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LIBRARY

OF THE

ARNOLD ARBORETUM

HARVARD UNIVERSITY

Digitized by the Internet Archive

in 2014

http://archive.org/details/gardensbulletins451 unse

nM MAY 0 4 200) ARNOLD

THE GARDENS’ BULLETIN

SINGAPORE

VOLUME 51

Singapore Botanic Gardens

1999

Singapore, 1999

Applications for reproduction should be

made to the Singapore Botanic Gardens,

1 Cluny Road

Singapore 259569

DATES OF PUBLICATION OF THE TWO PARTS WERE:

Part 1 Pp. 1-126 31 July 1999

Part2 Pp. 127-321 31 December 1999

Singapore Gardens Bulletin 51 (1999).

CONTENTS

Boyce, P.C.

The genus Rhaphidophora Hassk. (Araceae-Monsteroideae-Monstereae) in

Pemiigmiar WIAIAYSld Ai SING ADOLE 2...25..06.2 lac. geseiiesocecsiosdeentoctansctecsesnsuasevaeess 183 — 256

Hay, A.

The genus Alocasia (Araceae-Colocasieae) in the Philippines ............... ccc 1— 41

Holttum, R.E.

Tropical botanic gardens; past, present and TULUTE 6..........0ics. cece cicendeesesones 127 — 139

Johnson, D.V. and E. P. Tay

C.X. Furtado (1897—1980): contributions to the study of palms .............. 141 — 150

Kiew, R.

eae aad SHCCIES 11) WIBIESIA 12.8 i osescctienrvesuahtanocstvesassencasoredenwvarcenessoe 85 — 98

Kiew, R.

The Singapore Botanic Gardens Herbarium - 125 years of history. .......... 151 — 161

Kiew, R. Thismia goodii (Burmanniaceae), the blue-capped thismia, a new

Re nN OREN Poe ata ccc eB udder ss sneha Sencs tone ca dveew aavueed sdtvasantecduseresksadsancs 179 — 182

Kim-Lang Huynh

On some species of Pandanus and Freycinetia (Pandanaceae) in

Ra NE YON DO ci EM Occ re yc acssl Ua Lnsstitue oa rolvnine Vida cies nsedesvivicbnvdawstRtnohae 163 — 174

Mabberley, D.J. The importance to Indopacific botany of Baron Dumont

Ge! CGESE DS OMAIISEC CUI IICLEP bie soe sie ce ai essa outs soenssadiveycvsnnnirunncnsvcyeideercdiots 309 — 317

Nagamitsu, T., R.D. Harrison and T. Inoue

Beetle pollination of Vatica parvifolia (Dipterocarpaceae) in Sarawak,

by CEE I ee nee (UR rele sent ee, De a tel cane ee ee 43 — 54

Reynolds, D.R.

PIOUS A SOOMIVCCTSS Ge VICtINANIN ccc. cetsieetata vevncdscrnnnlgsbiausnsenanceddoaeerascceresenenss 71 — 84

Salma, I.

The taxonomic significance of trichome morphology in the genus

PR FM NaI SAN RI rd Mkt a sattcea cinch sends vary sa cas astavawavasboieada\ Supnaseacntacvonsebaneas> 55 — 70

S.M. Tam

Floristic diversity of Bukit Bauk (Terengganu), Peninsular Malaysia ...... 257 — 308

Teo, L.L. and R. Kiew

First record of a natural begonia hybrid in Malaysia ............cceceseeeeeeeeeee: 103 — 118

Turner, I.M.

Euphorbia heterophylla and E. cyathophora (Euphorbiaceae) in the

EAN OM SAIRAMIN Ps 0 KA cp oO. ste rele anon ad ipcidoans udev gpvedbevoeniaptasee 99 — 102

Singapore Gardens Bulletin 51 (1999) iv.

Turner, I.M. and P.F. Stevens

The transfer of Tripetalum cymosum K. Schum. (Guttiferae) to Garcinia........... 175 — 177

Veldkamp, J.F.

Eupatorium catarium, a new name for Eupatorium clematideum

Griseb., non Sch.Bip. (Compositae), a South American species

naturalized and spreading in SE Asia and Queensland, Australia

sxcnywanshctnenes snbivii ngpesesltaadellds bibs tanks dope en cicineapndelen ed Rte tains tan 119 — 124

BOOK REVIEWS

Wu Peng-cheng (ed.). Bryological Biology, Introduction and Diverse

BSEADCHES. oessvscscserivesecenescescecneosnsenecenessennsu onde 0s Ege Atee neeleaean ee ee 319 — 320

Rossignol, M. ef al. Struggle of Life .........2.::..:scsees.cpseaeceoeassevsn cqagene ieee en ee ae 321

OBITUARY

Kizhakkedathu Mathai Kochurmmen ........0.............0....cccccccceecccccsecccccsccccecccccssucceeenece 12=—. 176

The Gardens’ Bullega |

Singapore _ ae

VOL. 51 (Part 1) July 1999 | ane ISSN 0374-7859

NATIONAL PARKS BOARD © a ae

Singapore Botanic Gardens Cluny Road Singapore 259569 bia 4741165 Telefax: 4754295 Je

THE GARDENS’ BULLETIN

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 Ruth Kiew | Dr R.T. Corlett

(Editor) University of Hong Kong

Singapore Botanic Gardens

Dr T.W. Foong

(Assist. Editor)

Singapore Botanic Gardens

Dr S.C. Chin

Singapore Botanic Gardens

Dr M.J.E. Coode

Royal Botanic Gardens

Kew, U.K.

Hong Kong

Dr M.C. Roos

Riyksherbarium

Leiden, Netherlands

Dr E. Soepadmo

Forest Research Institute Malaysia

Kepong, Malaysia

Dr W.K. Tan

Singapore Botanic Gardens

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. —

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

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ry rN

KF. 5 NOV i° 199

The Gardens' Bulletin

Singapore

VOL. 51 (Part 1) July 1999 ISSN 0374-7859

CONTENTS

Hay, A.

The genus Alocasia (Araceae-Colocasieae) in the Philippines .............cceceeesseeseeseeeeeeseees 1

Nagamitsu, T., R.D. Harrison and T. Inoue

Beetle pollination of Vatica parvifolia (Dipterocarpaceae) in Sarawak, Malaysia......... 43

Salma, I.

The taxonomic significance of trichome morphology in the genus

eras OE AMID NCU US Fea cles etch eet sen eet cea Tes aCe Saclvasccdooesaoheasiabvas eddedeue otons dacdecbupeassaien 55

Reynolds, D.R.

PO cal is PrsC IM VU CLES, Go. Wie Miia se Nate Roach ca enpctidctocasiciteaodscncthasdd scudenncaesehiameshaiseseoss 71

Kiew, R.

BRC Aah at Sal OL OICE SPECIES WA IMIANESIA s,s o5- 53 sleactedschnabsonsicmesectepstsddeneannteassavaccnwteacsepengannsce 85

Turner, I. M.

Euphorbia heterophylla and E. cyathophora (Euphorbiaceae) in the

Pe Mm Oc SMORE SUNIL Pury Oe ne es Veeder eee ed ee, cE acl cies ead nepapsnabeacoeetecadebmddaedatidenacheceds Se)

Teo, L.L. and R. Kiew

Pips (ceord of anatural beponia hybrid i Malaysia 6!) 0. ,.05.cscucclssaeedesesetscontecsencnuceeders 103

Veldkamp, J.F.

Eupatorium catarium, a new name for Eupatorium clematideum

Griseb., non Sch.Bip. (Compositae), a South American species

naturalized and spreading in SE Asia and Queensland, Australia ..........ceceeeeeeeeeeeees 119

Obituary

Be at Re eat Fad UNM aint PG TEMA ba ccc baesasacadseotecsesdacststesessaeseseesnncriasonenedeeoensete 125

Date of Publication: 31 July 1999

Published by

National Parks Board

Singapore Botanic Gardens

Cluny Road

Singapore 259569

Printed by Oxford Graphic Printers Pte Ltd

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Gardens’ Bulletin Singapore 51 (1999) 1-41.

The Genus Alocasia (Araceae-Colocasieae) in the

Philippines

AL HAY

Royal Botanic Gardens Sydney, Mrs Macquaries Road, Sydney 2000, Australia.

Abstract

The genus Alocasia (Schott) G. Don (Araceae) is revised for the Philippine Islands. Fourteen

species are recognised, of which four are new to science. A key to the species is provided.

All except Alocasia macrorrhizos (L.) G. Don are endemic. Alocasia wenzelii Merr. is

placed in the synonymy of A. zebrina Schott ex van Houtte. Alocasia manilensis Engl. and

A. warburgii Engl. are synonyms of A. heterophylla (Presl) Merr. Alocasia reversa N.E.

Brown is Bornean, not Philippine as originally attributed. The new species (A. boyceana A.

Hay, A. clypeolata A. Hay, A. scalprum A. Hay and A. ramosii A Hay), the frequently

misinterpreted Alocasia heterophylla and the very rare A. atropurpurea Engl. are illustrated.

Brief notes are made on horticultural value, conservation status, local endemicity and

relationships of Philippines Alocasia. Where possible, cultivars recognised by the

international horticultural community are ascribed to species.

Contents

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Exetuded, dubious or doubthully Philippine species ..............:.....+-...0e00+9++: 36

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Introduction

The genus Alocasia includes about 65 species of small herbaceous to massive

arborescent pachycaul aroids occurring from Sri Lanka and India, through

Indochina to China and southern Japan, the Malesian Archipelago,

Australia and Oceania. Malesian, Australian and Oceanian species have

2 Gard. Bull. Singapore 51 (1999)

been listed in Hay et al. (1995). The Sri Lankan species were revised by

Nicolson (1987). The 14 species in New Guinea and Australia were revised

by Hay & Wise (1991). The genus has its principal centre of diversity in

Borneo where there are an estimated 23 species including two intractable

complexes of extreme variability (Hay, 1998). The Philippines is the second

most species-rich subregion in Malesia for this genus, followed by New

Guinea. However, New Guinea has three endemic infrageneric groups

(Hay & Wise, 1991; Hay, 1994) and represents a somewhat distinct

transwallacean enclave, while the Philippine species appear to be of West

Malesian and continental Asian affinity. More detailed biogeographical

evaluation must await molecular and cladistic analysis of the genus.

As with the genus in other parts of its range, the Philippines species

have come to be known scientifically not only through the work of Schott

(e.g. 1860), Engler (e.g. 1879) and Engler & Krause (1920), the main

botanical specialists in the family, but also through European horticultural

channels during the late 19th Century mania for exotic foliage plants and

to a lesser extent through general floristic work. A number of species,

including the spectacular A. sanderiana W. Bull and A. zebrina Schott ex

van Houtte - both once endangered by over-collecting, were described

from plants in British and Continental nurserymen’s collections, as was A.

sinuata N.E. Brown. More recently, further new species, such as A.

clypeolata and A. scalprum described here, have first come to light through

horticultural rather than botanical initiatives. The prodigious Philippine

floristic work of Merrill (1912, 1922) and the later and more eclectic Elmer

(1938, 1939) brought additional species to light, whereas the earlier

Philippine floristic accounts of Blanco (1837 and later editions), had

obfuscated matters by the use of superfluous or misapplied names.

Further discussion of general aspects of the genus can be found in

Hay (1998) and Hay & Wise (1991).

Conservation Status and Local Endemicity

Six Philippine Alocasia species are known from very few botanical

collections and appear to have extremely restricted distributions. Alocasia

atropurpurea Engl. is known from just three collections and two localities

from northern Luzon, and does not appear to have entered horticultural

collections extensively, if at all; A. portei Schott appears to be known only

from one wild collection from Laguna Province on Luzon, though this may

well be due to undercollecting of this massive plant; A. scalprum A. Hay,

from Samar, is not represented in preserved botanical collections at all,

except for its type, cultivated at the Munich Botanic Garden, though it is

Alocasia in the Philippines 3

well-known to horticultural collectors; A. sanderiana, restricted to northern

Mindanao, has at least in the past been over-collected for horticultural

purposes, and is known from only four preserved collections from the wild;

A. sinuata is known from a single botanical collection from Mindanao,

though material of horticultural origin is attributed also to Leyte and

Palawan; A. clypeolata A. Hay, also from northern Mindanao, is known

botanically only from its type, though this species again is well-known

horticulturally. The actual status of these species, with regard to

endangerment in the wild, requires verification by local botanists with field

access. Most of these species, however, except A. atropurpurea, are known

to be in horticulture, though more systematic measures for protection may

well be desirable.

Horticulture

Several Philippine Alocasia species are of considerable ornamental interest,

and are figured in the horticultural account by Burnett (1984). Some of

those cultivated entities he recognised appear to have no counterpart among

preserved botanical material and remain undescribed. Burnett illustrated

many species, though not always under the correct names, and recognised

a number of cultivars. Where correctly named useful illustrations of

Philippine species appear in that account, I have cited them.

Alocasia portei, of enormous stature and magnificent foliage, is widely

cultivated in the tropics and in tropical conservatories of public gardens

outside the tropics. A. sanderiana has long been in demand for its striking

leaves, and has been extensively used in hybridisation programmes. A.

zebrina is also a fine plant for use in tropical landscaping. The variable A.

micholitziana Sander has had a number of cultivars selected. Alocasia

scalprum is, according to Burnett (1984), extensively cultivated in the

Philippines as an ornamental pot plant. A. clypeolata has also been taken

up by specialist horticultural collectors, as have forms of A. sinuata, A.

heterophylla (Presl) Merr., A. boyceana A. Hay and A. ramosii A. Hay.

Some of these have now been tissue cultured (mainly in the U.S.A.) and

are commercially available.

ALOCASIA

Alocasia (Schott) G. Don in Sweet, Hort. Brit. ed. 3 (1839) 631; Schott,

Oester. Bot. Wochenbl. 2 (1852) 59; Engl. in A. & C. DC., Monogr.

Phanerogam. 2 (1879) 497, in Beccari, Malesia 1 (1882) 292, in Engl. &

Prantl, Pflanzenfam. 2 (3) (1887) 137, Bot. Jahrb. Syst. 25 (1898) 23; Naves,

4 Gard. Bull. Singapore 51 (1999)

Novis. App. (1882) 193; Merr., Fl. Manila (1912) 132, Enum. Philipp. FI.

Pl. 1 (1922) 184; K. Krause & Engl. in Engl., Pflanzenr. 71 (1V.23E) (1920)

71; Hay & Wise, Blumea 35 (1991) 503; Hay, Gards. Bull. Singapore 50

(1998) 236. - Colocasia Schott sect. Alocasia Schott in Schott & Endl.,

Melet. Bot. 1 (1832) 18. - Type: Alocasia cucullata (Lour.) G. Don.

Schizocasia Schott [Bonplandia 10 (1862) 148 (invalidly published); Engl.

in A. & C. DC., Monogr. Phanerogam. 2 (1879) 495 (invalidly published) |

ex Engl., Bot. Jahrb. Syst. 1 (1880) 186 & 25 (1898) 26, in Becc., Malesia 1

(1882) 295, in Engl. & Prantl, Nat. Pflanzenfam. 2, 3 (1887) 137; K. Krause

& Engl., Pflanzenr. 71 (IV.23E) (1920) 115; Bunting, Baileya 10 (1962)

112. - Type: Schizocasia acuta Engl. = Alocasia brancifolia (Schott) A.

Hay.

Massive arborescent pachycauls to small decumbent herbs; stems sympodial,

more or less cataphylliferous, fleshy, often starch-filled, smooth to tessellated

to clothed in marcescent leaf bases, usually bearing short, sometimes

branched, mostly subterranean stolons terminating in small cormels; foliage

leaves solitary or more usually clustered, glabrous to thickly pubescent,

sometimes enormous; petioles sheathing in the lower part, sometimes

glandular; leaf blades deeply pinnatifid to hastosagittate, to suborbicular,

to narrowly lanceolate, sometimes peltate (almost always in seedlings),

often strikingly coloured or of metallic appearance or bullate, membranous

to thickly coriaceous; major venation pinnate, running to a sometimes

conspicuous submarginal vein, usually with glands in the axils abaxially;

secondary venation reticulate, often joining between the primary veins to

form interprimary collective veins, but these quite commonly absent and

then secondary venation tending towards striate; inflorescences (solitary

to) paired and subtended by a prophyll and cataphyll, the pairs either

interspersed with foliage leaves (and then appearing lateral) or produced

in succession and then forming a cluster in the centre of the leaf crown;

peduncles mostly short at anthesis, rarely subequalling the petioles; spathe

divided into a persistent lower portion enclosing the ovaries, sterile interstice

and sometimes part of the male zone, the upper portion a deciduous,

variously coloured, membranous limb (this persistent in some New Guinea

species); spadix monoecious, protogynous, sometimes stipitate, shorter than

to equalling, rarely exceeding the spathe, sometimes partly adnate with the

spathe in the lower portion; basal female zone of naked pistils; ovaries

unilocular to apically incompletely 4-septate; stigma sessile or on a usually

short style, button-like to stellate; placentation basal; ovules several,

Alocasia in the Philippines 2

anatropous to suborthotropous; distal to the female zone a sometimes

attenuate, sometimes very short sterile interstice composed of (sometimes

massive) synandrodia; male zone composed of close-packed

rhombohexagonal synandria; thecae opening by apical pores, or concealed

from above by the expanded over-topping synconnective; spadix terminating

in a conspicuous appendix usually composed of elongate sinuous low

structures considered homologous with staminodes; fruiting peduncle usually

elongating; fruits usually red berries contained within the persistent and

enlarged spathe base which eventually dehisces to expose them; seeds ca.

3—5 mm in diam., albuminous, with thin testa.

Distribution: About 65 species from IndoMalesia with extratropical

extensions into the Himalayas, southern China, southern Japan and eastern

Australia. Alocasia macrorrhizos and its several varieties are virtually

pantropical, naturalised through introduction as ornamental plants and as

an inferior starch crop. However, even through much of its Asian range it

occurs in close association with human habitation and it may be a cultigen.

In Malesia there are 57 species currently recognised, 14 of them in the

Philippines.

Relationships of the Philippine species

In the present herbarium-based alpha-taxonomic study, most of the

Philippine species of Alocasia do not fall readily into groups either with

each other or with species outside the Philippines. No attempt has been

made to treat them under informal group headings, as has been done in

previous papers (Hay, 1998; Hay & Wise, 1991). Some affinities are to a

certain extent recognisable, however, though nevertheless tenuously at

present: Alocasia sanderiana and probably A. boyceana belong in the mainly

West Malesian Longiloba Group (see Hay, 1998); A. maquilingensis falls

rather clearly into the Puber Group, also West Malesian. Alocasia

atropurpurea, and other species with membranous peltate leaves — A.

micholitziana and A. clypeolata, may be allied with continental Asian A.

odora (Lodd.) Spach. The thick-textured sometimes peltate leaves of A.

sinuata suggest it may be related to the Bornean endemic Scabriuscula

Group. Alocasia portei is clearly related to A. macrorrhizos. The remaining

species do not suggest any particular affinity. Molecular studies are needed

throughout the genus to assist in proposing a firm infrageneric classification.

la.

1b.

Da.

2b.

ere

4a.

Ab.

5a:

5b.

6a.

6b.

Ta,

Tb.

8a.

Sb.

9a.

Gard. Bull. Singapore 51 (1999)

Key to the Species in the Philippines

Leaves all distinctly peltate, with the confluence of leaf blade across

the sinus exceeding S amma width ia)442)2ee hee eee 2

Leaves not peltate or only some on the plant peltate or confluence of

leaf blade across sinus not exceeding 4 mm in width ..... eee eee 5

Leaf margin deeply to shallowly sintiate 22.:.....2.00 0. 22ers 3

Leaf margin ROtSINUGte tae ee ee es 4

Leaf blades glossy, petioles subtended by _ cataphylls

PS ARPS RUDD Galiter Lan tle Maia! Abide bw h( iM ON orld a AE eal IE! 2 11. A. sanderiana

Leaf blades velvety, petioles not subtended by cataphylls

Rc AWE Oa EO Ee See eee 8. A. micholitziana

Blade cordato-sagittate, peltate, with the posterior lobes united only

aL ERC DASE cates. Lag ai os tint tp ete cee ee i 1. A. atropurpurea

Blade ovato-sagittate, peltate, with the posterior lobes united for most

Of theit envi. 2.5 24 2 eens aeeae ee 3. A. clypeolata

Blade deeply pinnatifid; massive"plants’ i... eee 9. A. portei

Blade with entire to shallowly undulate margin, massive to diminutive

PARAS sig ssres bcs cds tisttdy ait ai dhannsedaeenibs dm Nateeaans cage egiadan oa ahem ae ean 6

Blade very broadly cordato-sagittate, ca. 80 cm or more long ........... (i

Blade (sometimes very narrowly) sagittate to hastate, if broadly

cordato-sagittate, then small, less than 30 cm long... ee 8

Leaves glabrous; secondary venation not very prominent abaxially,

not forming interprimary collective veins ............... 6. A. macrorrhizos

Leaves densely to sparsely pubescent abaxially; secondary venation

prominent abaxially and forming well-defined interprimary collective

VMS PLL cone us setdeauiere dene raeig aca Supt n ce et to heed eRe 7. A. maquilingensis

All leaves with posterior costae with lamina to the sinus.................0.. 9

Non-peltate leaves with posterior costae (very reduced in A. scalprum)

faked my The SMEs ae pes cceesel ys acca ue devaoe aera ene een 10

Robust plants 1-1.8 m tall; leaf blade not bullate; secondary veins not

forming subsidiary Veins). jis iis scisigho toned ee 14. A. zebrina

Alocasia in the Philippines i

9b. Small plants 40 cm tall or less; leaf blade more or less bullate; secondary

Vem AMON TOPMII, SUDSICIALY VEUIS) yves.ccescccccsecicese bee seeetane 13. A. sinuata

10a. Posterior lobes about a third or less the length of the anterior lobe;

HV ETT SAC CONAN ois ce eceis oon sdeesvaecienesenennneeds 12. A. scalprum

10b. Posterior lobes more than a third the length of the anterior lobe;

Meets UMD EAN V LY AEIICOON AWE. .0..052. ioastdedsaseksncuyaresvoveempeue tab cadeatecesateces 11

11a. Sterile interstice composed of very large synandrodia occupying upper

part of lower spathe cavity; spathe constriction shallow .......0...... 12

11b: Sterile interstice not so; spathe constriction abrupt ...2.........sc.escsecee 13

12a. Posterior lobes diverging at an obtuse to very obtuse angle

ee eR elo Ncvyans dices bucie ues dtaekosetntso dens lier wnben case 4. A. culionensis

12b. Posterior lobes diverging at an acute angle ............. 5. A. heterophylla

13a. Posterior lobes ca. two thirds the length of the anterior; stigmas button-

bay QU ee Rein (ihe SESS R ne Pb BRL TO 10. A. ramosii

13b. Posterior lobes ca. half the length of the anterior; stigmas acutely

ee iicerss bi ies esp ane ics oH Sia gap owdwccuceacteweasteclae 2. A. boyceana

The Philippine Species

1. Alocasia atropurpurea Engl.

Alocasia atropurpurea Engl., Pflanzenr. 71 (IV.23E) (1920) 132; Merr.,

Enum. Philipp. Fl. Pl. 1 (1922) 184. - Type: Luzon, Ifugao Subprov., Mt

Polis, Feb 1913, McGregor BS 19663 (B, holo; K, US, iso).

Moderately robust herb; petiole to ca. 50 cm long, sheathing in the lower

third; blade membranous, ovato-sagittate, ca. 40 cm long, very shallowly

peltate; anterior lobe ca. 30 cm long x 30 cm wide at base, with about 4

primary lateral veins on each side of the anterior costa diverging at ca. 45-

70°; secondary veins flush on both sides of the blade, arising from the

primary veins at ca. 70° and then deflected to the margin, not or hardly

forming interprimary collective veins; posterior costae diverging at an obtuse

angle; posterior lobes ca. half the length of the anterior, rounded, with

little laminar tissue on the posterior side of the posterior costae, but this

confluent across the sinus; glands in the axils of the primary veins

Gard. Bull. Singapore 51 (1999)

Sh Cn,

CRC Ss .

Dare Feit ta

ae "4

EAL Wm LONE NEY

OL RS

ae ai

Lhe Sage hae

theeeegi(

& Cy

wc:

se!

O.0%6

HSAs ~

“«.

Figure 1. Alocasia atropurpurea Engl.

MacGregor BS19663 - A: leaf blade; B: inflorescence (part of spathe removed); C: pistils; D:

synandria. - Scale: bar to A,B = 5 cm, to C,D = 4 mm.

Alocasia in the Philippines 9

inconspicuous or absent; inflorescence arrangement [not known]; spathe

to ca. 14 cm long; lower spathe ca. 4 cm long, ovoid, asymmetric (due to

partial connation with spadix); limb broadly oblong-lanceolate, cucullate,

the basal ca. 1 cm deflected and somewhat inflated, then the rest erect,

dark purple brown; spadix somewhat shorter than the spathe; female zone

ca. 3 cm long, partly connate with spathe in lower third, tapering distally;

ovaries globose, rather large, ca. 2.5-3 mm diam. (dry); stigma strongly 3-

4-lobed on a very short style; sterile interstice slender, ca. 1 cm long,

corresponding with spathe constriction; male zone ca. 3 cm long x 1.3 cm

thick (dry); synandria rhombo-hexagonal, ca. 2 mm diam., more or less

capped by expanded synconnective; appendix slightly longer than to about

twice the length of the male zone, slightly constricted at junction with male

zone, thence cylindric and distally abruptly tapering; infructescence

unknown.

Distribution: Endemic to northern Luzon, known only from Mt Polis, and

an unspecified locality in Bontoc Subprovince.

Habitat: Not recorded.

Notes: This apparently highly distinctive species is poorly known. The

inflorescence seems large for the size of the leaves, though the latter may

have been collected from conveniently small examples. The female zone

partly adnate to the spathe, the large ovaries and purple, rather inflated

spathe limb appear diagnostic. Vanoverbergh 3684 has the appendix

relatively much longer than other specimens. Alocasia atropurpurea bears

some resemblance to continental Asian A. odora (Lodd.) Spach.

Other specimens seen: Luzon, Mountain Province, Mt Polis, Celestino PNH

8014 (GH); Luzon, Bontoc subprov., Vanoverbergh 3684 (P).

2. Alocasia boyceana A. Hay, sp. nov.

Ab Alocasia ramosii spadice stipitato, stigmatis acute lobatis, lobis posticis

foli lobum anticum circa dimidium aequantibus differt. - TYPUS: Cult.

Botanischer Garten Miinchen ex Philippines, Cebu (orig. coll.

Seidenschwarz s.n., voucher M), J. Bogner s.n. (NSW, holo).

[Alocasia heterophylla auct. non (Presl) Merr.: Engl. & K. Krause, Pflanzenr.

71 (IV.23E) (1920) 101, p.p. quoad specim. cit. Merrill 5328].

Moderately robust herb to ca. 60 cm tall; stem ca. 2 cm diam.; leaves few,

10 Gard. Bull. Singapore 51 (1999)

?1-3 together, often subtended by cataphylls; cataphylls papery, lanceolate,

to ca. 11 cm long; petiole to ca. 45 cm long, sheathing in the lower third,

sometimes maculate; blade rather narrowly sagittate to narrowly hasto-

Sagittate, to ca. 35 cm long; anterior lobe to 24 cm long, ca. 11 cm wide at

base, with 4—5 primary lateral veins on each side of the anterior costa

diverging at 45—80°; margin slightly undulate to entire; primary lateral veins

somewhat prominent adaxially and abaxially, with inconspicuous glands in

the axils on abaxial side; secondary venation arising from the primary at

ca. 70-90° then deflected marginally, not or hardly forming interprimary

collective veins; posterior lobes slender, about half the length of the anterior:

posterior costae diverging at an obtuse (hastate) to acute (sagittate) angle,

naked in the sinus for 1-2 cm; inflorescences paired or solitary; peduncle

to ca. 20 cm long; spathe ca. 7.5 cm long; lower spathe ca. 2 cm long, ovoid,

separated from the limb by a strong constriction; limb lanceolate, reflexed:.

spadix somewhat shorter than the spathe, stipitate for ca. 4 mm with the

stipe partly adnate to merely obliquely inserted on the spathe; female zone

6-8 mm long; pistils subglobose, ca. 1.5 mm diam.; stigma subsessile, rather

sharply 34-lobed; sterile interstice about equalling the female zone, tapering

distally, its top corresponding with spathe constriction; male zone ca. 12

mm long, cylindric, ca. 5 mm diam.; synandria rhombo-hexagonal, ca. 1

mm diam., opening by apical pores not overtopped by the synconnective;

appendix 1.5-1.7 cm long, tapering, composed of irregularly sinuate

staminodes, slightly narrower than to equalling the male zone in thickness;

infructescence broadly ovoid, ca. 2.4 cm diam.

Distribution: Philippines, on Luzon, Cebu, Negros.

Habitat: Rain forest at low to medium elevation, sometimes on limestone.

Notes: The stipitate spadix, sharply lobed stigmas and cataphylliferous stem

with few leaves suggest that A. boyceana is a segregate entity of the mainly

West Malesian A. longiloba Miq. complex.

This species is named for Peter Boyce (K), in recognition of his

continuing contribution to knowledge of tropical Asian Araceae.

Other specimens seen: Negros, Negros Oriental Province, Dumaguete

(Cuernos mts), Elmer 9702 (BO, E, G, K, L, NSW, NY, US); Luzon,

Hacienda de Julajala, Loher 2437 (K); Jolo, Mt Dajo, Merrill 5328 (US):

Mindanao, Colabato prov., Nutol, Ramos & Edano BS 84942 (H); Cebu,

Seidenschwarz s.n. (M); Cebu, Tagbao, University of San Carlos 145 (L);

Luzon, Bataan Province, Lamao R., Whitford 1362 (K, US); Central Luzon,

Bataan Province, Mt Mariveles, Lamao River, Williams 794 (NY).

Alocasia in the Philippines 1]

Figure 2. Alocasia boyceana A. Hay

Bogner s.n. - A: leaf and venation; B: inflorescence with part of spathe removed; C: pistils; D:

neuter organs of sterile interstice; E: synandria. - Scale: bar to A = 8 cm, to B = 2.4 cm, to C, D,

E:=.6.mm.

12 Gard. Bull. Singapore 51 (1999)

3. Alocasia clypeolata A. Hay, sp. nov.

Ab aliis speciebus Alocasiis Philippinarum lamina folii parva ovata vel

subrotunda valde peltata differt.- TYPUS: Philippines, Mindanao, Surigao

Province, Mt Kabatuan, 18 Mar 1949, Mendoza & Convocar PNH 10433

(GH holo, 2 sheets).

[? Alocasia cv. Green Shield; see Burnett, Aroideana 7 (1984) 87, fig. 20].

Small herb to ca. 30 cm tall; stem short, ca. 5 cm long x 2 cm diam. (dry);

leaves ca. 6 together; petiole ca. 17 cm long, sheathing in the lower quarter;

blade weakly coriaceous, darker near the main veins, ovate to very broadly

ovate, 16 x 6.5 cm to 12 x 8 cm; anterior lobe 7-11 cm long, widest ca.

quarter of the way from the base, the tip acute to obtuse and then apiculate;

anterior costa with 3 primary lateral veins on each side diverging at 45-60°;

secondary venation forming irregular interprimary collective veins in the

proximal part of the blade, these not formed in the distal portions; all

venation flush with the lamina on both sides (dry); posterior lobes ca. 4 cm

long, peltate for more than 80% of their length, with the posterior costae

subparallel; inflorescences paired amongst the leaves, subtended by oblong

lanceolate cataphylls to ca. 5 cm long; peduncle much shorter than the

petioles, ca. 5 cm long; spathe white, ca. 6 cm long; lower spathe ca. 1.8 cm

long, ovoid; spathe limb broadly lanceolate ca. 4.2 cm long x 1.8 cm wide at

the base, at male anthesis with the lower ca. 1 cm sharply reflexed and the

rest erect; spadix more or less equalling the spathe, stipitate for ca. 2 mm;

female zone subcylindric, ca. 1 cm long x ca. 6 mm diam.; ovaries globose,

ca. 1.2 mm diam.; style shorter than ovary, ca. 0.5 mm long; stigma rounded

and weakly lobed; sterile interstice ca. 6 mm long x 3 mm diam., composed

of rhomboid synandrodia ca. 1.3 mm diam.; male zone 1 cm long x 4 mm

diam., subcylindric, narrowed at the base and apex, the base corresponding

with the spathe constriction; synandria rhombo-hexagonal, ca. 1 mm diam.,

of 3-4 connate stamens; thecae opening by apical pores not overtopped by

the synconnective; appendix ca. 3 cm long, tapering to a point, covered in

low, elongate sinuous irregular sterile organs; infructescence with short

peduncle; fruiting spathe subglobose, ca. 2.5 x 1.5 cm.

Distribution: Known botanically from only the type collection made on

Mindanao.

Habitat: In rocky soil on a steep slope on forest edge, 80 m altitude.

Notes: This species bears strong resemblance to the plant known in

cultivation as Alocasia ‘Green Shield’, illustrated in Burnett, Joc. cit., and

Alocasia in the Philippines 13

Figure 3. Alocasia clypeolata A. Hay

Mendoza & Convocar 10433 - A: habit; B: venation; C: spadix with part of spathe removed; D:

synandria. - Scale: bar to A = 4 cm, to B = 2.5 cm, to C= 1 cm, to D = 2 mm.

14 Gard. Bull. Singapore 51 (1999)

also commonly grown as ‘Green Cuprea’. A. clypeolata appears to differ

from this cultivar only in its somewhat smaller leaf dimensions (about half

the size of the upper limit reported for the cultivated plant by Burnett).

The cultivated plant is said to have the leaf blades lime green with

‘dramatically contrasting black [sic] zones around the primary veins’

(Burnett, 1984: 87). Further colour notes made from the cultivated plant

include that the lower part of the spadix is cream to white and the appendix

is yellow (Burnett, loc.cit.)

The specific epithet alludes to the leaves shaped like small shields.

4. Alocasia culionensis Engl.

Alocasia culionensis Engl., Bot. Jahrb. Syst. 37 (1905) 135; K. Krause &

Engl., Pflanzenr. 71 (1V.23E) (1920) 102; Merr., Enum. Philipp. Fl. Pl. 1

(1922) 184. - Type: Philippines, Culion Island, Ilocano, Dec 1902, E.D.

Merrill 555 (B, holo; K (mixtum), US, iso).

Moderately robust herb to ca. 70 cm tall; stem erect, ca. 2 cm thick; leaves

several together, ?not interspersed with cataphylls; petiole to ca. 45 cm

long, sheathing in the lower third; blade sagittate to hastate, ca. 35-45 cm

long; anterior lobe narrowly triangular to triangular, sometimes with a

faintly undulate margin, 10-18 cm wide at base, with 3—5 primary lateral

veins on each side of the anterior costa diverging at 60-80°; axillary glands

inconspicuous; secondary venation fine, rather distant (ca. 3-4 mm apart),

arising from the primary at a wide angle, then rather abruptly deflected to

the margin and forming rather disorganised interprimary collective veins

only towards the margin, flush with the lamina on both sides; posterior

lobes diverging at an obtuse to very obtuse angle, spreading, tapering to

sub-rhomboid to narrowly round-ended, 11-20 cm long; posterior costae

naked in the sinus for 2—5 cm; inflorescences clustered, ca. 6-8 together,

subtended by lanceolate cataphylls ca. 12 cm long; peduncle to ca. 20 cm

long; spathe ca. 8-9 cm long; lower spathe 2—3 cm long, separated from the

limb by a long gradual constriction; limb narrowly lanceolate, white,

eventually reflexed, the tip mucronate for ca. 1 cm, the mucro straight;

spadix somewhat shorter than the spathe, stipitate for 2.5 mm, the stipe

free but obliquely inserted; female zone 5 mm long; ovaries more or less

flask-shaped, upturned, ca. 1 mm diam.; style ca. 0.5 mm long; stigma

weakly 2—3-lobed; sterile interstice short, ca. 4 mm long, composed of two

whorls of massive synandrodia filling the upper part of the lower spathe

cavity; male zone ca. 1.4 cm long, cylindric, ca. 5 mm diam. (dry); synandria

rhombo-hexagonal, ca. 1.5 mm diam., thecae opening by apical pores not

capped by synconnective; appendix 1.7 cm long, tapering; fruiting spathe

ovoid, ca. 3 cm diam.

Alocasia in the Philippines 15

Distribution: Endemic to the Philippines; Luzon, Palawan, Basilan, Culion,

Balabac, Mindoro, Mindanao, Leyte, Busuanga, Panay.

Habitat: Lowland rain forest.

Vernacular name: Urinkokuk (Yakan).

Notes: The isotype sheet at K has, in addition to Merrill 555, a specimen of

A. heterophylla with associated label notes of Loher 2442.

This species is very similar to A. heterophylla in inflorescence form,

sharing the weakly constricted spathe, flask shaped up-turned pistils beneath

massive synandrodia occupying the upper part of the lower spathe chamber.

It differs in the relatively shorter appendix, relatively longer male zone,

the overall larger size of the inflorescences arranged into larger

synflorescences, the larger leaves with finer secondary venation and widely

diverging posterior lobes that are never peltate. In the herbarium, the two

are easily separated in aspect and do not exhibit intergrading. However,

the listed differences are somewhat trivial, at least individually. While the

distributions of these two species, as recognised here, overlap, A. culionensis

has a more southerly emphasis. The matter of distinction of these species

may be more satisfactorily resolved by further field work.

Other specimens seen: Basilan, Isabela de Basilan, Ebalo 938 (H); Palawan,

Ebalo & Conklin 1275 (H); Sulu Archipelago, Jolo, Siasi, Kondo & Edano

PNH 38757 (H); Busuanga Isl., Malbato, Marche B229 (P); Palawan, Lake

Manguao, Merrill 9568 (K, US); Balabac, Dalawan Bay, Olsen 535 (L);

Leyte, Dagami, Ramos BS 15353 (US); Mindoro, Paluan, Ramos BS 39602

(K, P, US); Busuanga Isl., Ramos BS 41235 (US); Mindanao, Davao

Prov., Mati, Ramos & Edano s.n. (UC); Panay, Capiz Prov., Jamindan,

Ramos & Edano BS 31264 (BO, L); Balabac Isl, Ramos & Edano BS

49709 (UC); Luzon, Sorsogon Prov., Mt Bulusan, Sulit PNH 2649 (US).

5. Alocasia heterophylla (Presl) Merr.

Alocasia heterophylla (Presl) Merr., Philip. J. Sci. 3 (1908) 220, Fl. Manila

(1912) 133, Enum. Philipp. Fl. Pl. 1 (1922) 184; K. Krause & Engl., Pflanzenr.

71 (IV. 23E) (1920) 101, fig. 22. - Colocasia? heterophylla (Presl) Kunth,

Enum. Pl. 3 (1841) 40. - Caladium heterophyllum Presl, Relig. Haenk. 2

(1835) 148. - Type: Philippines, Luzon, Haenke s.n. (PR, holo).

Alocasia manilensis Engl., Bot. Jahrb. Syst. 25 (1898) 23; K. Krause &

Engl., Pflanzenr. 71 (I1V.23E) (1920) 95, fig. 19E & F; Burnett, Aroideana

16 Gard. Bull. Singapore 51 (1999)

7 (1984) 92, fig. 26. - Type: Philippines, Luzon, Manila Prov., Mt Alban,

Mar 1888, Warburg 12493 (B, holo).

Alocasia warburgi Engl., Bot. Jahrb. Syst. 25 (1898) 25; Merr., Philipp. J.

Sci. 1, Suppl. (1906) 33. - Type: Philippines, Warburg 13652 (B, presumed

destroyed); Celebes, Warburg 15723 (B, presumed destroyed). Neotype:

Philippines, Luzon, Tayabas Prov., Mt Binuang, Ramos & Edano BS 28495

(US; isoneo, BO, P, US).

Small herb to ca. 40 cm tall; stem to ca. 20 cm long, decumbent to creeping,

ca. 1.7 cm thick; leaves 3-5 together; petiole to ca. 35 cm long, sheathing in

the lower ca. quarter to third; blade to ca. 27 cm long, narrowly (hasto-)

sagittate, sometimes shallowly to deeply peltate in adult plants and then

some non-peltate leaves also present; anterior lobe to 20 cm long, to ca. 10

cm wide at base, narrowly triangular, with the margin sometimes shallowly

sinuate; anterior costa not very prominent abaxially, with 3-4 sometimes

opposite primary lateral veins diverging at ca. 45—60° and usually distally

deflected towards the leaf tip, then joining a conspicuous submarginal vein

situated ca. 3 mm from the margin; secondary venation rather widely spaced

(ca. 3 mm) arising from the primary at a high angle, thence mostly abruptly

(ca. right angle) deflected towards the margin and forming rather

disorganised interprimary collective veins; glands in the axils of primary

veins inconspicuous; posterior lobes diverging at an acute angle, narrow,

tapering, sometimes distally out-turned; posterior costae naked in sinus for

ca. 1.2 cm to peltate for ca. 20% of their length; inflorescences paired, to 4

together; peduncle to ca. 4 cm diam., ca. half the length of the petioles;

spathe 5-6.5 cm long; lower spathe narrowly ovoid, 2.5—3 cm long,

differentiated from the limb by a rather long gradual constriction; limb

narrowly lanceolate, eventually reflexed; spadix ca. three quarters of the

length of the spathe, shortly stipitate, with the stipe adnate to the spathe;

female zone 0.5—1 cm long, containing few to several flask-shaped pistils;

style ca. 0.5 mm, stigma globose; sterile interstice ca. 5 mm long, comprised

of massive synandrodia filling the upper part of the cavity of the lower

spathe; male zone ca. 5—8 mm long, subcylindric, ca. 3 mm thick; synandria

more or less hexagonal, ca. 1.2 mm diam., opening by apical pores not

capped by synconnective; appendix ca. 2 cm long, tapering; infructescence

ovoid, ca. 3 cm diam.

Distribution: Endemic to the Philippines; Luzon, Mindanao, Polillo.

Although Engler cited a specimen (now destroyed) from Sulawesi in the

protologue of A. warburgii, no other material from Sulawesi is conspecific

with the Philippines element.

Alocasia in the Philippines 17

Lesk

Eliane

Figure 4. Alocasia heterophylla (Presl) Merr.

Ramos & Edaiio BS 28495 - A: habit; B: venation; C: inflorescence (part of spathe removed).

=scale: Dar to.A = 7.5 'cm, to B = 6'cm)ito C = 2 cm.

18 Gard. Bull. Singapore 51 (1999)

Habitat: Lowland rain forest (dipterocarp forest) to ca. 300 m altitude.

Notes: This species is distinguished from A. ramosii by the leaves sometimes

peltate in mature plants, the gradually constricted spathe, the massive

synandrodia of the sterile interstice, the more widely separated secondary

venation, distally curved primary venation and the conspicuous submarginal

vein situated well in from the margin.

The type of A. manilensis consists of seedling leaves only, but it

matches A. heterophylla in shape and in the characteristic submarginal

vein. The syntypes of A. warburgii no longer survive at Berlin presumably

because Engler and Krause had reduced this to the synonymy of A.

heterophylla and Krause consequently did not take out a representative

specimen for protection from possible war damage which did, of course,

subsequently occur. My acceptance of the placement of A. warburgii in the

synonymy of A. heterophylla is based on the joint opinion of Engler and

Merrill, who examined the pertinent material of both species together at

Berlin, concluding that the two were indeed identical (Merrill’s annotation

on Haenke s.n. at PR - the holotype of A. heterophylla). Elmer’s

interpretation, however, also evidenced by specimen annotations, was

different, applying the name A. warburgii to material of what is here

recognised as a new species, A. boyceana. Elmer’s identifications of Araceae

were not always reliable and it is not apparent that he had examined the

relevant types; I therefore defer to Engler and Merrill’s view. The selected

neotype of A. warburgii is a widely distributed fertile collection that falls

well within the concept of A. heterophylla.

The illustration of this species in Krause & Engler (Joc. cit.) is of

poor quality, and it is perhaps as a consequence of this that the name

Alocasia heterophylla has been applied by various workers to material of

distinctly different species in Java and Borneo, where this species does not

occur. Almost all the material that Krause & Engler cited under A.

heterophylla was destroyed, but one extant specimen, Merrill 5328, is not

of this species. Together with the fact that the illustration of the spadix

does not show the characteristic enlarged synandrodia, this suggests that

the illustration is based on material of more than one species.

The cultivars Alocasia ‘Blue Prince’ and ‘Blue Lady’ (Burnett, 1984:

figs 17 & 18) appear to be derived from this species.

Other specimens seen: Luzon, Zambales Prov. Mt Pinatubo, Clemens 17362

(UC); Luzon, Pangasinan Prov., Labrador, Mt San Isidro, Fenix BS 29883

(US); Luzon, Quezon Prov., Quezon National Park, ca. 10 km W of

Atimonan, Nicolson 800 (K, US); Luzon, Rizal Prov., Mt Susong Balaga,

Ramos & Edano BS 29238 (US); Luzon, Rizal Prov., Mt Irig, Ramos &

Alocasia in the Philippines 19

Edano BS 48444 (UC); Mindanao, Davao Prov., Mati, Ramos & Edafio

BS 49310 (NY, UC); Polillo Island, Robinson BS 9199 (E, NSW); Oriental

Mindoro, Mansalay, Mt Yagaw (E slope), Sulit & Conklin PNH 16954

(Git>: 1).

6. Alocasia macrorrhizos (L.) G. Don

Alocasia macrorrhizos (L.) G. Don in Sweet, Hort. Brit. ed. 3 (1839) 631

(‘-rrhizon’); Schott in Oest. Bot. Wochenbl. 4 (1854) 409 (‘-rrhiza’); Engl.

in A. & C. DC., Monogr. Phanerogam. 2 (1879) 502; Naves, Nov. App.

(1882) 293; Merr., Philipp. J. Sci. 1, Suppl. 1 (1906) 33, Interpr. Herb.

Amboin. (1917) 103, Species Blancoanae (1918) 91, Enum. Philipp. FI. PI.

1 (1923) 185; K. Krause & Engl., Pflanzenr. 71 (IV.23E) (1920) 84, fig. 15;

- Arum macrorrhizon L., Sp. Pl. (1753) 965; - Colocasia macrorrhiza (L.)

Schott in Schott & Endlicher, Melet. Bot. 1 (1832) 18. - Type: Arum

maximum macrorrhizon zeylanicum Hermann, Parad. Bat. (1698) t. 73

(lecto; selected by Furtado, 1941).

Arum indicum Lour., Fl. Cochinch. (1790) 536, & ed. Willd. (1793) 655;

Roxb., Fl. Ind. ed. 2, 3 (1832) 498. - Colocasia indica (Lour.) Kunth,

Enum. Pl. 3 (1841) 39. - Alocasia indica (Lour.) Spach, Hist. Nat. Veg.

Phan. 12 (1846) 47; Schott, Oesterr. Bot. Wochenbl. 4 (1854) 410; Engl. in

A. & C. DC., Monogr. Phanerogam. 2 (1879) 501; Naves, Nov. App. (1882)

293; Usteri, Beitr. Ken. Philipp. Veg. (1905) 130; Merr., Fl. Manila (1912)

133; K. Krause & Engl., Pflanzenr. 71 (IV.23E) (1920) 87. - Type: Arum

indicum sativum Rumphius, Herb. Amboin. 5 (1747) t. 106 (lecto; selected

by Nicolson, 1979).

Calla maxima Blanco, FI. Filip. (1837) 658. - Arum grandifolium Blanco,

Fl. Filip. ed. 2 (1845) 458 & ed. 3 (1879) 63, t. 177 [nom. superfl. pro C.

maxima Blanco]. - Neotype : Philippines, Luzon, Rizal Prov., Antipolo,

Merrill, Species Blancoanae No. 630 (US; isoneo, BM, CAL, GH, K, NSW,

NY, P designated here).

[Calla badian Blanco, FI. Filip. (1837) 658, nom. nud.]}.

Alocasia indica var. heterophylla Schott ex Engl. in A. & C. DC. Monogr.

Phanerogam. 2 (1879) 502. - Alocasia indica var. diversifolia Engl.,

Pflanzenr. 71 (IV.23E) (1920) 88, [nom. superfl. pro A. indica var.

heterophylla\. - Type: Philippines, Luzon, Manila, Hiigel s.n. (W, holo;

presumed destroyed). Neotype: Schott Icones nos. 120, 122 (W, designated

here) [fiches 69: d9, and 70: al in the microfiche edition of the Icones].

20 Gard. Bull. Singapore 51 (1999)

Massive pachycaul with the stem decumbent or erect, to 4 m tall; petioles

to 1.3 m long, sheathing in lower third to half; blades ovato-sagittate,

bluntly triangular in general outline, held more or less erect, with the

margin entire to very slightly sinuate (distinctly sinuate in ‘var. heterophylla’

- see ‘Notes’ below); anterior lobe ca. 70 cm to over 1 m long, ca. 60-90 cm

wide at base, with ca. 9 rather distant primary lateral veins on each side of

the anterior costa diverging at ca. 60°; glands in axils of primary veins on

abaxial side distinct; secondary venation flush with the lamina or but slightly

raised abaxially, not forming interprimary collective veins or these poorly

defined; posterior lobes ca. third to half the length of the anterior, somewhat

rotund, often overlapping; inflorescences paired among the leaf bases,

subtended by membranous cataphylls; peduncles barely exceeding the

cataphylls at anthesis; spathe rather variable in length, ca. 13-35 cm long,

constricted about sixth of the way from the base; lower part green, ovoid;

limb broadly oblong-lanceolate, cowl-like at anthesis, later reflexed, then

deliquescent, membranous, pale yellow; spadix slightly shorter than the

spathe; female zone 1-2 cm long, c. 1.5 cm diam.; ovaries pale green, ca. 3

mm diam.; stigma sessile, 3—5-lobed, the lobes conic, yellow; sterile interstice

slightly shorter than to equalling the female zone, whitish, slightly narrowed

corresponding to the spathe constriction; male zone cylindric, c. 3-7 cm

long, c. 2 cm diam., whitish; synandria rhombo-hexagonal, convex-topped

due to cap-forming synconnective; appendix yellowish, slightly thicker than

the male zone at the base, thence tapering, equalling to considerably

exceeding half the length of the spadix, staminodial; fruiting spathe ca. 8

cm long, green.

Distribution: IndoMalesia to Oceania. It is not clear where, if anywhere,

this species occurs wild. It has evidently been widely distributed as a

subsistence crop and is now pantropical by introduction as an ornamental.

In the Philippines, widespread.

Habitat: Roadsides and clearings often near human habitation, at low to

medium elevation.

Vernacular names: Badian(g), Biga, Bilan mumpaha, Bungiang, Bira.

Notes: In designating a neotype for Calla maxima Blanco, of which no

original material was preserved, I have followed the uncontroversial

interpretation of Merrill (1918: 91) and used the exemplifying collection he

distributed. The drawings designated as the neotype of Alocasia indica var.

heterophylla Schott ex Engl. are annotated with this name, together with

the same collector’s name and provenance as the presumably destroyed

Alocasia in the Philippines 20

holotype specimen, indicating a high probability that the drawings were

prepared from that material.

In the Philippines, the most closely related species appears to be A.

portei, which differs in the divided leaf, more elongated spadix, tessellated

trunk, longer, brownish spathe and, perhaps most importantly, the

arrangement of inflorescences, which are clustered in large numbers in the

centre of the leaf crown in A. portei, whereas they are paired among the

leaves in A. macrorrhizos. A. maquilingensis (q.v.) is also similar, differing

in the smaller inflorescences, hairy leaves and abaxially rather prominent

secondary venation forming interprimary collective veins.

Several varieties have been described (see Furtado, 1941), mostly of

little botanical interest though some may be usefully recognised at the

level of cultivar. However, in the Philippines, a form that has rather deeply

sinuate leaf margins has been given varietal status (these specimens are

listed as ‘var. heterophylla’ below). Although slightly wavy leaf margins

are commonly found in A. macrorrhizos s.s. throughout its range, variation

to this extent seems confined to the Philippines. Merrill (1922: 185) regarded

this taxon as a synonym of A. portei. This status requires re-evaluation

following field work. It is possibly a hybrid between A. macrorrhizos and

A. portei, since horticultural crosses between A. portei and entire-leaved

species yield sinuate-margined hybrids. No data are yet available on the

arrangement of the inflorescences. In the meantime, I provisionally place

it in the synonymy of A. macrorrhizos, as it seems to intergrade with

somewhat wavy-margined forms that are typical of A. macrorrhizos

elsewhere. It does not intergrade with A. portei in leaf margin features.

This form is illustrated in the horticultural account of Burnett (1984, fig.

83), where it is regarded as a hybrid.

Other specimens seen: “Typical form’ - Luzon, Pampanga, Mt Arayat, Bolster

92 (UC); Luzon, Benguet, Clemens 18710 (UC); Luzon, Mountain Prov.

Baywinan, Banaue, Ifugao, Conklin & Buwaya I-951 (GH, L); Luzon,

Benguet, Baguio, Elmer 8818 (E, G, K, L, NY, US); Luzon, Irosin, Mt

Bulusam, Elmer 15455 (CAL, G, GH, K, L, NSW, NY, P, US); Luzon,

Laguna Prov., Los Bafios (Mt Maquiling) Elmer 18243 (CAL, GH, K, US);

Central Luzon, Loher 2438 (CAL, K, US); Luzon, Manila Province, Lueva

da Montalban, Loher 2436 & 7038 (both K); Luzon, Benguet, Loher 5029

(K); Luzon, Tayabas Prov., Loher 14020 (UC); Puerto Prince, Ile Paragua,

Marche 116 (P); Leyte, W of Tanauan, Malaquikay Barrio, Nicolson 815

(US); Luzon, Laguna Prov. College of Agriculture, behind Baker Hall,

Molawin Ck, Nicolson 838 (US); Luzon, Rizal Prov., Mt Irig, Ramos BS

41989 (US); Batan, Ramos BS 80647 (NY); Luzon, Sorsogon Prov., Pilar,

Regalado & Ugalde PNH 37494 (L, mixed with Cyrtosperma merkusii);

22 Gard. Bull. Singapore 51 (1999)

Polillo, Robinson BS 6997 (US); Luzon, Mt Polis pass, Walker 7513 (US);

Central Luzon, Bataan Prov., Mt Mariveles, Lamao R., Williams 320 (NY,

US);

‘Var. heterophylla’ - NE Polillo Island, Karlagan, Fox 280 (GH, L); Luzon,

Benguet, Yonglon, Loher 5029 (K); Luzon, Bataan Prov., Lamao R., Mt

Mariveles, Whitford 1278 (K, NY, US).

7. Alocasia maquilingensis Merr.

Alocasia maquilingensis Merr., Philippine J. Sci 13 (1918) 3, Enum. Philipp.

Fl. Pl. 1 (1923) 185; Elmer, Leafl. Philip. Bot. 10 (1938) 3615. - Type:

Philippines, Luzon, Laguna Prov., Mt Makiling, C.F. Baker 868 (?PNH,

presumed destroyed). Neotype: Philippines, Luzon, Laguna Prov., Mt

Makiling, E.D. Merrill 7153 (US; isoneo, BM, designated here).

[Alocasia vulcanica Elmer, mss.|

Robust caulescent herb; leaves several together, without interspersed

cataphylls; petiole to ca. 1 m long, thickly pubescent to glabrescent,

sheathing in the lower third to half; blade broadly ovato-sagittate ca. 55—90

cm long; anterior lobe 50-60 cm long, about as wide at base as long, with

ca. 8 primary lateral veins on each side of the anterior costa diverging at

ca. 60°, especially the proximal ones sometimes bearing subsidiary veins

intermediate in thickness between primary and secondary venation; primary

and subsidiary veins with conspicuous glands in their axils; secondary

venation rather prominent abaxially, numerous and close-spaced, arising

from the primary venation at ca. 70° then deflected marginally and organised

into distinct interprimary collective veins; primary and secondary veins

strongly pubescent to almost glabrous, but at least with some hairs on the

finest secondary veins especially near the blade margin; posterior lobes

mostly broadly rounded, sometimes squared off, ca. 20-30 cm long; posterior

costae naked in the sinus for ca. 2 cm; mflorescences ca. 10-30 clustered

together in the centre of the crown; peduncle ca. 15 cm long, pubescent;

spathe white, ca. 10 cm long; lower spathe ca. 2 cm long, separated from

the blade by a rather weak constriction slightly above the level of the

sterile interstice of the spadix; limb narrowly oblong lanceolate, the tip

obtuse to subtruncate and abruptly mucronate for ca. 5 mm; spadix equalling

to slightly exceeding the spathe in length, stipitate for ca. 6 mm; female

zone ca. 1 cm long; ovaries ovoid, ca. 1.5 mm tall; style slender, ca. 0.8 mm

long; stigma broad, weakly lobed; sterile interstice very short, not or hardly

attenuated, ca. 3 mm long (more or less one whorl of synandrodia); male

Alocasia in the Philippines 23

zone 1.7-2 cm long, cylindric, ca. 6 mm diam.; synandria mostly

rhombohexagonal, ca. 1.5—2 mm across, those in the middle of the male

zone longitudinally (with respect to spadix) compressed (?always); appendix

ca. 6-7 cm long, cylindric, distally tapering to a point, composed of low

sinuous compressed staminodia; fruiting spathe ?red, ovoid to pyriform,

4.5 x 2 cm, longitudinally dehiscent; fruits red.

Distribution: Endemic to the Philippines; Luzon, Mindanao, Leyte, Panay.

Habitat: On slopes in primary rain forests at low to medium elevations.

Notes: In the protologue of A. maquilingensis, Merrill (loc. cit.) cited

Baker 868 as the ‘type’, as well as citing a number of other collections. Of

all the specimens cited, it appears that only Merrill 7153, designated here

as neotype, is still extant. I infer that Merrill’s designation of one of the

cited specimens as the ‘type’ has the consequence that neotypification,

rather than lectotypification, is required under the circumstances.

The pubescent petioles and main veins are unique in Philippines

Alocasia, but also found (constantly or variably) in New Guinea A. nicolsonii

A. Hay, Sulawesi A. suhirmaniana Yuzami & A. Hay and A. celebica

Engl., West Malesian A. puber (Hassk.) Schott and A. inornata Hallier f,,

Sumateran A. arifolia Hallier f. and Bornean A. sarawakensis M. Hotta

(the latter only in juvenile plants), A. reginae N.E. Br. and A. scabriuscula

N.E. Br.. Plants from the type locality are much hairier than those from

elsewhere. This species can be mistaken for A. macrorrhizos (q.v.), which

differs in being totally glabrous, having hardly prominent secondary

venation that does not form interprimary collective veins, and much larger

inflorescences that do not form central clusters.

Other specimens seen: Luzon, Sorsogon Prov., Mt Bulusan, Elmer 16181

(BO, K, NSW, NY, US). Luzon, Laguna Prov., Mt Makiling, Elmer 17955

(BO, CAL, G, GH, K, L, NSW, P, US); Luzon, Laguna Prov. Los Bafios

(Mt Makiling), Elmer 18239 (BO); Luzon, Laguna Prov., Mt Makiling,

along path from College of Agriculture to summit of Mt Makiling, Nicolson

689 (US); Mindanao, Davao Prov., Upper Baractan (foot of Mt. Apo)

Nicolson 728 (US); Leyte, Vicinity of Dagami, nr Barrio Putok, Nicolson

814 (US); Luzon, Laguna Prov., Mt Makiling, along path from College of

Agriculture, Nicolson 846 (US); Panay, Capiz Prov., Jamindan, Ramos &

Edano BS 31472 (K, US); Mindanao, Zamboanga Dist., Malangas, Ramos

& Edano BS 36829 (K, US); Mindanao, Lanao Prov., Porog, Zwickey 791

(H).

24 Gard. Bull. Singapore 51 (1999)

8. Alocasia micholitziana Sander

Alocasia micholitziana Sander, Gard. Chron. ser. 3, 51 suppl. (1912) xv, fig.

9; N.E. Brown, Bot. Mag. (1913) t. 8522. K. Krause & Engl., Pflanzenr. 71

(IV.23E) (1920) 88; Merr., Enum. Philipp. Fl. Pl. 1 (1922) 88; Burnett,

Aroideana 7 (1984) 93, figs 28-31. - Type: Gard. Chron. ser. 3, 51 suppl.

(1912) fig. 9 [sterile]. Epitype: Cult. Kew ex hort. Sander (leg. ico

1911, N.E. Brown s.n. (K, designated here).

Moderately robust herb; stem erect to decumbent, to ca. 50 cm long, ca. 4

cm diam.; leaves ca. 4-7 together, not interspersed with cataphylls; petiole

to 45 cm long, sheathing in the lower ca. quarter to third, mottled brownish,

reddish and/or purplish; blade rich velvety matt deep green adaxially, paler

abaxially, sagittate, ca. 40 cm long x 13 cm wide, very shallowly to shallowly

peltate; costae and primary veins white adaxially; anterior lobe ca. 20 cm

long, rather broadly triangular, the margin very strongly to shallowly but

distinctly undulate; anterior costa with 4—5 primary lateral veins on each

side diverging at ca. 60—80° then curving distally to the margin; secondary

venation flush on both sides, arising from the primary venation at a wide

angle (ca. 90-100°) then curving to the margin; interprimary collective

veins not or hardly differentiated; posterior lobes diverging at ca. 60—90°,

narrowly triangular, the outer margins undulate, to ca. 11 cm long;

inflorescences to ca. 4 together; peduncle to ca. 20 cm long; spathe ca. 14

cm long; lower spathe ovoid, ca. 3 cm long, green, separated from the limb

by a rather abrupt constriction; limb ca. 11 cm long, basally somewhat

reflexed and distally cucullate at anthesis, opening wide; spadix ca. three

quarters the length of the spathe, sessile, ca. 9 cm long; female zone ca. 1

cm long (dry) x ca. 7 mm; ovaries globose, ca. 2.5 mm diam.; style ca. 1

mm, rather abruptly differentiated from ovary; stigma subglobose, weakly

lobed; sterile interstice slender, ca. 7 mm long x 2 mm diam. at narrowest,

the lower ring of synandrodia paler and larger than upper ones; male zone

more or less cylindric, ca. 1.2 cm long x 5 mm wide; synandria rhombo-

hexagonal, ca. 1.2 mm diam. (dry), opening by apical pores not capped by

synconnective; appendix ca. 6 cm long, tapering to a narrow blunt tip, not

or faintly constricted at junction with male zone, composed of irregular

low elongate staminodia; fruiting spathe ca. 4 cm x 2 cm; fruit ca. 5 m

diam.

Distribution: Endemic to Luzon.

Habitat: In primary and secondary forest and roadsides at ca. 1200-1500 m

altitude. This species is said to be common and widespread, and presumably

occurs over a somewhat wider altitudinal range than the limited available

data indicate.

Alocasia in the Philippines a5

Notes: The plant illustrated in the figure accompanying the protologue is

sterile and an epitype is needed to establish the application of this name

more firmly. Selection of the designated specimen is based on the facts

that it is fertile and preserved from a plant cultivated at Kew, annotated by

Brown as Alocasia micholitziana and as obtained from Sander’s nursery

and originally collected by Micholitz after whom the species was named.

Those facts would appear to form an adequate indirect link between the

specimen and the plant on which Sander based his description; indeed they

are quite probably clonally identical.

Madulid (pers. comm.) notes that this species occurs in several

varieties. These are not clearly evident from herbarium material, though

there is sufficient variation for a number of horticultural varieties to have

been recognised (Burnett, 1984). A. micholitziana is distinguishable from

A. sanderiana, with which it shares white major venation and undulate leaf

margins, by its paler, brighter, matt green leaf surface - not purple abaxially,

and by the leaf bases not subtended by cataphylls except at the beginning

of a sympodial module. Moreover, the leaf of A. micholitziana is less

deeply peltate than that of A. sanderiana.

Other specimens seen: Luzon, Ifugao Prov., Banaue, Bogner 1631 (M);

Ifugao prov., Banaue, Bayninan, Conklin & Buwaya PNH 80620 (GH, L);

Laguna Prov., Los Banos (Mt Maquiling), Elmer 17769 (BO, CAL, GH,

NY); Benguet Prov., Loher 2441 (K); Rizal Prov., Loher 15005 (UC);

Ifugao Prov., Banaue, Poitan, Liwang, Mendoza PNH 37182 (L); Laguna

Prov., Mt Makiling, along path from College of Agriculture, Nicolson 843

(L,:US).

9. Alocasia portei Schott

Alocasia portei Schott, Bonplandia 10 (1862) 148 [‘Alocasia? (Schizocasia)

portei’|; Beccari in Bull. Soc. Tose. di Ort. 4 (1879) 300 [n.v.]; Engl. in A. &

C. DC., Monogr. Phanerogam. 2 (1879) 645; Naves, Nov. App. (1882) 294;

Merr., Fl. Manila (1912) 132, Enum. Philipp. Fl. Pl. 1 (1922) 185; Burnett,

Aroideana 7 (1984) 127, fig. 89. - Schizocasia portei [Engl. in A. & C. DC.,

Monogr. Phan. 2 (1879) 495, invalidly published] Engl. in Beccari, Malesia

1 (1883) 295; K. Krause in Engl., Pflanzenr. 71 (IV.23E) (1920) 117. -

Neotype: Schott Icones no. 3085 (W; fiche 72:b5 in the microfiche edition;

designated here).

Schizocasia regnieri L. Linden & Rodigas, Ill. Hort. 34 (1887) descr. ad t. 6

& Gartenfl. 36 (1887) 333. - Type: Ill. Hort. 34 (1887) t. 6.

26 Gard. Bull. Singapore 51 (1999)

Massive arborescent pachycaul to 6 m tall; stem erect, to ca. 40 cm diam. at

base (thickening with age), distally to ca. 15 cm diam., older parts developing

tessellated ‘bark’; leaves several together, more or less erect; petiole to ca.

1.5 m long, yellowish to dark green mottled chocolate, sheathing in the

lower third; blade sagittate and deeply pinnatifid, dark green, coriaceous;

anterior lobe to ca. 1.5 m long, with up to 10 linear-lanceolate round-

tipped segments each with a primary lateral vein diverging from the anterior

costa at ca 80—90°; margins strongly crispate; primary veins prominent below,

tinged purple-brown or yellowish, with conspicuous axillary glands on

abaxial side; secondary venation flush, arising from primary at ca. 80—90°

and interspersed at ca. 2-3 cm intervals with abaxially slightly prominent

stronger veins (but nearer in size to secondary than primary) with glands

in their axils; inflorescences in numerous pairs clustered in the centre of

the leaf crown, subtended by conspicuous brown-mottled lanceolate

cataphylls: peduncle to ca. 30 cm long; spathe to 40 cm long; lower part

subcylindric, dark brownish green, ca. 5 cm. long; limb linear oblong-

lanceolate, at first erect and canoe-shaped and cucullate, thence reflexed

at the constriction, pale brownish, mottled and streaked chocolate brown,

membranous: spadix somewhat shorter than the spathe, to 32 cm long;

female zone sessile, ca. 4 cm long x 1.5 cm thick; ovaries subglobose, ca. 2.5

mm diam.; style very short; stigma weakly lobed; sterile interstice ca. 2 cm

long, narrowed, corresponding with spathe constriction, white; male zone

whitish, ca. 8 cm long, cylindric, ca. 1.5 cm diam.; synandria rhombo-

hexagonal, ca. 1.2 mm diam. (dry); thecae capped by synconnective;

appendix basally slightly thicker than male zone, then subcylindric and

distally tapering; fruiting spathe ovoid, ca. 10 cm long x 5 cm thick.

Distribution: Endemic to the Philippines; Luzon. However, A. portei is

anecdotally reported to be common and widespread in the Philippines,

although botanical collections have been made on only very few occasions,

probably on account of its enormous size.

Habitat: Secondary forest at low to medium elevations.

Notes: The protologue of Alocasia portei (Schott, loc. cit.) describes a leaf

only. No collector or collection was cited, and if type material was preserved

at W, it is presumably now destroyed. The designated neotype is a drawing

Schott had caused to be prepared, which, like the protologue, includes

only a leaf. However, it conforms unequivocally to the traditional and

current concept of this species, and it seems not unreasonable to suppose

that it was based on the original material.

S. regnieri appears to differ only in having a yellowish ground colour

Alocasia in the Philippines 27

to the petioles and costae. It was originally described from living material

imported to Europe from Thailand where it was presumably cultivated.

Other specimens seen: Cult. Kew ex Hort. Veitch, N.E. Brown s.n. (K):

Cult. Munich Bot. Gard., Bogner 1768 (M): Cult. Hort. Bot. Univ. Kiel.

Engler Araceae No 109 (GH, K, L, M, SING, US): Cult. Kew ex Hort.

Bull, N.E. Brown s.n. (K); Luzon, Laguna Prov., Mt Makiling, Sulit PNH

22895 (GH, L); Luzon, Laguna Prov., cult. at College of Forestry, Nicolson

794 (K, L, US).

10. Alocasia ramosii A. Hay, sp. nov.

Ab Alocasia heterophylla spatha abrupte constricta, synandrodiis minoribus.

nervulis densioribus, costis posticis minus acute divergentibus, vena

submarginali deficienti differt. - TYPUS: Philippines, Luzon, Laguna Prov..

Molawin Ck, path between Colleges of Agriculture and Forestry, 20 Oct

1960, D.H. Nicolson 752 (US, holo).

[Alocasia heterophylla auct. non (Presl) Merr.: Elmer, Leafl. Philipp. Bot.

10 (1938) 3613.]

Small herb to ca. 40 cm tall; leaves 2-several together, not each subtended

by cataphylls; petioles to ca. 25 cm long, sheathing in the lower third: blade

membranous, hastate to sagittate, to ca. 28 cm long: anterior lobe narrowly

to rather broadly triangular, the margin entire to very shallowly undulate:

anterior costa impressed adaxially, with 4-5 primary lateral veins on each

side diverging at ca. 60° and running more or less straight to the margin

(cf. A. heterophylla), with glands in the axils usually conspicuous; secondary

venation arising at a wide angle from primary veins, close-spaced (ca. 1.5-

2 mm apart; cf. A. heterophylla, A. culionensis), then abruptly deflected

into a somewhat disorganised zig-zag interprimary collective vein and/or

running to the margin; posterior lobes diverging at an obtuse (hastate) or

acute (sagittate) angle, narrow to rather broad, tapering to blunt points, 9-

13 cm long; posterior costae naked in the sinus for 1-3 cm, never peltate

except in seedlings; inflorescences 1-2 together; peduncle ca. 10-15 cm

long, slender; spathe ca. 5 cm long; lower spathe ovoid, ca. 1.3 cm long,

somewhat asymmetric due to adnation with spadix stipe, differentiated

from the limb by a strong constriction; limb lanceolate, somewhat canoe-

shaped, slightly deflected at base, thence up-curved; spadix stipitate for ca.

5 mm, with the stipe almost completely adnate to the spathe: female zone

ca. 8 mm long; ovaries subglobose, ca. 1 mm diam., stigma subsessile,

button-like; sterile interstice short, ca. 3-5 mm long, corresponding with

28 Gard. Bull. Singapore 51 (1999)

Figure 5. Alocasia ramosii A. Hay

Nicolson 752 - A: habit; B: venation; C: inflorescence (part of spathe removed). - Scale: bar to

A =10cm, to B=6cm, to C=2 cm.

Alocasia in the Philippines 29

spathe constriction; male zone ca. 8 mm long, subcylindric, ca. 4 m diam.;

synandria rhombo-hexagonal, ca. 1 mm diam., not capped by synconnective;

appendix tapering to a point, ca. 1.2 cm long, slightly curved to ventral side

of spathe; infructescence subglobose, ca. 1.7 cm diam.

Distribution: Endemic to the Philippines: Luzon, Negros, Panay.

Habitat: Lowland rain forest, often near streams, to 400 m altitude.

Notes: The specific epithet commemorates Maximo Ramos (died May 11

1932), prodigious collector of Philippine and Bornean plants, in the employ

of the Bureau of Science at Manila (van Steenis-Kruseman, 1950).

Little complete fertile material has been seen.

Other specimens seen: Luzon, Sorsogon Prov., Irosin (Mt Bulusan), Elmer

16344 (G, L, NY); Luzon, Laguna Prov., Los Bafios, (Mt Maquiling) E/mer

17769 (K, US - mixed; see also A. micholitziana); Luzon, Rizal Prov.,

Loher s.n. (UC); Luzon, Tayabas Prov., Umiray, Loher s.n. (UC); Panay,

Antique Prov. McGregor BS 32415 (K); Luzon, Laguna prov., resthouse

nr mud springs 400 m along path from College to summit of Mt Makiling,

Nicolson 684 (US); Luzon, Sorsogon Prov., along pathside around Lake

Bulusan, Nicolson 722 (US); Luzon, Laguna Prov., Molawin Ck, path

between Colleges of Agriculture and Forestry, Nicolson 753 (US); Negros

Oriental, Mt Canlaon, above Canlaon, Nicolson 832 (US); Luzon, Laguna

Prov., path from College to summit of Mt Makiling, Nicolson 845 (US);

Luzon, Camarines Prov., Mt Isarog, Ramos BS 22059 (US); Luzon, Rizal

Prov., Montalban, Robinson BS 6148 (US).

11. Alocasia sanderiana W. Bull

Alocasia sanderiana W. Bull, Retail List (1884) 11; Kew Bull. (1895) App.

2, 32; Merr., Fl. Manila: 132 (1912), Enum. Philipp. Fl. Pl. 1 (1923) 186; K.

Krause & Engl., Pflanzenr. 71 (I1V.23E) (1920) 104; Burnett, Aroideana 7

(1984) 100, fig. 40. - Schizocasia sanderiana (W. Bull) Engl. in Bot. Jahrb.

Syst. 25 (1898) 26. - Type: W. Bull, Retail List (1884) un-numbered fig. p. 3

(sterile). Epitype: Cult. Kew from material ex Philippines supplied by W.

Bull, 11 Aug 1887, N.E. Brown s.n. (K, designated here).

Moderately robust herb to 60 cm tall; stem decumbent to creeping, ca. 15

cm long, ca. 2 cm thick; leaves solitary to few together, interspersed with

cataphylls (precise arrangement unclear from herbarium material);

cataphylls papery, narrowly lanceolate; petiole to 60 cm long, sheathing in

30 Gard. Bull. Singapore 51 (1999)

lower fifth to quarter; blade sagittate, deep glossy green adaxially, purple

abaxially (?always), peltate for 25-30% of the length of the posterior lobes,

to ca. 40 cm long x 20 cm wide; margin deeply undulate to sub-pinnatifid;

anterior lobe to 24 cm long; anterior costa with 3-4 often but not always

opposite primary lateral veins on each side diverging at ca. 80—100°, these

somewhat acropetally deflected towards the tip of each segment; primary

veins and costae white to yellowish; secondary veins emerging from the

primary at a wide angle, thence deflected marginally, not or hardly forming

interprimary collective veins; inflorescence mostly paired; peduncle slender,

ca. 14-18 cm long; spathe 6-10 cm long, lower part 1.5—2 cm long, ovoid;

limb 4-8 cm long, more or less canoe-shaped; spadix somewhat shorter

than to half the length of the spathe, stipitate for ca. 5 mm; female zone ca.

1 cm long; ovaries ca. 2 cm diam., subglobose; stigma sessile, rather sharply

3—-4-lobed; sterile interstice corresponding to spathe constriction, ca. 5 mm

long, tapering distally; male zone 1.2 cm long, ca. 4-5 mm thick (dry);

synandria rhombo-hexagonal with the thecae ventrally but hardly laterally

joined; synconnective centrally impressed, marginally slightly exceeding

but not capping the thecae; appendix short, equalling the male zone,

cylindric then abruptly tapering in distal 3-4 mm, slightly narrower than

male zone (dry).

Distribution: Endemic to Mindanao.

Habitat: Old forest in damp ravines at low elevation.

Notes: The peltate dark green, white veined leaves interspersed with

cataphylls and the stipitate spadix and pointed-lobed stigmas indicate that

this species is a segregate of the Alocasia longiloba complex, of which A.

boyceana (q.v.) appears to be another facies.

Like A. zebrina, this species was CITES listed, but has now been

removed from CITES schedules. A. sanderiana was at one time much in

demand horticulturally. It has been used extensively in hybridisation (see

Burnett, 1984).

Other specimens seen: Cult. Kew, 4 Aug 1886, N.E. Brown s.n. (K); Surigao,

Loher 2439 (K); cult. Manila, Loher 2440 (K, US); Lanao Prov., Pantar,

Mearns 129 (US); Butuan sub prov., Agusan R., Merrill 7276 (US); Lanao

Prov., Cawit, Zwickey 413 (GH).

12. Alocasia scalprum A. Hay, sp. nov.

Ab aliis speciebus Alocasiis Philippinarum lamina folil parva tenuiter

Alocasia in the Philippines 31

lanceolata subfalcata, lobis posticis minutis, venis impressis differt. - TYPUS:

Cult. Botanischer Garten Miinchen, origin unknown, 1983, J. Bogner 1659

(M, holo).

[Alocasia cv. Samar Lance; see Burnett, Aroideana 7 (1984) 99, fig. 38].

Diminutive to small herb; leaves several together; petiole shorter than the

blade, to ca. 10 cm long, sheathing in the lower third, green mottled darker

green; blade narrowly lanceolate, slightly falcate, 15—25 cm long x 2-5 cm

wide; anterior lobe ca. 14-22 cm long, with the margin entire and slightly

revolute (dry); anterior costa with 2—3 primary lateral veins on each side

diverging at ca. 45°, like the costae and secondary venation deeply impressed

adaxially, prominent abaxially; secondary venation arising from the primary

at a low angle and running to margin, not or hardly forming interprimary

collective veins; posterior lobes much reduced, forming narrowly triangular

auricles 1.5—-3 cm long; posterior costae poorly developed, naked in the

sinus for ca. 1 cm to almost throughout their length; inflorescence solitary;

peduncle subequalling the petioles at anthesis; spathe ca. 6 cm long; lower

spathe ca. 1.5 cm long, subcylindric, differentiated from the limb by a

weak, gradual constriction; limb lanceolate, green, mucronate for ca. 5

mm; spadix somewhat shorter than the spathe, stipitate for 5 mm, the stipe

partly adnate to the spathe; female zone ca. 6 mm long, obliquely inserted

on the stipe, narrow, ca. 3 mm thick (dry); ovaries subglobose, ca. 1.2 mm

diam.; stigma sessile, discoid, almost as wide as the ovary, weakly 2-3-

lobed; sterile interstice ca. 9 mm long, the lowermost synandrodia reduced,

thence the interstice ca. 3 mm thick, slightly tapering distally; male zone

somewhat thicker than the upper part of the interstice, 4.5 mm diam, 8 mm

long; synandria rhombo-hexagonal, strongly wavy-edged, ca. 1.5 mm diam.;

appendix 1.6 cm long, tapering; infructescence unknown.

Distribution: Philippines, said to be known only from Samar.

Habitat: Unknown.

Notes: This species is apparently quite well-known in cultivation in the

Philippines, but is not represented by any wild-collected herbarium material.

Burnett (1984) notes that what is considered in the horticultural community

to be one species is highly variable and he lists four selected varieties all

said to have originated from the island of Samar. The only material I have

seen in flower, which forms the holotype, seems exactly to match the

cultivar known as ‘Samar Lance’. Although it seems probable that the

other forms mentioned and illustrated in Burnett (loc. cit.) are conspecific

32 Gard. Bull. Singapore 51 (1999)

Figure 6. Alocasia scalprum A. Hay

Bogner 1659 - A: leaf; B: leaf blade; C: inflorescence; D: spadix with part of spathe removed;

E: pistils; F: neuter organs of sterile interstice; G: synandria, from above. - Scale: bar to A, B,C

=2¢cm,io D = 1 em,to E.R G=2 mm

Alocasia in the Philippines 33

with A. scalprum, this remains to be verified.

The specific epithet alludes to the knife-shaped leaf blade.

13. Alocasia sinuata N. E. Brown

Alocasia sinuata N.E. Brown, Gard. Chron. n.s. 24 (1885) 678; K. Krause

& Engl., Pflanzenr. 71 (IV.23E) (1920) 102; Merr., Enum. Philipp. Fl. Pl. 1

(1922) 186. - Type: Cult. Kew ex Hort. W. Bull., 22 Oct 1885, N.E. Brown

s.n. (K holo).

[Alocasia Quilted Dreams - Burnett, Aroideana 7 (1984) 97, fig. 33].

[Alocasia bullata Hort., nom. inval.|

Herb to ca. 35 cm tall; stem [dimensions not known]; leaves several together;

petiole ca. 19 cm long, sheathing in the lower ca. fifth to quarter; blade

sagittate to ovato-sagittate, peltate or not, somewhat to markedly bullate,

somewhat to thickly coriaceous, dark green (sometimes darker about main

veins) adaxially, paler abaxially, with the margin sometimes very shallowly

sinuate, 22-27 cm long; anterior lobe widest at to ca. 4 cm from the base,

the apex broadly acute to obtuse and apiculate for 0.5—1 cm; anterior costa

with 2-3 primary lateral veins on each side, diverging at 60-80° and adaxially

impressed; axillary glands inconspicuous; subsidiary veins numerous (i.e.

ca. quarter to half the secondary venation thicker than the rest); secondary

and subsidiary venation forming rather ill-defined interprimary collective

veins running, like the primary venation, into a somewhat conspicuous

intramarginal vein 1-3 mm from the margin; posterior lobes acute, ca.

third to half the length of the anterior, connate for O-half of their length,

the inner sides more or less linear-lanceolate; posterior costae diverging at

45-90°, with lamina to the sinus when leaf not peltate; inflorescences

?solitary to paired to clustered, subtended by lanceolate cataphylls to ca. 7

cm long (in the type these with reduced petiole and leaf blade); peduncle

about three quarters to equalling the length of the petiole at anthesis;

spathe 7—7.5 cm long, green; lower spathe 1.5—2 cm long, ovoid; limb broadly

oblong-lanceolate, green, apiculate for ca. 0.5 cm; spadix somewhat shorter

than the spathe, 46.5 cm long, very shortly stipitate with somewhat oblique

insertion, white except for green ovaries; female zone ca. 6 mm long, 3-8

whorls of somewhat lax pistils; ovary globose, ca. 2 mm diam.; stigma

pronounced - almost as large as the ovary, subsessile, thickly discoid to

weakly 3-lobed; interstice ca. 7 mm long, the lower half of incompletely

connate neuter organs, the upper half of sterile synandria, somewhat

attenuate above and corresponding with spathe constriction, 2.5 mm diam.

34 Gard. Bull. Singapore 51 (1999)

(dry); male zone 1—1.5 cm long, somewhat obconic, distally ca. 5 mm diam.;

synandria rhombohexagonal, 4-5-merous, 2-3 mm diam., thecae not

overtopped by synconnective; appendix 1.5-4.5 cm long, basally isodiametric

with top of male zone then tapering distally; infructescence unknown.

Distribution: Endemic to the Philippines - Leyte, Mindanao, Palawan.

Habitat: Unknown.

Notes: Alocasia sinuata is botanically poorly known, though it has been in

cultivation for over a century. Two forms exist in cultivation today under

the names Alocasia Quilted Dreams and Alocasia Bullata (the latter used

as a pseudobinomial but it is botanically invalid). I have seen only one

collection from the wild, unfortunately without habitat notes.

The type was at first said to have originated in the ‘Malay

Archipelago’ (N.E. Brown, loc. cit.). However, the label has subsequently

been altered in Brown’s hand to indicate that the plant originated in the

Philippines. Observations on the species’ alleged large size made by W.

Bull which Brown (loc. cit.) quoted (‘a full grown plant would be about 4

or 5 feet in diameter’), notwithstanding the much smaller dimensions of

the type plant and matching material, are evidently incorrect (other material

examined is somewhat smaller than the type).

The collection cited below originating in Palawan is included with

some doubt as the sterile interstice is much attenuated and the secondary

venation seems not to form subsidiary veins. In other respects, however,

the specimen matches A. sinuata.

Other specimens seen: Cult. University of Florida Agricultural Research

Station ex Leyte Island, Croat 56900 (MO); Cult. Missouri Botanical Garden

ex cult. Manila Memorial Park, Croat 57106 (MO); Cult. Missouri Botanical

Garden ex Palawan, Croat 57188 (MO); Mindanao, Surigao Prov., without

specific locality, Ramos & Pascasio BS 34353 (K, US).

14. Alocasia zebrina Schott ex van Houtte

Alocasia zebrina Schott ex van Houtte, Fl. des Serres 15 (1863) t. 1541-

1542; C. Koch & Veitch, Wochenschr. Gartn. Pflanzenk. 6 (1863) 331;

Engl. in A. & C. DC., Monogr. Phanerogam. 2 (1879) 507; Naves, Novis.

App. (1882) 294; Merr., Fl. Manila (1912) 133, Enum. Philipp. Fl. Pl. 1

(1922) 186; K. Krause & Engl., Pflanzenr. 71 (IV.23E) (1920) 105; Elmer,

Leafl. Philip. Bot. 10 (1938) 3616; Burnett, Aroideana 7 (1984) 104, figs

46, 47. - Type: Fl. des Serres (1863) 15 t. 1541-1542.

Alocasia in the Philippines a5

Alocasia wenzelii Merr., Philipp. J. Sci. 13, Bot. (1918) 2, Enum. Philipp.

Fl. Pl. 1 (1922) 186. - Type: Philippines, Leyte, 28 Mar 1913, C.A. Wenzel

97 (?PNH, holo, presumed destroyed; BM, G, GH, US, iso).

Moderately robust to robust herb to 1.8 m tall; leaves several together, not

interspersed with cataphylls save at the beginning of a sympodial module;

petiole to 1.1 m long, sheathing in the lower third, pale green, obliquely

streaked brownish and/or darker green; blade rather narrowly sagittate,

ca. 45-100 cm long, rather coriaceous, rich glossy green; anterior lobe

triangular to somewhat ovate, to 80 cm long, with (4-)6—-10 primary lateral

veins diverging from each side of the anterior costa at ca. 80—90° and

running almost straight to the margin, bearing small axillary glands on the

abaxial side; secondary venation flush with the lamina and inconspicuous

on both surfaces of the blade, arising from the primary veins at a wide

angle then deflected to the margin without forming interprimary collective

veins; posterior costae diverging at an acute or right angle, bearing lamina

to the sinus and often minutely peltate; posterior lobes elongate, three

fifths to two thirds the length of the anterior lobe; inflorescences paired,

not in large clusters; peduncle ca. 25 cm long; spathe 13-16 cm long; lower

part 2.2—3 cm long, narrowly ovoid, separated from the limb by an abrupt

constriction corresponding with the sterile interstice of the spadix; limb

narrowly lanceolate, erect; spadix somewhat shorter than the spathe,

stipitate for ca. 7 mm; stipe conic; female zone 1.4—-2 cm long x ca. 8 mm

wide; ovaries globose; stigma weakly lobed, subsessile; sterile interstice ca.

1 cm long, distally attenuate; male zone cylindric, ca. 3-4.5 cm long x 5 mm

thick (dry); synandria rhombo-hexagonal, 1.5-2 mm diam., capped by

synconnective; appendix distinctly narrower (at least in dry state) than

male zone, 4.5—5 cm long, tapering; fruiting spadix globose, ca. 4.5 cm

diam.

Distribution: Endemic to the Philippines - Luzon, Mindanao, Leyte, Samar,

Biliran, Alabat.

Habitat: Rain forest at low to medium elevation.

Vernacular names: Badiang, Handuroi.

Notes: Easily distinguished from all other Philippine Alocasia species by

the rather narrowly sagittate leaves with striped petioles and rather long

acute posterior lobes bearing lamina to the sinus but with the blade not or

only very narrowly peltate.

36 Gard. Bull. Singapore 51 (1999)

Merrill (1922: 186) suggested that A. wenzelii was no more than a

robust form of A. zebrina, a view with which I concur. This species is

rather variable and a number of different forms are in cultivation.

Other specimens seen: Cult. Hort. Veitch ex Luzon, Anon. s.n. April 1863

(K); Luzon, Laguna Prov. Mt Makiling, Bartlett 15677 (GH, PNH); Luzon,

Sorsogon Prov., Mt Bulusan, Bogner 1648 (M); Leyte, Palo, Elmer 7292

(E); Luzon, Sorsogon Province, Irosin (Mt Bulusan), E/mer 14911 (BO, G,

GH, K, L, NY, P, US); Luzon, Laguna Prov. Los Bafios (Mt Maquiling)

Elmer 17790 (BM, BO, CAL, G, GH, K, L, NSW, NY, P, US); Biliran,

McGregor BS 18929 (US); Mindanao, Mt Apo, Todaya vill., Mearns s.n.

(US); Mindanao, Upper Baractan (base of Mt Apo), Davo City, Nicolson

727 (US); Samar, Loquilocan Barrio, Nicolson 810 (US); Leyte, Dagari

area, nr Putok R., Nicolson 823 (US); Luzon, Laguna Prov., Mt Makiling,

Nicolson 844 (US); Alabat Island, Ramos & Edano BS 48275 (BRI, NY);

N. Luzon, Ifugao Prov., Patukan Valley, Poblacion II, Kangan, Reynoso et

al. PPI 7222 (US); N. Luzon, Ifugao Prov., Bangaan View Point, Banaue,

Reynoso et al. PPI 7138 (US); Luzon, Laguna Prov. Los Banos, Robinson

BS 6731 (US)x:

Excluded, Dubious or Doubtfully Philippine Species

Alocasia liervalii Hérincq

Alocasia liervalii Hérincq, Hortic. Frang. (1869) 243. Type: none found.

The description is vague, of sterile material and not accompanied by an

illustration. Hérincq was a botanist and gardener associated with the Musée

d’Histoire Naturelle at Paris, but no pertinent preserved material has been

found there. Indeed, the whereabouts of his herbarium and types, if any

exist, is unknown (Stafleu & Cowan, 1979). This plant was said by Hérincq

to have been collected by Porte in the Philippines and introduced into

commercial horticulture by Lierval. The only clue to the identity of A.

liervalii is Hérincq’s allusion to the large size of the leaf blades and the

mottled petioles, which may suggest that this is a synonym of the earlier A.

zebrina.

Alocasia maxumiliani Maedicke

Alocasia maximiliani (‘maximiliana’) Maedicke, Mdéllers Deutsche Gart.-

Zeit. 26 (1911) 134; Burnett, Aroideana 7 (1984) 93, excl. descr. et fig. 27

Alocasia in the Philippines a}

i.e. Alocasia zebrina. - Type: Mollers Deutsche G§rt.-Zeit. 26 (1911) 133,

fig. I. ?= Xanthosoma sp.

This binomial is not in Index Kewensis, and I have not been able to find

any earlier use of it. Burnett (1984) attributed this species to the Philippines.

The basis for this is not clear to me, since Maedicke did not cite a

provenance. Maedicke’s description is of a sterile plant cultivated in the

Palmengarten at Frankfurt. The description and accompanying photograph

indicate it is a Xanthosoma. Although a new name was apparently erected,

there is no indication that the author was intentionally describing a new

species. It seems likely that this epithet would have been derived from

rather earlier usage by Schott, who worked under the patronage of

Archduke Ferdinand Maximilian of Austria, and did indeed name an East

Brazilian aroid Xanthosoma maximiliani Schott [Bonplandia 10 (1862)

322]. It would thus perhaps appear that Maedicke’s name was intended to

be a new combination. However, since no reference was made to any

earlier usage of the epithet, the name Alocasia maximiliani Maedicke must

in that case be regarded as entirely new and typified by Maedecke’s

illustration, not by the type of X. maximiliani Schott; it would need

epitypifying in the context of a revision of Xanthosoma. It seems more

plausible however, that use of the generic name Alocasia in this case was

simply a mistake rather than an actual taxonomic judgement.

Alocasia merrillii Engl. & K. Krause

Alocasia merrillii Engl. & K. Krause, Bot. Jahrb. Syst. 54, Beibl. 118 (1916)

124 = Schismatoglottis plurivenia Alderw. Type: Philippines, Mindanao,

Zamboanga Distr., Merrill 8072 (B, holo).

Alocasia reversa N.E. Brown

Alocasia reversa N.E. Brown, Gard. Chron. 8 (2) (1890) 38; Hook.f. in

Curtis’ Bot. Mag. Ser. 3, 52 (1896) t. 7498; Usteri, Beitr. Ken. Philip. Veg.

(1905) 130; K. Krause & Engl., Pflanzenr. 71 (IV.23E) (1920) 95; Merr.

Enum. Philipp. Fl. Pl. 1 (1922) 185; Hay, Gard. Bull. Singapore 50 (1998)

279. - Type: Cult. Kew ex Hort. Sander, Apr 1890, N.E. Brown s.n. (K,

holo).

Since its first description, subsequent authors have not ascribed any

additional Philippine material to this species. Material collected from the

wild and matching A. reversa is restricted to limestone substrate in Sarawak,

38 Gard. Bull. Singapore 51 (1999)

Borneo (Hay, 1998). Its initial ascription to the Philippines appears to

have been in error.

Alocasia talihan Elmer

Alocasia talihan Elmer, Leafl. Philipp. Bot. 10 (1938) 3614, nom. inval.,

descr. angl.; based on a collection from Luzon, Sorsogon Province, Irosin

(Mt Bulusan), Aug 1916, A.D.E. Elmer 17381 (NY). = Xanthosoma

sagittifolia Schott.

Alocasia urdanetensis Elmer

Alocasia urdanetensis Elmer, Leafl. Philipp. Bot. 10 (1939) 3699, nom.

inval., descr. angl.; based on a collection from Mindanao, Agusan Prov.

Cabadbaran (Mt Urdaneta), Sep 1912, Elmer 13857 (not located, presumed

lost at PNH).

The description suggests some resemblance to A. sanderiana.

Misapplied Names

The occurrence of several species of Alocasia was recorded for the

Philippines in Naves’ Appendix to the third edition of Blanco’s Flora de

Filipinas. Seven of these are not Philippine species. Since there is no

pertinent preserved material, it is not possible to interpret with any

confidence what Naves meant in applying these names.

Alocasia alba sensu auct. non Schott: Naves, Novis. App. (1882) 293.

Alocasia cucullata sensu auct. non (Lour.) G. Don: Naves, loc. cit. (but

possibly cultivated).

Alocasia longiloba sensu auct. non Miq.: Naves, op. cit., 294.

Alocasia lowii sensu auct. non Hook.f.: Naves, loc. cit.

Alocasia montana sensu auct. non (Roxb.) Schott: Naves, op. cit, 293.

Alocasia odora sensu auct. non (Lodd.) Spach: Naves, op. cit., 294.

Alocasia pubera sensu auct. non (Hassk.) Schott: Naves, op. cit., 293.

Alocasia in the Philippines 39

Acknowledgements

I am grateful to the Smithsonian Institution for awarding me a Visiting

Fellowship in 1996 and for providing me facilities to work in the Department

of Botany, US National Museum of Natural History and to Dr D.H.

Nicolson (US) for making that possible and for his encouragement, expert

advice and comment on the manuscript. I am also grateful to Peter Boyce

(K) for finding articles including protologues of Alocasia from European

horticultural journals and to Josef Bogner (Munich Botanic Garden) for

sending me preserved material and photographs of material he has

cultivated, to Dr Tom Croat (MO) for drawing my attention to informative

vouchers from the Missouri Botanical Garden living collection and to Clare

Herscovitch (NSW) for technical assistance. Special thanks are due to Lesley

Elkan (NSW) for the botanical drawings, which, due to circumstances

beyond our control, have had to be reproduced from photocopies. I thank

the curators and directors of the following herbaria for allowing me to see

Goulecions at BBM, BO, CAL. E.G, GH, K, L, M, MO. NSW, NY, P,

PNH, PR, UC, US.

Note: In Hay (1998) this paper was cited as ‘in press’ with Nordic Journal

of Botany. It has since transpired that the corrected and final version of

the paper and the original illustrations had become lost. In order to expedite

publication, the paper was withdrawn from that journal. No suggestion of

fault is directed at the editors of Nordic Journal of Botany. I thank Dr I.

Nielsen for his helpful comments on an earlier version of the manuscript,

and the editor-in-chief, Prof. Kai Larsen, for his forbearance.

References

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3635.

Elmer, A.D.E. 1939. Miscellaneous new species. Leaflets Philippine Botany.

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Engler, A. 1879. Araceae. In: A. & C. de Candolle (eds), Monographiae

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Engler, A & K. Krause, 1920. Araceae-Colocasioideae. In: A. Engler (ed.),

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Hay, A. 1994. Alocasia simonsiana - a new species of Araceae from New

Guinea. Blumea. 38: 331-333.

Hay, A. 1998. The genus Alocasia (Araceae-Colocasieae) in West Malesia

and Sulawesi. Gardens’ Bulletin Singapore. 50: 221-334

Hay, A. , J. Bogner, P.C. Boyce, W.L.A. Hetterscheid, N. Jacobsen & J.

Murata. 1995. Checklist and Botanical Bibliography of the Aroids of

Malesia, Australia and the Tropical Western Pacific Region. Blumea,

Supplement 8. - Rijksherbarium/Hortus Botanicus, Leiden University.

Hay, A. & Wise, R. 1991. The genus Alocasia (Araceae) in Australasia.

Blumea. 35: 499-545.

Merrill, E.D. 1912. A Flora of Manila. Bureau of Printing, Manila.

Merrill, E.D. 1918. Species Blancoanae. Bureau of Printing, Manila.

Merrill, E.D. 1922. An Enumeration of Philippine Flowering Plants. Vol. 1.

Bureau of Printing, Manila.

Nicolson, D.H. 1979. Araceae. In A.C. Smith (ed.), Flora Vitiensis Nova. 1:

438-460. Pacific Tropical Botanic Garden, Lauai.

Nicolson, D.H. 1987. Araceae. In M.D. Dassanayake & F.R. Fosberg (eds.),

Flora of Ceylon. 6: 17-101. Amerind, New Dehli.

Schott, H.W. 1860. Prodromus Systematis Aroidearum. Mechitarists’ Press,

Vienna.

Stafleu, F.A. & Cowan, R.S. 1979 Taxonomic Literature. 2: H-Le. Ed. 2.

[Regnum Vegetabile Vol. 98.] Junk, The Hague.

Steenis-Kruseman, M.J. van, 1950. Malaysian plant collectors and

collections. In: C.G.G.J. van Steenis (ed.), Flora Malesiana I. 1: 3-639.

Alocasia in the Philippines 4]

Index to Species

ee ANN MRE TRAN cee My esata chase ncaa vo dae a i deitvnsan tsa nonaesevesciinesvuasicweotoreconssics 3

BRE Ree Ee es ahs foc a aoa Sasa iecbeicewdeencavnescenaienaseie i

Saat aC Byars at ect I ekg 2 eed oahbe Pde digs ok tara Ssncsnesessowiesesscncvvaccuoses 9

ee Ne Ne Ih eailen ee die cdl od od snsepadanactesecncbshs-uceonavorchenianecss a2

SRN te age ede paisa lncur nny dalbwanesavodiessuccsansaaneneas 14

RI PORN le tls eed dials hdl av sednads pdnlisdcennveuteSigdasdoriuciiessssccearaaeehoees 15

OR lar MMs CUR etre 2a LE thas, Ea ee pees ds dala hu Sadiedacsveceotesancense 19

nN TIINIE MUM EASLa Dee he She ik shied sacasmibavensateriverades ae.

an cua Nia ORR shade ND aM, Rec AlN ESNet EK ARIK DN peavesecccesosentsoet 24

ae ae eee ease hats A Misra Ze cup user gncdoninédaatiattbuncnde lite wsedandases ett 25

NPN Na hs Ss IaH PNG dspam tak asupvosedateck secssinghadilusastidevcttcnensacb ease, ZA

Re RNa Neer hoe ac et bhp cee sa daha cusnindbnidevedsvdendssucseen> 29

SNES PERE AES A LEN Soak ered eases cph uve rnc aoaseevecuwdeneyinchrapetedndncusavesens 30

UN SEITE rete anh) 0 Sasi taas ab teaee se lonssede ba ttaatSenc.taslenddcodsesiansMicbesaldosddvaaterss a

een ee eo re ee ey eee or eee tA en es 34

Gardens’ Bulletin Singapore 51 (1999) 43-54.

Beetle Pollination of Vatica parvifolia

(Dipterocarpaceae) in Sarawak, Malaysia

TERUYOSHI NAGAMITSU

Hokkaido Research Centre, Forestry and Forest Product Research Institute

Hitsujigaoka 7, Toyohira, Sapporo 062-8516, Japan

RHETT D. HARRISON AND TAMIJI INOUE

Center for Ecological Research, Kyoto University,

Otsu 520-0105, Japan

Abstract

Pollination of a canopy tree of two species, Vatica aff. parvifolia and V. micrantha

(Dipterocarpaceae) was investigated in Sarawak, Malaysia. Flowers of the two species

open in the evening, and last for + 2 days. Rewards are petal tissue and pollen. In V. aff.

parvifolia, pollen was removed from anthers on the first day, and deposited on stigmas on

the second day, which suggests that the flowers are protandrous. Most (76%) flower visitors

were beetles (Chrysomelidae), 25% of which carried pollen. The beetles mated inside

flowers and often touched anthers and stigmas. These suggest that the beetles are pollinators.

Pollination of V. micrantha rarely occurred, because few pollen grains were removed from

anthers and deposited on stigmas. Dominant (71%) flower visitors were weevils (Apionidae

and Curculionidae), 9% of which carried pollen. The weevils laid eggs from outside flowers.

Bagging treatment increased fruit set in V. micrantha. This suggests the negative effects,

such as seed predation, of the weevils on fruit set.

Introduction

Subfamily Dipterocarpoidae (Dipterocarpaceae) in Asia comprises 13

genera and 470 species, classified into two tribes, the Dipterocarpae and

Shoreae, based on phenotypic characters (Ashton, 1982). Pollinators of

the Shoreae include various groups of insects, which differ in their ability

to transport pollen: thrips (Appanah & Chan, 1981), beetles (Sakai et al.,

1999), and bees (Appanah, 1990; Momose et al., 1996).

In the Dipterocarpae, pollinators known to date are butterflies, moths,

and bees, all of which are capable of transporting pollen long distances.

Honey bees have been reported to be the most effective pollinators of the

self-incompatible Vateria copallifera in Sri Lanka (Dayanandan et al., 1990).

Stingless bees visit flowers of Stemonoporus oblongifolius in Sri Lanka

(Ashton, 1988), which has a high (84%) outcrossing rate with potential of

apomixis (Murawski & Bawa, 1994). Butterflies and moths pollinate nectar-

44 Gard. Bull. Singapore 51 (1999)

secreting flowers of Dipterocarpus obtusifolius in Thailand (Ghazoul, 1997).

In Malaysia, geometrid moths pollinate D. pachyphyllus, and honey bees

pollinate D. geniculatus (Momose et al., 1998). However, pollinators of

Vatica, the largest genus (65 species) in tribe Dipterocarpae, were previously

unknown. Appanah (1987) predicted that pollinators of Vatica are tiny

flower-feeding insects, such as thrips, beetles, bugs, and hoppers, according

to floral traits of Vatica, such as nocturnal anthesis, poorly developed

anthers, and no nectar production.

In this study, we observed abundance and behaviour of flower visitors

of two species of Vatica, and examined the pollination process and the

effect of exclusion of flower visitors on fruit set, in order to determine

potential pollinators of Vatica.

Materials and Methods

The field research was conducted in the Lambir Hills National Park,

Sarawak, Malaysia (4°20'N, 113°50'E, 150-200 m elevation). Vatica parvifolia

Ashton is distributed in Sarawak and Burnei, and is rare on sandy ridges

below 600 m elevation (Ashton, 1982). One tree, which flowered in

September 1994, was investigated from the Operation Raleigh Tower in

the park, 25 m height above ground. This particular tree had leaves with

more distinct reticulation than V. parvifolia, so it is referred here as V. aff.

parvifolia (voucher: K. Momose 70, KYO and SAR; Nagamasu & Momose,

1997). Vatica micrantha Slooten (voucher: K. Momose 54, KYO and SAR)

in Borneo is widespread in mixed dipterocarp forest below 600 m elevation

(Ashton, 1982). One tree, which flowered in October 1994, was investigated

from the first aerial walkway in the Canopy Biology Plot at 20 m height

above ground (Inoue et al., 1995).

Flower visitors

Flower visitors of V. aff. parvifolia were observed and collected at 11.00h,

16.00h, and 21.00h on 13 September, and at 6.00h, 8.00h, 10.00h, 14.00h,

and 17.00h on 14 September, and those of V. micrantha at 9.00h, 13.00h,

15.00h, 17.00h, and 20.00h on 17 October, and at 6.00h on 18 October. We

collected flower visitors for 10 min using hand nets. At first, we caught

visitors flying around and coming to flowers, and later we swept visitors

staying on the flowers into the nets. These insects were identified to family

and superfamily levels. Pollen grains on their bodies were observed using

both light and scanning electron microscopes.

Pollination of Vatica 45

Pollination process

In order to describe the pollination process, flowers of V. aff. parvifolia

and V. micrantha were observed and collected after the collection of flower

visitors. An additional collection of V. micrantha flowers was made at

11.00h on 17 October.

Some insects visiting flowers left feeding marks on the petals. Feeding

marks were graded into four classes: Grade 0 = no feeding marks; Grade 1

= some petals with feeding marks; Grade 2 = all petals with feeding marks,

but the feeding area of each petal less than half the petal area; Grade 3 =

all petals with feeding marks, and at least one petal with more than half

the area eaten.

Pollen grains were extracted from anthers of single flowers in 3N

sodium hydroxide at 70°C, and dispersed in 1 ml of 20% ethanol. Pollen

grains in 0.0032 ml sampled from the | ml ethanol solution were counted

using a haematocytometer. Pollen grains on single stigmas were counted

using a light microscope. Pollen tubes in single styles were stained with

aniline-blue and counted using a fluorescine microscope (Alexander, 1987).

Fruit set

Effects of both exclusion of flower visitors and artificial self pollination on

fruit set of V. micrantha were examined by a pollination experiment. (We

did not conduct this experiment in V. aff. parvifolia, because the number

of the manipulated flowers was not enough). Branches with flower buds

were enclosed with fine mesh tetron bags (TORAY #9000). The number

of both flower buds and flowers on the bagged and unbagged branches

was counted on 10 October 1994 when flowering began. Artificial pollination

with pollen of the same tree was conducted for flowers in some of the bags

on 17 October. All bags were removed after flowering. To compare fruit

sets among 1) control (neither bagging nor pollination treatment), 2) bagging

treatment (no pollination treatment), 3) bagging and self-pollination

treatment, fruits on the monitored branches were counted on 30 November

and 23 December in 1994, and on 5 February in 1995.

Results

Flowers of Vatica aff. parvifolia (Fig. 1) have creamy-white petals, whereas

those of V. micrantha have yellowish-white petals, outside suffused with

red toward the base. In the two species, petals open from 16.00h to 21.00h.

Anther dehiscence occurs before the petals open. Petals and anthers begin

to fall two days after flower anthesis, depending on weather conditions.

46 Gard. Bull. Singapore 51 (1999)

Figure 1: a) A flowering tree of Vatica aff. parvifolia; b) A chrysomelid beetle, Oomorphus

sp. (Lamprosomatinae), collected from a V. aff. parvifolia flower. Pollen grains are found on

its ventral thorax.

Pollination of Vatica A7

Fresh and one-day-old flowers can be distinguished by the degree of petal

opening and connectivity of filaments to petals. No liquid reward was

observed in flowers of either species. Pollen and petal tissue are the rewards

for flower visitors.

Flower visitors

Most flower visitors collected from V. aff. parvifolia were beetles

(Chrysomelidae dominated by Oomorphus spp.; 76%, n = 179; Fig. 2a),

while those from V. micrantha were weevils (Apionidae and Curculionidae;

71%, n = 461; Fig. 2b). The composition of the flower visitors of the three

most abundant families differed between Vatica species (x° test, P < 0.001).

The composition of beetles and weevils temporally changed in V. micrantha,

but was stable in V. aff. parvifolia (x° test, P < 0.001 and P = 0.539,

respectively). Chrysomelid beetles on V. micrantha flowers were more

abundant at night and in the morning than in the afternoon (Fig. 2b).

Other visitors include species of Scarabaeidae and Cucujidae (Coleoptera);

Cicadellidae, Fulgolidae, Cimicidae, and Plataspidae (Hemiptera);

Blaconidae, Chalicidoidea, Trigonaloidea, Formicidae, and Vespidae

(Hymenopreta); Culicidae (Diptera); Blattellidae (Blattodea); and

Acromantidae (Mantodea). In addition to these insects, thrips

(Thysanoptera) were found in 10.2% (n = 177) and 1.4% (n = 140) of the

collected flowers of V. aff. parvifolia and V. micrantha, respectively.

On the bodies of these insects collected from the both Vatica species,

pollen grains were observed on 25% of beetles (Chrysomelidae, n = 103;

Fig. 1b), and 9% of weevils (Apionidae and Curculionidae, n = 142). No

pollen grains were found on the bodies of the other insects (n = 7). There

was no significant difference in the proportion of individuals with pollen

grains between beetles and weevils (Fisher’s exact probability test, P =

0.790 in V. aff. parvifolia and P = 0.127 in V. micrantha), and between

Vatica species (P = 0.259 in weevils and P = 0.052 in beetles). Female

chrysomelid beetles were observed feeding on both petals and pollen, and

staying inside the base of petal cups. On the other hand, male beetles were

observed feeding on petals, and waiting for females on the apical part of

the petals, where they mated. We found that apionid and curculionid weevils

laid eggs from outside the flowers into the petals and ovaries. The weevils

were observed feeding on both petals and pollen, but their mating was

rarely observed.

Pollination process

Feeding marks on petals of both V. aff. parvifolia and V. micrantha increased

with time after flower anthesis (Fig. 3a and 3b). Feeding marks in fresh

flowers of V. aff. parvifolia increased between 6.00h and 8.00h (Kraskal-

48 Gard. Bull. Singapore 51 (1999)

Wallis test, P = 0.039). One-day-old flowers had more feeding marks than

fresh flowers in both V. aff. parvifolia and V. micrantha (Mann-Whitney

test, P = 0.002 and P = 0.003, respectively).

The number of pollen grains in anthers of V. aff. parvifolia was

higher in fresh flowers than in one-day-old flowers, but that of V. micrantha

did not differ (Mann-Whitney test, P < 0.001 and P = 0.850, respectively;

Fig. 3c and 3d). The number of pollen grains in anthers of fresh flowers of

V. aff. parvifolia decreased between 8.00h to 14.00h (Kraskal-Wallis test, P

= 0.002; Fis: -3c):

The number of pollen grains on stigmas of V. aff. parvifolia was

higher in one-day-old flowers than in fresh flowers, but that of V. micrantha

did not differ (Mann-Whitney test, P < 0.001 and P = 0.121, respectively;

Fig. 3e and 3f). The number of pollen grains on stigmas of one-day-old

flowers of V. aff. parvifolia increased throughout the day (Kraskal-Wallis

a) Vatica aff. parvifolia \b) Vatica micrantha

Proportion (%)

VE 1G 28, GB) AO PA AR) Bite pi Rae Aan nT ee ee ee

13 Sep 14 Sep 17 Oct 18 Oct |

Time (date, hour)

Figure 2: Temporal changes in the proportion of four taxa of flower visitors (Chrysomelidae,

Apionidae, Curculionidae, and other taxa) collected from a) Vatica aff. parvifolia and b) V.

micrantha.

Pollination of Vatica 49

Vatica aff. parvifolia Vatica micrantha

Grade of

feeding mark

—_ —

No. pollen grains

in anthers (x103)

ie)

No. pollen grains

on a stigma

No. pollen tubes

ina style

Mather 6 8 1014 17 oo ae) 4 AB tz 20'S

13 Sep 14 Sep 17 Oct 18 Oct

Time (date, hour)

Figure 3: Temporal changes in the grade of feeding marks on petals (a-b), the number of

pollen grains in anthers (c-d), the number of pollen grains on a stigma (e-f), the number of

pollen tubes in a style (g-h), of flowers of Vatica aff. parvifolia (a,c, e, and g) and V. micrantha

(b, c, f, and h). Circles and squares show fresh and one-day-old flowers, respectively.

50 Gard. Bull. Singapore 51 (1999)

test, P = 0.014; Fig. 3e). Results of observations on pollen tubes in styles

were similar to those of pollen grains on stigmas (Fig. 3g and 3f).

Fruit set

We monitored 568 flower buds and flowers in control, 89 in bagging

treatment, and 90 in bagging and self-pollination treatment. Fruit set was

observed 50 and 72 days after flowering. In the bagging treatment (27.0%

and 20.2%) as well as the bagging and self-pollination treatment (15.6%

and 11.1%) fruit set was higher than that in the control (2.5% and 1.9%,

respectively; Fisher’s exact probability test, P < 0.001; Fig. 4). However,

fruit set in the bagging and self-pollination treatment did not differ from

that in the bagging treatment in the two periods (P > 0.154). Fruit set

observed 117 days after flowering was 0.35% in the control, 2.25% in the

bagging treatment, and 2.22% in bagging and self-pollination treatment.

Discussion

Pollination in Vatica aff. parvifolia

Pollination of V. aff. parvifolia was successful, because most of the pollen

grains were removed from anthers and on average about 20 pollen grains

were deposited on a single stigma, which are sufficient to fertilize one of

the six ovules per flower, from which one seed develops (Swarupanandan,

1986). |

Beetles (Chrysomelidae) are likely pollinators in V. aff. parvifolia,

based on the observations that 1) beetles were the most abundant flower

visitors, 2) only beetles and weevils (Apionidae and Curculionidae) were

observed carrying pollen grains on their bodies, and 3) according to their

behaviour observed at flowers, beetles had a greater chance of touching

anthers and stigmas than weevils, although the proportions of individuals

carrying pollen did not differ between beetles and weevils. However, further

studies are necessary to confirm beetle pollination in V. aff. parvifolia,

because only single tree was investigated in this study.

Fruit set in Vatica micrantha

Pollination of V. micrantha rarely occurred in this study, because only a

few pollen grains were removed from the anthers and deposited on stigmas.

The most abundant flower visitors were weevils, which appear to be seed

predators based on their behaviour observed at flowers.

The result of the pollination experiment showed that bagging

treatment increased fruit set. This unusual result suggests at least two

causes: 1) negative effects of flower visitors on fruit set, and 2) pollination

Pollination of Vatica a

mechanisms in the bagged flowers. One of the negative effects of flower

visitors is seed predation. Weevils, the dominant flower visitors, are common

seed predators of dipterocarps (Daljeet-Singh, 1974; Toy, 1991). Thus, it is

plausible that the bagging treatment prevented the weevils from predating

the seeds. Pollination in the bagged flowers might be achieved by thrips or

through autogamy, otherwise apomixis could occur. There is a possibility

that thrips entered and pollinated the bagged flowers (Appanah & Chan,

1981), because neither insecticide before bagging nor glue at the tied mouths

of the bags were used. Autogamy and apomixis have been shown in some

other dipterocarps (Chan, 1981; Kaur et al., 1986; Dayanandan et al., 1990;

Murawski & Bawa, 1994).

100

co 10

oN.

dp)

by ed

mB |

LL 1

0.1

0 20 40 60 80 100 S620

Days after flowering

Figure 4: Temporal changes in fruit set of Vatica micrantha. Fruit set is compared among

control (open circles), bagging treatment (closed circles), and bagging and self-pollination

treatment (closed squares). Different letters show significant differences (P < 0.001).

52 Gard. Bull. Singapore 51 (1999)

Beetle pollination syndromes in dipterocarps

Appanah (1987) predicted that Vatica flowers could be pollinated by tiny

flower-feeding insects, such as thrips, beetles, bugs, and hoppers. The

present study suggests that the potential pollinators of Vatica aff. parvifolia

are chrysomelid beetles, which supports the prediction of Appanah (1987).

Chrysomelid beetle pollination in subfamily Dipterocarpoidae has ~

also been demonstrated in Shorea parvifolia (Sakai et al., 1999), although

there are differences in pollinator behaviour and phenological traits of

flowers between Shorea and Vatica. Pollinators of S. parvifolia are

dominated by Monolepta spp. (Galerucinae), which visited flowers at night,

while those of V. aff. parvifolia and dominated by Oomorphus spp.

(Lamprosomatinae), which visited flowers throughout the day, and left

feeding marks on petals in the morning. Pollen removal from anthers of V.

aff. parvifolia was also observed in the morning. Pollen deposition on

stigmas was observed at night in S. parvifolia, but throughout the day in V.

aff. parvifolia. These lines of evidence suggest that beetle pollinators of S.

parvifolia exhibited clear nocturnal activity, whereas those of V. aff.

parvifolia seemed most active in the morning.

In accordance with the difference in daily activity of pollinators,

floral phenology differed between S. parvifolia and V. aff. parvifolia (Sakai

et al., 1999). The lifetime of most flowers is one day in the former, and two

days in the latter, even though flowers of the both species open in the

evening. Pollination of flowers of S. parvifolia occurs at the first night after

flowering, while flowers of V. aff. parvifolia seem protandrous with pollen

removal on the first day and pollen deposition on the second day, although

lack of stigma receptivity in fresh flowers has not been confirmed. These

two types of temporal matching between pollinator activity and floral

phenology observed in Shorea and Vatica suggest divergent syndromes in

chrysomelid beetle pollination in dipterocarps.

Acknowledgements

We would like to thank Dr. H. S. Lee and Mr. A. A. Hamid (Forest

Department Sarawak), and Dr. K. Ogino, (University of Shiga Prefecture)

for supporting and organizing this study; and Dr. P. S. Ashton (Harvard

University) for improving the early version of the manuscript. This study

was supported by Grants-in-Aid from the Japan Ministry of Education,

Science, and Culture (grant numbers 04041067, 06041013, and 09N1501)

and by JSTC Domestic Research Fellowship for TN.

Pollination of Vatica 53

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The Taxonomic Significance of Trichome Morphology in

the Genus Durio (Bombacaceae)

I. SALMA

Malaysian Agricultural Research and Development Institute, Kuala Lumpur,

Malaysia

Abstract

Scanning electron microscopy was used to examine the trichome morphology on the leaves

of 24 species of Durio. Glandular trichomes and non-glandular trichomes are present on

the abaxial leaf surface. Stellate hairs can be found on the adaxial leaf surface of D. affinis,

D. graveolens, D. oblongus, D. oxleyanus, D. singaporensis and D. zibethinus. Trichome

morphology can be used to distinguish the Durio species studied. D. carinatus can be

distinguished by the presence of only peltate scales on the abaxial leaf surface. The other

species can be divided into three major groups based on the density and distribution of the

peltate scales and stellate hairs.

Introduction

The genus Durio consists of 29 species, which are confined to tropical

Southeast Asia (Kostermans, 1958) and distributed from Sri Lanka, India

and Burma through Thailand, Peninsular Malaysia, Singapore, Borneo,

Sumatra, the Philippines to New Guinea.

Earlier taxonomic accounts of the genus have been evaluated and

revised based on their macromorphological characters (Beccari, 1889; King,

1891; Bakhuizen, 1924; Ridley, 1922; Corner, 1939; Wyatt-Smith, 1953;

Kostermans, 1958; Kochummen, 1972; Cockburn, 1976 and Ashton, 1988).

In Durio species, leaf morphology such as shape and size is variable even

within a single species making it difficult to distinguish the species based

on leaf characters alone (Kochummen, 1972; 1988). However, identification

of Durio species based on leaf characters is important since the flowering

and fruiting episodes are seasonal and for most of the year only sterile

specimens are available.

Micromorphological characters of the leaf epidermis are increasingly

being employed as a source of important diagnostic characters in taxonomic

determination (Metcalfe and Chalk, 1950; Stace, 1965, 1984). Baas (1972)

studied the epidermal cells of Kostermansia malayana. Durio species exhibit

a diversity of trichome types. All species bear several types of trichomes

on leaves, stem and flowers. Nor Azian (1992) who studied leaf epidermis

of eight Durio species revealed some diagnostic characters of trichomes

56 Gard. Bull. Singapore 51 (1999)

for their identification. The aim of the present study is to investigate the

microscopic variability of trichome characters of the leaf surface and to

assess their potential as taxonomic markers for Durio species.

Materials and Methods

Leaf samples from living plants as well as from herbarium materials of

only 24 Durio species were available and used for this investigation (Table

1). A5 mm section of dry leaf lamina (adaxial and abaxial surfaces) was

fixed onto double-sided adhesive tape on labelled stubs. The specimens

were coated with gold paladium at 260° A, and scanned under JOEL JSM-

35C scanning electron microscope at the Universiti Putra Malaysia.

A square of cellophane tape about | cm_ was pressed on the abaxial

leaf surface. It was then pulled free and pasted onto a glass slide. The

morphological features of the scales were observed under a light microscope.

Light microscope photographs were then taken.

Transverse sections of lamina were also made. Fresh samples were

fixed immediately in 50% FAA (formalin, acetic acid, alcohol). Dried

herbarium materials were revived in water at 60 C overnight. Specimens

were then dehydrated, sectioned and stained with safranin O and fast

green using the standard method (Sass, 1958).

Table 1: List of Durio species examined

Species Herbarium specimen Fresh specimen

(Locality)

15) yD. ais KEP 3945

2. D.carinatus FRI 3819 Rompin

3. D2. crassipes SAN 16810

4. D. dulcis SAN 15389

5.’ Drexcelsus A T3513

6a. D. grandiflorous SAN 4611

var. grandiflorus

6b. D. grandiflorus § 39199

var. tomentosus

7. D. graveolens KEP 65525 Jerangau

Trichomes in Durio

Species Herbarium specimen Fresh specimen

(Locality)

8a. D. griffithii var. KEP 7344 Semenyih

griffithii ;

8b. D. griffithii var. SAN 34716

acutifolius

9. D. johoricus MS 1674

10. D. kinabaluensis SAN 105554

11. D. kutejensis S 22848

12. D.lanceolatus SAN 15373

13. D.lowianus Serdang

14. D. macrolepis Haniff 3700

15. D.macrophyllus KEP 2731

16. D.malaccensis KEP 64878 Semenyih

17. D. oblongus S 43701

18. D. oxleyanus FRI 21576 Jerangau

19. D. pinangianus KEP 3780

20a. D. singaporensis Jerangau

var. singaporensis

20b. D. singaporensi MAR 6529

var. jerangauensis

21a. D. testudinarum A 3101

var. testudinarum

21b. D. testudinarum SAN 26217

var. crassifolius

22. D. perakensis KEP 28714

23. D.wyatt-smithii KEP 57451 Bt. Bauk, Dungun

24. D. zibethinus

Serdang

58 Gard. Bull. Singapore 51 (1999)

Results

The differences in shape, density and distribution of the trichomes are

illustrated in Figures la-k; Plates la-f and 2a-r. Data on trichome characters

are tabulated in Table 2. Two basic types of trichomes are found on the

leaf of Durio species: (a) glandular and (b) non-glandular .

Figure 1. Types of trichomes. (a) Two-celled stalk with unicellular hemispherical glandular

head; (b) Two-celled stalk with multicellular hemispherical glandular head; (c) One- to

two-celled stalk with ovoid multicellular glandular head; (d) Uniseriate stalk with single-celled

globular or hemispherical head; (e) Foot of peltate scale; (f) Complex peltate scale; (g) Simple

peltate scale (h) Cushioned stellate hair; (i) Flat stellate hair; (j) Four-armed stellate trichome

with a central cushion; (k) Simple non-glandular trichome

Trichomes in Durio 59

(a) Glandular trichomes

Glandular trichomes are found on the abaxial leaf surface of all Durio

species and these can be divided into three categories. The first type is the

glandular trichome with a one- or two- celled stalk and a hemispherical

glandular head, which is unicellular or multicellular. In transverse section

the short one-celled stalk has its foot embedded in the epidermal layer

(Figs.la-b) and the trichome base is supported by unmodified epidermal

cells (Fig. le). This type of glandular trichomes occurs abundantly on the

Plate 1. SEM micrographs of Durio trichomes showing their types and arrangement on the

abaxial leaf surface. (a) Complex peltate scale; (b) Simple peltate scale; (c) Cushioned stellate

hairs; (d) Flat stellate hairs; (e) Four-armed stellate trichomes with a central cushion; (f)

Arrangement of trichomes on the abaxial leaf surface. Scale bar = 100um.

EVE UPI ee RL ae ee ee | SS eRe ee |

60 Gard. Bull. Singapore 51 (1999)

abaxial leaf surface of all species.

The second type is the short-stalked trichome with a multicellular

glandular head. This type of trichome has a one- or two-celled stalk and an

ovoid multicellular glandular head (Fig.1c). The base of the trichome is

thickened with unmodified epidermal cells. This glandular trichome occurs

in D. affinis and D. carinatus.

The third type is the long-stalked trichome with a hemispherical

head consisting of a 2- to 9-celled uniseriate stalk terminating in a single-

celled globular or hemispherical head (Fig. 1d). The foot may arise from

the centre of the epidermal cell. The base of the trichome is thickened

with unmodified epidermal cells. This type of glandular trichome is observed

on the abaxial leaf surface of D. dulcis, D. testudinarum and D.

singaporensis.

(b) Non-glandular trichomes

Non-glandular trichomes are observed on the abaxial leaf surface of all

species and on the adaxial leaf surface of some species (Table 2; Plates 1 &

2). Six types of non-glandular trichomes are recognised:

(1) Complex peltate scales - sessile, foot uniseriate embedded in the

epidermis, uniseriate, multicellular head delimited by primary and secondary

radial walls, margin undulating, 3—5-lobed, or sometimes split, short- or

long-fimbriated (Fig. 1f and Plate la). The trichome base is generally

multicellular with polygonal, isodiametric, straight-walled cells (Fig. le).

This type of scale occurs on the abaxial leaf epidermis of all species.

(11) Simple peltate scales - sessile, foot uniseriate sunken in the

epidermis, multicellular head delimited only by primary radial walls, margin

strongly lobed, not split, long-fimbriated (Fig. 1g and Plate 1b), occurs in

all species.

(i111) Cushioned stellate hairs - sessile, rays more than five with a

central cushion (Fig. 1h and Plate 1c); occurs in all species except in D.

carinatus.

(iv) Flat stellate hairs - stalked or sessile, rays more than five, without

a central cushion (Fig. 1i and Plate 1d); occurs in all species except in D.

carinatus.

(v) Four-armed stellate trichome - sessile with a central cushion

(Fig. 1] and Plate le); occurs in all species except in D. carinatus

(vi) Simple - unicellular, long, tapered at the tip (Fig. 1k); occurs in

D. lowianus and D. singaporensis var. jerangauensis.

In most species (except D. oxleyanus, D. excelsus, D. perakensis, D.

griffithii var. griffithii and D. grandiflorus var. tomentosus), the two types

of peltate scales form three to four layers while the stellate hairs, the four-

Trichomes in Durio 61

armed trichomes and all the glandular trichomes lie beneath them (Table

2; Plates 1f and 2a-r). In D. oxleyanus, D. excelsus, D. perakensis, D.

griffithii var. griffithii and D. grandiflorus var. tomentosus, the abaxial leaf

surface is densely covered with the two types of stellate hairs and four-

armed trichomes with peltate scales scattered sparsely above them (Plate

2 e’and 1):

In some species, infraspecific variation of trichome characters is

observed. There is variation in the density of the type of trichomes on the

abaxial leaf surface of D. griffithii var. griffithii and D. griffithii var.

acutifolius. There are dense stellate hairs indicated by (+++) with very few

peltate scales (+) on the abaxial leaf surface of D. griffithii var. griffithii

while in D. griffithii var. acutifolius there are dense peltate scales (+++)

with dense stellate hairs (+++) underneath (Table 2). Similarly, in D.

grandiflorus var. grandiflorus the abaxial leaf surface possesses dense peltate

scales (+++) with dense stellate hairs (+++) underneath, while in D.

grandiflorus var. tomentosus there are dense stellate hairs (+++) with a few

peltate scales (+) scattered on top. D. singaporensis var. jerangauensis differs

from D. singaporensis var. singaporensis by having simple unicellular

trichomes on the abaxial leaf surface (Table 2). Whereas D. testudinarum

var. testudinarum can be distinguished from D. testudinarum var. crassifolius

by having a few long-stalked glandular trichomes with hemispherical head.

The shape of the outermost layer of the peltate scales varies between

species. Thus, in D. malaccensis the peltate scales are almost round or

have a slightly undulating margin, which is usually not split and 1s shortly

fimbriated (Plate 2j), while in D. macrophyllus the peltate scales are deeply

five-lobed and split and are distinctly fimbriated (Plate 2k). D. pinangianus

possesses similar trichome characters as D. malaccensis except that its

complex peltate scales are slightly lobed. However other leaf parameters

such as leaf size, texture and veination patterns are different.

In most species the complex peltate scales are split except in D.

malaccensis, D. oxleyanus, D. testudinarum var. crassifolius, D. grandiflorus

var. grandiflorus, D. griffithii var. acutifolius and D. johoricus where the

complex scales are mostly entire (Table 2).

The size of the outermost layer of complex peltate scales also varies

between species. D. graveolens, D. macrophyllus, D. oblongus and D.

singaporensis have large complex peltate scales with the mean diameter

greater than 800 mm (Table 2), whereas in D. excelsus, D. griffithii and D.

grandiflorus the scales are smaller and less than 200 mm in diameter.

The colour of the complex peltate scales on the abaxial surface of

both fresh and dried laminas is distinct in certain species. Thus in D.

graveolens, D. singaporensis, D. macrophyllus, D. oblongus and D.

kutejensis, the complex peltate scales are dark golden brown while in other

species the complex peltate scales are brown in colour.

Gard. Bull. Singapore 51 (1999)

F445

barn

Trichomes in Durio 63

Plate 2. SEM micrographs showing trichome shapes and their distribution on the abaxial

leaf surface of Durio species. (a) D. affinis (b) D. carinatus (c) D dulcis (d) D. grandiflorus

var. grandiflorus (e) D. griffithii var. griffithii (f) D. griffithii var. acutifolius (g) D. graveolens

(h) D. kutejensis (i) D. lowianus (j) D. malaccensis (k) D. macrophyllus (1) D. oxleyanus (m) D.

pinangianus (n) D. singaporensis (0) D. testudinarum (p) D. zibethinus; trichome shapes on

the adaxial leaf surface. (q) D. singaporensis and (r) D. graveolens. Scale bar= 100m.

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Gard. Bull. Singapore 51 (1999)

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Trichomes in Durio 67

In most species, the trichomes on the abaxial leaf surface are

appressed but in D. graveolens, D. macrophyllus and D. macrolepis the

trichomes are loosely attached, and the veins beneath them can clearly be

seen when the trichomes are removed.

On the adaxial leaf surface of D. affinis, D. graveolens, D. oblongus,

D. oxleyanus, D. singaporensis and D. zibethinus, the only trichomes present

are the sessile stellate hairs with more than five rays, and without a cushion

(Plates 2q and 2r).

Discussion and Conclusion

In general, the types of trichome present on the abaxial leaf surface of all

species are similar except for the presence of simple, non glandular

trichomes in D. lowianus, the long-stalked glandular trichome in D. dulcis,

D. kutejensis and D. singaporensis, and the multicellular-headed glandular

trichome in D. affinis and D. carinatus. Nor Azian (1992), who studied the

anatomy of eight Durio species, i.e. D. oxleyanus, D. zibethinus, D. lowianus,

D. kutejensis, D. malaccensis, D. graveolens, D. singaporensis and D.

griffithii, also observed the presence of glandular and non-glandular

trichomes on the abaxial leaf surface. However, she reported the presence

of only two types of glandular trichomes, unicellular and multicellular

uniseriate-stalked types. She did not report the presence of multicellular

glandular trichomes in D. griffithii nor the presence of simple trichomes in

D. lowianus. Density of the glandular trichomes is not a useful character to

distinguish the species.

Differences exist between the Durio species for density and

distribution of non-glandular trichomes. This was also noted by Nor Azian

(1992). Among trichome characters, the distribution and density of non-

glandular trichomes overall provide the most useful taxonomic character

in the identification of sterile Durio specimens. Based on the distribution

and density of the peltate scales and stellate hairs infraspecific variation

can be determined. D. griffithii var. acutifolius and D. grandiflorus var.

grandiflorus can be distinguished from D. griffithii var. griffithii and D.

grandiflorus var. tomentosus, respectively by having dense peltate scales

on the abaxial leaf surface. whereas very few complex peltate scales present

on the abaxial leaf surface of the latter two varieties. D. singaporensis var.

jerangauensis is differentiated from D. singaporensis var. singaporensis in

the presence of simple unicellular trichome.

The shape of the peltate scales in the outermost layer also provides a

useful character in distinguishing some species as they show differences in

shape and size. D. malaccensis is significantly different in having almost

68 Gard. Bull. Singapore 51 (1999)

circular and shortly fimbriated peltate scales while D. macrophyllus can be

recognised by its deeply lobed, 4—5-split, long-fimbriated peltate scales.

Although this character has not been used in the identification of the

species before, it can provide a good supplementary character in some

species particularly D. malaccensis and D. macrophyllus.

Another good trichome character is the mode of trichome attachment

to the surface of the leaf. The loose attachment of the trichomes in species,

such as D. graveolens, D. macrophyllus and D. macrolepis, also contributes

an important criterion for recognising these species.

In most species the colour of the peltate scales on the abaxial leaf

surface is brown. However, in certain species, such as D. graveolens and

D. macrophyllus, the dark coppery brown colour of the peltate scales

provides an important diagnostic character for their identification.

The presence of stellate hairs on the adaxial leaf surface is also an

important character, which can be used to distinguish some species, such

as D. affinis, D. graveolens, D. oblongus, D. oxleyanus, D. singaporensis

and individuals of D. zibethinus.

Based on density and distribution of trichomes on the adaxial leaf

epidermis, D. carinatus can be distinguished by having only peltate scales.

The other Durio species can be divided into three broad groups: (i) species

with a dense layer of stellate hairs with sparse peltate scales (such as D.

griffithii var. griffithii, D. grandiflorus var. tomentosus, D. oxleyanus, D.

perakensis and D. excelsus); (11) species with dense stellate hairs with peltate

scales moderately scattered above them, exposing stellate hairs beneath

(e.g. D. dulcis, D. graveolens, D. macrophyllus, D. macrolepis and some

individuals of D. zibethinus); and (iii) the remaining species which have

dense stellate hairs and dense peltate scales.

The use of a combination of all the trichome characters enables the

identification of most species as well as distinguishing spesies at infraspecific

level. Based on trichome characters of the abaxial leaf surface, species in

the first group one are distantly related to groups two and three, which are

closely related. The results of SEM study on the micromorphological

characters of the leaf surface support those obtained in the earlier study

based on macromorphological characters (Salma, 1996).

Leaf surface characters are under strong genetic control and are

little affected by the environment. Trichome structures are useful for the

delimitation of the Durio species. It can be concluded that these characters

provide significant systematic value for distinguishing Durio species.

Trichomes in Durio 69

Acknowledgements

I wish to thank Dr. Ruth Kiew and Dr. E. Soepadmo for their advice and

constructive criticisms on this paper. I also wish to thank the curators of

the herbaria of Kepong (KEP), Sandakan (SAN), Kuching (SAR) and

Singapore (SING) for the loan of specimens used in the present study. I

am also indebted to Universiti Putra Malaysia for allowing me to use the

Scanning Electron Microscope facilities. Thanks are also due to Mr. Tuan

Othman Tuan Abdullah, Mrs. Aminah and Mr. Ho Oi Kuan for their

technical assistance.

References

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ed. Sarawak: Dewan Bahasa Pustaka, Sarawak Branch. pp. 52-76.

Baas, P. 1972. The vegetative anatomy of Kostermansia malayana Soegeng.

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Jardine Botanique Buitenzorg Series II. 6: 162-163, 223-225.

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308.

Heywood V.H. 1984. The current scene in plant taxonomy. Jn: Heywood,

V.H. and Moore D.M. (eds.) Current Concepts in Plant Taxonomy.

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King, G. 1891. Materials for a Flora of the Malayan Peninsula. Journal

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Kochummen, K.M. 1972. Bombacaceae. Tree Flora of Malaya. 3: 100-116.

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Scientific Association Northern Branch and Universiti Sains Malaysia.

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Reinwardtia. 4: 47-150.

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Oxford: Clarendon Press. pp. 235-241.

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Clonal Variation in D. zibethinus. Ph.D. Thesis. Universiti Putra Malaysia.

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Stace, C.A. 1965. Cuticular characters as an aid to plant taxonomy. Bulletin

British Museum Natural History (Botany). 4: 1-78.

Stace, CA. 1984. The Taxonomic Importance of Leaf Surface. In: Heywood

V.H. and Moore D.M. (eds.) Current Concepts in Plant Taxonomy.

Academic Press. pp. 67 — 93.

Wyatt-Smith, J. 1953. Materials for the revision of Malayan Durio with

notes on Malayan species. Kew Bulletin.. pp. 513-532.

Gardens’ Bulletin Singapore 51 (1999) 71-84.

Foliicolous Ascomycetes 8: Vietnam!’

DON R. REYNOLDS

Natural History Museum of Los Angeles County

900 Exposition Boulevard, Los Angeles, California 90007, USA

Abstract

Records for Vietnam foliicolous ascomycetes (including the mitosporic species) are reviewed.

The current locations of previously cited specimens are noted. Recent collections from Ba

Vi and Cuc Phoung National Parks are annotated in the genera Appendiculella, Asteridiella,

Asterina, Atractilina, Hyalosphaeria, Leprieurina, Malacaria, Meliolaster, Polychaeton,

Sarcinella, Sirosporium and Trichothyrium. Seven of the twelve species are new records.

Introduction

Distinct lineages of foliicolous ascomycetes have evolved to live on plant

surfaces. They are distinguished by adaptive morphological attributes and

nutritional modes. These species constitute a guild found in close association

with vascular plants. The hypothesis underlying this report and related

work is that the diversity and distribution of this mycota reflects overall

biodiversity patterns of the associated flora.

The historical record for foliicolous fungi is rich in Southeast Asia.

The overall mycota of the Philippines and Indonesia is particularly well

known from mycological efforts beginning in the latter part of the 1800s.

Firsthand knowledge of foliicolous ascomycete fungi from field work over

a 35-year period by the author in Myanmar, Malaysia, the Philippines,

Singapore, Thailand as well as Taiwan and mainland China, has sustained

an impression of distinctive biodiversity patterns. Likewise, the historical

records from Vietnam suggest a distinct assemblage of foliicolous species.

An Overview of Vietnam Mycology

An inventory of Vietnamese fungi based on the literature comprises 365

species and 5 varieties in 143 genera. Vietnam type localities were cited for

46 species and 1 variety. A comparison of the Vietnam mycota with records

from the Philippines up to about 1905 (Teodoro, 1937) indicates a 45%

similarity.

‘Continued from: Reynolds, D.R. 1986. Foliicolous Ascomycetes 7: A teleomorphic phylogeny of the

Capnodiaceae. Mycotaxo. 27: 377-403.

72 Gard. Bull. Singapore 51 (1999)

The mycota of Vietnam was initially made known from collections

of J. B. Balansa and P. Bon and various others associated with a French

Scientific Mission in “Tonkin” and “Annan” at the turn-of-the century.

These collections were examined and published by Hennings (1895), von

Hohnel (1918), Karsten and Roumeguére (1890), Patouillard (1889, 1890,

1891, 1892, 1893, 1896, 1897, 1907, 1909), and Roumeguére (1886a, 1886b,

1890, 1892). Studies of mitosporic ascomycete fungi associated with plant

diseases were made by F. Bugnicourt (Mouchacca, 1997). Bui Xuan Dong

(1972, 1977, 1986) proposed a revision of Hyphomycete classification as a

doctoral thesis and subsequently undertook a survey of Vietnamese

Hyphomycetes, especially species of Aspergillus and Penicillium (Buti Xuan

Dong and Hoang Xuan Vinh, 1977). .

The locations of surviving specimens from Vietnam and adjacent

areas have been difficult to ascertain because of the destruction and

relocations that have occurred related to destructive world events in this

century and a subsequent lack of citation in recent literature. The difficulty

of obtaining specimen information is compounded by a dirth of accessible

fungal data, especially for type specimens. The alternative has been the

compilation of specimen information through personal visits to the major

herbaria of the world. These data are incorporated into this report.

Field Survey of Cuc Phoung and Ba Vi National Parks

The intent of recent collection efforts in Vietnam, the basis of this report,

was to resample the foliicolous mycota in order to gain better insight into

regional fungal biodiversity. This report is based on collections made in

December 1996 from the Cuc Phoung and Ba Vi National Parks in northern

Vietnam that have been designated as areas of high diversity worthy of

conservation (Khanh, 1994). Collections were made by the author

accompanied by Dr. Tran Ninh, Hanoi National University and Dr. Benito

Ching Tan, National University of Singapore.

One set of specimens is deposited in the University of Hanoi (HNU),

a second set in the Natural History Museum of Los Angeles County (LAM).

Specimens examined in this study are cited here after the species description.

For herbaria, the acronyms follow those in Index Herbariorum (Holmgren

et al., 1990). Specimens mentioned in historical reports are located in B,

FH (Pfister, 1977 for Patouillard’s type specimens), FLS, P, and S.

The collection sites

The World Wide Fund for Nature characterizes Vietnam as one of Asia’s

most biologically important countries (Anon., 1996). Ba Vi and Cuc Phuong

Foliicolous Ascomycetes in Vietnam 73

are two national parks cited by Khanh (1994) as major forest reserves of

Vietnam.

The vegetation of Cuc Phoung and Ba Vi National Parks was

described by Schmid (1989) and Nguyen Van Truong (1966) as Thailandian

monsoon forests. The 25,000-ha Cuc Phuong National Park is located about

100 km south of Hanoi in Ninh Binh, Hoa Binh and Thanh Hoa provinces,

20° 19’N, 105° 22’E. It comprises lowland and sub-montane seasonal

evergreen sub-tropical forest with lowland rain forest in the flatter parts of

the broad valley formed between two limestone ranges with elevations

from 150-637 m. The park flora is estimated at 2,000 flowering plant species.

The 2,140-ha nature preserve portion of Ba Vi National Park is

located about 50 km west of Hanoi in Ha Tay Province, 21° 25'N 105°

30°E. The three closed, moist forest types in this region are evergreen

tropical, evergreen sub-tropical, and broad-leaved and coniferous evergreen

sub-tropical forests. An “elfin forest” favouring the growth of bryophytes

is found in the high humidity, higher elevations of the sub-tropical areas.

The elevation in the park ranges from the lower boundary of the nature

preserve at 400 m to 1,270 m on Dihn Vua peak.

The sampling technique for foliicolous fungi utilized in this study

and the one apparent in the historical record is the haphazard, non-random

method that I call a “visual survey.” Collections were made by taking

leaves at ground level and within arm’s reach up to about 2.5 m that were

visibly colonized by fungi until there was an intuitive sense of diminished

return in a localized area.

Annotation of Recently Collected Foliicolous Species

1. Appendiculella Hohn., Stizb. K. Akad. Wiss. Wien, Math.-naturw. KI.

128 (1919) 556.

Ascocarp darkly pigmented, globose, with larviform appendages; ascus

globose, thin-walled; ascospores darkly pigmented, 4-septate; mycelium

darkly pigmented, superficial on living plant surfaces, hyphopodiate; without

setae.

Appendiculella arecibensis (F. L. Stevens) Toro, Mycologia 17 (1925) 144.

Ascocarp 194-213 um diam.; 4-8 erect larviform appendages brownish,

continuous, 113 x 26 um at base; ascospore ellipsoid, 4-septate, 44-48 x 20

um.

“N

74 Gard. Bull. Singapore 51 (1999)

Illustration: Hansford, 1963: Pl. LXX.

Specimen examined: Cuc Phoung National Park, Park Center, Don R.

Reynolds V325, 19 December 1996 (LAM).

This is a new record for Vietnam. The ascocarp and the setae are larger

and the ascospores are slightly smaller than those of the type specimen

from Puerto Rico, as well as other collections from Central America.

The other species reported from Vietnam, Appendiculella tonkinensis

(Karsten and Roumegueére) Toro, Mycologia 19 (1927) 71, was based on

Meliola tonkensis Karsten and Roumeguére, Rev. Mycol. 12 (1890) 77

described from B. Balansa 5944 (P) Thu Phap, distributed in Roumeguére

Fungi selecti exsiccati.

2. Asteridiella McAlpine, Proc. Linn. Soc. N.S.W. (1897) 38.

Ascocarp darkly pigmented, globose, glabrous; ascus globose, thin-walled;

ascospores darkly pigmented, 4-septate; mycelium superficial on living plant

surfaces, hyphopodate, without setae.

Asteridiella boni (Gaillard) Hansf.. Sydowia 10 (1962) 47.

Ascocarp to 230 um diam. with conoid surface cells measuring up to 20 um

high; ascospore ellipsoid, 3-septate, 60 x 20 um.

Illustration: Hansford, 1963: Pl. CCLXXXI.

Specimen examined: Ba Vi National Park, 1000 m. elevation, Don R.

Reynolds V301, 22 December 1996 (LAM).

The type specimen of A. boni differs from the other four species known

from Vietnam largely in the size of the ascospore, which measures 60 x 20

um and is most similar in morphological attributes to that known only

from a single Philippine collection.

Five species of Asteridiella are known from Vietnam. Asteridiella

boni (=Meliola boni Gail., Le Genre Meliola (1892) 39), was based on Bon

3319 (P) from Tonkin. Asteridiella duportii Hansf., Beih. Sydowia Ann.

Mycol., Ser. I. Beih. II. (1961) 239 was based on Duport 2 (FH) from

ChoGauh. The type specimen for A. gymnosporiae (Sydow) Hansf., Sydowia

10 (1957) 48, is PBS 7422 from the Philippines; this species is also known

from Bon 519] (FH) from Thinh Hoa Phu Dien (initially determined as

Meliola laevis Berk. and Curtis, J. Linn. Soc. London X (1869) 280). The

type specimen Balansa 25 of Asteridiella reticulata (Kars. & Roum.) Hansf.,

Sydowia 10 (1957) 50, was collected in Tu Phap and determined as Meliola

reticulata Kars. & Roum., Rev. Mycol. 12 (1890) 78. Asteridiella verrucosa

Foliicolous Ascomycetes in Vietnam 5 he

(Pat.) Hansf., Sydowia 10 (1957) 51 is based on Meliola verrucosa Pat., J.

Bot. Paris 11 (1897) 347, (=Irenina verrucosa (Pat.) F. L. Stevens, Ann.,

Mycol. 25 (1927) 457, with Bon 5840 (FH 7719) as its type collected from

Dinh Hoa; Bon 5851 was cited as an additional specimen in the original

description.

3. Asterina Lévy, Ann. Sci. Nat. Bot. sér. HI. (1845) 59.

Ascospore two-celled, dark brown; ascus ellipsoidal, with thickened apex;

paraphyses filamentous; ascocarp dark brown, initially forming a shield of

radiate tissue over hymenium, becoming rounded, the upper portion

breaking away with release of ascospores; mycelium darkly pigmented,

hyphodiate.

Asterina melatomatis Lév., Ann. Sci. Nat. Bot. sér. III. (1845) 59.

Ascospores 25 x 13 um; asci 40-50 x 24-30 um; paraphyses hyaline,

multicellular, 35 um; ascocarp up to 320 um diam.

Illustration: Miller & Arx, 1962: Fig. 31.

Specimen examined: Cuc Phoung National Park, Park Center, Don R.

Reynolds VV339, 19 December 1996.

The material of this new Vietnam record is similar to A. melatomatis Lév.,

Ann. Sci. Nat. Bot. ser III. 3 (1845) 59, described from Brazil, but with

ascocarps that measure 150 um diam. The smaller ascospore size

distinguishes A. melatomatis from previously reported Vietnam species.

Karsten and Roumegueére (1890) described several species of Asterina

based on B. Balansa’s Tonkin collections including A. balanseana Kars. &

Roum., Rev. Mycol. 12 (1890) 76, from Hanoi, A. insignis Kars. & Roum.,

Rev. Mycol. 12 (1890) 77, from Tu Phap, A. pauper Kars. & Roum., Rev.

Mycol. 12 (1890) 77, from Tonkin, A. setulosa Pat., J. Bot. 4 (1890) 62, and

A. sphaerotheca Kars. & Roum., Rev. Mycol. 12 (1890) 76, from Dong

Dong.

4. Atractilina Dearness & Bartholomew. Mycologia 16 (1924) 175.

Mitospores colourless to pale brown, fusiform, septate, with hilum, produced

from polyblastic, sympodial, denticles; mitosporophores brown pigmented,

erect, synemata forming; mycelium superficial, with pale brown

pigmentation.

Atractilina parasitica (Winter) Deigh. & Piroz., Mycol. Papers 128 (1972)

34.

76 Gard. Bull. Singapore 51 (1999)

Mitospores ovoid to clavate to lanceolate, roughened, 30-80 um x 3.5-9

um, terminal end narrowing into a small terminal swelling; fruit body to

550 um in height.

Illustration: Deigh. & Piroz., 1972: Fig. 17-19.

Specimens examined: Cuc Phoung National Park, Park Center, Don R.

Reynolds V309, V517, V328, V332, V337, V339, 19 December 1996; Cuc

Phoung National Park, trail from Park Center, Don R. Reynolds V342, 19

December 1996. Ba Vi National Park, 800 m elevation, Don R. Reynolds

V366, V374, V375, 21 December 1996 (LAM).

This new record for Vietnam is widely known from the tropics as a parasite

on species of Amazonia, Asteridiella, Balladyna, Irenopsis and Meliola and

is recognized by its pale brown mycelium, which forms directly over the

superficial mycelium. The species is also known from Babuyan, Puerto

Princessance, Palawan, Philippines (G. FE. Edaio PNH 20531 and IMI

76854b).

5. Hyalosphaeria F. L. Stevens, Trans. Ill. Acad. Sci. 10 (1917) 172.

Ascospores fusiform, clavate or cylindric, hyaline to pale cinnamon,

multiseptate; ascus fissitunicate, clavate; pseudoparaphysis present or absent;

ascocarp white, luteous or brick, not changing colour in KOH, subglobose

to discoid.

Hyalosphaeria miconiae F. L. Stevens, Trans. Ill. Acad. Sci. 10 (1917) 172.

Ascospore narrowly clavate to cylindric, ends rounded, 3-trans-septate,

36-57 x 4-5 um; ascocarps 100-140 um diam.; non-ostiolate but open at

maturity.

Illustrations: Pirozynski, 1977: Fig 4, A-C. Rossman, 1987: Fig. 34.

Specimen examined: Cu Phoung National Park, Park Center, Don R.

Reynolds V326, 19 December 1996 (LAM).

The type specimen of H. miconiae is F. L. Stevens 207 (ILL) from Puerto

Rico. The Vietnam collection is similar in most aspects to H. miconiae

except that the ascospores are smaller and have sublate rather than rounded

ends. Pseudoparaphyses are present as illustrated by Pirozynski (1977).

The ascocarps measure 81 um in length. The H. miconiae hyphae are

surrounded and perhaps parasitized by hyphopodate and darkly pigmented

hyphae. The identity of the hyphae as belonging to a Meliola sp. is suggested

by the typical 4-trans-septate spores sans ascocarp present in the

immediately close mycelial mat.

Foliicolous Ascomycetes in Vietnam vy

6. Leprieurina Arnaud, Ann. l’Ecole nat. d’Agric. Montpellier. 16 (1918)

210. |

Mitospores 2-celled, brown, pyriform; conidiophores short, lining upper

conidial centrum wall; pycnidum shield-shaped; mycelium brown, without

hyphopodia.

Leprieurina winteriana Arnaud, Ann. |’Ecole nat. d’Agric. Montpellier 16

(1918) 211.

Mitospores, granulate, with apical cell 70% larger than the second cell, 26

x 17 um; pycnidium 1-3 mm.

Illustration: Arnaud, 1918: Pl. 47.

Specimen examined: Cuc Phoung National Park, trail from Park Center,

Don R. Reynolds V341, 19 December 1996 (LAM).

This species is anew Vietnam record. Arnaud (1918) based his new species

on A. Puttemans 149 (P), Brazil S. Paulensis. He indicated that Prillieuxina

winteriana (Paschke) Arnaud, Ann. |’Ecole nat. d’Agric. Montpellier 16

(1918) 211 is the associated ascosporic state but this was not observed in

the recent Vietnam collections.

7. Malacaria H. Sydow, Ann. Mycol. 28 (1930) 69.

Ascospores clavate, fusiform or cylindric, multiseptate, pale grey; ascus

fissitunicate; pseudoparaphyses unbranched, septate; ascocarp superficial

with a thin stroma, dark luteous to brick red, unchanged in KOH, walls

smooth or with hairs.

Malacaria meliolicola H. Sydow, Ann. Mycol. 28 (1930) 69.

Ascospores narrowly clavate with elongate basal end that are bluntly

rounded, 3-septate, pale grey, 40-48 x 3-5 um; ascus clavate, 44-56 x 10-2

lum, pseudoparaphyses unbranched, up to 120 um; ascocarp 150-200 x

100-140 um in diam..

Illustration: Rossman, 1987: Fig. 4.

Specimen examined: Ba Vi National Park, 1000 m elevation, Don R.

Reynolds V520, 22 December 1996 (LAM).

The ascospore lengths of this new record for Vietnam fall within the 40-48

lum range described by Rossman (1987), but have a less tapering basal cell.

78 Gard. Bull. Singapore 51 (1999)

The ascospores of M. luxurians (Rehm) Rossman Mycological Papers. 157

(1987) 12, based on Paranectria luxurians Rehm, Leafl. Phil. Botany 8

(1916) 2924, found on C. F. Baker, Fungi Malayana 171 from the Philippines,

measure up to 175 um. Those of the Hansford type from Uganda measure

80-95 x 3-4 um including a “30-45 x 1 um basal appendage” (Hansford,

1961).

8. Meliolaster Hohn., Berich. Deut. Bot. Gesell. Berlin 35 (1918) 700.

Ascospores 3-celled, dark brown, fusiform; asci obpyriform, thickened at

apex; asci globose, apically thickened, paraphyses present; ascocarp a dark

brown thyriothecium, upper layer of radiate hyphae, opening with breaks

originating in a stellate pattern from ascocarp centre; mycelium of darkly

pigmented, hyphopodate.

There was some confusion concerning the name for this genus. Arnaud

(1917) used the name Patouillardina for a new, monotypic genus. Bresadola

(Rick, 1906) previously utilized Patouillardina for a name based on a

basidiomycete (Bresadola, 1925; Donk, 1958; Martin, 1939; Rogers, 1936).

Stevens (1927) regarded Melioaster described by Doidge (1920) as a

synonym of Amazonia Theissen, Ann. Mycol. 11 (1913) 499.

Meliolaster clavispora (Patouillard) Héhn., Berich. Deut. Bot. Gesell.

Berlin 35 (1918) 700.

Ascospores 40-50 x 10-16 um; ascus 60-100 x 44-80 um, paraphyses hyaline;

ascocarp 200 um in diam.

Illustrations: Arnaud, 1918: Pl. 37; Patouillard, 1890: Fig. 4.

Specimen examined: Cuc Phoung National Park, trail from Park Center,

Don R. Reynolds V348, 19 December 1996 (LAM).

This species was first described as Meliola clavispora Pat., J. Bot. 4

(1890) 61, with the type specimen collected at Tu-Phap (P). It was later

renamed as Patouillardina clavispora (Pat.) Arnaud, Comp. Rend. Acad.

Sci. Paris 164 (1917) 181. A Balansa specimen at P, issued in the exiccatum

set “Campignon du Tonkin 1887-1889”, is identified as the type. Both

Arnaud (1917, 1918) and von Hohnel (1918) critically examined this species

based on an isotype specimen that was issued as Roumeguere Fungi, selecti

exsiccati 5631. Both authors found the fungus to be microthyriaceaeous.

Patouillard (1890) remarked: “Espeéce bien distincte de toutes les congéneres

[Dimerosporium] par ses spores a deux cloisons”. Von Hohnel (1918) noted:

“Der Pilz ist ein Dimerosporium Fuckel mit drizelligen Sporen...”

——<e ss

Foliicolous Ascomycetes in Vietnam 719

9. Polychaeton (Persoon) Lév., Dict. Univ. d’Hist.Nat. 8 (1847) 454.

Fruit body comprised of an upright stalk formed of coalesed hyphae; a

mitosporic centrum is formed along its length; mitosporophores phialidic;

mitospores hyaline and unicellular.

Polychaeton citri (Per.) O. Kunze. 1891: 13.

Mitospores hyaline, aseptate, smooth, ovoid, 5-6 um; mitosporogenous

centrum in somewhat basal to midregion of column, 58 x 32 um; stalk may

extend to 31 um above the centrum, and below 289-6 um, almost becoming

sessile on the substratum.

Illustrations: Batista & Ciferri, 1962b: Figs 4, 5, Figs. I: 8, V: 26, 28;

Whew y tle 35.135) VIL 434 X> 58: Xl:.62; X64 XUE 71,72; XIV: 78;

Do a. 23: XOCIT.

Specimen examined: Ba Vi National Park, 800 m elevation, Don R. Reynolds

V338, 21 December 1996 (LAM).

This species is similar to Capnodium bambusae Roum., Rev. Bot. 12 (1890)

160. The type specimen was collected from Hai Phong in December 1889,

and distributed as C. Roumeguere Fungi exsiccati precipue Gallici. Centurie

LV. 5436.

10. Sarcinella Sacc.. Michelia 2 (1880) 31.

Mitospores solitary, subspherical, dark brown, smooth, 4-celled and

muriform, flattened; mycelium hyphodiate.

Sarcinella raimundoi Sacc., Ann. Mycol. 12 (1914) 313.

Mitospores dictyosporus as a 6-8 cell packet, 33-35 um diam., darkly

pigmented; mycelium darkly pigmented.

Specimens examined: Ba Vi National Park, park headquarters, Don R.

Reynolds V315, 19 December 1996; Ba Vi National Park, Don R. Reynolds

V356, 22 December 1966 (LAM).

This new record for Vietnam was originally described from M.B.

Raymundo-Baker 2016 (AM, NY and BPI) from Morong Valley in Rizal

Province, Philippines. The species was distributed as a C. F. Baker Fungi

Malayana 192, from Mt. Maquiling near Los Bafios, Laguna Province,

Philippines,

The Saccardo diagnosis included mention of fusoid, curved, 3-septate

SO Gard. Bull. Singapore 51 (1999)

mitospores characteristic of Questieriella Hughes, Can. J. Bot. 61 (1983)

1729. Hughes (1953, 1987) cited Questieriella as a “synapomorph” of S.

raimundii. The predicted pleomorphy of Schiffnerula Hohn. Sitz. Akad.

Wiss. Wien. Math. Naturwiss. Kl. Abt. 1, 118 (1909) 868 with Sarcinella

(Hughes, 1983) has not been established with S. raimundoi.

11. Sirosporium Bubak and Sereb. apud Bubak, Hedwigia 52 (1912) 272.

Mitospores solitary, cylindrical, brownish, smooth to verrucose, with

transverse and often longitudinal septa; conidiophores clustered, producing

conidia from apex and laterally, branched or unbranched; mycelium partly

immersed.

Sirosporium carissas Kapoor, Trans. Brit. Mycol. Soc. 51 (1968) 330.

The small fascicles of conidiophores produce brown, multiseptate

mitospores from conspicuous scars; mitospores obclavate, mostly trans-

septate with an occasional longiseptum, measuring up to 200 x 10 um.

Illustration: Ellis, 1976: Fig. 226.

Specimen cited: Cuc Phoung National Park, trail from Park Center, Don

R. Reynolds V353, 19 December 1996 (LAM).

This new record from Vietnam was originally described from India.

12. Trichothyrium Speg., Bol. Acad. Cienc. Cérdoba 11 (1889) 555.

Ascospore clavate, hyaline, 1-septate, with upper cell smaller than the

attenuated lower one; ascus obclavate with thickened wall, paraphysate;

ascocarp a thyriothecium, a darkly pigmented, orbicular, ostiolate shield;

mycelium comprised of septate, brown pigmented hyphae, most often

forming a plate that widely covers the hyphae and hyphodia of the

Meliolineae.

Trichothyrium reptans (Berk. & Curtis) Hughes, Mycol. Papers 50 (1953)

85.

Ascospore 15-20 x 5-7 um; ascus obpyriform, 30-40 x 11—16um, with apically

thickened wall; paraphyses hyaline 18 um; ascocarp up to 160 um in diam.;

mitospore termed an isthmospore, rounded to oblong in surface view, 12-

15 x 11-14 um; mycelial plate up to 75 um wide.

Illustration: Hughes, 1953: Figs. 48, 49.

Specimens examined: Cuc Phuong National Park, primary forest, Don R.

Foliicolous Ascomycetes in Vietnam 81

Reynolds V534, V300, V536, 17 December 1996; Cuc Phoung National

Park, trail from Park Center, Don R. Reynolds V318, V342, 19 December

1996; Cu Phoung National Park. Research Station Building, Don R.

Reynolds V034, V334, 20 December 1996.

This species and its mitospore were discussed in detail by Hughes (1953).

The fungus is known from herbarium collections at IMI from Malaya and

the Philippines and from the literature from the neotropics and Africa

(Hughes, 1953). The isthmospores (=/sthmospora glabra Stevens, Bot. Gaz.

45 (1918) 224) in this new record for Vietnam differ from those reported

from neotropical collections. The two pairs of pale brown cell components

of these complex mitospores are thick-walled, but each pair possesses more

than two upwardly and outwardly directed, rounded horns that are

irregularly scattered over their surface.

Acknowledgements

Acknowledgement is made to the United States National Science

Foundation, the Maggie Taylor Research Fund of the Los Angeles County

Museum of Natural History Foundation, and Florine Butler Reynolds for

financial support of field work in Southeast Asia.

References

Anonymous. 1996. WWF International Country Profile. Vietnam. World

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Gardens’ Bulletin Singapore 51 (1999) 85-98.

Reappraisal of Olea Species in Malesia

RUTH KIEW

Singapore Botanic Gardens, Cluny Road,

Singapore 259569

Abstract

One new species, Olea moluccensis Kiew from the islands of Buru and Taliabu, is described

and new combinations are made for three Philippine species, O. gitingensis (Elmer) Kiew,

O. obovata (Merr.) Kiew, and O. rubrovenia (Elmer) Kiew, all originally described as

Linocieras. Olea decussata (Heine) Kiew is synonymous with O. rubrovenia, Linociera

lauterbachii Lingelsh. with Olea paniculata R. Br.; L. longifolia Merr. with O. gitingensis,

and L. pallida (Merr.) Merr. and L. philippinensis Merr. with O. borneensis Boetl..

Introduction

Following the account of Olea for the Malesian region (Kiew, 1979), revision

of Chionanthus for New Guinea and the Philippines reveals that several

species previously described as Linociera have to be transferred to the

genus. Consequently, nomenclatural changes are required.

Neither Elmer (1913) nor Merrill (1915) reported Olea species in

their checklists of Philippine plants but both described Olea species under

Linociera (Linociera gitingensis and L. rubrovenia by Elmer and L.

longifolia, L. obovata and L. philippinensis by Merrill). The Philippines

has in fact the greatest number of Olea species compared with other areas

in Malesia (Fig. 1), including three endemics, O. gitingensis, O. obovata

and O. palawanensis (Kiew, 1993).

Lingelsheim (1927) also did not include any Olea species in his account

of the New Guinea Oleaceae but described Linociera lauterbachii, which is

in fact conspecific with Olea paniculata R. Brown (1810).

Olea in Malesia is represented by a ‘mixed bag’ of species, which can

be distinguished into four distinct species groups.

The first group is represented by a single species, Olea paniculata,

which is unique among Malesian Olea species in possessing terminal as

well as axillary inflorescences, an indumentum of large lepidote trichomes

(even on the petals), hermaphrodite (as opposed to polygamous) flowers,

corolla lobes that open widely and are not cucullate, stamens attached at

the top of the corolla tube with a distinct filament that fully exposes the

versatile anther, large and globose stigmas, and domatia on the leaves

including those of the seedling (Brouwer, 1979). These domatia are domed

86 Gard. Bull. Singapore 51 (1999)

Slacsine D heey

Pay

Figure 1. Distribution of Olea species in Malesia.

(Upper figure - number of endemic species, lower figure — total number of species)

with a wide mouth, which contains hairs, and so differ from those of

Jasminum, which are in the form of a dense tuft of hairs.

Olea paniculata belongs to Section Ligustroides (Bentham and

Hooker,1876), which includes an austro-african group of Olea species. In

Malesia, it extends from New Guinea westwards to E. Java.

No other Malesian species possesses domatia or large lepidote

trichomes, although they do have (as do Chionanthus species) small sunken

peltate trichomes on the lower leaf surface, which may appear minutely

punctate (Kiew & Che Su, 1982). All other species are polygamous with

subsessile stamens attached near the base of the corolla tube; cucullate

corolla lobes that when open expose only the tip of the anthers; and minute

stigmas.

The second group comprises species previously included in the genus

Tetrapilus. Among these is Olea brachiata (Lour.) Merr. (basionym: T.

brachiatus Lour.), which is characterised by its subserrate leaves. Several

other Malesian species, namely O. borneensis, O. palawanensis and O.

salicifolia exhibit this character, although it may not be very obvious due

to the teeth being small, distant and confined to the margin of the upper

half of the lamina.

The third group of species is distinct in possessing entire, coriaceous

Olea in Malesia 87

leaves that dry pale grey green or fawn and have obscure venation. This

group includes three Malesian endemics, viz. O. moluccensis (endemic to

the Moluccas Islands), and O. gitingensis and O. obovata (endemic to the

Philippines).

The fourth group comprises the two montane species, Olea javanica

and O. rubrovenia. These species differ markedly from the other Malesian

species in having entire leaves that dry brownish with the lateral veins

conspicuously darker beneath and unthickened petioles that dry black.

Blume described specimens of Olea javanica first in 1826 under the

genus Pachyderma and again in 1828 under the genus Stereoderma. Bentham

and Hooker (1876) reduced Tetrapilus and Pachyderma (including

Stereoderma) to synonomy with Olea. Johnson (1957), on the other hand,

reinterpreted Olea more narrowly to include only those species with lepidote

trichomes, entire leaves and terminal and axillary inflorescences, which in

Malesia would include only Olea paniculata. He then reinstated Tetrapilus

as a distinct genus, in which he included Pachyderma (Stereoderma), to

accommodate those species that had dentate or entire leaves and which

did not have lepidote trichomes nor terminal inflorescences. He included

O. javanica and O. rubrovenia, as well as Olea brachiata and O. dentata

(the latter synonymous with O. salicifolia Wall. ex G. Don) within Tetrapilus,

which as shown above represent two dissimilar groups of species.

While it is important to appreciate the morphological diversity within

Olea and the closely related Chionanthus, the division of Olea into several

narrowly defined genera would create more taxonomic problems within

the Oleaceae. If Johnson’s narrow generic delimitation is followed, the

two sections of Chionanthus in Malesia, namely Sect. Ceranthus and Sect.

EuLinociera, should also be raised to generic level as the differences

between them (Kiew, 1979) are as great as those between Olea and

Tetrapilus sensu Johnson. The proliferation of generic (and subgeneric)

names at this time, when the full diversity of tropical and subtropical Olea

and Chionanthus species worldwide is not fully understood, is premature.

This is particularly the case as different combinations of characters

are useful to distinguish species of Olea from those of Chionanthus in

different regions of the world. For example, in South Africa, Verdoorn

(1956) reported that Chionanthus (Linociera) species possess domatia but

lack endosperm in the seed, which instead has thick cotyledons. In contrast,

African species of Olea do not have domatia and the seeds are endospermic.

In Malesia, no Chionanthus species has domatia and apart from species in

Sect. Ceranthus, e.g. C. ramiflorus Roxb., the seed is endospermic.

For these reasons, Olea is here recognized in its traditional broad

sense to include Tetrapilus.

This account includes the nomenclatural changes for species

88 Gard. Bull. Singapore 51 (1999)

transferred from Linociera to Olea, updates synonomy and includes the

description of a new species from the Moluccas islands. A key to identify

the ten Malesian species is also provided. For completeness, reference is

made to recent publications with descriptions for all Malesian species.

Key to Malesian Species of Olea

la. Inflorescences terminal and axillary. Leaves with domatia and large

lepidote trichomes. Flowers hermaphrodite .............. 8. O. paniculata

1b. Inflorescences axillary, never terminal. Leaves without domatia and

lepidote trichomes (though sunken peltate glandular trichomes are

present). Flowers polyeamacatsye 4 oo os.4.i..040) ou ceeecsee a tule tet eee ee 2

2a. Leaf margin entire, lamina drying pale grey-green or fawn, lateral

veins obscure Deneathy aii ch cn saeencaee abel oper tae oe 3

2b. Leaf margin serrate or subserrate or, if entire then leaf dries brown

with the lateral veins conspicuously darker beneath...............e eee 5

3a. Lamina 3.5-7 cm long, about twice as long as wide, apex rounded....

ssibyed ah dietetic Adah Re NMOL ae ea a 6. O. obovata

3b. Lamina 14.5—28.5 cm long, about three times longer than wide, apex

ACUTE £O ACMI ALE spinnin geen ebsinpiisedynn gore celibate aches bacteutital 5 Onalaska en 4

4a. Inflorescences lax, 3.5—7 cm long. Pedicels 2-6 mm long. Fruits c. 5 x 4

mm with thin and: brittle pericarp. 22) oe 3. O. gitingensis

4b. Inflorescences condensed, 1—2 cm long. Pedicels c. 1 mm long. Fruits

c. 16 x 15 mm with thick and leathery pericarp ...... 5. O. moluccensis

Sa.. Lamina margin entire ici eee en he ee ee 6

Sb. , Lamina margin. serrate ‘or subsemrate oi. ee iz

6a. Axillary buds large and mammose. Lateral veins of leaves 6—10 pairs.

Pedicels 1-4 mm long, stamens always 2................0+- 9. O. rubrovenia

6b. Axillary buds minute and rounded. Lateral veins of leaves 4~7 pairs.

Pedicels 2-8. mm. stamens'2:) 3) OT 4 ji) a ee 4. O. javanica

7a. , Leet Dass COnt ates iia ce an Wee eee 7. O. palawanensis

7b. Leaf base not Cordates oo. sin sicicrencssiec uta ladsatghonet ier sald apes eee 8

8a. Petiole conspicuously thickened and drying pale fawn. Lamina

frequently obovate, drying grey-green. Flower very small (up to 1 mm

LOWE) scosoucaccsssierpatoncsidlecsoaveneee ane eee 1. O. borneensis

Olea in Malesia 89

8b. Petiole not thickened, drying black. Lamina lanceolate. Flowers larger

fee ym i a ee el ON es ae, Us elal uelh 9

9a. Lamina 2-2.5 times longer than wide, veins impressed above.

Inflorescence short (1-2.5 cm), glabrous. Flowers 1.5-2.5 mm long,

OE RE LON CORI es okt) ae PaaS 7 ae 2. O. brachiata

9b. Lamina 3 times longer than wide, lateral veins not impressed above.

Inflorescence longer (9-11 cm with a range of 3.5-19.5 cm long), finely

downy. Flowers 2—5 mm long, calyx finely downy ..............ccceesseeeeeees

1. Olea borneensis Boerl., Handl. Fl. Ned. Ind. 2 (1899) 332; Kiew, Blumea.

25 (1979) 308. Type: Korthals s.n., Borneo (L lecto)

Synonyms: Olea sp., Vidal, Phan. Cuming. Philip. (1885) 125, Rev. PI.

Vasc. Filip. (1886) 181. Mayepea pallida Merr., Gov. Lab. Publ. (Philip) 35

(1906) 58. Linociera pallida (Merr.) Merr., Philip. J. Sci. 1 (1906) Suppl.

116 non K. Schumann (1901). Linociera philippinensis Merr., Philip. J.

Sci.(Bot.). 4 (1909) 313. Type: Ahern’s Coll. 2874, Bosoboso, Rizal Prov.,

Philippines (K lecto).

Notes: Elmer (1913) and Merrill (1915) did not recognise Olea as being

present in the Philippines and Merrill (1906) renamed Vidal’s Olea species

as Linociera (Mayepea) pallida. Because K. Schumann (1901) had previously

described a species from New Guinea as L. pallida, Merrill renamed the

Philippine species L. philippinensis. The Philippine specimens match those

from Borneo and so L. philippinensis becomes a synonym of O. borneensis.

Olea borneensis most resembles O. brachiata and O. salicifolia in its

foliage, which is at least distantly toothed in the upper half of the lamina.

It is easily distinguished from these two species by its thickened petiole. In

addition, it is distinct among Malesian Olea species in possessing extremely

small flowers.

Its geographical distribution does not extend to the Asian mainland.

Endemic to Malesia, it is found as far north as the Guimaras Islands in the

Philippines. It has been most commonly collected from Luzon. Further

south it has been collected from Mindoro (Mt Yagaw), Mindanao (Mt

Urdaneta), Palawan (Victoria Mts) and Sibuyan Is. (Mt Giting-giting). In

Borneo, it is more common in the eastern parts with several collections

from Sabah and a few from E. Kalimantan (Martapoera).

It has been collected from the lowlands (170 m) but is most commonly

recorded from hill slopes, ridges and rocky ridge tops (up to 5000 m a.s.1.

in Sabah). On Mt Yagaw, Mindoro, it was reported as ‘badly battered by

wind’ (Sulit & Conklin PNH 17640). It has also been recorded from

90 Gard. Bull. Singapore 51 (1999)

ultrabasic soil on Mt Silam, Sabah.

Notes on Addurn 226 report that on Mt Ibi in Lagum, Luzon, its

wood was considered ‘good for house construction beams’.

2. Olea brachiata (Lour.) Merr., Lingn. Agr. Rev. 2 (1925) 127, Kiew,

Blumea. 25 (1979) 308. Basionym: Tetrapilus brachiatus Lour., Fl. CochinCh.

2 (1790) 611. Type: Louriero (BM lecto).

Synonym: O. maritima Wall. ex G.Don., Gen. Syst. 4 (1838) 49. Type:

Wallich Cat. 2813. Singapore. (K lecto)

Notes: Bentham & Hooker (1876) included Tetrapilus Lour. and

Pachyderma (Stereoderma) Blume as synonyms of Olea. In addition, they

listed Pachyderma javanica as synonymous with Olea maritima (Wall.) ex

G. Don and that Notelaea zollingeriana belonged to Olea. This was followed

by later authors until Knoblauch (1895) recognised O. javanica as a species

distinct from O. maritima. However, as he had not seen a specimen of N.

zollingeriana, he made no change to its position as a synonym of O. maritima

and there it has remained. (Olea maritima is currently considered a synonym

of O. brachiata (Lour.) Mertr.)

Javanese specimens referred to O. brachiata (Kiew, 1979) are atypical

for the species in having entire leaf margins and in their habitat (inland,

usually from mountain forest as opposed to being common on rocky shores).

Re-examination of these specimens leads to the conclusion that they fall

within the variation of O. javanica and that O. brachiata does not occur in

Java.

The other Javanese taxon linked with O. brachiata is O. graciliflora

Koord. & Valeton (Backer & Bakh.f., 1965). They distinguished this latter

species from O. javanica in their key by its more floriferous and finely

pubescent inflorescence and its long pedicels. Apart from its pubescence,

O. graciliflora does not differ from O. javanica, neither does it have the

serrate leaves of O. brachiata.

For these reasons, both N. zollingeriana and O. graciliflora are here

assigned to the synonomy of O. javanica (see below).

3. Olea gitingensis (Elmer) Kiew, comb. nov.

Linociera gitingensis Elmer, Leafl. Philip. Bot. 5 (1913) 1653. Type: Elmer

12290, Mt Giting-giting, Sibuyan Is. (US lecto; Gray, K iso).

Synonym: Linociera longifolia Merr., Philip. J. Sci. (Bot). 20 (1922) 431.

Type: Miranda FB 20626, Lanao, Mindanao (US lecto).

Olea in Malesia 9]

Notes: Olea gitingensis is endemic to the Philippines, not common but

widely distributed in the lowlands at about 100 m altitude. It is a most

distinctive species in its large, thick, broadly lanceolate leaves, which are

glossy above. It most resembles O. borneensis in its leaves, which have

obscure venation, dry greyish-green and in its thickened, fawn petioles, but

its lamina is never toothed.

Merrill (1922) noted its alliance with Olea, but his material comprised

only the type specimen, which was in young fruit, which did not allow him

to settle this point. The foliage is typical of Olea and, in addition, the

corolla lobes are shorter than the tube (a generic distinction for Olea, as

Chionanthus flowers in most cases have an extremely short corolla tube).

For these reasons, this species is transferred from Linociera to Olea.

This distinctive species has been collected from only five localities -

Mt Giting-giting, Sibuyan Is (Elmer 12168, 12290); Lanao (Miranda FB

20626) and Butuan (Mendoza PNH42269) from Mindanao; Lamao, Bataan

Prov., Luzon (Merrill 8626); and Mt Timbaban, Panay (Edano BS42391).

The single difference between the descriptions of L. gitingensis and L.

longifolia is in the number of lateral veins, 6-8 pairs and 13-15 pairs,

respectively. However, re-examination of the leaves of the type specimens

show that L. gitingensis has between 8 and 10 pairs and L. longifolia between

10 and 11, so this difference is not supported.

Blner (1913),7im describing £. gitingensis referred to. another

collection from Agusan Province, Mindanao (Elmer 13479), which in fact

belongs to O. borneensis.

At a glance, the foliage of O. gitingensis is closely similar in shape

and thickness to that of Chionanthus remotinervius (Merr.) Kiew. This

latter species is, however, typical of Chionanthus in its unthickened petiole

that dries black and in its thick pericarp. (The ripe fruit of O. gitingensis

remains unknown).

4. Olea javanica (Blume) Knobl., Bot. Centralbl. 61 (1895) 134; Koord.

& Valeton, Meded. Lands. Plant. 59 (1902) 251; Backer & Bakh.f., Fl. Java

2 (1965) 215; Kiew, Blumea 25 (1979) 311. Basionym: Pachyderma

javanicum Blume, Bijdr. (1826) 682;

Synonyms: Stereoderma javanicum (Blume) Blume, FI. Javae Praef. 7 (1828)

8. Type: Blume 2169a, Java (L lecto; K iso).

Notelaea zollingeriana Teijsm. & Binn., Nat. Tijdschr. Ned. Ind. 27 (1864)

33. Type: Binnendijk s.n. Hort. Bot. Bogor (L lecto, K iso).

Olea graciliflora Koord. & Valeton, Meded. Lands. Plant. 59 (1902) 254.

Type: Koorders 29339 Gunung Wilis, Java (BO lecto).

92 Gard. Bull. Singapore 51 (1999)

Notes: Previously known only from mountains in Java, post-war collections

have extended its geographical range to Flores and W. Sumbawa (Mt.

Batulanteh) to the east and Sumatra (with a single sterile specimen from

G. Pesagi) to the north.

Olea javanica is not a particularly variable species — the varieties

described and illustrated by Koorders & Valeton (1902) fall within the

range of variation of the species (Kiew, 1979). Their identical var.

acuminatissima and var. laxiflora are, however, striking in their narrow

leaves (three times longer than wide), which have extremely attenuated

apices, and long, slender pedicels (S-10 mm long) compared with most

specimens of this species, which have broader leaves up to 2.5 times longer

than broad, acuminate apices and pedicels 3-5 mm long. An additional

difference is recorded on Lanjouw 61: “fruits long-stalked hanging first

green later black violet.” (In all other varieties, and indeed in all other

Olea species, the fruits sit on upright pedicels). It is possible that this

difference has biological significance in terms of the dispersal agent. For

other species of Olea and Chionanthus with fruits of a similar size that

ripen purple black, birds are reported as dispersal agents for their fruits.

Perhaps a different bird species may disperse these pendant fruits but until

and unless there are data to support this, I refrain from recognising var.

acuminatissima as a distinct variety.

Incidentally, the type of Blume’s Pachyderma (Stereoderma) javanica

(Blume 2169a) has similar narrowly attenuated leaves. The type of Notelaea

zollingeriana also has narrowly lanceolate, entire leaves that are closely

similar to those of Blume 2169a. For this reason, it is here recognised as a

synonym of Olea.

5. Olea moluccensis Kiew, sp. nov.

Differt a Olea gitingensis, inflorescentia brevibus condensatis foliis obovatis

et fructis majoribus.

Typus: Hulstijn & Atje 364, Taliabu Is., Moluccas (BO (sheet no.1518-4) -

holo; BO (sheet no. 95014-14), L — iso).

Figure 2.

Habit and bark unknown. Twigs moderately stout, drying white, nodes

strongly flattened, glabrous, lenticellate. Petiole (0.8—)1-1.5 cm long and

3-5 mm thick, thickened and drying pale fawn, deeply grooved above.

Lamina thickly coriaceous, glabrous, drying light brown or grey-green,

slightly glossy above, slightly obovate, (14.5—)21(—28.5) x (5—)6.5(-9.5) cm,

base cuneate and decurrent, margin entire and recurved, apex acute to

Olea in Malesia 93

Figure 2. Olea moluccensis Kiew.

(A. twig; B. male flower; C. stamen and cucullate corolla lobes; D. fruit; E. T.S. fruit - Hulstijn

& Atje 364, Talibu Is., Moluccas).

94 Gard. Bull. Singapore 51 (1999)

acuminate, midrib slightly impressed above, prominent and glossy beneath,

lateral veins 11-14 pairs, slightly prominent above and beneath, intercostal

veins conspicuous and slightly prominent above in the dried state with a

distinct central vein parallel and midway between the lateral veins, marginal

vein c. 3-4 mm from margin.

Inflorescences axillary, a once-branched condensed panicle, solitary,

10-20 mm long, peduncle 2-3 mm long, lowermost branch 5-10 mm long,

slightly pubescent. Flowers crowded. Bracts scarious, lowermost c. 3 mm

long, uppermost c. 1 mm long, persistent, margin ciliate, apex pubescent.

Pedicel c. 1 mm long. Male flowers: buds almost globose, 1-1.3 mm diam.

Calyx c. 0.5 mm long, divided slightly more than halfway, lobes acute

opening widely, glabrous except for ciliate margin. Corolla 1-1.5 mm long,

divided slightly less than halfway, lobes oblong, cucullate, apex rounded,

opening to expose anther apices. Stamens 2, subsessile attached to the

base of corolla, anthers oblong, c. 0.5 mm long, connective broad. Bisexual

flowers not known. Fruit globose, c. 16 x 15 mm, drying with whitish

bloom, pedicel thickened, 2-3 mm long, pericarp drying leathery, c. 2 mm

thick. Seed unknown.

Specimens examined: Moluccas: Buru Is. de Vriese & Teysmann HB 1834

(BO, L), de Vriese s.n., sheet no. 1857-61 (L); Taliabu Is. Hulstijn & Atje

364 (BO, L). |

Notes: A most distinctive species of Olea in its large, thick leaves and much

condensed inflorescences. Its foliage closely resembles that of O. gitingensis

in its size, coriaceousness and entire margin but it differs from O. gitingensis,

which has lanceolate leaves, larger, lax inflorescences (3.5—7 cm long with

lowermost branches 1-2.5 cm long), longer pedicels (2-6 mm long) and

smaller fruits (c. 5 x 4 mm), which have a thin and brittle pericarp.

It is endemic to the Moluccas Islands (hence the specific epithet).

All specimens lack field notes so its habit and habitat are not known.

Unfortunately the specimens from Buru are sterile.

6. Olea obovata (Mertr.) Kiew, comb. nov.

Basionym: Linociera obovata Merr., Philip. J. Sci. (Bot.) 10 (1915) 338.

(Elmer, Leafl. Philip. Bot. 5 (1913) 1652 nomen nudum). Type: Ramos

15014, San Antonio, Laguna Prov., Luzon (K lecto).

Notes: As Merrill (1915) noted, this is a very distinctive species in its small,

thick, obovate leaves with a rounded apex and decurrent base, obscure

lateral veins, recurved margin and dense pyramidal inflorescences and it is

Olea in Malesia 95

quite unlike any other Malesian Olea species.

It is known only from Luzon, Philippines, with many of the collections

from mountainous areas (Mt. Alzapan, Mt. Binuang, Montalban, Mt.

Umingan). From Mt. Alzapan it is reported to grow in mossy forest at

2000 m (Ramos & Edano BS 45667, 45716). It has the habit of many trees

of mossy forest as judged from its much branched, bushy canopy with

bunches of upstanding, thick leaves with recurved margins at the tip of the

twigs.

7. Olea palawanensis Kiew, Blumea. 38 (1993) 127. Type: Ridsdale SMHI

1827, Narra Victoria Peaks, Palawan, Philippines (L holo).

Notes: Its cordate leaf base distinguishes this species from all the other

Malesian Olea species. It is endemic to Palawan, Philippines, where it

appears to be confined to ultramafic soils.

8. Olea paniculata R.Br., Prodr. (1810) 523; Koord. & Valeton, Meded.

Lands. Plant. 8 (1902) 256; Backer & Bakh.f., Fl. Java 2 (1965) 214; Kiew,

Blumea. 25 (1979) 312.

Synonym: Linociera lauterbachii Lingels., Bot. Jahrb. 61 (1927) 8; Kobuski,

J. Arn. Arbor. 21 (1940) 335. Type: Schlechter 16984, Minjem at Kelel, NE

New Guinea (B+; A lecto).

Notes: Lingelsheim (1927) did not record any species of Olea as occurring

in New Guinea. Although terminal inflorescences are atypical of Linociera,

he described specimens of Olea paniculata as Linociera lauterbachii. His

description differs from that of O. paniculata (Kiew, 1979) only in the

longer inflorescence (15 cm as oppposed to 6.5—8 cm) and the absence of

endosperm, which is unusual for Olea. Inflorescences of New Guinea

specimens are indeed sometimes longer (6-13 cm) than those of Australian

specimens. I have not seen any specimens that lack endosperm and it is

possible that the absence of endosperm observed by Lingelsheim was due

to immaturity of the fruit.

9. Olea rubrovenia (Elmer) Kiew, comb. nov.

Basionym: Linociera rubrovenia Elmer, Leafl. Philip. Bot. 2 (1909) 586, 5

(1913) 1652; Merr., Enum. Philip. Pl. 3 (1923) 305. Type: Elmer 10224

Negros Is., Philippines (US lecto; A, K, L iso).

Synonyms: Tetrapilus rubrovenius (Elmer) L.A.S. Johnson, Contrib. N.S.W.

Herb. 2 (1957) 408.

96 Gard. Bull. Singapore 51 (1999)

Ilex decussata Heine, Mitt. Bot. Staatssaml. Muenchen. 6 (1953) 209.

Olea decussata (Heine) Kiew, Blumea. 25 (1979) 309. Type: Clemens 28986,

G. Kinabalu, Sabah, Borneo (K lecto; L iso).

Notes: The discovery that Linociera rubrovenia is an Olea and is conspecific

with Olea decussata necessitates a name change as ‘rubrovenia’ is the earlier

name.

Olea rubrovenia is a montane species usually found above 1500 m

(but has been collected from as low as 400 m a.s.l. in mixed dipterocarp

forest on Bukit Mersing, Sarawak). It produces flushes of new leaves that

are reported as tinged pink. These new shoots frequently bear inflorescences

and their young green stems dry black.

Leaves of plants growing at high altitudes, e.g. on G. Kinabalu, or on

exposed summits have smaller (up to 9.5 x 5 cm), bunched, upstanding

leaves, which are thickly coriaceous, have recurved margins and the lateral

veins are finely impressed above compared with the larger (up to 22 x 8

cm), less coriaceous leaves of plants growing at lower altitudes.

The high altitude plants (Olea decussata (Heine) Kiew) on G.

Kinabalu are indeed distinctive but as the number of collections from this

mountain has increased, it becomes clear that the high altitude form

intergrades with the larger leaved one from lower altitudes and that Olea

decussata therefore cannot be maintained as distinct from O. rubrovenia.

Indeed, Green (1960) has already warned of ‘dangers in Malaysian

[Malesian] taxonomy which arise from phenotypic variation caused by

differences in altitude’ when he reduced the high altitude Osmanthus

sumatrana P.S. Green (Oleaceae) to the synonomy of O. scortechinii King

& Gamble. |

In Borneo, this species in Sabah is a common tree on G. Kinabalu up

to 3200 m altitude and has also been collected from G. Alab (Crocker

Range) and G. Trusmadi; in Sarawak from G. Mulu, G. Dulit, G. Murut

and Bk. Mersing; and in Kalimantan from G. Beratus. From the Philippines,

it has been collected from the Negros Is. (Cuernos Mts., type locality),

from Mindanao (Mt Apo, Mt. Camates, Mt. Matutum and Mt. Urdaneta),

and from Luzon (Mt. Nagapatan).

Olea rubrovenia closely resembles O. javanica in its entire leaves,

petioles that are not thickened and dry black, lateral veins that are

conspicuous beneath, particularly as they dry darker than the lamina,

foliaceous bracts (not always present because they are caducous), and the

frequent occurrence of globose galls replacing the male flowers. However,

the two species are distinct by a combination of the following characters:

Olea in Malesia Q7

Character O. javanica O. rubrovenia

axillary bud minute and rounded large and mammose

petiole length (mm) 410 7.5—25

no. lateral vein pairs 4—5(-7) 6—7(-10)

inflorescence length (cm) (3.5—)4.5-9 2.5—5.5

pedicel length (mm) 2-8 14

no. stamens 2.35.4 D.

Apart from being geographically separate, the clearest character that

distinguishes these two species is undoubtedly the shape and size of the

axillary buds.

10. Olea salicifolia Wall. ex G. Don., Gen. Syst. 4 (1837) 48. Type:

Wallich Cat. 2812, Silhet, India (K, lecto).

Synonyms: O. dentata Wall. ex DC Prod. 8(1844) 286. Kiew, Blumea. 25

(1979) 310. Type: Wallich 2840 (K lecto).

O. penangiana Ridl., J. Fed. Mal. States. Mus. 10 (1920) 148. Type: Curtis

950, Penang (SING lecto; K iso).

Notes: Olea salicifolia shows considerable variation from specimens with

smaller willow-shaped leaves from Silhet and Khasia, India, to specimens

with broader leaves from Thailand and Peninsular Malaysia.

In Malesia, this species is known only from Penang, where it was

once common on the rocky coasts at Batu Ferringi and Telok Bahang and

where its habitat is now much disturbed by beach resort developments,

and from a single sterile specimen (Mat 211, SING) from Singapore (precise

locality not stated) collected in 1893. This sterile specimen collected together

with a wood sample is recorded with the local name of penaga lilin, a name

usually used for the ironwood tree, Mesua ferrea (Guttiferae), an indication

that it has hard wood. Kurz (1877) recorded that in Myanmar, it (which he

called O. dentata) produces a ‘very heavy...close-grained wood’.

Acknowledgements

I am grateful to the curators of BM, BO, K, KEP, L, LAE, SAN, SAR,

SING and US for permission to examine specimens in their care. I

particularly appreciate Engkik Soepadmo’s helpful suggestions for

improving the manuscript and Rosemary Wise’s guidance in producing the

botanical drawing.

98 Gard. Bull. Singapore 51 (1999)

References

Backer, C.A. & R.C. Bakhuizen van der Brink. 1965. Oleaceae. In: Flora

of Java. 2: 212-218.

Bentham, G. & J.D. Hooker. 1876. Oleaceae. In: Genera Plantarum. 2:

672-680.

Brouwer, Y.M. 1979. Domatia in seedlings. Flora Malesiana Bulletin. 32:

3239-3246.

Elmer , A.D.C. 1913. Philippine Linociera. Leaflets Philippine Botany. 5:

1651-1657.

Green, P.S. 1960. Further notes on Malaysian Osmanthus. Notes from Royal

Botanic Garden Edinburgh. 23: 175-178.

Johnson, L.A.S. 1957. A review of the family Oleaceae. Contributions from

New South Wales Herbarium. 2: 395-418.

Kiew, R. 1979. The Oleaceae of Malesia. II. The genus Olea. Blumea. 25:

305-313.

Kiew, R. 1993. Olea palawanensis (Oleaceae), a new species from the

Philippines. Blumea. 38: 127-128.

Kiew, R. & Che Su Ibrahim. 1982. Comparative study of leaf anatomy of

Malaysian species of Chionanthus and Olea (Oleaceae) with special

reference to foliar sclereids. Botanical Journal of Linnean Society. 84:

79-101.

Knoblauch, E. 1895. Zur Kenntnis einiger Oleaceen-Genera. Botanisches

Centralblatt. 61: 134-135.

Kurz, S. 1877. Forest Flora of British Burma. 2: 157.

Lingelsheim, A. v. 1927. Beitrage zur Flora von Papuasien. 111. Die

Oleaceaeen Papuasiens. Botanischer Jahrbucher. 61: 1-22.

Verdoorn, I.C. 1956. The Oleaceae of Southern Africa. Bothalia. 4. 549-

639.

Gardens’ Bulletin Singapore 51 (1999) 99-102.

Euphorbia heterophylla and E. cyathophora

(Euphorbiaceae) in the Malay Peninsula

ILM. TURNER

Singapore Botanic Gardens

Singapore 259569

Abstract

The identities of two closely related species of Euphorbia naturalized in Southeast Asia

from the New World tropics are considered. The name Euphorbia heterophylla L. has been

consistently misapplied in the Malay Peninsula to plants of Euphorbia cyathophora Murray.

True Euphorbia heterophylla is found as a garden escape and weed of waste ground, but is

not as common in the Malay Peninsula as FE. cyathophora. The differences between the two

species are summarized.

Introduction

The botanical literature for the Malay Peninsula has generally referred to

the euphorbia with red bracteal leaves and alternately arranged foliage

leaves, that is relatively common on the upper parts of sandy beaches, as

Euphorbia heterophylla L. (e.g. Ridley 1924). However, these plants are

actually referable to a similar species, also native to tropical America,

Euphorbia cyathophora Murray. The similarity between the two species

has led some authors to consider E. cyathophora as a variety of E.

heterophylla or even to treat them as conspecific. However, most recent

treatments of the genus for different floras (Radcliffe-Smith in Airy Shaw

1977, Ma & Wu 1992, Burger & Huft 1995, Lin & Hsieh 1993, Chriztenhusz

1998), with the notable exception of Philcox (1997), have treated the two

as distinct species.

Distinguishing the Two Species

The consistent differences between Euphorbia cyathophora and E.

heterophylla are summarized in Table 1. Most authors report leaf shape to

be variable in both species and to be unreliable for separating them.

However, all the Malayan material of E. cyathophora that I have seen, has

some leaves with obvious lobing, usually forming a five-pointed outline,

whereas Malayan E. heterophylla has unlobed leaves, which generally have

a shallowly serrulate margin.

100

Gard. Bull. Singapore 51 (1999)

Table 1. Morphological differences between Euphorbia cyathophora and E.

herophylla (after Dressler 1961).

character E. cyathophora

E. heterophylla

stem cross section

indument of stem, petiole

and capsule

colour of foliage leaves

colour of bracteal leaves

cyathial gland

seed cross section

seed coat morphology

angled

glabrous

glossy green

green, often with red at

base

bilabiate with narrow,

elliptic-oblong opening

rounded

finely and sharply

tuberculate

rounded

more or less hairy

dull green

green, sometimes with

white or purplish at the

base

funnel-like with round

opening

angular

coarsely and bluntly

tuberculate

Distribution in the Malay Peninsula

Euphorbia cyathophora is relatively widespread in the Malay Peninsula,

and particularly common as a weed on the upper part of sandy beaches. It

probably naturalized from plants grown for ornamental purposes in gardens.

E. heterophylla is rarer, with records from Singapore and Pulau Tioman

only. This species is grown for medicinal purposes in Malay villages, and

has probably escaped from gardens also.

Citation and Synonymy

Euphorbia cyathophora Murray, Commentat. Soc. Regiae Sci. Gott. 7 (1786)

81; Radcliffe-Sm. in Airy Shaw, Kew Bull. 32 (1977) 75; Radcliffe-Sm., FI.

Mascareignes 160 (1982) 95; R.A. Howard, FI. Lesser Antilles 5 (1989) 48;

Huft in W.L. Wagner et al., Man. Fl. Pl. Hawaii (1990) 618; G.L. Webster

in Nicolson, Smithsonian Contr. Bot. 77 (1991) 85; J.S. Ma & C.Y. Wu,

Collect. Bot. 21 (1992) 102; S.C. Lin. & C.F..Hsieh, Fl) Taiwan, ed, 23

(1993) 458; W.C. Burger & Huft, Fieldiana, Bot. 36 (1995) 116.

Euphorbia heterophylla and E. cyathophora 101

Euphorbia heterophylla L. var. cyathophora (Murray) Griseb., Fl. Brit. W.

I. (1859) 54.

Poinsettia cyathophora (Murray) Klotzsch & Garcke, Monatsber. KGnigl.

Preuss. Akad. Wiss. Berlin 1859 (1859) 253; Dressler, Ann. Missouri Bot.

Gard. 48 (1961) 338.

Euphorbia heterophylla L. forma cyathophora (Murray) Voss in Vilmorin,

Vilm. Blumengaertn., ed. 3 1 (1895) 898.

Euphorbia heterophylla auct. non L., Ridl., Fl. Malay Penins. 3 (1924) 181;

Backer & Bakh. f., Fl. Java 1 (1963) 502; I.M. Turner, Gards. Bull.,

Singapore 50 (1995) 221.

Euphorbia heterophylla sensu lato, Philcox, Rev. Handb. Fl. Ceylon 11

(1997) 200.

Specimens examined: KEDAH, Md Haniff & Lebai Ishak SFN 10426, 5

April 1924 (SING). PERAK, Golden Sand Beach, L.E. Teo KL3051, 3

May 1972 (SING); Lumut, G.E.S. Cubbitt 9666, 22 May 1927 (SING).

PAHANG, Pulau Tioman, Lord Medway 4041, August 1962 (KLU); Pulau

Tioman, Kampong Tekek, Kadim & Noor KN676, 26 April 1962 (SING);

Pulau Tioman, L.H. Chua & K.E. Tian KL51, 4 November 1982 (SINU).

SELANGOR, Chemora, J. Carrick 610, 3 January 1959 (KLU). NEGERI

SEMBILAN, Tampin, /.H. Burkill 2296, 21 November 1916 (SING).

JOHMORE,Pencerang, 7.) feruya 2252, 12..March, 1933, (SING).

SINGAPORE, Pulau Sudong, J.F. Maxwell 81-147 (KLU); Cultivated in

Botanic Gardens, C.X. Furtado SFN 34862 (KEP); Pulau Ubin, Bukit

Macao, C.X. Furtado SFN 18341, 31 July 1927 (SING); Siglap at seashore,

J. Sinclair SFN 38875, 18 February 1950 (SING).

Euphorbia heterophylla L., Sp. Pl. (1753) 453; Radcliffe-Sm., FI.

Mascareignes 160 (1982) 94; R.A. Howard, Fl. Lesser Antilles 5 (1989) 49;

Huft in W.L. Wagner et al., Man. Fl. Pl. Hawaii (1990) 619; G.L. Webster

in Nicolson, Smithsonian Contr. Bot. 77 (1991) 85; J.S. Ma & C.Y. Wu,

Collect. Bot. 21 (1992) 102; S.C. Lin & C.F. Hsieh, Fl. Taiwan, ed. 2 3

(1993) 460; W.C. Burger & Huft, Fieldiana, Bot. 36 (1995) 118.

Poinsettia heterophylla (L.) Klotzsch & Garcke, Monatsber. Konig. Preuss.

Akad. Wiss. Berlin 1859 (1859) 253; Dressler, Ann. Missouri Bot. Gard. 48

(1961) 339.

Euphorbia geniculata Ortega, Nov. Pl. Descr. Dec. (1797) 18.

Euphorbia heterophylla L. var. geniculata (Ortega) Gomez, Anales Hist.

Nat. 23 (1894) 46.

102 Gard. Bull. Singapore 51 (1999)

Euphorbia prunifolia Jacq., Pl. Hort. Schoenbr. 3 (1798) 115; Backer &

Bakh. f., Fl. Java 1 (1963) 502; Mohamed Soerjani et al., Weeds of Rice in

Indonesia (1987) 288.

Specimens examined: PAHANG, Pulau Tioman, near Kampung Ayer

Hantu, L.H. Chua & K.E. Tian KL65, 5 November 1982 (SINU); Pulau

Tioman, near Kampung Tekek, F.J. Marret 12, 12 September 1985 (SINU).

SINGAPORE, Cluny Road, Md Nur SFN 37778, 28 May 1946 (SING, K).

Acknowledgements

Curators of KEP, KLU, SING and SINU very kindly obliged when I

requested access to material in their herbaria.

References

Airy Shaw, H.K. 1977. Additions and corrections to the Euphorbiaceae of

Siam. Kew Bulletin. 32: 69-83.

Burger, W.; 4c Huit,. MM. 1995: Flora Costaricensis Family #113

Euphorbiaceae. Fieldiana, Botany. 36: 1-169.

Chriztenhusz, M.J.M. 1998. Key for the Philippine species of Euphorbia.

Malesian Euphorbiaceae Newsletter. 9: 2-4. .

Lin, S.-C., & Hsieh, C.-F. 1993. Euphorbia. Flora of Taiwan, second edition.

3: 456-469.

Ma, J.S., & Wu, C.Y. 1992. A synopsis of Chinese Euphorbia L. s.l.

(Euphorbiaceae). Collectanea Botanica. 21: 97-120.

Philcox, D. 1997. Euphorbiaceae. Revised Handbook to the Flora of Ceylon.

11: 80-283.

Ridley, H.N. 1924. Euphorbia. Flora of the Malay Peninsula. 3: 80-82.

Gardens’ Bulletin Singapore 51 (1999) 103-118.

First Record of a Natural Begonia Hybrid in Malaysia

LAI-LAI TEO' AND RUTH KIEW?

‘School of Science, National Institute of Education (NTU), Singapore 25956

*Singapore Botanic Gardens, Cluny Road, Singapore 259569

Abstract

Study of three begonia populations in montane forest at Cameron Highlands, Pahang,

confirmed that hybridization had occurred between Begonia decora and B. venusta. The

hybrids are fertile and calculation of the hybrid index indicates that introgression had taken

place. Two populations are stable and are represented by hybrid swarms. Plants of the two

parents were not located in the vicinity of these two populations. The third, comprising

both parents and hybrids, is unstable and its composition has changed over the years. A

further two populations have been destroyed by habitat disturbance, which is prevalent at

Cameron Highlands.

Introduction

While hybrids of begonia are the norm in cultivation with over 10,000

hybrids and cultivars recorded (Mabberley, 1997), in contrast they are

rarely encountered under natural conditions. In Peninsular Malaysia, 53

native species are known (Kiew, in prep.), but only one putative hybrid

has been encountered. This was discovered at Cameron Highlands (Fig. 1)

in 1983 (Ruth Kiew RK1278) and became the basis of a more detailed

study, the results of which are reported below. The aim was to reassess

diagnostic characters that distinguish the two putative parents, Begonia

decora Stapf and B. venusta Ridl., to collect data from wild populations to

test whether hybrid plants indeed existed, and to assess the fertility of

hybrid plants.

Begonia decora was exhibited at the Royal Horticultural Society in

London in 1892, where it was described as “a pretty dwarf-growing plant,

with exceedingly ornamental foliage; the leaves are velvety-purple with

greenish-yellow veins, and very hirsute” and where it was awarded a First-

class Certificate (Anon., 1892). Towards the end of the nineteenth century,

it was used in hybridization with B. rex Putzeys to impart a wide range of

reds and a metallic sheen to the leaves (Thompson, M.L. and E.J.

Thompson, 1981). In contrast, B. venusta (Fig. 2a), which has the largest

flowers of any Peninsular Malaysian species, has yet to be introduced into

cultivation. It is a larger, more robust plant with glossy, fleshy leaves that

have earned it the common name of the cabbage-leaved begonia.

104 Gard. Bull. Singapore 51 (1999)

MT. 4

BERINCHANG

MT. JASAR

CAMERON HIGHLANDS

KUANTAN

KUALA

LUMPUR

MALAY

PENINSULA

00"E 105°E

Figure 1. The study sites of the hybrid populations of begonia at Cameron Highlands, Malaysia.

Natural Begonia Hybrid 105

Both species belong to the same group of species with markedly

oblique leaves within the Platycentrum section, which includes creeping,

rhizomatous species, which produce male flowers with four tepals and

fruits with two locules and three wings, one of which is enlarged.

Plants in the first hybrid population discovered (Fig. 2b) more closely

resembled plants of B. venusta (Fig. 2a) in having large cabbagey leaves

but they had scattered hairs and were slightly purplish beneath, characters

not seen in B. venusta. Other populations included hybrid plants (Fig. 2c)

that more closely resembled B. decora (Fig. 2d) in having smaller leaves,

which were more hairy and reddish-purple beneath, but not as small, densely

hairy and deeply coloured as B. decora itself (Fig. 2d).

Figure 2. Laminas of the begonia hybrids and parent species.

(a. Begonia venusta; b. venusta-like hybrid; c. decora-like hybrid; d. B. decora)

Materials and Methods

Morphological study

Herbarium material (including type specimens) of B. decora and B. venusta

was examined to draw up a list of clear and unambiguous diagnostic

characters that would distinguish these two parent species. These characters

were tested in the field to ensure that the complete range of variation was

106 Gard. Bull. Singapore 51 (1999) |

captured (Table 1). No hybrids were found among the herbarium collections.

Descriptions of the parent species are given in Appendix 1.

Table 1. Diagnostic characters used to differentiate Begonia decora from B.

venusta.

Character B. decora B. venusta

Plant height (cm) <20 28-55

Rhizome diam. (mm) <6 8-16

Petiole length (cm) <i >22

Petiole indumentum hairy glabrous

Ratio lamina width:length (mean) 1:1.6-1.8 1:1.2-1.5

Lamina undersurface colour red green

Lamina indumentum hairy glabrous

Lamina base overlapping not overlapping |

Lamina margin serrulate denticulate

Fruit wing broad and blunt tapered

Fruit wing texture thin thick and fibrous

Study sites

The hybrid is known only from Cameron Highlands, Pahang, a popular

highland tourist resort. Search was made along accessible forest paths and

along roads in the montane forest, which resulted in the discovery of six

populations. Unfortunately, two of these have since been destroyed (by

building construction and farming) before the more detailed study started

in 1996. |

All plants with both flowers and fruits in the remaining four

populations (Table 2) were scored for the hybrid index using the eleven

characters in Table 1.

Table 2. Number of individuals scored for the hybrid index in begonia

populations at Cameron Highlands

(Gunung=Mount)

Study sites B. decora hybrid B. venusta

Gunung Beremban NP if NP

Gunung Berinchang 11 30 4]

Gunung Jasar NP 20 NP

°99 Acre’ Plantation 2, NP NP

(NP=not present)

Natural Begonia Hybrid 107

In the ideal situation, populations comprising only one parent should

be scored. However, only one small population of B. decora could be

found, while none of just B. venusta was located. Sample size was also

limited by small population size and the fact that most plants were either

sterile or had flowers or fruits (not both).

The sites at Gunung Beremban, G. Berinchang and G. Jasar were in

upper montane forest in damp and deeply shaded conditions beneath a

closed canopy although the canopy was slightly more open above streams.

The G. Beremban site was relatively flat, while the other two were sited on

steep slopes. The site at 99 Acre’ Plantation was a forest fragment beside

the road close to farms and was a more exposed steep slope, which was,

however, shaded by overhanging trees.

Hybrid index

This was calculated based on the characters listed in Table 1. The presence

of each character state for B. decora was scored as ‘2’, those for B. venusta

‘0’, while an intermediate character state was scored as ‘1’. The score for

each plant was then totalled and results for each population expressed in a

histogram (Anderson, 1949). Thus, in theory plants of B. decora should

obtain a total of ‘22’ for the eleven characters scored, B. venusta ‘0’, while

hybrid plants are identified by intermediate scores.

Anatomical study

Since differences in leaf texture and indumentum were so pronounced, the

anatomy of the lamina of fresh leaves was also examined. Permanent slides

of lamina T.S. (30 samples each for the two parents and the hybrid) were

made by standard methods (Sass, 1958) being sectioned 15 um thick, stained

in 1% safranin and 0.5% fast green and mounted in permount.

Indumentum was examined after clearing the lamina (one sample

for each of the two parents and ten for the hybrid) with sodium hydroxide

and chloral hydrate and mounting in permount. The number of trichomes

visible in the optical field viewed at x40 magnification at eight points on

each lamina was recorded and the mean calculated.

Pollen and seed viability

To assess pollen viability, five samples each of about 150 fresh pollen

grains for each of the two parents and the hybrid were stained with

methylene blue and made into a temporary mount in glycerol. The

percentage viability was then calculated by counting the number of stained

108 Gard. Bull. Singapore 51 (1999)

(viable) and unstained (sterile) pollen grains for samples of B. decora, B.

venusta and the hybrid.

To test seed viability, 50 seeds of both parents and six hybrid plants

were sown on damp filter paper and the number of germinated seeds

recorded at 2-day intervals until there was no further germination.

Results

Morphology

Compared with the character states of the parent species (Table 1), the

hybrid plants displayed a complete range of morphological intermediacy in

all qualitative (Fig. 2) and quantitative (Table 3) characters.

Table 3. Range in quantitative characters displayed by hybrid plants

Character hybrid plants

Plant height (cm) 8-45

Rhizome diameter (mm) 4-14

Petiole length (cm) 9-41

Ratio lamina width:length (mean) | 1:1.1-1.8

Hybrid index

Calculation of the hybrid index gave a score for B. decora of 22 when it

grew alone (Fig 3d), and between 19 and 22 when it grew together with the

hybrid (Fig. 3a). No population comprised only plants of B. venusta and on

G. Berinchang, where it grew with hybrid plants, the distinction between it

and the venusta-like hybrid was not disjunct.

Scores for the hybrid ranged widely. For the G. Beremban population

(Fig. 3b), it ranged from 5 to 13 with a mean of 8 and a mode of 8-9, and

for the G. Jasar population (Fig. 3c), it ranged from 7 to 17 with a mean of

14 and mode of 16. The structure of the G. Berinchang population (Fig 3a)

was more complex. There was no clear distinction between scores for

plants of B. venusta and the hybrid, and hybrid scores ranged up to 16.

Anatomy

The hybrid also showed intermediacy in characters such as lamina thickness,

Natural Begonia Hybrid 109

a. Hybrid Index for Berinchang Population

no of plant

b. Hybrid Index for Berembum Population

no of plant

[e>)

fai

11 12) ts 14 15 16 Wey WIS AS Ais OZ ny «ee

c. Hybrid Index for Jasar Population

no of plant

0 1 2 3 4 5 6 7 8 9 10 11 Wad) Se: 14 #15 16 ewer eeon TeORM2T. «22

d. Hybrid Index for 99-Acre Plantation

—

[o)

no of plant

Figure 3. Hybrid index for begonia populations.

(a. Gunung Berinchang population; b. G. Beremban population; c. G. Jasar population; d. ’99

Acre’ Plantation population)

110 Gard. Bull. Singapore 51 (1999)

which was reflected by differences in the thickness of the tissue layers and

number of mesophyll layers (Table 4, Plate 1).

—_——“

00 MICRON

Plate 1. T.S. lamina of begonia species and hybrid.

(a. Begonia decora; b. hybrid; c. B. venusta)

Natural Begonia Hybrid ea!

Table 4. Differences in the lamina anatomy of the hybrid and its parents.

(meantstandard deviation)

Character B. decora hybrid B. venusta

Thickness (um)

Lamina 166425 234435 312+16

Upper epidermis 48+9 5 iese) 83414

Palisade mesophyll 29a5 40+7 5248

Spongy mesophyll 2948 43+14 215

Lower epidermis 3847 49+7 64+13

No. spongy mesophyll layers 1-2 2-3 3-4

The hairiness of B. decora 1s very pronounced. Not only are the hairs

dense but they are papillose (raised on a protrusion of the lamina) so that

the underneath of the lamina appears pitted (Fig. 4). This is in complete

contrast to B. venusta, which has glossy leaves that are completely smooth

and glabrous. When counted per x40 optical field, the lamina of B. decora

had (31-)35(-43) trichomes, B. venusta 0, ‘venusta-like hybrid plants (0-)

2(—5) and ‘decora-like’ hybrid plants (20—) 27(—36) trichomes.

Figure 4. T.S. lamina of Begonia decora.

(P — papillose hair base)

112 Gard. Bull. Singapore 51 (1999)

Pollen and seed viability

The pollen viability for both parent and hybrid plants was high: (87.5—) 96

(-100)% for B. decora, (91.9—)97.4(-100) % for B. venusta and (91.9—)97.4(—

100)% for hybrid plants.

Seed germination for all taxa was rapid (Fig. 5) beginning slightly

earlier in seeds of the hybrid at Day 6 compared with Day 8 for seeds of B.

decora and B. venusta, and while no further germination occurred after

Day 14 for the hybrid, seeds of the two parental species continued to

germinate up to Day 20. Percentage germination was high in all three taxa

- 100% for B. venusta and 98% for B. decora and the hybrids.

% of germination

Figure 5. Rate of seed germination in Begonia decora, B. venusta and hybrids.

Discussion

The data from the hybrid index and anatomical study support the view

that B. decora and B. venusta do produce hybrids in the wild and that the

hybrid plants are fertile and backcross with the parental species to produce

hybrid swarms. Arnold (1997), in reviewing cases of natural plant hybrids,

concluded that, for the majority, the hybrid exhibits equivalent or superior

fitness to the parents.

However, the structure of hybrid swarms is different at each site. On

G. Berinchang, hybrid plants show a wide range of intermediate scores

Natural Begonia Hybrid 113

between the two parents, while in the populations on G. Beremban and G.

Jasar, the scores for majority of hybrid plants are skewed towards B. venusta

and B. decora, respectively.

The high viability of seed from hybrid plants shows that there are no

genetic barriers between these two closely related species. Thus, internal

factors cannot play a role in isolating the two species.

In addition, flower characteristics and pollinator behaviour also do

not serve to keep the two species reproductively isolated. The tepals of

both species are similar in shape (broadly oval), size, colour (white or pale

pink) and open widely to expose a similar cluster of yellow anthers in the

male flower or twisted yellow stigmas in the female flower. Preliminary

observations on the populations at G. Beremban and G. Berinchang

indicated that a stingless bee, Trigona sp., is the most likely pollinator as it

visits the male flowers of both species and hybrid plants to collect pollen.

(No other flower visitors were observed). This bee also briefly visits the

female flowers (so briefly as to appear to ‘bump into’ the flower before

flying off). The female flowers of begonia do not offer any reward and

pollination is by deceit, the bee visiting the female flower in mistake for

the closely similar male flower. Agren and Schemske (1991) have shown

that in Costa Rica pollination by deceit in Begonia involucrata Liebmann

is also by a Trigona species and is very effective. Although female flowers

were visited infrequently and briefly (male flower were visited three times

as frequently and for ten times longer), three quarters of the ovaries set

fruit with about 50% seed set. In begonia species where a single

inflorescence is produced, the male and female phases do not overlap (B.

decora and B. venusta are protogynous; B. involucrata is protandrous),

pollen will therefore be carried from one plant to another (xenogamous

pollination). Therefore, when B. decora and B. venusta occur within the

flight range of a Trigona bee, it is likely that crossing between the two

species will occur.

This then raises the question as to what the isolating mechanisms are

that enable the two species to remain distinct at Cameron Highlands.

Begonia decora has a wider altitudinal range than B. venusta and at lower

altitudes, e.g., at ‘99 Acre’ Plantation, it occurs alone. B. decora also appears

to tolerate more exposed conditions at it can grow on steep roadside banks

at the edge of forest. Both species grow on damp, shaded steep slopes

within the forest. The venusta-like hybrid shows some propensity to become

weedy and colonises sandy banks or stream bottoms that frequently develop

due to siltation of streams when earth has been exposed by man’s activities.

In these circumstances, it can form pure stands. This is interesting as in

temperate floras, the occurrence of hybrids is often associated with man’s

disturbance of natural habitats, the greater variability of hybrid plants

114 Gard. Bull. Singapore 51 (1999)

compared with either parent is thought to contribute to their success in

invading unstable habitats (Anderson, 1949).

However, the exact ecological requirements of the parental species

and hybrid plants have yet to be tested experimentally because of the

difficulty of growing montane plants in the hotter lowlands. It therefore

cannot yet be proved that ecological factors are the major isolating factors

that maintain the parental species as distinct entities. Elsewhere, for example

on limestone, several species of begonia occur sympatrically and yet hybrids

are not found. However, on limestone, the distinction between microhabitats

is much clearer (Kiew, 1998).

The hybrid population on G. Berinchang appears to comprise both

F1 plants that score between 9 and 11, as well as the progeny of backcrossing

with scores ranging between 3 and 16. The wider scores for B. decora (19-

22) and B. venusta (0-2) than the theoretical 22 and 0 and the intergrading

of scores for B. venusta and the venusta-like hybrid (3-7) can be interpreted

as earlier hybridization and introgression that have caused the parent species

to be genetically less pure. Observation of this same population after three

years showed the number of plants of the hybrid and B. decora were

drastically reduced indicating that the composition of this population is

not stable. (That the previous year, 1998, was a particularly dry year due to

the El Nino effect may have been a contributing factor in this change.)

Long-term studies in temperate floras indicate that in many cases the

composition of hybrid populations is not stable (Stace, 1989).

The habitats beside the forest paths on G. Beremban and G. Jasar

are deep inside the forest and have experienced little disturbance other

than trampling along the narrow walking trail. These populations are

interesting as plants of the parental species could not be located in the

1996 study, although plants of B. decora were growing nearby the study

site on G. Jasar in 1983 (Ruth Kiew RK1276). It appears that in these two

populations hybridization is of longstanding and that introgressive

hybridization has occurred so that now plants of pure B. decora or B.

venusta no longer exist. It also indicates that the hybrids swarm with its

wide genetic variability may have superior fitness in this habitat compared

to the parents (Arnold, 1997). It is likely that the difference in these two

populations was due the numerical predominance of one of the two parent

species, B. decora on G. Jasar and B. venusta on G. Beremban.

In temperate floras, some authors (e.g., Stace, 1989) considered

that ‘in general the ability to hybridize is the usual situation’ and he cites

780 hybrids among the 2500 species in the British flora. Records of hybrids

in tropical floras are much rarer and quantitative studies are almost non-

existent. Two other genera, Nepenthes (Nepenthaceae) and Rhododendron

(Ericaceae), are well-know as forming hybrids readily, both under natural

Natural Begonia Hybrid HES

conditions or in cultivation because, like begonia, they have no internal

barriers to crossing. In Nepenthes, there are 280 hybrids recorded in

cultivation involving 34 of the 82 species known (Jebb and Cheek, 1997).

In Peninsular Malaysia, where 11 species occur, two hybrids are so well

known that they have been named (x hookeriana and x trichocarpa) and

several other informal hybrids are known to exist both among the lowland

as well as the montane species. Among the Vireya rhododendrons on G.

Kinabalu, Argent et al. (1988) have observed several hybrids and note that

these form hybrid swarms in inherently unstable or in disturbed habitats

where they may be common and abundant.

However, populations of hybrids of Nepenthes and Rhododendron

have not been subjected to detailed analysis. Only Globba hybrids have

been studied in detail when Lim (1973) studied three populations at

Cameron Highlands of hybrids between Globba patens Migq. and G. cernua

Baker (Zingiberaceae). The hybrid index indicated that introgressive

hybridization had occurred in the three populations and, similar to our

study, her G. Jasar population no longer included plants of one parent (G.

patens), although it was known earlier (in 1925) from that area.

Lim’s results are interesting as they too show that changes have

taken place in the montane flora at Cameron Highlands in the last 70

years, the period when it developed to become the largest highland resort

in Malaysia. Leong (1992) chronicled the forest clearance, which started

on a large scale in the 1930s when tea plantations and farms for temperate

vegetables were established, to the 1970s when large scale resort

development was encouraged and new areas were opened up to grow cut

flowers, an industry that by 1990 had surpassed the value of the vegetable

harvest. These developments have been at the expense of forest, which has

resulted not only in the obvious effect of destroying habitats but also has

had a more insidious effect in changing the environment. Leong reported

that by 1985 the temperature at Tanah Rata had risen by 4°C, which would

in turn affect other environmental factors, such as cloud formation, rainfall

and humidity.

Conclusion

Analysis of morphological intermediacy to produce a hybrid index is a

useful tool in confirming the occurrence of hybrids between B. decora and

B. venusta. Intermediacy in anatomical characters also supports their hybrid

status. That five hybrid populations have been found indicates that

hybridization will occur wherever these two species occur sympatrically.

The fertility of the hybrid plants enables introgression to take place resulting

in hybrid swarms.

116 Gard. Bull. Singapore 51 (1999)

Disturbance by man on the one hand appears to favour the spread of

the venusta-like hybrid, which grows lushly on wet sand resulting from soil

erosion, while on the other hand to threaten these populations with habitat

destruction as witnessed by two of the five populations having been cleared

since they were discovered in 1983.

Acknowledgements

We thank the curators of the herbaria at BM, K, KEP and SING for

permission to examine specimens in their care, to Mr S. Anthonysamy and

Miss S.M. Tam for help in field work, to Miss S. Madhavan for technical

advice on plant microtechnique and to IRPA Research Grant 52858 from

the Ministry of Science, Technology and the Environment, Malaysia, for

support and to Universiti Putra Malaysia for facilities to carry out this

research.

References

Agren, J. and D.W. Schemske. 1991. Pollination by deceit in a neotropical

monoecious herb, Begonia involucrata. Biotropica. 23: 235-241.

Anderson, E. 1949. Introgressive Hybridization. Wiley, New York, USA.

Anonymous, 1892. Begonia decora. Gardeners’ Chronicle. 5: 621.

Argent, G., A. Lamb, A. Phillipps and S. Collenette. 1988. Rhododendrons

of Sabah. Sabah Parks Publication, No. 8. Kota Kinabalu, Malaysia.

Arnold, M.L. 1997. Natural Hybridization and Evolution. Oxford University

Press, England.

Jebb, M. and M. Cheek. 1997. A skeletal revision of Nepenthes

(Nepenthaceae). Blumea. 42: 1-106.

Kiew, R. 1998. Niche partitioning in limestone begonias in Sabah, Borneo,

including two new species. Gardens’ Bulletin Singapore. 50: 161-169.

Kiew, R. (in prep.) The Begonias of Peninsular Malaysia. Natural History

Publications, Malaysia.

Leong, Y. K. 1992. Conservation and development of Cameron Highlands.

In: R.P. Lim and S.W. Lee (eds.). Hill Development. Malayan Nature

Society, Kuala Lumpur, Malaysia. pp. 20-32.

Natural Begonia Hybrid Ly

Lim, S. 1973. Cytogenetics and taxonomy of the genus Globba L.

(Zingiberaceae) in Malaya. V. Introgressive hybridization in hexaploids.

Journal of Linnean Society, Botany. 66: 143-156.

Mabberley, D.J. 1997. The Plant Book. Cambridge University Press, U.K.

2 ed:

Sass, J.F. 1958. Botanical Microtechnique. lowa, Iowa State College Press,

USA.

Stace, C.A. 1989. Plant Taxonomy and Biosystematics. Edward Arnold,

Le Te 22 ed:

Thompson, M.L. and E. J. Thompson. 1981. Begonias, the Complete

Reference Guide. New York Times Books, New York, USA.

Appendix 1: Description of the species

Begonia decora Stapf

Creeping plant with horizontal, slender reddish rhizomes, up to 6 mm diam. with internodes

5-7 mm long, foliage reaching up to 20 cm above ground. Stipules reddish, persistent,

slightly hairy, narrowly lanceolate, c. 12-13 x 4-6 mm, apex acute. Petiole reddish, 7-17 cm

long, slightly or densely hispid. Lamina malachite green or green-bronze and lighter green

along veins on upper surface, rosy-purple or magenta and lighter red along veins on lower

surface, obliquely ovate, 9-15 x 5.5—8.5 cm, base unequal, broadly rounded and overlapping,

margin minutely serrulate, apex acuminate to cuspidate, densely hirsute above, hairs curved,

magenta on raised whitish conical papilla, veins 7-9 radiating from junction with petiole,

densely hispid and prominent beneath. Inflorescence axillary, few-flowered, long-stalked

cyme, projecting above leaves, 1-3 flowers open simultaneously. Peduncle and pedicels

reddish green, peduncle 9-21 cm long, hispid. Bracts reddish, c. 19 x 14 mm, at base broadly

ovate, midway abruptly lanceolate narrowly to acute apex, caducous. Male flower: pedicel

c. 2 cm long, tepals 4, white tinged pink or pale pink, outer two oval with apex acute, 27-32

x c. 18 mm, inner two narrowly oval, c. 20 x c. 9 mm; stamens many, clustered, filament

short, anther golden yellow, c. 2 mm long. Female flower: pedicel 13-15 mm long, tepals 5,

pinkish white to rosy pink, broadly rounded, isomorphic, outermost c. 24 x 16 mm, innermost

c. 23 x 14 mm, ovary pale reddish-green, 3-winged, one wing wider, stigma pale yellow,

lobes twisted. Infructescent peduncle elongating to 24 cm long. Capsule pendant hanging

on a fine thread-like pedicel, 2-loculate, 10-12 mm long, two lateral wings rounded, 8-10

mm wide; posterior wing thin, broad with blunt apex, 17-26 mm wide.

Begonia venusta King

Creeping plant with slender rhizome 8-16 mm diam., at first with crowded internodes, then

internodes elongating to 9-17 cm long to produce a weakly erect plant 28-55 cm tall,

rooting at nodes, completely glabrous except for the bracts. Stipules persistent, lanceolate,

c. 20 x 10 mm, apex pointed. Petiole slightly reddish, 22-45 cm long. Lamina plain dark

118 Gard. Bull. Singapore 51 (1999)

green and glossy above, paler green beneath, fleshily leathery, obliquely ovate, 12-26 x 10-

17 cm, base unequally rounded not over-lapping, margin minutely and distantly denticulate,

apex acuminate, veins 7-9 radiating from junction with petiole, prominent beneath.

Inflorescence axillary, long-stalked cyme, at the level of or below the leaves, 14 flowers

open simultaneously, protandrous. Peduncle in male phase 10-26 cm long, elongating in

female phase to 45 cm long. Bracts in pairs, persistent, broadly lanceolate, 18-25 x c. 6 mm,

apex apiculate, sparsely hispid outside with trichomes c. 1 mm long. Male flower: pedicel

robust 2—2.5 cm long, tepals 4, white or white tinged pink, outer two broadly oval with

rounded apex, 18-33 x 17-26 mm, inner two oval with rounded apex, 16-30 x 7-10 mm,

stamens many, clustered, filament and anther yellow, filament 0.5-1 mm long, anther

narrowly oblong, 1—1.25 mm long. Female flower: pedicel 1-2 cm long, tepals 5, pinkish

white, broadly oval, isomorphic, outermost c. 18 x 14 mm, innermost c. 15 x 6 mm, ovary 3-

winged with one wing wider, style c. 2 mm long, stigma yellow, twisted lobes 6 mm long.

Peduncle of infructescence c. 29.5 cm long. Capsule c. 15-16 mm long, pendant on stiff

pedicel c. 3 cm long, two lateral wings ovate, c. 8 mm wide, posterior wing fibrous, oblong,

19-20 mm wide, tapered to rounded apex.

Gardens’ Bulletin Singapore 51 (1999) 119-124.

Eupatorium catarium, a New Name for Eupatorium

clematideum Griseb., non Sch.Bip. (Compositae), a South

American Species Naturalized and Spreading in SE Asia

and Queensland, Australia

J.F. VELDKAMP

Rijksherbarium/Hortus Botanicus,

P.O. Box 9514, NL-3200 RA Leiden

The Netherlands

Abstract

Eupatorium catarium Veldk. (Compositae) is a new name for Eupatorium clematideum

Griseb., non Sch.Bip., also known as Praxelis clematidea R.M. King & H. Rob.. The species

has been introduced in S. China and Queensland, Australia, where it appears to be spreading

rapidly.

Introduction

During a visit in 1998 to Hong Kong, Macao, and Queensland, I was

shown a recent introduction, Eupatorium clematideum Griseb.

(Compositae). This species is spreading rapidly there and now occurs in

mainland S China, at least in the Shenzhen region in Guangdong as well.

It has been regarded as a member of Eupatorium Tourn. section

Praxelis (Cass.) Baker, e.g. by Aristeguieta (1974), Cabrera and Vittet

(1954), Cabrera (1974), Cabrera et al. (1996) and Liogier (1996).

King and Robinson (1970, 1987) have split Eupatorium Tourn. into a

number of genera, of which Praxelis Cass. is one, and have called the

species Praxelis clematidea R.M. King & H. Rob.. Their elevation of

infrageneric groups in Eupatorium to genera is still under debate and is

not generally accepted.

Eupatorium clematideum was described by Grisebach (1879) based

on Argentine and Paraguay material, but this is an illegitimate name as

prior to that, in 1866, Schultz Bipontinus had made the same combination

for a Nepalese taxon: “Mikania clematidea DC.! pr. V. 191 n. 32 ist nach

einem von mir im britisch Museum untersuchten Originalexemplar ein

Eupatorium (clematideum Sz-Bip.)”. This specimen is a Wallich collection

(no. 3173; IDC microfiche 7394). The combination was reduced by Uniyal

(1995) to E. chinense L.. However, S.-W. Chung and C.-I Peng (1998) have

accepted E. clematideum Sch. Bip. as a distinct species with two varieties

in Taiwan.

120 Gard. Bull. Singapore 51 (1999)

Jackson (1895) mentioned another earlier Eupatorium clematideum

Less. cited as having been published by Baker (1876), but the latter had

included it as a synonymous manuscript name for Mikania phaeoclados

Mart. ex Baker and so it was invalidly published.

The combination made by Schultz Bipontinus makes that of Grisebach

a later homonym. Consequently, Praxelis clematidea R.M. King & H. Rob.

(1970) is not a ‘comb. nov.’, but a ‘nom. nov.’ [see ICBN Art. 58.1.(b) and

58.3] and correct in Praxelis. In Eupatorium, however, the species has no

available epithet, and I propose to call it E. catarium Veldk. The epithet

refers to the fairly disagreeable smell of the plant (‘of cats’).

As descriptions of Eupatorium sect. Praxelis and Eupatorium

clematideum sensu Griseb. and Praxelis clematidea King & H. Rob. are not

readily available in SE Asia and Australia, I give them here, together with

the synonymy.

Eupatorium L. sect. Praxelis (Cass.) Benth. ex Baker in Mart., Fl. Bras. 6,

2 (1876) 341. — Praxelis Cass. in Cuvier, Dict. Sc. Nat. 43 (1826) 261.

— Type: Praxelis villosa Cass. [= Praxelis diffusa (Rich.) Pruski =

Eupatorium pauciflorum Kunth; non E. diffusum Vahl].

Ooclinium DC., Prod. 5 (1836) 133. — Lectotype: Ooclinium grandiflorum

DC. [= Praxelis grandiflora (DC.) Sch.Bip.| [designated by R.M. King

& H. Robinson, Sida 3 (1969) 338].

Annual or perennial herbs or undershrubs. Capitules long-pedicelled,

solitary to few in a lax panicle. Phyllaries 15-25, in 3-5 indistinct series,

unequal, deciduous before the fruits, 3-6-nerved. Receptacle conical to

ellipsoid, glabrous. Corolla of marginal flowers without expanded outer

lobes. Anther collar [see Robinson & King (1977) t. 2] with the cell walls

with prominent ornate banding, below with many quadrate cells with

oblique or vertical banding, above with elongated cells with transverse

bands, exothecal cells usually as long as wide [see Robinson & King (1977)

t. 5], anther appendage large, serrate to lobed [see Robinson & King (1977)

t. 11]. Style without basal node. Achenes more or less flattened, 3- or 4-

costate; carpopodium distinct, highly asymmetric, cells clear, quadrate or

elongated, walls thin, firm. Pappus of 20-40 long, scabrous bristles, apical

cells pointed.

Distribution: 15 species in S. America [fide Bremer et al. (1994)]: Argentine,

Bolivia, Brazil, Colombia, Guyana, Paraguay, Peru, Venezuela; 1 recently

introduced in S China (e.g. Hong Kong, Macao, Shenzhen region in

Guangdong) and N Australia (Queensland).

Eupatorium catarium 121

Eupatorium catarium Veldk., nom. nov.

Eupatorium clematideum Griseb., Abh. Konigl. Ges. Wiss. Gottingen 24

(1879) 172, non Sch.Bip. (1866). — Syntypes: Balansa 936 (GOET?,

P?, n.v.), Lorentz s.n. ‘Cordoba’ (GOET?, n.v.). — Eupatorium

urticifolium L.f. (1781, non Reichard, 1780) var. clematideum Hieron.

ex Kuntze, Rev. Gen. 3 (1898) 148, comb. incorr. — Praxelis clematidea

R.M. King & H. Rob., Phytologia 20 (1970) 194, nom. nov.

Eupatorium urticifolium L.f. (non Reichard) var. nanum Hieron., Bot. Jb.

22 (1897) 783 (‘nana’), comb. incorr. — Type: Kuntze Sept. 1892 (B,

holo, extant?; NY?), S. Paraguay, Villa Florida.

Eupatorium pauciflorum auct. non Kunth.

Eupatorium urticifolium auct. non Reichard (1780) nec L.f. (1781).

Perennial (already flowering in its first season), 0.3—1 m tall, caudex woody,

erect, branched, branches terete, angular, septate hirsute, laxly foliate,

malodorous. Internodes 3-16 cm long. Petioles 0.3—2 cm long, blades ovate

to rhomboid, 2.5-6 by 1-4 cm, septate hirsute and gland-dotted on both

sides especially underneath, base rounded to cuneate, 3-nerved, margins

coarsely dentate, teeth 5-8 on each side, acute, apex acute. Inflorescence

of many capitules in terminal, dense, corymbiform, few-headed cymes.

Pedicels septate hirsute, 2-10 mm long. Involucre cylindrical campanulate,

7-10 by 4-5 mm. Phyllaries in 4 or 5 series, yellowish with 3-5 green

nerves, distally purplish, glabrous to distally appressed strigose, the outer

ones smallest, lanceolate, apex acuminate, the inner linear, acute. Flowers

25-30, somewhat purplish blue or lilac; corolla 3.5—4.8 mm long, shortly (4-

or) 5-dentate. Achenes 2—3 mm long, black, distally hispidulous; pappus

bristles 15—40, white. n = 311.

Distribution: N Argentina, Bolivia, S Brazil, Paraguay, Peru. Introduced

into Hong Kong around 1980, fide Corlett and Shaw (1995), Macao,

Shenzhen region in Guangdong, and no doubt elsewhere in S China

(Waterhouse and Corlett, 1996; not mentioned by Shi, 1985). The first

records in Australia date from November 1993 near S Johnstone,

Queensland. In 1994 it was found to be abundant and widespread in the

Tully and Bingil Bay Districts of N Queensland, where it may have been

introduced at least ten years earlier (perhaps already in the late 1960s) as

seeds from S. America. It now occurs from Townsville to the Daintree

region, and on the Atherton Tablelands.

Habitat: Roadsides, railway lines, disturbed areas, urban wasteland, beneath

fences, rural paths; altitude 0-700 m in Hong Kong, 0-800 m in Queensland,

(0—)350-3050 m in S America.

122 Gard. Bull. Singapore 51 (1999)

Ecology: In Australia, in 1998 it occurred in areas with a mean annual

rainfall of 945-4000 mm, and mean maximum January (summer)

temperatures between 28.1°C and 31.3°C, and mean minimum July (winter)

temperatures of 9.6°C to 17.0°C. There are occasional night frosts on the

Atherton Tablelands.

In Hong Kong, the mean annual rainfall is 1500-2500 mm, with mean

monthly temperatures at sea level varying between 15.8°C for January and

28.8°C for July. On the Peak of Hong Kong where night frosts occur, a

specimen with a well-developed woody caudex was collected (Veldkamp

8772, L) indicating it had been growing there for some time. Waterhouse

and Corlett (1996) reported that minimum temperatures of 3-4°C causes

minor damage to the flowers, but not to the leaves.

There seem to be no such observations for S America, but in Bolivia

it occurs up to 3050 m altitude suggesting frost-resistance.

Specimens collected at sea level on 18 September 1998 in Macao

(Veldkamp 8793, L) were dried and after arrival some days later in Leiden

were frozen to -30°C for about 48 hours. On 25 October about a dozen

seeds placed in a small container in my office germinated within two days,

indicating that they are quite frost and drought resistant. I put a plant

outside in March/April, 1999. There followed occasional night frosts with

some snow, and daily temperatures never reached 10°C. It grew slowly

and apparently was only slightly affected by these low temperatures; it was

more stocky and darker and the leaves were smaller than the rather etiolated

ones kept inside.

Extermination: Reported to be more resistant to herbicides than Ageratum

conyzoides L. because of its more robust rootstock and longer growing

and flowering seasons. |

Toxicity: Anecdotal reports suggest that it may be poisonous to livestock

and humans if ingested.

Notes: Eupatorium catarium, according to B.L. Robinson (1920) and which

he called E. clematideum: “is very closely related to E. pauciflorum Kunth

(‘H.B.K.’), but is considerably stouter and has much shorter pedicels and

in consequence decidedly denser cymes giving it a characteristic habit. It

seems best to accord it specific rank”.

In China and Australia, E. catarium has been mistaken for an odd

Ageratum conyzoides, which differs in its flat to convex receptacle, persistent

phyllaries in distinct rows, and 60-75, white or violet flowers per capitule.

Corlett and Shaw (1995) erroneously reported the presence of 50-65 flowers

per capitule in Hong Kong plants.

Eupatorium catarium f23

Acknowledgements

I want to thank Dr. R.T. Corlett, Hong Kong, for showing me around the

island, Ms. C.M. Kooij, Hong Kong, for taking me on a trip to Macao, Ms.

B.M. Waterhouse, Mareeba (Queensland), for allowing me to cite from

her manuscript and Corlett for discussions on the history and ecology of

the species in Hong Kong and Australia, and the anonymous referees of

this paper, who suggested crucial changes for the better.

References

Aristeguieta, L. 1974. Compositae. In: T. Lasser, Flora de Venezuela. 5, 1:

120, 180-184. Instituto Botanico, Caracas.

Baker, J.G. 1876. Compositae. In: C.F.P. von Martius, Flora Brasiliensis.

6, 2: 256. Mus. C.R. Pal. Vind.; Munchen.

Bremer, K., A.A. Anderberg, P.O. Karis & J. Lundberg. 1994. Tribe

Eupatorieae. In: K. Bremer (ed.), Asteraceae: Cladistics and Classification.

p. 656. Timber Press, Portland (OR).

Cabrera, A.L. 1974. Compositae. In: A. Burkart (ed.), Flora ilustrada de

Entre Rios. 6: 175, t. 85, f. i—l. INTA, Buenos Aires.

Chung, S.-W. & C.-I Peng. 1998. Eupatorium. In: T.-C. Huang. Flora of

Taiwan. 4: 953-956, t. 451,452, photo 450. Dept. Botany, National Taiwan

University, Taipei, Taiwan. 2" ed.

Cabrera, A.L. & N. Vittet. 1954. Catalogo de las Eupatorieas argentinas

(Compositae). Revista del museo de La Plata, Bot. 8: 242-243.

Cabrera, A.L., W.C. Holmes & S. McDaniel. 1996. Flora del Paraguay. 25:

63, t. 19-F/H, map 21. Conservatoire et Jardin Botaniques de la Ville de

Geneve, Switzerland & Missouri Botanical Garden, St. Louis, USA.

Corlett, R.T. & J.C. Shaw. 1995. Praxelis clematidea: yesterday South

America, today Hong Kong, tomorrow the world? Memoirs Hong Kong

Natural History Society. 20: 235-236.

Grisebach, A.H.R. 1879. Symbolae ad floram argentinam. Abhandlungen

der Koniglichen Gesellschaft der Wissenschaften zu Gottingen. 24: 172.

Jackson, B.D. 1895. Index Kewensis. 1: 916-a. Clarendon Press, Oxford.

King, R.M. & H. Robinson. 1970. Studies in the Eupatorieae (Compositae).

XXVIII. The genus Praxelis. Phytologia. 20: 193-195.

124 Gard. Bull. Singapore 51 (1999)

King, R.M. & H. Robinson. 1987. The genera of the Eupatorieae

(Asteraceae). Monograph Systematic Botany Missouri Botanic Garden.

22: 380-382, 535, 551.

Liogier, A.H. 1996. La flora de la Espanola. VUI. Universitie Central del

Este..72, Ser. Sci. 29120,

Robinson, B.L. 1920. The Eupatoriums of Bolivia. Contributions Gray

Herbarium. n.s. 61: 72.

Robinson, H. & R.M. King. 1977. Eupatorieae — systematic review. In:

V.H. Heywood and J.B. Harbourne.(eds.). The Biology and Chemistry

of the Compositae. pp. 446-448. Academic Press, London.

Schultz Bipontinus, C.H. 1866. Beitrag zur Geschichte und geographischen

Verbreitung des Cassiniaceae des Pollichiagebietes. Jahresbuch Pollichia.

22-24: 258.

Shi, C. 1985. Eupatorium L. In: Y. Ling & Y.-L. Chen, Flora Reipublicae

Popularis Sinicae. 74: 54-69. Science Press, Beijing.

Uniyal, B.P. 1995. Eupatorieae. In: P.K. Hajra, R.R. Rao, D.K. Singh &

B.P. Uniyal. (eds.). Flora of India. 12: 352-353. Botanical Survey of

India, Calcutta.

Waterhouse, B.M. & R.T. Corlett. 1996. Overlooked but still invading:

Praxelis clematidea, the unknown weed. Proceedings XIth Australian

Weeds Conference, Melbourne, 1996: 4 pp ms.

Gardens’ Bulletin Singapore 51 (1999) 125-126.

Obituary

Kizhakkedathu Mathai Kochummen

Kerala, India, 6 July 1931 - Kuala Lumpur, Malaysia, 31 March 1999

Mr Kochummen, better known as Koch, was unsurpassed in Malaysia in

his ability to identify a tree from a single leaf. It was his marvellous

photographic memory that enabled him to do this, honed during a decade

of botanical work at the Forest Research Institute at Kepong, where it was

his duty to confirm the identification of fallen leaves collected during the

large scale forest inventories that were being carried out by John Wyatt-

Smith throughout the Peninsula (Ng and Saw, 1999). This culminated in

his revisions of the Pocket Check List of Timber Trees (Wyatt-Smith, revised

by Kochummen, 1964 and 1979) and stood him in good stead when he was

involved in identifying the 335,240 trees with stem diameter 10 cm and

above to their 814 species in the 50-ha plot in the Pasoh Forest Reserve

(Kochummen, 1997).

Koch graduated with BSc from the University of Travancore in 1951

and, after coming to Malaya in 1952, obtained the post of Wyatt-Smith’s

Research Assistant in 1953. In 1955, he was promoted first to Assistant

Botanist, then to Forest Botanist in 1971 and Senior Forest Botanist in

1980. Retirement in 1986 only opened new horizons for him when he took

on consultancies (and he could name his price, for who else could identify

trees as accurately as he could!) and a Senior Fellowship (1986-1991) with

the Smithsonian Tropical Research Institute enumerating trees in the Pasoh

Plot. In 1992, he became Senior Botanist for the Tree Flora of Sabah and

Sarawak project, and before his untimely death had completed the revision

of the Moraceae, which included many new taxa (Kochummen, 1998).

Indeed, Koch had already gained a reputation for tackling large and

difficult families, such as the Lauraceae, Moraceae and Myrtaceae, from

his chapters in the Tree Flora of Malaya volumes where he revised 24 of

the 99 families.

To those who knew Koch personally, he will be greatly missed for

his cheerful help always freely given, his enthusiasm in identifying

specimens, his generous sharing of his experience in what are the vital

characters that identify a particular plant. His legacy of the knowledge of

practical and reliable vegetative characters of leaf, bole and bark are

recounted in his Tree Flora of Pasoh (Kochummen, 1997).

126 Gard. Bull. Singapore 51 (1999)

References

Wyatt-Smith, J. and Kochummen, K.M. 1964. Pocket Check List of Timber

Trees. Malayan Forest Record No. 17, Forest Research Institute,

Malaysia. 2" revised edition. (3" edition 1976).

Kochummen, K.M. 1997. The Tree Flora of Pasoh. Malayan Forest Records

No. 44, Forest Research Institute Malaysia, Kepong, Malaysia.

‘Kochummen, K.M. 1998. New species and variety of Moraceae from

Malaysia. Gardens’ Bulletin Singapore. 50: 197-219.

Ng, F.S.P. and Saw L.G. 1999. Kochummen, Kizhakkedathu Mathai. Flora

Malesiana Bulletin. 12: 251-253.

Ruth Kiew

Singapore Botanic Gardens

Singapore

eee ee

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The Gardens' Bulletin ©

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VOL. 51 (Part 2) Dec 1999 ISSN 0374-7859

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THE GARDENS’ BULLETIN

_ 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.

Dr Ruth Kiew

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Singapore Botanic Gardens

Dr T.W. Foong

(Assist. Editor)

Singapore Botanic Gardens

Dr S.C. Chin

Singapore Botanic Gardens

Dr M.J.E. Coode

Royal Botanic Gardens

Kew, U.K.

EDITORIAL BOARD

Dr R.T. Corlett

University of Hong Kong

Hong Kong

Dr M.C. Roos

Rijksherbarium

Leiden, Netherlands

Dr E. Soepadmo

Forest Research Institute Malaysia

Kepong, Malaysia

Dr W.K. Tan

Singapore Botanic Gardens

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| | ARNOLD

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The Gardens' Bulletin

Singapore

VOL. 51 (Part 2) Dec 1999 ISSN 0374-7859

CONTENTS

Holttum, R.E.

Mi eOicCa! DOLAINC PATCENS. PaASl, PRESEN ANG TUPUTE ..2,....0...:.-renrsonsnecnsnssarcencenesinereacesoeseses 27

Johnson, D.V. and E. P. Tay

Crurnado. (1369/1980); contributions to the study Of palais .......2i.2..c.csseessensnceoss 141

Kiew, R.

The Singapore Botanic Gardens Herbarium — 125 years of history ......... eects 131

Kim-Lang Huynh

On some species of Pandanus and Freycinetia (Pandanaceae) in Micronesia .............. 163

Turner, I.M. and P.F. Stevens

The transfer of Tripetalum cymosum K. Schum. (Guttiferae) to Garcinia ...........ee iW 2)

Kiew, R.

Thismia goodii (Burmanniaceae), the blue-capped thismia, a new species from

ane ee Ra ce ieee Fee enn forsee ache csieerre eae nero Sed rags vaca baadenebs cceapadansaaeeses 179

Boyce, P.C.

The genus Rhaphidophora Hassk. (Araceae-Monsteroideae-Monstereae) in

Ree aR LAL IVAN CE Sled ATAGH SN UMMSTeNIDES «feces on ccnccs kon sknbevasncensne dbs sanenecndasanverteceistesvonrssessSasssesensesees> 183

S.M. Tam

Floristic diversity of Bukit Bauk (Terengganu), Peninsular Malaysia .............. cee Zo7

Mabberley, D.J.

The importance to Indopacific botany of Baron Dumont de Courset’s

era meme eA IME oe neck Race sale pe Sada dss oytcuPAcah be oS Tolga ob Scviagnanviesevisesbacenes 309

BOOK REVIEWS

Bryological Biology, Introduction and Diverse Branches .................::cccceeeee Benito C. Tan - 319

MN ON re ee aes Sh tate ede esl ou Shas seh ray Uasegis chu aden sve dda lvlaededsvtessans Ian Turner - 321

Date of Publication: 31 Dec 1999

Published by

National Parks Board

Singapore Botanic Gardens

Cluny Road

Singapore 259569

Printed by Oxford Graphic Printers Pte Ltd

140" Anniversary of the Singapore Botanic Gardens

“History does not record the exact date. The nearest we can get 1s that the period of formation

[of the Botanic Gardens] was between the first half of November 1859 when a body of citizens

was formed with the express purpose of establishing a garden, and 1* January 1860 when the

first subscription became valid’”

Anon., 1959. Gardens’ Bulletin, Singapore. Frontispiece.

In 1859, the Singapore Agri-Horticultural Society was given a piece of derelict government

land at Tanglin for a pleasure garden but due to problems of debt, it was taken over by the

government in 1874 and gained the status of a botanic garden. (An earlier botanic garden had

been founded by Sir Stamford Raffles in 1822 on Fort Canning but it was closed in 1829).

Research activities covering plant taxonomy, horticulture and agriculture started in

earnest when H.N. Ridley became Director in 1888. It was Ridley who started the publication

of a journal by the Botanic Gardens in 1891 entitled The Agricultural Bulletin of the Straits

Settlements and Federated Malay States. This metamorphosed in 1913 into the Gardens’

Bulletin, Straits Settlements, which in 1947 became the Gardens’ Bulletin, Singapore.

It is therefore appropriate in the Singapore Botanic Gardens’ 140" year to include some

articles of a historic nature.

—— es

Gardens’ Bulletin Singapore 51 (1999) 127-139.

Tropical Botanic Gardens, Past, Present and Future!

RE. HOLTTUM

1895-1990

Assistant Director 1922-1925, Director 1925-1949

Singapore Botanic Gardens

Introduction

A botanic garden is essentially a museum of living plants. The word

‘museum’ is derived from the name of the Greek goddesses of learning

and the arts; thus a museum is a place devoted to the pursuit of such

studies. A botanic garden is primarily a place where plants are grown for

scientific study. But a garden differs from a museum in the fact that the

objects in it are living and growing, and thus need the attention of

horticulturists. Horticulture is in part applied botanical science: but it is

also in part an art, and the aesthetic aspects of horticulture therefore find

expression in any garden. But I would say that a garden cannot be accorded

the title ‘botanic’ unless it is a place where plant sciences of one kind or

another are studied, and where the results of such study are expressed

both in publications and in an arrangement and labelling, which conveys

information to visitors as well as aesthetic enjoyment. A park is a place

designed for recreation, not for study or education, and the plants in it are

selected and arranged solely for their aesthetic effect: here the horticulturist

alone has charge. Some of the older and smaller botanic gardens are now

in effect parks, and the distinction between botanic garden and park is not

generally understood by the public, sometimes also not by politicians. To

understand the present situation, one must look a little into the past.

History of Tropical Botanic Gardens

The principal interest in the early European botanic gardens (the first, at

Padua, Italy, dating from 1533) was medicinal plants. The study of such

plants in cultivation led to improved descriptions and illustrations being

published and thus to a wider interest in other plants and a gradual

development of botanical science. Plants from the New World were

introduced to such gardens from the late 16th century onwards, and also

carried across the Pacific to the Philippines by Spaniards, but tropical

Keynote Address presented at ‘The International Symposium on Botanic Gardens of the

Tropics’ in Penang, Malaysia in December 1984.

128 Gard. Bull. Singapore 51 (1999)

botanic gardens at which local and introduced species could be studied and

propagated were not founded until the 18th century. Apparently the earliest

was on the island of St Vincent in the West Indies in 1765. Its establishment

was due to interest in tropical plants by the Royal Society of Arts, and

probably in part to Kew, where the widowed Princess Augusta (mother of

King George III) began to develop a botanic garden in 1759 with advice

from the Earl of Bute who later acquired much influence with the young

King. The garden at St Vincent, like several other early tropical botanic

gardens (Calcutta, Bogor, Peradeniya) was under the control of a medical

man.

Soon afterwards the British initiative towards botanical exploration

and the establishment of tropical gardens passed to Sir Joseph Banks who

participated in Captain Cook’s first great voyage of exploration

(1768-1771). On his return, Banks met the King, who asked him to take a

personal interest in the royal gardens (of which there were two, hence the

plural) at Kew. Banks had been impressed by the wealth of previously

unknown plants, which he saw in Australasia and the Pacific, and the need

for their study both for a fuller understanding of plant taxonomy and for

the establishment of information about those that were useful in many

kinds of ways. He used his great influence, as President of the Royal

Society for 43 years, to promote botanical exploration in various parts of

the world, introducing living plants of a great number of species to Kew,

whence they were distributed widely. He also promoted the collection of

dried botanical specimens, which would serve as a basis for study of plants

which could not be cultivated at Kew. The best-known example of transfer

of useful plants from one part of the tropics to another through the influence

of Banks is that of breadfruit from Tahiti to the West Indies. The first

attempt by William Bligh in The Bounty in 1787 failed. The second and

successful voyage of Bligh in 1793, on which many other plants were also

carried, is less well-known. Bligh’s plants were received at the botanic

gardens in St Vincent and Jamaica, the latter established in 1775 through

the influence of Banks. An important example of botanical exploration

promoted by Banks is his engagement of Robert Brown as botanist on the

expedition under Captain Matthew Flinders for surveying the coast of

Australia (1801-1805). Brown’s account of his collections, published in

1810, is a work of basic importance. Banks was also actively concerned

with botanical developments in India.

W.J. Hooker came to London in 1805, a young man of modest

independent means who wished to devote himself to the study of plants.

He was welcomed by Banks, through whose influence he was later (1820)

appointed Regius Professor of Botany at Glasgow. Banks died in 1820

and botanical exploration based on Kew ceased. In 1838, the Government

Tropical Botanic Gardens 129

appointed a committee to consider the future of Kew, under the

chairmanship of Professor John Lindley who, like Hooker, had owed his

early progress to the patronage of Banks. The result was that from 1841

Kew became a national botanical institution with Hooker as its first

Director, and the tradition of Banks was revived there. During the period

1820-1841, Hooker had been in close contact with botanical exploration in

many parts of the world and had accumulated a large herbarium, also a

collection of specimens of useful plants. His advice was sought by the

Admiralty and by the Colonial Office, where the need for knowledge

about plants in remote parts of the world was a spur to action. At Kew,

Hooker and successive Directors were recognized as official advisers in all

such developments, and this process continued until the period of

decolonization from 1950 onwards. I owed my appointment as Assistant

Director of Gardens, Straits Settlements, in 1922 to the recommendation

of Kew, and Kew has remained my headquarters until the present time.

Thus Kew was concerned with the establishment of gardens at many

places in the tropics, received from all of them living plants, especially of

useful and ornamental kinds, and effected an exchange of such plants from

one part of the tropics to another. Publication of various kinds on the

culture of these plants was effected locally, dependent on the more or less

active interest of those in charge of the gardens and the local government

officials, and also through Kew.

Kew was also concerned to accumulate information about other native

plants with a view to the preparation of local floras. This information was

largely gathered from dried specimens, beginning with the personal

herbarium of William Hooker (estimated at one million specimens) and of

his friend, George Bentham, who wrote the first British colonial flora:

Flora Hongkongensis (1861). The colonial gardens were thus also asked to

take an interest in the local plants, which were not cultivated, the most

important of which were forest trees, and to despatch specimens and other

information to Kew. In this way, colonial botanic gardens became centres

of local knowledge on horticulture, agriculture, forestry and taxonomic

botany, and in many cases they were the centres where local departments

of agriculture were later established.

The two most important botanic gardens in tropical Asia, at Calcutta

and Bogor, had a different history. The Calcutta garden was first established

by the East India Company for the cultivation of teak trees, important for

shipbuilding, in 1787. It began to serve wider purposes with the appointment

of William Roxburgh as Superintendent in 1793. Roxburgh had been in

the medical service of the East India Company at Madras since 1776 and

there met J.G. Koenig who had been a pupil of Linnaeus. He thus became

interested in native plants and wrote the first Flora Indica, but he was also

130 Gard. Bull. Singapore 51 (1999)

interested in the culture of useful plants, native and otherwise, and

introduced nutmegs and cloves from the Moluccas to Penang. While still

at Madras, he engaged artists to make coloured illustrations of many local

plants and a selection of 300 of these were published in London through

the interest of Banks in three large folio volumes, as the Plants of the

Coast of Coromandel. Nathaniel Wallich, who succeeded Roxburgh in

1815, organized the collection of native Indian plant specimens over a

much wider area, and also in Singapore in 1822, accumulating a very large

herbarium from which many duplicates were distributed. This is still an

important basis for the taxonomy of Indian plants. During one of Wallich’s

absences from Calcutta, William Griffith took charge and also travelled in

Burma and the Malay Peninsula. He used a microscope and made the first

detailed morphological observations on many plants; these were published

after his death (at the age of 35) in 1845. The last active Superintendent at

Calcutta was George King (1871-1898). He made the first monographic

studies of Indian plants; his work on Ficus included the first thorough

study of the process of pollination in that genus. He established the

Botanical Survey of India in 1891.

The botanic garden at Bogor in Java was established by C.G.

Reinwardt in 1817; he was also Director of Agriculture, Arts and Sciences.

A Kew-trained horticulturist, James Hooper, who happened to be in Java

owing to a shipwreck, was appointed as first Curator, and remained until

1830. Reinwardt also promoted local botanical collecting. He left Java in

1822 and was succeeded as Director by C.L. Blume from. 1822-1826 in

which short period Blume undertook much botanical collecting and hastily

wrote descriptions of a large number of new species. After leaving Bogor

he became Director of the Rijksherbarium at Leiden, publishing more

detailed and beautifully illustrated descriptions of many plants from Java.

At Bogor, the post of Director was abolished and the Garden was

maintained by J.E. Teysmann, Curator, from 1831-1867. He travelled

widely and introduced a large number of plants to Bogor, where he arranged

them systematically. The result is that Bogor now has incomparably the

finest collection of woody plants to be found anywhere in the tropics. In

1880, Melchior Treub became Director and began modern scientific

botanical studies. He built a laboratory for visiting botanists in 1885, in

which 250 people worked for varying periods during the next fifty years.

Treub also founded other research institutes, which would be needed for a

department of agriculture. Such a department was founded in 1905 with

Treub as its first Director, but, after he retired in 1910, the research institutes

at Bogor became separated from the Department. Active botanical study

of various kinds has continued in which the living plants of the garden

have an important part, and the large herbarium of specimens from all

Tropical Botanic Gardens 131

parts of Indonesia is of basic importance for the continued production of

Flora Malesiana. Both garden and herbarium are now parts of Lembaga

Biologi Nasional.

Two other gardens in SE Asia have also been important and need

some further comment. The Peradeniya garden in Sri Lanka was established

in 1822. Active botanical study there began with the appointment of

George Gardner as Director in 1844, followed by G.H.K. Thwaites in

1859. From then onwards active floristic study of native plants continued,

also the introduction, through Kew, of exotic plants, among them Cinchona

and Para Rubber. H.J. Trimen, who succeeded Thwaites, established a

laboratory for visiting botanists in 1888. After retiring, he wrote a Flora of

Ceylon. The next Director, J.C. Willis, initiated modern botanical studies

of various kinds, but after his retirement in 1911 the Garden was made the

base of a new Department of Agriculture and purely botanical work ceased,

apart from Petch’s work on fungi until 1925 and Alston’s work as systematic

botanist from 1925-1930.

The garden in Singapore was effectively established in 1874 when

the Government took over the garden started by a local society. HJ.

Murton, trained in horticulture at Kew, was sent to take charge and received

the first plants of Para Rubber. He was energetic but unstable and was

replaced by Nathaniel Cantley in 1880. Cantley had training in forestry

and had the title Superintendent of Gardens and Forests, Straits Settlements.

He established the first forest reserves in Singapore, Malacca and Penang,

and the Garden at Penang in 1884, the latter primarily as a forest nursery.

Charles Curtis, Curator of the Penang Garden, had been a collector of

ornamental tropical plants for the firm of Veitch, and laid out the garden

with an eye to landscape effect. He was much interested in local plants of

all kinds, and helped H.N. Ridley, who succeeded Cantley in 1888, by

collecting specimens in Penang and elsewhere in the north of the Peninsula.

He was the first person to introduce Congea to cultivation. Ridley (at

Singapore 1888-1911) was a man of enormous energy who did an immense

and very varied work. He is best known for his pioneer experiments on

tapping Para rubber trees, which were the beginning of the plantation

rubber industry in Malaya, but he studied and wrote extensively also about

the useful plants of Malaya, including forest trees, and undertook much

botanical field work on which he largely based the Flora of the Malay

Peninsula which he wrote after retirement. During his service, the

Departments of Agriculture and Forests were established at Kuala Lumpur

but botanical activity continued at Singapore instead of being suppressed

as at Peradeniya. Penang was always complementary to Singapore where

a considerable herbarium and library were accumulated; Penang was a

base for observation of the plants in northern Malaya and their behaviour

132 Gard. Bull. Singapore 51 (1999)

in a climate distinctly different from that of Singapore. When Penang

became part of the Federation of Malaya, the Garden there was separated

from its headquarters and there was no local scientific base to which it

could be affiliated. The collections of interesting local plants brought

together with much care by Mr Cheang Kok Choy received no official

recognition. The local Government appeared to regard the Garden merely

as a park for recreation, not for study or educational purposes.

Future of Tropical Botanic Gardens

It is thus appropriate that this Symposium is held at Penang, where the

question of the future of a tropical garden is of immediate concern. I

suggest that a botanic garden must be equipped as a research institute in

its own right, or be closely affiliated with such an institute. All university

departments of botany have the need to cultivate plants for experiment

and for teaching material, and the botanic gardens of Europe are, in many

cases, affiliated to or constituent parts of universities. Tropical gardens

were started before local universities, but universities have now been

established and I suggest that formal connections of some sort between the

two are needed for the good of both. A garden cannot be called botanic

unless it is associated with a research institute and a library.

The nature of the study undertaken at any one garden must be limited

and will vary according to local circumstances and local climate. Though

much of it, under university auspices, will be purely scientific, some of it

may have a practical aim, or yield results of practical importance. There is

much more to be discovered about the possible uses of tropical plants;

and, for purposes of exploitation, breeding and selection involving

cytotaxonomic study are necessary. Basic studies could be planned to

facilitate such processes. And it must always be remembered that

horticultural skills and experience are necessary for success in producing

healthy well-grown plants. Different plants show many kinds of adaptation

to the environment and may need different cultural treatment; plants never

previously cultivated may present new problems. Plants vary also very

greatly in their response to different methods of vegetative propagation.

So an experienced horticulturist is needed to take charge, and should have

adequate status. Some experimental work in a garden may well concern

horticultural problems. There is still much to be learned about every

aspect of tropical plants. There is room for research on quite simple

horticultural processes such as the preparation of potting composts suited

to different climates. The development of the burnt earth technique in

Singapore as an adaptation to a climate of heavy rainfall is worth some

Tropical Botanic Gardens 133

critical experimental study.

Basic to all other studies of plants is a good taxonomy. Though the

taxonomy of plants native in most temperate latitudes is now well

understood, that of many tropical plants is not, and there is still need for

taxonomic study in tropical botanic gardens. This is not well understood

by many academic botanists, for which reason I am impelled to make a

statement upon it, based on my own experience. Until the second half of

the 19th century, taxonomy was the principal botanical study in Europe.

During that century, anatomical and then physiological studies gradually

developed and rightly led to the idea that the limited observations of

taxonomists were very inadequate for a good understanding of plant biology.

The new men wanted “scientific” botany. So far as European plants were

concerned, a fairly full taxonomic knowledge had by then developed, but

the new scientific botanists did not understand that this was not true of

tropical plants. They wrongly concluded that taxonomy was unscientific

and ceased to study it. When I was a student at Cambridge, it was clearly

indicated to me that taxonomy was an inferior discipline, and I was a little

ashamed of my interest in it.

When I went to Singapore in 1922, I had to use Ridley’s taxonomic

works in order to identify some of the plants which 1 saw around me, and I

was appalled by the inadequacy and often inaccuracy of his descriptions. I

gradually realized that what was needed was a better taxonomy, and that

this was basic for all other studies of local plants. Since that time 1 have

devoted the main part of my sustained study to taxonomy, while trying

also to gather information about plants as living organisms.

Taxonomy has two objectives. One is to enable people to identify

species, and this is important because information about plants is attached

to species names, and is unreliable unless the names have a definite meaning.

The other objective is to understand the inter-relationships of species; the

recognition of natural genera, families and groups of higher order.

The need to produce descriptions from which species may be

identified necessitates the use, where possible, of easily observable

characters. It makes no sense to require the observation of chromosomes

as the first step towards the identification of an unknown plant, There are

in fact many easily observable characters in most plants, so that in practice

the process of identification to the species level can usually be completed

with the use of a hand lens. Academic botanists tend to think that unless

one can use the latest, most complex and most expensive equipment one is

not being really scientific. This is a main reason for despising taxonomy

but it is not a valid reason. You must first use a hand lens before you can

decide the meaning of what you see with an electron microscope. And

there is still much more structure to be seen at low magnifications then is

134 Gard. Bull. Singapore 51 (1999)

realized by most botanists.

The plants of today are the result of a process of evolution: they

therefore have an inbuilt classification, and our problem is to find it. The

nearer a Classification is to being natural, the nearer it is to expressing

phylogeny. The difference between traditional taxonomy and cladistics is

that cladists analyse the significance of their taxonomic judgements in a

precise way. This analysis can give no guarantee that the judgements are

correct, still less does it tell us whether a further set of characters might

have modified the result of the analysis.

Another consideration is that you cannot understand how to delimit

a genus until you have seen all the species. The binomial system established

by Linnaeus, which is still followed as a basis for naming species, was only

effective because Linnaeus had first learned to know a large proportion of

European plants thoroughly and could therefore judge their natural

relationships. But those botanists (including Linnaeus) who had to deal

with the few early discovered tropical plants could not make such a

judgement. This has led to many changes in generic concepts and to much

confusion in synonymy. To be acceptable according to the present Code, a

name must be a binomial, so that one has to assign the species to a genus,

whether or not one is well enough informed to do so. An intelligent 18th

century taxonomist describing previously unknown plants, wrote Anonymos

in place of a generic name for some of them because he was uncertain of

their genera, but this wise man’s names are now declared illegitimate.

And because every species name is a binomial, morphologists and

experimental botanists tend to think that any species must be representative

of the generic name which it bears. A generic name (like any other name)

has only as much meaning, or as little, as the knowledge of a person who

uses it, whether orally or in print.

The nomenclature of ferns is much more confused than that of

angiosperms. In recent years, I have tried to make a survey of all species

of the fern genus Tectaria and its allies throughout Asia, Malesia and the

Pacific. One of these genera is Crenitis, which was first distinguished as a

natural group by Christensen when he studied the species of tropical

America. But the botanists who first tried to distinguish the species of

Ctenitis in Asia did not look as critically as Christensen had done, and in a

recent survey of them all, I discovered that 48 species had been wrongly

assigned to that genus. Those species belong properly to nine other genera:

and the fact that various authors assigned them to Ctenitis indicates that

those authors had also no clear concepts of the other genera.

The angiosperms with which I am best acquainted are some

monocotyledons, especially bamboos, orchids and Zingiberaceae. I believe

that in general monocotyledons are more difficult to understand from dried

Tropical Botanic Gardens 135

specimens than dicotyledons and that therefore it is more necessary to

study them from living plants and thus most conveniently in cultivation.

This is because they have no secondary growth in thickness and therefore

their primary vegetative structures must be of great precision. The precise

branching of inflorescences and the arrangement and nature of bracts or

bracteoles are of great significance in Zingiberaceae, as shown by Valeton,

but ignored by earlier authors. These bracts are often of delicate structure

and closely overlapping, so that, it is difficult to see them clearly in dried

specimens. There is much still to be learned about the taxonomy of this

family in Malesia, especially in New Guinea. In orchids there is much

greater complexity in the structure of flowers, and, when the flowers are

quite small, the structures are delicate and not well observable when dried,

but inflorescence structures in some genera are not well understood, notably

in Dendrobium and Eria. In both families, there is still much to learn

about the delimitation of genera.

Bamboos can only be understood from a study of living plants, and

very few taxonomists have made any considerable such study. I learned

how to look at a bamboo by studying Gamble’s great work on Indian

bamboos, but observation of Malayan species showed that some

modification of Gamble’s generic concepts was necessary; and I believe

that no one has yet discovered how to distinguish genera from non-flowering

material. When I went to New Guinea in 1963, I decided to try to

understand the bamboos there; those already recorded had been named

and described only from fragmentary flowering specimens. 1 found that

the commonest small bamboo near Lae had been given two different names,

both certainly in wrong genera. | had to remove it from those genera and

transferred it to Bambusa, but it is aberrant in Bambusa and (along with

some allied species) needs further study.

The above comments are made to show that tropical taxonomy is

still in need of more study, and that such study is best undertaken from

examination of living plants, most conveniently from plants in cultivation

and therefore easily available over a period necessary to display all stages

of their development. The opportunity for making such study is still an

important function of tropical botanic gardens. This has been recently

demonstrated at Penang by the cytological studies in Universiti Sains

Malaysia of a collection of plants of Zingiberaceae at the Botanic Garden,

brought together there over several years from neighbouring areas by Mr

Cheang Kok Choy.

In view of the present widespread destruction of tropical forests, the

conservation of tropical plants of all kinds is of increasing importance.

Tropical botanic gardens are places where public attention may be drawn

to this subject. But cultivation of native plants in such gardens can do little

136 Gard. Bull. Singapore 51 (1999)

towards conservation of species. The only practicable method is the

conservation of sufficient areas of natural forest, in which individual species

can find their natural habitats. Botanic gardens may do useful service by

cultivating limited range of species, but the total number is so large, and

their needs so varied, that to preserve them all in cultivation is not

practicable. The botanic gardens at Cibodas in Java and in Penang are

adjacent to areas of natural forest; the forest is an area where local species

are protected, and their presence adds to the value of the gardens, both for

research and public interest. Even the small area of natural forest within

the botanic garden in Singapore has this function and is being used by

local school teachers.

Early taxonomy was concerned with only a limited range of characters,

and it was some time before taxonomists realized that new observations

made by morphologists and anatomists might be pertinent to taxonomy.

There was some very inferior taxonomic work in the latter part of the 19th

century. But in more recent years taxonomists have taken a new range of

morphological and anatomical characters (including chromosomes) into

consideration. Every genus presents a different set of problems, and there

is endless opportunity for study of new kinds of characters, such as those

revealed by the scanning electron microscope. Cultivation together of a

group of allied species may lead to new insights. The selection of such

plants will depend on the interests of local staff members. I have indicated

above some structures needing investigation in orchids, bamboos and

Zingiberaceae. The maintenance of such a set of plants for comparative

study involves the concerned interest of a horticulturist. Living plants

cannot be left untended like specimens stored in a museum; unless someone

keeps a constant watch, some of them will soon be lost.

Floral biology is another study, which has many facets of

interest, relating to the development of flowers and their relationship to

pollinators, and is both of ecological interest and of practical significance

in plant breeding. There are many groups of tropical plants, which have

never been “improved” by scientific methods. The best edible tropical

fruits have been selected from time immemorial, for example, mangoes

and bananas, but modern methods of hybridization could produce radically

new developments. Why cannot a lychee be crossed with a rambutan?

Who can say what are the possibilities attainable by mangosteens with

other species of Garcinia? New races of Malayan mangoes might be

produced by crossing good Indian varieties, which do not yield well here,

with native Malayan species of Mangifera. Such experimental work would

take years to accomplish, and considerable areas of ground, but if never

started, they will never be realised. There are really great possibilities.

With the selection of new woody plants, whether fruit trees or

Tropical Botanic Gardens 137

ornamentals, comes the necessity for vegetative propagation. In some

cases micropropagation is the best, or indeed the only method, as with

palms. Oil palms have now been propagated in this way, but I believe that

more work is desirable on both oil palms and coconuts, which together are

the most important sources of vegetable oils. Rattans are another important

group of palms, which are now receiving more attention, and

micropropagation might be valuable for them. But though wonderful new

plants may be produced, it is necessary to maintain stock of old ones, so

that further breeding may be possible. Botanic gardens are places where

such stocks may be preserved.

A feature everyone expects to see in gardens is flowers, and surely

botanic gardens should not neglect this. In north temperate regions there

is a considerable change of temperature through the year, and flowers

have been developed to suit the various seasons. Many of them, e.g. roses,

owe their present condition to breeding and selection over a long period of

time. In the tropics, little breeding for production of new flowers has been

done and climatic situations are different. There is no cold season and

seasonal change is due to the distribution of rainfall through the year. In

those areas where a long dry season prevails there is an obligate succession

of flowers, though different from that in temperate regions, but in areas

such as Malesia, dry seasons are short and sometimes inconstant, so that

some plants can flower throughout the year. The extreme case is Singapore

where there is no dry season and slight spells of dry weather occur

erratically. In such conditions, woody plants from strongly seasonal areas

sometimes fail to flower. In Singapore, we had a plant of Bougainvillea

which was introduced from Kew in 1889 but never flowered from that time

onwards (at least until 1954) though it grew quite strongly. Many other

woody plants introduced from seasonal parts of the tropics, whether natural

species or hybrid cultivars, proved useless for floral display. This was

equally true of orchids, in which family hybrids had been raised mainly for

cultivation in greenhouses in temperate latitudes. Most local orchids did

not flower continuously but only in response to casual slight climatic change

as, for example, the commonest epiphytic orchid Dendrobium crumenatum.

The big breakthrough in Singapore came in 1893 with the production of

hybrid Vanda ‘Miss Joaquim’, one parent of which was one of the few

perpetual-flowering local orchids. Vanda ‘Miss Joaquim’ proved perpetual

in flowering and stronger than its parent. The production of further hybrids

in Singapore from 1928 onwards based on strong-growing local species,

showed that other hybrids could be perpetual-flowering, and nearly all

hybrids were stronger in growth and more free flowering than their parents.

Orchid breeding since that time has resulted in the present situation: orchid

flowers are now the most abundant and most varied garden flowers in

138 Gard. Bull. Singapore 51 (1999)

Singapore. It is my conviction that woody plants need the same treatment

if a better display of flowers is to be produced. The plants of the wet

tropics, apart from monocotyledons, are nearly all shrubs or trees: annual

or short-lived plants have no natural place. Garden plants of the wet

tropics grow throughout the year: why should they not also flower

throughout the year? Some such plants exist, for example, Plumeria, of

which there are now many hybrids, and Mussaenda in which genus hybrids

between Philippine and West African plants were developed at Los Banos,

and are now common in Malaysia. I suggest that experiments towards the

production of new hybrids in genera, which are productive of beautiful

flowers would not be out of place in a tropical botanic garden, and that

this, might be an important activity in the ever-wet tropics.

Most of the older tropical gardens were established at sea level, but

a few subsidiary gardens were added later at higher altitudes where a

different range of plants could be grown, notably at Cibodas in Java,

Hakgala in Sri Lanka and Darjeeling in northern India. There is much

room for further developments of this kind, possibly in relation to National

Parks, which are designed to conserve native mountain plants.

Botanic gardens in north temperate regions usually produce seed

exchange lists each year. But in Singapore there was no regular annual

production of seeds; plants mostly either produced seeds continuously or

not at all. Therefore we circulated a list which could be valid for a period

of years. I suggest that tropical gardens in general should do this. This

raises the question of seed banks. I am not sure whether such banks are of

value in the tropics where most plants are perennial in growth and flowering.

But if seed banks are attempted, there must be experiment to discover the

best means of preserving viable seeds of different kinds. Many seeds will

not withstand dry storage, and some tropical seeds have no resting period;

they either germinate at once, or die, as for example, durian seeds and

those of many dipterocarps, which are.such important timber trees in

Western Malesia.

As regards the scope for physiological experiment on cultivated

tropical plants, there seem to be endless possibilities and I am no expert in

this field. But I do suggest that plants in tropical gardens might be more

used in experiments. Studies in the physiology of photosynthesis and

respiration in orchids and ferns have been undertaken in Singapore. I

should like to know how the pattern of growth in fronds of the family

Gleicheniaceae is controlled; it depends on the permanent dormancy of

the apical growth of some branches of the frond. Penang would be an

ideal place to conduct such experiments, as a range of different species,

with different growth patterns, are here native. No such studies have ever

been made.

Tropical Botanic Gardens 139

My last word concerns horticulture, to which I revert because without

good horticulture you would have no gardens. And in botanic gardens

you will need to grow new plants, which may require new treatment; the

job is not a routine one like that of a commercial nursery. You must give

your horticulturist, who is a professional man in his own right, some chance

to show his talents for display of plants. But I must add a warning to

horticulturists, sets of plants for record or study must be maintained as

separate individuals, not allowed to form a thicket in which individual

plants of different species merge with each other, This happened a few

years ago at Kew when horticulturists insisted on landscaping in which

many species of Se/aginella were planted near together; most of them were

soon suppressed by the more vigorous ones and lost. Fortunately, someone

insisted on keeping a set of plants in pots in another place.

A botanic garden is primarily a place for the study of plants, and

horticulturists should remember this. John Smith, Kew’s first Curator

(himself a very able horticulturists) wrote in 1866 in Ferns, British and

Foreign (p. 43): “my long experience has shown that as soon as scientific

arrangement in any family of plants is lost sight of, and showy cultivation

made the first consideration, a rapid loss of species is the sure consequence”.

Acknowledgement

We are grateful to Dr Leong Yueh Kwong, organiser of ‘The International

Symposium on Botanic Gardens in the Tropics’ for permission to reproduce

this unpublished paper.

Gardens’ Bulletin Singapore 51 (1999) 141-150.

C.X. Furtado (1897-1980):

Contributions to the Study of Palms

DENNIS V. JOHNSON! AND ENG PIN TAY?

'3726 Middlebrook Avenue, Cincinnati OH 45208, USA

9 Romulea Mews, Livingston, Canning Vale, WA 6155, Australia

Abstract

An account of Caetano Xavier dos Remedios Furtado’s academic career, his taxonomic

work on Malayan palms and the African genus Hyphaene is given, together with a complete

list of his publications on palms and a resumé of his travels in connection with his research

on palms.

Introduction

Caetano Xavier dos Remedios Furtado was born in Goa, India, 14 October

1897. He attended the Poona Agricultural College in India and while an

undergraduate began to write technical articles, especially on the coconut

palm. The first article was published in 1919. After completing his B.Sc.

degree in 1921, Furtado obtained employment as an agronomist in Burma

where he continued his interest in coconuts. He joined the staff of the

Singapore Botanic Gardens in 1923 and within a few years began a lengthy

study of Malayan palms. Primarily on the basis of his publications on palm

research in the 1930s, Furtado was awarded a D.Sc. degree from the

University of Bombay in 1939. His thesis not only gave an account of these

researches but also highlighted the number of languages he mastered to

tackle palm taxonomy and also his interest in botanical nomenclature

(Appendix 3). Dr Furtado retired in 1952 but was re-employed as Botanist

until 1964 (Kiew, 1999). Even after this second retirement, he continued to

conduct research and to publish botanical articles for nearly another decade,

his last publication appearing in 1970.

Because his professional achievements in life and at his death were

overshadowed by the works of more eminent scientists, Furtado has not

been given the recognition he deserves as a botanist and palm specialist.

Furtado worked closely with E.J.H. Corner and R.E. Holttum when

they were Assistant Director and Director of the Singapore Botanic

Gardens. All three continued to work at the Gardens during World War II

in the period of Japanese military occupation of Singapore, during which

time Furtado was responsible for the administration of the Gardens. The

142 Gard. Bull. Singapore 51 (1999)

. |

Figure 1. Caetano Xavier dos Remedios Furtado.

C.X. Furtado 143

work of these two British botanists has tended to obscure Furtado’s

botanical research and publication.

At the time of his death on 13 June 1980, Furtado’s research and

writing on palms went almost unacknowledged because so much attention

was drawn to the death, four months earlier, of the preeminent world palm

expert H.E. Moore, Jr. Only two brief obituaries were published about

Furtado (Alphonso, 1980; Holttum and Dransfield, 1981) neither was

detailed nor provided a listing of his palm publications. A short informative

entry on Furtado appears in the Cyclopedia of Collectors (Steenis-

Kruseman, 1950).

Furtado’s Contribution to the Study of Palms

This article seeks to document the contributions C.X. Furtado made to the

study of palms, especially to Malayan palms and the African genus

Hyphaene. In addition to his work on palms, Furtado is known for his

contributions to nomenclature and Araceae systematics.

Appendix I provides full citations of Furtado’s 42 publications on

palms that could be located. The listing is believed to be complete, but it

was not possible to gain access to Furtado’s personal professional papers

to be certain. As the listing shows, Furtado’s early interest in the coconut

palm accounts for 11 publications from 1919 to 1927.

C.X. Furtado’s most significant palm research centred on an extensive

study of Malayan palms, which produced a series of 19 articles published

between 1934 and 1956. In retirement, Furtado embarked on a herbarium

study of the mostly African genus Hyphaene resulting in five articles.

Four new,palm genera were erected by Furtado:. Cornera,

Liberbaileya, Maxburretia, and Schizospatha. Subsequently, Cornera and

Schizospatha were reduced to synonymy with Calamus and Liberbaileya

combined with Maxburretia.

On the basis of a computer search of Index Kewensis, Furtado

described 104 palm species, mostly in the genera Calamus, Daemonorops,

Hyphaene, Licuala and Pinanga. Comparing the species Furtado described

with the February 1997 listing of palms provided by the World Conservation

Monitoring Centre, Cambridge, 53 species names remain valid. However,

further species names could be reduced to synonymy, especially in

Hyphaene, which is in need of revision.

A last and most appropriate recognition of Furtado’s palm research

is to be found in the binomials Maxburretia furtadoana J.Dransf. and

Korthalsia furtadoana J.Dransf. The first was named “in recognition of his

great contribution to our understanding of Malayan palms” (Dransfield,

144 Gard. Bull. Singapore 51 (1999)

1978); the second to recognize the herbarium research Furtado carried out

on that genus (Dransfield, 1981).

C.X. Furtado’s palm research extended over a period of more than a

half-century. As documented here, Furtado’s record of achievement

represents a major contribution to the scientific knowledge about the

important palm family.

Acknowledgements

The authors gratefully acknowledge the cooperation and assistance of:

Singapore Botanic Gardens’ Herbarium and Library; Royal Botanic

Gardens Kew for a computer search of Index Kewensis; World Conservation

Monitoring Centre, Cambridge; the Flora Malesiana Foundation for

permission to reproduce the portrait; Jose Ireneu dos Remedios Furtado;

John Dransfield and Ruth Kiew.

References

Alphonso, A.G. 1980. Caetano Xavier Furtado 1897-1980: an obituary.

Gardens’ Bulletin Singapore. 33: 301-02.

Dransfield, J. 1978. The genus Maxburretia (Palmae). Gentes Herbarum.

11(4): 187-199.

Dransfield, J. 1981. A synopsis of Korthalsia (Palmae-Lepidocaryoideae).

Kew Bulletin. 36: 163-94.

Holttum, R.E. and Dransfield, J. 1981. Furtado, Caetano Xavier. (Obituary)

Flora Malesiana Bulletin. 8(34): 3543-44.

Kiew, R. 1999. The Herbarium — 125 years of history. Gardenwise. 13: 6-9.

Steenis-Kruseman, M.J. van. 1950. Furtado, Caetano Xavier Dos Remedios.

In: Cyclopaedia of Collectors. Flora Malesiana. Ser. 1. 1: 183.

C.X. Furtado 145

Appendix I. C.X. Furtado: Publications on Palms

(GBSS Gardens’ Bulletin Straits Settlhements; GBS Gardens’ Bulletin

Singapore)

1919. Some tillage and labour problems of the coconut planter of Western

India. The Poona Agricultural College Magazine. 11(2): 105-107.

1920. Oryctes rhinoceros, a coleopteron injurious to the coconut palm in

Goa, Portuguese India. Boletim de Agricultura Nova Goa. 2(1/4):

82-85.

1920. Abnormal inflorescences. (of coconut palm) Poona Agricultural

College Magazine. 12(2): 81-83. (co-authored with P.C. Barreto)

1922. Vegetative reproduction in cocoanuts (sic). Poona Agricultural

College Magazine. 15(3): 144-146. (author listed in error as C.H. Furtado)

1923. Coconut tapering disease. Tropical Agriculturist. 61(2): 126. (letter

to editor)

1923. The coconut inflorescence. Poona Agricultural College Magazine.

14(4): 213-221.

1923. Ripening of coconut flowers. The Agricultural Journal of India. 18(5):

561. (letter to editor)

1924. Ripening of coconut flowers. Poona Agricultural College Magazine.

15(4): 240.

1924. A study of the coconut flower and its relation to fruit production.

GBSS. 3: 261-274.

1924. Branched coconut palms and their fertility. GBSS. 3: 274-279.

1927. Teratological notes. A. Abnormalities in coconut palms. B. Ananas

sativa. GBSS. 4: 78-86.

1933. The limits of the genus Areca Linn. and its sections. Feddes

Repertorium. 33(8-17): 217-239.

1934. Palmae Malesicae (I. Species of Jguanura and Pinanga). Feddes

Repertorium. 35(18-25): 273-283.

1935. Palmae Malesicae II. Nenga Wendlandiana Scheff. or Nenga pumila

(Mart.) Wendl? GBSS. 8: 159-163.

146 Gard. Bull. Singapore 51 (1999)

1935. Palmae Malesicae III. Notes on some Malaysian Calami. GBSS.

8: 241-261.

1935. Palmae Malesicae IV. Rattans described in Blanco’s Flora de Filipinas.

GBSS. 8: 321-338.

1935. Palmae Malesicae V. Notes on some Malayan Daemonorops. GBSS.

8: 339-367.

1937. Palmae Malesicae VI. Notes on some Daemonorops of the section

Cymbospathae. GBSS. 9: 152-181.

1937. Palmae Malesicae VII. Two new Calami from the Buitenzorg

Gardens. GBSS. 9: 182-186.

1939. Contributions to the Systematics of the Malaysian Palms. D.Sc. thesis

(Agriculture) Botany, University of Bombay.

1940. Palmae Malesicae VIII. The genus Licuala in the Malay Peninsula.

GBSS. 11: 31-73.

1941. Palmae Malesicae IX. Two new Coryphaceous genera in Malaya.

GBSS. 11: 236-243.

1949. Palmae Malesicae X. The Malayan species of Salacca. GBS. 12:

378-403.

1951. Palmae Malesicae XI. The Malayan species of Korthalsia. GBS. 13:

300-324.

1951. Palmae Malesicae XII. The Malayan species of Plectocomiopsis. GBS.

13: 325-338.

1951. Palmae Malesicae XIII. The genus Myrialepis. GBS. 13: 339-344.

1951. Palmae Malesicae XIV. The species of Plectocomia in Malaya. GBS.

13: 345-350.

1951. Palmae Malesicae XV. The genus Ceratolobus in Malaya. GBS. 13:

351-359.

1951. Palmae Malesicae XVI. The little-known Malayan genus Calospatha.

GBS. 13: 360-365.

1953. (Palmae Malesicae XVII). The genus Daemonorops in Malaya. GBS.

14: 49-147.

C.X. Furtado 147

1954:

Palmae Malesicae XVIII. Two new Calamoid genera of Malaysia.

GBS. 14: 517-529.

1956.

Palmae Malesicae XIX. The genus Ca/amus in the Malayan Peninsula.

GBS. 15: 32-265.

1959.

1960.

1963.

1964.

196].

1970.

Singapore’s contribution to the study of palms. GBS. 17: 195-198.

The philological origin of Areca and Catechu. Principes. 4: 26-31.

Malayan ornamental palms. Principes. 7: 31-32.

The origin of the word “Cocos”. GBS. 20: 295-312.

On the etymology of the word Cocos. Principes. 8: 107-112.

Some notes on Hyphaene. Garcia de Orta. 15(4): 427-460 + 19 plates.

Some Hyphaene species from the Botanic Gardens, Calcutta.

Principes. 14: 117-123.

1970.

The identify of Hyphaene natalensis Kuntze. GBS. 25: 283-297.

Asian species of Hyphaene. GBS. 25: 299-309.

A new search for Hyphaene guineensis Thonn. GBS. 25: 311-334.

148 Gard. Bull. Singapore 51 (1999)

Appendix 2. C.X. Furtado’s Palm Travels

Palms are difficult to interpret from herbarium specimens. Furtado made

two major expeditions to study palms in the field during which he collected

specimens that are deposited in the Singapore Botanic Gardens’ Herbarium

(SING).

In 1932, he joined the Clemens Expedition to Kinabalu and spent six

weeks between March and April making a particular study of palms and

aroids. He made extensive collections, which included several new species

he subsequently described.

In 1937, he spent a month (20 May to 19 June) travelling down the

west coast of Peninsular Malaysia collecting in Kedah, Perak, west Pahang

and Negri Sembilan. He also made extensive collections from SW Johore.

In addition to studying palms in the field, he also traveled to see type

specimens not available at SING. Taking half-pay for a year, from 17 April

1933 to 14 January 1934, he worked on the collections at the Berlin Botanic

Gardens. It was here he came into contact with Professor M. Burret, the

leading world specialist on palms at that time, and which explains why

subsequently Furtado published several articles in Feddes Repertorium. He

then proceeded to work in London from 16 January 1934. During this

period, he also took the opportunity to visit the herbaria at Vienna, Florence

and Paris. He resumed his duties at SING on 1 May 1934.

Another important visit was to the Bogor Botanic Gardens where he

spent a month in 1936 (April and May) studying the living palm collection

there, many of which had been named by O. Beccari, another great specialist

on Malesian palms. He was able to make herbarium specimens of several

of these, which are deposited at SING.

In the summer of 1967, Furtado went to Lisbon for health reasons.

Ever a scientist, he kept himself busy by carrying out a herbarium study of

the genus Hyphaene, with the result that his first publication on this genus

appeared in Garcia de Orta.

Ruth Kiew & Dennis V. Johnson

C.X. Furtado 149

Appendix 3. A History of My Work on Palms

(extract from C.X. Furtado’s DSc. dissertation)

When I came to Malaya in 1923, I began to study the biology of palms and

aroids cultivated in the Botanic Gardens, Singapore; but I could not make

much headway because the palms in this garden were badly named and

the herbarium material of the cultivated palms was totally unworked. A

difficulty of another nature arose from the fact that little of our material of

wild or cultivated palms had been submitted to specialists working in

Europe, nor was there in the Singapore herbarium any authentic material

received by way of exchange from any other institutions. a

Thus the only alternative left was to undertake systematic studies

independently of any external help or advice, and at such odd periods as

my duties of the Field Assistant would allow. This study obliged me_not

only to resuscitate my knowledge of Latin to the extent of being able to

read general and old systematic accounts of palms, but also to learn Italian

in order to read BECCARI’s work, Spanish to read the early works

published in Manila, French to read the accounts on palms from Indochina

and others written in that language and German to enable me to consult

systematic works in that language.

Not until I had made sufficient progress in these languages and

systematic studies did I realize yet another difficulty which arose because

of the lack of any material in the Singapore herbarium worked out by a

specialist on palms. This last difficulty was particularly felt whenever I had

reason to doubt the correctness of Mr. RIDLEY’s interpretation of species,

and also when the original descriptions were insufficient to guarantee a

correct identification of the species concerned.

In this dilemma, the Singapore Government came to my rescue,

after it was recognized that I had made sufficient progress in my studies in

palms, aroids and languages. I was sent to Europe in 1933 with letters to

several botanical institutions recommending them to give me facilities to

study the types of palms in their care.

I chose the Berlin Botanic Gardens as my headquarters in Europe

because Prof BURRET had promised to assemble there the types of

MARTIUS (from the Munich herbarium) and because it would be easy to

obtain there specimens I might require from most herbaria in Europe.

Further, the Berlin herbarium has a large number of holotypes, haptotypes

and neotypes of the Indo-Malaysian palms, nearly all the Indo-Malaysian

material there had been worked out by BECCARI. Further I expected the

cooperation of Prof BURRET in naming many of the American palms in

the Singapore Gardens, on the systematics of which I had worked very

150 Gard. Bull. Singapore 51 (1999)

little, and to learn from him about the new concepts he had introduced in

the classification of American palms. Also I hoped to improve my

knowledge of the German language and to profit by the discussions with

Prof BURRET who was the only palm specialist then known to the world.

During my study of the genus Areca and Palmae Malesicae III

(Calami), the results of which were prepared mainly in Europe, I naturally

profited much from the discussions with Prof BURRET on the subject of

nomenclature, and still more so from similar discussions with Prof H.

HARMS, also of Berlin, to whom I was introduced by Prof BURRET on

the plea that he was the better qualified to solve my nomenclatural

difficulties. And these discussions were specially valuable to me at the

time because the International Rules of Botanical Nomenclature ed. 3,

adopted at the International Botanical Congress, Cambridge, England,

1930, had not then been published. And no one was better able to discuss

those Rules that Prof HARMS, editor-in-chief of edition 3 of the Rules.

That I have not slavishly followed the opinions on nomenclature, so

kindly placed at my disposal in Berlin by the two professors named, may

be gathered from my papers and, once the Rules were published in 1934, I

commenced my own independent commentary upon them.

Gardens’ Bulletin Singapore 51 (1999) 151-161.

The Singapore Botanic Gardens Herbarium — 125 Years

of History

RUTH KIEW

The Herbarium, Singapore Botanic Gardens,

1 Cluny Road, Singapore 259569

Introduction

The Herbarium dates back to the establishment of the Botanic Gardens on

its present site. By 1875, H. J. Murton already had a collection of dried

specimens, which was temporarily housed in his office until the proposed

herbarium building was erected, and from the start he established a library

having ordered from England a ‘good collection of standard botanical

works’ (Annual Report for 1875). He reported that by 1879 the herbarium

comprised 3,000 named specimens (Annual Report for 1879). Most of his

collections were made from Singapore, Perak, Selangor and Malacca but

he acquired valuable old herbarium specimens, mostly of Indian and

Nepalese plants, that had been in J. Collin’s possession in Singapore but

which had originally been part of Ward’s herbarium held by the Linnaean

Society in London. These included Wallich’s specimens collected from

Penang, Malacca and Singapore, as well as the oldest specimens in the

Singapore herbarium, those collected by Moravian Missionaries in 1790

(Annual Report for 1889).

The First Phase

When N. Cantley was in charge of the Gardens, a new Office and Herbarium

was erected in 1882 at the cost of $1,500. This Herbarium occupied 650 sq

ft with a further 200 sq ft for a drying room (Annual Report for 1882).

(Drying specimens was always a problem in the humid tropics until the

advent of hot air drying ovens). In 1884, a herbarium keeper was employed

to cure and mount specimens as well as to look after the library (Annual

Report for 1884). From the efforts of Cantley and his collectors, such as

M. V. Alvins who made an interesting collection of ethnobotanical

specimens from the Malacca and Negri Sembilan region, the herbarium

collection was fast expanding and filled 18 cabinets. In addition, a small

stove had been acquired to ‘keep away the damp from the books, as well

as the herbarium specimens’ (Annual Report 1885). Duplicates were already

152 Gard. Bull. Singapore 51 (1999)

being sent to Kew. Even at that early date, Cantley reported in 1885 that

“The collecting of herbarium specimens, in order to afford material for

scientific investigation... had not been forgotten” (Annual Report for 1885).

H. N. Ridley took over the Herbarium in 1888 but his efforts were

hampered by the lack of a herbarium keeper (Annual Report for 1888). By

then, the Herbarium was regularly receiving specimens from C. Curtis,

Superintendent of the Botanic Garden in Penang; from G. King of the

Calcutta Botanic Garden including in 1892 a valuable gift of 5,000 specimens

of early collections from Peninsular Malaysia, such as those of Father B.

Scortechini, Kunstler (often referred to on herbarium labels as ‘King’s

Collector’) and L. Wray; the British Museum, which in 1895 gave 1,503

specimens collected by Wallich and Hance; as well as from Kew (Annual

Report for 1889).

Ridley’s plans for the Herbarium were ambitious: “I am ... attempting

to get specimens of every plant for each of the States [of the Malay

Peninsula], and the similarity of the flora induces me to add those of

Sumatra, Borneo and adjacent islands’ (Annual Report for 1889). He

therefore set about an ambitious collecting programme to many parts of

the Peninsula, then very remote. Many collecting trips were made when he

was on leave and could be free of his duties as Director. In addition, he

collected in Sumatra, Borneo, Java, peninsular Thailand and Christmas

Island (the last was at that time under the jurisdiction of Singapore). He

collected literally thousands of specimens and is credited with discovering

more than a thousand new species. |

In addition to his own collections, he continued to receive duplicate

specimens from the British Museum, Calcutta and Kew, as well as many

collections made by Europeans living locally, who were attached to

museums, the geological survey or were planters. He also regularly received

specimens from G.D. Haviland and G. F. Hose in Sarawak. Indeed, it was

extremely fortunate that Hose gave Singapore a duplicate set of his

collections as, on returning from home leave, he discovered his personal

collection had been totally consumed by termites (Steenis-Kruseman, 1950).

Later Ridley regularly received specimens collected from the Philippines,

Java and Sumatra.

In addition, Ridley built up a collection of wood, especially important

in those days when the timber trees were not well known, as well as a

collection of the less durable or more fragile specimens, which were

preserved in spirit.

And so the Herbarium grew. In 1890, the large quantities of plants

that Ridley was collecting required a specially constructed drying room

where specimens were dried on a corrugated iron platform heated from

beneath by chatties full of charcoal (Annual Report for 1890). In 1892 it

History of SING Herbarium 153

was necessary to spend $500 to enlarge the Herbarium (Annual Report for

1892). In 1889 Tassim Daud was employed as herbarium keeper followed

by Ahmad Kassim in 1895. With a herbarium keeper, the specimens could

be sorted into folders and the Herbarium to be rearranged to Ridley’s

satisfaction following the system used in Hooker’s The Flora of British

India (which included the Malay Peninsula). In addition, Ridley was able

to report that “the vermin which were very abundant have been destroyed”

(Annual Report for 1589).

By 1900, Ridley was boasting that the Herbarium was “now without

doubt the finest for the Malay Peninsula plants in the world and contains

many types and cotypes of plants from the Peninsula, Borneo, Sumatra

and Siam” (Annual Report for 1900). In 1910, the Herbarium was further

strengthened with the addition of Curtis’s collection when the Herbarium

of the Penang Botanic Garden was transferred to Singapore (Annual Report

for 1910).

In 1904 and 1905, the Herbarium moved to a new building (Annual

Report for 1905), but Ridley noted that it was not satisfactory as ‘the

whole building was very leaky and damp for a great part of the year’. In

1907, the Herbarium was one night invaded by termites. Fortunately, their

tunnels were discovered before they had got into the cabinets. In haste the

whole collection was re-poisoned taking four men from the Public Works

Department four months to complete (Annual Report for 1907).

In 1913, specimens collected from the Botanic Gardens of Penang

and Singapore were removed from the general collection and kept

separately (Annual Report for 1913). This ‘Gardens Herbarium’ was

important for checking that the names of plants grown in the Gardens

were correctly identified and could also be used as a basis for compiling a

catalogue of plants grown in the Gardens. This was very important for, as

Cantley had scathingly commented in the Annual Report for 1582: “A

catalogue purporting to be a list of plants contained in the Gardens was

published in 1879 and a supplement in 1880. But on my referring to the

catalogue for the names of the trees they could not be found: I next sought

for the plants contained in the list, and with the exception of a few was

equally unsuccessful”. However, although he began to catalogue the plants

it was not until 1912 that the first reliable Gardens’ catalogue was produced

(Anderson, 1912). (The next catalogue, The Checklist of Cultivated Plants

of Singapore Botanic Gardens by Tay et al., 1995, appeared very much

later).

Due to Ridley’s efforts, the collection was sufficiently comprehensive

to embark on taxonomic revisions for the “Materials for the Flora of the

Malay Peninsula” project under King at Calcutta. For this purpose large

quantities of specimens were sent as duplicates or on loan to specialists

154 Gard. Bull. Singapore 51 (1999)

who revised particular families. For example, in 1894 specimens were sent

to O. Beccari in Naples who was revising the palms. Ridley himself

undertook to revise the monocotyledons (Ridley, 1907). After Ridley

retired, he continued to work indefatigably at Kew to produce the five-

volume Flora of the Malay Peninsula (Ridley, 1922-1925).

The Second Phase

Ridley was succeeded as Director by I. H. Burkill, best remembered for

his two-volume A Dictionary of the Economic Products of the Malay

Peninsula (1.H. Burkill, 1936), for which the herbarium collection with

notes on local uses recorded on the herbarium label was an important

source of information. In addition, Burkill laid a sound basis for the scientific

investigation of the Malayan flora by creating the post of Curator of the

Herbarium and employing several talented taxonomists. With Ridley’s

Flora, which covered seed plants, complete, Burkill shifted the emphasis

onto the much neglected lower plants. In 1922, R.E. Holttum was appointed

Assistant Director and began to work on ferns; in 1923, C.X. Furtado was

appointed Assistant Botanist in charge of the naming of the living collection

and cultivated plants; in 1924, M.R. Henderson filled the new post of

Curator of the Herbarium (Table 1); and in 1929, E.H.J. Corner joined the

staff as Assistant Director and worked on fungi.

After the hiatus of WWI, when paper for mounting specimens was

not available, three women were employed to mount specimens (Annual

Report for 1922) and by 1924 the backlog was cleared and under the

direction of the new curator, the Herbarium was rearranged to follow

Ridley’s Flora, the folders following the Flora’s numbering of the families,

genera and species. In the course of this work, species not recorded in the

Flora came to Henderson’s attention, which he published a series of papers

(Henderson, 1927, 1929, 1930a,b). In 1926, the herbarium of the Federated

Malay States Museum was handed over and incorporated into the Singapore

collection (Annual Report for 1926).

In 1922, an active collecting programme started and continued up to

the Japanese Occupation. Burkill made extensive collections from both

Taiping Hills and Fraser’s Hill, Corner collected all over the Peninsula for

his book on Wayside Trees (Corner, 1940) as even many of the village fruit

trees were still improperly known (Annual Reports for 1936), Furtado

specialised in and made collections of aroids and palms (Johnson and Tay,

1999), Henderson concentrated on the limestone flora and visited many

hills, mountains and islands to collect, Holttum made collections notably

from Mount Kinabalu and several mountains in the Peninsula. In all these

History of SING Herbarium h5d

expeditions, the plant collectors employed in the Herbarium played an

important part. Corner started a dried collection of macrofungi having

successfully experimented with methods to keep them permanently free of

mould and beetle attack (Annual Report for 1931). Corner is also famous

for using beruk monkeys (pig-tailed macaques) to collect botanical

specimens (Corner, 1992).

C.E. Carr, a planter by profession, built up an excellent collection of

orchids, not only from the Peninsula but also 800 specimens from Mount

Kinabalu. After his early death in New Guinea, the Botanic Gardens bought

in 1936 his personal herbarium of over 3,000 orchid specimens, which

included nearly all his locally collected material, as well as many flowers in

spirit, besides a set of the 7,000 orchid specimens from New Guinea that

the Singapore Herbarium was given in return for processing them. This is

an invaluable resource for the study of Malesian orchid taxonomy.

Thousands of specimens were collected each year and large quantities

of duplicates were sent to regional and European herbaria; similar quantities

were received in exchange, such as H.H. Bartlett’s Sumatran specimens, a

set of Ledermann’s New Guinea specimens, and quantities of specimens

from the Forest Department, British North Borneo (Sabah). (The latter’s

herbarium was later to burn down).

By January 1930, the new herbarium was completed with two floors.

The upper with a gallery housed the herbarium of cultivated plants, museum

specimens and the spirit collection, while the ground floor was occupied by

the office and the general collection housed in seven double ranks of 12

two-unit teak cases (Annual Report for 1930).

The herbarium and library and their collections survived undamaged

even though the line of fire during the invasion by Japanese forces came

within half a mile of the Gardens’ boundary. (Some specimens, however,

on loan to German institutions did not survive). In 1948, the Hongkong

Herbarium collection arrived from Penang, where it had been sent for safe

keeping in 1940. (This was indeed fortunate as the building where it was

housed in Hongkong was destroyed during the war). After the war, the

collection was returned intact to Hongkong.

The Japanese Occupation brought an end to active collecting and

exchange of specimens. Both Corner and Holttum were interned in the

Gardens and, freed from administrative duties, devoted their time to

research, which turned out to be an extraordinarily productive time resulting

in taxonomic revisions and even a new theory of tropical botany — Corner’s

Durian Theory (Corner, 1949).

After the war, a notable addition to the herbarium was the acquisition

through repatriation of 5,000 specimens from the personal collection of Z.

Teruya, a Japanese planter once resident near Kota Tinggi, Johore. As

156 Gard. Bull. Singapore 51 (1999)

Secretary to the Japanese Planters Association, he had the opportunity to

travel all over the Peninsula. His collection was, however, only incorporated

in to the main collection in the 1960s (H.M. Burkill, pers. comm.).

The Third Phase

Furtado, Henderson and Holttum resumed work in the Gardens after the

Japanese Occupation ended. Furtado continued to work as Assistant

Botanist until he retired in 1952, when he was reappointed as Botanist.

(Even after this second retirement he continued to work in the Herbarium).

After Holttum left, Henderson became Director and, in 1948, J. Sinclair

was employed as Curator of the Herbarium (Table 1). By a curious twist

of fate, in 1954 exactly thirty years after Henderson had found his post in

Kuala Lumpur threatened and I.H. Burkill had created the post of Curator

for him, H.M. Burkill (I.H. Burkill’s son) found himself in the same situation

when funding for his post as botanist at the Rubber Research Institute in

Kuala Lumpur dried up. The posts of Assistant Director and the Botanist

both being vacant at that time, Henderson was able to offer H.M. Burkill a

secure post (H.M. Burkill, pers. comm.). H.M. Burkill immediately found

a niche in taking up the study of seaweeds, at that time a much neglected

field and one of economic importance.

With the setting up of the new university in Singapore, local botany

graduates came onto the job market. The first to be employed was Chew

Wee Lek in 1956, followed by Chang Kiaw Lan in 1959, by Hardial Singh

in 1963, and by Geh Siew Yin in 1970.

When H.M. Burkill became Director in 1957, he found the

administrative inflexibility of moving staff between the posts of Assistant

Director, Keeper of the Herbarium and Botanist made them difficult to fill

(H.M. Burkill, pers. comm.). He therefore arranged for them to be re-

graded as three botanists posts, so when Chang took over the curation

duties of the herbarium after Chew left, her post was designated Botanist

(Keeper).

After WWII, collecting in Peninsular Malaysia was severely restricted

by The Emergency. Sinclair continued to collect in Singapore and Johore.

Soon regular collecting trips were made to Sarawak, first by the Director,

J.W. Purseglove, then by Sinclair, Chew Wee Lek, Chang Kiaw Lan and

Geh Siew Yin. Chew was an active member of both Royal Society

Expeditions to Gunung Kinabalu. Burkill built up a collection of seaweeds.

After Independence, the Forest Departments of Peninsular Malaysia,

Sabah and Sarawak began to build up their own herbaria and regularly

send duplicates to the Herbarium. In contrast, the collecting activities by

History of SING Herbarium 157

Table 1. Curators/Keepers of the Herbarium

1924—1946 Murray Ross Henderson

The post of Curator of the Herbarium was created by I. H. Burkill in 1924 for

M.R. Henderson whose post in Kuala Lumpur had been abolished in a

government reorganisation. Henderson held the post (except during the war years

when he was in South Africa) until 1946, when he became Director of the Botanic

Gardens.

1948—1965 James Sinclair

In 1948, J. Sinclair was appointed Curator, a post he held until his retirement in

1963. He was then re-employed until 1965 to cover the duties of local staff

overseas for higher studies, continuing in an honorary capacity until 1967. It was

during his time in 1955 that, according to the Annual Reports, there was a change

in name from Curator to Keeper of the Herbarium but the reason for this was not

stated.

In 1959, the posts of Assistant Director, Keeper of the Herbarium, and

Botanist were all re-graded as Botanist, so that in 1960 the post became Botanist

(Keeper of the Herbarium).

1965—1970 Chew Wee Lek

In 1956, Chew Wee Lek was the first local graduate to be appointed as Botanist,

and, in 1965 on his return with a PhD from Cambridge, he took on the duties of

Keeper.

1970—1987 Chang Kiaw Lan

Chang Kiaw Lan, the second local graduate to work in the Herbarium, was

appointed as botanist in 1959 to take charge of the collection of fungal specimens.

In 1970, when Chew became Director, she took over his duties as Botanist

(Keeper).

1987—1993 Mrs Ng Siew Yin (née Geh)

When Chang retired in 1987, there were no taxonomists employed in the

Herbarium and the Keeper’s post effectively became defunct. Mrs Ng then took

on the duties of Keeper in addition to her own as Assistant Commissioner of the

Botanic Gardens.

1993—1996 Chin See Chung

Under the aegis of NParks, the post of Keeper of Herbarium and Library was

revived and held by Chin See Chung from 1993 until 1996 when he became

Director of the Gardens.

1997—present Ruth Kiew

In 1997, R. Kiew was appointed.

158 Gard. Bull. Singapore 51 (1999)

staff in Singapore declined and the SFN (Singapore Field Numbers) series,

which was started in 1924 and used by all staff to number their collections,

was discontinued at the end of 1958.

By 1949, the herbarium was already reported as “filled to overflowing”

in spite of 11 new units of herbarium cases being purchased. It was clear an

extension was urgently required (Annual Report for 1949). At that time,

the Herbarium ranked in size second in Asia and tenth in the world.

By the late 1960s, the herbarium building had become derelict. In

H.M. Burkill’s own words:

“The Singapore herbarium gave me endless anxiety. When the second

floor was added, iron girders (?cast iron, not steel) were bricked on to the

outside of the lower floor to carry its weight. I suspect, but without means

of knowing for sure, that the contractor used seashore sand not washed

free of salt for cement and mortar, resulting in serious corrosion of the

metal. There must have been settling of the upper floor on to the brickwork

of the lower. There was bulging and glass fillets cemented on were constantly

cracking. Year by year estimate proposals for a new building were rejected

by the Treasury. When the UNESCO Visiting Committee for Tropical

Herbaria came to Singapore I reported this most serious situation, and in a

courtesy call on high authority, they must have touched a receptive button

for soon after I received an urgent communication from the Director of

Public Works Department (PWD) to the effect that he would ‘repair’ the

building provided that I accept the repaired building standing on the same

outline footings of the old. The PWD would remove the roof and demolish

the rest down to the ground and build again up from there replacing the

old roof on top. This was the crux for retaining the original floor dimensions.

Into the bargain a third floor would be built in thus meeting my need for

more storage space. The work would be carried out on the PWD’s Upkeep

of Buildings vote, whereas any change in the ground dimensions would

have required a new OCSE (Other Charges Special Expenditure) vote

from the Treasury, application for which had been regularly rejected. In

effect the PWD gave me a new and extended building on a ‘repair’ vote!

Anyhow the situation was serious and I was given three weeks notice to

clear the building. No storage space was offered. Hamish Gilliland, Professor

of Botany, University of Singapore, came to my rescue with the loan of a

vacant Nissen hut in the University grounds. The ensuing massive and

complete clearout had some rewards. Many skeletons, put aside and long

since forgotten came to light, and were duly dealt with. Perhaps any

organisation with an accumulation of materials ought to have an upheaval

like this every few years!”

(H.M. Burkill, pers. comm.)

History of SING Herbarium 159

So through H.M. Burkill’s energies, a new three-floor herbarium

building was completed in 1964, to which was added in 1968 a new wing,

which also housed the library. In 1972, a third storey was added to the

wing, but what the herbarium gained from occupying the new floor on the

wing, it lost as the ground floor of the main herbarium building was given

over to administration. The Herbarium has remained little changed since

then, apart from experiencing an ever-increasing shortage of space as more

specimens are added to the collection.

In 1967, with the launching of the Garden City Campaign, the

botanists attached to the Herbarium were directed to apply their research

to more practical ends. On several occasions, H.M. Burkill deplored this

change, which was based on the misconception that a botanic garden could

function to international standards without the backing of a strong botanical

research programme (H.M. Burkill, 1993). He pointed out that the applied

fields of economic botany, education and conservation all need a strong

botanical base.

In 1973, the Botanic Gardens came under the administration of the

Parks and Recreation Branch of the Public Works Department. The stated

aims for botanical work included maintenance of the Herbarium and library,

collection of plants for the Herbarium, identification, botanical publication

and the preservation and maintenance of Nature Reserves in Singapore

(Tinsley, 1989). With the botanists’ posts transferred away, the Herbarium

was just able to tick over. In 1973, Mrs Ng moved to the Parks and

Recreation Branch as Assistant Commissioner, followed in 1975 by Hardial

Singh, leaving only Chang in the Herbarium. After 1970, for twenty years

no new taxonomists were recruited.

The Fourth Phase

The formation of the National Parks Board (NParks) in 1990 as a statuary

board under Tan Wee Kiat, Executive Director of NParks and Director of

the Botanic Gardens (the first for 18 years!) brought a wind of change as

Tan sought to restore a proper balance between research, education,

conservation and recreation and to revive systematic botany. Three new

posts for taxonomists (the Keeper of the Herbarium and Library being

one) were created. From 1990 to 1996, Tay Eng Pin held one post. In 1993,

Chin See Chung was appointed as Keeper to be followed in 1997 by the

seventh Keeper, Ruth Kiew (Table 1). She presented a set of about 2000

specimens, mostly forest herbs and plants collected from limestone and

mountains in Malaysia. In 1998, the second post went to I.M Turner

(Assistant Director, Horticulture) and the third is reserved for an orchid

taxonomist.

160 Gard. Bull. Singapore 51 (1999)

A new building for research, which included a spacious new

Herbarium, was planned. The last was overtaken by events when NParks

was enlarged with the merger with the Parks and Recreation Division, and

the research building was transformed into Gateway, the new headquarters

for NParks.

So it is that at present the Herbarium still occupies the old buildings,

although some improvements have been made in air-conditioning the

library, office and type collection. The Herbarium holds at least 600,000

specimens of which more than 4,000 are types and specimens are constantly

added through staff collections and by exchange. Space increasingly is a

constraint as the sturdy cupboards, many of solid teak from the days before

plywood, are filled to bursting, the overflow being accommodated in Leiden-

type boxes that double as cupboard space. Computerisation of the collection

is proceeding slowly using BRAHMS (Botanical Research and Herbarium

Management System) for databasing, label generation and bar-coding.

Concurrently plans are afoot for a new building that will take the Herbarium

into the new millennium, so with modern facilities, adequate staffing and

funding, the Herbarium aims to recapture its former reputation as a centre

of excellence in tropical taxonomy.

Acknowledgements

This paper is largely based on the illustrated account that appeared in

Gardenwise (1999) 13: 6-9. It is republished here with additions at Dr S.C.

Chin’s suggestion, Gardenwise being a more ephemeral publication. The

author is most grateful to Chin See Chung and Ng Siew Yin for helping to

fill gaps in the records and especially to H.M. Burkill for his time and

patience in answering queries and supplying information on the period he

served as Director of the Singapore Botanic Gardens.

References

Anderson, J.W. 1912. Index of Plants. Botanic Gardens, Singapore.

Methodist Publishing House, Singapore.

Burkill, H.M. 1993. The Singapore Botanic Gardens: A reputation to cherish

and preserve. In: Proceedings of The Singapore Botanic Gardens 130"

Anniversary Seminar, Singapore Botanic Gardens, Singapore. Pp.10-15.

Burkill, LH. 1935. A Dictionary of the Economic Products of the Malay

Peninsula. Gov. Straits Settlements and Federated Malay States, London.

History of SING Herbarium 161

Corner, E.J.H. 1940. Wayside Trees of Malaya. Government Printing Office.

Singapore.

Corner, E.J.H. 1949. The Durian Theory or the origin of the modern tree.

Annals Botany. 13: 367-414.

Corner, E.J.H. 1992. Botanical Monkeys. The Pentland Press, Edinburgh.

Henderson, M.R. 1927. Additions to the flora of the Malay Peninsula.

Gardens’ Bulletin, Straits Settlements. 4. 48—56.

Henderson, M.R. 1929. Additions to the flora of the Malay Peninsula.

Gardens’ Bulletin, Straits Settlements. 4: 411-414.

Henderson, M.H. 1930a. New species from the Malay Peninsula. Gardens’

Bulletin, Straits Settlements. 5: 72—80.

Henderson, M.R. 1930b. List of additions to the flora of Malay Penisula.

Gardens’ Bulletin, Straits Settlements. 5: 93-98.

Hooker, J.D. 1875-1897. The Flora of British India. Reeve & Co., London.

7 volumes.

Johnson, D.V. and E. P. Tay. 1999. C.X. Furtado (1897-1980): Contribution

of the study of palms. Gardens’ Bulletin, Singapore. 51: 141-150.

Ridley, H.N. 1907. Materials for a Flora of the Malay Peninsula. Methodist

Publishing House, Singapore.

Ridley, H.N. 1922-1925. The Flora of the Malay Peninsula. Reeve & Co..,

London. 5 volumes.

Steenis-Kruseman, M.J. van. 1950. Malaysian Plant Collectors and

Collections. Flora Malesiana. Series 1, Volume 1.

Tay, E.P., Nura Karim and S.C. Chin. 1995. Checklist of Cultivated Plants,

Singapore Botanic Gardens. Singapore Botanic Gardens, NParks Board,

Singapore.

Tinsley, B. 1989. Visions of Delight. Singapore Botanic Gardens.

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Gardens’ Bulletin Singapore 51 (1999) 163-174.

On Some Species of Pandanus and Freycinetia

(Pandanaceae) in Micronesia

KIM-LANG HUYNH

Phanerogamy Division, Botanical Institute,

University of Neuchatel, CH-2007 Neuchatel, Switzerland

Abstract

Two new species of Pandanus from the Caroline Islands are described: P. amissus (Kosrae

Island) and P. lorencei (Palau Island). New data on P. kanehirae and Freycinetia villalobosii,

both from Palau, are also reported. The staminate flower of P. whitmeeanus is described

for the first time. The taxonomic relationships of P. whitmeeanus within the genus Pandanus

are reassessed.

Introduction

The taxonomy of the family Pandanaceae in Micronesia was widely

investigated by Martelli (1934) and Kanehira (1935a-f, 1936a-b, 1937), who

described several new species in Pandanus Parkinson and some in

Freycinetia Gaudich. Recently, several specimens of Pandanus and

Freycinetia were collected in Micronesia by botanists of the National

Tropical Botanical Garden, Lawai (PTBG). In the course of their study,

two unknown species of Pandanus were discovered, and new data on some

previously described species of Pandanus and Freycinetia were obtained.

The new species and the new data are described below.

New Species

1. Pandanus amissus Huynh, sp. nov. (sect. Lophostigma)

Arbor 7-8 m alta, radicibus gralliformibus longissimis suffulta. Folia 2.5—3 m longa, 17 cm

lata in medio 15 cm prope basim, ensiformia, apicem versus abrupte attenuata, in basi

amplexicaulia; lamina in sicco brunnea, subcoriacea; plicis vix visibilibus; venis

longitudinalibus plerumque utrinque distinctis sed transversalibus omnibus invisibilibus;

denticulis marginalibus e 14-15 cm supra basim ad apicem praesentibus, omnibus antrorsis,

plerumque parallelis, generaliter 1.5-2 mm longis sed inferne paulo brevioribus, prope

basim 2-4 mm sursum versus 5-15 mm inter se separatis; denticulis costalibus in circiter 2/3

superis praesentibus, antrorsis, parallelis, + tam longis latis quam marginalibus proximis

sed multo magis inter se separatis (2-7 cm); vagina c. 14 cm longa 17 cm lata, utrinque fere

tote non nervata. Infructescentia monosyncarpica; syncarpio globoso, 22-23 cm diametienti;

pedunculo c. 45 cm longo. Drupae c. 10 cm longae 3.7 cm latae 3 cm crassae, maturitate in

164 Gard. Bull. Singapore 51 (1999)

6/10 superis liberae; stigmatibus unicis, c. 3 x 2 mm, reniformibus, verticalibus, c. 3 mm

stipitatis; stylo fere horizontali, atrobrunneo, corneo, subnitido, in vertice atrobrunneo fere

plano pileo excentrico, extrinsecus stigma ferenti; pileo c. 4 cm alto, convexiuscule pyramidali,

laevi, corneo sed non nitido, 5 vel 6 angulato, angulis vix visibilibus, non prominentibus nec

acutis; endocarpio c. 5 cm alto in axe (processu endocarpico supra loculum seminalem

incluso), apice ad c. 1.5 cm infra basim stigmatis basi c. 3.5 cm supra basim drupae: loculo

seminali ovoideo, c. 2.2 x 1.5 cm, apice plano, centro prope medium, parietibus endocarpicis

tenuis, superne processu endocarpico excentrico c. 1.5 cm longo productis; tubo germinationis

c. 7 mm longo infra loculum seminalem; mesocarpio supero percopiose medulloso,

abundanter et tenuiter fibroso, in medio caverna subsphaerica praedito; mesocarpio infero

copiose et grosse fibroso, ut videtur non medulloso. — Figs. 1-3.

Typus: Flynn & Lorence 6320 (holo PTBG, iso PTBG); Caroline Islands,

Kosrae Island, Tafunsak district, alt. c. 3 m, 13 September 1997; disturbed

lowland forest; large tree of 25 ft, with prop/stilt roots, leaves glossy dark

green, channeled; spines brown tipped, infructescence on pendulous

peduncle c. 18 inches long, head globose, with 100-200 drupes, these

glaucous blue-green, ripening with brownish-yellow base.

Tree 7-8 m tall, with very long stilt roots. Leaves 2.5—-3 m long, 17 cm wide

at the middle and 15 cm wide near the base, ensiform, abruptly attenuate

towards the apex, amplexicaul at the base, brown when dry, subcoriaceous;

pleats just visible; longitudinal veins mostly distinct on both surfaces, but

all transverse veins invisible; marginal prickles present from 14-15 cm

above the base up to the apex, all antrorse, mostly parallel, in general 1.5-

2 mm long but a little shorter below, 2-4 mm apart near the base, 5-15 mm

above; costal prickles present in about the upper 2/3, antrorse, parallel,

more or less as long and wide as the closest marginal prickles but much

more remote (2—7 cm); sheath c. 14 cm long, 17 cm wide, almost entirely

not nerved on both surfaces. Infructescence monosyncarpic; syncarp

globose, 22-23 cm in diameter; peduncle c. 45 cm long. Drupes c. 10 cm

long, 3.7 cm wide and 3 cm thick, free in the upper 6/10 when mature;

stigmas single, c. 3 x 2 mm, reniform, vertical, stipitate by c. 3 mm; style

almost horizontal, dark brown, corneous, subnitid, eccentric on the dark

brown, almost flat vertex of the pileus, bearing the stigma at the outside:

pileus c. 4 cm high, pyramidal but a little convex, smooth, corneous but not

nitid, 5—6-angled, the angles just visible, not prominent, not acute; endocarp

c.5 cm high on the drupe axis (incl. the endocarpic process above the seed

locule), the apex at c. 1.5 cm below the stigma base, the base at c. 3.5 cm

above the drupe base; seed locule ovoid, c. 2.2 x 1.5 cm, the apex flat, the

centre near the middle, the endocarpic walls thin, prolonged above in the

eccentric, c. 1.5 cm long, endocarpic-process; germination tube c. 7 mm

long below the seed locule; upper mesocarp very abundantly medullous,

abundantly and finely fibrous, with a subspheric cavern in the middle;

Pandanus and Freycinetia in Micronesia 165

a Gm

5, 6 4

2 cm 4

Figures 1-6: Pandanus amissus (1-3) (Flynn & Lorence 6320, holotype) and P. lorencei (4-6)

(Lorence et al. 8310, holotype):

1: Drupe in lateral view (the stigma, vertical and in profile, is at top right; dotted: pileus;

arrow: lower limit of free part). 2: Drupe in axial section (black: endocarp; fibres partially

shown in upper mesocarp). 3: Upper part of drupe, showing stigma (densely dotted) in front

view. 4: Upper part of leaf flattened horizontally, viewed by adaxial face. 5: Drupe in lateral

view (dotted: pileus and free part). 6: Same drupe in axial section (black: endocarp).

166 Gard. Bull. Singapore 51 (1999)

lower mesocarp abundantly and coarsely fibrous, apparently not medullous.

— Figs. 1-3.

Kosrae Island is at about 5°18’00"N 163°01’745"E.

Notes: Pandanus amissus is named in reference to its distribution remote

from New Caledonia. New Caledonia is the centre of diversity of sect.

Lophostigma (Brongn.) Warb., with 10 out of the 12 species in this section

(St. John, 1989d), and including the type of the section, P. viscidus (Brongn.)

Solms. Outside New Caledonia, P. syozoi Kaneh. is endemic to Palau and

P. amissus 18s known only from Kosrae Island. P. syozoi is the only species

of the section that has polysyncarpic infructescences (Kanehira, 1936b).

Pandanus amissus has very long prop roots, almost twice a man’s

height, as seen in a photograph of the type plant. In addition, it has the

largest drupes in sect. Lophostigma, up to 10 cm long and 3.7 cm wide,

compared with the New Caledonian species where the drupes do not exceed

6.7 cm in length and 1.6 cm in width (St. John, 1989d). Mature drupes of

P. syozoi have not yet been collected (Kanehira, 1936b).

2. Pandanus lorencei Huynh, sp. nov. (sect. Pandanus)

Arbor c. 2.5 m alta. Folia c. 90 cm longa, medio basim versus 3 cm lata, e circiter medio ad

apicem sensim attenuata, 6-8 cm flagellata; lamina in sicco coriacea, prope basim et infra

apicem revoluta; plicis inermibus, fere e basi ad infra apicem perspicue canaliculatis; venis

longitudinalibus utrinque visibilibus sed transversalibus invisibilibus; denticulis marginalibus

e c. 3 cm supra basim ad apicem praesentibus, omnibus antrorsis, densis (c. 2-5 mm inter se

separatis), in 1/3 infera divergentibus usque ad 1.5 mm longis, ceterum brevioribus minutis

parallelis vel applicatis; denticulis costalibus e c. 6 cm supra basim ad apicem praesentibus,

aliquot infimis retrorsis et multo brevioribus quam marginalibus proximis, ceterum antrorsis

et plerumque tam longis, inferne multo magis separatis superne multo creberrimis medio +

tam separatis; vagina c. 3 cm longa, 4 cm lata in dimidio infero 3.5 cm in apice, fere tote

longitudinaliter nervata in pagina abaxiali, fere tote laevi in adaxiali. Infructescentia

monosyncarpica; syncarpio globoso, 12-13 cm diametienti; pedunculo ignoto. Drupae 4.3-5

cm longae 2.8—4 cm latae 2.43.3 cm crassae, truncatae in apice cuneatae basi, maturitate in

1/2-3/5 superis liberae, 6 vel 7 loculares, carpellis exterioribus latioribus quam interioribus;

stigmatibus verticalibus vel obliquis, reniformibus vel + triangularibus, usque ad 2.5 mm

latis, plerumque centripetis, sulcum 4-5 mm longum in vertice carpelli locatum aspicientibus;

endocarpio c. 2.5 cm alto in axe, 0.5—1 cm peripheria; loculis seminalibus c. 1.5 cm longis,

basi prope medium; mesocarpio supero in 1/3 supera, copiose medulloso, abundanter fibroso;

mesocarpio infero in dimidio infero, medulloso, copiose fibroso. — Figs. 4-6.

Typus: Lorence et al. 8310 (holo PTBG, iso PTBG); Caroline Islands,

Palau (Republic of Belau), Oreor (Koror) State, unnamed limestone rock

island in vicinity of Omekang, W. of Bablomekang Island, 7°08’32"N

134°19°02"E, alt. 5-10 m, 17 February 1998; coastal palm forest; common,

Pandanus and Freycinetia in Micronesia 167

on coastal cliffs; tree, trunk 2.5 m tall with a whorl of branches, leaves

glaucous green, channeled, spines brown, heads 12-13 cm in diameter,

globose, drupes ripening orange then brown.

Tree c. 2.5 m tall. Leaves c. 90 cm long, 3 cm wide from the middle down

to the base, gradually attenuate from about the middle up to the apex, in

6-8 cm flagellate, coriaceous when dry, revolute near the base and below

the apex; pleats unarmed, distinctly channelled almost from the base up to

below the apex; longitudinal veins visible on both surfaces, but transverse

veins invisible; marginal prickles present from c. 3 cm above the base up to

the apex, all antrorse, close (c. 2-5 mm apart), in the lower third divergent

from the margins and up to 1.5 mm long, above shorter, minute, parallel or

applied to the margins; costal prickles present from c. 6 cm above the base

up to the apex, some below retrorse and much shorter than the closest

marginal prickles, the others antrorse and mostly as long as these, below

much more remote than the closest marginal prickles, above much closer,

in the middle more or less as remote as these; sheath c. 3 cm long, 4 cm

wide in the lower half, 3.5 cm wide at the apex, almost entirely nerved

longitudinally on the abaxial surface, almost entirely smooth on the adaxial

surface. Infructescence monosyncarpic; syncarp globose, 12—13 cm in

diameter; peduncle not known. Drupes 4.3—5 cm long, 2.8-4 cm wide, 2.4—

3.3 cm thick, truncate at the apex, cuneate at the base, free in the upper 1/

2-3/5 when mature, 6—7-celled, the outer carpels broader than the inner

carpels; stigmas vertical or oblique, reniform or more or less triangular, up

to 2.5 mm broad, centripetal for most of them, each facing a furrow 4—5

mm long on the carpel vertex; endocarp c. 2.5 cm high on the drupe axis,

0.5—1 cm high at the periphery; seed locules c. 1.5 cm long, the base near

the middle; upper mesocarp in the upper third, abundantly medullous,

abundantly fibrous; lower mesocarp in the lower half, medullous, abundantly

fibrous. — Figs. 4-6.

Notes: Pandanus lorencei is named after David H. Lorence, the principal

collector of this specimen.

In its smaller syncarp with fewer celled drupes and smaller leaves,

P. lorencei is distinct from the other pandans in Palau, especially from the

following species, which may be considered close relatives (Table 1).

168 Gard. Bull. Singapore 51 (1999)

Table 1. Comparison of P. /orencei with close relatives in Palau

Pandanus Syncarp No. cells Length of Length and

diameter in drupe drupes width of

(cm) (cm) leaves (cm)

lorencei 12-13 6-7 4.3-5 90 x3

divergens A 13-20 8 150 x 8

duriocarpus 16 16-18 4.5 (unknown)

erythrophloeus 19 7-13 5-6 150 x 8

okamotoi 18 12-16 7-8 200 x 6-7

Further notes on Pandanus kanehirae

Pandanus kanehirae Martelli is peculiar in that the carpels composing its

drupes are deeply separate (Fig. 7), a feature emphasized by Martelli (1934:

120, Fig. 3). It was by reason of this character in particular that sect.

Palauenses Huynh & B. C. Stone was defined to accommodate P. kanehirae

(Huynh, 1980: 404).

Recently, new data on P. kanehirae were obtained from Lorence

8202 (PTBG), collected on 10 February 1998 in Babeldaob (Palau), the

type locality of this species. As revealed by a field note, P. kanehirae is a

low tree 3-5 m tall. The infructescence comprises 4—6 syncarps (6 on the

infructescence that was collected, 3 of which had been removed); the

peduncle below the syncarps is at least 50 cm long. The structure of the

infructescence is most unusual, and provides further support to the

distinctness of sect. Palauenses. The rhachis of this infructescence is divided

into six branches each bearing or having borne a syncarp (Fig. 9). The two

lowermost syncarps (the 6th and Sth) have their pedicels at the same level,

at c. 15 cm from the base of the uppermost syncarp; one pedicel has no

node and is 6—7 cm long; the other pedicel was partly preserved and the

remnant is c. 3 cm long and shows a node at c. 1.5 cm above the base. The

two subsequent upper syncarps (the 4th and 3rd) also have their pedicels

at the same level, at c. 12 cm from the base of the uppermost syncarp.

These pedicels are separated from the two lowermost pedicels by three

internodes and have no nodes: one is c. 6 cm long, the other c. 4.5 cm long.

The subsequent upper syncarp (the 2nd) has its pedicel at c. 9 cm from the

base of the uppermost syncarp; the pedicel was partly preserved and is

separated from those of the 4th and 3rd syncarps by one internode, and

from the uppermost syncarp by six internodes. Apparently these nodes

have each borne a bract. No bracts were preserved, but bract scars are

Pandanus and Freycinetia in Micronesia 169

observed. What is unusual in this infructescence is that: firstly, two syncarps

may have their pedicels at one and the same level (instead of two); secondly,

a syncarp may be separated from the subsequent syncarp(s) by several

internodes (instead of one; in some polysyncarpic infructescences, two

internodes were observed between the uppermost syncarp and the

subsequent syncarp).

The specimen Lorence 8202 also provides more detailed knowledge

of the drupes of P. kanehirae. These are c. 3.5 cm long, (2.5—) 3-4 cm or

more wide, and 2.5—3 cm thick. Sometimes, two adjacent drupes intimately

fuse along their connate part (they are inseparable from each other), and

viewed from above appear to form a single drupe. There are (1—)3-7 stigmas,

which are vertical, eccentric, and very small. If two stigmas are close, they

are located together on a nitid, slightly sunken area. On those drupes

where it was possible to observe undamaged stigmas, these were found to

be more or less facing the same lateral face: this corroborates a feature

previously described of sect. Palauenses (Huynh, 1980: 404).

The specimen Lorence 8202 also makes possible a re-examination of

leaf characters of P. kanehirae. Leaves are c. 150-160 cm long, 7.5 cm wide

at the middle and 10.5 cm wide near the base, attenuate acuminate in the

apical part, and rigidly coriaceous in the lower half but subcoriaceous in

the upper half. The margins are armed from c. 8 cm above the base up to

the apex; the prickles are close, below up to c. 1 mm long along 20 cm, and

just visible above that level. The midrib is unarmed in the lower third but

armed above with minute prickles. The longitudinal veins are visible on

both surfaces, but not the transverse veins.

Further notes on Pandanus whitmeeanus

Pandanus whitmeeanus Martelli is the type species of sect. Coronata

Martelli, which is the only section of subg. Coronata Martelli (Stone, 1974:

521). It is unique in the genus Pandanus in that its stigmas are all

«centripetally» arranged as described and illustrated by Stone (1974: 488,

Fig. 13C) and St. John (1989a: Fig. 644; 1989b: Fig. 677; 1989c: Fig. 718).

In his taxonomic treatment, Stone (1974) divided the genus Pandanus

into eight subgenera: Pandanus B. C. Stone, Rykia (De Vries) B. C. Stone,

Lophostigma (Brongn.) St. John, Kurzia B. C. Stone, Vinsonia (Warb.) B.

C. Stone, Martellidendron (Pic. Serm.) B. C. Stone, Coronata, and

Acrostigma (Kurz) B. C. Stone. These were distributed in four groups, and

Stone (1974: 468, 527) attributed subg. Coronata and subg. Pandanus to

Group 3. No arguments, however, were given to explain these groupings.

With regard to subg. Coronata in particular, the staminate flower of

170 Gard. Bull. Singapore 51 (1999)

} 10, 11

5 mm

Figures 7-11: Pandanus kanehirae (7-9) (Lorence 8202) and P. whitmeeanus (10,11) (Whistler

6734):

7: Drupe with 5 stigmas in apical view. 8: Same drupe in lateral view (dotted: pileus and free

part). 9: Rhachis of a 6-syncarpic infructescence (dotted: location of the bases of the syncarps).

10: Staminate flower in lateral view (the peltate apex is dotted in margins). 11: Upper part of

staminate flower in axial section.

Pandanus and Freycinetia in Micronesia 171

P. whitmeeanus described below seems to indicate that it does not fall

within any of these groups and that it forms a group of its own.

According to Smith (1979: 478), P. whitmeeanus probably originated

in Vanuatu (the New Hebrides) and was an aboriginal introduction into

Fiji and other archipelagoes. These are Samoa, Tonga, the Cook Islands,

and the Horne Islands (St. John, 1989c: 6). In these islands, it is cultivated

as an economic plant. It is not known if it is cultivated also in the other

archipelagoes in Micronesia. The leaves are used for making mats and

plaiting handicraft articles (St. John, 1989b-c), while the fruits are used for

decoration as garlands (St. John, 1989a). Nevertheless, until recently only

the pistillate plant had been collected (Smith, 1979: 478).

However, a staminate plant of P. whitmeeanus has now been collected

from Tonga: Whistler 6734 (PTBG); Tongatapu Island, village of Ha’asini,

30 May 1989; tree 4 m high; local names paongo (the tree) and hingano

(the inflorescence).

The flowers of this staminate plant consist of a column whose upper

part comprises a peltate apex and a large number of stamens (Fig. 10). The

column varies in length (15-25 mm long), as do its stamen-bearing part (8—

12 mm) and its stalk (S-13 mm). The ratio between these two components

also varies. The peltate apex varies in width (4-6 mm). When viewed from

above, this latter appears clearly sunken (Fig. 11) or more or less flat. Its

shape also varies, being slightly triangular, elliptic, or rectangular. Its margins

are bordered with processes that are either very distinct or just perceptible,

but each generally bears a stamen. The processes are brown, like the peltate

apex, and contrast with the stamens, which are white; the similarity of the

processes with the peltate apex seems to indicate that they are parts of the

latter. In the stalk, 2 or 3 concentric circles of vascular bundles are observed,

but there are no other lignified elements. The stamens are 2—2.5 mm long

and have an acumen c. 0.4 mm long and a very thin filament (c. 0.1 mm).

The anthers have a connective that is lignified, except for the small cells

around the vascular bundle; the pollen-sac walls comprise 1 or 2 layers of

endothecial cells; the pollen is spinulose and oblong, with a pore generally

located on the longer axis.

To date, only two types of staminate flowers with a peltate apex

have been observed in Pandanus. In the first type, seen in sect. Solmsia B.

C. Stone (subg. Rykia), the peltate apex is spinulose at the upper face, and

the stamen filaments are thick and also spinulose (Stone, 1974: Fig. 2D). In

the second type, the peltate apex is smooth as are the stamen filaments,

which are thin. This type is found in some sections of subg. Lophostigma:

sect. Lophostigma (Huynh, 1982: Figs. 1 and 70); sect. Barrotia (Brongn.)

B. Castene (Stone, 19/4 Fic.6C;.Huynh,.1982: Fig, 7; St.John,. 1989d:

Figs. 741b and 744b); sect. Bernardia B. C. Stone (Stone, 1974: Fig. 6A; St.

172 Gard. Bull. Singapore 51 (1999)

John, 1989d: Fig. 748b); and sect. Brongniartia B. C. Stone (St. John, 1989d:

Fig. 756b).

In its smooth peltate apex and smooth, thin stamen filaments, the

staminate flower of P. whitmeeanus (Fig. 10) shows similarity with those of

subg. Lophostigma, and may indicate the existence of a close relationship

between subg. Coronata and subg. Lophostigma. However, the peltate

apex of P. whitmeeanus has marginal processes, which are absent in those

of subg. Lophostigma. This peculiarity of the staminate flower of

P. whitmeeanus seems to indicate that subg. Coronata forms a group of its

own within the genus Pandanus. By contrast, no staminate flowers with a

peltate apex have been observed in subg. Pandanus, which may indicate

that subg. Coronata and subg. Pandanus are not closely related.

Further notes on Freycinetia villalobosii

In F. villalobosii Martelli, new essential specific characters were observed

from specimen Lorence & Flynn 8280 (PTBG), which is a pistillate plant

collected on 16 February 1998 in Babeldaob (Palau), the type locality of

this species. The berries have a thick central sclerenchyma and thin,

fusiform-elliptic fibre-bundles (the presence/absence of a central

sclerenchyma and that of fusiform-elliptic fibre-bundles in berries are two

essential specific characters in Freycinetia: Huynh, 1997: 360). The leaves

are c. 55 cm long and 2 cm wide at the middle and gradually attenuate

from the middle to the apex where they are caudate along c. 2 cm; the

blade is strongly revolute in the upper half; the pleats are unarmed; the

longitudinal veins are very close, and distinct on the abaxial surface, less so

on the adaxial surface; the transverse veins are invisible on both surfaces;

the margins are armed from c. 4.5 cm above the base to the apex, the

prickles are antrorse, near the base up to 1 mm long and 1.5 mm apart,

above that level they are minute and much closer; the midrib is armed

from c. 15 cm above the base to the apex with very close prickles; at

intervals of c. 2-3 mm on the midrib, one prickle is 2—3 times longer and

wider than the adjacent upper and lower prickles, which is an unusual

feature.

Acknowledgements

The author is deeply indebted to the National Tropical Botanical Garden,

Lawai (PTBG) for the loan of materials, and the Swiss National Science

Foundation for a grant to study Pandanaceae (Grant no. 31-45707.95).

Pandanus and Freycinetia in Micronesia WE

References

Huynh, K.-L. 1980. Quelques espéces de Pandanus (Pandanaceae) peu

connues des archipels des Philippines, de Palaos et de Salomon. Candollea

35: 385-419.

Huynh, K.-L. 1982. La fleur male de quelques especes de Pandanus subg.

Lophostigma (Pandanaceae) et sa signification taxonomique,

phylogénique et évolutive. Beitrige zur Biologie der Pflanzen 57: 15-83.

Huynh, K.-L. 1997. The genus Freycinetia Gaudich. (Pandanaceae) in the

Solomon Islands. Candollea 52: 359-382.

Kanehira, R. 1935a. New or noteworthy trees from Micronesia IX.

Botanical Magazine Tokyo. 49: 60-68.

Kanehira, R. 1935b. New or noteworthy trees from Micronesia X.

Botanical Magazine Tokyo. 49: 103-114.

Kanehira, R. 1935c. New or noteworthy trees from Micronesia XI.

Botanical Magazine Tokyo. 49: 185-195.

Kanehira, R. 1935d. New or noteworthy trees from Micronesia XIII.

Botanical Magazine Tokyo. 49: 352-358.

Kanehira, R. 1935e. New or noteworthy trees from Micronesia XIV.

Botanical Magazine Tokyo. 49: 425-431.

Kanehira, R. 1935f. New or noteworthy trees from Micronesia XV.

Botanical Magazine Tokyo. 49: 525-532.

Kanehira, R. 1936a. New or noteworthy trees from Micronesia XVI.

Botanical Magazine Tokyo. 50: 520-525.

Kanehira, R. 1936b. New or noteworthy trees from Micronesia XVII.

Botanical Magazine Tokyo. 50: 541-549.

Kanehira, R. 1937. New or noteworthy trees from Micronesia XIX.

Botanical Magazine Tokyo. 51: 906-913.

Martelli, U. 1934. Pandanaceae. In: R. Kanehira 1934. New or noteworthy

trees from Micronesia V. Botanical Magazine Tokyo. 48: 116-130.

St. John, H. 1989a. Revision of the genus Pandanus Stickman (part 54):

Pandanus of Samoa. St. John’s private edition (Honolulu).

174 Gard. Bull. Singapore 51 (1999)

St. John, H. 1989b. Revision of the genus Pandanus Stickman (part 56):

Pandanus of Tonga, Polynesia. St. John’s private edition (Honolulu).

St. John, H. 1989c. Revision of the genus Pandanus Stickman (part 60):

Pandanus of the New Hebrides. St. John’s private edition (Honolulu).

St. John, H. 1989d. Revision of the genus Pandanus Stickman (part 61):

Pandanus of the New Caledonia. St. John’s private edition (Honolulu).

Smith, A. C. 1979. Flora Vitiensis Nova, vol. 1. Pacific Tropical Botanical

Garden (Lawai, Kauai, Hawaii).

Stone, B. C. 1974. Towards an improved infrageneric classification in

Pandanus (Pandanaceae). Botanische Jahrbiicher fiir Systematik,

Pflanzengeschichte und Pflanzengeographie 94: 459-540.

Gardens’ Bulletin Singapore 51 (1999) 175-177.

The Transfer of Tripetalum cymosum K. Schum.

(Guttiferae) to Garcinia

ILM. TURNER

Singapore Botanic Gardens, 1 Cluny Road, Singapore 259569

AND

PP STEVENS

Missouri Botanical Garden, 2345 Tower Grove Avenue, St. Louis, Missouri 63110,

USA

Abstract

The new combination Garcinia cymosa (K. Schum.) I.M. Turner & P.F. Stevens is made,

based on Tripetalum cymosum K. Schum., for a tree from New Guinea and the Bismarck

Archipelago. The oldest name available for the species is Leuconotis tenuifolia Engl., but

this epithet has already been used for another species of Garcinia. A columnar form of the

tree that is finding success in tropical horticulture, described as forma pendulum Lauterb.,

is also provided with a combination in Garcinia and is lectotypified.

Introduction

When describing the genus 7ripetalum, Schumann (Schumann in Schumann

& Hollrung 1889) included a single Papuasian species, T. cymosum K.

Schum. With the exception of a temporary misplacement of an African

species, this has remained the sole member of the genus. Schumann noted

that 7ripetalum differed from Garcinia in its basically trimerous flowers

and in having stamens in fascicles adnate to the petals. Reduction in number

of floral parts from four or five to three would alone be insufficient to

warrant recognition of a separate genus for a single species. The stamen

character is shared with other species from the Papuasian area, such as

those previously included in Pentaphalangium Warb., also reduced to

Garcinia (Kostermans 1976) and also differing in number of floral parts, as

well as Garcinia warrenii F. Muell. and, at least in the pistillate plant, G. .

platyphylla A.C. Sm. and G. hollrungii Lauterb. All these species have

distinctive, branched latex canals in the lamina.

Thus the separation of Tripetalum from Garcinia cannot be upheld.

Indeed, 7ripetalum has been accepted as a synonym of Garcinia by many

recent authors (e.g. van Steenis 1987, Brummitt 1992, Hoft 1992, Stevens

176 Gard. Bull. Singapore 51 (1999)

1995, Mabberley 1997) but surprisingly the formal transfer of the only

species in the genus, Tripetalum cymosum, to Garcinia has not yet taken

place. As the new combination is required for a forthcoming checklist of

the plants cultivated in the Singapore Botanic Gardens, this deficiency is

dealt with here.

A New Combination in Garcinia

The earliest available name for the species appears to be Leuconotis

tenuifolia Engler. The holotype of Leuconotis tenuifolia was lost in the

destruction of the Berlin Herbarium during the war, but a fragment of the

type is available in the herbarium of Wroclaw University. This consists of a

single leaf, small piece of stem and a drawing of a fruit. The leaf, when

viewed in transmitted light, has a branching system of latex canals typical

of Tripetalum cymosum. The specimen is actually annotated as Garcinia

tenuifolia Engler, but this name was never published and the combination

has subsequently been pre-empted by Ridley’s use of the name for a species

from the Malay Peninsula.

Garcinia cymosa (K. Schum.) I.M. Turner & P.F. Stevens, comb. nov.

Basionym: Tripetalum cymosum K. Schum. in K. Schum. & Hollrung, FI.

Kais. Wilh. Land (1889) 51.

Type: New Guinea, island near Hatzfeldthafen, M. Hollrung 384, November

1886 (holotype, B - destroyed).

Leuconotis tenuifolia Engl., Bot. Jahrb. 7 (1886) 470, non Garcinia tenuifolia

Ridl. (1928). Type: New Britain, Blanche Bay at 500 m, F.C. Naumann s.n.,

16 August 1875 (holotype, B - destroyed; isotype, WRSL - fragment).

The Columnar Form of the Tree

Some trees of this species have pendulous branches resulting in a columnar

crown form rather than the typical conical crown. These were described by

Lauterbach (1923) as Tripetalum cymosum forma pendulum. Peekel (1984)

reported that the columnar trees grown in villages in the Bismarck

Archipelago did not grow uniformly to type from seed - some seedlings

reverting to wild type or intermediate forms. Intergradation in the crown

architecture among individuals argues against recognising the columnar

tree at a formal taxonomic rank. However, this tree is now being used in

tropical landscape planting, e.g. Singapore, and horticulturists require a

name for this entity. A cultivar name may be appropriate, viz. Carallia

brachiata ‘Honiara’ (Wong & Whitmore 1994), but it is more expedient to

Garcinia cymosa rT

transfer the forma to Garcinia and use that for the cultivated plant, at least

in the interim. Interestingly, the columnar form in cultivation in Singapore

does remain true to type when grown from seed (Mrs Kartini Omar-Hor,

pers. comm..).

Garcinia cymosa (K. Schum.) I.M. Turner & P.F. Stevens forma pendula

(Lauterb.) I.M. Turner & P.F. Stevens, comb. nov.

Basionym: Tripetalum cymosum K. Schum. forma pendulum Lauterb., Bot.

Jahrb. Syst. 58 (1923) 44, sphalm. pendula.

Syntypes: Bismarck Archipelago, New Britain, Vunapope, G. Peekel 809d,

August 1911; Vakabur, G. Peekel 809b, August 1911 (lectotype, selected

here, WRSL).

References

Brummitt, R.K. 1992. Vascular Plant Families and Genera. Royal Botanic

Gardens, Kew.

Hoft, R. 1992. Plants of New Guinea and the Solomon Islands: dictionary

of the genera and families of flowering plants and ferns. Wau Ecology

Institute Handbook. 13: 1-166.

Kostermans, A.J.G.H. 1976. Notes on Clusiaceae of Sri Lanka and reduction

of Pentaphalangium Warb. Ceylon Journal of Sciences (Biological

Sciences). 12: 55-71.

Mabberley, D.J. 1997. The Plant-Book. 2": Edition. Cambridge University

Press, Cambridge.

Peekel, P.G. 1984. Flora of the Bismarck Archipelago for Naturalists. Transl.

E.E. Henty. Office of Forests, Lae.

Schumann, K. and M. Hollrung. 1889. Die Flora von Kaiser Wilhelms

Land.

Steenis, C.G.G.J. van. 1987. Checklist of Generic Names in Malesian Botany

(Spermatophytes). Flora Malesiana Foundation, Leiden.

Stevens, P.F. 1995. Guttiferae subfam. Calophylloideae. Handbooks of the

Flora of Papua New Guinea. 3: 61-126.

Wong Yew Kwan and T.C. Whitmore. 1994. Carallia brachiata cv. Honiara,

a beautiful fastigiate ornamental tree. Gardens’ Bulletin, Singapore. 46(2):

93-98.

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Gardens’ Bulletin Singapore 51 (1999) 179-182.

Thismia goodii (Burmanniaceae), the Blue-capped

Thismia, a New Species from Borneo

RUTH KIEW

Singapore Botanic Gardens,

1 Cluny Road, Singapore 259569

Abstract

A new species in Sect. Sarcosiphon of Thismia, T. goodii Kiew, with remarkable blue

perianth lobes, is described from Sabah, Borneo.

Introduction

Thismias are poorly known because, being ephemeral, they are rarely

encountered and, being succulent, they make poor herbarium specimens.

This new species was encountered on a sandstone outcrop where

two small populations were found growing among the leaf litter (Kiew,

1999). Its most striking feature is the bright blue cap comprised of the

perianth lobes that top the flower (Fig. 1). In its mitriform perianth lobes

without tentacles or appendages and in its coralline root system, it clearly

belongs to Sect. Sarcosiphon, which until now included three other Malesian

species: T. clandestina (Blume) Mig., T. crocea (Becc.) J.J. Smith and T.

episcopalis (Becc.) F. v. Muell. (Jonker, 1948).

It differs from all three species in its coloration and the presence of

the distinct three-lobed outer perianth (Fig. 2). In addition, it differs from

T. crocea in its linear (as opposed to spathulate) inner perianth lobes,

which border the large holes in the mitriform apex, and from the other two

species in the absence of flanges or hairs on the anthers. The other Bornean

species, 7. episcopalis, would not be confused with this new species being a

much larger plant (5-19 cm tall) with more flowers per stem (1-8 flowers),

which are yellow. While the Javanese T. clandestina is a smaller plant (up

to 4-5 cm tall) with 1-2 flowers per stem, it is different in its greyish-green,

smaller flowers 10 (rarely 13) mm long.

Description of Thismia goodii Kiew, sp. nov.

Inter species sectionis Sarcosiphonis ponenda sed abeis colore perianthii

loborum lazulino et lobis exterioribus perianthii conspicuis recedens.

180 Gard. Bull. Singapore 51 (1999)

Figure 1. a. Thismia goodii growing on leaf litter, b. the flower.

Photos: Ali Ibrahim.

Typus: Ruth Kiew RK 4611, Borneo, Sabah, Sipitang, Ulu Maligan, 19

March 1999 (holotypus SAN, isotypi L, SING).

Small succulent achlorophyllous herb with pale brown coralline root system.

Stems glabrous, pale fawn, 0.75—7 cm long and in life 2-3 mm thick, erect,

unbranched. Leaves spirally arranged, distant, 1.5—2 x c. 1 mm long, narrowly

lanceolate, appressed to and concolorous with stem. Bracts 3, at base of

flower forming an involucre covering the ovary, narrowly lanceolate, 7-9 x

3-4 mm, apex acuminate, pale brown, sparsely hairy especially on margin.

Flowers usually solitary, rarely 2 and then flowering in succession, (15-—)

18-24 mm long, erect, with slight unpleasant smell of fish oil. Perianth tube

urceolate, 12-15 mm long, c. 3-4 mm wide at base dilating to (6—)8(—9)

mm at top with slight constriction below anthers, white with faint dark

green tinge with narrow dark blackish green longitudinal stripes alternating

with faint ones, internally with low longitudinal ribs with faint cross bars,

caducous. Perianth lobes electric blue in colour. Outer perianth lobes 3, +

erect, slightly recurved, 2-3 x 5-9 mm, with minute point at centre. Inner

Thismia goodii 181

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Figure 2. Thismia goodii Kiew.

A. whole plant; B. flower viewed from the top showing outer and inner perianths; C. 3 stamens;

D. fruits with persistent stigma and style.

(Scale bar: 5 mm)

182 Gard. Bull. Singapore 51 (1999)

perianth lobes 3, narrow at base arching upwards to join forming a mitre

(S—)7(—9) mm high and making 3 large holes, thicker at apex and acuminate,

apex without appendages. Stamens 6, hanging from the inside of thickened

rim of perianth tube, filaments ribbon-shaped, free, 2—2.5 mm long, anthers

joined laterally, c. 3 mm long each with a central adaxial ridge, apex 3-

toothed, without flanges or hairs. Ovary obovoid, c. 3.5 x 3 mm, deep

purple, 1-celled with 3 placentas; style cylindric, c. 1.5 x 0.5 mm; stigmas 3-

fid, c. 1.5 mm long, minutely papillose. Fruit fleshy, urceolate, 3.5-5 x 3-4

mm, dark brown, persistent base of perianth tube forming a rim c. 0.5 mm

high, stigma and style persistent. Seeds numerous, oblong in outline, c.

0.15 mm long.

Distribution: Known only from the type locality.

Habitat: On thick leaf litter on a sandstone outcrop at 1260 m altitude.

Notes: This species is named for Mr Good Antok, who first discovered it

during an expedition to the Sipitang District in eastern Sabah.

Acknowledgements

The expedition to the Sitipang District, Sabah, was only possible with the

logistic support of the SAN Herbarium and the Sabah Agricultural Park.

The experience, competency and companionship of the field teams were

invaluable to the success of the expedition and the author is extremely

grateful to Diwol Sundaling, Good Antok and Kambira Loloh from SAN,

Herbert Lim, Jain Linton and Aninguh Surat from the Sabah Agricultural

Park, Ali Ibrahim and Saifuddin Suran from SING and Tan Jiew Hoe

from Singapore. The curators of E, L and SAN are thanked for permission

to examine specimens in their care, Mark J.E. Coode for correcting the

botanical latin and Ali Ibrahim for permission to reproduce his photographs.

References

Jonker, F.P. 1948. Burmanniaceae. Flora Malesiana. 4: 13-26.

Kiew, R. 1999. Blue-capped Thismia. Gardenwise. 13: (in press).

Gardens’ Bulletin Singapore SI (1999) 183-256.

The Genus Rhaphidophora Hassk.

(Araceae-Monsteroideae-Monstereae) in

Peninsular Malaysia and Singapore

P.C. Boyce

Royal Botanic Gardens, Kew,

Richmond, Surrey, TW9 3AE, U.K.

Abstract

An alpha-taxonomic account of Rhaphidophora in Peninsular Malaysia and Singapore is

presented as a precursor to the forthcoming Flora Malesiana Araceae treatment. Fifteen

species, two newly described, R. corneri, and R. nicolsonii, are recognized. An extensive

new synonymy is proposed: R. pteropoda (syn. Scindapsus pteropodus) is synonymized with

R. angustata,; R. celatocaulis (syn. Pothos celatocaulis), R. copelandii, R. korthalsii var.

angustiloba, R. latifolia, R. maxima, R. palawanensis Furtado non Merr., R. ridleyi (syn. R.

grandis Ridl. non Schott), R. tenuis, and R. trinervia with R. korthalsii; R. fluminea with R.

beccarii; R. apiculata Alderw. non K. Krause with R. maingayi; R. celebica with R. minor;

R. burkilliana with R. montana; R. batoensis, R. gracilipes, R. hallieri, R. kunstleri, R.

megasperma, R. pilosula, and R. scortechinii with R. puberula; R. gratissima (syn. R. sylvestris

var. obtusata), R. lingulata (syn. Monstera lingulata, Scindapsus lingulatus), R. motleyana,

R. wrayi, and Scindapsus aruensis with R. sylvestris. Dichotomous keys to genera and

species are provided. All species except R. falcata are illustrated.

! Contents

MT GUCNI AKG CMe Mn ceed he cat tea se caae ead acess aie dtnssht Cock de 55 Aaayeninn se aceedees 184

1 LIS AC) OR SSR ooo nes a? OR IRM ne En ee er ee a ee ee 184

AP EtOH VIC 80) AGING a. opt Aa scons ya notre sxevenecundaeei via iudapdspelncdancstnstsonbeche 185

PRAT NT Ve TINA NINO 6A yiiect.ccteta Aiudacdpund deans sos adeiiedeaUadsaesccdnesncendiesss 186

isis TIMING AUN) Tine Or etre n SNe meee SUNY. cadets ananncaubaet-siapardsichessoaseesesnves 188

Key to genera of Anadendreae and Monstereae in

Perret lly vib Stan AMI SANE AOC, 52 o 25st !ocnccsaeesesccoreetieeacencaceoceoetedeoats 190

EP SUC Ree) a RES Si Bas ge gC er ey Oa a ee 191

2 EO ONCE, ee ge, ee. a ee oa eae ee 197

Key to adult flowering Rhaphidophora plants in

Pre eta lean, WE aa yea ANG SING O Ee ce, 6.1 40ika snp - dence nendptuinncnperesndertepeneaaee 196

he SPeccies a cs.54. Sa ee ee ee ee ee ee ee 19%

DNDN OS as cg ek 2 eck oh eu UBER es Ganka chons ends uss ganas edenbnenap ae isnnceseanters 249

ae Ba Sat Petia Sie A Sci ta Unc ndvaniniopinun sshd bretadenrnnntesce~sennsitices 249

PRO fee NC hs cre wade norte esac pad sedin debra tions sn fdnicnnn cngen eds ex noponoenés 254

dems eet AE ae eRe eA ee eke cies oe aciivedan fosisys cap doeetsesedens ca skse ono nc¥sbesowtewennae 256

184 Gard. Bull. Singapore 51 (1999)

Introduction

Rhaphidophora Hassk. (including Afrorhaphidophora Engl.; c. 3 species in

tropical Africa) comprises. c. 100 species of small to large, occasionally

enormous, root-climbing lianes (sensu Schimper, 1903), rarely rheophytes,

distributed from tropical West Africa eastwards to the western Pacific,

north to southern Japan (Ryukyu Islands), and south to Northern Australia.

Rhaphidophora is one of the largest aroid genera in tropical Asia, and has

several nodes of diversity: the Himalaya (SE Nepal to NE Vietnam, roughly

17° — 23° N), West Malesia (including southernmost peninsular Thailand),

the Philippines, and East Malesia.

This is the first in a series of papers intended to present a complete

alpha-taxonomy of the genus Rhaphidophora in Asia. Accounts for each

of the Himalaya, Thailand, Indochina, the Philippines, Borneo, the

Indonesian archipelago, and Papuasia are being prepared and will be

published separately. All morphological terms employed follow Stearn

(1992).

History

Since establishment by Hasskarl (1842), Rhaphidophora has been dogged

by interpretative confusion arising from the generic descriptions published

by Hasskarl between 1842 and 1848. This confusion led to several papers

aiming to settle the nomenclatural identity of Rhaphidophora, e.g., Brown

(1882), Bakhuizen (1958), and Nicolson (1978). Although Nicolson (1978)

laid to rest the problem of interpreting Hasskarl’s genus, it is worth repeating

the basic arguments and details here.

Hasskarl’s original generic description (Hasskarl, 1842a) was sparse.

Another, more detailed description followed in Tijdschrift Nat. Gesch. &

Phys. (Hasskarl, 1842b), and a further very sparse description in Catalogus

plantarum...Horto Botanico Bogoriensi... (Hasskarl, 1844). These

descriptions agree on the main diagnostic feature of Hasskarl’s new genus:

a unilocular, uniovular ovary. However this matches exactly the diagnoses

of the earlier Scindapsus Schott (Schott & Endlicher, 1832), and also,

incidentally, the later Amydrium Schott (including Epipremnopsis Engl. —

see Nguyen & Boyce, 1999) However, by the time Hasskarl published an

extensive description of a Rhaphidophora species, R. lacera (Hasskarl,

1848), he had altered his generic description stating that his earlier

observations were in error, and that Rhaphidophora was characterized by

possessing a two locular ovary in which each locule has two ovules; exactly

the situation in the later Epipremnum Schott (Schott, 1857; Boyce, 1998).

Rhaphidophora in Peninsular Malaysia and Singapore 185

Indeed, specimens in BO annotated by Hasskarl as R. lacera are of the

common and widespread Epipremnum pinnatum (L.) Engl., (see Boyce,

1998). The above might lead one to conclude that based on the diagnoses

and annotated specimens the name Rhaphidophora (sensu Hasskarl, 1848)

is the correct name for Epipremnum (Schott, 1857). However, this is not

so. In publishing Rhaphidophora Hasskarl (1842a) cited Pothos pertusa

Roxb. [syn. Rhaphidophora pertusa (Roxb.) Schott] as a synonym of the

generic type of his new genus. This has two results. The first is that Hasskarl

should have taken up the epithet pertusa for his new species. Rhaphidophora

lacera Hassk. is thus is illegitimate. The second is that the element cited by

Hasskarl pertains to a species from South India and Sri Lanka, which has

two intrusive placentae each bearing numerous ovules, and thus does not

agree with any of Hasskarl’s diagnoses of Rhaphidophora. Nicolson (1978)

concluded that the specimen cited by Hasskarl should be regarded as typical

for the genus Rhaphidophora over the descriptive diagnoses. Therefore

Rhaphidophora as currently defined has two intrusive placentae each

bearing several to many ovules with the type R. pertusa of which R. lacera

is a synonym.

Since establishment of Rhaphidophora, almost all authors have

accepted it as a ‘good’ genus. The one exception is Miquel (1856) who

merged Rhaphidophora with Scindapsus at the rank of subgenus. No one

took up this new rank in subsequent treatments.

The last full revision of Rhaphidophora was that of Engler & Krause

(1908). Since then numerous regional accounts have been produced,

including China (Li, 1979), India (Sivadasan, 1982; Deb, 1983), Nepal (Hara,

1978), Bhutan (Noltie, 1994), Fiji (Nicolson, 1979), Sri Lanka (Nicolson,

1988) and Australia (Hay, 1993). To date no critical account of the genus

has been prepared for Peninsular Malaysia (but see Hooker, 1893; Ridley,

1907, 1925; Henderson, 1954), Borneo (but see Alderwerelt, 1920, 1922;

Miquel, 1856a & b; Ridley, 1905; Merrill, 1921), Java (but see Koorders,

1911; Backer & Backhuizen, 1968), the Philippines (but see Merrill, 1923),

New Guinea (but see Engler & Krause, 1910, 1912; Krause & Alderwerelt,

1924; Hay, 1981, 1990), the Himalaya (but see Hooker, 1893; Hara et al.,

1978; Boyce, in press) nor Thailand and Indochina (but see Gagnepain,

1942).

Geography and Endemism

Malesian Rhaphidophora species divide into two distributional groups. One

comprises taxa with limited distributions, sometimes narrowly endemic,

more usually restricted to one or more geographically adjacent landmasses,

186 Gard. Bull. Singapore 51 (1999)

and displaying limited morphological variation. The other group comprises

species with extensive distributions and, usually, a wide range of variation.

The species with restricted distributions occurring in Peninsular

Malaysia are Rhaphidophora corneri P.C. Boyce and R. nicolsonii P.C.

Boyce (both known from one location each in the Malay Peninsula); R.

angustata Schott and R. crassifolia Hook.f. (scattered but never abundant

in Malay Peninsula and possibly Sumatera); R. falcata Ridl. and R.

tetrasperma Hook.f. (Peninsular Malaysia and the far south of Thailand, R.

falcata known from two collections, R. tetrasperma from six) and R. maingayi

Hook.f. (Peninsular Malaysia, southern peninsular Thailand and Sumatera.)

Widespread taxa represented in the Malay Peninsula include R.

korthalsii Hassk. (common throughout Malesia and into the western Pacific,

extending north into southern peninsular Thailand), R. minor Hook.f.

(widespread but not common in western and central Malesia but absent

from Maluku and in the Philippines known only from Palawan), R. puberula

Engl. (localized in Peninsular Malaysia, Sumatera and Borneo), R.

foraminifera (Engl.) Engl. (uncommon in Malay Peninsula, Sumatera and

northern Borneo), R. beccarii Engl. (widespread and generally abundant

in west Malesia, north to southern peninsular Thailand and east to Borneo

but absent from Java and Nusa Tenggara), R. montana (Blume) Schott

(common throughout Malesia and extending into southern Thailand, though

absent from the Philippines), R. /obbii Schott (widespread but never

common in Peninsular Malaysia, Singapore, Borneo, Sumatera and

Palawan) and R. sylvestris (Blume) Engl. (relatively common throughout

Malesia and extending north to southern Thailand, but absent from the

Philippines, Sulawesi and Maluku). Within this group of widespread species,

R. minor and R. lobbii are remarkably unvarying, R. foraminifera, R. beccarii

and R. puberula are morphologically quite stable (although R. beccarii

displays varying degrees of neoteny and sporadic leaf lamina perforation

occurs in R. puberula), but R. montana, R. sylvestris and especially R.

korthalsii display a bewildering range of variation the taxonomic

implications of which are still very imperfectly understood.

Thus Peninsular Malaysia has two endemic species, two species

otherwise only in Sumatera, two species known also from southern Thailand

and eight species with distributions extending variously to Borneo, the

Philippines and the western Pacific.

Structure and Terminology

The position of the inflorescence and the architecture of the shoot it

Rhaphidophora in Peninsular Malaysia and Singapore 187

terminates are useful characters for assigning species names. Although

Monstereae (and Anadendreae) appear architecturally less diverse than

Potheae, shoot morphology nevertheless presents challenges for description

due to the plants’ often great size and complex shoot organisation, coupled

with an inherent architectural variability in some taxa. Broadly speaking

the shoot architecture of adult Rhaphidophora plants can be categorized

into three types:

1. Physiognomically monopodial clinging non-flowering stems rooting

along their entire length giving rise to variously elaborated free

sympodial lateral stems flowering terminally (e.g., R. lobbii, R.

puberula, R. angustata)

ii. | Physiognomically unbranched sympodial clinging stems flowering

terminally and free lateral stems usually not occurring other than as

a result of physical damage (e.g., pulled down by weight of foliage)

and then free stems usually soon climbing again (e.g., R. korthalsii)

ii. Physiognomically monopodial clinging stems and clinging sympodial

lateral stems but only sympodial lateral stems flowering (e.g., R.

foraminifera).

Species belonging to the second and third of these types are rare in

Peninsular Malaysia, with only R. korthalsii and R. nicolsonii present for 1.

and R. foraminifera for 1. The remaining fourteen Peninsular Malaysian

species all conform to the first type. When species of the first type are

examined it appears that they are not heterogeneous as a group, suggesting

that type 1. architecture has evolved repeatedly, but that phylogenetic

analysis is required. For the purposes of identifying species, it is a useful

character.

Leaf shape varies from uniformly oblong elliptic throughout the genus

(e.g., R. montana) to pinnately divided (e.g., R. tetrasperma), perforated

(e.g., R. corneri) or a combination of all three (e.g., forms of R. korthalsii).

If used with care, the shape of the leaves can be very useful in identifying

plants to species level. The leaf selected should be a mature leaf taken

either from an upper part of a clinging stem, but not a leaf subtending a

flowering event, or from the mid-point of a free lateral stem. Leaf

measurements given in the descriptions below are for leaves from these

positions but smaller leaves will often be found on herbarium specimens,

usually as a result of collecting a specimen of a convenient size for

preservation, and will often be considerably smaller than the dimensions

given here.

Some terms employed here and elsewhere may be unfamiliar. Below

is a list of definitions (taken from Boyce, 1998):

188 Gard. Bull. Singapore 51 (1999)

Physiognomically unbranched stems

Fertile stems, of variable length, with the appearance of being

unbranched but that are actually sympodia with growth terminating

by a, sometimes aborted, inflorescence. Such stems may be clinging

and orthotropous (or nearly so) or free and plagiotropic to pendent.

Clasping roots

Short specialized roots that anchor a climber, hemiepiphyte or

epiphyte to its substrate, generally a tree or rock.

Feeding roots

Specialized roots arising from aerial stems which, extending down

to the soil, transport nutrients to the plant.

Shingle climber

A type of juvenile morphology found in climbers, in which the

petiole is very short and the leaf blade relatively broad and more-

or-less overlapping with its neighbours to resemble the tiles (or

shingles) of a roof; such plants are found climbing up larger tree

trunks; e.g., R. korthalsii.

Interprimary veins

A vein approximately parallel to and situated between the primary

lateral veins and larger than secondary veins.

Longitudinal orientation (of stigmas)

Linear stigmas set parallel to the long axis of the spadix.

Trichosclereids

Literally a hair-like sclereid; fibre cells (cells with thick, lignified

walls) which are very slender and elongated so as to be visible to the

naked eye as hair-like structures. On tearing the leaf blade they can

be observed protruding from the torn edge.

Identification

Lianescent aroids are often collected under incorrect names in Asia (see,

e.g., Boyce, 1998; Nguyen & Boyce, 1999 for further discussion). Part of

the problem with accurate identification arises from an apparent lack of

readily observable critical characters, a difficulty exacerbated by the

Rhaphidophora in Peninsular Malaysia and Singapore 189

tendency of key-writers to concentrate on fertile characters, which are

difficult to observe, and interpretively ambiguous vegetative characters. I

have attempted where possible to concentrate on readily observable

characters in the keys presented here. In particular, I have employed the

feature typical of most monsteroid genera, trichosclereids. All monsteroid

genera (except Amydrium) have abundant trichosclereids (sparse and

scattered in Amydrium). These are readily observable by tearing a mature

leaf lamina and looking for ‘hairs’ protruding from the damaged edges and

are an extremely reliable character in Asia as a means of assigning a genus

to the tribe Monstereae. For a discussion on identifying lianes of the tribe

Potheae see Boyce (1998).

The keys presented below should go some way towards easing the

identification of the smaller-inflorescenced lianescent aroids to genus in

Asia. However, while it has proved relatively easy to key these out by this

method, species with large inflorescences have proved very intractable and

here I have had to resort to critical floral characters in order to produce a

reliable key for those taxa. As we gain a better insight into unvarying (or

at least unambiguous) vegetative and macrofloral characters I hope to be

able to produce more user-friendly keys for these larger flowered taxa.

Keys to species are less problematic, but earlier keys (e.g., Hooker,

1893; Ridley, 1905; Engler & Krause, 1908) tended to concentrate on leaf

shape as a means of separating Rhaphidophora species. This leads to

difficulties. Earlier monographers were generally dealing with fewer taxa

and far fewer collections than today and additionally it is frequently unclear

which leaf, from what position on the plant, is being employed in the key.

This latter problem is exacerbated by leaves collected for herbarium

specimens frequently being chosen for their suitability for drying and

mounting (i.e. small enough) rather than for their taxonomic usefulness.

This difficulty has led to the comment, not altogether unjustified, that

these early identification tools are keys to individual leaves rather than to

species. Having said that, there is little doubt that the ability to identify

aroid lianes by vegetative characters alone is very desirable given that

climbers are frequently encountered sterile in the field or, where fertile,

the inflorescences are seldom within easy reach and often, once brought

down from the canopy are found to be either too immature or too old to

be of much use. Thus, wherever possible, leaf characters, combined with

easily observed shoot architecture are employed together with inflorescence

characters in the keys provided here.

The key that follows is designed to help separate the monsteroid and

anadendroid lianes, genera that are frequently confused in herbaria.

190

la.

1b.

2a:

2b.

3a.

3b.

4a.

Gard. Bull. Singapore 51 (1999)

Key to the Genera of Anadendreae and Monstereae in

Peninsular Malaysia and Singapore

Inflorescence small; spathe mostly less than 9 cm long just prior to

Opening (A: Beek tadias in itwnt. eden gee ee ee 2

Inflorescence moderate to large; spathe mostly more than 9 cm long

just pronto Opening +.de.cdpuacs ia. caud lc ee, A eee 4

Spathe in bud slender with long and slender peduncle, conspicuously

long-beaked (beak to a third length of entire spathe), opening with

inside conspicuously glossy-waxy, white, greenish white or purple.

Flowers with a membranous perigon of fused tepals (i.e. flowers

perigoniate). Ripe fruits berry-like, dark red. Trichosclereids absent

a Aedes ee ae me ee boy: inane sel ee ele Anadendrum

Spathe in bud stout with short to long peduncle, not conspicuously

long-beaked, or if beak present then less than a sixth length of entire

spathe, opening with inside only moderately waxy yellow, orange,

greenish or white. Flowers naked (i.e. flowers aperigoniate). Ripe

fruits not berry-like, stylar region mostly abscissing to reveal ovary

cavity, if berry-like and stylar region not abscissing then fruits ripening

white or orange. Trichosclereids present (but sparse in Amydrium)

Trichosclereids abundant (many ‘hairs’ apparent when a mature leaf

lamina is torn). Leaf with petiole broadly canaliculate, sheath margins

broad, spreading, persistent, extending to apical geniculum; leaf lamina

thinly coriaceous, often variegated silvery grey. Ripe fruits with stylar

region abscissine:.13.2 3a tag s.cteeeee ee e, Scindapsus

Trichosclereids sparse (very few ‘hairs’ apparent when a mature leaf

lamina is torn). Leaf with petiole narrowly canaliculate or terete, sheath

margins narrow, erect or slightly inrolled, soon drying and degrading

into weak fibres, then falling to leave an obscure scar, sheath either

with free ligules extending beyond apical geniculum or sheath at most

extending only to half way along petiole, if the latter then remainder

of petiole terete with two prominent adaxial keels extending to apical

geniculum; leaf lamina variously textured. Ripe fruits with stylar region

NOt ADSCISSING 4.4.7 cc dere ethane ome ee Amydrium

Trichosclereids sparse (very few ‘hairs’ apparent when a mature leaf

lamina is torn). Higher order venation completely reticulate. Ripe

fruits with stylar region not abscissing Ovary 1-locular, placenta 1,

intrusive-parietal, OVUIES 2 ooiisusditiiicser ss -icecun eer Amydrium

Rhaphidophora in Peninsular Malaysia and Singapore 19]

4b. Trichosclereids abundant (many ‘hairs’ apparent when a mature leaf

lamina is torn). Higher order venation striate or reticulate. Ripe fruits

with stylar region abscissing. Ovary never as above combination..... 5

5a. Fruits with a solitary seed. Ovules solitary, placentation basal.

hE SN ES te BE, Mali OSA PEER URI OL Ee foc te eae ee Scindapsus

5b. Fruits with more than one seed. Ovules 4-6 or more, placentation

LGU S227 u uve, 1 las San CA ota et Wat St oe eee co ee eee 6

6a. Seeds many, ellipsoid, straight, 1.3-3.2 mm long, 0.6-1.0 mm wide;

testa brittle, smooth. Ovules 8 or more, superposed on 2 (rarely 3)

imlerUSIve spPanictal PIACCHEAS. ..2....0.eNA Nee eed Rhaphidophora

6b. Seeds few, curved, 3-7 mm long, 1.5-4.0 mm wide; testa bony and

ornamented. Ovules 4(-6) at base of a single intrusive parietal placenta.

Ne eM AN Sas Hc A Rican ccd CCA A REPTILE wok AC Maes Epipremnum

Field techniques

Specific taxonomy of Rhaphidophora is problematic in part because the

species are too often inadequately collected, and the specimens are often

sterile and almost invariably lacking the differing morphological phases

that the climbing aroids display at different stages of maturity. Moreover,

most species are rarely collected either because of natural scarcity or,

more often, because they are passed by as ‘too difficult’ to collect. While it

is true that aroids generally, and aroid lianes in particular, do not lend

themselves to convenient preparation for herbarium specimens, they are

no worse to deal with than many of the larger compound-leaved trees and

are considerably less tedious to handle than, for example, rattans, groups,

which are amply represented in herbaria worldwide.

As with all flowering plants the provision of fertile material is an

essential prerequisite to making a taxonomically useful herbarium voucher.

Before any material is collected from a particular plant, field notes should

be prepared, describing the life form of the plant (climber or rheophyte),

the position of the inflorescences (on adherent or free shoots), together

with geographical and ecological notes as appropriate. As the collecting

progresses, make notes of any characters that will be obscured or altered

by the drying process such as venation of leaves (impressed, raised), the

dimensions of fleshy or juicy organs, exudate, odours, and all colours should

be painstakingly noted. If photographic records can be made also, so much

the better, but nothing can replace the careful noting of field data.

When it comes to gathering plant material, collection of representative

192 Gard. Bull. Singapore 51 (1999)

pieces of the plant to display different phases of growth, e.g., shingling

shoots, foraging shoots, is very important. Leaves with extreme dimensions

along with those of average size and those displaying variations in

perforation and pinnation should also be gathered. Ideally inflorescences

in different stages should be taken, together with semi-mature and mature

fruits.

There is little point in gathering flowering material of aroids only to

dry the specimens in such a way that the spadix is invisible. If material

allows, two inflorescences, one at female receptivity and one at male anthesis

should be gathered in 70% alcohol in addition to collecting fertile specimens

for drying. If there are insufficient inflorescences for making spirit and

dried collections half the spathe should be removed longitudinally to reveal

the whole spadix but cut in such a manner as to leave the top the spathe

intact. Do not cut the spadix into longitudinal slices for drying unless

absolutely necessary. Material prepared in such a manner distorts badly

when dried and is usually useless for taking measurements.

Ideally you should end up with herbarium vouchers and spirit

collections that, combined with extensive field notes, permit one to visualize

the particular species in almost as much detail as if the plant were seen

growing.

RHAPHIDOPHORA

Rhaphidophora Hassk., Flora 25 (2) Beibl. 1 (1842) 11; Schott, Gen. Aroid.

(1858) 77 & Prodr. Syst. Aroid. (1860) 377-388; Miquel, Ann. Mus. Bot.

Lugd.-Bat. 3 (1867) 81-82; Engl. in A. & C. DC., Monogr. Phan. 2 (1879)

238-248; Engl. in Beccari, Malesia, vol. 1 (1882) 266-272, Tab. xix 6-9, xx

1-5; Benth. & Hook. f,, Gen. Pl. 3(2) (1883) 993-993; Engl. & Prantl, Nat.

Pflanzenfam. T. 2, Ab. 3 (1889) 119-120; Hook.f. in Hook.f, Fl. Brit. India,

vol. 6 (1893) 543-548; Engl. & Prantl, Nat. Pflanzenfam. Nachtr. 1 (1897)

58; Ridl., J. Straits Branch Roy. Asiat. Soc. 44: 185-187 (1905) & Mat. FI.

Malay Penins. 3 (1907) 39-46; Engl. & K. Krause in Engl., Pflanzenr. 37

(IV.23B) (1908) 17-53; Engl. & Prantl, Nat. Pflanzenfam. Nachtr. 3 (1908)

29; Koorders, Exkursfl. Java, vol. 1 (1911) 253-255; Merrill, J. Straits Branch

Roy. Asiat. Soc., special number (1921) 88-90 & Enum. Philipp. Flower.

Pl., vol. 1 (1923) 175-177; Ridl., Fl. Malay Penins., vol. 5 (1925) 120-124;

Henderson, Malayan Wild Flowers, Monocots, (1954) 238-239, Fig. 142;

Backer, Beknopte Fl. Java, vol. 17 (1957) 13-15; Backer & Bakh.f., FI.

Java, vol. 3 (1968) 106-107; Nicolson in A.C. Sm., Fl. Vitiensis Nova, vol. 1

(1979) 443-445, Fig. 88; Hay in R.J. Johns & Hay, Students’ Guide Monocot.

Papua New Guinea. Part 1 (1981) 68-72 , Fig. 29; Schott, Icones aroideae

Rhaphidophora in Peninsular Malaysia and Singapore 193

et reliquiae (IDC Microfilm) (1983) fiche nos. 28-31, 121; Hay, Aroids of

Papua New Guinea (1990) 83-87, Figs. 34, 35, Pl. XIVb, XV & Telopea 5

(1993) 293-300; Hay et al. Checklist & botanical bibliography of the aroids

of Malesia, Australia and the tropical western Pacific. Blumea, suppl. 8

(1995) 111-127; Mayo et al., Genera Araceae (1997) 118-121, Pl. 14, 109 D

— Scindapsus Schott subgen. Rhaphidophora (Hassk.) Migq., Flora Ned.

Indié 3 (1856) 185 — Type: Rhaphidophora lacera Hasskarl, nom. illeg.

pro. Pothos pertusus Roxb. [= Rhaphidophora pertusa (Roxb.) Schott]

Scindapsus Schott subgen. Pothopsis Miq., Flora Ned. Indié 3 (1856) 187

— Type: Scindapsus sylvestris (Blume) Kunth [= Rhaphidophora sylvestris

(Blume) Engl.|

[Raphidophora Hassk., Cat. Hort. Bogor. (1844) 58, orth .var.|

Medium-sized to very large, occasionally enormous, slender to robust,

leptocaul or pachycaul, homeophyllous or heterophyllous, rarely neotenic

(e.g., some populations of R. beccarii), root-climbing lanes, very seldom

clustering and rheophytic (e.g., R. beccarii); cut surfaces producing clear,

odourless sticky juice either drying + invisibly or coagulating into yellowish,

translucent jelly and eventually hardening to an brittle amber-like mass;

seedling stage mostly not observed but where known either leafy at

germination and probably skototropic (see Strong & Ray, 1975; c.f. Pothos,

Boyce, in press) by an alternating series of congested leafy and elongated

leafless shoots (e.g., R. angustata Schott) or germinating to give rise to a

non-skototropic shingling juvenile shoot (e.g., R. korthalsii Hassk); pre-

adult plants often forming modest to extensive terrestrial colonies of varying

morphological and physiological form such that descriptive generalizations

are nearly impossible, largest terrestrial colonies generally occurring in

places of less than optimum adult growth potential (e.g., depauperate tree

canopy, dry, exposed sites,); adult shoot architecture broadly divisible into

three types (but see R. corneri below): i. physiognomically monopodial

clinging non-flowering stems rooting along their entire length giving rise to

variously elaborated free sympodial lateral flowering stems (e.g., R. lobbii,

R. puberula, R. angustata), or 11. all stems physiognomically unbranched

sympodial clinging and flowering (e.g., R. korthalsii), or ii. physiognomically

monopodial and sympodial lateral stems clinging but only sympodial lateral

stems flowering (e.g., R. foraminifera). Stems with internodes of various

lengths separated by variously prominent leaf scars, smooth or asperous or

densely pubescent to ramentose (the last not in Malaysia), older stems

sub-woody or somewhat corky or with distinctive matt to sub-lustrous pale

brown papery epidermis, with or without variously textured prophyll,

194 Gard. Bull. Singapore 51 (1999)

cataphyll and petiolar sheath fibre either at the apices or along the newer

sections, rarely with cataphylls and prophylls deliquescing to black mucilage

later drying to leave fragmentary parchment-like remains on petioles,

developing laminas, inflorescences; flagellate foraging stems occurring in

some species, often exceedingly long, reaching the ground then rooting,

variously foraging and climbing again; clasping roots sparsely to densely

arising from the nodes and internodes, strongly adherent to substrate:

feeding roots rare to abundant, smooth, pubescent or prominently scaly,

later often becoming woody, strongly adherent to substrate or free; /eaves

distichous or spiralled, evenly distributed or scattered or clustered distally;

cataphylls and prophylls sub-coriaceous to membranaceous, either soon

drying and falling or degrading or deliquescing to variously textured sheaths

and fibres, these where present variously clothing upper stem before

eventually decaying and falling; petiole canaliculate to weakly carinate,

smooth or pubescent, with variously prominent apical and basal genicula;

petiolar sheath prominent, extending either partly to or fully to or

overtopping the geniculum, occasionally one side greatly expanded and

auriculate, especially in juvenile plants, at first membranaceous to

coriaceous, soon completely or along the margins drying chartaceous,

sometimes degrading to untidy variously netted or simple fibres and later

variously falling to leave a scar or disintegrating marginally or completely;

lamina sub-membranous to stiffly chartaceus or coriaceus, lanceolate or

oblong, + oblique, base decurrent to unequal or cordate, apex acute to

acuminate, entire to regularly pinnatifid or perforated, if pinnate then

divisions pinnatifid to pinnatisect (Stearn, 1992: 324), midrib often + naked

between segments, lamina occasionally with small to well developed

perforations adjacent to the midrib and primary veins, these sometimes

extending to lamina margin (fenestrations then occasionally additional to

fully developed pinnae), rarely abaxially pubescent when expanding, rarely

strongly concolorous at maturity (the last not in Malaysia); midrib usually

prominent, raised abaxially and prominently sunken, sometimes flush, rarely

slightly raised adaxially; primary venation + pinnate; interprimaries mostly

present, sub-parallel to primaries and sometimes indistinguishable from

them (e.g., R. monticola - Philippines) but usually less prominent and often

drying paler, usually glabrous, occasionally pubescent with domatia in the

axils of the primary and secondary veins; secondary venation striate (e.g.,

R. monticola — Philippines) to reticulate (e.g., R. korthalsii), variously

prominent, often very difficult to distinguish from primary venation (e.g.,

R. angustata); tertiary venation where visible reticulate to tessellate:

inflorescences solitary to several together, first inflorescence subtended by

a (usually fully developed) foliage leaf and/or a very rapidly disintegrating

cataphyll, subsequent inflorescences usually each subtended by a prophyll

Rhaphidophora in Peninsular Malaysia and Singapore 195

and cataphyll, more rarely by a prophyll and partially to almost fully formed

foliage leaf (but not in Peninsular Malaysia), inflorescences at anthesis

naked by disintegration of subtending cataphyll or partially to almost

completely obscured by netted and sheet-like fibres; peduncle terete to

laterally compressed; spathe ovate to narrowly or broadly canoe-shaped,

stoutly to rather weakly beaked, barely gaping to opening almost flat at

anthesis and then usually deciduous before anthesis is complete, occasionally

persisting into the early stages of infructescence development (e.g., R.

angustata), rarely drying and persistent (e.g., R. novoguiensis - New Guinea),

stiff to rather soft or stoutly coriaceous, dirty-white, greenish, cream or

yellow; spadix sub-globose to clavate-cylindrical, cylindrical or fusiform,

sessile or stipitate, often obliquely inserted on peduncle, tapering towards

the apex; flowers bisexual, naked; ovary 1- to partially 2-locular, lower part

+ bilaterally compressed, upper part variously cylindrical and variously

angled, most often rhombohexagonal, those upper- and lowermost on the

spadix often sterile and bereft of stigma, those uppermost frequently either

scattered or partially fused to each other and forming an appendix; ovules

few to many, anatropous, funicle long, placentae parietal to basal, sometimes

+ sub-axile, partial septa variably intrusive; stylar region well developed,

usually broader than ovary, usually truncate apically, rarely elongate-conic;

stigma sticky at female anthesis, punctiform, broadly elliptic or oblong and

then, orientation circumferential or longitudinal with respect to spadix;

stamens 4-6; filaments strap-shaped; anthers usually prominently exserted

from between pistils at male anthesis, rarely not exserted and pollen

extruded from between ovaries, dehiscing by a longitudinal slit;

infructescence with stylar.regions greatly enlarged, transversely dehiscent,

the abscission developing at the base of the enlarged to massive stylar

region and this falling to expose the ovary cavity with the many seeds

embedded in variously coloured sticky pulp; seeds oblong, testa thin, smooth,

embryo axile, straight, endosperm copious; pollen dicolpate, extended

monosulcate to fully zonate, ellipsoid or hamburger-shaped, medium-sized

(mean 33 um, range 24-55 um), exine foveolate, sub-reticulate, rugulate,

fossulate, scabrate, retiscabrate, verrucate, or psilate(Grayum, 1984);

chromosomes 2n = 60, 120 (42, 54, 56)(Petersen, 1989, 1993, 1994).

Distribution: Tropical and subtropical Africa, South and South East Asia,

Malesia, Australia and the tropical western Pacific.

Habitat: Lianescent bole-climbers, lithophytes, rarely rheophytes, usually

in well drained subtropical and tropical wet, humid, or seasonally

moderately dry primary and established secondary evergreen forest at low

to mid-montane elevations.

196 Gard. Bull. Singapore 51 (1999)

Etymology: Greek rhaphis, rhaphidos (needle) and pheré (1 bear); refers

to the macroscopic (to 1 cm long), needle-like unicellular trichosclereids

present in tissues.

Key to Adult Flowering Rhaphidophora plants in

Peninsular Malaysia and Singapore

la. Leaf lamina variously pinnately divided and/or perforated ............... 2

Ib. Leal Tatmane caiire »).:2..0..2.0.5.0..22 Se ee 8

2a. ‘Plants fowermp on adherent stems «i... aoa eee 3

2b. Planistiowertig on Tree stems rr 0/2A a. cd ceases tsccasene nese ee 5

3a. Flowers plants climbinp....2.0.4024 52n RR iene -

3b. Flowering plants rheophytic, not climbing.................. 2. R. beccarii

4a. Leaf lamina of mature plants slightly to extensively perforated,

perforations round to rhombic, extending c. '/s of lamina width on

each side of the midrib, abaxial surface of lamina with pubescent

veins; active shoot apices with black mucilage ........ 6. R. foraminifera

4b. Leaf lamina of mature plants pinnatisect, the pinnae often perforated

basally and appearing stilted, lamina always glabrous; active shoot

apices with sparse to copious netted fibre ......... 7. R. korthalsii

Sa. Abaxial surface of lamina and apical geniculum minutely pubescent

Wc EaR EER ints Nine vie > chisn does LOREM: te ee 12. R. puberula

5b. Lamina and apical geniculum not as above ............cceeeceeeeeeeeeeeeeeeeeeeeee 6

6a. Leaf lamina up to 53 x 47 cm, sparsely to + entirely deeply

pinnatipartite to nearly pinnatisect, occasionally with large rhombic

perforanons.ddjacent to midhid 2.10.4 occ tenth ee eee 7

6b. Leaf lamina up to 15 x 8 cm, entire or with a few, large ovate to

rhomboid or trapezoid perforations extending almost to the margin

dsl bine nines Seat ebeieN eel sia has Saeed so ls Bs HN 3. R. corneri

7a. Petiolar sheath extending '/2—*/4 along petiole; lamina coriaceous,

sparsely to rarely + entirely pinnatipartite, or pinnatisect......0.0.....0..

nsdeonsdenpis pobrenaarlilatardbpnguenettla ply eiicedaa iat asia eaten 11. R. nicolsonii

7b. Petiolar sheath extending to base of apical geniculum, lamina sparsely

to + entirely deeply pinnatipartite to nearly pinnatisect, occasionally

with large rhombic perforations adjacent to midrib .......... eee eee

nacvbons cuddnecddetussbion thuaddcee’levewerdsauD MW Oeee AURM ATIC ae Gea aa 14. R. tetrasperma

Rhaphidophora in Peninsular Malaysia and Singapore 197

8a.

Sb.

9a.

9b.

10a.

10b.

bbb.

12a.

12b.

13a.

Meloy

14a.

14b.

1D.

iSb,

Abaxial surface of lamina and apical geniculum pubescent...............

oh} VINER LU) tp We 3A ee eee 12. R. puberula

Flowering shoots consisting of scattered fans of large litter-trapping

leaves carried on short stout shoots and held at about 90° to the +

VIS Ss IN AIIN: SUCHIN eo teas tse che sec cceatiietouesecossonveescdeanl BOs aad! 1. R. angustata

Ey IM SIO OA AS AD OVE ooo. 52 2s) ec actguacadcevsteseaceseisssesescenoesects 10

Re Meare Him N Ne Be race ease eects DA, «NE ides vse ive ts avedenenseckeonesslly 11

Pict eriee lytic assert aol shah le P, ReaLalisee tons. 2. R. beccarii

Stems scabrid to asperous; spathe exterior minutely puberulent

eee ee a nee. vc tte ete cena Das ei ducdasee tae 8. R. lobbii

Stemsismooih: Spathe.extemor glalbPous s:cs40 ees. setiseccarivee nave 2

Apices of active stems with netted prophyll, cataphyll and petiolar

SNS Sls vn Poet Cy reage dasmenifews Miacshuin-snvnnmenpbantplede Meech cca einer 9. R. maingayi

Paces OF dative stems Make vin jesectee siiesocaraeeite tes intcscnsgtenesatensesney’: 13

Spadix at anthesis 3.5—6 cm long, cylindrical to weakly clavate .... 14

Spadix at anthesis 9-20 cm long, tapering apically ....10. R. montana

EVAN GRE 11 TES 010 2) AMM Bee A RPO, cee: ae oe ee ee 15

RS TEYELOUS, Se Pee ed Ac! © A021 cae Mien ce Se 13. R. sylvestris

Leaf lamina thickly coriaceous to almost fleshy, falcate-elliptic-

lanceolate to falcate-oblong or falcate-oblanceolate, 4.5—25.5 x 1.5-5

cm; margins slightly reflexed, this becoming greatly accentuated in

ELEC MY TRS ee) DCRR eh Sr) eae eee 4. R. crassifolia

Leaf lamina thinly coriaceous, narrowly falcate-elliptic to falcate-

lanceolate or falcate-oblanceolate, 2.5-16 x 1.2-3 cm, margins flat,

Fe AN Ale SULA W -CONOUII OC po. ou... coennzeiaanassaphceesaneandeneusse 9. R. minor

The Species

1. Rhaphidophora angustata Schott

Rhaphidophora angustata Schott, Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 128;

Engl. in A. & C. DC., Monogr. Phan. 2 (1879) 241; Engl. & K. Krause in

Engl., Pflanzenr. 37 (IV.23B) (1908) 25 — Type: Indonesia, ‘?Java’, P.W.

Korthals 206 (L, holo).

198 Gard. Bull. Singapore 51 (1999)

Scindapsus pteropodus Teijsm. & Binn., Natuurk. Tijdschr. Ned.-Indié. 27

(1864) 23; Engl. in A. & C. DC., Monogr. Phan. 2 (1879) 254 —

Rhaphidophora pteropoda (Teijsm. & Binn.) Engl., Bull. Soc. Tosc. Ortic.

4 (1879) 268 & in Beccari, Malesia 1 (1882) 268; Engl. & K. Krause in

Engl., Pflanzenr. 37 (1V.23B) (1908) 25-26; Ridl., Fl. Malay Penins. 5 (1925)

123, synon. noy. — Type: Indonesia, Sumatera, Loeboe-Aloeng, Teijsmann

s.n. (BO, holo; K, iso).

Rhaphidophora laetevirens Ridl., J. Bot. 40 (1902) 37 & Mat. Fl. Malay

Penins. 3 (1907) 43 — Type: Malaysia, Penang, Bk Penara, 1896, Ridley

s.n. (SING, lecto; selected here). Ridley cites two conspecific syntypes, the

other [Selangor, Kuala Lumpur Caves, Ridley s.n.| has not been traced,

and thus I have chosen the fertile Penang specimen as the lectotype.

Large, occasionally enormous, robust, semi-leptocaul, homeophyllous liane

to 20 m; seedling stage leafy at germination and skototropic by alternating

series of congested leafy and elongated leafless shoots; pre-adult plants

almost never forming terrestrial colonies; adult shoot architecture comprised

of greatly elongated, clinging, physiognomically monopodial, sparsely leafy,

non-flowering stems and short, free, sympodial, densely leafy, potentially

flowering stems; stems smooth, bright green, without prophyll, cataphyll

and petiolar sheath fibre, internodes to 20 x 3.5 cm on adherent shoots,

considerably shorter on free shoots, separated by prominent oblique leaf

scars, older stems sub-woody; flagellate foraging stems exceedingly long

with internodes to 50 cm or more and nodes with semi-persistent cataphylls;

clasping roots densely arising from the nodes and internodes of clinging

stems, prominently pubescent; feeding roots rare, adherent, pubescent; /eaves

weakly spiralled on adherent and flagelliform shoots, those on free shoots

distichous into dense few to many-leaved fans; cataphylls and prophylls

sub-coriaceous, soon drying and falling; petiole deeply canaliculate, 8-24 x

1.2-2.5 cm, smooth, apical and basal genicula prominent; petiolar sheath

prominent, extending to or occasionally, by means of a terminal brief

ligule, overtopping the apical geniculum, + long persistent and drying

chartaceous; /amina entire, falcate-lanceolate to falcate-oblong, oblique,

15-61 x 4-20 cm, sub-chartaceous, base acute to unequal, apex acute to

slightly acuminate; mid-rib prominently raised abaxially, slightly sunken

adaxially; primary venation pinnate, slightly raised abaxially, somewhat

impressed adaxially, the leaf appearing slightly quilted; interprimaries sub-

parallel to primaries, slightly raised abaxially, slightly impressed adaxially;

secondary venation weakly reticulate, slightly raised; tertiary venation

invisible; inflorescence solitary from the centre of the fanned leaves of a

free shoot, subtended by a fully developed foliage leaf; peduncle terete,

Rhaphidophora in Peninsular Malaysia and Singapore 199

LETTE.

L. GUAR.

Figure 1. Rhaphidophora angustata Schott

A. fertile shoot x zp B. foraging shoot x ae C. leaf lamina x gi venation detail x 4. All from

Nicolson 927.

Gard. Bull. Singapore 51 (1999)

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Figure 2. Rhaphidophora angustata Schott

A. flowering shoot x '/,; B. inflorescence, spathe fallen x '/,; C. spadix detail, late anthesis x 6;

D. spadix detail, early fruiting x 4. All from de Wilde & de Wilde-Duyfjes 13634.

Rhaphidophora in Peninsular Malaysia and Singapore 201

11-13 x 1-1.2 cm; spathe narrowly canoe-shaped, stoutly beaked, 16-29 x

2-3.5 cm, stiffly fleshy, greenish to white, gaping basally at anthesis and

then persistent partly into fruit development although eventually falling to

leave a large oblique scar at the base of the spadix; spadix cylindrical,

sessile, obliquely inserted on peduncle, 12—23 x 1.5—2 cm, dirty white; stylar

region rather well developed, mostly rhombohexagonal, 2—2.2 x 2 mm,

truncate; stigma punctiform, ca. 0.3 mm diam., but ovaries on lowermost

part of spadix with longitudinally orientated elongated stigmas ca. 1 x 0.25

mm; anthers not exserted at anthesis, pollen extruded from between ovaries;

infructescence 14—20 x 2.5-3.5 cm, dark green before ripening.

Distribution: Peninsular Malaysia (Penang, Perak, Selangor) and Sumatera.

Habitat: Primary to disturbed, moist to wet, lowland to upper hill dipterocarp

forest along rivers, on lofty trees and rocks on both acid and base substrates.

125 — 1500 m altitude.

Notes: 1. A frequently very high-climbing species unmistakable by the

scattered fans of large bright-green soft-textured litter-trapping leaves

carried on short stout shoots and held at about 90° to the + leafless main

stem. The large slender inflorescences arise from these fans.

2. Long known by the later name R. pteropoda, the epithet R. angustata

must now be applied to this species. The type of R. angustata is sterile (the

specimen is an immature fan of leaves but is unmistakably the same species

as that later described by Teijsmann and Binnendjik as Scindapsus

pteropodus) and of unknown provenance. In the protologue Schott cites

the type as ‘Sumatra occidentalis’ but the specimen label states ‘?Java’,

although R. angustata has never been collected wild in Java.

3. The syntype seen of Ridley’s name R. /aetivirens exactly matches R.

angustata. It is quite inexplicable that Ridley considered it to represent an

undescribed species, although later (Ridley, 1925) he merged it with R.

angustata (as R. pteropoda).

Other specimens seen: PENINSULAR MALAYSIA: Perak - Scortechini

s.n. (K); Bk. Larut Wray 2457 (SING), Derry sub. Curtis 3713, (K, SING),

Boyce 692, (K, KEP); Selangor - Hutan Lipur Kanching Boyce s.n. (sight

record — no voucher); Kepong, track to Bk. Lagong. ca. 1 km from GDZ

Guest House, Boyce 1064 (K, KEP).

202 Gard. Bull. Singapore 51 (1999)

2. Rhaphidophora beccarii (Engl.) Engl.

Rhaphidophora beccarii (Engl.) Engl. in Bot. Jahrb. Syst. 1 (1881) 181 & in

Beccari, Malesia 1 (1882) 270, Tab. xix 6 — 9; Hook.f, Fl. Brit. India 6

(1893) 546; Ridl., Mat. Fl. Malay Penins. 3 (1907) 44; Engl. & K. Krause in

Engl., Pflanzenr. 37 ([V.23B) (1908) 46; Alderw., Bull. Jard. Bot. Buitenzorg

III, 1 (1920) 382; Alderw., Bull. Jard. Bot. Buitenzorg HI, 4 (1922) 341;

Ridl., Fl. Malay Penins. 5 (1925) 124; Henderson, Malayan Wild Flowers,

Monocots, (1954) 238, Fig. 142 C, D — Epipremnum beccarii Engl. in Bull.

Soc. Tosc. Ortic. 4 (1879): 268 (1879) — Type: Malaysia, Sarawak, Kuching,

Oct. 1865, Beccari p.b. 833 (cited as ‘832’ by Engler, 1879) (FI, holo)

Rhaphidophora fluminea Ridl., J. Straits Branch Roy. Asiat. Soc. 44 (1905)

186; Engl. & K. Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 37, synon.

nov. — Type: Malaysia, Sabah, Bongaya, Dec. 1897, H.N. Ridley s.n. (SING,

holo)

Rhaphidophora borneensis Eng]., Araceae exsicc. et illustr. n. 195, see Engl,

Bot. Jahrb. Syst. 7, Beibl. 15 (1886) 1 — Type: Indonesia, Kalimantan,

Mindai to Pramassan, 19 June 1882, H. Grabowski s.n. (B+, holo; BM, iso).

Small to medium-sized, heterophyllous, sometimes homeophyllous, creeping

rheophyte, very rarely short liane, to 75 cm; seedling stage a non-skototropic

shingling shoot, soon becoming spreading-leafy; pre-adult plant initially

with + apressed shingle-leaves, later with leaves erect or spreading and at

this stage plants resembling adult in all but overall size and leaf division;

adult shoots all sympodial, clinging and flowering but main axis comprised

of longer modules than axillary axes; stems smooth, mid- to dark green,

with very sparse petiolar sheath fibre, internodes 1-7 x c. 1 cm, separated

by variously prominent slightly oblique leaf scars; flagellate foraging stems

absent; clasping roots very densely arising from the nodes and internodes

of clinging stems, pubescent; feeding roots very strongly adherent, densely

scaly; leaves distichous, apressed and shingling to erect or spreading and

scattered on pre-adult shoots, tending to become distally clustered on adult

shoots; cataphylls and prophylls membranaceous, soon drying and falling;

petiole narrowly canaliculate to carinate, 8-31 x 0.3-1.5 cm, smooth, apical

and basal genicula prominent; petiolar sheath prominent, extending to the

apical geniculum, variably persistent and mostly degrading into semi-

persistent weak fibres; /amina entire in seedling and pre-adult individuals,

entire, pinnatipartite or pinnatisect in adult plants, narrowly-lanceolate to

oblong-elliptic, slightly oblique, 21-51 x 2-23 cm, sub-coriaceous to slightly

fleshy, base decurrent, apex acuminate with a moderately prominent tubule;

Rhaphidophora in Peninsular Malaysia and Singapore 203

se a

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Figure 3. Rhaphidophora beccarii (Engl.) Engl.

A. habit x 1/ , B. leaf lamina x = > C. leaf lamina x '/ , D. venation detail x 4; E. inflorescence,

spathe fallen x 1; F. spadix detail, post female receptivity, pre-anthesis x 8. A, E & F from

Stone 9637;,B & D from Ridley s.n.;C from Kiew 1982.

204 Gard. Bull. Singapore 51 (1999)

mid-rib prominently raised abaxially, sunken adaxially; primary venation

pinnate, raised abaxially, slightly impressed adaxially; interprimaries sub-

parallel to primaries, slightly raised abaxially, + flush adaxially, often

forming a weak reticulum; secondary venation prominently reticulate,

slightly raised; tertiary venation a network of broadly spaced tessellate

veins arising at c. 90° from the mid-rib and crossing the primaries and

interprimaries; inflorescence one to three together, subtended by a

prominent cataphyll degrading to fibres before anthesis; peduncle terete,

8-12 x 0.2-0.4 cm; spathe narrowly canoe-shaped, stoutly beaked, 6.5-—7 x

1-1.5 cm, stiffly fleshy, greenish to dull white, swiftly falling at anthesis;

spadix cylindrical, sessile, inserted perpendicular to peduncle, 4.5—7 x 0.6—1

cm, dull whitish; sty/ar region rather well developed, mostly

rhombohexagonal, c. 1-1.5 x 1-1.5 mm, truncate; stigma elliptic,

longitudinally orientated, occasionally almost circular, c. 0.5 x 0.3 mm,

often very prominent especially in dried material; anthers exserted at

anthesis; infructescence 7-9 x 1-2 cm, mid-green when ripe.

Distribution: Peninsular Malaysia (Johore, Kelantan, Negri Sembilan,

Pahang, Penang, Perak, Selangor, Terengganu), Thailand (southern

peninsula), Sumatera and throughout Borneo.

Habitat: Rheophytic on rocky wooded streambanks, on rocks in streams

and rivers, on soft mud, sandy beds and bare rock on limestone or granite

in primary to disturbed old secondary lowland to hill and swamp forest.

70-900 m altitude.

Notes: 1. One of only two certainly obligate rheophytic species (the other

is R. wentii Eng. & K. Krause — New Guinea), R. beccarii is immediately

recognizable by its adult growth form, creeping along water courses or

attached on rocks in the water, and by the usually pinnately divided leaf

laminas.

2. Although the pinnately divided leaf is typical of adult plants, entire,

stenophyllous, lanceolate-leaved flowering plants are not rare. Such plants

have been referred to as R. fluminea and occur either as pure stands or as

mixed populations with the pinnate-leaved form (SFN 33/28 is an excellent

example of the latter phenomenon). They are treated here as a neotenic

manifestation of R. beccarii.

3. Rhaphidophora borneenis Engler, based on H. Grabowski s.n. from

Kalimantan, is a broad-leaved form of typical (i.e., divided-leaf) R. beccarit.

Rhaphidophora in Peninsular Malaysia and Singapore 205

Other specimens seen: PENINSULAR MALAYSIA: Johore - G. Pulai, Sg.

Ayer Hitam, above bridge at base of hill, Sinclair SFN 10832 (E, SING);

G. Panti Corner SFN 30961 (BO, SING), Bogner 366 (K, M), Kangka

Sedili Kechil, Corner SFN s.n. (SING); Sg Salat on Sg Endau, Kiew 1982

(KEP); Kelantan - Gua Musang, UNESCO Limestone Expd. 281 (K, L,

SING); Negri Sembilan, Ulu Bendol, FR north of road, Holttum SFN 9842

(SING), G. Beremban, Sg. Bendol, Furtado SFN 33128 (B, BO, K, L,

SING); Pahang - Bentong, Furtado SFN 33111 (K, P); Sg Talian, Ridley

2387 (K); Taman Negara, ALM, Zai & Ham s.n. (UKMB), Sg Tahan at

Lata Berkoh, Stone 13824 (KLU); Sg Yu, Hardial & Nor 27 (K, L, SING),

Sg Pertang, Ulu Tembeling, Henderson SFN 22120 (BO, SING); Sg Cheka,

Holttum SFN 24764 (BO, SING); Sg Jeletoh, Hardial & Nor 95 (L, SING );

Penang - Dalhousie s.n. (E), road from Penara Bukit to Ayer Hitam, Ridley

s.n. (SING); Perak - Scortechini s.n. (E, LE), Matang, Bubu F.R.,

compartment 77, banks of Sg Wang, Boyce 696 (K, KEP); Sg Tampang,

foot of G. Chantik, Gopeng, Furtado SFN s.n. (P, SING); ); Bk Larut

Besar, Kunstler 2874 (FI, K, P); Larut, Kunstler 3955 (K, L, P), Kunstler

3963 (L); Kunstler 10155 (BM, K), Temangoh, banks of Kertam river,

Ridley 14369, (BM, SING), Sg Chenderiang, Shah & Shuker 3414 (KEP,

SING); Selangor - Ulu Kerling, Hardial 13 (K, SING); Ulu Lui, Langat,

Hassan & Kasim UKMB 09039 (UKMB; Ulu Langat, Ulu Pansoon, Kasim

et al. 439 (UKMB); Sg Lalang F.R., Seminyih, Kiew 652 (KEP), Bk Lagong

F.R., Blanc 459 (KEP); stream half a mile before Klang Gates Resevoir,

Nicolson 1138 (L, SING, US);); Rawang, Ridley 7611 (K); Genting Bidai,

Ridley 7664 (K, SING); Ulu Gombak, Ridley s.n. (K), Ridley s.n. (K);

Templer Park, Samek UKMB 09947 (UKMB), Stone 9637 (KLU, L), Stone

12125 (KLU); 16th mile Ulu Gombak road at the Universiti of Malaya

Field Studies Centre, Stone FSC 321 (L), Stone 15384 (MO, KEP), Chin

1011 (L); G. Tangkau, Pansom, Ulu Langat, Umbai (for Hilliard) KL

1158 (KEP); Terengganu - Batu Biwa, Kiew 2348 (KEP).

3. Rhaphidophora corneri P.C.Boyce, sp. nov.

Rhaphidophora corneri differt a speciebus ceteris laminis foliorum parvis

oblongo-lanceolatis nervatura reticulata prominente plerumque

perforationibus nonnullis amplis ovatis vel rhomboideis vel trapezoideus;

inflorescentiis ex sympodiis floralibus elongatis distantibus orientibus;

sympodiis in apicibus ramorum lateralium liberorum positis — TYPUS:

Malaysia, Terengganu, Ulu Kajang, 13 Nov. 1935, E.J.H. Corner SFN 30441

(SING, holo).

206 Gard. Bull. Singapore 51 (1999)

Small, slender, heterophyllous?, leptocaul (?), liane of unknown ultimate

height; seedling stage and pre-adult plant unknown; adult shoot known

from flowering portions only, these either very long sympodia or

physiognomically monopodial (not clear from specimen), free but producing

one or two stout feeding roots from each node (adult plant thus perhaps

resembling Scindapsus scortechinii and therefore representing an

architectural type hitherto unrecorded in Rhaphidophora); stems smooth,

without prophyll, cataphyll and petiolar sheath fibre, internodes to 1-9 x

0.5—0.7 cm, separated by prominent corky leaf scars; flagellate foraging

stems not observed (absent?); clasping roots not observed; feeding roots

one or two per node, non-adherent (always?), stout, corky; /eaves distichous,

erect or spreading on adult shoots; cataphylls and prophylls membranaceous,

soon drying and falling; petiole narrowly canaliculate, 6-9 x 0.2-0.3 cm,

smooth, apical and basal genicula prominent; petiolar sheath obscure except

for basal-most portion, extending (always?) to the apical geniculum, sheath

of newest leaf degrading into a very few feeble fibres, sheath soon (?)

falling to leave a proportionally wide, corky scar basally on the petiole;

lamina entire or with a few, large ovate to rhomboid or trapezoid

perforations extending almost to the margin, 11-15 x 6-8 cm, oblong-

lanceolate, slightly oblique, sub-coriaceous, base obtuse to weakly cordate,

apex acute to acuminate with a slightly prominent apiculate tubule; mid-

rib prominently raised abaxially, slightly raised adaxially; primary venation

pinnate but distal-most veins becoming weakly reticulate and not reaching

leaf margin, raised abaxially, raised, but weakly so, adaxially; interprimaries

weakly reticulate to sometimes sub-parallel to primaries, slightly raised

abaxially, weakly raised adaxially, often forming a weak reticulum;

secondary venation reticulate, slightly raised abaxially, weakly raised

adaxially; tertiary venation prominently reticulate; inflorescence two (more?)

together, each subtended by a prominent swiftly falling cataphyll, and arising

sequentially on an elongated reiterative floral sympodia at the tip of

(probably) plagiotropic free lateral shoots; peduncle terete, 1—-1.3 x 0.2-

0.25 cm; spathe canoe-shaped, thick, stout-beaked, c. 2.7—2.9 x 0.7-0.8 cm;

spadix cylindrical, sessile, inserted slightly obliquely on peduncle, c. 2 x

0.42 cm; stylar region well developed, rounded-rhombohexagonal, c. 3—3.2

x 1.95-2 mm, convex to truncate, smoothly rounded; stigma impressed

irregularly elliptic, longitudinally orientated, c. 1 x 0.5 mm; anthers exserted

(?) at anthesis; infructescence not observed.

Distribution: Peninsular Malaysia (Terengganu). Endemic, known only from

the type collection.

Habitat. Unknown.

Rhaphidophora in Peninsular Malaysia and Singapore D207

NS AWS

Wh,

Na \ ay ly

(

L.GURR.

Figure 4. Rhaphidophora corneri P.C. Boyce

A. habit x '/,; B. leaf lamina x '/,; C. venation detail x 3; D. tip of flowering shoot with two

inflorescences, abaxial view x 1; E. tip of flowering shoot with two inflorescences, adaxial

view x 1'/,; F. spadix detail, post female receptivity, pre-anthesis x 6. All from Corner SFN30441.

208 Gard. Bull. Singapore 51 (1999)

Notes: 1. The manner of inflorescence production in R. corneri is unique in

the genus. The inflorescences arise sequentially from an elongated

reiterative floral sympodium on the tips of (probably) plagiotropic free

lateral shoots with each new inflorescence carried some distance from that

preceeding. The specimen studied bears the scars of several inflorescences

together with two developing inflorescences at different stages of maturity,

suggesting that the sympodia reiterate over a long period of time. The only

other species with a similar type of inflorescence production (but there on

the adherent orthotropic shoots and with the inflorescences congested) are

the New Guinea R. ledermannii Engl. & K. Krause and R. versteegii Engl.

& K. Krause.

2. Although represented by just the collection cited above I have no

hesitation in describing this species as new. Rhaphidophora corneri is a

remarkable species, unique in the genus by the manner of inflorescence

production (see note above), and small leaves with dense reticulate veins,

and large perforations (in the latter feature the species resembles Amydrium

hainanense (C.C. Ting & C.Y. Wu ex H. Li et al.) H. Li (see Nguyen &

Boyce, 1999). It would be very desirable to learn more about this plant

from additional field collections.

3. Rhaphidophora corneri is named for the late Prof. E.J.H. Corner, a

giant among botanists of Asia. Furtado first proposed the name in sched.,

but it was never published.

4. Rhaphidophora crassifolia Hook./.

Rhaphidophora crassifolia Hook.f., Fl. Brit. Ind. 6 (1893) 543; Ridl., Mat.

Fl. Malay Penins. 3 (1907) 42; Engl. & K. Krause in Engl., Pflanzenr. 37

(IV .23B) (1908) 22, Fig. 5; Ridl., Fl. Malay Penins. 5 (1925) 122 — Type:

Malaysia, Perak, Batang Padang District, Batang Padang, August 1884,

Kunstler 8111 (K, holo).

Large, occasionally very large, moderately robust, semi-pachycaul

homeophyllous liane to 10 m; seedling stage a non-skototropic shingling

shoot; pre-adult plants rarely forming small terrestrial colonies of apressed

shingling shoots; adult shoot architecture comprised of greatly elongated,

clinging, physiognomically monopodial, leafy, non-flowering stems and short

to moderately elaborated, free, sympodial, densely leafy, flowering stems;

stems smooth, climbing stems rectangular in cross section, the angles winged,

the surfaces between sulcate, the stem twisting to give a barley-sugar effect,

Rhaphidophora in Peninsular Malaysia and Singapore 209

Figure 5. Rhaphidophora crassifolia Hook.f.

A. Pre-adult shoot x */,; B. adult, sterile shoot x */,+; C. flowering shoot x '/,; D. leaf lamina x ee

E. venation detail x 4; F. inflorescence, just prior to spathe opening x 1; G. spadix detail, pre-

female receptivity x 6.A & B from Boyce 1226; C - E from Ngadiman SFN 36774, F & G from

Corner SFN 29318.

210 Gard. Bull. Singapore 51 (1999)

free stems more or less terete to weakly four-angled in cross section, often

branching extensively, growing to considerable lengths and then pendent

under their own weight, green, later dull brown, without prophyll, cataphyll

and petiolar sheath fibre but active apices coated with clear mucilage,

internodes to 10 x 2 cm on adherent shoots, usually shorter and less stout

on free shoots, separated by prominent slightly oblique leaf scars, older

stems woody; flagellate foraging stems absent; clasping roots densely arising

from the nodes and internodes of clinging stems, densely pubescent; feeding

roots rather rare, adherent, pubescent; /eaves weakly spiralled on adherent

and proximal portions of free shoots, densely distichous distally on flowering

shoots; cataphylls and prophylls membranaceous, very quickly drying and

falling; petiole shallowly canaliculate to grooved adaxially, 4-7 x 0.2-0.3

cm, smooth, with a slight apical and somewhat prominent basal geniculum;

petiolar sheath prominent, extending to and encircling the apical geniculum,

very swiftly drying and falling to leave a continuous scar from the petiole

base, around the top of the apical geniculum and back to the base; /amina

entire, falcate-elliptic-lanceolate to falcate-oblong or falcate-oblanceolate,

4.5-25.5 x 1.5-5 cm, thickly coriaceous to almost fleshy, upper surfaces

glossy, lower surfaces matt, base sub-ovate to acute or briefly decurrent,

apex sub-acute with a prominent apiculate tubule, margins slightly reflexed,

this becoming greatly accentuated in dried material; mid-rib barely raised

abaxially, slightly sunken adaxially; primary venation pinnate, slightly raised

abaxially and adaxially; interprimaries parallel to primaries, very slightly

raised abaxially and adaxially; secondary and tertiary venation + invisible

in fresh material, barely visible in dried specimens; inflorescence solitary,

subtended by a fully developed foliage leaf; peduncle compressed-cylindric-

clavate, 1.7—2.5 x 0.2-0.3 cm; spathe cigar-shaped, stoutly short-beaked,

4—6.5 x 1-1.7 cm, thickly fleshy, dull green to dull yellow, swiftly falling at

female receptivity; spadix cylindrical, sessile, inserted level on peduncle,

3.5-6 x 0.5-1.3 cm, dirty white; stylar region rather well developed, mostly

rhombohexagonal, 0.8—1 x c. 1 mm, truncate; stigma punctiform, c. 0.3 mm

diam.; anthers barely exserted at anthesis, pollen extruded from between

ovaries; infructescence not observed.

Distribution: Peninsular Malaysia (Johore, Negri Sembilan, Selangor) and

southern peninsular Thailand.

Habitat: Primary lowland rainforest, dry Dryobalanops forest, peat forest,

wet evergreen forest. 10-90 m altitude.

Notes: 1. Rhaphidophora crassifolia is one of the species here included in

Rhaphidophora in Peninsular Malaysia and Singapore DN

the informal Hongkongensis Group, an assemblage species defined by

clinging stems rectangular in cross-section, simple, lanceolate to elliptic,

frequently falcate leaves, usually rapidly falling cataphylls and prophylls

and petiolar sheaths, the latter shedding to leave a continuous scar around

the top of the apical geniculum, and small to medium-sized cigar-shaped

inflorescences carried on the tips of free lateral stems. The group is

taxonomically very difficult in part because the various species recognized

are separated by quantitative differences and in part because, being mostly

high climbing lianes, they are often woefully inadequately collected. I have

taken a pragmatic approach with their taxonomy for Peninsular Malaysia,

recognizing only those species that can be keyed out (even then, often with

some difficulty). While this is clearly far from satisfactory and has resulted

in a large number of synonyms, in particular with R. sylvestris and R.

montana, it means that identifications can be made with some confidence,

a situation near impossible if all names are regarded as ‘good’.

2. In Peninsular Malaysia, R. crassifolia is most easily confused with R.

sylvestris with which it shares decidedly falcate leaves and small (less than

7 cm long) inflorescences. It may be distinguished from R. sy/vestris in the

considerably thickened leaves, drying with prominently recurved margins,

by the glossy adaxial and matt abaxial leaf surfaces, and by the indistinct

venation, an artefact of the fleshy leaves and, in the field, by the prominently

angled, twisted adherent stems.

3. Confusion with R. montana is possible. Generally R. montana is a plant

far larger in all its parts, especially in the larger inflorescences and longer

spadix.

4. The similar R. maingayi Hook.f. may be distinguished by the shoot

apices covered with persistent untidy fibre derived from the degraded

prophylls, cataphylls and petiolar sheaths.

5. Work is underway with DNA studies of the Hongkongensis Group to

resolve the phylogeny of the group and, it is hoped, shed light on the

species boundaries.

Other specimens seen: PENINSULAR MALAYSIA: Johore - Pengkalan

Raja, Pontian, Ngadiman SFN 36774 (B, BO, L, K, P, SING); Sg Kayu

Ara, Mawai-Jemaluang road, Corner SFN 29318 (K, KEP, SING); Negeri

Sembilan - Pasoh FR La Frankie 2185 (KEP); Selangor - Genting Sempah

22 mile, Ulu Gombak FR, Stone 5738 (KLU).

212 Gard. Bull. Singapore 51 (1999)

5. Rhaphidophora falcata Ridl.

Rhaphidophora falcata Ridl., J. Asia. Soc. Straits Br. 44 (1905) 198; Engl.

& K. Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 37 — Type: Thailand,

Bangtaphan, Keith s.n. (SING, holo).

Distribution: Peninsular Malaysia (Penang) and southern Thailand.

Medium, slender, leptocaul homeophyllous lane(?) to unknown ultimate

height; seedling, pre-adult plants and adult shoot; stems smooth, terete,

without prophyll, cataphyll and petiolar sheath fibre, internodes 1-2 x 0.5-

0.7 cm, separated by rather well-defined, slightly oblique leaf scars, older

stems woody; flagellate foraging stems not observed; roots sparsely arising

from the nodes and internodes of stems, densely pubescent; cataphylls and

prophylls very quickly drying and falling; petiole adaxially basally

canaliculate, otherwise grooved, 11-19 x 0.2-0.5 cm, smooth, apical genicula

very prominent, basal geniculum poorly defined; petiolar sheath very

prominent basally, extending c. half way along the petiole, swiftly drying

and but seemingly persistent; /amina entire, lanceolate-falcate, very slightly

oblique, 16-20 x 2-2.5 cm, stiffly coriaceous, base acute to cuneate, apex

acute, with a prominent apiculate tubule; mid-rib prominently raised

abaxially, slightly sunken adaxially; primary venation densely pinnate,

slightly raised abaxially and adaxially; interprimaries parallel to primaries

and hardly less prominent, slightly raised abaxially and adaxially; secondary

and tertiary venation reticulate to sub-tessellate in dried specimens;

inflorescence solitary (always?), subtended by a fully developed foliage;

peduncle and_ spathe not observed; spadix cylindrical, sessile, inserted +

level on peduncle, c. 2.5 x 1.3 cm; stylar region, rounded, 1.5-1.7 x 1.6—2

mm, truncate; stigma punctiform, raised, c. 0.7 x 0.8 mm diam.; anthers not

observed; infructescence not observed.

Habitat: Unknown.

Notes: 1. Recognized as distinct, despite the dearth of material and

information, due to the falcate, stiffly coriaceous leaves, unique in the

genus. This species will be illustrated in the account of Thai Rhaphidophora.

2. The life form of R. falcata is not clear from the meagre material available.

The Cantley collection (erroneously labelled ‘Haniff?’) cited by Ridley

(1905) has traces of sandy mud on the stems and roots, suggesting that the

plant is terrestrial and perhaps rheophytic, and thus implying a relationship

to, for example, R. beccarii and R. wentit.

Rhaphidophora in Peninsular Malaysia and Singapore 21D

Other specimens seen: PENINSULAR MALAYSIA: Penang - Cantley s.n.

(SING).

6. Rhaphidophora foraminifera (Engl.) Engl.

Rhaphidophora foraminifera (Engl.) Engl., Pflanzenr. 37 (IV.23B) (1908)

45; Alderw., Bull. Jard. Bot. Buitenzorg III, 4 (1922) 197-198; Ridl., FI.

Malay Penins. 5 (1925) 123-124 — Epipremnum foraminiferum Engl.,

Bot. Jahrb. Syst. 25 (1898) 11 — Type. Neotype: Malaysia, Perak, Taiping,

based of Maxwell Hill Road, Nicolson 1047 (L neo; BH, BO, US isoneo

designated here — see discussion below).

Moderate to large, robust, pachycaul, homeophyllous liane to 15 m; seedling

stage not observed; pre-adult plants frequently (always?) forming extensive

terrestrial colonies; adult shoot architecture comprised of elongated, clinging,

physiognomically monopodial, densely leafy, non-flowering (always?) stems

and shorter clinging, sympodial, densely leafy, flowering stems; stems

smooth, mid-green, with cataphylls and prophylls deliquescing to black

mucilage drying to leave fragmentary parchment-like remains on petioles,

developing laminae, inflorescences, internodes 1-7 x 0.5—3.5 cm, separated

by prominent straight to slightly oblique leaf scars; flagellate foraging stem

absent; clasping roots densely arising from the nodes and internodes,

pubescent; feeding roots, adherent, exceedingly robust, densely pubescent

with dense verticels of prominent corky ramenta; leaves distichous;

cataphylls and prophylls membranaceous, soon deliquescing; petiole

canaliculate, 22-52 x 0.4-1 cm, smooth for the most part but upper 2-4 cm

and apical geniculum densely pubescent, apical geniculum prominent, basal

geniculum less so; petiolar sheath prominent, extending to apical geniculum,

+ short-persistent, degrading to weak, slightly netted fibres; /amina entire

to slightly or extensively perforated, perforations round to rhombic,

extending c. '/, of lamina width on each side of the midrib, ovate to oblong-

lanceolate or oblong-elliptic, slightly oblique, pubescent abaxially when

young, 7-53 x 6-19 cm, membranaceous to sub-coriaceous, base rounded,

acute to slightly decurrent, apex acute to acuminate; mid-rib prominently

raised and densely pubescent abaxially, + flush adaxially; primary venation

pinnate, slightly raised abaxially and adaxially, the leaf appearing slightly

quilted, pubescent in younger leaves, this indumentum mostly shed in older

leaves; interprimaries sub-parallel to primaries, less prominent, slightly raised

abaxially, slightly impressed adaxially; secondary venation reticulate, slightly

raised; tertiary venation tessellate, slightly raised; inflorescence mostly

subtended by one or two large cataphylls, these swiftly deliquescing into

214 Gard. Bull. Singapore 51 (1999)

) Sa ee ‘\ .

L GWAR.

Figure 6. Rhaphidophora foraminifera (Engl.) Engl.

A. pre-adult climbing shoot x '/,; B. pre-adult terrestrial shoot x '/,; C. leaf lamina x '/,.A & B

from Nicolson 941; C from Boyce 722.

Rhaphidophora in Peninsular Malaysia and Singapore 2AS

black mucilage, this drying and adhering patchily to developing spathe,

rarely subtended by a + fully developed foliage leaf; peduncle terete, 3-13

x 0.5-1.5 cm; spathe canoe-shaped, stoutly beaked, 10-27 x 1.5-13 cm,

stiffly fleshy, dull pale to dark yellow, wide gaping at anthesis and then

briefly persistent though maturation of the stamens, eventually falling to

leave a large + scar at the base of the spadix; spadix cylindrical, sessile,

slightly obliquely inserted on peduncle, 7-17 x 1.3-2.5 cm, dull greenish

yellow; stylar region moderately developed, rounded to rhombohexagonal,

1-2 x 1-1.5 mm, shortly conical when fresh, drying truncate; stigma

punctiform or elliptic and transversely orientated, raised at anthesis but

excavated in dry material, c. 0.3 mm diam.; anthers slightly exserted at

anthesis; infructescence 8-15 x 2—3 cm, dark green before ripening to

greenish yellow.

Distribution: Peninsular Malaysia (Pahang and Perak), Sumatera, and

throughout Borneo.

Habitat: Disturbed forest, moist hill dipterocarp forest, on rocks (granite

and limestone) in shaded position. 50-1035 m altitude.

Notes: 1. The type specimen of Epipremnum foraminiferum was sent to

Berlin from Bogor in 1897 (fide Engler, 1898). No such specimen is extant

in Berlin and the type must be considered lost. The collection chosen as

the neotype is fertile and matches Engler 5307, a sterile collection, which is

annotated as R. foraminifera in Engler’s handwriting and matches Engler’s

usage of the name R. foraminifera.

2. There are two Rhaphidophora species in Peninsular Malaysia with

perforated leaves and variously pubescent leaves: R. foraminifera and R.

puberula. They may be distinguished by the position of the inflorescence

(on short adherent shoots in R. foraminifera, on short free shoots in R.

puberula), by the presence of black mucilage produced by the deliquescing

cataphylls and prophylls (present in R. foraminifera, absent in R. puberula),

in leaf size of mature flowering-sized plants (to 53 x 19 cm in R. foraminifera,

34 x 10.5 cm in R. puberula), by the more or less rounded (R. foraminifera)

and oblong (R. puberula) stylar regions, and in overall size of the plant (R.

foraminifera is a bole climber to 15 m whereas R. puberula seldom climbs

higher than 3 m and frequently forms large, spreading masses on rocks).

3. Perforated leaves occur in a number of otherwise rather different looking

Rhaphidophora species (e.g. R. foraminifera. R. puberula, R. corneri, R.

subfalcata M. Hotta, R. ledermannii, R. veersteegii) suggesting that, while a

216 Gard. Bull. Singapore 51 (1999)

ES AS Lari ys

- OT Wara: ow

CMAQ)

. : iN WW ‘i \

OVA =

RA XN NN

4.Gure

Figure 7. Rhaphidophora foraminifera (Engl.) Engl.

A. flowering shoot x ’/ .; B. venation detail x 6; C. inflorescence, spathe fallen x */,; D. spadix

detail, anthesis x 6.A,C & D from Boyce 722; B from Croat 53118.

Rhaphidophora in Peninsular Malaysia and Singapore 2AT

useful diagnostic tool, lamina perforation cannot be used to specify groups

within Rhaphidophora.

Other specimens seen: PENINSULAR MALAYSIA: Pahang - Taman

Negara, Batu Luas (limestone) Kiew 13519 (KEP); Perak - Ridley s.n.

(BM); Bk Larut, Boyce 690 (K, KEP), Hay, Anthony & Banka 9073 (NSW,

KEP), Taiping, Maxwell Hill, Engler 5307 (B).

7. Rhaphidophora korthalsii Schott

Rhaphidophora korthalsii Schott, Ann. Mus. Bot. Lugd.-Bat. 1(1863) 129;

Engl. in A. & C. DC., Monogr. Phan. 2 (1879) 246; Hook.f, Fl. Brit. India

6 (1893) 548; Ridl., Mat. Fl. Malay Penins. 3 (1907) 45; Alderw., Bull. Jard.

Bot. Buitenzorg II, 4 (1922) 341; Engl. & K. Krause in Engl., Pflanzenr.

37 (IV.23B) (1908) 49-51, Fig. 21; Ridl., Fl. Malay Penins. 5 (1925) 123;

Henderson, Malayan Wild Flowers, Monocots, (1954) 238-239, Fig. 142 B

— Type: Indonesia, Java, P.W. Korthals s.n. (L, holo; L, P, iso).

Pothos celatocaulis N.E. Br., Gard. Chron. 13 (1880) 200, synon. nov. —

Rhaphidophora celatocaulis (N.E. Br.) Alderw., Bull. Jard. Bot. Buitenzorg

III, 1 (1920) 382; Alderw., Bull. Jard. Bot. Buitenzorg III, 4 (1922) 198,

synon. nov. — Type: Malaysia, NW Borneo (Sabah), Burbidge s.n., Hort.

Veitch no. 215 (K, holo; K, iso).

Rhaphidophora maxima Engl., Bull. Soc. Tose. Ortic. 4 (1879) 269 & in

Beccari, Malesia 1 (1882) 271, Tab. xx 1 — 5; Engl. & K. Krause in Engl.,

Pflanzenr. 37 (IV.23B) (1908) 48 — 49; K. Krause & Alderw., Nova Guinea

14 (1924) 214, synon. nov. — Type: Sarawak, Gunung Gading, July 1866,

Beccari PB2314 (FI, lecto; selected here). Engler lists two syntypes in

describing R. maxima. That chosen is the more complete of the two, clearly

conspecific, collections. The other is Beccari PP443 [Irian Jaya, Soron (FI)].

Rhaphidophora tenuis Engl., Bot. Jahrb. Syst. 1 (1881) 181 & in Beccari,

Malesia 1 (1882) 271-272; Engl. & K. Krause in Engl., Pflanzenr. 37

(IV.23B) (1908) 53, synon. nov. — Types: Malaysia, Sarawak, Beccari

PB1977 (FI lecto; B isolecto; selected here). Of the two conspecific syntypes

cited by Engler, that chosen is more complete. The other, Beccari PB2714

(B, FI) also originates in Sarawak.

Rhaphidophora korthalsii Schott var. angustiloba Ridl. ex Engl. & K. Krause

in Engler, Pflanzenr. 37 ([V.23B) (1908) 49, synon. nov. — Type: Malaysia,

218 Gard. Bull. Singapore 51 (1999)

Sarawak, Matang, July 1903, Ridley s.n. (SING, lecto; selected here). Engler

cites two syntypes, the other, Engler 5267 [Selangor, Batu Caves] was not

found in Berlin and is assumed destroyed.

Rhaphidophora copelandii Engl., Bot. Jahrb. Syst. 37 (1905) 115; Engl. &

K. Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 49, synon. nov. — Type:

Philippines, Mindanao, Davao, Mt Apo, April 1904, Copeland 1193 (PNH,

holot; B iso).

Rhaphidophora grandifolia K. Krause, Bot. Jahrb. 44, Beibl. 101 (1910) 11.

Type: Philippines, Negros, Negros Orientale, Dumaguete (Cuernos Mts),

March 1908, Elmer 9464 (PNH, holot; B, E, K, L, LE, MO, iso).

Rhaphidophora trinervia Elmer, Leafl. Philipp. Bot. 8 (1919) 3073, synon.

nov. — Type: Philippines, Laguna, Los Bafios (Mt Maquiling), June — July

L917; Elmer TS@e7 (PN, holo}; FijuK, 1b, MO, Prise),

Rhaphidophora ridleyi Merr., J. Str. Br. Roy. As. Soc. Special Edition

(Enum. Pl. Borneo) (1921) 90, synon. nov. — [Rhaphidophora grandis

Ridl., J. Straits Branch Roy. Asiat. Soc. 49 (1907) 51, nom. illeg., non

Schott 1858 (India)], synon. nov. — Type: Malaysia, Sarawak, Tambusan,

Sept. 1905, Ridley 12414 (SING, holo).

Rhaphidophora latifolia Alderw., Bull. Jard. Bot. Buitenzorg III, 4 (1922)

341; K. Krause & Alderw., Nova Guinea 14 (1924) 213, synon. nov. —

Type: Indonesia, Irian Jaya, Pionierbivak, 23 July 1920, Lam 711 (BO,

holo; L, iso). |

Rhaphidophora palawanensis Merr, Philipp. J. Sci. 26 (1925) 451, synon.

nov. — Type: Philippines, Palawan, Malampaya Bay, Oct. 1922, Merrill

BS 11570 (PNH, holot; B, K, P, iso)

Rhaphidophora trukensis Hosok., J. Jap. Bot. 13 (1937) 195, synon. nov. —

Type: Federated States of Micronesia, Chuuk (Truk) Island, near Orrip,

29 July 1939, Hosokawa 8334 (T1, holo).

[Epipremnum multicephalum Elmer, Leafl. Philipp. Bot. 10 (1938) 3624,

nom. nud., descr. Angl. — Based on: Philippines, Luzon, Sorsogon, Trosin

(Mt Bulusan), May 1916, Elmer 16061 (FI, K, L, MO, P, PNH7)].

Very large, occasionally enormous, slender to rather robust, pachycaul,

heterophyllous liane to 20 m; seedling stage a non-skototropic shingling

Rhaphidophora in Peninsular Malaysia and Singapore 219

U ey eo

Mee

Wes

(es

SWRA Re:

AAS

eff Z

een

= 2 = —s ~,

—— —— aN —>.

L. GUA,

Figure 8. Rhaphidophora korthalsii Schott

A. pre-adult shoot x '/,; B. pre-adult shingling shoot x '/,. A from Boyce 679; B from Nicolson

E12.

220 Gard. Bull. Singapore 51 (1999)

Figure 9. Rhaphidophora korthalsii Schott

A. flowering shoot, leaves removed x '/,; B. leaf lamina x '/,; C. venation detail x 2; D.

inflorescence, spathe removed x 1; E. spadix detail, post floral x 8. All from Kerr 15051.

Rhaphidophora in Peninsular Malaysia and Singapore Ze

juvenile shoot; pre-adult plants never forming terrestrial colonies; adult

shoot architecture comprised of greatly elongated, clinging,

physiognomically monopodial, densely leafy, flowering stems; stems smooth,

bright green, with sparse to copious prophyll, cataphyll and petiolar sheath

fibre, especially at the stem apices, internodes to 15 x 3.5 cm, separated by

prominent oblique leaf scars, older stems sub-woody; flagellate foraging

stems absent; clasping roots densely arising from the nodes and internodes,

prominently pubescent; feeding roots abundant, adherent and free, very

robust, densely ramentose-scaly; /eaves distichous; cataphylls and prophylls

membranaceous, soon drying degrading to intricately reticulate fibres, these

only very slowly falling; petiole shallowly grooved, upper part + terete, (1-)

9-65 x 0.2-1.5 cm, smooth, apical and basal genicula prominent; petiolar

sheath prominent, membranaceous, strongly to slightly unequal on one

side, extending almost to or reaching the apical geniculum, of + short-

duration, soon degrading into persistent netted fibres, these eventually

falling to leave a prominent, slightly corky scar; /Jamina of seedlings

overlapping in the manner of roof shingles, entire, lanceolate, 5-11 x 3.5-6

cm, base slightly cordate, lamina of pre-adult and adult plants free, entire,

pinnatipartite, pinnatisect or pinnatifid, 10-44 x 14—94 cm, broadly oblong-

elliptic to oblong lanceolate, slightly oblique, membranaceous to chartaceous

or sub-coriaceous, base truncate and very briefly decurrent, apex acute to

acuminate, individual pinnae 1—10 cm wide, frequently perforated basally

adjacent to the mid-rib, thus appearing stilted; mid-rib very prominently

raised abaxially, slightly sunken adaxially; primary venation pinnate, raised

abaxially, somewhat impressed adaxially, 2-4 primary veins per pinna;

interprimaries sub-parallel to primaries, slightly raised abaxially, slightly

impressed adaxially; secondary venation strongly reticulate, slightly raised;

tertiary venation invisible; inflorescence solitary to several together, first

inflorescence subtended by a membranaceous prophyll and one or more

cataphylls, these swiftly degrading to netted fibres, subsequent inflorescences

subtended by one or more swiftly degrading cataphylls, the whole forming

a mass of developing and open inflorescences and developing

infructescences partially concealed by persistent netted cataphyll and

prophyll remains; peduncle slightly laterally compressed to terete, 6-26 x

1-1.5 cm; spathe narrowly canoe-shaped, stoutly beaked, 10-30 x 3-5 cm,

stiffly fleshy, greenish to dull yellow, gaping wide at female receptivity and

then swiftly falling to leave a large straight scar at the base of the spadix;

spadix cylindrical, sessile, inserted + level on peduncle, 9-26 x 1.5—2 cm,

dull green to dirty white; stylar region rather well developed, mostly

rhombohexagonal, 1.5—2 x c. 2 mm, slightly conical; stigma punctiform to

slightly elliptic, if the latter then mostly longitudinally orientated, c. 0.3-0.5

x 0.2—0.4 mm; anthers barely exserted at anthesis; infructescence 14—27 x 3-

222 Gard. Bull. Singapore 51 (1999)

3.5 cm, dark green ripening to dull orange, stylar tissue abscising to reveal

orange ovary cavity pulp.

Distribution: Peninsular Malaysia (Kelantan, Pahang, Penang, Perak,

Selangor, Terengganu), Singapore. Widespread throughout tropical south

Asia from Sumatera and southern Thailand to Borneo and the Philippines

eastwards through the tropical western Pacific.

Habitat: Disturbed lowland primary or secondary dipterocarp forest, lower

and upper hill forest, wet pre-montane and montane forest, on granite,

sandstone, clay and limestone, occasionally in freshwater swamp forest.

10-1700 m altitude.

Notes: 1. Rhaphidophora korthalsii is a very widespread and variable species,

hence the extensive synonymy. However, as with Epipremnum pinnatum

(L.) Engl. (Boyce, 1998) there are several geographical elements that, given

more intensive study, might warrant formal taxonomic recognition.

Unfortunately current herbarium material is inadequate to confirm these

plants’ status and more field observations are needed.

2. Sterile herbarium material lacking the pre-adult stage may prove difficult

to distinguish from Epipremnum pinnatum. Mature leaves of ‘typical’ E.

pinnatum never have more than one primary lateral vein per pinna and the

stems of R. korthalsii lack the prominent irregular whitish longitudinal

crests and older stems the distinctive matt to sub-lustrous pale brown papery

epidermis typical of E. pinnatum. The feeding roots of R. korthalsii are

prominently scaly while those of E. pinnatum are lenticellate-corky. The

pre-adult stage of R. korthalsii is a shingle climber with oblong-elliptic to

ovate, slightly falcate upwards pointing leaves overlapping in the manner

of roof tiles.

3. Fertile material of R. korthalsii and Epipremnum pinnatum is readily

separated by the shape of the style apex (round versus trapezoid) and the

shape and orientation of the stigma (+ punctiform and circumferential

versus strongly linear and longitudinal) and, if fruits are mature, by seed

characters. The fruits of R. korthalsii each contain many small ellipsoid

seeds with a brittle, smooth testa whereas E. pinnatum has fruits with two

large, strongly curved seeds with a bony and ornamented testa.

4. In Peninsular Malaysia confusion with R. tetrasperma and R. nicolsonii

is possible. The former is readily separable by the generally smaller and

comparatively broader adult leaves (to 42 x 38 cm), and by the individual

Rhaphidophora in Peninsular Malaysia and Singapore 229

pinnae lacking the prominent parallel primary lateral veins and in never

being perforated close to the midrib. Rhaphidophora nicolsonii is

distinguished by the ovate leaves and by the petiolar sheath extending '/, —

*/, along the petiole.

Other specimens seen: PENINSULAR MALAYSIA: Kelantan - Tanah

Merah, Pergau Dam site, logging road to Sg Renyok Intake, Boyce 667 (K,

KEP); Pahang - Cameron Highlands, Henderson SFN 23475 (K, SING);

Fraser’s Hill, Henderson SFN 11426 (BO, SING), Nur SFN 11079 (BO,

SING); Taman Negara, between Sg Putih and Kuala Tahan, Soepadmo

809 (KLU); Pulau Tioman, Teker to Joara, Stone 11977 (KLU, MO); Bk

Koman, Zai & Kasim UKMB 02914 (UKMB); G. Senyum (limestone),

Henderson SFN 22318 (BO, SING); Penang - Government Hill, Maingay

2213 (K); Perak - Kledang Saiang Reserve, Arnot 33687 (KEP); Bk Larut,

Boyce 679 (K, KEP), Kunstler 6893 (CAL, K, UC), Ridley 11419 (K); G.

Hijau, Burkill & Haniff SFN 12586 (K, SING); Ipoh, above Perak Hill

Temple, Chin 997 (KLU) Tanjong Pondok FR, 12 miles north of Taiping,

Nicolson 1075 (SING, US); Selangor - Ulu Gombak, mile 22 on new

highway, Croat 53280 (MO); Dusan Mangis, Gombak FR, Farah s.n.

(UKMB); Selangor, between Fraser’s Hill and the Semangkok Pass, Burkill

FMS 7761 (BO, K, SING); Templer Park, Hassan 25 (UKMB); Anak

Takun (limestone), Templer Park, Hassan & Latif 140; Terengganu - Bk

Besi, Dunggun, Mk. Ar. UKMB 04958 (UKMB); (UKMB); SINGAPORE:

‘Singapore’, Wallich 9109 ( K); Bk Timah, Hullett 906 (K), 907 (K), Ridley

s.n. (SING); Seletar FR, near pipe line, Sinclair SFN 5480 (E, SING);

Seletar FR, behind Mee Soon village, Sinclair SFN 5516 (E, SING);

Northeast end of MacRitchie Reservoir, Sinclair SFN s.n (E, L, P, SING).

8. Rhaphidophora lobbii Schott

Rhaphidophora lobbii Schott, Prodr. Syst. Aroid. (1860) 379; Engl. in A. &

C. DC., Monogr. Phan. 2 (1879) 240; Hook.f., Fl. Brit. India 6 (1893) 544;

Ridl., Mat. Fl. Malay Penins. 3 (1907) 41-42; Engl. & K. Krause in Engl.,

Pflanzenr. 37 (1V.23B) (1908) 33-34; Ridl., Fl. Malay Penins. 5 (1925) 122;

Henderson, Malayan Wild Flowers, Monocots, (1954) 238, Fig. 142 A

[Rhaphidophora lobbii Schott, Bonplandia 5(2) (1857) 45, nom. tant.| —

Scindapsus lobbii (Schott) Ender, Index Aroid. (1864) 74 — Type:

Singapore, Lobb 44 (K, holo).

Small to moderate, somewhat slender, leptocaul, homeophyllous liane to 5

m; seedling leafy at germination and skototropic by alternating series of

224 Gard. Bull. Singapore 51 (1999)

Figure 10. Rhaphidophora lobbii Schott

A. portion of stem with two flowering shoots x '/,; B. pre-adult terrestrial shoots x '/,; C. stem

detail x 3; D. leaf lamina x */,; E. venation detail x 4; F. inflorescence, spathe fallen x 2; G.

spadix detail, early fruiting x 8. A & C from Kunstler 10571; B from Boyce 995; D & E from

Nauen s.n.; F & G from Alvins 270.

Rhaphidophora in Peninsular Malaysia and Singapore 225

congested leafy and elongated leafless shoots; pre-adult plants forming

diffuse terrestrial colonies; adult shoot architecture comprised of greatly

elongated, clinging, physiognomically monopodial, leafy, non-flowering

stems and long moderately elaborated, free, sympodial, densely leafy,

flowering stems; stems puberulent-scabrid to asperous, especially on older

growth, climbing stems weakly rectangular to + terete in cross section, free

stems + terete in cross section, often branching extensively and growing to

moderate lengths pendent under their own weight, dull brown, without

prophyll, cataphyll and petiolar sheath fibre, internodes to 13 x 0.6 cm on

adherent and free shoots, flowering shoots with much shorter internodes,

separated by weak to rather prominent, slightly oblique leaf scars, older

stems woody; flagellate foraging stems absent; clasping roots sparsely arising

from the nodes and internodes of clinging stems, densely pubescent; feeding

roots rather rare, sometimes adherent but often free, pubescent; /eaves

weakly spiralled and often sparsely arranged on adherent and proximal

portions of free shoots, densely spirally-distichous distally on flowering

shoots; cataphylls and prophylls membranaceous, very quickly drying and

falling; petiole grooved adaxially, 4-9.5 x 0.2-0.3 cm, smooth, with a

moderate apical and prominent basal geniculum; petiolar sheath slightly

prominent, extending beyond the apical geniculum by two ligules, very

swiftly drying and falling in strips to leave a continuous scar from the

petiole base, around the top of the apical geniculum and back to the base;

lamina entire, narrowly elliptic to elliptic-lanceolate to oblong or

oblanceolate, 6—24 x 2-10 cm, very softly coriaceous, upper surfaces slightly

glossy, lower surfaces pale satin-matt, drying markedly discolorous, dark

brown above, pale brown below, base cuneate to acute or sub-ovate, briefly

decurrent, apex acute to ovate-acuminate, with a prominent apiculate

tubule; midrib slightly raised abaxially, slightly sunken adaxially; primary

venation pinnate, slightly raised abaxially, prominent (dark veins against

pale lamina) in dried material; interprimaries parallel to, but much less

distinctive than, primaries, very slightly raised abaxially; secondary and

tertiary venation + invisible in fresh material, barely visible in dried

specimens, reticulate; inflorescence solitary, subtended by a fully developed

foliage leaf and a very quickly falling cataphyll; peduncle compressed-

cylindric, 1.5—-5 x 0.15-0.4 cm; spathe cigar-shaped, stoutly long-beaked,

3-5 x 0.4-1 cm, thickly fleshy, exterior minutely puberulent, dull green to

yellowish, swiftly falling at female receptivity to leave a substantial, slightly

oblique, scar; spadix slender cylindrical, sessile, inserted level on peduncle,

3-3.5 x 0.4-0.5 cm, dull yellow-white; stylar region rather well developed,

mostly rhombohexagonal, 1.9-2.4 x c. 2 mm, truncate; stigma punctiform,

c. 0.3 mm diam., prominent in dried material; anthers barely exserted at

anthesis, pollen extruded from between ovaries; infructescence oblong-

226 Gard. Bull. Singapore 51 (1999)

cylindric, 2.5-4 x 1-1.2 cm.

Distribution: Peninsular Malaysia (Kedah, Malacca, Pahang, Perak,

Selangor, Terengganu), Singapore, southern Thailand, Sumatera,

throughout Borneo extending to the Philippines (Palawan) and Sulawesi.

Habitat: Lowland to hill dipterocarp forest on wet to inundated soils, peat

swamp and fresh water swamp forest. 10-200 m altitude.

Notes: 1. A distinctive climber by the combination of the slender, asperous

stems and softly leathery leaves, a combination of characters unknown in

any other Malaysian Rhaphidophora. In dry material the strongly

discolorous leaves, combined with the primary veins prominently darker

than the abaxial leaf surface, are unmistakable.

2. Rhaphidophora lobbii is widespread and often common in Peninsular

Malaysia and that there are not more collections is puzzling. Only one

collection is recorded for Pasoh F.R. (Boyce 1338), particularly odd given

that in Pasoh substantial stands of R. /obbii occur in partially inundated

ground close to the boardwalk leading from the laboratory and guest

accommodation complex.

3. Rhaphidophora lobbii is habitually found in wet to inundated ground,

an unusual habitat for a monsteroid liane.

Other specimens seen: PENINSULAR MALAYSIA: Kedah - Ayer

Tunjang, Furtado s.n. (SING); near Keroh, Nauen s.n. (SING); Malacca -

‘Malacca’, Griffith 5961 (K), Maingay 3252 (K, L); Selandor, Alvins 270

(SING); Negeri Sembilan - Pasoh FR, trail to and environs of Canopy

Walkway, Boyce 1338 (K, KEP); Pahang - Sg Renoris, Taman Negara Fitt

9 (K); Sg Lembing, Hardial 2 (K, SING); Lancang, Bk Rengit, Zainudin

C18 (UKMB); Perak — ‘Perak’ Scortechini 135b (K); Ulu Bubong, Kunstler

10571 (K); Bk Larut, Kunstler 2019 (BM, K), Kunstler 2677 (K); Selangor -

Banting, Telok FR, Kassim & Zainudin s.n. (UKMB); Terengganu -

Kemaman, Ulu Kajang, Corner s.n. (SING). SINGAPORE: ‘Singapore’,

Schwabe s.n. (B); Bk Timah, Ridley s.n. (SING); Chan Chu Kang, Ridley

s.n. (SING); Krangi, Ridley s.n. (SING); Selitar, Ridley s.n. (SING).

9. Rhaphidophora maingayi Hook.f.

Rhaphidophora maingayi Hook.f., Fl. Brit. Ind. 6 (1893) 543; Ridl., Mat.

Rhaphidophora in Peninsular Malaysia and Singapore 227

Fl. Malay Penins. 3 (1907) 43; Engl. & K. Krause in Engl., Pflanzenr. 37

(I1V.23B) (1908) 37; Ridl., Fl. Malay Penins. 5 (1925) 123 — Type: Malaysia,

Malacca, 28 July 1868, Maingay 2972 (K, holo; L iso).

Rhaphidophora apiculata Alderw., Bull. Jard. Bot. Buitenzorg III, 1 (1920)

383, nomi. illeg., non K. Krause (1912, New Guinea), synon. nov. — Based

on: Indonesia, Sumatera, Sibolangit, Borem Bander Ceroe, 15 Feb. 1917,

Lorzing 4733 (BO).

Large, occasionally very large, robust, pachycaul homeophyllous liane to

20 m; seedling stage a non-skototropic shingling shoot; pre-adult plants

occasionally forming small terrestrial colonies of appressed shingling shoots;

adult shoot architecture comprised of greatly elongated, clinging,

physiognomically monopodial, leafy, non-flowering stems and long,

moderately elaborated, free, sympodial, densely leafy, flowering stems later

pendent under their own weight; stems smooth, climbing stems rectangular

in cross section, the angles often winged, the surfaces between slightly

convex, free stems spreading, irregularly four-angled in cross section,

sometime irregularly terete, branching little, growing moderate lengths,

green, later mid-brown, with very sparse to very copious untidy prophyll,

cataphyll and petiolar sheath fibre at the apices of active shoots, internodes

to 13 x 2.5 cm on adherent shoots, shorter and less stout on free shoots,

separated by large oblique leaf scars, older stems woody; flagellate foraging

stems infrequent, often of great length, + rectangular in cross section;

clasping roots densely arising from the nodes and internodes of clinging

stems, densely pubescent; feeding roots rare, adherent, pubescent; leaves

distichous to distichous-spiralled on adherent and free shoots, those distal

on flowering shoots densely so; cataphylls and prophylls membranaceous,

very quickly drying and degrading into netted and tangled fibres; petiole

deeply grooved adaxially, 5—23 x 0.25-0.5 cm, smooth, apical and basal

genicula weakly defined; petiolar sheath very prominent, extending to and

encircling the apical geniculum, briefly ligulate, very swiftly drying and

degrading into netted untidy fibres, eventually falling to leave a continuous

conspicuous scar from the petiole base, around the top of the apical

geniculum and back to the base; /amina entire, elliptic to elliptic-lanceolate

or falcate-oblong, 8-43 x 2.5—15 cm, coriaceous, upper surfaces semi-glossy,

lower surfaces matt, base sub-acute to decurrent, apex sub-acute with a

somewhat prominent apiculate tubule, margins very slightly reflexed in

dried material; mid-rib raised abaxially, slightly sunken adaxially; primary

venation pinnate, slightly raised abaxially and adaxially; interprimaries

parallel to primaries and only slightly less prominent, very slightly raised

abaxially and adaxially; secondary and tertiary venation + obscure in fresh

228 Gard. Bull. Singapore 51 (1999)

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A. flowering shoot x !/3; B. portion of adult, sterile shoot x '/o; C. stem cross section detail x '/s:

D leaf lamina x !/2; E. venation detail x 2; F. inflorescence, spathe fallen x 3/4; G. spadix detail,

post anthesis x 8 All from Boyce 664.

Rhaphidophora in Peninsular Malaysia and Singapore 229

material, visible as a faint reticulum in dried specimens; inflorescence

solitary, subtended by a fully developed foliage leaf and sparse to copious

netted petiolar sheath, prophyll and cataphyll fibre; peduncle compressed-

cylindric, 10-15 x 0.3-0.5 cm; spathe canoe-shaped, stoutly short-beaked,

12-22 x 2.5-6.5 cm, thickly stiff-fleshy, exterior dull green with black

bruising, white with yellow margins internally, swiftly falling at female

receptivity; spadix tapering-cylindrical, + sessile, inserted level on peduncle,

9-10 x 1.5-1.7 cm, cream; sty/ar region, mostly rhombohexagonal, 1.5—2 x

1.7-2.1 mm, truncate; stigma slightly longitudinally elliptic, raised, c. 0.3

mm diam.; anthers barely exserted at anthesis, pollen extruded from between

ovaries; infructescence not seen.

Distribution: Peninsular Malaysia (Kelantan, Malacca, Penang, Selangor),

southern Thailand and Sumatera.

Habitat: Open disturbed forest remnants on steep slopes, on sandstone.

755 m altitude.

Notes: 1. Plants of the R. montana complex with copious prophyll, cataphyll

and petiolar sheath fibre at the apices of active shoots are here referred to

R. maingayi but might represent an undescribed species since the type of

R. maingayi (Maingay 2972) bears only a very few such fibres. However,

while the typical form of these fibre-bearing plants is extremely distinctive,

there exists a range of intermediates with considerably less fibre that

encompass the plant represented by R. maingayi and almost intergrade

with the fibre-free plants of typical R. montana.

2. Rhaphidophora maingayi is virtually indistinguishable from R. montana

(q.v.) but for the presence of very sparse to copious netted prophyll,

cataphyll and petiolar sheath remains at the shoot apices. While this seems

a weak character it does serve to distinguish the species in habitat and,

indeed, R. maingayi and R. montana are sympatric but do not intergrade

in northern Peninsular Malaysia (e.g., Boyce 661 and Boyce 664) and

southern Thailand (Boyce, pers. obs.)

3. Alderwerelt’s R. apiculata, a nom. illeg. antedated by R. apiculata K

Krause (New Guinea), is almost certainly conspecific (and treated here as

such) with R. maingayi but is odd in drying with curiously olive-greenish

yellow leaves.

Other specimens seen: PENINSULAR MALAYSIA: Kelantan - Tanah

Merah, Pergau Dam site, logging road to Sg Long Intake 1, Boyce 664 (K,

230 Gard. Bull. Singapore 51 (1999)

KEP); Malacca - Panchor, Goodenough 1296 (SING); Penang - Pulau

Betong, Curtis 2885 (K, SING); Selangor - Kuala Selangor, Sg Tinggi, Nur

SFN s.n. (SING).

10. Rhaphidophora minor Hook,/.

Rhaphidophora minor Hook.f., Fl. Brit. Ind. 6 (1893) 544; Ridl., Mat. Fl.

Malay Penins. 3 (1907) 41; Engl. & K. Krause in Engl., Pflanzenr. 37

(IV.23B) (1908) 21, Fig. 4; Ridl., Fl. Malay Penins. 5 (1925) 121-122;

Henderson, Malayan Wild Flowers, Monocots, (1954) 238, Fig. 142 A —

Type: Malaysia, Malacca, Griffith 5988 (K, lecto; K, P, isolecto, selected

here). Hooker cited three clearly conspecific syntypes. That chosen is the

most complete specimen with developing and mature inflorescences and

infructescences. The other collections cited by Hooker are Scortechini s.n.

[Perak (CAL)] and Maingay 3041 [Malacca (K)].

Rhaphidophora celebica K. Krause, Notizbl. Bot. Gart. Berlin-Dahlem. 11

(1932) 331 —Type: Indonesia, Sulawesi, northwest of the island, near Toli-

Toh, Jan. 1910, Schlechter 20698 (B, holo).

[Rhaphidophora palawanensis Furtado, nom. nud. in sched. non. Merr.

(1925)]

Small to moderate, slender, leptocaul, homeophyllous liane to 6 m; seedling

not observed; pre-adult plants forming diffuse terrestrial colonies; adult

shoot architecture comprised of greatly elongated, clinging,

physiognomically monopodial, leafy, non-flowering stems and long

moderately elaborated, free, sympodial, densely leafy, flowering stems;

stems smooth, flexuous, climbing stems + terete, occasionally weakly 4-

angled in cross section, free stems somewhat laterally compressed in cross

section, often branching extensively, growing to considerable lengths and

pendent under their own weight with flowering tips upturned, without

prophyll, cataphyll and petiolar sheath fibre, internodes to 13 x 1.2 cm on

adherent and free shoots, flowering shoots with much shorter internodes,

separated by weak straight leaf scars, older stems woody; flagellate foraging

stems absent; clasping roots arising sparsely or singly from the clinging

stems, pubescent; feeding roots solitary from nodes, free, stout, slightly

pubescent; /eaves weakly distichous and sparsely arranged on adherent

and proximal portions of free shoots, moderately densely distichous distally

on flowering shoots; cataphylls and prophylls membranaceous, very quickly

drying and falling; petiole grooved adaxially, 3 — 6 x 0.1 — 0.25 cm, smooth,

231

Rhaphidophora in Peninsular Malaysia and Singapore

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Figure 12. Rhaphidophora minor Hook.f.

A. portion of adult, sterile shoot x '/,; B. flowering shoot x '/,; C. leaf lamina x 7/,; D. venation

detail x 2; E. inflorescence, spathe just beginning to absciss x 1'/,; F spadix detail, post female

receptivity, pre-anthesis x 8; G. spadix detail, early fruiting x 5. A, C— F from Kerr 14700; B,G

from Niyomdham et al. 1062.

232 Gard. Bull. Singapore 51 (1999)

with a slight apical and prominent basal geniculum; petiolar sheath slightly

prominent, extending beyond the apical geniculum by two small ligules,

very swiftly drying and falling in strips to leave a continuous scar from the

petiole base, around the top of the apical geniculum and back to the base;

lamina entire, narrowly falcate-elliptic to falcate-lanceolate or falcate-

oblanceolate, 2.5-16 x 1.2-3 cm, thinly coriaceous, drying pale straw-

coloured, base cuneate to acute or sub-ovate, apex acute with a prominent

tubule; midrib raised abaxially, slightly raised adaxially; primary venation

pinnate, slightly raised on both surfaces prominent (raised)in dried material;

interprimaries sub-parallel to, but much less distinctive than, primaries,

sometimes degrading into weakly reticulate venation, very slightly raised

abaxially; secondary and tertiary venation + invisible in fresh material, barely

visible in dried specimens, reticulate; inflorescence solitary, subtended by a

fully developed foliage leaf and a very quickly falling cataphyll; peduncle

compressed-cylindric, 3—4 x 0.3-0.5 cm; spathe cigar-shaped, stoutly long-

beaked, 3-9 x 1-1.5 cm, thin, dull green to dull yellow, swiftly falling at

female receptivity to leave a large, straight, scar; spadix slender cylindrical,

sessile, inserted level on peduncle, 2.5—7 x 0.5—0.6 cm, dull yellow-white;

stylar region rather well developed, mostly rhombohexagonal, 1.4-2 x c.

2 mm, truncate; stigma punctiform, c. 0.3 mm diam., slightly prominent in

dried material; anthers well-exserted at anthesis; infructescence oblong-

cylindric, 4.5—7 x 1-2.5 cm.

Distribution: Peninsular Malaysia (Johore, Malacca, Pahang, Perak),

Singapore, southern Thailand, Sumatera, throughout Borneo, into the

Philippines (Mindanao, Palawan), and Sulawesi.

Habitat: Lowland forest, along stream margins, sometimes in swampy soil.

10-100 m altitude.

Note: In the fresh state confusion with R. sylvestris is possible, although the

thinner, more prominently veined leaf and long beaked spathe of R. minor

are diagnostic. Dried material of R. minor is notable for the uniformly pale

straw coloured leaves.

Other specimens seen: PENINSULAR MALAYSIA: Johore - Pontian,

Kukup FR, Ahmed KEP 96213 (KEP); Jalan Mersing, Kadim & Noor 223

(K, L, SING); Sungai Mawai, off Sedili Besar river, near Mawai Baharu,

Kota Tinggi, Maxwell 78-274 (L, SINU); Sg Tebrau, Ridley 11631 (SING);

Kampong Kelantan, Kota Tinggi, Shah & Noor 860 (L, K, SING). Sinclair

SFN 10790 (E, K, L, SING); Johore river floodplain, Stone 8692 (KLU);

Malacca - Bk Kayu Luang, Alvins 5843 (SING); Pahang - Ayer Etam

Rhaphidophora in Peninsular Malaysia and Singapore 233

Pekai, Ridley s.n. (SING); Pulau Tioman, Joara Bay, south corner, Burkill

s.n. (SING); Perak - Sg Krian Estate, Spare SFN 33280 (K, SING).

SINGAPORE: Tuas, Ridley 4608 (BM, K); Pulau Sarimbun, Sinclair SFN

5a, (ESL, SING);

11. Rhaphidophora montana (Blume) Schott

Rhaphidophora montana (Blume) Schott, Ann. Mus. Bot. Lugd.-Bat. 1

(1863) 128; Engl. in A. & C. DC., Monogr. Phan. 2 (1879) 240-241; Engl.

& K. Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 40-41; Alderw., Bull.

Jard. Bot. Buitenzorg III, 1 (1920) 386-387 — Calla montana Blume,

Catalogus (1823) 62 — Scindapsus montanus (Blume) Kunth Enum. pl. 3

(1841) 64 (‘montana’) — [Rhaphidophora sylvestris (B1.) Engl. var. montana

(BI.) Nicolson, comb. ined. in sched. var.) — Type: Indonesia, Java, Antjal,

Blume s.n. (L, lecto; L isolecto, selected here). No type was explicitly cited

by Blume at the time of publication, nor in the later printing of the

protologue in Flora (1825) 147. At L there are two Blume sheets agreeing

very well with the protologue. There has been much unnecessary confusion

over Calla montana and the application of the name Anadendrum

montanum Schott (= A. microstachyum (de Vriese & Mig.) Alderw. &

Backer. See Alderwerelt, 1920.

Rhaphidophora angulata (Miq.) Schott, Prodr. Syst. Aroid. (1860) 379 —

Scindapsus angulatus Miq., Fl. Ned. Ind. 3 (1856) 219 — Type: Indonesia,

Sumatera, Doekoe, P.W. Korthals s.n. (U, holo).

Rhaphidophora fallax Schott, Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 128,

synon. nov. — Type: Indonesia, Java, Zippelius s.n. (L, holo; K, L, P, iso).

Rhaphidophora elliptica Ridl., J. Straits Branch Roy. Asiat. Soc. 44 (1905)

186; Engl. & K. Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 39 — 40,

synon. nov. — Type: Malaysia, Sarawak, Bau, July 1903, Ridley s.n. (SING,

holo).

Rhaphidophora burkilliana Ridl., Fl. Mal. Pen. 5 (1925) 121, synon. nov.

— Type: Malaysia, Selangor, Batu Caves, 2 Oct. 1922, Nur SFN 8965

(SING, holo; BM, K, iso).

Large, occasionally very large, robust, pachycaul homeophyllous liane to

15 m; seedling stage a non-skototropic shingling shoot; pre-adult plants

occasionally forming small terrestrial colonies of appressed shingling shoots;

234 Gard. Bull. Singapore 51 (1999)

adult shoot architecture comprised of greatly elongated, clinging,

physiognomically monopodial, leafy, non-flowering stems and long,

moderately elaborated, free, sympodial, densely leafy, flowering stems later

pendent under their own weight; stems smooth, climbing stems rectangular

in cross section, the angles often strongly winged, the surfaces between

more-or-less flat, free stems spreading, four-angled in cross section,

branching little, growing to short or moderate lengths, green, later mid-

brown, without fibre at the apices of active shoots, internodes to 16 x 2.5

cm on adherent shoots, usually shorter and less stout on free shoots,

separated by large oblique leaf scars, older stems woody; flagellate foraging

stems frequent, often of great length, + rectangular in cross section; clasping

roots densely arising from the nodes and internodes of clinging stems,

densely pubescent; feeding roots rare, adherent, pubescent; /eaves distichous-

spiralled on adherent and free shoots, those distal on flowering shoots

densely so; cataphylls and prophylls membranaceous, very quickly drying

and falling; petiole deeply grooved adaxially, 10-23 x 0.25—0.5 cm, smooth,

apical and basal genicula weakly defined; petiolar sheath very prominent,

extending to and encircling the apical geniculum, briefly ligulate, very swiftly

drying and falling to leave a continuous conspicuous scar from the petiole

base, around the top of the apical geniculum and back to the base; /amina

entire, elliptic to elliptic-lanceolate or falcate-oblong, 8-43 x 2.5-15 cm,

coriaceous, upper surfaces glossy, lower surfaces semi-matt, base sub-acute

to decurrent, apex sub-acute with a somewhat prominent apiculate tubule,

margins very slightly reflexed in dried material; mid-rib raised abaxially,

slightly sunken adaxially; primary venation pinnate, slightly raised abaxially

and adaxially; interprimaries parallel to primaries and only slightly less

prominent, very slightly raised abaxially and adaxially; secondary and tertiary

venation + obscure in fresh material, visible as a faint reticulum in dried

specimens; inflorescence solitary, subtended by a fully developed foliage

leaf and one or more cataphylls; peduncle compressed-cylindric, 10-15 x

0.3-0.5 cm; spathe canoe-shaped, stoutly very short-beaked, 12-22 x 2.5-

6.5 cm, thickly stiff-fleshy, dull yellow, paler internally, swiftly falling at

female receptivity; spadix tapering-cylindrical, + sessile, inserted level on

peduncle, 9-20 x 1.5-2 cm, cream; sty/ar region, mostly rhombohexagonal,

1.9-2.2 x 1.9-2.3 mm, truncate; stigma punctiform to slightly longitudinally

elliptic, raised, c. 0.3 mm diam.; anthers barely exserted at anthesis, pollen

extruded from between ovaries; infructescence 9-17 x 1.5—2.5 cm.

Distribution: Peninsular Malaysia (Kedah, Kelantan, Malacca, Pahang,

Selangor), Singapore, southern Thailand, Sumatera, Java, Nusa Tenggara,

throughout Borneo, Sulawesi, and Maluku.

Rhaphidophora in Peninsular Malaysia and Singapore 235

—

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ai (=Oe

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Figure 13. Rhaphidophora montana (Blume) Schott

A. flowering stem x ip B. stem cross-section x sib C. pre-adult terrestrial shoot x '/,; D. leaf

lamina x ae E. venation detail x 4; F. spadix detail, post female receptivity, pre-anthesis x 6.A

& B,D —F from Kanda s.n.; C from Nicolson 943.

236 Gard. Bull. Singapore 51 (1999)

Habitat: Open to closed, primary to disturbed secondary forest, often on

steep slopes on heavy soils over granite, sandstone or limestone. 90-1180

m altitude.

Notes: 1. A taxonomically difficult species in the Hongkongensis Group

that, although easy to recognize in its typical manifestation (shoot apices

without fibre, spreading stiff elliptic leaves and a tapering cylindrical spadix

to 9-16 cm long) exists in a range of forms displaying subtle differences in

leaf shape, inflorescence size, and mostly known from fewer than three,

usually inadequate, collections, all usually collected without shoot apices.

It is the extremes of these forms have been described as distinct species.

2. The use of traditional morphology has proved a most unsatisfactory

method for delimiting R. montana. With an alpha-taxonomy now being

established, there are excellent opportunities for further study of the R.

montana complex using macromolecular data, field observations, and

Statistical techniques to further resolve this group.

Other specimens seen: PENINSULAR MALAYSIA: Kedah - Pulau

Langkawi, Tanjong, Pulau Langgong, Boey 527 (KLU); Kelantan - Tanah

Merah, Pergau Dam site, logging road to Sg Long Intake 1, Boyce 661 (K,

KEP); Malacca, Ms de St Pollias et de La Croix 270 (P); Pahang - Taman

Negara, dam site, Latiff et al. 10 (UKMB); Selangor - Ulu Gombak, Ridley

s.n. (K), Mile 14, Kasim & Rahim 739 (UKMB), c. Mile 15, Croat 53313

(MO), Mile 22 on new highway, Croat 53282 (MO); Bk Takun (limestone),

Harun 32 (UKMB); Ulu Kuang, Provencher 1] (KLU); UKM Campus

FR, Bangi, Zainudin & Rahim s.n. (UKMB). SINGAPORE: Singapore

Botanic Gardens, ‘Garden Jungle’, Sinclair SFN 4831 (E, SING), Croat

53239 (MO) Nicolson 1006 (L, SING, US); Bk Timah, Ridley s.n. (SING),

Croat 53247 (MO); Chan Chu Kang, Goodenough s.n. (SING).

12. Rhaphidophora nicolsonii P.C. Boyce, sp. nov.

Rhaphidophora nicolsonii similis R. tetrasperma differt laminis foliorum

maioribus manifeste magis coriaceis, vagina petiolari circa tres quartae

longitudinid petioli attingente, spadice maiore, regione stylari rotundata

— TYPUS: Malaysia, Pahang, Cameron Highlands, milestone 43, near

Brinchang village, 21 July 1961, Nicolson 1186 (US, holo; B, BH, K, L,

SING, iso).

Large, robust, pachycaul, heterophyllous liane of unknown ultimate height:

Rhaphidophora in Peninsular Malaysia and Singapore 254

seedling stage not observed; pre-adult plants (occasionally?) forming

terrestrial colonies; adult shoot architecture comprised of elongated, clinging,

physiognomically monopodial, densely leafy, flowering stems; stems smooth,

without cataphyll and petiolar sheath fibre, internodes to 6 x 2-3 cm,

separated by prominent oblique leaf scars; flagellate foraging stems not

observed; clasping and feeding roots not observed; leaves weakly spiral-

distichous; cataphylls and prophylls membranaceous, soon drying and falling;

petiole shallowly grooved, upper part + terete, 45—70 x 0.5-0.7 cm, smooth,

apical and basal genicula very prominent; petiolar sheath prominent,

extending '/2 — °*/4 along petiole, soon falling to leave a prominent, slightly

corky scar; /amina sparsely to rarely + entirely pinnatipartite, or pinnatisect,

28-53 x 26-47 cm, broadly ovate to ovate-lanceolate to lanceolate,

coriaceous, base truncate and very weakly cordate, apex acute to acuminate,

individual pinnae up to 4 cm wide, but often leaves with only one division

per side and thus pinnae accounting from up to °*/4 of entire lamina, lamina

drying strongly discolorous, mid-brown adaxially, pale brown abaxially;

midrib very prominently raised abaxially, slightly sunken adaxially; primary

venation pinnate, raised abaxially, slightly impressed adaxially; interprimaries

sub-parallel to primaries and only slightly less prominent, slightly raised

abaxially, slightly impressed adaxially; secondary venation sub-parallel to

interprimaries, slightly raised; tertiary venation reticulate; inflorescence

solitary (always?); peduncle terete, to 19 x 1.5 cm; spathe not observed,

probably falling swiftly; spadix stoutly cylindrical, sessile, inserted

decurrently from 2 cm on peduncle, 14 x 2 cm, green; stylar region well

developed, mostly rhombohexagonal, c. 2 x 2 mm, truncate, margins

deflexed; stigma punctiform, c. 0.5 x 0.4 mm; anthers not observed;

infructescence not observed.

Distribution: Peninsular Malaysia (Pahang). Endemic.

Habitat: Sub to upper montane forest. 1500-2000 m altitude.

Notes: 1. In general aspect, R. nicolsonii resembles R. tetrasperma but 1s

readily distinguished by the larger, considerably more coriaceous leaves,

by the petiolar sheath extending to, at most, three quarters along the

length of petiole and in the larger spadix and rounded stylar region. If the

inflorescence of R. nicolsonii proves to be solitary per flowering event this

will be a further distinguishing character.

2. Outside of Malaysia R. nicolsonii appears to be related to R. pertusa (S

India, Sri Lanka), R. glauca (Roxb.) Schott and R. decursiva (Roxb.) Schott

(trans-Himalaya) and R. grandis Schott (western Himalaya) differing from

Gard. Bull. Singapore 51 (1999)

238

ort 4 iG

nN) a BOT.

oP CR.

Figure 14. Rhaphidophora nicolsonii P.C. Boyce

A. flowering shoot x */,; B. leaf lamina x '/,; C. leaf lamina x '/; D. venation detail x 3; E

inflorescence, spathe fallen x */,; F. spadix detail, post anthesis x 4. All from Nicolson 1186.

Rhaphidophora in Peninsular Malaysia and Singapore 239

these species in degrees of leaf division, venation and texture and in

inflorescence characters.

Other specimens seen: PENINSULAR MALAYSIA: Pahang - G.

Brinchang, Kiew RK4722 (SING); G. Berumban, Wray 1579 (SING).

13. Rhaphidophora puberula Engl.

Rhaphidophora puberula Engl., Bot. Jahrb. Syst. 1 (1881) 180 & in Beccari,

Malesia 1 (1882) 269-270; Engl. & K. Krause in Engl., Pflanzenr. 37

(1V.23B) (1908) 30; — Type: Indonesia, Sumatera, Padang, Ajer Mantjoer,

Aug. 1878, Beccari s.n. (FI, holo; B, iso).

Rhaphidophora scortechinii Hook.f, Fl. Brit. India 6 (1893) 545; Ridl.,

Mat. Fl. Malay Penins. 3 (1907) 42-43; Engl. & K. Krause in Engl.,

Pflanzenr. 37 (IV.23B) (1908) 23; Ridl., Fl. Malay Penins. 5 (1925) 122-

123, synon. nov. — Type: Malaysia, Perak, Scortechini 347 (K, holo; CAL,

iSO).

Rhaphidophora kunstleri Hook.f., Fl. Brit. Ind. 6 (1893) 546; 548; Ridl.,

Mat. Fl. Malay Penins. 3 (1907) 43; Engl. & K. Krause in Engl., Pflanzenr.

37 (IV.23B) (1908) 30; Alderw., Bull. Jard. Bot. Buitenzorg III, 4 (1922)

338; Ridl., Fl. Malay Penins. 5 (1925) 123, synon. nov. — Type: Malaysia,

Perak, Bk Larut, July 1883, Kunstler 4538 (K, lecto; selected here). There

are four conspecific collections labelled “Type’ in Hooker’s handwriting at

Kew. That chosen is in flower, with an expanded but still in situ spathe

with part of the spadix visible and therefore is the most informative. The

other collections are all by Kunstler from Perak: Kunstler 532 Goping (K);

6796 Larut (K) and 8426 Taiping (K, SING).

Rhaphidophora gracilipes Hook.f., Fl. Brit. Ind. 6 (1893) 545; Ridl., Mat.

Fl. Malay Penins. 3 (1907) 41; Engl. & K. Krause in Engl., Pflanzenr. 37

(IV.23B) (1908) 30, Fig. 10; Alderw., Bull. Jard. Bot. Buitenzorg III, 1

(1920) 384; Ridl., Fl. Malay Penins. 5 (1925) 121, synon. nov. — Type:

Malaysia, Perak, June 1886, Kunstler 10271 (K, lecto; BM, SING, isolecto,

selected here). The collection chosen is in flower with an intact spathe and

the spadix visible and thus most useful for determination purposes. The

other, clearly conspecific, syntypes of R. gracilipes are Kunstler 2102 |Perak,

Larut (BO, K)], Kunstler 7940 [Perak (K)] and Scortechini 1103 |Perak

(K)].

240 Gard. Bull. Singapore 51 (1999)

Rhaphidophora megasperma Engl., Bot. Jahrb. Syst. 25 (1881) 8; Engl. &

K. Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 29, Fig. 9; Alderw., Bull.

Jard. Bot. Buitenzorg II, 1 (1920) 383, synon. nov. — Type: Indonesia,

Kalimantan, Tumbang Hiang, 2 Sept. 1881, H. Grabowski 48 (B, holo).

Rhaphidophora batoensis Engl. & K. Krause in Engl., Pflanzenr. 37 (TV.23B)

(1908) 27, synon. nov. — Type: Indonesia, Sumatera, Kepulauan Batu, 25

Sept. 1896, Raap370, (BO, lecto; B, isolecto, selected here). Engler cites

two conspecific syntypes. The collection chosen is in flower (BO) and

mature fruit (B). The other syntype, Raap 325 (immature flower, BO), is

also from Kepulauan Batu.

Rhaphidophora hallieri Alderw., Bull. Jard. Bot. Buitenzorg III, 1 (1920)

385; Alderw., Bull. Jard. Bot. Buitenzorg III, 4 (1922) 196, synon. nov. —

Type: Kalimantan, Hallier 1187 (BO, holo; BO iso).

Rhaphidophora pilosula Alderw., Bull. Jard. Bot. Buitenzorg III, 1 (1920)

386; Alderw., Bull. Jard. Bot. Buitenzorg III, 4 (1922) 196, synon. nov. —

Type: Cult. Bogor Bot. Gard, June 1919, Alderwerelt s.n. (BO, holo).

Distribution: Peninsular Malaysia (Pahang, Perak, Selangor), Sumatera

(including Kepulauan Batu and Kepulauan Mentawi), Nusa Tenggara, and

throughout Borneo. |

Moderate to large, rather robust, semi-pachycaul homeophyllous liane to 5

m; seedling stage not observed; pre-adult plants often forming small

terrestrial colonies; adult shoot architecture comprised of elongated, clinging,

physiognomically monopodial, leafy, non-flowering stems and short, usually

unbranched, free, sympodial, densely leafy, flowering stems; stems smooth,

terete in cross section, with very sparse prophyll, cataphyll and petiolar

sheath fibre, this soon falling, internodes to 13 x 1.5 cm on adherent shoots,

usually shorter and stouter on free shoots, separated by large, straight,

corky leaf scars, older stems woody; flagellate foraging stems absent; clasping

roots sparsely arising from the nodes and internodes of clinging stems,

pubescent; feeding roots rare, adherent, pubescent; /eaves distichous on

adherent and free shoots; cataphylls and prophylls membranous, quickly

drying and degrading into very sparse fibres, these soon falling; petiole

deeply grooved to basally canaliculate, 9-33 x 0.2-3 cm, with the distal

portion and apical geniculum minutely pubescent, especially when newly

expanded, apical and basal geniculum moderately prominent; petiolar sheath

very prominent, extending to just below the apical geniculum, swiftly drying

and degrading into sparse, soon-falling; /amina entire to slightly perforated,

Rhaphidophora in Peninsular Malaysia and Singapore 241

L.GuUrR

Figure 15. Rhaphidophora puberula Engl.

A. flowering shoot x '/,; B. leaf lamina x '/,; C. venation detail x 6; D. inflorescence, spathe

removed; E. spadix detail, post anthesis x 6; spadix detail, early fruiting x 4.A & F from Ridley

14444; B —- E from Burkill 6267.

242 Gard. Bull. Singapore 51 (1999)

perforations round to rhombic, extending c. '/. — '/, of lamina width on

each side of the midrib, ovate to oblong-lanceolate or oblong-elliptic,

oblique to falcate, minutely pubescent abaxially when young, 8-34 x 3.5—

10.5 cm, thinly coriaceous to sub-chartaceous, base unequal, rounded, acute

to slightly decurrent, apex acute to acuminate; midrib prominently raised

and pubescent abaxially, + sunken adaxially; primary venation pinnate,

slightly raised abaxially and adaxially, the leaf appearing slightly quilted,

pubescent in younger leaves, indumentum mostly shed in older leaves;

interprimaries sub-parallel to primaries, very slightly less prominent, slightly

raised abaxially, slightly impressed adaxially; secondary venation tessellate

to weakly reticulate, slightly raised; tertiary venation not visible;

inflorescence solitary, very rarely two together, fragrance of frangipani and

pineapple (fide Hay et al. 9029), subtended by a fully developed foliage

leaf and, if more than one inflorescence, then second preceded by a large

cataphyll; peduncle strongly compressed-cylindric, 4-9 x 0.4—0.6 cm; spathe

canoe-shaped, stoutly long-beaked, 5.5-19 x 1-6.5 cm, stiff-fleshy, dull

green to dull yellow, pale yellow internally, falling (after?) female receptivity

to leave a large, straight scar; spadix very weakly clavate cylindrical,

sometimes slightly curved, sessile, inserted + level on peduncle, 3—11.5 x 1-

1.6 cm, cream; stylar region mostly hexagonal, 1.5—2 x 2.1—2.5 mm, truncate;

stigma punctiform, prominent, 0.5—0.75 x 0.45-0.75 mm; anthers exserted

at anthesis; infructescence stoutly oblong-cylindrical, 4.5-14 x 1.5—2 cm.

Habitat: On trees and rocks in primary and secondary lowland to mid-

elevation dipterocarp forest, often on steep slopes, on granite and limestone.

15-350 m altitude.

Notes: 1. Given the overall uniformity of the species, it is quite extraordinary

that R. puberula should have been redescribed no fewer than seven times

mostly on minor differences of leaf shape.

2. Superficially similar to R. foraminifera, especially in the perforate-leaf

forms, R. puberula can be readily distinguished from R. foraminifera by

the production of inflorescences on free lateral shoots and in having the

active shoot apices without the characteristic black mucilage of R.

foraminifera. Generally, flowering plants of the perforate-leaved forms of

R. puberula have smaller leaves that are noticeably Jess perforated than

those of R. foraminifera.

Other specimens seen: PENINSULAR MALAYSIA: Pahang - 8 miles south

of Kuala Lipis, Burkill & Haniff SFN 17065 (SING, UC); Perak - Batu

Togoh, Wray 2161 (SING); Upper Perak, Wray 3605 (SING); Kuala

Rhaphidophora in Peninsular Malaysia and Singapore 243

Kangsar, logging road up G. Bubu from Manong, Boyce 704 (KEP); Ipoh,

at 4th mile on Gopeng Road, Burkill SFN 6267 (SING); Tambun, near

Ipoh, on way to Tanjung Rambutan, Hay, Anthony & Banka 9029 (NSW,

KEP); Terong, Ridley 14434 (K, SING); Perak, Tong Temple, Chemor, 4

miles north of Ipoh, Sinclair SFN 9852 (E, L, SING); Selangor - Ulu Lui,

Langsat, Hassan & Kasim s.n. (UKMB); Ulu Langat, Bk Jangkang,

Kampong Pansom, Umbai (for Hilliard) KL 1681 (K, KEP, L).

14. Rhaphidophora sylvestris (Blume) Engl.

Rhaphidophora sylvestris (Blume) Engl. in A. & C., DC, Monogr. Phan. 2

(1879) 239 & in Beccari, Malesia 1 (1882) 268; Engl. & K. Krause in Engl.,

Pflanzenr. 37 (IV.23B) (1908) 22-25, Fig. 6 (‘silvestris’); Alderw., Bull.

Jard. Bot. Buitenzorg III, 1 (1920) 383; Ridl., Fl. Malay Penins. 5 (1925)

122 — Calla sylvestris Blume, Catalogus (1823) 62 — Scindapsus sylvestris

(Blume) Kunth, Enum. pl. 3 (1841) 64 — [Scindapsus angustifolius Hassk.,

Flora 25 (2), Beibl. 1 (1842) 12, nom. illeg. — Rhaphidophora angustifolia

(Hassk.) Schott in Bonplandia 5 (1857) 45, nom. illeg. based on the type of

Calla sylvestris Blume] — Type: Indonesia, Java, Blume 178 (L, lecto; 2

sheets; LE, isolecto, selected here). No type was explicitly cited by Blume

at the time of publication, nor in the later printing of the protologue in

Flora (1825) 147. At L there are two sheets collected by Blume under no.

178, and an un-numbered Reinwardt sheet, of appropriate chronology.

Given that the the protologue appears to have been compiled from both

the Blume and Reinwardt collections, these are here regarded as syntypes

and typified accordingly.

Scindapsus lingulatus Hassk., Flora 25(2) Beibl. 1 (1842) 12; Schott, Prodr.

Syst. Aroid. (1860) 378; Engl. in A. & C., DC, Monogr. Phan. 2 (1879) 248,

synon. nov.— Rhaphidophora lingulata (Hassk.) Schott, Bonplandia 5 (1857)

45, synon. nov. — Monstera lingulata (Hassk.) C. Koch ex Ender, Index

Aroid. (1864) 74, synon. nov. — Type: Indonesia, Java, Hasskarl s.n. (not

traced and, as with many Hasskarl names, it is quite possible that a specimen

was never prepared). Neotypification of S. lingulatus will be done when

revising Rhaphidophora for Java.

Scindapsus aruensis Engl., Bull. Soc. Tosc. Ortic. 4 (1879) 270, synon. nov.

— Type: Indonesia, Melaku, Aru Islands, Gabu-lengaw, May 1873, Beccari

s.n. (FI, holo).

Rhaphidophora wrayi Hook.f, Fl. Brit. India 6 (1893) 544; Ridl., Mat. FI.

Malay Penins. 3 (1907) 42; Engl. & K. Krause in Engl., Pflanzenr. 37

244 Gard. Bull. Singapore 51 (1999)

(1V.23B) (1908) 34; Ridl., Fl. Malay Penins. 5 (1925) 122, synon. nov. —

Type: Malaysia, Perak, Larut, Besar, April 1882, Kunstler 2939 (K, lecto;

selected here). Hooker cites several conspecific syntypes. Unfortunately,

the sheet (one of two sheets for the collection) of Wray 662 explicitly cited

by Hooker as ‘Wray mixed with 662’ (the other sheet of the mixed collection

Wray 662 is the holotype of Anadendrum montanum Schott var. longirostre

Hook.f.) is infertile and cannot be considered nomenclaturally useful despite

the eponymous epithet. The collection chosen is the only syntype in which

the spadix size and details can be readily seen and is the most useful for

purposes of identification. The other syntypes are: Scortechini 521 Perak

(K), Kunstler 6956 Perak (BM, K); Wray ‘mixed with 662’ Perak (BM, K).

Rhaphidophora gratissima Becc., Nelle Foreste di Borneo (1902) 604, synon.

nov. — Rhaphidophora sylvestris (Blume) Engl. var. obtusata Engl., Malesia

1 (1883) 268, synon nov. — Type: Malaysia, Sarawak, Kuching, Nov.1865,

Beccari PB952 (FI & FI spirit 423, holo; K, iso).

Rhaphidophora motleyana Engl. & K. Krause in Engler, Pflanzenr. 37

(IV.23B) (1908) 25, synon. nov. — Type: Indonesia, Kalimantan,

Bangarmassin, 1857-1858, Motley 741 (K, holo).

Distribution: Peninsular Malaysia (Negri Sembilan, Pahang, Penang, Perak,

Selangor), Sumatera, Java, Nusa Tenggara, throughout Borneo, and

Maluku.

Medium to large, occasionally very large, moderately robust, leptocaul to

semi-pachycaul homeophyllous liane to 20 m; seedling stage a non-

skototropic shingling shoot; pre-adult plants very seldom forming small

terrestrial colonies of shingling shoots; adult shoot architecture comprised

of greatly elongated, clinging, physiognomically monopodial, leafy, non-

flowering stems and long, moderately elaborated, free, sympodial, densely

leafy, flowering stems later pendent under their own weight; stems smooth,

climbing stems rectangular in cross section, the angles often slightly winged,

the surfaces between slightly concave, free stems rectangular to sub-terete

in cross-section, green, later mid-brown, without prophyll, cataphyll and

petiolar sheath fibre, internodes to 2.5—-5 x 0.5—1 cm on adherent shoots,

usually less stout on free shoots, separated by weakly defined, slightly

oblique leaf scars, older stems woody; flagellate foraging stems frequent,

often of great length, + rectangular in cross section; clasping roots densely

arising from the nodes and internodes of clinging stems, pubescent; feeding

roots very rare, adherent, pubescent; /eaves distichous on adherent and

free shoots, those distal on flowering shoots densely so; cataphylls and

Rhaphidophora in Peninsular Malaysia and Singapore 245

6-077

ISS :

5 an

Out:

2 Oc

oe!

Figure 16. Rhaphidophora sylvestris (Blume) Engl.

A. portion of adult sterile stem x '/,; B. stem cross-section x 1; C. flowering shoot x '/,; D. leaf

lamina x 1; E. venation detail x 4; F. inflorescence, spathe fallen x 1; G. spadix detail, post

anthesis x 4.A & B from Backer 11199; C, F & G from de Wilde & de Wilde-Duyfjes 13830; D

& E from Afriastini 2399.

246 Gard. Bull. Singapore 51 (1999)

prophylls membranous, very quickly drying and falling; petiole deeply

grooved adaxially, 1-8.5 x 0.15—0.3 cm, smooth, apical and basal genicula

weakly defined; petiolar sheath very prominent, extending to and encircling

the apical geniculum, briefly ligulate, very swiftly drying and falling to

leave a continuous conspicuous scar from the petiole base, around the top

of the apical geniculum and back to the base; /amina entire, lanceolate-

elliptic to falcate-lanceolate, slightly to markedly oblique, 4.5—32 x 1.75-8.5

cm, thinly coriaceous, upper surfaces slightly glossy, lower surfaces semi-

matt, base sub-acute to briefly truncate, apex acute to slightly attenuate,

with a prominent apiculate tubule; midrib slightly raised abaxially, slightly

sunken adaxially; primary venation pinnate, slightly raised abaxially and

adaxially; interprimaries parallel to primaries and only slightly less

prominent, very slightly raised abaxially and adaxially; secondary and tertiary

venation + obscure in fresh material, visible as a faint reticulum in dried

specimens; inflorescence solitary, subtended by a fully developed foliage;

peduncle compressed-cylindric, 2—8.5 x 0.15-0.5 cm; spathe cigar-shaped,

stoutly very short-beaked, 4.5—7.5 x 1-3 cm, thinly stiff-fleshy, dull yellow,

paler internally, swiftly falling at female receptivity; spadix weakly clavate-

cylindrical, sessile, inserted + level on peduncle, 3-6 x 1-2.5 cm; stylar

region, mostly rhombohexagonal, 1-2 x 1.5—2 mm, truncate; stigma

punctiform, raised, c. 0.25-0.3 mm diam.; anthers exserted at anthesis;

infructescence 6-8 x 2—2.5 cm.

Habitat: Damp to wet hills to sub-montane forest. 1100-1400 m altitude.

Notes: Rhaphidophora sylvestris is most similar to R. montana and R.

crassifolia. From R. montana it may be distinguished by the narrower leaves,

the lack of shoot tip fibre and by the shorter (3.5-6 cm compared with

9-16 cm in R. montana), clavate-cylindrical spadix. Overall R. crassifolia is

very similar in appearance but may be readily identified by the considerably

thicker leaves, especially notable in dried specimens, and the twisted primary

adherent stem. Additionally, R. crassifolia is a species of lowland habitats.

Other specimens seen: PENINSULAR MALAYSIA: Negeri Sembilan -

Langgang FR, Kiew 2379 (KEP); Pahang - Fraser’s Hill, on Selangor border,

Burkill & Holttum FMS 7859 (BO, K, SING), Burkill & Holttum SFN

7862 (SING); below Fraser’s Hill, Tras Valley, Holttum SFN 11487 (SING),

Richmond, Shah & Noor 671 (K, SING), trail down the slope after RISDA

office, Zainudin, Muzni & Sharif 4634 (UKMB); Cameron Highlands,

Henderson SFN 23271 (BO, SING), Boh Plantations, Nur SFN 32827 (KEP,

MO, SING, UC), Ulu Telom, Jaamat FMS 27284 (KEP); ravine 10 miles

from Berinchang, Bowen I] (KLU); Penang - Government Hill Road,

Rhaphidophora in Peninsular Malaysia and Singapore 247

Burkill SFN 4313 (K, SING); Perak — Maxwell Hill (Bk Larut), Boyce 680

(K, KEP), Burkill & Haniff SFN 12975 (K, SING), (Taiping Hills), Ridley

11418 (BM, K, SING); (Trees Cottage), Ridley 2960 (K); G. Hijau, Stone

14391 (KLU).

15. Rhaphidophora tetrasperma Hook,

Rhaphidophora tetrasperma Hook.f., Fl. Brit. India 6 (1893) 548; Ridl.,

Mat. Fl. Malay Penins. 3 (1907) 44-45; Engl. & K. Krause in Engl.,

Pflanzenr. 37 (IV.23B) (1908) 48; Ridl., Fl. Malay Penins. 5 (1925) 124 —

Type: Malaysia, Perak, Scortechini 169b (K, holo).

Distribution: Peninsular Malaysia (Kelantan, Perak), and southern Thailand.

Small to medium-sized, rather slender, semi-pachycaul, heterophyllous liane

to 5 m; seedling stage a non-skototropic shingling juvenile shoot; pre-adult

plants very rarely forming terrestrial colonies; adult shoot architecture

comprised of elongated, weakly clinging, physiognomically monopodial,

flexuous, moderately leafy, non-flowering stems and weakly adherent or,

more commonly, free lateral flowering stems; stems smooth, without

prophyll, cataphyll and petiolar sheath fibre, internodes to 14 x 1 cm,

separated by prominent straight leaf scars; flagellate foraging stems not

observed; clasping roots sparsely produced from nodes and internodes;

feeding roots stout, produced singly or in pairs from most nodes of free

shoots; leaves weakly spiral-distichous; cataphylls and prophylls

membranous, soon drying and falling; petiole shallowly grooved, 10-34 x

0.2-0.4 cm, smooth, apical and basal genicula slightly prominent; petiolar

sheath prominent, extending to base of apical geniculum, soon falling to

leave a prominent, slightly corky scar; /Jamina sparsely to + entirely deeply

pinnatipartite to nearly pinnatisect, occasionally with large rhombic

perforations adjacent to midrib, 12-42 x 9.5—38 cm, broadly ovate to ovate-

lanceolate, thinly coriaceous, base truncate or very weakly cordate, apex

acute to acuminate, individual pinnae up to 6 cm wide; midrib prominently

raised abaxially, slightly sunken adaxially; primary venation pinnate, raised

abaxially, slightly impressed adaxially; interprimaries diverging from

primaries, much less prominent, slightly raised abaxially, very slightly

impressed adaxially; secondary venation weakly reticulate, very slightly

raised; tertiary venation barely visible; inflorescence few together, subtended

by two prominent cataphylls, these soon falling; peduncle terete, 2—2.5 x

0.3-0.4 cm; spathe canoe-shaped, 3-3.5 x 0.8-1.5 cm, stiffly fleshy, apparently

falling swiftly, white with adherent black cataphyll remnants; spadix

248 Gard. Bull. Singapore 51 (1999)

Sy oS

ex \

a”

——™

Say)

SUT RS

SO }HOY sl

SANUS WN a

SEY \ ==

LOW y ) =

eee

SOU

SRY

Py SNA A\\ Ny

SIM

ecg BY) Z HD \ D

Figure 17. Rhaphidophora tetrasperma Hook.f.

A. pre-adult stem x '/,; B. leaf lamina x '/,; C. shingling shoot x '/,; D. shingling shoot x '/,. All

from Boyce 1224.

Rhaphidophora in Peninsular Malaysia and Singapore 249

cylindrical, sessile, inserted slightly decurrently on peduncle, 3-3.5 x 0.75-1

cm, white; stylar region well developed, mostly rhombohexagonal, c. 2 x 2

mm, truncate, margins deflexed; stigma elliptic, longitudinally orientated,

c. 1 x 0.2 mm; anthers exserted at anthesis; infructescence not observed.

Habitat: Disturbed rather dry to moist or wet forest on sandstone and

granite. 190-760 m altitude.

Notes: As noted under R. nicolsonii above, R. tetrasperma most closely

resembles R. nicolsonii but may be readily distinguished by petiole sheath

and leaf lamina characters.

Other specimens seen: PENINSULAR MALAYSIA: Kelantan - Tanah

Merah, Pergau Dam site, logging road to Sg Long Intake 1, Boyce 652 (K,

KEP); Perak, - Bidor Corner SFN s.n. (SING); Sg Batang Padang, Tapah

Hill Reserve, Furtado SFN 33100 (SING).

Acknowledgements

Thanks are due to Linda Gurr for skilfully executing the illustrations that

accompany this article. I also wish to thank Dr H. RoeBler, Miinchen, for

providing the Latin translation of the diagnoses.

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254 Gard. Bull. Singapore 51 (1999)

Index of Exsiccatae

angustata = 1 maingayi = 9

beccarii = 2 minor = 10

corneri = 3 montana = 11

crassifolia = 4 nicolsonii = 12

falcata = 5 puberula = 13

foraminifera = 6 sylvestris = 14

korthalsii = 7 tetrasperma = 15

lobbii = 8

Ahmed KEP 96213: 10: Alvins 5843: 10: Arnot 33687: 7

Beccari p.b. 952: 14; p.b. 1977, p.b. 2314, p.b. 2714, p.p. 443: 7; Blanc 459: 2;

Blume 178: 14; Boey 527: 11; Bogner 366: 2; Boyce 652: 15; 661: 11; 664: 9;

667, 679: 7; 680: 14; 690: 6; 692: 1; 696: 2; 704: 12; 1064: 1; 1338: 8; Burbidge

s.n. sub. Hort. Veitch no. 215: 7; Burkill FMS 7761: 7; SFN 4313: 14; SFN

6267: 14; Burkill & Haniff SFN 12586: 7; SFN 12975: 14; SFN 17065: 13;

Burkill & Holttum FMS 7859, SFN 7862: 15

Cantley s.n.: 5; Chin 997: 7; 1011: 2; Copeland 1193: 7; Corner SFN 3096: 2;

SFN 30441: 3; SFN 29318: 4; Croat 53239: 11; 53247: 14; 53280: 7; 53282,

53313: 11; Curtis 2885: 9; 3713: 1

Derry sub. Curtis 3713: 1;

Elmer 9464,16061,18057: 7; Engler 5267: 7; 5307: 6

Fitt 9: 8; Furtado SFN 33111, SFN 33128: 2; SFN 33100: 15

Goodenough 1296: 9; Grabowski 48: 13; Griffith 5961: 8; 5988: 10

Hallier 1187: 13; Hardial 2: 8; 13: 2; Hardial & Nor 27, 95: 2; Harun 32: 11;

Hassan 25: 7; Hassan & Latif 140:7; Hassan & Kasim UKMB 09039: 2;

Hay, Anthony & Banka 9029: 13; 9073: 6; Henderson SFN 11426: 7; SFN

22120: 2; SFN 22318, SFN 23271: 14; SFN 23475: 7; Hilliard KL 1158: 2;

Holttum SFN 9842: 2: SFN 11487: 14: SFN 24764: 2: Hosokawa 8334: 7:

Hullett 906, 907: 7

Rhaphidophora in Peninsular Malaysia and Singapore 255

Jaamat FMS 27284: 14

Kadim & Noor 223: 10; Kasim & Rahim 739: 11; Kasim et al. 439: 2; Kiew

652, 1982, 2348: 2; 2379: 14; 4722: 12; 13519: 6; Korthals 206: 1; Kunstler

532; 13,2019: S;.21012 13: 2677: 8: 2874: 22,2939: 14; 3955, 3963: 274538.

6766: 13; 6893: 7; 6956: 14; 7940: 12; 8111: 4; 8426: 13; 10155: 2; 10271: 13;

10571: 8

La Frankie 2185: 4; Lam 711: 7; Latiff et al. 10: 11; Lérzing 4733: 11

Maingay 2213: 7; 2972: 9; 3041: 10; 3252: 8; Maxwell 78-274: 10; Merrill

BoS 11570: 7; Mk. Ar. URMB 04958: 7; Motley 741: 14; Ms de St Pollias et

de La Croix 270: 11

Ngadiman SFN 36774: 4; Nicolson 1006: 11; 1047: 6; 1075: 7; 1138: 2; 1186:

12; Nur SEN 8965: 11; SEN 11079: 7; Nur SEN 32827: 14

Provencher 11: 11

Raap 325, 370: 13; Ridley 2387: 2; 2960: 14; 4608: 10; 7611, 7664: 2; 11418:

14; 11419: 7; 11631: 10; 14369: 2; 14434: 13

Samek UKMB 09947: 2; Schlechter 20698: 10; Scortechini 135b: 8; 169b: 15;

347: 13; 521: 14; 1103: 13; Shah & Noor 671: 14; 860: 10; Shah & Shuker

3414: 2; Sinclair SFN 4831: 11; SEN 5480, SFN 5516: 7; SEN 5833: 10; SEN

9852: 13; SEN 10790: 10; 10832: 2; Soepadmo 809: 7; Spare SFN 33280: 10;

plone Fou O21, 0/356. 426092: 10: 9637: 2711977. 7; 12125, 13824: 2; 14391:

14; 15384: 2

Umbai (for Hilliard) KL 1158: 2, KL 1681: 13; UNESCO Limestone Expd.

PGi, Zz

Wallich 9109: 7; Wray ‘mixed with 662: 14; 2161: 13; 2260: 12; 2457: 1; 3605:

Zai & Kasim UKMB 02914: 7; Zainudin C18: 8; Zainudin, Muzni & Sharif

4634: 14

256 Gard. Bull. Singapore 51 (1999)

Index to Rhaphidophora Species

angustata\..... -csageee $3.11...) ata a eee ooo ee 197

beccarib..... 22 ae S....8. 22. Lge. eee eee eee 202

COTTE... 25b2 BOR AS. ce ik.. SL RR. See ee ee 205

crassifolia ....:-253922t.. 2 i0..d.L LRA Bea ae 208

SNC 5 LI on on sais vn sac cewnaninstctenp een eitaie- ened Ree dee 212

fOTMmINi fer Doe. ke. Sos cates snide ee ee 213

Korthalsti-.. {ook sd. th. ae os a ee eee 217

i) ee <<) | ene ere terrae Wermree cee he to 223

mampoyi)....eoee Mancath...2.220%. 2 oe. 2 eee 226

FINO ohn TE | ee be A ek ae ee 230

LE) 11 c 1) eee S| neers ee me Cae Ay a 235

nicOlsOnits. eee eke. i511. den ae ee 236

puberilaS 2252) ...8..0 lee eS ee eee 239

sylvestris 2.462 RBS. Le ee RAO ee eee 243

(elT SPOT ING 5. AER. on. 1. RR i ee 247

Gardens’ Bulletin Singapore 51 (1999) 257-308.

Floristic Diversity of Bukit Bauk (Terengganu),

Peninsular Malaysia

S.M. TAM

Department of Plant Sciences, University of Cambridge,

Cambridge CB2 3EA, United Kingdom

Abstract

Bukit Bauk is a small, isolated, coastal hill range that is part of the Terengganu Hills

situated at the southern end of the East Range in Peninsular Malaysia. Four vegetation

types were distinguished on Bukit Bauk, namely, lowland dipterocarp forest, peat swamp

forest, hill dipterocarp forest and vegetation of disturbed areas. Its floristic diversity was

investigated leading to a checklist of plants. Its flora comprises at least 638 species (7.7% of

the total flora of the Peninsular Malaysia) belonging to 285 genera and 103 families. The

most speciose families are the Euphorbiaceae (59 species), Dipterocarpaceae (43 species),

Rubiaceae (36 species), Palmae and Guttiferae (each with 31 species). Of these, 91 taxa are

endemic (87 species, | subspecies and | variety), about 3.6% of the total number of endemic

taxa in the Peninsula. This includes 17.6% of the total number of tree species and 8.0% of

endemic tree species. This study confirms that Bukit Bauk Forest Reserve with a relatively

small area of 7,596 ha is a species-rich area with a high degree of endemism and is an

important component of the flora in Peninsular Malaysia that should be conserved by

being totally protected.

Contents

WS ALEC oa OAR eee Rm AME noe eo ee EL See oe ee! 258

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PVs aur. ne. Ponce ose nA cemietn Atbide. el Ada AR. 260

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Fy Che AON MaelIShUNUCE. AhedS tee 45, J hrct ..s5.)hupisresstsodeesseeseverdeesoroass 270

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wo LEAL SGT este RUE ATES Mi aca els i tlle 0 el Pa ae ano OO 274

VEL SuVON ys (SAMS ERI Saddell tel Soler vie Wn a ee 274

eT ae ee ee Oe ee Lt en dia nasi fanspbracersovvers ihe

258 Gard. Bull. Singapore 51 (1999)

Introduction

Bukit Bauk is a small, isolated coastal hill range of granite rock overlain

with deep loamy soil (Anon, 1954) located on the east coast of Terengganu

(4° 42’ N and 103° 25’ E), approximately 11 km south of Kuala Dungun.

Bukit Bauk proper covers an estimated area of 33.75 km’ with the highest

peak attaining 472.4 m altitude. It is part of the Terengganu Hills (Ulu

Kemaman), which are situated at the southern end of the East Range on

the east coast of Peninsular Malaysia. These hills are extremely interesting

botanically because they are known to possess a large number of species

within a relatively small area, of which a large number are endemic. Holttum

(1936, cited by Kiew, 1991a) first drew attention to its flora, which he

described as “extraordinarily rich and distinctly different from that of other

parts of the Peninsula”. Corner (1960) even suggested that this area

contained the richest and, maybe, the oldest component of the flora in the

Malay Peninsula.

In 1996, the Terengganu Hills were recognized as one of the

biodiversity hotspots in Peninsular Malaysia (Kiew, 1996). Its flora was

estimated to comprise approximately 1500 vascular plant species. Exact

figures were then not available as the Terengganu Hills had not been the

subject of any detailed botanical investigation since Corner mounted two

expeditions to Ulu Kemaman in 1932 and 1935 (Kiew, 1996). These forests

still remain undercollected and poorly known botanically.

Literature survey revealed that no botanical collecting had been

carried out on Bukit Bauk and its surrounding forest prior to 1927 (Burkill,

1927). The first detailed botanical investigation on Bukit Bauk was carried

out in 1955 in a one-acre ecological plot in Compartment 8B (Bukit Bauk

Virgin Jungle Reserve) by officers of the Forest Research Institute (Wyatt-

Smith, 1963). Kapur trees (Dryobalanops sumatrensis, Dipterocarpaceae)

were enumerated and subsequently re-measured in 1957 and 1959. This

one-acre plot yielded a comparatively high figure of 123 tree species (gbh

> 12 in) from 31 families.

Putz (1978) reported that the Forest Research Institute Malaysia

(FRIM) maintained the ecological plot in the Virgin Jungle Reserve (VJR);

and that in 1973 a Big Tree Plot had been set up in the lowland forest

situated beside the main road adjacent to Bukit Bauk. He also noted that

University of Malaya had visited the area as a study site.

Examination of herbarium records showed that since then a small

number of collections were made from Bukit Bauk but no complete floristic

checklist nor results of ecological studies concerning Bukit Bauk were ever

published. A systematic and comprehensive floristic study of the forest

Bukit Bauk was therefore compiled from existing information and as well

as field studies to verify its status as a biodiversity hotspot.

Flora of Bukit Bauk 259

Flora of Peninsular Malaysia

Peninsular Malaysia is recognized as one of the world’s biodiversity hotspots

(Aiken and Leigh, 1992). Its geographic position as the southernmost land

limit of the Asian mainland makes it a meeting place for several regional

floras (Wyatt-Smith, 1963). The lowland floristic province of Peninsular

Malaysia is generally recognized to be divided into four regions (Wyatt-

Smith, 1963; and Ashton, 1990) - that of the extreme northwest above the

Kangar-Pattani line (which is similar to that of peninsular Thailand), the

Perak subprovince, the generalised inland flora on udult ultisol soils on the

west coast and Main Range (which is similar to those of Sumatra and

Kalimantan), and the Riau Pocket beginning from the south east of

Peninsular Malaysia and extending as a belt up to 32-48 km wide as far

north as the Kelantan-Terengganu border (its flora is strongly influenced

by elements from Sarawak and the Riau province). Bukit Bauk falls within

this last region.

The tropical rain forests of Peninsular Malaysia harbour a great wealth

of species. Turner (1995) listed 8,239 vascular plant species belonging to

1,656 genera and 237 families, of which 632 species (34 families) are ferns

and fern allies, and 27 species (4 families) are gymnosperms. A checklist of

endemic trees for Peninsular Malaysia compiled by Ng et al. (1990) included

a total of 2,830 tree species in 532 genera and 100 families, of which 746

species representing 26.4% of the total are endemic. Kiew (1988) estimated

there are about 2600 species of herbs in Peninsular Malaysia belonging to

551 genera and 94 families. The level of endemism in herbaceous species is

considerably higher than for trees (Kiew, 1991b) as seen by several genera

with more than 40 species where 87% of Sonerila (Melastomataceae) species

are endemic, 88% for Argostemma (Rubiaceae), 94% for Didymocarpus

(Gesneriaceae), 96% for Begonia (Begoniaceae) and 100% for Didissandra

(Gesneriaceae).

Hill Forests

Mountains run parallel to the axis of Peninsular Malaysia with the Main

Range lying to the west of a central axis and the East Coast Mountains to

the east. The latter consist essentially of three major and several minor

granite coulisses in Terengganu, including Bukit Bauk, which is also formed

of granite material (Anon., 1954). Mountains in the Peninsula are

comparatively young with emplacement of the granite having occurred in

Permian times. However, Burgess (1975) reported that the east coast granite

may be older, dating at least from the Triassic period.

An abrupt change in slope is a conspicuous and consistent

topographical feature of the Peninsula. This zone of change has been called

steep land boundary (Leamy and Panton, 1966, cited by Burgess, 1975)

260 Gard. Bull. Singapore 51 (1999)

and is defined as the line separating land with average slopes less than 20°

from those with average slopes in excess of that figure dividing potential

agricultural land from land suited to protective and productive forest. By

this criterion, approximately 60% of land in Peninsular Malaysia falls within

the lowland zone (Lim, 1991).

Early floristic studies of hill forests in Peninsular Malaysia were

conducted as part of silvicultural research of economic resources. Timber

extraction in hilly terrain became increasingly important post-World War

II (Wyatt-Smith, 1963) due to increased demand and technological changes

in logging, which introducted heavier and more mechanized equipment

that enabled logging operators to log extensively in both lowland forest

and on slope land (Aiken and Leigh, 1992).

Wyatt-Smith (1963) provided detailed and comprehensive forestry

data on the economic hill forests of Peninsular Malaysia, which, apart from

coastal hills, centred mainly between elevations of 150—600 m. The

altitudinal range of hill forest begins from sea level along the coast to

about 1200 m in the Main Range.

Putz (1978) in his survey of all VJRs in Peninsular Malaysia gave

brief descriptions on their ecology, floristics and conservation importance.

Out of a total of 86 VJRs surveyed, 31 areas involved hill forests of which

one was the Bukit Bauk VJR covering 65 acres.

Kiew (1992), in listing the state of knowledge of mountain peak

flora, noted that only 16 floristic surveys were available for individual

peaks, out of which only 5 were post war, and most were based on short

periods of fieldwork.

Bukit Bauk

The only published botanical account of Bukit Bauk is that by Wyatt-

Smith (1963) who described kapur forest. Kapur trees are predominant

and characterize the lowland forest in the Bukit Bauk Forest Reserve

(FR). Kapur produces a medium-heavy hardwood timber. Its distribution

is restricted to two large blocks along the east coast of Peninsular Malaysia,

where it is fast disappearing due to logging activities. There is a smaller

block in Selangor, on the west coast. The 1955 enumeration recorded all

trees greater than 12 in (30 cm) gbh growing on a one-acre (0.4 ha) plot,

which numbered 222 indiduals in 123 species, 81 genera and 31 families

(Wyatt-Smith, 1963). The commonest species was kapur with 24 individuals

followed by Cleistanthus sumatranus (Euphorbiaceae; 8 individuals),

Nauclea officinalis (Rubiaceae; 7 individuals), Knema hookeriana

(Myristicaceae; 6 individuals) and Scaphium macropodum (Sterculiaceae;

5 individuals). The majority of species (82 species) were represented by a

single tree. It was also noted that, compared with species of Shorea, kapur

Flora of Bukit Bauk 261

was well represented in the smaller size classes, an indication of its shade

tolerant characteristics.

Family representation in the VJR Plot recorded the Dipterocarpaceae

as the commonest family for all size classes, except the small size classes

(minimum girth to 12 inches, 30 cm). In this range, the family Euphorbiaceae

was represented by the most genera with the Annonaceae, Guttiferae,

Myristicaceae, Myrtaceae and Rubiaceae also well represented.

In 1957 and 1959, the Bukit Bauk Jungle Plot was remeasured and

recruitment of trees that had reached a minimum 12 in (30 cm) gbh and

individual mortality was recorded. Sixteen new trees were enumerated, of

which six species had not previously been recorded in the 1955 survey.

Wyatt-Smith considered this an indication of the richness of the flora of

Bukit Bauk.

In addition, a full enumeration of all sapling growth greater than 5 ft

(1.5m) tall and less than 12 in (30 cm) gbh was undertaken in 1955 on a

quarter chain wide transect. Subsequently the saplings were re-enumerated

in 1959. Kapur saplings were found to be the commonest in both

enumerations. Results of the Big Tree Plot Study set up by FRIM at Bukit

Bauk (Anon,, 1973) also recorded kapur as the commonest tree.

Putz (1976) commented that Bukit Bauk was interesting ecologically

because it contained at least three distinct forest types: the most prominent

being the kapur forest, the well developed “palm break” above 150 m

altitude dominated by Livistona endauensis with the undergrowth

dominated by its juveniles, while a third was forest common on the hill

ridges made up of seraya (Shorea curtisii), balau (SS. foxworthyi) and

associated species of Calophyllum.

Methodology

Readily available information about species, their habitats and conservation

status is needed before management strategies can be developed to prevent

decline in plant diversity, resources and habitats. A preliminary checklist

was compiled from records in the literature, e.g., from Tree Flora of Malaya

(Whitmore 1972, 1973; Ng 1978, 1989) and from previous collections from

Bukit Bauk based on herbarium records at KEP, obtained from their

accession books and database, and the index catalogue developed by Ruth

Kiew.

To complete and update the checklist, five field trips were made to

Bukit Bauk between September 1997 and May 1998 to collect and identify

plant specimens of all plant groups except bryophytes and climbers (the

latter because of their inaccessibility). Sterile material was collected as

ecological specimens, while fertile material is deposited at KEP.

262 Gard. Bull. Singapore 51 (1999)

Methods employed in collecting specimens are as follows:

1. general survey of existing flora near to a path, road or forest track,

listing of plants occurring from the edge of the road to 20 m into the

forest at 50 m altitude intervals, and

3. circular plots (r = 5 m) at least 20 m from a road/path in visually

undisturbed forest at 35 m altitude intervals.

In the general survey and plant listings, all trees, shrubs and herbs

were recorded and collected where possible. For the circular plots, all

species present were identified and trees with 30 cm gbh and above were

enumerated. These three methods were conducted at three main sites:

1. Microwave Station Road (Compartment 11A, GPS Reading: 4° 42.251’N

and 103° 23.553’E) leads up to the nearest peak where the summit has

been cleared for a microwave station — general survey of plants along

the road from 50 m up to the summit at 350 m a.s.l., plant listing at 50

m altitude intervals, and circular plots;

2. Kampung Trail (GPS Reading: 4° 42.413’N and 103° 25.114’ E) — general

survey of plants on a self cleared trail 0 m a.s.l. to first summit at 210 m

a.s.l., plant listing, and circular plots; and

3. Quarry Trail (Compartment 2) — general survey of plants from 50 m

a.s.l. to 150 m altitude.

The nearest climatological station to Bukit Bauk is at Dungun

Hospital (4° 46’N and 103° 25’E at 3.1 ma.s.l.). For 1995, the mean annual

temperature was 27.5°C with an annual mean relative humidity at 0800

MST of 93% and at 1400 MST of 70%. The total rainfall for 1995 was 2640

mm with the wettest months from September to January with the maximum

in November and December, the North-East monsoon period. There were

short dry periods between February and March and between June and

August with the driest month in June (Malaysian Metereological Service,

1997).

Bukit Bauk FR covers a total area of 7,596 ha and consists of 46

compartments of various sizes (1-10, 11A, 11B, 11C, 12 to 36, 37A, 37B,

38-43). Compartment 8 is classified as Virgin Jungle Reserve (128 ha).

Bukit Bauk FR includes lowland forest that was logged prior to 1960 and

which has recovered to become regenerated forest. Other areas were logged

as recently as 1986 to 1990. Following foresters’ classification, there are

small areas of primary “poor” forest and primary “good” forest on

undulating terrain reaching 300 m a.s.l. where the predominant plant species

include medang (Lauraceae), kedondong (Burseraceae), nyatoh

(Sapotaceae) and kelat (Syzygium sp.) in the “poor” forest and meranti

merah muda (Shorea spp.), kempas (Koompassia malaccensis), medang

(Lauraceae) and kelat (Syzygium spp.) in the “good” forest (Chin et al.,

1997). There is also a small area of peat swamp forest with some parts

Flora of Bukit Bauk 263

logged before 1980, others as recently as 1990. The forest surrounding the

summit area is in Compartment 11A and is classified as regenerated forest.

Results and Discussion

Vegetation Types on Bukit Bauk

Several vegetation types on Bukit Bauk and the adjacent lowland forest

can be recognised based on habitat, altitudinal zonation and floristic

composition. These include lowland dipterocarp forest, peat swamp forest,

hill dipterocarp forest and secondary (disturbed) forest.

1. Lowland Dipterocarp Forest

Lowland dipterocarp forest at about 50-150 m altitude includes vegetation

on flatland, gentle undulating slopes and in valleys. On the inland face of

the hill (Bukit Bauk Microwave Station Road and Kampung Trail), the

forest has three distinct tree layers. The emergent storey (30-45 m tall) is

discontinuous and consists of gregarious trees mainly of kapur Dryobalanops

sumatrensis and other dipterocarps, such as Shorea parvifolia ssp. parvifolia,

S. acuminata, S. foxworthyi, S. macroptera, S. maxwelliana, S. pauciflora,

Dipterocarpus rigidus (which normally grows on coastal hills), Hopea

mengarawan, as well as non-dipterocarps Endospermum diadenum, Sapium

beccatum, Santiria griffithii, Lithocarpus cantleyanus, Cratoxylum formosum,

Calophyllum teysmanii var. teysmanii, Irvingia malayana, Litsea castanea,

Dialium platysepalum, Adenanthera malayana, Parkia speciosa, Artocarpus

elasticus, A. scortechinii, Parartocarpus bracteatus, Streblus taxoides, Bouea

oppositifolia, Dracontomelon dao, Gluta wallichii, Swintonia floribunda,

Atuna racemosa ssp. excelsa, Scorodocarpus borneensis, Nephelium

maingayi, Palaquium hexandrum, Pouteria maingayi, Heritiera javanica,

Scaphium macropodum, Pentace triptera, Aquilaria malaccensis, Parinari

oblongifolia and Syzygium glaucum.

The continuous second layer at about 20-30 m tall includes large-

sized trees of Gymnacranthera forbesii, Myristica cinnamomea, Castanopsis

rhamnifolia, C. schefferana, C. tungurrut, Lithocarpus cantleyanus, L.

leptogyne, L. maingayi, Shorea glauca (a coastal hill species), Dipterocarpus

verrucosus, D. sublamellatus, Hopea myrtifolia, H. sulcata, Vatica

scortechinii, Litsea costalis, Gluta wallichii, G. elegans, Calophyllum

teysmanii var. teysmanii, C. ferrugineum var. oblongifolium, Garcinia

malaccensis, G. parvifolia, G. wallichianum var. wallichianum, Mesua ferrea,

M. grandis var. grandis, M. racemosa, Beilsmiedia palembanica,

Cinnamomum iners, Barringtonia pendula, Adenanthera pavonina,

Cynometra ramiflora, Fordia albiflora, Parkia singularis, Memecylon

264 Gard. Bull. Singapore 51 (1999)

minutiflorum, Madhuca utilis, Elaeocarpus pedunculatus, E. nitidus var.

nitidus, E. stipularis, Artocarpus fulvicortex, A. nitidus, A. integer var.

silvestris, Streblus elongatus, Ficus microcarpa, Pimelodendron griffithianum,

Baccaurea reticulata, B. motleyana, Irvingia malayana, Anisophyllea corneri,

Diospyros clavigera, D. rufa, D. sumatrana, Parishia paucijuga, Buchanania

arborescens, Melanochyla angustifolia, M. fulvinervis, Cyathocalyx

pruniferus, Ilex cymosa, Canariumi littorale, Santiria griffithii, S. rubiginosa

var. rubiginosa, Lophopetalum floribundum, Blumeodendron tokbrai,

Drypetes laevis, Horsfieldia superba, Syzygium chloranthum, S. griffithii,

Strombosia javanica, Xanthophyllum affine, Gynotroches axillaris, Prunus

javanica, Maclurodendron porteri, Xerospermum noronhianum, Gordonia

singaporeana, Pentace strychnoidea, Schoutenia accrescens ssp. accrescens

and var. borneensis, Brackenridgea hookeri and Maranthes corymbosa.

The third and lowest layer comprises smaller trees less than 20 m tall

and is represented by Gironniera hirta, G. parvifolia, Alangium ebenaceum

var. ebenaceum, Sarcotheca laxa var. laxa and var. sericea, Aporusa aurea,

A. lunata, A. nigricans, A. maingayi, Abdulmajidia chaniana, Barringtonia

macrostachya, Diospyros apiculata, D. lanceifolia, D. pauciflora, D. rigida,

Symplocos adenophylla var. adenophylla, S. rubiginosa, Canarium

patentinervium, Agrostistachys longifolia var. leptostachya, Calophyllum

ferrugimeum var. ferrugineum, C. gracillimum, Garcinia eugeniifolia, G.

nervosa, G. malaccensis, G. rostrata, G. scortechinii, G. subhorizontale,

Mesua lepidota var. lepidota, Cinnamomum mollissimum, Cryptocarya

griffithiana, Litsea magnifica, Strombosia ceylanica, Knema malayana, K.

furfuracea, K. patentinervia, K. scortechinii, Baccaurea motleyana, B.

parviflora, B. racemosa, Croton laevifolius, Xylopia ferruginea var.

ferruginea, Blumeodendron tokbrai, Gynotroches axillaris, Vitex gamosepala,

Polyalthia macrophylla var. macrophylla, Xylopia caudata, X. ferruginea

var. ferruginea and var. oxyantha, X. malayana, Ilex macrophylla, Euonymus

Javanicus, Elaeocarpus palembanicus, E. petiolatus, Antidesma coriaceum,

Cleistanthus sumatranus, Drypetes longifolia, D. pendula, Glochidion

wallichianum, Neoscortechinia kingii, Paracroton pendulus, Trigonostemon

laevigatus, Ryparosa hullettii, Leea indica, Archidendron bubalinum, A.

ellipticum, Saraca declinata, S. cauliflora, Pternandra coerulescens,

Horsfieldia sucosa, Maesa ramentacea, Syzygium attenuatum ssp. attenuatum,

S. gratum, S. sp. 57, S. sp. 59, Tristaniopsis merguensis, Strombosia ceylanica,

Galearia fulva, Paropsia vareciformis, Aidia densiflora, Gardenia tubifera

var. tubifera, Porterandia anisophylla, Timonius flavescens, Guioa

pleuropteris, Lepisanthes tetraphylla, Leptonychia caudata, Rinorea

anguifera, Symplocos rubiginosa var. rubiginosa and the palms Oncosperma

horridum, Arenga hookeriana and A. westerhoutii,

Flora of Bukit Bauk 265

Beneath is a sparse understorey of treelets, shrubs, and tree saplings,

which is dominated by populations of the small to medium sized palms,

Licuala ahlidurii and L. terengganuensis together with smaller populations

of L. khoonmengii and L. glabra. Other palms include Arenga seedlings

spp., Plectocomia elongata, Calamus spp. and Daemonorops monticola.

Treelets include Anisophyllea disticha, Pentace strychnoidea, Thottea sp.,

Ixora congesta, Ficus parietalis, F. deltoidea var. deltoidea, Eurycoma

longifolia, Lasianthus densifolius, Phaeanthus optalamicus; shrubs such as

Chroesthes longifolia, Dracaena maingayi, Polyalthia bullata, Chloranthus

erectus, Kibara coriacea, Ficus deltoidea var. deltoidea, Glycosmis

chlorosperma var. chlorosperm and Sterculia coccinea, with saplings of

Mesua grandis, M. ferrea, Ardisia mystica, A. perakensis, Payena sp. A,

Durio wyatt-smithii, Ilex cymosa, Mallotus paniculatus, Calophyllum sp.,

Hopea sulcata and Shorea macroptera. Species of shrubs and herbs are

poorly represented with the shrubs Triuphetta grandidens and Premna

serratifolia, Pandanus sp. and the herbs Hedyotis philippinensis, Mapania

cuspidata var. cuspidata, Scaphochlamys rubromaculata, Molineria latifolia,

Alpinia javanica, Amischotolype griffith, Chroesthes longifolia and climbers,

Claderia viridiflora, Rhapidophora lobbii, Grenacheria amentacea, and

Uncaria cordata var. cordata..

Valley bottoms are richer in treelets, shrubs and herbaceous plants.

It is here that populations of four Henckelia species, H. puncticulata, H

miniata, H. rugosa, H. heterophylla are found. Other herbs include Globba

unifolia var. sessiflora, Sauvagesia serrata, Hanguana malayana, Zingiber

spp., Alpinia javanica and Alpinia sp.. The shrub Tarenna fragrans and

trees, such as Cynometra ramiflora and Fordia albiflora, were also found in

this habitat.

At about 150 m a.s.l., where the slopes become steeper the emergent

layer becomes less well defined and is dominated by species, such as Alstonia

angustiloba, Cryptocarya rugulosa, Shorea curtisii (a large tree usually

growing on hill ridges), $. maxwelliana, Hopea mengarawan and mature

individuals of the fan palm Livistona endauensis. The second canopy layer

is more continuous with species, such as Durio wyatt-smithii, Litsea

magnifica, Cynometra malaccensis, Memecylon excelsum, M. megacarpum,

Artocarpus lowii, A. lanceifolius, Knema hookeriana, Myristica cinnamomea,

Xanthophyllum affine, Sandoricum koetjape, Shorea multiflora, S.

teysmanniana, S. falcifera (a coastal hill species), Gluta wallichii (very

common on Bukit Bauk), Lithocarpus wallichianus and Palaquium

maingayl.

The third tree layer is made up of Rhodamnia cinerea, Garcinia

pyrifera, Actinodaphne pruinosa, Diospyros scortechinii, Castanopsis lucida,

Lithocarpus maingayi, Abdulmajidia chaniana (very common on Bukit

266 Gard. Bull. Singapore 51 (1999)

Bauk), Mesua grandis var. cordata, Archidendron bubalinum, Acmena

acuminatissima, Lepisanthes fruticosa and Actinodaphne pruinosa. The shrub

layer is dominated by the clustering palm Licuala ahlidurii, Prismatomeris

glabra and Urophyllum glabrum; treelets such as Eurycoma longifolia,

Lasianthus densifolius and saplings of Arenga, Diospyros rufa, Garcinia

nervosa, Hopea myrtifolia, Shorea sp. and Vatica scortechinii. The ground

layer is sparse with clusters of Pandanus sp., Dracaena conferta, Alpinia

javanica, Scaphochlamys rubromaculata, the climber Piper

porphyrophyllum, the palms Daemonorops monticola, Calamus sp. and

seedlings of Livistona endauensis and the fern Lygodium circinnatum.

Forest on the eastern side of Bukit Bauk directly facing the sea

grows on sandy and rocky (granitic) soil. At the base of the hill, lowland

forest has been cleared for quarrying activities. At about 50-150 m a.s.l.,

where the terrain is steep, the emergent layer is rather discontinuous and

is characterized by the presence of Shorea glauca (a coastal hill species),

Palaquium hexandrum, Swintonia floribunda and Sindora echinocalyx (a

hill forest species). Individuals of kapur and Livistona endauensis are

conspicuously absent.

The second canopy layer is dense and continuous (despite the

disturbance at the base of the hill) and consists of trees, such as

Erythroxylum cuneatum (a species normally found near the sea), Sapium

beccatum, Scaphium macropodum, Lophopetalum floribundum, Artocarpus

lanceifolius, A. elasticus, Xerospermum noronhianum, Mesua racemosa,

Litchi chinensis (a hill forest species) and the palm Arenga westerhoutii.

The third storey species are Barringtonia macrostachya, Cleistanthus

sumatranus, Paracroton pendulus, Baccaurea parviflora, Garcinia

scortechinii, Buchanania sessifolia, Knema furfuracea and Streblus taxoides.

Saplings, shrubs and seedlings occur in small numbers, namely Eurycoma

longifolia, Saraca declinata and Donax grandis. The vegetation on this part

of the hill reflects past disturbance because species usually found growing

in disturbed or open sites are encountered here, such as Vitex pubescens,

Callerya artopurpurea, Commersonia bartramia and Melastoma

malabathricum.

ii. Peat Swamp Forest

In Peninsular Malaysia, peat swamp forests occur just behind the coastline

(Wyatt-Smith, 1963). On Bukit Bauk, peat swamp forest is separated by

the main road from the west of the hill. The peat swamp forest here has

three canopy layers. The emergent layer is represented by scattered

individuals of Shorea acuminata, S. macroptera, Dryobalanops sumatrensis,

Mangifera quadrifida, Dialium platysepalum, Sarcotheca griffithii, Heritiera

javanica, H. simplicifolia, Scaphium macropodum and Intsia palembanica.

Flora of Bukit Bauk 267

The second canopy layer is fairly continuous with trees of Canarium

rufum, Dacroydes costata, D. rostrata, Santiria laevigata, S. rubiginosa, S.

tomentosa, Durio wyatt-smithii, D. griffithiit, Heritiera javanica, H.

simplicifolia, Euonymus javanicus, Gynotroches axillaris, Atuna racemosa

ssp. excelsa, Parinari elmeri, Dillenia reticulata var. psilocarpa, Dipterocarpus

costulatus, D. lowii, D. verrucosus, Shorea multiflora, S. singkawang ssp.

singkawang, Vatica pauciflora, Aquilaria malaccensis, Gironniera subequalis,

Castanopsis lucida, C. megacarpa, C. schefferana, Lithocarpus lucidus, Litsea

costalis, Actinodaphne macrophylla, Xerospermum noronhianum, Madhuca

utilis, Brackenridgea hookeri, Xanthophyllum affine, Ochanostachys

amentacea, Sarcotheca griffithii, Melanochyla angustifolia, Elaeocarpus

nitidus var. nitidus, Pimelodendron griffithianum, Garcinia griffithii, G.

parvifolia, Mesua ferrea, M. racemosa, Actinodaphne macrophylla,

Memecylon minutiflorum and Anisophyllea corneri.

The third layer is dense, consisting of many species of smaller trees,

such as Tabernaemontana corymbosa, Monocarpia marginalis, Polyalthia

lateriflora, Gironniera parvifolia, Pternandra coerulescens, Memecylon

megacarpum, Archidendron bubalinum, Barringtonia scortechinii,

Cryptocarya griffithiana, Cinnamomum mollissimum, Stemonurus

malaccensis, Ryparosa hullettii, Blumeodendron kurzii, Kopsia singapurensis,

Pseuduvaria macrophylla, Xylopia caudata, X. ferruginea var. oxyantha,

Diospyros adenophora, D. apiculata, D. argentea, Microcos fibrocarpa, M.

lanceolata, Agrostistachys longifolia var. leptostachya, Rinorea anguifera,

Aporusa aurea, A. bracteosa, A. falcifera, A. maingayi, Knema curtisii, K.

furfuracea, K. hookeriana, K. patentinervia, Baccaurea brevipes, B. minor,

B. parviflora, Lepisanthes tetraphylla, Blumeodendron kurzii (a swamp forest

species), Nephelium costatum, N. cuspidatum var. ophiodes, Ryparosa

hullettii (a swamp forest species), Mallotus griffithianus, Quassia indica (a

tidal swamp species), Stemonurus malaccensis, Leptonychia caudata,

Adenanthera clypearia, Ixora lobbii, Urophyllum glabrum and Timonius

flavescens (a peat soil species).

The shrub layer is dominated by clumps of four Licuala species, L.

ahlidurii, L. ferruginea, L. terengganuensis and L. spinosa, which grow on

dry land on the fringe of swamp land, as well as Tabernaemontana

corymbosa, Archidendron clypearia var. clypearia, Ixora lobbii and Sterculia

coccinea. The ground flora is sparse but Orchidantha ?fimbriata is common.

ii. Hill Dipterocarp Forest

Hill forest on Bukit Bauk begins above 150 m a.s.l. on much steeper slopes

(more than 45°), which ascend to ridges before reaching the summit area,

which is an exposed plateau. The emergent layer is dominated by large

tree species, such as Shorea curtisii (particularly common), S. foxworthyi,

268 Gard. Bull. Singapore 51 (1999)

S. maxwelliana, Anisoptera curtisii, Dryobalanops sumatrensis (which is

not as common as it is in lowland forest), Sindora echinocalyx, Gluta

wallichii, Swintonia floribunda, Alstonia angustiloba, Sapium beccatum,

Litsea castanea, and mature individuals of the fan palm Livistona endauensis.

The second canopy layer is discontinuous with small gaps where

natural landslides have occurred. This layer is rich in species including

Dacroydes rostrata, Scaphium macropodum, Castanopsis rhamnifolia,

Elaeocarpus nitidus, E. palembanicus, E. pedunculatus, Gluta wallichii,

Baccaurea maingayi, B. motleyana, Diospyros rufa, D. sumatrana,

Cleistanthus glaucus, Neoscortechinia nicobarica, Callerya artopurpurea,

Cratoxylum formosum, Shorea bracteolata and S. falcifera (the last two

both coastal hill species), S. multiflora, Dipterocarpus rigidus, Pentace

floribunda, Lithocarpus cantleyanus, L. rassa, Garcinia wallichianum var.

wallichianum, Parishia paucijuga (frequently encountered), Melanochyla

fulvinervis, Parinari elmeri, Litsea tomentosa, Fagrea elliptica, Memecylon

minutiflorum, Syzygium chloranthum, S. claviflorum var. claviflorum, S.

griffithii, Xanthophyllum affine, X. griffithii ssp. erectum, Litchi chinensis,

Palaquium maingayi, Pouteria maingayi, Pentace floribunda, P. strychnoidea,

Dillenia grandiflora, Horsfieldia ridleyana and Schoutenia accreascens ssp.

borneensis.

The third storey is denser with species, such as Gironniera hirta, G.

parvifolia, Aporusa maingayi, A. frutescens, A. nigricans, A. prainiana,

Polyalthia macrophylla var. macrophylla, Euonymus javanicus, Diospyros

apiculata, D. pauciflora, D. scortechinii, Agrostistachys longifolia var.

leptostachya, Antidesma coriaceum, Mallotus paniculatus, Castanopsis lucida,

Lithocarpus maingayi, Calophyllum gracillimum, Ardisia mystica,

Pternandra echinata, Cinnamomum mollissimum, Tristaniopsis merguensis,

Galearia fulva, Gardenia tubifera var. tubifera, Porterandia anisophylla,

Symplocos adenophylla, S. rubiginosa, Vitex longisepala, Rinorea anguifera,

Cleistanthus glaucus, Mesua lepidota, Calophyllum ferrugineum var.

ferrugineum, C. subhorizontale, Aglaia leucophylla, Santiria tomentosa,

Rhodamnia cinerea (a coastal species), Knema scortechinii, Memecylon

megacarpum, Garcinia eugeniifolia, G. pyrifera, and the palms Arenga

westerhoutii and Oncosperma horridum.

The shrub layer includes Aidia densiflora, Ardisia rosea var. 2, Maesa

ramentacea, Palaquium maingayl, Ixora pendula var. pendula, I. umbellata,

Lasianthus densifolius, Chloranthus erectus, Ficus deltoidea var. deltoidea,

Aporusa frutescens, Eurycoma longifolia, Smilax calophylla, saplings of

trees such as Drypetes longifolia and Paracroton pendulus, and many

juveniles of Livistona endauensis. At about 170 m a.s.l. on the Kampung

Trail the undergrowth layer is completely dominated by juveniles of the

fan palm Livistona endauensis (seedlings to individuals with short trunks).

Flora of Bukit Bauk 269

Litter on the forest floor is thick consisting of some broken branches and

leaves but mostly dried fronds of Livistona endauensis. Beneath the topsoil

is damp and loose. The ground layer is denser than in the lowland forest

with more and larger clusters of Pandanus spp., Licuala terengganuensis

(less dense than in lowland forest and decreasing as altitude increases),

juveniles of the palms Arenga sp., Calamus longispathus, Calamus sp. and

the herbs Molineria latifolia and Hedyotis philippinensis.

At about 200 m a.s.l., steep hill forest gives way to hill ridges with

exposed areas of dry, thin, nutrient-poor soil, usually of a sandy or rocky

nature. The emergent layer is very incomplete with occasional individual

large trees of Anisoptera curtisii, Shorea bracteolata, S. curtisii, S. glauca, S.

maxwelliana, Anisophylla griffithii, Alstonia angustiloba and crowns of

Livistona endauensis, the last common in this habitat. The second layer is

thinner and includes smaller tree species such as Cleistanthus glaucus,

Adenanthera malayana, Calophyllum wallichianum var. wallichianum,

Elaeocarpus pedunculatus, Shorea falcifera (a coastal hill species), Fagraea

wallichiana (a hill ridge species), Horsfieldia ridleyana, Pentace grandiflora

with a large number of Shorea curtisii adults and saplings.

The third storey is denser with species such as Santiria rubiginosa,

Castanopsis schefferana, Parishia paucijuga, Breynia coronata, Abdulmajidia

chaniana, Aporusa prainiana, Syzygium lineatum, Gordonia maingayi,

Diospyros confertiflora, Tristania merguensis, Macaranga triloba, Fagraea

fragrans and Greenea corymbosa (a coastal hill ridge species). Tree saplings

and shrubs are poorly represented on these hill ridges. In exposed areas,

e.g., forest edge, climbers, such as Uncaria cordata, Smilax setosa, Nepenthes

gracilis and N. ampullaria, as well as the orchid Bromheadia finlaysoniana

eTOW.

The first summit on the southern front of Bukit Bauk at 350 m a.s.l.

has been cleared to build a microwave station, telecommunication station

and a guardhouse. It is dry and exposed with rocky or sandy soil and is

mostly occupied by smaller-sized trees, shrubs and saplings. Canopy trees

on the fringe of the clearing include Blumeodendron tokbrai, Shorea

maxwelliana, and Parishia paucijuga. Smaller-sized trees include Rhodamnia

cinerea (a coastal species), Macaranga heynei, M. hypoleuca and M. triloba,

Fagraea fragrans, Canarium patentinervium, Vitex pubescens, Archidendron

ellipticum, Gardenia tubiflora var. tubiflora, Syzygium chloranthum, Garcinia

parvifolia, Gordonia maingayi, Castanopsis schefferiana and Cleistanthus

glaucus.

The shrub layer consists mostly of Shorea saplings, Rubus moluccanus

and Livistona endauensis juveniles and seedlings with the ground layer

colonized by Dicranopteris linearis, Sticherus truncatus vat. truncatus, Ardisia

rosea var. 2, Elephantopus mollis, Uncaria cordata, Stachytarpheta indica

270 Gard. Bull. Singapore 51 (1999)

and Bromheadia finlaysoniana.

iv. Vegetation of Disturbed Areas

The disturbed areas are mainly by the sides of the road leading up to the

Microwave Station and at the telecommunication towers. Forest edge

species include Mallotus philippensis, Melastoma malabathricum, Macaranga

heynei, M. triloba, Ficus fulva, F. aurata and F. deltoidea (an epiphyte),

Rhodamnia cinerea, Diospyros scortechinii, Commersonia bartramia, Vitex

pinnata, Croton laevifolius, Themeda villosa, Molineria latifolia, Fagraea

racemosa, Dissochaeta gracilis, Rubus moluccanus, var. moluccanus, Myrica

esculenta, Syzygium lineatum, Rhodomyrtus tomentosa, Sarcotheca laxa var.

laxa and var. sericea, Morinda elliptica, Porterandia anisophyllea, Nepenthes

gracillis and N. ampullaria, and the ferns Dicranopteris linearis var. linearis,

Sticherus truncatus var. truncatus, Taenitis blechnoides, Blechnum orientale

and Lygodium longifolium. On hill ridges on shaded, rocky roadside banks,

populations of Henckelia punticulata grow together with Selaginella.

Floristic Diversity of Bukit Bauk

The preliminary checklist included a total of 322 species from 65 families;

field work increased the total number of species to 638 in 285 genera and

103 families (Table 1). A total of 115 plant species from 37 families recorded

in the preliminary checklist were re-collected in this study. The complete

floristic checklist is given in Appendix 1. In general, the forests of Bukit

Bauk are poor in ferns and herbaceous plants.

Table 1. Floristic Composition of Bukit Bauk based on number of taxa.

Angiosperms | |

Dicotyledons Monocotyledons }Gymnosperms | Pteridophytes Total

| Family | 76 14 2 11 103

_ Genus | 238 32 2 | 13 285

_ Species 567 52 | 2 17 638

Apart from palms, the most speciose families are predominantly trees

(Table 2).

Flora of Bukit Bauk 271

Table 2. Ten Most Speciose Families at Bukit Bauk

Family No. Genera No. Species No. Endemics

Euphorbiaceae 24 a7 5

Dipterocarpaceae i 43 >

Rubiaceae pHs 36 2

Palmae 13 a1 | 10

| Guttiferae | 4 31 3

| Myrtaceae 6 26 1

Annonaceae 10 23 6

Anacardiaceae 10 20 | s)

Leguminosae 12 | 19 2

Lauraceae 8 18 e)

The Euphorbiaceae is the most speciose family with 59 species (9.2%

of the species recorded from Bukit Bauk). Wyatt-Smith (1963) had already

found this to be the case in sample plots at Bukit Bauk (one-acre plot),

Mata Ayer (Perlis) and Gunung Raya (Langkawi Island) and similar results

were reported for lowland forest at Pasoh FR, Negeri Sembilan

(Kochummen et al., 1990) and Sungai Menyala, Negeri Sembilan, and Bukit

Lagong, Selangor (Manokaran and Kochummen, 1990). In terms of number

of species, the Euphorbiaceae is the second largest family of dicotyledons

in Peninsular Malaysia with 70 genera and 364 species of trees, shrubs,

herbs and climbers (Turner, 1995).

The second most speciose family on Bukit Bauk is the

Dipterocarpaceae with 43 (67%) species, which represent more than a

quarter of the Peninsula’s 156 species. Wyatt-Smith (1963) reported the

Dipterocarpaceae as the third commonest family in the one-acre plot on

Bukit Bauk at the minimum size class gbh = 30 cm but ranked first both in

species and individuals at the minimum size class gbh = 60 cm. It is well

represented by big trees of the upper canopy and emergent layer. In terms

of total tree densities and total basal area, the Dipterocarpaceae was also

the most common family in Bukit Lagong and Sungai Menyala (Manokaran

and Kochummen, 1990) but in Pasoh FR the Dipterocarpaceae ranked

tenth in species diversity (Kochummen et al, 1990).

The Rubiaceae on Bukit Bauk is the third most speciose family with

36 species, whereas at Pasoh FR it ranked fourth with 46 species

(Kochummen et al., 1990), and in the Bukit Bauk one-acre plot for minimum

size class gbh = 30 cm (Wyatt-Smith, 1963) it ranked sixth. The Rubiaceae

is the most species rich dicot family in Peninsular Malaysia (Turner, 1995)

272 Gard. Bull. Singapore 51 (1999)

but it is better represented by shrubs and herbs (which are sparse on Bukit

Bauk) than by trees.

Palms are not usually included in forest enumerations. On Bukit

Bauk they rank fourth with 31 species and include the highest number of

endemic species of any family (Table 2). They form a conspicuous element

of the flora on Bukit Bauk and the Terengganu Hills as a whole, which

have been noted to be particularly rich in palm species (Kiew, 1996). Bukit

Bauk FR supports nine species of Licuala, four endemic to Terengganu

(Saw, 1997) and a large population of Livistona endauensis. This latter

palm is only common on Bukit Bauk, the adjacent Terengganu hills and at

Endau Rompin, Johore (Dransfield and Wong, 1987).

The fifth most speciose family, the Guttiferae with 31 species

representing more than a quarter of the Peninsula’s 120 species, owes its

diversity to the large number of Calophyllum and Garcinia species, each

with 11 species. Putz (1976) considered that on ridges at Bukit Bauk

Calophyllum formed a distinct type of hill forest growing in association

with Shorea curtisii and S. foxworthyi. Guttiferae ranked fourth in the

Bukit Bauk one-acre plot (Wyatt-Smith, 1963).

The other five families are each represented by less than thirty species.

Notable among them is the Anacardiaceae, which on Bukit Bauk is

represented by more than a quarter of the Peninsula’s 77 species.

Endemism

The number of endemic taxa occurring on Bukit Bauk is 98 (93 species, 1

subspecies and 4 varieties) representing 14.6% of the total number of

species (Table 3). Endemism is higher among monocots (26%), mainly

due to the high level of endemicity among palms. Thirteen species are

endemic to Terengganu and four are narrow endemics known only from

Bukit Bauk/Bukit Bauk FR (Table 4).

Table 3. Species Endemism at Bukit Bauk

| Category Dicots | Monocots Total

_ Terengganu only i) 6 | 13

+ -

_ Terengganu and one 15 3 | 18

other state |

Endemic to Si 5 62

Penininsular

Malaysia

Total No. Endemic 79 14 93 (14.6%)

Total No. Species 567 53 | 639

Flora of Bukit Bauk DIS

Table 4. Species endemic to Terengganu or Bukit Bauk/Bukit Bauk FR

Terengganu only Bukit Bauk only

Henckelia miniata

Kochummenia parviflora Pseudovaria cerina

Pentace grandiflora Dacroydes breviracemosa

Licuala bayana Gonocaryum sp. A

Licuala fractiflexa Licuala ahlidurii

Licuala khoonmengii

Licuala terengganuensis

Scaphochalmys rubromaculata

Comparison of species on Bukit Bauk with the total flora of

Peninsular Malaysia shows that Bukit Bauk is home to at least 7.8% of the

Peninsula’s species and to 16.5% of the tree species with a very high

representation of families (43.9% and 65%) and genera (18.3% and 36.8%)

for all and for tree species, respectively (Table 5). In terms of endemism,

Bukit Bauk harbours at least 3.7% of the total number of endemic taxa

and 8.0% of the total number of endemic tree species in Peninsular

Malaysia. These figures are high considering the small area (33.75 km’)

that Bukit Bauk covers.

Table 5. Comparison of Floristic Data from Bukit Bauk and Peninsular

Malaysia

Peninsular Malaysia Bukit Bauk

Total ‘Trees Total ‘Trees

(Turner, 1995) (Ng et al., 1990)

No. Families (% ) 22th 100 103.(435)4 :65.,.(65)

No. Genera (%) 1565 532 285 (18.2) 196 (36.8)

No. Species (%) 8239 2830 638 (7.7) 467 (16.5)

No. Endemics (%) 2547 746 (26.4) O38. HBT) rev: 60' nih 8)

These results confirm that the flora of Bukit Bauk is high in species

diversity and endemism, particularly when its small size is taken into

consideration, and it certainly deserves to be recognized as a biodiversity

hotspot in Peninsular Malaysia, being especially rich in the

Dipterocarpaceae (represented by 27% of the Peninsula’s species), the

Anacardiaceae and Guttiferae (each by 26%) and the Palmae (18%). In

addition, as Putz (1976) has already noted, the presence of the kapur and

274 Gard. Bull. Singapore 51 (1999)

fan-palm forests also contribute to Bukit Bauk’s conservation

importance.These all support its status as a biodiverse area of conservation

value for the flora of Peninsular Malaysia (Kiew, 1996). Detailed

conservation implications will be discussed in a separate paper.

Conclusions

Bukit Bauk is a small, isolated coastal hill with a high degree of floristic

diversity being not only species rich (comprising at least 638 plant species

belonging to 285 genera and 103 families) but also in having a high level of

endemism (89 species) with four species being known only from this hill.

Within the small area (7,596 ha), several forest types are found — the

lowland mixed dipterocarp forest with kapur as the dominant species, peat

swamp forest, hill forest where the fan palm Livistona endauensis is a

conspicuous feature, and secondary forest.

The most species-rich families are the Euphorbiaceae (9.2% of all

species recorded from Bukit Bauk), Dipterocarpaceae (6.7%), Rubiaceae

(5.6%), Palmae (4.9%), and Guttiferae (4.9%). The ranking of the first

three families is comparable with other floristic studies in lowland and hill

forests in Peninsular Malaysia. Palms are not usually included in forest

enumerations but are an important component of the flora at Bukit Bauk.

The Palmae is also the family with the highest number of endemic species

(11 out of 31 species). Other families that are well represented on Bukit

Bauk by more than a quarter of their Peninsular Malaysian species are the

Anacardiaceae, Dipterocarpaceae and the Guttiferae.

The flora of Bukit Bauk includes 89 species endemic to Peninsular

Malaysia, 13 of which are endemic to Terengganu and 4 to Bukit Bauk.This

study therefore supports the status of Bukit Bauk as a biodiversity hotspot

for plants, both on grounds of its species richness and the number of

endemic species. It is therefore an area of conservation importance and

should be accorded permanent legal protection, particularly as it has been,

and still is, threatened by habitat distruction or disturbance from road-

building, construction works associated with the microwave station,

quarrying, logging and the lack of a clear boundary that allows intrusion.

Acknowledgements

This study was carried out as a partial fulfillment of the requirements for a

MSc degree at Universiti Putra Malaysia and was supported by a research

assistantship from Universiti Putra Malaysia and IRPA Research Grant

No. 52858 awarded to Ruth Kiew by the Ministry of Science, Technology

a ee.

Flora of Bukit Bauk 2S

and the Environment. I am most grateful to my supervisors, Ruth Kiew

and Quah Soon Cheang, to my co-supervisor Saw Leng Guan, and to

Forest Research Institute Malaysia for allowing me use of herbarium

facilities. I am also much indebted to the KEP staff for their help, and

especially to Kamarudin Saleh, Baya Busu, Mustapha and S. Anthonysamy

for assistance with fieldwork and to Suleka Madhaven for assistance in the

laboratory. Finally, I should like to thank Ruth Kiew for help in the

preparation of this paper.

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Appendix. Floristic Checklist of Bukit Bauk

Species entry begins with * if the species is endemic to Peninsular Malaysia,

followed by species name, source of information, and habitat and

distribution, which follow Turner (1995).

Abbreviations for sources of information are:

TF - Tree Flora of Malaya, Vol. 1-4

AB - FRIM Assession Book

FR - FRIM Database

RK- Riuth Kiew’s index catalogue

98 - Collected and identified in this study

S - Swamp Forest Species identified in this study

Abbreviation for states in Peninsular Malaysia and Singapore follow Turner

(1995): Jh — Johor; Kd — Kedah; K1 — Kelantan; MI — Melaka; NS — Negeri

278 Gard. Bull. Singapore 51 (1999)

Sembilan; Ph — Pahang; Pk — Perak; Pn — Penang; Ps — Perlis; SI — Selangor:

Tg — Terengganu, Sp — Singapore; and P. - Pulau.

DICOTYLEDONS

Acanthaceae

Chroesthes longifolia (Wight) B. Hansen. RK, 98. Lowland forest.

Widespread,

Actinidaceae

Saurauia nudiflora DC. AB. Lowland and hill forest. Kd, Pk, Ph, Sl, Tg,

S. roxburghii Wall. FR. Disturbed hill forest to 1250 m. KI, Tg, Pk, Ph, SI.

Alangiaceae

Alangium ebenaceum (C.B. Clarke) Harms var. ebenaceum. FR, 98.

Lowland forest. Tg and Pk southward.

Anacardiaceae

Bouea oppositifolia (Roxb.) Meisn. 98. Lowland forest to 700 m.

Widespread.

Buchanania arborescens (Blume) Blume. 98. Common on sandy and rocky

coasts and river banks. Widespread.

B. sessifolia Blume. 98, S. Lowland forest to 750 m. Widespread.

Campnosperma auriculatum (Blume) Hook.f. S. Lowland and montane

forest to 1600 m, often in swampy places. Widespread.

Dracontomelon dao (Blanco) Merr. & Rolfe. 98. Lowland forest near rivers.

Kd, Kl Te: Pa Fe Pi, Sl, Ns. E.

*Gluta capituliflora Ding Hou. AB. Hill forest 100-1000 m. KI], Tg.

G. elegens Hook.f. AB, 98. Lowland forest to 300 m. Kd, Kl, Tg, Pk, Pn,

Ph, Sl.

G. pubescens (Ridl.) Ding Hou. AB. Lowland forest, swamps to 400 m, KI],

Tg, Ph, NS, MI, Jh.

G. wallichii (Hook.f.) Ding Hou. AB, 98, S. Lowland and hill forest to

1000 m. KI, Tg, Pk, Ph, Sl, NS, M1, Jh, Sp.

Mangifera griffithii Hook.f. AB. Wet lowland forest to 360 m. Kd, KI], Tg,

Pk, Ph, SI, Ml, Jh, Sp.

M. magnifica Kochummen. AB., Lowland and hill forest. Kd, Kl, Tg, Pk,

Ph, Sl, NS, MI, Jh.

M. quadrifida Jack. S. Lowland forest to 900 m. Kd, KI, Tg, Pk, Ph, Sl, M1,

Jh, Sp.

Melanochyla angustifolia Hook.f. AB, 98, S. Lowland forest to 600 m. Kd,

Flora of Bukit Bauk 279

Kl, Tg, Pn, Ph, Sl, NS, M1, Sh.

M. caesia (Blume) Ding Hou. AB. Lowland and hill forest. K], Tg, Pk, Ph,

Sl, NS; Jhb,

M. fulnivervis (Blume) Ding Hou. AB, 98. Lowland forest to hills 1350 m.

Ki, Tg;-Pk, Ply Sl,NS; Sh.

Parishia maingayi Hook.f. AB. Lowland and hill forest to 600 m on ridge

crests. Tg, Pk, Ph, Sl, Jh, Sp.

P. paucijuga Engl. AB, 98. Lowland to hill forest to 400 m. Tg, Pn, Pk, Ph,

Sl, NS, MI, Jh, Sp.

*Semecarpus trengganuensis Kochummen. TF, AB. Lowland forest. Tg,

Ph.

Swintonia floribunda Griff. 98, S. Lowland forest and hill forest to 1000 m.

Kd, Ki; Pa, Ph, Sl, NS, Jh, Tg.

S. schwenkii (Teijsm. & Binn.) Teijsm. & Binn. AB. Lowland and hill

forest to 700 m. Kd, Tg, Pn, Ph, Sl, NS, Ml, Jh, Sp.

Anisophylleaceae

Anisophylla corneri Ding Hou. 98, S. Lowland and hill forest to 1000 m.

Kid) Ki Te, Pk; PhySl, NS, MI, Jh:

A. disticha (Jack) Baill. AB, RK, 98, S. Lowland to montane forest.

Common and widespread.

A. griffithii Oliv. AB. Hill forest often on ridges. Kd, KI, Tg, Pn, Pk, Ph, Sl,

Ml, Jh, Sp.

A. scortechinii King. AB. Lowland forest. Kd, Tg, Pn, Pk, Ph, Jh.

Annonaceae

Cyathocalyx carinatus (Ridl.) J. Sinclair. AB. Lowland forest. Pk, Sl, Tg.

C. pruniferus (Maingay ex Hook.f. and Thomson) J. Sinclair. AB, 98.

Lowland and hill forest, Kl, Tg, Pk, Ph, Sl, Ml, Jh. E.

C. ridleyi (King) J. Sinclair. AB. Damp lowland forest. Ph, Jh, Tg, Sp.

*Fnicosanthum fuscum (King) Airy Shaw. AB. Lowland and hill forest.

Pk, Ph.

Fissistigma fulgens (Hook. f. & Thomson) Merr. AB. Lowland forest. K],

Tg, Pk, Ph, Sl, M1, Jh, Sp.

*Goniothalamus fulvus Hook.f. & Thomson. AB. Lowland forest. Tg, Ph,

Ml, Jh.

G. macrophyllus (Blume) Hook.f. & Thomson. 98. Common in lowland

forest. Kd, Kl, Tg, Pk, Ph, Sl, NS, MI, Jh, Sp.

G. tapis Mig. AB. Lowland and hill forest. Pn, Kl, Tg, Pk, Jh, Sp.

Mezzitia parviflora Becc. AB. Lowland forest to 300 m. Kd, Pn, Tg, Pk,

Ph, Sl, NS, MI, Jh, Sp.

Monocarpia marginalis (Scheff.) J. Sinclair. AB, S. Lowland forest. Kd,

280 Gard. Bull. Singapore 51 (1999)

Pn, KI, Tg, Pk, Ph, Sl, NS, Ml, Jh.

Phaeanthus optalamicus Roxb. ex G. Don. 98. Lowland forest. Kl], Pn, Pk,

SI, NS, MI, Jh, Tg, Sp.

Polyalthia bullata King. AB, 98. Lowland forest. Pk, Ph, Sl, NS, MI, Jh, Tg.

P. cauliflora Hook.f. & Thomson var. beccarii. AB. Lowland forest. Tg,

Ph, NS, Ml, Jh, Sp.

*P. dumosa King. AB. Montane forest. Tg, Pk.

P. lateriflora (Blume) King. S. Lowland forest to lower montane forest. Pk,

Ph; Sly NSYMi Ii Pe. Sp.

P. rumphii (Blume) Merr. AB. Lowland and hill forest. Kd, Tg, Pk, Ph, Sl,

NS, MI, Jh, Sp.

P. sumatrana (Miq.) Kurz. AB Lowland and hill forest. Kl, Tg, Pk, Ph, Sl,

NS, Jh; Sp:

*Pseuduvaria cerina J. Sinclair. FR. Hill forest. Tg, only Bukit Bauk.

P. macrophylla (Oliv.) Merr. var. macrophylla. 98, S. Lowland and hill

forest: Kd Pay Ri We, Pk,’ Ph, si NS/ ME

Xylopia caudata Hook.f. & Thomson. AB, 98, S. Lowland forest. Pk, Ph,

Sl; NS, Mi, ihyiie,: Sp.

X. ferruginea (Hook.f. & Thomson) Hook.f. & Thomson var. ferruginea.

AB. Lowland forest. Kd, Kl, Tg, Pk, Ph, Sl, NS, MI, Jh, Sp.

X. ferruginea (Hook. f. & Thomson) Hook.f’ & Thomson var. oxyantha

(Hook.f. & Thomson) J. Sinclair. 98, S. Lowland forest, Ph, Sl, NS, Jh, Tg,

5 0)

X. malayana Hook.f. & Thomson var. malayana. AB, 98. Lowland forest.

Te, PK; Ph, SIME ih Sp.

*X. subdehiscens (King) J. Sinclair. AB. Lowland and hill forest. Pn, Tg,

Pk, PS: ,

Apocynaceae

Alstonia angustiloba Wall. ex A.D.C. 98. Lowland to hills to 700 m. Common

and widespread.

Kopsia arborea Bl. RK. Lowland. Pn, Tg.

*K. macrophylla Hook.f. FR. Kl, Tg, Pk, Ph, NS, MI, Jh.

K. singapurensis Ridl. 98. Common in lowland swamp forest. Pk, Ph, SI,

NS, Ji, ‘TeySy:

Parameria laevigata (A. L. Juss.) Moldenke. 98. Lowland forest. Pn, Pk,

Ph, SI, NS, MI, Tg, Sp.

Tabernaemontana corymbosa Roxb. ex Wall. S. Lowland and hill forest to

900 m. Kd, Ki, Tg;iPn, Pk; Ph; Sly NS} Ml, FaySp:

*Urceola maingayi D. J. Middleton. FR. Ph, Tg.

U. torulosa Hook.f. AB. Lowland forest. Kl, Tg, Pk, Ph, Sl, MI, Jh, Sp.

Flora of Bukit Bauk 281

Aquifoliaceae

Ilex cymosa Blume. 98. Lowland forest, swamps and secondary forests.

Widespread.

I. macrophylla Hook.f. 98. Lowland, well drained forest. Widespread.

Araliaceae

Schefflera oxyphylla (Miq.) R. Vig. FR. Lowland and hill forest. Kl, Tg,

Pap Pk»PhySi).NS, Ml, Jh.

Aristolochiaceae

Thottea grandiflora Rottb. RK. Lowland and hill forest to 600 m. Tg, PK,

Ph, NS, M1, Sh, Sp.

Thottea sp. 98.

Burseraceae

Canarium caudatum King. AB. Lowland forest. Widespread.

C. littorale Blume. AB, 98, S. Lowland forest, occasionally in highlands.

Widespread.

C. patentinervium Maiq. 98. Lowland forest and hill forest, understorey. Kd,

Ki Vo"Pk, Ph; Sl, NS, Mi, Jh-Sp.

*Dacroydes breviracemosa Kalkman. TF. Lowland forest. Tg, only from

Bukit Bauk FR.

D. costata (Benn.) H.J. Lam. S. Lowland and hill forest. Kd, Tg, Pn, Pk, Sl,

INS, oh, Sp.

D. rostrata (Blume) H.J. Lam. 98, S. Hill forests to 600 m, swamps. Ps, Kd,

De, Pos Pk PhS NS, Mil, Jhy Sp:

Santiria apiculata Benn. var. rubra (Ridl.) Kalkman. AB. Lowland and hill

forest. Widespread.

S. griffithii (Hook.f.) Engl. AB, 98. Lowland forest to 300 m. Widespread

except for far north.

S. laevigata Blume. S. Lowland to montane forest. Kd, Kl, Tg, Pn, Pk, Ph,

MI, Jh, Sp.

S. rubiginosa Blume var. rubiginosa. AB, 98, S. Lowland to mountains.

Kokelp,. Pk, Ph St NS; th)Sp.

S. tomentosa Blume. 98, S. Swampy lowland forest to 250 m. Kd, KI, Tg,

PaePls, Ph; Sk: NS,\ Mb -Jh, Sp.

Bombacaceae

Durio affinis Becc. AB. Lowland forest. Tg, Jh.

D. griffithii (Mast.) Bakh. S. Lowland forest. Kd, Kl, Tg, Pk, Ph, Sl, NS,

Ml, Jh, Sp.

D. oxleyanus Griff. AB. Lowland forest, Kd, Tg, Pk, Ph, Sl, NS, MI, Jh.

282 Gard. Bull. Singapore 51 (1999)

D. wyatt-smithii Kosterm. AB, 98, S. Lowland forest. KI, Tg.

Celastraceae

Euonymus javanicus Blume. 98, S. Lowland and hill forest to 400 m.

Widespread.

Lophopetalum floribundum Wight. 98. Lowland forest to 360 m. KI], Pn,

Pk, Ph, NS, Jh, Tg.

Chrysobalanaceae

Atuna nannodos (Kosterm.) Kosterm. AB. Lowland and hill forest to 750

m. Pn, Tg, Pk, Ph, NS, Jh.

A. racemosa Raf. ssp. excelsa (Jack) Prance. AB, 98, S. Lowland and hill

forest to 750 m. Kd and Tg southwards.

Maranthes corymbosa Blume. 98. Mostly coastal but also in the mountains.

Ps, Kd, Pn, Kl, Pk, Ph, Sl, NS, Ml, Jh, Tg, Sp.

Parinari costata (Kofth.) Blume. AB. Lowland forest. Widespread except

for far north.

P. elmeri Merr. 98, S. Lowland and hill forest to 600 m. SI, M1, Jh, Tg.

P. oblongifolia Hook.f. 98. Lowland forest. Kd to Jh.

P. rigida Kosterm. AB. Lowland forest, swamps. Kl, Tg, Ph, NS, Jh, Sp.

Compositae

Elephantopus mollis Kunth. 98. Weed from tropical America.

Mikania cordata (Burm.f.) B.L. Rob. 98. Forest margins, waste places.

Widespread.

Crypteroniaceae

Crypteronia griffithii C.B. Clarke. AB. Lowland and hill forest to 600 m.

Kd, Tg, Pn, Pk, Ph, Sl, NS, Ml, Jh, Sp.

Connaraceae

Connarus semidecandrus Jack. AB. Common in open country. Widespread.

Chloranthaceae

Chloranthus erectus (Buch.-Ham) Verdc. 98. Lowland and hill forest. Pn,

Pk, Ph, SI, NS, Ml, Jh, Tg, Sp.

Combretaceae

Lumnitzera racemosa Willd. AB. Mangroves, all coasts but commoner on

the east coast.

Flora of Bukit Bauk 283

Dilleniaceae

Acrotrema costatum Jack.RK. Widespread.

Dillenia grandiflora Wall ex Hook.f. & Thomson. 98. Lowland forest and

hill forest to 1000 m. Kd, Pn, KI, Tg, Pk, Ph, Sl, NS, MI, Jh, Sp.

D. reticulata King var. psilocarpella Hoogl. S. Lowland forest. Kl, Pk, Ph,

Sl, NS, MI, Jh, Tg.

Dipterocarpaceae

Anisoptera curtisti Dyer ex King. AB, 98. Lowland and hill forest to 700 m.

Widespread.

A. laevis Ridl.. AB, FR. Widespread.

Dipterocarpus caudatus Foxw. ssp. penangianus (Foxw.) P. S. Ashton. FR.

Coastal hill forest. Pn, Kl, Pk, Ph, Jh, Tg, Sp.

. concavus Foxw. FR. Lowland forest. Pk, Ph northward.

. costulatus Slooten. FR, S. Lowland forest. Widespread.

. eurynchus Mig. FR, AB. Lowland forest. Tg, Ph, NS, Jh.

. gracilis Blume. FR. Lowland forest. Widespread.

. lowii Hook.f. S. Lowland forest. Scattered.

. rigidus Ridl. FR, 98. Coastal hill forest. Tg southwards.

. sublamellatus Foxw. 98. Lowland forest. Widespread.

. verrucosus Foxw. ex Slooten. AB, 98, S. Lowland forest. Commoner in

the south.

Dryobalanops sumatrensis (J.F. Gmel.) Kosterm.AB, 98, S. Gregarious in

sandy lowland forest. Tg, Ph, Sl, NS, Jh.

Hopea dryobalanoides Mig. AB. Lowland forest. Widespread

H. dyeri F. Heim. AB, FR. Lowland and hill forest to 1000 m. Widespread.

A. griffithii Kurz. FR. Lowland forest. Widespread.

H.. kerangasensis P.S. Ashton. FR. Lowland forest on sandy soils. Tg, Ph.

H. mengerawan Mig. 98. Swampy lowland forest. Tg, Ph and NS southward.

H. myrtifolia Mig. AB, FR, 98. Lowland forest. Pk and Tg, Ph southward.

H. nutans Ridl. FR. Swampy lowland forest, east coast, Tg southward.

*H. pubescens Ridl. AB, FR. Lowland forest, KI, Ph, Tg.

*H. sulcata Symington. 98. Lowland forest to 400 m. Tg, Pk, SI, Jh.

Parashorea stellata Kurz. AB. Lowland and hill forest to 650 m. Tg and Pk

northward.

Shorea acuminata Dyer. FR, 98, S. Lowland forest.Pk, Tg southward.

*S. bentongensis Foxw. FR. Lowland forest. Sl, Ph, Jh, Tg.

S. bracteolata Dyer. AB, 98. Coastal hills to 600 m. Kd, Pn, KI, Tg, Pk, Ph,

SI, MS, MI, Jh, Sp.

S. curtisii Dyer ex King. AB, 98. Ridges to 850 m. Widespread

S. faguetiana F. Heim. FR. Mostly in hills to 700 m. Kd, Pn, KI, Tg, Pk, Ph,

Sl. NS, MI, Jh.

Seoeees >

284 Gard. Bull. Singapore 51 (1999)

S. falcifera Dyer ex Brandis. AB, 98. Coastal hill. Tg, Ph

S. foxworthyi Symington. 98. Lowland and hill forest to 700 m. Kd, KI, Tg,

Pk; Ph, Shh:

S. glauca King. FR, 98. Coastal hills to 600 m. Ps, Kd, Pn, Kl, Tg, Pk, Ph,

NS, MI, Jh.

S. hypochra Hance. AB, FR. Lowland forest. Sl and Ph northward.

S. macroptera Dyer. AB, FR, 98, S. Lowland and hill forest to 900 m. Kd,

Pn, Kl, Tg,,Pk, Paes, NS, MI, Jh; Sp:

S. maxwelliana King. AB, 98. Lowland and hill forest to 700 m. Pn and Tg

southward.

S. multiflora (Burck) Symington. 98, S. Lowland and hill forest to 700 m.

Widespread.

S. parvifolia Dyer ssp. parvofolia . FR, 98. Lowland forest. Ph, Jh, Tg.

S. pauciflora King. AB, FR. 98. Lowland and hill forest. Widespread.

S. singkawang (Miq.) Mig. ssp. singkawang . AB, S. Lowland forest below

400 m. Kd, KI to Jh.

S. singkawang (Miq.) Mig. ssp. *scabrosa P.S. Ashton. AB, FR. Forest

near sea. Tg, Ph. )

S. teysmanniana Dyer ex Brandis. 98. Peat swamp forest. Sl, Tg.

*Vatica cuspidata (Ridl.) Symington. AB. Lowland ridge forest, near sea.

Widespread.

V. pauciflora (Korth.) Blume. S. Swampy lowland forest. Widespread.

*V. scortechinii (King) Brandis. 98. Lowland and hill forest to 1800m, Tg,

Pk, Ph, SI.

V. umbonata (Hook.f.) Burck. FR. Lowland and hill forest. Tg, Ph.

V. venulosa Blume. FR. Swampy lowland forest. Pk, Ph, Tg.

Erythroxylacaceae

Erythroxylum cuneatum (Miq.) Kurz. 98. Lowland and hill forest. Common

near sea, throughout.

Ebenaceae

*Diospyros adenophora Bakh. S. Lowland forest, also on limestone. K],

Ph, NS, to.

D. apiculata Hiern. AB, FR, 98, S. Lowland and hill forest. Pn, Kl, Tg, Pk,

Ph, NS, MI, Jh.

*D. argentea Griff. 98, S. Lowland and hill forest to 800 m, Tg, Pk, Ph, Sl,

NS, MI, Jh, Sp.

*D. clavigera C.B. Clarke. AB, FR, 98. Lowland forest to 500 m. Pn, Tg,

Pk, Ph, Sl, NSpMipdhy Sp:

D.confertiflora (Hiern) Bakh. 98. Lowland to montane forest at 1250m.

Kd, Pn, Kl; Tg; Ph, SI, Sh.

Flora of Bukit Bauk 285

*D. ismailiti Ng. AB, FR. Lowland and hill forest to 1000 m, Kd, Tg, Ph, Sl,

NS, Jh.

D. lanceifolia Roxb. 98. Lowland and hill forest to 700 m. Throughout.

D. pauciflora King & Gamble. 98. Lowland and hill forest to 600m. Tg, Pk,

Sh.

D. pendula Hasselt ex Hassk. AB. Lowland and hill forest to 1100 m. Kd,

Pa, Pe} Pk, Ph, Si, dh:

D. pyarrhocarpa Mig. AB. Lowland and hill forest to 900 m. Kd, Tg, Pk,

Sl, NS, MI, Jh.

*D. ridleyi Bakh. AB. Lowland and hill forest to 1200m. Pn, Tg, Ph, SI, Jh.

D. rigida Hiern. 98. Lowland forest. Kl, Tg, Pk, Ph, SI, Jh.

*D. rufa King & Gamble. 98. Lowland to lower montane forest at 1500m

Kd, Tg, K1, Pk, Ph, NS, MI, Jh.

*D. scortechinii King & Gamble. 98. Hill and montane forest at 400-1500m,

Ki, Tey Pk, Ph Ns.

*D. singaporensis Bakh. 98. Lowland forest to 600m. Kd, Pk, Ph, Sl, NS,

MI, Jh, Tg.

D. sumatrana Mig. 98. Lowland to montane forest at 1500m. Common.

*D. tristis King & Gamble. AB. Lowland to lower montane forest at 1200m.

Ka, Te PepPhesSl, th.

Elaeocarpaceae

Elaeocarpus nitidus Jack var. nitidus. 98, S. Lowland to montane forest at

1500m. Common throughout.

E. palembanicus (Mig.) Corner. AB, 98. Lowland and hill forest to 900m.

Kids Puy Kis he; Ph; Sly NS, Th:

E. pedunculatus Wall. ex Mast. 98. Lowland and hill forest to 670m.

Common throughout.

E. petiolatus (Jack) Wall. 98. Lowland and hill forest to 500m. Widespread.

*F’. polystacyhus Wall. ex Mull. Berol. AB. Lowland and hill forest to

100m, Kl, Tg; Pk) Ph, Sl,NS, JhySp:

E. stipularis Blume var. stipularis. 98. Lowland to lower montane forest at

1250m. Throughout.

Euphorbiaceae

Agrostistachys longifolia (Wight) Benth. var leptostachya (Pax & K. Hoffm.)

Whitmore. AB, 98, S. Lowland and hill forest. Kd, Tg, Pk, Ph.

Alchornea rugosa (Lour.) Mull. Arg. var rugosa. AB. Lowland forest. Ph

northward.

Antidesma coriaceum Tul. 98. Lowland to lower montane forest at 1500 m.

Except for far north.

A. cuspidatum Mull.. Arg. AB. Lowland forest. Common throughout.

286 Gard. Bull. Singapore 51 (1999)

A. leucocladon Hook.f. AB. Lowland forest. Ps, Kd, Kl, Tg, Pk, Sl, M1.

Aporusa aurea Hook.f. 98, S. Lowland and hill forest. Scattered throughout.

A. bracteosa Pax & K. Hoffman. AB, S. Lowland forest. K1, Ph. Southward.

A. falcifera Hook.f. 98, S. Lowland and hill forest to 540 m. Scattered

throughout.

A. frutescens Blume. AB, 98. Kd, Pn, KI, Pk, Jh, Tg, Sp.

A. lunata (Mig.) Kurz. 98. Lowland to montane forests at 1500m. Pn, Pk,

Ph, Jh, Tg, Sp.

A. microstachya (Tul.) Mull. Arg. AB. Hillsides in lowland forest to 400 m.

Kd, Tg southward.

A. nigricans Hook.f. 98. Lowland and hill forest to 600 m. Scattered

throughout.

A. prainiana King ex Gage. 98. Hillsides in lowland forest. Kd, Pn, Tg, Pk,

Ph, MI, Jh, Sp.

A. symplocoides (Hook.f.) Gage. AB. Lowland forest. Common throughout.

Baccaurea brevipes Hook.f. S. Damp lowland forest. Throughout.

B. maingayi Hook.f. S. Hill forest at 150-780 m. Pk, Tg, NS, Jh.

B. motleyana (Mull. Arg.) Mull. Arg. AB, 98. Lowland and hill forest to

900 m. Scattered throughout.

A.minor Hook.f. S. Lowland forest. KI, Pk, Ph, NS, Jh, Tg, Sp.

B. parviflora (Mull. Arg.) Mull. Arg. AB, 98, S. Widespread, commoner in

the north, lowland and hill forest to 600 m.

B. racemosa (Reinw.) Mull. Arg. AB, 98. Lowland forest, throughout,

commoner in the south.

B. reticulata Hook.f. 98. Lowland forest to 300 m. Tg and Pk southward.

B. sumatrana Mull. Arg. AB. Lowland forest. Common throughout.

Blumeodendron kurzii (Hook. f.) J.J. Sm. AB, S. Lowland forest. Kd, K],

Pk, Phy Sh,Fe¢:

B. tokbrai (Blume) J.J. Sm. AB, 98. Lowland forest to 450 m, often in

swamps. Tg, Pk, Ph, Sl, NS, Jh, Sp.

Breynia coronata Hook.f. 98. Lowland and mountains to 1200 m.

Widespread.

Cheilosa montana Blume. AB. Lowland forest. KI, Tg, Ph, Sl, Sp.

Cleistanthus decurrens Hook.f. AB. Lowlands, often near limestone. Pn,

TesPk, Pay NS ae

*C. glaucus Jabl. AB, 98. Limestone and rocky outcrops. Pk, Tg.

C. hirsutulus Hook. f- AB. Lowland forest, rarely on limestone. Scattered

throughout.

*C. podocarpus Hook.f. AB. Lowland and hill forest. Tg, Pk, Jh.

C. sumatranus (Miq.) Mull. Arg. 98. Lowland forest. Ph, Ml, Jh, Tg, Sp.

Croton laevifolius Blume. AB, 98, S. Lowland, forest margins. Common

throughout.

Flora of Bukit Bauk 287

Drypetes laevis (Mig.) Pax & K. Hoffm. 98. Lowland. Kd, Ph, NS, Jh, Tg.

D. longifolia (Blume) Pax & K. Hoffm. 98. Lowland forest. Widespread.

D. pendula Ridl. AB, 98. Lowland forest. Common throughout.

Endospermum diadenum (Miq.) Airy Shaw. 98, S. Lowland to lower

montane forest at 1000 m. Common throughout.

Glochidion superbum Baill. AB. Lowland forest. Common throughout

G. wallichianum Mull. Arg. 98. Lowland forest. Sl, Ph northward.

Hevea brasiliensis (Willd. ex A. Juss.) Mull. Arg. 98. Naturalised from

S.America.

*Macaranga amissa Airy Shaw. AB. Swampy lowland forest. Tg, Pu, MI,

oh.

M. heynei 1.M. Johnst. 98. Lowland forest and secondary forest. Ps, Kd,

Pik; Phy sh NS, Mi:cdh, Te, Sp.

M. hypoleuca Rchb. f. & Zoll. AB. Secondary forest, throughout.

M. laciniata Whitmore & Airy Shaw. AB. Secondary forest. Kl, Tg, Ph.

M. triloba (Blume) Mull. Arg. 98. Lowland forest and secondary growth.

Common throughout.

Mallotus dispar (Blume) Mull. Arg. AB. Usually on limestone. Scattered

in the north.

*M. griffithianus Hook.f. S. Lowland forest. Throughout.

M. paniculatus (Lam.) Mull. Arg. 98. Secondary forest in lowland and hill

forest to 900 m. Throughout.

M. philippensis (Lam.) Mull. Arg. 98. Lowland forest margins, sometimes

on limestone. Kd, Tg, Pk, Ph, NS.

Neoscortechinia kingii (Hook.f.) Pax & K. Hoffm. AB, 98, S. Lowland

forest’ Ps. Kad) Pn, Ki T 9, Pk, Ph, SLiNS; oh Sp.

N. nicobarica (Hook.f.) Pax & K. Hoffm. 98. Lowland or more frequently

on hill forest. Kd, Kl, Ph, Sl, Jh, Tg.

Paracroton pendulus (Hassk.) Miq. 98. Damp or swampy lowland forest.

Kd, Pn, Tg, Pk, Ph, NS, Ml, Jh, Sp.

Phyllanthus roseus (Craib & Hutch.) Beille. AB. Lowlands or hills to 750

m. P. Langkawi, Kd, Pn, Tg, Pk.

Pimelodendron griffithianum (Mull. Arg.) Benth. 98, S. Lowland forest.

bel Pes Pi; -PhyS1 NS, Mt, Ih, Sp:

Ptychopyxis javanica (J.J. Sm.) Croizat. AB. Lowland forest. Kd, Pn, Pk,

Tg.

Sapium beccatum Roxb. 98. Lowland and hill forest to 600 m. Throughout.

Suregada multiflora (Juss.) Baill. var multiflora. AB. Rocky and sandy

shores, inland in rocky places to 750 m. Widespread. |

Trigonostemon laevigatus Mull. Arg. 98. Coasts. P. Langkawi, Pn, Tg, Pk,

P. Tioman, Jh.

T. longifolius Baill. AB. Lowland and hill forest to 600 m. Throughout.

288 Gard. Bull. Singapore 51 (1999)

T. malaccanus Mull. Arg. AB. Lowland forest. Widely scattered.

Fagaceae

Castanopsis lucida (Nees) Soepadmo. 98. Lowland to lower montane forest.

Kd,.Pn, KE, Pk, Ph, Si, NS, M1, Jay Pesca:

C. megacarpa Gamble. S. Lowland and hill forest to 600 m. Kd, K1, Tg,

Pk, Ph,:Sig¢NS; Mile ih, Sp.

C. rhamnifolia (Miq.) A. DC. 98. Lowland to montane forest at 1600 m.

K1;:Pu, Phy Sire:

C. schefferana Hance. 98, S. Lowland forest. Kd, Tg, Ph, NS, Jh, Sp.

C. tunggurut (Blume) A. DC. 98. Lowland forest. S1, NS, M1, Tg.

Lithocarpus_ cantleyanus (King ex Hook.f.) Rehder. 98. Lowland and hill

forest to. 830 mimicieePk, Ph, 81,1 le: Sey

L. leptogyne (Korth.) Soepadmo. 98. Uncommon, lowlands. Pk, Jh, Tg.

L. lucidus (Roxb.) Rehder. S. Forest to 1600 m. Throughout.

*I. maingayi (Benth.) Rehder. 98. Hill forest at 600-1000 m. Kd, Pn, K1,

Tg,Pky Phy SipNar M11 J Sh.

L. rassa (Miq.) Rehder. 98. Lowlands but commoner in the mountains.

Kd, Pn; Ki; oP ky Ph; $1, NS, M1, dh.

L. wallichianus (Lindl. ex Hance) Rehder. 98. Lowland and hill forest.

Throughout.

L. wrayi (King) A. Camus. AB. Lowland to lower montane forest at 1350

m. Kd, K1, Tg, Pk, Ph, NS.

L. urceolaris (Jack) Merr. AB. Lowland to lower montane forest. Tg, Ph,

Shi:

Quercus argentata Korth. AB. Lowland to montane forest. Tg, Ph, MI, Jh,

Sp:

Flacourtiaceae

Casearia velutinosa Ridl. AB. Lowland forest often near streams. Tg, Pk,

Sd; She

Homalium dictyoneurum (Hance) Warb. AB. Lowland forest. Kd, Tg, Jh.

Hydnocarpus woodii Merr. AB. Lowland and hill forest to 1000 m. Ps, Tg,

Pk, Ph,Si, NSs th:

Ryparosa hullettii King. 98, S. Swampy lowland forest. Tg, Pk, Jh, Sp.

Gesneriaceae

Cyrtandra sp. Ridl. RK.

*Henckelia heterophylla (Ridl.) A. Weber. AB, 98. By streams in lowland

and hill forest. Ph, Jh, Tg.

*H. rugosa (Ridl.) A. Weber. 98. Lower montane forest. Kd, K1, Tg, Pk,

Ph, Si:

Flora of Bukit Bauk 289

*H. miniata (Kiew) A. Weber. RK. Lowland forest on sandstone. Tg (only

Bukit Bauk and Bukit Chabang).

H. platypus (C.B. Clarke) A. Weber. RK. 98. Lowland and hill forest to

1000 m. Widespread.

*H. puncticulata (Ridl.) A. Weber. AB. Lowland forest. Tg, Ph, Jh, Sp.

*H. salicinoides (Kiew) A. Weber. RK. Lowland forest. K1, Tg.

Guttiferae

Calophyllum calaba L. var. bracteatum (Wight) P.F. Stevens. AB, FR.

Lowland and hill forest. Throughout.

C. canum Hook.f. AB. Lowland forest. Kd, Pn, K1, Tg, Pk, Ph, S1, NS,

M1, Jh.

C. ferrugineum Ridl. var. ferrugineum. 98. Lowland forest. Tg, Jh, Sp.

C. ferrugineum Ridl. var. *oblongifolium (T. Anderson) P.F. Stevens. AB,

98. Lowland and hill forest to 760 m. K1, Tg, Pk, Ph, S1, NS, M1, Sh.

C. flavoramulum M.R. Hend. & Wyatt-Sm. AB. Lowland forest. Tg, Ph,

Sh.

*C. gracillimum M.R. Hend. & Wyatt-Sm. AB, 98. Lowland and hill forest

to 1000 m. Tg, Ph, S1.

C. inophylloide King var. singaporense Hend. & Wyatt-Sm. TF. Lowland

and hill forest, hillsides to montane forest at 1200 m. K1, Jh, Tg, Sp.

C. rufigemmatum M.R. Hend. & Wyatt-Sm. AB. Lowland forest. Tg, Ph,

Ih, Sp:

C. sclerophyllum Vesque. AB. Swamp forest. Kd, K1, Tg, Pk, Ph, Jh.

C. teysmanii Miq. var teysmanii AB, 98. Lowland to lower montane forest

at 1200m. K1, Tg, Ph, Jh, Sp.

C. teysmanii Mig. var inophylloide (King) P.F. Stevens. AB, FR. Lowland

to lower montane forest. Kd, Pn, Tg, Pk, Ph, S1, NS, Jh.

*C. subhorizontale M.R. Hend. & Wyatt-Sm. 98. Lowland forest. Ph, S1,

NS, Tg.

C. wallichianum Planch. & Triana var. *wallichianum. AB, 98. Lowland

and Inil forest: Kd, Pn, K1, Pk, NS, Tg.

C. wallichianum Planch & Triana var. incrassatum (M.R. Hend. & Wyatt-

Sm.) P.F. Stevens. AB. Lowland forest. K1, Tg, Ph, S1, M1, Jh, Sp.

Cratoxylum arborescens (Vahl) Blume. AB. Common in lowland, swamp

horest and hills. Pn, Ki, Tg, Pk, Ph, SI, NS, ML, Jh, Sp.

C. formosum (Jack) Dyer. 98. Lowland forest to 600 m. Throughout.

Garcinia eugeniifolia Wall. ex T. Anderson. AB, 98. Lowland to lower

montane forest. P. Langkawi, Kd, Pn, Tg, Pk, Ph, Jh, Sp.

G. griffithii. AB, S. Lowland and hill forest to 900 m Scattered throughout.

G. malaccensis Hook.f’ AB, 98. Lowland forest to 540 m. KI, Tg, Ph, Sl,

NS, Jh.

290 Gard. Bull. Singapore 51 (1999)

G. merguensis Wight. AB. Lowland forest. P. Langkawi, Kd, Pk, Ph, Tg.

G. murdochii Ridl. AB. Lowland forest, also on limestone. Kd, Tg, Pk, Jh.

G. nervosa Mia. var. nervosa. 98, S. Lowland forest. Scattered widely.

G. parvifolia (Miq.) Mig. AB, 98, S. Lowland forest to 600 m. Throughout.

G. penangiana Pierre. AB. Lowland forest to 300 m. Pn, Tg, Pk, Ph.

*G. pyrifera Ridl. 98. Lowland and hill forest, Pn, Kl, Tg, Pk, Sl, Jh, Sp.

G. rostrata (Hassk.) Mig. 98. Lowland and hill forest. Scattered throughout.

G. scortechinii King. 98, S. Lowland and hill forest to 700 m. Common

throughout.

Mesua assamica (King & Prain) Kosterm. AB. Lowland forest. Kd, KI], Tg,

Ph, NS, Jh.

M. ferrea L. AB, 98, S. Lowland forest to 480 m. Throughout but commoner

in the north.

M. grandis (King) Kosterm. 98, S. Lowland and hill forest to 1200 m. Tg,

Pk southward.

M. lepidota T. Anderson var. lepidota. 98. Lowland forest. Tg, Pk, Ph, Sl,

NS, MI, Jh.

M. racemosa (Planch. & Triana) Kosterm. 98, S. Lowland forest. Kd, Kl],

Tg, Pk, (Ph, Si Wit.

*M. sp.C. TF. Lowland forest. Ph, Tg. (Bukit Bauk and Gunung Tapis).

*M. sp. E. TF. Lowland forest, Tg, Ph.

Icacinaceae

Gomphandra quadrifida (Blume) Sleumer var. ovalifolia (Ridl.) Sleumer.

AB Lowland to lower montane forest to 1200 m. Kd, Kl, Tg, Pk, Ph, Sl,

NS... MI, Ih. So,

Gonocaryum sp. A. TF. Tg (Bukit Bauk only).

Stemonurus malaccensis (Mast.) Sleumer. S. Lowland and hill forest to

1300 m. Throughout.

Irvingiaceae

Irvingia malayana. 98. Lowland forest. Kd, Kl, Tg, Pk, Ph, Sl, NS, M1, Jh,

Sp.

Ixonanthacaeae

Allantospermum borneense Forman. AB, TF. Hill to lower montane forest.

Kd, Tg, Pk.

Ixonanthes reticulata Jack. AB. Lowland and hill forest. Throughout.

Junglandaceae

Engelhartia serrata Blume. AB. Lowland to montane forest at 2000 m. Kd,

Po, Pk, Phi, Jn, 72, a0.

Flora of Bukit Bauk 291

Lauraceae

Actinodaphne macrophylla (Blume) Nees. S. Lowland to montane forest.

KI 'Tg, Pk, Ph; Sl; Ml; Jh:

*A. pruinosa Nees. 98. Lowland and hill forest. Kd, Pn, Sl, NS, M1, Tg, Sp.

Alseodaphne insignis Gamble. AB. Lowland forest. Tg, Pk, Ph, SI.

A. pendulifolia Gamble. AB. Lowland forest. Tg, Pk, Ph, Sl, Jh.

Beilsmiedia palembanica (Miq.) Kosterm. 98. Lowland to montane forest.

Pky. Phys) MLE Sh, Fe:

B. perakensis Gamble. AB. Lowland and hill forest. KI, Tg, Pk, Ph, SI, NS.

Cinnamomum iners Reinw. AB, 98. Primary and secondary forest in lowland

and hills. Common throughout.

*C. mollissimum Hook.f. 98, S. Lowland and hill forest. Pn, Kl, Tg, Pk, Ph,

Sl, NS, Ml, Jh.

Cryptocarya griffithiana Wight. 98, S. Lowland forest. Pk, Ph, Sl, NS, MI,

jh, ‘Pe; Sp:

C. rugulosa Hook.f. 98. Lowland and hill forest. Kd, Pn, Pk, Tg, M1, Jh, Sp.

*Endiandra maingayi Hook.f. AB. Lowland to montane forest at 1200 m.

RD PK Phy SLi NS, Th, Tg.

Litsea castanea Hook.f. AB, 98, S. Lowland to montane forest. Kd, KI, Tg,

Pe Pre St NS, Mi, Jh.

L. costalis (Nees) Kosterm. 98, S. Lowland and hill forest to 500 m. Pk, Ph,

eS Wil, Sp.

L. johorensis Gamble. AB. Lowland to lower montane forest at 1200 m.

Rete, PPh, NS, Jh.

L. magnifica (Miq.) Villar. 98, S. Lowland and hill forest. Pn, Kl, Ph, MI,

ean.

L. sessiliflora Hook.f. AB. Lowland and montane forest to 1500 m. Kd, KI],

Te, PE, Pasi NS, Sh.

L. tomentosa Blume. 98. Lowland and hill forest to 750 m. Pn, Pk, Ph, Sl,

Tg.

Nothaphobe coriacea (Kosterm.) Kosterm. AB. Peat swamp forest, Tg, Ph,

Sl.

Lecythidaceae

*Abdulmajidia chaniana Whitmore. 98. Lowland forest to 570 m. Ph, Tg,

Sh.

*Barringtonia fusiformis King. AB. Lowland forest, often near rivers. Pk

to Jh.

B. macrostachya (Jack) Kurz. AB, 98. Lowland forest, rarely in

mountains. Throughout.

B. pendula (Griff.) Kurz. AB, 98. Lowland forest to 500 m. Throughout.

B. scortechinii King. AB, S. Lowland and hill forest to 1200 m. Throughout.

292 Gard. Bull. Singapore 51 (1999)

Leeaceae

Leea indica. 98. Lowland forest, particularly secondary regrowth.

Throughout.

Leguminosae

Adenanthera malayana Kosterm. 98. Lowland and hill forest to 900 m.

Widespread but scattered.

A. pavonina L. 98. Lowland forest usually near the sea. East coast and

offshore islands.

Archidendron bubalinum (Jack) I.C. Nielsen. AB, 98, S. Lowland forest,

often in secondary forest. Widespread.

A. clypearia (Jack) I.C. Nielsen ssp. clypearia var. clypearia. $. Lowland

forest to 1200 m. Widespread.

A. contortum (Martelli) I.C. Nielsen. AB. Common in secondary forest,

usually near the sea. Widespread.

A. ellipticum (Blume) I.C. Nielsen. 98. Lowland forest including secondary

forest. Widespread.

Callerya artopurpurea (Wall.) Schot. 98. Open country. Widespread.

Crudia curtisii Prain. AB, TF. Lowland and hill forest. Tg, Pk, Pn, Ph, Sl,

MI. E.

Cynometra malaccensis Meeuwen. 98, S. Lowland and hill forest to 600 m.

Pk, Ph, Si, Mi. -Te.

C. ramiflora L. 98. Riverbanks and swamps especially near coast.

Widespread.

Dialium platysepalum Baker. 98, S. Lowland forest, Scattered throughout.

*Fordia albiflora (Prain) Dasuki & Schot. AB, 98. Lowland forest often

near rivers: fe, Pk Phy SiyJir

Intsia palembanica Miq. S. Lowland forest. Common throughout.

Ormosia venosa Baker. TF. Hill forest. Tg, Sl, MI.

Parkia singularis Mig. AB, 98. Lowland and hill forest to 500 m. Tg, Pk,

Ph: Sis Sir

P. speciosa Hassk. 98, S. lowland and hill forest to 900 m. Widespread.

Saraca cauliflora Baker. 98, S. Lowland and hill forest, often riverine, M1

and Ph northward.

Saraca declinata (Jack) Miq. 98, S. Lowland and hill forest, often riverine.

Widespread.

Sindora echinocalyx (Benth.) Prain. 98, S. Hill forest to 700 m. Widespread.

Loganiaceae

Fagrea elliptica Roxb. 98. Montane forest. Kd, Ph, Tg.

F. racemosa Jack ex Wall. 98, S. Secondary forests. Widespread.

*F. wallichiana Benth. 98. Hill ridge forest. Kd, Pn, Kl, Tg, Jh.

Flora of Bukit Bauk 293

Magnoliaceae

Magnolia candolii (Blume) H. Keng var. obovata (Kort.) Noot. AB.

Lowland and hill forest. Kd, KI, Tg, Pk, Jh.

Meliaceae

Aglaia exstipulata (Griff.) W. Theob. AB. Lowland and hill forest to 1400

im Kd. Pu, Ki, te, Pk, Ph, Sl, NS, Ml, Jh, Sp.

A. lawii (Wight) Saldanha ex Ramamoorthy. FR. Lowland forest. Kl, Pk,

SL, Tg:

A. leucophylla King. 98. Lowland and hill forest to 700 m. Kd, Kl, Tg, Pk,

Si NS, Jh;

A. macrocarpa (Miq.) Dannell. AB. Lowland to montane forest at 1750 m.

Kd, Pa, Ki ie, Ph, Pk, SI,.NS, Jh, Sp.

A. odoratissima Blume. AB. Lowland and hill forest to 1370 m. Throughout.

A. spectabilis (Miq.) S.S. Jain & Bennet. AB. Lowland forest to 300 m. Tg,

Pk) PhySl, NS; Mi. Jh,Sp.

Chisocheton sp. AB.

Dysoxylum cauliflorum Hiern. AB, FR. Lowland and hill forest.

Throughout.

D. dumosum King. AB. Lowland and hill forest to 1000 m. Tg, Pk, Ph, Sl,

Sh.

Sandoricum koetjape (Burm.f.) Merr. 98, S. Lowland and hill forest.

Widespread.

Melastomataceae

Dissochaeta gracilis (Jack) Blume. 98. Forest margins. Common throughout.

Melastoma malabathricum L. 98. Secondary growth in lowlands and hills.

Throughout.

Memecylon dichotomum (C.B. Clarke) King var. dichotomum. AB. Pk,

Tg.

M. excelsum Blume. 98. Lowland and hill forest to 1200 m. Kd, Kl, Tg, Pk,

Eh, NS, Mi. dh.

M. garcinoides Blume. AB. Lowland and hill forest to 1200 m. KI, Pk, Ph,

Si NS, Mi Jh, Tg, Sp.

M. megacarpum Furtado. 98, S. Lowland to montane forest at 1800 m. Kd,

Po, Ki, Te, Pk, Ph, Sl, Ml, NS, Jh, Sp.

M. minutiflorum Mig. 98, S. Lowland to montane forest at 1800 m. Kd, Pn,

Tg, Pk, Ph, Sl, NS, M1, Jh, Sp.

M. oleifolium Blume. AB. Lowland and hill forest to 1200 m. Kd, K], Tg,

Bk Pit. sl, NS. .ML Jh, Sp.

M. pubescens (C.B. Clarke) King. AB. Lowland and hill forest to 1200 m.

Kd, Tg, Pk, Ph, Sl, NS, MI, Jh, Sp.

294 Gard. Bull. Singapore 51 (1999)

Pternandra coerulescens Jack. AB, 98, S. Lowland and hill forest. Kd, Pn,

Kl, Tg, Pk, Ph, Sl, NS, Ml, Jh, Sp.

P. echinata Jack. 98. Lowland and hill forest to 1200 m. Kd, Pn, KI, Tg, Pk,

Ph, Sl, NS, MI, Jh, Sp.

Sonerila integrifolia Stapf. RK. Lowland to montane forest. Common on

the Main Range.

Monimiaceae

Kibara coriacea (Blume) Tul. AB, 98. Lowland and hill forest to 1000 m.

Pn, Tg, Pk, Ph, Sl, NS, MI, Jh, Sp.

Moraceae

Artocarpus elasticus Reinw. ex Blume. 98., lowland forest and open country.

Common throughout.

A. fulvicortex F.M. Jarrett. 98. Lowland forest. Pk, Ph, NS, Ml, Tg, Sp.

*A. hispidus F.M. Jarrett. AB. Lowland forest. Pn, Tg, Pk, Ph, Sl, Ml, Sp.

A. integer (Thunb.) Merr. var. silvestris Corner. AB, 98, S. Lowland to

montane forest. Widespread.

A. lanceifolius Roxb. 98. Lowland and hill forest. Widespread.

A. lowii King. 98. Lowland forest. Widespread but uncommon.

A. nitidus Trecul ssp. griffithii King. 98. Lowland forest. Throughout.

A. scortechinii King. 98. Lowland forest. Widespread.

Ficus aurata Mig. var aurata. FR, 98. Secondary forest. Kl, Tg, Pk, Ph, Sl,

Ml, Jh, Sp. |

F. deltoidea Jack var. deltoidea. 98. Seashores to mountain tops.

F. fulva Reinw. ex Blume. 98. Secondary forests in lowlands and hills. Ml

northwards.

F. microcarpa L.f. 98. Swampy places. Common and widespread.

F. parietalis Blume. 98. Lowland forest. Widespread.

Paratocarpus bracteatus (King) Becc. 98. Lowland forest to 600 m. Pn, Sl,

NS, MI, Jh, Tg, Sp.

Streblus elongatus (Miq.) Corner. 98. Lowland forest. Kd, Pn, Tg, Pk, Ph,

SI, NS, Jh, Sp.

S. taxoides (K. Heyne) Kurz. 98. Lowland, hills also on limestone.

Widespread.

Myricaceae

Myrica esculenta Buch.-Ham. 98. Lowland forest, particularly secondary

forest on poor soil, mountain tops. Widespread, commoner in the south.

Myristicaceae

Gymunacranthera farquhariana (Hook.f. & Thomson) Ward var. eugeniifolia

Flora of Bukit Bauk 295

(A.DC.) R.T.A. Scouten. AB. Lowland and hill forest to 1300 m. P.

Langkawi, Kd, Pn, Tg, Pk, Ph, SI, M1, Jh, Sp.

G. forbesii (King) Warb. 98. Lowland forest. Pn, KI, Tg, Pk, Ph, Sl, NS, M1,

Jh, Sp.

Horsfieldia ridleyana (King) Warb. 98. Lowland forest on poor soils. K1,

Tg, Pk, Ph, Sl, MI.

H. sucosa (King) Warb. AB, 98. Lowland forest. Kl, Tg, Pk, Ph, Sl, M1, Jh,

Tg, Sp.

H. superba (Hook.f. & Thomson) Warb. 98. Lowland forest. Common

throughout.

Knema conferta (King) Warb. AB. Swampy lowland forest. Pk, Ph, MI, Jh,

Sp.

K. curtisii (King) Warb. S. Lowland forest. Pn, KI, Tg, Pk, Ph, Sl, NS, MI,

Sh.

K. furfuracea (Hook.f. & Thomson) Warb. AB, 98, S. Lowland forest. P.

Langkawi, Kd, Pn, KI, Tg, Pk, Ph, Sl, NS, Ml, Jh, Sp.

K. hookeriana (Wall. ex Hook.f. & Thomson) Warb. AB, 98. Lowland and

hiisrorest; Pn) Ki, Te) Pk, Ph, NS»Mb dh; Sp.

K. kunstleri (King) Warb. AB. Lowland and hill forest on poor soils. K],

Tg; Pk Ph; Si, NS, Jh.

K. laurina (Blume) Warb. var. laurina. AB. Lowland forest. P. Langkawi,

Kd, Te;:Pk) Ph,/Si, NS; Ml, Jh, Sp.

K. malayana Warb. AB, 98. Lowland forest. Kd, Kl, Tg, Pk, Ph, Sl, NS, M1,

jh, Sp:

K. patentinervia (J. Sinclair) W.J. de Wilde. 98, S. Lowland forest. Kl, Tg,

Pk, Ph, Ml, Jh.

*K. plumulosa J. Sinclair. AB. Swampy lowland forest. Pn, Tg, Pk, Sl, Jh.

K. scortechinii (King) J. Sinclair. 98. Lowland forest. Kd, Kl, Tg, Pk, Ph, SI,

NS, Ml, Jh.

Myristica cinnamomea King. 98. lowland and hill forest, throughout.

M. elliptica Hook.f. & Thomson. AB. Swampy lowland forest. Throughout.

M. maxima Warb. AB. Lowland forest. Commonest in the south.

M. maingayi Hook.f. AB. lowland forest. Common throughout.

Myrsinaceae

*Ardisia mystica B.C. Stone. FR, 98. Montane forest. Ph (Genting

Highlands), Tg.

*A. perakensis King & Gamble. 98. Lowland forest. Pk, Tg.

*A. rosea King & Gamble var 2. 98. Hill and montane forest above 800 m.

Pk, Phy SL Ye.

*A. solida B.C. Stone. AB, FR. Lower montane forest at 1000 m. Jh

(Gunung Besar), Tg.

296 Gard. Bull. Singapore 51 (1999)

A. villosa Roxb. var. villosa. AB, FR. Lowland forest. Kd, Pn, Pk, Ph, Tg.

Grenacheria amentacea (G.B. Clarke) Mez. 98. Lowland forest. Widespread.

Labisia pumila (Blume) Fern.-Vill var. pumila. RK. Lowland and hill forest.

Widespread.

Maesa ramentacea Wall. ex Roxb. 98. Lowland and hill forest to 1200 m.

Throughout.

Rapanea porteriana Wall ex A.DC. AB. Coasts, lowland, hill and montane

forests to 1700 m. Throughout.

Myrtaceae

Acmena acuminatissima (Blume) Merr. & L.M. Perry. 98. Lowland to

montane forest. Widespread.

Cleistocalyx nervosum (DC.) Kosterm. AB. Lowland forest, often near

streams) KuyPieiee Pk, Ph, Si, J.

Rhodamnia cinerea Jack. AB, 98. lowland secondary forest, often near the

sea. Common throughout.

Rhodomyrtus tomentosa (Aiton) Hussk. 98. Open sandy ground. Common

throughout, particularly on the east coast.

Syzygium attenuatum (Miq.) Merr. & L.M.Perry ssp. attenuatum var.

attenuatum. 98. Lowland to lower montane forest. Kd, Pn, Ph, Sl, M1, Jh,

RenSp:

S. cerasiforme (Blume) Merr. & L.M. Perry. AB. Lowland forest. P.

Langkawi to MI.

S. chloranthum (Duthie) Merr. & L.M. Perry. 98. Lowland and hill forest.

Kd to Sp.

S. claviflorum (Roxb.) Wall ex A.M. Cowan & Cowan var. claviflorum. 98.

Lowland and hill forest. Ps, P. Langkawi, Kd, Tg, Pn, Pk, Ph, Sp.

S. cinereum (Kurz) P. Chantaranothai & J. Parn. AB. Lowland forest,

sometimes in mountains. Widespread.

S. dyerianum (King) P. Chataranothai & J. Parn. AB. Lowland to lower

montane forest at 1300 m. Common throughout.

S. flosculiferum (M.R. Hend.) Sreek. AB. Lowland and hill forest. Kd, Pk,

Ph, Sl, Mi, thpmenSp:

S. glaucum (King) P. Chantaranothai & J. Parn. AB, 98. Lowland forest.

Kd) Pay Pig Pi, SNS, Mi Jh, Tg, Sp.

S. gratum (Wight) S.N. Mitra. AB, 98., mostly on rocky and sandy shore.

Widespread.

S. griffithii (Duthie) Merr. & L.M. Perry. AB, 98. Lowland and hill forest.

Kd to Sp.

*§. goodenovii (King) Masam. AB. Lowland forest. Pk, NS, Ml, Tg.

S. kiahii (M.R. Hend.) I.M. Turner var. kiahii. AB. Lowland forest including

swamps. Tg, Ph, Sl, Jh.

eT —

Flora of Bukit Bauk 200

S. lineatum (DC.) Merr. & L.M. Perry. 98. Secondary forests and open

country in lowlands. Common throughout.

S. nigicans (King) Merr. & L.M. Perry. AB. Lowland and hill forest. Kd,

Pa. Kl tie Pk Pia SL. NS; MI, Jh, Sp.

S. polyanthum (Wight) Walp. var. polyanthum. AB. Lowland forest. P.

Langkawi and KI to Sp.

S. rugosum Korth. var. rugosum AB. Lowland and hill forest. Ps, Kd, Tg,

Pk, Ph, Sl, Ml, Jh, Sp.

S. syzygioides (Miq.) Merr. L.M. Perry. AB. Lowland, often near sandy

coasts. Common throughout.

S. valdevenosum (Duthie) Merr. & L.M. Perry. AB. Lowland and hill

forest. Pn to Jh.

*Suepv 22;. AB. Lowland forest. Tg, Jh,

*§. sp. 57. 98 . Lowland forest. Sl] (once from Sg. Lallang FR), Tg.

*§. sp 59. 98. Tree to 18m tall, once from Tg,

Sap SS.

Tristaniopsis merguensis (Griff.) Peter G. Wilson & J. T. Westerh. AB, 98.

Rocky coasts and mountains. Widespread.

Nepenthaceae

Nepenthes ampullaria Jack. 98. Secondary forest, forest margins. Pn, Pk,

Ml, Jh, Tg, Sp.

N. gracilis Korth. FR, 98. Secondary forest, forest margins in the lowlands.

Key Po; Tieg.Pk Ph, NS, MI, Jh, Sp.

Ochnaceae

Brackenridgea hookeri (Planch.) A. Gray. 98, S. Lowland and hill forest to

1000 m. Throughout.

Sauvagesia serrata (Korth.) Sastre. 98. Damp shady places in lowland and

hill forests. Tg, Ph, Jh.

Olacaceae

Ochanostachys amentacea Mast. S. Lowland and hill forest to 900 m. Kd,

Pay kkt le. Pk. Ph, Si, NS, Ml, Jh, Sp.

Scorodocarp.s borneensis (Baill.) Becc. AB, 98. Lowland forest. Kd, Pn,

Kl, Tg, Pk, Ph, Sl, NS, MI, Jh, Sp.

Strombosia ceylanica Gardn. 98. Widespread.

S. javanica Blume. 98. Lowland forest. Kd, Pn, Kl, Pk, Ph, Sl, NS, Jh, Tg,

Sp:

Opiliaceae

Champereia manillana (Blume) Merr. AB. Lowland forest. Widespread.

298 Gard. Bull. Singapore 51 (1999)

Oxalidaceae

*Sarcotheca glomerula Veldkamp. AB. Lowland forest. Kd, Pk, Ph, Tg.

S. griffithii (Planch ex Hook.f.) Hallier f. S. Lowland forest. Ps, Kd, Pn, Tg,

Pk) Ph, Si, NS, Midas Sp.

S. laxa (Ridl.) Knuth var. */axa. 98. Swamps or forest margins. Ps, Kd, KI],

ibe; Pk)

S. laxa (Ridl.) Knuth var. *sericea (Ridl.) Veldkamp. AB, 98. Swamps and

forest margins. Tg, Ph, Jh, Sp.

Pandacaceae

Galearia fulva (Tul.) Mig. 98, S. Lowland and hill forest to 690 m.

Throughout.

Passifloraceae

Paropsia vareciformis (Griff.) Mast. AB, 98. Lowland and hill forest to 500

m. Ts, Ph: Pk SiS; M1.

Pentaphragmataceae

Pentaphragma horsfieldii (Mig.) Airy Shaw. RK. Lowland and montane

forest to 2000 m. Pk, Ph, SI, Ml, Tg.

Piperaceae

Piper porphyrophyllum N.E. Br. 98. Lowland forest. Widespread.

Polygalacaeae

Xanthophyllum affine Korth. ex Mig. AB, 98, S. Lowland and mountains.

Widespread.

X. griffithii Hook.f. ex A.W. Benn. ssp. erectum Meijden. 98. Lower montane

forest to 1400 m. Pk, Sl, NS, MI, Tg.

X. ngli Meijden. AB. Lowland forest. Tg, NS, MI.

X. obscurum A. W. Bennett. AB. Lowland forest. Kd, Pn, Kl, Tg, Pk, Ph,

SI, NS, MI, Jh, Sp.

Rhamnaceae

Ventilago malaccensis Ridl. AB. Lowland forest. Widespread.

Rhizophoraceae

Gynotroches axillaris Blume. 98, S. Lowland including swamps to mountains

at 1400 m. Throughout.

Rosaceae

Prunus arborea (Blume) Kalkman var. arborea. AB. Lowland to lower

Flora of Bukit Bauk 299

montane forest at 1300 m. Pk, SI, NS, MI, Tg, Sp.

P. grisea (Blume) Kalkman var. tomentosa (Koord. & Valeton) Kalkman.

98. Lowland forest to 600 m. Widespread.

P. javanica (Teij S.M. & Binn.) Mig. 98. Lowland forest. Kd, Pk, Ph, Tg.

Rubus moluccanus L. var. moluccanus. 98. Forest margins in lowlands and

hills. P. Langkawi, Pn, Tg, Pk, Ph, Sl, NS, Sp.

Rubiaceae

Aidia densiflora (Wall.) Masam. AB, 98. Lowland to montane forest.

Throughout.

*Canthium sp. 1. AB. Lowland forest. Pk, Ph, Sl, Tg.

C. strychnoides Craib. AB, FR. Lowland to lower montane forest. Tg, KI.

Gaertnera vaginans (DC.) Merr ssp. junghuhniana (Miq.) Beusekom. FR.

Lowland to montane forest. Throughout.

Gardenia tubifera Wall. var. tubifera. AB, 98, S. Lowland and hill forest.

Kid Kb Te ePk Phy Si NS MI, Jh, Sp.

G. tubifera Wall. var. subcarinata Corner. AB. Lowland and hill forest. Kd,

Ki Te. Poa, Pk, Ph, Sl Mil; dh, Sp.

Greenea corymbosa (Jack) K. Schum. 98. Rocky coasts to hill ridges to 700

m. Widespread.

Hedyotis philippinensis Willd. ex Spreng. 98. Lowland and hill forests.

Widespread.

*Hypobathrum venulosum (Hook.f.) K.M. Wong. AB. Lowland and hill

forest. Widespread.

Ixora congesta Roxb. FR, 98, S. Lowland and hill forest to 1000 m.

Throughout.

I. grandifolia Zoll. & Moritzi var. grandifolia. FR. Lowland and hill forest.

Throughout.

I. javanica (Blume) D.C. var. javanica. FR. Lowland and hill forest.

Widespread.

I. lobbit King & Gamble. S. Lowland to montane forest at 1200 m.

Throughout.

I, pendula Jack var. pendula. 98. Lowland and hill forest to 600 m.

Throughout.

I. umbellata Koord. & Valeton var. umbellata. 98. Lowland and hill forest,

also on limestone. Throughout.

*Kochummenia parviflora K.M. Wong. FR. Collected twice from Tg.

Lasianthus densifolius Mig. 98. Lowland and hill forest. Kd, Tg, Pk, Ph, Sl,

NS, MI, Jh, Sp.

L. griffithii Wight. AB, FR. Lowland and hill forest. Kl, Tg, Pk, Ph, Sl, NS,

Ml, Jh, Sp.

Lasianthus sp. RK.

300 Gard. Bull. Singapore 51 (1999)

Morinda elliptica (Hook.f.) Ridl. AB, FR, 98. Lowland secondary forest

and forest margins. Widespread, common.

Nauclea officinalis (Pierre ex Pit.) Merr. & Chun. AB. Lowland and hill

forest. Throughout.

Neonauclea pallida (Reinw. ex Havil) Bakh.f. ssp. malaccensis (Gand.)

Ridsdale. AB. Lowland to montane forest, often rheophyte. Widespread.

Ochreinauclea maingayi (Hook.f.) Ridsdale. AB. Tg, Pn, Pk, Ph, Sl, NS,

Kl, Jh, Sp.

Porterandia anisophylla (Jack ex Roxb.) Ridl. AB, FR, 98. Lowland and

‘hilkforestitd, Pu, Te, Pk, Ph, Si, NS, Mi, th. Sp:

Prismatomeris glabra (Korth.) Valeton. FR, 98. Lowland to montane forest.

Throughout.

Psychotria malayana Jack. AB, FR. Lowland to montane forest at 1600 m.

All states except Ps and Kd.

*P. penangiana Hook.f. FR. Lowland forest, common. Widespread,

P. viridiflora Reinw. ex Blume. AB, FR. Lowland forest also on limestone.

Throughout.

Rothmannia macrophylla (R.Br. ex Hook.f.) Bremek. AB. Lowland and

hill forest. Kd, Kl, Tg, Pn, Pk, Ph, Sl, NS, M1, Jh, Sp.

Tarenna fragrans (Nees) Koord. & Valeton. 98. Lowland forest, often on

river banks. Kd, Pn, Pk, Ph, Sl, NS, MI, Jh, Tg, Sp.

Timonius flavescens (Jack) Baker. 98, S. Peaty soils in the lowlands and

mountains. Throughout.

*T. wrayi King & Gamble. AB, FR. Lowland and hill forest. Tg, Pk, Ph,

Jh.

Uncaria cordata (Lour.) Merr var. cordata. AB, 98. Lowland forest.

Widespread.

Urophyllum glabrum Wall. FR, 98, S. Lowland to lower montane forest at

1400 m. Throughout.

U. griffithianum (Wight) Hook.f. FR. Lowland and hill forest. Pn, Tg, Pk,

Ph, ‘Sl, NSMibsihe Sp:

U. streptopodium Wall. ex Hook.f. FR. lowland to lower montane forest at

1200:m. Po, Kip Tei Pk, ‘Ph; Sls NS;M1, ah, Sp:

Rutaceae

Glycosmis chlorosperma Spreng. var. chlorosperma. 98. Lowland to montane

forest. Widespread.

G. decipiens B.C. Stone. AB. Lowland to lower montane forest. Tg, Ph,

P.Tioman, NS, Jh.

Maclurodendron porteri (Hook.f.) T.G. Hartley. AB, 98. Lowland to lower

montane forest. Throughout.

Flora of Bukit Bauk 301

Sapindaceae

*Glenniea penangensis (Ridl.) Leenh. AB. Lowland and hill forest to 1000

m. Widespread.

Guioa pleuropteris (Blume) Ridl. 98. Coastal and riverine forest.

Widespread.

Lepisanthes fruticosa (Roxb.) Leenh. AB, 98. Lowland forest, usually on

slopes. Kd, Pn, Kl, Pk, Ph, Tg, Sl, M1.

L. senegalensis (Poir.) Leenh. AB. Lowland and hill forest to 1000 m.

Widespread.

L. tetraphylla (Vahl.) Radlk. AB, 98, S. Lowland forest. Widespread.

Litchi chinensis Sonn. 98. Hill forest. Kd, Tg, Pk, Sl, NS.

*Nephelium costatum Hiern. S. lowland forest, Pk, Ph, Tg, Sl, NS, MI.

N. cuspidatum Blume var. ophiodes (Radlk.) Leenh. S. Lowland and hill

forest. Ph, NS, Jh, Tg.

N. maingayi Hiern. 98. Swampy lowland forest. Kl, Tg, Pk to Jh.

Xerospermum noronhianum (Blume) Blume. AB, 98, S. Lowland forest to

500 m. Widespread.

Sapotaceae

Ganua sp. AB.

Isonandra sp. AB.

* Madhuca. tomentosa H.J. Lam. AB, FR.. Swampy lowland forest. Ph, Tg,

Jh.

*M. tubulosa H.J. Lam. AB, TF, FR. Lowland forest. Tg, Jh.

M. utilis (Ridl.) H.J. Lam. AB, FR, 98, S. Lowland forest. Tg, Pk, Ph, Sl,

Jh.

Palaqium clarkeanum King & Gamble. AB, FR. Lowland forest, Kd, K1,

Tg, Pk, Ph, Sl, NS, Jh.

P. hexandrum (Griff.) Baill. AB, FR, 98. Mostly lowland forest. Kd, Pn,

Kl, Tg, Pk, Ph, Sl, NS, Ml, Jh, Sp.

P. leiocarpum Boerl. AB, FR. Lowland freshwater swamp forest. Kl, Tg.

*P. maingayi (C.B. Clarke) King & Gamble. 98. Lowland and hill crest.

no, KL Ts: Pk. Pho SL. NS, ML. Th.

P. obovatum (Griff.) Engl. AB, FR. Lowland forest. Kd, Pn, Kl, Tg, Ph, Sl,

NS, Ml, Jh, Sp.

P. rostratum (Miq.) Burck. AB, FR. Lowland to montane forest. Kd, Pn,

Kl, Tg, Pk, Ph, Sl, NS, MI, Jh, Sp.

P. semaram H.J. Lam. AB, FR. Lowland forest. Kl, Tg, Ph, Jh.

P. sukoei C.E.C. Fisch. FR. Lowland and hill forest. Tg, Pk, Jh.

P. xanthochymum (de Vriese) Pierre ex Burck. AB, FR. Swampy lowland

forest. Tg, Pk, Ph, Sl, Jh, Sp.

*Payena maingayi C.B. Clarke. AB, FR. Lowland forest. Kd, Pn, Tg, Pk,

302 Gard. Bull. Singapore 51 (1999)

Ph, Sl, NS, Ml, Jh, Sp.

Payena sp. A. TF, FR. Lowland forest. Tg (twice from Bukit Bauk).

Pouteria maingayi (C.B. Clarke) Baehni. AB, 98. Lowland and hill forest.

Throughout.

Schisandraceae

Kadsura scandens Blume. AB. lowland forest, widespread.

Simaroubaceae

Eurycoma longifolia Jack. 98. Lowland and hills. Throughout.

Quassia indica (Gaert.) Noot. S. Tidal swamps. Tg, Jh, Sp.

Sterculiaceae

Commersonia bartramia (L.) Merr. 98. Lowland disturbed sites. Widespread.

Heritiera javanica (Blume) Kosterm. 98, S. Lowland forest to 600 m. Kd,

Kl, Tg, Pk, Ph, Sl, NS, Jh.

H. simplicifolia (Mast.) Kosterm. S. Lowland forest. Kl, Tg, Pk, Ph, Sl, NS,

Ml, Jh, Sp.

Leptonychia caudata (wall. ex G. Don) Burret. 98, S. Lowland to lower

montane forest. Ps, Kd, Pn, Tg, Pk, Ph, Sl, MI, Jh, Sp.

Scaphium macropodum (Miq.) Beumee ex Heyne. 98, S. Lowland and hill

forest to 1200 m. Kd, Pn, K1, Tg, Pk, Ph, Sl, NS, M1, Jh, Sp.

Sterculia coccinea Jack. 98, S. Lowland forest. Common throughout.

S. cuspidata R.Br. AB. Lowland and hill forest. Kd, Pn, Tg, Pk, Ph, NS,

Ml, Jh.

Styracaceae

Styrax benzoin Dryand. var. hiliferum Steenis. AB. Lowland forest. Ph,

Tg.

Symplococaceae

Symplocos adenophylla Wall ex G. Don var. adenophylla. AB, 98. Sea

coasts to montane forest at 1500 m. Widespread.

S. barringtoniifolia Brand. AB. Lowland and hill forest to 750 m. Tg, Pk,

Ph, Sl, M1, Jh.

S. crassipes C.B. Clarke var. curtisii (Oliv.) Noot. AB. Lowland to montane

forest 300-1200 m. Tg, Pk, Ph, Sl.

S. rubiginosa Wall. ex DC. 98. Lowland to montane forest. Kd, Pn, Tg, Pk,

Ph, Sl, NS, M1, Jh, Sp.

Flora of Bukit Bauk 303

Theaceae

*Gordonia maingayi Dyer. AB, 98. Lowland to montane forest at 1600 m.

Kl, Tg, Pk, Ph, Sl, NS, MI.

*G. multinervis King. AB. Lowland forest below 800 m. Pn, Kl, Tg, Pk, Ph,

Ml, Jh, Sp.

*G. singaporeana Wall. ex Ridl. 98. Lowland forest. Pn, Pk, NS, Ph, NS,

Ml, Jh, Tg, Sp.

Pyrenaria acuminata Planch. AB. Lowland forest. Pn, Kl, Tg, Pk, Ph, Sl,

NS, Ml, Jh, Sp.

Tiliaceae

Microcos blattifolia Corner. AB. Lowland and hill forest to 1000 m.

Widespread.

*M. erythrocarpa (Ridl.) Airy Shaw. AB. Lowland forest to 600 m.

Widespread.

M. fibrocarpa (Mast.) Burret. S. Scattered widely.

M. lanceolata (Miq.) Burret. S. Lowland forest. Mostly west coast states.

*Pentace acuta Ridl. AB. Lowland forest. Tg, Pk.

P. floribunda King. 98. Hill forest. Kl, Pk, Ph, SI, Tg.

*P. grandiflora Kochummen. AB, 98. Ridge top forest. Tg only.

P. macrophylla King. AB. Lowland and hill forest. Tg, Pk, Sl, Jh.

*P. strychnoidea King. AB, 98. Lowland and hill forest. Kd, Kl, Tg, Pk, Ph,

SI, NS.

P. triptera Mast. AB, 98. Lowland forest. Kd, Pn, Kl, Tg, Pk, Ph, Sl, NS,

MI, Jh, Sp.

Schoutenia accrescens (Mast.) C.H. Curtis ssp. accresens. AB, 98. lowland

to montane forest. Common and widespread.

S. accrescens (Mast.) C.H. Curtis ssp. borneensis Roekmowati. 98. Lowland

forest. Tg, Jh.

Triumphetta grandidens Hance. 98. Sandy shores. East coast.

Thymelacaceae

Aquilaria hirta Ridl. AB. Lowland forest. Tg, Ph, Jh, Sp.

A. malaccensis Lam. 98, S. Lowland and hill forest to 750m. Kd, Pn, KI, Tg,

Pk, Ph, St, NS, MI, Ua, Sp.

Gonystylus brunnescens Airy Shaw. AB. Lowland forest, often near sea.

Pn, Tg, Pk,.Ph.

Ulmaceae

Gironniera hirta Ridl. 98. Lowland forest. Kl, Ph, Jh, Tg, Sp.

G. parvifolia Planch. AB, 98, S. Lowland and hill forest to 900 m. Kd, Pn,

KI 9, Pk, Ph, Sl, NS, Mil, Jh, Sp.

304 Gard. Bull. Singapore 51 (1999)

G. subaequalis Planch. S. Lowland and hill forest to 1100 m. Kd, Pn, KI],

Te, Pk, Ph,iSl NS; Bieta, Sp.

Verbenaceae

Clerodendron deflexum Wall. AB. Lowland to lower montane forest at

1300 m. Throughout.

Gmelina elliptica Sm. AB. Open country near the sea. Widespread.

Premna serratifolia L. 98. Sea shore. Common on all coasts.

Stachytarpheta indica (L.) Vahl. 98. Sandy places near the sea. Widespread

weed.

Teijsmanniodendron simplicifolilum Merr. TF. Lowland forest. Tg, Ph, Pk.

Vitex gamosepala Griff. 98. Lowland and hill forest to 1200 m. Throughout.

V. pinnata L . 98. Disturbed forest. Throughout.

*V. longisepala King & Gamble. 98. Mainly montane forest. Pn to MI.

Violaceae

Rinorea anguifera (Lour.) Kuntze. 98, S. Lowland and hill forest to 600 m.

Throughout.

R. lanceolata (Wall.) Kuntze var. lanceolata. AB. Lowland forest below

750 m. Widespread mostly in the south.

Vitaceae

Ampelocissus sp. AB.

MONOCOTYLEDONAE

Araceae

Aglaonema nitidum (Jack) Kunth. RK. Damp lowland forest. Widespread.

Alocasia denudata Endl. var. denudata. RK. Forest margins. Widespread.

Homalomena aromatica (Roxburgh) Schott. 98. Widespread.

Rhapidophora lobbii Schott. 98. Lowland forest. Pn, Sl, Ml, Tg, Sp.

Commelinaceae

Amischotolype griffithii (C.B. Clarke) I.M. Turner. 98. Lowland forest.

Widespread.

Cyperaceae

Mapania cuspidata (Miq.) Uittien var. cuspidata. 98. Primary lowland forest.

Kl, Pk, Ph, $i Ju, Fe;

Flora of Bukit Bauk 305

Dracaenaceae

Dracaena conferta Ridl. 98. Lowland and hill forest. Widespread.

*D. maingayi Hook.f. 98. Lowlands near the sea. Mostly in the south.

Gramineae

Themeda villosa (Poir) A. Camus. 98. Sandy waste places, river banks,

Hanguanaceae

Hanguana malayana (Jack) Merr. 98. Lowland and hill forest to 1500m, or

in lowland ponds and rivers. Widespread.

Hypoxidaceae

Molineria latifolia (Dryand.) Herb. ex Kurz var. latifolia. 98. Forest and

forest margins, lowland to mountain tops. Throughout.

Palmae

*Areca ridleyana Furtado. AB, FR. Hill forest. Tg, Ph, Jh.

Arenga hastata (Becc.) Whitmore. AB, FR. Lowland forest sometimes on

limestone.,T¢, Pk» Ph, Sl,.Jh-

A. westerhoutii Griff. 98. Locally gregarious in hillside forest including

limestone. Widespread.

A. hookeriana (Becc.) Whitmore. 98. Lowland forest including limestone.

Widespread.

Calamus blumei Becc. FR. Swampy or wet forest to 800 m. Pk, Ph, Sl, NS,

Tg, Jh.

C. diepenhorstii Mig. 98. Lowland and hill forest to 700 m. KI, Tg, Pn, Pk,

Ph, Sl, NS, M1, Jh, Sp.

*C. holttumii Furtado. FR. Hill forest. Tg (Bukit Bauk), Jh.

C. longispathus Ridl. FR, 98. Ridge tops in hill forests. Tg, Pk, Ph, Sl, NS,

Ml, Jh.

C. polystachys Becc. 98. Ph (Mentakab), P. Tioman, Tg.

C. scabridulus Becc. FR. Peat swamp forest. Tg, Sl, Jh.

Daemonorops longipes (Griff.) Martelli. FR. Damp spots in lowland forest.

Ph, NS, Ml, Jh, Tg, Sp.

*D. monticola (Griff.) Martelli. 98. Hill forest, Pn, Jh, Tg.

D. sabut Becc. FR. Alluvial or frestwater swamp forest to 250 m. Tg, Pk,

Ph, NS, Jh, Sp.

Iguanura wallichiana (Wall. ex Martelli) Hook.f. ssp. wallichiana var.

wallichiana. 98. Lowland and hill forest to 1000 m. North of Fraser’s Hill,

Ph, and Sg. Dungun, T¢.

306 Gard. Bull. Singapore 51 (1999)

Korthalsia sp. 98.

*Licuala ahlidurii L.G. Saw. 98. Lowland forest, Tg (only Bukit Bauk).

*T.. bayana L.G. Saw. 98. Tg only.

L. ferruginea Becc. AB, 98. Lowland forest. Tg, Ph, NS, Jh, Sp.

*L. fractiflexa L.G. Saw. 98. Tg only.

L. glabra Griff. var. glabra. AB, FR, 98. Lowland and mountains. KI], Pk,

Ph, S1):Te)Mi

*T.. khoonmengii L.G. Saw. AB, 98. Tg only.

*[T_. malajana Becc. 98. Lowland and hill forest. Kl, Tg, Ph, Sl, Jh.

L. spinosa Wurmb. AB, 98. Damp open sites. Throughout.

L. terengganuensis L.G. Saw. AB, 98 Tg only.

*Livistona endauensis J. Dransf. & K.M. Wong. FR, 98. Gregarious in hill

forest to 660 m. Tg, Jh.

Johannesteijmannia altifrons (Rchb.f. & Zoll) H.E. Moore. 98. Lowland

forest. Kl, Ph, Sl, Tg, Jh.

Oncosperma horridum. (Griff.) Scheff. 98. Lowland and hill forest to 500

m.Throughout.

Pinanga disticha (Roxb.) Blume ex H. Wendl. 98, S. Lowland and montane

forest to 1200 m. Widespread and common.

P. simplicifrons. AB, FR. Wet lowland forest. East of Main Range.

Plectocomia elongata Mart. ex Blume. 98. Lowland and montane forest to

2000 m. Kd, KI, Tg, Ph, Pn, Pk, Sl, NS, Ml, Jh, Sp.

Pholidocarpus macrocarpus Becc. 98. Swampy lowland forest. Widespread.

Lowiaceae

*Orchidantha ? fimbriata Holttum. 98, S. Lowland and hill forest. Tg, Pk,

Ph, Sl, Ju.

Marantaceae

Donax grandis (Miq.) Ridl. 98. Lowland forest. Widespread.

Orchidaceae

Bromheadia finlaysoniana (Lindl.) Miq. 98. Lowland secondary forest and

scrub. Common throughout.

Claderia viridiflora Hook.f. 98. Lowland and hill forest. Widespread.

Smilacaceae

Smilax setosa Mig. 98. Lowland forest margins. Widespread.

S. calophylla Wall. ex A. DC. 98. Lowland and hill forest. Widespread.

Flora of Bukit Bauk 307

Taccaceae

Tacca integrifolia Ker Gawl. 98. Lowland and hill forest. Widespread.

Zingiberaceae

Alpinia javanica Blume var. javanica. AB, 98. Open places in lowland

forest. Widespread.

*Globba unifolia Ridl var. sessifolia. 98. Lowland and montane forest. K],

Tg.

*Scaphochlamys rubromaculata Holttum. 98. Lowland forest at 200 m.Tg

(Ulu Bendong and Bukit Bauk).

GYMNOSPERMS

Cyacadaceae

Cycas rumphii Miq. 98. Mostly rocky shores. Widespread.

Gnetaceae

Gnetum gnemon L. var. gnemon. 98, S. Rocky coasts and islands. Mostly

east coast.

PTERIDOPHYTA

Adiantaceae

Taenitis blechnoides (Willd.) Sw. 98. Common at forest edges. Widespread.

Aspleniaceae

Asplenium nidus L. 98. Lowland and mountains. Throughout.

Blechnaceae

Blechnum finslaysonianum Wall. ex Hook.f. & Grew. 98. Forests to 500 m.

Widespread.

B. orientale L. 98. Common in open sites, lowland and mountains.

Widespread.

Dipteridaceae

Dipteris lobbina (Hook. & Grew.) Copel. 98. Rheophyte, lowland and hills

to 1000 m.

Dryopteridaceae

Tectaria semipinnata (Roxb.) C.V. Morton. 98. Lowland forest.

308 Gard. Bull. Singapore 51 (1999)

T. singaporeana (Hook. & Grew.) Copel. 98. Lowland forest.

Gleicheniaceae

Dicranopteris linearis (Burm.f.) Underw. var. linearis. 98. Lowland and

mountains to 1400 m. Throughout.

Sticherus truncatus (Willd.) Nakai var. truncatus. 98. Forest margins to

1600 m. Widespread.

Lycopodiaceae

Lycopodium sp. 98.

Polypodiaceae

Platycerium coronarium (D. Koenig ex O.F. Mull) Desv. 98. Lowland forest

and plantations. Widespread.

Schizaeaceae

Lygodium circinnatum (Burm.f) Sw. 98. Lowland to montane forest at

1500 m.

L. longifolium (Willd.) Sw. 98. Lowland forest margins.

Lygodium sp. 98. |

Schizaea dichotoma (L.) J. Sm. 98. Lightly shaded sandy soils to 1000 m.

Selaginellaceae

Selaginella sp. 98.

Thelypteridaceae

Pronephrium repandrum (Fee) Holttum. 98. Shady lowland and hill forest.

Gardens’ Bulletin Singapore 51 (1999) 309-317.

The Importance to Indopacific Botany

of Baron Dumont de Courset’s Botaniste Cultivateur

D. J: MABBERLEY

National Herbarium of The Netherlands, University of Leiden;

Royal Botanic Gardens Sydney,

Mrs Macquaries Road, Sydney 2000, Australia’

Abstract

An examination of the second edition of Dumont de Courset’s Botaniste Cultivateur

completes the listings of new taxa in the Australian Plant Name Index and not yet in Index

kewensis, where most are already found. Besides correction of the authority of the Chinese

Caragana pygmaea (Leguminosae) to ‘(L.) Dum. Cours.’ (and the North African Genista

ferox to ‘(Poir.) Dum. Cours.’), and restoration of the Malesian Pipturus asper Wedd.

(Urticaceae, lately referred to P. arborescens C. B. Robinson), and the superseding of the

Himalayan Rubus fragarioides Bertol. (Rosaceae, non (Michaux) Dum. Cours.) by R.

franchetianus Lévl., Dillenia crenata (A.C. Sm.) Hoogl. (Dilleniaceae, non (Andr.) Dum.

Cours.) of the Solomon Islands is here renamed Dillenia crenatifolia Hoogl. ex Mabb.,

nom. nov..

Introduction

In early nineteenth-century Europe, there was, following that for Cape

plants, a vogue for Australian ones, fuelled by contemporary British

voyages, particularly that of Flinders on the /nvestigator (1801-5), and also

those of the French, notably that of Baudin (1754-1803). Both also made

collections in present-day Indonesia. The French, particularly those

collecting for Josephine Bonaparte, were as quick as the British to get the

new plants into cultivation. With improved glasshouses and heating systems

it was possible to grow successfully more plants from the tropics, including,

for example, Josephinia imperiatricis Vent. (Pedaliaceae; see Heine, 1967),

named after Josephine and figured by Redouté in Ventenat’s Jardin de la

Malmaison (1804).

On the British side, most of these early introductions were

incorporated in the second edition of Hortus kewensis (1811-1813),

attributed to W.T. Aiton but partly written and edited by no less than

Robert Brown (1773-1858), who had been naturalist on the voyage of the

Investigator. Like the first edition of Hortus kewensis, the second one was

arranged according to the sexual system of Linnaeus, which, paradoxically,

' address for correspondence

310 Gard. Bull. Singapore 51 (1999)

Brown himself has much of the credit for ousting when re-introducing the

Natural System of classification. This was in his Flora Novae Hollandiae et

Insula Van Diemen (1810), the first, though incomplete, Flora of Australia,

and a book in which many common Malesian plants were first described.

Brown was effectively carrying the system of de Jussieu to British readers.

Not surprisingly, the French equivalent of Hortus kewensis, Baron Georges

Louis Marie Dumont de Courset’s Botaniste Cultivateur (ed. 1, 5 vols,

1802-5 — for sale in Paris, London and Haarlem, with a later Leipzig

edition in German) and ed. 2 (1811-4) almost exactly contemporaneous

with the second edition of Hortus kewensis, was arranged along Jussieuian

lines. That there is so little concordance between these two compendia

may be explained by their being written in isolation due to the Napoleonic

Wars.

Dumont de Courset (1764 — 1824) was born at the chateau de Courset

near Boulogne in northern France (Lair, 1814) and was destined for a

military career, but, whilst serving as a cavalry captain, was posted to the

Pyrenees, where the flora so captivated him that he turned to botany and

agriculture. Author of a number of works on agronomy, he set about

creating a private botanic garden at Courset, where, by 1814, he had some

3600 exotic perennial species in cultivation, extensive greenhouses complete

with the latest Australian plants as well as notable collections of Cape

heaths and pelargoniums. Widely respected for his skill and breadth of

knowledge, he was referred to as “ce nouveau Theophraste”. He died at

Courset. ,

His Botaniste Cultivateur was largely based on personal knowledge

of the living plants and the first six volumes of his second edition, which

appeared “vers la fin de 1811”, covers some 8700 species referred to 1400

genera. On top of this, a supplementary seventh volume was issued in

1814. Although Hortus kewensis dealt only with plants cultivated in the

London area, even the first edition of Botaniste Cultivateur claimed to

cover all plants in England as well as France, while the second edition had

in addition those of Italy and Austria (presumably in the old broad sense

and therefore covering many other present-day central European countries

too).

Australasia

The second edition deals with 520 plants from Australia alone (see vol. 7:

361) and contains hundreds of novelties from all over the world. Many

binomials published in the book are today’s accepted names of ornamental

and other economic plants, tropical as well as temperate, grown worldwide.

Botaniste Cultivateur 3t1

Strangely, although the new names in the early volumes of the book were

safely caught up in Index kewensis, many of those in the later ones — some

of them first published in the first edition — are not, though names in

Eucalyptus (vol. 7), for example, and those for many other plants (including

many based on sterile material), are. Some of the missing are listed in

Mabberley (1990, cf. also other neglected French work, notably that of

Chazelles, discussed in Mabberley, 1991), and some, referring to the Pacific,

are picked up in the synonymy of Araucaria (Abies colombaria, Pinus

colombaria) by Green (1994 : 543).

Many others are in the Australian Plant Name Index (1991) with a

suitable note, “not in Index Kewensis”. Some of these lead to ramifications

beyond Australia, e.g. Dillenia crenata (Andr.) Dum. Cours., 1.e. Hibbertia

grossulariifolia (Salisb.) Salisb. (Dilleniaceae), which according to the

catalogue appended to vol. 7 was being grown at Courset in 1814. The

name is an earlier homonym of D. crenata (A.C. Sm.) Hoogl., a therefore

illegitimate name for a tree from the Solomon Is. It is a pleasure to carry

out the wish of my late friend, Ruurd Hoogland (1922-1994), monographer

of the genus Dillenia and avid bibliographer, who had inadvertently created

the superfluous homonym and, as is apparent from an annotated sheet at

L, later realised his error:

Dillenia crenatifolia Hoogl. ex Mabb., nom. noy. — Dilleniaceae.

Wormia crenata A.C. Sm. in J. Arn. Arb. 22: 498 (1941). - Dillenia crenata

(A.C. Sm.) Hoogl., Blumea 7: 42 (1952), nom. illegit., non D. crenata

(Andr.) Dum. Cours., Bot. Cult., ed. 2, 7: 251 (1814) = Hibbertia

grossulariifolia (Salisb.) Salisb. (Dilleniaceae).

Type: Solomon Is., Santa Isabel., Tatamba, 4 Jan. 1933, Brass 3418 (A,

holo-, 1.V.; BMT, KT, 11).

Distribution: Solomon Islands.

It is not only Dumont de Courset’s Pacific plant-names that have

been omitted from Index kewensis but also those of many of plants from

other parts of the world, often the tropics. Those for Indomalesia so far

not picked up, plus, for completeness, those based on plants from the rest

of the world, are given in the Appendix and an attempt is made to dispose

of them there.

N.B. Several of Dumont de Courset’s new names were also published at

about the same time in the first part (23 Oct 1811) of Poiret’s supplement

vol. 2 to Lamarck’s Encyclopédie Méthodique, though according to Lair

(1814), Dumont de Courset’s book did not appear until late in the year, so

perhaps followed Poiret: e.g. Caragana jubata (Pallas) Poir. and Comocladia

brasiliastrum Poir. As Dumont’s herbarium is lost, those of his new species

312 Gard. Bull. Singapore 51 (1999)

described from sterile plants are not considered below, despite many such

being already in /ndex kewensis and/or Australian Plant Name Index.

Indomalesia

The only nomenclatural consequences are for Indopacific plants, though

two new combinations, in Caragana and Genista in current use, must

correctly be attributed to Dumont de Courset. The only adjustments

necessitated by setting the record straight in bringing all of Dumont de

Courset’s second edition into line with that followed for Australian plants

in the Australian Plant Name Index are the restoration of a name in Pipturus

(Urticaceae) and resurrection of a name (also inexplicably not yet in Index

kewensis) for a Himalayan Rubus (Rosaceae). A fortunate outcome is

that both of the correct ‘new’ names have extant holotype specimens,

unlike one, at least, of those they replace:

1. Pipturus asper Wedd. in Ann. Sci. Nat. 4, 1: 197 (1854). Type: Philippines,

Cuming 724 (P [‘Manille’], holo!; BM!, E!, G!, K!, L!, OXF!, iso) -

Urticaceae.

Urtica arborescens Link, Enum. Hort. Berol. 2: 386 (1822), nom. illegit.,

non Dum. Cours., Bot. Cult. 3 : 689 (1802) & ed. 2, 6: 370 (1811), Le.

Urera baccifera (L.) Wedd. (Urticaceae). - Pipturus arborescens C.B.

Robinson in Philip. J. Sci., Bot. 6: 13 (1911), nom. superfl., illegit.

Type: cultivated in Berlin from material sent from Manila, Philippines

(B, delet.).

Distribution : Ryukyus, Taiwan, Philippines, Borneo (teste Eichhorn MSS).

I am grateful to Henk den Bakker and Karl Eichhorn (L) for help in

clarifying the identities of these plants.

2. Rubus franchetianus Lévl. in Bull. Acad. Int. Géog. Bot. 20: 71 (1909);

Focke in Bibl. Bot. 19(83): 16 (1914). - R. fragarioides Bertol. var.

adenophora Franchet, Pl. Delavay.: 203 (1890); Lauener in Notes Roy.

Bot. Gard. Edinb. 30: 272 (1970). Type: Yunnan, HoKin, ‘Col de Koua-la-

po’, 3500m, 13 July 1886, Delavay 2160 (P!, holo; P!, iso) - Rosaceae.

Rubus fragarioides Bertol. in Mem. Accad. Sci. Ist. Bologna 12: 236, t. 5

(1861) & Misc. Bot. 22: 16, t. 5 (1862), nom. illegit., non R. fragarioides

(Michaux) Dum. Cours., Bot. Cult.,.ed. 2, 7: 293. (1814), 16.

Waldsteinia fragarioides (Michaux) Tratt. (Rosaceae) - R. arcticus L.

var. fragarioides Focke, Bibl. Bot. 17 (72): 24 (1910). Type: Sikkim,

‘11-12000 ped.’, J.D. Hooker s.n. in Herb. Hooker & Thomson

(holo-, BOLO, n.v.; BM!, E, K!, L!, P!).

Botaniste Cultivateur 313

Rubus fragarioides Bertol. var. pubescens Franchet, Pl. Delavay.: 203 (1890).

Type: Yunnan, ‘les bois de Fang-yang-tchang’, 19 July 1887, Delavay

svi! tole; P!, 180).

Distribution: Nepal to Bhutan, S Tibet, N Burma, W China.

I am grateful to Henry Noltie (E) for help here with literature

pertinent to the flora of Bhutan.

References

Green, P.S. 1994. Oceanic Islands I. Flora of Australia. 49.

Heine, H. 1967. “Ave Caesar, botanici te salutant”; L’épopée napoléonienne

dans la botanique. Adansonia I, 7: 115-140.

Lair, P-A. 1814. Description des Jardins de Courset, Situés aux Environs de

Boulogne-sur-Mer. Déterville, Paris.

Mabberley, D.J. 1990. The significance of the three independent ‘Kew’

editions of JOHNSON’s Gardener’s Dictionary. Fedde’s Repert. 101:

55-68.

Mabberley, D.J. 1991. The problem of ‘older’ names. Pp. 123-134 (Chapter

14) in Hawksworth, D.L. (ed.), Improving the Stability of Names: Needs

and Options [Regnum vegetabile 123]. Koeltz, KOnigstein.

Nelson, E.C. 1990. *...and flowers for our amusement’: the early collecting

and cultivation of Australian plants in Europe and the problems

encountered by today’s taxonomists. Pp. 265-296 in Short, P.S. (ed.),

History of Systematic Botany in Australasia. Australian Systematic Botany

Society, Inc., South Yarra, Victoria.

Appendix: Validly published binomials in neither Australian Plant

Name Index nor Index kewensis

Abies dammara (Lamb.) Dum. Cours., Bot. Cult. ed. 2, 6: 474 (1811),

based on Pinus dammara Lamb., = Agathis dammara (Lamb.) Rich.

(Araucariaceae)

Agriphyllum fruticosum (L.) Dum. Cours., Bot. Cult. 2: 519 (1801) & ed.

2, 4: 264, 627 (1811), based on Gorteria fruticosa L., = Berkheya

fruticosa (L.) Ehrh. (Compositae)

314 Gard. Bull. Singapore 51 (1999)

Ailanthus sinensis Dum. Cours., Bot. Cult., ed. 2, 6: 227 (1811), nom.

superfl. pro A. glandulosa Desf., = A. altissima (Mill.) Swingle

(Simaroubaceae)

Broussonetia tinctoria (L.) Dum. Cours., Bot. Cult. 3 : 686 (1802) & ed. 2,

6: 367 (1811), based on Morus tinctoria L., = Maclura tinctoria (L.)

Steud. (Moraceae)

Bunium semicompositum Dum. Cours., Bot. Cult., ed. 2, 4: 409 (1811),

based on a plant from Spain, = ? (Umbelliferae)

Caprifolium americanum (Mill.) Dum. Cours., Bot. Cult. 2: 577 (1801) &

ed. 2, 4: 338 (1811), based on Periclymenum americanum Mill. =

Lonicera x americana (Mill.) K. Koch (Caprifoliaceae)

Caprifolium implexum (Sol.) Dum. Cours., Bot. Cult., ed. 2, 7: 209 (1814),

based on (& =) Lonicera implexa Sol.

Caprifolium japonicum (Thunb.) Dum. Cours., I.c. (1814), based on (& =)

L. japonica Thunb.

Caragana halodendron (Pallas) Dum. Cours., Bot. Cult. 3: 513 (1802) &

ed. 2, 6: 144 (1811), based on Robinia halodendron Pallas, =

Halimodendron halodendron (Pallas) Voss (Leguminosae)

Caragana pygmaea (L.) Dum. Cours., Bot. Cult., ed. 2, 6: 144 (‘pigmaea’

1811), based on Robinia pygmaea L. (Leguminosae)

Ceanothus glaber Dum. Cours., op.cit. 6: 268 (1811), nom. superfl. pro C.

microphyllus Michaux (Rhamnaceae); non Spach (1834=?), nec

Trelease (1897, i.e. C. x lorenzenii (Jepson) McMinn). Note: Spach’s

plant has always been obscure so it is good to see the name disappear

as illegitimate.

Cotyledon hybrida Hort. Paris. ex Dum. Cours., Bot. Cult. 5: 416 (1805) &

ed. 2, 5: 277 (1811), based on Kalanchoe spathulata DC. = K. laciniata

(L.) DC. (Crassulaceae)

Coutarea portlandia Dum. Cours., Bot. Cult. 2: 561 (1801) & ed. 2, 4: 317

(1811), nom. superfl. pro Portlandia hexandra Jacq. = C. hexandra

(Jacq.) K. Schum. (Rubiaceae)

Botaniste Cultivateur 315

Crotalaria oxalidifolia (Sims) Dum. Cours., Bot. Cult., ed. 2, 7: 314 (1814),

based on Loddigesia oxalidifolia Sims, = Hypocalyptus oxalidifolius

(Sims) Baill. (Leguminosae)

Diosma gracilis Dum. Cours., op. cit. 5: 216 (1811), based on a plant

cultivated at Malmaison as D. imbricata, = Adenandra sp. (Rutaceae)

Diosma praecox Dum. Cours., op. cit. 5: 211 (1811), based on a plant

cultivated at Courset, = Rutacea?

Duhamelia coccinea (Aubl.) Dum. Cours., op. cit. 4: 328 (1811), based on

Guettarda coccinea Aubl., = Isertia coccinea (Aubl.) J. Gmelin

(Rubiaceae)

Genista ferox (Poir.) Dum. Cours., Bot. Cult. 3 : 458 (1802) & ed. 2, 6: 68

(1811), based on Spartium ferox Poir. (Leguminosae)

Geranium lineatum Dum. Cours., Bot. Cult. ed. 2, 5: 18 (1811), based on a

plant cultivated at Courset, = Pelargonium sp. (Geraniaceae)

Geranium modestum Dum. Cours., op. cit. 5: 29 (1811), based on a plant

cultivated at Courset, = Pelargonium sp.

Geranium mucronatum Dum. Cours., Bot. Cult. 5: 377 (1805) & ed. 2, 5:

25 (1811), based on a plant from the Cape, = Pelargonium sp.

Geranium pygmaeum Dum. Cours., Bot. Cult. ed. 2, 5: 35 (‘pigmaeum’,

1811), based on a plant cultivated at Courset, = Pelargonium sp.

Geranium rigidum (Willd.) Dum. Cours., op.cit. 5: 30 (1811), based on (&

=) Pelargonium rigidum Willd.

Geranium squarrosum Dum. Cours., Bot. Cult. 5 : 379 (1805) & ed. 2, 5: 14

(1811), based on a Cape plant, = Pelargonium sp.

Gomphrena fruticosa Hort. Paris. ex Dum. Cours., Bot. Cult. 1: 645 (1802),

ed. 2, 2: 488 (1811) = Alternanthera porrigens (Jacq.) Kuntze

(Amaranthaceae)

Helianthemum anglicum Dum. Cours., Bot. Cult. 3 : 128 (1802) & ed. 2, 5:

182 (1811), based on a plant from England,= H. appeninum (L.)

Mill. (Cistaceae)

316 Gard. Bull. Singapore 51 (1999)

Helianthemum myrtifolium Dum. Cours., Bot. Cult. 3 : 127 (1802) & ed. 2,

5: 182 (1811), nom. superfl. pro Cistus canus L., = H. oelandicum

(L.) Dum.-Cours. (Cistaceae)

Hibiscus aristatus (Cav.) Dum. Cours., Bot. Cult. 3 : 62 (1802) & ed. 2, 5:

92 (‘aristata’, 1811), based on (& =) Pavonia aristata Cav.

(Malvaceae)

Hibiscus cuneifolius (Cav.) Dum. Cours., Bot. Cult. 3: 61 (1802) & ed. 2,

5: 91 (1811), based on (& = ) Pavonia cuneifolia Cav. (Malvaceae)

Linaria speciosa Hort. Angl. ex Dum. Cours., op.cit., ed. 2, 7: 142 (1814),

baséd on‘asplant from Egypt =, ."sp.-) rion Jacad Yiste}i— &

bipartita (Vent.) Willd. (Scrophulariaceae)

Lupinus fruticans Dum. Cours., op. cit. 6: 90 (1811), nom. superfl. pro L.

arboreus Sims (Leguminosae)

Mespilus sanguinea (Pallas) Dum. Cours., op. cit. 5: 452 (1811), based on

(& =) Crataegus sanguinea Pallas (Rosaceae)

Oenothera candida Dum. Cours., op. cit. 5: 361 (1811), nom. superfl. pro

O. tetraptera Cav. (Onagraceae)

Pelargonium nobile Hort. Angl. ex Dum. Cours., op. cit. 7: 360 (1814),

based on a plant cultivated at Courset, = P. sp. (Geraniaceae)

Pinus romaniae Dum. Cours. op. cit. 6: 459 (1811), based on a plant from

the ‘Levant’, = P. nigra J.F. Arnold ssp. pallasiana (Lamb.) Holmboe

(Pinaceae)

Rubus fragarioides (Michaux) Dum. Cours., op. cit. 7: 293 (1814), based

on Dalibarda fragarioides Michaux, = Waldsteinia fragarioides

(Michaux) Tratt. (Rosaceae). See above

Sapindus paniculata (Laxm.) Dum. Cours., Bot. Cult. 2: 769 (1802) & ed.

2, 4: 550 (1811), based on (& =) Koelreuteria paniculata Laxm.

(Sapindaceae)

Sapium sebiferum (L.) Dum. Cours., Bot. Cult. 3: 651 (1802) & ed. 2, 6:

326 (1811, Euphorbiaceae), based on Croton sebiferum L., = Triadica

sebifera (L.) Small (Euphorbiaceae)

Botaniste Cultivateur cal,

Serissa buxifolia Dum. Cours., Bot. Cult., ed. 2, 4: 331 nom. superfl. pro

Lycium japonicum Thunb., = S. japonica (Thunb.) Thunb.

(Rubiaceae)

Serratula odoratissima (Willd.) Dum. Cours., op. cit. 4: 625 (1811) = Trilisa

odoratissima (Willd.) Cass. (Compositae)

Terebinthus atlanticus (Desf.) Dum. Cours., Bot. Cult. 3: 575 (1802) &

ede 2. a 222, (1611), based on (& =) Fistacia atlantica Dest.

(Anacardiaceae)

Terebinthus pistacia Dum. Cours., Il.cc., nom. superfl. pro P. trifolia L., =

P. vera ds.

Terebinthus vulgaris Dum. Cours., op. cit. 6: 222 (1811), based on (& = )

P. terebinthus L.

Trichilia palustris [Thouin ex] Dum. Cours., Bot. Cult. 2: 813 (1802) & ed.

2, 4: 609 (1811) = ? Meliacea

Urtica arborescens Dum. Cours., Bot. Cult. 3 : 689 (1802) & ed. 2, 6: 370

(1811) = Urera baccifera (L.) Wedd. (Urticaceae). See above.

Xylosteum flexuosum (Thunb.) Dum. Cours., Bot. Cult., ed. 2, 7: 208

(‘Xylosteon’, 1814), based on Lonicera flexuosa Thunb., = Lonicera

japonica Thunb. (Caprifoliaceae)

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Gardens’ Bulletin Singapore 51 (1999) 319-320.

BRYOLOGICAL BIOLOGY, INTRODUCTION AND DIVERSE

BRANCHES

Peng-cheng Wu (Editor). 1998. Science Publisher, Beijing, China. xxvi +

357 pp. (in Chinese). ISBN 7-03-006377-5. Hard-cover. Ren-min-bi Y 58.

Postage extra. Available from, Prof. P.-C. Wu, The Herbarium, Institute

of Botany, Academia Sinica. Xiangshan, 100093 Bejing, China.

This book was published last year by the Academia Sinica in Bejing. It is

edited by Prof P.-C. Wu, a respected and experienced Chinese bryologist

in mainland China. Both the editor and many of the chapter contributors

are Staff of the Institute of Botany in Bejing.

The new publication, which has seventeen chapters, deals with almost

all aspects of bryology and is dedicated to Prof Pan-chieh Chen, the founder

of modern Chinese Bryology. In addition, it has Chinese and English

forewords written by the most senior Chinese cryptogam botanist, Prof.

Cheng-kui Zeng of the Institute of Oceanography of Chinese Academy of

Sciences, and by the editor. The large volume concludes with a combined

index to the scientific names of bryophytes and English and Chinese

botanical terms mentioned in the text. The main text, however, is written

in Chinese. I recollect that the book was ready for publication in the early

90’s. The delay was due to the policy change of Chinese publishing houses

that gave priority to more profitable books. Consequently, the review of

the literature in several chapters does not include publications dated after

1994.

That the new book was finally published is indeed a great

accomplishment. Each of the seventeen chapters aims to provide a

comprehensive review of the history and a summary of the progress and

the state-of-the-art in all branches of modern bryology, with special focus

on Chinese accomplishments. In that sense, the book is fittingly the first

manual of bryology written in Chinese.

The published titles of the various chapters of the new book and

their respective authors are as follows: 1. Brief History of Bryological

Research (Peng-cheng Wu); 2. Taxonomy and Systematic Arrangement of

the Primary Taxa of Bryophytes (Peng-cheng Wu); 3. The Fundamental

Structure of the Gametophytes and Sporophytes of Bryophytes (Jiang-xin

Luo); 4. Spore Morphology of Bryophytes (Yu-long Zhang); 5.

Chromosomes and Cytotaxonomy of Bryophytes (De-yuen Hong); 6.

Physiology and Biochemistry of Bryophytes (Ding-ji Shi); 7. Molecular

Biology of Bryophytes (Ding-ji Shi); 8. Physiological Ecology of Bryophytes

(Yan-hong Li); 9. Chemical Constituents of Bryophytes (Guan-fu He); 10.

Reproductive Characteristics of Bryophytes (Yu Jia); 11. Tissue Culture of

Bryophytes (Mei-zhi Wang); 12. Bryophyte Ecology (Yan-hong Li); 13.

Flora, Communities and Distribution of Bryophytes in Antarctica (Shun-

320 Book review

shi Hu); 14. Bryoflora and Its Geographical Distribution Patterns (Peng-

cheng Wu); 15. Fossils of Bryophytes (Jiun-rong Tao and Peng-cheng Wu);

16. Bryophyte Indicators for Environment (Ming-jou Lai); and 17.

Relationship Between Bryophytes and Man (Peng-cheng Wu).

Seven of the seventeen chapters are contributed not by professional

bryologists but by well-known Chinese angiosperm specialists in the areas.

It is a laudable effort for these non-bryologist authors to produce such a

readable review of the development of a particular field of bryology.

Nonetheless, in terms of volume editorship, depth of information and quality

of graphic presentation, the new book pales somewhat in comparison with

the 1984 edition of The New Manual of Bryology edited by Prof. R. M.

Schuster and published in English by The Hattori Botanical Laboratory in

Japan.

After a quick perusal, I am quite impressed by the informative

contents of the chapters on ecology, metabolic physiology, cytology, spore

morphology and fossils of bryophytes. The two chapters on the biochemistry

and chemical constituents of bryophytes also look good to a non-chemist

like me. However, the book is somewhat spoiled by its lack of essays on

the recent developments in molecular systematics, the origin and evolution

of bryophytes and the ontogenetic development of spores and protonema

of bryophytes.

Since there is no other bryological book covering a similar broad

range of topics as yet published in China, this new book becomes the de

facto authoritative reference on bryology inside China. As an encyclopaedic

source of information about bryophytes, it definitely meets the needs of

the Chinese bryological community, including the amateur, student and

professional groups. And, because it is written in Chinese, it easily fills a

relevant niche in the publication markets inside and outside China, such as

in Hong Kong, Taiwan and Singapore, where there is a large Chinese-

language population of plant hobbyists who have difficulty reading non-

Chinese botanical publications.

The new manual, together with a previous textbook, Bryology, written

by Prof. Ren-liang Hu, which was published by the Advanced Education

Publisher in 1987 (also in Chinese), will, for many years to come, continue

to educate, stimulate and promote interest in the study of bryophytes in

China.

Benito C. Tan

Department of Biological Sciences

National University of Singapore

Singapore 119260

Gardens’ Bulletin Singapore 51 (1999) 321.

STRUGGLE OF LIFE or The Natural History of Stress and Adaptation

by Martial Rossignol, Line Rossignol, Roelof A.A. Oldeman and Soraya

Benzine-Tizroutine with contributions by A. Ambroise, E.A.P. de Bruijn

and C. Caisne. Treemail, Heelsum, The Netherlands (http://

www.treemail.nl). x + 237 pp. 1998. £24.99. Distributed by Natural History

Book Service (http://www.nhbs.com).

This Franco-Dutch production is a peculiar publication. From the title,

one might expect a gentle run through the myriad ways in which living

organisms are suited to their natural environments. Instead the book is a

curious mixture of plant molecular biology, plant morphology and systems

ecology. These somewhat disparate topics are glued together by speculations

on the importance of temporal phenomena such as lunar and solar cycles

in controlling or influencing biological systems at all levels of organisation

from molecules to the biosphere. I probably lack the breadth and depth of

perception required to appreciate the underlying message. Other readers

of the book may, for instance, be more impressed than I was to learn that a

common feature of architectural metamorphosis is 5 — it being the number

of base pairs in a half turn of the DNA helix and the frequent number of

orders in a complete branch system. Devotees of the molecular genetics of

potato tissue cultures, tree architecture or Professor Oldeman’s pedantic

brand of vegetation ecology (the term ‘succession’ is banned from the

book) may want to acquire a copy. Those with a philosophical turn of

mind and a penchant for the ‘unexplained’ may also care to dip in. There

is no particular reason why botanists working in, or on, Southeast Asia

should see the book.

Ian Turner

Singapore Botanic Gardens

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