.5—4.4 cm, below hairy all

over or on the veins, base cuneate to rounded, apex acuminate; midrib

above sunken into a narrow groove; lateral veins 8—10 pairs; intercostal

veins scalariform; petiole 0.4—0.5 cm long. Flowering on the twigs; raceme

axis highly condensed 0.1—0.2 cm long; florets c. 3 per umbellule; peduncle

practically absent (umbellules practically sessile). Fruits ellipsoid, c. 1 x 0.7

cm; cupule forming a cup c. 0.6 cm wide, c. 0.5 cm deep, non-pedicellate.

Distribution: Endemic in Borneo: Sabah, Gunung Kinabalu at 1500-2600 m.

Specimens examined: Sabah Chew & Corner RSNB 151, Chew & Corner

RSNB 4473, Clemens 29564, Clemens 31052, Clemens 32506, Clemens 33811,

Clemens 35117, Sugau JBS 128 and SAN 117234.

11. Litsea garciae Vidal

Rev. Pl. Vasc. Filip. (1886) 228. Type: Vidal 861, Philippines (holo not

seen; iso L).

New synonyms:

Litsea sebifera auct. non (Willd.) Pers. in Blume, Bijdr. Fl. Ned. Ind. (1825)

560; Gamble, J. As. Soc. Beng; 75 (1912) 176; Ridley, Fl. Malay

Peninsula 3 (1924) 128.

Litsea glutinosa auct. non (Lour.) C.B.Rob.: Burkill, Economic Products

Malay Peninsula 2 (1936) 1376.

Litsea aurea Kosterm., Reinwardtia 8 (1970) 86. Type: bb 8697, Sumatra,

Palembang (holo BO).

Litsea robusta auct. non Blume: Kochummen, Tree Flora Malaya 4 (1989)

163, pro parte.

Notes: This species has had a very confused taxonomic history although it

is clearly distinguished by its uniquely asymmetric leaf shape: oblong-ovate

to lanceolate, expanded more on one side than the other in the lower half

of the leaf. No other Litsea species has a leaf like this. Furthermore, the

apical buds, when resting are covered by large overlapping silvery-silky

reduced leaves. The fruits are among the largest in the genus, depressed

globose, up to 4 cm diameter when fresh. An edible variety is grown in

Sarawak (Bidayuh name engkalak), Brunei (Brunei Malay name

pengalaban), and the Philippines and Java. In Sabah and Peninsular

Malaysia, the species is represented by big forest trees (up to 50 m tall),

232 Gard. Bull. Singapore 57 (2005)

undoubtedly wild and indigenous, but the fruits are not eaten.

This species was first described by Blume (1825) as ‘Litsea sebifera

Pers.’ and this name was taken up by Gamble (1912) and Ridley (1924).

However, Litsea sebifera Pers. has turned out to be a different species. The

oldest valid name for this species is Litsea garciae Vidal. Gamble’s

description: ‘innovations silvery-silky’ and leaves ‘often unequal at base’

undoubtedly fits Litsea garciae. Ridley’s comment that trees are ‘occasionally

planted by the Javanese’ also points to Litsea garciae, because no other

Litsea species is cultivated in home gardens. Burkill (1936) redetermined

this species as ‘Litsea glutinosa’, but this was a mistake; the true Litsea

glutinosa (Lour.) C.B.Rob. has different leaves and much smaller fruits.

Kochummen (1989) did not recognise Litsea garciae in the Tree Flora of

Malaya because he confused it with Litsea robusta, which 1s similar in

inflorescence and fruit but different in leaf shape. On re-sorting

Kochummen’s specimens, about half were found to belong to L. garciae

and half to L. robusta. The latter is rare in Borneo.

12. Litsea globularia Ng, sp. nov.

(Latin, globularis=globe, referring to the fruit)

Folia opposita infra tomentosa. Fructus globosi ad 2.4 cm diam. cupula

patelliformi circulari ad & mm lata pedunculo 2—12 mm longo pedicello 2—5

mm longo articulato insidenti. Typus: Primack S 42435, Sarawak, Gunung

Mulu (holo KEP; iso A, K, L, SAR).

Figure 4

Medium-sized tree. Leaves opposite, chartaceous, tomentose below

especially on the midrib; blades obovate, 8-23 x (3—)5(-8) cm, base rounded,

apex acuminate; midrib sunken above and the groove sometimes filled

with hairs; lateral veins 9-14 pairs; intercostal venation laxly reticulate;

petiole 0.5—-1 cm long. Inflorescences borne on twigs; raceme axis highly

condensed, c. 0.1 cm long; florets 5 per umbellule; peduncles 0.2—1.2 cm

long. Fruits globose, to 2.4 cm diam.; cupule forming a circular plate up to

0.8 cm wide; pedicel thick, 0.2—-0.5 cm long.

Distribution: Endemic in Borneo: Sabah, Sarawak, Brunei and Kalimantan.

Selected specimens examined: Sabah (SAN 16397, SAN 28910, SAN 31923,

SAN 42841, SAN 50511, SAN 65214, SAN 86964 and SAN 128632), Sarawak

(S 4025, S 15093, S 23019, S 33214, S 42435, S 50511 and S 62309); Brunei

(Kirkup DK 942, Coode MC 6340); Kalimantan (Kostermans 10439).

Litsea, Lindera and Neolitsea in Borneo 235

13. Litsea grandis (Wall. ex Nees) Hook. f.

Fl. Brit. Ind. 5 (1886) 162. Basionym: Tetranthera grandis Wall. ex Nees in

Wallich, Pl. As. Rar. 2 (1831) 162 Type: ‘Wallich 2552’, Penang (holo K).

Notes: This species is now reorganized to have three varieties, linked by

intermediates:

1.. var. paludosa: leaves ovate or elliptic; glaucous below

li. var. grandis: leaves oblong-elliptic, elliptic, or round; not glaucous below;

indumentum on twigs and leaf undersurface mid-brown

lil. var. rufo-fusca: leaves ovate; not glaucous below; indumentum dark

blackish red-brown

i. Var. paludosa (Kosterm.) Ng, stat. nov.

Basionym: Litsea paludosa Kosterm., Reinwardtia 8 (1970) 103. Type:

Anderson § 14520, Sarawak, Simanggang (holo BO; iso SAR, SING).

Notes: This variety includes Litsea grandis in Anderson (1963).

ii. Var. rufofusca (Kosterm) Ng, stat. nov.

Basionym: L. rufofusca Kosterm., Reinwardtia 7 (1968) 352, lc. 8 (1970)

109. Type: bb. 33044, Kalimantan, Sampit area (holo L; iso BO).

14. Litsea jaswirti Ng, sp. nov.

(Named for Jaswir Singh, b. 1938, plant collector of the Sabah Forest

Department)

Folia elliptica basi attenuata apice acuminato. Fructus ovoidei ad obovoidei

ad 2.3 cm longi I cm diam. plerumque solitarii cupula non profunda

prominente stipitata stipite prominenti insidentes. Typus: Jaswir Singh SAN

28253, Sabah, Kundasang (holo KEP; iso SAN).

Figure 5

Small and medium-sized tree. Leaves alternate; blade elliptic 7—13.5 x 2—

4.5 cm, glabrous, chartaceous, base attenuate, apex acuminate, the upper

and lower halves almost mirror images of each other in size and shape;

midrib above flat, striate or shallowly impressed; lateral veins 5—7 pairs;

intercostal veins reticulate; petiole 0.8—1.5 cm long. Flowering on the twigs;

raceme axis 0.4—1.5 cm; florets 3—5 per umbellule; peduncles 0.3—1 cm

long. Fruit narrow, ovoid, ellipsoid or obovoid, to 2.3 x 1.0 cm; usually 1,

rarely 2—3 developed per umbellule; cupule a shallow cup to 0.5 cm wide,

pedicel 0.5—1.3 cm long.

234 Gard. Bull. Singapore 57 (2005)

Distribution: Endemic in Borneo: Sabah, Gunung Kinabalu.

Specimens examined: Sabah, Kinabalu (Carr SFN 27216, Chew RSNB 525,

Clemens 26367, SAN A 4532, SAN 15272, SAN 28253, SAN 28294, SAN

28917).

15. Litsea lancifolia (Roxb. ex Wall.) Hook. f.

Fl. Brit. Ind. 5 (1886) 159. Basionym: Tetranthera lancifolia Roxb. ex Wall.

ex Nees in Wallich, Pl. As. Rar. 2 (1831) 65. Type: ‘Wallich 2532’, India,

Silhet (holo K).

Notes: This is one of the four species in which the anther locules may be 4

or 2. The species varies greatly in leaf size, leaf shape, midrib prominence

and peduncle length, and has a wide distribution from India to Yunnan,

Sumatra, Peninsular Malaysia and Borneo. In this treatment, the Bornean

population is divided into four varieties, all linked by intermediates. The

species is characterised by leaves opposite to sub-opposite, intercostal

venation reticulate to sub-scalariform, fruits ellipsoid or ovoid, small (to 8

x 7 mm), with hypanthium undeveloped or forming a small cup or cone,

pedicellate or non-pedicellate.

The anthers are 4-locular except in var. i/iaspaiei where the number

is variable, 4 or (rarely) 2 locules.

The four varieties are distinguished as follows:

i. var. iliaspaiet: leaves broad, but small and shorter than 8 cm.

i. var. lancifolia: leaves broad, longer than 8 cm, with midrib sunken above.

iii. var. grandifolia: leaves broad, longer than 8 cm, with midrib prominent

above.

iv. var. rheophytica: leaves linear, willow-like.

i. Var. iliaspaiei Ng, var. nov.

(Named for Ilias Paie, 1936-1986, plant collector in the Sarawak Forest

Department)

A varietate typica foliis obovatis ad ellipticis parvis 3.5—7(—8.5) cm longis

1.3—3.2 cm latis differt. Typus: Ilias Paie S 40749, Sarawak, Bukit Sempadai,

(holo KEP; iso SAR).

Small trees to 10 m tall. Leaves opposite, glabrous to finely hairy below:

blade obovate to elliptic, small, 3.5—7(-8.5) x 1.3—3.2 cm, base cuneate,

Litsea, Lindera and Neolitsea in Borneo 235

CL ty

“a

(L5

mje

CL,

Figure 4. Litsea globularia Ng. A, fruiting (young) leafy twig; B, detail of venation and

indumentum on lower leaf surface; C, young fruit; D, umbellule of florets; E, female flower.

(A-B from S$ 42435, C from S 39856, D-E from S 47538.)

o®)

Gard. Bull. Singapore 57 (2005)

apex acuminate; midrib above sunken; lateral veins 5—10 pairs; intercostal

veins reticulate; petioles 0.3—0.4 cm long. Flowering on the twigs, raceme

axis highly condensed, 0.1—0.3 cm long; florets 1-6 per umbellule; peduncles

0.1—0.6 cm long. Fruits ovoid, c. 0.7 x 0.6 cm; cupule forming a shallow

plate c. 0.3 cm wide, on a short c. 0.1 cm pedicel.

The anthers may be 4locular (S 40749, SAN 1769 and SAN 88274)

or 2-locular ($ 52326). In the case of the specimen with two anther locules,

the number of florets was reduced to one, technically making this specimen

an Iteadaphne, but in all other respects, it conforms to L. lancifolia var.

iliaspaiel.

Distribution: Endemic in Borneo: Sabah and Sarawak. In hill and lower

montane forests at 950—1600 m elevation.

Selected specimens examined: Sabah, (SAN 31441, SAN 119768, SAN A

1769 and Sugau JBS 10), Sarawak (S 21788, S 33758, S 40749, S 47656 and

S 36855).

ii. Var. lancifolia

New synonyms:

Litsea varians (Blume) Boerl., l.c. (1900) 143. Basionym: Tetranthera varians

Blume, Mus. Bot. Lugd. Bat. 1 (1851) 376. Type: Korthals s. n.,

Borneo (holo not seen; iso BO Acc. No. 1249188).

ii. Var. grandifolia (Stapf) Ng, stat. nov.

Basionym: Lindera grandifolia Stapf, Trans. Linn. Soc. Bot. Ser. 2, 4 (1894)

220. Type: Haviland 1334. Sabah, Kinabalu (holo K; iso SAR).

New synonyms:

Litsea amaroideocarpa Kosterm., Reinwardtia 7 (1968) 345.

Litsea oppositifolia Gibbs, J. Linn. Soc. Bot. 42 (1914) 130. Type: Gibbs

3136, Sabah, Tenom (holo BM).

Notes: Stapf (1894) based Lindera grandifolia on a fruiting specimen.

Kostermans transferred Lindera grandifolia to Litsea but had to give it the

new epithet amaroideocarpa, because Litsea grandifolia had already been

used for another species (Litsea grandifolia Lecomte in Nouv. Arch. Mus.

Se Ser 5 (1913) 87). However, under its new status as a variety of L.

lancifolia, Stapf’s original epithet is hereby restored.

iv. Var. rheophytica (Kosterm.) Ng, stat. nov.

Basionym: Litsea rheophytica Kosterm., Reinwardtia 7 (1968) 350. Type:

Chai P.P.K. S 18951, Sarawak, Rejang River, Pelagus Falls (holo L; iso

Litsea, Lindera and Neolitsea in Borneo Rew |

Figure 5. Litsea jaswirii Ng. A, flowering and fruiting leafy twig; B, detail of venation and

indumentum on lower leaf surface: C, raceme of involucral buds; D, involucral bud: E, umbellule

of florets within an involucral bud; F, female flower. (A—F from SAWN 28253.)

238 Gard. Bull. Singapore 57 (2005)

BO, KEP, SAN, SAR).

16. Litsea machilifolia Gamble

Bull. Misc. Inform. Kew (1910) 320. Type: Curtis 795, Penang (lectotype,

here designated, SING);

New synonyms:

Litsea teysmannii Gamble, Bull. Misc. Inform. Kew (1910) 319. Type: Ridley

5845, Malacca (lectotype, here designated, SING).

Litsea panamonja auct. non Hook. f.: Gamble, J. As. Soc. Beng. 75, 2 (1912)

172; Ridley, Flora Malay Pen. 3 (1924) 127; Kochummen, Tree Flora

Malaya 4 (1989)162.

Litsea sp. ‘1’ and Litsea sp. ‘2’ in Kochummen, Tree Flora Malaya 4 (1989)

166.

Notes: This widespread and common species is easily distinguished by leaf

morphology, relatively long racemes, and relatively large fruits. The cupule

is a fleshy cup on a pedicel of variable length (O—3 cm long). This extreme

variation in fruit pedicel length has been a source of great confusion.

Gamble (1910) placed specimens with long pedicels in L. machilifolia and

those without pedicels in L. teysmannii. Kochummen (1989) recognised L.

teysmannii but associated it with swampy habitats. For the non-swamp

specimens, he placed those with 2—3 cm-long pedicels in L. machilifolia;

one specimen (FRI 26004) with a 0.7 cm-long pedicel into his Litsea sp. 2;

and specimens without pedicels into Litsea sp. 1 (FMS 6844, FRI 2045, FRI

3806, FRI 15688, FRI 19240, FRI 26003, KEP 110331 and SFN 30880).

Non-fruiting specimens were placed arbitrarily. The impossibility of sorting

out non-fruiting specimens indicates that the whole complex belongs to

one species. In Borneo, the fruiting pedicel is similarly variable although

not as extreme as in the Malay Peninsula.

Two Malayan specimens, from Perak (Curtis 2694) and Malacca

(Derry 2002), had previously been determined by Gamble (1912) as Litsea

panamonyja, a species otherwise known only in India and Myanmar. I have

redetermined these specimens as Litsea machilifolia, thereby excluding

Litsea panamonja from the Malay Peninsula.

17. Litsea magnifica (Miq.) F. Villar

in Blanco, Fl. Filip. 3, Nov. App. (1880) 181. Basionym: Lepidadenia

magnifica Mig. Fl. Ind. Bat. 1 (1858) 936. Type: Teysmann HB 1011,

Sumatra, Batang Baroes (holo not seen; iso U Acc. No. 38224).

New synonym: Litsea johorensis Gamble, Bull. Misc. Inform. Kew (1910)

315. Type: Ridley 13479, Johore (lectotype, here designated, SING).

Litsea, Lindera and Neolitsea in Borneo 239

Notes: Litsea magnifica and L. johorensis are cauliflorous and similar in

their large obovate leaves and small fruits, c. 1 x 0.7 cm. Kochummen

(1989) separated them by the degree of swelling of the petiole base. There

is also variation in the development of the cupule in fruit, but the number

of herbarium collections is few and the evidence insufficient to support

more than one species.

18. Litsea sessiliflora Hook.f.

Fl. Brit. Ind. 5 (1886) 160. Type: Maingay, KD 1511, Penang (holo K).

This is one of the four species in which the number of anther locules may

be 4 or 2. The variation is found within the new variety, var. othmanii.

i. Var. sessiliflora

New synonym: Litsea sandakanensis Merr., J. Str. Br. Roy. As. Soc. 85

(1922) 194. Type: Ramos 1507. Sabah (holo not seen; iso L Acc. No.

923246540).

Var. othmanii Ng, var. nov.

(Named for Haji Othman Ismawi, b. 1940, plant collector in the Sarawak

Forest Department)

A varietate typica foliis glabris vel infra minute adpresse pubescentibus (vs.

tomentosorum) 1.5—3(—4.5) cm latis (vs. 3—9.5 cm) differt. Typus: Othman

S 57189, Sarawak, Lundu, Gunung Putin (holo KEP; iso SAN, SAR).

Small tree. Leaves opposite, glabrous to finely appressed hairy below; blade

mostly narrow-elliptic (rarely narrow-ovate or narrow-obovate), (6—)7—14.5

x 15-3 cm, base cuneate, apex acuminate; midrib sunken above; lateral

veins 6-10 pairs; intercostal veins reticulate; petiole 0.3—0.8 cm long.

Flowering on the twigs, raceme axis 0.1—0.5 cm long; florets 1—3 per

umbellule; anther locules 2 (Wong WKM 1252) or 4 (S 2992, S 44528);

peduncles 0.1—0.2 cm long. Fruits ellipsoid, c. 0.8 x 0.6 cm; cupule a small

cup 0.3—0.4 cm wide, c. 0.2 cm deep, non-pedicellate.

Distribution: Endemic in Borneo: Sabah, Sarawak, Brunei and Kalimantan.

Lowland to lower montane forests at 1500 m elevation.

Selected specimens examined: Sabah (Chew RSNB 506, SAN 1156, SAN

29331, SAN 32930, SAN 46653, SAN 54383, SAN 78015, SAN 84387, SAN

91434 and SAN 120142); Sarawak (S 1145, S 13271, S 21517, S 29992, S$

35172, S 41851, S 44528, S 51475 S 83468); Brunei (BRUN 432, BRUN

1143, BRUN 15636, Wong WKM 785 and WKM 1252, Kalimantan (Church

240 Gard. Bull. Singapore 57 (2005)

582 and Kostermans 21440).

Notes: The narrow, elliptic leaves resemble Litsea subumbelliflora (Blume)

Ng which, however, has an attenuate leaf base. Also, the fruits of L.

subumbelliflora have distinct peduncles (0.2—0.6 cm long) and pedicels

(0.1—0.5 cm long).

When I first examined the specimen Wong WKM 1252 and found

uniflorous umbellules with 2-locular anthers, I thought I had a new species

of [teadaphne and a new generic record for Borneo. However, a single

male specimen does not make a species. I eventually found the females in

Litsea filed together with male specimens having 4-locular anthers and 2—3

florets per umbellule!

19. Litsea suboppositifolia Ng, sp. nov.

(Named for the subopposite leaves)

Folia subopposita coriacea ovata ad elliptico-oblonga plerumque plus quam

4 cm lata. Typus: Sugau JBS 116, Sabah, Tambunan (holo KEP; iso SAN).

Figure 6

Small tree. Leaves subopposite, coriaceous, glabrous, slightly glaucous

below; blade ovate to oblong-elliptic, 8.5—20 x 3.5—8 cm, but mostly broader

than 4 cm, base cuneate, apex acute to acuminate; midrib broad and flat,

impressed, or prominent above; lateral veins 6—12 pairs; intercostal veins

reticulate to vaguely scalariform; petiole 1—2.5 cm long, 0.2—0.4 cm thick

(relatively thick and stout). Inflorescences borne on the twigs; raceme axis

0.5—2.5 cm long; florets c. 3 per umbellule; peduncles 0.3—0.8 cm long.

Fruits ellipsoid, to 2 x 1.5 cm; cupule forming a cup to 1.5 cm wide and

0.5—0.8 cm deep; pedicel short and fleshy, 0.2—0.3 cm long.

Distribution: Endemic in Borneo: Sabah and Sarawak. Lower and upper

montane forest at 1100—3000 m elevation.

Selected specimens examined: Sabah (SAN 29195, SAN 38327, SAN 38349,

SAN 46569, SAN 95242, SAN 123342, SAN 123387, SAN 123532, Sugau

JBS 116); Sarawak (S 38135 and S 50427).

20. Litsea subumbelliflora (Blume) Ng, comb. nov.

Basionym: Laurus subumbelliflora Blume, Bijdr. Fl. Ned. Ind. 1le (1826)

554. Type: Blume s.n., Java (lectotype, here designated, L Acc. No.

951212095)

241

Litsea, Lindera and Neolitsea in Borneo

A i 3 cm

}

1 mm

Figure 6. Litsea suboppositifolia Ng. A, fruiting (young) leafy twig; B, part of infructescence

with an older fruit; C, umbellule of 3 fiorets (flowers) within an involucral bud; D, longitudinal

section of flower; E, a stamen with 4-locular anther. (A from J.B. Sugau JBS 116 from Carr

SFN 27101, C-E from Carr SFN 27724).

242 Gard. Bull. Singapore 57 (2005)

New synonyms:

Laurus pauciflora Blume, Fl. Ned. Ind. 1le (1826) 555. Type: Blume s.n.,

Java (holo L Acc. No. 905230213).

Iteadaphne confusa Blume, Mus. Bot. Lugd. Bat. 1 (1851) 365, nomen illeg.

Lindera subumbelliflora (Blume) Kosterm., J. Sci. Res. Indon. 1 (1952)

£27.

Litsea lanceolata (Blume) Kosterm., Reinwardtia 7 (1968) 348. Basionym:

Aperula lanceolata Blume lI.c. (1851) 367. Type: Blume s.n., Java

(holo not seen; iso U Acc. No. 190430B).

Notes: This is one of the four species in which male specimens with two

anther locules were placed in Lindera or Iteadaphne and those with 4

anther locules in Litsea. In Borneo, both conditions occur, but in the Malay

Peninsula only the 2-locular condition is found. The species includes Laurus

subumbelliflora Blume and Laurus pauciflora Blume that Blume (1851)

combined and renamed /teadaphne confusa Blume when he created the

genus /teadaphne. This procedure is incorrect. The situation was rectified

with the choice of the epithet subumbelliflora by Kostermans (1952) when

he reduced Iteadaphne confusa Blume to Lindera subumbelliflora (Blume)

Kosterm. With the merger of Lindera subumbelliflora with Litsea lanceolata,

the species becomes Litsea subumbelliflora.

21. Litsea umbellata (Lour.) Merr.

Philip. J. Sc. Bot. 14 (1919) 242. Basionym: Hexanthus umbellatus Lour.,

Fl. Cochinch. (1790) 195. Type: not seen.

New synonym: Litsea gracilis Gamble, Bull. Misc. Inform. Kew. (1910)

317. Type: Ridley 14603, Perak, Temango (holo not seen; iso SING).

Notes: In his account in the Tree Flora of Malaya, Kochummen (1989)

noted that he had not seen the rare species, Litsea gracilis, known only

from the type. However, the isotype in SING bears a note by him reducing

it to Litsea umbellata. | agree with this reduction.

22. Litsea unita (Blume) Boerl.

Handl. Fl. Ned. Ind. 3 (1900) 145. Basionym: Cylicodaphne unita Blume,

Mus. Bot. Lugd. Bat. 1 (1851) 387. Type: s. coll., s. n., Indonesia (holo L

Acc. No. 90523382).

New synonyms:

Litsea pallidifolia Merr., Univ. California Publ. Bot. 5 (1929) 81. Type:

Elmer 20808, Sabah, near Tawau (holo not seen; iso SING).

Litsea montis-dulit Airy Shaw, Bull. Misc. Inform. Kew (1939) 536. Type:

Synge 1649, Sarawak, Dulit Ridge (holo K; iso BO, SING).

Litsea, Lindera and Neolitsea in Borneo 243

Notes: This species has pale-coloured leaves with few, widely spaced lateral

veins. The isotype of Litsea montis-dulit in SING bears a note by

Kostermans reducing it to Litsea unita but which reduction he did not

publish.

Lindera Thunb.

23. Lindera bibracteata (Blume) Boerl.

Handl. Fl. Ned. Ind. 3 (1900) 146. Basionym: Laurus bibracteata Blume,

Bidr. Fl. Ned. Ind. (1825) 553. Type: Blume s.n., Java, Mt Salak (holo L

Acc. No. 905230346).

New synonyms:

Lindera rufa (Stapf) Gamble, J. As. Soc. Beng. 75, 2 (1912) 199. Basionym:

Lindera caesia (Reinwardt ex Blume) VillarBoerl. var. rufa Stapf,

Trans. Linn. Soc. Bot. 4 (1894) 220. Type: Haviland 1106, Borneo,

Sabah (holo not seen; iso SING, SAR).

Lindera turfosa Kosterm., Reinwardtia 7 (1969) 497. Type: Kostermans

$136, Borneo, Kalimantan, Sampit district (holo BO; iso L).

Notes: This is a distinctive and common species with triveined, ovate leaves

with a long-acuminate apex. There is variation in the degree of hairiness in

the twigs and underside of the leaves, but the variation is continuous.

24. Lindera lucida (Blume) Boerl.

Handl. Fl. Ned. Ind. 3 (1900) 147. Basionym: Litsea lucida Blume, Bijdr.

Fl. Ned. Ind. (1825) 562. Type: Blume s.n., Java (holo L Acc. No. 905230435).

New synonym: Lindera pedicellata Kosterm., Reinwardtia 8 (1970) 83.

Type: Clemens 28522, Borneo, Sabah, Gunung Kinabalu (holo BM)

Neolitsea Merr.

25 Neolitsea cassia (L.) Kosterm.

J. Sci. Res. Indon. 1 (1952) 152; Comm. For. Res. Inst. 57 (1957) 49 and 54.

Basionym: Laurus cassia L., Sp. Pl. (1753) 369. Type: ‘Fl. Zeylanica

No 146’ (BM)

Synonym: Neolitzea zeylanica (Nees) Merr., Philip. J. Sc. 1, Suppl. (1906)

57. Basionym: Litsea zeylanica Nees, Amoen. Bot. Bonn. | (1823)

58, t. 5. Type: Hermann’s Herb: Laurus zeylanica in BM, not seen

New synonyms:

Neolitsea latifolia (Blume) S.Moore, J. Bot. 43, Suppl (1925) 89. Basionym:

Litsea latifolia Blume, Mus. Bot. Lugd. Bat. 24 (1851) 394. Type: s.

coll., s.n. Sumatra (not seen).

244 Gard. Bull. Singapore 57 (2005)

Neolitsea coccinea B.C.Stone, Malaysian Forester 43 (1980) 245. Type: Kiew

RK 732, Gunung Ulu Kali (holo KEP).

Neolitsea sp. 1 in Kochummen, Tree Flora Malaya 4 (1989) 169.

Notes: Nees (1823), in publishing Litsea zeylanica Nees, listed Litsea cassia

L. as a synonym. When Merrill (1906) established the genus Neolitsea, he

transferred Litsea zeylanica (Nees) to Neolitsea zeylanica (Nees) Merr.,

and made it the type species of the new genus. Kostermans (1952) briefly

discussed the history of Neolitsea and made the following comment about

its type species: “As Neolitsea zeylanica (Nees) Merr. is identical with Laurus

cassia L. (p.p.), the type species was renamed in this paper: Neolitsea cassia

(L.) Kosterm.’

Kostermans (1957) later restated the change in a more formal

manner, but an explanation was only provided at the end of this paper,

under the heading “Additions to: A historical survey of Lauraceae’. The

addendum says “Laurus cassia L. is based on no. 146 in his Flora zeylanica,

which in turn is based on Hermann’s specimens. In Hermann’s Herbarium

(British Museum) these represent a mixture of Litsea zeylanica Nees and

the wild Cinnamomum zeylanicum (Trimen in J. Linn. Soc. Bot. 24: 140.

1887).

Litsea latifolia Blume was described in 1851 with three varieties,

var. areolata, vat. caesia and var. punctata in addition to the typical variety.

The species was reduced by Hooker 7. (1886) to Litsea zeylanica Nees. I

have seen a specimen labelled as Litsea latifolia Blume (BO Acc. No.

1278323), possibly the type of L. latifolia var. caesia Blume, and an image

of another, NY 355281, possibly the type of L. latifolia var. punctata Blume

(Herb. Meisner), from the NY website. Both specimens conform to Litsea

zeylanica, now Neolitsea cassia, hence I concur with Hooker f.

Moore (1925) transferred Litsea latifolia Blume to Neolitsea latifolia

(Blume) S.Moore apparently without realizing that Hooker f had already

reduced Litsea latifolia Blume to Litsea zeylanica.

Neolitsea zeylanica, now Neolitsea cassia, is variable in fruit shape

(globose or ellipsoid), fruit size (0.5—1.5 cm diam.) leaf texture (chartaceous

to coriaceous), and cupule development (distinct or indistinct). It covers a

vast geographic range, from the Himalayas to Australia, and from sea

coasts to high mountains. In the Malay Peninsula on Gunong Ulu Kali,

there is a population with coriaceous leaves and small ellipsoid fruits that

was described as Neolitsea coccinea B.C.Stone and which 1s identical to

montane specimens of N. cassia in Borneo. Then there are montane

specimens in the Malay Peninsula with ellipsoid fruits but leaves chartaceous

to coriaceous, which Kochummen (1989) called Neolitsea sp 1, and which

are also part of the N. cassia complex.

Litsea, Lindera and Neolitsea in Borneo ZAS

26. Neolitsea villosa (Blume) Merr.

Philip. J. Sc. Bot. 4 (1909) 261. Basionym: Litsea villosa Blume, Mus. Bot.

Lugd. Bat. 1 (1851) 349. Type: Zippelius s.n., Moluccas, Ambon (holo not

seen; iso L Acc. No. 905234108).

New synonyms: N. mollissima (Gamble) Gamble, Bull. Misc. Inform. Kew

(1911) 172. Basionym: Tetradenia mollissima Gamble, Bull.

Misc.Inform. Kew (1910) 366. Type: Wray 931, Perak, Gunung Batu

Putih (holo K).

Notes: This species differs from the more widespread Neolitsea cassia (L.)

Kosterm. by its densely hairy twigs. I cannot see any difference between N.

mollissima and N. villosa. Also, N. kedahense Gamble, existing as a single

population on Gunong Jerai, Kedah, is probably no more than a local

variant of N. villosa.

Excluded Species

Litsea casearioides Kosterm., Reinwardtia 7 (1968) 346. Type: James Ah

Wing SAN 19047 (holo L; iso SAN, SING, KEP).

This species has to be excluded from Lauraceae (J. Sugau and W.J.J.O. de

Wilde, pers. comm.). Its status will be clarified in a paper being prepared

by de Wilde.

Acknowledgements

I am grateful to the curators of the herbaria at BO, SAN, SAR and SING

for the loan of specimens, to the curators at K and BM for photographs of

type specimens, to Dr. J. F. Veldkamp for the botanical Latin, and to the

Tree Flora of Sabah and Sarawak Project for funding the study. Special

thanks are due to Prof. Jens Rohwer and Dr Henk van der Werff for

kindly reviewing the manuscript and to Joseph Pao for preparing the

botanical illustrations.

References

Anderson, J.A.R. 1963. The flora of the peat swamp forest of Sarawak and

Brunei, including a catalogue of all recorded species of flowering

plants, ferns and fern allies. Gardens’ Bulletin Singapore. 20: 131—228.

Bentham, G. and J.D. Hooker. 1880. Laurineae. Genera Plantarum 3: 146—

165.

246 Gard. Bull. Singapore 57 (2005)

Burkill, [.H. 1936. Litsea. Dictionary of the Economic Products of the Malay

Peninsula. 2: 1375—1379 (pagination according to ed 2, 1966).

Gamble, J. S. 1910. XLVII. New Lauraceae from the Malayan region III.

Bulletin of Miscellaneous Information from Kew. pp. 312-321.

Gamble, J. S. 1912,. Lauraceae. Journal of the Asiatic Society of Bengal.

75: 35-202.

Hooker, J.D.f. 1886. Laurineae. Flora of British India. 5: 116-188.

Kochummen, K.M. 1989. Lauraceae. Tree Flora of Malaya. 4: 98-178.

Kostermans, A.J.G.H. 1952. A historical survey of Lauraceae. Part III.

Journal for Scientific Research in Indonesia. 1: 141-159. 1

Kostermans, A.J.G.H. 1957. Lauraceae. Communications of Forest Research

Institute (Indonesia). 57: 1—64.

Kostermans, A.J.G.H. 1970. Materials for a revision of Lauraceae III.

Reinwardtia. 8: 21—196.

Kostermans, A.J.G.H. 1988. Materials for a revision of Lauraceae V.

Reinwardtia. 10: 439-469.

Merrill, E.D. 1906. The flora of Lamao forest reserve. Philippine Journal

of Science 1, suppl: 1-143.

Moore S. 1925. Dr H. O. Forbe’s Malayan plants. Journal of Botany. 43,

Suppl: 1-136.

Ridley H.N. (1924). Laurineae.,Flora of the Malay Peninsula. 3: 75—137.

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Transactions of the Linnean Society, Botany. 4: 69-263.

Gardens’ Bulletin Singapore 57 (2005) 247—251 247

An Unusual New Species of Begonia

(Begoniaceae) from Vietnam

NGUYEN QUANG HIEU

New York Botanical Garden, Bronx, NY 10458 and

The Arboretum, University of California-Santa Cruz, Santa Crux, CA 96060, USA

AND

MARK C. TEBBITT

Brooklyn Botanic Garden, 1000 Washington Avenue, Brooklyn, NY 11225, USA

Abstract

An unusual new Begonia species, Begonia cucphuongensis H. Q. Nguyen & Tebbitt, is

described from limestone ridges in northern Vietnam.

Introduction

In 2004, the first author discovered a very distinct species of Begonia in

Vietnam’s Cuc Phuong National Park, which is newly described and

illustrated here.

This species has an unusual combination of morphological characters

and cannot be classified within any of the 66 recognized sections of Begonia

(Doorenbos et al., 1998; Shui et al., 2002; Forrest and Hollingsworth, 2003;

de Wilde and Plana, 2003). In many respects the species is similar to B.

boisiana Gagnep., which also has never been assigned to a section despite

having been described as long ago as 1919. Both species share a combination

of characters not otherwise found in the Begoniaceae — both are caulescent

herbs with a distichous leaf arrangement, have palmate-pinnate leaf

venation, lunate stigmas, and entire placentae and lack cystotyles. Both

species are also restricted to shallow soils overlaying limestone in northern

Vietnam. Given these similarities, the two species may well be closely

related.

However, the two species differ considerably in several respects.

Most notably B. cucphuongensis has a rhizome, while B. boisiana is non-

rhizomatous, it also has protogynous rather than protandrous inflorescences,

female flowers with three rather than five tepals and a markedly different

fruit shape. Since many of the characters by which the two species differ

are considered taxonomically important in the sectional classification of

248 Gard. Bull. Singapore 57 (2005)

Begonia, we refrain from classifying either species into an existing section

or from creating any new sections to accommodate these species. This

must await completion of an ongoing molecular phylogenetic analysis of

these and other Indo-Chinese Begonia.

Begonia cucphuongensis H. Q. Nguyen & Tebbitt, sp. nov.

Haec species Begoniae boisianae affinis, sed ab ea habitu rhizomatoso, in

floribus femineis tepalis 3 et ovarii alis aequalibus distinguitur.Typus:

Vietnam. Ninh Binh Province: Cuc Phuong National Park, 19 June 1999,

shade on limestone-derived soil, forest dominated by Streblus macrophyllus,

M. A. Jaramillo et al. MAJ 493 (holo HN; iso CPNP, F, MO).

Figure 1, Plate 1A

Monoecious, shortly rhizomatous, perennial herb, with an upright stem

30-40 cm tall. Stems few-branched, internodes 1.5—6 cm long, sparsely

hairy. Stipules persistent, white, red or purple, elliptic to lanceolate, 1—1.5

x c. 0.3 cm, apex acute, ending in a short hair, margin entire. Leaves

distichous; lamina asymmetric, ovate-lanceolate to oblong-lanceolate, 7—

12.5 x 2-4.5 cm, above green, beneath paler green, often reddish tinged,

both surfaces glabrous, apex acuminate, base oblique-cordate, margin with

a few widely spaced short teeth-like multiseriate hairs where main veins

terminate, teeth-like hairs 1-3 mm long, venation palmate-pinnate; petiole

green, 1.5—4.5 cm long, with sparse microscopic uniseriate glandular hairs.

Inflorescences axillary towards apex, cymose, 2—3-branched, 10- to 15-

flowered, bisexual, protogynous; peduncle green, glabrous, 1—2 cm long;

bracts persistent, pale purple or white, ovate or elliptic, 2-5 x 2-4 mm,

margin dentate. Flowers white to pink. Staminate flowers: pedicel 0.8—1.7

cm long, glabrous; tepals 4, outer two ovate-cordate to cordate, 6—7 x 5—6

mm, margins entire; inner two oblong, c. 5 x 2 mm, glabrous, margin

entire; androphore absent; stamens 15—40, yellow; anthers obovate, c. 1

mm long, apex truncate or rounded, connective not projecting, filament c.

1 mm long. Pistillate flowers: bracteoles absent; pedicel c. 1 cm long,

glabrous; tepals 3, outer two obovate, c. 8 x 8 mm, margin entire, the inner

one elliptic, 7-8 x 3-6 mm, margin entire. Ovary pink, 3-locular, elliptic,

1—1.2 x 0.6—-0.7 cm long, glabrous, wings 3, equal, arcuate-deltoid, c. 8 mm

tall, projecting past locules for c. 1 mm at apex and c. 2 mm at base;

placentae axile, entire; styles 3, shortly fused at base, 2—3 mm long, yellow,

unbranched, stigmas lunate. Fruits becoming dry and dehiscing via slits

positioned next to the wings, c. 1.2 cm long.

Distribution: Vietnam, Ninh Binh province, Nho Quan District, Cuc Phuong

Begonia cucphuongensis from Vietnam 249

Figure 1. Begonia cucphuongensis H. Q. Nguyen & Tebbitt. (PK. Lok et al. P10408).

250 Gard. Bull. Singapore 57 (2005)

National Park; Thanh Hoa Province, Ba Thuoc District, Pu Luong protected

area.

Other specimens examined: Vietnam. Thanh Hoa Province, Ba Thuoc

District, Pu Luong protected area, 500—700 m, 20°25°48”N, 105°14°13”E,

13 April 2001, N. T. Hiep et al. HAL 937, 939 (CPNP, F, HN). Ninh Binh

Province, Cuc Phuong National Park, Bong 20°21’04'N, 105°35’°16" E, on

limestone ridge, 500 m, 20 July 2000, P. K. Loc et al. P10323, 10408 (CPNP,

F, HN); 9 Dec 1999, N. T. Hiep et al. NTH 4098 (HN); 18 Nov 1999, N. T.

Hiep & N. X. Tam 4093 (HN); 15 Mar 2000, N. T. Hiep 4188 (HN); 15 Mar

2000, N. T. Hiep 4189; D. D. Soejarto et al. 11080 (CPNP, F, HN).

Acknowledgements. This study was supported by Grant 1-UO1-TW01015,

International Cooperative Biodiversity Group (ICBG) through funds from

NIH, NSF, and Foreign Agricultural Service of the USDA. The authors

thank the personnel of the Cuc Phuong National Park and the Directors of

the Herbaria of the Cuc Phuong National Park, Institute of Ecology and

Biological Resources, and the Field Museum. Permits for the collection

and export of plant specimens were granted by the Vietnamese Ministry of

Agriculture and Rural Development, Hanoi, through a letter dated 15

September 1998, Ref. No. 3551/BNN/KHCN, and from the Cuc Phuong

National Park, through a letter dated 16 September 1998. We thank Roy

Gereau for help with the Latin diagnosis, Bui Xuan Chuong for the

illustration, D. Doel Soeyjarto, [Ihsan Al-Shehbaz, Peter F. Stevens, Daniel

Harder and Ruth Kiew for comments on an earlier draft of the manuscript.

References

Doorenbos, J., M. S. M. Sosef and J. J. F. E. de Wilde 1998. The Sections of

Begonia Including Descriptions, Keys and Species Lists (Studies in

Begoniaceae VI). Wageningen Agricultural University Papers. 98—2,

Netherlands.

Forrest, L. L. and P. M. Hollingsworth. 2003. A recircumscription of

Begonia based on nuclear ribosomal sequences. Plant Systematics and

Evolution. 241: 193-211.

Shui, Y. M., C.-I. Peng and C. Y. Wu. 2002. Synopsis of the Chinese

species of Begonia (Begoniaceae), with a reappraisal of sectional

delimitation. Botanical Bulletin of Academia Sinoca. 43: 313-327.

Begonia cucphuongensis from Vietnam Oi

Plate 1. Innflorescences of A. B. boisiana Gagnep. (NTH 3069) and B. Begonia cucphuongensis

H. Q. Nguyen & Tebbitt (NTH 4098). Photographs H. Q. Nguyen.

Milde, I, J. EF. Es de. and V: Plana. 2003. A new section of Begonia

(Begoniaceae) from West Central Africa. Edinburgh Journal of Botany.

60: 121-130.

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Gardens’ Bulletin Singapore 57 (2005) 253—261 ZS

Three New Species of Scaphochlamys

(Zingiberaceae) from Peninsular Malaysia

Y.Y. SAM AND L.G. SAW

Forest Research Institute Malaysia,

52109 Kepong, Selangor, Malaysia

Abstract

Three new species of Scaphochlamys are described from Peninsular Malaysia: S. abdullahii

Y.Y.Sam & Saw, S. cordata Y.Y.Sam & Saw and S. laxa Y.Y.Sam & Saw. All three are

endemic to the eastern part of Peninsular Malaysia, with S. abdullahii and S. cordata

confined to only one locality each in Terengganu.

Introduction

Scaphochlamys is a genus of small gingers only known from the West

Malesian forest. Twenty six species are known so far and Peninsular

Malaysia has the highest number of species; 19 species (Newman eft al.

2004). In recent years, collections made from newly accessible forested

area have revealed a number of new species. In this paper, we described

three, S. abdullahii, S. cordata and S. laxa.

The description below is based on herbarium specimens and fresh

material (especially important for floral parts) including plants in cultivation

in the nursery of Forest Research Institute Malaysia. In most cases, the

cultivated plants are similar in size to the wild population except for S.

cordata, which tends to be smaller.

1. Scaphochlamys abdullahii Y.Y.Sam & Saw sp. nov.

Scyphochlamide longifolia similis, folius infra pilis dense appressis (nec costa

pilosa solum) petiolo glabro (nec pilosus), bracteolis quam bracteis

brevioribus (nec longiores), bracteis plus quam unifloris (nec uniflorae)

differt. Typus: Peninsular Malaysia, Terengganu, Setiu, Ulu Setiu Forest

Reserve, 17 January 2005. Sam, Aidil & Arif FRI 49130 (holo KEP; iso

SAN, SING).

Figure 1.

Ground herb to 55 cm tall, rhizome creeping underground, cut rhizome

purple, 5-6 mm diam. Leafy shoots erect, close together, one leaf per

shoot, rarely 2; bladeless sheaths 3, red to brownish red, ridged

254 Gard. Bull. Singapore 57 (2005)

longitudinally, densely hairy especially when young, largest sheath 6—10.5

cm, drying during anthesis. Leaf sheath 1.5—3 cm, ligule bilobed, lobes

small; petiole (5.5—)9—29 cm, round in cross section, channelled only in the

terminal part, purplish brown at the base; lamina elliptic, 12—33 x 3.8—7.6

cm, unequal, apex narrowly acute, base narrowly rounded or decurrent,

sometimes oblique, with densely appressed hairs on lower surface, red to

purplish red when young. /nflorescence, 6.5—17 cm long, red when young;

peduncle 4-13 cm long, covered entirely by the bladeless sheath when

young; inflorescence head ovoid, 3—5.5 cm long. Bracts 8—25, spirally

arranged, 19-25 x 13—20 mm, basal ones larger, spathulate, apex blunt,

reflexed, margin thin and slightly wavy; rosy red when young, turning to

reddish brown or green, imbricate and hiding the rachis entirely. Bracteoles

shorter than the bracts, first bracteole largest, 10—16 x 5—7 mm, apex bilobed

and hairy, tinged red. Flowers 5—6 per bract, white except for the calyx.

Calyx tubular, 4~7 mm long, tinged red, apex blunt, split 2—3.5 mm on one

side from apex. Corolla tube 17—25 mm. Dorsal corolla lobe lanceolate,

10-12 x 34 mn, apex hooded, tinged red distally. Lateral corolla lobes

lanceolate, 7-10 x 2—4 mm, tip tinged pink to red. Staminodes oblong, 8.5—

11 x 2—3 mm, apex blunt, slight yellowish. Labellum 13—14 x 9-11 mm,

bilobed, recurved, c. 5 x 5 mm, median band yellow, both sides with purple

streaks, sparsely papillose on the abaxial surface. Stamen flushed purple,

covered with glandular hairs on the abaxial surface; filament c. 2 mm long;

thecae 2.5—4 mm long, yellow; anther crest c. 1.5 x 2 mm, recurved.

Epigynous glands 2, linear, c. 5 mm long, yellow. Ovary ellipsoid, trilocular.

Seed ellipsoid, c. 8 x 3 mm, dark brown.

Distribution: Endemic to Peninsular Malaysia. Very rare, only recorded

from Setiu District, Terengganu.

Habitat: Lowland forest, c. 41 m altitude. This species was found in forest

fringes.

Notes: Scaphochlamys abdullahii belongs to the single-leaf group and is

closest to S. longifolia, which has similar closely imbricating bracts and

elliptic leaves. However, it differs from S$. longifolia in having densely

appressed hairs on the lower leaf surface, a glabrous petiole and it produces

many flowers in the axil of each bract; S. /ongifolia has hairs on the midrib

and petiole and single-flowered bracts (Holttum, 1950).

Ridley described S. longifolia in 1924 but its type was missing when

Holttum (1950) revised the genus. The description given by Holttum was

based on a specimen collected from the type locality that he thought was

the same species. Although the details of the flower parts were not

Scaphochlamys in Peninsular Die Be

: a

Sats Ns y

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SS SES YE

aR Ye

Figure 1. Scaphochlamys abdullahii Y.Y.Sam & Saw. A. habit; B. floral bract with flower; C.

calyx; D. dorsal corolla lobe; E. lateral corolla lobes; F. staminodes; G. labellum; H. anther

front, side and back view; I. stigma. (A. from type specimen; B, C, D, E, F, G, H, I from living

collection of 2002—0072).

256 Gard. Bull. Singapore 57 (2005)

satisfactory, other characters, e.g. leaf, inflorescence structure and floral

bracts, are sufficient to differentiate S. abdullahii from S. longifolia.

Scaphochlamys abdullahii has a bright rosy red colour in its young

inflorescence. This is distinct from other Scaphochlamys species that usually

have dull blood-red or brownish red in their young floral bracts.

Scaphochlamys abdullahii is named in honour of the Prime Minister

of Malaysia, Datuk Seri Abdullah Ahmad Badawi, for his strong interest

in nature conservation and protection of the environment.

Other specimens examined: TERENGGANU, Setiu, Kuala Sungai Bok via

Kampung Seladang, 14 Mar 1975, Mohd. Shah & Ahmad MS 3514 (SING);

Ulu Setiu Forest Reserve, 16 Aug 2001, Khaw KSH 743 (KEP); Cultivated

material 14 July 2002 Sam FRI 47053 (KEP).

2. Scaphochlamys cordata Y.Y.Sam & Saw sp. nov.

Scaphochlamys atroviridi similissima, folii basi cordata (nec cuneata ad

decurrenti), petiolo longiore (31—55.5 vs. 6-10 cm longus), inflorescentia

longiore (21.5—25.3 vs. 5-10 cm longa) differt. Typus: Peninsular Malaysia,

Terengganu, Dungun, Jengai Forest Reserve, 17 October 2002. Sam &

Mustapa FRI 47155 (holo KEP; iso E, SAN).

Figure 2.

Ground herb, rhizome creeping on ground surface. Leafy shoots ascending,

close together, one leaf per shoot; bladeless sheaths 4, reddish brown, apex

mucronulate, largest sheath 11-17 cm. Leaf sheath 1.5—2 cm long, ligule

bilobed, lobes small, c. 1 mm; petiole slender, 31-55.5 cm long, round in

cross section; lamina cordate, 15.5—28.5 x 8.2—14.7 cm, slightly asymmetric,

apex acuminate, base strongly cordate, some with auricles overlapping;

secondary veins 8—10 pairs, first 6 pairs always originate from the base,

prominently raised on the upper leaf surface; leaf below densely hairy

proximally, brownish red when young. /nflorescence slender, 21.5—25.3 cm

long, red when young; peduncle 10—17.8 cm long, hidden within bladeless

sheath when young; rachis 5.5—8.5 cm. Bracts spirally arranged, 4—10, lax,

1.5—2 cm apart, red when young, turning to green, spathulate, 15—22 x 5.5—

12 mm when flattened, hairy towards the apex on both surfaces, distally

reflexed. Bracteoles shorter than the bracts, narrowly lanceolate, 6—11 x

1—3 mm with inflexed margin, apex pointed, hairy throughout, first bracteole

largest, enclosing subsequent bracteoles and flowers. Flowers more than 15

per bract, small, white except for the calyx. Calyx tubular, 7.5—9 x 1.5—2

mm, red, hairy, split 2.5—4 mm down on one side from apex. Corolla tube

11-14 mm. Dorsal corolla lobe lanceolate with strongly inflexed margin,

6—8.5 x 1.5—2 cm, apex hooded, pinkish distally, papillose on the abaxial

Scaphochlamys in Peninsular 257

Figure 2. Scaphochlamys cordata Y.Y.Sam & Saw. A. habit; B. inflorescence; C. bract front and

side view; D. dorsal corolla lobe; E. lateral corolla lobes; F. staminodes; G. labellum; H. anther

front, side and back view; I. stigma. (A from type specimen; B, C from living material of 2004-

0728; D, E, F G, H, I from FRI 49135).

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ioe)

Gard. Bull. Singapore 57 (2005)

surface; lateral corolla lobes lanceolate with strongly inflexed margin, 5—7

x 1—2 mm, pinkish distally, papillose on the abaxial surface. Staminodes

oblong, 3—7.5 x 1—2 mm, apex rounded, papillose on the adaxial surface.

Labellum 7—10 x 5—9.8 mm, bilobed, lobes overlapping, cleft c. 4 mm from

distal end, median band faintly yellow, purple blotches at both sides. Stamen

3.5—4.5 mm, tinged purple; filament short, c. 1 mm long; thecae c. 2.5 mm

long, anther crest short, less than 1 x 1.5 mm. Stigma small, cup-shaped,

yellowish, papillose.

Distribution: Endemic to Peninsular Malaysia; only known from Jengai

Forest Reserve in Terengganu.

Habitat: Lowland dipterocarp forest, at 80-140 m altitude. The species is

observed to favour ridges.

Notes: Scaphochlamys cordata belongs to the single leaf group with very

lax floral bracts in the inflorescence. This species is similar to S. atroviridis

but differs in the leaf shape and apex, petiole and inflorescence length, size

of floral bracts and labellum (Table 1).

Table 1. Diagnostic characters to distinguish Scaphochlamys cordata and

S. atroviridis

Characters S. cordata S. atroviridis

Leaf shape cordate elliptic with cuneate to

decurrent base

Leaf apex acuminate rounded

Petiole length (cm) 31-55.5 6-10

Inflorescence length (cm) | 21.5-25.3 5-10

Size of floral bracts (mm) | 15-22 x 5.5-12 30-35 x 15

Size of labellum (mm) 7-10 x 5-9.8 18x15

This species is named for its distinctly cordate leaves.

Other specimens examined: TERENGGANU, Dungun, Jengai Forest

Reserve, 9 Mar 2005, Sam FRI 49135 (KEP). Cultivated material 23 Aug

2000 Saw FRI 44306 (KEP).

3. Scaphochlamys laxa Y.Y.Sam & Saw sp. nov.

Scyphochlamide cordata similis, petiolo longiore (31-36 vs. 13-29 cm

259

Scaphochlamys in Peninsular

4y 7

VA Zu

g@erWe LL

i, i]

Myf

NG:

Figure 3. Scaphochlamys laxa Y.Y.Sam & Saw. A,B habit; C. inflorescence; D. calyx; E. dorsal

corolla lobe; F. lateral corolla lobes; G. staminodes; H. labellum; I. anther front, side and back

view; J. stigma. (A, C from living material of 2004-0718; B, D, E, FG, H,1,J from type specimen).

260 Gard. Bull. Singapore 57 (2005)

longus), foliis late ellipticis vel ovatis (nec cordata), labello maiore (11.5—

22 vs. 7-8 cm longum, 10—20 vs. 5—6 cm latum) distinguenda. Typus:

Peninsular Malaysia, Terengganu, Dungun, Jengai Foest Reserve, 9 March

2005. Sam & Apok FRI 49136 (holo KEP; iso SAN, SING).

Figure 3.

Rhizomatous herb; rhizome running horizontally on the ground, fleshy.

Leafy shoots erect, 30-40 cm tall, red when young; one leaf per shoot, 1—3

cm apart (sometimes much closer and appearing as a two-leaved shoot

when under stress or in cultivation); bladeless sheaths 3, red, ridged

longitudinally, largest 7—-13.5 cm. Leaf sheath 3.8—4.5 cm, ligule bilobed,

lobes not obvious; petiole 13—36 cm long, grooved in cross section, red

when young; lamina broadly elliptic or ovate, 13—22 x 10.5—14 cm, apex

cuspidate, base slightly decurrent on the petiole, never cuneate, margin

ciliate; secondary veins slightly raised on the upper surface, dark green,

some with white bands in between midrib and leaf margin; leaf underneath

pale green, red when young. /nflorescence 15—33 cm long; peduncle 9—12

cm, red when young; rachis 2.5—11 cm. Bracts spirally arranged, (2-4—)5—

10, lax, 0.4-2 cm apart, dark green, spathulate, broadly elliptic when

flattened, 1.7—2.5 x 0.8—1.6 cm, apex acute, distally reflexed, margin hairy

when young. Bracteoles smaller than the bracts, 5—7 x 4.5—6 mm, hairy

towards apex, first bracteole enclosing subsequent bracteoles and flowers.

Flowers more than 20 per bract, white except for the calyx. Calyx tubular,

6—7 x c. 2 mm, apex bilobed, tinged red, split 3-4 mm down on one side

from apex. Corolla tube 13-17 mm long. Dorsal corolla lobe lanceolate, 8—

13 x 3-4 mm with inflexed margin, apex hooded and pointed, faintly yellow.

Lateral lobes narrowly lanceolate with incurved margin, 7—11 x 2—3 mm,

apex also hooded and pointed. Lateral staminodes oblong, apex blunt, 9—

11.5 x 1-3 mm, papillose on the adaxial surface, faintly yellow towards the

apex. Labellum hairy, 11.5—22 x 10—20 mm, bilobed, papillose on the upper

surface, lobes 5.5—7 x 6—7 mm, cleft c. 6 mm from distal end, crisped,

median band yellow with purple streaks at both sides. Filament 2.5—3 mm,

some with purple blotches; thecae 2—2.5 mm long; anther crest 0.5—1 x 2.5—

3 mm, apex trilobed, strongly recurved. Stigma c. 1 mm wide, cup-shaped,

covered with brown papillae. Epigynous glands 2, linear, c. 4 mm, yellow.

Ovary c. 2 mm long, trilocular.

Distribution: Endemic to Peninsular Malaysia. Found on the Timur Range

of Pahang and Terengganu.

Habitat: The species is found on undulating land and slopes of lowland

dipterocarp forest, from 65 to 130 m a.s.l., where it grows in abundance

Scaphochlamys in Peninsular 261

under heavy forest shade.

Notes: Scaphochlamys laxa is unique in having a very broadly elliptic or

ovate lamina, unlike other Scaphochlamys species. S. laxa is similar to S.

cordata but is distinguished by its shorter petiole (13—29 vs. 31-36 cm),

grooved in cross section (round in S. cordata) and broadly elliptic or ovate

leaves. The other differences include the larger flowers and _ the trilobed

and reflexed anther crest of S. laxa.

The name ‘/axa’ refers to the bracts that are very lax on the rachis.

Other specimens examined: TERENGGANU, Dungun, Jengai Forest

Reserve, 17 Oct 2002, Sam & Mustapa FRI 47152 (KEP); Pasir Raja Selatan

Forest Reserve, 7 June 2004, Markandan FRI 49179 (KEP). PAHANG,

Jerantut, Gunung Aais Forest Reserve, Sungai Lurut, 5 Jul 2004, Sam FRI

49057 (KEP).

Acknowledgements

We wish to thank the Director-General of FRIM for his constant support

in plant research studies; M. Markandan, D. Mustapa, K. Apok and A.

Angan for field assistance; S. H. Khaw for providing her collection. We

also like to thank Dr. J.F. Veldkamp for the Latin diagnosis. M. Zainal

and M.N. Aidil illustrated the plates. This study is supported by the Timber

Levy Fund under the Flora Malaysiana Centre Project 1: Domestication of

Herbaceous Plants for Horticultural Introduction.

References

Holttum, R.E. 1950. The Zingiberaceae of the Malay Peninsula. The

Gardens’ Bulletin Singapore 13: 82-105.

Newman, M., Lhuillier, A. & Poulsen, A.D. 2004. Checklist of the

Zingiberaceae of Malesia. Blumea, Supplement 16: 145—150.

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Gardens’ Bulletin Singapore 57 (2005) 263—267 263

Malaxis inexpectata and Habenaria

paradiseoides (Orchidaceae),

New Records for Peninsular Malaysia

M. JUTTA!

AND

O.Z. FARIDAH!?

Institute of Bioscience, Universiti Putra Malaysia (UPM),

43400 Serdang, Malaysia.

"Biology Department, Faculty of Science, Universiti Putra Malaysia (UPM),

43400 Serdang, Malaysia.

Abstract

Two new records, Malaxis inexpectata (J.J.Sm.) J.B.Comber and Habenaria paradiseoides

J.J.Sm., are reported for the Orchidaceae of Peninsular Malaysia. Until now, both were

known only from Sumatra, M. inexpectata with two, and H. paradiseoides with one collection.

Malaxis inexpectata

Malaxis inexpectata (J.J.Sm.) J.B.Comber, Orchids of Sumatra (2001) 172.

syn. Pseudoliparis inexpectata (J.J.Sm.) Marg., Ann. Bot. Fennici. 40 (2003)

63. Microstylis inexpectata J.J.Sm., Bull. Jard. Bot. Buit. ser. 3, 10 (1928)

33.

Figure 1

Small herbs with closely spaced pseudobulbs, somewhat laterally compressed

stems measuring 1.5 cm long by 0.6 cm wide. Leaves 4—5, the upper ones

largest; lamina with three prominent veins, elliptic-ovate, apex acute-

acuminate, margins undulate, 4.5 x 1.7 cm; petiole to 1 cm long, ending in a

broad overlapping sheath. Inflorescence terminal; scape ~5 cm long; rachis

bearing up to 20 flowers, opening 2—3 at a time; floral bracts narrowly

lanceolate, acute, reflexed, 5 mm long. Flowers non-resupinate, orange.

Median sepal ovate, obtuse, 4 x 2.2 mm; lateral sepals a little smaller,

oblique; petals lanceolate, obtuse, 2.4 x 0.75 mm, recurved. Labellum

including auricles about 3.4 mm long and 2.8 mm wide, not toothed; side

lobes inconspicuous, midlobe curved up; fovea oblong with a distinct linear

264 Gard. Bull. Singapore 57 (2005)

median depression and a small raised callus at the base; auricles not

touching, apex rounded, 1.6 mm long. Column short, curved, winged, with

a distinct, papillose horn-like projection at the back below the dorsally

positioned anther; pollinia four, club-shaped.

Specimens examined: KEDAH: Langkawi, Gunung Machinchang 10 Jan

2004 Mariam Jutta LCO/F-H I (UPM-IBS Herbarium; flower spirit

specimen); Mariam Jutta LCO/F 148 (live collection, Astaka Orkid UPM).

Notes: The species was collected as a tiny bulb on the lower peak of

Gunung Machinchang and flowered in the live collection at Astaka Orkid,

Universiti Putra Malaysia (UPM). It could not be confidently identified

using the key in Seidenfaden & Wood (1992). Based on floral characteristics,

it keys out closest to Malaxis prasina (Ridl.) Seidenf. & Smitinand, the

only Peninsular Malaysian species with an untoothed apex of the labellum,

but it is distinct from all Malaxis species reported from Peninsular Malaysia

in having a ‘horn’ on the dorsal part of the column. The dorsal horn plus

the orange flower colour in M. inexpectata are characteristic of Sect.

Pseudoliparis, and make this collection the first record of this section for

Peninsular Malaysia.

Our specimen matches the original description by Smith (1928) in most

vegetative and floral aspects except: length of the lip 3.4 mm vs. Smith’s

2.4 mm; apex of labellum entire vs. ‘a little erose’; dorsal horn on column

papillose vs. hairy (Fig. 1). However, after discussion with Peter O’Byrne,

it was concluded that the Langkawi specimen merits inclusion in M.

inexpectata.

This paper applies the name Malaxis inexpectata (J.J.Sm.) J.B.Comber as

accepted by the World Checklist of Monocots (Govaerts, 2004). The name

Pseudoliparis inexpectata Marg. (Margonska, 2003) is treated as a synonym.

Habenaria paradiseoides

Habenaria paradiseoides J.J.Sm., Bull. Jard. Bot. Buit. ser. 3, 12 (1932)

105.

Figure 2

Small herb with closely spaced tufted stems, glabrous throughout. Stems

short, each bearing about 8—10 linear, acute leaves, to 10 cm long by 0.5

cm wide. Stem above leaves 20—25 cm long, bearing a few leaf-like bracts,

the lowest one up to 6 cm long; rachis to 10 cm long, with up to 20 white

265

Scaphochlamys in Peninsular

2mm

lateral sepal median sepal petal

: column side view

lip

T QW 0.4 men

pollinia

ilabichack back view column, front view

Figure 1. Flower parts of Malaxis inexpectata (J.J.Sm.) J.B.Comber.

266 Gard. Bull. Singapore 57 (2005)

flowers; floral bracts persistent, triangular acuminate, to 1.5 cm long. Flowers

predominantly white. Median sepal about 3 mm long, broadly elliptic,

white flushed with green at the back, and forming a hood with the obliquely

lanceolate petals; lateral sepals of two unequal halves, oblique, 5.5 mm

long, recurved, in-rolled. Labellum with conspicuous, winged, often laciniate

side lobes measuring about 2 cm across; midlobe linear, obtuse, 7 mm

long; spur 1.4 cm long, thickened towards the base, and curved forward.

Column 2.0 mm long by 2.5 mm wide; anther canals compact, short, bent

upwards.

Specimens examined: TERENGGANU: Pasir Raja Forest Reserve (FR) 8

June 2004 Mariam Jutta LCO/F 150 (live collection Astaka Orkid UPM); 5

April 2005 Mariam Jutta LCO/F-H 9 (UPM-IBS Herbarium); 11 April

2005 Mariam Jutta LCO/F 544 (live collection, material passed to Singapore

Botanical Garden June 05). PAHANG: Jerantut, Gunung Aais FR,

Markandan Moorthy FRI 49202 (KEP, SAN, SING).

Notes: Habenaria paradiseoides is considered as having no near allies within

the genus (Smith, 1932). Its closest relatives among the Peninsular Malaysian

species are H. reflexa Blume and H. kingii Hook.f., which have a similar

habit, but differ in being limestone species opposed to H. paradiseoides

that was found on granite. Flowers, though sharing some similar

characteristics in the lateral sepals and spur, show distinct differences

between H. paradiseoides and the other two species, especially in structure

of the labellum and spur (both H. kingii and H. reflexa) and of the anther

canals (H. reflexa).

This species was collected twice in Peninsular Malaysia from similar

habitats (exposed locations in narrow gaps between rocks lining the banks

of large rivers, often prone to strong currents) in Pahang and Terengganu.

Both the description and habitat recorded by Smith (1932) match these

specimens. Variations in the side lobes of the labellum are common, even

on the same plant (Fig. 2), ranging from almost entire to deeply laciniate

margins and either two- or three-sectioned lobes, the former being more

common.

Acknowledgements

We acknowledge the help and advice of Jaap J. Vermeulen, Sam Yen Yen,

Ahmad Ngah and Peter O’Byrne. Ruth Kiew assisted in the preparation of

the manuscript.

Scaphochlamys in Peninsular 267

References

Comber J.W. 2001. The Orchids of Sumatra. Natural History Publications,

Kota Kinabalu, Malaysia.

Govaerts R. 2004. World Checklist of Monocotyledons. Royal Botanical

Gardens, Kew. Published on the Internet; http://www.kew.org/

monocotChecklist/ (accessed 22"? August, 2005)

Margonska H.B. 2003. Materials towards a revision of the genus

Pseudoliparis (Orchidaceae, Malaxidinae) Annales Botanica Fennici 44:

63-66

Seidenfaden G. and J. J. Wood. 1992. The Orchids of Peninsular Malaysia

and Singapore. Olsen & Olsen, Denmark.

Smith J. J. 1928. Orchidaceae novae malayenses XIII. Bulletine Jardine.

Botanique. Buitenzorg. ser. 3, 10: 33-34

Smith J. J. 1932. Orchidaceae novae malayenses XIV. Bulletine Jardine.

Botanique. Buitenzorg. Ser. 3, 12: 105—106.

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Gardens’ Bulletin Singapore 57 (2005) 269—278 269

New Tristaniopsis Peter G.Wilson &

J.T.Waterh. (Myrtaceae)

From Borneo

P. S. ASHTON

Arnold Arboretum, Harvard University, 22 Divinity Avenue

Cambridge MA 02138, U.S.A.

and

Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, U.K.

Abstract

Three new species, Tristaniopsis kinabaluensis P.S.Ashton, T. microcarpa P.S.Ashton and

T. rubiginosa S.Teo ex P.S.Ashton, and three new subspecies, Tristaniopsis kinabaluensis

ssp. silamensis P.S.Ashton, T. merguensis ssp. tavaiensis P.S.Ashton and T. whitiana ssp.

monostemon P.S.Aston, are described from northern Borneo, in preparation for a treatment

of the Myrtaceae for the Tree Flora of Sabah and Sarawak.

Introduction

Species definition in Tristaniopsis Peter G.Wilson & J.T.Waterh. (formerly

Tristania R. Br.) has proven to be even more difficult in Borneo than

among the notorious and much larger myrtaceous genus Syzygium Gaertn.

Leaf size and shape is variable. The number of stamens, which are clustered

opposite the petals, is characteristic of each species but, although at most

10 per cluster, may vary in exceptional cases. Here, we adopt a conservative

species concept, awaiting regional monographic and phylogenetic research.

Eventually, and with further flowering collections, some at least of the

infrapecific taxa described here may be raised to species rank.

All specimens examined have been at the Kew herbarium unless otherwise

stated.

1. Tristaniopsis kinabaluensis P.S.Ashton, sp. nov.

T. merguensis affinis, foliis minoribus basim versus subsessilibus attenuatis

vel anguste obtusis haud auriculatis, subtus hebete glaucescentibus,

staminibus 3(-5) in fasciculis, fructibus ad 5 x 4 mm minoribus facile

270 Gard. Bull. Singapore 57 (2005)

distinguitur. Typus: A. Gibot SAN 60705, Sabah, Bukit Hampuan, Ranau,

in flower (holo K; iso SAN).

Tree to 20(-35) m tall, to 30 cm diameter. Bark rust-brown, cracked and

falling in small flakes; inner bark red-brown. Young parts densely yellowish

silvery to pale brown downy, caducous. Twig c. 3 mm diameter, stout,

eventually dark brown. Leaf subsessile, thickly leathery, dull, drying mauve

to dark yellowish brown, generally pale mauve glaucous beneath, pits more

or less apparent above, dots faint to obscure beneath; blade obovate-

oblanceolate, 4-12 x 1.5—4.5 cm, base tapering or abruptly terminating at

the c. 1 mm stout stalk, apex rounded or shortly broadly acuminate; venation

hardly or not raised-on either surface, main lateral veins c. 15 pairs,

ascending. Cyme to 8 cm long, terminal or axillary, doubly branched, the

branches and flowers clustered towards the end of the rachis. Flower bud

to 4 x 3 mm, obconical, subsessile; sepal lobes to 1 x 1 mm, ovate acute;

stamens 3(-5) per cluster. Capsule to 4 x 3 mm, ellipsoid or spherical.

Notes: As Tristaniopsis merguensis but leaf smaller, subsessile, base tapering

or narrowly obtuse not auriculate even in juveniles, dull more or less

mauve glaucous beneath; stamens 3(-5) per fascicle; capsule to 5 x 4 mm,

smaller. The floral stamens are 3 in a fascicle as in 7. whitiana (Griff.)

Peter G.Wilson & J.T.Waterh. but the leaf is smaller, thickly leathery, and

subsessile.

Subspecies kinabaluensis

Subspecies tomento rufescente-brunneo, costis foliorum obscuris, fructu

ad 5 x 4 mm lobis sepalorum ad 1 x 1 mm capsula amplexa valvis ad 4 x 2

mm ellipticis distinguitur.

Hairs dull rufous-brown; leaf venation obscure throughout; fruit to 5 x 4

mm, sepal lobes to 1 x 1 mm clasping base of to 4 x 2 mm elliptic capsule

valves following dehiscence.

Distribution and ecology: Known only in and near Kinabalu National Park.

Locally common in lower montane oak-laurel forest.

Other specimens examined: SABAH - Mikil SAN 33924 (K), Mikil SAN

33945 (K), Madani SAN 36776 (K, SAN), road to Kamburangoh, Ranau;

Meijer SAN 37990 (K, SAN), above Tenompok; Clemens 30242 (K),

Tenompok; Chew & Corner RSNB 4859 (K, SING); RSNB 4148 (K, SAN,

SING) Mesilau R.; Chew & Corner RSNB 4458 (K, SAN, SING)

Tristaniopsis in Borneo 271

Bambangan R.; Chew et al. 1862 (K, SAN, SING), Beaman 10699 (K),

Pinosuk Plateau; Beaman 9381 (K), above bank of E. Mesilau R.

Notes: Several collections from Sarawak, in fruit or young bud, differ in

their less coriaceous, dull but not glaucous leaf blade. These include Chai

S 30427 (K, SAN, SING), Anderson S 4289 (K, SAN, SING), Abg. Mochtar

et al. S 30818 (K, SAN), Gunung Api, Mulu N.P.; B. Lee S 38259 (K,

SAN), Bukit Berar, Mulu N.P.; Abg. Mochtar et al. S 49609 (K, SAN),

Mulu N.P.; Yi, P.C. S 41131 (K, SAN), Sabal F.R., Serian. They were

collected from organic soils on limestone to 100 m altitude, and on

sandstone. Good flowering material is needed to decide their identity.

Subspecies silamensis P.S.Ashton, ssp. nov.

Subspecies tomento argento-luteo, costis foliorum patentibus costis

lateralibus subtus elevatis, calyce in fructu tenue lobis ad 3 x 4 mm rotates,

capsula ad 4 mm diam. globosa valvis ad 4 x 3 mm latioribus differt. Typus:

G. Shea SAN 75187, Sabah, Gunung Silam, Lahad Datu, 800 m, in flower

(holo K; iso SAN, SING).

Hairs yellowish silvery; leaf venation visible, the main lateral veins slender

slightly raised beneath; fruit calyx to 8 mm diameter, shallowly cup-shaped,

the lobes spreading, to 3 x 4 mm; capsule to 4 mm diameter, spherical,

valves broadly elliptic, to 4 x 3 mm.

Distribution and ecology: Known only from Mt. Silam, coastal E. Sabah;

on ultramafic substrate at 800-900 m in lower montane kerangas.

Other specimens examined: Shea SAN 75180 (K), Mujin SAN 37845 (K,

SAN, SING), Beaman 6991 (K), Mt. Silam.

2. Tristaniopsis merguensis (Griff.) Peter G.Wilson & J.T.Waterh.

Austr. J. Bot. 30 (1982) 430. Basionym: Tristania merguensis Griff., PI.

Cantor.(1857) 18. Holotype: Griffith s.n., Mergui, peninsular Burma (holo

K).

Synonyms:

Tristania maingayi Duthie, Fl. Brit. Ind. 2 (1878) 467; T. subauriculata

King, J. As. Soc. Beng. 70, 2 (1901) 502.

New synonyms:

T. stellata Ridl., J. Bot. (1930) 38. Type: Haviland 1983, Sarawak, Kuching

ave Gard. Bull. Singapore 57 (2005)

(holo K); T. grandifolia Ridl., loc.cit., Tristaniopsis grandifolia (Ridl.) Peter

G.Wilson & J.T.Waterh., loc. cit. Type: Beccari 2489, loc. incert. (holo K).

Subspecies merguensis

Canopy tree to 30 m high, to 40 cm diameter. Bark red-brown, becoming

irregularly cracked and coarsely flaky in scroll-like pieces, fibrous; inner

bark pale red brown, rather crumbly. Sapwood rich red-brown, hard.

Inflorescence rachis persistently minutely yellow-brown pubescent, flower

buds caducously so, parts otherwise hairless. Twigs 3—5 mm diam., round,

glabrous, smooth soon thinly flaking. Leaf blade elliptic to obovate, (6—

)10(-17) x (2—)4(—7) cm, leathery, glabrous; apex sharply to broadly acute,

base without distinct stalk, distinctly auriculate in juvenile leaves but less

distinctly so in mature leaves; main lateral veins shallowly furrowed to

slightly raised above, raised below, (17—)28(—34) pairs, 2-8 mm apart,

ascending; intramarginal vein c. 1 mm within margin. Cyme to 12 cm long,

to 5-branched. Flower bud c. 3 x 2 mm; pedicel to 2.5 mm long; calyx lobes

c. 1 x 1 mm; petals c. 1.5 mm long; stamens (3—)5—10 per cluster, filament

1—2 mm long; anther c. 0.1 mm. Capsule to 1 x 0.8 cm, ellipsoid-globose,

pedicel 0.5—1 mm long; seeds c. 0.8 x 0.3 cm, relatively large, elliptic.

Distribution and ecology: East coast of Peninsular Malaysia (including

Singapore) and throughout Borneo. In lowland mixed peat swamp forest

over sandy alluvium, kerangas forest over ultramafic rock and rarely on

limestone hill and organic soil on high ridge tops to 1000 m.

Specimens examined from Borneo: SABAH — Ahmad Talip SAN 68396

(K, SAN, SING), Silam, Lahad Datu; Joseph B. & Kuntul SAN 12085 (K),

Mt Silam; A. Gibot SAN 66926 (K, SAN), Lumut, Beaufort; Saikeh SAN

73354 (K, SAN), Mesapol, Sipitang. SARAWAK — Bujang Tajai S 1997

(K), Smythies S 808 (K), Kayangeran F.R., Lawas; Haviland & Hose 3187

(K), ‘Baram’; Anderson S 30830 (K, SING), Gunung Api, Mulu N.P.; Brunig

S 0970 (K), Merurong Plateau, Bintulu; Othman & Abak S 8854 (K, SAN,

SING), Bukit Urang, Bintulu; Brain S 15947 (K, SING), Nyabau F.R.,

Bintulu; A. Muas S$ 13356 (K, SAN), Balai Ringin P.F., Serian; Hj. Bujang

S. 13440 (K, SAN, SING), Gunung Selang, Kuching; Yii, P.C. S 42956 (K,

SAN), Telok Gador, Bako N.P., Kuching; Haviland 1983 (K), near Kuching;

Beccari 3676 (K), 2489 (K), loc. incert. BRUNEI — Ashton S 5898 (K,

SING), Seria; Ashton BRUN 808 (K), Ashton 819 (K), Labu, Temburong.

E. KALIMANTAN — Kostermans 13137 (K, SING), Mt. Palimasan, W.

Kutel.

Tristaniopsis in Borneo 213

Notes: We agree with the synonymy recommended by Kochummen (1978),

but doubt whether 7Jristaniopsis pontianensis M.R. Henders. of southern

Johor is distinct either. We here formally reduce T. grandifolia Ridl., in

fruit, whose large leaves are typical of immature trees and T. stellata Ridl.

in which the leaves are at the small end of the observed range.

Subspecies tavaiensis P.S.Ashton, ssp. nov.

A species typico foliis ad 11 x 4 cm minoribus distincte petiolatis ad 5 mm

late acuminates, cymis ad 8 cm longis brevioribus. Typus: L. Madani SAN

81723, Tavai Forest Reserve, Telupid, Sabah (holo K; iso KEP, SAN,

SING). -

Differing from the type subspecies as follows:

Leaf blade to 11 x 4cm, smaller, with distinct 8-10 mm long stalk, to 5 mm

long broadly acuminate apex; cyme to 8 cm long, shorter.

Distribution and ecology: Known only from Bukit Tavai F.R., S. Meliau,

Karamuak, Telupid in forest overlying ultramafic rock.

Other specimens examined: SABAH — Soepadmo et al. FRI 41309 (K,

SING), Zainudin 5029 (K, SAN), Cheksum Tawan CST 285, Perumal &

Sundaling SAN 135176 (K, SAN), all from the same locality of the type.

3. Tristaniopsis microcarpa P.S.Ashton, sp. nov.

T. merguensis similis foliis maioribus haud auriculatis, floribus minoribus

staminibus paucis fructibus minimis differt.Typus: H. S. Martyn SAN 2/623,

Look Mengulang, P. Sakar, in flower (holo K; iso KEP, SAN, SAR, SING).

Canopy tree to 30 m tall, to 80 cm diameter. Bark at first smooth greenish

to yellow- or rust-brown, later exfoliating in large grey-brown scroll-like

strips, eventually becoming shaggy towards base. Living parts shortly greyish

puberulent, more or less persistent on cyme and exposed parts of flower

and fruit, elsewhere early glabrescent. Twig c. 5 mm diam., round, stout,

drying blackish at endings, becoming grey brown thinly peeling. Leaf blade

(9—)13—27 x (3—)5—10 cm, oblanceolate, thinly leathery, minutely densely

and larger sparsely dotted beneath, pitted above, drying dull olive-brown

darker above; subsessile or occasionally tapering into c. 15 mm long stalk;

apex shortly broadly acuminate or blunt; main lateral veins 25—30 pairs

with variably distinct intermediates, dense, spreading, raised throughout

more so beneath; tertiary veins lax distinct. Cyme to 15 cm long, to 3 mm

274 Gard. Bull. Singapore 57 (2005)

diam. at base, elliptic in section, slender, to 4-branched, with to 8 cm long

erect rachis. Flower to 3 x 3 mm, small; stamens 3(—4) in each cluster.

Capsule to 6 x 4 mm, to 6 mm wide when open, 3-celled; seeds many,

small.

Distribution and ecology: Known from Sabah where it is widespread, NE

Sarawak to the Rejang valley, and E and C. Kalimantan to the Schwaner

Range. Locally frequent in mixed dipterocarp forest to 1000 m, on clay

and sandy clay soils over both sedimentary, and also ultramafic rocks where

it is sometimes common.

Other specimens examined: SABAH — Sigin & Rahim SAN 99694 (K,

SAN, SING), Bukit Mensasau Beluran; Madani SAN 130694 (K, SAN),

Bidu-Bidu F.R., Beluran; Amin G. et al. SAN 93896 (K, SAN, SING),

Kiabau, Beluran; Amin G. SAN 70315 (K, SAN), Telupid; Davol, Pius et

al. SAN 124652 (K, SAN), S. Kim, Tankulap, Telupid; P.S. Shim SAN

134716 (K, SAN, SING), Bukit Tingka, Telupid; Leopold & Taha SAN

83534 (K, SING), Kodoh & Tarmijin SAN 83666 (K, SAN, SING), M.87

Telupid Rd., Sandakan; Meijer SAN 53317 (K, SAN, SING), Sungai Meliau,

Karamuak, Sandakan; Madani SAN 81176 (K, SING), Bintang Mas logging

area, Karamuak; A. Gibot SAN 36069 (K, SAN), Singh & Talip SAN

52602 (K, SAN, SING), A. Gibot SAN 36025 (K, SAN), Pulau Sakar;

Muin Chai SAN 26995 (K, SAN, SING), Muin Chai SAN 26655 (K, SAN,

SING), H.S. Martyn SAN 21613 (K), Look Mengulang, P. Sakar; Joseph

B. et al. SAN 120799 (K, SAN), Km 24 Taliwas, Lahad Datu. SARAWAK

— §. Tong S 32801 (K, SAN), Ulu Masia, Meligan range, Lawas; Yahud et

al. S 76338 (K, SAN), Sungai Kelepang, Remudu, Bario; P.J. Martin S

38913 (K, SAN), Gunung Mulu N.P.; Othman et al. S 41468 (K, SAN), Ulu

Balleh; Othman et al. S 62110 (K, SAN), Nanga Sebatu, Mengiong, Balleh,

Kapit. C. and W. KALIMANTAN -— Argent & Wilkie 943 (K, SAN), Km

48 from Sangai, Sungai Mentoya, Kota Waringin Timor; Jarvis & Ruskandi

6212 (K), Bukit Raka, Bukit Raya N. P.

Notes: This species is similar in leaf to Tristaniopsis merguensis, but the

leaf is larger, never auriculate at base even in juveniles; the flowers are

smaller with fewer stamens, and the fruit is much smaller.

4. Tristaniopsis rubiginosa S.Teo ex P.S.Ashton, sp. nov.

Species praestans foliis magnis subtus prominenti fusco-brunneo

puberulentibus staminibus 3—4 in fasculo facile distinguitur. Typus:

Purseglove 5053, fruit, Telok Asam Bako N.P. (holo K; iso SAR, SING).

Tristaniopsis in Borneo 275

Small tree to 7 m tall, to 15 cm diam., with open prominently chocolate-

brown-leaved crown. Bark pale grey and mauve peeling and scroll-marked;

inner bark whitish. Sapwood dark tallow-brown. Young parts densely dark

warm brown pubescent, caducous successively on leaf blade above, blade

beneath, leaf stalk, twig, cyme and floral receptacle, more or less persisting

into fruit. Twig c. 5 x 4 mm apically, somewhat flattened at first becoming

round, smooth, blackish when dry, eventually thinly flaking. Leaf blade

10-17(—28) x 5—9 cm, broadly elliptic to obovate, thickly leathery, more or

less persistently dark chocolate-brown pubescent beneath fading to mauve

in fallen leaves, drying dull mauve-brown above, densely pitted above,

dots obscure beneath; margin inrolled; base wedge-shaped narrowly tapering

down sides of 10-17 mm long stout stalk; apex to 1 cm long broadly

acuminate or blunt rounded; main veins c. 26 pairs with unequally dispersed

more or less equal intermediate veins, spreading, slender but distinctly

raised beneath, visible but hardly raised above; tertiary veins obscure, lax.

Cyme to 12 cm long, terminal or subterminal-axillary, to 3-branched, drying

flattened and ribbed. Flower bud to 5 x 3 mm, relatively large, ellipsoid,

receptacle obconical; stamens 3-4 per cluster. Capsule to 5 x 4 mm, to 7

mm diam. following dehiscence; seeds many, small.

Distribution and ecology: Endemic to northern Borneo. Very local, but at

Bako National Park abundant, on podsols in kerangas on sandstone plateaux

and raised beaches, and in the open ‘padang’ vegetation on the tableland,

and on ultramafic substrate in Sabah, at low altitude.

Other specimens examined: SARAWAK — Sipun Dominic & Dami S 81530

(K, SAN), Chai & Ilias S 17868 (K, SAN, SING), Corner & Brunig S

10495 (K), Brunig S 10408 (K), Anderson & Ashton S 12341 (K, SING),

Sinclair & Kadim SFN 10315, Bako N.P., Kuching. SABAH — Saw L.G.

FRI 36238 (K, SAN), Meliau Basin. BRUNEI — Ashton BRUN 647 (K,

SING), Bukit Puan.

5. Tristantopsis whitiana Peter G.Wilson & J.T.Waterh.

Austr. J. Bot. 30 (1982) 440. Basionym: Tristania whitiana Griff., Pl. Cantor.

(1837) 18. Type: White s.n., Singapore (CAL?, not seen). Synonym: Tristania

sumatrana Migq., Fl. Ind. Bat., Suppl. (1861) 308.

Subspecies whitiana

Canopy, occasionally shortly emergent tree to 45 m high, to 1.5 m diam.,

276 Gard. Bull. Singapore 57 (2005)

with concave rounded buttresses. Bark at first white to light greenish grey

with occasional hint of very pale orange, smooth, later peeling in scroll-

like strips; older unpeeled bark evenly pale grey-dull light olive; peeled

bark scrolls pale to dark grey with mauve-brown patches; inner bark whitish.

Sapwood yellowish. Leaf beneath, rachis, flower bud, fruit sparsely or

densely more or less persistently grey-brown puberulent, or sometimes

glabrous. Twigs c. 2 mm thick apically, slender, round, glabrous, smooth

eventually thinly peeling. Leaves shiny fresh green when alive, drying rich

dark olive-brown glistening beneath, oblong to lanceolate, (7.3—)12.9(—26)

x (2.5—)4(—5.5) cm, thinly leathery; base narrowly wedge-shaped, stalk

distinct, (0.5—)0.8(—1.2) cm long, slender; apex sharply acute; main lateral

veins subequal, (54—)68(—92) pairs, very many, dense, slender, slightly but

distinctly raised above and below; intramarginal vein to 1 mm within margin.

Cyme to 10 cm long, to 5-branched, with long slender rachis. Flower cream

with yellow stamens; bud c. 1.5 x 1 mm; pedicel c. 1 mm long; calyx minutely

warty, unribbed, lobes c. 0.5 x 1 mm; petals c. 1.5 mm long; stamens 3 per

cluster opposite each petal, filament c. 1.5 mm long, anther c. 0.2 mm.

Capsule c. 4 x 3.5 mm, ellipsoid; seeds many, c. 0.4 x 0.2 mm, elliptic.

Distribution and ecology: Sumatra, Peninsular Malaysia (including

Singapore) and Borneo. More abundant in Sarawak and SW Sabah than

elsewhere in Sabah. The paradigmatic late successional tree of landslips on

the steep inland hills. Locally abundant also in secondary forest and river

banks from the lowland to upper dipterocarp forest, to 1500 m on Mt.

Kinabalu; generally in dipterocarp forests on mostly clay soils, but on

sandy soils and the transition to kerangas in W. Sarawak.

Other Borneo specimens examined: SABAH — Lowe s.n. (K) ‘Borneo’;

Woods 31 (K), loc. incert.; Orolfo SAN 5488 (K, SING), Ulu Tawau; Orolfo

SAWN 23 (K, SING), Silimpopon R., Sta. Lucia (= Tawau) For. Dist.; Meijer

SAN 19526 (K, SING), Serudong, Tawau; Leopold & Taha SAN 83584 (K,

SAN), M 87!/2 Telupid road, Sandakan; Brand SAN 30876 (K, SAN, SING),

Berambangan, Kudat; A. Gibot SAN 79604 (K, SAN, SING), Kinabalu

N.P.; Saikeh SAN 72244 (K, SAN, SING), Beaufort; Dewol & Karim SAN

77725 (K, SAN, SING), Lumaku F.R., Sipitang. SARAWAK — Beccari

2773 (K), 2916 (K), loc. incert., Chai S 39731 (K, SAN), Sungai Mentawei

Mulu N.P.; Anderson S 4219 (K, SAN, SING), Melinau Paku path Gunung

Mulu; S. Tong S 34929 (K, SAN), Sungai Koyan Mengak, Belaga; Richards

2459 (K), Long Kapa, Dulit, Tinjar; Jacobs 5399 (K), Belaga, Daud &

Tachun SFN 35672 (K, SING), Nanga Pelagus, Kapit; Ilias S 25783 (K,

SING), Bukit Pantu, Kapit; Othman Ismawi S 40049 (K), Semengoh

arboretum, Kuching; Dg. Awa & Othman Ismawi S 47100 (K, SAN), Teluk

Tristaniopsis in Borneo yi

Bandung, Santubong; Sinclair & Kadim SFN 10394 (K, SING), Sempadi

F.R., Lundu. BRUNEI - Ashton & Whitmore BRUN 485 (K, SING),

Bangar; Ashton BRUN 881 (K), Ulu Tutong at first rapid.

Subspecies monostemon P.S.Ashton, ssp. nov.

A species typico floribus staminibus solo 5 differt. Typus: Dan S 3033,

Lambir Hills, Sarawak (holo K; iso SAN, SING).

Differing from the type subspecies as follows: Tree at most 25 m tall, 30

cm diam.; bark brilliant coppery-brown; flower with a ring of 5 single

stamens.

Distribution and ecology: Known from throughout Sarawak, Brunei, and

W. and C. Kalimantan. Locally gregarious; on banks, usually on sandstone

rocks, by white and black water rivers, in kerangas forest.

Other collections examined: SARAWAK -— B. Lee § 46502 (K, SAN); Ilias

S 15452 (K, SAN, SING), Sungai Sebiau, Bintulu; Ashton S 16457 (K,

SAN, SING), Sungai Jilai, Tatau; Jugah S 15224 (K, SAN, SING), Sungai

Sabal Tapang, Serian; S. Teo S 75432 (K, SAN), Matang Wildlife Centre;

Jugah § 51588 (K, SAN), Sungai Raya, Division I; Enjah et al. S 75426 (K,

SAN), Pueh F.R. BRUNEI — Niga et al. BRUN 15109 (K, SAN, SING),

Sungai Lumut, Belait; Coode 7102 (K, SAN, SING), Ulu Tutong; Ashton

BRUN 3303 (K, SING), Bukit Patoi. KALIMANTAN — Awmack 19 (K),

W. bank of R. Rekut, C. Kalimantan; Church et al. 1745 (K, SAN), 8 m

north of Desa Jelundung, Serawai, W. Kalimantan.

Notes: The single stamen opposite each petal is diagnostic. The tree is

conspicuously different from the type on account of its vivid bark colour,

but more collections in flower are needed to confirm whether bark and

stamen characteristics are always correlated.

Acknowledgements

Stephen Teo, formerly of the Sarawak Forest Department Herbarium,

early recognized many of the taxa, and prepared draft descriptions upon

which the present descriptions are partially based. The work is otherwise

based on collections at the Royal Botanic Gardens, Kew and Harvard

University herbaria. Eve Lucas at Kew and Emily Wood at Harvard have

been generous with their counsel. Staff at KEP, SAN, SAR and SING

provided information on specimens in their care.

278 Gard. Bull. Singapore 57 (2005)

Reference

Kochummen, K.M. 1978. Tristania. Tree Flora of Malaya. 3: 251-253.

Gardens’ Bulletin Singapore 57 (2005) 279-283 DD

Musa lokok (Musaceae), a New Species of

Banana from Bario, Borneo

CONNIE GERI

AND

FRANCIS S. P. NG

Sarawak Biodiversity Centre,

KM 20, Jalan Borneo Height, Semenggoh, =

Locked Bag 3032, 93990 Kuching Sarawak, Malaysia

Abstract

Musa lokok Geri & Ng, sp. nov. is described and illustrated. Locally, the young unfurled

leaves have been used as cigarette wrappers.

Musa lokok

Musa lokok Geri & Ng, sp. nov.

Musa suratiorum similis sed folius subtus non glauco-ceraceus et fructis

pendentis differt. Typus: Sarawak: Marudi District, Bario, Pa’Lungan C.

Geri & Pasen Paran TK1552, 16 Aug 2005 (holo SAR, iso KEP, E, SBC).

Plate 1

Clump forming herbaceous plant to c. 3 m tall to the tip of the leaves.

Suckers emerging from below ground level from the corm with up to 6

rhizomes radiating horizontally, pseudostems spaced 10—30 cm apart,

mature pseudostem 1.3—2 m tall, slender about 2.5 cm diam., gradually

swollen to 4 cm diam. at the base, covered with brown fibrous persistent

old leaf sheaths, emergent leaf sheaths yellowish green. Sap watery. Upper

margin of sheaths (shoulder) smooth. Fourth to last leaf: petiole about 30 x

1 cm, yellowish green, petiole closed almost to the lamina, the channel

about one third of the depth of the petiole, lamina 120-126 x 21-24 cm,

narrowly elliptic, broadest about the middle, apex acute, base unequal,

cuneate, not waxy underneath, light green above and paler underneath.

Inflorescence hanging vertically downwards. Peduncle glabrous, smooth

and without wax, at first white becoming black.} Female buds c. 37 x 3 cm,

bracts bright purple outside, white inside, lifting to 90°, straight not curling

back. First bract sterile, subsequent bracts with 1—3 flowers. Female flowers

c. 10 cm long, ovary pale cream, the compound tepal deep green, c. 5 cm

280 Gard. Bull. Singapore 57 (2005)

long, apically divided into 3 lobes c. 9 mm long, the free tepal c. 4 cm long,

translucent white, wrinkled along the margin, staminodes 5, up to 1 cm

long, style pale greenish cream, style and stigma c. 4 cm long, stigma

glabrous, pure white. Fruit bunch hanging vertically down, the fruit at

about 30° to the rachis, about 3 hands each with 1—3 bananas arranged in

one row. Fruits ripening pale yellowish green, not splitting open, ovules in

2 rows per loculus. Pedicel c. 2.5 cm long, the fruit 6-8 cm long, 1.5—1.7 cm

diam., straight, cylindrical, angled, contracted at the apex into a short ‘beak’.

Seeds black, barrel-shaped, c. 5 x 3 mm, at the inner end with a large

circular white patch, rough with irregular very low tubercles. Male axis

black, bract scars c. 1.5 cm apart on the same rank. Male buds slender,

oblong-ovoid, c. 22 x 3.3 cm. Unopened bracts white beneath the flowers,

rosy purple, slightly darker at the tip, straight, not curling back, glossy.

Male flowers c. 4.3 cm long, compound tepal translucent pale yellowish

green but white at the base, apically divided into 3 lobes c. 4 mm long, the

middle lobe divided into 3 smaller lobes, free tepal c. 3 cm long, white and

translucent, 3-pointed, stamens 5, pure white, c. 3 cm long, anther about 3/

4 of the stamen length, style slender, white, style and stigma c. 3.5 cm long,

longer than the stamens.

Other specimen: Sarawak: Marudi District, Bario, Pa’Lungan C. Geri &

Pasen Paran TK1553, 16 Aug 2005. (SAR, SING, SAN, E).

Distribution: Endemic to Borneo, known only from Bario in the Marudi

District.

Habitat: Swampy depressions in undisturbed forest at about 1167-1300 m

altitude

Vernacular name: Lokok (Kelabit) and Sukar (Kelabit).

Local people informed us that the young unfurled leaves were formerly

used as cigarette wrappers. The name ‘lokok’ appears to be a variation of

‘rokok’, the Malay word for cigarette. An elderly woman told us the original

Kelabit name is ‘sukar’ because compared with other bananas, the leaves

of this species are less liable to split and become tatty. Christensen (2002)

refers to the cigarette-wrapper banana of the Kelabits as ‘lukor’. We have

sometimes heard ‘lokok’ pronounced as ‘lukor’.

Notes: Musa lokok is unusual among Bornean bananas based on the

combination of the following characters: it is a small slender banana with a

narrowly elliptic leaf about 120-125 x 20-25 cm with an inflorescence

hanging vertically down, the male bud is slender and spindle-shaped and

Musa lokok from Borneo

Plate 1: Musa lokok Geri & Ng a. Clump showing the pendent inflorescence, b. female

inflorescence removed from the pseudostem, c. male flower, d. female flower, e. L.S. of fruit.

Photographs FSP Ng

282 Gard. Bull. Singapore 57 (2005)

up to 3 cm in diameter, neither the male or female bracts curl back.

Based on these characteristics, it resembles Musa suratii Argent from the

Tenom District, Sabah, which was also recently discovered in Usun Apau,

Baram District (S. Julia, pers. comm.). Like M. lokok, it is a highland

species growing at about 1000 m altitude. However, it is distinct from

Musa suratii in the characters noted in Table 1. Notable differences include

the absence of wax from the lower surface of the leaf and the bananas

pointing obliquely downwards.

Table 1. Morphological differences between Musa suratii and M. lokok

Musa suratii M. lokok

Petiole length (cm) 16—20 c. 30

Lower leaf surface Thick white wax No wax

Basal flowers Hermaphrodite, with 5 Female with 5 small

stamens ranging from staminodes

fully developed with

apparently fertile pollen

to tiny staminodes

Female with 5 small

staminodes

Fruit bunch With 8—10 hands of With 3—5 hands of

fruits, bearing 8—10 fruits, bearing 1—7

bananas in two rows in bananas in each

the second hand, hand, pointing

reflexed upwards obliquely downwards

Fruit diameter (mm) 8—10 15—17

Seed (mm) 25532 SS aS.

Male bract colour Orange with green tip Bright purple

The clump that T7K1/553 was collected from was somewhat more

variable, the leaves were 118—150 x 17—29 cm, the inflorescence had the

first one or two bracts sterile and bearing up to 5 hands of bananas with 3—

7 bananas per hand.

One population appeared to be a hybrid between Musa lokok and

M. textilis Nees because the male bud was broader, more like that of M.

textilis, and, in addition, it had a conspicuous green tip to the male bracts.

Acknowledgements

This work was carried out as part of the Traditional Knowledge

Documentation Project of Sarawak Biodiversity Center. We thank Datin

Musa lokok from Borneo 283

Eileen Yen Ee Lee for her strong support in the project and Pasan Paren

for his assistance and help in the collection of specimens. We should also

like to thank Dr Ruth Kiew for her advice in describing this species.

Reference

Christensen H. 2002 Ethnobotany of the Iban and the Kelabit. Forest

Department Sarawak, Malaysia and University of Aarhus, Denmark.

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Gardens’ Bulletin Singapore 57 (2005) 285—287 285

Dipterocarpus tempehes (Dipterocarpaceae) —

A New Record for Singapore

s. LEE AND AIDI ABDUL GHANI

National Parks Board,

Singapore Botanic Gardens, Cluny Road,

Singapore 259569

The extreme dry weather conditions in the early part of this year triggered

mast flowering of many trees in Singapore. Dipterocarps, many of which

are known to flower heavily once in 5—10 years, produced flowers by the

millions in many parts of Singapore. These particular trees were previously

mistaken for Dipterocarpus sublamellatus Foxw. because of their similar

leaves. When conducting regular surveys, in June 2005 the National Parks

Board staff of the Central Catchment Nature Reserve, came across a strange

and wingless dipterocarp fruit along MacRitchie Nature Trail.

On consulting the literature and the herbarium collection of the

Singapore Herbarium (SING), these specimens (MacRitchie Nature Trail

- Aidi A.G. SING2005-246 29 June 2005 (SING), SING 2005-259 11 July

2005 (SING); Lua H.K. SING2005-290 16 July 2005 (SING)) appeared to

be identical with Dipterocarpus tempehes Sloot. P.S. Ashton (pers. comm.)

confirmed its identity. However, according to Newman et al. (1995) and

Symington (2004), this species has never been recorded from Singapore or

Peninsular Malaysia, respectively.

Prior to this find, Dipterocarpus tempehes was known to be endemic

only in Borneo (Ashton, 1982, 2004; Meijer and Wood, 1964; Newman et

al., 1998), where it is found in periodic swamps and along stream banks.

This is a new significant record for Singapore. In total, there are

five mature trees towering above 35 m just off the nature trail. Similar in

morphology to the Bornean ones, its habitat in Singapore is different. It is

possible that the MacRitchie Nature Trail could have been swampy in the

past, judging by the species composition of the primary forest patch near

the streamlet. However, the area is now by and large moist and gently

sloping, with no periodic inundations.

Acknowledgements

Chew P.T. and Lua H.K. are thanked for providing information on the

ecology of the area, P.S. Ashton for promptly confirming its identity and

J.J. Vermeulen for drawing the botanical illustration.

286 Gard. Bull. Singapore 57 (2005)

a ij 5.cm

Figure 1. Dipterocarpus tempehes Sloot. a. fruit; b. leafy shoots with old infructescence. (from

Aidi SING2005-259).

Dipterocarpus tempehes in Singapore 261

References

Ashton, P.S. 1982. Dipterocarpus tempehes. Flora Malesiana, series I. 9:

297.

Ashton, P.S. 2004. Dipterocarpus tempehes. Tree Flora of Sabah and

Sarawak. 5: 125.

Meier, W. and G.H.S. Wood. 1964. Dipterocarps of Sabah. Sabah Forest

Record. 5: 270.

Newman, M.F., P.F. Burgess and T.C. Whitmore. 1995. Manuals of

Dipterocarps for Foresters — Singapore. Black Bear Press Ltd, Cambridge.

Newman, M.F., P.F. Burgess and T.C. Whitmore. 1998. Manuals of

Dipterocarps for Foresters — Borneo Island Medium and Heavy

Hardwoods. Black Bear Press Ltd, Cambridge.

Symington, C.F. 2004. Foresters’ Manual of Dipterocarps.’ Malayan Forest

Records 16.

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Gardens’ Bulletin Singapore 57 (2005) 289-290 289

Book Review: Christian Puff, Kongkanda Chayamarit and Voradol

Chamchumroon. 2001. Rubiaceae of Thailand. A Pictorial Guide to

Indigenous and Cultivated Genera. Forest Herbarium, National Parks,

Wildlife & Plant Conservation Department, Bangkok, Thailand. viii + 245

pp. ISBN 974-463-142-2. Price: EURO 40 or USD 50 (inclusive of airmail).

The Rubiaceae comprise one of the largest families of flowering plants in

the Southeast Asian tropics, and are therefore important in the Thai flora,

including about 110 genera and some 600 species. There are few consistent

workers in the family in Southeast Asia, because an introduction to such a

diverse array of groups and form is not an easy thing to grasp or teach, so

that collectors and botanists have had few guides to go by. The many taxa

that are either rare or restricted in occurrence are frequently incompletely

or little documented. Thus, the Rubiaceae, like other large and diverse

families, have attracted few students so far, and have languished among

the so-called “difficult and perhaps untidy families”.

The appearance of this book gives hope that all this is about to

change. With more than 20 years of experience in studying Thai and

Southeast Asian Rubiaceae, Christian Puff, joined by co-workers

Kongkanda Chayamarit and Voradol Chamchumroon who provide the

Thai text to complement the main English text, now present a well

organized account that is a truly effective guide to this family. As given in

the subtitle, the book is meant to introduce the family botanically, in terms

of its diversity of life form, vegetative and reproductive structure and

biological attributes such as pollination biology and dispersal. This

indispensable introduction is covered in the first two chapters of the book

(General notes; Selected character states), which throughout is lavishly

illustrated by colour photography and complemented by easy-to-understand

diagrams. |

The indigenous genera of Thailand are treated in the main part,

forming Chapter 3, where the arrangement by life form (trees/treelets/

shrubs; climbers; epiphytes and ant-plants; herbs) makes for a very user-

friendly introduction. The photography is actually splendid, mouth-

wateringly attractive in many cases. The last portion, Chapter 4, deals with

the non-indigenous taxa, grouped as ornamentals and cash crops, again

very informative. Most (more than 80) Thai genera are covered in this

book, and there is nearly always a single page of text facing a plate of well-

chosen illustrations of the genus concerned (exceptionally, genera such as

Psychotria, Morinda, Prismatomeris, Hydnophytum or Argostemma, where

variation in form and biology may be of particular interest, are given four

full pages of treatment).

The account is more than just a potential novice’s tool par excellence.

290 Gard. Bull. Singapore 57 (2005)

It also gives botanists specializing in other groups and other regions a

highly accurate interpretation of the many forms expressing the fascinating

diversity of plants encountered in the tropical forest in general. From the

peculiar hooked branches of the climbing Oxyceros, the near-recumbent

sprawling shrubby habit of Gardenia saxatilis, to the bizarre ant-inhabited

stem-base tubers of Hydnophytum and Myrmecodia epiphytes, and

herbaceous Argostemma species that have but a single conspicuous (in

fact, somewhat oversized) leaf developed, the array represents an instructive

panorama of plant form. Reproductive structure is equally varied and key

types are often the basis of group recognition: syncarps formed by fruit

fusion such as in Morinda, the peculiar cymes of Mouretia where scorpioid

branches are broadly flattened and bear four rows of flowers fused together

by their ovaries, and the show-calyces of Mussaenda and Schizomussaenda,

are but some of the many interesting variations of infructescence and

inflorescence form there can be in this large family.

The diversity of the Rubiaceae, as in any large and widely distributed

family, is easy to appreciate in the context of radiation and adaptation in

different directions, in response to climate, geography, soils or other

ecological factors. Thailand covers an interesting range of climatic and

floristic belts, from deciduous, seasonally dry forest in the north to wetter

evergreen forest in the Thai peninsula, from the Myanmar and Indo-Chinese

regions to the Malesian floristic region to its south, and includes influences

from both temperate and tropical regions. Within, it has landform and

vegetation from beaches to high mountains, and peatswamps to karst

limestone.

This book is highly recommended to all interested in the tropical

flora and represents an indispensable introduction for those wandering—by

choice or chance—into the fold of the Rubiaceae.

K.M. Wong

Institute of Biological Sciences,

University of Malaya,

Kuala Lumpur

Gardens’ Bulletin Singapore 57 (2005) 291—293 veh

OBITUARY

Arthur George Alphonso

A deep love of plants led Arthur George Alphonso to pursue studies

at the Serdang Agricultural College and thence to his appointment as

Horticultural Assistant with the Singapore Botanic Gardens on 1 January

1940. In 1954, the Gardens sponsored him for a two-year training course at

the Royal Botanic Gardens, Kews. He returned to assume the position of

Gardens’ Curator. George Alphonso developed a rewarding lifelong career

with the Gardens, rising through the ranks to be its chief administrator

before his retirement in 1976.

Upon his first appointment, he worked in the grounds, but during

the WW II Japanese occupation of Singapore, was placed in charge of the

potting yard. This was essentially the nursery and technical support area

for the Gardens. Early experimental research and simple planting trials

were carried out in the potting yard, with one of the mainstays being plant

introduction. An estimated 80% of the plant species cultivated for roadside

planting and in home gardens in the 1960s were introduced through the

efforts of the Gardens. This was actively built up by George through an

active plant exchange programme with other botanical institutions and

personal contacts that he had established all over the world. Gardening

enthusiasts, especially from the Australian, African and South American

tropics, often stopped by the Gardens to share information and exciting

new plant material they had discovered.

The Gardens’ collections relied on and grew steadily with this

goodwill exchange programme. There was scarce funding for things

botanical and horticultural in those times, as the nation had more pressing

issues to address. As the collections grew and were successfully propagated,

the Gardens found itself with surplus plant material, which could be shared

with the gardening community in Singapore, and a plant sales nursery was

established for the public. Gardening advice was frequently sought from

the Gardens’ staff and as Curator, George was the main contact for the

public. He was acknowledged as Singapore’s expert in the field. When

Singapore embarked on its enlightened greening programme and its Garden

City Campaign, George Alphonso was among the pioneers instrumental in

its successful implementation. Hundreds of thousands of planting material

was produced by the Gardens and supplied gratis to government

departments, schools, charitable institutions and other official organizations

for Singapore’s initial greening efforts.

Interest in gardening had always featured significantly among

292 Gard. Bull. Singapore 57 (2005)

George Alphonso with Queen Elizabeth II when she visited the Gardens in 1972. A VIP

orchid, Dendrobium Elizabeth, a hybrid developed by the Singapore Botanic Gardens, was

named in her honour.

Singaporeans. Those who could afford gardens were always keen to add

new species or hybrids to their collection; others flocked to flower shows

to admire the landscapes and prize blooms proudly displayed by the growers.

Annual flower shows were eagerly anticipated, and the Singapore Botanic

Gardens actively co-ordinated these shows, with George Alphonso taking

on the role of Show Manager, and subsequently Judge, for many decades.

The staging of the shows wrought a close bonding and camaraderie among

the growers, hobbyists and professionals and inculcated an appreciation of

plants in the visitors. George was active in the Singapore Gardening Society

and the Malayan Orchid Society, Singapore Branch, where he served as

both President and Editor of the Malayan Orchid Review. He was kept

busy especially when there was both an Orchid and a Garden Show held in

the same year.

George’s professional interest in orchids grew progressively from

the early fifties. Expeditions to the surrounding forests brought him into

close proximity with this fascinating family of plants, and his interest could

only grow as he gained more exposure to the range of forms and natural

habitat of the plants. Orchid hybridisation was already an important aspect

of research in the Gardens, with the precedence set by such eminent

293

botanists as RE Holttum and GH Addison. George Alphonso sustained

the Gardens’ special interest and furthered its reputation as a world famous

garden for the collection and conservation of orchid species and for

producing many award winning hybrids. He forged close friendships with

other orchid enthusiasts at local and international events and became much

sought after as a judge at orchid shows as his name and reputation grew

entwined with that of the Singapore orchid world. George’s counsel was

elicited when Singapore decided to have a national flower, and his advised

choice of Vanda Miss Joaquim was accepted.

George Alphonso’s long tenure with the Gardens will be fondly

remembered by those who have worked with him. His was an era when

the grounds were inspected on foot and hands-on advice was shared with

the staff. His world was that of open skies and field, and he had little

patience with air-conditioned meeting rooms. He felt that the world of

plants and the solutions to their problems were best dealt with where they

thrives — outdoors.

Geh Siew Yin,

Singapore

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INSTRUCTIONS TO AUTHORS

The Gardens’ Bulletin publishes original research findings and reviews of progress in the fields

of plant taxonomy, horticulture and allied subjects. Contributions must be original and the material

must not have been submitted for publication elsewhere.

Authors should look at the layout of articles recently published in the journal to ensure that

submitted manuscripts conform as closely as possible to the accepted format. Particular care

should be taken with the format of the references. Manuscripts may be submitted in electronic

form (PC-compatible form) together with a hardcopy and original drawings and illustrations as

appropriate.

Titles and authors: The title should give a concise description of the contents of the article. It

should include the family name if a taxon name is included in the title. The names(s) and

affiliation(s) of the author(s) must be given below the title. A short running title should also be

provided. Lengthy papers must have contents listed at the beginning of the paper. Avoid footnotes.

Abstract: An abstract of up to about 100 to 200 words should be provided. It should

comprehensively summarise the contents of the article as it is likely to be reproduced without the

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Scientific names: The complete scientific name - genus, species, authority with family in

parenthesis —- must be cited for every organism at the time of first mention.

Abbreviations: Standard abbreviations may be used in the text, but the full term should be given

on the first mention. Dates should be cited as: 1 Jan 2000. Units of measurement should be

spelled out except when preceded by a numeral where they should be abbreviated in standard

form: g, ml, km, etc. and not followed by stops.

Tables: All tables should be numbered and carry a heading with their content. These should be

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should be used to indicate magnification. Provide a photocopy of the illustrations to indicate the

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When grouping photographs, the page size of the journal must be taken into account to

optimize the space. Colour photographs should only be included where colour adds significantly

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For figures including photographs, type the captions in numerical order on a

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Literature citation: Citation in the text should take the form: King and Gamble (1886). If several

papers by the same author in the same year are cited, they should be lettered in sequence, 2000a,

2000b, etc. When papers are by three or more authors they should be cited as King et al. (1886)

in the text, but with all the authors’ names given in the reference section. All references must be

placed in alphabetic order according to the family name of the first author. The journal title must

be given in full, as in the following example:

Stone, B.C. 1994. Additional notes on the genus Glycosmis (Rutaceae). Gardens’ Bulletin

Singapore. 46: 113-119.

References to books and monographs should be cited according to the following form:

Ridley, H.N. 1930. The Dispersal of Plants Throughout the World. L. Reeve, Ashford, U.K.

For literature citations in taxonomic papers, the following style is required:

Medinilla alternifolia Blume, Mus. Bot. Ludg.-Bat. I:1 (1849) 19.

Sterculia acuminatissima Merr., Philip. J. Sci. 21 (1922) 524.

Offprints: Authors will be given 50 offprints gratis. Additional copies must be ordered and paid

for in advance of publication.

Manuscripts should be sent to: The Editor, Gardens’ Bulletin Singapore, The Singapore Botanic

Gardens, Cluny Road, Singapore 259569.

<N é

Field Guide to the Grasses of Singapore

(Excluding the Bamboos)

Helena Duistermaat

‘ ay hal ;

is :

2 Supplement of The Gardens' Bulletin Singapore

Volume 57, 2005 ISSN 0374-7859

THE GARDENS’ BULLETIN SINGAPORE

The Gardens’ Bulletin Singapore publishes papers on plant taxonomy (including

revisions), horticulture, phytogeography, floristics, morphology, anatomy and

related fields with emphasis on plants in the West Malesian region.

EDITORIAL BOARD

Dr R. Kiew Dr R.T. Corlett

(Editor) University of Hong Kong

Singapore Botanic Gardens Hong Kong

Dr P.Y. Tan Dr M.C. Roos

(Assist. Editor)

Singapore Botanic Gardens

Dr S.C. Chin

Singapore Botanic Gardens

Dr. M.J.E. Coode

Royal Botanic Gardens,

Kew, U.K.

National Herbarium of the Netherlands

Leiden University,

Leiden, The Netherlands

Dr E. Soepadmo

Forest Research Institute Malaysia

Kepong, Malaysia

Dr W.K. Tan

National Parks Board, Singapore

The Gardens’ Bulletin is published twice yearly by the National Parks Board,

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lbw

Field Guide to the Grasses of Singapore

(Excluding the Bamboos)

HELENA DUISTERMAAT

The Herbarium, Singapore Botanic Gardens,

1 Cluny Road, Singapore 259569

Drawings by

Jaap J. Vermeulen

Sections

Introduction

Grass Morphology

Making a Reference Collection of Grasses

Turf Grasses

Bamboos and Grasses

Grasses and Grass Look-alikes

Glossary

Key to the Flowering Grasses of Singapore

Descriptions and Generic Keys

Key to Sterile Grass Plants of Singapore

Literature

Acknowledgements

Index

Plates

142

151

LSS

157

166

Supplement of The Gardens’ Bulletin Singapore

Nol. 3/7, 2002

Date of Publication: 2 March 2005

Published by

National Parks Board

Singapore Botanic Gardens

1 Cluny Road, Singapore 259569

Printed by KaroCraft Pte Ltd

ISSN 0374-7859

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Supplement to Gardens’ Bulletin Singapore 57 (2005) 1-177 ]

Introduction

Grasses are generally neglected by the layman because they all look the same

and do not have obvious flowers. However, the family of the grasses, Gramineae

(also called Poaceae), is worthy of attention for a number of reasons. First,

they are of huge economic importance. They are widely used for food (cereals,

fodder), ornamentals (including lawns) and construction (bamboos), while other

members of the family are known as the world’s worst weeds. The second

reason is that grasses are very abundant. Not only in species number is the

grass family one of the largest of the world (with about 10,000 species); but

they also cover a vast area, approximately 20% of the earth’s land surface,

ranging from tropical to arctic regions. They occupy a wide variety of habitats

but they are most abundant in open habitats.

The original vegetation of Singapore consisted primarily of forests

(mainly lowland dipterocarp forest) where only a few specialized forest-grass

species in low densities occurred. In the permanently open habitats such as

beaches, a few other grass species probably grew. (For a description of the

original vegetation and the impact of human activities see Corlett 1992, 1993).

Large-scale clearance of forest in Singapore started only after the British arrived

in 1819. By the 1880s only 7% of the original primary forest was left. Today,

99.8% of the original forest cover has been cleared or severely disturbed. Rivers

and even forest streamlets have been canalized or replaced by concrete drains

and many of the original beaches have been replaced by reclaimed land.

A considerable number of the native grass species are now endangered

or extinct. Ridley (1907) noticed that thé majority of grass species of Malaya

(Peninsular Malaysia and Singapore) either occurred on sandy spots by the

sea or as weeds. At present, grasses are found mainly in unnatural open habitats,

such as turf, roadsides and wastelands. Turfed areas (including more than 1000

ha of golf courses) alone occupy about 18% of Singapore’s 574 km/ of land

surface (Corlett 1992: 415). From this it will be clear that grasses are indeed

an important part of Singapore’s present day flora.

Ridley (1907) provided the first account of the grass family for Malaya,

followed much later by Gilliland (1971). Recently, Keng et al. (1998)

summarised the family for Singapore. Recent new records are discussed in

Duistermaat (2004). The keys provided in all works (in the latest flora treatment

unfortunately not to the species level) are very technical. Also, some quite

common species rarely seem to flower in Singapore and so cannot be identified

with the available literature.

The present guide aims to provide all the necessary information with

which anyone interested in grasses should be able to identify the species that

occur in Singapore. All species that have been mentioned for the island state

are included in this guide. The text is fully illustrated, a main key to the flowering

grasses of Singapore is continued to the species level in the generic keys, and

Supplement of Gardens’ Bulletin Singapore 57 (2005)

a key that uses only the vegetative characters is provided. The identification

keys are designed to be user-friendly: the characters are easy-to-see (with the

aid of a handlens with 10x magnification) and are explained in a section on

Grass Morphology (p. 3) whereas descriptive terms are explained in the Glossary

(p. 19). The Key to Sterile Grass Plants is a novelty for the region. In Europe,

it has been proven possible to identify grasses on basis of vegetative parts

alone (e.g. Van der Meijden 1983). It seemed useful to develop one for Singapore

as a number of species rarely flower and roadsides and fields are mown

frequently. Species that are thought to be extinct or found only in cultivation

are, with a few exceptions, not included in this key. However, the majority of

species can be identified with this key and only in a few cases groups of species

remain (for the time being) unidentifiable when sterile. A shortened key to the

turf grasses is also included (p. 16) to facilitate the identification of turf material.

Bamboos were excluded from the present treatment, as the key in Keng et al.

(1998) seems quite satisfactory. The characters that differentiate these from

true grasses are shortly discussed. A separate section explains how to distinguish

a grass from other groups of plants that look like grasses.

The species are arranged according to their scientific name, rather than

by a common name as many do not have a common name. If an English and

Malay name is known from the literature mentioned above or from Henderson

(1954), they are included. In case more names are available priority has been

given to one that was mentioned by Gilliland (1971) or Keng et al. (1998).

English names have been checked in Mabberley (1997) to ensure they are not

in use for another species. Only one species (Pennisetum purpureum) is known

by two English names. The Malay names have been checked by staff of the

Singapore Herbarium. Only Padi Burung and Rumput Kumpai each refer to

two different species.

For reasons of convenience, the species descriptions are arranged

alphabetically, rather than taxonomically. A modern classification of the grass

family is found in Kellogg (2002). The descriptions are based on material present

in the Singapore Herbarium (SING) and the herbarium of the University of

Singapore (SINU). Almost all species descriptions are accompanied by an

illustration of a (group of) spikelet(s). A selection of species is illustrated in

colour.

The total number of accepted taxa (species, subspecies, varieties) is 134

(including 13 that have been found in cultivation only). Records are not included

in the account if no material is extant in SING or SINU or if they have only

been found cultivated in the Singapore Botanic Gardens. They are briefly

discussed under the genus. Also excluded are Eremochloa ciliaris (Buitenhuis

and Veldkamp 2001: 407; no material), Miscanthus floridulus and M. sinensis

(only cultivated in the Singapore Botanic Gardens).

It is remarkable that in the tropics grasses are largely ruderal with many

species being cosmopolitan. In relation to other plant families the grasses have

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 3

a high (if not the highest) number of exotics: 31% of Singapore grasses are not

native. Some of them have been introduced intentionally, in Singapore for

instance as fodder for the horses of the British army or as a part of new seed

mixes for turfs. Others where brought in unintentionally, e.g. with straw and

hay for cattle, with sand for land reclamation or with potting mix. It is unlikely,

however, that we have a complete record of all the species that have been

introduced in Singapore. Some of the exotics will have lived here for too short

a period to become noticed. If the climate is not (very) suitable for the introduced

‘species it cannot establish a viable population and when the import of the

fresh material ceases the species will (slowly) disappear. Other introductions,

however, are very successful. Today, 28% of the most common species are not

native whereas the commonest grass species of Singapore is probably the-exotic

Axonopus compressus. This shows that not only the number of exotic grass

species is high, but that they do actually form an important part of the present

day vegetation.

The introduction of new species is an ongoing process that is only to a

certain level predictable, depending for instance on the geographical origin of

the imported material. No attempt has been made to include species in the

account that might be expected in the near future. If grasses collected in

Singapore cannot be identified with the present Field Guide, there are several

options. It could be that the plant under study is not actually a grass, or if it is

indeed a grass its characters could have been misinterpreted or the species 1s

more variable than described here or it might be a new record for Singapore.

In the case of new introductions the following works are recommended:

Gilliland (1971), Bor (1960) and the World Grass Species Database at the

Kew website (http://www.rbgkew.org.uk/data/grasses/grasses.ink).

Grass morphology

Vegetative parts

Grasses do not have primary roots (as in carrots, Daucus carota), only a tuft of

adventitious roots. In general grass roots are rather uninteresting except for

the ones of Lophatherum, which form small edible tubers.

The stem of a grass 1s called a culm. It is jointed: divided into nodes and

internodes (Fig. 1la,b). Nodes can be recognized as short, (slightly) thickened

and solid parts of the culm between the much longer internodes. The nodes

bear the leaves and, if present, roots and branches (in the leaf-axil). Before

roots at the nodes start to grow they may be seen as tiny circular pits termed

root-eyes (Plate 28, 29). Root-eyes are obvious in Saccharum species. The

internodes are generally hollow, but are solid in a few grasses (e.g. Saccharum

Sugar Cane and Themeda Tasselgrass). At the base of the culm they can be

very short (about as long as the nodes) resulting in a concentration of leaves,

4 Supplement of Gardens’ Bulletin Singapore 57 (2005)

which cover all the nodes. In the upper part of the culm the internodes are

elongated.

The typical culm grows vertically: erect, or ascending from a geniculate

base. Some culms, however, grow horizontally. Horizontal culms that lie on

the soil surface are called stolons. They are generally green and the nodes root

or not and bear well-developed leaves. Culms growing horizontally beneath

the soil surface are called rhizomes. Rhizomes are generally whitish with rooting

nodes and reduced, scale-like leaves. The presence of nodes and reduced leaves

shows that rhizomes are not roots.

Branching of the culm always occurs at the nodes in a leaf axil between

the culm and the basal part of the leaf (sheath, see below). The new shoot

either grows upward to emerge at the sheath’s tip (intravaginal branching,

Fig. 2a) or else breaks through the base of the sheath and grows outward

(extravaginal branching, Fig. 2b). Rhizomes and stolons are the result of

extravaginal branching. Tufted plants, on the other hand, are the result of

intravaginal branching at the base of the culm.

Grass leaves always arise from the node, in two alternate (‘distichous’ )

ranks along the culm (Fig. 3), but this is often obscured due to the twisting of

the sheath or the culm inside the sheath. The leaves are composed of three

parts: sheath, ligule, blade (Fig. 1c,d, 4).

The sheath is a tube around the culm with (slightly) overlapping margins

(only in a few species non-Singaporean are the margins united). The sheath

may be shorter to longer than the internode. The midrib may be raised or winged,

and (minute) transverse veins may be present as well (best seen with back

lighting). The throat (top of the sheath) or the base of the blade may have ear-

shaped or triangular appendages, referred to as auricles (Fig. 4d). Auricles can

be persistent or deciduous, glabrous or hairy.

The leaf blade arises on the top of the sheath. It is generally flat and

elongated (much longer than wide) with more or less prominent longitudinal

veins (parallel-nerved; only rarely, as in Scrotochloa urceolata, the blades are

pinninerved: with veins arising from the midvein). Transverse veins are

sometimes present as well. However, the presence of transverse veins in the

blade does not predict their presence in the sheath, and vice versa. The midvein

is prominent or not, is sometimes broad and white at the upperside of the blade

Figure 1. The jointed culm (stalk) of a grass.

a. nodes, b. internodes, from the nodes arise c. the sheaths with d. their blades.

Figure 2. Branching of the culm.

a. intravaginal; b. extravaginal.

Figure 3. Grass leaves arranged in two ranks or distichous.

Figure 4. Grass leaf.

a. sheath, b. blade, c. ligule, d. auricle.

Figure 5. Blade base.

a. cuneate, b. rounded, c. cordate, d. amplexicaul.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 3

6 Supplement of Gardens’ Bulletin Singapore 57 (2005)

or distinctly winged underneath. The base can be cuneate, rounded, cordate or

amplexicaul (Fig. 5). If the blade at the base is narrower than the sheath, the

sheath is termed shouldered (Fig. 6a). Sometimes the blade is much narrowed

at the base and appears stalked. It is then termed pseudopetiolate (Fig. 6b).

(This is not a true petiole, which would be at the base of the sheath). The way

the young blades emerge is constant within a species: either inrolled with one

margin overlapping the other (Fig. 7a), or folded along the midrib with the

margins meeting (Fig. 7b), or plicate (pleated; Fig. 7c).

The ligule is located at the junction of sheath and blade (Fig. 4c) on the

side facing the culm (the adaxial side) and is either membranous and glabrous

or hairy or is just a row of hairs. Only in a few species is the ligule absent (e.g.

Echinochloa colona). A ligule even a few millimeters long is readily recognized.

Shorter ligules, however, are more difficult to see. To examine it, pull the culm

away from the blade and put your index finger on the underside of the blade’s

base while you prise open the sheath with your thumb. Pull back the blade

with the middle finger of the same hand. With your free hand you can now

hold the handlens to see the ligule (practice this first with one of the taller

grasses).

Sometimes (as in bamboos) a collar is present. This is a band of hardened

tissue on the abaxial side (this is the side furthest away from the culm) of the

top of the sheath, indicating that the blade will eventually break off at that

point (generally leaves have no callus, sheath and blade just wilt). An herbaceous

rim or a row of hairs at this place is called an external ligule (e.g. bamboos,

Paspalum conjugatum; Fig. 8).

The initial leaf (or leaves) of a culm or branch is (are) often different

from the upper leaves in having no blades and ligules. The first leaves of young

culms may be scale-like sheaths or cataphylls (Fig. 9). As growth proceeds,

normal leaves are produced. When culms branch, the first leaf of the new

shoot is produced from the adaxial side, between the new shoot and the main

culm. This prophyll is an elongated and thin sheath-like scale with two keeled

veins; its function is uncertain.

Various parts of the plant may be covered with hairs and these parts are

then called hairy. If margins or ligules have a fringe of hairs they are described

as ciliolate (with short hairs) or ciliate (with long hairs), but the difference is

gradual. ,

Figure 6. Leaf blade narrowed at base.

a. Sheath shouldered, b. blade pseudopetiolate.

Figure 7. Young leaf blades.

a. inrolled with one margin overlapping the other, b. folded along the midrib with the

margins meeting, c. plicate (pleated).

Figure 8. External ligule, a herbaceous rim or a row of hairs at the abaxial side of the

sheath’s apex.

Figure 9. Cataphylls (sheathlike and bladeless bracts) at the base of the culm.

Field Guide to the Grasses of Singapore (Excluding the Bamboos)

Fig. 6

Fig. 9

8 Supplement of Gardens’ Bulletin Singapore 57 (2005)

The inflorescence will generally develop from the terminal leaf (but

see below), which is then also known as the flag leaf. It may differ from the

leaves on sterile shoots in, for example, hairiness, size or ligule length.

Reproductive parts

Flowers

Grass flowers are highly reduced. They consist of only 1—6 stamens, a single

ovary with 1—3 feathery stigmas, and/or 2—3 minute fleshy scales or lodicules.

True sepals and petals are lacking (Fig. 10a—d). The lodicules are thought to

be a reduced perianth and their function has still to be ascertained. The grass

flower is protected by two straw-coloured to green bracts. The lower bract is

called the Jemma (Fig. 10e) and the upper is called the palea (Fig. 10f). The

palea has two keeled veins. The lemma and palea together with its flower is

called a floret (Fig. 10a—f). Florets are either bisexual (with stamens and ovary)

or unisexual (male with stamens only or female with ovary only) or sterile

(empty, both stamens and ovary are absent).

Bisexual florets may be cleistogamous, that is, they remain closed even

at flowering time, which results in self pollination. They are recognized by the

presence of anthers on the top of mature fruits in closed florets (e.g. Eragrostis,

Eriachne).

Spikelets

Florets are arranged in two alternate ranks on a central axis, the rachilla (Fig.

10g). At the base of the rachilla, below the lowest floret, are two empty bracts:

the lower glume (Fig. 10h) and the upper glume (Fig. 101). The structure of

two glumes and a rachilla with one to many florets is called a spikelet (Fig.

10a—1i). The structure of a spikelet can be compared with a flowering branch

(Fig. 11), with each flower in the axil of a leaf and accompanied by an extra

leaf. However, it is still unclear if the lemma and the palea are homologous to

(i.e. evolved from) these two leaves (Kellogg, 2002).

The spikelet is thus a miniature inflorescence. It is the basic unit in the

description of the grass inflorescence. Tiny as it may be, the basic pattern of a

spikelet is clear-cut. However, structures may be duplicated (as are the glumes

in bamboos) or reduced or even absent. This hampers the interpretation of the

structures and the determination of the number of florets in a spikelet (especially

the distinction between 1- vs 2-flowered). Therefore, in the main key to the

Figure 10. Grass flowers arranged in a spikelet (exploded view).

a. lodicule, b. anther, c. ovary, d. stigma, e. lemma, f. palea, g. rachilla, h. lower glume,

i. upper glume.

Figure 11. Branch with flowers in the axils of bracts and pedicels each with a bracteole.

Figure 12. Cross-section of a spikelet.

a. not flattened, b. laterally flattened, c. dorsiventrally flattened.

* = rachilla.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 9

ca” i Fig. Il

10 Supplement of Gardens’ Bulletin Singapore 57 (2005)

flowering grasses (p. 20) whenever possible the neutral term bracts is used

instead of glumes, lemmas and paleas but in the generic keys these terms are

used as the genus description has already clarified which structures are to be

expected.

Spikelets may be sessile (not stalked) or stalked; the stalk is called the

pedicel. Spikelets or groups of sessile spikelets may be surrounded by an

involucre, which is formed of bristles or a single bract attached at the base of

the spikelet(s) and at least half as long as the spikelet. The involucre together

with the spikelet(s) it encloses is known as a bur. Generally, spikelets on the

same plant are similar (or homomorphous), but sometimes they are

heteromorphous, very different from each other (e.g. Digitaria bicornis,

Scrotochloa urceolata, Zea mays). These different types of spikelets may be

found within the same inflorescence or in separate inflorescences on one plant.

Apart from the size (length, width) of the spikelet, the shape of the spikelet

is very important for the identification of the species. The outline of the spikelet

varies from rounded (orbicular) to linear. Looking at the cross-section of the

spikelet, it can either be flattened or not flattened. A spikelet is called not

flattened if the cross-section is rounded: all axes are equally long (Fig. 12a)

and the glumes and lemmas are generally distinctly rounded on the back. The

spikelet is called flattened if its cross-section is elliptic with one axis distinctly

longer than the other. Laterally flattened spikelets have boat-shaped glumes

and lemmas and the rachilla is (nearly) visible (Fig. 12b). When all the glumes

and lemmas are flattened to only slightly rounded on the back, the spikelet is

called dorsiventrally flattened and the rachilla is hidden deeply inside the

spikelet (Fig. 12c). Finally, the hunch-backed or distinctly lop-sided spikelet

is called gibbous (e.g. Cyrtococcum species).

The number of florets per spikelet varies from one to a few tens depending

on the species. For most species this number is fixed, e.g. spikelets strictly 1-

flowered or (most commonly in Singapore) strictly 2-flowered (note that florets

are referred to as flowers). However, spikelets with an additional floret may

be found as abnormalities within the same inflorescence (e.g. Jsachne species,

Panicum maximum). It is therefore advisable to always check several spikelets

within the same inflorescence. The number of florets is much less fixed in the

species that have 3- or more-flowered spikelets (for example, Eragrostis). The

lowest floret is referred to as the 1* floret (and 1“ lemma, 1|* palea), etc.

Awns are needle-like appendages or the much-narrowed continuations

of the glumes, the lemmas or the paleas. They arise either from the tip (as in

Dactyloctenium aegyptium) or in a sinus (referred to as the notch) between

two lobes at the tip of the bract (as in Chloris barbata), or from its back. They

vary in length and shape. In Singapore awns are always simple (unbranched),

either straight or geniculate (then often with a twisted basal part and terminal

bristle) and smooth or scaberulous. Bracts with very short awns (less than

0.5—1 mm long) are often called mucronate (if they are the continuation of

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 11

the midvein only) or apiculate (if they are the continuation of midvein and a

narrow part of the bract), but the difference between these and a true awn is

gradual. The number of awns per spikelet varies from none to six.

Spikelets or parts of spikelets may be articulated, which means that they

fall off or break up at maturity. A thickened structure (the callus) often indicates

the place of articulation where either the entire inflorescence or a branch or

entire spikelet or a part of the spikelet (with the rachilla articulating or not)

breaks off.

Inflorescence

The inflorescence is the arrangement of the spikelets without any leafy bracts.

It develops within the protection of a sheath, emerging only when it is almost

ready for flowering. The main axis of the inflorescence is called the rachis (if

unbranched) or central rachis (if branched). The part of the main axis below

the first branch or spikelet is called the peduncle. The peduncle is generally

long, with the inflorescence arising well above the flag leaf.

The spikelets are either arranged directly on the rachis, or on (primary,

secondary or tertiary) branches. They are single or arranged in groups of 2—5

spikelets often with | sessile and lor more pedicelled. An unbranched

inflorescence with only sessile spikelets is called a spike (Fig. 13a), or if all

spikelets are pedicelled, it is called a raceme (Fig. 13b), and if the spikelets

come in pairs of | sessile and | pedicelled spikelet it 1s called a spike-like

raceme (Fig. 13c). The branched inflorescence is called a panicle and has

branches either at intervals along the central rachis (Fig. 13d) or clustered at

the tip of the peduncle (a central rachis is absent in this case, Fig. 13e). The

(central) rachis and panicle branches can be rounded, triquetrous (triangular in

cross-section) or flattened and the branches can be simple (unbranched) or

branched further.

Generally, a single inflorescence arises from the terminal leaf of a culm

or lateral branch. Sometimes, however, the terminal leaf has two or more

inflorescences (Fig. 14) or the subterminal leaves have inflorescence(s) as well.

The ultimate in this range is a spatheate inflorescence where much branched

culms with spathes (sheath-like bracts or reduced leaves) all produce a separate

inflorescence (Plate 36). The final sheath is any sheath from which axil arises

an inflorescence without any leaves or spathes. It may be the sheath of the flag

leaf or a subterminal leaf or a spathe.

Fruit

The ovary of a grass contains a single ovule that will mature into a single seed

and the ovary wall into the fruit wall. The fruit is a grain or caryopsis with the

fruit wall permanently fused to the seed coat (except in Eleusine and

Sporobolus). Only grasses have this type of fruit. It 1s orbicular to linear-

lanceolate in outline and flattened to round in cross-section. The hilum, the point

sed Supplement of Gardens’ Bulletin Singapore 57 (2005)

of attachment of the grain, is facing the palea. The embryo is at the base and

the starchy mass making up the rest of the fruit is the endosperm or foodstore

for the germinating plant. This is the part we eat.

Diaspore

A grass diaspore, the part that is dispersed, may be the fruit alone (e.g.

Eragrostis) or fruits together with a floret (e.g. Centotheca), a spikelet (e.g.

Lophatherum), groups of spikelets (Cenchrus, Pennisetum) or even an entire

inflorescence (Scrotochloa) depending on where disarticulation takes place.

The presence of hooks, hairs, awns and spines increases the chance of

(long distance) animal or wind dispersal. The ability to float (as in Mnesithea,

Rottboellia, Thuarea), will enhance dispersal by water. The diaspores of, for

instance, Rottboellia cochinchinensis also seem to get carried away by ants

that are attracted by elaisomes, structures that contain oily substances.

Life Forms

The absence of pronounced seasons in Singapore often makes it difficult to

establish whether a plant is an annual or a perennial. However, if at flowering

time all shoots of a plant show an inflorescence it is considered an annual.

Some annuals have more than one generation a year. The presence of sterile

shoots at flowering time, a rootstock and/or cataphylls indicates the plant is

perennial.

Making a Reference Collection of Grasses

When studying grasses it is a good idea to build up a small reference collection

or herbarium for personal use. This enables one to compare species seen at

different times and places and makes it easier to remember their names.

Collected plants should be dried and mounted on thick paper.

It is quite simple to make good collections of the smaller-sized grasses:

just collect the whole plant, including the underground organs. A blunt knife,

a small triangular trowel or a screwdriver is a useful tool to dig up the roots

and/or rhizomes, especially when clayey soil has hardened after a long period

of drought. Remove as much soil as is possible when still in the field. It is

advised to gather several specimens (‘duplicates’) and to collect both flowering

Figure 13. Type of inflorescence.

a. spike with spikelets single and sessile, b. raceme with spikelets single and

pedicelled, c. spike-like raceme with spikelets in pairs of 1 single and 1 pedicelled, d.

panicle with central rachis present, e. panicle without central rachis.

Figure 14. Terminal leaf with two panicles.

Figure 15. Two types of bamboo leaves.

a. culm leaves, b. foliage leaves.

Figure 16. Culm leaf of bamboo.

a. sheath, b. ligule, c. blade and d. auricles.

Figure 17. Tip of the leaf blade or bract.

a. bifid, b. emarginate, c. truncate, d. obtuse, e. acute, f. acuminate, g. mucronate.

Field Guide to the Grasses of Singapore (Excluding the Bamboos)

ened

—

nage

14 Supplement of Gardens’ Bulletin Singapore 57 (2005)

and fruiting spikelets, so that both the different parts of the spikelet (including

the anthers) as well as its disarticulation can be studied (essential in Eragrostis).

A young, emerging blade should also be included, if present, especially when

sterile material is collected. All this will increase the chance of finding the

correct name.

If the plant is too big to collect whole (or if it is impossible to reach the

lower parts), at least the inflorescence with the flag leaf and its node should be

collected. Try also to get a piece of the culm with at least two leaves, for the

flag leaf may differ considerably from the ones lower down. Be careful when

handling the leaves, as some have very sharp edges that can make very nasty

wounds.

For future reference number the collections in a notebook with a

consecutive numbering, starting with |. Attach small pieces of paper with that

number to the plants. Date and place of collection should be recorded as well

as fieldnotes, such as habitat and any features that will be lost in the drying

process (e.g. coloration or three-dimensional shapes). At the same time, all

other parts that could not be included in the collection should be described,

such as whether the culm is single or tufted, erect or ascending, whether or not

stolons or rhizomes are present and the total length of the culm. Features that

can still be seen on the dried specimefi (e.g. presence of hairs, veins etc.) need

not be recorded. If, at a later moment, you think you have collected the same

species again, do give it anew number, not the one you have employed before.

Most likely it is from another locality and therefore a different gathering.

Besides, it might even turn out to be a different species.

Collected specimens are easily dried when pressed between a single

sheet of tabloid-size newspaper. Arrange the parts so that all details can be

studied (avoid overlapping the leaves) and fold or cut the parts to fit the size of

paper used for mounting (see below). Best results are obtained when dried for

a few hours to a day in an oven at 60 °C (maximum). Without an oven, the

drying process will take a few days and the newspaper needs to be changed

every day to prevent mould from growing and to preserve some of the original

colour.

After drying, the specimens are best mounted with narrow strips of glued

paper on sheets of thick paper (A3 size) adding a label with the name and the

field notes, as well as collector’s name and place and date it was collected

(Plate 1). Store them in boxes or plastic bags in a dark and dry place; either

add mothballs or freeze them at regular intervals to prevent insects from foraging

on the collection.

You might want to send (unmounted) duplicates with the labels to a

scientific institute, like a local herbarium. They might want to keep them as

scientific records and so your efforts could become part of the national and

international heritage. The collector’s name and numbers allow for quick and

unequivocal reference to the individual collections.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) fs)

Turf Grasses

A number of characteristics make grasses especially suitable for turf and lawns.

First of all, in a number of perennial grasses the growing point of the non-

flowering culm is close to the soil surface. When mown, these grasses can

thus resprout more quickly than other plants. Furthemore, their linear leaf

blades with long fibres are better able to resist being trodden on. Finally, the

ability to form horizontal culms (rhizomes and stolons) enables grasses to

-form mats. The more frequently mown the grass is, the denser the mat becomes.

When mown too low down, however, they cannot resprout and as a consequence

the turf will be damaged.

From the above it will be clear that some grass species are more suitable

for turf than others. Annual grasses, for instance, and grasses that form a single

erect stem or that grow in tufts are of no use, as they will never form a dense

mat. Only a few species, Axonopus compressus, Cynodon dactylon (including

the cultivars Tifway, Tif dwarf and Tif-fine that are commonly used in the

various parts of golf courses), Digitaria didactyla and Zoysia matrella, are

commonly used for lawns and fields in Singapore and a few more are (or

were) used infrequently. The fine-leaved grasses, such as Cynodon dactylon,

Digitaria didactyla and Zoysia matrella, produce beautiful lawns but need a

lot of maintainance. As it is difficult even for wiry weeds like Desmodium

triflorum (Lesser Clover-leaved Desmodium) to grow between the densely

arranged grass stems, they tend to cover the lawn and need to be weeded out

by hand. Also, these fine lawns are not tolerant to shade and tend to be less

resistant to intense use. Coarse turf with wide-leaved grass species, such as

Axonopus compressus and Stenotaphrum secundatum does not need regular

weeding as wiry weeds can grow underneath these grasses. Both grass species

are strong and will give good general purpose lawns. They will also grow in

light shade. It is impossible, however, to grow a lawn in deep shade. Eremochloa

ophiuroides (Munro) Hack. (Centipede Grass, Rumput Lipan, Rumput

Centipede) is also used for lawns in Peninsular Malaysia, but has not (yet)

been found in Singapore. This slow-growing species gives a rather coarse,

good lawn that is not very tolerant to wear.

The separate key to the species given here enables the user to quickly

identify turf grasses. The various cultivars of these species are, however, not

separated. As the lawn will normally be mown before the plants start flowering,

only vegetative characters are used. For older turfs in which weedy grass species

have invaded, The Key to Sterile Grass Plants of Singapore (p. 142) should be

used. Descriptions of the species (except for Eremochloa ophiuroides) can be

found in Descriptions and Generic Keys (from p. 31).

16 Supplement of Gardens’ Bulletin Singapore 57 (2005)

Key to the turf grasses

la. Leaf blade 0.6—2.5(—3.0) mm wide. 2

1b. Leaf blade 3—9 mm wide. 5

2a. Ligule as a row of hairs, longest hairs at least 0.5 mm long. 3

2b. Ligule membranous, glabrous or with hairs at the tip up to 0.1 mm long.

3a. Sheath rounded, midrib not winged. Blade at base cuneate.

Bermuda Grass, Cynodon dactylon

3b. Sheath flattened, midrib winged, wing 0.5 mm wide, Blade at base rounded,

Centipede Grass, Eremochloa ophiuroides

4a. Ligule (0.5—)1.5—2.2(—2.5) mm long, glabrous.

Serangoon Grass, Digitaria didactyla

4b. Ligule 0.2 mm long, ciliolate. Siglap Grass, Zoysia matrella

5a. Young leaf blade folded along the midrib. 6

5b. Young leaf blade inrolled. 8

6a. Ligule as a row of hairs, 0.5—0.7 mm long.

Centipede Grass, Eremochloa aphiuroides

6b. Ligule membranous, with hairs at the tip up to 0.1 mm long. 7

7a. Leaves glossy and (dark) green; sheath with margins in upper half glabrous

or glabrescent; blade hardly wider than sheath. Axonopus fissifolius

7b. Leaves glaucous; sheath with margins densely hairy in upper half; blade

distinctly wider than sheath.

St. Augustine Grass, Stenotaphrum secundatum

8a. External ligule present as a short-hairy fringe. 9

8b. External ligule absent. 10

9a. Nodes hairy. Ligule densely fringed. :

Carpet Grass, Axonopus compressus

9b. Nodes glabrous. Ligule glabrous,

Buffalo Grass, Paspalum conjugatum

10a. Sheath 1—2.5 cm long, without transverse veins. Ligule 0,05—O.1 mm

long. Love Grass, Chrysopogon aciculatus

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 17

10b. Sheath 4—8.5 cm long, with transverse veins. Ligule 0.5—0.8 mm long.

Creeping Panic Grass, Panicum repens

Bamboos and Grasses

Bamboos, which belong to the Bambuseae tribe in the Gramineae, are actually

grasses and not palms. Unlike palms but like the true grasses, they have jointed

culms with solid nodes and hollow internodes. Kellogg (2002) discusses their

relationship within the grass family.

Bamboos are recognized by a number of characters. First of all, they

have woody culms. Secondly, bamboos have leaves of two types (Fig. 15).

The young culm shoot is the part of the bamboo that we eat. It is covered with

scale-like leaves, which are brownish and rather rigid structures with the blade

absent or much shorter than the sheath and often triangular in shape (Fig. 15a

and 16). These culm-leaves are shed entirely when the culm hardens during

growth. The foliage leaves, on the other hand, are found on the branches higher

along the main culm (Fig. 15b). They have persistent sheaths with the blades

deciduous above the collar. The blades are green, elongate and pseudopetiolate.

Finally, the spikelets of bamboos are arranged in dense nodal clusters on a

leafless axis (Plate 2). The spikelet often has more than two glumes. However,

flowering of bamboos is rare, and therefore in their taxonomy much emphasis

is laid on the vegetative parts. If you see any flowering specimens it might be

wise to make a collection, for some flower only once in a human life time. Be

sure to collect the culm leaves, noting in your data the presence and direction

of the remnant of the blade (pointing up, down or horizontal). The hairs on the

sheath, if present, may stick to your hands. To remove the hairs, gently rub

your hands through your own hair.

The bamboos are excluded here because Keng et al. (1998) provided a

useful key to all 19 species of bamboo known from Singapore, including those

only known from cultivation. Further information can also be found in Wong

(1995a,b) and Ohrnberger (1999).

Grasses and Grass Look-alikes

The grass family, Gramineae, can be confused with a number of other groups.

For Singapore, the sedge family, Cyperaceae, is the most relevant one. Both

have strap-shaped leaves that are differentiated into sheath and blade and the

flowers are arranged in spikelets without true sepals and petals. There are a

number of differences between the two families, but most do not hold for all

members, certainly not worldwide (Table 1).

Supplement of Gardens’ Bulletin Singapore 57 (2005)

Table 1. Differences between grasses and sedges in Singapore.

Grasses Sedges

Culm Rounded to flattened, never Triangular in cross-section or

triangular in cross-section rounded to flattened

Leaves In two ranks (may be In three ranks, rarely in two

obscure)

Sheath Overlapping, not fused (in Fused

margins Singapore)

Ligule Generally present and short Absent or present (in Singapore

to long, sometimes absent always less than | mm long)

Flower Each generally subtended by Each always subtended by only

two bracts one bract (and a number of

perianth-bristles or scales can

be present as well)

Florets Arranged in two ranks Spirally arranged or in two

ranks

Fruit Fruit wall generally fused to Fruit wall free from the seed

seed coat

coat

Species from other families that could also be mistaken for grasses

because of their grass-like leaves include Blyxa alternifolia and B. auberti

(Hydrocharitaceae), Eriocaulon longifolium and E. truncatum (Eriocaulaceae),

Murdannia nudiflora and M. vaginata (and perhaps other Commelinaceae),

Philydrum lanuginosum (Philydraceae), Xyris complanata and X. pauciflora

(Xyridaceae). Most of them differ from grasses in that they have three sepals

and three colourful petals, but their petals tend to fall after flowering, and

some do have a lesser number or scale-like perianth parts. However, all these

non-sedge grass look-alikes have fruits that are different from the grasses:

namely a dehiscent capsule containing two to many seeds that are free from

the fruit wall as compared with always one seed per flower in grasses and

sedges.

If plants are not flowering or fruiting it, might be difficult to establish

beyond doubt whether they are grasses. If the sheath has free margins and a

ligule is present then the plant is a grass. On the other hand, if the sheath 1s

closed (the margins are fused) or a ligule is absent, and/or the distinction between

sheath and blade is unclear then it is probably not a grass.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 19

Glossary

Technical terms have been avoided as much as was possible. The use of some

general descriptive terms, however, could not be avoided. They follow in

alphabetical order. For the names of the different parts of a grass plant see the

section on Grass Morphology (p. 3).

acuminate Tapering to a point (Fig. 17f).

acute Sharply pointed (Fig. 17e).

amplexicaul Clasping the stem and the two lobes not fused (Fig. 5d).

annular Like a ring.

antrorse Pointing forwards or upwards.

ascending Gradually curving upward. y

bifid Divided in two parts (Fig. 17a).

bristles Long stiff hairs (longer than | mm).

caducous Falling off (eventualy).

cartilaginous Hard and tough.

ciliate Fringed with long hairs.

ciliolate Fringed with short hairs.

collar-shaped The outline a band much wider than long (Fig. 18a).

cordate Heart-shaped (Fig. 5c).

cuneate Wedge-shaped (Fig. 5a).

decumbent Horizontal and only the tip curving upward.

ellipsoid Three-dimensional body with elliptic outline.

elliptic The outline of a flattened circle, length-width ratio c. 1.5 and largest

width in the middle.

emarginate With V-shaped indentation at the top (Fig. 17b).

erect Upright.

fusiform Cigar-shaped and length-width ratio more than 2.

geniculate Abruptly bent upward, knee-like.

glabrescent Becoming glabrous.

glabrous Hairless.

glaucous With a waxy layer and a bluish glow.

globose Globe-shaped.

granulate Covered with small grains.

herbaceous Soft, coloured and not transparent.

heteromorphous Of dissimilar forms.

homomorphous Of similar forms.

hyaline Thin, colourless and transparent.

indument The covering of hairs.

indurated Hardened.

keeled With a ridge.

lanceolate Length-width ratio 3—5S.

20 Supplement of Gardens’ Bulletin Singapore 57 (2005)

linear Length-width ratio 10 or more and parallel sides.

membranous Thin and semi-transparent.

mucronate Ending abruptly in a sharp point that is the continuation of the

midvein only (Fig. 17g).

nodular With small rounded lumps.

nodule Small rounded lump.

oblong Length-width ratio c. 2.5 and largest width in the middle.

obtuse Blunt (Fig. 17d).

orbicular The outline circular, length-width ratio 1.

ovate The outline egg-shaped, length-width ratio c. 1.5 and largest width below

the middle (Fig. 18b).

ovoid Egg-shaped.

patent Spreading.

pectinate Margin with a row of relatively long hairs, comb-like.

punctate Covered with small dots.

reticulate Covered with intersecting lines forming the shape of a network.

rugose With irregular ridges as if wrinkled.

rugosity Irregular ridge like a wrinkle.

rugulose With irregular minute ridges as if finely wrinkled.

scaberulous With a slightly rough surface.

scabrous With a rough surface.

scrambling Climbing by sprawling over other plants or fences.

serrate Margin saw-toothed.

serrulate Margin minutely saw-toothed.

setaceous Bristle-like.

setae Short stiff hairs (up to | mm long).

setulose With short stiff hairs.

smooth Surface even, free from roughness, lumps, wrinkles etc.

striate Covered with narrow lines, ridges or furrows.

subglobose Almost globe-shaped.

terete Rounded in cross section.

triquetrous Triangular in cross-section.

truncate Appearing as if cut off (Fig. 17c).

undulate Wavy, going up and down.

winged With a flat elongation.

Key to the Flowering Grasses of Singapore

Transverse veinlets are best seen with back lighting (be sure not to look into

the sun or a lamp when using a hand lens). The terms leaves, sheaths and

blades do not include cataphylls, prophylls and spathes. Pedicel length is to be

measured on any spikelet except the uppermost of a branch where it may be

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 21

much longer than usual. Unless stated otherwise, the size of spikelets excludes

awns if present. The adaxial side of a structure is the side closest to the axis,

the abaxial side is the side furthest away from the axis; for leaves this axis is

the culm, for inflorescence branches it is the central axis and for spikelets it is

the rachis or the branch of the inflorescence they are directly attached to.

To facilitate back tracking of the followed path in the key, the couplet

number of origin is repeated in bold print at the beginning of the couplet it

referred to if it is more than 3 couplets away.

A. Plant woody. Internodes hollow. Culm-leaf sheaths deciduous, blade absent

to shortly triangular, shorter than the sheath. Foliage leaves with sheath

persistent; blade deciduous from sheath, linear to lanceolate,

pseudopetiolate (Fig. 6). Inflorescence with spikelets arranged in dense

nodal clusters without subtending bracts (Plate 2).

Bamboos (see Keng et al. 1998)

B. Plant herbaceous to semi-woody. Internodes hollow or solid. Culm-leaves

as foliage leaves, basal ones with sheath deciduous or not; blade

persistent, generally longer than sheath, ovate to linear, pseudopetiolate

or not. Inflorescence with spikelets single or in small groups and not

densely clustered, or spatheate and with clusters of spikelets in subtending

bracts. 1

(if not flowering see p. 142)

la. Inflorescence(s) from final sheath either an unbranched spike or (spike-

like) raceme (Fig. 13a—c), or branched with the longest branch up to 5

mm long and spikelet unawned. Ligule always present. 2

1b. Inflorescence(s) from final sheath branched (Fig. 13d,e), a panicle with

branches at least 10 mm long (sometimes with only 2 branches closely

together, as in Plate 13, 15); if panicle branches shorter then either ligule

absent or spikelet with 1—6 awns of at least 2 mm long. 24

2a. Spikelets (in groups) all surrounded by involucre of bristles or indurated

bract at least half as long as spikelet, and falling off together with them.

2b. Spikelets not or not all surrounded by an involucre, bristles when present

either less then half as long as spikelet or persistent on pedicel after

spikelet has fallen off (see Setaria parviflora). 4

3a. Involucre a dentate and indurated bract with bristles in lower half.

Cenchrus

3b. Involucre of many bristles only. Pennisetum

29 Supplement of Gardens’ Bulletin Singapore 57 (2005)

4a. Inflorescence at base with a hard bead-like involucre enclosing | female

and 2 sterile spikelets; rachis continuing through pore at tip of involucre,

with 2 pairs of male spikelets. Culm internodes solid.

Coix lacryma-jobi

4b. Inflorescence at base with spikelets exposed, either involucre absent or

consisting of sterile herbaceous spikelets. Culm internodes solid or

hollow. 5

Sa. Spikelets all either unawned, or awned with awn up to | mm long and

spikelet including awn shorter than 4 mm. 6

5b. Spikelets, at least part of them, awned, either awn at least 2 mm long or

spikelet including awn at least 9 mm long. 16

6a. Leaf blade 56—75 mm wide. All spikelets unisexual or sterile but never

bisexual, heteromorphous, 8—8.5 mm long. Tripsacum dactyloides

6b. Leaf blade 1—24 mm wide. At least lowest spikelet bisexual. Spikelets

homomorphous or heteromorphous, 1—6 mm long. 7

7a. Lowest bract of spikelet 0.9—1 times as long as spikelet. 8

7b. Lowest bract of spikelet 0.05—0.65 times as long as spikelet. 12

8a. Inflorescence rachis transversely articulating, base of joints with a central

knob. Spikelets paired. Lowest bract of sessile spikelet dorsiventrally

flattened, indurated. 9

8b. Inflorescence rachis not articulating. Spikelets single. Lowest bract of

spikelet either boat-shaped, or flattened and herbaceous. 11

9a. Young leaf blade folded along midrib. Sessile spikelet orbicular, c. 1 mm

long. Lower glume with squarish depressions. Mnesithea granularis

9b. Young leaf blade inrolled. Sessile spikelet ovate to elliptic, 3.9—5 mm

long. Lower glume either granulate or marginal keels at base with gland-

like warts. 10

10a. Lower glume smooth, marginal keels at base with glandlike warts, glabrous

or hairy. 1* floret of sessile spikelet sterile. Mnesithea glandulosa

10b. Lower glume minutely granular, marginal keels smooth. 1“ floret of sessile

spikelet male. Rottboellia cochinchinensis

11a. Nodes hairy. Rachis longitudinally folded (U-shaped), 3.6 mm wide with

| bisexual spikelet at base and 4 or 5 male spikelets above. Inflorescence

disarticulating as a single unit. Thuarea involuta

11b. Nodes glabrous. Rachis rounded, c. 0.2 mm diameter with many bisexual

spikelets, not articulating. Zoysia matrella

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 23

12a. (7) Young leaf blade folded along midrib. Spikelets 3.8—6 mm long.13

12b. Young leaf blade inrolled. Spikelets 1.7—3.4 mm long. 14

13a. Stolons absent. Branchlets absent. Rachis c. 1 mm wide. Spikelets 3—6-

flowered. Eleusine indica

13b. Stolons present. Branchlets each with 3 spikelets. Central rachis 3—4

mm wide. Spikelets 2-flowered. Stenotaphrum secundatum

‘14a. Pedicels (or at least half of them) with bristles 3—11 mm long. Setaria

14b. Pedicels all glabrous. 15

15a. Inflorescence a spike-like panicle; branches many, up to 5 mm long.

Spikelet not flattened, gibbous, 2-flowered. Sacciolepis

15b. Inflorescence a raceme; branches absent. Spikelets laterally flattened,

not gibbous, 1-flowered. Zoysia matrella

loa. (5) Culm with more than 20 inflorescences, forming a spatheate

inflorescence. Each inflorescence at base with an involucre of 2 pairs of

sterile or male, unawned spikelets, followed by | or 2 fertile awned

spikelets (and more sterile spikelets). Themeda

16b. Culm with 1—10 inflorescences, spatheate or not. Each inflorescence

either with fertile spikelets only, or if spikelets paired the pedicelled

ones may be sterile. 17

17a. Spikelets single, lowest bract with awn at least 2 mm long (a basal 0.3

mm-long bract at adaxial side should be ignored if present: see Lepturus).

17b. Spikelets in pairs or threes, lowest bract either unawned or awn up to 0.5

mm long. 20

18a. Rachis 1.5—2 mm wide, with cavities holding the spikelets, breaking up

at maturity. Lowest (abaxial) bract of spikelet flat, indurated.

Lepturus repens var. repens

18b. Rachis 0.1—0.4 mm wide, without cavities holding the spikelets,

persistent. Lowest bract of spikelet boat-shaped, hyaline to thinly

herbaceous. 19

19a. Nodes hairy. Spikelet 2-flowered, lowest bract (excluding awn) 0.6—0.7

times as long as spikelet. Oplismenus burmanni

19b. Nodes glabrous. Spikelet 1-flowered, lowest bract (excluding awn) as

long as spikelet. Perotis indica

20a. Young leaf blade inrolled. 21

24 Supplement of Gardens’ Bulletin Singapore 57 (2005)

20b. Young leaf blade folded along midrib. 23

21a. Spikelets in threes, the middle pedicelled and the 2 lateral sessile.

Polytrias indica

21b. Spikelets in pairs of | sessile and | pedicelled. 22

22a. Sessile spikelet 3.5—5 mm long, dorsiventrally flattened, with 1 awn.

Dichanthium

22b. Sessile spikelet 1.5—2 mm long, laterally flattened, with 2 awns.

Pogonatherum crinitum

23a. Blade at least at base with up to 7 mm-long hairs. Inflorescence | and at

tip of culm. Pedicel glabrous. Spikelets of lower 8 or 9 pairs all unawned,

followed by 5—8 pairs of | sessile awned spikelet and | pedicelled

unawned spikelet. Heteropogon contortus

23b. Blade glabrous. Inflorescences 2—10 scattered along culm. Pedicel hairy

on one margin. Spikelets all in pairs of 1 sessile awned and 1 pedicelled

unawned. Schizachyrium sanguineum

24a. (1) Culm internodes solid. Spikelets unisexual with male and female

spikelets in separate inflorescences on the same plant. Male inflorescence

a panicle from terminal sheath, female inflorescence (cob) in the axils

of lower leaves, completely enclosed in large bracts, with a stout central

axis and numerous fruits in rows. Zea mays

24b. Culm internodes hollow or solid. Spikelets either bisexual or unisexual

with male and female spikelets in the same inflorescence. 25

25a. Leaf blade with transverse veins present. Stolons absent. Blades 8.5—60

mm wide. 26

25b. Leaf blade without transverse veins. Stolons absent or present. Blades

0.5—50 mm wide. 30

26a. Spikelets heteromorphous, unisexual. 27

26b. Spikelets homomorphous, bisexual. 28

27a. Leaf blade 40—62 mm wide, pinninerved. Spikelets unawned; female

spikelet 6—7.5 mm long, not flattened, globose-obovoid; male spikelet

46 mm long, more or less laterally flattened, elliptic-ovate.

Scrotochloa urceolata

27b. Leaf blade up to 30 mm wide, parallel-nerved. Spikelets awned; female

spikelet c. 50 mm long (including awn), more or less laterally flattened,

lanceolate to linear; male spikelet c. 16 mm long (including awn), similar

in shape. Zizania latifolia

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 25

28a. Spikelets 4—12-flowered, awned with awn at least | mm long.

Lophatherum gracile

28b. Spikelets 2- or 3-flowered, unawned or mucronate with mucro up to 0.3

mm long. 29

29a. Plant 20—80 cm tall. Inflorescence up to 20 cm long. Longest branch

6—10 cm long, straight. Spikelets 4—5.5 mm long.

Centotheca lappacea

29b. Plant (40—)200—300 cm tall. Inflorescence at least (20O—)50 cm long.

Longest branch at least 20 cm long, wavy. Spikelets (1.6—)2—2.1 mm

long. Thysanolaena latifolia

30a. (25) Ligule at least 6 mm long, membranous, glabrous. Inflorescence

with central rachis present. Oryza

30b. Ligule either absent or 0.05—4 mm long, if longer than 4 mm then ligule

hairy or inflorescence without central rachis. 31

31a. Inflorescence from final sheath with simple branches only. 32

31b. Inflorescence from final sheath with at least longest branch further

branched. 60

32a. Spikelet 3—many-flowered. 33

32b. Spikelet 1- or 2-flowered. 36

33a. Spikelet unawned. 34

33b. Spikelet awned. 35

34a. Young leaf blade folded along midrib. Inflorescence with up to 11

branches, at least half the number at the tip. Eleusine

34b. Young leaf blade inrolled. Inflorescence a panicle with more than 20

branches scattered along central rachis, often in whorls of 2—4.

Leptochloa chinensis

35a. Stolons absent. Inflorescence branches with spikelets up to the tip. Spikelet

not flattened; awns 3—5 mm long. Chloris barbata

35b. Stolons present. Inflorescence branches without spikelets in the uppermost

1—2.5 mm (Plate 30). Spikelet laterally flattened; awns 1.5—2 mm

long. Dactyloctenium aegyptium

36a. (32) Ligule at least 0.5 mm long, either a row of hairs or basal part

membranous with hairs at tip as long as or longer than the membrane.

26 Supplement of Gardens’ Bulletin Singapore 57 (2005)

36b. Ligule either absent or up to 0.1 mm long or up to 7.5 mm long and

membranous with hairs at tip absent or shorter than the membrane.

37a. Spikelet either unawned or with a mucro up to 0.3 mm long, or awn

concealed and not exserted beyond the spikelet bracts. 38

37b. Spikelet awned, awn obvious and not concealed. 41

38a. Culms 1—20(—40) cm long. Spikelet 1—2.5 mm long, either 1-flowered

or 2™ floret abortive and hyaline or herbaceous. 39

38b. Culms 30—500 cm long. Spikelet 3—4.3 mm long, 2-flowered, 2™ floret

indurated. du

39a. Leaf blade 1.5—2.5 mm wide, cuneate at base. Inflorescence with branches

coming from one point, central rachis absent. Spikelets in two rows,

2.4—2.5 mm long. Cynodon dactylon

39b. Leaf blade 4—9 mm wide. Inflorescence with branches scattered along

central rachis. Spikelets not in rows, c. 1 mm long.

Sphaerocaryum malaccense

40a. Culm erect to ascending, stolons absent. Spikelet at base with a cup-

shaped callus, 0.2—0.7 mm long, often red-coloured. Eriochloa

40b. Culm geniculate to decumbent or creeping to scrambling, stolons absent

or present. Spikelet at base without a cup-shaped callus. Urochloa

41a. (37) Lowest bract of spikelet awned with awn at least 2 mm long.

Oplismenus

41b. Lowest bract of spikelet unawned or with mucro up to 0.5 mm long. 42

42a. Leaf blade setaceous, c. 0.5 mm wide. Spikelet with 2 or 6 awns, the two

lowest bracts as long as spikelet (excluding awns). Eriachne

42b. Leaf blade flat, 3—12 mm wide. Spikelet with 1 awn, lowest bract

distinctly shorter than spikelet. 43

43a. Panicle without central rachis. Spikelet 3—3.5 mm long. (Very dry places).

Alloteropsis cimicina

43b. Panicle with central rachis. Spikelet c. 4.7 mm long. (Marshy places).

Echinochloa picta

44a. (36) Culm with more than 20 inflorescences, spatheate. Inflorescence

without central rachis, branches 2. Cymbopogon

44b. Culm with 1(—10) inflorescences, not spatheate. Inflorescence with central

rachis absent or present, branches 2—many. 45

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 27

45a. Spikelets in pairs, heteromorphous; sessile spikelet (4—)5—7.5 mm long.

Pedicel 0.7—2 mm wide, as wide as or wider than rachis. Ischaemum

45b. Spikelets in groups of 1|—3(—5), homo- or heteromorphous, sessile and/

or pedicelled, 1.2—4.4 mm long. Pedicel 0.1—0.4 mm wide, narrower

than or as wide as rachis. 46

46a. Ligule absent. Echinochloa

46b. Ligule present, although sometimes a minute rim. 47

A7a. Spikelets not flattened or laterally flattened. 48

47b. Spikelets dorsiventrally flattened. 55

48a. Inflorescence without central rachis, branches 2—4. Z 49

48b. Inflorescence with central rachis, branches 3—many. 50

49a. Young leaf blade inrolled. Spikelet 2.3—2.8 mm long, either unawned or

awn 5.5—8.5 mm long. Dimeria ornithopoda

49b. Young leaf blade folded along midrib. Spikelet c. 1.2 mm long, awn minute

and mucronate. Eustachys tenera

50a. Spikelets with lowest bract 0.9—1 times as long as spikelet. 51

S5O0b. Spikelets with lowest bract 0.15—0.7 times as long as spikelet. 52

Sla. Ligule 0.05—0.1 mm long. Spikelets in threes (or the lower in pairs), 2-

flowered, callus needle-shaped, awn c. 4 mm long; the two lowest bracts

herbaceous. Chrysopogon aciculatus

51b. Ligule 2 mm long. Spikelets single, 1-flowered, callus more or less obtuse,

unawned (acuminate); both bracts indurated. Leersia hexandra

52a. Ligule 1.5 mm long. Spikelets gibbous. Cyrtococcum accrescens

52b. Ligule 0.2—O0.7 mm long. Spikelets not gibbous. 53

53a. Spikelets with 2" lowest bract 0.4—0.5 times as long as spikelet.

Ottochloa nodosa

53b. Spikelets with 2° lowest bract (almost) as long as spikelet. 54

54a. Blade at base cordate. Spikelets c. 4.3 mm long; lowest bract 0.7 times as

long as spikelet. Glumes and lemmas at their tips laterally flattened, as

if pinched. Acroceras munroanum

54b. Blade at base cuneate to rounded. Spikelets 2.1—2.4 mm long; lowest

bract 0.15—0.2 times as long as spikelet. Glumes and lemmas at their

tips not laterally flattened. Panicum trichocladum

28 Supplement of Gardens’ Bulletin Singapore 57 (2005)

55a. (47) Spikelet awned, awn 13—23 mm long. | 56

55b. Spikelet unawned. 57

56a. Inflorescence with 13—21 branches. Rachis of branches and pedicels

more or less flattened and furrowed. Bothriochloa bladhii

56b. Inflorescence with 2—5 branches. Rachis of branches rounded, pedicels

somewhat flattened, both not furrowed. Dichanthium annulatum

57a. Spikelets 1.1—1.6 times as long as wide. Paspalum

57b. Spikelets 2.2—3.3 times as long as wide. 58

58a. Spikelets in groups of 2 or 3(—5). Digitaria

58b. Spikelets single. 59

59a. Ligule 0.4 mm long, densely fringed. Inflorescences usually several from

the upper leaf. Spikelets 2—2.4 mm long. Axonopus

59b. Ligule 1 mm long, glabrous. Inflorescence | from the upper leaf. Spikelets

2.7—4.4 mm long. Paspalum vaginatum

60a. (31) Spikelets 3—many-flowered. 61

60b. Spikelets 1- or 2-flowered, rarely and then only a few 3-flowered. 64

6la. Culm 12—100(—160) cm long. Blades 0.5—7 mm wide. Spikelet

glabrous, or with hairs up to 0.6 mm long; 1* lemma 0.7—2 mm long.

Eragrostis

61b. Culm (80—)150—300 cm long. Blades 7—40 mm wide. Spikelet with

silky hairs 2—5 mm long; 1“ lemma 3.5—12 mm long. 62

62a. Ligule of flag leaf 0.4—0.7 mm long (excluding hairs on blade just behind

ligule). Rachilla internodes between the lemmas with long hairs. Lemmas

glabrous. Phragmites vallatoria

62b. Ligule of flag leaf 1.5—5 mm long. Rachilla internodes between the

lemmas glabrous or with short hairs at its tip only. Lemmas (except

first) with long hairs. 63

63a. Culm internodes hollow. Ligule 1.5—2 mm long, membraneous, tip

ciliolate. Leaf blade 15—40 mm wide. The two lowest bracts of spikelet

c. 0.8 times as long as spikelet (including awns). 1* lemma hairy.

Arundo donax

63b. Culm internodes solid. Ligule 3—5 mm-long row of hairs. Leaf blade

7—20 mm wide. The two lowest bracts of spikelet 0.2—O.3 times as

long as spikelet (including awns). 1“ lemma glabrous.

Neyraudia arundinacea var. zollingeri

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 29

64a. (60) Ligule absent. Echinochloa

64b. Ligule present, sometimes just a minute rim. 65

65a. Either lowest two bracts of spikelet indurated (in lower half) and as long

as spikelet (excluding awn when present), or each spikelet with 2 bracts

only and these indurated. 66

65b. Lowest two bracts of spikelet hyaline to herbaceous, shorter than or as

long as spikelet; each spikelet with 3 or more bracts. 67

66a. Leaf blade 3—7 mm wide, margins serrulate. Spikelets single, laterally

flattened, with 2 bracts (1 lemma and | palea), unawned. (Ponds and

marshes). Leersia hexandra

66b. Leaf blade 21—42 mm wide, margins smooth. Spikelets paired; sessile

spikelet dorsiventrally flattened to subterete, with 5 bracts (2 glumes, 2

lemmas and | palea), awned or unawned; pedicelled spikelets variously

reduced, dorsiventrally flattened. (Dry places). Sorghum bicolor

67a. Spikelets awned, awn exserted beyond spikelet bracts and (0.6—)1—14

mm long; ifawn 0.6—1 mm long then upper glume and 1* lemma densely

hairy, hairs 2—6.5 mm long and pinkish. 68

67b. Spikelets either unawned or awn not exserted beyond spikelet bracts and

up to 0.6 mm long and upper glume and |“ lemma glabrous or with

hairs up to 0.5 mm long and whitish or yellowish. 74

68a. Ligule either 0.05—O.1 mm long and densely fringed or membranous

with hairs at tip absent to shorter than membranous part. 69

68b. Ligule either a row of hairs or membranous with hairs at tip longer than

membranous part. 70

69a. Culms 40—110 cm long. Nodes hairy. Inflorescence with longest branches

4—7 cm long. Spikelets in pairs, the terminal in threes. Callus obtuse.

Awn 13—14 mm long. Bothriochloa bladhii

69b. Culms 15—35 cm long. Nodes glabrous. Inflorescence with longest

branches 1.5—3 cm long. Spikelets in threes, the lower sometimes in

pairs. Callus needle-shaped. Awn up to 4 mm long.

Chrysopogon aciculatus

70a. Ligule in basal third either membranous or of fused hairs. 71

70b. Ligule a row of free hairs. 72

71a. Inflorescence with central rachis absent or shortly developed, branches

(almost) in a single whorl. Lowest bract of spikelet unawned.

Alloteropsis cimicina

30 Supplement of Gardens’ Bulletin Singapore 57 (2005)

71b. Inflorescence with central rachis well developed, branches scattered.

Lowest bract of spikelet awned, awn at least 2 mm long.

Oplismenus compositus

72a. Spikelets densely hairy; hairs pink to red turning silvery at maturity, patent,

at least a few longer than 4(—6.5) mm; awn up to 2 mm long.

Melinis repens

72b. Spikelets (sparsely) hairy; hairs yellowish white, appressed to erecto-

patent, up to | mm long; awn at least 3 mm long. 73

73a. Leaf blade flat, c. 12 mm wide. Spikelet with 1 exserted awn, lowest

bract about half as long as spikelet, second as long as spikelet (excluding

awns). Echinochloa picta

73b. Leaf blade setaceous, c. 0.5 mm wide. Spikelet with 2 exserted awns, the

two lowest bracts as long as spikelet (excluding awns).

Eriachne pallescens

74a. (67) Spikelet at base with a cup-shaped, 0.2—0.7 mm long, often red-

coloured callus. Eriochloa

74b. Spikelet without a cup-shaped callus at the base. 75

75a. Lowest bract of spikelet 0.1—0.7 times as long as spikelet. 76

75b. Lowest bract of spikelet (0.8—)0.9—1 times as long as spikelet. 82

76a. Pedicels usually with 1—9 bristles longer than the spikelet and not falling

off with the spikelets. Setaria

76b. Pedicels without bristles, if hairs present then shorter than spikelet. 77

77a. Spikelets gibbous. Cyrtococcum

77b. Spikelets not gibbous. | 78

78a. Spikelet either 1-flowered or 2-flowered with 2™ floret hyaline and thinner

than |“ floret. 79

78b. Spikelet 2-flowered, 2" floret (more or less) indurated and thicker than

I1* floren: ~ 80

79a. Sheath with (few) transverse veins present. Spikelet (3—)3.9—4.9 mm

long, 2-flowered. (Semi-, aquatic). Hymenachne amplexicaule

79b. Sheath without transverse veins. Spikelet 1.1—2.3 mm long, 1-flowered.

(Terrestrial). . Sporobolus

80a. Second lowest bract of spikelet (= upper glume) 0.4—0.5 times as long

as spikelet. Ottochloa nodosa

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 31

80b. Second lowest bract of spikelet (= upper glume) 0.9—1 times as long as

spikelet. 81

8 1a. Inflorescence branches rounded to triquetrous, 0.1—0.7 mm wide, if patent

then with spikelets well-spaced. Spikelets abaxial, single or paired.

Panicum

81b. Inflorescence branches more or less flattened, 0.8—1.2 mm wide, patent,

with spikelets densely crowded. Spikelets adaxial, at least in middle of

branch paired or in threes. Urochloa mutica

82a. (75) Pedicels and/or spikelets with long silky hairs, hairs 2—12 mm long.

82b. Pedicel and spikelet either glabrous or with hairs up to 0.5 mm long. 84

83a. Leaves before flowering clustered at the base of the culm. Inflorescence

branches not articulated, not breaking up at maturity. Both spikelets of a

pair pedicelled, shortest pedicel at least 1 mm long. Imperata

83b. Leaves before flowering scattered along elongated culm. Inflorescence

branches articulated, breaking up at maturity. One spikelet of a pair

sessile, the other pedicelled. Saccharum

84a. Spikelet 3.5—5.2 mm long. Sheath at least 4 cm long. 85

84b. Spikelet 1.3—2.5(—3.3) mm long, if longer than 3 mm then sheath not

longer than 2 cm. 86

85a. Culm 120—250 cm long. Spikelets in pairs or threes of 1 sessile and | or

2 pedicelled, the pedicelled narrower and less hairy than the sessile

spikelet. Lower glume with many appressed hairs, acute.

Sorghum propinquum

85b. Culm 25—40 cm long. Spikelets in pairs, both pedicelled, similar. Lower

glume glabrous, apiculate. Urochloa glumaris

86a. Ligule row of hairs, 1—4 mm long. Leaf blade 2.5—9 mm wide, base

rounded. Isachne

86b. Ligule membraneous, 0.5 mm long, tip ciliolate. Leaf blade 15—20 mm

wide, base cordate. Panicum brevifolium

Descriptions and Generic Keys

Descriptions are kept short. If not mentioned otherwise, the following applies

to all species: internodes less than 3 mm diameter, transverse veins absent,

pseudopetiole absent, leaf blades parallel-nerved and not distinctly ribbed,

spikelets homomorphous and length measured excludes awns.

32 Supplement of Gardens’ Bulletin Singapore 57 (2005)

A key to the species is given if a genus (i.e. the first part of the scientific

name) is represented by two or more species. Characters that are shared by all

the species of this genus are included in the generic description. There is no

generic description if only one species is present.

Beside the common names, synonyms used in the earlier literature (as

cited in the Introduction) are mentioned. The figures depict the spikelets, unless

mentioned otherwise. The glumes (if present) are always grey and the bar

indicates | mm. The species description is followed by information on whether

it is native in Singapore or not, its global distribution, its first record for

Singapore (as present in the Singapore Herbarium (abbreviated as SING) and

the herbarium of the National University of Singapore (SINU)) and its present

day abundance (and date of last record if now extinct in Singapore). Botanic

Gardens refers to the Singapore Botanic Gardens. In a separate line the ecology

is given. The term Malesia is used for the biogeographical region ranging

from the Isthmus of Kra in Thailand to Bougainville Island in the Solomon

Islands and including Malaysia, Brunei, Indonesia, Timor Leste, Philippines

and Papua New Guinea, but excluding Australia. Finally, any other information

of relevance, including differences with species it has been or might be confused

with is given in notes. Herbarium material is cited as the collector’s name in

italic print followed by the collection number and/or year of collection.

Acroceras Stapf

Acroceras tonkinense (Balansa) C.E.Hubb. was mentioned for the Botanic Gardens (Ridley

1907, as Panicum oryzoides Sw.; Ridley 1925, as A. sparsum Ridl. ex Stapf) but no herbarium

or living material has been found. It differs from A. munroanum in having hairy nodes, blades

with midvein distinct and margins thickened, and spikelets with only the 2’ lemma pinched

at its tip.

Acroceras munroanum (Balansa) Henrard — Fig. 19.

Gilliland 1971: 165. Keng et al. 1998: 154.

A. ridleyi (Hack.) Stapf ex Ridl. 1925: 229.

Culms stoloniferous, up to 55 cm long; nodes glabrous. Sheath rounded, glabrous, margin

hairy. Ligule membranous, 0.2—0.3 mm long, entire or ciliolate. Young leaf blade inrolled.

Blade 3.5—7 cm long, 6—9 mm wide, glabrous, scabrous, base cordate. Inflorescence panicle,

central rachis 3—5 cm long, branches 3—5; longest branch 1.5—2.5 cm long, c. 0.5 mm

wide, simple. Pedicel c. 0.7 mm long. Spikelets in pairs and similar (or single), c. 4.3 mm

long, 1.8—2.5 mm wide, not flattened, 2-flowered, unawned; lowest bract c. 0.7 times as

long as and 2” as long as spikelet; glumes and lemmas at tip laterally flattened, glumes and 1“

floret herbaceous; lower glume c. 3.1 mm long, glabrous; upper glume c. 3.8 mm long, glabrous;

1 lemma and palea present, glabrous; 2" lemma and palea glabrous, indurated.

Figure 18. Lower glume of spikelets of Panicum.

a. collar-shaped, b. ovate.

Figure 19. Acroceras munroanum (Balansa) Henrard.

Figure 20. Alloteropsis cimicina (L.) Stapf.

Figure 21. Arundo donax L.

Field Guide to the Grasses of Singapore (Excluding the Bamboos)

‘ne a Ps

a i ~Be . rn

o_o

34 Supplement of Gardens’ Bulletin Singapore 57 (2005)

Not native; India and Myanmar to Malesia. First record 1899; extinct (1961).

It is probably adventive as it was only found in the Botanic Gardens; however, Ridley

(Ridley s.n., 1899, SING) reports it as wild.

Alloteropsis J.Presl

Alloteropsis cimicina (L.) Stapf — Fig. 20.

Ridley 1925: 223. Gilliland 1971: 198. Keng et al. 1998: 154.

Axonopus cimicinus (L.) P.Beauv.: Ridley 1907: 144.

Culms tufted, 40—60 cm long; nodes glabrous or hairy. Sheath rounded, glabrous to hairy,

external ligule hairy. Ligule membranous with hairs at tip, basal part 0.6 mm long, hairs 1 mm

long. Young leaf blade inrolled. Blade 3—7 cm long, 7—11.5 mm wide, glabrous, base cordate.

Inflorescence panicle, central rachis absent or shortly developed; branches 4—5, (almost) in

a single whorl, 8—14 cm long, c. 0.6 mm wide, lowest 3—7 cm naked, simple (or branched).

Pedicel 1—5 mm long. Spikelets in groups of 2—4, similar but unequally pedicelled, 3—3.5

mm long, 1.2—1.8 mm wide, dorsiventrally flattened, 2-flowered, awned; lowest bract 0.5—

0.7 times as long as and 2" as long as spikelet; glumes and 1*' lemma thin herbaceous; lower

glume 1.6—2 mm long, glabrous, acute to acuminate; upper glume 2.9—3.4 mm long, margin

densely hairy, hairs up to 1.2 mm long; 1“ lemma 2—2.5 mm long, glabrous; |“ palea absent;

2™ Jemma and palea glabrous, indurated; 2™ lemma awn 1—2.7 mm long, scabrous.

Native; Paleotropics. First record 1890. Rare.

Waste places, usually on damp soil.

Arundo L.

Arundo donax L. — Fig. 21.

Gilliland 1971: 51. Keng et al. 1998: 154.

Culms loosely tufted, erect, 150—300 cm long; internodes hollow; nodes glabrous. Sheath

rounded, glabrous. Ligule membranous, 2 mm long (1.5 mm in non-flowering shoot), ciliolate.

Young leaf blade inrolled. Blade c. 40 cm long, 15—40 mm wide, glabrous, sometimes with

white stripes (cv versicolor), base rounded. Inflorescence panicle, central rachis 35—-60 cm

long, branches many (in whorls); longest branch 20—30 cm long, c. 0.7 mm wide, branched.

Pedicel 3—8 mm long. Spikelets single, c. 13 mm long (including awn), c. 5 mm wide, laterally

flattened, 3- or 4-flowered (falling off at maturity), awned; rachilla internodes with short

hairs at tip, between the glumes 0.8 mm long, between the florets 1.7 mm long and articulated;

the two lowest bracts c. 0.8 times as long as spikelet; glumes and lemmas herbaceous, paleas

hyaline; lower glume c. 10.3 mm long, enveloping, glabrous; upper glume c. 10 mm long,

glabrous, midvein 0.5—0.8 mm-long protruding; 1“lemma c. 11.3 mm long, basal half densely

covered with up to 5 mm-long hairs, midvein protruding in c. 3 mm long awn; |* palea c. 5

mm long; 2"—4" florets as 1‘, gradually decreasing in size.

Not native; S. Europe, Asia, now worldwide. First record 1877. Cultivated.

Axonopus P.Beauv.

Culms with stolons, erect part 1O—40 cm long. Sheath more or less flattened, midrib raised,

glabrous, margin hairy. Ligule membranous, 0.4 mm long, densely fringed. Inflorescence

panicle, 1—4 per sheath, central rachis absent or present, at least 2 branches at tip; longest

branch 4—8 cm long, 0.8—0.9 mm wide, simple. Pedicel absent. Spikelets single,

Figure 22. Axonopus compressus (Sw.) P.Beauv.

a. facing the 1* lemma, b. facing the upper glume.

Figure 23. Axonopus fissifolius (Raddi) Kuhlm.

Figure 24. Bothriochloa bladhii (Retz.) S.T.Blake, pair of spikelets.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) B35

36 Supplement of Gardens’ Bulletin Singapore 57 (2005)

dorsiventrally flattened, 2-flowered, unawned; the two lowest bracts as long as spikelet; lower

glume absent; upper glume and 1* lemma herbaceous, (slightly) hairy; 1“ palea absent; 2"

lemma and palea indurated.

la. Nodes hairy. Young leaf blade inrolled. Blade flat, (sparsely) hairy, margin

undulate, with numerous long hairs. A. compressus

1b. Nodes glabrous or sparsely appressed-hairy. Young leaf blade folded along

midrib. Blade folded, glabrous, margin smooth, glabrous or with a few

hairs at the base. A. fissifolius

Axonopus compressus (Sw.) P.Beauv. — Fig. 22, Plate 4.

Carpet Grass, Rumput Tikar

Ridley 1925: 216. Gilliland 1971: 187. Keng et al. 1998: 154. Veldkamp 1997b: 511.

Paspalum platicaulon Poir.: Ridley 1907: 125.

Nodes hairy. External ligule a sparse row of short hairs. Young leaf blade inrolled. Blade 5—

15 cm long, 4—9 mm wide, flat, (sparsely) hairy, margin undulate, base rounded and with

many up to 2 mm-long hairs. Central rachis 1—2 cm long, lower branches 1 or 2. Spikelet

2.3—2.4 mm long, c. | mm wide; upper glume 5 veins, midvein present (sometimes obscure).

Not native; Neotropics, introduced in Africa and Asia. First record 1918 (Botanic

Gardens), in the wild 1955. Common.

In open to partly shaded places.

Introduced between 1895 and 1900 (Purseglove P4073, 1955, SING), but spread rapidly

only after it was used for edging beds in the Botanic Gardens (Ridley 1925: 216). It is now

probably the most common grass species in Singapore, widely used for lawns (including golf

courses). It is less usually called Cow Grass.

For differences with Chrysopogon aciculatus and Paspalum conjugatum see notes

under the respective species.

Axonopus fissifolius (Raddi) Kuhlm. — Fig. 23.

Keng et al. 1998: 154. Veldkamp 1997b: 511.

A. affinis Chase: Gilliland 1971: 189.

Nodes glabrous or sparsely appressed-hairy. External ligule absent. Young leaf blade folded

along midrib. Blade 7.5—18 cm long, 3—5 mm wide, glossy green, folded, glabrous, margin

smooth and glabrous or sparsely hairy at base, base cuneate to slightly rounded. Central rachis

absent to 1 cm long, lower branches 0 or 1. Spikelet c. 2 mm long, 0.6—0.9 mm wide; upper

glume 4 (or 5) veins, midvein absent or obscure.

Not native; Neotropics, introduced in Africa and Australasia. First record 1955. Rare.

In open to partly shaded and moist places, apparently associated with seepage.

Its blades are generally narrower and darker green than those of A. compressus.

Veldkamp (1997b) mentions that the nodes are glabrous.

Bothriochloa Kuntze

Bothriochloa bladhii (Retz.) S.T.Blake — Fig. 24.

Long-leaved Beard Grass

Keng et al. 1998: 157.

Amphilophis glabra (Roxb.) Stapf: Ridley 1925: 209 (excluding Singapore).

Andropogon intermedius R.Br.: Ridley 1907: 166 (excluding Singapore).

B. intermedia (R.Br.) A.Camus: De Wet and Harlan 1970: 339—340. Gilliland 1971: 281.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 37

Culms tufted, erect to geniculate, 40—110 cm long; nodes hairy, lower glabrescent. Sheath

rounded, midrib slightly raised, glabrous. Ligule membranous, 1—1.3 mm long, hairy at abaxial

side, ciliolate. Young leaf blade inrolled. Blade 15—21 cm long, 6—8 mm wide, sparsely

hairy with up to 5 mm-long hairs, base cuneate. Inflorescence panicle, central rachis 2.5—8.5

cm long; branches 13—-21, 4—7 cm long, c. 0.3 mm wide, more or less flattened, furrowed

(at least in upper part of branch), margins hairy, empty at basal 1 cm, simple or the lowest

sometimes branched. Spikelets in pairs of | sessile and 1 pedicelled, the upper in threes.

Sessile spikelet 3.5—3.8 mm long, c. 1 mm wide, dorsiventrally flattened, 2-flowered, awned;

callus obtuse, hairy; the two lowest bracts as long as spikelet; lower glume 3.7 mm long,

basal half and margin upper half hairy; upper glume c. 3.5 mm long, glabrous; 1“ lemma c.

2.4 mm long, hyaline, sterile; 1‘ palea absent; 2"? lemma 1.4—1.6 mm long, hyaline, with

13—14 mm long awn from tip; 2"! palea absent. Pedicel 2—2.5 mm long, furrowed, margins

hairy. Pedicelled spikelet reduced to 2 glumes, unawned.

Native; Paleotropics. First record 1955. Rare.

Weed of open habitats. es

It can be mistaken for Dichanthium annulatum, which differs as follows: branches

and pedicels solid, central rachis 0.5—1.5 cm long and inflorescence with 2—5 branches.

Hybrids with Dichanthium annulatum are known (de Wet and Harlan 1970), but have not

(yet) been found in Singapore.

Cenchrus L.

Weston 1974: 375—380. Doust and Kellogg 2002: 1203—1222.

Culms tufted, erect to geniculate, 20—60 cm long; nodes glabrous. Sheath more or less

flattened, midrib (slightly) raised, glabrous or hairy in upper half marginal area. Ligule either

membranous with hairs at tip or a row of hairs, basal part O—0.2 mm long, hairs 0.5—0.8 mm

long. Young leaf blade inrolled. Blade 7—26 cm long, 6—7 mm wide, base rounded.

Inflorescence spike-like raceme of subsessile burs, 2.5—7 cm long, 1.3—1.5 cm wide. Bur

deciduous; involucre indurated, at base covered with bristles, tip 8-lobed, spikelets 2—4.

Pedicel absent. Spikelets not flattened, 2-flowered, unawned; upper glume and 1“ lemma thin

herbaceous; 2"! lemma and palea herbaceous, glabrous, palea slightly shorter than lemma.

It is most closely related to Pennisetum, which has an involucre of bristles fused only

at base. Future research might show they are in fact one genus (Doust and Kellogg, 2002). In

Singapore Setaria differs from both Cenchrus and Pennisetum in that the bristles do not fall

off together with the spikelet.

la. Burs 1—2 mm apart; basal bristles 0.1—0.2 mm wide, erect, longest up to

7 mm long; lobes at margin with hairs up to 0.5 mm long. Spikelet 4—

5 mm long. C. brownii

1b. Burs 1.5—4 mm apart; basal bristles 0.3—0.5 mm wide, recurved, longest

up to 4 mm long; lobes at margin with at least a few hairs 1—1.5 mm

long. Spikelet 5—6.2 mm long. C. echinatus

Cenchrus brownii Roem. & Schult. — Fig. 25.

Weston 1974: 376. Keng et al. 1998: 157 (incl. C. echinatus). Duistermaat 2004: 30.

Inflorescence in middle dense, burs 1—2 mm apart. Involucre 4—5 mm wide (excluding

spines), 5—6 mm long, lobes at margin with hairs up to 0.5 mm long, bristles up to 7 mm

long, 0.1—0.2 mm wide, erect. Spikelet 4—5 mm long, 1 mm wide; lowest bract c. 0.75 and

2™ 0.9—1 times as long as spikelet; lower glume absent; upper glume c. 3 mm long, glabrous;

1“ lemma c. 3.5 mm long; 1* palea absent; 2° lemma c. 3.5 mm long.

3 8 Supplement of Gardens’ Bulletin Singapore 57 (2005)

Not native; Neotropics, introduced in Africa and tropical (Austral)Asia. First record

1950. Rare.

Weed of waste places.

Cenchrus echinatus L. — Fig. 26, Plate 3.

Weston 1974: 378. Veldkamp 1999b: 232. Duistermaat 2004: 30.

Inflorescence in middle more or less open, burs 1.5—4 mm apart. Involucre 4—6 mm wide

(excluding spines), 6—8 mm long, lobes at margin with at least few hairs 1—1.5 mm long,

bristles up to 4 mm long, 0.3—0.5 mm wide, recurved. Spikelet 5—6.2 mm long, 2 mm wide;

either lowest bract c. 0.85 and 2" 0.9—1 times as long as spikelet, or lowest c. 0.45 and 2™

0.9—1 times as long as spikelet; lower glume absent or c. 2.8 mm long, hyaline; upper glume

5—5.2 mm long, glabrous to minutely hairy at base; 1** lemma 5.3—5.8 mm long; 1* palea

5.3 mm long, thin herbaceous; 2™ lemma c. 6 mm long.

Not native; Neotropics, introduced in Africa, Asia and Australia. First record 1994.

Rather rare.

Weed of waste places.

Centotheca Desv.

Monod de Froideville 1971: 57—60.

Centotheca longilamina Ohwi was mentioned for Singapore by Gilliland (1971: 54), but no

herbarium or living material has been found. It differs from C. lappacea as follows: blades at

least 6.3 times as long as wide and gradually narrowed into symmetric base, spikelets 1—3-

flowered, and the 1“ lemma bristled (Monod de Froideville 1971).

Centotheca lappacea (L.) Desv. — Fig. 27, Plate 5.

Barbed Grass, Rumput Darah

Ridley 1907: 181; 1925: 253. Gilliland 1971: 53. Keng et al. 1998: 158.

Centotheca latifolia Trin.: Monod de Froideville 1971: 59.

Culms tufted, erect, 20—80 cm long; internodes (partially) filled with spongy pith, up to 2.5

mm diameter; nodes glabrous. Sheath rounded, glabrous to sparsely hairy at tip, basal sheaths

reduced. Ligule membranous, |—1.5 mm long. Young leaf blade inrolled. Blade 3.5—11 cm

long, 13—-28 mm wide, transverse veins present, underside glabrous to hairy, upperside

appressed-hairy, margins often transversely wrinkled, base abruptly and asymmetrically

narrowed, pseudopetiole 0.5—0.7 mm long. Inflorescence panicle, central rachis 11—20 cm

long, branches many; longest branch 6—10 cm long, c. 0.2 mm wide, rounded, scaberulous,

simple or branched. Pedicel 2—4 mm long. Spikelets single, 4—5.5 mm long, 1.4—1.9 mm

wide, not to slightly laterally flattened, herbaceous, 2- or 3-flowered, unawned; rachilla

articulate between the florets, 0.7 mm long between 1* and 2" floret, extension above last

floret 1.8 mm long; the two lowest bracts 0.6—0.8 times as long as spikelet; lower glume c.

2.5 mm long, enveloping, glabrous; upper glume 2.7—3 mm long, glabrous; |*' lemma 3.4—

3.5 mm long, glabrous, mucro 0.2 mm long; 1‘ palea 2.2—2.3 mm long; 2" and 3™ lemma

2.5—3.5 mm long, uppermost marginal area at maturity with up to 10 retrorse bristles of up to

0.8 mm long, mucro 0.3 mm long; 2" and 3" palea 1.8—2.7 mm long.

Native; Paleotropics. First record 1894. Rather rare.

Figure 25. Cenchrus brownii Roem. & Schult., bur with an involucre and several spikelets.

Figure 26. Cenchrus echinatus L., bur with an involucre and several spikelets.

Figure 27. Centotheca lappacea (L.) Desv.

Figure 28. Chloris barbata Sw.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 39

POPPE

40 Supplement of Gardens’ Bulletin Singapore 57 (2005)

In forests, partly shaded areas, on moist (to wet) soil.

Non-flowering plants of Lophatherum gracile differ in having blades that are not

wrinkled, pseudopetioles that are much longer (7—20 mm long) and roots with tubers.

Chloris Sw.

Chloris barbata Sw. — Fig. 28, Plate 6.

Plush Grass

Ridley 1907: 173; 1925: 250. Gilliland 1971: 86. Keng et al. 1998: 158.

Culms tufted, erect to geniculate, stolons absent, 35—70 cm long; internodes of non-flowering

plants very short; nodes glabrous. Leaves of non-flowering plants all basal and distinctly

distichous, shoot fan-shaped. Sheath 2.5—7.5 cm long, flattened, midrib winged, tip with

few hairs. Ligule membranous, c. 0.5 mm long, hairy at abaxial side, ciliolate. Young leaf

blade inrolled. Blade 1.5—13 cm long, 1—6 mm wide, in basal half with few up to 3 mm-

long hairs, midvein winged underneath, base cuneate. Inflorescence panicle, central rachis

absent; branches 7—14, 4—7.5 cm long, 0.1—0.2 mm wide, rounded, scaberulous, simple.

Pedicel c. 0.4 mm long, rounded, scaberulous. Spikelets single, in 2 rows, c. 2.3 mm long,

1—1.2 mm wide, not flattened, often tinged purple, 3-flowered with 1* floret bisexual and 2"

and 3™ sterile, awned; rachilla between glumes and 1“ lemma hairy; lowest bract 0.6—0.7

and 2" 0.8—1 times as long as spikelet; lower glume 1.4—1.6 mm long, glabrous, hyaline;

upper glume as lower, 1.7—2.5 mm long; 1* lemma 1.7—2.2 mm long, thick herbaceous,

along midvein and upper half marginal veins with up to 1 mm long hairs, truncate, awned just

below tip, awn 3—5 mm long and scaberulous; 1‘ palea 1.9—2 mm long; 2"? and 3 lemma

0.9—1.1 mm long, (thin) herbaceous, truncate, awn 3—5 mm long, scaberulous; 2™ and 3"

palea absent.

Not native; Neotropics, introduced in Africa, Asia and Australia. First record 1892.

Common.

Weed of waste places on well-drained soil, including reclaimed land, roadsides, etc.

Ridley (1925) already reported it as well established.

For differences with Eustachys tenera see note under that species.

Chrysopogon Trin.

Veldkamp 1999a: 503—533.

Vetiveria Bory

Sheath glabrous, at least one margin hairy. Young leaf blade inrolled. Inflorescence panicle,

central rachis present. Sessile spikelet 2-flowered; 1“ palea absent.

Veldkamp (1999a) argues that Vetiveria and Chrysopogon are best united. Although

the two species present in Singapore look very different, intermediate characters of a number

of African and Australian species seriously blur the generic delimitation.

la. Culms 15—35 cm long, with short stolons. Blade 1.5—9 cm long. .

C. aciculatus

1b. Culms 150—250 cm long, with short rhizomes. Blade 25—120 cm long.

C. zizanioides

Figure 29. Chrysopogon aciculatus (Retz.) Trin., group of three spikelets, one sessile and

two pedicelled. 7

Figure 30. Coix lacryma-jobi L., inflorescence.

a. involucre with exserting stigma, b. male spikelets.

Figure 31. Cymbopogon martini (Roxb.) Will.Watson.

a. sessile spikelet, b. pedicelled spikelet.

Field Guide to the Grasses of Singapore (Excluding the Bamboos)

42 Supplement of Gardens’ Bulletin Singapore 57 (2005)

Chrysopogon aciculatus (Retz.) Trin. — Fig. 29, Plate 17.

Love Grass, Kemucut

Ridley 1925: 207. Gilliland 1971: 236. Keng et al. 1998: 158. Veldkamp 1999a: 509.

Andropogon aciculatus Retz.: Ridley 1907: 166.

Culms with short and densely leaved stolons, erect part 15—35 cm long; nodes glabrous.

Sheath 1—2.5 cm long, rounded. Ligule 0.05—0.1 mm long, densely fringed. Blade linear-

lanceolate to linear, 1.5—9 cm long, 4—6 mm wide, glabrous, base cuneate to rounded with

margins (sparsely) hairy. Inflorescence with central rachis 6—9 cm long, branches in whorls

of 2—8; longest branch 1.5—3 cm long, c. 0.2 mm wide, rounded, simple or branched. Spikelets

in threes of 1 sessile and 2 pedicelled, lower sometimes in pairs. Sessile spikelet 3..—4 mm

long, 0.5 mm wide, not flattened, awned; callus needle-shaped, 3.6—6.4 mm long, short-

hairy; the two lowest bracts 0.9—1 times as long as spikelet; lower glume c. 3.5 mm long,

herbaceous, hairy on midvein and margins, tip bifid; upper glume as lower, c. 4 mm long, tip

acuminate, mucro c. 0.6 mm long; 1* lemma 2.5 mm long, sparsely hairy, hyaline; 2"? lemma

c. 2.2 mm long, glabrous, hyaline, awn c. 3.8 mm long, scaberulous; 2™ palea c. 0.4 mm long.

Pedicel c. 3 mm long, flattened, hairy. Pedicelled spikelet male, unawned.

Native; S. and E. Asia to Australia. First record 1880s. Common.

In open places, lawns, roadsides, etc.

Non-flowering plants of Axonopus compressus differ in the hairy nodes, the presence

of a hairy external ligule, the longer ligule, and the generally longer blades that are (sparsely)

hairy on both sides.

Chrysopogon zizanioides (L.) Roberty

Vetiver, Rumput Wangi

Veldkamp 1999a: 522.

Vetiveria odorata Virey: Ridley 1925: 208.

V. zizanioides (L.) Nash: Gilliland 1971: 232. Keng et al. 1998: 187. Ng 1999: 288.

Culms with short and scented rhizomes, internodes of non-flowering plants very short; nodes

unknown. Leaves of non-flowering plants all basal. Sheath c. 12 cm long, keeled, transverse

veinlets present. Ligule of basal leaves 2 mm long and lacerate, of higher leaves 0.2 mm long

and densely fringed. Blade very long-linear, 25—-120 cm long, 4—5 mm wide, glabrous, with

strong and white midrib, transverse veins present, margins serrate, base cuneate and glabrous

to densely hairy.

Not native; Africa to Indochina, introduced in America. First record 1929. Cultivated.

It is only known from the Botanic Gardens (at present in the Evolution Garden) where

it has never flowered. I have not seen the culms but according to Veldkamp (1999a) they are

150—250 cm long and with glabrous nodes.

This species is cultivated for the rhizomes that contain a fragrant oil, used in perfumes

and in curries. Because of its soil-binding capacity it has been planted along the new highways

in Malaysia (Ng 1999).

Coix L.

Coix lacryma-jobi L. — Fig. 30.

Job’s Tears, Jelai

Ridley 1907: 151; 1925: 191. Gilliland 1971: 304. Keng et al. 1998: 159.

Culms single or tufted, erect, 60—200 cm long; internodes solid; nodes glabrous. Sheath

rounded, somewhat inflated, glabrous. Ligule membranous, c. 0.5 mm long, ciliolate. Young

leaf blade inrolled. Blade 17—42 cm long, 24—40 mm wide, glabrous, margin with minute

hairs, base rounded. Inflorescence | per sheath, many per culm, spike-like raceme, rachis

3.5—6 cm long. Spikelets unisexual, unawned, basal group enclosed in involucre, upper group

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 43

free. Involucre of basal group much indurated bead-like sheath, c. 10 mm long, to 6.5 mm

wide, shiny grey, enclosing | female and 2 sterile spikelets. Rachis continuing through pore

at tip of involucre. Pedicel upper spikelets c. 3.5 mm long, glabrous, flattened. Upper spikelets

in 2 pairs of | sessile and | pedicelled, otherwise similar, 10.5—11 mm long, 3—3.5 mm

wide, dorsiventrally flattened, 2-flowered, male, unawned; the two lowest bracts (almost) as

long as spikelet; lower glume 10—10.5 mm long, herbaceous, glabrous, submarginally winged,

wings up to | mm wide and widest at tip; upper glume 8.5—9.7 mm long, herbaceous, narrowly

winged on midvein, upper half of wing with short hairs; 1‘ lemma 8.2—9.5 mm long, glabrous,

hyaline, tip bifid; 1“ palea 7.5—-8.8 mm long; 2™ lemma and palea as 1“, slightly shorter.

Not native; tropical Asia, now worldwide. First record 1889. Rare.

On roadsides, waste places, in secondary vegetation.

According to Ridley (1925) it is not wild. Cultivated as a cereal and escaping. Involucres

of lower spikelets are used as beads.

Cymbopogon Spreng.

Gilliland 1971: 294—299. Soenarko 1977: 225—375.

Culms tufted, erect, when flowering up to 300 cm long; nodes at base densely crowded and

covered by sheaths. Sheaths of non-flowering culms overlapping, rounded, swollen at base,

glabrous. Ligule membranous, |.5—2.5(—5) mm long. Young leaf blade inrolled. Blade up

to 100 cm long, 8—20 mm wide, glabrous except for few hairs behind ligule, margins sharply

serrate, base cuneate (cordate in C. martini). Spatheate inflorescence up to 50 cm long, much

branched with branches up to 20 cm long, final sheaths (spathes) up to 2.5 cm long.

Inflorescence panicle, central rachis absent; branches 2, up to 2 cm long, c. 0.2 mm wide,

simple. Spikelets in pairs of 1 sessile and | pedicelled. Sessile spikelet not to slightly

dorsiventrally flattened, 2-flowered, awned or unawned; the two lowest bracts as long as

spikelet; lower glume flat, with 2 narrow submarginal keels; upper glume boat-shaped, midvein

keeled. Pedicel 2.5—3 mm long, flattened, hairy. Pedicelled spikelet somewhat smaller,

unawned.

Not native; Paleotropics. Cultivated.

Several species are cultivated for aromatic essential oils. In Singapore they are rarely

seen flowering and only known from gardens. As plants might survive for a long time in

abandoned plots, an annotated key to the species is provided.

la. Leaf blade at base cordate. Lower glume of sessile spikelet with deep

longitudinal furrow in its lower half. (India, two forms cultivated in the

tropics). Grown for palmarosa oil and ginger oil. Ginger Grass.

C. martini (Roxb.) Will.Watson — Fig. 31.

lb. Leaf blade at base cuneate. Lower glume of sessile spikelet without such a

furrow, but sometimes broadly hollow near the base. 2

2a. Sessile spikelet unawned or awn reduced to a bristle and not visible from

outside spikelet. | 5

2b. Sessile spikelet awned, awn visible from outside spikelet. 4

3a. Lower glume of sessile spikelet 6 mm long, very narrow, without distinct

veins between the keels, basal part depressed. Basal sheaths whitish

green, persistent. (Origin unknown, cultivated throughout the tropics).

Used for flavouring food. Lemon Grass, Serai. C. citratus (DC.) Stapf

44 Supplement of Gardens’ Bulletin Singapore 57 (2005)

3b. Lower glume of sessile spikelet 4—5 mm long, with distinct veins between

the keels, basal part not depressed. Basal sheaths purple, caducous. (India

and Ceylon, introduced elsewhere in the tropics). Grown for citronella

oil. Citronella Grass, Serai Wangi.

C. nardus (L.) Rendle var. nardus — Fig. 32.

4a. Lower glume of pedicelled spikelet 7(—9)-veined, upper glume 3(or more)-

veined. Basal sheaths yellowish. (Malay Peninsula, Thailand, on

limestone). Unscented. C. calcicola C.E.Hubb. — Fig. 33.

4b. Lower glume of pedicelled spikelet 1—3-veined, upper glume 1-veined.

Basal sheaths green. (India, naturalized in Indonesia, cultivated in the

tropics). Malabar Grass.

C. flexuosus (Nees ex Steud.) Will.Watson — Fig. 34.

Cynodon Pers.

Nowack 1992: 477—478.

Gilliland (1971) mentioned Cynodon arcuatus J.Pres] (a later synonym for C. radiatus

Roth in Roem. & Schult.) in the synonymy of C. dactylon, but according to Nowack (1992)

they are different:

la. Rhizomes present. Culms 0.1—0.2 m long. Blades 1.5—2.5 mm wide. Spikes 3—6, 1—

6 cm by 1.2—2 mm, not wavy. Anthers 0.8—1.5 mm long. C. dactylon

1b. Rhizomes absent. Culms 0.2—0.4 m long. Blades 4—7 mm wide. Spikes 5—8, 6—12 cm

by 0.6—1.5 mm, wavy. Anthers 0.5—0.7 mm long. (Not yet known from Singapore.)

C. radiatus

Cynodon dactylon (L.) Pers. — Fig. 35, Plate 7.

Bermuda Grass, Rumput Bermuda

Ridley 1907: 173, p.p.; 1925: 249, p.p. Gilliland 1971: 90. Keng er al. 1998: 160.

Culms with long rhizomes or stolons, rarely tufted without stolons, erect part 10—25 cm

long; nodes glabrous. Sheath rounded, glabrous or throat with few hairs. Ligule row of hairs

of unequal length, 0.2—1 mm long with longest hairs at least 0.5 mm long. Young leaf blade

inrolled to seemingly folded along midrib. Blade 0.5—4.5 cm long, 1.5—2.5 mm wide,

glabrous or underside in middle short-hairy, base cuneate. Inflorescence panicle, central rachis

absent; branches 2—5, 1.5—5 cm long, c. 0.5 mm wide (0.8—1.7 including spikelets), flattened

to V-shaped, shortly winged, hairy at base, simple. Pedicel absent. Spikelets single, alternate

on branch, 2.4—2.5 mm long, 1.3—1.9 mm wide, laterally flattened, yawning or not, 1(—2)-

flowered, unawned; the two lowest bracts 0.6—0.7 times as long as spikelet; lower and upper

glume 1.4—1.7 mm long, often erecto-patent, herbaceous, sparsely hairy, acute; 1** lemma

2.1—2.4 mm long, thin herbaceous, hairy on midvein and margins; palea 2—2.2 mm long;

2™ floret absent or much reduced, on elongated rachilla. Anthers (0.5—) 0.8—1.3 mm long.

Native; tropical and temperate regions of the world. First record 1894. Common.

In lawns, open grasslands, on seashores and sandy soil.

Figure 32. Cymbopogon nardus (L.) Rendle var. nardus, pair of spikelets.

Figure 33. Cymbopogon calcicola C.E.Hubb, pair of spikelets.

Figure 34. Cymbopogon flexuosus (Nees ex Steud.) Will.Watson, pair of spikelets.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 45

Fig. 33

46 Supplement of Gardens’ Bulletin Singapore 57 (2005)

The branches of the inflorescence are generally much narrower than Nowack (1992)

reports. Plants that share with C. radiatus a tufted habit and anthers of 0.5—0.7 mm long but

are otherwise as C. dactylon are here included in C. dactylon.

Differences with Zoysia matrella are mentioned there.

Cyrtococcum Stapf

Culms with stolons, lower nodes rooting. Sheath rounded, not shouldered. Ligule

membraneous, glabrous. Young leaf blade inrolled. Blade (sparsely) hairy, base cuneate (to

rounded), margins with long hairs. Inflorescence a panicle, central rachis present; branches

0.1—0.4 mm wide. Pedicel glabrous, rounded. Spikelets in pairs with pedicels of unequal

length, more or less laterally flattened, gibbous, 2-flowered with 1“ floret sterile and 2"

bisexual, unawned; lowest bracts 0.5—0.7 and 2" lowest 0.6—0.8 times as long as spikelet;

both glumes thin herbaceous; I‘ lemma (thin) herbaceous; I“ palea absent; 2" floret thick

herbaceous.

Differences with /sachne are mentioned there. The spikelets of Panicum are not gibbous,

and its upper glumes are at least 0.9 times the length of the spikelet.

Cyrtococcum oxyphyllum, earlier mentioned for Singapore (Ridley 1925: 233, Gilliland

1971: 148, Keng et al. 1998: 161) has not been found. Differences with both C. accrescens

and C. patens include a contracted inflorescence and that the longest pedicel of the pair is

shorter than the spikelet.

Veldkamp (pers. comm.) provided the key to the species. However, intermediates occur

(Duistermaat 2004: 31).

la. Erect part of culms 30—100 cm long. Sheath usually hairy all over. Panicle

20—50 by 6—30 cm. Spikelets usually 1.35—1.5 mm long.

C. accrescens

1b. Erect part of culms 10—30 cm long. Sheath usually hairy along margins

only. Panicle 3—18 by 0.8—2.3 cm. Spikelets usually 1.5—1.8 mm

long. C. patens

Cyrtococcum accrescens (Trin.) Stapf — Fig. 36.

Diffuse Panic Grass, Rumput Telur Ikan

Ridley 1925: 234. Gilliland 1971: 149. Keng er al. 1998: 160.

Panicum patens auct., non L.: Ridley 1907: 142.

Culms erect part 30—100 cm long; nodes glabrous. Sheath 3—5.5 cm long, sparsely hairy all

over. Ligule 1.5 mm long. Blade 7—14 cm long, 10—14 mm wide. Inflorescence 20—35 cm

long, 8—17 cm wide; longest branches 11—18 cm long, simple or branched. Longest pedicel

of the pair 5.5—7.5 mm long. Spikelet 1.3—1.5 mm long, 0.7—1 mm wide.

Native; S. and E. Asia. First record 1885. Rare.

In shaded places on damp soil.

Figure 35. Cynodon dactylon (L.) Pers.

Figure 36. Cyrtococcum accrescens (Trin.) Stapf.

a. pair of spikelets, b. spikelet.

Figure 37. Cyrtococcum patens (L.) A.Camus.

a. pair of spikelets, b. spikelet.

Figure 38. Dactyloctenium aegyptium (L.) Willd.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 47

a ce

sate

48 Supplement of Gardens’ Bulletin Singapore 57 (2005)

Cyrtococcum patens (L.) A.Camus — Fig. 37.

Gilliland 1971: 148. Duistermaat 2004: 31.

C. carinatum Stapf ex Ridl. 1925: 233 (excluding Singapore).

Culms erect part 1O—30 cm long; nodes hairy at one side. Sheath 1.5—2 cm long, with hairy

margins only. Ligule 0.4—0.8 mm long. Blade 4—7 cm long, 5—8 mm wide. Inflorescence

4—9 cm long, 1—2.3 cm wide; longest branch 3—5 cm long, branched. Longest pedicel of

the pair 2.5—4 mm long. Spikelet 1.5—1.8 mm long, 0.9—1 mm wide.

Native; S.E. Asia. First record 1993. Rare.

In secondary or primary forest on damp soil.

Although it was found only rather recently, it is probably native.

Dactyloctenium Willd.

Dactyloctenium aegyptium (L.) Willd. — Fig. 38, Plate 30.

Egyptian Finger Grass

Ridley 1925: 251. Gilliland 1971: 81. Keng et al. 1998: 161.

Eleusine aegyptia (L.) Desf.: Ridley 1907: 174 (as E. aegyptiaca).

Culms with stolons, erect part 20—30 cm long; nodes glabrous. Sheath rounded, sparsely

hairy, margins hyaline and at throat with few long hairs. Ligule membranous, 0.6—0.8 mm

long (including hairs at tip), hairs at tip 0.2—0.3 mm long. Young leaf blade inrolled. Blade

5—15 cm long, 5—6 mm wide, (sparsely) hairy, base rounded and hairy at margin.

Inflorescence panicle, central rachis absent; branches 3—5, 1.5—4.5 cm long, 0.9 mm wide

(including spikelets 5—7 mm wide), sparsely hairy, naked at uppermost 1—2.5 mm, simple.

Pedicel absent. Spikelets single, alternately in 2 dense rows, the uppermost in | row, 3.5—4

mm long, 3—3.5 mm wide, laterally flattened, (thin) herbaceous, 3- or 4-flowered, awned;

the two lowest bracts 0.7—0.75 times as long as spikelet; lower glume 2.5—2.8 mm long,

midvein keeled in lower half, sparsely hairy, tip truncate; upper glume 1.8—2 mm long,

glabrous, truncate, midvein scaberulous and protruding into 1.5—2 mm long awn; 1“ lemma

3.1—3.4 mm long, glabrous, vein | and keeled in upper half, acuminate; 1“ palea 2—2.3 mm

long; 2" “*" lemma and palea as 1“, but gradually smaller.

Native; Paleotropics. First record 1893. Common.

Weed on well-drained soil on roadsides, beaches, in lawns, waste places.

Dichanthium Willemet

Sheath glabrous. Ligule membranous, hairy at abaxial side. Young leaf blade inrolled (unknown

for D. mucronulatum). Spikelets in pairs of 1 sessile and 1 pedicelled. Sessile spikelet

dorsiventrally flattened, 2-flowered, awned; callus short-hairy; the two lowest bracts as long

as spikelet; glumes (thin) herbaceous, lower flattened and submarginally keeled to winged in

upper half, upper boat-shaped; lemmas hyaline, glabrous; 2" lemma awned from tip, awn

scaberulous and twisted; paleas absent. Pedicel (somewhat) flattened, hairy on one or both

margins. Pedicelled spikelet slightly smaller than sessile, 2 glumes (and hyaline lemmas)

present, unawned.

la. Nodes (densely) hairy. Inflorescence a panicle, branches 2—5S.

D. annulatum

1b. Nodes glabrous to sparsely hairy. Inflorescence spike-like raceme, simple.

Figure 39. Dichanthium annulatum (Forssk.) Stapf, pair of spikelets.

Figure 40. Dichanthium caricosum (L.) A.Camus, pair of spikelets.

Field Guide to the Grasses of Singapore (Excluding the Bamboos)

(nmin ati nani soarn

eel

50 Supplement of Gardens’ Bulletin Singapore 57 (2005)

2a. Upper blades hairy, base rounded. Lower glume of sessile spikelet truncate

and marginal veins protruding. D. caricosum

2b. Upper blades glabrous, base long cuneate. Lower glume of sessile spikelet

with 3-pointed tip. D. mucronulatum

Dichanthium annulatum (Forssk.) Stapf — Fig. 39.

Gilliland 1971: 284. Duistermaat 2004: 32.

Culms tufted, geniculate to decumbent, 50—200 cm long; nodes (densely) hairy. Sheath 3—

7 cm long, more or less rounded, midrib not to slightly raised. Ligule 1.5 mm long. Blade 7—

17 cm long, 3—6 mm wide, at least upperside (sparsely) hairy, base rounded. Inflorescence

panicle, central rachis 0.5—1.5 cm long; branches 2—5, 3.5—7 cm long, 0.4 mm wide,

rounded, axils hairy, simple. Sessile spikelet 3.5—3.9 mm long, 1—1.2 mm wide; lower

glume 3.4—3.8 mm long, margin and lower half hairy, with 9 veins, truncate; upper glume

3.5—4 mm long, margin and marginal veins sparsely hairy, acute; 1‘ lemma 3 mm long; 2"

lemma 1.6 mm long, 0.1 mm wide, awn 21—23 mm long. Pedicel 1.8—2 mm long.

Native; tropical Africa and South East Asia. First record 1932. Rare.

Weed in lawns, on roadsides, brick walls.

Gilliland (1971) reported that it grows on limestone, probably because he included

Ischaemum beccarii (see D. mucronulatum) in the synonymy.

Differences with Bothriochloa bladhii are mentioned there.

Dichanthium caricosum (L.) A.Camus — Fig. 40.

Ridley 1925: 210 (excluding Singapore). Gilliland 1971: 284. Keng et al. 1998: 162.

Culms geniculate, up to 35 cm long; nodes sparsely hairy. Sheath 2 cm long, rounded and

keeled at tip. Ligule 0.5 mm long. Blade 2.5—4 cm long, 2—3 mm wide, (sparsely) hairy,

base rounded and with prominent midvein. Inflorescence spike-like raceme, 2.5—3 cm long.

Sessile spikelet 3.5 mm long, 0.8 mm wide; lower glume 3.5 mm long, hairy on lower 7",

with 5 veins, truncate and marginal veins protruding; upper glume 3 mm long, hairy, with 3

veins, emarginate; 1*' lemma 1.8 mm long; 2™ lemma 1.3 mm long, awn 6.5—8 mm long.

Pedicel 2 mm long.

Not native; Paleotropics, Pacific Islands to Fiji. Single record 1932 from Newton.

Extinct (1932).

It is probably not native, as it prefers a pronounced dry season (Gilliland 1971).

Dichanthium mucronulatum Jansen

Gilliland 1971: 283. Keng et al. 1998: 161.

Ischaemum beccarii auct., non Hack.: Ridley 1907: 160; 1925: 204.

Nodes glabrous. Ligule 1—1.5 mm long. Blade 8—17 cm long, 4—5 mm wide, glabrous,

base long cuneate. Inflorescence spike-like raceme, 4—5 cm long. Sessile spikelet 4.5—5

mm long; lower glume indurated, with few short appressed hairs in upper '/, tip bifid,

mucronate; 2"* lemma awn c. 3 cm long. Pedicel 1.3 mm long.

Not native; Malaysia: Langkawi, on limestone. Single record 1897 from the Botanic

Gardens. Extinct (1897).

Weed.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 51

Digitaria Haller

Veldkamp 1973: 1—80.

Sheath (more or less) rounded, midrib raised or not, submarginally hyaline, margin glabrous.

Ligule membranous, glabrous, erose. Young leaf blade inrolled. Blade midvein inconspicuous

or at upperside white and/or at underside raised. Inflorescence panicle, central rachis absent

or present with at least 2 branches from tip of peduncle or central rachis; branches winged,

simple. Pedicel glabrous. Spikelets in groups of 2—3(—5) on one side of branch, at least 1

pedicelled, more or less dorsiventrally flattened, 2-flowered with 1“ floret sterile and 2"

bisexual, unawned; lower glume absent or glabrous; upper glume and 1|* lemma (thin)

herbaceous; 1“ palea absent; 2°* lemma and palea indurated, glabrous.

Because the lowest spikelet of a group is sometimes aborted, it is important to check

the number of spikelets in several groups throughout inflorescence. The rachis is considered

distinctly serrate if the spicules are visible with 10x magnification.

la. Spikelets 2—3.5 mm long, all in pairs. Abscission of pedicelsruncate. 2

1b. Spikelets 1.3—2 mm long, at least in the middle of the branches in groups

of 3(—5), sometimes apparently in pairs, then pedicel of third spikelets

adnate to the rhachis and this spikelet apparently solitary, or pedicels

with an abortive spikelet at base. Abscission of pedicels discoid to cup-

shaped. 10

2a. Lower glume absent to very small, up to 0.25 mm long (see note D. ciliaris).

2b. Lower glume distinct, at least 0.25 mm long. (i

3a. Margins of rhachis smooth to serrulate, teeth up to 0.03 mm long.

D. radicosa

3b. Margins of rhachis distinctly serrate, teeth at least 0.05 mm long. ~

4a. Spikelets either homomorphous and all 1“ lemmas in the second interspace

with yellowish bristles (appressed at flowering time!), or at least upper

pair of branch heteromorphous, i.e. 1‘ lemma of the sessile spikelet either

glabrous and veins equidistant or slightly pubescent and veins not

equidistant, that of the pedicelled spikelet always more pubescent to

bristled, veins not equidistant. (The basal spikelets may be

homomorphous, then completely glabrous!). 5

4b. All spikelets homomorphous, with a hairy fringe, but never bristled. 6

Sa. Upper glume 1—2.75 mm long, 0.35—0.8 times as long as the spikelet.

Bristles of the 1“ lemma absent or obscured by the other pubescence,

especially during flowering time. D. bicornis

5b. Upper glume 0.3—1 mm long, 0.15—0.3 times as long as the spikelet.

Bristles of the 1** lemma present, at most with only a few hairs between

them. D. setigera var. calliblepharata

52 Supplement of Gardens’ Bulletin Singapore 57 (2005)

6a. Spikelet 2—2.5 mm long. Upper glume 0.4—0.8 times as long as the

spikelet, 1—1.5 mm long. Anthers 0.3—0.6 mm long. D. nuda

6b. Spikelet (2—)2.8—3.1 mm long. Upper glume 0.2—0.25(—0.4) times as

long as the spikelet, usually much less, 0.25—1.25 mm long. Anthers

0.65—1.25 mm long. D. setigera var. setigera

7a. (2) Blades up to 2 mm wide. Spicules of pedicels often more or less hair-

like. D. didactyla

7b. Blades 2—9 mm wide. Spicules of pedicels absent to triangular. 8

8a. Spikelets either homomorphous and bristled, or at least upper pair of branch

heteromorphous: i.e. 1** lemma of the sessile spikelet either glabrous

and veins equidistant or slightly pubescent and veins not equidistant,

that of the pedicelled spikelet always more pubescent to bristled, veins

not equidistant. D. bicornis

8b. Spikelets homomorphous, never bristled. 9

9a. Rhachis distinctly serrate, spicules at least 0.05 mm long. D. ciliaris

9b. Rhachis smooth to serrulate, spicules up to 0.03 mm long. D. radicosa

10a. (1) Spikelets glabrous. 2’ lemma protruding (at maturity), pale yellow to

yellow-brown in fruit, often with a bluish tip. D. fuscescens

10b. Spikelets puberulous to pubescent, (appressed) hairs present at least along

the margins of the upper glume and 1* lemma. 2"* lemma not protruding,

pale yellow to dark brown in fruit. 11

lla. Plant tufted, culms erect to geniculate, not stoloniferous. Inflorescence

with 2—10(—14) branches, central rachis usually well-developed, 0.3—

3 cm long. Pedicels sparsely to densely serrate. 2°? lemma dark brown to

black in fruit. . D. violascens

11b. Plant stoloniferous. Inflorescence with 2—4 branches, central rachis only

rarely developed, then up to 1.5 cm long. Pedicels smooth to sparsely

serrate. 2"' lemma pale yellow to dark yellow-brown in fruit. 12

12a. Spikelets 1.3—1.7(—1.9) mm long. Upper glume as long as the spikelet.

Figure 41. Digitaria bicornis (Lam.) Roem. & Schult., pair of heteromorphous spikelets.

a. facing lower glume, b. facing upper glume.

Figure 42. Digitaria ciliaris (Retz.) Koeler.

a. facing upper glume, b. facing lower glume.

Figure 43. Digitaria didactyla Willd.

a. facing lower glume, b. facing upper glume.

Figure 44. Digitaria fuscescens (J.Presl) Henrard.

a. facing 1 lemma, b. facing upper glume.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 33

54 Supplement of Gardens’ Bulletin Singapore 57 (2005)

1* lemma with 5—7 more or less inequidistant veins, obtuse.

D. longiflora

12b. Spikelets 1.8—2 mm long. Upper glume 0.7—1 times as long as the

spikelet. 1‘ lemma with 7 equidistant veins. D. mollicoma

Digitaria bicornis (Lam.) Roem. & Schult. — Fig. 41.

Veldkamp 1973: 30. Duistermaat 2004: 32.

Paspalum sanguinale (L.) Lam. var. ciliare auct., non Hook.f.: Ridley 1907: 125 (excluding

Singapore).

Culms tufted, erect to ascending, 30—60 cm long; nodes sparsely hairy. Sheath 4—9 cm

long, glabrous to hairy. Ligule 2 mm long. Blade 4—14 cm long, 2—9 mm wide, glabrous or

hairy at upperside, base rounded. Inflorescence with central rachis 0—5S cm long; branches

2—5, 7—11 cm long, 0.9 mm wide, glabrous, serrate. Pedicel 2 mm long, serrate. Spikelets

in pairs, homomorphous in lower part, heteromorphous in upper 1/2—1/3 of rachis. Sessile

spikelet 2.7—3.5 mm long; lowest bract 0.03—0.15 and 2" lowest 0.35—0.7 times as long as

spikelet; lower glume 0. 1—0.4 mm long; upper glume 1—2.5 mm long (excluding overtopping

hairs), with up to 1 mm long silky hairs; 1“ lemma 2.8—3 mm long, either glabrous and

equidistantly 7-veined, or slightly pubescent and inequidistantly veined (pedicelled spikelet

then always pectinate), acute; 2™ lemma 2.6—2.7 mm long, acuminate, yellowish in fruit.

Pedicelled spikelet with 1* lemma inequidistantly veined, pectinate, hairs 0.5—1 mm long,

bristles absent or 1.5—-2 mm long.

Native; (sub-)tropics. First record 1890. Rare.

In waste places, often on sand, also along the shore.

It is readily recognized when the upper pedicelled spikelets, which fall off early, are

still present; otherwise it closely resembles D. nuda (lower glume generally absent) and D.

ciliaris (anthers 0.6—1 mm long vs 0.5—0.6 mm in D. bicornis).

Digitaria ciliaris (Retz.) Koeler — Fig. 42.

Common Crab Grass

Gilliland 1971: 196. Veldkamp 1973: 32. Keng et al. 1998: 162.

D. marginata Link: Ridley 1925: 213.

D. marginata Link var. ciliaris Hook.f. ex Ridl. 1925: 214, p.p.

Paspalum sanguinale auct., non Lam. (incl. var. commutatum auct., non Hook.f.): Ridley

POOR? 5, pp.

Culms tufted, erect to ascending, 30—65 cm long; nodes glabrous. Sheath 7—11 cm long,

glabrous or the lower at base patent hairy. Ligule 1—3 mm long. Blade 3—15 cm long, 3

7(—10) mm wide, glabrous to hairy, base cuneate to rounded and with up to 5 mm long hairs.

Inflorescence with central rachis 0.6—4 cm long; branches 2—6, 5—15 cm long, 0.8 mm

wide, serrate. Pedicel 0.8—-3 mm long, serrate. Spikelets in pairs, homomorphous, 2.5—3.5

mm long, 0.6 mm wide; lowest bract 0.1—0.2 and 2™' 0.5—0.8 times as long as spikelet;

lower glume (0.15—)0.25—0.5 mm long (see note); upper glume 1—2.4 mm long, 3 veins,

sparsely hairy at margin, acute; 1“ lemma 2.5—-2.7 mm long, inequidistantly 7-veined, sparsely

hairy, acute; 2™ lemma 2.5 mm long, acute, yellowish in fruit.

Native; tropics. First record 1880s. Common.

Weed of lawns and open places.

The length of the lower glume varies within a branch, but at least a few are 0.25 mm

long or longer. Apart from the absence of the lower glume, D. nuda is different in that the 1

lemma is (almost) equidistantly veined. For difference with D. bicornis see there.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 55

Digitaria didactyla Willd. — Fig. 43.

Serangoon Grass, Rumput Serangoon

Gilliland 1971: 195. Veldkamp 1973: 44. Keng et al. 1998: 162.

D. marginata Link var. debilis auct., non Ridl.: Hook.f. ex Ridley 1925: 214, p.p.

Paspalum sanguinale Lam. var. debile auct., non Hook.f.: Ridley: 1907: 125, p.p.

Culms densely tufted and with stolons, 20—40 cm long; nodes glabrous. Sheath 4—5 cm

long, lower at least at base (densely) hairy and upper glabrous. Ligule (0.5—) 1.5—2.2(—2.5)

mm long. Blade (0.6—)3—8 cm long, 0.6—1.7(—2) mm wide, glabrous or upperside short-

hairy, base cuneate to rounded and short-hairy. Inflorescence with central rachis O—1 cm

long; branches 2—4, (1—)4—6 cm long, 0.5 mm wide, glabrous, at base smooth and upwards

serrate with hairlike spicules. Pedicel up to 1.5 mm long, serrate. Spikelets in pairs,

homomorphous, 2—2.5 mm long, 0.6 mm wide; lowest bract 0.1—0.2 and 2"* 0.5—0.7 times

as long as spikelet; lower glume 0.2—0.5 mm long; upper glume 1—2 mm long, hairy, 3

veins, acute; |‘ lemma 2.4—2.5 mm long, hairy, more or less inequidistantly 7-veined, acute;

2" Jemma 2.1—2.3 mm long, acute, yellowish to greyish in fruit.

Not native; Africa, elsewhere cultivated and naturalized. First record F892 from the

Botanic Gardens, 1932 in the wild. Rather rare.

In lawns, on roadsides.

Introduced and used as a lawn grass. Elsewhere known as Blue Couch.

Digitaria fuscescens (J.Presl) Henrard — Fig. 44.

Gilliland 1971: 193. Veldkamp 1973: 61. Keng et al. 1998: 162.

Paspalum longiflorum auct., non Retz.: Ridley 1907: 126 p.p. — D. longiflora auct., non

Pers.: Ridley 1925: 214, p.p.

Culms with long stolons, erect part 15—40 cm long; nodes hairy. Sheath 1.1—1.7 cm long,

glabrous or with few hairs at tip. Ligule 0.5—2 mm long. Blade 1—5 cm long, 2—5 mm

wide, glabrous to sparsely hairy, base rounded and at margin with few hairs. Inflorescence

with central rachis rarely developed (up to 0.5 cm long); branches 2 or 3, 3—5 cm long, 0.5—

0.8 mm wide, glabrous, minutely serrate. Pedicel 0.5—1.5 mm long, smooth. Spikelets in

threes, homomorphous, |.3—1.6 mm long, 0.5 mm wide, glabrous; the two lowest bracts

0.9—0.95 times as long as spikelet; lower glume absent (rarely up to 0.1 mm long); upper

glume |.1—1.5 mm long, 3—5 veins, acute; 1** lemma |.1—1.5 mm long, in- or equidistantly

7-veined, acute; 2™ lemma protruding (at maturity), 1.5 mm long, acute, pale yellow to yellow-

brown with bluish to purplish tip in fruit.

Native; Paleotropics. First record 1890. Rare.

Pioneer on damp sandy to rocky soil in disturbed places, on beaches, roadsides, etc.

Digitaria longiflora (Retz.) Pers. — Fig. 45.

Lesser Crab Grass

Ridley 1925: 214, p.p. Gilliland 1971: 192. Veldkamp 1973: 66. Keng et al. 1998:

162.

D. pertenuis Buse: Ridley 1925: 214, p.p. Gilliland 1971: 192, p.p.

Paspalum longiflorum Retz.: Ridley 1907: 126, p.p.

Culms tufted and with long stolons, erect part 10—50O cm long; nodes glabrous or hairy.

Sheath 0.7—4.5 cm long, sparsely to densely hairy. Ligule 1—2 mm long. Blade 1—5 cm

long, 2—S mm wide, glabrous or hairy, base rounded. Inflorescence with central rachis rarely

developed (up to 0.2 cm long); branches 2—4, 2—5 cm long, 0.5 mm wide, glabrous, serrate.

Pedicel up to 1.5 mm long, smooth. Spikelets in threes, homomorphous, |.3—1.7(—1.9) mm

long, 0.5—0.6 mm wide; either lowest bract c. 0.1 and 2°‘ 0.9—1 times as long as spikelet, or

the two lowest bracts 0.9—1 times as long as spikelet; lower glume absent or up to 0.15

56 Supplement of Gardens’ Bulletin Singapore 57 (2005)

mm long; upper glume 1.3—1.7 mm long, hairy at least at margins, 5 veins, acute; 1** lemma

1.3—1.9 mm long, hairy, more or less inequidistantly 5—7-veined, obtuse to acute; 2° lemma

1.5—1.8 mm long, acute, (pale) yellow to yellow-brown in fruit.

Native; Paleotropics. First record 1894. Rare.

Pioneer on moist sandy to rocky soil, in open grasslands, forest margins, on roadsides.

A collection earlier than that mentioned in Duistermaat (2004: 33) was found among

material at SING that was included in D. violascens (see note there).

It closely resembles D. mollicoma, which generally has longer spikelets and acute to

acuminate 1* lemmas.

Digitaria mollicoma (Kunth) Henrard — Fig. 46.

Veldkamp 1973: 69. Duistermaat 2004: 33.

D. pertenuis Buse: Ridley 1925: 214, p.p. Gilliland 1971: 192, p.p.

Paspalum longiflorum auct., non Retz.: Ridley 1907: 126, p.p. — D. longiflora auct., non

Pers.: Ridley 1925: 214, p.p.

Culms with stolons, 45—50 cm long; nodes (sparsely) hairy. Sheath 1.5—8 cm long, (sparsely)

hairy. Ligule 1—2 mm long, erose. Blade (1—)2—6 cm long, 2—5.5 mm wide, glabrous to

hairy, base rounded. Inflorescence with central rachis rarely developed (up to 1.5 cm long);

branches 2—4, 4—7 cm long, 1—1.1 mm wide, glabrous, serrate. Pedicel 0.5—2.5 mm long,

smooth or sparsely serrate. Spikelets in threes or fours, homomorphous, 1.8—2 mm long

(excluding hairs), 0.6 —0.7 mm wide; lowest bract c. 0.9 times as long as and 2™ bract as long

as spikelet; lower glume absent; upper glume 1.6—1.7 mm long (excluding hairs), with 3

veins, hairy between veins and at margin, overtopping at tip; 1“ lemma 1.7—1.8 mm long,

more or less equidistantly 7-veined, hairy, acute to acuminate; 2" lemma 1.8—1.9 mm long,

acute, (pale) yellow to dark yellow-brown in fruit.

Native; Penisular Malaysia to New Guinea, and Taiwan. First record 1890. Rather

rare.

On open damp, sandy to rocky soil, (shaded) roadsides.

Earlier collections than that mentioned in Duistermaat (2004) were found among

material at SING that was included in D. violascens (see note there).

Digitaria nuda Schumach. — Fig. 47.

Veldkamp 1973: 41. Tan 1995: 139.

Culms tufted, erect to decumbent, 30—60 cm long; nodes glabrous to sparsely hairy. Sheath

3—8 cm long, glabrous to hairy. Ligule 1—1.5 mm long. Blade 1.5—7 cm long, 2.5—5 mm

wide, glabrous, base cuneate to rounded and with few hairs. Inflorescence with central rachis

rarely developed, up to 1 cm long; branches 2—4(—8), 5—8 cm long, 0.4—0.5 mm wide,

glabrous or sparsely hairy with hairs up to 5 mm long, serrate, rarely with setae in lower part.

Pedicel 0.5—2.5 mm long, serrate. Spikelets in pairs, homomorphous, 2—2.5 mm long, 0.5—

0.7 mm wide; either lowest bract less than 0.1 and 2"? 0.5—0.6 times as long as spikelet, or

lowest bract 0.5—0.6 times as long as and 2" as long as spikelet; lower glume absent or

Figure 45. Digitaria longiflora (Retz.) Pers.

a. facing upper glume, b. facing 1“ lemma.

Figure 46. Digitaria mollicoma (Kunth) Henrard.

a. facing 1* lemma, b. facing upper glume.

Figure 47. Digitaria nuda Schumach.

a. facing 1* lemma, b. facing upper glume.

Figure 48. Digitaria radicosa (J.Presl) Miq.

a. facing lower glume, b. facing upper glume.

Field Guide to the Grasses of Singapore (Excluding the Bamboos)

eget

Fig. 48

GRILLES

itt 42 PTE BE EEA GES AR

Fig. 47

58 Supplement of Gardens’ Bulletin Singapore 57 (2005)

rarely up to 0.15 mm long; upper glume 1.2—1.4 mm long (excluding overtopping hairs),

hairy, 3 veins; 1‘ lemma 2.2—2.5 mm long, (almost) equidistantly 7-veined, sparsely hairy,

acute; 2" lemma 2—2.4 mm long, acute, yellowish to brownish greyish in fruit.

Native; tropics. First record 1955. Rare.

Waste places, preferring sandy soil.

It is not mentioned in Keng et al. (1998), but is included in Tan (1995: 139).

Differences with D. ciliaris are mentioned there.

Digitaria radicosa (J.Presl) Miq. — Fig. 48, Plate 8.

Veldkamp 1971: 35. Keng et al. 1998: 163.

D. marginata Link var. debilis auct., non Ridl.: Hook.f. ex Ridl. 1925: 214, p.p.

D. timorensis (Kunth) Balansa: Gilliland 1971: 198.

Paspalum sanguinale Lam. var. debile auct., non Hook.f.: Ridley 1907: 125, p.p.

Culms with (short) stolons, erect part 20—60 cm long; nodes glabrous to sparsely short-

hairy. Sheath 2—6 cm long, glabrous or hairy. Ligule 1—2 mm long. Blade 3—5 cm long,

2—5 mm wide, glabrous or hairy, base (slightly) rounded and hairy. Inflorescence without

central rachis; branches 2 or 3 (or 4), 3—7 cm long, 0.5—0.6 mm wide, glabrous, smooth (at

10x magnification). Pedicel 0.5—3 mm long, more or less smooth. Spikelets in pairs,

homomorphous, 2.5—3.5 mm long, 0.5—0.7 mm wide; either lowest bract 0.03—0.1 and 2"

0.3—0.5 times as long as spikelet, or lowest 0.3—0.5 times as long as and 2" as long as

spikelet; lower glume absent or 0.1—0.4 mm long; upper glume |.2—1.6 mm long, 3 veins,

hairy at least at margin, obtuse or bifid; 1 lemma 2.7—3.5 mm long, 3—S veins (if 5 these

inequidistant), hairy especially at margin, acute; 2"’ lemma 2.4—2.7 mm long, acute, yellowish

in fruit.

Native; Paleotropics. First record 1892. Rather common.

In open places, lawns, on roadsides.

Digitaria setigera Roem. & Schult.

Veldkamp 1973: 37. Keng et al. 1998: 163.

D. marginata Link var. commutata auct., non Ridl. and var. pruriens (Fisch. ex Trin.) Hook.f.

ex Ridl. 1925: 214.

D. microbachne (J.Presl) Henrard: Gilliland 1971: 194.

D. microstachya Henrard: Gilliland 1971: 195.

Paspalum sanguinale Lam. var. extensum Hook.f. and var. pruriens (Fisch. ex Trin.) Hook.f.

and var. commutatum auct., non Hook.f. p.p.: Ridley 1907: 125.

Culms tufted and erect to decumbent or with stolons, (20—)40—100 cm long. Sheath 4—10

cm long. Ligule 1.5—3.5 mm long. Blade 3—17 cm long, 4—9 mm wide, glabrous, base

cuneate to rounded and hairy. Inflorescence branches serrate. Pedicel 0.5—3 mm long. Spikelets

in pairs, homomorphous; upper glume hairy at least at margin, obtuse or bifid; 1** lemma

inequidistantly 5- or 7-veined, acute; 2™ lemma acute, yellowish to brownish in fruit.

This is the most robust of the Digitaria species.

Figure 49. Digitaria setigera Roem. & Schult. var. calliblepharata (Henrard) Veldk.

a. facing 1“ lemma, b. facing upper glume.

Figure 50. Digitaria setigera Roem. & Schult. var. setigera.

a. facing 1“ lemma, b. facing upper glume.

Figure 51. Digitaria violascens Link.

a. facing lower glume, b. facing upper glume.

Figure 52. Dimeria ornithopoda Trin.

Field Guide to the Grasses of Singapore (Excluding the Bamboos)

——— [a

60 Supplement of Gardens’ Bulletin Singapore 57 (2005)

la. Spikelets 3—3.2 mm long; 1* lemma with 1.5 mm-long bristles, otherwise

glabrous; 2" lemma 2.8—3 mm long. D. setigera var. calliblepharata

lb. Spikelets 2.8—3.1 mm long; 1*t lemma hairy, bristles absent; 2"* lemma

2.5—2.8 mm long. D. setigera var. setigera

Digitaria setigera Roem. & Schult. var. calliblepharata (Henrard) Veldk. — Fig. 49.

Veldkamp 1973: 40. Keng et al. 1998: 163.

D. marginata Link var. ciliaris auct., non Ridl.: Ridley 1925: 214, p.p. — Paspalum sanguinale

Lam. var. ciliare auct., non Hook.f.: Ridley 1907: 125.

D. microbachne subsp. calliblepharata Henrard: Gilliland 1971: 194.

Nodes glabrous. Sheath glabrous to sparsely hairy. Inflorescence with central rachis 1.5—2

cm long; branches 5—8, 8—10 cm long, 0.5—0.7 mm wide. Spikelets 3—3.2 mm long,

0.7—0.8 mm wide (excluding bristles); lowest bract 0.15—0.2 and 2" as long as spikelet;

lower glume absent; upper glume 0.5—0.7 mm long (excluding hairs); 1** lemma 3—-3.2 mm

long, inequidistantly 7-veined, with up to 1.5 mm long bristles in 2™ intervenium, otherwise

glabrous; 2"? lemma 2.8—3 mm long.

Native; tropical Asia. First record 1904. Extinct (1959).

Open places.

Digitaria setigera Roem. & Schult. var. setigera — Fig. 50.

Nodes glabrous to sparsely hairy. Sheath at least lowest (sparsely) hairy with up to 4 mm-long

patent hairs. Inflorescence with central rachis O—4 cm long; branches (3—)5—11, 6—16 cm

long, 0.6—0.7 mm wide, glabrous. Spikelets 2.8—3.1 mm long, 0.7—0.9 mm wide; either

lowest bract 0.03 and 2"? 0.2—0.25 times as long as spikelet, or lowest bract 0.2—0.25 times

as long as and 2™ as long as spikelet; lower glume absent to 0.2 mm long; upper glume 0.6—

0.7 mm long (excluding hairs); 1‘ lemma 2.8—3.1 mm long, sparsely hairy, inequidistantly

7-veined; 2™ lemma 2.5—2.8 mm long.

Native; tropical Asia. First record late 19" century. Common.

Weed in waste places.

Digitaria violascens Link — Fig. 51.

Gilliland 1971: 191. Veldkamp 1973: 63. Keng et al. 1998: 63.

D. caespitosa Ridl. 1925: 215.

D. chinensis (Nees) A.Camus, non Hornem.: Ridley 1925: 215.

D. pertenuis Buse: Gilliland 1971: 192 (excluding Colour pl.17).

Paspalum longiflorum auct., non Retz.: Ridley 1907: 126, p.p. — D. longiflora auct., non

Pers.: Ridley 1925: 214, p.p.

Culms tufted, erect to ascending, stolons absent, 25—-70 cm long; nodes glabrous or sparsely

hairy. Sheath 2—14 cm long, hairy at least in lower half. Ligule 1—2 mm long. Blade (1.5—)5—

13 cm long, (1.5—)2—6 mm wide, glabrous or hairy, base rounded. Inflorescence with central

rachis 0.3—2 cm long; branches 2—6, 4—11 cm long, 0.6—0.8 mm wide, glabrous, serrate.

Pedicel 0.5—3.5 mm long, sparsely to densely serrate. Spikelets in threes (fours or fives),

homomorphous, |.3—1.9 mm long, 0.5—0.7 mm wide; either lowest bract 0.03 and 2™ 0.9—

| times as long as spikelet, or lowest bract 0.9—1 times as long as and 2™ as long as spikelet;

lower glume absent or 0.1 mm long; upper glume |1.2—1.9 mm long, hairy, 3 veins, truncate

to acute; 1 lemma 1.3—1.9 mm long, (sparsely) hairy, inequidistantly 7-veined (or midvein

lacking), obtuse to acute; 2" lemma 1.2—1.7 mm long, acute, dark brown to black in fruit.

Native; tropics. First record 1889. Common.

Weed in waste places, on roadsides etc.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 61

Contrary to Veldkamp (1973), plants with stolons are excluded from D. violascens.

These stoloniferous plants differ from D. violascens in having glabrous to sparsely serrate

pedicels and (dark) yellow-brown 2™ lemmas when fruiting. Part of the stoloniferous material

is included in D. longiflora and part in D. mollicoma.

Dimeria R.Br.

Dimeria ornithopoda Trin. — Fig. 52.

Bird’s Foot Grass

Ridley 1907: 151; 1925: 192. Keng et al. 1998: 163. — var. ornithopoda: Gilliland

, 197%: 214.

Dimeria glabra Ridl. 1925: 192. — D. ornithopoda Trin. var. glabra (Ridl.) Jansen: Gilliland

1971: 216. Keng et al. 1998: 163.

Culms tufted, erect to decumbent, stolons absent, 12—80 cm long; nodes hairy. Sheath 0.7—

2(—4) cm long, rounded, midrib distinctly raised, glabrous or (sparsely) hairy, margins

glabrous. Ligule membranous, 0.4—0.7 mm long, ciliolate. Young leaf blade inrolled. Blade

1—7 cm long, 1—2 mm wide, midvein at upperside prominent and white and at underside

raised, glabrous or hairy, base rounded and glabrous. Inflorescence panicle, central rachis

absent; branches 2 or 3, 2.5—6 cm long, 0.8 mm wide, glabrous, triquetrous (to narrowly

winged), naked at lower 1.2—3 mm, simple. Pedicel 0.1 mm long. Spikelets single, alternate,

2.3—2.8 mm long, 0.5 mm wide, laterally flattened, 2-flowered, awned or unawned; callus

with few hairs; lowest bract 0.8 times as long as and 2" as long as spikelet; lower glume 1.9—

2.5 mm long, herbaceous, | vein, sparsely hairy at least on vein, acute; upper glume 2.2—2.7

mm long, herbaceous, 3 veins, keeled and sparsely hairy on midvein, truncate with minutely

protruding midvein; 1‘ lemma 0.6—0.9 mm long, hyaline; 1‘ palea absent; 2° lemma 1.8—

2 mm long, hyaline, bifid, unawned or awn arising just below tip and 5.5—8.5 mm long; 2"

palea absent.

Native; tropical Asia and Australia. First record 1889. Rare.

In (partly) shaded and (periodically) wet grasslands, marshes; usually on poor soil,

sometimes ephemeral.

The infraspecific taxa are not maintained because they seem to be the extremes of a

range.

Echinochloa P.Beauv.

Sheath rounded, glabrous. Young leaf blade inrolled. Blade glabrous to sparsely hairy at upper

side, base rounded (to cuneate in E. picta). Inflorescence panicle, central rachis present;

branches simple or branched, setulose and/or hairy. Spikelets single, not to slightly

dorsiventrally flattened, 2-flowered; lowest bract about half as long as spikelet, 2" (excluding

awn) as long as spikelet; glumes and 1“ lemma (thin) herbaceous; lower glume enveloping;

I palea hyaline; 2"' lemma and palea indurated, glabrous (to sparsely hairy at tip in E. picta).

la. Ligule row of hairs, 1.5—2.5 mm long. E. picta

1b. Ligule absent. | 2

2a. Inflorescence branches all simple or rarely with the lowermost with an up

to 0.8 cm-long secondary branch. Spikelets more or less clearly arranged

in four rows. 1“ lemma acuminate to mucronate, never awned. Anthers

purple or yellow. Stigmas blackish-purple. E. colona

62 Supplement of Gardens’ Bulletin Singapore 57 (2005)

2b. Inflorescence branches with the lower ones usually shortly branched at

base. Spikelets usually not clearly arranged in four rows. 1* lemma

acuminate, usually at least a few awned. Anthers yellow. Stigmas white

or red. E. crus-galli

Echinochloa colona (L.) Link — Fig. 53, Plate 9.

Jungle Rice, Padi Burung

Ridley 1925: 222. Gilliland 1971: 167. Keng et al. 1998: 164.

Panicum colonum L.: Ridley 1907: 132.

Culms single or tufted, erect to ascending, 25—100 cm long; nodes glabrous to sparsely

hairy. Sheath 3—6 cm long. Ligule absent. Blade 6—28 cm long, 4—8 mm wide, glabrous to

sparsely hairy. Inflorescence 6—22 cm long; branches 6—30, 0.6—6.5 cm long, 0.4 mm

wide. Pedicel up to 0.7 mm long, sparsely hairy. Spikelets more or less arranged in 4 rows,

2.3—3.7 mm long, 0.8 mm wide, unawned; lower glume 0.9—1.6 mm long, hairy in marginal

area, 3 veins, 0.2 mm-long mucronate; upper glume 2.2—3.5 mm long, hairy on veins and

margin, 5 veins, 0.3 mm-long mucronate; 1“ lemma 2.1—3.2 mm long, hairy at least on

veins, 5 veins, acuminate; 1‘ palea 1.5—2.5 mm long; 2™ lemma 1.8—2.9 mm long, acute.

Native; Paleotropics. First record 1880s. Rather common.

In waste places, often periodically wet places.

Echinochloa crus-galli (L.) P.Beauv. — Fig. 54.

Barnyard Millet, Padi Burung

Ridley 1925: 222. Gilliland 1971: 168. Keng et al. 1998: 164.

Panicum crus-galli L.: Ridley 1907: 132.

Culms erect to ascending, 50—100 cm long; nodes glabrous. Sheath c. 15 cm long. Ligule

absent. Blade 25 cm long, 7 mm wide, glabrous. Inflorescence 14—24 cm long; branches

11—18, longest 3—8 cm long, 0.5 mm wide. Pedicel up to 0.5 mm long, sparsely hairy.

Spikelets not arranged in 4 rows, 2.8—3.1 mm long, 1.3 mm wide, mucronate to awned;

lower glume 1—1.2 mm long, hairy, 3 veins, 0.2 mm-long mucronate; upper glume 2.6—3

mm long, veins and marginal area hairy, 5 veins, 0.3—0.4 mm-long mucronate; 1* lemma 2

mm long, hairy on veins, 5 veins, awn 0.5—4(—20) mm long with the longest at tip of

branch(let); 1** palea 1.4—1.5 mm long; 2"* lemma 2—2.1 mm long, acute to acuminate.

Native; Paleo(-sub)tropics, serious weed in temperate regions. First record 1932. Rare.

In waterlogged waste places, roadsides.

This species is also known as Cockspur.

Echinochloa picta (J.K6nig) P.W.Michael — Fig. 55.

Clayton 1989: 57.

E. stagnina auct., non (Retz.) P.Beauv.: Bor 1960: 311; Gilliland 1971: 166; Keng et al..1998:

164.

Culms rhizomatous, creeping or floating, up to 200 cm long; nodes unknown. Ligule 1.5—

2.5 mm-long hairs. Blade 26 cm long, 12 mm wide, glabrous, base cuneate to rounded.

Figure 53. Echinochloa colona (L.) Link.

a. facing upper glume, b. facing lower glume.

Figure 54. Echinochloa crus-galli (L.) P.Beauv.

a. facing upper glume, b. facing lower glume.

Figure 55. Echinochloa picta (J.K6nig) P.W.Michael.

a. facing upper glume, b. facing lower glume.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 63

64 Supplement of Gardens’ Bulletin Singapore 57 (2005)

Inflorescence 18 cm long; branches 8, longest c. 3 cm long, 0.7 mm wide. Pedicel up to 1.5

mm long, setulose. Spikelets not arranged in 4 rows, 4.7 mm long, 2.1 mm wide, awned;

lower glume 2.1 mm long, sparsely hairy, 5 veins, acuminate to mucronulate; upper glume

4.7 mm long, sparsely hairy, at base with 7—9 veins, acute to obtuse; 1“ lemma 4.5 mm long,

sparsely hairy, 7 veins, awn from tip and 3.5 mm long; 1“ palea 3.4 mm long; 2™ lemma 4 mm

long, acuminate.

Native; India and SE Asia. Single record 1883, locality unknown. Extinct (1883).

Streamsides, pond margins, growing in water, sometimes floating.

It was formerly known as E. stagnina, which is an African species with less plump

spikelets (Clayton 1989).

The culm nodes are not represented on the Singapore Herbarium material. Gilliland

(1971) mentioned that the nodes are ‘with a few scattered short hairs’, whereas Veldkamp

(pers. comm.) has seen some glabrous nodes as well.

Eleusine Gaertn.

Culms tufted, erect to geniculate, stolons absent; nodes glabrous. Sheath glabrous, margin

hairy at least at throat. Young leaf blade folded along midrib. Blade underside glabrous,

upperside sparsely hairy, base rounded. Inflorescence panicle (rarely spike); branches at least

half the number at tip, simple. Pedicel absent. Spikelets single, alternate on abaxial side of

branch, laterally flattened, 3—6-flowered, unawned; the two lowest bracts 0.45—0.65 times

as long as spikelet; glumes and lemmas herbaceous, with keeled midvein, hairy; lemmas

obtuse; paleas hyaline, 2 keeled veins, hairy; 2" and following florets as 1“, gradually smaller.

la. Leaves overtopping the inflorescence. Lower glume 2.4 mm long, truncate;

upper glume 3 mm long, obtuse. E. coracana

1b. Leaves shorter than to overtopping the inflorescence. Lower glume 1.8—

2 mm long, obtuse to acute; upper glume 2.4—2.6 mm long, acute.

E. indica

Eleusine coracana (L.) Gaertn. — Fig. 56.

African Millet

Gilliland 1971: 80. Keng et al. 1998: 164.

E. indica (L.) Gaertn. var. coracana Gaertn.: Ridley 1907: 174; 1925: 250.

Culms c. 60 cm long. Leaves overtopping inflorescence. Sheath 10—14 cm long, rounded.

Ligule either membranous and ciliolate, or row of hairs, 2 mm long. Blade 53 cm long, 13

mm wide, base at margin glabrous. Inflorescence central rachis rarely developed (up to 1 cm

long); branches 7—9, 6—8 cm long, 2.5 mm wide, margins hairy. Spikelet 5 mm long, 3.8

mm wide, 4-flowered; lower glume 2.4 mm long, 3 veins, truncate; upper glume 3 mm long,

5 veins, obtuse; 1“ lemma 3 mm long, 7 veins.

Not native; Africa, cultivated in tropical regions. First record 1880s. Cultivated (last

record 1963).

Eleusine indica (L.) Gaertn. — Fig. 57.

Goose Grass, Rumput Sambau

Ridley 1907: 174; 1925: 250. Gilliland 1971: 78. Keng et al: 1998: 165.

Culms 25—45 cm long. Leaves shorter than to overtopping the inflorescence. Sheath 5—10

cm long, more or less flattened, midrib raised. Ligule membranous, 0.5—1 mm long, glabrous

or with few hairs at tip. Blade 4.5—16(—40) cm long, 3—4.5 mm wide, base at margin

sparsely hairy. Inflorescence central rachis O—7 cm long; branches 1—11, 5—15 cm long, |

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 65

mm wide, only at axils hairy. Spikelet 3.8—6 mm long, 1.7—4 mm wide, 3—6-flowered;

lower glume 1.8—2 mm long, | or 3 veins, obtuse to acute; upper glume 2.4—2.5 mm long,

5—7 veins, acute; | lemma 2.7—3 mm long, 3 veins.

Native; worldwide in (sub-)tropical regions. First record 1880s. Common.

Weed of waste places, often after disturbance, on moist (not wet) soil.

Eragrostis Wolf

Lazarides 1997: 77—187. Veldkamp 2002: 157—204.

Culms single or tufted, erect to decumbent; nodes glabrous. Sheath rounded, external ligule

short-hairy. Young leaf blade inrolled. Inflorescence panicle, central rachis present; branches

0.3—0.5 mm wide, at least longest branched. Spikelets single, laterally flattened, 3—many-

flowered, unawned; the two lowest bracts 0.05—0.5 times as long as spikelet; palea 2 keeled

veins; 2™ and following florets as 1‘, but gradually smaller.

Eragrostis elongata (Willd.) J.Jacq., actually the correct name for the Papuasian E.

diandra (R.Br.) Steud. (Veldkamp 2002), has been so widely misapplied to species of the E.

brownii complex, e.g. E. atrovirens and E. brownii (in the wider sense), that Veldkamp (2002:

160) has regarded it as a confusing name to be disregarded.

The inclusion in Keng et al. (1998: 166) of Eragrostis nutans (Retz.) Steud., described

as ‘palea falling together with the lemma’, is probably a misidentification; the name refers to

an Indian species with persistent paleas (Veldkamp 2002: 175).

la. Palea keels setose. E. amabilis

1b. Palea keels glabrous to ciliolate. 2

2a. Paleas falling off early (Fig. 66b). 3

2b. Paleas longer persistent (Fig. 63). 6

3a. Culm (60—)80—160 cm long. Spikelet rachilla with 7 or 8 nodes per 3

mm length. Anthers 0.6—1 mm long. Fruit fusiform, terete, fruit wall

finely striate. E. atrovirens

3b. Culm 10—55(—80) cm long,—if longer than 55 cm then spikelet rachilla

with 13—16 nodes per 3 mm length. Anthers 0.1—0.5 mm long. Fruit

ellipsoid, laterally somewhat flattened, fruit wall smooth or finely

reticulate. 4

4a. Perennials. Culms rooting at lower nodes. Ligule a ciliolate rim. Spikelet

rachilla with 13—16 nodes per 3 mm length. Keels of palea ciliolate.

E. unioloides

4b. Annuals. Culms not rooting at lower nodes. Ligule a ciliolate rim or a row

of hairs. Spikelet rachilla with 7—10 nodes per 3 mm length. Keels of

palea scaberulous to ciliolate. 5

Sa. Ligule a ciliolate rim. Glumes subequal, lower glume 1-veined. Anthers 2.

Fruit wall finely reticulate. E. gangetica

5b. Ligule a row of hairs. Glumes unequal, lower glume without veins. Anthers

3. Fruit wall smooth. E. pilosa

66 Supplement of Gardens’ Bulletin Singapore 57 (2005)

6a. (2) Sheaths and blades glandular. E. cilianensis

6b. Sheaths and blades without glands. 7

7a. Rachilla articulated, ultimately breaking up from above downward (Fig.

58a). E. brownti

7b. Rachilla persistent, not articulated (Fig. 58b). E. montana

Eragrostis amabilis (L.) Wight & Arn. ex Nees — Fig. 59, Plate 10.

Feathery Eragrostis, Rumput Telur Kutu

Ridley 1907: 178, p.p.; 1925: 246, p.p. Keng et al. 1998: 165. Veldkamp 2002: 164.

E. tenella (L.) P.Beauv. ex Roem. & Schult.: Gilliland 1971: 71.

E. viscosa (Retz.) Trin.: Ridley 1907: 177; 1925: 246. Gilliland 1971: 71. Keng et al. 1998:

167.

Culms geniculate to prostrate, not rooting at the lower nodes, 15—40 cm long, green. Sheath

1—3.2 cm long, glabrous to sparsely hairy, margin glabrous or throat with up to 4 mm long

hairs. Ligule 0.3—0.4 mm long hairs, free or fused at basal 0.1—0.2 mm. Blade 1—9 cm

long, c. 2.5 mm wide, glabrous, base rounded and with 2—3 mm long hairs. Inflorescence

3—16 cm long; longest branches 1.5—3 cm long, scaberulous, with tuft of hairs in axils.

Pedicel 1—3 mm long, scaberulous. Spikelet 1.5—1.9 mm long, 0.9 mm wide, hyaline, 3—

6-flowered, palea and rachilla disarticulating from top to bottom; lower glume 0.4—0.7 mm

long, glabrous, 1 vein; upper glume 0.6—0.8 mm long, as lower; 1* lemma 0.7—0.9 mm

long, glabrous, 3 veins, truncate; 1‘ palea 0.7—0.9 mm long, glabrous, keels with 0.3—

0.4(—0.6) mm long setae. Anthers 3, 0.2 mm long.

Native; Paleotropics. First record 1893. Common.

Weed on roadsides, in waste places, etc.

Glandular and sticky plants (in the literature known as E. viscosa) have not been found

in Singapore.

Differences with Sporobolus tenuissimus are mentioned there.

Eragrostis atrovirens (Desf.) Trin. ex Steud. — Fig. 60.

Gilliland 1971: 68. Lazarides 1997: 97. Keng et al. 1998: 165. Veldkamp 2002: 167.

E. elegantula auct., non Steud.: Ridley 1907: 179; 1925: 248, p.p.

E. elongata auct., non (Willd.) J.Jacq.: Ridley 1907: 179; 1925: 248. Keng et al. 1998: 166,

p.p.

Culms erect or geniculate, sometimes rooting at the lower nodes, (60—)80—160 cm long,

glaucous. Sheath 7—12 cm, glabrous. Ligule 0.2—0.3 mm long, ciliolate. Blade 1O—17 cm

long, 2—5 mm wide, glabrous or upperside sparsely long-hairy, base cuneate. Inflorescence

13—25 cm long; longest branches 8—20 cm long, smooth, glabrous. Pedicel 1—4 mm long,

scaberulous. Spikelet 3.5—20 mm long, 1.3—2.2 mm wide, herbaceous, 4—38-flowered;

rachilla with 7 or 8 nodes per 3 mm, persistent; lemmas and paleas disarticulating

simultaneously from bottom to top; lower glume 1—1.5 mm long, glabrous, 1 vein; upper

glume 1.5—2 mm long, as lower; 1*' lemma 1.7—2 mm long, punctate, 3 veins, acute; 1*

Figure 56. Eleusine coracana (L.) Gaertn.

Figure 57. Eleusine indica (L.) Gaertn.

Figure 58. Eragrostis rachilla.

a. articulated, b. not articulated.

Figure 59. Eragrostis amabilis (L.) Wight & Arn. ex Nees.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 67

68 Supplement of Gardens’ Bulletin Singapore 57 (2005)

palea 1.5—1.8 mm long, glabrous, keels scaberulous to ciliolate. Anthers 3, 0.6—0.7 mm

long.

Native; Paleotropics. First record 1906. Rather common.

On moist to wet soil, usually unshaded.

Its tall size and bluish colour are very distinctive.

Eragrostis brownti (Kunth) Nees — Fig. 61, Plate 11.

Lazarides 1997: 101. Veldkamp 2002: 169. Duistermaat 2004: 34.

Eragrostis cumingii Steud.: Gilliland 1971: 65. Lazarides 1997: 109. — var. cumingii: Veldkamp

Ae: Dt,

E. elegantula auct., non Steud.: Ridley 1907: 179; 1925: 248, p.p.

E. elongata auct., non (Willd.) J.Jacq.: Ridley 1907: 179; 1925: 248. Keng et al. 1998: 166,

p.p.

Culms geniculate, not rooting at the lower nodes, 18—50 cm long, green to glaucous. Sheath

1.5—6 cm long, at throat with up to 4 mm-long hairs. Ligule 0.2—0.3 mm long, ciliolate.

Blade 4—8(—24) cm long, 1—3 mm wide, glabrous, base rounded, with few up to 4 mm-

long hairs. Inflorescence 5.5—18 cm long; longest branches 1.5—-5 cm long, scaberulous to

setulose, with few hairs in axils. Pedicel 0.5—1.7 mm long, scaberulous to setulose. Spikelet

2.5—20 mm long, |.2—2.2 mm wide, (thin) herbaceous, (3—)4—50-flowered; rachilla with

9—11 nodes per 3 mm, articulated, (ultimately) breaking up from above, lemmas disarticulating

from below and paleas more or less persistent; lower glume 1.1—1.5 mm long, glabrous, 1

vein; upper glume 1.5—1.7 mm long, as lower; 1 lemma 1.5—1.7 mm long, glabrous or

somewhat punctate, 3 veins, along marginal veins minute glands (25x magnification!), obtuse

to acute; 1“ palea 1.2—1.6 mm long, glabrous, keels scaberulous. Anthers 3, 0.15—0.4 mm

long.

Native; Tropical Asia and Australia. First record 1889. Common.

Open to slightly shaded places, roadsides, lawns, often on sandy soil.

Eragrostis brownii is treated in the wider sense, including E. cumingii var. cumingii

(Duistermaat 2004).

Eragrostis cilianensis (Bellardi) Vignolo ex Janch. — Fig. 62.

Lazarides 1997: 105. Veldkamp 2002: 171. Duistermaat 2004: 34.

Culms erect or geniculate, sometimes rooting at the lower nodes, 55 cm long. Sheath 3—5 cm

long, with translucent glands on veins in upper half, collar in marginal area with up to 2.5

mm-long hairs. Ligule row of hairs, 0.5—0.7 mm long. Young leaf blade unknown. Blade

12—14 cm long, 3.5—4.5 mm wide, glabrous, base cuneate to rounded, margin in basal half

with sessile glands. Inflorescence 12—15 cm long; longest branch 3—6 cm long, glabrous or

with few scattered glands. Pedicel 1.5—2 mm long, scaberulous. Spikelet 5—9.5 mm long,

2.1—2.7 mm wide, herbaceous, 9—20-flowered; rachilla with 8 or 9 nodes per 3 mm, persistent,

lemmas disarticulating from below and paleas long persistent; lower glume 1.5 mm long,

glabrous, 1 vein; upper glume 2 mm long, glabrous, 3 veins; 1‘ lemma 2 mm long, glabrous,

marginal and uppermost area minutely punctate, 3 veins, midvein protruding; 1* palea 1.7

mm long, glabrous, keels ciliolate. Anthers 3, 0.25—0.35 mm long.

Probably not native; Paleo(sub)tropics, especially in regions with a distinct dry season.

Single record 1941 from Yio Chu Kang. Extinct (1941).

Figure 60. Eragrostis atrovirens (Desf.) Trin. ex Steud.

Figure 61. Eragrostis brownii (Kunth) Nees.

Figure 62. Eragrostis cilianensis (Bellardi) Vignolo ex Janch.

Figure 63. Eragrostis gangetica (Roxb.) Steud.

Figure 64. Eragrostis montana Balansa.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 69

Fig. 62

70 Supplement of Gardens’ Bulletin Singapore 57 (2005)

Eragrostis gangetica (Roxb.) Steud. — Fig. 63.

Gilliland 1971: 68. Keng et al. 1998: 166. Veldkamp 2002: 175.

Culms erect, not rooting at the lower nodes, c. 40 cm long. Sheath 3—6 cm long, margin at tip

with a few hairs. Ligule 0.1—0.2 mm long, ciliolate. Blade 6—14 cm long, 2—4 mm wide,

underside glabrous, upperside with up to 1.5 mm long hairs, base cuneate. Inflorescence 14—

15 cm long; longest branches c. 5 cm long, glabrous. Pedicel 2—5 mm long, scaberulous.

Spikelet 4.5—5.5 mm long, 1.5 mm wide, herbaceous, 9—13-flowered; rachilla with 9 or 10

nodes per 3 mm, persistent, lemmas and paleas disarticulating from below upward; lower

glume unknown; upper glume | mm long, glabrous, | vein; 1“ lemma 1.5 mm long, glabrous,

smooth or minutely punctate, 3 veins, obtuse; |* palea 1.1 mm long, glabrous, keels scaberulous.

Anthers 2, 0.3 mm long.

Probably not native; tropical Africa to N Vietnam. Single record 1951 from Seletar

Reservoir. Extinct (1951).

In disturbed places.

According to Veldkamp (2002), the lemmas are acuminate.

Eragrostis montana Balansa — Fig. 64.

Keng et al. 1998: 166. Veldkamp 2002: 180.

E. malayana Stapf: Ridley 1907: 178; 1925: 247. Gilliland 1971: 65.

Culms erect or geniculate, not rooting at the lower nodes, 20—35 cm long. Sheath 1.8—3.2

cm long, glabrous. Ligule row of hairs, 0.2—0.3 mm long. Blade 4—6 cm long, 0.5—1.3 mm

wide, underside glabrous, upperside sparsely hairy with hairs up to 2 mm long, base rounded.

Inflorescence 5—10 cm long; longest branch 1—2.5 cm long, scaberulous, axils glabrous or

with | long hair. Pedicel 1—3 mm long, glabrous. Spikelet 2—4 mm long, 1.2—2.1 mm

wide, (thin) herbaceous, 3—18-flowered; rachilla with 13 nodes per 3 mm, persistent, lemmas

disarticulating from below and paleas long persistent; lower glume 0.6—0.7 mm long, glabrous,

1 vein; upper glume 0.9—1 mm long, as lower; 1*“' lemma |1.1—1.2 mm long, glabrous, 3

veins, acute; 1* palea 1 mm long, glabrous, keels scaberulous. Anthers 3, 0.2 mm long.

Native; Myanmar to western Malesia. First record 1889. Extinct (1963).

On water-logged, stony or sandy soil-in open forest, along roads, ditches, banks.

Eragrostis pilosa (L.) P.Beauv. — Fig. 65.

Indian Love Grass

Ridley 1907: 179; 1925: 247. Gilliland 1971: 69. Keng et al. 1998: 167. Lazarides

1997: 142. Veldkamp 2002: 183.

Culms erect or geniculate, not rooting at the lower nodes, 14—45 cm long. Sheath 3.5—9 cm

long, glabrous, sometimes glandular, throat at margin with up to 3 mm long caducous hairs.

Ligule row of hairs, 0.3—0.4 mm long. Blade 10—13 cm long, 2 mm wide, glabrous or

sometimes glandular, base rounded, with a few hairs. Inflorescence 9—17 cm long; longest

branch 3—8 cm long, glabrous, lowest axils with up to 3 mm long hairs (or rarely glabrous).

Pedicel 2—5 mm long, glabrous. Spikelet 3.3—5.5 mm long, 0.8—1.2 mm wide, hyaline to

thin herbaceous, 4—13-flowered; rachilla with 7—9 nodes per 3 mm, persistent, lemmas

disarticulating from below and paleas slightly longer persistent; lower glume 0.5 mm long,

glabrous, unveined; upper glume | mm long, glabrous, | vein; 1** lemma 1—1.5 mm long,

with scattered minute glands, otherwise glabrous, 3 veins, obtuse; 1* palea 0.9—1.1 mm

long, glabrous, keels scaberulous. Anthers 3, 0.2 mm long.

Native; temperate to tropical regions of the Old World. First ord 1904. Rather rare.

Drought-resistant weed, in open places like roadsides, flowerbeds etc.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) #1

Eragrostis unioloides (Retz.) Nees ex Steud. — Fig. 66.

Pink Eragrostis, Rumput Kolam Padang

Gilliland 1971: 66. Keng et al. 1998: 167. Lazarides 1997: 163. Veldkamp 2002: 187.

Eragrostis amabilis auct., non Wight & Arn. ex Nees: Ridley 1907: 178, p.p.; 1925: 246, p.p.

Culms geniculate to decumbent, rooting at the lower nodes, 12—50(—80) cm long. Sheath

2—4 cm long, glabrous, throat at margin with a tuft of up to 2 mm long hairs. Ligule 0.2 mm

long, ciliolate. Blade 1.5—14 cm long, 2—7 mm wide, glabrous or with long hairs on upperside

only, base rounded, collar with up to 3 mm long hairs. Inflorescence 5—17 cm long; longest

branch 1.5—6 cm long, glabrous. Pedicel 2—9 mm long, scaberulous. Spikelet 3.5—7.8 mm

long, 2.5—3.2 mm wide, herbaceous, 7—34-flowered (less in juvenile spikelet), whitish and

often tinged pinkish; rachilla with 13—16 nodes per 3 mm, persistent, lemmas and paleas

disarticulating simultaneously from below; lower glume I1—1.2 mm long, | vein (not in

middle); upper glume 1.5—1.7 mm long, glabrous, | vein; 1“ lemma 1.4—1.5 mm long,

glabrous, punctate, 3 veins, acute; |“ palea 1.2 mm long, glabrous, keels ciliolate. Anthers 2,

0.2—0.4 mm long.

Native; SE Asia. First record 1888. Common.

Open, unshaded to shaded places on moist to waterlogged soil, e.g. roadsides, fields,

swamps etc.

Eriachne R.Br.

Van Eck-Borsboom 1980: 127—138.

Culms tufted, erect; nodes glabrous. Sheath rounded. Ligule row of hairs. Young leaf blade

setaceous. Blade c. 0.5 mm wide, setaceous. Inflorescence panicle, central rachis present;

branches 0.2 mm wide, with few spikelets. Pedicel scaberulous. Spikelets single, more or less

laterally flattened, 2-flowered, florets articulating, awned; the two lowest bracts as long as

spikelet (excluding awns); glumes herbaceous; callus below each floret hairy; lemmas and

paleas hairy, 1* identical to 2".

According to van Eck-Borsboom (1980), most Malaysian and Singapore specimens

are cleistogamous.

la. Each spikelet with 2 awns, points of the palea not extended into awns.

E. pallescens

1b. Each spikelet with 6 awns, points of the palea extended into awns.

| E. triseta

Eriachne pallescens R.Br. — Fig. 67.

Slender Eriachne

Ridley 1907: 172; 1925: 240. Gilliland 1971: 94. Van Eck-Borsboom 1980: 133. Keng

etal, 1998: 167.

Culms 20—60 cm long. Sheath 2—3.5 cm long, glabrous or margin sparsely hairy. Ligule 0.5

mm long. Blade 2—6 cm long, hairy, or underside glabrous, base somewhat rounded, collar

at margin with few up to 3 mm long hairs. Inflorescence 2—7 cm long, branches 2—4; longest

3—5 cm long, scaberulous, with 2—4 spikelets, simple or branched. Pedicel 5—25 mm long.

Spikelet 4—4.4 mm long, |.1—2 mm wide, with 2 awns; lower glume 3.9—4.2 mm long, not

enveloping base, glabrous or with few hairs on midvein, 9 veins; upper glume as lower, 3.5—

3.9 mm long, 9—11 veins; 1‘ lemma 3—3.2 mm long, herbaceous, 5 veins, awn from tip and

3.5—3.7 mm long; 1* palea 3—3.4 mm long, bifid, unawned.

Native; tropical Australasia. First record 1889. Rather rare.

Dry, rocky or clayey soil, unshaded.

72 Supplement of Gardens’ Bulletin Singapore 57 (2005)

Eriachne triseta Nees ex Steud. — Fig. 68.

Ridley 1907: 172 1925: 240 (both excluding Singapore). Van Eck-Borsboom 1980:

136. Keng et al. 1998: 168.

Massia triseta Balansa in Morot: Gilliland 1971: 94.

Culms c. 40 cm long. Sheath 2—4(—6.5: flag) cm long, sparsely short-hairy. Ligule 0.4 mm

long. Blade 7—11 cm long, sparsely hairy at both sides, base cuneate. Inflorescence 5—9 cm

long; branches 2, c. 1.5 cm long, axil with up to 1 mm long hairs, with 2 spikelets, simple.

Pedicel 2—8 mm long. Spikelet 9.8 mm long, 1.8 mm wide, with 6 awns; lower glume 9 mm

long, almost entirely enveloping base, glabrous, 11 veins; upper glume as lower; 1“ lemma

18.3 mm long including the not clearly demarcated awn, indurated, 5 veins; 1** palea 17 mm

long including the two awns, indurated, 2 veins graduating into awns.

Probably not native; tropical Australasia. Single record 1983 near the National Parks

Pasir Panjang Nursery. Rare.

Partly shaded, open waste ground.

Eriochloa Kunth

Davidse 1987: 143—155. Thompson et al. 1990: 1463—1468.

Stolons absent. Nodes hairy. Sheath rounded. Ligule row of hairs. Young leaf blade inrolled.

Blade at underside sparsely hairy, base rounded. Inflorescence panicle, central rachis present,

branches simple or branched. Spikelets in pairs (sometimes obscurely so) with pedicels of

unequal length, not to slightly dorsiventrally flattened, 2-flowered, unawned or if awned awn

not exserted; callus cup-shaped, 0.2—0.7 mm long, often red-coloured; the two lowest bracts

as long as spikelet; lower glume absent; upper glume and 1* lemma herbaceous, with 5 veins,

transverse veinlets very obscure or absent; 2™ lemma and palea indurated, glabrous.

Thompson et al. (1990) showed that the callus is not a rudimentary lower glume.

Davidse (1987) suggested it might contain lipids to attract ants, but further research is needed

to verify this.

Eriochloa villosa (Thunb. in Murray) Kunth, reported by Keng et al. (1998), is excluded

because information on the herbarium labels of d’ Alleizette on which this record was based is

unreliable (Veldkamp pers. comm.).

la. Pedicel scaberulous. Upper glume glabrous or sparsely hairy at base and

margin, obtuse to acute; 1‘ palea 2.5—2.8 mm long; 2"? lemma 0.1 mm

apiculate. E. meyerana

1b. Pedicel scaberulous or with a few up to 2 mm-long hairs at tip. Upper

glume (sparsely) hairy, 0.1—0.3 mm-long mucronate; |“ palea absent;

2™ lemma with 0.4—0.6 mm-long awn, awn not exserted. E. procera

Figure 65. Eragrostis pilosa (L.) P.Beauv.

Figure 66. Eragrostis unioloides (Retz.) Nees ex Steud.

a. young spikelet, b. maturing spikelet.

Figure 67. Eriachne pallescens R.Br.

Figure 68. Eriachne triseta Nees ex Steud.

Figure 69. Eustachys tenera (J.Presl) C.E.Hubb.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 73

74 Supplement of Gardens’ Bulletin Singapore 57 (2005)

Eriochloa meyerana (Nees) Pilg. — Fig. 70.

Keng et al. 1998: 168. Veldkamp 1999b: 233.

Culms single, erect, c. 55 cm long. Sheath 5—12 cm long, glabrous to sparsely hairy at base,

margin in basal half with up to 1 mm-long hairs. Ligule 0.7—0.8 mm long. Blade 4—11 cm

long, 4—5.5 mm wide. Inflorescence 11—14 cm long, branches 5—9; longest branch 6—8

cm long, 0.6 mm wide, scaberulous. Longer pedicel of the pair 1.5—-2 mm long, scaberulous.

Spikelet 3.2—3.6 mm long, 1.1—1.3 mm wide; upper glume 2.8—2.9 mm long, glabrous or

sparsely hairy at base and marginal area, obtuse to acute; 1* lemma 2.7—2.8 mm long, sparsely

hairy, acute; 1“ palea 2.5—2.8 mm long; 2" lemma 2—2.3 mm long, smooth, 0.1 mm-long

apiculate.

Not native; (sub-)tropical Africa, introduced elsewhere. First record 1948 in Kew

Herbarium (not seen). Extinct (1966).

On bank of canal, University campus (the university campus adjacent to the Botanic

Gardens and the Bukit Timah Road).

Eriochloa procera (Retz.) C.E.Hubb. — Fig. 71, Plate 12.

Gilliland 1971: 202. Keng et al. 1998: 168.

E. annulata (Fliggé) Kunth: Ridley 1925: 223 (excluding Singapore).

E. polystachya auct., non Humb., Bonpl. & Kunth: Ridley 1907: 127 (excluding Singapore).

Culms single or tufted, erect or outer culms ascending, 50—120 cm long. Sheath 6—14 cm

long, glabrous. Ligule 0.7 mm long. Blade 5—20 cm long, 4.5—10 mm wide. Inflorescence

17—18 cm long, branches 9—14; longest 3.5—7 cm long, 0.4 mm wide, short hairy especially

in axil. Longer pedicel of the pair 2—3 mm long, at tip with up to 2 mm long hairs or rarely

scaberulous. Spikelet 3—3.5 mm long, 1 mm wide; upper glume 2.7—3.1 mm long, (sparsely)

hairy, 0.1—0.3 mm-long mucronate; 1‘ lemma 2.5—2.9 mm long, hairy; 1* palea absent to

0.5 mm long; 2"’ lemma |.8—2 mm long, rugulose, awn from tip 0.4—0.6 mm long and not

exserted.

Native; Paleotropics. First record 1928. Rather common.

Wet to moist waste places in coastal areas and more inland.

The cup-shaped callus distinguishes this species from Panicum repens, Urochloa

subquadripara and U. piligera. Also, P. repens has a lowest bract that is c. 0.2 times as long

as spikelet, whereas both Urochloa species have glumes and 1“ lemmas with transverse veins.

Eustachys Desf.

Eustachys tenera (J.Presl) C.E.Hubb. — Fig. 69.

Gilliland 1971: 88. Keng et al. 1998: 168.

Cynodon dactylon auct., non (L.) Pers.: Ridley 1907: 173, p.p.; 1925: 249, p.p.

Culms tufted, geniculate and with short stolons, 20—40 cm long; nodes glabrous. Sheath 4—

5 cm long, glabrous, broadly winged. Ligule 0.1 mm-long hairs. Young leaf blade folded

along midrib. Blade up to 9 cm long, 2—3.5 mm wide, glabrous, base cuneate. Inflorescence

panicle, central rachis absent; branches 2—4, longest 4—5 cm long, 0.4 mm wide, scaberulous,

Figure 70. Eriochloa meyerana (Nees) Pilg.

a. facing 1“ lemma, b. lateral view, c. pair of pedicels.

Figure 71. Eriochloa procera (Retz.) C.E.Hubb.

a. facing 1* lemma, b. pedicel.

Figure 72. Heteropogon contortus (L.) P.Beauv.

a. male spikelet, b. female spikelet.

Figure 73. Hymenachne amplexicaulis (Rudge) Nees.

Field Guide to the Grasses of Singapore (Excluding the Bamboos)

76 Supplement of Gardens’ Bulletin Singapore 57 (2005)

simple. Pedicel absent. Spikelets single, alternate on one side of branch, c. 1.2 mm long, 1.3

mm wide, somewhat laterally flattened, 2-flowered, mucronate; lowest bract as long as and

2™ (excluding mucro) 0.9 times as long as spikelet; lower glume | mm long, hyaline to thin

herbaceous, glabrous, | vein; upper glume 1.1 mm long, herbaceous, scaberulous, | vein, 0.5

mm mucronate; 1* lemma 1.1 mm long, herbaceous, 3 veins with 0.5 mm long hairs; 1* palea

0.9 mm long; 2"? lemma 0.6 mm long, hyaline, glabrous; 2™ palea absent.

Native; Indochina to Malesia. First record 1898. Extinct (1964).

Sandy soil.

It has been confused with Cynodon dactylon, but the latter has unwinged sheaths,

longer ligules, hairy blades and acute upper glumes that are not mucronate. Chloris barbata

differs in generally being a taller plant, and in having inrolled young leaf blades and distinctly

awned and 3-flowered spikelets.

Heteropogon Pers.

Heteropogon contortus (L.) P.Beauv. — Fig. 72.

Tangle-head

Ridley 1925: 213 (excluding Singapore). Gilliland 1971: 292. Keng et al. 1998: 169.

Andropogon contortus L.: Ridley 1907: 167 (excluding Singapore).

Culms tufted, erect, c. 60 cm long; nodes glabrous. Sheath 4—7 cm long, rounded, with up to

1.5 mm-long bulbous-based hairs, margin hairy. Ligule 1 mm long, basal part membranous

and 0.5 mm long, hairs at tip 0.5 mm long. Young leaf blade folded along midrib. Blade 10—

20 cm long, 2—7 mm wide, sparsely hairy to glabrescent, base cuneate and hairy. Inflorescence

spike-like raceme, 4—5 cm long. Pedicel 0.9 mm long, glabrous. Spikelets in pairs of | sessile

and | pedicelled, 2-flowered, unisexual, first 8 or 9 pairs male and unawned spikelets followed

by 5—8 pairs of 1 sessile female (or rarely bisexual) awned and 1 pedicelled male unawned

spikelet; the two lowest bracts as long as spikelet; both paleas absent. Male spikelet 6—7 mm

long, | mm wide; lower glume 5.8 mm long, herbaceous, almost glabrous at base, upwards

with hairs increasing in length up to 1.5 mm long, 13 veins; upper glume 6—7 mm long,

herbaceous, sparsely hairy on midvein and margins only, 3 veins; 1“ lemma 4.5 mm long,

hyaline; 2™ lemma as 1*, 4mm long. Female spikelet 5 mm long, falling off with 2.5 mm long

sharply pointed and short-hairy part of rachis; lower glume 5 mm long, slightly indurated,

with 0.2 mm long stiff hairs, 9 veins; upper glume as lower, 5 mm long, 3 veins; 1“ lemma 2.2

mm long, hyaline; 2 lemma 4.5 mm long, basal part very narrow and hyaline, upward widening

and somewhat indurated, awn from tip and 8.5 cm long.

Native; tropical America, Africa, India to S. China, Japan, Marianas Isl., Australia,

widespread in Malesia. First record 1936. Extinct (1957).

On open sandy soil, beach.

The long awns tend to become entangled, hence the name Tangle-head.

Hymenachne P.Beauv.

Pohl and Lersten 1975: 223—227.

Hymenachne and Sacciolepis are separated by the presence of culms filled with spongy

pith in the former, and gibbous upper glumes in the latter (Pohl and Lersten 1975).

Hymenachne amplexicaulis (Rudge) Nees — Fig. 73.

Swamp Panic Grass, Rumput Kumpai

Keng et al. 1998: 170.

H. acutigluma (Steud.) Gilliland: Gilliland 1971: 155.

H. myuros auct., non Humb., Bonpl. & Kunth: Ridley 1925: 230.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) yi |

Panicum myurus auct., non Humb., Bonpl. & Kunth: Ridley 1907: 135 (excluding Singapore).

Culms erect to ascending, stolons absent, c. 90 cm long; internodes c. 3 mm diameter, (partially)

filled with open-spongy pith; nodes glabrous. Sheath 8—11 cm long, rounded, with (few)

transverse veins present (only visible from inside), glabrous, margin hairy. Ligule membranous,

2 mm long. Young leaf blade inrolled. Blade 10—22 cm long, 6—11 mm wide, underside

glabrous, upperside sparsely hairy, base rounded to somewhat cordate. Inflorescence spike-

like panicle, central rachis 10—23 cm long, branches erect; longest branch up to 6 cm long,

0.6 mm wide, branched, 1* branchlets very short and flattened with hairlike spicules on margin.

Pedicel 0.2—0.5 mm long, with hairlike spicules. Spikelets single, (3—)3.9—4.9 mm long,

0.6 mm wide, not flattened, 2-flowered, unawned; lowest bract c. 0.25 and 2" 0.7—0.9 times

as long as spikelet; glumes herbaceous, midvein more or less keeled and scaberulous and up

to 0.4 mm protruding; lower glume 1—1.3 mm long, enveloping, glabrous, 1—3 veins; upper

glume 2.5—3.2 mm long, glabrous, 5 veins; 1** lemma 3.4—4.4 mm long, (thin) herbaceous,

glabrous, 5 veins, (long-)acuminate; 1‘ palea absent; 2°? lemma 2.4—2.7 mm long, hyaline;

2™ palea 2.4 mm long.

Native; India, Myanmar, Thailand, Peninsular Malaysia, Singapore, Borneo. First record

1904. Rare.

In open swamps and riversides, on muddy soil.

Also called Wick Grass, as the pith of the culm was used as a wick for oil lamps.

The length of the spikelets is more variable than mentioned by Gilliland (1971: 4.6

mm long).

Differences with Panicum auritum are mentioned there. Sacciolepis indica has gibbous

spikelets of up to 3.4 mm long, shorter panicle branches (up to 0.5 cm long), culms without

pith and sheaths without transverse veins.

Imperata Cirillo

Ng 1999: 288.

Culms single or tufted, with short rhizomes, erect, internodes of non-flowering plants very

short. Leaves of non-flowering plants all basal. Sheath rounded. Ligule membranous, dorsal

side hairy. Young leaf blade inrolled. Blade base cuneate, those of flag leaves 0.6—0.8 cm

long. Inflorescence panicle, silky hairy, central rachis present; branches numerous, scaberulous,

persistent, branched. Spikelets in pairs with pedicels of unequal length, not flattened, 2-

flowered, unawned; callus densely silky-hairy, hairs 6—10 mm long; the two lowest bracts as

long as spikelet; both glumes thin herbaceous, hairy; both lemmas and 2” palea hyaline; 1*

palea absent. :

Some Miscanthus species (not recorded from Singapore, but used as ornamentals),

Neyraudia, Phragmites and Saccharum also have silky-hairy inflorescences. However, all

these have non-flowering culms with much longer internodes and with all leaves scattered

along the culm. Also, in Saccharum species the inflorescence branches break up at maturity,

whereas the spikelets of Miscanthus species are distinctly awned. The spikelets of both

Neyraudia and Phragmites have 4 or more florets.

Recent (as yet unpublished) research has indicated that the two taxa generally accepted

cannot be distinguished. The anthers often seem to have 2 locules at the base and 4 at the tip

(study by Ms Delia Co; Veldkamp pers. comm.). For now the two species are maintained.

la. Flowering culm glabrous just below the nodes. Peduncle hollow. Panicle

28—52 cm long, more or less lax; longest branch 7.5—15 cm long,

drooping, without spikelets in lower 0.3—0.4 part. Anther 1.

I. conferta

78 Supplement of Gardens’ Bulletin Singapore 57 (2005)

1b. Flowering culm with long bulbous-based hairs just below the nodes.

Peduncle more or less solid. Panicle 4—20 cm long, more or less

contracted; longest branch 1.2—4 cm long, erect, with spikelets from

the base. Anthers 2. I. cylindrica var. major

Imperata conferta (J.Presl) Ohwi — Fig. 74.

Gilliland 1971: 222. Keng et al. 1998: 170.

I. contracta Hitche.: Burkill 1935: 1228.

I. exaltata auct., non Brongn.: Ridley 1907: 152; 1925: 193.

Culms 50—150 cm long; flowering culm glabrous just below the nodes. Sheath glabrous.

Ligule 0.7—2.5 mm long. Blade 65—110 cm long, 10—22 mm wide, glabrous. Peduncle

hollow. Inflorescence 28—52 cm long, 5—8 cm wide; longest branch 7.5—15 cm long,

drooping, naked in lower 0.3—0.4 part. Pedicel short 1.3—1.5 mm, long 2—3.5 mm long,

scaberulous. Spikelet 3—3.2 mm long; lower glume 3—3.5 mm long, hairs 3.5—-5 mm long,

3 veins; upper glume as lower, 2.8—3.2 mm long; 1“ lemma 2—2.1 mm long; 2™ lemma 1.9

mm long; 2" palea 1 mm long. Anther 1, 1.6—1.8 mm long.

Native; SE Asia. First record 1889. Rare.

Open sandy places, unshaded to partly shaded dry laterite soil, on roadsides.

The only recent location where /. conferta and I. cylindrica grew together yielded an

intermediate form with characters of both species. Most likely it is a hybrid between the two

species.

Imperata cylindrica (L.) P.Beauv. — Fig. 75.

Lalang, Lalang

Clayton and Renvoize 1982: 702.

I. cylindrica var. maior (Nees) C.E.Hubb. ex C.E.Hubb. & Vaughan: Gilliland 1971: 220.

Keng et al. 1998: 170.

I. arundinacea Cirillo: Ridley, 1907: 152; 1925: 193.

Culms 65—75 cm long; flowering culm with bulbous-based hairs just below the nodes. Sheath

glabrous or hairy. Ligule (0.5—)2 mm long, tip ciliolate. Blade 40—60 cm long, 6—14 mm

wide, base with up to 3 mm long hairs. Inflorescence 4—20 cm long, 0.6—2 cm wide; longest

branch 1.2—4 cm long, erect, with spikelets from the base. Pedicel short 1.3 mm, long 2.5

mm long, at base with up to 3 mm long hairs. Spikelet 3—4.5 mm long; lower glume 2.7—4.5

mm long, hairs 8—10 mm long, | vein; upper glume as lower, 2.7—4.2 mm long, 3 or 5

veins; 1 lemma 2.4—3 mm long; 2™ lemma 1.4—1.9 mm long; 2™ palea 1.1—1.2 mm long.

Anthers 2, 2—3 mm long.

Native; tropical Asia. First record 1880s. Common.

Unshaded to slightly shaded fields, roadsides etc.

Clayton and Renvoize (1982) argued that all varieties should be ignored. Although

there is geographical variation, the variants cannot be separated perfectly.

Lalang is one of the world’s worst weeds. Fields that have been burnt repeatedly are

dominated by it to such an extent that all other species are outcompeted. The rhizomes of this

fire climax species survive fire and immediately start new growth whereas other species are

often killed. When frequently mown, however, plants get smaller and produce smaller

inflorescences or stop flowering altogether. Eventually, this process kills them. For

intermediates, see note under J. conferta.

Figure 74. Imperata conferta (J.Presl) Ohwi.

Figure 75. Imperata cylindrica (L.) P.Beauv.

Field Guide to the Grasses of Singapore (Excluding the Bamboos)

RE inremenanionee re

rooernnemeeninaneny sty ear oe nih At ANAC Rn

ee WON oe

Fig. 74

80 Supplement of Gardens’ Bulletin Singapore 57 (2005)

Isachne R.Br.

Iskandar and Veldkamp 2004: 159—179.

Culms with stolons or tufted and geniculate. Sheath rounded, midrib not raised, glabrous,

margins hairy. Ligule row of hairs. Young leaf blade inrolled. Blade sparsely hairy at both

sides, at base at underside with prominent veins, base rounded. Inflorescence panicle, central

rachis present; branches glabrous, axils of the lowest with a gland, at least longest branched.

Pedicels present. Spikelets single, not flattened, 2(—3)-flowered, unawned; glumes (thin)

herbaceous; both paleas slightly shorter than lemmas, glabrous, texture as lemma.

Spikelets with 3 florets are very rare and in Singapore are known only from one

collection (see /. pulchella). The hairy ligule and the relatively long glumes distinguish the

species of Jsachne readily from Cyrtococcum species.

The key follows Iskandar and Veldkamp (2004).

la. Glumes longer than florets. Lower floret bisexual. Both lemmas hairy,

similarly indurated and convex. I. kunthiana

1b. Glumes shorter than or as long as florets. Lower floret male. Both lemmas

glabrous to hairy, the 1* lemma thinner and usually larger and flatter

than the 2". Z

2a. Culm with annular gland below the node. I. pulchella

2b. Culm without annular gland below the node. 3

3a. Nodes glabrous. Ligule (1.4—)2—4 mm long. Blade underside at base

7—9-veined, margins white cartilaginous or not. 2"? lemma indurated.

I. globosa

3b. Nodes (sparsely) hairy. Ligule 1—1.6 mm long. Blade underside at base

3—5-veined, margins not white cartilaginous. 2"’ lemma herbaceous.

I. minutula

Isachne globosa (Thunb.) Kuntze — Fig. 76.

Rounded Isachne, Rumput Minyak

Gilliland 1971: 123. Keng et al. 1998: 170. Iskandar and Veldkamp 2004: 165.

f, australis R.Br,: Ridley 1907; 129-1925: 239°

Culms erect part 30—60 cm long; nodes glabrous, annular ring below them absent. Sheath

1.8—6.5 cm long, marginal hairs 2—3 mm long. Ligule 2—4 mm long. Blade 4—7.8 cm

long, 3.5—7 mm wide, prominent veins 7—9; margin white cartilaginous or not, scaberulous,

glabrous or hairy. Inflorescence 5—14 cm long, longest branch 2.5—6 cm long. Pedicel 0.5—

1.7 mm long, glabrous. Spikelet 1.8—2.5 mm long, 1.2—1.4 mm wide, not yawning; the two

lowest bracts 0.9—1 times as long as spikelet, not longer than the florets; lower glume 1.7—

2.4 mm long, sparsely hairy, 7 veins; upper glume 1.6—2.2 mm long, sparsely hairy, 9 veins;

1 lemma 1.8—2.2 mm long, glabrous, slightly indurated, dorsally convex or grooved; 2"

lemma 1.2—1.6 mm long, glabrous, indurated.

Native; India to China, south to Malesia, Solomon Islands, New Caledonia, Australia,

New Zealand. First record 1889. Rare.

In wet and open places.

Figure 76. Isachne globosa (Thunb.) Kuntze.

Figure 77. Isachne kunthiana (Wight & Arn. ex Steud.) Mig.

Figure 78. Isachne minutula (Gaudich.) Kunth.

Figure 79. Isachne pulchella Roth in Roem. & Schult.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 81

82 Supplement of Gardens’ Bulletin Singapore 57 (2005)

Isachne kunthiana (Wight & Arn. ex Steud.) Mig. — Fig. 77.

Ridley 1907: 128. Gilliland 1971: 122. Keng et al. 1998: 171.

I. semitalis Rid]. 1925: 237.

Culms erect part 7—30 cm long; nodes short appressed hairy, annular ring below them absent.

Sheath 1.2—2 cm long, marginal hairs up to 2 mm long. Ligule 1—1.6 mm long. Blade 2—

5 cm long, 6.5—9 mm wide, prominent veins 5—7; margin white cartilaginous, scaberulous,

glabrous. Inflorescence 2—4 cm long, longest branch 2 cm long. Pedicel 1.2—1.5 mm long,

glabrous or with a single hair. Spikelet 2—3.3 mm long, yawning at maturity; the two lowest

bracts as long as spikelet, longer than the florets; lower glume 2.2—3 mm long, hairy, 7 or 9

veins; upper glume 1.9—2.9 mm long, hairy, 9 or 11 veins; 1* lemma 1.5—1.8 mm long,

hairy, indurated, convex; 2"? lemma as 1*.

Native; India to Japan, China, Malesia southward to Solomon Islands, New Caledonia,

Australia, New Zealand. First record 1892. Rare.

Damp open forest sites.

Isachne minutula (Gaudich.) Kunth — Fig. 78.

Iskandar and Veldkamp 2004: 167.

I. miliacea auct., non Roth: Ridley 1907: 129; 1925: 239.

I. pulchella auct., non Roth: Gilliland 1971: 120. Keng et al. 1998: 171.

Culms erect part c. 10 cm long; nodes (sparsely) hairy at least below the margins of the

sheaths, annular ring below them absent. Sheath 0.6—2.2 cm long, marginal hairs up to 1 mm

long. Ligule 1—1.6 mm long. Blade 1.6—3.3 cm long, 2.5—6 mm wide, prominent veins

3—5; margins not white cartilaginous, scaberulous, glabrous. Inflorescence 24.5 cm long,

longest branch 0.6—2 cm long. Pedicel 0.5—1.3 mm long, glabrous. Spikelet 1.4—1.8 mm

long, 1.2 mm wide, not yawning; the two lowest bracts 0.9—1 times as long as spikelet, not

longer than the florets; lower glume 1.3—1.7 mm long, sparsely hairy, 7 veins; upper glume

as lower, 1.2—1.7 mm long; 1* lemma 1.2—1.7 mm long, glabrous, thin herbaceous, dorsally

grooved; 2° lemma 0.8—1.2 mm long, densely covered with 0.15 mm long, patent hairs,

slightly indurated.

Native; India to Vietnam, Sumatra, Java, Borneo, Philippines, Celebes, Lesser Sunda

Islands, Moluccas, Australia. First record 1892. Extinct in the wild (1905).

Damp places in forest.

Recently only found in the Botanic Gardens at several different places where it seems

to grow abundantly as a weed in damp to wet places.

This species was previously known as /. pulchella (Gilliland 1971, Keng et al. 1998),

but that species has an annular gland below each node.

Isachne pulchella Roth in Roem. & Schult. — Fig. 79.

Iskandar and Veldkamp 2004: 168.

Culm erect part 20—35 cm long; nodes hairy to glabrescent, annular ring below them present.

Sheath 1—2.5 cm long, marginal hairs up to 0.5 mm long. Ligule c. 1 mm long. Blade 1—2.5

cm long, 3—5 mm wide, prominent veins 7; margins white cartilaginous, scaberulous, at base

hairy. Inflorescence 3—4.5 cm long, longest branch 0.8—1.5 cm long. Pedicel 0.3—0.7 mm

long, glabrous. Spikelet c. 1.4 mm long, c. 1 mm wide, yawning at maturity; the two lowest

bracts 0.8—0.9 times as long as spikelet, not longer than the florets; lower glume 1.3 mm

long, glabrous, 5 veins; upper glume as lower, 1.1—1.2 mm long; 1* lemma 1.2 mm long,

glabrous to sparsely short-hairy at tip, thin herbaceous, flattened, not dorsally grooved; 2”

lemma | mm long, densely short-hairy, indurated.

Native; India to SW China, Sumatra, Java, Borneo, Philippines, Celebes. First record

i905. Rare.

On clayey soil in a ditch with slow flowing clear water.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 83

This is the first time this species is recorded for Singapore (see the note under

I. minutula). The first record is from Bukit Timah, the second from the Western Catchment

area where it was found in 2004. Both the remoteness of the recent locality and the habitat

indicate that it is native. Some of the spikelets were 3-flowered with an additional floret

above the 2".

Ischaemum L.

Auricles triangular, fused with ligule, glabrous. Ligule membranous, tip entire or ciliolate.

Young leaf blade inrolled. Inflorescence panicle, central rachis absent (or up to 0.8 cm long);

branches 2(—4), one with spikelets from the base upward, the other with the basal 0.6 cm

naked, margins hairy (or glabrous in J. magnum), simple. Spikelets in pairs of 1 sessile and 1

pedicelled on one side of rachis, heteromorphous, not flattened, 2-flowered; callus with thin

part that is concave after flowering; the two lowest bracts as long as spikelet; lemmas and

paleas thinner than glumes; both paleas slightly shorter to slightly longer than the lemmas.

Sessile spikelet with lower glume flattened, basal 0.5—0.7 part indurated, upper 0.3—0.5

part herbaceous and submarginally keeled to winged; upper glume boat-shaped, midvein more

or less keeled. Pedicelled spikelet (slightly) smaller than sessile; lower glume boat-shaped,

smooth (or minutely rugulose), winged at one margin; 2" lemma unawned (or awn shorter

than in sessile spikelet).

Inflorescences with 3 or 4 branches occur rarely in /. ciliare, I. muticum and I. timorense.

Material of Ischaemum barbatum Retz., earlier mentioned as a weed in the Botanic

Gardens (Ridley 1925, Keng et al. 1998), has not been re-found. It differs from J. magnum in

having a geniculate awn.

The key follows Gilliland (1971).

la. Lower glume of sessile spikelet entire at the tip; upper glume unawned. 2

1b. Lower glume of sessile spikelet bifid at the tip; upper glume awned, awn

0.5—2 mm long. “|

2a. Lower glume of sessile spikelet not transversely rugose or nodular. Pedicel

and joint of rachis subequal to rachis below. I. muticum

2b. Lower glume of sessile spikelet transversely rugose or nodular at the

margins. Pedicel and joint of rachis inflated compared to rachis below.

3a. Lower glume with up to 4 transverse rugosities or marginal nodules.

Perennial. I. magnum

3b. Lower glume with about 6 transverse rugosities. Short-lived ‘annual’.

I. rugosum

4a. Lower glume obovate, wings 0.4—1 mm wide. ~ I. ciliare

4b. Lower glume ovate-lanceolate, wings 0—0.4 mm wide. I. timorense

Ischaemum ciliare Retz. — Fig. 80, Plate 13.

Smut Grass

Ridley 1907: 160. Veldkamp 1991: 180. Keng et al. 1998: 171.

I. aristatum auct., non L.: Ridley 1925: 203.

I. indicum auct., non (Houtt.) Merr.: Gilliland 1971: 263.

84 Supplement of Gardens’ Bulletin Singapore 57 (2005)

Culms with stolons or straggling, 20—90 cm long; nodes hairy. Sheath 2—10 cm long, rounded

at base and more or less flattened in upper part, with prominent midrib, at least the lower

hairy, margins glabrous. Ligule 1—2 mm long, hairy at abaxial side, tip ciliolate. Blade (2—)

4—7(—11) cm long, 4—8 mm wide, both sides hairy, base cuneate, pseudopetiole absent.

Peduncles 1 per sheath and from upper sheath. Inflorescence branches 3—12 cm long,

divergent. Sessile spikelet (4—)4.5—5.5(—7.5) mm long, 1.5—2 mm wide, awned; lower

glume 3.4—5.5 mm long, obovate, basal part smooth and glabrous, upper part 7—13 veins

and sparsely hairy, wings 0.4—1 mm wide, tip bifid; upper glume (4—)5.5—6 mm long, 5

veins, margins hairy, awn 0.9—1.5 mm long; 1*t lemma 3.8—4.5 mm long; 2"? lemma 3—4

mm long, upper half bifid, awned from notch, awn 6—16 mm long, geniculate. Pedicel c. 2.5

mm long, as thick as rachis, margins hairy.

Native; India to Japan, south to Malesia, Australia. First record 1890. Common.

Lawns, roadsides, open places.

Plants infected by smut fungus occur frequently, hence the English name. Their

inflorescences do not develop completely and often do not completely exert from their sheaths.

It is very similar to J. timorense with which it might be conspecific. In addition to

characters already mentioned in the key, /. ciliare is generally (but not always) a smaller plant

with shorter leaf blades.

Ischaemum magnum Rendle — Fig. 81, Plate 14.

Ridley 1907: 158; 1925: 200. Gilliland 1971: 261. Keng et al. 1998: 171.

I. laeve Ridl.: Ridley 1907: 158; 1925: 201.

Culms single or tufted, erect or geniculate, 110—200 cm long; nodes glabrous. Sheath 10—

17 cm long, rounded, glabrous or hairy, margin glabrous or hairy. Ligule 2—7.5 mm long,

glabrous or hairy at abaxial side, tip entire. Blade 1O—35 cm long, 1O—16 mm wide, glabrous,

base long cuneate, pseudopetiole 1—4 mm long and hairy. Peduncles 1—3 per sheath and

from upper and 1 or more lower sheaths. Inflorescence branches 6.5—14 cm long, closely

appressed or divergent. Sessile spikelet 5S—6.5 mm long, 1.5—1.8 mm wide, (minutely) awned;

lower glume 5—6.5 mm long, oblong, glabrous, basal part with 4 (or 5) transverse rugosities

or (rudimentary) marginal nodules, upper part 0 or 1 vein, wings O—0.4 mm wide, tip entire

(and mucro up to 0.1 mm long); upper glume 5—6.3 mm long, glabrous or margins hairy, 3—

5 veins, unawned; 1‘ lemma 4.5—5.5 mm long; 2"! lemma 2.5—5.1 mm long, upper half

bifid, awned from notch, awn 0.4—12 mm long, straight. Pedicel 1.5 mm long, as thick as

rachis, margins glabrous or one hairy.

Native; Myanmar to Borneo. First record 1878. Rather rare.

Margin of rivers, canals and lakes; in places with seepage; on beach.

Ischaemum muticum L. — Fig. 82, Plate 15.

Seashore Centipede Grass, Rumput Tembaga Jantan

Ridley 1907: 159; 1925: 201. Gilliland 1971: 255. Keng et al. 1998: 172.

Culms with stolons or straggling, 30—200 cm long; nodes glabrous. Sheath 2.5—4 cm long,

more or less flattened, midrib raised, glabrous or hairy at tip, at least one margin hairy. Ligule

0.5 mm long, glabrous, tip entire or ciliolate. Blade 3—12 cm long, 5—13 mm wide, glabrous

or underside sparsely hairy, base rounded, pseudopetiole 1—2 mm long. Peduncles | per

sheath and from upper sheath. Inflorescence branches 1.5—5 cm long, closely appressed.

Sessile spikelet 6.7—7.5 mm long, 2.2—2.3 mm wide, minutely awned, awn not protruding;

Figure 80. Ischaemum ciliare Retz., pair of spikelets.

Figure 81. Ischaemum magnum Rendle, pair of spikelets.

Figure 82. Ischaemum muticum L., pair of spikelets.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 85

sa Fig. 82

86 Supplement of Gardens’ Bulletin Singapore 57 (2005)

lower glume 6.7—7.5 mm long, oblong, glabrous, basal part smooth, upper part with 7—9

veins and transverse veins, wings c. 0.4 mm wide, tip entire; upper glume 6.7—7.5 mm long,

3 veins, glabrous, unawned; 1* lemma 5.5—6.5 mm long; 2"* lemma 4.5—6.6 mm long, tip

acute, awn 1.2—1.5 mm long, straight. Pedicel 5.3 mm long, as thick as rachis, margins hairy.

Native; India to E. Asia, south to Malesia and Polynesia. First record 1888. Common.

In lawns, open places, scrub, also near the sea (mangroves and beaches above high

tide mark).

It is readily distinguished from the other species of Ischaemum by the pale yellow

inflorescence with white stigmas sticking out laterally.

Ischaemum rugosum Salisb. — Fig. 83.

Wrinkled Centipede Grass, Rumput Ekor Cawi

Ridley 1907: 157; 1925: 200. Gilliland 1971: 259. Keng et al. 1998: 172.

Culms erect, c. 50 cm long; nodes hairy. Sheath 8—10 cm long, rounded, sparsely hairy at tip,

margins glabrous. Ligule 3 mm long, glabrous, tip entire. Blade (3—)15—17 cm long, (2—)

10 mm wide, (sparsely) hairy on both sides, base rounded, densely hairy behind ligule,

pseudopetiole absent. Peduncles 1 per sheath, from upper and up to 2 lower sheaths.

Inflorescence branches 5—7 cm long, divergent. Sessile spikelet 5 mm long, 1.7 mm wide,

awned; lower glume 5 mm long, ovate, glabrous, basal part with 6 or 7 transverse rugosities

or (rudimentary) marginal nodules, upper part 9 veins, wings 0.2 mm wide, tip entire; upper

glume 5 mm long, 3 veins, glabrous, unawned; 1“ lemma 4.8 mm long; 2"? lemma 4 mm long,

upper half bifid, awned from notch, awn c. 17 mm long, geniculate. Pedicel c. 2 mm long, as

thick as rachis, one margin hairy.

Native; India to east Asia, south to Malesia. Single record 1898 from ‘Galang’ (Kallang).

Extinct (1898).

According to (Ridley 1925), it grew in waste ground or in places where the ground

was not too dry (Gilliland 1971).

Ischaemum timorense Kunth — Fig. 84.

Common Centipede Grass

Ridley 1907: 160; 1925: 203. Gilliland 1971: 264. Keng et al. 1998: 172.

I. macrurum Stapf ex Ridl. 1925: 203.

Culms with short stolons, 60—120 cm long; nodes hairy. Sheath 4—7 cm long, rounded at

base and more or less flattened in upper part, hairy at least at base and tip, margins glabrous or

hairy. Ligule 2—4 mm long, hairy at back, tip ciliolate. Blade (2—)4—26 cm long, 4—11

mm wide, glabrous or hairy, base rounded and hairy, pseudopetiole absent. Peduncles 1 and

from upper sheath. Inflorescence branches 2—10 cm long, divergent. Sessile spikelet 4.5—

6.8 mm long, 1.3—1.4 mm wide, elliptic, awned; lower glume 3.5—6 mm long, ovate, basal

part smooth and glabrous, upper part 6 or 7 veins and glabrous or hairy, wings 0O—0.4 mm

wide, tip bifid; upper glume 4.5—6 mm long, 7 veins, glabrous, awn 0.5—2 mm long; 1*

lemma 3—4.8 mm long; 2™ lemma 2.6—4.3 mm long, upper half bifid, awned from notch,

awn 5.5—17 mm long, geniculate or straight. Pedicel 2.8—3.3 mm long, as thick as rachis,

hairy on margins.

Native; India to east Asia, Malesia and Polynesia. First record 1889. Rather common.

Figure 83. Ischaemum rugosum Salisb., pair of spikelets.

Figure 84. Ischaemum timorense Kunth, pair of spikelets.

Figure 85. Leersia hexandra Sw.

Figure 86. Leptochloa chinensis (L.) Nees.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 87

88 Supplement of Gardens’ Bulletin Singapore 57 (2005)

In open places, on forest margins or in scrub.

Differences with J. ciliare are mentioned there.

Leersia Sw.

Leersia hexandra Sw. — Fig. 85.

Rice Grass

Ridley 1907: 148; 1925: 253. Gilliland 1971: 97. Keng et al. 1998: 173.

Culms single or tufted, geniculate, 5|0—100 cm long; nodes hairy, hairs deflexed. Sheath 4—

9.5 cm long, rounded, midrib (slightly) raised, glabrous, scaberulous at base, margins glabrous.

Auricles fused with ligule, triangular, glabrous. Ligule membranous, 2 mm long, glabrous,

truncate or emarginate. Young leaf blade inrolled. Blade (3.5—)11—14 cm long, 3—7 mm

wide, sparsely hairy, margin serrulate, base cuneate. Inflorescence panicle, central rachis 4.5—

9.5 cm long, branches 5—14; longest branch 1.7—5 cm long, basal 1/3—2/5 part without

spikelets, simple or branched. Pedicel c. 0.5 mm long, scaberulous, tip cup-shaped. Spikelets

single, 3.3—3.9 mm long, c. 1.3 mm wide, laterally flattened, 1-flowered, unawned; callus

more or less obtuse; only 2 bracts present, almost equally long, indurated; glumes absent;

lemma 3.3—3.9 mm long, veins 5 and with stiff hairs most prominent on midvein, acuminate;

palea 3.2—3.7 mm long, 3 veins, acute.

Native; pantropical. First record 1889. Rare.

In wet or swampy areas.

The two glumes are reduced to minute auricles forming a cup-shape at the tip of the

pedicel.

Leptochloa P.Beauv.

Lazarides 1980: 247—269.

Leptochloa chinensis (L.) Nees — Fig. 86.

Ridley 1907: 175 (excluding Singapore); 1925: 249. Gilliland 1971: 73. Lazarides

1980: 258. Keng et al. 1998: 173.

Culms single or tufted, erect to geniculate or with stolons, 30—120 cm long; nodes glabrous

or hairy. Sheath 5.5—10 cm long, keeled, glabrous, margins glabrous. Ligule membranous

(i.e. fused hairs), 1—1.3 mm long, hairy at abaxial side. Young leaf blade inrolled. Blade

8.5—19 cm long, 6—13 mm wide, midvein winged underneath, glabrous or upperside sparsely

hairy, base cuneate. Inflorescence panicle, central rachis 10—45 cm long, branches at least

25 and often in whorls of 2—4; longest branch 3—14 cm long, with spikelets from the base,

simple. Pedicel 0.5 mm long, scaberulous. Spikelets single, 2.1—4.3 mm long, 0.7—1.3 mm

wide, more or less laterally flattened, 3—7-flowered, unawned; the two lowest bracts 0.5—

0.6 times as long as spikelet; glumes and lemmas herbaceous; lower glume 0.7—2 mm long,

1 vein, glabrous, acute; upper glume 1.1—2.3 mm long, | vein, glabrous, acute to acuminate;

1* lemma 1.4—1.7 mm long, 3 veins, hairy at least near the veins, truncate to obtuse; 1“ palea

c. 1.4mm long, hyaline; upper florets as 1“ but gradually smaller, last floret sometimes much

reduced.

Native; S and E Asia, Australia. First record 1906. Rather rare.

Waste areas on sand or clay, dry to waterlogged soil, or in water.

Lepturus R.Br.

Nowack and Veldkamp 2002: 385—389.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 89

Lepturus repens (G.Forst.) R.Br. var. repens — Fig. 87, Plate 16.

Nowack and Veldkamp 2002: 388. — L. repens (G.Forst.) R.Br.: Ridley 1907: 182;

1925: 254 (both excluding Singapore). Gilliland 1971: 83. Keng et al. 1998: 173.

Culms erect or with stolons, 20—30 cm long; nodes glabrous. Sheath 4—6.5 cm long, rounded,

glabrous, margin glabrous. Ligule membranous, 0.5—1 mm long, tip ciliolate. Young leaf

blade inrolled. Blade 2.5—9 cm long, 2—4.5 mm wide, underside glabrous, upperside distinctly

ribbed, ribs flat and scaberulous, margins in upper half scaberulous, base cuneate and glabrous

or sparsely hairy. Inflorescence spike, rachis 7—17 cm long, 1.5—-2 mm wide. Pedicel absent.

Spikelets single, alternately in rachis cavities, 9—14 mm long (including awn), dorsiventrally

flattened, 2-flowered, awned; lowest bract (including awn) as long as and 2™ c. 0.5 times as

long as spikelet (rudimentary lower glume ignored: see note); upper glume (including awn)

9—14 mm long, indurated, 7 veins, glabrous, at tip gradually tapering into scaberulous awn;

both lemmas 4.5—4.8 mm long, thin herbaceous; both paleas 4.7 mm long, thin herbaceous.

Native; East Africa, South and East Asia, Australia, Polynesia and Hawaii. First record

1941. Rather common.

Sandy beaches above high tide mark, preferring coral sands.

The lower glumes are easily overlooked. They are about 0.3 mm long and hidden

between the rest of the spikelet and the rachis (adaxial).

—

Lophatherum Brongn.

Lophatherum gracile Brongn. — Fig. 88, Plate 19.

Crested Grass, Rumput Jarang

Ridley 1907: 181; 1925: 253. Gilliland 1971: 55. Keng et al. 1998: 173.

Culms single or loosely tufted, erect to geniculate, 40—60 cm long; some roots partly thickened,

up to 3 mm diameter; nodes glabrous. Sheath 3.5—10 cm long, somewhat keeled, glabrous to

sparsely hairy, with transverse veins, one margin hairy. Ligule membranous, 0.7 mm long, tip

ciliolate. Young leaf blade inrolled. Blade S—16 cm long, 8.5—22 mm wide, with transverse

veins present, glabrous to sparsely hairy, pseudopetiole 7—20 mm long and densely short-

hairy. Inflorescence panicle, central rachis 15—-32 cm long, branches 6—14 in whorls of 1—

4; longest branches 7—12 cm long, simple or branched. Pedicel 0O—0.2 mm long, scaberulous

to sparsely hairy at tip. Spikelets single, 5.7—8 mm long, 1.4—1.6 mm wide, somewhat

laterally flattened, 4—12-flowered with 1* floret bisexual and 2™—12" sterile, awned; the

two lowest bracts 0.3—0.5 times as long as spikelet; rachilla not articulate, between 1“ and

2™ lemma 1—3.7 mm long and glabrous, extension beyond last lemma 1.5 mm long and

scaberulous; glumes and lemmas herbaceous; lower glume 2.9—3.2 mm long, slightly

enveloping, 3 or 5 veins, hairy at least at tip, truncate to obtuse; upper glume c. 4 mm long,

slightly enveloping, 5 veins, sparsely hairy, acuminate; 1“ lemma 4.7—5.5 mm long, 9—11

veins, glabrous, tip bifid and ciliolate, awn from notch 1.2 mm long and scaberulous; 1“ palea

2—4.3 mm long, hyaline; 2"-—12™ lemmas as first but gradually smaller, not visible in young

spikelets before flowering time; 2"°—12" paleas absent.

Native; S and SE Asia and Australia. First record 1880s. Rare.

Open spots on moist soil in primary and older secondary forests.

For the difference in its vegetative parts with Centotheca lappacea see there.

Melinis P.Beauv.

Melinis repens (Willd.) Zizka — Fig. 89, Plate 20.

Natal Grass

Keng et al. 1998: 173.

Rhynchelytrum repens (Willd.) C.E.Hubb.: Gilliland 1971: 150.

90 Supplement of Gardens’ Bulletin Singapore 57 (2005)

Culms tufted, erect to somewhat geniculate, 55-80 cm long; nodes (upper distinctly) hairy.

Sheath 7—17 cm long, rounded, lower ones densely hairy, upper glabrescent, margins glabrous.

Ligule row of hairs, 0.7—1 mm long. Young leaf blade inrolled. Blade (4—)9—14 cm long,

(1.5—)4 mm wide, sparsely hairy in basal half at underside, base rounded and sparsely hairy.

Inflorescence panicle, central rachis 10—17 cm long, branches 8—12; longest branch 3—7

cm long, scaberulous to sparsely hairy at base, branched. Pedicel 1—3.5 mm long, thinner

than branch, purplish, sparsely hairy near tip. Spikelets single, 3.5—4.2 mm long, 2 mm

wide, yawning, somewhat laterally flattened, 2-flowered with 1* floret male and 2™ bisexual,

awned; lowest bract 0.2 times as long as and 2" as long as spikelet; lower glume 0.9—1.2 mm

long, herbaceous, | vein, densely hairy, hairs up to 1.5 mm long and reddish, emarginate;

upper glume 3.2—3.6 mm long, herbaceous, 5 veins, densely hairy, hairs up to 6.5 mm long

and red, truncate to obtuse, awn from tip and 0.6—2 mm long; 1* lemma as upper glume,

3.1—3.5 mm long, awn 0.5—1.5 mm long; 1* palea 2.7—3 mm long, hyaline; 2"? lemma

1.8—2 mm long, thin herbaceous, 5 veins, emarginate; 2™ palea 1.7—2 mm long, thin

herbaceous.

Not native; Africa, introduced in Asia, Australia and America. First record 1896 from

the Botanic Gardens, 1939 in the wild. Rather rare.

Open spaces on dry sandy soil.

Mnesithea Kunth

Veldkamp et al. 1986: 281—307.

Culms erect. Inflorescence spike-like raceme, rachis articulated, joints at articulation with a

central knob. Spikelets in pairs of 1 sessile and 1 pedicelled. Sessile spikelet more or less

dorsiventrally flattened, 2-flowered with 1* floret sterile and 2™ bisexual, unawned; lowest

bract as long as spikelet, 2" slightly shorter but longer than the florets; lower glume flattened,

submarginally keeled, indurated; upper glume (thick) herbaceous; lemmas and paleas hyaline

(or 2™ palea absent).

The internodes of the inflorescence are corky, allowing them to float on water after

articulation. The central knob (or elaiosome) in M. laevis (and Rottboellia cochinchinensis as

well) is reported to contain an oily substance that attracts ants (Veldkamp ef al. 1986: 295).

This still needs to be confirmed for the two species included below.

Rottboellia differs from this genus in having a male 1* floret (Veldkamp et al. 1986).

la. Culms 90—210 cm long. Sessile spikelet 3.9—4.8 mm long. Lower glume

lanceolate, veins not thickened and raised, keels at base with 2—7 gland-

like warts, in upper half winged. M. glandulosa

1b. Culms c. 40 cm long. Sessile spikelet c. 1 mm long. Lower glume orbicular,

veins and transverse veins thickened and raised, making squarish

depressions, keels thickened and raised and without warts.

M. granularis

Figure 87. Lepturus repens (G.Forst.) R.Br. var. repens.

a. spikelet, b. rachis, c. exploded view of spikelet.

Figure 88. Lophatherum gracile Brongn.

Figure 89. Melinis repens (Willd.) Zizka.

Figure 90. Neyraudia arundinacea (L.) Henrard var. zollingeri (Biise) Henrard.

Field Guide to the Grasses of Singapore (Excluding the Bamboos)

Fig. 89

92 Supplement of Gardens’ Bulletin Singapore 57 (2005)

Mnesithea glandulosa (Trin.) De Koning & Sosef — Fig. 91.

Mat Grass

Veldkamp et al. 1986: 288. Keng et al. 1998: 174.

Coelorachis glandulosa (Trin.) Stapf ex Rid]. 1925: 204. Gilliland 1971: 270.

Rottboellia glandulosa Trin.: Ridley 1907: 161.

Culms single or tufted, 90—210 cm long; internodes hollow, 4—10 mm diameter; nodes

glabrous. Sheath 10—17 cm long, rounded, glabrous. Auricles triangular, hairy. Ligule

membranous, 1—1.5 mm long, ciliolate. Young leaf blade inrolled. Blade 29—45 cm long,

13—24 mm wide, glabrous, base cuneate to somewhat rounded and hairy. Peduncles 1—5

per sheath from upper and lower sheaths. Inflorescence 7—14 cm long. Pedicel 4.5—5 mm

long, not fused to rachis. Sessile spikelet 3.9—4.8 mm long, 1.4—1.6 mm wide; lower glume

3.9—4.8 mm long, 7 veins, glabrous or hairy, keels at base with 2—7 glandlike warts and in

upper half winged, wings 0.5—0.9 mm wide; upper glume 3.6—4.5 mm long, boat-shaped, 3

veins, midvein keeled and ciliolate; 1 lemma 2.6—3.2 mm long; 1“ palea 1.7—2 mm long;

2™ lemma 2.6—2.8 mm long; 2" palea c. 2.5 mm long. Pedicelled spikelet reduced, lower

glume winged on one side only, lemmas and paleas absent.

Native; Thailand, Peninsular Malaysia, Singapore, Sumatra, Java, Borneo, Philippines.

First record 1887. Rather rare.

Coast-near woodlands (abandoned plantations, forests etc.), on dry to moist soil.

Ridley (1907, 1925) mentioned it grew on riverbanks.

Rottboellia cochinchinensis is different in having hairy sheaths and upper surfaces of

blades; other differences are mentioned in note under Mnesithea and in the key to flowering

grasses (p. 22, lead 10).

Mnesithea granularis (L.) De Koning & Sosef — Fig. 92.

Veldkamp et al. 1986: 294. Keng et al. 1998: 174.

Hackelochloa granularis (L.) Kuntze: Gilliland 1971: 277.

Manisuris granularis L.f.: Ridley 1907: 163; 1925: 205 (both excluding Singapore).

Culms single, c. 40 cm long; basal internodes c. 3 mm diameter; nodes hairy. Sheath c. 2.5 cm

long, rounded, densely hairy, margin densely hairy, auricles absent. Ligule membranous, 1—

1.5 mm long, hairy. Young leaf blade folded along midrib. Blade 7—9 cm long, 6—8 mm

wide, hairy, base rounded to cordate and pectinate. Peduncles | per sheath, from 1—3 sheaths.

Inflorescence 1—2 cm long. Pedicel c. 0.8 mm long, furrowed, fused to rachis to form cavity

that holds sessile spikelet. Sessile spikelet c. 1 mm long, c. 1 mm wide; lower glume c. 1 mm

long, veins and transverse veins thickened and raised to make squarish depressions, glabrous,

wings absent; upper glume c. | mm long, more or less flattened, veins not obvious; 1* lemma

c. 0.8 mm long; 1“ palea absent; 2" lemma as 1“; 2"! palea absent (see note). Pedicelled

spikelet lower glume c. 1.2 mm long, winged on one side only, acute; upper glume c. 1.2 mm

long, boat-shaped, keel winged, acute.

Doubtfully native; pantropical. Single record 1949 from Kampung Kelapa at Pulau

Ayer Merbau, now part of Jurong Island. Extinct (1949).

According to Gilliland (1971) the 2" palea is 0.6 mm long.

In Peninsular Malaysia, it was also found only once. In Java it grows much bigger

(Gilliland 1971), suggesting that it might not be native in Singapore.

Figure 91. Mnesithea glandulosa (Trin.) De Koning & Sosef, pair of spikelets.

Figure 92. Mnesithea granularis (L.) De Koning & Sosef.

a. sessile spikelet, b. pedicelled spikelet.

Figure 93. Oryza sativa L.

Field Guide to the Grasses of Singapore (Excluding the Bamboos)

RAM Rlabetin te ane.

Fig. 93

94 Supplement of Gardens’ Bulletin Singapore 57 (2005)

Neyraudia Hook.f.

Neyraudia arundinacea (L.) Henrard var. zollingeri (Biise) Henrard — Fig. 90.

Veldkamp 1999b: 234. Duistermaat 2004: 42.

Neyraudia madagascariensis Hook.f. var. zollingeri Hook.f.: Ridley 1907: 176 (excluding

Singapore).

Triraphis madagascariensis sensu Ridl., 1925: 251 (excluding Singapore).

Neyraudia reynaudiana (Kunth) Keng ex Hitchc.: Gilliland 1971: 61.

Culms up to 3 m long; internodes solid; nodes glabrous. Sheath 9—16 cm long, rounded,

hairy at tip, margins glabrous. Ligule row of hairs, 3—5 mm long. Young leaf blade inrolled.

Blade c. 40 cm long, 7—20 mm wide, glabrous, base rounded and with short hairs. Inflorescence

panicle, silky hairy, central rachis c. 65 cm long, branches many and in groups of 1—4;

longest branch c. 18 cm long, glabrous, branched. Pedicel 3—4 mm long, scaberulous. Spikelets

single, c. 10.5 mm long (including awns), more or less laterally flattened, herbaceous, 7-

flowered with 1 floret sterile and 2°‘—7" bisexual, awned; rachilla articulated between 1“

and 2" floret; the two lowest bracts 0.2—0.3 times as long as spikelet; lower glume 2.5 mm

long, at base almost enveloping, 3 veins, glabrous, acute; upper glume 2.7 mm long, 3 veins,

midvein somewhat keeled and ciliolate, acute; 1 lemma 3.9 mm long, 3 or 5 veins, glabrous,

acute; 1*' palea absent; 2°’ lemma 3.5 mm long, margin with c. 2 mm-long silky hairs, midvein

protruding into c. 0.8 mm-long awn; 2™ palea 2.3 mm long; 3"—7" florets as 2"? with lemmas

up to 5 mm long and awns up to 2 mm long.

Doubtfully native; India to SW China, Peninsular Malaysia, Singapore, Sumatra, Java,

Lesser Sunda Islands; naturalized in North America and Caribbean. Single record 1959 from

Bartley Road. Extinct in the wild (1959).

It was reported as growing on infertile soil, in thickets and unshaded places (Veldkamp

1999). In Peninsular Malaysia it is more common in the north and grows in less wet places

than Phragmites (Gilliland 1971). It is not known whether in Singapore it was native. Today,

itis only known planted in the Botanic Gardens.

It is different from Phragmites in the solid culm internodes and hairy lemmas (hollow

respectively glabrous in Phragmites). Differences with other species with silky-hairy panicles

are mentioned in a note under /mperata.

Oplismenus P.Beauv.

Scholz 1981: 1—213. Veldkamp 1999b: 234.

Culms stoloniferous, 25—60 cm long; nodes (sparsely) hairy. Sheath rounded, hairy, margins

hairy. Auricles triangular, hairy. Ligule row of hairs, fused in basal part. Young leaf blade

inrolled. Blade hairy, base rounded. Inflorescence panicle, central rachis flattened to somewhat

triquetrous and hairy; branches 3—5, more or less flattened to triquetrous, 0.4 mm wide.

Pedicel rounded, hairy. Spikelets single, not to laterally flattened, 2-flowered with 1* floret

sterile and 2™ bisexual, awned; the two lowest bracts (excluding awns) 0.6—0.8 times as long

as spikelet; glumes and 1“ lemma (thin) herbaceous; lower glume not enveloping, 5. veins,

awned; upper glume shorter awned; 2" floret leathery, glabrous.

la. Awns antrorsely scaberulous, filiform, dull. O. burmanni

1b. Awns smooth, rather thick, viscid, shiny. O. compositus

Oplismenus burmanni (Retz.) P.Beauv. — Fig. 94.

Ridley 1907: 145; 1925: 221 (both excluding Singapore). Gilliland 1971: 172 (excluding

Singapore). Scholz 1981: 57. Veldkamp 1999b: 234. Duistermaat 2004: 42.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 95

Sheath 0.7—1.9 cm long. Ligule 0.8—1 mm long, basal part 0.3—0.4 mm long, free hairs

0.5—0.6 mm long. Blade 2.5—3.5 cm long, 5—9 mm wide. Inflorescence 2.5—5 cm long;

longest branch 0.5—1.2 cm long, simple. Spikelet 2.3—3.2 mm long, 1 mm wide; lower

glume 1.7—2 mm long, sparsely to densely hairy, awn 2—14 mm long and antrorsely

scaberulous; upper glume as lower, 1.5—1.8 mm long, awn 1.5—3.5 mm long; 1* lemma

2.2—2.8 mm long, 7 veins, sparsely to densely hairy, awn 0.7—0.9 mm long; 1* palea absent;

2™ lemma 1.6—2.4 mm long; 2™ palea 1.5—2.3 mm long.

Probably not native; pantropical. First record 1959. Rare.

Shaded places, between rocks, in weedy fields, plantations and lawns.

This species is in Singapore only known from a few roads bordering the Botanic

Gardens and in Peninsular Malaysia only from Kedah, Pahang and Penang.

Oplismenus compositus (L.) P.Beauv. — Fig. 95.

Common Wood Grass

Ridley 1907: 145; 1925: 221 (both excluding Singapore). Gilliland 1971: 171 (excluding

Singapore). Scholz 1981: 85. Veldkamp 1999b: 234. Duistermaat 2004: 42.

Sheath 1.5—3.5 cm long. Ligule 1.2 mm long, basal part 0.5 mm long, free hairs 0.7 mm

long. Blade 4.5—5.5 cm long, 5—7 mm wide. Inflorescence 2.5—9 cm long; longest branch

1—1.5 cm long, simple or branched. Spikelet 2.5—3 mm long, 1.2 mm wide; lower glume

2.4 mm long, sparsely to densely hairy, awn 4—12 mm long and smooth; upper glume as

lower, 2 mm long, awn 0.4 mm long; 1‘ lemma 2.2 mm long, 9 veins, glabrous to margins

ciliate, 0.1 mm apiculate; 1‘ palea 1.5 mm long, hyaline; 2"? lemma 1.9 mm long; 2™ palea 1.9

mm long.

Status unknown; pan(sub)tropical. First record 1959. Rare.

Moderately shaded soil, open places in primary and secondary forest.

The inflorescence branches and spikelets are shorter than mentioned in Veldkamp

(1999b), but Scholz (1981) recognized them as O. compositus var. rariflorus (J.Presl) Scholz.

The wider species concept of the latter is followed, but not the recognition of her overlapping

varieties.

The species is known throughout Peninsular Malaysia but is there more common in

the north (Gilliland 1971).

Oryza L.

Duistermaat 1987: 157—193.

Oryza rufipogon Griff. has been mentioned for Singapore by Duistermaat (1987: 171),

but material has not been found. As its occurrence is often associated with rice cultivation, it

is now probably extinct here. It is different from O. sativa as follows:

la. Spikelets articulating, falling off at maturity. Anthers 3.5—6.2 mm long. O. rufipogon

1b. Spikelets not articulating, remaining attached to pedicel at maturity. Anthers 0.8—2.5

mm long. O. sativa

Oryza sativa L. — Fig. 93.

Rice, Padi

Gilliland 1971: 99. Duistermaat 1987: 174.

Culms single or in small tufts, erect to ascending, 50—190 cm long; nodes glabrous. Sheath

glabrous, (obscure) transverse veins present. Auricles deciduous, linear-lanceolate, falcate,

1—5 mm long, 0.5—1 mm wide, with up to 2 mm long hairs. Ligule membranous, (6—)10—

96 Supplement of Gardens’ Bulletin Singapore 57 (2005)

36 mm long, glabrous. Young leaf blade inrolled. Blade 24—60 cm long, 6—22 mm wide,

glabrous, margins scabrous or rarely smooth. Inflorescence panicle, central rachis 9—30 cm

long, branches many; longest branches 2—13 cm long, simple or branched. Pedicel 1—7 mm

long, glabrous to slightly pubescent, at tip cup-shaped. Spikelets single, 7—10.9 mm long,

2.65—4.6 mm wide, not articulating, laterally flattened, 3-flowered, awned or unawned; the

two lowest bracts 0.2—0.4 times as long as spikelet; lower and upper glume absent; 1“ and 2"

lemma sterile, 1.4—4 mm long, herbaceous, glabrous, margin serrate; 1‘ and 2" palea absent;

3™ lemma fertile, 6—10 mm long, strongly indurated, finely reticulate, covered by glassy

hairs, 5 veins, acuminate, awn from tip and up to 60 mm long, antrorsely scaberulous, callose

at base; 3™ palea as 3" lemma, 5.8—9.5 mm long, 3 veins, awn 0.2—0.45 mm long.

Not native; Southeast Asia, now cultivated in (sub-)tropical regions. First record 1880s.

Cultivated (last record 1970).

Rice is no longer cultivated in Singapore but it can still be found as a casual from bird-

seed.

The glumes are reduced to minute auricles that form a cup at the tip of the pedicel.

Ottochloa Dandy

Ottochloa nodosa (Kunth) Dandy — Fig. 96.

Slender Panic Grass

Gilliland 1971: 143. Keng et al. 1998: 174.

Hemigymnia fusca Ridl. 1925: 228.

Hemigymnia multinodis Stapf; Ridley 1925: 228.

Panicum nodosum Kunth; Ridley 1907: 137.

Culms erect to geniculate or creeping to scrambling, 30—120 cm long; nodes glabrous or

hairy. Sheath 3—6.5 cm long, rounded, glabrous to densely patent hairy, at least one margin

hairy. Ligule membranous, 0.5—0.7 mm long, tip ciliolate. Young leaf blade inrolled. Blade

(S—)8—17 cm long, (S—)7—12 mm wide, glabrous to densely hairy, base rounded and

margin glabrous to sparsely hairy, pseudopetiole 0.7—2 mm long. Inflorescence panicle, central

rachis 10—26 cm long, branches (3—)6—9 in groups of 1—3; longest branch (1—)6.5—16

cm long, (simple or) branched, ultimate branches short and not or only slightly overlapping.

Pedicel 0.5—1.3 mm long. Spikelets single, 2.7—3.3 mm long, 0.8—1 mm wide, not flattened,

2-flowered with 1“ floret sterile and 2! bisexual, unawned; the two lowest bracts 0.4—0.5

times as long as spikelet; glumes and 1* lemma herbaceous; lower glume 1.2—1.4 mm long,

margin (at tip) ciliate, 3—5 veins, obtuse; upper glume as lower, 1.5—1.7 mm long, 7 veins;

1* lemma 2.5—3 mm long, glabrous or hairy, 7 or 9 veins, truncate to obtuse; 1* palea absent;

2™ lemma 2.3—2.6 mm long, glabrous, 5 veins, truncate, 0.1 mm long apiculate, somewhat

indurated; 2™ palea 2—2.3 mm long.

Native; India and west Malesia. First record 1889. Common.

In open to (partly) shaded places, forest margins, also on wet soil.

This species is very variable with respect to indument density on sheath and blade, the

number and the length of the panicle branches and the colour of the spikelets (green to purple).

Different forms may grow next to each other. A monstrosity with the upper floret having 4

ovaries each with 2 or 3 stigmas occurs rarely.

The upper glumes of Panicum species are 0.9—1 times as long as the spikelet.

Figure 94. Oplismenus burmanni (Retz.) P.Beauv.

Figure 95. Oplismenus compositus (L.) P.Beauv.

Figure 96. Ottochloa nodosa (Kunth) Dandy.

Figure 97. Panicum auritum J.Presl ex Nees.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 97

sh estates

SSS

Fig. 97

98 Supplement of Gardens’ Bulletin Singapore 57 (2005)

Panicum L. |

Veldkamp 1996a: 181—216. Aliscioni et al. 2003: 796—821. Duistermaat 2004: 35—37.

Sheath rounded. Auricles absent. Young leaf blade inrolled. Inflorescence panicle, central

rachis present, branches 6 or more (or less in P. laxum), rounded, branched (with only 2

spikelets in P. trichocladum). Pedicel present. Spikelets single or paired, not flattened, 2 (or

3)-flowered with 1‘ floret sterile or male and 2™ bisexual, unawned; 2™ lowest bract 0.9—1

times as long as the spikelet; glumes (thin) herbaceous; 1* lemma as upper glume; 1* palea

hyaline; 2"? lemma and palea (slightly) indurated, palea (almost) as long as lemma.

Panicum is retained here in its traditional wider delimitation (Duistermaat 2004: 35),

rather than following Aliscioni et al. (2003).

Inflorescences with a few 3-flowered spikelets have been seen in P. maximum.

The key follows Veldkamp (1996a).

la. Lower glume collar-shaped (Fig. 18a). 2

1b. Lower glume ovate (Fig. 18b) or triangular. 3

2a. Sheaths and blades glabrous. Ligule a hairy collar. Spikelets 3.3—4.5 mm

long. Upper glume 3.3—4.1 mm long. 1* floret sterile; palea up to 0.7

times as long as the lemma (or absent). P. paludosum

2b. Sheath margins and blades hairy. Ligule a membranous ciliolate collar.

Spikelets 2.6—3.25 mm long. Upper glume 2.6—3 mm long. 1|* floret

male; palea more than 0.8 times as long as the lemma. P. repens

3a. Ligule a membranous glabrous or ciliolate collar (hairs at the tip much

shorter than the basal part). +

3b. Ligule either a ciliate collar (hairs at the tip (almost) as long as the basal

part), or a row of hairs, or membranous with abaxially a row of long

hairs. 7

4a. Blades ovate-lanceolate, at base cordate, margin pectinate. Lower glume

1.6—3 mm long. P. brevifolium

4b. Blades linear, at base truncate to cordate, margin glabrous (or pectinate in

P. auritum). Lower glume 0.3—1.3 mm long. ~)

5a. Panicle branches naked at base. Lower glume triangular, 0.1—0.2 times as

long as the spikelet. 1‘ palea present; 2"* lemma tip apiculate, incurved.

P. trichocladum

5b. Panicle branches with spikelets (almost) to base. Lower glume ovate, 0.3—

0.7 times as long as the spikelet; 1‘ palea absent or present; 2"’ lemma

tip obtuse to acuminate, straight. 6

6a. Culms 80—120 cm long. Spikelet 2—3 mm long. 1“ palea up to 0.75 times

as long as 1* lemma. P. auritum

6b. Culms 20—45 cm long. Spikelet 1.3—1.4 mm long. I“ palea as long as |"

lemma. P. laxum

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 99

7a. (3) Culms scrambling. Nodes glabrous (to puberulous). Pedicels smooth,

sometimes viscid. Spikelets obtuse; 2" lemma apiculate.

P. sarmentosum

7b. Culms erect or geniculate at base. Nodes hairy. Pedicels scaberulous or

hairy under the spikelet, not viscid. Spikelets acute to caudate; 2"! lemma

obtuse, acute, or acuminate. 8

8a. Glumes with a distinct internode. Upper glume and 1* lemma with transverse

veinlets. 2" lemma sessile, smooth, shiny, obtuse to acute.

P. luzonense

8b. Glumes without a distinct internode. Upper glume and 1“ lemma without

transverse veinlets. 2" lemma stipitate, transversely rugulose, dull,

acuminate. P. maximum

Panicum auritum J.Presl ex Nees — Fig. 97.

Giant Panic Grass, Rumput Kumpai

Ridley 1907: 135; 1925: 226. Gilliland 1971: 137. Veldkamp 1996a: 187. Keng et al.

1998: 174.

Culms single, erect or with short rhizomes, 80—120 cm long; internodes hollow; nodes

glabrous. Sheath 7—9 cm long, glabrous, minute transverse veins present (visible only from

inside), one margin hairy. Ligule membraneous, 0.4—1 mm long, glabrous, tip entire or

ciliolate. Blade 18—30 cm long, 9—23 mm wide, glabrous, margin scaberulous, base rounded

to cordate and pectinate. Inflorescence spike-like, central rachis 16—40 cm long, branches

erect; longest branch 2—12 cm long, with spikelets almost from the base. Pedicel 0.5—1.8

mm long, scaberulous. Spikelet 2.2—3 mm long, 0.7—0.8 mm wide, acute to acuminate,

glumes without a distinct internode, 1“ floret sterile; lowest bract 0.4 times as long as spikelet;

lower glume 0.8—1.3 mm long, ovate, 3—5 veins, glabrous; upper glume 2—2.5 mm long, 5

veins, transverse veins present (obscure), glabrous; 1** lemma 1.6—2.8 mm long, acute; 1*

palea 0.6—1.7 mm long; 2" lemma 1.6—2.4 mm long, scaberulous at tip. Anthers 2, 0.7—

0.8 mm long.

Native; India to S. China and Indonesia, New Guinea. First record 1880s. Extinct

(1966).

Moist places, river banks, ditches.

The Singapore material is different from that of Veldkamp (1996a) in having sheaths

with one margin hairy and blades that are pectinate at base.

Hymenachne amplexicaulis is different in having culms filled with pith, blades hairy

on the upperside and long-acuminate spikelets (3—)3.9—4.9 mm long with the upper glume

0.7—0.9 times as long as the spikelet. Sacciolepis has blades up to 6 mm wide, panicles with

up to 0.5 cm-long branches and gibbous spikelets.

Panicum brevifolium L. — Fig. 98.

Short-leaved Panic Grass

Gilliland 1971: 139. Veldkamp 1996a: 189. Keng et al. 1998: 175.

P. hirtifolium Ridl. 1907: 141; 1925: 228 (both excluding Singapore).

P. ovalifolium Poir. in Lam.: Ridley 1907: 141; 1925: 227.

Culms creeping to geniculate, 20—5O cm long; internodes hollow; nodes glabrous. Sheath

1.5—5 cm long, transverse veins absent, glabrous, margins hairy. Ligule membranous, 0.5

mm long, glabrous, tip ciliolate. Blade 3—4.5 cm long, 15—20 mm wide, sparsely hairy,

100 Supplement of Gardens’ Bulletin Singapore 57 (2005)

base cordate and pectinate. Inflorescence open, central rachis 9—11 cm long, branches erecto-

patent; longest branch 5.5 cm long, empty at base. Pedicel 1.5 mm long, glabrous. Spikelet

1.6 mm long, 0.6 mm wide, acute, glumes with a distinct internode, 1“ floret sterile; lowest

bract as long as spikelet; lower glume 1.6 mm long, ovate, sparsely hairy; upper glume 1.5

mm long, 5 veins, transverse veins absent, sparsely hairy; 1** lemma 1.6 mm long, truncate; 1*

palea 1.3 mm long; 2" lemma 1.3 mm long, smooth. Anthers 3, 0.75—1 mm long.

Possibly native; Paleotropics, possibly introduced in at least part of Malesia. Single

record 1894 from Government Hill (now the Istana). Extinct (1894).

Shaded places.

The spikelets are shorter than reported by Veldkamp (1996a: 1.9—2.1 mm long).

Panicum laxum Sw. — Fig. 99.

Duistermaat 2004: 36.

Steinchisma laxa (Sw.) Zuloaga: Simon 2003: 562.

Culms single or tufted, geniculate to prostrate, 20—45 cm long; internodes hollow; nodes

glabrous. Sheath 2.5 minute transverse veins present, glabrous, margin hairy

at least at tip. Ligule membranous, 0.3—0.4 mm long, glabrous, tip ciliolate. Blade 4—11 cm

long, 2—10 mm wide, glabrous, base rounded. Inflorescence open, central rachis 5—13 cm

long, branches patent; longest branch 2.5—6 cm long, with spikelets almost from the base at

adaxial side only, branchlets appressed to 1* order branch (spike-like). Pedicel 0.5—1 mm

long, scaberulous. Spikelet 1.3—1.4 mm long, 0.7 mm wide, obtuse, glumes without a distinct

internode, |* floret sterile; lowest bract 0.4—0.5 times as long as spikelet; lower glume 0.5—

0.6 mm long, ovate, 1 or 3 veins, midvein near tip keeled and with few spicules; upper glume

as lower, 1.2 mm long, 5 veins, transverse veins absent; 1** lemma 1.2—1.3 mm long, 3 veins,

obtuse to acute; 1*' palea 1.2—1.3 mm long; 2™ lemma 1|.1—1.2 mm long, scaberulous at tip.

Anthers 2, 0.5 mm long.

Not native; America, introduced in tropical Africa and Australia and according to

Veldkamp (pers. comm.) recently also found in Borneo. First record 2002. Rare.

Open to shaded roadsides or fields, on clayey or sandy soil or in cracks of concrete

drain.

The inflorescence is reminiscent of that of Urochloa subquadripara, which differs in

the simple branches and much longer spikelets (3.5—4.3 mm). The 1“ paleas are reported to

enlarge (Aliscioni et al. 2003), but this could not be ascertained.

Plants with single and prostrate culms (rather than being tufted and geniculate) have a

potential as a turf grass for shaded conditions (Simon 2003).

Panicum luzonense J.Presl — Fig. 100.

Ridley 1907: 136. Veldkamp 1996a: 195. Keng et al. 1998: 175.

P. caesium Nees ex Hook.f.: Ridley 1925: 225.

P. cambogiense Balansa: Gilliland 1971: 141.

Culms tufted, geniculate, 40—50 cm long; internodes hollow; nodes hairy. Sheath 6—9 cm

long, transverse veins absent, densely patent hairy, margins hairy. Ligule row of hairs fused in

basal half, 1—2.1 mm long. Blade 9—14 cm long, 5—7 mm wide, hairy, base rounded and

pectinate. Inflorescence open, central rachis 15—30 cm long, branches patent; longest branch

Figure 98. Panicum brevifolium L.

Figure 99. Panicum laxum Sw.

Figure 100. Panicum luzonense J.Presl.

Figure 101. Panicum maximum Jacq.

Figure 102. Panicum paludosum Roxb.

Field Guide to the Grasses of Singapore (Excluding the Bamboos)

101

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Fig. 102

102 Supplement of Gardens’ Bulletin Singapore 57 (2005)

7.5—15 cm long, with spikelets from the base or not. Pedicel 0.8—3 mm long, scaberulous.

Spikelet 2—2.1 mm long, 0.7—1.1 mm wide, acute, glumes with a distinct internode, 1*

floret sterile; lowest bract 0.5 times as long as spikelet; lower glume 0.9—1 mm long, ovate,

5 or 7 veins, glabrous; upper glume 1.6—1.9 mm long, 7—9 veins, transverse veins present,

glabrous; 1“ lemma 1.6—1.8 mm long, 5—7 veins, obtuse; 1* palea 1.6—1.7 mm long; 2"

lemma |.3—1.4 mm long, smooth. Anthers 3, 0.7 mm long.

Native; Sri Lanka, India to Australia. First record 1894. Extinct (1959).

In open, unshaded to moderately shaded roadsides, waste areas, beaches, fields etc.

The upper glume and |“ lemma have fewer veins than reported in Veldkamp (1996a:

upper glume 9—11-veined, 1* lemma 7—11-veined).

Panicum maximum Jacq. — Fig. 101, Plate 18.

Guinea Grass, Rumput Benggala

Ridley 1907: 140; 1925: 226. Gilliland 1971: 132. Veldkamp 1996a: 197. Keng et al.

1998: 175.

Urochloa maxima (Jacq.) R.D.Webster: Aliscioni et al. 2003: 810.

Culms with short rhizomes, erect to geniculate, 100—200 cm long; internodes hollow or

solid upwards; nodes (densely) hairy. Sheath 10—34 cm long, transverse veins absent, glabrous

or hairy, at least one margin hairy. Ligule membranous, 1—3 mm long, hairs at tip 0.3—0.5

mm long, abaxial row of hairs 1—3 mm long. Blade (9—)25—50 cm long, 6.5—25 mm

wide, glabrous, margin scaberulous or hairy near base, base cuneate to slightly rounded and

short-hairy. Inflorescence open, central rachis 27—50 cm long, branches patent; longest branch

13—32 cm long, naked at basal 2—9 cm. Pedicel 0.3—1.3(—3) mm long, scaberulous or

with few long hairs. Spikelet 3.2—3.6 mm long, 1—1.2 mm wide, acute, glumes without a

distinct internode, 1* floret male; lowest bract 0.4—0.5 times as long as spikelet; lower glume

1.4—1.5 mm long, ovate, 3 (or 5) veins, glabrous; upper glume 2.4—3.3 mm long, 5 veins,

transverse veins absent, glabrous; |*' lemma as upper glume, 3—3.5 mm long; |* palea 2.6—

3 mm long; 2" lemma 2.3—2.5 mm long, transversely rugulose; 3" floret sometimes present,

reduced. Anthers 3, 1—1.5 mm long.

Not native; Africa, introduced (sub-)pantropically. First record 1901. Common.

In disturbed areas, grassy places, riverbanks, thickets, margins of secondary forest,

etc.

This species was originally introduced as a fodder crop.

Panicum paludosum Roxb. — Fig. 102.

Gilliland 1971: 134 (excluding Singapore). Veldkamp 1996a: 200. Duistermaat 2004: 35.

P. proliferum auct., non Lam.: Ridley 1907: 139; 1925: 225 (both excluding Singapore).

Culms floating in water, c. 70 cm long; internodes hollow; nodes glabrous. Sheath 7—9 cm

long, transverse veins absent, glabrous, margins glabrous. Ligule row of hairs fused at basal

0.5 mm, 1.5 mm long. Blade 12—16 cm long, 7—9 mm wide, glabrous, base rounded.

Inflorescence open, central rachis 11—15 cm long, branches erecto-patent; longest branch

6—9 cm long, naked at base. Pedicel 1.5—2 mm long, scaberulous. Spikelet 3.4—3.5 mm

long, 1 mm wide, acute, glumes without a distinct internode, 1“ floret sterile; lowest bract

0.15—0.25 times as long as spikelet; lower glume 0.6—0.8 mm long, collar-shaped, 0 veins,

glabrous; upper glume 3.3—-3.4 mm long, glabrous; 1‘ lemma c. 3.2 mm long, acute; 1* palea

1.5—2.1 mm long; 2"? lemma 2—2.2 mm long, smooth. Anthers 3, c. 1.2 mm long.

Possibly native; India, S$. China, southward to Australia. First record 1959. Extinct

(1965).

Aquatic, in shallow water, bogs, ditches, rice fields etc.

Its natural status is doubted, as in Peninsular Malaysia it is known only from the northern and

central states of Kedah, Pahang and Penang (Duistermaat 2004).

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 103

Panicum repens L. — Fig. 103, Plate 21.

Creeping Panic Grass

Ridley 1907: 139; 1925: 225. Gilliland 1971: 135. Veldkamp 1996a: 202. Keng et al.

1998),.175:

Culms with stolons, up to 85(—150) cm long; internodes hollow; nodes glabrous. Sheath 4—

8.5 cm long, transverse veins present, glabrous to scabrous, only one margin hairy. Ligule

0.5—0.8 mm long, basal 0.4 mm membranous, hairs at tip 0.1—0.4 mm long. Blade 13—20

cm long, 4—5 mm wide, (sparsely) hairy at least at upperside, base cuneate. Inflorescence

open, central rachis 15—20.cm long, branches erecto-patent; longest branch 10—13 cm long,

naked at base. Pedicel 1—3.5 mm long, scaberulous. Spikelet 2.5—3.1 mm long, 0.9—1 mm

wide, acute, glumes without a distinct internode, |* floret male; lowest bract c. 0.2 times as

long as spikelet; lower glume 0.6—0.7 mm long, collar-shaped, 0 veins, glabrous; upper

glume 2.9—3 mm long, glabrous; 1“ lemma 2.6—2.7 mm long, acute; 1“ palea 2.3—2.5 mm

long; 2"! lemma 2—2.2 mm long, smooth. Anthers 3, 1—1.4 mm long.

Probably not native; pantropical. First record 1880s. Common.

In lawns on damp sandy soil, sunny to slightly shaded, and surViving periodic

inundation.

According to Veldkamp (1996a), it is probably introduced 1n Malesia. He reports slightly

longer anthers (1.3—1.5 mm long).

It is usually a seashore plant, but it is also found inland on sandy places (Ridley 1907,

1925). It is used as a sandbinder and for lawns and turfs (Veldkamp 1996a). In America it is

known as Torpedo Grass.

Differences with Eriochloa procera and Urochloa subquadripara are mentioned under

the respective species.

Panicum sarmentosum Roxb. — Fig. 104.

Scrambling Panic Grass, Rumput Janggut Ali

Ridley 1907: 140; 1925: 227. Gilliland 1971: 139. Veldkamp 1996a: 203. Keng et al.

1998: 176. Veldkamp 1999b: 235.

Culms scrambling, 70—800 cm long; internodes solid; nodes glabrous or hairy. Sheath 9—

10 cm long, transverse veins absent, (densely) hairy, margins hairy. Ligule 0.6—0.8 mm

long, basal part membranous and 0.3—0.4 mm long, hairs at tip 0.3—0.4 mm long. Blade

18—33 cm long, 12—22 mm wide, hairy, base cuneate. Inflorescence open, central rachis

16—43 cm long, branches patent; longest branch 6—24 cm long, with spikelets almost from

the base. Pedicel 0.2—1 mm long, glabrous. Spikelet 1.9—2.4 mm long, 0.7—0.9 mm wide,

obtuse, glumes without a distinct internode, 1* floret sterile or male; lowest bract 0.5 times as

long as spikelet; lower glume |.1—1.2 mm long, ovate, 3 veins, glabrous; upper glume |.9—

2.4 mm long, 5 veins, glabrous; 1‘ lemma 1.8—2.1 mm long, acute; 1‘ palea 1.3 mm long; 2"

lemma |.5—1.8 mm long, smooth. Anthers 3, | mm long.

Native; India to S China and S to Australia. First record 1889. Rare.

Forest margins and scrub.

Veldkamp (1996a) reports that the 1*' floret is sterile.

Panicum trichocladum Hack. ex K.Schum. in Engl. — Fig. 105.

Veldkamp 1999b: 235. Duistermaat 2004: 35.

Culms scrambling, 50—250 cm long; internodes hollow; nodes glabrous. Sheath 4—6 cm

long, margin at tip short-hairy. Ligule membranous, 0.4—0.5 mm long, tip ciliolate. Blade

5—8 cm long, 5—8 mm wide, glaucous, hairy, base cuneate to rounded, pseudopetiole 1—2

mm long. Inflorescence open, central rachis 7—9 cm long, branches patent; longest branch c.

104 Supplement of Gardens’ Bulletin Singapore 57 (2005)

4.5 cm long, naked at basal 1.5—2 cm, each branchlet with 2 spikelets only. Pedicel c. 2.5

mm long, scaberulous. Spikelet 2.1—2.4 mm long, 0.9—1 mm wide, obtuse to acute, glumes

without a distinct internode, 1* floret male; lowest bract 0.15—0.20 times as long as spikelet;

lower glume 0.3—0.5 mm long, ovate, 0 veins, glabrous; upper glume 1.9—2.2 mm long, 5

veins, glabrous; 1 lemma 2—2.2 mm long, obtuse; 1* palea 2—2.1 mm long; 2"? lemma

1.8—2 mm long, smooth. Anthers 3, 1.4 mm long.

Not native; Africa: Zaire to Ethiopia, introduced in Surinam and Singapore. First record

1994. Rare.

In shaded places.

Paspalum L.

De Koning and Sosef 1985: 279—318.

Nodes glabrous. Ligule membranous, glabrous. Young leaf blade inrolled. Inflorescence

panicle; branches 2—11, 0.7—3 mm wide, distinctly winged, simple. Pedicel flattened to

more or less rounded. Spikelets single or paired, alternate in 2 rows on one side of branch,

dorsiventrally flattened, 2-flowered with 1* floret sterile and 2™ bisexual, unawned; the two

lowest bracts as long as spikelet; lower glume and 1* palea absent; upper glume and 1* lemma

outer veins marginal, thin herbaceous; 2" lemma and palea indurated, palea almost as long as

lemma.

Paspalum dilatatum, earlier mentioned for Singapore (Gilliland 1971: 183, Keng et

al. 1998: 176), is not included. The only Singapore collection of this American species

originated from material that was cultivated in the Botanic Gardens. As in P. conjugatum, it

has an upper glume with a fringe of hairs, but the inflorescence has 3—5(—9) branches and

the spikelets are paired and c. 3.3 mm long.

The key follows that of De Koning and Sosef (1985).

la. Upper glume with a marginal fringe of relatively long, white hairs fringing

the spikelet. P. conjugatum

1b. Upper glume without such a fringe. 2

2a. Spikelets paired, at least in the middle of the branch, one sometimes abortive

and reduced to minute glumes. 3

2b. Spikelets solitary, without an abortive 2" spikelet. 5

3a. Spikelets oblong to ovate or obovate, upper glume and 1“ lemma with a

conspicuous dark centre and paler margin; 2" lemma and palea dark

brown, shiny. P. plicatulum

3b. Spikelets broadly ovate to obovate; upper glume and 1* lemma of one

colour; 2™ lemma and palea pale to brown, dull. 4

4a. Branches (4—)6—11, 2—3 mm wide. Both spikelets of a pair usually

well-developed in the middle of the branch. Upper glume pubescent,

very rarely glabrous. P. longifolium

Figure 103. Panicum repens L.

Figure 104. Panicum sarmentosum Roxb.

Figure 105. Panicum trichocladum Hack. ex K.Schum. in Engl.

Figure 106. Paspalum conjugatum P.J.Bergius.

Figure 107. Paspalum longifolium Roxb.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 105

106 Supplement of Gardens’ Bulletin Singapore 57 (2005)

4b. Branches 2—4(—6), 1—1.5(—1.9) mm wide. One of the spikelets of a

pair usually reduced. Upper glume glabrous. P. orbiculare

5a. Culms with stolons. Spikelets oblong, acute. P. vaginatum

5b. Culms tufted, stolons absent. Spikelets ovate or obovate to suborbicular,

blunt. 6

6a. Spikelets usually broadly obovate, (1.25—-)1.3—1.8 times as long as wide,

yellow-brown to brown; upper glume and 1* lemma 3- or 5-veined,

smooth, veins distinctly darker than between the veins; 2" floret yellow-

brown to brown in fruit. P. orbiculare

6b. Spikelets usually suborbicular, 1.05—1.25(—1.5) times as long as wide,

brown to dark brown; upper glume 5- or 7-veined, somewhat rumpled;

1‘ lemma 5—9-veined, veins the same colour as between the veins; 2"

floret dark brown in fruit. P. scrobiculatum var. bispicatum

Paspalum conjugatum P.J.Bergius — Fig. 106, Plate 22.

Buffalo Grass, Rumput Kerbau

Ridley 1907: 124; 1925: 218. Gilliland 1971: 180. De Koning and Sosef 1985: 290.

Keng et al. 1998: 176.

Culms with long stolons, erect part 15—60 cm long. Sheath 1.5—13 cm long, narrowing

towards tip, more or less flattened, midrib raised, glabrous, margin with up to 1 mm-long

white hairs, external ligule short-hairy. Ligule 0.2—0.7 mm long. Blade (1—)2—9(—20) cm

long, (I—)3—9 mm wide, sparsely hairy, margin hairy, base cuneate. Inflorescence central

rachis 3—10 mm long, branches 2; longest branch 7—16.5 cm long, 0.7—0.8 mm wide, with

few long hairs at base only. Pedicel 0.5—0.8 mm long, glabrous. Spikelets single, 1.5—1.7

mm long, 1—1.2 mm wide, ovate; upper glume 1.5—1.7 mm long, 2 concolorous veins

(midvein suppressed), smooth, marginal hairs 1.5—2 mm long and white; 1“ lemma 2 veins,

glabrous; 2™ lemma |.4—1.5 mm long, pale green.

Not native; America, introduced in Africa, Asia and Australia. First record 1884.

Common.

In unshaded to shaded, dry to moist places as lawns, roadsides, open wasteland, etc.

De Koning and Sosef (1985) report a much longer ligule (0.7—1.6 mm long). Axonopus

compressus has hairy nodes, densely fringed ligules and a less dense external ligule.

Paspalum longifolium Roxb. — Fig. 107.

Ridley 1925: 217. Gilliland 1971: 182. De Koning and Sosef 1985: 297. Keng et al.

1998: 176, .

P. platycoleum Ridl. 1925: 217 (excluding Singapore).

Culms tufted, 5S0—125 cm long. Sheath S—24 cm long, more or less flattened, midrib raised,

glabrous to densely long-hairy, margins glabrous or hairy in upper half. Ligule 0.8—2.5 mm

long. Blade (3—)17—36 cm long, (3—)4—7 mm wide, base cuneate and hairy at upperside.

Inflorescence central rachis 5—10 cm long, branches (4—)6—11; longest branch 6.5—8 cm

long, 2—3 mm wide, margins at least at base hairy. Pedicel 0.5—1.5 mm long, glabrous.

Spikelets in pairs, 2—2.4 mm long, 1.3—1.7 mm wide, broadly (ob-)ovate; upper glume 2—

2.4 mm long, 3 concolorous veins, smooth, (sparsely) hairy; 1“ lemma 3 veins, (sparsely)

hairy; 2" lemma 1.6—2 mm long, pale brown.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 107

Native; India to Vietnam, and SE to the Pacific and N. Australia. First record 1880s.

Rare.

In moist to wet places or in up to 60 cm-deep water, sometimes floating on open water.

Paspalum orbiculare G.Forst. — Fig. 108.

Ridley 1925: 217. Gilliland 1971: 184. De Koning and Sosef 1985: 299. Keng et al.

19938: 176.

Culms single or tufted, erect to geniculate, 9—80 cm long. Sheath up to 20 cm long, more or

less flattened, midrib raised, glabrous. Ligule 0.7—1 mm long. Blade 3—13(—50) cm long,

2.5—5.5 mm wide, glabrous, base somewhat rounded and glabrous to long-hairy. Inflorescence

central rachis 1—4 cm long, branches 2—6; longest branch 2—6 cm long, 1—1.2 mm wide,

margins scaberulous, base hairy. Pedicel 0.5—0.6 mm long, scaberulous. Spikelets single or

in pairs with the inner often reduced to tiny glumes, 1.9—2.1 mm long, 1.6—1.9 mm wide,

(broadly) obovate (to suborbicular); upper glume 1.9—2.1 mm long, 3 or 5 dark-coloured

veins, smooth, glabrous; 1‘ lemma 5—9 veins, glabrous; 2" lemma 1.5—1.9 mm long, yellow-

brown to brown in fruit. ‘i

Native; India to Taiwan and Hong Kong, and S to Polynesia and N. Australia. First

record 1932. Common.

In dry to wet unshaded places or in water up to 25 cm deep.

It is difficult to distinguish this species from P. scrobiculatum var. bispicatum. The

way the upper glume fits around the spikelet (smooth vs rumpled) seems to be the easiest

character to use in the field.

Paspalum plicatulum Michx. — Fig. 109.

De Koning and Sosef 1985: 313. Duistermaat 2004: 37.

Culms tufted, 30—150 cm long. Sheath with midrib raised, hairy in marginal area or upper

half. Ligule 1.5 mm long. Blade 6—18 cm long, c. 10 mm wide, hairy on underside.

Inflorescence central rachis 3—15 cm long, branches c. 10; longest branch c. 9 cm long,

0.9—1 mm wide, sparsely hairy at base. Pedicel 0.5 mm long, glabrous. Spikelets in pairs, c.

2.5 mm long, c. 1.6 mm wide, obovate; upper glume 2.5 mm long, 5 concolorous veins,

smooth, glabrous, with dark red-coloured spot in the centre; 1‘! lemma as upper glume; 2"

lemma 2 mm long, dark brown.

Not native; (sub-)tropical America, introduced in temperate Asia and Australasia. Single

record 2003 from Pulau Sakijang. Rare.

The species is cultivated for its high palatability to cattle in (sub-)tropical America,

the Chinese province of Gansu and in Papua New Guinea.

Paspalum scrobiculatum L. var. bispicatum Hack. — Fig. 110, Plate 23.

Kodo Millet

De Koning and Sosef 1985: 305. Keng et al. 1998: 177.

P. cartilagineum J.Presl: Gilliland 1971: 184.

P. commersonii Lam.: Ridley 1925: 218.

P. scrobiculatum L.: Ridley 1907: 124. Gilliland 1971: 185. -

Culms tufted, 40—60 cm long. Sheath 8—13 cm long, more or less flattened, midrib raised,

glabrous. Ligule 0.5—1 mm long. Blade 8—40 cm long, 5—7 mm wide, glabrous, margin

glabrous or sparsely hairy, base cuneate and glabrous or hairy. Inflorescence central rachis

1.5—3 cm long, branches 2—6; longest branch 5—9.5 cm long, 1.5—2.5 mm wide, glabrous

or hairy at base. Pedicel 0.5 mm long, glabrous. Spikelets single, 2—2.3 mm long, 1.5—1.9

mm wide, suborbicular to broadly (ob-)ovate; upper glume 2—2.3 mm long, 5 concolorous

108 Supplement of Gardens’ Bulletin Singapore 57 (2005)

veins, rumpled, glabrous; 1“ lemma 5 or 7 veins, glabrous; 2"* lemma 1.7—2 mm long, dark

brown in fruit.

Native; tropical Africa, Asia, Pacific and Australia. First record 1889. Common.

In unshaded to shaded, dry to moist places, roadsides, wastelands; seems less common

in urban environments.

The difference with P. orbiculare is mentioned there.

Paspalum vaginatum Sw. — Fig. 111, Plate 24.

Saltwater Paspalum, Rumput Dawai

Gilliland 1971: 182. De Koning and Sosef 1985: 309. Keng et al. 1998: 177.

P. distichum Ridl.: 1907: 124; 1925: 218.

Culms much branched at base, with stolons up to 300 cm long, erect shoots 25—80 cm long.

Sheath 2—4.5 cm long, rounded to somewhat flattened, midrib not or slightly raised, glabrous,

margin at tip hairy. Ligule 1 mm long. Blade (2—)8—15 cm long, 1.5—3.5 mm wide, glabrous,

base cuneate and much narrower than sheath. Inflorescence central rachis absent, branches 2:

longest branch 2.5—5.5 cm long, |—1.6 mm wide, glabrous, empty at basal 4—6 mm. Pedicel

0.5—0.7 mm long, glabrous to scaberulous. Spikelets single, 2.7—4.4 mm long, 1.2—1.5

mm wide, oblong; upper glume 2.6—4.4 mm long, 5 or 6 concolorous veins, smooth, glabrous;

I“ lemma 5 or 7 veins, glabrous; 2™ lemma 2.2—3 mm long, pale green.

Native; pan(sub)tropical. First record 1890. Rather common.

Halophilous (i.e. growing in saline soil), in open habitats as beaches, mangrove margins,

tidal pools; can withstand flooding.

Pennisetum Rich.

Culms usually tufted, erect to ascending (or with stolons in P. polystachion); basal internodes

10—15 mm diameter; nodes glabrous or appressed hairy. Sheath rounded. Ligule membranous

at base, with hairs at tip. Young leaf blade inrolled. Blade at base (somewhat) rounded, margins

scaberulous, midvein at upperside at least at base prominent and white. Inflorescence dense

spike. Spikelets in groups of 1—3, each group surrounded by involucre of bristles falling off

together with the spikelet(s); involucral bristles (slightly) longer than the spikelets, fused

only at the very base, the longest in basal half plumose. Spikelet not flattened, 2-flowered

with 1‘ floret sterile or male and 2™ bisexual, unawned; glumes and 1“ palea hyaline (or

absent); lemmas and 2™ palea (thin) herbaceous; 2" floret glabrous, palea slightly shorter

than lemma. Anthers 3.

Pennisetum glaucum, P. macrostachyum and P. setaceum have been grown in the

Botanic Gardens and in the garden of the University of Singapore. They are not included

here.

Differences with Cenchrus and Setaria are mentioned under Cenchrus.

la. Inflorescences 2—4 per culm; central rachis with ridges or winged below

the spikelets, glabrous. Spikelets single, 3.5—5 mm long. Anthers 1.1—

1.4 mm long, glabrous. P. polystachion

Figure 108. Paspalum orbiculare G.Forst.

Figure 109. Paspalum plicatulum Michx.

Figure 110. Paspalum scrobiculatum L. var. bispicatum Hack.

Figure 111. Paspalum vaginatum Sw.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 109

Fig. 108

110 Supplement of Gardens’ Bulletin Singapore 57 (2005)

1b. Inflorescences | (or 2) per culm; central rachis smooth to inconspicuously

ribbed, densely hairy. Spikelets in groups of 1—3, 5—5.8 mm long.

Anthers 2.3—3 mm long, ciliolate at tip. P. purpureum

Pennisetum polystachion (L.) Schult. — Fig. 112, Plate 25.

Feather Pennisetum

Holm et al. 1977: 367—371. Veldkamp 1997a: 211. Keng et al. 1998: 177.

Culm 98—200 cm long, stolons up to 1.5 m long sometimes present. Sheath 9—15(—19) cm

long, glabrous to hairy, margins glabrous to hairy in upper half, external ligule an herbaceous

17—30 cm long, (2—)5—7 mm wide, hairy. Inflorescences 2—4 per culm; central rachis 9—

21 cm long, with prominent ridges to decurrently winged below the spikelets, glabrous.

Involucral bristles up to 24 mm long, hairs up to 0.5 mm long. Pedicel absent. Spikelets

single, 3.5—5 mm long, 0.7 mm wide; lowest bract as long as and 2" bract c. 0.6 times as

long as spikelet; lower glume absent; upper glume 3.5—5 mm long, 5 veins, glabrous to

short-hairy, acute; 1** lemma 2.8—3.2 mm long, tip 3-lobed; 1‘ palea absent; 2"? lemma 2—

2.3 mm long, tip truncate and ciliate. Anthers 1.1—1.4 mm long, glabrous.

Not native; probably native to Africa, now pantropical. First record 1929 from the

Botanic Garden, 1959 in the wild. Rather common.

In open or partly shaded waste land, on well-drained soil.

It was introduced as a fodder crop, but today in many countries around the world it is

regarded as a weed (see Veldkamp 1997a).

Pennisetum purpureum Schumach. — Fig. 113, Plate 27.

Elephant Grass, Napier Grass

Holm et al. 1977: 367—371. Gilliland 1971: 162. Keng et al. 1998: 177.

Culm up to 250 cm long. Sheath 9—16 cm long, glabrous, external ligule absent. Ligule basal

part 0.3—0.5 mm long, hairs at tip 3.2—4 mm long. Blade (16—)27—45 cm long, (7—)15—

17 mm wide, underside glabrous, upperside (sparsely) hairy. Inflorescence 1 (or 2) per culm;

central rachis 8—27 cm long, smooth to inconspicuously ribbed, densely hairy. Pedicel 0.3—

2.3 mm long, scaberulous. Spikelets in groups of 1—3, 5—5.8 mm long, 0.9—1.7 mm wide;

involucral bristles up to 19 mm long, hairs up to 1.2 mm long; either lowest bract c. 0.1 and

2™ (0.2—0.3 times as long as spikelet, or lowest 0.2—0.3 times as long as and 2" as long as

spikelet; lower glume absent or 0.4—0.5 mm long; upper glume 1.2—1.8 mm long, 0 veins,

glabrous, obtuse; 1‘‘ lemma 4.1—5 mm long, acute; 1“ palea 4.1 mm long or absent; 2"

lemma 4.7—5.5 mm long, acuminate. Anthers 2.3—3 mm long, ciliolate at tip.

Not native; Africa, now pantropical. First record 1931. Rather common.

In open or partly shaded waste land, generally on clayey soil.

Holttum (Holttum s.n., 1931, SING) records that this species was used as food for

Chinese carp and also as fodder for dairy cows. .

Perotis Aiton

Veldkamp and Van Steenbergen 1992: 609—614.

Figure 112. Pennisetum polystachion (L.) Schult., spikelet surrounded by involucre of

bristles. 7

Figure 113. Pennisetum purpureum Schumach., group of three spikelets surrounded by

involucre of bristles.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) Lid

Fig. 112

112 Supplement of Gardens’ Bulletin Singapore 57 (2005)

Perotis indica (L.) Kuntze — Fig. 114.

Cat’s Tail Grass, Rumput Ekor Kucing

Gilliland 1971: 113. Veldkamp and Van Steenbergen 1992: 610. Keng et al. 1998:

178.

P. latifolia Aiton: Ridley 1907: 149; 1925: 243.

Culms tufted, geniculate to decumbent, 20—40 cm long; nodes glabrous. Sheath 1—S cm

long, rounded, glabrous. Ligule membranous, 0.3 mm long, tip ciliolate. Young leaf blade

inrolled. Blade 1—3.5 cm long, 3—6 mm wide, glabrous, base cordate to amplexicaul and

pectinate. Inflorescence raceme, rachis 7—15 cm long, 0.1—0.2 mm wide, scaberulous to

bristle-hairy. Pedicel 0.3 mm long, bristle-hairy, persistent. Spikelets single, 1.7—-2 mm long,

0.4 mm wide, patent, easily falling off from base upwards leaving pedicel attached to rachis,

laterally flattened, 1-flowered, awned; callus 0.3 mm long, hispid, falling off with spikelet;

lowest bract as long as and 2" 0.9 times as long as spikelet; lower glume 1.7—2 mm long,

hyaline to thin herbaceous, | vein, short hairy, obtuse, awn from tip 7—10 mm long and

antrorsely scaberulous; upper glume |.5—1.9 mm long, thin herbaceous, | vein, glabrous,

obtuse to acute, awn from tip 6.5—9 mm long; lemma 0.5—1 mm long, hyaline; palea 0O—0.7

mm long, hyaline.

Native; India to E China, throughout Malesia to Australia. First record 1880s. Rare.

On sandy soil near the coast, under Casuarina species, in coconut plantations, dry

grasslands.

Phragmites Adans.

Phragmites vallatoria (Pluk. ex L.) Veldk. — Fig. 115.

Common Reed, Rumput Gedabung

Veldkamp 1992: 232. Keng et al. 1998: 178.

P. communis auct., non Trin.: Ridley 1925: 240 (excluding Singapore).

P. karka (Retz.) Steud.: Ridley 1907: 175 (excluding Singapore). Gilliland 1971: 49.

Culms up to 300 cm long; internodes hollow, up to 15 mm diameter; nodes glabrous. Sheath

10—12 cm long, rounded, glabrous. Ligule of younger leaves row of hairs and 4—6 mm

long, of flag leaf with basal part membranous and 0.1—0.3 mm long and hairs at tip 0.3—0.4

mm long; just behind ligule few up to | mm long hairs on blade. Young leaf blade inrolled.

Blade c. 32 cm long, c. 10 mm wide, glabrous, base rounded. Inflorescence panicle, silky

hairy, central rachis c. 26 cm long, branches many; longest branch c. 15 cm long, thin, branched.

Pedicel 3—4 mm long, thinner than branch, scaberulous. Spikelets single, at least 11 mm

long and 5 mm wide, 4—more-flowered with 1* floret sterile and others bisexual, unawned;

rachilla between the glumes and the 1‘ lemma 0.5 mm long and glabrous and persistent,

between the upper florets 0.5 mm long and with up to 5 mm-long hairs and articulated; the

two lowest bracts up to 0.3 times as long as spikelet; lower glume 3.3 mm long, enveloping,

thin herbaceous, glabrous, 3 veins; upper glume as lower, 3.6 mm long; 1“ lemma 7 mm long,

herbaceous, 3 veins, glabrous, acute; 1“! palea 3 mm long, thin herbaceous; 2" lemma as 1",

8.5 mm long; 2" palea as 1‘; 3" and higher florets as 2", but gradually smaller in size.

Native; Asia and Australia. Single record 1880s around Freshwater Isle (unknown

locality). Extinct (1880s).

Figure 114. Perotis indica (L.) Kuntze.

Figure 115. Phragmites vallatoria (Pluk. ex L.) Veldk., Spueerets with rachilla articulated

above the first floret.

Figure 116. Pogonatherum crinitum (Thunb. in Murray) Kunth.

Figure 117. Polytrias indica (Houtt.) Veldk.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 113

Fig. 114 y :

114 Supplement of Gardens’ Bulletin Singapore 57 (2005)

In wet places.

Differences with the other species with silky-hairy panicles are mentioned under

Imperata.

Pogonatherum P.Beauv.

Pogonatherum crinitum (Thunb. in Murray) Kunth — Fig. 116, Plate 26.

Bamboo Grass, Rumput Sembor Batu

Keng et al. 1998: 178.

P. paniceum (Lam.) Hack.: Gilliland 1971: 251.

P. saccharoideum P.Beauv. var. monandra Hack.: Ridley 1907: 154; 1925: 195 (both excluding

Singapore).

Culms tufted, sometimes woody at base, stolons absent, 12—-50 cm long; nodes hairy. Sheath

2—3(—5) cm long, rounded, glabrous, one margin hairy. Ligule membranous, 0.4—0.5 mm

long, tip ciliolate. Young leaf blade inrolled. Blade 3.5—5.5 cm long, 1—3 mm wide, glabrous,

margin smooth to scaberulous, base cuneate and long-hairy. Inflorescence spike-like raceme;

central rachis 1.5—3.5 cm long, hairy. Pedicel 0.7 mm long, flattened, hairy. Spikelets in

pairs of 1 sessile and 1 pedicelled. Sessile spikelet 1.5—2 mm long, 0.4—0.5 mm wide,

laterally flattened, 1- or 2-flowered with 1“ floret sterile or absent and 2" bisexual, awned;

callus very short, with up to | mm-long hairs; lowest bract c. 0.9 times as long as and 2" as

long as spikelet; lower glume |.1—1.5 mm long, thin herbaceous, glabrous, tip truncate and

ciliolate; upper glume 1.5—1.7 mm long, thin herbaceous, glabrous or hairy in upper half,

awn from tip 12—17 mm long and golden brown; 1|* lemma and palea absent or 1—1.4 mm

long and hyaline; 2" lemma 1—1.1 mm long, hyaline, glabrous, tip bifid and ciliolate, awn

from notch 23 mm long and golden brown; 2" palea 0.4—1 mm long, hyaline. Pedicelled

spikelet as sessile, 1.4 mm long, 1-flowered.

Native; India to Japan, S to Peninsular Malaysia and Singapore. First record 1950.

Rare.

On shaded to sunny granite outcrops, rocky places, and unplastered brick walls.

Contrary to what the English name suggests this is not a bamboo. It is much smaller

than bamboos and not woody. Moreover, real bamboos have two types of leaves (culm and

foliage), and clusters of spikelets scattered along a rachis (nodular inflorescence). It is cultivated

as an ornamental.

Polytrias Hack.

Polytrias indica (Houtt.) Veldk. — Fig. 117.

Brown-top Grass

Veldkamp 1991: 180. Keng ef al. 1998: 179.

Eulalia praemorsa (Nees) Stapf ex Rid]. 1925: 197.

P. amaura (Biise) Kuntze: Gilliland 1971: 244.

P. praemorsa Hack.: Ridley 1907: 164.

Culms geniculate or with stolons, 25—40 cm long; nodes hairy, upper ones glabrescent. Sheath

1.5—5.5 cm long, rounded, glabrous or sparsely hairy at tip, margins glabrous or hairy at

base. Ligule membranous, 0.5 mm long, tip ciliolate. Young leaf blade inrolled. Blade 2.5—

4 cm long, 2—3.5 mm wide, hairy, base rounded and with long hairs. Inflorescence spike-like

raceme; rachis 4—6.5 cm long, rounded, with a longitudinal row of hairs between the spikelets.

Pedicel 2 mm long, thinner than rachis, flattened, hairy. Spikelets in threes, both the lateral

sessile and the middle pedicelled but otherwise similar, 3.5—3.7 mm long, 0.8—0.9 mm

wide, dorsiventrally flattened, 1-flowered (but see note), awned; lowest bract c. 0.9 times as

long as and 2" as long as spikelet; lower glume 3.4—3.5 mm long, herbaceous, 4 or 5 veins,

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 115

hairy, tip truncate and with protruding marginal veins; upper glume 3.5—3.7 mm long, thin

herbaceous, 3 veins, hairy in upper half, obtuse; lemma |.2—1.5 mm long, hyaline, tip bifid

and sparsely hairy, awn from notch 10—13 mm long and geniculate with lower part twisted

and upper part antrorsely scaberulous; palea 0.6 mm long, hyaline.

Probably not native; possibly native to Java, now pantropical. First record 1889. Extinct

(1963).

In dry, sunny places, e.g. lawns, waste ground.

Technically speaking, the spikelets are 2-flowered, with the 1“ lemma and palea lacking.

The lemma that is present is positioned next to the upper glume; if the spikelet was truly 1-

flowered, the lemma would be positioned opposite the upper glume (adjacent to the lower

glume). .

It was probably introduced as a lawn grass. Veldkamp (1991) reports that it occurs

especially in areas with a strong East monsoon but that in moister areas it is outcompeted by

Axonopus.

Rottboellia L.f.

Veldkamp et al. 1986: 281—307.

The genus is said to be different from Mnesithea in having a male |* floret (Veldkamp

et al. 1986).

Rottboellia cochinchinensis (Lour.) Clayton — Fig. 118.

Veldkamp et al. 1986: 305. Duistermaat 2004: 38.

R. exaltata L.f.: Holm et al. 1977: 139.

Culms tufted, with short rhizomes and thick roots, c. 80 cm long; nodes glabrous. Sheath 6—

7 cm long, rounded, spiny-hairy, margin hairy. Ligule membranous, 1.5 mm long, tip ciliolate.

Young leaf blade inrolled. Blade 21—45 cm long, 11 mm wide, spiny-hairy at upperside,

margin scaberulous, base rounded. Inflorescence | per sheath, several from 1 culm, spike-

like raceme, 5—7 cm long; rachis articulating, glabrous, simple; internodes at articulation

with a central knob, basal part flattened and 2—3.8 mm wide, upper part rounded and 2—4.5

mm wide. Pedicel 3.5 mm long, glabrous, flattened, one margin fused with rachis. Spikelets

in pairs of | sessile and 1 pedicelled, unawned. Sessile spikelet 4.5—5 mm long, 1.7—4 mm

wide, not flattened, 2-flowered with 1* floret male and 2™ bisexual; the two lowest bracts as

long as spikelet; lower glume 4—5 mm long, flattened, indurated, glabrous, minutely granular,

submarginal keels smooth and in upper part winged or not, tip obtuse or minutely bifid; upper

glume 4.5—5 mm long, boat-shaped, indurated, glabrous, upper half minutely granular, tip

acute and keeled; 1‘ lemma 3.5—4.5 mm long, thin herbaceous; 1*' palea 3—4.3 mm long; 2"

lemma c. 3 mm long, hyaline; 2"! palea absent or c. | mm long. Pedicelled spikelet reduced to

2 flattened herbaceous glumes enclosing 0—2 hyaline bracts.

Not native; Paleotropics. First record 1995. Rare.

Found on reclaimed land.

Material was undoubtedly introduced with sand brought in for land reclamation. It is

likely to spread further as it is one of the world’s worst weeds (Holm et al. 1977).

The internodes of the inflorescence are corky, allowing them to float on water after

articulation. Dispersal by ants, which are attracted by the central knob (or elaiosome) that

contains oily substances, is reported by Veldkamp et al. (1986: 295).

The species is probably more variable than previously thought (Veldkamp et al. 1986,

Duistermaat 2004). Mnesithea glandulosa is different in having free pedicels and lower glumes

with 2—7 gland-like warts in the lower half of the keels.

116 Supplement of Gardens’ Bulletin Singapore 57 (2005)

Saccharum L.

Whalen 1991: 109—125. Hodkinson et al. 2002: 381—392.

Ripidium Trin., non Bernh.

Culms with rhizomes and tufted, 100—500 cm long; internodes solid, 5—45 mm diameter,

just above the node with 1 or more rows of root-eyes; nodes glabrous. Sheath 20—40 cm

long, rounded, margins glabrous or at throat hairy, shouldered, auricles triangular and glabrous.

Ligule membranous, 0.5—3 mm long, tip ciliolate. Young leaf blade inrolled. Blade 50—200

cm long, glabrous, margin serrulate, base cuneate. Inflorescence panicle, silky hairy, central

rachis present, branches many and articulated; longest branch widest part 3 mm wide, branched.

Spikelets in pairs of 1 sessile (falling off with rachis internode and pedicel) and | pedicelled,

otherwise similar, not flattened, 2-flowered, unawned; the two lowest bracts 0.9—1 times as

long as spikelet; both lemmas hyaline, 1‘ lemma almost as long as glumes, 2"* lemma absent

to 0.75 times as long as glumes; 1“ palea absent; 2" palea absent to as long as 2"? lemma,

hyaline.

Ripidium probably has to be split off (Hodkinson et al. 2002: 391), which would here

only include Saccharum arundinaceum.

Almost all modern-day commercial sugarcanes are complex hybrids of S. officinarum

with S. spontaneum and/or S. sinense (Whalen 1991). For this reason, S. sinense is included in

the key but hybrids are not included. Plants rarely seem to flower in Singapore. Unfortunately,

most can only be identified when flowering.

Differences with the other species with silky-hairy panicles are mentioned under

Imperata.

The key (leads 2—4) follows Whalen (1991).

la. Root-eyes in | row above node (Plate 28). Sheath glabrous. Callus glabrous.

Lower glume with c. 3 mm long silky hairs. S. arundinaceum

1b. Root-eyes in 2—9 rows above node (Plate 29). Sheath hairy or glabrous.

Callus with up to 12 mm long silky hairs. Lower glume glabrous or with

ciliate margin or tip. 2

2a. Sheath hairy. Peduncle and central rachis glabrous or sparsely hairy, hairs

up to 0.5 mm long. 2" lemma absent or much reduced. Culms 20—45

mm diameter, green to yellow, red or violet. S. officinarum

2b. Sheath hairy or glabrous. Peduncle and central rachis densely hairy, hairs

c. 2.5 mm long. Both lemmas present. Culms 5—30 mm diameter. 3

3a. Rhizome short (plant bunch forming). Culms 15—30 mm diameter, with

leaf sheaths falling off. Sheath hairy or glabrous. Blade 25—60 mm

wide. Lodicules glabrous. S. sinense

3b. Rhizome elongate. Culms 5—15 mm diameter, with persistent leaf sheaths.

Sheath glabrous. Blade 3—30 mm wide. Lodicules ciliate.

S. spontaneum

Figure 118. Rottboellia cochinchinensis (Lour.) Clayton, part of the rachis with a pair of

spikelets.

Figure 119. Saccharum arundinaceum Retz.

Figure 120. Saccharum spontaneum L.

117

Field Guide to the Grasses of Singapore (Excluding the Bamboos)

118 Supplement of Gardens’ Bulletin Singapore 57 (2005)

Saccharum arundinaceum Retz. — Fig. 119, Plate 28.

Tebu Salah

Ridley 1907: 153; 1925: 193. Gilliland 1971: 226. Keng et al. 1998: 179.

Ripidium arundinaceum (Retz.) Grass]

Culms with short rhizomes, erect to ascending; basal internodes c. 10 mm diameter, root-eyes

in | row. Sheath more or less persistent, glabrous. Ligule 1 mm long and abaxially with row

of 5 mm long hairs. Blade 10—42 mm wide. Inflorescence 6—30 cm wide, peduncle and

central rachis glabrous, central rachis 29—90 cm long; longest branch 9—45 cm long;

internodes ultimate branches 4—6.5 mm long, with up to 10 mm long silky hairs at base only.

Pedicel 2.5—3 mm long, sparsely hairy, hairs up to 2.5 mm long. Spikelet 2.3—4.2 mm long,

c. 1.5 mm wide; callus 0.4 mm long, glabrous; lower glume 2.8—4.2 mm long, herbaceous,

hairy, hairs c. 3 mm long; upper glume up to 3.9 mm long, hyaline, long-hairy; 1“ lemma with

hairs up to 1 mm long; 2" lemma up to 3.1 mm long. Lodicules ciliate.

Native; India to S. China, Malesia to New Guinea. First record 1900. Rare.

On river banks.

Saccharum officinarum L. — Fig. 121, Plate 29.

Sugar Cane, Tebu

Gilliland 1971: 227. Whalen 1991: 118. Keng et al. 1998: 179.

Culms with short rhizomes, erect; basal internodes 20—45 mm diameter, root-eyes in 2—9

rows. Sheath deciduous, hairy. Ligule 0.5—3 mm long. Blade 40—60 mm wide. Inflorescence

14—17 cm wide, peduncle and central rachis sparsely hairy with hairs appressed and c. 0.5

mm long, central rachis 40—60 cm long; longest branch c. 15 cm long; internodes ultimate

branches 4—6.5 mm long, with up to 10 mm-long silky hairs at base only. Pedicel 2.5—3 mm

long, ciliate at tip. Spikelet 3.2—3.6 mm long, 0.8—0.9 mm wide; callus 0.4 mm long, hairy,

hairs up to 2 mm long; lower glume 3.2 mm long, herbaceous, glabrous or hairy at tip; upper

glume herbaceous, glabrous, margins hairy, hairs up to 0.9 mm long; 1“ lemma margins hairy;

2™ Jemma absent or much reduced. Lodicules glabrous.

Not native; Melanesia, now cultivated throughout the tropics. First record 1921 (Botanic

Gardens). Cultivated.

In gardens and waste places where rhizomes were dumped.

This species is a hybrid between Miscanthus floridulus and S. robustum. It is only

known from cultivation, where it is grown for sugar and juice and is one of the few economically

important crops from Papuasia. In Singapore it rarely flowers.

Saccharum sinense Roxb.

India Cane

Whalen 1991: 113.

This species is used for modern cultivar hybrids (see note under Saccharum).

It is present in the Botanic Gardens, but the basal internodes are only 12 mm diameter

and the leaves are just 13 mm wide. It is probably one of the clones resembling S. spontaneum

(Whalen 1991: 115).

Saccharum spontaneum L. — Fig. 120.

Wild Cane

Gilliland 1971: 224. Whalen 1991: 112. Keng et al. 1998: 180.

Culms with long rhizomes, erect; basal internodes 5—15 mm diameter, root-eyes in 2 rows.

Sheath persistent, glabrous. Ligule c. 3 mm long. Blade 3—30 mm wide. Inflorescence c. 5

cm wide, peduncle and central rachis densely hairy with hairs appressed and c. 2.5 mm long,

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 119

central rachis 20—5S0 cm long; longest branch 4—5 cm long; internodes ultimate branches

2.5—3.5 mm long, with 3—10 mm-long silky hairs. Pedicel 1.2—1.6 mm long, sparsely

hairy, hairs up to 2.5 mm long. Spikelet 3.4—3.9 mm long, c. 0.7 mm wide; callus 0.1 mm

long, densely hairy, hairs up to 11.5 mm long; lower glume 3.3—3.8 mm long, basal part

indurated, upper part hyaline, glabrous, margins ciliate; upper glume 3.4—3.7 mm long,

margins and vein ciliate; 1‘ lemma ciliate in upper half; 2°’ lemma up to 2.7 mm long. Lodicules

ciliate.

Not native; Paleo(sub)tropics. First record 1946. Extinct (1954).

On sandy soil, near water.

According to Sinclair (Sinclair 40184, 1954, SING) it is not native to Singapore but

had spread throughout the island. However, it has never been collected since his time.

Sacciolepis Nash

Culms single to loosely tufted or with short stolons, erect; internodes hollow; nodes glabrous.

Sheath glabrous. Ligule membranous, tip ciliolate. Young leaf blade inrolled. Blade glabrous,

margins smooth. Inflorescence spike-like panicle, central rachis glabrous, branches many;

longest branch up to 0.5 cm long, simple. Pedicel 0.4—1 mm long, glabrous. Spikelets single,

not flattened, gibbous, 2-flowered with 1“ floret sterile and 2™ bisexual, unawned; lowest

bract 0.45—0.55 times as long as and 2™ as long as spikelet; glumes and 1* lemma herbaceous;

lower glume at base enveloping, cordate, glabrous, acute; 1‘ lemma glabrous or hispid at tip;

I“ palea hyaline.

la. Inflorescence 0.8—8 cm long. Spikelets elongate ovoid, 2—3.4 mm long,

acute. S. indica

lb. Inflorescence 12(—20) cm long. Spikelets subglobose to ovoid, up to 2

mm long, obtuse. S. myosuroides

Sacciolepis indica (L.) Chase — Fig. 122, Plate 31.

Short-spiked Sacciolepis

Ridley 1925: 232 (excluding Singapore). Bor 1960: 357. Keng et al. 1998: 180. —

S. indica var. indica: Gilliland 1971: 152.

Panicum indicum L.: Ridley 1907: 134.

S. angusta auct., non Stapf: Ridley 1925: 232.

S. turgida Ridl. 1925: 231. — S. indica var. turgida (Ridl.) Gilliland: Gilliland 1971: 154.

Culms 15—50(—90) cm long. Sheath 2.5—4.3 cm long, rounded to keeled at tip. Ligule

0.2—0.5 mm long. Blade 3—10.5(—19) cm long, 2—5.6 mm wide, base somewhat rounded.

Inflorescence 0.8—8 cm long, not interrupted. Spikelet elongate ovoid, (2—)2.6—3.4 mm

long, 0.7—1.5 mm wide, acute; lower glume |.1—1.7 mm long, 5 or 7 veins; upper glume

1.9—3.2 mm long, 9 or 11 veins, glabrous to hispid in upper half, obtuse to cuspidate; 1*

lemma |.7—3 mm long, 7 or 9 veins, truncate to acute; 1‘ palea 0.7—1.2 mm long; 2" lemma

1—1.5 mm long, hyaline to indurated; 2™ palea 0.9—1.4 mm long, hyaline to indurated.

Native; India to Australia. First record 1889. Common.

In unshaded to shaded, wet to moist grassland.

Gilliland mentioned both the length of the culm and leaf blade to distinguish this

species from S. myosuroides. However, the character that seems to differentiate these species

best is the shape of the spikelet (Bor 1960). Hymenachne amplexicaulis differs in having

(partially) filled culms, sheaths with transverse veins, longer panicle branches (up to 6 cm

long) and non-gibbous spikelets. Panicum auritum has blades 9—23 mm wide, inflorescences

with the longest branches at least 2 cm long and spikelets that are not gibbous.

1 20 Supplement of Gardens’ Bulletin Singapore 57 (2005)

Sacciolepis myosuroides (R.Br.) A.Camus — Fig. 123.

Long-spiked Sacciolepis

Ridley 1925: 232. Bor 1960: 358. Gilliland 1971: 152. Keng et al. 1998: 180.

Panicum myosuroides R.Br.: Ridley 1907: 134.

Culms 65 cm long. Sheath 8—11 cm long, rounded. Ligule 0.5 mm long. Blade c. 25 cm long,

3.5 mm wide, base cuneate. Inflorescence 12(—20) cm long, interrupted. Spikelet subglobose

to ovoid, 1.7—2 mm long, 0.9 mm wide, obtuse; lower glume c. 0.8 mm long, 3 veins; upper

glume c. 1.7 mm long, 7 veins, glabrous, obtuse; 1** lemma 1.7 mm long, 9 veins, obtuse. 1*

palea 1.6 mm long; 2" lemma 1.3 mm long, indurated; 2"! palea 1.3 mm long, hyaline.

Native; India to Australia. Single record 1890 from Kranji. Extinct (1890).

In damp places.

The difference from S. indica is discussed there.

Schizachyrium Nees

Schizachyrium brevifolium (Sw.) Nees, mentioned as ‘very shortlived in Singapore’

(Gilliland 1971: 291) has not been found. Compared with S. sanguineum, this is a smaller

plant with leaves up to 4 cm long and lower glume of the sessile spikelet up to 2.5 mm long.

Schizachyrium sanguineum (Retz.) Alston — Fig. 124.

Gilliland 1971: 289. Keng et al. 1998: 181.

Andropogon hirtiflorus auct., non Kunth: Ridley 1907: 166.

S. semiberbe Nees: Ridley 1925: 210.

Culms tufted, erect, 115 cm long; nodes glabrous. Sheath 5—8 cm long, glabrous, basal strongly

keeled, upper rounded. Ligule membraneous, 1.2 mm long, tip ciliolate. Young leaf blade

folded along midrib. Blade 14 cm long, 3.5 mm wide, glabrous, base rounded. Inflorescence

spike-like raceme, 2—10 per culm; rachis 4—6 cm long, articulated, hairy on the callus with

hairs up to 1.5 mm long, outside rounded, inside flattened. Pedicel 5 mm long, flattened,

hairy on one margin. Spikelets in pairs of 1 sessile and | pedicelled; sessile spikelet 5.8—6.5

mm long, 0.5—0.6 mm wide, not flattened, 2-flowered with 1* floret sterile and 2"? bisexual,

awned; callus hairy; lowest bract as long as and 2™ 0.9 times as long as spikelet; lower glume

5.8—6.5 mm long, herbaceous, glabrous, 5 veins, marginal ones keeled, tip bifid; upper glume

5.5—6 mm long, boat-shaped, hyaline, 1 vein, keeled, margins and keel ciliolate; 1“ lemma

5.5 mm long, hyaline; 1‘ palea absent; 2" lemma 5 mm long, deeply cleft for c. 4 mm, awn

from notch 10—16 mm long with basal half twisted and upper half antrorsely scaberulous; 2"

palea 1.2 mm long, hyaline. Pedicelled spikelet as sessile, but 3.5—5 mm long, unawned;

lower glume 3.5—5 mm long, aristate 0.5 mm; 2" lemma 3.2 mm long, unawned.

Native; India to China and Malesia. First record 1880s. Extinct (1959).

In sandy places near the sea: beach or woodlands.

Scrotochloa Judz.

Judziewicz 1984: 299—304.

Scrotochloa is split off from Leptaspis because the inflorescence articulates as a whole

and its central rachis bears only one node (Judziewicz 1984). In Leptaspis the inflorescence is

persistent and its central rachis bears more than one node.

Figure 121. Saccharum officinarum L.

Figure 122. Sacciolepis indica (L.) Chase.

Figure 123. Sacciolepis myosuroides (R.Br.) A.Camus.

121

(Excluding the Bamboos)

2 to the Grasses of

ELLEN

0 SOLES LEE REN NLS TOES

122 Supplement of Gardens’ Bulletin Singapore 57 (2005)

Scrotochloa urceolata (Roxb.) Judz. — Fig. 125.

Shield Grass, Rumput Babi

Keng et al. 1998: 181. Judziewicz 1984: 300.

Leptaspis urceolata (Roxb.) R.Br.: Ridley 1907: 150; 1925: 255. Gilliland 1971: 47.

Culms single or in small tufts, erect, rhizome c. 9 mm diameter, 30—100 cm long. Sheath c.

9 cm long, rounded, with transverse veins visible from the inside, sparsely hairy, margins

glabrous. Ligule basal part membranous and 0.3 mm long, hairs at tip 0.5 mm long. Young

blade inrolled. Blade 11—32 cm long, 40—62 mm wide, twisted (underside facing upward),

pinninerved and with transverse veins, glabrous, base abruptly narrowed, pseudopetiole 20—

50 mm long, tip acuminate. Inflorescence panicle, deciduous as a whole, central rachis absent,

branches 5—8; longest branch 10—18 cm long, densely hairy, simple or branched. Pedicel

4—6 mm long. Spikelets heteromorphous, unisexual, in pairs of 1 male and | female, 1-

flowered, unawned. Female spikelet 6—7.5 mm long, 4—4.5 mm wide, globose-obovoid,

not flattened; both glumes early caducous; lemma 6—7.5 mm long, indurated, 11 veins,

obovoid, constricted at basal 1.5 mm, with many hooked hairs, margins fused but for pore at

tip, tip beaked; palea 7—-8.2 mm long, protruding through pore at tip of lemma, thin herbaceous,

glabrous, tip bifid over 0.5—1 mm. Male spikelet 4#—6 mm long, 0.7—2.3 mm wide, more or

less laterally flattened, herbaceous; lower glume 1.5 mm long, glabrous, | vein, obtuse; upper

glume as lower, 2.8 mm long; lemma 5 mm long, 9 veins, glabrous, truncate; palea 5.8 mm

long.

Native; Sri Lanka, SE Asia, New Guinea, Solomon Islands. First record 1889. Rare.

In moist and deeply shaded places in primary and mature secondary forest.

Setaria P.Beauv.

Veldkamp 1994: 373—384.

Ligule basal part membranous, with hairs at tip. Inflorescence 1 per sheath and 1—4 per

culm, raceme or (spike-like) panicle, central rachis present. Pedicel present. Spikelets single,

most or all with involucre of 1 or more bristles, bristles persistent (not falling off with the

spikelet). Spikelet not flattened, 2-flowered with 2™ floret bisexual, unawned; lowest bract

0.25—0.45 times as long as spikelet; glumes and 1“ lemma (thin) herbaceous; 1* palea hyaline;

2™ Jemma and palea indurated, glabrous.

Setaria sphacelata (Schumach.) Stapf & C.E. Hubb. has been grown in the University

Garden (now university campus adjacent to the Singapore Botanic Gardens and the Bukit

Timah Road).

Differences with Cenchrus and Pennisetum are mentioned under Cenchrus.

The key is after Veldkamp (1994).

la. Blades plicate (Fig. 7c). Inflorescence a lax panicle, longest branch 1—16

cm long. 2

lb. Blades flattened. Inflorescence dense and bottle-brush-shaped, branches

absent or up to 0.8 cm long. 3

Figure 124. Schizachyrium sanguineum (Retz.) Alston, rachis with a pair of spikelets on

basal internode and one sessile spikelet on the upper internode.

Figure 125. Scrotochloa urceolata (Roxb.) Judz., pair of spikelets.

a. female spikelet, b. male spikelet.

Figure 126. Setaria barbata (Lam.) Kunth.

Figure 127. Setaria palmifolia (J.K6nig) Stapf var. palmifolia.

123

Field Guide to the Grasses of Singapore (Excluding the Bamboos)

124 Supplement of Gardens’ Bulletin Singapore 57 (2005)

2a. Blade margin at base hairy. Central rachis of inflorescence hairy; branches

hairy. 2"' lemma easily detachable from the spikelet, (at maturity)

transversally rugose. S. barbata

2b. Blade margin at base glabrous. Central rachis of inflorescence glabrous,

scaberulous; branches scaberulous, with short hairs at base only. 2°¢

lemma persistent, (at maturity) finely rugulose.

S. palmifolia var. palmifolia

3a. Involucre absent or consisting of 1—3 bristles. Lower glume enveloping;

2™ lemma easily detachable from the spikelet. S. italica

3b. Involucre consisting of 4—15 bristles. Lower glume half enveloping; 2"

lemma persistent. S. parviflora

Setaria barbata (Lam.) Kunth — Fig. 126, Plate 32.

Veldkamp 1994: 375; 1999b: 236. Keng et al. 1998: 182.

Culms creeping, geniculate, ascending or scrambling, 30—200 cm long; nodes short-hairy.

Sheath 3—9 cm long, keeled to winged, glabrous or hairy, margins hairy. Ligule basal part

0.3—1 mm long, hairs at tip 0.5—O.7 mm long. Young leaf blade plicate and more or less

folded along midrib. Blade 13—68 cm long, (6—)19—35 mm wide, plicate, both sides with

up to 1.5 mm-long hairs, base long cuneate (to pseudopetiolate) and at margin with up to 4.5

mm long hairs. Inflorescence 5.5—18 cm long, open, central rachis with c. 2 mm-long hairs;

longest branch 1—3 cm long, hairy, branched. Pedicel 0.5—0.7 mm long; bristle 1, up to 4

mm long. Spikelet 2.4—2.6 mm long, |.1—1.3 mm wide, |“ floret male; 2" lowest bract c.

0.65 times as long as spikelet; lower glume 0.7—0.8 mm long, enveloping, 3 veins, glabrous;

upper glume |.6—1.7 mm long, 7 veins, glabrous; 1“ lemma 2.3—2.5 mm long, 7 veins,

glabrous; 1‘ palea 2.1—2.2 mm long; 2™ lemma |.9—2.2 mm long, transversely rugose (at

maturity), easily detachable from the spikelet; 2"¢ palea 1.8—2 mm long, rugulose.

Not native; probably native to West Africa, now pantropical. First record 1971.

Common.

Weed in moist and (partly) shaded roadsides and margins of woodlands.

Setaria italica (L.) P.Beauv. — Big. 128.

Italian Millet, Sekoi

Ridley 1907: 144; 1925: 234 (both excluding Singapore). Gilliland 1971: 156.

Veldkamp 1994: 377. Keng et al. 1998: 182.

Culms single, erect, 25—90 cm long; nodes glabrous. Sheath 2—16 cm long, rounded, glabrous

or sparsely hairy towards tip, margins hairy. Ligule basal part 0.5—0.7 mm long, hairs at tip

1.2—1.5 mm long. Young leaf blade inrolled. Blade 5—40 cm long, 4—16 mm wide, flattened,

glabrous, margin scaberulous, base rounded and at margin with 1 mm long hairs. Inflorescence

(0.5—)8 dense, central rachis with c. 1.5 mm-long hairs; longest branch up to

0.8 cm long, hairy, branched. Pedicel 0.5—0.8 mm long; bristles O—3, up to 11 mm long.

Spikelet 2.5—2.8 mm long, 0.9—1.3 mm wide, |“ floret sterile; 2"? lowest bract 0.85—0.95

times as long as spikelet; lower glume 1.2—1.3 mm long, enveloping, 3 or 5 veins, glabrous;

Figure 128. Setaria italica (L.) P.Beauv.

Figure 129. Setaria parviflora (Poir.) Kerguélen.

Figure 130. Sorghum bicolor (L.) Moench, sessile spikelet.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 125

Seen

cst

126 Supplement of Gardens’ Bulletin Singapore 57 (2005)

upper glume 2.3—2.4 mm long, 5 or 7 veins, glabrous; 1* lemma 2.5—2.6 mm long, 5 or 7

veins, glabrous; 1“ palea 1—1.4 mm long; 2° lemma 2.4—2.5 mm long, scaberulous, easily

detachable from the spikelet; 2" palea c. 2 mm long, scaberulous.

Not native; probably native to warm-temperate subtropical Eurasia. First record 1962.

Extinct (1969).

On dry waste ground.

The species is grown for its grain and in Singapore has disappeared after its cultivation

ceased.

Setaria palmifolia (J.K6nig) Stapf var. palmifolia — Fig. 127.

Broad-leaved Bristle-grass

Veldkamp 1994: 379.

Panicum plicata auct., non Lam.: Ridley 1907: 136 (excluding Singapore).

S. palmifolia (J.K6nig) Stapf: Gilliland 1971: 157. Keng et al. 1998: 182.

S. plicata auct., non Cooke: Ridley 1925: 235 (excluding Singapore).

Culms tufted, erect, c. 85 cm long; nodes sparsely short-hairy. Sheath 20 cm long, rounded,

sparsely hairy, margins glabrous or hairy. Ligule basal part 1.4 mm long, hairs at tip 0.6 mm

long. Young leaf blade plicate and more or less folded along midrib. Blade 24—33 cm long,

14—38 mm wide, plicate, glabrous or upperside hairy, base cuneate and with up to 5 mm long

hairs. Inflorescence 25—37 cm long, open, central rachis scaberulous; longest branch 6—16

cm long, scaberulous, hairy at base, branched, ending in a bristle. Pedicel 0.5—0.8 mm long;

bristle 0—1, up to 9 mm long. Spikelet 3.3—3.5 mm long, 1.1—1.2 mm wide, 1|* floret

sterile; 2"? lowest bract c. 0.7 times as long as spikelet; lower glume 1.3—1.5 mm long, half

enveloping, 3 (or 5) veins, glabrous; upper glume 1.9—2.2 mm long, 7 veins, glabrous; |*

lemma 3—3.3 mm long, 6 veins, glabrous; 1‘ palea 1.4—1.6 mm long; 2" lemma 2.5—2.7

mm long, rugulose at maturity, persistent; 2"* palea 2.4—2.5 mm long, rugulose.

Probably not native; India to China, Japan, Malesia to New Guinea, not in Australia.

First record 1904 from the Botanic Gardens, 1962 in the wild. Extinct (1963).

Forest grass on lightly shaded and fairly moist but not wet ground.

The species was probably cultivated in the Botanic Gardens. In the wild it was found

only in one area adjacent to the Botanic Gardens from where it probably had spread.

Setaria parviflora (Poir.) Kerguélen — Fig. 129.

Veldkamp 1994: 379. Keng et al. 1998: 183.

S. glauca auct., non (L.) P.Beauv.: Ridley 1907: 144.

S. pallidifusca (Schumach.) Stapf & C.E.Hubb.: Gilliland 1971: 159 (‘pallide-fusca’ ).

S. pumila auct., non (Poir.) Roem. & Schult.: Tan 1995: 147.

S. rubiginosa Migq.: Ridley 1925: 234.

Culms tufted, with short rhizomes, 40—150 cm; nodes glabrous. Sheath 2—6 cm long, basal

flattened and midrib raised, upper rounded, glabrous. Ligule 0.8—1.5 mm long, basal part

0.3—1 mm long, hairs at tip 0.5—-1 mm long (hairs shorter to longer than membrane). Young

leaf blade inrolled. Blade S—17 cm long, 4—7 mm wide, flattened, glabrous, scaberulous,

base cuneate to somewhat rounded and scaberulous or with few up to 3 mm long hairs.

Inflorescence 2—17 cm long, dense, central rachis with c. 1 mm-long hairs; longest branch

0—0.4 cm long, hairy, simple. Pedicel 0.3—0.5 mm long; bristles 4—9, up to 7 mm long.

Spikelet 1.9—2.3 mm long, 0.9—1 mm wide, |“ floret sterile; 2"° lowest bract c. 0.5 times as

long as spikelet; lower glume 0.8—1 mm long, half enveloping, 3 veins, glabrous; upper

glume 1—1.3 mm long, 5 veins, glabrous; 1‘ lemma 1.7—2.3 mm long, 5 veins, glabrous; 1*

palea 1.5—2 mm long; 2™ lemma 1—2 mm long, rugulose, persistent; 2" palea 1.5—1.7 mm

long, rugulose.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) iy

Native; pan(sub)tropical. First record 1889. Rare.

On dry roadsides, fields, sandy sea shore.

The inclusion of S. pumila in Tan (1995: ‘rare’) most probably refers to S. parviflora.

Material of the former, which differs from S. parviflora in having spikelets of 2.8—3.5 mm

long (Veldkamp 1994), has not been found.

Sorghum Moench

Culms tall, erect; basal internodes at least 5 mm diameter. Sheath rounded, margins glabrous.

Ligule membranous, tip ciliolate to ciliate. Young leaf blade inrolled. Blade glabrous, midvein

at upperside at least at base prominent and white, base hairy. Inflorescence panicle, central

rachis present, branches many and branched. Pedicel flattened, margins hairy. Spikelets in

pairs or threes of | sessile and 1 or 2 pedicelled, 2-flowered. Sessile spikelet callus hairy; the

two lowest bracts c. as long as spikelet; lower glume flat to rounded, upper glume boat-

shaped; both lemmas and 2" palea hyaline; 1‘ palea absent. Pedicelled spikelet lanceolate,

3—5 mm long, 0.8—0.9 mm wide, dorsiventrally flattened, male or sterile, unawned.

la. Culms tufted. Spikelets nearly globose at maturity, sometimes awned.

S. bicolor

lb. Culms long stoloniferous. Spikelets oblong-lanceolate, dorsiventrally

flattened, not much enlarged at maturity, unawned. _ S. propinquum

Sorghum bicolor (L.) Moench — Fig. 130.

Great Millet

Keng et al. 1998: 183.

S. vulgare (L.) Pers.: Gilliland 1971: 231.

Culms tufted, 135 cm long; nodes glabrous. Sheath 9—16 cm long, glabrous. Ligule 2—2.8

mm long, tip ciliolate. Blade 20—70 cm long, 21—42 mm wide, margins smooth, base rounded

to cordate. Inflorescence 11—40 cm long, central rachis glabrous; longest branch 4—9 cm

long, sparsely hairy at base. Pedicel 1—1.2 mm long. Sessile spikelet subglobose, 4—5.8

mm long, c. 3 mm wide, dorsiventrally flattened to subterete, awned or unawned; lower glume

3.6—5.4 mm long, indurated, 15 veins and 2" from margin keeled, with appressed hairs in

basal half, tip truncate; upper glume 3.7—-5 mm long, indurated, 9 veins, glabrous to sparsely

hairy; 1“ lemma 3.5—4.5 mm long; 2™ lemma c. 3.6 mm long, 0.6 mm deep bifid, awn if

present from notch and 4.5—10 mm long with basal part twisted and upper part scaberulous;

2" palea c. 2 mm long. Pedicelled spikelet variously reduced.

Not native; Africa, now cultivated in tropical and warm-temperate regions throughout

the world. First record 1892. Cultivated (not found after 1965).

This species has many cultivars; it is more variable than described above.

Sorghum propinquum (Kunth) Hitche. — Fig. 131.

Tebu Tikus

Gilliland 1971: 229. Keng et al. 1998: 184.

S. affine auct.: Ridley 1925: 195 (excluding Singapore).

Culms single, with long stolons, 120—250 cm; nodes short-hairy. Sheath 19—2i cm long,

glabrous, external ligule a fringe of hairs. Ligule 1.3—-2.2 mm long, basal part 0.3—0.9 mm

long, hairs at tip 1—1.5 mm long. Blade 40—65 cm long, 11—21 mm wide, margins serrulate,

base rounded. Inflorescence 25—42 cm long, central rachis with hairlike spicules; longest

branch 8—19 cm long, densely hairy at base. Pedicel 1.4—2 mm long. Sessile spikelet oblong-

lanceolate, 4.2—5.2 mm long, 1.5—1.8 mm wide, more or less dorsiventrally flattened,

128 Supplement of Gardens’ Bulletin Singapore 57 (2005)

unawned; lower glume 3.7—4.9 mm long, herbaceous, 9 veins, marginal ones keeled, transverse

veinlets present, entirely with appressed hairs, tip acute; upper glume 4—5S mm long,

herbaceous, 5 veins, sparsely hairy in lower 2/3™; 1* lemma 3.2—4.5 mm long; 2"? lemma

2—3.2 mm long; 2" palea 1.6—2.5 mm long. Pedicelled spikelet 3.8—5S mm long.

Probably not native; India to China and SE Asia. First record 1954. Rather rare.

In wasteland and on roadsides.

According to Keng et al. (1998) it is introduced in Singapore.

Sphaerocaryum Nees ex Hook.f.

Sphaerocaryum malaccense (Trin.) Pilg. — Fig. 132.

Elegant Roundseed

Gilliland 1971: 126. Keng et al. 1998: 184.

S. elegans Nees ex Steud.: Ridley 1907: 170; 1925: 241.

Culms creeping, erect part 10—20 cm long; nodes hairy. Sheath c. 4 mm long, rounded, hairy,

one margin hairy. Ligule row of hairs, 0.5—1 mm long. Blade 1—2.5 cm long, 4—9 mm

wide, glabrous, base cordate and at margin hairy. Inflorescence panicle, 2—4 cm long, central

rachis glabrous, branches many; longest branch c. 1.2 cm long, glabrous, simple. Pedicel 1—

2.5 mm long, glabrous. Spikelets single, c. 1 mm long, c. 0.4 mm wide, not flattened, 1-

flowered, unawned; lowest bract c. 0.6 and 2" bract almost as long as spikelet; lower glume

0.6 mm long, early caducous, thin herbaceous, glabrous, obtuse; upper glume as lower, 1 mm

long, acute; lemma | mm long, thin herbaceous, short-hairy; palea as lemma, but narrower.

Native; India to Thailand, Peninsular Malaysia and Singapore. Twice recorded 1889

from Ang Mo Kio and Krangi. Extinct (1889).

In moist to wet places, swampy streams in forests, fields.

Sporobolus R.Br.

Baaijens and Veldkamp 1991: 393—458.

Nodes glabrous (or at base covered by sheaths). Sheath glabrous or with few hairs at throat

and/or margins. Ligule membranous, tip ciliolate. Young leaf blade inrolled. Blade glabrous,

base cuneate, glabrous (or with few hairs). Inflorescence panicle, central rachis present,

branches many and branched. Pedicel glabrous. Spikelets single, glabrous, 1-flowered,

unawned.

The key to the species follows Baaijens and Veldkamp (1991).

la. Stoloniferous plants. Lower glume 0.55—0.9 times as long as spikelet,

upper glume 0.75—1 times as long. S. virginicus

1b. Tufted plants. Lower glume 0.2—0.5 times as long as spikelet, upper glume

up to 0.67 times as long.

2a. Pedicel 0.3—0.7 mm long. Spikelet 1.4—1.9 mm long; lemma and palea

herbaceous. S. indicus

Figure 131. Sorghum propinquum (Kunth) Hitchc., a group of one sessile and two

pedicelled spikelets.

Figure 132. Sphaerocaryum malaccense (Trin.) Pilg.

Figure 133. Sporobolus virginicus (L.) Kunth.

Figure 134. Sporobolus indicus (L.) R.Br. var. pyramidalis (P.Beauv.) Veldk.

Figure 135. Sporobolus tenuissimus (Schrank) Kuntze.

Figure 136. Sporobolus indicus (L.) R.Br. var. flaccidus (Roem. & Schult.) Veldk.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 129

Fig. 132

Fig. 133 Fig. 134 | Fig. 135 Fig. 136

] 30 Supplement of Gardens’ Bulletin Singapore 57 (2005)

2b. Pedicel 2—6 mm long. Spikelet 0.9—1.25 mm long; lemma and palea

hyaline. S. tenuissimus

Sporobolus indicus (L.) R.Br.

Literature and synonymy: see varieties.

Culms single or tufted, rhizomes absent, erect, not rooting at lower nodes, basal internodes

almost absent and only elongating in flowering culm. Leaves of non-flowering plants all

basal, covering the nodes. Sheath 3—10 cm long, rounded, throat with up to 0.5 mm long

hairs, one margin hairy. Ligule 0.2—0.4 mm long. Blade 4—35 cm long, 2—4.5 mm wide,

base glabrous or sparsely hairy. Pedicel 0.3—0.7 mm long. Spikelet not flattened; glumes

hyaline; lemma |.2—1.9 mm long, herbaceous; palea 1.2—1.8 mm long, herbaceous.

Native; pantropical. First record 1877. Common.

In unshaded to partly shaded, dry to moist places, e.g. roadsides, fields, beaches.

Although fungus infections of the spikelets have been reported frequently, they are

not known from Singapore, possibly because mowing reduces the chances of infection (Baaijens

and Veldkamp 1991: 425).

This is a very variable species. At present three varieties are known for Singapore, but

these are difficult to distinguish. The key to the varieties in Singapore is after Baaijens and

Veldkamp (1991):

la. Upper glume truncate, less than half as long as the spikelet, slightly longer

than the lower glume. S. indicus var. pyramidalis

1b. Upper glume more or less acute, (0.4—0.67 times as long as spikelet,

distinctly longer than the lower glume. 2

2a. Panicle usually somewhat lax and spikelets well-spaced. Spikelets usually

1.4—1.6 mm long. Anthers usually 2, 0.5—0.8 mm long. Seed 0.6—0.9

mm long. S. indicus var. flaccidus

2b. Panicle usually contracted and spikelets crowded. Spikelets usually 1.8—

1.9 mm long. Anthers usually 3, 0.7—1 mm long. Seed 0.9—1.1 mm

long. S. indicus var. major

Sporobolus indicus (L.) R.Br. var. flaccidus (Roem. & Schult.) Veldk.

Lesser Dropseed — Fig. 136, Plate 34.

Baaijens and Veldkamp 1991: 433. Keng et al. 1998: 184.

S. diandrus (Retz.) P.Beauv.: Ridley 1907: 170; 1925: 244. Gilliland 1971: 106 (as diander).

Culms 20—100 cm long. Panicle 14—21 cm long, usually somewhat lax; longest branch 2—

4.5 cm long, erecto-patent to patent, spikelets well-spaced. Spikelets usually 1.4—1.6 mm

long; lowest bract 0.25—0.30 and 2™ 0.4—0.6 times as long as spikelet; lower glume 0.3—

0.5 mm long, obtuse to truncate; upper glume 0.5—0.9 mm long, acute, serrate. Anthers

usually 2, 0.5—0.8 mm long. Seed 0.6—0.9 mm long.

Native; Mauritius, Pakistan, India, Malesia to Polynesia and Australia. First record

1877. Common.

Sporobolus indicus (L.) R.Br. var. major (Biise) Baaijens

Common Dropseed ;

Baaijens and Veldkamp 1991: 437. Keng et al. 1998: 184.

S. fertilis (Steud.) Clayton: Gilliland 1971: 106.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 131

S. indicus (L.) R.Br.: Ridley 1907: 171; 1925: 244.

Culms 20—100 cm long. Panicle 15—32 cm long, usually contracted; longest branch 2—3.5

cm long, erect, spikelets crowded. Spikelets usually 1.8—1.9 mm long; lowest bract 0.25—

0.30 and 2" 0.4—0.6 times as long as spikelet; lower glume 0.4—0.5 mm long, obtuse to

truncate; upper glume 0.6—1 mm long, obtuse to erose and denticulate. Anthers usually 3,

0.7—1 mm long. Seed 0.9—1.1 mm long.

Native; India to Japan, Malesia to Australia. First record 1892. Rather common.

Sporobolus indicus (L.) R.Br. var. pyramidalis (P.Beauv.) Veldk. — Fig. 134.

Baaijens and Veldkamp 1991: 439. Duistermaat 2004: 39.

Culms c. 155 cm long. Panicle 30—40 cm long, lax; longest branch 9 cm long, patent, spikelets

crowded. Spikelet c. 1.9 mm long; lowest bract 0.15 and 2™ 0.3 times as long as spikelet;

lower glume 0.3 mm long, truncate; upper glume 0.6 mm long, truncate. Anthers 3, 0.9 mm

long. Seed 0.8—1 mm long.

Not native; Africa, S. America. First record 2004. Rare.

Unshaded, open vegetation, sandy soil.

Sporobolus tenuissimus (Schrank) Kuntze — Fig. 135, Plate 35.

Baaijens and Veldkamp 1991: 444. Veldkamp 1997b: 512; 2003: 500. Duistermaat

2004: 38.

Single, erect, stolons absent, not rooting at lower nodes, 15—70 cm long, basal internodes

well developed. Leaves scattered along culm. Sheath 9—10 cm long, rounded, midrib slightly

raised, glabrous. Ligule 0.1 mm long. Blade c. 15 cm long, 2 mm wide. Inflorescence 9—20

cm long, lax; longest branch c. 3.5 cm long, erecto-patent, spikelets well-spaced. Pedicel 2—

6 mm long. Spikelet c. 1.1 mm long, 0.7 mm wide, laterally flattened; lowest bract c. 0.15 and

2™ c. 0.45 times as long as spikelet; lower glume 0.15 mm long, hyaline; upper glume 0.5 mm

long, hyaline; lemma | mm long, hyaline; palea 0.9 mm long, hyaline. Anthers 3, 0.3—0.4

mm long. Seed 0.7 mm long.

Not native; tropical S America, Africa and India, introduced in Vietnam and Malesia.

First record 2003. Rare.

In flowerbeds and on roadsides, open vegetation, unshaded.

This species is likely to spread further in Singapore. It often grows 1n association with

Eragrostis amabilis, which differs in the more prostrate to ascending habit, the many-flowered

spikelets with a jagged and not globose profile, and the distinctly setose paleas.

Sporobolus virginicus (L.) Kunth — Fig. 133.

Beach Dropseed

Gilliland 1971: 108. Baaijens and Veldkamp 1991: 445.

Culms long stoloniferous, lower nodes rooting, erect part c. 20 cm long; basal internodes

shorter than sheaths; nodes covered by sheaths. Leaves scattered along culms. Sheaths

overlapping, rounded, margin at tip with few hairs. Ligule 0.5 mm long. Blade 8—9 cm long,

3.5 mm wide, base with a few hairs. Inflorescence c. 6 cm long, c. 0.5 cm wide; longest

branch c. 1 cm long, appressed. Pedicel 0.3—1.5 mm long. Spikelet c. 2.3 mm long, 0.5 mm

wide, somewhat laterally flattened; lowest bract c. 0.7 and 2" c. 0.95 times as long as spikelet;

lower glume 1.6 mm long, thin herbaceous; upper glume 2.1 mm long, thin herbaceous; lemma

2.1 mm long, thin herbaceous; palea 1.9 mm long, thin herbaceous. Anthers 3, 1.1 mm long.

Seed c. 0.9 mm long.

Native; pan(sub)tropical. Single record 1941 from Changi. Extinct (1941).

Sandy sea-beach above high-water mark.

In Peninsular Malaysia it is only known from Langkawi. However, the species might

132 Supplement of Gardens’ Bulletin Singapore 57 (2005)

have been overlooked, as it rarely seems to flower (according to Van Steenis on annotated

material from Bali; Baaijens and Veldkamp 1991: 448). Also, because of the unstable substrate

populations may be very temporary.

Stenotaphrum Trin.

Sauer 1972: 202—222.

Stenotaphrum secundatum (Walt.) Kuntze — Fig. 137, Plate 33.

St. Augustine Grass, Rumput St. Augustine

Sauer 1972: 210. Keng et al. 1998: 185.

S. dimidiatum auct., non (L.) Brongn.: Gilliland 1971: 205.

Culms with long stolons, erect part c. 20 cm long; nodes glabrous. Sheaths 2—5.5 cm long,

flattened, midrib raised, glabrous, margin (at tip) with 1—1.5 mm long hairs. Ligule

membranous, 0.5 mm long, tip ciliolate. Young leaf blade folded along midrib. Blade (0.7—)

5—15 cm long, 4—7 mm wide, glaucous, sparsely hairy, base cuneate to rounded with margins

glabrous or sparsely hairy, tip obtuse. Inflorescence spike-like panicle; central rachis 6—9

cm long, 3—4 mm wide, glabrous, sinuous and alternately broadly winged; branches many,

up to 0.5 cm long, simple and each with 3 spikelets. Pedicel 0.2—3.5 mm long, flattened,

margins sparsely hairy. Spikelets single, basal 2 on each branch sessile to shortly pedicelled

and uppermost long pedicelled (and vestigial), 4.3—4.8 mm long, 1.6 mm wide, not flattened,

2-flowered with 1* floret male and 2™ bisexual, unawned; lowest bract 0.25—0.3 and 2™

0.95—1 times as long as spikelet; lower glume 1.2—1.4 mm long, orbicular, hyaline, glabrous,

obtuse; upper glume 3.9—4.3 mm long, herbaceous, 7 veins, glabrous, acute; 1‘ lemma 4.3—

4.6 mm long, indurated, glabrous, acute; 1‘ palea 3.8—4.4 mm long, indurated; 2"* lemma 4

mm long, herbaceous, hairy at tip; 2" palea 3.8—4 mm long. Pollen irregularly shaped.

Not native; paleotropics. First record 1950. Common.

On roadsides, lawns.

Originally being a coastal pioneer, it seems to grow better in places with well-drained

soil. It only propagates vegetatively through its long stolons. Flowering plants are rare, pollen

is misshapen, and fruits have not been found in Singapore.

Themeda Forssk.

Culms erect, stolons absent, with drooping branches; internodes solid; nodes glabrous. Sheath

keeled, margins glabrous or hairy (in basal half). Ligule membranous, tip lacerate. Spatheate

inflorescence branched, with several spike-like racemes arising from spathes (much reduced

leaves). Ultimate spathe with a cluster of |—8 spike-like racemes; each spike-like raceme at

base with involucre of 2 pairs of sterile (or rarely male) and unawned spikelets, upper part

with | or 2 fertile and 0—3 sterile spikelet(s). Fertile spikelet not flattened, 2-flowered, awned;

callus densely hairy; the two lowest bracts about as long as spikelet; glumes indurated; lemmas

hyaline, 2" with awn from tip; paleas absent.

la. Young leaf blade folded along midrib. Sheath at least in upper half sparsely

to densely hairy. Ultimate spathe with cluster of 4—8 spike-like racemes

Figure 137. Stenotaphrum secundatum (Walt.) Kuntze.

Figure 138. Themeda arguens (L.) Hack., inflorescence with spathe.

Figure 139. Themeda villosa (Poir.) A.Camus.

a. spike-like raceme, b. pair of spikelets.

Figure 140. Thysanolaena latifolia (Roxb. ex Hornem.) Honda.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 133

Fig. 137

Fig. 140

134 Supplement of Gardens’ Bulletin Singapore 57 (2005)

with 4—8 awns. Spike-like raceme (nearly) sessile with peduncle up to

3 mm long, fertile spikelet 1. T. arguens

1b. Young leaf blade inrolled. Sheath glabrous or hairy on the margin. Ultimate

spathe with 1—2 spike-like racemes with 2—4 awns. Spike-like raceme

with peduncle 15—28 mm long, fertile spikelets 2. T. villosa

Themeda arguens (L.) Hack. — Fig. 138.

Lesser Tasselgrass, Rumput Misai Adam

Ridley 1925: 212. Gilliland 1971: 300. Keng et al. 1998: 185.

Anthistiria arguens Willd.: Ridley 1907: 168.

Culms single or tufted, 30—120 cm long. Sheath 3.5—5.5 cm long, at least in upper half

sparsely to densely hairy, margins glabrous or hairy in basal half. Ligule 1—1.4 mm long,

glabrous. Young leaf blade folded along midrib. Blade up to 45 cm long, 5—6 mm wide,

glabrous or hairy at margin and/or upperside, base rounded and hairy. Spatheate inflorescence

15—50 cm long. Ultimate spathe 3—4 cm long, hairy in basal half, with cluster of 4—8

spike-like racemes. Spike-like raceme sessile, upper part with | pedicelled fertile spikelet.

Involucral spikelets | glume each, sessile spikelet 5—8 mm long, pedicelled spikelet 0.4 mm

long. Pedicel of fertile spikelet 1.3 mm long, glabrous. Fertile spikelet 6—6.5 mm long, 1.5

mm wide; callus 2.5 mm long; lower glume 6—6.4 mm long, setose near tip; upper glume

6.3—6.5 mm long, upper half setose; 1*t lemma 3.1—3.5 mm long, glabrous; 2"? lemma 3—

3.2 mm long, threadlike, awn 65—85 mm long with basal 3 mm glabrous and upper part

twisted and setose.

Native; SE Asia and Australia. First record 1893. Rare.

Locally abundant in open dry places, on roadsides, railway tracks, sometimes as a

pioneer.

Themeda villosa (Poir.) A.Camus — Fig. 139, Plate 36.

Greater Tasselgrass, Rumput Riong

Ridley 1925: 212. Gilliland 1971: 301. Keng et al. 1998: 185.

Culms tufted, 200—300 cm long; internodes of non-flowering plants very short. Leaves of

non-flowering plants all basal and distinctly distichous, shoot fan-shaped. Sheath 12—23 cm

long, glabrous. Ligule 1—1.8 mm long, glabrous or hairy at the back. Young leaf blade inrolled.

Blade 60—100 cm long, 7—10 mm wide, glabrous, base cuneate to somewhat rounded and

margin sparsely hairy. Spatheate inflorescence up to 100 cm long. Ultimate spathe 4—8 cm

long, glabrous, scaberulous, with cluster of 1—2 spike-like racemes. Spike-like raceme

peduncled, upper part with a pair of | sessile fertile and | pedicelled sterile spikelet, and a

triplet of 1 sessile fertile and 2 pedicelled sterile spikelets. Involucral spikelets all similar, 2

herbaceous glumes (and sometimes 2 hyaline lemmas), 9—14.2 mm long. Fertile spikelets

7—7.8 mm long, 1.8—1.9 mm wide; callus 1.5 mm long; lower glume 7—7.8 mm long,

indurated, densely hairy; upper glume 7—7.6 mm long, densely hairy; 1“ lemma 4.8—5.5

mm long, glabrous; 2™ lemma 4.5—5.3 mm long, awn up to 32 mm long and twisted in basal

half. Pedicel of sterile spikelet 2 mm long, glabrous. Sterile spikelets as the involucral spikelets.

Native; SE Asia. First record 1880s. Rare.

In unkempt sandy places, margin of woodland, riverbanks, near the sea (just above

high tide mark).

Thuarea Pers. |

Thuarea involuta (G.Forst.) R.Br. ex Roem. & Schult. — Fig. 141, Plate 37.

Sea Nut-grass

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 135

Gilliland 1971: 207. Keng et al. 1998: 186.

T. sarmentosa Pers.: Ridley 1907: 146; 1925: 236.

Culms with stolons 40—110 cm long, erect part 6—10 cm long; nodes hairy. Sheath 1.5—3

cm long, rounded, basal part glabrescent, upper part hairy, margins hairy. Ligule row of hairs,

0.5—1 mm long. Young leaf blade inrolled. Blade 0.8—4.6 cm long, 2—10 mm wide, underside

narrowly and densely ribbed (when dry), hairy, margins hairy, base cuneate. Inflorescence

deciduous, spike, 1—1.5 cm long; rachis 3.6 mm wide, longitudinally folded (U-shaped),

with transverse veins, hairy, narrowing towards tip, at base with | bisexual spikelet followed

by 4 or 5 male spikelets, after flowering widening and bending down to enclose mature bisexual

spikelet. Pedicel absent. Bisexual spikelet persistent, single, 4.2—4.7 mm long, laterally

flattened, 2-flowered with 1* floret male and 2"! female, unawned; the two lowest bracts as

long as spikelet; lower glume absent; upper glume 3.5—4.5 mm long, thin herbaceous, 7

veins, hairy, obtuse; 1* lemma as upper glume, 4.1—4.6 mm long, 5 veins; 1“ palea 4—4.3

mm long, hyaline; 2" lemma 4—4.5 mm long, indurated, 5 veins, upper part sparsely hairy,

tip truncate and apiculate; 2" palea 3.9—4.3 mm long, indurated. Male spikelets deciduous

after flowering, as bisexual spikelet but gradually decreasing in size, bothflorets male, all

bracts herbaceous.

Native; Indo-Pacific. First record 1890. Rare.

On sandy beaches.

Its last record on the main island (Tanjong Rhu up to Changi) dates back to 1921.

Today, it is only known from some of the southern islands.

The incurved rachis forms a nut-like structure. It articulates as a whole and, being

watertight, will float in seawater thus aiding dispersal of the seed.

Thysanolaena Nees

Thysanolaena latifolia (Roxb. ex Hornem.) Honda — Fig. 140.

Tiger Grass, Buluh Tebrau

Keng et al. 1998: 186.

T. agrostis Nees: Ridley 1907: 143; 1925: 241 (both excluding Singapore).

T. maxima (Roxb.) Kuntze: Gilliland 1971: 45.

Culms tufted, erect, (40—)200—300 cm long; internodes solid, at least 3 mm diameter; nodes

glabrous. Sheaths 12 cm long, rounded, with transverse veins near margin at tip, submarginally

hairy. Ligule membranous, |—2 mm long, tip ciliolate. Young leaf blade inrolled. Blade 40—

63 cm long, 28—70 mm wide, with transverse veins present (best visible at underside),

glabrous, margin smooth to minutely scaberulous, base cuneate to rounded, pseudopetiole

2—3 mm long and glabrous or hairy. Inflorescence panicle, (20—)50—62 cm long, central

rachis glabrous, branches many; longest branch 15—35 cm long, 1.2 mm wide, glabrous,

straight or upward sinuous, branched; ultimate branchlets 0.5—1 cm long, 0.2 mm wide,

appressed, well-spaced to somewhat overlapping, spikelets crowded. Pedicel 0.5—1.4 mm

long, scaberulous. Spikelets single or in pairs, homomorphous, (1.6—)2—2.1 mm long, 0.6

mm wide, not flattened, (thin) herbaceous, 2-flowered (sometimes proliferating or abortive),

unawned; rachilla between 1* and 2" floret 0.4 mm long, at tip with 0.3 mm long hairs; the

two lowest bracts c. 0.3 times as long as spikelet; lower glume 0.4—0.5 mm long glabrous;

upper glume (0.5—)0.6—0.7 mm long, glabrous; 1‘ lemma (1.5—)1.8—2 mm long, glabrous;

1* palea absent; 2"? lemma (1—)1.5 mm long, with submarginal row of 0.7—1 mm long setae

on both sides; 2"* palea 0.7—0.9 mm long, glabrous, hyaline.

Not native; India to China and S to Peninsular Malaysia, usually above 300 m asl,

introduced in Africa and Singapore. First record 1959. Rather rare.

In open places, on rocky, sandy or clayey soil.

136 Supplement of Gardens’ Bulletin Singapore 57 (2005)

It is used as an ornamental plant. The leaves are used for wrapping glutinous rice and

the inflorescences for making brooms.

Tripsacum L.

Tripsacum dactyloides (L.) L. — Fig. 142.

Gama Grass

Gilliland 1971: 309. Keng et al. 1998: 186.

Culms tufted, outer culms geniculate, 110 cm long; internodes 30 mm diameter; nodes glabrous.

Sheath 13—20 cm long, rounded, glabrous. Ligule membranous, 1—1.2 mm long, hairy at

the back, tip ciliolate with hairs c. 0.2 mm long. Blade 45—140 cm long, 56—75 mm wide,

hairy at upperside, margin scaberulous, base rounded to cordate. Inflorescence raceme, | per

sheath and up to 6 along upper 20—35 cm of culm; rachis 5—6 cm long, at base 4—5.5 mm

wide and indurated and glabrous, in upper half c. 2 mm wide and herbaceous and hispid.

Pedicel 0.5—2 mm long, hispid. Spikelets unisexual, not flattened, 2-flowered, unawned,

basal 4—10 of the raceme single and female, upper ones in pairs and male; the two lowest

bracts as long as spikelet, acute; lemmas and paleas hyaline. Female spikelet 8 mm long, 4

mm wide, sunken into hollow rachis; lower glume dorsally flattened, indurated, glabrous;

upper glume boat-shaped, thin herbaceous, glabrous. Male spikelet 8.5 mm long, 2 mm wide;

lower glume dorsally flattened, herbaceous, 23—25 veins, submarginally keeled, keels hispid,

tip emarginate; upper glume boat-shaped, 5 veins, glabrous, tip acute.

Not native; Neotropics, introduced in Africa and Australasia. Cultivated (last record

1969).

It has only been grown in the 1960s in the Botanic Gardens. It is closely related to Zea

with which it has been hybridized.

Urochloa P.Beauv.

Veldkamp 1996b: 413—437.

Young leaf blade inrolled. Ligule basal part of fused hairs or membranous, hairs at tip as long

as to longer than basal part. Blade base rounded. Inflorescence panicle, central rachis present,

branches triquetrous to flattened with the spikelets adaxially. Pedicel present. Spikelets more

or less dorsiventrally flattened, 2-flowered with 1 floret sterile (or male in U. mutica) and 2"

bisexual, unawned; 2™ lowest bract (almost) as long as spikelet; glumes and 1* lemma (thin)

herbaceous; lower glume (half) enveloping; upper glume and 1“ lemma in upper part with

(obscure) transverse veins; 1‘ palea absent or hyaline; 2" lemma and palea indurated, glabrous,

transversely rugulose.

The key to the species follows Veldkamp (1996b).

la. Spikelets at least in middle of the branch paired. Lower glume margins not

overlapping. y.

1b. Spikelets solitary. Lower glume margins overlapping. 3

2a. Lower glume 5—7-veined, c. 0.9 times as long as spikelet. U. glumaris

2b. Lower glume O0- or 1-veined, 0.25—0.45 times as long as spikelet.

U. mutica

Figure 141. Thuarea involuta (G.Forst.) R.Br. ex Roem. & Schult.,-inflorescence with lower

spikelet bisexual and following spikelets male.

Figure 142. Tripsacum dactyloides (L.) L.

a. part of the rachis with female spikelet, b. pair of male spikelets.

Figure 143. Zea mays L., male spikelet.

Field Guide to the Grasses of Singapore (Excluding the Bamboos)

138 Supplement of Gardens’ Bulletin Singapore 57 (2005)

3a. Upper glume distally hairy. U. piligera

3b. Upper glume glabrous. U. subquadripara

Urochloa glumaris (Trin.) Veldk. — Fig. 147.

Veldkamp 1996b: 420. Keng et al. 1998: 186.

Brachiaria paspaloides (J.Presl) C.E.Hubb.: Gilliland 1971: 179.

Panicum distachyum auct., non L.: Ridley 1907: 133.

Culm decumbent or with short stolons, 25—40 cm long; internodes up to 2 mm diameter;

nodes hairy. Sheath 4.2—6.5 cm long, rounded, sparsely hairy at base (and tip), one margin

hairy. Ligule basal part 0.1—0.7 mm long, hairs at tip 0.4—0.7 mm long. Blade 5—16 cm

long, 3—5 mm wide, hairy, base at margin sparsely hairy. Inflorescence 4.5—12 cm long,

central rachis hairy, branches 3—4; longest branch 3.5—7 cm long, 0.6 mm wide, triquetrous,

hairy, simple or branched. Pedicel 0.5—1.8 mm long, setulose. Spikelets at least in middle of

branch paired, 3.5—4.2 mm long, 1.3 mm wide; lowest bract c. 0.9 times as long as spikelet;

lower glume 3—3.8 mm long, margins not overlapping, 5—7 veins, glabrous, tip acuminate

to minutely caudate; upper glume 3.3—4.2 mm long, 7 veins, glabrous, tip acuminate to

caudate and puberulous; 1* lemma 2.8—3.5 mm long, glabrous; 1* palea 0.1—0.6 mm long;

2™ lemma 2—2.6 mm long; 2™ palea 2.1—2.4 mm long.

Native; India to S China, Polynesia, Malesia, not in Australia. First record 1880s.

Rare.

Open moist places.

It is easily distinguished from the other Urochloa species by the long lower glume.

Urochloa mutica (Forssk.) T.Q.Nguyen — Fig. 144, Plate 38.

Para Grass

Veldkamp 1996b: 424. Keng et al. 1998: 187.

Brachiaria mutica (Forssk.) Stapf: Ridley 1925: 219. Gilliland 1971: 178.

Panicum muticum Forssk.: Ridley 1907: 133.

Culms decumbent, creeping or scrambling, 30—500 cm long; internodes at least 5 mm

diameter; nodes densely patent hairy. Sheath 5.5—20 cm long, rounded, (densely) patent

hairy, margins glabrous or one sparsely hairy. Ligule basal part 0.3—0.5 mm long, hairs at tip

0.5—1.5 mm long. Blade 10—20 cm long, 8—19 mm wide, hairy at least at underside, throat

hairy. Inflorescence 15—26 cm long, central rachis sparsely hairy, branches 14—20 and patent;

longest branch 3—12 cm long, 0.8—1.2 mm wide, more or less flattened, glabrous, simple or

branched. Pedicel 0.5 mm long, glabrous to sparsely hairy. Spikelets at least in middle of

branch paired or in threes, 3—3.3 mm long, 1.3—1.4 mm wide; lowest bract 0.25—0.45

times as long as spikelet; lower glume 0.7—1.4 mm long, margins not overlapping, | vein,

sparsely minute-hairy, tip obtuse to emarginate; upper glume 2.7—3.1 mm long, 5 veins,

glabrous, tip acute; 1*t lemma as upper glume, 2.9—3 mm long; |“ palea 2.8—3.1 mm long;

2™ Jemma 2—2.3 mm long; 2"? palea 1.9—2 mm long.

Not native; Brazil, introduced elsewhere. First record 1893. Common.

Weed on moist to wet soil in roadsides, wasteland, river banks, mangrove, beach.

Figure 144. Urochloa mutica (Forssk.) T.Q.Nguyen.

a. facing lower glume, b. facing upper glume.

Figure 145. Urochloa piligera (F.Muell. ex Benth.) R.D.Webster.

a. facing lower glume, b. facing upper glume.

Figure 146. Urochloa subquadripara (Trin.) R.D.Webster.

a. facing lower glume, b. facing upper glume.

Field Guide to the Grasses of Singapore (Excluding the Bamboos)

139

on aie,

Fig. 144

140 Supplement of Gardens’ Bulletin Singapore 57 (2005)

It was introduced as a fodder crop for cattle and is now naturalized throughout Malesia.

It flowers rarely, but is readily recognized by the long and rather stout stolons with patent-

hairy nodes and sheaths.

Urochloa piligera (F.Muell. ex Benth.) R.D.Webster — Fig. 145.

Veldkamp 1996b: 425; 1999b: 237. Duistermaat 2004: 40.

Brachiaria piligera (F.Muell. ex Benth.) Hughes

Culms tufted, decumbent to geniculate, 40—70 cm long; internodes up to 2 mm diameter;

nodes glabrous to sparsely hairy. Sheath 5—8 cm long, more or less flattened, midrib slightly

raised, glabrous, margins glabrous or one hairy. Ligule basal part 0.1—0.2 mm long, hairs at

tip 0.9—1 mm long. Blade 11—14 cm long, 7—8 mm wide, glabrous, margin sparsely hairy.

Inflorescence 10—20 cm long, central rachis glabrous, branches 2—5; longest branch 2.5—

5 cm long, 0.7—1 mm wide, flattened, glabrous, scaberulous, simple. Pedicel 0.3—0.5 mm

long, glabrous. Spikelets single, 3—3.4 mm long, 1.3—1.4 mm wide; lowest bract 0.35—0.5

times as long as spikelet; lower glume 1.2—1.5 mm long, margins overlapping, 7—13 veins,

glabrous, tip obtuse; upper glume 2.8—3.1 mm long, 7 veins, hairy, tip obtuse to acute; 1“

lemma as upper glume, 2.3—2.9 mm long, acuminate; 1“ palea absent; 2" lemma 2.2—2.5

mm long; 2™ palea 2—2.3 mm long.

Not native; Australia, Sulawesi, Moluccas, New Guinea, introduced in Singapore. First

record 1959. Rare.

In swampy places, on moist roadsides, on sandy clay.

Urochloa subquadripara (Trin.) R.D.Webster — Fig. 146, Plate 39.

Veldkamp 1996b: 429. Keng et al. 1998: 187.

Brachiaria distachya auct., non Stapf: Ridley 1925: 219. Gilliland 1971: 176.

Brachiaria subquadripara (Trin.) Hitchce.

Panicum distachyum auct., non L.: Ridley 1907: 133.

Culms geniculate or with short stolons, 30—90 cm long; internodes up to 2 mm diameter;

nodes glabrous to densely hairy. Sheath 3—9 cm long, rounded at base and more or less

flattened with midrib slightly raised at tip, giabrous or hairy, one margin hairy. Ligule basal

part 0.4—0.5 mm long, hairs at tip 0.4—1 mm long. Blade 5—15 cm long, 5—10 mm wide,

glabrous to sparsely hairy at upperside, base hairy. Inflorescence 6.5—13 cm long, central

rachis glabrous to sparsely hairy, branches 3—8; longest branch 3—6.5 cm long, 0.6—1 mm

wide, flattened, glabrous to hispid at base, scabrous, simple. Pedicel 0.5—0.7 mm long,

glabrous to sparsely hairy. Spikelets single, 3.5—4.3 mm long, 1—1.2 mm wide; lowest bract

c. 0.4 times as long as spikelet; lower glume 1.4—1.8 mm long, margins overlapping, 5—11

veins, glabrous, tip obtuse; upper glume 3—3.8 mm long, glabrous, 7 veins, tip obtuse to

acute; 1 lemma as upper glume, 2.8—3.5 mm long; 1* palea 0.1—3 mm long; 2™ lemma

2.5—2.7 mm long; 2™ palea 2.3—2.5 mm long.

Native; Kashmir to S China, throughout Malesia to Queensland. First record 1890.

Common. |

In open waste places, damp grass fields, on roadsides, drought resistant.

It has been confused with Eriochloa meyeriana, E. procera and Panicum repens. Both

Eriochloa species are distinguished by the presence of a cup-shaped callus at the base of the

spikelet and the absence of transverse veins in the upper glume and 1“ lemma. Panicum

repens differs in having primary panicle branches with at least secondary branches, abaxial

spikelets and lower glumes that are collar-shaped and much shorter (0.6—0.7 mm long).

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 141

Zea L.

Zea mays L. — Fig. 143.

Maize, Jagung

Gilliland 1971: 304. Keng et al. 1998: 187.

Culms single, erect, 100—200 cm long; internodes solid; nodes glabrous. Sheath at least 14

cm long, rounded, hairy, margin hairy. Ligule membranous, 4—5 mm long, tip ciliolate. Young

leaf blade inrolled. Blade 47 cm long, 50 mm wide, sparsely hairy at upperside, base rounded.

Male inflorescence terminal, a panicle, c. 25 cm long, central rachis hairy, branches 2—20;

longest branch 20—30 cm long, simple. Pedicel 1.5—3 mm long, hairy. Male spikelets single

or paired, 8.4 mm long, 3 mm wide, laterally flattened, 2-flowered with both florets male,

unawned; the two lowest bracts as long as spikelet; glumes thin herbaceous, lemmas and

paleas hyaline; lower glume 8.4 mm long, 7 veins, hairy in upper half, acute; upper glume 8.2

mm long, 7 veins, hairy, acute; 1‘ lemma 6.6 mm long; 1* palea 5.9 mm long; 2™ lemma, 5

mm long; 2™ palea, 5.7 mm long. Female inflorescence from lower sheath, enveloped by

several bracts; female spikelets in rows on stout central rachis.

Not native; probably from Mexico, cultivated worldwide. First record 1920. Cultivated.

In America this species is called Corn, whereas in English corn is used for all sorts of

grains, especially wheats, oats and maize.

Zizania L.

Zizania latifolia (Griseb.) Stapf — Fig. 148.

Manchurian Water Rice

Gilliland 1971: 104. Veldkamp 1997b: 512. Keng et al. 1998: 188.

Culms single, erect, 110—180 cm long; internodes 15 mm diameter; nodes glabrous. Sheath

c. 30 cm long, rounded, with transverse veins, glabrous. Ligule membranous, 7—11 mm

long. Young leaf blade inrolled. Blade 40—80 cm long, 19—27 mm wide, with transverse

veins present, glabrous, base cuneate. Inflorescence panicle, 40—60 cm long, central rachis

glabrous, branches many; longest branch 6—7 cm long, 0.3 mm wide, glabrous, branched.

Pedicel 1—2 mm long, glabrous, tip cup-shaped. Spikelets single, heteromorphous, more or

less laterally flattened, 1-flowered, unisexual, awned. Female spikelet in upper half of panicle,

c. 50 mm long (including awn), 1.5 mm wide; glumes absent; lemma c. 50 mm long including

c. 26 mm-long awn, herbaceous, glabrous, 5 veins, acuminate; palea as lemma, c. 24 mm

long, 3 veins. Male spikelet in lower half of panicle, as female, c. 16 mm long (including c. 5

mm-long awn); palea c. 12 mm long.

Not native; India to Japan, introduced in Europe and Australasia. First record 1941.

Cultivated and escaped (until 1950).

Aquatic, in wet or muddy places.

It was cultivated for its young shoots that are eaten as a vegetable. According to Sinclair

(SF 40615, 1955) it was commonly planted, but did not flower. Today it is probably no longer

grown.

Zoysia Willd.

Zoysia matrella (L.) Merr. — Fig. 149, Plate 40.

Siglap Grass, Rumput Zoysia

Gilliland 1971: 109. Keng et al. 1998: 188.

Z. pungens Willd.: Ridley 1907: 149; 1925: 243.

Culms decumbent or with long stolons, erect part 5—40 cm long; nodes glabrous. Sheath 1—

1.6 cm long, rounded, glabrous, margin glabrous or hairy at throat. Ligule membranous, 0.2

mm long, tip ciliolate, with long hairs on blade behind the ligule. Young leaf blade inrolled.

142 Supplement of Gardens’ Bulletin Singapore 57 (2005)

Blade 1.3—9 cm long, 1—2.5 mm wide, inrolled to flat, glabrous or sparsely hairy at upperside,

base cuneate and hairy just behind the ligule. Inflorescence raceme; rachis 1—3.5 cm long,

glabrous, with spikelets crowded. Pedicel 0.8—3.3 mm long, glabrous. Spikelets single, 3—

3.4 mm long (including awn), 0.6—0.8 mm wide, laterally flattened, glabrous, 1-flowered,

(minutely) awned; either lowest bract 0.05—0.3 times as long as and 2 as long as spikelet,

or lowest as long as and 2"! 0.6—0.7 times as long as spikelet; lower glume absent or 0.2—1

mm long, herbaceous, | vein; upper glume 3—3.4 mm long, indurated, 1—7 veins, midvein

slightly keeled, apiculate to awned, awn 0).3—0.9 mm long; lemma 2—2.2 mm long, hyaline,

1 vein, truncate to obtuse; palea absent or 0.1—1.5 mm long, hyaline.

Native; (sub)tropical Asia and Australia. First record 1878. Common.

In lawns and on roadsides with well-drained (sandy) soil.

In America this species is called Zoysia Grass.

Cynodon dactylon is very similar in appearance and can be found in similar habitats,

but differs in having a ligule of hairs of unequal length, an inflorescence with 2 or more

branches clustered at the tip of the peduncle and sessile spikelets of which the glumes are

often gaping.

A related species with leaf blades narrowly inrolled and 0.7—1 mm diameter and the

peduncle, even in fruit, not or hardly exserted from the uppermost sheath has been described

as Z. pacifica (Goudswaard) M.Hotta & Kuroki (1994). It forms a dense, short turf and is

widely cultivated as a lawn grass or in tennis lawns and golf greens. It is expected to occur in

Singapore.

Key to Sterile Grass Plants of Singapore

Note: species that are thought to be extinct or known from cultivation only are not included

in this key (except for a few that have perhaps been overlooked and/or seem to flower rarely).

la. Young blade either folded along midrib or plicate, or blades less than 1

mm wide. 2

1b. Young blade inrolled, blades at least | mm wide. 9

2a. Blades plicate. Setaria barbata

2b. Blades either flat or folded one midrib, or less than 1 mm wide. 3

3a. Ligule a row of free hairs. *

3b. Ligule membraneous, glabrous or hairy. 5

4a. Culms with long rhizomes or stolons, rarely tufted and stolons absent.

Blade 1.5—2.5 mm wide. Cynodon dactylon

Figure 147. Urochloa glumaris (Trin.) Veldk.

a. lateral view, b. facing upper glume.

Figure 148. Zizania latifolia (Griseb.) Stapf.

a. male spikelet, b. female spikelet.

Figure 149. Zoysia matrella (L.) Merr., upper glume lateral view, lemma and palea hidden

inside.

143

Field Guide to the Grasses of Singapore (Excluding the Bamboos)

144 Supplement of Gardens’ Bulletin Singapore 57 (2005)

4b. Culms tufted, erect, stolons absent. Blade c. 0.5 mm wide.

Eriachne pallescens

5a. Stolons absent. 6

5b. Stolons present. 7

6a. Sheath glabrous, margins hairy at least at tip. Eleusine indica

6b. Sheath at least in upper half with bulbous-based hairs, margins glabrous or

hairy in basal half. Themeda arguens

7a. Sheath rounded, midrib not raised. Blade 0.6—1.7(—2) mm wide.

Digitaria didactyla

7b. Sheath flattened, midrib raised. Blade 3—7 mm wide. 8

8a. Leaves glossy and (dark) green; sheath with margins in upper half glabrous

or glabrescent; blade hardly wider than sheath. Axonopus fissifolius

8b. Leaves glaucous; sheath with margins densely hairy in upper half; blade

distinctly wider than sheath. Stenotaphrum secundatum

Ya. (1) Leaf blade with transverse veins present, 8.5—-70 mm wide. 10

9b. Leaf blade without transverse veins, 1—60 mm wide. 13

10a. Leaf blade with pseudopetiole 0.5—3 mm long. 11

10b. Leaf blade with pseudopetiole 7—50 mm long. 12

(10c. Leaf blade without pseudopetiole, margins serrate. Cultivated. Chrysopogon zizanioides )

lla. Internodes up to 2.5 mm diameter. Blade 3.5—11 cm long, 13—28 mm

wide, hairy at least at upperside. Centotheca lappacea

11b. Internodes at least 3 mm diameter. Blade 40—63 cm long, 28—70 mm

wide, glabrous or pseudopetiole hairy. Thysanolaena latifolia

12a. Blade 8.5—-22 mm wide, pseudopetiole 7—20 mm long.

Lophatherum gracile

12b. Blade 40—62 mm wide, pseudopetiole 20—50 mm long.

Scrotochloa urceolata

13a. (9) Ligule absent. Echinochloa colona, E. crus-galli

13b. Ligule present, either membranous or as a shallow rim, or as a row of

hairs. 14

14a. Ligule 0.5—4 mm long, either of free hairs, or hairs at tip at least as long

as the membranous basal part. 15

14b. Ligule 0.05—7.5 mm long,—f longer than 0.5 mm then membranous

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 145

with hairs at tip absent or shorter than basal part. 30

15a. Ligule hairs free. 16

15b. Ligule either hairs fused at base, or basal part membranous. 22

16a. Nodes hairy. 17

16b. Nodes glabrous. 20

17a. Sheath 0.5—3 cm long. 18

17b. Sheath 6—17 cm long. 19

18a. Ligule 1—1.6 mm long. Blade slender, underside with 3—9 prominent

and well-spaced veins, base rounded. y Isachne

18b. Ligule 0.5—1 mm long. Blade stiffened, underside narrowly and densely

ribbed (when dry), base cuneate. Thuarea involuta

19a. Lower sheaths glabrous. Eriochloa procera

19b. Lower sheaths densely hairy. Melinis repens

20a. (16) Blade 1.5—2.5 mm wide. Cynodon dactylon

20b. Blade 3—7 mm wide. 21

21a. Sheath more or less flattened, midrib (slightly) raised, margins glabrous

(hairs submarginal if present). Ligule 0.7—1 mm long. Blade 7—26 cm

long. Cenchrus

21b. Sheath rounded, midrib not raised, margins hairy. Ligule |—4 mm long.

Blade 1—7.8 cm long. Isachne

22a. (15) Blade at base cordate. Alloteropsis cimicina

22b. Blade at base cuneate to rounded. 23

23a. Blade 2.5—5.5 cm long, 3.8—9 times longer than wide. Oplismenus

23b. Blade 5—65 cm long, at least 12 times longer than wide. 24

24a. Basal internodes at least 5 mm diameter. Midvein of blade at least at base

prominent at upper side, white and broad. 25

24b. Basal internodes up to 3 mm diameter. Midvein of blade not prominent at

upper side, green and/or narrow. 2/7

25a. Nodes densely patent hairy. Urochloa mutica

25b. Nodes glabrous or with appressed hairs. 26

26a. External ligule either as a herbaceous rim or absent (but sheath and

146 Supplement of Gardens’ Bulletin Singapore 57 (2005)

underside blade may be hairy). Pennisetum

26b. External ligule present as a fringe of hairs. Sorghum propinquum

27a. Sheath rounded, midrib not raised. 28

27b. Sheath at least at tip more or less flattened, midrib (slightly) raised. 29

28a. Nodes glabrous. Panicum paludosum

28b. Nodes hairy. Urochloa glumaris

29a. Blade at base distinctly rounded. Cenchrus, Urochloa

29b. Blade at base cuneate to slightly rounded. Setaria parviflora

30a. (14) Stolons absent (but rhizomes may be present). Leaves all basal, culm

elongating only upon flowering. 31

30b. Stolons absent or present. Leaves scattered along culm, not clustered at

the base. 34

31a. Leaves strictly distichous. Sheaths distinctly flattened. 32

31b. Leaves irregularly arranged. Sheaths rounded. 33

32a. Sheaths 2.5—7.5 cm long. Ligule c. 0.5 mm long. Chloris barbata

32b. Sheaths 12—23 cm long. Ligule 1—1.8 mm long. Themeda villosa

33a. Rhizomes present. Blade 6—22 mm wide. Imperata

33b. Rhizomes absent. Blade 2—4.5 mm wide. Sporobolus indicus

(33c. Rhizomes absent. Blade 8—20 mm wide, margins sharply serrate. Cultivated.

Cymbopogon)

34a. (30) Blade with pseudopetiole (0.7—)1—4 mm long. 35

34b. Blade without pseudopetiole. 38

35a. Culm erect or geniculate, not scrambling. Sheath 10—17 cm long. Ligule

2—7.5 mm long. Ischaemum magnum

35b. Culm generally creeping or scrambling. Sheath 1.2—6.5 cm long. Ligule

0.4—0.7 mm long. - 36

36a. Sheath more or less flattened, midrib raised. Auricles fused with ligule,

0.3—1 mm long. Ischaemum muticum

36b. Sheath rounded, midrib raised or not. Auricles absent. 37

37a. Nodes, sheaths and blades glabrous or hairy. Blades not glaucous.

Ottochloa nodosa

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 147

37b. Nodes glabrous. Sheath with a few hairs at tip only. Blades glaucous,

hairy. Panicum trichocladum

38a. (34) Ligule 0.2—4 mm long, membraneous, if 0.2—0.5 mm long then

glabrous. 39

38b. Ligule 0.05—0.5 mm long,—either as a row of free hairs or membraneous

and hairy. 68

39a. Sheath (including margins and auricles) entirely glabrous. 40

39b. Sheath at least partly hairy. 50

40a. Nodes glabrous. 41

40b. Nodes (sparsely) hairy. Z 45

41a. Internodes 5—30 mm diameter. Blades 50—200 cm long. Saccharum

41b. Internodes up to 3 mm diameter. Blades 1.5—20(—50) cmlong. 42

42a. Blades stiff, upperside with flat ribs. Lepturus repens var. repens

42b. Blades slender, upperside without ribs (or minutely ribbed when dry).

43a. Stolons absent or present. Sheath rounded. Digitaria

43b. Stolons absent. Sheath more or less flattened. da

44a. Rhizomes absent. Ligule glabrous. Paspalum

44b. Rhizomes present, short. Ligule with hairs at tip. | Setaria parviflora

45a. (40) Ligule hairy at abaxial side. to

45b. Ligule glabrous at abaxial side. — 47

46a. Sheath flattened. Blade glabrous or hairy at base only, midvein winged

underneath. Leptochloa chinensis

46b. Sheath rounded. Blade hairy at least at upperside, midvein not winged.

Bothriochloa bladhii, Dichanthium annulatum

47a. Hairs on nodes deflexed. Blade with margins serrulate.

Leersia hexandra

47b. Hairs on nodes appressed to patent. Blade with margins smooth to

scaberulous. 48

48a. Stolons present. Digitaria

48b. Stolons absent. 49

148 Supplement of Gardens’ Bulletin Singapore 57 (2005)

49a. Blade 2—9 mm wide. Ligule 1—3 mm long. _ Digitaria

49b. Blade 1—2 mm wide. Ligule 0.4—0.7 mm long. Dimeria ornithopoda

50a. (39) Nodes hairy. 51

50b. Nodes glabrous. 57

Sla. Sheath 0.5—2(—4) cm long. Ligule glabrous to ciliolate at tip, never

hairy at abaxial side. 52

51b. Sheath 2—34 cm long, if shorter than 4 cm, then ligule hairy at abaxial

side. 54

52a. Sheath with margins hairy. Cyrtococcum patens

52b. Sheath with margins glabrous (or submarginal hairs present). 53

53a. Stolons present. Digitaria

53b. Stolons absent, culm erect to decumbent. Dimeria ornithopoda

54a. Ligule glabrous. Digitaria setigera

54b. Ligule hairy at tip or abaxial side, or with a row of hairs at abaxial side.

55a. Auricles present, grown with the margins of the ligule.

Ischaemum ciliare, I. timorense

55b. Auricles absent. 56

56a. Culm erect to geniculate, stolons absent. Sheath (excluding basal bladeless

sheaths) 10—34 cm long. Blade (9—)25—50 cm long.

Panicum maximum

56b. Culm decumbent or with stolons. Sheath 4—7 cm long. Blade 5—16 cm

long. Urochloa glumaris

57a. (50) Basal internodes 15—45 mm diameter. Root-eyes obvious on lower

nodes, in 2—9 rows. Saccharum officinarum

57b. Basal internodes 3—10 mm diameter. Root-eyes absent.

58a. Ligule densely fringed or hairy. Ss

58b. Ligule entire (or torn once or twice) and glabrous. 62

59a. Culm erect, stolons absent. Blade 10—25 mm wide. 60

59b. Culm erect, ascending or decumbent, stolons absent or present. Blade

46 mm wide. 61

60a. Sheath with soft hairs at tip only. Blade with soft hairs at base only.

Mnesithea glandulosa

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 149

60b.

61a.

61b.

62a.

62b.

63a.

63b.

64a.

Sheath and blade upperside with stiff hairs. Rottboellia cochinchinensis

Sheath without transverse veins, margins glabrous (except for few long

hairs at tip). Blade at base rounded, margins hairy.

Dactyloctenium aegyptium

Sheath with transverse veins present, one margin hairy. Blade at base

cuneate, margins glabrous. Panicum repens

(58) Stolons absent. 63

Stolons present. 65

Sheath more or less flattened, midrib raised. Paspalum

Sheath rounded, midrib raised or not. 64

Internodes up to 2 mm diameter. Sheath without transverse veins.

Digitaria

64b. Internodes c. 3 mm diameter. Sheath with (minute) transverse veins present.

65a.

65b.

66a.

66b.

67a.

67b.

68a.

68b.

69a.

69b.

70a.

70b.

TVa

71b.

12h

72b.

Hymenachne amplexicaule

External ligule hairy. Paspalum conjugatum

External ligule absent. 66

Stolons 2.5—3 mm wide. Blade at base cuneate, much narrower than

sheath. Paspalum vaginatum

Stolons or basal part of creeping culm up to 2 mm wide. Blade at base

(slightly) rounded, narrower or up to as wide as sheath. 67

Sheath not shouldered. Cyrtococcum accrescens

Sheath shouldered. Digitaria

(38) External ligule hairy. i 69

External ligule absent. 70

Stolons present. Nodes hairy. Axonopus compressus

Stolons absent. Nodes glabrous. Eragrostis

Sheath hairy. vel

Sheath glabrous (or hairy at margins only). 72

Sheath 0.7—2(—4) cm long. Blade 1—2 mm wide. Dimeria ornithopoda

Sheath 4.2—6.5 cm long. Blade 3—5 mm wide. —-Urochloa glumaris

Sheath with (minute) transverse veins present. Panicum laxum, P. repens

Sheath without transverse veins. 73

] 50 Supplement of Gardens’ Bulletin Singapore 57 (2005)

73a. Stolons present. 74

73b. Stolons absent (but culms may be decumbent). 78

74a. Sheath (2—)4—6 cm long. Blade with distinct flat ribs at upper side,

stiff. (beach grasses). a5

74b. Sheath 1—4.5 cm long. Blade not ribbed at upper side (or only minutely

when dry), (rather) slender. 76

75a. Stolon culm 1—1.5 mm wide. Sheath glabrous.

Lepturus repens var. repens

75b. Stolon culm c. 3 mm wide. Sheath hairy, at least the margin at tip.

Sporobolus virginicus

76a. Stolons with leaves densely crowded. Blade 4—6 mm wide, margins

hairy at base. Chrysopogon aciculatus

76b. Stolons with leaves well-spaced. Blade |_—6 mm wide, margins glabrous.

77a. Blade 2—6 mm wide, base (somewhat) rounded, glabrous.

Sacciolepis indica

77b. Blade 1—2.5 mm wide, base cuneate, hairy just behind ligule.

Zoysia matrella

78a. (73) Blade 20—40 mm wide. Coix lacryma-jobi

78b. Blade 1—7 mm wide. 79

79a. Nodes hairy. 80

79b. Nodes glabrous. 81

80a. Sheath margins glabrous. Dimeria ornithopoda

80b. Sheath with one margin hairy. Pogonatherum crinitum

81a. Blade at base cordate to amplexicaul. Perotis indica

81b. Blade at base cuneate to rounded. $2

82a. Culms erect. Sporobolus tenuissimus

82b. Culms geniculate to decumbent. 83

83a. Blade 2—6 mm wide, base somewhat rounded, glabrous. Sacciolepis

83b. Blade 1—2.5 mm wide, base cuneate, hairy just behind ligule. Zoysia

Field Guide to the Grasses of Singapore (Excluding the Bamboos) 151

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Acknowledgements

Several people have in one way or another been involved in this project. First

of all, Dr S.C. Chin (Singapore Botanic Gardens), Dr R. Kiew and staff of the

SING herbarium and Prof. H.T.W.Tan, Ms. Morgany, and Mr. Chua of the

SINU herbarium are gratefully acknowledged. The collections present in the

two herbaria form the basis of this work and I enjoyed working there. Additional

collections have been made along the streets of Singapore (often in front of

very surprised citizens) and on several off-street field trips. It was a great

pleasure to be in the field with Tan Kai Xin, Prof. H.T.W. Tan, Ms. Morgany

(all SINU), Serena Lee, Paul K.FK. Leong, Gwee Aik Teck, Haji Samsuri Ahmad

(all SING), Joseph Lai, Ali Ibrahim (NParks) and Keith Hillier. I would like to

thank Mohd Noor Jumaat for his assistance with the preparation of the

herbarium material. The enthusiastic support of Dr J.F. Veldkamp (L), my

156 Supplement of Gardens’ Bulletin Singapore 57 (2005)

early tutor in the grasses, cannot go unmentioned. I am very grateful for his

updated keys for Flora Malesiana, and for the confirmation of the identification

of Isachne pulchella (collected by Paul K.F. Leong), Panicum laxum,

Sporobolus indicus var. pyramidalis and S. tenuissimus. Hassan Ibrahim, Paul

K.F. Leong, Serena Lee (SING) and Mr. Chua (SINU) are greatly acknowledged

for trying out both identification keys and Hassan Ibrahim and Haji Samsuri

Ahmad for help with the Malay names. The comments of the editor Dr R.

Kiew and the reviewers Dr J.F. Veldkamp and Dr S.C. Chin have been very

helpful in improving the manuscript. Dr Wan Mohamad Wan Othman

(University Sabah Malaysia) is acknowledged for his help with Malaysian

turf grasses. My sketches of the spikelets were redrawn by Dr Jaap J. Vermeulen.

The other line drawings and part of the photographs are from his hand as well.

I am very grateful for his help and for his assistance with photo-shopping

them; the illustrations make this guide so much more attractive and readable.

Last, but not least I would like to thank my husband for his moral support and

our children Judith, Max and Jos for their patience. It must have been difficult

at times to cope with my preoccupation on grasses.

Field Guide to the Grasses of Singapore (Excluding the Bamboos) L57

Index

(Accepted scientific names are in bold print. Bold page numbers refer to descriptions.

Names may appear more than once on a page. The Malay follows the new spelling

where ‘c’ is pronounced as ‘ch’ is in English.)

abaxial 6,21 bisexual 8

Acroceras 32 blade 4

Acroceras munroanum 32 ~~ Blue Couch 55

Acroceras sparsum 32. ~——-Blyxa alternifolia 18

Acroceras tonkinense 32 Blyxa auberti 18

acuminate 19 Bothriochloa 36

acute 19 Bothriochloa bladhii 36

adaxial 6,21 Bothriochloa intermedia ~ 36

Alloteropsis 34 — Brachiaria distachya 140

Alloteropsis cimicina 34. ~—- Brachiaria mutica 138

Amphilophis glabra 36 ~—s- Brachiaria paspaloides 138

amplexicaul 19 Brachiaria piligera 140

Andropogon aciculatus 42 Brachiaria subquadripara 140

Andropogon contortus 76 bract 10

Andropogon hirtiflorus 120 Bristle-grass, Broad-leaved 126

Andropogon intermedius 36 _ bristles ie

annual 12 Brown-top Grass 114

annular 19 Buffalo Grass 16, 106

Anthistiria arguens 134 = Buluh Tebrau 135

antrorse 19 . bur 10

Arundo 34 caducous 19

Arundo donax 34 callus 1]

Arundo donax cv versicolor 34 — Cane, India 118

ascending 19 ~~ Cane, Sugar 118

auricle 4 Cane, Wild 118

awn 10 Carpet Grass 16, 36

Axonopus 34,115 cartilaginous 19

Axonopus affinis 36 —_—caryopsis I]

Axonopus cimicinus 34 ~— Cat’s Tail Grass 112

Axonopus compressus 3, 15, 16, 36, 42, 106 —cataphyll 6

Axonopus fissifolius 16,36 Cenchrus WR ir

Bamboo Grass 114 = Cenchrus brownii 37

Barbed Grass 38 = Cenchrus echinatus 37, 38

Beard Grass, Long-leaved 36 Centipede Grass L540

Bermuda Grass 16,44 Centipede Grass, Common 86

bifid 19 Centipede Grass, Seashore 84

Bird’s Foot Grass 61 Centipede Grass, Wrinkled 86

158

Centotheca 12, 38

Centotheca lappacea 38

Centotheca latifolia 38

Centotheca longilamina 38

Chloris 40

Chloris barbata 10, 40, 76

Chrysopogon 40

Chrysopogon aciculatus 16, 42

Chrysopogon zizanioides 42

ciliate 6, 19

ciliolate 6, 19

Citronella Grass 44

cleistogamous 8

Cockspur 62

Coelorachis glandulosa 92

Coix 42

Coix lacryma-jobi 42

collar 6

collar-shaped 19

cordate 19

Corn 14]

Cow Grass 36

Crab Grass, Common 54

Crab Grass, Lesser 55

Crested Grass 89

culm 3

cuneate 19

Cymbopogon 43

Cymbopogon calcicola 44

Cymbopogon citratus — 43

Cymbopogon flexuosus 44

Cymbopogon martini 43

Cymbopogon nardus var. nardus 44

Cynodon 44

Cynodon arcuatus 44

Cynodon dactylon 15, 16, 44, 74, 76, 142

Cynodon radiatus 44, 46

Cyperaceae 17

Cyrtococcum 10, 46, 80

Cyrtococcum accrescens 46

Cyrtococcum carinatum 48

Cyrtococcum oxyphyllum 46

Cyrtococcum patens 46, 48

Supplement of Gardens’ Bulletin Singapore 57 (2005)

Dactyloctenium 48

Dactyloctenium aegyptium 10, 48

Daucus carota 3

decumbent 19

Desmodium triflorum 15

Desmodium, Lesser Clover-leaved 15

diaspore 12

Dichanthium 48

Dichanthium annulatum 37, 50

Dichanthium caricosum 50

Dichanthium mucronulatum 48, 50

Digitaria 51

Digitaria bicornis 10, 54

Digitaria caespitosa 60)

Digitaria chinensis 60)

Digitaria ciliaris 54

Digitaria didactyla 15, 16, 55

Digitaria fuscescens 55

Digitaria longiflora 55, 56, 60, 61

Digitaria marginata 54

Digitaria marginata vat. ciliaris 54, 60

Digitaria marginata var. commutata 58

Digitaria marginata var. debilis 55, 58

Digitaria marginata var. pruriens 58

Digitaria microbachne 58

Digitaria microbachne subsp.

calliblepharata 60)

Digitaria microstachya 58

Digitaria mollicoma 56, 61

Digitaria nuda 54, 56

Digitaria pertenuis 55, 56, 60

Digitaria radicosa 58

Digitaria setigera 58

Digitaria setigera var. calliblepharata 60

Digitaria setigera var. setigera «60

Digitaria timorensis 58

Digitaria violascens 56, 60

Dimeria 61

Dimeria glabra 61

Dimeria ornithopoda 61

Dimeria ornithopoda var. glabra 6]

Dimeria ornithopoda var. ornithopoda 61

distichous 4

Field Guide to the Grasses of Singapore (Excluding the Bamboos)

Dropseed, Beach 131

Dropseed, Common 130

Dropseed, Lesser 130

Echinochloa 61

Echinochloa colona 6, 62

Echinochloa crus-galli 62

Echinochloa picta 61, 62

Echinochloa stagnina 62, 64

Elephant Grass 110

Eleusine 11, 64

Eleusine aegyptia 48

Eleusine aegyptiaca 48

Eleusine coracana 64

Eleusine indica 64

Eleusine indica var. coracana 64

ellipsoid 19

elliptic 19

emarginate 19

endosperm 12

Eragrostis 8, 10, 12, 14, 65

Eragrostis amabilis 66, 71, 131

Eragrostis atrovirens 65, 66

Eragrostis brownii 65, 68

Eragrostis cilianensis 68

Eragrostis cumingii 68

Eragrostis cumingii var. cumingii 68

Eragrostis diandra 65

Eragrostis elegantula 66, 68

Eragrostis elongata 65, 66, 68

Eragrostis gangetica 70

Eragrostis malayana 710

Eragrostis montana 70

Eragrostis nutans 65

Eragrostis pilosa 70

Eragrostis tenella 66

Eragrostis unioloides 71

Eragrostis viscosa 66

Eragrostis, Feathery 66

Eragrostis, Pink 71

erect 19

Eremochloa ciliaris 2

Eremochloa ophiuroides 15, 16

Eriachne

Eriachne pallescens

Eriachne triseta

Eriachne, Slender

Eriocaulon longifolium

Eriocaulon truncatum

Eriochloa

Eriochloa annulata

Eriochloa meyerana

Eriochloa polystachya

Eriochloa procera

Eriochloa villosa

Eulalia praemorsa

Eustachys

Eustachys tenera

extravaginal

female

Finger Grass, Egyptian

flattened

flattened, dorsiventrally

flattened, laterally

flattened, not

floret

flower

fruit

fusiform

Gama Grass

geniculate

gibbous

Ginger Grass

glabrescent

glabrous

glaucous

globose

glume, lower

glume, upper

Goose Grass

granulate

Guinea Grass

Hackelochloa granularis

hairy

Hemigymnia fusca

Hemigymnia multinodis

herbaceous

159

160

heteromorphous

Heteropogon

Heteropogon contortus

hilum

homomorphous

hyaline

Hymenachne

Hymenachne acutigluma

Hymenachne amplexicaulis

Hymenachne myuros

Imperata

Imperata arundinacea

Imperata conferta

Imperata contracta

Imperata cylindrica

Imperata cylindrica var. maior

Imperata exaltata

indument

indurated

inflorescence

internode

intravaginal

involucre

Isachne

Isachne australis

Isachne globosa

Isachne kunthiana

Isachne miliacea

Isachne minutula

Isachne pulchella

Isachne semitalis

Isachne, Rounded

Ischaemum

Ischaemum aristatum

Ischaemum barbatum

Ischaemum beccarii

Ischaemum ciliare

Ischaemum indicum

Ischaemum laeve

Ischaemum macrurum

Ischaemum magnum

Ischaemum muticum

Ischaemum rugosum

Supplement of Gardens’ Bulletin Singapore 57 (2005)

Ischaemum timorense

Jagung

Jelai

Job’s Tears

Jungle Rice

keeled

Kemucut

Lalang

lanceolate

leaf, culm-

leaf, flag

leaf, foliage

Leersia

Leersia hexandra

lemma

Lemon Grass

Leptaspis

Leptaspis urceolata

Leptochloa

Leptochloa chinensis

Lepturus

Lepturus repens

Lepturus repens var. repens

ligule

ligule, external

linear

lodicule

Lophatherum

Lophatherum gracile

Love Grass

Love Grass, Indian

Maize

Malabar Grass

male

Manisuris granularis

Massia triseta

Mat Grass

Melinis

Melinis repens

membranous

Millet, African

Millet, Barnyard

83, 84, 86

141

Field Guide to the Grasses of Singapore (Excluding the Bamboos)

Millet, Great 127

Millet, Italian 124

Millet, Kodo 107

Miscanthus T1

Miscanthus floridulus 2, 118

Miscanthus sinensis 2

Mnesithea 12, 90, 115

Mnesithea glandulosa 92,115

Mnesithea granularis 92

Mnesithea laevis” 90)

mucronate 20

Murdannia nudiflora 18

Murdannia vaginata 18

Napier Grass 110

Natal Grass 89

Neyraudia 771, 94

Neyraudia arundinacea var. zollingeri 94

Neyraudia madagascariensis var. zollingeri 94

Neyraudia reynaudiana 94

node 3

nodular 20

nodule 20

notch 10

Nut-grass, Sea 134

oblong 20

obtuse 20

Oplismenus 94

Oplismenus burmanni 94

Oplismenus compositus 95

Oplismenus compositus var. rariflorus 95

orbicular 20

Oryza 95

Oryza rufipogon S

Oryza sativa 95

Ottochloa 96

Ottochloa nodosa 96

ovate 20

ovoid 20

Padi 95

Padi Burung 2, 62

palea 8

Panic Grass, Creeping 17, 103

Panic Grass, Diffuse 46

161

Panic Grass, Giant 99

Panic Grass, Scrambling 103

Panic Grass, Short-leaved 99

Panic Grass, Slender 96

Panic Grass, Swamp 76

panicle I]

Panicum 96, 98

Panicum auritum TT, 99, 119

Panicum brevifolium 99

Panicum caesium 100

Panicum cambogiense 100

Panicum colonum 62

Panicum crus-galli 62

Panicum distachyum - 138, 140

Panicum hirtifolium 99

Panicum indicum 119

Panicum laxum 98, 100

Panicum luzonense 100

Panicum maximum 10, 98, 102

Panicum muticum 138

Panicum myosuroides 120

Panicum myurus T]

Panicum nodosum 96

Panicum oryzoides 32

Panicum ovalifolium 99

Panicum paludosum 102

Panicum patens 46

Panicum plicata 126

Panicum proliferum 102

Panicum repens 17, 74, 103, 140

Panicum sarmentosum 103

Panicum trichocladum 98, 103

Para Grass 138

parallel-nerved =

Paspalum 104

Paspalum cartilagineum 107

Paspalum commersonii 107

Paspalum conjugatum 6, 16, 104, 106

Paspalum dilatatum 104

Paspalum distichum 108

Paspalum longiflorum 55, 56, 60

Paspalum longifolium 106

Paspalum orbiculare 107

162

Paspalum platycaulon 36

Paspalum platycoleum 106

Paspalum plicatulum 107

Paspalum sanguinale 54

Paspalum sanguinale var. ciliare 54, 60)

Paspalum sanguinale var. commutatum 54, 58

Paspalum sanguinale var. debile 55, 58

Paspalum sanguinale var. extensum 58

Paspalum sanguinale var. pruriens 58

Paspalum scrobiculatum 107

Paspalum scrobiculatum var. bispicatum —_ 107

Paspalum vaginatum 108

Paspalum, Saltwater 108

patent 20

pectinate 20

pedicel 10

peduncle 1]

Pennisetum 12, 37, 108

Pennisetum glaucum 108

Pennisetum macrostachyum 108

Pennisetum polystachion 108, 110

Pennisetum purpureum 2, 110

Pennisetum setaceum 108

Pennisetum, Feather 110

perennial 12

Perotis 110

Perotis indica 112

Perotis latifolia 112

Philydrum lanuginosum 18

Phragmites 77, 94, 112

Phragmites communis 112

Phragmites karka 112

Phragmites vallatoria 112

pinninerved -

Plush Grass 40

Pogonatherum 114

Pogonatherum crinitum 114

Pogonatherum paniceum 114

Pogonatherum saccharoideum var.

monandra 114

Polytrias 114

Polytrias amaura 114

Polytrias indica 114

Supplement of Gardens’ Bulletin Singapore 57 (2005)

Polytrias praemorsa

prophyll

pseudopetiolate

punctate

raceme

raceme, spike-like

rachilla

rachis

rachis, central

Reed, Common

reticulate

rhizome

Rhynchelytrum repens

Rice

Rice Grass

Rice, Jungle

Rice, Manchurian Water

Ripidium

Ripidium arundinaceum

root-eye

Rottboellia

Rottboellia cochinchinensis

Rottboellia exaltata

Rottboellia glandulosa

Roundseed, Elegant

rugose

rugosity

rugulose

Rumput Babi

Rumput Benggala

Rumput Bermuda

Rumput Centipede

Rumput Darah

Rumput Dawai

Rumput Ekor Cawi

Rumput Ekor Kucing

Rumput Gedabung

Rumput Janggut Ali

Rumput Jarang

Rumput Kerbau

Rumput Kolam Padang

Rumput Kumpai

Rumput Lipan

12, 90, 115

12, 90, 92, 115

Field Guide to the Grasses of Singapore (Excluding the Bamboos)

Rumput Minyak

Rumput Misai Adam

Rumput Tikar

Rumput Riong

Rumput Sambau

Rumput Sembor Batu

Rumput Serangoon

Rumput St. Augustine

Rumput Telur Ikan

Rumput Telur Kutu

Rumput Tembaga Jantan

Rumput Wangi

Rumput Zoysia

Saccharum

Saccharum arundinaceum

Saccharum officinarum

Saccharum robustum

Saccharum sinense

Saccharum spontaneum

Sacciolepis

Sacciolepis angusta

Sacciolepis indica

Sacciolepis indica var. indica

Sacciolepis indica var. turgida

Sacciolepis myosuroides

Sacciolepis turgida

Sacciolepis, Long-spiked

Sacciolepis, Short-spiked

scaberulous

scabrous

Schizachyrium

Schizachyrium brevifolium

Schizachyrium sanguineum

Schizachyrium semiberbe

scrambling

Scrotochloa

Scrotochloa urceolata

sedge

Sekoi

Serai

Serai Wangi

Serangoon Grass

serrate

141

aie

116, 118

118

116, 118

76, 99, 119

119

77, 119

119

119, 120

119

120

119

17, 18

serrulate

setaceous

setae

Setaria

Setaria barbata

Setaria glauca

Setaria italica

Setaria pallidifusca

Setaria palmifolia

Setaria palmifolia var. palmifolia

Setaria parviflora

Setaria plicata

Setaria pumila

Setaria rubiginosa

Setaria spacelata

setulose

sheath

sheath, final

Shield Grass

shouldered

Siglap Grass

smooth

Smut Grass

Sorghum

Sorghum affine

Sorghum bicolor

Sorghum propinquum

Sorghum vulgare

spathe

spatheate

Sphaerocaryum

Sphaerocaryum elegans

Sphaerocaryum malaccense

spike

spikelet

Sporobolus

Sporobolus diander

Sporobolus diandrus

Sporobolus fertilis

Sporobolus indicus

Sporobolus indicus var. flaccidus

Sporobolus indicus var. major

163

126

ot)

126

122

130

Sporobolus indicus var. pyramidalis 131

164

Sporobolus tenuissimus

Sporobolus virginicus

St. Augustine Grass

Steinchisma laxa

Stenotaphrum

Stenotaphrum dimidiatum

Stenotaphrum secundatum

sterile

stolon

striate

subglobose

Tangle-head

Tasselgrass, Greater

Tasselgrass, Lesser

Tebu

Tebu Salah

Tebu Tikus

terete

Themeda

Themeda arguens

Themeda villosa

throat

Thuarea

Thuarea involuta

Thuarea sarmentosa

Thysanolaena

Thysanolaena agrostis

Thysanolaena latifolia

Thysanolaena maxima

Tiger Grass

Torpedo Grass

Tripsacum

Tripsacum dactyloides

triquetrous

Triraphis madagascariensis

truncate

undulate

unisexual

Urochloa

Urochloa glumaris

Urochloa maxima

Urochloa mutica

Urochloa piligera

66, 131

131

16, 132

100

132

15, 16, 132

Supplement of Gardens’ Bulletin Singapore 57 (2005)

Urochloa subquadripara

Vetiver

Vetiveria

Vetiveria odorata

Vetiveria zizanioides

Water Rice, Manchurian

Wick Grass

winged

Wood Grass, Common

Xyris complanata

Xyris pauciflora

Zea

Zea mays

Zizania

Zizania latifolia

Zoysia

Zoysia Grass

Zoysia pacifica

Zoysia matrella

Zoysia pungens

74, 100, 140

136, 141

10, 141

14]

141

14]

142

15, 16, 46, 141

14]

168 Supplement of Gardens’ Bulletin Singapore 57 (2005)

Plates 6—12.

6. Chloris barbata Sw.

7. Cynodon dactylon (L.) Pers.

8. Digitaria radicosa (J.Presl) Miq.

9. Echinochloa colona (L.) Link.

10. Eragrostis amabilis (L.) Wight & Arn. ex Nees.

11. Eragrostis brownii (Kunth) Nees.

12. Eriochloa procera (Retz.) C.E.Hubb.

—~

Ke}

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>_>

—

)

omy

SS)

170) Supplement of Gardens’ Bulletin Singapore 57 (2005)

Plates 13—18.

13. Ischaemum ciliare Retz.

14. Ischaemum magnum Rendle.

15. Ischaemum muticum L.

16. Lepturus repens (G.Forst.) R.Br. var. repens.

17. Chrysopogon aciculatus (Retz.) Trin.

18. Panicum maximum Jacq.

Field Guide to the Grasses of Singapore (Excluding the Bamboos)

L7Z Supplement of Gardens’ Bulletin Singapore 57 (2005)

Plates 19—26.

19. Lophatherum gracile Brongn.

20. Melinis repens (Willd.) Zizka.

21. Panicum repens L.

22. Paspalum conjugatum P.J.Bergius.

23. Paspalum scrobiculatum L. var. bispicatum Hack.

24. Paspalum vaginatum Sw.

25. Pennisetum polystachion (L.) Schult.

26. Pogonatherum crinitum (Thunb. in Murray) Kunth.

Field Guide to the Grasses of Singapore (Excluding the Bamboos)

174 Supplement of Gardens’ Bulletin Singapore 57 (2005)

Plates 27—33.

27. Pennisetum purpureum Schumach.

28. Saccharum arundinaceum Retz. Culm node with | row of root-eyes.

29. Saccharum officinarum L. Culm node with 2 rows of root-eyes.

30. Dactyloctenium aegyptium (L.) Willd.

31. Sacciolepis indica (L.) Chase.

32. Setaria barbata (Lam.) Kunth , left inflorescence with spikelets fallen off.

33. Stenotaphrum secundatum (Walt.) Kuntze. a. vegetative part, b. detail of

inflorescence.

Field Guide to the Grasses of Singapore (Excluding the Bamboos)

5 Ngee oe

mA 2 Sal ;

prs ae

(iy

—

so hes,

nga 2°

176 Supplement of Gardens’ Bulletin Singapore 57 (2005)

Plates 34—40.

34. Sporobolus indicus (L.) R.Br. var. flaccidus (Roem. & Schult.) Veldk.

35. Sporobolus tenuissimus (Schrank) Kuntze.

36. Themeda villosa (Poir.) A.Camus.

37. Thuarea involuta (G.Forst.) R.Br. ex Roem. & Schult. Fruiting inflorescence

with incurved rachis.

38. Urochloa mutica (Forssk.) T.Q.Nguyen. a. inflorescence, b. detail of culm,

node and sheath.

39. Urochloa subquadripara (Trin.) R.D.Webster.

40. Zoysia matrella (L.) Merr.

—~

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—

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—

NY)

ras

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SS)

INSTRUCTIONS TO AUTHORS

The Gardens’ Bulletin publishes original research findings and reviews of progress in the fields of plant

taxonomy, horticulture and allied subjects. Contributions must be original and the material must not

have been submitted for publication elsewhere.

Authors should look at the layout of articles recently published in the journal to ensure that submitted

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margins on all sides. All pages should be numbered sequentially.

Titles and authors: The title should give a concise description of the contents of the article. It should

include the family name if a taxon name is included in the title. The names(s) and affiliation(s) of the

author(s) must be given below the title. A short running title should also be provided. Lengthy papers

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Abstract: An abstract of up to about 100 to 200 words should be provided. It should comprehensively

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Scientific names: The complete scientific name — genus, species, authority with family in parenthesis —

must be cited for every organism at the time of first mention.

Abbreviations: Standard abbreviations may be used in the text, but the full term should be given on the

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Literature citation: Citation in the text should take the form: King and Gamble (1886). If several papers

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When papers are by three or more authors they should be cited as King et al. (1886) in the text, but with

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Stone, B.C. 1994. Additional notes on the genus Glycosmis (Rutaceae). Gardens’

Bulletin Singapore. 46: 113-119.

References to books and monographs should be cited according to the following form:

Ridley, H.N. 1930. The Dispersal of Plants Throughout the World. L. Reeve,

Ashford, U.K.

For literature citations in taxonomic papers, the following style is required:

Medinilla alternifolia Blume, Mus. Bot. Ludg.-Bat. I:1 (1849) 19.

Sterculia acuminatissima Merr., Philip. J. Sci. 21 (1922) 524.

Offprints: Authors will be given 50 offprints gratis. Additional copies must be ordered and paid for in

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Manuscripts may be submitted in a PC-compatible electronic form, preferably as a single file

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a disc (floppy, CD-ROM or ZIP disc) and sent to the same aie a a | 9 0 g

The Author

The author was born in The Netherlands. She had a keen interest in plants

before she could read and at nine she knew she wanted to be a botanist. Having

completed secondary school, she studied Biology at the University of Leiden

where she achieved her Masters in 1986, with a major subject on Malesian and

Australian species of the grass genus Oryza supervised by Dr. J.F. Veldkamp

and in 1996 her PhD-thesis on the generic delimitation of Arctium (Asteraceae)

and a worldwide revision of its species. During these years she assisted and led

several botanical excursions for students, experiencing the difficulties faced by

relatively untrained people when using identification keys. These experiences

were intensified when in 1993 she started working for FLORON, a network

of mainly amateur naturalists investigating the species distribution of the flora of The Netherlands. Since

2000, she lives with her husband and three kids in Singapore. When the youngest of her kids went to

school in 2002 she was very happy to return (as a volunteer) to the taxonomy of grasses. Her aim in

writing this book is to stimulate more people to study the grasses.

About the Book

Grasses are very important to mankind, especially for food. They also form an important part of today’s

vegetation of Singapore, occupying at least 18% of its land surface. Nevertheless they are generally

neglected, as they are considered difficult to identify. The purpose of this book is to provide all the

necessary information with which anyone interested in grasses should be able to identify the 134 non-

woody species and varieties that occur in Singapore. The book includes 16 new records, which were

not mentioned for the country in any of the earlier accounts of the family. The characters used in the

identification keys are easy-to-see and are explained in a separate section on Grass Morphology and in

a Glossary. The Key to Sterile Grass Plants is a novelty for the region. It enables one to identify grasses

that are not yet flowering, which is especially useful when doing a botanical survey of an area within

a limited period of time. As turfs cover large areas, a shortened key to the grasses that are used for turf

is also included. The text is fully illustrated and the species descriptions are arranged alphabetically.

Separate indices to common (English and Malay) names and scientific names are included. The book

should prove useful to the laymen, the student as well as the professional, with information on origin,

abundance and ecology included. The book primarily focuses on Singapore. However, it is estimated

that it also covers about 90% of the grass species occurring in central and southern Peninsular Malaysia

(below 500 m altitude and not including limestone areas).

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