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THE GARDENS’ BULLETIN

SINGAPORE

VOLUME 52

Singapore Botanic Gardens

Singapore, 2000

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. 1-212 31 August 2000

Part2 Pp. 213-302 29 December 2000

Singapore Gardens’ Bulletin 52 (2000)

CONTENTS

Adema, F.

Notes on Malesian Fabaceae (Leguminosae-Papilionoideae).

ae i SINUS RAO EIITION a3) sc Sasevcecasssesisnunsascnescetivaiecondsdeocesssesaneasnsceetens 5 — 10

Argent, G.

Two interesting wild Musa species (Musaceae) from Sabah,

AP nr ree Recs ge ag Berieeel wt ae aex cute pies cd nas ab onal bnspbdomtdes rap roms eeneses 203 — 210

Boyce, P.C.

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

southem and’ western Indonesian Archipelago ...,......::s:csecieccseorsnnsencoseeoenees 101 — 183

Boyce, P.C.

The genus Rhaphidophora Hassk. (Araceae-Monsteroideae-

Deena Romie | MNT TIT eS A ENMU ooo cose ssa sbaaennekercaietal spneteldvecmnstih pa tueargedunionnn 213 — 256

Boyce, P.C. and J. Bogner.

An account of neotenic species of Rhaphidophora Hassk.

(Araceae-Monsteroideae-Monstereae) in New Guinea and Australia ........ 89 — 100

Mabberley, D.J. A tropical botanist finally vindicated 0.0.0.0... eeseseecesecesecesnecesneeeenes 1—4

Nguyen, Q.B. and M. Newman.

A new species of Alpinia (Zingiberaceae) from Vietnam ...............ccceeeees 211 — 212

Nguyen, T.H. and R. Kiew.

New and interesting plants from Ha Long Bay, Vietnam............. eee 185 — 202

O’Bymne, P. and J.J. Vermeulen

Notheria diaphana gen. nov., sp. nov. (Orchidaceae), a gem from

SULAWESE cok. scus3.% Be Ue Reese Sa alae, 2B Sse een, APT ee Ne SR TM Ben 285 — 288

Salma, I.

The significance of pollen morphology in the taxonomy of the genus Durio

EUS Sg G2 te SRS ER aha WD Cine Ce ai er eT ee mm ene Wy oe 261 — 271

Slik, J.W.F., Priyono and P.C. Welzen.

Key to the Macaranga Thou. and Mallotus Lour. species (Euphorbiaceae) of East

POPS EST NG OSE ARE aD Sa On Oe De CN en 11 — 88

Vermeulen, J.J.

Carr’s spirit collection of Kinabalu orchids recovered, with some

notes on Bulbophyllum species present in this collection «0.0.0... cece 273 — 284

Vermeulen, J.J. and P. O’Byrne

Bulbophyllum jiewhoei (Orchidaceae), a new species of unknown

aoa RONAN ce MER NP ca, iat Ua ae oe Sige id NO op dad end anatabvnorgstZansadeus 293 — 296

Vermeulen, J.J. and L.G. Saw

Bulbophyllum sect. Hirtula (Orchidaceae): a new species and a new

record for Peninsular Malays ..0.:2.1.0.c..0cc0sssceenesvunnnanntneessteeaetee ee 289 — 292

Zhu, H.

A new species of Lasianthus (Rubiaceae) from Peninsular

Malaysia .2:.:...:s<cceececsesssncensicennnnsseeieng nee neE RO at et de teen eee ene 257 — 259

BOOK REVIEWS

Turner, I.M. The Plants of the Singapore Botanic Gardens: an

Annotated Check -Iist ......csesseianexeaicss tgnnndoerdece aati mee nc acco aa ane ee 297 — 298

Soepadmo, E. and L.G. Saw (eds.) Tree Flora of Sabah and Sarawak

VOWIME 3B pe.csiccnnceosasseosdeusdssnedagdanpencedegqeetereen aausec easter 299 — 300

Shaw, A.J. and B. Goffinet (eds.) Bryophyte Biology ........ cece eeeceesteceseeeseeeeeeenee 310 — 302

The Garden Bulletin

Singapore

JOL. 52 (Part 1) July 2000 | | ISSN 0374-7859

NATIONAL PARKS BOARD

_ Singapore Botanic Gardens Cluny Road Singapore 259569 Tel: 4741165 Telefax: 4754295

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 Hong Kong

Dr T.W. Foong Dr M.C. Roos

(Assist. Editor) Riksherbarium

Singapore Botanic Gardens

Dr S.C. Chin

Singapore Botanic Gardens

Dr M.J.E. Coode

Royal Botanic Gardens

Kew, U.K.

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.

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

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

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

Singapore.

Instructions for contributing authors are found on the inside backcover.

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The Gardens' Bulletin

Singapore

VOL. 52 (Part 1) July 2000 ISSN 0374-7859

CONTENTS

Mabberley, D.J.

Pe UP ODIC A WOt Anis (TMA VIMCIC MLE 3..6.0.6024-ncnnecsebeseatans eco tieathvessavupasdexdesnnssacepusttenrsekeconesdhvens f

Adema, F.

Notes on Malesian Fabaceae (Leguminosae-Papilionoideae). 6. The

Serine MARLENE) EPI SOOEE ITE OVE Sece oe 3c) ole nc 285 ced add cs Beh +s s0 BL 0 «+ 0 Sobnsep aca Bere xk Sp ak ana vb ECan oto sbns Spends 5

Slik, J.W.F., Priyono and P.C. van Welzen

Key to the Macaranga Thou. and Mallotus Lour. species

(Euphorvinecac) Of Bast Kalimantan; TndOmesia ..2.c..222 ccs cscscescondvscccsdueteectbeses sen eccetsadeess 11

Boyce, P.C. and J. Bogner

An account of neotenic species of Rhaphidophora Hassk. (Araceae-

Monsteroideae-Monstereae) in New Guinea and Australia ......0......ccccssecescssteceseeeseeeees 89

Boyce, P.C.

The genus Rhaphidophora Hassk. ee Monsteroideae-

Monstereae) in the southern and western Indonesian Archipelago .............eeeeeeeeeee: 101

Nguyen, T.H. and R. Kiew

New and interesting plants from, Ha Long Bay; Vietnam «.....3....6.5.isi.....seeccesessecceee- 185

Argent, G.

Two interesting wild Musa species (Musaceae) from Sabah, Malaysia ............ eee 203

Nguyen, Q.B. and M. Newman

Pew spccicsol Alpmia (Zineiberaccac) TOM ViICIMAMM) ..:....:.6.s.ccsencsennsersesernncensneestenee 211

Date of Publication: 31 August 2000

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 52 (2000) 1-4.

A Tropical Botanist Finally Vindicated

DAVID J. MABBERLEY

The National Herbarium of the Netherlands, University of Leiden,

The Netherlands, and

The Royal Botanic Gardens, Sydney, Australia

Abstract

Proof of the existence of the note of authority from Governor, Sir Shenton Thomas, to the

Japanese, commending E.J.H. Corner to help conserve the Singapore Museum and Botanic

Gardens has come to light.

Professor E.J.H. (‘John’) Corner (1906-1996) was one of the most colourful

biologists of the century. He was Professor of Tropical Botany in the

University of Cambridge, UK, and one of the very few botanists to work

on both fungi and seed plants. His work was thought-provoking and often

controversial, notably his views on systematic mycology and his advocacy

of his Durian Theory of angiosperm evolution (Mabberley, 1983, 1999).

For many outside science, though, the most controversial part of his

work was in Singapore during the occupation by Japanese Forces in WWII.

Corner had come to Singapore in 1929 as Assistant Director of the Botanical

Gardens. He found there:

‘the main library and the main herbarium...The herbarium was a storehouse of

knowledge...Not only were there hundreds of thousands of specimens scientifically

arranged ...together with their invaluable field-notes...but also a large number of

unpublished drawings and paintings of plants, records of expeditions as far afield as

New Guinea, fieldbook manuscripts...it was a unique centre of botany. Similarly at

Raffles Museum in Singapore there were the zoological, archaeological, and

anthropological counterparts...as well as the Raffles Library and the library of the

Malayan Branch of the Royal Asiatic Society’. (Corner, 1981)

The British surrendered Singapore to the Japanese at 5 pm on Sunday

15 February 1942. The Governor, Sir Shenton Thomas, had his office in

the Fullerton Building. Here Corner, with many other officials in the

Colonial Service, sought refuge while the city fell prey to looters. Through

the ensuing restless night, Corner

‘began pondering by what means the treasure houses of knowledge which had no

military significance might be saved...If they were looted, I could not see that funds

would ever come to start them again. Looting was the imminent danger. We had

heard repeatedly during the last two months, as the [British] withdrawal [through

the Malay Peninsula] hastened to Singapore, how the looters broke into buildings

before the Japanese had gained control, and how they smashed doors, cupboards,

desks, safes, and boxes, scattering the contents, throwing books and files into heaps

2 Gard. Bull. Singapore 52 (2000)

or bonfires, slashing pictures, and wrenching off taps to let the water run over

books, papers, clothes, and carpets. I was thinking not just of the Botanical Gardens

and Raffles Museum, but of the libraries of Raffles College, the College of Medicine,

Government offices, even at Government House, and the service they could render

in the restoration of peace and order...The whole progress of science in the Malay

Peninsula seemed doomed’.

Next day, Corner

‘spoke with several foresters and agriculturalists, who were also on that floor of the

Fullerton Building, of the need to appeal to the Japanese to help look after scientific

collections and records, only to meet the resolve that the British must now stand

together, shoulder to shoulder, in custody and humility’.

The following morning Corner decided to put his proposition directly

to the Governor, but as he

‘opened the door to go downstairs, up came the Aide-de-Camp to order [Corner]

to Sir Shenton who had had the same idea. Sir Shenton wrote me a pencilled note

in which he requested the Japanese authorities to preserve the scientific collections,

libraries, and matters of historic interest, particularly at the Museum and Botanical

Gardens’.

Corner

‘returned from Sir Shenton Thomas to [his] companions, showed them the note,

and asked if any one would accompany [him], but in vain; there was the same

determination to suffer in unity, which [he] would not, and [he] saw that at heart

they were not scientists’.

Corner was able to get the dispatch through the Japanese authorities

who passed it to Professor Hidezo Tanakadate (1884-1951), vulcanologist

and geologist of Tohoku Imperial University at Sendai, Japan (Corner,

1946, 1951). Tanakadate was put in charge of the Gardens and the Museum:

with him, Corner and others secured the collections and brought together

the island’s libraries, such that the heritage was saved for what is now the

modern state of Singapore. A ship of Japanese technicians and officials

had been sunk and the Japanese authorities therefore arranged for suitably

qualified British people to take their places — and not to be interned in the

Military Camp at Changi. Thus Corner and others became Civil Internees

and continued their duties under Japanese orders. As is well known, most

of the British were Kept in disgusting conditions in the prison camp. Despite

Corner’s considerable success in organising food supplies, he was thereafter

despised as a collaborator by many. For some of those who survived Changi,

their loathing was to become a lifelong one. With the publication of Corner’s

account of the episode in The Marquis. A tale of Syanon-to (1981), that

loathing was re-awakened: Corner’s account, particularly of the Governor’s

note, was questioned.

Sadly for Corner, the all-important dispatch was lost in a fire in

Sendai in 1945, when all of Tanakadate’s belongings were destroyed. Only

very recently has it come to light in the West that Tanakadate had in fact

published the Governor’s despatch (pencilled, like many of those of Field

A Tropical Botanist Finally Vindicated 3

Marshal Montgomery, for example, so as to avoid smudging), in a

contemporary Japanese newspaper. We now have the Japanese version of

the events of the time.

Tanakadate had been based in Saigon, Vietnam, and wrote in his

article sent from Singapore, ‘Conservation of a culture. A month’s travel

in Singapore — Part 1’:

‘As I was often being told of the significance of the fall of Singapore in the Great

East Asian War, I stayed by the radio once I heard the news. I went to bed late.

Next morning, while I was having breakfast, I was told ...that I was to be at the

airport by 10 00. I got ready in five minutes....It was a memorable flight. We flew

over Johore at the southern tip of the Malay Peninsula...As we were landing we

saw damaged aeroplanes, army barracks shot through like bee-hives...In the direction

of Singapore, we saw black smoke from burning oil rising to the sky with red flames

flaring up inside it....

In the morning of 17 February we travelled to Singapore City by car. We arrived at

the city office and met people who worked there. They were chiefs of the military

administration section, the executive section, the industry section, and Mr Toyota,

who was the ex-Consul-general of Singapore. When they saw me, they said, “We

have just received this from the ex-Governor”, and handed me a letter.

The letter said “We truly hope that you will conserve the Museum and the Botanical

Gardens at least, and that you will engage Dr [sic] Corner, who was formerly the

Assistant Director of the Botanical Gardens for that purpose”. I replied, “I have

come here in order to conserve the cultural heritage. The requisitioning of the

Museum is my first objective”. At my words, everyone present, including the former

mayor of Singapore, British and Japanese alike, glanced at one another and seemed

pleased with my answer. Then they came to shake hands. It must have been a

surprise for them to discover that a scientist from a university had come to conserve

the local cultural heritage of an enemy country. The chief of the executive section

said to me, “I entrust you with all the cultural institutions” and asked, “Will you

accept the post of Director of the Botanical Gardens and Museum?” I answered

ROR aleve

On 18 February, Dr Corner came and helped me. I took over the Museum and the

Botanical Gardens and began their conservation.’

(Asahi Shinbun 4 April 1942; modified, partly with the help of Rie Shiraishi and

Peter Valder (Sydney), from a translation by Tadashi Kajita (Botanical Gardens,

University of Tokyo), who also obtained for the author a copy of an original in the

library of the National Diet in Tokyo).

And so John Corner is vindicated, at last.

References

Corner, E.J.H. 1946. Japanese men of science in Malaya during Japanese

Occupation. Nature. 158: 63.

Corner, E.J.H. 1951. Nature. Prof H. Tanakadate 167: 586—7.

Corner, E.J.H. 1981. The Marquis. A Tale of Syonan-to, pp. 21—39, 166—

4 Gard. Bull. Singapore 52 (2000)

168. Heinemann Asia, Singapore.

Mabberley, D.J. 1983. ‘Corner, Edred John Henry’, pp. 158—159. In A.

Bullock & R.B. Woodings (eds), The Fontana Biographical Companion

to Modern Thought. Fontana, London.

Mabberley, D.J. 1999. ‘Edred John Henry Corner, C.B.E.’, Biographical

Memoirs of Fellows of Royal Society. 45:77—93.

Gardens’ Bulletin Singapore 52 (2000) 5—10.

Notes on Malesian Fabaceae (Leguminosae-Papilionoideae)

6. The Rare Genus Burkilliodendron

FRITS ADEMA

Nationaal Herbarium Nederland, Leiden Universiteit Branch, P.O. Box 9514,

2300 RA Leiden, The Netherlands

Abstract

Comprehensive descriptions of the rare Malaysian genus Burkilliodendron Sastry and its

single species B. album (Ridl.) Sastry are outlined and its relationship to Fordia discussed.

Introduction

Burkilliodendron Sastry is a rare, monotypic Malesian genus of papilionoid

legumes. It was described by Ridley (1925) as Burkillia but since then has

been little studied (Whitmore, 1969; Geesink, 1984). The material of the

only species (B. album) is very scarce; presently it is known only from the

type and one other specimen. The original description is short and in part

rather uninformative. For this reason a comprehensive and more detailed

description is given below together with some discussion on its position

within the family.

Nomenclature

Burkillia Ridley (1925) is an illegitimate name because of the earlier

published name of a green alga Burkillia West & West (1907, Annals of

the Royal Botanic Garden Calcutta 6: 228). It was renamed independently

by two authors in 1969 (Sastry, 1969; Whitmore, 1969). The name

Burkilliodendron Sastry was published first (23 Jan 1969) with Alloburkillia

Whitmore published later (9 Aug 1969). Burkilliodendron is the correct

name with its type species now known as Burkilliodendron album (Rid1.)

Sastry.

Systematics

Ridley (1925) based his description and comments on detailed information

from unpublished notes by S.T. Dunn. Ridley considered his new genus to

6 Gard. Bull. Singapore 52 (2000)

be closely related to Fordia and Millettia (probably sect. Albiflorae, now

Fordia, see Dasuki & Schot, 1991). Hence, Burkilliodendron has been

included in the tribe Millettieae. Geesink (1984) discussed this relationship

in his Scala Millettiarum: ‘The short inflorescence, consisting of two

seemingly collateral racemes with rather small and feeble flowers remind

me of the Phaseoleae subtribe Glycininae, and more specificially of the

genus Glycine.’ However, his interpretation of the inflorescence is wrong:

it is a pseudoraceme, not a raceme (or a panicle, see his Table 6, 3).

‘Feeble’ flowers are found in Millettieae as well as in Phaseoleae. All my

attempts to identify the type specimen of B. album invariably lead to

Millettieae and Burkilliodendron. However, I agree with Geesink (1984)

that the separation of the tribes Millettieae and Phaseoleae may not be

based on firm ground. Within the tribe Millettieae, the genus seems quite

isolated but it is certainly not closely related to Fordia.

Comparison with Fordia

Superficially Burkilliodendron resembles unifoliolate species of Fordia.

However, the differences between these two genera are numerous (Table 1).

Table 1. Comparison between Burkilliodendron album, Fordia unifoliolata, and

unifoliolate forms of Fordia albiflora.

Burkilliodendron Fordia unifoliolata Fordia albiflora

Rachis present absent present

Stipellae present absent absent

Pulvinus 2 1 Z

Axillary buds 5 or 7 lor 3 lors

strongly flattened + globular + globular

Inflorescences per axil 1 to several 1 1, rarely 3

Calyx lobes in bud not imbricate imbricate imbricate

Standard, basal callosities absent present present

Pods glabrous velvety velvety

The rare genus Burkilliodendron v4

Specimens

Burkilliodendron album was until recently known only from the type

material. This specimen Burkill & Haniff 12493 (not Burkill 12498 as cited

by Ridley) at Kew consists of two duplicates, one with flowers and one

fruit, the other with flowers only. The first is selected as the lectotype, the

second as an isolectotype. There is one other duplicate specimen with

flowers at Singapore. Burkill & Haniff 12493 was collected from Durian

Fipit, Perak. _.

To my surprise, a specimen collected in 1993 during the Belum

Scientific Expedition (Latiff et al. 4012) also belongs to this species. This

specimen was collected from the Temenggor Forest Reserve, Gerik, Perak.

Burkilliodendron

Burkilliodendron Sastry, Bull. Bot. Surv. India 10 (1969) 243; Geesink,

Leiden Bot. Series 8 (1984) 82. — Burkillia Ridl., Fl. Mal. Penins. 5 (1925)

304, nom. illeg. — Alloburkillia Whitmore, Gards. Bull. Sing. 24 (1969) 4.

— Type: Burkillia alba Ridl. [= Burkilliodendron album (Rid1.) Sastry].

Shrubs or treelets. Leaves unifoliolate; stipules present, caducous; stipellae

present, persistent. Axillary buds collateral. Leaflets entire. Inflorescences

axillary, pseudoracemes, | to several per axil; flowers 1 or 2 on knob-like

brachyblasts. Bracts to inflorescences, brachyblasts and flowers present,

persistent. Bracteoles at the top of the pedicels, persistent. Calyx

campanulate, bilabiate, lobes not imbrictae; upper lip shortly bidentate;

lower lip with 3 longer teeth. Corolla: standard without callosities; wings as

long as the keel petals. Stamens monadelphous, basal fenestrae present;

anthers all equal, fertile. Disc inconspicuous. Ovary very shortly stipitate;

ovules 2; style hairy in lower half, stigma terminal, capitate. Pods falcate,

thin-woody, dehiscent. Seeds 2, discoid.

Distribution: One species. Peninsular Malaysia.

Burkilliodendron album (Ridl.) Sastry, Bull. Bot. Surv. India 10 (1969);

Geesink, Leiden Bot. Ser. 8 (1984) 82, pl. IV-26. — Burkillia album RidL.,

Fl. Mal Penins. 5 (1925) 305. — Alloburkillia alba (Ridl.) Whitmore, Gards.

~ Bull. Sing. 24 (1969) 4. — Type: Peninsular Malaysia, Perak, Burkill &

Haniff 12493, lecto (K — flowers and fruit), isolecto (K, SING — flowers).

8 Gard. Bull. Singapore 52 (2000)

von Os 98

Figure 1. Burkilliodendron album. a. branch with flowers and one fruit. b. flower. c. standard.

d. keel petal. e. wing. f. stamens. g. ovary.

The rare genus Burkilliodendron 9

Figure 1.

Shrubs or treelets, 5|0—150 cm high. Twigs terete, 4—S mm diam., hirsute.

Stipules acicular to narrowly triangular, 4—8 x 1 mm, caducous, outside

hirsute, inside glabrous. ‘Rachis’ 1—3 cm long, hirsute, pulvinus 3 mm

long. Stipellae acicular, 1—2 mm long, with few hairs. Leaflets elliptic to

narrowly obovate, 17—27.5 x 5.5—8.5 cm, index 2.9—3.5, base cuneate or

rounded, apex acuminate, acumen 20—30 mm long, above hirsute on midrib

and nerves, below thinly hirsute, midrib above slightly raised, nerves above

+ flat or sligthly raised, 11 or 12 per side, 10O—20 mm apart; pulvinus 2—4

mm long. /nflorescences 2.5—5 cm long, peduncle up to 1 mm long, hirsute.

Bracts to the inflorescence acicular, c. 3 x 0.5 mm, outside sericeous, inside

glabrous; to brachyblasts and flowers similar to those to the inflorescences,

1.5—1.9 x 0.4—0.9 mm. Pedicels 1.5—2.5 mm long. Bracteoles acicular to

narrowly ovate, 1.2—1.4 x 0.2—0.4 mm, outside sericeous, inside glabrous.

Calyx 3—4 mm long, tube 2—3 mm long; upper lip + ovate, 0.5—1 x 2—4

mm, teeth c. 0.4 mm long; lateral teeth triangular, 1 x 1.5 mm, median

tooth triangular, 1.5—2 x 2 mm; outside sericeous, inside of teeth sericeous.

Corolla white. Standard: claw 2.5—3 mm; blade broadly ovate to obovate,

c. 7 x 6 mm, apex slightly retuse, outside sericeous except margins, inside

glabrous. Wings: claw c. 2.5 mm long; blade elliptic, c. 6 x 1.5 mm, rounded,

upper auricle 0.75 mm long; outside with very few hairs, wrinkled in lower

part at dorsal side, inside glabrous. Keel petals: claw c. 2.5 mm long; blade

boat-shaped. c. 6 x 3 mm, rounded, lateral pocket rather wide, c. 2.5 mm

long, upper auricle, 0.75 mm long; out- and inside glabrous. Stamens 7—9

mm long, free part 3 mm long, glabrous; anthers 0.5 x 0.2—0.4 mm, glabrous.

Ovary 4 mm long, sericeous; stipe 0.5 mm long, sericeous; style 5 mm long,

sericeous in lower half. Pods + hatchet-shaped, 7 x 1.5 cm, beaked, valves

glabrous outside, finely obliquely striate. Seeds discoid, c. 8 x 7 mm, hilum

c. 1 mm long.

Distribution: Peninsular Malaysia - Perak.

Habitat & ecology: Lowland dipterocarp forest, up to 350 m altitude.

Flowering: June, Sept; fruiting: June.

Notes: Known only from two specimens from northern Perak; probably

very rare although Burkill & Haniff wrote on their label, ‘quite common’.

_ Additional specimen: Perak — Gerik, Temenggor F.R. A. Latif et al. 4012

(L, UKMB).

10 Gard. Bull. Singapore 52 (2000)

Acknowledgements

The loan of type material from K is gratefully acknowledged. Mr. Jan van

Os of the Rijksherbarium made the beautiful drawing.

References

Dasuki, U.A. & A.M. Schot. 1991. Taxonomy of Fordia Hemsley

(Papilionaceae: Millettieae). Blumea. 36: 191—204.

Geesink, R. 1984. Scala Millettiarum. Leiden Botanical Series 8.

Ridley, H.N. 1925. Flora Malay Peninsula. Volume 5. Reeve, London.

Sastry, A.R.K. 1969. Burkilliodendron — A new generic name in Fabaceae.

Bulletin Botanical Survey India. 10: 243.

Whitmore, T.C. 1969. III. Leguminosae. Gardens’ Bulletin Singapore. 24:

4—6,

Gardens’ Bulletin Singapore 52 (2000) 11-87.

Key to the Macaranga Thou. and Mallotus Lour. Species

(Euphorbiaceae) of East Kalimantan, Indonesia

J.W.F. SLIK', PRIYONO?’ ann P.C. VAN WELZEN'

'Nationaal Herbarium Nederland, Universiteit Leiden Branch, P.O. Box 9514,

2300 RA Leiden. The Netherlands

*Wanariset Herbarium, P.O. Box 319, Balikpapan 76100, Indonesia

Abstract

A key to all Macaranga (27 taxa) and Mallotus (19 taxa) species known to occur in East

Kalimantan (Indonesia) is provided in this paper. The key is mainly based on vegetative

characters proven useful in the field. Some reproductive characters are included if

identification is otherwise impossible. The taxa are described briefly by their diagnostic

characters. Drawings are provided for most of the treated species.

Introduction

Species of the genera Macaranga Thou. and Mallotus Lour. (Euphorbiaceae)

often form a characteristic part of secondary vegetation in Southeast Asia,

and many species are among the first colonisers of disturbed vegetation. A

considerable number can also be found in late secondary vegetation and

the understorey of primary forest. The wide ecological range of both genera

with the more or less well-defined ecological preferences of the various

species, makes them potentially very useful for recognising the disturbance

history of a forest.

However, a Key to the species is still much in need, especially one

based on vegetative characters in order to identify sterile specimens. Keys

presently available for East Kalimantan are either outdated (Pax and

Hoffmann, 1914), do not cover all the species (KeBler & Sidiyasa., 1994),

or are partly based on the availability of both flowering and fruiting material

(Airy Shaw, 1975; Whitmore, 1975).

The key presented here is based on collections from East Kalimantan,

which are present in the Nationaal Herbarium Nederland, Universiteit

Leiden Branch (L) and the Wanariset Herbarium, East Kalimantan (WAN).

Reproductive characters have only been added when identification is

otherwise impossible. It is important to note that this key is made specifically

_ for East Kalimantan specimens. Identification of plants from outside this

area may not work, even if the species are included in this key. Some

species show strong dimorphism between their juvenile and adult states.

These are treated briefly in the species descriptions but not in the key,

12 Gard. Bull. Singapore 52 (2000)

which is based on (sub-)adult individuals. In general, the key will work

well for individuals that are taller than 0.5 m. An overview and keys for

several seedlings can be found in Bodegom et al. (1999). Short descriptions

of ecology, distribution and habit of the species are given, based on

information on the labels of East Kalimantan specimens and literature

references (Airy Shaw, 1975; Argent et al., 1998; KeBler & Sidiyasa, 1994;

Kefler et al., 1995; Sidiyasa et al., 1999; Whitmore, 1975).

Key to the Genera

la. Glabrous or indumentum simple. Twigs solid to hollow (ant inhabited).

Leaves alternate, often peltate, often lobed, sometimes with few basal

macular glands on upper surface. Inflorescences mostly branched.

Staminate flowers usually completely surrounded by bracts. Bracts

sometimes fimbriate. Stamens up to c. 30, anthers clearly (3—)4 celled,

apicifixed, theca opening apically. Styles partly connate, stigmas free.

Fruits 2—6 locular, usually smooth, sometimes spiny........ Macaranga

1b. Glabrous or indumentum simple to tufted to stellate. Twigs solid.

Leaves alternate to opposite, when opposite usually unequal in size or

shape, sometimes peltate, rarely lobed, usually with few basal macular

glands on upper surface. Inflorescences usually unbranched. Staminate

flowers not completely surrounded by bracts. Bracts with entire

margins. Stamens up to c. 150, anthers 2 celled, basi- to dorsi-fixed,

theca opening extrorse. Styles completely fused, apically splitting into

(2)-3-(4) stigmas. Fruits (2)-3-(4)-locular, usually spiny ........ Mallotus

Macaranga Thou.

Shrubs to trees up to 35 m tall, dioecious. Glabrous or indumentum simple.

Bark sometimes prominently hooped (rings surrounding the stem). Wood

soft. Twigs sometimes hollow and ant-inhabited (small openings), sometimes

whitish-glaucous. Stipules small and narrowly triangular, to very large and

ovate to rounded, sometimes recurved and forming an enclosed hollow

inhabited by ants. Leaves alternate to spirally arranged, small to very large;

blades often peltate, often 3(—5)-lobed, pinnate to basally tripliveined to

palmativeined, upper surface sometimes with basal macular glands, lower

surface often gland-dotted. Inflorescences axillary to rarely sub-terminal.

Staminate inflorescences mostly branched; bracts large, usually surrounding

the flowers, sometimes fimbriate; flowers 2—150 per node, sepals 2—4,

stamens 1—30, anthers (3- or) 4-celled, apicifixed, disc absent, pistil absent.

Pistillate inflorescences usually branched; bracts persistent to caducous;

Key to Macaranga and Mallotus of East Kalimantan l

(yo

flowers 1 to several per node, calyx short, ovary 1—6-locular, styles long to

short, partly connate, stigmas free. Fruit a leathery to berry-like capsule,

smooth to spiny to horned, sometimes with conspicuous glandular patches,

dehiscing into bivalved parts. Seeds often with red to pink aril.

la.

1b.

| 9a.

9b.

Key to the Macaranga Taxa

(EES 2 Ea eg EE t enn en gee eRe Ce BO 2

Movies Tncvebepwy (LLCs cere STAR AD IA ) os ccm ne nt ene edennssnnnrecesdnncenensie- 26

Se Ee Ee Se eA RRR Sa Ree 3

Leaves basally tripliveined to palmativeined .............. ee eee 10

Plants completely glabrous on vegetative parts ................eeeeeeeeeeeeee -

Plants hairy on twigs, petioles, and/or leaf undersides........................ 2

Leaves whitish glaucous on under surface. Basal veins of equal length

RE TOR = gh Ca | | 5. M. conifera

Leaves not whitish glaucous on under surface. Basal veins shorter

than at the centre of the lamina ............ 18. M. lowii var. kostermansii

Vegetative parts with hairs longer than 1 mm ............ eee eee 6

Vegetative parts with hairs shorter than 1 mm... eee 7

Leaf lamina 5—7 cm long, upper surface with conspicuous secondary

oe A Sees Re COM Be i: Be) Se ee 9. M. fulva

Leaf lamina 7—17 cm long, upper surface with inconspicuous secondary

veins. Pedicels 1.5—3 cm long ............. 4. Macaranga repando-dentata

Basal veins shorter than at the centre of the lamina...........0..00......2... 8

Basal veins of equal length or longer than higher on the lamina...... 9

Twigs and petioles densely hairy. Petioles usually shorter than 2 cm

a Ri aaa Baal isc ta deni anit cuas ica 4. M. brevipetiolata

Twigs and petioles mostly sparsely hairy. Petioles usually longer than

I a i ah Bi Baal Baines hn 17. M. lowii var. lowii

Lower surface of leaves conspicuously whitish glaucous, velvety hairy,

with secondary veins not prominent .......... eee 5. M. conifera

Lower surface of leaves not glaucous, not velvety hairy, with prominent

CES ES ES ee eee 6. M. costulata

10a.

10b.

lla.

11b.

12a.

12b.

13a:

13b.

14a.

14b.

ile

15b.

loa.

16b.

Lia:

17b.

18a.

18b.

Gard. Bull. Singapore 52 (2000)

Leaves: peltate:.....ciid..cc ads. .ce Seva leh Sie ieee ee eee 11

Leaves \not peltate ..:tait.i1...an. eae See ee 20

Leaves not lobed iss... ee ei eee Denes 12

Leaves lobed.......0i4..44. 220. ee See, 18

Petiole insertion more than 1.4 cm from blade base. Upper surface of

leaves usually without basal macular glands (when present then more

150.11 aA) nanan Hei amP EERO MMErIE basi rrbrdses 2! WMore dently jem bck vee eae 13

Petiole insertion less than 1.4 cm from blade base. Upper surface of

leaves sometimes with 2 basal macular glands ..................ccsssccseseeeees 16

Under surface of leaves densely hairy on whole surface, few to

WUMIETOUSIN BIANG GOU GU scot wee teks yee ere 25. M. tanarius

Under surface of leaves glabrous to sparsely hairy on main veins,

MUMETOUSTY GLATCT CHOU oo see sera csen onan -s- bate unto Laccenae tne ceases 14

Leaves never more than 11 cm wide. Capsules berry-like ...................

phlei Marana asinine Lene loan (0 p05 pie Doe AM MI. 28 oy 22. M. puncticulata

Leaves tarely less Tham 1) "Crn Wide. Castres tty oa ical acerge ae pe

Stipules tough (leathery when dried). Leaves leathery when dried,

under surface glaucous, secondary veins not prominent ..............e

Lsgiduish sven nese'Vag intee CONG ac RNa RRA a ee ta ar aan 23. M. recurvata

Stipules thin (papery when dried). Leaves papery when dried, under

surface not glaucous, secondary veins prominent........ 25. M. tanarius

Leaf under surface conspicuously glaucous, velvety hairy, secondary

VETS ILO SPOUT Ge -conzieoctcsetv caret ieiees cet chendan aan emmenin 5. M. conifera

Leaf under surface not glaucous, hairy but not velvety, secondary

VEMNIS PROM INGE 0 Leek nek ee ee Ae ce 17

Leaf upper surface with two basal macular glands....... 6. M. costulata

Leaf upper surface without basal macular glands........... 8. M. endertii

Stipules up to 4 cm long, ovate, apex acute, persistent, usually dead

and dry on living plant. Leaf lamina usually longer than 30 cm..........

Sb Sha hapeibevEaicetess ius id. ita ce eae ee ee ee 10.M. gigantea

Stipules up to 1.5 cm long, broadly ovate to ovate, apex rounded to

acuminate to acute, persistent to caducous, alive on living plant. Leaf

lamina shorter ‘than SO CH 232.5005. .essdlelasaDhnoteeentn ae eee eee ee een eee 19

Key to Macaranga and Mallotus of East Kalimantan |,

19a.

1b.

20a.

20b.

Dhak

Zip.

22a.

22:

Daa.

24a.

24b.

25a:

25h;

20a:

26b.

Twigs hairy, not glaucous. Stipules ovate, apex acute. Petioles hairy.

Leaf base rounded to truncate, blade shallowly 3 (to 5)-lobed, under

PO RCN ce etisanerdnnssnigtbnndcasnc tbl ated 7. M. depressa var. strigosa

Twigs usually glabrous, often glaucous. Stipules broadly ovate, apex

rounded to acuminate (to acute). Petioles rarely hairy. Leaf base often

emarginate, blade deeply 3-lobed, under surface glabrous to hairy....

Penne MR Ree Sorerd ar erie ey RS oN AE dah Cas beh abe cs RISENER Gea Laan 24

Pe Ae eNO OC ost aidancauesac WSO el ae AU. J Pon S.. Zi

TES aoa rt tua. MAIO creeds COM CRAIG Ae Re lds 22

Beat upper suriace completely hairy i1is.i:..00,..0.00. 26. M. trichocarpa

Leaf upper surface glabrous or rarely sparsely hairy on veins ........ 22

Vegetative parts glabrous or with few to numerous hairs. Leaves below

conspicuously whitish glaucous, velvety hairy, secondary veins not

Cream Cut nis caiedt S16 eo.ooee.). Ok Ak 5. M. conifera

Vegetative parts with few to numerous hairs. Leaves below not

glaucous, not velvety hairy, secondary veins prominent.................

PRM re Wee eA art Bts oa icciassguicanvnancadeseseceseredtls 6. M. costulata

Leaves shallowly 3-lobed (cusped), upper surface completely hairy,

base acute to rounded to slightly cordate ............... 26. M. trichocarpa

Leaves deeply 3-lobed, upper surface not completely hairy, base deeply

Enea URN ae NTE coh he lho eects nen sac atencicnanscdsvicnns tadeast@benckeuneak 24

Stipules persistent (present on all the leaves !). Leaf under surface

CRORE Ws Oak ee ieee ek eR 12. M. hosei

Stipules caducous (if persistent then leaf under surface velvety hairy).

Leat under suntace glabrous towelvety hairy..Aceinaie.iilies.....k.. 29

Twigs conspicuously glaucous, often with ant openings. Stipules ovate,

apex acuminate to acute. Leaf base often with large protruding glands

along the margin, leaf under surface mostly glabrous or with sparse

Mammals Veme yin Ao) icieics, eet ee te A 20. M. pearsonii

Twigs usually not glaucous, without ant openings. Stipules broadly

ovate, apex rounded to acuminate. Leaf base without large protruding

glands along the margin, leaf under surface velvety hairy ...................

ce Ome Bi Coweta cd eS cath oe see as pert Ld ets 21. M. pruinosa

ARN ONO NG dud ae ooh ape bib as esa vah de tavundid cancteprcnn tebe SnacOR eS Zt

Rea vie ee tiring eh se A AR eee heise Ria Abe Ae. 28

30b.

Sila.

ai:

32a.

S20:

33a;

33D:

34a.

34b.

35a.

Gard. Bull. Singapore 52 (2000)

Stipules persistent (present on all the leaves!), apex rounded.............

intl Be MTs Ah bot Rs pA LE Oe 12. M. hosei

SUPUES CHMUCOUS, VAPEX ACULE. ici iviccnctedbendscndavendsantesvad 20. M. pearsonii

Leaves mot lobed 4. s20.2.0i0 Qu ce A EL, Fon,

sheaves lobedii:,. 2.050400. 25100. Se a ee. Se 34

. Stipules recurved, forming a hollow between twig and stipule........ 30

~ stipules erect, not forming a WOM QW. nc.csvensrate en ae eae 32

. Stipules longer than wide (like a pair of horns), 1.5 —2.5 cm long ....

Payee ET iM cecnnitocimadicrvnecnsee ie NOME ete ee 16. M. lamellata

Stipules shorter than wide, up to 1.5 cm long (mostly between 0.5 and

dy corp min): ch 502. A save mente any vce poentsteee et nkde ces ae 31

Internodes often slightly swollen, sometimes hairy. Stipules caducous

to persistent, green (to red). Leaves ovate to lanceolate (always clearly

longer than wide), under surface glabrous or with sparse hairs, usually

not gland dotted. Plants often growing in swamps and along river

SSS ih ssi 1101s LI ah peo ad aes ha A ea rern ins 13. M. hullettii

Internodes usually not to sometimes slightly swollen, glabrous. Stipules

often persistent, red. Leaves broadly ovate (equal or only slightly

longer than wide), under surface often velvety hairy (to glabrous),

gland dotted. Plants mostly growing in dry places... eeeeeeeeeeeeeees

pees be act Leak eee I Ue RA 2. M. bancana (formerly M. triloba)

Twigs conspicuously swollen, ant openings large (c. 2 mm diameter).

Stipules carly.caducomsuiieeiels. cele Bee 22. M. puncticulata

Twigs not conspicuously swollen, ant openings absent or small (mostly

c. 1 mm diameter). Stipules caducous to persistent ............eeeeeeeeee 33

Twigs without ant openings. Stipules caducous to persistent. Leaves

leathery when dry, under surface densely gland dotted, slightly

glaucous. Fruits larger than 3 mm diam.................0 23. M. recurvata

Twigs usually with ant openings. Stipules persistent (present with all

the leaves !). Leaves papery when dry, below sparsely gland dotted,

not glaucous, Fritts small}: 3: mmadianiit eee 27. M. winkleri

Stipules recurved, forming a hollow between stem.and stipule....... 35

Stipules erect, not forming a hollow between stem and stipule ...... 42

Leaf base with conspicuously large glands along margin (larger than

Key to Macaranga and Mallotus of East Kalimantan LT

35D:

36a.

36b.

37a.

ait,

38a.

38b.

39a.

39b.

40a.

AQb.

Ala.

41b

Ada.

42b.

43b.

ace eared oh sted). eee a il 36

Leaf with large glands of equal size along the whole margin ..............

Glands along leaf margin with a clearly visible opening. Calyx caducous

aa ay cepted Se eats Ble he AOS... 1. M. aétheadenia

Glands along leaf margin without an opening. Calyx persistent .........

asia shkediese. tink AS. Meee a Ae 11. M. glandibracteolata

Stipules longer than wide, 1.5—2.5 cm long, resembling a pair of horns

femencies Dantes ANA cel aii, cat sae ra dalinks cual iaieh ates OU Ra coe St 16. M. lamellata

Stipules shorter than wide, up to 1.5 cm long (mostly 0.5 to 1 cm long)

eee Petes aaa ats italic cee i hltt ede hee) Ce pact veal! 38

Stipules not surrounding the stem. Leaves clearly 3 to 5-lobed ...... Bs

Stipules completely surrounding the stem. Leaves often shallowly 3-

Ree eG ordeal rth fod ett Lett at. ete Lice 40

eat tmider surlace CIA DrOUs ...:....02...50..0eee. ee etl 19. M. motleyana

Leal under surtace:velvety hairy) ...:c::..i..244 11. M. glandibracteolata

Stipules green, rarely red. Leaves conspicuously longer than wide,

usually less than 14 cm wide, shallowly lobed (cusped), lower surface

glabrous or with sparse hairs, usually not gland dotted. Plants often

growing in Swamps and along FiVeTS ........... ee eeeeeeeeeeeeeees 13. M. hullettii

Stipules usually red. Leaves slightly longer than wide, often more

than 14 cm wide, shallowly to deeply lobed, lower surface glabrous to

densely (velvety) hairy, usually gland dotted. Plants usually growing

ipetepaearhet cic ome nie es. Bale clei ts Seaiiied. santero 41

RIG e ADIOS ihe. Aiki le 2. M. bancana (formerly M. triloba)

PRA GIT iste Aes 2. ees AR Ree i beet 15. M. indistincta

Stipules narrowly triangular, c. 3—5 mm wide, not surrounding the

TMs atte etl aie tl Acetone ibs .ceidaneeteads desdehandss 43

Stipules broadly ovate to rounded, more than 5 mm wide, nearly to

bommplciely surrounding the Stem ).uwiWar. us ina ail At

Leaves with central lobe 3.5—5 times longer than wide, base of central

lobe not overlapping lateral lobes, under surface inconspicuously

PRO MOOR yk hire teeth del lusleeietsl aA 3. M. beccariana

Leaves with central lobe 1.3—2 times longer than wide, base of central

18 Gard. Bull. Singapore 52 (2000)

lobe usually overlapping lateral lobes, under surface conspicuously

glaucous:(pure white)).200%.cc4 Jeno eee 14. M. hypoleuca

44a. Stipules persistent (present with all the leaves !), apex rounded ........

ak he ic BRR I ee ae 12. M. hosei

4Ab: Stipules caducous, apex: Acute nn.wsumeneienmmrccmcmsrantense 20. M. pearsonii

1. Macaranga aétheadenia Airy Shaw (Fig. 1)

Diagnostic characters: Trees up to 20 m tall, dbh up to 20 cm. Bark hooped,

glaucous, latex red. Twigs hollow, glaucous, usually with ant openings.

Stipules recurved, half to completely surrounding the twig, red. Leaves

peltate, 3(-5)-lobed, base with very large hollow glands along the margin.

Staminate inflorescences branched. Fruits more than 2-locular, with

glandular bands, without spines.

Habitat & Ecology: Up to 300 m altitude. In disturbed places in primary

forest (gaps) and in secondary (logged) forests, often along roads or skid

trails. Usually inhabited by ants.

Distribution: Endemic to Borneo. In East Kalimantan found north of the

Mahakam river in Kutai and Berau.

2. Macaranga bancana (Miq.) Mull.Arg. (Fig. 2)

Diagnostic characters: Small trees up to 16 m tall, dbh up to 20 cm. Bark

smooth, hooped. Twigs hollow, transversely ribbed, usually with ant

openings, glabrous. Stipules recurved, completely surrounding the twigs,

usually red. Leaves peltate, 3-lobed, in young specimens sometimes very

deeply lobed, in adults trees usually very shallowly lobed, (seedlings not

lobed) under surface glabrous to densely velvety hairy, usually conspicuously

gland dotted. Staminate inflorescences branched. Fruits more than 2-locular,

without spines.

Habitat & Ecology: Up to 400 m altitude. In disturbed places in primary

forest (gaps) and in secondary forest. Mostly found in dry places but

occasionally also in swampy areas. On sandy loam. Usually inhabited by

ants.

Distribution: From Thailand southward into Peninsular Malaysia, Sumatra,

and Borneo. Found throughout East Kalimantan.

Key to Macaranga and Mallotus of East Kalimantan 19

Note: Often mentioned for this area as M. triloba (Blume) Miill.Arg.

However, recent insights have made it clear that this species does not

occur in Borneo, and that the correct name for the species found in this

area should be M. bancana (T.C. Whitmore & S.J. Davies, pers. comm.).

3. Macaranga beccariana Merr. (Fig. 3)

Diagnostic characters: Trees up to 15 m tall, dbh up to 15 cm. Bole straight.

Bark smooth, hooped, glaucous, latex clear to red. Twigs hollow, glaucous,

usually with ant openings. Stipules erect, apex pointed (they look spine-

like on living individuals), same colour as stem or twigs. Leaves peltate,

deeply 3-lobed, central lobe 3.5 to 5 times longer than wide, not overlapping

with lateral lobes, undersurface glaucous, glabrous. Staminate inflorescences

branched. Fruits more than 2-locular, without spines.

Habitat & ecology: Up to 900 m altitude. In disturbed places in primary

forest (gaps) and in secondary forests, often along roads. Usually inhabited

by ants.

Distribution: Endemic to Borneo. In East Kalimantan only found in the

northern part (Berau).

4. Macaranga brevipetiolata Airy Shaw (Fig. 4)

Diagnostic characters: Trees up to 14 m tall. Twigs solid, densely short

hairy apically. Stipules needle like, erect, caducous. Petioles usually not

more than 2 cm long, densely short hairy. Leaves not peltate, not lobed,

penniveined, usually obovate, base without or with indistinct basal macular

glands. Staminate inflorescences unbranched. Fruits 2-locular, with spines.

Habitat & ecology: Up to 300 m altitude. Primary forest understorey species,

also present in selectively logged forest. On sandy loam, sandy clay or dry

sandstone soils.

Distribution: Endemic to Borneo. In East Kalimantan found in Kutai.

5. Macaranga conifera (Zoll.) Mull.Arg. (Fig. 5)

Diagnostic characters: Trees up to 30 m tall, dbh up to 60 cm. Bole straight.

Bark hooped, dippled, latex red. Inner bark red. Wood white. Twigs solid,

usually glabrous, rarely hairy apically. Stipules erect, caducous. Leaves not

peltate to sub-peltate in adults, but peltate in seedlings and saplings!, not

20 Gard. Bull. Singapore 52 (2000)

lobed, tripliveined in adults, ovate, upper surface usually without basal

macular glands, under surface conspicuously glaucous, usually glabrous,

densely gland dotted (good visible in young individuals, inconspicuous in

adults), rarely hairy. Staminate inflorescences branched. Fruits 2-locular,

not spiny.

Habitat & ecology: Up to 1100 m altitude. In disturbed places in primary

forest (gaps) and in secondary forest, including scrub vegetation. Along

streams, roads, skid trails. Dry to periodically inundated to marshy terrain.

Loam, sandy, and limestone soils.

Distribution: Peninsular Malaysia, Sumatra, Borneo and Sulawesi. Found

throughout East Kalimantan.

Note: This species has a very strong dimorphy between the juveniles and

the adults. The juveniles have bright red petioles, clearly peltate leaves

and conspicuously large green persistent stipules.

6. Macaranga costulata Pax & K.Hoffm. (Fig. 6)

Diagnostic characters: Trees up to 30 m tall, dbh up to 60 cm. Latex red.

Twigs solid, sometimes with flaky yellow-orange indumentum on young

parts. Stipules caducous, erect. Leaves not peltate (tripliveined) to peltate,

not lobed, broadly ovate to ovate, young leaves with or without flaky

yellow-orange indumentum, upper surface usually with 2 basal macular

glands, under surface gland dotted. Staminate inflorescences branched.

Fruits 2-locular, not spiny.

Habitat & ecology: Up to 1800 m altitude. In primary and secondary forest,

including scrub. On clay, dark brown, limestone and sandstone soils.

Distribution: Endemic to Borneo. In East Kalimantan only found north of

the Mahakam river.

7. Macaranga depressa (Miull.Arg.) Miill.Arg. forma strigosa Whitmore

(Fig. 7)

Diagnostic characters: Shrubs to small tree up to 7 m tall, dbh up to 7 cm.

Bark hooped, latex clear, turning pink to red. Twigs solid, long hairy.

Stipules erect, semi-persistent, green-brownish. Leaves peltate, 3—5-lobed

(seedlings not lobed), upper and lower surface long hairy. Staminate

inflorescences branched. Fruits more than 2-locular, often with horn like

Key to Macaranga and Mallotus of East Kalimantan pal

appendages, not spiny.

Habitat & ecology: Up to 1200 m altitude. In primary and secondary forests,

also in swamp forest. Along roads, streams, forest edges. On sandstone to

red clayey loam.

Distribution: Endemic to Borneo. Present throughout East Kalimantan.

8. Macaranga endertii Whitmore

Diagnostic characters: Trees up to 15 m tall. Twigs solid. Stipules erect,

caducous. Leaves peltate, not lobed, upper surface without basal macular

glands, lower surface hairy. Staminate inflorescences branched. Fruits 2-

locular, not spiny.

Habitat & ecology: Up to 700 m altitude. In primary forest or on exposed

places. Mostly found on limestone.

Distribution: Endemic to Borneo. In East Kalimantan found north of the

Mahakam river.

9. Macaranga fulva Airy Shaw

Diagnostic characters: Bush of 4 m tall. Twigs solid, long hairy. Stipules

erect, spine-like, caducous. Leaves not peltate, not lobed, penniveined,

elliptic, lamina not longer than 7 cm, upper and under surface long hairy.

Staminate inflorescences unbranched. Pistillate inflorescences with very

large, leaf-like bracts. Fruits 2-locular, spiny.

Habitat & ecology: Lowland. In primary forest understorey. Sandy soil.

Distribution: Endemic to Borneo. Known from only one collection from

northern East Kalimantan (Bulungan district).

10. Macaranga gigantea (Rchb.f. & Zoll.) Mull.Arg. (Fig. 8)

Diagnostic characters: Trees up to 30 m tall, dbh up to 50 cm. Bole straight.

Bark hooped, lenticelled, latex pink to red. Twigs solid, apically hairy.

' Stipules erect, persistent but dying on stem (brown colour), large, up to 4.5

cm long, hairy. Leaves peltate (but deeply cordate in seedlings), shallowly

to deeply 3-lobed, very large, up to 50 by 50 cm, under surface conspicuously

hairy. Staminate inflorescences branched. Fruits 2-locular, not spiny.

22 Gard. Bull. Singapore 52 (2000)

Habitat & ecology: Up to 1400 m altitude, but usually lower. In disturbed

places in primary forest (gaps) and secondary forests, especially common

after fire, also present in scrub to grass lands. On sandy clay to sandy soils.

Distribution: From Thailand southward into Peninsular Malaysia, Sumatra,

and Borneo. Found throughout East Kalimantan.

11. Macaranga glandibracteolata S.J.Davies (Fig. 9)

Diagnostic characters: Trees up to 25 m tall, dbh up to 30 cm. Bark hooped,

latex red. Twigs hollow, glaucous, usually with ant openings. Stipules

recurved, not surrounding the twigs. Leaves peltate, 3-lobed, base sometimes

with very large glands along the margin, under surface velvety hairy,

sometimes glaucous. Staminate inflorescences branched. Fruits more than

2-locular, not spiny.

Habitat & ecology: Lowland. In primary forest in disturbed sites (gaps)

and in secondary forest, often along roads. Usually ant inhabited.

Distribution: Endemic to Borneo. Found in the northern part of East

Kalimantan (Berau).

12. Macaranga hosei King ex Hook.f. (Fig. 10)

Diagnostic characters: Trees up to 25 m tall, dbh up to 45 cm. Bark hoop

marked. Twigs solid to hollow, sometimes glaucous, with or without ant

openings, glabrous. Stipules erect, persistent, surrounding the twigs, apex

rounded. Leaves deeply cordate to peltate, 3-lobed, under surface sometimes

glaucous. Staminate inflorescences branched. Fruits 2-locular, not spiny.

Habitat & ecology: Lowland. In disturbed sites in primary forest (gaps)

and in secondary forests. Dry and swampy soils. Sandy and sandstone

soils. Can be ant inhabited.

Distribution: From Thailand southward into Peninsular Malaysia and

Borneo. Found in northern part of East Kalimantan, north of the

Sangkulirang limestone mountain range.

13. Macaranga hullettii King ex Hook.f. (Fig. 11)

Diagnostic characters: Small trees up to 12 m tall, dbh up to 11 cm. Bark

smooth, hooped, sometimes dippled, latex red. Twigs hollow, transversely

ribbed, sometimes hairy, usually with ant openings. Stipules recurved,

Key to Macaranga and Mallotus of East Kalimantan 25

usually green, sometimes tinged red to dark red, completely surrounding

the twig. Leaves peltate, usually not lobed, sometimes slightly 3-lobed

(cusped), always much longer than wide, under surface usually not gland

dotted. Staminate inflorescences branched. Fruits more than 2-locular, not

spiny but with horn-like apical appendages.

Habitat & ecology: Up to 1230 m altitude. Primary to lightly disturbed

secondary forests, often in swamps or along streams, also along roads and

in open places. On sandstone to sandy loam. Usually ant inhabited.

Distribution: Peninsular Malaysia, Sumatra, Borneo. Found throughout East

Kalimantan.

14. Macaranga hypoleuca (Rchb.f. & Zoll.) Mull.Arg. (Fig. 12)

Diagnostic characters: Trees up to 30 m tall, dbh up to 50 cm. Bole straight.

Bark smooth, hooped, glaucous, latex red. Twigs hollow, glaucous, usually

with ant openings. Stipules erect, persistent, spine-like, glaucous. Leaves

peltate, deeply 3-lobed (seedlings not lobed), central lobe usually

overlapping the lateral lobes, under surface conspicuously glaucous (bright

white). Staminate inflorescences branched. Fruits more than 2-locular, not

spiny, glaucous, seeds with conspicuous red aril.

Habitat & ecology: Up to 2400 m altitude, but usually below 1000 m. In

primary forest (gaps) and secondary forests, also in scrub, along roads,

streams, sometimes on alluvial places. On clayey to sandy clay soils. Usually

ant inhabited.

Distribution: From Thailand southward into Peninsular Malaysia, Sumatra,

Borneo. Found throughout East Kalimantan.

15. Macaranga indistincta Whitmore

Diagnostic characters: Trees up to 15 m tall, dbh up to 15 cm. Twigs hollow,

hairy, usually with ant openings. Stipules recurved, completely surrounding

the twigs. Leaves peltate, shallowly 3-lobed, under surface hairy,

conspicuously gland dotted. Staminate inflorescences branched. Fruits more

than 2-locular, not spiny.

Habitat & ecology: Up to 900 m altitude. In primary forest? Usually ant

inhabited.

24 Gard. Bull. Singapore 52 (2000)

Distribution: Endemic to Borneo. In East Kalimantan only known from

Kutai.

16. Macaranga lamellata Whitmore (Fig. 13)

Diagnostic characters: Small trees up to 10 m tall, dbh up to 8 cm. Bole

straight. Bark smooth, hooped, latex red. Twigs hollow, usually with ant

openings. Stipules recurved, very long and horn-like, up to 2.5 cm long,

completely surrounding the twigs. Leaves peltate, not lobed to shallowly 3-

lobed (cusped), under surface glaucous or not. Staminate inflorescences

branched. Fruits more than 2-locular, not spiny, but glandular banded,

seeds with conspicuous bright red aril.

Habitat & ecology: Lowland. In primary forest. Usually ant inhabited.

Distribution: Endemic to Borneo. Found south of the Mahakam river in

East Kalimantan.

17. Macaranga lowii King ex Hook.f. var. lowii (Fig. 14)

Diagnostic characters: Trees up to 20 m tall, dbh up to 16 cm. Bole straight.

Bark smooth, shallowly cracked, latex red. Inner bark thin, dark red. Twigs

solid, apically short hairy. Stipules erect, spine-like, usually crowded at

twig apex. Leaves not peltate, not lobed, penniveined, elliptic to obovate,

upper surface basely with two conspicuous macular glands, under surface

usually short hairy. Staminate inflorescences unbranched. Pistillate

inflorescences with large leaf-like bracts. Fruits 2-locular, spiny, sometimes

with long red stigmas.

Habitat & ecology: Up to 1350 m altitude. Mainly in the understorey of

primary forests but also in secondary forests. On well drained, clay-rich

and sandy loam soils.

Distribution: Southeast Asia, Sumatra, Borneo and the Philippines. Found

throughout East Kalimantan.

18. Macaranga lowii King ex Hook.f. var. kostermansii Airy Shaw (Fig. 15)

Diagnostic characters: Trees up to 15 m tall, dbh up to 15 cm. Bark smooth,

latex red. Twigs solid, glabrous. Stipules erect, spine-like, apically crowded

on twig, red. Leaves not peltate, not lobed, penniveined, elliptic to obovate,

upper surface with basally two dark red macular glands, under surface

Key to Macaranga and Mallotus of East Kalimantan 29

glabrous. Staminate inflorescences unbranched. Fruits 2-locular, spiny.

Habitat & ecology: Up to 150 m altitude. In the understorey of primary

forest but also in secondary forest. On slopes or ridges, along skid trails.

On sandy loam.

Distribution: Endemic to Borneo. Found throughout East Kalimantan.

19. Macaranga motleyana (Mull.Arg.) Mull.Arg. (Fig. 16)

Diagnostic characters: Trees up to 25 m tall, dbh up to 30 cm. Bole straight.

Bark smooth, hooped, glaucous, latex clear turning red. Twigs hollow,

glaucous, usually with ant openings. Stipules recurved, not completely

surrounding the twigs, green or red. Leaves peltate, deeply 3-lobed (unlobed

in seedlings), under surface usually slightly glaucous, glabrous. Staminate

inflorescences branched. Fruits more than 2-locular, not spiny, glaucous,

with glandular bands.

Habitat & ecology: Up to 400 m altitude. In primary forest in disturbed

sites (gaps) and in secondary forest, along river banks and roads. On dry

to swampy places. On limestone to red loamy clay. Usually ant inhabited.

Distribution: From Thailand and Indo-China southward into Peninsular

Malaysia, Sumatra and Borneo. Found throughout East Kalimantan.

20. Macaranga pearsonii Merr. (Fig. 17)

Diagnostic characters: Trees up to 35 m tall, dbh up to 65 cm. Bole straight.

Bark slightly fissured, hooped, lenticelled, latex red. Twigs solid to hollow,

glaucous, glabrous to hairy (when young). Stipules erect, caducous, apex

acute, green to reddish. Leaves deeply cordate to peltate, deeply 3-lobed

(unlobed in seedlings), base with conspicuous large glands along the margin,

under surface usually slightly glaucous, sometimes hairy. Staminate

inflorescences branched. Fruits 2-locular, not spiny.

Habitat & ecology: Up to 1200 m altitude. In primary forest in disturbed

sites (gaps) and secondary forest to scrub, along roads. Sometimes in

periodically flooded areas, usually on dry places. Quite often inhabited by

- ants:

Distribution: Endemic to Borneo. Found throughout East Kalimantan.

26 Gard. Bull. Singapore 52 (2000)

21. Macaranga pruinosa (Miq.) Muill.Arg. (Fig. 18)

Diagnostic characters: Trees up to 20 m tall, dbh up to 25 cm. Bark smooth,

hooped, latex red. Twigs solid, sometimes hairy. Stipules erect, semi-

persistent, apex rounded. Leaves deeply cordate to peltate, deeply 3-lobed,

base without conspicuously large glands along the margin, lower surface

velvety hairy. Staminate inflorescences branched. Fruits 2-locular, not spiny.

Habitat & ecology: Lowland. Primary and secondary forests, usually in

swamps or along streams, rarely also on dry ground.

Distribution: Peninsular Malaysia, Sumatra, and Borneo. Found throughout

East Kalimantan.

22. Macaranga puncticulata Gage. (Fig. 19)

Diagnostic characters: Small trees up to 10 m tall. Twigs solid to hollow,

when hollow often with very large ant openings (up to 2 mm) and the

swollen internodes strongly constricted at the nodes. Stipules erect,

caducous. Leaves peltate, not lobed, small, not wider than 11 cm but usually

much narrower, under surface glaucous, densely gland dotted. Staminate

inflorescences branched. Fruits more than 2-locular, not spiny.

Habitat & ecology: Lowland. In swamp forest. Sometimes ant inhabited.

Distribution: Peninsular Malaysia, Sumatra and Borneo. Found south of

the Mahakam river in East Kalimantan.

23. Macaranga recurvata Gage. (Fig. 20)

Diagnostic characters: Trees up to 30 m tall, dbh up to 45 cm. Twigs usually

solid, rarely hollow, glabrous. Stipules erect, caducous. Leaves peltate, not

lobed, under surface glaucous, densely gland dotted, glabrous. Staminate

inflorescences branched. Fruits 2-locular, not spiny.

Habitat & ecology: Up to 500 m altitude. In primary (Agathis) and secondary

forest. In swampy places. On acid, sandy soil.

Distribution: Peninsular Malaysia and Borneo. Found throughout East

Kalimantan.

Key to Macaranga and Mallotus of East Kalimantan 26

24. Macaranga repando-dentata Airy Shaw (Fig. 21)

Diagnostic characters: Shrubs to small trees up to 7 m tall. Twigs solid,

long hairy. Stipules erect, caducous, spine like. Leaves not peltate, not

lobed, penniveined, lamina longer than 7 cm, long hairy. Staminate

inflorescences unbranched. Pistillate inflorescences with bracts very large

and leaf like. Fruits 2-locular, spiny.

Habitat & ecology: Up to 300 m altitude. In the understorey of primary

forest. On sandy soils.

Distribution: Endemic to Borneo. In East Kalimantan only found in Kutai.

25. Macaranga tanarius (L.) Mull.Arg. (Fig. 22)

Diagnostic characters: Shrubs to trees up to 15 m tall, dbh up to 20 cm.

Bole straight. Bark smooth, hooped, latex red. Twigs solid. Stipules erect,

persistent, green to reddish. Leaves peltate, not lobed, upper surface basally

sometimes with several macular glands, under surface glabrous to velvety

hairy, thinly to densely gland dotted. Staminate inflorescences branched.

Fruits 2-locular, with few curling spines, glaucous.

Habitat & ecology: Up to 2100 m altitude. Usually in very disturbed

vegetation like scrub or abandoned shifting cultivation areas, often along

roads, or along streams. On clay loam, sandy, and sandstone soils.

Distribution: From India and southern China to Australia and New Guinea.

Found throughout East Kalimantan.

26. Macaranga trichocarpa (Rchb.f. & Zoll.) Miull.Arg. (Fig. 23)

Diagnostic characters: Shrubs-to 6 m tall, dbh up to 10 cm. Bark smooth,

orange brown, latex clear turning red. Twigs solid, hairy apically. Stipules

erect, caducous. Leaves not peltate, not lobed to sometimes slightly 3-

lobed (cusped), base cordate to attenuate, upper surface basally with two

conspicuous macular glands, densely short hairy (slightly sandpaper-like),

under surface densely short hairy. Staminate inflorescences unbranched.

Fruits 2-locular, spiny.

Habitat & ecology: Up to 500 m altitude. Usually in very disturbed secondary

forest, especially scrub or abandoned shifting cultivation areas, but also

present in gaps in primary forest. Often along forest edges and road sides

28 Gard. Bull. Singapore 52 (2000)

where it can form pure stands. On sandy loam, sandy clay and red-yellow

podsolic soils.

Distribution: From Myanmar and Indo-China, south into Peninsular

Malaysia, Sumatra and Borneo. Found throughout East Kalimantan.

27. Macaranga winkleri Pax & K.Hoffm. (Fig. 24)

Diagnostic characters: Shrubs to small trees up to 15 m tall, dbh up to 18

cm. Bole straight. Bark smooth, hooped, latex clear turning red. Twigs

hollow, strongly angular when young, usually with ant openings, red when

young. Stipules erect, conspicuously longer than wide, red. Leaves peltate,

not lobed, basal veins usually at a 90 or more degrees angle with the

midrib. Staminate inflorescences branched. Fruits 2-locular, very small (c.

3 mm diameter), not spiny.

Habitat & ecology: Up to 1800 m altitude. In disturbed sites (gaps) in

primary forest and in secondary forests, including very disturbed forest

(after fires). Often along forest edges, road sides, river sides, in land slips.

On yellow sandy loam. Usually ant inhabited.

Distribution: Endemic to Borneo. Found throughout East Kalimantan.

Key to Macaranga and Mallotus of East Kalimantan 29

Figure 1. Macaranga aétheadenia Airy Shaw. Habitus of pistillate plant and detail of staminate

inflorescence.

30 Gard. Bull. Singapore 52 (2000)

Figure 2. Macaranga bancana (Miq.) Miull.Arg. Habitus of pistillate plant and detail of

staminate inflorescence. The plant shown here has shallowly lobed leaves, but in the field the

leaves can also be very deeply lobed.

Key to Macaranga and Mallotus of East Kalimantan

Figure 3. Macaranga beccariana Merr. Habitus plus details of pistillate and staminate

inflorescences.

cy,

1cm

Gard. Bull. Singapore 52 (2000)

Figure 4. Macaranga brevipetiolata Airy Shaw. Habitus of pistillate plant and detail of young

staminate inflorescence.

Key to Macaranga and Mallotus of East Kalimantan

(PRY 98

Sart

ior Le}

fe CED e -

ay \Q eee

<6 O

Figure 5. Macaranga conifera (Zoll.) Mill.Arg. Habitus of pistillate plant and detail of

staminate inflorescence.

34 Gard. Bull. Singapore 52 (2000)

Figure 6. Macaranga costulata Pax & K.Hoffm. Habitus of pistillate plant and detail of

staminate inflorescence.

Key to Macaranga and Mallotus of East Kalimantan 35

Figure 7. Macaranga depressa (Miill.Arg.) Miill.Arg. forma strigosa Whitmore. Habitus of

pistillate plant and detail of staminate inflorescence.

36 Gard. Bull. Singapore 52 (2000)

ae. ae | |

-a

Figure 8. Macaranga gigantea (Rchb.f. & Zoll.) Miill.Arg. Habitus of pistillate plant and detail

of staminate inflorescence.

Key to Macaranga and Mallotus of East Kalimantan

Figure 9. Macaranga glandibracteolata S.J.Davies.

inflorescence.

Habitus and detail of staminate

Gard. Bull. Singapore 52 (2000)

Figure 10. Macaranga hosei King ex Hook.f. Habitus of pistillate plant.

Key to Macaranga and Mallotus of East Kalimantan

aah)

SL) j

.> SS

ae.

Figure 11. Macaranga hullettii King ex Hook.f. Habitus of pistillate plant and detail of

staminate inflorescence.

40 Gard. Bull. Singapore 52 (2000)

Figure 12. Macaranga hypoleuca (Rchb.f. & Zoll.) Miull.Arg. Habitus of pistillate plant and

detail of staminate inflorescence.

Key to Macaranga and Mallotus of East Kalimantan

Figure 13. Macaranga lamellata Whitmore. Habitus and details of full grown and young

pistillate inflorescences.

42 Gard. Bull. Singapore 52 (2000)

Figure 14. Macaranga lowii King ex Hook.f. var. Jowii. Habitus of pistillate plant and detail of

staminate inflorescence.

Key to Macaranga and Mallotus of East Kalimantan 43

Figure 15. Macaranga lowii King ex Hook.f. var. kostermansii Airy Shaw. Habitus of pistillate

plant and detail of fruit and staminate inflorescence.

44 Gard. Bull. Singapore 52 (2000)

Figure 16. Macaranga motleyana (Mill.Arg.) Mill.Arg. Habitus of pistillate plant and detail

of staminate inflorescence.

Key to Macaranga and Mallotus of East Kalimantan 45

rae EF

S>=

Figure 17. Macaranga pearsonii Merr. Habitus of pistillate plant and detail of staminate

inflorescence.

46 Gard. Bull. Singapore 52 (2000)

Figure 18. Macaranga pruinosa (Miq.) Mill.Arg. Habitus and details of pistillate and staminate

inflorescences.

Key to Macaranga and Mallotus of East Kalimantan

ALT)

Ree

Figure 19. Macaranga puncticulata Gage. Habitus of pistillate plant. This is a specimen with

solid stem. They can also have ant inhabited hollow stems with large openings (c.2 mm across).

48 Gard. Bull. Singapore 52 (2000)

Figure 20. Macaranga recurvata Gage. Habitus of pistillate plant, with young fruits.

Key to Macaranga and Mallotus of East Kalimantan

Figure 21. Macaranga repando-dentata Airy Shaw. Habitus of pistillate plant.

50 Gard. Bull. Singapore 52 (2000)

Figure 22. Macaranga tanarius (L.) Mill. Arg. Habitus of pistillate plant and detail of staminate

inflorescence.

Key to Macaranga and Mallotus of East Kalimantan

we. ey

Figure 23. Macaranga trichocarpa (Rchb.f. & Zoll.) Miill.Arg. Habitus of pistillate plant and

detail of staminate inflorescence.

52 Gard. Bull. Singapore 52 (2000)

Figure 24. Macaranga winkleri Pax & K.Hoffm. Habitus and details of staminate and pistillate

inflorescences.

Key to Macaranga and Mallotus of East Kalimantan 53

Mallotus Lour.

Shrubs to trees up to 35 m tall, usually dioecious. Indumentum simple to

tufted to stellate. Bark sometimes prominently hooped (rings around stem).

Wood soft to hard. Stipules mostly small, often caducous. Leaves alternate

to opposite (when opposite often unequal in size or shape), mostly small;

blades sometimes peltate, usually not lobed (sometimes 3-cusped),

penniveined to tripliveined to palmativeined, sometimes with hairy domatia,

usually with basal macular glands on leaf uppersides, often gland dotted

on leaf undersides, sometimes gland dotted on leaf uppersides.

Inflorescences axillary to terminal, racemes to thyrses to panicles. Staminate

inflorescences mostly unbranched thyrsoid racemes; bracts small, often

caducous; flowers 1—9 per node, sepals 3 or 4, disc rarely present, stamens

15 to 150, anthers 2-celled, basi- to dorsi-fixed, connective often widened,

pistil absent. Pistillate inflorescences mostly unbranched racemes; bracts

small, often caducous; flowers 1, rarely 2 per node, sepals 2—9, ovary (2-)

3(-4) locular, style long to short, apically splitting into (2—)3(-4) stigmas,

papillae plumose. Fruit a capsule, smooth to spiny to winged, usually

dehiscent into bivalved parts.

Key to the Mallotus Taxa

sete OI ac ar es arent npeneng sh aan can aawasienied cer mendes SudheeanscesmensneTvskssee aut voredes Z

Oe BIND Na cic eon ecinesl ch aetaapanvacnandentutuannanlvend-sttatenie 4

2a. Petiole insertion usually more than 5 mm from lamina base. Plants

often smelling of fenugreek when dried. Inflorescences never branched

2b. Petiole insertion usually less than 5 mm from lamina base. Plants

never smelling of fenugreek when dried. Staminate inflorescences

DIPS YE OE oR 2 ie et LAE TSP ARC ORE Ot SERRE eae ee ee ee ne 13

3a. Vegetative parts glabrous. Leaf under surface with conspicuous hair

CALS cite AU SoTL WAS ACRVCS sg oc. 8socacinsnegiindss Vaanssatecdene 3. M. floribundus

3b. Vegetative parts extremely hairy. Leaf under surface without

conspicuous hair tufts in axils of basal nerves.................. 6. M. lackeyi

4a. Leaves opposite, unequal, one of each pair extremely small (less than

1 cm) and stipule-like (check carefully because the leaves look alternate

Galea SUC Re NG thik cA AN St BS Palin naidevdall aay eemssvesuaete vie 5

4b. Leaves alternate to opposite, if opposite and unequal, then the smaller

52:

Sb.

6a.

6b.

fice

8a.

Sb.

9a.

9b.

10a.

10b.

lla.

11b.

124.

12b.

Gard. Bull. Singapore 52 (2000)

leaf cordate or the same shape as the larger leaf, never stipule-like ..

antbssee Gnesisavelrsicseasensbodesadten inne fe Slee talaga iam Ria ate aac ae ie 9

Petioles of large leaves less than diem lomg../. 2200... 2iesiee Gee 6

Petioles. of large leaves donger sian) ear: Ace ae ee ee 7

Vegetative: parisshattys int: .tees ee e.. noe 2. M. eucaustus

Vegetative parts elabroms (40 es or ieee 15. M. penangensis

Twigs, petioles and leaf under surface hairy .............. 17. M. stipularis

Twigs and petioles glabrous; leaf under surface glabrous to sparsely

hairy 2.2..Aste nec eee dea eatot (St). De ee ee 8

Petioles usually more than 1 mm thick, up to 5(—8) cm long. Leaves

ovate to rarely elliptic. Staminate inflorescences conspicuously

branched, both pistillate and staminate inflorescences with numerous

reddish-orange glands producing a sticky yellow excudate .................

Linn olla paigassiole s sda sal ng Ada NE Ab 2S, cea 4. M. griffithianus

Petioles usually less than 1 mm thick, up to 2(—3) cm long. Leaves

elliptic to obovate. Inflorescences never branched, without sticky yellow

OUGES gogo ls Slee ame ete re ee a 15. M. penangensis

Majority of leaves alternate, sometimes apically on twigs also opposite

ivane tached aah bacco te ah ta eeete uenR eeREE Tac ce fon oph ck dv xse's Gon tacgiace sane ae 10

Majority of Teaves ape Gsite o.oo oo cs cc orcs crete 15

Leaves Penniveimewre oa sey et belt caren ae 14. M. peltatus

Leaves basally tripinvswve ch co -c.cpetncs caves ocnceet0-ecsrsnaccedp tuned ipee mee ceaein 11

All vegetative parts densely hairy. Twigs, petioles and leaf under

surfaces mostly creamish brown, contrasting sharply with green upper

119 ib: & <n! «AER AIM D0) (9. / Uv NNDB MMPRE NE Ene hs vee 12

All vegetative parts more or less glabrous (except sometimes petioles).

Twigs, petioles and leaf under surfaces greenish, never creamish brown.

(Note: The three species in section Polyadenii can only be distinguished

with certainty if pistillate inflorescences are present) ............ eee 18

Leaf base with more than 2 macular glands ............... 16. M. repandus

Leaf base without or with Zimacular glands (0.0.0.2 ee eens 13

Leaves sometimes 3-cusped, base (cordate to) cuneate to attenuate,

upper surface basally always with 2 conspicuous macular glands,

Key to Macaranga and Mallotus of East Kalimantan 55

13b.

14a.

14b.

t5a.

15b.

l6a.

16b.

rye:

17b.

18a.

18b.

19a.

without a blackish area around them. Fruits not woolly, with few

robust spine-like appendages ...............eeseeeseeeeeeeeee 13. M. paniculatus

Leaves rarely 3-cusped, base rounded to truncate to cordate, rarely

attenuate, upper surface basally often with 2 inconspicuous macular

glands with a blackish area around them. Fruits woolly, smooth or

with numerous soft, spine-like appendages. ............eeeeeeeeeeeeeeeeeeeeees 14

Latex colourless, not turning black. Fruits without spine-like

om a ca SE OE 0) SSG OSE eT ON SO Be SES 8. M. macrostachyus

Latex turning black. Fruits with numerous soft, spiny appendages ....

OAL Soa Mirah BE oh a ee, She 10. M. mollissimus

Leaves opposite, strongly unequal in size. Small leaves clearly differing

in shape compared with the large leaves. Petioles of the small leaves

eee Nate bil, Wiha niaeer Pe eile eas. Sie es ae ena 16

Leaves all opposite or majority of leaves opposite, (slightly) unequal

in size. Small leaves more or less the same shape as the large leaves.

Petioles of the small leaves longer than 5 mm. ...............eeeeseeeeeeeeeees 17

Petioles of large leaves always longer than 10 mm. Large leaves basally

fiphyenica to rarely penniveiied ..................1......-.-...00-0-+0- 1. M. dispar

Petioles of large leaves 1—7 mm long. Large leaves penniveined ......

a ene ceed hse bn ot Giese to cea Meech neha es Sine ht 9. M. miquelianus

Vegetative parts usually glabrous (except sometimes petioles). Stipules

early caducous. Majority of leaves opposite, leaves not to slightly

differing in size or petiole length, upper surface usually densely gland

dotted (Note: The three species in section Polyadenii can only be

distinguished with certainty if pistillate inflorescences are present) ...

Vite END are 5s Md TAL INE SR SOLE Oa SEES 2 eee ED 18

Vegetative parts glabrous to densely hairy. Stipules persistent to

caducous. Leaves opposite, conspicuously differing in size and petiole

length, upper surface not densely gland dotted .........0.. eee 20

Petioles glabrous or with only a few scattered hairs. Pistillate pedicels

1.5—2.5(—4) cm long. Fruits with rounded carpels ..............eeeee

I RS cases na care te acne nshinceentnpetccsiccnaasaees 7. M. leucodermis

Petioles glabrous to hairy. Pistillate pedicels c. 0.5 cm long. Fruits

Eee ES OE ALACRA ea eon neg een cccencee cee nnees 19

Petioles glabrous or with only a few scattered hairs. Fruits with angular

Calpe, spme-like wines absent ...._........2:......-.--.:.....-.---. 12. M. muticus

56 Gard. Bull. Singapore 52 (2000)

19b. Petioles (glabrous to) short hairy. Fruits with spine-like wings, 1.5—3

Gm dO 20... 23. ..ncasninccnonmecnernntimeccieans ee ae ae 18. M. sumatranus

20a. Leaf base with two conspicuous macular glands placed on the first

pair of ves ...2Unic..can eee ores. ele ee 19. M. wrayi

20b. Leaf base with several (more than 2) sometimes inconspicuous macular

glands placed on the lamina, between the VeIMS ............eeeeeeeeeeereeeeee 21

21a. Leaves with second pair of veins originating c. half way up or higher

onthe lamina; .:2 2. cee ee, Ss eee 5. M. korthalsii

21b. Leaves with second pair of veins originating clearly less than half way

TEE Lamy C1111! 16, Seem neni. eLetter eae 11. M. moritzianus

1. Mallotus dispar (Blume) Mull.Arg. (Fig. 25)

Diagnostic characters: Shrubs or small trees up to 15 m tall, dbh up to 10

cm. Most parts usually hairy (stellate or tufted). Leaves opposite (alternate

when young), strongly unequal in size and shape; petioles of small leaves

less than 5 mm long, of large leaves more than | cm long; small leaves

cordate; large leaves tripliveined to penniveined, upper surfaces usually

without or with inconspicuous basal macular glands. Staminate inflorescence

unbranched, several flowers per node. Pistillate inflorescences with several

fruits developing, fruits spiny.

Habitat & ecology: Lowland up to 750 m altitude. In the understorey of

primary forest or in lightly disturbed secondary forest (selectively logged).

Sandy loam and limestone soils.

Distribution: From Thailand southward into Peninsular Malaysia, Sumatra,

Java, Borneo and the Philippines. Found throughout East Kalimantan.

2. Mallotus eucaustus Airy Shaw (Fig. 26)

Diagnostic characters: Trees up to 25 m tall, dbh up to 25 cm. Bole straight.

Bark smooth, hooped. Most parts densely short simple-haired. Leaves

distichous, opposite, strongly unequal in size and shape, small leaves reduced

to stipule-like structures without petioles; large leaves penniveined, upper

surface without basal macular glands. Staminate inflorescences unbranched,

one flower per node. Pistillate inflorescences with several fruits developing,

fruits short spiny.

Key to Macaranga and Mallotus of East Kalimantan ay

Habitat & ecology: Lowland up to 300 m altitude. In primary forest

understorey or in lightly disturbed secondary forest (selectively logged).

On sandy, and clayey loam soils.

Distribution: Endemic to Borneo. In East Kalimantan found north of the

Mahakam river.

3. Mallotus floribundus (Blume) Mull.Arg. (Fig. 27)

Diagnostic characters: Shrubs to trees up to 20 m tall, dbh up to 15 cm.

Strongly smelling of fenugreek when dried. More or less glabrous with

some scattered simple to stellate to tufted hairs. Leaves alternate to apically

pseudo-opposite, peltate, upper surface with several basal macular glands,

under surface with conspicuous hairy domatia at petiole insertion, often

glaucous, usually densely gland dotted. Staminate inflorescences

unbranched, several flowers per node. Pistillate inflorescences with several

fruits developing, fruits with long spines.

Habitat & ecology: In disturbed sites in primary forest (gaps) and in

secondary forest. Often along rivers or in swampy areas but also on dry

places. On sandy to sandstone soils.

Distribution: From Thailand and Indo-China to the Solomon Islands. In

East Kalimantan found north of the Mahakam river.

4. Mallotus griffithianus (Mull.Arg.) Hook.f. (Fig. 28)

Diagnostic characters: Trees up to 15 m tall, dbh up to 15 cm. Bole straight.

Bark smooth, hooped. Mostly glabrous with some scattered simple hairs.

Leaves opposite, strongly unequal in size and shape; small leaves reduced

to stipule-like structures without petioles; petioles of large leaves strongly

kneed distally, blade penniveined, upper surface without basal macular

glands. Staminate inflorescences branched, densely gland-dotted, glands

excreting yellow sticky fluid, one flower per node. Pistillate inflorescences

with several fruits developing, densely gland dotted, glands excreting yellow

sticky fluid, pedicels very long, up to 2 cm, fruits long spiny, each spine

with a terminal gland.

Habitat & ecology: Up to 360 m altitude. In understorey of primary forest

or in lightly disturbed secondary forest (selectively logged). On sandy,

loamy or tuff soils.

58 Gard. Bull. Singapore 52 (2000)

Distribution: Peninsular Malaysia and Borneo. Found throughout East

Kalimantan.

5. Mallotus korthalsii Mull.Arg. (Fig. 29)

Diagnostic characters: Shrubs to trees up to 15 m tall, dbh up to 15 cm.

Bark smooth. More or less glabrous to simple to stellate to tufted hairy.

Leaves opposite (alternate when young), unequal in size, tripliveined,

second pair of veins arising in the upper half of the lamina, upper leaf

surface with several basal macular glands on the lamina between the

secondary veins. Staminate inflorescences unbranched, several flowers per

node. Pistillate inflorescences with several fruits developing, spiny.

Habitat & ecology: Up to 2000 m altitude, usually much lower. In the

understorey of primary forest and in secondary forest. On sandy clay,

sandy loam, and limestone soils.

Distribution: Peninsular Malaysia, Borneo, and the Philippines. Found

throughout East Kalimantan.

6. Mallotus lackeyi Elmer (Fig. 30)

Diagnostic characters: Shrubs to small trees up to 10 m tall, dbh up to 10

cm. Plants smelling strongly of fenugreek when dried. Most parts densely

tufted to stellate haired. Bole crooked. Bark smooth to knobby to flaky.

Leaves alternate to apically pseudo-opposite, peltate, upper surface with

several basal macular glands, usually gland dotted, under surface densely

hairy, gland dotted. Staminate inflorescences unbranched, several flowers

per node. Pistillate inflorescences with several fruits developing, fruits short

spiny.

Habitat & ecology: Up to 1200 m altitude. In the understorey of primary

forest to secondary forest. Often along streams and rivers, on inundated

terrain but also on dry slopes. On sandy loam, clay and limestone soils.

Distribution: Borneo and the Philippines. Found throughout East

Kalimantan.

7. Mallotus leucodermis Hook.f. (Fig. 31)

Diagnostic characters: Trees up to 35 m tall, dbh up to 55 cm. More or less

glabrous with some scattered simple to stellate hairs. Steep buttresses

Key to Macaranga and Mallotus of East Kalimantan 59

present to absent. Bole straight. Bark rough to smooth, dippled, brittle,

with small flakes. Wood hard. Leaves alternate to rarely opposite, when

opposite then only slightly unequal in size, tripliveined, upper surface usually

with several basal macular glands, usually densely gland-dotted. Staminate

inflorescences unbranched, with several flowers per node. Pistillate

inflorescences with several fruits developing, pedicels very long (up to 4.5

cm), fruits smooth, globose carpels.

Habitat & ecology: Up to 1440 m altitude. In primary forest, sometimes in

secondary forest as a primary forest remnant species. On swampy as well

as dry places. On loamy soils, sometimes mixed with limestone.

Distribution: Peninsular Malaysia to New Guinea. Found throughout East

Kalimantan.

8. Mallotus macrostachyus (Miq.) Mull.Arg. (Fig. 32)

Diagnostic characters: Shrubs to trees up to 12 m tall, dbh up to 15 cm. All

parts densely flaky brown-creamish hairy. Bole not straight. Bark smooth,

lenticelled. Leaves alternate, often slightly peltate (up to 5 mm free base)

to cordate, upper surface usually with two inconspicuous basal macular

glands surrounded by blackish coloured stain when dried, green upper

surface contrasting sharply with brown-creamish hairy under surface.

Staminate inflorescences branched, with several flowers per node. Pistillate

inflorescences with several fruits developing, fruits smooth, brown-creamish

woolly.

Habitat & ecology: Up to 900 m altitude. Mainly in severely disturbed

secondary forest (after fire and shifting-cultivation), rarely in primary forest.

On sandy and sandy loam soils.

Distribution: From Thailand southward into Peninsular Malaysia, Sumatra,

and Borneo. Found throughout East Kalimantan.

9. Mallotus miquelianus (Scheff.) Boerl. (Fig. 33)

Diagnostic characters: Shrubs up to 5 m tall, dbh up to 5 cm. Almost

glabrous to densley simple to stellate to tufted hairy. Forming sprouts

along root-stalks. Leaves opposite, strongly unequal in size and shape:

petiole of small leaves up to 5 mm long, blade cordate, tripliveined, upper

surface with two to several large basal macular glands; petiole of large

leaves up to 7 mm long, blade obovate, penniveined, upper surface with

60 Gard. Bull. Singapore 52 (2000)

several large basal macular glands. Staminate inflorescences unbranched,

several flowers per node. Pistillate inflorescences with several fruits

developing, fruits short spiny.

Habitat & ecology: Up to 600 m altitude. In the understorey of primary

forest and in secondary forests (selectively logged). Often in swampy places

but also on well drained soils. On sandy and limestone soils.

Distribution: From Thailand southward into Peninsular Malaysia, Sumatra,

Borneo and the Philippines. Found throughout East Kalimantan.

10. Mallotus mollissimus (Geisel.) Airy Shaw (Fig. 34)

Diagnostic characters: Shrubs to trees up to 15 m tall, 15 cm diameter.

Latex clear, turning black. All parts densely flaky brown-creamish hairy.

Leaves alternate, often slightly peltate (up to 5 mm free base) to cordate,

upper surface usually with two inconspicuous basal macular glands

surrounded by blackish coloured stain when dried, green upper surface

contrasting sharply with brown-creamish hairy under surface. Staminate

inflorescences branched, with several flowers per node. Pistillate

inflorescences with several fruits developing, fruits with numerous soft

spines, brown-creamish woolly.

Habitat & ecology: Up to 1500 m altitude. Mainly in secondary forest

(logged, burnt), rare in primary forest. Along roads, streams and rivers.

On sandy loam soils.

Distribution: From Myanmar and Indo-China to New Guinea and Australia.

Found throughout East Kalimantan.

11. Mallotus moritzianus Mull.Arg. (Fig. 35)

Diagnostic characters: Shrubs to trees up to 10 m tall, dbh up to 10 cm.

Bark smooth. Usually simple to stellate to tufted hairy. Leaves opposite

(alternate when young), unequal in size, tripli- to penni-veined, second

pair of veins arising in the lower half of the lamina, upper leaf surface with

several basal macular glands on the lamina between the secondary veins.

Staminate inflorescences unbranched, several flowers per node. Pistillate

inflorescences with several fruits developing, spiny.

Habitat & ecology: Up to 1000 m altitude. Primary and secondary forest.

On stream banks, periodically inundated land, but also well drained land.

Key to Macaranga and Mallotus of East Kalimantan 61

On sandstone, clayey loam and sandstone soils.

Distribution: Peninsular Malaysia, Sumatra, Java, Lesser Sunda Islands,

and Borneo. Found north of the Mahakam river in East Kalimantan.

12. Mallotus muticus (Mull.Arg.) Airy Shaw (Fig. 36)

Diagnostic characters: Trees up to 35 m tall, dbh up to 70 cm. More or less

glabrous with some scattered simple to stellate hairs. Buttresses up to 7 m

tall, 1 m wide at base. Bole straight, slightly fluted. Bark smooth, flaky.

Wood hard. Leaves alternate to rarely opposite, when opposite then only

slightly unequal in size, tripliveined, upper surface usually with several

basal macular glands, usually densely gland dotted, under surface often

with hairy domatia. Staminate inflorescences unbranched, with several

flowers per node. Pistillate inflorescences with several fruits developing,

pedicels short (up to c. 5 mm long), fruits smooth, indehiscent, angular

carpels.

Habitat & ecology: Up to 500 m altitude. In primary forest, swamp forest

and in secondary forest as a primary forest remnant. Often on periodically

inundated or swampy terrain, but also on dry terrain. On clay rich to sandy

soils.

Distribution: Peninsular Malaysia, Sumatra and Borneo. Found throughout

East Kalimantan.

13. Mallotus paniculatus (Lam.) Miuill.Arg. (Fig. 37)

Diagnostic characters: Shrubs to trees, up to 25 m tall, dbh up to 25 cm.

Bole straight. Bark cracked, lenticellate, latex red. All parts densely flaky

brown-creamish hairy. Leaves alternate, rarely slightly peltate (up to 5 mm

free base), usually tripliveined, base attenuate to obtuse, upper surface

usually with two conspicuous basal macular glands, not surrounded by

blackish coloured stain when dried, green upper surface contrasting sharply

with brown-creamish hairy under surface. Staminate inflorescences

branched, with several flowers per node. Pistillate inflorescences with several

fruits developing, fruits with few spine-like appendages, short brown-

creamish hairy.

Habitat & ecology: Up to 1500 m altitude. Mainly in heavily disturbed

secondary forest (after fire, abandoned shifting-cultivation), rarely in

primary forest. Along roads, streams, on land slides. On sandy clay soils.

62 Gard. Bull. Singapore 52 (2000)

Distribution: From south China to New Guinea and Australia. Found

throughout East Kalimantan.

14. Mallotus peltatus (Geisel.) Mull.Arg. (Fig. 38)

Diagnostic characters: Shrubs to trees up to 12 m tall, dbh up to 10 cm.

Strongly smelling of fenugreek when dried. More or less glabrous with

some scattered simple to stellate to tufted hairs. Leaves alternate to apically

pseudo-opposite, not peltate (outside Kalimantan often also peltate),

penniveined, upper surface with several basal macular glands, under surface

with conspicuous hairy domatia along the midrib. Staminate inflorescences

unbranched, several flowers per node. Pistillate inflorescences with several

fruits developing, fruits with long spines.

Habitat & ecology: Up to 450 m altitude. In primary and secondary forests.

Often along streams and rivers, along roads, and inundated terrains. On

sandy loam, and limestone soils. ;

Distribution: From east India and south China to New Guinea. Found

throughout East Kalimantan.

15. Mallotus penangensis Mull.Arg. (Figs 39 and 40)

Diagnostic characters: Trees up to 25 m tall, dbh up to 30 cm. Bole straight.

Bark smooth, hooped. More or less glabrous with some scattered short

simple hairs. Leaves opposite, strongly unequal in size and shape, small

leaves reduced to stipule-like structures without petioles; petioles of large

leaves very variable in length (from 1 to 4.5 cm long), blade penniveined,

upper surface without basal macular glands, under surface sometimes with

hairy domatia. Staminate inflorescences unbranched, one flower per node.

Pistillate inflorescences usually with only terminal fruit developing, fruits

long spiny.

Habitat & ecology: Up to 1200 m altitude. In the understorey of primary

forest and in lightly disturbed secondary forest (selectively logged). On

sandy loam or (red) clay soils, also found on limestone.

Distribution: From Thailand to New Guinea. Found throughout East

Kalimantan.

Key to Macaranga and Mallotus of East Kalimantan 63

16. Mallotus repandus (Willd.) Mull. Arg.

Diagnostic characters: Shrubs up to 3 m tall. Most parts densely brown-

creamish simple to stellate hairy. Leaves alternate, tripliveined, upper

surface with several basal macular glands, under surface densely gland

dotted. Staminate inflorescences branched, several flowers per node.

Pistillate inflorescences with several fruits developing, fruits 2-locular,

smooth.

Habitat & ecology: Up to 420 m altitude. Mainly in secondary forest and

scrub.

Distribution: India and Sri Lanka to New Guinea and Australia. Found

north of the Sankulirang limestone mountain range in East Kalimantan

(Berau).

17. Mallotus stipularis Airy Shaw (Fig. 41)

Diagnostic characters: Trees up to 20 m tall, dbh up to 20 cm. Bole straight.

Bark smooth, hooped. Most parts with long simple hairs. Stipules large,

margins with long simple hairs, apex usually round. Leaves opposite,

strongly unequal in size and shape, small leaves reduced to stipule-like

structures without petioles; blade penniveined, upper surface without basal

macular glands, under surface usually with hairy domatia. Staminate

inflorescences unbranched, one flower per node. Pistillate inflorescences

with several fruits developing, fruits short spiny.

Habitat & ecology: Up to 950 m altitude. In the understorey of primary

forest and in lightly disturbed secondary forest (selectively logged). On

clay rich soils.

Distribution: Thailand, Sumatra and Borneo. Only found north of the

Sankulirang limestone mountain range in East Kalimantan.

18. Mallotus sumatranus (Migq.) Airy Shaw (Fig. 42)

Diagnostic characters: Trees up to 12 m tall, dbh up to 20 cm. More or less

glabrous to simple to stellate hairy on petioles. Leaves usually opposite,

only slightly unequal in size, tripliveined, upper surface usually with several

basal macular glands, usually densely gland dotted, under surface often

with hairy domatia. Staminate inflorescences unbranched, with several

flowers per node. Pistillate inflorescences with several fruits developing,

64 Gard. Bull. Singapore 52 (2000)

pedicels short (up to c. 5 mm long), fruits smooth, indehiscent, with long

pointed wings up to 3 cm long.

Habitat & ecology: Up to 50 m altitude. In primary forest, often in swamp

forest.

Distribution: Sumatra and Borneo. Found along the Mahakam river in

East Kalimantan (Kuta).

19. Mallotus wrayi King ex Hook.f. (Fig. 43)

Diagnostic characters: Shrubs to trees up to 19 m tall, dbh up to 15 cm.

Bole sometimes crooked, branching midway up. Bark smooth. From almost

glabrous to densely simple to stellate to tufted hairy. Leaves opposite

(alternate when young), unequal in size, tripliveined, second pair of veins

arising in the lower half of the lamina, upper leaf surface with two

conspicuous basal macular glands on the first secondary veins. Staminate

inflorescences unbranched, several flowers per node. Pistillate inflorescences

with several fruits developing, spiny.

Habitat & ecology: Up to 1200 m altitude. In primary and slightly disturbed

secondary forest (selectively logged). However, the species sprouts after

fire and can thus also be present in burnt forest. On swampy to well

drained, sandy to loamy to clayey soils.

Distribution: Peninsular Malaysia, Sumatra and Borneo. Found throughout

East Kalimantan.

nga and Mallotus of East Kalimantan

Key to Macara

Figure 25. Mallotus dispar (Blume) Miill.Arg. Habitus of pistillate plant and detail of staminate

inflorescence

66 Gard. Bull. Singapore 52 (2000)

——————

Figure 26. Mallotus eucaustus Airy Shaw. Habitus of pistillate plant and detail of staminate

inflorescence.

Key to Macaranga and Mallotus of East Kalimantan 67

—_

Figure 27. Mallotus floribundus (Blume) Miill.Arg. Habitus of pistillate plant and details of

staminate flowers, a fruit and the domatia at the petiole on the leaf underside.

68 Gard. Bull. Singapore 52 (2000)

Figure 28. Mallotus griffithianus (Miill.Arg.) Hook.f. Habitus of pistillate plant and detail of

staminate inflorescence.

Key to Macaranga and Mallotus of East Kalimantan

Figure 29. Mallotus korthalsii Mill.Arg. Habitus of pistillate plant.

70 Gard. Bull. Singapore 52 (2000)

nei

ie eS

—

(CF

ll =

ese

\ \

neg

eE Sa

te.

ne \ ee

ye iz Be i:

rats

/ VS.

Figure 30. Mallotus lackeyi Elmer. Habitus of pistillate plant and detail of staminate

inflorescence.

Key to Macaranga and Mallotus of East Kalimantan va

Figure 31. Mallotus leucodermis Hook.f. Habitus of pistillate plant and detail of staminate

inflorescence.

72 Gard. Bull. Singapore 52 (2000)

Figure 32. Mallotus macrostachyus (Mig.) Mill.Arg. Habitus of young staminate plant and

detail of pistillate inflorescence.

Key to Macaranga and Mallotus of East Kalimantan 73

Figure 33. Mallotus miquelianus (Scheff.) Boerl. Habitus of pistillate plant and detail of

staminate inflorescence.

74 Gard. Bull. Singapore 52 (2000)

Figure 34. Mallotus mollissimus (Geisel.) Airy Shaw. Habitus of pistillate plant and detail of a

part of the staminate inflorescence.

Key to Macaranga and Mallotus of East Kalimantan 73.

Figure 35. Mallotus moritzianus Miill.Arg. Habitus of pistillate plant and detail of staminate

inflorescence.

76 Gard. Bull. Singapore 52 (2000)

Figure 36. Mallotus muticus (Mill.Arg.) Airy Shaw. Habitus and detail of pistillate

inflorescence.

Key to Macaranga and Mallotus of East Kalimantan TF.

a =:

sea

pee

Figure 37. Mallotus paniculatus (Lam.) Miill.Arg. Habitus of pistillate plant and detail of

staminate inflorescence.

78 Gard. Bull. Singapore 52 (2000)

Figure 38. Mallotus peltatus (Geisel.) Mill.Arg. Habitus of pistillate plant and detail of

staminate inflorescence.

Key to Macaranga and Mallotus of East Kalimantan 79

a =

Figure 39. Mallotus penangensis Mill.Arg. Habitus of pistillate plant and detail of staminate

inflorescence.

10) Gard. Bull. Singapore 52 (2000)

Figure 40. Mallotus penangensis Mill.Arg. Habitus of pistillate plant and detail of staminate

inflorescence.

Key to Macaranga and Mallotus of East Kalimantan 81.

Figure 41. Mallotus stipularis Airy Shaw. Habitus of pistillate plant and detail of staminate

inflorescence.

$2 Gard. Bull. Singapore 52 (2000)

Figure 42. Mallotus sumatranus (Miq.) Airy Shaw. Habitus of pistillate plant.

Key to Macaranga and Mallotus of East Kalimantan 83.

Figure 43. Mallotus wrayi King ex Hook.f. Habitus of pistillate plant and detail of staminate

inflorescence.

84 Gard. Bull. Singapore 52 (2000)

Acknowledgements

The identification and nomenclature problems we faced while working on

Macaranga could not have been solved without the invaluable help of T.C.

Whitmore and S.J. Davies. We are very grateful for their help. We would

also like to thank Birgitte Fiala and Jan Frits Veldkamp for their comments

on earlier versions of this manuscript. This study is supported by a grant

of the Netherlands Foundation for the Advancement of Tropical Research

(WOTRO), within the Priority Programme ‘Biodiversity of Disturbed

Ecosystems’.

References

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Hadiah, D.J. Middleton, C. Pendry, M. Pinard, M. Warwick and K.S.

Yulita, 1998. Manual of the Larger and More Important Nondipterocarp

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KeBler P.J.A., K. Sidiyasa, Ambriansyah and Arifin Zainal, 1995. Checklist

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Pax, F. and K. Hoffmann, 1914. Euphorbiaceae - Acalypheae -

Mercurialinae. In: A. Engler (ed.): Das Pflanzenreich IV. 147. vii, Wilhelm

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at the Wanariset Herbarium East Kalimantan - Indonesia. The

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Indonesia.

Whitmore, T.C., 1975. Macaranga. In: Airy Shaw, H.K. (ed.) The

Key to Macaranga and Mallotus of East Kalimantan 85.

Euphorbiaceae of Borneo. Kew Bulletin. Additional Series 4: 140—159

Index of Exsiccatae

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ADCP GMR TOW UE VAT. WOSTCTHIONISID «.c080ecs.cetccevsecesvesaaccettasnesetosneettonchecegnuanesss 18

Res PIED EE TIO ICY OPT fete arc Seo sects dhs intandatecaavent tetvanessonnnecestacccaseannerce 19

Peace REA ON CS SNIITEE Tri. Aare Nan Soren colcts relic dhtcessprsccbereastsincrnunctotuoornesetesns 20

MIN eeaN ARNG PE OTHER IVOSII bee eas FOP oes fester ccs iee ace relsesen tetas cenactetsitheseseeccsaeraascessigsens 21

A aM INTE TOTNES GULVET IM: bluse. Betthchc tet caktbaseassadeedhs vdehrracditesonhecytaeeecedendbeadenes 22

TT ES Eg glia 0 id Oe ie eT cee Cie er pe ee a ee le 23

ICRF APL ATOM ATO OMILAIEE - 350s. css beindisy (oohincSal> cad a4 ede odsa¥osedebiga es. adldegssadvanes 24

Tl IIA OSA AIGA IRES crite to, ts Ha as 0h lov5 sf ctlpn Rau DL 4GR sh CN lon MOG Hs ead betinn 6 BUR o0 Fa eeb 25

VE CRP AME PE CII CII ain as ag assays Sipe onr seiirvsYpad ads Pod ean epgg sath dath« MMe Lo Re 26

PI ets) a NA PUES) OMIT etek eh i! ah ces ice caylee novi cc sae Puce oxean de qitenedessandehdnaeasiioncse 21

TRS SUICRER 96 ak hs: A oI Re te ee LU oe 9 a Cn 28

Es ORI CURES RPI DS AMS, Mere SACD rade he ee UN ais, AR tala adds bid eTetecetbesecvesese 29

RIM AS SOELIIPITO SrA. Lee yite te hes ead tbe bak sibs eke Fubets sa cavessezeadscenatt 30

HOE CUTIE RE THEA 0. US Ore ed Rho Ay 6 ok na a Ct Oe 31

NOTES GES 0 | iene’ A ie Re ee ne Oe, Nee | ee ee ee 32

FR ME Oy eg ae See ets ca a civ it a el SLRIes nS oid nga dons uve <ntaciuasrtoenand 33

Ee II eee Pd forecasts dae ee ser cree os each od eo hos bdeovenaanenssteses meet 34

VAG OS RCT OSE NCHA Y Ut; IAN AO cB REYES iy acy UNe <eak 0 ONE. acids vacsacdede- week iars 35

NE OLS TACT GIES A OTN AE I ides 4G oo Ahi Saree Wet Tade casagrinaab don d 36

EOE ERIE UU ESS TIRIITS 02s on og ndicv eh wwe cane deicess janajdacssvenesndabesecsesecacseseededvetvonsees 37

86 Gard. Bull. Singapore 52 (2000)

Mallotus. morizianus, nics ieee ae, 6 2a 38

MeallOtes PGR OUS sees ives sacveonsonneeaneedicoues btiemaniraecde siete. teusbidaaia mecca een aan 39

Matllotaspaniculatis ccc RE Be Le eee 40

Mallotus peltateis 8. dicate I A EL a ee 41

Mallotus: penangensis: >. .cii0...01.0 ie eR Re ae, 42

Mallotus repandus 4.2... i RL ADA 2 a 43

Matloras stipularis is .J..d eG A ea de 44

Mallotus' sumatranus nc ncuun ii Lak cee 8 45

Matlotus wrayl oi...2%. 0. A a 46

Afandi Ma’roef AM 216: 36 — Ambri et al. AA 24: 34; 48: 16; 84: 16; 400:

38; 446: 45: 477: 26; 1187: 40; 1190: 40; 1200: 27; 1474: 38; 1592: 26; 1606:

40; 1616: 25; 1619: 25; 1628: 37; 1720: 35; 1742: 42 — Ambri et al. W 2:

Ad: 10: 10; 24: 2: 57: 17; 72: 3: 78: 33; 3: 42: 99: 42. oe A 6G. eS ee

25: 333: 42; 335: 42; 636: 13; 767: 27; 1001: 10; 1009: 14-.1010: 21: 1024.7

— Ambriansyah AA 663: 36; 689: 5; 716: 3; 774: 46; 1025: 5; 1155: 25;

1494: 14; 2064: 7; 2152: 34 — Ambriansyah et al. AA 186: 26; 187: 2; 207:

33; 242: 41; 288: 20; 675: 33; 991: 41; 1159: 40; 1676: 13; 1678: 33; 1696: 40

— Ambriansyah et al. Berau 589A: 29; 598: 31; 602: 44; 805: 33; 883: 38

— Ambriansyah et al. W 943: 14; 950: 2; 971: 25; 980: 14; 984: 5 —

Arbainsyah AA 1849: 40; 1869: 27; 1928: 27; 1934: 40 — Arbainsyah er

al. AA 2033; 2045: 25 — Arifin AA 627: 7 — Arifin et al. AA 1643: 26;

1645: 20; 1653: 38; 1708: 37; 1753: 35; 1787: 41; 1804: 26 — Arifin Berau

557: 3; 559: 44 — Arifin et al. Berau 556: 3; 561: 44; 563: 29; 725: 34; 999:

31.

Balgooy 5718: 26; 5902 A: 7; 6076: 7; 6124: 7 — Balgooy et al. 5882: 25;

5883: 19; 5884: 14; 5902: 35; 5912: 10; 5916: 14; 5920 A: 20; 5948 B: 20;

5965: 42 — bb 2126: 37; 11895: 34; 12129: 42; 16188: 5; 16901: 5; 25145:

10; 34248: 5. ;

Endert 1498: 45; 1534: 26; 1606: 40; 1845: 10; 1866: 36; 1989: 45; 2089: 38:

2196: 30; 2200: 27; 2434: 10; 3028: 41; 3089: 13; 3266: 15; 3488: 33; 3735:

46; 4021: 41; 4032: 27; 4081: 13; 4087: 41; 4671: 13; 4694: 33; 4772: 41;

4811: 26; 4812: 26; 4831: 29; 4919: 31; 4959: 19; 4982: 5; 5096: 38; 5098: 38;

5114: 36; 5153: 33; 5158: 20; 5167: 38; 5173: 46; 5185: 36; 5267: 8.

Forman 445: 46; 456: 46; 460: 26.

Geesink 8904: 25; 8927: 12; 8956: 6; 9044: 7; 9299: 13; 9315: 8 — Goverse et

al. Berau 429: 40; 430: 40; 461: 3; 473: 31.

Key to Macaranga and Mallotus of East Kalimantan 87

Hamdi AA 2113: 19; 2119: 40.

Iwatsuki et al. B 7216: 14; 7301: 36; 7302: 36.

Kato et al. B 4772: 46; 4781: 41; 5363: 41; 5524: 32; 6122: 17; 6531: 41; 10212:

aa 10214 1; 11254: 7. 11441: 46: 11760: 28; 11827: 8;°11888: 38 —

Kartawinata 1317: 26; 1475A: 6 — Kessler PK 611: 13; 619: 13 — Kessler

et al. Berau 28: 25; 31: 25; 36: 11; 38: 11; 52: 14; 56: 1; 61: 19; 81: 40; 101:

mea 2 tu tu az: 157. A 156.14 1o4: 29° 174: 29: 272: 14: 277:

20; 293: 19; 194: 29; 200: 31; 201: 31; 255: 38; 270: 40; 296: 19; 323: 3; 357:

oe 27. 3/0: 29-376 5, 37%: 16, 004: 42; 615: 29°'016: 18; 616: 31: 701:

38: 740: 17: 856: 46 — Kessler et al. PK 745: 25; 786: 25; 787: 25: 789: 37:

794: 37; 796: 37; 825: 40; 843: 19; 863: 26; 875: 40; 899: 20; 900: 10; 910: 27;

053: 27: 973: 27: 977: 20; 989: 20; 1002: 20; 1054: 40; 1077: 25; 1089: 40:

100:.33571 102: 37; 1114:-25;.1130: 25; 1131:.26;1159: 25:°1164:,40; 1203:

S72 31, ze. 1244" AG; 1291725: 1292° 25; 1298: 35: 1302: 37:°1303: 37;

io Ze 25rd 325: 20; 1329: 4, 1333: 1; 1350237:-1351¢-37;.1388: 26; 1405: 38;

1435: 40; 1446: 20; 1960: 38; 1968: 36; 1972: 19; 1988: 19; 1989: 37; 2031:

27: 2038: 6; 2071: 40; 2076: 40; 2119: 26; 2710: 22: 2281: 36; 2285: 25; 2347:

20; 2356: 40; 2383: 40 — Kostermans 217: 36; 4062: 17; 4121: 17; 4160: 17;

4321: 13; 4344: 18; 4940: 42; 5011: 19; 5344: 36; 5512: 42; 5616: 5; 5686: 34;

5950: 42; 6011: 36; 6023: 19; 6026: 6; 6229: 42; 6356: 42; 6428: 17; 6433: 4;

6685: 31; 6693: 42; 7019: 31; 7563: 42; 7625: 42; 8632: 42; 9134: 42; 9239: 9;

9320: 29; 9637: 36; 10026: 26; 10065: 17; 10155: 42; 10156: 18; 10178B: 42;

TOUS: 42: 10229: 42: 10531: 27: 10696: 36; 12644: 38; 12691: 2: 13113: 23:

13148: 24; 13156: 27; 13762: 28; 13985: 38; 14053: 42; 21010: 38; 21017: 41;

21077: 46; 21092: 46; 21132: 38; 21137: 41; 21262: 40; 21264: 33; 21345: 28;

Prat: 19° 71445 Se. 2154123: 21548: 38; 21571: 36; 21591- 33; 21595: 32:

21596: 36; 21643: 7; 21716: 38 — Kramadibrata 137: 35; 248: 19.

Leeuwenberg et al. 13107: 40 — Leighton 72: 17; 73: 42; 92: 46; 218: 26; 264:

27; 373: 26; 382: 40; 394: 14; 430: 36; 512: 34; 570: 10; 762: 33; 885: 25; 912:

2° 1025: 17: 1026: 42; 1084: 20: 1091: 42.

Marsemi 14: 26 — McDonald et al. 3456: 46 — Meijer 1915: 26; 2091: 36;

2452: 38 — Mogea 2668: 27 — Murata et al. B 455: 26; 479: 26; 572: 35;

ToT: Wa: 941- 46: 1123: 27. 1277: 27: 3028: 3S.

Nedi 733: 36.

Paymans 20: 14.

Rahayu 140: 20 — Ramlanto 16: 40; 28: 17.

88 Gard. Bull. Singapore 52 (2000)

Sauveur 3: 42; 25: 17; 26: 17; 57: 14; 100: 42; 132: 42 — Sidiyasa S 298: 40;

436: 36; 438: 33; 440: 28; 476: 17; 502: 27; 525: 28; 531: 27; 632: 5; 639: 40;

670: 20; 754: 18; 756: 18; 762: 17; 790: 21; 805: 13; 823: 5; 1140: 41; 1141:

Al: 1157: 33;,1157-A: 463.1171: 33; 1172; Sh; LES As 19-4214: 38 ie

35; 1246: 42; 1248: 17; 1282: 35 — Sidiyasa et al. S 557: 42; 829: 2 — Slik

Berau 578: 26; 579: 26; 580: 26; 581: 31; 582: 19; 584: 46 — Slik W 1043: 2;

1044: 17; 1045: 7; 1046: 2; 1047: 26; 1048: 17; 1049: 2; 1050: 19; 1051: 21:

1052: 13; 1053: 25; 1054: 40; 1055: 40; 1056: 5; 1057: 20; 1058: 35 — Slik er

al. FS 7: 33; 12: 28; 13: 46; 14: 36; 15: 36; 17: 33; 18: 41; 21: 27 — Slooten

2137: 36 — Soetisna 16: 40 — Soewanda 35396: 42.

TFB 1243: 40; 1254: 27.

Ueda et al. B 11586: 13.

Valkenburg JVV 1208: 35; 1261: 13 — Valkenburg et al. JVV 1065: 27.

Wiriadinata 289: 36; 324: 42: 612: 46; 653: 10; 694: 27; 715: 46; 811: 46; 816:

32; 1146: 25; 1182: 20 — Wiriadinata et al. 63: 19; 1073: 10.

Gardens’ Bulletin Singapore 52 (2000) 89-100.

An Account of Neotenic Species of Rhaphidophora

Hassk. (Araceae-Monsteroideae-Monstereae) in New

Guinea and Australia

P.C. Boyce! AND J. BOGNER?

‘Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, U.K.

*Botanischer Garten, Menzinger Strafe 63, Miinchen, D-80693, Germany

Abstract

An account of the neotenic Rhaphidophora Hassk. species in New Guinea and Australia is

presented as a precursor to the Flora Malesiana and Flora of Australia accounts. Three

species, two (R. hayi, and R. okapensis) new to science, are described, together with a brief

discussion of neoteny in monsteroid aroids. A key is provided. All species are illustrated.

Introduction

Rhaphidophora in New Guinea, the western tropical Pacific, and Australia

comprises in excess of 30 species of which all, except R. korthalsti Schott,

are endemic to the region (Hay, 1981, 1990, 1993; Hay et al., 1995; Nicolson

1978, 1979). Work for the Flora of Australia Araceae account (Hay, in

press) has revealed that the species commonly referred to as R. pachyphylla

K. Krause comprises two distinct taxa, one hitherto undescribed. It is this

undescribed element that is present in Australia, which was reported by

Hay (1993) and illustrated by Jones & Gray (1988: 322) as R. pachyphylla.

During preparation of the description of this new taxon, to be called R.

hayi, it became apparent that a remarkable second undescribed species, so

far known only from the Eastern Highlands of Papua New Guinea, was

represented among specimens on loan to K. This is here described as R.

okapensis.

Although Rhaphidophora hayi is certainly distinct from R.

pachyphylla, it is still not clear whether R. pachyphylla as here now

redefined represents a single taxon. A specimen from high altitude in New

Guinea differs, aside from representing an extreme altitudinal increase on

the other specimens seen, in a shortly stipitate spadix, and a markedly

-rounded spathe apex. More specimens of ‘R. pachyphylla’ from higher

elevation are required to resolve this plant’s status.

This is one of a series of papers intended to present a complete

alpha-taxonomy of the genus Rhaphidophora in Asia. Accounts for the

Himalaya, Thailand and Indochina, Peninsular Malaysia and Singapore

90 Gard. Bull. Singapore 52 (2000)

(Boyce, 1999), the Indonesian archipelago excluding Borneo and Irian

Jaya, the Philippines (Boyce, 2000), Borneo, and Papuasia are being

prepared and will be published separately. All morphological terms

employed follow Stearn (1992).

Neoteny in Monstereae

The three Rhaphidophora species discussed here display neotenic habit:

plants retaining juvenile vegetative morphology at sexual maturity. Neoteny

in Monstereae appears to be of two types, obligate or facultative, defined

as to whether a species is habitually neotenic, or whether certain individuals

display a neotenic habit, and others do not.

The obligate species can be further divided into those that undergo

no change in their vegetative morphology at flowering (complete neoteny),

and those that produce a token adult phase at flowering, e.g., a shingling

species producing slightly scattered leaves on flowering shoots.

Rhaphidophora okapensis belongs to the first group, R. hayi to the second.

Complete neoteny is a feature of a very few species of Rhaphidophora.

Aside from R. okapensis, only R. latevaginata M. Hotta (Borneo) has this

character. R. /atevaginata differs from R. okapensis in flowering at the tips

of adherent shingling shoots, by the much larger, oblong, truncate-based

leaf laminas, and by the considerably larger inflorescence. Elsewhere in

the Monstereae only Scindapsus lucens Bogner & P.C. Boyce (Sumatera,

Peninsular Malaysia), and neotropical Monstera tuberculata Lundell are

known to be completely neotenic.

Facultative neoteny is so far known to occur in the widespread R.

beccarii (Engl.) Engl. (neotenic plants lacking the leaf lamina division

prominent in the typical form have been described as R. fluminea Ridl.),

and is occasionally observed in Epipremnum pinnatum (L.) Engl.

It appears from the above that neoteny in the Monstereae may have

arisen independently several times.

Key to Neotenic Rhaphidophora in New Guinea and Australia

la. Leaf base cordate on flowering shoots. Stylar region conical..............

hii A een pep OAR A atiesichi a teh whi Pole. nun 2. R. okapensis

1b. Leaf base truncate, acute or cuneate on flowering shoots. Stylar region

RO UITRCAC. sais ccnmenibcc tec pen ceenise sare tate cei ec ee ee ge 2

2a. Flowering shoots with base of leaf lamina truncate; spadix stipitate,

stigma elongated, longitudinally orientated. Plant with disarticulating

Neotenic Rhaphidophora in New Guinea and Australia 91 -

side shoots functioning as vegetative propagation units ..... 1. R. hayi

2b. Flowering shoots with base of leaf lamina acute to cuneate; spadix

sessile, stigma punctiform. Plant without disarticulating side shoot ..

ih Roe ee ny ens - oer Sa ER 3. R. pachyphylla

1. Rhaphidophora hayi P.C. Boyce & Bogner, sp. nov.

Rhaphidophora pachyphylla auctt. Austr. non K. Krause

A R. pachyphylla \aminis latioribus basi truncatis, stigmatibus maioribus

elatis oblongis in spadice longitudinaliter positis; ramis lateralibus brevibus

liberis leviter in frangentibus ad propagationem vegetativam pertinentibus

differt— TYPUS: Australia, Queensland, Cooroo Lands, north Johnstone

River, near Innisfail, Nov 1965, Webb & Tracey 7066 (BRI, holo).

Figure 1.

Moderate-sized, slender to somewhat robust, semi-leptocaul, homeophyllous

neotenic liane to 5 m; seedling stage a non-skototropic shingling juvenile

shoot; pre-adult plants forming small terrestrial colonies; adult shoot

architecture comprised of clinging, physiognomically unbranched, mostly

densely leafy, sterile stems and abbreviated, free, fertile stems; stems

rectangular-terete in cross section, widest side prominently convex, smooth,

dark green, without prophyll and cataphyll fibre but with very thin, adherent,

petiolar sheath tissue, internodes to 8 x | cm, separated by slight + straight

scars, older stems sub-woody; flagellate foraging stems weakly developed,

usually at least partially leafy and mostly replaced by short, readily

disarticulating free side shoots functioning as vegetative propagation units;

clasping roots arising from the internodes, prominently pubescent; feeding

roots c.3 mm diam., brown, minutely pubescent, sparsely lenticellate; leaves

distichous, shingling on adherent shoots, densely arranged or slightly

scattered on free shoots, scattered leaves with internodes between carrying

a prominent cataphyll of short duration; cataphylls and prophylls

membranous, soon drying and falling; petiole deeply grooved, 1-2 x 0.2-0.3

cm, smooth, apical and basal genicula barely visible; petiolar sheath

prominent but soon drying and adhering to stem while falling from petiole,

membranous, ligulate, margins of ligule fused, the ligule extending up to 3

cm above base of lamina and enclosing shoot apex; lamina broadly to

narrowly ovate-elliptic, stiffly coriaceous, base truncate to cuneate or cordate

(the last not on flowering shoots), and briefly decurrent, apex acute with a

tiny tubule; midrib prominently raised abaxially, slightly raised adaxially;

primary venation densely pinnate, slightly raised abaxially, somewhat

impressed adaxially; interprimaries sub-parallel to primaries, slightly raised

92 Gard. Bull. Singapore 52 (2000)

Figure 1. Rhaphidophora hayi P.C. Boyce & Bogner

A. adult shoot with flowering branch x '/,; B. leaf lamina x +; C. venation detail x 3; D. pre-

adult climbing shoot x +; E. disarticulating side shoot x '/.; F inflorescence x 1; G. spadix detail

at female receptivity x 10; H. spadix detail at post anthesis x 10. A-C from Webb & Tracey

7066; D — H from Sands, Pattison & Wood 2384.

Neotenic Rhaphidophora in New Guinea and Australia o3.-:

on both leaf surfaces; secondary venation reticulate, slightly raised abaxially,

+ flush adaxially; inflorescence solitary, subtended by a membranaceous

prophyll and one or more cataphylls, these soon falling; peduncle slightly

laterally compressed, 2—3 x 0.6—1 cm; spathe canoe-shaped, stoutly beaked,

5.5-8 x 2-4 cm, stiffly fleshy, yellow, gaping wide at female receptivity and

then slowly falling to leave a large scar at the base of the spadix; spadix

stoutly cigar-shaped, shortly stipitate; stipe 4-6 x 3-3.5 mm; spadix inserted

+ level on stipe, 3.5-6 x 1-1.2 cm, yellow; stylar region weakly developed,

mostly irregularly rhombohexagonal, 1—-1.3 x 1—1.1 mm, truncate; stigma

prominently raised, elongated, longitudinally orientated, c. 0.3-0.5 x 0.2-

0.4 mm; anthers not exserted at anthesis; infructescence not seen.

Distribution: Irian Jaya, Papua New Guinea [including New Britain, New

Ireland, Bougainville and Muyua (Woodlark) Island], and Australia

(Eastern tropical Queensland).

Habitat: Primary, secondary and monsoonal rain forest on coralline

limestone and basalt, 20 — 600 m altitude.

Note: While resembling Rhaphidophora pachyphylla K. Krause, R. hayi is

immediately distinguishable by the flowering shoots with broader, truncate-

based leaf laminas, the raised, larger, elongated, longitudinally orientated

stigmatic region, the occasional foraging shoot, and by the presence of

free, disarticulating side shoots functioning as vegetative propagation units.

This last character is unique in the Monsteroideae.

Other specimens seen: INDONESIA: Irian Jaya, 2 km N of Manokwari,

Nicolson 1577 (B, K, L, US). PAPUA NEW GUINEA: Woodlark (Muyua)

Island, Kulumadau, Brass 28831 (GH, L); Milne Bay, Esa’ala subdistrict,

Normanby Island, Sewa Bay, 21 Oct 1971, Lelean & Streimann LAE 52541

(L, US); Cnetral Prov., Sogeri Plateau, 5—7 miles beyond Kokoda Trail

Monument, 30 miles east of Port Moresby, Nicolson 1431 (L, US);

Bougainville, Arawa, McKillup’s Plantation, 6 km west of Kieta, Nicolson

1512 (B, K, US), New Britain, Kandrian, along road from airport, Nicolson

1540 (L, US); New Ireland, Namatanai, Hans Meyer Range, above Mandih

river, near Mandih Lake, c. 6 km WNW of Taron, 30 Oct 1975, Sands,

Pattison & Wood 2384 (K & K (living collection accession no. 1975-5026)

& K (spirit no. 63938)). AUSTRALIA: Queensland, Cape Tribulation,

Rijkers 1484 (BRI). There are many collections from Australia (mostly at

BRI and ORS) but all, except that cited here, are sterile and are not

included here.

94 Gard. Bull. Singapore 52 (2000)

2. Rhaphidophora okapensis P.C. Boyce & Bogner, sp. nov.

A Rhaphidophora okapensis differt ab omnibus speciebus ceteris Novae

Guineae ramis rigide erectis, laminis foliorum parvis valde coriaceis ovato-

cordatis longe acuminatis, inflorescentiis amplis purpureis differt — TYPUS:

Papua New Guinea, Eastern Highlands, 5 miles NE of Okapa, 24 Sept

1964, Hartley TGH 13098 (CANB, holo; GH, L, iso).

Figure 2.

Moderate, lightly robust, semi-leptocaul, homeophyllous neotenic liane to

unknown ultimate height; seedling stage and pre-adult plants not observed;

adult shoot architecture comprised of clinging, physiognomically

unbranched, densely leafy, sterile stems, and adherent, leafy fertile stems:

stems terete in cross section, smooth, without papery adherent prophyll,

cataphyll and petiolar sheath remains, internodes to 5 x 0.75 cm, separated

by slightly swollen nodes with rather + sloping scars, older stems sub-

woody; flagellate foraging stems not observed; clasping roots arising sparsely

from the internodes, smooth to very slightly pubescent; feeding roots not

observed; /eaves distichous, those on adherent shoots weakly shingling to

slightly scattered, those on free shoots pendent to slightly spreading;

cataphylls and prophylls membranous, soon drying and falling; petiole

shallowly and broadly grooved, 1-4 x 0.1—-0.2 cm, smooth, apical and basal

genicula quite prominent; petiolar sheath prominent, membranous, ligulate,

ligule extending c. 1.5 cm beyond apical geniculum, soon drying and

adhering to stem, later disintegrating and falling; /amina ovate, mid-green

above, paler below, very stiffly coriaceous, base cordate to rounded and

slightly notched, apex long acuminate with a pronounced tubule; midrib

proximally raised abaxially, slightly impressed adaxially; primary venation

pinnate, slightly raised on both leaf surfaces; interprimaries sub-parallel to

and barely distinguishable from primaries, slightly raised on both leaf

surfaces; secondary venation reticulate, slightly raised abaxially and

adaxially; inflorescence solitary on short leafy shoots, subtended by a fully

developed or reduced foliage leaf, and a membranous, soon-degrading,

long-ligulate prophyll; peduncle terete, 4-6 x 0.15—0.2 cm; spathe broadly

canoe-shaped, rounded, minutely apiculate, 3—3.5 x 1.2—1.5 cm, stiffly fleshy,

yellow, turning purple, falling to leave a large scar at the base of the

spadix; spadix stout-cigar-shaped, stipitate, inserted level on stipe, 1.9-2.2

x 0.75-1 cm; stipe 4-4.5 x 1-1.2 mm; stylar region conical, regularly

rhombohexagonal in plan view, 0.8-1 x c. 1 mm; stigma slightly raised,

punctiform, c. 0.4 mm diam.; anthers exserted at anthesis; infructescence

not seen.

Neotenic Rhaphidophora in New Guinea and Australia 95 :

Figure 2. Rhaphidophora okapensis P.C. Boyce & Bogner

A. adult shoot with flowering branch x */,; B. leaf lamina x 1; C. venation detail x 4; D.

inflorescence x 1+; E. spadix detail at anthesis x 10; F. pistil, side view x 10. All from from

Hartley 13098.

96 Gard. Bull. Singapore 52 (2000)

Distribution: Papua New Guinea, known only from the type.

Habitat: Disturbed mixed forest on slope. c. 1500 m altitude.

Note: A very distinctive species notable for the stiffly erect, densely leafy

stems, the thickly coriaceous, ovate-cordate leaves with a long acuminate

tip, the relatively large inflorescences that turn purple at maturity, the

long-stipitate spadix, and the conical stigmatic region. Based on overall

appearance, it is not immediately obvious to what R. okapensis is related.

It is included here on account of its neotenic habit, a character that might

lead to confusion with the preceding species.

3. Rhaphidophora pachyphylla K. Krause

Rhaphidophora pachyphylla K. Krause, Bot. Jahrb. Syst. 49 (1912): 92 —

Type: Papua New Guinea, Madang Province, near Wabbe, 29 Aug 1907,

Schlechter 16463 (B, holo; P, iso)

Figure 3.

Moderate to rather large, somewhat robust, semi-leptocaul, homeophyllous

neotenic liane to 5 m; seedling stage a non-skototropic shingling juvenile

shoot; pre-adult plants forming small terrestrial colonies; adult shoot

architecture comprised of clinging, physiognomically unbranched, mostly

densely leafy, sterile stems and, free, leafy fertile stems; stems rectangular-

terete in cross section, widest side prominently convex, smooth, mid-green,

mostly without prophyll, cataphyll and petiolar sheath fibre although

flowering shoots occasionally and briefly with parchment-like remains,

internodes to 3 x 1 cm, separated by rather prominent + straight scars,

older stems sub-woody; flagellate foraging stems absent; clasping roots arising

sparsely from the internodes, pubescent; feeding roots not observed; leaves

distichous, those on adherent shoots shingling, those on free shoots slightly

spreading, all densely arranged; cataphylls and prophylls membranous, soon

drying and falling; petiole deeply grooved, 1.5—-4 x 0.15-0.2 cm, smooth,

apical and basal genicula quite prominent; petiolar sheath prominent,

membranous, ligulate, slightly unequal on one side, of short-duration,

degrading to very weak fibres and soon falling; /Jamina narrowly ovate-

elliptic, stiffly coriaceous, base acute to cuneate, apex acute with a short

tubule; midrib prominently raised abaxially, slightly impressed adaxially;

primary venation pinnate, slightly raised abaxially, somewhat impressed

adaxially; interprimaries sub-parallel to primaries, slightly raised on both

leaf surfaces; secondary venation weakly reticulate, slightly raised abaxially,

+ flush adaxially; inflorescence solitary on short to somewhat elongated

Neotenic Rhaphidophora in New Guinea and Australia 97

are,

One

Orn)

0-0"

a at

5.010

“ fi

fc)

@*9

Figure 3. Rhaphidophora pachyphylla P.C. Boyce & Bogner

A. section of adult shoot with flowering branch x +; B. section of adult climbing shoot x +;

leaf lamina x +; C. juvenile climbing shoot x '/,; D. pre-adult climbing shoot x +; E. leaf lamina

x +; F venation detail x 4; G. inflorescence, spathe fallen x 1+; H. spadix detail, post anthesis x

10. A, E-F, G-H from Schlechter 16436; B from Kalkman 3389; C—D from Nicolson 1416.

98 Gard. Bull. Singapore 52 (2000)

leafy shoots, subtended by a fully developed foliage leaf, and soon-degrading

membranous prophyll; peduncle terete, 4-7 x 0.2—0.25 cm; spathe broadly

canoe-shaped, obtuse to slightly pointed, 5—6.5 x 1.5—2.2 cm, stiffly fleshy,

yellow, gaping at female receptivity and then falling to leave a large scar at

the base of the spadix; spadix narrowly cigar-shaped, long stipitate; stipe 6—

10 x 2-2.4 mm; spadix inserted obliquely on stipe, 4-5.5 x 1.2-1.6 cm,

yellow; stylar region weakly developed, mostly irregularly rhombohexagonal,

1-1.2 x 1—1.4 mm, truncate; stigma slightly raised, punctiform, c. 0.2—0.3 x

0.3-0.35 mm; anthers slightly exserted at anthesis; infructescence not seen.

Distribution: Irian Jaya, Papua New Guinea.

Habitat: Monsoonal lowland rainforest at 10-30 m altitude (2380 m in

montane Nothofagus forest Hoogland & Schodde 6943).

Notes: 1. Rhapidophora pachyphylla differs from R. hayi in leaf shape and

form of the stigma. In addition, R. pachyphylla lacks the disarticulating

shoots unique to R. hayi and never has foraging shoots.

2. As noted above, it is still not fully clear whether R. pachyphylla as here

defined is a single taxon. The collection Hoogland & Schodde 6943 cited

below, and for which habitat and elevation data appear in parentheses

above, while vegetatively typical of R. pachyphylla, has a markedly rounded

spathe apex, a shorter stipe (c. 3 mm long) and also represents an enormous

altitudinal increase on the other specimens seen. More specimens of ‘R.

pachyphylla’ from higher altitude are required to resolve this plant’s status.

Other specimens seen: INDONESIA: Tamimonding, Ka/lkman s.n. (L);

Merauke, Bis, Agats, Widjaja 6344 (BO, K, L). PAPUA NEW GUINEA:

Western Highlands, Wabag subdistrict, near Poio village, west slopes of

lower Yaki valley, 6 July 1960, Hoogland & Schodde 6943 (BM, GH, L):

Brown River F.R., 20—25 miles northwest of Port Moresby, Nicolson 1416

(K,:L, US), |

Acknowledgements

Thanks are due to Linda Gurr for skilfully executing the illustrations that

accompany this article. We also wish to thank Dr H. RoeBler, Munich, for

providing the Latin translation of the diagnoses.

Neotenic Rhaphidophora in New Guinea and Australia 99 ;

References

Boyce, P.C. 1999. The genus Rhaphidophora Hassk. (Araceae-

Monsteroideae-Monstereae) in Peninsular Malaysia and Singapore.

Gardens’ Bulletin Singapore. 51: 183—255.

Boyce, P.C. 2000. The genus Rhapidophora Hassk. (Araceae-Monsteroideae-

Monstereae) in the Philippines. Gardens’ Bulletin Singapore. 52: (in press).

Boyce, P.C. (in prep). The genus Rhaphidophora Hassk. (Araceae-

Monsteroideae-Monstereae) in Sumatera, Java, Nusa Tenggara, Sulawesi,

and Maluku. Gardens’ Bulletin Singapore.

Hay, A. 1981. Araceae. In R.J. Johns & A. Hay (eds), A Students’ Guide to

the Monocotyledons of Papua New Guinea. Part 1: 31-81. Office of

Forests, Port Moresby, Papua New Guinea.

Hay, A. 1990. Aroids of Papua New Guinea. Christensen Research Institute,

Madang.

Hay, A. 1993. Rhaphidophora petrieana - a new aroid liane from tropical

Queensland; with a synopsis of the Australian Araceae-Monstereae.

Telopea 52: 293-300.

Hay, A. (in press). Araceae. In Flora of Australia. Vol. 39. ABRS/CSIRO,

Melbourne.

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. Blumea, suppl. 8.

210 pp.

Jones, D.L. & Gray, B. 1988. Climbing Plants in Australia. Reed, Sydney.

Nicolson, D.H. 1978. Araceae. In A.C. Smith (ed.), A precursor to a new

Flora of Fiji. Allertonia 1: 331-414.

Nicolson, D.H. 1979. Araceae. In A.C. Smith (ed.), Flora Vitiensis Nova. 1:

438-460.

Stearn, W.T. 1992. Botanical Latin. 4" edition. David & Charles, Newton

~ Abbot, England.

100 Gard. Bull. Singapore 52 (2000)

Index of Exsiccatae

hayi=1 okapensis = 2 pachyphylla = 3

Brass 28831: 1; Hartley 13098: 2; Hoogland & Schodde 6943: 3; Kalkman

s.n.: 3; Lelean & Streimann LAE 52541: 1; Nicolson 1416: 3; 1431, 1577: 1.

Rijkers 14841: 1; Sands, Pattison & Wood 2384: 1; Schlechter 16436: 3;

Webb & Tracey 7066: 1; Widjaja 6344: 3.

Gardens’ Bulletin Singapore 52 (2000) 101-183.

The Genus Rhaphidophora Hassk.

(Araceae-Monsteroideae-Monstereae) in

the Southern and Western Indonesian Archipelago

P.C. Boyce

Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, U.K.

Abstract

An alpha-taxonomic account of Rhaphidophora in Sumatera, Java, Nusa Tenggara, Sulawesi

and Maluku is presented as a precursor to the forthcoming Flora Malesiana Araceae

treatment. Twenty four species are recognized, of which five (R. araea P.C. Boyce, R.

balgooyi P.C. Boyce, R. floresensis P.C. Boyce, R. sabit P.C. Boyce and R. ustulata P.C.

Boyce) are newly described. One new synonomy (R. scaberula Alderw. into R. puberula

Engl.) is made. In addition, R. moluccensis Eng]. & K. Krause is treated as doubtful. Eight

informal morpho-taxonomic units (‘Groups’) are proposed and compared. A dichotomous

key to species is provided and 21 species are illustrated.

Contents

ES yh Be aE 5 he) EC eR eee an Pe 101

UI RTEN RRA RPE Dc Bic, SURAA ARM 2 So ies ie SR Rg ec cn )4 NousiinssS pial banat eepnteannsipnagies 102

nee MUD UINEE OES OA PAI WMUISUIS <5.) og... Ah fos de desde gativcnnaseipcataayarepieceantospjaryetbesniarents 102

ameremesRSRENRN SP URIEN 1 PEORCSTNUONANE U5. .2/00 nous doe ets capes bo ce coats ee cee sctcntacces 104

NTE Gig) 4 TRAE UR Oe ea Se UR 105

Key to adult flowering Rhapidophora species 2--7...............1cccsssccssosceeenes 108

Lo Ce MEN Rogge eee Nee oe eS SE Ong Ts Er eS fo 0 oe 112

ce a Las ah RRM a i ae 178

a et =, Se ae eR OL Op En ET OREC Et Ay ee 179

= as Ba BS DELLS Mi ieeteiae Men pte Boag daa SE OPE Cota a ee eee en ee 179

a es lees |e canes ee dei es ee ea See ee 181

a i Lane Cia os Sin eeen einen Siena s. See See ee TREE” Reet eee Oe ee ee 183

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.

102 Gard. Bull. Singapore 52 (2000)

Rhaphidophora is one of the largest aroid genera represented 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 second in a series of papers intended to present a complete

alpha-taxonomy of the genus Rhaphidophora in Asia. An account for

Peninsular Malaysia and Singapore has been published recently (Boyce,

1999) and accounts for each of the Himalaya, Thailand and Indochina, the

Philippine islands (Boyce, 2000), Borneo, and Papuasia are being prepared

and will be published separately. All morphological terms employed follow

Stearn (1992).

Synonomy cited is for the species, not for the review area. The reason

for this is that synonymic names based on types outside the review area

are frequently applied to species in local herbaria.

History

The last complete revision of Rhaphidophora was that of Engler & Krause

(1908). For the Malesian region, to date no critical account of the genus

has been prepared for Borneo (but see Alderwerelt, 1920, 1922; Miquel,

1856a & b; Ridley, 1905; Merrill, 1921), the Philippine Islands (but see

Merrill, 1923) and New Guinea (but see Engler & Krause, 1910, 1912;

Krause & Alderwerelt, 1924; Hay, 1981, 1990). A summary of the taxonomic

and nomenclatural history of Rhaphidophora was presented in Boyce

(1999).

Infrageneric Relationships

With an alpha-taxonomy of Rhaphidophora developing, potentially

informative higher morpho-taxonomic units are beginning to emerge.

Although it is too soon to begin rigorously testing phylogenetic relationships

within and between these groups, it is clear that they can be usefully given

informal names. The groups are named using the earliest accepted species

epithet belonging to the group. A summary of these groups for the review

region and elsewhere follows.

The Hongkongensis Group is defined by shingling, non-skototropic

seedling and shingling juvenile shoots, preadult and adult plants with

climbing stems square to rectangular in cross-section, simple, often

coriaceous leaf laminas, a petiolar sheath extending beyond the leaf base

by short to rather long ligules and the sheath soon falling to leave a

horseshoe-shaped scar around the top of the apical geniculum. All species

Rhaphidophora in southern and western Archipelago 103

flower on free lateral shoots that are either angular or more-or-less terete

in cross-section. Species in the Hongkongensis Group are often lofty

climbers and are frequently very inadequately collected, making working

from herbarium specimens tedious and unrewarding. The group contains

numerous species, many from Indochina and Thailand. Species in the review

area are R. conocephala Alderw., R. maingayi Hook.f., R. montana (Blume)

Schott, R. sylvestris (Blume) Engl. and R. talamauana Alderw.. Perhaps

the most easily recognizable, although paradoxically the most taxonomically

intractable of these informal groups, much remains to be done before the

Hongkongensis Group can be regarded as well understood.

The Hookeri Group has seedling and juveniles with spreading leaves,

the apical geniculum and abaxial surface of leaf lamina pubescent, the

primary lateral veins usually much more conspicuous than the

interprimaries, and long petioles (at least three quarters as long as the

lamina) with the sheath extending to the apical geniculum. The leaf lamina

can either be entire or perforated, occasionally (R. puberula Engl.) in the

same plant. All stems are terete in cross-section and are commonly glabrous,

rarely tomentose (R. hookeri Schott — E. Himalaya to S China). Flowering

is either on clinging or free shoots. Species in the review area are R.

foraminifera (Engl.) Engl. and R. puberula.

The Korthalsii Group comprises four species with non-skototropic

shingling seedlings, shingling juvenile growth and either heteroblastic leaf

development leading to spreading, variously pinnately divided and

perforated leaves, or neotenic development resulting in shingling fertile

adult plants. Petioles may be long (equalling or exceeding the lamina) or

much shorter, the petiolar sheath reaches the apical geniculum, overtopping

it by a short auriculate ligule on one or both sides. Sheaths are either long

persistent or soon degrade into persistent to swiftly falling fibres. All species

are notable for producing densely ramentose-scaly feeding roots and are

either high-climbing, e.g., R. korthalsii Schott, R. crassicaulis Engl. & K.

Krause (Indochina) and R. latevaginata M. Hotta (Borneo), or creeping

rheophytes, R. beccarii (Engl.) Engl.. Species in the Korthalsii Group

covered in this treatment are R. korthalsii and R. beccarii.

The Neoguineensis Group as presently defined is mostly Wallacean,

with only one species, R. cylindrosperma Engl. & K. Krause (Borneo) in

Sundaland. The plants have long, terete stems, rather small, often slightly

stiff leaf laminae, petioles with the sheath extending to the apical geniculum

_and overtopping it by two small ligules. Inflorescences are carried on the

tips of, often long, free shoots. Specimens are notable for drying black. It is

not clear what relationship, if any, exists between this and the next two

monospecific groups.

The Minor Group contains only R. minor Hook.f., a diffuse climber

104 Gard. Bull. Singapore 52 (2000)

with flexuous, smooth, + terete clinging stems, and flowering on the tips of

often very long free shoots, leaf laminas are chartaceous with primary

lateral and interprimary veins not differentiated from one another (leaf

lamina thus appearing densely veined), and short petioles in which the

petiolar sheath extends beyond the apical geniculum by two small ligules.

R. minor dries a characteristic straw colour. This and the next group are

notable for growing in swampy places, an unusual habitat for monsteroids.

The Lobbu Group, containing just R. /obbii Schott, is similar to the

Minor Group but differs in having puberulent-scabrid to asperous stems,

very softly coriaceous leaves not drying straw-coloured and with the primary

lateral veins conspicuously thicker than the interprimaries, flowering shoots

much shorter and the spathe exterior minutely puberulent. The seedling is

leafy at germination becoming skototropic by alternating series of congested

leafy and elongated leafless shoots. (The seedling of R. minor not known.)

The Hollrungii Group with one species, R. megasperma Engl., in

Borneo and the rest in Wallacea is probably not a natural group but it is

useful to recognise it at present until the majority of its constituent species

are revised. The group is defined by inflorescences occurring in clusters of

3 — 10, with each inflorescence subtended by and separated from the next

by a conspicuous chartaceous cataphyll. Inflorescences arise either at the

tips of a clinging primary or lateral axis; if the former then vegetative

growth is suspended for the flowering event and then reiterated, the

vegetative portions of the axis thus occurring as discrete leafy modules

separated by leafless portions bearing an inflorescence/infructescence (later

naked). The spathe is persistent and marcescent.

Plants of the Decursiva Group have foliage leaves interspersed with

up to seven cataphylls, giving plants a rather sparse appearance. Seedling

and juvenile plants are non-shingling, adult plants have pinnately divided,

occasionally perforated leaves. Stems are terete and glabrous, although in

R. glauca (Wall.) Schott (Himalaya, Thailand, Indochina) the most robust

stems are slightly muricate. Flowering occurs on the tips of clinging primary

axes, more rarely on clinging lateral shoots. As with the Hongkongensis

Group, many of the species occur outside the Malesian region. None is yet

recorded for the review region, although R. nicolsonii P.C. Boyce, is endemic

in Peninsular Malaysia.

Geography and Endemism

As noted in Boyce (1999), Malesian Rhaphidophora species divide into

two distributional groups. One comprises taxa with limited distributions,

sometimes narrowly endemic, more usually restricted to one or more

Rhaphidophora in southern and western Archipelago 105

geographically adjacent landmasses, and displaying limited morphological

variation. The other group comprises species with extensive distributions

and, usually, a wide range of variation.

The southern and western Indonesian archipelago is notable for the

high percentage of locally endemic Rhaphidophora species, with 13 of the

25 species recorded from only one island or island group. This compared

with Peninsular Malaysia with only two locally endemic species from a

total of 15.

While a paucity of botanical collections is undoubtedly a contributing

factor to this apparent high level of local endemism, it is quite possible

that the low number of botanical collections for many species is itself a

result of the rarity and local distribution of these species. Indeed, the

greater number of collections of widespread and common species such as

R. korthalsii support this hypothesis.

There are no data for the conservation status of any of these locally

endemic species and much remains to be done by way of field studies to

ascertain whether any or all of these species are threatened.

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, 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; Engl. & Prantl, Nat.

Pflanzenfam. Nachtr. | (1897) 58; Engl. & K. Krause in Engl., Pflanzenr.

37 (IV.23B) (1908) 17—53; Engl. & Prantl, Nat. Pflanzenfam. Nachtr. 3

(1908) 29; Koorders, Exkursfl. Java, 1 (1911) 253—255; Backer, Beknopte

Fl. Java, 17 (1957) 13—15; Backer & Bakh. f., Fl. Java, 3 (1968) 106—107;

Schott, Icones aroideae et reliquiae (IDC Microfilm) (1983) fiche nos.

28—31, 121; 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.) Miq., 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.]

106 Gard. Bull. Singapore 52 (2000)

| 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 lianes, very seldom

clustering and rheophytic (e.g., R. beccarii) and then always with a creeping

juvenile stage: cut surfaces producing clear, odourless, sticky juice either

drying + invisibly or coagulating into yellowish, translucent jelly and

eventually hardening to a brittle amber-like mass: seedling stage mostly not

observed but where known either leafy at germination and skototropic

(see Strong & Ray, 1975) by an alternating series of congested leafy and

elongated leafless shoots (e.g.. R. angustata Schott) or germinating giving

rise to a non-skototropic, shingling juvenile shoot (e.g., R. korthalsii); pre-

adult planis 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: 1. 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 ii. all stems physiognomically monopodial, 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 the review area), older stems subwoody or

somewhat corky or with distinctive matt to sublustrous pale brown papery

epidermis. with or without variously textured prophyll, cataphyll and

petiolar sheath fibre either at the tips or along the newer sections, rarely

with cataphylls and prophylls deliquescing to black mucilage which on

drying leaves fragmentary parchment-like remains on petioles, developing

laminae. 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; leaves distichous or weakly

spiralled, evenly distributed or scattered or clustered distally: cataphylls

and prophylls subcoriaceous to membranous, either soon drying and falling

or degrading or deliquescing to variously textured sheaths and fibres, these

Rhaphidophora in southern and western Archipelago 107

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 or fully or overtopping the geniculum,

occasionally one side greatly expanded and auriculate, especially in juvenile

plants, at first membranous 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; /amina submembranous to stiffly chartaceous or

coriaceous, 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, sometime flush, rarely slightly raised adaxially; primary

venation + pinnate; interprimaries mostly present, subparallel to primaries

and sometimes indistinguishable from them (e.g., R. monticola K. Krause -

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 swiftly disintegrating cataphyll,

subsequent inflorescences usually each subtended by a prophyll and

cataphyll, more rarely by a prophyll and partially to an 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. novoguineensis Engl. - New

Guinea), stiff to rather soft or stoutly coriaceous, dirty-white, greenish,

cream or yellow; spadix subglobose to clavate-cylindrical, cylindrical or

fusiform, sessile or stipitate, often obliquely inserted on peduncle, tapering

108 Gard. Bull. Singapore 52 (2000)

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 + subaxile, 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, orientation circumferential or longitudinal; stamens 4—6;

filaments strap-shaped; anthers usually prominently exserted from between

ovaries 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) (Mayo et al. 1997), exine foveolate, subreticulate, rugulate, fossulate,

scabrate, retiscabrate, verrucate, or psilate; chromosomes 2n = 60, 120 (42,

54, 56) (Mayo et al, 1997).

Distribution: About 100 species from tropical Africa, South and South

East Asia, Australia and the Pacific with extensions into the subtropical

Himalaya, southern China and the southernmost islands of Japan.

Habitat:, 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 as lianescent bole-climbers,

lithophytes, rarely rheophytes.

Etymology: Greek rhaphis, rhaphidos (needle) and phero (I bear); refers

to the macroscopic (to Icm long), needle-like, unicellular trichosclereids

present in tissues.

Key to Adult Flowering Rhaphidophora in the Southern

and Western Indonesian Archipelago

Note. Distributions given in the key refer to the review area only.

Rhaphidophora in southern and western Archipelago 109

la.

1b.

2a.

2b.

3a.

3b.

Aa,

Ab.

an:

5b.

6a.

6b.

7a.

7b.

8a.

Sb.

9a.

Leaf lamina variously pinnately divided and/or perforated ............ 2

ES i a cea ee ete eos, ae 5

Leaves variously pinnately divided and/or perforated. Plants

a rea Nt PEN PTI SUIS oop ientenstisacwnnnted UR clditwaslebatsiulblibdasessees 3

Leaves with scattered, small perforations. Plants flowering on free

lateral stems. Sumatera, Nusa Tenggara.................. 17. R. puberula

er te ean WEEE CS AOI, ANA eee 4

Flowering plants rheophytic, not climbing. Sumatera...

Leaf lamina of mature plants slightly to extensively perforated,

perforations round to rhombic, extending c. quarter of lamina width

on each side of the midrib, abaxial surface of lamina with pubescent

veins; active shoot tips with black mucilage. Sumatera .........

Ne SE 2) ice Ls te I EAA. 7. R. foraminifera

Leaf lamina of mature plants pinnatisect, the pinnae often perforated

basally and appearing stilted, lamina always glabrous; active shoot

tips with sparse to copious netted fibre. Sumatera, Java, Sulawesi

ORRIN Ss tk. Ne 9. R. korthalsii

Abaxial surface of lamina and apical geniculum pubescent. Sumatera,

Re WIS, Bg ROR OL 17. R. puberula

Panna and apicalpemiculiim glabrous). 01.0400. e ies 6

Flowering shoots consisting of scattered fans carried on short stout

shoots and held at about 90° to the + leafless main stem. Sumatera

PE eR ck pe Ls DAU ASE CET: BRM AMEE 1. R. angustata

Beet GOS MOL ASAD OVE 2o. Socici sacs neoresviisnaeedeocthosabtssaneacscaeadsuvdeess 7

leprae 0) Ce a ee 6 fe en Ee be eee ae Se 9

J SEI @ cg Ma ag ofc 8) 0) (1) Gene ey Pe Een che ee oe 8

Peduncle much shorter than petiole, spathe 6.5—7 cm, spadix 4.5—7

cm, sessile. Plant habitually rheophytic. Sumatera ...... 4. R. beccarii

Peduncle subequalling or exceeding petioles, spathe 4—5 cm, spadix

2.4—3.2 cm, stipitate. Plant terrestrial, perhaps occasionally

PHeephylicNSuimalera (emdemic) sy Alige.. iki lel iae sees 2. R. araea

Flowering plant with leaves appressed to substrate and shingling on

adherent shoots, slightly scattered on flowering shoots. Maluku

110

9b.

10a.

10b.

lla.

11b.

12a.

12b.

13a.

13b.

14a.

14b.

15a.

15b.

l6a.

17a.

17b.

18a.

Gard. Bull. Singapore 52 (2000)

(Ternate, Halmahera «¢ndeumt)aec 3, 16. R. parvifolia

Flowering plant not.as. aDOVE scpccreseracssesiinnc a Oe 10

Stems scabrid to asperous. Spathe exterior minutely puberulent.

Sumiatera., SUlaWeSh iscsi. dcccacseee ee Sele ote ee 10. R. lobbii

Adherent stems square or rectangular in cross-section; tips of active

stems with netted prophyll, cataphyll and petiolar sheath. Sumatera

snsnonnishgnmnatinhiancilel LRiph coef soe Misa Rp vad Oe Meg Rae 11. R. maingayi

Adicieas stems variously shaped in cross-section; tips of active stems

without fibrous. material .1ic.:.. 010054440 eee ee 12

Spadix shipitate ;...iiussepncstalsno.-ctidstiekal dl Ge ee 13

Spadin SCSBME 6: 45 -chnisiwnds hatin wae see ee 19

Leaf lamina thickly coriaceous. Sumatera ............ 5. R. conocephala

Leaf lamina variously textured but not thickly coriaceous. .......... 14

Lamina at least 7 times longer than wide, narrowly falcate-lanceolate

to narrowly falcate-oblanceolate. Sulawesi (endemic) .... 18. R. sabit

Lamina no more than 5 times longer than wide. .............:eeeeee 15

Spadix at anthesis.8-—15 cm JOng ccs bots RR Se 16

Spadix at anthesis no more than 7.5 cm ION .........e ee eeeeeeeeeeeeeeeees 18

Cataphyll, prophyll and petiolar sheath fibres present. Primary lateral

veins significantly more prominent than interprimaries. Stigma

elongate to rounded, raised. Maluku (Ternate, Ceram — endemic).

sons bachiandiebsamuietsias baidipmaiiaiolecana nde lgeenme Nt Sate, DRS 22. R. ternatensis

Cataphyll, prophyll and petiolar sheath fibres absent. Primary lateral

veins thicker and clearly differentiated from interprimaries. Stigmas

VALIOUS .o.-icassninansorvissonsiiondeieis > Meee Os Ae eee a eee AT

Spadix at anthesis 11—15 cm long; stigma round, flat. Maluku

(Téimate, Bure ~ endemie}iane. gies eee 3. R. balgooyi

Spadix at anthesis 8—10 cm long; stigma punctiform, raised. Sulawesi

(endéniic) 2.02223. 2408. Lae eee 19. R. sarasinorum

Leaf lamina on flowering shoots elliptic-lanceolate to oblong-elliptic,

stiffly coriaceous, base cuneate, apex acuminate to attenuate with a

prominent tubule. Sulawesi (endemic) ...................4- 8. R. koordersii

Rhaphidophora in southern and western Archipelago H11

18b.

19a.

19b.

20a.

Zia:

21.

oes

22d,

23a,

23b.

24a.

24b.

25a.

2ab;

Leaf lamina on flowering shoots lanceolate to oblong lanceolate, +

falcate, thinlycoriaceous, base acute, apex acute to long-acuminate

with a slight tubule. Maluku(Ternate, Tidore, Ceram - endemic) ...

es os citsi2s tm bk fT ian ta to eTComsinings tt 15. R. oligosperma

Sead at abiesis 3.5258!5 CrmbOUe eens. Me end clade eadee didaeens 20

Spadix at anthesis 9—20 cm long. Sumatera, Java, Nusa Tenggara,

Pe aN A WSU ede enchaceoceuadcwesvsevceacervustncesecnvbeeeen: 14. R. montana

ee CLAN ATCC Y MIMIC He TR opengeh . Saas when ea sila de bine cbadvianidiies 22

Bpacix cylinder! to tapering cylimdncal %....5..6.iBiicn. ns ccdee 24

Leaf lamina notably coriaceous. Stigmas c. Imm diam., the middle

excavated, the edges prominently raised. Sumatera (endemic)........

SRS EE ee Ree | oe meee) ee Rt are 12. R. megastigma

Leaf lamina not notably coriaceous, mostly rather thin. Stigmas not

as above or if as large then not excavated and lacking a prominently

eo ee ae ae AL: Rm Sh Oe Ree eRe eer RO Ere 22

Spadix not more than 3 cm long at anthesis. Sulawesi (endemic) ...

ee cnn ecckh? header nM. dist arti Gotit es wens. 23. R. teysmanniana

Spadix exceeding 3 cm, often up to 6 cm long, at anthesis ........... 23

Flowering shoots stems + rectangular in cross-section. Spathe beak

short. Sumatera,Java, Nusa Tenggara... eee 20. R. sylvestris

Flowering shoots terete in cross-section. Spathe beak long. Sumatera

FET Ve 170 BG i) ee Cae ee ee 21. R. talamauana

Leaf lamina 2.5—16 x 1.2—3 cm, drying uniformly pale straw-

coloured. Spadix slender cylindrical, 2.5—7 cm long. Sumatera,

EE OT PEC OMBEE Ce tt otetaee gan cs o ar Sat CE ane eo en 13. R. minor

Leaf lamina 21—34 x 4—7 cm, drying darker above, paler below.

Spadixitapering cylindrical; 6.5—8:5 cm lone... .2.s.scs.ceissccaelvesaes oS

Petiole drying pale with apical geniculum much darker, leaf lamina

drying strongly discolorous, pale yellow-brown abaxially, mid-brown

adaxially. Spadix strongly tapering-cylindrical; stigma discoid, flat.

Sumatera (Pulau Enggano - endemic).................:ee 24. R. ustulata

Petiole drying + uniformly mid-brown, leaf lamina drying barely |

discolorous. Spadix slightly tapering-cylindrical; stigma punctiform,

raised. Nusa Tenggara (Pulau Flores - endemic).....6. R. floresensis

112 Gard. Bull. Singapore 52 (2000)

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).

Scindapsus pteropodus Teysm. & Binn., Natuurk. Tijdschr. Ned.-Indié. 27

(1864) 23; Engl. in A. & C. DC., Monogr. Phan. 2 (1879) 254 —

Rhaphidophora pteropoda (Teysm. & 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 — Type: Indonesia, Sumatera,

Loeboe-Aloeng, Teysmann s.n. (BO, holo; K, iso).

Rhaphidophora laetevirens Ridl., J. Bot. 40 (1902) 37 — Type: Malaysia,

Penang, Bk Penara, 1896, Ridley s.n. (SING, lecto; selected by Boyce,

1999):

Figures 1 & 2

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 subwoody; flagellate foraging stems exceedingly long

with internodes to 50 cm or more and nodes with semipersistent cataphylls;

clasping roots dense, arising from the nodes and internodes of clinging

stems, prominently pubescent; feeding roots rare, adherent, pubescent; leaves

weakly spiralled on adherent and flagelliform shoots, those on free shoots

distichous into dense, few to many-leaved fans; cataphylls and prophylls

subcoriaceous, 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;

lamina entire, falcate-lanceolate to falcate-oblong, oblique, 15—61 x 4—20

cm, subchartaceous, base acute to unequal, apex acute to slightly acuminate;

Rhaphidophora in southern and western Archipelago £3

Figure 1. Rhaphidophora angustata Schott

A. fertile shoot x '/,; B. foraging shoot x +; C. leaf lamina x ’/,; D. venation detail x 4. All from

Nicolson 927.

114 Gard. Bull. Singapore 52 (2000)

Ah \

Sate

— _—

= aS

ley. \

ale Ce.

Chee OAS

of ons

ZITO}

A fa)

PSY ERAREY

~ adie! ‘

wane

ne Or O™

Dore TO

L.GURe.

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 southern and western Archipelago 13

midrib prominently raised abaxially, slightly sunken adaxially; primary

venation pinnate, slightly raised abaxially, somewhat impressed adaxially,

the leaf appearing slightly quilted; interprimaries subparallel 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, 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, c. 0.3 mm diam., but ovaries on lowermost part of spadix with

longitudinally orientated elongated stigmas c. 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: Sumatera and Peninsular Malaysia.

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 it is unmistakably the same

species as that later described by Teysmann 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.

Other specimens seen: SUMATERA. ‘comm. D. Habury 1866’, Binnendijk

s.n (K); Aceh Prov. - Bk. Lawang, Bohorok, Langkat, Soedarsono 283 (K,

L); G. Leuser N.R. c. 35 km NW of Kutatjane, 3—5 km upstream Lau

Ketambe de Wilde & de Wilde-Duyfjes 13634 (K, L), Ketambe Research

Station, Alas river valley, c. Guhru river, de Wilde & de Wilde-Duyfjes

116 Gard. Bull. Singapore 52 (2000)

18154 (BO, K, L, MO); West Sumatera - Bt Gajabuih, about 15 km east

from Padang City, Hotta 25568 (BO, KYO).

2. Rhaphidophora araea P.C. Boyce, sp. nov.

Rhaphidophora araea differt a pluribus speciebus generis habitu terrestri

vel rheophytico; a speciebus R. beccarii et R. wentii (semper rheophyticis)

pedunculo subaequilongo vel longiore quam petioli et spadice parvo stipitato

— TYPUS: Indonesia, Sumatera, Riau Prov., S. Tulang, 13 Oct. 1939, P.

Buwalda 7032 (BO, holo; GH, K, L, SING, iso).

Figure 3

Slender, heterophyllous (?), creeping rheophyte (?) to 45 cm; seedling and

pre-adult plants unknown; adult shoot sympodial, clinging main axis creeping

or briefly ascendant; stems minutely longitudinally ridged, without petiolar

sheath fibre, internodes 0.5--1 x 0.3—0.5 cm, separated by prominent very

slightly oblique leaf scars; dormant axillary and terminal buds comparatively

very large, matt black; clasping roots very densely arising from the nodes

and internodes of clinging stems, finely pubescent; feeding roots adherent,

densely pubescent; /eaves spiro-distichous, erect or spreading and scattered

and tending to become distally clustered on adult shoots; cataphylls and

prophylls unknown, (soon drying and falling ?); petiole very narrowly

canaliculate to almost carinate, 6—20 x 0.1—2.25 cm, smooth, apical and

basal genicula not prominent; petiolar sheath barely visible, but extending

to just below the apical geniculum, soon falling; /Jamina entire, very narrowly

lanceolate to narrowly lanceolate-elliptic or narrowly oblanceolate, slightly

to prominently oblique, 9—30 x 1.5—4 cm, coriaceous, semiglossy above,

matt beneath, margins minutely revolute, base long-decurrent, apex long-

acuminate with a moderately prominent tubule; midrib raised abaxially,

sunken adaxially; primary venation pinnate, slightly raised abaxially and

adaxially; interprimaries subparallel to primaries, slightly raised abaxially

and adaxially; secondary venation reticulate, very slightly raised;

inflorescence solitary, subtended by a fully developed foliage leaf and a

prominent, narrow prophyll; peduncle subequalling or exceeding petioles,

terete, 9—14 x 0.18—0.2 cm; spathe broadly canoe-shaped, abruptly stout-

beaked, 4—5 x 0.6—1.2 cm, stiffly fleshy, yellow, falling (at anthesis ?);

spadix cylindrical, stipitate, 2.4—3.2 x c. 0.5 cm, green; stipe c. 2 x 0.75 mm;

stylar region rather well developed, mostly rhombohexagonal, c. 1—1.5 x

1—1.5 mm, very slightly conical to truncate; stigma punctiform, c. 0.3 mm

diam., raised; anthers exserted at anthesis; infructescence unknown.

Distribution: Sumatera. Endemic.

Rhaphidophora in southern and western Archipelago

I WWD oN ee

= Pa Se a ee Set, Se —_—

6. GSS SSS CSRS See , B

Orn SES Ey Sy Se POT OAS: CRIT = —_

OE : Eo" cy, ees Benes SS SOF SS z

ETS ee ROC ERO ES

SIO?

SSS

—

—————

——

—

SEEEZA-=-

—S

Figure 3. Rhaphidophora araea P.C. Boyce

llen x 2; F. spadix detail, late

A. flowering shoot x '/,; B. leaf lamina x +; C. venation detail x 4+; D. leaf lamina x +; E. inflorescence, spathe fa

anthesis x 10; G. gynoecium, three quarter view, early fruiting x 10. All from Buwalda 7032.

118 Gard. Bull. Singapore 52 (2000)

Habitat: Primary forest, secondary forest on river, on rocks. 200—400 m

altitude.

Notes: 1. Rhaphidophora araea is immediately distinguished by its terrestrial

(probably rheophytic) habit (in west Malesia otherwise known only in the

probably unrelated R. beccarii), the peduncle subequalling or exceeding

the petioles and small, stipitate spadix.

2. The specific epithet comes from the Greek word anaéiio, meaning slender,

in allusion to the habit of this species.

Other specimens seen: SUMATERA. Riau Prov. - Tigapuluh Mts, 15 km

SW of Talanglakat on Rengat - Jambi road, S. Sesirih, Burley et al. 1719

(SING, GH); S. Akar, Buwalda 6922 (BO, K).

3. Rhaphidophora balgooyi P.C. Boyce, sp. nov.

Rhaphidophora balgooyi differt a R. ternatensis cataphyllo et prophyllo et

fibris vaginae petiolaris deficientibus, nervis lateralibus primariis manifeste

minus prominentibus, spadice maiore, stigmatibus planis; differt a R.

sarasinorum spadice minore stigmatibus minoribus punctiformiter elevatis

— TYPUS: Indonesia, Maluku, Pulau Buru, Waeha River, between

Waelanga and Lake Kunturun, 15 Nov. 1984, van Balgooy 4768 (L, holo;

GH, iso).

Rhaphidophora ternatensis Alderw. f. major Alderw, Bull. Jard. Bot.

Buitenzorg III, 4 (1922): 194 — Type: Indonesia, Maluku, Pulau Ternate,

Beguin 1141 (BO, holo).

Figure 4

Large, robust, semi-pachycaul homeophyllous liane to 8 m; seedling stage

& pre-adult plants not observed; adult shoot architecture comprised of

elongated, clinging, physiognomically monopodial, leafy, non-flowering

stems and long, stout, free, sympodial, densely leafy, flowering stems; stems

smooth, internodes 1—2 cm x 0.5—1.2 cm on free shoots, separated by

well defined, oblique leaf scars; flagellate foraging stems not observed;

roots not observed; leaves spiro-distichous on free shoots; cataphylls and

prophylls membranous, drying and falling; petiole deeply grooved adaxially,

12.5—30 x 0.4—1.2 cm, apical geniculum strongly defined and drying slightly

darker than the rest of the petiole; petiolar sheath very prominent, extending

to just below the apical geniculum, upper part ending abruptly, lower part

sheathing the emerging leaf or inflorescence; /amina entire, oblong, very

Rhaphidophora in southern and western Archipelago 119

Figure 4. Rhaphidophora balgooyi P.C. Boyce

A. flowering shoot x ope B. leaf lamina x +; C. venation detail x 2; D. inflorescence, x +; E.

spadix detail, late anthesis x 6. All from Van Balgooy 4768.

120 Gard. Bull. Singapore 52 (2000)

slightly oblique, 20—38 x 8.5—11 cm, thinly coriaceous, base cuneate to

acute, apex weakly falcate, acute to briefly acuminate, with an apiculate

tubule; midrib prominently raised abaxially, very slightly channelled

adaxially; primary venation pinnate, almost imperceptibly raised abaxially

and adaxially; interprimaries parallel to primaries, barely visible; secondary

and tertiary venation very obscure; inflorescence solitary, subtended by a

fully developed foliage leaf and one or more cataphylls; peduncle slightly

compressed-cylindric, 1—12 x c. 0.5 cm; spathe elongate canoe-shaped,

stoutly long-beaked, 12—19 x 1.5—3 cm; spadix cylindrical, tapering basally,

apex acute, stipitate; stipe 6—10 x c. 2 mm; spadix inserted + obliquely on

stipe, 11—15 x 1.5—1.75 cm; stylar region mostly rhombohexagonal, 1—1.6

x 1.2—1.7 mm, truncate; stigma rounded, flat, c. 0.35—0.4 mm diam.; anthers

exserted at anthesis; infructescence not observed.

Distribution: Maluku — Buru, Ternate. Endemic.

Habitat: Riverine forest. 350—400 m altitude.

Notes: 1. Described by Alderwerelt as a robust form of R. ternatensis, R.

balgooyi differs in lacking cataphyll, prophyll and petiolar sheath fibres, in

the significantly less prominent primary lateral leaf venation, the

considerably larger spadix and flat stigmas. Confusion with R. sarasinorum

is possible. The latter may be distinguished by the smaller (8—10 cm at

anthesis) spadix and smaller, punctiform raised stigma.

2. Named for Dr Max M.J. van Balgooy who collected the type specimen.

4. 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; Engl. & K. Krause in Engl.,

Pflanzenr. 37 (IV.23B) (1908) 46; Alderw., Bull. Jard. Bot. Buitenzorg III,

1 (1920) 382; Alderw., Bull. Jard. Bot. Buitenzorg III, 4 (1922) 341 —

Epipremnum beccarii Engl., Bull. Soc. Tosc. Ortic. 4: 268 (1879) — Type:

Malaysia, Sarawak, Kuching, Oct. 1865, Beccari p.b. 833 (cited as *832’ by

Engler, 1879) (FI, holo).

Rhaphidophora borneensis Engl, Araceae exsicc. et illustr.n. 195 & in Bot.

Jahrb. Syst. 7, Beibl. 15 (1886) 1 — Type: Indonesia, Kalimantan, Mindai

to Pramassan, 19 June 1882, H. Grabowski s.n. (B+, BM)

Rhaphidophora fluminea Ridl., J. Straits Branch Roy. Asiat. Soc. 44 (1905)

Rhaphidophora in southern and western Archipelago 121

186; Engl. & K. Krause in Engl., Pflanzenr. 37 (1V.23B) (1908) 37 — Type:

Malaysia, Sabah, Bongaya, Dec. 1897, Ridley s.n. (SING, holo).

Figure 5

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 + appressed shingle-leaves, later with leaves erect or spreading and at

this stage plants resembling the 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; /eaves distichous, appressed and shingling to erect or spreading and

scattered on pre-adult shoots, tending to become distally clustered on adult

shoots; cataphylls and prophylls membranous, 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

semipersistent 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,

subcoriaceous to slightly fleshy, base decurrent, apex acuminate with a

moderately prominent tubule; midrib prominently raised abaxially, sunken

adaxially; primary venation pinnate, raised abaxially, slightly impressed

adaxially; interprimaries subparallel 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 midrib and crossing the

primaries and interprimaries; inflorescence one to three together, subtended

by a prominent cataphyll degrading into 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; stylar region rather well developed, mostly

trhombohexagonal, 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.

122 Gard. Bull. Singapore 52 (2000)

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Figure 5. Rhaphidophora beccarii (Engl.) Engl.

A. habit x +: 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.

Rhaphidophora in southern and western Archipelago 123

Distribution: Southern Thailand, Peninsular Malaysia, Sumatera (including

Lingga Archipelago) and throughout Borneo

Habitat: Rheophytic on rocky, wooded streambanks, on rocks in streams

and rivers, in soft mud, sandy streambeds 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 two or three obligate rheophytic species (the others are

the New Guinea R. wentii Eng. & K. Krause and possibly the Sumatran R.

araea), R. beccarii is immediately recognized by its adult growth form, its

habit of creeping along watercourses or being 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

leaved stenophyllous to 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 33128 is

an excellent example of the latter phenomenon). They are treated here as

a neotenic manifestation of R. beccarii.

These narrow leaf forms have been confused with R. araea but may

readily be distinguished by the peduncle much shorter than the petiole and

the larger, sessile spadix.

Other specimens seen: SUMATERA. Riau Prov. - Lingga Archipelago,

Pulau Lingga, Resun, Biinnemyer 6785 (BO, K, L), Parbasiran, Marbau,

Bilah, near Bilah Pertama, Toroes 177 (L, US), Labuhan Batu, Kota Pinang,

Si Mandi Angin, S. Kanan, Toroes 4071 (US);; North Sumatera Prov. -

Sibolangit, Boren Bila to Aek Buru, Lorzing 9717 (BO), Lorzing 12240

(BO), Tapianoeli, Padang Si Dimpoean, Padang Lawas, G. Manuen, Toroes

4594 (US); Padang Si Dimpoean, Padang Lawas, Hatiran, Toroes 4936

(US); Padang Lawas, Sopsopan, Aek Si Olip, Toroes 5559 (L, US); West

Sumatera - Indragira, Taluk region near bivak Dewan I, Meijer 4254 (SING);

5. Rhaphidophora conocephala Alderw.

Rhaphidophora conocephala Alderw., Bull. Jard. Bot. Buitenzorg III, 1

(1920) 384 — Type: Indonesia, Sumatera, North Sumatera, Sibolangit, 10

May 1917, Lérzing 5137 (BO, holo; K, L, iso).

Figure 6

Large, moderately robust, semipachycaul homeophyllous liane to 15 m;

—

124 Gard. Bull. Singapore 52 (2000)

seedling stage a non-skototropic shingling shoot; pre-adult plants forming

small terrestrial colonies of appressed shingling shoots; adult shoot

architecture comprised of elongated, clinging, physiognomically monopodial,

leafy, non-flowering stems and moderately elaborated, free, sympodial,

moderately leafy, flowering stems; stems smooth, climbing stems rectangular

in cross-section, free stems more or less terete to very weakly four-angled

in cross-section, larger shoot systems pendent under their own weight,

without prophyll, cataphyll and petiolar sheath fibre, internodes to 7 x 1.5

cm on adherent shoots, shorter and less stout on free shoots, separated by

prominent 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 rare, adherent, pubescent:

leaves weakly spiralled on adherent shoots, slightly scattered-distichous on

flowering shoots; cataphylls and prophylls membranous, very quickly drying

and falling; petiole shallowly canaliculate adaxially, 4—7 x 0.1—0.2 cm,

smooth, with a slight apical and rather prominent basal geniculum; petiolar

sheath prominent, extending to and encircling the apical geniculum, very

swiftly drying and falling to leave a thin continuous scar from the petiole

base, around the top of the apical geniculum and back to the base,

occasionally newest leaves with parchment-like sheath remain briefly

adherent; /amina entire, falcate-lanceolate to falcate-oblong or falcate-

oblanceolate, 10—29.5 x 1.5—7 cm, coriaceous, upper surfaces slightly

glossy, lower surfaces less so, base minutely cordate to subovate to acute

or briefly decurrent, apex subacute with a prominent apiculate tubule,

margins slightly revolute in dried material; midrib raised abaxially, very

slightly sunken adaxially; primary venation pinnate, raised abaxially and

adaxially; interprimaries parallel to primaries, slightly raised abaxially and

adaxially; secondary and tertiary venation slightly raised in dried specimens;

inflorescence solitary, subtended either by a fully developed foliage leaf or

by one or more subfoliar (i.e., developed petiole but atrophied lamina)

cataphyll; peduncle slightly compressed-cylindric, 3—6 x 0.3—0.5 cm; spathe

cigar-shaped, stoutly short-beaked, 7—9.5 x 2—3.5 cm, thickly fleshy,

exterior light yellow, interior darker, swiftly (?) falling at female receptivity;

spadix cylindrical to slightly clavate, very shortly stipitate, light yellow, 4—

5.3 x 1.2—1.5 cm; stipe c. 2 mm long; stylar region well developed, mostly

rounded to rhombohexagonal, 1.2—1.3 x c. 1.2 mm, conical; stigma

conspicuously raised-punctiform, c. 0.2 mm diam.; anthers slightly exserted

at anthesis; infructescence stoutly cylindrical, 6.5—7.5 x 1.8—2 cm.

Distribution: Sumatera, Kalimantan and into the Philippines (Palawan).

Habitat: Damp primary and old secondary forest. 450—1000m altitude.

Rhaphidophora in southern and western Archipelago £25

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Figure 6. Rhaphidophora conocephala Alderw.

A. flowering shoot x +; B. inflorescence, spathe fallen x 1+; C. spadix detail, female receptivity

x 10; D. gynoecium, three quarter view x 6; E. spadix detail, post anthesis x 10; F. portion of

preadult sterile shoot x '/,; G. leaf lamina x +; H. venation detail x 3. A— E,G & H from Nur

SFN 7369; F from Lérzing 11750.

126 Gard. Bull. Singapore 52 (2000)

Note: Very close to R. sylvestris but consistent in the conical style topped

with a prominent, raised button-like stigma. Generally R. conocephala has

more coriaceous leaves compared with R. sylvestris.

Confusion is possible with R. crassifolia (Peninsular Malaysia and

southern Thailand) which differs by the considerably thicker, almost

succulent leaves and the prominently angled and elaborately twisted

adherent stems.

Other specimens seen: SUMATERA. North Sumatera - Sibolangit, Lérzing

11750 (BO, K, L). Sibolangit, Bk Semiak, Nur SFN 7369 (BO, K, SING).

6. Rhaphidophora floresensis P.C. Boyce, sp. nov.

Rhaphidophora floresensis simillima speciebus R. apiculata, R. hollrungii,

R. sabit et R. ustulata , quae petiolis longis et cataphyllis chartaceis distinctae

sunt; a his speciebus R. floresensis differt inflorescentiis in surculis lateralibus

adhaerentibus, spadice sessili, stigmatibus manifeste elevatis — TYPUS:

Indonesia, Nusa Tenggara, Flores, Sessok, 15 Sept. 1975, E. Schmutz 3837C

(L, holo).

Figure 7

Large, moderately robust, semi-leptocaul (?) homeophyllous (?) liane

(ultimate height unknown); seedling and pre-adult plants unknown; adult

shoot architecture not fully known but apparently comprised of clinging,

physiognomically monopodial, leafy, non-flowering stems and long,

moderately elaborated, free, sympodial, densely leafy, flowering stems;

stems smooth, free stems, + terete in cross-section, without fibre at the tips

of active shoots, internodes to 3 x 1 cm, separated by large + straight leaf

scars; roots unknown; /eaves spiral-distichous on free shoots; cataphylls

and prophylls conspicuous, membranous, very quickly drying and falling:

petiole deeply grooved adaxially, 11—22 x 0.2—3.5 cm, smooth, apical and

basal genicula well defined; petiolar sheath prominent, extending to the

apical geniculum, swiftly drying and leaving a slightly corky scar; lamina

entire, elliptic to falcate-lanceolate, 21—24 x 4—7 cm, very thinly coriaceous,

base acute, apex acuminate with a prominent apiculate tubule; midrib raised

abaxially, slightly sunken adaxially; primary venation pinnate, barely visible

in dried specimens; interprimaries parallel to primaries, almost invisible:

secondary and tertiary venation only just visible as a faint reticulum;

inflorescence solitary, subtended by a large papery cataphyll; peduncle

strongly compressed cylindric, c. 7 x 0.3—0.5 cm; spathe canoe-shaped,

stoutly very long-beaked, 9—12 x c. 2 cm, stiff-leathery, swiftly falling (at

Rhaphidophora in southern and western Archipelago

Figure 7. Rhaphidophora floresensis P.C. Boyce

- D. inflorescence. x

A. flowering shoot x +: B. leaf lamina x +: C. venation detail x

fg =e

spadix detail, late anthesi

500-7

3837C.

cium, three quarter view x 6. All from Schmut

x 6; F. gynoe

128 Gard. Bull. Singapore 52 (2000)

female receptivity?), yellow; spadix slightly tapering-cylindrical, 6.5—8.5 x

1—1.3 cm sessile, inserted level on peduncle; stylar region rhombohexagonal,

c. 1.8 mm diam., stigma punctiform, raised, c. 0.3 mm diam.; anthers barely

exserted at anthesis; infructescence c. 9 x 1.5 cm.

Distribution: Nusa Tenggara (Pulau Flores). Endemic.

Habitat: Forest. 610 m altitude.

Notes: 1. One of a group of species that includes R. apiculata Engl. (New

Guinea), R. hollrungii K. Krause (New Guinea), R. sabit P.C. Boyce and

R. ustulata P.C. Boyce, distinguished by long petioles and chartaceous

cataphylls. Species in this group can be distressingly similar in overall

appearance and are mostly separated on characters such as inflorescence

position, presence or absence of a spadix stipe and whether the top of

stylar region is truncate or conical. The meagre number of botanical

collections for each species exacerbates the problem of ‘sameness’.

Rhaphidophora floresensis may be distinguished by its flowering on adherent

side shoots, by the sessile spadix and the prominently raised stigma.

2. Named for Flores Island where it is endemic.

Other specimens seen: NUSA TENGGARA: Pulau Flores - Ruteng,

Verheijen 2973 (L), Adjang, Verheijen 3024, (L).

7. Rhaphidophora foraminifera (Engl.) Engl.

Rhaphidophora foraminifera (Engl.) Engl., Pflanzenr. 37 (I1V.23B) (1908)

45; Alderw., Bull. Jard. Bot. Buitenzorg III, 4 (1922) 197 — Epipremnum

foraminiferum Engl. in Bot. Jahrb. Syst. 25 (1898) 11 — Neotype: Malaysia,

Perak, Taiping, base of Maxwell’s Hill Station road, Nicolson 1047 (US,

neo; L, BH, BO isoneo; designated by Boyce, 1999).

Figure 8 & 9

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 laminas, inflorescences, internodes 1—7 x 0.5—3.5 cm, separated

130 Gard. Bull. Singapore 52 (2000)

< Guan

Figure 9. 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 southern and western Archipelago iat

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; Jeaves distichous;

cataphylls and prophylls membranous, 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;

lamina entire to slightly or extensively perforated, perforations round to

rhombic, extending c. quarter 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, membranous to subcoriaceous,

base rounded, acute to slightly decurrent, apex acute to acuminate; midrib

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 subparallel 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 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: Sumatera, Peninsular Malaysia and throughout Borneo.

Habitat: Disturbed forest, moist hill dipterocarp forest, on rocks in shaded

position. S0O—1035 m altitude.

Notes: 1. Confusion can occur between R. foraminifera and R. puberula.

They are distinguished by the position of the inflorescence (on short

adherent shoots in R. foraminifera, on short free shoots in R. puberula), by

132 Gard. Bull. Singapore 52 (2000)

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.)

2. Perforated leaves occur in a number of otherwise rather different-looking

Rhaphidophora species (e.g., R. foraminifera. R. puberula, R. ledermannii

Engl. & K. Krause, R. veersteegii Engl. & K. Krause) suggesting that while

a useful diagnostic tool, lamina perforation cannot be used to define groups

within Rhaphidophora.

Other specimens seen: SUMATERA. Bengkulu Prov. - Air Siman, DS

Napal Putih, Kec. Ketahun, Afriastini 2652 (BO); Aceh Prov. - between

Lau Simerah & Lau Penanggajan, Alston 14509 (BM); G. Leuser N.R., c.

35 km NW of Kutatjane, 3 - 5 km upstream Lau Ketambe, de Wilde & de

Wilde-Duyfjes 12061 (L), de Wilde & de Wilde-Duyfjes 14592 (L), G. Leuser

N.R., upper Mamas river valley, c. 15 km w of Kutacane, Camp Pawang,

de Wilde & de Wilde-Duyfjes 18426 (L); North Sumatera - Sibolangit

Loérzing 5521 (BO, L, K), Lorzing 12534 (BO, L) Sibolangit, Cult Bogor

ex Docters van Leeuwen (BO).

8. Rhaphidophora koordersii Engl.

Rhaphidophora koordersii Engl. in Bot. Jahrb. 25 (1898): 6; Engl. & K.

Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 43—44 — Type: Indonesia,

Sulawesi, Minahassa, Kawewatoe, 27 Feb. 1895, Koorders 16166 (B, holo;

BO, L, iso).

Figure 10

Slender, leptocaul, homeophyllous (?) liane (ultimate height unknown);

seedling and pre-adult plants not observed; 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 not

observed, free stems terete in cross-section, with weak prophyll, cataphyll

and petiolar sheath fibre, internodes to 2 x 0.7—0.8 cm on free shoots;

leaves weakly distichous; cataphylls and prophylls membranous, very quickly

drying and degrading to weak fibres and sheets of tissue; petiole weakly

canaliculate, 7—10 x 0.15—0.3 cm, smooth, with a slight apical and

Rhaphidophora in southern and western Archipelago 133

: SESS

Y j SA

L. GUAR

Figure 10. Rhaphidophora koordersii Engl.

A. flowering shoot x +; B. leaf lamina x +; C. venation detail x 2; D. inflorescence, x 1; E. spadix

detail, female receptivity x 8; F gynoecium, three quarter view x 8. All from Riedel s.n..

134 Gard. Bull. Singapore 52 (2000)

prominent basal geniculum; petiolar sheath very prominent, extending

beyond the apical geniculum by two very small ligules, swiftly drying and

degrading into weak fibre and soft strips, eventually falling to leave a

continuous scar from the petiole base, around the top of the apical

geniculum and back to the base; /amina entire, elliptic-lanceolate to oblong-

elliptic, stiffly coriaceous, 7—20 x 3—6 cm, base cuneate, apex acuminate

to attenuate with a prominent tubule; midrib raised abaxially, slightly sunken

adaxially; primary venation pinnate, slightly raised on both surfaces:

interprimaries subparallel to and about as prominent as primaries; secondary

venation + invisible in dried specimens; inflorescence solitary, subtended

by a fully developed foliage leaf; peduncle slightly compressed-cylindric,

4—]2 x 0.2—0.3 cm; spathe narrowly boat-shaped, stoutly long-beaked, c.

10 x 1.6 cm, coriaceous, falling to leave a large, straight, scar; spadix slender

cylindrical, 5—6 x c. 0.7 cm, stipitate; stipe 1.1—1.3 x c. 0.15 cm; stylar

region rather well developed, mostly rhombohexagonal, c. 1 x 1.2 mm,

conical; stigma punctiform, c. 0.3 mm diam.; anthers exserted at anthesis;

infructescence oblong-cylindric, 5.5—9 x 1.5—1.7 cm.

Distribution: Sulawesi (Minahassa Peninsula). Endemic.

Habitat: Unknown. 55 m altitude.

Note: Superficially similar to the widespread R. montana, Bornean R.

elliptica Ridl. and R. elliptifolia Merr., but readily distinguished by the

prominently stipitate spadix, raised stigma, the smaller, elliptic-lanceolate

to oblong-elliptic, stiffly coriaceous leaves with (in dried material) more

prominently raised primary venation and the fibrous cataphyll remains

subtending the inflorescence.

Other specimens seen: Sulawesi, Minahassa, Gorontalo, Riedel s.n. (BO,

K).

9. Rhaphidophora korthalsii Schott

Rhaphidophora korthalsii Schott, Ann. Mus. Bot. Lugd.-Bat. 1(1863) 129;

Engl. in A. & C. DC., Monogr. Phan. 2 (1879) 246; Engl. & K. Krause in

Engl., Pflanzenr. 37 (IV.23B) (1908) 49—S1, Fig. 21; Koorders, Exkursfl.

Java, 1 (1911) 255; Alderw., Bull. Jard. Bot. Buitenzorg III, 4 (1922) 341;

Backer, Beknopte Fl. Java, 17 (1957) 15; Backer & Bakh. f, Fl. Java 3

(1968) 107 — Type: Indonesia, Java, P.W. Korthals s.n. (L, holo; L, P, iso).

Pothos celatocaulis N.E. Br., Gard. Chron. 13 (1880) 200 — Rhaphidophora

Rhaphidophora in southern and western Archipelago 135

celatocaulis (N.E. Br.) Alderw., Bull. Jard. Bot. Buitenzorg II, 1 (1920)

382 & Bull. Jard. Bot. Buitenzorg III, 4 (1922) 198 — Type: Malaysia,

Sabah, Burbidge s.n., Hort. Veitch no. 215 (K, holo; K, iso).

Rhaphidophora maxima Engl., Bull. Soc. Tosc. 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 — Type: Sarawak, G. Gading, July

1866, Beccari PB 2314 (FI, lecto, selected by Boyce, 1999).

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 — Types: Malaysia, Sarawak, Beccari PB 1977 (FI

lecto; B isolecto; selected by Boyce, 1999).

Rhaphidophora korthalsii Schott var. angustiloba Ridl. ex Engl. & K. Krause

in Engler, Pflanzenr. 37 ([V.23B) (1908) 49 — Type: Malaysia, Sarawak,

Matang, July 1903, Ridley s.n. (SING, lecto; selected by Boyce, 1999).

Rhaphidophora copelandii Engl., Bot. Jahrb. Syst. 37 (1905) 115; Engl. &

K. Krause in Engl., Pflanzenr. 37 (1V.23B) (1908) 49 — Type: Philippines,

Mindanao, Davao, Mt Apo, April 1904, Copeland 1193 (PNH, holovt; 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, holoy; B, E, K, L, LE, MO, iso).

Rhaphidophora trinervia Elmer, Leafl. Philipp. Bot. 8 (1919) 3073 — Type:

Philippines, Laguna, Los Ba‘os (Mt Maquiling), June—July 1917, Elmer

18057 (PNH, holo7y; FI, K, L, MO, P, iso).

Rhaphidophora ridleyi Merr., J. Str. Br. Roy. As. Soc. Special Edition

(Enum. Pl. Borneo) (1921) 90 — Rhaphidophora grandis Ridl., J. Straits

Branch Roy. Asiat. Soc. 49 (1907) 51, nom. illeg., non Schott 1858 [India =

R. decursiva (Roxb.) Schott)] — Type: Malaysia, Sarawak, Tambusan, Sept.

1905, Ridley 12414 (SING, holo).

Rhaphidophorea latifolia Alderw., Bull. Jard. Bot. Buitenzorg III, 4 (1922)

341 —Type: Indonesia, Irian Jaya, Pionierbivak, 23 July 1920, Lam 711]

(BO, holo; L, iso).

Rhaphidophora palawanensis Merr, Philipp. J. Sci. 26 (1925) 451 — Type:

136 Gard. Bull. Singapore 52 (2000)

Philippines, Palawan, Malampaya Bay, Oct. 1922, Merrill BS 11570 (PNH,

holoy; B, K, P, iso)

Rhaphidophora trukensis Hosok., J. Jap. Bot. 13 (1937) 195 — Type:

Federated States of Micronesia, Chuuk (Truk) Island, near Orrip, 29 July

1939, Hosokawa 8334 (TI, 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)].

Figure 11 & 12

Very large, occasionally enormous, slender to rather robust, pachycaul,

heterophyllous liane to 20 m; seedling stage a non-skototropic, shingling

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 tips, internodes to 15 x 3.5 cm, separated by

prominent oblique leaf scars, older stems subwoody; 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

membranous, soon drying and degrading into 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, membranous, 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; /amina 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, membranous to

chartaceous or subcoriaceous, base truncate and very briefly decurrent,

apex acute to acuminate, individual pinnae 1—10 cm wide, frequently

perforated basally adjacent to the midrib, thus appearing stilted; midrib

very prominently raised abaxially, slightly sunken adaxially; primary

venation pinnate, raised abaxially, somewhat impressed adaxially, 2—4

primary veins per pinna; interprimaries subparallel to primaries, slightly

raised abaxially, slightly impressed adaxially; secondary venation strongly

reticulate, slightly raised; tertiary venation invisible; inflorescence solitary

Rhaphidophora in southern and western Archipelago 137

(ern

(eae

BAN ANNI SSS

+A vag

L. GUAR.

Figure 11. Rhaphidophora korthalsii Schott

A. pre-adult shoot x +; B. pre-adult shingling shoot x +. A from Boyce 679; B from Nicolson

gL em

138 Gard. Bull. Singapore 52 (2000)

Figure 12. 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 southern and western Archipelago 139

to several together, first inflorescence subtended by a membranous 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—3.5 cm, dark green ripening to dull orange,

stylar tissue abscissing to reveal orange ovary cavity pulp.

Distribution: Sumatera, Java, Sulawesi and Maluku (Pulau Batjam and

Ceram). Widespread in south tropical 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 submontane 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,

with an 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 the 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 sublustrous, 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

140 Gard. Bull. Singapore 52 (2000)

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 vs. trapezoid) and the

shape and orientation of the stigma (+ punctiform and circumferential vs.

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 iF. pinnatum has fruits with two large, strongly

curved seeds with a bony and ornamented testa.

4. Confusion is possible between R. korthalsii and Amydrium zippelianum

D.H. Nicolson although there is a suite of characters that distinguish them.

The leaflet tips of R. korthalsii are truncate, those of the Amydrium species

are acute to acuminate; the petiolar sheath in R. korthalsii extends to the

apical geniculum while in Amydrium the sheath only reaches to the top of

the basal geniculum, the remainder of the petiole being terete with two

sharply defined low keels running its length to merge with the base of the

leaf lamina. The feeding roots of R. korthalsii are prominently scaly, while

those of A. zippelianum are smooth. Fruiting material of R. korthalsii has

the stylar region abcissing to reveal a pulp cavity with numerous, small,

ellipsoid seeds, whereas A. zippelianum has one or two large reniform to

ovoid seeds in each indehiscent fruits.

Other specimens seen: SUMATERA. Bengkulu Prov. - Air Siman, Napal

Putih, Ketahun, Afriastini 2652A (BO); North Sumatera - Silo Maradja,

Asahan, Bartlett 7220 (US), Lau Kakar, Batten-Pooll SFN s.n. (SING),

Taloen na Oeli, east of Dolok Si Manoek-manoek, near headwaters of

Aek Mandosi, Rahmat si Boeea 10265 (L), Karohoogulake, north of

Brastagi, Lérzing 6787 (BO), Brastagi, Yates 1537 (K, UC), G. Sago,

Bukit Pinago, near Pajakumbuh, Meijer 4977 (SING); Aceh Prov. -

Wassemar, Batten-Pooll SFN s.n. (SING), Badjatinggi, south of

Tebingtinggi, L6rzing 7597 (BO), G. Leuser N.R., climbing G. Bandahara,

c. 5 km NE of Kg Seldok (Alas valley), c. 25 km N of Kutatjane, camp 1,

descent Bandahara, de Wilde & de Wilde-Duyfjes 15049 (L); West Sumatera

- G. Merapi, Biinnemyer 4999 (BO); Jambi Prov. - Palembang, Semangus,

Buwalda s.n. (BO, L), Berbak Reserve, Air Hitam Laut, Franken & Roos

249 (L); JAVA. ‘Java’, Herb. Hasskarl s.n. (L); West Java - Cibadak

(‘Tjibadak’), Bakhuizen van den Brink 3886 (BO, L), Bogor, G. Pangrangor,

Bakhuizen van den Brink 6254 (BO), Bogor, Campea (“Tjiampea’), Hallier

s.n. (BO), Koorders 30646B (BO, L), Koorders 30647B (BO), Koorders

30726B (BO), Koorders 30646B (BO, L), Koorders 30647B (BO), Koorders

Rhaphidophora in southern and western Archipelago 14]

30726B (BO); G. Boender, van Steenis 4012 (BO), Ciapus (‘Tjapoes’),

Hallier s.n. (BO), ‘near Batavia’, Kollmann s.n. (BM), Takokak, Koorders,

15124B (BO), G. Salak, Koorders 24132B (BO), Raap 160 (L), SULAWESI.

Minahassa - Kelelonde, Soputan Mts, Alston 15830 (BM, BO), Bivak

Pandak, Koorders 19641B (BO); G. Roroka - west slope, Balgooy 3299

(BO, L, GH, US); Lambasang, Biinnemyer 11485 (BO); Talaud Archipelago

- Pulau Karakelang, Pasir Malap, east of Lobo, bank of Kuala Bahewa,

Lam 3019 (L); Pulau Kaebana - Batu Sangia, c. 7 km west of Tangkeno,

McDonald & Ismail 4117 (BO, GH), Mt Nokilalaki, above Kg Toro, Meijer

9436, 9448 (BO); Tapalaeng, Noerkas 466 (BO). MALUKU. Pulau Batjan

- G. Damar, Masoeroeng, Nedi 45 (BO); Ceram, Treub s.n. (BO).

10. Rhaphidophora lobbii Schott

Rhaphidophora lobbii Schott, Prodr. Syst. Aroid. (1860) 379; Engl. in A. &

C. DC., Monogr. Phan. 2 (1879) 240; Engl. & K. Krause in Engl., Pflanzenr.

37 (1V.23B) (1908) 33—34 —Rhaphidophora lobbii Schott, Bonplandia

5(2) (1857) 45, nom. tant. — Scindapsus lobbii (Schott) Ender, Index Aroid.

(1864) 74 — Type: Singapore, Lobb 44 (K, holo).

Figure 13

Small to moderate, somewhat slender, leptocaul, homeophyllous liane to 5

m; seedling leafy at germination and skototropic by alternating series of

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, distally densely spirally distichous on flowering

shoots; cataphylls and prophylls membranous, 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

142 Gard. Bull. Singapore 52 (2000)

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 subovate,

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-cylindric, 2.5—4 x 1—1.2 cm.

Distribution: Southern Thailand, Peninsular Malaysia, Singapore, Sumatera,

throughout Borneo and into Sulawesi.

Habitat: On wet to inundated soils in lowland to hill dipterocarp forest,

peat swamp and fresh water swamp forest. 10—200 m altitude.

Notes: 1. A climber distinctive by the slender, asperous stems, softly leathery

leaves and minutely puberulent spathe exterior, a combination of characters

unknown in any other Malesian 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 habitually found in wet to inundated ground,

an unusual habitat for a monsteroid liane.

Other specimens seen: SUMATERA. North Sumatera - Asahan, Hoeta

Bagasan, Rahmat si Boeea 6930 (US); Aceh Prov. - S & SW of Gg Leuser

Rhaphidophora in southern and western Archipelago 143

Figure 13. 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 (‘Dr King’s Collector’) 10571; B from Boyce

995;D & E from Nauen s.n.;F & G from Alvins 270.

144 Gard. Bull. Singapore 52 (2000)

N.P., P.T. Hargas logging concession, south of Sibulussalam-Gelombang,

just N of crossing of approach road with Lae Batu Batu (a tributary of

Alas river), near Belingtang, de Wilde & de Wilde-Duyfjes 20523, 20523B

(BO, L, US); Jambi Prov. - Berbak Reserve, Air Hitam Laut, Franken &

Roos 249, 273, 291, 292A (L). SULAWESI. Morowali Prov. - 1 km from

Ranu base camp, Grimes 1150 (K).

11. Rhaphidophora maingayi Hook,

Rhaphidophora maingayi Hook.f., Fl. Brit. Ind. 6 (1893) 543; Engl. & K.

Krause in Engl., Pflanzenr. 37 (1V.23B) (1908) 37 — Type: Malaysia,

Malacca, 28 July 1868, Maingay 2972 (K, holo; L iso).

Rhaphidophora apiculata Alderw., Bull. Jard. Bot. Buitenzorg III, 1 (1920)

383, nom. illeg., non K. Krause (1912, New Guinea) — Based on: Indonesia,

Sumatera, North Sumatera, Sibolangit, Borem Bander Ceroe, 15 Feb. 1917,

Loérzing 4733 (BO).

Figure 14

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,

sometimes irregularly terete, little branched, growing moderate lengths,

green, later mid-brown, with very sparse to very copious untidy prophyll,

cataphyll and petiolar sheath fibre at the tips 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; /eaves

distichous to spiral-distichous on adherent and free shoots, those distal on

flowering shoots densely so; cataphylls and prophylls membranous, 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

Rhaphidophora in southern and western Archipelago 145

WP 2

a a =

Figure 14. Rhaphidophora maingayi Hook.f.

A. flowering shoot x '/,; B. portion of adult, sterile shoot x '/,; C. stem cross-section detail x '/,;

D. leaf lamina x +; E. venation detail x 2; F. inflorescence, spathe fallen x +; G. spadix detail,

post anthesis x 8. All from Boyce 664.

146 Gard. Bull. Singapore 52 (2000)

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 semiglossy,

lower surfaces matt, base subacute to decurrent, apex subacute with a

somewhat prominent apiculate tubule, margins very slightly revolute in

dried material; midrib 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 sparse to copious

netted petiolar sheath, prophyll and cataphyll fibre; peduncle compressed-

cylindric, 10O—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; stylar 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 unknown.

Distribution: Southern Thailand, Peninsular Malaysia and Sumatera.

Habitat: Open disturbed forest remnants on steep slopes, on sandstone.

755 m altitude.

Notes: 1. Rhaphidophora maingayi 1s 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 tips.

2. Plants of the R. montana complex with copious prophyll, cataphyll and

petiolar sheath fibre at the tips 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.

3. Alderwerelt’s R. apiculata [a later homonym pre-dated by R. apiculata

K Krause (New Guinea)] is almost certainly conspecific with R. maingayi

(and treated here as such) but is odd in drying with curiously olive-greenish

yellow leaves.

Other specimens seen: SUMATERA. Aceh Prov. - Kaban Dyake, Batten-

Rhaphidophora in southern and western Archipelago 147

Pooll s.n. (SING); Lampang, Bengkulu, South Sumatera borders - Ranau,

Ruttner 273 (BO); North Sumatera - Asahan, Goerach Batu, Yates 169]

(B.C);

12. Rhaphidophora megastigma Engl.

Rhaphidophora megastigma Engl., Bot. Jahrb. 1: (1881): 180 & in Beccari,

Malesia, vol. 1 (1882) 269; Engl. & K. Krause in Engl., Pflanzenr. 37

(1V.23B) (1908) 38—39 — Type: Indonesia, Sumatera, Ayer Montijoer,

360 m, Beccari s.n. (FI holo; B iso).

Robust homeophyllous liane (ultimate height unknown); seedling stage

and pre-adult plants not observed; adult shoot comprised of greatly

elongated, clinging, physiognomically monopodial, leafy, non-flowering

stems and long, free, sympodial, leafy flowering shoots; stems smooth,

subterete, internodes 1—6.5 x 0.7—1 cm, separated by weakly defined leaf

scars; clasping roots c. 2 mm diam., minutely pubescent; feeding roots not

observed; petiole narrowly but deeply channelled adaxially, 1O—18 cm x c.

2 mm, basal and apical geniculi prominent; petiolar sheath rather prominent,

extending 3.5—4 cm along petiole, soon falling to leave a prominent scar;

lamina oblanceolate to narrowly oblong-elliptic, 15—30 x 5—6.5 cm,

coriaceous and rather distinctly discolorous in dried material, apex acute

to very briefly attenuate, base briefly decurrent; midrib prominently raised

abaxially, slightly raised adaxially; primary lateral veins very faint, barely

discernible; peduncle slightly compressed cylindrical, 2.5—3 x 0.25 cm; spathe

ovate-boat-shaped, 4.5—6 x 2—2.5 cm, apex attenuate, falling at anthesis;

spadix sessile, cylindrical, 4—4.5 x c. 1 cm; stylar region c. 1.25 x 2 mm,

stigma + ovoid, | mm diam, + flush with the style, the middle excavated,

the edges prominently raised; anthers barely exserted at anthesis;

infructescence unknown.

Distribution: Sumatera. Endemic.

Habitat: Unknown. Sea level.

Note: Close to R. montana but differing by shorter (4.5—6 cm vs. 9—20

cm) cylindrical spadix and the large (1 mm diam.) deeply excavated stigma.

Other specimens seen: SUMATERA. West Sumatera, Padang, S. Bulu,

Beccari s.n. (F1).

148 Gard. Bull. Singapore 52 (2000)

13. Rhaphidophora minor Hook.f.

Rhaphidophora minor Hook.f., Fl. Brit. Ind. 6 (1893) 544; Engl. & K.

Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 21, Fig. 4 — Type: Malaysia,

Malacca, Griffith 5988 (K, lecto; K, P, isolecto, selected by Boyce, 1999).

Rhaphidophora celebica K. Krause, Notizbl. Bot. Gart. Berlin-Dahlem. 11

(1932) 331 —Type: Indonesia, Sulawesi, northwest of the island, near Toli-

Toli, Jan. 1910, Schlechter 20698 (B, holo).

[Rhaphidophora palawanensis Furtado, nom. nud. in sched. var., non. Merr.

(1925)]

Figure 15

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, distally moderately densely distichous

on flowering shoots; cataphylls and prophylls membranous, very quickly

drying and falling; petiole grooved adaxially, 3—6 x 0.1—0.25 cm, smooth,

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 subovate, 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;

Rhaphidophora in southern and western Archipelago 149

Wily

;

va:

a Ng

0 SON ao.

Patt

O's

adsi>>

o2:-

Figure 15. Rhaphidophora minor Hook,

A. portion of adult, sterile shoot x ok B. flowering shoot x +; C. leaf lamina x as D. venation

detail x 2; E. inflorescence, spathe just beginning to abscise 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.

150 Gard. Bull. Singapore 52 (2000)

interprimaries subparallel 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 fhombohexagonal, 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: Southern Thailand, Peninsular Malaysia, Singapore, Sumatera,

throughout Borneo, Sulawesi and into the Philippines (Mindanao, Palawan).

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 longer beak to the spathe of R.

minor are diagnostic. Dried material of R. minor is notable for the uniformly

pale straw-coloured leaves.

Other specimens seen: SUMATERA. North Sumatera - Tandjung Balai,

Alston 15358 (BM, BO, L, MO); Riau Prov. - S. Gangsal, between Kemang

and Djawi-Djawi, Buwalda 6823 (GH, K, L, SING)

14. 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 HI, 1 (1920) 386—387 — Calla montana Blume,

Catalogus (1823) 62 — Scindapsus montanus (Blume) Kunth Enum. pl. 3

(1841) 64 (‘montana’) — |Rhaphidophora sylvestris (Bl.) Engl. var. montana

(Bl.) Nicolson, comb. ined. in sched. var.| — Type: Indonesia, Java, Antjyal,

Blume s.n. (L, lecto; L, isolecto, selected by Boyce, 1999).

Rhaphidophora angulata (Miq.) Schott, Prodr. Syst. Aroid. (1860) 379 —

Scindapsus angulatus Migq., Fl. Ned. Ind. 3 (1856) 219 — Type: Indonesia,

Rhaphidophora in southern and western Archipelago 15%

Sumatera, Doekoe, P.W. Korthals s.n. (U, holo).

Rhaphidophora fallax Schott, Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 128 —

Type: Indonesia, Java, Zippelius s.n. (L, holo; K, L, P, iso).

Rhaphidophora burkilliana Ridl., Fl. Mal. Pen. 5 (1925) 121 — Type:

Malaysia, Selangor, Batu Caves, 2 Oct. 1922, Nur SFN 8965 (SING, holo;

BM, K, iso).

Figure 16

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;

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, little

branched, growing to short or moderate lengths, green, later mid-brown,

without fibre at the tips 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 spiral-

distichous on adherent and free shoots, those distal on flowering shoots

densely so; cataphylls and prophylls membranous, 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 subacute

to decurrent, apex subacute with a somewhat prominent apiculate tubule,

margins very slightly revolute in dried material; midrib 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

152 Gard. Bull. Singapore 52 (2000)

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; stylar 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: Southern Thailand, Peninsular Malaysia, Singapore, Sumatera,

Java, Nusa Tenggara, Maluku, Sulawesi and throughout Borneo.

Habitat: Open to closed, primary to disturbed secondary forest, often on

steep slopes over granite, sandstone or limestone. 10—600 m altitude.

Notes: 1. A taxonomically difficult species in the Hongkongensis Group

that, although easy to recognize in its typical manifestation (shoot tips

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 that are mostly known from fewer then

three, usually inadequate, collections, all usually collected without shoot

tips. It is the extremes of these forms that have been described as distinct

species.

2. Plants in cultivation at K from North Sumatera may possibly warrant

formal recognition. They are notable in the stems consistently with the

widest side strongly rounded-convex and the narrowest side somewhat

sulcate. In addition, the leaves are thicker-textured than those of typical R.

montana and have a distinctive bluish caste. However, these plants have

yet to flower and for the moment I prefer to refer them to R. montana s.l.

3. The use of traditional morphology has proved a most unsatisfactory

method for delimiting R. montana. With an alpha-taxonomy now proposed,

excellent opportunities exist for further study of the R. montana complex

using macromolecular data, field observations and statistical techniques to

further resolve this group.

Other specimens seen: SUMATERA. Riau, Prov. - Lingga Archipelago,

Pulau Singkep, Kp Raja, Biinnemyer 7114 (L), Aer Kandis (Radja Mas),

near Rantau Parapat, Bila, Toroes 250] (US); West Sumatera - Bk Gajabuih,

about 15 km east of Padang City, Hotta 25436 (KYO), Sidjundjang,

Pajakumbuh, 6 km N of S. Langsat, Meijer 4447 (SING);; North Sumatera

Rhaphidophora in southern and western Archipelago 153

ererAdy

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2,2, o- 642

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‘

Figure 16. Rhaphidophora montana (Blume) Schott

A. flowering stem x +; B. stem cross-section x fe C. pre-adult terrestrial shoot x +; D. leaf

lamina x '/,; 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.

154 Gard. Bull. Singapore 52 (2000)

- Sibolangit, Lérzing 15529 (BO, L). JAVA. ‘Java’, Blume 157 (L); Cult.

Bogor B.G., Y 14, Alston 12635 (BM), Y. 76, Nicolson 883 (US), Y. 76,

Nicolson 944 (BO, US), Kanda s.n. (BO); Tjianten, near Bogor, Backer

25974 (BO), Bogor, G. Pangrango, Bakhuizen van den Brink 7708 (BO),

G. Cibodas, near Campea (‘Tjiampea’), Hallier s.n. (BO), Campea

(‘Tjiampea’), Hallier s.n. (BO), Nicolson 857 (US); West Java, no further

data, Ploem s.n. (BO). Yogyakarta, Preanger, Nanggerang, SW of

Tarikmalaja, Backer 8784 (BO). NUSA TENGGARA. Pulau Flores -

Schmutz 128 (L). SULAWESI. Minahassa - Koorders 16154B (BO, L).

MALUKU. Pulau Sula - Sanana, Bloembergen 4492 (BO, L).

15. Rhaphidophora oligosperma Alderw.

Rhaphidophora oligosperma Alderw., Bull. Jard. Bot. Buitenzorg III, 4

(1922): 340 — Type: Indonesia, Maluku, Pulau Ternate, North Foramadiahi,

5 March 1921, V.M.A. Beguin 1457 (BO, holo).

Figure 17

Large, moderately robust, homeophyllous (?) liane to 15 m; seedling stage

and pre-adult plants unknown; adult shoot architecture comprised of

elongated, clinging, physiognomically monopodial, leafy, non-flowering

stems and long, moderately elaborated, free, sympodial, densely leafy,

pendent flowering stems; stems drying with prominent longitudinally ridges,

terete in cross-section, internodes to 1.2—7 x 0.5—1.5 cm, separated by

large oblique, very prominent leaf scars; flagellate foraging stems, clasping

roots and feeding roots unknown; leaves distichous on free shoots; cataphylls

and prophylls membranous, conspicuous, quickly drying and falling; petiole

broadly canaliculate, 7.5—18 x 0.3—0.4 cm, smooth, apical and particularly

basal genicula well defined, drying darker then petiole; 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 to oblong lanceolate, + falcate, 14—33 x

4—8 cm, thinly coriaceous, base acute, apex acute to long-acuminate with

a slight tubule; midrib raised abaxially, slightly sunken adaxially; primary

venation pinnate, raised abaxially and adaxially; interprimaries parallel to

primaries and only slightly less prominent; secondary and tertiary venation

reticulate, rather prominent in dried specimens; inflorescence solitary,

subtended by a fully developed foliage leaf and one (more?) conspicuous

cataphylls; peduncle compressed-cylindric, 8 x 9 cm; spathe narrowly cigar-

shaped, long stout-acuminate, 6—7 x c. 1 cm, exterior light green; spadix

very slightly tapering-cylindrical, stipitate, 3—4(—7) x 0.6—0.75 cm; stipe

Rhaphidophora in southern and western Archipelago 155

Figure 17. Rhaphidophora oligosperma Alderw.

A. flowering shoot x ’/,; B. leaf lamina x '/,, C. venation detail x 3; D. leaf lamina x '/,; E.

inflorescence, x 1+; E. spadix detail, late anthesis x 10.A & B from /djan & Mochtar 38; D—F

from Pleyte 46.

156 Gard. Bull. Singapore 52 (2000)

3—3.5 mm long; stylar region weakly rhombohexagonal, c. 0.8—0.9 mm

diam., truncate; stigma punctiform, + flat, c. 0.25 mm diam.; anthers well

exserted at anthesis; infructescence tapering-cylindrical, 4.5—9.5 x 0.75—1

cm, ovaries few-seeded.

Distribution: Maluku (Ternate, Tidore, Ceram). Endemic.

Habitat: In old secondary forest. 8300—1500 m altitude.

Notes: 1. Similar in appearance to R. koordersii (Sulawesi) but differing in

the thinner, broader and shorter leaf lamina. In Maluku, the stiptate spadix

and rather small habit make it distinctive.

2. Confusion with R. oligosperma (Sulawesi) is possible but the latter is

immediately distinguished by the much narrower, strongly falcate leaves,

the sessile spadix and conical style.

Other specimens seen: MALUKU: Pulau Ternate - Moseley s.n. (K), G.

Ake Abdos, [djan & Mochtar 38 (BO, GH, K, L), Pleyte 46 (L); Pulau

Tidore - Buku Mala-Mala, Lam 3778 (BO, L); Pulau Ceram - de Vreise &

Teysmann s.n. (L)

16. Rhaphidophora parvifolia Alderw.

Rhaphidophora parvifolia Alderw., Bull. Jard. Bot. Buitenzorg III, 4 (1922):

338 — Type: Indonesia, Maluku, Pulau Ternate, Laguna, 21 Dec. 1920,

Beguin 1259 (BO, holo).

Moderate-sized, slender to somewhat robust, semi-leptocaul, homeophyllous

neotenic liane to 4 m; seedling and pre-adult plants unknown; adult shoot

architecture comprised of clinging, physiognomically unbranched, mostly

densely leafy, sterile stems and abbreviated, free, fertile stems; stems

minutely punctate, rectangular-terete in cross-section, widest side

prominently convex, narrow side sulcate, internodes to 1—7 x 0.25—1 cm,

separated by slight + straight scars, older stems woody; clasping roots arising

from the internodes, pubescent; feeding roots unknown; /eaves distichous,

scattered, appressed to substrate and shingling on adherent shoots, slightly

scattered on free shoots, cataphylls and prophylls unknown; petiole deeply

grooved, 0.2—1.2 x 0.1—0.25 cm, smooth, groove extending into c. | cm

into abaxial lamina surface, apical and basal genicula barely visible; petiolar

sheath soon drying and falling, membranous, slightly ligulate; /amina broadly

subovate to ovate, thinly coriaceous, base rounded, briefly decurrent, apex

Rhaphidophora in southern and western Archipelago 157

obtuse to obtuse-acuminate with a tiny tubule; midrib prominently raised

abaxially, slightly sunken adaxially; primary venation densely pinnate,

slightly raised abaxially and adaxially; interprimaries subparallel to primaries

and barely distinguishable from them; secondary venation reticulate, barely

visible; inflorescence solitary; peduncle terete, c. 2.5 x 0.3 cm; spathe

unknown; spadix cigar-shaped, sessile, inserted + level on peduncle, c. 4 x

0.75 cm; stylar region mostly irregularly rhombohexagonal, 1—1.3 x c. 1.1

mm, truncate; stigma barely raised, rounded, c. 0.3—0.diam.; infructescence

stoutly cigar-shaped, c. 4.5 x 1.2 cm, orange-yellow at maturity.

Distribution: Maluku. Endemic.

Habitat: ‘Jungle’. 280 m altitude.

Note: Very similar to R. hayi P.C. Boyce & Bogner (New Guinea and

northern Australia) and R. pachyphylla K. Krause (New Guinea) (Boyce

& Bogner, 2000) but differing from both by the sessile spadix.

Rhaphidophora parvifolia may be further distinguished from R. hayi by

the thinner leaf laminas, the non-adherent petiolar sheaths and cataphylls,

and by the rounded stigma. From available material, it is not possible to

tell if R. parvifolia has the disarticulating side shoots typical of R. hayi. R.

pachyphylla is readily separable from R. parvifolia by the leaf lamina on

flowering shoots having an acute to cuneate base.

Other specimens seen: INDONESIA: Maluku - Pulau Halmahera, Pasih

Putih, Taylor 2841B (BO).

17. 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; Engl. &

K. Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 23 — Type: Malaysia,

Perak, Scortechini 347 (K, holo; CAL, iso).

Rhaphidophora kunstleri Hook.f., Fl. Brit. Ind. 6 (1893) 546; 548; Engl. &

K. Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 30; Alderw., Bull. Jard.

Bot. Buitenzorg III, 4 (1922) 338 — Type: Malaysia, Perak, Bk Larut, July

1883, Kunstler 4538 (K, lecto, selected by Boyce, 1999).

158 Gard. Bull. Singapore 52 (2000)

Rhaphidophora gracilipes Hook.f., Fl. Brit. Ind. 6 (1893) 545; Engl. & K.

Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 30, Fig. 10; Alderw., Bull.

Jard. Bot. Buitenzorg HI, 1 (1920) 384 — Type: Malaysia, Perak, June

1886, Kunstler 10271 (K, lecto; BM, SING, isolecto, selected by Boyce,

1999).

Rhaphidophora batoensis Engl. & K. Krause in Engl., Pflanzenr. 37 (IV.23B)

(1908) 27 — Type: Indonesia, Sumatera, Kepulauan Batu, 25 Sept. 1896,

Raap 370, (B, lecto; BO, isolecto, selected by Boyce, 1999).

Rhaphidophora hallieri Alderw., Bull. Jard. Bot. Buitenzorg III, 1 (1920)

385; Alderw., Bull. Jard. Bot. Buitenzorg II, 4 (1922) 196 — 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 — Type: Cult.

Bogor Bot. Gard, June 1919, Alderwerelt s.n. (BO, holo).

Rhaphidophora scaberula Alderw., Bull. Jard. Bot. Buitenzorg III, 4 (1922):

196, Fig. y., synon. nov. — Type: Indonesia: Sumatera, Deli, 20 March

1918, Lérzing 5563 (BO - not traced). Alderwerelt’s description, especially

of the leaf lamina being tomentose abaxially, suggests that R. scaberula is

either a form of R. foraminifera with unperforated leaf laminas, or more

likely an exceptionally robust form of the common and widespread R.

puberula. Although I have been unable to trace the type there is a fertile

specimen that is uncontroversially R. puberula, Lérzing 12538 (BO),

annotated as matching Loérzing 5563 and on which basis R. scaberula is

placed into synonymy.

Figure 18

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

159

Rhaphidophora in southern and western Archipelago

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Figure 18. 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.

160 Gard. Bull. Singapore 52 (2000)

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 blah-blah, soon-falling; lamina entire to

slightly perforated, perforations round to rhombic, extending c. a fifth toa

quarter 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 subchartaceous,

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 subparallel 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, smelling

of frangipani and pineapple (fide Hay et al. 9029) subtended by a fully

developed foliage leaf and, if more than one inflorescence, then the 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 to apricot-orange

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, glaucous.

Distribution: Sumatera, Peninsular Malaysia, Nusa Tenggara and throughout

Borneo.

Habitat: On trees and rocks in primary and secondary lowland to mid-

elevation dipterocarp rain forest, often on steep slopes, on granite and

limestone. 15—1200 m altitude.

Notes: 1. Given the overall uniformity of this species, it is quite extraordinary

that R. puberula should have been redescribed no fewer than seven times

based mostly on minor differences of leaf shape.

Rhaphidophora in southern and western Archipelago 161

2. Superficially similar to R. foraminifera, especially in the perforate-leaf

forms, R. puberula can readily be distinguished from R. foraminifera by

the production of inflorescences on free lateral shoots and in having the

active shoot tips 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 /ess perforated than those of R.

foraminifera.

Other specimens seen: SUMATERA. ‘Sumatera’, Soepadmo 257 (BO);

Aceh Prov. - between Lau Simerah and Lau Penanggajan, near Kutatjane,

Alston 14555 (BO, BM), Lau Kelawa, north of Kutatjane, Alston 14632

(BM), G. Leuser N.R., Ketambe, valley of Lau Alas, near tributary of Lau

Ketambe, c. 35 km NW of Kutatjane, de Wilde & de Wilde-Duyfjes 14592

(BO, L, K, KEP, MO, US), Ketambe R.S., Alas River, c. 35 km NNW of

Kutacane. along Guhra River, de Wilde & de Wilde-Duyfjes 18057 (L);

North Sumatera - Silo Maradja, Asahan, Bartlett 7266 (US), Sibolangit,

Karta 30 (B, BO, L); Borem Bandar Baru, Lorzing 5913 (BO, L), G. Sago,

Bukit Pinago, near Pajakumbuh, above Halaban, Meijer 4713 (SING),

Kepulauan Batu, Raap 325 (BO), Tapianoeli, Padang Si Dimpoean, Padang

Lawas, Hatiran, Toroes 4937 (US), Padang Si Dimpoean, Padang Lawas

Subdivision, Pulau Liman, Toroes 5356 (L).West Sumatera - Mentawi

Archipelago, Pulau Siberut, Batten Pooll SFN s.n. (SING), Boden-Kloss

SFN 14470 (K, SING), Mentawi, Pulau Sipora, Boden-Kloss SFN 14713

(BO, K, SING), Mentawi, Pulau Sioban, /boet 450 (BO, L, SING), Bk

Gajabuih, Ulu Gadat, about 15 km east from Padang City, Hotta 26035

(BO, KYO); NUSA TENGGARA.: Pulau Flores - Manggarai, near Ruteng,

Verheijen 592 (L).

18. Rhaphidophora sabit P.C. Boyce, sp. nov.

Similis R. oligosperma (Maluku), sed foliis valde angustioribus valde falcatis,

spadice sessili, stylo conico differt. Forma et proportio foliorum (lamina

saltem septies longior quam lata) in genera unicae — TYPUS: Indonesia,

Sulawesi, Kendari, G. Papalia, Kec. Ranomeeto, 13 Feb. 1986, Mor. Amir

144 (BO, holo; L, iso).

Figure 19

Moderate, robust, semi-pachycaul, homeophyllous (?) liane (?), (ultimate

height unknown); seedling stage and pre-adult plants not observed; adult

shoot architecture not fully known but seemingly comprised of clinging,

physiognomically, non-flowering stems and shorter clinging, sympodial,

162 Gard. Bull. Singapore 52 (2000)

densely leafy, flowering stems; stems drying deeply longitudinally ridged.

internodes on flowering stems 2—3 x c. 1 cm, separated by prominent

straight to slightly oblique leaf scars; clasping roots arising densely from

the nodes, pubescent; feeding roots, unknown; leaves distichous in specimen

seen; cataphylls and prophylls chartaceous, soon falling; petiole deeply

canaliculate, 14—31 x 0.3—0.4 cm, apical and basal geniculum prominent;

petiolar sheath prominent, extending to apical geniculum, + short-persistent,

degrading to chartaceous strips and a few weak fibres; /amina entire,

narrowly falcate-lanceolate to narrowly falcate-oblanceolate, oblique, drying

distinctly discolorous, dark brown above, orange-brown below, 32—46 x

4.5—7 cm, weakly coriaceous, base rounded, decurrent, apex acute with a

prominent apical tubule; midrib prominently raised abaxially, + sunken

adaxially; primary venation densely pinnate, raised abaxially, slightly raised

adaxially; interprimaries parallel to primaries and barely less prominent;

secondary venation reticulate, very slightly raised; inflorescence subtended

by a large chartaceous cataphyll, these swiftly degrading, and falling;

peduncle slightly compressed, distinctly enlarged distally towards spathe

insertion, 10.5—11 x 0.3—0.5 cm; spathe unknown, falling to leave a large

scar at the base of the spadix; spadix slightly tapering-cylindrical, sessile,

obliquely inserted on peduncle, 5 x 1 cm in the single collection known;

stylar region moderately developed, rounded-trapezoid to weakly

trhombohexagonal, c. 1 x 1.5 mm, shortly conical; stigma punctiform, raised,

c. 0.3 mm diam.; anthers not exserted at anthesis; infructescence unknown.

Distribution: Sulawesi. Endemic. Known only from the type.

Habitat: Unknown. 100 m altitude.

Notes: 1. This new species might be confused with R. oligosperma (Maluku)

but it is immediately distinguished by its much narrower, strongly falcate

leaves, the sessile spadix and conical style.

2. The specific epithet is derived from sabit the Malay word for a sickle,

which is in allusion to the long, narrowly falcate leaves.

19. Rhaphidophora sarasinorum Engl.

Rhaphidophora sarasinorum Engl., Bot. Jahrb. 37 (1905) 114; Engl. & K.

Krause in Engl., Pflanzenr. 37 (I1V.23B) (1908) 37, Fig. 15 — Lectotype

selected here: Indonesia, Sulawesi, Mt Maharau, Masarang, 10 May 1894,

Sarasin 232 (lecto, B).

Rhaphidophora in southern and western Archipelago 163

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Figure 19. Rhaphidophora sabit P.C. Boyce

A. flowering shoot x '/,; B. leaf lamina x '/,; C. venation detail x 3; D. inflorescence, spathe

fallen x 1; E. spadix detail, late anthesis x 10; F. gynoecium, three quarter view, early fruiting x

10. All from Amir 144.

164 Gard. Bull. Singapore 52 (2000)

Figure 20

Large, robust, homeophyllous liane to unknown ultimate height; seedling

stage and pre-adult plants unknown; stems smooth, terete in cross-section,

internodes to | x 1.5 cm long, separated by large oblique, very prominent,

slightly shiny leaf scars; flagellate foraging stems, clasping roots and feeding

roots unknown; leaves weakly spiralled on free shoots; cataphylls and

prophylls membranous, quickly drying and degrading into soft fibre, then

swiftly falling; petiole canaliculate, 16—22 x 0.25—0.4 cm, smooth, apical

and basal genicula well defined; petiolar sheath not observed, scar from

(swiftly?) fallen sheath extending to the apical geniculum; /amina entire,

oblique-oblong, 21—27 x 9.5—12 cm, coriaceous, upper surfaces semiglossy,

lower surfaces matt, base obtuse, apex acuminate with a prominent tubule;

midrib raised abaxially, very slightly sunken adaxially; primary venation

pinnate, barely raised abaxially and adaxially; interprimaries parallel to

primaries and barely less prominent; secondary and tertiary venation +

barely visible in dried specimens; inflorescence solitary, subtended by a

fully developed foliage leaf and one (more?) cataphylls; peduncle

compressed-cylindric, 12 x 0.5 cm; spathe ovate-canoe-shaped, stoutly long

beaked, 14.5 x 7 cm (flattened out), thickly stiff-fleshy, exterior green,

turning purple with age, yellow internally, persistent at least until anthesis;

spadix very slightly tapering-cylindrical, very briefly stipitate, 8—10 x 1—

1.5 cm; stipe c. 2 mm long; stylar region very weakly rhombohexagonal,

almost circular, c. 0.8—0.9 mm diam., truncate; stigma punctiform, raised,

c. 0.3 mm diam.; anthers well exserted at anthesis; infructescence unknown.

Distribution: Sulawesi. Endemic. Known only from vicinity of Masarang.

Habitat: Unknown.

Notes: 1. Superficially similar to R. montana, but readily distinguished by

the terete (not rectangular) cross-section of the adherent stems, the briefly

stipitate spadix, and the anthers exserted at anthesis. Further, in R.

sarasinorum the spathe is persistent (swiftly falling in R. montana), with

the spathe exterior turning purple prior to opening, a feature not found in

R. montana and one that is rare in Rhaphidophora as a whole, the species

generally having the spathe yellow, green or orange just prior to female

receptivity.

2. Confusion with R. ternatensis and R. balgooyi is possible. From R.

ternatensis the prophyll, cataphyll and petiolar sheath fibre, primary lateral

veins barely differentiated from the interprimaries and much shorter spadix

165

Rhaphidophora in southern and western Archipelago

wen

Figure 20. Rhaphidophora sarasinorum Engl.

A. flowering shoot x +; B. leaf lamina x +; C. venation detail x 2; D. inflorescence, spathe fallen

x 1; E. spadix detail, late anthesis x 10. All from Sarasin 232.

166 Gard. Bull. Singapore 52 (2000)

stipe are diagnostic. The larger spadix (11—15 cm) and flat stigmas

immediately distinguish Rhaphidophora balgooyi.

3. Engler cited two syntypes. The other, Sarasin 557, Indonesia, Sulawesi,

Mt Maharau, Masarang, October, 1894, is not in B and is presumed

destroyed.

20. 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’); Koorders, Exkursfl.

Java, 1 (1911) 254; Alderw., Bull. Jard. Bot. Buitenzorg III, 1 (1920) 383;

Backer, Beknopte Fl. Java, 17 (1957) 14; Backer & Bakh. f., Fl. Java 3

(1968) 106 — Calla sylvestris Blume, Catalogus (1823) 62 — Scindapsus

sylvestris (Blume) Kunth, Enum. pl. 3 (1841) 64; Mig., Flora Ned. Indié 3

(1856) 187—188 — 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; L, LE, isolecto,

selected by Boyce, 1999).

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

— Rhaphidophora lingulata (Hassk.) Schott, Bonplandia 5 (1857) 45 —

Monstera lingulata (Hassk.) C. Koch ex Ender, Index Aroid. (1864) 74 —

Type: Indonesia, Java, Hasskarl s.n. (not traced, see Boyce, 1999). Neotype

designated here: Indonesia, Java, East Java, Klakah (‘“Gebok Klakka’), 5

Nov. 1844, Zollinger 2500 (BO neo.; LE, P isoneo.) The collection selected

as the neotype is fertile and has narrow lingulate leaves, matching well

Hasskarl’s diagnosis.

Scindapsus aruensis Eng]., Bull. Soc. Tosc. Ortic. 4 (1879) 270 — Type:

Indonesia, Melaku, Aru Islands, Gabu-lengaw, May 1873, Beccari s.n. (FI,

holo).

Rhaphidophora wrayi Hook.f., Fl. Brit. India 6 (1893) 544; 42; Engl. & K.

Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 34 — Type: Malaysia, Perak,

Larut, Besar, April 1882, Kunstler 2939 (K, lecto, selected by Boyce, 1999).

Rhaphidophora gratissima Becc., Nelle Foreste di Borneo (1902) 604 —

Rhaphidophora sylvestris (Blume) Engl. var. obtusata Engl., Malesia 1 (1883)

Rhaphidophora in southern and western Archipelago 167

268 — Type: Malaysia, Sarawak, Kuching, Nov.1865, Beccari PB 952 (FI,

holo; FI spirit 423, K, iso).

Rhaphidophora motleyana Engl. & K. Krause in Engler, Pflanzenr. 37

(1V.23B) (1908) 25 — Type: Indonesia, Kalimantan, Bangarmassin, 1857—

1858, Motley 741 (K, holo).

[Scindapsus lanceolataus Miq. nom. nud. in sched. BO]

[Pothos cucculata Zipp. nom. nud. in sched. L]

Figure 21

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 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 slightly

winged, the surfaces between slightly concave, free stems rectangular to

subterete 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 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 subacute 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

168 Gard. Bull. Singapore 52 (2000)

venation + obscure in fresh material, visible as a faint reticulum in dried

specimens; inflorescence solitary, subtended by a fully developed foliage

leaf; 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.

Distribution: Peninsular Malaysia, Sumatera, Java, Nusa Tenggara,

throughout Borneo and Maluku.

Habitat: Damp to wet, hill to montane forest. 500—1630 m altitude.

Note: Rhaphidophora sylvestris is most similar to R. montana and R.

talamauana. From R. montana it can distinguished by the narrower leaves

and by the shorter, clavate-cylindrical spadix (3.5—6 cm compared with

9—16 cm in R. montana). The flowering shoots with much more prominent

leaf scars, scattered leaves, stems + rectangular (not terete) in cross-section,

and the much shorter spathe beak serve to distinguish R. sylvestris from R.

talamauana.

Other specimens seen: SUMATERA. North Sumatera - Sibual-buali via

Madurana, Sipirok crater margin, Afriastini 2399 (BO), Lau Kakar, Batten

Pooll SFN s.n. (SING), Sibolangit, Lé6rzing 8440 (BO): Bohorok, Schwabe

s.n. (B), Brastagi, Yates 1457 (BO, UC); Aceh Prov. - G. Leuser N.R., G.

Bandahara, track from kampung Seldok north-east to large ‘blang’ S of

summit, c. 25 km NNW of Kutatjane, camp 3, de Wilde & de Wilde-Duyfjes

13007 (K, KEP, L, US), G. Ketambe, 8 - 15 km SW from the mouth of Lau

Ketambe, c. 40 km NW of Kutatjane, camp 2, de Wilde & de Wilde-Duyfjes

13830 (L); West Sumatera - Bk Gajabuih, Ulu Gadat, about 15 km east

from Padang City, Hotta 2529] (BO, KYO), around Sirah Plot, near pass

of Padang - Solek road, Hotta 26414 (BO, KYO), on route from base camp

to Bk Gajabuih plot, about 15 km east of Padang City, Hotta et al. 157

(BO, KYO), Takenson, van Steenis 5957 (BO); West Sumatera, Kerinci

region, base of G. Tudjuh, Meijer 6589 (L). JAVA. ‘Java’ Anie 244 (L), de

Vriese s.n. (L), Horsfield s.n. (K), Junghuhn s.n. (BO), Korthals 155 (L),

Lobb s.n. (K), Reinwardt s.n. (L), Zippel 34 (K); 40 (L), Zollinger 1604

(P); West Java - Nirmala, Backer 11199 (BO, L), G. Ganisan, east of

Bogor, Bakhuizen van den Brink 6163 (BO, L), van Leeuwen 7561 (BO),

Sukabumi, G. Gombong, Greimgas, Dransfield 2055 (BO), Ciapus

169

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Figure 21. 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.

170 Gard. Bull. Singapore 52 (2000)

(‘Tjapoes’), Bogor, Hallier s.n. (BO), Cibodas, Koorders 31545 (BO, L),

Lotsy 364a (BO), Cibodas B.G., Nicolson 894 (BO, US), G. Mas Tea

plantation, west of Puncak Pass, van Steenis 12641 (BO); Yogyakarta -

Preanger, Bandoeng, Tjigenteng, Koorders 26437 (BO, L), G.

Pangentjangan, Koorders 26672 (BO, L), G. Goenten, Wanu (?) 6778 (BO);

Central Java - G. Besar, Tjipang, Koorders 32899 (BO), Banjaemas

Pringambo, Koorders 39169 (L), G. Moendjoek, G. Karay, Koorders 40094

(BO, L), G. Oengaran, van Leeuwen-Reijnvaan 2071 (BO); East Java -

Malang, Ranoe Daroengan, de Groot & Wehlburger 97 (BO);

21. Rhaphidophora talamauana Alderw.

Rhaphidophora talamauana Alderw., Bull. Jard. Bot. Buitenzorg III, 1

(1920) 384 — Lectotype selected here: Indonesia, Sumatera, G. Malintang,

18 July 1918, Biinnemeijer 3592 (BO lecto; L isolecto).

Figure 22

Medium to large, rather robust, semi-pachycaul (?) homeophyllous (?)

liane (ultimate height unknown); seedling stage and pre-adult plants

unknown; 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, climbing free stems terete in cross-section, internodes to

1.5—4.5 x 0.3—0.5 cm on free shoots, separated by very well defined,

strongly oblique leaf scars; flagellate foraging stems, clasping roots and

feeding roots not observed; leaves scattered, distichous on free shoots;

cataphylls and prophylls not observed and presumably very swiftly falling;

petiole grooved adaxially, 3—12 x 0.2—0.3 cm, smooth, apical and basal

genicula well defined; petiolar sheath not observed in entirety, presumably

very swiftly drying, patches of thick tissue observed and sheath 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, oblique-ovate-

lanceolate to obovate-lanceolate, 12—27 x 2—7.5 cm, thinly coriaceous,

upper surfaces slightly glossy, lower surfaces matt, base cuneate, apex acute

to long-acuminate, with a slender 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, faintly visible as a weak

reticulum in dried specimens; inflorescence solitary, subtended by a fully

developed foliage leaf; peduncle strongly compressed-cylindric, 3—5 x c.

0.4 cm; spathe cigar-shaped, stoutly beaked, 6—6.5 x 1—3 cm, green before

172 Gard. Bull. Singapore 52 (2000)

opening: spadix weakly clavate-cylindrical, sessile, inserted slightly obliquely

on peduncle, c. 6 x 1.5 cm; stylar region mostly rhombohexagonal, 1—2 x

1.5—2 mm, truncate; stigma punctiform to very slightly elongate, flat, c.

0.25—0.3 mm diam.; anthers apparently not exserted at anthesis:

infructescence c. 10 x 2.5 cm.

Distribution: Sumatera. Endemic. Known only from West Sumatera.

Habitat: Unknown. 1000—1150 m altitude.

Notes 1: Rhaphidophora talamauana forms part of the Hongkongensis

Group, which typically has climbing and often flowering stems rectangular

in cross-section, somewhat thickened, often almost succulent leaves, and

flower on free lateral shoots. The species of this group are universally

taxonomically intractable and there is still much to be done before they

are properly understood.

2. Very similar to R. sylvestris, but recognized by the flowering shoots with

considerably more prominent leaf scars, scattered leaves, stems terete (not

rectangular) in cross-section, and by the much longer spathe beak.

3. In describing R. talamauana, Alderwerelt cites two conspecific collections.

The other is Biinnemeijer 657 from West Sumatera, G. Talamau, 9 May

1917 (BO). The collection chosen as the lectotype is in flower ( Biinnemeijer

657 is a fruiting specimen) and is thus more useful for the purposes of

identification.

Other specimens seen: SUMATERA. West Sumatera, G. Talamau,

Biinnemeijer 657 (BO).

22. Rhaphidophora ternatensis Alderw.

Rhaphidophora ternatensis Alderw., Bull. Jard. Bot. Buitenzorg III, 4 (1922):

194 — Type: Indonesia, Maluku, Pulau Ternate, Ake Bobtja, 20 Oct. 1920,

Beguin 1004 (BO, holo).

Figure 23

Medium to large, moderately robust, leptocaul to semi-pachycaul

homeophyllous liane to 8 m; seedling stage and pre-adult plants not observed;

adult shoot architecture comprised of greatly elongated, clinging,

physiognomically monopodial, leafy, non-flowering stems and long,

moderately elaborated, free, sympodial, densely leafy, flowering stems

Rhaphidophora in southern and western Archipelago 173

Figure 23. Rhaphidophora ternatensis Alderw.

A. flowering shoot x */,; B. shoot apex with emerging inflorescence x '/,: C. leaf lamina x */,: D.

venation detail x 3; E. inflorescence, spathe fallen x 1; F. spadix detail, female receptivity x 12:

G. gynoecium, side view, female receptivity x 12. All from Eyma 2726.

174 Gard. Bull. Singapore 52 (2000)

pendent under their own weight; stems smooth, longitudinally rugulose

when dry, climbing free stems terete in cross-section, with slight prophyll,

cataphyll and petiolar sheath fibre, internodes 1—2 cm x 0.5—1 cm on

adherent shoots, usually longer and slightly less stout on free shoots,

separated by weakly defined, more-or-less straight leaf scars, older stems

woody; flagellate foraging stems not observed; roots not observed; leaves

spiral-distichous on adherent and free shoots; cataphylls and prophylls

membranous, drying and degrading into modest fibrous masses; petiole

deeply grooved adaxially, 7.5—26 x 0.4—1.5 cm, drying with weak

longitudinal costate, apical geniculum strongly defined and drying slightly

darker than the rest of the petiole; petiolar sheath very prominent, extending

to and merging with the apical geniculum, lower part strongly sheathing

the emerging leaf or inflorescence; /amina entire, lanceolate to ovate-

lanceolate, slightly falcate very slightly oblique, 19—42 x 6—13 cm,

coriaceous, base ovate-cuneate to acute, apex falcate, acute to acuminate,

with a prominent apiculate tubule; midrib raised abaxially, slightly

channelled adaxially; primary venation pinnate, slightly raised abaxially

and adaxially; interprimaries parallel to primaries, considerably less

prominent, very slightly raised abaxially and adaxially; secondary and tertiary

venation reticulate; inflorescence solitary, subtended by a fully developed

foliage leaf and moderate amounts of fibre; peduncle slightly compressed-

cylindric, 5—9 x 0.4—0.5 cm; spathe canoe-shaped, stoutly very short-

beaked, orange, 10—12 x 1.5—3 cm; spadix cylindrical, truncate basally,

stipitate; stipe 6—10 x c. 2 mm; spadix inserted + obliquely on stipe, 5—7.5

x 1—1.5 cm; stylar region mostly rhombohexagonal, 1—1.2 x 0.9—1.2 mm,

truncate, minutely roughened; stigma slightly elongate to rounded, raised,

c. 0.25—0.3 mm diam.; anthers exserted at anthesis; infructescence not

observed.

Distribution: Maluku (Ceram, Ternate). Endemic.

Habitat: Unknown.

Note: Similar to R. balgooyi, but readily distinguished by presence of

cataphyll, prophyll and petiolar sheath fibres, in the significantly more

prominent primary lateral leaf venation, the smaller spadix, and the raised

styles.

Other specimens seen: MALUKU. West Ceram - Riring to Batu Sore,

Sapalewa Cave, Eyma 2627 (BO, L).

23. Rhaphidophora teysmanniana Engl. & K. Krause

Rhaphidophora in southern and western Archipelago ive:

Rhaphidophora teysmanniana Engl. & K. Krause In Engl., Pflanzenr. 37

(IV.23B) (1908) 35, Fig. 13; Alderw., Bull. Jard. Bot. Buitenzorg HI, 1

(1920) 387 — Type: Indonesia, Sulawesi, Panghadjeno, Teysmann 11774

(BO, holo; BO, iso).

Medium slender, leptocaul homeophyllous (?) lane (ultimate height

unknown); seedling stage, pre-adult plants and overall adult shoot

architecture not observed but adult shoots probably consisting of

physiognomically monopodial, leafy, non-flowering stems and long, free,

sympodial, moderately leafy flowering stems pendent under their own

weight; stems smooth, free stems terete to subterete in cross-section, without

prophyll, cataphyll and petiolar sheath fibre, internodes to 1.2—1.8 x 0.4—

0.6 cm on free shoots, separated by weakly defined, slightly oblique leaf

scars; flagellate foraging stems, clasping and feeding roots not observed;

leaves spiral-distichous on free shoots; cataphylls and prophylls membranous,

very quickly drying and falling; petiole broadly canaliculate, 7—10 x 0.2

0.25 cm, smooth, basal genicula prominent; petiolar sheath fallen on all

specimens examined, extending to and encircling the apical geniculum,

presumably 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; /Jamina entire, oblong-lanceolate, subfalcate, markedly

oblique, 25—30 x 6—7 cm, thinly coriaceous, base acute slightly unequal-

acute, apex long-acuminate, with a prominent apiculate tubule; midrib

prominently raised abaxially, slightly sunken adaxially; primary venation

pinnate, prominently raised abaxially and slightly raised adaxially;

interprimaries subparallel to primaries, less prominent, very slightly raised

abaxially and adaxially; secondary and tertiary venation + obscure in fresh

material, visible as a very faint reticulum in dried specimens; inflorescence

solitary, subtended by a fully developed foliage leaf; peduncle terete, c. 2.5

x 0.15 cm; spathe cigar-shaped, stoutly long-beaked, c. 3.8 x 1 cm, thickly

fleshy; spadix weakly cylindrical, sessile, inserted + level on peduncle, c. 3

x 0.8 cm; stylar region mostly rhombohexagonal, 1—2 x 1.5—2 mm, truncate;

stigma punctiform, barely raised, c. 0.25—0.3 mm diam.; infructescence

unknown.

Distribution: Sulawesi. Endemic. Known only from the type.

Habitat: Unknown.

Note: Similar to R. novoguineensis and R. peekelii Engl. & K. Krause (both

New Guinea, Bismark Archipelago, New Ireland, New Britain, Solomon

Is.) but differing in the cylindrical rather than globose to elongate-globose

176 Gard. Bull. Singapore 52 (2000)

spathe and the cylindrical (not ellipsoid) spadix.

24. Rhaphidophora ustulata P.C. Boyce, sp. nov.

In specimimibus herbarii laminis foliorum manifeste discoloribus, petiolis

pallidis, geniculo apicali obscuro insignis est; a R. montana spatha minore

graciliore, spadice distincte breviore et distincte angustato, a R. floresensis

spadice gracili plus angustato, spathe minore, stigmate plano differt —

TYPUS: Indonesia, Sumatera, Bengkulu Prov., Pulau Enggano, Belowa

(‘Boea-Boea’), 6 June 1936, W.J. Liitjeharms 4473 (L, holo; K, iso).

Figure 24

Large, moderately robust, semi-leptocaul (?) homeophyllous (?) liane

(ultimate height unknown); seedling and pre-adult plants unknown; adult

shoot architecture not fully known but apparently comprised of clinging,

physiognomically monopodial, leafy, non-flowering stems and free,

sympodial, densely leafy, flowering stems; stems smooth, free stems, +

terete in cross-section, without fibre at the tips of active shoots, internodes

to 2 x 0.75 cm, separated by large + straight leaf scars; roots unknown;

leaves scattered-distichous on free shoots; cataphylls and prophylls

conspicuous, membranous, very quickly drying and falling; petiole deeply

grooved adaxially, 15—22 x c. 0.2 cm, smooth, apical and basal genicula

well defined, in dry material the apical geniculum drying much darker

(dark brown) than the petiole (pale orange-brown); petiolar sheath very

prominent, extending to the apical geniculum, briefly acute-ligulate, swiftly

drying and falling in chartaceous strips to leave a slightly corky scar; /amina

entire, elliptic to falcate-elliptic-lanceolate, 24—34 x 4—7 cm, thinly

coriaceous, drying strongly discolorous, pale yellow-brown abaxially, mid-

brown adaxially, base acute, briefly decurrent, apex acuminate with a

prominent apiculate tubule; midrib slightly raised abaxially, slightly sunken

adaxially; primary venation pinnate, very slightly raised abaxially and

adaxially; interprimaries parallel to primaries, less prominent, very slightly

raised abaxially and adaxially; secondary and tertiary venation just visible

as an extremely faint reticulum; inflorescence solitary, subtended by a fully

developed foliage leaf with a greatly expanded petiolar sheath is greatly

expanded; peduncle strongly compressed cylindric, 2—3 x c. 0.3 cm; spathe

narrowly canoe-shaped, slender very long-beaked, c. 8 x 1.2 cm, stiff-thinly

leathery; spadix strongly tapering-cylindrical, sessile, inserted level on

peduncle, 7.5—8.2 x c. 1.2 cm; stylar region rhombohexagonal, 1—1.2 mm

diam., truncate; stigma elongate to discoid, + flat, c. 0.3 mm diam.; anthers

not exserted at anthesis; infructescence unknown.

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L.GUAR.

Figure 24. Rhaphidophora ustulata P.C. Boyce

A. flowering shoot x '/,; B. leaf lamina x 1/2; C. venation detail x 4; D. inflorescence, spathe

fallen x 2; F. spadix detail, pre-female receptivity x 14. All from Liitjeharms 4473.

178 Gard. Bull. Singapore 52 (2000)

Distribution: Sumatera (known only from Pulau Enggano). Endemic

Habitat: Forest. 100 m altitude.

Notes: 1. Distinctive in herbarium specimens by the strongly discolorous

leaf lamina, the pale petioles and the dark apical geniculum. R. ustulata is

most similar to R. floresensis from which it differs in its slender, more

tapering spadix, the smaller spathe, and the flat stigma. Confusion with R.

montana is possible. However, R. ustulata differs in the smaller, more

slender spathe and the much shorter and more distinctly tapered spadix.

Herbarium specimens have notably thinner-textured leaf laminas than R.

montana.

2. The epithet, ustulata, Latin for burnt, is in allusion to the diagnostic

dark apical geniculum which, compared with the much paler petiole, gives

the impression of having been scorched.

Doubtful Species

Rhaphidophora moluccensis Engl. & K. Krause

Rhaphidophora moluccensis Engl. & K. Krause in Engl., Pflanzenr. 37

(IV.23B) (1908): 36, Fig. 21 — Types: Indonesia, Cult. Bogor. ex. Maluku,

Wokam, Feb. 1906, Engler s.n. (B+); Indonesia: Cult. Bogor. ex. Maluku,

Wahai, no voucher cited (Bf?).

Medium, moderately robust, semi-pachycaul (?) homeophyllous (?) lane

(ultimate height unknown); seedling stage, pre-adult and adult shoot

architecture unknown; stems smooth, terete in cross-section, internodes

1.5—2.5 cm; roots not observed; petiole deeply canaliculate adaxially, 12—

15 cm long; petiolar sheath prominent, extending to c. 1.5 cm from apical

geniculum, long persistent; /Jamina entire, falcate-lanceolate-elliptic, 25—

30 x 2.5—3.2 cm, thinly coriaceous, base acute, apex acuminate; primary

venation pinnate, slightly raised abaxially and adaxially; secondary and

tertiary venation + obscure parallel to primary lateral veins; inflorescence

unknown.

Distribution: Maluku. Endemic, known only from the type.

Habitat. Unknown.

Rhaphidophora in southern and western Archipelago 179

Note: Known only from two sterile collections, both lost. Based on the

illustration that accompanies the protologue, the long, narrowly falcate

leaf laminas recall R. sabit (q.v.) but differ in the primary lateral veins,

which are more prominently differentiated than the interprimaries (they

are barely differentiated in R. sabit). In the absence of a specimen and

with no fertile parts described, it is impossible to identify Engler’s R.

moluccanum confidently to any known species.

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 and Alistair Hay (NSW)

and Dan Nicoloson (US) for reading the manuscript and providing much

useful critcism.

References

Alderwerelt van Rosenburgh, C.R.W.K.van, 1920. New or noteworthy

Malayan Araceae. Bulletin Jardin Botanique Buitenzorg Ul, 1: 359—

389.

Alderwerelt van Rosenburgh, C.R.W.K.van, 1922. New or noteworthy

Malayan Araceae II. Bulletin Jardin Botanique Buitenzorg Ul, 4: 163—

229.

Boyce, P.C. 1998. The genus Epipremnum Schott (Araceae—

Monsteroideae—Monstereae) in west and central Malesia. Blumea. 43:

183—213.

Boyce, P.C. 1999. The Genus Rhaphidophora Hassk. (Araceae-

Monsteroideae-Monstereae) in Peninsular Malaysia, and Singapore.

Gardens’ Bulletin Singapore. 51: 183 — 256.

Boyce, P.C. & J. Bogner, J. 2000. An Account of Neotenic Species of

Rhaphidophora Hassk. (Araceae-Monsteroideae-Monstereae) in New

Guinea and Australia. Gardens’ Bulletin Singapore. 52: 89 — 100.

Engler, A. & K. Krause 1908. Araceae-Monsteroideae. In A. Engler (ed.),

Das Pflanzenreich. 37 (1V.23B) 4—139.

Engler, A. & K. Krause 1910. Araceae. Nova Guinea 8: 247—252.

180 Gard. Bull. Singapore 52 (2000)

Engler, A. & K. Krause 1912. Araceae. Nova Guinea 8: 805—809.

Hay, A. 1981. Araceae. In R.J. Johns & A. Hay (eds), A Students’ Guide to

the Monocotyledons of Papua New Guinea. Part 1: 31—81. Office of

Forests, Port Moresby.

Hay, A. 1990. Aroids of Papua New Guinea. Christensen Research Institute,

Madang.

Hay, A. 1993. Rhaphidophora petrieana - a new aroid liane from tropical

Queensland; with a synopsis of the Australian Araceae-Monstereae.

Telopea 52: 293—300.

Krause, K. & C.R.W.K.van Alderwerelt van Rosenburgh, 1924. Araceae.

Nova Guinea 14: 210—220.

Merrill, E.D. 1921. A bibliographic enumeration of Bornean Plants. Journal

Straits Branch Royal Asiatic Society, special number, 1—637.

Merrill, E.D. 1923 (‘1922’). Araceae. An Enumeration of Philippine

Flowering Plants. 1: 172—189. Bureau of Printing, Manila.

Merrill, E.D. 1925. Additions to our knowledge of the Philippine flora.

Philippine Journal of Science 26: 447—496.

Miquel, F.A.W. 1856a. Aroideae novae javanicae... Botanische Zeitung

(Berlin) 14: 561—565.

Miquel, F.A.W. 1856b. Araceae. Flora van Nederlandsch Indié. Vol. 3:

174—223. Van der Post, Amsterdam.

Ridley, H.N. 1905. The aroids of Borneo. Journal Straits Branch Royal

Asiatic Society 44: 169 —188.

Schimper, A.F.W. 1903. Guilds in Plant-geography upon a Physiological

Basis, part 2, ch. 2: 192—206. (English translation by W.R. Fisher, revised

and edited by P. Groom & I.B. Balfour.) Clarendon Press, Oxford. ©

Stearn, W.T. 1992. Botanical Latin, Fourth Edition, pp. xiv + 546. David &

Charles, Newton Abbot & London.

Strong, D.R. & Ray, T.S. 1975. Host tree location behavior of a tropical

vine (Monstera gigantea) by skototropism. Science 190: 804—806.

Rhaphidophora in southern and western Archipelago

18]

Index of Exsiccatae

angustata = 1

araea = 2

balgooyi = 3

beccarii = 4

conocephala = 5

floresensis = 6

foraminifera = 7

koordersii = 8

korthalsti = 9

lobbii = 10

maingayi = 11

megastigma = 12

minor=13

montana = 14

oligosperma = 15

parvifolia = 16

puberula = 17

sabit = 18

sarasinorum = 19

sylvestris = 20

talamauana = 21

ternatensis = 22

teysmanniana = 23

ustulata = 24

Afriastini 2399 = 20; 2652A = 9; 2652 = 7; Alston 12635 = 14; 15358 = 13;

15830 = 9; 14509 = 7; 14555, 14632 = 17; Amir 144 = 18; Anie 244 = 20

Backer 8784 = 14; 11199 = 20; 25974 = 14. Bakhuizen van den Brink 3886 =

9; 6163 = 20; 6254 = 9; 7708 = 14; Balgooy 3299 = 9; Bartlett 7220 = 9; 7266

= 17; Beguin 1004 = 22; 1141 = 3; 1259 = 16; 1457 = 15; Bloembergen 4492 =

14; Blume (?) 157 = 14; Blume 178 = 20; Boden-Kloss 14470, 14713= 17,

Rahmat si Boeea 6930 = 10; 10265 = 9; Biinnemeijer 657, 3592 = 21; 4999 =

9; 6785 = 4; 7114 = 14; 11485 = 9; Burley et al. 1719 = 2; Buwalda 6823 = 13;

692247032 = 2

Danser 7770 = 9; de Groot & Wehlburger 97 = 20; de Wilde & de Wilde-

Duyfes 12061 = 7; 13007 = 20; 13634 = 1; 13830 = 20; 14592 = 17; 15049 =

Oo ees, 24s 16154 = © 16426 = TF: 20525, 20525 B= 40 de Vrese &

Teysmann s.n. = 15; Dransfield 2055 = 20

Engler 4001 = 17; Eyma 2627 = 22

Franken & Roos 249, 273,291, 292A = 10

Grimes 1150 = 10

Hotta 25291 = 20; 25436 = 14; 25568 = 1; 26035 = 17; 26414 = 20; Hotta et al.

157 = 20

Iboet 450 = 17; Idjan & Mochtar 38 = 15

182 Gard. Bull. Singapore 52 (2000)

Karta 30 = 17; Koorders 15124B = 9; 16154B = 14; 16166 = 8; 1964/B,

24132B = 9; 26437; 26672 = 20; 30646B, 30647B, 30726B = 9; 31545, 32899,

39169, 40094 = 20; Korthals 155 = 20; 206 = 1

Lam 711, 3019= 9; Lanw3778 = 15;.L6rzing 4733 = 14;3137 = 5S; 5521 = 7;

5913 = 17; 6787 = 9; 7597 = 9; 8440 = 20; 9717411750 = 5; 12534 = 7;

12240 = 4; 15529 = 14; Lotsy 364a = 20; Liitjeharms 4473 = 24

McDonald & Ismail 4117 = 9; Meijer 4254 = 4; 4447 = 14; 4713 = 17; 4977 =

9; 6589 = 20; 9436 = 9; 9448 = 9; Moseley s.n. = 15

Nedi 45 = 9; Nicolson 857, 883 = 14; 886 = 17; 894 = 20; 927 = 1, 933 = 20;

94] = 7; 943 = 14; 944 = 14; Noerkas 466 = 9; Nur SFN 7369 =5

Pleyte 46 = 15

Raap 160 = 9; 325 = 17; 370 = 17, 458 = 14; Ruttner 273 = 11

Sarasin 232 = 19; Schlechter 20698 = 13; Schmutz 128 = 14, 3837C = 6;

Schwabe s.n. = 20; Soedarsono 283 = 1; Soepadmo 257 = 17

Taylor 2841B = 16; Teysmann 11774 = 23; Toroes 177 = 4; 2501 = 14, 4071,

4936 = 4; 4937 = 17; 4594 = 4; 5356 = 17; 5559 =4

van Balgooy 4768 = 3; van Leeuwen 7561 = 20; van Leeuwen-Reijnvaan

2071 = 20; van Steenis 4012 = 9; 5957 = 20; 12641 = 20; Verheijen 592 = 17;

2973, 3024 =6

Wanu (?) 6778 = 20

Yates 1457 = 20; 1537 = 9; 1691 = 11

Zippel 34, 40= 20; Zollinger 1604, 2500 = 20

Rhaphidophora in southern and western Archipelago 183

Index to Rhaphidophora Species

eee ee Pash an seen eect Saved wbbdevnsencccdesndaneces 112

Re USP etait eed Pe ras accra R ae sane tccpuactasd dats devecssreacaneres 116

Nk AUG Aut eae sity TERE ach esis svavveesaeecneseicee 118

Ne cet e y c See Baletissschc Sass ch dav dail divs sv sesauunevesd 120

Re oe aA oe a a hk Vedra psdenseut ts sany sn vindaashivdaveaeseseanectins 123

I Rs A cel ok ecg eds Sivoo bush cena vice cwuagstbninsdewnscdeades 126

TIN ET GOUT ea iy (Wr Ar ed bales AR Op 0 1) oe ee ee ore 128

TO Manali ies IN dal ht Sc lh andes cd haa i ash Gfidandics 42s ovnsdvinsocesseuss 132

Nk ea Frc rack pwsea anette viainn ath icien deeardeviddiveeseoseess 134

a at as catibed ced Pes unnsbOvusi ihdnnidddvolivientsvoinysnonnae 141

NR cagA ade fs 8 Wy Seah ceaiey ates aie roses xe in Adawi shee iade Hsunssvndeietiniss 144

a Rik be AN a ie hs tute ageing «oes xi'gny Xdansine ov azaniddnanctan 147

IIR IML 2 iil Bh Sac thai eit doa A dvbcaltes dun Seca edonge hh dadsvedsdavataveticcnendze 148

tS SAS ase kets | ee et ee ae | ee ee en 178

2 aie nol eh nds at aft ee an IE lbs eG Sie ate ed 150

i Mek celta tah skivcencjnli SAG Gide dN Penne 5 ls nen cpa sa ocr 154

STONES SEES TE OME ABIRE 01E 0) ile: Cate. Se A toe oon eae ne eee a eee ee ee 156

EEE NORE DE ROE ARE Dy ORR SE ee! [NR DO A? ee oe oe Lag

SPU Nia rce eas tec ts ald va vets Pa eaiiapddsivas vs avisadnncserindactesds 161

Tg ee ee te CER ENCE an Cre ae a ee Pee eee 163

ENED 208 sis a SURE Cae Cosi Reds sa Aas ei basbvnanased henscees sachs 166

ces CON a Ne EN Cy ed at.) Sac cna whe Sse ot Sct 22adavend tas 170

OE TG ES ES is. By tos Ca bony et cae ee ee en L/2

Senna RPh 59 oe Ad hie GA a ene aA Wd akecvietosd Wisgsthiainkas and sbse8s see: 174

ST EE niet War od OO peeks Oe Rete SE ete a he ae a 176

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Gardens’ Bulletin Singapore 52 (2000) 185-202.

New and Interesting Plants From Ha Long Bay, Vietnam

NGUYEN TIEN HIEP

Institute of Ecology and Biological Resources NCST,

Nghia Do, Cau Giay, Hanoi, Vietnam

AND

RUTH KIEW

Singapore Botanic Gardens, Singapore

Abstract

Six new species are described from Ha Long Bay, Vietnam: Chirita halongensis Kiew &

T.H. Nguyen, C. hiepii Kiew, C. modesta Kiew & T.H. Nguyen (Gesneriaceae), /mpatiens

halongensis Kiew & T.H. Nguyen (Balsaminaceae), Livistona halongensis T.H. Nguyen &

Kiew (Palmae) and Paraboea halongensis Kiew & T.H. Nguyen (Gesneriaceae). In addition,

notes on habit and habitat for Chirita drakei B.L. Burtt, C. gemella D. Wood and C.

hamosa R. Br. are provided.

Introduction

Ha Long Bay, a World Heritage Site, 165 km northeast of Hanoi, is

renowned for its spectacular seascape with tower karst limestone islands

dotting the wide bay. Of the nearly 2000 islands, 788 are included within

the 434 square kilometer World Heritage Site.

While there are some early collections from the area, there has not

been a focus on investigating its flora until the impetus to select 30

characteristic plant species for Wendy Gibbs to paint for a forthcoming

tourist guide of the wild plants of Ha Long Bay (T.H. Nguyen & Kiew,

2000). During the course of exploration, which covered many islands, several

new species were encountered, which are here named and described. A

new ginger, Alpinia calcicola, is described separately by Q.B. Nguyen et al.

(2000). In addition, some species previously poorly known are here provided

with a more detailed description, particularly for their habit, based on

living plants.

Flowering at Ha Long Bay is for most species seasonal with the

majority flowering in the hot, wet summer (May to October) when

temperatures average between 27° and 29°C and the rainfall in June, July

and August exceeds 300 mm a month. Rather few flower in the cool, dry

winter when temperatures drop to between 16° and 18°C and there is less

186 Gard. Bull. Singapore 52 (2000)

than 30 mm rainfall in each of December, January and February.

Early collectors named several species for Ha Long Bay, viz. Ficus

alongensis Gagnep., Jasminum alongense Gagnep. and Schefflera alongensis

R. Viguier. Of these only the jasmine appears to be endemic to Ha Long

Bay. Other endemics include Chirita drakei B.L. Burtt and C. gemella D.

Wood, as well as Alpinia calcicola Q.B. Nguyen & M. Newman, Chirita

halongensis Kiew & T.H. Nguyen, C. hiepii Kiew, C. modesta Kiew & T.H.

Nguyen, /mpatiens halongensis Kiew & T.H. Nguyen, Livistona halongensis

T.H. Nguyen & Kiew, and Paraboea halongensis Kiew & T.H. Nguyen

from our study. This high level of endemism illustrates the conservation

importance of Ha Long Bay.

Chirita (Gesneriaceae)

Seven species of Chirita Benth. are now known from Ha Long Bay, of

which three are new species described below. Apart from C. hamosa R.

Br. in Sect. Microchirita C.B. Clarke, which has a wide geographic range in

SE Asia, the rest belong to Sect. Gibbosaccus C.B. Clarke and are endemic

to Ha Long Bay.

Of these six, C. modesta is extremely rare and only one small

population was found growing around the mouth of one cave. It most

closely resembles to C. speluncae (Hand.-Mazz.) D.Wood, a Chinese species

known from a single cave in NE Yunnan. C. gemella is also extremely

local. We found it on a single island where it was abundant on one slope.

Previously it was known from a single collection without precise locality

being recorded as from ‘Tonkin’. C. hiepii is not as common as C.

halongensis, which at sea level grows in the same habitat, i.e. in crevices in

the limestone rocks in light shade. The distribution of C. halongensis extends

higher up slopes, where conditions are exposed. C. drakei, the most common

chirita at Ha Long Bay, is a remarkably beautiful shrubby plant with silver-

grey leaves and large, white tubular flowers with violet lobes. It is certainly

a handsome plant and is found everywhere growing on exposed limestone

rocks from sea level up exposed, rocky slopes. This species and C. gemella,

C. halongensis and C. hiepii, that all have attractive purple lobed flowers,

are potential ornamental plants. The seventh species, C. semicontorta

Pellegrin, was not refound during our survey.

Key to Chirita Species of Ha Long Bay

la. Plants large and bushy or with an erect stem and a single leaf or

New Plants from Ha Log Bay 187

ee TL ER ee ec 2

1b. Plants with a short subterranean rhizome and a tuft of leaves at the

ee 1 Ge ws 1 sot ie a eS eB 3

2a. Low bush with woody stem and terminalia branching with tufts of

silky silver-grey leaves at tip, flowers from leaf axils, c. 5 cm long,

WibHEOLTY WIGTET IGNES CLG. i ie. 1. C. drakei

2b. Erect, fleshy plant, at first with one leaf, then producing pairs of

leaves, flowers epiphyllous, c. 1 cm long, flowers dull white ...............

Pe ea ee I SA Bed Asda tdens. 4. C. hamosa

Ga iicaves with petiole lonper than latin 0520... ts lia ee oes ees =

Oe caves avith petiole shorter Mian lamilia..: eed) ev. el hel i ek >

4a. Petioles 4—8.5 cm long, lamina 3—5.5 x 1.8—4.5 cm, fruits 14—22

Dre Wns ALLY Geos. See ee Tai ate, Od aed ke 7. C. semicontorta

4b. Petioles 3.5—4.5 cm long, lamina 2.5—3.8 x 1.4—2.1 cm, fruit 7—12

aI Se ane: oper ch dye es ba Sc katie A hdd: 6. C. modesta

5a. Leaves horizontal in a tight rosette, flowers 1(—4) flowers per stalk

EET a et] Uo Vols, AMEE RA MERE NS cet SEO ay Oe Re Sot, Poe es 2. C. gemella

5b. Leaves upstanding in a lax tuft, inflorescence paniculate with 6—30

Se ee: Bre, cay ae | Ue Se eee 0 aie eee ee eee eee 6

6a. Upper surface of lamina glossy and glabrous, lateral veins on lower

Sitinee Piano land ODSeUre ANI 4... aL I. 3. C. halongensis

6b. Upper surface of lamina densely hirsute, lateral veins on lower surface

gr ay lat) Beate ee, ae ees eee ee ROE oe a a or oe 5. C. hiepii

1. Chirita drakei B. L Burtt. Notes R.B.G. Edinb. 23 (1960) 98.

Nguyen & Kiew, Wild Plants of Ha Long Bay (2000) Plate 22.

Plate Ic

This beautiful species was previously known from three specimens, all

collected from Ha Long Bay. In fact, it is one of the most common and

striking plants there with its long, white tubular, violet-lobed flowers and

silky, silver-grey leaves. It grows in exposed conditions on limestone cliffs,

rocks and screes from sea level to about 100 m altitude.

The early herbarium specimens are misleading as they represent

snippets of shoot tips of this most robust species. It is in fact a low, much

branched shrub rooted deeply in crevices with a gnarled and woody trunk

188 Gard. Bull. Singapore 52 (2000)

3.5—4 cm thick, frequently subterranean, producing thick, horizontal

branches spreading over the rock surface and forming a hemispherical

bush up to | m across and 60 cm tall. These branches can grow up to 45 cm

long and exhibit terminalia branching where a series of tufts of leaves are

produced at the tips. The ultimate twigs are c. 1 cm thick. The internodes

are very short, often with the nodes touching. Presuming only a few tufts

of leaves are produced each year, these plants must be very ancient.

It can be found in flower throughout the year and is very floriferous

in the summer months (July to November) compared with sparse flowering

in winter and early spring. Usually its inflorescences are two-flowered but

occasional three-flowered inflorescences can be found.

The wife of a fisherman told us that all parts of the plants are boiled

and the decoction is drunk after childbirth.

Specimens: Nguyen Tien Hiep & L. Averyanov NTH 2599 12 May 1999

(HN, LE, MO, SING), Nguyen Tien Hiep & Ruth Kiew NTH 2599 15 July

1999 (HN, SING), Nguyen Tien Hiep & Ruth Kiew NTH 4059 11 Nov 1999

(HN, K, LE, MO, SING).

2. Chirita gemella D. Wood. Notes R.B.G. Edinb. 33 (1974) 141.

Plate 1d

Previous to our study, C. gemella was known only from the unicate type

specimen of four plants on a single sheet, (Lemarie 123 - holo P, photo E).

The locality was recorded just as ‘Tonkin’. It is in fact a very rare species

and we found it on a single unnamed island in Ha Long Bay. As we were

able to observe it in both the flowering and fruiting seasons, Wood's

description is here supplemented by our observations of living plants.

Perennial, rosette herb, rhizome subterranean, c. 6—7 cm long x 1.5—1.8

mm wide, sometimes spreading by stolons up to 5 mm thick. Leaves in the

centre of the rosette almost sessile, but the petioles elongate and in the

outer, lower leaves the petioles reach 4.5 cm, /amina stiff and fleshy, dark

green above, whitish green beneath, indumentum of white hairs dense on

both surfaces, especially dense on margin and under surface of veins, 3.5—

5(—9) x 2.2—3.0 cm, lateral veins (2—)3(—4) pairs, in life impressed above,

prominent beneath. Flower solitary, from the axils of the outer, older leaves,

with up to 5 flowers open simultaneously on a single plant, pedicel pale

purple (4.5—)7.5—13.5 cm long, corolla 19—20 x 15—18 mm, white outside,

lobes mauve, inside tube white with bright yellow bilobed patch on upper

surface of the throat, stamens with pale green filaments with tuft of long

New Plants from Ha Log Bay 189

white hairs at top, anthers pale purple. Capsule with persistent calyx,

narrowly cylindrical, 31 x 1.5 mm, pale brown, splitting on upper surface,

style persistent.

Distribution: Vietnam, Quang Ninh Province, Ha Long Bay, only found on

one island.

Habitat: In rock crevices or on thin soil among grass tussocks on a steep

slope above sea level.

Specimens: Nguyen Tien Hiep & L. Averyanov NTH 2597 12 May 1999

(HN - flowers); Nguyen Tien Hiep & Ruth Kiew RK 4745 16 July 1999

(SING - fruits), NTH 4205 11 April 2000 (HN, BM, E, K, LE, MO, SING

- flowers).

Notes: Flowering freely in early spring (April and May), by July it is in

fruit. It differs from the other Chirita species in Ha Long Bay in its very

small, woolly leaves arranged in a tight rosette. All the plants in the

population we found had solitary flowers, as compared with the type

specimen that had up to four flowers on a pedunculate inflorescence, but

apart from this single character our specimens matched the type.

3. Chirita halongensis Kiew & T.H. Nguyen, sp. nov. ~

(Sect. Gibbosaccus C.B. Clarke)

Nguyen & Kiew, Wild Plants of Ha Long Bay (2000) Plate 25.

Chiritae semicontortae Pellegrin similis, sed foliis majoribus, petiolis quam

laminis brevioribus et floribus majoribus differt.

Typus: Ha Long Bay, Nguyen Tien Hiep & Ruth Kiew 4219 12 April 2000 —

flowers (HN holo; E, LE, MO, SING iso).

Plate le

Perennial herb, rhizome large, subterranean, + woody, to c. 11 x 2—2.5 cm

with a corky layer formed by leaf cushions, often branched, internodes

congested. Leaves subopposite, crowded at apex, upstanding and brittle

(snapping easily), glabrous (except for margin), petiole winged, (1—)2.5—

6 cm long, fleshy, flat and slightly grooved above, /amina narrowly elliptic

to slightly obovate, (3—)11—14 x (0.75—)2.5—3 cm, either plain dark

green above and whitish green and suffused deep rosy purple in lower two

thirds beneath or dark green above with midrib and lateral veins white and

completely whitish green beneath or, in totally exposed conditions, purple

190 Gard. Bull. Singapore 52 (2000)

green above and rosy purple beneath; succulent, slightly glossy, apparently

glabrous above but with microscopic, unicellular, eglandular trichomes,

base decurrent into winged petiole, margin finely ciliate with long uniseriate

trichomes, appearing recurved in dried state, entire or subcrenate, apex

acute, lateral veins 3—4 pairs, ascending, slightly raised above, plane

beneath. [Inflorescences up to 3 per axil from older leaves, erect, 24—36

cm, paniculate with second or third order branching and between 8 to 30

flowers, peduncle rosy purple or reddish brown, to 30 cm long; bracts 2,

foliaceous, 6—10 x 1.5—3 mm, green or purplish green, bracteoles similar

but 3—4 x 1—1.5 mm. Peduncle, pedicels, calyx and ovary densely pilose

with minute, glandular hairs, calyx with some short non-glandular trichomes

as well. Pedicels 9—20 mm long, calyx green, divided to base, lobes 1.5—

2.5 x 0.75 mm at base, tapering to acute apex, corolla campanulaie, slightly

saccate at base, minutely hispid outside with short glandular trichomes,

completely glabrous inside, tube 7—10 x 6.5—13 mm, white inside and out

or with 2 golden brown nectar guides inside and faintly yellowish outside,

lobes deep purple on inner surface, white tinged purple outside, upper 2

lobes recurved, lower 3 spreading, c. 5 x 6-7 mm, stamens and staminodes

attached c. 3—4 mm above base of corolla, staminodes c. 1 mm long,

filaments geniculate, white, 2—S mm long, completely glabrous, anthers

brown, c. 3 x 1 mm, fused face to face, disc annular, c. 0.3 mm deep, ovary

pale green, c. 7—8 x 1 mm, style white c. 2 mm long, stigma spathulate and

slightly bifid at apex, exerted just beyond corolla mouth. Fruit with pedicel

13—30 mm long, calyx persistent, capsule narrowly cylindric, reddish brown,

(16—)32—37 x 1 mm, style persistent.

Distribution: Vietnam, Quang Ninh Province, Ha Long Bay, on several

islands.

Habitat: It grows in cracks or among rocks from sea level in semi-shade to

exposed screes near summit of limestone islands.

Notes: Another species, C. semicontorta Pellegrin, was early described from

Ha Long Bay and C. halongensis is similar to it in habit (thick rhizome

with conspicuous leaf scars), in the leaves being more or less glabrous, and

the inflorescences, which have long peduncles and relatively short branches.

C. halongensis 1s clearly different from this species in a number of characters.

The leaves of C. semicontorta are smaller and relatively broader (3—5.5 x

1.8—4.5 cm), the petioles are longer than the lamina, the inflorescence is

much shorter (to 9 cm long) with fewer flowers (3—9 flowers), and the

flowers are very much smaller (10 x 2.5 mm).

Chirita halongensis is a very attractive plant with variegated leaves

New Plants from Ha Log Bay 19]

and many-flowered inflorescences with dangling, purple-lobed flowers. It

flowers in spring. It grows in rock crevices or on rock screes. In exposed

conditions, its leaves are much smaller (c. 4 x 0.75 cm) and are distinctly

purplish on both surfaces, compared with larger plants that grow in light

shade. Populations on different islands exhibit differences in flower colour.

Most have flowers with a completely white corolla tube, but in some

populations the corolla tube is yellowish with two golden brown nectar

guides in the throat.

Specimens: type and Nguyen Tien Hiep & L. Averyanov NTH 2617 11 May

1999 (HN), Nguyen Tien Hiep & Ruth Kiew NTH 4069 10 November 1999

fruits (HN, SING).

4. Chirita hamosa R. Br., Cyrtandreae 117 (xii 1839).

Nguyen & Kiew, Wild Plants of Ha Long Bay (2000) Plate 27.

This species is remarkable for its annual habit and rapid growth. At Ha

Long Bay, we found it growing in soil in the shaded mouth of one cave. In

July, it formed drifts of hundreds of seedlings, ranging in size from plants

c. 4 cm tall with a single leaf c. 2.7 x 1.9 cm with 7—9 pairs of veins to

plants up to 25 cm tall with the single lower leaf c. 21 x 13.5 cm with c. 12

pairs of veins and with an additional pair of small, expanding leaves above.

Even one-leaf plants with a leaf c. 8 x 7 cm were in flower and fruit. By

November, they had grown to robust plants c. 25 cm tall, but on returning

to the cave in April, they had completely died down and not a trace of

them was to be found!

Chirita hamosa is a widespread and variable species (Wood, 1974).

However, there appear to be regional differences in fruit size as plants

from Ha Long Bay have much longer fruits (S—7.75 cm long) than the

many specimens from Thailand, which have fruits c. 2 cm long. One

specimen from Laos, Poilane 15951 (K), has fruits 3—4 cm long and the

type specimen from Martaban, Myanmar, has fruits 6 cm long (B.L. Burtt,

pers. comm.).

The Ha Long population is also different in flower size and colour. It

has flowers 15—25 mm long and 7—9 mm wide (compared with 7—11 mm

long and 4 mm wide in C. hamosa s.s.) and, while some plants have white

flowers with a lemon yellow patch in the throat, others have very pale

mauve ones. There appear to be seasonal differences in flower colour as in

July all the plants had white flowers, while in November some had mauve

flowers.

Long fruits (7 cm long) and lilac blue flowers are characteristic of C.

192 Gard. Bull. Singapore 52 (2000)

aratriformis D. Wood from Khaumoi, Langson Province, Tongkin, but the

Ha Long plants would not be confused with this species, which has axillary

(not epiphyllous) flowers that are larger (28 by 12 mm) and besides it is a

large plant to 60 cm tall with proportionately smaller leaves up to 11 by 9

cm.

Specimens: 16 July 1999 Nguyen Tien Hiep & Ruth Kiew NTH 3906 (HN),

NTH 3927 (HN), NTH 3937 (HN), RK 4744 (SING): 9 Nov 1999 NTH

4051 (E, HN, LE, MO, SING).

5. Chirita hiepii Kiew, sp. nov.

(Sect. Gibbosaccus) C.B. Clarke

Nguyen & Kiew, Wild Plants of Ha Long Bay (2000) Plate 25.

Chiritae halongensi Kiew & T.H. Nguyen similis, sed foliorum pilis densis

et albis, nervis prominentibus et fructibus brevioribus differt.

Typus: Ha Long Bay, Nguyen Tien Hiep & Ruth Kiew NTH 4208 11 April

2000 (HN holo: BM, E, K, LE, MO, SING iso)

Plate If

Perennial herb, softly hairy with straight white, uniseriate, eglandular

trichomes, c. 4—6 mm long, particularly dense on petiole, leaf margin,

undersurface of lamina, and sparse and 1 mm long on upper surface of

lamina, peduncle and calyx; peduncle and calyx with dense layer of short,

glandular hairs; rhizome subterranean, + woody, c. 3—5 cm long and 2

2.5 cm thick, internodes congested and the leaves forming a compact tuft

at the apex. Leaves subopposite, at first sessile, petiole elongating to 3—5.5

cm in the outer leaves, fleshy, flat or grooved above, /amina lanceolate,

7.5—9 x 3.5—4.2 cm, mid-green above, whitish green beneath, succulent,

base narrowing or decurrent, margin subcrenate to entire, apex acute or

blunt, lateral veins 34 pairs, plane or slightly impressed above, midrib

and veins very prominent beneath. /nflorescence one per older leaf, erect,

13—25.5 cm long, paniculate with second or third order branching and up

to c. 6 or 14 flowers respectively, peduncle 2—5.5 cm long, pale brownish-

green; bracts in whorl of 3 at top of the peduncle, mid-green or pale

brownish-green, linear, c. 9 x 1.5 mm, apex rounded, trichomes short

glandular, bracteoles similar but c. 3 x 0.75 mm. Pedicels of lower flowers

15—30 mm long, of upper 7—16 mm long, calyx densely hairy, mid or pale

green, divided to base, lobes c. 2—3 mm long, tapering to an acute apex,

corolla straight, slightly pouched, outer surface with sparse layer of short

glandular trichomes, tube white, 5—9 x 5—7 mm, inner lower surface of

New Plants from Ha Log Bay 193

throat with two bands of glandular hairs, lobes purple, spreading, rounded,

isomorphic, 4—6 x 3—6 mm, stamens and staminodes attached c. 1 mm

above base of corolla tube, staminodes c. 2—3 mm long, filaments

geniculate, c. 6 mm long, sparsely hairy below anther, anthers brown, c. 2.5

x 1 mm, fused face to face, disc annular, c. 0.5 mm deep, ovary reddish

brown, pilose with dense layer of short glandular trichomes, c. 6—10 mm

long with slender style c. 4 mm long, stigma spathulate, shallowly bifid at

apex, white or pale green. Fruit with pedicel 1.1—4.5 cm long, calyx

persistent, capsule narrowly cylindric, 13—15 x c. 1.5 mm, stigma and style

persistent.

Distribution: Vietnam, Quang Ninh Province, Ha Long Bay, rare, found

on only one island.

Habitat: Its rhizome roots into cracks in limestone rocks above the high

tide mark in semi-shade.

Notes: This species is not as common or as widespread as C. halongensis,

which it most closely resembles in habit (thick rhizomes that penetrate

rock crevices), crowded, narrowly lanceolate, fleshy leaves and panicles of

pretty purple-lobed flowers held above the leaves. It is distinct from C.

halongensis in several characters, the most striking of which is the dense,

soft, white indumentum on both leaf surfaces and the very prominent

veins on the lower surface. (The leaves of C. halongensis, in contrast, are

on the upper surface glabrous to the naked eye and the veins are plane on

the lower surface). In addition, the flowers of C. hiepii do not have reflexed

upper corolla lobes, there are two bands of short glandular hairs in the

throat of the corolla tube, the stamens are long and attached near the base

of the corolla, the filaments are hairy at the top, and the capsule is short.

In contrast, C. halongensis has reflexed upper corolla lobes, a glabrous

inner surface of the corolla, short stamens (2—S mm long), which are

attached c. 3—4 mm above the base of the corolla, the filaments are

glabrous, and the fruits are 16—37 mm long. In life, these two species are

distinct in the colour of their leaves: mid-green and concolourous above

and whitish green beneath in C. hiepii, while in C. halongensis the leaves

are dark green above, frequently the midrib and veins are white on the

upper surface, and the lamina is frequently rosy purple beneath.

This species is named for Dr Nguyen Tien Hiep, senior botanist and

veteran plant collector of The Herbarium, Institute of Biology and

Biological Resources, Hanoi.

Specimens: Ha Long Bay - type and Nguyen Tien Hiep & Ruth Kiew NTH

4205 (HN).

194 Gard. Bull. Singapore 52 (2000)

6. C. modesta Kiew & T.H. Nguyen, sp. nov.

(Sect. Gibbosaccus C.B. Clarke)

Chiritae speluncae (Hand.-Mazz.) D. Wood similis, sed foliis majoribus,

inflorescentiis paniculatis et floribus albis differt.

Typus: Ha Long Bay, Nguyen Tien Hiep & Ruth Kiew NTH 4078 10 Nov

1999 (HN, holo, SING, iso).

Perennial herb, rhizome slender c. 5 mm thick embedded in rock fissures,

internodes congested. Indumentum of short, uniseriate, glandular and

eglandular trichomes, c. 0.2 mm long, dense on lower surface of lamina

and petiole, sparse on upper leaf surface with many unicellular trichomes

with a raised base. Leaves tufted, spirally arranged, petiole brownish, at

first short, elongating to (28—)35—45(—55) x 2—3 mm in the outer leaves,

terete, lamina oval, (16—)25—30(—38) x (7—)14—17(—21) mm, dark

green above, pale beneath, thick and fleshy, base decurrent, margin entire,

slightly recurved, apex slightly acute, midrib slightly impressed above, lateral

veins (1—)2(—3) pairs, in life completely plane above and beneath, in

dried state midrib and lateral veins slightly prominent beneath. /nflorescence

erect, 4—11 cm long, paniculate with third order branching and up to c. 16

flowers, indumentum of peduncle, bract, pedicel and calyx of dense,

glandular trichomes, c. 0.75 mm long, peduncle (3.5—)5.5(—10.5) cm long,

purplish brown, bracts linear, c. 1—1.5 mm long. Pedicel 9—15 mm long,

calyx divided to base, lobes narrowly linear, c. 1.5 mm long, corolla

campanulate, minutely pilose outside, glabrous inside, white, outside tube

slightly greenish at base with 3 faint purple lines behind the upper lobes, c.

6—10 x 4—6 mm long, lobes rounded, upper lobes recurved, 1.5 x 2 mm

long, lateral lobes 2.5 x 2 mm, and lower lobe 3 x 3 mm, spreading, stamens

and staminodes attached c. 2 mm from base of corolla, staminodes c. 1 mm

long, filaments geniculate, strongly curved in upper half, c. 2.5 mm long,

green, glabrous, anthers white, c. 2.5 x 0.5 mm, fused face to face, disc

annular, c. 0.3 mm deep, ovary green, densely pilose with short glandular

trichomes, c. 3—4 mm long, style 3.5 mm long, stigma spathulate, exerted

just beyond upper corolla lobes. Infructescence curling backwards on itself.

Fruit with persistent calyx, capsule narrowly cylindric, initially splitting

along upper side, subsequently splitting into 4 longitudinal parts, (S—)7—

12 x c. 1.5 mm, style sometimes persistent c. |_—4 mm long.

Distribution: Vietnam, Quang Ninh Province, Ha Long Bay, narrow endemic

found only at Tam Cung Cave.

New Plants from Ha Log Bay 195

Habitat: It grows in small crevices with the rhizome deeply embedded and

with only the small tuft of leaves above the rock surface. It appears to

grow only on the vertical cliff faces close to the cave mouth in light shade.

Notes: This is an atypical species of Chirita in its spathulate stigma and its

fruits that eventually split into four longitudinal parts. However, Wood

(1974) has already drawn attention to a few other Chirita species (e.g. C.

integra Barnett and C. lacei (W.W. Sm.) B.L. Burtt) that have a spathulate

stigma. Of all the Chirita species, this new species shares a remarkable

resemblance with C. speluncae (Hand.-Mazz.) D. Wood from north-east

Yunnan in its small size, rather few, small hairy petiolate leaves in a lax

tuft, small flowers and short fruits. C. modesta differs from C. speluncae in

its larger leaves, paniculate inflorescences, white flowers and spathulate

stigma. (C. speluncae has petioles 2—10 mm long, laminas 9—22 x 5—9

mm, and solitary flowers with violet-blue lobes and a shallowly two-lobed

stigma).

Chirita modesta occupies a very narrow niche (see Habitat above)

and this may explain why it is so rare as very few caves meet its exacting

requirements. The single locality of C. speluncae, as its name suggests, is

recorded as a cave, but details of its habitat are lacking.

Specimens: Ha Long Bay — type and Nguyen Tien Hiep & Ruth Kiew NTH

3905 11 July 1999 (HN, SING).

Paraboea halongensis Kiew & T.H. Nguyen, sp. nov.

(Gesneriaceae)

Paraboea rabilii Xu & B.L. Burtt similis, sed foliis oppositis et inflorescentiis

terminalibus differt.

Typus: Ha Long Bay, Nguyen Tien Hiep & Ruth Kiew NTH 4055 9 Nov

1999 (HN holo; BM, E, K, LE, MO, SING iso).

Thin, wiry, perennial shrublet, branching close to base, stems at first felty

becoming glabrous and pale grey with age, 13—18 cm x c. 2 mm, stems

weak becoming decumbent and rooting, nodes annular, lower internodes

1—1.5 cm long, upper congested. Jndumentum on upper surface of lamina

of glistening, translucent, long, uniseriate, glandular trichomes with dense

layer of short unicellular hairs; young stem, lower leaf surface, bracts,

rachis of inflorescence with a tangled felt of long, unicellular, eglandular

trichomes, ferrugineous at first becoming white with age, on lower surface

of veins with sparse long, dark brown, uniseriate trichomes that wear off

196 Gard. Bull. Singapore 52 (2000)

with age. Leaves opposite, lower pairs distant, upper clustered at top of

stem, petiole (6—)12—17 mm, lamina lanceolate to slightly ovate, 26—43 x

15—25 mm, in life deep green above, dark fawn or grey-fawn beneath,

base rounded to slightly cordate, sometimes slightly unequal, margin serrate,

outlined in white (ciliate), apex acute, lateral veins ascending 4—5 pairs,

impressed above, prominent beneath. /nflorescence terminal, paniculate

with fourth order branching, peduncle c. 0. 5 cm long, branches 6.5—10 cm

long, 2—5 spreading, rachis white, extremely fine, rachis to second order

branching c. 4 cm long; bracts foliaceous, 12 x 6 mm with stalk 4 mm long.

Pedicel c. 9 mm long, green, calyx divided to base, lobes narrowly linear, c.

2 x 1 mm, light green, corolla campanulate, 8 x 7—8 mm, completely white

inside and out, tube slightly saccate at base, mouth oblique, lobes +

isomorphic, broadly rounded, c. 2 x 4—5 mm, recurved, stamens strongly

curved to position anthers c. halfway along corolla tube, filaments pale

rosy purple, c. 3—4 mm long, anthers connivent, white, c. 1.5 mm long,

ovary white, c. 5 mm long, style straight c. 3 mm long, stigma positioned at

the corolla mouth, minute, globose and glistening. Capsule narrowly

cylindric, 12—20 x 1—1.5 mm, twisted when mature and splitting on both

upper and lower sides.

Distribution: Vietnam, Quang Ninh Province, Ha Long Bay, at Me Cung

Cave and one other island.

Habitat: It grows in soil-filled cracks in bare, exposed rocks near the summit

of limestone islands.

Notes: The habit - a thin, wiry plant with weak stems, branching near the

base - is rather unusual in Paraboea and resembles that of P. rabilii Xu &

B.L. Burtt from Thailand. In addition P. rabilli has similar small, lanceolate

leaves, slender inflorescence branches, and short, white flowers and short

fruits. However, it would not be mistaken for P. halongensis as P. rabilii

has alternate leaves with 8—10 pairs of veins, and axillary inflorescences.

The individual branches of P. halongensis are monocarpic. The leaves

and shoot die after fruiting and are replaced by a new shoot, which

contributes to the bushiness of the shrublet.

Impatiens halongensis Kiew & T.H. Nguyen, sp. nov.

(Balsaminaceae)

Nguyen & Kiew, Wild Plants of Ha Long Bay (2000) Plate 28.

Impatiens bonii Hook. f. maxime similis, sed foliorum nervis lateralibus

magis numerosis, floribus albis apice luteis et alis recurvis differt.

New Plants from Ha Log Bay 197

Typus: Ha Long Bay, Nguyen Tien Hiep and Ruth Kiew NTH 4056 11 Nov

1999 (holo HN, iso SING)

Plate 1b

Glabrous, perennial balsam. Stem succulent, erect, unbranched (sometimes

bifurcating), smooth and glossy, reddish-brown, to c. 40 cm tall, in life c.

12.5 mm thick at base, tapering to 5 mm at stem apex, leaf scars conspicuous.

Leaves spirally arranged, clustered at stem apex, petiole reddish brown,

(1.7—)3.5—4.5 cm long, grooved above, /amina broadly lanceolate, thin,

glossy, dark green above, pale green beneath with dark green veins, (3.2—)

7—10.5 x (1.9—)4.5—6.7 cm, base rounded slightly unequal with pair of

small, pale green glands at base of midrib, margin crenate, apex acuminate,

lateral veins 5—7 pairs, slightly impressed or plane above, slightly prominent

beneath. Flowers solitary, 2 per axil, flowering asynchronously, basically

white with edges of petal lobes yellow or greenish yellow, the lower centre

of throat with a bright yellow patch, inside the throat pale green with lines

of minute red spots sometimes coalescent, c. 14 x 12 mm; pedicel slender,

shorter than petiole, 2—2.5 cm long, pale brownish green; ebracteate; sepals

4, outer lateral sepals broadly ovate, c. 6 x 5 mm, united into an acuminate

tip, upper lateral sepals fleshy, acinaciform, c. 4 x 1 mm, lip naviculate, c.

21 mm long of which c. 7 mm is the spur, c. 8 mm across mouth and c. 7

mm deep, spur simple, recurved, inflated, c. 6 x 2.5 mm; standard obovate

with a Keel-like pouch, c. 9 x 6 mm with 2 rounded lobes c. 3 mm across,

pouch c. 3 mm wide, lateral petals connate, c. 8 x 13 mm; distal lobes c. 4 x

13 mm, strongly recurved, basal lobes rounded c. 7 x 6.5 mm, shallowly

notched, notch c. 2.5 mm deep, inner margin with raised bright yellow

central ridge; ovary c. 3 mm long, stigma sessile, globose, c. 2 mm across.

Capsules obovate, c. 12 x 4 mm, narrowing abruptly to an apicula c. 3 mm

long, glabrous, strongly ribbed, dark green, almost white between ribs.

Seeds discoid, flat, slightly obovoid, c. 2 x 1 mm, minutely rugose.

Distribution: Vietnam, Quang Ninh Province, endemic to Ha Long Bay.

Habitat: In light shade, on ground in soil derived from limestone rocks.

Notes: The common balsam at Ha Long Bay is /mpatiens verrucifer Hook.

f., which is conspicuous as it grows on vertical limestone cliffs and rocks

close to sea level. It is a massive balsam growing to | m in height with

tough succulent stems about 3 cm thick. It produces large, deep pink flowers,

though some populations have paler flowers. In contrast, the new species

is not as common and has only been found on a few islands where it grows

198 Gard. Bull. Singapore 52 (2000)

in light shade under the tree canopy.

Impatiens halongensis belongs to a group of Vietnamese balsams,

which includes /. boni Hook. f. and I. verrucifer Hook. f., that are glabrous,

have axillary flowers, four lateral sepals of which the outer ones are free, a

short spur and connate lateral petals. It would not be mistaken for J. boni

and J. verrucifer, which have large 3-cm wide, pink flowers with a deeply

divided lip and a pedicel that is longer than the petiole.

Some plants of J. boni (formerly described as distinct and called J.

pygmea Hook. f.) have basically white flowers and pedicels shorter than

the petioles, but /. halongensis is clearly distinct from them in the number

of lateral veins (3—5 pairs in I. boni), in flower shape (the distal lobes of

the lateral petals are not recurved in J. boni), and colour (the white flowers

of I. boni have violet apices).

Livistona halongensis T.H. Nguyen & Kiew, sp. nov.

(Palmae)

Nguyen & Kiew, Wild Plants of Ha Long Bay (2000) Plate 24.

Forma foliorum et magnitudine fructuum Livistonae endauensi J. Dransf.

& K.M. Wong similis, sed inflorescentiis erectis et multo longioribus

praecipue differt.

Typus: Ha Long Bay, Nguyen Tien Hiep, L. Averyanov & Nguyen Van

Can NTH 2630 12 May 1999 flowers (holo HN, iso SING)

Plate la

Solitary palm. Stem to c. 10 m tall and c. 20 cm diam., flowering at c. 5 m

tall, leaf scars conspicuous and close. Leaf with petiole longer than lamina.

Leaf base and leaf sheath clasping the stem, c. 15 cm wide at base, thickly

woody; ligule rich brown, triangular and fibrous, the proximal 35—38 cm

stretched to form a fibrous network, the distal c. 70 cm tattered into ribbons.

Petiole glabrous, 1.25—1.30 m long, c. 2 cm wide at base, c. 1.5 cm wide

distally, adaxially flat and green with light yellow band along the margin,

abaxially green and rounded, proximal 15 cm without spines, distally with

77—86 irregularly spaced spines per side; spines dull orange, paler green

towards centre, 10—12 mm long, woody, hooked, underside strongly keeled.

Adaxial hastula persistent, papery, broadly ovate, c. 3 x 2.5 cm (hastula

absent on the abaxial surface). Lamina unsplit at base, then splitting into

one-veined segments which ultimately split along the single vein to become

bifid, segments slightly pendant; near the margin, lamina c. 47 cm long and

splitting c. 6 cm from base, in the centre c. 77 cm long and splitting c. 23—

25 cm from base, veins c. 64; lower surface at base with pale fawn scurfy

New Plants from Ha Log Bay 199

indumentum wearing off distally; intercostal veins fine and slightly darker

green, just visible to the naked eye; margin at base with a c. 2 cm-long row

of very fine teeth. Old leaves forming a conspicuous skirt beneath the

crown, ultimately marcescent. /nflorescence solitary, interfoliar, green,

glabrous, c. 3.4 m long, more or less erect and projecting above crown;

prophyll and 10 upper bracts tubular and thick leathery-fibrous, prophyll

2-keeled, c. 33 x 5 cm at base tapering to 3 cm wide; bracts each c. 50 cm

long and terete, the upper 7 with partial inflorescences; peduncle c. 1.6 m x

2 cm with 5 bracts; partial inflorescences divaricate, softly pale fawn velvety,

with one bracteole c. 27 cm long, proximally stalk flat, terete distally, c. 95

cm long of which stalk is 60 cm long, arching outwards and downwards

with c. 10 second order rachillae, lower with third order rachillae, ultimate

rachillae c. 1—1.5 mm thick. Flowers minute in distant pairs on common

stalk c. 0.3 mm long, bud obovoid, flowers pale creamy yellow, c. 2 mm

long; calyx c. 1 mm long, glabrous, tubular dividing c. halfway, lobes 3

acute; corolla c. 2 mm long, divided c. halfway, lobes 3, cucullate; stamens

6, c. 1 mm long, staminal tube rich brown, narrowing abruptly into short

filament, anthers subglobose, glistening white, c. 0.3 mm long; ovary

obovoid, deeply ridged, dark reddish brown, c. 1 mm long; single united

style, pale brown, c. 0.3 mm long; stigma minute. Fruit stalk to (2—)3—S

mm long, fruit globose 10—12 mm across, smooth, glossy bright dark green,

epicarp and mesocarp leathery, endocarp crustaceous and brittle, c. 0.75

mm thick, single-seeded, endosperm homogeneous with an intrusion of

testa in the centre.

Distribution: Vietnam: Quang Ninh Province, endemic to Ha Long Bay.

Habitat: Limestone islands, in soil-filled crevices on rocky limestone

substrate, sometimes gregarious but it is not present on all islands.

Specimens: collections from same plant, all numbered NTH 2630 (12 May

1999 — type); Nguyen Tien Hiep & Ruth Kiew 15 July 1999 (fruits); 11

April 2000 (immature inflorescences).

Notes: This is the first Livistona species to be described from limestone. It

flowers in May, when it produces spectacular large, cream inflorescences

above the green leaf canopy. It fruits in July.

It is clearly different from the other Vietnamese species of Livistona

as indicated in the key.

200 Gard. Bull. Singapore 52 (2000)

Key to the Vietnamese Species of Livistona

la. Lamina irregularly divided into broad segments that almost reach the

lasinla .eo eeS L. saribus Merrill ex A. Cheval.

1b. Lamina unsplit at base, segments regularly divided ............. ee 2

2a. Tips of leaf segments conspicuously pendulous ...... L. chinensis R.Br.

2b. Tips of leaf segments not or only slightly pendulous ...........0000.... 3

3a. Inflorescences as long as leaves, fruits ellipsoid, 25—35 x 20— 25 mm

(adbis ele wenindecet RL Se Ss ee L. speciosa Kurz

3b. Inflorescences longer than the leaves, erect and held above the crown,

fruits globose; 10-12 am: wide 222255-...200,2e se L. halongensis

Livistona fengkaiensis X.W. Wei & M.Y. Xiao from southern China has a

short, pendulous inflorescence and larger, ellipsoid fruits and so would not

be mistaken for L. halongensis. John Dransfield (pers. comm.) suggests

that this Chinese species is in fact conspecific with L. speciosa.

Livistona halongensis, in its relatively small fruits and its leaves, which

do not split to the base and which have single-veined segments that bifurcate

near the margin, most resembles Livistona endauensis J. Dransf. & K.M.

Wong, which grows on sandstone ridges in southeast Peninsular Malaysia.

However, apart from the large erect inflorescences (which are more than

twice the size of those of L. endauensis, which has inflorescences to c. 1.5

m long), it differs in several other characters: the leaves have a shorter

petiole (they are c. 1.75 m long in L. endauensis), a conspicuous hastula (it

is only 1 cm long in L. endauensis) and the leaf is more deeply split (more

than a third of the way down, compared with only halfway in L. endauensis).

In addition, its fruit is globose and slightly smaller than that of L. endauensis,

which is pyriform and 16 by 14 mm in size.

Acknowledgements

The authors are indebted to the Ha Long Bay Authority for its total

support in providing boats and boatmen without which we would not have

been able to explore the islands, in particular to Mr Nguyen Van Tuan,

Head of the Management Department of Ha Long Bay, for his interest in

and support of our field work, to IUCN Vietnam for initiating and co-

ordinating the project, and especially to Dinh Thi Minh Thu of IUCN

Vietnam for her most efficient handling of all the logistics, to the Royal

Netherlands Embassy for financial support and to Wendy Gibbs, without

New Plants from Ha Log Bay 201

whom there would never have been the project, for her patience in being

greeted with yet another plant and being told “You must paint this!” and

to Leonid Averyanov of Komarov Botanical Institute for taking part in

the May 1999 field trip. We also thank the Curators of the Herbaria at E,

K and BM for permission to examine specimens in their care, John

Dransfield for discussions on Livistona and for providing us with relevant

literature, B.L. Burtt for sharing information on Indo-Chinese Gesneriaceae,

and M.J.E. Coode for correcting the botanical latin.

References

Dransfield, J. and K.M. Wong. 1987. A new species of Livistona (Palmae)

from Peninsular Malaysia. Malayan Nature Journal. 41: 11—123.

Nguyen, Q.B. and M. Newman. 2000. A new species of Alpinia

(Zingiberaceae) from Vietnam. Gardens’ Bulletin Singapore. 52: 211—

4 VA

Nguyen, T.H. and R. Kiew. 2000. Wild Plants of Ha Long Bay. 1UCN

Vietnam, Hanoi, Vietnam. I[llustrations by W. Gibbs.

Wood, D. 1974. A revision of Chirita (Gesneriaceae). Notes Royal Botanic

Garden Edinburgh. 23: 123—205.

202 Gard. Bull. Singapore 52 (2000)

Plate 1. Endemic plants of Ha Long Bay.

a. Livistona halongensis, b. Impatiens halongensis, c. Chirita drakei, d. Chirita gemella, e. Chirita

halongensis, f. Chirita hiepii.

Gardens’ Bulletin Singapore 52 (2000) 203-210.

Two interesting wild Musa species (Musaceae) from

Sabah, Malaysia.

G. ARGENT

Royal Botanic Garden, Edinburgh, EH3 5LR, U.K.

Abstract

Two new species of Musa are described and illustrated. M. suratii Argent is described from

Sabah with an additional locality in Sarawak. M. monticola Hotta ex Argent is known only

from the two montane localities in Sabah, Mt. Kinabalu and the Sinsuron road in the

Crocker Range.

Introduction

There is still a lot to be learnt about the relatively small genus Musa as the

wild plants are very often neglected by botanists in the field and herbarium

specimens will always be poor material to work from. Our knowledge of

the Bornean species goes back to the classic work of Odoardo Beccari

(1902) based on his field work from 1865 but the species of this genus had

been all but neglected on the island until Mitsuro Hotta made a series of

expeditions and described several additional species. One of the new species

described here was recognised by Hotta and given the name, which has

been adopted here, but he never described it.

Terminology is inevitably somewhat specialised in a group of large

and unwieldy plants that have enormous economic importance. The

descriptive terms here follow the tradition of banana taxonomy as used by

Simmonds (1962, 1966) and as modified in Argent (1976), which 1S

particularly important in standardising leaf measurements to the 4" last to

emerge before the inflorescence.

Musa suratii Argent sp. nov.

Ab omnibus aliis speciebus generis Musae pseudocaule tenuissimo usque

ad 4 cm diametro, gemma mascula tenuissima usque ad 3 cm lata et semine

in genere minimo cognito 2.5 mm tantum diametro differt.

Typus: Sabah: Kallang, Tenom District Andy Surat & A. Lamb 268/89, 8

Sept. 1989 (holo SAN; iso E).

204 Gard. Bull. Singapore 52 (2000)

Fig. 1.

Clump forming herbaceous plant to c. 4m high. Suckers emerging from

below ground level on the corm, initially near vertically, up to 10 cm from

the parent; suckers slightly waxy, vertical in young clumps but becoming

angled outwards in the larger older clumps. Mature pseudostem up to 3.5

m high, slender to 4 cm in diameter at 30 cm above the ground, tapering

gradually upwards and spreading at between 30-45° from the vertical; dark

purplish black and somewhat shiny in the basal half, dull yellowish green

in the upper half, swollen at junction with the corm which is c.10 cm in

diameter. Inner sheath cream tinged with purple, undersheath yellow,

shiny. Sap watery. Upper margin of sheath (shoulder) smooth, appressed,

not scarious. Fourth last leaf: right handed to 3 cm; petiole 16—20 cm x 8

mm, dull yellowish with a pinkish edge to the margins, petiole channel

closed in the proximal half, open distally, the channel small, about one

quarter of the depth of the petiole; blade 120—150 x c.20 cm; more or less

narrowly elliptic, broadest about or slightly above the middle, broadly

pointed but often tattered at the apex, cuneate at the base, very white

waxy underneath, dull green not waxy above.

Inflorescence hanging vertically downwards, peduncle slightly waxy,

glabrous, smooth. Female bud, 40 x 4 cm, orange to slightly pink, pruinose

with wax outside, imbricate for nearly half the length. Bracts lifting to

c.45°, straight, not curling back but somewhat channelled, bright shining

orange underneath. Basal flowers mostly fully hermaphrodite, c. 9 cm

long, the ovary cream, the compound tepal green, pale proximally, darker

distally, the free tepal unwrinkled, or wrinkled irregularly, more or less

truncate but with three short points. Stamens 5, variously ranging in size

from fully developed with apparently fertile pollen to tiny staminodes.

Style green, as long as the flower, stigma globose, cream. Fruit bunch

hanging vertically down, fairly lax, the fruits reflexed upwards, 8—10 ‘hands’

with the bananas in two rows with 8—10 fruits in the second ‘hand’. Fruits

pale whitish-green, ripening pale yellow, not splitting open, ovules in two

rows per loculus. Pedicel c.10 x 5 mm, fruit 6—9 x 0.8—1.0 cm, straight or

curved, almost smoothly cylindrical hardly angled, contracted at the apex

to a darker green ‘beak’ for 5—6 mm. Seeds dark greyish-black, slightly

elongated, small c.2.5 x 2 mm, almost smooth but with irregular very low

tubercles.

Male axis vertical, glabrous, pruinose, pinkish-purple, 30—140 cm

long, the scars somewhat prominent, the peduncle almost smooth and only

very weakly vertically striate, the bract scars well-spaced 1.5—2.5 cm apart

in the same rank. Male bud slender, from 15—22 x 1.8—3 cm, broadest at

about one quarter of its length from the base, imbricate for between one

Two wild bananas from Sabah 205

Figure 1. Musa suratii: A. inflorescence with submature fruit bunch, B. male bud, C. male

flower. Male flower parts: D. compound tepal, E. free tepal, F. stamen, G. style and stigma.

Hermaphrodite flower parts: H. stamen. I. compound tepal, J. style and stigma, K. flower (free

tepal missing). L. TS. of petiole, M. single fruit, N. TS. fruit. Seed: P. in LS, Q. entire.

206 Gard. Bull. Singapore 52 (2000)

third to one half of its length, bright orange except for the tips of the

bracts which are green. Male bracts lifting to 45°, straight, not curling

back but the margins slightly inrolled longitudinally; slightly pruinose-waxy,

orange with green tips outside, bright, somewhat shiny not waxy orange

inside. Male flowers 30—45 mm long, creamy at the base, green in the

distal half, compound tepal with the three major lobes up to 4 mm acutely

pointed and irregularly fimbriate with long hairs to c.1 mm long, the two

minor lobes free for c.1 mm and minutely mucronate, free tepal c.4 mm

shorter than the compound tepal, translucent white, irregularly three-

pointed the lateral points broad the central point smaller, firmer, greenish-

yellow. Stamens 5, creamy white, the anther over half the length of the

stamen. Style slender, the length of the flower, broadened slightly towards

the stigma.

Additional specimen: Sarawak: River Delok, Lubok Antu District

Christensen 1066 (AAU, E!, SAR).

Notes: Named in honour of Aninguh (Andy) Surat, plant collector for the

Orchid Centre, the Agricultural Park, Tenom, Sabah, who first found this

species.

This delicate species looks from a distance more like a Heliconia

than a Musa as, in addition to the very slender habit, the leaves although

actually spirally arranged are displayed more or less distichously in a single

plane. The original population was along a logging road that had opened

up the forest canopy of mixed lower montane forest with some

Dipterocarpaceae at about 750 m altitude. This population, the only one

known in Sabah, is rapidly shrinking as the forest re-grows and the canopy

is closing. A second population was reported from Sarawak by Dr.

Christensen (pers. comm.) and confirmed from herbarium material. She

reported it to be common and used as a food plant by the local people. It

is speculated that, in the absence of any further reports from Sabah, the

population may have been introduced by logging equipment, used

elsewhere, with the minute seed brought in on clinging soil. It is now

being cultivated in the Tenom Agricultural Park. The tiny seeds, by far

the smallest in the genus, give no clue as to its identity at sectional level.

Musa monticola [Hotta ex] Argent sp. nov.

Musae textili similis sed pseudocaulibus multo minoribus 1—2 m tantum

altis, pedunculo masculo brevissimo usque ad 12 cm longo et gemma

mascula anguste acuta cum imbricatione 1—2 cm longo apicali differt.

Two wild bananas from Sabah 207

Typus: Sabah: Kinabalu Park Headquarters, Kinabalu, Ranau District

Argent 2195, 20 Sept.1995 (holo SAN; iso Kinabalu Park Herbarium, E).

Fig. 2.

Pseudostem 1—2 m, dark shiny brown or mottled green and brown often

with a little violet, waxless, sometimes almost completely covered with

pale brown withered sheaths, under-sheath pale green, inner sheath white,

15 cm girth at the base. Sap watery or slightly creamy. Rhizomes short,

the suckers erect arising up to 10cm away from the parent pseudostem.

Suckering frequent, the plants mostly forming small clumps. Upper leaf

sheath margin (shoulder), appressed, smooth with a dark brown edge c.2

mm wide, sometimes narrowly scarious for 1—1.5 mm. Fourth last leaf:

petiole 36—48 x 1.4—1.6 cm, mostly dark rather lustrous brown, sometimes

flushed with pink, without wax, the channel c. one third of the depth, (TS

ratio 1/2), margins erect distally, completely enfolded in the lowest one

third to one half and edged with a black or scarious margin c 1mm wide;

leaf blade left-handed to c.1—1.5 cm, sometimes almost symmetrical,

rounded to auriculate at the base, rarely broadly tapering, dull green above,

yellowish green below, often with brown discoloured areas, sometimes

with light pink on the midrib above and below, without wax, PB ratio

3.5—4, the blades being 104—160 x 32—45 cm broadest about the middle.

Inflorescence with glabrous, green peduncle. Bunch held semi-erect,

very dense, the fruits two-rowed, in 5—7 ‘hands’, the second ‘hand’ with

10—13 fruits from very short c. 5 mm pedicels. Fruits more or less

ageotropic disposed almost radially from the axis. Basal bracts, long,

lingulate, c.23 x 9 cm, shiny, glossy purplish-brown outside, sometimes

with a little pink; passing to yellow for 1—2 cm at the base and with green

margins at the tip, inside creamy yellowish with a pinkish purple edge, sub-

persistent to persistent in a withered brown state, trapped between the

tightly appressed fruit as they develop. Basal flowers hermaphrodite with

a full compliment of fully developed stamens, ovary c.3 cm, compound

tepal c. 3.3 cm, cream with yellowish tips; the free tepal white, strongly

keeled and wrinkled; stamens cream; stigma white; style cream. Young

fruits glabrous, green, 40—75 x 16—25 mm; ovules in two rows per loculus

often strongly angled in cross section due to the compression between

fruits. Mature fruit ripening dirty cream in the basal part, dirty creamy-

green in the upper part, (clear pale yellow in cultivation), indehiscent, not

strongly aromatic. Seeds small, 4—5 mm diameter, sub-spherical, only

weakly angled, with a distinctly warty surface, the hilum small c.1.5 mm,

smooth, apical chamber small.

Male peduncle growing diagonally downwards, weak, 5—12 cm long,

glabrous, often terminating whilst the fruits are still very immature,

208 Gard. Bull. Singapore 52 (2000)

——

Figure 2. Musa monticola: A. inflorescence with submature fruit bunch, B. male bud, C. TS.

petiole. Hermaphrodite basal flower: D, flower, E. compound tepal, F. free tepal, G. style and

stigma, H. stamen. I. TS. Fruit, J. fruit. Seed: K. LS, L. entire. Male flower: M. whole flower, N.

compound tepal, O. free tepal, P. stamen, Q. style and stigma.

Two wild bananas from Sabah 209

frequently broken and appearing to be absent. Male bud conical with a

very acute tip, 6—9 x 2.2—4 mm, imbricate for just 1—2 cm at the tip,

creamy orange with a dark purple flush and blackish purple at the bract

margins; bracts lifting rather irregularly to the perpendicular or slightly

higher, often several opening together, intensely shiny, orange brown on

the adaxial surface, not reflexing but the margins tending to inroll slightly

with age, persistent or tardily deciduous. Male flowers up to 3 cm, cream,

the tips of the compound tepal yellowish green, strongly reflexed and

rolled, to 5 mm long when unrolled. Free tepal translucent white with a

yellow spot at the apex, nearly as long as the compound tepal, concave and

acutely pointed, sometimes with a small subapical wrinkle.

Additional specimens: Sabah: Mt. Kinabalu, Ranau District - Kiau View

Trail Cockburn SAN 76808 vernacular name ‘Pahuo’ in Dusun Bunda

Tuan (SAN!), Bembangan Camp Chew & Corner RSNB no.4576 27

Feb.1964 (K!, SAN!), Cultivated specimen at RBGE; Crocker Range,

Sinsuron Road, Argent 989 (E!), 19891874 (E!).

Notes: This interesting species was first noted as an annotation by Professor

Hotta to a specimen in the Sandakan Herbarium. It is mentioned in his

paper on the distribution of the genus Musa in Malesia but without any

description or diagnosis of the plant (Hotta, 1987). I have used his name,

which clearly relates to the somewhat unusual montane habitat of this

species. It is common in the garden area around the Park Headquarters

on Mt. Kinabalu where it occurs both in the gardens and in open areas of

the forest from about 1200—1700 m altitude. It also occurs on the Crocker

Range and can be seen from the Sinsuron Road at similar altitudes in the

secondary roadside vegetation. At present, these are the only two known

locations for this species, both of which were known to Hotta.

Its sectional position within Musa is unclear. Its seed shape and

anatomy rule out its inclusion in section Callimusa and the male axis habit

rules out section Rhodochlamys. Without a chromosome count it is difficult

to assess whether it should be assigned to section Australimusa or section

Musa. On the whole, the polished bracts and lack of wax favour its inclusion

in section Australimusa as Hotta inferred (Hotta, 1987).

The fairly extreme montane environment of this species would tend

to lend weight to the observation of Simmonds (1962) that Musa conforms

to ‘Bakers Law’ - the theory that extreme environments encourage the

development of hermaphrodite basal flowers instead of female, making

self pollination the likely norm. In this respect M. suratii from lower altitude,

with its partially hermaphrodite flowers appears to be a halfway house.

Musa monticola is a very distinct species. The short pseudostems,

210 Gard. Bull. Singapore 52 (2000)

rarely more than 2 m high, contrast with those of Musa textilis Née, which

in Sabah are almost invariably over 3 m. The very short male peduncle

only up to 12 cm, again contrasts with the very vigorous male phase in M.

textilis, whose male peduncle usually grows to 1 m and often to more than

2m. The very acutely pointed male bud with only 1—2 cm of imbrication

at the tip differs significantly from the much more broadly pointed and

strongly imbricated male bud of M. tfextilis, whose area of imbrication is

usually more than 3 cm. The hermaphrodite basal flowers and subspherical

warty seeds are other distinctive differences.

Acknowledgements

The fieldwork involved in the description of the above species was

sponsored by the Royal Botanic Garden, Edinburgh, and supported in

Sabah by Tony Lamb, the Agricultural Park, Tenom, and the Kinabalu

Park. I would also thank the Curator of SAN for facilities to study

specimens. Maureen Warwick ably executed the illustrations, and I am

indebted to Dr Robert Mill for the Latin diagnoses. Mr B.L. Burtt, as

always, gave sound and much valued advice.

References

Argent, G.C.G. 1976. The Wild Bananas of Papua New Guinea. Notes

Royal Botanic Garden Edinburgh. 35: 77—114.

Beccari, O. 1902. Nelle Foreste di Borneo. Firenze, Italy.

Hotta, M. 1987. Distribution of the genus Musa in Malesia. Acta Phytotaxa

Geobotanica. 38: 292-.302.

Simmonds, N.W. 1962. The Evolution of the Bananas. Longmans, U.K..

Gardens’ Bulletin Singapore 52 (2000) 211-212.

A New Species of Alpinia (Zingiberaceae) from Vietnam

NGUYEN QUOC BINH

Institute of Ecology and Biological Resources NCST,

Nghia Do, Cau Giay, Hanoi, Vietnam

AND

MARK NEWMAN

Royal Botanic Garden, Edinburgh EH3 5LR, U.K.

Abstract

Alpinia calcicola Q.B. Nguyen & M.F. Newman, a new species of subsect. Alpinia from Ha

Long Bay,Vietnam, is described.

Introduction

While working on a guide to plants of Ha Long Bay (N.T. Nguyen &

Kiew, 2000), a species of Al/pinia was collected that does not match any of

those cited in Q.B.Nguyen (1994). Further study indicates that it is a new

species, which is described here:

Alpinia calcicola Q.B.Nguyen & M.F. Newman sp. nov. Alpiniam maclurei

Merr. tangit, ob laminam minorem, (22—31 x 3—7 cm), glabram, bracteolam

2 vel 3 flores gerentem, tubum calycis tubum corollae aequantem, antheram

ecristatam inter species subsectionis Alpiniae ponenda.

Typus: Vietnam, Quang Ninh Province, Ha Long Bay, Nguyen Tien Hiep

& Ruth Kiew, NTH 4220 12 April 2000, flowering (holo HN; iso SING).

Herb 1.5—2 m. Leaf sheath glabrous, striate; ligule 8—10 mm, emarginate,

shortly hairy, ciliate at margin; petiole 2—7 mm, reddish brown, glabrous;

lamina lanceolate to linear-lanceolate, 22—31 x 3—7 cm, light green above,

yellowish green beneath with reddish brown midrib, glabrous on both

surfaces; margin with sparse, tooth-like hairs; base acute; apex acuminate.

Inflorescence terminal, yellowish brown, 26—32 cm long; rachis slender,

2—3 mm diameter, sparsely hairy; cincinni diffuse, becoming denser towards

apex, axes hairy; bract subtending inflorescence pale pinkish brown, to 18

x 2 cm, caducous; cincinni without bracts, usually with a very short flap of

free tissue at the join with the rachis; bracteoles broadly ovate, split to the

a12 Gard. Bull. Singapore 52 (2000)

base, pinkish white to pink, hairy at tip and margin, usually subtending 2—

3 flowers, first bracteole 12—14 x 8—10 mm, subsequent ones smaller;

ovary globose, hirsute; epigynous glands c. 3 mm long, fused into a truncated

cone; calyx tubular, 12—14 mm, adpressed silky hairy outside, split into

three short ciliate teeth above; corolla tube as long as calyx, very sparsely

hairy outside, dorsal corolla lobe 14—19 x 4—6 mm, oblong, slightly hooded,

lateral corolla lobes smaller, 14—16 x 5—7 mm; labellum c. 14 x 18 mm,

broader than long, shortly bifid at apex, margins frilled, lateral lobes pure

crimson with white margin, centre with longitudinal crimson lines; /ateral

staminodes 4—5 mm, fleshy; filament 13—14 mm, anther 4—S mm long,

sparsely short hairy on back, thecae parallel, connective sometimes produced

as a small flap of tissue; style filiform, stigma funnel-form, ciliate; fruit

globose, 6—7 mm.

Other specimens seen: Vietnam, Quang Ninh Province, Ha Long Bay:

Nguyen Tien Hiep & Leonid Averyanov NTH2600 12 May 1999 (HN, LE,

SING); Nguyen Tien Hiep & Ruth Kiew NTH3931 15 July 1999 (HN,

SING).

Habitat: In rock cracks and crevices in open places from sea level almost to

the summits of limestone islands.

Notes: This species is illustrated in colour by Wendy Gibbs in Plate 17 of

Wild Plants of Ha Long Bay (T.N. Nguyen and Kiew, 2000).

Acknowledgements

We should like to thank Nguyen Tien Hiep (HN), Ruth Kiew (SING) and

Leonid Averyanov (LE) for bringing this new species to our attention and

providing us with material. This species was discovered during the fieldwork

for the guide to Wild Plants of Ha Long Bay, a project initiated and co-

ordinated by IUCN Vietnam.

References

Nguyen, Q.B. 1994. The genera Alpinia Roxb. and Kaempferia L.

(Zingiberaceae) in the flora of Vietnam. Journal of Biology (Viet Nam)

16(4): 143-145.

Neuyen, T. H. & R. Kiew. 2000. Wild Plants of Ha Long Bay. IUCN

Vietnam, Hanoi, Vietnam. Illustrated by W. Gibbs.

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VOL. 52 (Part 2) December 2000 ISSN 0374-7859

CONTENTS

Boyce, P.C.

The genus Rhaphidophora Hassk. (Araceae-Monsteroideae-

Denials 00) VS NTP NO Soe, Bob oy en edeiasers onthe hekesexitvavesUuyriccbicntoscasoappenctiaaconns svusvdveacsd 213

Zhu, H.

A new species of Lasianthus (Rubiaceae) from Peninsular Malaysia ................:cceeeee ZW

Salma, I.

The significance of pollen morphology in the taxonomy of the genus

MOB es 5 EA OND A IG Yel icc nace mu aeeet at soo ae ets niea ce ctragna sa dev etek des deescavasenvestnencntideUcvnevancers 261

Vermeulen, J.J.

Carr’s spirit collection of Kinabalu orchids recovered, with some

notes oa Bulbophyllum species present im Chis COMSCHION. 2.02... .0i.cicsccsecscnsscessncasenccceconss pH)

O’Byrne, P. and J.J. Vermeulen

Notheria diaphana gen. nov., sp. nov. (Orchidaceae), a gem from

BUS Ae MUN ER et RM ode Poe a hoe cas eg aaciedie snr ck ev enetha Wiss eab dhe uerdvstadswactacees seeenbaven veal 285

Vermeulen, J.J. and L.G. Saw

Bulbophyllum sect. Hirtula (Orchidaceae): a new species and a new

TURN CO re RUMEN ULSD IVA ANAS Sista date esos antserooece se sade Nan entuctnbuptiiacsveeeilaxaonsaonesvedsensiiasosbs 289

Vermeulen, J.J. and P. O’Byrne

Bulbophyllum jiewhoei (Orchidaceae), a new species of unknown

Ba aR ee re a eee intake Bays Bete eae use sta as Wyn RE $e ny dp dean cds Gaede ToOANlow nan cvanadesrdevsUouabubeds 205

BOOK REVIEWS

The Plants of the Singapore Botanic Gardens: an Annotated Check-list ................ R. Kiew - 297

Tree Rlora.of Sabah. amd Sapawalkk \V OWI Biiiiecsccses ccecondcnarsvnnedsedepnstesnessandenerscacsannseses R. Kiew - 299

Bryophyte La 6 (allan "a stintene. iRaA EN Cale bei Reece AS cet aOR Ah DREN nn AUP A B.C. Tan - 301

Date of Publication: 29 December 2000

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 52 (2000) 213-256.

The Genus Rhaphidophora Hassk. (Araceae-

Monsteroideae-Monstereae) in the Philippines

P.C. Boyce

Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, U.K.

Abstract

An alpha-taxonomic account of Rhaphidophora Hassk. in the Philippine Islands is presented

as a precursor to the forthcoming Flora Malesiana Araceae treatment. Eleven species are

recognized, of which Rhaphidophora cretosa is newly described and R. banosensis is proposed

as a nomen novum for R. stenophylla Elmer non K. Krause. R. bulusanensis is reduced to

the synonymy of R. monticola, R. rigida is merged with R. philippinensis, and R. lagunensis

is synonymized with R. acuminata. A dichotomous key to species is provided. All species

are illustrated.

Contents

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Key to adult flowering Rhaphidophora in the Philippine Islands ......... 218

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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 represented in tropical

Asia and has several nodes of diversity; the Himalayas (SE Nepal to NE

Vietnam, roughly 17°—23° N), West Malesia (including southernmost

peninsular Thailand), the Philippines, and East Malesia.

The last complete revision of Rhaphidophora was that of Engler &

214 Gard. Bull. Singapore 52 (2000)

Krause (1908). A summary of the taxonomic and nomenclatural history of

Rhaphidophora was presented in Boyce (1999).

This is the fourth in a series of papers intended to present a complete

alpha-taxonomy of the genus Rhaphidophora in Asia. Accounts for

Peninsular Malaysia and Singapore (Boyce, 1999) and the southern and

western Indonesian archipelago have been published recently (Boyce, 2000),

togther with an account of neotenic species in New Guinea (Boyce &

Bogner, 2000). Accounts for each of Borneo, Papuasia, the Himalayas,

Thailand, and Indochina are being prepared and will be published

separately. All morphological terms employed follow Stearn (1992).

Endemism and Relationships

Of the 11 Philippine species recognized, just four are recorded from outside

the region. One of these is the widespread Rhaphidophora korthalsii Hassk..

Of the other three non-endemics, R. minor Hook.f. and R. conocephala

Alderw. are recorded in the Philippines only on Palawan, an island that

while politically part of the Philippines has a flora with numerous Bornean

elements, while R. perkinsiae Engl. has been found in Taiwan (Lanyu

Islands) and in the Ryukyus. Lanyu is notable for having Philippine floristic

elements.

Most of the seven endemic Philippine species, plus non-endemic R.

perkinsiae, seem to be closely related to one another and considerably

more distant to species in neighbouring countries. Certain species (e.g. R.

acuminata Merr., R. perkinsiae and R. philippinensis Engl. & K. Krause, R.

elmeri Engl. & K. Krause and R. banosensis P.C. Boyce) are so similar that

it seems probable that these are of recent evolution. The paucity of

collections and inadequacy of the specimens exacerbate the problem of

separating these similar taxa and it is not yet clear whether more exhaustive

fieldwork will reduce the number of recognized Philippine species or support

recognizing a considerable number of further novelties.

A few Philippine endemics seem to be morphologically similar: to

species from neighbouring countries rather than to other Philippine species.

Of these R. cretosa belongs to the Hongkongensis Group (see Boyce, 2000)

and is morphologically most similar to the widespread Rhaphidophora

sylvestris (Blume) Engl. and may represent a link to the Bornean flora. R.

todayensis is unique in the Philippines by the pubescent abaxial surface of

the leaf lamina and on the basis of this character is probably allied to R.

foraminifera (Engl.) Engl. (Sumatera, Peninsular Malaysia and throughout

Borneo), R. puberula Engl. (Peninsular Malaysia, Sumatera, Nusa Tenggara

and throughout Borneo) and R. hookeri Schott (N. India to SW China),

Rhaphidophora in the Philippines 215.

which form the Hookeri Group (see Boyce, 2000). R. monticola is an

enigma. In the fertile condition it is morphologically distinct from any

other species of Rhaphidophora, while the leaf, especially in the lamina

venation, resembles R. philippinensis. The extraordinarily long peduncle

and the uniform pale yellow-brown colour of dried material renders it

instantly recognizable and distinct.

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. Fl. Malay Penins. 3 (1907) 39—46; Engl. & K. Krause in Engl.,

Pflanzenr. 37 (IV.23B) (1908) 17—S53; Engl. & Prantl, Nat. Pflanzenfam.

Nachtr. 3 (1908) 29; Koorders, Exkursfl. Java, vol. 1 (1911) 253—255;

Merr., 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 FI. Java, vol. 17 (1957) 13

15; Backer & Bakh-f., Fl. 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 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.) Miq., 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.|

216 Gard. Bull. Singapore 52 (2000)

[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

(not in the Philippines), root-climbing lianes, very seldom clustering and

rheophytic (not in the Philippines) and then always with a creeping juvenile

stage; 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 skototropic (see Strong

& Ray, 1975) by an alternating series of congested leafy and elongated

leafless shoots (not in the Philippines) or germinating to give rise to a non-

skototropic shingling juvenile shoot (e.g., R. korthalsii); 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: 1. physiognomically monopodial clinging non-flowering stems

rooting along their entire length giving rise to variously elaborated free

sympodial lateral flowering stems (e.g., R. minor), or u. all stems

physiognomically monopodial clinging and flowering (e.g., R. korthalsii),

or iii. physiognomically monopodial and sympodial lateral stems clinging

but only sympodial lateral stems flowering (e.g., R. acuminata); stems with

internodes of various lengths separated by variously prominent leaf scars,

smooth or asperous or densely pubescent to ramentose (the last not in the

review area), older stems subwoody or somewhat corky or with distinctive

matt to sublustrous pale brown papery epidermis, with or without variously

textured prophyll, 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; leaves distichous or weakly spiralled, evenly distributed

or scattered or clustered distally; cataphylls and prophylls subcoriaceous to

membranous, 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

Rhaphidophora in the Philippines OTT.

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

membranous 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; /Jamina submembranous to stiffly chartaceous or coriaceous,

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; midrib usually

prominent raised abaxially and prominently sunken, sometime flush, rarely

slightly raised adaxially; primary venation + pinnate; interprimaries mostly

present, subparallel to primaries and sometimes indistinguishable from

them (e.g., R. monticola) 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)

to reticulate (e.g., R. korthalsii), variously prominent, often very difficult

to distinguish from primary venation (e.g., R. monticola); 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 swiftly disintegrating cataphyll, subsequent inflorescences

usually each subtended by a prophyll and cataphyll, more rarely by a

prophyll and partially to almost fully formed foliage leaf (but not in West

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 and then rotting (i.e., R. elmeri) rarely persistent through to

_ fruit maturity (1.e., R. monticola ), stiff to rather soft- or stoutly coriaceous,

dirty-white, greenish, cream or yellow; spadix subglobose 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

218 Gard. Bull. Singapore 52 (2000)

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 a appendix; ovules few to many, anatropous,

funicle long, placentae parietal to basal, sometimes + subaxile, 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, orientation circumferential

or longitudinal; stamens 4—6; filaments strap-shaped; anthers usually

prominently exserted from between ovaries 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,

subreticulate, rugulate, fossulate, scabrate, retiscabrate, verrucate, or psilate;

chromosomes 2n = 60, 120 (42, 54, 56).

Distribution: About 100 species from tropical Africa, South and South

East Asia, Australia and the Pacific with extensions into the subtropical

Himalayas, southern China and the southernmost islands of Japan.

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.

Etymology: Greek rhaphis, rhaphidos (needle) and pher6é (1 bear); refers

to the macroscopic (to lcm long), needle-like unicellular trichosclereids

present in tissues.

Key to Adult Flowering Rhaphidophora in the Philippine Islands

la. | Leaf lamina variously pinnately divided and perforated. Stem apices

with copious netted fibres. Juvenile plant shingling ..........0..

1b. Leaf lamina entire. Stem apices without netted fibres. Juvenile plant

PRONE SEMA NII ai iin os n ina ap esnah vest Vannes green’ date aieectie aaa ee 2

Rhaphidophora in the Philippines 219.

2a:

2b.

Da.

3b.

4a.

Ab.

pa.

Sb.

6a.

6b.

Va,

Fy at

8a.

Sb.

9a.

10a.

Abaxial surface of lamina and apical geniculum minutely pubescent

SSR CEM Cc La Ee cl Bre ce OES EP 8 11. R. todayensis

Pamina anc mpicalivemiculiniglabrous ...16. bie. desec icles ese &

Beteae AeA MNIN cl) EMI RBIS aia cases carseat ntidend yp Shadsavtneweeadetienseasnene +

Leal lamina (tainly COMACEOUS Of ChALtACEOUS ....15..05...2.dsscceesesecenee. i

Leaf lamina falcate to falcate-oblong, stiffly coriaceous. (Palawan)

Sh eRe ent et parse Mate rode dnabtind valbqo ca eeeas waa teten pubes cya htocee te 3. R. conocephala

Leaf lamina oblong to ovate-oblong or ovate, never falcate, variously

LLG ICEL 1 0 aah. ieee pk alibi at er ge pr 4 leet 20 co ie AA eA 5

BP STATIS RIMES, FO 05) co cl cccantnendosvatersnaetvvenes 10. R. philippinensis

2 ENR 6 We) 0) or oh Re: eae ee ee ee ee 6

Petiole 5—11 cm; leaf lamina oblong to oblong-lanceolate, up to 23

x 8 cm. Entire plant drying dark brown. Peduncle 7—26 x 0.3—0.6

cm; spathe falling (at female receptivity 7?).............. 1. R. acuminata

Petiole 30—108 cm; leaf lamina ovate to ovate-oblong, up to 80 x 41

cm. Entire plant drying pale yellow-brown. Peduncle 30—117 x 1.5—

2 cm. Spathe persistent though to maturation of fruit...

sprig tahoe perimeter Mile. c3 tele lat hs telat i eo caine Meer 8. R. monticola

By OMI NCI oce MIN Re Soe ee FUMER SS See edenateaeecerstactaneectecnes 8

LORVOR ION SIMONI RC Cees mn Teme e cre saith re esas laa seigiatcasveeudastendeces 9

Primary lateral veins on the abaxial leaf surface significantly more

visible than interprimaries and never less than 5 mm apart. Spathe

persistent during fruit development, eventually rotting away. Spadix

Se hag a eu aaah Ua deat ah gees ee aan can Ae 5. R. elmeri

Primary lateral veins on the abaxial leaf surface + indistinguishable

from interprimaries, hence with the appearance of being less than 2

mm apart. Spadix 10—12 cm long. Spathe shed during anthesis .....

Poe Po AE On ek ips 5s ean Rela a Sees A eee abe 9. R. perkinsiae

Spadix more than 10 cm long (10—13 x 1—2cm). Leaf lamina falcate-

lanceolate to falcate oblong, up to 32.5 x 7 cm........ 2. R. banosensis

Spadix no more than 7 cm long. Leaf lamina ovate to falcate-elliptic,

Deere nt? MMO Me AN ee Ree ete, cree Ae teehee isc sactiasscctneleunaseoustcheaste 10

Leaf lamina non-falcate, 5.5—16 x 2—5 cm, thinly coriaceous with

veins not particularly prominent. Leaves drying strongly discolorous,

220 Gard. Bull. Singapore 52 (2000)

dark above; pale below, 2... eee 4. R. cretosa

10b. Leaf lamina falcate, 2.5—16 x 1.2—3 cm, thinly coriaceous to

chartaceous with veins prominent. Leaves drying pale straw-coloured

ee SARIS Gate iA ek ee 7. R. minor

The Species

1. Rhaphidophora acuminata Merr.

Rhaphidophora acuminata Merr., Philipp. J. Sci. 10 (1915) 265, Enum.

Philipp. Flower. Pl., 1 (1923) 175 — Type: Philippines, Leyte, 10 Sept.

1914, C.A. Wenzel 1139 (PNH7, holo; BM, MO, US, iso).

Rhaphidophora lagunensis Elmer, Leafl. Philipp. Bot. 8 (1919) 3072, Enum.

Philipp. Flower. PI., 1 (1923) 175, synon. nov. — Type: Philippines, Luzon,

Laguna, Los Banos (Mt Maquiling), June-July 1917, A.D.E. Elmer 17812

(PNH7, holo; BM, FI, K, L, MO, P, US, iso).

Figure 1

Moderately robust, semi-pachycaul homeophyllous liane to 4 m; seedling

stage and pre-adult plants not observed; adult shoot architecture comprised

of elongated, clinging, physiognomically monopodial, leafy, non-flowering

stems and shorter, clinging, sympodial, densely leafy, flowering stems; stems

smooth, terete in cross-section, with briefly persistent cataphyll remains at

the apices of active shoots, internodes to 6 x 1 cm on adherent shoots,

usually shorter and less stout on free shoots, drying very dark brown,

separated by large oblique leaf scars; flagellate foraging stems not observed;

clasping roots densely arising from the nodes and internodes of stems,

pubescent; feeding roots not observed; leaves distichous; cataphylls and

prophylls large and conspicuous, membranous, very quickly drying and

falling; petiole deeply channelled adaxially, 5—11 x 0.2—0.3 cm, smooth,

apical and basal genicula weakly defined; petiolar sheath very prominent,

extending to overtopping the apical geniculum, ligulate, clasping the newly

emerged leaf or inflorescence, swiftly drying and soon falling whole to

leave a conspicuous scar; /amina entire, oblong to oblong-lanceolate, slightly

oblique, 6—23 x 2—8 cm, coriaceous to stiffly coriaceous, drying discolorous,

base subacute, apex caudate-acuminate with a slightly prominent apiculate

tubule; midrib raised abaxially, slightly sunken adaxially; primary venation

densely pinnate, slightly raised abaxially and adaxially; interprimaries

parallel to primaries and only slightly less prominent, very slightly raised

abaxially and adaxially; secondary venation + obscure in fresh material,

Rhaphidophora in the Philippines totod

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Figure 1. Rhaphidophora acuminata Merr.

A. flowering shoot x '/,; B. leaf lamina x 7/,; C. venation detail x 3; D. inflorescence, spathe

fallen x 1; E. spadix detail, post anthesis x 8: F. gynoecium. three quarter view x 8. A Wenzel &

Mendoza 7857, B—C Mendoza 7859, D—F from Wenzel 1139.

PbS Gard. Bull. Singapore 52 (2000)

visible as a faint reticulum in dried specimens; inflorescence solitary,

subtended by a fully developed foliage leaf and several degraded cataphylls;

peduncle slightly compressed-cylindric, 7—26 x 0.3—0.6 cm; spathe canoe-

shaped, stoutly short-beaked, 8—14 x 1.2—5 cm, stiff-fleshy, green, falling

(at female receptivity ?); spadix cylindrical, sessile to short stipitate, inserted

obliquely on peduncle or stipe, 5—12 x 1—2 cm, white; stylar region weakly

rhombohexagonal, often barely angled, 0.9—1.2 x 1—1.1 mm, weakly

conical; stigma cylindrical, raised, c. 0.2 mm diam.; anthers strongly exserted

at anthesis; infructescence 10O—17 x 2—4 cm.

Distribution: Philippines (Leyte, Luzon, Mindanao). Endemic.

Habitat: Remnants of primary forest, invaded by secondary elements, 110—

700 m altitude.

Notes: 1. Very similar to R. perkinsiae but distinguished by the stiffer,

smaller leaves, the denser primary lateral and interprimary venation, the

wide ligulate petiolar sheath, the semi-persistent cataphylls at the active

shoot tips and the prominent cylindrical style.

2. Rhaphidophora lagunensis is treated here as a synonym of R. acuminata.

It differs primarily in its smaller stiffer leaves. However, all specimens

pertaining to R. /agunensis are in mature ripe fruit and have lost much of

their pre-fertilization detail in the preserving process, thus it is not possible

to say whether the distinctive stigma of typical R. acuminata is present in

R. lagunensis. Placing it in synonomy is provisional.

Other specimens seen: LUZON. Sierra Madre Mountains, NNW of

Dingalan, Jacobs 7857 (K, L, PNH); Batangas, Ramos & Deroy BS 22669

(K, P, US). MINDANAO. Bukidnon, Mt Katanglad, south slope of south

peak, So. Calambra, Bo Kibangay, Sulit PNH 10042 (L, PNH).

2. Rhaphidophora banosensis P.C. Boyce, nom. nov.

[Rhaphidophora stenophylla Elmer, Leafl. Philipp. Bot. 8 (1919) 3073; Merr.,

Enum. Philipp. Flower. Pl., 1 (1923) 176, nom. illeg.; non R. stenophylla K.

Krause, Bot. Jahrb. Syst. 49 (1912) 94 — Type: Philippines, Luzon, Laguna,

Los Banos (Mt. Maquiling), June — July 1917, A.D.E. Elmer 18449 (PNH7,

holo; MO, iso)]

Figure 2

Medium to large, moderately robust, semi-pachycaul homeophyllous liane

Rhaphidophora in the Philippines

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Figure 2. Rhaphidophora banosensis P.C. Boyce

A. flowering shoot x 7/,; B. leaf lamina x '/,, C. venation detail x 4; D. leaf lamina x 13. Beat

lamina x '/,; F. inflorescence, spathe fallen x */,; G. spadix detail, post female receptivity x 6; H.

spadix detail, early fruiting x 6. A-E,G & H from Ramos & Edano BS 75472; E from Elmer

839858.

224 Gard. Bull. Singapore 52 (2000)

to unknown ultimate height; seedling stage and_ pre-adult plants not

observed; adult shoot architecture comprised of elongated, clinging,

physiognomically monopodial, leafy, non-flowering stems and simple to

slightly elaborated, free, sympodial, moderately leafy, flowering stems; stems

smooth, climbing stems terete in cross-section, free stems more or less

terete to very weakly four-angled in cross-section, without prophyll,

cataphyll and petiolar sheath fibre, internodes to 2.5 x 0.75 cm free shoots,

separated by prominent oblique leaf scars; flagellate foraging stems not

observed; roots not observed; leaves scattered-distichous on flowering

shoots; cataphylls and prophylls membranous, very quickly drying and

falling; petiole very shallowly canaliculate adaxially, 9—10 x 0.3—0.4 cm,

smooth, with a slight apical and prominent basal geniculum; petiolar sheath

prominent, extending to and encircling the apical geniculum, very swiftly

drying and falling to leave a thin continuous scar from the petiole base,

around the top of the apical geniculum and back to the base; /amina entire,

falcate-lanceolate to falcate-oblanceolate, 13.5—32.5 x 2.5—7 cm, thinly

coriaceous, base to acute or slightly rounded, apex acuminate with a

prominent apiculate tubule; midrib raised abaxially, almost flush adaxially;

primary venation pinnate, raised abaxially and slightly less so adaxially;

interprimaries parallel to primaries, less prominent, slightly raised abaxially

and adaxially; secondary venation weakly reticulate, slightly raised abaxially

and adaxially, but not at all prominent; inflorescence solitary, subtended by

a fully developed foliage leaf and one or more soon-falling cataphylls;

peduncle compressed-cylindric, swollen at insertion of spadix, 9 x 0.7 cm;

spathe not observed; spadix cigar shaped, sessile, inserted obliquely on

peduncle, c. 10 x 1 cm; stylar region well developed, rhombohexagonal,

1.5—2 x 1.5—2 mm, truncate; stigma slightly raised-punctiform, c. 0.3 mm

diam.; anthers not exserted at anthesis; infructescence stoutly cylindrical, c.

ks -& 2 con,

Distribution: Philippines (Catanduanes, Luzon). Endemic.

Habitat: Margins of forests.

Notes: 1. Immediately identifiable by the long, narrow falcate leaf lamina

and relatively short petiole.

2. Elmer published the name R. stenophylla for this species apparently

unaware that name was already occupied for a species described by Krause

from New Guinea.

Other specimens seen: CATANDUANES. Catanduanes, Ramos & Edano

Rhaphidophora in the Philippines 225-

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Figure 3. Rhaphidophora conocephala Alderw.

A. flowering shoot x '/,; B. inflorescence, spathe fallen x 1'/,; C. spadix detail, female receptivity

x 10; D. gynoecium, three quarter view x 6; E. spadix detail, post anthesis x 10; F. portion of

preadult sterile shoot x ’/,; G. leaf lamina x '/,; H. venation detail x 3. A-E, G & H from Nur

SFN 7369; F from Lérzing 11750.

226 Gard. Bull. Singapore 52 (2000)

BS 75472 (SING). LUZON. Bulacan Province, Angat, Ramos & Edajio

BS 34035 (US); Zambales Mts, Pinaglubo, Loher 7050 (K).

3. Rhaphidophora conocephala Alderw.

Rhaphidophora conocephala Alderw., Bull. Jard. Bot. Buitenzorg III, 1

(1920) 384 — Type: Indonesia, Sumatera, Sibolangit, 10 May 1917, Lérzing

5137 (BO, holo; K, L, iso).

Figure 3

Large, moderately robust, semi-pachycaul homeophyllous liane to 15 m;