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THE GARDENS’ BULLETIN

SINGAPORE

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(int 5 raat . ©

eek, Pe Bice #

fort Sat & Ser F

VOLUME 54

Singapore Botanic Gardens

2002

Singapore, 2002

Applications for reproduction should be

made to the Singapore Botanic Gardens,

1 Cluny Road

Singapore 259569

DATES OF PUBLICATION OF THE TWO PARTS WERE:

Part 1 Pp. 1-164 31 July 2002

Part2 Pp. 165-294 20 December 2002

Singapore Gardens Bulletin 54 (2002).

CONTENTS

Argent, G.C.G.

New taxa and combinations in the genus Diplycosia (Ericaceae) of

SEO Aa CIRITIS INT AT PUP IN AY GR ooo oa oa an casnceopcvets vaactvoiaesévedsitesscsensbeasseseevebenes 217—238

Ashton, P. S.

Tazonomic notes on the tree flora Of Brunei 1 .,..cccdccvccorcosesasecclasdsveccocesvcessss 207-216

Burtt, B.L.

New Gesneriaceae: a Chirita from Vietnam and a Monophyllaea from

SANA ae Bette ION Mince cli chnc8y Valdes ce cn ick has dab cw cinocdass/e dab reureascoraresnsoueces 239—242

Chung, R.C.K.

Leaf epidermal micromorphology of Grewia L. and Microcos L.

(filiacear) in Peninsular Malaysia and BOrme -.........2..<./se.it-.c.sc0seseescedcseceess 263—286

Gideon, O.G.

Coprosma bougainvilleensis (Rubiaceae), a new species from

Bougdmmvile island. Papia NOW GiUs6a \0.0:.4.J.0c5.ecrecscincoaccdencesavnokitndpseserteresne’ 165—170

Hay, A and C. Herscovitch

Two remarkable new West Malesian Homalomena (Araceae) species ....... 171—178

Kiew, R.

Mangifera odorata (Anacardiaceae) iS a HYD ............ceseseeseeseeeeeeeeeeeeeeeees 205—206

Kiew, R.

A new species of Monophyllaea (Gesneriaceae) from Borneo..............c60 243—246

Kloet, S.P. Vander

Re-examination of Vaccinium dialypetalum (Ericaceae) ..........ccceseeseeeeeeeeees 179—183

hee, C.C:

Nepenthes platychila (Nepenthaceae), a new species of pitcher plant

TR GN iO OR N15 Oe OA MRS Jy wn OR MR Ae cle ee, 6a 257—261

Mabberley, D.J.

Limau hantu and limau purut: the story of lime-leaves (Citrus hystrix DC.,

UMA MeL) evga nee ie Gate, Saad 42512) (aac toy vat acer ed bes too gsiocednaman tage 185—197

Mabberley, D.J.

The identity of Swietenia alternifolia (Mill.) Steud. (‘Meliaceae’) ............. w. 253—255

Rustiami, H.

Two new species of Daemonorops (Palmae) from Sumatra.............cceeeeee 199-—204

Saw, L.G.

A new species of Vatica (Dipterocarpaceae) from Peninsular Malaysia .....247—251

Singapore Gardens Bulletin 54 (2002) iv.

Vermeulen, J.J.

A taxonomic revision of Bulbophyllum (Orcidaceae). 2: Sections

Altisceptrum, and Hirtiila ..g....1.:.s.cei-+.-na<axdesseentsanenps tudes ee 1—151

Vermeulen, J.J.

Bulbophyllum praetervisum J.J. Verm. (Orchidaceae), an overlooked

species close to B. macranthum Lindl ...::,..::.2,.73 153—154

Vermeulen, J.J.

Updates on sections of Bulbophyllum (Orchidaceae) revised in

Vermeulen, (1993) ..ccspseatdsscesdanenacceaegecseisp es seeeses nee nee ene aim 287—288

BOOK REVIEWS

The Ecology of Trees in the Tropical KainBorest 22. R.B. Primack - 155—157

Orchids of Sumatra:.....45235.2 9 J.J. Vermeulen, J.J. and P. O’Byrne - 159—160

World Checklist and Bibliography of Sapotaceae i ee R. Kiew - 161

Vegetation and Vascular Flora of Doi Sutep-Pui National Park,

Northern Thailand: <:......4.25¢n0 00 ee ee R. Kiew -163—164

Tree Flora of Sabah and Sarawak: Vol4.......23 ee P.S. Ashton - 289—292

A Field Guide to the Rattans of Lao PDK 2. 2 eee R. Kiew - 293—294

SE eee en eee

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54 (Part 1) July 2002

Singapore Botan

VOL.

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

Riyksherbarium

Leiden, Netherlands

Dr E. Soepadmo

Forest Research Institute Malaysia

Kepong, Malaysia

Dr W.K. Tan

Singapore Botanic Gardens

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

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

conclusions expressed by the contributing authors.

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

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

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

Singapore.

Instructions for contributing authors are found on the inside backcover.

£ AE Re den x15. \ a ee ~ BS i

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The Gardens’ Bulletin

Singapore

ae as oe

¢ — 17 2003 PONE

VOL. 54 (Parts 1) July 2002 ISSN 0374-7859

CONTENTS

Vermeulen, J.J.

A taxonomic revision of Bulbophyllum (Orchidaceae) 2: Sections Altisceptrum and

MeN Ne A a ee ae if ide OM cE Ca a st ashe tbat cadevind sania Zausislcn 1

Vermeulen, J.J.

Bulbophyllum praetervisum J.J. Verm. (Orchidaceae), an overlooked

Be Cies erse GOT, MUGEMORUIIIIIN VAIL so icc oecsintacncvcineneovecectansnanvedepcectinnnd nentadecoonen asta nsnddeeendos 3

BOOK REVIEWS

the Ecology of Trees in the TropitalfRaim Forest .........5..0......00-.0+00<00 R.B. Primack - 155

COULSON DUAN ea. Secs gap snstadcnccnincstenedounitanadeannanous J.J. Vermeulen and P. O’Byrne - 159

World Checkhst.and- Bibliography of Sapotacede iio cucs tees. fp bce diee sivas R. Kiew - 161

Vegetation and Vascular Flora of Doi Sutep-Pui National Park,

ES a ena Tara NAAN I SN ax ccs Se asec erase sotpecwinetlaocstnp/seng <wovisremp aeanaslid ipmgrotcnnsntpars ivi R. Kiew - 163

Date of Publication: 31 July 2002

Published by

National Parks Board

Singapore Botanic Gardens

Cluny Road

Singapore 259569

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Gardens’ Bulletin Singapore 54 (2002) 1-151.

A Taxonomic Revision of Bulbophyllum (Orchidaceae)

2. Sections Altisceptrum and Hirtula*

J.J. VERMEULEN

Singapore Botanic Gardens

Singapore 259569

Abstract

Bulbophyllum sect. Altisceptrum and sect. Hirtula (Orchidaceae) are revised. Both sections

are distributed in South and East Asia, and Malesia. The two sections together include 49

species and | subspecies. The investigations leading to this paper yielded 25 new species.

Many of these have been described in precursors to this revision, eight are described here

as new: B. sororculum, B. lanuginosum, B. grotianum, B. scaphiforme, B. comberipictum,

B. ochthodes, B. debrincatiae, and B. pilosum. All species (except one) are illustrated.

Introduction

The present revision is a continuation of Vermeulen (1993); consequently,

the introductory chapters 2 to 6 in that first instalment of the series largely

apply to this revision. The numbering of the species is also continued. This

revision therefore starts with species number 111. Whereas the sections of

Bulbophyllum revised in the first instalment occur almost exclusively in

East Malesia, sect. A/tisceptrum and sect. Hirtula are predominantly from

South and East Asia, and West Malesia. The species in both sections have

racemose inflorescences.

Section Altisceptrum includes 9 species and 1 subspecies. These are

large plants that are rather similar to the racemose series of sect. Sestochilus,

but they generally differ in having long and narrow, 1-veined petals. On

the whole, the delimitation of the species is rather straightforward, although

a re-evaluation of B. disjunctum sensu Vermeulen (1991: 21) led to its

division into two distinct species and one subspecies. Bulbophyllum

gymnopus Hook.f. is included in the section; it had recently been ge Fast in

Drymoda Lindl. by Garay et al. (1994: 641).

Section Hirtula comprises 40 species. These are generally

characterised by the presence of 1-celled hairs on the flowers, particularly

on the petals and the lip. Most species are rather rare in nature, and only

sparsely represented in collections. This tends to make a taxonomist’s life

deceptively easy, but in fact the reliability of the revision suffers because

the delimitation of the species relies on the taxonomist’s intuition rather

than on the evaluation of a representative set of specimens. Only one of

* continued from Orchid Monographs 7 (1993): 1—324

2 Gard. Bull. Singapore 54 (2002)

the commoner species, B. hirtulum Ridl., was found to display more than

average infraspecific variability. Apart from this, confusion about the

identity of B. nigrescens Rolfe, B. nigripetalum Rolfe, and B. secundum

Hook.f. (see Seidenfaden, 1979) has been resolved, and a re-evaluation of

the Thai material of B. penicillium Par. & Rchb.f. in Rchb.f. has led to its

description as a new species.

Although this revision includes only 8 new species, no fewer than 25

species out of the 49 included have been described as new by the present

author over the last 10 years. The present revision presents the first full

plates of many of these.

Extinct species. Several species are marked in this revision as ‘probably

extinct’. This is defined here as not having been found for at least 50 years,

following the IUCN definition of the term ‘extinct’. However, many

supposedly extinct plant species that were collected long ago occur in

areas that are only rarely visited by botanists. The species may be extinct,

particularly if they were found in lowland areas, but they may still be

present, unnoticed, in remote remnants of Asia’s forests.

Section Altisceptrum J.J. Sm.

Bulbophyllum sect. Altisceptrum J.J. Sm., Bull. Jard. Bot. Buitenzorg, s. 2, 13 (1914)

34. - LECTOTYPE: Ephippium elongatum Blume [= Bulbophyllum elongatum

(Blume) Hassk.], here designated.

Rhizome creeping, scales usually covering rhizome, pseudobulbs and

leaf bases before disintegrating. Roots predominantly sprouting below the

pseudobulbs. Pseudobulbs 1-leafed, not or hardly laterally flattened,

wrinkling along pre-determined longitudinal lines or in a random pattern

with age. Inflorescence with many flowers. Rhachis not swollen when

compared to the peduncle. Floral bracts patent to reflexed. Flowers spirally

arranged, sometimes partly in whorls (penduliscapum, elongatum), not

resupinate, or resupinate (gymnopus), not fully opening, many

simultaneously. Pedicel and ovary: basal node usually more or less flush

with the surface of the rhachis, or on a stump up to 2 mm high (gymnopus).

Median sepal porrect or somewhat recurved with an incurved top part, 3-

veined. Lateral sepals free, recurved to spreading, 3-veined. Petals linear,

often gradually tapering from a wide base, or elliptic to obovate with a

linear, long drawn-out top part (nemorale), margins usually erose-

denticulate, sometimes entire or papillose-ciliolate; the lower margin often

decurrent along the column foot, 1-veined, or 2—3-veined (nemorale). Lip

usually without a median slit, rarely with a median slit (fulvibulbum),

glabrous or finely papillose locally, thin to thick but not fleshy. Column:

Bulbophyllum Sect. Altisceptrum and Hirtula 3

stigma not protruding at its base; column foot elongated in several species,

leaving a gap between the median and lateral sepals that is not filled

entirely by the petals (distinctly present in gymnopus; moderately distinct

in elongatum, slight in placochilum and lissoglossum); column foot otherwise

without accessories. Stelidia distinct, triangular to linear, often subulate,

often with a small tooth along the upper margin and a narrow wing along

the lower, at the base. Anther abaxially with a papillose ridge near its base;

frontal part not or hardly projecting, front margin glabrous to papillose.

Pollinia 4; the inner + half as long as the outer or longer, flattened on both

sides; the outer flattened on one side. Stipes absent.

Distribution. Bhutan (1 taxon), India (1), Thailand (2), Peninsular

Malaysia (3), Sumatra (2), Java (1), Borneo (7, of which 5 are endemics),

Philippines (3), New Guinea (2).

Delimitation of the section. Section Altisceptrum 1s very similar to the

racemose series of sect. Sestochilus. The main distinctive characters lie in

the petals: although slightly varying in shape, they all consist of a wide

basal part tapering into a long, linear top part. They usually have one vein.

While most have petals with 3 or more veins, some racemose species

of sect. Sestochilus also have 1-veined petals: Bulbophyllum pachyphyllum

J.J. Sm., B. tahanense Carr, B. univenum J.J. Verm., and B. wrayi Hook.f.

These have distinctly shorter and wider petals than sect. Altisceptrum, and

entire and glabrous margins. Bulbophyllum nemorale L.O. Williams, of

sect. Altisceptrum, has petals with three veins, and B. elongatum (Blume)

Hassk., has petals with entire and glabrous margins. On account of overall

similarity, both have been included in sect. Altisceptrum.

Garay et al. (1994) transferred Bulbophyllum gymnopus to the genus

Drymoda Lindl., on account of the presence of a gap between the petals

and the lateral sepals. Drymoda otherwise includes minute plants with

deciduous leaves, 1-flowered inflorescences, and flowers with a lip of a

different shape, and a column with projecting teeth along the lower margin

of the stelidia. Bulbophyllum gymnopus is not obviously similar; it fits

better in sect. Altisceptrum. The displacement of the basal node of the

pedicel, the resupinate flowers, and the wide gap between the petals and

lateral sepals are unique characters for the species within the section; the

last character occurs, less distinctly, in several other species of sect.

Altisceptrum.

Bulbophyllum phreatiopse J.J. Verm., here included in sect. Hirtula,

has petals with papillose-ciliolate margins, but these are shorter and wider

than in sect. A/tisceptrum.

In spite of these species, sect. Altisceptrum possesses an overall

distinctness when compared with sect. Sestochilus, mainly through its rather

small, spidery flowers. It is therefore best to maintain it as a separate

4 Gard. Bull. Singapore 54 (2002)

group. J.J. Smith originally listed three species: Bulbophyllum elongatum

(Blume) Hassk., B. penduliscapum J.J. Sm., and B. pachyphyllum J.J. Sm.

The latter is best included in the racemose series of sect. Sestochilus for

reasons given above. Otherwise J.J. Smith’s concept of the section is

followed.

Monophyly. It seems reasonable to assume monophyly a priori on

account of a rather marked overall similarity of the species. Section

Sestochilus seems the most likely outgroup; however, sect. Altisceptrum

may well be nested within sect. Sestochilus. If this were true, its status as a

separate section would leave Sestochilus as a paraphyletic group.

Cultivation. Five species have been cultivated in recent years. The

general rules for growing Bulbophyllum apply here: semi-shade, a substrate

with some ‘soft’ organic matter that is never allowed to dry out entirely,

high air humidity and abundant air movement. High air humidity seems

particularly important except, possibly, for B. gymnopus. The plants grow

rather easily on tree-fern slabs, with some sphagnum or other soft material

among the roots when mounted to help them settle, or in baskets with a

potting mix containing sufficient soft organic matter. However, B.

penduliscapum is growing well outdoors in Singapore in a small pot mainly

filled with medium-sized charcoal chips. Bulbophyllum disjunctum often

occurs abundantly in its natural habitat but is rarely found in flower. It

may continue this habit of shy flowering when taken into cultivation.

Key to Species of Bulbophyllum sect. Altisceptrum

1. Leaf tip rounded to obtuse (check undamaged leaves)

2. Inflorescence pendulous. Veins of rhizome bracts persistent as a

whorl of stiff, erect bristles around the pseudobulb ...............0.0.

een ares ee ee sosddanbnshensieip -phsdesactrinionpte, tlc Tc

2. Inflorescence erect to patent. Veins of rhizome bracts more or less

decaying with the bract itself

3. Lateral sepals 5—8 mm long. Flowers resupinate: column foot

directed, towards the base.of the inMOresCe nie icc cbrctpseesent uence

a a lal a 112. B. gymnopus

3. Lateral sepals 10—22 mm long. Flowers not resupinate: column

foot directed towards the tip of the inflorescence

4. Median sepal 2—3.2 mm wide. Lateral sepals 2.2—3.8 mm

WUE cs onsn'siasiia tnancooss cesta te eae ae eae 113. B. farinulentum

4. Median sepal 3.4—5.3 mm wide. Lateral sepals 4.5—5 mm

11s | reer oe ene oem CE tate Ee 114. B. fulvibulbum

1. Leaf tip acute to acuminate

Bulbophyllum Sect. Altisceptrum and Hirtula >

5. Petals with 3 veins, the laterals sometimes not reaching the base.

Petals with an elliptic to obovate basal half and a linear top half ...

Pes EI TER ccTLD cre UR MUIETS 32 FATIH Aach veroabaohe 115. B. nemorale

5. Petals with 1 vein. Petals gradually tapering from a wide base, or a

from widest part close to the base

6. Keels on the adaxial side of the lip at the proximal end with two

erect10 autrorsehobtuserknobsy. ci sninss..v,-- 116. B. disjunctum

6 Keels on the adaxial side of the lip without knobs, or no keels

present

7. Lip orbicular to transversely elliptic in outline (do not spread

UE Siete tack oint ce ME re cael, matey eg 117. B. placochilum

7. Lip + triangular in outline, with or without a projecting,

linguiform top

8. Median sepal 12.5—17.5 mm long. Lip 2.4—3 mm wide.

Leaves in flowering plants 6—16 cm wide ........... eee

ie) .Lipeiel Sone ett brag .onv-ecia 118. B. elongatum

8. Median sepal 8—11.2 mm long. Lip 1—1.8 mm wide. Leaves

in flowering plants 2.1—4.5 cm wide... 119. B. lissoglossum

111. Bulbophyllum penduliscapum J.J. Sm. — Fig. 1, plate 1.

Bulbophyllum penduliscapum J.J. Sm., Icon. Bogor. 2 (1903) 101. — TYPE: Indonesia,

Sumatra, northern part, Heldt s.n. (BO, holo., not seen).

Bulbophyllum macrophyllum F. Kraenzl., Bot. Jahrb. Syst. 34 (1905) 249. — TYPE:

Malaysia, Sarawak, Beccari PB 3113 (FI, lecto.; K, isolecto., indicated by

Vermeulen, 1991), PB 1344 (FI).

Bulbophyllum jarense Ames, Orch. 6 (1920) 301. — TYPE: Philippines, Leyte, Jaro,

Wenzel 761 (PNH holo., destroyed; AMES, K, NY, S, iso.).

Secondary roots erect, growing in dense tufts surrounding the plant.

Rhizome 6—10 mm diam., sections between pseudobulbs 1.8—4 cm long;

bracts soon withering but veins persistent as a whorl of stiff, erect bristles

around the pseudobulbs. Pseudobulbs close to distant, discoid, 0.5—0.7 x

0.5—1.2 cm. Petiole 11—24 cm. Leaf blade elliptic, 21—38 x 7.5—15.5 cm,

index (length/width) 1.6—2.8; rounded to obtuse. Inflorescence a rather

dense raceme, with the flowers + in whorls of 4—8 towards the tip of the

rhachis, pendulous, 23—65 cm, 50—250-flowered. Peduncle 9—14 cm, bracts

5—6, the longest 40—60 mm long. Rhachis 12—58 cm. Floral bracts

triangular, 4—7 x 1.5—2 mm, acute. Pedicel and ovary 3—5S mm long.

Median sepal ovate to triangular, 8—12 x 1.5—2 mm, index 4.7—6.7; acute

to acuminate, margins entire; glabrous or finely papillose locally, base

broadly attached; rather thin. Lateral sepals slightly oblique, 8—12.5 x 1—

2.5 mm, index 4.1—8; otherwise as the median sepal. Petals porrect, often

falcate, gradually tapering from a wide base, 5—9 x 0.7—0.9(—1.2) mm,

6 Gard. Bull. Singapore 54 (2002)

index 7.1—11.3; acute, margins finely erose; glabrous or finely papillose

locally, base broadly attached; rather thin. Lip recurved in the basal half.

triangular, 2—3.5 x 1—1.6 mm, index 2—2.5; obtuse, margins entire or

slightly erose and somewhat finely papillose; rather thin; adaxially concave

near the base, with 2 distinct, obtuse ridges diverging towards the base and

running to about half-way up the lip, space in between channelled, surface

glabrous except for a papillose strip proximally along the median line,

abaxially with a weak, rounded ridge up to about 1/3 of the length of the

lip, surface glabrous. Column 2.2—3 mm long. Stelidia falcate or not,

subulate, 0.8—1.2 mm long, acute, often with a small, antrorse, deltoid,

obtuse to acute tooth along the upper margin.

Colours: Sepals pale yellow at the base, yellow at the tip, with large

dark purple spots. Petals translucently yellow with similar spots. Lip

brownish at the base, yellow at the tip. Column yellowish. Flowers sweetly

fragrant, attractive to flies.

Habitat & ecology: Found in kerangas forest, forest on limestone,

and lower montane forest. Epiphyte growing in large clumps on tree trunks

and major branches, catching leaf litter in the dense tangle of secondary

roots. Alt. 300-1100 m. Flowering in Feb—Apr, Nov, Dec.

Distribution: MALAYSIA: Peninsula ( 2 living plants seen); Sarawak

(3 specimens seen); Sabah (8). PHILIPPINES: Leyte (2). INDONESIA:

Sumatra (2).

Notes: Unmistakable, even without flowers, because of the clustered

pseudobulbs, the persistent fibres of the rhizome scales, and the tufts of

erect catch-roots. Well-grown, flowering plants can be quite spectacular.

Some variability occurs in the stelidia; these are porrect in some

specimens (e.g. the type of B. jarense) but strongly curved downwards in

others (e.g. the type of B. macrophyllum).

112. Bulbophyllum gymnopus Hook.f. — Fig. 2.

Bulbophyllum gymnopus Hook.f., Fl. Brit. Ind. 5 (1890) 764; id., Hooker’s Icon. Pl. 21

(1892) t. 2040; Seidenf., Dansk Bot. Ark. 33, 3 (1979) 100. — Phyllorchis gymnopus

(Hook.f.) O. Kuntze, Rev. Gen. Pl. 2 (1891) 677. — Drymoda gymnopus

(Hook.f.) Garay, Hamer & Siegerist, Nordic J. Bot. 14 (1994) 641; Seidenfaden,

Opera Bot. 124 (1995) 47, colour photograph pl. 9b. — TYPE: Bhutan, Griffith

5133 (K, lecto., indicated by Seidenfaden, 1979); India, Khasia Hills, Griffith

5134 (K, W); India, Assam, Jowai, Clarke 42549 (K).

Figure 1. Bulbophyllum penduliscapum J.J. Sm. (111) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial

side, right: abaxial side; e. Column and lip, lateral view; f. Anther, above: abaxial side,

below: adaxial side; g. Pollinia, above: single pair, below: two pairs. — All from Vermeulen

1147 (living plant).

Bulbophyllum Sect. Altisceptrum and Hirtula

rhe rs ii i

BW hee Uuehinnd paren ‘}

+ aan

“HA CSS

Figure 1. Bulbophyllum penduliscapum J.J. Sm. (111)

8 Gard. Bull. Singapore 54 (2002)

Rhizome 2.5—4.5 mm diam., sections between pseudobulbs 3—22

cm long; bracts, including veins, soon withering. Pseudobulbs distant, ovoid,

1.2—4 x 0.8—1.5 cm. Petiole 1.0—3.5 cm. Leaf blade elliptic, 4.8—17 x 1—

3.2 cm, index (length/width) 4.3—6.3; obtuse. Inflorescence a lax raceme,

erect to patent, 7—23 cm, 7—28-flowered. Peduncle 1.8—7 cm; bracts 4—

6, the longest 5—17 mm long. Rhachis 5—17 cm. Floral bracts ovate to

triangular, 1.8—3.5 x 1—1.8 mm, acute. Flowers resupinate. Pedicel and

ovary 8—10 mm long, basal node on a 1—2 mm-long stump. Median sepal

ovate to triangular, 4.5—7 x 1.4—2.2 mm, index 2.2—3.9; emarginate,

rounded to acute, margins entire; glabrous, base narrowly attached; thin,

surface glabrous or slightly papillose towards the tip abaxially. Lateral

sepals falcate, 5—8 x 1.4—2.5 mm, index 3—4.3; obtuse, otherwise as the

median sepal. Petals recurved, falcate, linear with a wide base, 3—5.5 x

0.4—0.7 mm, index 4.2—11; obtuse to acuminate, margins erose towards

the tip; glabrous, base broadly attached; thin. Lip recurved slightly below

half-way, linguiform, 2—4 x 0.8—1.3 mm, index 1.6—5; obtuse, margins

entire, glabrous or finely papillose locally; thick, surface + glabrous; adaxially

concave near the base, often with 2 inconspicuous, obtuse ridges about

half-way up the lip, space in between the two ridges somewhat channelled;

abaxially with a distinct, deeply emarginate ridge up to about 1/3 of the

length of the lip. Column 2—3 mm long, foot elongated so that a gap exists

between the petals and the lateral sepals. Ste/idia triangular, 0.4—1 mm

long, acute, often with an inconspicuous, deltoid, obtuse tooth along the

upper margin.

Colours: Rhizome scales in young shoots densely spotted with purplish

brown. Sepals and petals white, pale greenish white or yellowish. Lip yellow

or orange. Column white to orange. Flowers with an unpleasant smell.

Habitat & ecology: Epiphytic or lithophytic in evergreen forest

(Seidenfaden, 1979), in montane forest. Alt. 600-2000 m. Flowering in

Jan—Mar and Oct—Dec.

Distribution. BHUTAN (1 specimen seen), INDIA: Sikkim (1);

Khasia Hills (1); Naga Hills (1); Assam (5). THAILAND (2).

Notes. The densely spotted scales of the young shoots are very

characteristic in this species.

Figure 2. Bulbophyllum gymnopus Hook.f. (112) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial

side, right: abaxial side; e. Column and lip, lateral view; f. Anther, above: adaxial side,

below: abaxial side; g. Pollinia, left: two pairs, right: single pair. — a. From King’s

collectors s.n. (herbarium specimen); b—e. From Seidenfaden & Smitinand GT 8643

(spirit sample); f. From Herb. Hooker f. s.n. (herbarium specimen); g. From Hort. Kew

EN 596-68 (spirit sample).

Bulbophyllum Sect. Altisceptrum and Hirtula 9

3mm

Il mam

2mm

Figure 2. Bulbophyllum gymnopus Hook ¢. (112)

10 Gard. Bull. Singapore 54 (2002)

113. Bulbophyllum farinulentum J.J. Sm.

Rhizome 4—8 mm diam., sections between pseudobulbs 1.2—5 cm

long: bracts, including veins, soon withering. Pseudobulbs close to distant,

ovoid, 0.8—2.2 x 0.8—2.2 cm. Petiole 2.5—18 cm. Leaf blade elliptic to

obovate, 8.5—27 x 3—7.5 cm, index (length/width) 2.2—4.2; rounded to

obtuse. Inflorescence a lax to dense raceme, erect to patent, 19—55 cm,

5—150-flowered. Peduncle 10—25 cm; bracts 5—46, the longest 18—40 mm

long. Rhachis 3.5—22 cm. Floral bracts ovate to triangular, 6.8—12 x 1.5—

2 mm, acute. Flowers not resupinate. Pedicel and ovary 4—15 mm long.

Median sepal ovate to triangular, 9.5—19 x 2—3.2 mm, index 4.2—8.7;

acute, margins entire, + glabrous, base broadly attached; rather thin;

adaxially finely papillose to pubescent except for the basal part. Lateral

sepals 10—22 x 2.2—3.8 mm, index 3.9—8.8; otherwise as the median sepal.

Petals recurved, tapering from a wide base, 6.5—13.5 x 0.9—1.6 mm, index

4—10.4; acute to acuminate, margins + entire, papillose to ciliolate, base

rather narrowly attached; thin, top somewhat thickened; adaxially finely

papillose to pubescent except for the basal part. Lip recurved in the basal

half, + triangular, 2.8—4.6 x 1—1.7 mm, index 2.1—3; subacute, margins

entire, finely papillose locally; thick; adaxially slightly concave near the

base, with 3 ridges: 2 distinct, obtuse ridges proximally diverging and often

protruding like two retrorse, conical, obtuse knobs and distally running

parallel up to about 2/3—3/4 of the length of the lip, space in between the

two ridges deeply channelled and proximally with an inconspicuous to

rather distinct, rounded median ridge with longitudinal rows of minute

papillae on its crest, adaxial surface otherwise finely papillose locally;

abaxially with rather inconspicuous, retuse to truncate ridge up to about 1/2

of the length of the lip, surface glabrous. Column 2.3—3 mm long. Stelidia

narrowly triangular to subulate, 0.8—1.2 mm long, acute, often with a

small, antrorse, deltoid, obtuse to acute tooth along the upper margin;

often with an inconspicuous, deltoid, obtuse to broadly rounded wing along

the lower margin, at the base

Notes: The material allows two subspecies to be distinguished, mainly

on account of the size of the vegetative parts and the number of flowers in

Figure 3. Bulbophyllum farinulentum J.J. Sm. ssp. farinulentum (113a) — a. Habit; b.

Flower; c. Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; d.

Lip, left: adaxial side, right: abaxial side; e. Column and lip, lateral view; f. Anther,

left: abaxial side, right: adaxial side; g. Pollinia, left: single pair, right: two pairs. —

Bulbophyllum farinulentum J.J. Sm. ssp. densissimum (Carr) J.J. Verm. (113b) — h.

Lip, left: adaxial side, right: abaxial side; 1. Column and lip, lateral view. - a—e. From

Hort. Leiden 20771, f, g. From Hort. Leiden 26544 (all spirit samples); h. From Synge

98 (herbarium specimen); i. From Hort. Leiden 914076 (spirit sample).

Bulbophyllum Sect. Altisceptrum and Hirtula

Figure 3; a—g. Bulbophyllum farinulentum J.J. Sm. ssp. farinulentum (113a);

h—i. Bulbophyllum farinulentum J.J. Sm. ssp. densissimum (Carr) J.J. Verm. (113b)

12 Gard. Bull. Singapore 54 (2002)

the inflorescence. Further collecting, however, may reveal intermediate

forms.

Vermeulen (1991: 21) incorrectly regarded this as synonymous with

B. disjunctum. The differences are given below the latter.

Key to the subspecies

re Inflorescence 5—25-flowered. Flowers 1n a lax raceme...........:.........

ie, Tasnipeteieti A SN Jay MMR: ceplontc: Santas Mts on a. subsp. farinulentum

1. Inflorescence 60—150-flowered. Flowers in a dense raceme.............

a. subsp. farinulentum -— Fig. 3.

Bulbophyllum farinulentum J.J. Sm., Bull. Jard. Bot. Buitenzorg, s. 3, 2 (1920) 87. —

TYPE: Indonesia, Sumatra, Kerinci, Jacobson cult. 1413 (Leg. Groenenveldt)

(BO, holo., not seen).

Bulbophyllum noeanum Kerr, Bull. Misc. Inform., Kew (1927) 219. — TYPE: Thailand,

Pattani, Kerr 0113 (K, holo.).

Bulbophyllum disjunctum sensu J.J. Verm., Orch. Borneo 2 (1991) 21 (pro parte), non

Ames & C. Schweinf.

Rhizome 4—6 mm diam. Pseudobulbs 1—2.2 x 0.8—2.2 cm. Petiole

2.5—10 cm. Leaf blade, 8.5—20 x 3—6.5 cm. Inflorescence a lax raceme,

19—55 cm, 5—25-flowered. Rhachis 3.5—22 cm. Pedicel and ovary 10—15

mm long. Lip 3—4.6 x 1—1.7 mm, often with the ridges proximally

protruding like two inconspicuous to distinct knobs. Stelidia narrowly

triangular, 0.8—1.2 mm, with or without a wing along the lower margin, at

the base.

Colours: Sepals and petals (pale) yellow to pale ochre, with red veins.

Lip yellow, reddish brown in the centre, or in the apical half.

Habitat & ecology: Epiphyte growing in shaded places in kerangas

forest in lowland and montane conditions. Alt. 400—2000 m. Flowering in

Jan, Feb, May, Sep—Nov.

Distribution: THAILAND: Peninsula (1 specimen seen).

MALAYSIA: Peninsula (3); Sarawak (1); Sabah (5). INDONESIA:

Sumatra (fide J.J. Smith); Kalimantan (1).

b. subsp. densissimum (Carr) J.J. Verm., stat. nov. — Fig. 3, plate 2.

Bulbophyllum densissimum Carr, Gard. Bull. Straits Settlem..8 (1935) 116. — TYPE:

Malaysia, Sarawak, Dulit Range, Synge 98 (K, holo.).

Bulbophyllum disjunctum sensu J.J. Verm., Orch. Borneo 2 (1991) 21 (pro parte), non

Ames & C. Schweinf.

Bulbophyllum Sect. Altisceptrum and Hirtula 13

Rhizome 6—8 mm diam. Pseudobulbs 0.8—1.5 x 0.8—1.5 cm. Petiole

9—18 cm. Leaf blade, 17—27 x 6.5—7.5 cm. Inflorescence a dense raceme,

21—36 cm, 60—150-flowered. Rhachis 8—18 cm. Pedicel and ovary 4—8

mm long. Lip 2.8—3.2 x 0.9—1.1 mm, with the ridges proximally protruding

like two distinct knobs. Stelidia subulate, c. 1.2 mm, + without a wing along

the lower margin.

Colours: Peduncle bracts yellowish green or red. Flowers entirely

creamy yellow, slightly sweetly scented.

Habitat & ecology: Epiphyte in lowland (kerangas) forest and

probably lower montane forest. Alt. 900 m. and probably higher. Flowering

in Mar, Aug.

Distribution: MALAYSIA: Sarawak, Hose Mts. (3 specimens seen).

Notes. Forms of B. disjunctum with dense-flowered inflorescences, and

occurring in the same area, may look similar but have acute leaves.

114. Bulbophyllum fulvibulbum J.J. Verm. — Fig. 4, plate 3.

Bulbophyllum fulvibulbum J.J. Verm., Orch. Borneo 2 (1991) 23. — TYPE: Malaysia,

Sabah, Ulu Padas, Vermeulen 603 (L, holo.; K, iso.).

Rhizome 12—14 mm diam., sections between pseudobulbs 3—4 cm

long; bracts, including veins, soon withering. Pseudobulbs distant, ovoid,

3—4 x 2.5—2.8 cm. Petiole 18—25 cm. Leaf blade elliptic to obovate, 37

40 x 8.5—10 cm, index (length/width) 4—4.4, obtuse. Inflorescence a rather

lax raceme, erect to patent, 28—82 cm, 18—62-flowered. Peduncle 13—29

cm; bracts 5—6, the longest 25—50 mm long. Rhachis 12—53 cm. Floral

bracts ovate, 6—15 x 2—3 mm, acute. Flowers not resupinate. Pedicel and

ovary 14—25 mm long. Median sepal ovate, 10—14.3 x 3.4—5.3 mm, index

2—4.3; obtuse, margins entire, finely papillose or ciliolate towards the tip,

base broadly attached; rather thick, top thickened; surface finely papillose

towards the tip. Lateral sepals 12—17 x 4.5—5 mm, index 2.5—3.6; otherwise

as the median sepal. Petals recurved, tapering from a wide base, 8—11 x

1.5—2.2 mm, index 4.2—5.4; obtuse, margins entire, finely ciliolate towards

the tip, base broadly attached; thick, top thickened, surface adaxially very

finely papillose towards the tip. Lip recurved slightly below half-way,

linguiform, 3.5—4.6 x 1.8—2.5 mm, index 1.7—2.1; obtuse, margins entire,

glabrous or finely papillose locally; thick; adaxially slightly concave near

the base, with 3 ridges: 2 distinct, obtuse ridges proximally diverging and

protruding backwards like two conical, obtuse horns and distally running

parallel up to about 2/3 of the length of the lip, space in between the two

ridges deeply channelled and with a distinct, narrow, obtuse often finely

papillose median ridge with or without a thin median slit over its crest,

adaxial surface otherwise glabrous but finely papillose towards the base;

14 Gard. Bull. Singapore 54 (2002)

abaxially with a distinct, truncate ridge up to about 1/3 of the length of the

lip, surface glabrous. Column 3—5 mm long. Stelidia triangular, 0.5—1.3

mm long, acute, often with a small, antrorse, deltoid, obtuse to acute tooth

along the upper margin.

Colours: Sepals yellow or greenish, adaxially with or without dark

red spots, abaxially (sometimes also adaxially) with dark red flushing

towards the base. Petals yellow or greenish. Lip yellow or greenish, suffused

with dark red towards the base, or lip entirely dull red with a pale green

tip. Flowers scentless.

Habitat & ecology: Found as an epiphyte in montane forest with

Agathis, and in kerangas forest growing near the forest floor. Alt. 1100—

1500 m. Flowering observed in Oct (Mar, Apr in greenhouse).

Distribution: MALAYSIA: Sarawak, Bario area (1 specimen seen);

Sabah, Ulu Padas (3).

Notes: Close to the preceding species; differs in having flowers with

wider parts. The median ridge on the adaxial side of the lip is always very

distinct, whereas it is usually inconspicuous in B. farinulentum.

115. Bulbophyllum nemorale L.O. Williams — Fig. 5.

Bulbophyllum nemorale L.O. Williams, Bot. Mus. Leafl., Harvard University 9 (1938)

100. — TYPE: Philippines, Leyte, Jaro, Masaganap, Wenzel 871 (PNH, holo.,

destroyed; AMES, iso.).

Rhizome 5—8 mm diam., sections between pseudobulbs 2—5 cm

long. Pseudobulbs close to distant, ovoid, 1—1.3 x 0.8—1.5 cm. Petiole

7.5—17 cm. Leaf blade elliptic, 15—35 x 4.5—10.5 cm, index (length/width)

2.8—5; acute to acuminate. Inflorescence a lax to rather dense raceme,

erect to patent, 21—49 cm, 13—40-flowered. Peduncle 9—32 cm; bracts

6—8, the longest 20—48 mm long. Rhachis 9—19 cm. Floral bracts elliptic

to ovate, 7—12 x 2—3.5 mm, acute to slightly acuminate. Flowers + secund.

Pedicel and ovary 7—11 mm long. Median sepal ovate to triangular, 13—

17 x 2.4—4 mm, index 3.5—5.6; acute, margins entire; glabrous, base rather

broadly attached; rather thin. Lateral sepals falcate, 13—15 x 4.2—5 mm,

index 2.6—3.7, base broadly attached; otherwise as the median sepal. Petals

porrect, falcate, basal half elliptic to obovate, top half linear, 11—14 x

1.5—3, index 4—7.3; acute, margins finely erose + half-way; glabrous; base

rather broadly attached; rather thin with a thickened top part; with 3 veins,

Figure 4. Bulbophyllum fulvibulbum J.J. Verm. (114) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: abaxial

side, right: adaxial side; e. Column and lip, lateral view; f. Anther, above: abaxial side,

below: adaxial side; g. Pollinia, left two pairs, right: single pair. — All from Vermeulen

603 (living plant).

Bulbophyllum Sect. Altisceptrum and Hirtula 15

2mm

~ = y

LEZ : $<

NR Sa

iN ae = .

Figure 4. Bulbophyllum fulvibulbum J.J. Verm. (114)

16 Gard. Bull. Singapore 54 (2002)

the laterals sometimes incomplete and not reaching the base. Lip recurved

in the basal half, margins entire; glabrous or finely papillose locally; 3-

lobed; midlobe + triangular, 4.2—4.8 x 2.2—3.8 mm, index 1.2—2.2; obtuse,

rather thick; adaxially concave and with 2 knob-shaped ridges near the

base, with 2 more distinct, obtuse ridges proximally diverging and running

+ parallel from about 1/3 to 2/3 of the length of the lip, the space in

between channelled; abaxially with a retuse ridge up to about 1/2—2/3 of

the length of the lip; /ateral lobes attached along the proximal third of the

lip, retrorse, ovate to semi-circular, 1.4—1.5 mm high, rounded, margins

incurved, thin. Column 3—5S mm long. Stelidia subulate, 0.8—2:2 mm long,

acute, with an antrorse, deltoid, obtuse tooth along the upper margin.

Colours: flowers (pale) yellow with purple spots. Lip green.

Habitat & ecology: Forest, epiphytic or as a terrestrial. Alt. 700—

1300 m. Flowering in Feb, May, fruiting in Feb, Apr.

Distribution: PHILIPPINES: Luzon (1 specimen seen); Leyte (4).

INDONESIA: Irian Jaya (1).

Notes: The Irian Jaya specimen only has partly peloric flowers, with

rudiments of pollinia below the disfigured stelidia. The flowers are slightly

larger, and have slightly narrower parts than the Philippine material, but

are otherwise quite similar.

116. Bulbophyllum disjunctum Ames & C. Schweinf. — Fig. 6, plate 4.

Bulbophyllum disjunctum Ames & C. Schweinf., Orch. 6 (1920) 176; J.J. Verm., Orch.

Borneo 2 (1991) 21 (pro parte, see also under B. farinulentum and its subspecies),

colour photograph 4a. TYPE: Malaysia, Sabah, Gunung Kinabalu, Marei Parei

Spur, Clemens 254 (AMES, holo.; BM, BO, K, NY, SING, iso.).

Rhizome 5—10 mm diam., sections between pseudobulbs 3.5—8 cm

long. Pseudobulbs distant, ovoid, 1—2 x 0.8—1.5 cm. Petiole 6—16 cm.

Leaf blade elliptic to obovate, 13—28 x 3—6.5 cm, index (length/width)

2.9—7.4; acute to acuminate. /nflorescence a lax raceme, erect to patent,

19—62 cm, 10—40-flowered. Peduncle 10—23 cm; bracts 5—8, the longest

28—42 mm long. Rhachis 9—39 cm. Floral bracts elliptic to ovate, 9—20 x

1.8—4 mm, acute. Pedicel and ovary 5—12 mm long. Median sepal ovate-

triangular, 10.5—23 x 1.8—3.5 mm, index (3—)6.1—7.3; long-caudate,

margins entire or slightly erose-papillose, base broadly attached; rather

Figure 5. Bulbophyllum nemorale L.O Williams (115) — a. Habit; b. inflorescence; c.

Flower; d. Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; e.

Lip, above: adaxial side, below: abaxial side; f. Column and lip; lateral view; g. Anther,

above: adaxial side, below: abaxial side; h. Pollinia, left: single pair, right: two pairs. —

a,c, d. From Wenzel 871; b, h. From Kalkman & Tissing 4187; e—g. From Wenzel 922

(all herbarium specimens).

Bulbophyllum Sect. Altisceptrum and Hirtula

6mm

lmm

6mm

Figure 5. Bulbophyllum nemorale L.O Williams (115)

18 Gard. Bull. Singapore 54 (2002)

thin, surface glabrous, rarely slightly papillose adaxially. Lateral sepals

(11.5—)16—32 x 2.5—4 mm, index (2.8—)6.4—8.3; otherwise as the median

sepal. Petals recurved, tapering from a wide base, 7.5—16 x 1—2.5 mm,

index (3—)7—9.4; acute, margins finely erose-denticulate, rarely almost

entire, base rather broadly attached; thin, top somewhat thickened, surface

glabrous, adaxially occasionally finely papillose towards the tip; with 1

vein. Lip recurved in the basal half, ovate to triangular, 3—6 x 1—2(—3)

mm, index (1—)2—3; obtuse to acute, margins entire, finely papillose locally;

rather thin; adaxially concave near the base, with 3 ridges: 2 distinct, obtuse

to acute ridges proximally diverging and protruding like two erect to

antrorse, conical, obtuse knobs and distally running parallel up 3/4 of the

length of the lip or more, the space in between channelled and with a

rather distinct, rounded median ridge with longitudinal rows of minute

papillae proximally on its crest, adaxial surface otherwise glabrous or finely

papillose locally; abaxially with rather inconspicuous, retuse to truncate

ridge up to about 1/3 of the length of the lip, surface glabrous. Column

2.2—4 mm long. Stelidia + subulate, 1—1.8 mm long, acute, with a small,

antrorse, deltoid, obtuse to acute tooth along the upper margin; often with

a rather inconspicuous, deltoid, obtuse to subacute wing along the lower

margin, at the base.

Colours: Peduncle and floral bracts usually suffused with red,

sometimes yellowish green. Pedicel and ovary pale green. Sepals and petals

pale yellow or creamy white, usually with red dots partially aligned along

the veins. Lip yellow with a paler base, with or without red markings.

Flowers sweetly scented.

Habitat & ecology: Terrestrial or understorey epiphyte, often growing

in moss, in montane forest. Also found in lowland kerangas forest. Alt.

500-3000 m. Flowering mainly in Mar, Apr, Oct—Dec, but incidental

flowering recorded in most months.

Distribution: MALAYSIA: Sarawak (13 specimens seen); Sabah (22).

Notes: Bulbophyllum farinulentum differs from B. disjunctum in

having rounded to obtuse leaf tips, adaxially papillose to pubescent sepals,

papillose-ciliolate (rather than erose-denticulate) petals, and in lacking

erect or antrorse knobs on the adaxial surface of the lip. The colours are

also slightly different: B. farinulentum has sepals and petals with red veins,

B. disjunctum has scattered red spots, more or less aligned along the veins.

Figure 6. Bulbophyllum disjunctum Ames & C. Schweinf. (116) — a. Habit; b. Flower;

c. Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left:

abaxial side, right: adaxial side; e. Lip, left: adaxial side, right: lateral view; f. Column

and lip, lateral view; g. Anther, above: abaxial side, below: adaxial side; h. Pollinia, left

two pairs, right: single pair. —- a. From Carr 3212; b—d, f—h. From De Vogel 8218; e.

From Beaman 11128 (all herbarium specimens).

Bulbophyllum Sect. Altisceptrum and Hirtula

anernen,

2mm

Figure 6. Bulbophyllum disjunctum Ames & C. Schweinf. (116)

20 Gard. Bull. Singapore 54 (2002)

Locally in Sarawak (Hort. Leiden 914075 & 980182, from the Hose

Mts. and the Bario area respectively), a form with short, sturdy, rather

dense inflorescences and relatively short and wide flower parts is found.

The aberrant measurements are given between brackets in the description

above. See plate 4, and compare with the normal form (Vermeulen 1991,

colour photograph 4a; see above). The first specimen listed above also has

the keels on the lip with inconspicuous knobs only. This form may be

mistaken for B. farinulentum ssp. densissimum, but see below that

subspecies.

117. Bulbophyllum placochilum J.J. Verm. — Fig. 7. = (probably extinct)

Bulbophyllum placochilum J.J. Verm., Orch. Borneo 2 (1991) 29. — TYPE: Malaysia,

Sabah, Gunung Kinabalu, Clemens 34056 (L, holo.; BM, NY, iso.).

Rhizome 5—10 mm diam., sections between pseudobulbs 4—11 cm

long. Pseudobulbs distant, ovoid, 1—2 x 0.8—1 cm. Petiole 10—20 cm.

Leaf blade elliptic, 23—38 x 4.5—11 cm, index (length/width) 3.2—5S.1;

acuminate. Inflorescence a dense raceme, erect to patent, 24—36 cm, 15—

20-flowered. Peduncle 21—32 cm; bracts 7—9, the longest 30—42 mm

long. Rhachis nodding near its base, slightly thickened, 3—4.5 cm. Floral

bracts ovate, 8—10 x 2—3 mm, acute. Pedicel and ovary 6—10 mm long.

Median sepal elliptic, 11—14 x 4—5.5 mm, index 2.5—2.6; rounded to

obtuse, margins entire; glabrous, base broadly attached; rather thick. Lateral

sepals slightly falcate, 10.5—14 x 4.5—4.8 mm, index 2.3—2.9; otherwise as

the median sepal. Petals porrect, tapering from a wide base, 11—13 x 2—

2.2 mm, index 5.5—5.9; obtuse, margins entire or minutely erose towards

the tip; glabrous, base broadly attached; rather thin, somewhat thickened

towards the tip; with 1 vein. Lip recurved near the base and reflexed near

the tip, orbicular to transversely elliptic, 3—3.5 x 3.5—4 mm, index 0.7—1;

acute, margins entire; glabrous, rather thin along the median line, thin

elsewhere; adaxially slightly concave and with a central callus near the

base, with 2 distinct, obtuse ridges without protruding knobs, diverging

proximally and running from about 1/4 to 3/4 of the length of the lip, the

space in between channelled; abaxially with a retuse ridge up to about 1/2

of the lip. Column 4.5—4.8 mm long. Stelidia somewhat falcate, triangular,

1.3—2 mm long, acute, with a broadly rounded wing along the lower margin,

at the base.

Figure 7. Bulbophyllum placochilum J.J. Verm. (117) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, above: adaxial

side, below: abaxial side; e. Column and lip, lateral view; f. Anther, above: abaxial

side, below: adaxial side. — All from Clemens 34065 (herbarium specimen).

Bulbophyllum Sect. Altisceptrum and Hirtula

————

10mm NY,

5mm

Imm

-—_-——H

2mm

Figure 7. Bulbophyllum placochilum J.J. Verm. (117)

Scm

22 Gard. Bull. Singapore 54 (2002)

Colours: Flowers creamy yellow, sepals and petals minutely speckled

pale purple in the top half; lip minutely speckled pale purple except for the

top. Column white, front with two crimson streaks.

Habitat & ecology: Montane forest. Alt. 1500—1800 m. Flowering

and fruiting in Jun—Aug.

Distribution: MALAYSIA: Sabah, Gunung Kinabalu only. (3

specimens seen). Not collected since 1933.

Notes. Without flowers, this could be mistaken for a small form of B.

elongatum.

118. Bulbophyllum elongatum (Blume) Hassk. — Fig. 8.

Bulbophyllum elongatum (Blume) Hassk., Cat. Bogor. 2 (1844) 39. — Ephippium

elongatum Blume, Bijdr. (1825) 309. — Cirrhopetalum elongatum (Blume) Lind1.,

Edward’s Bot. Reg. 29 (1843) sub 49. — Phyllorchis elongata (Blume) Kuntze,

Rev. Gen. Pl. 2 (1891) 677. — TYPE: Indonesia, Java, Gunung Salak, Blume 195

(L, lecto.), 287 (L, syn.), s.n. LHB 902,322-934 (L, syn.).

Bulbophyllum gigas Ridl., J. Linn. Soc., Bot. 32 (1896) 277. - TYPE: Malaysia, Perak,

Maxwell’s Hill, Ridley (1893) (SING, holo.).

Bulbophyllum sceptrum Rchb.f., Walp. Ann. 6 (1861) 254. — Phyllorchis sceptrum

(Rchb.f.) Kuntze, Rev. Gen. Pl. 2 (1891) 678. — TYPE: Indonesia, ‘Sundaic’ Is.

(W, holo., not seen).

[Not B. elongatum sensu F. Kraenzl., Bot. Jahrb. Syst. 34 (1905) 248; J.J. Sm., Repert.

Spec. Nov. Regni Veg. 32 (1932) 310. — Specimen: Beccari PS 10239 (FI,

similar to the nominal taxon B. korinchense Ridl., sect. Aphanobulbon). |

Rhizome 5—13 mm diam., sections between pseudobulbs 3—17 cm

long. Pseudobulbs distant, ovoid, 0.8—1.6 x 0.6—1.5 cm. Petiole 5.5—22

cm. Leaf blade elliptic, 20—SO x 6—16 cm, index (length/width) 3.1-3.9;

acute to acuminate. /nflorescence a dense raceme, with the flowers in

irregular whorls, erect to patent, 24—65 cm, 30—100-flowered. Peduncle

14—53 cm; bracts 7—8, the longest 25—80 mm long. Rhachis 3—12 cm,

nodding near its base. Floral bracts ovate, 7—13 x 1.5—4 mm, acute. Pedicel

and ovary 5—6 mm long. Median sepal elliptic, 12.5—17.5 x 3.8—4 mm,

index 3.3—4.6; obtuse to acuminate, margins entire; glabrous, base rather

narrowly attached; rather thin. Lateral sepals somewhat falcate, 12—16 x

3.5—4.5 mm, index 3.2—4; acute to acuminate, otherwise as the median

sepal. Petals porrect, falcate, linear, tapering from a wide base, 10.5—15 x

Figure 8. Bulbophyllum elongatum (Blume) Hassk. (118) — a. Habit; b. Flower; c.

Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, above:

adaxial side, below: abaxial side; e. Lip, adaxial side; f. Column and lip, lateral view; g.

Anther, above: adaxial side, below: abaxial side; h. Pollinia, above: two pairs, below:

single pair. - a—d, f—h. From Hort. Leiden 26216 (spirit sample); e. From Sands 5285

(herbarium specimen).

Bulbophyllum Sect. Altisceptrum and Hirtula

Smm

2mm

Figure 8. Bulbophyllum elongatum (Blume) Hassk. (118)

Scm

24 Gard. Bull. Singapore 54 (2002)

1.2—1.8 mm, index 7.2—10; obtuse to acute, margins entire; glabrous, base

broadly attached; thin, top thickened; with 1 vein. Lip reflexed in the basal

half, triangular with a projecting, linguiform top, 2.7—3.3 x 2.4—3 mm,

index 1—1.2; rounded, margins entire; glabrous, rather thin, top thick;

adaxially flat near the base, with 3 ridges without protruding knobs: 2

weak, rounded ridges, most conspicuous distally, the space in between

channelled and with a inconspicuous, rounded median ridge with

longitudinal rows of minute papillae on its crest proximally; abaxially with

rather inconspicuous, truncate ridge up to about 1/2 of the length of the

lip. Column 4.4—5 mm long. Stelidia narrowly triangular, 1.5—2.2 mm

long, acute, with a small, antrorse, deltoid, obtuse to acute tooth along the

upper margin; with an often inconspicuous, deltoid, obtuse tooth along the

lower margin, at the base.

Colours: Sepals and petals white with purple spots. Lip yellow with

fine red dots.

Habitat & ecology: Epiphyte in damp forest, near streams; also found

as a terrestrial, creeping over the forest floor. Alt. 300—2000 m. Flowering

in Jan—Mar, Aug—Dec.

Distribution: MALAYSIA: Peninsula (7 specimens seen); Sarawak

(1); Sabah (1). BRUNEI (1). PHILIPPINES: Luzon (1); Negros (1); Leyte

(2). INDONESIA: Sumatra (1); Java (4); Irian Jaya (1). PAPUA NEW

GUINEA (2).

Notes: Plants from the Philippines are relatively small.

119. Bulbophyllum lissoglossum J.J. Verm. — Fig. 9.

Bulbophyllum lissoglossum J.J. Verm., Orch. Borneo 2 (1991) 25, colour photograph

pl. 4b; J.J. Wood et al., Plants of Mt. Kinabalu 2 (1993) 111, colour photograph

pl. 14b. — TYPE: Malaysia, Sabah, Gunung Kinabalu, Vermeulen & Chan 390

(LL, holo.; K, iso).

Rhizome 4—6 mm diam., sections between pseudobulbs 1—3 cm

long. Pseudobulbs close to distant, ovoid, 1.8—3 x 1—2.5 cm. Petiole 4—15

cm. Leaf blade elliptic, 15—27 x 2.1—4.5 cm, index (length/width) 3.8—

8.5; acute to slightly acuminate. /nflorescence a rather dense raceme, erect

to patent, 26—40 cm, 22—45-flowered. Peduncle 12—27 cm; bracts 6—7,

the longest 18—32 mm long. Rhachis erect or arching, 8—16 cm. Floral

bracts elliptic to ovate, 5.5—8 x 1.5—2 mm, acute. Pedicel and ovary 9—10

Figure 9. Bulbophyllum lissoglossum J.J. Verm. (119) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, above: adaxial

side, below: abaxial side; e. Column and lip, lateral view; f. Anther, above: adaxial

side, below: abaxial side; g. Pollinia, left: two pairs, right: single pair. — All from

Vermeulen & Chan 390 (living plant).

Bulbophyllum Sect. Altisceptrum and Hirtula Do

16mm

4mm

k——————_—_——xoIH«

3mm

Figure 9. Bulbophyllum lissoglossum J.J. Verm. (119)

26 Gard. Bull. Singapore 54 (2002)

mm long. Median sepal ovate to triangular, 8—11.2 x 2—3 mm, index 3.2—

4.5; subacute, margins entire, finely papillose locally, base rather broadly

attached; rather thick, top thickened, surface glabrous. Lateral sepals

somewhat falcate, 9—12 x 2—3 mm, index 3.3—4..5; otherwise as the median

sepal. Petals porrect, falcate, linear, 5.5—6.5 x 0.8—1 mm, index 5.2—46.5;

obtuse, margins slightly erose towards the tip; glabrous, base broadly

attached; rather thin, thickened towards the tip; with 1 vein. Lip reflexed

half-way, + triangular, 2—3.5 x 1—1.8 mm (without spreading), index 1.5—

2.4; obtuse, margins entire; glabrous; rather thick; adaxially without ridges,

surface + flat; abaxially with a weak, rounded ridge up to about 1/3 of the

length of the lip. Column 2.5—3.5 mm long. Stelidia falcate linear, 1—1.5

mm long, acute, with a small, antrorse, deltoid, obtuse to acute tooth along

the upper margin; with a broadly rounded wing along the lower margin, at

the base.

Colours: Inflorescence purple or green, suffused with purple. Bracts

pale green, spotted purple. Pedicel and ovary yellowish with some purple

spots. Sepals white, slightly greenish at the base, the median spotted purple,

the laterals only so in the basal half. Petals as the median sepal. Lip green,

median part yellowish, top part suffused with purple but extreme tip bright

green. Column white with a few purple spots. Flowers without smell.

Habitat & ecology: Understorey epiphyte, or terrestrial in moss

cushions. Montane (kerangas) forest. Alt. 1000—1900 m. Flowering in

May—Jul, Nov, Dec.

Distribution: MALAYSIA: Sabah, Gunung Kinabalu to Ulu Padas

(8 specimens seen).

Notes: The Sarawak record in Vermeulen (1991: 25) is based on the

erroneous identification of the specimen SAR (Chai) 33968, which is B.

disjunctum.

Section Hirtula Ridl.

Bulbophyllum sect. Hirtula Ridl., Mat. Fl. Mal. Penins. 1 (1908) 68; J.J. Verm., Orch.

Borneo 2 (1991) 231; Garay et al., Nordic J. Bot. 14 (1994) 628. - LECTOTYPE

(see Vermeulen, 1991: 231): Bulbophyllum hirtulum Ridl.

Bulbophyllum sect. Barbula J.J. Sm., Repert. Spec. Nov. Regni Veg. 32 (1932) 328. —

TYPE: Bulbophyllum trichoglottis Ridl. (= B. hirtulum Rid1.).

Rhizome creeping. Roots predominantly sprouting below the

pseudobulbs. Pseudobulbs 1-leafed, usually not or hardly laterally flattened,

sometimes moderately flattened (penicillium, ochthodes), wrinkling with

age in a random pattern. /nflorescence with many flowers, rarely 1-flowered

(sometimes in rariflorum). Rhachis either not swollen when compared to

Bulbophyllum Sect. Altisceptrum and Hirtula 27

the peduncle, or swollen into a spindle-shaped, cylindrical, or bilaterally

flattened body tapering towards both ends, with an inconspicuous to distinct

depression below each flower. Floral bracts recurved to reflexed. Flowers

usually spirally arranged (but [partly] distichous in tremulum, xenosum

and secundum; partly in incomplete whorls in polycyclum), not resupinate,

or resupinate (secundum), one to many open simultaneously. Pedicel and

ovary: basal node either more or less flush with the surface of the rhachis,

or on a stump up to 2 mm high. Median sepal: margins glabrous to distinctly

ciliate; 1—3-veined. Lateral sepals free, margins glabrous to distinctly ciliate,

the upper often with longer cilia than the lower; 1—5-veined. Petals +

porrect, margins entire or slightly erose, sparsely (cerebelum) to distinctly

ciliate, rarely papillose (phreatiopse); 1-veined. Lip: margins entire, usually

with 1-celled cilia, hairs or spindle-shaped appendages, rarely sparsely ciliate

(cerebellum, limbatum, rariflorum) or glabrous (phreatiopse); rather thin

to thick, often fleshy, at the base usually with two small, sometimes

inconspicuous (/indleyanum, phreatiopse, parviflorum), retrorse, semi-elliptic

to triangular, obtuse auricles flanking the ligament. Column: stigma often

protruding or with distinct teeth at its base, column foot sometimes

somewhat thickened but otherwise without accessories. Stelidia often

distinct, triangular to linear, often subulate, often with teeth along the

upper and lower margins that may be so large that they together form an

oblong structure from which only the tip of the stelidia protrude. Anther

abaxially usually with a papillose to hairy ridge near its base; frontal part

not or hardly projecting, front margin usually long ciliate, less often

papillose, rarely + glabrous (parviflorum). Pollinia 4; the inner + half as

long as the outer or longer, flattened on both sides; the outer flattened on

one side. Stipes absent.

Distribution. China (3 species), Nepal (2), Bhutan (2), India (4 species,

of which 1 is endemic), Myanmar (3), Thailand (8, 1 endemic), Vietnam (5,

2 endemic), Peninsular Malaysia (5, 2 endemic), Sumatra (7, 1 endemic),

Java (1 endemic), Borneo (19, 14 endemic), Sulawesi (1 endemic),

Philippines (4 endemic), New Guinea (1 endemic).

The use of the term ‘lip margin’. Towards the tip of the lip, the

margins may only be visible as a very thin flange or even a thin line,

usually with hairs implanted. In several species with a very thick lip this

line does not follow the full extension of the lip; but traverses the lip over

the adaxial or abaxial surface. The opposite surfaces are then described as

‘distinctly convex’. In several species a median ridge on the adaxial side

overtops the distinctly convex adaxial surface.

Delimitation of the section. Sect. Hirtula includes a suite of species

with usually rather ‘hairy’ flowers. The petals and/or the lip are indeed

ciliate or otherwise set with 1-celled hairs in all species, although only

28 Gard. Bull. Singapore 54 (2002)

sparingly so in some. Moreover, a series of species (B. eublepharum Rchb..,

B. insulsoides Seidenf., B. levinei Schltr, and B. scabratum Rchb.f.),

seemingly constitute a grading of the section into racemose species with a

less ‘hairy’ aspect. An extended character set is necessary to delimitate

sect. Hirtula:

a. Pseudobulbs wrinkling in a random pattern with age. Occurs in all

species of sect. Hirtula. It also occurs elsewhere, but not in the

species listed above with B. eublepharum; these all have pseudobulbs

wrinkling along pre-determined longitudinal lines.

b. Inflorescence racemose. Occurs in all species of sect. Hirtula, although

in B. rariflorum the inflorescence may abort after producing a single

flower.

C. Petals ciliate with 1-celled hairs. Occurs in all species, although B.

cerebellum has only sparsely ciliate petals, and B. phreatiopse has

papillose rather than ciliate petals. Elsewhere, ciliate petals are found

in B. eublepharum, but this character is rarely found among SE

Asian racemose Bulbophyllum. Species in and around sect.

Cirrhopetalum all have fimbriate petals (with many-celled hairs).

d. Lip with long hairs. Occurs in most species, although B. cerebellum,

B. hirtulum, B. limbatum, and B. rariflorum, have only sparsely hairy

lips, and B. phreatiopse has a glabrous lip. A long-haired lip is a rare

character in other SE Asian racemose Bulbophyllum (present in,

for instance, B. refractum Rchb.f. from Java).

é: Two small auricles at the base of the lip, one on each side of the

ligament connecting the lip to the column foot. These occur in all

species, but are inconspicuous in B. hirtulum, B. lindleyanum, B.

phreatiopse, and B. parviflorum. This character is unique within SE

Asian racemose Bulbophyllum, although it occurs in some sections

with 1-flowered species, for instance in a series of New Guinean

species of sect. Polyblepharon.

If a polythetic set of a., b., and e. is chosen as diagnostic, this allows

the exclusion of the species listed with B. eublepharum above, because

these lack two out of the three characters. |

Bulbophyllum phreatiopse displays two out of the three diagnostic

characters, but its petals have papillose rather than ciliate margins, and the

lip is glabrous. It does not fit well in sect. Hirtula, and has been included

with doubt.

Bulbophyllum hastiferum Schltr, B. pubiflorum Schltr, B. crassissimum

J.J. Sm., and B. uvaeflorum O’Byrne have shortly ciliate petals. Because

the lateral sepals form a disc or spoon-shaped structure below the lip,

these are better included in sect. Osyricera. O’Byrne originally included

Bulbophyllum uvaeflorum in sect. Hirtula.

Bulbophyllum Sect. Altisceptrum and Hirtula 29

Several other similar-looking SE Asian species do not display the

character set mentioned and are therefore excluded. Bulbophyllum

inconspicuum Maxim. has fimbriate petals with many-celled hairs; B.

dayanum Rchb.f. and similar species have ciliate petals, as well as two

lateral lobes near the base of the lip. These lateral lobes, however, are

attached in a much more distal position and are therefore not considered

to be homologous with the auricles in sect. Hirtula.

Bulbophyllum vanum J.J. Verm. from Africa, as well as several

Madagascar species included in the sections Micromonanthe,

Panthoblepharon, and Trichopus, have ciliate petals, or a ciliate lip, or

both. After analysing several species (B. microglossum Perr., B. muscicolum

Schltr, B. nigriflorum Perr., and B. pleurothallopsis Schltr), I assume that

none have auricles similar to the ones in sect. Hirtula.

Monophyly. The diagnostic set of characters is sufficiently sturdy to

assume monophyly a-priori, possibly after exclusion of Bulbophyllum

phreatiopse. For an outgroup, the first choice would be the series mentioned

above with B. eublepharum.

Taxonomic views on sect. Hirtula. Ridley lists four taxa under his

sect. Hirtula: B. tenerum Ridl. [a species close to B. planibulbe (Ridl.)

Ridl.], B. hirtulum Ridl., B. cincinnatum Ridl. (is B. hirtulum), and B.

limbatum Lindl. His section includes species with ‘hairy’ flowers (except

for the probably erroneously included B. tenerum), with an elongated or

subumbellate inflorescence. This general idea was retained in Vermeulen

(1991). Garay et al. (1994: 628) state that only Bulbophyllum carinilabium

and B. hirtulum (with some of its synonyms listed as separate taxa) can be

included because of the chosen lectotype species of sect. Hirtula. They do

not offer any further explanation, and I can only assume that the presence

of a subumbellate inflorescence, present in both species, has been given

priority over other characters in the delimitation of sect. Hirtula. The

subumbellate B. gemmareginae and B. cerebellum, however, show that

several flower morphologies found in the series with an elongated

inflorescence are repeated in the series with a subumbellate inflorescence.

I prefer a wider circumscription of sect. Hirtula, as proposed here, because

it is more likely to result in a monophyletic group (see above).

Species delimitation. Many species of sect. Hirtula appear rare in

existing collections; they are represented by only a few samples, or even by

a single one. This puts a limit to the reliability of this revision, because a

proper delimitation of the species was difficult and infraspecific variability

could not be assessed with the material now available. So far, only

Bulbophyllum hirtulum displays variability exceeding the average. Some

species pairs have been found that could well be sister species: B.

lasioglossum-—B. negrosianum, and B. jolandae—B. comberipictum. The three

30 Gard. Bull. Singapore 54 (2002)

species B. aithorhachis—B. sororculum—B. setuliferum are rather similar

and may be closely related. Their status may need revision once more

material of all three becomes available, although the moderately rich

material now available of B. setuliferum displays no signs of grading into

any of the others. All taxa assumed to be part of a group of sister species

have been given specific rank, ad hoc considerations as to the phylogeny

of the group have not entered the taxonomy of the revision.

Cultivation. About 25 species have recently been in cultivation. Most

are plants from the understorey of primary tropical forests, and have been

grown successfully under conditions usually recommended for

Bulbophyllum: a substratum that never dries out entirely, semi-shade, high

air humidity and good air movement. A fair number of plants have been

lost because of bacterial rot, turning the victims from green to brown

within a few days. Proper ventilation or even a permanent, rather strong

airflow over the plants seems to prevent this to a certain extent. It appears

that several species found at medium altitudes do well in lowland conditions

as long as shading and a continuous airflow keep the plants cool, although

they are, of course, best grown at a temperature similar to that in their

natural habitat. Species with long rhizomes, like B. limbatum and B.

rariflorum, grow well on thin slabs of fern root, a tree fern trunk or an

upright wooden support. Most other species prefer some soft organic matter

among their roots, either planted on a slab or in a pot. Flowering is often

irregular, and years may pass before plants decide that it is time to please

their owner. On the whole, most species are interesting rather than

spectacular, although B. gemmareginae and B. xenosum are striking plants

with flowers of delicate colours and shapes.

Key to Species of Bulbophyllum sect. Hirtula

1. Inflorescence subumbellate (rhachis half as long as the width of the

inflorescence, or shorter); or inflorescence with 1 flower, and

pseudobulbs touching

2., Lateral sepals reflexed. upper smarcin. lone ciliate... 2

LP LS TRICE LY INE. MOR ITE Serie 120. B. gemma-reginae

2. Lateral sepals somewhat recurved; upper margin glabrous to finely

papillose or ciliolate

3. Lip distally with mobile c. 2 mm long, flat appendages along the

EE att 7 ee 2 SO I A RRS DI He 121. B. carinilabium

3. Lip distally ciliate, without flat appendages

4. Lateral sepals acute to acuminate. Lip without lateral lobes

(next to inconspicuous basal auricles)............ 122. B. hirtulum

Bulbophyllum Sect. Altisceptrum and Hirtula 31

4. Lateral sepals rounded. Lip with lateral lobes attached along

the proximal thind ofits dength......04.....1.0. 123. B. cerebellum

1. Inflorescence an elongated raceme; or inflorescence with 1 flower, and

at least some pseudobulbs widely distant

5. Rhachis spindle shaped, or otherwise swollen when compared to the

peduncle: (in some species difficult to see in herbarium material)

6. Petal index (length /width) 0.6—1.6; petals reniform, (transversely)

elliptic to ovate

7. Lip ovate-subtriangular, distally rather thin and tapering

towards a subacute tip

Pseudobulbs discoid. Lip ovate in the proximal third, then

abruptly narrowed and drawn out into a slender, tapering

top part. Lip margins only with long, thin, wavy hairs .......

Pin dty ae itlseniecreheee cde UR eS La) 124. B. clipeibulbum

. Pseudobulbs ovoid. Lip ovate-subtriangular, gradually

tapering towards the tip. Lip margins with stiff hairs, some

of these slightly swollen distally ............ 125. B. penicillium

7. Lip subspathulate, obovate, elliptic, or oblong, distally very

thick and with a truncate to rounded tip (tip defined by the

line along which the marginal hairs are inserted, excluding

crests if present)

Median ridge on the adaxial side of the lip protruding

beyond the tip of the lip as a high crest

10. Inflorescence + erect, or with the peduncle patent, and

the rhachis (slightly) nodding. Lip distally glabrous on

the adaxial side (hairs implanted along the margins may

cover adaxial surface)

11. Rhachis, pedicel and ovary without blackish dots.

Petal index (length/width) 0.6—0.7. Lip concave near

the tip (a depression present on both sides of the

MS IAM SIAC ec sd ontecdaceoardgnwed «nobis 126. B. setuliferum

11. Rhachis, pedicel and ovary with minute blackish dots.

Petal index 1.2—1.6. Lip not concave near the tip ..

BPMs OAL. coh Rp MIL oe YAEL dca: 127. B. sororculum

10. Inflorescence entirely pendulous. Lip distally irregularly

hirsute on the adaxial side.............. 128. B. aithorhachis

9. Median ridge on the adaxial side of the lip merging with

the lateral ridges well before the tip of the lip, not protruding

beyond the tip of the lip

12. Pedicel and ovary with minute blackish dots. Rhachis c.

5 mm thick. Longest hairs on lip straight, slightly club-

SEC abe csshe the: -ehetemcctee--Pedun gas 129. B. echinochilum

32 Gard. Bull. Singapore 54 (2002)

12. Pedicel and ovary without blackish dots. Rhachis c. 1.5

mm thick. Longest hairs on lip wavy, thin ...............0...

iii. ee eee 130. B. lanuginosum

6. Petal index more than 1.6, or petals hastate

13.Lip abruptly narrowed at c. 1/3 of its length, then drawn out

into a slender, tapering top part. Lip without a median ridge

towards:the ip:.;.:a.s..0dd sabes dee gt 124. B. clipeibulbum

13.Lip rounded at the tip; or lip with a median ridge overtopping

the rounded front

14.Column foot with an obtuse, papillose spur protruding from

between the base of thedateralsepdlsiimedj nbchace. Digh

14.Column foot without a spur between the lateral sepals

15. Petal tips acute

16. Rhachis with the top 2—7 cm with only minute, sterile

bractssand. noflewetssasent.oindes 131. B. atratum

16. Rhachis with flowers to the very tip

17. Whole inflorescence pendulous. Rhachis 2—3 mm

thick +2.43)...t42ener. Aee-: 132. B. groeneveldtii

17. Peduncle erect or patent, rhachis nodding. Rhachis

6—-7 mm thick «...;/¢ecesags 133. B. grotianum

15. Petal tips rounded to obtuse

18. Rhachis distinctly flattened, the top 3—3.5 cm with

only minute, sterile bracts and no flowers

19.Sepals abaxially with scattered coarse papillae.

Lateral sepals ovate, subacute. Lip adaxially finely

hirsute: .leogteye-deteebs. 2evirp. sci... 134. B. Janus

19.Sepals abaxially glabrous. Lateral sepals

triangular, acuminate. Lip adaxially glabrous

except for the finely papillose top part ...............

AR AONE Fo LY RT RS ekornuebae 135. B. xiphion

18. Rhachis + circular in section, with flowers to the

very tip

20. Flowers dimorphic: flowers along the basal part

of the rhachis not much opening, with the lip

1.8—2.3 mm long; flowers along the top part

opening wide, with the lip c. 4.2 mm long (flowers

at the very tip of the rhachis smaller and with a

smaller lip) :acth: dosti 136. B. mirabile

20. Flowers all of + the same shape

Bulbophyllum Sect. Altisceptrum and Hirtula 33

21. Lip adaxially with a distinct, narrow median

ridge embedded in a furrow over the distal 2/3

of the lip, and overtopping its tip as a high

POESOCIN IR. OLS RAD 127. B. sororculum

21. Lip adaxially flat or convex towards the tip,

but without median ridge

22. Lip adaxially papillose. Pseudobulbs

SIS TAME LR 137. B. spadiciflorum

22. Lip adaxially glabrous or irregularly

wrinkled. Pseudobulbs touching

23. Rhachis 1—1.8 cm long; flowers in a

very dense raceme. Lip with lateral

lobes attached along the proximal third

ofats 1éngiieareus.: 123. B. cerebellum

23. Rhachis 33—36 cm long, flowers in a

rather lax raceme. Lip without lateral

lobes (next to basal auricles flanking

the ligament)........ 138. B. polycyclum

5. Rhachis about as thick as the peduncle

24.Sepals adaxially partly or entirely pubescent or pilose

25. Rhachis, pedicel, ovary, and abaxial surface of sepals densely

hispid with short, stiff hairs ....... ee. 139. B. lindleyanum

25. Rhachis, pedicel, ovary and abaxial surface of sepals glabrous

26.Lip abaxially with numerous spindle-shaped hairs arising

Leola DIN Pits Cass MOP LN 140. B. tremulum

26.Lip abaxially glabrous, pubescent or velutinous

27.Lip pouch-shaped, deeply concave on the adaxial side

BIGGAR TAR ERAS, BL tL 141. B. scaphiforme

27.Lip only somewhat concave near the base on the adaxial

side

28.Lip 2.8—4.2 x 1—1.6 mm, abaxial surface convex

towards the tip. Column 1—1.3 mm long..................

WE. AU QUEA-TAM) AO Bie. Ue 142. B. nigripetalum

28.Lip 5—7 x 2—3.3 mm, abaxial surface + flat to

concave towards the tip. Column 1.4—1.8 mm long

PAS RIEL. UG Boh lcd he 143. B. nigrescens

24. Sepals adaxially glabrous to finely papillose (margins often ciliate)

29. Petal index (length/width) 3—4.6

30. Lip 1.7—3 mm long (without stretching)

31.Sepals abaxially hirsute. Lip adaxially with a circular

Ge pression: Near TNE LIP s.0ci-,.t0seceneccess 144. B. trulliferum

31.Sepals abaxially glabrous. Lip adaxially convex near the

tip

34 Gard. Bull. Singapore 54 (2002)

32.Rhizome sections between pseudobulbs 0.5—1.8 cm

hehe 2s. Riis Be ND ee 154. B. secundum

32.Rhizome sections between pseudobulbs 2—10 cm

LOG kid LEE 153. B. parviflorum

30. Lip 4—7.5 mm long

33.Anthesis starting at the base of the rhachis. Median

sepal triangular, tapering towards the tip or caudate,

margins glabrous or finely papillose

34. Pedicel and ovary 48—50 mm long. Median sepal c.

18.5 mm long. Lip margin with the longest hairs +

Siaight Stig ASE. ARSE SoS es 145. B. jolandae

34. Pedicel and ovary 8—20 mm long. Median sepal c. 7

mm long. Lip with the longest hairs distinctly wavy,

Cig ieee tes ta 2 ra 146. B. comberipictum

33.Anthesis starting at the top of the rhachis. Median sepal

elliptic, acute to shortly acuminate, margins ciliate

35.Lip adaxially with a median ridge continuing over c.

1/3 of the lip. Pedicel and ovary 7.5—14 mm long.

Median sepal 6.5—9 mm long.. 147. B. lasioglossum

35.Lip adaxially with median ridge continuing over c. 3/4

of the lip. Pedicel and ovary 5.5—6.5 mm long.

Median sepal 5.5—5.8 mm lOng..........eeeeeeeeeeeeeeeeeeees

Sie tets bade Bes wees ese 148. B. negrosianum

29.Petal index less than 3

36.Lip adaxially with 2—4 rows of distinct warts (calli)

stretching over most of its length

37. Pseudobulbs touching. Lip with slightly club-shaped

hairs. Stelidia with a strap-shaped to somewhat

spathulate tooth with a 2—3-furcate tip along the lower

mare, near the Bases: !.220 Se 149. B. dracunculus

37.Pseudobulbs distant. Lip without club-shaped hairs.

Stelidia with a deltoid to triangular, obtuse or acute

tooth along the lower margin, near the tip

38. Lip 6—6.8 mm long, adaxially with 4 rows of

irregularly shaped calli ......... 150. B. polygaliflorum

38. Lip c. 3.1 mm long, adaxially with 2 rows of

irregularly shaped calli... 151. B. ochthodes

36.Lip adaxially without calli, or with a single callus near its

base

Bulbophyllum Sect. Altisceptrum and Hirtula 35

39. Lip adaxially near the base with a triangular callus

40. Rhizome sections between pseudobulbs 4—19 cm

long. Margins of lip glabrous.... 152. B. phreatiopse

40. Rhizome sections between pseudobulbs 0.5—1.8 cm

long. Margins of lip ciliate ........... 154. B. secundum

39. Lip adaxially near the base without a callus

41. Lip 1.7—3 mm long (without stretching)

42. Rhizome sections between pseudobulbs 2—10

ET FOE rece necneac tte ete cinr che 153. B. parviflorum

42. Rhizome sections between pseudobulbs 0.5—1.8

CUS MOUS Menace Seteteat encase cern 154. B. secundum

41. Lip 3.5—7.5 mm long

43. Lip margins with long hairs

44.Lip margins glabrous half-way. Petals

triangular-ovate.............. 155. B. debrincatiae

44. Lip margins with hairs from base to tip. Petals

obovate

45.Median sepal index (length/width) 2.3—

ZA, WEP WIL MONS WAITS ........006-0.20-02-2-

2S ge pct tem thre. See andl 156. B. pilosum

45.Median sepal index 7—8, margins ciliate

Rvp ded Rimubeetin mdash a: 157. B. xenosum

43. Lip margins papillose-ciliolate

46. Floral bracts triangular, not clasping the

rhachis. Inflorescence with many flowers open

SMMUGETICOUSLY 222-2. ..82..2hen--- 158. B. limbatum

46. Floral bracts tubular, clasping the rhachis.

Inflorescence with a single flower open only;

each flower developing only after anthesis of

ie Previ0Us ONO L....o6..:.-s; 159. B. rariflorum

36 Gard. Bull. Singapore 54 (2002)

120. Bulbophyllum gemma-reginae J.J. Verm. — Fig. 10, plate 5.

Bulbophyllum gemma-reginae J.J. Verm., Blumea 41 (1996) 21. — TYPE: Malaysia,

Sabah, Ulu Padas, Jongejan cult. 1856 (L, holo.).

Rhizome c. 1.3 mm diam., sections between pseudobulbs 0.2—0.7 cm

long. Pseudobulbs touching, aoe to + globose, 0.5—1.2 x 0.3—0.7 cm.

Petiole 0.3—0.4 cm. Leaf blade elliptic, 5—6.7 x 1.5—3.3 cm, index (length/

width) 1.9—4.1; subacute. Jnflorescence a subumbellate raceme, patent to

+ pendulous, c. 3 cm, 7—10-flowered. Peduncle c. 2.7 cm; bracts c. 6, the

longest c. 3 mm long. Rhachis porrect or arching, spindle-shaped, c. 0.3 x

0.12 cm. Floral bracts triangular, 0.8—1.2 x 0.6—0.8 mm, acute to shortly

acuminate. Flowers opening wide, many simultaneously. Pedicel and ovary

7.5—10 mm long, basal node on a c. 0.5 mm-long stump. Median sepal

reflexed, triangular, c. 5.8 x 1.9 mm, index c. 3; acute, margins long ciliate,

base broadly attached; thin, surface glabrous. Lateral sepals ovate, c. 6.3 x

2.7 mm, index c. 2.3; subacute, otherwise as the median sepal. Petals porrect,

ovate, c. 4.3 x 1.2 mm, index c. 3.6; acuminate, margins ciliate, base rather

broadly attached; thin, surface shortly pubescent towards the tip. Lip

abruptly recurved near the base, spathulate, c. 5 x 1.4 mm, index c. 3.6;

rounded, margins ciliate with the cilia longest c. half-way up the lip; thick,

surface + glabrous; adaxially slightly concave and with two calli near the

base, with a median slit up to c. 1/3 of the lip, with two distinct, obtuse

ridges proximally diverging and distally continuing parallel and close

together over 2/3—3/4 of the lip, with the distal half slightly concave;

abaxially with a distinct, rounded ridge near the base, elsewhere + flat.

Column c. 1.1 mm long, stigma protruding at its base, column foot with

two longitudinal, deltoid, rounded lamellae. Stelidia falcate, c. 0.8 mm long,

acute, + rectangular in outline with a large, antrorse, triangular, acute

tooth along both margins. Anther: front margin ciliate, abaxial ridge +

glabrous.

Colours: Sepals creamy yellow, suffused with pale reddish brown

towards the tip. Petals creamy yellow, with almost black lines bordering

the midvein and the margins. Lip white, ridges on adaxial side greyish with

minute white specks, top part with greyish black margins, hairs yetloyagy.

Column yellow, foot with reddish purple spots.

Habitat & ecology: Understorey epiphyte in lower montane forest.

Alt. c. 1100 m.

Figure 10. Bulbophyllum gemma-reginae J.J. Verm. (120) — a. Habit; b. Flower, including

rhachis; c. Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; d.

Lip, above: adaxial side, below: abaxial side; e. Column and lip, lateral view; f. Anther,

above: abaxial side, below: adaxial side; g. Pollinia, left: single pair, right: two pairs. —

All from Jongejan cult. 1856 (spirit sample).

Bulbophyllum Sect. Altisceptrum and Hirtula

GUase

SSS

sisaze ‘eat eles Shy

SSS

Figure 10. Bulbophyllum gemma-reginae J.J. Verm. (120)

38 Gard. Bull. Singapore 54 (2002)

Distribution: MALAYSIA: Sabah, Ulu Padas (1 specimen seen).

121. Bulbophyllum carinilabium J.J. Verm. — Fig. 11, plate 6.

Bulbophyllum carinilabium J.J. Verm., Orch. Borneo 2 (1991) 233. — TYPE: Malaysia,

Sabah, Ulu Padas, Vermeulen 641a (L, holo.; K, iso.).

Rhizome 2.5—4 mm diam., sections between pseudobulbs 0.4—1 cm

long. Pseudobulbs (almost) touching, ovoid to + globose, 0.8—1.8 x 0.5—

1.2 cm. Petiole 0.3—2.3 cm. Leaf blade elliptic to ovate, 5.5—15 x 2.5—5

cm, index (length/width) 2—5.4; acute. Inflorescence a subumbellate raceme,

4—16.4 cm, 4—11-flowered. Peduncle patent to + pendulous, 3.7—15.7

cm, bracts 3, the longest 3.5—6 mm long. Rhachis porrect or arching,

spindle-shaped, 0.3—0.7 x 0.1—0.18 cm. Floral bracts triangular, 1—1.3 x

0.6—0.8 mm, subacute. Flowers not much opening, few to many

simultaneously. Pedicel and ovary 3.5—5 mm long, basal node almost flush

with the surface of the rhachis. Median sepal porrect, ovate, 7—9 x 1.5—

2.8 mm, index 2.8—6; (sub)acute, margins ciliate, base rather broadly

attached; thin, surface glabrous. Lateral sepals somewhat recurved, 7—10 x

2—3 mm, index 2.3—5; obtuse to acute, margins glabrous; rather thin,

otherwise as the median sepal. Petals porrect, triangular, 2—2.2 x 1—1.3

mm, index 1.6—2; (sub)acute, margins ciliate, base broadly attached; thin,

surface glabrous. Lip + straight but with a deep transverse fold near the

base, elliptic to ovate with a narrow claw, 4.8—5.7 x 1.5—2.8 mm, index

1.9—3.8; rounded to obtuse, margins ciliate but with mobile, c. 2 mm long,

linear, flat appendages near the tip; thick; adaxially concave near the base,

with a narrow median ridge in the proximal 1/3, with two distinct calli

clasping its distal end, with the distal 2/3 flat or slightly concave but with a

small callus near the tip, with the surface glabrous except for a scabrous

median strip over the distal 2/3; abaxially with a weak, rounded ridge over

the distal three-quarters, surface glabrous. Column 1.6—2 mm long, column

foot with two longitudinal, subtriangular lamellae clasping the median ridge

on the adaxial side of the lip. Stelidia semi-elliptic, 0.7—1 mm long, rounded.

Anther: front margin densely papillose with elongate papillae, abaxial ridge

+ glabrous.

Colours: Sepals purple, base whitish or yellow with purple veins.

Petals purple. Lip dark purple, sometimes yellowish green towards the

base; the cilia white, the appendages near the tip purple or yellowish with

Figure 11. Bulbophyllum carinilabium J.J. Verm. (121) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial

side, right: abaxial side; e. Column and lip, lateral view; f. Anther, left: abaxial side,

right: adaxial side; g. Pollinia, above: two pairs, below: single pair. — All from Vermeulen

641a (living plant).

Bulbophyllum Sect. Altisceptrum and Hirtula 39

Simm

3mm

ees. Se See a et POZE

» lmm

‘

Figure 11. Bulbophyllum carinilabium J.J. Verm. (121)

40 Gard. Bull. Singapore 54 (2002)

purple margins.

Habitat & ecology: Understorey epiphyte in montane forest. Alt.

1500—3600 m. Flowering in Mar—May, Oct.

Distribution: MALAYSIA: Sabah, Gunung Kinabalu to Ulu Padas

(5 specimens seen).

122. Bulbophyllum hirtulum Ridl. — Fig. 12.

Bulbophyllum hirtulum Ridl., J. Bot. 38 (1900) 71, J.J. Verm., Orch. Borneo 2 (1991)

237, colour photograph pl. l6e. — TYPE: Malaysia, Penang Hill, Ridley &

Curtis s.n. (K holo.).

Bulbophyllum cincinnatum Ridl., J. Straits Branch Roy. Asiat. Soc. 39 (1903) 75. —

TYPE: Malaysia, Perak, Batu Tujoh, Ahmad s.n. (K, holo.).

Bulbophyllum trichoglottis Ridl., J. Fed. Malay States Mus. 4 (1909) 66. — TYPE:

Malaysia, Pahang, Telom, Hort. Singapore (Ridley) 1909 (SING, holo.)

Bulbophyllum trichoglottis Ridl. var. sumatranum J.J. Sm., Bull. Jard. Bot. Buitenzorg,

s. 3, 2 (1920) 98. - TYPE: Indonesia, Sumatra, Barisan Range, Ajoeb (Exp.

Jacobson) 807 (BO, holo.; K, L, SING, iso.).

Bulbophyllum salaccense sensu Beaman, T.E., et al., Orch. Sarawak (2001) colour

photograph pl. 7c, non Rchb.f.

Rhizome 1.2—2.5 mm diam., sections between pseudobulbs 0.15—

0.7 cm long. Pseudobulbs touching, ovoid to + globose, 0.2—1 x 0.2—0.7

cm. Petiole 0.3—2.2 cm. Leaf blade elliptic to obovate, 2.2—17 x 0.7—6

cm, index (length/width) 2—3.5; obtuse to acute. Inflorescence hidden below

the leaves, a subumbellate raceme, 4.5—10 cm, (1—)2—14-flowered.

Peduncle patent to + pendulous, 4.3—9.5 cm, bracts 4—5, the longest 3—7

mm long. Rhachis porrect or arching, spindle-shaped, 0.3—0.7 x 0.1—0.18

cm. Floral bracts triangular, 1—1.3 x 0.6—0.8 mm, subacute. Flowers not

much opening, many simultaneously. Pedicel and ovary 2—4 mm long,

basal node on a 0.2—0.5 mm-long stump. Median sepal porrect, sometimes

reflexed, ovate to triangular, 4.5—6.5 x 1.8—3.3 mm, index 1.4—2.5; acute

to acuminate, margins finely papillose to ciliolate; base broadly attached;

rather thin, adaxial surface glabrous, abaxial surface glabrous or finely

papillose. Lateral sepals somewhat recurved, 4—6 x 2.5—5 mm, index 1.1—

2.2; otherwise as the median sepal. Petals porrect, ovate, 2—4 x 0.7/—2

mm, index 2.1—3.6; obtuse to acute, margins ciliate, base broadly attached;

thin, surface glabrous or adaxially slightly papillose towards the tip. Lip

Figure 12. Bulbophyllum hirtulum Ridl. (122) — a. Habit; b. Flower; c. Flower analysis,

from left to right: median sepal, petal, lateral sepal, lip; d. Lip, above: adaxial side,

below: abaxial side; e. Column and lip, lateral view; f, g, h. Lip, adaxial side; 1, j, k.

Column, lateral view; |. Anther, left: adaxial side, right: abaxial side; m. Pollinia, left:

single pair, right: two pairs. - a—e, 1, m. From TOC cult. 1203 (living plant); f, 1. From

Carr s.n.; g,}. From Chan s.n.; h, k. From Lewis 29] (all spirit samples).

Bulbophyllum Sect. Altisceptrum and Hirtula 41

Imm

Figure 12. Bulbophyllum hirtulum Ridl. (122)

42 Gard. Bull. Singapore 54 (2002)

without lateral lobes, abruptly recurved in the proximal half or half-way,

elliptic to obovate, 2.6—4 x 1.5—2.8 mm, index 1.3—2.6; rounded,

sometimes apiculate, margins ciliate; thick; adaxially (slightly) concave and

with 1—2 calli near the base, with or without a median slit up to c. 1/2 of

the lip, with two inconspicuous to distinct, proximally diverging, obtuse

ridges, with the surface distally convex, with or without two tiny pits close

to the tip, with the surface smooth to coarsely, often transversely wrinkled,

often finely papillose all over or with a patch of hairs or papillae distally

along the median line; abaxially with a retuse to rounded ridge near the

base, surface distally convex, smooth to coarsely wrinkled; basal auricles

inconspicuous. Column 1.2 column foot often with two

weak, longitudinal ridges. Stelidia semi-elliptic or triangular, 0.6—0.9 mm

long, rounded to acute, if triangular with a patent to antrorse, semi-circular

to triangular, rounded to acute tooth along the upper margin, and with a

similar tooth along the lower margin, sometimes with a second, retrorse

tooth along the lower margin. Anther: front margin densely ciliate or

papillose with elongated papillae, abaxial ridge + glabrous.

Colours: Sepals whitish, creamy yellow, or yellowish green; margins,

veins and sometimes transverse veins reddish purple. Petals the same, but

without coloured transverse veins. Lip dark red purple, white near the

base, and adaxially with a white, cream, or pink strip along the midvein.

Flowers sometimes faintly scented.

Habitat & ecology: Understorey epiphyte in mixed dipterocarp forest

and kerangas forest. Alt. 200—1400 m. Flowering in Jan—Mar, Jul, Sep,

Oct.

Distribution: MALAYSIA: Peninsula (7 specimens seen); Sarawak

(5); Sabah (3). INDONESIA: Sumatra (2); Kalimantan (1).

Notes: Bulbophyllum hirtulum shows infraspecific variability in the

following characters:

- The median sepal may be reflexed instead of porrect (Sarawak).

- The lip may have a patch of hairs distally on the adaxial side of the lip

(Sarawak).

- The lip may have two tiny pits near the tip, on the adaxial side (Sarawak,

Peninsular Malaysia). The wall separating the pits may develop into a

triangular, acute tooth forming a hood over the pits (Peninsular Malaysia).

- The stelidia may have a triangular, subacute tooth along their lower

margin (Peninsular Malaysia, Sarawak, and Sumatra).

- The median slit on the adaxial side of the lip may extend over c. 1/5 of

the length of the lip or more.

- The upper and the lower surface of the lip are entirely smooth, without

corrugations (Sabah).

All these characters occur single, or in any number, and in various

Bulbophyllum Sect. Altisceptrum and Hirtula 43

combinations. Therefore, they cannot be used to distinguish infraspecific

taxa.

123. Bulbophyllum cerebellum J.J. Verm. — Fig. 13, plate 7.

Bulbophyllum cerebellum J.J. Verm., Blumea 41 (1996) 350. - TYPE: ‘Borneo’ no

further details known, Giles 998 (K, holo.).

Rhizome 2—2.5 mm diam., sections between pseudobulbs 0.4—1.3

cm long. Pseudobulbs touching, ovoid, 1.2—2.5 x 0.7—1.2 cm. Petiole 1.5—

2.5 cm. Leaf blade elliptic to (ob)ovate, 5—14 x 1.5—3.5 cm, index (length/

width) 3.3—4; acute. /nflorescence a very short and dense or a subumbellate

raceme, 4—12 cm, 10—29-flowered. Peduncle patent to pendulous, 3.8—

10 cm, bracts c. 5, the longest 3—6 mm long. Rhachis pendulous, spindle-

shaped, 1—1.8 x c. 0.3 cm, glabrous. Floral bracts triangular, 1—1.5 x 0.8—

1 mm, acute. Flowers not much opening, many simultaneously. Pedicel and

ovary 2.6—3.5 mm long, basal node on a 0.2—0.3 mm-long stump. Median

sepal + porrect, triangular, 3.6—4.6 x 2—3 mm, index 1.2—1.9; acute,

margins with scattered very fine papillae, base rather broadly attached;

rather thin, surface glabrous. Lateral sepals somewhat recurved, 4—5.5 x

3.2—4.3 mm, index 1.2—1.3; rounded, upper margin glabrous, lower

ciliolate, base broadly attached; otherwise as the median sepal. Petals

porrect, ovate, 2.8—3.2 x 1.5—1.9 mm, index 1.5—1.9; rounded, margins

sparsely and very finely ciliate, base rather narrowly attached; rather thin,

surface glabrous. Lip abruptly recurved c. half-way, 3-lobed; midlobe +

elliptic, 2.6—3.4 x 1.4—3 mm, index 1.4—1.9; rounded, margins shortly

and sparsely ciliate towards the tip; very thick, surface irregularly wrinkled

except for the glabrous basal part; adaxially concave and with a short

median slit near the base, with two distinct erect ridges diverging proximally,

continuing over c. 1/3 of the lip and distally truncate, surface distally

distinctly convex; abaxially with a wide, rounded ridge towards the base,

distinctly convex towards the tip; lateral lobes attached along the proximal

1/3 of the lip, antrorse, falcate, ovate, rounded, entire, glabrous, broadly

attached, rather thin. Column 1—1.5 mm long, column foot somewhat

swollen, abaxially without a spur near the tip. Stelidia 0.7—1 mm long,

acute, + rectangular in outline, with or without a distinct, antrorse, deltoid,

obtuse tooth along the upper margin and with a similar, obtuse to acute

tooth along the lower margin. Anther: front margin papillose with elongated

papillae, abaxial ridge + glabrous.

Colours: Peduncle purplish or dark red, rhachis green. Pedicel and

ovary yellowish green. Sepals yellow or yellowish green, with some brownish

stains towards the tip or with brown veins. Petals yellow, with pale purple

or brownish spots towards the tip. Lip bright yellow, base and lateral lobes

44 Gard. Bull. Singapore 54 (2002)

pale greenish, sometimes with brownish stains. Column yellow.

Habitat & ecology: Epiphyte in primary forest. Alt. c. 1300 m.

Flowering in May, Oct.

Distribution: MALAYSIA: Sabah (3 specimens seen).

124. Bulbophyllum clipeibulbum J.J. Verm. — Fig. 14, plate 8.

Bulbophyllum clipeibulbum J.J. Verm., Malayan Orch. Rev. 35 (2001) 51. — TYPE:

no locality, Hort. Salzburg OR 637/94 (SZU, holo.; SING, iso.).

Rhizome 5—6 mm diam., sections between pseudobulbs 3.5—4.5 cm

long. Pseudobulbs touching, dorsiventrally discoid, 1.0—1.8 x 3.5—5.5 cm.

Petiole 38—60 cm. Leaf blade elliptic-ovate, 14.5—22 x 4—6 cm, index

(length/width) 2.8—4.4; acute. Inflorescence an elongated, lax raceme, c.

50-flowered. Rhachis arching or nodding, spindle-shaped, c. 19 x 0.7 cm (at

an early stage of anthesis), slightly and finely rugose-verruculate. Floral

bracts ovate to triangular, 4.5—S x c. 1.5 mm, acute. Flowers not fully

opening, several simultaneously. Pedicel and ovary c. 3.6 mm long, basal

node on a 0.2—0.4 mm-long stump. Median sepal recurved at the tip,

ovate-triangular, c. 9.8 x 2 mm, index c. 4.9; acuminate, margins very finely

ciliate, base narrowly attached; rather thick, surface adaxially glabrous,

abaxially very finely papillose. Lateral sepals triangular, c. 10.2 x 3 mm,

index 3.4; upper margin glabrous but shortly papillose locally, base broadly

attached; otherwise as the median sepal. Petals porrect, ovate, c. 1.5 x 0.9

mm, index 1.6—1.7; subacute, margins erose, long ciliate, base broadly

attached: rather thin, surface glabrous but with a few hairs distally on both

sides. Lip recurved and ovate in the proximal third, distally abruptly

narrowed and drawn out into a slender, gradually tapering top part; c. 6.3 x

1.2 mm, index 5.2—5.3; tip subacute, margins ciliate near the base, elsewhere

with long, thin, wavy hairs; lip rather thick in the proximal third, rather

thin elsewhere; adaxially concave near the base, with two rounded ridges

separated by a median furrow c. half-way up the proximal third, adaxial

surface glabrous but very finely papillose at about 1/3 of its length; abaxially

with a wide, rounded ridge towards the base, slightly convex elsewhere,

surface glabrous but with a tuft of long, thin wavy hairs at about 1/3 of its

length. Column c. 2.2 mm long, stigma protruding at its base. Stelidia

subulate with a knob-shaped, rounded tip, c. 1.5 mm long; with an

inconspicuous, rounded wing along the lower margin, at the base. Anther:

Figure 13. Bulbophyllum cerebellum J.J. Verm. (123) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, above: adaxial

side, below: abaxial side; e. Column and lip, lateral view; f. Anther, left: abaxial side,

right: adaxial side; g. Pollinia, left: two pairs, right: single pair. — a. From a photograph

by A. Kocyan; b. From Kocyan s.n. (spirit sample); c—g. From Giles 998 (spirit sample).

Bulbophyllum Sect. Altisceptrum and Hirtula 45

4mm

ik:

Imm

2mm

3mm

eed OEE = j

Figure 13. Bulbophyllum cerebellum J.J. Verm. (123)

46 Gard. Bull. Singapore 54 (2002)

front margin papillose, abaxially with many stiff hairs distally.

Colours: Plant medium green. Rhachis greenish, suffused with blackish

purple. Sepals yellowish towards the base, cream-coloured towards the tip,

suffused with reddish purple. Petals yellowish near the base, blackish purple

elsewhere. Lip cream-coloured, marked with reddish purple, hairs yellow.

Habitat & ecology: Found as an epiphyte in open or closed primary

evergreen forest on granite bedrock. Alt. 700—1500m. Flowering in Dec.

Distribution: VIETNAM: southern part (1 specimen seen).

Notes: All data on habitat, ecology and distribution are from L.

Averyanov (pers. comm.). According to him, the inflorescence is up to 60 cm

long and pendulous. The only specimen available to me has an incomplete

inflorescence: the basal part is missing.

125. Bulbophyllum penicillium Par. & Rchb.f. in Rchb.f. — Fig. 15, plate 9.

Bulbophyllum penicillium Par. & Rchb.f. in Rchb.f., Trans. Linn. Soc. London, Bot. 30

(1874) 151; King & Pantling, Ann. Roy. Bot. Gard. (Calcutta) 8 (1898) 79, t.

108. — Phyllorchis penicillium (Par. & Rchb.f. in Rchb.f.) O. Kuntze, Rev. Gen.

Pl. 2 (1891) 677. — TYPE: Myanmar, Moulmein, Parish 303 (K, holo.; W iso.).

Bulbophyllum inopinatum W.W. Sm., Notes Roy. Bot. Gard. Edinburgh 8 (1915) 346.

— TYPE: Hort. Edinburgh s.n. (E, holo.).

[Not B. penicillium sensu Rolfe, J. Bot. 23 (1885) 215; Rolfe in Vidal y Soler, Phaner.

Cum. Philipp. (1885) 80 & 149; Vidal y Soler, Rev. Pl. Vasc. Filip. (1886) 267;

Ames, Philipp. J. Sci. 7 (1912) 128; Ames, Enum. Philipp. Fl. Pl. 1 (1925) 392. -

Specimen: Cuming 2076 (BM, W; = B. flavescens [B1.] Lindl., sect.

Aphanobulbon).|

[Not B. penicillium sensu Seidenfaden, Opera Bot. 124 (1995) 47 (= B. lanuginosum).|

Rhizome 2.5—4 mm diam., sections between pseudobulbs 1.2—3.5

cm long. Pseudobulbs close or distant, ovoid, 1.8—3.5 x 1.5—2 cm, laterally

flattened(?). Petiole 4—7.5 cm. Leaf blade elliptic to (ob-)ovate, 12.5—20.5

x 1.8—3.6 cm, index (length/width) 4.7—8.2; acute. /nflorescence an

elongated, lax raceme, 28—65 cm, 16—65-flowered. Peduncle + patent,

15—29 cm, bracts 4—6, the longest 10—15 mm long. Rhachis arching or

nodding, spindle-shaped, 12—S0 x c. 0.5 cm, glabrous. Floral bracts ovate

to triangular, 4—8 x 2—4 mm, acuminate. Flowers not fully opening, several

simultaneously. Pedicel and ovary 5.5—6.5 mm long, basal node. + flush

with the surface of the rhachis. Median sepal + porrect, ovate to triangular,

Figure 14. Bulbophyllum clipeibulbum J.J. Verm. (124) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial

side, right: abaxial side; e. Column and lip, lateral view; f. Anther, left: abaxial side,

right: adaxial side; g. Pollinia, left: single pair, right: two pairs. — a (plant). From Hort.

Vienna 099-B-160-2 (living plant); a (inflorescence)—g. From Hort. Salzburg OR 637/

94 (spirit sample).

Bulbophyllum Sect. Altisceptrum and Hirtula 47

CMM ae as |

Smm

| —maeane

DEER

e oe

~~. JEN

SSS a tig LD

Figure 14. Bulbophyllum clipeibulbum J.J. Verm. (124)

48 Gard. Bull. Singapore 54 (2002)

7.5—12.5 x 1.8—3 mm, index 3.4—5.5; acute, margins very finely ciliate,

base broadly attached; rather thin, surface glabrous. Lateral sepals recurved,

8—12 x 2.5—3.3 mm, index 2.9—4; upper margin glabrous, otherwise as

the median sepal. Petals porrect, elliptic to ovate, 1—1.5 x 0.8—1.2 mm,

index 1.2—1.4; rounded, margins long ciliate, base rather narrowly attached;

rather thick, surface glabrous. Lip somewhat recurved in the basal half,

ovate-subtriangular, 4—8 x 0.9—1.3 mm, index 4—6.7; gradually tapering

towards a subacute tip, margins with stiff hairs over its entire length, those

in the proximal third short, thin, those elsewhere short or long, slightly

swollen; lip rather thick near the base, rather thin elsewhere; adaxially

somewhat concave near the base, with three ridges, the lateral ridges low,

rounded, towards the base almost touching over a deep slit before diverging,

distally slightly diverging and continuing up to c. 1/2 of the lip, the median

ridge starting near the ligament as a thick patch and continuing up to 1/3

of the lip as low, narrow, obtuse ridge, adaxial surface glabrous but slightly

finely hirsute near the base; abaxially with a retuse ridge near the base,

surface otherwise flat, glabrous. Column 1.8—3 mm long, stigma slightly

protruding at its base. Stelidia subulate, 1—1.7 mm long, tip rounded; with

an inconspicuous, rounded wing along the lower margin, at the base. Anther:

front margin papillose and with some stiff hairs, abaxial ridge with scattered

stiff hairs.

Colours: Rhachis dark brownish purple. Flowers greenish, marked

with purple: sepals with purple veins, the lateral sometimes also with a

purple blotch c. half-way; petals entirely purple, or with a purple top part,

or with a purple vein and blotches along the margins; lip marked with

purple or entirely purple; hairs on lip purple or pink with purple segments

at irregular intervals.

Habitat & ecology: Alt. up to 2000 m. Flowering in Jan, Aug, Sep

(May, Jun in cultivation).

Distribution: BHUTAN (1 specimen seen). INDIA: Assam (1).

MYANMAR: Tenasserim (1).

Notes: In herbarium specimens the rhachis shrivels to about the same

diameter as the peduncle; its thickness in fig. 15 has been estimated from

the plate in King and Pantling (1898).

Figure 15. Bulbophyllum penicillium Par. & Rchb.f. in Rchb.f. (125) — a. Habit; b.

Flower; c. Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; d.

Lip, adaxial side; e. Lip, abaxial side; f. Column and lip, lateral view; g. Anther, above:

abaxial side, below: adaxial side; h. Pollinia, left: two pairs, right: single pair. — a

(plant). From Parish 303; a (inflorescence). From Hort. Glasnevin s.n.; b—h. From

O’Brien, d.d. 6-1895 (all herbarium specimens).

Bulbophyllum Sect. Altisceptrum and Hirtula

AL ALE

te m\y NWA Sf

3mm

1 fp p bi; Y tg _

2mm

ANN <NAASSS

\ NARS WS NS

AVS

ay SN R Se

ROTA ON LWRAQAS

\ \ \ AS =

Figure 15. Bulbophyllum penicillium Par. & Rchb.f. in Rchb.f. (125)

50 Gard. Bull. Singapore 54 (2002)

126. Bulbophyllum setuliferum J.J. Verm. & L.G. Saw — Fig. 16, plate 10.

Bulbophyllum setuliferum J.J. Verm. & L.G. Saw, Gard. Bull. Singapore 52 (2000) 289.

— TYPE: Malaysia, Pahang, Cameron Highlands, near Tanah Rata, Hort.

Singapore SBG-O 490 (SING, holo.; KEP, KLU, iso.).

Rhizome 2.5—5S mm diam., sections between pseudobulbs 1.4—4 cm

long. Pseudobulbs close or distant, depressed conical, 1.4—2.5 x 1.3—2.8

cm. Petiole 3—6.5 cm. Leaf blade elliptic to (ob-)ovate, 10—17 x 2—3.2

cm, index (length/width) 3.7—6.1; acute to slightly acuminate. /nflorescence

an elongated, lax raceme, 33—80 cm, 22—68-flowered. Peduncle + patent,

22—39 cm, bracts 5—6, the longest 6—9 mm long. Rhachis (slightly)

nodding, spindle-shaped, flattened or not, 11—41 x 0.4—0.9 cm, glabrous,

without blackish dots. Floral bracts triangular, 5—6.6 x 2—3.2 mm, acute.

Flowers not much opening, one at a time or a few simultaneously. Pedicel

and ovary 3—4 mm long, without blackish dots, basal node + flush with

the surface of the rhachis. Median sepal porrect or little incurved, triangular,

6—7.5 x 1.8—2 mm, index 3—3.8; acute, margins very finely ciliate, base

rather broadly attached; rather thin, surface adaxially finely hirsute,

abaxially glabrous. Lateral sepals recurved, 6—8 x 2.1—2.5 mm, index

2.6—3.3, upper margin glabrous; otherwise as the median sepal. Petals

porrect, transversely elliptic to reniform, 0.7—0.8 x 0.9—1.1 mm, index

0.6—0.7; rounded, margins long ciliate, base narrowly attached; rather thin,

surface glabrous. Lip slightly curved in the proximal half, subspathulate,

3.5—4 x c. 1.2 mm, index 2.9—3.3; truncate to rounded, margins ciliate

along the proximal 2/3 of the lip and with long, slightly club-shaped hairs

along the distal 2/3, the latter increasing in length and thickness towards

the tip of the lip; very thick; adaxially concave near the base and near the

tip, slightly convex in between, with a thin median slit up to 1/4—1/3 of the

lip, with a distinct, narrow median ridge over + the entire length of the lip

and overtopping its front as a high crest, adaxial surface glabrous but

finely hirsute near the base; abaxially with a retuse ridge towards the base,

with a distinctly convex top part with scattered club-shaped hairs arising

from tiny pits, surface otherwise glabrous. Column 1.5—2.1 mm long,

column foot slightly swollen. Ste/idia falcate, triangular, 0.5—1 mm long,

acute, with a deltoid, rounded wing along the lower margin, at the base.

Anther: front margin and abaxially towards the tip papillose with elongated

papillae.

Figure 16. Bulbophyllum setuliferum J.J. Verm. & L.G. Saw (126) — a. Habit; b. Flower;

c. Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip,

below: adaxial side, above: abaxial side; e. Column and lip, lateral view; f. Anther,

above: abaxial side, below: adaxial side; g. Pollinia, left: single pair, right: two pair. — a.

From Hort. Singapore SBG-O 490; b—g. From Hort. Singapore SBG-O 625 (both

spirit samples).

Bulbophyllum Sect. Altisceptrum and Hirtula a4

“4 wail,

2mm

~ =

srs pee

pn Len

Figure 16. Bulbophyllum setuliferum J.J. Verm. & L.G. Saw (126)

52 Gard. Bull. Singapore 54 (2002)

Colours: Inflorescence and sepals pale green to whitish, suffused

with dull reddish purple; the sepals proximally around the veins only. Petals

translucent with a purple margin. Lip cream-coloured, with a purple margin,

and often with some purple spotting on both surfaces. Hairs on lip shiny

reddish purple, sometimes partly green. Column yellowish green, foot and

wings below stelidia marked with purple.

Habitat & ecology: In (lower) montane forest, on tree trunks and

branches, in shade; also found in moss at the foot of a tree. Alt. 1500—

1800 m. Flowering in Feb—Jun, Nov (May, Sep, Oct in cultivation).

Distribution: MALAYSIA: Peninsula (7 specimens seen).

127. Bulbophyllum sororculum J.J. Verm., sp. nov. — Fig. 17, plate 11.

Bulbophyllum sororculum J.J. Verm., a Bulbophyllo aithorhachide in inflorescentiis

erectis, labello adaxialiter glabro differt. — TYPE: Indonesia, Sulawesi, central

part, M. Brans s.n. (WU, holo.).

Rhizome 2.2—4 mm diam., sections between pseudobulbs 1.4—3 cm

long. Pseudobulbs distant, (depressed) conical, 1—1.4 x 0.8—2.2 cm. Petiole

1.4—2.5 cm. Leaf blade elliptic, 6—13.5 x 1.4—2.6 cm, index (length/width)

3.3—5.8; acute. Inflorescence an elongated, lax raceme, erect to patent, c.

17 cm (not fully developed), c. 22-flowered. Peduncle c. 9.5 cm, bracts c. 4,

the longest c. 8 mm long. Rhachis spindle-shaped, c. 7.5 x 0.4 cm (not fully

developed), with many minute, blackish dots. Floral bracts triangular, c.

4.5 x 2.8 mm, acute. Flowers opening wide, a few simultaneously. Pedicel

and ovary c. 2.5 mm long, with minute blackish dots, basal node on a c. 0.8

mm-long stump. Median sepal refiexed, ovate, c. 5.2 x 2 mm, index c. 2.6;

acute, margins very finely ciliate, base rather narrowly attached; rather

thick, surface adaxially sparsely hirsute, abaxially with few minute blackish

dots near the base. Lateral sepals incurved to reflexed, triangular, c. 5 x 2.5

mm, index c. 2; acuminate, upper margin glabrous; otherwise as the median

sepal. Petals porrect, (ob-)ovate, 1.2—2 x 1-1.2 mm, index 1.2—1.7; rounded,

margins long ciliate towards the tip, base rather narrowly attached; rather

thin, surface glabrous. Lip recurved in the proximal half, subspathulate, c.

3.2 x 1.1 mm, index 2.4—2.5; rounded, margins with long hairs increasing

in length and thickness towards the tip of the lip; very thick; adaxially

concave near the base, convex elsewhere, with a distinct, narrow median

ridge embedded in a furrow over the distal 2/3 of the lip, protruding from

Figure 17. Bulbophyllum sororculum J.J. Verm. (127) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: abaxial

side, right: adaxial side; e. Column and lip, lateral view; f. Anther, left: abaxial side,

right: adaxial side; g. Pollinia, above: single pair, below: two pairs. — All from M. Brans

s.n. (spirit sample).

Bulbophyllum Sect. Altisceptrum and Hirtula

Imm

Figure 17. Bulbophyllum sororculum J.J. Verm. (127)

3mm

54 Gard. Bull. Singapore 54 (2002)

the furrow in the distal 1/3 and overtopping its front as a high crest, adaxial

surface glabrous but finely hirsute near the base; abaxially with a retuse

ridge towards the base, with a distinctly convex top part with scattered

club-shaped hairs, surface otherwise glabrous. Column c. 2 mm long, column

foot slightly swollen, abaxially without a spur near the tip. Stelidia slightly

falcate, subulate, c. 1.1 mm long, acute, with a deltoid, obtuse wing along

the lower margin, at the base. Anther: front margin and abaxial ridge

densely papillose with elongated papillae.

Colours: Rhachis dull purplish red with tiny green spots. Floral bracts

green, suffused with dull purple. Sepals greenish, heavily spotted with dull

purple. Petals greenish at base, white in the middle, grading into a blackish

purple top part; hairs yellow. Lip greenish at the base, yellowish elsewhere,

margins and median crests almost black, elsewhere with almost black spots.

Habitat & ecology: Epiphyte in primary forest, on the bole of a tree

c.3 m above the ground, at c. 500 m alt. Flowering in Oct, Nov in cultivation.

Distribution: INDONESIA: Sulawesi, central part (1 specimen seen).

Notes: Bulbophyllum sororculum is in some respects intermediate

between B. setuliferum and B. aithorhachis. The former differs in having a

lip with a concave top part, the latter in having a pendulous inflorescence

and a lip that is hirsute distally on the adaxial side; and both differ in

having less elongated petals.

One of the flowers available had the two petals of different lengths.

The species is remarkable for the asymmetrically opening flowers:

these are kept so close to the rhachis that the lateral sepal facing the

rhachis cannot spread properly.

128. Bulbophyllum aithorhachis J.J. Verm. — Fig. 18, plate 12, 13.

Bulbophyllum aithorhachis J.J. Verm., Blumea 41 (1996) 349. — TYPE: Brunei, Sungei

Ingei, surroundings of Batu Melintang, Jongejan cult. 2219 (L, holo.).

Rhizome 3—4 mm diam., sections between pseudobulbs 1—2.5 cm

long. Pseudobulbs close or distant, depressed conical, 0.6—1.4 x 1.1—2.2

cm. Petiole 0.8—3.8 cm. Leaf blade elliptic to ovate, 9—16 x 1.7—4 cm,

index (length/width) 4.0—5.3; acute. Inflorescence an elongated, lax raceme,

pendulous, 27—S0 cm, 22—50-flowered. Peduncle 16—28 cm, bracts 4—5,

the longest 7—12 mm long. Rhachis spindle-shaped, 10—22 x 0.8—1.2 cm,

with many minute blackish dots. Floral bracts triangular, 3.5—4 x 4—5

Figure 18. Bulbophyllum aithorhachis J.J. Verm. (128) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, above: adaxial

side, below: abaxial side; e. Column and lip, lateral view; f. Anther, above: abaxial

side, below: adaxial side; g. Pollinia, left: two pairs, right: single pair. — All from

Jongejan cult. 2219 (spirit sample).

Bulbophyllum Sect. Altisceptrum and Hirtula 55

3mm

2mm

Figure 18. Bulbophyllum aithorhachis J.J. Verm. (128)

56 Gard. Bull. Singapore 54 (2002)

mm, acute. Flowers opening wide, a few simultaneously. Pedicel and ovary

c.4 mm long, with many minute blackish dots, basal node + flush with the

surface of the rhachis. Median sepal recurved to reflexed, ovate, 5.5—6 x

2—2.5 mm, index 2.2—3; obtuse, margins very finely ciliate, base rather

broadly attached; rather thick, surface adaxially sparsely hirsute, abaxially

with few minute blackish dots near the base. Lateral sepals porrect to

spreading, triangular, 5.2—6.2 x 2.2—3 mm, index 1.7—2.9, upper margin

glabrous; otherwise as the median sepal. Petals porrect, transversely elliptic

to reniform, 0.8—1.2 x 1.3—1.6 mm, index 0.6—0.8; rounded, margins long

ciliate towards the tip, base narrowly attached; rather thick, surface glabrous.

Lip abruptly recurved in the proximal half, obovate, 3.3—4 x c. 1.3 mm,

index 2.4—2.5; rounded, margins with long hairs increasing in length and

thickness towards the tip of the lip; very thick; adaxially hardly concave

and with two proximally diverging calli near the base, with a median slit up

to 1/4 of the lip, with three distinct, obtuse ridges almost from base to tip,

the lateral low and wide, diverging proximally and distally, running parallel

and close together + half-way up the lip, the median ridge high and narrow,

starting in the proximal half and overtopping the front of the lip as a high

crest, adaxial surface irregularly hirsute except near the base; abaxially

with a wide, retuse ridge towards the base, convex towards the tip, surface

finely velutinous distally, glabrous proximally. Column c. 2 mm long, column

foot slightly swollen. Stelidia falcate, triangular, c. 0.7 mm long, acute, with

a deltoid, obtuse wing along the lower margin, at the base. Anther: front

margin and abaxial ridge densely papillose with elongated papillae.

Colours: Rhachis, pedicel and ovary pale greenish, suffused with

reddish purple. Flowers pale greenish or yellow, the parts marked with

dark reddish purple: the sepals spotted, particularly towards the margins

and near the veins, the petals along the top margin, the lip adaxially towards

the margins, abaxially towards the tip. Cilia along the petals and thicker

hairs along the top part of the lip dark reddish purple; but hairs on the

midrib and fine hairs along the margins of the lip white.

Habitat & ecology: Primary forest. Alt. 50—800 m. Found with mature

fruits in Apr.

Distribution: BRUNEI (1 specimen seen). INDONESIA: Sumatra

Gb):

Notes: As in the previous species, the flowers cannot open completely

because they are held so close to the rhachis that the lateral sepal facing

the rhachis is trapped in between.

The Sumatra specimen only has fruits and could either be this or B.

setuliferum. On account of the very thin peduncle (indicating a pendulous

inflorescence), the rhachis structure, and the length of the floral bracts, it

has been included here.

Bulbophyllum Sect. Altisceptrum and Hirtula ba

129. Bulbophyllum echinochilum F. Kraenzl. — Fig. 19. (probably extinct)

Bulbophyllum echinochilum F. Kraenzl., Repert. Spec. Nov. Regni Veg. 17 (1921) 385;

Ames, Sched. Orch. 6 (1923) 81. — TYPE: Philippines, Manila, Hort. Muenchen

(Loher s.n.) d.d. 1916 (B[?], holo., destroyed; AMES, iso.).

Pseudobulbs close or distant, depressed conical, 1.2—2.5 x 1.6—2.0

cm. Petiole c. 1.1 cm. Leaf blade elliptic, 7.5—10 x 1.4—2 cm, index (length/

width) 5—5.3. Inflorescence an elongated, lax raceme c. 17 cm, 18—20-

flowered. Peduncle + patent, c. 9 cm. Rhachis arching or nodding, spindle-

shaped, c. 8 x 0.5 cm. Pedicel and ovary c. 3—4 mm long, with scattered

minute blackish dots. Median sepal recurved to reflexed, ovate, c. 5.8 x 1.7

mm, index c. 3.4; acute, margins very finely ciliate, base rather broadly

attached; rather thin, surface adaxially finely hirsute, abaxially with few

minute, blackish dots near the base. Lateral sepals recurved, subtriangular,

c. 6 x 2 mm, index c. 3, upper margin finely papillose; otherwise as the

median sepal. Petals porrect, elliptic, c. 1.2 x 1 mm, index c. 1.2; rounded,

margins long ciliate, base narrowly attached; thin, surface glabrous except

for a small tuft of hairs on the abaxial side. Lip almost straight, + elliptic, c.

3.7 x 0.9 mm, index c. 4.1; rounded, margins ciliate except near the base

and near the tip and with long, + straight, slightly club-shaped hairs along

the distal 2/3, the latter increasing in length towards the tip of the lip; very

thick; adaxially concave near the base, with three distinct, obtuse ridges,

the two lateral ridges wide, diverging proximally, the median higher and

narrow, all three ridges fusing into a single ridge well before the front of

the lip, adaxial surface glabrous but finely hirsute near the base; abaxially

with a weak, retuse ridge towards the base, with a distinctly convex top

part with scattered club-shaped hairs arising from tiny pits, surface otherwise

glabrous. Column c. 1.8 mm long, column foot slightly swollen. Stelidia

slightly falcate, triangular, c. 0.7 mm long, acute.

Colours: Pseudobulbs reddish. Rhachis dull purple with white spots.

Sepals yellowish green, with tiny reddish brown dots. Petals white with

reddish brown margins. Lip yellowish green, hairs yellowish brown.

Habitat & ecology: Flowering in Dec in cultivation.

Distribution: PHILIPPINES: Luzon (1 specimen seen). Not collected

since-c. 1910.

Notes: Only known from an incomplete isotype specimen. The

description of the vegetative parts has been copied from Kraenzlin’s

description and his notes with the isotype material.

Ames (1923) doubted its provenance on account of a superficial

resemblance to African species. As it fits well in sect. Hirtula, however, it

is clearly of Asian origin.

58 Gard. Bull. Singapore 54 (2002)

130. Bulbophyllum lanuginosum J.J. Verm., sp. nov. — Fig. 20.

Bulbophyllum lanuginosum J.J. Verm., a Bulbophyllo penicillio \abello adaxialiter

crista distincta, pilis lanatis secus margines differt. — TYPE: Thailand, purchased

on Bangkok market, Thaitong 1395 (C, holo.).

Bulbophyllum penicillium sensu Seidenfaden, Opera Bot. 124 (1995) 47, colour

photograph pl. 9b, non Par. & Rchb.f. in Rchb.f.

Rhizome c. 2 mm diam., sections between pseudobulbs c. 0.6 cm

long. Pseudobulbs ?close, ovoid, c. 1.2 x 1.2 cm. Petiole 1.5 cm. Leaf blade

elliptic, c. 9 x 1.5 cm, index (length/width) c. 6; acute. Inflorescence an

elongated, lax raceme, 9.5—14 cm, 16—20-flowered. Peduncle + erect, 3—

4 cm, bracts c. 4, the longest 4—4.5 mm long. Rhachis erect, swollen, +

circular in section, 6.5—10 x c. 0.15 cm, glabrous. Floral bracts triangular,

2.5—3 x 1.5—1.8 mm, acute. Flowers opening wide, one at a time or a few

simultaneously. Pedicel and ovary 2—2.2 mm long, without minute blackish

dots, basal node on a 0.7—1 mm-long stump. Median sepal spreading to

reflexed, triangular, c. 6.5 x 1.9 mm, index c. 3.4; acute, margins very finely

ciliate, base rather broadly attached; rather thin, surface adaxially very

finely hirsute locally, abaxially glabrous. Lateral sepals c. 6 x 2 mm, index

c. 3, upper margin + glabrous; otherwise as the median sepal. Petals porrect,

transversely elliptic, c. 0.9 x 1.2 mm, index 0.7—0.8; subacute, margins long

ciliate, base narrowly attached; thin, surface glabrous. Lip somewhat

recurved in the basal half, + oblong, c. 3 x 0.9 mm, index 3.3—3.4; rounded,

margins ciliate near the base and with long, thin, wavy hairs near the tip,

with shorter, thin, + straight hairs in between; very thick; adaxially slightly

concave and with two calli near the base, with a thin median slit up to c. 1/3

of the lip, with three distinct, obtuse ridges fused into a crest that increases

in height into a high and narrow wedge distally, the two lateral ridges

wide, diverging proximally, forming the edges of the wedge distally, the

median ridge narrow, ending just before the end of the wedge, adaxial

surface glabrous but finely hirsute near the base; abaxially with a retuse

ridge towards the base, with a distinctly convex top part with long, thin,

wavy hairs, surface otherwise glabrous. Column c. 1.6 mm long, stigma

protruding at its base. Stelidia subulate, c. 0.9 mm long, acute, with a large,

antrorse, triangular, acute tooth along the upper margin. Anther: front

margin papillose, abaxially towards the tip with elongated papillae.

Colours: Rhachis, bracts, pedicel and ovary greenish. Flowers

greenish; sepals with brownish purple veins, petals with brownish purple

Figure 19. Bulbophyllum echinochilum Kraenzl. (129) — a. Pseudobulb and leaf; b.

Flower; c. Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; d.

Lip, above: adaxial side, below: abaxial side; e. Column and lip, lateral view. — All

from Hort. Muenchen (Loher s.n.) d.d. 1916 (poor quality herbarium specimen).

Bulbophyllum Sect. Altisceptrum and Hirtula 59

= s 8 a

SEPSAS ne

a’ e . Sad !

ss :

3mm

—_ = =

To Z

—-s =

i/ SS

B E

44);

2mm

SSSA

Ty Py 4 , : Fe

‘as i aS poco : : re S | Scm

SOV

;

< SR,

Figure 19. Bulbophyllum echinochilum Kraenzl. (129)

60 Gard. Bull. Singapore 54 (2002)

hairs, and lip with the lateral crests and the abaxial side similarly coloured.

Shorter, stiff hairs on lip purple, the long, wavy hairs white.

Habitat & ecology: Flowering observed in Jun.

Distribution: THAILAND: ‘probably Kanburi area’ (1 specimen

seen).

Notes: Only known from the type specimen. The description of the

vegetative parts is not complete.

Erroneously identified by Seidenfaden (1995) as B. penicillium; it

differs from that species in its smaller stature, shorter peduncle, its crested,

obtuse lip with woolly, wavy hairs (hence the specific epithet).

131. Bulbophyllum atratum J.J. Sm. — Fig. 21, plate 14.

Bulbophyllum atratum J.J. Sm., Bull. Jard. Bot. Buitenzorg, s. 2, 25 (1917) 73. - TYPE:

Indonesia, Sumatra, Barisan Range, Hort. Bogor (leg. Ajoeb, Exp. Jacobson)

862 (BO, holo., not seen, likely isotypes in L).

Rhizome 1—2 mm diam., sections between pseudobulbs 0.4—0.8 cm

long. Pseudobulbs touching, ovoid, 0.5—1.0 x 0.6—1 cm. Petiole 2—3 cm.

Leaf blade elliptic to ovate, 10—14.5 x 2.5—4.3 cm, index (length/width)

3.3—4.6; obtuse. Inflorescence an elongated, rather lax to rather dense

raceme, entirely pendulous, 35—50 cm, 8-40-flowered. Peduncle 25—33

cm, bracts 4—5, the longest 6—10 mm long. Rhachis swollen, + circular in

section, 9.5—16.5 x 0.3—0.5 cm, glabrous, top 2—7 cm with minute, sterile

bracts only. Floral bracts triangular, 0.8—1 x 1—1.2 mm, acuminate. Flowers

not fully opening, many simultaneously. Pedicel and ovary 2.3—3 mm long,

papillose-hirsute, basal node on a 0.2—0.3 mm-long stump. Median sepal +

porrect and incurved distally, triangular, 5—5.5 x 1.5—2 mm, index 2.5—

3.7; acute, margins ciliolate, base broadly attached; rather thin, surface

adaxially glabrous, abaxially papillose-hirsute. Lateral sepals (slightly)

recurved, falcate, 4—4.2 x 2.5—2.9 mm, index 1.3—1.7; otherwise as the

median sepal. Petals porrect, falcate, ovate with or without two minute

teeth proximally along the lower margin, 3.5—4 x 1.2—1.5 mm, index

2.5—2.9; acute, margins ciliate with cilia longer towards the tip, base

narrowly attached; thin, surface glabrous except for a few hairs near the

tip. Lip abruptly recurved c. half-way, elliptic to obovate, 1.8—2.2 x 1—1.3

mm, index 1.5—2; rounded, margins ciliate except near the base; very

thick; adaxially hardly concave and with 2 inconspicuous calli and a short

Figure 20. Bulbophyllum lanuginosum J.J. Verm. (130) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial

side, right: abaxial side; e. Column and lip, lateral view; f. Anther, left: adaxial side,

right: abaxial side; g. Pollinia, left: two pairs, right: single pair. — All from Thaitong

1395 (spirit sample).

Bulbophyllum Sect. Altisceptrum and Hirtula 61

LUBU

—

SSSA AN . R

\ NOSSRR .

LEZ

YY 2

OSA

PES \ Ne

(SEZ NE

ZEIT asp WAVE

LIES %é AY SAN Z5

Z \

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f “py

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i Pu ‘ “ ,

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7 f ua

as we Ba - Imm

N X SSS

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a ee

Imm ;

ASS SF AK)4.,

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AW “=

NSN

yf |}

4 V

Figure 20. Bulbophyllum lanuginosum J.J. Verm. (130)

62 Gard. Bull. Singapore 54 (2002)

median slit near the base, with two weak, obtuse ridges that are proximally

diverging and running parallel and close to the median line up to near the

tip, adaxial surface glabrous; abaxially with a wide, retuse ridge and glabrous

towards the base, convex and papillose-hirsute towards the tip. Column c.

1.5 mm long, stigma slightly protruding at its base, column foot abaxially

without a spur near the tip. Stelidia c. 0.6 mm long, acute, + rectangular in

outline with a distinct, antrorse, deltoid, obtuse to acute tooth along the

upper margin and often along the lower margin. Anther: front margin

papillose with elongated papillae, abaxial ridge + glabrous.

Colours: Peduncle dark green at the base, blotched with maroon

towards the tip, rhachis brownish or dark maroon, almost black. Flowers

yellowish or cream coloured near the base, whitish towards the tip, sepals

with partially or entirely blackish purple veins and margins, the lateral

sometimes with large blackish purple spots instead of coloured veins; petals

without blackish markings. Lip blackish towards the tip, white towards the

base, with fine red spots adaxially. Column and anther creamy white.

Habitat & ecology: Epiphyte in montane forest. Alt. c. 1000 m.

Flowering in Mar, Nov.

Distribution: MALAYSIA: Peninsula (2 specimens seen).

INDONESIA: Sumatra (2).

Notes: Specimens from Peninsular Malaysia with wider leaves than

in the description above have been seen in cultivation.

132. Bulbophyllum groeneveldtii J.J. Sm. — Fig. 22, plate 15.

Bulbophyllum groeneveldtii J.J. Sm., Bull. Jard. Bot. Buitenzorg, s. 3, 2 (1920) 95. —

TYPE: Indonesia, Sumatra, Padang Highlands, Jacobson cult. 951 (leg.

Groenenveldt) (BO, holo).

Rhizome c. 3 mm diam., sections between pseudobulbs c. 0.4 cm

long. Pseudobulbs touching, (depressed) globose, 0.4—0.7 x 0.4—1.2 cm.

Petiole 0.3—0.5 cm. Leaf. blade elliptic to obovate, 8—28 x 0.8—3.3 cm,

index (length/width) 8.3—12.5; obtuse. Inflorescence an elongated, lax

raceme, entirely pendulous, 7—18 cm, up to 45-flowered. Peduncle 4—7

cm, bracts c. 4, the longest c. 2 mm long. Rhachis swollen, + circular in

section, 3.5—12 x 0.2—0.3 cm, papillose-hirsute, to the very tip with flowers.

Floral bracts triangular, c. 1.5 x 1 mm, acute. Flowers not much opening, a

few simultaneously. Pedicel and ovary 1.5—2.6 mm long, papillose-hirsute,

Figure 21. Bulbophyllum atratum J.J. Sm. (131) — a. Habit; b. Flower; c. Flower analysis,

from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial side, right:

abaxial side; e. Column and lip, lateral view; f. Anther, left: adaxial side, right: abaxial

side; g. Pollinia, above: two pairs, below: single pair. — All from Hort. Singapore SBG-

O 87 (spirit sample).

Bulbophyllum Sect. Altisceptrum and Hirtula

Imm

Se er eT

Som

;

—_

weet cower hoe OPE e be cegere™

Figure 21. Bulbophyllum atratum J.J. Sm. (131)

64 Gard. Bull. Singapore 54 (2002)

basal node + flush with the surface of the rhachis. Median sepal + porrect,

triangular, 3.2—3.3 x 1.3—1.5 mm, index 2.1—2.5; acute, margins papillose-

hirsute, base rather broadly attached; rather thick, surface adaxially

glabrous, abaxially papillose-hirsute. Lateral sepals slightly recurved, 3.3—

3.5 x 2.2—3 mm, index 1.1—1.5; otherwise as the median sepal. Petals

porrect, falcate or not, hastate with an ovate, acute top part with ciliate

margins, and a triangular, acute tooth along upper and lower margin, near

the base; 1.6—2.2 x 1.2—1.5 mm, index 1—1.8, base rather broadly attached:

rather thin, surface glabrous. Lip abruptly recurved in the basal half, elliptic,

2—2.4 x 0.9—1.5 mm, index 1.6—2.2; rounded, margins ciliate in the

proximal third; very thick; adaxially not concave and with a short median

slit near the base, with two weak, obtuse ridges proximally diverging, and

running parallel and close to the median line up to near the tip, adaxial

surface glabrous but hirsute near the base and in the top half; abaxially

with a wide, rounded ridge towards the base, convex and with scattered

coarse hairs towards the tip, surface otherwise glabrous. Column 1—1.3

mm long, stigma with an inconspicuous, rounded callus at its base, column

foot with two inconspicuous wings distally, abaxially without a spur near

the tip. Srelidia = triangular, c. 0.6 mm long, acute, with 2 small, antrorse,

deltoid, acute teeth along the upper margin and 1 = similar tooth along

their lower margin. Anther: front margin papillose with elongated papillae,

abaxial ridge + glabrous.

Colours: Flowers yellow or pale green, sepals and petals with partially

or entirely brown or blackish purple veins and margins, sometimes with

additional spots; petals with brown or blackish purple cilia. Lip entirely

blackish, or blackish towards the tip only, except for the midvein.

Habitat & ecology: Understorey epiphyte in high montane forest and

in low and open kerangas forest. Rooting in small humus deposits and

moss. Alt. 1100—1300 m. Flowering in Jan, Oct—Dec.

Distribution: MALAYSIA: Sabah (4 specimens seen). INDONESIA:

Sumatra (1).

Notes: When sterile, it is characterised by the pendulous, almost

strap-shaped leaves.

133. Bulbophyllum grotianum J.J. Verm., sp. nov. — Fig. 23, plate 16, 17.

Bulbophyllum grotianum J.J. Verm., a Bulbophyllo atrato et B. groeneveldtii

Figure 22. Bulbophyllum groeneveldtii J.J. Sm. (132) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal; lip; d. Lip, above: adaxial

side, below: abaxial side; e. Column and lip, lateral view; f. Anther, above: adaxial

side, below: abaxial side; g. Pollinia, left: two pairs, right: single pair. — All from

Vermeulen 1107 (living plant).

Bulbophyllum Sect. Altisceptrum and Hirtula

MAMI ASSES NER

DEAN PTT

~~.

~—----

Figure 22. Bulbophyllum groeneveldtii J.J. Sm. (132)

66 Gard. Bull. Singapore 54 (2002)

inflorescentia erecta, rhachide nutanti 0.6—0.7 cm crassa differt. — TYPE:

Malaysia, Sarawak, G. Mulu area, G. Pala near Batu Bulang, Hort. Leiden

970530 (L, holo.).

Rhizome 4—5 mm diam., sections between pseudobulbs 0.4—1 cm

long. Pseudobulbs touching, ovoid, 0.4—0.8 x 0.4—0.8 cm. Petiole 1.2—2.2

cm. Leaf blade elliptic, 8—16.5 x 2.8—3 cm, index (length/width) 2.3—5S.7;

rounded. /nflorescence an elongated, rather dense raceme, 17.5—23.5 cm,

45—65-flowered. Peduncle + erect, 13—16 cm; bracts c. 4, the longest c. 5.5

mm long. Rhachis nodding, spindle-shaped, 4.5—7.5 x 0.6—0.7 cm, papillose-

hirsute, to the very tip with flowers. Floral bracts triangular, 0.9—1.3 x c. 1

mm, cuspidate. Flowers not much opening, several simultaneously. Pedicel

and ovary 1.8—2 mm long, hirsute, basal node + flush with the surface of

the rhachis. Median sepal + porrect, triangular, 3—3.4 x 1.2—1.9 mm, index

1.7—2.5; acute, margins ciliate, base rather broadly attached; rather thick,

surface adaxially glabrous, abaxially hirsute. Lateral sepals slightly recurved,

3—3.2 x 2.2—2.5 mm, index 1.2—1.4, acuminate-cuspidate, otherwise as

the median sepal. Petals porrect, ovate-subhastate with an inconspicuous,

deltoid, acute tooth along upper and lower margin, 2.2—2.8 x 0.7—1 mm,

index 2.8—3.2; acute, margins ciliate except near the base, base rather

broadly attached; rather thin, surface glabrous. Lip abruptly recurved c.

half-way, obovate, 1.6—1.8 x 1.1—1.3 mm, index 1.3—1.5; broadly rounded,

margins ciliate; very thick; adaxially somewhat concave and with a short

median slit near the base, with two weak, rounded ridges starting in the

proximal half and slightly diverging close to the tip, with a small cavity in

between, adaxial surface glabrous but very finely papillose towards the

base; abaxially with a wide, retuse ridge towards the base, convex towards

the tip, surface glabrous. Column 1—1.2 mm long, column foot abaxially

without a spur near the tip. Ste/idia + triangular, 0.4A—0.5 mm long, acute,

with an antrorse, minute, triangular, acute tooth along the upper margin,

and 2 larger, narrowly triangular, acute teeth along the lower margin, the

proximal falcate. Anther: front margin ciliate, abaxial ridge somewhat

papillose.

Colours: Flowers cream-coloured or yellowish, marked with blackish

purple: the median sepal along the veins and margins; the lateral sepals

also with a blotch near the tip; the lip near the base with dots, the tip

entirely blackish purple abaxially. Column and anther cream-coloured,

Figure 23. Bulbophyllum grotianum J.J. Verm. (133) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial

side, right: abaxial side; e. Column and lip, lateral view; f. Anther, above: adaxial side,

below: abaxial side; g. Pollinia, above: single pair, below: two pairs. — All from Hort.

Leiden 970530 (spirit sample).

Bulbophyllum Sect. Altisceptrum and Hirtula

iff

be a

2mm

Page

rong

“tag KS Pe os a ay

[BAM

‘%

«a

COUN NN

“yn r]

Imm

2mm

Figure 23. Bulbophyllum grotianum J.J. Verm. (133)

68 Gard. Bull. Singapore 54 (2002)

spotted maroon. Pollinia yellow.

Habitat & ecology: Epiphyte in forest on a steep limestone hill. Alt.:

sea level to 250 m. Flowering in Jul—Sep in cultivation.

Distribution: MALAYSIA: Sarawak (1 specimen seen).

Notes: Differs from Bulbophyllum atratum and B. groeneveldtii in

having an erect inflorescence with a nodding, much thicker rhachis.

Named in honour of the late Mr. P.S.J. Groot, Borculo, the Netherlands,

an avid orchid grower and friend of thirty years.

134. Bulbophyllum janus J.J. Verm., nom. nov. — Fig. 24. (probably extinct)

Bulbophyllum janus J.J. Verm. — Basionym: Bulbophyllum platyrrhachis Ridl., J. Fed.

Malay States Mus. 8 (1917) 96. - SYNTYPES: Indonesia, Sumatra, Kerinchi,

Siolak Daras, Robinson & Boden Kloss s.n. (BM, lecto.); Sumatra, central part,

Sarik, Alahan Panjang, Micholitz s.n. (K).

[Not Bulbophyllum platyrhachis (Rolfe) Schltr, Bot. Jahrb. Syst. 38 (1905) 15 (= B.

maximum [Lindl.] Rchb.f.).]

[Not Bulbophyllum platirhachis De Wild., Miss. Laur. (1906) 223 (= B. acutebracteatum

De Wild. var. acutebracteatum).|

Rhizome c. 5 mm diam., sections between pseudobulbs 1—1.2 cm

long. Pseudobulbs ?close, ovoid, 2.2—3 x 1.4—1.8 cm. Petiole 6—7.5 cm.

Leaf blade elliptic to ovate, 16—19 x 4.8—5.5 cm, index (length/width)

3.3—3.5, rounded. /nflorescence an elongated, lax raceme, c. 65 cm, c. 100-

flowered. Peduncle erect, c. 52 cm, bracts c. 5, the longest 23—25 mm long.

Rhachis nodding, swollen and flattened, elliptic-ovate, c. 12.5 x 1.1 cm,

densely papillose, top c. 3 cm with minute, sterile bracts only. Floral bracts

triangular, 2—2.5 x c. 2 mm, acute. Flowers not much opening. Pedicel and

ovary c. 4.5 mm long, with scattered coarse papillae, basal node on a 0.2—

0.3 mm-long stump. Median sepal porrect, subtriangular, c. 4.3 x 2 mm,

index 2.1—2.2; acute, margins shortly ciliolate towards the tip, base broadly

attached; rather thick, surface adaxially glabrous but very finely pubescent

near the tip, abaxially with scattered, coarse papillae. Lateral sepals slightly

recurved, ovate, c. 4.3 x 3.8 mm, index 1|.1—1.2; subacute, otherwise as the

median sepal. Petals porrect, falcate, spathulate, c. 3.4 x 1.2 mm, index

2.8—2.9; truncate, margins ciliate, base broadly attached; rather thin; surface

glabrous but adaxially hirsute near the tip. Lip rather abruptly recurved c.

half-way, oblong, c. 3 x 1.9 mm, index 1.5—1.6; rounded, margins ciliate

with the longest hairs about half-way up the lip; very thick; adaxially concave

Figure 24. Bulbophyllum janus J.J. Verm. (134) — a. Habit; b. Flower; c. Flower analysis,

from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: abaxial side, right:

adaxial side; e. Column and lip, lateral view. — a (plant)—e. From Robinson & Boden

Kloss s.n.; a (inflorescence). From Micholitz s.n. (all herbarium specimens).

Bulbophyllum Sect. Altisceptrum and Hirtula 69

3mm

Figure 24. Bulbophyllum janus J.J. Verm. (134)

70 Gard. Bull. Singapore 54 (2002)

and with a basal callus continuing up to c. half-way up the lip and with a

short median slit near the base, with 2 distinct, rounded ridges diverging

proximally and continuing over c. 2/3 of the length of the lip, adaxial

surface finely hirsute; abaxially with a wide, rounded ridge towards the

base, convex towards the tip, surface glabrous towards the base and finely

hirsute towards the tip. Column c. 2.3 mm long, stigma with a retuse callus

at its base, column foot abaxially without a spur near the tip. Stelidia +

triangular, c. 0.6 mm long, obtuse, with a small, antrorse, obtuse tooth

along the upper margin, and a larger, triangular tooth along the lower

margin, near the base.

Colours: Rhachis greenish. Flowers yellow.

Habitat & ecology: Alt. 600—1000 m. Flowering observed in Mar.

Distribution: INDONESIA: Sumatra (2 specimen seen). Not collected

since 1914.

Notes: The name platyrrhachis is sufficiently close to platyrhachis

and platirhachis to warrant a new name, following ICBN 2000, art. 53.3.

The material available does not allow detailed description of the

development of the inflorescence during anthesis. According to the original

description the rhachis is 5 cm wide. The only rhachis available (Micholitz

s.n., Syntype) is, dry, 1.1 cm wide. Allowing for shrivelling, the reconstructed

width would be c. 1.5 cm, as drawn in fig. 24. The width given in the

original description 1s probably a typographical error.

The name refers to the two-sided rhachis.

135. Bulbophyllum xiphion J.J. Verm. — Fig. 25, plate 18, 19.

Bulbophyllum xiphion J.J. Verm., Blumea 41 (1996) 354. — TYPE: Indonesia,

Kalimantan Timur, Apo Kayan, Gunung Sungei Pendan area, Hort. Leiden

913241 (L, holo.).

Rhizome 5—6 mm diam., sections between pseudobulbs 0.9—1.5 cm

long. Pseudobulbs (almost) touching, ovoid, 1—1.6 x 1—1.6 cm. Petiole

4—6 cm. Leaf blade elliptic to obovate, 20—24 x 5—6.5 cm, index (length/

width) 3.5—4; rounded to obtuse. /nflorescence an elongated, lax raceme,

c. 42 cm, 60—70-flowered. Peduncle patent, c. 28 cm, bracts c. 4, the longest

c. 16 mm long. Rhachis nodding, swollen and flattened, elliptic-ovate, 11—

15 x 1.2—1.5 x 0.6—0.8 cm, finely rugose-papillose, top 3.5 cm with minute,

sterile bracts only. Floral bracts deltoid to triangular, 0.5—1.2 x 0.9—1.2

Figure 25. Bulbophyllum xiphion J.J. Verm. (135) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, above: adaxial

side, below: abaxial side; e. Column and lip, lateral view; f. Anther, left: adaxial side,

right: abaxial side; g. Pollinia, above: two pairs, below: single pair. — All from Hort.

Leiden 91324] (spirit sample).

Bulbophyllum Sect. Altisceptrum and Hirtula 71

2mm

aan

1mm

2mm

Figure 25. Bulbophyllum xiphion J.J. Verm. (135)

72 Gard. Bull. Singapore 54 (2002)

mm, acute. Flowers not much opening, a few simultaneously, anthesis from

the top of the rhachis downwards. Pedicel and ovary 2—2.4 mm long, basal

node = flush with the surface of the rhachis. Median sepal porrect, triangular,

5—5.5 x 2—2.4 mm, index 2.3—2.5; acute, margins shortly ciliolate, base

broadly attached; thick, surface adaxially very finely papillose towards the

margins, abaxially glabrous. Lateral sepals slightly recurved, slightly falcate,

5.5—6.2 x 3.2—3.5 mm, index 1.7—1.8; acuminate, adaxially glabrous,

otherwise as the median sepal. Petals porrect, obovate to elliptic, 3—3.2 x

c. 1.5 mm, index 2—2.2; rounded, margins long ciliate except near the

base, base broadly attached; thin, adaxially very finely papillose and with

some long hairs towards the tip, abaxially glabrous. Lip rather abruptly

recurved c. half-way, ovate, c. 3 x 1.7 mm, index 1.4—1.5; rounded, margins

long ciliate with the longest hairs about half-way up the lip; very thick;

adaxially somewhat concave near the base, with 2 inconspicuous, rounded

ridges diverging proximally and continuing over c. 2/3 of the length of the

lip, adaxial surface glabrous but very finely papillose towards the tip:

abaxially with a wide, truncate ridge towards the base, convex towards the

tip, surface glabrous but with scattered hairs implanted in tiny pits towards

the tip. Column 2—2.2 mm long, column foot abaxially without a spur near

the tip. Stelidia c. 0.6 mm long, acute, + rectangular in outline with an

antrorse, truncate wing along the upper margin, and a small, narrowly

triangular, acute tooth along the lower margin. Anther: front margin

papillose with elongated papillae, abaxial ridge papillose.

Colours: Rhachis yellowish green, either spotted with bronze purple

or not. Sepals off-white, adaxially spotted with purple. Lip purple.

Habitat & ecology: Epiphyte in primary forest on a ridge and on a

tree overhanging a river. Alt. 650—1050 m. Flowering observed in Oct.

Distribution: INDONESIA: Kalimantan Timur (2 specimen seen).

136. Bulbophyllum mirabile Hallier — Fig. 26, 27, plate 20, 21.

Bulbophyllum mirabile Hallier.. Ann. Jard. Bot. Buitenzorg 13 (1896) 316. — TYPE:

West Kalimantan, Hort. Bogor (Leg. Jaheri) (BO, holo., not seen).

Pseudobulbs touching, ovoid-conical, 2.5—3 x 1.5—2 cm. Petiole 3—

5 cm. Leaf blade elliptic, 19—20 x 7.4—8.5 cm, index (length/width) 2.3—

2.6; rounded. Jnflorescence an elongated, lax raceme, entirely pendulous,

11.3—29 cm, 75—120-flowered. Peduncle 2—9 cm, bracts 2—3, the longest

Figure 26. Bulbophyllum mirabile Hallier. (136) — a. Habit; b. Flowers, above: from

the basal part of the rhachis, below: from the top part of the rhachis; c. Flower analysis.

flower from the basal part of the rhachis, from left to right: median sepal, petal, lateral

sepal, lip; d. Flower analysis, flower from the top part of the rhachis, from left to right:

median sepal, petal, lateral sepal, lip — All from Jongejan cult. 4031 (spirit sample).

Bulbophyllum Sect. Altisceptrum and Hirtula 73

Figure 26. Bulbophyllum mirabile Hallier. (136)

74 Gard. Bull. Singapore 54 (2002)

11—13 mm long. Rhachis slightly papillose, basal part spindle-shaped, 4—

10.5 x 0.7—0O.8 cm, gradually tapering into a cylindrical top part of 4.3—14

x 0.3—0.45 cm. Floral bracts ovate to triangular, 2—2.5 x 12—2 mm,

(sub)acute. Flowers many open simultaneously, dimorphic, flowers along

the basal part of the rhachis different from those along the top part, anthesis

Starting at the transition between the two types, continuing upwards and

downwards. Flowers along the basal part of the rhachis not much opening.

Pedicel and ovary 2—3 mm long, locally papillose-hirsute, basal node +

flush with the surface of the rhachis. Median sepal + porrect, ovate, 3—5 x

1.1—1.8 mm, index 2.3—3.4; obtuse, margins ciliolate, base rather broadly

attached: rather thick, surface adaxially glabrous, abaxially papillose-hirsute.

Lateral sepals slightly recurved, slightly falcate, 2.7—5 x 2—3 mm, index

1.2—1.7; otherwise as the median sepal. Petals porrect, falcate, elliptic to

ovate, 1.3-3.5 x 0.3—0.9 mm, index 3.2—5; obtuse, margins ciliate, base

broadly attached: rather thick with a very thick top, surface glabrous,

adaxially sometimes hirsute towards the tip. Lip (rather abruptly) recurved

c. half-way, ovate, 1.8—2.3 x 0.9—1.3 mm, index 1.7—2: rounded, margins

ciliate with the cilia longest near the tip: very thick; adaxially concave and

with 1—2 basal calli sometimes continuing up to c. half-way up the lip and

with a short median slit near the base, with two distinct ridges diverging

proximally, continuing over 1/2—4/5 of the lip and distally fused to a wide,

rounded or flat median ridge overtopping the tip of the lip, adaxial surface

glabrous, distally sometimes verrucose along the median line; abaxially

with a wide, rounded to truncate, glabrous to papillose ridge towards the

base, convex and hairy towards the tip; basal auricles sometimes

inconspicuous. Column 1.1—1.8 mm long, stigma with two small, obtuse

lateral teeth near its base, column foot abaxially without a spur near the

tip. Stelidia 04—0.5 mm long, acute, + rectangular in outline with an

antrorse, deltoid, obtuse to acute tooth along their upper margin and a

similar but often patent tooth along their lower margin. Anther: front margin

ciliate, abaxial ridge + glabrous. Flowers along the top part of the rhachis

similar, but opening wide. Pedicel and ovary 0.5—1.5 mm long. Median

sepal somewhat recurved, 3.3—4.2 x 1.2—1.3 mm, index 2.5—3.5. Lateral

sepals spreading, 3.4—4.8 x 1.8—2.2 mm, index 1.8—2.2: otherwise as the

median sepal. Petals 1.3-2.2 x c. 0.5 mm, index 2.6—4.4. Lip recurved near

the base, elliptic, 34.2 x 1.5—2 mm, index 2.1—2.5; margins ciliolate near

Figure 27. Bulbophyllum mirabile Hallier. (136) — Flower from the basal part of the

rhachis: a. Lip, left: adaxial side, right: abaxial side; b. Lip, adaxial side: c. Column and

lip, lateral view: d. Anther, left: adaxial side, right: abaxial side: e. Pollinia, left: single

pair, right: two pairs. — Flower from the top part of the rhachis: f. Lip, left: adaxial side,

right: abaxial side: g- Column and lip, lateral view. — b. From Bogor comm. 153: a. c—

g. From Jongejan cult. 4031 (both spirit samples).

Bulbophyllum Sect. Altisceptrum and Hirtula

Figure 27. Bulbophyllum mirabile Hallier. (136)

76 Gard. Bull. Singapore 54 (2002)

the base, elsewhere glabrous to sparsely ciliate, surface glabrous; adaxially

slightly concave and with two calli near the base, convex beyond two short

ridges together forming a ‘v’; abaxially with a rounded ridge near the base,

convex towards the tip.

Colours: Rhachis and sepals pale greenish with pale purple dots.

Petals translucent whitish or yellowish, with purple dots and cilia. Lip

whitish, adaxially suffused with reddish purple, hairs purple.

Habitat & ecology: Epiphyte. Alt. 200—300 m.

Distribution: MALAYSIA: Peninsula (1 specimen seen, a doubtful

record, see below). BRUNEI (1). INDONESIA: Kalimantan (1).

Notes: The only Bulbophyllum with dimorphic flowers. The two forms

occupy separate parts of the rhachis; a short transitional area carries a few

flowers of intermediate shape. The flowers along the basal part of the

rhachis seem to be fertile; those along the top part often (not always) have

a malformed and incomplete column with neither pollinia nor stigma. In

such flowers, the lip and the petals may be immovably fused with the

column. Near the tip of the rhachis the flowers gradually but conspicuously

decrease in size. |

The specimen Hort. Glasnevin s.n. is marked ‘Malaya’. In the past,

this was used for what is now Peninsular Malaysia, but the species has not

been collected there since. The record is considered doubtful.

In spite of having been in cultivation on three independent occasions, no

vegetative parts have been preserved except for one leaf. Details have

been copied from Hallier’s original description.

137. Bulbophyllum spadiciflorum Tixier

Bulbophyllum spadiciflorum Tixier, Adansonia 6 (1966) 449, incl. line drawing; Tran

Hop, Orch. Vietnam (1998) 126, colour photograph 33. — Osyricera spadiciflora

(Tixier) Garay, Hamer & Siegerist, Nordic J. Bot. 14 (1994) 643. — TYPE:

Vietnam, Quang Duc, piste de Fyan, Hort. Paris 174/65 (P, holo., not seen).

Rhizome c. 8 mm diam., sections between pseudobulbs 1.5—2 cm

long. Pseudobulbs distant, ovoid, c. 1.2 x 1 cm. Petiole c. 2 cm. Leaf blade

elliptic to obovate, c. 25 x 2.5—3 cm, index (length/width) 8.3—10; obtuse.

Inflorescence an elongated, lax raceme, c. 50 cm. Peduncle erect to patent,

c. 30 cm, bracts c. 5, the longest c. 20 mm long. Rhachis nodding, swollen, +

circular in section, c. 20 x 0.5—0.6 cm. Floral bracts c.2 mm long, acuminate.

Flowers: anthesis from the top of the rhachis downwards. Median sepal

porrect, triangular, c. 4 x 0.8 mm, index c. 5, margins ciliate. Lateral sepals

c.4x 2 mm, index c. 2; acuminate. Petals porrect, obovate, c. 1.6 x 0.6 mm,

index c. 2.7; rounded, margins ciliate except near the base, base rather

narrowly attached. Lip recurved in the proximal half, obovate, c. 3.5 mm

Bulbophyllum Sect. Altisceptrum and Hirtula 77

long, rounded, margins with long hairs towards the tip; adaxially somewhat

concave near the base, without a median ridge, surface papillose; abaxial

surface glabrous. Column c. 1.7 mm long, column foot abaxially without a

spur near the tip. Stelidia obtuse. Anther: front margin ciliate, abaxially

papillose.

Colours: Rhachis green. Sepals and petals yellowish green with purple

veins. Lip purple.

Habitat & ecology: Alt. 5|00—1000 m. Flowering observed in May.

Distribution: VIETNAM: Central part.

Notes: No material available, although the species has been found

again recently (Tran Hop, 1998). The type could not be found in P. The

information above has been compiled from Tixier’s description and

illustration.

138. Bulbophyllum polycyclum J.J. Verm. — Fig. 28, plate 22.

Bulbophyllum polycyclum J.J. Verm., Blumea 41 (1996) 351. — TYPE: Indonesia,

Kalimantan Timur, Apo Kayan, Gunung Sungei Pendan area, Hort. Leiden

913457 (L, holo.).

Rhizome 3—4 mm diam., sections between pseudobulbs 0.9—1.2 cm

long. Pseudobulbs touching, ovoid to depressed-conical, 1.2—1.5 x 1.3—

1.8 cm. Petiole 3—4 cm. Leaf blade elliptic to ovate, 12.5—16 x 4.5—6.5

cm, index (length/width) 2.2—2.4; obtuse. Jnflorescence an elongated, rather

lax raceme with the flowers spirally arranged towards the base of the

inflorescence, arranged in often imperfect whorls of 5—8 or in interrupted

spirals towards the tip, entirely pendulous, 50—75 cm, 70—90-flowered.

Peduncle 17—40 cm, bracts 5—6, the longest 8—10 mm long. Rhachis

swollen, + circular in section, with the flowers on low-conical protuberances,

33—36 x 0.3—0.4 cm, + glabrous. Floral bracts triangular, 1.2—1.5 x 1—1.3

mm, acute. Flowers not much opening, a few simultaneously. Pedicel and

ovary 2.2—3 mm long, basal node on a c. 0.5 mm-long stump. Median

sepal + porrect, triangular, 3.6—4.5 x 1.5—2 mm, index 1.9—2.7; obtuse,

margins very finely papillose, base rather narrowly attached; rather thick,

surface adaxially glabrous, abaxially very finely papillose-hirsute. Lateral

sepals slightly recurved, 3.3—4 x 2.5—3 mm, index 1.2—1.6; otherwise as

the median sepal. Petals porrect, ovate to spathulate, with a widened base,

1.9—2 x 0.6—1 mm, index 2—3.2; rounded, margins ciliate, base rather

broadly attached; thin, surface glabrous. Lip abruptly recurved slightly

over half-way, elliptic-obovate, 2.2—2.9 x 1.5—1.8 mm, index 1.5—2,

rounded; margins ciliate with the cilia longest c. half-way up the lip: very

thick, surface + glabrous; adaxially concave and with two weak calli

continuing to half-way up the lip as weak ridges near the base, with a

78 Gard. Bull. Singapore 54 (2002)

median slit over most of its length ending in a small, circular plug that can

be removed and leaving a deep conical pit, with two distinct ridges diverging

proximally and continuing over c. 1/2 of the lip, surface distally distinctly

convex and overtopping the front margin; abaxially with a wide, truncate

ridge towards the base, convex towards the tip. Column 1.8—2 mm long,

stigma with 2 large lateral calli at its base, column foot swollen, abaxially

without a spur near the tip. Stelidia falcate, triangular, c. 0.7 mm long,

acute, with an antrorse, deltoid, obtuse tooth along their upper margin and

a similar but triangular, acute tooth along their lower margin. Anther:

front margin ciliate, abaxial ridge glabrous.

Colours: Rhachis dark green. Sepals and petals greenish, heavily

stained with blackish purple, particularly towards the margins and near the

veins. Lip yellowish, stained with blackish purple.

Habitat & ecology: Epiphyte in kerangas forest of c. 20 m high, with

a mossy floor. Alt. 850—1700 m.

Distribution: MALAYSIA: Sabah (1 specimen seen). INDONESIA:

Kalimantan (1).

139. Bulbophyllum lindleyanum Griff. — Fig. 29, plate 23.

Bulbophyllum (“Bolbophyllum”) lindleyanum Gritf., Notulae 3 (1851) 287. — Phyllorchis

lindleyana (Griff.) O. Kuntze, Rev. Gen. Pl. 2 (1891) 677. — TYPE: Myanmar,

Mergui, Griffith 697 (not seen).

Bulbophyllum rigens Rchb.f., Hamburger Garten- Blumenzeitung 21 (1865) 296. —

Phyllorchis rigens (Griff.) O. Kuntze, Rev. Gen. Pl. 2 (1891) 677. — TYPE:

locality unknown, Herb. Reichenbach 2252 (Day 63) (W, holo.).

Bulbophyllum caesariatum Ridl., Bull. Misc. Inform., Kew (1924) 202. -SYNTYPES:

Thailand, southwest coast, Kopah Jan-Jan, Haniff & Nur SFN 2702 (SING,

lecto.; K, isolecto., the latter not seen); 2750 (K, not seen).

Rhizome 2.5—3.5 mm diam., sections between pseudobulbs 1.2—2.5

cm long. Pseudobulbs touching, ovoid to globose, 1.2—3 x 1—2.5 cm. Petiole

1.8—3.5 cm. Leaf blade elliptic to ovate, 6—18 x 1.5—3 cm, index (length/

width) 4—6.8; acute. Inflorescence an elongated, rather lax raceme, 11—28

cm, 9—25-flowered. Peduncle erect to patent, 6—19 cm, bracts c. 6, the

longest 4.8—13 mm long. Rhachis nodding, not thickened, 3.5—10 cm,

hispid with short, stiff hairs. Floral bracts triangular, 5—8 x 1.2—2.5 mm,

acuminate. Flowers opening fully, many simultaneously. Pedicel and ovary

7—11 mm long, hispid as the rhachis, basal node on a 0.3—1 mm-long

Figure 28. Bulbophyllum polycyclum J.J. Verm. (138) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, adaxial side; e.

Lip, abaxial side; f. Column and lip, lateral view; g. Anther, above: abaxial side, below:

adaxial side; h. Pollinia, above: single pair, below: two pairs. — All from Hort. Leiden

913457 (spirit sample).

Bulbophyllum Sect. Altisceptrum and Hirtula

Figure 28. Bulbophyllum polycyclum J.J. Verm. (138)

80 Gard. Bull. Singapore 54 (2002)

stump. Median sepal recurved, ovate, 5.1—6.5 x 2.2—3.5 mm, index 1.7—

2.5; acute to acuminate, margins somewhat papillose, sparsely ciliate, base

broadly attached; rather thin, surface locally very finely papillose, adaxially

pilose with long, thin hairs, abaxially hispid as the rhachis. Lateral sepals

ovate-triangular, 6—8 x 3—4 mm, index 1.5—2; acuminate, upper margin

without cilia; otherwise as the median sepal. Petals somewhat recurved,

elliptic to ovate, 1.5—2.3 x 1.2—1.5 mm, index 1.1—2; (sub-)acute, margins

ciliate, base broadly attached; thin; surface glabrous but abaxially pilose

near the tip. Lip abruptly recurved slightly below half-way, ovate-oblong,

2—3 x 1.2—1.8 mm, index 1.3—2.1 (all without artificial spreading);

rounded, margins ciliate in the distal 2/3; rather thin, surface + glabrous;

adaxially almost flat proximally, somewhat convex distally; abaxially with

a weak, rounded ridge proximally, slightly convex distally; basal auricles

absent or inconspicuous. Column 2.5—3 mm long, foot slightly swollen.

Stelidia subulate, 1—1.7 mm long, subacute, finely papillose distally, near

the base with a minute, obtuse tooth along their upper margin, and a

distinct, deltoid, rounded to obtuse wing along their lower margin. Anther:

front margin and abaxial side papillose.

Colours: Sepals and petals greenish yellow to white, the sepals with

blackish purple to dark brown veins and sometimes a similarly coloured

tip, the petals with a similarly coloured or a greenish vein. Lip greenish

yellow with greyish hairs.

Habitat & ecology: In forest, also found as an epiphyte in old rubber

plantations among limestone hills. A lowland species. Flowering observed

in Jan, Nov, Dec.

Distribution: MYANMAR: Moulmein (2 specimens seen); Mergui

(fide Griffith); ‘S part, close to Thai border’ (1). THAILAND: Peninsula

(6).

Notes: Two sheets: Herb. Reichenbach (Foerstermann) 15563, and

Foerstermann 08 (both W) are marked “Sarawak”. On the first sheet another

locality: “Birma, Bhamo” is also given but crossed out. Foerstermann has

collected both in Myanmar (Burma) as well as in Sarawak, nevertheless I

assume that both sheets are wrongly labeled because the species has never

been found again in Sarawak, and the locality is far outside its known

range.

Figure 29. Bulbophyllum lindleyanum Griff.(139) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, adaxial side; e.

Lip, abaxial side; f. Column and lip, lateral view; g. Anther, left: adaxial side, right:

abaxial side; h. Pollinia, above: two pairs, below: single pair. — All from FF 48 (spirit

sample).

Bulbophyllum Sect. Altisceptrum and Hirtula

3mm

Oe Se

4mm

J , i S P ~

nik, Uy é

A Y F

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Figure 29. Bulbophyllum lindleyanum Griff. (139)

82 Gard. Bull. Singapore 54 (2002)

140. Bulbophyllum tremulum Wight — Fig. 30, plate 24.

Bulbophyllum (“Bolbophyllum”) tremulum Wight, Icon. Pl. Ind. or. 5, 1 (1851) t. 1749;

Joseph, Rec. Bot. Surv. India 22 (1982) 92. — Phyllorchis tremula (Wight) O.

Kuntze, Rev. Gen. Pl. 2 (1891) 678. — TYPE: India, Nilgiri Hills, Jerdon s.n.

(not seen).

Rhizome 1.8—3.2 mm diam., sections between pseudobulbs 1.2—3

cm long. Pseudobulbs close or distant, ovoid or low-conical, 0.7—1.5 x

0.8—1.7 cm, surface somewhat corrugated. Petiole 0.1—0.7 cm. Leaf blade

ovate, 2—8.5 x 0.6—1.6 cm, index (length/width)2.7—4.3; obtuse.

Inflorescence an elongated, lax raceme, 7—26 cm, 9—13-flowered. Peduncle

erect to patent, 5—9 cm, bracts 3—4, the longest 4—8.5 mm long. Rhachis

erect to arching, not thickened, 2—7 cm, glabrous. Floral bracts triangular,

1.2—3 x 0.7—1.2 mm, acute. Flowers + distichous, secund, opening wide,

many simultaneously. Pedicel and ovary 7—8 mm long, basal node on a

0.3—0.6 mm-long stump. Median sepal spreading to reflexed, triangular,

7—12 x 1.7—2.8 mm, index 3.7—5.5; acute, margins ciliate, base rather

narrowly attached; thin, adaxial surface pubescent towards the base,

abaxially glabrous. Lateral sepals 8—13.5 x 2—4.2 mm, index 2.6—4.5;

upper margin sparsely ciliolate; otherwise as the median sepal. Petals

porrect, elliptic-ovate with a wide base, 1.8—2.6 x 0.8—0.9 mm, index 2—

2.9; obtuse to acute, margins ciliate, base broadly attached; thin, surface

glabrous but pubescent adaxially near the tip. Lip recurved near the base,

oblong, 5.5—8 x 1.7—2 mm, index 3.2—4.7; rounded to obtuse, margins

ciliate proximally; lip thick, very thick near the base; adaxially somewhat

concave and with two calli near the base, with two distinct ridges diverging

proximally and continuing parallel and close together over c. 1/2 of the lip,

adaxial surface smooth to distinctly verrucose, glabrous; abaxially with a

rounded ridge near the base, surface otherwise + flat, with numerous

spindle-shaped hairs arising from tiny pits, surface otherwise glabrous.

Column 1.5—2 mm long, stigma distinctly protruding at its base. Stelidia

0.5—0.7 mm long, obtuse, + rectangular in outline with an antrorse, deltoid,

obtuse tooth along the upper margin, and a deltoid, rounded to obtuse

wing along the lower margin. Anther: front margin and abaxial ridge with

distinct, elongated papillae.

Colours: Sepals and petals yellowish or greenish, veins and margins

partially or entirely purple; the petals sometimes entirely suffused with

Figure 30. Bulbophyllum tremulum Wight. (140) — a. Habit; b. Flower; c. Flower analysis,

from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial side, right:

abaxial side; e. Column and lip, lateral view; f. Anther, above: abaxial side, below:

adaxial side; g. Pollinia, above: single pair, below: two pairs. — All from Jongejan cult.

4544 (spirit sample).

Bulbophyllum Sect. Altisceptrum and Hirtula 83

4mm

Y Ui, Yip AA | | ! f ) o% pat = é Bs Ba

“if ey HI | aN? Pe .

j ty Waly, :

Yyf iy | WY ; y | I

y fy AWAY 4 Y } Y Y

y Wy Lif j Hf, Hy

YY yi MN lyf

Figure 30. Bulbophyllum tremulum Wight (140)

84 Gard. Bull. Singapore 54 (2002)

purple. Lip white, yellow or brownish, sometimes suffused with purple

near the base, or entirely dark purple. Hairs along the margins white,

those on the abaxial side of the lip purple.

Habitat & ecology: Alt. 800—1100 m. Flowering in Jan, Feb, Sep,

Dec (Feb, Apr in cultivation).

Distribution: INDIA: south-west part: Nilgiris, Silent Valley (4

specimens seen).

Notes: Data from Joseph (1982) have been included in the description.

141. Bulbophyllum scaphiforme J.J. Verm., sp. nov. — Fig. 31.

Bulbophyllum scaphiforme J.J. Verm., a Bulbophyllo nigrescenti labello marsupiiformi

differt. - TYPE: Thailand, Doi San Yao, Seidenfaden & Smitinand (GT) 7464

(C, holo.).

Bulbophyllum nigrescens sensu Seidenf. & Smitin., Orch. Thailand 3 (1961) 426 (partly,

incl. fig. 319); Seidenf., Dansk Bot. Ark. 33, 3 (1979) 104 (partly, incl. fig. 64);

Tran Hop, Orch. Vietnam (1998) 113, colour photograph 30; S.C. Chen et al.,

Native Orch. China (1999) 35 (upper left and right colour photographs only),

non Rolfe.

Rhizome: sections between pseudobulbs 1.5—2.5 cm long.

Pseudobulbs close, depressed conical, 1.5—2.5 x 1.5—3 cm. Petiole 1—1.5

cm. Leaf blade elliptic, 7—9.2 x c. 1.8 cm, index (length/width) c. 3.9;

?subacute. Inflorescence an elongated, rather lax raceme, erect to patent,

28—54 cm, 23—33-flowered. Peduncle 14—25 cm. Rhachis not thickened,

14—22 cm long, glabrous. Floral bracts triangular, 3—4 x c. 2 mm, acute.

Flowers spreading to reflexed, secund, not fully opening, many

simultaneously. Pedicel and ovary 5.5—9 mm long, basal node on a 0.8—2

mm-long stump. Median sepal porrect to recurved, ovate, 4.5—7.5 x 2—4.2

mm, index 2.3—3.8; acute, margins ciliate, base rather broadly attached;

rather thin, adaxially pubescent, abaxially glabrous. Lateral sepals recurved,

elliptic to ovate, 6—8 x 2.5—4.2 mm, index 1.9—2.4; acute to acuminate,

otherwise as the median sepal. Petals + porrect, spathulate, 2—2.5 x 0.8—

1.5 mm, index 1.6—2.5; acute to acuminate, margins ciliate distally, base

broadly attached; rather thick with a thick top, surface pubescent distally.

Lip recurved in the proximal half, pouch-shaped distally, obovate in outline,

4.8—5.5 x 1.8—3.2 mm, index 1.6—2.8; rounded, margins ciliate; rather

thin; adaxially somewhat concave and with a median slit reaching up to c.

Figure 31. Bulbophyllum scaphiforme J.J. Verm. (141) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial

side, right: abaxial side; e. Column and lip, lateral view; f. Anther, left: adaxial side,

right: abaxial side; g. Pollinia, above: a single pair, below: two pairs. — a (plant).

Modified after Seidenfaden & Smitinand (1961); a (inflorescence). From Tixier s.n.

(herbarium specimen); b—g. From GT 7464 (spirit sample).

Bulbophyllum Sect. Altisceptrum and Hirtula

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Figure 31. Bulbophyllum scaphiforme J.J. Verm. (141)

86 Gard. Bull. Singapore 54 (2002)

1/3 of the lip near the base, with 2 distinct, obtuse ridges diverging

proximally, running + parallel over c. 3/4 of the lip, and ending in some

irregularly shaped calli, surface with irregular, radiating furrows distally

but otherwise glabrous; abaxially with a wide, rounded ridge near the base,

surface otherwise convex, pubescent, or only so in front of the basal ridge

and towards the margins. Column 1—1.6 mm long, stigma with a distinct,

obtuse tooth slightly below its base. Stelidia triangular, 0.4A—0.7 mm long,

acute. Anther: front margin and abaxially towards the tip densely papillose.

Colours: Sepals blackish purple, yellowish towards the base, or

yellowish with blackish purple veins, margins and scattered spots. Petals

blackish purple distally, yellowish proximally, often with a blackish purple

vein. Lip blackish purple with green or yellowish ridges adaxially. Anther

yellow.

Habitat & ecology: Alt. 1100-1400 m. Flowering in Mar—May, Jul.

Distribution: CHINA: Yunnan (fide S.C. Chen et al., under B.

nigrescens). THAILAND (5 specimens seen). VIETNAM: southern part

(Ly,

Notes: This species has been confused with Bulbophyllum nigrescens

by Seidenfaden & Smitinand (1961), and Seidenfaden (1979). It is clearly

distinct from this species in having a pouch-shaped (or boat-shaped) lip.

The specific epithet refers to that character.

No vegetative parts are available, the description has been completed

from Seidenfaden & Smitinand (1961, fig. 319, sub B. nigrescens).

142. Bulbophyllum nigripetalum Rolfe — Fig. 32, plate 25.

Bulbophyllum nigripetalum Rolfe, Bull. Misc. Inform., Kew (1891) 197; Summerh.,

Kew Bull. 6 (1952) 471. — SYNTYPES: “said to have been imported (...) from

the West Coast of Africa”, later shown to originate from Asia by Summerhayes

(1952), O’Brien cult. (1891) (K, lecto.), O’Brien cult. (1889) (K).

Bulbophyllum secundum sensu Seidenf., Dansk Bot. Ark. 33, 3 (1979) 103 (partly, incl.

syn. B. nigripetalum), non Hook.f.

Bulbophyllum nigrescens sensu S.C. Chen et al., Native Orch. China (1999) 35 (lower

left colour photograph only), non Rolfe.

Rhizome 1.5—3 mm diam., sections between pseudobulbs. 0.6—2.5

cm long. Pseudobulbs close, ovoid or low-conical, 0.9—1.7 x 0.9—1.8 cm.

Petiole 0.8—2 cm. Leaf blade elliptic, 5.2—17 x. 0.6—1.5 cm, index (length/

Figure 32. Bulbophyllum nigripetalum Rolfe (142) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial

side, right: abaxial side; e. Lip, adaxial side; f. Column and lip, lateral view; g. Anther,

left: abaxial side, right: adaxial side; h. Pollinia, above: two pairs, below: single pair. —

a. From Henry 13055 (herbarium specimen); b—d, f—h. From Morrison 1032 (spirit

sample); e. From Maxwell 89-524 (herbarium specimen).

Bulbophyllum Sect. Altisceptrum and Hirtula 87

2mm

ee eee

lmm

Figure 32. Bulbophyllum nigripetalum Rolfe (142)

88 Gard. Bull. Singapore 54 (2002)

width) 6—11.4; obtuse. /nflorescence an elongated, rather lax raceme, erect

to patent, 15—49 cm, 13—32-flowered. Peduncle 8—25 cm, bracts c. 5, the

longest 4—9 mm long. Rhachis not thickened, 5.5—25 cm long, glabrous.

Floral bracts triangular, 1.8—4.5 x 1—2 mm, acuminate. Flowers spreading

to reflexed, secund, not fully opening, many simultaneously. Pedicel and

ovary 3.8—6.5 mm long, basal node on a 0.5—0.8 mm-long stump. Median

sepal porrect to recurved, ovate to triangular, 3.1—6 x 1.3—2.2 mm, index

2—3.6; obtuse to acute, margins ciliate, base rather narrowly attached;

rather thin, adaxially pubescent, abaxially glabrous. Lateral sepals recurved,

3—6 x 2.2—3 mm, index 1.3—2.2; otherwise as the median sepal. Petals +

porrect, elliptic to obovate or spathulate, 1.6—2.2 x 0.5—1 mm, index

1.8—3.6; obtuse to acuminate, margins ciliate distally, base broadly attached:

rather thin with a thick tip, surface pubescent distally. Lip recurved in the

proximal half, obovate to elliptic, 2.83—4.2 x 1—1.6 mm, index 1.8—3;

rounded to obtuse, margins ciliate; thick and fleshy; adaxially somewhat

concave near the base, with 3 ridges: 2 distinct, obtuse ridges diverging

proximally, continuing over 1/3—3/4 of the lip, and a thin, low, sometimes

inconspicuous median ridge continuing over c. 1/2 of the lip, surface

glabrous, finely velutinous towards the margins; abaxially with a wide,

rounded ridge near the base, surface elsewhere convex, partly or entirely

velutinous. Column 1—1.3 mm long, stigma distinctly protruding at its

base, or with a tooth slightly below its base. Stelidia triangular, 0.3—0.6

mm long, acute. Anther: front margin and abaxially towards the tip densely

papillose.

Colours: Sepals cream-coloured or yellow, near the base often with

blackish purple veins, sometimes blackish purple distally, or blackish purple

all over. Petals blackish purple, sometimes yellowish with a purple vein

proximally. Lip blackish purple, often yellowish at the base. Flowers with a

lemon or turpentine scent (1 obs.)

Habitat & ecology: Epiphyte or lithophyte in forest. Alt. 1000-1300

m. Flowering in Mar—May.

Distribution: CHINA: Yunnan (1). THAILAND (3 specimens seen).

Notes: Erroneously synonymised with Bulbophyllum secundum by

Seidenfaden (1979); it differs in having non-resupinate flowers, and adaxially

pubescent sepals. It is also of larger stature.

Similar to B. nigrescens, it differs in having a shorter lip that is bi-

convex and fleshy towards the tip. The adaxial surface is ?always (sometimes

difficult to see in herbarium specimens) very finely velutinous towards the

margins (glabrous in B. nigrescens). An additional difference may be that

B. nigripetalum has a thin, sometimes inconspicuous median ridge instead

of a median slit, as in B. nigrescens. The differences between the two seem

Bulbophyllum Sect. Altisceptrum and Hirtula 89

minor, but so far all material can be assigned unequivocally to one or the

other species.

143. Bulbophyllum nigrescens Rolfe — Fig. 33, plate 26, 27.

Bulbophyllum nigrescens Rolfe, Bull. Misc. Inform., Kew (1910) 158; Seidenf. & Smitin.,

Orch. Thailand 3 (1961) 426 (partly, excl. fig. 319 [is B. scaphiforme]); Seidenf.,

Dansk Bot. Ark. 33, 3 (1979) 104 (partly, excl. fig. 64 [is B. scaphiforme]). —

TYPE: Thailand, Chiangmai, Doi Suthep, Kerr 84 (K, holo.).

Bulbophyllum angusteellipticum Seidenf., Nordic J. Bot. 1 (1981) 209. - TYPE: Thailand,

Doi Inthanond, Seidenfaden & Smitinand (GT) 8310 (C, holo., not seen).

[Not Bulbophyllum nigrescens Schltr, Repert. Spec. Nov. Regni Veg. Beih. 1 (1913)

839 (= Bulbophyllum rubromaculatum W. Kittr., Bot. Mus. Leafl., Harvard

University 30 [1981] 100).]

[Not Bulbophyllum nigrescens sensu Tran Hop, Orch. Vietnam (1998) 113, col. phot.

30 (= B. scaphiforme); S.C. Chen et al., Native Orch. China (1999) 35 (= B.

nigripetalum and B. scaphiforme).|

Rhizome 2—3.5 mm diam., sections between pseudobulbs 0.6—2.5

cm long. Pseudobulbs close, ovoid or low-conical, 0.8—1.7 x 1—2 cm.

Petiole 0.8—2 cm. Leaf blade elliptic to ovate, 6—13 x 0.9—1.7 cm, index

(length/width) 4—8.4; obtuse. /nflorescence an elongated, rather lax raceme,

erect to patent, 23—44 cm, 13—27-flowered. Peduncle 8—22 cm, bracts c.

5, the longest 6.5—9 mm long. Rhachis not thickened, 9—20 cm long,

glabrous. Floral bracts triangular, 1.8—4.5 x 1—2 mm, acuminate. Flowers

spreading to reflexed, secund, not fully opening, many simultaneously.

Pedicel and ovary 5—11 mm long, basal node on a 0.5—0.8 mm-long stump.

Median sepal porrect to recurved, ovate to triangular, 5.5—95 x 1.8—3.2

mm, index 1.8—5; obtuse to acute, margins ciliate, base rather narrowly

attached; rather thin, adaxially pubescent, abaxially glabrous. Lateral sepals

recurved, 6.5—10.5 x 2.7—4.2 mm, index 1.6—3.4; otherwise as the median

sepal. Petals + porrect, elliptic to obovate or spathulate, 1.5—2.5 x 0.8—1.5

mm, index 1.3—3.2; obtuse to acuminate, margins ciliate distally, base

broadly attached; rather thin with a thick top, surface pubescent distally.

Lip recurved in the proximal half, obovate to elliptic, 5—7 x 2—3.3 mm,

index 1.8—3.5; rounded to obtuse, margins ciliate; rather thin; adaxially

somewhat concave near the base and with a median slit or furrow reaching

up to 1/3—2/3 of the lip, with 2 distinct, obtuse ridges diverging proximally,

continuing over 1/3—3/4 of the lip, and often ending in a patch of irregularly

shaped verrucae, surface glabrous; abaxially with a wide, rounded ridge

near the base, surface elsewhere concave to flat, glabrous or partly

pubescent. Column 1.4—1.8 mm long, stigma distinctly protruding at its

base, or with a distinct, obtuse to acute tooth slightly below its base. Stelidia

triangular, 0.5—0.8 mm long, acute. Anther: front margin and abaxially

90 Gard. Bull. Singapore 54 (2002)

towards the tip densely papillose.

Colours: Sepals yellowish, stained with blackish purple and with

blackish purple veins; sometimes entirely blackish purple adaxially and

pale reddish brown abaxially. Petals and lip almost entirely blackish purple;

the petals proximally sometimes yellow with a blackish purple vein.

Habitat & ecology: Epiphyte or lithophyte in deciduous or evergreen

forest. Alt. 700-1500 m. Flowering in Jan—May, Jul.

Distribution: THAILAND (8 specimens seen). VIETNAM (1).

Notes: This has been confused with Bulbophyllum scaphiforme by

Seidenfaden & Smitinand (1961), and Seidenfaden (1979), and with B.

scaphiforme and B. nigripetalum by S.C. Chen et al. (1999). The differences

are given below the species mentioned.

The identity of B. angusteellipticum has been surmised from an almost

identical sample from the type locality.

144. Bulbophyllum trulliferum J.J. Verm. & A. Lamb - Fig. 34, plate 28.

Bulbophyllum trulliferum J.J. Verm. & A. Lamb, Blumea 38 (1994) 337. — TYPE:

Malaysia, Sabah, Crocker Range, Tenom Orchid Centre (T.O.C.) cult. 2321 (L,

holo.).

Rhizome 1.8—2.5 mm diam., sections between pseudobulbs 0.6—1

cm long. Pseudobulbs + touching, depressed conical to almost discoid,

0.4—0.7 x 1.2—2 cm. Petiole c. 1 cm. Leaf blade elliptic, 8.5—17 x 0.8—1.2

cm, index (length/width) 10—14.2; acute. Inflorescence an elongated, rather

dense raceme, 29—32 cm, 10—14-flowered. Peduncle erect to patent, 27—

29 cm, bracts c. 5, the longest c. 5 mm long. Rhachis nodding, not thickened,

2—3 cm, glabrous. Floral bracts ovate, c. 1 x 0.8 mm, acute. Flowers not

fully opening, many simultaneously. Pedicel and ovary 2.5—3 mm long,

finely hirsute, basal node + flush with the surface of the rhachis. Median

sepal porrect, ovate, 3—3.3 x 2—2.6 mm, index 1.2—1.5; rounded to obtuse,

margins ciliate, base rather broadly attached; rather thin, surface adaxially

glabrous, abaxially finely hirsute. Lateral sepals recurved, 3—3.8 x 2—3

mm, index 1.3—1.5; obtuse, margins papillose; otherwise as the median

sepal. Petals porrect, triangular, c. 1.5—2.5 x 0.5—0.7 mm, index 3—4.3;

subacute, margins ciliate, base broadly attached; thin; surface glabrous.

Figure 33. Bulbophyllum nigrescens Rolfe (143) — a. Habit; b. Flower; c. Flower analysis,

from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial side, right:

abaxial side; e, f, g. Lip, adaxial side; h, i. Column and lip, lateral view; j. Anther, left:

abaxial side, right: adaxial side; k. Pollinia, above: single pair, below: two pairs. — a—d,

h, j, k. From Jongejan cult. 259] (spirit sample); e. From Cumberlege 102] (herbarium

specimen); f, i. From Hort. Aarhus 78-B-632 (spirit sample); g. From Maxwell 89-523

(herbarium specimen).

Bulbophyllum Sect. Altisceptrum and Hirtula 9]

Bren dee el 266 RKeS

week pee rept ®

oo taeps

Smm

3mm

Imm

Figure 33. Bulbophyllum nigrescens Rolfe (143)

92 Gard. Bull. Singapore 54 (2002)

Lip abruptly recurved c. half-way, oblong to subspathulate, 2—2.5 x 1.1—

1.4 mm, index 1.5—2.1; rounded, margins ciliate except near the base and

near the tip; very thick; adaxially concave and with a bifid callus near the

base, with 2 distinct, rounded ridges diverging proximally and distally,

distally enclosing a circular depression, adaxial surface glabrous; abaxially

with a wide, retuse ridge proximally, convex and with thick, stiff, long hairs

towards the margins distally, surface otherwise glabrous. Column 1.6—1.7

mm long, stigma protruding at its base. Stelidia narrowly triangular, c. |

mm long, acute, with a small, deltoid, obtuse tooth along the upper margin,

and a larger, antrorse, triangular obtuse tooth along the lower margin,

near the base. Anther: front margin ciliate, abaxial surface with an

inconspicuous ridge only, + glabrous.

Colours: Flowers cream-coloured; sepals with a reticulate pattern of

pale purple veins; petals with the vein, margin and cilia pale purple; lip

orange towards the base, adaxially with a large, blackish spot near the tip;

hairs white.

Habitat & ecology: Epiphyte in primary forest. Alt. c. 900 m. Flowering

observed in Sep.

Distribution: MALAYSIA: Sabah, Crocker Range (3 specimen seen).

145. Bulbophyllum jolandae J.J. Verm. — Fig. 35.

Bulbophyllum jolandae J.J. Verm., Orch. Borneo 2 (1991) 241, colour photograph pl.

16f. — TYPE: Malaysia, Sabah, Ulu Padas, Vermeulen 602 (L, holo.).

Rhizome c. 3 mm diam., sections between pseudobulbs 0.6—0.8 cm

long. Pseudobulbs touching, ovate, 1.2—1.5 x 1—1.2 cm. Petiole 2—2.5 cm.

Leaf blade elliptic, 8.5—9.5 x 2—2.3 cm, index (length/width) 44.3; acute.

Inflorescence an elongated, very lax raceme, 25—42 cm, 4—14-flowered.

Peduncle erect to patent, 20—22 cm, bracts c. 4, the longest c. 6 mm long.

Rhachis not thickened, erect or arching, 5—21 cm long, glabrous. Floral

bracts triangular, 2.5—3 x 1 mm, acuminate. Flowers opening wide, several

simultaneously, anthesis from the base of the rhachis upwards. Pedicel and

ovary 48—S5S0 mm long, basal node on a c. 1.5 mm-long stump. Median

sepal reflexed, triangular, c. 18.5 x 2 mm, index 9.2—9.3; caudate, margins

ciliolate near base, base rather broadly attached; thin, surface glabrous.

Lateral sepals ovate-triangular, c. 20 x 4.5 mm, index 4.4—4.5; acute-

Figure 34. Bulbophyllum trulliferum J.J. Verm. (144) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial

side, right: abaxial side; e. Column and lip, lateral view; f. Anther, left: adaxial side,

right: abaxial side; g. Pollinia, above: two pairs, below: single pair. — a. Modified after a

sketch and a photograph of A. Lamb; b—g. All from Tenom Orchid Centre cult. 2321

(spirit sample).

Bulbophyllum Sect. Altisceptrum and Hirtula 93

RUC PRs

2mm

—— — ee ee eo

Figure 34. Bulbophyllum trulliferum J.J. Verm. (144)

94 Gard. Bull. Singapore 54 (2002)

acuminate, upper margin glabrous; otherwise as the median sepal. Petals

porrect, falcate, triangular, c. 4.2 x 1.2 mm, index 3.5; acuminate, margins

ciliate except near the tip, base broadly attached; thin, surface glabrous.

Lip curved in the basal half, + elliptic to ovate, c. 7.5 x 2.8 mm, index 2.6—

2.7; rounded, margins ciliate, and with long, thin, + straight hairs except

near the base and near the tip; lip rather thin with a thick base, surface

glabrous; adaxially concave near the base, with a median slit proximally

opening into a small cavity and distally continuing over c. 1/3 of the lip,

with two rounded ridges continuing over c. 1/3 of the lip, proximally

diverging and almost fused to a single median ridge distally; abaxially with

a retuse ridge near the base. Column c. 2 mm long, stigma protruding at its

base. Stelidia c. 0.7 mm long, tip acute; + rectangular in outline with an

antrorse, deltoid, subacute tooth along the upper and lower margin. Anther:

front margin long ciliate, abaxial surface papillose.

Colours: Sepals proximally somewhat glaucous green, distally

yellowish, medium green in between. Petals greenish, purplish near the

base, margins and cilia purple. Lip purplish except for a paler patch near

the base, short hairs white, long hairs purple. Column greenish, blotched

purple.

Habitat & ecology: Epiphyte in moss, in open montane kerangas

forest. Alt. c. 1500 m. Flowering observed in Oct.

Distribution: MALAYSIA: Sabah, Ulu Padas (1 specimen seen).

146. Bulbophyllum comberipictum J.J. Verm., sp. nov. — Fig. 36.

Bulbophyllum comberipictum J.J. Verm.,a Bulbophyllo jolandae pedicello cum ovario

8—20 mm longo, labello pilis tenuibus lanatis differt. - TYPE: Malaysia, Sarawak,

Hose Mts., Hort. Leiden 914194 (L, holo.).

Bulbophyllum sp. Comber, Orchids of Sumatra (2001) 816, colour photograph.

Rhizome c. 3 mm diam., sections between pseudobulbs 0.6—1.2 cm

long. Pseudobulbs close or touching, ovate-conical, c. 1.5 x 1.5 cm. Petiole

c. 2.5 cm. Leaf blade elliptic, c. 10.5 x 3.1 cm, index (length/width) c. 3.3;

acute. Inflorescence an elongated, rather dense raceme, c. 32 cm, 26—31-

flowered. Peduncle erect to patent, c. 18.5 cm, bracts c. 4, the longest c. 7

mm long. Rhachis not thickened, c. nodding, c. 13.5 cm long, glabrous.

Floral bracts triangular, c. 2.5 x 1 mm, acuminate. Flowers opening wide,

several simultaneously, anthesis from the base of the rhachis upwards.

Pedicel and ovary 8—20 mm long, basal node on a c. 1 mm-long stump.

Figure 35. Bulbophyllum jolandae J.J. Verm. (145) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial

side, right: abaxial side; e. Column and lip, lateral view; f. Anther, above: abaxial side,

below: adaxial side. — All from Vermeulen 602 (living plant).

Bulbophyllum Sect. Altisceptrum and Hirtula

AGE

Figure 35. Bulbophyllum jolandae J.J. Verm. (145)

96 Gard. Bull. Singapore 54 (2002)

Median sepal c. porrect to reflexed, triangular, c. 7 x 1.4 mm, index c. 5;

tapering towards the tip and acute, base rather broadly attached; thin,

glabrous. Lateral sepals ovate-triangular, c. 7 x 2.2—2.4 mm, index 2.9—

3.2; subacute; otherwise as the median sepal. Petals porrect, falcate,

triangular, 2.7—2.8 x 0.6—0.9 mm, index 3.1—4.5; acuminate, margins ciliate

except near the base and near the tip, base broadly attached; thin, surface

glabrous. Lip curved in the basal half, + elliptic, c. 4.5 x 1.2—1.3 mm, index

3.4—3.8; truncate, margins ciliate near the base and at the tip, in between

with long, thin, distinctly wavy hairs; lip rather thin with a thick base,

surface glabrous; adaxially concave near the base, with a median slit

proximally opening into a small cavity and distally continuing over c. 4/5 of

the lip, with two rounded ridges continuing over c. 4/5 of the lip, diverging

proximally and almost fused to a single median ridge distally; abaxially

with a retuse ridge near the base. Column c. 1.3 mm long, stigma protruding

at its base. Stelidia c. 0.4 mm long, tip acute; + rectangular in outline with

an antrorse, deltoid, subacute tooth along the upper and lower margin.

Anther: front margin long ciliate, abaxial surface + glabrous.

Colours: Sepals creamy yellow, the median flushed with some purple

along the margins. Petals blackish purple, with purple cilia. Lip whitish,

marked with dark purple towards the tip and along the margins; hairs

purple.

Habitat & ecology: Recorded from submontane kerangas forest, and

from a steep, densely forested ravine cut into volcanic soil. Alt. 800—1300

m. Flowering observed in Nov.

Distribution: MALAYSIA: Sarawak, Hose Mts. (1 specimen seen).

INDONESIA: Sumatra (photo Comber, see below).

Notes: Similar to Bulbophyllum jolandae but distinct because of its

much denser inflorescence with much smaller flowers on shorter pedicels.

The lip has a more distinct and distinctly longer median ridge (reaching up

to 4/5 of the length of the lip), and distinctly woolly rather than straight

hairs.

For a while this species was known only from a photograph by

Comber in Sumatra; the specific name refers to this. It was not found again

in spite of an extensive search of the same locality. The photograph shows

a plant almost identical to the Sarawak specimen, but with narrower leaves.

Figure 36. Bulbophyllum comberipictum J.J. Verm. (146) — a. Habit; b. Flower; c.

Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, above:

adaxial side, below: abaxial side; e. Column and lip, lateral view; f. Anther, left: abaxial

side, right: adaxial side; g. Pollinia, above: two pairs, below: single pair. — All from

Hort. Leiden 914194 (spirit sample).

Imm

3mm

Bulbophyllum Sect. Altisceptrum and Hirtula

ii his Yih Li

NN x \\

Ss)

Figure 36. Bulbophyllum comberipictum J.J. Verm. (146)

98 Gard. Bull. Singapore 54 (2002)

147. Bulbophyllum lasioglossum Rolfe ex Ames — Fig. 37, plate 29.

Bulbophyllum lasioglossum Rolfe ex Ames, Orch. 1 (1905) 100. — TYPE: Philippines,

Luzon, Bataan, Mt. Mariveles, Lamao River, Borden 797 (PNH, holo., destroyed;

AMES, K, iso.).

Rhizome 1.8—3 mm diam., sections between pseudobulbs 0.6—0.8

cm long. Pseudobulbs + touching, ovoid, 0.7—1.5 x 0.5—0.8 cm. Petiole

1.4—2.6 cm. Leaf blade elliptic to ovate, 5.8—12 x 1.5—3.2 cm, index

(length/width) 2.6—4.7; (sub-)acute. Inflorescence an elongated, rather lax

raceme, 16—33 cm, 21—44-flowered. Peduncle erect to patent, 11.5—19

cm, bracts c. 5, the longest 6—8 mm long. Rhachis nodding, not thickened,

5—14.5 cm, glabrous. Floral bracts triangular, 1.5—3 x 0.8—1.2 mm, acute.

Flowers opening wide, several simultaneously, anthesis from the top of the

rhachis downwards. Pedicel and ovary 7.5—14 mm long, basal node on a

0.4—1 mm-long stump. Median sepal porrect to recurved, elliptic, 6.5—9 x

2—2.2 mm, index 3.2—4.1; acute to acuminate, margins ciliate, base rather

narrowly attached; thin, surface glabrous. Lateral sepals recurved or

spreading, ovate-triangular, 6.5—9 x 2.5—2.6 mm, index 2.6—3.6; margins

glabrous, base broadly attached; otherwise as the median sepal. Petals

porrect, ovate, 2.5—3.2 x 0.6—1 mm, index 3—4.6; acute, margins ciliate,

base broadly attached; thin, surface glabrous. Lip curved near the base,

obscurely 3-lobed; midlobe ovate, 5—7 x 1.2—2.2 mm, index 2.7—4.2;

rounded to obtuse, margins distally ciliate, otherwise with long, thin, straight

or slightly wavy hairs; rather thin with a thick base, surface glabrous;

adaxially concave near the base and with a median slit reaching up to 1/3

of the lip, with two obtuse ridges diverging proximally towards the distal

end of the lateral lobes, distally fused to a single median ridge and

continuing over c. 1/3 of the lip, surface otherwise slightly concave; abaxially

with a wide, retuse ridge near the base; /ateral lobes attached along the

proximal 1/6 of the lip, erect, semi-circular, margins ciliate, broadly attached,

rather thick, adaxially pubescent towards the margins, abaxially glabrous.

Column 1.2—1.5 mm long, stigma with a distinct, deeply bifid callus at the

base. Stelidia semi-elliptic, c. 0.5 mm long, tip acute, margins distally slightly

erose. Anther: front margin deeply bifid, papillose, abaxial surface distally

papillose. |

Colours: Rhachis brownish purple. Pedicel and ovary green. Sepals

and petals pale (yellowish) green. Lip dark purplish brown, or ?yellowish.

Figure 37. Bulbophyllum lasioglossum Rolfe ex Ames (147) — a. Habit; b. Flower; c.

Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left:

abaxial side, right: adaxial side; e. Column and lip, lateral view; f. Anther, left: adaxial

side, right: abaxial side; g. Pollinia, above: two pairs, below: single pair. — a, c—g. From

Borden 797; b. From Topping s.n. (all herbarium specimens).

Bulbophyllum Sect. Altisceptrum and Hirtula 99

5 cm

3mm

Se a et

2mm 3

leet »

BoE Sts

MOY

AVG

SURE RETEST TPT anny

meta \\N 4mm

: RS

s SSS

RSS

tify

ours a

PARTON tea

Figure 37. Bulbophyllum lasioglossum Rolfe ex Ames (147)

100 Gard. Bull. Singapore 54 (2002)

Habitat & ecology: Epiphyte in montane forest. Alt. c. 1000 m.

Flowering in May—Jun.

Distribution: PHILIPPINES: Luzon (3 specimens seen).

148. Bulbophyllum negrosianum Ames - Fig. 38, plate 30.

Bulbophyllum negrosianum Ames, Leafl. Philipp. Bot. 5 (1912) 1586. — TYPE:

Philippines, Negros, Negros Oriental, Cuernos Mts., Dumaguete, Elmer 982]

(PNH, holo., destroyed; AMES, NY, iso.).

Rhizome 3—4 mm diam., sections between pseudobulbs 0.8—2 cm

long. Pseudobulbs + touching, ovoid, 1.5—2.5 x 1—1.6 cm. Petiole 2—S

cm. Leaf blade elliptic, 8.5—16 x 3.2—4.6 cm, index (length/width) 2.2—

4.7; subacute. Inflorescence an elongated, rather dense raceme, + patent, c.

42 cm, c. 45-flowered. Peduncle c. 23.5 cm, bracts c. 5, the longest c. 9 mm

long. Rhachis not thickened, c. 18.5 cm, glabrous. Floral bracts triangular,

1.5—2 x 0.8—1 mm, acute. Flowers secund, several open simultaneously,

not fully open, anthesis from the top of the rhachis downwards. Pedicel

and ovary 5.5—6.5 mm long, basal node on a 0.3—0.5 mm-long stump.

Median sepal recurved, elliptic, 5.5—5.8 x 2.2—3 mm, index 1.8—2.7; acute-

acuminate, margins ciliate, base rather narrowly attached; thin, surface

glabrous. Lateral sepals recurved, ovate, 5.5—5.8 x 3—3.5 mm, index 1.5—

2; margins glabrous, base broadly attached; otherwise as the median sepal.

Petals porrect, elliptic to ovate, 3—3.5 x 0.7—1 mm, index 3.5—4.3; subacute,

margins ciliate, base broadly attached; thin, surface glabrous. Lip curved

near the base, obscurely 3-lobed; midlobe elliptic to obovate, c. 4 x 1.5—

2.3 mm, index 1.7—2.7; obtuse, margins with long, thin, straight or slightly

wavy hairs; rather thin with a thick base, surface glabrous; adaxially concave

near the base and with a median slit reaching up to 1/4 of the lip, with two

obtuse ridges diverging proximally towards the distal end of the lateral

lobes, distally fused to a distinct, thick callus with two knobs at about 1/3

the way up of the lip, a third, obtuse median ridge issuing distally from this

callus, bordered by distinct furrows and continuing over c. 3/4 of the lip,

surface otherwise concave; abaxially with a wide, retuse ridge near the

base; lateral lobes attached along the proximal 1/5 of the lip, erect, semi-

circular, margins with hairs, broadly attached, rather thick, adaxially

pubescent towards the margins, abaxially glabrous. Column 1.3—1.6 mm

Figure 38. Bulbophyllum negrosianum Ames (148) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial

side, right: abaxial side; e. Lip, adaxial side; f. Column and lip, lateral view; g. Anther,

above: adaxial side, below: abaxial side; h. Pollinia, above: two pairs, below: single

pair. — a—d, f—h. From Elmer 982] (herbarium specimen); e. From Hort. Vienna 176-

90 (spirit sample).

101

Bulbophyllum Sect. Altisceptrum and Hirtula

RO .

Figure 38. Bulbophyllum negrosianum Ames (148)

102 Gard. Bull. Singapore 54 (2002)

long, stigma with a distinct, deeply bifid callus at the base. Stelidia semi-

elliptic, c. 0.5 mm long, tip obtuse, margins distally erose. Anther: front

margin deeply bifid, papillose, abaxial surface distally papillose.

Colours: Sepals dull purple, the median whitish between the veins in

the centre; or sepals orange yellow with dark red veins proximally, orange

red distally. Petals dark red, with a yellowish midvein; cilia yellow. Lip

yellow, adaxial side reddish brown in the centre, abaxial side with a dark

red patch about half-way, pale yellow proximally. Hairs along lip margins

yellow.

Habitat & ecology: Epiphyte in mossy montane forest. Alt. 1200—

1400 m. Flowering in Apr, May.

Distribution: PHILIPPINES: Luzon (1 specimen seen); Negros (1).

Notes: Flowers rather similar to those of the preceding species. They

differ in being smaller, and in the presence of a distinct callus adaxially on

the lip, where the two lateral keels meet.

The Luzon specimen has wider floral parts than the type from Negros,

but is otherwise not different.

149. Bulbophyllum dracunculus J.J. Verm. — Fig. 39, plate 31.

Bulbophyllum dracunculus J.J. Verm., Gard. Bull. Singapore 52 (2000) 276. — TYPE:

Malaysia, Sabah, NE flank of Gunung Alab, Hort. Leiden 30978 (L, holo.,

SING, iso.).

Bulbophyllum polygaliflorum sensu J.J. Wood, Orch. Borneo 3 (1997) 247 (colour

photograph only), non J.J. Wood.

Rhizome 2.5—2.7 mm diam., sections between pseudobulbs 0.4—1

cm long. Pseudobulbs touching, ovoid, 0.7—1.6 x 0.4—1.4 cm. Petiole 0.4—

1.3 cm. Leaf blade elliptic to ovate, 5.1—8.1 x 1.2—3 cm, index (length/

width) 2.7—5; obtuse to acute. /nflorescence an elongated, lax raceme, +

patent, 15—24 cm, 11—20-flowered. Peduncle 10—18 cm, bracts c. 4, the

longest 4—6 mm long. Rhachis not thickened, 5—7 cm long, glabrous.

Floral bracts triangular, 1.5—2.8 x 2—2.2 mm, slightly acuminate. Flowers

not fully opening, one at a time. Pedicel and ovary 4—4.5 mm long, basal

node + flush with the surface of the rhachis. Median sepal + porrect,

triangular, 4—5 x 1.3—2 mm, index 2.5—3.1; acute to acuminate, margins

ciliate, base rather broadly attached; rather thick, surface glabrous. Lateral

sepals recurved, ovate, 4.5—6.3 x 2.8—4.2 mm, index 1.2—1.6; rounded to

Figure 39. Bulbophyllum dracunculus J.J. Verm. (149) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial

side, right: abaxial side; e. Column and lip, lateral view; f. Anther, left: abaxial side,

right: adaxial side; g. Pollinia, above: two pairs, below: single pair. — All from Hort.

Leiden 30978 (spirit sample).

103

Bulbophyllum Sect. Altisceptrum and Hirtula

Figure 39. Bulbophyllum dracunculus J.J. Verm. (149)

104 Gard. Bull. Singapore 54 (2002)

obtuse, otherwise as the median sepal. Petals porrect, spathulate, 1.6—2.3

x 1.1—1.5 mm, index 1.1—1.9; truncate to retuse, margins ciliate, base

rather broadly attached; rather thick, adaxial surface glabrous but with

two hirsute patches near the top. Lip recurved near the base, oblong, 3.5—

4.2 x 0.8—1.2 mm, index 3.5—4.3; obtuse, margins with slightly club-shaped

hairs increasing in length and thickness towards the tip of the lip; very

thick; adaxially + flat near the base and with a median slit up to 1/3 of the

lip, with 2 rows of irregularly shaped, fleshy calli, those on opposite rows

partly fused c. half-way up the lip, adaxial surface finely papillose; abaxially

with a truncate ridge distally divided in two rows of fleshy calli, surface

glabrous. Column 1.2—1.8 mm long, column foot with two longitudinal,

rounded lamellae. Stelidia triangular, 0.5—0.7 mm long, obtuse, with a

distinct, downwards projecting, strap-shaped to somewhat spathulate tooth

with a 2—3-furcate tip along the lower margin and sometimes with a second,

minute, deltoid, obtuse to acute tooth just in front of this. Anther: front

margin long-papillose, abaxial ridge papillose.

Colours: Peduncle dark purple, rhachis paler. Bracts pale green,

suffused pale lilac. Sepals yellowish or pale green, suffused brownish red

adaxially, margins with white or pink cilia. Petals translucent white, spotted

with purple adaxially, margins with dark purple hairs. Lip creamy white,

yellowish or pinkish, calli blackish purple, hairs purple.

Habitat & ecology: High montane forest, secondary forest, shrubby

vegetation on soils derived from ultrabasic rock. Alt. 800-1800 m. Flowering

in Jan, Apr, Jun—Aug.

Distribution: MALAYSIA: Sabah (4 specimens seen).

150. Bulbophyllum polygaliflorum J.J. Wood — Fig. 40.

Bulbophyllum polygaliflorum J.J. Wood, Kew Bull. 39 (1984) 96; J.J. Wood, Orch.

Borneo 3 (1997) 247 (excl. colour photo, is B. dracunculus J.J. Verm.). —

TYPE: Malaysia, Sarawak, Hansen 499 (C, holo.; K, iso.).

Rhizome c. 6 mm diam., sections between pseudobulbs 1.5—4.5 cm

long. Pseudobulbs distant, ovoid, 2—4.2 x 0.8—1.6 cm. Petiole 1.8—4 cm.

Leaf blade elliptic to ovate, 9—18 x 3.2—4.7 cm, index (length/width)

2.5—4.2; acute. Inflorescence an elongated, rather dense raceme, erect to

patent, 78—86 cm, 30—45-flowered. Peduncle 63—70 cm, bracts 5—9, the

Figure 40. Bulbophyllum polygaliflorum J.J. Wood (150) — a. Habit; b. Flower; c.

Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left:

adaxial side, right: abaxial side; e. Column and lip, lateral view; f. Anther, above:

adaxial side, below: abaxial side; g. Pollinia, left: single pair, right: two pairs. — a

(plant), b—g. From SFN 27956 (herbarium specimen), = Carr 3595 (spirit sample); a

(inflorescence). From Hansen 499 (herbarium specimen).

Bulbophyllum Sect. Altisceptrum and Hirtula 105

hae

Ae RO

Ns ANS

1 _

ANS

MISSES

i S Imm

SENN TON

4 ty

Ar: BR

De. AS

Te. BN

CE seme

3mm

SEAN

—e

—_.

ATT

eet AT

Figure 40. Bulbophyllum polygaliflorum J.J. Wood (150)

106 Gard. Bull. Singapore 54 (2002)

longest 10 mm long. Rhachis not thickened, 15—18 cm long, glabrous.

Floral bracts ovate, 2—3 x 1.5—2 mm, acuminate. Flowers not fully opening,

several simultaneously. Pedicel and ovary 5—6.5 mm long, basal node +

flush with the surface of the rhachis. Median sepal + porrect, triangular,

6—6.4 x 1.5—2.4 mm, index 2.6—4; acute to acuminate, margins ciliolate,

base broadly attached; thick, surface glabrous. Lateral sepals somewhat

recurved, ovate to elliptic, 7.5—9 x 4.8—5 mm, index 1.5—1.8; obtuse,

otherwise as the median sepal. Petals porrect, spathulate with a widened

base, 3—3.5 x c. 2 mm, index 1.5—1.8; rounded, margins ciliate, base rather

broadly attached; rather thick, surface hirsute towards the tip. Lip recurved

near the base, elliptic to ovate, 6—6.8 x 2—2.5 mm, index 2.7—3; obtuse.

margins densely ciliate; very thick; adaxially concave near the base, with a

median slit over most of its length, with 4 rows of irregularly shaped, fleshy

calli, the two inner rows higher and longer than the two outer, adaxial

surface finely and shortly hirsute; abaxially with a rounded ridge near the

base, surface otherwise convex with a median slit, pubescent in the centre

and along the margins. Column 1—1.2 mm long, column foot with two

longitudinal, rounded lamellae. Srelidia triangular, c. 0.5 mm long, acute,

with a large, antrorse, deltoid, acute tooth along the upper margin, and a

similar but much smaller, slightly falcate, triangular tooth along the lower.

Anther: front margin ciliate, abaxial ridge papillose.

Colours: Sepals greenish, marked with pale purple, or purple, cilia

white. Petals translucent near the base, blackish purple near the tip, cilia

yellow. Lip greenish, upper surface with blackish green warts and yellow

indument, sides with yellow cilia. Column whitish, spotted purple.

Habitat & ecology: Epiphyte in montane forest. Alt. 1700—1800 m.

Distribution: MALAYSIA: Sarawak (1 specimen seen). Sabah (2).

151. Bulbophyllum ochthodes J.J. Verm., sp. nov. — Fig. 41, plate 32.

Bulbophyllum ochthodes J.J. Verm., a Bulbophyllo polygalifloro labelli latere adaxiali

tantum callis in seriebus duabus differt. - TYPE: Malaysia, further details

unknown, Hort. Singapore SBG-O 616 (SING, holo.; K, KLU, iso.).

Rhizome 4—7 mm diam., sections between pseudobulbs 2—6 cm

long. Pseudobulbs distant, ovoid, 1.8—3.5 x 0.7—1.5 cm. Petiole 1—3.5 cm.

Leaf blade elliptic, 7.5—14 x 2.7—4.5 cm, index (length/width) 2.6—4; acute.

Inflorescence an elongated, rather dense raceme, + erect, 26—58 cm, 40—

Figure 41. Bulbophyllum ochthodes J.J. Verm. (151) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial

side, right: abaxial side; e. Column and lip, lateral view; f. Anther, left: adaxial side,

right: abaxial side; g. Pollinia, left: single pair, right: two pairs. — All from Hort. Singapore

SBG-O 616 (spirit sample).

Bulbophyllum Sect. Altisceptrum and Hirtula 107

3mm

Imm

Figure 41. Bulbophyllum ochthodes J.J. Verm. (151)

108 Gard. Bull. Singapore 54 (2002)

60-flowered. Peduncle 19—46 cm, bracts c. 6, the longest 4.8—5.5 mm

long. Rhachis not thickened, 6.5—12 cm long, glabrous. Floral bracts

triangular, 1.8—2 x c. 1 mm, acute. Flowers not fully opening, few

simultaneously. Pedicel and ovary 2.8,—4 mm long, basal node on a 0.2—

0.5 mm-long stump. Median sepal + porrect, triangular, 3.8—4 x 1.5 mm,

index 2.5—2.7; acuminate, margins ciliolate, base rather narrowly attached;

thick, surface glabrous. Lateral sepals somewhat recurved, ovate, 4.5—5 x

2.8—3 mm, index c. 1.6; subacute, otherwise as the median sepal. Petals

porrect, spathulate with a widened base, c. 2 x 1—1.1 mm, index c. 1.8;

rounded, margins ciliate, base rather broadly attached; rather thin, surface

hirsute towards the tip. Lip abruptly recurved near the base, elliptic-oblong,

3—3.1 x 1.2—1.5 mm, index 2—2.6; rounded, margins densely ciliate with

the hairs thicker and = stiff towards the tip; very thick; adaxially concave

near the base, with a median slit over most of its length, with 2 rows of

irregularly shaped, fleshy, finely and shortly hirsute calli, adaxial surface

otherwise + glabrous; abaxially with a rounded ridge near the base, surface

otherwise convex with scattered, slightly stiff, thick hairs more or less

arranged in two rows, surface otherwise glabrous. Column 1.2—1.3 mm

long, stigma with an inconspicuous, slightly cleft basal callus, column foot

swollen. Stelidia deltoid, c. 0.4 mm long, obtuse, with a small, deltoid,

obtuse teeth along the lower margin. Anther: front margin ciliate, abaxial

ridge papillose.

Colours: Peduncle dark red. Sepals reddish purple, pale yellowish

green near the base, cilia along the margins white. Petals whitish near the

base, blackish purple towards the tip, hairs blackish purple. Lip blackish

purple except for a pale yellowish patch near the base abaxially, and the

yellowish furrows between the warts adaxially. Column yellowish.

Habitat & ecology: Montane forest. Alt. c. 1500 m.

Distribution: MALAYSIA: Peninsula (1 specimen seen).

Notes: Most similar to Bulbophyllum polygaliflorum, but differs in

the flowers having a shorter lip with only two rows of calli adaxially.

Only a single specimen is known, from a nursery in Cameron

Highlands, Malaysia, which was said to have been collected locally.

Figure 42. Bulbophyllum phreatiopse J.J. Verm. (152) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, above: adaxial

side, below: abaxial side; e. Column and lip, lateral view; f. Anther, left: adaxial side,

right: abaxial side; g. Pollinia, above: single pair, below: two pairs. — All from Hoogland

& Craven 10899 (herbarium specimen).

Bulbophyllum Sect. Altisceptrum and Hirtula 109

---

2mm

—_——=—=-- = <a oy

~-- = =

-—---—-

LOGY,

See?

lle

Rossi sieht,

= :

Figure 42. Bulbophyllum phreatiopse J.J. Verm. (152)

110 Gard. Bull. Singapore 54 (2002)

152. Bulbophyllum phreatiopse J.J. Verm. — Fig. 42.

Bulbophyllum phreatiopse J.J. Verm., Blumea 38 (1993) 151. - TYPE: Papua New

Guinea, East Sepik District, Mt. Hunstein, Hoogland & Craven 10899 (L, holo..

AMES, LAE iso.).

Rhizome 2.5—5 mm diam., sections between pseudobulbs 4—19 cm

long. Pseudobulbs widely distant, ovoid, 1.2—1.8 x 0.8—1.2 cm. Petiole 3—

5.5 cm. Leaf blade elliptic, 10—13.5 x 2—3.3 cm, index (length/width)

4.5—5.5; obtuse. Inflorescence an elongated, dense raceme, erect to patent,

27—35 cm, 40—55-flowered. Peduncle 18—26 cm, bracts 8—11, the longest

15—25 mm long. Rhachis not thickened, 7—13 cm long, glabrous. Floral

bracts ovate, 2.5—5 x c. 2 mm, acute. Flowers secund, not fully opening,

many simultaneously. Pedicel and ovary 1.8—3 mm long, basal node c.

flush with the surface of the rhachis. Median sepal porrect, ovate to elliptic,

c. 4.6—5.2 x 1.8—2 mm, index 2.5—3; obtuse to subacute, margins and

adaxial surface finely papillose distally, base broadly attached; rather thin.

Lateral sepals recurved, ovate, 5—5.5 x 2.5—3 mm, index 1.8—2, acute;

otherwise as the median sepal. Petals porrect, ovate, 3—3.8 x 1.2—1.5 mm,

index c. 2.5; obtuse, margins and adaxial surface papillose distally, base

rather narrowly attached; rather thin. Lip recurved in the proximal half,

ovate, 3—3.5 x 1—1.5 mm, index 2.3—3; rounded, thick, glabrous; adaxially

concave near the base, and with a median ridge with a tooth on its crest

and flanked by furrows, this median ridge proximally widening to a

triangular callus forming a roof over the flanking furrows, in front of the

median ridge two more ridges converging distally, adaxial surface distally

convex; abaxially with a wide, retuse ridge towards the base. Column 1.5—

1.8 mm long. Stelidia triangular, 0.6—0.9 mm long, tip acute; with or without

a small, deltoid, obtuse tooth along the upper margin. Anther: front margin

and abaxial ridge papillose.

Colours: Sepals and petals white. Lip white, or orange-yellow.

Habitat & ecology: Epiphyte in primary forest. Alt. 1300-1500 m.

Flowering in Oct—Nov.

Distribution: PAPUA NEW GUINEA: East Sepik Dist., Southern

Highlands District (2 specimens seen).

Notes: Bulbophyllum phreatiopse is tentatively included in sect.

Hirtula, see the note below the description of the section.

Figure 43. Bulbophyllum parviflorum Par. & Rchb.f. in Rchb.f. (153) — a. Habit; b.

Flower; c. Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; d.

Lip, left: adaxial side, right: abaxial side; e. Lip, adaxial side; f. Column and lip, lateral

view; g. Anther, left: adaxial side, right: abaxial side; h. Pollinia, above: single pair,

below: two pairs. — a. From Pantling 245 (herbarium specimen); b—d, f—h. From

Hort. Leiden 960104 (spirit sample); e. From GT 8549 (spirit sample).

Bulbophyllum Sect. Altisceptrum and Hirtula 111

NSN =

SS SNS ran .

—a\bsi l

7 eds, en..

Imm

-——————

2mm

Scm

SST FLEE XQ

ge SS SS i) \ : IMSS

VE a IIIS Ret f/ ™

Figure 43. Bulbophyllum parviflorum Par. & Rchb.f. in Rchb.f. (153)

112 Gard. Bull. Singapore 54 (2002)

The original description of B. phreatiopse erroneously stated that the

petals are 3-veined.

153. Bulbophyllum parviflorum Par. & Rchb.f. in Rchb.f. — Fig. 43, plate 33.

Bulbophyllum parviflorum Par. & Rchb.f. in Rchb.f., Trans. Linn. Soc. London, Bot.

30 (1874) 152. — Phyllorchis parviflora (Par. & Rchb.f. in Rchb.f.) O. Kuntze,

Rev. Gen. Pl. 2 (1891) 677. — TYPE: Myanmar, Tenasserim, Parish 305 (K,

holo.; C, W, iso.).

Bulbophyllum thomsoni Hook.f., Fl. Brit. Ind. 5 (1890) 764; id., Hooker’s Icon. Pl. 21

(1892) t. 2041; King & Pantling, Ann. Roy. Bot. Gard. (Calcutta) 8 (1898) 83, t.

116. — Phyllorchis thomsonii (Par. & Rchb.f. in Rchb.f.) O. Kuntze, Rev. Gen.

Pl. 2 (1891) 678. — TYPE: Sikkim, Thomson s.n. (1850) (K, holo.).

Rhizome 2.8—5 mm diam., sections between pseudobulbs 2—10 cm

long. Pseudobulbs widely distant, depressed conical, 0.8—2 x 1.2—1.8 cm.

Petiole 1—3 cm. Leaf blade ovate, 7.2—16 x 1.2—2.4 cm, index (length/

width) 5.4—7.9; obtuse to acute. Inflorescence an elongated, dense raceme,

erect to patent, 15—30 cm, 35—80-flowered. Peduncle 6—12 cm, bracts c.

6, the longest 9—14 mm long. Rhachis not thickened, 9.5—19.5 cm long,

glabrous. Floral bracts ovate to triangular, 1.3—4 x 1—2 mm, acuminate.

Flowers sometimes secund, not fully opening, many simultaneously. Pedicel

and ovary 2.2—4 mm long, basal node + flush with the surface of the

rhachis or on a c. 0.2 mm-long stump. Median sepal porrect, triangular,

2.8—6 x 0.8—1.7 mm, index 2.4—3.8; acute, margins ciliolate, base rather

broadly attached; rather thin, surface very finely papillose locally. Lateral

sepals slightly recurved, 2.8—6.5 x 1.1—2.5 mm, index 1.5—3.2, upper

margin glabrous to ciliolate, lower margin ciliolate to ciliate; otherwise as

the median sepal. Petals porrect, (ob-)ovate to elliptic, 1.5—2.4 x 0.6—1

mm, index 1.8—3; obtuse to acuminate, margins ciliate, base rather broadly

attached; thin, glabrous. Lip recurved in the proximal half, oblong to

subspathulate, 1.7—2.5 x 0.8—1l1 mm, index 1.8—2.5; rounded, margins

ciliate except near the tip; thick, surface finely papillose, or only so distally;

adaxially concave near the base, without a basal callus, with 2—3 ridges,

the lateral distinct, rather sharp, diverging proximally and continuing over

c. 2/3 of the lip, the median ridge, if present, low, inconspicuous, rounded

and continuing over 1/2 of the lip; abaxially with a wide, retuse ridge

towards the base; basal auricles inconspicuous. Column 1—1.3 mm long.

Stelidia triangular, 0.4—0.7 mm long, tip obtuse; with a deltoid, rounded to

obtuse tooth along the upper margin, and often an inconspicuous, rounded

wing along the lower margin, at the base. Anther: front margin glabrous,

abaxial surface somewhat finely papillose.

Colours: Sepals and petals creamy white, the petals sometimes with

red cilia. Lip white, yellow or green.

Bulbophyllum Sect. Altisceptrum and Hirtula 113

Habitat & ecology: Tropical forest. Alt. 400-1500 m. Flowering in

Jul—Oct.

Distribution. BHUTAN (1). INDIA: Sikkim (3 specimens seen).

MYANMAR: Moulmein (1). THAILAND (2).

Notes: The flower colour is partly copied from King & Pantling (1898).

On average, the Myanmar and Thailand specimens (including the

type of Bulbophyllum parviflorum) are smaller in all parts than those from

the eastern Himalayas (including the type of B. thomsoni). As far as can

be ascertained from herbarium material, they also have a more distinct

median ridge on the adaxial side of the lip. Distinction of subspecies,

however, is impossible with the material now available.

154. Bulbophyllum secundum Hook.f. — Fig. 44, plate 34, 35.

Bulbophyllum secundum Hook..f., Fl. Brit. Ind. 5 (1890) 764; id., Hooker’s Icon. PI. 21

(1892) t. 2042; Seidenf., Dansk Bot. Ark. 33, 3 (1979) 103 (partly, excl. syn. B.

nigripetalum [is B. nigripetalum]). — Phyllorchis secunda (Hook.f.) O. Kuntze,

Rev. Gen. Pl. 2 (1891) 678. — TYPE: India, Naga Hills, Prain 41 (K, holo.).

Bulbophyllum subparviflorum Z.H. Tsi & S.C. Chen, Acta Phytotax. Sin. 32 (1994)

555. — TYPE: China, Yunnan, Menghai, 7’si 9/-544 (PE, holo.; K, iso.)

Rhizome (0.8—1.2 mm diam., sections between pseudobulbs 0.5—1.8

cm long. Pseudobulbs close, depressed conical to lenticular, 0.4—1 x 0.6—

1.5 cm. Petiole 0.7—1.5 cm. Leaf blade elliptic, 3.2—10 x 0.4—1 cm, index

(length/width) 5—12.5; obtuse to acute. /nflorescence an elongated, rather

lax raceme, erect, 7.5—22 cm, 8—23-flowered. Peduncle 5—13.2 cm, bracts

c. 4, the longest 2—4.5 mm long. Rhachis not thickened, 2.2—11 cm long,

glabrous. Floral bracts triangular, 1—1.8 x 0.8—1.2 mm, acute to acuminate.

Flowers distichous or scattered, resupinate, secund, not fully opening, several

simultaneously. Pedicel and ovary 1.8—6 mm long, basal node + flush with

the surface of the rhachis or on ac. 0.3 mm-long stump. Median sepal +

porrect, ovate to triangular, 2—3.2 x 0.9—1.3 mm, index 2.1—2.5; obtuse

to acute, margins ciliolate, base rather narrowly attached; rather thin, surface

glabrous. Lateral sepals recurved, triangular, 2.2—3.8 x 1.3—2.2 mm, index

1.4—1.9; (sub-)acute, upper margin glabrous to sparsely ciliolate; otherwise

as the median sepal. Petals porrect, obovate to spathulate, 1.5—2.2 x 0.6—

0.8 mm, index 2.2—3; rounded to obtuse, margins ciliate, base broadly

attached; thin, glabrous or slightly papillose towards the tip. Lip recurved

in the proximal half, ovate, 1.8—3 x 0.9—1.2 mm, index 1.8—2.8; rounded,

margins ciliate except near the tip; thick; adaxially slightly concave near

the base and with a deeply cleft, triangular basal callus, with 2 very weak,

rounded ridges diverging proximally and fused to a single slightly convex

body in the distal half of the lip, surface finely papillose to shortly pubescent

proximally and glabrous distally; abaxially with an inconspicuous, retuse

114 Gard. Bull. Singapore 54 (2002)

ridge towards the base, surface glabrous. Column 1.1—1.3 mm long. Stelidia

triangular, 0.3—0.5 mm long, tip acute; with or without an often antrorse,

deltoid to triangular, obtuse to acute tooth along the upper margin, with or

without a deltoid, rounded to obtuse wing along the lower margin, at the

base. Anther: front margin papillose, abaxial surface papillose distally.

Colours: Sepals pale green, often suffused with dull reddish brown

or purple towards the tip, or almost entirely of that colour except for the

base. Petals pale green, similarly suffused with reddish brown or purple:

hairs purple. Lip reddish purple, or brownish green, proximally dark brown

along the margins. Column greenish, often dark brownish or purplish at

the foot. Flowers also reported to be light blue (unlikely, see Samuel 13549)

or pink (Grey-Wilson & Phillips 93).

Habitat & ecology: Epiphyte in montane temperate or tropical

evergreen (shrub) forests; also reported from depleted evergreen forest.

Alt. 1200-2500 m. Flowering in Jan, Feb, May—Jul, Sep.

Distribution: CHINA: Yunnan (1 specimen seen). NEPAL (2).

INDIA: Sikkim (1); Assam (1); Naga Hills (1). MYANMAR: (1): northern

part (2). THAILAND (4). VIETNAM: southern part (1).

Notes: Seidenfaden (1979) has confused this with Bulbophyllum

nigripetalum; see the note under the latter.

155. Bulbophyllum debrincatiae J.J. Verm., sp. nov. — Fig. 45, plate 36.

Bulbophyllum debrincatiae J.J. Verm., a Bulbophyllo piloso in calcari brevi e sepalorum

lateralium basi protrudenti, labelli parte apicali pilis longis differt. - TYPE:

Philippines, Luzon, Nueva Viscaya, Debrincat, B. cult. (leg. Cootes, J.E.) 001

(SING, holo.).

Rhizome 3—4 mm diam., sections between pseudobulbs 1.2—1.6 cm

long. Pseudobulbs close, ovoid, 1.3—2.5 x 1—1.6 cm. Petiole 2—3.5 cm.

Leaf blade ovate-elliptic, 9.2—14.5 x 3.3—5 cm, index (length/width) 2.7—

2.9: rounded. Inflorescence an elongated, rather dense raceme, erect, c. 43

cm, c. 34-flowered. Peduncle c. 24.5 cm, bracts c. 5 (4 present on portion of

inflorescence available), the longest c. 10 mm long. Rhachis slightly

thickened, c. 15 x 0.3 cm, finely papillose. Floral bracts triangular, c. 2.2 x

1.2 mm, acute to acuminate. Flowers secund, not fully opening, several

simultaneously, anthesis from the top of the rhachis downwards. Pedicel

and ovary c. 5.5 mm long, finely papillose distally, basal node + flush with

Figure 44. Bulbophyllum secundum Hook.f. (154) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial

side, right: abaxial side; e. Column and lip, lateral view; f. Anther, above: abaxial side,

below: adaxial side; g. Pollinia, above: single pair, below: two pairs. — a. From Jongejan

s.n. (living plant); b—g. From Jongejan cult. 889 (spirit sample).

Bulbophyllum Sect. Altisceptrum and Hirtula 115

ee og age ’

—<S is

: PP

lmm ~ iD

2mm

U -

JIcm

y) i

I\ 6

RY N

me Of

af\

2mm

Figure 44. Bulbophyllum secundum Hook.f. (154)

116 Gard. Bull. Singapore 54 (2002)

the surface of the rhachis. Median sepal porrect to recurved, ovate-

triangular, c. 7.5 x 2.2 mm, index c. 3.4; acute, margins ciliate, base rather

broadly attached; rather thick, surface glabrous adaxially, finely papillose

abaxially. Lateral sepals recurved, triangular, c. 7.5 x 3.6 mm, index c.2.1,

upper margin ciliolate with scattered longer hairs in between; otherwise as

the median sepal. Petals porrect, triangular-ovate, c. 3.4 x 1.4 mm, index

2.4—2.5; subacute, margins long ciliate, hairs longer towards the tip, base

rather broadly attached; rather thin, surface glabrous. Lip curved close to

the base, oblong, c. 7.5 x 2 mm, index 3.7—3.8; rounded, margins ciliolate

proximally, long ciliate distally, glabrous in between; thick; adaxially slightly

concave near the base, without a basal callus, with a median furrow reaching

up to c. 1/4 of the lip, slightly convex elsewhere, surface glabrous; abaxially

with a rounded ridge near the base, surface slightly convex, glabrous but

with a finely pubescent patch in front of the basal ridge. Column c. 1.8 mm

long, foot slightly swollen, abaxial side with a short, obtuse spur near the

tip. Stelidia c. 0.8 mm long, acute, obliquely. rectangular in outline with a

distinct, antrorse, triangular, acute tooth along the upper margin, and a

similar, but obtuse tooth along the lower. Anther: front margin pilose,

abaxial surface finely papillose.

Colours: Rhachis yellowish, slightly suffused with red. Bract, pedicel

and ovary yellowish green, marked brownish red distally. Sepals yellowish

green, marked with dark red (appearing almost black) along veins and

margins; hairs around edge greyish. Petals similarly coloured, hairs blackish.

Lip shiny black, hairs greyish. Column green with black blotches.

Habitat & ecology: Epiphyte in montane forest. Alt. c. 1200 m.

Flowering in cultivation in Oct.

Distribution: PHILIPPINES: Luzon, Nueva Viscaya (1 specimen

seen).

Notes: Bulbophyllum debrincatiae is most similar to B. pilosum, but

differs in having triangular-ovate petals, glabrous margins along the middle

section of the lip, and a remarkable, small, obtuse papillose spur protruding

from between the base of the lateral sepals.

Named in honour of Mrs Bev Debrincat, who acquired the plant, as

part of Mr Jim Cootes’s orchid collection from the Philippines in 1998, and

who was the first to flower it in cultivation.

Figure 45. Bulbophyllum debrincatiae J.J. Verm. (155) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial

side, right: abaxial side; e. Column and lip, lateral view; f. Anther, above: adaxial side,

below: abaxial side; g. Pollinia, left: single pair; right: two pairs,. — All from Debrincat,

B. cult. (leg. Cootes, J.E.) 001 (Photographs, notes from J. Cootes, spirit sample).

Bulbophyllum Sect. Altisceptrum and Hirtula 117

Imm

ANS

3mm

STII INOIIN 2s tore

eas

ers

Figure 45. Bulbophyllum debrincatiae J.J. Verm. (155)

118 Gard. Bull. Singapore 54 (2002)

156. Bulbophyllum pilosum J.J. Verm., sp. nov. — Fig. 46.

Bulbophyllum pilosum J.J. Verm., a Bulbophyllo lasioglosso in sepalis lateralibus ciliatis,

stigmatis basi sine callo bifido differt. - TYPE: Malaysia, Sabah, Ulu Padas,

Vermeulen 562 (L, holo.).

Rhizome 1.5—2 mm diam., sections between pseudobulbs 0.4—0.7

cm long. Pseudobulbs touching, ovoid, 1.4—2.5 x 0.6—0.9 cm. Petiole 2—

3.2 cm. Leaf blade elliptic to ovate, 11—13 x 2—2.5 cm, index (length/

width) 5.2—5.8; acute. Inflorescence an elongated, rather dense raceme, c.

19 cm, c. 10-flowered. Peduncle + patent, c. 17 cm, bracts c. 5, the longest

c. 4 mm long. Rhachis not thickened, nodding, c. 2 cm long, glabrous.

Floral bracts triangular, c. 1.6 x 1 mm, acuminate. Pedicel and ovary c. 6

mm long, basal node on a c. 0.2 mm-long stump. Median sepal porrect to

?recurved, triangular, c. 5.8 x 2.1 mm, index 2.3—2.4; acute-acuminate,

margins with long hairs, base rather broadly attached; rather thin, surface

glabrous. Lateral sepals ?recurved, ovate, c. 7.3 x 3.1 mm, index 2.3—2.4;

subacute, otherwise as the median sepal. Petals porrect, obovate, c. 3.3 x

1.8 mm, index 1.8—1.9; rounded, margins long ciliate, hairs longer towards

the tip, base rather broadly attached; rather thick, surface glabrous. Lip

curved in the proximal half, obscurely 3-lobed; midlobe ovate, c. 4.2 x 1.8

mm, index 2.3—2.4; rounded, margins with long, thin, wavy hairs; rather

thin with a thick base, surface glabrous; adaxially concave near the base,

without a basal callus, with a median slit reaching up to 1/2 of the lip, with

two obtuse ridges diverging proximally towards the distal end of the lateral

lobes, distally thick and together forming a small callus continuing over c.

1/4 of the lip, surface otherwise + flat; abaxially with a wide, retuse ridge

near the base; /ateral lobes attached along the proximal 1/4—1/5 of the lip,

erect, semi-circular, margins ciliate, broadly attached, rather thin, surface

glabrous. Column c. 1.5 mm long, foot slightly swollen. Stelidia c. 1 mm

long, acute, obliquely rectangular in outline with a distinct, antrorse,

triangular, acute tooth along the upper and lower margin.

Colours: Perianth.on top of a fruit purplish.

Habitat & ecology: Understorey epiphyte in tall forest on leached

sandstone soil. Alt. 1300-1500 m. Fruiting in Oct.

Distribution: MALAYSIA. Sabah, Ulu Padas (1 specimen seen).

Notes: The floral details have been reconstructed from the perianth

persistent on an almost mature fruit. Some details may be inaccurate.

Most similar to Bulbophyllum lasioglossum; the latter differs in having

Figure 46. Bulbophyllum pilosum J.J. Verm. (156) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial

side, right: abaxial side; e. Column and lip, lateral view. — All from Vermeulen 562

(herbarium specimen).

Bulbophyllum Sect. Altisceptrum and Hirtula 119

4mm

2mm

3mm

Figure 46. Bulbophyllum pilosum J.J. Verm. (156)

120 Gard. Bull. Singapore 54 (2002)

lateral sepals with glabrous margins, much narrower, acute petals, and a

deeply bifid callus at the base of the stigma.

The name refers to the pilose sepals.

157. Bulbophyllum xenosum J.J. Verm. — Fig. 47, plate 37.

Bulbophyllum xenosum J.J. Verm., Blumea 41 (1996) 353. - TYPE: Brunei, Temburong,

upper zone of Bukit Belalong, Jongejan cult. 4425 (L, holo.).

Rhizome 2.5—3 mm diam., sections between pseudobulbs 1.5—4.5

cm long. Pseudobulbs distant, depressed-conical to lenticular, 1.2—1.7 x

1—1.4 cm. Petiole 1.2—2 cm. Leaf blade elliptic, 8.5—13 x 0.8—1.2 cm,

index (length/width) 10—11; subacute. Inflorescence an elongated, lax

raceme, c. 44 cm, c. 30-flowered. Peduncle patent, c. 19 cm, bracts c. 4, the

longest 5 mm long. Rhachis not thickened, arching, c. 25 cm long, glabrous.

Floral bracts triangular, 2—2.5 x c. 1 mm, acute. Flowers imperfectly

distichous but scattered at the base and the tip of the rhachis, secund,

many open simultaneously, fully open. Pedicel and ovary 4.5—5 mm long,

basal node on a c. 0.3 mm-long stump. Median sepal porrect to recurved,

ovate, c. 11 x 1.5 mm, index 7—8; acute, margins ciliate, base rather broadly

attached; rather thin, surface glabrous. Lateral sepals recurved, triangular,

9.5—10 x c. 2.2 mm, index 4.3—4.6; upper margin glabrous, base broadly

attached; otherwise as the median sepal. Petals porrect, obovate, c. 1.5 x

0.8 mm, index 1.8—1.9; acute, margins long ciliate towards the tip, base

broadly attached; thin, surface glabrous. Lip curved near the base, ovate-

triangular, c. 5.5 x 1.5 mm, index 2.6—2.7; obtuse, margins slightly erose,

with long, straight, rather thin hairs increasing in length and thickness

towards the tip of the lip; lip rather thin, base thick; adaxially slightly

concave near the base, without a basal callus, with a median slit continuing

over about 1/3 of the lip, surface otherwise flat or somewhat convex,

glabrous; abaxially with a retuse ridge near the base, surface otherwise flat

or somewhat convex, glabrous except for two longitudinal rows of hairs

about half-way up the lip. Column c. 2.2 mm long. Stelidia falcate, triangular,

c. 0.9 mm long, tip acute; with an inconspicuous, deltoid, acute tooth along

the upper margin, and an inconspicuous, deltoid, broadly rounded wing

along the lower margin. Anther: front margin long ciliate, abaxial ridge

with hairs.

Colours: Rhachis, pedicel and ovary suffused with reddish purple.

Figure 47. Bulbophyllum xenosum J.J. Verm. (157) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: abaxial

side, right: adaxial side; e. Column and lip, lateral view; f. Anther, above: abaxial side,

below: adaxial side; g. Pollinia, left: two pairs, right: single pair. — All from Jongejan

cult. 4425 (spirit sample).

Bulbophyllum Sect. Altisceptrum and Hirtula 121

4mm

3mm

Imm

2mm

Scm

Figure 47. Bulbophyllum xenosum J.J. Verm. (157)

122 Gard. Bull. Singapore 54 (2002)

Sepals greenish white, abaxially suffused with pale purple, adaxially heavily

stained with purple and with large, blackish purple blotches. Lip yellowish

white, suffused with reddish purple towards the margins, and with large,

more or less transverse blackish purple blotches; hairs reddish at the base,

yellow towards the tip.

Habitat & ecology: Primary forest. Alt. 650—900 m.

Distribution: BRUNEI (1 specimen seen).

158. Bulbophyllum limbatum Lindl. — Fig. 48, plate 38.

Bulbophyllum limbatum Lindl., Edward’s Bot. Reg. 26, misc. (1840) 74

(“Bolbophyllum”); J.J. Smith, Repert. Spec. Nov. Regni Veg. 32 (1932) 305;

Seidenfaden, Dansk Bot. Ark. 33, 3 (1979) 100. — Phyllorchis limbata (Lindl.)

O. Kuntze, Rev. Gen. Pl. 2 (1891) 677. - TYPE: Singapore, Loddiges (K, holo.).

Bulbophyllum blepharosepalum Schltr, Bull. Herb. Boiss. s. 2, 6 (1906) 462. - TYPE:

Indonesia, Sumatra, Indragiri, Sungei Lalah, Schlechter 1324] (B, holo..,

destroyed).

[Not Bulbophyllum limbatum sensu Par. & Rchb.f. in Rchb.f., Trans. Linn. Soc. London,

Bot. 30 (1874) 152 (= B. rufilabrum Par. ex Hook.f., sect. Careyana, fide

Seidenfaden 1979).]

Rhizome 2—4 mm diam., sections between pseudobulbs 2—26 cm

long. Pseudobulbs (widely) distant, ovoid to depressed-conical or lenticular,

1—2.5 x 1.2—2.5 cm. Petiole 1—4 cm. Leaf blade ovate, 7.5—16 x 2—3.7

cm, index (length/width) 3.1—5.3; acute. /nflorescence an elongated, rather

dense raceme, erect to patent, 6.5—26 cm, 7—22-flowered. Peduncle 5—

16.5 cm, bracts 5—6, the longest 4.5—6 mm long. Rhachis not thickened,

1.4—10 cm long, glabrous. Floral bracts triangular, not clasping the rhachis,

1.2-1.5 mm long, acute. Flowers scattered, secund, not opening fully, many

simultaneously. Pedicel and ovary 45—6.2 mm long, basal node on a 0.2—

0.4 mm-long stump. Median sepal porrect, ovate, 4—6.5 x 1.8—3.4 mm,

index 1.5—2.6; obtuse to acute, margins ciliate, base broadly attached;

rather thin, surface glabrous or finely papillose locally. Lateral sepals

recurved, ovate-triangular, 4.8—7.5 x 2.5—5 mm, index 1.1—2.2:; obtuse to

acuminate, upper margin slightly papillose; otherwise as the median sepal.

Petals porrect, elliptic to spathulate, 2.4—3.8 x 1—1.9 mm, index 1.7—2.8:

rounded to obtuse, margins ciliate distally, base broadly attached; rather

thin, surface finely papillose to pubescent distally, otherwise glabrous. Lip

curved near the base, (ob-)ovate to elliptic, 3.5—6 x 1.2—3 mm, index

Figure 48. Bulbophyllum limbatum Lindl. (158) — a. Habit; b. Flower; c. Flower analysis,

from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: abaxial side, right:

adaxial side; e. Column and lip, lateral view; f. Anther, above: adaxial side, below:

abaxial side; g. Pollinia, left: two pairs, right: single pair. — a. From Jongejan cult. 3995

(spirit sample); b—g. From Hort. Leiden 960104a (spirit sample).

Bulbophyllum Sect. Altisceptrum and Hirtula 123

4mm

lmm

3mm

Figure 48. Bulbophyllum limbatum Lindl. (158)

124 Gard. Bull. Singapore 54 (2002)

1.6—3; rounded to obtuse, margins papillose distally; thick; adaxially slightly

concave near the base, without a basal callus, with two inconspicuous,

rounded, sometimes finely pubescent, proximally diverging ridges in the

proximal 1/3 of the lip, with a median strip widening c. half-way up the lip

into a vaguely outlined, large, somewhat convex, ovate, glabrous or papillose

patch, with the surface otherwise glabrous proximally and often finely

papillose or hirsute distally; abaxially with a + truncate ridge near the

base, surface otherwise slightly concave, glabrous. Column 1.7—2.3 mm

long, Stelidia triangular, 0.4—0.8 mm long; acute. Anther: front margin

erose and fimbriate, abaxial ridge papillose.

Colours: Sepals yellowish to greenish, with reddish-purple veins and

margin, proximally often with a few purplish blotches; cilia white. Petals

yellowish, with a reddish purple vein and margins; proximally often with a

purplish blotch. Lip bright yellow, or reddish purple, adaxially with a

yellowish or whitish median line, abaxially with a yellowish to white base.

Habitat & ecology: Rainforest; also in low and open kerangas forest

and similar forest on ultrabasic soil. Alt.: sea level to 1000 m. Flowering in

Jan, Apr, Aug, Oct, Nov.

Distribution: THAILAND (fide Seidenfaden). MALAYSIA:

Peninsula (6); Sarawak (4); Sabah (4). SINGAPORE (1). BRUNEI (1).

INDONESIA: Sumatra (fide Schlechter); Riau Archipelago, Bangka (fide

J.J. Smith); Kalimantan (1).

Note: The flowers are almost identical to those of the next species.

The differences between the two are mainly in the vegetative parts, and

the mode of anthesis.

159. Bulbophyllum rariflorum J.J. Sm. — Fig. 49.

Bulbophyllum rariflorum J.J. Sm., Bull. Dep. Agric. Indes Neerl. 15 (1908) 20. — TYPE:

Indonesia, Kalimantan, Pontianak, Hort. Bogor s.n. (BO, holo., not seen).

Rhizome 1.7—2.7 mm diam., sections between pseudobulbs 0.8—9.5

cm long. Pseudobulbs close to widely distant, depressed-conical to lenticular,

0.7—1.2 x 1.2—1.8 cm. Petiole 1—2.6 cm. Leaf blade elliptic to ovate, 7—

14 x 0.8—1.6 cm, index (length/width) 6.5—10; acute. Inflorescence an

elongated, very lax raceme, patent, 11—41 cm, 1—9-flowered. Peduncle

10.6—32.5 cm, bracts 3—4, the longest 3—5 mm long. Rhachis not

thickened, slightly arching, 0.4—9.5 cm long, glabrous. Floral bracts tubular,

Figure 49. Bulbophyllum rariflorum J.J.sm. (159) — a. Habit; b. Flower; c. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial

side, right: abaxial side; e. Column and lip, lateral view; f. Anther, above: adaxial side,

below: abaxial side; g. Pollinia, above: two pairs, below: single pair. — All from Jongejan

cult. 2214 (spirit sample).

Bulbophyllum Sect. Altisceptrum and Hirtula 125

3mm

Figure 49. Bulbophyllum rariflorum J.J.Sm. (159)

126 Gard. Bull. Singapore 54 (2002)

clasping the rhachis, 1.8—2.9 mm long, acute to acuminate. Flowers

imperfectly distichous or scattered, not opening fully, each one developing

only after anthesis of the previous. Pedicel and ovary 5—5.6 mm long,

basal node on a c. 0.4—0.7 mm-long stump. Median sepal porrect, ovate,

5—6.3 x 2—2.8 mm, index 1.9—2.5; rounded to obtuse, margins ciliate,

base rather broadly attached; rather thin, adaxial surface finely papillose

distally, otherwise glabrous. Lateral sepals recurved, ovate-triangular, 5.3—

7.2 x 2.8—4 mm, index 1.4—2.2; obtuse to acute, upper margin slightly

papillose, base broadly attached; otherwise as the median sepal. Petals

porrect, spathulate, 3—3.2 x 1.5—2 mm, index 1.5—2; rounded, margins

ciliate distally, base rather broadly attached; thin, adaxial surface finely

papillose distally, otherwise glabrous. Lip curved near the base, obovate to

elliptic, 5.2—6.8 x 2.5—3.3 mm, index 2—2.3; rounded, margins papillose-

ciliolate distally; thick; adaxially slightly concave near the base, without a

basal callus, with a median slit continuing over about 1/4 of the lip, with

two inconspicuous, rounded, proximally diverging ridges in the proximal 1/3

of the lip, with a median strip widening c. half-way up the lip into a vaguely

outlined, large, somewhat convex, ovate, glabrous or papillose patch, with

the surface otherwise glabrous proximally and often finely papillose or

hirsute distally; abaxially with a + truncate ridge near the base, surface

otherwise slightly concave, glabrous. Column 1.5—2 mm long, Stelidia

triangular, 0.5—0.6 long; acute. Anther: front margin erose, abaxially

papillose distally.

Colours: Sepals yellowish, distally with purplish veins, proximally

with a purplish blotch. Petals yellowish, suffused purplish distally. Lip

adaxially yellow or purplish with a dark purple median line, abaxially

yellowish.

Habitat & ecology: Lowland kerangas forest.

Distribution: MALAYSIA: Sarawak (2 specimens seen).

INDONESIA: Kalimantan (1).

| Species fere Cognita

Bulbophyllum sarcoscapum Teijsm. & Binn. (probably extinct)

Bulbophyllum sarcoscapum Teijsm. & Binn., Natuurk. Tijdschr. Ned.-Indie 29 (1867)

242.— TYPE: Indonesia, Java, ‘in forests of Mt. Madoer, near Bogor’, (collector

not indicated) (not seen).

Rhizome c. 2 mm diam., sections between pseudobulbs c. 3 cm long.

Pseudobulbs distant, depressed-conical to depressed-globose. Petiole 2.2—

3 cm. Leaf blade elliptic-obovate, 11—23 x 2.3—2.8 cm, index (length/

Bulbophyllum Sect. Altisceptrum and Hirtula 127

width) 4.7—9.1; acute to acuminate. Inflorescence an elongated, lax raceme,

pendulous, 22—26 cm, c. 25-flowered. Peduncle c. 16 cm, bracts c. 5, the

longest c. 12 mm long. Rhachis thickened, c. 0.8 cm diam., glabrous. Floral

bracts ovate, c. 3 x 2 mm, acute. Pedicel and ovary c. 3 mm. Flowers

opening wide. Sepals reflexed, c. 4 mm long, surface adaxially finely

pubescent, abaxially glabrous. Petals much shorter, semi-orbicular, margins

ciliate. Lip ?elliptic, c. 4 mm long, obtuse, margins with hairs; adaxially

furrowed, surface glabrous; abaxially with long hairs from base to tip.

Colours: Inflorescence purple. Flowers with blackish purple hairs.

Habitat & ecology: Montane forest.

Distribution: INDONESIA: Java (1 specimen seen). Not collected

since c. 1860.

Notes: No type material could be located, in spite of a search in LE

and W, where some of the Teysmann collections are stored. A sterile

specimen, Zollinger 947, from Bandung, Java (W) is likely to be the same

species. The description above is compiled from Teijsmann & Binnendijk,

and from the W specimen.

Bulbophyllum sarcoscapum is of uncertain taxonomic status; the

chances ever to find it again on Java seem remote. The information available

suggests a species rather similar to B. aithorhachis, but no median ridge on

the lip is mentioned in the original description.

Acknowledgements

I owe a debt of gratitude to many people and institutions who have helped

me (and still do) in many ways with my self-imposed task of revising as

many sections of Bulbophyllum as I possibly can. Here, I can mention only

a few. The spirit samples made from the extensive living collections in the

Botanic Gardens in Leiden, and the by now almost legendary, but

unfortunately no longer existing, living collection of Bulbophyllum of P.

Jongejan, Amersfoort (both in the Netherlands, the material kept in L)

must be mentioned. The herbaria AAU, AMES, B, BM, BRI, BO, C, F,

FOKEP, RLUVE, LAP, NY, PPE, S; 8ZU;,SAN; UKMB, W, WU ‘have

made material available for study. Jim Cootes (Australia), Bob & Chittima

Cowen (Thailand), Bev Debrincat (Australia), Gunter Fischer (Austria),

Alexander Kocyan (The Netherlands), Alvin Lok (Singapore), Peter

O’Byrne (Singapore), Anton Sieder (Austria), Heok Hui Tan (Singapore)

provided material of several rare species. J.F. Veldkamp (The Netherlands)

provided the Latin diagnoses.

128 Gard. Bull. Singapore 54 (2002)

References

Garay, L.A., Hamer, F., & Siegerist E.S., 1994. The genus Cirrhopetalum

and the genera of the Bulbophyllum alliance. Nordic Journal Botany 14:

609—646.

Seidenfaden, G. 1979. Orchid genera in Thailand 8: Bulbophyllum Thou.

Nordic Journal Botany 33, 3: 1—228.

Vermeulen, J.J. 1993. A taxonomic revision of the genus Bulbophyllum,

sections Adelopetalum, Lepanthanthe, Macrouris, Pelma, Peltopus and

Uncifera. Orchid Monographs 7: 1—324.

Bulbophyllum Sect. Altisceptrum and Hirtula 129

Identification lists

This identification list includes all collections examined for this revision.

Identification list sorted according to the species:

111 - Bulbophyllum penduliscapum - Beccari, O. PB 3113: PB 3144: Hort. Bogor

HLB 905,234-4797; HLB 924,325-179: Hort. Bogor . comm. nr. 008; 064; 091: 134:

420: Hort. Leiden 026209; Jacobson, E.R. s.n.; Jongejan, P. cult. 2990; Lamb, A.

1989/1105; SAN (Amin & Sigin) 069164: (Lamb, A.) 092332: Tenom Orchid Centre

(T.O.C.) cult. 1787: Vermeulen, J.J. 1158; 1302: Vermeulen, J.J.. & Duistermaat.

H. 1147; Wenzel, C.A. 761; 865.

112 - Bulbophyllum gymnopus - Clarke, C.B. 42549: F_.K.W. 7705: Herb. Hooker f.

s.n.; Gnffith 5133; 5134; Hort. Kew EN 596-68; King’s collectors s.n.; Pantling, R.

357; Parry, N.E. 484: Seidenfaden, G. & Smitinand, T. (GT) 8643: 8869: Tessier-

Yandell 02.

113a - Bulbophyllum farinulentum subsp. farinulentum - Beaman, J.H. (et al.)

07006; Carr, C.E. 0106; FRI (Ng, F.S.P.) 27102: Hort. Leiden 020771: 026544:

914306; Hort. Singapore SBG-O 0594; Kerr, A.F.G. 113; Lamb, A. s.n.; (Supat,

A.) 1989/ 1101; SAN (Fidilis Krispinus) 118814.

113b - Bulbophyllum farinulentum subsp. densissimum - Hort. Leiden 914076:

SAR (Ashton, P.S.) 16531: Synge, P.M. 98.

114 - Bulbophyllum fulvibulbum - Hort. Leiden 930900; 970299: Jongejan, P. cult.

1814; Vermeulen, J.J. 0603.

115 - Bulbophyllum nemorale - BSC (Fenix, E.) 28356: Kalkman. C., & Tissing,

M.O. 4187; PNH (Edano, G.E.) 12707: Wenzel. C.A. 871; 922: 928.

116 - Bulbophyllum disjunctum - Argent, G. & Coppins, B. 1167; Bakhtiar Effendi

bin Yahya 56; Beaman, J.H. (et al.) 06764; (et al.) 07238; (et al.) 11128; Carr, CE.

3212: Clemens, J.. & Clemens, M.S. 00254: 27361; 32554; 32787: 40226: Collenette,

S. 1002: Comber, J. 103; FRI (Saw, L.G.) 71675; Lamb, A. 1983/ 158; 1990/ 1246:

Mahadimenakbar 64: Nor Hashimah Abu Hani 44: SAN (Aban Gibot) 056304:

(Lamb, A.) 087497: SAR (Native coll.) 05049; (Asah Ak Luang) 22741: (Chai, P.)

33968; (Martin, P.J.) 38167; (Ilias Paie) 40717: (Awa, D., & Yii, P.C.) 46827: (Yu.

P.C., & Abu Talib) 58237; (Yu, P.C., & Abu Talib) 58476: Sato, T. 0793: (et al.)

1849: SFN (Moulton, J.C.) 06664; UKM (Mohd. Kasim) 23704; Vermeulen, JJ... &

Duistermaat, H. 0697: Vogel, E.F. de 8212.

117 - Bulbophyllum placochilum - Clemens, J.. & Clemens, M.S. 33922: 34056:

SFN (Carr, C.E.) 27938.

118 - Bulbophyllum elongatum - Blume, C.L. 195; 287: s.n.; HLB 902,322-934:

BSC (Loher, A.) 14730; Carr, CE. (1929/03) s.n.; Curtis, G. s.n.; Elmer, A.D.E.

09823; Grieve, H.G. 32: Hort. Leiden 026216: Mohd. Shah 2788; Reeve, T.M.

1098; 6025; Ridley, H.N. (1893) s.n.; (1893) s.n.; (1908/11) s.n.; Royen, P. van,

Sleumer, H., & Schram, F. 7795; Sands, M.J.S. 5285; SAR (Lai, S.T., Rantai, &

130 Gard. Bull. Singapore 54 (2002)

Awg Enjah) 75308; SFN (Holttum, R.E.) 21582; Wenzel, C.A. 351; 704.

119 - Bulbophyllum lissoglossum - Carr, C.E. 3371; Clemens, J., & Clemens, M.S.

29820: Griswold J.A. 79; SAN (Lamb, A., “KL. 3021”) 087149; (Lamb, A., “LKC

3190”) 091580: SFN (Carr, C.E.) 27249; Vermeulen, J.J.. & Chan, C.L. 0390;

Vermeulen, J.J., & Duistermaat, H. 1086.

120 - Bulbophyllum gemma-reginae - Jongejan, P. cult. 1856.

121 - Bulbophyllum carinilabium - Beaman, J.H. 08952: Hort. Leiden 026443;

Sato. T. 0265; SFN (Carr, C.E.) 26987; Vermeulen, J.J. 0641a.

122 - Bulbophyllum hirtulum - Ahmad (1894) s.n.; Ajoeb (Exp. Jacobson, E.R.)

807: Carr, C.E. 0194; (1933/03) s.n.; Chan, C.L. s.n.; Collenette, S. 0719: Hort.

Bogor HLB 920,342-4: Hort. Leiden 913046; 914195; 970483; Hort. Singapore

(Carr, C.E., & Holttum, R.E.) s.n.; (Ridley) (1909) s.n.; (Tan, H.H.) SBG-O 0316;

SBG-O 2166; Korthals, P.W. s.n.; Lewis, G.P. 291; Ridley, H.N. & Curtis, G.

(1899/03) s.n.; Tenom Orchid Centre (T.O.C.) cult. 1203; Vermeulen, J.J. 0636.

123 - Bulbophyllum cerebellum - Carr, C.E. 3470; Giles 998; Kocyan, A. (1999/10/

22) s18-

124 - Bulbophyllum clipeibulbum - Hort. Salzburg OR 637/94: Hort. Singapore

SBG-O 2087.

125 - Bulbophyllum penicillium - Hort. Glasnevin s.n.; (1917/05/24) s.n.; O’Brien,

J. cult. (1895/06) s.n.; Pantling, R. 251; Parish, C. 303; Prain’s collector 89.

126 - Bulbophyllum setuliferum - FRI (Saw, L.G.) 44299: Hort. Singapore SBG-O

0490: SBG-O 0499; SBG-O 0625; Lok, F.A. cult. 053.

127 - Bulbophyllum sororculum - Brans, M. s.n.

128 - Bulbophyllum aithorhachis - Jongejan, P. cult. 2219: Schrage, A.H. (1954/03/

01) s.n.

129 - Bulbophyllum echinochilum - Hort. Muenchen (Loher) (1916/12) s.n.

130 - Bulbophyllum lanuginosum - Thaitong 1395.

131 - Bulbophyllum atratum - Chang, C.K. cult. s.n.; Hort. Bogor s.n.; HLB 921,250-

380; Hort. Singapore SBG-O 0087; Jacobson, E.R. s.n.

132 - Bulbophyllum groeneveldtii - Hort. Leiden 026335; Jacobson, E.R. cult.

(Groeneveldt) 951; Jongejan, P. cult. 1818; Vermeulen, J.J.. & Duistermaat, H.

0971; 1107.

133 - Bulbophyllum grotianum - Hort. Leiden 970530.

134 - Bulbophyllum janus - Micholitz, W. s.n.; Robinson, H.C., & Boden Kloss, C.

(1914/03/15) s.n.

135 - Bulbophyllum xiphion - Hort. Leiden 913241; Vogel, E.F. de, & Cribb, P-J.

9062.

136 - Bulbophyllum mirabile - Hort. Bogor HLB 908,100-122; HLB 924,325-695:

Hort. Bogor , comm. nr. 006; 036; 153; Hort. Glasnevin s.n.; Hort. Leiden 949339;

Jongejan, P. cult. 4031.

138 - Bulbophyllum polycyclum - Hort. Leiden 913457; Jongejan, P. cult. 1870.

Bulbophyllum Sect. Altisceptrum and Hirtula 131

139 - Bulbophyllum lindleyanum - Day, J. 63: F.F. 48: Foerstermann, LF. 08:

Herb. Reichenbach, H.G. (Foerstermann) 15563: Hort. Singapore SBG-O 2185:

SBG-O 2186; Lok, F.A. cult. 057; Seidenfaden, G. & Smitinand, T. (GT) 6139:

SFN (Haniff & Nur) 02702: (Haniff & Nur) 10301.

140 - Bulbophyllum tremulum - Beddome, R.H. 8179: Herb. Reichenbach, H.G.

(Wight) 49425; Hort. Kew (1925/04/27) s.n.; Hort. Leiden 022802: Jongejan, P.

cult. 4544: Lawrence, T. cult. (1901/02/28) s.n.: Meebold, A. 8533.

141 - Bulbophyllum scaphiforme - Hort. Vienna 1992/ 182: Seidenfaden, G. &

Smitinand, T. (GT) 2099; 7464; 8332: 8332a: Tixier, P. s.n.

142 - Bulbophyllum nigripetalum - Henry, A. 13055; Hort. Glasnevin (1892/05)

s.n.; Hort. Kew (1915/01) s.n.; (1916/03/13) s.n.; (1919/05/22) s.n.; Lawrence, T.

cult. (1899/04) s.n.; Maxwell, J.F. 1989/ 524: Morrison, A. 1032; O’Brien, J. cult.

(1888) s.n.; (1889/04) s.n.; (1891/05/18) s.n.: Seidenfaden, G. & Smitinand, T. (GT)

7656.

143 - Bulbophyllum nigrescens - Cumberlege 1021: Hort. Aarhus 1978/ B-232:

Hort. Vienna 099B 104; 1995/ 446; Jongejan, P. cult. 2591; Kerr, A.F.G. 084:

(1915/03/14) s.n.; Lawrence, T. cult. (1898/02) s.n.; Maxwell, J.F. 1989/ 523; Menzies,

D., & Dupuy, D. 328; Seidenfaden, G. & Smitinand, T. (GT) 8332b; Smitinand, T.

10168.

144 - Bulbophyllum trulliferum - Tenom Orchid Centre (T.O.C.) cult. 2321.

145 - Bulbophyllum jolandae - Vermeulen, J.J. 0602.

146 - Bulbophyllum comberipictum - Hort. Leiden 914194.

147 - Bulbophyllum lasioglossum - Borden, T.E. 797; Elmer, A.D.E. 17687:

Topping, D. LeRoy (1904/05) s.n.

148 - Bulbophyllum negrosianum - Elmer, A.D.E. 09821: Hort. Vienna 1990) 176.

149 - Bulbophyllum dracunculus - Carr, C.E. 3647; Clemens, J.. & Clemens, MLS.

32908; Collenette, S. 0535; Hort. Leiden 030978.

150 - Bulbophyllum polygaliflorum - Carr, C.E. 3595: Hansen, C. 499: SFN (Carr,

Fh any Ap iD

151 - Bulbophyllum ochthodes - Hort. Singapore SBG-O 0616.

152 - Bulbophyllum phreatiopse - Hoogland, R.D. & Craven, L.A. 10899: Reeve,

T.M. 5772.

153 - Bulbophyllum parviflorum - Cooper, R.E. 1239; Hort. Glasnevin (1914/10)

s.n.; (1915/10/13) s.n.; Hort. Leiden 960104; Jongejan, P. cult. 1974; King, G. s.n.;

Pantling, R. 245; Parish, C. 305; Seidenfaden, G. & Smitinand, T. (GT) 8524; 8549:

Thomson (1850) s.n.

154 - Bulbophyllum secundum - Beusekom, C.F. van, & Phliengkhlai, C. 1271:

Evrard, F. 1018; Garrett, H.B.G. 795; Grey-Wilson & Phillips 93; Hort. Glasnevin

(1904/07) s.n.; Jongejan, P. cult. 0303; 0889; Kerr, A.F.G. 180; Maxwell, J.F. 1988

685; Pantling, R. 214; Prain, D. 41; Samuel, A. 13549; Sander cult. (1895/06) s.n.;

Swinhoe, R. 122; Tsi, Z.H. 1991/ 544; White, K. 406.

155 - Bulbophyllum debrincatiae - Debrincat, B. cult. (leg. Cootes, J-E.) O01.

132 Gard. Bull. Singapore 54 (2002)

156 - Bulbophyllum pilosum - Vermeulen, J.J. 0562.

157 - Bulbophyllum xenosum - Jongejan, P. cult. 4425.

158 - Bulbophyllum limbatum - Burkill, H.M. 1789; Carr, C.E. (1920/08) s.n.;

Hort. Bogor HLB 914,150-360; HLB 921,250-248; Hort. Leiden 960104; 970478;

Hort. Singapore (1932/08) s.n.; Jeppesen, H.L. (1920/08) s.n.; Jongejan, P. cult.

3995; 4256; Lamb, A. 1988/ 925; Loddiges, C. s.n.; Mohd. Shah & Kadim 383; SAN

(Dewol Dundaling) 134960; SFN (Moulton, J.C.) 06757; (Moulton, J.C.) 06758:

Tenom Orchid Centre (T.O.C.) cult. (Lamb, A.) (1989/01) s.n.; Teruya, Z. 1890.

159 - Bulbophyllum rariflorum - Hort. Bogor (1909) s.n.; HLB 913,114-114; I,

M.C. 133; Hort. Bogor , comm. nr. 116; 128; 130; 136; Jongejan, P. cult. 2013; 2214.

* - Bulbophyllum sarcoscapum - Zollinger, H. 947.

Bulbophyllum Sect. Altisceptrum and Hirtula 133

Identification list sorted according to the specimens:

The identifications are given as numbers between square brackets. These numbers

correspond with the species numbers in this revision. To facilitate the use of the

identification lists, an enumeration of the taxa with their numbers is given.

[111] B. penduliscapum J.J. Sm. M351 b. xiphion:s.J. Verm.

[112] B. gymnopus Hook.f. [136] B. mirabile Hallier.

[113] B. farinulentum J.J. Sm. [137] B. spadiciflorum Tixier.

[113a] ssp. farinulentum [138] B. polycyclum J.J. Verm.

[113b] ssp. densissimum (Carr) J.J. Verm. [139] B. lindleyanum Griff.

[114] B. fulvibulbum J.J. Verm. [140] B. tremulum Wight

[115] B. nemorale L.O Williams [141] B. scaphiforme J.J. Verm.

[116

]

] B. disjunctum Ames & C. Schweinf. [142]

117] B. placochilum J.J. Verm.

. nigripetalum Rolfe

. nigrescens Rolfe

elongatum (Blume) Hassk. 144] B. trulliferum J.J. Verm.

lissoglossum J.J. Verm. . jolandae J.J. Verm.

. comberipictum J.J. Verm.

. lasioglossum Rolfe ex Ames

. nhegrosianum Ames

. dracunculus J.J. Verm.

. polygaliflorum J.J. Wood

. ochthodes J.J. Verm.

[

[120] B. gemma-reginae J.J. Verm.

[

[126] B. setuliferum J.J. Verm. & L.G. Saw [152] B. phreatiopse J.J. Verm.

[

121] B. carinilabium J.J. Verm. [147]

122] B. hirtulum Ridl. [

123] B. cerebellum J.J. Verm. [

124] B. clipeibulbum J.J. Verm. [

127] B. sororculum J.J. Verm. [153] B. parviflorum Par. & Rchb.f. in Rchb.f.

128] B. aithorhachis J.J. Verm. [154] B. secundum Hook.f.

129] B. echinochilum Kraenzl. [155] B. debrincatiae J.J. Verm.

130] B. lanuginosum J.J. Verm. [156] B. pilosum J.J. Verm.

131] B. atratum J.J. Sm. [157] B. xenosum J.J. Verm.

132] B. groeneveldtii J.J. Sm. [158] B. limbatum Lindl.

133] B. grotianum J.J. Verm. [159] B. rariflorum J.J.Sm.

134] B. janus J.J. Verm. [*] | B. sarcoscapum Teysm. & Binn.

Ahmad (1894) s.n. [122]; Ajoeb (Exp. Jacobson, E.R.) 807 [122]; Argent, G. &

Coppins, B. 1167 [116].

Bakhtiar Effendi bin Yahya 56 [116]; Bans, M. s.n. [127]; Beaman, J.H. (et al.)

06764 [116]; (et al.) 07006 [113a]; (et al.) 07238 [116]; 08952 [121]; (et al.) 11128

[116]; Beccari, O. PB 3113 [111]; PB 3144 [111]; Beddome, R.H. 8179 [140];

Beusekom, C.F. van, & Phlengkhlai, C. 1271 [154]; Blume, C.L. 195 [118]; 287

[118]; s.n. [118]; HLB 902,322-934 [118]; Borden, T.E. 797 [147]; BSC (Loher, A.)

14730 [118]; (Fenix, E.) 28356 [115]; Burkill, H.M. 1789 [158].

Carr, C.E. 0106 [113a]; 0194 [122]; 3212 [116]; 3371 [119]; 3470 [123]; 3595 [150];

3647 [149]; (1920/08) s.n. [158]; (1929/03) s.n. [118]; (1933/03) s.n. [122]; Chan, C.L.

s.n. [122]; Chang, C.K. cult. s.n. [131]; Clarke, C.B. 42549 [112]; Clemens, J., &

Clemens, M.S. 00254 [116]; 27361 [116]; 29820 [119]; 32554 [116]; 32787 [116];

32908 [149]; 33922 [117]; 34056 [117]; 40226 [116]; Collenette, S. 0535 [149]; 0719

[122]; 1002 [116]; Comber, J. 103 [116]; Cooper, R.E. 1239 [153]; Cumberlege 1021

[143]; Curtis, G. s.n. [118];.

134 Gard. Bull. Singapore 54 (2002)

Day, J. 63 [139]; Debrincat, B. cult. (leg. Cootes, J.E.) 001 [155].

Elmer, A.D.E. 09821 [148]; 09823 [118]; 17687 [147]; Evrard, F. 1018 [154].

F.F. 48 [139]; F.K.W. 7705 [112]; Foerstermann, I.F. 08 [139]; FRI (Ng, F.S.P.)

27102 [113a]; (Saw, L.G.) 44299 [126]; (Saw, L.G.) 71675 [116].

Garrett, H.B.G. 795 [154]; Giles 998 [123]; Grey-Wilson & Phillips 93 [154]; Grieve,

H.G. 32 [118]; Griffith 5133 [112]; 5134 [112]; Griswold J.A. 79 [119].

Hansen, C. 499 [150]; Henry, A. 13055 [142]; Herb. Hooker f. s.n. [112]; Herb.

Reichenbach, H.G. (Foerstermann) 15563 [139]; (Wight) 49425 [140]; Hoogland,

R.D. & Craven, L.A. 10899 [152]; Hort. Aarhus 1978/ B-232 [143]; Hort. Bogor

s.n. [131]; (1909) s.n. [159]; HLB 905,234-4797 [111]; HLB 908,100-122 [136]; HLB

913,114-114 [159]; HLB 914,150-360 [158]; HLB 920,342-4 [122]; HLB 921,250-248

[158]; HLB 921,250-380 [131]; HLB 924,325-179 [111]; HLB 924,325-695 [136]; II,

M.C. 133 [159]; Hort. Bogor , comm. nr. 006 [136]; 008 [111]; 036 [136]; 064 [111];

O91 [111]; 116 [159]; 128 [159]; 130 [159]; 134 [111]; 136 [159]; 153 [136]; 420 [111]:

Hort. Glasnevin s.n. [125]; s.n. [136]; (1892/05) s.n. [142]; (1904/07) s.n. [154]:

(1914/10) s.n. [153]; (1915/10/13) s.n. [153]; (1917/05/24) s.n. [125]; Hort. Kew

(1915/01) s.n. [142]; (1916/03/13) s.n. [142]; (1919/05/22) s.n. [142]; (1925/04/27)

s.n. [140]; EN 596-68 [112]; Hort. Leiden 020771 [113a]; 022802 [140]; 026209

[111]; 026216 [118]; 026335 [132]; 026443 [121]; 026544 [113a]; 030978 [149]; 913046

[122]; 913241 [135]; 913457 [138]; 914076 [113b]; 914194 [146]; 914195 [122]: 914306

[113a]; 930900 [114]; 949339 [136]; 960104 [158]; 960104 [153]; 970299 [114]; 970478

[158]; 970483 [122]; 970530 [133]; Hort. Muenchen (Loher) (1916/12) s.n. [129];

Hort. Salzburg OR 637/94 [124]; Hort. Singapore (Carr, C.E., & Holttum, R.E.)

s.n. [122]; (Ridley) (1909) s.n. [122]; (1932/08) s.n. [158]; SBG-O 0087 [131]; (Tan,

H.H.) SBG-O 0316 [122]; SBG-O 0490 [126]; SBG-O 0499 [126]; SBG-O 0594

[113a]; SBG-O 0616 [151]; SBG-O 0625 [126]; SBG-O 2087 [124]; SBG-O 2166

[122]; SBG-O 2185 [139]; SBG-O 2186 [139]; Hort. Vienna 099B 104 [143]; 1990/

176 [148]; 1992/ 182 [141]; 1995/ 446 [143].

Jacobson, E.R. s.n. [131]; s.n. [111]; Jacobson, E.R. cult. (Groeneveldt) 951 [132]:

Jeppesen, H.L. (1920/08) s.n. [158]; Jongejan, P. cult. 0303 [154]: 0889 [154]; 1814

[114]; 1818 [132]; 1856 [120]; 1870 [138]; 1974 [153]; 2013 [159]; 2214 [159]; 2219

[128]; 2591 [143]; 2990 [111]; 3995 [158]; 4031 [136]; 4256 [158]; 4425 [157]; 4544

[140]. |

Kalkman. C., & Tissing, M.O. 4187 [115]; Kerr, A.F.G. 084 [143]; 113 [113a]; 180

[154]; (1915/03/14) s.n. [143]; King’s collectors s.n. [112]; King, G. s.n. [153]; Kocyan,

A. (1999/10/22) s.n. [123]; Korthals, P.W. s.n. [122].

Lamb, A. s.n. [113a]; 1983/ 158 [116]; 1988/ 925 [158]; (Supat, A.) 1989/ 1101

[113a]; 1989/ 1105 [111]; 1990/ 1246 [116]; Lawrence, T. cult. (1898/02) s.n. [143]:

(1899/04) s.n. [142]; (1901/02/28) s.n. [140]; Lewis, G.P. 291 [122]; Loddiges, C. s.n.

[158]; Lok, F.A. cult. 053 [126]; 057 [139].

Mahadimenakbar 64 [116]; Maxwell, J.F. 1988/ 685 [154]; 1989/ 523 [143]: 1989/

524 [142]; Meebold, A. 8533 [140]; Menzies, D., & Dupuy, D. 328 [143]: Micholitz,

W. s.n. [134]; Mohd. Shah 2788 [118]; Mohd. Shah & Kadim 383 [158]; Morrison,

A. 1032 [142].

Bulbophyllum Sect. Altisceptrum and Hirtula 135

Nor Hashimah Abu Hani 44 [116].

O’Brien, J. cult. (1888) s.n. [142]; (1889/04) s.n. [142]; (1891/05/18) s.n. [142]:

(1895/06) s.n. [125].

Pantling, R. 214 [154]; 245 [153]; 251 [125]; 357 [112]; Parish, C. 303 [125]; 305

[153]; Parry, N.E. 484 [112]; PNH (Edano, G-E.) 12707 [115]; Prain’s collector 89

[125]; Prain, D. 41 [154].

Reeve, T.M. 1098 [118]; 5772 [152]; 6025 [118]; Ridley, H.N. (1893) s.n. [118];

(1893) s.n. [118]; (1908/11) s.n. [118]; Ridley, H.N. & Curtis, G. (1899/03) s.n

[122]: Robinson, H.C., & Boden Kloss, C. (1914/03/15) s.n. [134]; Royen, P. van,

Sleumer, H., & Schram, F. 7795 [118].

Samuel, A. 13549 [154]; SAN (Aban Gibot) 056304 [116]; (Amin & Sigin) 069164

[111]; (Lamb, A., “KL. 3021”) 087149 [119]: ou ae A.) 087497 [116]: (Lamb, A..,

“LKC 3190”) 091580 [119]; (Lamb, A.) 092332 [111]: (Fidilis Krispinus) 118814

[113a]; (Dewol Dundaling) 134960 [158]; Sander cult. (1895/06) s.n. [154]; Sands,

M.J.S. 5285 [118]; SAR (Native coll.) 05049 [116]; (Ashton, P.S.) 16531 [113b]:

(Asah Ak Luang) 22741 [116]; (Chai, P.) 33968 [116]; (Martin, P.J.) 38167 [116]:

(Ilias Paie) 40717 [116]; (Awa, D., & Yii, P.C.) 46827 [116]; (Yu, P.C., & Abu

Talib) 58237 [116]; (Yui, P.C., & Abu Talib) 58476 [116]: (Lai, S.T., Rantai, & Awg

Enjah) 75308 [118]; Sato, T. 0265 [121]: 0793 [116]: (et al.) 1849 [116]: Schrage,

A.H. (1954/03/01) s.n. [128]; Seidenfaden, G. & Smitinand, T. (GT) 2099 [141]:

6139 [139]; 7464 [141]; 7656 [142]; 8332 [141]; 8332a [141]; 8332b [143]: 8524 [153]:

8549 [153]: 8643 [112]: 8869 [112]: SFN (Haniff & Nur) 02702 [139]; (Moulton.

J.C.) 06664 [116]; (Moulton, J.C.) 06757 [158]; (Moulton, J.C.) 06758 [158]: (Haniff

& Nur) 10301 [139]; (Holttum, R.E.) 21582 [118]; (Carr, C.E.) 26987 [121]: (Carr,

C.E.) 27249 [119]; (Carr, C.E.) 27938 [117]: (Carr, C.E.) 27956 [150]; Smitinand, T.

10168 [143]; Swinhoe, R. 122 [154]; Synge, P.M. 98 [113b].

Tenom Orchid Centre (T.0.C.) cult. 1203 [122]; 1787 [111]; 2321 [144]: (Lamb,

A.) (1989/01) s.n. [158]; Teruya, Z. 1890 [158]: Tessier-Yandell 02 [112]: Thaitong

1395 [130]; Thomson (1850) s.n. [153]; Tixier, P. s.n. [141]; Topping, D. LeRoy

(1904/05) s.n. [147]; Tsi, Z.H. 1991/ 544 [154].

UKM (Mohd. Kasim) 23704 [116].

Vermeulen, J.J. 0562 [156]; 0602 [145]; 0603 [114]; 0636 [122]; 0641a [121]; 1158

[111]; 1302 [111]; Vermeulen, J.J., & Chan, C.L. 0390 [119]; Vermeulen, J.J., &

Duistermaat, H. 0697 [116]: 0971 [132]; 1086 [119]: 1107 [132]: 1147 [111]; Vogel,

E.F. de 8212 [116]; Vogel, E.F. de, & Cribb, P.J. 9062 [135].

Wenzel, C.A. 351 [118]; 704 [118]; 761 [111]; 865 [111]; 871 [115]; 922 [115]; 928

[115]; White, K. 406 [154].

Zollinger, H. 947 [*].

in)

oS)

Gard. Bull. Singapore 54 (2002)

Bulbophyllum penduliscapum J.J.Sm. (111).—Jongejan cult. 2990

(P. Jongejan).

Bulbophyllum farinulentum J.J. Sm. ssp. densisimum (Carr) J.J.

Verm. (113b).— Hort. Leiden 914076 (A. Schuiteman).

Bulbophyllum fulvibulbum J.J. Verm. (114). — Hort. Leiden

930285 (A. Meivogel).

Bulbophyllum disjunctum Ames & C. Schweinf. (116). — Hort.

Leiden 980182 (A. Schuiteman).

—~

ie)

Ae)

138 Gard. Bull. Singapore 54 (2002)

5 Bulbophyllum gemma-reginae J.J. Verm. (120). — Jongejan cult.

1856 (P. Jongejan).

6 Bulbophyllum carinilabium J.J. Verm. (121).— Hort. Leiden 26443

(A. Schuiteman).

7 Bulbophyllum cerebellum J.J. Verm. (123). — Kocyan s.n. (A.

Kocyan).

8 Bulbophyllum clipeibulbum J.J. Verm. (124). — Hort. Vienna

O99B160-2 (G. Fischer).

3 Bulbophyllum penicillium Par. & Rchb.f. in Rchb.f. (125). — (P.

Jongejan).

Bulbophyllum Sect. Altisceptrum and Hirtula

140

Gard. Bull. Singapore 54 (2002)

Bulbophyllum setuliferum J.J. Verm. & L.G. Saw (126). — Hort.

Singapore SBG-O 499 (P. O'Byrne).

Bulbophyllum sororculum J.J. Verm. (127).— M. Brans s.n. (M.

Brans).

Bulbophyllum aithorhachis J.J. Verm. (128).— Jongejan cult. 2219

(P. Jongejan).

Bulbophyllum aithorhachis J.J. Verm. (128).— Jongejan cult. 2219

(P. Jongejan).

Bulbophyllum atratum J.J. Sm. (131). — Hort. Singapore SBG-

O 087 (P. O’Byrne).

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as)

Lena

Ss)

142

Gard. Bull. Singapore 54 (2002)

Bulbophyllum groeneveldtii J.J. Sm. (132). — Jongejan cult. 1818

(P. Jongejan).

Bulbophyllum grotianum J.J. Verm. (133).— Hort. Leiden 970603

(A. Schuiteman).

Bulbophyllum grotianum J.J. Verm. (133).— Hort. Leiden 970603

(A. Schuiteman).

Bulbophyllum xiphion J.J. Verm. (135). — Hort. Leiden 913241

(P. Jongejan).

Bulbophyllum xiphion J.J. Verm. (135). — Cribb 9062 (P. Cribb).

Bulbophyllum Sect. Altisceptrum and Hirtula

es Baia

cae

alge ’ pied

(os)

Gard. Bull. Singapore 54 (2002)

Bulbophyllum mirabile Hallier, flowers near the base of the

inflorescence (136). —Jongejan cult. 4031 (P. Jongejan).

Bulbophyllum mirabile Hallier, flowers near the top of the

inflorescence (136). —Jongejan cult. 4031 (P. Jongejan).

Bulbophyllum polycyclum J.J. Verm. (138).—Jongejan cult. 1870

(P. Jongejan).

Bulbophyllum lindleyanum Griff. (139). — Hort. Vienna s.n. (A.

Sieder).

Bulbophyllum tremulum Wight (140). — Jongejan cult. 4544 (P.

Jongejan).

Bulbophyllum nigripetalum Rolfe (142). — Hort. Vienna O2044

(A. Sieder).

~—

Ss)

cv)

bee

—

146

Gard. Bull. Singapore 54 (2002)

Bulbophyllum nigrescens Rolfe (143). — Jongejan cult. 2591 (P.

Jongejan).

Bulbophyllum nigrescens Rolfe (143).— Hort. Vienna 446.95 (A.

Sieder).

Bulbophyllum trulliferum J.J. Verm. (144). — Tenom Orchid

Centre (T.0.C.) cult. 2321 (A. Lamb).

Bulbophyllum lasioglossum Rolfe ex Ames (147). — (M.

Clements).

Bulbophyllum negrosianum Ames (148).— Hort. Vienna cult. (A.

Sieder).

Bulbophyllum Sect. Altisceptrum and Hirtula

148

Gard. Bull. Singapore 54 (2002)

Bulbophyllum dracunculus J.J. Verm. (149).— Hort. Leiden 30978

(P. Jongejan).

Bulbophyllum ochthodes J.J. Verm. (151). — Hort. Singapore

SBG-O 616 (P. O’Byrne).

Bulbophyllum parviflorum Par. & Rchb.f. in Rchb.f (153). —

Jongejan cult. 1974 (P. Jongejan).

Bulbophyllum secundum Hook.f. (154). — Jongejan cult. s.n. (P.

Jongejan).

Bulbophyllum secundum Hook.f. (154). — Jongejan cult. s.n. (P.

Jongejan).

Bulbophyllum Sect. Altisceptrum and Hirtula 149

150 Gard. Bull. Singapore 54 (2002)

36 Bulbophyllum debrincatiae J.J. Verm. (155).— Debrincat cult. (leg.

Cootes) 001 (D. Titmuss).

37 Bulbophyllum xenosum J.J. Verm. (157). — Jongejan cult. 4425

(P. Jongejan).

38 Bulbophyllum limbatum Lindl. (158). — Jongejan cult. (P.

Jongejan).

Ss)

is)

—

cv)

—

cal

Gardens’ Bulletin Singapore 54 (2002) 153-154.

Bulbophyllum praetervisum J.J. Verm. (Orchidaceae),

an Overlooked Species Close to B. macranthum Lindl.

J.J. VERMEULEN

Singapore Botanic Gardens

Singapore 259569

Abstract

Bulbophyllum praetervisum sp. nov. is described, a species very similar in general aspect to

Bulbophyllum macrantum Lindl., but differing in the shape of the column and the lip

While revising Bulbophyllum sect. Sestochilus, | found that specimens of

two different species were included under B. macranthum Lindl. in

Vermeulen (1991: 263). Vegetatively and in the general aspect of the flowers

the two are remarkably similar. However, they clearly differ in the shape

of the column and the lip. One is doubtless B. macranthum Lindl., but the

other is new. It is formally described below; full information will be given

in the forthcoming revision of the section.

Bulbophyllum praetervisum J.J. Verm., sp. nov.

Bulbophyllum praetervisum J.J. Verm., a B. macrantho ob dentem triangularem secus

stelidiorum marginem inferiorem et labelli apicem valde recurvatum differt. —

TYPE: Malaysia, Sabah, Bukit Hampuan, Tenom Orchid Centre (T.O.C.) cult.

934 (K, holo.)

Other material seen: BRUNEI. No exact locality, Van Niel 3845 (L).

Notes: The holotype specimen is depicted in Vermeulen (1991: 262).

Bulbophyllum praetervisum differs from B. macranthum Lindl. in having a

triangular, acute tooth along the lower margin of each stelidium. The lip is

also different, particularly in lateral view: in B. praetervisum it is almost

straight proximally, with only the tip recurved in a hook-like manner; in B.

macranthum it is gradually curved over most of its length.

While B. macranthum is almost restricted to lowland conditions, B.

praetervisum also tolerates higher altitudes: in Sabah it has been found at

1500 m alt.

154 Gard. Bull. Singapore 54 (2002)

References

Vermeulen, J.J. 1991. Orchids of Borneo 2: Bulbophyllum. Royal Botanic

Gardens, Kew, and Toihaan Publishing Company Sdn. Bhd., Kota

Kinabalu, Sabah, Malaysia.

Gardens’ Bulletin Singapore 54 (2002) 155-157.

Book Review: Ian M. Turner. 2001. The Ecology of Trees in the Tropical

Rain Forest. 298 pages. Cambridge University Press, Cambridge, United

Kingdom. ISBN 0-521-80183-4. Price: S$75.90

This book by Ian Turner represents an outstanding introduction to the

biology of tropical rainforest trees, and represents an ideal background

text for researchers and students wanting to dig more deeply into the

subject. It would be ideal for research students in plant ecology prior to

conducting more field work, it would be essential background for students

in animal ecology to understand the system in which animals are living,

and it would represent the logical next book for those people who enjoyed

reading the more general books by T.C. Whitmore (An Introduction to

Tropical Rain Forests) and P.W. Richards (The Tropical Rain Forest). This

new book has its focus on the individual tree, rather than the forest itself

as in these previous two books. As such, the book presents material,

which connects with these books, but does not duplicate their focus. The

book would also make an excellent discussion book for a graduate level

course on tropical trees.

The book is organized according to the life cycle of the tree, beginning

with the growing tree, with good coverage of the trunk, the roots and

leaves. The next chapter covers tree performance, considering growth and

mortality. Reproductive biology comes next, considering breeding systems,

pollination and fruit dispersal. Logically following is a chapter on seed

size, and seedling form and ecology. The final chapter considers the

characteristics and usefulness of systems of tree classifications, including

pioneer versus climax trees, level of shade tolerance, and size and canopy

position at the time of flowering.

Within each of these chapters, the material is organized by more

specific topics. The overall effect is very satisfying. Individual topics are

introduced, the major issues are presented, controversies are presented,

the most relevant research is referenced and summarized, and then areas

requiring further research are outlined. In effect, much of the presentation

is organized as mini-reviews of the most important topics in tropical tree

ecology. How does sap ascend tropical trees in a high humidity

environment? Why do trees have buttresses? Do mineral nutrients, light

or other factors limit tree growth? Why are the leaves of so many tropical

trees similar in shape? Do trees have clumped or scattered distributions,

and are patterns caused by density-dependant effects? Can seedlings and

adult trees be classified into shade-tolerant and shade-intolerant species,

or are they better classified as pioneer and climax species?

The book will be useful to researchers with both general and specific

156 Gard. Bull. Singapore 54 (2002)

interests. For example, I have wondered whether tropical legume trees in

the forest fix nitrogen. I have periodically asked people about this, but

I have not gotten a specific or authoritative answer. This specific topic is

dealt with here with a clear and efficient one-and-half page summary of

the latest ideas and literature. Information is presented that 23% of

caesalpinioid legumes, which are trees in the forest, have been recorded to

form root nodules with nitrogen-fixing bacteria; much less than the 97% of

papilionoid legumes which form nodules. It may be that low phosphorus

availability, soil acidity, and abundant aluminium inhibit nitrogen fixation

in these systems. However, work with stable isotopes does indicate nitrogen

fixation in many tropical tree legumes, casuarina trees and cycads. The

section also includes a brief mention of the presumed evolutionary origins

of this relationship. I mention this example in detail because it was one

that I was particularly interested in, and here found such a concise and

informative discussion.

I think that for many students and researchers the extended discussion

of the evidence for shade intolerance and shade tolerance, pioneer and

climax species, and tree size would well be worth the price of the book.

There is abundant research on this topic, but Turner summarizes the key

evidence and points out the correlations among these various ways of

looking at trees, and the preponderance of continuity of species along axes

of characteristics rather than species fitting into neat categories.

There is one way in which I think the book fails to live up to its

promise. On the final page, in his final section, Where does this leave us?

Turner states, “One of my objectives when I set out to write this book was

to see if there was a new synthesis of the comparative ecology of tropical

trees awaiting discovery amidst the voluminous literature on tropical rain

forests. I have failed to find a new synthesis.” [| think Turner is wrong

here on two counts. First, he has provided numerous syntheses of significant

controversies in tropical ecology, which will guide research projects for

years to come. People will read this book and obtain ideas about what are

likely to be productive areas of research.

But more importantly, I think that Turner has failed to outline the

ways in which tropical trees will respond to the ever-increasing threats to

forests posed by human activity. These problems represent the next major

round of research in this field, but Turner does not discuss them. For

example, what will be the impact on tree biology of global climate change?

If the climate becomes warmer or drier, or the atmosphere has a higher

carbon dioxide concentration, how will this affect individual trees and the

forest in general? Will this tip the scale to trees with a certain type of

physiology or growth form or shade intolerance? If forests are increasingly

Book Review 157

fragmented by human activities, including logging, agriculture, roads,

ranching, and fire, how will this affect trees? And lastly, how will tropical

trees respond to human-caused changes in their reproduction? Introduced

Africanized honeybees are taking over pollination systems throughout Latin

America, seed dispersing animals are being hunted out of the forest

throughout the tropics to the point that many trees are no longer having

their seeds dispersed, and cattle and fire in the forest understorey are

eliminating the forest understorey in many places. These topics considered

together represent the new direction of research in the ecology of tropical

trees. Even though Turner did not outline this next major area of research,

his book represents a tremendous advance in the field and is to be greatly

appreciated.

Richard B. Primack

Biology Department

Boston University

5 Cummington Street

Boston, MA 02215, U.S.A.

Gardens’ Bulletin Singapore 54 (2002) 159-160.

Book Review: J.B. Comber 2001. Orchids of Sumatra. x + 1026 pages. The

Royal Botanic Gardens, Kew, U.K. ISBN 1 84246 027 7. Price: GBP 95.00,

RM 450.00.

Until the publication of this book, the orchid flora of Sumatra had been

badly neglected, particularly when you consider all the recent publications

on the orchids of Java, Borneo, Peninsular Malaysia and Thailand. The

main source of information has long been a checklist compiled by J.J.

Smith and published in 1933. Any attempt to identify a Sumatran orchid

usually started with a rather discouraging check in J.J. Smith’s list, it

continued with a lengthy search through a lot of old journals and the

literature on the orchid flora of surrounding areas, and it all too often

proved indecisive. It could be species X, but it could equally well be species

Y. This changed when Jim Comber decided to combine the information

present in the old literature with his excellent knowledge of west Malesian

orchids and the new information (mainly photographs) that he had gathered

during the years he lived on Sumatra. The first orchid flora of Sumatra is

now a fact.

The book is written following the lines of a normal flora: there is a

short introduction to the island of Sumatra, its people, its history and its

vegetation. This is followed by a key to the orchid genera, and keys to all

the orchid species. Each genus and species is shortly described; similarities

and taxonomic problems are shortly discussed below each taxon. For a

relatively small number of species, analytical drawings have been included,

most of these copied from originals by Rudolf Schlechter. And then, of

course, there are the numerous large colour photographs of magnificent

quality, most of which show close-ups of the flowers. While many of the

photographs show the plant growing in situ, a large number show plants

that were growing elsewhere in SE Asia, so they may not represent the

precise form found in Sumatra. We understand that much of the delay in

publishing this long-awaited volume was due to Comber’s insistence that

the photographs appear throughout the book alongside the relevant text,

rather than the much cheaper option of grouping them together and

inserting them as a series of colour plates. This treatment has resulted in a

book that has attained a much higher standard of presentation than most

floras; not only is the book informative, but it is a pleasure to use.

Of course, this volume also has a number of shortcomings, most

obviously those that result from the fact that there are still so many gaps in

our knowledge of the orchid flora of Sumatra. For example, many species

are known from the type material only, often collected long ago and in

several cases unfortunately now lost. For Comber, it must have been

160 Gard. Bull. Singapore 54 (2002)

frustrating to deal with such species, and problematic attempting to give

them their proper place in the enumeration. Another frustration must

have been Comber’s inability to collect fresh material; this has resulted in

several undescribed species being illustrated by photographs, but lacking a

formal name, description or herbarium specimen. It is a real shame that

these species could not have been described properly. There are also some

shortcomings of a more technical nature; it would have been useful if more

analytical sketches had been included. J.J. Smith’s drawings are available

to only relatively few people. They could have been copied just like

Schlechter’s drawings, and they would have facilitated identification.

As will be clear from the above, and as stated by Comber himself,

this book is a preliminary study. Much work remains to be done, and large

areas still need to be explored properly. Comber adds modestly *... but at

least it is a start’. It is definitely more than that. It is an excellent work of

reference for everybody interested in the orchid flora of West Malesia.

Jaap J. Vermeulen and Peter O’Byrne

Singapore.

Gardens’ Bulletin Singapore 54 (2002) 161.

Book Review: R. Govaerts, D.G. Frodin and T.D. Pennington 2002. World

Checklist and Bibliography of Sapotaceae. xi + 361 pages. The Royal

Botanic Gardens, Kew, U.K. ISBN 1 900347 94 6. Softback. Price: GBP

d7.50:

The fifth in the series, this volume covers the Sapotaceae, an important

tropical family particularly in the Malesian region. This volume includes 54

genera and 1,250 species as well as 3,689 synonyms below generic rank.

The layout, as with previous volumes, is clear and well organized making it

quick and easy to retrieve information. Unlike other checklists, the

bibliography gives a brief summary of the important conclusions for the

major references and the species lists aim to be taxonomically validated

based mainly on an assessment of the literature but where necessary

consulting specialists and the herbarium. Where there is doubt about

identity, this is indicated.

The volume is essential reference for botanists working away from

major libraries and will save a great deal of time for individual taxonomists

to drawn up species lists from scattered literature. In this connection,

providing the distribution of species is particularly useful.

I ‘tested’ its usefulness in comparing the names of the Bornean

Sapotaceae in the latest volume of Tree Flora of Sabah and Sarawak (April,

2002). The Kew Sapotaceae had captured all names but had ‘misspelt’

Madhuca sarawahensis (as ‘sarawakensis’). On the other hand, they were

correct in the spelling of Palaquium cochleariifolium, which was misspelt

in the Tree Flora. The notable omission was the five new species and four

other name changes made by Yui and Chai (2001). This is a serious limitation

of printed work in that it can very soon be out of date.

References

Yu, P.C. and P.P.K. Chai. 2001. New combinations, new names and new

species of Madhuca (Sapotaceae) from Sabah and Sarawak, Borneo.

Gardens’ Bulletin Singapore. 53: 343—356.

Chai, P.P.K. and P.C. Yui. 2002. Sapotaceae. Tree Flora of Sabah and

Sarawak. 4: 203—345.

Ruth Kiew

Singapore Botanic Gardens

Singapore

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Gardens’ Bulletin Singapore 54 (2002) 163-164.

Book Review: J.F. Maxwell and S. Elliott 2001. Vegetation and Vascular

Flora of Doi Sutep-Pui National Park, Northern Thailand. Biodiversity

Research and Training Program, Bangkok, Thailand. x + 205 pages, 36

colour photographs. ISBN 947 7360 61 6. Price: US$ 10.

The Doi Sutep-Pui National Park west of Cheng Mai city was established

in 1981, its higher peak reaches 1,685 m. It is the site of the famous 600-

year old Doi Sutep Temple.

This is a landmark publication and sets a high standard as a model

for biodiversity inventories of National Parks. In SE Asia and Malesia,

there is almost a complete dearth of accurate and complete information

for the plant species and associations found in National Parks. Part II by

Maxwell provides a detailed annotated enumeration of vascular plants (2,247

species in 195 families) in the Doi Sutep-Pui National Park based on

extensive fieldwork over a period of 13 years. The strength of this section

is in the care taken over accuracy of identification based on an extensive

collection of herbarium specimens.

Plant associations in the Park are described in Part II by Maxwell

and Elliott based on data from transects. The account details the vegetation

structure, species composition for all life forms and their abundance, and

the correlation between altitude, soil type and disturbance for the six

associations found in the Park. Their work clearly shows that the primary

evergreen seasonal forest without pine is the most biodiverse association,

both in number of species and in having more than double the number of

rare and endangered species, and that the summit vegetation has almost

completely been destroyed. Interwoven into the account is an interesting

discussion of the relative abundance of deciduous and evergreen associations

and the effect of disturbance, particularly of fire, on them. This section

also includes a historical account of plant collecting in the Park area from

the first collections made by C. Hosseus in 1904 as well as listing species

originally described from the Park (confusingly referred to in the text as

‘new species’ although many were described about a hundred years ago!).

Of particular value is Elliott’s description in Part IV of management

in the Park to maintain its integrity and biodiversity when 6,460 Hmong

villagers live within the Park and another 51,109 on the boundary. Already

43 per cent of the Park’s land has been converted to agriculture. Elliott’s

active involvement with the Forest Restoration Research Unit is a practical

example of how local communities can be involved in tree planting for

reforestation and for fire prevention; how in planting 40 indigenous tree

species (of the 600 tree species found in the Park) research is needed on

their basic biology in the selection and generation of seed and the

164 Gard. Bull. Singapore 54 (2002)

establishment of seedlings; and how the Park serves as a living laboratory

for education and research; and as a vital seed source of wild and

endangered trees.

Maxwell in Part I takes the opportunity to provide a detailed

historical review of the various systems in the literature for classifying the

Thai flora into vegetation types. Based on his own field work, he then

makes a plea for vegetation types to be based on a holistic approach that

takes into account a broader range of species (not just a few economically

important timber trees or a few dominant tree species) as well as vegetation

structure, that distinguishes phases of plant succession (especially to

appreciate the effect of fire on vegetation), and that places more emphasis

on altitude as well as seasonality. He stresses that all forest types in Thailand

are seasonal vegetation types and that describing the seasonal forest in

southern Thailand as ‘tropical rain forest’ is an incorrect application of the

term. Whether his conclusions will indeed be widely accepted or are widely

applicable in Thailand can now be tested against the clear framework that

he has provided, besides setting standards for the detailed ecological work

required to make this assessment.

The conclusion from reading the book is that the Doi Sutep-Pui

National Park is a flagship for biodiversity in Thailand with 20 per cent of

the Thai flora in just 261 km’. It also points to the urgent necessity of

conservation management as already 55 species recorded from the Park

were not recollected by Maxwell and Elliot and presumably no longer

occur there, while another 53 are critically endangered. Scientifically, it is

especially important as the type site of 512 species, more than any other

National Park or locality in Thailand. The problems the National Park

faces from fire, lack of watershed management, encroachment by

agriculture, and ‘development’ for tourism are common to many National

Parks in SE Asia and Elliott’s positive approach therefore has wide

applications elsewhere. On the other hand, Maxwell’s antagonist reference

to eco-pornography is counter-productive.

The book ends with a comprehensive list of references and an index

to species. It contains very few typographical errors, though page 7 records

average rainfall as 1000—2000 m! For the wealth of information, the book

is inexpensive and should not only be in the libraries of all botanical

institutions in the region but also in the resource centres of National Parks

and environmental NGOs.

Ruth Kiew

Singapore Botanic Gardens

Singapore

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Gardens' Bulle tin

"APR 1 @ 2003 Singapore ~ a

ARNOLD ~

ARBORETUM

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The Gardens' Bulleti

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VOL. 54 (Part 2) December 2002 ISSN 0374-7859

CONTENTS

Gideon, O.G.

Coprosma bougainvilleensis (Rubiaceae), a new species from Bougainville Island,

Be AO NS NNN A eae rs et ats sande rie TBE tad Precun Poem Beals RUM ae aoteablenemaopoen Me eos iudatamebirsvae Recent 165

Hay, A and C. Herscovitch

Two remarkable new West Malesian Homalomena (Araceae) SpeCles ..........:ccccceeceeeee iyo

Kloet, S.P. Vander

Re-cxammation of Vaccinium dialypetalum (ETICACCAC) «...1..is00.csveenssaseseceseoesoncarneaseenee 179

Mabberley, D.J.

Limau hantu and limau purut: the story of lime-leaves (Citrus hystrix DC.,

UL SAS ices oe eet reine a JB ee ae Aa em Nhe em 3. EN SnR n er O 185

| Rustiami, H.

Two new species of Daemonorops (Palmae) from Sumatra ....1.........cccccecccsesescsepecesentees 199

Kiew, R.

Wan yerd OAOTAIG (AMACATOLACCAE) IS AY DIIG .-2..100.0..0..-c-.0-cagenciecstepeensescesnessacennectesicesies 205

Ashton, P. S.

Laxonemionotes onsthe tree flora ot Brie Ls sscstherertitis nesta diiccave nas scedeactaeancecedscdenves 207

Argent, G.C.G.

New taxa and combinations in the genus Diplycosia (Ericaceae) of

oor cre eA SUAT IMI ANA SIA. cco. c0.0e-s0stxcadescnessvnden Vonstasuossenesessansncsacssereaescsroaenevansnseeks ZA

Burtt, B.L.

New Gesneriaceae: a Chirita from Vietnam and a Monophyllaea from Sulawesi ....... 239

Kiew, R. )

A new species of Monophyllaea (Gesneriaceae) from Borneo ............csseseesseeeseeeeeeees 243

Saw, L.G.

A new species of Vatica (Dipterocarpaceae) from Peninsular Malaysia ................0 247

Mabberley, D.J.

The identity of Swietenia alternifolia (Mill.) Steud. (‘“Meliaceae’) ...........ccceseeeseesseeeeeees 253

Lee, C. C.

Nepenthes platychila (Nepenthaceae), a new species of pitcher plant from

Sarawak, Bored ici. xaicabsxscieesscteans es saseoereea paneer eee 257.

Chung, R.C.K.

Leaf epidermal micromorphology of Grewia L. and Microcos L. (Tiliaceae) in

Penmsular Malaysia and Borne svcsescoss.cscs nesses Absdtacdee has ee acnctine-tuoc pe cente sean aa 263

Vermeulen, J.J. |

Updates on sections of Bulbophyllum (Orchidaceae) revised in Vermeulen (1993) ...287

BOOK REVIEWS

Free Flora of Sabah and Sarawake VOWS 5. tscjccst--cse-cctccetes cease eee P.S. Ashton —289

A Field Guidete the Rattans of Lao PDR 2.0. ti eee tee eee R. Kiew —293

Date of Publication: 20 December 2002

Published by

National Parks Board

Singapore Botanic Gardens

Cluny Road

Singapore 259569

Printed by Oxford Graphic Printers Pte Ltd

Gardens’ Bulletin Singapore 54 (2002) 165-170.

Coprosma bougainvilleensis (Rubiaceae), A New Species

from Bougainville Island, Papua New Guinea

O.G. GIDEON

Division of Biological Sciences, School of Natural and Physical Sciences,

The University of Papua New Guinea, PO Box 320, University PO,

National Capital District 134, Papua New Guinea

Abstract

With the description of a new species, C. bougainvilleensis Gideon, from Bougainville, the

genus Coprosma (Rubiaceae) is reported for the first time from the Solomons. The species

is described and illustrated. Infrageneric relationships and biogeography are briefly discussed.

Introduction

In May 1988, I had the opportunity to collect on Bougainville and Buka

islands. One of the most interesting collecting sites was Mt Balbi, a dormant

volcano, whose summit at 2,590 m is the highest point on the island of

Bougainville. Amongst the many plants collected from this mountain was

a curious species of Coprosma. The discovery of Coprosma on Bougainville

was exciting because, firstly, Coprosma was not previously known from

the Solomons Group or the Bismarck Archipelago, and secondly, it showed

no resemblance to any of the Coprosma species from New Guinea, plants

that I am familiar with. Examining the collection at LAE, I was able to

establish that it represents a distinct new species, and that it shows affinities

with a Pacific group of Coprosma. This species is in striking contrast to the

New Guinea species, which are generally low spreading shrubs with small,

obscurely veined leaves.

Coprosma bougainvilleensis Gideon sp. nov.

A Coprosma novaehebridae in foliis omnino glabris, stipulis edentatis

eciliatis, inflorescentis terminalis et baccis comparate maioribus (9-10.5

mm longis contra 5 mm longos) differt. Typus: Papua New Guinea -

Bougainville Island: Mt Balbi, Emperor Range, Gideon LAE 78543, 8

May 1988 (LAE, holo; L, UPNG, iso).

166 | Gard. Bull. Singapore 54 (2002)

Figure 1

Erect dioecious shrub, 2—5 m tall, young branchlets tetragonous, glabrous.

Leaves opposite, narrowly elliptic, with 6-8 prominent veins on each side

of the midrib, blade entirely glabrous, 8-13 by 2-3.5 cm, base cuneate,

apex acuminate, acumen 6-15 mm; petiole glabrous, terete, 1-2 cm long.

Stipules triangular, connate with the bases of the petioles, 5-7 mm long,

caudate, acumen 3-4 mm long, base 5-6 mm broad, entirely glabrous.

Inflorescence terminal, in few- to several-flowered cymes, either on the

main axes or on lateral branches of a higher order. Flowers small, pale

greenish. Male flowers: calyx green, glabrous, 4- or 5-lobed, lobes deltoid

or triangular, 0.5—1 mm long; corolla funnel-shaped to campanulate, tube

3-5 mm long, lobes 4 or 5, narrowly ovate, 4-6 by 1.5-4 mm, apex acute;

stamens 4 (or 5), inserted at the base of the corolla tube, filaments filiform,

15-20 mm long, anthers dorsifixed, greatly exserted and dangling, 5v6 mm

long, base sagittate, apex acute, opening with two introrse slits. Female

flowers: calyx similar to that of male flowers; corolla tubular, 6—-7.5 mm

long, entirely glabrous, lobes 4 or 5, narrowly ovate, 3-4 by 0.5-1.5 mm,

apex acute, staminodes absent. Ovary ovoid-globose, 2-locular, ovules

solitary, 1-2.5 by 1-2 mm, glabrous; style with 2 long stigmatic lobes, stylar

part 3-3.5 mm long, stigma lobes greatly exserted, 26-30 mm long,

hispidulous. Fruit a globose drupe, 9-10.5 by 9-10 mm, green, ripening

bright red and crowned by the persistent calyx; pyrenes 2, stony, ellipsoid,

6-7 by 3-4 mm.

Distribution: So far known only from the type locality.

Habitat: A very common plant in moderately open situations in the forest,

along walking tracks and along creek banks. Growing on volcanic ash soils

between 1900 and 2300 m altitude.

Notes: The affinity of this new species appears to lie with the western

Polynesian group “Persicaeifoliae” (subgenus Lucidae C.J. Webb), which

includes three species, each endemic to a single island, Samoa, Fiji or

Vanuatu. The group is characterised by lanceolate leaves with pilose veins,

triangular, ciliate, dentate stipules, and shortly branched peduncles. Among

these, it most nearly resembles the Vanuatu species, Coprosma

novaehebridae W.R.B. Oliv., which has a sparsely pilose lamina and pilose

petiole; ciliate, dentate stipules; and a 5-mm long fruit. (Oliver was unable

to describe the flowers as none was available). In contrast, C.

bougainvilleensis is entirely glabrous, the stipules are neither ciliate nor

dentate, and the fruits are 9-10.5 mm long.

Coprosma bougainvilleensis 167

Figure 1. Coprosma bougainvilleensis Gideon

A - female flowering twig; B - Fruits; C - Male flower; D - stipule.

168 Gard. Bull. Singapore 54 (2002)

Although C. bougainvilleensis most resembles Coprosma in tribe

Anthospermeae, it is anomalous in having terminal cymes; all the rest of

the genus have axillary inflorescences and inflorescence position is generally

quite stable in tribe Anthospermeae. Its generic position is therefore

regarded as provisional until the Australasian members of the

Anthospermeae are better understood. Molecular data may prove useful in

shedding light on its affinities.

Other specimens examined: Mt Balbi, Emperor Range, Bougainville Island,

Papua New Guinea. Gideon LAE 78596 (LAE); Gideon LAE 78597 (LAE).

(Duplicates of these collections will shortly be distributed from LAE).

Taxonomic and Biogeographic Notes

Coprosma J.R. & G. Forst. is a large, mainly southern temperate genus of

about 90 species (Darwin, 1979), recently expanded to include Nertera

Banks & Sol. ex Gaertn. and Leptostigma Arn., bringing the total to 128

species (Heads, 1996). The genus is centred in New Zealand, and has a

secondary centre of diversity in the Hawaiian Islands. Coprosma is placed

within the south temperate amphi-pacific subtribe Coprosminae Puff in

tribe Anthospermeae Cham. & Schltdl. The Anthospermeae are

characterised by their unisexual, anemophilous flowers, undoubtedly a

derived condition in the predominantly zoophilous Rubiaceae. The stigmatic

lobes are usually long and filiform, and are completely covered with

hispidulous hairs, significantly increasing their receptive surfaces. They are

usually longer than the style.

In his treatment of the New Zealand Coprosma, Hooker f. (1864)

recognised two main groups, based on whether the female flowers are

clustered or solitary, a distinction generally accepted by later workers at

subgeneric level (Oliver, 1935; Allan, 1961; Heads, 1996). Oliver, the only

monographer of the genus, recognised seven informal groups. Heads (1996)

reduced the genera Nertera and Leptostigma to sections of Coprosma, and

somewhat modified the subgeneric concepts to accommodate the changes.

Coprosma bougainvilleensis differs from all New Guinea species of

Coprosma by its generally larger stature and larger leaves (8-13 by 2-3.5

cm) with 6-8 prominent lateral veins. The mainland New Guinea species

are usually low spreading shrubs with small (3-35 by 1-10 mm), obscurely

veined, needle-tipped leaves, and all occur in the alpine regions. Heads

(1996) placed most of the New Guinea species in the informal group

“Malesia” within the subgenus Coprosma.

Heads (1996) provided an in-depth discussion of the evolution and

Coprosma bougainvilleensis 169

biogeography of Coprosma, and interested readers are referred to that

paper. Prior to the inclusion of Nertera and Leptostigma in Coprosma, the

two clearly distinct groups (subgenera) were western (subgenus Coprosma)

or eastern (subgenus Lucidae) in distributional range, with both groups

overlapping in New Zealand and Hawaii (see Heads, 1996, Fig.1), both

important centres of species diversity for the genus. The evolution and

biogeography of the Coprosminae remains controversial. Puff (1986)

suggested that the present distribution and diversity of the Coprosminae in

the southern regions was a result of long distance dispersal from the north

followed by secondary speciation, while Heads (1996) argued that the

present distribution of Coprosma is a result of vicariance rather than

dispersal. We must look to molecular biology to provide a more precise

estimation of the geographic origin and the evolutionary relationships of

the Coprosminae. The intrusion of the subgenus Lucidae into Papuasia is

probably best explained by (short distance) dispersal, and birds are the

most likely candidates as dispersers.

Acknowledgements

I wish to thank Dr J.F. Veldkamp for commenting on an earlier draft of

this paper and also for translating the Latin diagnosis, and to an anonymous

reviewer for useful comments. R. Kiapranis (LAE) and Dr M. Heads of

the University of Goroka (Papua New Guinea) are thanked for their help

and encouragement, especially the latter for persuading me to publish the

new species, as are Simeon Obedi and Yakas Lelean, former staff of LAE,

who accompanied me on the plant collecting expedition.

References

Allan, H.H. 1961. Rubiaceae. Flora New Zealand. 1: 559-593.

Darwin, S.P. 1979. A synopsis of the indigenous genera of Pacific Rubiaceae.

Allertonia 2: 1-44.

Heads, M. 1996. Biogeography, taxonomy and evolution in the Pacific

genus Coprosma (Rubiaceae). Candollea. 51: 381-405.

Hooker, J.D. 1864. Rubiaceae. Handbook New Zealand Flora. pp. 110-

122.

Oliver, W.R.B. 1935. The genus Coprosma. Bernice P. Bishop Museum

Bulletin. 132: 1-237.

170 Gard. Bull. Singapore 54 (2002)

Puff, C. 1986. A biosystematic study of the African and Madagascan

Rubiaceae Anthospermeae. Plant Systematics & Evolution, Suppl. 3: 1-

585:

Gardens’ Bulletin Singapore 54 (2002) 171-178.

Two Remarkable New West Malesian Homalomena

(Araceae) Species

A. HAY AND C. HERSCOVITCH

Royal Botanic Gardens Sydney,

Mrs Macquaries Road,

Sydney 2000, Australia

Abstract

Two very distinctive new species of Homalomena Schott (Araceae-Homalomeneae) are

described from Sumatera (H. elegantula A. Hay & Herscovitch) and Borneo (H. expedita

A. Hay & Herscovitch). Both are illustrated.

Introduction

Homalomena Schott is a genus of about 100 species of terrestrial or

lithophytic forest-dwelling herbs. Most are Malesian, with a few in

continental SE Asia and the neotropics. The genus is most closely allied to

and possibly congeneric with Furtadoa M. Hotta (West Malesia), the two

genera forming the tribe Homalomeneae. Furtadoa is distinguished by its

monandrous male ‘flowers’ with the stamens aligned in a regularly

transverse pattern (in relation to the spadix axis) and with a pistillode

associated with each (versus at least diandrous male flowers without

pistillodes in Homalomena). However, Sumateran H. monandra M. Hotta

is intermediate in having monandrous male ‘flowers’ in the same

configuration as those of Furtadoa but without associated pistillodes. The

tribe is currently considered most closely allied to neotropical Philodendreae

(Mayo et al., 1997).

Homalomena has been recently revised for New Guinea (Hay, 1999) and

Java (Yuzammi, 2000). Progress with revising the genus for the rest of the

Flora Malesiana region has unfortunately slowed, however, and therefore

we are publishing the more distinctive taxa as they come to light.

1. Homalomena elegantula A. Hay & Herscovitch, sp. nov.

A Homalomena bellula Schott habitu repenti, foliis distantibus, petiolo ad

apicem geniculato, lamina anguste elliptica longe cuspidata, inflorescentia

valde minore differt. Typus: Indonesia, Sumatera, West Sumatera, East

of Pajakumbuh, Taram, sandstone region of River Tjampo, 24 Aug 1957,

W. Meijer 6928 (L, holo).

12 Gard. Bull. Singapore 54 (2002)

Figure 1

Slender creeping herb to c. 25 cm tall. Rhizome very slender, elongate,

creeping, c. 5 mm diam. Leaves few, distant, about 5 cm apart on the

rhizome, subtended and enclosed by cataphylls in bud; petiole 15-22 cm

long, erect, apically sharply bent holding the blade at right angles, geniculate

at the junction with blade, sheathing only at the extreme base; blade

narrowly elliptic, slightly falcate, 17-20 cm long, c. 3 cm wide, the base

acute, the tip cuspidate-acuminate for c. 4 cm and apiculate for c. 2 mm,

drying greyish adaxially, reddish brown abaxially; midrib slender and very

prominent abaxially, impressed adaxially, with 6-8 primary lateral veins on

each side; primary lateral veins very weakly differentiated from the

secondary venation, diverging at c. 30-45° and somewhat prominent

abaxially and adaxially. Inflorescences several together, subtended by

sublinear cataphylls to c. 3 cm long; peduncle slender, to 4 cm long, declinate

after anthesis. Spathe unconstricted, spindle-shaped, c. 1 cm long, c. 2.5

mm thick (before anthesis), c. 1.2 cm long, c. 5 mm thick (after anthesis).

Spadix very shortly stipitate for c. 0.5 mm; female zone 2 mm long, c. 1.5

mm diam.; pistils crowded; ovary subglobose-ovoid, drying rust-brown, c.

0.3 mm diam.; stigma button-like, somewhat narrower than the ovary, raised

on a short style; infra-pistillar staminodes somewhat shorter than the ovary,

clavate on a very short stipe, c. 0.2 mm diam., ivory; male zone contiguous

with the female, cylindric, distally tapering to a point, drying ivory yellow;

male flowers 2-staminate; stamens truncate, c. 0.5 mm across; thecae opening

through large subapical pores; connective slightly elevated beside and

between the pores. Fruiting spathe broadly spindle-shaped.

Distribution: Malesia: endemic to Sumatera (West Sumatera; known only

from the type).

Habitat: Terrestrial in hill forest over sandstone, 500—1000 m alt.

Notes: Homalomena elegantula, though its minute inflorescence closely

resembles those of H. humilis (Jack) Hook.f and allies, is distinguished

from that group by its remarkable habit. Shoot organisation appears

comparable to that of H. hastata M.Hotta and H. bellula Schott, with

flowering being both functionally and anatomically terminal and renewal

taking place by the release of a dormant lateral bud. The architecture of

the rhizome thus resembles that of many gingers and most Schismatoglottis

species allied to S. calyptrata (Roxb.) Zoll. & Moritzi (Hay & Yuzammi,

2000). This organisation is in marked contrast to that of other Homalomena

species where the shoot consists of a physiognomically unbranched

Two New Homalomena Species L73

desley

ra har1.0/

Figure 1. Homalomena elegantula A. Hay & Herscovitch

W. Meijer 6928 — A. habit; B. leaf venation; C. detail of vegetative tip of rhizome showing leaf

base and cataphylls; D. rhizome with synflorescence; E. inflorescence with part of spathe

removed; F. pistils and staminodes; G. stamens.

Scale: bar to A = 12 cm, to B = 2.7 cm, to C=3 cm, to D= 6 cm, to E= 7 mm, to F & G=1.5 mm.

174 Gard. Bull. Singapore 54 (2002)

sympodium typical of most herbaceous Araceae. It has not previously

been recorded in this genus. The leaves of H. elegantula are preceded and

enclosed by a series of cataphylls and evidently the bud containing the

next foliage leaf is extended some distance from the previous one giving

the plant its long creeping habit. However, the exact architectural

organisation is impossible to observe from the single dried specimen known

to us. The elegant leaves (to which the epithet refers) are remarkable for

appearing to be held, through the action of a geniculum, such that the

blades all face the same way (and probably held tip down, but this cannot

be ascertained from the specimen as it cannot be orientated with

confidence), and for their elongate cuspidate tips.

2. Homalomena expedita A. Hay & Herscovitch, sp. nov.

Ab aliis speciebus Homalomenae in habitu stolonifera, heliophila,

paludicola, inflorescentia solitaria, spadice cum interstitio nudo longo

verruculoso differt. — Typus: Cult. RBG Sydney Acc. No. 940562 ex

Malaysia, Sarawak, Lundu, near bridge on Kuching Road (orig. coll. Hay,

Yahud, Saupel & Chan 9409), C. Herscovitch s.n. (NSW, holo; iso SAR).

Figures 2 & 3

Colony-forming stoloniferous herb to c. 60 cm tall. Stem an erect to creeping

rhizome to c. 30 cm long, c. 4 cm thick emitting cataphylliferous stolons to

c. 40 cm long, 1 cm thick, these eventually upturned, becoming rhizomatous,

leafy, emitting further stolons from the base of the rhizomatous portion.

Leaves clustered, to c. 10 together; petiole to c. 45 cm long, somewhat

spongy within, pale mid-green with broken darker green longitudinal

striations, sheathing in the lower 1/3, with an anise odour when crushed;

blade mid-green on both sides, glossy at first, becoming matt, very broadly

ovato-sagittate, leathery, c. 15-20 cm long and wide, the apex broadly

obtuse, very abruptly and shortly acuminate for c. 1 cm, finally stiffly

apiculate for c. 3 mm, the base shallowly cordate to almost truncate, usually

distinctly asymmetric, with widely spreading rounded to subtriangular

posterior lobes 7-10 cm long; midrib adaxially flat, abaxially slightly

prominent, with c. 5 adaxially impressed, abaxially slightly prominent

primary lateral veins on each side (plus a cluster of 2 or 3 on each side

running to the posterior lobes), alternating with fainter interprimaries and

diverging at c. 60°. Inflorescence solitary; peduncle c. 7 cm long, rather

thick, c. 8 mm diam., erect. Spathe green, c. 5 cm long, 1.5 cm across and in

bud slightly inflated at level of female zone, thence subcylindric-tapering,

but not constricted, later very narrowly ovoid, apiculate for c. 4mm. Spadix

stipitate for 6 mm, 4.5 cm long; female zone more or less cylindric, 0.8—1.2

cm long (irregular length around circumference of spadix), 1.2 cm wide;

Two New Homalomena Species ee

Lese

£ kan Y L00l

Figure 2. Homalomena expedita A. Hay & Herscovitch

Herscovitch s.n. — A. young shoot with runner; B. leaf blade outline; C. flowering shoot detail;

D. inflorescence.

scaleibarito A.B = 12 cn.ito.C, D=7.5'cm.

176 Gard. Bull. Singapore 54 (2002)

pistils subglobose, 1-2 mm diam.; stigma subsessile, discoid or very weakly

2—3-lobed, slightly narrower than the ovary, papillate; infrapistillar

staminodes absent; sterile interstice conspicuous, 1.5 cm long, naked, pale

green and c. 6 mm diam. in the lower 1 cm, with more or less regular

spirals of low domed warts c. 0.5 mm diam., the upper part (comprising the

base of the male zone) 9 mm diam., ivory and clothed in irregular sterile

stamens; fertile male zone elongate-bullet-shaped, 1.5 cm long, tapering to

a blunt acute tip, ivory; male flowers irregular (1—)2—4-staminate; stamens

truncate, irregular in size, 1-1.5 mm across, irregularly polygonal, slightly

sinuous on the abaxial side, the thecae overtopped by connective. Fruit

unknown.

Distribution: Malesia: endemic to Borneo (Sarawak); known from the type

locality and sighted near Sibu along the main road to Miri.

Habitat: Forming large, probably clonal colonies in open swamps and

ditches at low elevation, sometimes in tidal mud with Cryptocoryne ciliata

(Roxb.) Schott and mangroves.

Notes: The spadix of Homalomena expedita is unique in the genus, having

a very conspicuous naked warty interstice above the female zone as well as

a zone of sterile stamens at the base of the male zone. Homalomena expedita

is also very remarkable, in this genus of shade-loving terrestrial and

rheophytic plants, for its occupation of open swampy sites, even tidal

mudflats, in full sun. It appears to spread rapidly through the production

of stolons, and is certainly very vigorous (though seldom-flowering) in

cultivation when provided with large amounts of nutrients. At Lundu it

forms dense swards in shallow muddy ponds and from a distance rather

resembles water hyacinth (Eichhornia). It appears less vigorous, though

nonetheless invasive and persistent, in tidal, brackish conditions.

This species, in relation to the rest of its genus, is rather analogous to

the open swamp-dwelling, colony-forming Aglaodorum, contrasted with

Aglaonema (both Aglaonemateae). Both Homalomena expedita and

Aglaodorum griffithii Schott, besides their similar habitat preferences,

produce solitary inflorescences where their immediate forest-dwelling, non-

proliferating relatives generally produce complex synflorescences (except

in some of the most diminutive species).

In addition to the reduced number of inflorescences, it seems, on the

basis of limited observations of H. expedita in cultivation, that there is

other evidence pointing to depressed or suppressed sexual fertility: we

have never observed the spathe to open (but we are at present unable to

Two New Homalomena Species ty?

iSals cA Sem

eneee

3 ee

C2)

Figure 3. Homalomena expedita A. Hay & Herscovitch

Herscovitch s.n. — A. habit of cultivated plant; B. spadix; C. pistils; D. abortive pistils on interstice;

E. sterile stamens from lower part of male zone; F. stamens from upper part of male zone.

Scale: bar to A = 18 cm, to B=2cm,to C,D & E=6 mm.

178 Gard. Bull. Singapore 54 (2002)

verify that it remains closed throughout the flowering process); the papillae

on the interstice appear to be abortive pistils; the staminal thecae contain

some pollen, but are partially empty in most anthers; senescent male flowers

appear not to have shed pollen; the ovaries however, are filled with ovules

which appear normal in form.

The leaf, very broadly ovate with a near-truncate cordate base, is

similar to that of H. havilandii Ridl., also from Sarawak, but that species

does not share the habit and habitat preference of H. expedita nor the

prominent sterile interstice of the spadix.

The epithet means ‘set free’, or ‘foot-loose’, alluding to the departure

from confinement to shaded conditions otherwise typical for the genus,

and the invasive, stoloniferous habit.

Acknowledgements

A.H. thanks Yahud bin Hj. Wat, Saupel B. Atot and Chang Ban Sem

(Sarawak Forestry Department) for their kind assistance in the field in

1994, and Conrad D. Fleming for generous financial support of field work.

We also thank various Royal Botanic Gardens Sydney nursery staff, most

recently Phil Pettitt, for maintaining the living collection of Araceae over

the years and for expertly bringing Homalomena expedita into flower.

Thanks are also due to the Riksherbarium for the loan of herbarium

specimens among which we found H. elegantula. Last but not least we

thank Lesley Elkan for her skillful botanical illustrations.

References

Hay, A. 1999. Revision of Homalomena (Araceae-Homalomeneae) in

New Guinea, the Bismarck Archipelago and Solomon Islands. Blumea

44: 41-71.

Hay, A. and Yuzammi 2000. Schismatoglottideae (Araceae) in Malesia I —

Schismatoglottis. Telopea 9: 1-177.

Mayo, S.J., J. Bogner and P.C. Boyce 1997. The Genera of Araceae. Kew.

Yuzammi, 2000. A Taxonomic Study of the Terrestrial Aroids of Java.

Unpublished MSc Thesis, The University of New South Wales, Australia.

Gardens’ Bulletin Singapore 54 (2002) 179-183.

Re-examination of Vaccinium dialypetalum (Ericaceae)

S.P. VANDER KLOET

Department of Biology, Acadia University, Wolfville

Nova Scotia BOP 1X0, Canada

Abstract

Morphological re-examination of Vaccinium dialypetalum sensu lato supports the view that

it includes three distinct taxa: V. dialypetalum J.J.Sm., V. micranthum (Ridl.) Vander Kloet

and V. perakensis (Ridl.) Vander Kloet, all first described by Ridley under Agapetes but

transferred by Sleumer to Vaccinium. Keys and descriptions are provided to distinguish

the three taxa and new combinations are made for two taxa.

Introduction

Ridley (1923) recognized three species of Agapetes (Ericaceae) based on

their distinct floral development patterns. In all three species, floral axis

development is sequential. First, the rachis emerges from the perennating

bud, followed by the development of the pedicel and calyx tube at the first

node (indicated by a tiny bract on the rachis) and finally the corolla, style

and stamens are initiated. Thus in A. perakensis Ridl., the rachis and

pedicels continue to lengthen and thicken in tandem until anthesis when

rachis growth stops but the pedicel continues to elongate until the fruit

ripens, producing a rachis 3—5 cm long with many pedicels 1-2 cm long. In

A. micrantha Ridl., the development of the rachis is arrested but the pedicels

continue to lengthen and thicken until the berry is ripe, resulting in a

slender, 3-flowered rachis but A. wrayii Ridl., has a short stout rachis

bearing many flowers on long pedicels that continue to elongate as the

fruit matures.

Sleumer (1967) did not recognize these differences as significant and

united them into a single species, Vaccinium dialypetalyum J.J. Sm. While

Sleumer’s transfer of Ridley’s three Agapetes species to Vaccinium §

Galeopetalum (J.J.Sm.) Sleumer is widely accepted, his broad species

concept of V. dialypetalum was called into question when Ng (1976)

separated specimens with shorter calyx lobes and stamens with glabrous

filaments and short dorsal spurs as a new species, V. pseudodialypetalum.

This latter species is, however, identical with Ridley’s A gapetes micrantha.

Indeed, to place all three taxa in V. dialypetalum is quite untenable

since at least two qualitative features separate each taxon, viz., corolla

shape and stamen architecture separate V. micrantha from V. dialypetalum

180 | Gard. Bull. Singapore 54 (2002)

and V. perakensis, whilst anther fusion and the shape of the staminal awns

separate V. perakensis from V. dialypetalum. This morphological gap is

sufficient to recognize each of Ridley’s taxa at the specific level.

Taxonomic Treatment

Vaccinium § Galeopetalum (J.J.Sm.) Sleumer, Notizbl. Berlin-Dahl. 13

(1936) 115. Subgenus Galeopetalum, J.J.sm. Icones Bogoriensis 4 (1912)

101. Type: Vaccinium dialypetalum J.J.Sm.

In Malesia, according to Sleumer, this section is restricted to northern

Sumatra and adjacent Peninsular Malaysia. Species, such as V. petelotii

Merr., V. lanigerum Sleumer, and V. dunalianum Wight from northern

Vietnam, adjacent Yunnan and the Tibet-Burma border, have also been

referred to this section by Stevens (1969). However, Vander Kloet and

Paterson (2000) found that V. petelottii is genetically quite similar to V.

tonkinense Dop of sect. Conchophyllum Sleumer, hence Stevens’ extension

of sect. Galeopetalum into SE Asia needs to be reassessed.

Keys to Identify Malesian Taxa of Vaccinium § Galeopetalum

Key for flowering material

Corollas globose; calyx lobes shorter than the tube; stamens c. 1 mm long

ML. BOOS SE eed, SURI, Ce Pe BO ee Ee, BO wesseeee 2. V. micrantha

Corollas campanulate; calyx lobes much longer than the tube; stamens 3-4

mm long

Antherstiubed:,.2oc..cs Aer... se Bees 1. V. dialypetalum

Arithers test. 20. Aten . 2A ae ee 3. V. perakensis

Key for fruiting material

Rachis much shorter tian the Peqicels «.:...--ctc. cn erewe 1. V. dialypetalum

Rachis as long as or much longer than the pedicels

Rachis 1—2 cm long, fruits per raceme 3 or fewer ...2. V. micrantha

Rachis 3—10 cm long, fruits per raceme 4 or more . 3. V. perakensis

Vaccinium dialypetalum 181

1. Vaccinium dialypetalum J.J.Sm. Icones Bogoriensis 4 (1912) 99, t. 331.

Type: Indonesia, Java (cult.) Rant & Smith 513_in 1911. (holo, BO).

Synonyms: Agapetes pubescens Ridl., J. Bot. 62 (1924) 298. Type: Burkill

& Holttum 7828 (SING); A. wrayii Ridl., Fl. Malay Pen. 2 (1923) 205.

Type: Wray 8054 (SING);

Vaccinium longipes Sleumer, Bot. Jahrb. 71 (1941) 424. Type: not seen.

Epiphytic shrub with a well-developed tuber; stems 2 or 3, sinuous, 1-3 m

long; innovations initially reddish and pubescent; leaves ovate-lanceolate,

narrowly acuminate, 6-12 x 1.5-4.5 cm, margin entire, petioles 1-2 mm

long; racemes axillary, 6—10-flowered; rachis much shorter than the pedicels;

pedicels 2-4 cm long articulated with the calyx tube; calyx lobes longer

than the tube at anthesis; corolla campanulate, whitish-green, 6 mm long, 7

mm in diameter, 5-lobed, lobes 4-5 mm long; stamens 7 mm long, anthers

fused into ring, awns and tubules well developed; style glabrous, slender 6—

7 mm long; berry black, 7-9 mm diam., seeds many and small.

Distribution and ecology: Northern Sumatra and Peninsular Malaysia (Main

Range), in montane forests 900-1300 m altitude.

2. Vaccinium micranthum (Ridl.) Vander Kloet, comb. nov.

Type: Malaysia, Selangor, Sempang Mines, Ridley 15768 (1911). (holo

SING!).

Basionym: Agapetes micrantha Ridl., Fl. Malay Pen. 2 (1923) 205.

Synonyms: A. parviflora sensu Ridl., J. Roy. As. Soc. S. Br. 61 (1912) 26

non A. parviflora Dunn; Vaccinium ridleyii Sleumer, Bot. Jahr. 71 (1941)

424. Type: Ridley 15768 (SING); V. pseudodialypetalum Ng, Gard. Bull.

Sing. 28 (1976) 233. Type: KEP 56673.

non Vaccinium parviflorum Andrews, Bot Rep. (1800) t. 125 = Gaylussacia

resinosa (Aiton) T & G.

Epiphytic shrub with a well-developed tuber; stems 2 or 3, slender, sinuous

1-2 m long; innovations initially reddish and softly pubescent; leaves deltoid-

ovate, long caudate, 6-8 x 3-4.2 cm, margin entire, petioles 1-2 mm long;

racemes slender, 3-flowered, often obscured by the leaves; rachis 1-2 cm

long; pedicels 1-2 cm long, calyx lobes shorter than the tube; corolla globose,

greenish-white, 3 mm long, 3-4 mm diam., 5-lobed; stamens | mm long,

anther tubules and awns vestigial; style stout, glabrous, cylindrical, 3 mm

long;berry black, 8-9 mm diam.; seeds small, 25—40; testa pale brown.

182 Gard. Bull. Singapore 54 (2002)

Distribution and ecology: Known only from Peninsular Malaysia (Pahang

and Selangor), in montane forests between 1000-1600 m altitude.

3. Vaccinium perakensis (Ridl.) Vander Kloet, comb. nov.

Type: Malaysia, Perak, Thaiping, Maxwell’s Hill, 750-1200 m altitude.

Ridley 5532 (holo SING!).

Basionym: Agapetes perakensis Ridl., Fl. Malay Pen. 2 (1923) 205.

Synonyms: A. griffithii sensu K&G, J. As. Soc. Beng. 74 (1905) 59 non

C.B. Clarke in Hook.f. Type: King 6363 (SING); Vaccinium longilingue

Sleumer, Bot. Jahr. 71 (1941) 424. Type: Ridley 5532 (SING); V.

urophyllum Merr., Pap. Mich Ac. Sc. 19 (1934) 184. Type: King 6363 (iso

SING).

Epiphytic shrub with a well-developed tuber; stems few, slender and sinuous,

up to 5 m long; innovations reddish and pubescent, rarely glandular; leaves

ovate-lanceolate and often caudate, 8-14 x 2.2-5.5 cm wide, margin entire,

petioles 2-3 mm long; racemes axillary, well-developed, glabrous, pubescent

or glandular, 5—15 flowered; rachis invariably longer than the pedicels;

pedicels 1-3 cm long, calyx lobes as long as or longer than the tube at

anthesis; corolla campanulate, slightly 5-angled, creamy white often pink

inside, 5—6 mm long, obtusely 4 or 5 lobed; stamens c. 6 mm long, anthers

free, with 2 dorsal spreading and ultimately upwardly curved echinate

arms, tubules slender and erect; style slender and glabrous, 6-7 mm long;

berry reddish-black, often pubescent, 7-9 mm diam., seeds small, testa

blackish.

Distribution and ecology: Northern Sumatra and Peninsular Malaysia,

where it is uncommon, in montane forest, or riverine at lower elevations,

600-1200 m altitude.

References

Ng, F.S.P. 1976. Notes on the Systematy of Malayan Phanerogams. XX1V

Ericaceae. Gardens Bulletin Singapore 28: 231-234.

Ridley, H.N. 1912. New and rare Malayan Plants. Journal Royal Asiatic

Society Straits Branch. 61:19-31.

Ridley, H.N. 1923. Ericaceae. Flora of the Malay Peninsula. 2: 211-222.

Sleumer, H. 1967. Vaccinium (Ericaceae). Flora Malesiana. 6: 746-878.

Vaccinium dialypetalum 183

Stevens, P.F.1969. Taxonomic Studies in the Ericaceae. Unpublished D.Phil.

thesis, University of Edinburgh, U.K.

Vander Kloet, S.P. and I.G. Paterson. 2000. RAPD assessment of novelties

resulting in a new species of Vaccinium L. (Ericaceae) from Vietnam.

Botanical Journal Linnean Society. 134: 575-586.

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Gardens’ Bulletin Singapore 54 (2002) 185-197.

Limau Hantu and Limau Purut: the Story of Lime-Leaves

(Citrus hystrix DC., Rutaceae)?

D. J. MABBERLEY

Nationaal Herbarium Nederland, University of Leiden, The Netherlands;

Royal Botanic Gardens Sydney,

Mrs Macquaries Road, Sydney 2000, Australia*

Abstract

Limau purut (Citrus hystrix DC.), cultivated throughout SE Asia, appears to be a selected

form of the wild limau hantu (C. macroptera Montr., i.e. C. auraria Michel), though its

earliest scientific name may be C. fusca Lour. Complete synonymy with types is presented

in a provisional arrangement of ‘wild’ plants and cultivars. Suggestions for further work on

C. hystrix and its relations with other cultivated citrus are made. X Citroncirus is formally

reduced to Citrus and a new name proposed for the citrange root-stock, Citrus x insitorum

Mabb. A diagram of the relationships through hybridity of cultivated citrus is presented.

Introduction

Characteristic of Thai cooking, worldwide, are lime-leaves (Jimau purut,

Citrus hystrix DC.), chopped fine better to release their oils. The fruits are

not used for food, because, unlike those of species and hybrids placed in

‘subg. Citrus’, those of C. hystrix and other species placed in ‘subgen.

Papeda (Hassk.) Swingle’ are almost inedible due to the acrid oil in the

vesicles surrounding the seeds (Mabberley, 1997). They have been used

medicinally, and in Sri Lanka the English name is leech-lime because they

are used as a leech-repellent. In the Malay Peninsula the fruits were a soap

substitute and sold for this purpose (Burkill, 1931), a practice still prevalent

in Cambodia (Boeun Sok, Royal Botanic Gardens Sydney, pers. comm.)

and elsewhere in SE Asia.

Limau purut and limau hantu

A plant of Jimau purut grown in Ambon, Indonesia, was drawn for

Rumphius (1628—1702) and specimens were collected 1771—2 by Pierre

Sonnerat (1748—1814) in the ‘Indes’ and noted some decades later in

Lamarck’s Encyclopédie méthodique (1796). It was first grown in Europe

at the end of the eighteenth century when it was introduced to France, by

an amateur, from Mauritius, where it was being cultivated. A fruit was

given to a M. Rollljand, through whom a plant reached the botanic garden

* Address for correspondence

186 Gard. Bull. Singapore 54 (2002)

in Montpellier, France, in 1808. However, it did not flower for many years

and it was formally described in the sterile state in 1813 (as ‘C. histrix’). By

1816 it was in the trade, being grown by the Audibert nursery at Tonelle,

near Tarascon (Michel, 1816: 42, t. 18).

In cultivation in Singapore and elsewhere today are forms differing

in the amount of toothing of the lamina and the degree of roundedness of

its apex, but these features can vary markedly within a single cultivated

plant (pers. obs.). The pericarp of the fruit is generally somewhat lumpy in

appearance. A form with a particularly lumpy pericarp is known as var.

torosa (Blanco) Merr. and was also described from a cultivated plant,

grown in the Philippines.

In recent accounts, these cultivated plants have been considered allied

to, but distinct from, C. macroptera Montr., described from New Caledonia

though not native there. Citrus macroptera, which was also figured by

Rumphius (the plate being the basis of C. auraria Michel, an apparently

overlooked earlier name* for C. macroptera - see below), is found

throughout Malesia, and in Peninsular Malaysia it is known as Jimau hantu.

On the whole, such plants tend to have rather more acute leaf apices.

Having examined a wide range of material of /imau hantu and limau purut

held at BM, F, K, L, NSW, P, PNH, SING and UC, US, however, I am

forced to conclude that the cultivated plants called /imau purut appear to

be merely selected forms (cultivars) of limau hantu.

This conclusion is not original: Merrill (1923), studying just the

Philippine material available to him, amalgamated the ‘species’, though he

used varietal rather than cultivar status for the pomological forms.

Moreover, the ‘wild’ plants throughout their range were formally included

in C. hystrix by Engler and Harms (in Engl., Nat. Pflanzenfam. Ed. 2, 19a:

336, 1931), but these authors have not generally been followed in recent

floristic accounts.

The state of affairs is somewhat comparable with that of the cultivar

‘Etrog’ of Citrus medica L., the citron. That cultivar was first formally

described as C. tuberosus Mill. and has a markedly tuberculate berry surface

like Jimau purut, compared with the smoother-skinned ‘typical’ forms of C.

medica similar to those in Jimau hantu. Many forms of citrus with lumpy

or grotesquely formed fruits were also selected in Europe, notably in Italy,

and were fashionable in the orangeries of the rich where they were known

as ‘agrumi’.

Another ancient cultivar of C. medica, for example, is ‘Fingered’

(the ‘Buddha’s hand’), where the fruit is split into a number of finger-like

sections. Because of its form, in the past it has been afforded not merely

varietal (var. sarcodactylis (Noot.) Swingle) and specific (C. sarcodactylis

*As are his Citrus ventricosa (see below) and Citrus mammosa and C. nipis Michel, Traité

Citronier: 44 (1816) and in Duhamel, Traité Arbres Arbustes 7:111 (1819)=C.medica L. &

C. x aurantiifolia (Christm.) Swingle, respectively.

Limau Hantu and Limau Purut 187

Noot.) ranks, but even accommodated in a separate genus (as Sarcodactilis

helicteroides Gaertn.f.)!

A Provisional Classification for Wild and Cultivated Forms of

Citrus hystrix

Note: names not found in Index kewensis or other lists are marked with an

asterisk.

Citrus hystrix DC., Cat. Hort. Monsp.: 19, 97 (‘histrix’, 1813); Michel, Traité

Citronier: 42, t. 18 (1816) & in Duhamel, Traité Arbres Arbustes 7:108, t.

39 fig 1 (‘histrix’, 1819); DC., Prodr. 1: 539 (1824); Merr., Enum. Phil. FI.

Pl. 2: 342 (1923) excl. var. macrophylla (Wester) Merr. [cf. C. maxima

(Burm.) Merr. or C. x aurantium L. Grapefruit group]; Guillaumin in

Gagnep., Suppl. Fl. Gén. Indochine: 653 (1946); Staples & Kristiansen

(1999: 27-9), q.v. for description. - C. echinata St-Lag. in Ann. Soc. Bot.

Lyon 7: 122 (1880), nom. superfl.

Type: cultivated in Montpellier, France, “Frutex spectabilis [N.B. It was

sterile when first described and had still not flowered by 1824 (DC., l.c.)]

olim ex insula Mauritiana (ubi forsan cultus) merit. Mercatori Nemauensi

[a merchant of Nimes, France] Roland a navarcha quodam allatus, et anno

1808 a D®. Roland horto Monspeliensi humanissime missus” (G-DC fiche!

‘de M. Roland’, holo?).

(Northeastern India and southern China?), Burma and Thailand to Sumatra,

east to New Guinea, though natural distribution probably obscured by

cultivation, its having been carried far into the Pacific (A.C. Smith, FI.

Vitiensis Nova 3: 186, 1985), for example. Selected forms are cultivated

throughout the warm parts of the world for culinary (leaves - lime-leaves)

and medicinal (fruit - leech-lime) use. Indeed all named taxa seem to have

been based on cultivated plants, though perhaps the type of C. celebica

and some other Philippine taxa were truly wild ones.

As with Clerodendrum chinense (Osb.) Mabb. (Labiatae), Cupressus

lusitanica L. (Cupressaceae), Kerria japonica (L.) DC. (Rosaceae),

Rhododendron stenopetalum (Hogg) Mabb. (Ericaceae; Mabberley, 1995)

and Melia azedarach L. (Meliaceae; Mabberley, 1984), for example, the

name C. hystrix covers both the wild plant and the cultivar (the type, so far

apparently without a formal cultivar name, though Guillaumin’s name for

it among his ‘formes culturales’ [in Lecomte, Fl. Gén. Indochine 1: 676,

1911, under C. aurantium| could be construed as such, though being

‘Hystrix’, this would not be allowed by the Cultivated Code [Art. 25.2]). It

188 | Gard. Bull. Singapore 54 (2002)

is notable that the scrappy type at G-DC has leaflets rather more acute

than is seen in many cultivated specimens. The only cultivar name published

for any plant referable to this group seems to be ‘copahan’ (see below).

The plants are arranged below as ‘wild’ (Jimau hantu) followed by limau

purut (lime-leaves) and other cultivars.

‘Wild’ plants (limau hantu)

[Lemoen Amas Valentijn, Amboina 3(1): 190 (1726)]

*Citrus auraria Michel, Traité Citronier: 43 (1816) & in Duhamel, Traité

Arbres Arbustes 7:109 (1819). - [Limonellus aurarius Rumph., Herb.

Amb. 2: 109, t. 30 (1741)] - C. limetta Risso var. auraria Risso in

Risso & Poit., Orangers: t. 59 (1818).

Type [icon]: Rumph, Herb. Amb. 2: t. 30 (1741).

Citrus macroptera Montr. in Mém. Acad. Lyon 10: 187 (1860). - C. combara

Raf. var. macroptera (Montr.) Tanaka in J. Soc. Trop. Agric. 10: 352

(1938) & Stud. Citr. 9:2 (1939).

Type: New Caledonia, [le Art “juxta domos indigenarum”, Montrouzier

s.n. (LY, lost [F, photo!]).

Citrus celebica Koord. in Meded. ‘sLands Plant. 19: 370, 839 (1898).

Type: Indonesia, Sulawesi, Minahassa, Menado, 250-275m, Koorders 18751

(BO, holo-, n.v.; K!, P!, PNH!, iso-).

Citrus papuana F.M. Bailey in [Ann.] Rep. Brit. New Guinea 1901-2: 1

(1902) & Contrib. Fl. Brit. New Guinea: 1 + t. (1903).

Type: Papua New Guinea, Milne Bay, Le Hunte s.n. (BRI, n.v. holo?; P,

fragm.!).

Citrus x aurantium L. subsp. saponacea Saff. in Contrib. U.S. Nat. Herb. 9:

226 (1905), e descr.

Type: [Guam, cult.] not indicated.

Citrus southwickii Wester in Phil. Agr. Rev. 8: 16, tt. 3c [‘C. limao’], 4c

(1915). - C. hystrix DC. var. southwickii (Wester) Merr., Enum. Phil.

Fl. Pl. 2: 343 (1923). - C. macroptera Montr. var. southwickii (Wester)

Tanaka in Trans. Nat. Hist. Soc. Formosa 22: 430 (1932). - C. celebica

Koord. var. southwickii (Wester) Swingle in J. Wash. Acad. Sci. 28:

533 (1938).

Limau Hantu and Limau Purut 189

Type: Philippines, cult. Luzon, Lamao, Mar 1915, Wester 2049 (PNHf#,

holo?; K!, PNH!).

Citrus hystrix DC. var. boholensis Wester in Phil. Agr. Rev. 8: 19, tt.. 4a, 5a

(1915). - C. macroptera Montr. var. boholensis (Wester) Tanaka in

Trans. Nat. Hist. Soc. Formosa 22: 430 (1932). - C. combara Raf.

var. boholensis (Wester) Tanaka, Stud. Citr. 9: 3 (1939). - *C.

boholensis (Wester) Tanaka, Syst. Pomol. : 140 (1951), nom.nud.,

synon. nov.

Type: Philippines, Bohol, cult. Lamao, Mar. 1915, Wester 2525 (PNHf,

syn?; PNH!, isosyn.), Bohol, cult., 1914, Wester 4824 (PNHf?, syn?;

PNH! isosyn).

Citrus vitiensis Tanaka in Bull. Soc. Bot. Fr. 75: 715 (1928).

Type: Fiji, Viti Levu, Namoso/Namura R., 1866, Seemann 58 (K!, holo [F,

photo!]; BM!, P!, iso).

Citrus macroptera Montr. var. annamensis Tanaka in Bull. Mus. Hist. Nat.

Paris II, 2: 164 (1930). - C. hystrix DC. var. annamensis (Tanaka)

Guillaumin in Gagnep., Suppl. Fl. Gén. Indochine: 654 (1946) .

Type: Vietnam, massif Co Inh, pres de Nhatrang, 18 Sept. 1922, Poilane

4650 (P!, holo [F, photo]; UC!).

Citrus macroptera Montr. var. kerrii Swingle in J. Wash. Acad. Sci. 32: 34

(1942) - *C. kerrii (Swingle) Tanaka, Syst. Pomol.: 140 (1951), synon.

nov.

Type: Thailand, Nalawn Sawan, Kampeng Pét, Mé Lamung, 7 June 1922,

A.F.G. Kerr 6081 (ABD, holo; BM!, K!, iso)

Rumphius (l.c.) discussed the disarticulating of the lamina from the broad

winged petiole and the use of the fruit in washing, recording that it occurs

in Sulawesi and that it is the Lemoen Amas of Valentijn. It should be

noted that although the plants arranged above resemble morphologically

the presumed wild ancestor of the cultivars below, several of them are

described from plants in cultivation, often far beyond the putative natural

range, and in that sense are perhaps cultivars in their own right. The

concept of ‘wildness’, so obscured in cultivated plants such as these, is

clearly linked to human perceptions of human actions as somehow ‘above’

or ‘different from’ the dispersal activities of other animals’ selecting

‘superior forms in terms of fruit or seed size, colour, taste, texture, etc.

and is therefore perhaps an inappropriate one (Mabberley, 1999).

190 Gard. Bull. Singapore 54 (2002)

N.B. Merrill (1917) suggested that yet another plant described and figured

by Rumphius should be referred to C. hystrix (i.e. Herb. Amb. 2: t. 26 f. 3),

for which the flowers and leaves look right, but not the fruit (cf. C.

ventricosus below, where the opposite is true). Moreover Burkill (1931)

also referred to it, the Indochinese ‘C. cavalierei Léveillé’, a nomen nudum,

and the Chinese C. ichangensis Swingle, of which var. latipes Swingle has

been given specific rank as C. latipes (Swingle) Tanaka.

However, there are a number of more distinctive taxa, all apparently

certainly cultivars, the first now widely spread in the Old World Tropics,

the others grown principally in central Malesia, notably the Philippines.

‘Copahan’ (and unnamed cultivar[s]?) - /imau purut, lime-leaves, makrut

lime, leech lime, Kaffir lime)

Citrus hystrix DC. ‘Copahan’, Wester in Phil. Agric. Rev. 8: t. 5b (1915).

Citrus hystrix DC. (1813), sensu stricto. - C. limetta Risso var. OC Risso &

Poit., Hist. Nat. Orangers: 123 (1819). - C. aurata histrix Risso, Hist.

Nat. Eur. Medit. 1: 409 (1826). - C. x aurantium histrix Jacquemont-

Bonnefont, Cat. Prix Courants: 51 (1833). - C. x aurantium L. hystrix

[‘forme culturale’], Guillaumin in Lecomte, Fl. Gén. Indoch. 1: 676

(1911).

[Lemoen Porot Valentijn, Amboina 3(1): 189 (1726)]

? Citrus fusca Lour., Fl. Cochinch: 467 (1791).

Type: not found (see below).

Citrus decumana L. *4. Le citronnier de Combara ou citron a la grecque’

Poir. in Lam., Enc. Méth. 4: 580 (1796), synon. nov. - Citrus combara

Raf., Fl. Tell.: 142 (1838) e descr., synon. nov. - C. macroptera Monttr.

var. combara (Raf.) Tanaka in J. Ind. Bot. Soc. 16: 238 (1937) nom.

superfl. pro var. annamensis.

Type: [Indes, Sonnerat s.n.|, (P-LAM!).

?* Citrus ventricosa Michel, Traité Citronier: 43 (1816) & in Duhamel, Traité

Arbres Arbustes 7:109 (1819); - [ Limo ventricosus Rumph., Herb.

Amb. 2: 102 (1741)] - Risso in syn. C. x bergamia Risso & Poit. var.

ventricosa M. Roem., Fam. Nat. Syn. Monogr. 1: 61 (1846, nom

superfl. (C. limetta var. auraria Risso in syn.).

Limau Hantu and Limau Purut 19]

Type: not indicated. Note that Rumphius's associated plate (t. 26 fig 2,

1741) does not match his (or Michel's) text at all and apparently

represents the sambal, C. amblycarpa (Hassk.) Ochse, a plant whose

relationships are as yet unclear.

Papeda rumphii Hassk. in Flora 25, Beibl. 2: 42 (1842). - Citrus papeda

Migq,., Fl. Ind. Bat. 1(2): 530 (1859), non C. rumphii Risso (‘rhumphiv’,

1844), i.e. C. x aurantium L. cv. - [Limo agrestis Rumpf, Herb. Amb.

2: 104, t. 27 (1741)].

Type [icon]: Rumph., Herb. Amb. 2: t. 27 (1741).

*Citrus tuberoides J.W. Benn., Sel. Rare Fr. Ceylon: t. 1 (1842) & Ceylon:

142 [tt.] (1843).

Type: not indicated. The plate should serve as iconotype in the absence of

any specimen found.

Citrus x bergamia Risso var. unguentaria M. Roem., Fam. Nat. Syn. Monogr.

1: 61 (1846). - [Limo unguentarius Rumph., Herb. Amb. 2: 103, t. 26,

fig 1 (1741)].

Type: not indicated.

The commonly seen cultivated plants tend to have more rounded leaf

apices and bumpier fruit than the ‘wild’ forms do. It is possible that a

number of distinctive cultivars, so far none named, are included here.

Rumphius described the use of his Limo agrestis as a condiment, and both

Limo unguentarius and Limo ventricosus for shampooing and washing, for

example.

Notes: Citrus fusca Lour. may be an earlier name for C. hystrix. Loureiro

cited a Rumphian plate (Herb. Amb. 2: t. 33, 1741) which is referable to C.

x aurantium L. Bitter Orange group, so that later authors have referred C.

fusca to that. However he said that it is widespread in ‘Cochinchina’ but

rarer in China and described its contorted branches, its unpleasantly scented

leaves, the characteristic wide petiole and rough green-brown fruits, all

typical of C. Aystrix, and discusses the medicinal properties of the peel. As

no Loureiro specimen is known, it is perhaps best to consider it a nomen

dubium: if it should prove conspecific, it will be in the interests of

nomenclatural stability to propose its rejection.

Citrus combara Raf. appears to be based on information in Lamarck’s

Encyclopédie Méthodique, hence its specific epithet and placement here.

192 Gard. Bull. Singapore 54 (2002)

‘Torosa’ (kolobat)

Citrus torosa Blanco, FI. Filip.: 609 (1837); cf. Merr., Sp. Blanc.: 204 (1918).

- C. hystrix DC. var. torosa (Blanco) Wester in Phil. Agric. Rev. 8:

19 (1915).

Type: Philippines, Luzon, Batangas Prov., Aug. 1914, Merrill, Sp.

Blancoanae 46 (US [sheet 903713 ‘Sept. 1914’ - Dan Nicolson pers.

comm.]|, neo, designated here [Nicolson and Arculus (2001) have

argued the case for the US set of Merrill’s Species Blancoanae being

considered as appropriate choices for neotypes]; F!, K!, L!, P!, iso).

‘Balincolong’ (balinkolong)

Citrus micrantha Wester “Balincolong’, Wester in Phil. Agric. Rev. 8: t. 7c

(1915).

Citrus micrantha Wester in t.c. : 16, tt. 5c, 6b (1915). - C. hystrix DC. var.

micrantha (Wester) Merr., Enum. Phil. Pl. 2: 343 (1923). - C.

macroptera Montr. var. micrantha (Wester) Tanaka in Trans. Nat.

Hist. Soc. Formosa 22: 430 (1932). - C. combara Raf. var. micrantha

(Wester) Tanaka, Stud. Citr. 9: 3 (1939).

Type: Philippines, Wester s.n. (PNH 7, photo F! [? = (Lamao), Mar. 1915,

Wester 2049 (P, ?iso!)]).

Citrus hystrix var. balincolong Tanaka in Trans. Nat. Hist. Soc. Formosa

22: 429 (1932). - Citrus balincolong Tanaka, Syst. Pomol.: 139 (1951),

nom. nud. synon. nov.

Type: ‘Phlippines’, Bohol, 1914, Wester 4834 (PNH7; PNH, iso!).

A cultivar with small flowers. Although it was referred to C. hystrix by

Merrill (1923), he conceded that it might be of hybrid origin: this

has yet to be tested. I have seen material from only the Philippines

so far.

‘Samuyao sa Amoo’ (samuyau)

Citrus micrantha Wester ‘Samuyao sa Amoo’, Wester in Phil. Agric. Rev.

10: t. 7d (1917).

Citrus micrantha Wester var. microcarpa Wester in op. cit. 8: 21, t. 7b

(1915). - C. hystrix DC. var. microcarpa (Wester) Merr., Enum. Phil.

Fl. Pl. 2: 343 (1923). - *C. westeri Tanaka, Syst. Pomol.: 139-140

(1951, ‘westerii’), non C. x microcarpa Bunge (1833), synon. nov.

Limau Hantu and Limau Purut 193

Type: Philippines, Wester s.n. (PNH?# [F, photo!]).

A cultivar with small leaflets and small aromatic fruits, which in overall

appearance resembles the Chinese C. ichangensis included in the

synonymy of C. macroptera by Burkill (1931).

Further work

The hypothesis above is that from a widespread ‘wild’ form (/imau purut)

a number of cultivars have been selected and that the greatest diversity of

these is in central Malesia. With DNA techniques now available, it should

be relatively easy to test this hypothesis and to show whether or not any of

the cultivars has arisen through hybridization with other species.

It also needs to be ascertained whether C. ichangensis is conspecific

with C. hystrix or not and, if it is, whether it would best be treated as a

geographical subspecies of C. hystrix. According to Needham (1986: 375—

6), C. ichangensis was crossed with C. maxima long ago to produce ‘C.

hsiangyuan’ (i.e. C. wilsonii Tanaka [1932], ? = C. x junos Sieb. ex Tanaka

[1924]), a plant discussed in Han Yen Chih’s Chii Lu of AD 1178.

If this is true, the value of recognizing C. hystrix and its allies as a

distinct subgenus comes into question, particularly with the re-inclusion in

Citrus of Eremocitrus Swingle, Fortunella Swingle and Microcitrus Swingle

(Mabberley, 1998). Recent chemical work (Samuel et al., 2001) has

confirmed the close-knit nature of these plants and also brings back in

Poncirus Raf., long excluded because of its trifoliolate leaves. Elsewhere

in Aurantieae Rchb. (Citreae), e.g. Luvunga Wight & Arn. (Mabberley,

1998), there are also both unifoliolate and trifoliolate species. With the

inclusion of P. trifoliata (L.) Raf., so comes in x Citroncirus, the last surviving

of the hybrid genera recognized by Swingle and followers with a narrow

generic concept:

Citrus L., Sp. Pl.: 782 (1753).

Type: C. medica L.

Poncirus Raf., Sylva Tell.: 143.

Type: P. trifoliata (L.) Raf. = C. trifoliata L.

X Citroncirus J. Ingram & H. Moore in Baileya 19: 171 (1975), synon. nov.

Type: x C. webberi J. Ingram & H. Moore, l.c., non Citrus webberi Wester

(?= C. hystrix/C. ichangensis x C. x aurantium L.), = Citrus x insitorum

Mabb., nom. nov.

194 Gard. Bull. Singapore 54 (2002)

Type [icon]: USDA Yearbook 1904: 228, tt. XI n. 716, XII f. 1-3 (‘Rusk’).

The new epithet means ‘grafters’ citrus’ because this is the citrange, an

important tristeza-resistant rootstock for citrus. The cross was first made

in 1897 and cultivars include ‘Morton’ and ‘Rusk’, which were grown from

the same individual fruit (Swingle 1948: 371).

Within the cultivated citrus, it is now possible to draw up a scheme linking

- putative wild species with their hybrid offspring and to point up where

there are still uncertainties and areas for further analysis (Fig. 1). These

include:

e What the putative unknown parent species of the lemon and the lime are

e Whether or not the Tahiti Lime (C. x J/atifolia Tanaka) is a cross

between the lime and the lemon or citron

e Whether or not the Meyer lemon (C. “Meyeri’) and sweetie (C. limetta

Risso) are crosses between C. x aurantium and the citron (in which

case they are cultivars referable to C. x bergamia) or the lemon.

e Where C. amblycarpa (sambal) fits in this scheme.

Acknowledgments

I am grateful for help from the following who facilitated access to collections

in their care or answered specific Citrus questions for me: William Burger

(F), Larry Dorr (US), Mme S. Hul (P), Ruth Kiew (SING), Jim McCarthy

(Manila), Domingo Madulid (PNH), Dan Nicolson (US), Ohn Set (SING)

and Peter Valder (NSW). I am also grateful to Dan Nicolson (US) for

information on Citrus torosa typification, and to both Rudi Schmid (UC)

and Donna Herendeen (National Agricultural Library, Beltsville, USA)

for photocopies of US literature unavailable in Australia or Western

Europe.

Note added in proof: According to O. Gulsen & M.L. Roose (2001),

‘Chloroplast and nuclear genome analysis of the parentage of lemons’,

Journal American Horticultural Science 126: 210-215, the unknown parent

(CITRUS SP 1 in Fig. 1) of C. x limon is C. x aurantium. If this is so, C. x

bergamia is best treated as a cultivar group of C. x limon to which C. x

taitensis,C. x limetta and C. x 'Meyeri' would also be referred as cultivar

groups, if they have the lemon rather than the citron as a parent.

Hybridization experiments should now be performed to confirm the DNA

work.

195

Limau Hantu and Limau Purut

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References

Burkill, LH. 1931. An enumeration of the species of Paramignya, Atalantia

and Citrus, found in Malaya. Gardens Bulletin Straits Settlements. 5: 212—

220 [see also Index, 1-7, 1932].

Mabberley, D.J. 1984. A monograph of Melia in Asia and the Pacific: the

history of white cedar and Persian lilac. Gardens Bulletin Singapore 37:

49-64.

Mabberley, D.J. 1995. Plants and Prejudice. [Inaugural address as

Professor], University of Leiden, the Netherlands.

Mabberley, D.J. 1997. A classification for edible Citrus. Telopea. 7: 167-

Bi2s

Mabberley, D.J. 1998. Australian Citreae with notes on other Aurantioideae

(Rutaceae). Telopea. 7: 333-344.

Mabberley, D.J. 1999. Where are the Wild Things? Pp. 1-11 in Paradisus:

Hawaiian plant watercolors by Geraldine King Tam. {Text by David J.

Mabberley]. Honolulu Academy of Arts, Honolulu.

Mabberley, D.J. 2001. Citrus reunited. Australian Plants. 21: 5-4.

Merrill, E.D. 1917. An interpretation of Rumphius’s Herbarium

Amboinense. Dept. Agric. Nat. Res., Bur. Sci. Publ.-9.

Merrill, E.D. 1923. An Enumeration of Philippine Flowering Plants, vol. 2.

Bureau of Printing, Manila, the Philippines.

Michel, E. 1816. Traité du citronier. Bertrand, Paris, France.

Needham, J. 1986. Science and Civilisation in China 6(1). Botany. Cambridge

University Press, Cambridge, U.K.

Nicolson, D.H. and D. Arculus. 2001. Candidates for neotypification of

Blanco’s names of Philippine plants: specimens in the U.S. National

Herbarium. Taxon. 50: 947-54. |

Samuel, R., F. Ehrendorfer, M.W. Chase and H. Greger. 2001. Phylogenetic

analyses of Aurantioideae (Rutaceae) based on non-coding plastid DNA

sequences and phytochemical features. Plant Biology. 3: 77-87.

Staples, G.W. and MLS. Kristiansen 1999. Ethnic culinary herbs. University

of Hawaii Press, Honolulu, U.S.A.

Limau Hantu and Limau Purut 197

Swingle. W.T. 1948. The botany of citrus and its wild relatives of the

orange subfamily (family Rutaceae, subfamily Aurantioideae). Pp. 129-

474 in H.J. Webber & L.D. Batchelor (eds), The Citrus Industry, vol. 1.

University of California Press, Berkeley & Los Angeles, U.S.A.

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Gardens’ Bulletin Singapore 54 (2002) 199-204.

Two New Species of Daemonorops (Palmae) from

Sumatra

HIMMAH RUSTIAMI

Herbarium Bogoriense, Jalan Ir. H. Juanda 22,

Bogor, Indonesia

Abstract

Two new species of Daemonorops (Palmae) from Sumatra are described and illustrated, D.

dransfieldii Rustiami from Batang Palupuh, Bukit Tinggi, West Sumatra and D. acehensis

from Takengon, Aceh.

Introduction

While working on a revision of Daemonorops sect. Piptospatha, | came

across herbarium material of two new species from Sumatra. They belong

to the dragon’s blood group, a group within sect. Piptospatha known by

the presence of red resin on the fruit scales. In all, 11 species have been

described in this group (Beccari, 1911; Furtado, 1935; Burret, 1940;

Dransfield,1990).

Daemonorops dransfieldii Rustiami sp. nov.

Species nova a ceteris speciebus fructibus ellipsoideis, petiolis geniculatis

singulariter plicatis differt. Typus: West Sumatra: Bukit Tinggi, Batang

Palupuh, J. Dransfield 2757, 1972. (holo BO).

Figure 1 & 2

Clustering small rattan up to 6 m tall. Stems without sheaths up to 15 mm

diam., with sheaths up to 25 mm diam.; internodes up to 150 mm long.

Leaf sheaths green, covered with black, oblique spines joined at base,

some bulbous—based, up to 3 cm long, sheath surface ridged with caducuous

black indumentum, leaf sheath mouth armed as the rest of sheath; knee

conspicuous, singly plicate, armed as the rest of sheath. Leaves up to 3 m

long including petiole to 25 cm, armed adaxially with short, erect, scattered

spines up to 4 mm long, abaxially armed with erect, rarely solitary spines,

up to 15 mm long; rachis unarmed, or armed only slightly proximally;

cirrus up to 1.5 m long, armed with regularly arranged groups of grapnel-

200 Gard. Bull. Singapore 54 (2002)

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Figure 1. Daemonorops dransfieldii Rustiami

A. Leaf sheath with young infructescence, B. Part of cirrus, C. Part of prophyll. All from

Dransfield 2757.

Two New Sumatran Daemonorops Species 201

‘A

> P,

VES

SAE

Giz

Whe Gaos

eek ite: eae

SI VES

Figure 2. Daemonorops dransfieldii Rustiami

A. Infructescence, B. Mature fruit,C. Seed, D. Seed in transverse section. All from J. Dransfield

ZIT

202 Gard. Bull. Singapore 54 (2002)

like spines, blackish at the tip; leaflets mostly regularly arranged, slightly

irregular above, 30 on each side of the rachis, stiff, horizontal; leaflets

lanceolate, papery, acute, 25-31 cm long, c. 2.5 cm wide, armed with

scattered, reddish, short bristles along the main nerve on both surfaces.

Inflorescences pendulous to 35 cm long, peduncle 13-16 cm long, armed

distally with groups of spines; peduncular bracts leathery, erect, c. 17 cm

long, 3 cm wide, ellipsoid oblong, covered by rusty indumentum, armed

with very rare, scattered spines; partial inflorescences 6, each bearing up to

11 rachillae; involucre pendulous, flat, just above the involucrophore, c. 5

mm long; involucrophore short, papery, c. 2 mm long. Female and male

flowers unknown. Fruits broadly oblong, c. 2.5 x 1 cm, covered in 9 vertical

rows of scales encrusted with reddish brown dragon’s blood. Seed ovoid, c.

18 x 7 mm, surface smooth.

Distribution: Sumatra - known only from Batang Palupuh, Bukit Tinggi,

West Sumatra.

Habitat: Found on a hill slope in disturbed forest at 800 m altitude.

Notes: Named in honour of John Dransfield, palm systematist who collected

this species. Daemonorops dransfieldii differs from the other dragon’s blood

species by its broadly oblong fruit and singly plicate knee. The description

is based only on the type specimen. As yet, this species is known to occur

only in the Batang Palupuh area, but may be found elsewhere in West

Sumatra if further fieldwork is carried out.

Daemonorops acehensis Rustiami sp. nov.

Species nova a ceteris speciebus fructibus resinosis habitu parvo, vaginis

foliorum spinulis confertissimis, longis erectis armatis et cirro non nisi

usque ad 40 cm longo differt. Typus: North Aceh, Takengon, Burlintang,

J. Dransfield & D. Saerudin 2015, 1971 (holo BO).

Figure 3

Clustering small rattan up to 5 m tall. Stems without sheaths up to 10 mm

diam., with sheaths up to 25 mm diam.; internodes up to 50 mm long. Leaf

sheaths dark green, covered with black, very dense, single spines, up to 2.5

cm long, sheath surface covered with very dense black indumentum, leaf

sheath mouth papery, armed as the rest of sheath, knee conspicuous, armed

as the rest of sheath. Leaves up to 2 m long including petiole to 35 cm;

abaxial and adaxial surfaces armed with long, erect spines to 3 cm long;

rachis armed as petiole, sparsely covered with black indumentum; cirrus to

40 cm long, armed with regularly arranged groups of grapnel-like spines,

Two New Sumatran Daemonorops Species 203

HI? «

UL Tirtet Sskoke

foce .

Figure 3. Daemonorops acehensis Rustiami.

A. Leaf sheath with infructescence, B. Part of leaves, C. Part of cirrus. All from Dransfield

& Saerudin 2015.

204 Gard. Bull. Singapore 54 (2002)

blackish at the tip, proximally armed with single hook-like spines; leaflets

mostly regularly arranged, slightly irregular towards the leaflet tip, 25 on

each side of the rachis; leaflets lanceolate, papery, acute, 24-26 cm long, 2

cm wide, armed with scattered, reddish, short bristles along the main nerve

on adaxial surface. Inflorescences pendulous to 20 cm long; peduncle to 4

cm long, armed with scattered, single spines; peduncular bracts papery,

bearing scattered solitary spines, partial inflorescences 4; each bearing up

to 6 rachillae; involucre pendulous, flat, just above the involucrophore, 5

mm long; involucrophore short, papery, 2 mm long. Female and male

flowers unknown. Fruits rounded, c. 2.2 x 1.8 cm, covered in 12 vertical

rows of scales encrusted in reddish brown dragon’s blood. Seed ovoid, c.

18 x 7 mm, surface reticulate.

Distribution: Sumatra - endemic to Birungen, Takengon, North Aceh.

Habitat: Found on steep hill slopes in relatively dry montane forest at

1800 m altitude.

Notes: This species, the only dragon’s blood species recorded from Aceh,

differs from other members of the group by its small habit, the leaf sheath

armed with very dense, long, erect spines and with the cirrus only up to 40

cm long.

Acknowledgements.

I should like to thank Dr. John Dransfield, Kew, for critical comments on

the material and assisting with the Latin diagnosis. I have also to thank

Iskak Syamsoedin and Tirto, who prepared the illustrations.

References

Beccari, O. 1911. The Species of Daemonorops. Annals Royal Botanic

Garden. Calcutta. 12: 1-233.

Burret, K. E. M. 1940. Indomalayische Palmen. Notizblatt desbotanischen

Gartens und Museums Zu Berlin-Dahlem. 15: 191-192.

Dransfield, J. 1990. Notes on rattans (Palmae: Calamoideae) occuring in

Sarawak, Borneo. Kew Bulletin. 45: 76.

Furtado, C. X. 1935. Notes on some Malayan Daemonorops. Gardens’

Bulletin Straits Settlement. 8: 339-365.

Gardens’ Bulletin Singapore 54 (2002) 205-206.

Mangifera odorata (Anacardiaceae) is a Hybrid

R. KIEW

Singapore Botanic Gardens,

Singapore 259569

Abstract

Mangifera xodorata Griff. (pro sp.) is confirmed as a hybrid between M. foetida Lour. x M.

indica L.

Mangifera odorata

Ding Hou (1978) first suggested that the polymorphous Mangifera odorata

Griff., a well-known Malesian fruit tree locally known as kuwini in

Peninsular Malaysia, was a hybrid possibly between M. foetida Lour. and

M. indica L. Corner (1988) also considered this possible because kuwini is

difficult to separate from M. foetida in the sterile state. They had both

noted that kuwini is only known from villages, whereas M. foetida is not

only grown in villages but has also been collected wild from the forest. M.

indica is the commercial cultivated mango. However, Kostermans and

Bompard (1993) disputed its hybrid status and considered M. odorata a

‘good species’ that could be distinguished from M. foetida on leaf venation.

Molecular analysis by Teo et al. (2002) using AFLP (amplified fragment

length polymorphism) produced 28 monomorphic bands specific to M.

foetida of which 20 occurred in kuwini; while M. indica had 22 monomorphic

species-specific bands of which 16 occurred in kuwini. Kuwini, in contrast,

had no species-specific bands but had additively inherited bands from M.

foetida and M. indica. This provides evidence for the hybrid status of

kuwini and that its parents are indeed M. foetida and M. indica.

Further, UPGMA (unweighted pair group method with arithmetic

mean) analysis showed that kuwini has closer genetic similarity to M. foetida

(76% similarity) than to M. indica (66 % similarity).

Kuwini therefore cannot be maintained as a distinct species and is

here given taxonomic status as a hybrid. |

Mangifera xodorata Griff. (pro sp.)

= M. foetida Lour. x M. indica L.

Griffith, Notul. 4 (1854) 417; Ding Hou, FI. Mal. 1,8 (1978) 437; Kostermans

& Bompard, The Mangoes. (1993) 167.

Type: Griffith 1098 Malacca. (holo K, iso P).

206 Gard. Bull. Singapore 54 (2002)

Acknowledgements

Advice on nomenclature from J.F. Veldkamp is much appreciated.

References

Corner, E.J.H. 1988. Mangifera. Wayside Trees of Malaya. 1: 115-121.

Ding Hou. 1978. Mangifera. Flora Malesiana. 1, 8: 423-440.

Kostermans, A.J.G.H. & J.M. Bompard. 1993. The Mangoes. Academic

Press, London. p. 167.

Teo, L.L., R. Kiew, O. Set, K. Lee & Y.Y. Gan. 2002. Hybrid status of

kuwini, Mangifera odorata Griff. (Anacardiaceae) verified by amplified

fragment length polymorphism. Molecular Ecology. 11: 1465-1469.

Gardens’ Bulletin Singapore 54 (2002) 207-216.

Taxonomic Notes on the Tree Flora of Brunei: 1

P.S. ASHTON

Arnold Arboretum, Harvard University, U.S.A.

and

Royal Botanic Gardens, Kew, U.K.

Abstract

As a precursor to The Field Guide to the Trees of Brunei Darussalam, the following taxonomic

changes and novelties are presented: Gluta wallichii (Hook.f.) Ding Hou subsp. lafrankiei

P.S.Ashton subsp. nov. and Parishia coriacea P.S.Ashton, sp. nov. (Anacardiaceae);

Canarium latistipulatum Ridley subsp. mitus P.S.Ashton subsp. nov.and Dacryodes

patentinervia (Leenh.) P.S.Ashton, stat. nov. (Burseraceae); Hopea rugifolia P.S.Ashton,

sp. nov. and Vatica patentinervia P.S.Ashton sp. nov. (Dipterocarpaceae).

Introduction

Brunei Darussalam has become the most intensively collected region, for

its area, of Borneo. The tree flora is particularly well collected. Some 7,600

tree collections are cited in Kirkup (1996). This checklist, however, omitted

some 300 old collections in the herbaria of the Sarawak Forest Department

(SAR) and Forest Research Institute Malaysia (KEP), as well as most of

the c. 3,000 ecological voucher specimens collected during the course of

the Universiti Brunei Darussalam (UBD)-Royal Geographical Society

(RGS) Rainforest Project 1991-1992; by Japanese scientists associated with

the Brunei Forest Department’s Sungei Liang Research Station supported

by the Japan International Collaboration Agency (JICA); by S.J. Davies

in the course of establishing permanent plots on behalf of UBD during

1994-1995, and by myself during ecological research on behalf of the Brunei

Forest Department during 1958-1959. With these and recent Forest

Department collections included, Brunei has achieved a collecting intensity,

for trees alone, of almost 30 numbers per 100 km”.

Brunei is estimated to have a tree flora of almost 3,000 species,

roughly equal to the whole of Peninsular Malaysia and Singapore. Brunei

is not a formal participant in the Tree Flora of Sabah and Sarawak project

(Soepadmo, 1995), but the occurrence of species in Brunei is noted in that

Flora. Initiated in 2001, a ten-year project sponsored by the Brunei Shell

Petroleum Company, Universiti Brunei Darussalam, and the Brunei Forest

Department under the leadership of Dr Kamariah Abu Salim aims to

complete a comprehensive field guide to the tree flora of Brunei based on

208 Gard. Bull. Singapore 54 (2002)

the format of the Audubon Guides to U.S. Wildlife and the Collins Guides

in Europe. This project is planned to move in tandem with and to

complement the Tree Flora of Sabah and Sarawak project. All species

recorded from Brunei, and those expected owing to their presence in

adjacent regions, will be included in field keys and provided with short

field diagnostic descriptions and line illustrations. About one third of the

1,100 species recorded in the 52 ha tree demography plot in nearby Lambir

National Park, Miri, Sarawak, for instance, have yet to be found in Brunei,

and these will be included, either as descriptions or in footnotes to recorded

species.

This enterprise provides an opportunity to observe and collate more

detailed information on the natural history of individual species than is

possible in floras of larger, less intensively collected regions such as Sabah

and Sarawak. It is, therefore, providing opportunity for identification of

discontinuities in variation within currently recognised species, meriting

specific or infraspecific recognition. In a few cases, novel species are being

discovered. This communication will be the first in a series, in which these

new entities and taxonomic changes, will be discussed and formalised prior

to publication of the volumes of the guide. Mark Coode is thanked for

assisting wih the Latin.

New Taxa and Status Changes

Anacardiaceae

1. Gluta wallichii (Hook. f;) Ding Hou subsp. lafrankiei P.S.Ashton,

subsp.nov.

A Gluta wallichii subspecie typica petiolo et foliorum nervis subtus badio-puberulentibus

lamina subtiliter coriacea margine tenu revoluta basi cuneata costis utrinsecus 13—18

satis distincta. Typus: Awang Yakup S 8903 Borneo, Sarawak, Semengoh Forest Reserve

(holo K, flowers).

Notes: J.V. LaFrankie, in the course of identifying the trees of the 52-ha

tree demographic plot within Lambir National Park recognised two distinct

entities within the range of variation in Gluta wallichii described by

Kochummen (1996). In one entity, conforming to the typical subspecies,

the bark is pale brown, shallowly cracked and flaking, the leaf glabrescent,

the lamina thinly coriaceous, wavy on drying, with shortly tapering base,

hardly revolute at the margin (visible by lens only), and with at least 19

pairs of secondary veins. In the other, the bark is red-brown and fissured,

the leaf venation beneath and petiole brownish puberulent, the lamina

Tree Flora of Brunei 209

coriaceous, drying flat, with straight visibly revolute margin, strictly cuneate

base and 13-18 secondary pairs of veins. Further, the typical subspecies

mostly occurred on clay and sandy clay soils, while the new subspecies

occurred on humic yellow sands, and we have found this to be the case in

Brunei and elsewhere in the region. Nevertheless, the distinctly narrowed

leaves with glaucous undersurface and slender petioles, as well as flower

and fruit characters of G. wallichii are shared by both entities, so I chose

to recognise them as subspecies until more is known about patterns of

variation throughout the range of G. wallichii.

Other specimens examined: BRUNEI - Bukit Patoi Wood et al. SAN 17147

(BRUN, fruit). SARAWAK - Bidi, Bau Burley & Lee 330 (A, K, SAR),

Semengoh Forest Reserve Asah Unyong Tree No.156 (K, SAR, young

flowers), Rosli S 15774 (K, SAR, young fruit), Rosli S 15746 (K, SAR,

fruit), Anderson S 14932 (K, SAR, flowers), S 9387 (K, SAR, flowers).

2. Parishia coriacea P.S.Ashton, sp. nov.

Parishiae maingayi Hook. f. affinis, sed sepalis in fructu brevioribus, foliolis utrinsecus

3 costis lateralibus prominentibus distinguitur. Typus: Niga Ningkat NN 230, Borneo,

Brunei, Bukit Besong, Ukong, (holo, K iso BRUN fruit).

Tree to 30 m tall, 80 cm diam.; bark pale grey-brown, shallowly powdery

flaking. Young parts shortly rust-brown hairy, the hairs caducous except

on panicle, calyx and nut. Twigs stout c. 1 cm diam., rugose. Leaf rachis up

to 40 cm long, petiole up to 18 cm long; leaflets c. 3 pairs, opposite, lamina

elliptic, 7.5-20 x 4.5-7 cm, coriaceous, drying grey-brown; base obtuse or

cuneate, margin slightly revolute, apex acuminate, acumen to | cm long;

midrib prominent beneath, narrowly elevated and furrowed above;

secondary veins 12-25 pairs with many shorter intermediate veins, slender

but prominent beneath, evident but hardly elevated above; the reticulate

tertiary veins likewise; petiolules 5-10 mm long, stout. Panicle up to 60 cm

long, terminal. Flower bud up to 6 mm diam. Male flower: perianth 4-

merous, sepals pubescent, c.l1 mm long, petals glabrous, c.2 mm long,

stamens 4, surrounding the densely pubescent ovary vestige. Female flower

unknown. Fruit: sepals to 5.5 x 1 cm, lorate; nut ovoid, up to 2 x 1 cm,

shortly beaked, dark chestnut-brown velutinate.

Distribution and habitat: Endemic to mixed dipterocarp forests on deep

yellow coastal sands of the Neogene coastal hills of Brunei and north-east

Sarawak, a habitat of exceptional richness and endemism.

Notes: This species is easily distinguished by its relatively short fruit sepals,

210 Gard. Bull. Singapore 54 (2002)

at first shorter than the ripening nut, and by its 3 pairs of coriaceous

leaflets drying distinctly pale grey-brown, with slender but prominent

venation beneath..

Other specimens examined: BRUNEI - Andulau Forest Reserve Ashton

BRUN 3272 (BRUN, K fruit), BRUN 18568 (BRUN sterile), Sungei Liang

Dransfield JD 7022 (BRUN, K fruit), Bukit Teraja Coode 6959 (BRUN, K

fruit), Loagan Merimbun Suzuki K13400 (BRUN sterile). SARAWAK -

Bintulu, Labang Forest Reserve Ilias Pa’ie S 15817 (K, L fruit), Limbang,

Ulu Medamit Wright & Othman Ismawi S 32313 (K flowers).

Burseraceae

1. Canarium latistipulatum Ridl. subsp. mitus P.S.Ashton, ssp. nov.

A Canario latistipulato typico arbor grandis tomento persistenten scabrido, foliolis

latioribus, fructibus vastis suave esculentibus differt. Typus: Coode 7107 Borneo, Brunei,

Bukit Bahak, Rambai (holo K, iso BRUN, young fruit).

An elegant columnar prominently buttressed village tree with dark

hemispherical crown. Young twigs, leaflets beneath, leaf stalks and

inflorescences sparsely shortly tawny scabrous tomentose. Stipules reniform,

up to 15 x 10 mm, attached near the base of the petiole, tardily caducous.

Leaflets usually 7, lamina broadly elliptic, c. 11 x 5 cm, applanate, drying

dark reddish to greenish brown; base broadly cuneate to obtuse, margin

shallowly dentate, apex acuminate, acumen prominent; secondary veins 7-

-10 pairs, arched, somewhat anastomosing within margin, without

intermediate veins, prominent beneath, not so to shallowly depressed above

as also the subscalariform tertiary veins. Flowers unknown. Infructescence

paniculate. Fruit drupaceous, broadly ellipsoid, up to 8 x 6 cm, lustrous,

ripening yellow-brown; mesocarp orange-brown, fleshy and fibrous,

resinous; endocarp woody, sharply triangular in cross section.

Notes: It is surprising that this fruit tree, known to Kedayan and Dusun

farmers as mitus, should have so long remained unrecognised. Admittedly,

we only have information of its cultivation in northern Brunei from Bandar

Seri Begawan westward to the Tutong valley. |

In Kirkup (1996, p. 44), this plant had been mistaken for Canarium

littorale Blume, but the distinct reniform stipules are placed towards the

base of the petiole, which is a distinguishing character of C. latistipulatum

Ridl., and not on the adjacent twig as in C. littorale. The fruits, which have

yet to be collected for the herbarium, are the size of a mango. They are

buried in order to rot off the highly resinous mesocarp, then the seed

within its striking triangular woody endocarp is cleaned and marketed.

Tree Flora of Brunei 7 |

The fruit of the typical subspecies, which is said to be a native forest tree

of Sabah and Sarawak (Kochummen, 1995), is unknown, but the narrow

leaflets, powdery puberulent tomentum and apparently small size serve to

distinguish it.

Other specimens examined: BRUNEI — Tutong: Pak Engalau, Tasek

Merimbun Bernstein JHB 394 (BRUN, K sterile), Panchang K.M. Wong

s.n. (BRUN sterile), Kampong Mitus P.S. Ashton s.n. (BRUN, K sterile).

2. Dacryodes patentinervia (Leenh.) P.S.Ashton, stat. nov.

Basinym: Dacryodes macrocarpa (King) H.J. Lam var. patentinervia Leenh., Flora

Malesiana 1, 7 (1976) 821; Kochummen, Tree Flora of Sabah and Sarawak 1 (1996) 71.

Typus: Sinclair & Kadim 10492 Borneo, Brunei, Labi Forest Reserve (holo L, iso K,

SAN, SAR fruit).

Tree to 35 m tall, 80 cm diam., with narrow buttresses and pale yellow-

brown thinly flaky bark. Bud and young shoots puberulent, other parts

glabrous. Leaves: rachis up to 23 cm long and 2 mm diam., slender, terete

or somewhat angular towards base; leaflets 5—7; lamina narrowly elliptic to

lanceolate, 7-14 x 4-7 cm, drying bright coppery brown with the midrib

_ darker beneath, equal to subfalcate, base cuneate, margin slightly revolute,

apex acuminate, acumen tapering, slender, c. 1 cm long; midrib slender

beneath, applanate above; secondary veins 10-16 pairs, patent, hardly raised

below, evident but applanate above as also the many short intermediate

veins and densely reticulate tertiary veins; petiolules 12-35 mm long on

lateral, up to 6 cm long on terminal leaflets, adaxially flattened at base

otherwise terete, prominently swollen at each end. Inflorescence up to 24

cm long, laxly paniculate, shortly branched, glabrous. Male flower: perianth

3-merous, glabrous; sepals united into a basal cup; stamens 6, anther deltoid;

ovary vestige columnar, tapering. Female flower unknown. Fruit broadly

ellipsoid, up to 5 x 3 cm, asymmetric with eccentric stigma, ripening lustrous

apple red.

Distribution and habitat: Endemic to northwest Borneo from the Rejang

valley eastwards, where it is among the most abundant Burseraceae in its

chosen habitat, in mixed dipterocarp forest on the deep yellow sands of

the Neogene coastal hills; it also occurs in upper dipterocarp forest on

shale ridges at 650-800 m, and on ultramafic substrates at Bukit Hampuan,

Sabah at 1500 m.

Vernacular names: Dacryodes patentinervia is known as sabal (Brunei, Iban),

212 Gard. Bull. Singapore 54 (2002)

sibut (Tutong, Dusun), and seladah (Sarawak).

Uses: The fruit is collected from the forest and eaten as a popular laxative.

Notes: This entity has been confused with Dacryodes expansa (Ridl.) H.J.

Lam - holotype Haviland 2271 Sarawak, Kuching (K) - and was formally

described as a variety of the well-known D. macrocarpa (King) H. J. Lam -

holotype King’s Collector 7298 Peninsular Malaysia, Perak (K). All three

taxa share a distinctive coppery brown tinge to the dry leaf. Leenhouts

(1956) considered D. macrocarpa, with the present entity as a subspecies,

to differ from D. expansa in petal shape, filaments united with the disc,

and 6—10(—13) of secondary veins as opposed to 10—13 pairs in D.

expansa. Later, in 1972, he distinguished the two species according to

whether the petiole is terete except for the flattened base (D. expansa), or

prominently flattened at base (D. macrocarpa), and only in 1976 did he

redefine variation within D. macrocarpa adding var. patentinervia, and

drew up a new description of D. expansa.

Dacryodes macrocarpa, sometimes a large tree of the mixed peat

swamp forest, is here distinguished by fruits ripening dull green; leaflets

unequal, broadly ovate, of often markedly irregular shape, shortly abruptly

acuminate; midrib prominent on both surfaces; secondary veins 7—9 pairs;

and inflorescence up to 20 cm long, much branched and stout.

Dacryodes expansa, with which the present entity was originally

confused by Hasan and Ashton (1962) and Ashton (1964) seems to be

known only from the type from west Sarawak. It differs from D.

patentinervia in having larger, ellipsoid flower buds up to 4 x 3 mm; 3—4

(not 2) pairs of thinly coriaceous leaflets with 10—13 pairs of ascending

secondary veins prominent beneath, and remote tertiary veins; and a

flattened, sharp edged, adaxial side to the stout petiole, which is up to 4

mm diam. The fruit is unknown.

Other specimens examined: BRUNEI - Bukit Teraja Dransfield JD 6867

(BRUN, K fruit), Coode 6892(K fruit); Andulau Forest Reserve Ashton

BRUN 253 (BRUN, K fruit), BRUN 1567] (BRUN sterile), BRUN 15272

(BRUN sterile), BRUN 16193 (BRUN fruit), Wyatt-Smith KEP 80093 (K

fruit), Wood et al. SAN 17487 (BRUN fruit), ecological specimen Ashton

PS 3966 (BRUN sterile), (K); Bukit Kukub BRUN 16781 (BRUN fruit);

Sukang BRUN 18665 (BRUN fruit); Luagan Lalak, Labi Niga Ningkat NN

185 (K fruit); Merangking-Buau, Belait BRUN 16220 (BRUN fruit), Bukit

Batu Patam, Ulu Ingei, Belait Wong WKM 1038 (BRUN, K flowers);

Bukit Biang Wood et al. SAN 17123 (BRUN fruit); Bukit Belalong west

ridge Wong WKM 1534 (BRUN, K fruit); Bukit Belalong south ridge

Tree Flora of Brunei 213

Wong WKM 1356 (BRUN, K young fruit). SARAWAK - Miri, Lambir

National Park: Dan Hj. Bakar S 4370 (K fruit); George S 40503 (K fruit),

Sungai Jangkang, Bakam Abang Mohtar S 47111 (K fruit), path to Bukit

Lambir Yeo & Jugah Kudi S 38496 (K young fruit), Mount Lambir Jlias

Paie S 16602 (K young fruit); Gunong Mulu National Park (Camp 3) S$

58412 (K young fruit); Jugah Kudi S 23696 (K fruit), Bukit Iju, Balingian

Sibat Luang S 23655 (K fruit); Ulu Kenyana, Mukah Ashton S$ 19494 (K

young fruit).

Dipterocarpaceae

1. Hopea rugifolia P.S.Ashton, sp. nov.

Hopeae mesuoides Ashton, H. sphaerocarpae (Heim) Ashton, H. subalatae Sym. affinis

sed lamina coriacea rugulescenti grisea nervis subtus magis prominentibus basi cuneata,

petiolo brevi satis distincta. Typus: Wong WKM 1414 Borneo, Brunei, Bukit Belalong

south ridge, flowers (holo K, iso BRUN, flowers).

Small monopodial understorey tree up to 20 m tall, 20 cm diam., with low

buttresses and stilt roots; bark smooth. Young parts densely evenly minutely

grey-brown hairy, the hairs persisting sparsely on leaf venation beneath,

midrib above at base, and on petiole. Stipules acicular, up to 6 mm long,

caducous. Lamina broadly lanceolate, 5—9 x 2—3.5 cm, chartaceous, drying

grey-brown and wrinkling; base broadly wedge-shaped to obtuse, apex

acuminate, acumen slender, prominent up to 1.5 cm long; midrib slender

but elevated beneath, obscure and depressed above; secondary veins c. 10

pairs, slender but distinctly raised beneath, with short intermediate veins;

tertiary veins densely scalariform, evident beneath; petiole to 4 mm long,

slender. Panicle axillary, up to 4 cm long, slender, when dried up to 0.7 mm

diam. at base, terete, glabrous, singly branched; bracteoles deltoid, up to 1

mm long, glabrous, subpersistent. Flower bud to 1.5 mm diam. at anthesis,

small, broadly ovoid; calyx sparsely pubescent outside, sepals suborbicular,

to 1 x 1 mm diam., outer two subacute or obtuse, inner three mucronate;

corolla purple, petals oblong, obtuse, to 3 mm long, puberulent on parts

exposed in bud; stamens 15, filaments short, broad at base, tapering, anthers

ellipsoid, connectival appendage c. twice length of anther, slender; ovary

and stylopodium short, narrowly cylindrical, truncate, medially constricted,

glabrous, style conical, short, glabrous. Fruit glabrous; pedicel up to 5 mm

long, 0. 5 mm diam., very slender; sepals ovate, subequal, up to 8 x 7 mm

when ripe, acute, saccate, incrassate and appressed to nut; nut up to 10x 8

mm, broadly ovoid, with up to 2 mm-long prominent stylopodium remnant.

214 Gard. Bull. Singapore 54 (2002)

Distribution and habitat: It occurs, like many dipterocarps with wingless

fruits, in dense clusters in the forest understorey, on sandy clay soils at low

altitude and also skeletal humic clay soils in upper dipterocarp forest at

600-800 m altitude. At Lambir National Park, north-east Sarawak, it shares

the forest with Hopea mesuoides P.S.Ashton, which is, however, confined

there to yellow humic sandy soils.

Notes: Hopea rugifolia is named for its distinctively wrinkled fallen leaf in

a family where leaves are generally more coriaceous. This species belongs

to a group of closely similar monopodial understorey species comprising

H. mesuoides (Sarawak north-east of S. Balingian and Brunei), H.

sphaerocarpa (Heim) P.S.Ashton (with which H. rugifolia has heretofore

been confused but which is now confirmed only from Sarawak west of

Batang Lupar), and H. subalata Symington. The last is a point endemic of

Kanching Forest Reserve, Selangor, where it is locally gregarious on the

Klang Gates quartzite in association with the only known west coast locality

of Dryobalanops aromatica Gaertn. f. H. subalata has been shown to be

triploid and likely partially apomictic though adventive embryony (Kaur et

al., VITG):

The present species is distinguished from these three species by its

chartaceous leaf drying grey-brown and wrinkling, with cuneate rather

than obtuse base, prominent secondary veins beneath, and short petiole.

Other specimens examined: SARAWAK - Bukit Mersing, Anap Sibat Luang

S 25028 (K fruit), Lambir National Park S 46439 (K fruit).

2. Vatica patentinervia P.S.Ashton, sp. nov.

Vaticae congestae Ashton similis sed costis foliorum lateralibus utrinsecus

7-8 acumine prominenti tomento plano distinguitur, a V. endertii Sloot.

inflorescentibus ramifloris brevioribus, fructibus maioribus, ramulis

tomentosis, lamina coriacea costis paucioribus differt. Typus: //ias Pa’ie S

15148 Borneo, Sarawak, Bintulu, Segan Forest Reserve (holo K, iso SAR

fruit).

Subcanopy tree, to 30 m tall. Young parts densely evenly buff pubescent,

the hairs persistent on twig, petiole and inflorescence. Twig slender, ribbed.

Stipule narrowly deltoid, c. 5 x 3 mm. Lamina elliptic-obovate, 7-15 x

3.5v6 cm, thinly coriaceous, applanate to narrowly revolute, drying pale

buff-grey; base cuneate to obtuse, apex acuminate, acumen up to | cm

long, prominent; midrib hardly elevated above, prominent beneath;

secondary veins 7-8 (up to 12 in juveniles) pairs, evident above, prominent

beneath; tertiary veins reticulate; petiole 1.2-2.5 cm long, rugulose.

Tree Flora of Brunei 215

Inflorescence axillary to ramiflorous, fascicled, up to 3 cm long, up to 2

mm diam., singly branched. Flowers unknown. Fruit: pedicel up to 4 mm

long, slender; sepals unequal, free to base; longer two up to 12 x 2.2 cm,

lorate, obtuse, prominently 3-veined; shorter three up to 4.5-1.5 cm,

narrowly ovate, acute, recurved. Nut ellipsoid, up to 12 x 8 mm, tapering

to 6 mm-long style remnant, glabrous.

Notes: This species, which is only known from clay soil in mixed dipterocarp

forest from two localities, shares the large fruit and fascicled ramiflorous

inflorescences of Vatica congesta P.S.Ashton, but is distinguished by the

distinct even buff pubescence of twig, petiole and inflorescence, fewer (7-

8) more slender pairs of secondary veins (more in juveniles; 9-11 pairs in

V. congesta), and prominently acuminate, not obtuse or retuse, less

coriaceous lamina. V. endertii Slooten differs in having glabrescent young

parts, solitary terminal or axillary inflorescences, smaller fruit, and a more

coriaceous lamina with 11-13 pairs of secondary veins. Saplings, whose

leaves may have up to 12 pairs of secondary veins, can also be confused

with those of V. odorata (Griff.) Symington subsp. mindanensis (Foxw.)

P.S.Ashton, but the leaves dry distinct buff-grey, which with the dense

even buff pubescence on twigs and petioles serves to distinguish them

from all three species.

Other specimens examined: BRUNEI - Kuala Belalong Ashton S 5744 (SAR

sterile), Balslev et al. 62, 68, 84, 91, 123, 133, 160, 188, 208, 249, 255, 286,

359, 363, 365, 369, 372, 396, 437 (UBD, sterile ecological specimens).

References

Ashton, P.S. 1964. Ecological Studies in the Mixed Dipterocarp Forest of

Brunei State. Oxford Forestry Memoirs. 24: Appendix.

Kirkup, D.M. 1996. A Checklist of the Flowering Plants and Gymnosperms

of Brunei Darussalam. Forest Dept. Brunei Darussalam & Royal Botanic

Gardens, U.K.

Hasan Pukol and P.S. Ashton. 1962. Checklist of Brunei Trees. State Forest

Office, Brunei.

Kaur, A., C.O. Ha , K. Jong, V.E. Sands, H.T. Chan, E. Soepadmo and

P.S. Ashton.1978. Apomixix may be widespread among the trees of the

climax rain forest. Nature, Lond., 271: 440-442.

216 | Gard. Bull. Singapore 54 (2002)

Kochummen, K.M. 1995. Burseraceae. Tree Flora of Sabah and Sarawak.

1: 45-100.

Kochummen, K.M. 1996. Anacardiaceae. Tree Flora of Sabah and Sarawak.

2: 1-92.

Leenhouts, P.W. 1956. Burseraceae. Flora Malesiana. 1, 5: 209-296.

Leenhouts, P.W. 1972. Agenda, corregenda and emendenda: Burseraceae.

Flora Malesiana. 1, 6: 917-928.

Leenhouts, P.W. 1976. Agenda, corregenda and emendenda: Burseraceae.

Flora Malesiana. 1, '7: 820-822.

Soepadmo, E. 1995. Background to the Tree Flora of Sabah and Sarawak

Project. In: (eds.) E. Soepadmo and K.M. Wong. Tree Flora of Sabah

and Sarawak. 1: XIII-XIX.

Gardens’ Bulletin Singapore 54 (2002) 217-238.

New Taxa and Combinations in the Genus Diplycosia

(Ericaceae) of Borneo and Peninsular Malaysia.

G.C.G. ARGENT

Royal Botanic Garden Edinburgh,

Edinburgh EH3 5LR, U.K.

Abstract

Seven new species of Diplycosia and one new form are described: D. abanii; D. mogeana;

D. rhombica; D. mantorii; D. othmanii; D. paulsmithii; D. ngii and D. salicifolia Sleumer

forma gigantea. The genus Pernettyopsis King & Gamble is reduced to synonymy with

Diplycosia and the relevant new combinations are made. Advice on collecting material of

the genus is given.

Introduction

The genus Diplycosia Blume was effectively monographed for Flora

Malesiana (Sleumer 1966—1967), as all but two of the 97 species then

recognised occurred in the Malesian region. Since then there has been

little interest in a genus of small epiphytic shrubs with rather insignificant

flowers and no known economic value. A few new species have been

described (Argent 1982, 1989) and new accounts of the Kinabalu species

have been published (Argent in Wong & Phillipps, 1996; Beaman et al.,

2001), but there is little there to advance our understanding of the

relationships within the genus. Molecular work (Kron, 2001) suggests that

the genus is nested within Gaultheria although morphologically it remains

quite distinct and only very rarely is there any problem in referring even

sterile plants to one or other of these genera. The genus is most diverse in

Borneo with well over half the total number of described species and Mt

Kinabalu remains by far the richest single locality with 27 species known

from its environs. In the present paper, one new species from Peninsular

Malaysia and six new species and one new form from Borneo are described

emphasizing Borneo as the centre of diversity of the genus.

The genus is under-collected partly because of the small flowers but

also because many species are epiphytes growing in peaty accumulations

on large trees and are thus inaccessible. Even when they are collected the

resulting specimens are often poor. Collections made with field descriptions

of flowers often lack them in the herbarium as the flowers easily fall off in

drying and careless handling. Details of indumentum (of great importance

in discriminating taxa) are often difficult to interpret because young stems

218 Gard. Bull. Singapore 54 (2002)

(which do not carry flowers) are not collected and old ones lose bristles

and hairs or conversely appear to gain them with the erect branches of

algal epiphytes. Field descriptions of the fruit may eventually lead to

useful subdivisions within the genus but these are rarely adequate and

need to be made preferably with field sketches of the fruit structure which,

because of its fleshy nature, is lost on pressing and drying. Variations in

both wild and cultivated specimens observed have led to the proposal in

this paper to reduce the genus Pernettyopsis King & Gamble to synonymy

with Diplycosia. There are undoubtedly many more species to be recognised

and described especially from the mountains in Borneo but more, carefully

prepared collections are needed. Specimens should preferably possess

open flowers, some of which should be separated on collection into a

numbered packet so that they are less likely to be lost on drying and

handling. Next most important is to look carefully for some new young but

fully or nearly fully expanded leafy shoots which will show the indumentum

structure clearly. If there are mature fruits, a quick sketch showing which

parts are fleshy and the relative position and length of calyx lobes to ovary

might ultimately prove illuminating. Sterile specimens with the indumentum

in good condition are still worth collecting as these can often still be

identified to a species.

Diplycosia species are not especially easy to cultivate although we

have maintained at least one of our collections in the Royal Botanic Garden

Edinburgh for more than 25 years. It would undoubtedly give interesting

information as to variability if more species could be grown satisfactorily.

Some species are extremely attractive plants with a dense golden-brown

indumentum but unfortunately these have proved to be the most difficult

to keep alive in cultivation.

New Taxa

1. Diplycosia abanii Argent, sp. nov.

(Named after the collector of the species from the Forestry Department in

Sandakan, Sabah.)

Diplycosiae clementium Sleumer similis sed setis in pedicellis calyceque

adpressis, foliis laevibus (haud rugosis) cum reticulo obscuro, venis

lateralibus abaxialibus haud elevatis et setis in pagina inferiore foliorum

brevioribus magis sparsis et semiadpressis (non patentibus) praecipue differt.

Typus: Borneo: Sabah, Beluran District, Bukit Monkobo, 15 March 1982,

Aban Gibot SAN 95227 (holo SAN; iso A nv, BO, K nv, KEP nv, L, SAR

nv, SING nv).

Diplycosia in Borneo and Peninsular Malaysia 219

Figure 1

Shrub 1-2 m tall, stems covered in coarse, brown, semi-appressed bristles

but with no obvious finer indumentum. Petiole 4-7 x 1-2 mm, with semi-

appressed bristles, becoming glabrescent with age. Leaf blade ovate to

broadly elliptic, 6-10 x 2.5-4.5 cm, glabrescent above, sparsely setose below

with variable length setae which are more or less appressed, these breaking

off to leave a black punctate surface where old and abraded, not especially

setose on the midrib; coriaceous, base rounded; margin entire, slightly

revolute at first fringed with rather fine semi-erect bristles; apex shortly

acuminate, usually with a fine acute point but occasionally rounded, the

terminal gland small and not especially conspicuous, midrib impressed

above, distinctly prominent beneath, lateral veins plane or minutely

impressed above, plane beneath, pinnate, 3-7 per side, spreading at c. 45

degrees and then arching round to link with the one above and thus

creating a rather indistinct, looping intramarginal vein. Flowers fasciculate

or solitary, 1-2 per axil; pedicels at anthesis 10-12 mm long and less than 1

mm thick, densely covered with appressed bristles that are up to 2.5 mm

long. Bracteoles c. 1 mm long, almost semi-circular, the backs and edges

with long semi-appressed to appressed bristles. Calyx c. 4 mm long with

erect, appressed bristles, tube c. 1.5 mm long, lobes c. 2.5 mm long. Corolla

c. 5.5 mm long with a slightly verrucose surface and some crimped hairs

outside, glabrous inside, lobes 3.5 mm long, triangular with broadly acute

points. Stamens c. 4 mm long, filaments sigmoid, glabrous, c. 2 mm long;

anthers c. 2.75 mm long, echinulate. Disc glabrous; ovary glabrous, style c.

3.5 mm long, cylindrical, glabrous. Fruit not seen.

Notes: This distinctive species keys out in Sleumer (1966-67) with the

larger leaved, coarsely hairy species. The type specimen does not have

any young shoots and so it is not absolutely certain that there is not a fine

puberulous indumentum in the young state but none of the stems examined

show any evidence of this. In any case this specimen does not agree with

any of the species that have this character. Diplycosia abanii appears most

closely related to D. clementium Sleumer, a common species on Mt.

Kinabalu and also recorded from N Sarawak. Both have the slender

pedicels, hairy corolla and a similar general venation pattern, but D. abanii

differs in its longer patent bristles on all conspicuous parts, rugose leaves

with much more pronounced venation, the main lateral veins on the

underside of the leaves being strongly raised. D. aurea Sleumer has

appressed bristles but these are much denser on the undersides of the

leaves and the pedicels are quite different, being much thicker, 1.5-2 mm

wide and the bracteoles are larger and more densely hairy. Bukit Monkobo

220 Gard. Bull. Singapore 54 (2002)

Figure 1. Diplycosia abanii. All from Aban 95227: A. habit x 1; B. flower x 3; C. abaxial leaf

surface x 25; D. half flower x 3; E. stamens x 5.

Diplycosia in Borneo and Peninsular Malaysia 224

is an isolated peak away from the main mountain ranges that might well be

expected to have other endemic species. The fruits are described as hairy

and brownish on the label but this undoubtedly refers to the bristly calyx

which becomes fleshy and forms the ‘false fruit’; the ovary itself is glabrous.

The collection is described as coming from “hill top mossy forest” and is

probably restricted to the small area of this type of forest about the summit

of Bukit Monkobo where it is probably an epiphyte.

2. Diplycosia mogeana Argent, sp. nov.

(Named in honour of the collector of the type material, Dr Joannis Mogea,

formerly curator of Herbarium Bogoriense).

Diplycosiae clementium Sleumer et D. abanii Argent (supra) affinis sed ab

ambabus foliorum basibus cuneatis, calyce sine setis et corolla glabra differt.

Typus: Borneo: Central Kalimantan, Bukit Raya and upper Katingan

(Mendayai) River area C. 10 km. NNW of Tumbang Tosah at camp 2, c.

112° 50’E 0° 30’°S, 1 Dec. 1982, Mogea 3814 (holo BO; iso L).

Figure 2

Semi-epiphytic shrub c. 1 m tall, stems at first covered in coarse, brown,

patent bristles but with no finer indumentum. Petiole 6-8 x c. 2 mm, with

irregularly patent bristles which often persist for some time. Leaf blade

elliptic, 6-11.3 x 2.5v3.5 cm, glabrescent above leaving the surface obscurely

punctate, sparsely setose below on the veins, the surface distinctly punctate

with hair bases and a few much larger black gland-like structures; coriaceous,

base cuneate; margin entire but with rather persistent setae along the edge,

flat or very slightly revolute, apex narrowly acute or slightly acuminate,

with a fine acute point, the terminal gland small and not especially

conspicuous, midrib very slightly impressed above and distinctly prominent

beneath, lateral veins plane above, distinctly prominent beneath, venation

pinnate with 3-7 lateral veins per side, spreading at c. 45° in the basal half

but much more widely in the upper part, each arching round to link with

the one above. Flowers fasciculate or solitary, up to 3 per axil; pedicels 2—

6 mm at anthesis, c. 1 mm thick and apparently increasing in thickness in

fruit, densely covered with semi-erect bristles that are often curved.

Bracteoles c. 1 mm diam., almost semi-circular, the backs with a few bristles

at the base and the edges glandular fimbriate. Calyx c. 4 mm, tube c. 1.5

mm long, the lobes triangular, c. 2.5 mm long, glandular fimbriate along

the edges otherwise glabrous. Corolla c. 6 x 5 mm, glabrous, lobes 2 x 2

mm with rather thick blunt points, glabrous inside, lobes 3.5 mm long,

triangular with broadly acute points. Stamens c. 4 mm long, filaments

sigmoid, glabrous, c. 2 mm long; anthers c. 2.75 mm long, echinulate. Disc

222 Gard. Bull. Singapore 54 (2002)

Figure 2. Diplycosia mogeana All from Nooteboom 4586: A. habit x 1;B. flower x 3; C. abaxial

leaf surface x 25; D. half flower x 3; E. stamens x 3.

Diplycosia in Borneo and Peninsular Malaysia 2735

glabrous; ovary glabrous, style c. 4.5 mm, swollen about the middle, glabrous.

Fruit not seen.

Notes: This species keys out in Sleumer (1966-67) to couplet 24 but the

characters do not fit either species remaining beyond this lead: Diplycosia

rufescens Schltr. from Papua New Guinea and D. triangulanthera J.J.Sm.

from Sulawesi are unlikely allies purely from a geographical point of view

as very few Diplycosia species occur on more than one island. Both of

these species have much smaller leaves and flowers than D. mogeana. Of

the Bornean species, it is most similar to D. clementium Sleumer and D.

abanii Argent (described above) but it differs from both in having cuneate

leaf bases and lacking bristles on the calyx and hairs on the corolla. The

type is described as a semi-epiphytic shrub growing in primary mossy rain

forest at 1600 m, c. 1 m tall with pale greenish flowers and the leaves very

pale beneath. The Nooteboom collection, however, is described as a treelet

3 m tall with white flowers growing at 1500 m.

Specimen examined: Borneo, Kalimantan, Bukit Raya, 112° 42’E 0°39°S,

24 Jan 1983, Nooteboom 4586 (L),

3. Diplycosia rhombica Argent, sp. nov.

(Named from the distinctive narrowly rhomboidal shape of the leaves).

Diplycosiae microsalicifoliae Argent similis sed ambitu folii rhomboideo et

pedicellis multo (plus quam triplo) longioribus, petiolis cum indumenti

vestigiis praeditis recedit.

Typus: Borneo, Sabah, Lahad Datu District, on the ridge top of Mt

Tribulation, 4° 51’N 117° 39°E, 16° Aug 1976, Cockburn SAN 84884 (holo

K; iso BO nv, KEP nv, L nv, SAR nv, SING nyv,).

Figure 3A

Small shrub to c. 60 cm tall. Branches densely leafy, grooved and angled

when dry, glabrous or with traces of a very fine, minute, whitish pubescence.

Petiole 2-4 x c. 1 mm, weakly grooved above, sometimes slightly verruculose

and with few small simple white hairs. Leaf blade narrowly rhomboid or

narrowly elliptic, (25—)35-53 x (5—)8-13 mm, somewhat coriaceous; base

narrowly attenuate; margin entire, flat and smooth or sometimes slightly

undulate; apex acute with a prominent apical gland which often forms a

mucronate tip, glabrous and smooth above, punctate with black glands

below but otherwise hairless; veins + obsolete except for the slender midrib

which is not raised above and only minutely so beneath. Flowers solitary

224 Gard. Bull. Singapore 54 (2002)

or in pairs from the leafy axils. Pedicels, glabrous, up to 16 x 0.3 mm.

Bracteoles broadly ovate, c. 8 mm long, fimbriate, with glands along the

margins. Calyx c. 2.3 mm long, lobes 0.8 mm long, deltoid, fimbriate, with

glands along the margins, and occasionally on the abaxial surface, otherwise

glabrous, lacking a conspicuous terminal gland but with a subapical swelling

on the abaxial side of each lobe. Corolla (submature) to c. 2 mm long,

pink, subglobose, glabrous, lobes c. 0.3 mm. Filaments linear, flattened,

glabrous, c. | mm long; anther cells c. 0.8 mm long, finely granular

echinulate, tubules narrow, c. 0.3 mm long. Disc glabrous; ovary glabrous

c. 1 mm long, style, c. 1mm, glabrous, slightly swollen near the middle.

Fruit not seen.

Notes: This very distinct species unfortunately lacks fully open flowers and

there are no young branches to unequivocally display the indumentum.

Most branches examined are completely glabrous, but there are traces of a

fine white indumentum in a few places on the stems and several of the

petioles have traces of a simple, short white indumentum, which is usual

only in species which have hairy stems when young. Superficially Diplycosia

rhombica looks similar to some forms of D. microsalicifolia Argent (1982)

but the very long filiform pedicels more than 10 mm long are quite different

to the short, 3-5 mm pedicels of D. microsalicifolia. It keys out in Sleumer

(1966-1967) on the basis of the stem having a fine white puberulence to D.

kemulensis J.J.Sm. but that species has much larger triplinerved leaves and

short pedicels only 4 mm long and is described as having 4-merous flowers.

If keyed as if the branches are completely glabrous, D. rhombica agrees

with neither alternative at couplet 103 as the pedicels are completely

glabrous and it is quite different from the three Bornean species beyond

this point in the key, D. urceolata Stapf, D. commutata Sleumer and D.

heterophylla Blume var. latifolia (Blume) Sleumer, as in these species all

three have prominent lateral veins in the leaves, which this new species

does not. Mt Tribulation, from which D. rhombica comes, is an isolated

ultramafic mountain both features commonly associated with endemism

and it is not surprising to find this novelty growing near the summit at c.

800 m (2,600 ft.). It is reminicent of D. sphenophylla Sleumer on Kinabalu

that also appears to be restricted or at least most common in ultramafic

areas.

4. Diplycosia mantorii Argent, sp. nov.

(Named after the collector, Asik Mantor).

Caulibus pubescentia humili rigida brevi patenti tantum instructis, foliis

usque ad 26 x 12 mm venis lateralibus fere perfecte obscuris, pilis longis

Diplycosia in Borneo and Peninsular Malaysia 225

Figure 3. Diplycosia rhombica All from Cockburn SAN 84884: A. habit x 0.8; B. abaxial leaf

surface x 25; C. flower x 3; D. half-flower x 3; E. stamens x 5. Diplycosia mantorii Type: FE.

habit x 1; G. abaxial leaf surface x 25; H. stem surface x 25; I. flower lacking stamens and

corolla x 3; J. partial l.s. section to show ovary and style x 3; K. margin of calyx lobe x 25.

226 Gard. Bull. Singapore 54 (2002)

ovarll e parte superiore exorientibus insignis.

Typus: Borneo, Sabah, Penampang District, Tunggol FH. Km 45, Jln. K.

Kinabalu, c. 6° 14°N. 116° 25°E 17 July 1986, Asik Mantor SAN 113779

(holo K; iso KEP, SAN).

Figure 3B

Climber to 4 m tall. Branches laxly leafy. Stems covered in a very short

(c. 0.02 mm long) rigid, patent, pubescence together with a few much

longer caducous, glandular hairs on the youngest stems. Petiole 1.5-2 x c. 1

mm, weakly grooved above, with the same short pubescence as the stems.

Leaf blade elliptic, 13-26 x 5-12 mm, glabrous above with a distinctly

finely undulate surface when dry, somewhat irregularly punctulate below,

puncta probably representing gland bases, coriaceous; base broadly cuneate;

margin more or less entire when mature and very slightly revolute but

finely denticulate when young with glandular hairs in the angles, these

sometimes leaving minute puncta after being shed; apex rounded, sometimes

mucronate and/or retuse, the terminal gland occasionally conspicuous but

mostly obscure,; midrib slightly depressed but mostly rather obscure above,

slightly raised and much more distinct below, lateral veins almost entirely

obscure, occasionally 1-2 traces from near the base arching upwards.

Flowers solitary, mostly from upper leaf axils; pedicels short and slender,

2-3 x 0.5 mm, covered with fastigiate glandular hairs. Bracteoles small, c.

0.5 mm wide, + semi-circular and fimbriate round the edges. Calyx c. 4mm

long, tube c. 2 mm long, glabrous, lobes c. 2 mm long, triangular with

broadly acute points, densely hairy and glandular along the margins

otherwise glabrous, without any distinctively large gland at the apex.

Corolla and stamens not seen. Disc glabrous; ovary with long but rather

sparse erect hairs on the upper half, style c. 2.5 mm long, with a few

appressed hairs near the base, otherwise glabrous. Fruit not seen.

Notes: If the anthers were known this species might key out in Sleumer

(1967) to D. consobrina Becc., which is recorded from low altitude on

Gunung Serapi (Mt. Mattang) in Sarawak, but that species has bristles on

the stems and has distinctly 3- or 5-veined leaves. Ignoring stamen

morphology, it keys out with some difficulty in both Sleumer (1967) and

Argent (1989) to D. kemulensis J.J.sm. but that species is from much

higher altitudes and has larger triplinerved leaves and a glabrous ovary.

This very distinctive new species looks superficially like D. elliptica Ridl.

but is totally lacking in the long bristles that clothe the stem in that species,

it also lacks the stipule-like axillary perulae that are usually distinctive in

D. elliptica, and D. mantorii has dense glandular hairs on the pedicels

Diplycosia in Borneo and Peninsular Malaysia 227

compared with the almost glabrous pedicels of D. elliptica. D. mantorii is

from one of the low hills rising from the coastal plain between Kota

Kinabalu and Kota Belud and although described as a climber, it is almost

certainly an epiphyte probably rooting along its stems into moss or peaty

accumulations as opportunities occur. More material with good flowers is

needed to complete the description but D. mantorii is unlikely to be

confused with any other Diplycosia species.

Another collection Argent 25108518, also from Sabah, but from much

further south at Long Pa Sia, Sipitang District, agrees very closely with the

above specimen differing only in the larger leaves up to 3 cm long. It is

also incomplete, having only old pedicels with the bracteoles still attached

although the field note states “fruits grey-blue with short pedicels”.

5. Diplycosia salicifolia Sleumer forma gigantea Argent, forma nov.

A forma salicifolia foliis multo latioribus (usque ad 5 cm latis versus 2 cm

latis) et caulibus novellissimis minute pubescentibus differt.

Typus: Malaysia, Sabah, Long Pa Sia, Sipitang District, c. 4° 26’N 115°

4A4V’E. 25 Oct. 1985 Argent & Lamb 2510859 (holo SAN; iso A nv, E, L).

Notes: The type specimen was growing with vigorous hanging stems to 3 m

long in submontane forest as an epiphyte on a tree just above the flood

zone of a river at c. 1100 m altitude. It is without flowers but has pedicels

and bracteoles in good condition. In all observable respects it agrees with

Diplycosia salicifolia except in the very much larger leaves up to 5 cm wide

(up to 2 cm in the type form) and the puberulous stems (glabrous in forma

salicifolia). The vigorous stems and broader leaves might be accounted for

by the lower altitude and different habitat from that of the type form,

which was from a ridge on Gunung Mulu between 2010-2200 m. However,

another collection from the Long Pa Sia area in the Sandakan herbarium,

Lamb & Dolois 69, from a similar altitude to this new form, is much more

similar to the typical form. These collections represent new records of D.

salicifolia from Sabah and reinforce the relationship of the flora of the

Long Pa Sia region with that of the G. Mulu and G. Murud complex in

northern Sarawak.

6. Diplycosia othmanii Argent, sp. nov.

(Named after the senior collector of the type specimen, Hj.Othman of the

Sarawak Forest Department).

Foliis majusculis ad apicem caudatis venis arcuatis supra basem exorientibus,

pedicellis brevissimis et antheris sagittatis bracchiis basalibus longis distincta.

228 Gard. Bull. Singapore 54 (2002)

Typus: Borneo, Sarawak, Ulu Sg. Mengiong, Balleh, Kapit District, 19

Oct 1988, Othman & Rantai Jawa $55787 (holo SAR; iso E, K, KEP nv.,

Ba},

Figure 4

Epiphytic shrub. Branches laxly leafy, covered in long gland-tipped bristles

and with a short, white, patent indumentum. Petiole 3-4 x c. 1 mm, with

both long glandular hairs and short, white, patent bristles. Leaf blade

broadly elliptic to ovate, 4.5-5.3 x 2.1-3.3 cm, glabrescent above, laxly

covered with rufous bristles, with dark bases beneath; leathery, base rounded

or broadly tapering, shortly contracted into the petiole, margin flat, minutely

crenulate and at first fringed with bristles, apex mucronate to shortly

caudate, the terminal gland small, major lateral veins 2—3 on each side,

curved ascending, mostly arising above the blade base, narrowly impressed

above, very slightly raised beneath. Flowers axillary, solitary or in pairs.

Pedicels very short c. 1.5 mm long. Bracteoles almost sessile, semicircular,

c. 1 mm wide, glabrous except for the shortly white ciliate margin. Calyx

up to c. 2.7 mm long, glabrous outside, lobes deltoid, c. 1 mm long, fringed

with very short, white hairs, without any trace of terminal or marginal

glands. Corolla c. 4.8 mm long, glabrous. Stamens c. 3.8 mm long; the

filaments c. 2 mm long, flattened, glabrous, expanding just above the base

and then tapering upwards; anthers 2.4 mm long, thecae c. 1.2 mm long,

dark brown, coarsely echinulate and with long basal free ‘arms’ to c. 0.5

mm long, tubules smooth, paler brown to c. 1.2 mm long, opening with a

broad oblique pore which extends to half the tubule length. Disc glabrous;

ovary glabrous, subspherical, to c. 1.6 mm, style glabrous, c. 2.4 mm long.

Fruit not known.

Notes: Similar to Diplycosia pilosa Blume from West Java at least in some

key characteristics, but D. othmanii has very different leaf venation with

its weakly raised pinnate lateral veins compared to the very distinct strongly

abaxially raised and arching lateral veins of D. pilosa, D. othmanii also

lacks the distinct revolute edge of D. pilosa. The most distinctive features

of D. othmanii are the very short pedicels, the caudate leaf apices and the

long spreading prolongations at the base of the anthers. It keys out in

Sleumer (1966-1967) with D. microphila Becc. and D. kostermansii Sleumer,

but it has much larger leaves than either of these species. Alternatively, it

would key out in Argent (1989) to D. memecyloides Stapf, but that species

has no long glandular bristles on the stems. The leaf venation, where the

main veins usually arise from well above the base of the blade, is also

distinctive.

Diplycosia in Borneo and Peninsular Malaysia 229

Figure 4. Diplycosia othmanii All from Othman & Jawa S 55787: A. habit x 1.2; B. flower bud

x 4; C. stamens x 6; D. half flower bud x 4; E. abaxial surface of the leaf x 25; K. surface of the

stein'*-25.

230 Gard. Bull. Singapore 54 (2002)

A second collection, S$ 8679, agrees well with the type although it has

no flowers. It is slightly more slender than the type with the leaves up to

21 mm wide and with less pronounced caudiform apices and broadly

tapering rather than rounded bases. It is possible that the description of

green fruits on the label actually applies to the flowers as the specimen has

no fruits. The type was described by the collector as an epiphyte at 5 m in

mixed dipterocarp forest, it is unclear whether 5 m refers to altitude or the

height up the host tree. That it grows in dipterocarp forest suggests that

this species is found at low altitudes and the Brunig collection confirms

this as it was collected at c. 15 m altitude (50 ft) in very wet kerapah

(swampy heath forest) at the foothill of a terrace in a mossy sheltered

position.

Specimen examined: Borneo, Sarawak, Bintulu District, Sungai Penyilam,

17° Feb 1961, Brunig S 8679 (SAR).

7. Diplycosia paulsmithii Argent, sp. nov.

(Named after Paul Smith, plant collector, keen observer and explorer in

SE Asia who was one of the collectors of the type specimen).

A Diplycosia aurea Sleumer foliis anguste lanceolatis plus quam triplo

longioribus quam latioribus (in D. aurea ovatis, praecipue minus quam

duplo longioribus quam latioribus), indumento abaxiali sparso fugacio,

marginibus planis, floribus plerumque solitariis, haud in fasciculos densos

agsregatis differt. |

Typus: Borneo, Sabah, Gunung Tambuyukon, Kota Marudu District, 9"

Oct 1995, Argent & Smith 395 (holo SNP; iso A, E, K, KEP, L, SAN,

SING).

Figure 5

Spreading shrub to 70 cm tall. Branches laxly leafy. Stems laxly covered

in appressed bristles. Petiole 24 x c. 1 mm, grooved above and with a few

course appressed bristles. Leaf blade lanceolate or narrowly elliptic, 50—-

100 x 12-27 mm, coriaceous, glabrous and smooth above when dry, with

rather sparse appressed bristles when very young and later punctulate

from the remaining bases below; base rounded occasionally broadly cuneate:

margin flat, entire, often with a persistent fringe of bristles, apex acute

without a prominent terminal gland, midrib narrowly and slightly depressed

above, distinctly raised below, lateral veins pinnate, arching upwards, mostly

rather obscure, although with one or two clearly distinct in larger leaves.

Diplycosia in Borneo and Peninsular Malaysia

231

;

{

Figure 5. Diplycosia paulsmithii All from Argent & Smith 395: A. habit x 1 B. flower x 3: C.

half flower x 3; D. stamens x 6; E. surface of young stem x 25; F. young abaxial leaf surface x

25; G. pedicel indumentum x 25.

2352 Gard. Bull. Singapore 54 (2002)

Flowers mostly from upper leafy axils, solitary, 2 or 3 per axil; pedicels 7-9

x c. 0.5 mm, covered with appressed bristles, 4- and 5-merous flowers on

the same plant. Bracteoles c. 1.5 mm wide, + semi-circular and fimbriate

with glandular hairs round the margins and a few long coarse bristles

abaxially. Calyx c. 2.5 mm long, tube c. 1 mm long, lobes, c. 1.5 mm long,

triangular with broadly acute points, moderately densely setose abaxially

with bristles up to 3 mm long, shortly glandular along the margins, no

distinctively large gland at the apex. Corolla white c. 4x 5 mm, tube with a

very few sparse bristles outside, the lobes deltoid with a touch of purple at

the apex, erect on opening but becoming reflexed, c. 1 x 1 mm, a distinct

cluster of small bristles on the outside. Stamens 4.6 mm long, the filaments

c. 2.1 mm long, glabrous, with anthers c. 2.5 mm long, tubules straight, c.

0.6 mm long. Disc glabrous; ovary glabrous, style, c. 3.3 mm, tapering

upwards and glabrous. Fruit not known.

Notes: this is distinctive with the fully appressed stem bristles contrasting

with the spreading bristles on the calyx. If the bristles on the undersides of

young leaves can be observed, it keys directly in Argent (1989) to Diplycosia

aurea Sleumer, but it differs both in leaf shape and in the much laxer

indumentum that is fugaceous and not persistent. The inflorescences are

very different in the two species. They are fasciculate tufts with matted

golden hairs completely obscuring the short pedicels in D. aurea. In D.

paulsmithii the flowers tend to be solitary or where fasciculate the individual

pedicels are clearly discernable as they are more slender and covered with

much less dense semi-appressed hairs. In Sleumer (1966—1967), D

paulsmithii keys out with the bristly species but at couplet 4, it does not

agree with either lead as the bristles on the calyx are up to 3 mm long but

they point in all directions and are clearly not appressed or suberect. Having

bristles on the underside of the leaf (before they fall off) that are appressed,

it again keys out to D. aurea. The second half of couplet 4 leads to D.

lilianae J.J.Sm. from West New Guinea and D. loheri Merr. from the

Philippines, both unlikely candidates given the local distributions of most

montane Diplycosia species. D. paulsmithii, however, differs from D. lilianae

as the bristles are not gland-tipped, and D. paulsmithii lacks the fine white

hairs on the pedicels characteristic of D. loheri. 3

Diplycosia paulsmithii is relatively common on the upper part of G.

Tambuyukon where it grows in semi-open areas amongst the ultramafic

rocks, often as an epiphyte. It may well be endemic to this interesting

mountain.

Specimens examined: Gunung Tambuyukon, camp III to summit Nais et al.

SNP 4798, 5" Oct 1990 (KEP, SNP); cultivated specimen, the same locality,

Diplycosia in Borneo and Peninsular Malaysia 239

flowering 20° Feb 2002 Accession No. 19952784 (E, SAN, SNP).

8. Diplycosia ngii Argent, sp. nov.

(Named after the collector Dr, Francis $.P.Ng, sometime Deputy Director

General, Forest Research Institute Malaysia, Kepong, who has made an

outstanding contribution to botany in Malaysia).

Diplycosiae pseudorufescenti similis sed indumento tenui albo sub setis

grossis brunneis carenti, calyce abaxialiter breviter piloso (haud omnino

glabro), foliis majoribus et pedicellis longioribus differt. "

Typus: Peninsular Malaysia, Pahang, Gunung Jasar, summit area, 23

Aug 1977, Ng FRI 27145 (holo KEP).

Figure 6

Shrub to 2 m tall. Branches laxly leafy. Stems terete, densely and subpatently

setulose, covered with rufous semi-appressed bristles of differing lengths,

without any finer pubescence. Petiole appressed setulose, 2-4 x 1-1.5 mm.

Leaf blade ovate to ovate-elliptic, 23-35 x 12-19 mm. Young leaves densely

appressed rufous-setulose on both sides, the mature leaves becoming

glabrous above but mostly remaining appressed setulose below, finally in

old leaves all the bristles are denuded to give a punctate surface, coriaceous;

base rounded or more rarely broadly tapering; margin more or less revolute,

at first with small bristles but these mostly early caducous to leave it

minutely crenulate with impressed bristle bases; apex shortly acuminate,

occasionally acute, terminal gland obscure, midrib impressed above,

prominent beneath, lateral veins mostly two, basal or supra-basal, ascending,

usually becoming obscure in the lower half of the leaf and before they

reach the margin, occasionally visible to just over half way, rather obscure

abaxially. Flowers axillary, solitary or paired pedicels c. 3 mm at flowering,

later up to c. 6 mm long when in fruit, puberulous with crisped hairs, basal

bracts, several, minute. Bracteoles c. 2 mm across, ovate, with involute

sides and so appearing acute, shortly hairy abaxially. Calyx to c. 3 mm

long, shortly hairy abaxially, lobes c. 2 mm long, acute with involute margins.

Corolla tubular campanulate, glabrous, c. 4 mm long, lobes reflexed, to c.

0.5 mm long. Stamens c. 3.5 mm long, filaments slender, sigmoid, glabrous,

c. 1.5 mm long, anthers c. 2.5 mm long, the cells c. 1.5 mm long, tubules c. 1

mm long. Disc glabrous; ovary rather densely grey-pubescent with erect

hairs, style c. 3.5 mm long, glabrous. Fruit not known.

Notes: Diplycosia ngii keys out in Sleumer (1966-67) to D. pseudorufescens

Sleumer, but differs from this species in not having the fine white patent

puberulous tomentum under the coarse brown setose hairs, the leaves are

234 Gard. Bull. Singapore 54 (2002)

Figure 6. Diplycosia ngii All from Ng 27145: A. habit x 1; B. half flower x 9; C. flower x 9.

Diplycosia in Borneo and Peninsular Malaysia 239

larger and the apical gland is not distinct, the calyx is shortly hairy abaxially

(not glabrous) and the pedicels become much longer, up to 6 mm in fruit

whereas in D. pseudorufescens they are not more than 3 mm. Type

specimens of both subspecies of D. pseudorufescens have been examined

but it has not been possible to locate the specimen collected on Gunung

Korbu (Kerbau), Perak (Sleumer, 1966-67). Sleumer implied that this

collection has the same characteristics of indumentum as the Kinabalu

plants, the type locality of both subspecies of D. pseudorufescens. Clemens

29282 from Mt. Kinabalu is described with fruit, which suggests a real

difference in the final pedicel length between D. pseudorufescens and D.

ngil. The indumentum of bristles on the stems is reminiscent of D.

sumatrensis Merr. from Sumatra (Indonesia) but both leaves and flowers

are much smaller in this new species, which at present is only known from

the type collection.

Reduction of Pernettyopsis to Synonymy with Diplycosia.

Pernettyopsis was described by King & Gamble (1905) with two species (P.

malayana King & Gamble and P. subglabra King & Gamble) from Penisular

Malaysia. The second species was later reduced to synonymy by Sleumer

(1966-1967) along with a third (P. breviflora (Ridl.)Ridl., which Ridley

(1915) had originally described as a species of Diplycosia! Sleumer stated

that “the wooly indumentum on the calyx lobes is apparently detersile and

early gone and the length of the pedicels varies” and concluded that there

was material of only one species. Pernettyopsis was described as different

from Diplycosia in that the fruit was a berry, the calyx was unchanged in

fruit, 1.e. Was not accrescent and succulent, and was about as long as the

corolla. A further species, P. megabracteata Argent, was described from

Borneo in 1982. This latest species technically agreed with the description

of Pernettyopsis but was very different to the original species having

extraordinarily long acuminate bracteoles and very differently shaped

flowers: ovoid cylindrical and more than twice as long as broad in P.

megabracteata; short urceolate and hardly longer than broad in P. malayana.

The distinction between the fruits had already been questioned

(Argent, 1982) and the definition of Diplycosia fruits as thin-walled capsules

(Sleumer, 1966-1967) cannot be maintained. D. acuminata in Peninsular

Malaysia clearly has an ovary that becomes enlarged and fleshy and behaves

as a true berry. The calyx lobes of this species, although also accrescent,

are much less so than in most other Diplycosia species and show a clear

transition to the state in Pernettyopsis. Of many other species observed in

the wild, the ovary never appears to split naturally and the fruit behaves

236 Gard. Bull. Singapore 54 (2002)

like a false berry although most of the ‘attractive’ flesh is associated with

the calyx. That the calyx lobes are as long as the corolla in the two

Pernettyopsis species would appear to be of no great significance as the

flowers are of totally different shapes. Some of the very bristly Diplycosia

species in Borneo, such as D. rufa Stapf, D. aurea Sleumer and D. hirtiflora

Argent, also have long calyx lobes. There thus seems little point in

maintaining the genus Pernettyopsis that is so weakly supported

morphologically. Of the two currently recognized species, the original one

(sensu Sleumer, 1966-1967) is frequently confused with Diplycosia and the

second, although superficially very distinct from all other known Diplycosia

species (Argent, 1982), is at least as different from the original P. malayana.

Both these Pernettyopsis species share all the important structural

characteristics of inflorescence and flower with Diplycosia. We do not

have the anatomical evidence of Middleton (1989) but the conclusions

drawn are parallel to those of Middleton & Wilcock (1990) to justify the

inclusion of the genus Pernettya Gaudich. within Gaultheria Kalm ex L.

Thus the genus Pernettyopsis is formally synonymised below with two

resulting new combinations.

New Combinations

Diplycosia Blume, Bijdr. (1826), 857. Type species: D. heterophylla Blume,

Bidr 858 (1826).

Synonym: Pernettyopsis King & Gamble, J. As. Soc. Bengal, 74, 2 (1905)

79.

Type species: P. malayana King & Gamble, J. As. Soc. Bengal, 74, 2 (1905)

72:

Diplycosia malayana (King & Gamble) Argent, comb. nov.

Basionym: Pernettyopsis malayana King & Gamble, J. As. Soc. Bengal, 74,

2.44905 ) 9:

Lectotype: Scortechini 402, Peninsular Malaysia, Gunung Batu Puteh.

Synonyms: Pernettyopsis subglabra King & Gamble, J. As. Soc. Bengal,

74, 2 (1905) 79.

Type: Scortechini 752, Perak.

Diplycosia breviflora Ridl., J. Fed. Mal. St. Mus. 6 (1915) 158.

Type: Ridley s.n. Pahang, Gunung Tahan.

Pernettyopsis breviflora (Ridl.) Ridl., Fl. Mal. Pen. 2 (1923) 222.

Diplycosia in Borneo and Peninsular Malaysia 257

Diplycosia megabracteata (Argent) Argent, comb. nov.

Basionym: Pernettyopsis megabracteata Argent, Bot. J. Linn. Soc. 85 (1982)

10-12 f. 4. (1982).

Type: Argent 824, Sarawak, Gunung Mulu.

Acknowledgements

The research programme at the Royal Botanic Garden Edinburgh is

supported by the Scottish Executive Committee and Rural Affairs

Department. I thank the curators of the following herbaria who have

loaned specimens: Bogor, Harvard, Kew, Kuching, Leiden and Sandakan.

I am grateful to Christina Oliver for the illustrations and to Dr Robert Mill

for translating the Latin diagnoses and for very helpful comments.

References

Argent, G.C.G.1982. New taxa and combinations and comments on

Ericaceae. Botanical Journal of the Linnean. Society. 85: 1-17.

Argent, G.C.G. 1989. Diplycosia in Borneo. Notes Royal Botanic Garden

Edinburgh. 46: 17-26.

Beaman, J.H., C. Anderson C. and R.S. Beaman 2001. The Plants of

Mount Kinabalu. 4, Dicotyledons families Acanthaceae to Lythraceae.

Natural History Publications (Borneo), Kota Kinabalu, Malaysia.

King, G. and J.S. Gamble. 1905. Pernettyopsis. Journal of the Asiatic Society

of Bengal. 74: 11 79-80.

Kron, K. 2001. An analysis of the phylogenetic relationships in the

wintergreen group (Diplycosia, Gaultheria, Pernettya, Tepuia; Ericaceae).

Systematic Botany. 26: 808-817.

Middleton, D.J. 1989. Taxonomic Studies in the Genus Gaultheria L. and

Related Genera (Ericaceae). Unpublished Ph.D thesis, University of

Aberdeen, Scotland, U.K

Middleton, D.J. and C.C. Wilcock, 1990. A critical examination of the

status of Pernettya as a genus distinct from Gaultheria. Edinburgh Journal

of Botany. 47: 291-303.

Ridley, H.N. 1915. The Botany of Gunung Tahan. Journal Federated Malay

States Museum. 6: 158.

238 Gard. Bull. Singapore 54 (2002)

Sleumer, H. 1966-67. Ericaceae. Flora Malesiana. 6: 469-914.

Wong, K.M. & A.Phillipps 1996. Kinabalu, Summit of Borneo. The Sabah

Society in association with Sabah Parks, Kota Kinabalu, Malaysia.

Gardens’ Bulletin Singapore 54 (2002) 239-242.

New Gesneriaceae: a Chirita from Vietnam and a

Monophyllaea from Sulawesi

BLL, BURTE

Royal Botanic Garden Edinburgh,

Edinburgh EH3 5LR, U.K.

Abstract

Chirita bogneriana B.L.Burtt is described from Ha Long Bay, Vietnam and Monophyllaea

chinii B.L.Burtt from Maros, Sulawesi.

Chirita bogneriana

A few years ago, living material of a species of Chirita (Gesneriaceae) was

sent to the Royal Botanic Gardens Edinburgh by Josef Bogner (Munich

Botanic Garden, Germany) for identification, by no means the first

interesting gesneriad that we have received from him. The plant originally

came from Ha Long Bay in northern Vietnam. I was unable to identify this

plant, but, learning of the survey being made there by Nguyen Tien Hiep

and Ruth Kiew, I decided to wait and see if they too had collected this

plant. Their results have now been published (Nguyen and Kiew, 2000)

and they record seven species of Chirita: one of them is C. hamosa R. Br.,

which belongs to the distinctive section Microchirita; the remainder are

referable to section Gibbosaccus, as is the Munich plant: but, although

three new species were recognized, none of the species found by them fits

this, which is accordingly described here as C. bogneriana.

The following description is based on a well-grown pot-plant. It is

likely that it was larger and more floriferous than would be found in the

wild, where the species almost certainly grows in crevices of limestone

rock.

Chirita bogneriana B.L. Burtt, sp. nov.

(Sect. Gibbosaccus C.B.Clarke) |

Typus: Vietnam, Ha Long Bay, spec. cult. in RBG Edinburgh (ex Bot.

Gart. Muenchen comm. J. Bogner) under no. 19990640 (holo E; iso HN,

K, M, SING).

Chirita hiepii Kiew (species in sinu Ha Long etiam incola) affinis sed foliis

superne pilis brevibus inter se distantibus (nec pilis 4—6 mm longis

240 Gard. Bull. Singapore 54 (2002)

praecipue in marginibus) indutis; corollae limbo albo (nec purpureo), tubo

ad basin laete luteo-viridi glabro (nec lineis duabus glanduloso-pilosis

instructo) distinguenda.

Rosette herb with numerous ascending leaves and a vertical rootstock 2.5

cm diam. at top and rather abruptly narrowed at base. Leaves (including

petiole) up to 10 cm long and 3 cm broad, lanceolate, gradually tapering

into the 3.5 cm long flat petiolar part, the blade shortly pubescent above,

the hairs well spaced (not overlapping) slightly longer and distinctly denser

and appressed below, margin entire very slightly revolute, veins 4—5 on

each side of midrib scarcely visible above, prominent below ascending at

angle of 30°. Inflorescences compound pair-flowered cymes, solitary in the

leaf axils, common peduncle 9—12 cm then branched, branches subtended

by linear bracts 8 mm long, bracts and peduncle and whole inflorescence

pilose with white gland-tipped hairs; branches usually branched again; whole

inflorescence (in cultivated plant) 20—30 cm long, pedicels slender up to 6

cm long. Calyx tube 0.5 mm, lobes 8 x 1.2 mm, on the inside with minute

globular glands and short (0.3 mm) acute hairs, on the outside with patent

gland-tipped hairs 1—1.8 mm long. Corolla limb white, lobes glabrous

inside, anterior lobes 6.5 x 5 mm, posterior 6 x 6 mm; tube | cm long, it

and the outside of the lobes glandular-puberulous. Stamens arising 2.5 mm

above base of corolla; the anterior pair fertile 5 mm long, rather thick,

glabrous except for a few gland-tipped hairs near the connective, anthers

3.2 mm with widely divergent thecae; posterior pair 3 mm, sterile; dorsal

staminode 0.5 mm. Ovary 6 x 1.3 mm, passing smoothly into 3 mm rather

thick style, both with short hairs. Stigma with lower lobe bifid, upper

aborted. Fruit not seen (seed is apparently not set on these plants).

Notes: Chirita bogneriana is most closely allied to C. hiepii Kiew, also from

Ha Long Bay, but lacks the 4—6 mm long hairs on leaf margin and petiole

which are a feature of that species. It also lacks the deep violet corolla

lobes of C. hiepii, and the corolla tube is glabrous inside without the pair

of glandular hairy bands of C. hiepii. From C. halongensis Kiew and T.H.

Nguyen, it differs in the leaves being shortly pubescent, not glabrous, above

and the veins on the under surface being distinctly prominent. C. halongensis

shares with C. hiepii violet corolla lobes whereas the flowers of C.

bogneriana are white, with just a greenish tinge at the base of the tube.

Monophyllaea chinii

This new species of Monophyllaea grows on limestone cliffs near Maros in

New Gesneriacceae 244

SW Sulawesi. It was first collected in 1986 by Dr S.C. Chin, well known for

his revision of M.R. Henderson’s account of the limestone flora of the

Malay Peninsula. Ten years later, a second collection from the same general

area, but from a different locality, was made by Dr. W. de Wilde and Dr B.

de Wilde-Duyfjes. M. chinii belongs to subgen. Monophyllaea and is the

first record of that subgenus from Sulawesi, the other monophyllaeas

previously found on Sulawesi all belong to subgen. Moultonia.

Within subgen. Monophyllaea, M. chinii is to be associated with a

small group of delicate species that has a rather scattered distribution and

comprises species 13-16 in the revision of the genus (Burtt, 1978). The

species are M. glabra Ridl. from S Thailand and N Malaysia (Langkawi

Islands), M. hirticalyx Franch. from C. Malaysia (Perak and Selangor), M.

musangensis A. Weber (S. Kelantan — after the revision), M. tenuis B.L.Burtt

from the First (Kuching) Division of Sarawak, and the ill-known M. longipes

Kraenzl. from the Philippine Island of Luzon.

The new Monophyllaea chinii is almost certainly an annual, as is M.

glabra, which comes from an area with a distinctly seasonal climate. M.

tenuis, certainly, and M. hirticalyx almost certainly, last for more than one

flowering period. There remains much to be learnt about the duration of

Monophyllaea and its control.

Although I have classified these delicate plants close to one another,

it remains to be discovered whether they are really interrelated or whether

each is more closely akin to more robust species in its own neighbourhood.

Monophyllaea chinii B.L.Burtt, species nova ex affinitate M. hirticalycis

Franch. et M. tenuis B.L.Burtt sed plantis celebicis multo minoribus ut

videtur annuis, caulibus pedunculis pedicellis pilis 1.5—2 mm longis indutis,

pedicellis 8—12 mm longis patentibus, pilis in calyce 1.5 mm longis, interdum

glandulosis distinguenda.

Typus: SW Sulawesi, Maros, Leangleang Prehistoric Park S.C. Chin 3408,

8 June 1986 (holo E; iso L) ’limestone hills, on vertical limestone cliff,

flowers pale purple or white; small plants, longest leaf to 15 cm, stem to 15

’

cm .

Stem (hypocotyl) slender, c. 3—10 cm long, pilose with patent hairs 2 mm

long. Leaf solitary (macrocotyledon) 5—12 [—15 cm fide Chin] x 3—7.5

cm, more or less ovate, cordate at the base, thinly shortly hairy above, but

with long hairs around the base of the peduncles which arise at the base of

the midrib; lower veins matching the leaf outline, the lower ones strongly

curved the upper ones straight. Peduncles 1—3, pilose with spreading hairs.

Bracts absent. Inflorescence (incl. peduncle) up to c. 7 cm long, bearing

242 Gard. Bull. Singapore 54 (2002)

about 4—6 pairs of flowers, the lowermost pairs c. 1 cm apart. Pedicels

strongly patent, 8—12 mm long. Calyx with tube 1 mm long, lobes 5

subequal c. 2.2 x 1.3 mm with slender hairs on the back c. 1.5 mm long

some of them gland-tipped. Corolla bilabiate 10 mm long, tube 5 mm;

lower lip 6.5 mm across minutely papillose inside, median lobe 3.5 x 3 mm,

anterior laterals 3 x 3 mm, posterior 0.5 x 2.2. mm, glabrous; tube 5 mm

long with a ring of hairs all round the inside just below the base of the

filaments. Stamens 4, the fifth (dorsal) represented by a minute staminode;

anterior pair with filaments 2.5 mm and anthers 0.8 mm long, posterior

pair with filaments 1.5 mm and anthers 1 mm long, all filaments with a

minute projection just below the anther, all anthers coherent. Disc annular

1.2 x 0.3 mm margin lobulate. Ovary conical 1 x 0.8 mm narrowed into 2

mm style, both glabrous; stigma 0.2 mm, slightly oblique, papillate. Capsule

1.5 x 1.5 mm, 2-(?finally 4-)valved, brown. Seeds 0.3 x 0.2 mm, light brown,

coarsely reticulate.

Additional material: S W Sulawesi. Maros, NE of Wong Padang, c. 15 m

alt., W. de Wilde & B. de Wilde-Duyfjes 21847 15 July 1996 (E, L) ‘limestone

rock area, on rather dry half-shaded rock, locally gregarious. Plants rather

palish green, more or less juicy; flowers pale lilac.’

Acknowledgements

The floral dissections and sketches by O.M. Hilliard are acknowledged

with thanks.

References

Burtt, B.L. 1978. Studies in the Gesneriaceae of the Old World. XLV: A

preliminary revision of Monophyllaea. Notes Royal Botanic Garden

Edinburgh. 37: 1-59. —

Nguyen T.H. and R. Kiew. 2000. New and interesting plants from Ha Long

Bay, Vietnam. Gardens Bulletin Singapore. 52: 185-201.

Gardens’ Bulletin Singapore 54 (2002) 243-246.

A New Species of Monophyllaea (Gesneriaceae) from

Borneo

R. KIEW

Singapore Botanic Gardens,

Singapore 259569

Abstract

Monophyllaea burttiana Kiew is described from Sabah, Malaysia.

Introduction

Since Burtt’s 1978 monograph on the genus, which described 32 species

from Malesia, a further three species have been described from Peninsular

Malaysia (Weber, 1998). There are undoubtedly more species to be

discovered in remote parts of the region. Monophyllaea burttiana Kiew

(described below) is the second species to be recorded from Sabah.

Monophyllaea in Sabah

Unlike Sarawak, where there is a wealth of Monophyllaea species (14

species), until now only one species, the widespread M. merrilliana Kraenzl.

was recorded from Sabah. Indeed, while in Sarawak monophyllaeas are a

characteristic feature of limestone hills, Sabah is in marked contrast with

all except two of the 60 odd hills being devoid of them. M. merrilliana only

occurs on limestone in the Tabin Wildlife Sanctuary in eastern Sabah,

where it does not grow on cliff faces but instead on and among the scattered

limestone boulders in the damp, deep shade around the hill (Kiew, 2001).

In Sabah, it is more common and has been collected from non-limestone

substrates, e.g. from Gunung Kinabalu, Gunung Trusmadi and the Danum

Valley.

In contrast to Monophyllaea merrilliana, this new species covers a

vertical limestone cliff face of the unique below-ground, bowl-shaped gorge

at Batu Urun. It is immediately distinguished from M. merrilliana by its

uniseriate (not branched) hairs and unbranched inflorescences although it

does belong to the same subgenus Moultonia in possessing anchor-shaped

placentas and non-valvate fruits.

Among the species in subgen. Moultonia, most of which occur in

244 Gard. Bull. Singapore 54 (2002)

Borneo, it keys out with M. glauca C.B. Clarke in having unbranched

inflorescences at the base of the midrib and a short way up it, in having

unbranched hairs, and in the fruit where the style is not exerted beyond

the calyx. It is distinct from this species in its shorter peduncle (up to 11

cm long as opposed to 15—25 cm long in M. glauca), which is three times

longer than the flowering part (as opposed to only a fifth or less in M.

glauca), as well as its longer calyx, glabrous corolla lobes, larger anthers,

and leaf base that is only slightly cordate.

Observations in the field show that the differences between the two

subgenera are strongly correlated with seed dispersal. As Burtt (1978)

pointed out, the fruit of subgen. Monophyllaea with long pedicels and four

valves are censer fruits where the seeds are dispersed when shaken out by

wind or other means. On the other hand, the fruits of subgen. Moultonia

are typical splash cups, a mode of dispersal common to many other species

of rainforest understorey herbs (Kiew, 1988), such as species of Argostemma

(Rubiaceae), Epithema (Gesneriaceae), Phyllagathis and Sonerila

(Melastomataceae), in having short erect pedicels, calyx lobes that open to

about 45° and the top of the pericarp that falls off to expose the minute

seeds in the splash cup. In this new species, the style base is inflated and

hollow and breaks down to expose the seeds. The ballistic force of large

water drops dripping through the canopy can bounce the seeds several

meters from the parent plant (Savile and Hayhoe, 1978). When dispersal

does not occur, the seeds germinate within the open splash cup.

Monophyllaea burttiana Kiew, sp. nov.

A Monophyllaea glauca C.B.Clarke inflorescentiis brevioribus, ad 11 cm

tantum (nec 15—25 cm), corollae lobis glabris et folio non nisi leviter

cordato differt.

Type: Borneo: Sabah — Pensiangan District, Batu Urun (lat 4’ 48”N long

116’ 38”E) J.J. Vermeulen 2100 16 Feb 2001 (holo SING, iso E, L, SAN).

Stem fleshy, up to 35 cm long (not encrusted). Leaf pendent, c. 22—23 cm

long but much decomposed at the tip, broadly ovate and up to 26 cm wide

or broadly oval and c. 22 cm wide, base slightly cordate, thin, slightly

rough and densely hairy above, less hairy beneath, hairs uniseriate c. 1 mm

long and raised on a hemispherical base, veins plane above, slightly raised

beneath. Inflorescences arising from the base of the midrib and c. 3.5 cm

up the midrib, unbranched with green peduncle (2.5—)11 cm long, densely

pilose with uniseriate glandular hairs, flowering part scorpioid with c. 4

rows of flowers, upto 4.5 cm long, not persistent; bract pair reflexed c. 3 x 2

Monophyllaea burttiana from Borneo 245

mm; pedicel pilose, erect, 3—5 mm long. Calyx c. 8 mm long, deep purple

or purple brown, densely pilose outside, tube c. 1.5 mm long, lobes

isomorphic, apex acute. Corolla c. 9 mm long, 4 mm wide, white with

green upper lobes and yellow patch in the throat, tube c. 4.5 mm long with

a ring of sparse hairs inside about midway up the tube, hairs extending

towards upper lobes, lobes glabrous, upper lobes strongly reflexed, c. 3.5 x

3.5 mm, lateral and median lobes isomorphic, c. 4.5 x 3.5 mm. Stamens

with green filaments, both pairs arising c. 2.5 mm from base of corolla

tube, anterior c. 3 mm long and curved, posterior c. 1.5 mm and straight,

anthers yellow, c. 1 mm diam., connate and filling the mouth of the corolla

tube. Disc c. 0.5 mm high. Ovary c. 1 mm long, glabrous; style c. 2 mm

long, placentas anchor-shaped in TS. Calyx closes over ovary after

pollination, opening with lobes at 30° on fruit maturity and forming a

splash cup. Pedicel lengthening to 11 mm. Fruit globose, c. 1.5 x 1.25 mm,

pericarp thin topped by hollow inflated style base, style c. 2.5 mm long

enclosed in the calyx in developing fruit, caducous in mature fruit. Seeds

many, ellipsoid, tiny c. 0.3 mm long, smooth and jet black.

Distribution: Endemic to Sabah, Borneo, known only from Batu Urun.

Habitat: Deeply shaded limestone cliff face at c. 600 m altitude.

Notes: This species is extremely local being known from a single population

that covers a rock face measuring about 20 by 15 m.

The species is named for B.L. Burtt, who for a life-time has worked

assiduously on the genus and the family and been an inspiration to others.

Other specimens examined: Lamb B85 (E), Kiew RK 4444 20 May 1997

(SAN, SING).

Acknowledgements

The author is indebted to Anthony Lamb and his staff at the Tenom

Orchid Centre for taking us to the inaccessible Batu Urun, to the staff at

the Forest Research Centre, Sandakan for their expert help in the field

and to B.L. Burtt for improving the latin diagnosis.

246 Gard. Bull. Singapore 54 (2002)

References

Burtt, B.L. 1978. Studies in the Gesneriaceae of the Old World. XLV: A

preliminary revision of Monophyllaea. Notes Royal Botanic Garden

Edinburgh. 37: 1-59.

Kiew, R. 1988. Herbaceous flowering plants. In: (ed.) E. Cranbrook. Key

Environments: Malaysia. pp. 56-76.

Kiew, R. 2001. Towards a limestone flora of Sabah. Malayan Nature

Journal. 55: 77-93.

Savile, D.B.O. and H.N. Hayhoe. 1978. The potential effect of drop size on

efficiency of splash-cup and springboard dispersal devices. Canadian

Journal Botany. 56: 127-128.

Weber, A. ‘1997’ published 1998. New taxa of Monophyllaea (Gesneriaceae)

from Peninsular Malaysia. Beitraege Biologie Pflanzen. 70: 405-406.

Gardens’ Bulletin Singapore 54 (2002) 247-251.

A New Species of Vatica (Dipterocarpaceae) from

Peninsular Malaysia

L.G. SAW

Forest Research Institute Malaysia,

Kepong, 52109 Kuala Lumpur, Malaysia

Abstract

A new species of Dipterocarpaceae, Vatica yeechongii L.G. Saw is described from Peninsular

Malaysia. The species is currently known from only two localities: the type locality at

Sungai Lalang Forest Reserve, Selangor, and the Setul Forest Reserve, Negri Sembilan.

Introduction

The genus Vatica comprises about 65 species (Ashton, 1982) distributed in

Sri Lanka, southern and eastern India, Myanmar, Thailand, Indochina,

south China and through Malesia. In the most recent revision for the

genus for the Malesian region, Ashton (1982) recognised 21 species in

Peninsular Malaysia. Based on fruits, the genus can be divided into two

sections, viz., sect. Vatica (with fruit calyx lobes equal) and sect. Sunaptea

(with fruit calyx lobes unequal). In the recent mast flowering in Peninsular

Malaysia (mid-2002), a Vatica was collected from the Sungai Lalang Forest

Reserve that proved to be a new species belonging to sect. Vatica.

The species has unusually large leaves and the fruits are similar to

those of Vatica havilandii Brandis, V. venulosa Blume and V. chartacea

P.S. Ashton but its size, the leaves and fruits are much larger than those of

these three related species (Table 1).

Vatica yeechongii L.G. Saw, sp. nov.

Vaticae venulosae similis, sed foliis crasse coriaceis supra bullatis

oblanceolatis 44-84 cm longis 10-16.5 cm latis, basi anguste contracta in

basi cordata, acumine 1.5-2.5 cm longo, calycis lobis 3.5—4 cm longis, et

nuce ovoidea 15-18 mm diam. differt. Typus: Peninsular Malaysia, Selangor,

Sungai Tekala Recreational Forest in Sungai Lalang Forest Reserve, fr. 11

July 2002, Chung FRI 40344 (holo. KEP; iso. A, K, L, KLU, SAN, SAR,

SING).

248 Gard. Bull. Singapore 54 (2002)

Table 1. Comparison of Vatica yeechongii, V. chartacea, V. havilandii and V.

venulosa.

V. yeechongii V.chartacea V.havilandii V. venulosa

Leaves:

petiole

length (mm) 20-30 10-22 10-12 5-9

blade

texture thickly coriaceous, thinly chartaceous thinly coriaceous — thinly coriaceous

bullate above

shape oblanceolate oblong to obovate narrowly oblong to elliptic to ovate-

obovate lanceolate

length (cm) 44-84 11-25 8-17 4-12

width (cm) 10-16.5 3-10 2.5-5 1.5-5

base tapering narrowly broadly cuneate cuneate cuneate

to acordate base or obtuse

acumen length 1.5-2.5 1 1 up to 0.5

(cm)

no. of secondary 28—30 pairs 16-20 pairs 15-20 pairs 7-12 pairs

veins

Fruits pedicel

length (mm) 2-3 up to 6 up to 5 up to 2

Calyx lobes

length (cm) 3.54 up to 6 up to 2.5 up to 3

width (cm) 1-1.5 up to 1.5 up to 1.5 up to 1.3

shape ovate lanceolate ovate ovate

Nut

shape ovoid ellipsoid globose globose

diameter (mm) 15-18 up to 11 up to 12 up to 10

Figure 1

Small tree, 8-15 m tall, 9-13 cm diam., without buttreses. Bark smooth

with horizontal rings, greyish white with lichen patches; inner bark yellowish-

brown; sapwood pale, hard, exuding creamy white resin when cut. Twigs

robust, 1-2 cm diam., densely rufous stellate hairy when young. Stipules

narrowly triangular in shape, often curving to one side, 20-35 x 5-8 mm,

subpersistent. Leaves spreading horizontally to slightly drooping, thickly

coriaceous, bullate above, glossy dark green above, pale green below when

fresh, glabrescent; petiole 20-30 mm long, 9-13 mm thick, swollen, cracking

when dried, densely rufous stellate hairy; blade oblanceolate, 44-84 x 10-

16.5 cm, tapering narrowly to a cordate base, margin entire, apex acuminate,

acumen 1.5-2.5 cm long; midrib very stout and prominent below, raised

above; secondary veins 28-30 pairs, prominent below, distinctly raised

Vatica yeechongii

Figure 1. Vatica yeechongii. A. leafy fruiting twig; B. details of lower surface of leaf; C. fruit; D.

fruit with two calyx lobes removed. (All from Chung FRI 40344)

250 Gard. Bull. Singapore 54 (2002)

above; intermediate veins 2—4, with the middle one longer than the rest, 3—

3.5 cm from the midrib; tertiary veins scalariform-reticulate, conspicuous

above; linear domatia sometimes present along distal portion of secondary

veins at junction joining preceding veins. Flowers unknown. Infructescence

axillary, 1 per node, along branches of the leafy shoots, branching to only 1

order, short, densely rufous stellate hairy; rachis 4-8 cm long, c. 2 mm

thick at base and tapering to c. 1 mm thick apically. Fruits along main

rachis and on first order branches; pedicel 2-3 mm long, c. 1 mm thick,

densely rufous stellate hairy; calyx lobes green to reddish brown when

fresh, 3.5—4 x 1-1.5 cm, ovate, acute, connate up to 4-5 mm from the base,

glabrescent; nut ovoid, 15-18 mm in diameter, glabrescent, completely

hidden in, but free from, the calyx lobes. Germination epigeal.

Distribution: Endemic to Peninsular Malaysia. Rare, known from the Sungai

Lalang Forest Reserve, Selangor, and the Setul Forest Reserve, Negri

Sembilan.

Habitat: It was found in lowland dipterocarp forest, at about 50 m altitude,

on a gentle slope near the riverbank in the Sungai Lalang Forest Reserve

in association with Saraca cauliflora Baker (Leguminosae). The fruits appear

to be water dispersed. Of the nine mature trees the Forest Research Institute

Malaysia (FRIM) team managed to locate, all were within 5 m of the

stream bank, where the gradient of the stream is gentle. We were unable

to locate trees on the upper slopes of the river valley nor on the banks

further upstream where the stream is torrential. In the Setul Forest Reserve,

the species was also observed in a similar habitat.

Notes: The growth pattern on shoots is rhythmic: on the stem there are

obvious resting periods where nodes are close together, followed by

elongation periods of leaf-free growth with only stipules, then followed by

a cluster of spirally arranged leaves. Largest leaves are found on the

proximal ends of these clusters, the distal ones are smaller.

Chan Yee Chong, FRIM Research Assistant, had noted this species

as an unnamed Vatica sometime ago and during the recent mast flowering

and fruiting, he managed to collect fruiting material, which enabled the

description of this new species. The species epithet honours Chan Yee

Chong.

Other specimen examined: Peninsular Malaysia — Selangor: Sungai Tekala

Recreational Forest in the Sungai Lalang Forest Reserve, close to ranger’s

office and camping site, fr. 3 July 2002, Y.C. Chan FRI 46657 (KEP); Negri

Vatica yeechongii Zot

Sembilan: Setul Forest Reserve, fr. 20 Aug 2002, Y.C. Chan FRI 46668

(KEP).

Acknowledgements

I thank Azid Adam, Mohd. Jantan, Borhan Mat Saad and Abu Kasim

Omar, Selangor District Office, Cheras, Kuala Lumpur; Peter S. Ashton

for advice on the affinity of this new species; J. F. Veldkamp, Nationaal

Herbarium Nederland, for the Latin diagnoses; R. C. K. Chung, FRIM; E.

Soepadmo for help in editing the final draft; Tetriana Ahmed Fauzi for the

line drawing; Damahuri Sabri, Mohd. Aidil Noordin, Fakhrul Effendi

Othman and Mustapa Data, FRIM, for their help in the field; and especially

to Chan Yee Chong who first pointed out the new species and made the

first collection. Support for the research came from the Flora Malaysiana

Centre Fund, Ministry of Primary Industries, Malaysia under Project 6:

Conservation Monitoring System for Threatened Plants.

Reference

Ashton, P. S. 1982. Dipterocarpaceae. Flora Malesiana. Ser. 1, 9: 237-552.

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Gardens’ Bulletin Singapore 54 (2002) 253-255.

The Identity of Swietenia alternifolia (Mill.) Steud.

(‘Meliaceae’)

D.J. MABBERLEY

Nationaal Herbarium Nederland, University of Leiden, The Netherlands ;

Royal Botanic Gardens Sydney

Mrs Macquaries Road, Sydney 2000, Australia*

Abstract

Swietenia alternifolia (Mill.) Steud., formerly referred to Meliaceae, is a species of Luehea

Willd. (Malvaceae, s./.): the appropriate new combination, L. alternifolia (Mill.) Mabb.

(syn. L. speciosa Willd.), is therefore proposed.

Introduction

In the disposing of names to be excluded from Meliaceae in an account of

that family being prepared for Kubitzki’s Families and Genera of Vascular

Plants, the binomial Swietenia alternifolia (Mill.) Steud., i.e. Cedrus

alternifolia Mill. (Cedrela alternifolia (Mill.) Steud.), has had to be

considered. That species described by Philip Miller (1691-1771) was based

on the field notes made in 1730-1 in Campeche, Mexico, by William

Houstoun (1695-1733) - the British surgeon who trained under Boerhaave

in Leiden - combined with observations on plants raised by growers in

England, including plants grown in the Chelsea Physick Garden from seeds

sent by Houstoun. In his description of the new species, Miller (Gard.

Dict. ed. 8, Cedrus n. 3, 1759) noted, “The third Sort [of Cedrus, 1.e.

Cedrela| was discovered by the late Dr Houstoun at Campeachy, from

whence he sent Seeds to England, which succeeded in several Gardens;

when the Doctor first observed these Trees, they were destitute of Leaves,

but were loaded with ripe Fruits; and on his second Visit to the Place he

found the Trees in full Verdure”.

The plant is recorded by George Don in his General History (1831;

see below) as if still being cultivated in England, but it seems never to have

flowered there, and, in the Flora neotropica monograph dealing with the

mahogany genus, Swietenia Jacq., Styles (in Pennington and Styles 1981:

385, 405) rejected it from the family Meliaceae, though he did not suggest

any other disposition for it.

Houstoun’s MS description quoted by Miller, “Arbor excelsa Coryli

folio ampliore”, is his ‘C[ampichianus ager]. Coryli folio arbor’ of f. 101 in

* Address for correspondence

254 Gard. Bull. Singapore 54 (2002)

Catalogus plantarum in America observatarum by Mr Wm HOUSTON

(MS Banks 67, The Natural History Museum, London). This entry is

accompanied by the note, ‘spec.’, indicating that a specimen was preserved.

Houstoun’s herbarium specimens were bequeathed to Miller (Dandy 1958 :

139-140) and, in 1774, Miller’s collection was bought by Sir Joseph Banks,

whose own herbarium came in 1827 to the British Museum, now the

herbarium (BM) of The Natural History Museum, London.

The MS description, combined with Houstoun’s other observations

and records set out above, show that he had found in the Campeche area a

large, deciduous tree with simple, alternate, large, hazel-like leaves and

Cedrela-type fruits - and that he had collected a specimen of it.

The description is very strongly reminiscent of the plant more recently

grown, albeit only occasionally, in some tropical botanic gardens (e.g.

Bogor, Indonesia [Danimihardja & Notohardjo 1978: 153] and Penang,

Malaysia [Cheang Kok Choy s.n, Nov. 1966 — L!, SING]) as Luehea speciosa

Willd. (Malvaceae, s./.), a tree native in the Neotropics. This is a species

known from Yucatan and, sure enough, an excellent flowering specimen,

collected by Houstoun, complete with his MS label, ‘Coryli folio arbor

Clampeachy] MSS’, is filed with other Mexican material of L. speciosa at

BM. The correct disposition of Swietenia alternifolia is therefore under

Luehea.

New Combination

Luehea alternifolia (Mill.) Mabb., comb. nova

Cedrus alternifolia Mill., Gard. Dict., ed. 8, Cedrus n. 3 (“alternifolius”,

1759) & ed. 9: Cedrus n. 3 (‘alternifolius’, 1768); Chaz., Dict. Jard. 3: 332

(‘“alternifolius”, 1785) - Cedrela alternifolia (Mill.) Steud., Nomencl. 1: 170

(1821); DC., Prodr. 1: 625 (1824); G. Don, Gen. Hist. 1: 687 (1831); M.

Roem., Fam. Nat. Syn. Monogr. 1: 136 (1846); C.DC., in DC., Monog.

Phan. 1: 747 (1878); Styles, Fl. Neotropica 28: 385 (1981).- Swietenia

alternifolia (Mill.) Steud., Nomencl., ed. 2: 653 (1841); Styles, op.cit. : 405

(1981).

Type: Mexico, Yucatan, Campeche, 1730-1, Houstoun s.n. (BM !, neo,

selected here. In his description, first printed in ed. 7 (1757) of his Gardeners

Dictionary, Miller made no mention of flowers, strongly suggesting that

this Houstoun specimen was not taken into account when the description

was originally drawn up — indeed, if it had been, Miller would no doubt not

have named it as he did !).

Swietenia alternifolia 255

Luehea speciosa Willd. in Neue Schr. Ges. Naturf. Fr. Berlin 3: 410, t. 5

(‘Liihea’, 1801); Burret in Notizbl. Bot. Gart. Mus. Berlin-Dahlem 9: 831

(1926), synon. nov.

Type: [Venezuela,] ‘Auf dem Gipfel des hohen Gebirges zwischen la

Gauyana [sic] und Caracas’, January, Bredemeyer s.n. in Herb. Willdenow

14376 (B-WILLD, fiche!).

Acknowledgements

I am grateful to Sandy Knapp and Charlie Jarvis (BM) for guidance in

matters concerning Houstoun’s collections and to Neil Chambers of the

Banks Archive Project, The Natural History Museum, London, for reading

over a draft of this paper.

References

Dandy, J.E. 1958. The Sloane Herbarium. British Museum (Natural History),

London, U.K.

Danimihardja, S. and D. Notohardjo. 1978. An Alphabetical List of Plant

Species Cultivated in the Hortus Botanicus Bogoriensis. Indonesian

Institute of Sciences, Indonesia.

Pennington, T.D. and B.T. Styles. 1981. Meliaceae. Flora neotropica 28.

Organisation for Flora Neotropica, New York, U.S.A.

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Gardens’ Bulletin Singapore 54 (2002) 257-261.

Nepenthes platychila (Nepenthaceae),

a New Species of Pitcher Plant from Sarawak, Borneo

CHAEN OC LEE

Malesiana Tropicals, Kuching, Malaysia

Abstract

Nepenthes platychila Chi.C. Lee, a new species from the Hose Mountains in central Sarawak,

Borneo, is described and illustrated.

Introduction

The Hose Mountains is a remote and relatively isolated mountain range

located in central Sarawak between the watersheds of Sungai Baleh and

Sungai Balui. Botanically, the region was poorly known in the last century

largely due to difficulty of access and the extremely rugged topography.

Recent expeditions, however, indicate that the region is a very important

centre of diversity for Nepenthes in Sarawak from where a total of eight

taxa have been so far recorded (Lee, 2002).

During an expedition to the Hose Mountains in December 2001, on

the lower slopes of Gunung Bukit Batu, plants of an unusual taxon of

Nepenthes were found that did not match any described in Clarke (1997).

The extraordinary form of the pitcher peristome immediately suggested

that these plants belong to a previously unrecognized species. Specimens

collected on this expedition were deposited in the Kuching Herbarium

(SAR) and comparison with material from previous collections and other

possibly related species, confirmed that it is new. The description below

follows the conventions of Danser (1928) as maintained by Cheek and

Jebb (2001).

Nepenthes platychila Chi.C. Lee, sp. nov.

Nepenthi fuscae Danser similis sed peristomio lato expanso, operculo lato

ovato, facie inferiore plana (sine appendice lateraliter applanata) differt —

TYPUS: Gunung Bukit Batu, Hose Mountains, Sarawak, 30 Nov 2001, C.

Lee S 87071 (SAR).

address for correspondance: Malesiana Tropicals Sdn. Bhd.,1° Floor, Lot 4909, Sect. 64

KTLD, Upland Shop House, Jalan Upland, 93300 Kuching, Sarawak, Malaysia

258 Gard. Bull. Singapore 54 (2002)

Figure 1

Terrestrial or epiphytic climber to 4 m tall. Indumentum present on all

parts, very dense on stems, tendrils, young pitchers and inflorescence,

consisting of red-brown hairs 0.5—1 mm long, branched at the base; upper

surface of leaf lamina fairly densely pubescent with short white stellate

hairs 0.1-0.3 mm long. Climbing stems terete, 5-8 mm diam., internodes

6.5-8.5 cm long, axillary buds spike-like. Leaves of the climbing stems

coriaceous, blade oblong-elliptic, 10.5-18 by 3.8-4.6 cm, dark greyish-green

occasionally purple-green, upper pitchers light-green with abundant red

streaks, peristome yellowish-green with numerous purple-red streaks of

varying width, apex usually obtuse, occasionally sub-peltate, base clasping

the stem for 1/2—2/3 its circumference and decurrent for more than half the

internode, gradually attenuate into the winged petiole; longitudinal veins 2

or 3 on each side of the midrib, pennate veins reticulate and inconspicuous.

Lower pitchers unknown. Upper pitchers narrowly infundibular in lower

half, broadly infundibular in upper half; 12.5—16.5 by 5.1-6.5 cm; ventral

ridges indistinct; mouth sub-orbicular to slightly ovate, horizontal in front

and elevated towards lid; peristome flattened and expanded, to 3.3 cm

wide, ribs indistinct; inner surface glandular throughout; spur inserted 1

mm from base of lid, 2 mm long; lid ovate or orbicular-ovate, rounded or

slightly cordate at base, to 4.8 x 4.4 cm, lower surface without appendages,

often slightly keeled in the basal part of the midrib, with scattered small

crater-like glands most abundant on either side of the midrib. Bracts

absent. Male inflorescence: peduncle 8 cm long, rachis 23 cm !ong; partial

peduncles 2-flowered, to 4 mm below the branch, with each branch to 12

mm long; sepals ovate-elliptic, 3-4 mm long; staminal column 4-5 mm

long. Female inflorescence not seen. Infructescence similar in structure to

the male inflorescence, bearing up to c. 100 fruits; peduncle to 12.5 cm

long, rachis to 16 cm long; mature fruits 3.54.5 cm long.

Distribution: Borneo: SARAWAK - Kapit Division, Hose Mountains,

Gunung Bukit Batu and Bukit Sindap.

Habitat: Growing epiphytically in moss forest or terrestrially on steep

sandstone slopes among Dicranopteris and Dipteris ferns, 900-1400 m

elevation.

Notes: This species is readily distinguished by the unique form of its

peristome, which is greatly expanded and almost completely smooth on its

upper surface. The size of the peristome is variable within the species, but,

in its most extreme form, it is so wide that the entire pitcher is hidden

Nepenthes platychila from Sarawak 259

2cm

Figure 1. Nepenthes platychila Chi.C. Lee

a. Habit with upper pitchers; b. male inflorescence; c. male flower; d. detail of peristome, internal

view; e. upper surface of lid; f. underside of lid; g. detail of glands of lower lid surface; h. detail

of hairs on undersurface of lid; i. detail of hairs; j. fruit. (a-i from S 87071, | from S 87074).

260 Gard. Bull. Singapore 54 (2002)

beneath it when viewed from above. The only other species in the genus

bearing a similar peristome structure is Nepenthes jacquelineae C.Clarke,

T.Davis & Tamin of West Sumatra, but these two species appear to be

otherwise unrelated. It is unclear what purpose the unusual structure of

the peristome serves, but it may provide a convenient landing platform for

potential insect prey or as suggested by Clarke (2001) for N. jacquelineae

assist in luring insects by making the pitcher mouth appear brighter than

the surroundings.

Nepenthes platychila is clearly a member of Danser’s Regiae group,

and within this group it is probably most closely related to N. fusca Danser

by the similar pitcher shape and other vegetative features. However, the

distinctive peristome and broad lid without appendages separate these two

taxa, and indeed serve to distinguish this species from all others in the

Regiae group. Other unusual characteristics of this species include the

long pedicels and extraordinarily large mature fruits.

Observations in the field, and the fact that no rosette plants or lower

pitchers have yet been collected, suggest that this species enters a climbing

stage relatively early in growth.

This species is sympatric with Nepenthes fusca, N. pilosa Danser, N.

reinwardtiana Miq., N. tentaculata Hook f., and N. veitchii Hook f. A

single plant at the type locality displaying characteristics intermediate

between N. platychila and N. fusca may be a natural hybrid between these

two species.

Although so far known from only a few localities, it is unlikely that

this species is restricted to Gunung Bukit Batu and Bukit Sindap, since

suitable habitat conditions occur extensively throughout the Hose

Mountains.

Other specimens examined: SARAWAK: Hose Mountain, Sungai Temiai,

Mujong, 1260 m, 6 Dec 1991, Lai, Jugah, Awg. Enjah S 64084 (KEP,

SAR); Bukit Sindap, Hose Mountains, 1100 m, 1 Dec 2001, C. Lee S 87074

(K, SAR).

Acknowledgements

The author thanks YB Datuk Amar Haji Abdul Aziz Haji Husaian, CEO/

Chairman of the Sarawak Biodiversity Centre; Mr. Cheong Ek Choon,

Director of Forestry Department Sarawak; Datuk Amar [Dr.] Leonard

Linggi Tun Jugah, Chairman of Malesiana Tropicals Sdn. Bhd.; L. C. J.

Julaihi (SAR) for his generous assistance: Charles Clarke for his critical

review of the manuscript; Joseph Pao for preparing the line drawing; and

Nepenthes platychila from Sarawak 261

Jeland ak Kisai and Mael ak Lete for their invaluable assistance in the

field.

References

Clarke, C. M. 1997. Nepenthes of Borneo. Natural History Publications

(Borneo), Kota Kinabalu, Malaysia. xi + 207 pp.

Danser, B. H. 1928. The Nepenthaceae of the Netherlands Indies, Bulletin

Jardin Botanique Buitenzorg. Ser. Ill. 9: 249-438.

Cheek, M. and M. Jebb. 2001, Nepenthaceae, Flora Malesiana. 15: 1-157.

Lee, C. 2002. Nepenthes species of the Hose Mountains in Sarawak, Borneo,

Proceedings of the 4" International Carnivorous Plant Conference,

Hiroshima University ,Tokyo. Pp. 25-30.

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Gardens’ Bulletin Singapore 54 (2002) 263-286.

Leaf Epidermal Micromorphology of Grewia L. and

Microcos L. (Tiliaceae) in Peninsular Malaysia and

Borneo

RC. Ky CHUNG

Forest Research Institute Malaysia

Kepong, 52109 Kuala Lumpur, Malaysia

Abstract

Leaf epidermal morphology of 5 species of Grewia L. and 32 species of Microcos L. (including

their type species) were examined. Grewia and Microcos both have glandular and non-

glandular trichomes. Trichome characters alone cannot be used for delimiting Grewia from

Microcos or for distinguishing species within each genus. Five epidermal characters were

useful for distinguishing the two genera in Peninsular Malaysia and Borneo. Grewia species

differ from Microcos species in having radiating cuticular striation of epidermal/subsidiary

cells, predominantly anomocytic stomata, stomata elliptic to broadly elliptic in outline with

mean length 18.6—22.9 um and average length-width (L/W) ratios of 1.2-1.4. The Microcos

species were characterised by the absence of radiating cuticular striation of epidermal/

subsidiary cells (except in M. tomentosa), predominantly paracytic and anisocytic stomata,

stomata broadly elliptic to oblate in outline with mean length 12—16.4 um and average L/W

ratios of 0.9-1.1.

Introduction

The genus Grewia consists of about 200 species of small trees and shrubs

or rarely scandent shrubs, distributed from tropical Africa northwards to

the Himalayas, China and Taiwan, south eastwards to India, Sri Lanka,

Myanmar, Thailand, Indo-China, Malesia, Tonga, Samoa and the northern

parts of Australia. In Malesian region about 30 species are known, of

which 4 occur in Peninsular Malaysia and Borneo. Microcos is a genus of

about 80 species of trees and shrubs, occurring in tropical Africa, India, Sri

Lanka, Myanmar, Indo-China, S China, Hainan, Thailand, and throughout

Malesia except for the Lesser Sunda Islands. In Malesia, some 42 species

are known, of which 31 occur in Peninsular Malaysia and Borneo (Boer

and Sosef, 1998; Phengklai, 1998; Chung, 2001). For species in Peninsular

Malaysia and Borneo, a total of 13 macromorphological characters have

been identified as diagnostic for distinguishing Microcos species from those

of Grewia (Chung, 2001).

The taxonomic value of leaf epidermal structures is well documented

(Wilkinson, 1979; Theobald et al., 1979). In the order Malvales, to which

264 Gard. Bull. Singapore 54 (2002)

the Tiliaceae belong, epidermal characters, particularly the type and

distribution of trichomes and stomata, are reported useful for delimiting

and/or distinguishing taxa at subfamily levels (Metcalfe and Chalk, 1950;

Inamdar and Chohan, 1969; Rao and Ramayya, 1977; 1983; Inamdar et al.,

1983; Salma, 1999).

Work on organography as well as taxonomic significance of leaf

epidermal characters in the order Malvales is meagre (Solereder, 1908;

Metcalfe and Chalk, 1950). In the Tiliaceae, studies on the taxonomic

significance of epidermal characters are very scanty and limited to very

few genera from India, 1.e. Berrya (Inamdar et al., 1983; 1 species),

Corchorus (Rao and Ramayya, 1981; 1984; 1987; Inamdar et al., 1983; Rao,

1990; Sharma, 1990; Singh and Dube, 1993; 8 species), Grewia (Tiwari,

1978; Rao and Ramayya, 1981; 1984; 1987; Inamdar et al., 1983; Sharma,

1990; 32 species), Microcos (Inamdar et al., 1983; Sharma, 1990; 1 species),

Triumfetta (Rao and Ramayya, 1981; 1984; 1987; Inamdar et al., 1983; Rao,

1990; Sharma, 1990; 5 species).

Likewise, the significance of trichomes in plant systematics has long

been recognised in the classification of angiosperms (Davis and Heywood,

1963; Edmonds, 1982). Previous studies on the trichome morphology and

distribution in the Tiliaceae were conducted by Metcalfe and Chalk (1950),

Kostermans (1972), Morawetz (1981), Rao and Ramayya (1987), Rao

(1990), and Sharma (1990).

The aim of the present study is to investigate the micromorphological

variability of the leaf epidermis and to assess its taxonomic significance in

the Peninsular Malaysian and Bornean Grewia and Microcos.

Materials and methods

Most leaf material investigated was obtained from herbarium specimens

available at the Forest Research Institute Malaysia (KEP), unless stated

otherwise (herbarium abbreviations as in Holmgren et al., 1990). The species

and specimens studied are listed in the Appendix.

Fully developed leaves were selected from dried herbarium specimens.

For each leaf, an area from the middle of the lamina, including midrib and

leaf margin, was taken. All material was rehydrated in water prior to

cuticular maceration in Jeffrey’s solution (a mixture of equal volumes of

10% chromic acid and concentrated nitric acid), stained in a solution of

Sudan IV in alcohol 70% and mounted in glycerin-jelly. For scanning

electron microscopy (SEM), samples of dried leaf specimens (adaxial and

abaxial surfaces) were directly mounted on stubs and coated with gold

and observed under a JOEL JSM-6400 scanning electron microscope.

Leaf micromorphology of Grewia and Microcos 265

Figure 1. Trichome types in Grewia and Microcos.

A, short unicellular stalk with multicellular elliptic glandular head; B, short unicellular stalk

with multicellular broadly elliptic glandular head; C, short unicellular stalk with multicellular

rounded glandular head; D, short unicellular stalk with multicellular oblong glandular head;

E, simple; F, 2-armed tufted; G, 3-armed tufted; H, 4-armed tufted; I, 4-armed stellate; J, 6-

armed stellate; K, 4-armed cushioned stellate; L, 4-armed cushioned stellate; M, 8-armed

cushioned stellate; and N, 7-armed cushioned stellate trichomes.

Gard. Bull. Singapore 54 (2002)

266

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Leaf micromorphology of Grewia and Microcos

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268 Gard. Bull. Singapore 54 (2002)

Micrographs were made of both trichomes and stomata.

The pattern of stomatal distribution was classified as random (i.e.,

low density of stomata distributed randomly without regular pattern) or

regular (1.e., high density of stomata in a regular pattern). Stomatal sizes

were measured directly using a calibrated linear eyepiece micrometer.

Density mm was counted from 25 unit areas on the abaxial surface and

stomatal index was determined using the formula adapted from Wilkinson

(1979: 114). Trichomes were observed in epidermal peels as well as

stereoscopic observations of herbarium specimens. The descriptive

terminology of epidermal features is based on Van Cotthem (1970): Radford

et al. (1974); Wilkinson (1979); Theobald et al. (1979) and Baranova (1987).

Trichome types are illustrated in Fig. 1 and leaf epidermal characters are

summarised in Table 1.

Results

GREWIA L.

(Figure 1; Plates 1 and 2; Table 1)

Stomatal Complex

Epidermal and subsidiary cells

In the species investigated, the anticlinal walls of the epidermal cells

on both the leaf adaxial and abaxial surfaces are straight. This particular

character, therefore, has no taxonomic value for distinguishing Grewia

species.

Cuticular striation occurs only on the epidermal cells adjacent to the

stomata. The striae radiating from the external edges of guard cells are

either long (G. occidentalis, G. huluperakensis and G. multiflora) or, less

frequently, short (G. /aevigata and G. polygama).

Stomata

The stomata are mostly surrounded by a limited number of

unspecialised cells that are indistinguishable in size and shape from the

epidermal cells. However, a few stomata are occasionally accompanied on

either side by one or more subsidiary cells parallel to the long axis of the

pore and guard cells. Hence, the majority of the stomata are anomocytic

or very rarely anisocytic and paracytic types.

The stomatal outline varies from elliptic, broadly elliptic to rarely

rounded (G. huluperakensis and G. laevigata) to exclusively broadly elliptic

(G. multiflora) or broadly elliptic to rarely rounded (G. polygama). The

stomata are elevated slightly above the surrounding epidermal cells in G.

polygama, but positioned at the same level in the other three species.

Leaf micromorphology of Grewia and Microcos 269

The stomata are 15-—27.2 um long and 12-—22.4 um wide with a mean

length of 18.6 um in G. polygama, 19.5-19.8 um in G. huluperakensis and

G. multiflora, and 22.9 um in G. laevigata. Mean width ranges from 14.4—

15.2 um in G. huluperakensis, G. multiflora and G. polygama to 17 um in

G. laevigata. The range of length-width ratios (L/W) is (1—)1.2-1.4(-1.7).

Stomata distribution

Leaves are either hypostomatous (with stomata restricted to the

abaxial surface) in G. huluperakensis, G. laevigata and G. multiflora or

amphistomatous (stomata on both sides) in G. polygama. In all four species,

the stomata are randomly distributed.

The stomatal density ranges from 80 to 180 mm”, with means ranging

from 95 to 157 mm”. In G. laevigata, the stomatal density, (80—)95(-120),

is slightly lower than that of G. huluperakensis, G. multiflora and G.

polygama, (136—)147-157(-180). Stomatal index ranges between 11.3 and

29, with a mean of 14.1 in G. laevigata and 25.9-27.3 in G. huluperakensis,

G. multiflora and G. polygama.

Trichomes

Non-glandular trichomes

On the adaxial leaf surface, non-glandular trichomes are sparse in G.

huluperakensis, and sparse to none in G. laevigata, G. multiflora and G.

polygama. On the abaxial surface, dense to moderately dense non-glandular

trichomes occur in G. polygama, and sparse to none in G. laevigata. In G.

huluperakensis and G. multiflora, the abaxial surface is moderately densely

covered with non-glandular trichomes.

In Grewia, four main types and nine subtypes of non-glandular

trichomes are recognised (Fig. 1). These are: (1) simple trichomes, which

are long and tapered at the tip; (ii) tufted trichomes, which are either 2-, 3-

or 4-armed; (iii) stellate trichomes, which are either 4-, 5- or 6-armed and

without a central cushion; and (iv) cushioned stellate trichomes, which are

4—8-armed and with a central cushion. The cushioned stellate trichomes

are formed by a multicellular hair base (Figure 1L).

These were observed on either the abaxial, adaxial, or both sides of

the leaf (Table 1). Simple trichomes are present in G. huluperakensis

(sparse), G. multiflora (dense) and G. polygama (sparse), but absent in G.

laevigata. The 2—4-tufted non-glandular trichomes are dense in G.

huluperakensis, but are absent in the other species. Four-armed stellate

trichomes are sparsely present in G. occidentalis (the type species) but

absent in the other species, while the 5—6-armed stellate trichomes are

present in G. /aevigata but absent in the others. The 4-8-armed cushioned

stellate non-glandular trichomes occur in G. laevigata and G. polygama,

270 Gard. Bull. Singapore 54 (2002)

but are absent in the other species .

Glandular trichomes

Glandular trichomes are present on both leaf surfaces, varying from

sparse to moderately dense. Each glandular trichome consists of a

multicellular head which is either sessile or with a short unicellular stalk.

The outline of the multicellular head can be elliptic as in G. huluperakensis

and G. multiflora, or broadly elliptic as in G. huluperakensis, G. laevigata

and G. multiflora, or rounded as in G. laevigata, or oblong as in G. polygama.

All Grewia species examined possess glandular trichomes with short

unicellular stalks, except G. laevigata which has sessile or short-stalked

glandular trichomes.

Remarks

Apart from minor differences, the stomatal characters and trichome

micromorphology of these species conform to those of G. occidentalis (the

type species of the genus Grewia).

The leaves of G. occidentalis are glabrous on both surfaces except

the midrib and secondary veins, which are sparsely covered with 4—6-armed

stellate trichomes.

Based on trichome micromorphology, the four species can be

segregated into three broad groups as follows:

Group 1. G. multiflora: Simple non-glandular trichomes only

Group 2. G. huluperakensis: Simple and tufted 2-4-armed non-glandular

trichomes

Group 3. G. laevigata and G. polygama: 4—-8-armed stellate and/or

cushioned stellate trichomes.

This study mostly confirms earlier investigations (Metcalfe and Chalk,

1950; Inamdar et al., 1983; Rao and Ramayya, 1987; Rao, 1990; Sharma,

1990), but with some differences. In G. polygama from India, Sharma

(1990) observed trichomes with short basal cell, peltate apical cell and 5-

10-armed tufted trichomes, but these types were not observed in G.

polygama from Peninsular Malaysia. This is probaby the same as 4-8-

armed stellate trichomes as defined by the author. Similary, Rao also

reported simple unicellular and 5-10-armed tufted trichomes in G. glabra

Blume (now G. multiflora). In the present study, only the simple trichomes

were observed in Bornean material. The simple unicellular filiform

trichomes commonly found by Inamdar et al. (1983) in the vegetative and

floral parts of G. asiatica L. were not observed in the Peninsular Malaysian

and Bornean Grewia species. Similarly, mucilaginous cells observed by

Rao and Ramayya (1984) in the adaxial surface of G. subinaequalis DC.

and G. tenax (Forsk.) Aschers. & Schwf. were not encountered in any

Leaf micromorphology of Grewia and Microcos 271

Plate 1. Stomatal complex of Grewia species

A: G. laevigata (Abdul Samat 265), LMs of abaxial leaf surface showing straight anticlinal

wall of epidermal cells. — B: G. occidentalis (type species, Relinhoonk 483), LMs of abaxial

leaf surface with stomata showing stomatal type, stomatal shape in outline, and orientation of

striae over epidermal cells. — C-F: LMs (C—E) and SEMs (F) of abaxial leaf surface showing

stomatal distribution. C: G. occidentalis (Relinhoonk 483); D: G. laevigata (Abdul Samat 265);

E: G. huluperakensis (Turner 94-94); F: G. multiflora (Motley 240). — G: G. huluperakensis

(Burkill & Mohd. Haniff SFN 12410), SEMs of abaxial leaf surface with stomata showing

orietation of striae over anomocytic epidermal cells. — Bar equals 10 um in B, F—G; 50 um in

A, C-E.

272 Gard. Bull. Singapore 54 (2002)

Peninsular Malaysian and Bornean material.

MICROCOS L.

(Figure 1; Plates 3 and 4; Table 1)

Stomatal Complex

Epidermal and subsidiary cells

In most species of Microcos investigated, the anticlinal walls of the

epidermal cells on the adaxial surface of the leaf are straight or curved. In

M. dulitensis, M. fibrocarpa, M. gracilis, M. malayana, M. pearsonii, M.

reticulata, and M. sumatrana, however, these cell walls are usually undulate.

On the abaxial surface, the anticlinal epidermal cell walls are straight to

curved as in M. antidesmifolia (var. antidesmifolia and var. hirsuta), M.

borneensis, M. cinnamomifolia, M. erythrocarpa, M. fibrocarpa, M.

globulifera, M. henrici var. acuta, M. kinabaluensis, M. lanceolata, M.

latistipulata var. latistipulata, M. laurifolia, M. membranifolia, M. ossea, M.

riparia, M. stylocarpoides, M. subepetala, M. tomentosa, and M. triflora

(var. triflora and var. longipetiolata), or undulate as in M. crassifolia, M.

dulitensis, M. gracilis, M. hirsuta, M. latifolia, M. malayana, M. opaca, M.

pachyphylla, M. pearsonii, M. phaneroneura, M. reticulata, M. subcordifolia,

and M. sumatrana. Cuticular striation of epidermal or subsidiary cells is

absent in most species, but short striae are frequently present in M.

tomentosa.

Stomata

Stomata of Microcos species found in Peninsular Malaysia and Borneo

are mainly paracytic and anisocytic or occasionally anomocytic. The outline

of the stomata varies from broadly elliptic, rounded to oblate in most

species of Microcos but broadly elliptic to rounded in M. stylocarpoides.

Stomatal size varies from 10 x 9 um to 18 x 18 um. Mean stomatal

length ranges from 12 um in M. pearsonii to 16.4 um in M. dulitensis, and

width from 12 um in M. subcordifolia to 16.1 um in M. borneensis. The

stomatal length-width ratios (L/W) range from 0.8 to 1.5 with mean length-

width ratios of 0.9 in M. kinabaluensis, M. malayana and M. tomentosa, 1.1

in M. antidesmifolia var. hirsuta, M. fibrocarpa, M. globulifera, M. gracilis,

M. hirsuta, M. phaneroneura, M. riparia, M. stylocarpoides, M. subepetala,

and M. triflora var. triflora, and equal to 1 in the other species.

In M. dulitensis, M. malayana, M. phaneroneura, and M. tomentosa,

the stomata are raised above the level of the epidermal cells, but in the

other species the stomata are at the same level with the epidermal cells.

Leaf micromorphology of Grewia and Microcos ZFS

Plate 2. Non-glandular trichomes of Grewia species.

A-B: SEMs of abaxial leaf surface showing non-glandular trichomes distribution, A: G.

laevigata (Asik SAN 113171), sparse; B: G. polygama (Soepadmo & Mahmud ES 1219), dense.

— C-F: SEMs of abaxial leaf surface showing main types and subtypes of non-glandular

trichomes. C: G. multiflora (Motley 240), simple trichomes; D: G. huluperakensis (Burkill &

Ma. Haniff SFN 12410), tufted trichomes of 2- or 3- armed form; E: G. laevigata (Mohd. Nur

SFN 32966), 5-armed stellate trichomes; F: G. laevigata (Aban SAN 49197), cushioned stellate

trichomes of 7-armed form. — Bar equals 10 um in E-F; 100 um in A-D.

274 Gard. Bull. Singapore 54 (2002)

Stomatal distribution

In M. antidesmifolia var. hirsuta, M. gracilis, M. latifolia, M. subepetala,

M. sumatrana, and M. tomentosa, stomata occur on both surfaces of the

leaf, but in the other species they are confined to the abaxial surface. The

stomatal distribution pattern in Microcos varies from random to regular.

The random distribution pattern is predominant in Microcos species, while

the regular type occurs in only a few species, such as M. globulifera, M.

kinabaluensis, M. pachyphylla, M. riparia, M. subepetala, and M. tomentosa.

2 The stomatal density varies from 84 mm _ in M. malayana to 316

mm in M. riparia, while the mean stomatal density ranges from 92 mm

in M. malayana to 291mm _ in M. globulifera and M. riparia. The stomatal

index ranges from 15.5 in M. borneensis to 30.6 in M. pearsonii, while the

mean stomatal index ranges from 17 in M. opaca to 28.4 in M. globulifera.

Trichomes

Non-glandular trichomes

Non-glandular trichomes are absent on both the adaxial and abaxial

leaf surfaces (including the midrib and veins) of M. antidesmifolia var.

antidesmifolia, M. crassifolia, M. kinabaluensis, M. latistipulata var.

latistipulata, M. pachyphylla, and M. triflora (var. triflora and var.

longipetiolata).

On the adaxial leaf surface, non-glandular trichomes are moderately

dense in M. latifolia, M. phaneroneura and M. reticulata, sparse in M.

fibrocarpa, M. globulifera, M. gracilis, M. subcordifolia, and sparse or absent —

in M. lanceolata and M. tomentosa. In M. antidesmifolia var. hirsuta, M.

dulitensis, M. erythrocarpa, M. henrici var. acuta, M. hirsuta, M. laurifolia,

M. malayana, M. membranifolia, M. opaca, M. pearsonii, and M.

stylocarpoides, the adaxial side of the lamina is glabrous, but the midrib

and secondary veins are sparsely to densely covered with non-glandular

trichomes.

On the abaxial leaf surface, non-glandular trichomes are dense in M.

fibrocarpa, M. phaneroneura, M. reticulata, and M. tomentosa, moderately

dense in M. antidesmifolia var. hirsuta, M. dulitensis, M. globulifera, M.

gracilis, M. hirsuta, M. malayana, M. subcordifolia, and M. sumatrana, and

sparse to none in M. henrici var. acuta, M. lanceolata, M. membranifolia,

M. riparia, and M. stylocarpoides. In M. borneensis, M. cinnamomifolia, M.

erythrocarpa, M. latifolia, M. laurifolia, M. ossea, M. pearsonii, and M.

subepetala, the abaxial surface of the lamina is glabrous, but non-glandular

trichomes are present on the midrib and/or secondary veins.

As in Grewia, four main types of non-glandular trichomes were

observed in Microcos, i.e. simple, tufted, stellate, and cushioned stellate

trichomes. The leaf surface of M. dulitensis, M. erythrocarpa, M. fibrocarpa,

Leaf micromorphology of Grewia and Microcos 275

Plate 3. Stomatal complex of Microcos species.

A-B: LMs of abaxial leaf surface showing straight or curved (A) and undulate (B) anticlinal

wall of epidermal cells. A: M. antidesmifolia var. antidesmifolia (Stone & Chin 13853); B: M.

malayana (Whitmore FRI 15328). — C-D: LMs of abaxial leaf surface showing stomatal

distribution. C: M. lanceolata (Kochummen KEP 79086), random; D: M. pachyphylla (Elmer

21704), regular. — E: M. dulitensis (Pickles S. 3689). LMs of abaxial leaf surface with stomata

showing stomatal type and stomatal shape in outline. — F—H: SEMs of abaxial leaf surface

with stomata showing variation in elevation of the guard cells relative to epidermal cells (F—

G: same level; H: raised). F: M. ossea (Tong S. 33866); G: M. antidesmifolia var. hirsuta

(Scortechini s.n.);H: M. malayana (Chelliah KEP 104372). — Bar equals 1 um in G—H; 5 um

in F; 10 um in E; 50 um in A-D.

276 Gard. Bull. Singapore 54 (2002)

M. globulifera, M. latifolia, M. malayana, and M. opaca is covered with

simple and 2—4-armed tufted trichomes, whereas that of M. antidesmifolia

var. hirsuta and M. reticulata is covered with simple, 2—4-armed tufted

trichomes and 4—8-armed cushioned stellate trichomes.

A combination of simple and 4—-8-armed cushioned stellate trichomes

is found in M. henrici var. acuta, M. hirsuta, M. membranifolia, and M.

subcordifolia, while a combination of simple, 4—-7-armed cushioned stellate

and 4-armed stellate trichomes is observed in M. phaneroneura. The midrib

and secondary veins of M. ossea on the abaxial surface bear simple and 5-

armed stellate trichomes, while those of M. pearsonii possess dense 5-

armed stellate trichomes. Microcos borneensis, M. cinnamomifolia, M.

gracilis, M. lanceolata, M. laurifolia, M. riparia, M. stylocarpoides, M.

subepetala, M. sumatrana, and M. tomentosa, on the other hand, have 4-8-

armed cushioned stellate trichomes on the lamina or on the midrib and

secondary veins.

Glandular trichomes

In most species (except M. kinabaluensis, which has completely

glabrous leaves), glandular trichomes of varying density are found on both

leaf surfaces. In M. borneensis and M. membranifolia, glandular trichomes

are in clusters of 2-4.

The glandular trichomes of most Microcos species investigated possess

short unicellular stalks, except in M. borneensis, M. ossea and M. triflora

var. triflora where the glandular trichomes are sessile. The outline of the

multicellular head varies from elliptic (M. crassifolia, M. henrici var. acuta,

M. lanceolata, M. pachyphylla, M. pearsonii, M. riparia, and M. tomentosa)

to broadly elliptic (M. dulitensis, M. erythrocarpa, M. gracilis, M. latistipulata

var. latistipulata, M. laurifolia, M. malayana, M. opaca, M. ossea, M.

phaneroneura, M. stylocarpoides, and M. subcordifolia), rounded (M.

globulifera, M. hirsuta, M. membranifolia, and M. reticulata), or oblong

(M. latifolia and M. triflora var. triflora) or varying from broadly elliptic to

rounded in M. antidesmifolia var. antidesmifolia and var. hirsuta, M.

cinnamomifolia, M. fibrocarpa, M. subepetala, and M. triflora var.

longipetiolata, or rounded to oblong in M. borneensis and M. sumatrana.

Remarks

The characters of the stomatal complex in the species examined match

those of M. paniculata (the type species of the genus Microcos), except in

the absence of cuticular striation on the epidermal cells in most of the

Peninsular Malaysian and Bornean species. |

In trichome micromorphology, they also match well with M.

paniculata. The leaves of M. paniculata are glabrous on both surfaces,

Leaf micromorphology of Grewia and Microcos 277

Plate 4. Non-glandular trichomes of Microcos species.

A: M. fibrocarpa (Lesmy FRI 33964), simple trichomes and 2-armed tufted trichomes. — B:

M. tomentosa (Spare SFN 36245), SEMs of abaxial leaf surface showing dense non-glandular

trichomes distribution. — C: M. henrici var. acuta (Ilias S. 36520), 8-armed cushioned stellate

trichomes. — D: M. paniculata (type species, Cramer 3405), 8-armed cushioned stellate

trichomes. — E: M. paniculata (Clemens 4122), 8-armed stellate trichomes. — Bar equals 100

um in A-E.

278 Gard. Bull. Singapore 54 (2002)

except the midrib and secondary veins in the abaxial surface which are

sparsely covered with 8-armed stellate and cushioned stellate trichomes.

Based on trichome micromorphology, the Peninsular Malaysian and

Bornean species of Microcos can be separated into five broad groups:

Group 1. M. kinabaluensis: Glandular and non-glandular trichomes

absent.

Group 2. M. antidesmifolia var. antidesmifolia, M. crassifolia, M.

latistipulata var. latistipulata, M. pachyphylla, M. triflora (var.

triflora and var. longipetiolata): Glandular trichomes present;

non-glandular trichomes absent.

Group 3. M. antidesmifolia var. hirsuta, M. dulitensis, M. erythrocarpa,

M. fibrocarpa, M. globulifera, M. henrici var. acuta, M. hirsuta,

M. latifolia, M. malayana, M. membranifolia, M. opaca, M.

ossea, M. phaneroneura, M. reticulata, and M. subcordifolia:

Glandular and non-glandular trichomes of simple, tufted,

stellate and cushioned stellate hairs present.

Group 4. M. pearsonii: Glandular and non-glandular trichomes of stellate

hairs present.

Group 5. M. borneensis, M. cinnamomifolia, M. gracilis, M. lanceolata,

M. laurifolia, M. riparia, M. stylocarpoides, M. subepetala, M.

sumatrana, and M. tomentosa: Glandular and non-glandular of

cushioned stellate trichomes present.

Results of previous studies on the trichomes of Microcos (Inamdar et

al., 1983; Sharma, 1990) differ slightly from those of the present study.

Sharma (1990) reported the occurrence of 5—10-armed tufted stellate

trichomes and three-celled unequally 2-armed trichomes with short stalks

in G. microcos L. (now M. paniculata). These two types of trichomes,

however, were not observed in any species investigated in the present

study.

Discussion

Comparative stomatal complex characters of Grewia and Microcos

The differences between the leaf epidermal characters of Grewia and

Microcos species occurring in Peninsular Malaysia and Borneo are

summarised in Table 2. In Grewia, the anomocytic stomata are the

predominant type, whereas in Microcos the paracytic and anisocytic stomata

are the prevalent type. Radiating cuticular striation is present on the

epidermal cells immediately adjacent to the guard cells in Grewia species,

but absent on the subsidiary cells or epidermal cells adjacent to the guard

cells in Microcos species, except in M. tomentosa. The stomatal outline in

Leaf micromorphology of Grewia and Microcos 279

Grewia species is mainly elliptic to broadly elliptic with average L/W ratios

of 1.2—-1.4, whereas in Microcos species it is mostly broadly elliptic, rounded

to oblate with average L/W ratios of 0.9-1.1.

The leaves of Grewia species are either amphistomatic, e.g. G.

occidentalis, G. polygama, and G. subinaequalis DC. (Rao and Ramayya,

1981), or hypostomatic, e.g. G. huluperakensis, G. laevigata Vahl, G.

multiflora, and G. tenax (Forsk.) Aschers. & Schwf. (Rao and Ramayya,

1981). Likewise, the leaves of Microcos species may be amphistomatic (M.

antidesmifolia var. hirsuta, M. gracilis, M. latifolia, M. subepetala, M.

sumatrana, and M. tomentosa), or hypostomatic (M. paniculata and other

species from Peninsular Malaysia and Borneo). In the Tiliaceae,

amphistomatic leaves have been reported in a few species of Corchorus L.

and Triumfetta L. (Rao and Ramayya, 1981; Singh and Dube, 1993). For

related families in the Malvales, Rao and Ramayya (1981) reported

exclusively amphistomatic leaves for Malvaceae, exclusively hypostomatic

leaves for Elaeocarpaceae, and both amphistomatic and hypostomatic leaves

for Bombacaceae and Sterculiaceae.

Table 2. Differences between the epidermal characters of Grewia and Microcos

in Peninsular Malaysia and Borneo.

Genus Grewia Microcos

Characters

Epidermal/Subsidiary cells

Radiating cuticular striation present absent (except in M.

tomentosa)

Stomata

Type predominantly anomocytic mainly paracytic and

anisocytic

(occasionally paracytic (occasionally

and anisocytic) anomocytic)

Outline elliptic, broadly elliptic broadly elliptic, rounded,

(occasionally rounded) oblate

Average length (tm) 18.6—22.9 12-16.4

Average L/W ratio 1.2-1.4 0.9-1.1

Metcalfe and Chalk (1950) stated that in the Tiliaceae, the stomata

were exclusively anomocytic (ranunculaceous), but Kundu and Sen (1958),

Sen and Paul (1961), and Singh and Dube (1993) demonstrated the presence

of anisocytic, paracytic and tetracytic stomata in Indian Corchorus species.

Likewise, the present study also discovered that the stomata of Grewia

280 Gard. Bull. Singapore 54 (2002)

species in Peninsular Malaysia and Borneo were predominantly anomocytic

and occasionally paracytic, and those of Microcos species were mainly

paracytic and occasionally anomocytic. For other families in Malvales,

Inamdar and Chohan (1969) reported the occurrence of anomocytic,

anisocytic and paracytic stomata in the Bombacaceae (Adansonia digitata

L. and Bombax ceiba Burm.f.) and the Malvaceae (Althaea rosea Cav.:

Gossypium herbaceum L.; Hisbiscus rosa-sinensis L., H. schizopetalus

Hook.f, H. syriacus L.; Malvaviscus arboreus Cav.; Sida acuta Burm.f, S.

alba Cav.; and Thespesia populnea Sol. ex Corr.Serr.). Hussin and Sani

(1998), on the other hand, reported that the stomata of 12 species of

Sterculia L. (Sterculiaceae) were exclusively anomocytic.

The occurrence of different types of stomatal apparatus in the leaves

of a single taxon or in different species of a genus or different genera of a

family has been noted by many authors (Metcalfe and Chalk, 1950; Kundu

and Sen, 1958; Sen and Paul, 1961; Inamdar and Chohan, 1969; Rao and

Ramayya, 1983; Singh and Dube, 1993), where usually a few stomata of

one type occur along with the predominant stomata of another type. Less

often, two or more stomatal types occur together as “codominants” on the

same leaf and/or species, as for example in the Convolvulaceae (Pant and

Banerji, 1965), and Bombacaceae, Sterculiaceae and Elaeocarpaceae (Rao

and Ramayya, 1983).

Taxonomic value of trichome characters in Grewia and Microcos

The present study on the leaf trichomes of Peninsular Malaysian and

Bornean Grewia and Microcos species reaffirms earlier observations on

Tiliaceae species (Solereder, 1908; Metcalfe and Chalk, 1950; Tiwari, 1978;

Rao and Ramayya, 1987; Sharma, 1990) as well as on related families of

the order Malvales (Ramayya and Rao, 1976; Rao and Ramayya, 1977,

1983; Inamdar et al., 1983; Salma, 1999), namely, the existence of several

different trichomes types.

Based on the type, distribution and density of non-glandular

trichomes, species of Grewia can be divided into three broad groups (i.e.,

non-glandular trichomes exclusively of simple hairs; non-glandular trichomes

both simple and 2—4-armed tufted hairs; and 4-8-armed stellate and/or

cushioned stellate trichomes). Likewise, five broad groups of species in

Microcos can be recognised (i.e., glandular and non-glandular trichomes

absent; glandular trichomes present but non-glandular trichomes absent;

glandular and non-glandular trichomes of simple, tufted, stellate and

cushioned stellate hairs present; glandular and non-glandular trichomes of

stellate hairs present; and glandular and non-glandular cushioned stellate

trichomes present).

Leaf micromorphology of Grewia and Microcos 281

The infraspecific variability may be due to genetic variability or

ecological factors (Baas, 1975). In M. triflora, the two varieties differ in

the the body/stalk of glandular trichomes (sessile in var. triflora; short

unicellular in var. longipetiolata) and in M. antidesmifolia, the two varieties

differ in the occurrence of non-glandular trichomes (absent in var.

antidesmifolia; simple, tufted and cushioned stellate hairs in var. hirsuta).

However, this infraspecific variation could not be correlated with ecological

data, as information from the herbarium labels was insufficient.

Conclusions

On the basis of leaf epidermal characters, the genus Microcos in Peninsular

Malaysia and Borneo can be distinguished from Grewia by a combination

of the following characters: stomatal type, outline, mean length and L/W

ratio, and the occurrence of cuticular striation on the epidermal cells

immediately surrounding the guard cells. To some extent, this study supports

Burret’s view (1926) for segregating Microcos from Grewia. However,

trichome micromorphological characters alone cannot be used for

distinguishing genera, and species within each genus. According to the

ordinal classification defined by APG (1998), stomatal types and trichomes

are just minor variations on a pattern common to most Malvaceae, which

now includes Tiliaceae.

Acknowledgements

I thank the Director-General of the Forest Research Institute Malaysia

(FRIM) for support and encouragement; the Curators of the herbaria BM,

BO, K, KEP, KLU, L, SAN, SAR, SING, and UKMB for the loan of

specimens; A. L. Lim, University of Malaya, E. Soepadmo (FRIM), and

anonymous reviewers for their advice and comments on this paper; and

Patricia Loh for technical assistance. This study was financially supported

by the Tree Flora of Sabah and Sarawak Project (IRPA grant 01—04—01-

0024), and University of Malaya postgraduate research grants 489/97, 225/

98, and 209/99.

282 Gard. Bull. Singapore 54 (2002)

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Appendix: Source of Materials Studied

The species and specimens studied are given below in alphabetical order;

(1) = maceration only; (2) = trichomes only

Grewia occidentalis L. (type species): Moss 3959 (BM); Relinhoonk 483

(L); Smook 6553 (BM) — G. huluperakensis I.M.Turner: Burkill & Md.

Haniff SFN 12410 (SING); Turner 94-94 (SING) — G. laevigata Vahl:

Aban SAN 49197 (SAN); Abdul Samat 265 (KLU); Amin & Joseph

SAN 60068 (KEP); Asik SAN 11317 (KEP, 2); Mohd. Nur SFN 32966

(KEP) — G. multiflora Juss.: Blume s.n. (L sheet no. 908.253.1458; L);

Motley 240 (K) — G. polygama Roxb.: Chin CSC 1832 (KEP, 2); Chung

RC 10 (KEP, 1); Kamarudin & Mustapa FRI 42255 (KEP); Kiah SFN

35286 (KEP); Soepadmo & Mahmud ES 1219 (KEP).

Microcos paniculata L. (type species): Bhargave 1787 (L); Clemens 4122

(BM); Cramer 3405 (L) — M. antidesmifolia (King) Burret var.

antidesmifolia: Everett FRI 14442 (KEP); King’s Collector 4085 (L); Stone

& Chin 13853 (KLU); var. hirsuta (King) Burret: Creagh s.n. (K); Elmer

20911 (SING); Enggoh FMS 48988 (KEP); Scortechini s.n. (BO) — M.

borneensis Burret: Ashton S. 17956 (KEP); Smythies et al. S. 5877 (KEP)

— M. cinnamomifolia Burret: s.c. FMS 34460 (KEP); Haviland 2256

(SAR); Nicholson & Ping SAN 17701 (KEP) — M. crassifolia Burret:

Amin & Lideh SAN 70341 (KEP); Amin et al. SAN 67317 (KEP, 1);

Ramos 1704 (K) — M. dulitensis Airy Shaw: Pickles S. 3689 (SAR);

Richards 1767 (SING) — M. erythrocarpa (Ridl.) Airy Shaw:

Kochummen FRI 16417 (KEP); Kochummen FRI 29363 (KEP) — M.

fibrocarpa (Mast.) Burret: Kiah SFN 35064 (KEP); Lesmy FRI 33964

(KEP) — M. globulifera (Mast.) Burret: Chan FRI 23806 (KEP); Ogata

KEP 105036 (KEP) — M. gracilis Stapf ex Ridl.: Chai SAN 26066 (SAN);

Yi et al. S. 37752 (KEP) — M. henrici (Baker f) Burret var. acuta

R.C.K.Chung, (ined).: Ilias S. 36520 (KEP); Ilias S. 45121 (KEP); Lai et

al. S. 75359 (KEP) — M. hirsuta (Korth.) Burret: Haviland 1471 (SING);

Korthals s.n. (L sheet no. 944.56.122; L) — M. kinabaluensis

R.C.K.Chung, (ined.): Chew & Corner RSNB 4216 (SAN); Chew &

Corner RSNB 4994 (SAN) — M. lanceolata (Miq.) Burret: Chelliah

KEP 104621 (KEP); Kochummen KEP 79086 (KEP) — M. latifolia

Burret: Whitmore FRI 8931; Wong WKM 1545 (SING) — M. latistipulata

(Ridl.) Burret var. latistipulata: Burkill SFN 7826 (SING); Chai SAN

286 Gard. Bull. Singapore 54 (2002)

29741 (KEP) — M. laurifolia (Hook. ex Mast.) Burret: Kochummen FRI

16338 (KEP); Razali 2809 (UKMB) — M. malayana R.C.K.Chung,

(ined.): Loh FRI 19249 (KEP); Chelliah KEP 104372 (KEP): Whitmore

FRI 15328 (KEP) — M. membranifolia R.C.K.Chung, (ined.): Kadir &

Jiran SAN A 3367 (KEP); Singh SAN 30672 (KEP) — M. opaca (Korth.)

Burret: Jlias S. 39135 (KEP); Yi S. 41139 (KEP) — M. ossea Burret:

Clemens 21071 (SAR); Dayang Awa & Ilias S. 45689 (KEP): Tong S.

33866 (KEP) — M. pachyphylla Merr.: Elmer 21704 (SING); Salleh et al.

BRUN 15315 (KEP) — M. pearsonii (Merr.) Burret: Dewol & Donggop

SAN 129474 (KEP); Meijer & Pereira SAN 47225 (SAN) — M.

phaneroneura Burret: Chai S. 36100 (KEP): Hallier 2868 (BO); Ong

OHC 716 (KEP, 1) — M. reticulata Ridl.: Ashton S. 17800 (KEP); Creagh

s.n. (K, 2): Saikeh SAN 69284 (SAN, 2): Wood SAN A 4788 (KEP) —

M. riparia (Boerl. & Koord.) Burret: Hallier 1314 (BO): Yeob FMS 3190

(KEP) — M. stylocarpoides Burret: Hallier 756 (BO); Haviland 1685

(SAR) — M. subcordifolia R.C.K.Chung, (ined.): Kirkup & Thomas

DK 727 (KEP); Niga NN 118 (KEP) — M. subepetala Stapf ex Ridl.:

Haviland 1885 (K) — M. sumatrana (Baker f) Burret: Forbes 2684 (L)

— M. tomentosa Sm.: Carrick 565 (KEP, 1); Chung RC 9 (KEP); Everett

FRI 13785 (KEP): Spare SFN 36245 (KEP): Tachun FMS 23227 (KEP,

1) — M. triflora (Blanco) R.C.K.Chung var. triflora, (ined.): Forestry

Student PNH 35319 (KEP); Lazo PNH 33406 (KEP): var. longipetiolata

(Merr.) R.C.K.Chung, (ined.): Aban & Soinin SAN 66898 (SAN):

Haviland 2332 (SAR, 1); Wong WKM 1244 (KEP).

Gardens’ Bulletin Singapore 54 (2002) 287-288.

Updates on Sections of Bulbophyllum (Orchidaceae)

Revised in Vermeulen (1993)

J.J. VERMEULEN

Singapore Botanic Gardens

Singapore 259569

Abstract

Some errors and omissions in the revision of Bulbophyllum, sections Macrouris and Uncifera

(Vermeulen, 1993), are addressed.

Bulbophyllum sect. Macrouris

Bulbophyllum subpatulum J.J. Verm., nom. nov.

Bulbophyllum subpatulum J.J. Verm. — Bulbophyllum muscicola Schltr., Repert.

Spec. Nov. Regni Veg.. Beih. 1 (May 1 . 1913) 870: J.J. Verm.. Orch. Mon.

7 (1993) 106. —TYPE: Papua New Guinea, Carr 10252 (neo SING, iso

AMES, CANB, K, NY).

[Not Bulbophyllum muscicolum Rchb.f., Flora 55 (1872) 275 (= close to B.

retusiusculum Rchb.f., from continental South and East Asia.)]

[Not Bulbophyllum muscicola Schitr., Ann. Mus. Col. Marseille 3 (1913, exact

date not traceable) 36 (= sect. Trichopus, from Madagascar]

Notes: Bulbophyllum muscicolum Rchb.f., a taxon overlooked by

Vermeulen (1993), renders his attempts to properly date the original

publications of both B. muscicola Schltr., both from 1913, superfluous. A

new name for the New Guinea B. muscicola Schltr. is warranted, because

B. equivestigium Gilli has to be deleted as a junior synonym, see under B.

cylindrobulbum, below.

Bulbophyllum trifilum J.J. Sm. subsp. filisepalum (J.J. Sm.) J.J. Verm..,

comb. nov.

Bulbophyllum trifilum J.J. Sm. ssp. filisepalum (J.J. Sm.) J.J. Verm., Orch. Mon. 7

(1993) 117, nom. inval. — Bulbophyllum filisepalum J.J. Sm., Meded. Herb.

Leiden 23 (1915) 15. —- TYPE: Indonesia, Irian Jaya, Janowsky 412 (holo L,

iso BO).

Notes: Paul Ormerod kindly informed me that the combination made in

Vermeulen (1993) is invalid according to the International Code of

Botanical Nomenclature (2000) Art. 33.5, because no reference is made to

the original description of B. filisepalum. The error is corrected here.

288 Gard. Bull. Singapore 54 (2002)

Bulbophyllum sect. Uncifera

Bulbophyllum cylindrobulbum Schltr.

Bulbophyllum cylindrobulbum Schltr. in Schum. & Laut., Nachtr. (1905) 200; J.J. Verm..,

Orch. Mon. 7 (1993) 178. —TYPE: Papua New Guinea, Reeve 1015 (neo K, iso

L).

Bulbophyllum equivestigium Gilli, Ann. Nat. Mus. Wien 84 (1983) 16; J.J. Verm..,

Orch. Mon. 7 (1993) 107 (in synonymy). - TYPE: New Guinea, Gilli 566 (holo

Notes: Based on the description of Bulbophyllum equivestigium, Vermeulen

(1993) incorrectly synonymised it with B. muscicola Schltr. Now that the

type is available at W, it is clear that B. equivestigium is identical with B.

cylindrobulbum.

Reference

Vermeulen, J.J., 1993. A taxonomic revision of the genus Bulbophyllum,

sections Adelopetalum, Lepanthanthe, Macrouris, Pelma, Peltopus and

Uncifera. Orchid Monographs 7: 1-324.

Gardens’ Bulletin Singapore 54 (2002) 289-292.

Book Review: Soepadmo, E., L.G. Saw and R.C.K. Chung, (eds.), 2002.

Tree Flora of Sabah and Sarawak. Volume 4. Sabah Forestry Department.

Forest Research Institute Malaysia, Sarawak Forestry Department. xii +

388 pp. ISBN 983-2181-27-5 (Vol. 4). Price: RM 100 or US$ 100.

The ambitiousness of this project should not be underestimated. Northern

Borneo contains, for its area, the largest lowland tree flora of anywhere in

the Old World, and possibly anywhere. When the work is completed, the

75,000 square miles of Sabah and Sarawak may have been found to contain

in excess of 5,000 tree species. This is the first flora for a major region of

Borneo. Merrill’s Bibliographic Enumeration of Bornean Plants (Journal

Straits Branch Royal Asiatic Society, Special Number, 1921), extended by

Masamune (Enumeratio Phanerogamarum Bornearum, 1942) contains but

a rudimentary and outdated base. The regional Flora Malesiana provides

an invaluable foundation, but hardly more than one third of the families

have yet been treated by that great enterprise, while a close regional scrutiny

is permitting more discriminating taxon recognition. Tim Whitmore, K.M.

Kochummen and Francis Ng’s groundbreaking Tree Flora of Malaya

(Longmans, 1972-89), which treated just under 3000 species over 24 years,

was based on a vastly greater previous botanical corpus including King’s

Materials for a Flora of the Malay Peninsula (1896) and Ridley’s Flora of

the Malay Peninsula (Reeves, 1922-25).

At a time when biological research in Malaysia is somewhat in the

doldrums, Engkik Soepadmo and his colleagues are succeeding, overall, in

a genuinely indigenous enterprise while determinedly holding to

international standards. They have used the Flora project to train a new

cadre of taxonomists, who have contributed many of the families. The

editors are also committed to meet the needs of both professional

taxonomists and the wider professional and public user, by minimizing

technical jargon, by including identification keys primarily or entirely based

on readily observable characters, and by providing summary information

on ecology and uses.

Overall, while recognizing that shortcomings in a project with such

ambitious goals are inevitable, the editors have succeeded to a remarkable

extent. To my knowledge, no comparable indigenous project for a field-

based and user-friendly critical flora has been undertaken elsewhere in the

tropics. Particularly commendable are the keys, which, to varying extent

admittedly according in part to the field knowledge of the authors, put

major emphasis on vegetative over reproductive characters. This is vital

for a region in which flowering among canopy trees fluctuates vastly between

years, and in which reproduction is often scant for at least half the year.

290 Gard. Bull. Singapore 54 (2002)

The insistence on citation of types, and on substantiating species concepts

and geographical range by citation of selected specimens, provides an

invaluable test of ngour. High quality whole page botanical illustrations of

selected species from each genus are provided. There are indices to both

vernacular and scientific names.

Volume 4 is dedicated to our late friend and colleague Tim Whitmore:

the gracious dedication identifies Tim as both creative spirit and driving

force behind the Tree Flora of Malaya, which provided the model for the

current flora. This volume includes treatments of 6 families, 24 genera and

321 species; of the latter, 45 (14%) are new to science, thereby indicating

the inadequate extent of previous knowledge in spite of the achievements

of Flora Malesiana. Overall, 47% of the species treated are endemic to

Borneo and just under half of these are endemic to Sabah and Sarawak,

offering dramatic evidence of the validity of northern Borneo as a global

biodiversity hotspot. Four of the families, including the three largest, are

authored by Malaysian specialists.

The true test of the quality of a flora must be by its use in the field,

as well as the herbarium. This reviewer knows the families treated as a

naturalist rather than a specialist, and it would be presumptuous to offer a

detailed critique before my next visit to the field.

Ebenaceae, by Francis Ng, includes the 75 ebonies of Sabah and

Sarawak. An informative introduction provides, after generic definition,

extensive details on geography, ecology, biology and uses, and is clearly

the work of someone who knows his material as living organisms. Curiously,

bark and architecture descriptions are hidden within a section on taxonomy

rather than under the generic definition. The key relies substantially on

fruit, as well as vegetative characters, but not at all on flowers. This implies

that vegetative characters alone are insufficient. It would have been useful

to have included discussion of the best means to distinguish the most

vegetatively similar species in footnotes under their descriptions, if

impractical in the key. Previously recognized subspecies have often been

suppressed. For instance it may be, as the author states, that continuous

variation between Diospyros sumatrana subspecies decipiens and sumatrana

exists in the herbarium, but my experience is that they are sufficiently

distinct both in habitat and morphology in the field to merit sustainment

as subspecies. More discussion of such variation, which seems to occur in

several Diospyros, would have been useful even if a conservative decision

is finally made. Overall, though, I found this to be a masterly treatment,

and an excellent spur for us field biologists to further advance knowledge

of the family.

Tree Flora Sabah and Sarawak 4 291

Sapotaceae, with 11 genera and 121 species in Sabah and Sarawak, is

treated by a team of eight botanists: seven from the Sarawak Herbarium,

Kuching, and Joan Pereira from Sandakan Herbarium, under the overall

editorship of Paul Chai and P.C. Yui at Kuching with varying but clearly

substantial input from senior editor Engkik Soepadmo. Sapotaceae is a

notoriously difficult family sharing, with Lauraceae, complex and difficult

generic definition, which must rely on reproductive characters. The

treatment suffers from the substantial disadvantage, therefore, of having a

generic Key that starts with and continues to use reproductive characters at

major divisions, preventing its use in field ecological and other surveys. In

my experience, at least the majority of entities are recognizable in the

field, albeit not through resort to generic characters. | remain convinced

that an additional key to all 121 species, based on field and if necessary

ecological characters, would have been feasible. Of the larger genera

Madhuca, 47 species treated, is by Yui and Chai; Palaquium, 41 species, by

Abang Mohtar with assistance from Soepadmo, Payena, 12 species, by

Pereira, and Pouteria, 7 species, by Stephen Teo. In all, seemingly workable

keys are provided, based mostly or entirely on vegetative characters.

Infraspecific categories, we are informed, are eliminated, being either raised

to species rank or rejected; but nowhere in this family treatment are

taxonomic explanatory footnotes provided beneath the species descriptions;

these could have also helped overcome difficulties presented by the

problematic generic key. It does seem curious that species new to science

are described only in the quite small genus Payena, with a remarkable 5,

and Madhuca, with 7. This might reflect differing species concepts among

the authors, alternatively differing knowledge and attention to detail; the

texts overall suggest the latter.

Oleaceae, with 6 genera and 47 species by Ruth Kiew, is a masterly

account of a family that is the author’s speciality. Again, it appears not to

be possible to reliably identify the three tree genera Chionanthus, Ligustrum

and Olea on vegetative characters. Some explanatory comments would

have again been helpful here: Do not the inflorescences, or at least their

abscission scars, remain on the twigs? The species descriptions are

particularly clear in this family and brief footnotes are often provided,

though rather little is offered on ecology. This author alone described new

species and combinations with extensive taxonomic notes in a useful

appendix in the volume

The treatments of the remaining, small, families are all competent.

Aquifoliaceae, by Susyn Andrews, has a valuable introductory field key.

There is no habit description of //ex, nor allusion to the distinctive upper

surface of the dry leaf. As might be expected from an author from outside

the region, the ecological information is variable. Lecythidaceae, by Michelle

292 Gard. Bull. Singapore 54 (2002)

Pinard, is serviceable, with keys primarily relying on field characters. It

had the advantage of Hans Payens’ prior treatment in Flora Malesiana. I

find Barringtonia species often difficult to identify; more footnotes would

have been welcome. Proteaceae, by Richard Chung, aided by Hans

Sleumer’s treatment in Flora Malesiana, provides a competent and clearly

useable account of this rather easy family, enriched by a surprising number

of novelties.

Peter S Ashton,

Arnold Arboretum,

Harvard University, USA.

and

Royal Botanic Gardens,

Kew, UK.

Gardens’ Bulletin Singapore 54 (2002) 293-294.

Book Review: Evans, T.D., K. Sengdaia, O.V. Viengkham and B.

Thammavong. 2001. A Field Guide to the Rattans of Lao PDR. Royal

Botanic Gardens, Kew, U.K. 96 pp. ISBN 1-84246-009-0. Price: 15 GBP.

Rattans are one of the most important forest products in generating income

for local communities, but in spite of their long history of usage, many

countries still do not have books that enable rattans to be reliably identified.

With the rapid rate of forest exploitation, it is critical to know which

species are the most valuable, how widely distributed they are, and whether

they have potential for cultivation. For this to be possible, the first step is

to know the scientific name of each rattan and this is what this field guide

aims to do.

This little book has taken a new direction in rattan research in

producing a user-friendly guide that is attractive and easy to use by a wide

range of readers from the botanists to the local forester, agriculturalist or

conservationist. Solidly based on both fieldwork and scientific herbarium

research, it combines a great deal of information into an attractive format.

It begins with a general section on basic information on rattans, which is

followed by field keys and the heart of the book, information on the 31

main species and 20 similar allied species found in Lao and the surrounding

region. The book finishes with an illustrated glossary of terms, which makes

the often alien structures and terminology peculiar to rattans easily

understood, advice on how to collect these spiny plants, the management

and conservation of rattans, and a section on planting and cultivating rattans.

Particularly useful are the novel field keys set out like a spreadsheet

so that it is possible to compare several characters simultaneously rather

than being confined to the usual dichotomous Keys that many non-botanists

find uncomfortable to use. In fact throughout the book, careful thought

has been taken to explain how taxonomy works and to give helpful advice

on how to go about identifying plants.

The 31 Lao species are each given a double spread that includes a

colour map to show its distribution within Lao and the surrounding region,

as well as the distribution of similar species, and a colour photograph of

the living stem and leaf sheath. Information includes the scientific and

vernacular names, key diagnostic characters and other characters to look

for when identifying them, its habitat, phenology, uses, cultivation and a

description illustrated by line drawings. A box entitled ‘Compare’ includes

similar or allied species and their diagnostic characters, which provides a

quick way of checking the differences between similar species. The clear

layout makes it easy to see the salient characters of each rattan.

It would have been more helpful to have included the glossy in the

294 Gard. Bull. Singapore 54 (2002)

introductory section so that a quick glance through would familiarise the

reader with the various structures before attempting identification. I also

found the terminology ‘equivalent, alternative names’ confusing. Are these

synonyms? Elsewhere the intricacies of taxonomy have been very simply

explained, why not here? It would also be clearer to include synonyms in

the checklist of species indicating to which species they belong.

This is really a landmark publication and it will have wider use in the

neighbouring countries of Thailand, Cambodia and Vietnam, where similar

publications are not yet available. It is also a model for other user-friendly

guides for the non-specialist besides setting a new trend of involving local

botanists in its production, so that the project has not only produced a

publication but has also encouraged local expertise. This latter is essential

if rattans are to have a future, either from the viewpoint of their sustainable

production or their continued existence.

Ruth Kiew

Singapore Botanic Gardens