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

(Editor)

Singapore Botanic Gardens

Dr T.W. Foong

(Assist. Editor)

Singapore Botanic Gardens

Dr S.C. Chin

Singapore Botanic Gardens

Dr M.J.E. Coode

Royal Botanic Gardens

Kew, U.K.

Dr R.T. Corlett

University of Hong Kong

Hong Kong

Dr M.C. Roos

Rijksherbarium

Leiden, Netherlands

Dr E. Soepadmo

Forest Research Institute Malaysia

Kepong, Malaysia

Dr W.K. Tan

Singapore Botanic Gardens

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

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

conclusions expressed by the contributing authors.

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Instructions for contributing authors are found on the inside backcover.

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The Gardens' Bulletin

Singapore

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VOL. 53 (Parts 1 & 2) July 2001 ISSN 0374-7859

CONTENTS

Argent, G.

Contributions to the flora of Mount Jaya VI. A new banana species, Musa johnsii

‘(ERS S VEN sere ye oud USS al Gr BGT TE ye at el es ee Se 8 1

Bastmeijer, J.D. and R. Kiew

A new Cryptocoryne hybrid (Araceae) from the Bukit Timah Nature

[PEGSSST ASS STUEYEG I YO) 1 Di we Re Sees ts ko re ee di ee aR ee 9

Boyce, P.C.

The genus Rhapidophora Hassk. (Araceae-Monsteroideae-

LL CLES SETERNE ) FURY J BY OY REEL) ns gece SRN RN IR a a ee ee ene eee 19

Boyce, P.C.

The genus Rhaphidophora Hassk. (Araceae-Monsteroideae-

Monstereae) in New Guinea, Australia and the tropical Western

JPPEYES ABTS Aceepanse te se i es ey aga Ra as pt cree ee 75

Jong, K. and G. Argent

Cytology of two new species of Musa (Musaceae) from Borneo and

PRE MES CHOTA mel ATI OTSH peat ook ese sas sree ce cote t cs ae -conecscesostressacensesuecerescveusasenseeicecaestsesseseteees 185

Khaw, S.H.

The genus Evlingera (Zingiberaceae) in Peninsular Malaysia

(UECE PECTED Gh INE AS ES (BS ee i cence rece ee Ey Se ee 191

Kiew, R.

The limestone begonias of Sabah, Borneo — flagship species for

COTS EVE ON reese eR oa Se a cel 241

Kiew, R.

Begonia conipila Irmsch. ex Kiew (Begoniaceae) from the Gunung

MulurNattonallbark (Sara wakes ial ayslays ccs. tee tsscasccecnesccesucvsedvectessestencnsesvensennsevasccveeeusooeae 287

Ng, F.S.P.

New species, varieties and reductions in Diospyros (Ebenaceae) in Borneo and

Peninsular Malaysia including peninsular Thailand .......0.........csssscssseseseseseseeseseseeeseseeees 291

Tan, B.C. and E. H. Mandia

New and noteworthy records of mosses from Mindoro, the

Philippines, and thei bioseopraphicallimp lication ccccecceeescecsssceceesaceeteerteee seater 315

Turner, I. M.

New combinations of MallestampWiyii aCe ae ese eeensnaaaes cane eecee seta eee eee eens 323

Wong, C., R. Kiew, A. Lamb, Ohn Set, S. K. Lee, L. H. Gan and Y. Y. Gan

Sectional placement of three Bornean species of Musa (Musaceae) based on

Amplitied Fragment Length) Polymorphism (ABIOR) ie ccecectereceresp eters cereee tere eee 327

Yii P.C. and P.P.K. Chai

New combinations, new names and new species of Madhuca

(Sapotaceae) from Sabahand! Sarawak, BOmme Occ certs eeteeeeteeerte eee OAS

Corrigenda — I. Salma, The significance of pollen morphology in the

taxonomy of the eenus Durioy(Bombacacac)) ec reececee ceteerseee tere teen cetera cence 357

BOOK REVIEW

Secondary Forest Trees of Kalimantan Indonesia. A Manual for 300

Selected! SPE CieS -.....cccscccseccsencarenencbewadstescssasencetve aepaeanc tesa cree ees taser o eee ees F.S.P. Ng - 359

Obituary Dr Gunnar Seidentadem\ cere cece errata eee eee ee nee J. J. Vermeulen - 363

Date of Publication: 31 July 2001

Published by

National Parks Board

Singapore Botanic Gardens

Cluny Road

Singapore 259569

Printed by Oxford Graphic Printers Pte Ltd

Gardens’ Bulletin Singapore 53 (2001) 1-7.

Contributions to the Flora of Mount Jaya VI.

A New Banana, Musa johnsii (Musaceae) from

New Guinea

G. ARGENT

Royal Botanic Garden Edinburgh, EH3 5LR, U.K.

Abstract

A new species of banana, Musa johnsii Argent, is described and illustrated from Papua

(formerly Irian Jaya), Indonesian New Guinea, and its possible position in the genus is

discussed.

Introduction

New Guinea is an important centre of diversity for the genus Musa. With

the addition of this new species, there are ten wild species recorded most

of which are endemic. The area is also arguably the richest place in the

world for indigenous cultivars (Simmonds 1966), many of which are diploids

and potentially important as a gene reservoir. This new banana has all the

attributes of a wild species in that it has a non-parthenocarpic seedy fruit

and regenerates naturally from seed distributed by animals, yet it has a

sterile, non-seedy terminal portion and was cultivated in at least one village

(Utekini) for its use as a vegetable.

Terminology and description follow previous tradition as used by

Simmonds (1962, 1966) and Argent (1976).

Musa johnsii Argent sp. nov.

Species insignis propfer infructescentia densum paene sphaericum fructuum

subsessilium primo ad apices truncatis demum irregulariter schizocarpicis.

Ab omnibus species Musae differt fructu maturo cum tertia parte distali

medullae sterilis composita.

Typus: Indonesia: Papua, Freeport mining concession above Timika. ike

Nov 2000 Argent et al. 00562 (holo BO; iso K, E).

Fig 1 & 2.

Clump forming herbaceous plant. Pseudostem to 4 m tall, 28 cm diam.

near the base. Predominantly brown with dead clinging pseudostems,

Gard. Bull. Singapore 53 (2001)

Figure 1. Musa johnsii, vegetative features: A. petiole bases x '/,, B. T.S. petiole x 7/s, C. leaf

base x '/,.

otherwise green with some dark brown coloration in the upper parts, no

wax; undersheath cream to white, juice milky white or creamy. Suckering

moderate, arising alongside or up to 20 cm from parent, erect or nearly so.

Sucker leaves mostly auriculate. Shoulder brown or green, entire, smooth

and appressed, without any scarious margin. Fourth last leaf 242 x 88 cm,

right handed to c. 8 mm, (almost symmetrical), the base cordate to weakly

auriculate. Other leaves often left-handed and predominantly strongly

auriculate, all green, hardly different in colour above and below, slightly

paler beneath and with the prominent midrib, mostly pale yellow, sometimes

with a little brown proximally without obvious wax. Petiole 52 x 4.5 cm,

the adaxial channel green, open with only slightly incurved or erect margins

which are green, not or only narrowly (to c. 1 mm) scarious, abaxially the

petiole dark chocolate brown TS ratio 0.78 (see Argent 1976). PB ratio 4—

Peduncle stout, green, glabrous, densely scarred. Bunch horizontal, diagonal

or occasionally held completely vertically downwards. The female bracts

lanceolate to 54 x 12 cm, yellow, shiny outside, dull yellow and slightly

paler inside, acuminate the apical 12 cm with the margins strongly inrolled,

completely and quickly deciduous. Female flowers with a few staminodes

Musa johnsii 3

Figure 2. Musa johnsii, fertile features: A. habit of fruit bunch with male bud x '/,, B. T.S. fruit

x */3,C. LS. fruit x */;,D. whole fruit x ?/,, E. female bract after dehiscence x '/,, F. free tepal of

female flower x 1, G. style and stigma of female flower x 1, H. compound tepal of female

flower x 1, I. male flower x 1, J. free tepal of male flower x 1, K. stamen from male flower x 1,

L. compound tepal of male flower x 1, M. style and stigma from male flower x 1.

4 Gard. Bull. Singapore 53 (2001)

mostly less than half the length of the style, free tepal flushed purple.

Ovary trilocular, each locule with the ovules in two rows.

Male peduncle growing vertically downwards. The male bud up to 14 x 11

cm, shiny yellow, imbricate for c. 2 cm from the tip. Male bracts 13 x 7 cm,

shiny rich yellow outside, slightly darker near the margins, the tips pale

green, shiny yellow inside although becoming dull inside after falling, with

broadly rounded, obtuse apices, lifting to a high angle c. 45° to the axis;

after falling only recurved at the base not revolute from the apex or margins.

Male flowers, two-rowed, falling in a group, cream, the free tepal translucent

white, with a rounded, erose upper margin and no wrinkle, just over half

as long as the compound tepal. Compound tepal cream with pale yellow

apices, the two longest points with irregular papillose to subdenticulate

margins.

Fruit bunch dense, sub-spherical in shape to 17 x 18 cm. Fruit two-rowed,

the second hand with 14 fruits. The fruits irregular, apparently ageotropic

as they show no curvature in any part of the bunch, ripening orange but

the surface becoming mostly blackish by full maturity, 8—9 x 2.5—3.5 cm,

3-, 4- or 5-angled, broadly truncate at the apex with large scars up to 2 cm

diam., splitting irregularly to reveal pale pinkish orange pith and similarly

coloured or yellowish flesh around the seeds in the carpel chambers, the

pith white at the base of the fruit; yellow latex exuding from the cut skin of

the immature fruit. Pedicel c. 2—3 mm, the fruits almost sessile. Seeds

dark brown, 4—5 mm diam., irregular with rounded angles and with a

distinctly domed boss opposite the hilum which is c. 2 mm in diam. and

vertically striate, also with a raised band on one side from hilum to boss.

Vernacular name: Mel in the Amungkal language of the Amungme Tribe.

Uses: The Amungme people eat this species raw as a vegetable, slicing the

young stalks and mixing them with local Begonia leaves (nilmanep) and

salt.

Notes: Named in honour of Professor R.J. Johns who has made a lifetime

study of the plants of New Guinea and who first drew my attention to this

species.

This new species was a very surprising find on the southern side of

the main range north of Timika. It is very distinctive in the dense subspherical

bunch of almost sessile fruits, which are rather irregularly schizocarpic, and

are truncate at the apex. Unlike any other Musa species, it has a sterile

mucilaginous pith chamber in the distal third of the mature fruit.

Musa johnsii >

Figure 3. Professor R.J.“Bob’ Johns with the fruit bunch and male bud of Musa johnsii.

6 Gard. Bull. Singapore 53 (2001)

Two populations were seen. The first where it was semi-cultivated

for the edible leaf stalks near Utekini below Tembagapura at about 1600

m. The second was on the Darnell Ridge below the Hanekam Tunnel

between 1000—1500 m altitude, where several groups of plants occurred

along the roadside well away from any indigenous habitation. The largest

of these populations of more than a hundred plants displayed all stages

from abundant small seedlings to young clumps, plants flowering in the

female phase and older ones with mature fruit and male inflorescences.

They thus appear to be totally non-seasonal.

The fruits are puzzling. The distal pith chamber could be an attractive

lure to the fruit but, for a wild banana, the number of seeds in the very

restricted loculi is very low (an average of 46 for the 5 fruits for which

seeds were counted) compared with many hundreds in most other wild

Musa species. The flesh of this species in either the pith chamber or

around the seeds in the loculi did not seem to be particularly palatable and

the local people are not reported to eat the fruit at any stage. Animals

were certainly visiting the ripe fruits and removing the contents and many

of the seeds were damaged in the process. The fact that the female bracts

are completely deciduous is remarkable in that the mature fruits are tightly

packed in a very dense bunch in the sense of Argent (1976). This is no

doubt achieved because of the strong revolution of the bracts from both

base and apex, which results in their falling in tight rolls.

This species grows at the highest altitude of the three wild bananas

encountered in the Freeport project area around Timika. Musa banksii F.

Muell. was common in the lowland areas not too swampy and waterlogged.

M. lolodensis Cheesman occurred mainly at the upper margin of the heath

forest at about 500 m altitude, although one clump was seen growing with

Musa banksii at less than 100 m. Both these species can be easily

distinguished from M. johnsii even in the sterile state, as they both have

broad scarious margins at the upper edge of the leaf sheath or ‘shoulders’

(Argent 1976). The intense green colour of the foliage and orange colour

of the fruit indicate that this species probably belongs in section

Australimusa. It would appear to be closely related to M. lolodensis. The

seeds of the two species are remarkably similar. Apart from the vegetative

difference mentioned, the fruit of these two species are very different. M.

lolodensis has a lax bunch of distinctly long pedicellate fruit, which splits in

a much more regular fashion on ripening than this new species.

There was another wild banana reported from Mt Jaya, which could

be Musa ingens Simmonds, but this was not seen by the author. It is very

distinct from this new species as it has glaucous waxy pseudostems, a

reflexed ‘shoulder’ and a long pedicillate fruit with much larger seeds 8—9

mm in diameter.

Musa johnsii 7

Acknowledgements

This work was made possible with the logistical and financial assistance

from PT Freeport Indonesia and Rio Tinto in conjunction with Herbarium

Bogoriense, Kebun Raya Bogor and The Biodiversity Centre, Cenderawasik

University, Manokwari. I am also grateful to the Royal Botanic Gardens,

Kew for permission to join their expedition and to Prof R. J. Johns and Dr

T. Utteridge for their companionship and tireless support in organising the

expedition. Dr Carolyn Cook provided the local names and much valued

help at Tembagapura. The Royal Botanic Garden, Edinburgh gave

permission for me to participate in the fieldwork and Ms Christina Oliver

ably interpreted the specimens, photographs and crude field drawings. I

am indebted to Dr Robert Mill for the translation of the Latin diagnosis.

References

Argent G.C.G. 1976. The wild bananas of Papua New Guinea. Notes

Royal Botanic Garden Edinburgh. 35: 77—114.

Simmonds, N.W. 1962. The Evolution of the Bananas. Longmans, U.K.

Simmonds, N.W. 1966. Bananas. 2" ed. Longmans, U.K.

Gardens’ Bulletin Singapore 53 (2001) 9-17.

A New Cryptocoryne Hybrid (Araceae) from the Bukit

Timah Nature Reserve, Singapore

JAN D. BASTMEIJER

Oude Roswinkelerweg 72, 7822 AG Emmen, The Netherlands

AND

RUTH KIEW

Singapore Botanic Gardens, | Cluny Road, Singapore 259569

Abstract

A Cryptocoryne from the Bukit Timah Nature Reserve, Singapore, identified for many

years as C. griffithii Schott, is recognized to be a new hybrid species: Cryptocoryne xtimahensis

Bastmeijer. It is difficult to accurately determine its parents, but good candidates are C.

nurii Furtado and C. cordata Griff., both native to Johore, Peninsular Malaysia.

Introduction

The first collections of Cryptocoryne from Singapore were made in 1890

by H.N. Ridley (1904, 1907, 1925), the first director of the Botanic Gardens

Singapore. Two species were recorded: C. griffithii Schott from several

freshwater localities and C. ciliata (Roxb.) Schott from tidal areas. Today,

C. griffithii is still present in the MacRitchie Reservoir and Nee Soon

swamp forest but C. ciliata is reported to have disappeared from Singapore

(Tan, 1995). The latter is, however, a widespread species, occurring from

India to Papua New Guinea, while C. griffithii is a rather narrow endemic

being known from the south of Peninsular Malaysia (Malacca and Johore),

as well as in Singapore. However, today there may be only a few localities

left in southern Johore of the Malaysian population.

The main interest in Cryptocoryne is as an ornamental plant for

aquascaping. This hobby of cultivating tropical plants and keeping fish in

aquaria has become very popular in Europe, the USA, and Japan. From

the 1950s, Singapore was the main port for shipping wild collected plants

to these regions, and plant nurseries developed to satisfy this demand.

The attention of the first author was drawn to the problem of the

identity of the Bukit Timah plant by a photograph of C. griffithit on the

website of the Singapore Science Centre from an article by Tan et al.

10 Gard. Bull. Singapore 53 (2001)

(1994). The photograph presented there of C. griffithii did not match the

European idea of C. griffithii very well. The same photograph appeared in

Foulis et al. (1998). In June 1999, there was an opportunity to study the

plant in question more closely in the Bukit Timah Nature Reserve. We

found the plant flowering abundantly and collected some specimens for

further investigations.

A Short History of Cryptocoryne griffithii Schott

The identity of the real C. griffithii has been in dispute, especially because

of the erroneously named drawing in Hooker (1900). Ridley (1904) pointed

out that the drawing actually represented his newly described C. purpurea

Ridl.. In 1920, Engler gave a good description of both species. De Wit

(1961) pointed to this problem again, and he was fortunate to get live

plants from Malacca, from where Griffith had originally collected his plants.

A detailed description of the true C. griffithii together with a nice drawing

by Ike Zewald is found in De Wit (1961, 1971, 1990) and which closely

matches Schott’s unpublished drawing in Vienna (W) of C. griffithii.

One of the first commercial collectors of C. griffithii from Singapore

was Y.W. Ong (pers. comm.), who collected the plants in the reservoirs

and shipped them to Europe as far back as 1948. However, all publications

in Europe on this plant in the hobby magazines we could find refer to C.

purpurea. At that time there was also much confusion about C. cordata

Griff., which was not well understood, see for example Hoedeman (1948),

Wendt (1952), Oskam & Van Ramshorst (1954) and De Wit (1951, 1958a,

b). The reason for the lack of notes on C. griffithii is probably very simple:

the species is very difficult to grow and neither hobbyists nor professionals

were able to grow them at that time. But from 1961 onwards, C. griffithii

was at last a well-known species.

Around the 1970s, taxonomy in Cryptocoryne seemed to be resolved

(De Wit, 1971), but in the following years the opposite became true.

Jacobsen (1982, 1987) suggested a hybrid nature for C. purpurea. Jacobsen

& Bogner (1986, 1987) gave a detailed review on the Cryptocoryne of the

Malay Peninsula. Today, with more collections available and many

experiments on cultivation, it appears that within some groups there is a

near continuous variation, making it in some extreme examples, more or

less a matter of taste how to interpret a newly imported plant. Furthermore,

polyploidy and natural hybrids have proved to be rather common in

Cryptocoryne. Today we know a couple of C. griffithii-like plants that

show some variation. A recent review on C. griffithii and Cryptocoryne in

general is found in Bastmeijer (2000).

Cryptocoryne xtimahensis from Singapore 11

With the variation of C. griffithii now understood, it is possible to be

certain that the Bukit Timah plant is a distinct new hybrid differing, among

other characters, from C. griffithii in its narrowly ovate leaves, the long

apex of the spathe and large, irregular protuberances of the limb. The fact

that its pollen fertility was less than 10% and that it had never been found

in fruit strongly suggests that it is a hybrid.

Cryptocoryne xtimahensis Bastmeijer notosp. nov. (Araceae)

Lamina spathae protuberationibus irregularibus obtecta, longicaudata (in

C. griffithii brevicaudata et dense verrucosa), basis laminae plus minusve

auriculata (similiter in C. nurii), color laminae flavus protuberationibus

atrorubis, fauce modice angusta (similiter in C. nurii). Folia eis speciei C.

cordata simillima (sed C. cordata differt lamina spathae leavi tota flava) —

Typus: Singapore, Bukit Timah Nature Reserve, Jungle Fall Stream. 18

January 2001, dammed pools. Alt. c. 120 m., R. Kiew, S. Saifuddin, S. Teo

& A.T. Gwee RK 5127. (holo SING; iso C, K).

Rhizome creeping, 0.4—0.8 cm diam., stoloniferous, loosely rooted in the

mud or firmly fixed in gravel. Leaves in a rosette; petiole (2.5—)10—15 cm

(depending on water depth), dark green, sheathed in the lower part for

nearly one fifth of its length; leaf blade ovate, 4.5—8 cm long and 2—4 cm

wide, base rounded, margin flat to slightly undulate, apex acute, upper

surface dark green, sometimes with dark purple transverse markings, or

submerged leaves brownish green; pale green or sometimes tinged reddish

beneath.

Peduncle 2—4 cm long. Spathe 10—15 cm long; limb of the spathe

4—5 cm long and c. 1.5 cm wide, apex long caudate, base auriculate and

margins reaching each other, upper surface pale yellow with distinct dark

red irregular protuberances; collar distinct with a rim and forming a narrow

opening to the tube; tube 3—6 cm long and c. 0.5 cm in diameter, outside

and inside white; kettle 2—2.5 cm long, and c. 0.7 cm in diameter, outside

white, inside white without purple markings. Spadix usually with 6 female

flowers at the base, these pale green; stigmas white, more or less elliptic;

olfactory bodies small, yellow; naked axis between male and female flowers

c. 0.5 cm long, white; male part cylindrical, yellow, male flowers 50 — 60,

each male flower usually consisting of two stamens; appendix cone shaped,

c. 2mm long, white; flap above the male flowers white. Fruit and seeds not

known.

Emersed cultivated plants much smaller; leaves c. 10 cm long;

peduncle c. 1 cm long; spathe c. 6 cm long.

“ Gard. Bull. Singapore 53 (2001)

la. Dammed pools on the Jungle Fall Stream lb. Limb of the spathe of Cryptocoryne

in the Bukit Timah Nature Reserve, xtimahensis.

Singapore. photo Bastmeijer

photo Bastmeijer

lc. Cultivated plant of Cryptocoryne ld. Opened kettle showing the female

xtimahensis. (bottom) and male flowers (top).

photo van Wijngaarden photo van Wijngaarden

Figure 1. A new Cryptocoryne hybrid from the Bukit Timah Nature Reserve, Singapore

Cryptocoryne xtimahensis from Singapore 13

Chromosome number: 2n = 54. Pollen fertility < 10%. Vouchers at

Copenhagen (C).

Other collections examined: All from Bukit Timah in Jungle Fall Valley —

the type and J. Bastmeijer 807 (L), P. Blanc 91-1 (SING), K.S. Chua 333

(SINU), D. Nicolson 1367 (US) and J. Sinclair 5200 (E).

Discussion

While plants identified as Cryptocoryne griffithii are known from several

localities from Singapore, this new hybrid has only been collected from

- Bukit Timah. (There is just one collection, Ridley s.n. 1892 (SING), from

Bukit Timah that may be C. griffithii). All collections of C. xtimahensis are

from the Jungle Fall Valley, where the plant grows in two pools formed

up- and down-stream by an artificial dam in the tiny stream (Fig. la).

Constructed during the Japanese Occupation, it is currently silted up. The

hybrid appears to be restricted to these pools.

The first systematic analyses of pollen fertility and chromosome

numbers in Cryptocoryne were made by Jacobsen (1977) and Arends et

al.(1982). With pollen fertility near zero for this plant, we are no doubt

dealing with a hybrid. In addition, compared with C. griffithii in Singapore,

for which half the herbarium specimens were collected in fruit, none of the

herbarium specimens of this Bukit Timah population have fruits nor have

fruits been observed in the field.

The hybrid status is not, however, unique for Cryptocoryne. The

popular aquarium species C. xwillisii Reitz from Sri Lanka forms a complex

of hybrids in which C. parva De Wit is one of the parents, the other parent

may be C. beckettii Trimen, C. walkeri Schott or C. undulata Wendt

(Jacobsen 1981a,b). Another example is found in Tasek Bera, Peninsular

Malaysia, where a very big population of C. purpurea grows (often

erroneously ascribed to C. griffithii), which also has a pollen fertility of

about zero and is today regarded as a hybrid between C. cordata and C.

griffithii (Jacobsen 1987).

In classifying Cryptocoryne species, the main characters used are

those of the limb of the spathe. C. xtimahensis has large, irregular, red

protuberances on the limb of the spathe, the collar is very pronounced and

the base of the limb is cordate (Fig. 1b). These characters are distinct from

those of C. nurii, where the limb of the spathe is rather elongated and has

a yellow background, characters shared by C. cordata.

In addition, the leaves of C. xtimahensis are narrowly ovate with a

rounded to cordate base and an acute tip as are those of C. cordata. The

14 Gard. Bull. Singapore 53 (2001)

transverse markings on the upper leaf surface of most plants of C.

xtimahensis (Fig. 1c) are also a character of C. nurii.

The combination of these characters makes C. griffithii less likely as

one of the parents. The limb of C. griffithii is verrucose with small, regular

bumps, although the collar is very pronounced too. However, the leaves of

C. griffithii are broadly ovate. Both C. nurii and C. cordata grow in the

southern part of Johore, Peninsular Malaysia but have never been recorded

from Singapore. An artificial cross between these two species would be

very interesting but would be very difficult to perform, because neither of

the species is easy to cultivate. DNA analysis should be able to elucidate

the exact parentage.

Base chromosome numbers in Cryptocoryne are x = 10, 11, 14, 15, 17

and 18. Polyploidy is rather common in Cryptocoryne and even rather long

polyploid series exist (Arends et al., 1982). The chromosome number of

this new hybrid, 2n = 54, is remarkable because it could indicate that this

plant is of triploid origin with base number 18. This number is known only

from three Sri Lankan species, the C. crispatula-group from mainland Asia,

and the tidal species C. lingua Engl. from Sarawak. All known species

from the Malay Peninsula have base number 17, except for the distinct C.

longicauda Becc. ex Engl. (also known from Sumatra and Borneo), which

has base number 15 and the tidal species C. ciliata, which has base number

11. The number 2n=54 may be explained as originating from a triploid

number with 2n = 51 after which a second generation of aneuploid

segregation may have occurred (Orgaard et al. 1995).

The Future of Cryptocoryne Habitats in Singapore

The status of the Bukit Timah Nature Reserve provides a safe habitat for

this plant. However, C. xtimahensis occupies a very restricted area c. 10 m

long and 3 m wide. It is therefore extremely vulnerable to any hydrological

changes to this one particular stream. C. griffithii seems to be rather common

in the reservoirs (Chan, 2000), and provided there is no change in land use

will not be threatened.

C. ciliata, the other native Cryptocoryne of Singapore, is said to

have been re-introduced with material originating from southern Johore.

On the other hand, several species of Cryptocoryne have been

introduced in the past, for example C. wendtii De Wit, C. xwillisii Reitz

from Sri Lanka, and C. lingua from Sarawak (Chan, 2000). They probably

‘escaped’ by accident from the numerous nurseries, which produced - from

the 1960’s onwards - large quantities of these popular plants for aquascaping,

mainly for export. It will be interesting to investigate the flora of Singapore

in this respect.

Cryptocoryne xtimahensis from Singapore 15

Acknowledgements

The National Parks Board of Singapore is acknowledged for permission to

collect in the Bukit Timah Nature Reserve and to J. Lai and S. Saifuddin

for showing JDB the population. The aquatic plant firms Tropica

(Denmark) and Oriental (Singapore) were of great help during JDB’s stay

in Singapore. Gitte Petersen (Copenhagen, Denmark) made the

chromosome counts. Dr. H. Roessler (Munich, Germany) translated the

diagnosis into Latin. Piet van Wijngaarden cultivated the plants in the

Netherlands and was able to flower them after a couple of months. Niels

Jacobsen (Agricultural University, Copenhagen, Denmark) and Josef

_ Bogner (Botanical Garden, Munich, Germany) are acknowledged for their

great support in discussing taxonomic problems in Cryptocoryne, and for

comments on the manuscript.

References

Arends, J.C., J.D. Bastmeijer and N. Jacobsen. 1982. Chromosome numbers

and taxonomy in Cryptocoryne (Araceae).II. Nordic Journal of Botany.

2: 453—463.

Bastmeier, J.D. 2000. The Crypts Pages. http://users.bart.nl/~crypts/

index.html

Chan, S.Y. 2000. The Freshwater Macrophytes of Singapore. http://

molluscan.com/macrophytes

Engler, A. 1920. Araceae - Aroideae Cryptocoryne. Das Pflanzenreich.

IV.23.F: 232—249.

Foulis, L. et al. (eds.) 1998. Botanica. 2" ed. Periplus Editions, Singapore.

Hoedeman, J.J. 1948. Het geslacht Cryptocoryne Fischer, 1928. Het

Aquarium 19(3): 61—64.

Hooker, J.D. 1900. Cryptocoryne griffithii. Botanical Magazine 126, pl.

eho)

Jacobsen, N. 1977. Chromosome numbers and taxonomy in Cryptocoryne

(Araceae). Botanisher Notiser. 130: 71-87.

Jacobsen, N. 1981a. Cryptocoryne undulata Wendt und Bemerkungen zu

anderen Arten. Aqua-Planta. 6(2): 31—38.

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Jacobsen, N. 1981b. Cryptocoryne undulata Wendt und Bemerkungen zu

anderen Arten — Ein Nachtrag. Aqua-Planta. 6(4): 92—94.

Jacobsen, N. 1982. Cryptocorynen. Kernen Verlag. Stuttgart, Germany.

Jacobsen, N. 1987. Cryptocoryne purpurea Ridley. Aqua-Planta 12(2): 61—

62.

Jacobsen, N. and J. Bogner. 1986. Die Cryptocorynen der Malatischen

Halbinsel (1.Teil). Aqua-Planta. 11(4): 135—139.

Jacobsen, N. and J. Bogner. 1987. Die Cryptocorynen der Malaiischen

Halbinsel (2.Teil). Aqua-Planta. 12(1): 13—20.

Ong, Y.W. 1999. Private communication. Singapore.

@Mrgaard, M., N. Jacobsen and J.S. Heslop-Harrison. 1995. Molecular

cytogenetics in the genus Crocus L. In: P.E. Brandham and M.D. Benneth

(eds.), Kew Chromosome Conference IV, pp. 291-299. Royal Botanic

Gardens, Kew, U.K.

Oskam, H.C. and J.D. van Ramshorst. 1954. Cryptocoryne griffithii Schott.

Het Aquarium. 24(7): 146—149.

Ridley, H.N. 1904. New Malayan Plants. Journal of the Straits Branch of

the Royal Asiatic Society 41: 44—45.

Ridley, H.N. 1907. Cryptocoryne. Materials for a Flora of the Malayan

Peninsula. Pp. 3-6.

Ridley, H.N. 1925. Cryptocoryne. The Flora of the Malay Peninsula. 5:

86—88.

Schott, H.W. 1856. Aroideae. Synopsis Aroidearum. 1: 1—3.

Tan, H.T.W. 1995. A Guide to the Threatened Plants of Singapore. Singapore

Science Centre, Singapore.

Tan, H.T.W., ILM. Turner, Y.C. Wee and K.S. Chua. 1994. Plants in the

Singapore Red List. Singapore Scientist. 72: 26—32.

Wendt, A. 1952-1958. Die Gattung Cryptocoryne Fischer. Die

Aquarienpflanzen in Wort und Bild. Stuttgart, Germany.

Wit, H.C.D. de. 1951. Cryptocoryne cordata, Cryptocoryne griffithii. Het

Aquarium. 22(6): 128—129.

Cryptocoryne xtimahensis from Singapore |

Wit, H.C.D. de. 1958a. Cryptocoryne griffithii. Fishkeeping. January: 138.

Wit, H.C.D. de. 1958b. Cryptocoryne griffithii and its allies. Fishkeeping.

February: 230.

Wit, H.C.D. de. 1961. Het genus Cryptocoryne (17). Het Aquarium. 31(10):

232—233.

Wit, H.C.D. de, 1971. Aquarienpflanzen. Ulmer, Stuttgart, Germany.

Wit, H.C.D. de, 1990. Aquarienpflanzen. 2° ed. Ulmer, Stuttgart, Germnay.

Gardens’ Bulletin Singapore 53 (2001) 19-74.

The Genus Rhaphidophora Hassk. (Araceae-

Monsteroideae-Monstereae) in Borneo

P.C. BoYcE

Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, U.K.

Abstract

An alpha-taxonomic account of Bornean Rhaphidophora is presented as a precursor to the

forthcoming Flora Malesiana Araceae treatment. Thirteen species are recognized, none

novel. Rhaphidophora elliptica Ridl. 1905 (non 1908 = R. elliptifolia Merr.) and R.

megasperma Engl., treated respectively as synonyms of R. montana (Blume) Schott and R.

puberula Engl. by Boyce (1999), are resurrected as Bornean endemics. New synonyms are:

Scindapsus havilandii Ridl., Rhaphidophora jaculiformis Alderw., R. subfalcata M. Hotta

into R. megasperma, and R. nigrescens Ridl. into R. sylvestris (Blume) Engl. A dichotomous

key to species is provided. All species are illustrated.

Contents

USSG Oa Bk ee 19

SL SURI Diy S202 GTEC TST 7 Eee ae ee ee ee 20

EG OD) 0 ee Ne a tg ee ee ee 21

Key to adult flowering Rhaphidophora in Bore ..........cccceceseeseeeeeeteeees 24

SUE OA eee Rn an ok ce et ee i Ce ee 26

ALES DAU US 176 1 oe ie ee eee ee 72

ELS NST WES be as By ee ane ee oo ee ee eee 42

[UES RSTO es gE oa ue eR Se no er 73

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 Himalaya (SE Nepal to NE

Vietnam, roughly 17°—23° N), West Malesia (including southernmost

peninsular Thailand), the Philippines, and East Malesia.

This is the fifth in a series of papers intended to present a complete

20 Gard. Bull. Singapore 53 (2001)

alpha-taxonomy of the genus Rhaphidophora in Asia. Accounts for

Peninsular Malaysia and Singapore, the southern and western Indonesian

archipelago and the Philippines have been published recently (Boyce, 1999,

2000a, 2000b) together with a partial account for New Guinea (Boyce &

Bogner, 2000) and an account for Papuasia and the tropical Western Pacific

appears elsewhere in this journal (Boyce, 2001). Accounts for the Himalaya,

and Thailand and Indochina are being prepared and will be published

separately. All morphological terms employed follow Stearn (1992).

The last complete revision of Rhaphidophora was that of Engler &

Krause (1908). A summary of the taxonomic and nomenclatural history of

Rhaphidophora was presented in Boyce (1999) and a partial infrageneric

classification with an overview of informal morpho-taxonomic groups was

proposed by Boyce (2000a). Boyce (1999) provides a detailed discussion of

structure, generic limits, together with keys to the genera of monsteroid

and anadendroid aroids in Malesia.

Synonymy cited is for the species, not for the review area. The reason

for this is that synonymic names based on types from outside the review

area are frequently applied to specimens in local herbaria.

Geography and Endemism

Borneo has a relatively poor Rhaphidophora flora, with comparatively few

species recorded, although how much this is a reflection of inadequate

collecting is not clear. However, of the 13 species recognized for the island

five are endemic (compared with 2/15 in Peninsular Malaysia; 4/15 in

Sumatera) and thus Borneo has, with the exception of the Philippine islands

(7/11), the highest level of endemism in Sunda.

Two of the five Bornean endemics, R. elliptica Ridl. and R. elliptifolia,

belong to the Indo-Malesian Hongkongensis Group (see Boyce, 2000a).

Both Bornean endemics are morphologically close to west Malesian species

(R. elliptica to R. montana (Blume) Schott; R. elliptifolia to R. maingayi

Hook.f.) and are presumably endemic segregates. Another species from

this group, the widespread R. sylvestris (Blume) Engl. (Sumatera, Peninsular

Malaysia, Thailand, east to Borneo) is widespread and common in Borneo.

The other Bornean endemics are R. latevaginata M. Hotta, R. megasperma

Engl. and R. cylindrosperma Engl. & K. Krause.

Rhaphidophora latevaginata M. Hotta is a neotenic shingling species

that might conceivably have arisen from a species with a pronounced

heteroblastic development, e.g., R. korthalsii Schott (see Boyce, 2000a).

Rhaphidophora megasperma is of uncertain affinity in Sundaic Asia

but has morphological similarities, e.g., clustered inflorescences subtended

Rhaphidophora in Borneo 21

by chartaceous cataphylls, to certain New Guinea species (e.g., R. versteegii

Engl.) and is tentatively included in the Hollrungii Group (note that this

group had its name changed in Boyce, 2001).

Rhaphidophora cylindrosperma is another morphologically isolated

species in Sunda but with similar species (e.g., R. schlechteri K. Krause)

east of Wallace’s Line and belongs to the Neoguineensis Group. It is

interesting that two of the Borneo endemic Rhaphidophora appear to be

morphologically closer to species in Wallacean New Guinea than to any

Sundaic species.

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. 1 (1897) 58; Ridl., J. Straits Branch Roy. Asiat. Soc.

44: 185—187 (1905); Engl. & K. Krause in Engl., Pflanzenr. 37 (IV.23B)

(1908) 17—S3; Engl. & Prantl, Nat. Pflanzenfam. Nachtr. 3 (1908) 29;

Merrill, J. Straits Branch Roy. Asiat. Soc., special number (1921) 88—90;

Schott, Icones aroideae et reliquiae (IDC Microfilm) (1983) fiche nos.

28—31, 121; 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.]

[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 (Engl.) Engl.], 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 + invisible or coagulating into yellowish, translucent

jelly and eventually hardening to a brittle amber-like mass; seedling stage

in)

Gard. Bull. Singapore 53 (2001)

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

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 (descriptive generalisations

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: i. physiognomically unbranched clinging non-flowering stems

rooting along their entire length giving rise to variously elaborated free

lateral flowering stems (e.g., R. lobbii Schott, R. puberula Engl.), or 11. all

stems physiognomically unbranched clinging and flowering (e.g., R.

korthalsii), or ii. physiognomically unbranched leader and lateral stems

clinging but only 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 later drying to leave fragmentary parchment-like remains

on petioles, developing laminas and 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, clinging to substrate;

feeding roots rare to abundant, smooth, pubescent or prominently scaly,

later often becoming woody, clinging to substrate or free; /eaves 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 canaliculate to weakly carinate, smooth or

pubescent, with variously prominent apical and basal genicula; petiolar

sheath prominent, nearly reaching to overtop 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

Rhaphidophora in Borneo 23

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 - 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 soon 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, inflorescences at male 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 male anthesis and then usually

deciduous before male anthesis is complete, occasionally persisting into

the early stages of infructescence development (e.g., R. angustata), rarely

drying and persistent (e.g., R. megasperma), stiff to rather softly 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 + cylindrical and variously angled, most often rhombohexagonal, those

upper- and lower-most on the spadix often sterile and bereft of stigma,

those uppermost frequently either scattered or partially fused to each other

and forming a rudimentary 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

male 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, thecae dehiscing

24 Gard. Bull. Singapore 53 (2001)

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: 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 phero (1 bear); refers

to the macroscopic (to 1 cm long), needle-like unicellular trichosclereids

present in tissues.

Key to Adult Flowering Rhaphidophora in Borneo

la. Mature leat lamima,pimately divided co... cee eee ee az

lb. Mature leaf lamina entire, without or without perforations, but never

pinmately, Givided.c0 a. c.bece un ea eee gence ee 8}

2a. Flowering plants usually rheophytic, rarely low-climbing on trees

beside torrential streams. Plants always associated with sandy or

ROCKY: POLES USUGEAINS esac raster 1. R. beccarii

2b. Flowering plants high-climbing on forest trees. Plants not specifically

ASSOCIALEG Wath, WATCTCOUMESES gc ccccs press expences seers 7. R. korthalsii

3a. | Geniculum and abaxial surface of lamina pubescent ................06 4

3b. | Geniculum and abaxial surface of lamina glabrous................:0.008 5

4a. Flowering on clinging stems. Leaves of mature plants extensively

perforated. Active shoot tips with black mucilage ...........c:eseeeeees

Rhaphidophora in Borneo D5

4b.

Sa.

Sb.

6a.

6b.

7a.

7b.

8a.

8b.

9a.

9b.

1a:

10b.

OS re A ole) eras OP RS Soe ene eee Lc ea 6. R. foraminifera

Flowering on free lateral stems. Leaves of mature plants lacking or

with only with scattered, perforations. Active shoot tips lacking black

(G1 eS es nk er ha cee er ach ele ee ee Soe 12. R. puberula

Leaves always shingling, even in flowering individuals; leaf laminas

stiffly coriaceous, broadly oblong-ovate-elliptic, 8—48 x 6.5—20.5

cm, bright green, slightly to markedly glaucous, base truncate-cordate

to broadly cuneate. Flowering on clinging shoots «0.0.0.0... eee

Wee Nae ae ee EE BEE IEEE tds Lisl Bos ictehowne 8. R. latevaginata

Leaves spreading in adult and flowering individuals; leaf laminas

variously coloured but never glaucous. Flowering on free or clinging

SNGVETS, a caeeces dient SER ee ye ea la ee ese ee 6

Stems scabrid to asperous. Spathe exterior minutely puberulent ....

MM tO Se oe Mic de Sogn aed Gen caret ouetbon sss Sapdoocsessyeee 9. R. lobbii

Stems smooth: Spathe extenor SlAaDLOUS -..-7..:..0..-c.scceresecseracecnseavserees iil

Clinging stems square or rectangular in cross-section; tips of active

stems with netted prophyll, cataphyll and petiolar sheath fibre.......

So eee ee 4. R. elliptica

Clinging stems variously shaped in cross-section, including square

or rectangular; tips of active stems without fibres ........... eee 8

Spadix stipitate; leaf lamina thickly coriaceous.... 2. R. conocephala

Spadix sessile; leaf lamina variously textured but never thickly

OID EEO US Gee ora nce een pes aero ee ee ee 9

Inflorescences two, three or more together, each subtended by a

prominent chartaceous prophyll and one or more chartaceous

cataphyll; leaf lamina oblong-lanceolate or oblong-elliptic, entire to

SHED PEELOLALE. PLADTOUS ...... ccccccceceenesaceaeereeoees 10. R. megasperma

Inflorescences almost always solitary, if two together then

inflorescences not subtended by prominent chartaceous prophylls

and cataphylls; leaf lamina variously shaped and almost never

perforated, if perforated then apical geniculum and abaxial surface

Eo Le TEIN TV2 20) Oe 61 212) (Re ee 10

Chingitig stems rectanpular Im ‘CrOSS-SECUION «..........-...21--seces-ceorseneese tel

Clinging stems more-or-less terete 1M CrOSS-SECTION ..........c cc eeeeeees 12

26 Gard. Bull. Singapore 53 (2001)

lla. Leaf lamina lanceolate-elliptic to falcate-lanceolate, 4.5—32 x 1.75—

8.5 cm. Spadix cylindric to weakly clavate-cylindrical, 3—6 cm long

SERS Re os ee a eee 13. R. sylvestris

11b. Leaf lamina narrowly elliptic to narrowly elliptic-oblong, 20—29 x

4.5—7 cm. Spadix cylindrical, c. 5 x 1 cm ............ 5. R. elliptifolia

12a. Leaf lamina narrowly falcate-elliptic to falcate-lanceolate or falcate-

oblanceolate, 2.5—16 x 1.2—3 cm, drying uniformly pale straw-

coloured. Spadix slender cylindrical, 2.5—7 cm long... 11. R. minor

12b. Leaf lamina subfalcate-lanceolate or oblong-elliptic, oblique, 10—

19.5 x 2.5—6.5 cm, drying very dark brown. Spadix cylindrical-obtuse,

3—10 x 1 cm (fruiting specimen only).............. 3. R. cylindrosperma

The Species

1, 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 (I1V.23B) (1908) 46; Alderw., Bull. Jard. Bot. Buitenzorg III,

1 (1920) 382; Merrill, J. Straits Branch Roy. Asiat. Soc., special number

(1921) 88; 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 PB 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)

186; Engl. & K. Krause in Engl., Pflanzenr. 37 (1V.23B) (1908) 37; Merrill,

J. Straits Branch Roy. Asiat. Soc., special number (1921) 89. — Type:

Malaysia, Sabah, Bongaya, Dec. 1897, Ridley s.n. (SING, holo).

Figure 1

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 adults in all but overall size and leaf division;

Rhaphidophora in Borneo 27

Figure 1. 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 anthesis, pre-male anthesis x 8. A, E & F from

Stone 9637;B & D from Ridley s.n.;C from Kiew 1982.

28 Gard. Bull. Singapore 53 (2001)

adult shoots 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 clinging, densely scaly; leaves 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 semi-persistent weak fibres;

lamina 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 apicule; 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; inflorescences one to three

together, subtended by a prominent cataphyll degrading to fibres before

male 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, soon falling at male anthesis; spadix cylindrical, sessile, inserted

perpendicular to peduncle, 4.5—7 x 0.6—1 cm, dull whitish; stylar region

rather well developed, mostly rhombohexagonal, c. 1—1.5 x 1—1.5 mm,

truncate; stigma elliptic, longitudinally orientated, occasionally almost

circular, c. 0.5 x 0.3 mm, often very prominent especially in dried material;

anthers exserted at male anthesis; infructescence 7—9 x 1—2 cm, mid-

green when ripe.

Distribution: Southern Thailand, Peninsular Malaysia, Sumatera (including

the Lingga Archipelago) and Borneo (Sarawak, Brunei Darussalam, Sabah,

West and East Kalimantan).

Habitat: Along rocky or sandy stream banks, often on rocks in stream or at

waterfalls in primary to slightly disturbed or old secondary lowland forest,

peatswamp forest, occasionally persisting briefly in logged over areas,

occurring on a variety of substrates. Sea level to 300 m altitude.

Rhaphidophora in Borneo 29

Notes: 1. One of possibly two obligate rheophytic species (the other is

Rhaphidophora araea P.C. Boyce - Sumatera), R. beccarii is immediately

recognizable by its adult growth form, creeping along watercourses or

attached on rocks in the water, and by the usually pinnately divided leaf

laminas. Occasionally plants are seen climbing up tree trunks on the bank

of torrential streams.

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. They are

treated here as a neotenic manifestation of R. beccarii.

3. Rhaphidophora borneensis, based on a Grabowski specimen from

Kalimantan, is a broad-leaved form of typical (1.e., divided-leaf) R. beccarii.

Other Bornean specimens seen: SARAWAK. 1° Div.: Setapok F.R., 6 miles south of Kuching,

Bogner 1507 (K, M); Sg. Entabai, Bogner 1562 (M, US); Matang Family Park, Sg. Cina,

Boyce 719 (K); Lundu, Brooke 8409 (L):13th Mile, Matang, Brooke 9459 (L): Lundu, G.

Gadin, Clemens & Clemens 21934 (K); Matang F.R., 10 miles west of Kuching, Nicolson

1282 (US); Setapok F.R., 6 miles south of Kuching, Nicolson 1343 (L, US); Matang, Ridley

s.n. (BM); Padawan, G. Merubong, Ulu Ss: Sluba, Yii $51363 (K, L, SAR, US); 2™ Div.:

Simanggang. Brooke 10729 (BM, L): 3° Div.: Kapit District, S. Bena area, Burtt 13001,

12938 (E); 4" Div.: Lambir N.P., Mile 18, Chai 539439 (K. KEP, L, SAR, US);5 Div.: 8 - 9

miles from Limbang, Sg. Bakol, Bogner 1504 (M, US); 7" Div.: En route (survey highway)

from Sg. Mah to Sg. Shinonok, Ulu Sg. Minah, Bintulu District, Hirano & Hotta 14082

(KYO): Sg. Bejangung, a branch of the Sg. Anap, Bintulu District, Hirano & Hotta 1174

(KYO); not located: Sg. Entabai, Bogner 1346 (K, M, US); Sg. Engkramut, Bogner 1393

(M, US). BRUNEI DARUSSALAM. Belait: Labi, Kg. Teraja, path along Sg. Teraja,

Boyce 245 (BRUN, K, L); Temburong: Sg. Temburong at Kuala Belalong, Boyce 396

(BRUN, K, L); Sg. Temburong near Kuala Belalong, Jacobs s.n. (BRUN, K, L); Sg.

Temburong, just downstream from Kuala Belalong, Wong 242 (BRUN, K, L). SABAH.

Kudat: Ranau, Kg. Puas area, Amin et al. SAN 94681 (K, KEP, L, SAN, SAR, SING):

Pantai Barat: Keningau, Ulu Sg. Pingas Pingas, Jimpin SAN 122024 (K, KEP, SAN):

Keningau, Shang Lian logging area, Lanas, Krispinus SAN 118444 (K, SAN): Keningau,

Camp C area, Tiulan, Lantoh SAN 102053 (K, KEP, L, SAN); Keningau, Ulu Sg. Matud,

Tangki SAN 119596 (K, L, SAN, SAR); Sandakan: Lamag, Ulu Sg. Lokan, Aban & Petrus

SAN 90675 (K, SAN): Sg. Lokan, Amin et al. SAN 97483 (K, L, SAN, SAR); VJH 45 A

Lungmanis, Dewol et al. SAN 118075 (K, KEP, L, SAN); Beluran, Sg. Baba Gibot SAN

90040 (K, L, SAN); Labuk and Sugut, western side of Bt. Doji and pass way from Telupid

to Ulu Karamuak, Kokawa & Hotta 471 (KYO, L); Telupid, Labuk and Sugut, along Sg.

Meliau, foot of G. Tawai, Kokawa & Hotta 110 (KYO, L); Mile 45 Labuk road, Meijer

SAN 44014 (K, L, SAN); KALIMANTAN. West Kalimantan: Selimban Kapuas, Main

(sub. Polak) 2092 (L); Bt. Ubili, Niewwenhuis 97 (BO): East Kalimantan: W. Koetai, No. 29

near L. Liah Leng, Endert 3013 (BO, K, L); W. Koetai, No. 5 near Lahoem, Endert 1860

(BO, K, L); Kelassar (Kelasen?), Hallier 1555 (BO); Pulau Nibung, W. Koetai, Sg. Loewai,

near Padang Loewai, Posthumus 2173 (BO); Samarinda, mouth of Batang Mahakam, Sg.

30 Gard. Bull. Singapore 53 (2001)

Mukun, near Sanga Sanga, Meijer 1136 (BO, L); No further data, Hallier 999 (BO); Batu

Lesoena, Nieuwenhuis 219 (BO).

2. Rhaphidophora conocephala Alderw.

Rhaphidophora conocephala Alderw., Bull. Jard. Bot. Buitenzorg III, |

(1920) 384. — Type: Indonesia, Sumatera, North Sumatera, Sibolangit, 10

May 1917, Lérzing 5137 (BO, holo; K, L, iso).

Figure 2

Large, moderately robust, semi-pachycaul homeophyllous liane to 15 m;

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 unbranched,

leafy, non-flowering stems and moderately elaborated, free, 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

clinging 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, clinging, pubescent; leaves

weakly spiralled on clinging shoots, slightly scattered-distichous on flowering

shoots; cataphylls and prophylls membranous, very soon 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 soon

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; lamina

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 apicule, 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,

Rhaphidophora in Borneo

Figure 2. Rhaphidophora conocephala Alderw.

A. flowering shoot x '/,; B. inflorescence, spathe fallen x 1'/,; C. spadix detail, female anthesis

x 10; D. gynoecium, three quarter view x 6; E. spadix detail, post-male anthesis x 10; F. portion

of pre-adult sterile shoot x '/;; G. leaf lamina x '/,; H. venation detail x 3. A — E, G. & H from

Nur SFN 7369; F from Lérzing 11750.

SR Sea ee race

<Q?

hay

Laue Ove :

FSET ES

Sey)

32 Gard. Bull. Singapore 53 (2001)

stoutly short-beaked, 7—9.5 x 2—3.5 cm, thickly fleshy, exterior light yellow,

interior darker, soon (?) falling at female anthesis; 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 male anthesis;

infructescence stoutly cylindrical, 6.5—7.5 x 1.8—2 cm.

Distribution: Sumatera, Kalimantan (East) and into the Philippines

(Palawan).

Habitat: Damp primary and old secondary forest. 450—1000 m altitude.

Note: Very close to and possibly indistinguishable from Rhaphidophora

sylvestris but consistent in the conical style topped with a prominent, raised

button-like stigma and the notably more coriaceous leaves.

Other Bornean specimens seen: KALIMANTAN. East Kalimantan: Batu Penalong,

Mahakam, Wiriadinata 850 (BO, L).

3. Rhaphidophora cylindrosperma Engl. & K. Krause

Rhaphidophora cylindrosperma Engl. & K. Krause, Pflanzenr. 37 (IV.23B)

(1908) 28; Merrill, J. Straits Branch Roy. Asiat. Soc., special number (1921)

88. — Type: Indonesia, northern West Kalimantan, between the greater

and lesser arms of the S. Sambar, Hallier 1164 (BO, holo).

Figure 3

Moderate leptocaul homeophyllous liane to unknown ultimate height;

seedling stage and pre-adult plants not observed; adult shoot architecture

not fully observed, flowering stems, moderately branched and leafy,

apparently pendent; stems smooth, terete, with cataphylls and prophylls

soon falling, internodes 2—3.5 x 0.8—1.2 cm, separated by prominent

straight to slightly oblique leaf scars; flagellate foraging stem not observed;

roots not observed; leaves spiro-distichous; cataphylls and prophylls

membranous, soon falling; petiole canaliculate, 5—14 x 0.2—0.4 cm, basal

geniculum very large and prominent, apical geniculum less so; petiolar

sheath narrow and not prominent, extending to apical geniculum, + short-

persistent; /amina entire, subfalcate-lanceolate or oblong-elliptic, oblique,

10—19.5 x 2.5—6.5 cm, membranous, base acute, apex acuminate; midrib

prominently raised abaxially, slightly impressed adaxially; primary venation

pinnate, prominently raised abaxially, impressed adaxially; interprimaries

ies)

Rhaphidophora in Borneo

Figure 3. Rhaphidophora cylindrosperma Engl. & K. Krause

A. flowering shoot x '/;; B. flowering shoot x '/,; C. leaf lamina x */,; D. venation detail x 2; E.

inflorescence, spathe fallen x 1; F. spadix detail, post-male anthesis x 6. A, C— D from Church

et al. 1573; B, E—F from Sidiyasa PBU 650.

34 Gard. Bull. Singapore 53 (2001)

subparallel to primaries, slightly less prominent; secondary venation

reticulate, slightly raised abaxially; inflorescence solitary, subtended by a +

fully developed foliage leaf and one or more cataphylls; peduncle terete,

2—4 x 0.3—0.4 cm; spathe slender, 9—11 x 0.5 cm, apically tapering and

ultimately stout beaked; spadix cylindrical-obtuse, sessile, inserted + on

peduncle, 3—10 x | cm (fruiting specimen only); stylar region moderately

developed, slightly rhombohexagonal, c. 1.8 mm diam., + truncate; stigma

punctiform, raised, c. 0.3 mm diam.; anthers not observed; infructescence

3—10 x 1—1.5 cm.

Distribution: Endemic to Borneo. Sarawak ae Division), Brunei Darussalam

(Belait), Kalimantan (West, Central and East).

Habitat: Lowland primary to lightly disturbed forest. 25—200 m altitude.

Notes: 1. Superficially similar to Rhaphidophora puberula but differing in

leaves completely glabrous and never perforated, by the smaller, more

slender spadix and in flowering on long, free shoots.

2. Among the Sundaic Asian species, it is not immediately apparent to

what R. cylindrosperma is related. The superficial similarity to species in

the Hookeri Group (e.g., R. puberula and R. foraminifera) is not borne out

by closer examination. The growth form with inflorescences often carried

at the tips of short shoots arising from much longer pendent stems is

reminiscent of R. lobbii, but R. cylindrosperma differs in more coriaceous

leaves, different leaf venation and in having a longer, externally glabrous

spathe. The long, pendent terete stems with upturned tips and small leaves

recall some New Guinea species, in particular R. schlechteri K. Krause and

I have provisionally included R. cylindrosperma in the Neoguineensis Group

to which those species are assigned (see Boyce 2000a).

3. Boyce 345 is provisionally included in R. cylindrosperma although it has

much more coriaceous leaf laminas with closer venation. It may represent

an undescribed species but the material is inadequate to make a decision.

Other specimens seen: SARAWAK. 1° Div: Kuching, Hewitt s.n. (SING). BRUNEI

DARUSSALAM. Belait: Sg. Deriam, Boyce 345 (BRUN, K, L). KALIMANTAN. West

Kalimantan: Serawai, 8 km NE of Desa Jelundung, Batu Lintang, 1 km S of camp along

hunting trail, Church, Mahyer & Afriastini 1573 (BO, E, GH, K); Bt. Raya, Nooteboom

4327 (BO, L); Central Kalimantan: Barito Ulu, P.T. Pamenang logging concession road,

km 20, Sidyasa PBU 650 (BO, K, L); East Kalimantan: Wanriset research area, road

Balikpapan to Samarinda, km 15, Sg. Wain area, Ambri & Arifin 354 (BO, L, K).

Rhaphidophora in Borneo

4. Rhaphidophora elliptica Ridl.

Rhaphidophora elliptica Ridl., J. Straits Branch Roy. Asiat. Soc. 44 (1905)

186; Engl. & K. Krause in Engl., Pflanzenr. 37 (1V.23B) (1908) 39—40;

Merrill, J. Straits Branch Roy. Asiat. Soc., special number (1921) 88. —

Type: Malaysia, Sarawak, 1” Div., Bau, July 1903, Ridley s.n. (SING, holo).

Figure 4

Large, occasionally very large, robust, pachycaul homeophyllous liane to

30 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 unbranched, leafy, non-flowering stems and long,

moderately elaborated, free, leafy, flowering stems later pendent under

their own weight; stems smooth, climbing stems broadly rectangular in

cross-section, the angles often slightly winged, the surfaces between convex,

free stems spreading, rectangular in cross-section, branching, growing to

considerable lengths, green, later mid-brown, without fibre at the tips of

active shoots, internodes to 16 x 1—2.5 cm on clinging 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,

clinging, pubescent; /eaves distichous and ascending on clinging and free

shoots; cataphylls and prophylls membranous, very soon drying and falling;

petiole deeply grooved adaxially, 10—21.5 x 0.2—1 cm, smooth, apical and

basal genicula weakly defined; petiolar sheath very prominent but soon

falling, extending to and encircling the apical geniculum, ligulate, very

soon 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;

lamina entire, elliptic to elliptic-lanceolate or falcate-oblong, weakly

channelled along midrib, 16—25 x 2.5—8 cm, coriaceous, upper surfaces

semi-glossy, lower surfaces semi-matt, base subacute to rounded or very

slightly cordate, apex subacute with a somewhat prominent apiculate

apicule, margins very slightly revolute in dried material; midrib raised

abaxially, slightly sunken adaxially; primary venation pinnate, slightly raised

abaxially and adaxially but barely visible in fresh material; interprimaries

parallel to primaries and only slightly less prominent, very slightly raised

abaxially and adaxially, + obscure in fresh material; 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 and one or more membranous, soon falling cataphylls; peduncle

36 Gard. Bull. Singapore 53 (2001)

Figure 4. Rhaphidophora elliptica Ridl.

A. flowering shoot x ’/, B. leaf lamina x '/,; C. venation detail x 3; D. portion of adult sterile

stem x '/,; E. inflorescence, spathe fallen x '/,; F. spadix detail, post-male anthesis x 4. A —C,E

—F from Mamit S$ 37665; D. from Kew LCD acc. no. 1965-49804.

Rhaphidophora in Borneo 37

compressed-cylindric, 4—11 x 0.3—0.5 cm; spathe canoe-shaped, stoutly

very short-beaked, 6.5—16 x 2—3.5 cm, thickly stiff-fleshy, dull yellow,

paler internally, soon falling at female anthesis; spadix tapering-cylindrical,

+ sessile, inserted level on peduncle, 8—14 x 1.5—2 cm; stylar region weakly

rhombohexagonal, 1.8—2.2 x 1.9—2.1 mm, + truncate; stigma punctiform,

raised, c. 0.3 mm diam.; anthers barely exserted at male anthesis, pollen

extruded from between ovaries; infructescence 10—15 x 2—2.5 cm.

Distribution: Endemic to Borneo. Sarawak (a and 3° Divisions), Brunei

Darussalam (Muara, Temburong), Sabah (Sandakan).

Habitat: On trees and cliffs in primary to disturbed primary lowland

dipterocarp and swamp forest, on a variety of substrates including limestone

and shale. Sea level to 115 m altitude.

Notes: 1. Boyce (1999) treated Rhaphidophora elliptica as a synonym of R.

montana, and comparison of the description above with that of R. montana

will reveal rather few morphological differences. However, more detailed

study of R. elliptica has demonstrated subtle but consistent differences

between it and R. montana such that I am now recognizing the Bornean

plants to be a distinct, endemic species. In particular, R. e/liptica is distinctive

in having the leaves distichous and ascending with the lamina shallowly

channelled along the midrib (leaves spiro-distichous and spreading with

the lamina flat in R. montana) and in having clinging stems considerably

wider then deep (c. 4:1 v. c. 2:1 in R. montana). These differences, so

obvious in the field, are almost completely obscured in herbarium specimens

unless, as is seldom the case, they are recorded by comprehensive field

data.

2. Rhaphidophora elliptica belongs to the taxonomically difficult the

Hongkongensis Group defined by clinging, occasionally free, stems square

to rectangular in cross-section, by simple, rather stiff leaves and conspicuous,

membranous ligulate petiolar sheaths extending to the apical geniculum

and soon falling to leave a horseshoe-shaped scar at the junction of the

petiole and lamina. The use of traditional herbarium morphology has proved

a woefully unsatisfactory method for delimiting species in this group and it

is hoped that the establishment of an alpha-taxonomy will provide a stimulus

for further study of the complex using macromolecular data, field

observations and morphometric techniques.

Other specimens seen: SARAWAK. 1 Div.: Along road between Kuching and Padawan,

10 miles SW of main Kuching - Serian highway Croat 53186 (MO); Kuching, Setapok F.R.,

38 Gard. Bull. Singapore 53 (2001)

Mamit S 37665 (K, KEP, L, US); One mile west of Bau, Nicolson 1293 (US); Setapok E.R.,

6 miles south of Kuching, Nicolson 1348 (US); Bau, Purseglove 4473 (K, L, SING); 3" Div.:

Kapit, Upper Batang Rejang, Clemens & Clemens 21929 (GH, K, MO). BRUNEI

DARUSSALAM. Muara: Hotta 13214 (KYO, L); Temburong: Batu Apoi F.R., ridgetop

west of Kuala Belalong Field Studies Centre, Poulsen 150 (AAU, BRUN, K). SABAH.

Sandakan: Sg. Lantoh, Lantoh SAN 87848 (K, KEP, L, SAN, SAR): Kinabatangan Road,

Batu Puteh, Pin-Supu F.R., Bt. Supu, Lim et al. LSP 766 (SAN, SING).

5. Rhaphidophora elliptifolia Merr.

Rhaphidophora elliptifolia Merrill, J. Straits Branch Roy. Asiat. Soc., special

number (1921) 88. — [Rhaphidophora elliptica Ridl., J. Straits Branch

Roy. Asiat. Soc. 49 (1908) 52, nom. illeg., non Ridl. (1905)]. — Type:

Malaysia, Sarawak, 1” Div., Kuching, Oct. 3 1905, Hewitt 12 (SING, holo).

‘Figure 5

Large, robust, pachycaul homeophyllous liane to 10 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 unbranched,

leafy, non-flowering stems and long, moderately elaborated, free, densely

leafy, flowering stems; stems smooth, climbing stems prominently

rectangular in cross-section, the surfaces slightly convex, free stems

spreading, irregularly four-angled in cross-section, sometimes irregularly

terete, with untidy prophyll, cataphyll and petiolar sheath fibre at the tips

of active shoots, internodes to 7 x 1.5 cm on clinging shoots, shorter and

less stout on free shoots, separated by large oblique to almost straight leaf

scars, older stems sub-woody; flagellate foraging stems not observed

(absent?); clasping roots densely arising from the nodes and internodes of

clinging stems, pubescent; feeding roots rare, clinging, pubescent; leaves

spiro-distichous on clinging and free shoots; cataphylls and prophylls

membranous, soon drying and degrading into netted and tangled fibres;

petiole deeply channelled adaxially, 10—13 x 0.3—0.4 cm, smooth, apical

geniculum well defined, basal genicula rather weakly defined; petiolar sheath

prominent, extending to and encircling the apical geniculum, briefly ligulate,

soon drying and degrading into netted fibres, eventually falling to leave a

continuous slight scar from the petiole base, around the top of the apical

geniculum and back to the base; /amina entire, narrowly elliptic to narrowly

elliptic-oblong, 20—29 x 4.5—7 cm, coriaceous, upper surfaces semi-glossy,

lower surfaces matt, base acute to decurrent, apex attenuate with a slight

apiculate apicule, margins revolute in dried material; midrib raised abaxially,

sunken adaxially; primary venation densely pinnate, raised abaxially and

adaxially; interprimaries parallel to primaries and barely less prominent,

Rhaphidophora in Borneo 39

Figure 5. Rhaphidophora elliptifolia Merr.

A. portion of adult sterile stem x ?/,: B. portion of older, leafless stem x '/;; C. leaf lamina x '/,;

D. venation detail x 4; E. inflorescence, spathe fallen x 1; F. spadix detail, post-female anthesis,

pre-male anthesis x 8. A & B from Meijer 2101; C — F from Hewitt s.n.

40 Gard. Bull. Singapore 53 (2001)

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 and netted

petiolar sheath, prophyll and cataphyll fibre; peduncle compressed-cylindric,

5.5 x 0.3 cm; spathe broadly canoe-shaped, stoutly short-beaked, 6—6.5 x

2—2.5 cm, thickly stiff-fleshy, dull yellow, persistent into early fruiting;

spadix cylindrical, + sessile, inserted level on peduncle, c. 5 x 1 cm; stylar

region rhombohexagonal, 1.7—2 x 1.7—2 mm, truncate; stigma rounded,

raised, c. 0.3 mm diam.; anthers not exserted at male anthesis; infructescence

c. 7 cm long (known from partial material only).

Distribution: Endemic to Borneo. Sarawak (1° Division), Kalimantan (East).

Habitat: Primary dipterocarp forest. Less than 100 m altitude.

Notes: 1. Rhaphidophora elliptifolia resembles Peninsular Malaysian/

Sumateran R. maingayi (see Boyce, 1999) in the persistent fibrous remains

at the tips of active stems and the stems rectangular in cross-section, but

differs in the smaller inflorescences (12—22 v. 6—6.5 cm), the spathe

persistent into fruiting and the narrower, more coriaceous leaves.

Rhaphidophora elliptifolia replaces R. maingayi in Borneo.

2. Confusion with R. elliptica is possible, although the smaller inflorescences

and persistent fibre at the shoot tips readily distinguish R. elliptifolia.

Other specimens seen: SARAWAK. No further data, Hewitt s.n. (SING). KALIMANTAN.

East Kalimantan, Pulau Nunukan, north of Tarakan, Meijer 2101 (L, BO).

6. Rhaphidophora foraminifera (Engl.) Engl.

Rhaphidophora foraminifera (Engl.) Engl., Pflanzenr. 37 (1V.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).

Figures 6 & 7

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 unbranched, densely leafy, non-flowering (always?) stems

and shorter clinging, densely leafy, flowering stems; stems smooth, mid-

{i IM

Yy, 1D,

\ Yy

Q\ Y V}, \\

. N Y,

: -)

foraminifera (Engl.) Engl.

'/,: B. pre-adult terrestrial shoot x '/;; C. leaf la

42 Gard. Bull. Singapore 53 (2001)

4. GuRra

Figure 7. Rhaphidophora foraminifera (Engl.) Engl.

A. flowering shoot x ’/,;; B. venation detail x 6; C. inflorescence, spathe fallen x */,; D. spadix

detail, male anthesis x 6.A,C & D from Boyce 722; B from Croat 53118.

Rhaphidophora in Borneo 43

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 by

prominent straight to slightly oblique leaf scars; flagellate foraging stem

absent; clasping roots densely arising from the nodes and internodes,

pubescent; feeding roots clinging, exceedingly robust, densely pubescent

with dense verticils of prominent corky ramenta; /eaves 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; /amina entire to slightly

or extensively perforated, perforations round to rhombic, extending c. + of

lamina width on each side of the midrib, ovate to oblong-lanceolate or

oblong-elliptic, slightly oblique, pubescent abaxially when young, 7—53 x

6—19 cm, 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 soon 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 male 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 mn, shortly conical when fresh, drying truncate; stigma

punctiform or elliptic and transversely orientated, raised at male anthesis

but excavated in dry material, c. 0.3 mm diam.; anthers slightly exserted at

male anthesis; infructescence 8—15 x 2—3 cm, dark green before ripening

to greenish yellow.

Distribution: Sumatera, Peninsular Malaysia, Borneo - Sarawak (ia and 6"

Divisions, but doubtless more widespread), Brunei Darussalam (Belait)

and Sabah (Pantai Barat).

44 Gard. Bull. Singapore 53 (2001)

Habitat: On trees, rocks and cliffs in primary to disturbed secondary lowland

to moist hill dipterocarp forest, on a variety of substrates including limestone

and shale. 10—700 m altitude.

Notes: 1. Confusion can occur between Rhaphidophora foraminifera and

R. puberula. They may be distinguished by the position of the inflorescence

(on short clinging shoots in R. foraminifera, on short free shoots in R.

puberula), by the presence of black mucilage produced by the deliquescing

cataphylls and prophylls (present in R. foraminifera, absent in R. puberula),

in leaf size of mature flowering-sized plants (to 53 x 19 cm in R. foraminifera,

to 34 x 10.5 cm in R. puberula), by the more or less rounded (R.

foraminifera) v. 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. versteegit)

suggesting that while a useful diagnostic tool, lamina perforation cannot be

used to circumscribe taxonomically meaningful groups within

Rhaphidophora.

Other Bornean specimens seen: SARAWAK. 1” Div.: Road between Kuching and Padawan,

10m SW of main Kuching to Serian highway, Croat 53182 (MO); 1 mile from Bau, Nicolson

1288 (US); 6" Div: Ulu Balingian, road from Sibu to Bintulu, Boyce 722 (K, M). BRUNEI

DARUSSALAM. Belait: Labi, Kp Teraja, path along the Sg. Teraja, Boyce 235 (BRUN,

K, L, MO). SABAH. Pantai Barat: Road from Kota Kinabalu to Tambunan, Ulu Moyog,

Boyce 1391, 1400 (K, SNP); Tambunan District, Road to Kg Tondulu, Boyce 14/4 (K,

SNP); Moyog District, along road between Kota Kinabalu and Tambunan, 21 miles SE of

Kota Kinabalu, W slopes of Crocker Range, Croat 53118 (MO); Kinabatangan, Tamegang

Timber Camp near Kg Pangkaian, Hotta 1416 (KYO, L).

7. Rhaphidophora korthalsii Schott

Rhaphidophora korthalsii Schott, Ann. Mus. Bot. Lugd.-Bat. 1(1863) 129;

Engl. in A. & C. DC., Monogr. Phan. 2 (1879) 246; Engl. & K. Krause in

Engl., Pflanzenr. 37 (1V.23B) (1908) 49—S1, Fig. 21; Alderw., Bull. Jard.

Bot. Buitenzorg III, 4 (1922) 341. — Type: Indonesia, Java, P.W. Korthals

sn. (eu, holen i. Piso):

Pothos celatocaulis N.E. Br., Gard. Chron. 13 (1880) 200. — Rhaphidophora

celatocaulis (N.E. Br.) Alderw., Bull. Jard. Bot. Buitenzorg HI, 1 (1920)

382 & Bull. Jard. Bot. Buitenzorg III, 4 (1922) 198. — Type: Malaysia,

Rhaphidophora in Borneo 45

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 (I1V.23B) (1908) 48—49; Merrill, J. Straits Branch Roy. Asiat.

Soc., special number (1921) 89. — 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

(1V.23B) (1908) 53; Merrill, J. Straits Branch Roy. Asiat. Soc., special

number (1921) 90. — Types: Malaysia, Sarawak, Beccari PB 1977 (FI lecto;

B isolecto; lecto selected by Boyce, 1999).

Rhaphidophora korthalsii Schott var. angustiloba Ridl. ex Engl. & K. Krause

in Engler, Pflanzenr. 37 (IV.23B) (1908) 49; Merrill, J. Straits Branch Roy.

Asiat. Soc., special number (1921) 89. — Type: Malaysia, Sarawak, Matang,

July 1903, Ridley s.n. (SING, lecto; lecto; selected by Boyce, 1999).

Rhaphidophora copelandii Engl., Bot. Jahrb. Syst. 37 (1905) 115; Engl. &

K. Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 49. — Type: Philippines,

Mindanao, Davao, Mt Apo, April 1904, Copeland 1193 (PNH, holoy; B

isO).

Rhaphidophora grandifolia K. Krause, Bot. Jahrb. 44, Beibl. 101 (1910) 11.

— Type: Philippines, Negros, Negros Orientale, Dumaguete (Cuernos

Mts), March 1908, Elmer 9464 (PNH, holot; B, E, K, L, LE, MO, iso).

Rhaphidophora trinervia Elmer, Leafl. Philipp. Bot. 8 (1919) 3073. — Type:

Philippines, Laguna, Los Banos (Mt Maquiling), June—July 1917, Elmer

18057 (PNH, holoy; FI, K, L, MO, P, iso).

Rhaphidophora ridleyi Merr., J. Straits Branch Roy. Asiat. Soc., special

number (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).

Rhaphidophora latifolia Alderw., Bull. Jard. Bot. Buitenzorg II, 4 (1922)

341; K. Krause & Alderw., Nova Guinea 14 (1924) 213. — Type: Indonesia,

Irian Jaya, Pionierbivak, 23 July 1920, Lam 711 (BO, holo; L, iso).

46 Gard. Bull. Singapore 53 (2001)

Rhaphidophora palawanensis Metr., Philipp. J. Sci. 26 (1925) 451. — Type:

Philippines, Palawan, Malampaya Bay, Oct. 1922, Merrill BS 11570 (PNH,

holo 5. Ke Paso):

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. inval., descr. Angl. — Based on: Philippines, Luzon, Sorsogon, Trosin

(Mt Bulusan), May 1916, Elmer 16061 (sheets seen at FI, K, L, MO, P,

PNH?*)].

Figures 8 & 9

Very large, occasionally enormous, slender to rather robust, pachycaul,

heterophyllous lane 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

unbranched, 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, clinging and free, very robust, densely

ramentose-scaly; /eaves distichous; cataphylls and prophylls membranous,

soon drying and degrading to intricately reticulate persistent fibres; 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

Rhaphidophora in Borneo

L.GUAR.

Sas = — : =

7 , SZ ———S ———S = Ss —— < y

Dy SSS SS

)

(

A. pre-adult shoot x '/,; B. pre-adult shingling shoot x '/,. A from Boyce 679; B from Nicolson

Figure 8. Rhaphidophora korthalsii Schott

1712.

48 Gard. Bull. Singapore 53 (2001)

venation strongly reticulate, slightly raised; tertiary venation invisible;

inflorescence solitary to several together, first inflorescence subtended by a

membranous prophyll and one or more cataphylls, these soon degrading

to netted fibres, subsequent inflorescences subtended by one or more soon

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—S cm, stiffly fleshy, greenish to dull yellow,

gaping wide at female anthesis and then soon 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, |.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 male

anthesis; infructescence 14—27 x 3—3.5 cm, dark green ripening to dull

orange, stylar tissue abscising to reveal orange ovary cavity pulp.

Distribution: Widespread in south tropical Asia from Sumatera and southern

Thailand to Borneo and the Philippines eastwards through the tropical

western Pacific. In Borneo, widespread in Sarawak, Brunei Darusslam and

Sabah. In Kalimantan only recorded from East Kalimanatan.

Habitat: Primary to disturbed secondary lowland, hill and montane forest,

moss forest, peatswamp forest, on trees, rocks and cliffs on a variety of

substrates including limestone and ultrabasic soils. 20—1800 m altitude.

Notes: 1. Rhaphidophora korthalsii is a very widespread and variable species,

hence the extensive synonymy. However, as with Epipremnum pinnatum

(L.) Engl. (Boyce, 1998) there are several geographical elements that, given

more intensive study, might warrant formal taxonomic recognition.

Unfortunately, current herbarium material is inadequate to confirm these

plants’ status and more field observations are needed.

2. Sterile herbarium material lacking the pre-adult stage may prove difficult

to distinguish from the Epipremnum pinnatum (in Borneo known wild

only from east Sabah). 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.

Rhaphidophora in Borneo 49

£.GuURR

Figure 9. Rhaphidophora korthalsii Schott

A. flowering shoot, leaves removed x '/,; B. leaf lamina x '/,; C. venation detail x 2; D.

inflorescence, spathe removed x 1; E. spadix detail, post-male anthesis x 8. All from Kerr

15051.

50 Gard. Bull. Singapore 53 (2001)

korthalsii is a shingle climber with oblong-elliptic to ovate, slightly falcate

upwardly 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 vy. trapezoid) and the

shape and orientation of the stigma (+ punctiform and circumferential v.

strongly linear and longitudinal) and, if fruits are mature, by seed characters.

The fruits of R. korthalsii each contain many small ellipsoid seeds with a

brittle, smooth testa, whereas E. pinnatum has fruits with two large, strongly

curved seeds with a bony and ornamented testa.

Other Bornean specimens seen: SARAWAK. 1” Div.: Bau, Brooke 8998 (BM); Lundu,

Foxworthy 40 (SING); Three miles from Kuching, Haviland & Hose 3605 (K); Kuching,

Hewitt 40 (SING); Semengoh F.R., 6 miles west of Kuching, Nicolson 1252 (US); 1 mile

west of Bau, Nicolson 1301 (US): Bako N.P., 20 miles northeast of Kuching, Lintang Path,

Nicolson 1329 (US); Setapok F.R., 6 miles west of Kuching, Nicolson 1341 (US); Kuching

Ridley s.n. (SING); 3" Div.: Hose Mountains, gorge of Sg. Simpurai, Burtt & Martin 4926

(E); Bt. Raya, Kapit, Soepadmo & Chai S$28148 (KLU, SAR); 4" Div.: Niah, Ahmed 64

(SAR, SING); Miri District, Niah, Sg. Sekaloh, G. Subis, Anderson $31956 (E, K, L, SAR,

SING, US); Kelabit Highlands, Apu Batu Buli, Nooteboom & Chai 2216 (L, SAR, US); a

Div.: Bintulu, Bt. Urang, Brunig S 12093 (K, SAR); Eastern ridge of Bt. Kans, Bintulu

district, Hirano & Hotta 1432 (KYO); Sg. Ma’au, Dataran Tinggi Merurong, Tubau, Othman

et al. S 49050 (K, SAR); 9" Div.: Tebedu, mile 15, Mohtar et al. S 49245 (K, SAR, US);

Serian, G. Penrissen; Paie S 16001 (K, LE, SAR). BRUNEI DARUSSALAM. Belait: Ulu

Ingei, Bt. Batu Patam, lower slopes near Sg. Ingei, Boyce 312 (BRUN, K); Temburong: Sg.

Temburong above Kuala Belalong, Argent et al. 9136 (E, K); Bt. Biang, Ashton 154 (K,

US); Sg. Temburong at Wong Nguan gorge, Wong 1727 (BRUN, K). SABAH. Kudat:

Ranau, Bt. Kulung, Meijer SAN 122414 (SAN); Ranau, Bt. Kulung, Meijer SAN 122429

(SAN); Ranau, Kinabalu road to Mesilau, Mikil SAN 38691 (K, SAN): Mesilau, Poore 131

(K); Sandakan: Paitan F.R., Ampuria SAN 32773 (K, SAN, SING); Elopura, mile 15 Labuk

Road, Cuadra A 2237 (K, KEP, SAN, SING); Sandakan, Segalid Lokan F.R., Gibot SAN

80977 (K, L, SAN); Lamag, Batu Puteh, Majawat & Lasan SAN 88017 (KEP, SAN); Pantai

Barat: Tambunan, road to Kg Tondulu, Kg Tondulu valley, Boyce 1421 (K, SING, SNP);

Papar, Kimanis road, Boyce 1443 (K, SNP); Dallas, Clemens & Clemens 26405 (BM, SING);

Dallas, Clemens & Clemens 26495 (BM, L, UC); Tenompok, Clemens & Clemens 28813

(BM, K, SING); Upper Kinabalu, Clemens & Clemens 29146 (BM, BO, K, L, SING, UC);

Upper Kinabalu, Lilau Basin, Mesilau, Meneringtan, Clemens & Clemens 29146a (BM, K,

L, SING); Penibukan, Clemens & Clemens 31150 (BM, K, SING); Penataran river, Clemens

& Clemens 34298 (BM, BO, K, L, UC); Mount Kinabalu N.P., along road to summit, Croat

53131 (MO); Lamag, Sg. Pin, Dewol & Harum SAN 89922 (K, SAN); Ulu Segama, Danum

Valley, path to Rhino Ridge, Dransfield 6302 (K); Pedalaman: Tenom, north ridge of G.

Malutut, c. 15 km north of Tenom, Kokawa & Hotta 2455 (KYO, L); Tawau: cutting area

of Luasong Camp, N.B.T. about 60 km NW of Tawau, Kokawa & Hotta 631 (K, KYO),

Kokawa & Hotta 797 (K, KYO, P); Lahad Datu, Ulu Sg. Sanum, south of camp 111, Stone

et al. SAN 85269 (KLU, SAN); Lahad Datu, Ulu Sg. Sanum, just north of camp 111, Stone

et al. SAN 85310 (KLU, SAN). KALIMANTAN. East Kalimantan: Wanariset research

area, road Samboja to Semoi, km 11, Ambri & Arifin 331 (L); Berau, Tandung Redeb, Sg.

Kelai near Long Lanuk, Kostermans 21125 (BO, L); Berau, near Tandung Redeb,

Rhaphidophora in Borneo 51

Kostermans 21725 (BO, L, US); Danan Pentulak, Bt. Pegah, Main (sub. Polak) 1957 (BO):

Pulau Nunukan, Tarakan, Meijer 2600 (BO): North of Tarakan, Simengkadu, Meijer 2389

(BO).

8. Rhaphidophora latevaginata M. Hotta

Rhaphidophora latevaginata M. Hotta, Acta Phytotax. Geobot. 22 (1966)

4. — Type: Malaysia, Sarawak, 4" Division, Bintulu District, about 4 km

east from Minah Camp, Sg. Kakus, 4 Oct. 1963, Hirano & Hotta 140 (KYO,

holo).

Figure 10

Moderate to very large, robust, pachycaul, homeophyllous neotenic liane

to 12 m; seedling stage a non-skototropic shingling juvenile shoot; pre-adult

plants forming small terrestrial colonies of shingling closely appressed leaves;

adult shoot architecture comprised of clinging, physiognomically unbranched,

shingling to very densely leafy, sterile stems and almost identical fertile

stems; stems weakly compressed-terete to weakly rectangular in cross-

section, smooth, dark green, without prophyll and cataphyll fibre but with

at least upper parts with thin, black adherent petiolar sheath tissue,

internodes to 12 x 2 cm, separated by prominent straight scars, but scars

obscured by leaf bases on all but the oldest stems, lower parts of stem later

sub-woody with slightly shiny cracking epidermis; flagellate foraging stems

moderately well developed although often somewhat short and leafy;

clasping roots arising densely from the nodes and internodes, prominently

scaly; feeding roots c. 3 mm diam., brown, minutely pubescent; Jeaves

distichous, appressed, ascending and shingling, becoming slightly scattered

and spreading (often litter-trapping) towards fertile tips: cataphylls and

prophylls membranous, soon drying black and persisting briefly before

falling; petiole deeply canaliculate and winged, 3—22 x 0.5—2 cm, smooth,

apical and basal genicula almost obscure in young leaves, later becoming

prominent, especially the basal geniculum; petiolar sheath very pronounced,

up to 2.5 cm wide, rather thickly membranous, prominently ligulate, outer

sheath (away from climbing surface) greatly expanded and partly to

completely obscuring stem, sheath turning black but persisting some

considerable time, eventually rotting away to reveal two large scars

extending to the top of the petiole; /Jamina broadly oblong-ovate-elliptic,

8—48 x 6.5—20.5 cm stiffly coriaceous, bright green, slightly to markedly

glaucous, base truncate-cordate to broadly cuneate, very briefly decurrent,

apex rounded to acute with a tiny apicule; midrib prominently raised

abaxially, slightly sunken adaxially: primary venation pinnate, slightly raised

yA Gard. Bull. Singapore 53 (2001)

abaxially, more so adaxially; interprimaries sub-parallel to primaries, slightly

raised on both leaf surfaces; secondary venation tessellate-reticulate, shghtly

raised abaxially, + flush adaxially, all veins much more prominent in dried

material; inflorescence solitary on a clinging shoot, subtended by a fully

developed foliage leaf and one or more cataphylls; peduncle laterally

compressed-cylindrical, 6.5—11 x 0.5—0.7 cm; spathe not observed; spadix

stoutly cigar-shaped, sessile; spadix inserted + level on stipe, 17.5 x 1.5 cm,

pale green; stylar region rhombohexagonal, c. 2 x 1 mm, truncate; stigma

slightly raised, elongated, longitudinally orientated, c. 0.75 x 0.2 mm; anthers

not exserted at male anthesis; infructescence stoutly cigar-shaped, 15 x 2

cm, stylar region becoming convex at fruit maturity.

Distribution: Endemic to Borneo. Sarawak (lf: and 7" Division), Brunei

Darussalam (Temburong), Sabah (Pantai Barat and Sandakan), Kalimantan

(localities not traced ).

Habitat: Primary to secondary moist lowland to hill dipterocarp forest on

clay and sandstone. 50—840 m altitude.

Notes: 1. Despite its remarkable appearance with densely glaucous-leafy

shingling shoots ascending, often spirally, high into the canopy, and

furthermore being widespread and not uncommon in Borneo, R. latevaginata

was described only relatively recently and, remarkably, is known from

only 13 herbarium specimens only two of which are more than 50 years

old. Given the plant’s striking appearance, it seems inconceivable that it

was not noticed by early fieldworkers although quite possibly its overall

similarity to the juvenile stages of the common and widespread R. korthalsii

may have led to it being passed by as not worth collecting.

2. It is not at all clear to what Rhaphidophora latevaginata is related. It is

the only large neotenic lianescent aroid in Borneo, although neoteny in

Rhaphidophora occurs in three groups (Boyce, 1999; Boyce & Bogner,

2000). The similarity of the plant to the juvenile stage of R. korthalsii

might seem compelling and the shared scaley feeding roots suggest a

relationship. However, the inflorescences are quite different; the style of

R. latevaginata is rhombohexagonal and the stigma longitudinally orientated

(v. style round and stigma + punctiform in R. korthalsii.)

Other specimens seen: SARAWAK. i Dive Matang F-.R., 10 miles west of Kuching, Nicolson

IZOZK(E O'S): 7” Div.: Sibu-Bintulu road, 2.3 km after bridge over the Batang Kemena,

Boyce 728 (K, M); Bintulu, about 2 - 4 km east from of Minah Camp, Sg. Kakus, Hirano &

Hotta 41 (KYO), Hirano & Hotta 149 (KYO); Vicinity of Minah Camp (timber camp of

Rhaphidophora in Borneo 58

<~}

yy)

pean

Figure 10. Rhaphidophora latevaginata M. Hotta

A. flowering shoot x */,; B. adult leaf lamina x '/,; C. venation detail x 3; D. inflorescence,

spathe fallen x ’/,; E. spadix detail, early fruiting x 4; F. pre-adult terrestrial shoot x '/;; G. pre-

adult terrestrial shoot x !/,;H. pre-adult terrestrial shoot x '/;.A,D & E from Hirano & Hotta

140,B & C from Dewol & Meijer 131653; F & G. from Hirano & Hotta 149; H from Hirano &

Hotta 41.

54 Gard. Bull. Singapore 53 (2001)

L.L.B.T.C.), Sg. Kakus, Hotta 15717 (KYO). BRUNEI DARUSSALAM. Temburong: Sg.

Temburong at Kuala Belalong, Boyce 402 (BRUN, K, L, SING, MO); Batu Apoi F.R.,

ridge west of Kuala Belalong Field Studies Center slope, between Danish Plot and Sg.

Belalong, Poulsen 165 (AAU, BRUN, K). SABAH. Pantai Barat: Road from Kota Kinabalu

to Tambunan, Ulu Moyog, Boyce 1389 (K, SNP); Sandakan: Phenological trail above sawmill,

Dewol & Meijer SAN 131653 (SAN); Kinabatangan District, Kg. Bilit, Bt. Bilit, Lim et al.

1272 (SING). KALIMANTAN. Locality not traced: Batu Makele, Batu Eilander, Raap 118

(BO); G. Dansu, Pentaclale, Pegah, Main (sub. Polak) 1958 (BO).

9, Rhaphidophora lobbii Schott

Rhaphidophora lobbii Schott, [Bonplandia 5(2) (1857) 45, nom. nud.], Prodr.

Syst. Aroid. (1860) 379; Engl. in A. & C. DC., Monogr. Phan. 2 (1879)

240; Engl. & K. Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 33—34;

Merrill, J. Straits Branch Roy. Asiat. Soc., special number (1921) 89. —

‘Scindapsus lobbii (Schott) Ender, Index Aroid. (1864) 74. — Type:

Singapore, Lobb 44 (K, holo).

Figure 11

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 unbranched, leafy, non-flowering

stems and long moderately elaborated, free, 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 clinging 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 clinging but often free, pubescent; /eaves weakly spiralled and

often sparsely arranged on clinging and proximal portions of free shoots,

densely spirally-distichous distally on flowering shoots; cataphylls and

prophylls membranous, very soon drying and falling; petiole grooved

adaxially, 4—9.5 x 0.2—0.3 cm, smooth, with a moderate apical and

prominent basal geniculum; petiolar sheath slightly prominent, extending

beyond the apical geniculum by two ligules, very soon 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; /amina entire, narrowly elliptic

Rhaphidophora in Borneo 55

“=

Figure 11. 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.

56 Gard. Bull. Singapore 53 (2001)

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 apicule; 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 soon falling cataphyll;

peduncle compressed-cylindric, 1.5—S5 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, soon falling at female anthesis 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 male anthesis, pollen extruded from between

ovaries; infructescence oblong-cylindric, 2.5—4 x 1—1.2 cm.

Distribution: Southern Thailand, Peninsular Malaysia, Singapore, Sumatera,

North Borneo and into Sulawesi.

Habitat: Primary to disturbed secondary lowland dipterocarp and

peatswamp forest on wet to inundated soils. Sea level to 250 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. In Borneo, confusion with R. cylindrosperma is possible although the

latter differs in its more coriaceous leaves, different leaf venation and in

having a longer, externally glabrous spathe.

3. Rhaphidophora lobbii is habitually found in wet to inundated ground,

an unusual habitat for a monsteroid liane, cf. Epipremnum amplissimum

Schott, and E. giganteum (Roxb.) Schott.

Other Bornean specimens seen: SARAWAK. 1 Div.: Setapok F.R., 4 miles south of Kuching,

a... - i ees

ake

Rhaphidophora in Borneo 57

Anderson § 23496 (SAR); Semengoh F.R., 6 miles west of Kuching. Nicolson 1258 (US):

Setapok F.R.. 6 miles south of Kuching. Nicolson 1359 (US): 2™ Div.: Simanggang. Triso

P.F., Anderson 14547 (SING): 8° Div.: Binatang. Pulau Bruit, Anderson 8032 (SING): Sg.

Kelepu, Pulau Bruit, Anderson 8007 (L, SING): Baram. Anderson S 3084 (SAR, SING).

BRUNEI DARUSSALAM. Temburong: Selapon, village margin, Dransfield 6917 (BRUN.

K. L); Batu Apoi F.R., on ridge between Kuala Belalong and Bt. Belalong. in permanent

Plot 2, Poulsen 362 (AAU, BRUN, K): Belait: Melilas, Ulu Ingei. path from Ulu Ingei to

Sg. Tupai, Sands 5919 (BRUN, K, L): Rasau, Van Niel 4250 (L. MO). SABAH. Sandakan:

Sepilok F.R.. Gambating SAN 94588 (K. L. SAN): Sg. Dagat. George et al. SAN 120521 (K.

SAN): Kinabatangan, Tamegang Timber Camp near Kg. Pangkaian. Kokawa & Hotta 1417

(KYO, L): Labuk Road, Meijer SAN 38784 (K, L. SAN): Below waterfall Kebun Cina F.R..,

Meijer & Dewol SAN 131749 (SAN): Bongaya F.R.. Kodoh & Aban SAN 82024 (K, L.

SAN, SAR. SING). KALIMANTAN. West Kalimantan: Pontianak, Polak 308 (BO): Sg.

Banghong. Polak 645 (BO, K): Bt. Raya. Tumbang Riang. Veldkamp 7910 (BO, L). East

Kalimantan: Wanariset. Balikpapan, Afriastini 116 (BO); Km 79 on road from Sangui Sg.

Mantaya. Kab. Kotawaringin Timur, Argent et al. 93138 (B, E): Berau, Tandung Redeb. Sg.

Kelai, near Long Lanuk. Kostermans 21133 (BO. L): Munukan, north of Tarakan, Meijer

2034 (BO): ‘Borneo’. Korthals 149 (L) & Korthals s.n. (L).

10. Rhaphidophora megasperma Engl.

Rhaphidophora megasperma Engl., Bot. Jahrb. Syst. 25 (1881) 8: Engl. &

K. Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 29, Fig. 9: Alderw., Bull.

Jard. Bot. Buitenzorg III, 1 (1920) 383; Merrill, J. Straits Branch Roy.

Asiat. Soc., special number (1921) 89. — Type: Indonesia, Kalimantan,

Central Kalimantan, Tumbang Hiang, 2 Sept. 1881, Grabowski 48 (B, holo).

Scindapsus havilandii Ridl., J. Straits Branch Roy. Asiat. Soc. 44 (1905)

184; Engl. & K. Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 75: Merrill,

J. Straits Branch Roy. Asiat. Soc.. special number (1921) 91, synon. nov. —

Type: Malaysia, Sarawak, 2™° Div.. Saribas, July 1892, Haviland 2089 (K,

holo: SING, iso).

Rhaphidophora jaculiformis Alderw., Bull. Jard. Bot. Buitenzorg III, 4

(1922) 197, synon. nov. — Type: Indonesia, Kalimantan, no further data,

1893, Jaheri s.n. (BO, holo; BO. iso).

Rhaphidophora subfalcata M. Hotta. Acta Phytotax. Geobot. 22 (1966) 6,

synon. noy. — Type: Malaysia. Sarawak, 7 Div., Bintulu, along Sg. Kakus

from Sg. Tinggili to Sg. Puteh, 13 Nov. 1963, Hirano & Hotta 1068 (KYO,

holo).

Figure 12

Moderate sized, slender to slightly robust semi-leptocaul homeophyllous

creeping and climbing liane to 5 m: seedling stage not observed: pre-adult

58 Gard. Bull. Singapore 53 (2001)

plants forming extensive terrestrial colonies; adult shoot architecture

comprised of greatly elongated, clinging, physiognomically unbranched,

sparsely leafy, non-flowering stems and short, little-branched, free, densely

leafy, flowering stems; stems smooth, terete in cross-section, internodes to

10 x 1.7 cm on clinging shoots, much shorter on free shoots, separated by

slightly oblique leaf scars, older stems subwoody; flagellate foraging stems

absent; clasping roots sparsely arising from the nodes and internodes of

clinging stems, pubescent; feeding roots rare, clinging, pubescent; leaves

spiro-distichous on clinging shoots, distichous on free shoots; cataphylls

and prophylls membranous, soon drying chartaceous and persisting at the

tips of flowering shoots, then falling; petiole grooved canaliculate, 11—29 x

0.2—0.4 cm, apical and basal geniculum moderately prominent; petiolar

sheath wide and membranous but very soon falling and thus usually not

prominent, extending almost to or reaching apical geniculum, very soon

‘drying and degrading into very sparse, soon-falling fibres; /amina entire to

slightly perforated, perforations round to rhombic, extending c. '/;—'/, of

lamina width on each side of the midrib, oblong-lanceolate or oblong-

elliptic, oblique, 12.5—42 x 3—10.5 cm, thinly coriaceous, base unequal,

rounded, weakly acute to subrounded, apex acuminate; midrib prominently

raised abaxially, + sunken adaxially; primary venation pinnate, slightly raised

abaxially and adaxially; interprimaries subparallel to primaries, much less

prominent, slightly raised abaxially and adaxially; secondary venation

prominently reticulate, raised abaxially, less so adaxially; inflorescence two,

three or more, together, each subtended by a prominent chartaceous

prophyll and one or more chartaceous cataphylls; peduncle slender to

somewhat stout, terete, 4—1 x 0.2—0.3 cm; spathe broadly canoe-shaped,

stout-beaked, 4—6 x 2—3.5 cm, stiff-fleshy, dull yellow on opening,

persistent into early fruiting, eventually falling to leave a prominent scar;

spadix stoutly to somewhat slender cylindrical, sessile, inserted obliquely

on peduncle, 2.5—4 x 0.5—0.7 cm, dull cream; stylar region mostly

hexagonal, 1.1—1.2 x 1—1.1 mm; stigma punctiform, very prominent, 0.1—

0.2 x c. 0.3 mm; anthers exserted at male anthesis; infructescence stoutly

oblong-cylindrical, 3.5—S x 1.2—1.5 cm.

Distribution: Endemic to Borneo (Sarawak ae. 3 7 candy Divisions)

and Kalimantan (West and Central).

Habitat: Primary to disturbed secondary lowland, often riverine, forest on

clay-loam. 40—130 m altitude.

Notes: 1. Distinctive in Borneo by the short leafy free side shoots bearing

clusters of inflorescences subtended and interspersed by prominent

Rhaphidophora in Borneo 59

N\A

——= SS \\\"\\

— Li\\\

———— PN

Se = |

SSS SSS = \

SS —S

—=

———— eS i

Figure 12. Rhaphidophora megasperma Engl.

A. flowering shoot x '/,; B. leaf lamina x '/,; C. venation detail x 2: D. leaf lamina 7 HES

inflorescence, spathe removed x 1'/,; E. spadix detail, post-male anthesis x 6; spadix detail,

early fruiting x 4. A & F from Ridley 14444: B—E from Burkill 6267.

60 Gard. Bull. Singapore 53 (2001)

chartaceous prophylls and cataphylls and by the spathe drying and persistent

into early fruiting. The clustered inflorescences subtended by chartaceous

cataphylls recall some New Guinea species, notably Rhaphidophora

versteegil.

2. Boyce (1999) merged R. megasperma with R. puberula. However, study

of further Bornean specimens of both taxa clearly shows that this was in

error and that R. megasperma is the earliest name for a Bornean endemic

hitherto called Scindapsus havilandii, Rhaphidophora jaculiformis and R.

subfalcata.

3. Hotta’s name R. subfalcata has been applied to large plants of R.

megasperma with perforated leaves. The presence of perforated and

unperforated leaves in individuals of same species is not uncommon in

Rhaphidophora; R. puberula is typical of such a phenomenon.

4. Both type sheets of Ridley’s Scindspaus havilandii are fertile; so it is odd

that Ridley should have considered it to belong to Scindapsus since the

generic diagnostic characters, number and position of ovules (solitary and

basal v. many on one or two parietal placentas), are readily observable.

5. The types of Aldrwerelt’s R. jaculiformis match R. megasperma perfectly.

Other specimens seen: SARAWAK. 1. Div.: Matang, Ridley s.n. (K); 3" Div.: Kapit, Sung,

upper Sg. Katibas, Sg. Joh trail, Zainudin et al. 4537 (K, UKMB). 7 Div.: Sg. Bejangung, a

branch of the Sg. Anap, Bintulu District, Hotta 15578 (KYO); 8 Div.: Bandar Sri Aman -

Sibu road, 156 km before Sibu below bridge over the Sg. Undup, Boyce 721 (K, M); Serian,

Kg. Tebekang, Othman & Munting S55097 (K, L, SAR). KALIMANTAN. West Kalimantan:

Serawai, 8 km NE of Desa Jelundung, Batu Lintang, next to camp, Church, Mahyer &

Afriastini 1668 (BO, GH, K); Serawai, Winkler 238 (E, L); Serawai, near Talok Nibung,

Winkler 1250 (E, L). Lake Tajan, Main (sub. Polak) 1758 (BO, L); Upper Samba river, 60-

80 km NNW of Tumbang Samba, Tumbang Habangoi to Tumbang Riang, Mogea 3550

(BO, K, KEP, L); Central Kalimantan: Bt. Raya, Nooteboom 4120 (L, BO):

11. Rhaphidophora minor Hook.f.

Rhaphidophora minor Hook.f., Fl. Brit. Ind. 6 (1893) 544; Engl. & K.

Krause in Engl., Pflanzenr. 37 (1V.23B) (1908) 21, Fig. 4; Merrill, J. Straits

Branch Roy. Asiat. Soc., special number (1921) 89. — 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

Rhaphidophora in Borneo 61

Toli-Toli, Jan. 1910, Schlechter 20698 (B, holo).

| Rhaphidophora palawanensis Furtado, nom. nud. in sched. var., non. Merr.

(1925)}

Figure 13

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

unbranched, leafy, non-flowering stems and long moderately elaborated,

free, 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 clinging 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 clinging and proximal portions of free shoots, moderately densely

distichous distally on flowering shoots; cataphylls and prophylls

membranous, very soon 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 soon 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; /amina 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 apicule; midrib raised abaxially, slightly raised

adaxially; primary venation pinnate, slightly raised on both surfaces

prominent (raised) in dried material; 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 soon 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, soon falling at female anthesis 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

62 Gard. Bull. Singapore 53 (2001)

rhombohexagonal, 1.4—2 x c. 2 mm, truncate; stigma punctiform, c. 0.3

mm diam., slightly prominent in dried material; anthers well-exserted at

male anthesis; infructescence oblong-cylindric, 4.5—7 x 1—2.5 cm.

Distribution: Southern Thailand, Peninsular Malaysia, Singapore, Sumatera,

throughout Borneo (widely scattered), Sulawesi and into the Philippines

(Mindanao, Palawan).

Habitat: Lowland peatswamp forest, riverine forest, rear of freshwater

mangrove. Sea level to 10 m altitude.

Notes: 1. In the fresh state confusion with Rhaphidophora sylvestris is

possible, although the thinner, more prominently veined leaf and longer

spathe beak of R. minor Hook.f. are diagnostic. Dried material of R. minor

is notable for the uniformly pale straw coloured leaves.

2. In Borneo, R. minor is habitually found in swampy habitats whereas

elsewhere in its range it tends to be in drier places.

Other Bornean specimens seen: BRUNEI DARUSSALAM. Belait: Kuala Belait, Van Niel

3944 (L); Seria, Van Niel 4138 (L); Sg. Belait, Van Niel 4634 (L). SARAWAK. L Div.:

Kuching, Hewitt s.n. (K); Kuching, Ridley s.n. (SING); Kuching, Ridley s.n. (K). 7” Div.:

Ulu Bawan, Balingian, Ashton S 19585 (GH, K, L, MO, SAR, SING, US); Sg. Tengah,

Brooks 1016 (BM); Batang Lassa, near Rhuma Jumbau, Mamit S 33623 (K, L, MO, SAR,

US). SABAH. Sandakan: Beluran, Sg. Mangkayok, near Sg. Kalagan, Meijer SAN 51617

(SAN); Ulu Dusum, Meijer SAN 122736 (E, SAN, US). KALIMANTAN. West Kalimantan:

Mempaura, Teysmann 8105 (BO); Central Kalimantan: Sampit, Buwalda 7917 (BO); East

Kalimantan, 20 km north of Wanariset, Kg. Padang, Samboja, Sidiyasa 1111 (K, L); Cape

Genderen, Pladjoe, Amdjah 7 (BO, K, L, SING), Amdjah 32 (BO, L), Amdjah 951 (BO, K,

L); G. Samenggaris, Amdjah 1084 (BO, K, L); South Kalimantan: no further data, Winkler

3448 (L): Locality not traced: Hallier 35, 66 (BO); Long Lansiah (?), Teysmann 11338 (BO, L);

12. 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

(IV.23B) (1908) 30. — Type: Indonesia, Sumatera, West 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 (I1V.23B) (1908) 23. — Type: Malaysia,

Perak, Scortechini 347 (K, holo; CAL, iso).

Rhaphidophora kunstleri Hook.f., Fl. Brit. Ind. 6 (1893) 546; 548; Engl. &

Rhaphidophora in Borneo 63

Figure 13. Rhaphidophora minor Hook.f.

A. portion of adult, sterile shoot x '/,; B. flowering shoot x '/,; C. leaf lamina x */;; D. venation

detail x 2; E. inflorescence, spathe just beginning to abscise x 1'/,; F. spadix detail, post-female

anthesis, pre-male anthesis x 8; G. spadix detail, early fruiting x 5. A & D from Winkler 238; B

& from Church et al. 1668; E & F from Hotta 1068.

64 Gard. Bull. Singapore 53 (2001)

K. Krause in Engl., Pflanzenr. 37 (1V.23B) (1908) 30; Alderw., Bull. Jard.

Bot. Buitenzorg III, 4 (1922) 338. — Type: Malaysia, Perak, Bt. Larut, July

1883, Kunstler 4538 (K, lecto, selected by Boyce, 1999).

Rhaphidophora gracilipes Hook.f., Fl. Brit. Ind. 6 (1893) 545; Engl. & K.

Krause in Engl., Pflanzenr. 37 (1V.23B) (1908) 30, Fig. 10; Alderw., Bull.

Jard. Bot. Buitenzorg III, 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 (1V.23B)

(1908) 27. — Type: Indonesia, Sumatera, North Sumatera, Kepulauan Batu,

25 Sept. 1896, Raap 370, (B, lecto; BO, isolecto, selected by Boyce, 1999).

-Rhaphidophora hallieri Alderw., Bull. Jard. Bot. Buitenzorg II, 1 (1920)

385; Alderw., Bull. Jard. Bot. Buitenzorg III, 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 II, 4 (1922) 196. — Type: Cult.

Bogor Bot. Gard, June 1919, Alderwerelt s.n. (BO, holo).

Figure 14

Moderate to large, rather robust, semipachycaul 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 unbranched, leafy, non-flowering stems and short, usually

unbranched, free, 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 clinging 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, clinging, pubescent; /eaves distichous on clinging and

free shoots; cataphylls and prophylls membranous, soon 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, soon drying and

degrading into sparse, soon-falling fibres; /Jamina entire to slightly

perforated, perforations round to rhombic, extending c. '/;—'/, of lamina

Rhaphidophora in Borneo 65

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, fragrant of frangipani

and pineapple (fide Hay et al. 9029) subtended by a fully developed foliage

leaf and, if more than one inflorescence, then second preceded by a large

cataphyll; peduncle strongly compressed-cylindric, 4—9 x 0.4—0.6 cm; spathe

canoe-shaped, stoutly long-beaked, 5.5—19 x 1—6.5 cm, stiff-fleshy, dull

green to dull yellow, pale yellow to apricot-orange internally, falling (after?)

female anthesis 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 male 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 lower/

upper montane transition forest, often on steep slopes, on granite and

limestone. 90—1550 m altitude.

Notes: 1. Given the overall uniformity of this species, it is quite extraordinary

that Rhaphidophora puberula should have been redescribed no fewer than

seven times based mostly on minor differences of leaf shape.

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 Jess perforated than those of R.

foraminifera.

66 Gard. Bull. Singapore 53 (2001)

Other Bornean specimens seen: SARAWAK. 1” Div.: Matang, Ridley s.n. (K); 4" Div.:

Baram District, G. Mulu N.P., Hidden Valley, Argent et al. 887 (L); 7” Div.: Bintulu, Segan

F.R., Chai $31999 (K, L, SAN, SAR, SING, US). BRUNEI DARUSSALAM. Belait: Labi,

track to Rampayoh waterfall, Ahmad 15 (BRUN, K). SABAH. Sandakan: Bettohan near

Sandakan, Kloss SFN 19099 (K, L, SING); Lamag, Kinabatangan, Ulu Sg. Pin, Madani &

Saigol SAN 90180 (K, L, SAN, SAR); Pantai Barat: Keningau, Sg. Labou, Mantor SAN

110980 (GH, K, SAN); Tuaran District, below summit of Lawa Mandau, Telekom station

east of Telipok, Meijer et al. SAN 136186 (GH, SAN); Pedalaman: Kiau, Mount Kinabalu,

Clemens 10142 (UC); Gurulau Spur, Mount Kinabalu Clemens 10772 (UC); Dallas, Clemens

& Clemens 26453 (BM, K, L, SING); Dallas, Clemens & Clemens 26781 (BM); Dallas,

Clemens & Clemens 26876 (BM, BO, GH, K, L, SING, UC); Tenompok, Clemens &

Clemens 26900 (BM, BO, K, L, SING, UC); Kinabalu N.P., along the road between Park

H.Q. and Tenompok, Kokawa & Hotta 3089 (KYO); Tawau: Elmer 21593 (BO, K, L, MO,

SING, UC); Sabah Foundation logging area Umas-Umas, Krispinus SAN 87213 (K, KEP,

L, SAN, SAR); Kalabakan, km 41 Imbak road, Luasong, Krispinus SAN 95680 (K, L, SAN,

SAR); Lahad Datu, Madai Caves, Madai Baturong F.R., Kokawa & Hotta 1097 (KYO, L).

Locality not traced: Pandawan, Sg. Mesopo, Fidilis & Sumbing SAN 113389 (K, KEP,

SAN); KALIMANTAN. Cult. Bogor, Y70, from Kalimantan collection, Ashton 12626 (BM,

BO) & Nicolson 942 (BO, L, US).

13. Rhaphidophora sylvestris (Blume) Engl.

Rhaphidophora sylvestris (Blume) Engl. in A. & C., DC, Monogr. Phan. 2

(1879) 239 & in Beccari, Malesia 1 (1882) 268; Engl. & K. Krause in Engl.,

Pflanzenr. 37 (IV.23B) (1908) 22—25, Fig. 6 (‘silvestris’); Alderw., Bull.

Jard. Bot. Buitenzorg III, 1 (1920) 383; Merrill, J. Straits Branch Roy.

Asiat. Soc., special number (1921) 90 (‘silvestris’). — 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).

Scindapsus aruensis Engl., Bull. Soc. Tosc. Ortic. 4 (1879) 270. — Type:

Indonesia, Melaku, Aru Islands, Gabu-lengaw, May 1873, Beccari s.n. (FI,

holo).

Rhaphidophora in Borneo 67

cs aa

CxS

See

% cr

‘eB

Figure 14. Rhaphidophora puberula Engl.

A. flowering shoot x '/,; B. leaf lamina x '/,; C. venation detail x 6; D. inflorescence, spathe

removed x 2; E. spadix detail, post-male anthesis x 6; F. spadix detail, early fruiting x 4.A & F

from Ridley 14444, B—E from Burkill 6267.

68 Gard. Bull. Singapore 53 (2001)

Rhaphidophora wrayi Hook.f., Fl. Brit. India 6 (1893) 544; 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 var. obtusata Engl., Malesia 1 (1883) 268 — Type:

Malaysia, Sarawak, 1 Div., Kuching, Nov.1865, Beccari PB 952 (FI, holo;

FI spirit 423, K, iso).

Rhaphidophora nigrescens Ridl., J. Straits Branch Roy. Asiat. Soc. 44 (1905)

185 (‘migrescens’), synon. noy. — Type: Malaysia, Sarawak, 1" Div., Matang,

July 1903, Ridley s.n. (SING, lecto, selected here). Ridley cites two

conspecific syntypes, that chosen is the best specimen for identification

purposes. The other syntype is: Malaysia, Sarawak, 19 June 1893, Bartlett

s.n. (SING).

Rhaphidophora motleyana Engl. & K. Krause in Engler, Pflanzenr. 37

(I1V.23B) (1908) 25; Merrill, J. Straits Branch Roy. Asiat. Soc., special

number (1921) 89. — Type: Indonesia, Kalimantan, South Kalimantan,

Bangarmassin, 1857—1858, Motley 741 (K, holo).

[Scindapsus lanceolataus Mig. nom. nud. in sched. BO]

|Pothos cucculata Zipp. nom. nud. in sched. L]

Figure 15

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 unbranched,

leafy, non-flowering stems and long, moderately elaborated, free, 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 .S—1 cm on clinging 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 i in cross-section; clasping roots densely

Rhaphidophora in Borneo 69

SOE

BAIS

4 .GUAR.

Figure 15. 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-male

anthesis x 4.A & B from Backer 11199; C, F & G. from de Wilde & de Wilde-Duyfjes 13830; D

& E from Afriastini 2399.

70 Gard. Bull. Singapore 53 (2001)

arising from the nodes and internodes of clinging stems, pubescent; feeding

roots very rare, clinging, pubescent; /eaves distichous on clinging and free

shoots, those distal on flowering shoots densely so; cataphylls and prophylls

membranous, very soon 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 soon 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 apicule; midrib slightly raised abaxially, slightly sunken

_adaxially; primary venation pinnate, slightly raised abaxially and adaxially;

interprimaries parallel to primaries and only slightly less prominent, very

slightly raised abaxially and adaxially; secondary and tertiary venation +

obscure in fresh material, visible as a faint reticulum in dried specimens;

inflorescence solitary, subtended by a fully developed foliage 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, soon falling at female anthesis; spadix cylindric to 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 male anthesis;

infructescence 6—8 x 2—2.5 cm.

Distribution: Peninsular Malaysia, Sumatera, throughout Borneo, Java, Nusa

Tenggara and Maluku.

Habitat: Primary to disturbed secondary lowland to lower montane forest

on a variety of substrates including sandy loam and limestone. 120—1800

m altitude.

Note: Rhaphidophora sylvestris is more variable in Borneo than in other

parts of its range (e.g. Peninsular Malaysia, Java) but the paucity of

collections coupled with more or less continuous variation makes it

impossible to be confident that more than one species is involved. Of

particular note is the occurrence of plants with coriaceous leaf laminas that

have variously been described as Rhaphidophora gratissima Becc. (syn. R.

sylvestris var. obtusata Engl.) and R. nigrescens Ridl.. Such plants approach

R. conocephala in appearance (and also resemble non-Bornean R.

crassifolia), although the flat-topped styles readily distinguish them from

Rhaphidophora in Borneo v4

R. conocephala, while the juvenile shoots without conspicuously twisted

stems separate them from R. crassifolia. Lamina thickness is not a consistent

character, even between duplicates of the same collection.

Other Bornean specimens seen: SARAWAK. 1" Div.: G. Pueh (Mt Poi), base camp, Clemens

& Clemens 21935 (K); Matang, Kochummen FRI 29145 (KEP, GH, K, L, SING); Kuching,

Ridley s.n. (K); Puak, Ridley 12413 (BM, K, SING); Kuching, Tiang Bekap, G. Maja, W.L.

Chew 714 (L, SING). SABAH. Pantai Barat: Ranau, Bt. Hampuan, Amin & Jarius SAN

121155 (K, SAN); Mt Kinabalu, Bungal trail, Clemens & Clemens s.n. (SING); Dallas,

Clemens & Clemens 26720 (K, SING), Clemens & Clemens 26921 (BM, BO, K, L, SING),

Clemens & Clemens 27030 (BM, BO, K, L, SING, US), Clemens & Clemens 27312 (BM,

BO, K, SING); Upper Kinabalu, Mementong, Mesilau Basin, Clemens & Clemens 28487

(BM, K, L, SING); Tenompok, Clemens & Clemens 29227 (BM, BO, K, L, SING, UC);

Ranau, Mt Kinabalu, Mesilau valley, Cockburn SAN 70108 (SAN); Kinabalu N.P., along

Sg. Mesilau from Mesilau Camp to Mesilau Cave, Kokawa & Hotta 3995 (KYO); Kinabalu,

Penibukan, Sg. Tahubang, Nooteboom & Aban 1582 (L). BRUNEI DARUSSALAM. Belait:

Sg. Keduan, off Sg. Belait, Forman 1174 (BRUN, K). KALIMANTAN. Central Kalimantan:

Sintang, HPH km 70, west camp off main (new) logging road, Church et al. 985 (BO, GH,

K), ‘Borneo’, Korthals 153, Korthals s.n. (L).

Acknowledgements

Thanks are due to Linda Gurr for skillfully executing the illustrations that

accompany this article. Thanks also to Dr Alistair Hay (NSW) who critically

reviewed the manuscript and offered much useful advice.

References

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. 2000a. The genus Rhaphidophora Hassk. (Araceae-

Monsteroideae-Monstereae) in the Southern and Western Indonesian

Archipelago. Gardens Bulletin Singapore. 52: 101—183.

Boyce, P.C. 2000b. The genus Rhaphidophora Hassk. (Araceae-

Monsteroideae-Monstereae) in the Philippines. Gardens Bulletin

Singapore. 53: 213—256.

12 Gard. Bull. Singapore 53 (2001)

Boyce, P.C. and J. Bogner. 2000. An account of neotenic species of

Rhaphidophora Hassk. (Araceae-Monsteroideae-Monstereae) in New

Guinea and Australia. Gardens Bulletin Singapore. 52: 89—100.

Boyce, P.C. 2001. The genus Rhaphidophora Hassk. (Araceae-

Monsteroideae- Monstereae) in New Guinea, Australia and the Tropical

Western Pacific. Gardens Bulletin Singapore. 53: 75—183.

Engler, A. and K. Krause 1908. Araceae-Monsteroideae. In A. Engler (ed.),

Das Pflanzenreich 37 (1V.23B) 4—139. Engelman, Leipzig.

Mayo, S.J., J. Bogner and P.C. Boyce. 1997. Genera of Araceae. Royal

Botanic Gardens, Kew, U.K.

Schimper, A.F.W. 1903. Guilds in Plant-Geography upon a Physiological

Basis. Pt 2, ch. 2: 192—206. (English translation by W.R. Fisher, revised

and edited by P. Groom & I.B. Balfour.) Clarendon Press, Oxford,

WK.

Stearn, W.T. 1992. Botanical Latin, 4" ed., pp. xiv + 546. David & Charles,

Newton Abbot & London, U.K.

Strong, D.R. & Ray, T.S. 1975. Host tree location behavior of a tropical

vine (Monstera gigantea) by skototropism. Science. 190: 804—806.

Index Exsiccatorum

beccarii = 1 latevaginata = 8

conocephala = 2 lobbii = 9

cylindrosperma = 3 megasperma = 10

elliptica = 4 minor = 11

elliptifolia = 5 puberula = 12

foraminifera = 6 sylvestris = 13

korthalsii = 7

Aban & Petrus SAN 90675 = 1; Afriastini 116 = 9; Ahmad 15 = 12, 64 = 7;

Ambri & Arifin 331 = 7, 354 = 3; Amdjah 7, 32, 951, 1084 = 11; Amin &

Jarius SAN 121155 = 13; Amin et al. SAN 94681, SAN 97483 = 1; Ampuria

SAN 32773 = 7; Anderson S 3084, 8007, 8032, 14547, S 23496 = 9; S 31956 =

7; Argent et al. 93-138 = 9; 887 = 12; 9136 = 7: Ashton 154 = 7: 12626 = 12: S

19585 = 11

Beccari PB 833 = 1, PB 952 = 13; PB 1977, PB 2314, PB 2714 =; Bogner

1346, 1393, 1507, 1504, 1562 = 1; Boyce 235 = 6; 245 = 1; 312 = 7; 345 = 3;

Rhaphidophora in Borneo 73

396 = 1; 402 = 8; 719 = 1; 721 = 10; 722 = 6; 728, 1389 = 8; 1391, 1400, 1414

= 6; 1421, 1443 = 7; Brooke 8409, 9459 = 1; 8998 = 7; 10729 = 1; Brooks

1016 = 11; Brunig S 12093 = 7; Burbidge sub Veitch no. 215 = 7; Burtt

12938, 13001 = 1: Burtt & Martin 4926 = 7; Buwalda 7917 = 11

Chai S 31999 = 12: S. 39439 = 1; Chew 714 = 13; Church et al. 985 = 13:

1573 = 3; 1668 = 10; Clemens 10142, 10772 = 12; Clemens & Clemens 21929

eee pose. ie 219395 = 43° 26405 = 7; 26453 = 12; 26495 = 7; 26720 = 13;

26781, 26876, 26900 = 12; 26921, 27030, 27312, 28487 = 13; 28813, 29146,

29146a = 7; 29227 = 13; 31150, 34298 = 7; Cockburn SAN 70108 = 13: Croat

Silla — 6; 33131 = 7; 53182 = 6: 53186 = 4; Cuadra A 2237 =7

Dewol & Harum SAN 89922 = 7; Dewol & Meijer SAN 131653 = 8; Dewol

et al. SAN 118075 = 1; J.Dransfield 6302= 7; 6917 =9

Elmer 21593 = 12; Endert 1860, 3013 =1

Fidilis & Sumbing SAN 113389 = 12; Forman 1174 = 13; Foxworthy 40 = 7

Gambating SAN 94588 = 9; George et al. SAN 120521 = 9; Gibot SAN

80977 = 7: SAN 90040 = 1; Grabowski 48 = 10

Hallier 35, 66 = 11; 999 = 1; 1164 = 3; 1187 = 12; 1555 = 1: Haviland & Hose

3605 = 7: Hewitt 12 = 5: 40 = 7; Hirano & Hotta 41, 140, 149 = 8: 1068 = 10:

Oi — 46-1252 —7- 17082 = 1: Hotta 1716 = 6; 13214 = 4; 15578 = 10: 15717

= §

Jimpin SAN 122024 =1

Kloss SFN 19099 = 12; Kochummen FRI 29145 = 13: Kodoh & Aban SAN

82024 = 9: Kokawa & Hotta 110, 471 = 1. 631, 797 = 7; 1097 = 12; 1417 = 9:

2455 = 7; 3089 = 12; 3995 = 13; Korthals 149 = 9; 153 = 13; Kostermans

eieZy — 7; 21133 = 9; 21725 = 7; Krispinus SAN 87213, SAN 95680 = 12;

SAN 118444 =1

Lantoh SAN 87848 = 4; SAN 102053 = 1; Lim et al. LSP 766 = 4; 1272 = 8

Madani & Saigol SAN 90180 = 12; Main (sub. Polak) 1758 = 10; 1957 = 7;

1958 = 8; 2092 = 1; Majawat & Lasan SAN 88017 = 7; Mamit S 33623 = 11;

S 37665 = 4; Mantor SAN 110980 = 12; Meijer 1136 = 1; 2034 = 9; 2389 = 7;

2101 = 5; 2600 = 7, SAN 38784 = 9; SAN 44014 = 1; SAN 51617 = 11; SAN

74 Gard. Bull. Singapore 53 (2001)

122414, SAN 122429 = 7; Meijer SAN 122736 = 11; Meijer & Dewol SAN

131749 = 9; Meijer et al. SAN 136186 = 12; Mikil SAN 38691 = 7; Mogea =

3550 = 10; Mohtar et al. S 49245 ='7; Motley 741 = 13

Native collector D34 = 12; 966 = 9; Nicolson 942 = 12; 1252 = 7; 1258 = 9.

262871282 =1- 1288 3621293 = 41301 1329. 14 sas ea

= 4; 1359 = 9: Nieuwenhuis 97, 219 = 1; Nooteboom 4327 = 3; Nooteboom

4120 = 10; Nooteboom & Aban 1582 = 13; Nooteboom & Chai 2216 = 7

Othman & Munting S 55097 = 10; Othman et al. S 49050 = 7

Paie S 16001 = 7; Polak 308, 645 = 9; Poore 131 = 7; Posthumus 2173 = 1;

Poulsen 150 = 4; 165 = 8; 362 = 9; Purseglove 4473 = 4

Raap 118 = 8; Ridley 2089 = 10; 12413 = 13; 12414 =7

Sands 5919 = 9; Sidyasa PBU 650 = 3; 1111 = 11; Soepadmo & Chai S

28148 = 7; Stone et al. SAN 85269, SAN 85310 =7

Tangki SAN 119596 = 1; Teysmann 8105, 11338 = 11

Van Niel 3944, 4138 = 11; 4250 = 9; 4634 = 11; Veldkamp 7910 = 9

Winkler 238, 1250 = 10; 3448 = 11; Wiriadinata 850 = 2; Wong 242 = 1; 1727

Vig S 51303 =A

Zainudin et al. 4537 = 10

Gardens’ Bulletin Singapore 53 (2001) 75-183.

The Genus Rhaphidophora Hassk. (Araceae-

Monsteroideae- Monstereae) in New Guinea, Australia

and the Tropical Western Pacific

P.C. Boyce

Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, U.K.

Abstract

An alpha-taxonomic account of Rhaphidophora in New Guinea, Australia and the tropical

western Pacific is presented as a precursor to the forthcoming Flora Malesiana Araceae

treatment. Thirty species are recognized, of which 12 (R. cravenschoddeana, R. cryptantha,

R. fortis, R. gorokensis. R. guamensis, R. intonsa, R. intrusa, R. jubata, R. kokodensis, R.

mima, R. pilosa and R. waria) are novel. Twenty new synonomies are made: R. hollrungii

and R. iboensis = R. australasica; R. pallidivenia = R. conica; R. oreophila = R. discolor, R.

wentii = R. geniculata; R. nutans = R. microspadix; R. buergersii, R. conferta, R. drepanophylla,

R. obliquata and R. peekelii = R. schlechteri; R. apiculata; R. engleri [syn. R. palauensis

Engl. & K. Krause (1921) non Koidz. (1916)], R. forbesii, R. kanehirae and R. palauensis

Koidz. (1916) = R. spathacea; R. graeffei and R. reineckei = R. spuria; R. ledermannii = R.

versteegii. One species (R. dahlii Engl.) is treated as doubtful. A dichotomous key to

species is provided. All species are illustrated.

Contents

VLE LCS OE ae ee 2 ee ee 75

Mat MO UR EE EIEN E SIRI 000202) 8 SE 86 23 ccc eseallcce cnccznesesch desaescdcetelsence 76

Sea PAIR ALC CLC OMMIS IN cohen 2 oon: nace Sacre cd ohende-cocpodcontecadsscdedeencnssesteeasedeaeieas= 78

NaN PMREEELONPINTLS EOD Ree he ee op See re ei 5 Na Hagined soso naga vscebioa neat eb vieeictonagivdesdscs 78

Key to adult flowering Rhaphidophora in New Guinea,

Pema Alien Alia) Be TLOPICAl WEStCEE PACHIG — 2 oe. cie.ncctsassusseonscassceseesest 82

LE SUS SSS eg LE Eo aise en 88

aS STAVES ee ee ee 179

ULE LENSD FEL I Ce pe ee 179

Pati sel Sead Coc CVA REA Po) ec ee ea) 8D. Stipes oa deat ees tan sbeceseoanteds 180

I SA lh aca ou 2 canner ne vnsScakdsndcvavnennens dt 183

Introduction

Rhaphidophora Hassk. (including Afrorhaphidophora Engl.; at least 3

species in tropical Africa) comprises c. 100 species of small to large,

occasionally enormous, root-climbing lianes (sensu Schimper, 1903), rarely

76 Gard. Bull. Singapore 53 (2001)

rheophytes, distributed from tropical West Africa eastwards to the western

Pacific (Samoa), 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 Himalaya

(SE Nepal to NE Vietnam, roughly 17°—23° N), West Malesia (including

southernmost peninsular Thailand), the Philippines, and East Malesia.

Thirty species occur in New Guinea, Australia and the tropical Western

Pacific region.

The last complete revision of Rhaphidophora was that of Engler and

Krause (1908) and a summary of the taxonomic and nomenclatural history

of Rhaphidophora was presented in Boyce (1999).

This is the sixth in a series of papers intended to present a complete

alpha-taxonomy of the genus Rhaphidophora. Accounts for Peninsular

Malaysia and Singapore (Boyce, 1999), south and west Indonesia (Boyce,

2000a), the Philippines (Boyce, 2000b), Borneo (Boyce, 2001) and for

neotenic species in New Guinea and Australia (Boyce & Bogner, 2000)

have been published to date. Accounts for each of India, Sri Lanka and

the Himalaya, Thailand and Indochina, and Africa are being prepared and

will be published separately.

Synonymy cited is for the species, not for the review area. The reason

for this is that synonymous names based on types from outside the review

area are frequently applied to specimens in local herbaria.

Infrageneric Relationships

Boyce (2000a) proposed eight informal morpho-taxonomic groups for the

Rhaphidophora of Indochina and Thailand, Sunda and Wallacea, each

group named with the earliest accepted species epithet belonging to that

group. It should be noted that these groups are a first, pragmatic attempt

at creating morpho-taxonomic units. Molecular work underway at

Cambridge will result in a better, infrageneric classification.

Of the eight groups proposed by Boyce (2000a), three occur in the

New Guinea and Pacific region:

1. Korthalsii Group (R. korthalsii Schott).

2. Neoguineensis Group (R. neoguineensis Engl., R. intrusa P.C. Boyce,

R. mima P.C. Boyce, R. schlechteri K. Krause).

3. Spathacea (Hollrungii) Group (R. australasica F.M. Bailey, R.

guamensis P.C. Boyce, R. spathacea Schott, R. versteegii Engl. & K.

Krause, R. waria P.C. Boyce). The Hollrungii Group is here renamed

the Spathacea Group, following recognition of R. spathacea as the

earliest described attributable species.

Rhaphidophora in New Guinea, Australia and Western Pacific nT

In addition, a further eight indigenous New Guinea/Pacific groups

are proposed and described here:

Brevispathacea Group (R. brevispathacea Engl. & K. Krause, R.

stolleana Engl. & K. Krause, perhaps R. geniculata Engl.):- Long

pendent stems (erect and clinging in R. geniculata) with much

abbreviated inflorescence-bearing shoots arising in the leaf axils. Spathe

marcescent.

Discolor Group (R. discolor Engl. & K. Krause, R. stenophylla K.

Krause, R. conica Engl., R. petrieana A. Hay):- Flowering on free

shoots. Spadix long stipitate. Rather to markedly narrow leaves.

(Perhaps not a good group because free shoots and stipitate spadices

recur throughout the genus but morphologically it is quite

homogeneous.)

Fortis Group (R. fortis P.C. Boyce):- Extremely stout, clinging

flowering shoots and short-petioled leaves. Inflorescences with

appearance of being borne in the leaf axils.

Jubata Group (R. jubata P.C. Boyce):- Robust, pachycaul, flowering

on adherent shoots. The crested reniform seeds are unique in the

genus.

(Similar in appearance to R. intonsa P.C. Boyce and R.

cravenschoddeana P.C. Boyce, which are different in not having crested

reniform seeds.)

Microspadix Group (R. microspadix K. Krause):- Long, very slender

shoots flowering at the tip. Spathe small, marcescent.

(May be part of the Neoguineensis Group).

Okapensis Group (R. gorokensis P.C. Boyce, R. kokodensis P.C. Boyce,

R. okapensis P.C. Boyce & Bogner, R. pilosa P.C. Boyce):- Montane

species with stiff coriaceous leaves with prominent petiolar sheathes

that degrade into fibres, or fall completely. Inflorescences on adherent

or free shoots. Spathe marcescent where known (perhaps early

marcescent in R. okapensis).

Pachyphylla Group (R. cryptantha P.C. Boyce & C.M. Allen, R. hayi

P.C. Boyce & Bogner, R. pachyphylla K. Krause (+ R. parvifolia

Alderw. — Maluku):- Defined by neotenic shingling habit with petioles

much shorter than the lamina.

Spuria Group (R. intonsa, R. cravenschoddeana, R. spuria (Schott)

Nicolson):- Fibre-covered shoot apices, large inflorescences, often

several together as in the Spathacea Group (q.v.) , but inflorescences

subtended by fibrous, not chartaceous, remains.

(Possibly heterogeneous).

78 Gard. Bull. Singapore 53 (2001)

Geography and Endemism

New Guinea, Australia and the western pacific region is remarkable for

the high number of Rhaphidophora species, coupled with very high diversity

and a high percentage of endemics (30 species of which 29 are endemic;

only the widespread and common R. korthalsii is not endemic to the area).

Apart from R. guamensis (endemic to Guam) and R. petrieana (endemic

to Australia), all the rest occur on the island of New Guinea, of which 16

species are endemic to the island. These figures, compared with those for

the other main nodes of diversity for the genus in tropical Asia -Borneo (5

endemics out of a total of 13 species), Peninsular Malaysia (2 out of 15),

Sumatera (4 out of 15) and the Philippine islands (7 out of 11) - suggest

Rhaphidophora is a genus of Gondwana origin.

There are no data for the conservation status of any of these endemic

species and much remains to be done by way of field studies to ascertain

whether any or all of these species are threatened. Nevertheless, it seems

that several species have very restricted distributions (e.g., R. kokodensis,

R. gorokensis, R. jubata), others are known from a single or very few

collections (e.g., R. pilosa, R. intrusa).

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, | (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. 1 (1897) 58; K. Schum. & Lauterbach, Fl. Schutzgeb.

Siidsee (1900) 211; Engl. & K. Krause in Engl., Pflanzenr. 37 (1V.23B)

(1908) 17—53; Engl. & Prantl, Nat. Pflanzenfam. Nachtr. 3 (1908) 29;

Nicolson in A.C. Sm., Fl. Vitiensis Nova, 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 subgen. Rhaphidophora (Hassk.) Miq., Flora Ned. Indié 3 (1856)

185. — Type: Rhaphidophora lacera Hasskarl, nom. illeg. pro. Pothos

Rhaphidophora in New Guinea, Australia and Western Pacific 79

pertusus Roxb. [= Rhaphidophora pertusa (Roxb.) Schott].

Scindapsus Schott subgen. Pothopsis Migq., Flora Ned. Indié 3 (1856) 187.

— Type: Scindapsus sylvestris (Blume) Kunth [= Rhaphidophora sylvestris

(Blume) Engl.].

[Raphidophora Hassk., Cat. Hort. Bogor. (1844) 58, orth. var.].

Medium-sized to very large, occasionally enormous, slender to robust,

leptocaul or pachycaul, homeophyllous or heterophyllous, rarely neotenic,

root-climbing lianes, very seldom clustering and rheophytic and then 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 or germinating to give rise to a non-

skototropic shingling juvenile shoot; pre-adult plants often forming modest

to extensive terrestrial colonies of varying morphological and physiological

form (descriptive generalisations 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: i. physiognomically

unbranched clinging non-flowering stems rooting along their entire length

giving rise to variously elaborated free lateral lateral flowering stems, or 11.

all stems physiognomically unbranched, clinging and flowering, or ii.

physiognomically unbranched leader and sympodial lateral stems clinging

but only sympodial lateral stems flowering: 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 sub-woody or somewhat corky or with distinctive matt to sub-lustrous

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 both cataphylls and prophylls deliquescing to black

mucilage and later drying leaving fragmentary parchment-like remains on

petioles, developing laminas and 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:

80 Gard. Bull. Singapore 53 (2001)

cataphylls and prophylls subcoriaceous to membranous, caducous or

degrading or deliquescing to variously persistent, variously textured, sheaths

and fibres, these where present variously clothing upper stem before

eventually decaying and falling; /eaves distichous or weakly spiralled, evenly

distributed or scattered or clustered distally; petiole canaliculate to weakly

carinate, smooth or pubescent, with variously prominent apical and basal

genicula; petiolar sheath prominent, nearly reaching to overtopping the

geniculum, occasionally one side greatly expanded and auriculate, especially

in juvenile plants, at first membranous to coriaceous, then soon drying

chartaceous completely or along the margins, sometimes degrading to untidy

variously netted or simple fibres and later variously falling leaving 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; midrib

usually prominently raised abaxially and prominently sunken, sometime

flush or rarely slightly raised adaxially; primary venation + pinnate;

interprimaries mostly present, sub-parallel to primaries and sometimes

indistinguishable from them 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 to reticulate,

variously prominent, often very difficult to distinguish from primary

venation; fertiary 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, inflorescences at male 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 male anthesis, usually caducous

before male anthesis is complete, occasionally marcescent into the early

stages of infructescence development, 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;

Rhaphidophora in New Guinea, Australia and Western Pacific 81

ovary 1- to partially 2-locular, lower part + bilaterally compressed, upper

part cylindrical and variously angled, most often rhombohexagonal, those

upper- and lower-most on the spadix often sterile and bereft of stigma,

those uppermost frequently either scattered or partially fused to each other

and forming a rudimentary appendix; ovules few to many, anatropous,

funicle long, placentas 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

male 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 male anthesis,

rarely not exserted and pollen extruded from between ovaries, thecae

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 shedding 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 (1 bear); refers

to the macroscopic (to 1cm long), needle-like unicellular trichosclereids

present in tissues.

Gard. Bull. Singapore 53 (2001)

Key to Adult Flowering* Rhaphidophora plants in New Guinea,

Australia and the Tropical Western Pacific

* juvenile vegetative phases included for species where confusion is possible

la.

1b.

2a.

2b.

3a.

3b.

4a.

4b.

Shingle climber (petiole much shorter than lamina, lamina appressed

to: substrate) cl eeceoics CAR ee ee eee 2

Not a8 above tisctines sc Sc. ccnGeeaghl a5. Rieeecee dace oes ean She eee ea eae eee 6

Leaf lamina cordiform, chartaceous, base cordate, posterior lobes

overlapping; primary venation strongly reticulate «0.0.0.0... 2)

Leaf lamina variously shaped (not cordiform), membranous to stiffly

coriaceous, base slightly cordate to truncate; primary venation pinnate

Primary venation conspicuously silver-grey; inflorescences arising

from behind appressed shingling leaves, carried on abbreviated shoots

arising from the leaf axil. Papua New Guinea (not known with

certainty,).2:..54. 0 2a ae ee eee 5. R. cryptantha

Primary venation not silver grey; plant never flowering in the

shingling phase. Indonesian Papua, Papua New Guinea (including

New Ineland)e...c..820-2 eee 29. R. versteegii (juvenile phase)

Leaf lamina membranous, with c. 3 prominent veins per side, these

running from near the base to the upper margin or tip and crossing

over the reticulate minor venation; plant never flowering in the

shingling phase; ‘Lhroughoutsresiones 02-2 ee eee

iL ed Bild dee ee 16. R. korthalsii (juvenile phase)

Leaf lamina coriaceous to stiffly coriaceous, without prominent veins

running from the base to the upper margin or tip; plant fertile in the

Shingling: phase. 2.2 <.sicsiszcseceeteceade eo ee 3

Leaf lamina of flowering shoots with base truncate; spadix stoutly

cigar-shaped, shortly stipitate, inserted + level on stipe; stipe 4—6 x

3—3.5 mm; stigma elongated, longitudinally orientated; plant with

disarticulating side shoots functioning as vegetative propagation units.

Indonesian Papua, Papua New Guinea [including New Britain, New

Ireland, Bougainville and Muyua (Woodlark) Island], and Australia

(Eastern-tropical Oucensland) i... pee, eee ee ee 11. R. hayi

Leaf lamina of flowering shoots with base acute to cuneate; spadix

narrowly cigar-shaped, long stipitate, inserted obliquely on stipe;

stipe 6—10 x 2—2.4 mm; stigma punctiform; plant without

Rhaphidophora in New Guinea, Australia and Western Pacific 83

6a.

6b.

Va:

7b.

8a.

8b.

9a.

9b.

10a.

10b.

Lika,

11b.

12a.

12b.

disarticulating side shoots. Indonesian Papua, Papua New Guinea.

EPR Sm eNE epee iA cyst Shc, Lae niaceal PiecnddsGeodanedencnsee 21. R. pachyphylla

Leaf lamina variously pinnately divided and/or perforated; roots

SOO Clio) (OTe) DULG LeSTe es 11 Be eres Pine et eRe eee a ee 7

Leaf lamina entire, if pinnate then with densely ramentose-scaly

Nt oe aoe a een aa Sago ctceceiannaivinegdeage aceavdseciesedectess 8

Leaf lamina of mature plants pinnatisect to slightly to extensively

perforated, perforations round to rhombic, extending c. + of lamina

width on each side of the midrib; shoot tips with prominent

chartaceous prophyll and cataphyll remains. Indonesian Papua, Papua

New Guinea (including New Ireland) ............00.0... 29. R. versteegii

Leaf lamina of mature plants pinnatisect, the pinnae often perforated

basally; active shoot tips with sparse to copious netted fibre.

PiiouehOut the LEPIOM ss... 5020) cs. -.che se ckench nsaseteenecbsvnces 16. R. korthalsii

Inflorescencesjanismig on clingimg ShOOtS: <2-.2sc.20s2-s cceveneeeteeeecceeneeee 9

Patloreseciices Anise (Om Fre SHOOUS) 2 ites cedeeeced. .dedehan tae cdcseceens-s 17

Shoot tips without variously textured cataphyll, prophyll and petiolar

sheath fibre; lamina ovate, 2.5—8.5 x 0.9—4 cm, very stiffly

coriaceous, base cordate to rounded and slightly notched, apex long-

acuminate with a pronounced tubule. Papua New Guinea...............

ee ee aes Ac ek RG a heed thee a vende. secniealilees tee 20. R. okapensis

Shoot tips with variously textured cataphyll, prophyll and petiolar

sheath fibre; leaf laminas various, but not the above combination of

BA eRe Spee eee oh, ae etl he steal Not ge vatieds suescecbeetioeeennee 10

Leaf lamina pinnatisect; feeding roots densely ramentose-scaly.

‘Werte ira ney 0 ret (0) eee ene ee eee 16. R. korthalsii

Leaf lamina entire; feeding roots smooth or pubescent ................ fet

JO ES VEY SES CY) SOG ie Soe ne en ee 2

CMGI SSL OUI eu en he bec eee Cones ee eee Soe ee eee 16

Inflorescences solitary; stylar region conical, stigma not prominent.

New Guinea and Australia (eastern tropical Queensland) ...............

ees Bache be festa wae shack chee ate go Bsa iedice sues 1. R. australasica

Inflorescences two or more together; stylar region truncate, stigma

SYST ICTS 0 Bs Pie Se, Neocon ye ee EE Re Pe 13

14a.

14b.

l6a.

16b.

Gard. Bull. Singapore 53 (2001)

Inflorescences subtended by netted fibre and copious sheet-like

tissue, arising two or more sequentially between foliage leaves. Papua

New Guinea 4.6.0 nce eee eres oe eee ee 7. R. fortis

Inflorescences each subtended by prominent chartaceous prophyll

and one or more large cataphylls, the whole forming a synflorescence

not mterspersed) with foliage leaves. e...es 2 ee eee eet 14

Prophylls and cataphylls subtending inflorescences soon degrading

and falling; spathe caducous, 16—24 cm long, stiff, thin; spadix slender

cylindrical, 13—21 x 1—3 cm, long decurrent (to 2 cm) at peduncle/

spathe insertion. New Britain and Bougainville ........... 14. R. jubata

Prophylls and cataphylls subtending inflorescences drying chartaceous

and persisting into fruit maturation; spathe marcescent, 5—13.5 cm

long, stiff-fleshy, very thick-walled (up to | cm at tip); spadix stoutly

cylindrical, 3—16.5 x 1—1.5 cm, inserted almost level on peduncle

sede dhe vawadennisivesateve tune ceueusta alee ikuca ata Bra Seance eae e cat Se EG ER ee ee ne 15

Leaf lamina drying pale grey-green, often (but not always) profoundly

perforated; juvenile growth phase shingling. Indonesian Papua, Papua

New Guinea (including New Ireland) ..................... 29. R. versteegii

Leaf lamina drying strongly bicolored, adaxially dark brown, abaxially

bright orange-brown with copious, minute tannin cells, never

perforate; juvenile growth phase not shingling. Palau, Indonesian

Papua, Papua New Guinea (including Woodlark (Muyua) Island),

Federated States of Micronesiai\( Yap) <.:......20<-0- 25. R. spathacea

Inflorescence solitary, terminating short shoots arising singly in the

axils of several adjacent leaves, each inflorescence subtended by a

prophyll and several degraded, netted cataphylls; spathe ovoid-

ellipsoid, 3.5—6.5 cm long; spadix ovoid-ellipsoid, 2.5—3.5 x 1.3—

1.7 cm; stylar region prominently conical, hexagonal in plan view;

Stigma punctiform. New 'Giineay,..ennee eee 8. R. geniculata

Inflorescences several close together in a synflorescence, each

subtended by a prominent chartaceous prophyll and one or more

chartaceous cataphylls, the entire synflorescence emerging from a

mass of chartaceous cataphylls; spathe canoe-shaped, 4—5.5 cm long;

spadix ylindrical, c. 3.2 x 0.9 cm; stylar region rounded-conical, almost

circular in plan view; stigma deeply excavated. Indonesian Papua

(Mimika Prov.) and Papua New Guinea (East Sepik Prov.) ............

CT eee Ee erred ha Sic ae 29. R. waria

Rhaphidophora in New Guinea, Australia and Western Pacific 85

17a.

17b.

18a.

18b.

19a.

19b.

20a.

20b.

21a.

21b.

22a.

Shoot tips with variously textured cataphyll, prophyll and petiolar

Senne eke os eaeor ter ar ne Cee 16

Shoot tips without cataphyll, prophyll and petiolar sheath fibre .....

Seer peace ie a ae Sl a sian ats asain enn 24

LS Ue TSN SSE SEES Ee 20 ne 19

en ee 20

Flowering shoots extremely abbreviated, arising from axils of leaves,

all but obscured by netted petiolar sheath and cataphyll fibre: spathe

ovoid-cylindric, 3—4 cm long, thinly coriaceous: spadix slender

cylindrical, 2.9—3.4 x 1.7—1.9 cm: stylar region depressed centrally:

anthers exserted at male anthesis. Indonesian Papua (Mimika/Digul

Soe Ey 5 gee es ae ee 2. R. brevispathacea

Flowering shoots elongated and leafy, not as above: spathe canoe-

shaped, 20—23 cm long, stiffly very fleshy: spadix cylindrical to

slightly cigar-shaped, 13.5—19 x 2.2—4 cm: stylar region truncate

with a raised rim; anthers not exserted at male anthesis. Indonesian

Papua (Yapen), Papua New Guinea (East & West Sepik, Milne Bay

TC ER 8 | a ne ee 6. R. discolor

Leaf lamina narrowly lanceolate to narrowly lanceolate-elliptic, not

pee enn ASX OP CARE SEEN, CANIAC COIS 5. ne teed aad 21

Leaf lamina ovate to oblong-elliptic or oblong lanceolate, slightly

oblique, frequently exceeding 30 x 10 cm, up to 76 x 32,

submembranous, subcoriaceous, only rarely coriaceous and then

oie AE Sete gO ha th ee BP

Cataphylls, prophylls and petiolar sheath degrading to simple robust

fibres. Papua New Guinea (Kokoda & Port Moresby).................-..-.

0 REET 2 ie ae gee ee ee 15. R. kokodensis

Cataphylls, prophylls and petiolar sheath degrading to weak fibres

and strips of soft felt-like tissue. Indonesian Papua (Kepala Burung

emus Ice MURA es Sexo SE Pei at th a at, Sat ab tintacs 23. R. pilosa

Leaf lamina drying strongly bicolored, adaxially mid-brown, abaxially

pale brown: inflorescences usually several together, occasionally

solitary each subtended by a soon-degrading membranous prophyll

and cataphyll; spathe 9.5—12 (-16 cm) long. Papua New Guinea

(Western Prov.). Solomon Islands (Guadalcanal, New Georgia, San

Cristobal and the Santa Cruz Group), Fiji, Western and American

lk ae ta ect tea 26. R. spuria

220:

123i

24a.

24b.

25a.

25D:

26a.

26b.

iar

27b.

Gard. Bull. Singapore 53 (2001)

Leaf lamina not drying bicolored or if bicolored then abaxially

greenish brown, adaxially dark brown with dense faint dark speckling;

inflorescence always solitary, subtended by a + fully developed foliage

leaf and copious netted fibre and sheet-like tissue or a + fully

developed foliage leaf and one or more degrading papery cataphylls;

Spathe 17—22: lomo... ccicaieasictsvtectseerssnescebeeopene ve Meceeecende one daneenetieeeaeee 23

Leaf lamina subcoriaceous, drying concolorous, base subacute to

slightly decurrent; inflorescence subtended by a + fully developed

foliage leaf and copious netted fibre and sheet-like tissue; spathe

17—20 cm long; spadix 10.5—18.5 x 1.7—2.3 cm; anthers not exserted

at male anthesis. Papua New Guinea (Central & Morobe) ..............

beans ac he AISA aa ee 12. R. intonsa

Leaf lamina submembranous, drying bicolored, abaxially greenish

brown, adaxially dark brown and densely faint dark speckled, base

oblique, one side rounded to almost truncate, the other subacute;

inflorescence subtended by a + fully developed foliage leaf and one

or more degrading papery cataphylls; spathe c. 22 cm long; spadix

17—19 x 1.5—2 cm, yellow green; anthers long-exserted at male

anthesis. Papua New Guinea (East Sepik, Morobe & Bougainville)

widen ttbncabietuedesadshdnde chon snp dceat dee mee belt Soe aan ene een 4. R. cravenschoddeana

Spadix stipitate:.. .cscsc LE a

Spadix Sessile, 2857... 2 ee as)

Stylar region conicallt....2 ee eee 26

Stylar resiomtruncate wate eee ee 28

Spadix base with cochleate insertion; stigma glossy, black, at tip of

long (c. 2 mm) slender stipe; anthers not exserted at male anthesis.

Guam Si04..52h Rae ee 10. R. guamensis

Spadix base tapering to almost truncate; stigma not as above; anthers

exserted at male anthesis. Papua New Guinea (including Louisiade

Archipelago); Australia (Oweensland)) we. een ee eee eee Dy

Petiolar sheath soon drying and eventually falling more-or-less entire

leaving a conspicuous scar; inflorescence solitary, subtended by a

partially developed foliage leaf and a membranous cataphyll; spathe

6—8 cm long; spadix stipe c. 1 x 0.6 cm. Australia (Queensland) ...

DpaddebSh oles Ue sncsstaduie ease TA Ace 22. R. petrieana

Petiolar sheath persistent through to leaf fall; inflorescence not

solitary, subtended by a + fully developed foliage leaf; spathe 11—

Rhaphidophora in New Guinea, Australia and Western Pacific 87

28a.

28b.

29a.

29b.

30a.

30b.

31a.

31b.

37a.

13 cm long; spadix stipe 2.5—3 x 3—0.4 cm. Papua New Guinea

(including Louisiade Archipelago)..............00 27. R. stenophylla

Petiolar sheath extending beyond the apical geniculum by two ligules

adaxially and by fused extensions along the abaxial midrib for up to

half the leaf lamina length; leaf lamina oblong-elliptic, up to 16.5 cm

long, submembranous; spathe apex truncate and extending into a

very long, slender beak up to 1/3; length of entire spathe: spadix

cigar-shaped; anthers not exserted at male anthesis. Indonesian Papua

(LEER NT Ee ee eee See ee 13. R. intrusa

Petiolar sheath extending beyond the apical geniculum only by two

tiny ligules; leaf lamina, lanceolate to oblong-lanceolate, up to 34

cm long and usually exceeding 18 cm, thinly coriaceous; spathe apex

stoutly short-beaked, beak not exceeding '/s length of entire spathe;

spadix cylindrical; anthers strongly exserted at male anthesis. Papua

New Guinea (including Bismarck Archipelago), Solomon Islands..

Se ere oe La BIE a ee Oa es, th as 24. R. schlechteri

Stylar region conical. Primary lateral veins drying much paler than

abaxial leaf lamina. Indonesian Papua (Kepala Burung & Mimika

Prov.), Papua New Guinea (Muyua, New Britain) ........ 3. R. conica

Stylar region truncate. Primary lateral veins not as above............ 30

Leaf lamina stiffly coriaceous, narrowly lanceolate; spathe 6—6.5

cm long: spadix cylindrical, 3.3—3.5 x c. 0.7 cm. Papua New Guinea

(Eastern Highlands — Goroka; Southern Highlands — Mendi) .........

teas Mee Me A ctv cnosivhusvand tao tinesoeneeenscvowen 9. R. gorokensis

Leaf lamina membranous to thinly coriaceous, variously shaped but

not narrowly lanceolate, or if narrowly lanceolate then not stiffly

coriaceous and inflorescences borne on the tips of long, slender,

pendent stems; spathe 2.5—3 cm long: spadix ovoid-globose to ovoid-

EME (eB ree pcg a My 2 |b 0 7 di an 31

Inflorescences arising from long, pendent shoots ..............0.:cce 32

Inflorescences arising from short, spreading shoots ..............:::00+ 35

Leaf lamina narrowly lanceolate to lanceolate falcate, 4—19 x 1—3

cm, thinly coriaceous, weakly bicolored, adaxially dull mid-brown,

abaxially paler brown with dense tannin cells; inflorescences borne

at tips of very long, slender flowering stems (c. 2 mm diam. in dried

material); spathe ovoid-ellipsoid, stoutly long-beaked, 2.5—3 x 0.7—

0.9 cm; spadix ovoid-globose to ovoid-cylindric, 1—1.5 x 0.5—0.6

88 Gard. Bull. Singapore 53 (2001)

cm. Indonesian Papua (Mimika Prov.), Papua New Guinea (Madang

SOMOrObEe Prov.) svscsissssvecesd heise ssscese- tesco cane ee ae ee

digta Peele Sale Bele ica Ben hse sa calges Si as nae, ee ee 17. R. microspadix

32b. Leaf lamina elliptic to oblong-elliptic, 20—29 x 8—13 cm,

membranous, markedly bicolored, adaxially mid-brown, abaxially

pale orange brown; inflorescences borne on short shoots arising from

axils of leaves along long, pendent flowering stems c. 1 cm diam. in

dried material; spathe elongate-cylindric, stoutly short-beaked, c.

10.2 x 1.7 cm; spadix slender cylindrical, 8 x 1.1 cm. Papua New

Guineai(East Sepik Provs),. 2. ee ee 28. R. stolleana

33a. Spathe cigar-shaped, 1—4.5 x 0.7—1.2 cm, caducous; spadix

cylindrical, 1.5—2.5 x 0.5—0.8; anthers not exserted at male anthesis.

Papua New Guinea (Bougainville & Manus), Solomon Islands.......

asiage Dasicecagd dodo wualoubctbealalgcordSeetidan le SOEs ramet ee geen ae a 18. R. mima

33b. Spathe globose to ovoid-ellipsoid, 1—2 x 1—2.5 cm, marcescent;

spadix globose to ellipsoid-cylindrical, 1—1.5 x 0.7—1 cm; anthers

well-exserted at male anthesis. Mainland New Guinea. ...............04

oh aaa vtulouitct tea ce le ca dipallgthtc Cada Una Uee tee cere ene 19. R. neoguineensis

The Species

1. Rhaphidophora australasica F.M. Bailey

Rhaphidophora australasica F.M. Bailey, Queensland Agric. J. 1 (1897)

452 & Fl. Queensland 5 (1902) 1697; Engl. & K. Krause in Engl., Pflanzenr.

37 ([V.23B) (1908) 28; Williams, Native Pl. Queensland 1 (1979) 252,

unnumbered plate; Jones & Gray, Climbing Pl. Australia (1988) 316,

unnumbered plate p. 322; Hay, Telopea 5 (1993) 296—297. — Type:

Australia, Queensland, Cairns, Cowley s.n. (BRI, holo; K, iso).

Rhaphidophora hollrungii Engl. in K. Schum. & Hollrung, Fl. Kais.-Wilh.

Land (1889) 19; Engl. in Bot. Jahrb. Syst. 25 (1898) 9; K. Schum. &

Lauterbach, Fl. Schutzgeb. Stidsee (1900) 211; Engl. & K. Krause in Engl.,

Pflanzenr. 37 (1V.23B) (1908) 44, Fig. 18, synon. noy. — Type: Papua New

Guinea (‘Kaiser Wilhelmsland’), West Sepik Prov., Augusta River, Sept.

1897, Hollrung 746 (B, lecto; selected here). Engler cited two syntypes, the

other (Papua New Guinea (‘Kaiser Wilhelmsland’), Morobe Prov.,

Sattelberg, July 1890, Lauterbach 588) is missing from Berlin and presumed

destroyed.

Rhaphidophora in New Guinea, Australia and Western Pacific 89

Rhaphidophora iboensis K. Krause, Bot. Jahrb. Syst. 49 (1912) 93, synon.

noy. — Type: Papua New Guinea (‘Kaiser Wilhelmsland’), Madang Prov.,

Ibo Range, 17 Dec. 1908, Schlechter 18985 (B, holo).

Figure 1

Moderately robust, pachycaul, homeophyllous liane to 15 m; seedling stage

not observed; pre-adult plants forming sparse terrestrial colonies; adult

shoot architecture comprised of clinging, physiognomically unbranched,

densely leafy flowering stems; stems smooth, later with lustrous papery

epidermis, cataphylls and prophylls soon degrading into long, untidy fibres,

internodes 1—5 x 0.5—1 cm, separated by prominent slightly oblique corky

leaf scars; flagellate foraging stem absent; clasping roots densely arising

from the nodes and internodes, minutely pubescent; feeding roots not

observed; /eaves spiro-distichous; cataphylls and prophylls chartaceous but

soon degrading into long fibres; petiole deeply grooved, 9—22 x 0.25—0.3

cm, smooth, apical and basal geniculum prominent and drying slightly

darker; petiolar sheath extending to apical geniculum, short-persistent, at

first, chartaceous, soon degrading to fibres; /amina entire, lanceolate to

elongate-lanceolate or elongate-oblanceolate, slightly falcate, 19—46 x 2.5—

9.5 cm, subcoriaceous, base rounded to subacute, apex acute to acuminate;

midrib raised abaxially, + sunken adaxially; primary venation pinnate,

slightly raised abaxially, much less-so adaxially; interprimaries sub-parallel

to primaries, and almost indistinguishable from primary lateral veins;

secondary venation reticulate, slightly to barely visible; inflorescence solitary,

subtended by an incompletely developed leaf and much fibrous cataphyll

remains; peduncle slightly robust, terete, 3.5—9.5 x 0.2—0.35 cm; spathe

slender canoe-shaped, long beaked, 5—9.5 x 2—2.5 cm, stiff-fleshy, cream,

caducous leaving a prominent scar; spadix cylindrical, sessile, inserted

slightly obliquely on peduncle, 3.75—8 x 1—2 cm, cream; stylar region

long-conical, weakly hexagonal in top view, drying longitudinally ribbed,

1.1—1.2 x 1—1.1 mm; stigma punctiform, minutely excavated, 0.1—0.2 x c.

0.3 mm; anthers exserted at male anthesis; infructescence oblong-cylindrical,

8—9 x 1.5—2 cm.

Distribution: New Guinea (widespread but scattered: often abundant where

it occurs), Australia (eastern tropical Queensland).

Habitat: Rain forest. Sea level to 760 m altitude.

Other specimens seen: INDONESIAN PAPUA. Jayapura Prov.: Cyclop Mts, above Ifaar

(‘Ifar’), McKee 1877 (L). PAPUA NEW GUINEA. Morobe Prov.: Lae, White et al. NGF

1555 (L). AUSTRALIA. Queensland: Cook District, Mossman, entrance to Mossman Gorge,

Gard. Bull. Singapore 53 (2001)

/ ,

If AN \ \

Mf \.\\\"

L GUAR

Figure 1. Rhaphidophora australasica F.

A. flowering shoot x '/,; B. leaf lamina x

M. Bailey

'/,, C. venation detail x 4; D. inflorescence, spathe

fallen x '/,; E. spadix detail, post-male anthesis x 8; F. stylar region and stigma, side view x 8.

A-C from McKee 1877; D-F from Hollrung 746.

Rhaphidophora in New Guinea, Australia and Western Pacific 9]

Blake 19756 (K); c. 20 miles NNW of Daintree, Boyland 399 (K); Along road from Atherton

to Main Coastal Highway, along Palmerston Highway, Henrietta Falls, Croat 52614 (MO);

Wright Creek, Lake Eacham State Park, 20 km E of Atherton, Croat 52575 (K, MO);

S.F.R. 756, East Downey L.A., Hyland 5608 (K, L); Wright Creek, c. 10 miles E of Atherton,

Melville 3693 (K, L, US); Lacey’s Creek, Mission Beach area, Smith 10189 (K).

Notes. 1. Almost all New Guinea specimens previously identified as R.

australasica (incl. R. hollrungii) belong to either R. spathacea or to one of

several new species described elsewhere in this paper. The combination of

solitary inflorescences arising on clinging shoots, fibrous cataphyll remains

and conical style is diagnostic for R. australasica.

2. The two widely separated New Guinea localities suggest a species that is

widespread but uncommon in New Guinea.

2. Rhaphidophora brevispathacea Engl. & K. Krause

Rhaphidophora brevispathacea Engl. & K. Krause, Bot. Jahrb. Syst. 54

(1916) 79; K. Krause & Alderw., Nova Guinea 14 (1924) 212. — Type:

Indonesian Papua, Mimika/Digul Prov. boundary, Lorentz River, near

Kloofbivak, 31 Oct. 1912, Pulle 254 (B, holo; BO, K, L, iso).

Figure 2

Moderate, somewhat slender, leptocaul, homeophyllous (?) liane of

unknown ultimate height; seedling and pre-adult plants not observed; adult

shoot architecture not completely known, but observed to be comprised of

greatly elongated, spreading to strongly pendent, physiognomically

unbranched, leafy, non-flowering stems giving rise to extremely short, free,

sympodial, (foliage) leafless, flowering stems; stems smooth, climbing stems

not observed, free stems weakly rectangular to + terete in cross-section,

dull yellow brown, internodes to 5 x 0.7 cm, flowering shoots extremely

abbreviated, arising from axils of leaves and, where stems pendent then

twisting to present inflorescences upwards, stems all but obscured by netted

petiolar sheath and cataphyll fibre; flagellate foraging stems not observed;

clasping roots arising singly from the nodes and internodes, slightly

pubescent, later glabrescent; feeding roots not observed; leaves weakly

spiralled, scattered; cataphylls and prophylls membranous, very quickly

degrading leaving semi-persistent netted remains; petiole grooved adaxially,

7—9.5 x 0.2—0.3 cm, smooth, with a weakly defined apical and slightly

prominent basal geniculum; petiolar sheath slightly prominent, extending

beyond the apical geniculum by two ligules, caducous leaving a slight

continuous scar from the petiole base, around the top of the apical

92 Gard. Bull. Singapore 53 (2001)

geniculum and back to the base; /amina entire, elliptic to elliptic- or

lanceolate-elliptic, 6.2—20 x 2.5—8.75 cm, coriaceous, upper surfaces very

slightly glossy, drying strongly bicolorous, adaxially dark brown, abaxially

bright orange-brown with copious, minute tannin cells, base subovate, very

briefly decurrent, apex acute to briefly acuminate, with a minute apiculate

tubule; midrib slightly raised abaxially, sunken adaxially; primary venation

pinnate, slightly raised abaxially, prominent (darker 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 dried specimens, very weakly reticulate; inflorescence solitary,

subtended by several reduced leaves (lamina absent) with rapidly degrading

petiolar sheaths and netted cataphylls, the whole obscuring the peduncle;

peduncle terete, 3—4 x 0.25—0.3 cm; spathe ovoid-cylindric, stoutly very

short-beaked, 3—4 x 2—2.3 cm, thinly coriaceous, marcescent to fruiting,

then abcissing basally and pushed off by developing fruits, leaving a slight

scar; spadix slender cylindrical, stipitate, inserted more or less level on

stipe, 2.9—3.4 x 1.7—1.9 cm; stipe slender terete, c. 3 x 1.5 mm; stylar

region rhombohexagonal, 1.7—2.1 x c. 2 mm, depressed centrally; stigma

punctiform, c. 0.4 mm diam., prominent in dried material; anthers exserted

at male anthesis; infructescence not observed.

Distribution: Indonesian Papua (Mimika/Digul Prov. boundary). Known

only from the type.

Habitat: Not recorded. 100 m altitude.

Note: Distinctive by the long pendent stems with much abbreviated, netting-

sheathed flowering shoots and a small, marcescent spathe. Rhaphidophora

stolleana shares the habit of long pendent stems with abbreviated of

inflorescence-bearing shoots arising in the leaf axils, but is readily

distinguished by a much longer spathe (c. 10.2 cm long) and spadix (c. 8 cm

long). To date R. brevispathacea is known only from Mimika/Digul Prov.

and R. stolleana only from East Sepik. In the small, marcescent spathe, R.

brevispathacea resembles R. neoguineensis, although the latter has a quite

different growth habit and inflorescence presentation.

3. Rhaphidophora conica Engl.

Rhaphidophora conica Engl., Bot. Jahrb. Syst. 1 (1881) 181. — Type:

Indonesian Papua, Kepala Burung Prov., Ramoi, Feb. 1875, Beccari PP

409 (FI, holo).

Rhaphidophora in New Guinea, Australia and Western Pacific 93

LGOURR

Figure 2. Rhaphidophora brevispathacea Engl. & K. Krause

A. adult shoot with flowering branches x '/,; B. leaf lamina x '/,; C. venation detail x 2; D.

inflorescence, spathe fallen x 1'/,; E. spadix detail, post-male anthesis x 8; F. stylar region and

stigma, side view x 8. All from Pulle 254.

94 Gard. Bull. Singapore 53 (2001)

Rhaphidophora pallidivenia Alderw., Bull. Jard. Bot. Buitenzorg III, 4

(1922) 339, synon. nov. — Type: Indonesian Papua, Mimika Province,

Varen (Lorentz) River, 15 June 1907, Djibda s.n., ex cult. Bogor B.G.

Y.40, April 1921, Alderwerelt s.n. (BO, holo).

Figure 3

Moderately robust, semi-leptocaul, homeophyllous (?) liane to 5 m; seedling

and pre-adult plants not observed; adult shoot architecture comprised of

elongated, clinging, physiognomically unbranched, leafy, non-flowering

stems and elongated, free, leafy, flowering stems; stems smooth, flexuous,

stems + terete in cross-section, without prophyll, cataphyll fibre, internodes

5—10 x 0.5—1 cm, separated by weak straight leaf scars, older stems woody;

flagellate foraging stems not observed; clasping roots arising singly from

nodes; feeding roots solitary from nodes; leaves weakly spiro-distichous,

moderately densely arranged; cataphylls and prophylls membranous,

caducous; petiole grooved adaxially, 7—30 x 0.6—1 cm, smooth, with a

prominent apical and basal geniculum; petiolar sheath prominent, extending

to base or top of apical geniculum, soon drying chartaceous, then degrading

very slightly to weak fibres before falling leaving a feeble scar; lamina

entire, elliptic to elliptic-lanceolate, 1O—40 x 5—12 cm, thinly coriaceous,

base cuneate to acute, apex acute-acuminate, with a minute tubule; midrib

raised abaxially, slightly sunken adaxially, drying conspicuously lighter than

surrounding lamina; primary venation pinnate, slightly raised on both

surfaces in dried material, drying lighter; interprimaries sub-parallel to

primaries and only slightly less conspicuous; secondary venation reticulate,

fine but conspicuous; inflorescence solitary, subtended by a fully developed

foliage leaf; peduncle slightly compressed-terete, 2.5—10 x 0.2—0.35 cm;

spathe broadly cone-shaped to ellipsoid, truncate basally at insertion on

petiole, apex briefly to rather long-beaked, 5—12 x 24 cm, thinly fleshy,

greenish yellow, caducous, drying pale brown, falling leaving a thin scar;

spadix cylindrical, sessile, inserted level on peduncle, 2—7 x 0.7—1.2 cm,

obtuse, white to yellow; stylar region rounded-rhombohexagonal, 0.9—1.1

x 0.9—1.1 mm, slightly blunt-conical; stigma punctiform, c. 0.3 mm diam.,

rather prominent in dried material; anthers slightly exserted at male anthesis;

infructescence not observed.

Distribution: Indonesian Papua (Kepala Burung and Mimika Prov.), Papua

New Guinea (Muyua, New Britain).

Habitat: Rain forests, sometimes on coralline limestone. No altitudes

recorded.

Figure 3. Rhaphidophora conica Engl.

A. flowering shoot x '/,; B. leaf lamina x '/;;C. venation detail x 3; D. portion of creeping adult

stem x '/,; E. inflorescence x ?/,; F. spadix detail, pre-anthesis x 8. All from Furtado s.n.

96 Gard. Bull. Singapore 53 (2001)

Notes: 1. Brass 28608, while much smaller than the other collections, is

morphologically identical.

Other specimens seen: INDONESIAN PAPUA. Mimika Prov.: Varen (Lorentz) River,

Cult. Bogor, Y40, Furtado s.n. (SING), Nicolson 926 (US). PAPUA NEW GUINEA.

Woodlark (Muyua) Island: Kulumadu, Brass 28608 (GH, L); West New Britain Prov.: 2

miles northeast of Kandrian, Nicolson 1552 (K, L, US).

4. Rhaphidophora cravenschoddeana P.C. Boyce, sp. nov.

Rhaphidophora cravenschoddeana facile cum R. intonsa confunditur (species

etiam cum inflorescentiis grandibus et apicibus surculorum fibrosis) sed

fibris simplicibus neque reticulatis, lamina foliae textu tenuiore, costa

abaxialiter prominenti, venis lateralibus primariis eis interprimariis magis

‘diversis, antheris longe exsertis seorsa. — TYPUS: Papua New Guinea,

North Solomons Prov., Bougainville, near Koniguru No.1, c. 11 miles north

of Buin, 18 July 1963, Craven & Schodde 14 (CANB, holo; GH, K, iso).

Figure 4

Large, robust, pachycaul (?), homeophyllous liane to unknown ultimate

height; seedling stage not observed; pre-adult plants consisting of sparse

terrestrial colonies; adult shoot comprised of elongated, clinging,

physiognomically unbranched, non-flowering stems and free lateral, leafy

flowering stems; stems with cataphylls and prophylls degrading to long

sparse ragged fibres and weak sheets of tissue, internodes 0.5—3 x 0.5—2

cm, separated by prominent to very large unevenly oblique corky leaf

scars; flagellate foraging stem and clasping roots not observed; feeding roots

stout, arising singly from the node, densely velvety-hairy; /eaves spiro-

distichous; cataphylls and prophylls quickly degrading to long sparse ragged

fibres and weak sheets of tissue; petiole canaliculate, 6—46.5 x 1.3—1.5 cm,

apical geniculum very large, prominent, basal geniculum very weakly

defined; petiolar sheath very prominent, extending to apical geniculum,

swiftly degrading to long ragged fibres and weak sheets of tissue, later

falling leaving a jagged scar; lamina entire, ovate to oblong-elliptic or

oblong-ovate, strongly oblique, 8—S3 x 6—26 cm, submembranous, drying

discolorous, adaxially dark brown, densely faint dark speckled, abaxially

greenish brown, base oblique, one side rounded to almost truncate, other

subacute, apex subacute to acuminate or briefly attenuate; midrib very

prominently raised abaxially, slightly sunken adaxially; primary venation

pinnate, prominently raised abaxially, very slightly raised adaxially, drying

paler than lamina; interprimaries parallel to primaries, much less prominent,

slightly raised abaxially, very slightly raised adaxially; secondary venation

Rhaphidophora in New Guinea, Australia and Western Pacific 97

Figure 4. Rhaphidophora cravenschoddeana P.C. Boyce

A. part of flowering shoot x '/,; B. leaf lamina x '/;;C. venation detail x 2; D. entire inflorescence

x '/,; E. spadix detail, post-male anthesis x 2'/,; F. portion of creeping juvenile stem x 7/; G.

portion of pre-adult stem x '/,;. A-E from Craven & Schodde 14, F & G from Croat 52760.

98 Gard. Bull. Singapore 53 (2001)

parallel to interprimaries and only slightly less prominent, very slightly

raised abaxially and adaxially; tertiary venation reticulate, weakly raised

abaxially, invisible abaxially; inflorescence solitary, subtended by a + fully

developed foliage leaf and one or more degrading papery cataphylls;

peduncle laterally compressed, stout, 13—15 x 0.7—1 cm; spathe slender

canoe-shaped, 22 x 4 cm, stoutly long-beaked, caducous before male

anthesis, falling leaving a large scar; spadix stoutly cylindrical, sessile,

inserted more or less level on peduncle, 17—19 x 1.5—2 cm, yellow green;

stylar region hexagonal, 1.5—2 x 1.2—1.5 mm, truncate; stigma punctiform

to slightly ellipsoid, very slightly raised, 0.4 x 0.4—0.5 mm diam.; anthers

long-exserted at male anthesis; infructescence not observed.

Distribution: Papua New Guinea (East Sepik and Morobe Prov.),

Bougainville.

Habitat: Primary and secondary riverine rain forest. 100-1000 m altitude.

Notes: 1. Easily confused with Rhaphidophora intonsa (also with large

inflorescences and fibrous shoot tips) but separated by the simple, not

netted, fibres, thinner-textured leaf lamina with, on the abaxial surface, a

conspicuously raised midrib and much more pronounced differentiation

between the primary lateral and interprimary veins, and by the long-exserted

anthers.

2. The specific epithet is for the collectors of the type, Lyndley Craven and

Richard Schodde now at CANB, who made many collections in Papua

New Guinea.

Other specimens seen: PAPUA NEW GUINEA. East Sepik Prov.: Vicinity of Wewak,

along trail beyond Boys Town (Reform School operated by S.V.D. Missionaries), Croat

52760 (MO), Croat 52763A (MO); Angoram, Upper Karawari River, Domstreich 77 (L):

Morobe Provy.: Sattelberg, Clemens 8064 (B), 2066 (B); Along road to Sankewap, 1 km in

from Sankewap River, beginning 2.7 km south of Lutheran School, Croat 52818 (MO);

Along Busu River, 22.6 km by road SE of Lae via road past PNG University of Technology

and Igam road past Army Base, beginning 3.8 km from asphalt road at Army Base, Croat

52842 (MO): North Solomons Prov.: Bougainville: Maide River gorge, lower south slopes

of Lake Lolow crater, c. 15 miles north of Buin, Craven & Schodde 286 (CANB, GH); Near

Barilo village, c. 6 miles north of Buin, Craven & Schodde 375 (CANB, GH, K).

Rhaphidophora in New Guinea, Australia and Western Pacific 99

5. Rhaphidophora cryptantha P.C. Boyce & C.M. Allen, sp. nov.

Architectura surculi R. cryptanthae eam R. hayi ob inflorescentias in apicibus

surculorum abbreviatorum facile depulsorum ex axillis surculorum

scindulantium exorientium feruntur arctissime revocat. R. cryptantha

inflorescentias post folia ferenti, forma atque colore folii laminae, spatha

minore marcescenti, spadice ellipsoidali atque stylo stigmatico longo

nihilominus differt. Inflorescentiis in surculis post folia appressa scindulantia

vectis orientibus ac aperientibus R. cryptantha distincta est ab omnibus

speciebus ceteris neotenicus (R. hayi, R. pachyphylla et R. parviflora) in

quibus inflorescentiae in surculis a foliis porrectis portatae sunt, atque in

genere unica est. — TYPUS: Cultivated Fairchild Tropical Garden (acc.

no. 74407), ex Papua New Guinea, Botanic Garden, Lae, 24 Sept. 2000,

Boyce 1447 (FTG, holo: K. M, MO, SING, iso).

Figures 5 & 6

Moderate-sized, slender to somewhat robust, semi-leptocaul, homeophyllous

neotenic liane to 3 m: seedling stage unknown: pre-adult plants shingling,

hardly or not forming terrestrial colonies; adult shoot architecture comprised

of clinging, physiognomically unbranched, densely leafy, sterile stems and

very abbreviated. free, flowering stems arising from the axils and remaining

concealed behind the leaves of the main stem; stems rectangular in cross-

section, widest side slightly convex, smooth, dark green, without prophyll

and cataphyll fibre but with newest parts very thin, adherent, petiolar

sheath tissue, internodes 1—5 x 0.2—0.5 cm, separated by + straight scars:

flagellate foraging stems rather weakly developed, usually at least partially

leafy: clasping roots arising from the internodes, prominently pubescent.

spreading but usually not extending much beyond the leaf span: feeding

roots not observed (absent?): eaves distichous, shingling and ascending on

adherent shoots, densely arranged and slightly spreading on free shoots,

on flagella shoots leaves scattered with internodes between carrying a

prominent cataphyll of short duration: cataphylls and prophylls

membranous, caducous; petiole very shallowly grooved, 0.75—1.5 x 0.3—

0.4 cm, smooth, apical and basal genicula barely visible: petiolar sheath

prominent, membranous, long-ligulate, margins of ligule fused. the ligule

extending up to 3.5 cm above base of lamina and enclosing shoot apex.

caducous but adhering to stem: /amina broadly ovate, coriaceous, base

cordate, apex rounded with a tiny tubule; midrib barely visible abaxially

and adaxially: primary venation reticulate, barely visible abaxially, slightly

raised and silver-grey adaxially: interprimaries more-or-less absent:

secondary venation reticulate, flush abaxially, very weakly raised adaxially.

silver-grey: inflorescence sequentially produced from a shoot arising from

100 Gard. Bull. Singapore 53 (2001)

Figure 5. Rhaphidophora cryptantha P.C. Boyce & C. Allen

A. abaxial view of adult stem with two flowering branches, x 7/,; B. entire flowering branch

with inflorescence, spathe cut away to show spadix x 2. All from Boyce 1447.

Rhaphidophora in New Guinea, Australia and Western Pacific 101

Figure 6. Rhaphidophora cryptantha P.C. Boyce & C. Allen

A. adaxial view of adult stem, x 7/;; B. adaxial view of juvenile stem, x 7/;; C. flagellate stem x 7/3.

All from Boyce 1447.

102 Gard. Bull. Singapore 53 (2001)

behind the leaf, each subtended by a very small membranous, caducous

prophyll and one or more very reduced leaves; peduncle terete, 0.75—1 x

0.2—0.3 cm; spathe ellipsoid, weakly beaked, 3.5—4 x 1.2—1.5 cm, spongy-

fleshy, yellow, inflated and gaping at female anthesis and then closing and

drying onto developing infructescence; spadix ellipsoid, stipitate, inserted

+ level on stipe, 2.1—2.2 x 0.7—0.8 cm, creamy white; stipe 5—6 x 2—2

mm, white; stylar region irregularly rhombohexagonal, 2—2.2 x 2.1—2.3

mm, weakly conical; stigma very prominently raised on a c. 1 mm stipe,

punctiform to globose, c. 0.5 mm diam.; anthers exserted at male anthesis;

infructescence not seen.

Distribution: Not known with certainty. The specimen used to make the

type was introduced into cultivation from Lae Botanic Garden and it seems

quite possible that it originated in Papua New Guinea. The plant in

cultivation at Kew was received from Ted Green, also ex Papua New

Guinea.

Habitat: Unknown. In cultivation the plant is very intolerant of cool

conditions, suggesting that it originates from low elevations.

Notes: 1. A species of extraordinary appearance, both in flowering in a

neotenic state (otherwise known from R. hayi, R. parvifolia (Maluku) and

R. pachyphylla) and especially by the inflorescences arising and opening

on shoots carried behind the appressed shingling leaves. Often the mature

inflorescences are completely obscured by the leaves, although occasionally

the tip of the spathe is visible. In this flowering mode R. cryptantha is

distinct from the neotenic species listed above (in which the inflorescences

are carried on shoots held clear of the leaves), and unique in the genus.

The shoot architecture of R. cryptantha most closely recalls that of

R. hayi in that the inflorescences are borne on the tips of easily dislodged

abbreviated shoots arising from the axils of shingling shoots. However, R.

cryptantha differs in bearing the inflorescences behind the foliage, in the

shape and coloration of the leaf lamina, the smaller, marcescent spathe,

ellipsoid spadix and long style.

2. In cultivation R. cryptantha has been confused with Monstera dubia, a

species with a similar shingling juvenile stage and variegated leaves. The

two are readily separable by the orientation of the leaves, those of R.

cryptantha are ascending, those of M. dubia descending. Dissection of the

ovary of R. cryptantha reveals two intrusive parietal placentae and numerous

ovules, assigning it to Rhaphidophora.

Rhaphidophora in New Guinea, Australia and Western Pacific 103

3. Despite the as yet unconfirmed origin of this plant, we have no hesitation

in publishing it as a new species as it is of such distinctive appearance.

4. The specific epithet alludes to the inflorescences that open hidden or

barely emerging from behind the shingling, substrate-clasping leaves.

Other specimens seen: CULTIVATED. Ex Papua New Guinea, Cult. Kew acc. no. 1983-

4495. Green s.n. (K).

6. Rhaphidophora discolor Engl. & K. Krause

Rhaphidophora discolor Engl. & K. Krause, Bot. Jahrb. Syst. 54 (1916) 80.

— Type: Papua New Guinea, East Sepik Prov., April River, 7 Sept. 1912,

Ledermann 8569 (B & B spirit, holo).

Rhaphidophora oreophila Engl. & K. Krause, Bot. Jahrb. Syst. 54 (1916)

80, synon. nov. — Type: Papua New Guinea, Morobe Prov., Erap

(Erappenberg), 31 Oct. 1912, Ledermann 9590 (B & B spirit, holo).

Figure 7

Medium-sized, robust, pachycaul, homeophyllous (?) liane to 25 m; seedling

stage and pre-adult plants not observed; adult shoot architecture not fully

observed but apparently comprised of elongated, clinging, physiognomically

unbranched, non-flowering stems and free, sympodial, leafy flowering stems;

stems smooth, cataphylls and prophylls caducous, internodes 1—3 x 0.5—1

cm, separated by wide but not especially prominent slightly oblique corky

leaf scars; flagellate foraging stem, clasping roots and feeding roots not

observed; /eaves distichous to very weakly spiro-distichous; cataphylls and

prophylls caducous; petiole deeply canaliculate, 17—37 x 0.4—1 cm, apical

geniculum prominent, basal geniculum large but weakly defined; petiolar

sheath prominent, extending to apical geniculum, slowly degrading to

chartaceous strips and a very few simple fibres, then falling leaving a pale

scar; lamina entire, ovate to oblong-lanceolate or oblong-elliptic or

oblanceolate, slightly oblique, 18—47 x 9—16 cm, coriaceous, base rounded

to oblique and very weakly cordate on one side and slightly decurrent on

the other, apex acute to briefly acuminate, drying adaxially dull olive-

green and abaxially dark brown in the one known collection: midrib

prominently raised abaxially, sunken adaxially; primary venation pinnate,

raised abaxially and adaxially; interprimaries sub-parallel to primaries, only

slightly less prominent, slightly raised abaxially and adaxially; secondary

venation reticulate, slightly raised abaxially and adaxially in dry specimens;

104 Gard. Bull. Singapore 53 (2001)

inflorescence solitary, subtended by a fully developed foliage leaf; peduncle

stout, terete, 9—10 x 1.7—2 cm; spathe canoe-shaped, stoutly acuminate,

20—23 x 2.5—2.5 cm, stiffly very fleshy, marcescent, eventually falling

leaving a very large (c. 1.5 cm wide) scar; spadix cylindrical to slightly

cigar-shaped, slightly curved, long-stipitate, inserted more or less level on

stipe, 13.5—19 x 2.2—4 cm; stipe slender-terete, 1.5—3 x 0.4—1.5 cm; stylar

region rhombohexagonal, 1.75—2.5 x 1.5—3 mm, truncate with a raised

rim; stigma punctiform, flat, c. 0.4 mm diam.; anthers not exserted at male

anthesis; infructescence cylindrical, c. 17 x 2 cm (known from fragments).

Distribution: Indonesian Papua (Yapen), Papua New Guinea (East anf

West Sepik, Milne Bay and Morobe Provinces).

_ Habitat: Lowland swamp forest to upper hill Araucaria forest. 20-900 m

altitude.

Notes: 1. A remarkable and readily identifiable species with a large, very

long-stipitate spadix. Confusion with Rhaphidophora stenophylla (also

flowering on free shoots and with long-stipitate spadix) is possible, although

the latter is readily identifiable by the much narrower leaf laminas, persistent

petiolar sheath not degrading to strips and fibre, smaller (6—9 x 1.5—2.2

cm) spadix and long, pointed style

2. While Nicolson 1475 seems clearly to belong here on morphology, it

occurs in very different habitat (upper hill Araucaria forest) from that of

the type (lowland swamp).

3. Rhaphidophora oreophila Engl. & K. Krause is a smaller manifestation

of this species.

Other specimens seen: INDONESIAN PAPUA. Pulau Yapen, Kamuda, near Serui, Aet &

Idjan s.n. (L). PAPUA NEW GUINEA. West Sepik Prov.: Telefomin, Sandaun, Hak

Valley, contour transect on south bank of Nenem river, opposite community school, Frodin

& Morren 3209 (K):; East Sepik Prov.: Angoram, Latoma village, Wogupmeri river, Leach

NGF 34337 (L):; Morobe Prov.: Araucaria forest plantation 2 miles west of Wau, Nicolson

1475 (B, L, K, US); Milne Bay Prov.: Raba Raba, Mayu River, near Mayu Island, Streimann

NGF 28712 (L, US).

7. Rhaphidophora fortis P.C. Boyce, sp. nov.

Rhaphidophora fortis in Nova Guinea singularis est characteribus

sequentibus concatenatis: folia breviter petiolata, surculi florentes

Rhaphidophora in New Guinea, Australia and Western Pacific 105

Ose.

70s

ISR

oGe.

S27

2-9?

©,

520,

Figure 7. Rhaphidophora discolor Engl. & K. Krause

A. flowering shoot x '/;; B. leaf lamina x '/;;C. venation detail x 4; D. inflorescence, early stages

of fruiting, spathe fallen x */,; E. lower spadix detail, early fruiting x 6; F. upper spadix detail,

early fruiting x 3. All from Streimann & Kairo NGF 39190.

106 Gard. Bull. Singapore 53 (2001)

adhaerentes valde robusti, inflorescentiae ex axilliis foliorum enascentes ut

videtur. — TYPUS: Papua New Guinea, West Sepik Prov., Vanimo

SubProv., Ossima, 27 Jan. 1969, Streimann & Kairo NGF 39190 (K, holo;

L, LAE, CANB, US, iso).

Figure 8

Robust, pachycaul (?), homeophyllous (?) liane to unknown ultimate height;

seedling stage and pre-adult plants not observed; adult shoot architecture

not fully observed but apparently comprised of elongated, clinging,

physiognomically unbranched, flowering stems; stems with epidermis drying

smooth and yellowish, with cataphylls and prophylls degrading to weak

netted fibres and sheets of chartaceous tissue at the stem tips, internodes

1—2.5 x 1.5—2 cm, separated by weakly defined leaf scars; flagellate foraging

_ stem and feeding roots not observed; clasping roots arising from nodes and

internodes, smooth with dark chartaceous epidermis; /eaves distichous;

cataphylls and prophylls very conspicuous, extending almost to apical

geniculum of newly emerging leaves, quickly degrading to netted fibres

and strips of chartaceous tissue; petiole stout, broadly and shallowly

canaliculate, 14—16 x 0.8—1 cm, apical geniculum prominent, basal

geniculum weakly defined and obscured by cataphyll remains; petiolar sheath

very prominent, extending to apical geniculum, but very swiftly degrading

to netted fibres, later falling leaving a smooth, scar; lamina entire, elliptic

to oblong-elliptic or lanceolate elliptic, slightly to markedly oblique, 24—

31 x 8—12 cm, subcoriaceous, base rounded to subacute, apex acute to

briefly acuminate, falcate; midrib raised abaxially, sunken adaxially; primary

venation pinnate, slightly raised abaxially and adaxially; interprimaries

parallel to primaries, less prominent, slightly raised abaxially and adaxially;

secondary venation tessellate-reticulate, slightly raised abaxially and

adaxially in dry specimens; inflorescences not known but apparently arising

two or more sequentially between foliage leaves, subtended by netted

fibre and copious sheet-like tissue; infructscences with peduncle laterally

compressed, c. 5 x 0.6 cm, more or less obscured by cataphyll remains;

spathe not observed; spadix massively ovoid-cylindrical, sessile, inserted

more or less level on peduncle, 8—9 x c. 3 cm; stylar region rounded-

rhombohexagonal, c. 2.3 x 2.7, truncate; stigma punctiform, slightly raised,

c. 0.6 mm diam.; anthers not observed.

Distribution: Papua New Guinea (West Sepik Prov.). Known only from

the type.

Habitat: Lowland forest. 25 m altitude.

Rhaphidophora in New Guinea, Australia and Western Pacific 107

ARO

y} 8

Ny)

iat

inc

\\v

£ GURR

Figure 8. Rhaphidophora fortis P.C. Boyce

A. flowering shoot x '/,; B. leaf lamina x '/,; C. venation detail x 3: D. inflorescence. spathe

fallen x '/,; E. spadix detail, post-male anthesis x 4. F spadix detail, early fruiting x 4. All from

Nicolson 1475.

108 Gard. Bull. Singapore 53 (2001)

Notes: 1. An extraordinary species with extremely stout, clinging flowering

shoots and short-petioled leaves, the whole shoot resembling a plant of

Bornean Scindapsus beccarii Engl.. Inflorescences are borne in the leaf

axils. It is not clear from the material available whether the inflorescence

terminates the main axis which then reiterates, displacing the inflorescence,

or whether the inflorescence is borne on a much abbreviated shoot arising

from the axil.

2. The specific epithet comes from the Latin, fortis, strong or stout, in

allusion to the plant’s relatively massive form.

8. Rhaphidophora geniculata Engl.

Rhaphidophora geniculata Engl., Bot. Jahrb. Syst. 25 (1898) 7; K. Schum.

& Lauterbach, Fl. Schutzgeb. Siidsee (1900) 211; Engl. & K. Krause in

Engl., Pflanzenr. 37 (I1V.23B) (1908) 25, Fig. 7; Engl. & K. Krause, Nova

Guinea 8 (1910) 249. — Type: Papua New Guinea, Madang Prov. (?),

Bismarck Range, 9 Sept. 1896, Lauterbach 827 (B, neo; designated here).

In describing R. geniculata, Engler cited three syntypes [Papua New Guinea

(‘Kaiser Wilhelmsland’), Morobe Prov., Sattelberg, July 1890, Lauterbach

6l6a (By); Papua New Guinea (‘Kaiser Wilhelmsland’), Madang Prov.,

Gogol river, Nov. 1890, Lauterbach 976 (B+) & 1058a (B*)]. The specimen

chosen here as the neotype was determined as R. geniculata by Engler and

matches the protologue well.

Rhaphidophora wentii Engl. & K. Krause, Nova Guinea 8 (1910) 248 &

Nova Guinea 8 (1912) 805, synon. nov. — Type: Indonesian Papua,

Noordfluss, 2 June 1907, Versteeg 1191 (B, holo; BO, L, K, iso).

Figure 9

Rather small, slender, semi-pachycaul homeophyllous liane to 3 m; seedling

stage and pre-adult plants not observed; adult shoot architecture comprised

of elongated, clinging, physiognomically unbranched, leafy, sterile stems

and very short, adherent (always?) flowering stems arising from the axils

of the leaves; stems smooth, terete in cross-section, drying deeply

longitudinally sulcate; with very sparse to copious netted prophyll, cataphyll

and petiolar sheath fibre, internodes 1—4 x 0.5—1.2 cm, separated by

large, oblique, slightly corky leaf scars; flagellate foraging stems absent;

clasping roots arising from the nodes and internodes, pubescent; feeding

roots not observed; leaves distichous; cataphylls and prophylls membranous,

quickly drying and degrading into netted fibres, these later falling; petiole

shallowly grooved, 1.4—27 x 0.2—0.7 cm, apical geniculum long but not

Rhaphidophora in New Guinea, Australia and Western Pacific 109

Figure 9. Rhaphidophora geniculata Engl.

A. flowering shoot x '/;; B. leaf lamina x */,; C. venation detail x 3; D. inflorescence, spathe

removed x 1; E. spadix detail, pre-anthesis x 6; F. stylar region and stigma, side view x 6. All

from Johns 9870.

110 Gard. Bull. Singapore 53 (2001)

especially prominent, basal geniculum barely visible to rather large

(dependent on age), moderately prominent; petiolar sheath extending to

just below the apical geniculum degrading into semi-persistent netted fibres,

eventually falling; Jamina entire, oblanceolate or oblong-elliptic, slightly to

markedly oblique, oblique, 1.8—35 x 3—7 cm, thinly coriaceous to weakly

chartaceous, usually drying conspicuously discolorous, adaxially dark olive-

brown, abaxially pale olive-green to reddish brown, base decurrent, apex

acute, falcate-acute, acuminate or rounded-acuminate; midrib prominently

raised and usually darker abaxially, + flush adaxially; primary venation

arching-pinnate, slightly raised abaxially, almost flush adaxially;

interprimaries sub-parallel to primaries, slightly to much less prominent,

slightly raised; secondary venation weakly reticulate, slightly raised; tertiary

venation reticulate-striate, barely visible; inflorescence solitary, terminating

a short shoot arising in the axils of leaves, occasionally arising from axils of

' fallen leaves, and usually inflorescences arising from several adjacent leaves,

and thus sections of stems carrying several inflorescences in diffuse clusters,

each inflorescence subtended by a prophyll and several degraded, netted

cataphylls; peduncle strongly compressed-terete, 5—9 x 0.3—0.6 cm; spathe

ovoid-ellipsoid, base decurrent and oblique, apex rounded, slender-

acuminate, 3.5—6.5 x 1.5—2.5 cm, white, marcescent falling at fruit

maturation leaving a large oblique scar; spadix ovoid-ellipsoid, stipitate,

inserted obliquely on stipe, 2.5—3.5 x 1.3—1.7 cm; stipe 2—8 x 1—2 mm;

stylar region mostly hexagonal, 1.5—2 x 2.1—2 mm, prominently conical;

stigma punctiform on a long (c. 2 mm) stipe, c. 0.2—0.35 diam.; anthers not

exserted at male anthesis. /nfructescence stout ellipsoid-cylindrical, c. 4 x 2

cm.

Distribution: New Guinea. Widespread but, in view of the rather few

collections, perhaps uncommon.

Habitat: Primary and slightly disturbed lowland to upper hill forest and

lower montane forest, 150-790 m altitude.

Notes: 1. In flowering on clinging shoots and by the conical style

Rhaphidophora geniculata resembles R. australasica from which it may be

distinguished by the ovoid-ellipsoid, stipitate spadix (cylindrical and sessile

in R. australasica) and the lack of fibrous petiolar sheath remains. It also

differs from that species by the inflorescences borne on short shoots arising

from the leaf axils. In this respect it is reminiscent of R. brevispathacea and

R. stolleana, although in these the stems are pendent and not clinging.

2. The protologue of R. wentii states the spadix to be sessile; examination

Rhaphidophora in New Guinea, Australia and Western Pacific 111

of the fertile types (B, BO, L) shows this to be incorrect; all are stipitate.

The illustration accompanying the protologue of R. geniculata is a good

match for the B holotype and L isotype of R. wentii.

Other specimen seen: INDONESIAN PAPUA. Digul Prov.: Merauke, Branderhorst 324

(L); Mimika Prov.: Freeport Concession Area, Golf Course surrounds, Johns 9970 (BO, K,

MAN): Kali Kopi, between Kali Kopi levee and the Kopi River, Ufteridge 79 (BO, K,

MAN). PAPUA NEW GUINEA. West Sepik Prov.: Vanimo, Vanimo hinterland, Streimann

LAE 52964 (LAE, US): Southern Highlands Prov.: Mt Bosavi, northern side, 2—4 km west

of Ludesa mission station, Jacobs 9331 (L); Wasu, NE slope of Mt Bosavi, Moi et al. 196

(BFC, L, LAE); Morobe Prov.: vicinity of Kajabit Mission, Clemens 10662 (GH): Vicinity

of Lae, along logging road to Busu River, from 3.8 km E of Igam road and Military Base to

c. 6 km up the road junction, Croat 52787 (MO), Croat 52788 (MO); Left off of Igam road

past Military Reserve. Croat 52803 (MO).

9. Rhaphidophora gorokensis P.C. Boyce, sp. nov.

Rhaphidophora gorokensis R. kokodensi R. okapensi, R. pilodi atque R.

wariae similis, species omnes monticolae foliis parvis rigide coriaceis sunt.

A R. waria atque R. okapensi in surculis liberis lateralibus florenti et spadice

sessili prompte distinguibilis est; porro a R. waria vaginis petiolaribus plus

minusve omnio cadentibus neque in fibris fatiscentibus, atque a R. okapensi

forma laminae foliae (anguste lanceolate vel elliptics, basi acuta, apice

longe acuminata tubulo minuto instructa, neque ovata, basi ovata, apice

longe acuminata tubulo manifeste) distinguitur. A R. pilode atque R.

kokodensi (ambo in surculis liberis lateralibus florentes) R. gorokensis

fibra vaginae petiolaris carenti distinguitur. — TYPUS: Papua New Guinea,

Eastern Highlands Prov., Goroka, Marafunga logging area, Upper Asaro

Valley, 6 Sept. 1961, Womersley & Sleumer NGF 13971 (GH, holo; L,

EAE, iso).

Figure 10

Small, homeophyllous (?) liane to 12 m; seedling stage and pre-adult plants

not observed; adult shoot architecture not fully observed but presumably

comprised of elongated, clinging, physiognomically unbranched, leafy, non-

flowering stems and (observed), free, little branching, sympodial, densely

leafy flowering stems; stems terete in cross-section, without cataphyll,

prophyll and petiolar sheath fibre, internodes to 1—4 x 0.5—0.8 cm,

separated by slightly oblique, corky leaf scars; flagellate foraging stems,

clasping roots and feeding roots unknown; leaves spiro-distichous on free

shoots; cataphylls and prophylls chartaceous, caducous; petiole deeply

canaliculate, 6.5—11 x 0.15—0.25 cm, smooth, genicula weak; petiolar sheath

prominent, chartaceous, extending to apical geniculum, briefly persistent,

112 Gard. Bull. Singapore 53 (2001)

then partially caducous, later falling leaving an irregular, corky scar; lamina

entire, narrowly lanceolate to elliptic, oblique, falcate, 7—16 x 2—5 cm,

stiffly coriaceous, base acute, oblique, apex long-acuminate with a minute

tubule; midrib raised abaxially, flush to slightly sunken adaxially; primary

venation densely pinnate, raised abaxially and adaxially; interprimaries sub-

parallel to primaries, indistinguishable from them; secondary and tertiary

venation conspicuously parallel-reticulate; inflorescence solitary, subtended

by an under-developed foliage leaf; peduncle compressed-terete, 5—7 x

0.2—0.2.5 cm; spathe narrowly canoe-shaped, stout-acuminate, 6—6.5 x c.

1.5 cm, marcescent(?); spadix cylindrical, sessile, 3.3—3.5 x c. 0.7 cm; stylar

region rhombohexagonal, c. 0.9—1 x 0.8—1.1 mm, truncate; stigma

punctiform, flush with a raised rim, c. 0.5 mm diam.; anthers not exserted

at male anthesis; infructescence not observed.

- Distribution: Papua New Guinea (Eastern Highlands Prov. - Goroka and

Mendi).

Habitat: Lower montane rain forest, Fagaceae and Nothofagaceae forest

on limestone. 1600-2440 m altitude.

Note: Similar to Rhaphidophora kokodensis, R. okapensis, R. pilosa and R.

waria in being a montane species with small, stiffly coriaceous leaves. It is

readily distinguishable from R. okapensis and R. waria by flowering on

free lateral shoots and by the sessile spadix. It can be further distinguished

from R. waria by the petiolar sheaths falling more or less entire and not

degrading into fibres, and from R. okapensis by the shape of the leaf

lamina (narrowly lanceolate to elliptic, base acute, apex long-acuminate

with a minute tubule v. ovate, base cordate, apex long acuminate with a

pronounced tubule). From R. kokodensis and R. pilosa (both flowering on

free lateral shoots) R. gorokensis is distinguished in lacking petiolar sheath

fibre.

Other specimens seen: PAPUA NEW GUINEA. Southern Highlands Prov.: Mendi, Det

Mission, 16 km SSW of Mendi, Vinas 151 (GH, K); Eastern Highlands Prov.: Goroka,

Collins Mill, Omahaiga River valley, Mt Otto area, Robbins 870 (L).

10. Rhaphidophora guamensis P.C. Boyce, sp. nov.

Dum Rhaphidophoram spathaceum simulans, R. guamensis inflorescentia

solitaria apicem surculi liberi lateralis ferenti neque e mole prophyllorum

chartaceorum cataphyllorumque exorienti, etiam specice robuste stipitato

statim distinguibilis est. Praeterea, stipes stigmaticus egregie longus insignis

Rhaphidophora in New Guinea, Australia and Western Pacific 113

tty

Sat

Pies

ert

‘ E

all

L GUAL

Figure 10. Rhaphidophora gorokensis P.C. Boyce

A. flowering shoot x '/,; B. leaf lamina x ”/;; C. leaf lamina x ’/,; D. venation detail x 4; E.

inflorescence x 1; F. spadix detail, pre-anthesis x 8. A & B, D-F from Womersley & Sleumer

NGF 13971:C from Robbins 870.

114 Gard. Bull. Singapore 53 (2001)

est. — TYPUS: Guam, slopes of Mt Almagosa, Naval Magazine, near

Fena Dam, 28 April 1962, B.C. Stone 4101 (L, holo; GUAM, US, iso).

Figure 11

Moderately robust, medium-sized semi-leptocaul, heterophyllous(?) liane

to unknown ultimate height; seedling stage not observed; pre-adult plants

forming scattered terrestrial colonies; adult shoot architecture comprised of

elongated, clinging, physiognomically unbranched, scattered-leafy, non-

flowering stems and free, sympodial, leafy flowering stems; stems smooth,

mid-green, cataphylls and prophylls briefly persistent then falling leaving

bare stems, internodes 0.5—3 x 0.3—1 cm, separated by prominent straight

leaf scars; flagellate foraging stem not observed; clasping roots solitary to

somewhat densely produced arising from the nodes and internodes adjacent

. to nodes; feeding roots not observed; /eaves more-or-less distichous;

cataphylls and prophylls drying chartaceous, briefly persistent; petiole deeply

canaliculate, 9—15 x 0.2—0.4 cm, smooth, apical geniculum large but not

especially prominent, basal geniculum almost invisible; petiolar sheath

extending to base of apical geniculum, broad, chartaceous, short-persistent,

degrading to very sparse papery strips, then falling; /amina entire, oblong-

elliptic to oblong-lanceolate or oblanceolate, slightly oblique to markedly

oblique, 15—32 x 4—8.5 cm, submembranous to coriaceous, base rounded

to acute, apex acute to weakly acuminate; midrib raised abaxially, + flush

adaxially to weakly raised abaxially; primary venation pinnate, slightly raised

abaxially, flush adaxially, drying darker than lamina; interprimaries sub-

parallel to primaries, much less prominent, slightly raised abaxially, barely

visible adaxially; secondary venation reticulate, raised, especially notable

in dry material; inflorescence solitary, subtended by a fully to partially

developed foliage leaf and one to several degraded and soon-falling

chartaceous cataphylls; peduncle rather stout, terete, 6—8 x 0.3—0.5 cm;

spathe broadly canoe-shaped, briefly stout-beaked, 10—12 x 2—7 cm,

spongy-fleshy, very thick-walled, pale yellow, caducous, falling leaving a

very prominent oblique scar; stipe stoutly terete, 1—1.5 x 0.4—0.5 cm;

spadix slender to somewhat stout-cylindrical, stipitate, cochleate at insertion

on stipe, 7—9 x 1—1.5 cm, white at male anthesis; stylar region strongly

conical, mostly hexagonal in top view, 0.9—1.2 x 1—1.1 mm; stigma

punctiform on the tip of a long (c. 2 mm) stipe, 0.5—0.2 x c. 0.3 mm, glossy

and black in dried material; anthers not exserted at male anthesis;

infructescence not observed.

Distribution: Guam.

Habitat: Mixed forest, coconut plantation, on limestone. c. 400 m altitude.

Rhaphidophora in New Guinea, Australia and Western Pacific 115

Ns er Oy

Figure 11. Rhaphidophora guamensis P.C. Boyce

A. flowering shoot x '/,; B. leaf x '/;; C. pre-adult creeping shoot x '/;: D. spadix detail at post

male anthesis x 6. All from Stone 4101.

116 Gard. Bull. Singapore 53 (2001)

Note: While resembling Rhaphidophora spathacea, R. guamensis 1s

immediately distinguishable by the solitary inflorescence carried on the tip

of a free lateral shoot, and which does not emerge from a mass of

chartaceous prophyll and cataphylls, and also by the robustly stipitate spadix.

In addition, the remarkably long stigmatic stipe is notable.

Other specimens seen: GUAM. Mt Lamlam summit, near lighthouse, Anderson 147 (GUAM,

US); Almagosa Springs area, Naval Magazine, Rinehart 15568 (GUAM, US).

11. Rhaphidophora hayi P.C. Boyce & Bogner

Rhaphidophora hayi P.C. Boyce & Bogner, Gard. Bull. Singapore 52 (2000)

91, fig.1. — Type: Australia, Queensland, Cooroo Lands, north Johnstone

River, near Innisfail, Nov. 1965, Webb & Tracey 7066 (BRI, holo).

Figure 12

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, flowering stems; stems

rectangular to 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; /eaves distichous, shingling and ascending 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, caducous; petiole deeply grooved,

1—2 x 0.2—0.3 cm, smooth, apical and basal genicula barely visible; petiolar

sheath prominent, caducous but adhering to stem, membranous, ligulate,

margins of ligule fused, the ligule extending up to 3 cm above base of

lamina and enclosing shoot apex; /amina broadly to narrowly ovate-elliptic,

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;

Rhaphidophora in New Guinea, Australia and Western Pacific wily

Mt | Wi

YY,

INN

Figure 12. 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 anthesis x 10; H. spadix detail at post-male anthesis x 10. A-C from Backer

11199; D-H from Sands et al. 2384.

118 Gard. Bull. Singapore 53 (2001)

interprimaries sub-parallel to primaries, slightly raised on both leaf surfaces;

secondary venation reticulate, slightly raised abaxially, + flush adaxially;

inflorescence solitary, subtended by a membranous, caducous prophyll and

one or more similar cataphylls; 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 anthesis and then slowly falling

leaving a large scar at the base of the spadix; spadix stoutly cigar-shaped,

shortly stipitate, inserted + level on stipe, 3.5—6 x 1—1.2 cm, yellow; stipe

4—6 x 3—3.5 mm; stylar region weakly developed, mostly irregularly

rhombohexagonal, 1.1—1.3. x 1—1.1 mn, truncate; stigma prominently

raised, elongated, longitudinally orientated, c. 0.3—0.5 x 0.2—0.4 mm;

anthers not exserted at male anthesis; infructescence not seen.

_ Distribution: Indonesian Papua, Papua New Guinea (including New Britain,

New Ireland, Bougainville and Muyua (Woodlark) Island), and Australia

(eastern tropical Queensland).

Habitat: Primary and secondary monsoon or rain forest on coralline

limestone and basalt. 20-600 m altitude.

Notes: 1. While resembling Rhaphidophora pachyphylla, 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 occurs also in R. cryptantha (q.v.).

2. Very similar to R. parvifolia Alderw. (Maluku: Pulau Ternate) but

differing by the stipitate spadix. From available material it is not possible

to tell if R. parvifolia has the disarticulating side shoots typical of R. hayi.

3. There are many collections of R. hayi originating from Australia (mostly

in BRI and ORS) but all except those cited here are sterile.

Other specimens seen: INDONESIAN PAPUA. Kepala Burung Prov.: 2 km N of Manokwari,

Nicolson 1577 (B, K, L, US). PAPUA NEW GUINEA.Woodlark (Muyua) Island:

Kulumadau, Brass 28831 (GH, L); Central Prov.: Sogeri Plateau, 5 - 7 miles beyond Kokoda

Trail Monument, 30 miles east of Port Moresby, Nicolson 1431 (L, US); Milne Bay Prov.:

Esa’ala , Normanby Island, Sewa Bay, 21 Oct. 1971, Lelean & Streimann LAE 52541 (L,

LAE, US); North Solomons Prov.: Bougainville, Arawa, McKillup’s Plantation, 6 m west of

Kieta, Nicolson 1512 (B, K, US); West New Britain Prov.: 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 et al. 2384 (K, K

Rhaphidophora in New Guinea, Australia and Western Pacific 119

(living collection acc. no. 1975-5026, K spirit no. 63938). AUSTRALIA. Queensland: Cape

Tribulation, Rijkers 1484 (BRI).

12. Rhaphidophora intonsa P.C. Boyce, sp. nov.

Rhaphidophora intonsa apicibus surculorum reticulo fibroso denso inclusis

distinguitur, porro inflorescentiae grandes in surculis liberis diagnostici

sunt. Cum R. spuria eam confundere potest, sed haec basi laminae truncata

vel leniter cordata atque apicibus surculorum multo minus fibrosis facile

distinguitur. Cum R. australasica eam etiam confundere potest, sed ab illa

R. intonsa in surculis liberis florenti et inflorescentias multo maiores (spatha

17—20 cm nec 5—9.5 cm, spadice 10.5—18.5 cm nec 3.75—8 cm) procreanti

differt; praetera, apex regionis stylaris truncata est, neque manifeste conica.

— TYPUS: Papua New Guinea, Central Prov., Boridi, 30 Sept. 1935, Carr

14313 (SING, holo; BM, L, iso).

Figure 13

Medium to large, robust, pachycaul (?), homeophyllous (?) liane to 12 m;

seedling stage and pre-adult plants not observed: adult shoot architecture

not fully observed but apparently comprised of elongated, clinging,

physiognomically unbranched, non-flowering stems and free lateral, leafy

flowering stems; stems with cataphylls and prophylls degrading to dense

ragged fibres and sheets of tissue and forming dense matting at the stem

tips, internodes 1—3 x 0.5—1.4 cm, separated by prominent slightly oblique

corky leaf scars; flagellate foraging stem, clasping roots and feeding roots

not observed; /eaves spiro-distichous; cataphylls and prophylls quickly

degrading to dense ragged fibres and sheets of tissue; petiole shallowly

canaliculate, 12—43 x 0.4—0.9 cm, apical geniculum prominent, basal

geniculum weakly defined and obscured by fibre; petiolar sheath very

prominent, extending to apical geniculum, swiftly degrading to copious

netted fibres, later falling leaving a smooth, corky scar; /amina entire,

ovate to oblong-elliptic, slightly oblique, 17—47 x 9—20 cm, subcoriaceous,

base subacute to slightly decurrent, apex acute to briefly acuminate; midrib

prominently raised abaxially, very slightly raised adaxially:; primary venation

pinnate, slightly raised abaxially and adaxially; interprimaries parallel to

primaries, slightly less prominent, raised abaxially, slightly impressed

adaxially; secondary venation reticulate, slightly raised abaxially and

adaxially in dry specimens; inflorescence solitary, mostly subtended by a +

fully developed foliage leaf and copious netted fibre and sheet-like tissue;

peduncle laterally compressed to terete, 9—13 x 0.9—2 cm: spathe canoe-

shaped, 17—20 x 1.5—2.5 cm, stoutly long-beaked, stiffly fleshy, caducous

leaving a large scar at the base of the spadix: spadix cylindrical, slightly

120 Gard. Bull. Singapore 53 (2001)

curved, sessile, inserted more or less level on peduncle, 10.5—18.5 x 1.7—

2.3 cm, cream; stylar region rhombohexagonal, 1—2 x 1—1.5 mm, truncate;

stigma punctiform, raised at male anthesis flattened in dry material, c. 0.3

mm diam.; anthers not exserted at male anthesis; infructescence curved-

cylindrical, c. 17 x 2 cm.

Distribution: Papua New Guinea (Central & Morobe Prov.).

Habitat: Montane forest. 1290-1800 m altitude.

Notes: 1. The densely netted fibre-encased shoot tips and large inflorescences

are diagnostic. Nicolson 1473 is smaller and less robust than the type but

otherwise matches very well.

2. Confusion with R. spuria is possible although the latter is readily

distinguished by the truncate to weakly cordate lamina base and in having

very much less fibre at the shoot tips. Confusion with R. australasica is

possible. Rhaphidophora intonsa differs by flowering on free shoots and in

producing much larger inflorescences (spathe 17—20 cm/spadix 10.5—18.5

cm vs. spathe 5—9.5 cm/spadix 3.75—8 cm in R. australasica). Additionally,

the stylar region is truncate-topped in R. intonsa and prominently conical

in R. australasica.

3. The specific epithet comes from the Latin intonsa, unshaven, in allusion

to the dense, untidy prophyll and cataphyll fibre clothing the stem tips.

Other specimens seen: PAPUA NEW GUINEA. Morobe Prov.: Ogeramnang to Malang,

Clemens 4637 (GH); Sambangan, Clemens 7779 (B): Edie Creek road, above Wau, Nicolson

1473 (B, K, L, P, US).

13. Rhaphidophora intrusa P.C. Boyce, sp. nov.

Dum Rhaphidophoram schlechteri simulans, R. intrusa apice spathae longe

extenso (usque ad tertiam partem longitudinis spathae toto) atque ligulis

vaginae petiolaris valde elongatis secundum costam abaxialem usque ad

dimidiam longitudinis laminae extensis et secum adnatis statim cognoscibilis.

Hic character etiam, in spatione minore, in R. hayi repertus est, a qua R.

intrusa habitu non scindulanti, petiolis longioribus, lamina foliorum tenuiore

atque longiore differt. — TYPUS: Indonesian Papua, Kepala Burung Prov.,

Kabupaten Manokwari, Kecamatan Manokwari, Arfak Plains, close to road

from SP & to Sg. Wariori crossing, 22 April 1994, Sands 6276 (K, holo;

BO, MAN, iso).

Figure 13. Rhaphidophora intonsa P.C. Boyce

A. flowering shoot x '/,; B. leaf lamina x '/,; C. venation detail x 2: D. portion of clinging adult

stem x '/,; E. inflorescence, spathe fallen x !/,: F. spadix detail, male anthesis x 6; F A-C,E & F

from Carr 14313; D from Nicolson 1473.

122 Gard. Bull. Singapore 53 (2001)

Figure 14

Slender, leptocaul, homeophyllous liane to 5 m; seedling and pre-adult

plants not observed; adult shoot architecture comprised of elongated,

clinging, physiognomically unbranched, leafy, non-flowering stems and

somewhat lengthened mostly unbranched, free, sympodial, spreading to

pendent, rarely ascending, leafy, flowering stems; stems smooth, flexuous,

climbing stems + terete, free stems somewhat longitudinally flattened,

without prophyll, cataphyll and petiolar sheath fibre, internodes 1—6 x 0.6

cm on clinging shoots, those on free shoots much more slender, separated

by weak straight to slightly oblique leaf scars, older stems woody; flagellate

foraging stems not observed; clasping roots arising sparsely from nodes of

clinging stems, slightly pubescent; feeding roots not observed; leaves

distichous, somewhat scattered; cataphylls and prophylls not observed;

' petiole shallowly to rather deeply grooved adaxially, 2.5—5 x 0.1—0.2 cm,

smooth, with very slight apical and basal geniculum; petiolar sheath

prominent, chartaceous, extending beyond the apical geniculum by two

ligules adaxially and by ligules fused along the abaxial midrib for up to

half the leaf lamina length, sheath eventually falling leaving a continuous

scar from the petiole base, around the top of the apical geniculum and

back to the base and remnants along the abaxial midrib; /amina entire,

oblong-elliptic, 6—16.5 x 3.5—6 cm, submembranous, base subovate, apex

weakly acuminate, with a minute tubule; midrib raised abaxially, slightly

sunken adaxially in fresh material, slightly sunken abaxially and adaxially

in dry material; primary venation pinnate, slightly raised on both surfaces

in dried material; interprimaries sub-parallel to, but much less distinctive

than, primaries; secondary venation very feebly reticulate, almost invisible;

inflorescence solitary, subtended by a fully developed foliage leaf; peduncle

slender-terete, 3.5—4 x 0.15 cm; spathe cigar-shaped, cuneate basally at

insertion on petiole, apex truncate and extending into a very long slender

beak up to '/3 length of entire spathe, 7—8 x 1.3—1.5 cm, thick fleshy,

marcescent(?), later pushed off by developing infructescence(?) leaving a

large scar; spadix cigar-shaped, stipitate, inserted level on stipe, 4.5—5.5 x

0.9—1.2 cm, obtuse; stipe slender terete, c. 1 x 0.2 cm; stylar region dorso-

ventrally compressed-rhombohexagonal, 0.9—1.5 x 0.85—1.2 mm, truncate;

stigma punctiform, c. 0.45 mm diam., prominent in dried material; anthers

not exserted at male anthesis; infructescence stoutly cigar-shaped, c. 5 x 1.5

cm.

Distribution: Indonesian Papua (Kepala Burung Prov.). Known only from

the type.

Rhaphidophora in New Guinea, Australia and Western Pacific 123

Figure 14. Rhaphidophora intrusa P.C. Boyce

A. flowering shoot x ?/;; B. leaf lamina x ?/;; C. inflorescence x 7/3; D. inflorescence, spathe

partly removed to show spadix x 7/;. E. spadix detail, male anthesis x 6. All from Sands 6276.

124 Gard. Bull. Singapore 53 (2001)

Habitat: Partially disturbed lowland rain forest. Sea level to 20 m altitude.

Notes: 1. While resembling Rhaphidophora schlechteri, R. intrusa is

immediately recognizable by the long extended spathe apex (up to '/3 the

length of the entire spathe) and by the greatly elongated petiolar sheath

ligules that extend fused along the abaxial midrib for up to half the leaf

lamina length. The latter character is also found, to a much lesser extent,

in R. hayi, from which R. intrusa differs in the non-shingling habit, longer

petioles and thinner, longer leaf lamina.

2. The specific epithet comes from the Latin, intrusus, to intrude, in allusion

to the extraordinary ligules described above.

. 14. Rhaphidophora jubata P.C. Boyce, sp. nov.

Rhaphidophora jubata species valde distincta est magnitudini grandi,

pedunculo longo, apicibus stylaribus parvis stigmatibus prominentibus atque

seminibus cristatis reniformibus notabilis (his adhuc Rhaphidophorae non

perscriptis). Cum R. spuria eam confundere potest, sed haec stylares maiores

(1.6—2.4 x c. 2 mm diametro) atque stigmata punctiforma vel leviter

ellipsoidea non manifeste elevata neque siccitate nigra seminaque

ellipsoidea ecristata habet. — TYPUS: Papua New Guinea, North Solomons

Prov., Bougainville, path from Arawa to Korpei, 10 miles southwest of

Kieta, 1 Nov. 1964, Nicolson 1529 (L, holo; B, K, P, US, iso).

Figure 15

Very robust, pachycaul, homeophyllous liane to 10 m; seedling stage not

observed; pre-adult plants forming extensive terrestrial populations; adult

shoot architecture comprised of elongated, clinging, physiognomically

unbranched, densely leafy, flowering; stems smooth, with cataphylls and

prophylls degrading into weak slightly fibrous patches and then falling,

internodes to 4—13 x 1—3.5 cm; flagellate foraging stem not observed;

clasping roots not observed; feeding roots arising singly from nodes, robust;

leaves spiro-distichous; cataphylls and prophylls subcoriaceous, degrading

to weakly fibrous patches, then falling; petiole deeply canaliculate, (6—)

40—85 x 0.35—0.5 cm, smooth, apical and basal genicula very large though

not especially prominent; petiolar sheath very prominent, extending from

between */, petiole length up to the apical geniculum, short-persistent,

falling leaving thin scar; lamina entire, ovate to oblong-ovate, slightly

oblique, 21.5—90 x 10.5—35 cm, thinly coriaceous to submembranous,

drying pale yellow-brown to brown, base oblique, broadly rounded to

Rhaphidophora in New Guinea, Australia and Western Pacific 125

‘a4

4),

(UF

Ae)

ling 4

ja)

Liven Wicaina'

Gil

LGURR E 5

Figure 15. Rhaphidophora jubata P.C. Boyce

A. flowering shoot x '/;.; B. leaf lamina x 7/,; C. venation detail x 2; D. inflorescence, plus lower

peduncle x */,; E. spadix detail, pre-anthesis x 6; F. mature seed, side view x 20. A from Nicolson

1496; B-C from Nicolson 1530; D-F from Nicolson 1529.

126 Gard. Bull. Singapore 53 (2001)

shallowly and minutely cordate, more rarely subacute, apex rounded-

acuminate with a small apical tubule; midrib very prominently raised and

abaxially, + impressed adaxially; primary venation pinnate, raised abaxially

and adaxially; interprimaries parallel to primaries, much less prominent;

secondary venation reticulate, hardly visible; inflorescence several together,

each subtended by a prophyll and one or more chartaceous cataphylls,

these soon degrading; peduncle terete, 11—25 x 0.3—2 cm; spathe slender

canoe-shaped, stoutly attenuate-beaked, 16—24 x 2.5—3 cm, stiffly thin-

fleshy, caducous, yellow; spadix slender cylindrical, sessile but long decurrent

(to 2 cm) on peduncle/spathe insertion, 13—21 x 1—3 cm; stylar region,

weakly rhombohexagonal to trapezoid, 1—1.2 x 1—1.3 mm, truncate; stigma

punctiform, prominently raised, c. 0.25 mm diam., drying glossy black;

anthers not exserted at male anthesis; infructescence slender cylindrical,

- 14.5—17.5 x 2.5 cm; seed reniform, laterally compressed, conspicuously

crested along dorsal side (i.e., the side opposed to the insertion of the

funicle), c. 1.2 x 2.5 mm at maturity, c. 8—12 per fruit arranged on two

parietal placentae.

Distribution: Papua New Guinea (New Britain and Bougainville).

Habitat: Primary and secondary lowland to upper hill forest, along streams

and paths. 90-980 m altitude.

Notes: 1. A very distinct species notable for its overall large size, long

peduncle and small stylar tops with prominently raised stigmas. Confusion

with Rhaphidophora spuria is possible although the latter has larger stylar

tops (1.6—2.4 x c. 2 mm diam.) and punctiform to slightly ellipsoid stigmas

that are not prominently raised and do not dry glossy black, and ellipsoid

seeds lacking a crest.

2. The specific epithet is derived from the Latin, jubatus, crested, in allusion

to the crested reniform seeds of this species, a form of seed not hitherto

recorded for Rhaphidophora. This seed form is common in Epipremnum

Schott, which differs from Rhaphidophora by having 4 (— 6) seeds at base

of a single intrusive parietal placenta. Furthermore, the testa in

Rhaphidophora (including R. jubata) is brittle, while that of Epipremnum

is tough and bony.

Other specimen seen: PAPUA NEW GUINEA. North Solomons Prov.: Bougainville, path

up Dakao Creek near Korpei, 11 miles southwest of Kieta, Nicolson 1530 (B, K, L, P, US);

Pavairi, Ridsdale & Lavarack NGF 30628 (LAE, US); Kapikavi, Ridsdale & Lavarack

NGF 31600 (LAE, US); East New Britain Prov.: Kareeba road, 2 miles west of Kerevat,

Nicolson 1496 (US).

Rhaphidophora in New Guinea, Australia and Western Pacific bay

15. Rhaphidophora kokodensis P.C. Boyce, sp. nov.

Rhaphidophora kokodensis R. australasicae similis videtur sed in surculis

liberis lateralibus florenti, stylis apice planis, et foliis minoribus angustoribus

magis coriaceis prompte sejuncta. Porro species montana est igitur

altitudinaliter sejuncta. Cum R. pilode (etiam in surculis liberis lateralibus

florenti) eam confundere potest, quamquam R. kokodensis non fibras

aspectu coacto proprio ut in R. pilode habet. — TYPUS: Papua New

Guinea, Central Prov., Kokoda , eastern side of Lake Myola no. 1, 23 July

1974, Croft et al. LAE 61974 (GH, holo; BRI, CANB, L, LAE, iso).

Figure 16

Small, homeophyllous (?) liane to unknown ultimate height; seedling stage

and pre-adult plants not observed; adult shoot architecture comprised of

elongated, clinging, physiognomically unbranched, leafy, non-flowering

stems and long, moderately elaborated, free, sympodial, densely leafy

flowering stems; sfems terete in cross-section, internodes to 1—4 x 0.5—0.8

cm, separated by slightly oblique, prominent leaf scars; flagellate foraging

stems, clasping roots, and feeding roots unknown; leaves spiro-distichous

on free shoots; cataphylls and prophylls chartaceous but very soon degrading

into long tough fibres; petiole deeply canaliculate, 5—16 x 0.2—0.25 cm,

smooth, apical geniculum quite well defined, basal geniculum weak; petiolar

sheath prominent, chartaceous, extending to apical geniculum, very swiftly

degrading to fibres; /Jamina entire, narrowly lanceolate to narrowly

lanceolate-elliptic, occasionally slightly falcate, 13—-25 x 1.75—6 cm, stiffly

coriaceous, base acute, apex acuminate to long-acuminate with a minute

tubule; midrib slightly raised abaxially, slightly sunken adaxially; primary

venation densely pinnate, slightly raised abaxially and adaxially:

interprimaries parallel to primaries indistinguishable from them; secondary

and tertiary venation very faintly reticulate; inflorescence solitary, subtended

by an underdeveloped foliage leaf and much degraded cataphyll fibre;

peduncle compressed-terete, 7—12 x 0.2—0.3 cm; spathe narrowly cigar-

shaped, long stout-acuminate, (4—)8—9 x c. 1.3 cm, marcescent(?): spadix

cylindrical, sessile, 2—6 x 0.8—1 cm; stylar region rhombohexagonal, c.

0.9—1 x 0.8—1.1 mm, truncate; stigma punctiform, very slightly raised, c.

0.4 mm diam.; anthers not exserted at male anthesis; infructescence

cylindrical, c. 11 x 1.3 cm.

Distribution: Papua New Guinea (Central Prov. - Kokoda and Port

Moresby).

‘128 Gard. Bull. Singapore 53 (2001)

Habitat: Submontane rain forest on dark brown loam. 1500-2000 m altitude.

Note: Superficially similar to Rhaphidophora australasica but readily

separated by its flowering on free lateral shoots, by the flat-topped styles

and the smaller, narrower, much more coriaceous leaves. In addition, being

a moniane species, R. kokodensis is separated altitudinally. Confusion with

R. pilosa (also flowering on free lateral shoots) is possible, although R.

kokodensis has petiole and shoot fibres without the distinctive felted

appearance of those of R. pilosa.

Other specimens seen: PAPUA NEW GUINEA. Central Prov.: Port Moresby, Boridi, Carr

13237 (BM, K, L, SING), Carr s.n. (BM); East slope of Lake Myola No. 2, Croft & Lelean

NGF 34561 (GH, K, L).

16. Rhaphidophora korthalsii Schott

Rhaphidophora korthalsti 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; Alderw., Bull. Jard.

Bot. Buitenzorg III, 4 (1922) 341; Hay, Aroids of Papua New Guinea, pl.

XV, a—c. — Type: Indonesia, Java, P.W. Korthals s.n. (L, holo; L, P, iso).

Pothos celatocaulis N.E. Br., Gard. Chron. 13 (1880) 200. — Rhaphidophora

celatocaulis (N.E. Br.) Alderw., Bull. Jard. Bot. Buitenzorg III, 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; Beccari,

Malesia 1 (1882) 271, Tab. xx 1—5; K. Schum. & Lauterbach, Fl. Schutzgeb.

Siidsee (1900) 211; Engl. & K. Krause in Engl., Pflanzenr. 37 (IV.23B)

(1908) 48—49; K. Krause & Alderw., Nova Guinea 14 (1924) 214. — Type:

Sarawak, G. Gading, July 1866, Beccari PB 2314 (FI, lecto, selected by

Boyce, 1999).

Rhaphidophora tenuis Eng]., Bot. Jahrb. Syst. 1 (1881) 181; 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 var. angustiloba Ridl. ex Engl. & K. Krause in

Engler, Pflanzenr. 37 (IV.23B) (1908) 49. — Type: Malaysia, Sarawak,

Matang, July 1903, Ridley s.n. (SING, lecto; selected by Boyce, 1999).

Rhaphidophora in New Guinea, Australia and Western Pacific 129

L. GUAR

Figure 16. Rhaphidophora kokodensis P.C. Boyce

A. flowering shoot x !/;; B. leaf lamina x >; C. leaf lamina x '/,; D. venation detail x 3: E.

inflorescence, spathe fallen x 2/3; F. spadix detail, post-male anthesis x 4. A & D from Croft et

al. LAE 61974; B-C, E-F from Carr 13237.

130 Gard. Bull. Singapore 53 (2001)

Rhaphidophora copelandi Engl., Bot. Jahrb. Syst. 37 (1905) 115; Engl. &

K. Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 49. — Type: Philippines,

Mindanao, Davao, Mt Apo, April 1904, Copeland 1193 (B holo; PNH

isot).

Monstera latevaginata Engl. & K. Krause in Engl., Pflanzenr. 37 (I1V.23B)

(1908) 11. — Type: Cult. Bot. Gard. Berlin, Engler & Krause s.n. (B, holo).

Rhaphidophora grandifolia K. Krause, Bot. Jahrb. 44, Beibl. 101 (1910) 11.

— Type: Philippines, Negros, Negros Orientale, Dumaguete (Cuernos Mts),

March 1908, Elmer 9464 (B, holo; E, K, L, LE, MO, PNH? iso).

Rhaphidophora trinervia Elmer, Leafl. Philipp. Bot. 8 (1919) 3073. — Type:

_ Philippines, Laguna, Los Banos (Mt Maquiling), June—July 1917, Elmer

18057 (PNH, holojsFip K, EL, MO; Piso):

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).

Rhaphidophora latifolia Alderw., Bull. Jard. Bot. Buitenzorg III, 4 (1922)

341; K. Krause & Alderw., Nova Guinea 14 (1924) 213. —Type: Indonesian

Papua, Bonggo Range, Mamberamo, Pionierbivak, 23 July 1920, Lam 711

(BO, holo; L, iso).

Rhaphidophora palawanensis Merr., Philipp. J. Sci. 26 (1925) 451. — Type:

Philippines, Palawan, Malampaya Bay, Oct. 1922, Merrill BoS 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. inval., descr. Angl. — Based on: Philippines, Luzon, Sorsogon, Irosin

(Mt Bulusan), May 1916, Elmer 16061 (FI, K, L, MO, P, PNH7*)].

eae, eS SS a eee ee

Rhaphidophora in New Guinea, Australia and Western Pacific eu

Figures 17 & 18

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

unbranched, 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; leaves distichous; cataphylls and prophylls membranous,

soon drying degrading to intricately reticulate fibres, these only very slowly

falling; petiole shallowly grooved, upper part + terete, (I—) 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 leaving a

prominent, slightly corky scar; shingling lamina entire, falcate-lanceolate,

5—11 x 3.5—6 cm, base slightly cordate, pre-adult and adult lamina

spreading, 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 sub-parallel to primaries, slightly

raised abaxially, slightly impressed adaxially; secondary venation strongly

reticulate, slightly raised; tertiary venation invisible; inflorescence solitary

to several together, first inflorescence subtended by a 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 anthesis and then caducous leaving 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,

1S6Z Gard. Bull. Singapore 53 (2001)

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 male

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: Widespread in south tropical Asia from Sumatera and southern

Thailand to Borneo and the Philippines eastwards through the tropical

western Pacific.

Habitat: Disturbed lowland, lower and upper hill forest primary, riverine

or secondary forest, on basalt, granite, clay and coralline limestone. 10-800

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. korthalsti are

prominently scaly while those of E. pinnatum are lenticellate-corky. The

pre-adult stage of R. korthalsii is a shingle climber with oblong-elliptic to

ovate slightly falcate upwards pointing leaves overlapping in the manner

of roof tiles and with c. 3 prominent veins per side, running from near the

base of the leaf to the upper margin or tip and crossing over the minor

venation,.

3. Fertile material of R. korthalsii and E. pinnatum is readily separated by

the shape of the style apex (round v. trapezoid) and the shape and

orientation of the stigma (+ punctiform and circumferential v. strongly

linear and longitudinal) and, if fruits are mature, by seed characters. The

fruits of R. korthalsii each contains many small ellipsoid seeds with a brittle,

smooth testa whereas EF. pinnatum has fruits with two large, strongly curved

seeds with a bony and ornamented testa.

Rhaphidophora in New Guinea, Australia and Western Pacific

CEH

SS3

Figure 17. Rhaphidophora korthalsii Schott

L.GUAg.

133

A. pre-adult shoot x '/,; B. pre-adult shingling shoot x '/,. A from Boyce 679; B from Nicolson

LTD.

134 Gard. Bull. Singapore 53 (2001)

4. Confusion is possible between R. korthalsii and Amydrium zippelianum

(Schott) Nicolson although there is a suite of characters that distinguish

them. The leaflet tips of the Amydrium species are acute to acuminate,

those of R. korthalsii are truncate, 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: INDONESIAN PAPUA. Kepala Burung Prov.: surroundings of

Ayawasi, Ave 4330 (BO, L); Manokwari Prov.: Pengunungan Maoke (Nassua Mts), Docters

van Leeuwen 10750 (BO); Wariori River, river valley west of camp between Wariori and

Mangopi rivers, c. 11 km inland, Johns 8197 (BO, K, MAN); Fanindi, 1 km west of

Manokwari, Nicolson 1573 (B, K, L, P, US); Arfak Mountains, Mupi Dessa, trail from

Mupi village to G. Humibou, near S. Mupi, c. 3 km from Kali Umera, between Kali Ureda

and the confluence of Kali Ngwes and S. Mupi, Sands 6846 (K); Mimika Proy.: Freeport

Concession Area, along road from bridge to Kuala Kenchana, along track near left turn by

river, Barker 145 (BO, K, MAN); Kali Kopi, between Kali Kopi levee and the Kopi River,

Johns 9796 (BO, K, MAN): Lorentz River, near Bivak Island Pulle 66 (L); Beaufort (Van

der Sande) River Pulle 353 (K, L); Biak Island, hill northeast of Mokmer airport, Nicolson

1567 (B, K, L, US). PAPUA NEW GUINEA. East Sepik Prov.: vicinity of Wewak, along

trail beyond Boys Town (Reform School operated by S.V.D. Missionaries), Croat 52763

(MO); Lordberg, Ledermann 10319 (B spirit); Malu, Ledermann 10826 (B spirit); Central

Proy.: Port Moresby, northeast of Manumu village, sles & Vinas NGF 33829 (US), Isles &

Vinas NGF 34488 (L); Manus Prov.: Manus, | km SW of Kabuli village on south coast in

western Manus, Sands 2779 (K); Western Prov., Kiunga, Streimann & Womersley LAE

51847 (L, US); Morobe Prov.: vicinity of Lae, along logging road to Busu River, from 3.8

km E of Igam road and Military Base to c. 6 km up the road junction, Croat 52792 (MO),

Croat 52793 (MO); c. 20 km SE of Lae, along logging road to Busu River (left off Igam

road past Military Reserve), Croat 52800 (MO); Along Busu River, 22.6 km by road SE of

Lae (via road past PNG University of Technology and Igam road past Army Base), beginning

3.8 km from asphalt at Army Base, Croat 52840 (MO); Lae B.G. Croat 52849 (MO);

Bumbu logging area, 7 miles north of Lae, Nicolson 1385 (L, US); Oomsis Creek, 20 miles

from Lae on Bulolo road, Nicolson 1483 (B, K, L, P, US); Madang Prov.: no further data,

Ledermann 6641 (B spirit); Lower Ramu-Atitau area, SE of Aiome Patrol Post, along

Apenam track, east side of Tiganants River (loka Creek), Pullen 949 (CANB, L); North

Solomons Prov.: Bougainville, McKillip’s Arawa Plantation, 6 miles west of Kieta, Nicolson

1520 (B, K, L, US); West New Britain Prov.: Kandrian, along road to airport, Nicolson

1542 (B, K, L, US); East New Britain Prov.: Kareeba road, 2 miles west of Kerevat,

Nicolson 1498, (B, K, L, P, US). FEDERATED STATES OF MICRONESIA. Chuuk

(Truk), Toi Island, Suiyota, Takamatsu 40 (K, L); Winipwoot, C.C.Y. Wong 278 (GH, US).

SOLOMON ISLANDS. Rennell Island: Dissing 2776 (K); Kolombangara: 2 miles NNW of

Kuzi village, Hunt RSS 2398 (K, US); San Cristobal: Ridge between Warahito and Pegato

river, Whitmore 6204 (K, L, US). VANUATU. Erromango: Poututu, Bourdy 228 (P);

eee

Rhaphidophora in New Guinea, Australia and Western Pacific 135

Figure 18. 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.

136 Gard. Bull. Singapore 53 (2001)

Ipota, Bourdy 244 (K, P); Portnarvin, Rautop, Cabalion 2370 (P); vicinity of Nouankao

Camp, Green RSNH 1267 (K); Forestry route, km 16.5, Raynal RSNH 16222 (K); Espiritu

Santo: Bank of Achone River, opposite side from Casevaia village, Curry 939 (K); Kuvutant,

Ludvigson 20 (L); Anatom (Aneityum): Anelgauhat Bay, Kajewski 830 (K, P), Morrison

s.n. (K).

17. Rhaphidophora microspadix K. Krause

Rhaphidophora microspadix K. Krause, Bot. Jahrb. Syst. 49 (1912) 92. —

Type: Papua New Guinea (‘Kaiser Wilhelmsland’), Madang Prov., Bismarck

Range, 11 Nov. 1908, Schlechter 18678 (B, holo).

Rhaphidophora nutans Ridl., Trans. Linn. Soc. Lond. 9 (1916) 239, synon.

nov. — Type: Indonesian Papua, Mimika Prov., Puncak Jaya (Mt.

- Carstensz), Camp 6a, 16 Jan. 1913, Boden Kloss s.n. (BM, holo; K iso).

Figure 19

Small, very slender, leptocaul, homeophyllous (?) liane to unknown ultimate

height; seedling and pre-adult plants not observed; adult shoots comprised

of greatly elongated, clinging, physiognomically unbranched, leafy, non-

flowering stems and long, free, sympodial, leafy flowering stems; stems

smooth, somewhat flexuous, stems terete in cross-section, branching little,

growing to considerable lengths and pendent with flowering tips upturned,

without prophyll, cataphyll and petiolar sheath fibre, internodes to 4 x c.

0.2 cm, separated by weak slightly oblique leaf scars, older stems woody;

flagellate foraging stems absent; clasping roots arising singly from each

node; feeding roots not observed; leaves weakly distichous and sparsely

arranged; cataphylls and prophylls membranous, caducous; petiole deeply

grooved adaxially, 2—S x 0.15—0.25 cm, smooth, with a slight apical and

basal geniculum; petiolar sheath slightly prominent, extending to the apical

geniculum, caducous in strips leaving a slender scar; /amina entire, narrowly

lanceolate to lanceolate falcate, 4—19 x 1—3 cm, thinly coriaceous, drying

adaxially dull mid-brown, abaxially paler brown with dense, well-defined

to somewhat obscure tannin cells, base cuneate, apex long-acuminate with

a slender prominent tubule; midrib slightly raised abaxially and adaxially;

primary venation pinnate, very slightly raised on both surfaces; interprimaries

sub-parallel to, but much less distinctive than, primaries, sometimes

degrading into weakly reticulate venation, very slightly raised abaxially;

secondary venation + invisible in dried specimens, parallel-reticulate;

inflorescence solitary, subtended by a fully developed foliage leaf and a

caducous cataphyll; peduncle compressed-terete, 3—7.3 x 0.2—O0.25 cm;

spathe ovoid-ellipsoid, stoutly long-beaked and tipped with a fine tubule,

ee es. ae a a” ae

Rhaphidophora in New Guinea, Australia and Western Pacific [37

Figure 19. Rhaphidophora microspadix K. Krause

A. flowering shoot x '/,; B. leaf lamina x 1'/,; C. venation detail x 6; D. inflorescence, spathe

fallen x 4. All from Boden Kloss s.n.

138 Gard. Bull. Singapore 53 (2001)

2.5—3 x 0.7—0.9 cm, marcescent until early fruiting, then falling leaving a

large, oblique scar; spadix ovoid-globose to ovoid-cylindrical, sessile,

inserted slightly obliquely on peduncle, 1—1.5 x 0.5—0.6 cm; stylar region

rhombohexagonal, 1.4—2 x c. 2 mm, truncate; stigma punctiform, c. 0.4

mm diam., prominent in dried material; anthers well-exserted at male

anthesis; infructescence ovoid-globose, c. 1.2—2.5 x 1—1.5 cm.

Distribution: Indonesian Papua (Mimika Prov.), Papua New Guinea

(Madang and Morobe Provinces).

Habitat: Lower montane forest. 945-1700 m altitude.

Notes: 1. The type specimen of Rhaphidophora microspadix has the abaxial

_ leaf lamina with dense conspicuous tannin cells visible to the naked eye.

These cells are also present on the type of R. nutans, but much less clearly

visible (x10 lens is required).

2. Rhaphidophora microspadix appears to be allied to R. neoguineense,

differing in the much narrower leaf lamina with a long acuminate tip and

in flowering habitually on long, pendent shoots.

Other specimens seen: PAPUA NEW GUINEA. Morobe Prov.: Matap, Clemens 11151,

41197 (GH); Gumi Divide, 25 km west of Bulolo, Kairo 757 (L, LAE).

18. Rhaphidophora mima P.C. Boyce, sp. nov.

Rhaphidophora mima multum cum R. neoguineensi (non in Insulas

Solomonenses inventa) persimili confusa est, quamquam ab ea spatha

fusiformi caduca atque spadice cylindrica (spatha globosa marcescensque

et spadix globoso-ellipsoideus in R. neoguineensi) prompte distinguitur. —

TYPUS: Solomon Islands, Guadalcanal, Wanderer Bay area, 23 Oct. 1968,

Mauriasi et al. BSIP 12268 (SING, holo; BSIP, K, iso).

Figure 20

Slender, leptocaul, homeophyllous liane to 4 m; seedling and pre-adult

plants not observed; adult shoot architecture not fully observed but seemingly

comprised of elongated, clinging, physiognomically unbranched, leafy, non-

flowering stems and slightly lengthened mostly unbranched, free, leafy,

flowering stems; stems smooth, flexuous, climbing stems + terete,

occasionally weakly angled, free stems terete, to similarly sulcate, without

prophyll, cataphyll and petiolar sheath fibre, internodes 0.3—5 x 0.2—0.3

cm on free shoots, flowering shoots with shorter internodes, separated by

cd 6 a nnn

Rhaphidophora in New Guinea, Australia and Western Pacific 139

weak straight leaf scars, older stems woody; flagellate foraging stems not

observed; clasping roots and feeding roots not observed; leaves spiro-

distichous, slightly scattered; cataphylls and prophylls membranous,

caducous; petiole grooved adaxially, 1.5—5 x 0.1—0.18 cm, smooth, with a

slight apical and basal geniculum; petiolar sheath slightly prominent,

extending beyond the apical geniculum by two ligules, caducous leaving a

continuous scar from the petiole base, around the top of the apical

geniculum and back to the base; /amina entire, elliptic to elliptic-lanceolate

or oblanceolate, slightly falcate, 5—16 x 2.2—7 cm, thinly coriaceous, base

cuneate subovate or minutely cordate, apex acute to briefly acuminate,

with a small tubule; midrib raised abaxially, sunken adaxially; primary

venation pinnate, slightly raised on both surfaces in dried material;

interprimaries reticulate to sub-parallel to, but much less distinctive than,

primaries, degrading into weakly reticulate venation, very slightly raised

abaxially; secondary and tertiary venation reticulate; inflorescence solitary,

subtended by a fully developed foliage leaf and a caducous cataphyll:

peduncle slightly compressed-terete, |—3 x 0.2—0.3 cm; spathe cigar-shaped,

apex beaked, 1—4.5 x 0.7—1.2 cm, fleshy, yellow-green to yellow-cream,

caducous leaving a large scar; spadix cylindrical, sessile, inserted level on

peduncle, 1.5—2.5 x 0.5—0.8 cm, obtuse, white; stylar region rounded-

rhombohexagonal, 0.9—1 x 0.85—1.1 mm, truncate; stigma punctiform, c.

0.3 mm diam., slightly prominent in dried material; anthers not exserted at

male anthesis; infructescence not observed.

Distribution: Papua New Guinea (Bougainville, Manus), Solomon Islands

(Fauore, Guadalcanal, Kolombangara, Malaita, San Jorge, Small Malaita).

Habitat: Well-drained primary and secondary forest on flat and hilly terrain,

occasionally in lowland swamp forest. 3—1000 m altitude.

Notes: 1. In herbaria Rhaphidophora mima is much confused with the very

similar R. neoguineensis (absent from the Solomon Islands) although it is

readily distinguished by a caducous, cigar-shaped spathe and cylindrical

spadix (spathe globose and marcescent, spadix globose-ellipsoid in R.

neoguineensis ).

2. The specific epithet is the adjectival form of mimus, from the Greek,

mimos, actor, in allusion to the confusing similarity of this species to

Rhaphidophora neoguineensis.

3. The Manus collection (Foreman & Katik LAE 59291A), while outside

the main geographical range of the species, is undoubtedly R. mima.

140 Gard. Bull. Singapore 53 (2001)

Other specimens seen: PAPUA NEW GUINEA. Manus Prov.: Manus, near Pelikawa,

Foreman & Katik LAE 59291A (L, US); North Solomons Prov.: Bougainville, Sulka wide

bay, Bateson 89, 91, 92, 93 (K); Buin, Kugumaru, Kajewski 1891 (BM, GH, K, SING);

Koniguri, Kajewski 2162 (GH, P); Pavairi, Lavarack & Lavarack NGF 31091 (K, L, US);

McKillup’s Arawa Plantation, 6 miles west of Kieta, Nicolson 1516 (US), Nicolson 1522 (B,

K, L, US); vicinity of Aku village, c. 10 miles west of Buin, Schodde 4070 (BRI, CANB,

GH, K, L, LAE, US); Siwai, Waterhouse 207 (GH, K), Waterhouse 778 (K). SOLOMON

ISLANDS. Malaita: SW Malaita, Wairokai River area, Gafui et al. BSIP 10215 (BSIP, K,

SING); Sw’u area, Mauriasi et al. BSIP 13632 (BSIP, L, SING); Tantalau - Kwalo trail, near

Kwalo, Stone 2366 (BISH, K, US); Small Malaita: Palasu’u, east of Rota School, Gafui

BSIP 17296 (BSIP, K, SING); San Jorge: Talise village, Hunt RSS 2722 (K, US);

Kolombangara: Ridge west of Vila river, Mauriasi et al. BSIP 8429 (BSIP, K, SING);

Fauore: Halluma River, Mauriasi et al. BSIP 13977 (BSIP, K, L, SING).

_19. Rhaphidophora neoguineensis Engl.

Rhaphidophora neoguineensis Engl. in K. Schum. & Hollrung, Fl. Kais.

Wilh. Land (1889) 19 (‘neo-guineensis’); K. Schum. & Lauterbach, FI.

Schutzgeb. Siidsee (1900) 211; Engl. & K. Krause in Engl., Pflanzenr. 37

(IV.23B) (1908) 20—21, Fig. 3 (‘novo-guineensis’); Engl. & K. Krause,

Nova Guinea 8 (1912) 805; Alderw., Bull. Jard. Bot. Buitenzorg III, 1

(1920) 386 (‘novo-guineensis’); Alderw., Bull. Jard. Bot. Buitenzorg III, 4

(1922) 337—338 (‘novo-guineensis’); K. Krause & Alderw., Nova Guinea

14 (1924) 212 (‘novo-guineensis’). — Type: Papua New Guinea (‘Kaiser

Wilhelmsland’), Madang Prov., Adelbert Range, near Hatzfeldthafen, Oct.

1886, Hollrung 372 (B, holo).

Figure 21

Slender, leptocaul, homeophyllous liane to 5 m; seedling not observed; pre-

adult plants forming small terrestrial colonies; adult shoot architecture

comprised of elongated, clinging, physiognomically unbranched, leafy, non-

flowering stems and very short to somewhat lengthened mostly unbranched,

free, sympodial, leafy, flowering stems; stems smooth, flexuous, climbing

stems + terete, occasionally weakly sulcate on two opposing sides, free

stems terete, to similarly sulcate, without prophyll, cataphyll and petiolar

sheath fibre, internodes 1—9 x 0.2—0.6 cm on clinging and free shoots,

flowering shoots with shorter internodes, separated by weak straight leaf

scars, occasionally disarticulating into 2—4 internode lengths and these

later rooting and forming independent plants, older stems woody; flagellate

foraging stems frequent, terete in cross-section with reduced leaves basally,

terminal portion with caducous cataphylls; clasping roots arising sparsely

from the clinging stems, very slightly pubescent; feeding roots solitary from

Rhaphidophora in New Guinea, Australia and Western Pacific

141

Pe,

299

—

Lop!

ms <,

ic SA on

Figure 20. Rhaphidophora mima P.C. Boyce

A. flowering branch, spathe intact x '/;; B. flowering branch, spathe fallen x '/,; C. inflorescence,

spathe fallen, x 2; D. spadix detail, post-anthesis x 4. A from Kajewski 1891; B from Foreman

& Katik LAE 59291A; C-D from Nicolson 1522.

142 Gard. Bull. Singapore 53 (2001)

nodes, clinging to climbing surface; /eaves weakly spiro-distichous,

moderately densely arranged; cataphylls and prophylls membranous,

caducous; petiole grooved adaxially, 3—12 x 0.1—0.2 cm, smooth, with a

slight apical and basal geniculum, although older leaves often with genicula

enlarged and cracking-corky; petiolar sheath slightly prominent, extending

beyond the apical geniculum by two ligules, caducous leaving a continuous

scar from the petiole base, around the top of the apical geniculum and

back to the base; /amina entire, elliptic to elliptic-lanceolate or oblanceolate,

6—25 x 2—9 cm, thinly coriaceous to submembranous, base cuneate to

acute or subovate, apex acute to weakly acuminate, with a minute tubule;

midrib raised abaxially, slightly sunken adaxially in fresh material, slightly

sunken abaxially and adaxially in dry material; primary venation pinnate,

slightly raised on both surfaces in dried material; interprimaries reticulate

. to sub-parallel to, but much less distinctive than, primaries, degrading into

weakly reticulate venation, very slightly raised abaxially; secondary and

tertiary venation reticulate; inflorescence solitary, subtended by a fully

developed foliage leaf and a caducous cataphyll; peduncle slightly

compressed-terete, 1.3—2.5 x 0.15—0.3 cm; spathe globose to ovoid-

ellipsoid, truncate basally at insertion on petiole, apex briefly beaked, 1—2

x 1—2.5 cm, thick fleshy, dark yellow, marcescent, drying brown, later

pushed off by developing infructescence leaving a large scar; spadix globose

to ellipsoid-cylindrical, sessile, inserted level on peduncle, 1—1.5 x 0.7—1

cm, obtuse, yellow-white; stylar region rounded-rhombohexagonal, 0.9—

1.5 x 0.85—1.2 mm, truncate; stigma punctiform to very slightly elliptic, c.

0.45 x 0.3 mm diam., prominent in dried material; anthers well-exserted at

male anthesis; infructescence not observed.

Distribution: New Guinea. Widespread but, by the exclusion of many

collections here recognized as new species, not as common as was once

thought.

Habitat: Primary to disturbed secondary lowland, gallery and hill monsoon

and rain forest on various substrates including coralline limestone. 3—700

m altitude.

Note: Rhaphidophora neoguineensis is characterized by a small (1—2 x 1—

2.5 cm), marcescent, globose spathe and ellipsoid-cylindrical spadix. In

herbaria it is much confused with R. mima but which is readily separated

by the cigar-shaped, caducous spathe and cylindrical spadix. Confusion

with R. schlechteri is also possible, although this has a larger spathe (4—6 x

c. 2.4 cm) and a stipitate spadix.

Rhaphidophora in New Guinea, Australia and Western Pacific 143

a) =

[ri6

Figure 21. Rhaphidophora neoguineensis Engl.

A. adult shoot with flowering branch x */,; B. adult clinging shoot x '/;; C. flagellate shoot x */;

D. leaf lamina x '/,;E. venation detail x 3; F. inflorescence, spathe removed x 4; G. spadix detail,

male anthesis x 6. A-C, F-G from Hay K2 [cult. Kew (Acc. no. 1982-5010) & K spirit no.

58061] x '/;; D-E from Nicolson 1402.

144 Gard. Bull. Singapore 53 (2001)

Other specimens seen: INDONESIAN PAPUA. Bonggo Range, Mamberamo, Pionierbivak,

Lam 763 (L). Jayapura Prov.: Taritatau (Idenburgh) River, Bernhard Bivak, Meyer Drees

279 (L); Biak, hill northeast end of Mokmer airport, Nicolson 1565 (L, US). PAPUA NEW

GUINEA. East Sepik Prov.: Wewak, 2 miles west of But village, Essig & Martin LAE

55124 (L, LAE); West Sepik Prov.: Vanimo, Wutung, on Papua New Guinea/Indonesian

Papua border, Streimann LAE 52789 (LAE, US); Sepik Prov.: Sepik River, Gjellerup 330

(L); Kelel, Schlechter 16329 (P); Keneyia, Schlechter 18298 (P); Western Highlands Prov.:

Mt Hagan, near Ruti E.L.G., Henty & Streimann NGF 38835 (US); Morobe Prov.: Lae,

Markham Bridge, Kairo & Streimann NGF 30715 (GH, K, L, SING, US); Lae B.G., Millar

NGF 9938 (GH, K, L), Nicolson 1402 (B, BM, K, L, P, SING, US); Near Markham River, 9

miles southwest of Lae on Bulolo road, Nicolson 1480 (US); Central Proy.: Port Moresby,

Brown River F.R., 20-25 miles northwest of Port Moresby, Nicolson 1427 (US).

20. Rhaphidophora okapensis P.C. Boyce & Bogner

Rhaphidophora okapensis P.C. Boyce & Bogner, Gard. Bull. Singapore 52

(2000) 94, fig.2. — Type: Papua New Guinea, Eastern Highlands, 5 miles

NE of Okapa, 24 Sept. 1964, Hartley TGH 13098 (CANB, holo; GH, K, L,

iSO).

Figure 22

Moderate, slightly 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 clinging, leafy flowering stems; stems terete

in cross-section, smooth, without prophyll, cataphyll and petiolar sheath

remains, internodes to 5 x 0.75 cm, separated by slightly swollen nodes

with + 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; leaves distichous, those

on adherent shoots weakly shingling to slightly scattered, those on free

shoots pendent to slightly spreading; cataphylls and prophylls membranous,

caducous; 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;

lamina ovate, adaxially mid-green, abaxially paler, very stiffly coriaceous,

2.5—8.5 x 0.9—4 cm, 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;

Rhaphidophora in New Guinea, Australia and Western Pacific 145

Figure 22. Rhaphidophora okapensis P.C. Boyce & J. 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 male anthesis x 10; F. pistil, side view x 10. All from

Hartley 13098.

146 Gard. Bull. Singapore 53 (2001)

inflorescence solitary on short leafy shoots, subtended by a fully developed

or reduced foliage leaf, and a soon-degrading, membranous, 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, (early marcescent?) leaving 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 male anthesis; infructescence

not seen.

Distribution: Papua New Guinea (Eastern Highlands and Central

Provinces).

Habitat: Disturbed mixed forest on slope. 1200-1600 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 turning purple at maturity, the long-

stipitate spadix, and the conical stigmatic region.

Other specimens seen: PAPUA NEW GUINEA. Central Prov.: Boridi, Carr 14863 (BM, L,

SING); Eastern Highlands Prov.: Kainantu, Arau-Andandara road, Streiman NGF 23963

(US).

21. Rhaphidophora pachyphylla K. Krause

Rhaphidophora pachyphylla K. Krause, Bot. Jahrb. Syst. 49 (1912) 92; P.C.

Boyce & Bogner, Gard. Bull. Singapore 52 (2000) 94, fig.2. — Type: Papua

New Guinea, Madang Prov., near Wabbe, 29 Aug. 1907, Schlechter 16463

(B, holo; P, iso).

Figure 23

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 flowering stems; stems rectangular-

terete in cross-section, widest side prominently convex, smooth, mid-green,

without prophyll, cataphyll and petiolar sheath fibre although flowering

shoots occasionally with parchment-like remains, internodes to 3 x | cm,

Rhaphidophora in New Guinea, Australia and Western Pacific 147

Figure 23. Rhaphidophora pachyphylla K. Krause

A. 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-male anthesis x

10. A, E —F from Krause 108; B from Kalkman 3389: C-D from Nicolson 1416; G—-H from

Schlechter 16436.

148 Gard. Bull. Singapore 53 (2001)

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, caducous; 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 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 anthesis and

then caducous leaving a large scar at the base of the spadix; spadix narrowly

cigar-shaped, long stipitate, inserted obliquely on stipe, 4—5.5 x 1.2—1.6

cm, yellow; stipe 6—10 x 2—2.4 mm; stylar region weakly developed, mostly

irregularly rhombohexagonal, 1—1.2 x1—1.4 mm, truncate; stigma slightly

raised, punctiform, c. 0.2—0.3 x 0.3—0.35 mm; anthers slightly exserted at

male anthesis; infructescence not seen.

Distribution: Indonesian Papua, Papua New Guinea. The scattered localities

and few collections suggest a widespread but uncommon species.

Habitat: Lowland monsoon or rain forest at 10—30 m altitude.

Notes: 1. Long confused with Rhaphidophora hayi, R. pachyphylla differs

in leaf shape, and form of the stigma. Additionally, R. pachyphylla lacks

the disarticulating shoots unique to R. hayi, and never has foraging shoots.

2. It is still not fully clear whether R. pachyphylla as here defined is a

single taxon. Hoogland & Schodde 6943 (Western Highlands Prov., Wabag,

near Poio village, west slopes of lower Yaki valley, 6 July 1960, in montane

Nothofagus forest, 2380 m (BM, GH, L), while vegetatively matching typical

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.

—_

Rhaphidophora in New Guinea, Australia and Western Pacific 149

Other specimens seen: INDONESIAN PAPUA. Tamimonding, Kalkman s.n. (L); Digul

Prov.: Merauke, Bis, Agats, Widjaja 6344 (BO, K, L). PAPUA NEW GUINEA. Central

Prov.: Brown River F.R., 20 - 25 miles northwest of Port Moresby, Nicolson 1416 (K, L,

US).

22. Rhaphidophora petrieana A. Hay

Rhaphidophora petrieana A. Hay, Telopea 5 (1993) 295, fig. 1. — Type:

Australia, Queensland, National Park Reserve 904, Palmerston Highway,

30 Nov. 1982, B. Gray 2862 (QRS, holo).

Rhaphidophora sp. aff. australasica (Qld) in Jones & Gray, Climbing PI.

Australia (1988) 316, unnumbered plate p. 322.

Figure 24

Medium to large, moderately robust, semi-leptocaul homeophyllous liane

to 20 m; seedling and pre-adult plants not observed; adult shoot architecture

comprised of greatly elongated, clinging, physiognomically unbranched,

sparsely leafy, non-flowering stems and long, moderately elaborated, free,

sympodial, densely leafy, flowering stems; stems smooth, climbing and free

stems terete in cross-section, green, later mid-brown, without prophyll,

cataphyll and petiolar sheath fibre, internodes to 2.5—5 x 0.5—1 cm on

clinging shoots, usually less stout on free shoots, separated by well defined,

slightly oblique, corky leaf scars, older stems subwoody; flagellate foraging

stems absent; clasping roots sparse, arising from the nodes of clinging stems,

minutely pubescent; feeding roots not observed; leaves spiro-distichous on

clinging shoots, disticho-secund on free shoots; cataphylls and prophylls

membranous, very quickly drying and falling; petiole grooved adaxially,

4.5—8 x 0.15—0.3 cm, smooth, apical and basal genicula weakly defined;

petiolar sheath very prominent, extending to just below apical geniculum,

swiftly drying and eventually falling more-or-less entire leaving a

conspicuous scar; /amina entire, narrowly ovate to lanceolate, somewhat

falcate, slightly oblique, 4.5—22.5 x 1—4.5 cm, thinly coriaceous, base acute,

apex acuminate, with a tiny tubule; midrib raised abaxially, slightly sunken

adaxially; primary venation pinnate, slightly raised abaxially and adaxially;

interprimaries parallel to primaries and barely less prominent, very slightly

raised abaxially and adaxially; secondary venation + obscure in fresh

material, visible as a faint reticulum in dried specimens; inflorescence

solitary, subtended by a partially developed foliage leaf and a membranous

cataphyll; peduncle compressed-terete, 3—10 x 0.15—0.8 cm; spathe canoe-

shaped, stoutly short- to rather long-beaked, 6—8 x 1—3 cm (8 cm wide

flattened out), thickly stiff-fleshy, creamy yellow, caducous at female

150 Gard. Bull. Singapore 53 (2001)

Figure 24. Rhaphidophora petrieana A. Hay

A. flowering shoot x */;; B. inflorescence, longitudinal section x 2/,; C. spadix detail, male anthesis i]

x 6. All from Hyland 9162.

Rhaphidophora in New Guinea, Australia and Western Pacific 151

anthesis; spadix bluntly tapering-cylindric, stipitate, inserted + level on

stipe, 3—6 x 1—2.5 cm; stipe terete, c. 1 x 0.6 cm; stylar region mostly

rounded-hexagonal, c. 2 x 1.8—2 mm, conical; stigma punctiform and

prominently raised, c. 0.25—0.3 mm diam.; anthers exserted at male anthesis;

infructescence not observed.

Distribution: Australia (wet tropical regions of eastern Queensland).

Habitat: Lowland to lower montane rain forest. Sea level to 800 m altitude.

Note: Long confused in herbaria and literature with Rhaphidophora

australasica but readily distinguished by flowering on free lateral shoots

and by the stipitate spadix. Further, in R. petrieana the petiolar sheaths fall

more-or-less intact and do not degrade into semi-persistent fibres as they

do in R. australasica.

Other specimens seen: AUSTRALIA. Queensland: Foot of Mt Demi, Flecker 9005 (QRS);

Timber Reserve 55, Whyanbeel, Gray 202 (QRS); State Forest Reserve 310, Upper

Goldsborough Logging Area, Gray 1055 (QRS); Cooper Creek, Hind 246 (NSW); State

Forest Reserve 755, Barong Logging Area, Hyland 9162 (BRI, L, NSW, QRS).

23. Rhaphidophora pilosa P.C. Boyce, sp. nov.

Rhaphidophora pilosa R. gorokensi R. kokodensique simillima est, ab illa

apices surculorum fibrosos (fibram in R. gorokensi deest) habenti atque ad

hac natura coacta (non simpliciter fibrosa) reliquiarum apicum surculorum

differens. Cum R. waria eam confundere potest, quamquam illa species

Magis major est in surculis adhaerentibus non liberis florens. — TYPUS:

Indonesian Papua, Kepala Burung Prov., Arfak Mts, Minjambau, 20 May

1962, C. Versteegh BW 12647 (L, holo; MAN, iso).

Figure 25

Small, homeophyllous (?) liane to unknown ultimate height; seedling stage

and pre-adult plants not observed; adult shoot architecture comprised of

elongated, clinging, physiognomically unbranched, leafy(?), non-flowering

stems and short, free, sympodial, flowering stems; sfems terete in cross-

section, internodes to at least 5 x 0.5 cm, separated by straight oblique,

weak leaf scars; flagellate foraging stems, unknown; clasping roots solitary,

stout, arising from nodes; feeding roots not observed; leaves weakly spiro-

distichous on free shoots; cataphylls and prophylls chartaceous, very soon

degrading into weak fibres and patches of soft felt-like debris; petiole weakly

canaliculate, 4.5—9 x 0.2—0.25 cm, smooth, apical and basal geniculum

G2 Gard. Bull. Singapore 53 (2001)

Figure 25. Rhaphidophora pilosa P.C. Boyce

A. flowering branch, spathe intact x */;; B. flowering branch, spathe fallen x */,; C. spadix detail,

post-anthesis x 5. All from Versteegh BW12647.

— "=

Rhaphidophora in New Guinea, Australia and Western Pacific 153

well defined petiolar sheath prominent, chartaceous, extending to apical

geniculum, very swiftly degrading to weak fibres and strips of soft felt-like

debris; /amina entire, narrowly lanceolate to narrowly lanceolate-elliptic,

occasionally very slightly falcate, 6—18 x 2—3.5 cm, stiffly coriaceous,

base rounded to acute, apex long-acuminate with a prominent apicule;

midrib raised abaxially, slightly flush to very slightly sunken adaxially;

primary venation densely pinnate, very slightly raised abaxially and

adaxially; interprimaries parallel to primaries, slightly less pronounced;

secondary venation very faintly open-reticulate; inflorescence solitary,

subtended by a foliage leaf and felted debris; peduncle stout, compressed-

terete, 8—11 x 0.3—0.5 cm; spathe canoe-shaped, very long stout-acuminate,

9.5 x c. 1.8 cm, marcescent; spadix cigar-shaped, sessile, 5.5 x 1 cm; stylar

region trhombohexagonal, c. 0.9—1 x 1.2—1.5 mm, truncate; stigma

punctiform, flush, c. 0.4 mm diam.; anthers not exserted at male anthesis;

infructescence cylindrical, c. 8 x 1.8 cm.

Distribution: Indonesian Papua (Kepala Burung Prov.). Known only from

the type.

Habitat: Secondary submontane forest. 1250 m altitude.

Notes: 1. Rhaphidophora pilosa is one of several montane species with

stiffly coriaceous leaves (Okapensis Group). It is most similar to R.

gorokensis and R. kokodensis, differing from the former in possessing

fibrous shoot tips (R. gorokensis lacks fibre) and from the latter in the

felted (not simply fibrous) nature of the shoot tip debris. Confusion with

R. waria is possible although this is a much larger species flowering on

clinging, not free, shoots.

2. The specific epithet is from the Greek ‘pilosa’, felt-like, in allusion to

the unique manner in which the cataphylls, prophylls and petiolar sheath

degrade into felted fibres.

24. Rhaphidophora schlechteri K. Krause

Rhaphidophora schlechteri K. Krause, Bot. Jahrb. Syst. 49 (1912) 94. —

Type: Papua New Guinea, Eastern Highlands Prov., Tanqueti camp, 27

Nov. 1908, Schlechter 18888 (B, holo).

Rhaphidophora conferta K. Krause, Bot. Jahrb. Syst. 49 (1912) 95, synon.

noy. — Type: Papua New Guinea, Keneyia Camp, 2 Oct. 1908, Schlechter

18308 (B, holo; P, iso).

154 Gard. Bull. Singapore 53 (2001)

Rhaphidophora peekelii Engl. & K. Krause, Bot. Jahrb. Syst. 54 (1916) 78;

Peekel ex Henty, Fl. Bismarck Archipelago (1984) 66—67 fig. 110, synon.

nov. — Type: Papua New Guinea, New Ireland, Namatanai, Peekel 296 (B,

holo).

Rhaphidophora buergersii Engl. & K. Krause, Bot. Jahrb. Syst. 54 (1916)

78 (‘biirgersii’), synon. noy. — Type: Papua New Guinea, Sepik Prov.,

Lordberg, 9 Dec. 1912, Ledermann 10241 (B, holo).

Rhaphidophora drepanophylla Alderw., Bull. Jard. Bot. Buitenzorg III, 4

(1922) 340; K. Krause & Alderw., Nova Guinea 14 (1924) 213, synon. nov.

— Type: Indonesian Papua, Bonggo Range, Mamberamo, near Prauwen

Bivak, 1 Sept. 1920, Lam 1014 (BO, holo; L, iso).

Rhaphidophora obliquata Alderw., Bull. Jard. Bot. Buitenzorg III, 4 (1922)

340; K. Krause & Alderw., Nova Guinea 14 (1924) 212, synon. nov. —

Type: Indonesian Papua, Bonggo Range, Mamberamo, near Pionier Bivak,

30 June 1920, Lam 483 (BO, holo).

Figures 26 & 27

Slender to moderately robust, leptocaul, homeophyllous (?) liane to 20 m;

seedling and pre-adult plants not observed; adult shoot architecture comprised

of elongated, clinging, physiognomically unbranched, leafy, non-flowering

stems and very short to somewhat lengthened, unbranched, free, sympodial,

leafy, flowering stems; stems smooth, flexuous, terete, without prophyll,

cataphyll and petiolar sheath fibre, internodes 1—9 x 0.5—1.5 cm, separated

by thin, slightly oblique, leaf scars, older stems woody; flagellate foraging

stems absent; clasping roots arising singly from the node; feeding roots not

observed; Jeaves spiro-distichous, scattered; cataphylls and prophylls

membranous, caducous; petiole grooved adaxially, 7—14 x 0.2—0.3 cm,

smooth, with a very slight apical and prominent basal geniculum; petiolar

sheath very prominent on newest leaves, very broad basally, narrowing c. +

way along petiole and extending beyond the apical geniculum by two tiny

ligules, caducous leaving a continuous scar from the petiole base, around

the top of the apical geniculum and back to the base; /amina entire,

lanceolate to oblong-lanceolate, slightly to prominently oblique, 10—34 x

4—9.5 cm, thinly coriaceous, base obtuse-cuneate to rounded and minutely

cordate, apex falcate, acute to long-acuminate, with a tiny tubule; midrib

slightly raised abaxially, flush to very slightly sunken adaxially; primary

venation pinnate, slightly raised on both surfaces; interprimaries sub-parallel

and barely less distinctive than primaries; secondary and tertiary venation

reticulate, conspicuous in dry material; inflorescence solitary, subtended by

Rhaphidophora in New

Guinea, Australia and Western Pacific

155

Figure 26. Rhaphidophora schlechteri K. Krause

. flowerin

A. flowering shoot x

E. spadix detail, pre-anthesi

fallen x 1;E.

etal. BSIP 914].

'/;; B. leaf lamina x '/,; C. venation detail x 4; D. inflorescence, spathe

sx8.A & C from Schlechter 18888; B, D-E from Boraule

156 Gard. Bull. Singapore 53 (2001)

a incompletely developed foliage leaf and one or more prominent caducous

cataphylls; peduncle slightly compressed-terete, 5—9 x c 0.5—1 cm; spathe

cigar-shaped, apex stoutly beaked, 4—11 x c. 3.5 cm (flattened out), thick,

marcescent, falling as fruits ripen, green at male anthesis; spadix cylindrical,

stipitate, inserted slightly obliquely on stipe, 6—8.5 x 1.5—2 cm, creamy

white; stipe terete, 0.5—1 cm long; stylar region depressed-

rhombohexagonal, c. 1.2 x 1.3 mm, truncate; stigma punctiform, c. 0.3 mm

diam., raised at male anthesis but barely prominent in dried material;

anthers strongly exserted at male anthesis; infructescence stoutly cylindrical,

c. 8 x 3 cm, with dry spathe persisting.

Distribution: Widespread from Papua New Guinea (including Bismarck

Archipelago) to the Solomon Islands.

"Habitat: Primary to disturbed secondary broadleaf and coniferous

(Araucaria) forest on humus, alluvium and lava. Sea level to 1500 m altitude

Notes: 1. As defined here Rhaphidophora schlechteri is a variable and

widespread species. The broad species concept adopted is based on

examining all the numerous duplicates of the Nicolson collections cited

here that exhibit wide variation, which supports the merging of elements

recognized as separate species by earlier workers.

2. In most of its medium to large manifestations R. schlechteri is similar in

overall appearance to R. mima, but is distinguishable by the marcescent

spathe and stipitate spadix. Small forms of R. schlechteri maybe confused

with R. neoguineensis, although the globose spathe and sessile ellipsoid

spadix readily distinguish the latter.

Other specimens seen: PAPUA NEW GUINEA. West Sepik Prov.: Telefomin, Sanduan,

track on leaf bank of Mai (Yuwa) river, to 0.5 km south of Fiak airstrip, Frodin et al. 2553

(K); Central Prov.: Sogeri Plateau, Rouna Waterfall, 20 miles east of Port Moresby, Nicolson

1432 (B, BM, K, L, P, SING, US); Morobe Prov.: Wau, road half way to Yamap, Kairo

NGF 44078 (L, NGF, US); 10 miles north of Bulolo, Nicolson 1477 (B, K, L, P, US);

Madang Provy., southern slopes of Finisterre Range, near Budemu, Pullen 5996 (BM, L,

LAE); Saidor, Matafuma village, vicinity of Pukie Primary School, Vandenberg & Katik

NGF 42372 (GH, K, L, US); North Solomons Prov.: Bougainville, vicinity of Barilo village,

c. 6 miles north of Buin Station, Schodde 3935 (GH, L); New Ireland: Mussau, Kgie &

Olsen 1377 (L); Namatanai, coastal region, near Tamul River, c. 0.5Skm NW of Taron, east

coast, Sands et al. 2044 (GH, K, L, US); SOLOMON ISLANDS. Guadalcanal: West

Guadalcanal, Wanderer Bay, Vuragoba area, Boraule et al. BSIP 9141 (BSIP, K, SING);

North central Guadalcanal, Tina River, Nakisi & Babala BSIP 8222 (BSIP, K, SING); San

Cristobal: Puepue River, Brass, 2794 (BO, GH, L); South Ridge west of Tetere village,

Gafui et al. BSIP 1 2575 (BSIP, K, SING); Fauore (‘Fauo’) Island: Guppy 196 (K); Ulawa:

Moh, Teona BSIP 6303 (BSIP, K, L, SING).

‘

Rhaphidophora in New Guinea, Australia and Western Pacific

Figure 27. Rhaphidophora schlechteri K. Krause

157

A. flowering shoot x '/;; B. leaf lamina x '/>; C. venation detail x 2: D. inflorescence, spathe

fallen x 1; E. spadix detail, post-male anthesis x 6. All from Ledermann 10241.

158 Gard. Bull. Singapore 53 (2001)

25. Rhaphidophora spathacea Schott

Rhaphidophora spathacea Schott, Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 129.

— Type: ‘New Guinea’, Zippelius s.n. (L, holo).

Rhaphidophora apiculata K. Krause, Bot. Jahrb. Syst. 49 (1912) 93, synon.

noy. — Type: Papua New Guinea (‘Kaiser Wilhelmsland’), Madang Prov.,

Finisterre Range, 2 Sept. 1908, Schlechter 18152 (B, holo; P, iso).

Rhaphidophora palauensis Koidz., Bot. Mag. Tokyo 30 (1916) 400, synon.

nov. — Type: Palau, Angaur Island, Kayima 154 (TI, holo).

Rhaphidophora forbesii Rendle, J. Bot. 61, Suppl. (1923) 58, synon. nov.

- — Type: Papua New Guinea, Sogeri Prov., 1885-1886, Forbes 323 (BM,

holo).

Rhaphidophora engleri Kanehira, Fl. Micrones. (1933) 409. —

Rhaphidophora palauensis Engl. & K. Krause, Bot. Jahrb. Syst. 56 (1921)

433, nom. illeg., non Koidz. (1916), synon. nov. — Type: Palau, Babelthuap

(‘Babelthaob’), Dorfe Ngarsul, 21 Feb. 1914, Ledermann 14336 (B, holo).

Rhaphidophora kanehirae Hatusima, J. Japanese Bot. 15 (1939) 19, fig. 1,

g—j, synon. nov. — Type: Federated States of Micronesia, Yap, Kanehira

1188 (TI, holo).

Figures 28 & 29

Moderately robust, medium-sized pachycaul, homeophyllous liane to 15

m; seedling stage not observed; pre-adult plants forming scattered terrestrial

colonies; adult shoot architecture comprised of clinging, physiognomically

unbranched, densely leafy flowering stems; stems smooth, mid-green, with

cataphylls and prophylls persistent and drying dark yellow, then degrading

into fibres and parchment-like remains, internodes 1—6 x 0.4—1.5 cm,

separated by prominent slightly oblique leaf scars; flagellate foraging stem

absent; clasping roots densely arising from the nodes and internodes, smooth

and drying with parchment-like epidermis; feeding roots not observed; leaves

spiro-distichous to distichous; cataphylls and prophylls chartaceous, later

degrading into strips of tissue and fibres, especially at tips of flowering

shoots; petiole deeply canaliculate, 8—S53 x 0.4—0.8 cm, smooth with faint

to rather prominent dark dense speckling, apical geniculum prominent,

basal geniculum slightly less so; petiolar sheath extending to apical

geniculum, broad, chartaceous, short-persistent, degrading to sparse fibres

and papery strips, then falling; /Jamina entire, ovate-elliptic to oblong-

Rhaphidophora in New Guinea, Australia and Western Pacific 159

Za \

SSSS5Q

ZS = SS

Figure 28. Rhaphidophora spathacea Schott

A. flowering shoot x '/,; B. leaf lamina x '/;; C. venation detail x 3; D. inflorescence, spathe

removed x */,; E. spadix detail, post-male anthesis x 8; F. stylar region and stigma, side view x

6. All from Forbes 323.

160 Gard. Bull. Singapore 53 (2001)

lanceolate or oblong-elliptic, slightly oblique, 11—35 x 2.5—18 cm,

chartaceous to coriaceous, base rounded to acute, apex acute to acuminate;

midrib prominently raised abaxially, + flush adaxially; primary venation

pinnate, raised abaxially, less so adaxially; interprimaries sub-parallel to

primaries, slightly to much less prominent than the primaries, slightly raised

abaxially, barely visible adaxially; secondary venation reticulate, raised,

especially conspicuous in dry material; tertiary venation minutely reticulate,

hardly visible; inflorescences two, three or more together, each subtended

by a prominent chartaceous prophyll and one or more chartaceous

cataphylls, the entire synflorescence emerging from a mass of dried,

chartaceous cataphyll remains; peduncle slender to stout, terete, partially

to completely obscured by cataphylls, 4—12 x 0.2—0.8 cm; spathe slender

canoe-shaped, hardly to stoutly beaked, 5—13.5 x 2—2.5 cm, stiff-fleshy,

_ lower 0.7—1 cm with connate margins (fide Nicolson 1580, but not observed

elsewhere), yellow to yellow green, marcescent to early fruiting, eventually

falling leaving a prominent scar; spadix stoutly cylindrical, sessile, inserted

almost level on peduncle, 3—16.5 x 1—1.5 cm, white at male anthesis;

stylar region conical, mostly hexagonal in top view, 0.9—1.2 x 1—1.1 mm;

stigma punctiform, very prominently raised, those at the tip of the spadix

even more raised, 0.1—0.2 x c. 0.3 mm, glossy and almost black in dried

material; anthers exserted at male anthesis; infructescence stoutly oblong-

cylindrical, 6—9 x 1.4—2 cm.

Distribution: Palau, Indonesian Papua, Papua New Guinea (including

Woodlark (Muyua) Island), Federated States of Micronesia (Yap).

Habitat: Lowland to lower montane primary to disturbed secondary forest

on a variety of substrates including granite, coralline limestone, volcanically-

derived soils. Sea-level to 1500 m altitude.

Notes: 1. Most similar to Rhaphidophora versteegii but readily separated

by the non-shingling juvenile phase and on drying the strongly discolorous

leaf laminas of which the abaxial surface is bright orange-brown with

copious, minute tannin cells. Further, R. spathacea never has perforated

leaf laminas.

2. In publishing R. spathacea, Schott stated that it was from Java, although

both type sheets in L are labelled ‘Nov. Guinea.’ The types of R. spathacea

match very well collections from New Guinea made under the various

synonyms proposed above.

Rhaphidophora in New Guinea, Australia and Western Pacific 161

Figure 29. Rhaphidophora spathacea Schott

A. flowering shoot x '/;; B. leaf lamina x '/;; C. venation detail x 2: D. pre-adult clinging shoot x

'/4; E. inflorescence, spathe sectioned x 1: F. spadix detail, post male anthesis x 8; G. stylar

regions and stigmas at spadix tip x 8. All from Brass 23847.

162 Gard. Bull. Singapore 53 (2001)

3. The type of R. engleri has rather narrow leaves, although other collections

from Palau match collections of R. spathacea from New Guinea.

4. The holotype of R. apiculata K. Krause is notable for the ovate-elliptic

leaf laminas with primary and interprimary veins almost indistinguishable.

However, the isotype (P) is of a much longer, narrower leaf, which is much

closer in appearance to the type of, for example, R. versteegii and R.

spathacea.

5. The type of R. forbesii is very incomplete. Nonetheless, the leaf lamina

shape and spadix match the type of R. apiculata almost exactly.

Other specimens seen: PALAU: Babelthuap: trail along Japanese pipeline between Ngardmau

and waterfront, Bowden-Kerby 5983 (US); Lake Ngardok Fosberg 32572 (US); Aulupse’el,

~ Direbachal Beach, Evans 581 (US); Yap: Talgo n Bibau village agroforest, Falanrum &

Faimaw 6795 (US); Angaur: East coast, Fosberg 31986 (US); Angaur, Koidzumi s.n. (T1).

INDONESIAN PAPUA. Kepala Burung Prov.: Surroundings of Ayawasi, Ave 4736 (BO,

L); Triton Bay, Le Guillou ‘Triton 48° (P); 2 km north of Manokwari, west end of Tafelberg

F.R., Nicolson 1580 (B, K, L, P, US); Mimika Proy.: Freeport Concession Area, path east at

Mile 50 on road to Tembagapura, Coode 8037 (BO, K, MAN); Golf Course surrounds,

Johns 9967 (BO, K, MAN); Freeport Concession Area, Joins 10393 (BO, K, MAN); Golf

Course surrounds, Johns 10431 (BO, K, MAN). PAPUA NEW GUINEA. East Sepik

Prov.: vicinity of Malu and April river, Ledermann 7384 (B spirit); Western Proy., Palmer

River, 2 miles below junction of Black River, Brass 7100 (GH), Brass 7380 (GH); Central

Prov.: Port Moresby, above Boridi village, Foreman & Vinas LAE 60099 (BRI, L, LAE,

US); Morobe Prov.: Along road to Sankwep SE of Lae, c. 10 km beyond Sankwep, Croat

52808 (MO); North slopes of Mt Missim (Misson), Croat 52952 (K, MO); Bumbu Logging

Area, 7 miles north of Lae, Nicolson 1386 (B, P, SING, US), Nicolson 1409 (SING, US);

Lae, Sankwep, 10 miles east of Lae, Streimann NGF 47054 (US); Milne Bay Prov.: Bibiguni

Camp, Gwariu River, Brass 23847 (GH, L, US); Alotau, Kulumadau Island, Kairo 237

(GH, L); Woodlark (Muyua) Island: Kulumadau, Brass 28618 (L).

26. Rhaphidophora spuria (Schott) Nicolson

Rhaphidophora spuria (Schott) Nicolson, Allertonia 1 (1978) 348. —

Cuscuaria spuria Seem. ex Schott, Bonplandia 9: 260, nom. nud.; Schott,

Bonplandia 9 (1861) 367; Seeman, Viti (1862) 444 & FI. Vit. (1868) 287;

Engl., DC, Monogr. Phan. 2 (1879) 251, in syn. pro Cuscuaria marantifolia

Schott; Engl. & K. Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 68, in

syn. pro Scindapsus cuscuaria (Aubl.) Presl. — Type: Fiji, Viti Levu, 1860,

Seeman 655 (K, holo).

Rhaphidophora storckiana Schott, Bonplandia 10 (1862) 346; Seeman, FI.

Vit. (1868) 287; Engl. & K. Krause in Engl., Pflanzenr. 37 (I1V.23B) (1908)

Rhaphidophora in New Guinea, Australia and Western Pacific 163

43; Parham, Pl. Fiji Isl. (1964) 267; ed.2 (1972) 363. — Rhaphidophora

peepla var. storckiana (Schott) Engl., DC, Monogr. Phan. 2 (1879) 243;

Drake, Ill. Fl. Ins. Mar. Pac. (1892) 326. — Type: Fiji, Ovalau, 1862, Storck

911 (K, holo; BM, iso).

Rhaphidophora reineckei Engl., Bot. Jahrb. Syst. 25 (1898) 9; Engl. & K.

Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 41, synon. noy. — Type:

Western Samoa, Savai’i, 1894, Reinecke 594 (B, holo).

Rhaphidophora graeffei Engl., Bot. Jahrb. Syst. 25 (1898) 9; Engl. & K.

Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 40, synon. nov. — Type:

Western Samoa, Upolu, Graeff 73 (B, holo).

Figure 30

Moderate to large, robust, semi-pachycaul homeophyllous liane to 15 m;

seedling stage not observed; pre-adult plants forming small terrestrial

colonies; adult shoot architecture comprised of elongated, clinging,

physiognomically unbranched, leafy, non-flowering stems and short, usually

unbranched, free, sympodial, densely leafy, flowering stems; stems smooth,

terete in cross-section, with sparse prophyll, cataphyll and petiolar sheath

fibre, this soon falling, internodes 0.5—8 x 0.5—2 cm on clinging shoots,

usually shorter and stouter on free shoots, separated by large, straight,

corky leaf scars, older stems woody; flagellate foraging stems absent; clasping

roots densely arising from the nodes and internodes of clinging stems,

notably pubescent; feeding roots not observed; leaves weakly spiro-distichous

on clinging and free shoots; cataphylls and prophylls membranous, quickly

drying and degrading into sparse fibres, these soon falling; petiole deeply

canaliculate, 14—66 x 0.3—0.1.5 cm, apical and basal geniculum moderately

prominent; petiolar sheath prominent, extending to the apical geniculum,

swiftly drying and degrading into sparse, soon-falling fibres; /amina entire,

ovate-oblong to oblong-lanceolate or oblong-elliptic, sometimes slightly

oblique, occasionally falcate, 5.7—76 x 2.5—32 cm, thinly to quite coriaceous

(larger laminas tending to be thinner textured), often drying strongly

discolorous, adaxially mid-brown, abaxially pale brown, base unequal,

cuneate to rounded, subtruncate or weakly cordate, where present basal

‘lobes’ more developed on one side, apex acute to obtuse, acuminate with

a prominent, short tubule; midrib prominently raised abaxially, + sunken

adaxially; primary venation pinnate, raised abaxially and adaxially;

interprimaries sub-parallel to primaries, hardly less prominent, slightly raised

abaxially and adaxially; secondary venation reticulate, slightly raised;

inflorescence solitary to several together, strongly sweet-fragrant, if solitary

164 Gard. Bull. Singapore 53 (2001)

then subtended by a partially to fully developed foliage leaf, if more then

one than subsequent inflorescences each subtended by a soon-degrading

membranous prophyll and cataphyll; peduncle compressed-terete, often

with a deep longitudinal sulcus on the shoot side, 5—19 x 0.25—1 cm;

spathe broadly canoe-shaped, stoutly beaked, 9.5—16 x 2.5—5.5 cm (up to

c. 7 cm wide when flattened out), stiff-fleshy, cream at male anthesis,

caducous leaving a large, straight scar; spadix cylindrical, inserted + level

to somewhat obliquely on peduncle, 6.2—15.5 x 1.2—1.5 cm, creamy white

at male anthesis; stylar region mostly hexagonal, 1.6—2.4 x c. 2 mm diam.,

truncate, area around stigma sunken; stigma punctiform to slightly ellipsoid,

c. 0.5—0.7 mm x 0.5 mm; anthers exserted at male anthesis; infructescence

stoutly oblong to tapering-cylindrical, 10—12 x 1.1—2.5 cm.

Distribution: Papua New Guinea (Western Prov.), Solomon Islands

(Guadalcanal, New Georgia, San Cristobal and the Santa Cruz Group),

Fiji, Western and American Samoa.

Habitat: Well-drained primary and secondary lowland to upper hill forest,

on rocky, red soils. 30— 1000 m altitude.

Notes: 1. Rhaphidophora spuria is the only entire-leaved Rhaphidophora

indigenous to Fiji and Samoa. Examination of the types of R. spuria, R.

graeffei and R. reineckei reveals no characters separating these species

hitherto recognized as endemic for Fiji (R. spuria) and the Samoan islands

(R. graeffei and R. reineckei). The earliest name, R. spuria, is adopted

here.

2. Rhaphidophora spuria is most similar to R. intonsa from Papua New

Guinea (Morobe and Central Prov.) but may be readily distinguished by

the truncate to weakly cordate lamina base and in having very much less

fibre at the shoot tips.

3. Despite the geographical disjunction, the single collection from mainland

Papua New Guinea is of this species, matching the Pacific plants in

vegetative and floral characters.

Other specimens seen: PAPUA NEW GUINEA. Western Prov.: Fly River, Madiri Plantation,

Womersley & Simmonds 5047 (GH). SOLOMON ISLANDS. New Georgia: Hovoro,

Cowmeadow’s collectors BSIP 3791 (BSIP, K, SING); San Cristobal: Ridge west of Warahito,

Hunt RSS 2297 (BSIP, K, L); Santa Cruz Group: Vanikolo (Vanikoro) Island, Kajewski 584

(GH, K); Guadalcanal: adjacent to Tina River, 12 miles inland from coast, Womersley &

Whitmore BSIP 1110 (BSIP, K, L, SING). FIJI. No further data, Capt. Wilkes Exped. s.n.

(US); Viti Levu: Nausori Highlands, 16 km from Bukika Sawmill, Melville & Melville

Rhaphidophora in New Guinea, Australia and Western Pacific 165

Figure 30. Rhaphidophora spuria (Schott) Nicolson

A. flowering shoot x '/,; B. leaf x 3/\9; C. spadix detail, male anthesis x 10. A from Kajewski 584;

B from Teraoka & Kennedy 64; C from Smith 8653.

166 Gard. Bull. Singapore 53 (2001)

71.998 (K); Naitasiri, Colo-I-Suva, Parham 11246 (K); MBA, vicinity of Nalotawa, eastern

base of Mt Evans Range, Smith 4443 (US); Namosi, northern base of Korombasambasanga

Range, in drainage of Wainavindrau Creek, Smith 8653 (K, L, P, US); Namosi, hills east of

Wainikoroiluva River, near Namuamua, Smith 8947 (K, L, P, US). Kadavu (Kandavu),

Namalata Isthmus region, Smith 29 (K, P, US); WESTERN SAMOA. Upolu: Lake Lanuto’o,

Christophersen 402 (BISH, K); Above Vaipouli, Christophersen & Hume 1902 (BISH, K, P,

US); Tapatapa, Cox 34 (K); Near Lotofaga, Lafulemu Ranch (property of Fay Ala’ilima),

west of Fagatola River, c. 0.5 km NW of the bridge, Teraoka & Kennedy 64 (US); Near Mt

Leou’e, Whistler 743 (B, US); Savai’1: Salailua,Christophersen 2956 (BISH, K, P); La Vai,

above Salailua Christophersen 3004 (BISH, US); Above Asau, Block 28, Whistler 1036 (B,

US). AMERICAN SAMOA. Pago Pago, Meebold 8209 (K).

27. Rhaphidophora stenophylla K. Krause

Rhaphidophora stenophylla K. Krause, Bot. Jahrb. Syst. 49 (1912) 94. —

. Type: Papua New Guinea (‘Kaiser Wilhelmsland’), East Sepik Prov., Djamu,

22 April 1908, Schlechter 17581 (B, holo#; P, iso).

Figure 31

Large, rather robust, semi-leptocaul, homeophyllous liane to unknown

ultimate height; seedling stage very slender terrestrial to climbing, leaves

arranged in two ranks; pre-adult plants clinging, slender, with very narrow

leaves; adult shoot comprised of elongated, clinging, physiognomically

unbranched, non-flowering stems and free lateral, leafy flowering stems;

stems terete to strongly compressed, but not rectangular in cross-section,

without cataphyll, prophyll and petiolar sheath fibre, internodes 1—3 x

0.2—2 cm, separated by prominent almost straight, slightly corky leaf scars;

flagellate foraging stem absent; clasping roots arising from nodes and

internodes; feeding roots not observed; leaves spiro-distichous in mature

plants; cataphylls and prophylls caducous; petiole deeply canaliculate, 2—

16 x 0.2—1 cm, apical geniculum small and very obscure, basal geniculum

weakly defined; petiolar sheath very prominent, broadly winged and

extending to apical geniculum, persistent almost through to leaf fall; /amina

entire, linear-lanceolate to narrowly lanceolate or narrowly oblong-

lanceolate, slightly falcate, 9—38 x 1.5—4 cm, subcoriaceous, base subacute

to slightly decurrent, apex acute to briefly acuminate with a prominent

tubule; midrib raised abaxially, sunken adaxially; primary venation pinnate,

slightly raised abaxially, almost flush adaxially; interprimaries reticulate-

parallel to primaries, almost indistinguishable from them, slightly raised

abaxially, almost flush adaxially; secondary venation reticulate, hardly

visible; inflorescence solitary, subtended by a + fully developed foliage

leaf; peduncle slightly laterally compressed to terete, 6—10 x c. 1 cm; spathe

broadly canoe-shaped, stoutly long-beaked, 11—13 x 1.5—2.5 cm, stiffly

Rhaphidophora in New Guinea, Australia and Western Pacific 167

Figure 31. Rhaphidophora stenophylla K. Krause

A. adult shoot with flowering branch x '/;; B. leaf lamina x '/>; C. venation detail x 3; D. leaf

lamina x '/;; E. pre-adult clinging shoot x '/,; F. pre-adult creeping shoot x */;; G. inflorescence,

spathe fallen x 1; H. spadix detail, male anthesis x 6. A-C, E & F from Nicolson 1430; D, G-H

from Streimann NGF 44247.

168 Gard. Bull. Singapore 53 (2001)

fleshy, yellow, caducous leaving a large scar at the base of the spadix;

spadix cylindrical, slightly curved, long stipitate, 6@—9 x 1.5—2.2 cm, inserted

level to almost truncate on stipe, white; stipe slender to stoutly terete,

2.5—3 x 3—0.4 cm; stylar region rhombohexagonal, 1—2 x 1—1.5 mm,

conical; stigma punctiform, prominently raised, c. 0.3 mm diam.; anthers

strongly exserted at male anthesis; infructescence not observed.

Distribution: Papua New Guinea (including the Louisiade Archipelago).

Habitat: Monsoon lower montane forest to submontane primary forest on

moderate to steep slopes, sometimes on limestone. 20-940 m altitude.

Notes: 1. Unmistakable by the long, narrow leaves, persistent winged

_ petiolar sheath, long stipitate spadix and prominently raised stigma. There

exist plants with broader laminas than typical (Brass 27691 & 28311 are of

this broad-leaf type) that may be confused with Rhaphidophora discolor

(also flowering on free shoots and with long-stipitate spadix), although the

latter is readily distinguished by the broader leaf laminas, petiolar sheath

degrading to strips and fibres, larger (13.5—19 x 2.2—4 cm) spadix and flat

stigmas.

2. Scindapsus schlechteri K. Krause is superficially similar in its narrow leaf

lamina and persistent-winged petioles but differs, aside from a single ovule

per ovary (and thus one-seeded fruits), by the sessile spadix and thicker

leaf lamina with almost no visible venation.

Other specimens seen: PAPUA NEW GUINEA. West Sepik Prov.: Telefomin, Sandaun,

Hak Valley, Gentry transect on slope above Nenem, SE of Mianmin Airstrip, Frodin &

Morren 3155 (K); East Sepik Prov.: vicinity of Malu & April river, Ledermann 8614 (B

spirit); Central Prov.: Port Moresby, Koitaki (c. 30 km WNW of Port Moresby), Carr 12218

(BM, SING); Sogeri Plateau, 5-7 miles beyond Kokoda Trail Monument, 30 miles east of

Port Moresby, Nicolson 1430 (K, L, P, SING, US); Milne Bay Prov.: Louisiade Archipelago,

Rossel and Misima Island, Narian, Brass 27691 (GH, L); Abaleti, Brass 28311 (L); Morobe

Prov.: Wau, Bulolo, Middle L.A., Streimann NGF 44247 (GH, K, LAE, SING, US).

28. Rhaphidophora stolleana Engl. & K. Krause

Rhaphidophora stolleana Engl. & K. Krause, Bot. Jahrb. Syst. 54 (1916)

79. — Type: Papua New Guinea, East Sepik Prov., April River, 24 May

1912, Ledermann 7382 (B, holo).

Figure 32

Moderate, somewhat robust, semi-leptocaul, homeophyllous (?) lane

ee ee

Rhaphidophora in New Guinea, Australia and Western Pacific 169

Figure 32. Rhaphidophora stolleana Engl. & K. Krause

A. adult shoot with flowering branch x '/,; B. leaf lamina x '/;; C. venation detail x 3; D.

inflorescence, spathe sectioned x 1'/,; E. spadix detail, post-male anthesis x 8. All from

Ledermann 7382.

170 Gard. Bull. Singapore 53 (2001)

unknown ultimate height; seedling and pre-adult plants not observed; adult

shoot architecture not completely known, but observed to comprise of greatly

elongated, strongly pendent, physiognomically unbranched, leafy, non-

flowering stems giving rise to abbreviated short, free, sympodial, flowering

stems; stems smooth, climbing stems not observed, free stems weakly four

angled to + terete in cross-section, occasionally with two close longitudinal

keels along one side, dull pale yellow brown, without prophyll, cataphyll

and petiolar sheath fibres, internodes to 13.5 x 1 cm, flowering shoots

abbreviated, arising from just above the axils of leaves and twisting to

present inflorescences upwards; flagellate foraging stems not observed;

clasping roots arising singly from the nodes and internodes, slightly

pubescent; feeding roots not observed; leaves very weakly spiralled,

scattered; cataphylls and prophylls membranous, caducous; petiole shallowly

_ canaliculate, 13—14 x 0.35—0.7 cm, smooth, with a well defined apical and

large, slightly prominent basal geniculum; petiolar sheath very prominent,

extending to apical geniculum, swiftly caducous leaving a slight scar; /amina

entire, elliptic to oblong-elliptic, 20—29 x 8—13 cm, membranous, drying

discolorous, adaxially mid-brown, abaxially pale orange-brown, base

subovate, very briefly decurrent, apex acute to briefly acuminate, slightly

falcate with a minute apiculate tubule; midrib slightly raised abaxially,

sunken adaxially; primary venation pinnate, slightly raised abaxially;

interprimaries parallel to primaries, occasionally slightly reticulated, slightly

raised abaxially; secondary and tertiary venation reticulate, prominent

abaxially in dried specimens; inflorescence solitary, subtended by a foliage

leaf; peduncle terete, c. 6 x 0.3 cm; spathe elongate-cylindric, stoutly short-

beaked, c. 10.2 x 1.7 cm, thinly stiff-coriaceous, duration unknown; spadix

slender cylindrical, sessile, inserted very obliquely on to the peduncle, c. 8

x 1.1 cm; stylar region rhombohexagonal, 1—1.2 x c. 1 mm, truncate; stigma

slightly longitudinally elongate to punctiform, c. 0.4 x 0.25 mm; anthers

exsertion not observed; infructescence not observed.

Distribution: Papua New Guinea (East Sepik Prov.). Known only from the

type.

Habitat: Lowland riverine forest. 20-50 m altitude.

Note: In its growth form, with long pendent stems with abbreviated flowering

shoots arising in the leaf axils, Rhaphidophora stolleana resembles R.

brevispathacea. The inflorescences are, however, quite different, with the

spathe ovoid-cylindrical, 3—4 cm long, spadix slender cylindrical, 2.9—3.4

cm long in R. brevispathacea v. the spathe c. 10.2 cm long, spadix c. 8 cm

Rhaphidophora in New Guinea, Australia and Western Pacific 171

long in R. stolleana. To date, R. stolleana is known only from East Sepik

and R. brevispathacea only from Mimika/Digul Province.

29. Rhaphidophora versteegii Engl. & K. Krause

Rhaphidophora versteegii Engl. & K. Krause, Nova Guinea 8 (1910) 248,

Nova Guinea 8 (1912) 805; K. Krause & Alderw., Nova Guinea 14 (1924)

213. — Type: Indonesian Papua, Mimika/Digul border, near Sabang, 25

June 1907, Versteeg 1308 (B, holo; BO, L, K, iso).

Rhaphidophora ledermannii Engl. & K. Krause, Bot. Jahrb. Syst. 54 (1916)

81; Hay, Aroids of Papua New Guinea (1990) Pl. XIV, b, synon. nov. —

Type: Papua New Guinea, West Sepik Prov. (Felsspitz), 7 Aug. 1913,

Ledermann 12684 (B, lecto; selected here). Engler & Krause cited two

syntypes, the other, Papua New Guinea, West Sepik (Felsspitz), Aug. 1913,

Ledermann 12722, is missing from Berlin and presumed destroyed.

Figures 33 & 34

Robust, large, pachycaul, heterophyllous liane to 20 m; seedling stage not

observed; pre-adult plants shingling; adult shoot architecture comprised of

clinging, physiognomically unbranched, densely leafy flowering stems; stems

smooth, mid-green, with cataphylls and prophylls persistent and drying

dark yellow, degrading into parchment-like remains, internodes 1—4 x

0.4—2.5 cm, separated by very prominent slightly oblique leaf scars;

flagellate foraging stem absent; clasping roots densely arising from the nodes

and internodes, smooth and drying with parchment-like epidermis; feeding

roots not observed; leaves distichous; cataphylls and prophylls chartaceous,

degrading into strips of tissue and weak fibres at tips of flowering shoots;

petiole deeply canaliculate, 83—48 x 0.4—0.8 cm, smooth, with faint to rather

prominent dark dense speckling, apical geniculum prominent, basal

geniculum very large but not prominent; petiolar sheath extending to apical

geniculum, broad, chartaceous, short-persistent, degrading to papery strips

and sparse fibres, then falling; shingling lamina entire, cordiform, 2.5—7.5

x 3—6.5, chartaceous, base cordate, posterior lobes overlapping, apex obtuse

and minutely apiculate; adult lamina entire to slightly or extensively

perforated, perforations elliptic to rhombic, extending from c. + to entire

width of lamina on each side of the midrib, lamina ovate-elliptic to oblong-

lanceolate or oblong-elliptic, slightly oblique, 33—57 x 9.5—27 cm, sub-

membranous, drying pale grey-green to bright green, base rounded to acute,

apex acute to acuminate; midrib prominently raised abaxially, + flush

adaxially; primary venation pinnate, prominently raised abaxially, weakly

172 Gard. Bull. Singapore 53 (2001)

so adaxially; interprimaries sub-parallel to primaries, less prominent than

interprimaries, slightly raised abaxially and adaxially; secondary venation

feebly reticulate to subtesselate; inflorescence very rarely solitary, usually

several together, each subtended by a prominent chartaceous prophyll and

one or more chartaceous cataphylls, the entire synflorescence emerging

from a mass of dried, chartaceous cataphyll remains; peduncle slender to

stout, terete, partially to completely obscured by cataphylls, 4—12 x 0.2—

0.8 cm; spathe slender canoe-shaped, hardly to stoutly beaked, 5—10.5 x

1—2 cm, stiff-fleshy, very thick-walled (up to 1 cm at tip), yellow to yellow

green, marcescent to early fruiting, eventually falling leaving a prominent

scar; spadix stoutly cylindrical, sessile, inserted almost level on peduncle,

3—9.5 x 1—1.5 cm, white at male anthesis; stylar region conical, mostly

hexagonal in top view, 0.9—1.2 x 1—1.1 mm; stigma punctiform, very

_ prominently raised, those at the tip of the spadix even more raised, 0.1—

0.2 x c. 0.3 mm, glossy, almost black in dried material; anthers exserted at

male anthesis; infructescence stoutly oblong-cylindrical, 6—9 x 1.4—2 cm.

Distribution: Indonesian Papua, Papua New Guinea (including New

Ireland). Seemingly widespread but, based on known collections, perhaps

not common.

Habitat: Lowland to lower montane primary to secondary rain forest on

clays and silts. Sea level to 1500 m altitude.

Notes: 1. Rhaphidophora versteegii is most similar to R. spathacea in having

clusters of inflorescences subtended and interspersed by chartaceous

prophylls and cataphylls. It is readily identifiable, and distinguished from

R. spathacea, by the distinctive pale grey-green colour of dried material

and the form of the juvenile growth phase. Further, leaf lamina perforations

occur in most (but not all) individuals of R. versteegii; laminas of R.

spathacea are never perforated. The arrangement and shape of the juvenile

shingling leaves of R. versteegii are unique in the genus although known

from only one collection (Croat 5252).

2. The type of R. ledermannii differs primarily in the perforation of the leaf

lamina (R. ledermannii has profoundly perforated laminas) and as such

appears to represent distinct species. However, the occurrence of perforated

and unperforated laminas in different plants of the same species is not

rare, e.g. R. puberula Engl., R. megasperma Engl. (see Boyce 1999, 2001)

and its occurrence in R. /edermannii is not sufficient to warrant segregating

the two species. Furthermore, a suite of characters — clustered inflorescences

arising on clinging stems and subtended by chartaceous, later weakly fibrous

Rhaphidophora in New Guinea, Australia and Western Pacific 173

Figure 33. Rhaphidophora versteegii Engl. & K. Krause

A. flowering shoot x 7/,; B. leaf lamina x '/;; C. venation detail x 3; D. juvenile shingling stage,

adaxial view x '/,; E. juvenile shingling stage, abaxial view x '/,; F. inflorescence, spathe removed

x 1; G. spadix detail, post-male anthesis x 8; H. spadix detail, early fruiting x 8; J. stylar region

and stigma, side view x 4. All from Croat 52752.

174 Gard. Bull. Singapore 53 (2001)

prophylls and cataphylls, stoutly cigar-shaped spathes, prominently raised

and, compared with style diameter, large, glossy black stigmas and petioles

with small speckles (tannin cells?) - leaves little doubt that one species is

involved.

3. Perforated leaf laminas occur in a number of otherwise unrelated

Rhaphidophora species (e.g., R. foraminifera (Engl.) Engl., R. puberula, R.

versteegii, R. pertusa (Roxb.) Schott) suggesting that while a useful diagnostic

tool, lamina perforation cannot be used to circumscribe taxonomically

meaningful groups within Rhaphidophora.

Other specimens seen: INDONESIAN PAPUA. No locality, Docters van Leeuwen III74

(L); Mimika/Digul Proy. boundary: Lorentz River, near Bivakeiland, Pulle 44 (B spirit, L,

K), Pulle 67 (L, K). PAPUA NEW GUINEA. East Sepik Prov.: vicinity of Wewak, along

- trail beyond Boys Town (reform school operated by S.V.D. Missionaries), Croat 52752

(MO); Southern Highlands Prov.: Aria, near Veta, Aet 375 (L); Gulf Prov.: Baimuru,

Vailala River, 70 km at 110 from Baimuru, 50 km at 318 from Kerema, Croft et al. LAE

61251 (L, LAE); New Ireland: Lamet, NW Lavongai (New Hanover), 5km S of Lai Bay,

Croft & Lelean LAE 65461 (E, GH, K, L, LAE, M).

30. Rhaphidophora waria P.C. Boyce, sp. nov.

Vicinitate in donatione inflorescentiae condita, Rhaphidophora waria R.

spathaceae proxima est, sed foliorum laminis minoribus angustioribus magis

rigidioribus venis lateralibus primatiis eis interprimariis clare inter se

diversis, spadice stiptato, stigatibus profunde excavatis differt. — TYPUS:

Papua New Guinea, East Sepik Prov., Ambunti, along Yapa (Hunstein)

River, | Aug. 1966, Hoogland & Craven 10796 (L, holo; GH, K, L, LAE,

CANB, iso).

Figure 35

Small but robust pachycaul, homeophyllous (?) liane to unknown ultimate

height; seedling stage not observed; pre-adult plants climbing with leaves

arranged in two ranks and distinctive by the conspicuous yellow chartaceous

petiolar sheaths; adult shoot architecture comprised of clinging,

physiognomically unbranched, densely leafy flowering stems; stems with

conspicuous smooth papery epidermis and cataphylls and prophylls

persistent and drying dark yellow, then degrading into parchment-like

remains and fibres, internodes 1—2.5 x 0.3—0.9 cm, separated by almost

straight leaf scars; flagellate foraging stem absent; clasping roots densely

arising from the nodes and internodes, minutely pubescent; feeding roots

not observed; leaves distichous; cataphylls and prophylls thinly but stiffly

chartaceous, eventually degrading into strips of tissue and fibres, particularly

Rhaphidophora in New Guinea, Australia and Western Pacific 175

ye: L GURK

Figure 34. Rhaphidophora versteegii Engl. & K. Krause

A. flowering shoot x ‘/,; B. leaf lamina x ?/,; C. venation detail x 4: D. inflorescence. spathe

removed x 2: E. spadix detail, pre-anthesis x 8. A from Versteeg 1308: B-E from C roft & Lelean

LAE 65461.

176 Gard. Bull. Singapore 53 (2001)

at tips of flowering shoots; petiole deeply canaliculate, 3—15 x 0.2—0.4 cm,

smooth, apical geniculum weakly prominent, basal geniculum slightly less

so and mostly obscured by cataphyll remains; petiolar sheath extending to

apical geniculum, persistent, eventually degrading in strips leaving fibres;

lamina entire, lanceolate to elliptic, falcate, 6—23 x 1.5—5 cm, stiffly

coriaceous, apex acute, briefly acuminate; midrib raised abaxially, sunken

adaxially; primary venation pinnate, raised abaxially, slightly less raised

adaxially; interprimaries sub-parallel to primaries and less conspicuous;

secondary venation tessellate-reticulate, raised abaxially, nearly flush

abaxially; tertiary venation similar to secondary venation although more or

less invisible; inflorescences several together, each subtended by a prominent

chartaceous prophyll and one or more chartaceous cataphylls, the entire

synflorescence emerging from a mass of cataphyll remains; peduncle stout,

terete to laterally compressed, obscured by cataphylls, 2—4 x 0.3—0.35

cm; spathe canoe-shaped, slightly stoutly beaked, 4—5.5 x 1.5—2 cm, stiff-

fleshy, duration unknown; spadix cylindrical, stipitate, c. 3.2 x 0.9 cm, cream;

stipe 2—3 mm; stylar region rounded-conical, almost circular in plan view,

0.25—0.4 mm diam.; stigma deeply excavated, c. 0.2 mm diam.; anthers not

exserted at male anthesis; infructescence not observed.

Distribution: Indonesian Papua (Mimika Prov.) and Papua New Guinea

(East Sepik Prov.).

Habitat: Stunted forest and mixed heath forest on steep slopes. 200-580 m

altitude.

Notes: 1. Based on inflorescence presentation, Rhaphidophora waria 1s

close to R. spathacea but differs in the smaller, narrower, much stiffer leaf

laminas with the primary lateral and interprimary veins clearly differentiated

from one another, the stipitate spadix and the deeply excavated stigmas.

2. The specific epithet is taken from the Wasuk name ‘waria’ for the plant,

which is recorded on the type specimen label. It does not refer to the

Waria valley.

Other specimen seen: INDONESIAN PAPUA. Mimika Prov.: Freeport Concession Area,

Garcinia site on road, Johns 10418 (BO, K, MAN).

Figure 35. Rhaphidophora waria P.C. Boyce

A. flowering shoot x '/,; B. leaf lamina x 7/;; C. venation detail x 3; D. juvenile creeping stage x

\/,: E. inflorescence, spathe sectioned x 1; F. spadix detail, post-male anthesis x 8. All from

Hoogland & Craven 10796.

————————————————

178 Gard. Bull. Singapore 53 (2001)

Insufficiently known species

Rhaphidophora dahlii Engl.

Rhaphidophora dahlii Engl., Bot. Jahrb. Syst. 25 (1898) 8; K. Schum. &

Lauterbach, Fl. Schutzgeb. Siidsee (1900) 211; Engl. & K. Krause in Engl.,

Pflanzenr. 37 (IV.23B) (1908) 33. — Type: Papua New Guinea, East New

Britain, Gazelle Peninsula, Ralum, Jan. 1897, Dahl s.n. (B7, holo).

With no extant type, no spirit material in B (where the rich spirit collection

is occasionally a source of type material in the absence of a dried type

specimen), no specimens annotated by Engler or Krause (who presumably

would have been familiar with, and have annotated, authentic material),

_ and an ambiguous protologue, it is impossible to unequivocally ascribe the

name R. dahlii to any known species.

On the face of it, attempts to match Engler’s protologue to known

Rhaphidophora in the Bismarck Archipelago should be straightforward.

There are only five Rhaphidophora species present on New Britain (R.

conica, R. hayi, R. jubata, R. korthalsii and R. mima) and two (R. hayi and

R. schlechteri) on nearby New Ireland. Of these, R. korthalsii and R. hayi

may be immediately discounted since neither is remotely similar to the

vegetative characters described for R. dahlii. Of the remaining four species,

R. schlechteri can be ruled out because of its stipitate spadix (that of R.

dahlii is described as sessile), and R. jubata because it is far too large

(petiole to 85 cm long, lamina to 90 cm long, spathe 16—24 cm, spadix

13—21 v. petiole to 14 cm, lamina to 24 cm, spathe c. 4 cm, spadix to 3 cm

in R. dahlii). The remaining species, R. conica and R. mima, are both

possible candidates, but there remain several problems, not least of which

is that while in description rather similar to R. dahlii, R. conica and R.

mima are Clearly distinguishable from one another on characters such as

spathe persistence and shape of the stylar region, characters that are either

not or only scantily recorded by Engler for R. dahlii. Thus it is impossible

to be certain if either or neither is attributable to R. dahilii.

Excluded species

Rhaphidophora amplissima Schott = Epipremnum amplissimum (Schott)

Engl.

Rhaphidophora carolinensis (Volk.) Fosberg = Epipremnum carolinense

Volk.

Rhaphidophora cunninghamii Schott = Epipremnum pinnatum (L.) Engl.

Re —

Rhaphidophora in New Guinea, Australia and Western Pacific 179

Rhaphidophora koidzumii Kanehira = Epipremnum carolinense

Rhaphidophora loveilae F.M. Bailey = Epipremnum pinnatum

Rhaphidophora neocaledonica Guillaumin = Epipremnum pinnatum

Rhaphidophora pinnata (L.) Schott = Epipremnum pinnatum

Rhaphidophora pertusa var. vitiensis (Schott) Engl. = Epipremnum

pinnatum

Rhaphidophora vitiensis Schott = Epipremnum pinnatum

Rhaphidophora zippeliana Schott = Amydrium zippelianum (Schott)

Nicolson

Acknowledgements

Thanks are due to Linda Gurr, Pat Davis and Emmanuel Papadopoulos

for skillfully executing the illustrations that accompany this article, and to

Dr Alan Radcliffe-Smith for the Latin translations of the diagnoses. Thanks

also to Dr Alistair Hay (NSW) who critically reviewed the manuscript and

offered much useful advice and to Dr Dan Nicolson (US) for advice on

nomenclature.

References

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183—213.

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Monsteroideae-Monstereae) in Peninsular Malaysia, and Singapore.

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Boyce, P.C. & J. Bogner. 2000. An account of neotenic species of

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Rhaphidophora Hassk. (Araceae-Monsteroideae-Monstereae) in New

Guinea and Australia. Gardens’ Bulletin Singapore. 52: 89—100

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Das Pflanzenreich 37 (1V.23B) 4—139. Engelman, Leipzig.

Mayo, S.J., J. Bogner & P.C. Boyce. 1997. Genera of Araceae. Royal Botanic

Gardens, Kew, U.K.

Schimper, A.F.W. 1903. Guilds in Plant-geography upon a Physiological

Basis. Pt 2, ch. 2: 192—206. (English translation by W.R. Fisher, revised

and edited by P. Groom & I.B. Balfour.) Clarendon Press, Oxford, U.K.

Stearn, W.T. 1992. Botanical Latin, 4" Ed. David & Charles, Newton Abbot

& London, U.K.

| Strong, D.R. & Ray, T.S. 1975. Host tree location behavior of a tropical

vine (Monstera gigantea) by skototropism. Science. 190: 804—806.

Index Exsiccatorum

australasica = 1 hayi = 11 pachyphylla = 21

brevispathacea = 2 intonsa = 12 petrieana = 22

conica = 3 intrusa = 13 pilosa = 23

cravenschoddeana = 4 jJubata = 14 schlechteri = 24

cryptantha = 5 kokodensis = 15 spathacea = 25

discolor = 6 korthalsii = 16 spuria = 26

(Onis microspadix = 17 stenophylla = 27

geniculata = 8 mima = 18 stolleana = 28

gorokensis = 9 neoguineensis = 19 versteegii = 29

guamensis = 10 okapensis = 20 waria = 30

Aet 375 = 29; Aet & Idjan s.n. = 6; Anderson 147 = 11; Ave 4330 = 16, 4736

= 925)

Barker 145 = 16; Bateson 89, 91, 92, 93 = 18; Beccari PP 409 = 3, PP 443 =

16, Blake 19756 = 1; Boden Kioss s.n. = 17; Boraule et al. BSIP 9141 = 24;

Bourdy 244, 228 = 16; Bowden-Kerby 5983 = 25; Boyce 1447 = 5; Boyland

399 = 1; Branderhorst 324 = 8; Brass 2794 = 24, 7100, 7380, 23847 = 25,

27691, 28311 = 27, 28608.='3, 28015 = 25 260k

Cabalion 2370-= 16; Carr 12218 = 27, 13237 = 15, 14315 — 12 1400s 20.

s.n. = 15; Christophersen 402, 2956, 3004 = 26; Christophersen & Hume

Rhaphidophora in New Guinea, Australia and Western Pacific 181

1902 = 26; Clemens 2066 = 4, 4637 = 12, 7779 = 12, 8064 = 4, 10662 = 8,

11151, 41197 = 17; Coode 8037 = 25; Cowley s.n. = 1; Cowmeadow’s

collectors BSIP 3791 = 26; Cox 34 = 26; Craven & Schodde 14, 286, 375 = 4:

Groat 52575, 52614 = 1, 52752 = 29, 52760 = 5, 52763 = 16, 52763A = 4,

59707 = 25. 92/6/, J2/06 = &, 92792, 52793, 52800'= 16, 52803 = 8, 52808 =

25, 52818 = 4, 52840 = 16, 52842 = 4, 52849 = 16, 52952 = 25; Croft &

Lelean LAE 65461 = 29, NGF 34561 = 15; Croft et al. LAE 61251 = 29,

LAE 61974 = 15; Curry 939 = 16

Dissing 2776 = 16; Dyjibdja s.n. = 3; Docters van Leeuwen 10750 = 16,

11048, III74 = 29; Domstreich 77 = 4

Essig & Martin LAE 55124 = 19; Evans 581 = 25

Falanrum & Faimaw 6795 = 25; Flecker 9005 = 22; Forbes 323 = 25; Foreman

& Katik LAE 59291A = 18; Foreman & Vinas LAE 60099 = 25; Fosberg

31986, 32572 = 25; Frodin & Morren 3155 = 27, 3209 = 6; Frodin et al. 2553

= 24; Furtado s.n. = 3

Gafui BSIP 17296 = 18; Gafui et al. BSIP 10215 = 18, BSIP 12575 = 24;

Gielerup. 330 =A9: Graeffe 73 = 26, Gray 202, 1055, 2862 = 22; Green

RSNH 1267 = 16; Green s.n. = 5; Guppy 196 = 18

Hartley 13098 = 20; Henty & Streimann NGF 38835 = 19; Hind 246 = 22;

Hollrung 372 = 19, 746 = 1; Hoogland & Craven 10796 = 30; Hoogland &

Schodde 6943 = 21. Hosokawa 8334 = 16; Hunt RSS 2297 = 26, RSS 2398 =

16, RSS 2722 = 18; Hyland 5608 = 1, 9162 = 22

Isles & Vinas NGF 33829, NGF 34488 = 16

Jacobs 9331 = 8; Johns 8197 = 16, 9967 = 25, 9796 = 16, 9970 = 8, 10393 =

25, 10418 = 30, 10431 = 25

Kairo 237 = 25, 757 = 17, NGF 44078 = 24: Kairo & Streimann NGF 30715

= 19; Kajewski 584 = 26, 830 = 16, 1891, 2162 = 18; Kalkman 3389 = 21;

Kayima 154 = 25; Koidzumi s.n. = 25; Kgie & Olsen 1377 = 25

Lam 711 = 16; 763 = 19; 1014 = 24; Lauterbach 827 = 8; Lavarack &

Lavarack NGF 31091 = 18; Le Guillou ‘Triton 48’ = 25; Leach NGF 34337

= 6; Ledermann 6641 = 16, 7382 = 28, 7384 = 25, 8569 = 6, 8614 = 27, 9590

= 6, 10241 = 24, 10319, 10826 = 16, 12684 = 29, 14336 = 25; Lelean &

Streimann LAE 52541 = 12; Ludvigson 20 = 16

182 Gard. Bull. Singapore 53 (2001)

Mauriasi BSIP 8429, BSIP 12268, BSIP 13632, BSIP 13977 = 18; McKee

1877 = 1; Meebold 8209 = 26; Melville 3693 = 1; Melville & Melville 71.998

= 26; Meyer Drees 279 = 19; Millar NGF 9938 = 19; Moi et al. 196 = 8;

Morrison s.n. = 16

Nakisi & Babala BSIP 8222 = 24; Nicolson 926 = 3, 1385 = 16, 1386 = 25,

1402:=19. 1409 = 25; 1416 =21, 1927-99, 19302] 277 14S = 2a

1473 = 12, 1475 = 6, 1477 = 24, 1480 = 19, 1483 = 16, 1496 = 14, 1498 = 16,

1512 = 11, 1516 = 18, 1520 = 16, 1522 = 18, 1529, 1530=14; 1540 =11 1972

= 116, 1552.= 3, 1564= 12, 1565: =A9; 1567/1573 =1651577 = 41 bsb = 25

Parham 11246 = 26; Peekel 296 = 24; Pulle 44 = 29, 66 = 16, 67 = 29, 254 =

2, 353 = 16; Pullen 949 = 16, 5996 = 24

Raynal RSNH 16222 = 16; Reinecke 594 = 26; Ridsdale & Lavarack NGF

30628, NGF 31600 = 14; Rijkers 14841 = 11; Rinehart 15568 = 10; Robbins

870 =9

Sands 2779 = 16, 6276 = 13, 6846 = 14; Sands et al. 2044 = 24, 2384 = 11;

Schlechter 7785 = 25, 16329 = 19, 16436 = 21, 17581 = 27, 18152 = 25, 18298

= 19, 18308 = 24, 18678 = 17, 18888 = 24, 18985 = 1; Schodde 3935 = 24,

4070 = 18; Seeman 655 = 26; Smith 29, 4443, 8653, 8947 = 26, 10189 = 1;

Stone 2366 = 18, 4101 = 10; Storck 911 = 26; Streimann LAE 52789 = 19,

LAE 52964 = 8, NGF 23963 = 20, NGF 28712 = 6, NGF 44247 = 27, NGF

47654 = 25; Streimann & Kairo NGF 39190 = 7; Streimann & Womersley

LAE 51847 = 16

Takamatsu 40 = 16; Teona BSIP 6303 = 24; Teraoka & Kennedy 64 = 26

Utteridge 79 =8

Vandenberg & Katik NGF 42372 = 24; Versteeg 1191 = 8, 1308 = 29;

Versteegh BW 12647 = 23; Vinas 151 =9

Waterhouse 207, 778 = 18; Webb & Tracey 7066 = 11; Whistler 743, 1036 =

26; White et al. NGF 1555 = 1; Whitmore 6204 = 16; Widjaja 6344 = 21;

Capt. Wilkes Exped. s.n. = 26; Womersley & Simmonds 5047 = 26; Womersley

& Sleumer NGF 13971 = 9; Womersley & Whitmore BSIP 1110 = 26;

C.C.Y. Wong 278 = 16

Zippelius s.n. = 25

Rhaphidophora in New Guinea, Australia and Western Pacific 183

Index to species

Names in bold are accepted, names in italics are synonyms, numbers refer

to accepted name

GHSCHAYIO SPUTIG SCUOU ---<0c.0-..22-.0ce-0sas0creseees 162

Epipremnum multicephalum Elmet ........... 130

Pathosicélatocaulis N.E. BY ~ wo. ..c28c.i220c0--00+3- 128

WIONISIEF A TALE ORTNOED a2; 222050 ceec2s00c0sesaeseszus 130

Rhaphidophora apiculata K. Krause .......... 158

australasica F.M. Bailey .................:::0:000 88

UDI CESS LUFT peso Beene ee 158

brevispathacea Engl. & K. Krause ........... 91

buergersii Engl. & K. Krause .................. 154

celatocaulis (N.E. Br.) Alderw. ............... 128

CO TUREN LAD Wee INT AUSE occ ccs ecesncncsrsovsessienesseoos 153

CUS ety BR ee 92

BC TICLOTECIOIES IND aoa ecesaascdaca deste scdecaaiessere 130

cravenschoddeana P.C. Boyce .................. 96

cryptantha P.C. Boyce & C. Allen ........... 99

CEVA re Bee eee nee a enn one ae 178

discolor Engl. & K. Krause .................-+- 103

drepanophylla Alderw. .............::-:-+-20000+ 154

REPLI PUNC AIC cesta as tagcts ag Caiseronstivacsssascssent 158

FPEGABYECE TEAS (NOU a nner ee ee 158

ICU LEGS 80) (ee 104

PRETTIEST Oe oe. carccenracrecssceccsorane-voveteocoss 108

POROMEMSIS EC HE OY CE ctiee tt ensesceveteseseasetete: 111

SED tel haved CR oe oe see ce ee eee 163

BICUT CE OLE Ke KT AUISE scczvatsnee:asvatecsavetvosees 130

PETAAPECLESMERAC cts set cost noscectspetacursvarcuey eee eee 130

SUAMMIONISIS PC. BOYCE 2 oiecce-cecces-cscstcsveres 112

aya k © BOyce Sc BOPNEL .:eecsesesc-nscrctees 116

PIU GHTE CLINI Ls yee cece ws. stestsavesden eerariesteea6s 88

LS OLIISISM COUT AUSC tes tornco. -sschavc asco oerisvexcass 89

Heh TT EM Geni 8 (0) (el ae eee ee 119

MMNERINS BH ECCS ON CON 1 sce 25. ccege sete sereceusenteoeseass 120

AUN AEDES SOV CO: ac soc c se ponceeeaeacvacneeescnseres 124

Keene niy ae EXACUS WIA 2 cose: sectecccsescocctsasseese 158

Kokodensis Pi C. "BOYCE: firtes..d..ccbtecswsevenss 127

MOrpitalshi(SCHOU estes. eater. tte ctse<ccteetes 128

var. angustiloba Ridl. ex Engl. &

SPOp IS ANLS © ge seta eevee Ye hose cen ccepaes.etcn Sass 128

LFS) POLES NWGVES 3 eceacepecrereeereorter ter eEe Pee 130

ledermannii Engl. & K. Krause................ 171

OPUS PIPL CEA BUS sock cen secavees cu ctor ccsecaestset ec 128

MMIETOSPadlix KK [KG AUSE 22. 2ee-ceceneten-t- ccc -s- 136

AUBIN AEs CB OY CE cence ace cos tee cnc aon s-tececneoou 138

FICO RULHEETISES EMIS. eatet tk cesaees saseeeee.octeve 140

TOV OREUTCETISIS ELD eve cet caccorscteoesenesreses oe 140

TELA LETTS DER GUS Sonu as vicac soestcoescncssteasPoetcvceieaste 136

ODNGU TAMA GET We eracsripceccscseeececcicoseseetes 154

okapensis P.C. Boyce & Bogner............ 144

oreophila Engl. & K. Krause .................. 103

pachyphiylia K- Krause’: :..--22-1<..2--2-002<02- 146

palauensis Engl. & K. Krause..............-.- 158

FPTELIATECT ISIS) WCONAZ sarc ooees eres p-eoseas es ne aeeee eases 158

PLA GTI ETISTS| NICTTe percneecen cee ere eeatcerese eee 130

PALIF IV eri AP ANCE GWE ee rica cex otter -ceeesce-ceetee- 94

peekelit Eupl. & K Krause ..2.......-...0000: 154

peepla var. storckiana (Schott) Engl...... 163

Petrie amay AC Ay) ins csecesedescevszecvoseestnacses 149

PUlOSAIENC, BOYCE: s.cccceocvencsascsacecsusosavensssses 151

PEMECKEL SIGN yee op ccctcccsnnsccoceneneersscasseseres= 163

TIGIEVIANIGGI Bras revtctecssce- ce trcencreenteoenceeeseere 130

SEMIS CITEKWIE NT AUSE cree c-ncscer cscs esecnnees 153

Spathiaceal SCHOtlr sc -vseaaseceeecescsesere ese 158

spuria (Schott) Nicolson.............:::000 162

STEMI PO NVA KRAUSE recrar.cesvesccweractexcesnec 166

stolleana Engl. & K. Krause.................... 168

SLORCKAAIUA SCHOUEG tee .taes-atxaseeescn=cotsresee=-beceee 162

LEPEEEES NEMA Seance cecaser tenses. vce cocveevavacon sad eWsewonnera 128

DT INI ETAT EAU CTs scares sanxiescsocscesansevereseatestnaxam 130

UF DICETISES VIA OS OK AW A ie ercssccscunvensevccstene-vevenne 130

versteegii Engl. & K. Krause.................. 171

WU cRUN AA Est Gas OY CE tecseseccsececstsecrecretnacecnasnorse 174

wentii Engl. & K. Krause ...........:.:e0 108

Gardens’ Bulletin Singapore 53 (2001) 185-189.

Cytology of Two New Species of Musa (Musaceae) and

Their Sectional Relationship

K. JONG! AND G. ARGENT

Royal Botanic Garden Edinburgh, Edinburgh, Scotland, UK.

‘Also, Dept. Plant & Soil Science, University of Aberdeen, Scotland, UK

Abstract

Cytological observations were made on two recently described new Bornean species of

Musa L., M. monticola and M. suratii. Both have 2n = 20. Taxonomic and biogeographic

implications are discussed.

Introduction

Two new Bornean species of Musa (Musaceae) were recently described by

Argent (2000), M. suratii Argent and M. monticola |Hotta ex] Argent,

both from Sabah, Malaysia.

Their sectional placement, however, was unconfirmed. As cytological

information has historically been of major value in helping to determine

the sections and relationships in Musa taxonomy (Simmonds, 1962),

chromosome preparations of these two species were made.

Materials and Methods

Seeds collected in the field were germinated in garden compost under

glass at 28°C. Young seedlings were potted up for growing on. Root tips

were pre-treated in alpha-bromonaphthalene for 22 h at 4.5°C or in

paradichlorobenzene for 4 h at room temperature (c. 20°C), fixed in 3:1

ethanol:glacial acetic acid and stored in this fixative until required. After

hydrolysis in 5M HCl at room temperature for 30 min, the roots were

transferred to the Feulgen Reagent (prepared according to Fox, 1969) for

2 h. After washing with tap water, roots were placed in an 1:1 enzyme

mixture of 2% pectinase and 2% cellulase for 60 min. Squashing was done

in 45% acetic acid or in 0.4% aceto-carmine. Permanent slides were

prepared according to a modified freezing method outlined in Jong (1997).

186 Gard. Bull. Singapore 53 (2001)

Cytological Observations and Discussion

Musa has a range of basic chromosome numbers that is associated with the

traditional sections (Stover and Simmonds 1987) as follows:

Section Basic Number

Musa (formerly Eumusa) Baker 9. Geil i

Rhodochlamys Sagot xen

Australimusa Cheesman = NG)

Callimusa Cheesman x — 10

[Musa beccarii| Incertae Sedis Simmonds > Gam)

Ingentimusa Argent a

One section, /ngentimusa is represented only by a single species, M.

ingens Simmonds. The genus Ensete Horan has a basic number of 9, the

same as Musa beccarii N.W. Simmonds, although there is no question that

that species belongs with Ensete, its position within Musa is still problematic

since molecular and morphological data (Jarret & Gawel 1995) appear to

be in conflict. In fact, there has been a progressive tendency to amalgamate

sections Musa and Rhodochlamys (Simmonds 1962, Jarret and Gawel 1995,

Shepherd 1999). Further, Jarret and Gawel (1995) imply that Musa coccinea

Andr. is sufficiently different from the other Musa species to be placed in

a section of its own. The only basic number missing from this dysploid

series of x = 7 tomas 38)

The somatic chromosome number of both M. suratii and M. monticola

is 2n = 20 (Fig. la-ld). There are, however, certain cytological differences

between the two species that are worth noting. The chromosomes of M.

suratii are larger than those of M. monticola, and at least two pairs of

chromosomes bear satellites. There is also a gradual gradation of

chromosome size not obvious in M. monticola, where only one satellited

chromosome pair is visible. Interphase nuclei are finely granulose, with no

distinctively stained chromocentres, in contrast to those of M. monticola,

where chromocentres are obvious and numerous. Unlike M suratii, there

is a characteristic tendency for chromosomes in M. monticola to associate

in twos, often so closely together that they might be mistaken as single

chromosomes; quite frequently there is one association of three as illustrated

in Fig le and 1d.

In terms of chromosome number, both new species fall within either

sect. Callimusa or sect. Australimusa whose basic chromosome number is x

= 10. At present we have insufficient evidence to say whether the difference

in the chromosome morphology and behaviour noted above are consistent

Cytology of two new Musa Species 187

Figures la and 1b Musa suratii, metaphase, 2n = 20. 1b is a drawing of 1a, solid arrowheads

indicating satellited chromosomes. Open arrow points to chromosome partially out-of-focus.

Figures lc and 1d Musa monticola, metaphase, 2n = 20 from different roots. Note marked

close association of certain chromosomes in twos, and one in a group of three. Open arrow

points to out-of-focus chromosome. Scale bar = 10 um

or taxonomically significant between these two sections.

Section Callimusa has distinctive seed morphology with elongated,

barrel-shaped seeds that have a large oil store, which is represented in old

seed by an air space. Most of the species also have well-developed

corrugated, scarious auricles at the apex of the leaf sheath (‘shoulder’ as

defined by Argent 1976). Australimusa seeds by contrast are variable from

globose to highly angled, the leaves have variable ‘shoulders’ on the leaf

188 Gard. Bull. Singapore 53 (2001)

sheaths, which may occasionally be loosely corrugated but are never

expanded into auricles. On these morphological grounds, both species are

clearly associated with sect. Australimusa rather than sect. Callimusa.

Hotta (1987) clearly realised the relationship between Musa monticola

and the local form of M. textilis Née in Sabah and the seed, although

slightly larger and more distinctly mammilose, is very similar to that of

cultivars of M. textilis. Musa suratii has the smallest seed recorded in the

genus but the morphology is even closer to that of M. textilis, which is the

type species of sect. Australimusa. However, Musa suratii has extraordinary

morphology with green flowers, orange bracts, a very slender habit and

copious wax. All of these characters considerably expand the concept of

sect. Australimusa. The description of two more species in sect.

Australimusa from Borneo moves the emphasis of the distribution a little

-more to the west as previously five of the six known species occurred in

New Guinea and the Solomon Islands. Now with the uncertain status of

M. muluensis Hotta and M. tuberculata Hotta, it is looking as if Borneo

may actually encompass more variation in this section than is found in

New Guinea.

Acknowledgements

We wish to thank all the horticultural staff who have looked after the

living banana collections, in particular Mr D. Mitchell and Mr P. Smith

who have them in their overall charge. Mary Mendum has provided

important technical assistance for which we are most grateful.

References

Argent, G.C.G. 1976. The wild bananas of Papua New Guinea. Notes

from the Royal Botanic Garden Edinburgh. 35: 77-114.

Argent, G. 2000. Two interesting wild Musa species (Musaceae) from

Sabah, Malaysia. Gardens’ Bulletin Singapore. 52: 203-210.

Fox, D. 1969. Some characteristics of the cold hydrolysis technique for

staining plant tissues by the Feulgen reaction. Journal Histochemistry

and Cytochemistry. 17: 226-232.

Hotta, M. 1987. Distribution of the genus Musa in Malaysia. Acta

Phytotaxonomica et Geobotanica. 38: 292-302.

Jarret, R.L. and N. Gawel. 1995. Molecular markers, genetic diversity and

Cytology of two new Musa Species 189

systematics in Musa. In: Gowen, S. (ed.). Bananas and Plantains.

Chapman & Hall, U.K. Chapter 3.

Jong, K. 1997. Laboratory Manual of Plant Cytological Techniques. Royal

Botanic Garden Edinburgh, Edinburgh, U.K. vi+96 pp.

Shepherd, K. 1999. Cytogenetics of the Genus Musa. International Network

for the Improvement of Banana and Plantain, Montpellier, France.

Simmonds, N.W. 1962. The Evolution of the Bananas. Longmans, U.K..

Stover, R.H. and N.W. Simmonds.1987. Bananas. 3" ed. Longmans Scientific

& Technical, U.K.

Gardens’ Bulletin Singapore 53 (2001) 191-239.

The Genus Etlingera (Zingiberaceae) in Peninsular

Malaysia Including a New Species

S.H. KHAW

23 Jalan SS2/81

47300 Petaling Jaya, Selangor, Malaysia

Abstract

Etlingera pieeae Khaw from the Temenggor Forest Reserve in Perak, Peninsular Malaysia,

is described as a new species. Including this new species, there are 12 Etlingera species in

Peninsular Malaysia for which a key and descriptions are provided.

Introduction

Etlingera species are recognized by the presence of an involucre of large,

sterile bracts, elongate and tubular bracteoles (Fig. 1), flowers with a distinct

tube formed by the bases of the labellum and the filament above the

attachment of the corolla lobes (Fig. 2), and the labellum inrolling as the

flower withers (Burtt and Smith, 1986).

The first comprehensive systematic account of Peninsular Malaysian

species now included in Eflingera was given in the classic monograph of

Holttum (1950). His detailed account included four Phaeomeria species and

six Achasma species, now all considered as species of Etlingera (Table 1).

Holttum (1950) credited Valeton for recognizing that Phaeomeria is

closely related to Achasma. The two major characters that Holttum (1950)

used to distinguish between these two genera were the peduncle (long and

erect in Phaeomeria, short and usually subterranean in Achasma) and the

labellum (short in Phaeomeria and elongate in Achasma).

Subsequently, Burtt and Smith (1986) united the two genera Achasma

and Nicolaia Horan., which replaced Phaeomeria, an invalid name (Burtt

and Smith, 1986) with a third, Geanthus Valeton, to form a composite

genus, Etlingera Giseke. (Geanthus is not found in Peninsular Malaysia).

In uniting the three into a single composite genus, Burtt and Smith (1986)

maintained that the two characters conventionally used to divide Achasma,

Geanthus and Phaeomeria into distinct genera were inadequate. In addition,

they observed that the anther is held at an angle to the filament in Achasma,

whereas it is erect in Geanthus and Phaeomeria.

192 Gard. Bull. Singapore 53 (2001)

Table 1. Etlingera species in Peninsular Malaysia, including the names

used by Holttum.

Etlingera species Name in Holttum (1950)

E. elatior Phaeomeria speciosa

E. maingayi P. maingayi

E. fulgens P. fulgens

E. venusta P. venusta

E. pauciflora Achasma pauciflorum

E. punicea A. macrocheilos

E. subterranea A. subterraneum

E. metriocheilos A. sphaerocephalum

E. triorgyalis A. triorgyale

E. littoralis A. megalocheilos

E. corneri

E. pieeae

Further to Holttum’s enumeration of the species, while making an

inventory of gingers of the Temenggor Forest Reserve, Perak, Peninsular

Malaysia, in 1994 and 1995, two unnamed species of Etlingera were collected

(Khaw, 2000). One, E. corneri J. Mood & H. Ibrahim, was recently described

(Mood & Ibrahim 2000); the other, E. pieeae Khaw, is described here.

The new species, E. pieeae, is particularly interesting as it is in some

respects intermediate between Holttum’s Phaeomeria and Achasma. In

Peninsular Malaysia, the two are easily recognized based on inflorescence

characters (the flower head of the Phaeomeria group being either cup-

shaped on a long, aerial stalk or in the Achasma group sub-cylindric on a

short, subterranean stalk). However, the inflorescences of E. pieeae do not

have the typical appearance of either group. Instead, because its sterile

bracts are not broad but oblanceolate, spreading slightly and loosely

imbricating at the bases, a wide cup-like or a narrow sub-cylinder of sterile

bracts is not obvious. In this aspect, its inflorescence resembles that of E.

pauciflora (formerly a species of Achasma), which has the typical

subterranean inflorescence. In addition, E. pieeae is intermediate between

the two groups in the flower head having a relatively long peduncle. The

result is that the distal part of its peduncle together with the flower head

are aerial, a character of the Phaeomeria group. These features, coupled

with flowers with a long labellum, (a character of Achasma) make it distinct

from other Eltingera species in Peninsular Malaysia. The fact that it is

intermediate between the two groups adds support for uniting Phaeomeria

Etlingera (Zingiberaceae) in Peninsular Malaysia 193

and Achasma into a single genus.

Illustrated accounts of Peninsular Malaysian Etlingera were given by

Weber (1995), Larsen er al. (1999) and Lim (2000).

The attractive and colourful inflorescences and infructescences of

Etlingera species are of horticultural potential. In Peninsular Malaysia,

only kantan, E. elatior (Jack) R.M. Smith, is grown as an ornamental and

commercially for culinary purposes. The rest are not widely cultivated

horticulturally or for the cut-flower trade, although they are gaining

popularity in subtropical Australia and U.S.A..

Materials and Methods

This account is based on the examination of the living specimens. Field

observations on the morphology of the living plants and the development

of the inflorescence and infructescence were made for all the Etlingera

species, except for E. subterranea and the fruits of E. littoralis, for which

material was not available. In addition, observations were made of plants

cultivated from rhizomes and seed collected from the field. Material fixed

in Copenhagen mixture, colour photographs, as well as dried specimens

were prepared for study. Observations of fresh materials, unless otherwise

stated, form the basis of the characters used in the key and the descriptions

of the species. Drawings of dissected Etlingera flowers follow the style of

Burtt and Smith (1986). Herbarium material from the Evlingera collections

at KEP, KLU and SING were examined. Types not seen are indicated

(s.v.). Photographs of the type specimen of E. venusta at K were made by

Dr Saw Leng Guan.

Compared with Holttum’s account, several additional characters have

proved useful in identifying the species, such as the production of stilt

roots, number of flowers, the aspect of the flower head in lateral view,

shape of the basal lobes of the labellum, the form of the epigynous glands

and the scent of the crushed leaf sheaths. Examination of the minute

epigynous glands that Holttum (1950) called stylodes (nectary glands)

reveals that there are two main types in Eftlingera (Fig. 3), the one with a

rounded or irregular apex is found in the Phaeomeria group of species, the

other type with a narrowed apex with the sole exception of E. metriocheilos

is found in the Achasma group.

The description of the fruit of E. metriocheilos in the present work

differs from that of Holttum. He described the fruit as smooth based on

the examination of a single specimen (Ridley s.n. Maxwell’s Hill, June

1893), whereas examination of living material shows the fruits to be ridged

longitudinally, each ridge with a shoulder-like apical mound, covered with

194 Gard. Bull. Singapore 53 (2001)

=—erTes

Figure 1. Inflorescence types in Etlingera. A, E. maingayi; B, E. elatior, C, E. fulgens; D, E.

triorgyalis; E, E. pauciflora; F, E. punicea; G, E. metriocheilos; H, E. littoralis; (from living

material).

195

Etlingera (Zingiberacea

196 Gard. Bull. Singapore 53 (2001)

0.7 cm

0.9 cm

Figure 2. Flower and labellum types in Etlingera. A, B, E. maingayi; C, D, E. elatior; E, F, E.

fulgens; G, H, E. triorgyalis; 1, J, E. pauciflora, K, L, E. metriocheilos; M, N, E. punicea; O, P, E.

littoralis; (left, flower in lateral view; right, dissected flower; from spirit material).

Etlingera (Zingiberaceae) in Peninsular Malaysia

198 Gard. Bull. Singapore 53 (2001)

0.1 cm

0.2 cm

Figure 3. Epigynous glands in Etlingera. A, E. maingayi; B, E. fulgens; C, E. elatior; D, E. corner;

E, E. venusta; F, E. pieeae; G, E. triorgyalis; H, E. pauciflora;1, E. punicea; J, E. metriocheilos; K,

E. littoralis; (A—K, left, adaxial view; right, abaxial; from spirit material).

Etlingera (Zingiberaceae) in Peninsular Malaysia

199

200 Gard. Bull. Singapore 53 (2001)

1.5 cm

Figure 4. Infructescence types in Etlingera. A, E. maingayi; B, E. elatior; C, E. fulgens; D, E.

triorgyalis; E, F, G, E. pauciflora; H, E. punicea;1, E. metriocheilos; (from spirit material).

201

Etlingera (Zingiberaceae) in Peninsular Malaysia

202 Gard. Bull. Singapore 53 (2001)

prominent warts (Fig. 41). In addition, infructescences of three species (E.

triorgyalis, E. pauciflora and E. punicea) stated as unknown by Holttum,

are described and illustrated here (Fig. 4D, E-G, H).

Description of the Genus Etlingera in Peninsular Malaysia

Rhizomes at, just below, or deeper in the ground (E. pauciflora, E. punicea

and E. littoralis); elements long and slender between the leafy shoots in

the smaller species (E. maingayi and E. pauciflora) or stout in the larger

species; with stilt-roots above the soil, leaf-litter or swampy ground (E.

maingayi and E. corneri). Leafy shoots tall, to 6 m in the larger species,

often with swollen, bulbous bases; close together forming clumps or widely

spaced to about | m apart; sheaths ribbed longitudinally with cross-bars

- covered by tufts of short, white hairs (often clearer on young sheaths);

scent of crushed sheaths either a pleasant sour scent (all species in the

Phaeomeria group and including FE. corneri) or strong and peppery (in E.

triorgyalis and E. pieeae) or without a distinct odour; ligule bilobed or

entire; petiole usually present or lamina subsessile (E. venusta); laminas

many, large, to about | m long, often when young pink or with brownish-

pink broad bars or deep purple-red beneath (EF. fulgens and E.

metriocheilos). Inflorescence on a separate leafless shoot, arising from the

base of the leafy shoot. Peduncle covered by bladeless sheaths in two

ranks, widely spaced and not overlapping (the Phaeomeria group) or

overlapping (the Achasma group); long, slender with a short horizontal

subterranean axis, which turns upwards, forming a much longer, erect,

aerial stalk (Fig. SA) with the flower head raised well above the ground (in

the Phaeomeria group and E. corneri) or with the horizontal subterranean

part of the peduncle relatively longer (Fig. 4D), the erect part only a few

cm long with the flower head just below ground level, so that only the

opened flowers are displayed on the soil surface (the Achasma group) or

the peduncle is at first horizontal and subterranean but then erect and

producing a flower head well above the ground level (E. pieeae, Fig. 7A,

B). The flower head enclosed by broad sterile bracts; usually tightly

imbricating and forming either a capitate, wide cup-like structure (the

Phaeomeria group, Fig. 1A—C, 5B, 6B) or a narrower, sub-cylindric structure

(the Achasma group, Fig. 1D, F—H). In both cases, in side view the sterile

bracts hide a portion of the flower-head proper except in E. pauciflora, E.

subterranea and E. pieeae where the narrow lanceolate sterile bracts are

loosely imbricating at the bases only, so do not form a sub-cylindric structure

(Fig. LE, 7B), their inflorescence heads proper thus in side view are visible

between the bracts. Receptacle of inflorescence with apex inflated, rounded

Qe Oe Se aI ws,

Etlingera (Zingiberaceae) in Peninsular Malaysia 203

or flat, usually elongating only a few cm but up to 17 cm in E. elatior; with

numerous (55—200 or more) flowers per flower head in the Phaeomeria

group, 30—60 in the Achasma group, 20—35 in E. pieeae and 1—3 flowers

in E. pauciflora and E. subterranea. Sterile and fertile bracts: the outermost

sterile bracts show a transition from the uppermost sheath of the peduncle,

bracts usually not persisting until fruit maturity, coloured or whitish, erect

or spreading, much broader than the fertile bracts; outermost fertile bracts

show a transition from the sterile bracts, decreasing in size towards the

centre of the flower-head; each fertile bract producing one flower. Bracteole

tubular, two-lobed, deeply slit down the opposite side. Calyx tubular, 3-

lobed, deeply slit down the opposite side. Corolla tubular, often densely

hairy at entrance to tube; with three erect lobes (petals), the dorsal lobe

slightly wider than the lateral ones. Labellum with a short or elongated

blade, broadening at the base to form distinct basal lobes (Fig. 2, 7H) or

without basal lobes (E. venusta and E. corneri, Fig. 5H, 6J); the lowest part

of the labellum joined to the base of the stamen to form a short, fleshy

staminal tube above the bases of the corolla-lobes; usually with the basal

lobes erect on either side of the anther or folded together, partially or

entirely covering the anther; after flowering, the lower part of the blade

inrolling while the distal part withers; with the blade erect or spreading

horizontally and the apex slightly reflexed; the apex entire or bilobed.

Staminodes lacking. Filament short or in E. venusta and E. corneri almost

sessile. Anther erect or bent forward towards the labellum. Epigynous

glands as two fleshy, rounded mounds surrounding the base of the style,

each mound irregularly lobed at the apex in the Phaeomeria group (Fig.

3A-E) and E. metriocheilos (Fig. 3J) or as two fleshy, compressed blades,

not encircling the base of the style, each blade not further lobed, apex

shortly pointed in most species of the Achasma group (Fig. 3F-I, K). Ovary

glabrous or hairy, stigma rather large. /nfructescence large, usually globular

but elongate in E. elatior; fruits close together, pericarp smooth and each

with a long persistant calyx in the Phaeomeria group (Fig. 4A—C, 51, 6M)

or ridged longitudinally and with warts in the Achasma group (Fig. 4D, H,

I) or in E. pauciflora (Fig. 4E-G) and E. littoralis (Holttum, 1950) smooth;

indehiscent, fleshy or dry; maturing within about two months. Seeds

irregularly angular and truncate, each surrounded by thin, whitish,

translucent pulp.

204

la.

1b.

2a.

3b.

4a.

Ab.

Gard. Bull. Singapore 53 (2001)

Key to Etlingera species in Peninsular Malaysia

Inflorescence raised well above the ground on peduncles 40-200 cm;

sterile bracts showy, aerial, tightly imbricating, with their upper half

spreading horizontally, forming a very shallow or deep, wide, cup

(with diameter exceeding or nearly the same as the height); lip with

a'very short: blade. tc... 62.05).ceg2 te ee ee eee f)

Inflorescence almost entirely subterranean on peduncles 5—20 cm or

rarely (only in E. pieeae) raised above the ground on peduncles to

48 cm long; sterile bracts not showy, usually embedded in the ground

except for their distal parts, usually tightly imbricating, then forming

a long narrow sub-cylinder (with diameter always less than the height)

or loosely imbricating at the bases only, thus not forming a sub-

cylinder; lip wathvelongated! blade? 722.222 .22=..25. cee eee eee 6

Sterile bracts tightly imbricating at the bases only, forming a very

shallow, inconspicuous cup, with the upper half of the bracts

spreading horizontally; receptacle of inflorescence elongating

markedly during flowering forming a cone-shaped flower head within

the involucralrcup!. 2: ee. cet ete eee ee 1. E. elatior

Sterile bracts tightly imbricating, forming a firm cup; receptacle of

inflorescence hardly elongating during flowering forming a flat or

bowl-like flower head atiside view s).-t2-ee.cee.sseee enn cee 3

Involucral cup up to c. 3 cm high and c. 4 cm wide, in side view the

cup hiding a third of the flower-head; sterile bracts densely appressed

white-hairy all Over SUniaCe sem... rte ere, 2. E. maingayi

Involucral cup up to c. 9 cm high and c. 7.5 cm wide, in side view the

cup hiding almost entirely the flower-head; sterile bracts glabrous or

only sparsely white-haunyn x) ce cs5 etcetera e eee 4

Leaves with upper surfaces glossy, dark green, lower surfaces

conspicuously purple-red when young; lamina smooth, not

prominently ribbed, margin markedly crisped; sterile bracts

subrotund, strongly incurved at apex; lip with blade widening at

base to form distinct basal lobes, red with yellow margin; fruits pale

preen, densely appressed! sliomt=hanty arecrcceee see eeeee ee 3. E. fulgens

Leaves entirely green on both surfaces even when young; lamina

prominently ribbed, margin not markedly crisped; sterile bracts

oblong-elliptic, recurved at apex; lip with blade narrowed to base,

not widening to form distinct basal lobes, white with red median

band; fruits bright red ‘amd! plalonOuUs tice -e- cen cenceenoeenseeseer er aresnente 3S)

Etlingera (Zingiberaceae) in Peninsular Malaysia 205

Sa.

5b.

6a.

6b.

Ta.

7b:

8a.

8b.

9a.

Ob:

10a.

10b.

Stilt-roots absent; petiole always inconspicuous (leaf subsessile); leaf

base always cuneate; sterile and fertile bracts with acute apices,

purple-pink, pale green at the base; the involucral cup completely

a@ecayed Oclore runt matUnityn Lee eee 4. E. venusta

Stilt-roots 10-25 cm long; petiole 1.5-4 cm long; leaf base mostly

rounded (sometimes cuneate); sterile and fertile bracts with

subtruncate apices, bright red, white at the base; involucral cup

Pac Sis CML OLE MAAUUMILE Yio heehee a see ec ekeeeeascecvecseceett 5. E. corneri

Peduncles to 30 cm; fertile part of inflorescence with all sterile bracts

raised well above the ground; bracts loosely imbricating at the bases

only, not forming a sub-cylinder, in side view the flower head visible

MeV eRe RACES eae can csadenr. Aiacsnostietsceiesieonvedsdecpiceacaneedeass 6. E. pieeae

Peduncles 3-15 cm; fertile part of inflorescence always subterranean

except for distal parts of sterile bracts; bracts tightly imbricating,

forming a long, narrow sub-cylinder, in side view hiding one to two

PMMA GAO EML Ds MLO CEN CAG not e...5cscc se sadsvnsteccescsocveasvstsoneseasnceccesoesueseotses 7

MEINEM S Ol) PCr MINT OUESCOMCE,......0alusacaticevesesdareodersacevendssssossessncesesscones 8

PUOWEES IO —-OU PEK MIMOTESCEMCE, .....2....--saccccsncoveeocesedeceececeeccsereceeesecees 9

Corolla lobes 1-2 cm longer than calyx, dorsal lobe 34 x c. 1.5 cm;

lip yellow at centre with red margins. .................0 7. E. pauciflora

Corolla lobes almost same length as calyx, dorsal lobe 1-1.5 x c. 0.5

cm; lip crimson with white margins. ................0 8. E. subterranea

Corolla c. 2 cm longer than calyx; dorsal corolla lobe hiding the

entire anther; stigma hidden under dorsal corolla lobe; lip with yellow

MMCOtan And ANG TE MAT PINS. 2-....:2-...cccesscccsensseesoreseee-- 9. E. punicea

Corolla about the same length as calyx; dorsal corolla lobe hiding

only the basal part of anther; stigma clearly visible; lip entirely red

or with yellow, orangey-red or white margins, always without a yellow

EN CON ALUM TAM Preset eee eee ree RCTY .eanod es ies Soveateesttiecectesveressienats cinta 10

Leaf sheaths purple-red; lower surface of young leaves purple-red;

sterile bracts deep-red; lip purple-red with white margin; stigma

PUPle TEA aAlMOSt DIACK I... iccccscevececesscstcseecvseernes 10. E. metriocheilos

Leaf sheaths green; lower surface of young leaves entirely green or

green flushed pale-pink; sterile bracts white or pale green with pink

apices; lip entirely red or with a yellow margin; stigma pale-pink or

[DATIISHIIT TE ves saad ia Ae, ae a we eS HO 11

206 Gard. Bull. Singapore 53 (2001)

lla. Outer sterile bracts with apices recurved; lip entirely pink-red; stigma

pale-pink; crushed leaf sheaths with a distinct peppery scent; inner

surface of staminal tube densely hairy and conspicuously warty; fruits

with longitudinally ridged pericarp. .............e eee 11. E. triorgyalis

11b. Outer sterile bracts with apices not recurved; lip entirely red or with

a yellow or orange-red margin; stigma bright red; crushed leaf sheaths

without any distinct scent; only inner surface of staminal tube densely

hairy, not warty; fruits with smooth pericarp. .......... 12. E. littoralis

Description of the Species

I. __Etlingera elatior (Jack) R.M. Sm., Notes Roy. Bot. Gard. Edinb. 43

(1986) 244; Lim, Folia malaysiana 1 (2000) 4. Syn.: Alpinia elatior Jack,

‘Misc. mal. 3, 7 (1822) 2 (s.v.); Elettaria speciosa Blume, Enum. Pl. Javae

(1827) 51; Alpinia magnifica Roscoe, Monandr. Pl. (1828) t. 75 (s.v.);

Phaeomeria imperialis Lindl., Nat. Syst. Ed. 2 (1836) 446 (s.v.); Ridl., FI.

Malay Penins. 4 (1924) 272; Nicolaia imperialis Horan., Prodr. Monogr.

Scitam. (1862) 32, t. 1. (s.v.); Nicolaia speciosa Horan., Monogr. Scitam.

(1862) 32 (s.v.); Valeton, Bull. Jard. Bot. Buitenz. 3" Ser. 3 (197d SS:

Hornstedtia imperialis Ridl., in J. Str. Br. Roy. Asiat. Soc. 32 (1899) 148;

Mat. Fl. Malay. Penins. (1907) 40; Phaeomeria magnifica (Roscoe) K.

Schum., Pflanzenr. Zingib. (1904) 262; Phaeomeria speciosa (Blume) Merr.,

Enum. Philip. Pl. 1 (1922) 241; Holttum, Gard. Bull. Sing. 13 (1950) 181;

Henderson, Malayan Wild Flowers, Monocots. (1954) 153. Type: Sumatra,

Jack s.n., (s.v.) — specimen lost?

Figures 1B, 2C & D, 3C, 4B

Rhizomes just below ground; stout, 3-4 cm diam.; scale-leaves overlapping,

green. Leafy shoots 5-6 m tall; 10-18 cm apart, forming clumps; young

sheaths longitudinally ribbed, with cross-bars not covered by tufts of short,

white hairs; crushed sheaths with a pleasant, sour fragrance; ligule c. 2 cm

long, apex broad, entire or slightly bilobed, margin with short brown hairs;

petiole 2.5-3.5 cm long, surface longitudinally ribbed. Laminas c. 17 pairs;

(topmost) c. 33 x 4.5 cm, (middle) 66-81 x 15-18 cm, (basal) c. 36 x 14 cm;

upper surface with slightly raised lateral veins; entirely green on both

surfaces, (sometimes flushed pink in young leaves); lower surface glabrous;

base variable, mostly broadly rounded or cordate, sometimes cuneate or

unequal. Peduncle to 100-200 cm; shortly horizontal, then erect, aerial,

close to base of leafy shoot; sheaths c. 13, shortest at base 5—20 cm long,

apex rounded with subapical tooth 0.1 cm long, pale green at apex, dark

green towards base; not overlapping, widely spaced. Inflorescence raised

Etlingera (Zingiberaceae) in Peninsular Malaysia 207

well above the ground. Fertile apical part of inflorescence with well

developed sterile bracts tightly imbricating at the bases only, forming a

very shallow inconspicuous cup-like structure c. 2.5 cm high, 6 cm wide

and with the upper half of the bracts reflexed; in side view the cup hiding

c. 1/3 or less of the conical inflorescence head proper; the cup not persistent

until fruit maturity. Receptacle of inflorescence 4-9 cm long, lengthening to

17 cm (usually shorter); apex inflated and rounded; bearing up to 200 or

more flowers; 11-13 flowers open simultaneously; flower (from base of

ovary to apex of labellum) c. 4.5 cm long. Sterile bracts c. 13; 5.5-10 x 1.5-

3.5 cm; soft (not rigid), spreading outwards from a loosely imbricating

base; oblong-elliptic, strongly recurved; apex rounded, with subapical tooth

c. 0.5 cm long; glabrous; pink with white margins. Outer floral bracts

resembling the sterile bracts, c. 7.5 x 1-2 cm; inner floral bracts smaller c. 4

x 0.7 cm. Bracteole 22.5 x c. 1.5 cm (when flattened); unequally 2-lobed,

apex of lobe acute with c.1 cm slit on opposite side; glabrous, translucent

with a pink median band. Calyx 2.5-3 x c. 1.5 cm (when flattened); 3-

lobed, each lobe with an apical tuft of short, brown hairs; tube with c.1.5

cm slit on opposite side; glabrous; white, deep pink at apex. Corolla tube

1—1.5 cm long; inner surface densely covered with long hairs at entrance to

tube; lobes (petals) longer than calyx, 2-2.5 cm long, apices rounded, slightly

hooded, white with pink tips; dorsal lobe not hiding the anther and stigma,

c. 0.7 cm wide, lateral lobes narrower, c. 0.4 cm. Labellum held erect;

blade 1.8-2 cm long, c. 0.8 cm wide; deep red with a yellow margin except

at basal lobes; margin of blade plain; apex rounded and retuse. Staminal

tube 0.5-1 cm long above the base of the corolla-lobes; inner surface of

tube sparsely covered with long hairs, more hairy below anther, (not warty).

Filament short, c. 0.5 x 0.3 cm wide, adaxially hairy. Anther c. 0.7 x 0.3 cm;

dark red, with white hairs on pollen sacs. Epigynous glands 0.2-0.4 x 0.2-

0.3 cm; comprising 2 rounded mounds, surrounding base of style, each

mound deeply and irregularly lobed, apices glabrous, surface conspicuously

warty. Ovary 0.4-0.6 x 0.3-0.5 cm; densely covered with appressed, white,

short hairs; style white, hairy; stigma dark red, 0.3 cm across. /nfructescence

elongate, up to c. 19 cm but usually shorter x 10 cm in diam.; surmounted

by withered flowers; withered, sterile bracts persistent until fruit maturity

at base of fruit head; floral bracts persistent with fruits at their axils; number

of fruits per head 15-25 (or more). Fruits obovoid; c. 2.5 x c. 3 cm wide at

the top; densely covered with appressed short hairs; apex with remnant of

calyx c. 3 cm long; pericarp smooth, not ridged longitudinally; pericarp

0.2-0.3 cm thick; buff or pale green at maturity. Seeds irregularly angular,

truncate; c. 0.4 x 0.3 cm wide.

208 Gard. Bull. Singapore 53 (2001)

Distribution: This species is widely cultivated and has been collected from

several places in the Peninsula. Ridley (1924) reported the species wild in

forests in Rawang, Selangor and in Tambun near Ipoh and the Taiping

Hills in Perak. The species is described as native to the Peninsula, Java and

Sumatra (Ridley, 1899).

Notes: Locally known as kantan, this is commonly cultivated as a flavouring,

the young inflorescences and fruits being used in curries. The newly opened

inflorescence is beautiful but the bracts quickly turn black and rot, rendering

it unsuitable for the cut-flower trade.

One of the most conspicuous features of this species is the lengthening

of the receptacle of the inflorescence during flowering, the spike thus

forming an elongating cone within the involucral cup.

Ridley (1899 p: 148 & 149) described the lip as white edged in

Hornstedtia imperialis var. imperialis and the margins as yellow in var.

speciosa. Ibrahim (1986) reported a form with white bracts and yellow

flowers from Pulau Tioman.

Specimens seen: PERAK: Tanjong Malim, Burkill & Haniff 14002 (SING); Ipoh, Ampang

limestone cliffs, Burkill & Haniff 13934 (SING); Taiping, Batu Tegoh, Henderson 10139,

10143, 10312 (SING); Larut Hills, King’s Collector 3075 (SING); Telok Anson, Durian

Sebatang, Haniff 15293 (SING); Taiping Waterfall, Ridley s.n. (SING); Bubu F.R., Jalong,

Symington SFN 39556 (KEP). SELANGOR: Rawang, Ridley 7811 (SING); Kanching, Bukit

Takun, Stone 8787 (SING). JOHOR: Kota Tinggi, Ridley s.n. (SING). PAHANG: Pelangai

or Manchis, Burkill & Haniff 16793 (SING); Ulu Tembeling, Sg. Sat, Henderson 22068

(SING). TERENGGANU: Ulu Brang, Moysey & Kiah SFN 33873 (SING); Kuala

Terengganu, Bukit Berangan, Sinclair & Kiah SFN 40891 (SING).

2. Etlingera maingayi (Baker) R.M. Sm., Notes Roy. Bot. Gard. Edinb.

43 (1986) 247; Lim, Folia malaysiana | (2000) 5. Syn.: Amomum maingayi

Baker, in Hook. f., Fl. Brit. India 6 (1892) 235; Hornstedtia maingayi (Baker)

Ridl., in J. Str. Br. Roy. Asiat. Soc. 32 (1899) 150; Mat. Fl. Malay. Penins. 1

(1907) 41; Phaeomeria maingayi (Baker) K. Schum., Pflanzenr. Zingib. 20

(1904) 266; Ridl., Fl. Malay Penins. 4 (1924) 272; Holttum, Gard. Bull.

Sing. 13 (1950) 180; Henderson, Malayan Wild Flowers, Monocots. (1954)

155; Nicolaia maingayi (Baker) K. Larsen, Nat. Hist. Bull. Siam Soc. 23:

574 (1970). Type: Malacca, Maingay 1586, s.v. (K).

Figures 1A, 2A & B, 3A, 4A

Rhizomes long, slender, creeping, c. 2.5 cm diam.; supported by stilt-roots

10-30 cm long; scale-leaves overlapping; green flushed orange-pink. Leafy

shoots 34 m tall; 10-20 cm apart, forming clumps; young sheaths slightly

longitudinally ribbed, with cross-bars covered by tufts of very short, white

Etlingera (Zingiberaceae) in Peninsular Malaysia 209

hairs; crushed sheaths with a pleasant, sour fragrance; ligule to 1.5 cm,

apex entire or slightly bilobed, margin and surface with short, light-brown

hairs; petiole to 0.5 cm, surface longitudinally ribbed. Laminas c. 17 pairs,

(topmost) 25-62 x 3-5 cm, (middle) c. 66 x 10-12 cm, (basal) 20-43 x 5.5—

8.5 cm; upper surface smooth, entirely green (young leaves sometimes

brownish-pink all over on both surfaces or with brownish-pink bars on

green lamina), lower surface glabrous, base cuneate, unequal. Peduncle up

to c. 100 cm long, shortly horizontal, then erect, close to base of leafy

shoot; sheaths c. 9, shortest at base 2-7 cm long, apex rounded, with

subapical tooth c. 0.2 cm long, pink or purplish-red at apex, green towards

base, not overlapping, widely spaced. /nflorescence raised well above the

ground. Fertile apical part of inflorescence with well developed sterile bracts

tightly imbricating, forming a firm, cup-like structure, 2—3 x c. 4 cm; in side

view the cup hiding c. 1/3 of the inflorescence head proper, the cup persistent

until fruit maturity. Receptacle of inflorescence 1.5—2 cm long; apex inflated

and conical; bearing c. 70 flowers; c. 9 flowers open simultaneously; flower

(from base of ovary to apex of labellum) c. 4.5 cm long. Sterile bracts c. 8;

3-4 x 4.5—5 cm; rigid, erect, subrotund, the whole concave towards the

inside; apex broadly rounded, retuse, with a subapical tooth 0.2-0.5 cm

long; outer surface with densely appressed, white hairs, red. Outer floral

bracts like sterile bracts; c. 3.5 x 1.5—3 cm; inner floral bracts smaller, 3—3.5

x 0.5-1.5 cm. Bracteole c. 2 x 1 cm (when flattened); apex 2-lobed and

rounded with 0.5 cm slit on opposite side; densely, tawny hairy on entire

surface. Calyx 2.5-3 x c.1 cm (when flattened); 3-lobed, each lobe with a

glabrous, subapical tooth 0.2 cm long; tube with c.1 cm slit on opposite

side; surface densely covered with appressed, tawny hairs; red. Corolla

tube c. 1.5 cm long; inner surface densely covered with long hairs at entrance

of tube; lobes (petals) shorter than calyx, c. 1.5 cm long, apices rounded,

not hooded, red; dorsal lobe not hiding the anther and stigma, c. 0.5 cm

wide, lateral lobes narrower, c. 0.2 cm. Labellum held erect; blade 1.5-2

cm long, 0.6—0.8 cm wide; blade and basal lobes pink with a red median

band not reaching the apex, margins of base white; margins of blade crisped;

apex rounded and bilobed. Staminal tube c. 1 cm long above the base of

the corolla-lobes; inner surface of tube densely covered with long hairs,

not warty. Filament very short, c. 0.1 x 0.2 cm, adaxially hairy. Anther c. 0.8

x 0.2 cm, red, hairy on pollen sacs. Epigynous glands c. 0.3 x 0.2 cm;

comprising 2 rounded mounds, surrounding base of style, each mound

minutely lobed, apices hairy, (not warty). Ovary c. 0.3 x 0.3 cm, glabrous;

style red, hairy; stigma red, c. 0.2 cm wide. Infructescence spherical, small,

5-7 x 5.5-7 cm in diameter; surmounted by withered flowers; sterile bracts

persisting until fruit maturity at base of fruit-head; floral bracts persistent

with fruits at their axils; 12—15 fruits per head. Fruits obovoid, c. 2.5 x 2.5—

210 Gard. Bull. Singapore 53 (2001)

3.5 cm wide at the top; glabrous; apex with remnant of calyx c. 3 cm long;

pericarp smooth, not ridged longitudinally; pericarp 0.40.8 cm thick; bright

red, shiny. Seeds irregularly angular, truncate, c. 0.4 x 0.4 cm.

Distribution: In Peninsular Malaysia, this species had been collected from

many localities in lowland forests. It is also found in Thailand (Smith,

1986).

Notes: This widely distributed species has involucral/sterile bracts of variable

widths and apices. The inflorescence is recognizable from its red peduncle,

small subglobose flower-heads and pink outer bracts entirely covered with

appressed silvery-white hairs and rose-red flowers.

Lim (2000) published a new variety, var. ovata (type at KEP), which

has much broader leaves, 18.5 cm wide, compared with c. 12 cm wide and

sessile, (not petiolate) in var. maingayi. Unfortunately, vegetative characters

are less reliable than those of the inflorescence in Etlingera taxonomy.

Within the same species, some leafy shoots may produce both petiolate

and sessile leaves, as in E. pauciflora. Lim did not give any measurements

of the inflorescence or its organs for a comparative study although his

photographs showed specimens in flower. A more complete study is

required for a taxonomic decision on the status of this variety.

In the same publication, Lim (2000) rendered Holttum’s var.

longibracteata (type at SING) synonymous with var. maingayi. In his variety,

Holttum (1950) described the involucral bracts as c. 3 x 1.5 cm (v. c. 3 x 2.5

cm in var. maingayi), with the apex narrowly rounded instead of broadly

rounded. Without stating sound reasons, Lim declared that var.

longibracteata, “may be an aberration, and has so far not been found again

near the type location — where the usual form is not uncommon”. However,

not being able to relocate a taxon at the type-site (moreover, “Tembeling’

covers a large area) is insufficient grounds for eliminating Holttum’s var.

longibracteata. The variety should be maintained until detailed comparative

studies with var. maingayi are made.

Specimens seen: PERLIS: Lubuk Sireh, Hutan Simpan Mata Ayer, Wan Fadhilah HI 931

(KLU). KEDAH: Pass to Kroh from Baling, Corner SFN 31569 (SING); Yan, G. Jerai,

Batu Hampar, Saw FRI 44640 (KEP). PERAK: Upper Perak, Wray s.n. (SING); Temengor,

Ridley 14419 (SING). SELANGOR: Dusun Tua, Ridley 7801 (SING). JOHOR: Sedili

River, below Mawai, Corner SFN 36969 (2 sheets) (KEP, SING,); Batu Pahat, Ridley s.n.

(SING); Ulu Kahang, Holttum 10902 (SING); Kota Tinggi, G. Panti F.R., West, Saw FRI

37746 (KEP). KELANTAN: Kuala Lebir, Gimlette s.n. (SING); Gua Musang, Saw FRI

37403 (KEP). PAHANG: Kota Glanggi, Furtado s.n. (SING); Ulu Sg. Sat, Mohd. Shah &

Mohd. Noor MS 1772 (KEP, SING); Genting Highlands, Awana Trail, Saw FRI 37406

(KEP); Taman Negara, Sg. Relau, Chua, Mustapa & Apok FRI 40607 (KEP).

Etlingera (Zingiberaceae) in Peninsular Malaysia yA

3. Etlingera fulgens (Ridl.) C.K. Lim, Folia malaysiana 1 (2000) 6. Syn.:

Hornstedtia fulgens Ridl., J. Str. Br. Roy. Asiat. Soc. 32 (1899) 149; Mat.

Fl. Malay. Penins. (1907) 40; Phaeomeria fulgens (Ridl.) K. Schum.,

Pflanzenr. Zingib. (1904) 262; Ridl., Fl. Malay Penins. 4 (1924) 272; Holttum,

Gard. Bull. Sing. 13 (1950) 180; Nicolaia fulgens (Ridl.) K. Larsen, Nat.

Hist. Bull. Siam Soc. 23 (1970) 574. Type: Perak, Ridley s.n. (SING, iso).

Figures 1C, 2E & F, 3B, 4C

Rhizomes just below ground; c. 3 cm diam.; scale-leaves overlapping,

creamy-white. Leafy shoots 4-5 m tall; 10-14 cm apart, forming clumps;

young sheaths faintly ribbed longitudinally, with cross-bars not covered by

tufts of short, white hairs; crushed sheaths with a pleasant, sour fragrance;

ligule 1.5—2 cm, apex broad, entire or slightly bilobed, margin with short,

light brown hairs; petiole 1.5—2 cm, surface longitudinally ribbed. Laminas

c. 15 pairs; (topmost) c. 31 x 4 cm, (middle) 64-96 x 14-19 cm, (basal) 12-

28 x 4.5-8.5 cm; upper surface smooth, glossy, dark green; lower surface

glabrous; young leaves with lower surface conspicuously dark purple-red,

older leaves with only petiole and midrib purple-red; base variable, blunt

or rounded, unequal. Peduncle up to c. 100 cm, shortly horizontal at first,

then erect, aerial, close to base of leafy shoot; sheaths c. 12, shortest at

base 2.5—12 cm long, apex rounded with subapical tooth 0.1 cm long, green,

not overlapping, widely spaced. /nflorescence raised well above the ground.

Fertile apical part of inflorescence with well developed sterile bracts tightly

imbricating, forming a firm, broad cup-like structure c. 8.5 x c. 7.5 cm, in

side view the cup almost entirely hiding the inflorescence head proper, the

cup not persisting until fruit maturity. Receptacle of inflorescence c. 1.5 cm

long, apex inflated and rounded; bearing c. 60 flowers; 3-6 flowers open

simultaneously; flower (from base of ovary to apex of labellum) c. 5 cm

long. Sterile bracts c. 7; 5-7 x 3.5-5.5 cm; rigid, erect, subrotund, strongly

incurved at apex; apex broadly rounded, slightly retuse, with subapical

tooth hardly 0.1 cm long; outer surface glabrous to sparsely hairy all over

with white hairs, deep pink with green margins. Outer floral bracts

resembling sterile bracts, 4-5 x 2-3.5 cm; inner floral bracts smaller, 34 x

0.5-1.5 cm. Bracteole 2.5 x c. 1 cm (when flattened); 2-lobed, apex of lobe

acute with 1.5 cm slit on opposite side; glabrous, translucent, tinged red.

Calyx c. 4 x 1 cm (when flattened); 3-lobed, each lobe with a glabrous

subapical tooth less than 0.1 cm; tube with c. 2 cm slit on opposite side,

glabrous, red. Corolla-tube c. 1.5-2 cm long, inner surface densely covered

with long hairs at entrance into tube; lobes (petals) longer than calyx, 2 cm

long, apices rounded, slightly hooded, red with white tips; dorsal lobe not

hiding the anther and stigma, c. 0.5 cm wide, lateral lobes c. 0.3 cm. Labellum

PAD Gard. Bull. Singapore 53 (2001)

held erect; blade 1.8—2 cm long, c. 0.8 cm wide; blade and basal lobes pink-

red with a yellow margin except at basal lobes; margins of blade plain,

apex rounded and retuse. Staminal tube c. 0.5 cm long above the base of

the corolla-lobes; inner surface of tube sparsely covered with long hairs,

more hairy below the anther, (not warty). Filament short, c. 0.3 x 0.2 cm,

adaxially hairy. Anther c. 0.8 x 0.4 cm, pink-red, hairy on pollen sacs.

Epigynous glands 0.5-0.6 x 0.2 cm; comprising 2 rounded mounds,

surrounding base of style, each mound deeply and irregularly lobed, apices

hairy, surface conspicuously warty. Ovary c. 0.6 x 0.3 cm; densely appressed

hairy with white, short hairs; style red, hairy; stigma pink-red, c. 0.3 cm

across. Infructescence globular, flat-topped, large; c.11 x 13 cm in diameter;

not surmounted by withered flowers; sterile and floral bracts completely

disintegrated at fruit maturity; 30—35 fruits per head. Fruits ovoid; c. 4-5 x

2.5-3.5 cm at the middle; densely covered by appressed short hairs; apex

with remnant of calyx 3-4 cm long; pericarp smooth, (not ridged

longitudinally); pericarp 0.2—0.4 cm thick; pale green at maturity. Seeds

irregularly angular, truncate; c. 0.4 x 0.3 cm.

Distribution: In Peninsular Malaysia, this species is not common. Holttum

(1950) reported it was in cultivation in Singapore. Larsen (1970) recorded

it occurs in southern Thailand.

Notes: This species is recognized by its shiny leaves that are dark green

above and when young are conspicuously purple-red beneath but turn

green when old and have wavy margins, and by its inflorescences with

rounded, sterile bracts, strongly incurved at the apex, rose-red with greenish

white margins, forming a shallow, wide cup. The young plants sometimes

produce short stilt-roots but these are not observed in adult plants.

Smith (1986) had combined this taxon with a Javan one as Etlingera

hemisphaerica (Blume) R.M. Sm.. Lim (2000) rejected this decision and

resurrected the epithet fulgens for the Peninsular Malaysian taxon. Among

the arguments he presented were that Smith (1986), in making the

combination E. hemisphaerica, did so without having viewed the types of

either Ridley’s Malayan Hornstedtia fulgens or Blume’s Javan Elettaria

hemisphaerica. However, Lim did not make clear whether he had actually

seen the types of these two species himself nor did he give a critical

assessment of the inflorescence characters of both the species. The

arguments presented by Lim for his decision are therefore not conclusive.

Since I have not seen Blume’s type from East Java, and to avoid any

unnecessary changes until the types of both species are examined, E. fulgens,

is used in the present work for the taxon from Peninsular Malaysia.

Etlingera (Zingiberaceae) in Peninsular Malaysia 213

Specimens seen: PERAK: Jor. Batang Padang, Henderson 10854 (SING). PAHANG: Krau

Wildlife Reserve, Hulu Sg. Lompat, Chua FRI 41742 (KEP). TERENGGANU: Ulu Brang,

Moysey & Kiah SFN 33670 (SING).

4. Etlingera venusta (Ridl.) R.M. Sm., Notes Roy. Bot. Gard. Edinb. 43

(1986) 250. Syn.: Hornstedtia venusta Ridl., in J. Str. Br. Roy. Asiat. Soc.

32 (1899) 149: Mat. Fl. Malay Penins. (1907) 40: Phaeomeria venusta (Ridl.)

K. Schum., Pflanzenr. Zingib. (1904) 264; Ridl., Fl. Malay Penins. 4 (1924)

272; Holttum, Gard. Bull. Sing. 13 (1950) 182: Nicolaia venusta (Ridl.) K.

Larsen, Nat. Hist. Bull. Siam Soc. 23 (1970) 575. Type: Selangor, Genting

Bidai, Ridley 7810 (K, holo: SING, iso).

Figure 5

Rhizomes just below ground; 2.5-4.5 cm diam.; scale-leaves overlapping,

green. Leafy shoots 2.54 m tall; 10-18 cm apart, forming clumps; young

sheaths longitudinally ribbed, with cross-bars not covered by tufts of short,

white hairs; crushed sheaths with a pleasant, sour fragrance; /igule 2-3 cm:

apex deeply bilobed; margin and surface densely covered with short, brown

hairs; thin, papery, upper portion breaking off easily; petiole inconspicuous,

always subsessile. Laminas c. 10 pairs; (topmost) 51-64 x 9.5—13 cm, (middle)

66-80 x 14-20 cm, (basal) 45-54 x 13-14.5 cm; lamina prominently ribbed;

entirely green on both surfaces; lower surface glabrous; base cuneate always.

Peduncle to 60-80 cm; shortly horizontal, then erect, aerial, close to base

of leafy shoot; sheaths c. 12, shortest at base 2-9 cm long, apex rounded

with subapical tooth 0.2 cm long, flushed red at apex, green towards base,

not overlapping, widely spaced. /nflorescence raised well above the ground.

Fertile apical part of inflorescence with well developed sterile bracts, tightly

imbricating, forming a firm, deep cup-like structure 8—9 x c. 7.5 cm: in side

view the cup almost entirely hiding the inflorescence head proper; cup not

persisting until fruit maturity. Receptacle of inflorescence 2-3 cm long;

apex inflated and conical; bearing 60-70 flowers; 1-5 flowers open

simultaneously; flower (from base of ovary to apex of labellum) 5—5.3 cm

long. Sterile bracts c. 8; 7.5-12 x 4-5.5 cm; ngid, erect, oblong-elliptic,

strongly recurved at apex: apex acute, with subapical tooth c. 0.5 cm; outer

surface glabrous or sparsely covered with white hairs: bracts entirely purple-

pink or with some white towards apex, pale green at base. Outer floral

bracts resembling inner sterile bracts but narrower, 7.5-8 x 2.5-3.7 cm;

inner floral bracts smaller, 2-6 x 0.5—1.5 cm. Bracteole c. 3 x 1.5 cm (when

flattened); 2-lobed, apex of lobe acute; with c. 2 cm slit on opposite side;

glabrous, translucent, pink towards apex. Calyx 44.3 x c. 1.5 cm (when

flattened); 2- or 3-lobed, each lobe with a glabrous or hairy subapical tooth

0.1 cm; tube with 2-2.5 cm slit on opposite side; glabrous, with brown hairs

214 Gard. Bull. Singapore 53 (2001)

Figure 5. Etlingera venusta. A, base of peduncle; B, inflorescence; C, sterile bract; D, floral

bract; E, flower with bracteole, side view; F, bracteole; G, calyx; H, flower dissected with style

displaced; I, infructescence; (A from dry material, B-I from spirit material of Khaw KSH

244).

Etlingera (Zingiberaceae) in Peninsular Malaysia

2AS

216 Gard. Bull. Singapore 53 (2001)

at apex only; white, deep purple-red towards apex. Corolla-tube 1-1.2 cm

long, inner surface with long hairs at entrance into tube; lobes (petals)

shorter than calyx, c. 1.5 cm long, apices rounded, slightly hooded, white

with pink tips; the dorsal lobe not hiding the anther and stigma, 0.5—0.7 cm

wide, lateral lobes 0.4-0.5 cm. Labellum held erect; blade narrowed to

base without widening to form basal lobes; 2—2.2 x 1.2-1.3 cm; white with a

red median band not reaching the apex, without red lines radiating from

the red band towards the margin; margin slightly crisped or plain; apex

rounded, (not retuse). Staminal tube 1.2-1.3 cm long above the base of the

corolla-lobes; inner surface of tube with long hairs, (not warty). Filament

very short, c. 0.1 x 0.2-0.3 cm wide, adaxially hairy. Anther c. 1 x 0.3 cm,

pale yellow or white, with white hairs on pollen sacs. Epigynous glands c.

0.5 x 0.2 cm; comprising 2 rounded mounds, surrounding base of style,

each mound deeply and irregularly lobed, apices hairy, surface warty

towards the apices. Ovary 0.4—0.7 x 0.4—0.7 cm, glabrous; style white, hairy;

stigma pale pink, 0.3 cm across. Infructescence globular; large, c. 9 x 13-15

cm diam.; not surmounted by withered flowers; sterile and floral bracts

completely disintegrated at fruit maturity; 15—20 fruits per head. Fruits

ovoid, c. 5 x c. 3.5 cm wide at the middle; glabrous, apex with remnant of

calyx 3-4.5 cm long; pericarp smooth, not ridged longitudinally; pericarp

0.4-0.5 cm thick; bright red or pink, shiny. Seeds irregularly angular,

truncate; c. 0.4 x 0.3-0.5 cm wide, widest at apex.

Distribution: In Peninsular Malaysia, this species is collected mainly from

Perak, Selangor and Pahang. As it has not been recorded elsewhere, P.

venusta is therefore endemic to Peninsular Malaysia.

Notes: This species is distinguished from EF. corneri by the absence of stilt-

roots, its subsessile, broader leaves with cuneate bases; its rose-pink

inflorescences formed by bracts with acute apices and infructescences of

red fruits not surrounded by bracts, which completely decay during fruit

development.

Specimens seen: PERAK: Ulu Batang Padang, Ridl. 13835 (SING); Tapah Hills, Sg. Woh,

Ng FRI 1330 (KEP, SING); Tapah Hills, S. Keteh, Whitmore FRI 15668 (KEP); Hulu

Perak, Temenggor F.R., Sg. Samlor, Khaw KSH 378 (KEP). PAHANG: Genting Highlands,

Awana Hotel Trail, Saw FRI 36448 (KEP).

5. Etlingera corneri J. Mood & H. Ibrahim, Nord. J. Bot. 20 (24. 8. 2000)

278-283. Syn.: Etlingera terengannuensis C.K. Lim, Folia malaysiana | (2000)

9. Type: Thailand, Narathiwat Province, 1991, Vitoon, Lyon 91.433, s.v.

(KLU, SING).

Etlingera (Zingiberaceae) in Peninsular Malaysia Palys

Figure 6

Rhizome above leaf-litter or just below soil level, 3-4.5 cm diam.; supported

by pinkish-orange stilt-roots, 10-25 cm long; scale-leaves overlapping, green

tinged dull red. Leafy shoots 2.54 m tall; close together, 6-14 cm apart,

forming clumps; sheaths longitudinally ribbed, with cross-bars not covered

by tufts of short, white hairs, when crushed emitting a pleasant, sour

fragrance; ligule 1.5—2 cm, apex deeply bilobed, margin and surface densely

covered with short hairs; petiole 1.54 cm, surface reticulate. Laminas 10-

11 pairs; (topmost) 36-47 x 5.5-8 cm, (middle) 57-65 x 13-16 cm, (basal)

14-35 x 5-10 cm; apex shortly-pointed, c.1 cm long; base variable, mostly

rounded, occasionally unequal, sometimes cuneate; entirely green on both

surfaces when young; upper surface prominently ribbed; lower surface

glabrous; margin red, crisped, with short white or light brown hairs. Peduncle

c. 90 cm, shortly horizontal to c. 4 cm, then erect, close to base of leafy

shoot; sheaths c. 12, shortest at base, 2.5-13 cm long, apex bilobed with

subapical tooth to c. 0.5 cm long, surface with short hairs, red at apex, pale

green towards base; not overlapping, widely spaced. Inflorescence raised

well above the ground. Fertile apical part of inflorescence with well

developed sterile bracts tightly imbricating, forming a firm, deep, cup-like

structure 8-9 x c. 7.5 cm; the cup almost entirely hiding the inflorescence-

head proper in side view; the cup persistent until fruit maturity. Receptacle

of inflorescence 1-1.5 cm long; apex inflated, conical or rounded; bearing

50-60 flowers; 1-5 flowers open simultaneously; flower (from base of ovary

to apex of labellum) 5.3-6 cm long. Sterile bracts 8-14; 6.5-8.5 x 2-5.5 cm;

rigid, erect, oblong-elliptic, recurved at apex; apex subtruncate, retuse,

with subapical tooth c. 0.5 cm long; outer surface of bract glabrous, densely

covered with appressed white hairs at base only; bracts bright red, white or

pale green at base. Outer floral bracts resembling the sterile bracts but

smaller, 5.5-7.5 x 24 cm; inner floral bracts smaller still, 3.54 x 0.5-1.5

cm. Bracteole tubular at base; c. 3.5 x 1.5 cm when flattened; unequally 2-

lobed, apex of lobe acute or rounded with c. 2 cm slit on opposite side;

surface with white hairs; white with red apex. Calyx tubular at base; 44.4

x c. 1.5 cm when flattened; 3-lobed, each lobe with a glabrous or hairy

subapical tooth c. 0.1 cm long; tube with c. 2.5 cm slit on opposite side:

glabrous, with appressed white hairs at base only; white, red at apex.

Corolla-tube 1.5-2 cm long, inner surface densely covered with long hairs

at entrance into tube; lobes (petals) shorter than calyx, c. 2 cm long, apices

rounded, not hooded, pink; dorsal lobe not hiding the anther and stigma,

c. 0.5 cm wide, lateral lobes narrower, c. 0.4 cm wide. Labellum held erect,

blade narrowed to base without widening to form basal lobes; c. 2 x 1.2-1.3

cm; white with a red median band not reaching the apex, with red lines

218 Gard. Bull. Singapore 53 (2001)

Figure 6. Etlingera corneri. A, leaf base and tip; B, inflorescence; C, sterile bract, adaxial

view; D, sterile bract, lateral view; E, floral bract, adaxial view; F, flower with bracteole, lateral

view; G, flower with bracteole, adaxial view; H, bracteole, flattened, adaxial view; I, calyx,

flattened, adaxial view; J, flower dissected with style displaced; K, epigynous glands, adaxial

view; L, epigynous glands, abaxial view; M, infructescence with persistent involucral and floral

bracts displaced; (A, B from fresh material, C-L, spirit, M, dry; from Khaw KSH 601 ).

220 Gard. Bull. Singapore 53 (2001)

radiating from the red band towards the margin; margin slightly crisped;

apex rounded, slightly retuse. Staminal tube 0.9-1 cm long above the base

of the corolla-lobes, inner surface of tube densely covered with long hairs,

(not warty). Filament very short, c. 0.1 x 0.2-0.3 cm, adaxially hairy. Anther

c. | x 0.3 cm, red at apex, white at base, with yellow hairs on pollen sacs.

Epigynous glands c. 0.4 x 0.2 cm; comprising two rounded mounds,

surrounding base of style, each mound deeply and irregularly lobed, apices

glabrous, surface warty towards the apices. Ovary 0.4-0.6 x c. 0.4 cm,

glabrous, white; style white, hairy; stigma pink, c. 0.2 cm across.

Infructescence globular, large, c. 9 x 10-11 cm, not surmounted by withered

flowers, sterile bracts surrounding fruits persisting to fruit maturity, 8—17

fruits per fruit head. Fruit ovoid; c. 4 x 2.5 cm at the middle; glabrous; apex

with remnant of calyx 3-4.5 cm long; bright red or pink, shiny; pericarp

smooth, not ridged longitudinally, 0.3-0.4 cm thick. Seeds 35-45 in each

fruit; covered with a white, pulpy aril; dark brown; irregularly angular

truncate; c. 0.4 x 0.3-0.5 cm, widest at apex.

Distribution: This attractive species of horticultural potential has been

recorded in the Peninsula from the states of Kedah, Perak, Pahang and

Terengganu. It is not common and there are few collections. It is also

found in southern Thailand (Mood & Ibrahim, 2000).

Notes: The confusion over Etlingera corneri and E. venusta is discussed in

detail by Mood & Ibrahim (2000). E. corneri is distinguished from E.

venusta by its stilt roots, petiolate leaves usually with rounded bases, its

bright red inflorescence in the form of a deep cup; the outer bracts that are

strongly recurved and have subtruncate apices, and the infructescence of

red fruits surrounded by persistent bracts.

The plants from northern Perak, on which the above description is

based, have leaves that are entirely green on both surfaces, even when

young. This character differs from that described by Mood & Ibrahim

(2000): “dark green above, ... green (or purple) below ...”. For two specimens

from Johor, which were previously annotated as Phaeomeria venusta and

which were cited by Mood & Ibrahim as being E. corneri, the field notes

read: “leaves dark green, glossy with undulate edge, purple beneath, though

old leaves may lose the purple” (Corner SFN 32778, KEP, SING) and

“dark green above, purple beneath, glossy on both surfaces” (Sinclair 10596,

SING). This leaf character is reminiscent of E. fulgens. As I am uncertain

of the identity of Corner SFN 32778 and Sinclair SFN 10596, I have excluded

them from the list below of specimens seen. The field notes in Corner SFN

32778 also mention “stilts 2-4” high”, shorter than those described here.

Until Mood and Ibrahim (2000) clarified the confusion over the identities

—_

Etlingera (Zingiberaceae) in Peninsular Malaysia 2

of E. venusta and E. corneri, these two species were often misidentified.

The photographs in Weber (1995) showing the inflorescence and

infructescences of E. venusta from Sekayu, Terengganu are actually those

of E. corneri. This was noted by Mood and Ibrahim (2000). Similarly,

photographs of E. venusta infructescences in Larsen et al. (1999) are in fact

those of E. corneri, while the photograph showing “clusters of horn-like

Etlingera fruits” are the fruits of E. venusta.

Specimens seen: KEDAH: Ulu Muda F.R., Baling, Waduk, K.H. Lim, LKH 158 (KEP).

PERAK: Hulu Perak,Temenggor F.R., Pos Chiong, Sg. Sara, Khaw KSH 373 (KEP); Sg.

Tiang, Khaw KSH 586, 601, 606 (KEP). PAHANG: Kuantan, Sg. Pohoi, Mohd. Shah,

Sidek & Samsuri MS 3787 (KEP). TERENGGANU: Ulu Brang, Moysey & Kiah SFN

33721 (KEP, SING); Hulu Terengganu F.R., Hutan Lipur Sekayu, Anthonysamy SA 648

(KEP): Ulu Terengganu (Extension) F.R., Cpt. 43, Saw FRI 39880 (KEP); Ulu Terengganu

F.R., C.K. Lim LIM 4229 (KEP).

6. Etlingera pieeae Khaw, sp. nov.

Etlingera triorgyali affinis sed inflorescentia supra terram elevata, pedunculo

saltem duplo longiore, bracteis involucralibus tantum basin laxe imbricatis

ergo cupulam involucralem conspicuam non formanti, inflorescentia inter

bracteas a latere visibili, bracteis rigidis leviter expansis, oblanceolatis,

longitudinaliter inflexis non recurvatis, pallide viridis marginibus et apicibus

rubris et longitudinaliter rubro-striatis, apice inflato axis florifero conico,

flore, bractea involucrale, bracteis, bracteolis et calycibus brevioribus et

angustioribus, corolla calyce longiore loba dorsali antheram et stigma

occultanti, tubo corollae multo breviore, labello erecto loba media breviore

et angustiore, tubo staminali breviore, infructescentia minore vestigtis

bractearum involucrarum persistentibus, fructibus paucis minoribus,

pericarpio laeve non porcato recedit. Typus: Perak, Hulu Perak, Belum

F.R., Sg. Beruar, Khaw KSH 420 (holotypus KEP; isotypus KLU).

Figure 7

Rhizomes just below soil level; 3-4 cm diam.; scale-leaves overlapping,

green. Leafy shoots to 4-5 m tall; 7-18 cm apart, forming clumps; sheaths

conspicuously longitudinally ribbed with cross-bars covered by short, white

hairs; when crushed with a strong, peppery scent; ligule 2—2.5 cm long,

apex acute, margin and surface densely covered with short, brown hairs;

petiole 2.54 cm long, surface reticulate. Laminas 13-14 pairs; (topmost)

68-73 x 7-9 cm, (middle) 87-93 x 16-21 cm, (basal) c. 55 x 13 cm; when

young entirely green on both surfaces; upper surface prominently ribbed;

lower surface and margin with soft, short hairs; base cuneate, unequal;

Gard. Bull. Singapore 53 (2001)

1.6 cm

—_——

Figure 7. Etlingera pieeae S.H. Khaw, sp. nov. A, young inflorescence; B, inflorescence with

sheaths and flower-head; C, sterile bract, abaxial view; D, sterile bract, adaxial view; E, floral

bract, abaxial view; F, bracteole, flattened, abaxial view; G, calyx, flattened, abaxial view; H,

flower dissected with style displaced; I, stigma, abaxial view; J, stigma, lateral view; K, epigynous

glands, adaxial view; L, epigynous glands, abaxial view; M, infructescence with fruits surrounded

by remnants of sheaths and bracts with skeletonized veins. (A-L from spirit material of Khaw

KSH 420, M from dry material.)

sular Malaysia

Etlingera (Zingiberaceae) in Penin

224 Gard. Bull. Singapore 53 (2001)

apex shortly pointed, c. 1 cm long. Peduncle to c. 48 cm long, horizontal,

then erect, aerial, close to base of leafy shoot; sheaths c. 12, shortest at

base from 1.5-13 cm, apex rounded with a sharp, hard, subapical spine c.

0.1 cm, overlapping, green, striped deep pink, dull purple-red at apex.

Inflorescence raised well above the ground. Fertile apical part of inflorescence

with well developed sterile bracts, loosely imbricating at the bases only,

(not forming any conspicuous sub-cylindric structure), in side view the

inflorescence head proper visible between the bracts. Receptacle of

inflorescence c. 1 cm long, apex inflated and conical; bearing 20-35 flowers;

7-12 flowers open simultaneously; flower (from base of ovary to apex of

labellum) c. 9 cm long long. Sterile bracts 5-8; 9-12 x 1.5-3 cm; rigid,

spreading slightly, oblanceolate, longitudinally inflexed but not recurved;

apex rounded, retuse, crisped, with subapical hairy spine c. 0.2 cm long;

outer surface with white hairs, densely appressed hairy at apex and base;

surface pale green, apex and margin red and longitudinally striped reddish-

pink. Outer floral bracts like inner sterile bracts; 8.5—9.5 x 1.5—2.5 cm; inner

floral bracts smaller, c. 6 x 0.5—1 cm. Bracteole 5-6.5 x c. 1.5 cm (when

flattened); 2-lobed, apex of each lobe rounded with c. 3 cm slit on opposite

side; surface densely covered with tawny hairs; pink. Calyx 4.5-6 x c. 1.5

cm (when flattened); 3-lobed, each with a glabrous, subapical tooth c. 0.1

cm; tube with c. 2 cm slit on opposite side; surface densely covered with

appressed white hairs. Corolla tube c. 4 cm long, inner surface densely

covered with long hairs at entrance of tube, rest of tube glabrous; lobes

(petals) 1-2 cm longer than calyx, c. 2 cm long, apices rounded, not hooded,

margins crinkled, red; dorsal lobe hiding the anther and stigma, c. 1 cm

wide, lateral lobes narrower, c. 0.5 cm. Labellum held erect, blade elongate,

3.5—4 cm long, c. 2 cm wide; blade and basal lobes entirely bright pink-red;

margin of blade crisped and crinkled; apex broadly rounded, slightly retuse.

Staminal tube c. 0.5 cm long above the base of the corolla-lobes; inner

surface of tube densely covered with long hairs especially below the anther,

rest of surface densely covered by warts (glands?). Filament short, c. 0.3 x

0.4 cm, adaxially hairy. Anther c. 1 x 0.3 cm, white, hairy on pollen sacs.

Epigynous glands 0.5—-0.6 x c. 0.2 cm; comprising 2 fleshy, compressed

blade-like structures, not encircling base of style, each blade not further

lobed, apex glabrous, shortly-pointed, (surface not warty). Ovary c. 0.5 x

0.4 cm, densely covered with appressed, white, short hairs; style white or

pale pink, hairy; stigma pink, c. 0.4 cm across. /nfructescence a small spherical

head, c. 2.5 x 5 cm (when dry), not surmounted by withered flowers,

surrounded by remnants of sterile bracts with skeletonized veins persistent

at fruit maturity, c. 10 fruits per fruit head. Fruit obovoid; c. 1.5 x 1 cm at

the top (when dry); buff coloured; densely covered with appressed, short

hairs; apex with remnant of calyx base only, c. 0.5 cm long (when dry);

Etlingera (Zingiberaceae) in Peninsular Malaysia

pericarp smooth, not ridged longitudinally; pericarp 0.1—0.2 cm thick. Seeds

irregularly angular, truncate.

Distribution: Endemic to Peninsular Malaysia: Perak and Kedah (in Ulu

Muda F.R., Baling, Waduk, sighted only, not collected).

Ecology: By banks of shaded as well as more open, unshaded, small streams

in lowland forest at c. 300 m altitude.

Notes: Etlingera pieeae closely resembles E. triorgyalis in vegetative

characters. Both have tall leafy-shoots with sheaths that emit a strong

peppery scent when crushed. They are often found growing near each

other. However, its inflorescence shows that E. pieeae is a distinct species.

It has a combination of floral characters of both the Phaeomeria and

Achasma groups of Etlingera. It resembles the Phaeomeria group in having

a long peduncle with its flower-head held well above the ground, but has

overlapping peduncle-sheaths and a long labellum like those of the Achasma

group. Its flower-head is neither cup-shaped (typical of the Phaeomeria

group) nor sub-cylindric (most of the Achasma inflorescences). Instead, its

inflorescence with slightly spreading oblanceolate, sterile bracts, loosely

imbricating at the bases only, resembles that of E. pauciflora of the Achasma

group. This combination of features in E. pieeae from the two groups

makes it an interesting species, lending support to the union of Phaeomeria

and Achasma into a single genus, Etlingera.

Table 2 summarises the differences between E. pieeae and E.

triorgyalis, its closest relative.

This species is named for Mr. Abdullah Piee, who first discovered

this new species, in appreciation for his assistance and companionship in

ginger field trips.

Specimens seen: PERAK: Hulu Perak, Temenggor F.R., Sg. Emban, Khaw KSH 603 (KLU),

KSH 604 (KEP): Belum F.R., Sg. Beruar, Khaw KSH 420 (KEP, KLU).

7. Etlingera pauciflora (Ridl.) R.M. Sm., Notes Roy. Bot. Gard. Edinb.

43 (1986) 248. Syn.: Hornstedtia pauciflora Ridl., in J. Str. Br. Roy. Asiat.

Soc. 32 (1899) 144; Mat. Fl. Malay. Penins. (1907) 38; Fl. Malay Penins. 4

(1924) 270; Achasma pauciflorum (Ridl.) Holttum, Gard. Bull. Sing. 13

(1950) 187. Type: Selangor, Gua Batu, Ridley 8174, s.v. (K).

Figure 1E, 21 & J, 3H, 4 E-G

Rhizomes 6-13 cm below the ground; c. 0.6 cm diam.; scale leaves

overlapping, white. Leafy shoots 24 m tall; close together or widely spaced,

Gard. Bull. Singapore 53 (2001)

Table 2. Characters that distinguish Etlingera pieeae from E. triorgyalis.

E. pieeae E. triorgyalis

Peduncle length (cm) to c. 48 5-15

Inflorescence raised well above the — subterranean except for

ground apices of sterile bracts

flower-head

number of flowers

flower length (cm)

sterile bracts

form

width (cm)

colour

corolla lobes

tube length (cm)

sterile bracts loosely

imbricating at the bases

only, not forming any

conspicuous subcylinder

the flower head visible

between the bracts in side

view

20-35

rigid, spreading slightly,

oblanceolate,

longitudinally inflexed

but not recurved

1.5-3

pale green, apex and

margin red, longitudinally

striped red

1-2 cm longer than calyx

dorsal lobe hiding anther

and stigma

c. 4

labellum length (cm) 3.5—4

staminal tube

length (cm)

Infructescence size (cm)

persistence of

sterile bracts at

fruit maturity

number of fruits

Fruit

colour

size (cm)

pericarp

Co05

CALEY KS) (Ary)

the bracts partially decay,

remnants of bracts with

skeletonized veins

surrounding infructescence

ome 0)

buff

é: LS x1 (dry)

smooth, not ridged

longitudinally

and flowers

sterile bracts tightly

imbricating, forming a

narrow subcylinder,

the subcylinder hiding

c. 2/3 of the flower-head

in side view

40-50

Ge 15:5

rigid, erect, oblong-

obovate, recurved at

apical part, concave

3-6

white or pale green,

deep pink towards apex

equal to calyx

dorsal lobe not hiding

anther and stigma

5.8-6.5

(a)

eral,

em (0) >¢ 110)

bracts completely

decayed

20-35

dark red or pink

G3 % 3.Cma

ridged longitudinally,

each ridge with many

irregular fine spine-

like processes

Etlingera (Zingiberaceae) in Peninsular Malaysia 224

9-80 cm apart; sheaths slightly longitudinally ribbed with cross-bars covered

by tufts of short, white hairs (clearer in young sheaths), without distinct

scent when crushed; /igule 1-2.5 cm; apex rounded, entire; margin with

short, brown hairs; surface glabrous; lamina often sessile or with petiole

0.5—1 cm long in laminas with unequal bases, surface reticulate. Laminas c.

11 pairs; (topmost) c. 37 x 6.5 cm, (middle) 44-65 x 9-11 cm, (basal) c. 40 x

12.5 cm; upper surface prominently ribbed; entirely green on both surfaces

when young; lower surface glabrous; margins crisped, red with brown hairs;

base cuneate, sometimes unequal. Peduncle 5-10 cm long; horizontal, then

erect, entirely subterranean at base of leafy shoot or some distance away;

sheaths c. 8, shortest at base 0.5—5.5 cm, apex acute with stiff apical spine c.

0.1 cm, overlapping, white. /nflorescence entirely subterranean except for

apices of sterile bracts and flowers. Fertile apical part of inflorescence with

1-3 well developed sterile bracts loosely imbricating at the bases only, not

forming any conspicuous, sub-cylindric structure; in side view the

inflorescence head proper visible between the bracts; the sterile bracts not

persisting to fruit maturity. Receptacle of inflorescence c. 0.5 cm long; apex

inflated and rounded, bearing 1-3 flowers; 1 or 2-3 flowers open

simultaneously; flower (from base of ovary to apex of labellum) 11.5-13

cm long. Sterile bracts 1-3; 5—6 x 1-1.5 cm; rigid, spreading slightly outwards

from a loosely imbricating base; lanceolate, concave, apex acute, with a

stiff, sharp point; outer surface glabrous; white. Floral bracts like sterile

bracts, c.5 x 1 cm. Bracteole 4.5—5 x 1.5—-2 cm (when flattened); unequally

2-lobed; apex of lobe rounded with c. 3 cm slit on opposite side; glabrous;

red. Calyx 6.5-7 x 1.5 cm (when flattened); 3 lobed, each lobe with an

apical tuft of long, brown hairs; tube with c. 2.5 cm slit on opposite side;

glabrous, tawny hairs at apex only; pink, red at apex. Corolla tube 4.5—6 cm

long; inner surface densely covered with long hairs at entrance into tube,

rest of tube glabrous; lobes (petals) c. 1-2 cm longer than calyx, 34 cm

long; apices rounded, not hooded, red; the dorsal lobe hiding the anther

and stigma, c. 1.5 cm wide, lateral lobes narrower, c. 1 cm. Labellum held

horizontal; blade elongate, c. 5 cm long; c. 1.5 cm wide; blade yellow with

red margins, basal lobes yellow tinged red; margin of blade crisped and

crinkled, apex rounded, entire or slightly retuse. Staminal tube c. 1.5 cm

long above the base of the corolla lobes; inner surface of the base of the

tube densely covered with long hairs, (not warty). Filament short, c. 0.5 x

0.6 cm wide, back glabrous. Anther c. 1 x 0.5 cm; red, hairy on pollen sacs.

Epigynous glands 0.6-0.7 cm by 0.2-0.3 cm; comprising 2 fleshy, compressed,

blade-like structures, not encircling base of style, each blade not further

lobed, apex glabrous, shortly-pointed, surface not warty. Ovary c. 0.5 x 0.5

cm, with white, short hairs; style pale-pink, hairy; stigma white, 0.3 cm

across. Infructescence spherical; small, c. 4 x 4 cm in diameter; not

228 Gard. Bull. Singapore 53 (2001)

surmounted by withered flowers; bracts completely disintegrated at fruit

maturity; 1—3 fruits per head. Fruits spherical; c. 2-4 x 3-4 cm wide at the

middle; densely covered with appressed white, short hairs; apex with scar

of calyx only, hardly 0.1 cm long; pericarp smooth, not ridged longitudinally;

pericarp c. 0.1 cm thick; white. Seeds irregularly angular, truncate, c. 0.6 x

0.4 cm wide at the top.

Distribution: Endemic to Peninsular Malaysia. Although there are few

collections, where this species occurs, it is common. Weber (1995) mentions

that it has been found in Selangor, Pahang, Terengganu and Johor. It has

also been found in Perak (Khaw, 2000).

Notes: This species is easily recognized in being few-flowered, usually with

_ only 1-3 flowers. In flower coloration, it is similar to Eflingera punicea in

that the lip has a yellow centre but it is a distinct species as the leafy

shoots, inflorescence (Fig. 1E, F), flowers (Fig. 21, J, M, N) and

infructescence (Fig. 4, E-G, H) of E. pauciflora and E. punicea are entirely

different.

Specimens seen: JOHOR: Kota Tinggi, Holttum s.n. (SING); Sg. Segun, G. Panti, Corner

SFN 30890 (2 sheets) (SING). TERENGGANU: Ulu Kajang, Kemaman, Corner SFN

30430 (SING).

8. Etlingera subterranea (Holttum) R.M. Sm. Notes Roy. Bot Gard. Edinb.

43 (1986) 250. Syn.: Achasma subterraneum Holttum, Gard. Bull. Sing. 13

(1950) 187. Type: Pahang, Cameron Highlands, 5000 ft., Holttum, SFN

31212 (SING).

Distribution: This species is endemic to Peninsular Malaysia.

Notes: I have not been able to locate this species in the field. It has only

been collected twice, on both occasions from Cameron Highlands, Pahang.

According to Holttum (1950): “This is like A. sphaerocephalum in flowers,

but like A. pauciflorum in inflorescence. In leaf-characters it is rather

between A. sphaerocephalum and A. macrocheilos, having a petiole of

moderate length. ... The fruits agree exactly with the fruits of A.

megalocheilos as described in Java by Valeton (Ic. Bog. 2: t. 199), but are

only 2 or 3 instead of 12 or more”.

Other specimen seen: PAHANG: Cameron Highlands, 4,600 feet, Henderson SFN 23562

(SING).

Etlingera (Zingiberaceae) in Peninsular Malaysia 229

9. Etlingera punicea (Roxb.) R.M. Sm. Notes Roy. Bot. Gard. Edinb. 43

(1986) 250. Syn.: Alpinia punicea Roxb., Fl. Indica 1 (1820) 73; Achasma

macrocheilos Griff., Notul. Plant. Asiat. 3 (1851) 429, Ic. Plant. Asiat.

(1851) t. 357; Holttum, Gard. Bull. Sing. 13 (1950) 188; Amomum

macrocheilos Baker, FI. Brit. India 6 (1892) 235; Hornstedtia macrocheilos

Ridl., in J. Str. Br. Roy. Asiat. Soc. 32 (1899) 147; Fl. Malay Penins. 4

(1924) 271; Hornstedtia metriocheilos (Griff.) Ridl., in J. Str. Br. Roy.

Asiat. Soc. 32 (1899) 147; Fl. Malay Penins. 4 (1924) 271 (not of Griffith).

Type: Malacca, Griffith s.n., s.v. (specimen lost?).

Figure 1F, 2M & N, 31, 4H

Rhizomes to c. 12 cm below ground; slender, c. 1.5 cm diam.; overlapping

scale-leaves, green tinged dull red or pink. Leafy shoots 2-5 m tall; close

together or widely spaced, 11-40 cm apart; sheaths conspicuously ribbed

with cross bars covered by tufts of short, white hairs (clearer in young

sheaths), sometimes smooth, green or yellowish with purple blotches,

crushed sheaths without distinct scent; /igule 0.5-1 cm, apex broadly

rounded, entire, margin and surface densely covered by short, brown hairs;

petiole c. 0.5 cm or subsessile, surface reticulate. Laminas c. 10 pairs;

(topmost) c. 58 x 9 cm, (middle) 62-91 x 14-17 cm, (basal) 18-22 x 7 cm;

upper surface smooth or with slightly raised lateral veins; entirely green

(sometimes with brown bars on lower surface of young leaves); lower

surface glabrous; base cuneate, sometimes with auricles. Peduncle 5—10

cm; horizontal, then erect, entirely subterranean at base of leafy-shoot or

some distance away; sheaths c. 13, shortest at base 0.5—5 cm long, apex

acute, with a stiff spine 0.4 cm long, overlapping, pale green. /nflorescence

entirely subterranean except for apices of sterile bracts and flowers. Fertile

apical part of inflorescence with well developed sterile bracts tightly

imbricating forming a narrow sub-cylindric structure, 10-13 x c. 4 cm wide,

in side view hiding c. 2/3 or more of the inflorescence head proper; sterile

bracts persisting to fruit maturity. Receptacle of inflorescence c. 0.5 cm

long; apex inflated and rounded; bearing c. 30 flowers; 7-10 flowers open

simultaneously; flower (from base of ovary to apex of labellum) 10.5—12.5

cm long. Sterile bracts c. 5; 5.5—6 x 2-2.5 cm; rigid, erect, lanceolate, broadest

at middle, strongly concave; apex acute, with a stiff, sharp point; outer

surface sparsely covered with white hairs, more densely hairy at base;

white, pink towards apex. Outer floral bracts resembling the sterile bracts;

c. 6 x 2 cm; inner floral bracts smaller c. 5 x 1 cm. Bracteole c. 4 x 1.5 cm

(when flattened); 2-lobed, apex of lobe rounded with 1.5 cm slit on opposite

side; surface with light brown hairs; pink. Calyx c. 7 x 1.5 cm (when

flattened); 3-lobed, each lobe with a hairy, subapical tooth c. 0.1 cm; tube

with c. 3 cm slit on opposite side; glabrous, with tawny hairs only at apex.

230 Gard. Bull. Singapore 53 (2001)

Corolla-tube c. 6 cm; inner surface with hairs mostly at upper 1/3 of tube;

lobes (petals) c. 1-2 cm longer than calyx, c. 3 cm long, apices rounded, not

hooded, red; dorsal lobe hiding the anther and stigma, c. 0.8 cm wide,

lateral lobes narrower, c. 0.5 cm. Labellum held horizontal, blade elongate,

5.5-6 cm long, c. 1.5 cm wide; blade and basal lobes yellow with a narrow

scarlet margin broadening to the spatulate apex, scarlet margins of the haft

infolded so that the ‘margin’ appears to be yellow; margin of blade crisped

and crinkled; apex rounded, retuse or deeply bilobed. Staminal tube 0.8-1

cm long above the base of the corolla-lobes; inner surface of tube glabrous

or sparsely covered with long hairs, (not warty). Filament short, c. 0.3 x 0.3

cm, adaxially glabrous. Anther c. 1 x 0.5 cm; red, hairy on pollen sacs.

Epigynous glands c. 0.5 x 0.2-0.3 cm; comprising 2 fleshy compressed blade-

like structures, not encircling base of style, each blade not further lobed,

apex glabrous, shortly-pointed, surface not warty. Ovary c. 0.5 x 0.4 cm;

covered with appressed, white, short hairs; style white, hairy; stigma pink,

0.3 cm across. Infructescence globular; large c. 6 x 9 cm in diameter; not

surmounted by withered flowers; sterile bracts at base of fruit head persisting

to fruit maturity; floral bracts persistent with fruits at their axils; S5—15

fruits per head. Fruits obovoid; 3-3.5 x 3-4.5 cm wide at the top; densely

covered with appressed short hairs; apex with remnant of calyx base only,

less than 0.5 cm long; pericarp ridged longitudinally, each ridge with many

irregular, coarse, spine-like processes; pericarp c. 0.2 cm thick (excluding

ridges), pinkish-red. Seeds irregularly angular, truncate; c. 0.4 x 0.3 cm

wide at the top.

Distribution: A common species found in lowland forests throughout the

Peninsula. It also occurs in Thailand, Borneo, Java and Sumatra (Smith,

1986).

Notes: The conspicuous character of this species is the colour of the lip,

yellow in the middle, the rest scarlet. There is variation in the details of the

lip, especially the tip of the mid-lobe.

Specimens seen: PERAK: Tanjong Malim, Burkill & Haniff 13476 (SING). SELANGOR:

Genting Bidai, Ridley 7812 (SING); Genting Simpah, Hume 9722 (SING); Klang, Teluk

Reserve, Burkill SFN 5997, Bukit Rajah Forest, Burkill 15512 (SING); Ulu Gombak, Yong

FRI 99256 (KEP). NEGRI SEMBILAN: G. Tampin, Burkill 3179 (SING); Seremban, G.

Telapak Buruk, Saw FRI 40052 (KEP). JOHOR: B. Tinjau Laut, Corner SFN 37061 (2

sheets) (SING); Sg. Pelepah, G. Panti, Md. Nur 20012 (SING); Kuala Tebing Tinggi, Ridley

s.n. (SING); Batu Pahat, Ridley s.n. (SING); Sg. Kayu Ara, Mawai-Jemaluang Rd., Corner

SFN 31476 (SING); G. Pulai, Ridley s.n. (SING); G. Panti, Ridley s.n. (SING); 12 Mile

Mawai-Jemaluang Road, Corner SFN 29993 (SING); Sg. Tebrau, Ridley s.n. (SING).

TERENGGANU: Sg. Kemaman, Kampong Ayer Puteh, Corner SFN 30717 (KEP, SING).

Etlingera (Zingiberaceae) in Peninsular Malaysia 231

10. Etlingera metriocheilos (Griff.) R.M. Sm. Notes Roy. Bot Gard. Edinb.

43 (1986) 247. Syn.: Achasma metriocheilos Griff., Notul. Plant. Asiat. 3

(1851) 427, Ic. Plant Asiat. (1851) t. 356; Amomum sphaerocephalum Bak.

in Hook. f., Fl. Brit. India 6 (1892) 234; Amomum metriocheilos Bak. in

Hook. f., Fl. Brit. India 6 (1892) 234; Hornstedtia albomarginata Ridl., in

J. Str. Br. Roy. Asiat. Soc. 32 (1899) 145; Fl. Malay Penins. 4 (1924) 269;

Hornstedtia sphaerocephala (Baker) K. Schum., Pflanzenr. Zingib. (1904)

192; Achasma sphaerocephalum (Bak.) Holtt., Gard. Bull. Sing. 13 (1950)

189. Type: Malacca, Griffith 5758, s.v. (K).

Figure 1G, 2K & L 3J, 4I

Rhizomes just below ground or creeping at soil surface; stout, 24.5 cm

diam.; scale leaves overlapping, conspicuously pink, bright red or dull

purple-red. Leafy shoots 2-6 m tall; close together or widely spaced, 10-80

cm apart; sheaths dull red or green, conspicuously ribbed longitudinally

with cross-bars covered with tufts of short, white hairs or with sheaths

bright red, surface slightly reticulate; crushed sheaths without distinct scent;

ligule c. 1 cm long, apex broadly rounded, entire, margin and surface with

short, brown hairs (densely, silky-hairy in var. major); petiole 0.5-1.5 cm

(c. 3.5 cm in var. petiolata), surface reticulate. Laminas c. 15 pairs; (topmost)

22-55 x 3-4.5 cm, (middle) 57-80 x 9.5-14.5 cm, (basal) 9-25 x 5.5-8.5 cm;

smooth (prominently ribbed in var. petiolata); entirely dark green above,

when young dark purple-red beneath (or with brown-red bars on upper

surface in var. rubrostriata); lower surface glabrous (with rather rough,

short, brown hairs in var. major); base variable on the same leafy shoot,

rounded or cuneate. Peduncle 4-10 cm; horizontal, then erect, entirely

subterranean at base of leafy shoot or more usually some distance away;

sheaths c. 7, shortest at base 14 cm long, apex acute with subapical spine

to 0.2 cm, overlapping, red. /nflorescence entirely subterranean except for

apices of sterile bracts and flowers. Fertile apical part of inflorescence with

well developed sterile bracts tightly imbricating forming a narrow sub-

cylindric structure 4-6 x c. 5 cm, hiding c. 1/3 of the inflorescence head

proper in side view; sterile bracts persisting to fruit maturity. Receptacle of

inflorescence c. 2 cm long; apex inflated and conical; bearing 25—60 flowers;

5-6 flowers open simultaneously; flower (from base of ovary to apex of

labellum) 8-10.5 cm long. Sterile bracts c. 8; 3-4.5 x 2-3 cm: rigid, erect,

oblong-elliptic, broadest below middle, strongly concave; apex acute, with

a stiff spine 0.2 cm long; outer surface densely covered with appressed,

brown hairs, hairs golden brown at base of bract; entirely red, or red at

base and deep purple-red at apex. Outer floral bracts resembling the sterile

bracts, c. 4.5 x 1.5 cm: inner floral bracts smaller, c. 4.x 1 cm. Bracteole 2.5—

3 x c. 1.5 cm (when flattened); unequally 2-lobed, apex of lobe acute with

o>)

Gard. Bull. Singapore 53 (2001)

c. | cm slit on opposite side; surface with brown hairs, red. Calyx 5-6.5 x

1.5 cm (when flattened); 3-lobed, each lobe with a glabrous, subapical

tooth c. 0.2 cm long; tube with 2.5-3 cm slit on opposite side; glabrous,

only the apex with tawny hairs; red. Corolla tube 2.5-3.5 cm long; inner

surface of tube at the entrance sparsely hairy, rest of tube glabrous; lobes

(petals) variable: equal to, shorter or longer than calyx, 2-2.5 cm long,

apices rounded, not hooded, red; dorsal lobe not hiding the anther and

stigma, c. 0.5 cm wide, lateral lobes narrower, c. 0.4 cm. Labellum held

horizontal, blade elongate, 3.5-5 cm long, c. 1.5—-1.8 cm wide; blade and

basal lobes crimson, margins of lobes white; margin of blade plain or slightly

crisped; apex retuse or deeply bilobed. Staminal tube 1.5—-2 cm long above

the base of the corolla-lobes, inner surface of tube glabrous or sparsely

covered with long hairs, (not warty). Filament short, c. 0.6 x 0.3 cm wide,

adaxially glabrous. Anther c. 1 x 0.5 cm; crimson, hairy on pollen sacs.

Epigynous glands 0.3-0.4 x 0.3 cm; comprising 2 rounded mounds,

surrounding base of style, each mound minutely lobed, apices hairy, surface

not warty. Ovary c. 0.5 x 0.4 cm; densely covered with appressed, white,

short hairs; style pink, hairy; stigma dark purple, almost black, 0.4 cm

across. Infructescence globular; large, c. 7 x 9 cm diam.; not surmounted by

withered flowers; sterile bracts at base of fruit head and floral bracts

persisting to fruit maturity; 25—45 fruits per head. Fruits obovoid; 3+4 x c.

2.5 cm wide at the top; densely covered with appressed, short hairs; apex

with remnant of the base of the calyx only, less than 0.5 cm long; pericarp

ridged longitudinally, each ridge with a shoulder-like apical mound, covered

in prominent warts; pericarp c. 0.2 cm thick (excluding ridges); dark purple.

Seeds irregularly angular, truncate, c. 0.4 x 0.3 cm wide at the top.

Distribution: Found in many parts of the Peninsula, in lowland forests and

at moderate elevations on the mountains. It is also found in Borneo (Smith,

1986).

Notes: This species is variable and Holttum (1950) described four varieties,

ranging from the few-flowered, small-sized var. rubrostriata, to the robust

var. petiolata. The distinguishing characters of this species are: (1) the

striking red or purple lower surface of young leaves and leaf-sheaths except

for var. rubrostriata, which has a red-barred upper surface and is green

beneath, (2) a sub-globose or sub-cylindric inflorescence, (3) the deep red

or crimson involucral bracts that are distinctly shorter than the calyces,

and (4) the deep crimson lip with white (never yellow) margins.

While noting it was possible that Achasma sphaerocephalum (Baker)

Holttum was identical with A. metriocheilos Griff., Holttum (1950 p. 191)

nevertheless thought it better to exclude Griffith’s name. The reason he

Etlingera (Zingiberaceae) in Peninsular Malaysia 233

cited was that the details given by Griffith were inadequate for him to

decide with certainty whether the two species were conspecific.

Smith (1986 p. 247) combined Achasma metriocheilos Griff. with

Amomum sphaerocephalum Bak. as Etlingera metriocheilos (Griff.) R.M.

Sm.. However, she included in the combination, Hornstedtia metriocheilos

(Gniff.) Ridl., the species which Ridley (1899 p. 147) described as having a

crimson lip with a yellow centre. However, this is a different species and,

as Holttum had noted, it is in fact Etlingera punicea.

Types seen: Syntypes of Hornstedtia albomarginata Ridl., J. Str. Br. Roy.

Asiat. Soc. 32 (1899) 145. — Penang, road to Penara Bukit, Ridley s.n.

(SING); Selangor, Petaling, Ridley s.n. (SING).

Achasma sphaerocephalum (Bak.) var. petiolatum Holttum, in Gard. Bull.

Sing. 13 (1950) 191. — Johor, Ulu Segun, G. Panti, Corner SFN 30745

(SING).

var. rubrostriatum Holttum, in Gard.

Bull. Sing. 13 (1950) 190. — Selangor, K. Kubu side of the Gap, Corner

SFN 30776 (SING).

var. grandiflorum Holttum, in Gard.

Bull. Sing. 13 (1950) 191. — Terengganu, Kemaman, Bukit Kajang, Corner

SFN 30234 (SING).

var. majus Holttum, in Gard. Bull. Sing.

13 (1950) 190. — Terengganu, Kemaman, Bukit Kajang, Corner SFN 30205

(SING).

Other specimens seen: PERAK: Maxwell's Hill, Coll.? s.n. (SING); Tea Gardens, Curtis s.n.

(SING); Kuala Kangsar, Kg. Hitam Jong, Sg. Plus F.R., Hashim Pendek KLU 119 (KEP).

SELANGOR: Petaling, Ridley s.n. (SING); UPM campus, Anthonysamy SA 199 (KEP).

11. Etlingera triorgyalis (Baker) R.M. Sm. Notes Roy. Bot. Gard. Edinb.

43 (1986) 250. Amomum triorgyale Baker, in Hook. f., Fl. Brit. India 6

(1892) 237; Hornstedtia triorgyale (Baker) Ridl., in J. Str. Br. Roy. Asiat.

Soc. 32 (1899) 144; Mat. Fl. Malay. Penins. 1 (1907) 38; Fl. Malay Penins. 4

(1924) 269; Achasma triorgyale (Baker) Holttum, Gard. Bull. Sing. 13 (1950)

186. Type: Perak, Larut, King’s Collector 2105, s.v. (K).

Figure 1D, 2G & H, 3G, 4D

Rhizomes just below ground; stout, 2.5—4.5 cm diam.; scale leaves

overlapping, green tinged dull red or pink. Leafy shoots 5-6 m tall: close

together or widely spaced to c. 1 m apart; sheaths conspicuously ribbed

longitudinally with cross-bars covered by tufts of short, white hairs

(especially in young sheaths), when crushed with a strong scent suggestive

234 Gard. Bull. Singapore 53 (2001)

of pepper; ligule 1-2.5 cm; apex rounded, entire; margin and surface with

short, brown hairs; petiole 1.5—3 cm, surface reticulate. Laminas c. 15 pairs;

(topmost) c. 43 x 3.5 cm, (middle) 80-100 x 6—21.5 cm, (basal) c. 25 x 7 cm;

smooth or with slightly raised lateral veins, when young entirely green on

both surfaces; lower surface softly hairy with short hairs; margin broadly

crisped, red with brown hairs; base broadly cuneate, unequal, slightly

decurrent. Peduncle short, 5-15 cm; horizontal, then erect, entirely

subterranean at base of leafy shoot; sheaths c. 14, shortest at base 4-7.5 cm

long, apex acute with subapical spine 0.2—1 cm long, white or pale green

tinged deep pink, overlapping. /nflorescence subterranean except for distal

parts of sterile bracts and flowers. Fertile apical part of inflorescence with

well developed sterile bracts tightly imbricating, forming a narrow sub-

cylindric structure 10-14 x c. 6 cm wide, in side view hiding c. 2/3 or more

of the inflorescence-head proper, the sterile bracts not persisting to fruit

maturity. Receptacle of inflorescence c. 1 cm, apex inflated and rounded;

bearing 40-50 flowers; 8-17 flowers open simultaneously; flower (from

base of ovary to apex of labellum) c. 13.5 cm long. Sterile bracts c. 7; 9-11 x

3-6 cm; rigid, erect, oblong-obovate, recurved at apex, concave, apex

broadly rounded, retuse, with subapical spine c. 0.1 cm; outer surface with

white hairs, hairs densely appressed at base; white or pale green, deep

pink towards apex. Outer floral bracts resembling the sterile bracts, c. 10.5

x 2.5 cm; inner floral bracts smaller, c. 8.5 x 1 cm. Bracteole c. 8 x 2 cm

(when flattened); 2-lobed, apex of lobe rounded with c. 2 cm slit on opposite

side; surface with light-brown hairs; white with red apex. Calyx 7.5-9 x c. 2

cm (when flattened); 3-lobed, each lobe with a hairy, subapical tooth c. 0.1

cm long; tube with c. 4.5 cm slit on opposite side; glabrous, with tawny-

hairs at apex only; white, pink-red at apex. Corolla tube 5.8-6.5 cm long,

inner surface densely covered with long hairs at entrance of tube, rest of

tube glabrous; lobes (petals) equal to calyx, c. 2.8 cm long, apices rounded,

not hooded, red; dorsal lobe not hiding the anther and stigma, c. 1 cm

wide, lateral lobes narrower, c. 0.7 cm. Labellum held horizontal, blade

elongate, c. 5 cm long, c. 2.5 cm wide; blade and basal lobes entirely bright

pink-red; margin of blade crisped and crinkled, apex broadly rounded and

retuse. Staminal tube c. 1.2 cm long above the base of the corolla lobes;

inner surface of tube densely covered with long hairs below anther only,

rest of surface densely covered by warts. Filament short, 0.4 x 0.3 cm,

adaxially hairy. Anther c.1 x 0.5 cm, red. Epigynous glands c.1 x 0.2-0.3 cm;

comprising 2 fleshy compressed blade-like structures, not encircling base

of style, each blade not further lobed, apex glabrous, shortly-pointed, surface

not warty. Ovary c. | x 0.5 cm, densely covered with appressed, white short

hairs; style pale pink, hairy; stigma pale pink, c. 0.5 cm across. Infructescence

globular, large, c. 6 x 10 cm diam.; not surmounted by withered flowers;

Etlingera (Zingiberaceae) in Peninsular Malaysia 235

bracts completely disintegrated at fruit maturity; 20—35 fruits per head.

Fruits obovoid; c. 3 x 2.5-3 cm wide at the top; densely covered with

appressed, short hairs; apex with remnant of calyx base only, less than c.

0.5 cm long; pericarp ridged longitudinally, each ridge with many irregular,

fine spine-like processes; pericarp c. 0.2 cm thick (excluding ridges), dark

red or pink. Seeds irregularly angular, truncate c. 0.4 x 0.3 cm wide at the

top.

Distribution: In Peninsular Malaysia, recorded from Perak, Selangor and

Pahang. It also occurs in Borneo and Sumatra (Smith, 1986).

Notes: The very broad bracts, recurved at the deep rose apices, together

with the cherry red or deep rose lips are striking characters of this species.

Specimens seen: PERAK: Ipoh, foot of limestone rocks, Curtis SFN 3317 (SING).

SELANGOR: Genting Peras, Ridley 7806 (SING). PAHANG: Ulu Sg. Sat, Mohd. Shah &

Mohd. Noor MS 1880 (KEP, SING); Temerloh, Sg. Nering, Henderson 10583 (SING).

12. Etlingera littoralis (K6nig) Giseke. Syn.: Hornstedtia megalochilus Ridl.,

in J. Str. Br. Roy. Asiat. Soc. 32 (1899) 146; Mat. Fl. Malay. Penins. 1

(1907) 38; Fl. Malay Penins. 4 (1924) 270; Amomum megalocheilos Baker,

Fl. Brit. India 6 (1892) 236; Achasma megalocheilos Griff., Notul. Plant.

Asiat. 3 (1851) 426, Ic. Plant. Asiat. (1851) t. 355; Valeton., Ic. Bog. 2

(1903) t. 188, 199.

Figure 1H, 20 & P, 3K

Rhizomes at surface or below ground to c. 10 cm deep; thick, 3-3.5 cm

diam.; scale-leaves overlapping, green tinged dull red. Leafy shoots 3-6 m

tall; close together or widely spaced, 12-48 cm apart; sheaths conspicuously

ribbed longitudinally, with cross-bars covered by tufts of short, white hairs

(clearer in young sheaths); crushed sheaths without distinct scent; ligule

1.5-2 cm long; apex broadly rounded, entire; margin with short brown

hairs, surface with purple blotches, glabrous; petiole 1.5-4.5 cm; surface

reticulate. Laminas c. 13 pairs; (topmost) c. 69 x 8 cm, (middle) c. 101 x 18

cm, (basal) c. 27 x 7 cm; smooth; entirely green, in young leaves sometimes

flushed pink beneath; mostly glabrous, sometimes with short hairs; base

variable on the same leafy shoot, cuneate, rounded or truncate, always

unequal. Peduncle 4-10 cm; horizontal, then erect, entirely subterranean,

at base of leafy shoot or some distance away; sheaths c. 11, shortest at base

from 1.5—6 cm, apex acute with a stiff subapical spine 0.1 cm, overlapping,

pale green or white tinged pink. /nflorescence subterranean except for

apices of sterile bracts and flowers. Fertile apical part of inflorescence with

236 Gard. Bull. Singapore 53 (2001)

well developed sterile bracts tightly imbricating forming a narrow sub-

cylindric structure c. 12 x 5 cm, in side view hiding c. 2/3 or more of the

inflorescence-head proper. Receptacle of inflorescence c. 0.5 cm; apex

inflated and rounded; bearing 30-35 flowers; 5-9 flowers open

simultaneously; flower (from base of ovary to apex of labellum) 13-16 cm

long. Sterile bracts 5 or 6; 6-8.5 x 2-5 cm; rigid, erect, oblong-obovate,

broadest at middle, concave; apex acute, with a stiff spine 0.1 cm long;

outer surface glabrous, densely covered with appressed, white hairs at base

only; white, pink towards apex. Outer floral bracts resembling the sterile

bracts; c. 8.5 x 1.5-2.5 cm; inner floral bracts smaller, c. 8 x 0.5 cm. Bracteole

7-7.5x c.1.5cm wide (when flattened); 2-lobed, apex of lobe acute with 3

cm slit on opposite side; surface with brown hairs; pale pink. Calyx 7-9 x c.

1.5 cm (when flattened); 3-lobed; subapical tooth inconspicuous; tube with

_ c.3 cm slit on opposite side; glabrous, with tawny hairs at apex only; pink.

Corolla tube 6-6.8 cm long, inner surface densely covered with long hairs

at entrance oftube, rest of tube sparsely hairy; lobes (petals) equal to calyx,

c. 2.5 cm long, apices rounded, not hooded, pink; dorsal lobe not hiding

the anther and stigma, c. 1 cm wide, lateral lobes narrower, c. 0.5 cm.

Labellum held horizontal, blade elongate, c. 7 cm long, c. 2 cm wide; blade

and basal lobes entirely red or with a yellow margin sometimes not reaching

the broad apex; margin of blade plain or crisped and crinkled; apex rounded,

entire or slightly retuse. Staminal tube c. 1.5 cm long above the base of the

corolla-lobes; inner surface of tube densely covered with long hairs, (not

warty). Filament short, c. 0.5 x 0.4 cm wide, adaxially hairy. Anther c. 1 x

0.5 cm; red, hairy on pollen sacs. Epigynous glands c. 1 x 0.2-0.3 cm;

comprising 2 fleshy, compressed, blade-like structures, not encircling base

of style, each blade not further lobed, apex glabrous, shortly pointed, surface

not warty. Ovary 0.5—0.6 x 0.5 cm; densely covered with appressed, yellow,

short hairs; style pink, hairy; stigma red, 0.3 cm across. Fruit not known.

Distribution: Common in lowland forests throughout the Peninsula. It is

also found in China, Thailand, Borneo, Java and Sumatra (Smith, 1986).

Notes: There is variation in lip colour in this species. Most commonly

encountered are plants with scarlet lips with a yellow margin. However,

plants growing nearby may have scarlet lips edged orange-red or a third

type with much narrower lips are scarlet with a thin yellow margin.

The description of the infructescence by Holttum (1950) was based

on fruits described from Java by Valeton. I have attempted in the field to

find the infructescence of this common species but without success. The

inflorescences in a large patch growing in the Temenggor F.R., Perak,

Etlingera (Zingiberaceae) in Peninsular Malaysia 235]

were tagged and the site revisited two months later to look for the fruits

but none was found. The fact that it has never been found in fruit in

Peninsular Malaysia warrants further investigation.

Specimens seen: PERLIS: Wang Kelian, Wan Fadhilah HI 919 (KLU): Mata Ayer F.R.,

Saw, Kamarudin, Jamaludin & Baya FRI s.n. (KEP). PERAK: Tambun limestone cliff,

Burkill SFN 6294 (SING); Upper Perak, Wray 3444 (SING); SELANGOR: Gua Batu,

Ridley s.n. (SING); Kuala Lumpur, Ridley s.n. (SING). MALACCA: Bukit Sadanen, Ridley

s.n. (SING). JOHOR: Castleroad, Ridley s.n. (SING); Sg. Segun, G. Panti, Corner SFN

30891, 30892 (SING); Batu Pahat, Ridley s.n. (SING); Kota Tinggi, Ridley s.n. (SING).

KELANTAN: Sg. Lebir, Ben. Stone & Mahmud Sidek 12507 (KEP). PAHANG: Jerantut,

Corner SFN 30768 (SING); Temerloh, Titi Bungor, Henderson 10674 (SING); Telom, Ridley

13832 (SING); Tanjong Gajah, Ridley s.n. (SING); Sg. Pertang, Bentong, Burkill & Haniff

SFN 16515 (SING); Pekan, Ridley 1632 (SING); Tahan Woods, Ridley s.n. (SING).

TERENGGANU: Kemaman, Sg. Nipa, Corner SFN 30581 (SING), Bukit Kajang, Corner

SFN 30378 (SING).

Acknowledgements

I thank Dr Saw Leng Guan, (KEP) for his encouragement, advice and

providing photographs of the type and other specimens of Phaeomeria

venusta from Kew and to the Curators of KEP, KLU and SING for

permission to use their herbaria. I thank Dr J. Dransfield (Royal Botanic

Gardens, Kew) for the Latin diagnosis. I also acknowledge with thanks Dr

Saw Leng Guan, Dr Ruth Kiew and unnamed reviewers for advice on this

paper. Thanks are due to the Malaysian Nature Society for help during the

Malaysian Heritage and Scientific Expedition: Belum phase 1 (1993/1994)

and phase 2 (1998). Advice from Dr E. Soepadmo (FRIM) during the

Belum Expeditions is also gratefully acknowledged. Help and

companionship in the field from Ms Rosema Abdullah and Mr Abdullah

Piee are very much appreciated.

References

Burtt, B.L. and R.M. Smith. 1986. Etlingera: The inclusive name for

Achasma, Geanthus and Nicolaia (Zingiberaceae). Notes Royal Botanic

Garden Edinburgh. 43: 235-241.

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

Bulletin, Singapore. 13: 1-249.

Ibrahim, H. 1986. Notes on gingers of Pulau Tioman. Nature Malaysiana.

11(4): 10-13.

238 Gard. Bull. Singapore 53 (2001)

Khaw, S.H. 2000. A checklist of the Gingers (Zingiberaceae) of the Belum

and Temenggor Forest Reserves in Perak, Peninsular Malaysia. Malayan

Nature Journal. 54: 227-232.

Larsen, K. 2000. Selection of Etlingera corneri J. Mood & H. Ibrahim

(Zingiberaceae) as a future correct name. Nordic Journal of Botany. 20:

475-476.

Larsen, K., H. Ibrahim, S.-H. Khaw and L.G. Saw. 1999. Gingers of

Peninsular Malaysia and Singapore. Natural History Publications

(Borneo), Kota Kinabalu, Sabah. Malaysia. 135 pp.

Larsen, K.1970. The genus Nicolaia in Thailand. Natural History Bulletin

Siam Society. 23: 574-575.

Lim, C.K. 2000. Taxonomic notes on Etlingera Giseke (Zingiberaceae) in

Peninsular Malaysia: the “Nicolaia” taxa. Folia malaysiana. 1: 1-12.

Mood, J. & H. Ibrahim. 2000. A new species of Etlingera (Zingiberaceae)

from Peninsular Malaysia and southern Thailand. Nordic Journal of

Botany. 20: 279-283.

Ridley, H.N. 1899. The Scitamineae of the Malay Peninsula. Journal Straits

Branch Royal Asiatic Society. 32: 85-184.

Ridley, H.N. 1924. Zingiberaceae. Flora of the Malay Peninsula. 4: 233-

285. Reeve & Co., London.

Smith, R.M. 1986. New combinations in Etlingera Giseke (Zingiberaceae).

Notes Royal Botanic Garden Edinburgh. 43: 243-254.

Weber, A. 1995. Etlingera gingers (Zingiberaceae) of Peninsular Malaysia.

Nature Malaysiana. 20(3): 80-89.

Postscript: While the account above was in press, the note by Latiff (2001)

appeared discussing the question of priority of E. terengannuensis C.K.

Lim and E. corneri J. Mood & H. Ibrahim. As noted above, not all the

specimens cited by Mood & Ibrahim (2000) belong to a single taxon and

Lim confirms that Corner SFN 32778 and “the Curtis specimen of 1890”

belong to E. fulgens based on the purple undersides of their leaves.

However, as Latiff points out, this does not affect the validity of the name

E. corneri, which is based on the type so the fact that the specimens cited

included more than one taxon is not relevant to the problem of the priority

of names.

Etlingera (Zingiberaceae) in Peninsular Malaysia 239

Detailed examination of the species in question based on living

material has allowed accurate description of the species (see main text)

based only on specimens of E. corneri.

Reference: Latiff, A. 2001. Notice of priority of E. terengganuensis C.K.

Lim. Folia malaysiana. 2(2): 75—68.

Gardens’ Bulletin Singapore 53 (2001) 241-286.

The Limestone Begonias of Sabah, Borneo — Flagship

Species for Conservation

RUTH KIEW

Singapore Botanic Gardens, 1 Cluny Road. Singapore 259569

Abstract

The 18 Begonia (Begoniaceae) species now known from limestone and associated substrates

in Sabah, Malaysia. are listed and a key provided for their identification. Twelve are described

as. new species: Begonia anthonyi Kiew. B. baturongensis Kiew. B. berhamanii Kiew,. B.

diwoliu Kiew. B. heliostrophe Kiew. B. keeana Kiew. B. lambii Kiew. B. layang-layang

Kiew, B. madaiensis Kiew. B. melikopia Kiew, B. punbatuensis Kiew and B. urunensis

Kiew. All these new species belong to sect. Petermannia (Klotzsch.) A-.DC.. except for B.

diwolii that belongs to sect. Diploclinium (Lindl.) A-DC.. Reasons for placing B. amphioxus

Sands in sect. Petermannia and B. gueritziana Gibbs in sect. Platycentrum (Kliotzsch.) A-DC.

are given. Begonia is the most speciose genus on limestone in Sabah. Of the 18 species.

only one, B. gueritziana, is widespread and also found on non-limestone substrates. Of the

others. 12 species are known from single limestone hills and 5 are from hills within the

Same area. Habitat disturbance. which increases the risk of fire, therefore poses the greatest

threat to the conservation of these species.

Introduction

As flagship species for the conservation of the limestone flora of Sabah,

begonias meet all the criteria. Most are readily recognised as relatives of

the begonia house plants, the majority of the limestone species are attractive

(Kiew, 2000) and therefore have not only aesthetic appeal but also have

commercial potential in horticulture as ornamental plants, as well as

exemplifying the need to conserve the limestone flora to ensure its continued

existence.

Begonias are instantly recognised by their leaves, which are usually

very asymmetric. Although the Sabah species have relatively small flowers,

they make up for this deficiency either in their decorative habit or the

great variety of leaves. Some are small rosette plants, like B. gueritziana

and B. lambii; others are tall cane-like plants, like B. keithii Kiew with

bright red, glossy stems: some have leaves coloured magenta underneath.

like B. keeana; or have pink spots, like B. malachosticta Sands, or purple-

magenta hairs, like B. Jambii: others have unusually shaped leaves. such as

the swallow begonia. B. layang-layang, with leaves curved like a swallow’s

wing, or B. amphioxus with bizarre, narrow peltate leaves pointed at both

ends.

242 Gard. Bull. Singapore 53 (2001)

These attractive or unusual-looking species hold potential as

ornamental plants and several have been successfully grown locally in the

Agricultural Park at Tenom under Anthony Lamb’s supervision.

That the limestone flora in Sabah is under threat is not in doubt. The

major threat is from accidental fires that occur in the periodic drought

periods, which are particularly severe in El Nino years. Of the 59 limestone

hills in Sabah, only 22 fall within protected forest or a Virgin Jungle Reserve

(Lim & Kiew, 1997). The other hills are either no longer surrounded by

forest or else lie within commercial forest that has been, is or will be

logged. The vegetation on these hills is particularly vulnerable to fire.

Beaman et al., (1985) showed that in the Great Burn of 1982/83, the

frequency of burning was five times greater in logged-over forest than

undisturbed forest. Indeed, the limestone vegetation that burned then, for

example, on Bukit Dulong Lambu (Gomantong Cave), Bukit Batangan

and Gunung Madai, has not yet recovered.

Other hills are under threat from quarrying as a source of cement,

such as Pulau Balambangan and Bukit Tengar (Segarong Cave), or for

marble, as at the Borneo Marble Quarry on the Segama River, or for road

metal, for example, Batu Pang, Supu and Temambong. Yet others suffer

disturbance from birdnest collecting activities, such as Gunung Madai and

Bukit Dulong Lambu, where the latter has a village built on the summit. A

few have become tourist attractions and are suffering degradation, such as

Batu Tulug (Batu Putih) and Batu Punggul. Even protected status does

not necessarily ensure protection as is demonstrated by the Virgin Jungle

Reserve surrounding Bukit Baturong that has been logged twice (illegally)

in the last ten years.

Conservation of the limestone flora therefore requires a strategy to

protect its biodiversity and Begonia serves as a useful indicator as it is one

of the most biodiverse groups of the limestone flora in Sabah being one of

the ten most speciose families and the most speciose genus (Kiew, 1998a).

In addition, it exhibits extremely high endemism. All its 18 species are

endemic to Sabah. Only one species, B. gueritziana, is widespread; the rest

are narrow endemics. Twelve species are found on single hills and five

species on adjacent hills (Table 1). In addition, their distribution reflects

the general regional differences of the limestone flora as a whole in Sabah

because their distribution is mirrored by the distribution of other species

(Kiew, 1998a). Therefore, by using the distribution of begonia species, a

pattern emerges of those hills that are of greatest importance in order to

protect the maximum biodiversity of the limestone flora. (The only

limestone localities without begonia species are the northern islands of

Balembangan and Banggi). Begonia is therefore an obvious choice as a

conservation icon to illustrate the need for conserving a network of hills in

Limestone Begonias of Sabah, Borneo 243

order to include maximum biodiversity

Table 1. Distribution of Begonia species on limestone in Sabah.

District Locality Begonia Species

(a) species found on single hills

Pensiangan Batu Punggul/Batu Tinahas amphioxus, anthonyi

Pensiangan Sapulut layang-layang

Pensiangan Batu Urun urunensis

Pensiangan Pun Batu punbatuensis

Kinabatangan Bukit Dulung Lambu gomantongensis,

malachosticta

Kinabatangan Melikop melikopia

Lahad Datu Gunung Madai madaiensis

Lahad Datu Bukit Baturong baturongensis,

berhamanii

Semporna Batu Tengar keithii

(b) species found on several hills in the same area

Pensiangan Batu Punggul/Tinahas, Sapulut = lambii

Kinabatangan Kinabatangan Valley (3 hills) ——postarii

Kinabatangan Kinabatangan Valley (2 hills) —heliostrophe

Lahad Datu Segama River (5 hills) diwolii, keeana

Two hills are outstanding for their variety of begonia species. Both

Bukit Dulong Lambu and Batu Punggul are each home to four begonia

species (Kiew, 1998b) as, besides those listed in Table 1, B. guweritziana

also grows there.

While 17 species are narrow endemics, the eighteenth species, B.

gueritziana, is not only widespread but is also the only species that is not

confined to limestone. It grows on a variety of rock types usually by streams

ranging from 30—750 m altitude, the latter on Gunung Kinabalu. However,

although widespread, it has not been recorded from limestone in Sarawak.

The distribution of begonias on limestone in Sarawak shows the same

pattern of high endemism as in Sabah with species being restricted to

244 Gard. Bull. Singapore 53 (2001)

particular areas that also reflects the biodiversity patterns of other limestone

species (Kiew, 1991), namely species are confined to the Bau area, the

Subis area (Niah Cave) or to limestone in the Gunung Mulu National

Park. The fact that only eight begonia species have been described from

limestone in Sarawak, as opposed to 18 for Sabah, may be an indication of

the lack of systematic botanical study than actual lower biodiversity.

Of the 18 species now known from limestone in Sabah, 12 are new

species described below. Apart from B. diwolii in sect. Diploclinium and

the widespread B. gueritziana in sect. Platycentrum, the other 16 species all

fall within sect. Petermannia. Section Petermannia is said by Doorenbos et

al. (1998) to have male flowers typically with two tepals. However, as more

species from Borneo are described, it is likely that this character will prove

not to be diagnostic for the section as the majority of the cane-like begonias

on limestone in fact have male flowers with four tepals.

Section Petermannia is well represented in Borneo and includes

several groups of similar species. One is the group of relatively short

begonias recognised by Sands (1990) that have bristly stems, leaves that

are narrow, obovate and with the midrib in line with the short petiole, the

stipules persistent and becoming papery, and the flowers crowded into

short inflorescences. From Sabah limestone, two species, B. anthonyi and

B. berhamanii, belong to this group. These species do not grow directly on

limestone but are common on the soil around the base of cliffs or on the

steep slopes leading up to the hill, where they frequently form drifts.

Another group includes the cane-like begonias that grow directly on

the limestone. All these species are extremely decorative, particularly those

with an unusual leaf shape (narrow and acutely pointed at both ends), with

daintily scalloped leaf margins often coloured bright red, or with attractive

variegation. In addition, B. keithii has striking red, glossy stems and the

many small carmine heart-shaped male flower buds (Kiew, 1998c). Some

can withstand full sun and live on the exposed summits, such as B.

baturongensis, B. keithii, and B. madaiensis. These species do not flower in

deep shade. Of the other species that grow in light shade beneath the tree

canopy, B. amphioxus and B. layang-layang have narrow leaves with a

pointed base, B. keeana, B. heliostrophe and B. malachosticta, have, in

contrast, broader leaves with a large rounded basal lobe. These cane-like

species have great potential as ornamental plants (Kee, 2000). Indeed,

Sands (1990) reported that B. amphioxus grows well in hanging baskets.

However, the affinity of some species in sect. Petermannia, such as

B. lambii, B. gomantongensis Kiew, B. melikopia, B. postarii Kiew, B.

punbatuensis and B. urunensis, is obscure. There is still a great deal to be

learnt about the begonia flora of Borneo where the undescribed species

outnumber those already named.

Limestone Begonias of Sabah, Borneo 245

la.

1b.

Za.

2b.

3a.

3b.

4a.

Ab.

Sa.

Sb.

6a.

6b.

Ta.

7b.

8a.

8b.

9a.

9b.

Key to Limestone Begonias in Sabah

Small rhizomatous begonias with prostrate stems ...............00:.:0000 2

PE DUS oo 2 11 eee 3

Internodes up to 7.5 cm long, leaves well spaced; lamina margin

ashy, fruit with S cq tal Wines (2.2 ibecesece- scsi io dcen esc ceescenees 5. B. diwolii

Internodes very short, leaves tufted; lamina margin glabrous, fruit

with one wing longer than the other two.................. 7. B. gueritizana

Stem short up to 13 cm tall with few (up to 4) leaves, lamina deeply

ena ie te eh Go BE pss ecte Rebatelts sh aasactydcennesennes 11. B. lambii

Stem more than 25 cm tall with many leaves, lamina plane ........... ~

Mature stem conspicuously bristly, leaf with pinnate venation, basal

lobe scarcely developed; male inflorescence compact, up to 2 cm

ee ee ee ee ee 5

Mature stem glabrous; leaf with palmate-pinnate venation, basal

lobe well developed; male inflorescence a lax cymose panicle....... 7;

Lamina lanceolate, up to 3 cm wide, basal lobe cuneate or slightly

sounded, fruit up to-12 mm wide :..........22:<.0...00.-..0.-- 4. B. berhamanii

Leaf obovate, 3 cm or more wide, basal lobe cordate or auriculate,

seem MADE Uh EE ARRON -T, UADBIAINT NCI 22 ic 56) Lae, hac dasasewcewalocks Ledesclatassecdeedeweds 6

Leaf more than 2.5 times longer than wide, basal lobe cordate up to

4 mm long, fruit wing tip pointed... eee 2. B. anthonyi

Leaf up to 2.5 times longer than wide, basal lobe auriculate up to 10

mm long, fruit wing tip rounded 1.0.0.0... 18. B. urunensis

CARRERA VEE Y MIASTIADOVE Yack sas cadences cecssc tte ncttccens sceensee 16. B. postarii

ee 1a Se 5S ee ee ee ee eee 8

Lamina at least 4 times longer than wide, base narrowed to a point,

DUNG TSE SDE 20 ay ee ee a 9

Lamina less than 4 times longer than wide, base broad and rounded,

DE SP (PE Te 2 eo en eee ee 11

Lamina red-spotted; fruit stalk less than 2.5 mm long, fruit oblong

Co ES I A ae ee 1. B. amphioxus

Lamina plain green or grey mottled; fruit stalk more than 12 mm

long, fruit ovoid or deltoid, 18 x 15 mm or larger .............:.e 10

246

10a.

10b.

tellae

jal los

12a.

12b.

13a.

14a.

14b.

Sa:

15b.

l6a.

16b.

17a.

7b:

Gard. Bull. Singapore 53 (2001)

Stem and petiole crimson; petiole up to 1.5 cm long; lamina not

peltate; fruit ovoid and narrower, up to 25 mm wide .. 10. B. keithii

Stem and petiole pale green; petiole more than 2.5 cm long; lamina

peltate; fruit deltoid and wider, more than 29 mm wide ................4..

eR er Per ee ty aR Bea nsic ee ccocnea couse 12. B. layang-layang

Lamina less than 6 cm wide ::.: ee oes eee 12

Lamina:6 cm or wider. 2202.0... a ek Se ee 14

Lamina more than 3 times longer than wide, pink spotted; tips of

fruit wings pommbted! 35) aece css ee eee 14. B. malachosticta

Lamina less than 3 times longer than wide, not spotted; tips of fruit

WINGS TOUMGEM »..- fcc BLUE Sees taste stone cae. eee eee ee ee 13

Petiole 3—4 cm long; fruit deltoid, shorter than wide, 22—28 mm

WIE so 4 Ee A ce 3. B. baturongensis

Petiole 1—3 cm long; fruit rhomboid (narrowed distally), longer

than wide, 13=20 mmr wide Wa2is0 2: ene eee 13. B. madaiensis

Lamina more than twice as long as wide, apex pointing upwards ...

wa cend beck lic lyecsus tlre te La Oe 8. B. heliostrophe

Lamina less than twice as long as wide, apex pointing downward...

sesoheswnntedsnudiadaseuibelobtesctod te Oat ode SOLON oP Re TD Re ER 15

Lamina obliquely rotund; fruit oblong with pimply surface..............

we ssbtslalch cuebudoduabtOHest acs dati Succ DIgoneane At aaLeN GOR, Seem Ree set 6. B. gomantongensis

Lamina obliquely ovate; fruit deltoid with smooth surface........... 16

Lamina less than 10 cm long; young stems minutely hispid; male

flowers with outer 2 tepals 14—15 mm long ...... 17. B. punbatuensis

Lamina more than 10 cm long; young stems glabrous; male flowers

with outer tepals 7—8' mama! lomgsth ces..c22 cet -cetect est ctens ete eeecenteensecet jl

Lamina up to 16 x 8.5 cm; petioles up to 7 cm long; female flowers

solitary, male flowers with 4 tepals; fruit to 2.5 cm long ..............08

alee basics ae ead este tOaeenae UR Nee REST REEE To ane ee eRe ee Se 9. B. keeana

Lamina 15 x 12 cm or larger; petiole more than 8 cm long; female

flowers in pairs; male flowers with 2 tepals; fruit 4—5.5 cm long ....

vehdduvaaesath Eis DEL Ce EE, RS ee Re eI) CO are

Limestone Begonias of Sabah, Borneo 247

1. Begonia amphioxus Sands

Section Petermannia

Sands, Kew Magazine. 7 (1990) 81 & Plate 149; Kiew, Gardenwise. 15

(2000) 13.

TYPE: Batu Punggul Sands 4045 (not seen).

Distribution: Borneo — SABAH: Pensiangan District, Batu Punggul and

Batu Tinahas.

Habitat: Not common, base of limestone cliffs and in light shade on flat

tops of subpeaks below the canopy.

Notes: Begonia amphioxus is a striking begonia with reddish stems, peltate

leaves narrowing sharply to a point at both ends, beautifully marked with

deep crimson spots. It shows some variation in leaf shape (peltate or not),

number of tepals in the female flower (3 to 5) and in the number of wings

in the fruit (2 or 3). It is also unusual in having joined tepals in the female

flower.

Sands (1990) referred this species to sect. Platycentrum on account

of the specimens he collected having female flowers with 2 locules and

styles and 3—5S tepals, the male having 4 tepals, and the fruit being 2-

winged although he noted that ‘a third wing may occasionally develop in

some fruits ... it tends to be smaller.’

Collecting on Batu Punggul allowed the population at the type site

to be reexamined as well as that on nearby Batu Tinahas. The majority of

fruits on plants at the type site did indeed have fruits with two equal wings

and two locules, but on the same plant, a few fruits with three wings (the

third wing being smaller) could be found. On Batu Tinahas, plants mostly

had fruits with three locules and three equal wings and only a few (on the

same plant) had two locules and two wings. In all other respects the two

populations were the same. The weak development of the third wing

indicates a reduction from the typical state with three equal wings rather

than the character state of sect. Platycentrum where the third wing is well

developed and much larger.

It is certainly an aberrant species for sect. Platycentrum, which, as

Doorenbos et al. (1998) point out, is a section that includes plants that

always have fruits with three wings, one of which is markedly larger than

the other two. They drew attention to this anomalous situation by listing

B. amphioxus under the heading ‘Species not attributable to any existing

section’. As they noted, apart from the single character of the fruit being

two locular, B. amphioxus is typical of sect. Petermannia in its erect habit,

dichasial male inflorescences, solitary female flowers (which may have 4, 3

248 Gard. Bull. Singapore 53 (2001)

or 2 tepals but usually has 5) and the style, which is caducous in the fruit.

On morphological grounds, there is therefore no doubt that B. amphioxus

is a member of sect. Petermannia. In addition, recently Tebbitt (1999)

using molecular data showed that it is affiliated with other species in sect.

Petermannia and Tebbitt and Maciver (1999) demonstrated the presence

of perforate base plates in its endothecial cells, again confirmation that it

belongs to sect. Petermannia.

Begonia amphioxus is easy to grow. In the Agricultural Park at

Tenom, Sabah, it seeds freely and spreads on a lightly shaded limestone

rockery. At Kew, it has been propagated from nodal cuttings, grows on all

types of soil mix but best on a fibrous mix and it flourishes in hanging

baskets (Sands, 1990). It certainly deserves to be more widely known in

cultivation.

Specimens examined: SABAH: Pensiangan District - Batu Punggul Ruth Kiew & S.

Anthonysamy RK4379 (KEP, L, SAN, SING), Batu Tinahas Ruth Kiew & S. Anthony

RK4337 (SAN, SAR, SING).

2. Begonia anthonyi Kiew, sp. nov.

Section Petermannia

A Begonia cauliflora Sands foliis basi cordatis et tepalis integris glabris

differt — TYPUS: Batu Punggul R. Kiew & S. Anthonysamy RK 4352

(holo SAN; iso K, SAR, SING).

Figure 1

Low, erect, unbranched herb, decumbent and rooting at nodes. Stem

ferrugineous towards apex, 18—24 cm tall x 3—4(—6) mm diam., internodes

2—5.5 cm long, becoming woody, nodes not swollen. Stipules glabrous,

narrowly lanceolate, 13—16 x c. 4 mm, margin entire, apex attenuate,

persistent. Leaves alternate, distant, in life upstanding; petiole 2—4(—15)

mm long, slightly grooved above; lamina very dark plain green, papery

when dry, glossy above, obovate, sometimes slightly falcate, (12—)14(—

17) x (3—)4.5(—6.2) cm, slightly asymmetric, narrow side curving inwards

then rounded at base, broad side rounded (2—)3(—4) cm wide and cordate

at base, basal lobes 2—4 mm long, margin dentate and sparsely fringed

with bristly hairs, apex attenuate; midrib not at an angle to petiole, venation

pinnate, lateral veins 6—7 pairs, branching towards margin, sometimes

with | vein in basal lobe, impressed above and slightly prominent beneath.

Plant protogynous. Male inflorescences from upper axils, erect, short

flattened spike, 5—20 mm long of which peduncle is 5—7 mm, longer than

petioles, continuously growing with one or sometimes two flowers open at

Limestone Begonias of Sabah, Borneo 249

ye ee

Figure 1. Begonia anthonyi Kiew

A Habit x '/,, B & C Male inflorescences x 1!/,and x */;, D Male bud x 3/,, E Open male flower

x 1, F Androecium x 4, G Stamen x 8, H Female flower x 1, I Open female flower x 1, J Style

and stigma x 4, K T.S. ovary x 1'/,, L Fruit x */,. (from RK 4438)

250 Gard. Bull. Singapore 53 (2001)

a time; bracts distichous, pale green, persistent. Male flower with pedicel

4—13 mm; tepals 2, totally white and scintillating, glabrous, slightly ovate,

9—12 x 6—7 mm, margin entire, apex rounded; stamens c. 30, in a sessile,

globose cluster c. 4 x 3 mm, filaments c. 1.25 mm long, anthers pale lemon

yellow, ellipsoid, c. 1 mm long, apex emarginate. Female inflorescences

with | or 2 flowers produced from lower leaf axils; bracts 2, reddish brown.

Female flower with pedicel 4—6 mm, sparsely hispid; ovary pinkish with

minute red hairs, narrowly deltoid, 9—16 x 6—16 mm, wings 3, equal,

locules 3, placentas axile, bilamellate with many ovules on both surfaces;

tepals 5, white or rosy red, elliptic, 6—10 mm long, outermost c. 6 mm

wide, innermost c. 4 mm wide, margin entire, apex rounded; styles 3, c. 5

mm long, joined c. half way then bifurcating; stigma papillose forming a

continuous twisted band. Fruit pendent, pedicel stiff, 3—10 mm long, capsule

broadly deltoid, 12—15 x 14—20 mm, locules 3, dehiscing between locules

and wings, wings 3, 4—5 mm wide, thinly fibrous, tips pointed; seeds brown,

minute, broadly ellipsoid, c. 0.25 mm long, base truncate, distally rounded.

Distribution: Borneo: SABAH - Pensiangan District, Batu Punggul, Batu

Tinahas.

Habitat: On rocks or on steep earth slopes up to the base of the limestone

hill, often gregarious.

Notes: Begonia anthonyi belongs to sect. Petermannia in being an erect,

protogynous begonia with the ovary and fruit with three equal wings. Within

sect. Petermannia, it conforms to the group of species recognised by Sands

(1990) when he described B. cauliflora from Sabah that has hispid stems,

short petioles, laminas that are widest at or above the middle and narrow

to the base (rather than having a well developed basal lobe), has a midrib

that is more or less in line with the petiole, and male flowers that are

crowded in short axillary inflorescences. B. berhamanii described below

also belongs to this group. In addition to the characters listed above, species

in this group frequently have an unbranched stem, two tepals in the male

flower, rather few stamens (to about 30), and the fruit stalk is thick and

rigid. This last character contrasts with the other limestone species in this

section where the pedicel is long and at fruit maturity is dry and thread-

like so that the fruit is dangling.

Begonia anthonyi and B. berhamanii grow on soil close to the cliff

base or on the steep slope up to the hill. They tend to grow gregariously

forming carpets but are local and not found away from the limestone hill.

Begonia anthonyi is distinct from B. cauliflora in having a more

asymmetric leaf with the broader side more rounded and it is cordate at

Limestone Begonias of Sabah, Borneo 250

the base (not decurrent) and the lamina is green beneath (not crimson to

brownish as in B. cauliflora), in the male tepals being white and glabrous

(as opposed to pink and hirsute outside), and the female tepals glabrous

and entire (hairy outside and serrate in B. cauliflora). In addition, in B.

anthonyi flowers open one at a time, whereas B. cauliflora is shown as

having several flowers open simultaneously on a single inflorescence.

The species is named for S. Anthonysamy, for many years herbarium

assistant at Universiti Pertanian Malaysia, who accompanied me and helped

with the preparation of plant specimens on many field trips, including the

one to Batu Punggul and Batu Tinahas.

Specimens examined: SABAH: Pensiangan District - Batu Punggul the type and L. Kuntil

SAN 135786 (SAN), Sumbing Jimpim SAN 136091 (SAN); Batu Tinahas R. Kiew & S.

Anthonysamy RK 4438 (SAN, SING).

3. Begonia baturongensis Kiew, sp. nov.

Section Petermannia

A Begonia keithii Kiew petiolis longioribus, foliis latioribus basi rotundatibus

et fructibus latioribus differt — TYPUS: Batu Baturong R. Kiew et al. RK

5026 (holo SAN; iso K, SAR, SING).

Cane-like, glabrous begonia. Stem reddish-brown, glossy, erect, 30—50 cm

tall and 5—10 mm thick, internodes (5—)9.5 cm long, stems becoming

horizontal with many leafy side shoots held horizontally, eventually falling

and becoming pendulous, woody, slightly thicker and distinctly annular at

nodes. Stipules pale green, slightly obovate to lanceolate, 25—28 x 10—11

mm, entire, midrib keeled, apex acute, caducous. Leaves alternate, distant,

pendant; petiole 3—4 cm, slender, slightly grooved above; lamina of young

plants with light green or grey spots, adult unfolding leaves yellow becoming

plain mid-green above and slightly flushed reddish-crimson, sometimes

light green beneath, margin red, slightly succulent in live state, thinly

leathery when dry, slightly glossy, obliquely ovate, asymmetric, (5.5—)7 x

4.25—5.5 cm, narrow side lanceolate, broad side broadly lanceolate 3.25—

4 cm long, base slightly cordate with basal lobe markedly rounded 2.5—4

cm long, margin red, shallowly dentate, apex shortly acuminate; venation

palmate-pinnate, midrib and lateral veins 4, + equal-sized, branched c.

midway to margin, c. 3 in basal lobe, prominent beneath.

Plant protogynous. Male inflorescence axillary in distal leaf axils, glossy

red, erect, cymose panicle to simple cyme in uppermost axil, longer than

petioles, 3—4 cm long of which peduncle is 1.25—2 cm, much branched.

Male flower with pedicel 2.5—5 mm long; tepals 4, outer surface of outer 2

252 Gard. Bull. Singapore 53 (2001)

cerise or deep rosy red, pinkish red inside, glabrous, outer two slightly

cordate and concave, 6—7 x 6—8 mm, margin entire, apex acute, inner

two pale pink, elliptic, 5—6 x 2.5—3.5 mm, apex acute; stamens more than

30, cluster globose, sessile, 2.5—4 x 3—3.5 mm, filaments c. 0.5—0.75 mm,

anthers obovate, c. 0.75 mm long, apex emarginate. Female flowers solitary,

up to 5 each produced at successive axils, pedicel erect, 11—14 mm long,

enclosed by 2 large bracts green tinged red, caducous; ovary green, obovate,

16—20 x 12—17 mm, wings 3, equal, edges tinged red, locules 3, placentas

axile, bilamellate with many ovules on both surfaces; tepals (4—)5, rosy

red, subrotund, slightly convex, isomorphic, 10 mm long, entire, apex

rounded, outer 8—10 mm wide, innermost one c. 6 mm wide; styles 3,

golden yellow, 4—5 mm long, free almost to base, bifurcating; stigma

papillose forming a continuous twisted band. Fruit dangling on fine thread-

like pedicel, 1O—30 mm, capsule broadly ovate, (16—)20(—24) x (22—)

24(—28) mm, glabrous, locules 3, dehiscing between upper half of the

locule and wing, wings 3 isomorphic, thinly leathery, truncate, c. 5 mm

wide, thin and fibrous; seeds brown, minute, broadly ellipsoid, c. 0.3 mm

long, base truncate, rounded distally.

Distribution: Borneo: SABAH — Lahad Datu District, Bukit Baturong.

Habitat: Begonia baturongensis grows on limestone rock below the canopy

in light shade or on exposed rock ledges and crags.

Notes: This species is typical of sect. Petermannia in its cane-like habit and

the ovary and fruits having three equal wings. In addition, the female

flowers produced in the lower leaf axils with the male inflorescences in the

upper ones is a condition found in species of this section.

Like Begonia madaiensis (described below), B. baturongensis has

relatively small leaves, which decrease markedly in size towards the apex,

which are well-spaced on long petioles with their apex pointing downwards

resulting in the twigs having a dainty zigzag appearance, especially on the

lateral horizontal branches. The juvenile leaves of both are variegated, but

become plain green with age. The strikingly variegated juvenile leaves are

illustrated by Kiew (2000). Male inflorescences are only produced after the

female flowers have already become fruits.

In addition, these two species share the same niche growing fully

exposed or in light shade on the summit and shoulders of cliff faces and,

indeed, they do not flower in shade.

The two species have very different fruits: those of B. baturongensis

are broader than long and have truncate wing tips, while those of B.

madaiensis are longer than wide and are narrowed distally. The leaves are

Limestone Begonias of Sabah, Borneo 255

also different, those of B. baturongensis have longer petioles (3—4 cm

long) and the leaf margin is shallowly dentate, while the leaves of B.

madaiensis have short petioles (up to 3 cm long) and the margin is scalloped.

The flowers of B. baturongensis are more decorative being deep rosy red

to cerise compared with the white flowers of B. madaiensis.

Specimens examined: SABAH: Lahad Datu District - Bukit Baturong: the type and S.P.

Lim et al. LSP 737 (SAN, SING).

4. Begonia berhamanii Kiew, sp. nov.

Section Petermannia

A Begonia cauliflora Sands foliis angustioribus et tepalis integris glabris

differt — TYPUS: Batu Punggul R. Kiew & S. Anthonysamy RK 5046

(holo SAN; iso SING).

Figure 2

Low, erect herb, becoming procumbent and rooting at nodes, indumentum

of short hooked hairs, appressed and dense on stem, petiole and lower

surface of veins. Stem reddish-brown, up to 25 cm tall and 2—3 mm diam.,

unbranched, woody, internodes 1.5—2 cm long, nodes swollen. Stipules

lanceolate, 13—14 x 6—7 mm, midrib dorsally strongly keeled with minute

scattered hairs, margin entire with a row of fine hairs, apex narrowly

attenuate, persistent. Leaves alternate, distant, held horizontally; petiole

3—5 mm long, shallowly grooved above; /amina plain dark green and

glabrous above, whitish green (sometimes faintly reddish) beneath, papery

when dry, glossy above, slightly asymmetric, narrowly lanceolate and slightly

falcate, 12.5—16 x 2.5—3.2 cm of which broad side 1.5—1.8 cm wide, basal

lobe cuneate sometimes rounded, 2—3 mm long, margin shallowly dentate,

apex attenuate; venation pinnate, midrib and lateral veins concolorous with

lamina, 7—8 pairs, branching c. midway to margin, 1 vein in basal lobe,

slightly impressed above, slightly prominent beneath.

Plant protogynous. Male inflorescences from upper leaf axils, sessile, to 2

cm long, shorter than petioles, sometimes bifurcating, flowers congested

and covered by distichous, overlapping bracts, only one or two flowers

open at a time; bracts deep rosy or brown red, lanceolate, c. 12 x 3.5—4

mm, margin dentate with fringe of hairs, persistent. Male flower with

minutely hairy, pale pink pedicel 4—11 mm long, tepals 2, opening rosy

red, fading to white faintly tinged pink outside, elliptic, 6—8 x 6—7 mm,

margin entire, apex rounded; stamens 20—30, in hemispherical sessile cluster

c. 3 mm across, filaments 0.5—1.5 mm long, anthers dull white, rotund, c.

0.75 mm long, apex emarginate. Female flowers solitary from lower leaf

254 Gard. Bull. Singapore 53 (2001)

axils; pedicel 2—3 mm long, with densely appressed hairs; ovary white,

deltoid, 8—12 x 11—16 mm, locules 3, outer surface minutely hairy, wings

3, equal, 4—5 mm wide, glabrous, placentas axile, bilamellate with many

ovules on both surfaces; tepals 5, rosy red, broadly elliptic, isomorphic, 5—

10 mm long, apex rounded, outer 4.5—6 mm wide, inner 3—4.5 mm wide,

margin entire; styles 3, 4—6 mm long, divided to base, bifurcating; stigma

papillose forming a continuous twisted band. Fruit with stiff, decurved

pedicel c. 3—4 mm long, the fruit becoming bent backwards and parallel

to stem, capsule broadly deltoid, 11—13 x 10—16 mm, sparsely hispid, 3-

locular, dehiscing between locule and wing, wings 3, isomorphic, 2.5—8

mm wide, tips slightly acute, thin almost papery; seeds brown, minute,

almost globose, c. 0.2 mm long, narrowed to base, rounded distally.

Distribution: Borneo: SABAH -— Lahad Datu, Bukit Baturong.

Habitat: Shaded base of limestone cliff on damp soil.

Notes: Like Begonia anthonyi, this species belongs to the group of species

within sect. Petermannia with hispid stems, short petioles, slightly

asymmetric laminas with a poorly developed basal lobe and with the midrib

almost in line with the petiole. B. berhamanii more resembles B. cauliflora

Sands in its narrowly lanceolate leaves with a slightly rounded basal lobe

and in its dense clusters of male flowers. It is readily distinguished from B.

cauliflora by its narrower leaves with an attenuate apex that are whitish

green beneath (in B. cauliflora the leaves are 4.2—6.4 cm wide, the apex is

acute-acuminate and they are crimson to brownish beneath) and the

glabrous, entire tepals of the male and female flowers (those of B. cauliflora

are hairy outside and those of the female flower are serrate).

The species is named for Berhaman Ahmad who, while he was a

research officer at the Forest Research Centre, Forest Department Sabah,

ably organised several expeditions to explore and collect the limestone

flora.

Specimens examined: SABAH: Lahad Datu District - Bukit Baturong the type and S.P.

Lim et al. LSP 715 (SAN, SING).

5. Begonia diwolii Kiew, sp. nov.

Section Diploclinium

A Begonia speluncae Ridl. caulibus pilosis, internodiis longioribus et foliis

non-peltatis differt — TYPUS: Tempadong, Segama River Ruth Kiew et

al. RK 4767 (holo SAN, iso BRUN, K, L, SAR, SING).

Limestone Begonias of Sabah, Borneo 255

At. wR

Figure 2. Begonia berhamanii Kiew

A Habit x '/,, B Stipules x 2,C Male flowers x 2, D Fruit x 1'/,. (from LSP 715)

256 Gard. Bull. Singapore 53 (2001)

Figure 3

Creeping, prostrate herb attached to substrate at each node by fibrous

roots, old stems setose, indumentum of appressed, concolorous, uniseriate,

eglandular hairs c. 1.5 mm long on the young stems, stipules, petiole, leaf

margin, lower surface of lamina and particularly dense on veins. Stem pale

purplish becoming deep reddish brown with age, up to c. 1 m long and

1.5—2 mm diam. near apex and to 4 mm in old stems, succulent, not

swollen at nodes, internodes 4—12 mm apart at apex lengthening to 5.5—

7.5 cm, stem slightly zigzag with short lateral prostrate branches, c. 10 cm

long produced at wide intervals. Stipules pale reddish purple, broadly

lanceolate, 9—19 x 3.5—8 mm, margin entire, apex distinctly setose, setae

4—5 mm long, persistent. Leaves alternate, distant, lamina positioned flat

against rock surface; petiole brownish or reddish purple, more hispid than

stem, 3—7 cm long up to 14.5 cm on pendent branches, 1—3 mm diam.,

terete; Jamina rich deep plain green above, deep magenta beneath, thinly

fleshy, matt, upper surface glabrous except for a few setose hairs on the

veins, orbicular to reniform, smaller leaves slightly asymmetric, large leaves

very asymmetric, (2—)4(—6.5) cm long and (2.5—)5(—6.5) cm wide of

which the broader side is (1.5—)3(—4.5) cm wide, base slightly cordate

but not overlapping, basal lobes shallowly rounded (0.5—)1.5(—3.5) mm

long, margin slightly wavy and fringed by long hairs, apex rounded in

orbicular leaves, acute in reniform leaves; venation palmate, midrib and

lateral veins concolorous with lamina (occasionally paler), (2—)3, equal

sized, bifurcating twice, first near petiole or more than half way towards

margin, second close to margin, 1—2 veins in basal lobe, slightly impressed

above, slightly prominent beneath.

Inflorescence protandrous, axillary from second node, cymose panicle, pale

magenta, sparsely setose, erect, longer than petioles, 7—15 cm long,

peduncle 6.5—10.5 cm long, each branch with | female flower and several

male flowers eventually with up to 3 branches; bracts paired at base, brown

and papery, narrowly acute with conspicuous midrib, 6—9 x 2.5—3 mm,

margin entire, apex setose, caducous. Male flowers with pale pink pedicel

7—10 mm long; tepals 4, outer 2 with the upper tepal deep rosy pink

outside, both suffused pink inside, glabrous, broadly oval, 6—9 x 5—7 mm,

margin entire, apex rounded to acute, inner 2 almost completely white,

oval, narrower, 7—8 x (2.5—)4 mm, apex bluntly rounded; stamens c. 30,

cluster subglobose with torus c. 1 mm long, filaments c. 0.4 mm long,

anthers lemon yellow, obovate, c. 0.6 x 0.5 mm, apex emarginate. Female

flower with pale pink to magenta pedicel c. 6 mm long; ovary pale pink

with margin of wings tinged green, sparsely pustulate with occasional short

hairs, cordate, c. 3.5 x 3.5 mm, wings 3, equal-sized, c. 1.25 mm wide, base

Limestone Begonias of Sabah, Borneo 257

Figure 3. Begonia diwolii Kiew

A Habit x '/,, B Male flower, C Open male flower, D Androecium. E Stamen. F Female flower,

G Style and stigma, H TS. ovary, I Fruit. (from RK 5073)

258 Gard. Bull. Singapore 53 (2001)

rounded, narrowed to apex, locules 3, placentas axile, bilamellate with

many ovules on both surfaces; tepals 4, c. 4.5—8 x 4—5 mm, margin entire,

outer 2 slightly smaller, tinged magenta with outer surface paler inside,

scintillating, microscopically pustulate, apex rounded; inner 2 slightly longer,

pure white, apex truncate; styles 3, c. 3.5 mm long, joined at the base for c.

1.5 mm, then bifurcating; stigmas green-yellow, with a lunate, papillose

band on each branch. Fruits up to 3 per infructescence, pendent, peduncle,

branches and pedicels becoming thin, dry and hair-like, 12.5—16.5 cm

long, capsule 10—13 x 3.5—5 mm, whole fruit becoming dry and papery,

glabrous, locules 3, dehiscing between locule and wing, wings 3, equal-

sized, broadly rounded, c. 4—5 mm wide; seeds brown, minute, broadly

ellipsoid, c 0.25—0.3 mm long, base narrowed, apex rounded.

Distribution: Borneo — SABAH: Lahad Datu District, Segama River

(Tempadong, Batu Belas, Borneo Marble Quarry and an unnamed hill by

the Lahad Datu road).

Habitat: On vertical and horizontal rock faces from the cliff base in deep

shade to light shade near the summit of limestone hills.

Notes: This new species is most distinctive in its habit of producing long

prostrate, frequently zigzag stems that creep and root over the rock surfaces

(illustrated in Kiew, 2000). The long internodes mean that the leaves are

distant and so contrast with other rhizomatous limestone species with leaves

with palmate venation, such as B. gueritziana in Sabah and B. speluncae

Ridl. in Sarawak, which both have very short internodes so that their

leaves are in a tuft. Furthermore, it cannot be mistaken for B. gueritziana,

which has much larger and thicker leaves, and fruits with one wing much

larger than the other two. In fruit shape (three equal, narrow, rounded

wings), leaf shape (small and frequently orbicular) and the inflorescence

being longer than the petiole, B. diwolii most closely resembles B. speluncae

but it is readily distinguished from the latter, which not only has short

internodes and tufted leaves, but is also glabrous, and the leaves are

frequently peltate. In addition, B. speluncae is placed in sect. Reichenheimia

by Doorenbos et al. (1998), a section that has placentas with one (not two)

lamellae.

This new species belongs to sect. Diploclinium Group | of Doorenbos

et al. (1998), which includes the rhizomatous species with palmately veined,

often symmetric leaves, bisexual inflorescences that are usually protandrous,

male flowers with four tepals and obovate anthers, female flowers with

four tepals, and three-loculate fruits with three equal-sized wings and

bilamellate placentas.

Limestone Begonias of Sabah, Borneo 259

Many of these characters are shared by sect. Petermannia, although

species in that section are less usually rhizomatous or palmately veined.

However, the one character that does exclude B. diwolii from sect.

Petermannia and places it in sect. Diploclinium is its protandry.

The fact that this species is protandrous is not always apparent from

herbarium specimens. Observations in the field show that the inflorescences

produce three branches in sequence, each of which produces one or two

male flowers that open first, followed by a single female flower, then further

male flowers, the entire inflorescence therefore eventually producing a

maximum of three fruits. Unless a plant is observed in the initial male

phase, it appears protogynous with a developing fruit below and young

male flowers above.

This charming species with its dainty round leaves magenta beneath

and bright pale flowers deserves to be introduced into cultivation.

Experimentation in the Singapore Botanic Gardens shows that it grows

well on old coral (Kee, 2000).

It gives me great pleasure to name this decorative species for Diwol

Sundaling, Senior Herbarium Officer at SAN, who organised the expedition

to the Segama River (and many others) and whose enthusiasm and support

in the field is much appreciated.

Specimens examined: SABAH: Lahad Datu District - Tempadong the type and Ruth Kiew

et al. RK5073 Unnamed hill on Lahad Datu road (E, SAN, SING).

6. Begonia gomantongensis Kiew

Section Petermannia

Kiew, Gardens’ Bulletin Singapore 50 (1998) 164.

TYPE: Bukit Dulang Lambu James Awing SAN 47257 (holo SAN).

Distribution: Borneo: SABAH — Kinabatangan District, Bukit Dulong

Lambu (Gomantong Cave).

Habitat: Base of limestone hill in deep shade on boulders or the foot of

cliff faces.

Notes: A member of sect. Petermannia, it is distinct from the other limestone

species in Sabah in its subrotund leaves, which are held horizontally on

long petioles, and its oblong capsules with a pimply surface.

It is known only from Bukit Dulong Lambu.

Specimens examined: SABAH: Kinabatangan District - Bukit Dulong Lambu James Awing

260 Gard. Bull. Singapore 53 (2001)

SAN 47257 (SAN), S.P. Lim & Ubaldus LSP 785 (SAN, SING), R. Kiew & S.P. Lim BDL

3 (SAN, SING).

7. Begonia gueritziana Gibbs

Section Platycentrum

Gibbs, Linn. Soc. Bot. 42 (1914) 82 & figure.

TYPE: Kayoh Hills, Tenom L.S. Gibbs 2892 (holo BM).

As more specimens are available, including ones with female flowers,

additional information is provided here to supplement the original

description:

Rhizome with crowded internodes. Petiole 6—13 cm long; lamina dark

green above, often reddish-purple beneath, S—8 x 5.5—7 cm, basal lobes

scarcely developed 3—18 mm long. /nflorescences protandrous, longer than

petiole, 11—22 cm long of which peduncle is 10—20 cm. Male flower outer

two tepals broadly elliptic, 11—12 x 7.5—8 mm, inner two narrowly elliptic,

c. 11 x 3 mm; stamens joined at base, torus c. 1 mm long. Female flowers

with ovary pale green, glabrous, c. 5—9 mm long, locules 2, placentas

axile, bilamellate, wing 3 unequal, longer wing c. 4—7 mm wide, two shorter

c. 2 mm wide; tepals 4, rosy pink, outer two almost rotund, 5—9 x 5—8

mm, margin entire, apex rounded, inner two elliptic, c. 4—8 x 4—5 mm,

apex cucullate; styles yellow, 2.5—3 mm long, bifurcating; stigma a twisted

continuous papillose band. Capsule c. 10 mm long, longer wing 10—12 mm

wide, broadly rounded, slightly concave, two shorter wings 4—5 mm wide,

thin, apex slightly acute.

Distribution: Borneo — SABAH: Apart from islands and the coastal areas

of the west and north coasts, B. gueritziana is widespread in Sabah both on

and off limestone having been collected from the Tenom, Keningau, Kota

Merudu, Ranau, Labuk Sugut, Kinabatangan and Tawau districts. However,

it has not been collected further west than the Kallang Waterfall in Tenom

(Ruth Kiew & S.P. Lim RK 4280). It has not been reported from Brunei

(Sands, 1996) nor has it been collected from Gunung Api, in the Mulu

National Park, Sarawak, the closest limestone to Sabah. It therefore is a

Sabah endemic.

It is the only widespread begonia species on limestone in Sabah and

is particularly common on all the Kinabatangan limestone hills (even on

the smaller, disturbed ones like Supu and Batu Tulug) and has been

collected from the Kelabangan Ridge, Batu Urun and Lian Cave. On the

other hand, it is absent from limestone in the south (Segama River and

Limestone Begonias of Sabah, Borneo 261

Madai-Baturong limestone) and in the west (Melikop, Pun Batu, Batu

Punggul and Sungai Pangi).

Habitat: On limestone, B. gueritziana grows in light shade and is most

frequent in soil-filled crevices and is particularly common on humus covered

ledges (Kiew, 1998b).

Of the begonia species that grow on limestone in Sabah, it is the only

species that has been collected from non-limestone substrates (basalt,

gabbro and sandstone are cited on herbarium specimens). It has usually

been collected from rocks by the edge of streams from near sea level (30

m) up to 750 m altitude on Gunung Kinabalu.

Notes: Gibbs (1914) originally placed B. gueritziana in sect. Reichenheimia

as do Doorenbos et al. (1998). However, it is atypical of this section in the

ovary having unequal wings, being 2-loculate and each locule having two

placentas. These ovary characters clearly place it within sect. Platycentrum,

with which it is in accordance for all other diagnostic characters, such as

the rhizomatous habit, palmate leaf venation, the axillary, protandrous

inflorescence (illustrated in Kiew, 2000), male flower with 4 tepals, and the

nodding fruit, which dehisces on both sides of the narrower wings.

As in B. diwolii, examination of herbarium specimens can be

misleading, because if the initial stage with only male flowers is not

represented, the inflorescence may appear to be protogynous.

Compared with the begonia flora of Peninsular Malaysia, sect.

Platycentrum is poorly represented in Sabah, the only other species being

B. adenostegia Stapf from Gunung Kinabalu.

Although widespread in terms of geography, substrate and altitude,

it is rather uniform in habit and leaf shape. However, the population on

limestone at Kelabangan is different in having pure white flowers compared

to the usual rosy pink ones.

It is apparently an easy species to grow, as at the Agricultural Park,

Tenom, it seeds freely and has gone wild on shaded rocks in the Orchid

entre.

Specimens examined: The type and from limestone: SABAH: Kinabatangan District —

Kinabatangan Valley, Batu Batangan Ruth Kiew & S.P. Lim RK 4291 (SAN, SING),

Meijer SAN 23103 (locality misnamed Batu Bilit) (SAN); Batu Temambong Besar Ruth

Kiew & S.P. Lim RK 4192 (SAN, SING), Lim et al. LSP 1151 (SAN, SING); Keruak FR

Ruth Kiew & S.P. Lim RK 4183 (SAN, SING); Bukit Dulong Lambu (Gomantong) Meijer

SAN 20754 (K), S.P. Lim et al., SPL 616 (SAN, SING); Supu J. Singh & Eging SAN 51826

(SAN), Batu Tulug (Batu Putih) Dewol & Harun SAN 89912 (SAN, SING), Puasa SAN

10115 (K), S.P. Lim et al. LSP 625 (SAN, SING), LSP 771 (SAN, SING); Pensiangan

District - Batu Urun Ruth Kiew & S. Anthony RK4463 (SAN, SING) - also observed on

262 Gard. Bull. Singapore 53 (2001)

Baladut and Sarupi (Kinabatangan Valley); Kelabangan (Pensiangan District).

8. Begonia heliostrophe Kiew, sp. nov.

Section Petermannia

A Begonia malachosticta Sands foliis majoribus sursum spectantibus differt

— TYPUS: Batu Batangan R. Kiew & Lim S.P. RK 4293 (holo SAN; iso

BRUN, K, KEP, L, SAR, SING).

Figure 4

Bushy, cane-like, glabrous begonia, usually branched at base from a

prostrate rhizome. Stem reddish brown or purple, up to 1.25 m x c. 7.5 mm

diam., becoming woody, flowering at 60 cm, internodes 5—7 cm long,

erect, branching at c. 30 cm, nodes swollen and almost articulate. Stipules

pale green, lanceolate, 10—15 x 4—6 mm, margin entire, apex setose,

caducous. Leaves alternate, distant, positioned with the apex held upwards;

petiole reddish brown or purple, 5—7 cm long, grooved above; lamina

sometimes with a silvery hue when immature, becoming flecked silver and

finally uniformly mid-green at maturity, except for deep crimson patch at

junction with petiole, beneath slightly reddish-green between veins, thinly

succulent in life, thinly chartaceous dried, glossy above, asymmetric,

narrowly ovate (10.5—)14—16(—19) x (3.5—)6—6.5 cm of which broad

side is (2.5—)4.5—5 cm wide, basal lobe rounded, (2—)3.5—5 cm long,

base cordate but not overlapping, margin reddish, serrulate, apex attenuate;

venation palmate-pinnate, midrib and lateral veins 3—S pairs along midrib

and 1 pair at base, bifurcating once about halfway to margin and again

close to the margin, 2—3(—4) veins in basal lobe, slightly raised above,

plane and pale green beneath.

Inflorescences axillary, protogynous, much branched cymose panicle, bright

deep magenta, erect, 7.5—10 cm long of which peduncle is 1.75—3.5 cm

long, longer than petioles, with 1 or 2 female flowers proximally and many

male flowers distally on thin, branches; bracts lanceolate, pale green or

reddish green, c. 7 x 4 mm, margin entire, apex acute, caducous. Male

flower with reddish pedicel c. 5 mm long; tepals 4, white or very pale green,

suffused crimson towards base, inner surface glistening, margin entire, apex

rounded, outer two concave, almost rotund 4—6 x 4.5—6 mm, inner two

broadly elliptic 2—4 x 1.5 mm; stamens c. 60, cluster broadly conical, c. 3

mm long including a short torus c. 0.25 mm long, filaments c. 0.75 mm long,

anthers lemon yellow, broadly oblong, c. 0.5 mm long, apex emarginate.

Female flower pendent in bud then straightening so the open flower is held

horizontally; pedicel 1.75—2 cm long, crimson, ovary glabrous, oblong, c.

Limestone Begonias of Sabah, Borneo 263

Figure 4. Begonia heliostrophe Kiew

A Habit x '/,, B Branchlet of male inflorescence x 2, C Male bud x 2, D Open male flower x 2 '/5, E

Androecium x 5, F Stamen x 10, G 1&2 Female flower x 1, H Open female flower x 1'/,, I

Style and stigma x 5, J T.S. ovary x 1. (from RK 4293)

264 Gard. Bull. Singapore 53 (2001)

15 x 9 mm wide proximally and 14 mm wide distally, wings 3, isomorphic,

reddish when immature becoming green with slight reddish tinge on the

margins at maturity, locules 3, placentas axile, bilamellate, many ovules on

both surfaces; tepals 5(—6), pale yellowish green with a peachy tinge,

elliptic, isomorphic, c. 10 mm long, outermost 6—9 mm wide, innermost c.

4 mm wide, margin entire, apex rounded; styles 3, c. 3 mm long, divided to

base, bifurcating, stigma bright lemon yellow, papillose forming a continuous

twisted band. Fruit dangling on a fine hair-like pedicel, 2—3 cm long;

capsule broadly deltoid, 17—23 x 17—22 mm, glabrous, locules 3, dehiscing

between locule and wing, wings isomorphic, rounded at base, 7—10 mm

wide, tips rounded, thinly fibrous; seeds brown, minute, broadly ellipsoid,

c. 0.25—0.3 mm long, base truncate, rounded distally.

Distribution: SABAH: Kinabatangan District, Kinabatangan Valley (Batu

Batangan and Keruak).

Habitat: On soil at the base of cliffs or in crevices in vertical rock faces

from the base to the crest of cliffs in deep or light shade, particularly

common on soil-covered ledges in light shade.

Notes: In its cane-like habit, its female flowers with 5 tepals, 3 bifurcating

styles that are caducous in the fruit, 3-loculate ovary and fruit with 3 equal

wings and bilamellate placentas, it is typical of sect. Petermannia but like

the majority of limestone begonias, it differs in having 4 tepals in the male

flower instead of the typical 2.

It is unique among the limestone begonias in Sabah in that the leaf

grows with its apex pointing upwards compared with the usual position

where the apex points downward. The species epithet highlights this unusual

feature.

It belongs to a group of cane-like begonias that grow in light shade

on rock faces that have asymmetric, narrowly elliptic leaves with one basal

lobe well-developed and rounded. On limestone, this group includes

Begonia malachosticta, B. keeana and the above species. In addition to the

orientation of the leaf, B. heliostrophe differs from B. malachosticta in its

broader, plain (non-variegated leaves), many-flowered male inflorescences

and rounded (non-arcuate) wings of the fruit, and from B. keeana, besides

the orientation and non-variegation of the leaf, by its larger leaves (up to

12.5 x 8.5 cm in B. keeana) and yellowish green flowers (compared with

the rosy red ones of B. keeana).

Specimens examined: SABAH: Kinabatangan District - Batu Batangan the type; Keruak

Amin & Matin SAN 108091 (SAN), Ruth Kiew & Lim S.P. RK 4158 (E, K, L, SAN, SAR,

SING), RK 4175 (SAN, SING).

Limestone Begonias of Sabah, Borneo 265

9. Begonia keeana Kiew, sp. nov.

Section Petermannia

A Begonia malachosticta Sands foliis latioribus et fructibus latioribus

pedicello multo longiore suffultis — TYPUS: Tempadong R. Kiew et al.

RK 4766 (holo SAN; iso K, SING).

Cane-like begonia, glabrous, root stock woody with several stems. Stem

reddish brown flecked white, smooth and glossy, to 1.25 m tall and 5—7

mm diam., flowering at c. 30 cm tall, internodes 2—8.5 cm, erect or pendent

if growing on edge of cliff face, older stems branching sparingly in the

upper 50 cm, branches at 45 to main stem, woody, slightly swollen at

nodes. Stipules pale green, broadly lanceolate, 14—23 x 7—10 mm, margin

entire, apex setose, persistent. Leaves alternate, distant, lamina pendent

held at 45° on erect petioles; petiole concolorous with stem, in lower leaves

c. 3—7 cm x 3—4 mm long, in upper leaves 1—1.5 cm x c. 2 mm long,

terete; Jamina variegated, margin silvery grey, veins dark green, lamina

between veins either completely silvery grey to the base of the veins or

with a few, large grey-green blotches, beneath uniform deep purple magenta,

fleshy (brittle and snapping in live state), thinly chartaceous in dried state,

glossy above, obliquely narrowly ovate, asymmetric, 11.5—16 x 7—8.5 cm,

the broader side 5—S.5 cm wide, base cordate but not overlapping, basal

lobes rounded, 4—5 cm long, margin slightly scalloped between veins and

minutely serrulate, apex attenuate; venation palmate-pinnate, midrib and

lateral veins 3—4, equal-sized, branching midway to margin, 2 veins in the

basal lobe, slightly indented above, slightly prominent and concolorous

beneath.

Plant protogynous. Male inflorescence produced after and in the same axil

as a fruit or from the upper leaf axils, an erect, cymose panicle with third

order branching, reddish brown or deep purple red, longer than subtending

petiole, 4—15.5 cm long of which peduncle is 2.5—7.5 cm long; bracts pale

green, ovoid, 6—9 x 5—6 mm, margin entire, apex acute, persistent. Male

flowers with pedicel rosy red, 13—18 mm in open flower, very slender,

tepals 4, pale rosy red deeper red towards centre, margin entire, outer two

+ rotund, cucullate, 7—8 x 7—9 mm, glabrous, inner two oblong, apex

rounded, 6—8 x 2—3.5 mm; stamens more than 40, cluster hemispherical,

3(—4) x 3(—4) mm including torus c. 0.5 mm long, filaments c. 0.75—1

mm long, anthers pale yellow, 0.75—1 mm long, apex emarginate. Female

flowers solitary, produced at 1 or 2 (—3—4) successive leaf axils; pedicel c.

15—16 mm long; ovary deep rosy red, oblong, 16—24 x 15—22 mm, wings

3, equal, locules 3, paler, placentas axile, bilamellate with many ovules on

both surfaces; tepals 5, outermost glossy, rosy red (paler than wings), broadly

266 Gard. Bull. Singapore 53 (2001)

oval c. 9—10 x 8—9 mm, innermost isomorphic and slightly smaller c. 7 x 6

mm, sometimes narrowly elliptic c. 8 x 4 mm, margin entire, apex rounded;

styles 3, pale yellow, c. 4 mm long, joined at base for c. 1.5 mm, bifurcating;

stigma papillose forming a continuous twisted band. Fruit dangling, pedicel

drying to fine thread, 23—30 mm long, capsule broadly deltoid, 25—31 x

25—29 mm, glabrous, locules 3, dehiscing on upper half between locules

and wing, wings 3, equal, 13—15 mm wide, very thin and fibrous, tip

rounded; seeds brown, minute, ellipsoid, c. 0.3 mm long, truncate at base,

rounded distally.

Distribution: Borneo — SABAH: Lahad Datu District, Tempadong and

Batu Belas on Segama River and Tabin Wildlife Reserve.

Habitat: At base of cliffs, in crevices in rock face or on exposed shoulders

of cliffs below tree canopy in light shade.

Notes: It is a typical member of sect. Petermannia in its erect cane-like

habit, asymmetric leaves, in being protogynous with solitary female flowers

produced before the distal, many flowered male inflorescences, in having a

3-loculate ovary with the placentation axile, bilamellate placentas, and the

fruit having three equal wings and caducous style.

It most resembles B. malachosticta Sands in its habit and leaf shape

but differs in the pattern of variegation (B. malachosticta has pink spotted

leaves), much wider leaves (they are less than 5 cm wide in B. malachosticta),

much branched male inflorescence (that of B. malachosticta is a simple

cyme with three flowers or is once branched and just 4 cm long with about

six flowers), larger male flowers (3—7.5 mm long in B. malachosticta) and

the broader fruit with rounded wings (fruit is narrower, 1O—17 mm wide,

and has arcuate wings in B. malachosticta).

Local people eat the young, succulent shoots of this new species as a

vegetable. Most begonia leaves have a sourish taste, but in Sabah relatively

few are actively gathered for food. Another such esculent species, Begonia

lazat Kiew & Reza Azmi, was discovered during the course of an

ethnobotanical study (Reza Azmi and Kiew, 1998). The fact that species

known to local people as food plants are still unknown to science indicates

how little is still known about the begonia flora in Sabah.

This is a handsome begonia illustrated in Kee (2000) and Kiew (2000),

which has lush, glossy leaves with silvery variegations on the upper surface

and a deep magenta underside. It grows well on old porous coral in the

nursery at Singapore Botanic Garden (Kee, 2000). That it adopts a

pendulous habit when growing on the edge of cliffs indicates its potential

for being grown in hanging baskets.

Limestone Begonias of Sabah, Borneo 267

This new species is named for Andrea Kee, horticulturalist at

Singapore Botanic Gardens, who was a member of the expedition that

discovered this new species and who has carried out trials on its cultivation.

Specimens examined: SABAH: Lahad Datu District - Segama River — Batu Belas Ruth

Kiew et al. RK 4765 (SAN, SING): Tempadong the type: Tabin Wildlife Reserve Ruth

Kiew RK 5118 (SAN, SING, University of Sabah).

10. Begonia keithii Kiew

Section Petermannia

Kiew, Gardens’ Bulletin Singapore 50 (1998) 189 & Fig. 4.

TYPE: Batu Tengar Cave Kiew, Anthony & Lim RK4327 (holo SING: iso

K, KEP, L, SAN, SAR).

Figure 5

Distribution: Borneo - SABAH: Semporna District, Batu Tengar Cave

(Segarong Cave).

Habitat: In light shade or full sun, growing in crevices on the limestone hill,

dominating exposed shoulders and summit of the hill, where it forms thickets

with its woody cane-like stems, illustrated in Kiew (2000).

Notes: This is one of the most decorative of Sabah begonias with its polished

red stems that give them a lacquered appearance, the dainty pointed leaves

with a crimson scalloped margin and the many carmen, heart-shaped male

buds set on slender sprays. It would be a fine addition to the cultivated

begonias and, considering the harsh conditions under which it grows, would

probably mean it is easier to cultivate than the forest species that grow in

deep shade.

Specimens examined: SABAH: Semporna District - Batu Tengar Cave (Segarong Cave)

Symington & Agama 9312 (K, SING). Keith A9416 (K, SING). Kiew et al. RK 4327 (SAN.

SING).

11. Begonia lambii Kiew, sp. nov.

Section Petermannia

A Begonia inostegia Stapf foliis dense hirsutis obovatis brevipetiolatis et

fructibus oblongis differt — TYPUS: Batu Tinahas R. Kiew & S.

Anthonysamy RK 4405 (holo SAN; iso K, SING).

268 Gard. Bull. Singapore 53 (2001)

Figure 5. Begonia keithii Kiew

A Habit x '/,, B & C Branchlet of male inflorescence x '/, and x 1, D Male flower bud x 1!/,, E

Male flower x 1'/,, F Androecium x 5, G Stamen x 10, H Female flower x 1, 1 Open female

flower x 1'/,, J Style and stigma x 5, K T.S. ovary x 2, L Fruit x 1. (from RK 4327)

Limestone Begonias of Sabah, Borneo 269

Figure 6

Rosette herb with 34 leaves held = flat on the leaf litter layer. Stem and

petioles with brown and scurfy indumentum giving a felted appearance

with additional sparse long hairs. Stem unbranched, becoming woody. erect.

to 10—13 cm tall and 3—5 mm diam.., internodes 1.25—2.5 cm long. Stipules

pale green, lanceolate, c. 14 x 14 mm, margin entire, midrib keeled, apex

cuspidate, caducous. Leaves alternate, distant, held horizontally: petiole

1—2.5 cm, shallowly grooved above: /amina plain dark green above, green

or reddish brown or maroon beneath, hairs on upper surface upstanding,

eglandular, uniseriate, c. 1.5 mm long, translucent in completely green

leaves, reddish brown, purple or magenta where lamina is coloured beneath,

glabrous beneath, thin, matt, broadly obovate, strongly asymmetric, 12—

14.5 cm x 7.5—11cm, narrower side slightly concave, broader side 5—7.5

cm wide rounded, base unequally cordate, basal lobe 1—3 cm long, margin

denticulate with each tooth tipped by long hair, apex rounded or shortly

acuminate (acumen 4—5 mm long); venation palmate, midrib and veins

brown or deep purple and densely hairy beneath, 5—7 veins, midrib +

same size as the laterals, branching dichotomously 3—4 times before

reaching margin, c. 3 veins in larger basal lobe, veins deeply impressed

above giving the lamina a corrugated appearance.

Inflorescence protogynous, from axil of new leaf, densely hairy, erect, longer

than petioles, at first compact and cymose 1—3 cm long with one female

flower, then producing a spike 7.5—10 cm long of distant cymules with

crowded male flowers, peduncle 1.25— 2.5 cm; bracts pale green, narrowly

lanceolate, 5 x 1.5 mm, entire, persistent. Male flower with white pedicel

4—5 mm long, densely hairy; tepals 2, pure white, densely hairy on outer

convex central portion, broadly elliptic, 5—9 x 4.5—8, entire, apex rounded:

stamens c. 30, cluster conical, 2—3.5 x 3—3.5 mm, usually sessile, filaments

0.5—0.75 mm long, anthers obovate, c. 1.5 x 0.75 mm, apex not emarginate.

Female flower: pedicel c. 4—7 mm long, white, densely hairy; ovary green,

sparsely hairy on the wings, densely hairy on the locules, oblong with

wings expanding slightly distally, S—9 x 3—8 mm, wings 3, reddish,

isomorphic, wings c. 1—2 mm wide, locules 3, placentas bilamellate: tepals

5, white, outer surface with long hairs, + isomorphic, innermost slightly

narrower, elliptic, 9—12 x 5—8 mm in outer 4 tepals and 4—5 mm wide in

innermost tepal, entire, apex acute; styles 3, greenish, c. 4 mm long, joined

for c. 1 mm at base, distally bifurcating, stigma papillose forming a

continuous twisted band. Fruit decurved and pendent, pedicel rigid, 1—2

cm long, capsule oblong, rounded at base, truncate or rounded at apex,

15—18(—25) x 10—13(—14) mm. hispid particularly on locules, style

caducous, locules 3, dehiscing between locule and wing, wings 3, equal,

270 Gard. Bull. Singapore 53 (2001)

thin, 2—4 mm wide; seeds brown, minute, broadly ellipsoid, c. 0.2—0.25

mm long, tapered to base, rounded distally.

Distribution: Borneo — SABAH: Pensiangan District, Batu Punggul and

Batu Tinahas; Sapulut.

Habitat: Steep earth slopes up to or on flat areas at the base of limestone

cliffs.

Notes: The species is striking in several respects. Its deeply corrugate,

obovate leaves with a dense covering of magenta or purple hairs contrasting

with the dark green lamina make it a distinctive species. Its habit too,

where the dark green leaves lie on and blend with the leaf litter, is also

unusual. It is illustrated in Kiew (1998d, 2000).

It is a typical member of Section Petermannia in having an upright

habit, being protogynous, the female flowers having 5 sepals, a 3-loculate

ovary with bilamellate placentas, the male flowers having 2 tepals, obovate

anthers that are not emarginate, and the capsule having 3 isomorphic wings

and caducous styles.

In inflorescence structure, it is closely similar to B. inostegia Stapf

from Gunung Kinabalu. Both at first have a short inflorescence, which

produces the single female flower, as this matures into the fruit, an erect

axis is produced with a few distant, short lateral branches that produce

bunches of crowded male flowers. The inflorescence is thus protogynous

and the single female flower is basal to the many male flowers. However,

the leaves of these two species are very different, those of B. inostegia are

long-petioled, ovate with a cuspidate apex, and the capsule is obovoid. In

addition, B. lambii also does not have the peculiar ovate, fimbriate stipules

characteristic of B. inostegia.

This decorative species is named for Anthony Lamb of the

Agricultural Park, Sabah, whose deep interest in Sabah plants has inspired

him to bring this species (and many others of horticultural potential) into

cultivation.

Specimens examined: SABAH: Pensiangan District - Batu Punggul Ali Ibrahim AI 135

(SING), Sumbing Jimpin SAN 135991 (SAN), Batu Tinahas the type and R. Kiew & S.

Anthonysamy RK 4408 (K, SAN, SING); Sapulut (near Kampung Naaturan) R. Kiew & S.

Anthonysamy RK 4345 (SING). It is also cultivated at the Agricultural Park, Tenom.

Limestone Begonias of Sabah, Borneo 27

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it int Lee

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Figure 6. Begonia lambii Kiew

A Habit x '/,,B Male inflorescence x 1,C Male bud x */,,D Open male flower x 1,E Androecium

x 4, F Stamen x 10, G Female flower x 1, H Open female flower x 1,1 Style and stigma x 4, J

TS. ovary x 1!/,, K Fruit x 1. (from RK 4408)

272 Gard. Bull. Singapore 53 (2001)

12. Begonia layang-layang Kiew, sp. nov.

Section Petermannia

A Begonia amphioxo Sands foliis latioribus et fructibus majoribus pedicello

longiore suffultis — TYPUS: Sapulut R. Kiew & S. Anthonysamy RK 4441

(holo SAN; iso BRUN, K, L, SAR, SING)

Cane-like, glabrous begonia. Stem pale green at first, becoming brown, c. 1

m tall and 3—6 mm diam., woody, internodes up to 4 cm long, erect,

branching dichotomously, young branches conspicuously zigzag, nodes

annular. Stipules pale green, narrowly lanceolate, to 12—14 x 2—4 mm

with prominent midrib, margin entire, apex acute, caducous. Leaves

alternate, distant, pendent; petiole pale green, (2.5—)3.5(—S.5) cm and in

dried state 1 mm thick, slightly grooved above; lamina plain dark green

above except for the red patch at point of attachment with petiole, pale

green or sometimes reddish beneath, succulent drying slightly leathery,

slightly glossy above, peltate, slightly asymmetric, narrowly elliptic, (8—)

10(—11.5) x (2—)2.2(—4.5) cm, the broader side (1.7—)2(—2.7) cm wide,

basal lobe acute to pointed sometimes slightly oblique, length of basal lobe

(3.5—)4.2(—5.5) cm, margin red with fine distant teeth becoming closer

and more prominent towards apex, apex attenuated; venation palmate-

pinnate, midrib and lateral veins concolorous with lamina, 1—4 unbranched

pairs, 3 veins in basal lobe, plane above, slightly prominent beneath.

Plant protogynous. Male inflorescences from upper leaf axils, green, erect,

longer than petioles, a slender, lax three times-branched panicle with c. 15

flowers, 3.7—S.2 cm long of which peduncle is 2.2—3.7 cm; bracts lanceolate,

thin, green, lower ones c. 12 x 3 mm, becoming smaller towards stem apex,

midrib prominent, margin entire, caducous. Male flowers with whitish pedicel

(3—)5—7 mm long, tepals 4, outer two pinkish outside, white inside,

glabrous, subrotund, c. 6 x 5.5 mm, entire, apex rounded, inner two white,

narrowly elliptic, c. 5 x 1.75 mm, apex rounded; stamens over 50, cluster

conical, sessile, c. 3 x 2.5 mm, filaments c. 1—1.25 mm, anthers golden

yellow, obovate, c. 0.75—1 mm long, apex emarginate. Female flowers

solitary (rarely 2 per axil) in lower leaf axils; pedicel 17—27 mm long;

ovary deltoid, 17—27 x 19—30 mm, wings 3 equal reddish, c. 4—8 mm

wide, locules 3, placentas axillary, bilamellate with many ovules on both

surfaces; tepals 5, free, greenish white, outer four broadly elliptic, slightly

tapered to base, 9—15 x 5—9 mm, entire, apex rounded, inner one narrower,

slightly obovate, 10—12 x 4—5S mm, apex rounded; styles 3, dark yellow,

4—S5 mm joined for c. 1.75 mm at base, distally bifurcating; stigma papillose

forming a continuous twisted band. Fruit dangling, pedicel extremely fine,

(1.3—)2.8(—3.7) cm, capsule broadly deltoid, distal edge straight, (20—)

Limestone Begonias of Sabah, Borneo 275

24.5(—29) x (22—)26(—33) mm, locules 3, dehiscing between upper half

of wing and locule, wings 3, isomorphic, rounded proximally, truncate

distally, 7(—8) mm wide, thinly fibrous, style caducous; seeds brown, minute,

broadly ellipsoid, c. 0.3 mm long, base narrowed, rounded distally.

Distribution: Borneo — SABAH: Pensiangan District, Sapulut.

Habitat: In shade below tree canopy on rocks at base of cliff and on

subsummits.

Notes: It is a typical member of sect. Petermannia having a cane-like habit,

being protogynous and having fruits with three equal wings and a caducous

style.

In its peltate and relatively narrow leaves, it most resembles B.

amphioxus. However, it is clearly distinct from this species in its non-

variegated leaves (those of B. amphioxus are red spotted), which are broader

and less than four times longer than wide (compared with 8—28 mm wide

and at least five times longer than wide in B. amphioxus), the tepals of the

female flower are always free (frequently joined in B. amphioxus), and the

much larger fruits on long pedicels with three broad equal wings. The

fruits of B. amphioxus are smaller, 9—13 x 5—14 mm, frequently are two-

winged, the wings are only 2—5S mm wide and the pedicels 2—2.5 cm long.

(Sands, pers. comm., informs me that 2—2.5 mm given as pedicel length in

his type description is in error and should read 2—2.5 cm).

Both B. layang-layang and B. amphioxus grow in moderate shade on

rock faces quite close to the base of the cliff. In this they differ from B.

baturongensis, B. keithii and B. madaiensis that grow fully exposed or in

light shade and do not flower in deep shade.

This species takes its name from the local one, /ayang-layang meaning

swallow in Malay, because of the resemblance of the leaf shape to a

swallow’s wings. It is illustrated in Kiew (1998d) as ‘the swallow begonia’.

Specimens examined: SABAH: Pensiangan District - Sapulut (near Kampung Labang) the

type and Diwol Sundaling SAN 13522 (SAN), Aban & Singh SAN 107961 (SAN).

13. Begonia madaiensis Kiew, sp. nov.

Section Petermannia

A Begonia baturongensi Kiew fructibus longioribus quam latioribus apice

acutis — TYPUS: Gunung Madai R. Kiew et al. RK 5057 (holo SAN; iso

K, SAR, SING).

274 Gard. Bull. Singapore 53 (2001)

Figure 7

Cane-like, glabrous begonia with several stems arising from prostrate

rhizome. Stem reddish brown when young, at first erect, then becoming

prostrate or pendent if growing on cliffs, to 1.7 m tall x 4—10 mm diam.,

internodes 3.5—10 cm long, woody, slightly thicker at nodes much branched,

twigs zigzag to 60 cm long, maroon. Stipules pale rosy purple, narrowly

lanceolate, c. 20—21 x 5—6 mm, margin entire, midrib keeled, apex

attenuate, caducous. Leaves alternate, distant, pendent; petiole maroon,

1.2—5 cm x 1—3 mm long, terete; Jamina green, dark green or brown-

green, beneath maroon or pinkish light green, margin red, veins red above

and beneath, young leaves variegated above with oval whitish-green or

silvery spots of varying size between the veins, sometimes coalescing, and

with tiny spots along the margin, veins deep ruby red above and rosy

purple beneath, expanding leaf strongly pleated, succulent and brittle in

live state, membranous when dried, surface slightly scintillating and

appearing finely velvety, obliquely ovate, asymmetrical, (S—)10(—13.5) x

(1.7—)4(—6) cm, the broader side being (1—)2.2(—3.2) cm wide, basal

lobe broadly rounded (1.2—)2.5(—5) cm long, margin scalloped, apex

acuminate; venation palmate-pinnate, midrib and lateral veins red beneath,

(2—)3 veins branching c. midway to the margin with (1—)2 veins in the

basal lobe, plane above, slightly prominent beneath.

Plant protogynous. Male inflorescences about 5 produced from upper axils,

brownish red, erect, slightly longer than petioles, once-branched cyme with

up to 7 flowers, 3.7—4 cm long of which 2—2.7 cm is the peduncle; bracts

narrowly lanceolate, c. 12—14 x 3 mm, caducous. Male flower with pink

pedicel 7—9 mm; tepals 4, outer two white or rosy pink, glabrous, subrotund,

(S—)7(12) x (6—)7(—9) mm, margin entire, apex rounded, inner two pure

white, narrowly obovate, 6—7 x 2 mm, margin entire, apex rounded; stamens

c. 40, cluster broadly ovoid, sessile, c. 4 x 3 mm, filaments 0.5—0.75 mm,

anthers golden or pale yellow, obovate, c. 1 mm long, apex emarginate.

Female flowers solitary from lower leaf axils, up to 5 produced; pedicel 9—

11 mm long, pale red; ovary pale green-white with edge of wings reddish,

oblong, c. 11—12 x 9 mm, wings 3, equal, locules 3, placentas axile,

bilamellate with many ovules on both surfaces; tepals 5, white faintly pink

towards edge, elliptic, margin entire, apex acute, isomorphic, the inner

slightly narrower, outer four c. 9 x 6 mm, inner one c. 9 x 4 mm; styles 3,

yellowish green or golden yellow, 4 mm long joined for c. 0.5 mm at base,

bifurcating; stigmas papillose forming a continuous twisted band. Fruit

pedicel withering and becoming thread-like so the fruit dangles, (6—)14(—

20) mm long, capsule obovate, (17—)25(—29) x (10—)18(—20) mm,

glabrous, locules 3, dehiscing between locule and wing in the upper half of

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capsule, wings 3, equal, rhomboid, (3—)5(—7) mm wide, thinly fibrous;

seeds brown, minute, broadly ellipsoid, c. 0.25—0.3 mm long, base truncate,

distally rounded.

Distribution: Borneo — SABAH: Lahad Datu District, Gunung Madai.

Habitat: Growing directly on limestone rocks and cliff faces in light shade.

Notes: In its erect habit, being protogynous with solitary female flowers

below and male inflorescences above, in the 3-loculate ovary with

bilamellate placentas and capsule with 3 equal wings and a caducous style,

it is typical of sect. Petermannia. However, the male flowers have four

tepals.

Among the cane-like begonias on limestone in Sabah, B. madaiensis

is most similar to B. baturongensis in their leaves being broader and having

a rounded basal lobe compared with those of B. amphioxus and B. keithii,

and in being smaller than those of B. heliostrophe, B. keeana and B.

malachostica. B. madaiensis can easily be told apart from B. baturongensis

by its fruit, which narrows to the apex, giving it a rhomboid outline. Besides

fruit shape, it also differs from B. baturongensis in several other features,

such as its scalloped leaf margin, smaller bracts, less branched male

inflorescence, white or pale pink flowers, tepals of the male flower that are

longer than broad, and the leaves, which are strongly pleated before they

expand.

The leaves of young plants are larger (about twice the size of those

on fertile shoots) and can be conspicuously variegated, but this coloration

is not expressed in older leaves. Plants growing in deeper shade tend to

have greener, less maroon leaves and plants in deep shade, 1.e. at the base

of cliffs, do not flower. Flowering on a single twig is effectively unisexual

as the female phase does not overlap with the later male phase.

Specimens examined: SABAH: Lahad Datu District - Gunung Madai the type and W.

Meijer SAN 37950 (SAN), SAN 37959 (SAN), S.P. Lim et al. LSP 672 (SAN, SING), LSP

702 (SAN, SING).

14. Begonia malachosticta Sands

Section Petermannia

Sands, Kew Magazine. 7 (1990) 64 & Plate 145.

TYPE: Bukit Dulong Lambu Sands & Young - Sands 3933 (not seen)

Distribution: Borneo — SABAH: Kinabatangan District, Bukit Dulong

Lambu (Gomantong Cave).

Limestone Begonias of Sabah, Borneo OAT

Habitat: In rocky crevices in the sheer cliffs where the tree canopy begins

to open up, as well as on exposed jagged outcrops. It is no longer common

on the summit, which is greatly disturbed by the village of birdnest collectors

that is established on the summit plateau.

Notes: This is a strikingly beautiful species with pink spotted leaves with a

red underside. It has been introduced into cultivation at Kew where it is

grown in a 50:50 mix of peat and sand. It is propagated easily from nodal

cuttings (Sands, 1990).

It is eaten as a vegetable but is reportedly very sour (Kamideh SAN

66578).

Specimens examined: SABAH: Kinabatangan District - Bukit Dulang Lambu (Gomantong

Cave) James Ah Wing SAN 38108 (SAN), SAN 47263 (SAN); Kamideh Bangillan SAN

66578 (SAN).

15. Begonia melikopia Kiew, sp. nov.

Section Petermannia

A Begonia erythrogyna Sands foliis latioribus, inflorescentiis minoribus,

tepalis rubris et fructibus angustioribus differt — TYPUS: Gua Melikop

Ruth Kiew et al. RK 5011 (holo SAN; iso BRUN, K, L, SAR, SING).

Cane-like, glabrous begonia with prostrate rhizome with several spaced

erect stems to 1.5 m tall x 5—7 mm diam., base becoming woody, young

stems brittle at swollen nodes, reddish above nodes, otherwise green, leaf

scars very conspicuous, internodes 2.5—5.5 cm long, slightly ribbed, stems

with 1—2 branches. Stipules green, strongly keeled, lanceolate, 23—25 x

9—11 mm, margin entire, apex acute, caducous. Leaves alternate, distant,

in a bunch at top of stem, held horizontally; petiole completely reddish or

reddish proximally and distally but green in between, 8.5—13 cm x 4—6

mm, grooved above; /amina plain mid-green above, beneath completely

magenta or with magenta blotches and green veins, seedling leaves with

white spots, succulent and brittle in life, thinly chartaceous in dried state,

glossy above, asymmetric, broadly ovate, sometimes almost reniform, 15—

19.5 x 12—18.5 cm of which broader side is 7.5—11.5 cm wide, basal lobes

broadly rounded, 3.5—7.5 cm long, base cordate but not overlapping, margin

scalloped between veins and minutely and distantly dentate, apex cuspidate;

venation palmate-pinnate, midrib and lateral veins concolorous with lamina

above, 5—7 pairs, branching twice dichotomously, first less than halfway

to margin, then c. halfway to margin, 1—2 veins in basal lobe, impressed

above, prominent beneath.

278 Gard. Bull. Singapore 53 (2001)

Inflorescence axillary, protogynous with 2 female flowers at base of the

erect cymosely branched panicle of male flowers, shorter than petioles,

6—7.5 cm long of which peduncle is 1.7—2.5 cm with diam. c. 2 mm,

branching c. 3—4 times, c. 4—8 main branches 4—15 mm long, ultimate

branches 8—20 mm long, knobbly from congested scars of fallen flowers,

each terminating in 1 bud or 1 bud and 1 one flower. Male flower with

slender pedicel 7—12 mm long, tepals 2, flame-coloured darker towards

base, glabrous, oval, 8—11 x 4—5 mm, margin entire, apex rounded; stamens

c. 25, cluster conical, 3.5—4 x 2.5—3 mm, sessile, filaments c. 0.75 mm long,

anthers pale yellow, spathulate, c. 1.2 mm long, apex not emarginate. Female

flower not known. Fruit pendent and dangling on fine and thread-like

pedicel 25—38 mm long, capsule narrowly deltoid narrowed into pedicel,

40—55 x 10—27 mm, glabrous, locules 3, placentas axile, bilamellate with

many seeds on both surfaces, dehiscing between wing and locule, wings 3,

equal, 6—7 mm wide, becoming thin and papery, tip acute or sometimes

rounded; seeds brown, minute, almost globose, c. 0.2 mm long, base tapered,

rounded distally.

Distribution: Borneo: SABAH -— Kinabatangan District, Gua Melikop.

Habitat: Gua Melikop is an exposed cliff face on a hillside at about 400 m

altitude. Its summit is covered by a deep layer of peat, which does not

support a limestone flora.

Begonia melikopia grows rooted in cracks in the sheer limestone

rock face in deep shade below the canopy but c. 1.5—2 m above base of

limestone cliff. (Some seedlings were found growing in soil near the base

of the cliff but no adult plants were found on soil, suggesting that these

seedlings might not survive to maturity).

Notes: In its large, broadly ovate leaves, paniculate male inflorescence and

large deltoid fruits pendent on long pedicels, it resembles B. erythrogyna

Sands. However, it is distinct from this species in its much broader leaves

(those of B.erythrogyna are less than 16 cm wide), its shorter, axillary male

inflorescence (terminal and 7—14 cm long in B. erythrogyna), the male

flowers are flame-coloured (they are white to pale green in B. erythrogyna),

and the fruit, which is more than twice as long as wide (in B. erythrogyna it

is less than twice as long as wide).

Begonia melikopia is typical of sect. Petermannia in its cane-like

habit, in being protogynous, having male flowers with two tepals, and the

fruits with three equal wings and three locules each with bilamellate

placentas.

Limestone Begonias of Sabah, Borneo 279

Specimens examined: Known only from the type population.

16. Begonia postarii Kiew

Section Petermannia

Kiew, Gardens’ Bulletin Singapore. 50 (1998) 165 & Fig. 1.

TYPE: Bukit Panggi Kiew & Lim RK 4221 (holo SAN; iso SING).

Figure 8

Distribution: Borneo — SABAH: Kinabatangan District, Kinabatangan

Valley (Bukit Panggi, Bukit Dulong Lambu (Gomantong Cave) and Subuk

Estate).

Habitat: At base of limestone hills (but not on cliff faces) growing in deep

shade in sheltered damp habitats on soil, low limestone boulders or the

base of tree trunks.

Notes: This softly hairy begonia with hairy tepals in the male flower and

the hairy fruit is quite unlike any other limestone species in Sabah.

Specimens examined: SABAH: Kinabatangan District - Bukit Dulong Lambu Joseph B. et

al. SAN 122763 (SAN), Lim S.P. & Ubaldus LSP 802 (SAN, SING), Ruth Kiew & Lim S.P.

BDL 4 (K, SAN, SAR, SING); Bk. Panggi Ruth Kiew & Lim S.P. RK 4221 (SAN, SING):

Subak Estate (Lower Kinabatangan River) J. Dransfield et al. JD 5770 (SAN).

17. Begonia punbatuensis Kiew, sp. nov.

Section Petermannia

A Begonia burbidgei Stapf fructibus minoribus, pedicellis filiformibus 2.5—

3 cm longis differt — TYPUS: Pun Batu R. Kiew & A. Berhaman RK 4260

(holo SAN; iso BRUN, K, L, SAR, SING).

Figure 9

Cane-like begonia. Stem robust, dark brown, up to 1.3 m tall and 6 mm

diam., erect, little branched, woody, internodes up to 4—S.5 cm long, nodes

conspicuously swollen with conspicuous leaf scars, eglandular uniseriate

hairs c. 0.5—0.75 mm long, scattered on the young stem, becoming

glabrescent. Stipules pale green, glabrous except for a few scattered hairs

on the outer surface of the midrib, slightly obovate, 2—3.5 x 1—1.3 cm,

margin entire, apex acute, persistent. Leaves alternate, distant; petiole green

with brown bristly hairs, 2.5—6 cm long, grooved above; /amina plain mid-

green above and beneath except for crimson patch on upper surface at the

junction with the petiole, succulent and brittle in life, thinly leathery in

280 Gard. Bull. Singapore 53 (2001)

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A Habit x '/;, B Branchlet of male inflorescence x 1'/,, C Male bud x 1, D Male flower x 2, E

Androecium x 4'/,, F Stamen x 8, G Fruit x 1. (from RK 3838)

Limestone Begonias of Sabah, Borneo 281

dried state, glossy above, obliquely ovate, strongly asymmetric, 5—9 x 6—

10 cm, narrow side lanceolate, broad side broadly ovate 4.5—7 cm wide,

base cordate, not overlapping, basal lobe 1.7—4 cm long, margin shallowly

dentate, faintly red in young leaves, apex acute; venation palmate, midrib

and lateral veins concolorous with lamina, sparsely hairy beneath, 3—4, +

equal-sized, branching dichotomously twice before reaching the margin,

first close to petiole, second towards the margin, | vein in basal lobe, plane

above, prominent beneath.

Inflorescences protogynous, in the axils of upper leaves, pale green, glabrous,

+ erect, longer than adjacent petiole, cymose, with up to 4 inflorescences

at successive leaf axils, each branched at base, one axis with 2 female

flowers on peduncle 2.5—3 cm long, the other axis bears the male

inflorescence with c. 4 lateral branches, conspicuously jointed, 5.5—9 cm

long of which peduncle 3.2—4.5 cm long; bracts pale green, obovate, those

at base c. 20 x 15 mm, those subtending male flowers c. 10 x 9 mm, persistent.

Male flowers with whitish green, glabrous pedicel 17 mm long; tepals 4,

rosy or pale pink, glabrous, outer two broadly oval, 14 x 13 or 15 x 10 mm,

inner two narrowly elliptic, c. 9 x 5 mm, margin entire, apex rounded;

stamens c. 50, cluster conical, sessile, 9 x 4 mm, anthers subsessile, golden

yellow, narrowly oblong, c. 2 x 1 mm, apex emarginate. Female flower with

pale green, glabrous pedicel; ovary pale greenish white, glabrous, ovoid, 15

x 15—17 mm, wings 3, isomorphic, 3 locules, placentas axile and bilamellate

with many ovules on both surfaces; tepals 5, pale rosy pink, broadly elliptic,

outer 17 x 10 mm, inner 14 x 7 mm margin entire, apex slightly acute; styles

3, golden yellow, c. 5—6 mm long, bifurcating; stigma papillose forming a

continuous twisted band. Fruits pendent, dangling on fine, thread-like

pedicel 2.5—3 cm long, capsule 17—22 x (12—)20—28 mm, glabrous,

locules 3, dehiscing between upper half of locule and wing, wings 3,

isomorphic, truncate distally (4—)6—7 mm wide, thinly fibrous; seed brown,

minute, ellipsoid, c. 0.5 mm long, base truncate, distally rounded.

Distribution: Borneo — SABAH: Keningau District, Pun Batu.

Habitat: Summit of the limestone hill growing on exposed peat-covered

rocks.

Notes: Pun Batu is an isolated tower karst hill and its summit is covered in

a deep peat layer. Its unique begonia grows in this habitat and as yet has

not been found on any other limestone hill. The surrounding forests are

actively being logged but when we visited in 1996, its surrounding foothills

were still pristine.

In its upright habit, in being protogynous, and with the fruit being 3-

282 Gard. Bull. Singapore 53 (2001)

Figure 9. Begonia punbatuensis Kiew

A Habit x '/,, B Branchlet male inflorescence x 1, C Male bud x 1'/,, D Male flower x 1, E

Androecium x 3, F Stamen x 10, G Fruit x 1. (from RK 3898)

Limestone Begonias of Sabah, Borneo 283

loculate with three equal wings and a bilamellate placenta and caducous

style, it falls within sect. Petermannia.

Begonia punbatuensis resembles B. burbidgei from Gunung Kinabalu

in its robust, woody stems with swollen nodes, large stipules on the young

stems and in its leaf shape and venation. However, B. burbidgei is a larger

plant growing to c. 2—2.7 m tall, its leaf apex is acuminate to cuspidate, its

inflorescences have larger bracts, the male flowers have much larger tepals

(the outer two tepals measuring c. 25 x 20 mm), and the fruits are also

much larger (up to 27 x 46 mm), the wings are not truncate and the fruit

stalk is stiff and short (c. 15 mm long).

Although its bracts are quite conspicuous, B. punbatuensis does not

belong to sect. Bracteibegonia sensu Doorenbos et al. (1998), which includes

just three species from Java and Sumatra. These differ from species in sect.

Petermannia in having pinnate venation, few flowered bisexual

inflorescences and persistent styles in the fruit. Doorenbos et al. therefore

transferred the Bornean species with conspicuous bracts, previously placed

in sect. Bracteibegonia (such as B. burbidgei Stapf and B. imbricata Sands),

to sect. Petermannia, to which they conform in their venation, inflorescence

structure and caducous style.

Specimens examined: SABAH: Pensiangan District — Pun Batu the type and Ruth Kiew &

A. Berhaman RK 4279 (E, KEP, SAN, SING).

18. Begonia urunensis Kiew, sp. nov.

Section Petermannia

A Begonia anthonyi Kiew and B. berhamanii Kiew caulibus ramosis, foliis

latioribus et inflorescentiis masculis paniculatis differt — TYPUS: Batu

Urun R. Kiew & S. Anthonysamy RK 4473 (SAN, SING).

Low, erect, bushy herb, trichomes uniseriate c. 0.5 mm long, dense on

young stems, stipules, petioles, lower surface of veins, inflorescence and

pedicels, sparse on lamina. Stem to 33 cm tall and 5—7 mm diam., woody

towards the base, lowermost internodes 4.5—10 cm long, lower nodes

swollen, upper internodes 3—4.5 cm long, producing short branches which

sometimes branch again. Stipules lanceolate, 5.5—17 x 2.5—3.5 mm, margin

entire, apex attenuate to setose, caducous. Leaves alternate, distant, towards

apex some leaves subopposite; petiole 6—15 mm long in lower leaves, 4—7

mm in the upper; /amina plain mid-green above, (12.5—)15—(15.5) x (S—)

6.5(—8) cm, slightly succulent in life, papery when dry, obovate, sometimes

rhomboid and widest midway, slightly asymmetric, narrow side concave

towards the base, broad side rhomboid, 3—4.2 cm wide, base unequally

284 Gard. Bull. Singapore 53 (2001)

cordate, basal lobe 2.5—7.5 mm long and auriculate, margin minutely and

distantly dentate, apex narrowly attenuate; venation pinnate, lateral veins

6—7 pairs, branching once, sometimes twice before reaching the margin, 1

vein in basal lobe, prominent above and beneath.

Plant protogynous with 2 female flowers and a male inflorescence from

each of upper leaf axils. Male inflorescence erect, 0.8—2.5 cm long of

which peduncle is to 1 cm long longer than petioles, central axis with

several tiers of three times branched side branches c. 7 mm long; bracts

lanceolate, c. 7 x 2 mm, margin entire, apex setose, caducous. Male flowers

with pedicel 3—5 mm long; tepals 2, pinkish, densely hispid outside, elliptic,

c. 5 x 3.5 mm, margin entire, apex rounded; stamens c. 40—SO in conical,

sessile cluster; filaments c. 0.5 mm long; anthers orange, spathulate, c. 0.75

mm long, apex not emarginate. Female flower not known. Fruit with

decurved, stiff pedicel (4—)7(—9) mm long; capsule broadly oblong, (14—)

18(—21) x (15—)17(—19) mm, glabrous, locules 3, placentas axile,

bilamellate with many seeds on both surfaces, dehiscing along entire length

between locule and wing, wings 3, isomorphic, tip rounded, (4—)6(—7)

mm wide, thinly fibrous; seed brown, minute, broadly ellipsoid, c. 0.2 mm

long, base truncate distally rounded.

Distribution: Borneo — SABAH: Pensiangan District, Batu Urun.

Habitat: Deeply shaded forest on soil on steep slopes up to and at the base

of limestone.

Notes: Its erect habit, three locular ovary and fruit with three equal wings

place this species within sect. Petermannia. Among the limestone species,

it groups with B. anthonyi and B. berhamanii, which are short begonias

with bristly stems, their leaves are only slightly asymmetric, the petioles

short and the midrib more or less in line with the petiole, the male flowers

have two tepals, and the fruits do not have long, thin, thread-like pedicels.

However, it is not typical of this group of begonias in two respects. Firstly,

it differs in habit. In B. urunensis, the stem branches once or sometimes

twice to produce flat sprays of leaves giving it a bushy appearance. (The

stems of B. anthonyi and B. berhamanii are unbranched). Secondly, the

male inflorescences are produced from the same leaf axil as the female

flowers and are erect and paniculate, whereas in B. anthonyi and B.

berhamanii, the female flowers are produced from the lower axils and the

short, spicate male inflorescence from the upper. B. urunensis can also be

told apart from these two species by its broader leaf, which is less than 2.5

times longer than broad as compared with 2.7—4 and 5 times, respectively

for B. anthonyi and B. berhamanit.

Limestone Begonias of Sabah, Borneo 285

Batu Urun is one of the most remarkable limestone formations in

Sabah being unique in its cave, which has its mouth on the side of a deep

and wide basin below soil level. From the mouth of the cave a dangerously

steep scree slopes precipitously down to a subterranean river, which to

judge from the large tree trunks lodged in it carries strong currents in rainy

weather. On one side of the basin is a vertical wet cliff face that is covered

in an undescribed species of Monophyllaea (Gesneriaceae), one of two

Monophyllaea species to be found on limestone in Sabah (Kiew,1998a).

Above soil level surrounding this basin is a wide area where the limestone

projects just above the soil surface, forming an uneven and pitted rock

pavement, or it outcrops as low cliffs up to 4—5m high or as boulders of

various sizes, all in deep shade below the canopy. It is on soil over limestone

that this new begonia species is found.

Specimens examined: SABAH: Pensiangan District — Batu Urun, the type and Ruth Kiew &

S. Anthonysamy RK 4472 (K, SAN, SING).

Acknowledgements

This paper forms part of the results from ‘The Biodiversity and Conservation

of the Limestone Flora of Sabah’ project carried out in collaboration with

the Forest Research Centre (FRC), Sabah Forest Department. The author

thanks the Singapore Botanic Gardens and the Ministry of Science,

Technology and Environment, Malaysia (under IRPA Grant No. 52858)

for sponsorship; the staff at FRC, Robert C. Ong, K.M. Wong, Diwol

Sundaling, A. Berhaman and S.P. Lim, for their logistic help and enthusiastic

support; Anthony Lamb, Herbert Lim and Andy Surat of Agricultural

Park, Tenom, for showing us localities in the Pensiangan District; J.H. Tan

with whose support Melikop was located; Mdm P.H. Yap and Jaap J.

Vermeulen for their excellent botanical illustrations; Andrea Kee for

carrying out horticultural trials; to the Curators of the BM, E and SAN

herbaria for permission to examine specimens in their care; to Mark J.E.

Coode for comments on the manuscript and for correcting the Latin and

to Martin J.S. Sands for his detailed and helpful comments.

References

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

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Kee, A. 2000. Collecting begonias from the wild. Gardenwise. 15: 14—15.

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Conservation in Malaysia. Malayan Nature Society, Malaysia. pp. 42—

50.

Kiew, R. 1998a. Towards a limestone flora of Sabah. Paper presented at

B.C. Stone Memorial Symposium, Kuala Lumpur. Malaysia.

Kiew, R. 1998b. Niche partitioning in limestone begonias in Sabah, Borneo,

including two new species. Gardens’ Bulletin Singapore. 50: 161—169.

Kiew, R. 1998c. The unique elements of the limestone flora of Batu Tengar

Cave (Segarong), Sabah, Malaysia. Gardens’ Bulletin Singapore. 50: 185—

196.

Kiew, R. 1998d. Wanderings in the great forests of Borneo. Gardenwise.

11: 8—11.

Kiew, R. 2000. Decorative limestone begonias from Sabah. Gardenwise.

15: 12—13, 15.

Lim, S.P. & R. Kiew. 1997. Gazetteer of limestone localities in Sabah,

Borneo. Gardens’ Bulletin Singapore. 49: 111—118.

Reza Azmi and R. Kiew. 1998. Begonia lazat (Begoniaceae), a new culinary

begonia from Borneo. Gardens’ Bulletin Singapore. 50: 43—48.

Sands, M.J.S. 1990. Six new begonias from Sabah. Kew Magazine. 7: 57—

85.

Sands, M.J.S. 1996. Begoniaceae. In: M.J.E. Coode, J. Dransfield, L.L.

Forman, D.W. Kirkup and Idris M. Said (eds.) A Checklist of the

Flowering Plants and Gymnosperms of Brunei Darussalam. Forestry

Department, Brunei & Royal Botanic Gardens, Kew. pp. 38—40.

Tebbit, M.C. 1999. A revised classification of selected Asian begonias based

on evidence from morphology and molecules. In: S. Andrews, A.C. Leslie

and C. Alexander (eds.), Taxonomy of Cultivated Plants. Royal Botanic

Gardens, Kew. pp. 401—402.

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203—221.

Gardens’ Bulletin Singapore 53 (2001) 287-289.

Begonia conipila Irmsch. ex Kiew (Begoniaceae) from

the Gunung Mulu National Park, Sarawak, Malaysia

RUTH KIEW

Singapore Botanic Gardens, | Cluny Road, Singapore 259569

Abstract

Irmscher’s name, Begonia conipila, is validated for the striking species that grows on the

Melinau limestone in Sarawak, Borneo.

Introduction

The scientific name of this most distinctive species was in fact never validly

published although, following Irmscher’s annotated specimens, it has

become the accepted name and specimens of it are correctly identified in

herbaria and the name has appeared in print (Anderson & Chai, 1982;

Jangowe, 1999). It is timely therefore to validate the name of his species,

particularly because it is seen by most visitors who climb Gunung Api in

the Gunung Mulu National Park.

Low’s specimen is selected as the lectotype as it is the earliest

collection and was annotated as a new species by Irmscher.

Begonia conipila Irmsch. ex Kiew

Section Petermannia

A Begonia havilandii Ridl. foliis minoribus, tepalis masculis 2 et fructibus

oblongis differt - LECTOTYPUS: ‘Mooloo’ (Gunung Mulu National Park)

Low s.n. (K).

Creeping herb or low root climber, roots fibrous produced at nodes in

contact with substrate, indumentum of stiff, white, uniseriate, eglandular

trichomes, c. 2—5S mm long, each on a raised conical base, dense on young

stem, petiole, on stipule margin and outer surface of stipule midrib, on the

lamina dense on the upper surface, margin and lower surface of veins,

some on lower lamina surface. Leafy stem 15—25 cm long and 2—3(—4)

mm diam., little branched, internodes 2.5—S cm long. Stipules lanceolate

with a distinct midrib, 7—12 x 4—7 mm, margin entire, apex setose,

persistent. Leaves alternate, distant, petiole 1—2 cm in upper leaves

elongating to 2—S5 cm in lower leaves, /amina somewhat bullate, variegated,

288 Gard. Bull. Singapore 53 (2001)

upper surface dark green to blackish purple, paler between veins, the lower

surface reddish, rarely light green, in dried state thin and papery, conical

hair bases giving a pustulate appearance above, asymmetric, broadly ovate

to reniform, 4—7.5 x 3.5—8.5 cm, broader side (2.25—)3.5—5S cm wide,

base rounded to cordate, lobes slightly overlapping, basal lobes rounded,

(0.75—)2—3.5 cm long, margin irregularly serrate, each tooth setose, apex

cuspidate; venation palmate, main veins 3, equal-sized, branching c. halfway

to margin, with another 1—2 veins in basal lobes, impressed above and

prominent beneath.

Inflorescence axillary, whitish or light green, hispid, erect, longer than

petioles, racemose, 3.5—9.5 cm long of which peduncle is 1.5—6 cm,

protogynous with a single basal female flower and many male flowers

distally on 2 short lateral branches c. 3—11 mm long and crowded in a

terminal cluster; bracts similar to stipules, c. 5—6 x 3 mm, persistent;

bracteoles light green or whitish, broadly ovate, 0.75 x 0.25 mm, apex

setose, persistent. Flowers with white, translucent tepals. Male flower with

whitish or light green pedicel, c. 1—3 mm long; tepals 2, sparsely hispid on

upper half of outer surface, broadly elliptic, c. 5 x 3.5—4.5 mm, margin

entire, apex rounded; stamens yellow, c. 10—11, in hemispherical cluster

joined in a torus c. 0.5—0.75 mm, filaments between 0.25 mm on lower

stamens to 1 mm long in the distal ones, anthers bright yellow, obovate, c.

0.75—1 mm long, emarginate, opening by pores. Female flower with pedicel

c. 3—5S mm long, light green, finely hispid; ovary narrowly oblong, c. 7.5—

11 x 3—5S mm, wings 3, subequal, two shorter 1.5 mm and broader one 2

mm wide, locules 3, placentas axile, bilamellate with many ovules on both

surfaces; tepals 5, isomorphic, narrowly lanceolate, c. 7—7.5 x 3—4 mm,

margin entire, apex acute, sparsely hispid on outer surface; styles 3, free to

base, 5 mm long, bifurcating; stigma greenish, papillose forming a continuous

twisted band. Fruit pedicel stiff, decurved, (2—)4—6(—8) mm long, capsule

oblong, c. (11—)18—20 x 10—15 mm, sparsely hispid, locules 3 but one

scarcely developed, dehiscing between wing and locule, wings thin, narrowed

to base, truncate distally, two narrower wings, c. 3—S mm wide, broader

wing c. 5—7 mm wide, style not persistent.

Distribution: Borneo - SARAWAK: endemic to the Melinau limestone in

the Gunung Mulu National Park (on Gunung Api and low limestone

outcrops along the Sungai Melinau).

Habitat: It grows from the base of limestone outcrops to c. 1170 m in lower

montane limestone forest on Gunung Api, usually on limestone boulders

but also on steep screes in deeply shaded habitats.

Begonia conipila from Sarawak 289

Notes: Begonia conipila is typical of section Petermannia in possessing

upright (although weak) stems (as opposed to prostrate rhizomes),

protogynous inflorescences with the female flower basal and the male

flowers distal, male flowers with two tepals and obovate anthers that open

by pores, and female flowers with five tepals, three free styles and a 3-

loculate ovary with bilamellate placentas. The fruit is interesting in that

one locule is hardly developed (it is about a quarter the size of the other

two). A similar situation is seen in B. amphioxus (Kiew, 2001).

In its creeping stem with erect branches, broadly obovate leaves with

stiff pale hairs, serrate leaf margin with each tooth tipped by a long hair,

the few stamens in the male flowers, and the 3-loculate ovary with

bilamellate placentas, it resembles B. havilandii Ridl. from Sarawak.

However, it 1s distinct from this species as B. conipila has smaller leaves

(they are c. 6—10 cm long in B. havilandii), male flowers with two tepals

(four in B. havilandii) and an oblong capsule (orbicular in B. havilandii).

With its small, variegated leaves, it is as Low noted on his specimen,

‘a charming plant’. Jangowe (1999) illustrates this species with a fine colour

photograph.

Specimens examined: SARAWAK: Gunung Mulu National Park - Low s.n. “Mooloo’ (K);

Gunung Api - Abang Moktar $49606 (SAN, SAR), Anderson S4287 (K, SAR), $30732

(SAR, SING), B.L. Burtt B8354 (SAR), P. Chai S30054 (SAR), $35790 (SAR), C. Hansen

102 (SAN, SAN), P.J. Martin S$38257 (SAR), I. Nielsen 502 (SAR), B.C. Stone 13693

(SAR), Yii P.C. & Abu Talib $58845 (SAR); limestone on Sungai Melinau Chew Wee Lek

CWL 1055 (SAR, SING), B.C. Stone 13598 (KLU, SAR).

Acknowledgements

The author thanks the curators of the herbaria at BM, E, K, KLU, SAN

and SAR for permission to examine specimens in their care, to Mark J.E.

Coode for his patience in correcting the Latin, and to M.J.S. Sands for

comments on the manuscript.

References

Anderson, J.A.R. and P.P.K. Chai. 1982. Gunung Mulu National Park,

Sarawak. 9. Vegetation. Sarawak Museum Journal. 30: 195—206.

Jangowe, J. 1999. Begonias of the national parks of Malaysian Borneo.

Begonian. July/Aug: 144—145.

Kiew, R. 2001. The limestone begonias of Sabah, Borneo — Flagship species

for conservation. Gardens’ Bulletin Singapore. 53: 241—286.

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Gardens’ Bulletin Singapore 53 (2001) 291-313 .

New Species, Varieties and Reductions in Diospyros

(Ebenaceae) in Borneo and Peninsular Malaysia

including Peninsular Thailand

FRANCIS S.P. NG

*/o Forest Research Institute Malaysia, Kepong, 52109 Kuala Lumpur, Malaysia

Abstract

In the genus Diospyros, seven new species (D. beccarioides Ng, D. brainii Ng, D. crockerensis

Ng, D. keningauensis Ng, D. lunduensis Ng, D. multinervis Ng and D. parabuxifolia Ng)

and six new varieties (D. curranii Merr. var. kalimantanensis Ng; D. ferruginescens Bakh.

var. rufotomentosa Ng; D. lanceifolia Roxb. var. iliaspaiei Ng, var. renageorgei Ng, var.

saliciformis Ng; D. penibukanensis Bakh. var. scalarinervis Ng) are described. Thirty species

or varieties are reduced to synonymy.

Introduction

In revising the genus Diospyros for the Tree Flora of Sabah and Sarawak,

I took the opportunity to review the genus for Borneo and Peninsular

Malaysia. This has resulted in the recognition of seven new species and six

new varieties, and the reduction of 30 species or varieties to synonymy.

New Species

1. Diospyros beccarioides Ng, sp. nov.

Arbor ad 20 m alta; rami dense rubro-brunnee pubescentes demum

glabrescentes. Folia membranacea ad chartacea, glabra, oblonga ad ovato-

oblonga 16-30 cm longa 5.5—11 cm lata, basi cuneata leviter attenuata rare

rotundata, apice acuminato, costa supra immersa plana vel ‘marginibus’

elevatis provisa, infra nervis lateralibus prominentibus paribus 7-13

incurvatis ante margines anastomosantibus venam intra-marginalem plus

minusve distinctam formantibus, venatione intercostali prominula laxe

scalariformi; petiolus 0.8-1.5 cm longus. Inflorescentia mascula cymis

subsessilis condensatis floribus ut videtur 3 vel plus sed ignotis. Fructus 1—

3 pedicellis 0.3-0.8 cm longis suffulti, globosi ad 2.5 cm diam. glabri, calyce

in lobis 4 vel 5 erectis folioso-coriaceis plicatis accrescentibus ad c. 2 cm

longis 1.5 cm latis diviso. Typus: Yii et al. S 50325, Sarawak (KEP holo,

SAR iso).

292 Gard. Bull. Singapore 53 (2001)

Figure 1

Tree to 20 m tall. Twigs densely reddish brown pubescent, becoming

glabrous with age. Leaves membranaceous to chartaceous, glabrous; oblong

or oblong-obovate, 16-30 x 5.5—-11 cm, base cuneate and slightly attenuate,

rarely rounded, apex acuminate; midrib above sunken, flat, or sunken with

raised sides; lateral veins prominent below, 7-13 pairs, inarching and

anastomosing at the margins to form a more-or-less distinct intramarginal

vein; intercostal venation prominulous below, laxly scalariform; petiole 8—

15 mm long. Male inflorescences subsessile condensed cymes of 3 or more

flowers. Flowers unknown. Fruits 1-3, on 3-8 mm long stalks, globose, to c.

2.5 cm diam., glabrous. Fruit calyx divided into 4-5 valvate lobes, the lobes

erect, leafy-coriaceous, plicate, accrescent, to c. 2 x 1.5 cm.

Distribution and habitat: Sumatra (Jacobs 8341, Lampung Prov.); Borneo,

Sarawak (S 8633, S 16177, S 31817, S 32625, S 50325, S 65187; Chew CWL

1100, 1141); Kalimantan (Endert 5164, 5380; Kessler et al. 824, 969); Sulawesi

(Vogel & Vermuelen 6981). In Sarawak, the species is found on limestone

hills, up to 200 m altitude.

Notes: Bakhuizen in Bull. Jard. Bot. Btz. Ser. iii. 15 (1938) 239 had included

specimens of three species under Diospyros beccarii Hiern. The two

syntypes of D. beccarii Hiern belong to D. sumatrana Migq., hence D. beccarii

has to be reduced to D. sumatrana. At least two other of his cited specimens,

bb. 12051 and bb. 14750, belong to D. daemona Bakh.. The third element,

represented by, for example, Endert 5164 belongs to this new species, D.

beccarioides, which can be distinguished from D. sumatrana by its larger

fruits with accrescent calyx lobes up to 2 cm long, and larger leaves, with a

stronger tendency to form an intramarginal vein.

2. Diospyros brainii Ng, sp. nov.

Arbores mediocres ad 29 m altae. Folia mediocra ad magna, oblongo-

ovata, 11.5—19.5 x 2.5-6 cm, apice acuminata, basi cuneata, lamina chartacea,

glabra, costa supra immersa; nervi secundarii infra prominuli ad prominentes

8-10 pares intus curvati in nervo intramarginali inconspicuo ad circa tertiam

spatiam inter marginem costamque anastomosantes; nervi tertiarii

inconspicui reticulati; petiolus 0.5—1 cm. Flores ignoti. Fructus solitarius,

sessilis, globosus ad 4.5 cm diam., glaber, in sicco valde rugosus. Calyx

accrescens, in lobis 4 vel 5 coriaceis valvatis 1.5 cm longis profunde divisus,

in quoque axe principali sursum curvato fructus curvaturam secundo sed

marginibus inter se deorsum curvatis. Typus: Brain anak Tada, S 15932,

293

rioides Ng. A. leafy twig: and B. fruit. (A-B from S 50325).

Figure 1. Diospyros becca

294 Gard. Bull. Singapore 53 (2001)

Figure 2. Diospyros brainii Ng. A. leaty twig and B. fruit. (A—B from S 15932).

Diospyros (Ebenaceae) in Borneo and Peninsular Malaysia 295

Borneo, Sarawak (SAR holo, SAN, SING iso).

Figure 2

Medium trees to 29 m tall. Leaves medium-sized to large, oblong-obovate,

11.5-19.5 x 2.5-6 cm, apex acuminate, base cuneate; lamina chartaceous,

glabrous; midrib sunken above; lateral veins prominulous to prominent

below, 8-10 pairs, inarching and anastomosing to form a faint looping

intramarginal vein at one third of the distance from the margin to the

midrib; intercostal venation faint, reticulate; petiole 5-10 mm long. Flowers

unknown. Fruit solitary, sessile, globose, to 4.5 cm diam., drying much

wrinkled and shrivelled, glabrous. Fruit calyx accrescent, divided deeply

into 4-5 coriaceous valvate lobes, 1.5 cm long, the main axis of each lobe

curved upwards following the curvature of the fruit base, but the sides

bent backwards towards each other.

Distribution and habitat: Endemic to Borneo: Sarawak (S 15932 and S 27961).

Lowland forest at Bintulu.

Notes: The location of the intramarginal vein at some distance from the

margin is reminiscent of Diospyros neurosepala Bakh., which differs in

having leafy calyx lobes. The fruiting calyx recalls that of D. sulcata

Kosterm., which differs in the absence of an intramarginal vein in the leaf.

3. Diospyros crockerensis Ng. sp. nov.

Arbores parvae ad 11 m altae; ramuli dense velutini. Folia mediocra,

elliptica, 7-17 x 2.5-8.3 cm, apice acuminata, basi cuneata, lamina chartacea

infra dense pubescentia, costa supra immersa sulco puberulo; nervi

secundarii subtus prominentes 6—9 pares; nervi tertiarii subtus prominuli

laxe scalariformes; petioli c. 0.5 cm longi dense velutini. Flores masculi 3-9

vel plus ad extremos cymarum elongatarum 24.5 cm dispositi, calyx in

lobis 4 anguste oblongis imbricatis profunde divisus, corolla salveriformis

ad 0.8 cm longa. Flores feminei c. 3-floris ad extremos cymarum elongatarum

0.82.8 cm dispositi, calyx in lobis 4 imbricatis profunde divisus. Pedicellus

in fructu 0.8-2.8 cm. Calyx non-accrescens, lobis c. 0.2 x 0.1 cm. Fructus

plerumque solitarii, oblongo-ovoidei ad 3.5 x 1.5 cm, velutini, apice in

acumine acuto gradatim contracto, pericarpio tenui lignoso. Typus: Dewol

& Karim SAN 78389, Borneo, Sabah, Crocker Range (SAN holo, SING

isO).

296 Gard. Bull. Singapore 53 (2001)

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Figure 3. Diospyros crockerensis Ng. A. leafy twig; B. fruit; C. female flowers after shedding of

corollas; D. male flower bud; E. male flower bud in longitudinal section. (A—C from SAN

78389, D-E from § 25273).

Diospyros (Ebenaceae) in Borneo and Peninsular Malaysia 297

Figure 3

Small trees to 11 m tall. Twigs densely velvety. Leaves medium-sized, elliptic,

7-17 x 2.5-8.3 cm, apex acuminate, base cuneate; lamina chartaceous,

densely hairy below; midrib sunken above, the groove covered with short

hairs; lateral veins prominent below, 6-9 pairs; intercostal venation

prominulous below, laxly scalariform:; petiole c. 5 mm long, densely velvety.

Male flowers in clusters of 3-9 or more, at the ends of elongated 24.5 cm

long cymes; calyx divided deeply into 4 narrowly oblong imbricate lobes;

corolla salverform, to 8 mm long. Female flowers in clusters of about 3, at

the ends of elongated 8-28 mm cymes; calyx divided deeply into 4 imbricate

lobes. Fruits usually solitary, on 8-28 mm stalks, oblong-ovoid, with the

apex gradually tapered to a sharp point, to 3.5 x 1.5 cm, velvety, with thinly

woody wall. Fruit calyx non-accrescent, the lobes c. 2 x 1 mm.

Distribution and habitat: Endemic to Borneo: Sabah (SAN 78389, SAN

11589, SAN 130109) and Sarawak (S 25273). In hill forests.

4. Diospyros keningauensis Ng, sp. nov.

Arbores mediocres ad magnae, ad 40 m altae. Folia minuta ad mediocra,

anguste elliptica ad ovata 3.5—13 x 1.5-4 cm, apice acuminata, basi cuneata

ad rotundata, lamina chartacea glabra ad infra sparse appresse pubescentia,

proprie rugosa in sicco, costa supra prominula in juventute appresse

pubescens; nervi secundarii subtus prominentes 5-8 pares; nervi tertiarii

subtus prominuli scalariformes; petioli 0.5—-1 cm. Flores masculi 3—5 in

cymis 0.5-1.5 cm dispositi, quae in quoque nodo 2 vel plus glomeratae

possunt; calyx in lobis 4 valvatis divisus; corolla salveriformis 1.4 cm longa.

Flos femineus solitarius pedunculo 0.5—1 cm; calyx lobis valvatis 4 (5) divisus.

Pedicellus in fructu 0.5—1 cm. Calyx non-accrescens, lobi c. 0.5 x 0.5 cm.

Fructus solitarius, globosus ad 4.5 cm diam., in juventute dense pubescens,

in sicco pericarpio crasso irregulariter rugosus. Typus: Leopold & Saikah

SAN 74495, Borneo, Sabah (KEP holo, SAN, SING iso).

Figure 4

Medium and big trees to 40 m tall. Leaves small to medium-sized, narrowly

elliptic to ovate, 3.5-13 x 1.5-4 cm, apex acuminate, base cuneate to

rounded; lamina chartaceous, glabrous to sparsely appressed hairy below,

drying characteristically wrinkled; midrib prominulous above, appressed

hairy when young; lateral veins prominent below, 5-8 pairs; intercostal

veins prominulous below, scalariform; petiole 5-10 mm long. Male flowers

298 Gard. Bull. Singapore 53 (2001)

Figure 4. Diospyros keningauensis Ng. A. leafy twig; B. female inflorescence; C. female flower;

D. female flower in longitudinal section; E. as D but after removal of ovary to show staminodes;

F ovary in transverse section. G. male inflorescence; H. male flower bud; I. male flower bud in

longitudinal section; J. fruit. (A-F from SAN 26704, B—D from SFN 27136, G-I from SFN

27136, J from SAN 74495).

Diospyros (Ebenaceae) in Borneo and Peninsular Malaysia 299

in clusters of 3-5 on 5-15 mm long cymes, which may be clustered in 2 or

more to each node; calyx divided into 4 valvate lobes; corolla salverform,

c. 1.4 cm long. Female flowers solitary on a 5-10 mm peduncle; calyx

divided into 4(—5) valvate lobes; corolla 4(—5)-lobed; staminodes 8; ovary

bearing 8 ovules. Fruits solitary, on 5—10 mm long stalks, globose, irregularly

wrinkled on drying, thick-walled, to 4.5 cm diam., densely hairy when

young. Fruit calyx not accrescent, the lobes about 5 x 5 mm.

Distribution and habitat: Endemic to Borneo: Sabah (Carr SFN 27136;

SAN 26704, SAN 33783, SAN 71930, SAN 74495). Lower montane and

montane forests at 1300-1500 m in the vicinity of Mt Kinabalu (Keningau,

Ranau).

5. Diospyros lunduensis Ng, sp. nov.

Arbores mediocres ad 25 m altae. Folia mediocra, elliptica ad oblongo-

elliptica 5.6-14 x 2.8-6 cm, apice breviter acuminata vel acuta, basi rotundata

ad subcordata, lamina coriacea, glabra, supra in sicco ad nigra vergentia,

costa supra immersa; nervi secundarii subtus prominentes 5-7 pares; nervi

tertiarii supra prominuli laxe reticulati; petioli 0.5—0.8 cm. Flores masculi

ignoti, feminei solitarii pedicellis 0.2—0.4 cm longis suffulti, calyce in lobis 4

rotundatis imbricatis pubescentibus profunde diviso. Pedicelli in fructu 0.2—

0.4 cm longi. Calyx non-accrescens, lobis ad 0.6 x 0.6 cm patentibus ad

reflexis. Fructus solitarius, globosus ad 2 cm diam., glaber vel sparse appresse

pubescens. Typus: Othman S 49996, Borneo, Sarawak, Lundu, G. Pueh

(KEP holo, SAN iso).

Figure 5

Medium-sized trees to 25 m tall. Leaves medium-sized, elliptic to oblong-

elliptic, 5.6-14 x 2.8-6 cm, apex shortly acuminate or acute, base rounded

to subcordate; lamina coriaceous, glabrous, tending to dry black above;

midrib sunken above; lateral veins prominent below, 5-7 pairs; intercostal

venation prominulous below, laxly reticulate; petiole S-8 mm long. Male

flowers unknown. Female flowers solitary, on 2-4 mm long stalks; calyx

deeply divided into 4 rounded imbricate hairy lobes; corolla 4-lobed;

staminodes 4; ovary with 4 ovules. Fruits solitary, on 2-4 mm long stalks,

globose, to 2 cm diam., glabrous or sparsely appressed hairy. Fruit calyx

not accrescent, the lobes to 6 x 6 mm, spreading to reflexed.

Distribution: Endemic to Borneo: Sarawak (S 49996 and S 25482).

300 Gard. Bull. Singapore 53 (2001)

Figure 5. Diospyros lunduensis Ng. A. leafy flowering twig; B. female flower bud; C. female

flower bud in longitudinal section; D. as C but with ovary removed to show staminodes; E.

ovary in transverse section; F. fruit. (A—E from § 25482, F from S$ 49996).

Diospyros (Ebenaceae) in Borneo and Peninsular Malaysia 301

6. Diospyros multinervis Ng, sp. nov.

Arbores mediocres ad 20 m altae. Folia magna, oblongo-lanceolata 21-32

x 5.8-9.2 cm, apice acuminata, basi subcordata, lamina chartacea glabra,

costa supra immersa: nervi secundarii subtus prominentes 15-17 pares:

nervi tertiarii subtus prominentes incerte scalariformes; petioli 0.8—1 cm.

Flores ignoti. Cymae in fructu laxe ramosae 2.5-3.5 cm longae: pedicelli 1—

1.5 cm. Calyx patelliformis, 4-radiatus, in lobis 4 valvatis divisus marginibus

coreaceis reflexis c. 2 cm diam. Fructus 3—5, globosi 2-3 cm diam., glabri.

Typus: Burley et al. 2726, Borneo, Kalimantan (KEP holo).

Figure 6

Medium-sized trees to 20 m tall. Leaves large, oblong lanceolate, 21—32 x

5.89.2 cm, apex acuminate, base subcordate; lamina chartaceous, glabrous;

midrib sunken above; lateral veins prominent below, 15-17 pairs; intercostal

venation prominent below, vaguely scalariform: petiole 8-10 mm long.

Flowers unknown. Fruits 3-5, with 1-1.5 cm pedicels on laxly branched

cymes 2.5—3.5 cm long, globose to elliptic, 2—3 cm diam., glabrous, puckered

on drying. Fruit calyx c. 2 cm diameter divided into 4 valvate lobes, forming

a 4-pointed plate, with the sides of the lobes coriaceous and reflexed.

Distribution and habitat: Endemic to Borneo: Kalimantan (Burley et al.

2472 and 2726, G. Bentuang area of W. Kalimantan). Lowland forest.

Notes: This species is very close to Diospyros wallichii King & Gamble,

differing in the glabrous fruits borne on individual pedicels 1—-1.5 cm long,

which arise from peduncles 1-2 cm long (adding up to an infructescence c.

3.5 cm long excluding the fruits). Overall, the inflorescence/infructescence

is much longer than that of D. wallichii.

The local Malay name tuba monyet (monkey tuba) indicates that the

fruits are used as a kind of fish poison.

7. Diospyros parabuxifolia Ng. sp. nov.

Arbores mediocres, ad 15 m altae. Folia minuta, obovata 1.5—2 x 0.8-1.5

cm, apice rotundata acuta vel breviter acuminata, basi cuneata ad attenuata,

lamina chartacea glabra, costa supra immersa ad prominens; nervi secundarii

et tertiarii invisibiles; petioli 0.1—-0.2 cm. Flores ignoti. Pedicelli in fructu

elongati 1.5-2 cm. Calyx non-accrescens, in lobis 4 imbricatis rotundatis

0.2 x 0.2 cm profunde divisus. Fructus solitarius, ellipsoideus c. 2 x 1 cm,

pericarpio angusto in sicco corrugato fere glabro. Typus: //ias Paie S 33017,

Gard. Bull. Singapore 53 (2001)

302

Figure 6. Diospyros multinervis Ng. A. leafy twig and B. fruit. (A-B from Burley et al. 2726).

Diospyros (Ebenaceae) in Borneo and Peninsular Malaysia 303

Borneo, Sarawak, Lawas, Bukit Batanga (KEP holo; SAN, SAR iso).

Figure 7

Medium-sized trees to 15 m tall. Leaves small, obovate, 1.5—2 x 0.8-1.5 cm,

apex rounded, acute or shortly acuminate, base cuneate to attenuate; lamina

chartaceous, glabrous; midrib sunken to raised above; lateral veins invisible;

intercostal venation pattern invisible; petiole 1-2 mm long. Flowers

unknown. Fruits solitary, on elongated 1.5—2 cm long stalks, ellipsoid, c. 2 x

1 cm, thin-walled and puckered on drying, almost glabrous. Fruiting calyx

not accrescent, divided deeply into 4 imbricate rounded lobes, c. 2 x 2 mm.

Distribution and habitat: Endemic to Borneo: Sarawak, Lawas and Kapit

(S 33017, S 60062). Lower montane forests at 1030-1500 m altitude.

Notes: This species differs from Diospyros buxifolia (Blume) Hiern mainly

in its elongated fruit stalk. From D. graciliflora Hiern, which also has an

elongated fruit stalk, it differs in the fruit, which is ellipsoid (not globose).

In addition, the fruit stalk of D. graciliflora is much more slender.

New Varieties

1. Diospyros curranii Merr., Phil. J. Sc. Bot. 4 (1909) 306.

var. curranii

Synonym: D. curraniopsis Bakh., Gard. Bull. S. S. 7 (1933) 168, syn. nov.;

types: Endert 4883, Borneo, Kalimantan (BO), Elmer 21464, Borneo, Sabah

(BO).

var. kalimantanensis Ng, var. nov.

A varietate typica in calyce frutescenti patenti nec reflexo multo amplificato

quo fundibulum fructu centrali formanti differt. Typus: Veldkamp 8148,

Borneo, Kalimantan, Tumbang Tubus S of Bukit Raya (KEP holo, BO, L

iso).

Distribution: Borneo, Kalimantan, at Bukit Raya (Mogea 3569, 3585, 3621;

Veldkamp 8148).

Notes: This differs from the typical variety in the fruiting calyx being

outstretched instead of reflexed, and much enlarged, so that the calyx

304 Gard. Bull. Singapore 53 (2001)

Figure 7. Diospyros parabuxifolia. A. leafy twig and B. fruit. (A-B from S 33017)

Diospyros (Ebenaceae) in Borneo and Peninsular Malaysia 305

forms a 5-lobed funnel with the fruit in the centre. This variety appears to

replace var. curranii in the vicinity of Bukit Raya, so male and sterile

collections from Bukit Raya can be assumed to belong to this variety

rather than the type variety.

2. Diospyros ferruginescens Bakh., Bull. Jard. Bot. Btz. Ser. iii, 15 (1941)

409.

var. rufotomentosa Ng, var. nov.

A varietate typica praecipue in foliis infra dense rufe tomentosis differt:;

folia etiam latiora (ad 5.5 cm lata) possunt, nervis secundariis

prominentioribus distincte incurvatis. Typus: Maikin et al. SAN 132942,

Borneo, Sabah, Lumaku FR (SAN holotype).

Medium-sized trees to 20 m tall. Twigs densely rusty hairy. Leaves medium-

sized, oblong-elliptic, 6.5—11.5 x 3-5.5 cm, apex acuminate, base acute;

lamina chartaceous, densely rusty hairy below, tending to dry black above;

midrib sunken above, the groove often filled with hairs; lateral veins

prominent below, 6-10 pairs, inarching strongly at the margins; intercostal

veins prominent below, reticulate; petiole c. 5 mm long. Male flowers in

clusters of 3—9, shortly pedicelled, sharing a common 5-7 mm peduncle;

calyx divided into 4 small triangular valvate teeth: corolla salverform, 5—7

mm long. Female flowers not known. Fruits solitary, on 1-2 mm long stalks,

ovoid to globose, to 1.5 cm diam., densely rusty hairy. Fruit calyx accrescent,

divided deeply into 4-5 triangular valvate lobes, forming a 4-5 pointed

star, each lobe c. 5 x 4 mm.

Distribution and habitat: Endemic to Borneo: Sabah (SAN 55535, SAN 66013,

SAN 102122, SAN 113977, SAN 132942, SAN 139235), Kalimantan

(Kostermans 8778). Lowland and hill forests up to 700 m.

Notes: This differs from the typical variety mainly in the underside of the

leaves, which is densely rusty-hairy. The leaves may also be broader (to 5.5

cm wide) with the lateral veins more prominent and distinctly inarching.

3. Diospyros lanceifolia Roxb., Hort. Beng. (1814) 93.

var. iliaspaiei Ng, var. nov.

Arbores mediocres ad 27 m altae; ramuli dense ferrugineo-pubescentes.

306 Gard. Bull. Singapore 53 (2001)

Folia mediocra ad magna, oblonga ad oblongo-lanceolata 13.5-35 x 3.5-14

cm, apice acuminata ad acuta, basi cuneata ad rotundata interdum fovearum

glandulosarum pare uno instructa, lamina coriacea infra dense ferrugineo-

pubescentia inter venas bullata, costa supra immersa; nervi secundarii subtus

prominentes, supra immersi, 15—24 paribus incurvatis ad marginem in nervo

intramarginali conspicuo anastomosantibus; nervi tertiarii subtus

prominentes supra immersi transverse reticulati; petioli 1-2.4 cm. Flores

feminei c. 3, in cymis brevibus 0.2—0.5 cm dispositi. Pedunculi in fructu

breves, 0.2-0.5 cm longi. Calyx lobis c. 1 x 0.8 cm marginibus reflexis.

Fructus 1-3, globosi nitide nigri, ad 2 cm diam, in juventute velutini. A var.

consanguinea in nervis secundariis laminae quoque latere in nervo

intramarginali valde evoluto anastomosantibus differt. Typus: /lias Paie S

50180, Borneo, Sarawak (KEP holo, SAN, SAR iso).

Medium-sized trees to 27 m tall. Twigs densely rusty hairy. Leaves medium-

sized to large, oblong to oblong-lanceolate, 13.5-35 x 3.5-14 cm, apex

acuminate to acute, base cuneate to rounded, sometimes bearing a pair of

pit-glands; lamina coriaceous, densely rusty hairy below, bullate between

the veins; midrib sunken above; lateral veins sunken above, prominent

below, 15-24 pairs, inarching and joining at the margins to form a prominent

intramarginal vein; intercostal venation sunken above, prominent below,

reticulo-transverse; petiole 1—2.4 cm long. Female flowers in clusters of c. 3

on short 2-5 mm long cymes. Fruits 1-3, on short 2-5 mm long stalks,

globose, shiny black, to 2 cm diam., velvety when young. Fruit calyx: lobes

c. 10 x 8 mm, with sides reflexed.

Distribution and habitat: Endemic to Borneo: Sarawak (S 24803, S 39109, S$

41224, § 50180, AZ 5671). Lowland forests.

var. renageorgei Ng, var. nov.

Arbores mediocres ad 21 m altae. Folia mediocra, elliptica ad ovata 6—19.5

x 4-8.5 cm, apice acuminata, basi rotundata paullo attenuata saepe fovearum

glandulosarum pare uno provisa, lamina chartacea glabra supra plerumque

subtus interdum nitentia, inter venas paullo bullatum, costa supra immersa;

nervi secundarii supra immersi, subtus prominentes 6—12 pares; nervi tertiaril

supra immersi subtus prominuli transverse reticulati; petioli 0.8—1.5 cm.

Flores feminei 1-5 in cymis ad 0.6 cm longis dispositi. Pedicelli in fructu

0.3-0.6 cm. Calyx lobis ad 1 x 1 saltem marginibus reflexis. Fructus circa 3,

globosi 1.3 cm diam., velutini. A var. /anceifolia in laminae basi rotundata

nec cuneata differt. Typus: Rena George S 43068, Borneo, Sarawak,

Limbang (KEP holo, SAN iso).

Diospyros (Ebenaceae) in Borneo and Peninsular Malaysia 307

Medium-sized trees to 21 m tall. Leaves medium-sized, elliptic to ovate, 6—

19.5 x 4-8.5 cm, apex acuminate, base rounded, slightly attenuate, and

often bearing a pair of pitted glands: lamina chartaceous, glabrous, usually

shiny on the upper surface and sometimes on the lower surface, slightly

bullate between the veins; midrib sunken above: lateral veins sunken above,

prominent below, 6—12 pairs: intercostal venation sunken above, prominent

below, reticulo-transverse; petiole 8-15 mm long. Female flowers in clusters

of 1—5 on cymes up to 6 mm long. Fruits about 3, on 3-6 mm long stalks,

globose, c. 1.3 cm diam., velvety. Fruit calyx with lobes up to 1 x 1 cm, the

lobes or at least their sides reflexed.

Distribution and habitat: Endemic to Borneo: Sabah (SAN 99668), Sarawak

(S 17862, S 17897, S 42254, S 43068, S 47134) and Brunei (Coode 7050).

Lowland forests.

Notes: Named in honour of Rena George, 1956—1994, botanist of the

Sarawak Forestry Department, who sacrificed her life for the sake of

advancing botanical knowledge.

Specimen S 36348 from Kapit at 950 m altitude is placed here with

some doubt; its leaves are not shiny like the others.

var. saliciformis Ng. var. nov.

Arbores mediocres ad 20 m altae. Folia elongata, saliciformia 5.5—11.5 x 1-

3.2 cm, apice gradatim contracta, basi gradatim contracta et attenuata

(glandes desunt), lamina chartacea glabra, costa supra immersa: nervi

secundarii prominuli ad fere invisibiles 5—7 pares; nervi tertiarii subtus fere

invisibiles; petioli 0.3-0.5 cm. Flores ignoti. Pedicelli in fructu 0.1—-0.2 cm.

Calyx stellae 4-radiatae similis, lobis ad 0.6 x 0.4 cm. Fructus solitarius,

subsessilis, globosus ad 1.5 cm diam., velutinus glabrescens ad glaber. A

varietatibus ceteris in laminis angustis saliciformibus differt. Typus: Coode

7826, Borneo, Brunei (KEP holo, BRUN, SAN, SING iso).

Medium-sized trees to 20 m tall. Leaves elongated, willow-like, 5.5-11.5

x 1-3.2 cm, apex gradually tapered, base gradually tapered and attenuate,

without pit-glands; lamina chartaceous, glabrous: midrib sunken above;

lateral veins prominulous to almost invisible, 5—7 pairs; intercostal

venation practically invisible below; petiole 3-5 mm long. Flowers

unknown. Fruits solitary, subsessile on 1-2 mm stalks, globose, velvety

becoming glabrous, to 1.5 cm diam. Fruit calyx a 4-pointed woody star

with the lobes up to 6 x 4 mm.

308 Gard. Bull. Singapore 53 (2001)

Distribution and habitat: Endemic to Borneo: Sarawak (Chew 1170, S 49201)

and Brunei (Coode 7826). Lowland forest.

Notes: The leaves of Diospyros lanceifolia var. saliciformis are superficially

similar to those of D. eucalyptifolia Bakh. and the narrow-leafed riverine

form of D. andamanica (Kurz) Bakh.

3. Diospyros penibukanensis Bakh., Bull. Jard. Bot. Btz. Ser. 111, 15 (1937)

124.

var. scalarinervis Ng, var. nov.

A var. penibukanensi in folis subtus glabris nec dense pubescentibus, nervis

tertiariis scalariformibus remotioribus, ramulorum et foliorum pilis tenuibus

appressis nec patentibus differt. Varietates etiam geographicaliter disjunctae

prima Sarawak secunda Sabah incola. Typus: S 56607, Borneo, Sarawak

(KEP holo, SAN, SAR iso).

Small trees to 10 m tall, rarely to 25 m. Twigs velvety on young parts.

Leaves medium-sized to large, elliptic to broadly obovate or oblong-obovate,

11-38 x 4-15 cm, apex acuminate, base subcordate, rarely only rounded;

lamina chartaceous, glabrous except the veins; midrib sunken above; lateral

veins prominulous below and finely appressed hairy, 5—24 pairs; intercostal

venation prominent below, scalariform; petiole 8-5 mm long. Male flowers

up to 10 in a condensed cyme c. 5 mm long; calyx with 4 imbricate triangular

lobes. Female flowers up to 10 in a condensed cyme c. 5 mm long; calyx

with 4 imbricate semicircular lobes. Fruits in clusters of 1-3, on c. 5 mm

long stalks, globose, to 2.5 cm diam., velvety. Fruit calyx not accrescent,

the calyx lobes rounded, coriaceous, 4-6 x 5-8 mm.

Distribution and habitat: Endemic to Borneo: Sarawak (S 13586, S 13681,

S 13702, S 18639, S 34482, S 34935, S 34962, S 35271, S 41850, S 49997,

S 56607). Lowland, hill and heath forests.

Notes: Variety scalarinervis differs from var. penibukanensis in its leaves,

which are glabrous below (instead of densely hairy), the scalariform

intercostal venation being spaced further apart, and the hairs on its twigs

and leaves being fine and appressed (instead of spreading). They are also

geographically disjunct, the former in Sarawak and the latter in Sabah.

Diospyros (Ebenaceae) in Borneo and Peninsular Malaysia 309

Reductions

1. Diospyros andamanica (Kurz) Bakh., Bull. Jard. Bot. Btz. Ser. iii, 15

(1937) 74, (1941) 382.

Synonyms: D. tahanensis Bakh., Gard. Bull. S.S. 7 (1933) 163, syn. nov.;

type: Ridley s.n., Peninsular Malaysia, Pahang (SING). D. longepedunculata

Kosterm., Blumea 23 (1977) 462, syn. nov.; type: SAN 29807, Borneo,

Sabah (KEP, SAN, SAR, SING).

2. Diospyros areolata King & Gamble, J. As. Soc. Beng. 74, 2 (1905) 228.

Synonyms: D. bantamensis Koord. et Valeton ex Bakh., Gard. Bull. S.S. 7

(1933) 165, syn. novy.; type: Koorders 1718, Java (BO). D. malam Bakh.,

Gard. Bull. S.S. 7 (1933) 176, syn. noy.; types: bb. 6335, bb. 8037, Borneo,

Kalimantan (BO).

3. Diospyros blancoi A.DC., Prod. 8 (1844) 151.

Synonym: D. durionoides Bakh., Gard. Bull. S.S. 7 (1933) 169, syn. nov.;

types: bb. 7978, 10050 and 12456, Borneo, Kalimantan (BO).

4. Diospyros buxifolia (Blume) Hiern, Trans. Cambr. Phil. Soc. 12, 2 (1873)

218.

Synonym: D. elegantissima Bakh., Gard. Bull. S.S. 7 (1933) 169, syn. nov.;

type: Ridley 16349, Peninsular Malaysia, Pahang (SING).

5. Diospyros elliptifolia Merr., Phil. J. Sci. 30 (1926) 247.

Synonym: D. kinabaluensis (Bakh.) Kosterm., Blumea 23 (1977) 460, syn.

nov. [basionym: D. elliptifolia forma kinabaluensis Bakh., Bull. Jard. Bot.

Btz. Ser. iti, 15 (1935) 233]; types: DeVoogd 1486, Sumatra (BO); Haviland

1322, Borneo, Sabah (SING, SAR).

6. Diospyros foxworthyi Bakh., Gard. Bull. S.S. 7 (1933) 171.

Synonyms: D. /evigata Bakh., Gard. Bull. S.S. 7 (1933) 175, syn. nov.;

types: bb. 8053, 11089, 11255, Borneo, West Kalimantan (all at BO). D.

cylindrocarpa Kosterm., Blumea 23 (1977) 456, syn. nov.; type: SAN 7119,

Borneo, Sabah (SAN, SING).

7. Diospyros korthalsiana Hiern, Trans. Cambr. Phil. Soc. 12, 2 (1873)

168.

Synonym: D. myrmecocalyx (Hiern) Bakh., Gard. Bull. S.S. 7 (1933) 178,

syn. nov. [basionym: Maba myrmecocalyx Hiern, Trans. Cambr. Phil. Soc.

12, 2 (1873) 139]; type: Beccari 3568, Borneo, Sarawak (K).

310 Gard. Bull. Singapore 53 (2001)

8. Diospyros lateralis Hiern, Trans. Cambr. Phil. Soc. 12, 2 (1873) 167.

Synonym: D. crassipes Bakh., Gard. Bull. S.S. 7 (1933) 167, syn. nov.; type:

Beccari 2041, Borneo (K).

9. Diospyros mindanaensis Merr., Phil. J. Sci. 4 (1909) 309.

Synonym: D. endertii Bakh., Gard. Bull. S.S. 7 (1933) 169, syn. nov.; type:

Endert 4017, Borneo, Kalimantan (BO).

10. Diospyros montana Roxb., Pl. Corom. 1 (1795) 37.

Synonym: D. pubicalix Bakh., Gard. Bull. S.S. 7 (1933) 182, Bull. Jard.

Bot. Btz. 15 (1955) Plate 39, syn. nov.; type: Hamid C.F. 3827, Peninsular

Thailand (BKF, BO, KEP, SING).

Notes: Diospyros pubicalix has been an enigmatic species known only from

its type specimen. It is well illustrated by Bakhuizen (1955). The shoots

grow in flushes from resting buds, with the leaves exhibiting a characteristic

sequence of changes. The first leaves are small and have cordate to

subcordate bases. They are followed by larger leaves with rounded bases

and finally leaves with cuneate bases. On some specimens, only parts of

the sequence are evident, misleading authors not aware of this source of

variation to create new species.

11. Diospyros pilosanthera Blanco, FI. Filip. Ed. 1 (1837) 304.

var. pilosanthera

Synonym: D. cubica Bakh., Gard. Bull. S.S. 7 (1933) 168, syn. nov.; type:

Beccari 830, Borneo, Sarawak (K)

var. elmeri (Merr.) Ng, Malay. Forester 40 (1977) 238.

Synonym: D. nidus-avis Kosterm., Blumea 23 (1977) 464, syn. nov.; type: S

32181, Borneo, Sarawak (SAN, SAR, SING).

12. Diospyros plectosepala Hiern, Trans. Cambr. Phil. Soc. 12, 2 (1873)

ZO;

Synonyms: D. poiensis Bakh., Gard. Bull. S.S. 7 (1933) 181, syn. noy.; type:

Clemens 20304 (BO, SAR). D. setosa Bakh., Gard. Bull. S.S. 7 (1933) 184,

syn. nov.; type: Haviland 2324, Borneo (K).

13. Diospyros ridleyi Bakh., Gard. Bull. S.S. 7 (1933) 354.

Synonym: D. dajakensis Bakh., Gard. Bull. S.S. 7 (1933) 168, syn. nov.;

type: bb. 10490, Borneo, Kalimantan (BO).

14. Diospyros rufa King & Gamble, J. As. Soc. Beng. 74, 2 (1905) 228.

Diospyros (Ebenaceae) in Borneo and Peninsular Malaysia 311

Synonym: D. swingleri Kosterm., Blumea 23 (1977) 172, syn. nov.; type: S

26969. Borneo, Sarawak (SAR, SING).

15. Diospyros simaloerensis Bakh., Gard. Bull. S.S. 7 (1933) 185.

Synonym: D. paraoesi Bakh., Gard. Bull. S.S. 7 (1933) 180, syn. nov.; type:

bb. 10162, Borneo, Kalimantan (BO).

16. Diospyros styraciformis King & Gamble, J. As. Soc. Beng. 74, 2 (1905)

216.

Synonyms: D. styraciformis var. sarawakana (Bakh.) Ng, Malay. Forester

40 (1977) 241, syn. nov. [basionym: Diospyros sarawakana Bakh., Gard.

Bull. S.S. 7 (1933) 184]; type: Haviland 2316, Borneo, Kuching (K, SAR,

SING). D. clavipes Bakh., Gard. Bull. S.S. 7 (1933) 166, syn. nov.; type:

Teysmann 11397, Borneo, Kalimantan (BO).

17. Diospyros subrhomboidea King & Gamble, J. As. Soc. Beng. 74, 2

(1905) 209.

Synonyms: D. confusa Bakh., Gard. Bull. S.S. 7 (1933) 167, syn. nov.; type:

Baker s.n., Singapore (KEP, SING). D. jaheri Bakh., Gard. Bull. S.S. 7

(1933) 173, syn. nov.; type: Jaheri 128, Borneo, Kalimantan (BO).

18. Diospyros sumatrana Miq., P|. Jungh. (1851-55) 203.

Synonyms: D. monticola Kosterm., Blumea 23 (1977) 463, syn. nov.; type:

Clemens 32541, Borneo, Sabah (BO). D. beccarii Hiern, Trans. Cambr.

Phil. Soc. 12, 2 (1873) 204, syn. nov.; types: Beccari 2492 and 259], Borneo

(K).

19. Diospyros venosa Wall. ex A. DC., Prod. Syst. Nat. 8 (1844) 233.

var. venosa

Synonym: D. bintuluensis Kosterm., Blumea 23 (1977) 453, syn. nov.; type:

S 15912, Borneo, Sarawak (SAN, SAR).

var. olivacea (King & Gamble) Ng, Malay. Forester 40 (1977) 246.

Synonym: D. olivacea (King & Gamble) Kosterm., Blumea 23 (1977) 465,

syn. and stat. noy., [basionym: Maba olivacea King & Gamble in Williams,

Bull. Herb. Boiss. II, 5 (1905) 227]; type: King’s Collector 7877, Peninsular

Malaysia, Perak, Batang Padang (K, SING).

20. Diospyros wallichii King & Gamble in Williams, Bull. Herb. Boiss. 2,5

(1905) 429.

Synonym: D. pulchrinervia Kosterm., Blumea 23 (1977) 460, syn. nov.;

3 IBY Gard. Bull. Singapore 53 (2001)

type: Kostermans 21583, Borneo, Kalimantan (KEP, SAR, SING).

Acknowledgements

I am very grateful to the Director-General of Forest Research Institute

Malaysia for providing research funds and facilities under the Tree Flora

of Sabah and Sarawak Project, to the curators of the herbaria at BO, K,

KEP, L, SAN, SAR and SING for the loan of specimens and/or the excellent

help and facilities given during visits to their herbaria, to Dr. J.F. Veldkamp,

National Herbarium of the Netherlands, Leiden Branch, for the Latin

descriptions and to Mr Joseph Pao, Sarawak Forestry Department, for

meticulously preparing the illustrations.

Alphabetic List of Diospyros Species and Varieties Reduced

. bantamensis Koord. et Valeton ex Bakh. to D. areolata King & Gamble

. beccarii Hiern to D. sumatrana Miq.

. bintuluensis Kosterm. to D. venosa Wall. ex A.DC. var. venosa

. clavipes Bakh. to D. styraciformis King & Gamble

confusa Bakh. to D. subrhomboidea King & Gamble

. crassipes Bakh. to D. lateralis Hiern

cubica Bakh. to D. pilosanthera Blanco. var. pilosanthera

curraniopsis Bakh. to D. curranii Merr. var. curranti

cylindrocarpa Kosterm. to D. foxworthyi Bakh.

dajakensis Bakh. to D. ridleyi Bakh.

. durionoides Bakh. to D. blancoi A.DC.

elegantissima Bakh. to D. buxifolia (Bl1.) Hiern

. endertii Bakh. to D. mindanaensis Merr.

jaheri Bakh. to D. subrhomboidea King & Gamble

kinabaluensis (Bakh.) Kosterm. [Basionym: D. elliptifolia forma

nabaluensis Bakh.]| to D. elliptifolia Merr.

. levigata Bakh. to D. foxworthyi Bakh.

. longepedunculata Kosterm. to D. andamanica (Kurz) Bakh.

. malam Bakh. to D. areolata King & Gamble

. monticola Kosterm. to D. sumatrana Miq.

. myrmecocalyx (Hiern) Bakh. [Basionym: Maba myrmecocalyx Hiern]

to D. korthalsiana Hiern

D. nidus-avis Kosterm. to D. pilosanthera Blanco var. elmeri (Merr.) Ng

D. olivacea (King & Gamble) Kosterm. [Basionym: Maba olivacea King &

Gamble] to D. venosa Wall. ex A.DC. var. olivacea (King & Gamble) Ng

HOOSUEFHOHHHOHHHHyHyyNsssyd

Diospyros (Ebenaceae) in Borneo and Peninsular Malaysia 313

D. paraoesi Bakh. to D. simaloerensis Bakh.

D. poiensis Bakh. to D. plectosepala Hiern

D. pubicalix Bakh. to D. montana Roxb.

D. pulchrinervia Kosterm. to D. wallichii King & Gamble

D. setosa Bakh. to D. plectosepala Hiern

D. styraciformis var. sarawakana (Bakh.) Ng [Basionym: D. sarawakana

Bakh.] to D. styraciformis King & Gamble

D. swingleri Kosterm. to D. rufa King & Gamble

D. tahanensis Bakh. to D. andamanica (Kurz) Bakh.

Gardens’ Bulletin Singapore 53 (2001) 315-322.

New and Noteworthy Records of Mosses from Mindoro,

the Philippines, and Their Biogeographical Implication

BENITO C TAN

Department of Biological Sciences, National University of Singapore,

Singapore 119260

AND

EMELINA H MANDIA

Biology Department, De La Salle University, Taft Ave., Manila, The Philippines

Abstract

A recent, small collection of mosses from Mt. Halcon, Mindoro Island, the Philippines, has

yielded two new records for the country (Rhacocarpus alpinus (Wright) Par. and

Dicranoloma daymannianum Bartr.) and eight new records for the island. Relevant

ecological, morphological and biogeographical notes for the 15 species collected are provided.

Introduction

Floristic exploration was conducted from 1996—1997 around the summit

of Mt. Halcon in Mindoro, the seventh largest island in the Philippines

(Fig. 1). The actual summit stands at 2,582 m asl and has a grid coordinate

of 13° 15°46” N and 120° 59°29” E. The exploration was carried out

principally to classify and describe the vegetation types found on the

northeastern summit zone (Mandia, 1998). In addition to a good number

of new records of vascular plants for the island and the Philippines that

will be reported separately by the second author, the exploration has yielded

two species of mosses new to the country (Rhacocarpus alpinus and

Dicranoloma daymannianum), and eight new records for the island. In the

case of little known moss flora of Mt. Halcon, these additions are rather

significant. Furthermore, the new records highlight the Mindoro-Palawan

connection, a noteworthy feature of the island’s biogeography.

Summit Vegetation

The summit vegetation of Mt. Halcon extends from 1,950 to 2,582 m altitude

and consists largely of open, low growing species of Leptospermum,

Gard. Bull. Singapore 53 (2001)

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podocarps, ericoids, sedges, lycopods, ferns, ground lichens and bryophytes,

interrupted by patches of shrubs and mossy forest. Vascular cushion plants

and geophytes, such as the sundews (Drosera), Centrolepis philippinensis

Merr. (Centrolepidaceae), Trachymene saniculifolia Stapf (Umbelliferae),

Aletris foliolosa Stapf (Melanthiaceae), Oreobulus ambiguous Kukenth. &

Steenis (Cyperaceae), Patersonia lowii Stapf (Iridaceae), Gentiana

luzoniensis Merr. (Gentianaceae), Schoenus melanostachys R. Br. and S.

curvulus F. Muell. (Cyperaceae) abound. The soil is shallow, acidic and

oligotrophic, deriving from mica schist and white quartz.

Geological Origin and Biogeography of Mindoro

Little is known about the geology and geological history of Mt. Halcon,

except that the mountain mass consists of mainly metamorphic rocks of

continental crust origin uplifted since mid-Miocene (Fernandez, 1982; JICA-

MMASJ, 1984). Geologically, Mindoro, with Mt. Halcon, is interesting,

being ascribed to be part of a Tertiary micro-continent, the North Palawan

Block (Holloway, 1982). Others, such as Hamilton (1981), included only

the southern Mindoro in the North Palawan Block, giving the northern

portion a separate origin. The arrival of North Palawan Block from its

pre-drift position contiguous with the south China mainland to its present

day position in the Philippine archipelago has been postulated to be in

mid- to late Pleiocene (Holloway, 1982; Hall, 1996, 1998). The event

became a crucial step in the evolution of the modern Philippine biota

because the resulting island chain provided the necessary land habitats for

two-way migrations of plants and animals between Borneo and Luzon.

Furthermore, the drifting of the ancient North Palawan Block across the

South China Sea might also have carried with it some continental Asiatic

plants and animals, which have become incorporated into the Philippine

biota (Tan et al., 1988). Seen in this light, the study of Mindoro mosses

becomes relevant in understanding the biogeography of the island.

Floristic Affinity of Mindoro Mosses

Floristically, the vascular plant flora of Mindoro has been identified more

with the Luzon flora than with the Palawan flora (Merrill, 1928). Its

incompletely known moss flora also supported a strong Luzon connection

(Tan and Iwatsuki, 1991). It is therefore of phytogeographical significance

that Rhacocarpus alpinus, Dicranoloma daymannianum and Acroporium

johannes-winkleri were identified among the new collection of mosses from

Mt. Halcon. These three Malesian taxa are widespread in distribution, at

318 Gard. Bull. Singapore 53 (2001)

least in Borneo, but had either no prior Philippine record in the case of R.

alpinus and D. daymannianum, or were not previously known from

Mindoro, like Acroporium johannes-winkleri Broth. For all three taxa,

Mt. Halcon represents the northernmost penetration of their ranges in

Malesia north of the equator.

Rhacocarpus is a widespread austral genus of 7 species (Frahm, 1996),

with R. alpinus distributed from Sumatra, Borneo, Sulawesi to New Guinea

(Koponen and Norris, 1986). Its presence on Mt. Halcon shows the

Gondwana influence on the mountain flora, albeit a tenuous one, which

hitherto had not been apparent. The Mt. Halcon record also foretells the

likely presence of this species in Palawan. Likewise, the new Mindoro

record of Acroporium johannes-winkleri reinforces the important role of

Palawan as a Pleistocene land bridge in the island hopping of mosses in the

area. Earlier, Tan (1994) reported Acroporium johannes-winkleri from

two localities in Palawan, and one locality each from Mindanao (Mt. Apo)

and Sibuyan Island (Mt. Giting-Giting).

It is worthwhile to note that eight moss taxa reported from Mindoro

(see below) have no Palawan record. Under-collection is one reason for

this phenomenon. Compared with Mindoro, however, Palawan has a better

known moss flora owing to the four recent explorations undertaken between

1987—1993 by the senior author (Tan, 1996). An alternative explanation

for the dissimilar distribution of taxa in these two islands may lie in the

relatively “short” time available for the full exchange of biota since the

late Pliocene when the two islands drifted close to each other geographically.

Annotation of the new collection of Mt. Halcon mosses

Below we present the list of 15 moss species collected from Mt. Halcon,

together with brief comments on their morphology and ecology. The

collection numbers belong to the second author, with voucher specimens

deposited at SINU and herbarium of De La Salle University-Manila campus.

The single asterisk (*) indicates a new record for Mindoro, and double

asterisk (**), new to the Philippines.

* Acroporium johannis-winkleri Broth. [EHM 78]. Epiphytic on branches,

this is a medium-sized Acroporium with rather stiff-looking and

spreading leaves. Stems measure to 2 cm tall. It is a West Malesian

endemic, being common in Peninsular Malaysia and Borneo.

Braunfelsia dicranoides (Dozy & Molk.) Broth. [EHM 82]. This

dicranaceous moss is easily recognized among its congeners by the

New moss records from Mindoro, the Philippines 319

strongly falcate, non-plicate and unicostate leaves. Its long, sheathing

perichaetial leaves are also diagnostic. It is a common mossy forest

epiphyte throughout Malesia.

*Breutelia arundinifolia (Duby) Fleisch. [EHM 76, 83]. This large moss is

recognized by its big antheridial head, widely spreading to somewhat

squarrose and sheathing leaves, and tomentose stems (cf. Virtanen,

1997). Widespread in East Asia and Oceania, the species is a

common ground dweller on open sites at high elevations in the

Philippines. It has no Palawan record (Tan, 1996).

*Campylopus exasperatus (Nees & Blume) Brid. [EHM 77]. This is an

easily recognized Campylopus in the Philippine mountains. The

leaves are often broadly lanceolate, acute to blunt, with a percurrent

and broad costa that is poorly defined. The plants from Mt. Halcon

are large, measuring to 12 cm tall and carpeting the soil underneath

the Falcatifolium forest and heath vegetation.

*C. hemitrichius (C. Muell.) Jaeg. [EHM 86, 92]. Distinctive in having

narrowly lanceolate leaves with concolorous, excurrent costa, this

species is a ground dweller in open sites at the summit of Mt. Halcon.

There seem to be two ecotypes on this mountain, one with erect,

appressed leaves and the other with somewhat secund leaves. Tan

(1983) clarified the taxonomic confusion of this species vis-a-vis other

related Philippine congeners. At present, Campylopus hemitrichius

has no Palawan record.

Dicranoloma assimile (Hampe) Par. [EHM 84]. This is a widespread

Dicranoloma in Philippine mountains growing mainly on trunks and

branches, sometimes on soil.

** Dicranoloma daymannianum Bartr. [EHM 85]. Found attached to the

base of trunks and branches of shrubs in the Podocarpus-

Falcatifolium scrub, the present species resembles a small form of

D. braunii without the filamentous propagules. The presence of a

central strand in stem cross-section and the short, upper laminal

cells further distinguish it from D. braunii. Klazenga (1999) discussed

and illustrated well this uncommon Malesian taxon. Its presence in

Palawan can be predicted.

Leucobryum javense (Brid.) Mitt. [EHM 81]. This is the largest Leucobryum

found commonly in Philippine forests.

L. sanctum (Brid.) Hampe [EHM 80]. This is another common forest

species in Malesia including the Philippines. The opening of

perichetial bud of this epixylic and epipetric species are thickly

covered with highly branched rhizoids that arise mainly from the

abaxial side, but toward the base, of inner perichaetial leaves. The

320 Gard. Bull. Singapore 53 (2001)

larger outer perichaetial leaves, oddly, do not form any rhizoidal

outgrowth. Yamaguchi (1993) illustrated and labeled this structure

as a “tomentum” without any taxonomic comment.

* Macrothamnium javense Fleisch. [EHM 75]. The species is found attached

to prostrate branches. The strongly spinose and toothed leaf margin

of this species is characteristic among its congeners. All Philippine

records, thus far, are from Luzon island.

*Racomitrium lanuginosum [EHM 90]. Growing in open heath vegetation

at the summit, the species is identified by its whitish and strongly

erose leaf apices. Mt. Halcon is the second locality in the Philippines

for this nearly cosmopolitan moss. Its first Philippine record came

from Mt. Giting-Giting in Sibuyan Island in central Philippines (Tan,

1993):

** Rhaccocarpus alpinus (Wright) Par. [EHM 72]. This is the northernmost

locality of this species in the old world tropics. The closest population

to Mt. Halcon is on Mt. Kinabalu in NE Borneo. Plants are abundant

on Mt. Halcon forming extensive carpets under cushion-forming

seed plants. Its presence at high mountain peaks in Palawan should

be sought for.

Schlotheimia wallisii C. Muell. [EHM 91]. Epiphytic on trunk bases and

prostrate branches, this Malesian endemic is known from Borneo,

the Philippines and New Guinea (Vitt ef al., 1993). The other

Philippine localities include Luzon, Negros and Mindanao, but not

Palawan.

Sphagnum junghuhnianum Dozy & Molk. [EHM 88]. Plants are abundant

on wet soil under the thick growth of Miscanthus grasses. It is the

most common species of Sphagnum in Philippine mountains.

S. sericeum C. Muell. [EHM 87]. This is the other species of Sphagnum

found on wet soil beneath heath vegetation on Mt. Halcon. In the

Philippines, this species is known also from Negros and Mindanao.

Acknowledgements

We are grateful to Dr. R. Seppelt for his help in making available the

article on a revision of Rhacocarpus by J.-P. Frahm. We are equally thankful

to Dr. N. Klazenga for his critical review of the manuscript.

New moss records from Mindoro, the Philippines 521

References

Fernandez, J.C. 1982. Geology and Mineral Resources of the Philippines.

Vol. One, Geology. Bureau of Mines and Geo-Sciences, Ministry of

Natural Resources, Manila, the Philippines.

Frahm, J.-P. 1996. Revision der Gattung Rhacocarpus Lindb. (Musci).

Cryptogamie, Bryologique et Lichénologique. 17: 39—6S.

Hall, R. 1996. Reconstructing Cenozoic SE Asia. In: R. Hall & D. Blundell

(eds.) Tectonic Evolution of Southeast Asia. The Geological Society,

London, U.K. pp. 153—184.

Hall, R. 1998. The plate tectonics of Cenozoic SE Asia and the distribution

of land and sea. In: R. Hall and J.D. Holloway (eds.). Biogeography

and Geological Evolution of SE Asia. Backhuys Publishers, Leiden,

Netherlands. pp. 99—131.

Hamilton, W. 1981. Tectonics of the Indonesian Region (second printing).

Geological Survey Professional Paper 1078. United States Government

Printing Office, Washington DC. U.S.A.

Holloway, N.H. 1982. North Palawan Block, Philippines — its relation to

Asian mainland and role in the evolution of South China Sea. American

Association of Petroleum Geologists Bulletin. 66: 1355—1383.

JICA-Metal Mining Agency of Japan (MMAJ). 1984. Report on the

Geological Survey of Mindoro Island. Philippine Bureau of Mines,

Manila, the Philippines.

Klazenga, N. 1999. A revision of the Malesian species of Dicranoloma

(Dicranaceae, Musci). Journal of Hattori Botanical Laboratory. 87. \—

130.

Koponen, T. and D. H. Norris. 1986. Bryophyte flora of the Huon Peninsula,

Papua New Guinea. XVII. Grimmiaceae, Racopilaceae and

Hedwigiaceae (Musci). Acta Botanical Fennica. 133: 81—106.

Mandia, E.H. 1998. The Vegetation on the Northeastern Summit Zone of

Mt. Halcon, Mindoro Island, Philippines. PhD Dissertation. University

of the Philippines at Los Banos, Laguna. 210 pp.

Merrill, E.D. 1928. Flora of the Philippines. In: R-E. Dickerson. Distribution

of Life in the Philippines. Bureau of Printing, Manila, the Philippines.

pp. 130—167.

322 Gard. Bull. Singapore 53 (2001)

Tan, B.C. 1983. The status of Campylopus hemitrichius (C. Muell.) Jaeg.

Cryptogamie, Bryologique et Lichénologique. 4: 357—361.

Tan, B.C. 1993. Noteworthy range extension of Malesian mosses. Journal

of Hattori Botanical Laboratory. 74: 227—233.

Tan, B.C. 1994. The bryophytes of Sabah (North Borneo) with special

reference to the BRYOTROP transect of Mount Kinabalu. XIX. The

genus Acroporium (Semaophyllaceae, Musci) in Borneo, with notes on

species of Java and the Philippines. Willdenowia. 24: 255—294.

Tan, B.C. 1996. Biogeography of Palawan mosses. Australian Systematic

Botany. 9: 193—203.

Tan, B.C. & Z. Iwatsuki. 1991. A new annotated Philippine moss checklist.

Harvard Papers in Botany. 3: 1—64.

Tan, B.C., Z.-H. Li and P.-C. Lin. 1988. The Hainan-Mindoro connection,

an obscure pathway for plant migration in Southeast Asia. Natural

History Bulletin of Siam Society. 36: 7—15.

Vitt, D.H., T. Koponen and D.H. Norris. 1993. Bryophyte flora of the

Huon Peninsula, Papua New Guinea. LIII. Ulota and Schlotheimia

(Orthotrichaceae, Musci). Acta Botanica Fennica. 148: 5—25.

Virtanen, V. 1997. The genus Breutelia (Bartramiaceae, Musci) in Southeast

Asia and Oceania. Bryologist. 100: 324-348.

Yamaguchi, T. 1993. A revision of the genus Leucobryum (Musci) in Asia.

Journal of Hattori Botanical Laboratory. 73: 1—123.

Gardens’ Bulletin Singapore 53 (2001) 323-325.

New Combinations in Malesian Myricaceae

I.M. TURNER

Singapore Botanic Gardens, 1 Cluny Road, Singapore 259569

Abstract

In line with recent opinion favouring the splitting of the genus Myrica L. sensu lato, new

combinations in Morella Lour. (Morella esculenta (Buch.-Ham.) I.M. Turner and M. javanica

(Blume) I.M. Turner) are provided for the two Malesian species of Myricaceae.

Introduction

The genus Myrica L. as traditionally circumscribed consists of some 40—

50 extant species of trees and shrubs ranging from the arctic to the tropics.

The generally accepted type of the genus, circumboreal Myrica gale L.,

and the Californian endemic Myrica hartwegii S. Watson differ significantly

from the rest of the species. This is manifested principally in having

inflorescences inserted at the ends of the previous year’s growth rather

than on the old wood, and smooth fruits with a pair of adnate bracteoles

rather than papillate fruits lacking adnate bracteoles (MacDonald 1989,

Wilbur 1994). Chromosome numbers and essential oil chemistry also

distinguish the groups (MacDonald 1989). This division has often been

recognised at infrageneric rank (e.g. Kubitzki 1993) but, despite long-

standing reluctance by botanists to split Myrica, the tide seems to have

turned to favour the recognition of at least two genera. Unfortunately as

Myrica is typified by Myrica gale L. the generic name has to remain with

the numerically minor element and the many warm temperate and tropical

species have to be placed in a different genus. A proposal to conserve

Myrica with a conserved type (Verdcourt and Polhill 1997) was made in

order to reduce the number of necessary name changes. This was rejected

(Brummitt 1999, p. 367) largely on the grounds that, as the split was

unsettling to the nomenclature of important species whichever element

was maintained as Myrica, accepted typification and nomenclatural priority

should be upheld.

The next oldest generic name available for the tropical segregate is

Morella Lour., typified by Morella rubra Lour., better known as Myrica

rubra (Lour.) Sieb. & Zucc., a relatively important fruit tree in China and

Japan. New combinations in Morella have already been published for species

from North American (Wilbur 1994) and Africa (Killick, Polhill and

324 Gard. Bull. Singapore 53 (2001)

Verdcourt 1998). Even if the proposal to conserve Myrica had been accepted

it might not have prevented the resurrection of Morella because the

proposed conserved type was to be Myrica cerifera L.. This is one of the

waxy-fruited North American species that are possibly generically distinct

from the fleshy-fruited African and Asian species.

In Malesia, two species of Myrica have long been recognised: Myrica

esculenta Buch.-Ham. and Myrica javanica Blume. See Backer (1951) and

Noorsiha (1996) for extensive synonymies, description, illustrations and

notes on ecology and uses. Both species are widespread, common and

ecologically important, as well as possessing traditional utility. Both are

clearly best placed in Morella rather than Myrica sensu stricto and new

combinations effecting this change are made below:

Morella esculenta (Buch.-Ham.) I.M. Turner, comb. nov.

Basionym: Myrica esculenta Buch.-Ham. in D. Don, Prodr. Fl. Nep. (1825)

56.

Type: Nepal, Hamilton s.n. (K).

Distribution: India and Nepal, China, Indo-China, Malesia (excluding New

Guinea).

Morella javanica (Blume) I.M. Turner, comb. nov.

Basionym: Myrica javanica Blume, Bijdr. Fl. Ned. Ind. (1825) 517.

Type: Java, Mt. Gede, Reinwardt s.n. (L).

Distribution: Malesia except Malay Peninsula.

References

Backer, C.A. 1951. Myricaceae. Flora Malesiana, series I. 4: 277—279.

Brummitt, R.K. 1999. Report of the committee for Spermatophyta: 48.

Taxon. 48: 359—371.

Killick, D.J.B., R.M. Polhil! and B. Verdcourt. 1998. New combinations in

African Myricaceae. Kew Bulletin. 53: 993—995.

Kubitzki, K. 1993. Myricaceae. In: K. Kubitzki, J.G. Rohwer & V. Bittrich

(eds.) The Families and Genera of Vascular Plants. Vol. II Flowering

Plants - Dicotyledones. Magnoliid, Hamamelid and Caryophyllid Families.

Springer Verlag, Berlin, Germany. Pp. 453—457.

New combinations in Malesian Myricaceae 525

MacDonald, A.D. 1989. The morphology and relationships of the

Myricaceae. In: P.R. Crane & S. Blackmore (eds.) Evolution, Systematics,

and Fossil History of the Hamamelidae, Volume 2: ‘Higher Hamamelidae’.

Clarendon Press, Oxford, U.K. Pp. 147—165.

Noorsiha, A. 1996. Myricaceae. Tree Flora of Sabah and Sarawak. 2: 245—

249.

Verdcourt, B., and R. Polhill. 1997. (1291-1292) Proposals to conserve the

names Myrica and Gale (Myricaceae) with conserved types. Taxon. 46:

347—348.

“Wilbur, R.L. 1994. The Myricaceae of the United States and Canada: genera,

subgenera, and series. Sida. 16: 93—107.

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Gardens’ Bulletin Singapore 53 (2001) 327-341.

Sectional Placement of Three Bornean Species of Musa

(Musaceae) based on Amplified Fragment Length

Polymorphism (AFLP)

CAROL WONG!, RUTH KIEW?, ANTHONY LAMB’, OHN SET”,

SING KONG LEE', LEONG HUAT GAN! AND YIK YUEN GAN"

' Natural Sciences Academic Group, National Institute of Education, Nanyang

Technological University, 1 Nanyang Walk, Singapore 637616

* Singapore Botanic Gardens, 1 Cluny Road, Singapore 259569

> Agricultural Park, Tenom, Sabah, Malaysia

Abstract

The traditional approach to the classification of Musa species (Musaceae) is the separation

into four sections (Musa, Rhodochlamys, Callimusa and Australimusa) based on chromosome

number and morphological characters. The sectional placing of Musa beccarii N.W.

Simmonds is still unresolved due to its unique chromosome number. The sections of two

new species from Sabah, Malaysia, M. monticola [Hotta ex] Argent and M. suratii Argent,

were also undetermined. This study employs Amplified Fragment Length Polymorphism

(AFLP) as a molecular tool to determine the sectional placement of these three species

within Musa. Eight primer combinations generated 17 genetic markers, which confirmed

M. monticola and M. suratii as distinct species. AFLP data support the inclusion of M. beccarii

and M. monticola in sect. Australimusa, while results show M. suratii falling between

sect. Callimusa and sect. Australimusa suggesting that the two sections can no longer be

maintained as distinct.

Introduction

Banana, Musa L. (Musaceae), is an important perennial crop cultivated in

the tropics for food, fibre and as ornamental plants. Plagued by a complex

genetic system of sterility, interspecific hybrid constitutions, heterozygosity

and polyploidy (Novak, 1992), taxonomic classification of Musa is still in

need of improvement. While the genetic diversity and classification of

cultivated edible Musa are extensively studied (Simmonds and Shepherd,

1955; Bhat et al., 1992; Kaemmer et al., 1992), those of wild Musa are less

emphasised.

The traditional approach to wild Musa classification is the separation

of Musa into four sections (Musa Baker, Rhodochlamys Sagot, Callimusa

For correspondence: Associate Professor Gan Yik Yuen

328 Gard. Bull. Singapore 53 (2001)

Cheesman and Australimusa Cheesman) based on chromosome number

and morphological characters (Cheesman, 1947; Simmonds, 1962). A fifth

section, sect. Ingentimusa Argent, was recognised by Argent (1976) to

include Musa ingens N.W. Simmonds from New Guinea. Section Musa is

the largest and most diversified section ranging from South India to Japan

and Samoa. Most edible banana cultivars, including their progenitors

M. acuminata Colla and M. balbisiana Colla, belong to this section. Derived

from wild species in sect. Australimusa are the Fe’i banana cultivars mainly

distributed in New Guinea. The popular Filipino abaca originates from M.

textilis Née, also of this section. Section Callimusa is distributed from

Indochina to Indonesia and consists mainly of ornamental species, the best

known being M. coccinea Andr. Section Rhodochlamys, ranging from India

to Indochina, also consists of ornamental species, best represented by

M. ornata Roxb..

However, for some species it is not clear to which section they belong.

The sectional placing of M. beccarii N.W. Simmonds, M. monticola [Hotta

ex] Argent and M. suratii Argent are yet to be determined.

Simmonds (1960) described M. beccarii from Sabah, Malaysia. He

noted it superficially resembled M. coccinea (sect. Callimusa) but is distinct

from this species in the bracts, male buds, fruits and seeds, which are more

similar to species from sect. Rhodochlamys. Its chromosome number, n = 9

(Shepherd, 1959), is unlike any other Musa species: n = 10 in sect. Callimusa

and sect. Australimusa; n = 11 in sect. Musa and sect. Rhodochlamys; and

n = 7 in sect. Ingentimusa, although the chromosome number n = 10 has

also been reported for M. beccarii (Shepherd, 1959). Hence, the formal

position of M. beccarii within Musa is still undetermined.

Musa monticola, found in montane regions of Sabah (Mt. Kinabalu

and the Crocker Range) is a new species of undetermined section (Argent,

2000). Argent eliminated the possibility of it belonging to sect. Callimusa

and sect. Rhodochlamys on the grounds of it being distinct in seed structure

and anatomy, and the male axis habit, respectively. Instead, the polished

bracts and absence of wax suggested placement in sect. Australimusa (Hotta,

1987). Recent cytological examination of the species shows its chromosome

number to be n = 10 (Jong and Argent, 2001), which supports its placement

in sect. Australimusa.

Another new species described by Argent (2000) is M. suratii, from

Sabah and Sarawak, Malaysia. This species is morphologically distinct within

Musa in possessing minute seeds, giving no clue as to its formal placement

within Musa. Jong and Argent (2001) determined its chromosome number

as n = 10.

Amplified Fragment Length Polymorphism (AFLP) is a reliable and

robust fingerprinting technique widely used in genetic diversity studies of

Sectional placement of Bornean Musa species 329

plants and animals (Vos ef al., 1995). Applications of AFLP include inferring

phylogenetic relationships (Aggarwal et al., 1999), analysing genetic

diversities of populations and cultivars (Paul er al., 1997; Loh et al., 1999),

evaluating gene flow and dispersal (Travis et al., 1996), introgression (Tohme

et al., 1996) and hybridisation (Beismann er al., 1997). AFLP has the

advantage over other molecular techniques, such as RFLP, RAPD and

microsatellites, in that it is highly reproducible, requiring no prior knowledge

of genome sequence, relatively fast and easy to use, and in being able to

generate multiple loci per assay. In 1999, Crouch er al. reported

polymorphism in Musa generated by AFLP markers to be as high as that

obtained using microsatellites, contrary to previous reports utilising soybean

~ (Powell er al., 1996). Consequently, Crouch etal. (1999) concluded that

AFLP is most effective and suitable for genetic diversity studies of Musa.

Hence, this study aims to resolve the sectional placement of

M. beccarii, M. monticola and M. suratii within Musa using AFLP. (Section

Ingentimusa could not be included in this analysis due to lack of material).

Materials and Methods

Plant materials

This study employed a total of fifteen Musa species with sample sizes

ranging from one to five (Table 1). The materials included representatives

from four sections of Musa of both wild and cultivated origin, and from a

variety of introductions (material of the fifth section, sect. Jngentimusa,

was not available). The samples were collected from wild populations, and

plants grown in the Singapore Botanic Gardens (Singapore), the Royal

Botanic Garden Edinburgh (UK) and the Agricultural Park at Tenom

(Sabah, Malaysia). Voucher specimens are deposited in the herbaria at

Singapore Botanic Gardens and Royal Botanic Garden Edinburgh.

Leaf tissue was used for AFLP analysis. The leaves were surface

sterilised using a procedure from Zhang et al. (1997). The leaves collected

were swirled in 95% ethanol for 1 min, 5% bleach (NaOCl) for 5 min, and

then re-immersed in fresh 95% ethanol for 30 sec, after which they were

blotted dry and stored in sealed plastic bags at —80°C until needed for

DNA extraction.

DNA extraction

Plant DNA was extracted using the CTAB method according to Reichardt

and Rogers (1993). Briefly, leaf tissue was pulverised using liquid nitrogen

prior to the addition of 4 ml of Solution I (2% w/v CTAB (Sigma), 100

mM Tris-HCl, 20 mM EDTA, 1.4 M NaCl, pH 8.0) per gram of leaf tissue

Gard. Bull. Singapore 53 (2001)

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Sectional placement of Bornean Musa species 331

and incubated for 60 min at 65°C. The homogenate was then extracted

with an equal volume of chloroform/isoamy] alcohol (24:1) and centrifuged

at 12,000 rpm for 5 min. The upper aqueous phase was recovered and

incubated with 1/10 volume of Solution II (10% w/v CTAB, 0.7 M NaCl),

prewarmed to 65°C. The aqueous phase was then extracted with 1 volume

of chloroform/isoamyl alcohol (24:1) and recovered as before. To the

recovered aqueous phase, 1 volume of Solution III (1% w/v CTAB, 50

mM Tris-HCl, 10 mM EDTA, pH 8.0) was added and incubated overnight

at 37°C. The mixture was then centrifuged for 5 min at 3,500 rpm and

supernatant removed. The DNA pellet was then re-dissolved in Solution

IV (10 mM Tris-HCl, 0.1 mM EDTA, 1 M NaCl, pH 8.0) at 0.5 to 1 ml per

' gram starting material, followed by ethanol precipitation of the DNA. It

was then washed with 70% ethanol, dried and re-suspended in minimal

volume of TE buffer at 0.1 to 0.5 ml per gram starting material.

AFLP analysis

AFLP analysis was carried out according to Vos et al. (1995) with minor

modifications. Restriction digests of genomic DNA with EcoRI and Msel

were carried out at 37°C for 1 h. Following heat inactivation of the

restriction endonucleases, the genomic DNA fragments were ligated to

EcoRI and Msel adapters overnight at 16°C to generate template DNA

for amplification. PCR was performed in two consecutive reactions. The

template DNA generated was first pre-amplified using AFLP primers each

having one selective nucleotide. The PCR products of the pre-amplification

reaction were then used as template after 5-fold dilution in sterile water.

for selective amplification using two AFLP primers, each containing three

selective nucleotides. A total of eight primer combinations were used in

this study (Table 2). The final PCR products were run on a 6% denaturing

polyacrylamide gel in 1X TBE buffer. The EcoRI primers used were not

radioactively labelled as in the original protocol. Instead, a modified silver

staining method was used (Loh et al., 1999).

Data analysis

For the diversity analysis, bands were scored as present (1) or absent (0) to

form a raw data matrix. A square symmetric matrix of similarity was then

obtained using Jaccard’s Similarity Coefficient [x / (y-z)]. where x is the

number of fragments in common between two taxa, y is the total number

of fragments scored, z is the number of fragments absent in both taxa,

from the raw data matrix. Genetic diversity estimates (GDEs) were then

calculated as 1 minus Jaccard’s Similarity Coefficient and used for cluster

analysis using the Unweighted Pair Group Method with Arithmetic mean

332 Gard. Bull. Singapore 53 (2001)

(UPGMA) technique of the NEIGHBOR program in PHYLIP version

3.5c (Felsenstein, 1993).

Table 2. Sequences of the primers and adapters used for AFLP analysis

Name / Abbreviation Enzyme Type Sequence (5’-3’)

GYY 101/ EA+ EcoRI Adapter + CTCGTAGACTGCGTACC

GYY 102/ EA- EcoRI Adapter - AATTGGTACGCAGTCTAC

GYY 103/ MA+ Msel Adapter + GACGATGAGTCCTGAG

GYY 104/ MA- Msel Adapter - TACTCAGGACTCAT

GYY 105/ E-A EcoRI Primer +1 GACTGCGTACCAATTCA

GYY 107/ E-AAC EcoRI Primer +3 GACTGCGTACCAATTCAAC

GYY 108/ E-AAG EcoRI Primer +3 GACTGCGTACCAATTCAAG

GYY 109/ E-ACA EcoRI Primer +3 GACTGCGTACCAATTCACA

GYY 110/ E-ACT EcoRI Primer +3 GACTGCGTACCAATTCACT

GYY 111/ E-ACC EcoRI Primer +3 GACTGCGTACCAATTCACC

GYY 112/ E-ACG EcoRI Primer +3 GACTGCGTACCAATTCACG

GYY 113/ E-AGC EcoRI Primer +3 GACTGCGTACCAATTCAGC

GYY 114/ E-AGG EcoRI Primer +3 GACTGCGTACCAATTCAGG

GYY 106/ M-C Msel Primer +1 GATGAGTCCTGAGTAAC

GYY 115/ M-CAA Msel Primer +3 GATGAGTCCTGAGTAACAA

GYY 116/ M-CAC Msel Primer +3 GATGAGTCCTGAGTAACAC

GYY 117/ M-CAG Msel Primer +3 GATGAGTCCTGAGTAACAG

GYY 118/ M-CAT Msel Primer +3 GATGAGTCCTGAGTAACAT

GYY 119/ M-CTA Msel Primer +3 GATGAGTCCTGAGTAACTA

GYY 120/ M-CTC Msel Primer +3 GATGAGTCCTGAGTAACTC

GYY 121/ M-CTG Msel Primer +3 GATGAGTCCTGAGTAACTG

GYY 122/ M-CTT Msel Primer +3 GATGAGTCCTGAGTAACTT

Results

Identification of species using AFLP markers

AFLP fingerprinting of 15 species of Musa with eight primer combinations

(Table 2) yielded a total of 276 unambiguous bands of size 50—500 base

pairs. Of these, 262 (95%) are polymorphic across all samples, while the

number of polymorphic loci generated per primer combination is 33. Figure

1 illustrates an AFLP profile generated using primer pair 1 (E-AAC, M-

CAA).

A total of 17 genetic markers unique to each species were observed

for nine species of Musa (Table 3), including M. monticola and M. suratii.

No unique bands distinguished M. beccarii from the rest of the Musa species.

M. suratii is distinct due to two unique bands, while another unique marker

Sectional placement of Bornean Musa species 333

23°45 6 7.8 910 1112131

Figure 1. AFLP profile obtained with primer pair 1 (E-AAC, M-CAA) illustrating banding

patterns for some species examined in this study. Lane 1: M. acuminata ssp. truncata, Lane 2:

M. balbisiana, Lane 3: M. textilis, Lane 4: M. violascens, Lane 5: M. ornata, Lane 6: M. coccinea,

Lane 7: M. nagensium, Lane 8: M. campestris, Lane 9: M. velutina, Lane 10: M. jackeyi, Lane

11: M. beccarii, Lane 12: M. suratii, Lane 13: M. monticola, Lane 14: M. borneensis, Lane M:

pUC19/HpalI molecular weight marker.

334 Gard. Bull. Singapore 53 (2001)

Table 3. Genetic markers observed for nine species of Musa

Primer Pair EcoRI Msel 2 Total

S 3 a Ee a 8 number of

S = £ a ~ > ‘

~ So Vea ees ae Sie unique

S $ 5 § 5 z : = K S markers

CSR eee Se OS ‘SS per primer

Sf 2) SSeS soso. i

1 AAC* CAA** : = — er ae i : 0

10 AAG CAC = 4 5 Bela oe : 1

19 ACA CAG - - = = a 2 = = ES D)

28 ACE ‘CAT = = ae | sake , 1

37 ACG CTA 1 = Sen Oe en oe . 3

46 ACT Gre = : 1) ial ees ee 1 3

55 INGE (GING - 1 {| ~ 2 a oe. 1 6

64 AGG (Gil = E 62k ey r 1

Total 1 My, “Zhe Si) Ae ete hae 17

*EcoRI : EcoRI-adapter based primer; the selective nucleotides added at the 3’ end are

indicated

**Msel : Msel-adapter based primer; the selective nucleotides added at the 3’ end are

indicated

was observed for M. monticola. The unique bands support the distinctiveness

of both M. suratii and M. monticola as separate species. The number of

unique bands observed for the remaining species of Musa ranged from one

to four.

Genetic relationships between species of Musa

The cluster analysis using values of GDEs (Table 4) generated a phenogram

(Fig. 2) depicting genetic relationships between species of Musa. Two main

clusters were observed, each made up of the traditional sectional alliance

of Musa-Rhodochlamys and Callimusa-Australimusa respectively. The

cluster of the Musa-Rhodochlamys alliance revealed both subspecies of M.

acuminata, ssp. truncata and ssp. malaccensis, to be closely related as

anticipated. M. velutina H. Wendl. & Drude and M. ornata of sect.

Rhodochlamys clustered closely with M. acuminata, while M. nagensium

Prain and M. balbisiana in sect. Musa formed the remaining members of

this cluster.

The cluster of the Callimusa-Australimusa alliance showed M.

violascens Ridl., M. borneensis Becc. and M. coccinea, which all belong to

sect. Callimusa, separated from the other species in this cluster. The cluster

was further divided into a sub-cluster consisting of M. textilis, M. beccarii,

355

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Sot ot) eee eee oe Pe te PO i Rom a eer eee

ee a ee ee ee oe ee Gee

a 8 > = = 8 = 5 ® 9 > =. a Ae 8S §

cise Ge oe fees ce Gi 2 kee Se eae. =.

. ace See, aie Es 4s s @8s &

Sg ee S| 5 5 Sy S

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suoneurquios s9uutid yYysI9 Jo (S|qH) soyewnso APISIOAIP SOUNS URI “p AGEL

(nov “PW OF poyerasiqgqe si vipulUNgD “P)

oS)

Gard. Bull. Singapore 53 (2001)

Musa velutina

Musa ornata

M. acuminata ssp. truncata

M. acuminata ssp. malaccensis

Musa nagensium

Musa balbisiana

Musa jackeyi

Musa textilis

Musa beccarii

Musa monticola

Musa suratii

Musa campestris

Musa violascens

Musa borneensis

Musa coccinea

0.26 0.13 0

Figure 2. Genetic relationships of 15 species of Musa by UPGMA cluster analysis. Scale

depicts genetic diversity estimates (GDEs).

M. monticola, M. suratii and M. campestris Becc., while M. jackeyi Hill in

sect. Australimusa was distant from this subcluster.

Based on the phenogram, M. beccarii is closest to M. textilis of

sect. Australimusa. GDE values between M. beccarii and both species of

sect. Australimusa examined ranged from 0.249 (between M. beccarii and

M. textilis) to 0.391 (between M. beccarii and M. jackeyi), as compared

with species of sect. Callimusa where values ranged from 0.336 (between

M. beccarii and M. campestris) to 0.446 (between M. beccarii and

M. violascens). These data suggest M. beccarii to be closer genetically to

species of sect. Australimusa than to those of sect. Callimusa.

Musa monticola showed a similar pattern in that it was also genetically

most similar to M. textilis. GDE values between M. monticola and species

Sectional placement of Bornean Musa species 337

of sect. Australimusa ranged from 0.276 (between M. monticola and M.

textilis) to 0.462 (between M. monticola and M. jackeyi), while values for

species of sect. Callimusa ranged from 0.410 (between M. monticola and

M. campestris) to 0.491 (between M. monticola and M. coccinea). This,

too, suggests a higher similarity between M. monticola and members of

sect. Australimusa than between M. monticola and members of sect.

Callimusa.

Musa suratii clustered closely with M. campestris, which belongs to

sect. Callimusa and had GDE values ranging from 0.357 (between M. suratii

and M. campestris) to 0.526 (between M. suratii and M. borneensis) when

compared with species in sect. Callimusa. On the other hand, species of

- sect. Australimusa possessed GDE values of between 0.358 (between

M. suratii and M. textilis) to 0.405 (between M. suratii and M. jackeyi). GDEs

between M. suratii and M. textilis, and M. suratii and M. campestris are

highly similar, giving no indication of whether M. suratii is genetically closer

to members of sect. Callimusa or sect. Australimusa.

Discussion

AFLP is highly informative and reliable in providing valuable insight into

genetic similarities between species. In addition, it is useful in generating

unique molecular markers for identification purposes and the development

of these is important in the improvement of banana breeding programmes.

In assessing unique bands, results confirm both M. monticola and M. suratii

to be distinct species as described by Argent (2000).

The separation of the Rhodochlamys-Musa and Callimusa-

Australimusa taxa in this molecular result is in agreement with previous

morphological data and supports the traditional chromosomal grouping of

Musa species into the n = 11 group and n = 10 group respectively.

Musa beccarii falls within a cluster comprising M. textilis and

M. monticola. Simmonds and Weatherup (1990) suggested that M. beccarii

belonged in sect. Callimusa based on morphological characters. Our results

disagree with that placement as genetically its alliance with

sect. Australimusa is much stronger.

In addition, the chromosome numbers reported for M. beccarii are

n = 9 and n = 10, this latter number being due to multivalent formation in

meiosis (Shepherd, 1959) and which supports the inclusion of M. beccarii

in sect. Australimusa. The subglobose seed of M. beccarii resembles that of

sect. Australimusa and is unlike the cylindrical seed found in sect. Callimusa.

Based on its seed structure, chromosome number and the AFLP results

obtained, M. beccarii is confidently placed within sect. Australimusa.

338 Gard. Bull. Singapore 53 (2001)

Argent (2000) suggested the inclusion of M. monticola in

sect. Australimusa based on morphological characters. AFLP results are in

agreement with this placement, as M. monticola clusters closely with

M. textilis and M. beccarii. Values of GDEs support the inclusion of

M. monticola in sect. Australimusa, rather than in sect. Callimusa. Although

M. monticola is distinct from M. textilis morphologically, e.g. in pseudostem

height, the length of the male peduncle, imbrication of the male bud, and

characteristics of the basal flower and seeds (Argent, 2000), they are

genetically the most closely related.

Interestingly, M. suratii clustered closely with M. campestris of sect.

Callimusa, but GDEs indicate high genetic similarity with M. textilis

suggesting M. suratii to be intermediate between sect. Callimusa and sect.

Australimusa. Argent (2000) noted that its unique tiny seeds provide no

clue as to its sectional alliance. Jong and Argent (2001) determined its

chromosome number as n = 10, which further supports its position within

the Callimusa-Australimusa group. AFLP results show M. suratii falling

between sect. Callimusa and sect. Australimusa suggesting that the two

sections can no longer be maintained as distinct.

Conclusion

AFLP has shown M. monticola and M. suratii to be distinct species as

described by Argent (2000). In addition, results unambiguously show that

these three species fall within the Callimusa-Australimusa alliance of Musa,

thus disproving any notion of them belonging to the Musa-Rhodochlamys

alliance. Their chromosome numbers also support this placement.

M. beccarii and M. monticola possess high genetic similarities with M. textilis

in sect. Australimusa, while M. suratii is intermediate between M. campestris

in sect. Callimusa and M. textilis in sect. Australimusa.

UPGMA cluster analysis illustrates why the problem has arisen as

the three species in question, M. beccarii, M. monticola and M. suratii, in

fact cluster between taxa in sect. Australimusa (with M. jackeyi being most

distant) and taxa in sect. Callimusa (with M. coccinea, M. borneensis and

M. violascens being most distant) and are intermediate between

M. campestris in sect. Callimusa and M. textilis in sect. Australimusa. While

M. beccarii is definitely closer to M. textilis and can therefore be said to fall

within sect. Australimusa, M. monticola and M. suratii are intermediate

and could equally belong to either section. The single character that

separates the two sections is the seed type and, while that of M. monticola

is more like that of sect. Australimusa, that of M. suratii is unique and

conforms to neither section. This calls into question the validity of keeping

Sectional placement of Bornean Musa species 339

sections Australimusa and Callimusa distinct. This problem is dealt with in

a later study.

Acknowledgements

This research was funded by the Academic Research Fund, National

Institute of Education, Nanyang Technological University, Singapore, RP

12/98/GYY. We thank the Director of Singapore Botanic Gardens for

permission to collect leaf samples, George Argent (Royal Botanic Garden

Edinburgh, UK) for providing plant materials and useful comments on the

manuscript and Kwiton Jong (University of Aberdeen, Scotland, UK) for

making available his data on cytology.

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New Combinations, New Names and New Species of

Madhuca (Sapotaceae) from Sabah and Sarawak, Borneo

P.C..YIT AND PAUL, PK... CHAI

‘/, Sarawak Forestry Department

Kuching, Malaysia

Abstract

Five new species of Madhuca, M. engkikiana, M. markleeana, M. multinervia, M. ochracea,

' and M. silamensis, are described and two new combinations, M. daemonica and M. prolixa,

and two new names, M. cheongiana and M. kuchingensis, are proposed as a consequence of

the reduction of Ganua to Madhuca.

Introduction

In revising the family Sapotaceae for the Tree Flora of Sabah and Sarawak,

we accepted Pennington’s reduction (1991) of Ganua Dubard (1908) to

Madhuca Buch.—Ham. ex J.F. Gmelin (1791). The enlarged genus Madhuca

comprises 47 species and one variety in Sabah and Sarawak out of a total

of 50 species found in Borneo.

The Sabah and Sarawak account includes two new combinations (M.

daemonica and M. prolixa), two new names (M. cheongiana and M.

kuchingensis), and five new species (M. engkikiana, M. markleeana, M.

multinervia, M. ochracea, and M. silamensis) here described.

New Combinations and New Names

1. Madhuca cheongiana Yii & P. Chai, nom. nov.

Synonyms: Ganua sarawakensis Pierre ex Dubard, Bull. Mus. Hist. Nat.

Paris 14 (1908) 409. Type: Beccari PB 3105, Borneo, Sarawak, Kuching

(holotype FI, n.v.); Van den Assem, Blumea 7 (1953) 375: Ganua attenuata

Griffioen & H.J. Lam, nom. ined., Anderson, A Checklist of the Trees of

Sarawak (1980) 314, nom. nud.

Notes: In transferring Ganua sarawakensis to the genus Madhuca, the species

name “sarawakensis” cannot be used because it is already preoccupied by

Madhuca sarawakensis (Pierre ex Dubard) H.J. Lam, Bull. Jard. Bot.

Buitenz., 3, 7 (1925) 180, based on Kakosmanthus sarawakensis Pierre ex

344 Gard. Bull. Singapore 53 (2001)

Dubard, Bull. Mus. Hist. Nat. Paris 14 (1908) 407, and typified by Beccari

PB 423, Borneo, Sarawak, Kuching (holotype P, n.v.; isotypes FI, L, n.v.).

The species is endemic to Borneo (Brunei Darussalam, Kalimantan,

Sabah, and Sarawak). In Sabah and Sarawak, it is uncommon and scattered

in a few localities (SAN 29561, SAN 33601, SAN 36601A, SAN 86191,

SAN 126691; S 2162,S 2222, S- 7559, S 13043, S 15437, S 29477, and S

32947), and occurs mainly in lowland kerangas and mixed dipterocarp

forests below 200 m altitude.

The species is named in honour of Mr. Cheong Ek Choon, the

Director of the Sarawak Forestry Department, for his interest and

continuous support in the study of the diversity, conservation and

sustainable management of forest tree resources in Sarawak.

2. Madhuca daemonica (Van den Assem) Yii & P. Chai, comb. nov.

Basionym: Ganua daemonica Van den Assem, Blumea 7 (1953) 394. Type:

Egar A 0932, Borneo, Sarawak, Setapok FR (holotype KEP!).

Notes: A species endemic to Borneo and found in a few localities in Sabah

(SAN 17437, SAN 17448), and more commonly in Sarawak (Egar A 0932,

S 2618, S 2722, S 2768, S 4423, S 4666, S 5866, S 7088, S 12393, S 12965, S

14462, S 30039, and S 36452). Its habitats include peatswamp, kerangas

and mixed dipterocarp forests at altitudes up to 700 m.

3. Madhuca kuchingensis Yii & P. Chai, nom. nov.

Based on Ganua beccarii Pierre ex Dubard, Bull. Mus. Hist. Nat. Paris 14

(1908) 408. Type: Beccari PB 2241, Borneo, Sarawak (holotype FI, n.v.).

Notes: In the genus Madhuca, there exists M. beccarii (Engl.) H.J. Lam,

Bull. Jard. Bot. Buitenz., 3, 7 (1925) 177, based on Payena beccarii Engl.,

Bot. Jahrb. 12 (1890) 508, and typified by Beccari PB 1598, Borneo,

Sarawak, Kuching (holotype FI, n.v.; isotypes K, P, n.v.). The new name

Madhuca kuchingensis Yii & P. Chai is, therefore, proposed.

The species is endemic to Borneo and confined to the central and

western parts of Sarawak (S 3356, S 4307, S 14846, S 24344, S 24514, §

24516, S 24545, S 24602, S 27064, S 32394, S 32472, S 37044, and S 37861),

and found mainly in lowland kerangas and mixed dipterocarp forests at

altitudes up to 100 m.

New combinations, names and species of Madhuca in Borneo 345

4. Madhuca prolixa (Pierre ex Dubard) Yii & P. Chai, comb. nov.

Basionym: Ganua prolixa Pierre ex Dubard, Bull. Mus. Hist. Nat. Paris 14

(1908) 409. Type: Beccari PB 2446, Borneo, Sarawak, Kuching (holotype

FI, n.v.; isotype L, n.v.).

Notes: A species occurring in Peninsular Malaysia and Borneo. Scattered

throughout Sarawak in lowland mixed dipterocarp forest at altitudes up to

200 m (S 4099, S 18771, S 25258, and S 29249). It also occurs in Brunei

(BRUN 2476 and BRUN 3317).

New Species

1. Madhuca engkikiana Yii & P. Chai, sp. nov.

(Engkik Soepadmo, Coordinator and Chief Editor of the Tree Flora of

Sabah & Sarawak project)

Madhucae kuchingensi arcte similis, sed in ramulis glabris foliis bene

dispositis, venatione intercostali reticulata supra non impressa, venis paucis

e costa enascentibus venis lateralibus parallelis differt. Typus: Kodoh Tarodop

SAN 83612, Borneo, Sabah, Sandakan, Telupid (holotypus SAR!; isotypi

AA, K, KEP!, L, SAN!, SING!).

Figure 1

Tree up to 25 m tall, 42 cm diameter. Bark reddish grey; inner bark yellowish

red. Sapwood yellow. Twigs slender, terete, glabrous. Terminal buds c. 3

mm long, puberulous. Stipules triangular, c. 1 x 0.4 mm. Leaves spirally

arranged, scattered and well-spaced along the twigs, coriaceous, glabrous

on both surfaces; blade elliptic to elliptic-obovate, 9-13 x 3.5-4.8 cm, base

cuneate, slightly decurrent and oblique, margin entire and plane, apex

short-acuminate; midrib raised on both surfaces; lateral veins 11—15 pairs,

ascending at an angle of c. 80° from the midrib, distinctly connected by the

thickened intercostal veins to form intramarginal vein-loops, prominent on

both surfaces; intercostal venation reticulate, with a few veins arising from

the midrib and parallel to the lateral veins; petioles 2—3.5 cm long, flat to

slightly raised on the adaxial side, thickened, black and puberulous at the

base. Inflorescences axillary, 6—8-flowered; pedicels up to 0.3 cm long,

yellowish appressed hairy. Flowers: calyx biseriate, consisting of two whorls

of two sepals; sepals suborbicular, c. 3 x 2.5 mm, tufted hairy at the apex;

corolla c. 2 mm long, tube c. 0.75 mm long, lobes 7, oblanceolate, densely

pubescent with tufted yellowish hairs at the apex; stamens 16, in two whorls,

346 Gard. Bull. Singapore 53 (2001)

Figure 1. Madhuca engkikiana. A, leafy flowering twig; B, flower bud; C, outer sepal; D, inner

sepal; E, longitudinal section of flower bud; F, petal; G, petals and stamens; H, stamens in

different views; I, gynoecium; J, fruit. (A—I from SAN 83612, J from SAN 131963.)

New combinations, names and species of Madhuca in Borneo 347

filaments very short, anthers lanceolate, c. 1 x 0.5 mm, slightly hairy; ovary

subconical, c. 2 mm long, hairy, 6—7-loculed, style stout, c. 1.5 mm long.

Fruits (young) ellipsoid, laterally compressed, 1.5 x 0.6-1.2 cm, densely

brownish tomentose. Seeds laterally compressed, pointed at both ends;

testa thin, smooth; scar linear.

Distribution: Endemic to Borneo. Known only from four collections from

Sabah (SAN 53982, SAN 54205, SAN 131963, and the type).

Ecology: In lowland mixed dipterocarp forest, up to 250 m altitude.

Notes: Similar to Madhuca kuchingensis but differs by its glabrous twigs,

leaves well-spaced along the twigs, and reticulate intercostal venation that

is not impressed above and with a few shorter veins arising from midrib

and parallel to lateral veins.

2. Madhuca markleeana Yii & P. Chai, sp. nov.

(Mark Lee Hua Seng, Deputy Director, Forestry Department, Sarawak)

Madhucae borneensi et M. sarawakensi in characteribus vegetativis similis,

sed in ramulis laminisque foliorum glabris, stipulis orbicularibus, sepalis

exterioribus magnis suborbicularibus et recurvatis, pedicello fructifero

longiore (7-8 cm longo) differt. Typus: Yii S 72728, Borneo, Sarawak,

Bukit Meluku (holotypus SAR!; isotypi K, KEP!, L, SAN!).

Figure 2

Tree up to 15 m tall, 25 cm diameter, with very low buttresses. Bark

chocolate brown with greyish green mottles, smooth to finely fissured;

inner bark c. 4 mm thick, dull orange, granular. Twigs terete, rusty-brown

velvety hairy at the tips, glabrescent. Terminal buds up to 8 mm long; bud-

scales elliptic, 10-14 x 9-11 mm. Stipules lanceolate, c. 10 x 3 mm, velvety

hairy, caducous. Leaves spirally arranged, scattered and well-spaced along

the twigs, coriaceous, glabrous on both surfaces; blade lanceolate or oblong,

25-35 x 7-8 cm, base broadly cuneate and slightly oblique, margin entire

and plane, apex obtuse or acuminate with a sharp tip; midrib broadly

crested above, prominent below; lateral veins 28-35 pairs, ascending at an

angle of 75-85° from the midrib, straight at first and then curving and

joining at their tips to form vein-loops rather far from the leaf margin,

impressed above, prominent below; intercostal venation slender, scalariform,

with at least one vein descending from the margin and parallel to the

lateral veins, faint above, distinct below; petioles 1.5-2.5 cm long, flat on

348 Gard. Bull. Singapore 53 (2001)

Figure 2. Madhuca markleeana. A, fruiting leafy twig; B, fruit; C, longitudinal section of fruit;

D, part of inflorescence; E, flower bud; F, petal; G, flower bud with sepals and a few petals

removed; H, longitudinal section of flower bud; I, petals and stamens; J, stamens in different

views; K, gynoecium. (A-C from S$ 72729, D-K from § 72728.)

New combinations, names and species of Madhuca in Borneo 349

the adaxial side, round on the abaxial side, velvety hairy, glabrescent.

Inflorescences axillary, 3-5-flowered; pedicels subangular, 5-6 cm long, pale

green, sparsely pubescent. Flowers: calyx biseriate, consisting of two whorls

of two sepals; sepals suborbicular, c. 10 x 14 mm, outer pair with recurved

margin, inner pair smaller, margin not recurved, pubescent; corolla white,

c. 12 mm long, lobes 8, lanceolate, 7-8 mm long, apex acute, tube c. 4 mm

long, slightly pubescent at the throat; stamens 19, in two whorls, filaments

subulate, c. 5 mm long, anthers sagittate, yellowish, c. 5 mm long; ovary

disciform, c. 3 x 5 mm, 8-loculed, pilose, style filiform, pale green, tapering

towards stigma. Fruits ellipsoid, c. 7 x 4 cm, 1-seeded, tapering at both

ends; pericarp thick and fleshy, green, densely brown tomentose; stalk 7-8

cm long. Seeds ellipsoid, 4.5 x 1.2—2 cm, pointed at both ends; testa thin;

scar narrowly linear, c. 43 x 3 mm.

Vernacular name: Sarawak—nyatoh gasing (Malay).

Distribution: Endemic to Sarawak; known only from two collections, the

type and Yii S 72729, Borneo, Sarawak, Bukit Meluku (K, KEP!, L, MO,

SAN!, SAR!, SING!).

Ecology: Understorey tree on steep slopes of lowland mixed dipterocarp

forest, up to 400 m altitude.

Notes: Resembles Madhuca borneensis and M. sarawakensis in vegetative

characters but differs in having glabrous twigs and leaves, orbicular stipules,

large suborbicular and recurved outer sepals, and longer fruit stalk (7-8

cm long).

3. Madhuca multinervia Yii & P. Chai, sp. nov.

(Latin, multi = many, nervis = nerves; alluding to the many, closely parallel

lateral veins of the leaves)

Madhucae elmeri arcte similis in characteribus vegetativis, sed in ramulis

novellis foliis petiolisque glabris, venis lateralibus 28-45 paribus arcte

approximatis ascendentibus distinguenda. Typus: Dewol SAN 97008, Borneo,

Sabah, Tongod district, Bt. Pantagaluang (holotypus SAR!; isotypi AA,

BO!, K, KEP!, L, OX, SAN!, SING!).

Figure 3

Tree up to 20 m tall, 40 cm diameter. Bark pale brown with greyish mottles,

smooth; inner bark pale reddish, with sticky white latex. Sapwood whitish.

350 Gard. Bull. Singapore 53 (2001)

Twigs slender, solid, 2-4 mm diameter, subangular with distinct stipular

scars, rusty-brown velvety hairy at the tips, quickly glabrescent and

becoming blackish. Terminal buds c. 5 mm long. Stipules broadly ovate, c.

6 x 4 mm, truncate, rusty-brown velvety. Leaves spirally arranged and

scattered along ends of twigs, papyraceous to subcoriaceous, glabrous or

with appressed silvery hairs on the midrib above, glabrous or with remnants

of appressed silvery hairs and subappressed ferrugineus hairs on the midrib

below; blade oblong-elliptic, 16.5—19 x 5-12 cm, base obliquely cuneate to

rounded, margin entire and plane, apex obtuse; midrib impressed and

slightly crested above, strongly prominent and rounded below; lateral veins

28-45 pairs, ascending at an angle of 65-85° from the midrib, straight or

slightly curved, arching and joining into vein-loops at 1-2 mm from the

leaf margin, impressed above, prominent below; intercostal venation

. scalariform, faint; petioles 2-4 cm long, thickened, narrowly grooved on

the adaxial side, rounded on the abaxial side, rugose and black at the base,

glabrous. Inflorescences axillary, 3—10-flowered; pedicels 1-2 cm long,

slender, velvety hairy. Flowers: calyx biseriate, consisting of two whorls of

two sepals; sepals free, imbricate, orbicular, c. 6 mm across, velvety hairy,

inner pair thinner, crested and hairy; corolla 7-10 mm long, 8-lobed, lobes

elliptic or ovate, c. 3.5 x 1.5 mm, apex acute, densely hairy at the throat;

stamens 16-24, in two or three whorls, filaments very short, anthers sagittate,

c. 3.5 mm long; ovary subconical, c. 1 mm across, 8-loculed, glabrous, style

c. 8 mm long, glabrous. Fruits ellipsoid, up to 2.1 x 1.2 cm, 1-seeded;

pericarp thin, glabrous; stalk up to 2 cm long. Seeds laterally compressed,

ellipsoid to obovoid, 1.7 x 1.1 x 0.8 cm, obtuse at both ends; scar as long as

seed, c. 5 mm wide.

Distribution: Endemic to Borneo. Scattered throughout the eastern parts

of Sabah (SAN 88312, SAN 93874, SAN 96918, SAN 97008, SAN 99692,

SAN 99726, SAN 111764, SAN 124566, and SAN 133478).

Ecology: Usually on hillsides and ridges in primary lowland mixed

dipterocarp forest.

Notes: Closely allied to Madhuca elmeri in vegetative characters but can be

distinguished by its glabrous young twigs, leaves and petioles, and 28-45

pairs of closely set ascending lateral veins.

4. Madhuca ochracea Yii & P. Chai, sp. nov.

(Latin, ochraceus = pale yellowish brown; referring to the indumentum)

New combinations, names and species of Madhuca in Borneo 351

Figure 3. Madhuca multinervia. A, flowering leafy twig: B, flower bud; C, longitudinal section

of flower bud: D, outer sepal: E, inner sepal: F, petals: G, part of flower with petals and stamens:

H. stamen; I, gynoecium. (All from SAN 97008.)

352 Gard. Bull. Singapore 53 (2001)

Madhucae dubardii similis in foliis infra persistentiter pallide flavido-brunneo

tomentosis, sed in folii margine recurvata, apice caudato, nervis intercostalibus

a margine descendentibus venis lateralibus parallellis differt. Typus: Wright

S 29130, Borneo, Sarawak, Niah National Park (holotypus SAR!; isotypi

K, L, SAN!, SING!).

Figure 4

Tree up to 25 m tall, 70 cm diameter. Bark greyish brown, scaly; inner bark

dull orange. Twigs angular, glabrous. Terminal buds c. 5 mm long. Stipules

broadly ovate, c. 5 x 4 mm, glabrous, persistent. Leaves spirally arranged

and crowded at the ends of twigs, coriaceous, glabrous above, covered

with persistent pale yellowish brown tomentum below; blade elliptic or

elliptic-obovate, 8.5—-19 x 3.5-8.2 cm, base cuneate, margin recurved, apex

caudate or acuminate, acumen c. | cm long; midrib shallowly grooved and

crested above, rounded and prominent below; lateral veins slender, 26-34

pairs, ascending at an angle of 80—85° from the midrib, curved, diminishing

and becoming inconspicuous toward the leaf margin, faint and impressed

above, distinct below; intercostal venation slender, descending from the

margin and parallel to the lateral veins, laxly reticulate towards the leaf

margin, indistinct above, faint below; petioles 2-4.8 cm long, narrowly

grooved on the adaxial side, thickened and rugose at the base. /nflorescences

axillary, 2—7-flowered; pedicels c. 1.2 cm long, glabrous, angular and enlarged

at the apex. Flowers: calyx biseriate, consisting of two whorls of two sepals;

sepals orbicular, c. 6 mm across, apex rounded, velvety hairy; corolla c. 8

mm long, 10-lobed, lobes obovate, c. 4 x 3 mm, apex rounded and ciliate;

stamens 16, in two whorls, filaments c. 2.5 mm long, anthers sagittate, c. 2.5

mm long; ovary subconical, c. 2 mm across, 6—8-loculed, velvety hairy, style

c. 7 mm long, subangular, glabrous. Fruits (immature) ovoid, c. 1 cm across,

base rounded, apex flattened and topped by stout remnant of style, pale

yellowish brown hairy. Seeds unknown.

Vernacular name: Sarawak—nyatoh kelabu (Iban).

Distribution: Endemic to Borneo. Known from only two collections from

Sarawak, the type and §$ 23/00 from the Mentagai Hills, Marudi, Miri

Division.

Ecology: In primary lowland mixed dipterocarp forest on clay loam soils,

up to 100 m altitude.

Notes: Similar to Madhuca dubardii in the persistently pale yellowish brown

tomentose leaf undersurface but differs by its recurved leaf margin, caudate

New combinations, names and species of Madhuca in Borneo 353

Figure 4. Madhuca ochracea. A, flowering leafy twig: B, flower bud; C, longitudinal section of

flower bud: D, outer sepal; E, inner sepal: F, petal: G, petal and stamens: H, stamen: I, gynoecium;

J, young fruit. (A-I from § 29/30, J from S 23100.)

354 Gard. Bull. Singapore 53 (2001)

leaf apex and intercostal veins descending from the margin and parallel to

the lateral veins.

5. Madhuca silamensis Yii & P. Chai, sp. nov.

(of Mt. Silam, Sabah)

Ab ullis speciebus borneensibus cognitis in foliis crasse coriaceis argenteo-

brunneo tomentosis secus ramos crassos subangulares dispersis distincta.

Typus: Mujin SAN 37849, Borneo, Sabah, Mt. Silam (holotypus SAR!;

isotypus SAN!).

Figure 5

Tree up to 9 m tall, 15 cm diameter. Bark shallowly fissured or scaly,

’ greyish brown; inner bark brittle, with white latex. Twigs stout, subangular,

glabrous. Terminal buds up to 8 mm long. Stipules triangular, c. 4 x 4 mm,

crested, glabrous. Leaves spirally arranged, scattered and well-spaced along

the twigs, thickly coriaceous, silvery brown tomentose on both sides; blade

elliptic to oblong-elliptic, 16-25 x 6-8 cm, base narrowly cuneate, margin

entire and plane, apex blunt or rounded; midrib raised on both sides,

stronger below; lateralveins 15-19 pairs, ascending at an angle of 70—80°

from the midrib, curved, diminishing and becoming inconspicuous near the

leaf margin, distinctly raised on both surfaces; intercostal venation laxly

reticulate; petioles 3-4 cm long, grooved on the adaxial side, thickened at

the base, glabrous. Inflorescence axillary, 2—3-flowered; pedicels 1—-1.5 cm

long, angular and slightly enlarged at both ends. Flowers: calyx biseriate,

consisting of two whorls of two sepals; sepals broadly ovate, 6 x 4 mm,

rusty-brown tomentose (mature flowers not seen). Fruits unknown.

Distribution: Endemic to Sabah; known only from the type collection from

Mt. Silam, Lahad Datu district and SAN 51742 from Bt. Tawai,

Kinabatangan district.

Ecology: In forest on ultrabasic soils, up to 850 m altitude.

Notes: The spirally arranged leaves and the calyx comprising two whorls of

two sepals confirm that the specimens belong to Madhuca. The species is

distinct from any known species of Madhuca from Borneo in having thickly

coriaceous, silvery-brown tomentose leaves that are scattered along the

subangular and stout glabrous twigs.

New combinations, names and species of Madhuca in Borneo 355

Figure 5. Madhuca silamensis. A, flowering leafy twig; B, flower bud; C, longitudinal section

of flower bud; D, gynoecium. (All from SAN 37849.)

356 Gard. Bull. Singapore 53 (2001)

Acknowledgements

The work was supported financially by the Tree Flora of Sabah and Sarawak

project jointly executed by the Forest Research Institute Malaysia (FRIM),

Sabah Forestry Department and Sarawak Forestry Department. We are

deeply indebted to the Director General and Directors of these institutions

for their support and encouragement. Grateful acknowledgements are also

due to the Directors/Curators of the Herbarium Bogoriense (BO), and the

herbaria of the Forest Research Institute Malaysia (KEP), Sabah Forestry

Department (SAN), Sarawak Forestry Department (SAR), Royal Botanic

Gardens, Kew (K), National Herbarium of Netherlands, Leiden Branch

(L), and Singapore Botanic Gardens (SING) for the loan of specimens

and/or hospitality accorded to us during our visit to their institutions. We

_are extremely grateful to Dr. J.F. Veldkamp for his kind help in providing

Latin translations for the diagnoses of the new species. Finally we would

like to express our thanks to Mr. Joseph Pao of the Sarawak Forestry

Department for diligently preparing the illustrations, and to Dr. E.

Soepadmo, Dr. Francis $.P. Ng, and Dr. W. Vink for their constructive

criticisms and comments on the manuscript.

References

Dubard, M.M.M. 1908. Les Sapotacées du groupe des Illipées. Revue

Génerale Botanique. 20: 201.

Gmelin, J.F. 1791. Systema Naturae. 2, 1: 772 & 799.

Pennington, T.D. 1991. The Genera of Sapotaceae. Royal Botanic Gardens,

Kew, U.K.

Corrigenda

Gardens’ Bulletin Singapore 52 (2000) 261-271.

The Significance of Pollen Morphology in the Taxonomy

of the Genus Durio (Bombacaceae)

I. SALMA

replace page 269 with the following:

graveolens, D. kutejensis, D. oxleyanus, were reported as oblate spheroidal.

Sharma (1970) showed D. lowianus and D. pinangianus had a similar pollen

shape, but that pollen of D. griffithii and D. singaporensis were oblate and

that of D. carinatus oblate spheroidal.

Generally, in the majority of species the thickness of the pollen wall

varied from 2—4 um. However, the pollen wall of D. griffithii and D.

excelsus was thin (less than 2 um) while that of D. testudinarum was thick

(greater than 4 um).

Size varied from 46 to 145 um for the polar diameter (P) and 50 to

120 um for the equatorial diameter (E) (Table 2). According to the

classification of the pollen size by Erdtman (1952), Durio pollen can be

divided into three groups, i.e. medium (25—S0 um), large (SO—100 um)

and very large (100—200 um). Most Durio species fell within the large

group except for D. griffithii, which had medium to large pollen, and D.

affinis, D. oxleyanus, D. pinangianus, D. testudinarum and D. zibethinus,

which possessed large to very large pollen (Table 2). Similar results were

obtained by Abang Mokhtar (1991), except that pollen of his sample of D.

griffithii and D. acutifolius fell within the medium-sized group and D.

graveolens in the very large group. This difference in size was probably

due to whether herbarium or fresh specimens were used. Since there was

variation in pollen size within a species and variation between species was

not distinct, pollen size is therefore not a good character for distinguishing

the species.

Conclusions

Only in D. testudinarum are pollen characters species-specific (Table 3)

and the pollen of this species can clearly be distinguished by a combination

of its large size, thick wall and verrucose exine. Although there are

differences in pollen morphological characters between the species,

intraspecific variation also occurs. A combination of pollen characters,

such as the exine sculpture, size and shape, can, in some cases (Table 3),

provide supplementary information, which should, however, be used

together with other characters for the identification of Durio species.

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Gardens’ Bulletin Singapore 53 (2001) 359-361.

Book Review: Paul Kessler (Ed) 2000. Secondary Forest Trees of

Kalimantan, Indonesia. A Manual to 300 Selected Species. 404 pp. MOFEC

Tropenbos, PO Box 319, Balikpapan, Indonesia. ISBN 90-5113-044-9.

Paperback. Price US$35, available from The National Herbarium

Netherlands, University of Leiden Branch, PO Box 9514, 2300 RA, Leiden,

The Netherlands.

This book is a welcome addition to the slowly growing botanical literature

on Borneo. The work now being undertaken on the Tree Flora of Sabah

and Sarawak (the Malaysian part of Borneo) has shown that the floristic

congruence between Indonesian Borneo (Kalimantan) and Malaysian

Borneo (Sabah and Sarawak) is over 80%. Any book on the flora of

Kalimantan is useful to Sabah and Sarawak and vice versa.

About half of the book (p 23-198) is taken up by descriptions of the

trees, in alphabetical order by family and species. Each description is

followed by concise notes on habitat and ecology, distribution and uses.

For each family, there is an identification key to the genera treated, and

under each genus, there is an identification key to the species. The other

half of the book (p 204403) is taken up by line drawings, one page per

species, illustrating about two-thirds of the 300 species.

There is no overall key to families or to the whole secondary forest

flora. Instead, the book offers a comprehensive list of spot characters under

which genera are listed where applicable. For example, under spot character

4: armed plants, are listed Cratoxylum, Flacourtia, Gmelina, Leea, Oxyceros

and Ziziphus. Under spot character 7: white sap, are listed Alstonia,

Artocarpus, Cerbera, Dyera, Ficus, Homalanthus, Parartocarpus, Prainea

and Tabernaemontana. Altogether 58 spot characters are compiled. I find

spot characters very useful, in fact, more useful than the keys. The keys

work only for the selection of species recognised in this manual, whereas

spot characters, working at the genus level, tend to cover all species of the

listed genera. One can query the list at any point instead of proceeding

along the pre-set path of a key. With slight changes, this spot character list

can be used anywhere in SE Asia.

The descriptions and illustrations are of a high professional standard.

Floral dissections, tertiary venation and indumentum details have been

omitted by the artists, but the form and habit of each plant are effectively

depicted. From my own experience, I know that getting the drawings done

for a flora can be a real hassle. This approach sacrifices the fine details, but

saves a lot of time. Otherwise, it would have been very difficult to get such

a high proportion of the species illustrated.

The plant identified as Phyllanthus emblica is actually Phyllanthus

360 Gard. Bull. Singapore 53 (2001)

pectinatus, as indicated by the position of the fruits towards the ends of the

leafy twigs. In P. emblica, the fruits would be close to the base. P. emblica

ranges across mainland Asia from India to South China but stops at Perlis

in north Peninsular Malaysia. P. pectinatus replaces it in Sumatra and

Peninsular Malaysia south of Perlis (Ng, F.S.P. 2000. Malaysian Naturalist

53(3): 32—35). All Bornean plants previously identified as P. emblica are

likely to be P. pectinatus.

I expected a discussion of the nature and origin of secondary forests

in Kalimantan, but the authors, P.J.A. Kessler, P.B. Pelser, C.E. Risdale

and K. Sediyasa have carefully avoided this topic. Instead, they offer this

cryptic introduction: “After the completion of our manual “Trees of the

Balikpapan — Samarinda area, East Kalimantan, Indonesia’ the production

of a manual to selected tree species of the secondary forest was considered

.an urgent necessity. Deforestation in Kalimantan was proceeding at an

alarming rate and had led to millions of hectares of more or less severely

degraded forest. The recent immense forest fires (1997-1998) added to the

loss and made a publication even more urgent.” In this way, the authors

try to convey a sense of urgency, but do not explain how deforestation and

burning impact upon forests, primary and secondary, in Kalimantan. A

discussion might have been politically difficult in a book sponsored by the

Indonesian Ministry of Forestry and Estate Crops.

I think that what is happening in Kalimantan is too important to

ignore. When I went to South Kalimantan recently, I found that the forests

there had been replaced by /mperata grassland. The sheer scale of

grasslands, in the absence of a cattle industry, took me by surprise. In the

humid tropics, it needs work to convert woody vegetation to grassland.

Logging alone does not result in grasslands. In 1994, I was in West

Kalimantan and was shocked to see that for hundreds of kilometres, fires

were burning all over the countryside. These were individually small fires

that did not obstruct road traffic, but the smoke and haze filled the skies.

It was obvious that these fires were the work of the people of the

countryside. Their little houses were spaced out so that each could lay

claim to many hectares of land — far more land than the settlers could farm

manually. To maintain their claims, the settlers had to keep their lands

cleared of natural vegetation. Annual burning during the dry season was

the only way to do this. Eventually, the woody vegetation would give way

to grassland. Why do the settlers clear more land than they can cultivate?

They were hoping that plantation companies would eventually acquire the

land for oil palm, pulpwood or other industrial crops, and pay compensation.

The loggers had indeed opened the way into the forests, but the settlers

did the rest. In this scheme of things, secondary forests in Kalimantan are

Book Review 361

not forests gradually recovering from logging, but a brief transient phase

between forest and /mperata grassland. West Kalimantan will soon look

like South Kalimantan and the rest of Kalimantan is not far behind.

Francis S.P. Ng

Kepong, Malaysia

Gardens’ Bulletin Singapore 53 (2001) 363.

Obituary

DR GUNNAR SEIDENFADEN

To our great sadness we learned that Dr Gunnar Seidenfaden passed away

in February 2001 at the respectable age of 92. His career as an orchid

taxonomist has been a long and extremely fruitful one. He continued to be

productive right up to his death, leaving several unfinished manuscripts.

His impact on the taxonomy of the orchids in the Indo-Chinese to

Malaysian region has been enormous. The core of his orchidological oeuvre

is his revision of the orchids of Thailand. In order to have at least a chance

of completing this project he developed his own, slightly unusual but

certainly concise style: full bibliography, full citation of specimens, good

illustrations but no descriptions. Making use of the information gathered

for this project, he produced a checklist for the orchids of Indo-China, and,

together with Jeffrey Wood, he revised R.E. Holttum’s Orchids of Malaya.

For the latter project he visited Singapore on several occasions to consult

the Singapore Botanic Gardens’ herbarium.

Seidenfaden can be ranked among the greatest in the field of South

East Asian orchidology, on par with Smith, Schlechter and Holttum. While

all these persons were professional plant taxonomists; for Seidenfaden,

however, orchid taxonomy was only one of his many fields of interest. One

could say that he ‘did’ the orchids of Thailand as a hobby, which makes his

achievements all the more impressive.

Gunnar Seidenfaden has left an admirable scientific legacy. His death

is much regretted.

Reference

Seidenfaden, G and J.J. Wood. 1992. The Orchids of Peninsular Malaysia

and Singapore. Olsen & Olsen, Denmark.

Jaap J. Vermeulen

Singapore Botanic Gardens

Singapore

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of the (first) author and the journal title be given in full, as in the following example:

Stone, B.C. 1994. Additional notes on the genus Glycosmis Correa (Rutaceae). Gardens’

Bulletin Singapore. 46: 113-119.

References to books and monographs should be cited according to the following form:

Ridley, H.N. 1930. The Dispersal of Plants Throughout the World. L.Reeve, Ashford,

Kent.

For literature citations in taxonomic papers the following style is required:

Medinilla alternifolia Blume, Mus. Bot. Lugd.-Bat. I:2 (1849) 19.

Sterculia acuminatissima Merr., Philip. J. Sci. 21 (1922) 524.

Illustrations: Drawings should be done in indian ink. Authors should indicate where individual

illustrations receive first mention in the text.

Offprints: Authors will be given 50 offprints gratis. Additional copies must be ordered and

paid for, prior to publication.

Manuscripts should be sent to: THE EDITOR, GARDENS’ BULLETIN SINGAPORE,

SINGAPORE BOTANIC GARDENS, CLUNY ROAD, SINGAPORE 259569

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