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_ THE GARDENS’ BULLETIN SINGAPORE

The Gardens’ Bulletin Singapore publishes original papers on plant taxonomy

(including revisions), horticulture, phytogeography, floristics, morphology,

anatomy and related fields with emphasis on plants in the West Malesian

region.

’ Dr. B.C. Tan Dr. Jana Leong-Skorniékova

Singapore Botanic Gardens Singapore Botanic Gardens

(Editor) (Assistant Editor)

Dr. PY. Tan Ms. C. Soh

National Parks Board Singapore Botanic Gardens

(Assistant Editor) (Journal Business Manager)

EDITORIAL BOARD

Dr. S.C. Chin Dr. M.C. Roos

Singapore Botanic Gardens National Herbarium Netherlands

Singapore Leiden University, The Netherlands

Dr. M.J.E. Coode

Royal Botanic Gardens

Kew, U.K.

Prof. Sir P. Crane

Yale University

Dr. E. Soepadmo

Forest Research Institute Malaysia

Kepong, Malaysia

Prof. T. Stuessy

University of Vienna

New Haven, USA Austria

Dr. R.T. Corlett Dr. W.K. Tan

University of Hong Kong National Parks Board

Hong Kong Singapore

Dr. W.J. Kress Dr. I.M. Turner

Department of Botany, NUNH Research Associate

Smithsonian Institution Singapore Botanic Gardens

Washington DC, USA

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

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

conclusions expressed by the contributing authors.

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

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

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

Singapore.

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

[Cover photo : Rafflesia aurantia (see p. 20); photo by Danilo S. Balete]

M DEC 04 2008

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The Gardens' Bulletin

Singapore

VOL. 61 (1) 2009 ISSN 0374-7859

CONTENTS

Argent, G.

Rhododendron sojolense Argent (Ericaceae), A New Species of Rhododendron Subgenus Vireya

AEG Ad STN AES UMN OME Serer encase tos ane sans Sean far sas tence cosecasencaateasateoedestees evevedessvetenvensdstenstesvinss |

Ashton, P.S.

Additional Novel Taxa of Syzygium and Tristaniopsis from Sabah and Sarawak, and a Surprising

DIS CONME GN MOIMS OTC ICON LOL am aretee ae cee cea Me oS ese e Set Soh Snook vie 8See5s en 00s sea oe enn Coe ace sees SOR eas sR 7

Barcelona J.F., L.L. Co, D.S. Balete and N.A. Bartolome

Rafflesia aurantia (Rafflesiaceae): A New Species from Northern Luzon, Philippines .............. 17

Hughes, M., Deden Girmansyah, Wisnu Handoyo Ardi and Nurainas

SGVEHINE WES DECIES: OL FCC OMIGmil OM SWIM AA nczs.ces-scnss-ceneesse5esesses22-: <<< croasdesseeeee cane sanstastos «santsse 29

Huyen, D.D.

A New Record for the Flora of Vietnam: Geodorum citrinum Jacks. (Orchidaceae) ................. 45

Iwatsuki, Z., K.-T. Yong and T. Suzuki

A New Species of Fissidens (Bryopsida, Fissidentaceae) from Peninsular Malaysia ................ 49

Kato, M. and S. Koi

Taxonomic Studies of Podostemaceae of Thailand. 3. Six New and a Rediscovered Species ... 55

Kiew, R.

Three New Species of Gesneriaceae from Kelantan, Malaysia ..................::cesceccesseeseeeeeeeeeseees 18

Kurzweil, H.

A’Review of the Genus Plocoglottis (Orchidaceae) im Thatland ..............2..<..<c.0000esessasoesesssoesnees 81

Lee, C.C.,S. Mcpherson, G. Bourke and M. Mansur

Nepenthes pitopangii (Nepenthaceae), a New Species from Central Sulawesi, Indonesia ........ 95

Low, Y.W. and K.M. Wong

Old Hats are Better: New Considerations and Taxonomic Changes in the Southeast Asian Gardenia

EUDT CGE OMIplex (RUDIACE AG) ooo 5-2 cs hens nen nn nnce ines svasssnteszisascascoasnseaessotensseesisdabeensacescdesssseaa=ees 101

Neamsuvan, O., T. Seelanan and J.F. Veldkamp

A Revision of Bothriochloa Kuntze (Poaceae) in Thailand .....................s.seseseeseeeeceecccececceeeeeeee 129

Neamsuvan, O., T. Seelanan and J.F. Veldkamp

A Revision of Hemisorghum (Poaceae, Sorghinae) in Thailand ............cccccesccseeseeseeteeeseeeseees 145

Rosing, W.C.

CorticolousiMyxomycefesiom sin ga pone meres escort tentee en ae eee ee een eee 151

Suksathan, P. and P. Triboun

Ten New Species of Impatiens (Balsaminaceae) from Thailand ................e:cesccecesecesreeseeereeees 159

Turner, I.M.

A New Species of Alphonsea (Annonaceae) from BOrmme6 ..-<.....-.--<..:-2-c00ceeeceeu-eooeeorerueeseeeneeses 185

Turner, I.M. and J.F. Veldkamp

AEs tony iol Cananeal(AnmOnaceae) escesess:-eesecsectesersecenee none sear eeneeese re eee ae eee 189

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

Miscellaneous*@ucurbitiNe ws: lll csscecccsteccncrsssrcestectecc enon rener eee ee Re cn nS ene en ene 205

Wong, S.Y., P.C. Boyce and J. Bogner

Studies on Schismatoglottideae (Araceae) of Borneo VIII: A Review of Piptospatha elongata in

SALAWAK \secccracese cossasazecrossesuccusvisvenecesceesscessetisnssceeeee ¥1n0 cs De eNa cate Sete t eevee ene eee RT ee 217

Book Reviews

Ferns of Kinabalu, an Introduction by J.H. Beaman and P.J. Edwards.

BoC Vann ove beae sass ccsepeuescensnesaceacnaenstcssegusodtscaeucs icdesusens uss. carecetyctectsometecterec ester acereet eieeeeaeenenees 235

An Annotated Checklist of Southeast Asian Begonia by M. Hughes

Date of publication: October 2009

Published and copyrighted by

National Parks Board

Singapore Botanic Gardens

1 Cluny Road

Singapore 259569

Printed by Oxford Graphic Printers Pte Ltd

Gardens’ Bulletin Singapore 61 (1): 1-6. 2009 il

Rhododendron sojolense Argent (Ericaceae),A New Species of

Rhododendron Subgenus Vireya from Sulawesi, Indonesia

G. ARGENT

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

Abstract

Rhododendron sojolense Argent (Ericaceae) is described as a new species

of subgenus Vireya section Euvireya from Mt. Sojol on the northern arm

of the island of Sulawesi, Indonesia. Comparisons with related species are

made.

Introduction

There has been considerable recent botanical fieldwork on the island of

Sulawesi in recent years, which has produced a number of new taxa. It is an

island of extraordinary complexity in its geological history and biological

diversity (Mendum and Atkins, 2004). Professor Sleumer’s Flora Malesiana

account (Sleumer, 1966) described 25 species of Vireya Rhododendron

of which 19 were considered endemic. This number was increased to 28

species in a review of the Rhododendrons of Sulawesi (Argent, 2007), but

remarkably this increase in number was entirely with records of non-endemic

species, which underlined biogeographical links with Borneo, Philippines

and Maluku. However the species described here is clearly a new endemic.

It keys in Sleumer (1966) most closely to Rhododendron lompohense J.J.Sm.

also from Sulawesi, but R. Jompohense differs from R. sojolense in having

white tubular-funnel shaped flowers and rounded not tapering leaf bases. In

Argent (2007), it keys to R. leptobrachion Sleumer to which it would appear

to be more closely related. It differs markedly from this species in that the

leaves are revolute, not involute, with shorter and much broader apices; the

blades are much broader, about twice as long as broad and not three times

as long as broad as in R. leptobrachion, and the leaf base although acute,

is not decurrent as it is in R. leptobrachion (for about half the length of

the petiole in that species). The lateral buds, which are prominent in both

species, are broad and rounded in the new species, whereas they are slender

and very acutely pointed in R. leptobrachion, and the flowers are larger,

mat pink rather than glossy red. The bracts of the new species are glabrous

outside, not hairy as described for R. leptobrachion, although the cultivated

material of R. leptobrachion at RBGE also has the bracts glabrous outside

Gard. Bull. Singapore 61 (1) 2009

as in this new species. Cultivated material of R. leptobrachion in Edinburgh

agrees in all other respects with the type specimen of this species held in the

Riksherbarium, Leiden. R. /eptobrachion has been recorded from Mt. Sojol

(Binney, 2003) and it has not been possible to check these claims. It would

appear likely that these plants too (probably only collected as living plants)

were this new species as all previous records of R. leptobrachion are from

the central mountains in Sulawesi.

Rhododendron sojolense flowered for the first time in cultivation at

the Royal Botanic Garden Edinburgh in May 2005, but it was not considered

sufficiently mature to describe until it was seen to be flowering regularly. The

cultivated plants are now nearly | m high. They grow easily and at present

flower just once a year in late Spring or early Summer. Three seedlings were

collected in the wild all from the same general locality. The two additional

(non-type) collections differ only in having green rather than pink flower

buds. Mt. Sojol the locality for this new species is an isolated peak on the

northern arm of Sulawesi. It is a remote location requiring several days walk

to reach the montane forest and it has not been well collected. The lower

slopes have been greatly denuded of vegetation by agriculture, but there is

still a good area of humid mossy forest around the summit.

Rhododendron sojolense Argent, sp. nov.

Rhododendroni leptobrachioni similis, a qua alabastris lateralibus brevibus

rotundatis haud tenuibus acutis, foliis revolutis non involutis apicibus multo

latioribus brevioribusque et corolla longiore rosea non rubra differt. —

Holotypus: Indonesia, Sulawesi, Mt Sojol, in mossy forest, 2000-—3000m,

locally common as an epiphyte (BO; iso, E,L. SING). Figs. 1-3.

Figure 1. Colour illustration of the inflorescence of Rhododendron sojolense (based on type

specimen).

Rhododendron sojolense Argent (Ericaceae) 3

Figure 2. A. Habit with inflorescence; B. Pistil with pedicel: C. Stamen; D. Corolla viewed

from the inside with stamens; E. Inflorescence bud: F. Close-up to show scales on the

underside of the leaf. All drawings based on type specimen.

4 Gard. Bull. Singapore 61 (1) 2009

EHT = 5.00 kV Signal A = SE: Date :22 Apr 2008 2s

File Name = 20000498 young 1.tif

* = Pa

EHT = 5.00kV Signal A

1 = SE2 8

= WD= 10mm __ File Name = 20000498 young 2.tif ays

Figure 3. A. Scanning electron microscopic view of the leaf underside showing the

distribution of the scales; B. SEM close-up showing the shape of a typical leaf scale (based

on type specimen).

Rhododendron sojolense Argent (Ericaceae)

Subgenus Vireya, section Euvireya

Erect shrub. Twigs rounded, 3-5 mm in diameter, laxly silvery scaly, becoming

glabrescent, without simple hairs, purplish-red when fresh, with prominent

rounded lateral buds in the axils of the upper leaves and occasionally along

the stems; internodes 5-12 cm. Leaves 3-6 together in loose pseudo-whorls

at the upper 1-3 nodes. Blade 65-180 x 29-55 mm, elliptic, broadly elliptic,

or slightly obovate; apex broadly acute to obtuse, occasionally broadly

apiculate; margin entire, strongly and broadly revolute, often to half the

width between midrib and margin; base broadly to narrowly tapering,

densely scaly on both sides when young, glabrescent above, more persistently

and laxly to subdensely scaly beneath. Scales silvery, with a relatively wide

transparent marginal zone star-shaped to subcircular, centre minute and

slightly impressed, brown. Mid-vein narrow raised above in the proximal

1/3, impressed distally, strongly and obtusely prominent beneath for most

of its length, often purplish; lateral veins 6-10 per side, irregular, spreading

at ca 45°, obscurely inarching, with additional shorter intermediate veins all

minutely raised, minutely grooved above, more slender and hardly raised

beneath. Petiole 14-25 x 3-4mm,rounded above, purple, densely scaly. Flower

buds to 32 x 15mm, ovoid to conical-ovoid, smooth, dull red, glabrous apart

from some scales on the bract margins. Outer bracts to 25 x 12 mm, ovate

to ovate-oblong, the outer often splitting to become emarginate, inner ones

spathulate, glabrous inside. Bracteoles 15-20 x ca 0.3 mm, linear, densely

hairy. Inflorescence 7-10-flowered, an open umbel, the flowers half-hanging

to hanging. Pedicels 15-24 x ca 1.25 mm, pink, slender, laxly to densely scaly,

without hairs. Calyx 3-4 mm in diameter, obliquely disc-shaped, densely scaly

outside. Corolla 45-50 x 30-35 mm, tubular, bright matt pink; tube 25-38 x

9-12 x 10-12mm, cylindrical, straight, very prominently and deeply sulcate

in the proximal 42, glabrous outside, sub-densely shortly hairy inside with

retrorse (proximally pointing) hairs; lobes 12-15 x 12-15 mm, sub-circular or

emarginate, overlapping to 3/4, half-spreading. Stamens irregularly arranged,

exserted to ca 10 mm; filaments linear, pink, densely sub-patently hairy in

the proximal 3/4, glabrous distally; anthers brown, ca 3 x 1 mm, oblong. Disc

prominent, yellow, deeply lobed, long hairy on the upper side otherwise

glabrous. Ovary 8 x 4 mm, sub-cylindrical, green, but densely covered in

long appressed silver hairs, which completely cover small scales, tapering

distally; style slender, deflected to the lower side when young, pale pink, ca

26mm, hairy in the proximal 2/3 not scaly; stigma capitate. Fruit 20-30 x 5-6

mm, cylindrical, slightly curved, sub-densely hairy and scaly. Seeds 3.8 mm,

without tails 0.8-1 mm, the longest tail 1.8mm, the tails highly crimped.

The type specimen was collected by Smith and Galloway (# 101,

RBGE Accession 20000498) as living material on 27 Feb 2000. The plant

6 Gard. Bull. Singapore 61 (1) 2009

flowered in cultivation in Edinburgh, May-June. Additional materials:

RBGE accessions 20080948 & 20080949. Both from the same locality.

Acknowledgements

I would like to thank Louise Galloway, Mary Mendum and Paul Smith for

their companionship on the expedition on which this species was collected

and for safely collecting and growing the living material upon which the

description is based. I am grateful to Anna Dorwood for ably drawing Fig.

1; Frieda Christi for producing the scanning electron micrographs and to

Robert Mill for rendering the diagnosis into Latin. The colour photograph

was taken by Lynsey Wilson. The expedition on which this plant was collected

was supported by LIPI the Indonesian Institute of Sciences (Lembaga

IImu Pengetahuan Indonesia) and funded by the Royal Botanic Garden

Edinburgh.

References

Argent. G.C.G. 2007. Rhododendrons of subgenus Vireya. Royal

Horticultural Society, London, pp. 388.

Argent, G.C.G. 2007. Rhododendrons of Sulawesi. Buletin Kebun Raya

Indonesia 10(1): 20-23.

Binney. D. 2003. Rhododendron Collecting in Sulawesi, Indonesia, pp. 111-

114. In:G. Argent & M. McFarlane (eds.), Rhododendrons in Horticulture

and Science. Royal Botanic Garden, Edinburgh.

Mendum, M. and Atkins, H. J. 2004. The Gesneriaceae of Sulawesi I.

Edinburgh Journal of Botany 60(3): 299-304.

Sleumer, H. 1966. Rhododendron. Flora Malesiana 6: 474-668.

Gardens’ Bulletin Singapore 61 (1): 7-15. 2009 4

Additional Novel Taxa of Syzygium and Tristaniopsis

from Sabah and Sarawak, and a Surprising Discovery

of Shorea contorta

P.S. ASHTON

Arnold Arboretum, Harvard University and Royal Botanic Gardens, Kew

Abstract

A further five species and two subspecies of Syzygium, and one subspecies

of Tristaniopsis (Myrtaceae) are described as a precursor to the account of

the family in the Tree Flora of Sabah and Sarawak. Shorea contorta Vidal is

here recorded form Borneo for the first time.

Introduction

Since publishing new species of Syzygium (Ashton, 2007) and Tristaniopsis

(Ashton, 2007) as precursors to my forthcoming account of Myrtaceae for

the Tree Flora of Sabah and Sarawak (Soepadmo et al. (eds.), continuing),

I have had the opportunity to examine the unidentified material at the

herbaria of Singapore (SING) and Forest Research Institute Malaysia

(KEP). Additional novelties, published here, have inevitably resulted, though

reassuringly few. Striking is the addition of one further entity apparently

endemic to the astonishing ultramafic Bukit Tawai Forest Reserve of NE

Sabah, already replete with the endemics, S. cornuflorum P.S.Ashton, S.

soepadmo P.S.Ashton and Tristaniopsis merguensis (Griff.) PeterG. Wilson &

J.T. Waterh. ssp. tavaiensis P.S.Ashton. All these species have morphologically

similar sister entities, often in nearby forests. A further novel species confined

to ultramafics is more widespread in NE Sabah. These distributions, and

many others published in the Tree Flora in other families, emphasize the

important opportunities that Bukit Tawai provides for molecular and other

approaches to examine speciation processes in tropical rain forest, and the

vital necessity of satisfactory conservation and protection from fire, which

has been a scourge of smaller ultramafic sites in Northeast Borneo.

This communication also provides me with the opportunity to

make an unexpected new addition to the Dipterocarpaceae of Sabah and

Sarawak, whose account in the Tree Flora was already published (Ashton,

2004): Shorea contorta Vidal |syn. Pentacme contorta (Vidal) Merrill &

Rolfe, SAN 127272, Awang Amin Segun, Kepayang, Kuala Penyu, Sabah

(K)]. The label states the habitat solely as ‘lowland’. The tree was 10 m tall.

8 Gard. Bull. Singapore 61 (1) 2009

The specimen consists of the leafy shoot of an immature individual, with

the distinct narrow raceme of the species, and equally distinct fallen fruit,

glabrous throughout again suggesting immaturity. Kuala Penyu is on the

coast of SW Sabah, at the northernmost tip of the Klias peninsula. The Klias

peninsula is close to the northernmost extension in the coastal lowlands of

the Neogene basin, which, running southwest to the Lambir Hills in Miri

District, NE Sarawak, is so rich in tree species endemic to yellow sandy

soils. This is the habitat which, with many mixed peat swamp entities,

characterizes the geographical distribution named by Corner (1960) the

Riau Pocket. This habitat is therefore quite different from the fertile clay

loams so characteristic of the Philippine lowland mixed dipterocarp forest

within which S. contorta was one of the most widespread and characteristic

species. It is possible though that the collecting locality lies further inland,

on clay loams more similar to many Philippine soils. With its recalcitrant

seed and demanding seedling, it seems unlikely to have been introduced,

though that cannot be ruled out. Otherwise, this is an example of a rare

sweepstakes dispersal event, as unlikely in its way as that of the baobab,

which have dispersed from Madagascar onto Australia. I beseech any visitor

to Kuala Penyu to seek out this plant again: molecular genetic evidence

might be revealing.

1. Syzygium claviflorum (Roxb.) Walpers ex Steudel, Nomencl. Bot., ed 2

(1841) 657.

Basionym: Eugenia claviflora Roxb., Fl. Ind. Ed. 2,2 (1832) 488. — Lectotype:

Maynmar, E. Chittagong, Wallich 3575B (K).

-Syzygium longiflorum Wall., Cat. (1831) 3575B, nom. nud.

Note: E. claviflora Roxb. Is, first, a validly published name, as is evident

from the specimen of Wallich 3575B in the East India Company Herbarium

at K. There are two sheets: one with two specimens in early post-anthesis,

numbered A and B, the other a single fine specimen in anthesis numbered

3575B and annotated Syzygium claviflorum Wall., and Eugenia claviflora

Roxb. in Roxburgh’s handwriting. A further label is attached in another

hand, noting E. Chittagong as provenance. This second specimen is therefore

formally recognized here as lectotype.

la. Syzygium claviflorum subsp. tavaiense P.S.Ashton, subsp. nov.

Subspecies novum a species typico ramulis crassioribus lamina ca 13 x

6 cm ampliore coriacissima petiolo crassiore costis tertiariis supra magis

prominentibus elevatis floribus minoribus differt. — Holotypus: Borneo,

Sabah, Sandakan, Telupid, Bukit Tawai Forest Reserve, Zainudin 5012, in

flower (KEP)

Additional Novel Taxa of Syzygium and Tristaniopsis from Sabah and Sarawak )

Small smooth-barked tree. Differing from ssp. claviflorum as follows: Twig

at first ca 3 mm diameter, stout; leaf blade ca 13 x 6 cm, broader, thickly

leathery, base obtuse to broadly wedge-shaped hardly tapering into stalk,

stalk ca 3 mm diameter; secondary veins not furrowed above, tertiary veins

more distinctly raised above than beneath; flower to 8 mm long smaller.

Distribution and ecology: Based ontwo collections apparently simultaneously

gathered from the same tree, the type and Bojo & Cheksum OB 040, from

Tawai F.R., on ultrabasic substrate.

2. Syzygium flagrimonte P.S.Ashton, sp. nov.

Syzygio napiformi (Koorders & Valeton) Merr. & L.M. Perry affinis sed frutex

lamina minima obtusa petiolo 4 mm longo caudice receptaculi gracillimo

facile distinguitur. — Holotypus: Borneo, Sarawak, Mulu National Park,

Gunung Api, summit ridge, 1600 m, Argent & Jermy 1017 (A).

Shrub to 80 cm tall. Parts hairless. Twig ca 1 mm, round at first, round,

wrinkled, drying blackish. Leaf blade 15-30 x 10-22 mm, elliptic, thinly

leathery, black dotted beneath, pitted above, drying pale ochreous brown;

base wedge-shaped terminating abruptly at ca 4 mm long, slender black-

drying stalk; apex obtuse to subacuminate: veins dense, subequal, main veins

8-10 pairs, ascending, obscure above, slender but distinctly elevated beneath:

tertiaries obscure; intramarginal vein within | mm of margin. Flower (post-

anthesis) to 11 x 4 mm, comprising a 3 x 4 mm tapering, pale, buff, drying

receptacle on aca 1 mm diameter, slender, tapering pseudostalk; sepal lobes

4.0.8 x 0.8 mm, small, | deltoid distant around the calyx rim; style extended

to ca 4 mm. Fruit unknown.

Distribution and ecology: Known only from the type, in upper montane

forest on the exposed karst and rocky summit ridge of Gunung Api (‘fire

mountain’ — probably on account of occasional lightning fires implying

occasional severe drought): at 1600 m.

Note: A distinctive upper montane, apparent a point endemic within the S.

attenuatum group, distinguished from S. napiforme, which appears to be its

sister species, by its habit and leaves, and unusually slender pseudostalk.

3. Syzygium georgeae P.S.Ashton, sp. nov.

Syzygio castaneo (Merr.) Merr. & L.M. Perry affinis sed lamina ovata cordata

nervis supra depressis lobis calycis in flore distinctis ovatis valde differt. —

Holotypus: Borneo, Sabah, S. Meliau, Telupid, Tawai Forest Reserve, E.

Soepadmo, T: Khalkausas, R. George, B.S. John & S. Davol (FRI 41256, SING).

10 Gard. Bull. Singapore 61 (1) 2009

A small tree with many scrambling branches, to 10 m tall, 15 cm diameter,

with smooth grayish white bark. Parts hairless. Twig 1-2 mm diameter

apically, slender, at first distinctly narrowly 4-winged. Leaf blade 4.5-6 x

2-3.2 cm, ovate-lanceolate, thinly leathery, satiny, drying golden-brown and

finely black dotted beneath, dark yellow-brown and densely minutely pitted

above; base shallowly cordate, subsessile with a stalk ca 1 mm long, drying

black; apex acute to subacuminate; venation distinct beneath, drying blackish,

forming a net with the tertiaries, veins subequally slender and hardly raised

beneath, distinctly furrowed above, ca 11 main pairs with many irregular

intermediates branching among the tertiaries; intramarginal vein ca | mm

within margin, hardly arched. Panicle to 3-terminal, to 5 cm long, ca | mm

diameter in flower, quadrangular; bracts minute caducous. Flower bud ca 4

x 3 mm, small, club shaped with slender pseudostalk; sepal lobes 4, ovate,

subacute, hyaline towards margins, clasping corolla; stamens many; style ca

3 mm long. Fruit not seen.

Distribution and ecology: Known only from the type. Common along a stony

occasionally flooded river bank on ultramafic substrate.

Notes: Clearly related to S. castaneum, I only recognize it from that variable

species owing to its conspicuously different leaf shape with its venation

distinctly furrowed on its upper surface. This is one more distinct sister

ecospecies of ultramafic substrates, for which the Tawai Forest Reserve

provides the richest repository.

Named in honour of Rena George, a young Sarawakian field

botanist of extraordinary promise who died in the field while following her

avocation.

4. Syzygium oblanceolatum (C.B. Robinson) Merr., Philip. J. Sc. 79 (1951)

405.

Basionym: Eugenia oblanceolata C.B. Robinson, Philip. J. Sc. 4, C (1909) 400.

— Lectotype: Philippines, Samar, /oc. incert., Cuming 1676 (BM).

Key to subspecies -

1. Leaf blade ca 23 cm long, oblanceolate; main veins ca 14 paifs .............. Je

Leaf blade to 17 cm long, elliptic, base cuneate; main veins 6-10 pairs..........

visa’ anosn dSebaaga ell UStam sa unes ae ceo CES a EER eens eee ssp. kinabaluense

2. Leaf base subcordate to cordate ............ ssp. oblanceolatum (Philippines)

Leaf base abruptly tapering, occasionally cuneate ............ ssp. kihamense

Additional Novel Taxa of Syzygium and Tristaniopsis from Sabah and Sarawak 11

4a. Syzygium oblanceolatum ssp. kihamense (Merrill & Perry) P.S. Ashton,

stat. nov.

Basionym: S. kihamense Merrill & Perry, Mem. Amer. Ac. 18 (1939) 150;

Burgess, Timbers of Sabah (1966) 412. — Holotype: Borneo, Kalimantan, W.

Kutei, near Kiham, Batu Bong, Endert 2341 (BO; iso, K).

-S. petakense Merrill & Perry, loc. cit., Beaman, Pl. Mt Kinabalu 5 (2004)

223.— Holotype: Borneo, E. Kalimantan, W. Kutei, near Long Petak, Endert

4063 (BO).

Tree. Parts hairless. Twig ca 3 mm diameter, stout with long internodes,

sharply 4-angled, smooth, dark red-brown. Leaf blade ca 23 x 6 (12-24 x

4-8) cm, large, oblanceolate, densely minutely pitted above, sparsely black

dotted beneath, drying dull rust brown, paler above: base wedge-shaped,

tapering into ca 5 mm long stout stalk; acumen to 5 mm long, broad,

tapering, twisted down; veins unequal, main veins ca 14 pairs, prominent

and distinctly raised more so beneath, furrowed along their crests above,

ascending, with few shorter intermediates; tertiaries evident throughout;

intramarginal vein 1-4 mm within margin, hardly raised, weakly arched.

Panicle axillary to ramiflorous, with to 10 cm long straight erect rachis, to 6

cm long basa! branches and shorter branches above, quadrangular. Flowers

in dense terminal clusters, subtended by 2 pairs of bracteoles; buds to 4 x 3

mm, obovoid to spherical, without distinct pseudostalk; sepal lobes 4,2 x 3

mm, broadly ovate acute, more or less warty, hyaline along margin; stamens

many; style extended 5 mm. Fruit ripening white, to 15 x 12 mm, ellipsoid,

sessile, with 7 mm diameter, prominent apical calyx crown.

Ecology: In mixed dipterocarp forest on fertile clay loams including black

volcanic soils, at low altitude, below 800 m.

Distribution: Northeast Borneo, E. Kalimantan, Pulau Laut and throughout

Sabah [Balong Tawau (San 30139); Kalabakan (San 9/444); Diwata, Lahad

Datu (San 39968): Segama Rd, Lahad Datu (San 31616); Sandakan (San

10186); Sepulut, Keningau (San 101325); Pangi (San 27439): slopes of Bt

Batanga, Sipitang (San 16622)).

Note: Related to Syzygium papillosum (Duthie) Merr. & L.M. Perry of

Peninsular Malaysia and southern Kalimantan, in which the twig is round,

leaf more leathery with deeply cordate base. S. kihamense and S. petakense,

both collected by Endert in West Kutei, East Kalimantan differ in no

consistent way.

12 Gard. Bull. Singapore 61 (1) 2009

4b. Syzygium oblanceolatum ssp. kinabaluense P.S.Ashton, ssp. nov.

A species typico lamina elliptica basim versus cuneata 6.7-17 x 3.5-7.5 cm

minore costis lateralibus utrinsecus 6-10 differt. —Holotypus: Borneo, Sabah,

Ranau, Km 58 Kinabalu-Tambunan Rd, 5°47°N, 116°20°E, 1450 m asl, in

flower, J.S. Beaman 7937 (K).

Differing from the two other subspecies as follows: leaf blade 6.5-17 x 3.5-

7.5 cm, generally elliptic, base cuneate; veins 6-10 pairs, lax in arrangement.

Ecology: Apparently quite common in lower montane forest, at 1450-1700 m.

Distribution: Endemic to Gunung Kinabalu and the northern Crocker Range

| Kundasang (San 28884); Mesilau, Kinabalu (San 48001, 42835, RSNB 4214);

Tenompok (SFN 26942); Ranau, Kinabalu N.P. (San 79580)].

5. Syzygium praestantilimbum P.S.Ashton, sp. nov.

Syzygio rosulento (Ridl.) Merr. & L.M. Perry clare affinis sed arbuscula

partibus glabris lamina enorme coriacissima petiolo ca 8 mm longo paniculis,

4m longis conspicue distinguitur.— Holotypus: Borneo, Brunei Darussalam,

Labi, path to Bukit Teraja, Peng. Mohamad (BRUN 18339, SING).

Treelet to 8 m tall, occasionally reaching the canopy, 60 cm diameter, with

drooping branches bearing the huge leaves on long internodes. Parts hairless.

Twig ca 3 mm diameter, round to compressed, dark warm brown. Leaf blade

ca 25 x 14 (20-35 x 12-16) cm, elliptic, leathery, dull, drying mauve-brown

obscurely minutely pitted above, rust-brown obscurely minutely dotted

beneath; base wedge-shaped terminating abruptly at the 7-12 mm long, 3

mm diameter, stout black-drying stalk; apex to 12 mm, slender acuminate;

veins unequal, main veins ca 17 pairs with a few shorter less prominent

intermediates, deeply furrowed above, prominent beneath; tertiaries lax,

more or less distinctly raised beneath, hardly sunken above; intramarginal

veins 1(2) pairs, somewhat arched, ca 6 mm within margin. Inflorescence to

4 cm long, 3-terminal or 1-axillary, round, singly branched, flowers clustered

at branch endings; bracts and bracteoles to 2 x 2 mm, ovate-deltoid, acute,

subpersistent. Post-anthesis flower calyx 5 x 4 mm, obovoid to top-shaped

without distinct pseudostalk, with 4,2 x 2 mm deltoid sepal lobes, hyaline

at margins, becoming reflexed; stamens many; style extending 4 mm. Young

fruit to 6 mm diameter, spherical, subsessile and densely clustered, wrinkled

on drying, ripening pinkish white.

Ecology: Local but sometimes abundant as an understorey tree, in mixed

dipterocarp forest on deep yellow sandy soils, in a habitat shared with other

o>)

Additional Novel Taxa of Syzygium and Tristaniopsis from Sabah and Sarawak 1

understorey trees sharing large leathery leaves, including S. velutinum A.P.

Davis.

Distribution: Known from the type, Andulau Forest Reserve in Brunei

Darussalam (BRUN 18283) and from sterile collections (Field herbarium

numbers AG 02747, AG 03847, AG 3210-058) from the large research plot in

Lambir N.P. at Miri in Sarawak.

Note: Closely resembling S. rosulentum in the herbarium but a small

subcanopy tree with much larger leathery leaf and shorter inflorescence.

The inflorescence in both species is, as Merrill defines it in his key to the

genus (1939, 141), with flowers in dense heads, each flower ‘subtended by

two decussate and approximate pairs of bracts’. Examination of atypical

inflorescences of species with this character persuades me that it is caused

by extreme shortening of the terminal branchlets of a panicle. The two taxa

co-exist in the same stands.

6. Syzygium tubiflorum P.S.Ashton, sp. nov.

Syzygio zeylanico (L.) DC.,S. kinabaluense (Stapf) Merr. & L.M. Perry late

affinis sed lamina nervis obscuris calyce in flore fructuque leve terete satis

distincta. — Holotypus: Boreno, Sabah, Maliau Basin, top ridge, 1220 m, coll.

Maig, Sidikin & Jeprin (MB 904, SING).

Treelet to 5 m tall. Parts hairless. Twig 0.5-1 mm diameter apically, very

slender, narrowly but distinctly 4-winged, drying dark brown to blackish,

smooth. Leaf blade 15-40 x 3-15 mm, lanceolate, small, leathery, drying pale

grey-brown, faintly densely pitted above, densely prominently black dotted

beneath; base wedge-shaped, tapering into ca 4 mm long, 0.5 mm diameter,

very slender stalk; apex to 12 cm long slender acuminate; venation obscure

above the sunken midrib excepted, unequal, the ca 6 pairs of irregularly

disposed main veins evident beneath only, slender and hardly raised.

Inflorescence ca 15 mm long, | mm diameter, erect, singly branched, round

terminal or axillary panicle; bracts to 7 x 1 mm, lorate-lanceolate, falling

early. Flowers unknown. Young fruit 5 x 2 mm, torch-shaped, subsessile,

without distinct pseudostalk, without ribs, smooth, drying black; sepal lobes

4, deltoid, minute, falling early; bracteoles ca 3 mm long, linear, fugaceous.

Distribution and ecology: Known only from the type, from the Meliau

summit ridge at 1220 m.

Note: A distinct entity in the S. zeylanicum group, but at once distinguished

by its leaf venation, smooth, round young fruit, and early falling bracts and

14 Gard. Bull. Singapore 61 (1) 2009

bracteoles. As a member of that group and as the fruit indicates, the flower

for which it is named, though as yet unknown, must possess an elongate

tubular psedostalk devoid of the sharp angles characteristic of the elongate

pseudostalks of sister species.

7. Syzygium valentissimum P.S.Ashton, sp. nov.

Lamina magna coriacissima costis lateralibus subaequalibis ramulis

quadrangularibus paniculis erectis crassis gemmuis obconicis facile distinguitur.

— Holotypus: Borneo, Sabal-Balai Ringin Forest Reserve, Klingkang Range,

Sri Aman, in flower bud, Julathi & al. (S. 83369, KEP).

Canopy tree to 30 m tall; bark brown. Parts glabrous. Twig ca 4 mm diameter

apically, at first quadrangular, smooth, dark chocolate-brown. Leaf blade

7-15 x 5-6 cm, ovate to lanceolate, thickly leathery yet margin hardly inrolled,

drying dark chocolate-brown dull below, somewhat glistening above;

base broadly wedge shaped tapering into ca 15 mm long, 3 mm diameter,

stout black-drying stalk; apex to 8 mm long down-turned acuminate; veins

subequal, main veins ca 18 pairs with many intermediates, tertiary veins

reticulate, venation overall finely but distinctly more or less equally raised

on both surfaces. Panicles to 5-terminal or subterminal axillary, to 11 mm

long (immature), ascending, stout, quadrangular, 2-branched. Flower bud

(young) to5 x 4mm, obconical without distinct pseudostalk; sepals 4, ca 3 x

4 mm, broadly hemispherical, obtuse, cupped, broadly imbricate and tightly

clasping corolla; stamens many. Fruit unknown.

Distribution and ecology: On the lower sandstone slopes, in mixed dipterocarp

forest, of the Kingkang range (the type) and Gunung Buri (S. 4341/4, Ilias

Pa’ie), respectively to the north and south of the Serian-Sri Aman road in

West Sarawak.

Note: At once distinguished by its relatively large thickly leathery leaf with

subequal veins, quadrangular twigs, and stout erect panicles with obconical

buds. The leaves recall S. selukaifolium P.S.Ashton of NE Sarawak in

which the twig is terete and the panicle axillary with torch-shaped buds,

and S. merrillii (C.B. Rob.) Merr., which shares the buds and panicle of S.

selukaifolium, but in which the twig is also quadrangular.

8. Tristaniopsis microcarpa P.S.Ashton, Gard. Bull. Sing. 57 (2005) 273.

— Holotype: Borneo, Look Mengulang, P. Sakar, in flower, H.S. Martyn (San

DNOD SKS):

Additional Novel Taxa of Syzygium and Tristaniopsis from Sabah and Sarawak 15

8a. Tristaniopsis microcarpa ssp. corymbosa P.S.Ashton, ssp. nov.

Tristaniopside microcarpa late similis sed lamina coriacissima, ramuli caduci

lamina infra inflorentia alabastrique persistenter alutaceo-cinerascentibus.

— Holotypus: Borneo, Sabah, Telupid, mile 872, Hap Seng logging area, in

flower, Kodoh & Tarmiyji (San 8366, K).

Small tree to 20 m tall, to 40 cm diameter, with red brown peeling bark.

Twig at first, leaf beneath, inflorescence and flower bud persistently densely

buff cinereous. Leaf blade (8-)16-35 = (3.5-) 6-12 cm, oblanceolate, thickly

leathery.

Distribution and ecology: Endemic to ultramafic substrates in eastern Sabah,

on rocky sites often beside rivers [the type; Telupid, Bt. Tawai (SAN 13451,

FRI 41309); Telupid, S. Ruku-Ruku (SAWN 703/5)|; also at 850 m on the Mt.

Silam summit (SAN 63945) and in lower montane kerangas at 1000 m on G.

Lutong, Meliau basin (FRI 36238).

Notes: Differing from subspecies microcarpa only in leaf texture and

indumentum, and possibly smaller stature at maturity. It appears to occur on

shallower, more humic soils. A specimen (FRI 4/309) was misidentified as T-

merguensis ssp. tavaiensis in the original description of that entity (Ashton,

2006).

References

Ashton, P.S. 2004. Dipterocarpaceae. In: Soepadmo, E., Saw, L.G. & Chung,

R.C.K, Tree Flora of Sabah and Sarawak 4: 63-388.

Ashton, PS. 2006. New Tristaniopsis Peter G.Wilson & J.T. Waterh

(Myrtaceae) from Borneo. Gardens’ Bulletin Singapore 57: 269-278.

Ashton, P.S. 2007. New Syzygium (Myrtaceae) from northern Borneo. Kew

Bulletin 61: 107-144.

Corner, E.J.H. 1960. The Malayan flora, pp. 21-24. In: Purchon, R.D. (ed.),

Proceedings of the Centenary and Bicentenary Congress of Biology,

Singapore.

Merrill, E.D. 1939. The Myrtaceous genus Syzygium Gaertner in Borneo.

Memoirs of the American Academy of Arts and Science 18(3): 133-202.

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Gardens’ Bulletin Singapore 61 (1): 17-27. 2009 77

Rafflesia aurantia (Rafflesiaceae): A New Species from

Northern Luzon, Philippines

HEEBARCELONA IAL. CO! . DS. BALERE >

AND N.A. BARTOLOME ”

‘Philippine National Herbarium (PNH), Botany Division, National Museum of the

: Philippines, P. Burgos St., Manila, P.O. Box 2659, Philippines.

~ Conservation International-Philippines, 6 Maalalahanin St., Teacher’s Village,

Diliman, Quezon City 101, Philippines.

* Herbarium, Institute of Biology, College of Science, Diliman,

Quezon City 1101, Philippines.

“Institute of Biology, University of the Philippines, Diliman 1100,

Quezon City, Philippines.

* Department of Zoology, Field Museum of Natural History, 1400 South Lake

Shore Drive, Chicago, IL 60605, U.S.A.

Abstract

A new Philippine species of Rafflesia from the Sierra Madre Mountain

Range of northeastern Luzon is described. Rafflesia aurantia is the ninth

presumed endemic species thus far described from the Philippines, and the

fifth reported from the island of Luzon. This species is apparently allied to

R. tengku-adlinii of Sabah - both are small-sized and their overall color is

similar. Biogeographical considerations and the morphological differences

between our new species and R. tengku—adlinii, however,strongly support the

recognition of two distinct evolutionary lineages/species. The conservation

status of the fast disappearing lowland dipterocarp forests in northeastern

Luzon, particularly the type locality in the Quirino Protected Landscape

(QPL) is also discussed and suggests that the new species may be highly

threatened.

Introduction

The Philippines is fast emerging as the global center of Rafflesia diversity

relative to its land area with the recent description of five more new species

after the publication of R. speciosa Barcelona & Fernando (2002) from

Antique Province in Panay Island. These newly described species are, R.

mira Fernando & Ong (= R. magnifica Madulid, Tandang & Agoo), from

Compostela Valley Province in Mindanao reported in 2005, R. baletei

Barcelona & Cajano from Camarines Sur Province in Luzon (Barcelona

et al., 2006), R. lobata Galang & Madulid (2006) from the Central Panay

18 Gard. Bull. Singapore 61 (1) 2009

Mountains, and R. /eonardi Barcelona & Pelser from Cagayan Province in

Luzon (Barcelona et al., 2008).

The recentrediscoveries of R. manillanaTeschem.(MadulidandAgoo,

2008; Madulid et al., 2008) and R. philippensis Blanco [as R. banahawensis

Madulid, Villariba-Tolentino & Agoo (2007) and as R. banahaw Barcelona,

Pelser & Cajano (2007; see also Barcelona et al.,2009)] in their type localities

as well as new populations of the presumed extinct R. schadenbergiana

in Mindanao (Lays, 2006; Barcelona et al. 2008a, 2009) has tremendously

increased our knowledge of the diversity and geographic distribution of this

enigmatic plant group. A summary of our current knowledge on Philippine

Rafflesia including their taxonomy, ecology, geographical distribution, and

conservation status is detailed in Barcelona et al. (2009).

In 2004, a biodiversity survey that we conducted in the lowland

forests of the western edge of the central portion of the Sierra Madre

Mountain Range in northeastern Luzon, now designated as the Quirino

Protected Landscape (QPL) in Quirino Province, resulted in the discovery

of yet another population of Rafflesia. Comparison with currently known

Philippine Rafflesia species and with that of the morphologically similar R.

tengku-adlinii of Borneo strongly supports the recognition of a new species

from Luzon, which we describe below.

Rafflesia aurantia Barcelona, Co & Balete, sp. nov.

Rafflesiae tengku-adlinu Mat-Salleh & Latiff (ex Sabah) colori aurantae

perigonii propemodum mentiens, sed ab ea sufficienter differt processibus

disci centralis florum marium polymorphis et varie ramosis complanatis

dispositis horizontaliter, processibus florum feminarum confertis complanatis

dispositis verticaliter, ramentis longioribus (7-10 mm longis) glabris varie

ramosis apicibus capitatis nec clavatis, antheris insigniter paucioribus (12-14).

—Holotypus: Philippines. Luzon, Quirino Province, Sierra Madre Mountain

Range, Quirino Protected Landscape, Nagtipunan Municipality, Barangay

Matmad, Sitio Mangitagud, 16° 03' 22.5" N, 121° 28' 39.7" E, ca 450 m,

disturbed lowland dipterocarp forest, ca 5 m away from embankment of

boulder-strewn creek, 11 Jun 2004, Bartolome & Balete 2543 (PNH, spirit

collection, one male and one female flowers; isotypes, US, PUH, SING,

two dissected mature buds and four immature buds in different stages of

development). Figs. 1-3.

Mature flower buds 8.5-9 cm in diameter covered with atrocastaneous bracts;

cupule of mature flowers light brown, 2.2 cm tall, 6.3 mm wide. Open flower

ca 20 cm in diameter when fresh. Perigone lobes orange, 4.5-5.5 cm long, 6.3-

7.7 cm wide, distantly disposed, outer surface uniformly covered with fine

Rafflesia aurantia (Rafflesiaceae) 19

Figure 1. Rafflesia aurantia. A. Lanate ramenta en masse; B. Variably-sized and branched

ramenta; C. Magnified outer surface of diaphragm and perigone lobe showing ridged,

areoles-forming ornamentations; D. Partially-open male flower; E. Male flower disk showing

polymorphic processes with central ones vertically-flattened and horizontally oriented; F.

Semi-globular anthers embedded in deep sulci; G. Longitudinal section of a female flower

showing lunate ovary and dense, vertically-oriented, and laterally-flattened processes.

20 Gard. Bull. Singapore 61 (1) 2009

Figure 2. Rafflesia aurantia, a fully expanded flower next to a senescent one (Photo: Danilo

S. Balete).

Figure 3. Rafflesia aurantia, an early senescent flower next to a developing bud (Photo: Dani-

lo S. Balete).

Rafflesia aurandia (Raifiesiaceae ) 21

warts that are of sharp-edged_ areoles-formmg omamentations when fresh.

these turnmg into blunt ridges im alcohol-preserved specimens. Diaphragm

concolorous and similarly omamented as perigone lobes (except along the

aperture rim). pentangular. ca 10 cm across: windows absent: aperture 3-3.6

cm m diameter. Lower surface of diaphragm (except along the aperture

Margi) covered with uniformly lanate. glabrous ramenta. that are rather

sparse towards the aperture. becoming dense towards the base of the floral

cavity, 7-10 mm long. slender. unbranched to furcate. entirely glabrous. tips

swollen. Disk 55-6 cm across. ca 6 mm thick: neck of column ca 3 cm tall

Disk processes flattened. polymorphic. centrally disposed ones 5-10 mm long.

variably branched_horizontally-onented in the male flowers. dense. flattened.

and vertically-oriented in the female flower. peripheral ones much smaller.

052 mm long. narrowly lanceolate. spmose. or markedly reduced into

tubercles. tips tufted with golden brown hairs. Anthers of male flowers 12-14.

semi-slobular. each hidden im a deep sulcus; anther pores ca 25 mm across.

Ovary of female flowers lunate. ca 35 mm long. ca 65 mm wide (fig. 1A-G).

Etymology: This species is named after the orange coloration of the

flowers.

Ecology: Plants of R. aurantia have all been found parasitic on prostrate

stems and underground roois of the host lana. Tetrastigma (which

unfortunately was not collected) and has never been found yet to grow on

the climbing stems of the host. Some murid rodents that may possibly be the

seed dispersers of Rafflesia recorded on-site include Apomys, Bullimus, and

Rattus everettii, whereas flower visitors observed were metallic bottle flies.

Distribution and habitat. So far known only from the type locality. The

population was concentrated in disturbed lowland dipterocarp forest along

the lower slopes of Mungiao Mountains. in the vicinity of the headwaters

of Cagayan River. northeastern Luzon’s principal drainage system and the

Philippines longest river. With a prevailing canopy of 20-25 m and occasional

emergents to ca 40 m in height. this forest is dominated by Shorea spp. and

Anisoptera thurifera (Dipterocarpaceae). Several species of Lithocarpus

(Fagaceae). Syzygium (Myrtaceae). Litsea, and Cinnamomum (Lauraceae)

dominate the subcanopy layer. Open grasslands and agricultural areas are

located merely half a kilometer away.

Conservation status: Though observed to be common locally. this new

species is thus far known only from a single population. The sporadic

collections of plant specimens by R-C. McGregor in 1912 at nearby Dupax

to the north. and Ramos and Edafio in 1925 at Mt Alzapan to the east. are

22 Gard. Bull. Singapore 61 (1) 2009

the only historical botanical records notable in this part of Luzon. Small-

scale logging for certain premium timber species as well as forest clearings

for swidden agriculture are rather prevalent in the area. However, it is

the commercial open-pit mining (chiefly for gold and copper) that poses

the gravest threat for this Rafflesia habitat. Despite the establishment of

an approximately 206,875 ha new protected area, the Quirino Protected

Landscape (Presidential Decree No. 548 dated 9 February 2003), about

30,930 ha of these were recently excised in favor of mining, resulting in the

fragmentation of QPL into three parcels.

Taxonomic and biogeographical notes: Rafflesia aurantia closely resembles

R. tengku-adlinii of Borneo in size, flower color, habit, and the absence

of windows. It is, however, different from the latter by its lanate, entirely

glabrous, and substantially longer (7-10 mm, longest in the genus) ramenta

that are sparse towards the diaphragm and absent near the aperture rim,

its polymorphic, flattened, irregularly and sometimes horizontally disposed,

and multi-branched processes, and fewer anthers (12-14). In R. tengku-

adlinii, the ramenta extend to the diaphragm (in fact, they are visible near

- the diaphragm aperture), are significantly shorter (3-5 mm), and covered

with fine bristles. Ramenta characters, such as disposition, morphology, and

length, show little variation within species and are therefore useful characters

for distinguishing species of Rafflesia (Nais, 2001:18-19; Mat Salleh, 1991:10).

In addition to being different in characters of the ramenta, R. aurantia 1s

also distinguished from R. tengku-adlinii by disk process morphology. The

processes of R. tengku-adlinii are much reduced relative to those of R. aurantia

and they are more or less solitary and regular in disposition. Furthermore,

R. tengku-adlinii has more anthers than R. aurantia (20 in the holotype of

the former, the only specimen described and measured in the protologue).

Determining the anther number in most species of Rafflesia (except those in

R. manillana and R. lobata where the wide diaphragm aperture exposes the

anther impressions on the floor of the floral tube) is destructive in nature.

Perhaps because of its rarity, only the holotype of R. tengku-adlinii was

measured and described in detail. In addition, the fine, sharp-edged, areoles-

forming ornamentations (warts?) on the outer surface of the perigone lobes

and diaphragm in fresh flowers of R. aurantia are different from the quite

distinctive pustular warts on the perigone lobes of fresh flowers of R. tengku-

adlinii. In addition, the geographic distributions of R. tengku-adlinii (Sabah,

Borneo) and R. aurantia (northern Luzon), coupled with the generally low

dispersal ability of Rafflesia do not strongly support gene flow between these

disjunct populations to occur. The short anthesis, pollen germinability and

longevity (Nais and Wilcock in Nais 2001: 58), the low seed viability (Nais

and Wilcock, 1999 in Nais 2001: 78-80), and the obligate parasitic nature of

Rafflesia aurantia (Rafflesiaceae)

Rafflesia make gene flow unlikely, although long distance dispersal events in

the past cannot be discounted. We therefore presume that both populations

are reproductively isolated. Although Luzon may have been connected to

Northern Borneo and Sabah via the Sulu-Cagayan Arc before the Miocene,

i.e. ca. 50 million years (MY) ago (Hall, 2002: 381), there is no evidence

of the existence of Rafflesia (or its immediate ancestor) during this time.

The age of the stem-lineage of Rafflesiaceae was estimated by Davis et al.

(2007) to be 46 MY old, whereas the age of the crown group/genus Rafflesia

was estimated by Barkman ef al. (2008) to be 12MY old, with most species

divergences having occurred within the last 1-2 MY. Hence, the scenario

that both Borneo and Northern Luzon populations may represent relicts of

a more widely distributed species (vicariance) in the past is unlikely.

Some Rafflesia species, (e.g., R. arnoldii in Peninsular Malaysia,

Borneo, & Indonesia and R. speciosa in Panay & Negros Islands) do defy

transoceanic barriers. However, these islands were historically connected

during the Pleistocene when the sea level was much lower than today and,

in the case of Panay and Negros, represent land bridge islands that, together,

form a single Pleistocene aggregate island complex sharing many endemic

fauna (Wikramanayake ef al., 2002).

Thus, both R. tengku-adlinii and R. aurantia are likely examples of

morphologically semi-cryptic species, and possibly, an example of extreme

convergence in distantly related lineages. Studies on the accelerated rates of

floral evolution using molecular markers in 15 species (out of the 26 currently

recognized species) of Rafflesia by Barkman ef al. (2008) reveals that

genetic affinity is closer among those species found in the same geographic

region than those that are morphologically similar but found in different

geographic regions. Likewise, preliminary maximum likelihood analyses

of both nuclear and mitochondrial markers of five species of Philippine

Rafflesia (R. manillana, R. leonardi, R. schadenbergiana, R. speciosa, and R.

lobata) and Rafflesia species from Peninsular Malaysia and Borneo have

recovered the Philippine species as a monophyletic clade sister to all the

other species of Rafflesia (B. van Ee, pers. comm.).

Biodiversity in Luzon’s Sierra Madre Mountain Range

The Sierra Madre Mountain Range constitutes the largest block of lowland

forest remaining anywhere in the Philippines. It also boasts one of the coun-

try’s richest bird fauna, including some of the most rare and threatened,

such as the Philippine Eagle (Pithecophaga jeffery) and the Isabella Oriole

(Oriolus isabellae) (Danielsen et al., 1992; Mallari et al., 2002). Mammal di-

versity is also remarkably high and recent surveys strongly suggest that the

Sierra Madre is a unique center of mammal endemism (Danielsen ef al.,

24 Gard. Bull. Singapore 61 (1) 2009

1992; Rickart et al., 1998; Heaney, pers. comm.). Herpetological diversity in

the Sierra Madre appears similarly high, and distinct from that of the Cor-

dillera Central, southern Luzon, or the Bicol Peninsula (Brown et al., 2000;

Diesmos et al., 2005; Diesmos and Brown, unpublished data).

Many Philippine endemic plants are also found to occur in the Sierra

Madre mountain range, most notable of which is Podosorus angustatus

Holttum, a monotypic endemic fern genus known only from the type. At

the 16-ha Palanan Forest Dynamics Plot, one of the large-scale forest plots

coordinated by the Smithsonian Center for Tropical Forest Science (CTFS)

and Arnold Arboretum of Harvard University, 142 out of 323 tree species

recorded are Philippine endemics (Co et al., 2006), with 5 species thought to

be possibly new to science.

The discovery of Rafflesia in the Sierra Madre Mountain Range

and on other similar lowland habitats in the Philippines highlights the

importance of this fast disappearing forest type in the conservation of the

country’s threatened biodiversity (Tan et al., 1986).

Acknowledgements

We thank the indigenous Bugkalot community of Barangay Matmad; the

Municipal Government of Nagtipunan, Mayor R.K. Camma and Municipal

Councilor Y. Ebenga for hosting us; the regional, provincial, and municipal

offices of the Department of Environment and Natural Resources (DENR)

in Quirino under Foresters W. Malvar and H. Toribio; Forester J. Corpuz

joined us at the study site; Ms. E. Pasion (Community Organizer for

Quirino Province of the Sierra Madre Biodiversity Corridor Program, CI-

Philippines) helped in the preparatory logistics especially for securing the

collecting permit.

We thank M. Roy (‘Aloy’) M. Duya, L. Valenzuela-Duya

(Conservation International- Philippines), P. Alviola (UPLB-MNH),

J. Sarmiento (Laksambuhay Conservation, Inc.), W. Reyes (Wildlife

Conservation Society of the Philippines), and A. Benedict Oli (University of

the Philippines at Los Banos). Their comradeship and company in the field

is indispensable. To them, we dedicate the discovery of this new species. We

also thank Mr. N. Diego Jr. (PNH) for the line drawing illustration, Mr. M.G.

Price who provided the Latin diagnosis, and Drs. P.B. Pelser, L.R. Heaney,

and R.M. Brown for reviewing earlier drafts of the manuscript.

Financial support was provided by the Critical Ecosystem Partnership

Fund (CEPF), and the United States Agency for International Development

(USAID) for the series of biological surveys conducted by CI-Philippines

with partners in Quirino Province as well as other parts of the Sierra Madre

Biodiversity Conservation Corridor.

Rafflesia aurantia (Rafflesiaceae) Ds

References

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Gardens’ Bulletin Singapore 61 (1): 29-44. 2009 29

Seven New Species of Begonia from Sumatra

M. HUGHES , DEDEN GIRMANSYAH

WISNU HANDOYO ARDI AND NURAINAS—

‘Author for correspondence:

Singapore Botanic Gardens (Herbarium), 1 Cluny Road, Singapore 259569

Current address: Royal Botanic Gardens Edinburgh, 20a Inverleith Row,

Edinburgh EH3 5LR, U.K. E-mail: m.hughes@rbge.ac.uk

- Herbarium Bogoriense, Cibinong Science Center, Java, Indonesia.

* Bogor Botanic Garden, Jl. Ir. H. Juanda No. 13, P.O.Box 309, Java, Indonesia.

* Universitas Andalas Herbarium, Kampus Limau Manis, Padang 25163, Sumatra,

Indonesia.

Abstract

Seven new Begonia species are described from northern and western Sumatra:

Begonia gracilicyma Irmsch. ex M.Hughes (unplaced to section), Begonia

laruei M.Hughes (sect. Petermannia), Begonia multijugata M.Hughes (sect.

Petermannia), Begonia pasamanensis M.Hughes (sect. Reichenheimea),

Begonia puspitae Ardi (sect. Reichenheimea), Begonia tuberculosa

Girmansyah (sect. Platycentrum) and Begonia verecunda M.Hughes (sect.

Bracteibegonia). The conservation status of each species 1s assessed.

Introduction

There are currently 45 species of Begonia known from Sumatra (Hughes,

2008a). This is certainly a vast underestimate of the true number, largely as

a result of a lack of recent work on the genus and the propensity Begonia

has for generating narrow endemics (Hughes, 2008b), but also due to the

relatively low collection density over much of the island (Laumonier, 1997).

However the extensive collections by de Wilde and de Wilde-Duyfies have

provided much insight into Begonia diversity in the Gunong Leuser environs.

Given the relentless pressure on forest habitats throughout Southeast Asia

(Sodhi, 2004), there is an urgent need to provide taxonomic data as a basis

for conservation efforts.

During recent expeditions to Sumatra, seven new species of

Begonia have been discovered and are described here. They belong to

sect. Petermannia (2 species); Reichenheimea (2 species); Bracteibegonia (1

species) and Platycentrum (1 species); there is a further rather enigmatic

species which is unplaced to section. In common with most other Begonia

from Sumatra, these new taxa are found in the mountainous spine of the

30 Gard. Bull. Singapore 61 (1) 2009

island along the Barisan Range, and also from the mountains in North

Sumatra and Aceh (Fig 1). Herbarium material indicates that there are

still a considerable number of undescribed taxa in sections Bracteibegonia

and Reichenheimea from these areas, which may represent largely endemic

radiations. The Tigapulu Mountains are also likely to harbour further

endemic taxa, although the areas which remain most under-explored are the

mountains and limestone karst regions in northern Aceh, from which hardly

any material has been seen. All specimens cited are available as digital

images via Hughes & Pullan (2007); Doorenbos ef al. (1998) was consulted

to assist with sectional placement.

Begonia gracilicyma Irmsch. ex M.Hughes, sp. nov. (not placed to section)

B. divaricatae maxime similis, sed foliis plus elongatis et minus dentatis,

floribus parum rubescentibus non albis neque roseis differt. — Typus:

Sumatra, Padang, Ajer Mantjoer, O. Beccari PS610 (holo, FI; iso, B, FI, K,

L). Fig. 2B.

Plant erect, branching herb, 40 to 70 cm high. Stem woody, especially at base,

glabrous, ca 5 mm wide, internodes 5-10 cm apart. Stipules lanceolate, 10-

12 x 3 mm, glabrous, with a filiform extension at the tip, deciduous. Leaves

alternate; petiole 1.5-5 cm long, glabrous; lamina elongate-lanceolate,

strongly asymmetric, basifixed, cordate at base, lobes not overlapping, one

lobe much larger giving an angular appearance, 10-18 x 2.5-5.5 cm, midrib

8-13 cm long, venation palmate-pinnate, upper surface matt green, glabrous;

underside pale green sometimes marked with red, glabrous; margin glabrous,

denticulate; apex acuminate. Inflorescence appearing adnate on the petioles,

protandrous, bisexual; bracts 1-3 mm long, margin entire, deciduous. Male

flowers: pedicel 5 mm, glabrous; tepals 4, outer tepals orbicular, white with

a reddish tinge on the reverse, glabrous, ca 4 mm in diameter, margin entire;

inner tepals oblong-obovate, white, 4 x 2 mm; androecium yellow, symmetric;

stamens ca 30; filaments slightly fused at base, 0.75 mm long; anther about

as long as the filament, dehiscing through slits about half the length of the

anther, hooded, connective not extended. Female flowers: pedicel 10 mm

long; ovary 3 locular, with three equal wings, placentae entire; tepals 5,

pale green, 5 mm long, margin entire; stigma with three styles joined at the

base, U-shaped, persistent. Fruit brown, dehiscent, pendulous on a hair-like

pedicel when dry, rounded at base, truncate to retuse at the apex; wings

extending along the pedicel, equal, 14 x 5 mm; capsule shape oval, length 8-9

mm, width 5 mm. Seeds unknown.

Other specimens examined: Sumatera Barat: Mt. Tandikat, 23 vii 1955,

W. Meijer 3812 (L); ibid., 1955, W. Meijer 391? (BM); Talaman, H.A.B.

Seven New Species of Begonia from Sumatra 3

Bunnemeijer 372 (B, BO); Anai Nature Reserve, 9 xi 1991, Anda collectors

25 (ANDA); ibid., 9 xi 1991, Anda collectors 37 (ANDA)); ibid., 23 xii 1983,

Niniek & Wardi 458 (BO); ibid., 21 11 1990, Anda collectors 90 (ANDA);

Anak Air Ambacang Badak, 15 viii 1995, H. Okada 2004 (ANDA); Bukit

Tambun Tulang, 8 xi 1998, Anda collectors s.n. (ANDA); ibid., 29 x 1988 -

31 x 1988, Anda collectors sn. (ANDA)); ibid., 7 xi 1998, Anda collectors 15

(ANDA),; ibid., v 2006, Anda collectors 17 (ANDA)); ibid., 8 xi 1998, Anda

collectors 18 (ANDA\)): ibid., 8 xi 1998, Anda collectors 23 (ANDA); ibid..,

28 ii 1987, Anda collectors 24 (ANDA)); tbid., 8 xi 1998, Anda collectors

42 (ANDA)); ibid., 26 v 1991, Anda collectors 45 (ANDA)); ibid., 10 xi 1991,

Anda collectors 47 (ANDA); Bungus—Cindakir, 25 v 2002, Anda collectors

35 (ANDA); Desa Sipisang, 19 xii 1992, Anda collectors 21 (ANDA)); ibid.,

6 iv 1997, Anda collectors 23B (ANDA)); ibid., 19 xii 1992, Anda collectors

25 (ANDA); ibid., 19 xii 1992, Syofyan et al. 31 (ANDA); ibid., Anda

collectors 33 (ANDA),; ibid., 5 iv 1997, Anda collectors 35 (ANDA); ibid., 5

iv 1997, Anda collectors 41 (ANDA)); ibid., 17 viii 1995, Anda collectors 503

(ANDA); Gunung Gadut, 15 xii 1987, H. Okada 4629 (ANDA); Gunung

Gadut, Bt. Batu Bajolang, 12 1 1983, M. Hotta, et al. 1326 (A, L); ibid., 121

1983, M. Hotta, et al. 1320 (ANDA); Gunung Gadut, Bukit Gambir, 15 xii

1987, H. Okada 4625 (ANDA); Kandang Ampat, Kabupaten Padang Pisang,

27 xi 1994, Anda collectors 10 (ANDA)); ibid., 27 xi 1994, Anda collectors 5

(ANDA); Muko-muko, 5 x 1986, Witnarti 24 (ANDA [3]); Padang Pariaman,

30 iv 2004, D. Girmansyah 380 (BO); Road to Rimbo Panti, 27 v 2007, M.

Hughes & D. Girmansyah MH1407 (BO, E); Taman Hutan Raya, Ladang

Padi, 16 v 1993, Anda collectors 112 (ANDA): ibid., 18 xii 2004-19 xii 2004,

Anda collectors 112? (ANDA)); ibid.,22 v 2007, M. Hughes & D. Girmansyah

MH1403 (BO, E [3]); ibid., 24 v 2003, Anda collectors 23 (ANDA)); ibid.,

24 v 2003, Anda collectors 23 (ANDA); ibid., 4 v 2002, Anda collectors 27

(ANDA)); ibid., 24 v 2003, Anda collectors 27? (ANDA); ibid., 14 vi 1998,

Anda collectors 29 (ANDA)); ibid., D. Girmansyah, et. al. 3 (BO, E); ibid., 16

v 1993, Anda collectors 38 (ANDA); ibid.,4 v 1998-5 v 1998, Anda collectors

46 (ANDA),; ibid., 5 v 2002, Anda collectors 72 (ANDA); Tambun Tulang,

10 xi 1991, Anda collectors sin. (ANDA)); ibid., 29 x 1983, Eliwiratma 09

(ANDA)); ibid., 10 xi 1991, Anda collectors 43 (ANDA)); tbid., 10 xi 1991,

Anda collectors 51 (ANDA); ibid., 10 xi 1991, Anda collectors 52 (ANDA).

Distribution and ecology: Sumatera Barat: Agam, Padang Pariaman, Tanah

Datar, Padang. At altitudes of 150 to 750 m (Fig 1). Terrestrial forest floor

herb found in steep lower and mid-montane forest.

IUCN category: LC. Begonia gracilicyma is found at a number of sites within

the Gunung Singgalang Protection Forest and the Kerinci Seblat National

B. gracilicyma

B. laruei

| B. pasamanensis

“ B. verecunda

@ 8B multijugata

B. puspitae

B. tuberculosa

0 100200 400 Kilometers

Gard. Bull. Singapore 61 (1) 2009

Figure 1. Distribution of B. gracilicyma, B. laruei (top); B. pasamanensis, B. verecunda

(middle); B. multijugata, B. puspitae, B. tuberculosa (bottom).

ies)

io)

Seven New Species of Begonia from Sumatra

Park.

Notes: Acording to unpublished manuscripts in the Berlin herbarium

archives, Irmscher first coined the name Begonia biinnemeijerii for this

species, based on a collection by H.A.B. Biinnemeijer from “Talaman”.

However, this epithet was later crossed out on the manuscript and specimens,

and replaced with B. gracilicyma, referring to the hair-like nature of the

peduncles and pedicels when dry. For unknown reasons, the taxon was left

out of a paper by Irmscher (1953) which included several other new species

from Sumatra, hence the name is validated here. The species is distinct from

all other species currently described from Sumatra in its elongate leaf shape

with a very large basal lobe which gives the lamina a distinctive outline,

especially when pressed (Fig. 2). The leaves are tissue-thin and translucent

when dry. Its highly branched growth form is suggestive of an affinity with

B. divaricata Irmsch., also from the Padang region, which grows at much

higher altitudes (around 1500-1700 m). The combination of caulescent

habit, protandrous inflorescences and entire placentae makes B. gracilicyma

impossible to place comfortably in any of the existing Asian sections of the

genus. Irmscher annotated his manuscript with the unpublished section

Merrillia, although we refrain from validating this name here; further study,

including molecular work, is required before allocating B. gracilicyma to a

section. “Anda collectors” refers to the large number of student collectors

from the University of Andalas in Padang.

Begonia laruei M.Hughes, sp. nov. Sect. Petermannia

Ab omnibus speciebus Begoniae sectionis Petermanniae insulae Sumatrae

foltis lobatis et habitu majore distincta. — Typus: Sumatra, North Sumatra

Prov., Gunung Sibayak, 12 v 2007, 3° 13' 38"' N 98° 31' 2"' E M. Hughes & D.

Girmansyah MH1389 (holo, E: iso, BO, ANDA). Fig. 2A.

Plant erect, 50-100 cm high. Stem woody, especially at the base, glabrous,

ca. 8 mm wide, internodes 4-10 cm apart. Stipules lanceolate, ca. 8 mm

long, caducous. Leaves alternate: petiole 2.5—8 cm long, glabrous; lamina

lanceolate, strongly asymmetric, usually with 2+ pointed lobes extending

to around 1-5 cm, basifixed, cordate at base, lobes not overlapping, 11—25 x

3.5—10 cm, midrib 9-15 cm long, venation palmate: upper surface dark green

with silver markings between the veins or uniform green, glabrous; underside

pale green, glabrous; margin glabrous, with small teeth between the lobes,

apex acuminate. Inflorescence protogynous, bisexual, terminal, cymose,

branching up to 6 times; bracts 4 x 5 mm, entire or slightly retuse, deciduous.

Male flowers: pedicel 4 mm, glabrous, tepals 4; outer tepals reddish or dirty

pink, more so towards the base, sometimes white, orbicular, truncate at base,

34 -

J Gard. Bull. Singapore 61 (1) 2009

Figure 2. A. Begonia laruei (main picture and flowers, Ketambe Research Station environs;

variegated leaf, inset, Gunung Sibayak; female flowers, top left, male, top right); B. Begonia

gracilicyma (Padang, Ladang Padi; ripe fruit, top right; male flower, botom right; female

flower after fruit set, bottom left); C. Begonia puspitae (cultivated in Bali Botanic Garden

(main picture) and Bogor Botanic Garden (inset flowers); male flowers, left, front and side

view; female, right); D. Begonia verecunda (Ketambe Research Station environs; male flower,

top left; female flower, bottom right; ripe fruit, bottom left).

ca 5 x 5 mm, margin entire; inner tepals ca 4 x 2 mm, paler than the outer;

androecium symmetric, yellow; stamens ca 30; filaments short, on a column;

anthers hooded, ca. | mm long, lower ones sub-sessile, upper ones on a short

filament. Female flowers: pedicel ca 4 mm long; ovary 3 locular, with three

Seven New Species of Begonia from Sumatra 35

equal wings, placentae bifid; tepals 5, obovate, ca 8 x 4 mm, margin entire;

stigma with three styles, deciduous. Fruit truncate to slightly retuse at base,

pale brown, dehiscent, pendulous, usually borne in pairs; wings equal, bases

not extending along the pedicel, rounded at the tips, 14 x 6 mm; capsule

shape broadly oval, 12 x 9 mm. Seeds barrel shaped, 0.3 mm long.

Other specimens examined: Aceh. Gunung Ketambe, 16 v 1972, de Wilde &

de Wilde-Duyfjes 12006 (BO, L [2]): ibid., 19 vii 1972, de Wilde & de Wilde-

Duyfjes 13814 (BO, L); Gunung Leuser Nature Reserve, Gunung Mamas,

7 ii 1975, de Wilde & de Wilde-Duyfjes 14632 (L); Gunung Leuser Nature

Reserve, Ketambe Research Station, 28 vii 1979, de Wilde & de Wilde-Duyfjes

19206 (BO); Lau Alas, 6 vi 1972, de Wilde & de Wilde-Duyfjes 12627 (BO, L);

Mamas River, 27 vi 1979, de Wilde & de Wilde-Duyfjes 19164 (BO [2], L);

Blang Kedjeren, 15 11 1937, C.G.G.J/.v. Steenis 337 (BO); Bur ui Papandji, 23

vi 1930, Frey-Wyssling 45 (BO); Gajolanden, 21 111 1937, CG.G.J.v. Steenis

9914 (BO, L). Sumatera Utara. Brastagi, 3-17 iv 1925, H.S. Yates 1400 (BO);

Dolok Singgalang, 25 v 1922, J.A. Lorzing 8863 (BO); Gunung Sibayak, 7 xii

1988, PJ_A. Kessler 105 (B, L [2]); ibid., 5 x 1928, A. Lorzing 14038 (BO);

Gunung Sinabung, 14 v 2007, M. Hughes & D. Girmansyah MH1398 (BO,

E); Lae Banalsal, 17 xi 1941, H. Surbeck 554 (L); Sarinembah, 28 vi 1918,

H.H. Bartlett & C.D. La Rue 200 (A, L); Karoland, Petjeren, 22 vi 1928, C.

Hamel & Rahmat Si Toroes 782 (A); Bandar Baru, Sungai Tepi, 20 v 1981, W.

Meijer 15803 (BO, L).

Distribution and ecology: Aceh: Gayo Lues, Aceh Tengara. Sumatera Utara:

Langkat, Karo, Simalungen. At altitudes of 400-1400 m. Terrestrial forest

floor herb found in lower and mid-montane forest.

IUCN category: LC. The bulk of the distribution of Begonia laruei lies within

the Gunung Leuser National Park, where it is known from a number of

different localities.

Notes: Irmscher annotated a specimen (BO) of this taxon with the

unpublished name B. bartlettii. However, as there is already a B. bartlettiana

it has been decided to commemorate the second collector Carl Downey

La Rue, who made the first herbarium collection of this species with H.H.

Bartlett whilst working as a botanist for the U.S. Rubber Co. in Asahan. In

common with many species in sect. Petermannia, B. laruei is usually found

as single plants scattered throughout the forest, though small colonies

can occasionally be found. On the herbarium sheet its leaves are instantly

recogniseable, being lobed to a greater or lesser extent (Fig. 2). The fruit

shape is also quite distinctive, being wider than long and fairly woody, and

36 Gard. Bull. Singapore 61 (1) 2009

often slightly retuse at both ends. There is variation in the leaf variegation,

with some populations (e.g., Gunung Sibabyak) having leaves which are

heavily blotched with silver on a dark green background with a thin silver

margin, whereas others have paler leaves with no markings (e.g., Gunung

Sinabung vicinity) or red veins on the upper surface (Ketambe environs).

Begonia multijugata M. Hughes sp. nov. Sect. Petermannia

A B. atricha fructibus minoribus et multijugatis diagnoscenda. — Typus:

Sumatra, Aceh, Gunung Leuser Nature Reserve, Air Panas, 18 111 2008, 3°

41' 44" N 97° 39' 31" E P. Wilkie, M. Hughes, A. Sumadijaya, S. Rasnovi,

Marlan & Suhardi PW768 (holo, BO; iso, E, SING). Fig. 3B.

Erect glabrous herb to 50 cm high. Stem slightly woody when dry, glabrous,

5-8 mm wide, internodes 5-20 cm apart, red. Stipules lanceolate, ca 20 x 6

mm, with minute glandular hairs on the reverse, with a very small exten-

sion at the tip, deciduous. Leaves petiole 1.5-6 cm long, glabrous; lamina

ovate-lanceolate, asymmetric, basifixed, cordate at base, lobes not overlap-

ping, length 12-22 cm, width 5-9 cm, midrib 10-17 cm long, venation pal-

mate-pinnate; upper surface dark green with white spots between the veins,

glabrous, underside wine-red, glabrous; margin glabrous, toothed at the end

of the veins with smaller teeth between, apex acuminate. Inflorescence very

compressed, protogynous, bisexual; bracts translucent white. Male flowers:

pedicel length 10 mm, glabrous; tepals 2, white, glabrous, 6 mm long, obo-

vate, margin entire; androecium pale yellow, symmetric, almost conical; sta-

mens 30, filaments shorter than the anther at the base, becoming slightly

longer toward the apex, anther 0.75 mm long, obovate, hooded, dehiscing

through short slits less than half the length of the anther, connective retuse.

Female flowers: pedicel up to about | cm long; ovary 3 locular, with three

equal wings, wings rounded-triangular, 2-3 mm wide, placentae bifid; tepals

5, white, obovate-orbicular, 6 mm long, margin entire; stigma yellow, with

three styles, once spirally twisted, deciduous. Fruit pale brown, dehiscent on

all 3 faces, borne in a cluster of up to 5 pairs, each pair ca 5 mm apart; wings

narrow, capsule broadly elliptic, 7-9 x 6-7 mm. Seeds rounded-barrel shaped,

0.25 mm long.

Other specimens examined: Aceh. Gajolanden, 25 11 1937, C.G.G.J/.v. Steenis

9291 (BO, L); Lau Alas, 2 vi 1972, de Wilde & de Wilde-Duyfjes 12537 (L [2]);

Kloet Nature Reserve, 10 vii 1985, de Wilde & de Wilde-Duyfjes 19910 (BO, L).

Distribution and ecology: Aceh: Gayo Lues, Aceh Tenggara, Aceh Seletan.

At altitudes of less than 400 m (Fig. 1). Terrestrial herb found in lowland

forest.

Seven New Species of Begonia from Sumatra 37

IUCN category: LC. As the entire known range of this species is within the

Gunung Leuser National Park, it should be considered as Least Concern as

long as the Park remains intact.

Notes: Vegetatively this glabrous species closely resembles B. atricha, both

in leaf shape and colouring (dark green with evenly spaced white spots

between the veins). However the infructescence differs considerably, being

a congested cluster of small, paired fruits which make the species instantly

recogniseable when fertile (Fig. 3): B. atricha has large bell-shaped fruits

borne singly on long thin pedicels. B. multijugata is usually found as solitary

plants on the forest floor. The eipthet refers to the paired fruits, being derived

from jugatus, meaning yolked together in pairs.

Begonia pasamanensis M.Hughes, sp. nov. Sect. Reichenheimea

A B. stictopoda foliis plus late ovatis et adpressis, pedicellis pilosis differt. —

Typus: Sumatra, West Sumatra, Road to Padang, 29 v 2007, 0° 2' 32'' N 100°

13' 5" E M. Hughes & D. Girmansyah MH1419 (holo, E: iso, BO, ANDA)

Fig. 3A.

Repent herb, ca 20 cm tall. Stem rhizomatous, 3 mm wide, internodes 0.3-

1 cm apart. Stipules persistent, triangular, ca 7 x 3 mm, glabrous, with a

filiform extension at the tip, semi persistent. Leaves alternate: petioles

densely hairy, 3-20 cm long, unequal on the same plant, being much longer on

older leaves; lamina broadly ovate, basifixed, asymmetric, base cordate with

lobes overlapping in vivo, length 4-9 cm, width 3.5-8 cm, midrib 3-6 cm long,

venation palmate, upper surface mid green to blackish green, reddish when

young, usually paler along the veins, glabrous, smoothly bullate between

the veins, underside wine-red, hairy on veins only, margin with occasional

short hairs, entire to denticulate, apex obtuse. Inflorescence cymose, axillary,

protogynous, bisexual; bracts sub-orbicular, 1.5-3 mm long, 1.5-3 mm wide,

margin fimbriate, deciduous. Male flowers: pedicel length 6 mm, hairy, tepals

4; outer tepals sub-orbicular, white or pale pink, with scattered colourless

hairs, truncate at base, margin entire; inner tepals oblong-obovate, white:

androecium yellow, symmetric; stamens ca. 50; filaments fused at base into a

short column, 1 mm long, subequal; anther shorter than the filament, length

0.75-1 mm long, oblong, dehiscing through slits longer than half the length

of the anther, slightly hooded, connective retuse. Female flowers: pedicel:

ovary with three equal wings, 3 locular, placentae entire; bracteoles absent:

tepals 3 or 4, white or pale pink with scattered colourless hairs; outer sub-

orbicular 8-10 mm in diameter, often deeper pink at the base externally:

inner tepals oblong-obovate, 7-10 mm long, ca 3 mm wide; stigma with three

styles, styles deep yellow, u-shaped, stigmatic surface spiral. Fruit truncate

38 Gard. Bull. Singapore 61 (1) 2009

at base, recurved and held horizontally at maturity; ca 8 x 14 mm in total,

capsule orbicular, ca 6 mm in diameter; wings rounded, equal to subequal.

Seeds barrel shaped, 0.25 mm long.

Other specimens examined: Sumatera Barat. Rimbo Panti National Park, 28

v 2007, M. Hughes & D. Girmansyah MH1411 (BO, E)); ibid., 28 v 2007, M.

Hughes & D. Girmansyah MH1411A (BO, E); ibid., 28 v 2007, M. Hughes

& D. Girmansyah MH1416 (BO, E); ibid., 28 v 2007, M. Hughes & D.

Girmansyah MH1417 (BO, E); ibid., 12 vi 1953, J.v. Borssum Waalkes 1730

(BO, L); Road to Padang, 29 v 2007, M. Hughes & D. Girmansyah MH1421

(BO, E); Road to Rimbo Panti, 27 v 2007, M. Hughes & D. Girmansyah

MH1410 (BO, EB).

Distribution and ecology: Sumatera Barat, endemic to Pasaman, at altitudes

of 200-500 m (Fig. 1). Lithophytic herb, on steep rocky banks with clay soil

or on rocks near streams.

IUCN category: LC. Although B. pasamanensis has quite a narrow

~ distribution, it is found within the Rimbo Panti Nature Reserve and the

Malampah Alahan Panjang Nature Reserve. It also has the potential to

colonise roadside banks where there is canopy cover and suitable substrate

(clay soil on limestone).

Notes: In life this species has leaves which are slightly raised between the

veins, with the veins on the upper leaf surface being a shade paler than the

lamina. It grows gregariously on steep banks along the main road through

the Rimbo Panti Nature Reserve (Fig. 3), where it co-occurs with a species

tentatively identified as B. stictopoda (Miq.) A.DC. Although superficially

similar, B. pasamanensis can be distinguished by its smaller, more broadly

ovate leaves which are usually much darker and have a blunter apex; those

of B. stictopoda are more elongate, especially on younger plants when their

size is more similar. B. pasamanensis also tends to have its petioles and

leaf laminas appressed against the substrate. As well as these vegetative

characters, B. pasamanensis has hairy pedicels, as opposed to glabrous in

B. stictopoda. Several plants along the roadside through Rimbo Panti were

intermediate between the two species, and it is possible that this artificial

environment has brought the species together where they would otherwise

not naturally co-occur. However, large hybrid swarms were not observed.

Observations in the field show that B. pasamanensis and some other

Sumatran species in sect. Reichenheimea have fruits which are recurved at

maturity, with one wing pointing downwards and the other two forming a

small splash-cup.

Seven New Species of Begonia from Sumatra 39

Figure 3. A. Begonia pasamanensis (Rimbo Panti Nature Reserve; female flowers, inset); B.

Begonia multijugata (Gunung Leuser Nature Reserve, Air Panas; male flower, top left; female

flower, bottom left: ripe fruits, bottom right); C. Begonia tuberculosa (Gunung Sinabung:

male flower, top right; female flower, bottom right).

Begonia puspitae Ardi, sp. nov. Sect. Reichenheimea

Ab omnibus speciebus Begoniae sectionis Reichenheimeae insulae Sumatrae

foliis supra pilosis et stipulis gaudentibus pilis plumosis differt. — Typus:

Sumatra, West Sumatra, Gunung Silungkang, Suaka Alam Batang Pangean,

28 vili 2005, D.M. Puspitaningtyas DM1742 (holo, BO). Fig. 2C.

Repent herb ca 20 tall. Stem rhizomatous, glabrous, internodes ca 5 mm

apart. Stipules persistent, broadly triangular ca 13 x 7 mm, covered with

branched hairs 0.5-1.5 mm long. Leaves, petiole 7-22 cm long, green, with

dense white to pink hairs up to 3 mm long; blade very asymmetric 8-22 x

5—13.5 cm, reddish green or green between the veins above and pale green

below, broadly ovate, base cordate, margin hairy, denticulate, tip acuminate,

venation palmate, pilose on both surfaces, hairs white, 1-2 mm long.

Inflorescence axillary, cymose, covered with white dense hairs 0.5-1 mm,

primary peduncle longer than the leaves, ca 23-28 cm long, protandrous;

bracts ovate, ca 3 x 5 mm, white at base and pale green at tip, outer surface

covered with dense pink hairs, ca 1 mm long, margin serrate and hairy, tip

acuminate ending with a hair, falling before the male flowers open. Male

40 Gard. Bull. Singapore 61 (1) 2009

Flowers: pedicel 12-17 mm; tepals 2, white, sub-orbicular 7-8 x 8-9 mm,

margin entire, tip rounded, with reddish stiff hairs 0.5-1 mm long externally;

androecium yellow, symmetric, column ca 0.5 mm; stamens ca. 35; filaments

ca | mm long; anthers yellow, elliptic, 0.8 mm long, tip truncate, opening by

slits as long as the anther sac. Female flowers: pedicel 15-20 mm, greenish

red; ovary pale pink to white, ca 4 mm long, wings 3, unequal, locules 3,

placentation axile, placentae entire; tepals 2, white, glabrous, sub-orbicular

7-9 x 8-9 mm, margin entire, tip rounded; styles 3, styles and stigma yellow,

ca 3.5 mm. Fruits with pedicel 2 cm long; dehiscent, capsule oval ca 7 mm

long, hairless, locules 3, wings 3, unequal, splitting between locules and wings.

Seeds barrel shaped, ca 0.5 mm long, collar cells ca 1/5 length of the seed.

Distribution and ecology: Sumatera Barat, only known from the type locality,

Gunung Silungkang, between 600-700 m altitude (Fig 1).

IUCN category: DD. There is not enough information on the current

condition and protection status of the type locality.

Notes: B. puspitae is unusual in having branched hairs covering the

stipules, and thus, differs from the other known Sumatran species in sect.

Reichenheimea. The presence of a fairly dense indumentum of short hairs

on the upper leaf surface also marks it as distinct from other Sumatran

species in this section (Fig. 2). The epithet is after the collector, Dwi Murti

Puspitaningtyas.

Begonia tuberculosa Girmansyah, sp. nov. Sect. Platycentrum

A B. areolata petiolis albopilosis, foliis mollibus et variegatis, radicibus

tuberculosis differt. —-Typus: Sumatra, North Sumatra, Gunung Sinabung, 3°

10' 33"' N 98° 23' 31'' E M. Hughes & D. Girmansyah MH1394 (holo, E; iso,

E [2], BO, ANDA). Fig. 3C

Repent herb ca 45 cm high, with small ca 5 mm diameter tubercles on the

roots. Stem rooting at the nodes, hairy, more so towards internodes, ca 6

mm wide, internodes up to 20 cm apart though usually shorter. Stipules

lanceolate, 10 mm long, 5 mm wide, hairy, with a filiform extension at the

tip, persistent. Leaves alternate but appearing opposite subtending the

inflorescence; petioles around 10 cm long, with dense grey-white hairs

becoming very light brown on drying; lamina ovate with an extended tip,

basifixed, cordate at base lobes not overlapping, ca 13 x 8 cm, midrib ca 9

cm long, venation palmate, asymmetric, upper surface mid green with dark

purple or sometimes dark green patterning along the veins, upper surface

hairy all over, underside with red hairs denser on the veins, fading with age,

Seven New Species of Begonia from Sumatra 4]

margin fimbriate—denticulate. Inflorescence cymose, terminal, bisexual;

bracts ca 5 mm long, margin fimbriate, caducous. Male flowers: pedicel 15

mm, hairy; tepals 4, outer tepals sub orbicular, white with pale pink on the

reverse, hairy, hairs denser towards the base, truncate to rounded at base,

ca. 10 x 10 mm, margin entire, inner tepals oblong-elliptic, white, ca 15 x 8

mm; androecium yellow, symmetric; stamens ca. 80-100; filaments, 1.5-2 mm

long, equal, fused at the base into a short column; anther shorter than the

filament, ca 1.5 mm long, oblong elliptic, dehiscing through slits longer than

half the length of the anther, connective extended. Female flowers: pedicel

ca 2 cm long; ovary 3 winged, one wing much enlarged, covered in quite

stout pinkish hairs, 2 locular; tepals 5, white, ovate ca 10 x 8 mm, with pinkish

hairs on the reverse; stigmas 2, pale yellow-green or yellow, stigmatic surface

convulted. Fruit not seen mature. Seeds unknown.

Other specimen examined: Aceh. Mamas River, 11 vii 1985, de Wilde & de

Wilde-Duyfjes 19075 (L).

Distribution and ecology: Aceh: Aceh Tenggara. Sumatera Utara: Karo. At

altitudes of 1200-1400 m (Fig 1). Terrestrial herb of mid-upper montane

forest, found in rich humus and often at the base of tree trunks.

IUCN category: VUD2. Begonia tuberculosa is currently known from only

2 sites, one of which (Gunung Sinabung) is being encroached despite being

in the Leuser Ecosystem Conservation Area. The other site in the Gunung

Leuser National Park is considered safe. It is recommended that (1) protection

of the forest on Gunung Sinabung, especially at the lower elevations, be

made a priority and (ii) the species be brought into cultivation.

Notes: B. tuberculosa is instantly recognisable as a member of sect.

Platycentrum, with its two-locular fruit and large androecium on a short

column. Its closest affinity is probably with B. areolata Miq., a widespread

species at 1000—1800m in Sumatra and Java, as it shares the unusual character

of having a pair of opposite leaves subtending the inflorescence. As well

as having tubercles on the roots, B. tuberculosa differs from B. areolata

in having variegated leaves (Fig. 3) and grey-white (not red) hairs on the

petiole, which contrast with the claret-red hairs on the underside of the

lamina. The lamina margin is also less lobed than B. areolata, and feels very

different to the touch, being soft and velvety rather than bristly and bumpy.

Sect. Platycentrum is especially species rich in Peninsular Malaysia (Kiew,

2005), and B. tuberculosa and other species in the section from Sumatra and

Java probably represent a link with that flora, especially species with bullate

leaves such as B. wyepingiana and B. vallicola.

42 Gard. Bull. Singapore 61 (1) 2009

Begonia verecunda M.Hughes sp. nov. Sect. Bracteibegonia

Differt a B. bracteata stipulis angustis lanceolatis acuminatis (haud late

ovatis); a B. lepidella fructibus recurvatis. — Typus: Sumatra, Aceh, Gunung

Leuser Nature Reserve, Ketambe Research Station, 7 111 2008, 3° 40' 46"'

N 97° 38' 37" E P. Wilkie, M. Hughes, A. Sumadijaya, S. Rasnovi, Marlan &

Rabusin PW623 (holo, BO; iso, E, SING). Fig. 2D.

Low growing herb, 20 cm high. Stem slightly woody, densely hairy, ca 3 mm

wide, internodes 1-2 cm apart. Stipules narrowly lanceolate, 10 mm long,

with a filiform extension at the tip, laxly deciduous, appearing very narrow

and almost filiform when dry. Leaf petiole around 1| cm long, densely hairy;

lamina ovate-lanceolate, asymmetric, basifixed, cuneate to rounded on one

side, lobed on the other, the short basal lobe crossing the petiole slightly,

total length 6.5-10 x 3-4 cm, venation palmate-pinnate, upper surface dark

green, usually glabrous but sometimes with one or two short fleshy hairs,

underside deep red or pale green, with short fairly robust hairs on the veins,

appressed in the direction of the apex, becoming sparser on the smaller

veins, margin entire to minutely denticulate, apex acute. Inflorescence a

short and compact cyme, protogynous, bisexual; bracts persistent at the

base. Male flowers: pedicel length 5-10 mm, glabrous; tepals 4; outer tepals

sub-orbicular, glabrous, 9 mm long, white tinged with deep pink, colouring

zygomorphic, with top of flower being darker; margin entire; inner tepals

white, ca 6 x 3 mm; androecium yellow, symmetric; stamens 30; filaments

slightly fused at base, unequal, shorter or longer than the anther; anther

about | mm long, oblong obovate, dehiscing through slits less than half the

length of the anther, hooded, connective slightly retuse. Female flowers:

pedicel ca 6 mm long; tepals 5, white tinged with pink, much deeper on the

reverse, colouring zygomorphic, darker towards the top, elliptic-ovate, 7 mm

long, margin entire; persistent and closed during fruit maturation, eventually

deciduous; ovary green with the adaxial wing tinged deep pink, 3 locular,

placentae bifid; stigma yellow, deciduous. Fruit dehiscent, recurved at

maturity; wings subequal, 3-4 mm wide, the adaxial wing the larger; capsule

shape narrowly elliptic, slightly curved, length 10 mm, width 4 mm. Seeds

barrel shaped, 0.25 mm long.

Other specimens examined: Aceh. Gunung Leuser Nature Reserve, 16 v 1972,

de Wilde & de Wilde-Duyfjes 12010 (K, L); Gunung Leuser Nature Reserve,

Air Panas, 18 111 2008, P. Wilkie, M. Hughes, A. Sumadijaya, S. Rasnovi, Marlan

& Suhardi PW779 (BO, E, SING); Gunung Leuser Nature Reserve, Gunung

Mamas, 7 ii 1975, de Wilde & de Wilde-Duyfjes 14640 (L); Gunung Leuser

Nature Reserve, Ketambe Research Station, 7 111 2008, P. Wilkie, M. Hughes,

A. Sumadijaya, S. Rasnovi, Marlan & Rabusin PW617 (BO, E, SING); ibid.,

Seven New Species of Begonia from Sumatra 43

7 111 2008, P. Wilkie, M. Hughes, A. Sumadijaya, S. Rasnovi, Marlan & Rabusin

PW621a (SING); Lau Alas, 28 v 1972, de Wilde & de Wilde-Duyfjes 12370

(L); ibid., 4 11 1975, de Wilde & de Wilde-Duyfjes 14487 (L); ibid., 21 iti 1975,

de Wilde & de Wilde-Duyfjes 15683 (L).

Distribution and ecology: Aceh: Aceh Tenggara. At altitudes of around 200-

400 m (Fig 1). Terrestrial herb found scattered on the forest floor in lowland

forest.

IUCN category: LC. As the entire known range of this species is within the

Gunung Leuser National Park, it should be considered as Least Concern as

long as the Park remains intact.

Notes: The epithet is derived from verecundus meaning shy or demure,

referring to the way the female flowers close their tepals and bow away

from sight after pollination (Fig. 2). The short petioles and peduncles, dense

indumentum, bicoloured flowers and elongate fruit with quite narrow wings

are characteristic for sect. Bracteibegonia. Judging from other herbarium

collections, the variation in this section on Sumatra appears to be at once

both complex and subtle, and there are undoubtedly many other species

waiting to be described. Tepal number, colour and shape as well as fruit

position at maturity are important characters in this group.

Acknowledgements

The support of the M.L. MacIntyre Trust, the Sibbald Trust, the Royal

Botanic Garden Edinburgh Small Projects Fund, the Davis Expedition

Fund, the Percy Sladen Memorial Trust and SYNTHESYS (grants NL-TAF

1608, FR-TAF 1416 and DE-TAF 2181) is gratefully acknowledged. This

research would not have been possible without the support of the Indonesian

Ministry of Research and Technology (RISTEK), the Indonesian Institute

of Sciences (LIPI), Direktorat Jenderal Perlindungan Hutan dan Konservasi

Alam (DITJEN PHKA), Bandan Pengolola Kawasan Ekosystem Leuser

(BPKEL), Universitas Sumatera Utara and Universitas Syiah Kuala. A.

Sumadiaya and S. Rasnovi are thanked for their assistance in the field. The

curators of herbaria A, ANDA, B, BM, BO, E, K, L, MICH, P,SING and U are

thanked for facilitating access to specimens. This research was supported by

the National Parks Board, Singapore. Two anonymous referees are thanked

for their comments and improving the manuscript.

44 Gard. Bull. Singapore 61 (1) 2009

References

Doorenbos, J., M.S.M. Sosef and J.J.FE. de Wilde. 1998. The sections

of Begonia including descriptions, keys and species lists (Studies in

Begoniaceae VI). Agricultural University Wageningen Papers 98(2):

1-266.

Hughes, M. and M. Pullan. 2007. Southeast Asian Begonia Database.

Electronic publication accessible via: www.rbge.org.uk.

Hughes, M. 2008a. An annotated checklist of Southeast Asian Begonia. Royal

Botanic Garden Edinburgh, UK.

Hughes, M. 2008b. Population genetic divergence corresponds with species

level biodiversity patterns in the large genus Begonia. Molecular Ecology

17: 2643-2651.

Irmscher, E. 1953. Neue Begoniaceen von O. Beccari in Malesia gesammelt.

Webbia 9: 469-509.

Kiew, R.2005. Begonias of Peninsular Malaysia. Natural History Publications,

Borneo.

Laumonier, Y. 1997. The Vegetation and Physiography of Sumatra. Kluwer,

Dordrecht.

Sodhi, N.S., L.P. Koh, B.W. Brook and P.K.L Ng. 2004. Southeast Asian

biodiversity: an impending disaster. Trends in Ecology and Evolution 19:

654-660.

Gardens’ Bulletin Singapore 61 (1): 45-48. 2009 45

A New Record for the Flora of Vietnam:

Geodorum citrinum Jacks. (Orchidaceae)

DD HUYEN

Institute of Ecology and Biological Resources,

18 Hoang Quoc Viet Road, Cau Giay, Hanoi, Vietnam

Abstract

Geodorum citrinum Jacks. is a new record for the orchid flora of Vietnam.

Introduction

Geodorum Jacks. (Orchidaceae) is a small genus in Vietnam, represented by

five species (Averyanov and Averyanova, 2003; Ban et al., 2005).

During the years 2004-2008 several expeditions have been organized

in the Thai Nguyen province. Many plants have been added to the orchid

living collection of the Institute of Ecology and Biological Resources (IEBR).

Among them a new record in the genus Geodorum (Orchidaceae) for the

Flora of Vietnam has been made. Thus the genus Geodorum in Vietnam

consists of six species.

Geodorum citrinum Jacks. Andr. Bot. Rep. 3, t. 626 (1811); Seidenf. in

Opera Bot. 72:55, fig. 28 (1983). Figs. 1A-G, Plate 1A-D.

Terrestrial herb, stem short and erect, with ovate pseudobulb; 2-4 alternate

leaves with sheathing footstalks, ovate or oblong, 19-24 x 5-6.5 cm, apex

acute; scape lateral, erect, racemes terminating in a cernuous spike, with

5-6 flowers; flowers 4.5-5 cm in diameter; pedicel plus ovary 1.3-1.5 cm long:

sepals lanceolate, apex obtuse or acute, white green, 2.5-2.7 x 0.8-0.9 cm;

petals ovate-oblong, apex obtuse, white, 2.1-2.4 x 0.8-1.1 cm; labellum ovate,

2.1-2.3 x 1.5-1.7 cm, entire, base concave and glabrous, surface yellow, with

red-purple longitudinal stripes on epichile; column 3 mm long, column-foot

somewhat curved, with purple veins, 5 mm long; operculum white with two

violet prickles. Pollinia 2, ovaliform, wax-yellow.

Specimen examined: Vietnam. Thai Nguyen Province, Dinh Hoa

district, terrestrial in evergreen forest, 500-600 m alt., flowering from April

to June, Huyen 518 (HN).

Geodorum citrinum is here newly reported for the Flora of Vietnam.

The plant was identified using Seidenfaden's (1983) treatment and Pham

46 Gard. Bull. Singapore 61 (1) 2009

Figure 1. Geodorum citrinum Jacks. A. Plant habit; B. Flower front view; C. Dorsal sepal; D.

Lateral sepal; E. Petal; F. Lip; G. Operculum and pollinia.

A New Record for the Flora of Vietnam 47

rT." oe

Vee ea S.

Plate 1. Geodorum citrinum Jacks. A-B. Plant habit; C-D. Close-up of a single flower.

48 Gard. Bull. Singapore 61 (1) 2009

(2000). Our material has white-green flowers, turning yellow with age and

with some purple markings on the lip and column.

The species is also distributed in the neighbouring countries Thailand

and Malaysia, and in Bangladesh and Myanmar.

Acknowledgments

The authors are grateful to Prof. Le Xuan Canh at IEBR for assistance in the

field work, and to Dr. Chin See Chung and Dr. B.C. Tan for providing access

to the Singapore Herbarium (SING). This work was partially supported by

a Singapore Botanic Gardens Research Fellowship.

References

Averyanoyv, L. V. & Averyanova A. L. 2003. Updated checklist of the Orchids

of Vietnam. Vietnam National University Publishing House. Hanoi.

Ban, N. T., Averyanov L.V. & Huyen D. D. 2005. Orchidaceae. Checklist of

Plant Species of Vietnam, 3. Agriculture Publishing House. Hanoi.

Pham Hoang Ho, 2000. Illustrated Flora of Vietnam 3. TRE Publications,

Hochiminh City.

Seidenfaden, G. 1983. Orchid genera in Thailand XI. Cymbidieae Pfitz.

Geodorum. Opera Botanica 72: 47-64.

Gardens’ Bulletin Singapore 61 (1): 49-53. 2009 49

A New Species of Fissidens (Bryopsida, Fissidentaceae)

from Peninsular Malaysia

ZUWAISUKI K-T YONG AND

TSUZUKI

The Hattori Botanical Laboratory, Okazaki Branch, 10-3 Mutsuna-shin-machi,

: Okazaki- shi, Aichi-ken 444-0846, Japan

~ Institute of Biological Sciences, Faculty of Science, University of Malaya, 50603

: Kuala Lumpur, Malaysia

" The Hattori Botanical Laboratory, Shimada Branch, 6480-3 Takasago-cho,

Shimada-shi, Shizuoka-ken 427-0054, Japan

Abstract

Anew moss species collected from Peninsular Malaysia, Fissidens benitotanii

Z.Iwats., K.-T. Yong & Tad.Suzuki is described. The species belongs to

subgenus Fissidens section Fissidens. Most of the members in sect. Fissidens

have smooth laminal cells, except for a few species, which included this newly

described species, possess unipapillose laminal cells. Fissidens benitotanii

is easily distinguished from other Fissidens species in the region by the

following characteristics: narrowly lanceolate leaves with shortly excurrent

costa, thin limbidia that disappear near leaf apex, and unipapillose laminal

cells with distinctive sharp papilla.

Introduction

Iwatsukiand Mohamed (1987) reported 24 species of Fissidens for Peninsular

Malaysia and Singapore in a revision of the genus. Recently, we discovered an

interesting species while studying some of the Fissidens specimens collected

by one of the co-authors, K.-T. Yong from Peninsular Malaysia. After

studying the specimens carefully, and comparing them with related known

species, we reached the conclusion that one of the specimens represents a

new species belonging to subgen. Fissidens sect. Fissidens. However, this

plant specimen in question is different from the typical members of sect.

Fissidens in having distinctly unipapillose laminal cells.

Fissidens benitotanii Z.lwats., K.T. Yong & Tad.Suzuki, sp. nov.

Sterilis. Caulis 4.5-5.0 mm longus, cum foltis 2.0-2.5 mm latus. Folia 9-13 juga,

anguste lanceolata, 1.3-1.7 mm longa, 0.25-0.35 mm lata; apice acuto; costa

excurrenta; cellulae laminarum unipapillosa; marginibus limbatus.

50 Gard. Bull. Singapore 61 (1) 2009

— Holotypus: Peninsular Malaysia, State of Perak, Belum Royal Park, Upper

Belum Region, upstream of Kenarong River, lowland forest, alt. 200-300 m,

on soil, 30 Jul 2003, coll. K.-T. Yong 4660 (KLU; isotype in NICH, SING).

Figs. 1, 2.

Medium-sized plant for the genus. Stems simple, 4.5-5.0 mm long, 2.0-2.5 mm

wide with leaves; axillary hyaline nodules not differentiated; in cross-section

cortical cells small and thick-walled, central strand weakly differentiated.

Leaves 9-13 pairs, more or less densely arranged; middle to upper leaves

narrowly lanceolate, 1.3-1.7 mm long, 0.25-0.30 mm wide, narrowly acute

at apex; base of dorsal lamina rounded, not decurrent; costa stout, shortly

excurrent; margin almost entire, but finely crenulate by projection of cells

near apex; vaginant lamina about 1/2 of the leaf length. Limbidia usually

1-2 rows of elongate cells near apical laminae, 3-4 rows of cells on vaginant

laminae; in cross-section 1-2 cells thick; lacking near leaf apices. Laminae in

cross-section 1 cell thick; cells of apical lamina quadrate to hexagonal, 8-12

um long, moderately thick-walled, sharply unipapillose; cells of vaginant

laminae similar to those of apical laminae, but larger and longer toward

base. Dioicous?; male plants and sporophytes not found.

The species name, Fissidens benitotanii, is dedicated to Dr. Benito

C. Tan, an active bryologist in Asia who has spent many years studying

the tropical Asiatic bryophytes, particularly the moss species, and have

contributed many important papers to the study of Asiatic moss flora. In

addition, the honoree, B.C. Tan, is a mentor of K-T. Yong, one of the co-

authors of this paper, in his study of the moss flora of Malaysia.

The new species, Fissidens benitotanii, is placed within subgenus

Fissidens sect. Fissidens (cf. Suzuki and Iwatsuki, 2007) that is characterized

by the presence of limbidia on the apical, dorsal and vaginant laminae. By

and large, the species in sect. Fissidens are known to have smooth laminal

cells, but there are exceptional cases where a small number of them have

papillose laminal cells. Those species with unipapillose laminal cells are

thus far only been reported from the tropical areas; for example, Fissidens

angustifolius Sull. (syn. FE. dixonianus E.B. Bartram) is found in Fiji Islands,

Central and South America (cf. Iwatsuki and Suzuki, 1996; Pursell, 2007),

and F raiatensis E.B. Bartram (cf. Whittier and Miller, 1967) from Society

Islands.

The new species, F benitotanii, can be distinguished from F

angustifolius and F. raiatensis by its limbidia disappearing at the leaf apex,

its narrowly lanceolate leaves with acute apices, and the shortly excurrent

costa (Fig. 1g), whereas F. angustifolius has percurrent costa, and limbidia

always reaching the leaf tips. Also, papilla on the laminal cell of F benitotanii

is much sharper comparing with that of F angustifolius. Hyaline nodules are

A New Species of Fissidens from Peninsular Malaysia 51

Figure 1. Fissidens benitotanii Z.1wats., K.-T. Yong & Tad.Suzuki. A. Plant. B-D. Leaves. E-F.

Leaf apices; G. Cells at leaf apex; H. Marginal cells of apical lamina; I. Cells of apical lamina.

J. Cells at margin of vaginant lamina; K-L. Cross-sections of leaf; M. Cross-section of stem.

All figures drawn from the isotype specimen (K.-T. Yong 4660 in NICH). Scale bars in ym.

SZ Gard. Bull. Singapore 61 (1) 2009

present on the stems of F. angustifolius, but are absent in F. benitotanii.

The type material for F benitotanii is sterile, with no sporophytic

structure. We made a few attempts to search for the archegonia and antheidia

of this species, but were not successful. Sexuality of this species is probably

dioicous.

5 é ;

SUL d : i, Pad sere 20m

Figure 2. Fissidens benitotanii Z.Iwats., K.-T. Yong & Tad.Suzuki. A. Plant; B. Leaf apex; C.

Median part of leaf; D. Basal part of leaf; E. Papillose cells at middle of leaf; F. Cross-section

of leaf; G. Cross-section of stem; All figures were from the isotype specimen (K.-7: Yong 4660

in NICH).

A New Species of Fissidens from Peninsular Malaysia 55

Acknowledgements

Grateful acknowledgement is made to the Japan Society for the Promotion

of Science for financial support made available through a Grant in Aid

for Scientific Research © (no. 19570098) and a research grant provided

by University of Malaya, Malaysia. The authors also like to express their

appreciation to Madam Patricia Loh who helps us to improve the English

text.

References

Iwatsuki, Z. and M.A. Haji Mohamed. 1987. The genus Fissidens in

Peninsular Malaysia and Singapore (a preliminary study). Journal of

Hattori Botanical Laboratory 62: 339-360.

Iwatsuki, Z. and T. Suzuki, 1996. Fissidens in the Fiji Islands. Journal of

Hattori Botanical Laboratory 79: 139-162.

Pursell, R., 2007. Fissidentaceae, 278 pp. In: Flora Neotropica Monograph

101. New York Botanical Garden Press, New York.

Suzuki, T. and Z. Iwatsuki, 2007. A new approach to the infrageneric

classification of the genus Fissidens (Fissidentaceae, Bryopsida). Hikobia

15: 67-85.

Whittier, H.O. and H.A. Miller, 1967. Mosses of the Society Islands: Fissidens.

Bryologist 70(1): 76-93.

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Gardens’ Bulletin Singapore 61 (1): 55-72. 2009

Taxonomic Studies of Podostemaceae of Thailand.

3. Six New and a Rediscovered Species

M. KATO AND S. KOI

Department of Botany, National Museum of Nature and Science,

Tsukuba 305-0005, Japan. E-mail: sorang@kahaku.go.jp

Abstract

Podostemaceae is an ecologically and morphologically unusual aquatic

plant family. By examination of new collections from Thailand, we describe

seven species, two of which are new species assigned to TJerniopsis of

Tristichoideae (7? chanthaburiensis, T: minor), four are new species assigned

to Hydrobryum and Polypleurum of Podostemoideae (H. phetchabunense,

P. insulare, P. prachinburiense, P. sisaketense), and one is Zeylanidium

lichenoides rediscovered. In total, two subfamilies, 10 genera, and 42 species

with four varieties occur in Thailand. A key to all the species is provided.

Introduction

Podostemaceae is an aquatic angiosperm family distributed in the tropics

and subtropics of the world. The family is unusual ecologically and

morphologically. Plants grow submerged on rock surfaces during the rainy

season, and they flower and set fruits, while protruding and drying above

the water, during the following dry season when the water level becomes

low. The plants either adhere to the rocks by ribbon-like or crustose roots

developed from the hypocotyl in most species with rudimentary or no

primary shoots and roots, or by crustose or multi-branched adventitious

shoots in rootless species. Kato (2004, 2006) and Koi er al. (2008) reported

nine genera and 35 species from Thailand, the largest species number in

Asia. In this paper, we describe six new species referred to four genera and a

poorly known species, based on new molecular and morphological data and

new collections. Molecular phylogenetic data (S. Koi, unpubl. data) are useful

to identify species that are morphologically slightly different from its close

relatives. As a result, 10 genera and 42 species assigned to two subfamilies

occur in Thailand, indicating again the highest diversity of Podostemaceae

in Asia. A key to all the species of Thailand is provided.

56 Gard. Bull. Singapore 61 (1) 2009

Taxonomy

Terniopsis chanthaburiensis M. Kato & Koi, sp. nov.

Terniopsis malayana ramulis ramosis similis, sed pedicellis longioribus

differt; T. brevis plantis parvis, ramulis usque brevibus, ad 3.5 mm longis

similis, sed pedicellis longioribus differt. — Holotypus: Southeastern Thailand.

Chanthaburi Prov., Klong Yai, Pong Nam Ron District, 150 m alt., 12°56' N,

102°28' E, fl. Mar 2005, 7. Wongprasert 771601 (BKF; isotype, TNS). Fig. 1.

Figure 1. Terniopsis chanthaburiensis. A. Ramuli on flank of root; B. Reproductive shoot unit

(at right), a flower comprises lobed calyx, two stamens and an ovary on pedicel; Based on

Kato et al. TL-1402 (1A), Wongprasert 771601 (1B). Scale bars = 1 mm.

Root creeping, monopodially branched, flattened-subcylindrical, 0.5-1

mm wide; ramuli on both flanks of root, 3-6 mm long, simple or to 5 times

branched; leaves in 3 ranks, oblong-elliptic, univeined, middle leaf to 1 mm x

0.5 mm, lateral leaf to 1-2 mm = 0.5-0.8 mm. Flowering shoots 1-4 per shoot

Taxonomic Studies of Podostemaceae of Thailand S7/

unit, associated with 4-5 ramuli to 5 mm long; pedicel with 2 bracts at base,

5-10 mm long; calyx membranaceous, 3-lobed 1/2 to base, 2/3-3/4 length of

ovary; stamens 2, as long as ovary or shorter, ca 1.5 mm long; ovary obovoid-

ellipsoid, ca 2 mm long, ca 1 mm thick, 3-locular; stigmas 3, separated from

each other, cristate, 0.3 mm long; capsule stalked (stalk to 10 mm long),

trigonous, ribs 9.

Other specimens examined: Southeastern Thailand. Chanthaburi Prov.:

Klong Yai, Pong Nam Ron District, 150 m alt., 12°56' N, 102°28' E, st. Mar, M.

Kato et al. TL-1402 (BKF, TNS); fl.-buds Jan, M. Kato et al. TL-1607 (BKF,

TNS).

Ecology and distribution: On rocks in stream; vegetative plants submerged

in the rainy season, and reproductive plants exposed in the air in the dry

season; known only from the type locality.

Notes: Phylogenetically, this is a sister taxon to 7? malayana, T: brevis

and 7? minor, and together, sister to the clade of 7: ubonensis in eastern

Thailand and 7. sessilis in eastern central China (S. Koi, unpubl. result). This

indicates that small morphological changes occurred in the diversification

of Terniopsis: Thai species, in particular, 7? brevis, T: chanthaburiensis, and T.

minor, are little distinguished from the Chinese 7° sessilis (Chao, 1980; Kato

and Kita, 2003), but 7? chanthaburiensis is distinct in the long pedicel. Also,

T. chanthaburiensis is similar to T: malayana in the branched ramuli on the

floriferous root, bud differs in having the long pedicel.

Terniopsis minor M. Kato & Kol, sp. nov.

A T. brevis simile, sed radicibus latioribus, ramulis longioribus, ad 3.5 mm

longis, stigmatibus linearis differt. — Holotypus: Southeastern Thailand.

Trat Prov., Klong Kaeo waterfall, Bo Phloi village, Bo Rai Distr., 12°37'29"

N, 102°34'54" E, 170 m alt., fl. Jan 2007, M. Kato, S. Koi, N. Katayama & T.

Wongprasert TL-1609 (BKF, isotype, TNS). Fig. 2.

Root creeping, monopodially branched, flattened-subcylindrical, 0.7-1.5 mm

wide; ramuli on both flanks of root, 4-7 mm long, simple; leaves in 3 ranks,

oblong-elliptic, middle leaf to 1 mm x 0.3 mm, lateral leaf to 1.1 mm x 0.3 mm.

Flowering shoot associated with single ramulus 1-3 mm long; pedicel with

2 bracts at base, 1.5-3 mm long; calyx membranaceous, 3-lobed 1/4 to base,

as long as ovary; stamens 2, as long as ovary, 1-1.5 mm long; ovary obovoid-

ellipsoid, 1-1.5 mm long, ca 0.7 mm thick, 3-locular; stigmas 3, separated

from each other, linear-oblong, apex cristate, to 0.4 mm long; ovules ca 20

per locule; capsule stalked (stalk 2-3 mm long), trigonous, ribs 9.

58 Gard. Bull. Singapore 61 (1) 2009

Figure 2. Terniopsis minor. A. Ramuli on flank of root; B. Two reproductive shoot units; left

one with flower comprising lobed calyx, two stamens and ovary on pedicel; right one with

flower bud. Based on Kato et al. TL-1609. Scale bars = 1 mm.

Other specimens examined: Nakhon Ratchasima Prov., Haew Narok

Waterfall, Khao Yai National Park, 14°17' N, 101°24' E, 360 m, st. Dec, Jan,

Kato et al. TL-60, TL-309 (BKE, TNS); fi. Jan, TL-1535 (BKE TNS); st., fi.

& fr. Jan, R. Imaichi et al. TKF-25 (BKF, TNS). Chanthaburi Prov., Klong

Yai, Pong Nam Ron District, 150 m alt., 12°56' N, 102°28' E, fl.-buds Jan, M.

Kato et al. TL-1608 (BKF, TNS). Trat Prov., Sato stream, Klong Sato, Bo Rai

District, 12°42' N, 102°25' E, st., fl. & fr. Jan, S. Koi et al. TK F-22 (BKF, TNS);

Pong Nam Ron stream, Klong Yai, Pong Nam Ron District, 12°56' N, 102°28'

E, st. Jan, S. Koi et al. TK F-23 (BKF, TNS).

Ecology and distribution: On rocks in stream; vegetative plants submerged

in the rainy season, and reproductive plants exposed in the air during the

dry season; known only from the type locality and its vicinity.

Note: This species is phylogenetically sister to 7: brevis and morphologically

close to it also. However, it differs from 7: brevis only in having wider roots,

longer ramuli, and linear-oblong stigmas.

Taxonomic Studies of Podostemaceae of Thailand 59

Polypleurum sisaketense M. Kato & Koi, sp. nov.

A congenris stigmatibus semiorbiculatis differt, P. wongprasertii, P. longifoliis,

P. phuwuaensis et P. rubroradicans floris sessilibus, bracteis filiformibus, basi

vanigatis simile, sed a P. wongpraserti ovulis plura, a P. longifoliis foliis

brevioribus, a P. phuwuaensis spathellis brevioribus, a P. rubroradicans

ovullis plura differt. — Holotypus: Eastern Thailand. Si Sa Ket Prov.,

Phoolaor waterfall, Kantharalak Dist., 14°27'17" N, 104°39'21" E, 180 m

alt., fl. Jan 2006, M. Kato, S. Koi & T: Wongprasert TL-1502 (BKF; isotype,

TNS). Fig. 3.

Figure 3. Polypleurum sisaketense. A. Flower buds enclosed by spathellas subtended by

bracts at sinuses of root branches; two spathellas are rupturing; B. Flower comprising two

tepals (one seen), one stamen and one ovary on pedicel with spathella and bracts at base; C.

Fruit with spathella and bracts; D. Seeds in capsule with valve removed. All figures based on

Kato et al. TL-1502. Scale bars = 1 mm.

Root ribbon-like, 2-3 mm wide, branched, with tufts of leaves on dorsal

surface exclusively at sinuses between root branches; leaves 2-4 per tuft, 5-20

mm long, base sheath-like, scars forming mounds, distal part needle-like,

relatively thick (ca 0.2 mm), caducous. Flowering shoot on dorsal surface at

sinuses between root branches; bracts 2-4, sheath-like, papillate, needle-like

with sheath-like base or ovate with needle-like part caducous; flower 1, bud

covered by spathella, spathella 1.5-2 mm long, papillate, ruptured near apex

60 Gard. Bull. Singapore 61 (1) 2009

and also split longitudinally at anthesis, ellipsoid, narrowed to base; pedicel

4-8 mm long; tepals 2, 1 on each side of stamen, ca 0.5 mm long; stamen 1,

inserted at base of ovary, 1.5-1.8 mm long, as tall as ovary; ovary 2-locular,

globose-ellipsoid, slightly flattened, 1-1.5 mm long; stigmas 2, hemicircular,

flattened, 0.1-0.2 mm long and wide, subequal (stigma facing stamen larger);

ovules covering entire septum surface, 40-70 per locule; capsule stalked

(stalk 5-10 mm long), subsymmetric, ribs ca 12.

Ecology and distribution: On rocks near waterfall; vegetative plants

submerged in the rainy season, and reproductive plants exposed in the air

in dry season; known only from the type locality.

Notes: Although it is difficult to distinguish this species from other congeners

in most characters, it differs mainly in having semicircular, flattened stigmas.

Phylogenetically it forms a clade together with P. longifolium, P. phuwuaense

and P. erectum, but the clade has no obvious sister clade (S. Koi, unpubl.

data).

~ Polypleurum prachinburiense M. Kato & Koi, sp. nov.

A congenris alabastris globosis differt. - Holotypus: Southeastern Thailand.

Prachingburi Prov., Kaeng Wung Sai, Kaeng Hin Phoeng waterfall, Na Dee

Dist. Prachinburi Prov., 14°10' N, 101°43' E, 70 m alt., fl. & fr. Jan 2007, M.

Kato, S. Koi, C. Tsutsumi, N. Kitagawa & T: Wongprasert TL-1601-1 (BKF;

isotype, TNS). Fig. 4.

Root ribbon-like, 2-4 mm wide, branched, with tufts of leaves on dorsal

surface exclusively at sinuses between root branches; leaves 2-4 or more per

tuft, 10-25 mm long, base sheath-like, ovate, papillate, distal part needle-like.

Flowering shoots on dorsal surface at sinuses between root branches; flower

buds, along with bracts, globose; bracts 2(-4), ovate, mucronate or rarely

needle-like, papillate, thick; flower 1, bud covered by spathella, spathella

papillate, 1-1.5 mm long, ruptured near apex at anthesis; pedicel 4-8 mm long;

tepals 2, 1 on each side of stamen, 0.7-1 mm long; stamen 1, inserted above

base of ovary, 1.5-2 mm long, slightly longer than ovary; ovary 2-locular,

ellipsoid, slightly flattened, 1-1.5 mm long; stigmas 2, narrowly conical,

pointed, 0.2-0.4 mm long, subequal; ovules covering all or most of septum

surface, 15-20 per locule; capsule stalked (stalk 5-9 mm long), subsymmetric,

ribs 12-14.

Other specimens examined: Southeastern Thailand. Prachingburi Prov.,

Takro waterfall, 35 m alt., 14°11' N, 101°36' E, fl. & fr. Mar, M. Kato et al.

TL-1404 (BKF, TNS); fl. Jan, M. Kato et al. TL-1611, TL-1612 (BKF, TNS);

Taxonomic Studies of Podostemaceae of Thailand 61

Figure 4. Polypleurum prachinburiense. A. Flower buds enclosed by spathellas subtended

by bracts at sinuses of root branches; B. Flowers with spathellas and bracts at sinuses of root

branches: a flower comprises two tepals (one seen), one stamen and one ovary on pedicel

with spathella and bracts at base: C. Fruit with spathella and bracts; D. Seeds in capsule with

valve removed. Based on Imaichi et al. TIK-20 (4A) and Kato et al. TL-1601-1 (4B-D). Scale

bars = 1 mm.

Kaeng Hin Phoeng waterfall, Na Dee Dist., 14°10' N, 101°43' E, 70 m alt., fl.

Jan, M. Kato et al. TL-1534 (BKF,.TNS): fl. & fr. Jan, M. Kato et al. TL-1601-2,

TL-1601-3, TL-1602, TL-1603, TL-1604 (BKF, TNS): fl. Jan, R. Imaichi et al.

TIK-20 (BKF, TNS).

Ecology and distribution: On rocks in stream; vegetative plants submerged

in the rainy season, and reproductive plants exposed in the air in the dry

season; known only from the type locality and its vicinity.

62 Gard. Bull. Singapore 61 (1) 2009

Notes: This species is characterized by the globose flower-buds with usually

a single pair of mucronate bracts. In this character it differs from the most

closely related P. wongprasertii whose bracts are attenuate at the apex. In

the other characters it is most similar to P. wongprasertii. P. prachinburiense

forms a clade together with P. wongprasertii, P. ubonense,P. sisaketense, P.

longifolium, P. phuwuaense and P. erectum, but their inter specific relationship

are not clean (S. Koi, unpubl. data).

Polypleurum insulare M. Kato & Kol, sp. nov.

A P. wallichio, P.schmidtiano er P. longistyloso foliis et floribus supra radicibus

ad omnem ramificatinem differt, a P. wongprasertii radicibus angustioribus,

bracteis parvioribus, pedicellis brevioribus, ovartis parivioribus differt.

— Holotypus: Southeastern Thailand. Trat Prov., Khlong Phu waterfall, Ko

Chang, 12°04'04" N, 102°18'53" E, 125 m alt., fl. & fr. Jan 2004, M. Kato, S. Koi

& T: Wongprasert TL-1512 (BKF, isotype, TNS). Fig. 5.

Figure 5. Polypleurum insulare; A. Flowers with spathellas and bracts at sinuses of root

branches; a flower comprises two tepals (one seen), one stamen and one ovary on pedicel

with spathella and bracts at base; B. Fruit with spathella and bracts. Based on Kato et al. TL-

1512. Scale bars = 1 mm.

Taxonomic Studies of Podostemaceae of Thailand 63

Root ribbon-like, 1-2 mm wide, branched, with tufts of leaves on dorsal

surface exclusively at sinuses between root branches; leaves 3-5 per tuft,

5-15 mm long, base sheath-like, ovate, papillate, persistent, forming a mound

around leaves, distal part needle-like, caducous. Flowering shoot on dorsal

surface at sinuses between root branches; bracts 2-4, papillate, ovate, obtuse,

small (to 0.7 mm long) or rarely with needle-like tips; flower 1, bud covered

by spathella, spathella 1.5-2 mm long, papillate, appressed, ruptured near

apex and also split longitudinally at anthesis; pedicel 2-3 mm long; tepals 2,

1 on each side of stamen, ca 1 mm long; stamen 1, ca 1 mm long, as tall as

ovary; ovary 2-locular, ellipsoid, slightly flattened, ca 1 mm long; stigmas 2,

linear, narrowed to apex, 0.2-0.3 mm long; ovules on septum surface except

in central area, 10-12 per locule; capsule stalked (stalk 2.5-4 mm long),

subsymmetric, ribs 8-12.

Other specimens examined: Southeastern Thailand. Trat Prov., Khlong Phu

waterfall, Ko Chang, 12°04'04" N, 102°18'53" E, 90 m alt., st. Feb, M. Kato et

al. TL-1304 (BKF, TNS); Tharn Mayom waterfall, Ko Chang, 12°04'17" N,

102°20'57" E, 160 m alt., st. Jan, M. Kato et al. TL-1521(BKF, TNS); Tharn

Mayom waterfall, Ko Chang, 12°04'17" N, 102°20'42" E, 235 m alt., st. Jan, M.

Kato et al. TL-1526 (BKF, TNS); Klong Phloo, Ko Chang, fr. Feb, F Konta &

T. Wongprasert s.n. (BKF, TNS); Ko Kut, 12°35' N, 101°31' E, Charoenphol et

al. 5115 (AAU, BKEF, K).

Ecology and distribution: On rocks in streams; vegetative plants submerged

in the rainy season, and reproductive plants exposed in the air during the

dry season; known from Ko Chang and Ko Kut, the two offshore islands in

southeastern end of Thailand.

Notes: Kato (2006) wrongly identified specimens (fF: Konta & T: Wongprasert

sn.) from Ko Chang and Charoenphol et al. 5115 from Ko Kut to be P.

wongprasertit. Charoenphol et al. 5115 was also wrongly identified as P.

schmidtianum (Cusset, 1992). A comparison with new collections and a

molecular phylogenetic result shows that plants of the islands are distinct

from the continental species and are to be separated as a new species. In

a phylogenetic tree, P. insulare is isolated from P. wongprasertii as equally

as from P. erectum, P. longifolium, P. phwuaense, P. prachinburiense and P.

sisaketense, and distant from P. schmidtianum. Polypleurum insulare differs

from P. wongprasertii by its narrower roots, very short bracts, shorter pedicels,

and smaller ovaries, and from those closely or distantly related species by the

diagnostic characters shown in Key. There are now two species, P. insulare

and P. schmidtianum, occurring on the small offshore islands in southeastern

Thailand.

64 Gard. Bull. Singapore 61 (1) 2009

Zeylanidium lichenoides (Kurz) Engler

Nat. Pflanzenfam. 2nd ed. 18a (1930[1928]) 62; van Royen, Danks Bot. Arkiv

23 (1965) 165; Cusset, Adansonia 14 (1992) 31; Mathew & Satheesh, Aquat.

Bot. 57 (1997) 265, f. 15, 16. — Typus. Parish s.n., Martaban, Myanmar (CAL,

not seen).

Roots ribbon-like, 1-2 mm wide, branched, with tufts of leaves or flowering

shoots at sinuses of root branches; leaves linear, 20-30 mm long, ensiform,

ca 5 per tuft in 2 files. Flowering shoots solitary, appressed; bracts 4-5 in 2

files, uniform but basal ones smaller, basal part ovate to ovate-lanceolate,

0.8-1.5 mm long, distal part linear, 5-7 mm long, caducous; each flower bud

enclosed by spathella, spathellas ellipsoidal, 1.2-1.8 mm long, longitudinally

split at anthesis, persistent: flowers erect; pedicels horizental at base, upright

upwards, 1.2-1.5 mm long; tepals 2, one on each side of stamen, linear, 0.5-

0.8 mm long; stamens 2 with flattened andropod, branched 1/3-1/4 from tip,

1.8-2.0 mm long, as long as pistil; anthers ellipsoidal, ca 0.5 mm long; ovaries

single, sessile, unilocular, ellipsoidal, 1.2-1.8 mm long, ca 1.0 mm wide;

stigmas 2, forked above or at base, equal, narrowly triangular, entire, ca 0.5

mm long; ovules 82-115 per ovary, born on whole flat placentas; capsule

stalked (stalk 1.6-2.0 mm long), ellipsoidal, 1.5 mm long, ca 1.0 mm wide,

8-ribbed, dehiscing by 2 unequal valves.

Other specimens examined: Northern Thailand. Chiang Mai, Huay Kaew

stream, Mae On, 600 m alt., 18°51.8' N, 99°18.0' E, st. fl.-buds, fl. & fr. Dec., S.

Koi & T: Wongprasert TK-02, TK-04, TK-05 (BKF,TNS); Huay Kaew stream,

Mae On, 650 m alt., 18°52.9' N, 99°17.5' E, fl. & fr. Feb, M. Kato et al. TL-1703,

TL-1704 (BKF, TNS).

Ecology and distribution: On rocks in stream; sterile plants submerged and

flowering and fruiting plants exposed; N Thailand, SE Myanmar, NE and S

India.

Notes: This is the second report of Zeylanidium from Southeastern Asia, and

northern Thailand is the eastern margin of distribution. It was recorded for

the first time from Doi Suthep and Doi Inthanon, Chiang Mai, in northern

Thailand (van Royen, 1965). But we have not collected the genus in these

areas and did not examine specimens cited by van Royen. The identification

here is based on our morphological observation and molecular data (S. Koi,

unpubl. data).

Species identification is tentative, because we did not examine the

type and authentic specimens, particularly from southeastern Myanmar

and northeastern India. Comparison of Cusset’s (1992) and Mathew

Taxonomic Studies of Podostemaceae of Thailand 65

and Satheesh’s (1997) descriptions and our collection suggests that the

Thai specimens may be referable to Zeylanidium lichenoides, but final

identification requires detailed comparison. The species delimitation is

another issue, because our preliminary phylogenetic data suggest a large

molecular difference between Thai and southern Indian plants.

Hydrobryum phetchabunense M. Kato & Koi, sp. nov.

Hydrobryum loeicum bracteis 2-4, spathellis irregulariter ruptis, stigmatibus

linearibus, curvatis, acutissimibus, ovulis 5-8 in quoque loculo, capsulis

16-20-costatis simile, sed paginis radicibus circum folia annularibus,

protrudentibus, pedicellis apicipus bracteis altioribus differt. — Holotypus:

Northeastern Thailand. Phetchabun Prov., Thadphramba waterfall, Nam

Nao Natl. Park, 750 m alt., 16°45' N, 101°39' E, fl. & fr. Feb 2004, M. Kato &

T. Wongprasert TL-1102 (BKF; isotype, TNS). Fig. 6.

Root crustaceous, irregularly lobed, raised annually around tufts of patent

leaves; leaves 1-4 per tuft, needle-like, terete, 2-3.5 mm long. Flowering

shoots appressed, with flower solitary at tip; bracts uniform, 2-4, ovate,

1-1.5 mm long; spathella smooth-surfaced, irregularly ruptured near apex

at anthesis; ovary-stalk (pedicel) 0.5-1.0 mm long, young one shorter; tepals

2, one on each side of stamen, linear, ca 1.5 mm long; stamens 2, branched

below middle, 2-3.5 mm long, common andropod (filament) shorter than

ovary; ovary 2-locular, ellipsoid, 1.5-2 mm long; stigmas 2, equal, ca 0.5 mm

long, linear, pointed, curved; ovules on marginal surface of septum, 5-8 per

locule; capsule stalked (stalk 1-1.5 mm long, higher than top of bracts),

ellipsoid, flattened, ribs 16-20.

Other specimen examined: Northeastern Thailand. Phetchabun Proyv.,

Thadphramba waterfall, Nam Nao Natl. Park, 750 m alt., 16°45' N, 101°39' E,

fl. & fr. Jan, S. Koi et al. TK F-0] (BKF,TNS).

Ecology and distribution: On rocks in stream; vegetative plants submerged

in the rainy season, and reproductive plants exposed in the air in the dry

season; known only in type locality.

Notes: The plant TL-//02 was treated as Hydrobryum loeicum on the basis

of its few and large ovules (4-8 per locule), many (16-20) ribs on the capsule,

and linear, pointed stigmas (Kato, 2004). However, H. phetchabunense is

distinguished from H. loeicum in the raised rings on the upper surface of

root around the tufts of leaves, and the higher pedicels than the top level

of the uppermost bracts. A molecular phylogenetic analysis shows that

H. petchabunense is sister to the clade of H. loeicum and Diplobryum

66 Gard. Bull. Singapore 61 (1) 2009

Figure 6. Hydrobryum phetchabunense. A. Leaves scattering on crustose root; B. Flower buds

enclosed by spathellas subtended bracts appressed to crustose root; C. Flower comprising

two tepals, two stamens with forked filament, and one ovary on short pedicel (not seen) sub-

tended by spathella and bracts; D. Fruits with spathellas and bracts on crustose root; E. Fruit

with spathella and bracts; F. Seeds (six on placenta) in capsule with valve removed. Based on

Kato & Wongprasert TL-1102. Scale bars = 1 mm.

vientianense, but not very close to H. loeicum (S. Koi, unpubl. result). The

three species are similar in having linear stigmas and up to 20 ribs on the

capsule. However, in H. phetchabunense, like in H. loeicum, the spathella is

smooth-surfaced (vs. papillate in H. vientianense), the stamens are shorter

(vs. 4-5 mm), the ovules are fewer (vs. 9-27 per locule), and the stigmas are

shorter (vs. 0.6-1 mm) (see also Key below). Thus, H. phetchabunense is

similar to H. loeicum, although the latter is sister to H. vientianense.

Taxonomic Studies of Podostemaceae of Thailand 67

Key to the species of Podostemaceae of Thailand

The key to all species of Podostemaceae of Thailand combines the results of

studies of Kato (2004, 2006) and the present study.

Flowers 3-merous (perianth 3-lobed, stamens 2 or 3, ovary 3-locular,

stigmas 3): leaves flattened, oblong or ovate, univeined ...............:ceeeeee

cane oh ee UE es Se ee Subfamily Tristichoideae (2)

Flowers 2-merous (tepals 2, filiform, stamens 1-2, ovary 1-2-locular,

stigmas 2): leaves filiform or vertically flattened, not veined ..............0.

I a a RL ccczicesbeculisediensencesee Subfamily Podostemoideae (12)

Root subcylindrical or ribbon-like: shoot subcylindrical, simple or

ramified; cupule absent, flower bud instead embraced by bracts ........... 3

Root absent; shoot crustose or rarely broadly ribbon-like, leafy on dorsal

surface and at margin (not on ventral surface); flower bud covered by

WELLE BD LD sade ie es Ae er ae 9

Reproductive shoot-complex comprising 3 branches, middle vegetative, 2

laterals floriferous; flowering shoot 4-5 mm long with leaves below flower

eaaaay sii) TABKS. GATANALS oi 05. och ee bese Cussetia diversifolia

Reproductive shoot-complex comprising 1-4 floriferous and 1-5

vegetative branches, not arranged in above mode: flowering shoot short

with 2 or several bracts thick or membraneous, flat ............eeeeeeeeees -

Vegetative shoots to 5 cm long, many times branched, sparsely leafy,

distal part comprising ramuli; bracts several ........... Terniopsis ramosa

Vegetative shoots to 2 cm long, composed of ramuli with tristichous

imbricate leaves, proximal part of shoot a few times branched; bra ..... 5

Root 2-10 mm wide; ramuli 3-90 mm long; stamens 5-6 mm long; ovules

25 TS RC GL lcs ies ke ee Terniopsis ubonensis

Root 0.2-1.5 mm wide; ramuli less than 20(-30) mm long: stamens 1-4

ee ote cy PEST CO ces 7c Ill Ae Se er eee eee ee a ee 6

Pedicels5-10mmlong..........................+.... lerniopsis chanthaburiensis

BANGER MED 9 HEPAT INE Scie oo rs Gd ds Lis oS oe cace aad stu Receauececencuscauceecceueccuseuveesane J

Roots 0.2-1 mm wide; ramuli 2.2-3.5 mm long ................ Terniopsis brevis

Roots 0.7-4 mm wide; ramuli 3-20 mm long .......... 0.0.0... se eee eee ee ee ees 8

Ramuli 4-7 mm long; stamens 1-1.5 mm long............... Terniopsis minor

10.

—

Gard. Bull. Singapore 61 (1) 2009

Ramuli 3-20 mm; stamens 2-4 mm long .............. Terniopsis malayana

Shoot 3-10 mm wide or wider; dorsal leaves arranged in branched

longitudinal rows; pedicel 5-8 mm long, 2.5-4 times as long as ovary... 10

Shoot to 2.5 mm wide; dorsal leaves in 1-2 inconspicuous rows; pedicel

{5-4 mm long, 1-2 timesrasilong as OVany x. sess eee eee

Dorsal leaves dense, fimbriate; lateral leaves narrowly deltoid, to 1.5 mm

longsovulesica 30; perlocule:.t.---- eee Dalzellia ubonensis

Dorsal leaves sparse, separate; lateral leaves deltoid-lanceolate, to 2 mm

long ovulesa0-60iperlocule’...........2-.-t eee Dalzellia ranongensis

. Shoots 1.5-2.5 mm wide; pedicel 1.5-2 mm long, as long as ovary; ovules

50-60 perloculeesenan tui ee Dalzellia kailarsenii

. Shoots 1-1.5 mm wide; pedicel 2-7 mm long, longer than ovary (1.3-1.5

mumn)<ovules 30-50iperlocule steerer ee ee Dalzellia angustissima

Roots subcylindrical or mibbon-likesacetee ee ee

ROOUS CLUSTOSE « sisi sinsseadh ty Rae Meee ee eee

13. Bracts 3-4-lobed or digitate; capsules globose, smooth (or weakly striped)

13;

14.

live

or- ellipsoid; ribbed s2........c00GoR ie ee ec 14

Bracts simple; capsules ellipsoid, slightly flattened, 8-15-ribbed .......... 17

Holdfasts present on ventral surface of root under tufts of leaves; tufts

of leaves and flowering shoots borne on flank of root between successive

root branches and at sinuses of root bramches = ic:kersees ene 15

. Holdfast absent; tufts of leaves and flowering shoots only at sinuses of

root branches; bracts 3-4-lobed or digitate co - aes ee aa

. Bracts linear-oblong, with 2 small lateral basal lobes; capsules globose,

SHO O thc eee acta Paracladopus chiangmaiensis

. Bracts digitate; capsules ellipsoid; 12-14=tibbe Giiirc eerste -cce eee

i aga Sotlubaca bane SOO aoe reenact ee Paracladopus chathaburiensis

. Bracts 3-4-lobed, lobes semicircular, thin, smooth ...... Cladopus taiensis

. Bracts digitate, segments finger-like, thick, papillate with silica ................

er Re sesh. sees seo no sor aca so doa Roccaccbese0% Cladopus fallax

Stamens 2; capsullé=mibs, Syisgee eee tesco eee ae 18

Stamen 1: capsulle silos S25 cesses ees cc satcnescecee to cen eco 20

Taxonomic Studies of Podostemaceae of Thailand 69

18.

29):

£9.

20.

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pupae

2S.

2B.

24.

Tufts of leaves and flowering shoots at sinuses between root branches;

bracts ovate or ovate-lanceolate base, apex portion linear, caducous;

capsule-valves unequal ............................... Zeylanidium lichenoides

Tufts of leaves and flowering shoots on flanks of root between successive

root branches; bracts lanceolate or ovate-lanceolate, apex obtuse or

sometimes acute; capsule-valves equal ......... Polypleurum wallichii (19)

Roots ca 5 mm wide; leaves to 5(-10) mm long; pedicels 5-8 mm long;

SER TE 2S P0100 TCR Sa a var. wallichii

Roots ca 3 mm wide; leaves to 2.5 mm long; pedicels to 2-4 mm long;

Mia TA) SEUERENG LONER EY sen ceo o won cn vaca sav'nn Sosenscecedensncectassesessuace var. parvum

Tufts of leaves borne between successive root branches; capsule-ribs 8-12

se csg ee ESSE eae oe a at Al

Tufts of leaves or shoots borne exclusively at sinuses of root branches;

SADE LLSSOTI SS Les SS ee A er eer es 22

Roots 2-4 mm wide; pedicels 6-7 mm long; ovary protruding from

spathella at anthesis, 2-locular; stigmas much shorter than ovary; capsule-

Soe Gee eS eee ae ee Polypleurum schmidtianum

Roots 1-1.5 mm wide; pedicels to | mm long; ovary mostly enclosed in

spathella at anthesis, 1-locular;style plus stigmas as long as ovary or longer;

capsule-ribs 10-12, inconspicuous ........... Polypleurum longistylosum

Shoots prominent with leaves exposed on elongate stems; flowers always

Biercea Moncey MAN ple PET SHOOE .-.._...-..-:-.s0.0.e-nc-cced ccd acusenacedcesecbeceedeeneeD 3

Shoots comprising tufts of leaves, lacking stems; leaf-bases embedded

Minin FOOt; Hower always | per SHOOL .........ccisecceccesecceccosnccdeeesoeseessescesess 24

Roots 1-1.5 mm wide; shoots to 5 cm long, simple, erect; leaves 15-30 mm

Merete OWES 15-50) PEE LOCUIE io. .2c. 80s icckees caste lcs Polypleurum erectum

Roots 2.5-4 mm wide; shoots to 18 cm long, branched, bent at base and

floating; leaves 30-70 mm long; ovules 50-70 per locule ......... eset eeeeeeee

a ee aeons ease oso cannescvicn Polypleurum longicaule

Bracts 4-6; pedicels 10-15 mm long; spathellas 4-6 mm long ..................

eR ert PE MAN. S15 Feed $5ey Pass: Se sarah de Reena phuwuaense

Bracts 2-4(-6); pedicels 2-8 mm long; spathellas 2-3 mm long ..............25

Complex of flower bud and bracts not globose; bracts usually obvious,

ALLeUUALe at AEX OFALlENMate Lip CACUCOUS .cc5..2-c0ca-ssbeescnecacencnecesnesecetovecesees

28.

jas).

Gard. Bull. Singapore 61 (1) 2009

. Complex of flower bud and bracts globose; bracts inconspicuous (ovate

or elliptic), not attenuate to apex, basal bracts rarely attenuate ......... 26

26. Roots 1-2 mm wide; stalk of capsule 2.5-4 mm IONG .............:.essescesceeeseeceenes

Jessa ules acetate eae oe ee ee Polypleurum insulare

. Roots 2-3 mm wide; stalk of capsule 4-10 mm long ..................ee ee pz) |

. Stigmas semicircular; ovules 40-70 per locule ... Polypleurum sisaketense

. Stigmas narrowly conical, subdeltoid or deltoid-ovate; ovules 15-22 per

loCUIG Jsictiiitcndiices te ee EE eS . 28

. Leaves 20-40 mm long;stigmas subdeltoid or deltoid-ovate ...................

s dhad dave Retna Gek kee Sea Polypleurum longifolium

Leaves 10-25 mm long; stigmas narrowly conical, pointed ..................

i Re LAS Polypleurum prachinburiense

Flowering shoots erect; bracts in 4 ranks, dimorphic, ventral (facing root)

bilobed, dorsalssimple::23..252..:223. Se ee ee ee 30

. Flowering shoots appressed or oblique or only flowers erect; bracts in 2

ranks, unifommssmmple.or trilobed & 2.2.0..¢2cnces5stees econ

. Capsules 2-2.5 mm long, ribs 12-16 ............... Hanseniella heterophylla

. Capsules 1.5-2 mm long, ribs 8 ........................ Hanseniella smitinandii

. Bracts trilobed with middle lobe much ee than lateral lobes ...............

.. Thawatchaia trilobata

: Bee ae in one Ghee or sometimes acute or acuminate ...... 32

. Stamens: 2 with forked filament +353. ee

. Stamen 1 with simple filament }.0.52.22.25. ssc: 2k oc entce eee

. Bracts linear-lanceolate, acuminate; stigmas unequal ................:.-++. 34

. Bracts deltoid, elliptic or ovate, obtuse or acute; as Sana or

Suibeqiball. 322222 Wik Becness cnn annaveussehcenlonnecace vane ebe ug Rca ns eee ee 25)

. Roots (crusts) ca 0.5 mm thick, not markedly raised around tufts of

leaves; bracts 2-3; stigmas unequal, forked below middle ...............::::000

ssssundnas'ssdbilttes, SESE AY OR Sea ea ee eee Hydrobryum bifoliatum

. Roots (crusts) to ca 1 mm thick, prominently raised around tufts of

leaves; bracts 4-6; stigmas markedly unequal, forked above middle ........

seiepetUaeaact wocaceeneels ke ee eee Hydrobryum kaengsophense

Taxonomic Studies of Podostemaceae of Thailand 71

41.

. Leaves oblique or appressed. tough: bracts deltoid, acute at apex: stigmas

subequal ... pSiuee veseseeseesees-s---- Hydrobryum tardhuangense

. Leaves ae soft (but tough i in H. eee aiense): bracts ovate or elliptic.

TE TIS ET Ts as 0 eo | Me en ae eee 36

. Stigmas obovate, cristate ....................-.- .. Hydrobryum ee

2 SST PSR OLS

oe PELE LATTE | ee ee

| ELDEST ee eee |

| TLE Ts 1b ae Hydrobryum somranii

2 LIES TELE SO SOS nek nn 39

. Ovules 48 per locule ...................................--- Hydrobryum loeicum

PeMMRE Ss es Pee OEE MOINS ne se ant Hydrobryum vientianense

. Ribs on capsule 16-20 ...........................Hydrobryum phetchabunense

OE TEED STEWS 0 Sa re Se ene ee eee ee eee eee | |

Stigmas subentire, oblong or narrowly deltoid-lanceolate, emarginated,

truncate, or obtuse at tip; ovules 12-20 on septum surface except in small

eae eee aes al area lew 10) 70) rannme borage an aes

oo Ee Hydrobryum chiangmaiense

Stigmas entire, linear, pointed at tip; ovules 11-15 on marginal surface of

septum: leaves to 3 mm long ..................------------ Hydrobryum japonicum

2. Ovary 2-locular; placentation axile: leaves somewhat tough ......................

oo REE eee Hydrobryum khaoyaiense

- Ovary 1-locular: placentation pseudo-central: leaves soft .................-.--.--

eS ee Hydrobryum micrantherum (43)

- Roots (crusts) 0.1-0.2 mm thick: bracts 2-4: ovary ca 1.5 mm long: ovules

9-13 per placenta .. zs ......-.... War. micrantherum

. Roots (crusts) 0. 2.03 3 mm anes facies 3-5; ovary Az 5-2 mm long: ovules

i Re eR AIDE Seer A Si Sc edie eee oe es) VR

Acknowledgments

We thank T. Santisuk, T. Wongprasert, S. Suddee, H. Akiyama, R. Fujiami,

R. Imaichi, N. Katayama, Y. Kita, and C. Tsutsumi for their help in the field

studies, and curators of the Herbaria, Aarhus University (AAU), Royal

72 Gard. Bull. Singapore 61 (1) 2009

Forest Department (BKF), Bangkok, and Royal Botanic Gardens (K), Kew

for helping me examine specimens. Plant illustrations were drawn by M.

Kinoshita. This study was supported by a Grant-in-Aid for Scientific Study

from the Japan Society for the Promotion of Science.

References

Chao, H.-C. 1980. A new genus (Terniopsis, gen. nov.) of Podostemonaceae

from Fujian, China. Acta Botanica Yunnanica 2: 296-299.

Cusset, C. 1992. Contribution a I’ étude des Podostemaceae: 12. Les genres

asiatiques. Bulletin du Muséum National d'Histoire Naturelle Paris, #

Série, Section B, Adansonia 14: 13-54.

Engler, A. 1930(1928). Reihe Podostemales, pp. 1-68, 483-484. In: A. Engler

and K. Prantl (eds.), Die Naturlichen Pflanzenfamilien. 2nd ed. Vol. 18a.

Engelmann, Leipzig.

Kato, M. 2004. Taxonomic studies of Podostemaceae of Thailand. 1.

Hydrobryum and related genera with crustaceous roots (subfamily

Podostemoideae). Acta Phytotaxonomica et Geobotanica 55: 133-165.

Kato, M. 2006. Taxonomic studies of Podostemaceae of Thailand. 2.

Subfamily Tristichoideae and subfamily Podostemoideae with ribbon-

like roots. Acta Phytotaxonomica et Geobotanica 57: 1-54.

Kato, M., and Y. Kita. 2003. Taxonomic study of Podostemaceae of China.

Acta Phytotaxonomica et Geobotanica 54: 87-97.

Koi, S., Y. Kita, and M. Kato. 2008. Paracladopus chanthaburiensis sp. nov.

(Podostemaceae) from Thailand, with notes on its morphology, phylogeny

and distribution. Zaxon 57: 201-210.

Mathew, C.J. and V.K. Satheesh. 1997. Taxonomy and distribution of the

Podostemaceae in Kerala, India. Aquatic Botany 57: 243-274.

van Royen, P. 1965. Studies in the Flora of Thailand 29: Podostemaceae.

Dansk Botanisk Arkiv 23: 183-185.

Gardens’ Bulletin Singapore 61 (1): 73-79. 2009 73

Three New Species of Gesneriaceae from Kelantan,

Malaysia

R. KIEW

Forest Research Institute Malaysia,

52109 Kepong, Malaysia

Abstract

During a botanical expedition in 2007 to the Gunung Tera area in Kelantan,

NE Peninsular Malaysia, three new species of Gesneriaceae were discovered:

Henckelia kelantanensis Kiew, H. pauziana Kiew, and Ridleyandra

kelantanensis Kiew.

Introduction

A botanical expedition to the little known Gunung Tera area in Kelantan,

the northeastern state of Peninsular Malaysia, was organised in February

2007 as part of the Flora of Peninsular Malaysia Project. The area lies within

the Stong State Forest Park and includes several small peaks of which the

highest is Gunung Tera (521'N 101 54'E) at 1556 m. The terrain is dissected

by steep-sided valleys with torrential rocky streams. Granite cliff faces are

common in the area. During the expedition 332 species of vascular plants

were collected, of which 21% were species endemic to Peninsular Malaysia

and 16% were new records for the state of Kelantan (Chew et al., 2009).

Gesneriaceae was well represented with 16 species, only Orchidaceae (29

species), Rubiaceae (27 species), Gramineae (22 species) and Zingiberaceae

(17 species) were more speciose. Three species of Gesneriaceae proved new

to science and are described below.

1. Henckelia kelantanensis Kiew, sp. nov.

Ab Henckelia nivea (Kiew) A. Weber floribus majoribus (30 mm longis et

distaliter 15 mm latis nec 21 mm longis et 5 mm latis), paucinervis (paribus

17-18 nec 20-31) et capsulis longioribus (5-8 cm nec 2-3.5 cm longis) differt.

— Typus: Peninsular Malaysia, Kelantan, Gunung Stong, 18 May 1988, Kiew

RK 2736 (holotype, SING: isotype, KEP).

Herb, unbranched or sometimes with a short branch. Indumentum of

uniseriate, multicelluar, eglandular hairs, on the young stem and lower

lamina surface and on veins dense and ca 1 mm long, on upper lamina

surface and calyx ca 0.5 mm long; uniseriate, multicelluar, glandular hairs on

74 Gard. Bull. Singapore 61 (1) 2009

the pedicels dense and ca 0.5 mm long, on the outside of the corolla sparse

and ca 0.5 mm long, and on the ovary and capsule dense and ca 0.5 mm long.

Stem woody, 30-50 cm tall, 3-4 mm diam. Leaves alternate, internodes 1-1.5

cm long, lamina softly hairy, pale beneath, narrowly lanceolate or slightly

oblanceolate, 12-15.75 x 2.75-4 cm, base narrowed ultimately to a narrow

deeply serrate wing or sometimes with a petiole ca 6-8 mm long, margin finely

serrate, apex narrowly acute, lateral veins 17-18 pairs. Flowers epiphyllous,

solitary or in a series of 1-3 flowers, on the midrib and displaced 1-4 mm

from the leaf axil; pedicels 5.5-7.75 mm long; calyx divided almost to base,

lobes 5, narrowly acute, 2.5-3 mm long; corolla white with 2 yellow nectar

guides in the throat, gibbous, ca 3 cm long, base narrowly tubular ca 1.5 cm

long and ca 2 mm wide, distal ca 1.5 cm dilating to ca 1 cm wide across the

mouth, lobes 5, rounded, upper two erect, ca 7 mm diam., lower 3 spreading,

7-8 x 7-9 mm; ovary purple, slender, narrowing towards the stigma, ca 2.5

cm long, stigma peltate, discoid, ca 1 mm diam. Capsules slender, curved

downwards, 5-8 cm long, 1.5 mm thick.

Habitat: Hill dipterocarp forest, 740-800 m altitude, very common on banks,

slopes and the base of granite rock faces.

Additional specimen examined: Kelantan, Gunung Stong Tengah Forest

Reserve, Batu Hampar, Yao et al. FRI 55768 (KEP).

Distribution: Endemic in Peninsular Malaysia, known only from the Gunung

Stong area in Kelantan.

Notes: Henckelia kelantanensis belongs to a group of species [H. corneri

(Kiew) A.Weber, H. lilacina (Ridl.) A.Weber and H. nivea (Kiew) A.Weber]

that have soft, distantly spaced, alternate leaves and epiphyllous flowers.

Among these, it most resembles H. nivea in its large, gibbous, white corolla.

However, it is distinct from this species in its larger flowers (21 mm long and

5 mm wide in H. nivea), fewer-veined leaves (20-31 pairs in H. nivea) and

longer capsules (2-3.5 cm long in H. nivea).

2. Henckelia pauziana Kiew, sp. nov.

Ab Henckelia craspedodroma (Kiew) A. Weber foliis paucinervis (paribus

21-29 nec 39-51) et floribus infundibularibus majoribus 25-30 mm longis

orificio, 10-15 mm lato (nec tubularibus, ca 12 mm longis orificio, ca 5 mm

lato) differt. -Typus: Kelantan, Stong Tengah Forest Reserve, Batu Hampar

to Cobra Camp (5° 20.03'N, 101° 55.14'E), 8 Feb 2007, Chew et al. FRI 53513

[holotype, KEP (flower): isotypes KEP (fruits), K,L,SAN,SING]. Fig. 1.

=s. Inflorescences

—

Sh)

}

im]

: u

)

iF)

fi \

)

. i |

' ey

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76 Gard. Bull. Singapore 61 (1) 2009

with 2 conspicuous yellow nectar guides, 2.5-3 cm long, basal 0.7-1 cm

narrowly tubular and c. 2 mm wide, above dilating to 1-1.5 cm wide, lobes

5, upper two erect, rounded, ca 6-7 mm diam., lower 3 spreading, broadly

oblong with a rounded apex, ca 6-8 x 5 mm; nectary forming a broad ring

around the base of the ovary, ca 0.5 mm high; ovary slender, to 15 mm long,

style thread-like 8-10 mm long, stigma peltate, ca 0.5 mm diam. Capsules

slender, straight, 3-5 cm long, ca 1 mm thick.

Habitat: Upper hill dipterocarp forest at ca 895 m altitude, locally common,

on a steep slope with accumulation of leaf litter.

Distribution: Endemic in Peninsular Malaysia, known only from the type

locality.

Notes: Henckelia pauziana resembles H. craspedodroma (Kiew) A.Weber

in its foliage with short petioles and relatively narrow, glossy laminas with

parallel, deeply impressed lateral veins that end between the teeth of the

serrate margin and dense shaggy hairs along the upper surface of the midrib.

However, it is distinct from this species by a combination of characters

shown in Table 1.

Table 1. Differences between Henckelia pauziana and H. craspedodroma

Character Henckelia pauziana Henckelia craspedodroma

Stem ht (cm) to 12 to 100

Lateral veins (pairs) 21-29 39-51

Teeth (mm) 3-4 x 2-3 1-2 x 2-4

Bract shape foliose ligulate

length (mm) 6-8 3-3.5

No. flowers l l-several

Corolla shape funnel-shaped tubular

colour pale lilac pale violet cream

length (mm) 25-30 Gan

width at mouth (mm) 10-15 ca 5

This species is named in honour of Pauzi Husin, eco-guide whose

intimate knowledge of the Stong State Forest Park led to the discovery of

several new species, including this one.

3. Ridleyandra kelantanensis Kiew, sp. nov.

Ab Ridleyandra longisepala (Ridl.) A.Weber laminis foliorum brevioribus

Three New Species of Gesneriaceae from Kelantan

usque 14 x 4.5 cm (nec 15-30 x 4-7 cm), pedunculis usque 7 cm longis (nec

8-12 cm), sepalis usque 5 mm longis (nec 10-15 mm) et corollis ca 3 cm

longis (nec usque 5 cm) differt. — Typus: Malaysia, Kelantan, Stong Tengah

Forest Reserve, path to Camp Cobra (5° 20.07'N, 101° 55.16'E), 8 Feb 2007,

Chew et al. FRI 53518 (holotype, KEP; iso E,L). Fig. 2.

Figure 2. Ridleyandra kelantanensis Kiew, Gesneriaceae (photo: Chew M.-Y.).

Unbranched herb. Stem woody, becoming decumbent, erect portion 15-19(-

30) cm tall, ca 4 mm diam. Indumentum of reddish brown, dense, uniseriate,

multicellular hairs, ca 0.5 mm long, dense and hispid on upper portion

of stem, petioles, lower surface of midrib, peduncles, pedicels and sepals.

Leaves in pairs, clustered in a rosette at the top of the stem, above up to 3-4

mm apart, below to 4.5 cm apart; petioles 1-1.75 cm long; lamina mid-green

above, slightly paler beneath, glabrous, slightly bullate, oblanceolate, 8.5-14

x 3-4.5 cm, narrowed to base, margin dentate, teeth falcate, 4-5 x 3-5 mm

and 2-6 mm apart, apex acute; midrib and veins slightly impressed above,

prominent beneath, lateral veins 14-20 pairs. Flowers solitary; peduncles,

pedicels and sepals maroon and covered in fine white hairs, peduncles 4.75-

7 cm long; bract pair broadly lanceolate, 2-3 x 1-1.5 mm, pedicels (4-) 7-8

mm; sepals free, lanceolate, ca 5 x 3 mm, tip acute; corolla funnel-shaped,

tube ca 2 cm long, ca 4 mm wide at base, dilating to 10-12 mm wide at the

mouth, upper lobes deep blue and recurved, rounded, ca 10-11 x 9-10 mm,

78 Gard. Bull. Singapore 61 (1) 2009

lower lobes white with 3 major and a few minor blue stripes, oblong, 8-12 x

9-10 mm, apex rounded; stamens 4 in 2 pairs; ovary 1.75-2 cm long, stigma

slightly bilobed, 2 mm across. Capsules slightly curved downwards, glabrous,

4-4.5 cm long, ca 1.5 mm thick; sepals caducous.

Habitat: Locally common in upper hill dipterocarp forest on steep banks

with peaty soil at 900-1000 m altitude.

Distribution: Endemic in Peninsular Malaysia, Kelantan, known only from

the type collections.

Notes: In flower colour and pattern it resembles Ridleyandra longisepala

(Ridl.) A.Weber but it differs in its overall smaller size: R. longisepala grows

to 1 m tall, has laminae 15-30 x 4-7 cm long, peduncles 8-12 cm long, sepals

10-15 mm long and a corolla ca 5 cm long. It is also different in its leaf margin

(R. longisepala has a margin that is doubly serrate) and in the presence of

a distinct pedicel (absent in R. longisepala). Weber (1998, ‘1997’) used this

last as a key character in separating Ridleyandra species, that is, whether the

bracts were immediately below the sepals (pedicel not discernable) or were

some distant below the sepals (1.e., the flower has a distinct pedicel).

The resemblance of the corolla markings to those of Ridleyandra

longisepala is remarkable. Weber has drawn attention to several unrelated

species of Ridleyandra having the same corolla coloration, namely deep

purple with an almost black throat, and suggested that this be due to

having the same pollinator. This is likely to be the case with Ridleyandra

kelantanensis and R. longisepala although, as in the case of all Ridleyandra

species, the pollinators remain unknown.

The two species also have different habitats and distributions:

Ridleyandra longisepala, is known only from the Main Range at Cameron

Highlands, Pahang, and grows in upper montane forest at 1700 m, whereas

R. kelantanensis grows in upper hill dipterocarp forest to 1000 m.

Acknowledgements

The expedition was funded by the Ministry of Science, Technology and

Innovation under the Flora of Peninsular Malaysia Project No. 01-04-01-

000 Khas at Forest Research Institute Malaysia (FRIM). The author is

indebted to the State Forestry Department of Kelantan for permission to

make botanical collections; to the FRIM expedition team (Kamarudin

Saleh, Chew M.-Y., Yao T.L., Mustafa Data and Angan Atan) and to Pauzi

Husin and the Baha Adventure Team whose help in the field contributed

in large measure to the success of the expedition; to the curators of the

Three New Species of Gesneriaceae from Kelantan

herbaria of the Royal Botanic Gardens at Edinburgh and Kew, and the

Singapore Botanic Gardens, for permission to examine specimens in their

care; and to Chew M.Y. for photographing the new species under difficult

field conditions and to M.J.E. Coode for correcting the botanical Latin.

References

Chew M.-Y., T.L. Yao, S. Kamarudin and R. Kiew. 2009. Exploration,

vegetation and a survey of plant life of the Gunung Tera Area, Kelantan,

Malaysia. Malaysia Nature Journal 61: 67-120.

Weber, A. 1998 (‘1997’). Revision of the genus Ridleyandra (Gesneriaceae).

Beitrage zur Biologie der Pflanzen 70: 225-273.

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Gardens’ Bulletin Singapore 61 (1): 81-93. 2009 $1

A Review of the Genus Plocoglottis (Orchidaceae)

in Thailand

H. KURZWEIL

Smegapore Botanic Gardens. 1 Cluny Rd. Singapore 259569

E-mail: hubert_kurzweil@nparks sovss

Abstract

A review of the genus Plocogloiis m Thailand ts presented. Five species are

recognised. Four of them. P. javanica, P. low, P. quadrifolia and P. gigantea.

are widespread in Malesia and have their northernmost limit of distribution

m Penmsular Thailand. and P javanica is also known im Vietnam. The

fifth. P bokorensis is also found in Cambodia. Laos and Vietnam and is in

Thailand known from a few widely scattered localities in the central. eastern.

southeastern. southwestern and northem regions.

Introduction

Plocoglottis Blume is a terrestrial orchid genus of 40 to 45 species and is

centred in Malesia. ranging from the Andaman Islands. southern Myanmar.

Thailand. Cambodia and Vietnam eastwards as far as New Guinea. Bismarck

Archipelago and the Solomon Islands. Plants of this genus are found in

lowland_ hill and lower montane forest up to about 1200 m. Based on all of

the available evidence (including morphological. anatomical and molecular

data) the genus was placed in tribe Collabieae of subfamily Epidendroideae.

although it was pointed out that further studies on this tribe are needed

(Pridgeon et al.. 2005: 116).

Plocoglotts is represented in Thailand by five uncommon species. In

the first detailed treatment of the orchids of Thailand [see Seidenfaden and

Smitinand 1959-1965: The Orchids of Thailand - A Preliminary List]. only

two species were listed. namely P. javanica Blume and P. quadrifolia IJ. Sm.

(incorrectly identified as Phaius longipes (Hook-£) Holttum). In the years

following this initial publication a further three species were added, and in a

brief overview of the Thai species of the genus (Seidenfaden. 1972). all five

species that are currently accepted were already listed.

A detailed review of the older literature and a discussion of the

taxonomic history of all of the species is provided m a later detailed treatment

(Seidenfaden. 1986). Though mvaluable to the professional botanist as

well as the orchid-enthusiast, Seidenfaden's contributions did not include

82 Gard. Bull. Singapore 61 (1) 2009

taxonomic descriptions which are provided in the present review.

Four of the Thai species, P. javanica, P. lowii Rchb. f., P. quadrifolia

and P. gigantea (Hook. f.) J.J. Sm. are widespread in the western part of

Malesia (with P. /owii even ranging further eastwards to New Guinea in

eastern Malesia), and have the northernmost limit of their distribution area

in Peninsular Thailand and Vietnam (P. javanica only). The distribution area

of the fifth Thai species, P bokorensis (Gagnep.) Seidenf., is very different.

This species is only found in Thailand and Indochina, and while widespread

(but widely scattered) in the northern half of Thailand it does not seem to

occur in the Peninsula of Thailand.

The plants of the genus Plocoglottis are sympodial herbs with or

without pseudobulbs, and arise from short creeping rhizomes. Two different

plant habits can be distinguished. In P. javanica and P. lowii the plants have

narrowly-fusiform pseudobulbs with one apical leaf (rarely two), while the

other species have elongate reed-like stems with several leaves. The erect

or spreading leaves are generally plicate and have often long petioles.

Inflorescences are lateral and arise at the base of the leafy stem. They are

usually as long as to clearly longer than the leafy shoot. The medium-sized

flowers have free sepals and petals, and a fleshy lip with an interesting

explosive mechanism. This lip is united with the gynostemium base by two

elastic flanges and snaps upwards when touched.

The present review is a precursory paper for a later contribution

to the Flora of Thailand. No attempt has been made to provide a complete

monographic treatment as very little material from outside of Thailand has

been seen.

Key to Plocoglottis in Thailand

1. Plant with one leaf (rarely two leaves) on top of a narrowly-fusiform

PSCUGODUID 2.0... ciccsesdacesntsscctcoe cccecueeeeecenece decee eee ee 2

1. Stems reed-like with four or more leaves 32. eee 3

Petioles mostly 20-40 cm long. Side- margins of the lip entire ...................

wabs'soeuancaustondecdsteadvacteabesensdijecstce ress uce Stee Ret ERZUES Ae Caan nS e 1. PR. javanica

2. Petioles shorter. Lip side margins erose-serrate, two conical calli near

the lip front margin ...1.5.20.5. ee eee ee ree 2.P. lowii

3. Lip side margins fimbriate Or TOS .............:eseeeeeeeeeeeeeee 0 Quadrifolia

3. Lap side miareims' Smite 2.06.52. cec. te icee eaten tteec ence ateacteceacesu cee enceececenee aan -

4. Stems up to | m tall, without pseudobulbs. Sides of lip much recurved, lip

broadest near the middle 7... 22a see cee 4. P. gigantea

: = ; 9

A Review of the Genus Plocoglottis (Orchidaceae) in Thailand roy)

4. Stems up to 50 cm tall, with prominent bottle-shaped basal pseudobulbs

to 8cm tall. Sides of lip not much recurved, lip broadest near the apex........

ee ee ee eS 5 E98 ak 5 Eg Scczcbsact eds acgedceeseades--.025 ed DOKOFENSIS

1. Plocoglottis javanica Blume

Bijdr. (1825) 381; Seidenfaden & Smitinand, Orch. Thailand (1961) 348;

Seidenfaden, Bot. Tidsskr.67 (1972) 108; Opera Bot.89 (1986) 69:Seidenfaden

& Wood, Orch. Penins. Malaysia & Singapore (1992) 181; Comber, Orch.

Java (1990) 104; Orch. Sumatra (2001) 292. -Types: Indonesia, Java, Seribu,

Blume s.n. (syn, L); Indonesia, Java, Mt. Salak, Blume s.n. (syn, L). Fig. 1.

Figure 1. Plocoglottis javanica Blume with solitary leaf (photo taken in Singapore). A. Plant:

B. Flower. Photo: H. Kurzweil (HK 2774 voucher in SING).

Terrestrial herbs, evergreen, pseudobulbous. Pseudobulbs narrowly conical,

terete, 4-10(-12) x 0.5-1 cm, ca 4 cm apart on a creeping rhizome to | cm

thick, enclosed by fibrous leaf sheaths. Leaf 1,on the apex of the pseudobulb,

elliptic-lanceolate, acute or acuminate, 46-61 x 6.8-9.4(-11.1) cm, plicate,

glabrous; with petioles (13.5-)20-40 cm long. Inflorescences 46-70(-81) cm

long, arising from the base of the leafy shoot, usually extending beyond the

leaves, glabrous below, pubescent or rarely glabrous above; peduncles to 50

cm long; sterile bracts 3-5, lanceolate-oblong, tubular, acute, 1.5-3 cm long,

subglabrous, two or three of them clustered at the stem base and the rest

scattered higher up; raceme lax or semi-dense, 10- to 15-flowered; rachis

84 Gard. Bull. Singapore 61 (1) 2009

gradually elongating to 30 cm long; floral bracts ovate-lanceolate, acute,

much shorter than the pedicel plus the ovary, 6.5-10 x 2-2.3 mm. Flowers

to 17 mm broad; yellow, blotched reddish, also reported as brown, lip

largely yellow. Pedicel plus ovary ca 15 mm long, glabrous or pubescent.

Sepals ovate-lanceolate, shortly acuminate, spreading, concave; median

sepal 5-veined, 10-15 x 4-6 mm; lateral sepals 7-veined, 13-15 =x 2-5 mm.

Petals linear-falcate, acute, 5-veined, 10-15 x 2.2-2.9 mm, spreading, their

tips curved forwards. Lip ovate-lanceolate, shallowly 3-lobed in the apical

part, 8-9 x 7-8 mm, basally joined with the gynostemium, margins entire;

midlobe an acute tip 0.5-2.5 mm long; side-lobes flabellate, obtuse, ca 2 x 3.5

mm. Gynostemium straight, stout, 5.5-6.2 mm long, apically broadly winged,

stigma cavity large.

Specimens examined: Peninsular Thailand, Pattani Province, Kao Kalakiri,

Kerr 0554 (K [fragment]); ibid., Songkhla Province, Klawng Pi, Kerr 0614

(BK, C, K); ibid., Yala Province, Betong, Maknoi 130 (QBG).

Illustrations: Seidenfaden (1986): fig. 34a-d; Seidenfaden & Wood (1992):

fig. 78a; Comber (1990): pl. on page 104; Comber (2001): pl. on page 292;

Pridgeon et al. (2005): pl. 45.

Habitat and flowering time: Rare in evergreen forest up to about 700 m.

Flowering in April and from July to August.

Distribution: Peninsular Thailand (3 collections seen); also in Vietnam, the

Andaman Islands, Malaya, Java, Sumatra and Borneo.

Notes: This species is similar to the much more common Plocoglottis lowit.

Differentiating characters of the two species are the lip side margins which

are entire in P. javanica but erose or fimbriate in P. /owii. The length of the

petiole of the leaves can in most cases also be used to distinguish between

the two species as P. javanica has usually an elongate petiole of (13.5-)20-

40 cm while the leaf petiole of P. /owii is usually much shorter (up to 14

cm). Also the hairiness of the rachis has been used to differentiate between

the two species (Seidenfaden, 1986: 69), but was here found not to be very

significant as a few exceptions occur.

2. Plocoglottis lowii Rchb.f.

Gard. Chron. (1865) 434; Seidenfaden, Bot. Tidsskr. 67 (1972) 109; Opera

Bot. 89 (1986) 70; Seidenfaden & Wood, Orch. Penins. Malaysia & Singapore

(1992) 181; Comber, Orch. Sumatra (2001) 292; Beaman & al., Orch. Sarawak

(2001) 383. — Type: Borneo, without locality, Low s.n. (W).

A Review of the Genus Plocoglottis (Orchidaceae) in Thailand 8

-Plocoglottis porphyrophylla Ridl., Trans. Linn. Soc. London, Bot. 3 (1893)

368. -Type: Peninsular Malaysia, Pahang, Pekan, Ridley 394 (holo, SING!).

Terrestrial herbs, evergreen, pseudobulbous. Pseudobulbs narrowly conical,

green to dark violet, 5-8(-15) x 1.1-1.5(-2) cm, ca 3 cm apart on a creeping

rhizome; enclosed by fibrous leaf sheaths. Leaves 1(-2)(-3?, see ‘Notes’

below), on the apex of the pseudobulb, elliptic-lanceolate, acute, 30-59 x 4.5-

12 cm, plicate, glabrous, green or purple above, purple underneath; petioles

(3-)6-14 cm long. Inflorescences 62-121 cm long, arising from the base of

the leafy shoot, much taller than the leaves, pubescent, violet, once here

observed with a side-branch; peduncles 48-84 cm long; sterile bracts 5-8,

scattered along the flowering stem and a few clustered at its base, lanceolate-

oblong, lower tubular, upper spreading, acute, (1.2-)1.6-4.5 cm long, glabrous

or pubescent; raceme lax, 20- to 30-flowered, producing flowers for many

weeks with 1-2 open at a time; rachis 14-36.5 cm long; floral bracts ovate-

lanceolate, acuminate, green-violet, 6-11.7 x 1.8-6 mm, much shorter than the

pedicel plus the ovary, pubescent. Flowers to 30 mm in diameter; yellowish

and greenish, with red or brown lines and blotches, bulging area on the inside

of the lateral sepals mostly shiny red, petals also reported as reddish purple,

lip epichile sometimes red, gynostemium light yellow; all parts except lip

pubescent on both sides though gynostemium usually sparsely so. Pedicel

and ovary 10-15 mm long, pubescent. Sepals oblong-lanceolate, subacute or

acuminate, spreading; median sepal 7-veined, 10-16(-17) x 4-5.6 mm; lateral

sepals 9-13 x (4.9-)5-7 mm, deflexed, tips inwards-curved, with a more or less

pronounced bulging area on the sides that face each other. Petals narrowly

triangular-lanceolate, spreading, gradually narrowed towards the tip, acute,

5- or 7-veined, 9.5-16.3 x 2-4.5 mm. Lip ovate, 3-lobed in its upper part, (5-

)5.5-8 x 5-8.3 mm, spreading, united with the gynostemium at the base, with

two pronounced apical conical calli, margins fimbriate or erose; midlobe

an acute tip, (1-)1.3-2.5 x 0.7-1.3 mm; side-lobes triangular, obtuse, 1.5-1.8

x 1.5-2 mm. Gynostemium 4.5-8 mm long, straight, stout, pubescent. Fruit

elongate-ellipsoid, 22-40 x 3-8 mm, pubescent.

Illustrations: Seidenfaden (1972): fig. 27a-c; Seidenfaden (1986): fig. 35a-d;

Seidenfaden & Wood (1992): figs. 78b-e; Comber (2001): pl. on page 292:

Beaman et al. (2001): pl. 28D.

Habitat and flowering time: In evergreen forest, freshwater swamp forest,

bamboo forest or sometimes in swampy grassland to 300 m elev. The few

detailed collector's notes indicate limestone as bedrock. Flowering in April,

June-July and October-February.

86 Gard. Bull. Singapore 61 (1) 2009

Specimens examined: Peninsular Thailand, Chumphon Province, Langsuan,

Kerr 0652 (C, C [spirit], K); ibid., Surat Thani Province, Chaiya District,

Maxwell 87-151 (L);ibid., Krabi Province, Nivomdham 1746 (C spirit]); ibid.,

Krabi Province, Bang Kram Forest Reserve, Larsen et al. 43403 (AAU);

ibid., Krabi Province, Bang Kram Forest Reserve, Larsen & al. 43782 (C

[spirit]); ibid., Krabi Province, Kow Pra-Bahng Krahm Wildlife Sanctuary,

Maxwell 06-38 (Sc.ChM. Univ. = Herbarium of the Science Faculty of Chiang

Mai University, Sc.ChM.Univ. [spirit]); ibid., Krabi Province, Thaithong

1794 (BCU [spirit]); ibid., Krabi Province, Khao Nop Ju Jee, Thorut 245

(BCU [spirit]); ibid., Nakhon Si Thammarat Province, Williams et al. 1361

(BKF [spirit]); ibid., Trang Province, Thung Khai, Larsen & al. 43982 (AAU,

C [spirit], SING) [in bud, identification uncertain]; ibid., Trang Province,

Thung Khai, Larsen & al. 43985 (AAU); ibid., Trang Province, Thung Khai

Botanical Garden [cultivated ?], Mauric 5 (BKF); ibid., Trang Province,

Tung Kai, QBG 10003 (QBG [spirit]); ibid., Trang Province, Thung Khai,

Suddee 391 (BKF [s]); ibid., Songkhla Province, Khao Keo, Kerr 0627 (K);

Tako, Langsuan, Put 1753 (K).

Distribution: Peninsular Thailand (16 collections seen); also in the Andaman

Islands, Malaya, Sumatra, Borneo, Moluccas and New Guinea.

Notes: This species is very distinct with its mostly solitary short-petioled

leaf and the erose or fimbriate side margins of the lip. This and further

distinguishing characters are also listed under the preceding species. The Thai

plants listed here clearly match Reichenbach's (1865) original description.

The bulging area on the lateral sepals referred to above is, however, subject

to variation in the species.

A specimen from Trang (Larsen & al. 43982, AAU, C [spirit], SING)

was collected in the bud stage, and was interpreted as a representative

of a new species because of its unusual habit with three leaves on top of

the pseudobulb (G. Seidenfaden, notes in his working files). While it will

probably never be possible to identify the plant accurately due to the

absence of flowers, it appears that the habit is not really unique as specimens

with two leaves occur occasionally (though rarely) in this species. A flower

bud that Seidenfaden had opened was found in the spirit collection at C, and

exhibits the fringed side margins of the lip typical of this species. It therefore

appears likely that the specimen is merely a slightly unusual representative

of Plocoglottis lowii.

3. Plocoglottis quadrifolia J.J.Sm.

Bull. Jard. Bot. Buitenzorg III: 8 (1926) 36; Seidenfaden, Bot. Tidsskr. 67

(1972) 109; Opera Bot. 89 (1986) 70; Seidenfaden & Wood, Orch. Penins.

A Review of the Genus Plocoglottis (Orchidaceae) in Thailand 87

Malaysia & Singapore (1992) 181; Comber, Orch. Sumatra (2001) 290; N.

Vaddhanaphuti, Wild Orch. Thailand (2005) 208. — Type: Indonesia, Sumatra,

Agam, Boekit Batoe Banting, Groeneveldt 873 (holo, L!).

-Phaius longipes auct., non (Hook.f.) Holtt.; Seidenfaden & Smitinand,

Orch. Thailand (1961) 329.

Terrestrial herbs, evergreen, stems reed-like, 30-40 cm tall, with leaves in the

upper half, lower half with sheathing leaves. Leaves 5-7, lanceolate-elliptic,

acute, acuminate, blades 11.5-21 x (1.6-)2-3.5 cm, plicate, glabrous, petiole to

1.5 cm long; leaves generally green with some purple. Inflorescences 67-92

cm tall, arising from the base of the leafy shoot, much taller than the leafy

shoots, softly-pubescent; peduncles to 58 cm long; sterile bracts 7-8, scattered

along the flowering stem and a few clustered at its base, lanceolate-oblong,

acute, 1.6-5 cm long, glabrous or hairy, sheathing at the base and their

upper part spreading; raceme lax, 10- to 25-flowered; rachis 14-37 cm long,

pubescent; floral bracts ovate-lanceolate, acuminate, 8-10 x 2-4 mm, mostly

much shorter than the pedicel plus the ovary, sometimes yellowish purple.

Flowers to 29 mm in diameter; mostly yellow with a red tinge, lateral sepals

distally yellow and basally shiny red, lip yellow with red markings, apical half

often pink, violet or red, gynostemium yellow or white. Pedicel and ovary

7-18 mm long, densely pubescent. Sepals ovate-lanceolate, subacute, 7- or

9-veined, spreading, hairy on the outside; median sepal 18-25 x 5.2-8 mm;

lateral sepals 13-22 x 5-10 mm, with a shiny bulging area on the sides that

face each other. Petals very narrowly ovate-lanceolate, subacute, spreading,

3- to 7-veined, 18-25 x 3-5 mm, glabrous. Lip quadrangular but narrowed

towards the base, 3-lobed in the apical half, 8-12 x 8-12 mm, united with the

gynostemium at the base, glabrous, margins fimbriate; midlobe a reflexed

apical tooth, 0.95-2 x 1.2-2 mm; sidelobes triangular or square, obtuse, 2-5

x 3.5-5 mm, with obscure or prominent apical calli. Gynostemium straight,

9-11 mm long, slender, rostellum a short blunt beak.

Specimens examined: Peninsular Thailand, Surat Thani Province, Koh

Pa-ngan, Put 756 (B, BK, C, C [spirit], K, P); ibid., Nakhon Si Thammarat

Province, Khao Luang, Geesinck & Santisuk 5483 (C, C [spirit], L); without

locality, Thaithong 1374 (BCU [spirit]).

Illustrations: Seidenfaden & Smitinand (1961): fig. 252 [as Phaius longipes

(Hook.f.) Holtt.]; Seidenfaden (1986): fig. 36a-e; Seidenfaden & Wood

(1992): fig. 78f; Comber (2001): pl. on page 290; N. Vaddhanaphuti (2005):

pl. on page 208.

Habitat and flowering time: Found in rock crevices in disturbed evergreen

88 Gard. Bull. Singapore 61 (1) 2009

forest, often along waterfalls. The plants are usually found in lower hill

forests up to 500 m. Flowering in May.

Distribution: Peninsular Thailand (3 collections seen), also in Malaya and

Sumatra.

Notes: Characteristic features of this species are the reed-like stems with

several leaves, the median sepals which are about 18-25 mm long and the

fimbriate lip side margins. As already pointed out by Smith (1926: 37), the

species is close to Plocoglottis lowii but differs from it vegetatively.

4. Plocoglottis gigantea (Hook.f.) J.J.Sm.

Repert. Spec. Nov. Regni Veg. 32 (1933) 228; Seidenfaden, Bot. Tidsskr.

67 (1972) 109; Opera Bot. 89 (1986) 70; Seidenfaden & Wood, Orch.

Penins. Malaysia & Singapore (1992) 183; Comber, Orch. Sumatra (2001)

291. -Alismorkis gigantea (Hook.f.) Kuntze, Revis. Gen. Pl. 2 (1891) 650.

-Basionym: Calanthe gigantea Hook.f., Fl. Brit. India 5 (1896) 856. — Types:

Peninsular Malaysia, Perak, Kings collector 10277 (syn, K!; isosyn, BM);

Peninsular Malaysia, Perak, Wray 2932 (syn, K!).

-Plocoglottis foetida Ridl., J. Linn. Soc., Bot. 32 (1896) 319. —Type: not

designated.

Terrestrial herbs, evergreen, stems reed-like, to 100 cm tall, with leaves in its

upper half, lower half with sheathing leaves only, stems pubescent. Leaves

6-12, lanceolate, acuminate, blades 19-30 x (1.9-)4.2-7.3 cm, plicate, shortly

hairy, sheathing at the base, sessile. Inflorescences erect, to over | m tall,

arising from the base of the leafy shoot, slightly longer or shorter than the

leafy stem, pubescent; sterile bracts 7-9, tubular, lanceolate-oblong, acute,

2.3-2.5 cm long, glabrous or sparsely pubescent; raceme lax, many-flowered;

rachis 18-23 cm long; floral bracts triangular, acuminate, acute, 10-15 x 8-9

mm, shorter than the pedicel plus the ovary, pubescent. Flowers to 38 mm

long, yellow blotched reddish, lip cream or white and turning yellow as the

flower ages; sepals and petals fleshy, densely pubescent on the outside and

sparsely so on the inside; flowers reported to have a foetid smell. Pedicel and

ovary 18-20 mm long, densely pubescent. Sepals lanceolate, acute, 20-23 x

8-10 mm, pubescent; median sepal spreading; lateral sepals deflexed. Petals

similar to the petals, slightly smaller, facing forwards. Lip elliptic, shallowly

3-lobed in its upper part, 9-10 mm long, united with the gynostemium at the

base, concave, sides much recurved, lip broadest near the middle; midlobe

a triangular tooth, deflexed, to 3 x 2 mm long; side-lobes obscure, obtuse, to

2 mm long. Gynostemium 6-9 mm long, straight, stout. Fruit ellipsoid, ca 5.2

x 1.4 cm.

A Review of the Genus Plocoglottis (Orchidaceae) in Thailand 89

Specimen examined: Peninsular Thailand. Narathiwat Province, Hala-Bala

Forest Reserve, Nivomdham 7189A (BKF [spirit]).

Illustrations: Seidenfaden (1986): fig. 37a-c; Seidenfaden & Wood (1992): fig.

78g-h.

Habitat and flowering time: The habitat of the single Thai collection has not

been recorded. However, in Malaya the plants are found in moist places in

lowland forest and flower in February.

Distribution: Peninsular Thailand (1 collection seen), also in Peninsular

Malaysia, Borneo and Sumatra.

Notes: Plocoglottis gigantea is well-characterised by its tall stems which

lack pseudobulbs at the base. The hairy flowers have a concave lip, which is

broadest near the middle and has strongly recurved side margins.

This species is in Thailand known only from a single collection made

in the Province of Narathiwat in the extreme south of the country in 2004.

The species was originally included in the list of the Thai Plocoglottis species

(Seidenfaden, 1972: 109; 1986: 70, fig. 37a-c) because of a literature reference

made by Haniff (1916: 353) who recorded the species for "Setol" (= Satun).

Haniff was, however, not sure about the identity of his plant.

The flowers have been reported to have a foetid smell, which

Ridley based his species name Plocoglottis foetida on (which is considered

synonymous with P. gigantea).

5. Plocoglottis bokorensis (Gagnep.) Seidenf.

Dansk Bot. Ark. 33 (1979) 219; Opera Bot. 89 (1986) 71; Opera Bot. 114

(1992) 105. —Basionym: Bulbophyllum bokorense Gagnep., Bull. Mus. Natl.

Hist. Nat. II: 22 (1950) 399. — Type: Cambodia, Bokor, Poreé-Maspero s.n.

(P). Fig. 2.

-Plocoglottis mirabilis Seident., Bot. Tidsskr. 65 (1970) 336; Bot. Tidsskr. 67

(1972) 109. —Type: Thailand, Khao Yai National Park, 1300 m, Cumberlege

926 (C!).

Terrestrial herbs, evergreen, stems reed-like, 40-50 cm tall, with several

leaves in its upper half, lower half with sheathing leaves, arising from distinct

pseudobulbs. Pseudobulbs bottle-shaped to conical, 7-8 = 3-4 cm, with

internodes 3.5-4 cm, dark olive-green and becoming purplish brown when

old. Leaves (5-)8-10, elliptic-lanceolate, acute, without prominent petioles,

blades 8-16.5 x 1.9-3.5 cm, plicate, glabrous, basally sheathing, blades wavy.

Inflorescences 30-51 cm long, arising from the base of the pseudobulb, mostly

Gard. Bull. Singapore 61 (1) 2009

Figure 2. Plocoglottis bokorensis (Gagnep.) Seidenf. with reed-like stem (photo taken in

Khao Yai National Park, Thailand). A. Plant, note the prominent pseudobulbs (arrowhead);

B. Inflorescence; C. Flower. Photo: S. Chantaranorrapint (Buakhlai 70 voucher in BCU).

A Review of the Genus Plocoglottis (Orchidaceae) in Thailand 9]

slightly taller than the leafy stem, pubescent; sterile bracts 5-8, scattered

along the flowering stem and a few clustered at its base, lanceolate-oblong,

tubular, acute, 2.7-4 cm long, glabrous; raceme lax, 7- to 15-flowered; rachis

(7-)13-20 cm long; floral bracts ovate-lanceolate, acuminate, 8-12 x 3-5.5

mm, shorter than the pedicel plus the ovary, pubescent, glossy whitish

green. Flowers pale pink or yellow with red blotches, lip uniformly yellow

or whitish-yellow with purple markings at the base. Pedicel and ovary 15-28

mm long, pubescent. Sepals oblong-lanceolate, navicular, subacute or acute,

5- or 7-veined, pubescent on the outside, glabrous and glossy on the inside,

spreading; median sepal hooded, 13-20 x 5-7 mm; lateral sepals similar but

slightly narrower, mucronate. Petals narrowly oblong, obtuse, 3- or 5-veined,

(11.5-)12.3-18.5 x 2.6-5 mm, glabrous. Lip ovate, shallowly 3-lobed, 7.5-11

x 8.5-12 mm, widest in its recurved upper part, sides not much recurved,

base with a distinct cavity formed by the flanges (which join the lip to the

gynostemium) and the bases of lip and gynostemium; midlobe triangular, 0.6-

1 x 0.5-1.5 mm; side-lobes triangular, obtuse, ca 3 x 2-2.5 mm. Gynostemium

4.5-7 mm long, glabrous, stigma triangular or heart-shaped.

Specimens examined: Northern Thailand, Nan Province, Doi Phu Kha

National Park, Srisanga 2584 (QBG); Eastern Thailand, Nakhon Ratchasima

Province, Khao Yai National Park, Cumberlege 926 (C [spirit]); Southwestern

Thailand, Phetchaburi Province, Kaeng Krachan National Park, Middleton

& al. 3291 (BKF [spirit]); Central Thailand, Nakhon Nayok Province, Khao

Yai National Park, Buakhlai 70 (BCU [spirit]); Southeastern Thailand, Trat

Province, Kao Kuap, Put 2978 (K).

Illustrations: Seidenfaden (1970): fig. 14a-e [as Plocoglottis mirabilis Seidenf. |;

Seidenfaden (1986): fig. 38a-e, pl. Vb; Seidenfaden (1992): figs. 56a-e.

Habitat and flowering time: The only available detailed collector's record

(Srisanga 2584, QBG!) indicates that the plants grew in dry evergreen

forest at an altitude of 800 to 1100 m. Plants in Khao Yai National Park

are reported to grow in a more humid environment. Flowering takes place

between May and July.

Distribution: In several widely scattered localities in northern, eastern,

southwestern, central and southeastern Thailand, also in Cambodia, Laos

and Vietnam.

Notes: This species is characterized by its reed-like stems which have

prominent pseudobulbs at the base. The lip shape is a distinguishing feature

from Plocoglottis gigantea, as it is generally widest in its apical part and has

92 Gard. Bull. Singapore 61 (1) 2009

entire and only little decurved side margins. The Thai plants were originally

described as P. mirabilis, which was later found to be conspecific with

Bulbophyllum bokorense Gagnep.

Plocoglottis bokorensis is interesting from a phytogeographical point

of view. It is known from several widely scattered localities in northern,

eastern, southwestern, central and south-eastern Thailand, and is therefore

the only Thai Plocoglottis species which is apparently absent from the

Peninsula.

Acknowledgements

I am grateful to the curators of the herbaria AAU, B, BK, BCU, BKF, C,

K, L, QBG, and the Science Faculty of Chiang Mai University for making

their collection available for study. Staff in Thai herbaria have helped in

many ways, particularly Dr. O. Thaithong and Miss S. Wongpakam (both

BCU), Dr. S. Suddee (BKF) and Dr. S. Watthana and Dr. P. Suksathan (both

QBG). Thanks to S. Chantaranorrapint for letting me use his photographs

of Plocoglottis bokorensis. 1 further thank the National Research Council of

Thailand for issuing a research permission.

References

Beaman, T-E., J.J. Wood, R.S. Beaman and J.H. Beaman. 2001. Orchids of

Sarawak. Natural History Publications, Kota Kinabalu.

Comber, J.B. 1990. Orchids of Java. Bentham-Moxon Trust, Royal Botanic

Gardens, Kew.

Comber, J.B. 2001. Orchids of Sumatra. Natural History Publications, Kota

Kinabalu.

Haniff, M. 1916. Record of a few orchids. Agricultural Bulletin of the Straits

and Federated Malay States 1: 353-355.

Pridgeon, A.M., P.J. Cribb, M.W. Chase and E.N. Rasmussen. 2005. Genera

Orchidacearum, vol. 4 (Epidendroideae, part one). Oxford University

Press.

Reichenbach, H.G. 1865. Plocoglottis lowii. Gardeners' Chronicle 1865: 434.

A Review of the Genus Plocoglottis (Orchidaceae) in Thailand 93

Seidenfaden, G. 1970. Contributions to the orchid flora of Thailand III.

Botanisk Tidsskrift 66: 303-356.

Seidenfaden, G. 1972. Contributions to the orchid flora of Thailand IV.

Botanisk Tidsskrift 67: 76-127.

Seidenfaden, G. 1986. Orchid genera in Thailand XIII. Thirty-three

epidendroid genera. Opera Botanica 89: 1-216.

Seidenfaden, G. 1992. The orchids of Indochina. Opera Botanica 114: 1-502.

Seidenfaden, G. and T. Smitinand. 1959-1965. The Orchids of Thailand — A

preliminary list. The Siam Society, Bangkok.

Seidenfaden, G. and J.J. Wood. 1992. The Orchids of Peninsular Malaysia

and Singapore. Olsen & Olsen, Fredensborg.

Smith, J.J. 1926. Orchidaceae Novae Malayenses, XI. Bulletin du Jardin

Botanique de Buitenzorg, ser. U1, 8: 35-70.

Vaddhanaphuti, Nantiya. 2005. A field guide to the wild orchids of Thailand,

ed. 4. Silkworm Books, Chiang Mai.

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Gardens’ Bulletin Singapore 61 (1): 95-99. 2009 9

‘nN

Nepenthes pitopangii (Nepenthaceae), a New Species from

Central Sulawesi, Indonesia

C.C.LEE ,S.R.MCPHERSON .G. BOURKE |

AND M. MANSUR-

: - Peti Surat 2507, Kuching, Sarawak 93750, Malaysia.

61 Lake Drive, Hamworthy, Poole, Dorset BH15 4LR, England.

* P.O. Box 117, Corrimal, New South Wales 2518, Australia.

: Botany Division, Research Center for Biology, Indonesian Institute of Sciences,

Cibinong Science Center, West Java, Indonesia.

Abstract

A new Nepenthes species, N. pitopangii, from Sulawesi Tengah, is described.

Introduction

Cheek and Jebb (2001) recognised eight species of Nepenthaceae for Sulawesi,

of which four species are endemic. These numbers are comparatively low

compared to the neighboring island of Borneo, where there are more

than 30 species, of which 70% are endemic. The lack of species richness in

Nepenthaceae in Sulawesi is yet to be examined in detail, but may be due to

a combination of biogeographical and ecological factors, or simply a lack of

detailed botanical exploration of the island.

In 2006, photos by Jonathan Newman of an unidentified taxon of

Nepenthes from Central Sulawesi, which were not readily assignable to

any presently known species, first appeared on the internet (http://www.

cpukforum.com/forum/index.php?showtopic=17326&hl=). Field visits to

the site in 2007 (S. McPherson and G. Bourke) and in 2008 (C. Lee and

S. McPherson), and examination of herbarium materials at Universitas

Tadulako, Palu (CEB) indicated that this plant represents a new species of

Nepenthes, which we describe below. The description that follows combines

measurements taken from the herbarium specimens at Universitas Tadulako

(which are sterile), with measurements of the inflorescence made from living

material in the field.

Nepenthes pitopangii Chi.C. Lee, S. McPherson, G. Bourke & M. Mansur,

spec. nov.

Nepenthi glabratae similis sed ascidiorum superiorum infundibuliformium

96 Gard. Bull. Singapore 61 (1) 2009

alis reductis differt - Holotypus: Lore Utara District, Poso Regency, Central

Sulawesi, 30 May 2007, RP 2054 (CEB). Fig. 1.

om Sy

1 A

AAU UU NONNNANNN'Y

Fig. 1. Nepenthes pitopangii. A. Climbing stem with upper pitcher and male inflorescence; B.

Male flower; C. Peristome teeth. All from living specimens. (drawing by Joseph Pao).

Nepenthes pitopangii (Nepenthaceae) 97

Terrestrial climber to ca 2 m tall. Climbing stems cylindrical to slightly

triangular in cross-section, particularly towards the developing shoot,

3-5 mm diam., internodes (4.1-) 6.4-9 cm long. Leaves of the climbing

stems chartaceous, sessile, lanceolate, apex acute, (10.1)11.5-14.4 (15.6)

cm long, (2)2.5-2.9 (3.4) cm wide; base clasping stem for about 2/3 its

circumference; longitudinal veins 3-4 on each side of the midrib, pennate

nerves inconspicuous; tendrils with a curl in the middle, 24-25 cm long.

Rosette pitchers unknown. Upper pitchers wholly infundibuliform, 3.8-4.5

mouth horizontal in front and slightly elevated towards the lid attachment;

peristome cylindrical, 1-3 mm wide, with distinct ribs, ca 0.45 mm apart, each

rib terminating in a blunt rounded tooth on the inner margin; interior of

pitcher without waxy zone, glandless immediately below peristome (ca 4

mm), otherwise evenly covered with glands ca 200 per cm’, yellow in color

above pitcher fluid, black in color where immersed; lid suborbicular, 2.9 by

2.8 cm, held horizontally over pitcher mouth, sides somewhat raised, lower

surface without appendages, evenly distributed with small crater-like rimmed

glands; spur simple, ca 1.5 mm long. Male inflorescence (measurements

taken from living specimens) a raceme, 37 by 2.5 cm; peduncle 18 cm long,

0.3 cm diam. at base, pedicels 7-9 (-11) mm long, each bearing a single flower,

bracts usually absent but occasionally towards base of inflorescence pedicels

with small filiform bract, ca 0.5 mm long inserted at about half the length of

the pedicel, tepals elliptic, ca 2 mm long, staminal column 2.5-3.0 mm long.

Female inflorescence and fruit unknown. Indumentum absent on all parts

of plant except for the developing pitcher and tip of the tendril, which are

covered with caducous silver-brown hairs. Color of living specimens: leaves

light green with red margins and red midrib on upper and lower surface;

stems dark red to purple with green spots; tendril red; upper pitchers pale

yellow with orange-red stripes, interior if pitcher and underside of lid pale

yellow; peristome orange-red.

Habitat. Secondary vegetation in submontane forest at an altitude of 1800 m.

Distribution: Lore Lindu National Park, Central Sulawesi.

Conservation Status: The geographical range of N. pitopangii is not yet

established (see below). Although the population from which all collections

and observations have been made consists of a single plant, it is unlikely

that this species does not grow elsewhere. We therefore propose an IUCN

Red List classification of “data deficient” (DD) until further detailed field

surveys can be conducted.

98 Gard. Bull. Singapore 61 (1) 2009

Notes: This species has anumber of characteristics in common with N. glabrata

Turnbull & Middleton, another montane species from Central Sulawesi.

These species share a similar leaf shape, pitcher lid, inflorescence, colouration

and lack of indumentum in nearly all parts. They appear to be very closely

related, but N. pitopangii differs in the unusual, widely infundibular upper

pitchers, as well as being more robust in all parts. Communication with R.

Pitopang, collector of the type specimen, confirmed that the specimens in

CEB were collected from the same individual plant as the one on which

the field observations were made by the present authors, this being the only

plant at the type locality. Given that no other plants of N. pitopangii have

been found to date, nothing is known about intraspecific variation within the

taxon and the possibility that this plant is a natural hybrid cannot be ignored.

However, we have discounted a hybridogenic origin for N. pitopangii, as

it does not display any morphological characteristics that suggest one.

All natural hybrids of Nepenthes recorded to date bear some obvious

similarities to either parent, but although N. pitopangii is sympatric with N.

maxima and N. tentaculata, it has little in common with these. The only other

species in Sulawesi with infundibular upper pitchers are N. maxima and N.

eymae and all known natural hybrids involving these bear petiolate leaves,

triangular lids, and/or appendages on the undersurface of the lid (C. Lee,

pers. observation), none of which are present in N. pitopangii. Moreover, the

upper pitchers of N. pitopangii are more widely infundibuliform than those

of N. maxima or N. eymae. Based on these observations, we have no doubt

that N. pitopangii is not a natural hybrid.

Field observations in the surrounding area and nearby mountains

(S. McPherson & G. Bourke in 2007 and C. Lee & S. McPherson in 2008)

did not reveal any additional plants of N. pitopangii, despite the presence of

extensive mossy forest in which N. maxima and N. tentaculata occurred. It

is therefore presumed that the principal habitat for this species may be at

lower elevations (below 1600 m) or perhaps, as with N. glabrata, it occurs

in submontane to montane scrub on ultramafic outcrops. Many species of

Nepenthes are opportunistic colonizers of disturbed areas, such as road

embankments with seedling recruitment primarily taking place before the

heavy growth of other plants. It is possible that the type specimen for this

species originated from a stray seed that germinated in a habitat, which had

only recently been cleared. The thick growth of ferns and other plants at this

site would normally impede the successful growth of a Nepenthes seedling.

The phenology, ecology and geographical range of N. pitopangii is

difficult to ascertain until more wild plants are discovered, but observation

conducted at the type locality showed that prey items in the upper pitchers

consisted primarily of small dipterans (particularly midges) and small

numbers of other insects such as wasps, ants, earwigs, and beetles.

Nepenthes pitopangii (Nepenthaceae) 99

Nepenthes pitopangii represents an interesting contribution to the

flora of Sulawesi, suggesting that more detailed explorations of the island

may yield further insights into patterns of diversity and endemism in

Nepenthaceae there, as well as further discoveries of new taxa. Detailed

field observations on other populations of N. pitopangii, should these be

found, will add greatly to our knowledge of this taxon.

This species is named for Dr. Rahmadanil Pitopang, curator of the

herbarium of Universitas Tadulako, who has studied the flora of Central

Sulawesi for over 18 years.

Specimens of other species examined: -Nepenthes glabrata: Res. Manado.

O. aft. Poso. Tusschen, Biv. III N uitlooper, van G. Loemoet, 3 Sep 1938,

Eyma 3585 (BO); E. Celebes, G. Loemoet, North Spur, 1938, Eyma 3585a

(BO); Menado O.A. Péso, B6zo—Péena, 1700-1800m, 10 Aug 1937, Eyma

1604 (BO). -Nepenthes hamata: Res, Menado. O. aft. Kolonedale, tusschen

Tomongkobae, Eyma 3969 (BO); Gunung Loemut, Eyma 3573 (BO);

Poso, Gunung Loemut, Eyma 3643 (BO); Tomongkobae, Eyma 3970 (BO):

Gunung Poka Pindjang, Kjellberg 1492 (BO): Mt. Roroka Timbu, summit

region, alt 2450 m, Balgooy 3335 (BO).

Acknowledgements

The authors would like to extend their gratitude towards Dr. Rahmadanil

Pitopang, for kindly providing access to study the type specimens, and to

Jonathan Newman for assisting with location data and maps. Special thanks

to Dr. Andreas Fleischmann for providing the latin diagnosis.

References

Kurata, Sh. 1984. New Species of Nepenthes from Sulawesi, Indonesia.

Journal of Insectivorous plant Society (Japan) 35:41-45.

Turnbull, JR. & A.T. Middleton. 1984. Three New Nepenthes from Sulawesi

Tengah. Reinwardtia 10:107-111.

Jebb, M. & M. Cheek. 2001. Nepenthaceae. Flora Malesiana 15: 1-157.

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Gardens’ Bulletin Singapore 61 (1): 101-128. 2009 101

Old Hats Are Better:

New Considerations and Taxonomic Changes

in the Southeast Asian Gardenia tubifera Complex

(Rubiaceae)

Y.W. LOW AND K.M. WONG

Institute of Biological Sciences,

University of Malaya,

50603 Kuala Lumpur,

MALAYSIA

E-mail: lowyeewen@yahoo.com

Abstract

Gardenia tubifera sensu Corner is revised over its entire range in Southeast

Asia. It is a heterogeneous complex of three distinct taxa, G. elata Ridl.,

G. subcarinata (Corner) Y.W.Low (elevated from varietal status) and G.

tubifera Wall., mainly distinguished by calyx form, fruit size and ecological

distribution. Two new varieties are described, G. elata var. longipedicellata

K.M.Wong (fromthe Philippines) and G.subcarinatavar.sumatrana Y.W.Low

(from Sumatra). A key for identification, descriptions, nomenclatural notes,

illustrations and exsiccatae examined for all recognised taxa are provided.

Gardenia elata is lectotypified.

Introduction

Gardenia tubifera Wall. sensu Corner (1939) is a heterogeneous taxon

including elements that are small to medium-sized trees occurring in

swamps and hill forests. Two varieties were recognised, namely, G. tubifera

var. subcarinata Corner (with a narrowly keeled calyx tube) and G. tubifera

var. tubifera (with a smooth calyx tube). Subsequently, G. tubifera var.

tubifera was further divided (Wong 1982, 1989) into two forms, G. tubifera

var. tubifera forma tubifera and G. tubifera var. tubifera forma elata (Ridl.)

K.M.Wong.

For the present work, taxonomic treatments by Corner (1939) and

Wong (1982, 1989) pertinent to the G. tubifera complex were reviewed. This

led to the distinctions reported here, including the resurrection of G. elata

Ridl.,a narrower circumscription of G. twbifera and the elevation of Corner’s

G. tubifera var. subcarinata to species level.

In this study, specimens of G. tubifera s./. from 14 herbaria (A, BKF,

102 Gard. Bull. Singapore 61 (1) 2009

BO, BRUN, IBSC, K, KEP, KLU, L, NY, SAN, SAR, SING and US) were

examined. A new variety is proposed for G. subcarinata from Sumatra in

Indonesia and another new variety for G. elata from Luzon in the Philippines.

Also, field observations of G. elata var. elata, G. subcarinata var. subcarinata

and G. tubifera were made to complement the findings from the herbarium

study.

Characters and species distinctions

Corner (1939) was convinced that G. tubifera was a very variable species

with regard to leaves, flowers and fruits. He adopted a broad concept for

G. tubifera, in which he included G. elata Ridl., G. resinifera Korth. and G.

speciosa Hook.f. as synonyms. His justification for this was largely based

on the calyx tube length, corolla tube length, and diameter across corolla

lobes in the open flower, the last of which he referred to as the ‘limb’ (more

precisely, limb span), which overlapped in the several taxa he considered. In

reaching this conclusion, Corner had considered earlier distinctions using

these characters by Hooker (1880), King and Gamble (1903) and Ridley

(1923), using measurements given by these authors, as well as from additional

specimens that were available. In so doing, and without the benefit of a

broader character survey including more species of Gardenia, Corner did

not attempt to use other characters for species distinction.

In the same paper, Corner (1939) also named another taxon with

keeled calyx tubes as a variety of G. tubifera. This taxon had corolla tube

length and ‘limb span’ that also fell within the range accepted by him for G.

tubifera.

The status of Gardenia tubifera var. subcarinata Corner

The most general characters that easily distinguish many species of Gardenia

appear to be those associated with calyx morphology. For example, in

Malaysia, there are three species of Gardenia with keeled calyces, namely,

G. carinata Wall., G. chanii Y.W.Low and G. pterocalyx Valeton. They can be

easily differentiated based on the form of the keels, for example, G. carinata

has keels resembling large triangular wings at the top of the subcylindrical

tube; G. chanii has keels resembling narrow elongate wings stretching down

the length of the tube to the top of the hypanthium only, the tube being

widely flared towards its apex; and G. pterocalyx has keels resembling

narrow elongate wings stretching down both the tube and the hypanthium,

the tube tightly ensheathing and not flared-out. The same characters were

useful in Hawaii (St. John and Kuykendall, 1949), where two species of

Gardenia with keeled calyces, G. mannii St.John & Kuykendall (keels with

New Considerations and Taxonomic Changes in the Southeast Asian Gardenia tubifera Complex 103

a laterally prolonged, narrowed proximal part ending in a spathulate apical

lobe) and G. remyi Mann. (keels resembling butterfly wings and without a

prolonged, narrowed proximal part), were distinguished.

We have also found that distinctive keel features are useful in

recognising individual species in our study. Additionally, there are also

consistent differences in other characters. For example, in distinguishing G.

chanii from G. pterocalyx (Low and Wong, 2007), leaf apex shape, domatia

type, leaf size, fruit pedicel length and corolla tube length provided consistent

differences in addition to calyx keel form. In fact, other than the placement

of G. tubifera var. subcarinata (with a slightly keeled calyx tube that often

shows clear marginal lobes) in a taxon that otherwise has a completely

smooth and subtruncate calyx tube, Wong (1982, 1989) also distinguished

species effectively based on calyx morphology, in particular, the tube and

keel form. It is true that a very small number of specimens of typical G.

tubifera sometimes display faint, rib-like traces along the calyx tubes, but

these are never laminate structures protruding from the calyx surface as are

keels, as defined here: similarly, dried fruits can be ribbed but not keeled.

We know of no other instances where Gardenia species have so great

a Variation in calyx form to include both smooth (unkeeled) and keeled

calyx tubes. In examining the probability that G. mbifera var. subcarinata has

been injudiciously grouped with typical G. mbifera (and therefore should be

recognised as a distinct species), we searched for other differences between

them. As, indeed, a number of such consistent differences exists (Table 1).

We conclude that G. tuwbifera var. subcarinata, with an easily distinguished

calyx form, should be recognised as a distinct species from G. tubifera. This

is here recognised as G. subcarinata (Corner) Y.W.Low comb. et stat. nov.

The two forms of Gardenia tubifera var. tubifera sensu Corner

Notwithstanding the distinctiveness (and usefulness) of calyx morphology

in distinguishing taxa, pairs of species with very similar calyx morphology

are known to occur that, nevertheless, can be well differentiated by other

characters. For example, G. coronaria Buch.-Ham.and G. thailandica Tirveng.

(both in the Myanmar-Thailand area but with overlapping distribution in

the isthmus region of the Thai-Malay Peninsula) cannot be distinguished

by their calyces as both have membraneous, sheathing calyx tubes with

oblique mouths. However, G. coronaria has longer corolla tubes (5.6-8.5 cm

long) and bigger corolla lobes (3-6 cm long, 2.2-3.4 cm broad), whereas G.

thailandica has shorter corolla tubes (3-5.8 cm long) and smaller corolla

lobes (1.5-2.1 cm long, 1-1.6 cm broad).

Another example is provided by G. beamanii Y.W.Low (north and

northwest parts of Borneo) and G. griffithii Hook.f. (Malay Peninsula,

104 Gard. Bull. Singapore 61 (1) 2009

Sumatra and Borneo), both with long and smooth calyx tubes sheathing the

corolla tube and with flared mouths. These species are distinguished by the

black dried exudate at shoot tips, puberulent lower leaf surface and longer

flower pedicels (0.5-0.8 cm long) of G. beamanii and the yellowish amber-

brown exudate at shoot tips, short-hairy lower leaf surface and shorter

flower pedicels (0.2-0.4 cm long) of the latter species.

We therefore conclude that it is possible for well-differentiated

species to share very similar calyx forms, but we wish to highlight that in

such cases it should still be possible to find a number of other consistent,

easily observable, differences in both vegetative and reproductive parts. In

other words, the distinctions between species are not slight or restricted to

a single character. Corner (1939) did not emphasize fruit size as a potential

species difference, especially in the case of G. tubifera sensu stricto (with

smaller fruits not exceeding 3 cm across) and G. elata (with larger fruits (3-

)4-6.5 cm across). We have conducted numerous surveys of these two species

in both Peninsular Malaysia and Borneo and find the fruit size difference is

highly consistent with other distinguishing features as outlined above (Table

1). In addition, the ecology may also be indicative. G. tubifera var. tubifera

forma elata is generally a lowland to hill forest taxon preferring drier sites

although they do (more rarely) occur in swamp forest areas, as observed

by Corner (1939). They can grow into trees of impressive size (to over 30

m tall: Ridley 11332). In contrast, G. tubifera var. tubifera forma tubifera is

a typically coastal estuarine and swamp forest species, and a much smaller

tree (up to about 13 m tall).

The two forms of G. tubifera var. tubifera recognised by Wong (1982,

1989) were distinguished also based on the hairiness of veins on the lower

leaf surface, leaf width, calyx tube length, corolla tube length, fruit size and

habitat. Table 1 shows that, whereas leaf width, calyx tube length and corolla

tube length are not consistent in their differences, further differences in

flower and fruit pedicel thickness, calyx mouth width and fruit size (verified

through the present study) indeed help in distinguishing these two rather

distinct taxa.

G. tubifera var. tubifera forma elata (previously G. elata) has a

puberulent midrib and secondary veins on the lower leaf surface, thicker

(2-3 mm) flower pedicels, a much wider calyx mouth [8-13(-15) mm], thicker

fruit pedicels [(2-)4-10 mm], and larger fruits [(3-)4-6.5 cm]. In contrast, G.

tubifera var. tubifera forma tubifera was distinguished by having a glabrous

midrib and secondary veins on the lower leaf surface, more slender flower

pedicels (1-2 mm), a narrower calyx mouth (5-8 mm), more slender fruit

pedicels (1.5-4 mm), and smaller fruits (2.4-3 cm).

In summary, these two taxa are differentiated by a suite of consistent

characters (Table 1), including various features of the reproductive parts,

New Considerations and Taxonomic Changes in the Southeast Asian Gardenia tubifera Complex 105

Table 1. A comparison of habitat and morphological characters among G.

subcarinata, G. elata and G. tubifera.

G. subcarinata

G. elata

G. tubifera

Provenance

Ecology

Leaf blade, length (cm)

Leaf blade, width (cm)

Leaf blade, pairs of sec-

ondary veins

Lower leaf surface,

pubescence on midrib &

secondary veins

Flower pedicel, length

(cm)

Flower pedicel, thickness

(mm)

Calyx tube, length (cm)

Calyx tube, width at mouth

(mm)

Calyx tube, pubescence

Calyx tube, keels presence

Corolla tube, length (cm)

Corolla tube, width at

middle (mm)

Corolla lobes, length (mm)

12-21

Corolla lobes, width (mm)

ameter at or near maturity

(cm)

Fruit pedicel, length (cm)

Fruit pedicel, thickness

(mm)

Malay Peninsula &

Sumatra

lowland forest

(4.5-)7-14

(1.9-)2.2-4.6

TA,

puberulent

0.1-0.5

1.2-1.8

puberulent

8-9 narrow keels

present along the

tube

4.6-6.8

1.5-2.5(-2.7)

0.1-0.4

Nicobar Islands,

Thailand, Malay

Peninsula, Sumatra,

Borneo, Sumbawa &

Philippines

lowland forest

(4.3-)10-22.5(-27)

(2-)4-11(-12)

(7-)14-22

puberulent

(0.6-)1.4-2.5(-3.5)

8-13(-15)

puberulent

keels absent

Q).2-1(-1.5) or more

(2-)4-10

Thailand, Malay

Peninsula,

Sumatra & Borneo

(Kalimantan only)

coastal estuarine &

swamp forest

9-15(-17)

generally glabrous

(very rarely sparsely

minute puberulent)

0.2-0.5

1:9

0.6-1.5(-1.9)

-8

sparsely puberulent

to subglabrous

keels absent

(2.4-)3.9-9.4

0.2-1.5

1.5-4

106 Gard. Bull. Singapore 61 (1) 2009

instead of just smaller differences that would be expected among forms of

the same species or variety. The different ecological distributions of these

taxa also support a more fundamental distinction. We propose that their

original distinction as species, G. elata Ridl. and G. tubifera Wall., should be

restored.

Two new varieties in the Gardenia tubifera complex

Based on similar considerations, we propose a new variety of G. elata from

Luzon material, and also a new variety of G. subcarinata from Sumatra.

These two new taxa differ from their respective typical varieties in only

minor details and distribution, rather than by the larger suites of characters

that consistently separate taxa at the species level (see key below).

The new variety of G. elata has longer flower pedicels (0.7-1 cm)

compared to the typical G. elata (only 0.1-0.5 cm long). On the other hand,

although the new variety of G. subcarinata has the keeled calyx tube form

of typical G. subcarinata, it differs in that the keels do not protrude beyond

the calyx tube margin (in typical G. subcarinata, the calyx keels protrude, as

distinct lobes, about 2 mm beyond the calyx tube margin). Otherwise, these

two taxa are indistinguishable from their typical forms.

Key to taxa in the Gardenia tubifera complex

1. Calyx outer surface keeled). 5.1... ciios -ocecneees - cece tech ae eee eee eee 2

1. Calyx outer surface smooth or at most very faintly ribbed upon drying (the

ribs resembling longitudinal veins, not laminate keel structures) ............. 3)

. Calyx keels ending in distinct spur-like expansions protruding up to 2mm

beyond the calyx tube margin (Malay Peninsula and Singapore) ...............

Bee es ete aren Pe them aath 7H ese Sundae G. subcarinata var. subcarinata

. Calyx keels not forming expanded apical portions and not protruding

beyond the calyx tube mareun (Sumatra) 22... oce cece r<sc5encc-e: eee

Fh enta pa etnec ees tes une «once t ee ten Cee erent ee eee an Ee ._ G. subcarinata var. sumatrana

im)

102)

. Veins on the lower leaf surface glabrous (very rarely sparsely minute

puberulent); calyx mouth of open flowers ca 5-8 mm wide; mature fruit ca

2.4-3 cm across (coastal estuarine to swamp forests only) ..... G. tubifera

. Veins on the lower leaf surface puberulent; calyx mouth of open flowers ca

8-13(-15) mm wide; mature fruit ca (3-)4-6.5 cm across (typically lowland

to hill forests, very rarely swamp LOreSt) (2325.2 yes ses ane 4

(Oe)

New Considerations and Taxonomic Changes in the Southeast Asian Gardenia tubifera Complex 107

4. Pedicels of open flowers ca 0.1-0.4(-0.5) cm long (Borneo, Malay Peninsula,

Nicobar Islands, Philippines, Singapore, Sumatra, Sumbawa, Thailand) ....

score oseceesiacec dEsSSccoae Reo A G. elata var. elata

4. Pedicels of open flowers ca 0.7-1 cm long (Philippines: known only in

Luzon from the Cagayan, Isabela and Rizal provinces) .............cccescesseeeeees

~ce sno ge pesnetgec See aac Eee ES ee G. elata var. longipedicellata

Taxonomic enumeration

1. Gardenia elata Rid. var. elata

J. Straits Branch Roy. Asiat. Soc. 79 (1918) 81. — Type: Singapore, Bukit

Timah, 1898, Ridley 11332 (lecto, K, hic designatus; isolecto, SING). Fig. 1.

-Gardenia tubifera var. tubifera forma elata (Ridl.) K.M.Wong, Gard. Bull.

Singapore 35 (1982) 22, Tree Fl. Malaya 4 (1989) 349; Coode et al., Checkl.

Fl. Pl. Gymnosperms Brunei Darussalam 270 (1996), quoad Ashton BRUN

1008, Ashton S 7834, Niga 52, Niga 63, Sands 5869, Simpson 2007, Wong

WKM 571.

-Randia speciosa Hook., Icon. Pl. 5 (1852) t. 824, nom. illeg., haud Randia

speciosa DC., Prodr. 4 (1830) 388; Gardenia speciosa Hook.f., Fl. Brit. Ind. 3

(1880) 117, King & Gamble, J. Asiat. Soc. Beng. 72 (2) (1903) 220, Ridley, FI.

Malay Penin. 2 (1923) 83, nomi. illeg., haud Gardenia speciosa Salisb., Prodr.

Stirp. Chap. Allerton (1796) 63, nec Gardenia speciosa Roxb. ex Wight &

Arn., Prodr. Fl. Ind. Orient. (1834) 422. -Type: Hook., Icon. Pl. 5 (1852) t. 824

(Randia speciosa Hook.).

-Gardenia lobbii Craib, Fl. Siam. 2 (1932) 120. (Craib proposed this as nom.

nov. for G. speciosa Hook.f.). See above.

-Gardenia longiflora S.Vidal, Revis. Pl. Vasc. Filip. (1886) 153, nom. illeg.;

Merrill, Enum. Philipp. Fl. Pl. (1923) 530; haud Gardenia longiflora Ruiz &

Pav., Fl. Peruv. 2 (1799) 67, t.219, nec Gardenia longiflora (Salisb.) Dryander

in Aiton, Hortus Kew., ed. 2, 1 (1810) 368. -Type: Luzon, Camarines Province,

Paracale, Jan 1884, Vidal 832 (open flower & fruit) (isotype, K).

-Gardenia longituba Ridl., J. Bot. 72 (1934) 274. -Type: British North Borneo,

Kudat, Jul 1885, Fraser 164 (flower bud) (holotype, K).

-“Gardenia glutinosa” auct. non Teijsm. & Binn. (1866): Elmer, Leafl. Philipp.

Bot. 4 (1912) 1331: quoad Elmer 13064.

-“Gardenia tubifera” auct. non Wall. ex Roxb. (1824): Corner, Gard. Bull.

Straits Settlem. 10 (1939) 46, pro parte: quoad G. speciosa Hook.f. & G.

elata Ridl. in syn.; Corner, Wayside Trees of Malaya | (1952) 541; Anderson,

Checkl. Trees Sarawak 297 (1980); Kessler et al., Secondary Forest Trees of

Kalimantan, Indonesia (2000) 135, quoad Fig. 134.

Tree to ca 30 m high, trunk to ca 119 cm diameter, not buttressed. Bark

108 Gard. Bull. Singapore 61 (1) 2009

Figure 1. Gardenia elata var. elata. A, Flowering leafy branch. B; Detail of puberulent veins

on lower leaf surface; C, Longitudinal section of fruit, calyx shown intact. [A & B from

Symington 24190 (SING); C from Sigin & Ismail SAN 100264 (L)].

New Considerations and Taxonomic Changes in the Southeast Asian Gardenia tubifera Complex 109

smooth, light grey-brown to dark brown. Stipules connate into a cylindrical

tube, (0.4-)0.6-1.5 cm long, apex 2-lobed to subtruncate, outside puberulent

and generally coated with resin (this sloughing off in older material), inside

with a mixture of dark-coloured colleters and scattered fine translucent

trichomes densely covering the basal half (trichomes slightly longer than

the colleters) and glabrous in the upper half. Petiole (0.3-)1.1-3.5(-3.8) cm

long, 1-2(-2.5) mm thick, evenly covered with puberulent hairs, sometimes

conspicuously resin-coated. Leaf lamina obovate to rarely elliptic, (4.3-)10-

22.5(-27) cm long, (2-)4-11(-12) cm wide; leaf base cuneate to rarely oblique;

leaf apex cuspidate; thinly coriaceous; conspicuously coated with resin when

young; midrib flat to sunken and minutely puberulent to subglabrous on

upper side, prominent and puberulent on lower side; secondary veins

(7-)14-22 pairs, flat and subglabrous on upper side, prominent and puberulent

on lower side, vein axils on the lower side with ciliate tuft-domatia to hairy

pocket domatia; tertiary venation scalariform. Flowers solitary. Pedicel 0.1-

0.4(-1) cm long and 2-3 mm thick in open flowers, reaching 0.2-1(-1.5) cm

long and (2-)4-10 mm thick at fruit maturity. Calyx narrowly obconical to

somewhat spindle-shaped, the apex often slightly oblique and torn into two

acute portions with corolla emergence, the tube subsequently slightly flared

outward and often appearing subtruncate in the open flower; medium green;

(0.6-)1.4-2.5(-3.5) cm long, 4-6 mm wide at the base, becoming 8-13(-15)

mm wide at the apex; outside densely puberulent at the base and sparsely

puberulent to subglabrous in the upper part, often coated with resin; inside

glabrous for most of the upper part to about 0.7 cm from the tube margin,

densely covered with a mixture of dark-coloured colleters and translucent

trichomes at the basal half (trichomes longer than colleters and especially

conspicuous as a dense fringe at the very base of the calyx); without lobes;

surface smooth, without keels or ribs. Corolla hypocrateriform, cream

turning light yellow, then deep to orange yellow; tube (4.5-)7-15 cm long, 2.5-

5 mm wide at the mid-portion, 6-13 mm wide at the throat, outside glabrous

to sparsely puberulent, inside largely glabrous except for a zone of dense

ribbon-like translucent hairs from the throat to around the middle of the

tube; lobes 8-10, oblanceolate to obovate, (16-)32-45(-50) mm long, 10-20(-

24) mm wide, contorted to the left in the bud stage, glabrous on both sides.

Stamens 8-10, inserted just below the corolla throat and between corolla

lobes, dorsifixed; filaments very short to inconspicuous; anthers 7-10 mm

long, ca a third to half of its length exserted; pollen in tetrads. Style (4.5-

)9-15.5 cm long, glabrous; stigma club-like with 5-8 lobes initially cohering

together, (4-)6-8 mm long, 3-5 mm wide, wholly exserted; ovary with several

parietal placentas. Fruits globose, rarely depressed globose or obovoid,

(3-)4-6.5 cm long, (3-)4-7 cm wide, surface in mature specimens smooth;

calyx persistent at fruit apex, the tube to 1-2.5 cm long, 0.8-1.5 cm wide at

110 Gard. Bull. Singapore 61 (1) 2009

the mouth; when ripe splitting irregularly to expose dark coloured seeds

embedded in a bright yellow-orange pulp. Seeds many, irregularly angular-

elliptic, flattened, 5-6 mm long, 4-7 mm wide, testa surface fine-areolate.

Habitat and ecology: Lowland forests preferring drier sites (including

ultramafic and volcanic soils in north Borneo and the Philippines), very

rarely in freshwater swamp forest.

Distribution: Widespread from the Nicobars, Thailand, Malay Peninsula,

Bangka, Sumatra, Sumbawa, Borneo to the Philippines (including Palawan)

(Fig. 2).

Proposed IUCN conservation assessment: Gardenia elata is a widespread

species although not commonly encountered,so that the status Least Concern

(LC) is still appropriate. However, with forest conversion rates increasing

steadily in the region, this status needs reassessment periodically.

MYANMAR

THAILAND

NICOBKARS *

__BORNEO

KALIMANTAN

— Gardenia elata var. elata

Seanase Gardenia elata var. longipedicellata

--- i

1,000 kilometres

—SSSSSSSSSSSSS=====_

Figure 2. Range of Gardenia elata var. elata (indicated by solid line, with uncertain parts

represented by dashed lines) and the area of occurrence of G. elata var. longipedicellata on

Luzon island, the Philippines (within dotted line).

New Considerations and Taxonomic Changes in the Southeast Asian Gardenia tubifera Complex 111

Notes: When Ridley first published G. elata, he cited only three specimens,

namely, Ridley 11332 (Bukit Timah, Singapore), Wray 4266 (Selama, Perak)

and Hose 229 (Baram, Borneo). As there was no particular type indicated,

all three specimens cited are syntypes. Corner (1939) considered G. elata a

synonym of G. tubifera Wall. In recognising G. elata as a form of G. tubifera

var. tubifera, Wong (1982) recorded Wray 4265 (Selama, Perak, SING) as

one of the syntypes, instead of Wray 4266 as stated by Ridley (1918). King

and Gamble (1903) had also listed Wray 4265 under G. speciosa Hook.f.,

also a synonym of G. elata. It has now been confirmed that Wray 4266

(SING) is a species of Brachylophon (Malpighiaceae) (Serena Lee, pers.

comm. ). As lectotypification is required for this species under Article 9.9 of

the International Code of Botanical Nomenclature (McNeill et al., 2006), we

have avoided choosing the Wray specimen but instead select Ridley 11332

(K) as the lectotype.

Gardenia longituba Ridl., overlooked by Corner (1939), was

considered closely related to G. longiflora S.Vidal by Ridley; in fact, both

these names are synonyms of G. e/ata. Ridley’s description of G. longituba as

having an unusually hairy stigmatic head was erroneous; close examination

of the type material, Fraser 164 (Kudat, British North Borneo, K), shows that

the otherwise smooth stigmatic head was in fact occluded by external fibres,

possibly from paper or cardboard material used when preparing herbarium

specimens. This taxon perfectly matches G. elata.

Specimens examined: BRUNEI: Belait District. Andulau Forest Reserve,

compartment 5, 10 Jun 2008, Low et al. LYW 180, fruits (BRUN, KLU):

Belait river, upstream from the Malayan river, 23 Oct 1988, Wong WKM

571, flower (A, BRUN, KEP), flowers (L, SAN); Bukit Sawat, Sungai Mau,

along Sungai Belait, 15 Oct 1991, Simpson 2007, fruit (A, BRUN, KEP, L,

SAN, SING): Sukang, Kampong Sukang, 21 Jul 1993, Sands et al. 5869, fruit

(A, BRUN, SAN, SING); Sungai Mau, 18 Aug 1988, Nangkat NN 52, fruit

(A, BRUN, SAN, SING): ibid., 28 Jan 1989, Nangkat NN 63, flower (KEP. L,

SAN, SING), flowers (A, BRUN), fruit (SING); Muara District. Berakas

Forest Reserve, 31 Sept 1959, Ashton BRUN 1008, fruit (BRUN, SING):

ibid., 12 May 1957, Ashton S 7834, flower (A, BO, BRUN, KEP, SAR).

INDIA: Nicobar Islands. North Nicobars, Katchall Island, 17 May 1975,

Chakrakanly 2557, flower (L). INDONESIA: Java. Cult. Hort. Bogor

V.10.49, 1903, Anon. s.n., flower (A); Tjibodas, sine date, Anon. s.n., flowers

(IBSC). Kalimantan. East Borneo. Berau, Inhutani area, Km 37 near transect

I, plot 6, 7 Oct 1997, Ambriansyah et al. Berau 841, fruit (A), fruits (L);

Commisi Kap. Genderen Stort., Gunung Samenggaris, Dec 1912, Amdjah

1094, flower (K, SING), flowers (A, BO 2 sheets), fruit (BO); Lojanan to

Tenggarong road, Kampung Rempaya, 26 Oct 1995, Ambri et al. AA 1416,

A Gard. Bull. Singapore 61 (1) 2009

flower (A, K, KEP, L, SAN); Sei. Seluang, 20 km from Wanariset, Waduk

road, 11 Jul 1995, Ambri et al. AA 1291, fruit (A, BO,SAN), fruits (L); South

Borneo. Sungai Wain region, North of Balikpapan, Oct 1950, Kostermans

4323, immature fruit (A); West Borneo. Pontianak, Bentiang, Gunung

Sekaju, West of Kampung Madamang, 6 Nov 1980, Shea 27578, fruit (A);

West Koetai, 21 Aug 1929, Endert 2765, fruit (A, BO). Sumatra, Bangka,

Lombok Besar, 1 Sep 1949, Kostermans & Anta 282, fruit (A, KEP), fruits

(L, NY, SING); East Coast, Asahan, Kuala Masihi, Apr 1927, Yates 2397,

flower (L, US), flowers (A, NY, SING); North Siberut Island, Gunung

Simapipit, 26 May 1994, A friastini 2737, fruit (K), fruits (L); Palembang, 16

Dec 1916, Lambach 1354, flowers (BO); Riau, Tigapuluh Mountains, 15 km

Southwest of Talanglakat, Rengat-Jambi road, vicinity of Sungai Serisih, 24

Nov 1988, Burley et al. 1680, fruit (KEP, L, NY, SING), fruits (A); South

Sumatra, Barisan Range, Seleman Enim, Bukit Seburong near Muara Dua,

15 Mar 1972, de Vogel 1299, fruits (L); Southeast Sumatra, Lampung, Way

Kambas, 3 Feb 1972, Mochtan 24A, fruits (L); West of North Sumatra,

Simaloer Island, 1 Nov 1918, Achmad 709, fruit (L). Sumbawa, Central

Sumbawa, Dompu, Raba Baka Trail to Matuatoi, 6 Jun 1961, Soejarto 60,

fruits (BO); West Sumbawa, Semongkat Atas, 17 km South of Sumbawa

Besar, 2 May 1961, Kuswata 112, fruit (BO, SING), fruits (A, BO).

MALAYSIA: Peninsular Malaysia. Johor, Kota Tinggi, Sungei Bang, 13 Mar

1966, Sinclair 10863, flowers (US 2 sheets); Mawai to Jemilang Road, Sungai

Berassau, 6 Feb 1935, Corner 28736, fruits (SING), Sungai Kayu near Sungai

Sedili, 10 Mar 1937, Kiah SFN 32368, fruits (A, SING), Sungai Sedili, 28 Mar

1937, Corner 32440, flower (BO), flowers (A, SING 3 sheets). Kedah, Koh

Mai Forest Reserve, 3 Apr 1938, Kiah SFN 35148, flowers (A, SING 2 sheets);

Ulu Muda Forest Reserve, 21 Jan 1969, Chan FRI 6777, fruit (A). Kelantan,

Kuala Krai, Taman Negara, Kuala Koh Headquarters, 30 Mar 1995, Latiff et

al. 4168, flower (K, L); Ulu Lebir Forest Reserve, 12 Aug 1970, Suppiah FRI

11681, fruit (L 2 sheets). Melaka, Chaban, 28 Sep 1885, Alvin 2364, fruits

(SING); Kemandore, 14 Jul 1917, Burkill 2509, fruits (SING). Negeri

Sembilan, Pasoh Forest Reserve, 6 Jul 1988, LaFrankie 3032, fruits (A); ibid.,

16 Jul 2008, Wong & Zulkapli s.n., leafy branch (KLU). Pahang, Rompin,

Pulau Tioman, Sungai Asah to waterfall, 29 Apr 1995, Zainudin & Bedul

5477, flowers (L), Ulu Sungai Sat, 11 Jul 1970, Mohd Shah & Mohd Noor MS

1833, fruit (A, L, US), fruits (SING). Perak, Gopeng, Apr 1884, King's

collector 5830, flower (L); Gunong Bubu via Trong, 27 Apr 1970, Suppiah

FRI 11673, fruit (K, L), near Selangore, Apr 1886, King’s collector 8736,

flowers (K); Selama, 1894, Wray 4265, flowers (SING). Selangor, Gombak,

27 Jun 1960, Poore 185, fruit (KLU); Kajang, Bukit Enggang, 9 Apr 1930,

Symington 24190, flower (SING); Sungai Buloh, 1891, Ridley s.n., fruits

(SING); Sungai Buloh Reserve, 25 Mar 1919, Abu 3313, flowers (SING).

New Considerations and Taxonomic Changes in the Southeast Asian Gardenia tubifera Complex 113

Terengganu, Kuala Trengganu, Jerteh, Gunung Tebu Forest Reserve,

compartment 65, 12 Oct 1971, Zainuddin FRI 17945, fruit (A, L, SING),

Kuala Trengganu, logging school area, 14 Nov 1978, Suppiah FRI 28251, fruit

(A). Sabah. Beaufort, Membakut, Kampung Binsulok Forest Reserve, 23

Apr 1984, Ag. Amin & Haya SAN 102465, flower (A, KEP, L, SAN, SING),

flowers (SAN); Beluran, Tongod, Ulu Sungai Pinangah, 16 Oct 1984, Amin

et al. SAN 107143, fruit (SAN 2 sheets); Kalabakan, Benaword logged over

area, 11 Apr 1980, Fedilis & Sumbing SAN 91785, fruit (SAN); Maliau Basin,

Rafflesia Camp to Resak Island, 26 Apr 2000, Ming et al. MB 283, fruit (KEP.

SAN); Keningau, Shang Lian logging area, LANAS, 16 Oct 1986, Mantor

SAN 118392, fruit (SAN) fruits (SAN); Kinabatangan, Gunung Rara Forest

Reserve, Maliau river, 11 Apr 1996, Puff 960411-1/2, flowers (SAN); Lamag,

Gunong Lotung, 5 miles Southeast of Inarat, 7 May 1976, Cockburn SAN

83039, flower (A, KEP, SAN, SING); Lamag, Sogo-sogo, Kampung Tongod,

22 Nov 1979, Madani SAN 91125, flower (K, KEP. SING), flowers (SAN);

Lamag, Tanegang Kechil, 26 May 1965, /. Singh & Eging SAN 51864, flowers

(K, SAN, SING); Sukau, Sungai Menanggul, 13 May 1996, Azmi et al. RA

512, flower (L), flowers (KEP), flower & flower bud (K, SAN); Kuala Penyu,

Kepayan, 20 Aug 1993, Ag. Amin SAN 127290, fruit (K), fruits (SAN);

Mempakul, Malikai, 25 Feb 1937, Mail 7060, flower (A, SING); Kudat,

without locality, 7 Aug 1885, Fraser 164, flowers (K); Berambangan, 9 Jul

1962, Brand SAN 30870, fruit (SAN); Dumpirit, Balajadia, 7 Feb 1933,

NBFD 2843, flower (BO), flowers (A); Lahad Datu, Mile 17.7 of Kalumpang-

Tawau road, 16 Sep 1962, Chai SAN 29828, fruit (BO, KEP, SAN, SING);

Pulau Sakar, 16 Mar 1961, H.S.M. & D. Brand SAN 24552, flower (SING),

flowers (SAN), flowers & fruit (BO, KEP.); Nabawan, Sepulut, Sepulut

Forest Reserve, Labang, 17 Oct 1988, Fedilis & Sumbing SAN 125652, fruit

(SAN); Sungai Tibow, 18 Jul 1984, Fedilis & Sumbing SAN 105342, fruit

(SAN); Papar, Mandahan Forest Reserve, 8 Jul 1987, Ag. Amin SAN 103348,

immature fruit (SAN), fruits (K); Ranau, without locality, 23 Feb 1990,

Majawat SAN 125800, fruit (KEP, L); Bongkud, 26 Mar 1986, Amin et al.

SAN 105640, flower (A, K, KEP, SAN, SING); Sandakan, Kretam, Sungai

Kulamba, 9 Apr 1984, Sundaling SAN 55998, flower (A, L, SAN 2 sheets):

Labuk Road Forest Reserve, 11 Feb 1993, Wong WKM 2600, flowers (SAN);

Sepilok Forest Reserve, Jalan Hg. Tanjong Cpt. 13,24 Sep 1968, Patrick SAN

63508, fruit (L), fruits (SAN); Sungai Dagat, 14 Jul 1987, George et al. SAN

120736, fruit (K, KEP), fruits (SAN); Sungai Malikop, 25 Aug 1984, Sigin &

Ismail SAN 100264, fruit (L, SAN); Telupid, Kampung Wonod, 19 Mar 1974,

Aban & Saikeh SAN 79413, flower (A, K, SING) flowers (A, KEP, SAN):

Semporna, Bodgaya, 6 May 1939, Valera SHN 10263, flower (SING), flowers

(KEP); Semporna, Mile 25 of Pagagau Road, 11 Mar 1965, J. Singh et al.

SAN 48883, flower (NY), flowers (SAN); Sipitang, Melaliah, 19 Oct 1961,

114 Gard. Bull. Singapore 61 (1) 2009

Md. Thaufeck SAN 27148 No. 19, fruit (BO, SING), fruits (SAN); Tawau,

Bombay Burmah Timber Company Concession, Sub-compartment no. 2 of

Compartment no. 1,28 Nov 1954, Wood SAN A3973, fruit (KEP), flowers &

fruits (L); Elphinstone Province, Oct 1922-Mar 1923, Elmer 20544, fruit (A

2 sheets, IBSC, L, NY, SING); Mostyn, Tengkayu Waterfall, Sabah Timbers

Company, 28 May 1965, Madani SAN 47171, flower (K), flowers (SAN);

Kalabakan Road, Mile 12, 26 Jul 1962, Aban SAN 30557, fruit (L, SAN),

fruits (KEP); Tenom, Mandalom Forest Reserve, 17 Sep 1986, Mantor SAN

116647, fruit (SAN 2 sheets). Sarawak. without locality, 1865-1868, Beccari

3250, flower (K), sine date, Native Coll. 214, flowers (US); 1st Division,

Kuching, Matang, Aug 1912, Anderson 6, flowers (SING); Kuching, Matang

Road, 10 Jul 1964, Salleh 12092, flower (A, K, NY); Mount Matang, 27 Oct

1929, J. & M.S. Clemens 22334, flower (K,SAR), flowers (A, K, NY); Kuching,

Santubong, 19 Nov 1904, Egon 252, fruit & flower (SAR); Simunjan, Serian

to Simanggang Road, Ulu Simpang Sabal Aping, Gunong Gaharu, 9 Oct

1974, Ilias & Azahari S 35687, fruit (KEP, L, SAN, SAR); 26th Mile Bau/

Lundu Road, Sampadi Forest Reserve, 17 Jun 1968, Jugah S 24948, flower

(SAN) flowers (A, K, SAR); 2nd Division, Sri Aman, 95th Mile, Kampong

Pungor Tapang, 9 Mar 1981, Ilias S 42712, fruit (KEP, L, SAN); 3rd Division,

Kapit, Balleh, Ulu Sungai Mengiong, Apan Entelit, 14 Mar 1996, Rantai et al.

S 74211, fruit (K, KEP, SAN, SAR, SING); 4th Division, Bintulu, Nyabau

Catchment Area, 22 Jun 1966, Sibat S 24617, flower (A, BO, KEP, SAN,

SING), flowers (K, SAR); 5th Division, Baram district, Miri river, Feb 1895,

Hose 506, flowers (K, L). PHILIPPINES: Busuanga. without locality, Sep

1922, Ramos Bur. Sci. 41218, immature fruit (K, L); NE of Coron, 2 km north

of San Nicolas, along Wayan Creek, 29 Jun 1984, Bourell 2439, fruit (A).

Culion, without locality, 29 Apr 1931, Herre 1085, flowers (A), flower &

flower buds (NY); Apr 1931, Herre 1088, fruit (NY). Luzon, Camarines

Province, Paracale, Jan 1884, Vidal 832, flower & fruit (K); Laguna Province,

Dahican River, Sep 1912, Ramos 1325, fruit (A, L, NY, SING); Tayabas

Province, May-Jun 1916, Cailipan For. Bur. 25640, fruit (K, US); Lucban,

May 1907, Elmer 7732, fruit (A). Mindanao, Zamboanga, Feb 1908, Whitford

& Hutchinson For. Bur. 9492, flower (NY, US). Palawan, without locality,

May 1913, Merrill 1360, flower bud & fruit (A, NY, SING); Bataraza, Bgy.

Sumbiling, Sitio Gamayon, Bulanjao Range, 8°33’N 117°24’E, 21 Mar 1995,

Soejarto & Madulid 9030, fruit (A, PNH); Puerto Princesa, Irawan, Impapai

hills above BFD Field Station, 9°51°N 118°37°E, 26 Jun 1992, Soejarto &

Fernando 7750, flower & flower buds (K, NY); Irawan, Irawan River Valley,

Tatanarom, road to Benguet mine, Mt Beaufort, 9°50°N 118°40°E, 16 Jul

1988, Soejarto & Madulid 6066, fruit (NY, SING, US); Irawan R. valley head,

19 Mar 1984, Ridsdale SMHI 145, flower bud (A, BO, K, KEP, L); lower

slopes of Mt Beaufort, 30 Mar 1984, Ridsdale SMHI 291, flowers & flower

New Considerations and Taxonomic Changes in the Southeast Asian Gardenia tubifera Complex IES)

bud (A, BO, K, L, SAN); Puerto Princesa, Mt Pulgar, Apr 1911, Elmer 13064,

flower (NY, US), flower & immature fruit (A); Pulot, Massin River, 12 km N.

Brooks Point, 23 Oct 1985, Ridsdale 998, fruit (A, L 2 sheets); Taytay, May

1913, Merrill 1279, flower (A, NY, SING); Taytay, island on Lake Manguao,

ca 10 km SE of Taytay town, 10°50’°N 119°33’E, 30 Jan 1991, Soejarto &

Fernando 7419, fruit (A, PNH); valley stream leading into NNW bay of lake,

7 Apr 1984, Ridsdale SMHI 357, fruit (A, BO, KEP, L, SAN). Panay, Capiz

Province, Oct-Nov 1925, Edano Bur. Sci. 46123, fruits (A, BO, NY, SING).

Sibuyan, Capiz Province, Magallanes, Mt Giting-Giting, Mar 1910, Elmer

12103, flower & flower buds (A) flower (NY, US). Tawi-tawi, Sulu Province,

Jul-Aug 1924, Ramos & Edano Bur. Sci. 44127, fruit (A, NY, SING, US).

SINGAPORE: Bukit Timah, 1898, Ridley 11332, flower (K, SING), Bukit

Timah Reserve, tree no. 166, 21 Jul 1938, Ngadiman SFN 35595, fruit (A);

Gutta Valley, 1907, Ridley s.n., fruit (SING). THAILAND: Peninsular

Thailand, Narathiwat, Waeng, Klong A-re-ma, 3 May 1999, Puudja 561,

flower (BKF); Pattani, Banang, 22 Jul 1923, Anon. 7275, fruit (K); Sukinin

District, Tomo Mine, 25 Dec 1999, Wongprasert 9912-38, fruit (BKF); Trang,

Talay Songkong, 19 Mar 1915, Vanpruk 661, flower (BKF), flowers (K).

la. Gardenia elata var. longipedicellata K.M.Wong, var. nov.

A var. typica pedicellis floris longioribus (7-10 mm longis) differt. -Typus:

Luzon, Cagayan Province, May 1921, Ponce For. Bur. 28435 [holo, A (sheet

1 of 2); isotypi, A (sheet 2 of 2), US]. Fig. 3.

Habitat and ecology: Lowland forests (including on ultramafic soils).

Distribution: Endemic to the Philippines, Luzon (Cagayan, Isabela and Rizal

provinces) (Fig. 2).

Proposed IUCN conservation assessment: Vulnerable due to very few known,

small or restricted populations (VU D2).

Note: This is recorded as a tree reaching about 15 m high. It shares much

similarity with the typical variety, differing only by its longer (0.7-1 cm)

flower pedicels compared to those of G. elata var. elata (only 0.1-0.4(-0.5)

cm long).

Specimens examined: PHILIPPINES: Luzon, Cagayan Province, May

1921, Ponce For. Bur. 28435, open flower (A, 2 sheets; US); ibid., Jan-May

1915, Velasco For. Bur. 24116, open flower & fruit (US); Isabela Province,

Mar 1910, Bernardo For. Bur. 15478, flower (L), Kapuntian, San Jose, San

Mariano, 16°59.7°N 122°2.3’E, sine date, Barbon et al. PPI 13137, immature

116 Gard. Bull. Singapore 61 (1) 2009

Snowy

“MencOuo

METRIC Wy 2 | 3 c) 5 6) ‘ 7 8 on:

“pedicel

Figure 3. Close-up of flower of Gardenia elata var. longipedicellata from Bernardo For. Bur.

15478 (L).

fruit (L); Isabela Province, Palanan, Digallorin, Divinisa camp site, 16°30°N

122°26°E, 10 Apr 1992, Ridsdale et al. ISU 479, flower bud (A, BO, K, L);

Rizal Province, Feb 1905, Ahern’s Collector For. Bur. 2673, fruit (NY, SING,

US); ibid., May 1907, Ramos Bur. Sci. 2689, open flowers (US).

2. Gardenia subcarinata (Corner) Y.W.Low, comb. et stat. nov.

Basionym: Gardenia tubifera var. subcarinata Corner, Gard. Bull. Straits

Settlem. 10 (1939) 48; Gard. Bull. Singapore 35 (1982) 22; Tree Fl. Malaya

4 (1989) 349. — Type: Penang, Government Hill, Feb 1889, Curtis 686 (holo,

SING 0048397; iso, SING 0048383). Fig. 4A-C.

-“Gardenia resinifera” auct. non Roth, Nov. Pl. Sp. (1821): Ridley, Fl. Malay

Penin. 2 (1923) 83.

-“Gardenia tubifera” auct. non Wall. ex Roxb. (1824): King & Gamble, J.

Asiat. Soc. Beng. 72 (2) (1903) 219 (as “Form 2”).

Tree, to ca 15 m high, trunk to ca 76 cm diameter, not buttressed. Bark

smooth, light grey-brown to dark brown. Stipules connate into a cylindrical

tube, 0.2-0.4 cm long, apex 2-lobed to subtruncate, outside puberulent and

New Considerations and Taxonomic Changes in the Southeast Asian Gardenia tubifera Complex 117

generally coated with resin (this sloughing off in older material), inside

with a mixture of dark-coloured colleters and scattered fine translucent

trichomes densely covering about three quarters of the surface from the base

(trichomes slightly longer than the colleters) and glabrous in the upper part.

Petiole (0.5-)0.6-1.5 cm long, 1-2 mm thick, evenly covered with puberulent

hairs, sometimes conspicuously resin-coated. Leaf lamina obovate (4.5-)7-

14 cm long, (1.9-)2.2-4.6 cm wide, leaf base cuneate, apex cuspidate, thin-

coriaceous, conspicuously coated with resin when young: midrib slightly

raised to flat and puberulent on upper side, prominent and puberulent on

lower side; secondary veins 7-12 pairs, flat to sunken and glabrous on upper

side, prominent and puberulent on lower side, vein axils on the lower side

with hairy pocket domatia; tertiary venation scalariform. Flowers solitary.

Pedicel 0.1-0.3 cm long and 1.2-1.8 mm thick in open flowers, reaching 0.1-0.4

cm long and 2.5-3 mm thick at fruit maturity. Calyx obconical, slightly flared

outwards at the apex: medium green: 0.4-0.7 cm long, 2-3.5 mm wide at the

base, becoming 4-8 mm wide at the apex: outside densely puberulent, often

coated with resin; inside glabrous for most of the upper half, densely covered

with a mixture of dark-coloured colleters and translucent trichomes at the

basal half (trichomes longer than colleters and especially conspicuous as a

dense fringe at the very base of the calyx); marginal lobes 7-9, subtriangular

to rounded, about 0.5-1 mm high: keels present, generally alternating with

the calyx lobes, forming narrow spurs at the apex that protrude up to 2

mm beyond the calyx tube margin, narrowing gradually towards the base

of the calyx limb but not extending downward to the hypanthium. Corolla

hypocrateriform, cream turning light yellow, then deep to orange yellow;

tube to 4.6-6.8 cm long, 1.5-3 mm wide at the mid-portion, 7-8 mm wide at

the throat, outside glabrous to puberulent, inside largely glabrous except for

sparse ribbon-like translucent hairs in narrow zones between stamens from

the throat to just below the anthers; lobes 6-9, oblanceolate to obovate, 12-

21 mm long, 6-13 mm wide, contorted to the left in the bud stage, glabrous on

both sides. Stamens 6-9, inserted just below the corolla throat and between

corolla lobes, dorsifixed; filaments very short to inconspicuous: anthers 5-7

mm long, c. a third to half of its length exserted; pollen in tetrads. Style 7.5-8

cm long, glabrous; stigma club-like with 3-4 lobes initially cohered together,

2-5 mm long, 1.5-3 mm wide, wholly exserted: ovary with several parietal

placentas. Fruits subglobose, 1.5-2.5(-2.7) cm long, 1.8-3 cm wide, surface

in mature specimens smooth; calyx persistent at fruit apex, the tube to 0.5-

0.7 cm long, 0.5-0.6 cm wide at the mouth, with low keels from its mouth to

the base, not extending downward to the fruit proper, forming apical spurs

projecting 0.5-1 mm beyond the calyx margin; when ripe splitting irregularly

to expose dark coloured seeds embedded in a bright yellow-orange pulp.

Seeds many, irregularly angular-elliptic, flattened.

118 Gard. Bull. Singapore 61 (1) 2009

Figure 4. Gardenia subcarinata. A-C. Gardenia subcarinata var. subcarinata: A. Leafy

branches, one terminated by a solitary flower; B. Fruit with persistent calyx; C. Calyx with

protruding apical keels of the less common (left) and the more common condition (right). D.

Gardenia subcarinata var. sumatrana, calyx without protruding apical keels. [A from 7. & P.

389 (KL 2989) (L); B from Curtis 686 (SING barcode no. 0048386); C from Ngadiman SFN

34926 (A) (left) and Zahir KEP 99132 (A) (right); D from Rahmat 1727 (A)].

New Considerations and Taxonomic Changes in the Southeast Asian Gardenia tubifera Complex 119

Habitat and ecology: Lowland forest (sea level to about 1750 ft [533 m)).

Distribution: Endemic to the Malay Penimsula (Fig. 5).

Proposed IUCN conservation assessment. Vulnerable due to fragmented

and declining area (VU A4ac).

Note: The type material for this taxon requires clarification. Corner

(1939) designated Curtis 686 in the Singapore Herbarium. collected from

Government Hill (=Penang Hill) m Penang. Peninsular Malaysia, as

the type of his G. mbifera var. subcarinata (the basionym of the species).

However, in the Singapore Herbarium there are six sheets with this number,

but differently dated. Two of the sheets are dated February of 1889. one

of which has Corner’s handwriting on it, stating “Type”. These are the two

sheets considered to represent the type material of the species, and the

sheet bearing Corner’s type annotation is considered the holotype, the other

regarded as an isotype sheet (see designations above). Another two sheets

are dated March, 1890. from the same locality but without any annotation

by Corner. The remaining two sheets are dated July. 1893, also without

annotation by Comer. Quite clearly, Curtis had assembled material from

three different gatherings from Government Hill and given them all his

number 686. Only the two sheets dated February. 1889. are to be accepted

as type material.

Specimens examined: MALAYSIA: Peninsular Malaysia. Johor, Bandar

Tenggara. Linggiu Forest Reserve. 23 Jul 1991, Lesmy FRI 35917, flower (A,

L. SING). Kedah, Bukit Enggang. Bukit Enggang Forest Reserve,4 Dec 1969.

Everett FRI 13766, fruit (L). fruits (A); Pedu. road from Pedu to Belatik, 15

May 1995. Zainudin & Bedul Moh 5541, flowers (L). Pahang, Raub. Sungai

Sempam. 15 Apr 1970. Soepadmo 666, fruit (BO, KUL): Taman Negara, path

to Terangan hide, 1 Apr 1975, Chan FRI 23851, flower (A. L. SING): Taman

Negara. trail from Terenggan to Kumbang Salt Lick. 1 May 1975, Balgooy

2603. flower (NY). flowers (L); Ulu Krau, Gunung Benom Game Reserve.

22 Apr 1967, Zahir KEP 99132. flower (A). Penang, without locality, Mar

1881, King’s collector 1474, flower (L); Government Hill, Feb 1889. Curtis

686, flowers & fruit (SING [sheet nos. 0048397 and 0048383}): ibid., Mar

1890, Curtis 686, flowers & fruits (SING [sheet nos. 0048386 and 0048520}):

ibid., Jul 1893, Curtis 686, flowers (SING [sheet nos. 0048524 and 0048394}):

Telok Bahang. Pantai Acheh Forest Reserve, 15 Sep 1966, Chelliah KEP

98143, fruits (A): Tiger Hill, 18 Nov 1950, Sinclair SFN 39095, flowers &

fruit (L. SING): ibid.. 18 Nov 1950, Sinclair 6705, fruit (US). Perak, Sungai

Kerian Estate. 29 May 1938. Spare SFN 34574, fruits (A): Taiping. 30 Oct

120 Gard. Bull. Singapore 61 (1) 2009

1969, Everett FRI 13596, fruits (A), Taiping, Maxwell Hill, 27 Feb 1983,

Khairuddin FRI 31835, flower & fruit (A); Maxwell Hill road, 30 Oct 1969,

Kochummen FRI 2919, fruit (A). Selangor, Kanching Forest Reserve, 25 Oct

1979, Kochummen FRI 11496, flower & fruit (A); Kuala Selangor, Sungai

Tinggi, 18 Oct 1937, Md Nur 34129, flower & fruits (A), fruit (L); Kuala

Lumpur to Kuala Selangor, 27 May 1971, T. & P. 389 (KL2989), flowers (L,

SING). Terengganu, Dungun, Bukit Bauk Forest Reserve, 19 Nov 1978,

Chan FRI 25155, fruit (A), Bukit Bauk, 27 May 1986, 7. & P. 1017 (KL

3517), fruit (L); ibid., 25 Jul 2006, Low et al. LYW 131, leafy branch (KLU).

SINGAPORE: Garden Jungle, 9 Dec 1889, Ridley 2588, flowers (SING);

Bukit Timah, 4 Apr 1938, Ngadiman SFN 34926, flowers (A, L).

2a. Gardenia subcarinata var. sumatrana Y.W.Low, var. nov.

A var. typica carinis calycis non extentis supra apicem tubus calycis differt.

— Typus: North Sumatra, Bila, Estate Aek-Buro, 15 Oct 1928, Lorzing 14218

(holo, SING; isotypi, A, L). Fig. 4D.

Habitat and ecology: Lowland forest (sea level to about 3281 ft [1000 m]).

Distribution: Endemic to Sumatra (Fig. 5).

Proposed IUCN conservation assessment: Vulnerable due to very few known,

small or restricted populations (VU D2).

Note: This is recorded as a tree reaching about 20 m high. It shares much

similarity with the typical variety, differing only in the keels on the flower

calyx tube, which do not expand into spur-like projections apically. In

contrast, in G. subcarinata var. subcarinata, the keels on the calyx tube form

conspicuous spur-like expansions at their apex.

Specimens examined: INDONESIA: Sumatra, East Coast Sumatra, vicinity

of Rantau Parapat, Bila, 28 Mar-10 May 1932, Rahmat Si Toroes 1727, flower

(A, NY, US); North Sumatra, Bila, Estate Aek-Buro, 15 Oct 1928, Lorzing

14218, flowers (A, SING), flowers & fruit (L); North Sumatra, Sibolangit,

1-4 Apr 1918, Bruinier 4, flowers (L); Baven Bandarbarat, 11 Aug 1918,

Lorzing 5914, flowers (L); Palembang, Banjoeasin, 16 Nov 1915, Grashoff

826, flowers (L).

3. Gardenia tubifera Wall. ex Roxb.

FI. Ind. ed. Carey & Wall. 2 (1824) 562 — Type: Singapore, Oct 1822, Wallich

Catalogue no. 8266, (holo, K-W; isotypes, K, sheets no. K000173277 &

K000173278). Fig. 6.

New Considerations and Taxonomic Changes in the Southeast Asian Gardenia tubifera Complex 121

. MALAY PENINSULA

SINGAPORE

\ \ 2A)

=> s

} SUMATRA Re

)

Figure 5. Distribution of Gardenia subcarinata var. subcarinata and G. subcarinata var.

sumatrana in West Sundaland area.

-Gardenia tubifera “form 1” & “form 3” sensu King & Gamble, J. Asiat. Soc.

Beng. 72 (2) (1903) 219. non “form 2” (= G. subcarinata).

-Gardenia tubifera sensu Comer, pro parte, Gard. Bull. Straits Settlem. 10

(1939) 46, Wayside Trees of Malaya 1 (1952) 541, excl. G. speciosa Hook.f.

& G. elata Ridl.

-Gardenia resinifera Korth., Ned. Kruidk. Arch. 2 (1851) 191, nom. illeg., non

Gardenia resinifera Roth, Nov. Pl. Sp. (1821) 150, nec Gardenia resinifera

sensu Ridley, Fl. Malay Penin. 2 (1923) 83. — Type: Borneo, Korthals s.n.

(syntype, L. 4 sheets).

-Gardenia glutinosa Teijsm. & Binn., Cat. Hort. Bot. Bogor. (1866) 119. nom.

invalid.

122 Gard. Bull. Singapore 61 (1) 2009

Tree toca 13 m high, trunk to ca 46 cm diameter, not buttressed. Bark smooth,

light grey-brown to dark brown. Stipules connate into a cylindrical tube, 0.4-

0.7 cm long, apex 2-lobed to subtruncate, outside puberulent and generally

coated with resin (this sloughing off in older material), inside with a mixture

of dark-coloured colleters and scattered fine translucent trichomes densely

covering the basal half (trichomes longer than the colleters) and glabrous

in the upper half. Petiole 0.2-1.5 cm long, 1-1.3 mm thick, subglabrous to

evenly covered with puberulent hairs, sometimes conspicuously resin-

coated. Leaf lamina obovate; 4.5-12.5(-21) cm long, 2.3-5.6(-6.3) cm wide;

leaf base cuneate; leaf apex cuspidate to rarely rounded; thinly coriaceous;

conspicuously coated with resin when young; midrib flat to sunken and

glabrous on upper side, prominent and glabrous to very rarely sparsely,

minutely puberulent on lower side; secondary veins 9-15(-17) pairs, flat and

glabrous on upper side, prominent and glabrous to minutely puberulent on

lower side, vein axils on the lower side with hairy pocket domatia; tertiary

venation scalariform. Flowers solitary. Pedicel 0.2-0.5 cm long and 1-2

mm thick in open flowers, reaching 0.2-1.5 cm long and 1.5-4 mm thick at

fruit maturity. Calyx narrowly obconical to somewhat spindle-shaped, the

apex often slightly oblique and torn into two acute portions with corolla

emergence, the tube subsequently slightly flared outward and often appearing

subtruncate in the open flower; medium green, 0.6-1.5(-1.9) cm long, 2-4

mm wide at the base, becoming 5-8 mm wide at the apex; outside densely

puberulent at the base and sparsely puberulent to subglabrous in the upper

part, often coated with resin; inside glabrous for most of the upper part to

about 0.5 cm from the tube margin, densely covered with a mixture of dark-

coloured colleters and translucent trichomes at the basal half (trichomes

longer than colleters and especially conspicuous as a dense fringe at the very

base of the calyx), without lobes, surface smooth, without keels but rarely

(upon drying) with a few faintly visible longitudinal ribs (probably the main

vascular traces). Corolla hypocrateriform, cream turning light yellow, then

deep to orange yellow; tube (2.4-)3.9-9.4 cm long, 1.5-3 mm wide at the mid-

portion, 5-9 mm wide at the throat, outside glabrous, inside largely glabrous

except for narrow patches of sparse ribbon-like translucent hairs from the

throat to just below the anthers; lobes 6-9, oblanceolate to obovate, 14-29

mm long, 7-18 mm wide, contorted to the left in the bud stage, glabrous on

both sides. Stamens 6-9, inserted just below the corolla throat and between

corolla lobes, dorsifixed; filaments very short to inconspicuous; anthers 6-8

mm long, ca a third to half of its length exserted; pollen in tetrads. Style (2.5-

)4.3-9.7 cm long, glabrous; stigma club-like with 4-5 lobes initially cohering

together, (3-)5-7 mm long, 2-4 mm wide, wholly exserted; ovary with several

parietal placentas. Fruits subglobose, 2.3-3.3 cm long, 2.4-3 cm wide, surface

in mature specimens smooth; calyx persistent at fruit apex, the tube to 0.5-

New Considerations and Taxonomic Changes in the Southeast Asian Gardenia tubifera Complex 12

Figure 6. Gardenia tubifera. A. Flowering leafy branch: B. Leafy branch with fruit: C.

Longitudinal section through lower part of flower including hypanthium: D. Transverse

section of fruit. [A from Low LYW 228: B from Low & Zulkapli LYW 163:C & D from Low

et al. LYW 35 (all KLU)].

124 Gard. Bull. Singapore 61 (1) 2009

1.4 cm long, 0.6-0.8 cm wide at the mouth; when ripe splitting irregularly to

expose dark-coloured seeds embedded in a bright yellow-orange pulp. Seeds

many, irregularly angular-elliptic, flattened, 4-5 mm long, 4-6 mm wide, testa

surface fine-areolate.

Habitat and ecology: Confined to coastal estuarine and swamp forest.

Distribution: Widespread from Thailand (Chonburi, Kao Sabab and

Peninsular Thailand), Malay Peninsula, Sumatra to Borneo (Kalimantan

only) (Fig. 7).

Proposed IUCN conservation assessment: Vulnerable due to fragmented

and declining area (VU Ad4ac).

Note: In the Kew Herbarium, the type material Wallich Catalogue no. 8266

is represented on three sheets. The first sheet, from Wallich’s Herbarium (K-

W: a fruiting specimen from Singapore) is not barcoded, but represents the

holotype, in accordance with Recommendation 9A.4 of the International

Code of Botanical Nomenclature (Vienna Code) (McNeill et al. 2006). The

second sheet, incorporated from Hooker’s Herbarium and bearing Kew

barcode K000173277 (also a fruiting specimen), is an isotype. The third

sheet contains two different collections; the specimen on the lower half of

the sheet (a fruiting specimen) is a duplicate of Wallich Catalogue no. 8266,

barcoded K000173278, and also represents an isotype, whereas the specimen

on the upper half of the sheet is Hervey’s collection of 1886 (including a

fruit and a flower) and is not a part of the type material.

Specimens examined: INDONESIA: Kalimantan. Central Borneo, Kec.

Mentaya Hilir Utara Sei Sampit, Bagendang, 25 Feb 1982, Afriastini 341,

flower (L); Kumai, Sungai Bekunyir, 14 Mar 1975, Anderson (1975)2, flowers

& fruit (BO 2 sheets); East Borneo, Pesiangan Bengka and Muara Kaman,

Mahakam river, 24 Aug 2000, Adriansyah AA 3002, fruit (A, K); Samarinda,

Sungai Pedang Kota Bangun Ulu, 23 Feb 1992, Ambri & Arifin AA 443, fruit

(A, K); South Borneo, Bangarmassing, 1857-1858, Motley 341, flowers (K);

Maharanda, 1918, Anon. 15b, flowers (BO); Pleihari, 22 Aug 1965, Sauwveur

965, flowers (L); Poeloe Lampei, sine date, Korthals s.n., fruits (L); 1bid.,

sine date, Korthals 2415, leafy branch (L); ibid., sine date, Korthals 2416,

leafy branch (L); ibid., sine date, Korthals 2417, leafy branch (L); Rantau

to Maugasari, 14 Dec 1988, Giesen 70, fruit (L), fruits (L); Tanah Laut

District, Hutan Kintap base camp, 20 Apr 1985, Leeuwenberg & Rudjiman

13460, flowers & fruit (L); Z. O. Borneo (Southeast Borneo), Veenbosch

bij Tamban, 10 Oct 1939, Polak 480, fruit (A, BO, L, SING); West Borneo,

New Considerations and Taxonomic Changes in the Southeast Asian Gardenia tubifera Complex #25

THAILAND

PALAWAN

MALAY PENINSULA

BORNEO

KALIMANTAN

@ Gardenia tubifera

1,000 kilometres

——

Figure 7. Distribution of Gardenia tubifera in the Sundaland area.

Kenepai, sine date, Teysmann 8332, fruits (L); Lake Tajan, 22 Oct 1949, Main

1746,immature fruit (A, K) fruit (BO): Salimbauw, sine date, Hallier B 1257,

flowers (L 2 sheets) flower & fruit (L); Soeka Lanting, sine date, Hallier B

183, fruit (L); Soengai Kenepai, sine date, Hallier B 1906, fruit (L); Sungei

Batang Putus, South of Danau Sentarum Wildlife Reserve, 31 Mar 1986,

Giesen 1, flowers (L) flower & fruit (L); West Koetai, No. 3 near Kampong

Sabentoeloeng, 20 Jun 1925, Endert 1518, fruits (L 2 sheets), flower & fruit

(A, BO, SING). Sumatra, Palembang, Sematang, 19 Jan 1920, Endert E 808,

flowers (L); Palembang, Kajoeagoeng, 29 Mar 1928, de Voogd 139, flowers

(L 2 sheets); ibid., 23 Jan 1929, de Voogd 283, flower (BO); Upper Riau,

Pekanbaru, Tenajan Reserve, 11 Aug 1960, Soepadmo 20, fruit (K, SING).

126 Gard. Bull. Singapore 61 (1) 2009

MALAYSIA: Peninsular Malaysia. without locality, 1862-1863, Griffith KD

2816, flower (L), flowers (K). Johor, Bekok River, 1880, Ridley 11139, flower

& fruit (SING); Kluang, 5 Mar 1973, Hardial Singh & Samsuri HS 1092, fruit

(US), flower & fruit (A); Kota Tinggi, Dec 1892, Ridley 4209, flower & fruit

(SING), Kota Tinggi, Sungai Bang, 13 Mar 1966, Sinclair 10863, flower (A,

SING); Kuala Sembrong, 1892, Lake & Kelsall 4088, flowers & fruit (SING);

Sungai Sedili, Bagan Limau, 18 Feb 1931, Corner SFN 24625, flower & fruit

(SING), Sungai Sedili, Danau, 27 Mar 1932, Corner SFN 25971, flower

& fruits (BO, SING). Kuala Lumpur, University of Malaya, Rimba Ilmu

Botanic Garden, cultivated opposite the medicinal plants section near the

front entrance, 24 Dec 2008, Low LYW 228, flower (KLU). Melaka, without

locality, sine date, Anon. s.n., flower (SING), 1845, Griffith s.n., flowers &

fruit (K, NY), Aug 1886, Hervey s.n., flower & fruit (K), sine date, Lobb

347, flowers (K), 1871, Maingay KD 838, flowers & fruit (K 2 sheets, L),

1871, Wright s.n., flowers (K); Bt. Sadanau, Feb 1890, Derry 360, flowers &

fruit (SING); Merlimau, Jun 1889, Derry 199, fruits (SING); Sungai Tebong,

South of Bukit Putus, 30 Jan 1916, Burkill SFN 1434, flower (SING); 5 miles

South of Malacca, Batu Berendam Road, 29 Apr 1961, Burkill HMB 2640,

fruits (A, SING). Negeri Sembilan, 12 miles Simpang Pertang-Kuala Pilah

roadside, 2 Sept 1977, Asri FRI 25738, fruit (A). Pahang, Lubuk Paku, 28

Nov 1924, Ngadiman SFN 16114, flower & fruits (SING); Muazam Shah to

Menchali Road, road to Kampung Sedaik Asal, | May 2008, Low et al. LYW

167, fruit (KLU); Pekan, 28 Nov 1929, Burkill & Haniff SFN 17129, flowers

(A), flowers & fruit (SING); ibid., 20 Aug 1909, Ridley s.n., flower (SING);

ibid., May 1890, Ridley 1375, flower (BO), flowers (BO, SING); Pekan,

Nenasi Forest Reserve, 17 Apr 2004, Ong EL 36, flower & fruits (KLU 2

sheets); Pekan, Sungai Bebar, 10 Apr 2005, Low et al. LYW 35, flower & fruit

(KLU); ibid., 30 Oct 2007, Low & Zulkapli LYW 163, fruit (KLU); Rompin,

Menchali Forest Reserve, | May 2008, Low et al. LYW 169, fruit (KLU);

Tasek Bera, 24 Apr 1980, Gianno 370, flower & fruit (KLU); ibid., 7 Sept

2005, Low et al. LYW 70, fruits (KLU); ibid., 10 Nov 1975, Stone et al. 12264,

flower (KLU), flowers (KLU), fruit (NY); Tasek Bera, Kota Iskandar, 1 Feb

1962, Anon. 1061, flowers & fruit (KLU); Tasek Bera, near Sungai Bera, 15

Oct 1930, Henderson SFN 24126, fruit (NY). Perak, Sungai Kerian Estate,

29 May 1938, Spare SFN 34574, fruits (SING). Perlis, Kangar, Jalan Batu

Pahat (cultivated?), 22 Aug 1991, Zainudin et al. AZ 3825, fruit (L), leafy

branch (K). Selangor, Petaling Jaya, Section 16, cultivated inside a bungalow

compound (private property),3 Mar 2006, Low etal. LYW 164,flowers (KLU);

Sungei Buloh Reserve, Kuala Lumpur, 3 Mar 1923, Foxworthy 7960, flowers

(SING). SINGAPORE: without locality, sine date, Burkill 324, fruits (US),

Oct 1822, Wallich Catalogue no. 8266, fruit (K 2 sheets, K-W). THAILAND:

Chonburi, Sriracha, 4 Feb 1927, Collins 1407, fruits (US); ibid., 7 Dec 1927,

New Considerations and Taxonomic Changes in the Southeast Asian Gardenia tubifera Complex 27

Collins 1833, flower (US 2 sheets); Kao Sabab, 18 Jan 1958, Sorensen et al.

506, fruit (BKF, L); Peninsular Thailand, Chumpon, Langsuan, 10 Jan 1977,

Santisuk 908, fruit (A); Satul, Kuan La Long, 30 Jan 1961, Ploenchitr 1602,

flower (BKF).

Acknowledgements

We thank the Keepers and Curators of various herbaria for loans and

permission to examine materials in their care: Arnold Arboretum, Harvard

University (A), Forest Herbarium, Thailand (BKF), Herbarium Bogoriense

(BO), Brunei National Herbarium (BRUN), South China Botanical

Garden, Guangzhou (IBSC), Royal Botanic Gardens, Kew (K), Forest

Research Institute, Malaysia (KEP), University of Malaya (KLU), Nationaal

Herbarium Nederland. Leiden (L), University of Michigan (MICH), New

York Botanical Garden (NY), Forest Research Centre, Sandakan (SAN),

Forest Research Centre, Sarawak (SAR), Singapore Botanic Gardens

(SING) and the United States National Herbarium, Washington (US). Help

in obtaining literature and digital images was obtained through Datuk C.L.

Chan (Kota Kinabalu); Prof. J.H. Beaman, J.J. Wood and Marie Briggs (K):

Dr C. Anderson and H. Huggins (MICH). S. Lee (SING), Dr B.W. Eko,

Dr Irawati and H. Arief (BO), and A. Bond, Dr A. Davis and S. Dawson

(K) kindly assisted with specimen loans and information. Field logistic

support was provided through J.T. Pereira, J. Sugau and M. Postar (Sabah);

M. Sugumaran, I. Zulkapli, K.T. Yong and I.S. Shanmugaraj (Peninsular

Malaysia); H.J. Jamilah, A.K. Muhd. Ariffin, E. Jangarun, P. Azlan and M.

Jakaria (Brunei). We are grateful to Prof. C. Puff (University of Vienna)

for providing the Latin diagnoses used here. Zainal Mustafa (Rimba I]mu

Botanic Garden) prepared the botanical drawings. This work results from an

MSc programme (Y.W.L.) at the Institute of Biological Sciences, University

of Malaya, and was supported in part by university research grants PS046-

2007C and FS266-2008C.

References

Corner, E.J.H. 1939. Notes on the systematy and distribution of Malayan

phanerogams, I. Gardens’ Bulletin, Straits Settlements 10(1): 46-48.

Hooker, J.D. 1880. The Flora of British India, Vol. 3. L. Reeve, London.

(Gardenia, pp. 115-120.)

King, G. and J.S. Gamble. 1903. Rubiaceae. In: Materials for a Flora of the

Malayan Peninsula, Gamopetalae. Journal of the Asiatic Society of Bengal

128 Gard. Bull. Singapore 61 (1) 2009

72(2): 216-221.

Low, Y.W. and K.M. Wong. 2007. Two new species of Gardenia (Rubiaceae)

from Borneo and notes on Gardenia pterocalyx. Edinburgh Journal of

Botany 64(1): 25-36.

McNeill, J., ER. Barrie, H.M. Burdet, V. Demoulin, D.L. Hawksworth, K.

Marhold, D.H. Nicolson, J. Prado, P.C. Silva, J.E. Skog, J.H. Wiersema and

N.J. Turland (eds.). 2006. International Code of Botanical Nomenclature

(Vienna Code). A.R.G. Gantner Verlag KG, Ruggell, Liechtenstein.

Ridley, H.N. 1923. The Flora of Malay Peninsula, Vol.2.L. Reeve & Co., Ltd,

London. (Gardenia, pp. 79-84.)

St. John, H. and J.R. Kuykendall. 1949. Revision of the native Hawaiian

species of Gardenia (Rubiaceae). Hawaiian Plant Studies, No. 15.

Brittonia 6(4): 431-449.

Wong, K.M. 1982. Notes on Gardenia and Acranthera (Rubiaceae) from

Peninsular Malaysia. Gardens’ Bulletin, Singapore 35(1): 21-32.

Wong, K.M. 1989. Rubiaceae. In: F.S.P. Ng (ed.), Tree Flora of Malaya, Vol. 4.

Longman Malaysia Sdn. Bhd., Kuala Lumpur. (Gardenia, pp. 348-350.)

Gardens’ Bulletin Singapore 61 (1): 129-143. 2009 129

A Revision of Bothriochloa Kuntze (Poaceae) in Thailand

O.NEAMSUVAN .J.VELDKAMP’ AND T.SEELANAN

‘Department of Botany, Faculty of Science, Chulalongkorn University,

d Bangkok 10330, Thailand.

‘National Herbarium of The Netherlands, Leiden University,

P.O. Box 9514, 2300 RA Leiden, The Netherlands.

_ E-mail: tosak.s@chula.ac.th

Abstract

Bothriochloa Kuntze (Poaceae) has 3 species in Thailand: B. bladhii,

B. ischaemum, and B. pertusa. A key, descriptions, and illustrations are

provided.

Introduction

The genus Bothriochloa was established by Kuntze in 1891. Its name is

derived from the Greek words fpo8etov (bothrion, a small pit) and yAon

(chloé, young shoot, grass), which refers to the pitted lower glumes of the

spikelets in some species of this genus (Bor, 1968). It belongs to the family

Poaceae, tribe Andropogoneae, subtribe Sorghinae and comprises about 33

pan(sub)tropical species (Clayton, Harman and Williamson, 2008).

The genus is characterised by a translucent median channel in the

joints and pedicels of the raceme. Its nearest relatives appear to be Capil-

lipedium Stapf and Dichanthium Willemet, and because B. bladhii (Retz.)

S. T. Blake can hybridise with some species of these two genera. Some have

therefore advocated to unite the three into Dichanthium, the oldest name

(De Wet & Harlan, 1966), but then infrageneric taxa are recognised, e.g.,

by Roberty (1960), who enumerated these three amongst no less than 12

sections in Dichanthium: Dichanthium sect. Amphilophis (Trin.) Roberty,

Dichanthium sect. Bothriochloa (Kuntze) Roberty, and Dichanthium sect.

Dichanthium.

Bothriochloa can be distinguished from the other two genera as

follows:

-Inflorescence composed of subdigitate racemes, rarely paniculate, racemes

PAIL ARMIOKe Mame SiS PikClOL PAINS: cic5..cins.ceseescescescstestecesaceateseeeehece Bothriochloa

-Inflorescence a panicle, racemes with 1-5(-8) spikelet pairs «0.0.0... eeeeeeeeeees

130 Gard. Bull. Singapore 61 (1) 2009

-Joints of the rachis and pedicels with translucent longitudinal channels; all

spikelet pairs heterogpamOus 1s ee Bothriochloa

-Joints of the rachis and pedicels without translucent longitudinal channels;

spikelet pairs heterogamous, excepting homogamous at basal spikelet pairs

ssosodecnsduddeanutitvs dole dvucegd gees dando tbac Gee Amma tesa a een eee Dichanthium

For Thailand, Bor (1965) reported B. glabra (Roxb.) A. Camus and

B. pertusa (L.) A. Camus without giving a key or descriptions. Sathagul

(1990) in her Master’s thesis recorded 6 species: B. caucasica (Trin.) C.E.

Hubb., B. glabra, B. insculpta (Hochst. ex A. Rich.) A. Camus, B. intermedia

(R. Br.) A. Camus, B. ischaemum (L.) Keng, and B. pertusa and provided a

key and descriptions. Nanakorn and Norsangsri (2001) reported 5 species: B

caucasica, B. glabra, B. intermedia, B. ischaemum, and B. pertusa. However,

because this was a mere checklist for Thai grasses there was neither a key

nor a description. Later studies have shown that B. caucasica, B. glabra,

and B. intermedia are to be considered part of the very polymorphic

(“compilospecies”: De Wet & Harlan, 1966) B. bladhii (Retz.) S.T. Blake.

In the present study in the framework of the Flora of Thailand

Project, three species could be distinguished: B. bladhii, B. ischaemum, and

B. pertusa. A key, descriptions, and illustrations are presented.

Materials and methods

This study was based on an intensive search through available taxonomic

literature and a study of specimens kept at the herbaria in Thailand: the

Bangkok Herbarium (BK), the Forest Herbarium, National Park, Wildlife

and Plant Conservation (BKF), the Kasin Suvatabhandhu Herbarium,

Department of Botany, Chulalongkorn University (BCU), The Herbarium,

Department of Biology, Chiang Mai University (CMU), Prince of Songkhla

University Herbarium (PSU) and The Herbarium of Department of Botany,

Kasetsart University. Several herbaria abroad also were visited: the National

History Museum, London (BM), the Royal Botanic Gardens, Kew (K),

the Museum National d‘Histoire Naturelle, Paris, France (P), the National

Herbarium of The Netherlands, Leiden, The Netherlands (L), the Botanical

Museum, Natural History Museum of Denmark, University of Copenhagen

(C), and the Herbarium, Institute of Biological Sciences, University of

Aarhus (AAU). Field work was carried out throughout Thailand during

which additional ecological data and localities were recorded.

Bothriochloa Kuntze

Revis. Gen. Pl. 2: 762. 1891. -[Bothriochloa sect. Eubotriochloa Ohwi, Acta

Phytotax. Geobot. 11: 166. 1942, nom. inval.]. -Dichanthium Willemet sect.

A Revision of Bothriochloa Kuntze (Poaceae) in Thailand 131

Bothriochloa Roberty, Boissiera 9: 159. 1960. — Type species: Bothriochloa

anamitica Kuntze [= Bothriochloa bladhii (Retz.) S.T. Blake].

Andropogon L.sect. Amphilophis Trin., Mém. Acad. Imp. Sci. St. Pétersbourg,

VI, Sci. Math. 2: 285. (1832) (rank indicated on p. 279). -Andropogon

L. subgen. Amphilophis Trin. ex Hack. in Mart., Fl. Bras. 2(3): 291. 1883.

-Amphilophis Nash in Britton, Man. Fl. N States: 71. 1901. -Bothriochloa

sect. Amphilophis Ohwi, Acta Phytotax. Geobot. 11: 166. 1942.-Dichanthium

Willemet sect. Amphilophis Roberty, Boissiera 9: 167. 1960.—Lectotype

species (not resolved). Not Amphilophis torreyanus Nash [=Bothriochloa

laguroides (DC.) Herter var. torreyana (Steud.) M. Marchi & Longhi-

Wagner, Bol. Inst. Bioci. Univ. Fed. Rio Grade do Sul 57: 52. 1998, f. 6, 19],

fide ING (http://ravenel.si.edu/botany/ing/ingForm.cfm), see note.

Tufted perennials. Culms slender. Leaf sheath keeled, glabrous; ligules

membranous; leaf-blade flat. Inflorescence subdigitate or paniculate,

composed of many racemes, each consisting of several pairs of sessile and

pedicelled spikelets; joints and pedicels with a translucent longitudinal

groove, flattened, hairy on both margins. Sessile spikelets dorsally compressed,

elliptic, oblong to lanceolate, 2-flowered, callus short, shortly bearded; lower

glumes as long as spikelet, smooth or pitted, 7-11-nerved, chartaceous to

membranous, pubescent below the middle, laterally 2-keeled, pectinately

setose on keel near tip, apex acute, margin inflexed; upper glumes equally

long or somewhat shorter, boat-shaped, 3-nerved, 3-keeled, subchartaceous

to hyaline, apex acute, margin inflexed; lower lemmas nerveless, hyaline;

upper lemmas linear, hyaline, continuous with the geniculate and twisted

awn; lower paleas absent; upper paleas small or absent. Pedicelled spikelet

1- or 2-flowered, the lower male or neuter, the upper neuter or more often

suppressed; lower glume chartaceous, glabrous, margin inflexed; upper

glume hyaline, glabrous, apex acute, margin inflexed; lower lemmas hyaline,

glabrous; awnless.

Distribution: 33 species, throughout the (sub)tropics, 3 in Thailand.

Note: We have not seen Nash (1901), but this lectotypification seems

incorrect. Amphilophis torreyanus was based on Andropogon torreyanus

Steud. [(1840), nom. nov. pro Andropogon glaucus Torr. (1824), non

Retz. (1789)]. This species is not to be found in Trinius (1832), where the

apparently unranked (but see rank indicated on p. 279) groups are a medley

of taxa, now attributed to Bothriochloa, Chrysopogon Trin. (Vetiveria Bory),

Sorghastrum Nash, and Sorghum Moench. A recent paper by Soreng &

Pennington (2003) also did not resolve the question of the type species.

1B2 Gard. Bull. Singapore 61 (1) 2009

Key to the species-

1. Panicle, lowest raceme shorter than main axis of inflorescence ...................

sidisanideebl mete eee ee eee eee 1. B. bladhii

1. Subdigitate, lowest raceme longer than main axis of inflorescence ......... Zs

2. Upper glumes hairy on the upper part of nerves. Lower glume of sessile

spikelet:without a cisculan pie: eee ee ee 2. B. ischaemum

2. Upper glumes glabrous. Lower glume of sessile spikelet with a distinct

circular pit 3.22 eee eee ee eee 3. B. pertusa

1. Bothriochloa bladhii (Retz.) S.T. Blake

Proc. Roy. Soc. Queensland 80: 62. 1969. - Andropogon bladhii Retz., Observ.

Bot. 2:27. 1781.-Andropogon annulatus Forssk. var. bladhii (Retz.) Hack. in

A. DC., Monogr. Phan. 6: 572. 1889. -Dichanthium bladhii (Retz.) Clayton,

Kew Bull. 32: 3. 1978. — Typus: China, Bladh s.n. in Herb. Retzius (LD, holo,

94/019-0745, SI, photo). Fig. 1.

-Andropogon intermedius R. Br., Prodr. 1: 202. 1810. -Andropogon

intermedius R. Br. var. genuinus Hack. in A. DC., Monogr. Phan. 6: 485, 1889.

nom. inval.-Sorghum intermedium (R. Br.) Kuntze, Rev. Gen. Pl. 2:792.1891.

-Amphilophis intermedia (R. Br.) Stapf, Agric. News (Barbados) 15: 179.

1916; in Prain, Fl. Trop. Afr. 9: 174. 1917. -Bothriochloa intermedia (R. Br.)

A. Camus, Ann. Soc. Linn. Lyon I, 76: 164. 1931. -Dichanthium ischaemum

(L.) Roberty subvar. intermedium (R. Br.) Roberty, Boissiera 9: 160. 1960.

nom. inval. -Dichanthium compilospecies intermedium (R. Br.) De Wet &

J.R. Harlan, Amer. J. Bot. 53: 97. 1966. -Type: Australia, R. Brown 6184 (BM,

holo, photo in BRI, K!).

-Andropogon glaber Roxb. [Hort. Bengal.: 7. 1814. nom. nud.] Fl. Ind. 1: 271.

1820. -Andropogon intermedius R. Br. subvar. glaber (Roxb.) Hack. in A.

DC., Monogr. Phan. 6: 487. 1889.-Amphilophis glabra (Roxb.) Stapf in Prain,

Fl. Trop. Afr. 9: 172. 1917. -Bothriochloa glabra (Roxb.) A. Camus, Ann. Soc.

Linn. Lyon I, 76: 164. 1931. -Dichanthium ischaemum (L.) Roberty subvar.

glabrum (Roxb.) Roberty, Boissiera 9: 159. 1960. nom. inval. -Bothriochloa

bladhii (Retz.) S.T. Blake subsp. glabra (Roxb.) B.K. Simon, Austrobaileya

3:79. 1989. -Type: India, Bengal, Roxburgh, s.n .(BM, holo; BR, G; Icon. ined.

1194: CAL, K).

-Andropogon punctatus Roxb., Hort. Beng.: 82. 1814, nom. nud., Fl. Ind.

1: 268. 1820. -Andropogon perfossus Nees & Meyen ex Steud., Nomencl.,

ed. 2, 1: 92. 1840. nom. nud. -Andropogon intermedius R. Br. var. punctatus

(Roxb.) Hack. &. subvar. perfossus Hack. in A. DC., Monogr. Phan. 6: 487.

1889. -Bothriochloa intermedia (R. Br.) A. Camus var. punctata (Roxb.)

Keng, Clav. Gen. Sp. Gram Prim. Sin.: 244. 1957. nom. inval. -Dichanthium

A Revision of Bothriochloa Kuntze (Poaceae) in Thailand 133

Figure 1. Bothriochloa bladhii. A. Habit; B. Spikelet pair; C-J. Sessile spikelet: C-D. Lower

glumes; E.-F. Upper glumes; G. Lower lemma; H. Upper lemma; I. Pistil; J. Stamens; K-O.

Pedicelled spikelet: K-L. Lower glume; M-N. Upper glumes; O. Lower lemma. [Drawing was

based on O. Neamsuvan 166 (BCU)].

134 Gard. Bull. Singapore 61 (1) 2009

ischaemum (L.) Roberty subvar. punctatum (Roxb.) Roberty, Boissiera 9:

160. 1960. nom. inval. -Bothriochloa bladhii (Retz.) S.T. Blake var. punctata

(Roxb.) R.R. Stewart, Kew Bull. 29: 444. 1974. -Lectotype: India, “mountain

grass” Roxburgh s.n. (G, holo), designated by Roberty [Boissiera 9 (1960)

160]; Herb. Hornemann s.n.: C; Roxburgh s.n. ex Herb. Hornemann in Herb.

Trinius 283.1: LE, fragments only, IDC microfiche BT-16/1]; Icon. ined. 892

(CAL, K).

-Andropogon haenkei J. Presl in C. Presl, Reliq. Haenk. 1: 340. 1830.

-Andropogon intermedius R. Br. var. haenkei (J. Presl) Hack. in A. DC.,

Monogr. Phan. 6: 486. 1889. -Bothriochloa glabra (Roxb.) Stapf subsp. haenkei

(J. Presl) Henrard, Blumea 3: 456. 1940. -Dichanthium ischaemum (L.)

Roberty subvar. haenkei (J. Presl) Roberty, Boissiera 9: 159. 1960. nom. inval.

-Lectotype: Philippines, Luzon, Haenke s.n. (PR, holo), here designated.

-Andropogon caucasicus Trin., Mém. Acad. Imp. Sci. St. Pétersbourg, VI, Sci.

Math. 2: 286. 1832.-Sorghum caucasicum (Trin.) Griseb. in Ledeb., Fl. Ross.

4: 476. 1853. -Andropogon intermedius R. Br. var. caucasicus (Trin.) Hack.

in A. DC., Monogr. Phan. 6: 486. 1889. -Bothriochloa caucasica (Trin.) C.E.

Hubb., Bull. Misc. Inform. Kew 1939: 101. 1939. -Dichanthium caucasicum

(Trin.) S.K. Jain & Deshp., Bull. Bot. Surv. India 20: 133. 1979 (“1978”).

-Type: E. Caucasus, Wilhelms s.n. A° 1827 in Herb. Trinius 178.1(0LE, IDC

microfiche BT-16/1).

-Rhaphis stricta Nees in Hooker’s J. Bot. Kew Gard. Misc. 2: 99. 1850.

-Andropogon leptanthus Steud., Syn. Pl. Glumac. 1: 391. 1854, non

Andropogon strictus Host., 1802. -Chrysopogon strictus Nees ex B.D. Jacks.,

Index Kew. 1: 95. 1893; 2: 704. 1895; nom. inval., in syn. -Type: Cuming 1400

(@GE, holo:G,GOET. KE LEW):

-Andropogon intermedius R. Br. subvar. puberulus Hack. in A. DC., Monogr.

Phan. 6: 487. 1889. -Type: Not indicated, material in W to be studied.

-Andropogon pertusus (L.) Willd. var. vegetior Hack. in A. DC., Monogr.

Phan. 6: 481. 1889. -Type: Sudan, Schweinfurth 1027 (W, holo; K!).

-Andropogon odoratus Lisboa, J. Bombay, Nat, Hist. Soc. 4: 123. 1889.

-Amphilophis odorata (Lisboa) A. Camus, Rev. Int. Bot. Appl. Agric. Trop.

1: 305. 1921.-Bothriochloa odorata (Lisboa) A. Camus, Ann. Soc. Linn. Lyon

IT, 76: 165. 1931. -Type: India, Bombay, Lisboa s.n. (BLATT, holo).

-Amphilophis glabra (Roxb.) Stapf. var. paupera Stapf ex Ridl., Fl. Malay

Penins. 5: 209. 1925. -Lectotype: not resolved, material in SING to be

studied.

Culms erect, stout, up to 2 m high, nodes glabrous or pubescent, internodes

terete or grooved on one side. Leaf sheaths terete, keeled in the upper part;

ligules ca 1 mm long; leaf blades up to 43 by | cm, lower surface glabrous,

upper surface scabrous and covered with long hairs at basal part, base

A Revision of Bothniochloa Kuntze (Poaceae) in Thailand 135

subcordate, apex long acuminate, margin scaberulous. Inflorescence a large

panicle, 12-17 by 4-5 cm, axis up to 15 cm long, primary branches whorled,

simple or divided, racemes up to 5 cm long, the lowest raceme shorter

than the central axis, joints 1.5-2.3 mm long. Sessile spikelets elliptic, ca 2.5

mm long, callus 0.2-0.5 mm long: lower glumes elliptic to oblong, greenish

yellow; 2.5-3 mm long, obscurely 7-9-nerved, occasionally 1-pitted, upper

glumes 2.5-3 by 1-1.2 mm, sparsely hairy on upper part of keel, otherwise

glabrous; /Jower lemmas lanceolate, 2-2.5 by 0.5-0.7 mm, glabrous, apex

obtuse; upper lemmas ca 1.5 mm long, awn ca 1.5 mm long; lodicules c. 0.2

mm long; anthers ca 1.5 mm long; pedicelled spikelets 2.5-3 mm long, callus

short; lower glumes lanceolate, ca 3 by 1 mm, 6-nerved, greenish yellow,

pectinately setose on keels, apex acute; upper glumes lanceolate, 2-2.8 by

0.5-0.8 mm, 3-nerved; lower lemmas lanceolate, ca 2 by 0.5 mm, apex acute;

anther ca 1 mm long, sometimes barren.

Specimens examined: S. Legaard & M. Norsangsri 21875 (AAU); K. Larsen,

S.S. Larsen, C. Nivomdham, W. Ueachirakan & P. Sirirugsa 42528 (AAU); K.

Larsen, S.S. Larsen, A.S. Barfod, W. Nanakorn, W. UVeachirakan & P. Sirirugsa

41757 (AAU): S. Legaard & M. Norsangsri 21693 (AAU); S. Legaard 21758

(AAU); J.-E Maxwell 74-597 (AAU, BK); J.-E Maxwell 85-928 (AAU, PSU);

O. Neamsuvan 166 (BCU): J/. Sadakorn 208 (BK): T: Smitinand 3416 (BKF):

M. Lazarides 7445 (BKF, K, L); A. Marcan 1815 (BM); A.EG. Kerr 6955,

7987, 9354, 11334, 13558 (BM.K): Th. Sorensen, K. Larsen & B. Hansen 5481,

5891 (C, K); J.-F Maxwell 92-567 (CMU, P): Ch. Charoenphol, K. Larsen & E.

Warncke 4878 (K): T: Smitinand 2035, 6087 (K); A. Marcan 1589 (K): A.FG.

Kerr 9331, 19781 (K); Put 2049 (BM, K); A.EG. Kerr 3842 (C, K):; K. Larsen

9993 (C.K) G. Murata, N. Fukuoka & C. Phengklai T. 16986 (P).

Ecology: Along roadside, open area, abandon field. Alt. 0-1500 m.

Uses: As a forage grass.

Vernacular name: Ya khaem khok, Ya khi ma

Trade names: Australian bluestem, Forest bluegrass, Long-leaved beard

grass.

Distribution: THAILAND. Northern, Chiang Mai, Sukhothai; North-

Eastern: Loei; Central: Bangkok, Saraburi, Lop Buri, Nakhon Nayok;

Eastern: Nakhon Ratchasima; South-Eastern: Chon Buri, Chantaburi, Sa

Kaeo; Southern: Chumphon, Songkhla, Krabi, Yala. Also tropical Africa

and Asia, introduced elsewhere.

136 Gard. Bull. Singapore 61 (1) 2009

2. Bothriochloa ischaemum (L.) Keng

Contr. Biol. Lab. Chin. Assoc. Advancem. Sci., Bot. 10: 201. 1936; Henrard,

Blumea 3: 457.1940. isonym; Mansf. ex Cuénod., Fl. Tunisie: 56. 1954. isonym.

-Andropogon ischaemum L., Sp. Pl.: 1047. 1753. -Andropogon ischaemum

var. genuinum Hack. in A. DC., Monogr. Phan. 6 (Apr 1889) 475, nom. inval.

-Andropogon digitatus St.-Lag. in Cariot, Etude FI., ed. 8, 2: 898. learly

1889. nom. superfl. -Sorghum ischaemum (L.) Kuntze, Revis. Gen. PI. 2:

792.1891. -Amphilophis ischaemum (L.) Nash, N Amer. FI. 17, 2: 124. 1912.

-Dichanthium ischaemum (L.) Roberty, Boissiera 9: 160. 1960. -Lectotype:

S. Europe, Herb. Linn. 1211.26 (LINN, holo), designated by Marchi &

Longhi-Wagner [Bol. Inst. Bioci. Univ. Fed. Rio Grande do Sul 57: 41. 1998]

= Andropogon gerardii Vitm.; a better choice for stability is Herb. Burser I,

101 (UPS, holo, microfiche IDC 1064), designated by Scholz [in Cafferty, et

al., Taxon 49: 245. 2000]. Fig. 2.

-Andropogon angustifolius Sibth. & Sm., Prodr. Fl. Graec. 1: 47. 1806. -Type:

Greece, Sibthorp s.n. (OXF, holo).

-Andropogon radicans Lehm., Sem. Hort. Bot. Hamburg.: 16. 1827.

-Andropogon ischaemum L. var. radicans (Lehm.) Hack. in A. DC., Monogr.

Phan. 6: 476. 1889. -Type: Cultivated, extant?

-Andropogon ischaemum L. var. songaricus Rupr. ex Fisch. & Meyen, Enum.

Pl. Nov.: 2. 1841. -Bothriochloa ischaemum (L.) Keng var. songarica (Fisch. &

Meyen) Celarier & J.R. Harlan, J. Linn. Soc. Bot. 55: 758. 1958. -Andropogon

ischaemum L. forma songaricus (Fisch. & Meyen) Kitag., Jap. J. Bot. 36: 20.

1961. -Type: Songaria, Schrenk s.n. (LE, holo; K).

-Andropogon ischaemum L. var. fallax Hack. in A. DC., Monogr. Phan. 6:

476. 1889. -Type: Timor (W, holo; “A. annulatus” Kunth Herb. ex p.).

-Andropogon taiwanensis Ohwi, J. Jap. Bot. 12: 652. 1936. -Type: Taiwan,

Shimada 4766 (KYO, holo).

Culms 20-60 cm high, nodes usually bearded. Leaf sheaths 4-6 cm long;

ligules ca 1 mm long; leaf-blades 3-10 cm by 2-4 mm, hairy on both surfaces,

margin scaberulous. Inflorescence subdigitate of 3-8 racemes, axis 0.5-1.5

cm long, the lowest raceme longer than the axis, racemes 4-6 cm long, joints

2-2.5 mm long. Sessile spikelets lanceolate, ca 4 mm long, callus hairy, ca

0.5 mm long; lower glumes lanceolate, 3.8-4 by 0.7-1 mm, 7-nerved, green;

upper glumes oblong, 3.5-4 by | mm, hairy on the upper part of nerves; lower

lemmas lanceolate, ca 3 by 0.5 mm, glabrous, apex acute; upper lemmas ca 2

mm long, awn brown, ca 1.3 cm long, short hairy; lodicules ca 0.3 mm long;

anthers 1-1.5 mm long; pedicelled spikelets ca 3 mm long; pedicel ca 3 mm

long, hairy on both margins; lower glumes oblong, 3 by 0.8-1 mm, 9-nerved,

hairy on upper half part of keel, apex acute; upper glumes oblong, ca 3 by 1

mm, 3-5-nerved, margin ciliolate; lower lemmas obovate, 2-2.5 by 1-1.5 mm,

A Revision of Bothriochloa Kuntze (Poaceae) in Thailand 137

Figure 2. Bothriochloa ischaemum. A. Habit; B. Spikelet pair; C-H. Sessile spikelet: C-D.

Lower glumes; E-F. Upper glumes; G. Lower lemma; H. Upper lemma; I.-N. Pedicelled

spikelet: I. Pedicel; J-K. Lower glumes; L-M. Upper glumes; N. Lower lemma. [Drawing was

based on Y. Sirichamorn 24 (BCU)].

138 Gard. Bull. Singapore 61 (1) 2009

apex obtuse to truncate; anthers 0.8-1 mm long, or barren.

Specimens examined: P. Sirirugsa 71 (BCU); Y. Sirichamorn 24 (BCU);

A.FEG. Kerr 19651, 19701 (K).

Ecology: Open or shady deciduous forest. Alt. 0-300 m.

Uses: Used for erosion control and forage.

Trade names: Old World bluestem, Plains bluestem, Yellow bluestem.

Distribution: THAILAND. Eastern: Buri Ram; Central: Bangkok, Suphan

Buri; South-western: Kanchanaburi. Also from S Europe to China,

introduced elsewhere.

Notes: This has been introduced in Thailand, but apparently it does not

persist. A form with pubescent nodes has been distinguished as var. songarica

(Rupr. ex Fisch. & C.A. Mey.) Celarier & J.R. Harlan. This seems hardly

worth of any recognition.

3. Bothriochloa pertusa (L.) A. Camus

Ann. Soc. Linn. Lyon II, 76: 164. 1931; Maire, Bull. Soc. Hist. Nat. Afrique

N. 31: 45. 1940. isonym. —Holcus pertusus L., Mant. Pl. 2: 301-302. 1771. -

Andropogon pertusus (L.) Willd, Sp. Pl. 4(2): 922. 1806. -Lepeocercis pertusa

(L.) Hassk., Pl. Jav. Rar.: 52. 1848. -Elionurus pertusus (L.) Nees ex Steud.,

Syn. Pl. Glumac. 1: 364. 1854. -Andropogon pertusus (L.) Willd. var. genuinus

Hack. in A. DC., Monogr. Phan. 6: 480. 1889. nom. inval. -Amphilophis

pertusa (L.) Nash ex Stapf, Agric. News (Barbados) 15: 179. 1916; Fl. Trop.

Afr. 9: 175. 1917. - Dichanthium ischaemum (L.) Roberty subvar. pertusum

(L.) Roberty, Boissiera 9: 160. 1960. nom. inval. -Dichanthium pertusum (L.)

Clayton, Kew Bull. 32: 4. 1977. — Lectotype: “India orientalis”. Herb. Linn.

1212.16 (LINN, holo, designated by Clayton, 1977). Fig. 3.

-Andropogon panormitanus Parl., Diar. 9 Congress. Scienz. Ital. Venezia.

sine pag. 1847; Fl. Ital. 1: 140. 1848. -Andropogon pertusus (L.) Willd.

var. panormitanus (Parl.) Hack. in A. DC., Monogr. Phan. 6: 481. 1889.

-Bothriochloa panormitana (Parl.) Pilger in Engler & Prantl, Nat.

Pfanzenfam., ed. 2, 14e: 161. 1940; Brullo, Giorn. Bot. Ital. 129: 173. 1995.

isonym. - Lectotype: Sicily, not resolved.

-Andropogon angustifolius auct. non Sibth. & Sm.: Parl., Fl. Palermo 1: 269.

1845.

A Revision of Bothriochloa Kuntze (Poaceae) in Thailand 139

Figure 3. Bothriochloa pertusa. A. Habit: B. Spikelet pair; C-H. Sessile spikelet: C-D. Lower

glumes; E-F. Upper glumes: G. Lower lemma; H. Upper lemma: I-L. Pedicelled spikelet: I-J.

Lower glumes: K. upper glume: L. lower lemma. [Drawing was based on O. Neamsuvan 167

(BCU)].

140 Gard. Bull. Singapore 61 (1) 2009

Culms up to 80 cm high, grooved on one side, nodes bearded. Leaf sheaths

3-5 cm long; ligules 0.5 mm long, tufted hairs on both sides of ligules 3 mm

long; leaf blades linear, 3-20 cm by 3 mm, sparsely short hairy on both

surfaces, base subcordate to rounded, apex acuminate, margin scaberulous.

Inflorescence digitate of 3-10 racemes, axis 1-3.5 cm long, racemes 4-5 cm

long, the lowest raceme longer than the central axis; joints ca 3 mm long.

Sessile spikelets elliptic to oblong, ca 3 mm long, callus 0.4-0.5 mm long;

lower glumes oblong, 2.5-2.8 by 1 mm, 9-11-nerved, shiny, 1-pitted; upper

glumes oblong, 3-3.5 by 1 mm, glabrous; lower lemmas broadly ovate, 1.5-

2.5 by 0.8 mm, |-nerved, apex obtuse to truncate, margin ciliate at the upper

part; upper lemmas ca 2 mm long, 1-nerved; awn 1.5-2 cm long, short hairy;

lodicules ca 0.2 mm long; anthers 1-1.5 mm long; pedicelled spikelets ca 3

mm long; callus ca 0.5 mm long, hairy; pedicel ca 3 mm long, covered by up to

3.5 mm long hairs on both margins; lower glumes oblong, 3.5 by 1.0-1.2 mm,

10-nerved, dorsally with a purple stripe near apex, 0-3-pitted, apex obtuse;

upper glumes elliptic, ca 2.5 by 1 mm, 3-nerved; lower lemmas obovate, ca 2

by 1 mm, apex acute; anthers 0.8-1 mm long.

Specimens examined: S. Lagaard and M. Norsangsri 21874, 21885 (AAU);

BRD 57 (AAU); K. Larsen, S.S. Larsen, A.S. Barfod, W. Nanakorn, W.

Ueachirakan and P. Sirirugsa 41014, 41566 (AAU); J.-E Maxwell 74-654

(AAU, L); K. Larsen, T: Smitinand and E. Warncke 1112 (AAU, K); O.

Neamsuvan 167 (BCU); C. Chermsirivathana 143, 201 (BK); Put s.n. (BK);

Y. Paisooksantivatana 813-82 (BK); A.EG. Kerr 13432 (BK, BM, K); M.

Lazarides 7434 (BKF, C, K, L); K. Larsen 8304 (C); Th. Sorensen, K. Larsen

and B. Hansen 2061 (C); Th. Sorensen, K. Larsen and B. Hansen 2477 (C, L);

Th. Sorensen, K. Larsen and B. Hansen 2516 (C,L,P); Th. Sorensen, K. Larsen

and B. Hansen 2031 (C, P); G. Murata, K. Iwatsuki and C. Phengklai T-14816

(L, P); A. G. Kerr 3858 (K); Dee 8523 (K); Sommai 63 (The Herbarium of

Department of Botany, Kasetsart University).

Ecology: Open areas, along road sides. Alt. 0-1,500 m.

Uses: Forage, resistant to trampling, drought, and grazing.

Vernacular name: Ya tot lueat, Ya hom, Ya hang ma

Trade names: Pitted bluestem, Pitted bluegrass, Seymour grass, Sourgrass,

Wiregrass.

Distribution: THAILAND. Northern: Nakhon Sawan; North-Eastern:

Phetchabun, Loei Khon Kaen; Eastern: Nakhon Ratchasima, Buri Ram;

A Revision of Bothniochloa Kuntze (Poaceae) in Thailand 141

Central: Bangkok, Lop Buri, Ayutthaya, Samut Prakan, Sarabumn; South-

Western: Prachuap Khiri Khan, Kanchanaburi; Southern: Chumphon,

Songkhla, Trang. S Africa to Thailand, introduced elsewhere.

Note: In Thailand, this species is very similar to B. ischaemum, but it differs

from B. ischaemum by the pit in the lower glume of the sessile spikelet.

Discussion

Because of the genetical complexity, most recent authors, e.g., Chen and

Phillips (2006) have regarded B. bladhii as a polymorphic species that

includes all forms with an elongate inflorescence axis. Yet, they thought that

2 varieties can be distinguished.

la. Lower glume of sessile and pedicelled spikelet without a pit on the back

aR Rs RS eee oe Sedat naw dec vcedvecegdeenneduscedeescusnecestcceensas var. bladhii

1b. Lower glume of sessile and pedicelled spikelet with 1-3 pits on the back

i are oes Saco Sawaal ce snc asennad dsl ccdauciNeassessatbese var. punctata

However, this is not supported by some specimens found in Thailand

where pitted and pitless glumes occurred in the same inflorescence. It is

congruent with Bhutan specimens (Noltie, 2000). Moreover, the many forms

of B. bladhii represent a complex species where all grades of intermediate

forms appear to be present (Celarier and Harlan, 1955), indicating that this

distinction is rather dubious.

For B. ischaemum, it is distinguished from B. pertusa by its lower

glume of the sessile spikelet without a pit. Interestingly, Celarier and Harlan

(1955) stated that glumes of the sessile spikelet in B. ischaemum may be

slender without pits but with a slight tendency toward pitting. It means there

is a gradual transition form to B. pertusa.

Sathagul (1990) reported B. insculpta for Thailand, but her specimen

(Sommai 63) turned out to belong to B. pertusa. From intensively literature

review, the morphological characters of these 2 species are very similar,

and therefore Celarier and Harlan (1955) regarded them as part of the B.

pertusa complex.

As mentioned above, it seems that the species boundary within

Bothriochloa is blurred. Possibly a molecular study of these 3 species

groups, B. bladhii, B. ischaemum and B. pertusa, might clarify taxonomic

delimitations, but in view of the great number of forms and the widespread

distribution of the “species”, it seems unlikely that the seminal experiments

of Celarier and Harlan can be feasibly repeated.

Bothriochloa bladhii and B. pertusa are common and widely

142 Gard. Bull. Singapore 61 (1) 2009

distributed throughout Thailand. On the contrary, B. ischaemum seems to

be rare. Only a few specimens were available in the herbaria and only ona

few occasions were seen in the field. As in Thailand it is not considered to be

a valuable forage grass, there is a risk that it may become extinct.

Acknowledgements

We would like to thank the Center of Excellence in Biodiversity, Faculty of

Science, Chulalongkorn University (CEB_D_11_2006), the 90° * Anniversary

of Chulalongkorn University Fund and the Development and Promotion

of Science and Technology Talents Project of Thailand (DPST) for funding

of this research. We also thank the directors and curators of AAU, BCU,

BK, BKF, BM, C, CMU, K, L, P, PSU and The Herbarium of Department

of Botany, Kasetsart University for making the specimens available for this

study.

References

Bor, N.L. 1965. Studies in the Flora of Thailand: Gramineae. Dansk Botanisk

Arkiv 23: 156.

Bor, N.L. 1968. Flora of Iraq vol. 9: Gramineae. The Ministry of Agriculture

of the Republic of Iraq, Bagdad.

Celarier, R.P. and J.R. Harlan.1955. Studies on Old World Bluestems.

Department of Botany and Plant Pathology and Agronomy, USA.

Chen, S.-L. and S.M. Phillips. 2006. Flora of China. vol. 22, pp. 607-609.

Science Press, Beijing & MBG Press, St. Louis.

Clayton, W.D., K.T. Harman and H. Williamson. 2008. The online world

grasses flora. [Online]: http://www.kew.org/data/grasses-db/sppindex.

htm [2009, August 12].

De Wet, J.M.J. and J.R. Harlan. 1966. Morphology of the compilospecies

Bothriochloa intermedia. American Journal of Botany 53: 94-98.

Kuntze, C.E.O. 1891. Bothriochloa Kuntze. Revisio generum plantarum 2:

762. Felix, Leipzig, etc.

A Revision of Bothniochloa Kuntze (Poaceae) in Thailand 143

Nanakorn, W. and M. Norsangsri. 2001. Species enumeration of Thai

Gramineae, pp. 33-34. Herbarium Queen Sirikit Botanic Garden, Chiang

Mai, Thailand.

Nash, G.V. 1901. Gramineae, pp. 71. In: N.L. Britton (ed.), Manual of the

Flora of the Northern States and Canada. Holt & Co., New York.

Noltie, H.J. 2000. Flora of Bhutan vol. 3(2), pp. 796-798. Royal Botanic

Garden Edinburgh, Scotland.

Roberty, G. 1960. Monographie systématique des Andropogonées du globe.

Boissiera 9: 167.

Sathagul, S. 1990. The taxonomic study of the genus Dichanthium Willemet

and its allies in Thailand. Master’s thesis. Department of Botany, Graduate

School, Kasetsart University.

Soreng, R.J. and S.J. Pennington. 2003. Catalogue of New World Grasses

(Poaceae). 3. Subfamilies Panicoideae, Aristoideae, Arundinoideae, and

Danthonioideae. Contribution of US National Herbarium 46: 84.

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Gardens’ Bulletin Singapore 61 (1): 145-149. 2009 145

A Revision of Hemisorghum (Poaceae, Sorghinae) in

Thailand

O. NEAMSUVAN .J.E. VELDKAMP ANDT. SEELANAN ©

‘Department of Botany, Faculty of Science. Chulalongkorn University.

: Bangkok 10330, Thailand

“National Herbarium Nederland, Leiden University, P.O. Box 9514,

2300 RA Leiden, The Netherlands.

“E-mail: tosak.s@chula.ac.th

Abstract

Hemisorghum C.E. Hubb. (Poaceae, Sorghinae) has a single species in

Thailand: H. mekongense (A. Camus) C.E. Hubb. It occurs from Burma to

Laos and Cambodia. In Thailand it is rare along riverbanks. A lectotype is

designated.

Introduction

According to recent literature the genus Hemisorghum C.E. Hubb. in Bor

(Poaceae, Sorghinae) would contain two species (Clayton and Renvoize.

1986). One, H. venustum (Thwaites) C.D. Clayton, occurs in Sn Lanka and the

Western Ghats of India, the other, H. mekongense (A. Camus) C.E. Hubb.,

in S Burma, NE Thailand, W Laos, and W Cambodia. However, during the

present revision the two turned out to be so different in so many aspects

that the first has recently been placed in a new genus, Lakshmia Veldk.

(Veldkamp, 2009). Clayton and Renvoize (1986) regarded it as basal in the

Sorghinae, but no molecular studies seem to have been done to support this.

At least H. mekongense is very similar to Sorghum Moench:

la. Panicle branches bearing long, lax racemes, rachis internodes and pedicels

scabrid: lower glume of sessile spikelet thinly coriaceous, flat on the back,

margins 2-keeled for most of its length ......0000 Hemisorghum

1b. Panicle branches bearing short, dense racemes, rachis internodes and

pedicels ciliate: lower glume of sessile spikelet coriaceous, convex on the

Baas apically 7-Keeled and wittged 2-2 ca occicccn he eecese ee nses Sorghum

Nanakorn and Norsangsri (2001) is the first and only work where

Hemisorghum mekongense is reported for Thailand. Because this is a

checklist no description was given. The present work extends this record

with a description, notes, and illustrations.

146 Gard. Bull. Singapore 61 (1) 2009

Materials and methods

This study was based on taxonomic literature and a study of herbarium

specimens at the Faculty of Science, Chiang Mai University (CMU), Royal

Botanic Gardens, Kew (K), National Herbarium of The Netherlands, Leiden

University (L), Herbier, Laboratoire de Phanérogamie, Paris (P). Field

work was carried out throughout Thailand with specimens deposited in the

herbarium of the Chulalongkorn University, Bangkok (BCU).

Hemisorghum C.E. Hubb. in Bor, Grass. Burma, Ceylon, India and Pakistan:

686. 1960. — Type species: Hemisorghum mekongense (A. Camus) C.E.

Hubb.

Tufted annuals. Ligule collar-shaped, fringed or ciliolate, membranous;

leaf-blades linear, broad, flat. Inflorescence a lax, espatheate panicle; the

lowermost branches solitary to whorled, tenacious, glabrous; racemes long,

with numerous pairs of sessile and pedicelled spikelets; rachis internodes

and pedicels filiform, scaberulous, articulation transversal. Sessile spikelets

tardily deciduous, with an obtuse, glabrous callus, dorsally compressed,

2-flowered; lower floret epaleate, neuter; upper floret perfect; lower glumes

thinly coriaceous, flat on the back, 7-11-nerved, laterally 2-keeled, the

margins sharply inflexed; upper glumes dorsally rounded, becoming |-keeled

upwards, 7-nerved, margins inrolled; lower lemmas hyaline; upper lemmas

finely l-nerved, awnless; upper palea hyaline or suppressed; lodicules

cuneate, glabrous; stamens 3. Pedicels free of the rachis. Pedicelled spikelets

very much reduced to | or 2 glumes rarely with a much reduced lemma,

barren, deciduous, dorsally compressed, awnless.

Distribution: Monotypic, in Myanmar, Thailand, Laos, Cambodia; one in

Thailand.

Hemisorghum mekongense (A. Camus) C.E. Hubb. in Bor

Grass. Burma, Ceylon, India and Pakistan: 162, 687. 1960. -Sorghum

halepense var. mekongense A. Camus, Bull. Mus. Hist. Nat. (Paris) 25: 497.

1919. -Sorghum mekongense (A. Camus) A. Camus, Fl. Indo-Chine 7: 323,

f. 35. 1922. -Lectotypus: Muong Mai, Laos, Thorel s.n. (P!; designated here).

Fig. 1.

Culms up to 2 m high, nodes minutely puberulous, with prop roots. Leaf-

sheaths terete, distally keeled, 10-15 cm long, glabrous; ligules 1-2 mm

long; leaf-blades 25-60 by 1-4 cm, glabrous, margin scaberulous, apex long-

acuminate. Panicles 30-50 cm long, primary branches 5-15 cm long, branched

A Revision of Hemisorghum (Poaceae, sorghinae) in Thailand 147

Figure 1. Hemisorghum mekongense: A. habitat: B. inflorescence: C. a part of raceme showing

sessile and pedicelled spikelets; D. sessile spikelet dissection: I. lower glume, II. upper glume,

III. lower lemma, IV. upper lemma, V. upper palea, VI. stamens, VII. pistil.

again, bearing 5-10 racemes; racemes 2-6 cm long, each with 2-7 spikelet

pairs; rachis internodes slightly shorter than the sessile spikelet, filiform.

Sessile spikelets 4-5 mm long, incl. callus; lower glumes ovate-lanceolate,

4-5 by 1.5-2 mm, 7- or 9-nerved, puberulous, keels serrate, apex acute:

upper glumes lanceolate, 4-4.8 by 1.3-1.5 mm, 7-nerved, apically 1-keeled,

chartaceous, puberulous, apex acute; lower lemmas ovate-lanceolate, 3.5-4

by ca 1 mm, hyaline, 2-nerved, apex acute; upper lemmas ovate, 2.5-3 by

0.8-1.2 mm; upper paleas narrowly ovate-lanceolate, ca 2.3 by 0.3 mm, apex

narrow, acute: lodicules ca 0.3 mm long; anthers 1.5-1.8 mm long. Pedicels

filiform, 3-4 mm long, 0.6-0.8 times as long as the sessile spikelet, serrulate

on the edges. Pedicelled spikelets usually very much reduced, rarely more

148 Gard. Bull. Singapore 61 (1) 2009

or less developed; lower glumes narrowly ovate-lanceolate, 0.5-4 by 0.5-1

mm, 0-7-nerved, laterally 2-keeled, keels serrulate, chartaceous, glabrous

to sparsely puberulous, margin inflexed, apex acute; upper glumes ovate-

lanceolate, 0.5-4 by 0.6 mm, 0-5-nerved, membranous, apex acute, margin

hyaline, ciliolate.

Specimens examined: MYANMAR: Griffith 6825 (K), Maung Po Khant

13417 (K). THAILAND: Kerr 21356 (K); Neamsuvan 262, 263 (BCU).

LAOS: Thorel s.n. (P); Maxwell 98-477 (CMU, L). CAMBODIA: Maxwell

07-459 (CMU).

Collector’s notes: Annual. Culms tufted, pale green to stramineous. Blades

green to dull dark green above, pale to dull green beneath. Inflorescence

axes green. Spikelets cream, very pale green, with green nerves. Stigmas,

styles pale light green.

Vernacular name: Ya Phong (Nong Khai)

Ecology: Open sandy, weedy area along river, seasonally submerged, 75 m

alt., flowering August-May.

Distribution: Myanmar (Tenasserim), Thailand, Northeastern (Nakhon

Phanom, Nong Khai), Laos (Attopeu, Bolikhamsai, Champasak,

Khammouane, Sayaboury, Vientiane), Cambodia (Kratie).

Notes: In Thailand, Hemisorghum mekongense is similar to Sorghum

halepense (L.) Pers., but the two species can be differentiated in the following

key —

la. Panicles 30-50 cm long. Rachis internode and pedicel serrate. Spikelets

rather slender; lower glume of sessile spikelet 2-keeled throughout, keels

SCADMOUS :c2) han eee ind: Ae Re eee ene H. mekongense

lb. Panicles 10-30 cm long. Rachis internode and pedicel ciliate. Spikelets

stout; lower glume of sessile spikelet becoming 2-keeled near the tip,

keelwwinced, ciltolate eee S. halepense

Acknowledgements

We would like to thank the Development and Promotion of Science and

Technology Talents Project (DPST), the Center of Excellence in Biodiversity,

Faculty of Science, Chulalongkorn University (CEB_D_11_2006) and the

90° Anniversary of Chulalongkorn University Fund for supporting this

A Revision of Hemisorghum (Poaceae, sorghinae) in Thailand 149

study. We are grateful to the curators and staff of K, L,and P for making their

collections available for this study, and to Mr. James F. Maxwell (CMU) for

information on the occurrence of this species in Cambodia. Many thanks to

Miss Angkhana Inta for making photographs of the CMU specimens.

References

Clayton, W.D. and S.A. Renvoize. 1986. Genera graminum: Grasses of the

world: 339. Her Majesty’s Stationery Office, London, U.K.

Nanakorn, W. and M. Norsangsri. 2001. Species enumeration of Thai

Gramineae: 36. Herbarium Queen Sirikit Botanic Garden, Chiang Mai.

Veldkamp, J.F. 2009. Lakshmia (Gramineae), a new genus from Sri Lanka

and W India. Rheedea 18: 81-85.

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Gardens’ Bulletin Singapore 61 (1): 151-157. 2009 151

Corticolous Myxomycetes of Singapore

W. C. ROSING

Department of Biology,

Middle Tennessee State University

Murfreesboro, TN 37132 USA

E-mail: wrosing@mtsu.edu

Abstract

The moist chamber culture technique was employed to detect myxomycetes

(plasmodial slime molds) associated with the bark surface of living trees.

Twenty-five species of myxomycetes in 13 genera were identified from

moist-chambered bark samples collected at three localities in Singapore.

Seventeen species are new records for Singapore. One species, Comatricha

pseudonigra was previously known only from Mitchell River National Park

in the state of Victoria, Australia.

Introduction

All previous published reports of myxomycetes (plasmodial slime molds)

from Malaysia and Singapore have dealt with species that produce fruiting

bodies which are large enough to be detected in the field (Chip, 1921;

Sanderson, 1922; Emoto, 1931; Lister, 1931; Nawawi, 1973). Fruiting bodies

of “corticolous” (bark-inhabiting) myxomycetes are usually too minute to be

detected in nature but can be recovered from the bark of living trees by using

the moist chamber technique of Gilbert and Martin (1933). The purpose

of this study was to survey the myxomycetes of the bark microhabitat in

Singapore.

List of localities

Localities from which bark samples were obtained are listed below. All

samples were collected, in Singapore in October of 2007, by Dr. D.R.

Reynolds, Mycologist at the University of California, Berkeley, while he was

on a Research Fellowship at the Singapore Botanic Gardens.

Locality 1: Singapore Botanic Gardens (SBG)

Locality 2: MacRitchie Reservoir Forest Reserve (MRR)

Locality 3: Bukit Timah Nature Reserve (BT)

152 Gard. Bull. Singapore 61 (1) 2009

Materials and Methods

Samples of dead outer bark were collected from unidentified trees at each

locality at a trunk height of approximately 1.5 m and placed in legal-sized

paper envelopes. Once dried, they were shipped to the author for moist

chamber culturing. There was only enough bark in each sample for a single

moist chamber.

Bark samples were placed on filter paper in disposable Petri dishes

(9 cm diam.) and flooded with sterile distilled water adjusted to pH 7.0

with KOH. Because substrate pH sometimes affects the abundance and

distribution of myxomycetes (Stephenson, 1989; Wrigley de Basanta, 2004),

excess water remaining after 24 hours was poured into clean plastic beakers

where pH was determined using an Orion model 210A pH meter and low

maintenance electrode. Moist chambers were incubated at room temperature

(22-25 C) ina laboratory exposed to diffuse daylight and examined daily for

a period of five weeks, longer if the bark was still producing fruiting bodies

or plasmodia. Small amounts of water were periodically added to each

culture to maintain moist conditions. If fruiting bodies of the same species

of myxomycete appeared more than once on the same moist chambered

bark sample, they were considered to be a single collection.

When mature, fruiting bodies were removed from the moist chambers

along with the bit of substrate upon which they developed. Once air-dried,

specimens were glued into herbarium quality boxes for permanent storage.

Identifications were made using keys by Martin and Alexopoulos (1969)

and Mitchell (2004).

Results

Twenty-five species of myxomycetes, many representing new records

for Singapore, developed from the moist chambered bark. In the list that

follows, myxomycetes are arranged alphabetically by genus and then species.

One species, Comatricha pseudonigra, was previously known only from the

Mitchell River National Park in the state of Victoria, Australia (Moreno et

al., 2007).

Nomenclature of Myxomycetes follows Hernandez-Crespo and

Lado (2005) and uses the conserved names of a number of genera (Lado

et al., 2005) recently approved by the Committee for Fungi (Gams, 2005) of

the IAPT. Collection numbers are those of the author (WCR). Specimens

are curated at SING Herbarium. Where duplicate specimens exist, they are

housed in the MTSU Herbarium (MTSU).

o>)

Corticolous Myxomycetes of Singapore 5

Annotated list of species

Arcyria cinerea (Bull.) Pers., Syn. Fung. (1801) 184. - Locality 1 (SBG):

WCR 9787, 9788 and 9789, at pH 3.4 - 5.4; Locality 2 (MRR): WCR 9790,

9791, 9792 and 9793, at pH 3.4-4.8; Location 3 (BT): WCR 9794, at pH 4.1.

Arcyria minuta Buchet, Pat., Mem. Acad. Malgache 6 (1927) 42. - Locality

1 (SBG): WCR 9795 (slide only), at pH 6. Not previously known from

Singapore.

Arcyria pomiformis (Leers) Rostaf., Mon. (1875) 271. - Locality 1 (SBG):

WCR 9796, at pH 3.4; Locality 2 (MRR): WCR 9797 and 9798, at pH 3.4-4.8.

Not previously known from Singapore.

Clastoderma debaryanum A. Blytt, Bot. Zeit. 38 (1880) 343. - Locality 1

(SBG): WCR 9799, at pH 6; Locality 2 (MRR): WCR 9800, 9801 and 9802, at

pH 4.1-4.8; Locality 3 (BT): WCR 9803, at pH 4.1.

Collaria arcyrionema (Rostaf.) Nann.-Bremek. ex Ing, Ruizia 9 (1991) 26.

- Locality 1 (SBG): WCR 9804 (slide only), at pH 5.4.

Comatricha elegans (Racib.) G. Lister,Guide Brit. Mycet. ed. 3 (1909) 31. -

Locality 2 (MRR): WCR 9806 and 9807, at pH 3.4 - 4.1.

Comatricha pseudonigra G. Moreno, W.C. Rosing, D.W. Mitch. & S.L.

Stephenson, Bol. Soc. Micol. Madrid 31 (2007) 172. - Locality 2 (MRR):

WCR 9808, at pH 3.7. Previously known only from Mitchell River National

Park, Victoria in Australia (37 40°03”S, 149°21°29”E).

Cribraria confusa Nann.-Bremek. & Y. Yamam., Proc. Kon. Ned. Akad.

Wetensch., C. 86 (1983) 212. - Locality 1 (SBG): WCR 9809, at pH 4.6;

Locality 2 (MRR): WCR 9810, 9811, at pH 3.8-4.5; Locality 3 (BT): WCR

9812, at pH 3.8. Not previously known from Singapore.

Cribraria microcarpa (Schrad.) Pers.,Syn. Fung. (1801) 190,emend. Nann.-

Brem. K. Ned. Akad. Wet. Proc. C. 69 (1966) 340. - Locality 2 (MRR): WCR

9813, at pH 4.8. Not previously known from Singapore.

Cribraria minutissima Schwein., Trans. Am. Phil. Soc. II. 4 (1832) 260.

- Locality 2 (MRR): WCR 98/4, at pH 3.4. Not previously known from

Singapore.

154 Gard. Bull. Singapore 61 (1) 2009

Diderma chondrioderma (deBary & Rostaf.) G. Lister in Lister, Mycet. ed.

3 (1925) 258. - Locality 1 (SBG): WCR 9815, at pH 6; Locality 2 (MRR):

WCR 9816, at pH 6.5. Not previously known from Singapore.

Diderma saundersii (Massee) Lado, Cuad. Trab. Fl. Micol. Iber. (2001) 35.

- Locality 1 (SBG): WCR 9817, at pH 4.7; Locality 2 (MRR): WCR 9878, at

pH 4.1. Not previously known from Singapore.

Didymium squamulosum (Alb. & Schwein.) Fr., Symb. Gast. (1818) 19. -

Locality 3 (BT): WCR 9819, at pH 5.7.

Echinostelium minutum de Bary in Rost., Mon. (1874) 215. - Locality 2

(MRR): WCR 9820, 9821, 9822 and 9823, at pH 3.4 — 4.8. Not previously

known from Singapore.

Hemitrichia calyculata (Speg.) M.L. Farr, Mycologia 66 (1974) 887.- Locality

2 (MRR): WCR 9824 at pH 4.8. Not known from Singapore but as noted in

Martin and Alexopoulos (1969), this species is often mistakenly reported as

H. clavata (Pers.) Rostaf., which has been reported from Singapore.

Licea biforis Morgan, J. Cinc. Soc. Nat. Hist. 15 (1893) 131. - Locality 2

(MRR): WCR 9825, at pH 6.5. Not previously reported from Singapore.

Licea eleanorae Ing, Myxomycetes Britain and Ireland (1999) 50. - Locality

| (SBG): WCR 9826, at pH 6.2. Not previously reported from Singapore.

Licea kleistobolus G.W. Martin, Mycologia 34 (1942) 702.- Locality 1 (SBG):

WCR 9827, at pH 4.7. Not previously reported from Singapore.

Licea operculata (Wingate) G.W. Martin, Mycologia 34 (1942) 702. - Locality

| (SBG): WCR 9828, at pH 6.2. Not previously reported from Singapore.

Licea rugosa Nann.-Bremek. & Y. Yamam., Proc. Kon. Ned. Adad. Wetensch.,

C. 90 (1987) 326. - Locality 1 (SBG): WCR 9829, 9830 and 9831, at pH 4.1-7.

Not previously reported from Singapore.

Macbrideola decapillata H.C. Gilbert, Univ. lowa Stud. Nat. Hist. 16 (1934)

158, emend. Alexop., Mycologia 59 (1967) 113.- Locality 1 (SBG): WCR 9832

at pH 6.2; Locality 2 (MRR): WCR 9833 and 9834 at pH 4.8. Not previously

reported from Singapore.

Physarum album (Bull.) Chevall., Fl. Gen. Env. Paris 1 (1826) 336. - Locality

Corticolous Myxomycetes of Singapore 15

3 (BT): WCR 9836, at pH 4.1.

Physarum crateriforme Petch, Ann. Bot. Gard. Peradeniya 4 (1909) 304. -

Locality 1 (SBG): WCR 9837, at pH 7; Locality 2 (MRR): WCR 9825, at pH

6.5. Not previously known from Singapore.

Stemonitis fusca Roth, Mag. Bot. Romer & Usteri | (1787) 26. - Locality 1

(SBG): WCR 9839, 9840 and 9841, at pH 4.4-6.2.

Stemonitis herbatica Peck, Ann. Rep. N.Y. State Mus. 26 (1874) 75; Ing,

Myxomycetes of Britain and Ireland (1999) 202. - Locality 3 (BT): WCR

9838, at pH 4.4.

Discussion

Seventeen myxomycete species are reported here as new records for

Singapore. Most of these species produce minute fruiting bodies, which

at 2 mm or less in total height, are unlikely to be detected in the field. A

similar but more extensive study involving moist-chambered bark samples

from 12 localities in the state of Victoria, Australia (Rosing et al., 2007)

produced twenty-nine new records for Victoria, eight new records for

Australia, and one species (Comatricha pseudonigra) then new to science,

but now known to exist in Singapore. It is unlikely that this species exists

only in two locations, southeastern Australia and Singapore, some 6000 km

apart. Tropical regions like Southeast Asia remain under-investigated for

Myxomycetes. The moist chamber technique for the recovery of corticolous

forms remains under-utilized. Further investigations will undoubtedly show

that C. pseudonigra has an extensive range in the region and that many other

species of myxomycetes remain to be discovered.

Stephenson (1989) noted that while most myxomycete species of

upland temperate forests of Virginia (USA) appeared to tolerate a wide

pH range, members of the Stemonitales tended to develop on more acidic

bark than did members of the Physarales and Trichiales, predominating on

the acidic bark of two species of conifers. The pH range (3.4-7) of bark

samples utilized for the present study is almost identical to that reported by

Stephenson (3.3-7.4) for his Virginia study.

Based on the present study, ten bark samples from Singapore

produced members of the Stemonitales, ten produced members of the

Trichiales, and eight produced members of the Physarales. The mean pH of

bark that produced members of the Trichiales was actually lower (4.35) than

that producing the bark members of the Stemonitales (4.74), though the

pH ranges were almost identical (3.4-6.2) for the bark materials producing

156 Gard. Bull. Singapore 61 (1) 2009

members of the Stemonitales and members of the Trichiales (3.4-6). The

eight bark samples that produced members of the Physarales had the highest

mean pH (5.57) with a pH range of (4.1-7).

The author recently collected myxomycete fruiting bodies, tree

bark, ground and aerial litter from a number of sites in Singapore while

undertaking a research fellowship at the Singapore Botanic Gardens. It will

be interesting to see whether any apparent correlation can be seen between

the bark pH and distribution patterns of corticolous myxomycetes in the

tropical rain forests of Singapore. Definitely, additional new records of

myxomycetes for the island nation will be forthcoming.

Acknowledgements

The author wishes to thank Dr. D.R. Reynolds for collecting the bark

materials used in this study. Appreciation is also extended to Drs. S.L.

Stephenson and D.W. Mitchell for their assistance with the identification of

a number of specimens.

References

Chipp, T.F. 1921.A list of the fungi of the Malay Peninsula. Gardens’ Bulletin

Singapore 2: 311-417.

Emoto, E. 1931. The Malayan myxomycetes. Journal of Botany, British and

Foreign 69: 38-42.

Gams, W. 2005. Report of the Committee for Fungi: 13. Taxon 54: 828-830.

Gilbert, H.C. and G.W. Martin. 1933. Myxomycetes found on the bark of liv-

ing trees. University of lowa Studies in Natural History 15: 3-8.

Hernandez-Crespo, J.C. and C. Lado. 2005. An on-line nomenclatural

information system of Eumycetozoa. See _ http://www.nomen.

eumycetozoa.com

Lado, C., U. Eliasson, S.L. Stephenson, A. Estrada-Torres and M. Schnit-

tler. 2005. (1688-1691) Proposals to conserve the names Amaurochaete

against Lachnobolus, Ceratiomyxa against Famintzinia, Cribraria Pers.

against Cribraria Schrad. ex J.F. Gmel. and Hemitrichia against Hypor-

hamma (Myxomycetes). Taxon 54; 543-545.

Corticolous Myxomycetes of Singapore 157

Lister, G. 1931. Notes on Malayan mycetozoa. Journal of Botany, British and

Foreign 69: 42-43.

Martin, G.W. and C.J. Alexopoulos. 1969. The Myxomycetes. University of

Iowa Press, Iowa City.

Mitchell, D.W. 2004. A key to the corticolous myxomycetes. Systematics and

Geography of Plants 74: 261-285.

Moreno, G., W.C. Rosing, D.W. Mitchell and S.L. Stephenson. 2007.

Comatricha pseudonigra, a new corticolous myxomycete from Australia.

Boletin dela Sociedad Micologia Castellana, Madrid 31: 171-175.

Nawawi, N. 1973. A new species of Licea from Malaysia. Transactions of

British Mycological Society 60: 153-154.

Rosing, W.C., D.W. Mitchell and S.L. Stephenson. 2007. Corticolous

Myxomycetes from Victoria. Australasian Mycologist 26: 9-15.

Sanderson, A.R. 1922. Notes on Malayan mycetozoa. Transactions of British

Mycological Society 7: 239-256.

Stephenson, S.L. 1989. Distribution and ecology of myxomycetes in

temperate forests. I]. Patterns of occurrence on bark surface of living

trees, leaf litter, and dung. Mycologia 81: 608-621.

Wrigley de Basanta, D. 2004. The effect of simulated acid rain on corticolous

myxomycetes. Systematics and Geography of Plants 74: 175-181.

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Gardens’ Bulletin Singapore 61 (1): 159-184. 2009 159

Ten New Species of Impatiens (Balsaminaceae) from

Thailand

P SUKSATHAN' AND P TRIBOUN-

‘Herbarium, Queen Sirikit Botanic Garden, Chiang Mai, Thailand

* Bangkok Herbarium, Princess Sirindhorn Herbarium Building, Plant Variety

Protection Division, Department of Agriculture, Bangkok, Thailand

Abstract

Ten new species of /mpatiens (Balsaminaceae), /. adenioides Suksathan &

Keerat.,/. charisma Suksathan & Keerat.,/. daraneenae Suksathan & Triboun,

I. doitungensis Triboun & Sonsupab, /. jiewhoei Triboun & Suksathan,

I. oreophila Triboun & Suksathan, /. ruthiae Suksathan & Triboun, /.

sirindhorniae Triboun & Suksathan, /. spectabilis Triboun & Suksathan, and

I. tigrina Suksathan & Triboun from Thailand are described and illustrated.

Introduction

The genus /mpatiens consists of over 1,000 species of annual or perennial

herbs widely distributed in tropical and sub-tropical regions of Africa and

Asia. A few members of the genus extend into Europe, and Central and

North America (Fischer, 2004). In contributions to the Flora of Southeast

Asia I, Shimizu (1970) recognized 39 plus 3 doubtful species as native to

Thailand. Subsequently many new species were discovered, and were added

into the account (Shimizu, 1977, 1991, 2000; Shimizu and Suksathan, 2004)

raising the total number of native Thai species to approximately 50. In

connection with a revision of the family Balsaminaceae for the Flora of

Thailand, intensive collecting in recent years in poorly explored mountain

areas as well as material studied in herbaria worldwide have revealed

several more undescribed species. Therefore, our assessment of the species

diversity in the country may be underestimated, and more species probably

await discovery.

In this paper we describes 10 new species using the descriptive

terminology of Grey-Wilson (1980), except for the terms upper pair and

lower pair of lateral sepals for which we prefer to use inner pair and outer pair

respectively. Impatiens sirindhorniae is, by gracious permission, dedicated

to Her Royal Highness Princess Maha Chakri Sirindhorn, in appreciation

of her enormous contributions to the conservation of indigenous Thai

wildlife.

160 Gard. Bull. Singapore 61 (1) 2009

The new species

1. Impatiens adenioides Suksathan & Keerat., sp. nov.

Impatiens mirabilis Hook.f. affinis, a qua differt lamina minori elliptica ad

oblanceolata coriacea, margine acute serrata, bracteis florium eglandulatis —

Typus: S Thailand, cultivated in Bangkok, originally from limestone area in

Phatthalung Province, 15 Apr 2009, K. Keeratikiet & C. Punpreuk 146 (holo,

QBG; iso, BK, SING). Fig. 1 & Plate 1A.

Lithophytic, succulent perennial herbs, up to 100 cm tall, with swollen

caudiform base, up to 30 cm in diameter, wholly glabrous; stems 4-7(-14)

upright, 2-3 cm in diameter at the base, moderately branched in upper part.

Leaves spirally arranged, crowded on the upper part of stem, petiole ca 1 cm

long; lamina elliptic to oblanceolate, 6-8 x 3-4 cm, coriaceous, apex acuminate,

base cuneate, margin acutely serrate, with two glands in the lower half; lateral

veins 7-10(-12) pairs. Inflorescence a stout 6-10(-20)-flowered terminal to

subterminal raceme, simple or 2-3(-6) branched, up to 15 cm long, waxy green;

the branch subtended by a small lanceolate, biglandular, + caducous bract, ca

5 mm long. Flowers waxy yellow, 3-3.4 cm long, pedicels rather thick, ca 0.5

cm long, subtended by a green, ovate, mucronate, caducous bract, ca 4 mm

long; lateral sepals 4, the outer pair rather thick, ca 7 x 5.5 mm, ovate, shortly

mucronate, the inner pair ca 4 mm long, linear-lanceolate, acute, lower sepal

ca 13 mm long, ca 9 mm deep, deeply navicular, entirely yellow, abruptly

constricted into a | cm-long, slender straight spur, incurved only at the tip;

dorsal petal ca 10 x 9 mm, suborbicular, shallowly cucullate, with a green

crest which is broadest in the middle, apex slightly emarginate; /ateral united

petals connate, entirely yellow; upper petals ca 7 x 10 mm, semicircular, apex

round; lower petals connate for about half the length, obovate in outline, ca

12 x 10 mm, apex deeply bilobed, the lobe apically shallowly emarginated.

Ovary glabrous. Fruit ca 2 cm long, clavate; seed unknown.

Additional specimen examined: Thailand, Chumphon Prov., R. Pooma, V.

Cheamchumroon & P. Chantaboon 3514 (BKF).

Distribution and habitat: Endemic to peninsular Thailand (Chumphon,

Phatthalung). Scattered in lowland, shaded, rugged limestone hills.

Etymology: The specific epithet, adenioides refers to its growth habit, which

resembles that of Adenium (Apocynaceae).

Notes: Impatiens adenioides was introduced to the first author by Kaweesak

Keeratikiat and Chuyos Punpreuk, plant lovers from Bangkok who

Ten New Species of Impatiens (Balsaminaceae) from Thailand 161

Figure 1. /mpatiens adenioides Suksathan & Keerat. A. Habit: B, C. Flower: D. Dorsal sepal:

E. Outer lateral sepals: F. Inner lateral sepals: G. Lower sepal: H. Lateral united petals. (B-C

and D-H share the same scale bar). [Drawn by P. Suksathan.]

162 Gard. Bull. Singapore 61 (1) 2009

received the plant from local people in Phatthalung province. Among the

other caudex species found in peninsular Thailand, 1.e., /. mirabilis Hook.

f., and I. opinata Craib, I. adenioides is easily recognized by its elliptic to

oblanceolate, thick-leathery lamina, with acutely serrated margin, and also

its eglandular flower bracts.

2. Impatiens charisma Suksathan & Keerat., sp. nov.

I. phengklaii Shimizu & Suksathan affinis, a qua differt cauli alata, petiolis et

pedunculis planis, flore petalis cum pari superiori lateralibus connatis et per

pares petalorum inferiorium superpositis. — Typus: W Thailand, Kanchanaburi

Province, Sai Yok district, 750 m alt, 5 Aug 2008, P. Suksathan, P. Triboun, H.

Boonnuang & K. Keeratikiat 4679 (holo, QBG; iso, BK, SING). Fig. 2 &

Plate 1B.

Tuberous rooted perennial herbs, completely glabrous, tuber up to 15 cm

in diameter; stems annual, 1-2, erect, unbranched, arising from points at the

top of the tuber, to 20 cm long, 4-6 angled, distinctly winged along stem

angles. Leaves spirally arranged, petiole 3-8 cm long, slightly flat, winged;

lamina up to 21 x 10 cm, elliptic, apex acuminate, base cuneate to rounded,

sometimes oblique, tapering into narrow wings along petiole and stem,

margin shallowly crenate, with two fusiform glands just above the base;

lateral veins 7-10 pairs. Inflorescence (1-) 2-flowered subumbellate raceme,

borne in the upper leaf axils; peduncles up to 10 cm long, flat, terminating

in 3-4, light green, narrowly elliptic to lanceolate, persistent bracts. Flowers

yellow, 4.5-5 cm long, with red tint at spur base; pedicels ca 2.5 cm long,

slender; lateral sepals 4, the outer pair greenish white, ca 1.6 x 1.3 cm long,

obliquely ovate, mucronate, the inner pair strongly reduced into yellow, ca

3 mm long, obliquely ovate scales; lower sepal ca 3 cm long, ca 3.5 cm deep,

bucciniform, abruptly constricted into a narrow, ca 1.5 cm long incurved spur,

the tip of spur minutely bifid; dorsal petal ca 2.3 x 2 cm, slightly cucullate,

broadly obovate to suborbicular, strongly reflexed in the upper half, apex

obcordate, with sub-apical mucro; lateral united petals connate; upper petals

ca 1.6 x 1.3 cm, broadly ovate, short-apiculate at apex; lower petals ca 2.7

x 2.4 cm, obovate, shortly apiculate at apex. Ovary glabrous. Fruit ellipsoid;

seeds unknown.

Flowering during the rainy season, between August and October.

Distribution and habitat: Endemic to W Thailand (Kanchanaburi). Growing

in shaded limestone crevices along cliffs in mixed bamboo forest, ca 750 m

alt., rather rare.

Ten New Species of Impatiens (Balsaminaceae) from Thailand 163

Figure 2. /mpatiens charisma Suksathan & Keerat. A. Habit; B, C. Flower; D. Dorsal sepal: E.

Outer lateral sepals; F. Inner lateral sepals: G. Lower sepal; H. Lateral united petals. (B-C and

D-H share the same scale bar). [Drawn by P. Suksathan.]

164 Gard. Bull. Singapore 61 (1) 2009

Entymology: The specific epithet, charisma - respect and loyalty earned by the

performance of good deeds, is dedicated to our beloved king, His Majesty King

Bhumibol Adulyadej, Rama IX of Thailand, who has been a tireless advocate

for the nature and environment proposing guidelines on the principles of

sustainable conservation and the development of natural resources.

Notes: This rare elegant balsam is one of the only two representative of

tuberous species found in SE Asia (the other 1s /mpatiens phengklaii Shimizu

& Suksathan). In both species, the annual stems die back to the tuberous

caudex at the end of the growing season (May-November). /mpatiens

charisma is easily distinguished from J. phengklaii by having winged stems,

flat petioles and peduncles, and the broader upper pair of lateral united

petals that at the base overlap to the lower one.

3. Impatiens daraneenae Suksathan & Triboun, sp. nov.

Impatiens hongsonensis Shimizu affinis, a qua differt floribus distortis roseis,

sepalo infero angustiore bucciniformi, petalo inferiori connato oblongo. —

Typus: N Thailand, Chiang Mai Province, Chiang Dao district, limestone

outcrops along roadside to Mueang Kong, ca 850 m alt., 7 Oct 2008, P

Suksathan & M. Wongnak 4672 (holo, QBG; iso, AAU, BK, BKF, SING).

Fig. 3 & Plate 1C.

Annual herbs, wholly glabrous; stem decumbent to erect, dark purplish

brown, rather thin, up to 50 cm long, laxly branched, slightly zigzag in upper

part. Leaves alternate, petiole up to 3.6 cm long; lamina up to 6.7 x 3.9 cm,

ovate, apex acuminate, base cuneate to rounded, sometimes oblique, margin

crenate-dentate, with two distinct, short-stalked glands at the base; lateral

veins 6-7 pairs. Flower axillary, solitary or 2-3-fascicled, pendulous under

the leaves, distorted, light pink, 4.0-4.2 cm long; pedicels ca 1.3 cm long,

minutely bracteate at base; lateral sepals 4, the outer pair whitish green,

ca 5 x 5 mm, obliquely broadly ovate, mucronate, the inner pair strongly

reduced into purple, subelliptic scales, ca 2 mm long; lower sepal 1.5 cm

long, 2.5 cm deep, broadly bucciniform, pinkish white, with pink reticulate

veins, clearly visible on the inner surface, abruptly constricted into a narrow,

ca 8 mm long incurved cylindrical spur, the tip of the spur minutely bifid;

dorsal petal ca 1.1 x 0.8 cm, obovate, strongly reflexed in the upper half,

minutely mucronate, base with a short, green, conical crest; lateral united

petals connate, pink, with two yellow patches at the base of lower petals,

unequally distorted; upper petals 7-13 x 8-10 mm, obliquely ovate; lower

petals connate about two-third of the length, distorted oblong in outline,

ca 20 x 12 mm, apex deeply bilobed, the lobe apically emarginated. Ovary

glabrous. Fruit ellipsoid; seeds unknown.

Ten New Species of Impatiens (Balsaminaceae) from Thailand 165

Figure 3. /mpatiens daraneenae Suksathan & Triboun. A. Habit: B, C. Flower; D. Dorsal sepal:

E. Outer lateral sepals; F. Inner lateral sepals: G. Lower sepal: H. Lateral united petals. (B-C

and D-H share the same scale bar). [Drawn by P. Suksathan.]

166 Gard. Bull. Singapore 61 (1) 2009

Flowering during the rainy season, August-October.

Distribution and habitat: Endemic to N Thailand (Chiang Mai). Growing on

limestone outcrop in mixed bamboo forest, ca 850 m alt. Locally common,

but as yet known only from the type locality.

Etymology: The plant is named in honour of Mrs. Daranee Derojanawong, a

plant lover from Bangkok who kindly helped us finding research funding.

Notes: Impatiens daraneenae was firstly discovered by the second author

during his field work in connection with the revision of the family

Gesneriaceae for the Flora of Thailand. It is easily distinguished by its

distorted, light pink flower, narrow bucciniform lower sepal, and oblong

lower connate petals with two basal yellow patches.

4. Impatiens doitungensis Triboun & Sonsupab, sp. nov.

Impatiens lacinulifera Y.L.Chen affinis, a qua differt floribus pupureo-roseis,

sepalis lateralibus laevibus, calcari apice obtusa, petalo dorsaliad basin cristato,

marginis foliorum cum dentibus sinuum prominentibus. — Typus: Thailand,

Chiang Rai Province, Mae Sai-Mae Fa Luang district, Doi Tung, limestone

crevices near summit, 1,300 m alt., 26 Jul 2008, P. Triboun, P. Blenchitra & B.

Sonsupab 3850 (holo, BK; iso, QBG). Fig. 4 & Plate 1D.

Small lithophytic annual herb, completely glabrous; stems light green, erect,

slender, 10-20 cm tall, simple to laxly branched. Leaves alternate, crowed on

the upper part of stem, petiole 7-10 mm long, light green, with many small

red patches; lamina ovate to lanceolate, ca 5.0 x 2.5-2.9 cm, apex mucronate,

base obtuse to cuneate, sometimes slightly oblique, margin serrulate with

prominent sinus-teeth, the two black-tipped basal glands obtuse; lateral veins

6 pairs. Flowers axillary, solitary or 2-fascicled, pendulous under the leaves,

purplish pink; pedicels ca 4 cm long, minutely bracteate at base; lateral sepals

4, the outer pair greenish white to pale pink, ca 7 mm long, broadly ovate,

mucronate, the inner pair strongly reduced into pink small scales, ca 2 mm

long; lower sepal ca 2.4 cm long, ca 2 cm deep, obliquely bucciniform, pale pink,

with red veins, clearly visible on the inner surface, abruptly constricted into a

narrow, |.1-1.3 cm-long, white, incurved cylindrical spur; dorsal petal ca 1.4 x

1 cm, obovate, strongly reflexed in the upper half, apically emarginate, base

with a short, green, subtriangular, pointed crest; lateral united petals connate,

entirely pink except for two yellow patches at the base of lower petals; upper

petals 1.1-1.2 x 0.8 cm, ovate, apiculate; lower petals ca 2 x 1 cm, connate

about three-quarters of the length, narrowly obovate in outline, apex 2 lobed,

the lobe apiculate. Ovary glabrous. Fruit linear, slender; seeds unknown.

Ten New Species of Impatiens (Balsaminaceae) from Thailand 167

Figure 4. Impatiens doitungensis Triboun & Sonsupab. A. Habit: B.C. Flower: D. Dorsal sepal:

E. Outer lateral sepals: F. Inner lateral sepals: G. Lower sepal: H. Lateral united petals. (B-C

and D-H share the same scale bar). [Drawn by P. Suksathan.]

168 Gard. Bull. Singapore 61 (1) 2009

Flowering in rainy season, between July and August.

Distribution and habitat: Endemic to Northern Thailand (Chiang Rai).

Growing in open limestone scrub vegetation, on the summit of limestone

mountain ca 1,300 m alt. As yet known only from the type locality.

Etymology: The specific epithet is named from the type locality at Doi Tung

where it was collected.

Notes: Impatiens doitungensis resembles the Chinese species J. lacinulifera

Y.L.Chen but clearly differs in its purplish pink flower with smooth lateral

sepals, blunt tipped spur, basally crested dorsal petal, and especially a

presentation of prominent sinus-teeth along leaf margin.

5. Impatiens jiewhoei Triboun & Suksathan, sp. nov.

Impatiens nalampooni Shimizu affinis, a qua differt floribus violaceo-

caeruleis sepalo infero rubro-striato, calcari curvato ad basin adnato

cylindrico. — Typus: Thailand, Kanchanaburi Province, Mueang distric, Khao

Pun cave, mixed bamboo forest in shaded limestone area, 11 Oct 2008, P

Triboun, H. Boonnuang & K. Keeratikiat 3893, (holo, BK; iso, QBG, SING).

Fig. 5 & Plate 2A.

Lithophytic annual herb, completely glabrous; stem decumbent to erect,

green to reddish green, rather thin, to 70 cm long, laxly branched, slightly

zigzag in upper part. Leaves alternate, petiole (1-)2-7.5 cm long; lamina 5-12

x 2.5-6.5 cm, ovate to lanceolate, apex acute to acuminate or blunt, base

cuneate to cordate, margin serrate, with two distinct, deltoid glands near

the base; lateral veins (4-)5-6 pairs. Flowers axillary, solitary or 2 fascicled,

pendulous under the leaves, 3-3.2 cm long, violet-blue, with a narrow long,

yellow patch at the base of the lower connate petal; pedicels 0.5-1.5 cm long,

minutely bracteate at base; lateral sepals 4, the outer pair pale green tinged

purple, ca 10 x 5 mm long, obliquely ovate, mucronate, the inner pair strongly

reduced into narrowly lanceolate, ca 3.5 mm-long scales; lower sepal 1.6 cm

long, 1.8 cm deep, broadly saccate, pale pink with red striped veins, clearly

visible on the inner surface, abruptly constricted into a narrow, cylindric,

green, incurved, 1.3 cm long spur, the spur tip dark red; dorsal petal blue, ca

1.9 x 1.7 cm, obcordate, slightly cuculate, strongly reflexed in the upper half,

base with a short, subtriangular crest; lateral united petals connate, violet-

blue, with a narrow longitudinal yellow patch in the middle of the connate

lower petal; upper petals ca 9 x 7 mm, obovate, apex slightly bilobed; lower

petals connate about two-third of the length, broadly ovate in outline, ca 2.3

Ten New Species of Impatiens (Balsaminaceae) from Thailand 169

Figure 5. /mpatiens jiewhoei Triboun & Suksathan. A. Habit; B. Flower; C. Dorsal sepal; D.

Outer lateral sepals; E. Inner lateral sepals; F. Lower sepal; G. Lateral united petals. (B-C and

D-H share the same scale bar). [Drawn by P. Suksathan. ]

170 Gard. Bull. Singapore 61 (1) 2009

x 1.8 cm, apically bilobed, the lobe shallowly emarginated. Ovary glabrous.

Fruit linear, 1.5-2.5 cm long; seeds unknown.

Flowering during the rainy season, between June and October.

Additional specimens examined: Thailand, Kanchanaburi, Kao Tok, below 50

m. alt., 13 Jul 1930, A.EG. Kerr 19549 (BK, K);same province, Tha Salao, ca

100 m alt., 10 Jul 1930, A.FEG. Kerr 19562, (K); ibid., 10 Jul 1930, A. Marcan

2497 (K).

Distribution and habitat: Endemic to W Thailand (Kanchanaburi). Growing

in shaded mixed bamboo to dry evergreen forest in limestone area, 20-100

m alt., locally abundant, but as yet known only from the type locality.

Etymology: The specific epithet is named in honour of Mr. Tan Jiew-Hoe,

the president of Singapore Gardening Society, who kindly encouraged the

authors to revise the genus /mpatiens in Thailand.

Notes: Impatiens jiewhoei is one of the most beautiful species, closely related

to the N Thailand endemic, /. nalampoonii Shimizu. However, the violet-

blue flower with red striped lower sepal, and basally adnate, cylindric, curved

spur, leave no doubt about its separate identity.

6. Impatiens oreophila Triboun & Suksathan, sp. nov.

Impatiens chinensis L. affinis, a qua differt floribus albis, calcari recto vel

leviter curvato ad apicem obtuso pagina interiori venatio rubro-reticulati. —

Typus: Thailand, Kanchanaburi Province, Sangkhla Buri, Khao Radar-Khao

Yai, open grassy summit, 1 ,300-1,400 m. alt., 13 Nov 2008, P. Triboun, W.J.J.O.

De Wilde and B.E.E. Duyfjes 4031 (holo, BK; iso, L, QBG). Fig. 6 & Plate

2B.

Terrestrial or lithophytic annual herb, completely glabrous; stems light

green, unbranched, erect, 15-45 cm tall, with long internodes, quadrangular

in transection. Leaves opposite, decussate, 6-9 pairs, scattered along the stem,

sessile; lamina narrowly lanceolate, (6-)8.5-11.5 x 1.3-1.8 cm, coriaceous,

apex acute or cuspidate, shortly cuneate at base, + amplexicaul, margin

serrate, with two green, retrorse glands at the base; lateral veins 4-6 pairs.

Flower subterminal axillary, solitary or 2 fascicled, ca 3.5 cm long, white;

pedicels slender, green, ca 4 cm long; lateral sepals 2; the outer pair linear,

falcate, white to light green, ca 10 x 1 mm, apex acute; the inner pair absent;

lower sepal navicular, ca 1.4 cm long, 7 mm deep, gradually tapering into

a 1.7-2 cm-long, straight or slightly curved spur, white with red reticulated

Ten New Species of Impatiens (Balsaminaceae) from Thailand imal

Figure 6. Impatiens oreophila Triboun & Suksathan. A. Habit; B, C. Flower: D. Dorsal sepal:

E. Outer lateral sepals: F. Lateral united petals. (B-C and D-H share the same scale bar).

[Drawn by P. Suksathan.]

172 Gard. Bull. Singapore 61 (1) 2009

veins , clearly visible on the inner surface, spur tip blunt, ca 0.5 mm in

diameter; dorsal petal white, ca 8 x 7 mm, obcordate, slightly cuculate, apex

mucronate, base shortly and slightly crested; lateral united petals free, white,

with two violet patches at the base of the lower petals; upper petals ca 2 x

2.5 mm, subtriangular, apex round; lower petals obovate, ca 1.2 x 0.7 cm.

Ovary glabrous. Fruit ellipsoid, 1.5-2.5 cm long; seeds unknown.

Flowering in late rainy season, between August-October.

Distribution and habitat: Endemic to W Thailand (Kanchanaburi), as yet

known only from the type locality. Growing on exposed granite rock crevices

as well as among grasses in open grassland near the summit, from 1,300-

1,400 m alt.

Etymology: The specific epithet oreophila means mountain-loving, which

refers to its habitat near the summit of the high mountain.

Notes: Impatiens oreophila is rather closely related to J. chinensis L. in

having an erect, simple stem, and opposite sessile leaves. However, it is

easily distinguished by its white flower, and its straight or slightly curved,

blunt-tipped spur, with red reticulated veins on the inner surface.

7. Impatiens ruthiae Suksathan & Triboun, sp. nov.

Impatiens nalampoonii Shimizu affinis, a qua differt flori rubro-purpureo

distorto, sepalis lateralibus viridibus, sepalo inferiori luteo-aurantiaco, anguste

bucciniformi. — Typus: Thailand, Loei Province, Nong Hin district, limestone

mountain, 715 m alt., 6 Jul 2008, PR. Suksathan, M. Wongnak, H. Boonnuang

and K. Keeratikiat 4553 (holo, QBG; iso, AAU, BK, BKF, SING). Fig. 7 &

Plate 2C.

Lithophytic annual herb, completely glabrous; stems decumbent to erect,

dark purplish brown, rather thin, to 50 cm long, laxly branched, slightly zigzag

in upper part. Leaves alternate, petiole up to 4 cm long, with two distinctly,

short-stalked, filiform glands at the middle or at the apex; lamina up to 10.5

x 5 cm, ovate, apex acuminate, base cuneate to rounded, sometimes oblique;

lateral veins 5-7 pairs; margin crenate-dentate. Flower axillary, solitary or

2-3-fascicled, pendulous under the leaves, distorted, ca 4 cm long, purplish

red, with orange-yellow lower sepal; pedicels ca 1.3 cm long, minutely

bracteate at base; lateral sepals 4, the outer pair whitish green, ca 8 mm long,

broadly ovate to suborbicular, mucronate, the inner pair strongly reduced

into subelliptic, 0.5 mm-long scales; lower sepal ca 1.5 cm long, 2.2 cm deep,

narrowly bucciniform, orange-yellow, with reticulate red veins, clearly

Ten New Species of Impatiens (Balsaminaceae) from Thailand 173

Figure 7. Jmpatiens ruthiae Suksathan & Triboun. A. Habit: B, C. Flower; D. Dorsal sepal; E.

Outer lateral sepals: F. Inner lateral sepals; G. Lower sepal: H. Lateral united petals. (B-C:

D-H share the same scale bar). [Drawn by P. Suksathan].

174 Gard. Bull. Singapore 61 (1) 2009

visible on the inner surface, abruptly constricted into a narrow, cylindric, ca

5 mm long incurved spur; dorsal petal pink, ca 1.4 x 1.1 cm, obovate, slightly

cuculate, strongly reflexed in the upper half, apex emarginate, forming a

subapical mucro, base with a short, green, subtriangular crest, ca 1.5 mm high;

lateral united petals connate, purplish red, marked with a broad yellow patch

at the base of the lower petals, unequally distorted; upper petals 1-1.2 x 1.3-

1.4 cm, obliquely ovate, apex rounded; lower petals 1.7-2 x 1-1.1 mm, oblong,

apically emarginated. Ovary glabrous. Fruit ca 15 x 4mm, clavate, constricted

in the upper part; seeds ca 1.5 mm long ellipsoid, seed coat pustulate.

Flowering during rainy season, July-September.

Distribution and habitat: Endemic to NE Thailand (Loei). As yet known

only from the type locality. Growing on limestone in moist, shaded limestone

foothill at ca 715 m alt., rather rare.

Etymology: The plant is named in honour of Dr. Ruth Kiew in Malaysia, a

botanist and a limestone plant expert who has supported this project.

Notes: Impatiens ruthiae is one of the most peculiar species with its burgundy

red flowers, a colour rarely found in other Asian species.

8. Impatiens sirindhorniae Triboun & Suksathan, sp. nov.

Impatiens walleriana Hook.f affinis, a qua differt caulibus decumbentibus

usque ad pendentibus, foliis griseo-viridis ovatis coriaceis, floribus calcaribus

profunde navicularibus. — Typus: Thailand, Krabi Province, Plaai Phraya

district, vertical limestone cliff, 20-150 m alt., 25 Jun 2007, P. Triboun, P.

Blenchitra and P. Yothakaew 3801 (holo, BK, iso, AAU, QBG, SING). Fig.

8, Plates 2D, 3C & D.

Lithophytic perennial herb, forming basal tillers, up to 15 stems, wholly

glabrous; stems succulent, decumbent to pendulous, greyish to brownish

green, glaucous, to 40 cm long, rarely branched. Leaves spirally arranged,

somewhat crowded on the upper part of stem, petiole 6-7.5 cm long, with two

distinct, short-stalked glands at the apex; /amina 3.2-4 x 2.8-3.5 cm, ovate,

thickly coriaceous, greyish green, apex acute, base truncate to subcordate,

margin roughly serrate; lateral veins 3-6 pairs. Flowers axillary, solitary,

rarely 2-fascicled, large, rather flat, 4.5-4.8 cm wide, light purple; pedicels

slender, suberect, 3-6.5 cm long, + minutely bracteate at base; lateral sepals

4, the outer pair whitish green, 6-7 x 3-5 mm long, obliquely ovate, acute,

the inner pair strongly reduced into whitish green, cordate to rotund, ca

2.5 mm scales; lower sepal 1.6 x 1.6 mm, deep navicular, pale to whitish

Ten New Species of Impatiens (Balsaminaceae) from Thailand 175

Figure 8. /mpatiens sirindhorniae Triboun & Suksathan. A. Habit: B. C. Flower: D. Dorsal

sepal: E. Outer lateral sepals: F. Inner lateral sepals: G. Lower sepal: H. Lateral united petals.

(B-C and D-H share the same scale bar). [Drawn by P. Suksathan.]

176 Gard. Bull. Singapore 61 (1) 2009

green, gradually constricted into a narrow, cylindric, curved, ca 6 cm long

spur; dorsal petal, ca 2 x 2.5 cm, depressed obovate, strongly reflexed just

above the base, apex cuspidate, base with a short, white, subtriangular crest,

ca 2 mm high; lateral united petals connate, forming 3 subequal lobes, which

are also subequal to the dorsal one; upper petals ca 2.5 x 2 cm, obovate, apex

cuspidate; lower petals connate for about two-third of the length, depressed

obovate in outline, ca 2.0 x 2.5 cm, apically bilobed. Ovary glabrous. Fruit

ellipsoid; seeds unknown.

Additional specimen examined: Thailand, Surat Thani Province, Kao Ken, 50

m alt.,27 Mar 1927, A.FEG. Kerr 12446 (BK, K).

Flowering in rainy season, between June-October.

Distribution and habitat: Endemic to peninsular Thailand (Krabi and Surat

Thani). Growing on open or shaded vertical limestone cliffs, 20-150 m alt.

Etymology: The specific epithet is given in honour of Her Royal Highness

Princess Sirindhorn for her dedication and encouragement to the

environmental sciences and biodiversity conservation in Thailand.

Notes: Impatiens sirindhorniae is one of the most beautiful perennial species,

easily recognized by its pendulous habit, succulent thick leaves and large

light purple flower.

9. Impatiens spectabilis Triboun & Suksathan, sp. nov.

Impatiens walleriana Hook.f. affinis, a qua differt herba annua, androeceo

et gynoecio occultis per bases petalorum, cum apertura angusta, sepalis

latioribus. — Typus: Thailand, Kanchanaburi Province, Thong Phaphum,

Wat Tha Khanun, on limestone, ca 135 m alt., 10 Oct 2008, P. Triboun, H.

Boonnuang and K. Keeratikiat 3889, (holo, BK; iso, BKF, K, QBG, SING).

Fig. 9 & Plate 3A.

Moderately branched, terrestrial or lithophytic, annual herb; stems

decumbent to erect, fragile, up to 40 cm long, often rooting at the lower

nodes. Leaves alternate, scattered along the stems, petiole 1-3.5 cm long,

dull red, finely pubescent to glabrescent; lamina 5-11 x 2-5.5 cm, ovate to

lanceolate, apex acute to acuminate, base cuneate to subcordate, sometimes

oblique, margin serrate, with two distinct, short-stalked, clavate glands near

the base; lateral veins 6-8 pairs. Flowers axillary, solitary, rarely 2 fascicled,

large, flat, ca 4 cm wide, bright pink; pedicels slender, suberect, ca 3 cm long,

minutely bracteate at base; lateral sepals 2(-4), the outer pair dull red, 6-10

Ten New Species of Impatiens (Balsaminaceae) from Thailand 7,

x 6 mm long, obliquely broadly ovate, mucronate, the inner pair absent,

or strongly reduced into small scales, less than 1 mm long; lower sepal

shallowly navicular, 14mm long, 6 mm deep, white to reddish green, abruptly

constricted into a narrow, cylindric, curved, 4.5-4.8 cm long spur; dorsal petal

ca 1.3 x 1.8 cm, obovate, minutely apiculate, strongly reflexed just above the

base, base with a short, red, subtriangular crest, ca 2 mm high; lateral united

petals partly connate, forming 4 subequal lobes, which are also subequal to

the dorsal one; upper petals ca 1.7 x 1.3 cm, obovate, minutely apiculate;

lower petals connate for about half of the length, slightly obliquely obovate,

ca 2.3 x 1.5 cm, minutely apiculate; the base of all petals positioned closed to

each other forming a small aperture hiding the androecium and gynoecium

underneath. Ovary glabrous. Fruit clavate; seeds many, 1.5-2.1 « 1-1.2 mm.

Additional specimens examined: Thailand, Kanchanaburi Province, Thong

Pha Phum distric, Kwai river valley, ca 100 m alt.,21 Oct 1984, /. Dransfield

6218 (K); same district, near Neeckey, near Wangka, 150 m alt., 13 Jun 1946,

G. den Hoed 945 (K); same district, I-Thong, along gas pipe-lines, 21 Nov

1997, K. Chayamarit 1002 (BKF); same district, Kao Leam dam, 29 Nov

1982, K. Hiroshike, T: Hiroshi, C. Nivomdham, & T: Wongprasert T-30452

(BKF); same locality, 10 Jul 1981, 7. Santisuk s.n. (BKF).

Flowering in rainy season, between July-October.

Distribution and habitat: W Thailand (Kanchanaburi). A very common

species forming large colonies on mountain slopes and limestone outcrops

in lowland dry evergreen and mixed deciduous forests along roadside from

Thong Phaphum to Sangkla Buri district. Also observed in Myanmar.

Etymology: The specific epithet refers to its spectacular large pink flower

that are easily spotted from a distance.

Notes: A very common but peculiar species, /mpatiens spectabilis has a very

characteristic flower, flat, with a large dorsal petal, and lateral united petals

almost equal in size and shape, and especially the androecium and gynoecium

that are hidden behind a very narrow aperture formed by petal bases. It

would be very interesting to study its floral morphology and pollination.

10. Impatiens tigrina Suksathan & Triboun, sp. nov.

Impatiens charani Shimizu affinis, a qua differt flore luteo, sepalis lateralibus

libris, calcari simplici. — Typus: Thailand, Udon Thani Province, Ban Paeu

district, shaded sandstone table, ca 300 m alt., 8 Jul 2008, P. Suksathan, M.

Wongnak, H. Boonnuang and K. Keeratikiat 4567 (holo, QBG; iso, BK,

178 Gard. Bull. Singapore 61 (1) 2009

Figure 9. /mpatiens spectabilis Triboun & Suksathan. A. Habit; B, C. Flower; D. Dorsal sepal;

E. Outer lateral sepals; F. Lower sepal; G. Lateral united petals. (B-C and D-H share the same

scale bar). [Drawn by P. Suksathan. |

Ten New Species of Impatiens (Balsaminaceae) from Thailand 179

SING). Fig. 10 & Plate 3B.

Lithophytic annual herb, with a green, erect cylindrical main stem, to 30 cm

long, ca 1.3 cm in diameter, richly branched at the top, completely glabrous;

branches of stem up to 30 cm long, purplish red or purplish brown, rather

thin, up to 30 cm long, slightly zigzag in upper part. Leaves ovate to broadly

lanceolate, apex acuminate, base cuneate to round, sometimes oblique, with

two distinct, short-stalked, fusiform glands at the base; lateral veins 12-

15 pairs; margin crenate-dentate; the basal ones larger, spirally arranged,

petiole 8-10 cm long and lamina up to 25 x 10 cm in basal leaves; the upper

leaves congested on top of the main stem, smaller, alternately arranged

along branches, petiole 5-10 mm long and lamina 7-16 x 2.5-6 cm. Flowers

axillary, solitary or 2-3-fascicled, pendulous under the leaves, yellow, ca 4

cm long, with orange-red transverse stripes on lower sepal: pedicels ca 1

cm long, minutely bracteate at base; lateral sepals 4, the outer pair pale

green, ca 10 x 7 mm long, obliquely broadly ovate, mucronate, the inner pair

strongly reduced into small sublanceolate, light green scales, ca 2 mm long;

lower sepal ca 1.6 cm long, 2 cm deep, broadly bucciniform, with orange-red

transverse stripes, clearly visible on the inner surface, abruptly constricted

into a narrow, 8 mm-long incurved claviform spur, swollen toward the tip;

dorsal petal yellow, ca 1.6 x 1.2 cm, cuculate, obovate, retuse, with a thick,

green, blunt crest in the lower half; lateral united petals connate, entirely

yellow; upper petals ca 1.5 x 1.6 cm, depressed orbicular; lower petals

connate, suborbicular in outline, ca 1.8 x 2 cm, with a prominent basally bi-

lobed keel, apex emarginate. Ovary glabrous, 4-carpellate. Fruit subfusiform,

ca 2 cm long; seeds many, ca 3 mm long, oboviod.

Flowering in rainy season, between July-September.

Distribution and habitat: Endemic to NE Thailand (Udon Thani), as

yet known only from the type locality. Growing on sandstone in shaded

sandstone tables in dry evergreen forest ca 300 m alt.

Etymology: The specific epithet, tigrina, means like a tiger, referring to its

yellow flower with red striped lower sepal.

Notes: Impatiens tigrina was first introduced to the first author by Mr. Methee

Wongnak, a geographer from Queen Sirikit Botanic Garden. Its growth

habit strongly resembles /. charanii Shimizu, but the yellow flower with free

outer lateral sepals and simple spur leave no doubt of its identity.

180 Gard. Bull. Singapore 61 (1) 2009

Figure 10. /mpatiens tigrina Suksathan & Triboun. A. Habit; B, C. Flower; D. Dorsal sepal; E.

Outer lateral sepals; F. Inner lateral sepals; G. Lower sepal; H. Lateral united petals. (B-C and

D-H share the same scale bar). [Drawn by P. Suksathan.]

Ten New Species of Impatiens (Balsaminaceae) from Thailand 181

Acknowledgements

The authors wish to gratefully acknowledge H. R. H. Princess Maha Chakri

Sirindhorn for kindly granting us the permission to name the new /mpatiens

after her. We are grateful to Dr. Benjamin Ollgaard for his kind suggestions

and for translating the Latin diagnoses. Thanks are due to the curators and

staff of the following herbaria: AAU, BK, BKF, BM, K, P, QBG, and SING for

making material available for study and also to Methee Wongnak, Chuyos

Punpreuk, Hassachai Boonnuang, and Kaweesak Keratikiat for supplying

specimens and participating in the field. Special thanks are also given to

Tan Jiew-Hoe and Daranee Derojanawong for financial support and kind

advice.

References

Fischer, E. 2004. Balsaminaceae. pp. 20-25. In: K. Kubitzki (editor), The

Families and Genera of Vascular Plants, Vol. 6. Flowering Plants:

Dicotyledons. Celastrales, Oxalidales, Rosales, Cornales, Ericales. Springer,

Berlin.

Grey-Wilson, C. 1980. Impatiens of Africa. Balkema, Rotterdam. 235 pp.

Shimizu, T. 1970. Contributions to the Flora of Southeast Asia I. /mpatiens

of Thailand and Malaya. Southeast Asian Studies 8(2): 187-217.

Shimizu, T. 1977. Some additional note on /Jmpatiens (Balsaminaceae) of

Thailand. Acta Phytotaxonomica et Geobotanica 23(1-3): 31-34.

Shimizu, T. 1991. New species of the Thai /mpatiens (1). The Journal Japanese

Botany 66: 166-171.

Shimizu, T. 2000. New species of Thai /mpatiens (Balsaminaceae) 2. Bulletin

of the National Science Museum, Series B (Botany) 26(2): 35-42.

Shimizu T. & Suksathan, P. 2004. Three new species of the /mpatiens

(Balsaminaceae). Part 3. Bulletin of the National Science Museum, Series

B (Botany) 30(4): 165-171.

Gard. Bull. Singapore 61 (1) 2009

Plate 1. A. Impatiens adenioides Suksathan & Keerat.; B. /. charisma Suksathan & Keerat.;

C. I. daraneenae Suksathan & Triboun; D. /. doitungensis Triboun & Sonsupab. (Photos: H.

Boonnuang).

> 79

Ten New Species of Impatiens (Balsaminaceae) from Thailand 183

Plate 2. A. Impatiens jiewhoei Triboun & Suksathan; B. /. oreophila Triboun & Suksathan;

C. I. ruthiae Suksathan & Triboun; D. I. sirindhorniae Triboun & Suksathan. (Photos: H.

Boonnuang).

Gard. Bull. Singapore 61 (1) 2009

Plate 3. A. Impatiens spectabilis Triboun & Suksathan; B. J. tigrina Suksathan & Triboun;

C-D. Impatiens sirindhoriae Triboun & Suksathan (Photos: H. Boonnuang).

Gardens’ Bulletin Singapore 61 (1): 185-188. 2009 185

A New Species of Alphonsea (Annonaceae) from Borneo

I.M.TURNER

Research Associate, Singapore Botanic Gardens and

Royal Botanic Gardens, Kew

Abstract

Alphonsea borneensis 1.M. Turner is described as a new species. It is a

medium-sized tree of lowland forest recorded from Sabah and Central

Kalimantan.

Introduction

In revising the genus A/phonsea for the Tree Flora of Sabah and Sarawak,

it became apparent that a number of distinctive fruiting specimens in

various collections, notably the Sandakan Herbarium, did not match any of

the species included in the revision of the genus by Kessler (1995). A new

species is therefore described.

Alphonsea borneensis 1.M. Turner, sp. nov.

Alphonsea javanicae similis, foliis chartaceis, fructibus maioribus differt. —

Typus: Borneo, Kalimantan, Central Kalimantan, Sintang, HPH km 83-87,

along old logging road east of camp (0°49'25.8", 112°3'38.8"), 6 May 1994,

A.C. Church et al. 1344 {holo, K (barcode: K000580481); iso, A [x2], BO, K,

L[x2]]. Figs. 1 & 2.

Tree to 16 m tall, 20 cm dbh. Twigs pale grey or grey-brown, drying

longitudinally wrinkled, often with raised lenticels, youngest twigs red-brown

with adpressed pale hairs. Leaves chartaceous, drying fairly uniformly brown

or grey-brown, glabrous, except for short hairs along midrib above; midrib

flush to slightly sunken above, prominent below in dry leaves, lateral nerves

slightly raised on both surfaces in dry leaves, lamina ovate to (narrowly)

elliptic, 4-13 x 1.5-5 cm, base acute to obtuse, apex acuminate, lateral nerves

10-18 pairs, obscure to the naked eye but visible under magnification, looping

well within the margin with secondary loops outside, tertiary and higher

orders of venation visible under magnification from below, more obscure

from above; petiole 2-5 mm long, 0.5-1 mm thick, drying black or dark brown,

rugose. Flowers unknown. Fruiting pedicel, 8-18 mm long, 3-3.5 mm thick,

calyx sometimes persisting. Monocarps to 6, ellipsoidal to globose, to 6 x 4

186 Gard. Bull. Singapore 61 (1) 2009

ere |

ROM, BOTANIC GAMDENS HWY

HERB. HORT. KEW. HAVEDNET OO

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Church, AC. Mamryar, U.W. India and Hamuran

Fociay, May 08. 1804

HERBARIUM BOGORIENSE

HARVARD UNIVERSITY HERBARIA

Figure 1. Photograph of the herbarium specimen that forms part of the holotype of Alphonsea

borneensis, sp.nov.© The Board of Trustees of the Royal Botanic Gardens, Kew. Reproduced

with the consent of the Royal Botanic Gardens, Kew.

A New Species of Alphonsea (Annonaceae) from Borneo 187

37.0_[INNONACERE (5.0)

Alphonse $-

KR H ANTAL Claryrola (3 toy

Figure 2. Photograph of the carpological specimen that forms part of the holotype of

Alphonsea borneensis, sp. nov.

cm, drying black, apex rounded or with short broad nipple, smooth, minutely

warty sometimes with very short brown hairs visible under magnification,

stipe 3-10 mm long, 6 mm thick. Seeds ca 8, lunar, ca 20 x 10 x 4mm.

Specimens examined: MALAYSIA, Sabah. Ulu Sungai Pingas-Pingas,

Keningau District, Interior Division, Sumbing Jimpin, SAN 122061 (SAN);

Upper Sg Miau near Mt Muruk Miau, Sipitang District, Interior Division,

Pius & Dauni, SAN 143151 (SAN); Hiaw Fok Logging Area, Marak-Parak

on the National Park Boundary, Kota Merudu District, Kudat Division,

Aban Gibot, SAN 100010 (SAN [x2]); Sepilok, Sandakan District, Sandakan

Division, de Wilde et al. SAN 144001 (K,SAN); Permodalan Plantation, Sg

Kertam, Sandakan District, Sandakan Division, Aban Gibot, SAN 96834

(SAN [x2]); Mostyn, Madai Forest Reserve, Lahad Datu District, Tawau

Division, S. Lantoh, SAN 67746 (SAN); Madai, Kunak District, Tawau

Division, Mansus & Suali,SAN 117726 (SAN); about 8 miles from Kampung

Merungin, Ranau District, West Coast Division, Leopold & Saikeh, SAN

82589 (SAN).

188 Gard. Bull. Singapore 61 (1) 2009

Note: This is a species of lowland forest, with one collection (SAN 100010)

from lower montane forest on ultrabasic substrate at 1200 m. Quite widely

collected in Sabah, once from Central Kalimantan (type).

A pertinent question is why, in the absence of flowers, is this species

described in the genus A/phonsea? The arborescent form and multi-carpellate

and multi-ovulate state of the specimens limits the choice of genus among

those occurring in Borneo to Alphonsea and its near miliusoid relatives

Mitrephora and Pseuduvaria. Species of the latter two genera typically

have leaves with regular ‘herring-bone venation’ of distinct sets of lateral

nerves arching in parallel, whereas Alphonsea borneensis has fairly obscure

venation, very similar to other Bornean Alphonsea species. A number of the

specimens listed above have been previously determined as ‘A/phonsea sp.’

The large monocarps of Alphonsea borneensis that dry black

and apparently glabrous to the naked eye are distinctive. The only other

species of Alphonsea in Borneo with fruits similar is A. javanica Scheff.,

but it has more cylindrical monocarps to 2.5 cm in diameter whereas those

of A. borneensis are ellipsoidal, or more rarely globose, and reach 4 cm in

diameter. The obvious difference between A. javanica and A. borneensis is

in the leaves. Alphonsea javanica has coriaceous leaves with a shiny upper

surface, whereas A. borneensis has chartaceous leaves with a smooth but not

markedly shiny upper surface. The foliage of A. borneensis 1s very similar to

that of A. kinabaluensis J. Sinclair but the monocarps are much larger and

not brown hairy.

Acknowledgements

I am grateful to the Arnold Arboretum, Forest Research Institute of

Malaysia and Singapore Botanic Gardens for financial assistance towards

this research and Royal Botanic Gardens, Kew, for facilities essential to its

completion. Dr J. F Veldkamp (L) kindly translated the diagnosis into Latin.

John Sugau, Joan Pereira and Suzana Sabran are thanked for their help

during my visit to Sandakan and for the subsequent loan of material.

Reference

Kessler, P. J. A. 1995. Studies on the tribe Saccopetalae (Annonaceae) —

IV. Revision of the genus A/phonsea Hook.f. & Thomson. Botanische

Jahrbiicher fiir Systematik, Pflanzengeschichte und Pflanzengeographie

118: 81-112.

Gardens’ Bulletin Singapore 61 (1): 189-204. 2009 189

A History of Cananga (Annonaceae)

I.M.TURNER AND JE VELDKAMP>

‘Research Associate, Singapore Botanic Gardens and

Royal Botanic Gardens, Kew

Correspondence: Fairfield, Pett Level Road, Winchelsea Beach,

, East Sussex TN36 4ND, UK

National Herbarium of the Netherlands, Leiden University

P.O. Box 9514, 2300 RA Leiden, The Netherlands

Abstract

The history of the name cananga in botanical nomenclature is outlined

and clarified. Cananga odorata (Lam.) Hook.f. & Thomson, is lectotypified.

Cananga brandisiana (Pierre) I.M. Turner is proposed for Cananga latifolia

(Hook.f. & Thomson) Finet & Gagnep., nom. superfl. The infraspecific

taxonomy of Cananga odorata (Lam.) Hook.f. & Thomson is reviewed and

the cultivar group names for plants producing ylang-ylang and cananga oil

are corrected.

Introduction

Cananga odorata (Lam.) Hook.f. & Thomson (Annonaceae) is a tree native

to the tropical Indo-Pacific. It has long been cultivated for its strongly

perfumed flowers and is grown widely in the tropics, including in plantations

for the production of essential oil. The species and its genus have a long

and rather confusing nomenclatural history that has yet to be resolved

completely. In this paper we outline that history and attempt to clarify some

of the issues and to rectify some of the mistakes.

Etymology and Pre-Linnean Taxonomy

Vernacular names for Cananga odorata in two Asian languages have been

used internationally in both common and scientific nomenclature. The

Malay name kenanga (or cananga in pre-standardised spelling) has been

borrowed for the generic name. The Tagalog name alang-ilang or ilang-ilang

(now the Filipino standard, B.C. Tan, pers. comm.) has been used widely in

the Spanish spelling variant ylang-ylang as a common name for the plant

or more particularly for the essential oil, which is widely used in perfume

manufacture.

190 Gard. Bull. Singapore 61 (1) 2009

The earliest mention of the species now known as Cananga

odorata in the western scientific literature is in the account of the trees

of Luzon island in the Philippines by Georg (Jiri) Josef Kamel (or Camel

or latinised to Camellus or Camelli). This was published as an appendix,

Herbarium aliarumque stirpium in insula Luzon e Philippinarum, to John

Ray’s third volume of Historia Plantarum (Ray, 1704). Kamel, a Moravian

Jesuit missionary and apothecary, referred to alanguilang (variant spelling

of ilang-ilang) under Zhampacae species (p. 83). Kamel’s manuscript was

accompanied by pen-and-ink drawings of most of the species. The originals

of these are divided between the Maurits Sabbe Library, Catholic University

of Leuven, Belgium, and the Natural History Museum, London. There are

two copies of the drawing for alanguilang in a bound volume in the Sir Hans

Sloane collection at the Botany Library of the Natural History Museum. As

Figure | shows, they very clearly depict Cananga odorata.

Georg Everhard Rumphius was the first to employ the name cananga

in botanical writings. He had started the manuscript for what later became

the Herbarium amboinense around 1663, for in August of that year he wrote

a request to the Lords XVII, the Board of the Dutch East Indian Company

(VOC), in which he announced that he had started on a work in which plants

and animals of ‘India’ (Dutch East Indies) would be described and politely

requested that the necessary literature be shipped by VOC vessels from

Amsterdam to Amboina ( Veldkamp, 2002, p. 12). Because of the political and

commercial sensitivity of the subjects treated in the subsequent six-volume

manuscript the VOC suppressed it until around 1735, and the publication of

the first volume edited by Johannes Burman appeared in 1741.

In the second volume of his Herbarium Amboinense, Rumphius

(1741) refers to cananga, clearly deriving the name from the Malay bonga

cananga (bungah kenanga [kenanga flower] in current spelling). Cananga,

cananga domestica or cananga vulgaris as it is referred to in the text and

legend to the plate (tab. 65) respectively, is clearly Cananga odorata, though

the four-part calyx in the drawing is erroneous. Rumphius also described

cananga sylvestris (p. 197), a wild form, including under it three entities.

None of Rumphius’s three ‘varieties’ of cananga sylvestris has been directly

linked to Cananga odorata (Merrill, 1917). Cananga sylvestris trifolia (p.

197; t. 66, f. 1) is possibly a species of Goniothalamus and is the sole basis

for Uvaria tripetala Lam. (=Unona tripetaloides Dunal, Unona tripetala

(Lam.) DC.). Cananga sylvestris angustifolia (p. 197; t. 66, f.2) has yet to be

identified with certainty but has been validated as Uvaria ligularis Lam. (=

Unona ligularis (Lam.) Dunal). Finally, cananga sylvestris latifolia (p. 198)

is the basis of Unona latifolia Dunal, the basionym of Fissistigma latifolium

(Dunal) Merr.

Francois Valentijn, the son-in-law of Rumphius, used parts of the

A History of Cananga (Annonaceae) 19]

Herbarium Amboinense manuscript in producing his Oud en Nieuw Oost-

Indién. This work was influential for nearly two centuries as a general guide

to the Far East and was long used in the training of the employees of the

Dutch East Indies Civil Service. Cananga appears for the first time in print in

volume 3 (Valentijn, 1726, p.213) where it is described (Appendix | contains

an English translation of the text) and figured (tab. 42, see Figure 2). There

can be little doubt that Valentijn described Cananga odorata.

The genus Cananga

Aublet (1775, p. 607, t. 244) was the first botanist to use Rumphius’s name

at generic rank. He mistakenly considered his new species from South

America, Cananga ouregou now Guatteria ouregou (Aubl.) Dunal, to

belong to the same genus as Rumphius’s entities. As none of the Rumphian

elements was validated when Aublet published Cananga, the genus must be

typified by Cananga ouregou [ICBN (McNeill et al., 2006) Art. 10.3, Ex. 3].

Unfortunately Cananga Aubl. pre-dated Guatteria Ruiz & Pav. (1794) and

so had priority over it as the correct name for the very large neotropical

genus. Rafinesque (1815, p. 175) employed Cananga as a generic name but

without any description or indication of its derivation.

In his taxonomic treatment of Unona, Dunal (1817) recognised

two ranks of infrageneric taxa without formal indication. One of the lower

subdivisions containing nine species, including Unona odorata (Lam.) Dunal,

he called Cananga. Later in the same year de Candolle (1817) followed

Dunal’s classification, using Cananga in exactly the same way as Dunal,

though he named both the taxon (Unonaria) and rank (section) of the level

above Cananga. It has generally been considered that Cananga here would

be at subsectional rank (viz. McNeill et al.,2006; Jessup, 2007) but de Candolle

did not indicate a definite rank anywhere in the work. Blume (1830) appears

to have been the first to use Cananga for a definite infrageneric rank when

he referred to Uvaria section Canangae. As he stated (p.12) that his sections

were based on Dunal, this can be regarded as an orthographic variant to be

corrected to Cananga.

Hooker and Thomson (1855) included a single species, Cananga

odorata (Lam.) Hook.f. & Thomson, when they described the genus Cananga

Rumph. ex Hook.f. & Thomson. Despite the absence of a direct reference to

Dunal, Hooker and Thomson must be considered to have transferred Dunal’s

name to generic rank [ICBN Art. 33.3, viz. Ex. 9]. However, the new genus

was a later homonym of Cananga Aubl. Baillon (1868) clearly recognised

this and proposed Canangium as a replacement name [Cananga Rumph.,

nec Aubl.], but he used it for a section of Unona (p. 213), not at generic rank

as later authors have assumed. It was King (1892) who first used Canangium

192 Gard. Bull. Singapore 61 (1) 2009

for the name of a genus recognising Canangium odoratum (Lam.) Baill. ex

King and describing a second species, Canangium scortechinii, which has

subsequently been reduced to the synonymy (Corner, 1939) of Cananga

odorata.

y a8.

C fy NArberis Manguilang ermuce

as , ry eee

: f= cf fructus rR cosng, g

fA, Bn Ca

f ewe eee acne S |

D5, a8 Pre ees

‘

M4,

85.2.

Figure 1. Drawing of alanguilan from G.J. Kamel’s manuscript of Herbarium aliarumque

stirpium in insula Luzon e Philippinarum © Natural History Museum, London.

A History of Cananga (Annonaceae) 193

The Nineteenth and early Twentieth Centuries saw confusion

among botanists regarding the application of the generic name Cananga. It

was used both in the sense of Guatteria Ruiz & Pav. and Cananga, though

latterly in the period Canangium was generally used for Asian taxa. As early

as the International Botanical Congress in Brussels in 1910 (Briquet, 1912),

Guatteria was proposed and accepted for conservation against Cananga

Aubl. Later Cananga Hook.f. & Thomson was formally conserved against

Cananga Aubl. (Lanjouw et al., 1952). Conservation of a name leads to

automatic rejection of homotypic names of the same rank [ICBN Art. 14.4]

so Canangium Baill. ex King is rejected as well.

There isstillconfusion andinaccuracy among the majornomenclatural

references (Farr et al., 1979; Greuter et al., 1993; van Setten and Maas,

1999; McNeill et al/., 2006) on the citation of Cananga and its synonyms, its

typification and the status of Canangium. The latter is often referred to as an

orthographic variant of Cananga implying that combinations in Canangium

can be corrected to Cananga, but as we have shown above it was clearly

introduced as a substitute name and was not a spelling mistake. Below we

give the full citation of the generic name.

Cananga (Dunal) Hook.f. & Thomson, FI. Ind. (1855) 129, nom. cons., non

Cananga Aubl., nom. rejic.

Basionym: Unona [unranked] Cananga Dunal, Monogr. Anonac. (Aug-Nov

1817) 96. — Typus: Canangium odoratum (Lam.) Baill. ex King (lectotype,

designated by Hutchinson, 1923) [Uvaria odorata Lam., Unona odorata

(Lam.) Dunal, Cananga odorata (Lam.) Hook.f. & Thomson].

Homotypic synonyms:

-Uvaria section Cananga (Dunal) Blume, Fl. Javae Anonaceae (1830) 13 as

‘Canangae’.

-Unona subsection Cananga (Dunal) G. Don, Gen. Hist. 1 (1831) 94.

-Unona section Canangium Baill., Hist. Pl. (Baillon) 1 (1868) 213, nom.

superfl.

-Unona section Cananga (Dunal) Pierre, Fl. Forest. Cochinch. (1881) t. 19.

-Canangium Baill. ex King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 61 (1892)

39, nomi. rejic.

Heterotypic synonym:

-Fitzgeraldia F. Muell., Fragm. (Mueller) 6 (1867) 1. -Type: Fitzgeraldia

mitrastigma F. Muell., Fragm. (Mueller) 6 (1867) 1.

Gard. Bull. Singapore 61 (1) 2009

194

Figure 2. Drawing of a branch of the Cananga-boom from Francois Valentijn’s Oud en Nieuw

Oost-Indién.

The species of Cananga (Dunal) Hook.f. & Thomson

The first published post-Linnean mention of Cananga odorata is in

the account of the Endeavour voyage by John Hawkesworth (1773).

Hawkesworth used the travel journals of Captain James Cook and the

naturalist Sir Joseph Banks to write the popular account of the expedition.

A section on the natural products available in Batavia (Jakarta) in Java,

‘Some account of Batavia’, is only very slightly changed from the passage in

Banks’s journal (see Hooker, 1896; Banks, 1980; or http://nla.gov.au/nla.cs-

A History of Cananga (Annonaceae) 195

ss-jrnl-banks_remarks-353 ef seq.) and includes the following entry:

‘The Cananga, or Uvaria Cananga, is a green flower, not at all

resembling the blossom of any tree or plant in Europe: it has indeed more

the appearance of a bunch of leaves than a flower, its scent is agreeable, but

altogether peculiar to itself.

There are no specimens of Uvaria cananga in the Banks herbarium

(BM). However, Uvaria cananga appears in the unpublished manuscript

of plants of Java (Plantae Javanenses) by Daniel Solander in the Botany

Library of the Natural History Museum, London. The entry reads “Uvaria

cananga Fig Pict. Cananga Rumph. Amb. 2 p. 195 t. 65. Valent. ind. n°. 42.

‘Fig Pict.’ refers to an outline drawing (pencil with watercolour) of Uvaria

cananga by Sydney Parkinson (J3 of Diment et al., 1987). The reference to

Rumphius’s description and illustration of Cananga and Valentijn’s plate

(Figure 2) and the presence of the Parkinson drawing whichis labelled Uvaria

cananga in pencil leaves little doubt that Banks and Solander intended a

new species to include the Rumphian element cananga and using the name

as the specific epithet in the Linnean genus Uvaria. Because of the rule on

tautonymy [ICBN Art. 23.4], this epithet cannot be employed in Cananga

and the entirely overlooked Uvaria cananga poses no threat to the widely

used Cananga odorata.

The first extensive description of Rumphius’s cananga was by

Lamarck (1785, p. 595) in his account of Uvaria odorata. He referred to

three elements, in order: Cananga. Rumph. Amb. 2 p. 195. tab 65; Alanguilan

de la Chine, Sonnerat, and Arbor saguisan. Raj. Suppl. Luz. 83. The first is

Rumphius’s description of cananga. The second is a herbarium specimen

collected by Sonnerat. The third refers to Kamel’s Luzon trees, but Lamarck

made a mistake in citing arbor saguisan, or arbor sagnisan as Kamel actually

spelled it. This is Goniothalamus amuyon (Blanco) Merr. not Cananga

odorata, which Kamel described on the same page under Zhampacae

species. The identification of arbor sagnisan as Goniothalamus amuyon and

not Cananga odorata is supported by Kamel’s use of amuyong or amoyong

as the vernacular name of arbor sagnisan, clearly the same name as Blanco

(1837, p.463) employed for the specific epithet of Uvaria amuyon; the details

of Kamel’s description, e.g. scentless flowers; and the presence of a Kamel

specimen labelled amuyong in the Sloane Herbarium (Figure 3) which

is Goniothalamus amuyon. Thanks to Lamarck and a paper by Fliickiger

(1881), ‘arbor saguisan’ is often cited, we now know wrongly, as the earliest

name for Cananga odorata. Below Uvaria odorata Lam., the basionym of

Cananga odorata, is lectotypified by the Sonnerat specimen. The origin of

this specimen remains uncertain. It has generally been assumed to be from

China, but then why did Sonnerat employ a Filipino name for the plant?

A second species of Cananga, ranging from Burma, through

196 Gard. Bull. Singapore 61 (1) 2009

Indochina, to the seasonal parts of northern Malay Peninsula, was first

described by Hooker and Thomson (1872, p. 60) as Unona latifolia. Pierre

(1881, t. 19) noted that this was a later homonym of Unona latifolia Dunal,

the basionym of Fissistigma latifolium (Dunal) Merr. Pierre provided a

substitute name Unona brandisiana Pierre, honouring Sir Dietrich Brandis,

the collector of the type. Surprisingly it seems to have escaped attention that

Pierre’s name has not been transferred to Cananga as it has priority over

the widely used Cananga latifolia |Hook.f. & Thomson] Finet & Gagnep.

An ay oy ;

Figure 3. Specimen of amuyong (volume HS 233, p. 81) from the Sloane Herbarium (BM-

SIE):

1. Cananga odorata (Lam.) Hook.f. & Thomson, FI. Ind. (1855) 130.

Uvaria odorata Lam., Encycl. (Lamarck) 1 (1785) 595. Unona odorata (Lam.)

Dunal, Monogr. Anonac. (1817) 45, 93, 97, 108, 143 as ‘Uvaria odorata’;

Uvaria javanica Thunb. [Widmark], Fl. Jav. 2 (1825) 14, 19, nom. superft.;

Canangium odoratum (Lam.) Baill. ex King, J. Asiat. Soc. Bengal, Pt. 2, Nat.

Hist. 61 (1892) 41. — Type: ?China, Sonnerat s.n., annot. Alanguilan de la

Chine (lectotype, designated here, P-LAM, barcode no. P00286083).

-Canangium scortechinii King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 61 (1892)

42. -Type: Perak, Scortechini 1925 (lectotype, designated here, K, iso, BM,

TCALSSING):

-Fitzgeraldia mitrastigma F. Muell., Fragm. (Mueller) 6 (1867) 1.-Canangium

mitrastigma (F. Muell.) Domin, Biblioth. Bot. 89 (1925) 670. -Type: Australia,

Queensland, Rockingham Bay, J. Dallachy. [MEL has several specimens

from the type location, including Dallachy specimens labelled Cananga

odorata and poor specimens from the von Mueller herbarium annotated

A History of Cananga (Annonaceae) 197

‘Fitzgeraldia mitrasacmoides’ and ‘Fitzgeraldia calyptrostigma’ with no

indication of the collector].

-Unona ? leptopetala Dunal, Monogr. Anonac. (1817) 98, 114, 143. Type:

Timor, Leschenault (?G,?P) Depicted in Delessert (Icon. Sel. 1 (1821) 23, t.

88).

-Unona odorata auct. non (Lam.) Dunal: Blume, Bijdr. (1825) 14. Unona

cananga Spreng., Syst. Veg. 4(2) (1827) 215. -Type: Java, Kuhl & van Hasselt

in Herb. Blume s.n. (lectotype, designated here, L, sheet no. 898.60-258).

-Unona odoratissima Blanco, Fl. Filip. (1837) 467. -Type: Philippines, Luzon,

Province of Rizal, Antipolo, M. Ramos 466 |Merrill: Species Blancoanae

466] (neotype, designated here, US, barcode no. 688555, isotypes, A, B, BM,

BOWCALE GH; KL; MO;NSW, NY, PU, UC, W).

-Unona ossea Blanco, FI. Filip. (1837) 467. Uvaria ossea (Blanco) Blanco, FI.

Filip. ed. 2 (1845) 322. -Type: Philippines, no material extant.

-Uvaria axillaris Roxb., [Hort. Bengal. (1814) 94, nom. nud.] Fl. Ind. ed. 1832

(Roxburgh) 2 (1832) 667. -Type: Mauritius, Grand Rivere, M. Rosselle’s

Garden, Hardwicke 6 (lectotype, designated here, BM, barcode no. BM

000895029).

-Uvaria cananga Banks in Hawkesworth, Voyages in the Southern

Hemisphere 2 (1773) 742. -Type: Outline drawing Uvaria cananga (del.

S. Parkinson) J3 (Diment ef al., 1987, reproduced on microfiche 1 of Java

Drawings) (lectotype, designated here, BM). Anon., cultivated tree in BO

no X1.B (iii) 20 (L, epitype, designated here, sheet no. 932.47--467; iso, BO).

[Dr. Irawati, Director of the Kebun Raya Bogor, informed us that in April

2008 the tree was still living].

-Uvaria cananga auct. non Pers.: Pers., Syn. Pl. 2 (1806) 94, excl. Aubl. ref.

-Uvaria farcta Wall., Numer. List (1831) no. 6460, nom. nud. Voucher: Burma,

Kogun near the river Salween, N. Wallich 1283B [Herb. Wall. no. 6460] (K-

W), 21 March 1827.

-Uvaria hortensis Norona, Verh. Batay. Genootsch. Kunsten 5 (1790) 28,

nom. nud.

-Uvaria? subcordata Mig., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 9.

-Canangium odoratum var. velutinum |Blume]| Koord. & Valeton, Meded.

Lands Plantentuin 61 (1903) 282 (Bijdr. Boomsoort. Java 9 (1903) 282)

as ‘velutina’. -Type: Java, [near Kuripan, not found in L]. Reinwardt s.n.

(neotype, designated here, L, sheet no. 898.63-533). [Leaves and branch only.

Labelled as ‘Uvaria velutina’ by Blume, and annotated by Miquel as Uvaria

? subcordata Miq., and ‘Uvaria velutina Bl. non Dun.’, ‘Jav? Moluccae?’].

Pekalongan, Soebah, Koorders 22570 (epitype, designated here, L; iso BO),

11 May 1896.

-Uvaria trifoliata Gaertn., Fruct. Sem. Pl. 2 (1790) 156, t. 114 f. 2. -Uvaria

gaertneri Dunal, Monogr. Anon. (1817) 40, 41, 87, 89, nom. superfl. -Type: E

198 Gard. Bull. Singapore 61 (1) 2009

collect. sem. hort. lugdb. (not located in B, L, TUB); Gaertner’s illustration

(t. 114 f.2) (lectotype, designated here). Java, Udjong Kulon Nature Reserve,

Nenga Wirawan 8 (epitype, designated here, L (barcode no. L0185668); iso,

A, BO, K, LAE, P, SING), 15 April 1963.

-Uvaria undulata cited erroneously in Index Kewensis (Index Kew. 2(4)

(1895) 1161) attributed to Lam., Encycl. 1 (1785) 595; a mistake possibly

originating with de Candolle [Prodr. 1 (1824) 90].

2. Cananga brandisiana (Pierre) I. M. Turner, comb. nov.

Basionym: Unona brandisiana Pierre, Fl. Forest. Cochinch. (1881) t. 19 as

‘brandisana’. — Type: Brandis s.n. [K, holotype (not found), isotype, CAL],

Burma, Tenasserim, Martaban, hill forests of the Saluen (Salween) River.

-Canangium brandisianum (Pierre) Saff., Bull. Torrey Bot. Club 39 (1912)

504 as ‘brandesianum’.

-Unona latifolia Hook.f. & Thomson, FI. Brit. India 1 (1872) 60, non U.

latifolia Dunal.; Cananga latifolia [Hook.f. & Thomson] Finet & Gagnep.,

Bull. Soc. Bot. Fr. Mém. 4 (1906) 84. Canangium latifolium [Hook.f. &

Thomson] Pierre ex Ridl., Fl. Malay. Pen. 1 (1922) 44, nom. superff.

Infraspecific taxa of Cananga odorata

Despite its cultural and economic importance there has only been one,

rather limited, attempt to provide a coherent taxonomic framework for

infraspecific variation in Cananga odorata (van Steenis in Koolhaas, 1939).

Here we can only review the possibilities and names available for various

entities.

It is probable that Cananga odorata is typified by a cultivated tree as

Sonnerat’s label noted the use of the flowers for pommade. Though scarcely

distinguishable morphologically, the wild tree, at least in parts of its range,

may differ from the widely cultivated tree in characteristics such as flower

scent (Corner, 1939). The only infraspecific name that might be employed

for a wild form seems to be Canangium odoratum var. velutinum Koord. &

Valeton, though Koorders and Valeton (1903) appeared to use this variety

to distinguish specimens with more densely pubescent foliage than between

wild against cultivated taxa. The weakness of this variety is reflected by

Koorders omitting it from his Excursionsflora von Java (Koorders, 1912) and

the very brief mention (not keyed) in Flora of Java (Backer and Bakhuizen

van den Brink, 1964, p. 105).

Among cultivated Cananga odorata there are several entities which

have been named. There is a widely grown garden shrub that is a dwarf

form that seems never to set seed. This was first described from cultivated

material in Thailand by Craib as Canangium fruticosum and subsequently

A History of Cananga (Annonaceae) 199

transferred to variety and forma ranks. Although probably never formally

validated, the name “Cananga fruticosa” is found in horticultural catalogues

and on the Internet.

There are two main commercial products from Cananga odorata,

ylang-ylang oil and cananga oil. Ylang-ylang production developed in the

Philippines and has been commercialised most notably in Madagascar

and islands of the Indian Ocean (Yusuf and Sinohin, 1999). Cananga oil

is a traditional product from Java and as it is inferior to ylang-ylang in

perfumery it seems not to have been grown much outside Java. In a paper

by Koolhaas (1939) on the cultivation of Cananga odorata (referred to as

Canangium odoratum) in Java, van Steenis presented an annotated key for

the material giving names at the rank forma for various entities. Ylang-ylang

trees were referred to forma genuinum, with some complication of subforms,

and cananga-oil trees to forma macrophyllum. Unfortunately, as published

after 1 January 1935, all but one of the new combinations are invalid in the

absence of a Latin diagnosis [ICBN Art. 36.1]. Recently Yusuf and Sinohin

(1999) have used van Steenis’s formae as the basis for names in horticultural

terms, transferring them to cultivar groups with forma genuinum being

referred to as Ylang-ylang Group and forma macrophyllum as Cananga

Group. This seems a sensible approach as clearly these plants are cultigens,

however nomenclaturally the change of name was unwarranted and the use

of Cananga as a cultivar name, or part thereof, is prohibited under the genus

Cananga [ICNCP (Brickell et al., 2004) Art. 20.5 (viz. Art. 19.23)]. ICNCP

does not preclude the use of cultivar names based on names published prior

to 1 January 1959 invalid under ICBN [ICNCP Art. 19.7], so we can use van

Steenis’s formae to provide replacement group names.

A list of infraspecific taxa of Cananga odorata

Cananga odorata var. fruticosa (Craib) Sincl., Sarawak Mus. J. 5 (1951)

599. -Canangium fruticosum Craib, Bull. Misc. Inform. Kew 1922 (1922) 166.

-Canangium odoratum var. fruticosum (Craib) Corner, Gard. Bull. Straits

Settlements 10 (1939) 15. -Canangium odoratum forma pumilum Steenis

in Koolhaas, Landbouw (Buitenzorg) 15 (1939) 590 as ‘pumila’. — Type:

Thailand, Bangkok, Kerr 4435 (lectotype, designated here, BM, barcode no.

000546881(BM), iso, ABD), 29 April 1920.

Cananga odorata var. odorata

Cananga odorata Genuina Group

-Canangium odoratum forma genuinum Steenis in Koolhaas, Landbouw

(Buitenzorg) 15 (1939) 590 as ‘genuina’, nom. inval. sub. ICBN Art. 36.1.

-Cananga odorata Ylang-ylang Group, Yusuf & Sinohin in Oyen & X. D.

200 Gard. Bull. Singapore 61 (1) 2009

Nguyen, PROSEA 19 (1999) 71, 73

Cananga odorata Macrophylla Group

-Canangium odoratum forma macrophyllum Steenis in Koolhaas, Landbouw

(Buitenzorg) 15 (1939) 590 as ‘macrophylla’, nom. inval. sub. ICBN Art.

362

-Cananga odorata Cananga Group, Yusuf & Sinohin in Oyen & X. D.

Nguyen, PROSEA 19 (1999) 71, 73

Acknowledgements

Thanks to Dr Robert Vogt (B), Dr P. Lakshminarasimhan (CAL), Dr Pina

Milne and Nimal Karunajeewa (MEL), Cornelia Dilger-Endrulat (TUB) for

information on specimens in their respective herbaria, Dr Irawati (Kebun

Raya Bogor) for the presence of Cananga in the Garden, Judy Warnement

and Dr Kanchi Gandhi (HUH) for continuing assistance, and John Hunnex

(BM) for help with the Sloane Herbarium.

References

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A History of Cananga (Annonaceae) 201

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the Asiatic Society of Bengal, Part 2. Natural History 61: 1-130.

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204 Gard. Bull. Singapore 61 (1) 2009

Appendix 1. Translation of the description of ‘Cananga-boom’ from

Valentijn’s Oud en Nieuw Oost-Indién.

The flowering trees

The Cananga-tree

In this country there are also large trees which at certain times produce

very beautiful fragrant flowers. One of these is the Cananga-tree, which is

rather high, beautiful of trunk (which sometimes may be a fathom around),

and greyish of bark.

The leaves are in rows on long twigs arranged against each other,

about as long as wide, similar to a peach leaf, acute, and full of ribs.

The blossom, or flowers, show themselves here and there between the leaves,

in bunches, well similar to a somewhat yellow or orange ribbon, which hangs

down, consisting of six long narrow weak leaflets , about a little finger long,

and as wide.

Her true colour at first is pale yellow, having on top as a small

peppercorn, from which the fruit comes.

They are very strong in scent, which they spread along the whole road where

this tree stands, especially towards the night; yet their colour is the most

pleasant.

Most of the flowers fall without producing a fruit (a few excepted), and

were collected, for nosegays for the young damsels, to carry them in their hair.

The native dries it, too, to smoke it under his tobacco, or to eat them

with betel-nut, putting a leaflet with the betel quid.

They also serve to decorate with them the vegetable-bows at the

large festivals, and especially an oil has been extracted, which is very strong,

fragrant, and useful against many ailments originating from frigidness. The

natives also use this oil below their betel-nut, also they mix it together with

their bobori’.

The wild does not differ much from the tame, of which we show a

branch on No XLII.

' certainly refers to ‘petals’. cf. Banks’s description of Uvaria cananga

* petal

* Sundanese name for coloured, fragrant ointment, applied to brides, grooms, and against

diseases.

Gardens’ Bulletin Singapore 61 (1): 205-216. 2009 205

Miscellaneous Cucurbit News III

W.J.J.O. DE WILDE AND B.E.E. DUYFJES

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

2300 RA Leiden, The Netherlands

Abstract

The miscellaneous notes on Cucurbitaceae comprise: (1) the description

of a new species in Kedrostis from Peninsular Malaysia; (2) three new

combinations in Neoachmandra from Africa, five new combinations in

Pilogyne from New Guinea and the Pacific; and (3) a modern description of

Zehneria baueriana, from Norfolk Isl., western Pacific.

Introduction

Now that the preliminary studies in Cucurbitaceae for the treatment in

Flora Malesiana come to an end,some more corollary results are presented

here. Previous miscellaneous notes were published in Reinwardtia in 2007

and 2009. A recent, second collection of a Kedrostis in central Peninsular

Malaysia, deviating from the widespread K. bennettii (Miq.) W.J.de Wilde &

Duyfjes, prompted us to recognize a new species. Thus, the following three

items are addressed here:

(1) Kedrostis monosperma W.J.de Wilde & Duyfjes, a new species from

Peninsular Malaysia.

(2) New combinations in African Neoachmandra, and Pilogyne from

Malesia and the Pacific.

(3) Assessment on the morphology of the male flower of Zehneria baueriana

Endl., the type species of the genus Zehneria Endl.

(1) A new species of Kedrostis from Peninsular Malaysia.

Kedrostis monosperma W.J.de Wilde & Duyfjes, sp. nov.

Kedrostidi bennettii similis, infructescentibus ramosis breviter pedunculatis,

fructibus plus gracilibus ovato-oblongis monospermis nitidis (i.s.) differt. —

Typus: Peninsular Malaysia, Pahang, Gunung Benom game reserve, Ulu

Krau, 23 Apr 1967, Rahim Ismail KEP 100114 (holo, KEP; iso, K, L, SING).

Figs. 1, 2.

Climber 5-7 m long, glabrous (except ovary), cystolyths not obvious;

monoecious. Tendrils unbranched. Leaf petioles 1.5-2 cm long; blade (sub)

entire, (broadly) ovate-oblong, 9-13 by 4-7.5 cm, glabrous at both surfaces,

Gard. Bull. Singapore 61 (1) 2009

Figure 1. Kedrostis monosperma W.J.de Wilde & Duyfjes. A. Female inflorescence with

flowers and young fruits; note hairy ovary; B. Infructescence with one ripe fruit. (Both photos

taken at Pahang, Ulu Krau, Rezab Hidupan Liar Krau by de Wilde).

Miscellaneous Cucurbit News III 207

Figure 2. Kedrostis monosperma W.J.de Wilde & Duyfjes. A. Apex of branch with female

inflorescences with immature and mature fruits; B. Node with male inflorescence (all flowers

already fallen off); C. Node with compound female inflorescence with female flower buds

and immature fruits. (A & C based on Siti Munirah et al. FRI 65736; B based on Rahim Ismail

KEP 100114. All illustrations drawn by Jan van Os).

208 Gard. Bull. Singapore 61 (1) 2009

base + rounded or truncate or faintly hastate, margin (sub)entire, apex 1-1.5

cm, acuminate. Male raceme: peduncle 2-5 mm long, co-axillary with female

inflorescence; raceme spike-like, 10(-15) mm long, 10-20-flowered, flowers +

densely set. Pedicels of male flowers 1-1.5 mm long, persistent; bract lacking;

mature flower not known. Female inflorescence: an axillary or terminal few-

branched short-shoot 10-30 mm long, the branches 5-10 mm long, each with

5(-10) flowers of different stages of development in loose clusters; bracts

lacking or minute and caducous, subulate, 0.5 mm long. Pedicels of female

flowers 1—2 mm long; ovary solid, apparently 2-loculed, each with 1 ovule,

ovoid-oblong, 5.5 mm long, ca 3 mm wide, base rounded, gradually narrowed

to the apex in upper half, sparsely hairy, hairs 0.2 mm long of many serial cells;

receptacle-tube 2(-2.5) mm across, inside glabrous; sepals (long) triangular,

0.5-1 mm long; petals imbricate, pale yellow, obovate, ca 3 by (1.5-)2 mm,

apex broadly rounded, both surfaces papillose hairy, hairs 0.1 mm long; style

ca 3 mm long, stigmas 2, together 2-2.5 mm wide, each + lacerate, papillose;

staminodes minute, fleshy, 0.2-0.3 mm long, inserted near receptacle-throat,

two paired and one solitary. Fruits ripening glossy orange-red, drying glossy,

ovate-oblong, ca 15 mm long, 10-12 mm wide, glabrous; fruiting pedicel ca

2 mm long. Seed | per fruit, globose, (pale) brown, ca 7 mm across, faintly

low-margined, smooth.

Other specimens examined: Malaysia, Pahang, Ulu Krau (Rezab Hidupan

Liar Krau), N 3° 51' 33.4"; E 102° 12' 22.6", 27 Nov 2008, Siti Munirah et al.

FRI 65736 (KEP, L).

Distribution: Only known from Peninsular Malaysia, Pahang, Ulu Krau.

Habitat and Ecology: Dense primary lowland forest on hillsides on rich soil,

to 214 m altitude; flowering and fruiting in April and November.

Notes. Kedrostis monosperma is similar to K. bennettii, the latter with a

wider distribution in western Malesia, but not known from Peninsular

Malaysia. In K. bennettii the disposition of the fruit is different, single or few

subaxillary to leaves, the fruits are dull on drying, globose or transversely

ellipsoid, 1- or 2-seeded, with a longer fruiting pedicel, 5-10 mm long; the

male racemes are much longer, to 15 cm long with longer peduncle, 3-7 cm

long; the petioles are generally longer, 2-6 cm long. As far as can be judged

from the few female flowering collections of K. monosperma, the ovary is

hairy, the perianth glabrous inside, in K. bennettii the ovary is subglabrous

and the perianth hairy in the throat.

In K. monosperma the older main stem of the liana is ca 0.5 cm thick,

the bark has whitish corky patches (see Fig. 1).

Miscellaneous Cucurbit News III 209

The collection, van Balgooy 7551 of K. bennettii from Bali (Lesser

Sunda Islands), has in the male raceme caducous bracts, 1-3 mm long, in

other specimens bracts are absent.

(2) Three new combinations in African Neoachmandra and five new

combinations in Pilogyne from Malesia and the Pacific.

The genus Neoachmandra was one of several new genera proposed by De

Wilde and Duyfjes (2006), as an outcome of morphological studies in the

flowers of a large group of SE Asian species till then generally referred to

a large variable genus Zehneria. The validity of these genera on molecular

grounds is currently tested (Cross ef al., in preparation) with a broad

sampling of species, including three Neoachmandra species from Africa,

newly combined below:

Neoachmandra capillacea (Schumach.) W.J.de Wilde & Duyfjes, comb.

nov.

-Basionym: Bryonia capillacea Schumach., Beskr. Guin. Pl. (1827) 430.

Melothria capillacea (Schumach.) Cogn. in A. & C. DC., Monogr. Phan.

(1881) 600; - Zehneria capillacea (Schumach. ) C. Jeffrey, Kew Bull. 15 (19

366. -Typus: Ghana, Thonning (holo, LE, not seen).

Os 7

62)

Neoachmandra cordifolia (Hook.f.) W.J.de Wilde & Duyfjes, comb. nov.

- Basionym: Melothria cordifolia Hook.f., Fl. Trop. Afr. [Oliver et al. 2 (1871)

563. —Typus: Gabon, Mann (holo, K, not seen).

- Synonym: Zehneria gillettii (De Wild.) C. Jeffrey, Kew Bull. 15 (1962) 366.

Neoachmandra deltoidea (Schumach.) W.J. de Wilde & Duyfjes, comb.

nov.

- Basionym: Bryonia deltoidea Schumach. Beskr. Guin. Pl. (1827) 449. -

Melothria deltoidea (Schumach.) Benth., Niger Flora [W.J. Hooker] (1849)

368; - Zehneria hallii C. Jeffrey, J.W. African Sci. Assoc. 9 (1965) 93, nom. nov.

[non Zehneria deltoidea Migq., Fl. Ned. Ind. 1, 1 (1856) 655 = Neoachmandra

leucocarpa (Blume) W.J.de Wilde & Duyfjes, Blumea 51 (2006) 23, based on

Bryonia leucocarpa Blume, Bijdr. Fl. Ned. Ind. (1826) 924]. -Typus: Ghana,

Thonning (holo, C, not seen).

The uncertain taxonomic status of the genus Zehneria Endl., based on DNA

sequencing of Z. baueriana, made us to accept Zehneria as a monotypic

genus restricted to its type, from Norfolk Isl. and New Caledonia in the

western Pacific. There are but a few collections of true Zehneria from

Norfolk Isl. and New Caledonia (see next item in the present publication).

210 Gard. Bull. Singapore 61 (1) 2009

Five deviating species in Zehneria were still kept under Zehneria. But since

the re-instatement of the genus Pilogyne (de Wilde and Duyfjes, 2009),

the five taxa now appear better included in Pilogyne as well, and the new

combinations are made below.

Pilogyne erythrobacca (W.J.de Wilde & Duyfjes) W.J.de Wilde & Duyfijes,

comb. nov.

- Basionym: Zehneria erythrobacca W.J.de Wilde & Duyfjes, Blumea 51

(2006) 55. — Typus: Papua New Guinea, Milne Bay, Menapi, Cape Vogel

Peninsula, Brass 21720 (holo, L;1so, A).

Pilogyne neocaledonica (W.J. de Wilde & Duyfjes) W.J. de Wilde & Duyfjes,

comb. nov. - Basionym: Zehneria neocaledonica W.J. de Wilde & Duyfjes,

Blumea 51 (2006) 67. — Typus: New Caledonia, Ddniker 3139 (holo, L; iso,

LZ, NOUSEEM):

Pilogyne samoensis (A. Gray) W.J. de Wilde & Duyfjes, comb. nov.

— Basionym: Karivia samoensis A. Gray, U.S. Expl. Exped. Phan. (1854) 643.

— Lectotypus: Samoa, without further locality, US exploring expedition s.n.

(US, barcode US00147706 see Fosberg & Sachet, 1981).

- Homotypic synonyms: Zehneria grayana (Cogn.) Fosberg & Sachet,

Smithsonian Contr. Bot. 47 (1981) 12; W.J. de Wilde & Duyfjes, Blumea 51

(2006) 59. - Melothria grayana Cogn. in A. & C. DC., Monogr. Phan. 3 (1881)

591, p.p., excluding specimens from Tahiti.

Notes: The epithet samoensis could not be used at the time by Cogniaux

(1881) in Melothria because the combination was already preoccupied

by Melothria samoensis A. Gray, a different species, now Neoachmandra

samoensis (A. Gray) W.J. de Wilde & Duyfjes.

Pilogyne tahitensis (W.J. de Wilde & Duyfjes) W.J. de Wilde & Duyfjes,

comb. nov.

— Basionym: Zehneria tahitensis W.J. de Wilde & Duyfjes, Blumea 51 (2006)

76. — Typus: Tahiti, Vesco s.n., anno 1847, male (holo, P, barcode P00218593;

iso, P,5 duplicates).

Pilogyne viridifolia (W.J.de Wilde & Duyfjes) W.J.de Wilde & Duyfjes,

comb. nov.

— Basionym: Ze/ineria viridifolia W.J.de Wilde & Duytjes, Blumea 51 (2006)

77. — Typus: Papua New Guinea, Milne Bay, Biniguni Camp, Gwariu River,

Brass 23914 (holo, A; iso, CANB, L).

Miscellaneous Cucurbit News III 211

(3) Assessment on the morphology of the male flower of Zehneria baueriana

Endl., the type species of the genus Zehneria Endl.

The genus Zehneria was described by Endlicher (1833), with a single species,

Z. baueriana from Norfolk Island, but the generic name became commonly

used for a large number of small-flowered species of the Old World (Jeffrey,

1962). De Wilde and Duyfjes (2006) recognized within the large genus

Zehneria some smaller genera for SE Asia, i.e., Neoachmandra, Scopellaria,

Urceodiscus, and Zehneria. Currently, the validity of these genera has been

tested molecularly (Cross et al., in preparation), and preliminary results

indicate that Neoachmandra and Zehneria (in the restricted sense, as defined

by De Wilde and Duyfjes, 2006) could only be partly held up, because DNA

sequencing of Z. baueriana (as well as of some related species in the Pacific)

revealed two different clones, one clone belonging to Zehneria as newly

defined, the other to Neoachmandra, rendering Zehneria paraphyletic, and

indicating that Z. baueriana could be of hybrid origin.

Zehneria baueriana is a local endemic of Norfolk Island and New

Caledonia, represented in herbaria by only few collections. Its original

description (Endlicher, 1833, 1838 with good figures) is only in Latin. The

only other description of the species, also in Latin, is by Cogniaux (1881),

under the name Melothria baueriana, but this latter was mixed up with

material from two different species: Zehneria grayana (Cogn.) Fosberg &

Sachet, and Z. guamensis (Merr.) Fosberg. Therefore, a more recent good

collection of Z. baueriana made in 1967 (Hoogland 11220) from the type

locality, with ample male fiowering and fruiting elements, offered the

opportunity to present an accurate description, at the same time, assessing

some ambiguities in Endlicher’s original description. Also, it appeared that

the species could not be readily identified with our key (De Wilde and

Duyfjes, 2006).

At present the true state of certain characters of Z. baueriana, now

needed for an understanding of the genus Zehneria in the restricted sense,

notably characters of the male inflorescence, and the place of insertion of

the stamens in the receptacle tube in male flowers, is assessed in order to

contrast these to those resembling species from the Malesian area (De

Wilde and Duyfijes, 2009).

Zehneria baueriana Endl., Prodr. Fl. Norfolk. (1833) 69; W.J.de Wilde &

Duyfjes, Blumea 51 (2006) 51.- Melothria baueriana (Endl.) F.Muell., Fragm.

(Mueller) (1868) 188; Cogn. in A. & C. DC., Monogr. Phan. 3 (1881) 610; in

Engl., Pflanzenr. 66, 4.275.I (1916) 109. — Lectotypus: Norfolk Island, Bauer

in Herb. Endlicher s.n., male (W; iso B+, K, LE see De Wilde & Duyfijes,

2006). Figs. 3, 4.

PND, Gard. Bull. Singapore 61 (1) 2009

Branched climber 5(-10) m long; stems 2-4 mm diam; subglabrous, generally

drying brown; dioecious. Pobract oblong-linear, 4-5 mm long. Tendrils

unbranched. Leaf petioles, 1.5-3.5 cm long; blade ovate, sometimes faintly

3-lobed, 5-10 by 4.5-7.5 cm, base deeply cordate, margin remotely minutely

(sometimes irregularly) denticulate, apex acute or broadly acuminate, 2-3

mm mucronate; cystoliths minute. Inflorescences in male and female a

subsessile many-flowered fascicle, in male 10-20, in female 5-12-flowered,

outer flowers oldest, in male sometimes divided into several sub-fascicles,

suggesting a reduced leafless short-shoot; common peduncle lacking; bracts

lacking; flowers outside glabrous, pedicel with few sparse, short hairs. Pedicels

of male flowers 2.5-3 mm long; receptacle tube ca 3 mm long, 3.5 mm wide,

densely long-haired in the throat and upper third inside; sepals ca 1 by 0.6

mm, acute; petals ovate, 2(-2.5) by 1.8(-2) mm, subacute, minutely papillose-

hairy adaxially; stamens inserted halfway in the receptacle tube, filaments

ca 1 mm long, glabrous; anthers circular in outline, ca 1-1.2 mm across,

thecae curved, at apex nearly touching, forming nearly a ring, connective

not produced, broad in the middle, thickened, hairy along the thecae; disc

depressed, bluntly 3-lobed, ca 1.5 mm wide. Pedicels of female flowers (after

Endlicher) 1-2 mm long; ovary narrowly ovoid(-oblong), much narrowed

into a neck to the perianth, 15 mm long, 4 mm wide, neck to 5 mm long, all

glabrous; perianth as in male flowers; style long-cylindrical, ca 3 mm long,

style-arms 3, each ca 0.5 mm long, stigmas reniform, 2-lobed, papillose; disc

consisting of 3 separate lobes at base of style, ca 1 mm long; staminodes

inserted ca. halfway the receptacle tube, subulate, slender, ca. 1 mm long

with some coarse hairs at base. Fruits 5-10 in a cluster, at first greenish with

5 longitudinal lines and densely scale-like paler flecked (netted), ripening

(orange-) red, drying with irregular surface, collapsing over the seeds, oblong,

12-20 mm long, 8-10 mm wide, glabrous; fruiting pedicel 5-7 mm long. Seeds

15-30, compressed, elliptic, 4 mm long, (2.5-) 3 mm wide, pale, rather broadly

but indistinctly margined, edge rounded, faces not ornamented.

Fieldnotes: Petals pale yellow; fruit red (Endlicher); fruit pale green

(Hoogland).

Other specimens examined: Norfolk Isl., Bauer s.n., fruit (W); Norfolk Isl.,28-

10-1967, Hoogland 11220,2 sheets, flowers & fruit (CANB). New Caledonia,

27-11-1970, Guillaumin & Baumann-Bodenheim 8604 (L); ibid., 7-02-1951,

Guillaumin & Baumann-Bodenheim 10315 (L).

Distribution: Norfolk Island, New Caledonia.

—

oS)

Miscellaneous Cucurbit News IIT

Figure 3. Details of two herbarium sheets of Zehneria baueriana: upper details of fruit showing

one seed: lower details of female inflorescences (based on Hoogland 11220, CANB).

214 Gard. Bull. Singapore 61 (1) 2009

Figure 4. Male flower of Zehneria baueriana (based on Hoogland 11220; drawn by Jan van Os).

Habitat and Ecology: Forest edges and roadsides, 200-300 m; flowering and

fruiting in October.

Notes: The stamens are inserted halfway in the receptacle tube, not towards

the apex as written by Endlicher (1833). The filaments are about as long

as the anthers, and the stamens are largely included. The anthers are

comparatively much larger as depicted by Endlicher (1838). The disc of the

male flowers is distinctly 3-lobed, but not consisting of 3 separate globose

parts as stated by Endlicher. The disc in the female flower is 3-parted. The

corolla was described by Endlicher as fused, but the petals are free.

As described above, the male flowers link up easily with flowers of

species of Zehneria recognized for SE Asia (De Wilde and Duyfjes, 2006),

or later, as Pilogyne Schrad. (De Wilde and Duyfjes, 2009). However, for

Miscellaneous Cucurbit News III ZS

molecular reasons, the type species of the genus Zehneria, i.e., Z. baueriana,

can better be regarded as separate from Pilogyne, and the minor differences

between these two genera are primarily in the male inflorescence,

pedunculate in Pilogyne, fasciculate and not pedunculate in Zehneria. This

latter condition is also found in Pilogyne neocaledonica and P. erythrobacca

p.p. (De Wilde and Duyfjes, 2006, fig. 15), but both these species differ

considerably in various other characters from Ze/ineria baueriana.

For Z. baueriana the stout habit, the 3-parted disc in the female

flowers, the lengthwise 5-lined fruits, the yellow or pale yellow petals are

unique. Unfortunately, female flowers of Pilogyne neocaledonica are

unknown as yet. As remarked by De Wilde and Duyfjes (2006, 2009), most

of the then under Ze/neria recognised species in eastern New Guinea and

southern Pacific possess characters blurring the clear distinction between

Neoachmandra and Zehneria in this area, presumably because these species,

like Z. baueriana, are (partly) of hybrid origin as well.

Acknowledgements

This research was carried out as part of the Flora of Peninsular Malaysia

Project (Project no. 01-04-01-0000 Khas) at the Forest Research Institute

Malaysia funded by the Ministry of Science, Technology and Innovation

(MOSTI). Material from BM, CANB, K, KEP, L, SING, and W was studied.

The permission from the curator of CANB to use a herbarium leaf fragment

of Hoogland 11220 for DNA sequencing is gratefully acknowledged. We

thank Ruth Kiew (KEP) for facilitating fieldwork, and also our enthusiastic

colleagues in the field, Siti Munirah and Hok Lim Kueh (both KEP),

without their help, the second collection of Kedrostis monosperma would

not have been found. As usual, J.F. Veldkamp (L) translated the description

of the new species into Latin, Jan van Os (L) prepared the drawings and

Ben Kieft (L) scanned the drawings and prepared the photographic plates.

This research also received support from the SYNTHESYS Project which

is financed by European Community Research Infrastructure Action under

the FP6 “Structuring the European Research Area” Programme.

References

Cogniaux, C.A. 1881. Cucurbitaceae. In: A. & C. de Candolle, Monographiae

Phanerogamarum Prodromi 3: 325-951. Masson, Paris.

De Wilde, W.J.J.O. and B.E.E. Duyfjes. 2006. Redefinition of Zehneria and

four new related genera (Cucurbitaceae), with an enumeration of the

216 Gard. Bull. Singapore 61 (1) 2009

Australasian and Pacific species. Blumea 51: 1-81.

De Wilde, W.J.J.O. and B.E.E. Duyfjes. 2007. Miscellaneous South East Asian

Cucurbit news. Reinwardtia 12, 4: 267-274.

De Wilde, W.J.J.O. & B.E.E. Duyfjes. 2009. Miscellaneous South East Asian

Cucurbit news II. Reinwardtia 12: 405-414.

Endlicher, S.L. 1833. Prodromus Florae Norfolkicae. Vienna.

Endlicher, S.L. (1837-)1838(-1841). Iconographia Generum Plantarum.

Vienna.

Jeffrey, C. 1962. Notes on Cucurbitaceae, including a proposed new

classification of the family. Kew Bulletin 15: 337-371.

Gardens’ Bulletin Singapore 61 (1): 217-233. 2009 Dia.

Studies on Schismatoglottideae (Araceae) of Borneo VIII:

A Review of Piptospatha elongata in Sarawak

WONG SIN YENG |, PC. BOYCE’ AND J. BOGNER *

‘Faculty of Resource Science and Technology, Universiti Malaysia Sarawak,

. 94300 Samarahan, Sarawak, Malaysia

‘Forest Herbarium (BKF), The office of Forest and Plant Conservation Research,

National Park, Wildlife and Plant Conservation Development, Chatuchak,

‘ Bangkok 10900, Thailand

“Augsburgerstrasse 43a, D-86368 Gersthofen, Germany

Abstract

A review of Piptospatha elongata (Engl.) N.E. Br.in Sarawak is presented. The

species is shown to comprise three morphologically and ecologically distinct

taxa, two of which are new to science and herewith described as Piptospatha

impolita S.Y.Wong, P.C. Boyce & Bogner and P. viridistigma S.Y.Wong, P.C.

Boyce & Bogner. A new key to Piptospatha is presented and all species of

the Elongata group are illustrated. Additionally, new observations on the

morphologies separating the Piptospatha elongata Group and P. grabowskii

Group sensu Bogner and Hay (2000) are presented.

Introduction

Bogner and Hay (2000) provides a comprehensive and excellent revision of

the Schismatoglottideae, including Piptospatha N.E.Br., for which six species

are recognized in Sarawak. Since the publication, the first two authors have

had the opportunity to undertake comprehensive field studies, and the first

author in detailed molecular analyses of most of the presently recognized

Sarawak species and this has produced a number of new observations

resulting in the removal of Piptosptha lucens (Bogner) Bogner & Hay into

a new genus (Boyce and Wong, in press) and compelling evidence that P

elongata (Engl.) N.E. Br. sensu Bogner and Hay (2000) is heterogeneous.

Piptospatha elongata Group

The distribution and habitat data and a considerable portion of the

description given in Bogner and Hay (2000) for P. elongata incorporates five

or possibly more taxonomic elements. One of these elements is restricted

to very hard sandstones in the coastal areas of Lundu (NW Sarawak) and

218

Gard. Bull. Singapore 61 (1) 2009

is here described as Piptospatha impolita S.Y.Wong, P.C. Boyce & Bogner.

Another element is endemic to the Serian & Padawan areas as far NW as

the Sungai Tegora (SE flanks of the Bungo range) and the southernmost Bau

limestones and is usually (but not exclusively) associated with limestone.

This element is also novel and is described as Piptospatha viridistigma

S.Y.Wong, P.C. Boyce & Bogner.

Key to Piptospatha species-

ile

Spathe limb caducous;spadix above female flower zone shed directly after

male anthesis; peduncle becoming erect and infructescence erect at fruit

dispersal; fruits dispersing via ‘splash-cup’ MechaNlCs ..............ecccsseeeseereeeees

waa aia aduilop its wag get Sanne pears Sat cane ae Re ene (P. elongata group) 2

. Spathelimbalmostfullypersistentuntilfruit dispersal;spadixaxispersistent

until well after fruit dispersal; peduncle declinate and infructescence

pendentatfruitdispersal;fruitdispersalvarious(mostlyunstudiedexceptfor

Pgrabowski) butnotvia splash-cup mechanics <2... -s-ee nese ee

Peduncle shorter than spathe; W Kalimantan .......... P. brevipedunculata

Peduncle muchiexceeding the Spathe 2... ses. carer se eee 3)

Sterile interstice well-defined; NW Borneo ...............:cceeeeee P. burbidgei

Sterile mterstice absent or not well-defined x ce ee 4

Connective extended into a pronounced elongate beak; Sabah...................

sebacaseauute sass evewoaaavey es xccaetentedde st cite ae evan ceauc ste we teete eater aes nee ee eee P. insignis

Connective not raised above the thecae or shortly elevated and obtuse ...

velous guabassebecdabeed rita eryordsseedtccassegacecacetn@e techs sey eceaate ras ee center 5

Anthers pubescent to papillOse 22c. reve. acces seemetnncesseeecces ne eer eee eeeeee eee 6

Anthers Gla brOus accecsca. -cczecessessa tease sucece coos tae cesbee Soceee erace Ne tenes 8

Connective swollensPenmsulan Malay Siayeye ss scetcs oye sere P. ridleyi

Connective MOt SWONEM ..na...cccociec.cecccne een crctceeene soutex- cesses cettesconseceaee ee eee 7

Anthers in closely appressed regularly arranged pairs; Malay Peninsula

and southern peninsular Uhatland sence ee P. perakensis

Anthers (seen from above) irregularly arranged; E Kalimantan ...............

Erne ce ee ree Poon 8 dy fia orom acad oade node Saacnoshccuasacuness P. manduensis

Spadix usually with slightly narrowed sterile appendix (male zone rarely

Studies on Schismatoglottideae (Araceae) of Borneo VIII 219

fertile to apex); petiole with narrow crispate wings distally; Sarawak ........

ees os chad a dlaed hee caacovendoncedsedencceeonseseeee P. truncata

8. Male zone fertile to apex; petiole without crispate wings; Sarawak,

Pa A BERNARD ee one cosas vn na Wen couecd ene decacenonsehewasecanivabeceese 9

9. Spadix bullet-shaped, the male portion tapering towards the apex;

lower part of male zone comprised of larger flowers, that may be sterile,

intermixed adjacent to the pistils with white staminodes; thecae broadly

excavated, the excavations of adjacent anthers forming a butterfly-shaped

depression; stigmas bright green; spathe at anthesis shading proximally to

distally from deep olive-green through very pale pink to medium pink, the

interior of the spathe tip rostrum with 5-7 conspicuous keels; persistent

fruiting spathe wide-flared; plants frequently limestone associated ...........

ea Pe A aE oy Es eadaanvesdsnddneudenaveccennesdesstee P. viridistigma

9. Spadix cylindrical; male flowers uniform throughout zone; thecae

longitudinally sulcate with the pores ventral and dorsal to the sulcae;

stigmas pink or dirty whitish. Spathe at anthesis shading proximally to

distally from deep plum purple through medium pink to deep pink, the

interior of the spathe tip rostrum with 2-3 conspicuous keels or keels

absent. Persistent fruiting spathe narrowly obconic; plants of sandstone

SCAG cet a a an 11

10. Sugmas mid-deep pink; anthers with connective flat; spathe tip rostrum

almost straight or only weakly refiexed (ca 45°) relative to spathe axis at

anthesis, inside with 2-3 conspicuous longitudinal keels; plants exclusively

SE DUSTED ese ops meena en P. elongata

10. Stigmas dirty whitish; anthers with a short acute-triangular connective on

each side, spathe tip rostrum strongly reflexed (ca 130°) relative to spathe

axis at anthesis, inside without keels or these only very vaguely defined;

Plame xClUStyCly @l-SaMdStONe «.....c..20..c.-feseesteneceteessctasncouesees P. impolita

11. Female zone free, stigmas as wide as ovary, usually on a distinct style;

basal sterile zone partly adnate to spathe; widespread in Borneo .............

nce voces leagadcuee ye oR R eee ane ae Re ae Oo ee P. grabowskii

11. Lower part of female zone adnate to the spathe; stigmas narrower than

AES SESS TSSIN IM BY 0s 2 Ce. 0 ae eon on neo oe P. kinabaluensis

Piptospatha elongata (Engl.) N.E. Br.

Curtis’s Bot. Mag. 51, in descr. ad tab. 7410 (1895); Engl., Pflanzenr. 55

(IV.23Da): 124, fig. 75 (1912); Ridl., J. Bot. 51: 202 (1913); Bogner & Hay,

Telopea 9(1): 205 (2000). Schismatoglottis elongata Engl., Bull. Soc. Tosc.

Ortic. 4: 298 (1879). Rhynchopyle elongata (Engl.) Engl., Bot. Jahrb. Syst.

220 Gard. Bull. Singapore 61 (1) 2009

1: 184 (1881) & in Becc. , Malesia 1: 289, pl. 23, figs 3-15 (1882). — Typus:

Malaysia, Sarawak, Kuching Division, Lundu, Gunung Gading, June 1867,

O. Beccari P.B. 2308 (holo, FI-B). Plates 1A & 2.

-Schismatoglottis marginata Engl., Bull. Soc. Tose. Ortic 4: 298 (1879).

Rhynchopyle marginata (Engl.) Engl., Bot. Jahrb. Syst. 1: 184 (1881) & in

Becc. , Malesia 1: 288, pl. 23, figs 1-2 (1882). Piptospatha marginata (Engl.)

N.E. Br., Curtis’s Bot. Mag. 51. in descr. ad tab. 7410 (1895); Engl., Pflanzenr.

55 (I[V.23Da) (1912) 125. - Type: Malaysia, Sarawak, no further data, O.

Beccari P.B. 3838 (holo, FI-B; iso, B).

-Gamogyne pulchra N.E. Br., Kew Bull. (1910) 197 & Curtis’s Bot. Mag. 135

t. 8330 (1910). -Type: not designated (see Bogner and Hay, 2000: 205).

-Piptospatha rigidifolia Engl., Pflanzenr. 55 (I1V.23Da): 127 (1912). - Type:

Malaysia, Sarawak, Kuching Division, Lundu, Sept. 1905, H.N. Ridley s.n.

(lecto, SING, selected by Bogner and Hay, 2000: 205).

?Piptospatha angustifolia Engl. ex Alderw., Bull. Jard. Bot. Buitenzorg

III, 4: 193 (1922); Bogner, Pl. Syst. Evol. 142: 52 (1983). - Type: Indonesia,

Kalimantan, H. Hallier 614 (holo, BO), prov. syn.

Rheophytic herb 9-20 cm high. Stem condensed, 2-6 cm long, 0.5-1.1 cm

diam., with robust pale green to more-or-less reddish-tinged roots 1.5-2

mm diam. Leaves several to 20 together; petiole 6-11 cm long, 1.5-2 mm

diam., slightly canaliculate adaxially, sheathing only at the extreme base,

the remainder of the sheath ligular, the ligules extended into a narrowly

triangular, purple to reddish portion 3-5 cm long, this drying dark brown and

not long-persisting; blade very narrowly elliptic to narrowly elliptic-oblong to

oblanceolate, coriaceous, dark green adaxially, paler abaxially, 15-24 cm long

x 1-3.5 cm wide, the base cuneate, the apex acute and tubular-apiculate for

2-3 mm; midrib robust, abaxially prominent, adaxially bluntly raised (fresh),

impressed (dry), with (6-)7-10 primary lateral veins on each side, diverging

at 35°-45° and more or less regularly alternating with lesser interprimary

veins especially in the lower half of the blade; secondary venation adaxially

more or less obscure, abaxially fine and dense; tertiary venation obscure.

Inflorescence solitary to 2-3 together and then often alternating with foliage

leaves; peduncle shorter than to equalling, rarely exceeding the length of

the whole leaf, 11-22 cm long, 1.8-2.2 mm diam., purple to reddish. Spathe

nodding at anthesis, subcylindric-obovoid, 3-4 cm long, apically rostrate for

3-4 mm, the rostrum conspicuously 2-3-keeled internally, straight at female

anthesis but recurving up to 45° at the onset of male anthesis; spathe at

anthesis shading proximally to distally from deep plum purple through

medium pink to deep pink, inflating and the distal-most part opening at

female anthesis, at the end of male anthesis the opening increasing by the

reflexing of the terminal rostrum and then at late male anthesis the upper

221

toglottideae {raceae f Borne VIII

Plate 1. Spadices of five Sarawak Piptospatha species. A. Piptospatha elongata showing the

sinuate arrangement of the stamens and diagnostic pink pistils: B. P. impolita,note the carinate

connectives and dirty white pistils; C. P. viridistigma, showing the butterfly-shaped excavated

thecae and diagnostic green pistils. Note the neuter flowers below the fertile female flowers

comprise centrally-depressed white pistillodes (positioned ventrally) and yellow staminodes

(positioned dorsally); D. P. burbidgei, note the distinctive zone of staminodes separating the

male and female flower zones; E. P. grabowskii, note the conspicuous zone of pistillodes

positioned below the female flower zone, and the pubescent stamens.

ap)

222 Gard. Bull. Singapore 61 (1) 2009

Plate 2. Piptospatha elongata (Engl.) N.E.Br. A. Flowering plant in habitat. Of note is the

nodding inflorescence with almost straight rostrum (A2) and the erect infructescence (A1);

B. Inflorescence at female anthesis. Note the almost straight rostrum; C. Spadix at male

anthesis with spathe artificially removed. Note the pollen masses; D. Plant in habitat with

inflorescences at a variety of anthetic stages: D1 = inter-anthetic period; D2 = onset of male

anthesis; D3 = post anthesis; E. inflorescence during late male anthesis. Note the spathe limb

is already shed and pollen has been released; F. Infructescence approaching maturity. Note

the narrowly obconic persistent lower spathe; G. Seedlings in habitat.

Studies on Schismatoglottideae (Araceae) of Borneo VIII 223

third opening further before becoming caducous. Spadix cylindric, more-

or-less sessile, 2-2.5 cm long, 0.4-0.5 cm diam.; female zone cylindric, 5-7

mm long, 5-7 mm diam.; ovary subcylindric to subprismatic, ca 0.9 mm

diam., mid-deep pink; stigma sessile, as broad as the ovary, thinly discoid,

mid- to deep pink; staminodes confined to 1-3 irregular and somewhat

oblique rows at the base of the female zone, truncate, more-or-less sessile,

0.7-0.9 mm diam., about as high as the pistils, ivory; male zone cylindric,

approximately equalling diam. of female zone, 1.3-1.7 cm long, apically

obtuse; stamens crowded, more or less rectangular from above, glabrous, at

anthesis longitudinally sulcate with the thecae pores dorsal and ventral to

the sulcae, 1.2-1.4 mm across, connective flat, pale cream; pollen extruded

in masses. Fruiting spathe narrowly funnel-shaped, erect, 1.2-1.5 cm diam.;

berry obovoid,ca 3.5 mm long x 1.2-2 mm diam.:; seed cylindric, very slightly

ribbed, 1.4-1.6 mm long, brown but outer integument translucent, with a

long curved micropylar appendage rotting away in dispersed seeds.

Other specimens seen: BORNEO. Sarawak, Kuching Division, Lundu, Sungai

Sebako, 14 Apr 1984, Dayang Awa & Paie 8.46973 (K, KEP, L, SAN, SAR):

Lundu, Kampung Sebako, Sungai Sebako, 01° 43' 18.9"; 109° 42' 53.8", 3 Feb

2005, P.-C. Boyce & Jeland ak Kisai AR-989 (SAR + spirit); Lundu, Gunung

Gading, trail to waterfalls, 01° 41' 28.3"; 109° 50' 43.6", 14 Nov. 2006, P.-C.

Boyce & Wong Sin Yeng AR-2052 (SAR + spirit); Lundu, 2 May 1954,

Brooke 8410 (L, SAR); Lundu, Gunung Gading, 19 Jul 1963, Chai S.18484

(GH, K, L, SING); Lundu, Gunung Gading, Oct 1929, Clemens & Clemens

21924 (GH, K, SAR); Lundu, Gunung Gading, Foxworthy 326 (SAR,

SING); Lundu, Sebuluh, 21 Jan 1989, Othman Ismawi et al. S.62244 (K, L,

KEP, MO, SAN, SAR); ibid, 15 Aug 1990, (K, KEO, L, SAN, SAR); Lundu,

Gunung Pueh, Sungai Batu, 2 Mar 1989, Othman Ismawi et al. S.56652 (K,

KEP, L, SAN, SAR, US); Lundu, Gunung Pueh, Sungai Batu, 2 Oct 1974,

Mamit S.35218 (K, L, SAN, SAR, US); Lundu, Gunung Gading, Micholitz

s.n. (SING); Lundu, Gunung Gading, 19 Sep 1955, Purseglove & Shah P.4534

(K, L, SING); Lundu, Gunung Gading, 16 Aug 1960, Sinclair & bin Tassin

10365 (E,K,L,SAR, SING); Lundu, Sematan, Pueh, close by Pueh mulberry

plantation (Silkworm Farm), 7 Feb 1996, S.Teo & Awg Enjah, S.68066 (K,

KEP, SAR): Lundu, Gunung Gading, Sungai Sebuluh, 14 Jun 1991, Yahud

et al. S.61955 (K, KEP, SAR, US); Lundu, Kampung Pasir Ulu, Sungai Pasir

Ulu, Yahud et al. 8.61925 (K, SAR); Lundu, Gunung Gading, 26 Mar 1980,

Yui Puan Ching S.42018 (L, SAR, US).

Habitat: Rheophytic on granite rocks in lowland to lower hill forest in light

to medium shade. 10-400 m asl.

224 Gard. Bull. Singapore 61 (1) 2009

Distribution: Borneo, Sarawak, Kuching Division, endemic to the Lundu

area, centred on Gunung Gading. It is quite likely also present in adjacent

Kalimantan but we have not been able to re-examine the Kalimantan Barat

specimens cited by Bogner and Hay [Church et al. 2787 (K); Niewwenhuis 432

(B) and Winkler 798 (HBG); by the vague locality data that accompanies

these specimens we are unable to place them adjacent to the Sarawak

distribution of P. elongata or P. viridistigma.

Notes: Piptospatha elongata as here defined is endemic to the Lundu area

centred on Gunung Gading where it is restricted to granite substrates. It

is readily separated in flower from the other elongata Group Piptospatha

in west Sarawak (P. viridistigma and P. impolita) by the combination of a

cylindrical spadix, unexcavated thecae, mid- to bright pink stigmas, a spathe

tip rostrum conspicuously 2-3-keeled internally, and remaining straight or

reflexing by only ca 45° during anthesis, flat anther connectives and the

spathe at anthesis shading deep plum purple proximally through medium

pink to deep pink distally and, in fruit, by the narrowly obconic persistent

lower spathe.

From P. impolita, P. elongata 1s readily distinguished by the mid- to

dark pink stigmas, the spathe tip rostrum conspicuously 2-3-keeled internally

and remaining straight or reflexing by only ca 45° during anthesis, and the

flat anther connectives.

Piptospatha elongata is immediately separated from P. viridistigma

by acylindrical (vs. bullet-shaped) spadix, longitudinally sulcate unexcavated

thecae, 2-3 (vs. 5-7 keels) on the interior of the spathe tip rostrum and a

narrowly obconic (vs. wide flared) persistent lower spathe. The pollen of P.

elongata (and P. impolita) is released en masses whereas that of P. viridistigma

is extruded in strings.

Piptospatha elongata is restricted to granite, whereas P. viridistigma

is mainly, although not exclusively, found on limestones.

Piptospatha impolita S.Y.Wong, P.C. Boyce & Bogner, sp. nov.

A Piptospatha elongata affinis, sed carinae intus spathorum nullis (vs. 2-3),

rostrum spathorum valde reflexis (dum 130°) at staminis connectivo acute

producto differt. — Holotypus: Malaysia, Kuching, Lundu, Sempadi, Sg.

Limau, Bukit Kankar, 25 Aug 2007, PC. Boyce, Wong Sin Yeng & Jipom

Tisai AR-2141 (SAR, + spirit). Plates 1B & 3.

Rheophytic herb 9-11 cm high. Stem condensed, 1-4 cm long, 0.5-0.9 cm

diam., with slender pale green 0.5-1.5 mm diam. Leaves several to 10

together; petiole 2-6 cm long, 1-1.5 mm diam., very slightly canaliculate

adaxially, sheathing only at the extreme base, the remainder of the sheath

plottideae (Araceae) of Borneo VIII “4.

Plate 3. Piptospatha impolita S.Y.Wong, P.C. Boyce & Bogner. A. Flowering plant in habitat:

B. Inflorescence at late female anthesis. Note that the orifice of the spathe lacks keels; C.

Inflorescence at late female anthesis; D. Inflorescence at onset of male anthesis with spathe

limb beginning to shed; E. Infructescence at mid-maturity. Note that the persistent lower

spathe is broadly narrowly obconic; F. Detail of male flower zone just prior to male anthesis

showing the sulcate stamens and lateral, beaked connective; G. Detail of leaf lamina abaxial

venation

226 Gard. Bull. Singapore 61 (1) 2009

ligular, the ligules extended into a narrowly triangular, purple to reddish

portion 2-3 cm long, drying dark brown and moderately long-persisting;

blade very narrowly elliptic to narrowly elliptic-oblong to oblanceolate,

thinly coriaceous, matte medium-green adaxially, paler to slightly dull-pink

flushed abaxially with the mid-rib and occasionally main venation markedly

reddish, 6-13 cm long x 0.5-1.5 cm wide, base cuneate, apex acute and

tubular-apiculate for 2-3 mm; midrib robust, abaxially prominent, adaxially

bluntly raised (fresh), impressed (dry), with (6-)7-10 primary lateral veins

on each side, diverging at ca 35° and more or less regularly alternating

with lesser interprimary veins; secondary venation adaxially more or less

obscure, abaxially fine and dense; tertiary venation obscure. Inflorescence

solitary to 2-3 together and then alternating with foliage leaves; peduncle

shorter than to equalling, the whole leaf, 9-14 cm long, 1-2 mm diam., purple

to reddish. Spathe nodding at anthesis, subcylindric-obovoid, 2-3 cm long,

apically rostrate for 3-4 mm, the rostrum internally smooth or with ca 2 very

ill-defined keels, straight at female anthesis but recurving strongly by up to

130° at the onset of male anthesis; spathe at anthesis shading proximally to

distally from deep plum purple through medium pink to deep pink, inflating

- and the distal-most part opening at female anthesis, at the end of male

anthesis the opening increasing by the reflexing of the terminal rostrum and

then at late male anthesis the upper third opening further before becoming

caducous. Spadix cylindric to very weakly, 1.5-2 cm long, 0.4-0.5 cm diam.;

female zone cylindric, 5-7 mm long, 5-7 mm diam.; ovary subcylindric to

subprismatic, ca 0.9 mm diam., dirty whitish; stigma sessile, as broad as the

ovary, thinly discoid, dirty whitish; staminodes confined to 1-2 irregular

and somewhat oblique rows at the base of the female zone, truncate, more

or less sessile, 0.7-0.9 mm diam., about as high as the pistils, ivory; male

zone weakly fusiform, isodiametric to female zone, 1.3-1.7 cm long, apically

obtuse; stamens crowded, more or less rectangular from above, glabrous, at

anthesis longitudinally sulcate with the thecae pores dorsal and ventral to

the sulcae, 1.2-1.4 mm across, connective short acute-triangular, pale cream;

pollen extruded in masses. Fruiting spathe narrowly funnel-shaped, erect,

1-1.3 cm diam.; berry obovoid, ca 2.5-3 mm long x 1.5-2 mm diam.; seed

cylindric, 1.4-1.6 mm long, brown but outer integument translucent, with a

short curved micropylar appendage rotting away in dispersed seeds.

Other specimen seen: BORNEO. Sarawak, Kuching Division, Lundu,

Sempadi, Sg. Limau, Bukit Kankar, 01°39' 44.2"; 109°59'56.5", 26 Mar 2004,

PC. Boyce & Jeland ak Kisai AR-269 (SAR + spirit).

Habitat: Rheophytic on very hard sandstones in seasonally dry, but perhumid,

lowland and lower hill forest between 50-150 m asl.

im)

Studies on Schismatoglottideae (Araceae) of Borneo VIII

Distribution: Borneo, Sarawak, Kuching Division, endemic to the Lundu

area along the coast.

Notes: Piptospatha impolita is most similar to P. elongata, but readily

distinguished by the anthers with a short acute-triangular connectives (one

on each side of the stamen and held parallel to the longitudinal sulcae),

the spathe tip rostrum without internal keels, or the keels only very weakly

defined, and the rostrum becoming strongly reflexed (ca 130° vs. 45°) relative

to spathe axis at anthesis, and the dirty whitish pistils and stigmas.

Etymology: from the Latin (impolitus - unpolished) in reference to leaf

lamina markedly matte adaxially.

Piptospatha viridistigma S.Y.Wong, P.C. Boyce & Bogner, sp. nov.

Ab alli Piptospatha gregis elongatae borneensibus combinatio spadice

conoideo, antherae excavates, excavatio papilio similis, spathae intus distalis

5-7 carinae instructa et spathae fructiferorum cyathiformis differt.— Holotypus:

Kuching Division, Siburan, Kampung Giam, Air Terjun Giam, 01° 19' 11.2";

110° 16' 11.4", 7 Feb 2006, PC. Boyce, Jeland ak Kisai & Wong Sin Yeng AR-

1687 (holo, SAR + spirit). Plates 1C & 4.

Rheophytic herb 16-40 cm high. Stem condensed, 2-6 cm long, 0.8-2 cm

diam. with robust more or less reddish-tinged roots 1.5-2.5 mm diam. Leaves

several to 15 together; petiole 6-15 cm long, 1.5-2.5 mm diam., D-shaped

in cross-section, minutely aperous, sheathing only at the extreme base, the

wings extended into a narrowly triangular purple to reddish ligular portion

5-8 cm long drying dark brown; blade very narrowly elliptic to narrowly

elliptic-oblong to oblanceolate, coriaceous, dark green adaxially, paler

abaxially, 10-24 cm long x 1.5-3.5 cm wide, the base cuneate, the apex acute

and apiculate for 2-3 mm; midrib robust, abaxially prominent, adaxially

impressed, with (6-)7-10 primary lateral veins on each side diverging at 35-

45° and more or less regularly alternating with lesser interprimary veins;

secondary venation adaxially more-or-less obscure, abaxially fine and dense:

tertiary venation obscure. Inflorescence solitary; peduncle shorter than to

equalling, or rarely exceeding the length of the whole leaf, 11-24 cm long,

1.8-2 mm diam., purple to reddish, minutely asperous. Spathe nodding at

anthesis, subcylindric-obovoid, 3-4 cm long, apically beaked for 3-4 mm,

spathe at anthesis shading proximally to distally from deep olive-green

through very pale pink to medium pink, the interior of the spathe tip rostrum

with 5-7 conspicuous keels, opening in the upper third, then the upper part

caducous. Spadix the bullet-shaped, with the male portion tapering towards

the apex, anthers with the thecae broadly excavated, and the excavations of

)

228 Gard. Bull. Singapore 61 (1) 2009

Plate 4. Piptospatha viridistigma S.Y.Wong, P.C. Boyce & Bogner. A. Inflorescence at

female anthesis; B. Detail of spadix at male anthesis. Note the extruded pollen in strands;

C. Inflorescence at late female anthesis. Note the conspicuous keels at the orifice of the

spathe; D. Inflorescence at onset of male anthesis with spathe limb beginning to shed; E.

Inflorescence at late male anthesis with spathe limb shed. Note the copious pollen strands;

F. Infructescence at mid-maturity. Note that the persistent lower spathe is broadly funnel-

form.

Studies on Schismatoglottideae (Araceae) of Borneo VIII 229

adjacent anthers forming a butterfly-shaped depression, and the diagnostic

bright green stigmas and pistils, 2-2.5 cm long, 0.4-0.5 cm diam.; female zone

fusiform, 5-7 mm long, 5-7 mm diam.; ovary subcylindric to subprismatic, ca

0.9mm diam.; stigma sessile, as broad as the ovary, thinly discoid; interpistillar

staminodes absent from among the pistils; neuter organs (an admixture of

staminodes and pistillodes) confined to 1-2 irregular and somewhat oblique

rows at the base of the female zone, staminodes truncate and centrally-

depressed positioned ventrally, white, pistillodes truncate or weakly convex,

positioned dorsally, yellow, all more or less sessile, 0.7-0.9 mm diam.,

about as high as the pistils; sterile interstice comprised of on in complete

row of white staminodes and 1-2 rows of the lowermost male flowers larger

than the fertile flowers and apparently sterile; male zone stoutly fusiform-

cylindric, isodiametric with female zone, and often with a distinct ledge at

the base that truncately-overhangs the female zone, 1.3-1.7 cm long, apically

obtuse; stamens crowded, truncate, the connective not raised, more-or-less

rectangular from above, glabrous, 1.2-1.4 mm across, thecae excavated,

adjacent thecae excavations forming a butterfly-shaped deporession, deep

yellow; pollen extruded in strings. Fruiting spathe broadly funnel-shaped,

erect, 1.2-1.5 cm diam.; berry obovoid, ca 3.5 mm long x 1.2-2 mm diam.; seed

cylindric, very slightly ribbed, 1.4-1.6 mm long, brown but outer integument

translucent, with a long curved micropylar appendage which rots away in

old seeds.

Other specimens examined: BORNEO. Sarawak, Kuching Division, Padawan,

Bukit Manok, 01° 12'; 110° 18', 1 Apr 2004, PC. Boyce, Jeland ak Kisai &

A.Shafreena AR-14 (SAR, + spirit); ibid, 18 Mar 2004, PC. Boyce, Jeland

ak Kisai & A.Shafreena AR-250 (SAR, + spirit); Siburan, Kampung Giam,

Air Terjun Giam, 01° 19' 11.2"; 110° 16' 11.4", 7 Feb 2006, PC. Boyce, Jeland

ak Kisai & Wong Sin Yeng AR-1687 (SAR, + spirit); Bau, Bongo Range,

trail to Tegora Mine, 01° 19' 41.5"; 110° 09' 19.0", 8 Sept. 2007, PC. Boyce,

Wong Sin Yeng & Alexander Kocyan AR-2185 (SAR, + spirit); Bau, Gunung

Lanyang, 11 Apr 2001, Julia S., et al. SBC 2683 (SAR, SBC); Padawan,

Gunung Gayu, 1° 12' 52"; 110° 19' 59", 14 Dec 1995, Kato et al. (SAR);

Padawan, Stabut, 16 Jan 1970, Mamit S.29953 (A, BO, E, K, L, MEL, SAR):

Padawan, Jalan Kampung Annah Rais, hot springs, 12 Jan 1996, Mohizah et

al. S. 66820 (K, KEP, L, SAN, SAR, US); Padawan, Kampung Sadir (*Sadil’),

Mini Hydro Station, Yii Puan Ching S.51335 (K, SAR); Gunung Siruruh,

Sarawak/Kalimantan border, 21 Sep 1987, Yii Puan Ching S. 55240 (K, KEP.

L, SAN, SAR, US). Sarawak, Samarahan Division, Serian, Sungai Ranchan

(‘Renchang’), Feb 1963, Ashton S.21298 (K, L, SAR, SING); Tebakang,

(‘Tabakang’), Bukit Alak, Awa & Paie S.45730 (K, SAR); Serian, Pichin,

Umon Murut, Tiab Belanting, 01° 08' 03.7"; 110° 27' 00.3"S, 22 Jun 2005,

230 Gard. Bull. Singapore 61 (1) 2009

P.C. Boyce, Jeland ak Kisai & A.Shafreena AR-1256 (SAR, + spirit); Serian,

Pichin, Bung Biringan, 28 Oct 2004, PC. Boyce & Simon Kutuh ak Paru

AR-733 (SAR, + spirit); Serian, Mongkos, Kampung Batuh Mawang, Labak

Ebang, Utak Samat, 5 Jan 2006, PC. Boyce & Simon Kutuh ak Paru AR-

1658 (SAR, + spirit); Serian, Gunung Niyat, Ulu Sungai Majat, 27 Feb 2002,

Jemree & Enjah S. 85506 (K, KEP, L, SAN, SAR); Serian, Ranchan Falls,

26 Feb 1993, Lai Chak Teck, Rantai et al. S. 66046 (K, KEP, SAR); Gunung

Sirang, nr Sarawak/Kalimantan border, Mamit $.35875 (L,SAR).

Habitat: Rheophytic, usually on limestone, occasionally on sandstone (pers.

obs.) or basalt (fide Ashton S.21298) along small forest streams and waterfalls

in light to medium shade, 100 — 350 (950) m asl.

Distribution: Borneo, Sarawak, Kuching & Samarahan Divisions, endemic to

the Serian & Padawan areas as far NW as the Sungai Tegora (SE flanks of

the Bungo range) and the southernmost Bau limestones. As with P. elongata

it is very likely that P. viridistigma extends into adjacent Kalimantan but we

have not been able to re-examine the Kalimantan Barat specimens cited by

Bogner and Hay (2000).

Notes: Piptospatha viridistigma is readily differentiated from P. elongata and

P. impolita by the bullet-shaped spadix, with the male portion deep yellow

(vs. pale cream) tapering towards the apex, the anthers with the thecae

broadly excavated with the excavations of adjacent anthers together forming

a butterfly-shaped depression, and the diagnostic bright green pistils (from

whence the trivial epithet is derived) and stigmas. Other characters include

the minutely puberulent petioles and peduncle, pollen extruded in strings (vs.

masses) and the spathe at anthesis shading proximally to distally from deep

olive-green through very pale pink to medium pink. In fruit the persistent

fruiting spathe is wide-flared rather then narrowly conical.

Sterile plants of P. viridistigma are very similar to P. elongata, although

the uniformly minutely asperous petioles (D-shaped in cross-section) and

longer persistent petiolar sheath are stable morphologies to differentiate P

viridistigma from P. elongata. To date P. elongata has never been collected

away from granite.

Despite the long standing obfuscation with P. elongata we believe that

the closest morphologically similar species is probably P. burbidgei (N.E.Br.)

M.Hotta based on the presence of a sterile interstice comprised of the

lowermost male flowers being larger than the fertile flowers and apparently

sterile and intermixed with an complete row of white staminodes (and thus

morphologically similar to the staminodal interstice that is diagnostic of P.

burbidgei) and also the bullet-shaped spadix. However, it must be noted that

Studies on Schismatoglottideae (Araceae) of Borneo VIII 231

vegetatively P burbidgei is quite different in appearance to P. viridistigma

(indeed different to all other Sarawak Piptospatha) and that the apparent

similarities in the inflorescences may well be an evolutionary parallel or

convergence.

Etymology: from the Latin for green (viride) and stigma in allusion to the

strikingly green coloured stigmas.

Additional elements included in P. elongata sensu Bogner

and Hay (2000)

Other elements cited by Bogner and Hay (2000) as belonging to P. elongata,

but which need to be excluded, or for which doubt exists are:

Nicolson 1264 (K+photo, L+photo, SAR+photo, US+photo), Kuching

Division, Matang FR, 10 mi W of Kuching, is unequivocally P. grabowskii

(Engl.) Engl.

Argent et al. 692 (L) from ~Miri Division (“4th Divn’), Gunung Mulu NP.

Sungai Lansat”, is referable to P burbidgei. The first two authors have

undertaken extensive fieldwork in the lowlands of Mulu, including the

Sungai Lansat, and are confident that P. elongata is not present in the area

(nor, indeed, anywhere in NE Sarawak), whereas P. burbidgei is a common

species on the exposed shales throughout the Mulu area, including on the

Sungai Lansat.

McDonald & Ismail 3615 (GH) from “Kalimantan Timur. Bulungan

(“‘Pujungan’) Distr.. Kayan-Mentarang Reserve....” may belong to either

P. burbidgei or P. manduensis Bogner & A.Hay. or be referable to a novel

taxon but requires re-examination. We are confident that by the considerable

geographical disjunction that it is not referable to any species dealt with in

detail in this paper.

The depauperate Hullett sn. (SING) collection from Sri Aman Division,

Lingga, Batu Gajah, is interesting in that it may be referable to the very

poorly known P. remiformis Ridl., the type of which is missing but originates

from Lingga. At present Lingga is notably under-collected but, based on

the few aroids so far gathered from there, harbours an intriguing mixture

of otherwise highly localized west Sarawak endemics including Aridarum

nicolsonii Bogner & M.Hotta and A. crassum S.Y.Wong & P.C. Boyce.

However, neither of these species in their main distribution occur in the same

locality nor on the same geology as P. elongata. The Lingga area needs to be

DBP Gard. Bull. Singapore 61 (1) 2009

extensively investigated before a confident identification can be made.

Informal Groups

One of the significant contributions in the Bogner and Hay (2000)

Piptospatha account is the proposal of two groups, the P. elongata Group

and the P. grabowskii Group defined by the persistence or otherwise of the

spathe limb into fruiting. We have observed a suite of additional characters

that further reinforce the morphological and perhaps phylogenetic validity

for these two groups.

Elongata Group

The spathe limb is caducous during anthesis as reported by Bogner and Hay

(2000). Based on our observations, spathe limb senescence begins mid-way

through the period of male anthesis, when the somewhat soft-textured limb

degrades and becomes increasingly fragile, while at the same time abscising

from the leathery and persistent funnel-form lower spathe. By the end of

male anthesis the inflorescence is comprised of an erect to sub-erect peduncle

and a nodding spathe/spadix from which the spathe limb is already shed and

the spadix is still intact. With the completion of male anthesis, providing

that the female flowers have been successfully fertilized, the spadix above

the female zone is shed — if fertilization has not been successful the entire

spadix decays and is shed. The shedding of the distal portion of the spadix

is accompanied by the peduncle becoming more or less erect and the axis at

the attachment of the spathe twisting so as to bring the funnel-form lower

spathe orthotropic to the peduncle; in such a posture the lower spathe later

functions as a‘splash-cup’ dispersal unit. At or shortly after the spathe being

bought vertical, the lower spathe perceptibly thickens and turns green.

Grabowskii Group

Bogner and Hay (2000) defined the Grabowskii Group by (in successfully

pollinated inflorescences) the spathe limb persistent throughout and after

anthesis with only the very distal-most portion decaying. We are able to add

to these observations and confirm that the spathe persists, with the very

distal-most portion decaying only after anthesis, until during fruit dispersal

at which point the spathe abscises at the junction of the peduncle and is lost

along with the dispersing fruits. Other notable differences in the Grabowskii

Group are that the inflorescence post-anthesis is declinate by deflexing of

the lower part of the peduncle and also the spadix is persistent until post-

dispersal, the spent male flowers and various neuter structures are shed, but

the axis remains even after the fruits and persistent spathe have been shed.

Based on on-going research it appears that the shedding of the distal-most

o>)

Studies on Schismatoglottideae (Araceae) of Borneo VIII 23

part of the spathe serves to enlarge the opening at the end of the now much

thickened spathe and is perhaps associated with facilitating the shedding of

the spent male flowers and neuter organs. It also appears that the spadix axis

is perhaps involved with fruit dispersal mechanics but more observations

are needed to confirm this. Whatever the outcome of studies on these

morphological interactions, it is clear that fruit/seed dispersal mechanics of

the Elongata and Grabowski groups is fundamentally different.

Pistillodes & Staminodes

Bogner and Hay (2000) noted several instances in Piptospatha where it was

not possible from dried and alcohol-preserved material to unequivocally

assign neuter organs as staminodes or pistillodes, in particular with regard

to such neuter organs situated below the female flower zone (i.e., basal-

most on the spadix). We have been fortunate to observe a large number of

fresh inflorescences of five species that are indigenous to Sarawak; based

on these observations we are confident that the lowermost organs in P

grabowskii (Plate 1E), P. elongata (Plate 1A), P. impolita (Plate 1B) and P.

burbidgei (Plate 1D) are pistillodes, while those present in P. viridistigma

are comprised of pistillodes and a few staminodes (see Plate 1C).

References

Bogner, J. and A. Hay. 2000. Schismatoglottideae in Malesia II — Aridarum,

Bucephalandra, Phymatarum and Piptospatha. Telopea 9(A): 183-194.

Boyce, P.C. and S.Y. Wong. 2009. Studies on Schismatoglottideae (Araceae)

of Borneo VII: Schottarum and Bakoa, two new genera from Sarawak,

Malaysian Borneo. Botanical Studies: in press.

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Gardens’ Bulletin Singapore 61 (1): 235-236. 2009 235

Book Review: J. H. Beaman and P. J. Edwards. 2007. Ferns of Kinabalu, an

Introduction. Natural History Publications (Borneo). 198 pp. ISBN: 983-

812-122-3. Price: Sing $53.

Mt. Kinabalu in Sabah State of Malaysia is a paradise of plant diversity.

In recent years, there have been many introductory pictorial guidebooks

published for various groups of plants grown on this highest mountain in

the Asian tropic. To my knowledge, this seems to be the first one on the fern

diversity.

This beautifully executed book with vividly coloured photo plates

of the more commonly seen fern species around the Kinabalu Nature Park

Headquarter and the several popular trails from 600 to 2,800 m, is very well

written by J. H. Beaman and P. J. Edwards, two knowledgeable botanists

who have studied the flowering plant and fern floras of Mt. Kinabalu for a

number of decades.

The book starts aptly with a succinct introduction describing the life

cycle of fern, a discussion of the latest system of classification of ferns based

on the molecular phylogeny, and a brief summary of the fern distribution in

the region from the point of view of the Mt Kinabalu flora.

More than a hundred pages of coloured photos and illustrative line

drawings, with short description for each of the species chosen to represent 29

recognized families arranged in alphabetical order, follow the Introduction.

And the book ends with a useful and updated checklist of all known ferns

from Mt Kinabalu, a Glossary, the Acknowledgements and an Index to the

Scientific Names.

What is new in phylogenetic interpretation published in this

guidebook is the inclusion of Psilotum and Equisetum in the ferns, and not

the traditional fern allied plant group. For Equisetum, the authors reported

that the recent DNA evidence indicated a close relationship to the Marattia

ferns. Eight endemic Bornean ferns are shown in very nicely reproduced

photos, namely Aglaomorpha brooksii Copel., Alsophila ramispina Hook.,

Elaphoglossum heterolepium Alderw., Grammitis reinwardtioides Copel.,

Loxogramme ensifrons Alderw., Mesophlebion dulitense Holttum, Selliguea

murudensis (C. Chr.) Parris, and Sphaeropteris capitata (Copel.) R.M.

Tryon. Likewise, five Mt. Kinabalu endemic fern, Dicranopteris clemensiae

Holttum, Diplazium atrosquamosum (Copel.) C. Chr., D. poiense C. Chr.,

Odontosoria veitchii (Baker) Parris and Sphaerostephanos lithophyllus

(Copel.) Holttum, are included with well taken closed up photos showing

also the fertile fronds.

Other nomenclatural and taxonomical novelties encountered in

this book are the use of Odontosoria for the genus Sphenomeris. The split

genera in the family Cyatheaceae, Hymenophyllaceae, Thelypteridaceae

236 Gard. Bull. Singapore 61 (1) 2009

and Grammitidaceae are recognized, while the recently proposed new

genera of Vittariaceae and the Polypodiaceous genera of Phymatorus (=

Microsorum) and Crypsinus (= Selliguea) are not accepted. Diplazium and

Athyrium are place in the family Woodsiaceae.

Although the book illustrated only 100 taxa of common ferns seen

on Mt. Kinabalu out of the 590 known taxa of the mountain fern flora, the

book is a useful guide to have while visiting the mountain either on an

excursion at leisure or on a more seriously planned botanical exploration

visit. Since the majority of the species are widespread in the region, sure

enough, the book can also serve as a good reference, used with monographs

and floristic publications, in the identification of the ferns in tropical SE

Asia. The authors and the publisher are to be thanked for producing this

timely and useful pictorial guide to the Kinabalu ferns.

B. C. Tan

The Herbarium

Singapore Botanic Gardens

Gardens’ Bulletin Singapore 61 (1): 237-239. 2009 DEG

Book Review: Hughes, M. 2008. An Annotated Checklist of Southeast Asian

Begonia. Royal Botanic Garden Edinburgh. 164 pp. Price: Sing $60 (£25).

This useful and informative publication aims to stimulate, as stated by the

author, the production of much needed local floras of Begonia through the

provision of a regional checklist with extensive bibliography.

The main body of the publication consists of a short and information-

rich Introduction, followed by a lengthy section of Annotated Species

List and a Geographic List, and ended with the References and Index of

Names.

According to the author, the main annotated list accounts for 521

species of Begonias reported from 18 Botanical Countries (see Brummit,

2001) in SE Asia based on some 5,827 specimens housed in various herbaria.

The political countries included in this checklist include Myanmar, Thailand,

Vietnam, Laos, Kampuchea, Malaysia, Philippines, Indonesia, Brunei, Papua

New Guinea, and the Pacific Island groups, Vanuatu and Fiji. Surprisingly,

based on current information, Philippines has the highest number of

species recorded (104 species) and also the greatest number of endemic

species reported (103 species). This is followed by Borneo (95 species) and

New Guinea (79 species). The high number of species of Begonias in the

Philippines probably reflects more on the long historically preference and

frequency of collections of this genus in the country.

For every species listed in this publication, sufficient information is

provided for the original publication and important taxonomic bibliography,

the complete synonymy, the type information, the sectional classification,

the enumeration of specimens reported in publications and where deposited

today, and most importantly, the author’s proposed IUCN status of the

species endangerment. According to the author, readers of the publication

can also find the digital images of the plant and the protologues of each

species by consulting the website of the Royal Botanic Garden Edinburgh.

The species names in the main annotated list are arranged

alphabetically for ease of searching. Likewise, the names of the taxa placed

under a country in the Geographic List are also alphabetically listed, but

with additional listing of excluded taxa from the country/island groups based

on the author’s many years of taxonomic and nomenclatural researches.

Some nomenclatural novelties published in this checklist include

Begonia cristata Warb. ex L.B.Sm. & Wassh. (syn. nov., p. 7), Begonia

yunnanensis H. Lév. (syn. nov., p. 83), B. sootepensis var. thorelii Gagnep.

(syn. nov., p. 83), B. lushaiensis C.E.C. Fish. (syn. nov., p. 83), Casparya

robusta var. glabriuscula A. DC. (syn. nov., p. 86), and B. modestiflora var.

sootepensis (Craib) Z. Badcock ex M. Hughes, comb. nov. (p. 84).

Additionally, two species of Begonia, B. kaniensis Irmsch. and B.

238 Gard. Bull. Singapore 61 (1) 2009

demissa Craib, are beautifully illustrated.

What I find most elucidating and informative about this publication

is the many comments of the author provided for many species throughout

the publication as Notes to explain the confusing history of a name, the past

errors made in reporting a species, the author’s taxonomic interpretation, the

alleged and corrected range information, as well as the justification of revised

sectional classification. All these notes show the profound knowledge of the

author for the genus, taxonomically, nomenclaturally and bibliographically.

Having been involved in the study of the flora of the Philippine

Archipelago for many years, I did find a few small errors in the spelling

of locality and the collector names that deal with the Philippine taxa. For

examples, the collector name of “Edafo” has been written inaccurately as

“Edano”, and the town name, “Penablanca” in Cagayan Province is mis-

spelled as “Panablanca”. Moreso, the locality name of Mt. Makiling is the

same as Mt. Maquiling, and Mt. Banahao is the same as Mt. Banajao. Mt.

Susong-Dalaga (female breast) in Rizal Province (see p. 37) 1s mis-spelled

as Mt. Susong-Dalanga. “Baranguay San Jose” on p. 26, the type locality

under B. chloroneura P. Wilkie & Sands, should be written as “Barangay

San Jose”. The type locality of B. longibractea Merr. (see p. 72) is Siargao

Island, and not Siargo Island as printed. Also, the type locality of Begonia

subtruncata Merr. in Union Province (see p. 125) is today known as “La

Union Province”. Likewise, the type locality, “Banos” town, of B. aequata A.

Gray in Laguna Province (p. 4), is to be “Los Bafios” town.

I have two nomenclatural points to make. (1) The listing of the

accepted name for the homotypic synonymy between Begonia wenczelii

Merr. (1915) and B. leytensis Merr. (1914), in favour of the former (p. 138),

is not clearly stated. The latter name is actually a junior homonym of B.

leytensis Elm. (2) Naked names, like B. bulusanensis Elmer ex Merr. and B.

hemicardia Elmer ex Merr.on p. 15, B. neopurpurea L.B.Sm. & Wassh. on p.

89, and B. sorsogonensis (Merr.) Elmer on p. 121, lack a type specimen, but

have an original specimen.

The workers and students of Begonia in SE Asia should be grateful

to the author for compiling such a useful publication that builds the

bibliographical groundwork for the documentation of the richness of the

species of this genus in the region; many of them are local endemic and

have high ornamental value. Seen in this light, this publication should be

considered essential and a bible to plant growers, landscapers and gardeners

who need to check on the correct name of a species of Begonia from SE

Asia used in horticulture.

Literature Cited: Brummit, R.K. 2001. World Geographical Scheme for

Recording Plant Distributions. Hunt Institute for Botanical Documentation,

Rhododendron sojolense

Pittsburgh.

B. C. Tan

The Herbarium

Singapore Botanic Gardens

239

The Gardens’ Bulletin

Singapore

VOL. 61 (2) 2010 | ISSN 0374-7859

A Commemorative Volume of the

150th Anniversary of the Singapore Botanic Gardens

Singapore Botanic Gardens DBOQuears © oe

necting PI —<——

THE GARDENS’ BULLETIN SINGAPORE

The Gardens’ Bulletin Singapore publishes original papers on plant taxonomy

(including revisions), horticulture, phytogeography, floristics, morphology,

anatomy and related fields with emphasis on plants in the West Malesian

region.

| Dr. B. C. Tan Dr. Jana Leong-Skorni¢ékova

Singapore Botanic Gardens Singapore Botanic Gardens

(Editor) (Assistant Editor)

Dr 2 Yotan Ms. C. Soh

National Parks Board Singapore Botanic Gardens

(Assistant Editor) (Journal Business Manager)

EDITORIAL BOARD

Dr. S.C. Chin Dr. M.C. Roos

Singapore Botanic Gardens National Herbarium Netherlands

Singapore Leiden University, The Netherlands

* Dr. M.J.E. Coode Dr. E. Soepadmo

Royal Botanic Gardens Forest Research Institute Malaysia

Kew, U.K. Kepong, Malaysia

Prof. Sir P. Crane Prof. T. Stuessy

Yale University University of Vienna

New Haven, USA Austria

Dr. R.T. Corlett Dr. W.K. Tan

DBS, National University of Singapore National Parks Board

Singapore Singapore

Dr. W.J. Kress Dr. I.M. Turner

Department of Botany, NUNH Research Associate

Smithsonian Institution Singapore Botanic Gardens

Washington DC, USA Singapore

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

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

conclusions expressed by the contributing authors.

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

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

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

Singapore.

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

[Cover photo : Musa arfakaina with mature fruit (see p. 245); photo by G. Argent]

fe y) MA UT 20 REED LESORE LAG

ite 4 UBRARY

The Gardens' Bulletin

Singapore

IN)

VOL. 61 (2) 2010 ISSN 0374-7859

CONTENTS

Congratulations to the Singapore Botanic Garden Sesquicentennial 2009

By Dr. C. Samper, Prof. P.H. Raven, Prof. S. Hopper and Prof. Sir P. Crane

Argent, G.

A New Species of Wild Banana Musa arfakiana (Musaceae) from Papua (Formerly Irian Jaya) of In-

RIES I Auer ne een Pe ce see eeas cers Reece noet Vo vnause cn sbssusocnnestoeonevessenvesnvetbenssustsoocsetuentenss sostessosunvenrecetous D4

Boyce, P.C., 1.B. Ipor and W.L.A. Hetterscheid

AReview of the White-flowered Amorphophallus (Araceae: Thomsonieae) Species in Sarawak .............

_apeeceeeenstccacenacesicest Sac Sc cei Sue SoECBEE GEESE SRE SEER eo cE Sc er ne 249

Boyce, P.C.,S.Y. Wong and Fasihuddin B.A.

Studies on Homalomeneae (Araceae) of Borneo II: The Homalomena of Nanga Sumpa (Batang A1) —

Noveliand) Pre-existing: Paxa-andJNotes om Iban WSages <......c.--<-cqceeece--coneeceeenc sees cece encer dest eeneaeness=s 269

Green, P.T., J. Claussen and D. J. O’Dowd

Lost for a Century: Rediscovery of the Endemic Ridley’s Jewel Orchid, Zeuxine exilis Ridl., on

CTR TETRIS err a Via VETO LOSE TAY 2 eRe BS Se on aE enn 319

Kloppenburg, R. and S.V. Siar

New Species of Hoya (Apocynaceae) from Brunei and the Philippines ..................:ceeseesseeseeeteeeeeees S27

Kumar, P. and J.F. Veldkamp

Pecteilis rawatii (Orchidaceae), a New Species from India 335

Liang, J., H. Zhu and Y.-X. Ma

Land Use, Land Cover Change and Conservation in the Dipterocarp Rain Forest Area of Souther

BUEREATDCATVEN Gs NE TA CANM Ore See ec RN 3 esto dtc Seren nt wee gat aie Coace cnigenony Noses n Ce seuceuscaun darouecguete a8

Ly, N.S.,S. Hul and J. Leong-Skorniékova

Siliquamomum oreodoxa (Zingiberaceae): a New Species from Southern Vietnam ..................... 359

Middleton, D.J.

Three New Species of Wrightia (Apocynaceae: Apocynoideae) from Thailand .....................0000 369

Miettinen, O. and N. Hernawati

Two Basidiomycetes New to Indonesia, Prerygellus armeniacus and Rimbachia Leucobryi ........ 379

Nathi, Y., B.C. Tan and T. Seelanan

Ten New Records of Mosses from Doi Inthanon National Park in Thailand .............cccccceceeeceeeeeeee 386

Reynolds, D.R.

Epitoliar Fungi of Singapore ise) 2 metre crete eee ate need gn caine sale cor no 40)

Sipman, H.J.M.

AVConspectusottheeichens| (eichenized Fun ei) Totisin gapOle eccereesessesese ee eeeeeee sees seamen ee 43°

Staples, G.W.

A Checklist of Merremia (Convolvulaceae) in Australasia and the Pacific ...........cc00ccccceeeeeeeeeeeeeees 48:

Widodo, P., A. Hartana and T. Chikmawati

Syzygium subscandens (Myrtaceae), a New Species from Sumatra ..............::cceccesesescesssesecescessensee DD.

Wong, S.Y.

Studies on Schismatoglottideae (Araceae) of Borneo XII: Three New Species of Schismatoglottis 11

the: Multiflora Grou pinta. cies hcrtes eet esis As acces reev e DD

Wong, S.Y. and P.C. Boyce

Studies on Schismatoglottideae (Araceae) of Borneo X. Pichinia, a New Genus from Sarawak

Malaysian BOmmne) 2..vivcccsccheccvssseassscesed coacee vedvaaeoses vencnessvevenesvausoorerncsse ce oree eee eo 54

Book Reviews

A Guide to the Mosses of Singapore by B.C. Tan and B.C. Ho

Parasitic Flowering Plants by H.S. Heide-J@rgensen

RO) 0) CCR one ee ee Re eee cone EES Sooo ce sooo eee daced aod donconosoconcennenbonscoabsc000000% 55)

Forest Formations of the Philippines by E.S. Fernando, Min Hwan Suh, Jaeho Lee and Don Koo Le

TANS Bie. vccccesccccssesecssconccescunaeone acess vausedecevsceat ev een te et sedans ee tone ones ateceee dec Cae eee ee 35:

Date of publication: March 2010

Published and copyrighted by

National Parks Board

Singapore Botanic Gardens

1 Cluny Road

Singapore 259569

Printed by Oxford Graphic Printers Pte Ltd

Gardens’ Bulletin Singapore 61 (2): 241-242. 2010 241

Congratulations to the Singapore Botanic Garden

Sesquicentennial 2009

Throughout 2009, universities, museums and botanical gardens in countries

across the world, have celebrated the bicentenary of the birth of Charles

Darwin. Most recently, on November 24" 2009, the 150° anniv ersary of the

publication of Darwin’s classic book “The Origin of Species ...” was the

opportunity for further reflection on the impact of his work. Through his

ideas and experiments Darwin established the foundation for our modern

understanding of biological diversity and how it has come to be through the

processes of evolution.

But in many celebrations of Darwin’s life it is sometimes forgotten

that his insights had as much meaning for those interested in the variety

of plant life as those concerned with the origin of diversity in the animal

kingdom. Indeed, Darwin devoted several books to different aspects of

plant evolution, and the mid-nineteenth century scientific milieu in which

he worked was a golden age of botanical exploration. This was a time when

new plants were being discovered and brought into cultivation from the

most remote corners of the globe.

The Singapore Botanic Gardens, founded in 1859 by a group of

agriculturalists and horticulturalists in Tanglin, was borne in this era. Its

origin lies in an era of fascination with plants and their potential economic

importance, and at a time when the value of public parks, for leisure and

beauty, was first being realized. Ever since, the Gardens has played a major

role in the life of Singapore and the surrounding region. Its luxuriant

planted landscapes are echoed throughout the city. Over 150 years, the

Singapore Botanical Gardens has evolved into one of the great botanical

gardens of the world. Both a major tourist destination and a national icon,

it continues to enrich the lives of the people of Singapore. Open every day,

free of charge, from early in the morning until late at night, the Singapore

Botanic Garden is a well-loved national treasure.

Plants have beenimportantin the history and development of Singapore

since Sir Stamford Raffles founded the first “Botanical and Experimental

Garden” on Government Hill in 1822 almost at the birth of the colony itself.

Since then, and especially since the early scientific work of H. N. Ridley, the

Singapore Botanical Gardens has been at the center of botanical research

on the plant life of tropical South-east Asia. That leadership continues

today, with even more urgency, as the Gardens coordinates with regional and

global partners on the conservation and sustainable use of tropical plants.

In 1859, the same year that the Singapore Botanic Gardens was

242 Gard. Bull. Singapore 61 (2) 2010

founded, on the other side of the world, Henry Shaw established the Mis-

souri Botanic Garden in St. Louis. A hundred years earlier, in 1759, the

Royal Botanic Gardens, Kew began its transition from a Royal estate to

a scientific collection of plants. The past year, 2009, has therefore been

a time for celebration at all three of these great gardens. A celebration

given greater significance by the recognition that all three now work ever

more effectively as part of a global network of more than 2,000 botanical

gardens - all dedicated to the cultivation and conservation of plant spe-

cies, and increasing knowledge and public enjoyment of botanical diversity.

We send our congratulations to the Singapore Botanic Gardens

on 150 years of remarkable achievements and outstanding accomplish-

ments. We wish it well at its sesquicentennial, and in the coming decades,

as it continues its important work of “Connecting People and Plants”.

Dr. Cristian Samper

Director, National Museum of Natural History, The Smithsonian

Institution

Professor Peter H. Raven

President, Missouri Botanical Garden

Professor Steven Hopper

Director, The Royal Botanic Gardens, Kew

Professor Sir Peter Crane

Carl W. Knobloch, Jr., Dean, School of Forestry and Environmental Sciences

Yale University

Gardens’ Bulletin Singapore 61 (2): 243-248. 2010 243

A New Species of Wild Banana Musa arfakiana (Musaceae)

from Papua (Formerly Irian Jaya) of Indonesia

G. ARGENT

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

Abstract

Musa arfakiana is described and illustrated as a new species from Papua,

Indonesia.

Introduction

New Guinea is an important centre of diversity for the genus Musa (Argent,

1976, 2001). With the addition of this new record, there are 11 wild species

recorded, most of which are endemic. The area is also arguably the richest

place in the world for indigenous cultivars (Simmonds, 1966), many of which

are diploides and potentially important as a gene reservoir.

Terminology and description below follows Simmonds (1962, 1966) and

Argent (1976).

Musa arfakiana Argent, sp.nov.

Musae johnsii Argent similis sed fructu minore distaliter acuto haud truncato

non schizocarpo, fructuum fasciculo erecto, tepalis compositis cum apicibus

viridibus non cremeis et canale petiolari paene clauso haud late aperto differt.

— Typus: INDONESIA. Papua, West Papua, Kampung Siobri, Mokwam

District, Arfak Mts. Mt. Sembiedip, 01° 07° S. 133° 54° E. 2 Feb 2009, Argent

et al ABEG 106 (holo, BO; iso, E & Manokwari Forestry Herbarium). Figs.

ie 3.

Clump forming herbaceous plant, suckers erect and close to the base

of the parent plant but mostly only one or two, the clumps small. Sucker

leaves mostly auriculate at the base. Pseudostem to 1.5-2 m,9-12 cm diameter

near the base, predominantly black, otherwise green with some dark brown

coloration in the upper parts, no wax; undersheath green, inner sheath white

with purple streaks, sap clear. Shoulder brown or green, entire, smooth and

appressed, the margin black but not scarious. Fourth last leaf ca 160 x 45

cm, right handed to ca 10 mm, the base auriculate, slightly broader in the

lower half. Other leaves often with a truncate or even slightly tapering base,

244 Gard. Bull. Singapore 61 (2) 2010

Figure 2. Musa arfakaina with mature fruit.

245

A New Species of Wild Banana Musa arfakiana from Papua of Indonesia

lowe GGNER

Figure 3. Musa arfakiana. A. Habit of fruit bunch, peduncle and male bud; B. Whole fruit

and fruit in |.s. showing distal sterile pith chamber: C. Compound and free tepal: D. Stigma

with anthers; E. Leaf base: F. Petiole bases showing upper shoulder margins; G. Petiole (t.s.)

at the mid-point.

246 Gard. Bull. Singapore 61 (2) 2010

all bright green, hardly different in colour above and below, slightly paler

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

a little brown proximally, without wax. Petiole 76-80 x ca 3 cm, green, the

adaxial channel green, almost completely closed, the wings black or green

distally, TS ration 0.33 (see Argent, 1976). PB ratio 2-3.

Peduncle thick, green, glabrous, smooth. Bunch erect. The female bracts

lanceolate, yellow, shiny outside, dull yellow and slightly paler inside,

acuminate the apices with the margins strongly inrolled, quickly deciduous

or sometimes trapped between the fruit. Female flowers hermaphrodite with

fully fertile anthers, cream with green tips. Ovary trilocular, each loculus

with the ovules in two rows. Fruit bunch dense, sub-spherical in shape to ca

30 x 30cm. Fruits in two rows, the second hand with ca 10 fruits. The fruits

irregular, apparently ageotropic, showing no curvature in any part of the

bunch, ripening bright orange, up to 8 x 5 cm, very strongly 2 or 3-angled,

tapering in the distal half to a broadly acute apex with a prominent scars ca

1 cm in diameter, not splitting, sterile in the distal third, the seeds confined to

the proximal two-thirds of the fruit, with orange pith and similarly coloured

or yellowish flesh around the seeds in the carpel chambers. Pedicel very short

ca 4-5 mm, the fruits almost sessile. Seeds dark brown 5-7 mm in diameter,

irregularly angled, and with only an indistinct domed boss opposite the

impressed hilum, which is ca 2.5-3mm in diameter and smooth.

Male peduncle growing vertically upwards or mostly angled upwards at

about 30° to the vertical, rough with a dense covering of bract and flower

scars. The male bud up to 15 x 9 cm, pale yellow or green, shiny yellow,

convolute or only imbricate for ca 1 cm from the tip. Male bracts up to

16 x 9cm, shiny pale yellow outside, shiny yellow inside but becoming dull

inside after falling, with broadly rounded, obtuse apices, lifting to a wide

angle to ca 45° below the horizontal; after falling only recurved at the base

not revolute from the apex or margins. Male flowers, two-rowed, cream, the

free tepal translucent white, with a rounded, but irregular upper margin and

no distinct sub-apical wrinkle, about half as long as the compound tepal.

Compound tepal cream with green apices.

Musa arfakiana is similar to Musa johnsii Argent, in the dense sub-spherical

bunch of almost sessile fruits, with sterile distal pith chambers. This new

species differs from M. johnsii in its smaller stature; in having the petiole

canals almost closed with the ‘wings’ inflexed not broadly open; in having

erect, not hanging bunches of fruit; the fruits tapering distally to broad

points, not expanding distally to broad truncate apices and the fruit is not

schizocarpic as it is in M. johnsii.

Vernacular name ‘Bulada’ in the local language of Kampung Siobri, where

A New Species of Wild Banana Musa arfakiana from Papua of Indonesia 247

the people did not eat any part of the plant in contrast to the the Amungme

Tribe whose people eat Musa johnsii as a vegetable (Argent, 2001).

Notes: Named after the mountain range on which it was found.

This very distinctive new species was a very surprising find above the

village of Siobri in the Arfak Mountains and may have a very restricted

distribution as it was not found around Anggra village which was very close

to Siobri, effectively on the other side of a main ridge of mountains. A single

population of Musa arfakiana was seen on the side of Mt. Sembiedip from

ca 1500-1800m. It occurred in the upper area of secondary forest, which was

regenerating after timber extraction and gardening but also occurred in the

relatively undisturbed lower montane forest, sometimes in quite deep shade.

The population comprised numerous plants in various stages of flowering

and fruiting and like M. johnsii, it would appear to be non-seasonal. There

was no evidence of anything eating the fruit, which remained very tough and

hard even when the seeds were apparently mature, but the seeds brought

back to Edinburgh failed to germinate. Nor was there any evidence of the

fruit splitting open when ripe and this was confirmed by our local guide from

the village, *Zeth’ who had a good knowledge of the local plants and readily

confirmed that the fruit of some other species did split open at maturity.

Nothing appeared to eat the fruit and thus the dissemination of the seeds

remains a mystery.

This species is undoubtedly related to M. johnsii having a similar sterile

pith chamber in the distal portion of the fruit, which until the discovery of

this new species was thought to be a character unique to M. johnsii. It is

also similar in its waxless foliage and spherical bunch shape. M. arfakensis

significantly differs from M. johnsii in the almost closed petiolar canals, the

shorter petiole to blade ratio having relatively longer petioles. M. arfakensis

also has an erect fruit bunch with semi-erect male axis, unlike the half-

hanging bunch with vertically descending the male axis in M. johnsii. The

fruit, although with a similar sterile distal portion, tapers distally, unlike the

truncate distal apices of M. johnsii and there was no evidence that the fruit

ever naturally splits open. The basal flowers in M. arfakensis are functionally

hermaphrodite, not female, and the tips of the compound tepals are green

and not cream as in M. johnsii.

Our local guide at Kampung Siobri claimed to recognise four other

different wild bananas, all growing in the vicinity. ‘Binput’ was Musa ingens

Simmonds; ‘Bunkan’ was M. acuminata Colla subsp. banksii but the other

two were not seen and could not be identified from the local descriptions.

Musa ingens was commonly seen on the road at higher elevations, distinctive

in its large size, tapering pseudostems and waxy leaves. It was also seen

248 Gard. Bull. Singapore 61 (2) 2010

on Yappen island, both these records extend the known distribution of

this species away from the main range that forms the spine of mountains

east-west along the island of New Guinea. Additionally Musa schizocarpa

Simmonds was seen on the road to Siobri (Arfak Mts.).

Acknowledgements

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

Giner has ably drawn the pen and ink illustration. Zeth our local guide freely

shared his local knowledge with our expedition and to the other members of

the team who participated in the collection of this plant: Louise Galloway,

Sadie Barber and Andrew Ensoll and our Indonesian counterparts: Dr.

Charlie Heatubon and Pak Rustandi. Financial assistance was provided by

the Royal Botanic Garden Edinburgh, especially “The Members Committee’,

The Royal Horticultural Society and the James and Eve Bennet Trust. We are

grateful to LIPI the Indonesian Institute of Sciences for their support.

References

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

Botanic Garden Edinburgh 35: 77-114.

Argent, G. 2001. Contributions to the Flora of Mount Jaya VI. A New

Banana, Musa johnsii (Musaceae) from New Guinea. Gardens’ Bulletin

Singapore 53: 1-7.

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

Simmonds, N.W. 1966. Bananas, 2” Ed. Longmans, U.K.

Gardens’ Bulletin Singapore 61 (2): 249-268. 2010 249

A Review of the White-flowered Amorphophallus (Araceae:

Thomsonieae) Species in Sarawak

PC.BOYCE . LB. IPOR AND WL.A. HETTERSCHEID >

‘Visiting Scientist, Pusat Pengajian Sains Kajihayat

(School of Biological Sciences), Universiti Sains Malaysia

1180 USM, Pulau Pinang, Malaysia

Email: phymatarum@gmail.com

“Herbarium (SAR), Forest Research Centre, Km 10 Jalan Datuk Amar Kalong

Ningkan, 93250 Kuching, Sarawak, Malaysia

* Wageningen University Botanical Gardens, Gen. Foulkesweg 37

6703 BL Wageningen, Netherlands

Email: wilbert.hetterscheid@wur.nl

Abstract

A review of the white-flowered Amorphophallus species in Sarawak is

presented. A total of five species are recognized, four of which belong to

the Eburneus Group and are restricted to limestone, and moreover, locally

endemic: A. eburneus Bogner (Padawan and Tebedu areas), A. brachyphyllus

Hett. (Bau), A. juliae sp. nov. (Merirai) and A. niahensis sp. nov. (Niah).

A fifth species, Amorphophallus infundibuliformis Hett., A.Dearden &

A.Vogel, of doubtful affinity, is widespread and locally abundant on a variety

of substrates excluding limestone. A key to the white-flowered species in

Sarawak is presented and all species are illustrated.

Introduction

Fieldwork on forested limestone areas of Sarawak is proving remarkably

productive in revealing hitherto undescribed species of Amorphophallus.

As this paper exemplifies, even supposedly well-botanized areas can be

revealed to have new taxa and, thus, it is no great surprise that when remote

and not easily accessible limestone areas are scrutinized, these, too, prove to

have their complement of novel species.

Batu Niah is a significant limestone formation in Miri Division,

northeast Sarawak. Bukit Merirai is a smaller but more remote limestone

formation on the border of Kapit and Bintulu Divisons. Both areas are

geographically separated from the limestones of Mulu (Miri and Limbang

Divisions) and from the isolated Bukit Sarang limestones on the Bintulu/

Kapit Division border, and from areas further east in Sarawak and Sabah,

as too from the complex limestone formations in western Sarawak that

250 Gard. Bull. Singapore 61 (2) 2010

have received the most attention in recent years. Fieldwork in these areas

has shown that while such formations have aroid species in common, there

are also for each formation, suites of closely related but morphologically

distinct, presumably vicariant, local endemics. In addition to examples

in the genus Schismatoglottis (see especially Hay and Yuzammi, 2000),

and Alocasia (see Hay, 1998, 2000), this phenomenon is particularly well

exemplified by Amorphophallus, in which the species on limestone in

western Sarawak [A. eburneus Bogner (Bau limestone), A. brachyphyllus

Hett. (Padawan limestone), and from Serian (A. ranchanensis Ipor, Tawan,

A.Simon, Meekiong & Faud)] are mirrored in eastern Sarawak by the

recently described Mulu-endemic, A. julaihii Ipor, Tawan & P.C. Boyce, the

two novel species here described, and thence to Sabah where A. tinekeae

Hett. & A. Vogel is restricted to the limestone at Gua Gomontong.

The two novelties described here take to 17 the number indigenous

endemic Amorphophallus species recorded for Borneo (see also Mayo and

Widjaja, 1982; Bogner et al., 1985; Bogner, 1989; Bogner and Hetterscheid,

1992; Hetterscheid, 1994; Hetterscheid and van der Ham, 2001; Ipor et al.,

2004, 2007). Remarkably, 15 of these species have been described within the

. past 30 years. Additionally, a further four species recorded for Borneo are

either introduced or of doubtfully native provenance: A. konjac K.Koch,

A.paeoniifolius (Dennst.) Nicolson, A. prainii Hook. f.and A. muelleri Blume.

Taxonomy

The limestone-obligate white-flowered Amorphophallus in Sarawak are

seemingly closely related, and are currently referred to as informal Eburneus

Group. The common characters, admittedly polythetic, for the Eburneus

Group, are: irregularly bulging tubers, presence of corky outer tuber skin,

turgid petioles, at least upper male flowers vertically aligned, inflorescence

with a fishy smell at anthesis, and large (to 2 cm long) elongate fruits that,

except for those of A. titanum (Becc.) Becc. (5-6 cm long), are the largest in

the genus.

The systematic position of non-limestone A. infundibuliformis

is unclear. The leaf, especially the intricate morphology of the petiole

ornamentation, is quite different to the turgid, smooth, usually unmarked

light green, rarely with smooth paler green or reddish circular markings

typical of the Eburneus Group. The tuber is also markedly different and

striking by the inside flesh deep red.

The use of aninformal subordinate grouping is in line with the approach

used in other taxonomically intractable groups (e.g., Alocasia G.Don.,

Schismatoglottis Zoll. & Moritzi, the Potheae Engl., and Rhaphidophora

Hassk.), where the establishment of informal groups has become a standard

A Review of the White-flowered Amorphophallus Species in Sarawak 75)

approach until such time as phylogenetic testing can be undertaken leading

to the establishment of evolutionarily robust groups (see Boyce and Wong,

2008 for commentary on this approach). Other species in the Eburneus

Group are A. julaihii (Mulu), A. hottae Bogner & Hett., and A. palawanensis

Bogner & Hett. in the Philippines (Palawan). All except A. hottae are

limestone obligates.

Key to the white-flowered Amorphophallus species in Sarawak

1. Petioles turgid, smooth, rachises of fully developed leaf spreading.

Spadix appendix smooth, rugulose or echinate but never with hooked

Ree OCC SME AIMS Ol MIMIC SEOMEC 2.2, ca seceecceuhecdoe ee oe sctecdsucusetenesceeeeetlocdecs Goins 2

1. Petioles not turgid, conspicuously white-warty, rachises of fully developed

leaf ascending. Spadix appendix with conspicuous hooked staminodes.

Plants of a variety of substrates but never on limestone .............eceseeeeeeeeee

are eee Re eR eect eos ccean eect escesadesnesscnesivennsa Os Ae INPUNGIDULIFOTMIS

2. Spadix appendix echinate. Male flowers not fused into longitudinal rows.

Spathe limb reflexing during anthesis. C and N.E. Sarawak ....... eee 3

2. Spadix appendix smooth or weakly rugulose. Male flowers fused into

longitudinal rows. Spathe limb remaining erect throughout anthesis. S.W.

ea eee ae see cea cone sud ansdd inarhauinsdeddebadeuedunedccasueasecuueiee 4

3. Petioles dull pale red with strongly demarcated irregular reddish brown

spots and elongated streaks. Opening of lower part of spathe strongly

recurving to form a conspicuous collar ca 1cm wide during anthesis, spathe

limb margins recurving markedly. Spadix appendix stongly echinate.

ee NaINIc a) MP Ne a an ee leew cz oi cais savvedoc usc unodhedeecbweddennatihee 4. A. juliae

3. Petioles pale to medium green, concolorous or very occasionally with

obscure paler green circles. Opening lower part of spathe not or only

minutely recurved, spathe limb incurved or planate, margins not recurving.

aa MN cle re ire esc ade ge eeccs eo tdec, «ccsnkedveasacencs.2iededntestiecesteess Dn) Ae IQHENSIS

4. Petiole to 120 cm, long relative to the lamina diameter; lamina little divided,

leaflets to 60 cm long, never petiolulate; cataphylls greyish brown. Ovaries

densely congested, dark purple, stigma diameter equalling ovary. Padawan

LUTTE OUIUGS, ope eda sam Re ee 2. A. eburneus

4. Petiole to 50 cm, very short relative to the lamina diameter; lamina much

divided, leaflets to 35 cm long, terminal leaflets petiolulate; cataphylls off

white. Ovaries distant, pale purple, stigma diameter distinctly smaller

PAMvOVALY. I> all WMESLOMES c5ncseseegs sarees sccewcsrneosethsthecs 1. A. brachyphyllus

Dy: Gard. Bull. Singapore 61 (2) 2010

1. Amorphophallus brachyphyllus Hett., Blumea 46(2): 258 (2001 ).— Type:

Malaysia, Sarawak, Kuching, Division, Bau district, exact locality unknown,

(described from plant cultivated in Hortus Botanicus, Leiden); orig. coll. P.

Kessler EVK 246 sub. Hetterscheid H.AM.032C-T (holo, L, spirit coll.). Fig.

Medium-sized, robust aseasonally dormant geophytic herb to 90 cm. Tuber

depressed globose, not offsetting, with irregular raised areas, to 32 cm diam.

x15 cm high, surface with a grey, corky layer. Leaf solitary; petiole short, to

50 x 5 cm diam., uniformly green, very turgid; lamina to 188 cm diam., highly

dissected, rachises naked; leaflets elliptic-lanceolate, to 35 x11 cm, those on

the most proximal parts of the rachises petiolulate, upper surface mid-green,

slightly glossy or dull, slightly coriaceous. Inflorescence solitary, rarely two

together, short pedunculate; cataphylls off-white; peduncle 8-13 x1- 2.2 cm

diam., entirely subterranean, white with a faint greenish flush, smooth, very

tightly enveloped by the cataphylls; spathe erect, suborbicular, often broader

than long, 10-13.5 x11.5-16 cm diam., limb obliquely spreading at female

anthesis, erect at male anthesis, lower part tubular, strongly convolute, largely

hidden in cataphylls, spathe exterior off-white, occasionally flushed pale reddish

purple on the inside limb margins, interior with base reddish purple, and with

scattered small warts, or coarsely grooved, with grooves distinctly verruculate.

Spadix longer than spathe, stipitate, 13.5-21 cm long; stipe massive, oblique, off-

white, 0.6-1 x 1.6 cm diam. (base); female flower zone 1.5-2.5 cm x1.5-2.3 cm

diam., slightly conic, flowers in vertically separate sinuous chains; male flower

zone conic, 3-4.5 x 1.1-2.2 cm diam., flowers arranged as female flowers but

chains closer together, or partly or entirely fused vertically, sometimes forming

vertical chains; appendix fusiform, 8-14 x 1.2-2.8 cm diam., slightly laterally

compressed, subacute, yellowish white, surface rugulose and with narrow,

shallow grooves, producing a strong smell of fried fish and oozing out droplets

at female anthesis. Pistil (female flower) with an ovate or slightly depressed

ovary, 2-3 mm diam. x 2.5-3 mm high, base off-white, top dirty reddish brown,

unilocular, one basal ovule; style excentrically placed, consisting of three acute

branches, two acroscopic small ones and one basiscopic longer one, pale dirty

reddish brown, ca 1.5 mm diam. x 0.3-0.8 mm long; stigma thin, ca 1.5 mm

diam. x ca 0.5 mm high, irregularly, shallowly lobed-sinusoid, surface very

pale dirty brownish, verruculate. Male flower consisting of ca 3 stamens but

pattern often obscured by lateral and vertical fusion of flowers; stamens ca 1

mm high, x ca 1-2 mm diam., often fused with adjacent stamens; filaments ca

0.5 mm long, entirely connate; anthers ca 0.5 mm long, truncate, often entirely

connate, ivory-white; pores apical, rounded, oval or variously elongate, often

confluent with adjacent pores in various ways. Fruit a very large, elongate,

slightly angulate berry, up to 4 x 2 cm, ripening red, 1-seeded. Seeds elongate-

A Review of the White-flowered Amorphophallus Species in Sarawak

Figure 1. Amorphophallus brachyphyllus Hett. A. Flowering plant in habitat, note the erect

spathe limb and rugulose spadix appendix; B. Inflorescence with spathe artificially opened.

Note the scattered pistils, and the male flowers fused into longitudinal rows; C. Mature

leaf. Note the complex lamina division and the distal-most leaflets are petiolate; D. Newly

emerging leaf. Note the unmarked, turgid petiole; E. Mature infructescence. [Images: A, C-E

Gard. Bull. Singapore 61 (2) 2010

oblongo-conical, ca 3 cm x 1.2 cm diam. at the point of germination, 0.7 at the

opposite end, skin brownish.

Distribution: East Malaysia, Sarawak,endemic to Karst limestone formations

in the Bau area of Kuching Division.

Ecology: On rocky lowland forested limestone slopes under perhumid to

everwet evergreen forest, in humus layer or humus-filled pockets, less often in

clay at the base of limestone formations, 15-60 m asl.

Notes: Amorphophallus brachyphyllus is without doubt closely related to A.

eburneus and the inflorescences are deceptively similar, although separable

by these characters: stigma of A. brachyphyllus is half the size of that of

A. eburneus, while the pistils are much more regularly placed and more

congested in A. eburneus. In addition to these small but consistent differences

there is a marked difference in leaf morphology. In A. eburneus the leaf

has a long petiole (to 120 cm) relative to the lamina diameter (the reverse

in A. brachyphyllus). Moreover, the lamina in A. eburneus is considerably

- less strongly divided, with the leaflets distinctly larger (to 60 cm) and

never petiolulate. Cataphylls in A. eburneus are greyish brown, those in A.

brachyphyllus off-white. These vegetative differences remain constant, and

coupled with geographical separation, and differences in the inflorescence,

strongly support the recognition of two discrete but closely related, probably

vicariant, taxa. Amorphophallus niahensis, described elsewhere in this paper

is also in this complex, as too is A. juliae.

Etymology: The epithet brachyphyllus (Greek: brachy - short; phyllus - leaf)

refers to the very short petiole length relative to the diameter of the lamina.

Other specimens seen: MALAYSIA. Sarawak. Kuching Division: Bau, Gua

Angin, 5 Jun 1999, C.C. Lee AM-21.1 (SAR); Bau, Jambusan, 26 May 2004, PC.

Boyce & Jeland ak Kisai AM-31 (SAR); Bau, Gunung Bidi, 01° 23’ 27.0”; 110°

07° 07.6”, 7 Dec 2004, P.C. Boyce & L. Jenkins AM-88 (SAR); Bau, without

further locality data: Hetterscheid H.AM.031A (L).

2. Amorphophallus eburneus Bogner, Willdenowia 18: 441 (1989). — Type:

Malaysia, Sarawak, ‘near Padawong, north of Baw’ (see notes below), Bogner

1772 [described from plant cultivated in Munich Botanical Garden; orig.

coll. York Meredith s.n. sub. Bogner 1772 (holo, M)]. Fig. 2.

Medium-sized to large, robust, aseasonally dormant geophytic herb to 1.3

m. Tuber depressed-globose, not offsetting, to ca 35 cm diam. x ca 20 cm high,

in)

A Review of the White-flowered Amorphophallus Species in Sarawak

Figure 2. Amorphophallus eburneus Bogner. A. Flowering plant in habitat, note the erect

spathe limb and rugulose spadix appendix; B. Inflorescence with spathe artificially opened.

Note the scattered pistils, and the male flowers fused into longitudinal rows; C. Mature leaf.

Note the simple lamina division and no leaflets are not petiolate; D. Mature leaf, infructescence

and seedling. Note the unmarked, turgid petiole; E. Mature infructescence. [Images A-C & E

256 Gard. Bull. Singapore 61 (2) 2010

pale greyish brown, skin corky, broken up in numerous, small, angulate fields

separated by narrow grooves. Leaf solitary; petiole very turgid, to ca 120 x ca

10 cm diam., smooth, uniformly pale green or occasionally with a few scattered

whitish greenish spots; lamina moderately dissected, up to ca 240 cm, diam.,

rhachises winged distally from the basal branches; leaflets elliptic or elliptic-

lanceolate, 12-60 cm long, 5.5-21 cm diam., mid or pale green, coriaceous,

long acuminate. Inflorescence solitary, short pedunculate, largely hidden in

cataphylls; cataphylls ca 6, lanceolate, pale olive brown to whitish or greenish

with pale brownish flushes, largest up to ca 35 x 10.cm;peduncle 15-20 x ca2.5cm

diam., pale green, entirely hidden by cataphylls; spathe erect, infundibuliform,

ovate, base strongly convolute and hidden by cataphylls, separated from limb

by a shallow constriction, limb slightly spreading, acute, margins revolute, 20-

23 x 19-21 cm, outside creamy white, upper margins sometimes flushed with

purple, inside base dark purple and then cream, limb cream with or without a

pale purple flush, base within with numerous irregular, shallow warts and some

shallow grooves. Spadix slightly longer than spathe, shortly stipitate, 21-26 cm

long; female flower zone slightly conic, 2.5-3 x 2-2.5 cm diam., flowers slightly

or variably distant; male flower zone cylindric, 4-5 x 1.5-2 cm diam., flowers

- congested; appendix 14-18 x 1.5-3 cm diam., terete or laterally compressed,

acute, cream, turning pale yellowish during male anthesis, surface rugulose and

with scattered, short, shallow grooves, producing a smell of fish during female

and male anthesis. Pistil (female flower) comprised of a depressed pyriform

or oblong ovary, 2.5-4 diam.x ca 4-5 mm high, uni- or rarely (?) bilocular, top

divided in three narrow lobes, base white or pale green, remainder dark purple;

stigma thin, strongly sinuous, stellate, + 2-lobed, partly sunken in between the

three lobes emanating from the ovary, surface densely scaberulate, dirty grey-

ish stained with purple, ca 1.5-2.5 diam. x ca 0.5-1 mm high, + quadrangular in

cross-section. Male flower consisting of 4-6 stamens, often elongate parallel to

the spadix-axis and connate with upper and/or lower flowers; stamens ca 1 mm

long x ca 1-2 mm diam.; filaments ca 0.5 mm long, connate; anthers ca 0.5 mm

long, free or connate, truncate or subtruncate, pores apical, free or connected

within one flower or with other flowers and then irregularly elongate. Fruit a

berry, very large, elongate, slightly angulate, 4.5 x 2 cm, red, 1-seeded. Seeds

elongate conical, 3.5 cm long x 1.2 cm diam. at the point of germination, 0.7 at

the opposite end, testa brownish.

Distribution: East Malaysia, Sarawak, endemic to Karst limestone formations

of Padawan (Kuching Division) and Tebedu (Samarahan Division).

Ecology: Perhumid lowland forested limestone, growing deep in limestone

cracks with leaf litter 25- 350 m asl.

A Review of the White-flowered Amorphophallus Species in Sarawak Day,

Notes: There are two errors in the type locality cited by Bogner (1989).

Padawan (Padawong, sic) is southeast, not north, of Bau. The limestones of

the Padawan area are both geographically and floristically distinct from the

Bau limestones and extend down to the Kalimantan border in the direction

of Tebedu and Serian (Samarahan Division), with their northern boundary

to the east of the Bungo range. Amorphophallus eburneus is restricted to

these limestones. For differences between A. eburneus and A. brachyphyllus

see under that species.

Etymology: The epithet eburneus (Latin: ivory white) is in allusion to the

white spathe and spadix.

Other specimens seen: MALAYSIA. Sarawak. Kuching Division: Padawan,

Gunung Braang, 2 May 2001, CC. Lee AM-67.1, AM-67.2 (SAR); Padawan,

Bukit Manok, 01° 12’; 110° 18’, 18 Mar 2004, PC. Boyce AM-67.3 (SAR);

Padawan, Vogel 940011 (L, cult. in Hortus Botanicus, Leiden, sub. Hetterscheid

H.AM.402); Vogel 940012 (L, cult. Hortus Botanicus, Leiden, sub. Hetterscheid

H.AM.394): Vogel 970618 (L, cult. Hortus Botanicus, Leiden, sub. Hetterscheid

H.AM.893); Padawan, Gunung Penrissen, Vogel s.n. (L, cult. Hortus Botanicus,

Leiden, sub. Hetterscheid H.AM.311). Samarahan Division: Serian, Pichin,

Umon Murut, Tiab Belanting, 01° 08’ 03.7”; 110° 27° 00.3”, 15 Jun 2005, PC.

Boyce s.n. (image record SAR); Serian, Mongkos, Kampung Batuh, Gunung

Selabur, 00°57’ 26.2”; 110° 30’ 15.8”, 15 Mar 2006, PC. Boyce s.n. (image

record SAR);

3. Amorphophallus infundibuliformis Hett., A.Dearden & A.Vogel, Blumea

39(1-2): 259 (1994). — Type: Malaysia, Sarawak, Kuching Division, without

further locality, 1990, Dearden s.n. (holo, L). Fig. 3.

Medium-sized, slender, aseasonally dormant geophytic herb to 1.3 m. Tuber

depressed-globose to subglobose, pinkish externally, internally deep red,

without offset development. Leaf solitary; petiole, pale grey-green to dark

green to dark green-purple, with dirty whitish with numerous, confluent,

irregular, green spots and scattered white punctiform dots and patches, these

more or less raised; lamina weakly dissected, up to 35 cm diam., thinly

coriaceous, adaxial surface deep, sometimes weakly metallic, glossy green,

abaxial surface paler, rachises naked, ascending, very shallowly and narrowly

canaliculate, greyish-green with irregular dark green patches; leaflets

lanceolate, margin with numerous small undulations, main veins impressed,

2-11 x 2-5 cm, shortly petiolulate, petiolule 0.2-1 mm long, very slightly

canaliculate adaxially, lowermost pair of leaflets symmetrical, all others

asymmetrical and obliquely inserted on petiolule, apex shortly acuminate to

258 Gard. Bull. Singapore 61 (2) 2010

Figure 3. Amorphophallus infundibuliformis Hett., A.Dearden & A.Vogel. A. Flowering

plant in habitat, note the funnel-form erect spathe; B. Inflorescence with spathe artificially

opened. Note hooked staminodes covering much of the spadix appendix; C. Detail of petiole

to show the diagnostic raised white warts; D. Mature leaf. Note ascending rachises; E. Mature

infructescence. [Images A-E © Peter Boyce].

A Review of the White-flowered Amorphophallus Species in Sarawak 759

long acuminate, up to 1.5 long; margins of larger leaflets very slightly sinuate;

5-8 pairs of primary lateral veins, venation forming distinct submarginal

veins; interprimary veins much less defined, secondary veins forming a

weak reticulum. Inflorescence solitary, short pedunculate; cataphylls whitish

with numerous small, pinkish dots and many larger, blackish green, irregular

spots; peduncle 4 cm x ca 0.8 cm diam. (base), lengthening in fruit; spathe

strongly convolute, funnel-form, obconic in side-view, transversely orbicular-

elliptic, 6 x ca 8 cm diam., very widely acute, limb and base poorly differen-

tiated, outside dirty whitish with pale brownish venation and scattered, small,

angulate, blackish green spots, inside whitish, the lower half dark maroon, base

within strongly, longitudinally ridged. Spadix sessile, very obliquely inserted,

slightly longer than spathe; female flower zone oblique, annuliform, 1 mm

(dorsal side)-5 mm (ventral side) long x ca 1 cm diam., flowers congested;

male flower zone cylindric, base oblique, 0.8-1.3 cm ca 1 cm x diam., flowers

congested; appendix cylindric, obtuse, whitish, 6.5 x 1 cm diam., entirely or

at least almost entirely, densely covered with hooked staminodes, these in

the lower third shortly conical to aristate, sometimes uncinate and with long,

narrowly decurrent, ridge-like bases, upwards shorter or reduced to only the

base, up to 2 mm long, bases longitudinally confluent. Pistil (female flower)

with ovaries depressed, irregular or cubic, angulate in cross-section, 0).9-1.5

x 1.2-1.5 mm, reddish brown, near the style insertion maroon, unilocular;

style absent or only basiscopically developed; stigma sessile or partly sessi-

le, acroscopically orientated, reniform, a shallow depression in the middle,

one conic lobe on the outward facing margin, ().8-1 mm x 0.3-0.5 mm high,

dark greyish brown, surface densely verruculate. Male flower upwards fused

into longitudinal chains, otherwise consisting of 3-5 stamens, upper flowers

confluent with the lowermost staminodial ridges; stamens ca 1-1.3 x ca 0.7

mm long, rounded, oval or irregular in cross-section, white; filaments absent or

nearly so, entirely connate; anthers truncate; pores apical, rounded or elongate

(confluent). Infructescence with few to rather many berries, irregular; berries

elongate, ca 2 x 0.75 cm, slightly conic, top truncate, orange-red, one-seeded.

Seed elongate ellipsoid, ca 1.8 x 0.5 cm, testa pale brown.

Distribution: East Malaysia, Sarawak (Kuching, Sri Aman & Kapit Divisions

but probably throughout the state and overlooked); Indonesia, Kalimantan

Barat.

Ecology: Perhumid to everwet lowland mixed dipterocarp to upper hill forest,

mainly on sandstones, occasionally on shale, rarely on raised podzols, 50-875

m asl.

Notes: Amorphophallus infundibuliformis cannot be mistaken for any other

260 Gard. Bull. Singapore 61 (2) 2010

species in the genus in Sarawak by virtue of the spadix appendix with hooked

staminodes. The petiole, with ascending rachices and intricately ornamented

with dark patches and conspicuous raised, white warty areas is approached

by that of A. ranchanensis,in which the raised areas are scutteliform and pale

grey-green. The inflorescences of A. infundibuliformis and A. ranchanensis

are profoundly dissimilar.

Etymology:The specific epithet is from the Latin for funnel-shaped, referring

to the shape of the spathe.

Other specimens seen: MALAYSIA. Sarawak: Kuching Division: Lundu,

Gunung Gading, 01° 42'; 109° 50', 3 Mar 2004, P.C. Boyce & Jeland ak Kisai

AM-3 (SAR); Bau, Segong, Sungai Adis, 11 Mar 2004, P.C. Boyce & Jeland

ak Kisai AM-4.1(SAR); Bau, Kampung Jugan, 26 Mar 2004, PC. Boyce &

Jeland ak Kisai AM-4.2 (SAR); 22 May 2004, PC. Boyce, Jeland ak Kisai

& Jipom ak Tisai AM-33 (SAR); Padawan, Puncak Borneo, trail behind

Malesiana Tropicals Nursery to Hornbill Resort golf course maintenance

kampong, 01° 07' 35.1"; 110° 13° 28.8”, 10 Jun 2004, PC. Boyce & Jeland ak

~ Kisai AM-36 (SAR); 01° 07' 35.1"; 110° 13' 28.8", 30 Nov 2004, PC. Boyce

AM-87 (SAR); Sematan, Teluk Selabang Ulu Sungai, Selabang, 8 Oct 2004,

P.-C. Boyce & Jipom ak Tisai AM-S1I (SAR); Bau, Gunung Noka, 11 Oct

2004, P.-C. Boyce & Jeland ak Kisai AM-84 (SAR); Lundu, Brungea, 8 Jan

2005, P.C.Boyce & Jipom ak Tisai AM- 92 (SAR); Bau, Kampung Duyoh,

Sungai Duyoh, 01° 20' 45.6"; 110° 02' 36.9", 8 Jun 2005, PC. Boyce & Jeland

ak Kisai AM- 95 (SAR); Matang, Kubah N.P., Waterfall Trail, 7 Mar 2009,

P.C. Boyce (SAR, image record). Sri Aman Division: Lubok Antu, Batang

Ai, Nanga Sumpa, 01° 12' 02.3"; 112° 03' 09.3", 27 Jul 2004, PC Boyce, Jeland

ak Kisai & N.Lembang AM-43 (SAR); Lubok Antu Batang Ai, Nanga

Sumpa, Sungai Pedal, 01° 111" 58.9"; 112° 0322702, 7 Apr 2005S F @beyee

et al. AM-94 (SAR); Sri Aman, Lubok Antu, Batang Ai, Nanga Sumpa,

Wong Ensalai, 01° 11' 51.0"; 112° 03' 39.9", 26 May 2008, PC. Boyce, Wong

Sin Yeng & Jipom ak Tisai AM-200 (SAR). Kapit Division: Kapit, Taman

Rekreasi Sebabai, 01° 56’ 45.6"; 112° 34" lo:8", 113 Dee 20047 A GeBovec

Jeland ak Kisai & M. Gibernau AM-89 (SAR). INDONESIA. Kalimantan:

Kalimantan Barat, Semeng, Sizemore 960031 (L, image record).

4. Amorphophallus juliae P.C. Boyce & Hett., sp. nov.

Ab omnibus speciebus in habitu calcicola lithophytica Borneensibus borealis

combinatio appendice spadicis echinatis et petiolorum foliis pallide rubro

et valde brunneis maculatis distinguitur; ab A. niahensis spathae lamina

marginem valde revolutis et reflexis et appendice spadicis profunde echinatis

differt.— Type: Sarawak, Kapit Division, Belaga District, Bukit Merirai, path

A Review of the White-flowered Amorphophallus Species in Sarawak 261

to Gua Tiang & Gua Spring, 2° 46' 07" N; 113° 38' 58" E, 6 Jul 2005, P. Leong,

R. Kiew, S. Julia et al. PL 135 (Holotypus, SAR; isotypus, SING.). Fig. 4.

Medium-sized, moderately robust, aseasonally dormant geophytic herb to

1.2 m tall. Tuber depressed globose, with irregular slightly raised areas, up

to 7 cm diam., 4 cm high, surface pale brown, interior white. Leaf solitary,

petiole proportionately short compared to the lamina diameter; petiole up to

35 cm long, ca 2 cm diameter at base, moderately turgid, cylindrical, smooth,

uniform dull pale red with strongly demarcated irregular reddish brown

spots and elongated streaks, subtended by 2-3 marcescent cataphylls, these

extending ca 1/3 the length of the petiole; lamina moderately dissected,

up to 55 cm diam., thinly coriaceous, adaxial surface mid green, abaxial

surface paler, rachises naked, narrowly canaliculate, pale pinkish green;

leaflets elliptic-lanceolate, 2-13 x 2-5 cm, more-or-less sessile, lowermost

pair of leaflets symmetrical, all others asymmetrical and obliquely inserted

or rhachis, apex shortly acuminate to 1.5 long; margins of larger leaflets

very slightly sinuate; 5-7 pairs of primary lateral veins, these adaxially

flush to slightly impressed, abaxially very slightly raised; venation forming

distinct submarginal veins; interprimary veins less defined, secondary veins

forming a weak reticulum. Inflorescence solitary, occasionally two together,

flowering before emergence of foliage leaves but in any one colony mature

plants at all stages of growth (emerging leaves to ripe infructescences)

present simultaneously; peduncle and lower part of spathe encased in sub-

fleshy cataphylls; peduncle cylindrical, up to 13 cm long, 5-6 mm diam., pale

green, pinkish where exposed to light; cataphylls several per inflorescence;

elongate-ovate to linear, 2-15 cm x 1.5 -3.5 cm, pale greenish white, sub

fleshy at anthesis, then soon withering and then decaying, drying mid brown.

Spathe broadly oblong-ovate, funnel-form, up to 9 cm long x 3 cm wide

(fresh), pressed material up to 7 cm diam.; spathe limb accounting for ca

1/3 or less of the spathe length, margins recurved at anthesis and at first

somewhat conspicuously green-veined; interior white, smooth, somewhat

glossy, exterior very pale greenish white to white; lower spathe convolute

and much inflated at anthesis, 2-7 cm long, interior muricate-verrucate,

deep reddish purple, exterior pale greenish white, sometimes slightly pink-

tinged, the margins recurving to form a conspicuous collar ca 1cm wide, this

somewhat glossy and frequently tinged and veined pale green at anthesis.

Spadix exceeding spathe, 9-12 cm long. Appendix up to 11 cm long, slender

elongate-fusiform, ca 9 mm diam. (fresh), ca 4 mm diam, (dried), white to

dull cream, very pronounced-echinate, producing a mild odour of rotten

fish during anthesis. Flowers unisexual; male flower zone weakly fusiform-

cylindrical, up to 3 cm x 6 mm diam., cream; stamens ca | mm long, ca

0.5 mm broad across, pores paired, mostly solitary, the reduction in pore

Gard. Bull. Singapore 61 (2) 2010

Figure 4. Amorphophallus juliae P.C.Boyce & Hett. A-B. Flowering plants in habitat, note

the strongly echinate spadix appendix, the recurved lower spathe margins and recur

spathe limb; C. Mature leaf; D. Emerging leaf. Note the conspicuous petiole markings; E.

Mature infructescence. [Images A-C, E © Julia anak Sang (used with permission). Image D

A Review of the White-flowered Amorphophallus Species in Sarawak 263

number proceeding upwards towards the appendix, pollen pale yellow.

Female flower zone cylindrical, contiguous with the maleflower zone, ca 4.5-

7 x ca 7 mm diam.; male flowers dense, not arranged in longitudinal rows;

pistils (female flowers) few, densely arranged, ovaries compressed globose,

ca 1.5 x 1.5 mm, dark purple, mostly bilocular; stigma sessile, conspicuously

three lobed. Infructescence with up to 50 berries, peduncle up to 18 cm long,

but mostly buries, ca 7 mm diam. at base, 9 mm diam. at apex, with a dark

brown V-shaped scar from the marcescent spathe. Fruit an ellipsoid berry

15-21mm x 8-10 mm, apex rounded, with conspicuous impressed blackish

stigma remnants, one seeded, deep orange at maturity. Seeds ellipsoid, 12-17

mm x 7-8.2 mm wide, testa smooth, thin, yellowish green.

Distribution: Endemic to Sarawak, so far recorded only from Bukit Merirai,

Belaga, Kapit Division.

Ecology: Lowland limestone forest, shady areas in humus-filled fissures and

holes in limestone, ca 60 m asl.

Notes: By the leaf with a proportionately short petiole compared with the leaf

lamina diameter, and in bearing numerous small leaflets Amorphophallus

juliae is vegetatively most similar A. brachyphyllus and A. niahensis. From

both, A. juliae differs in the pale reddish petioles with large darker reddish

brown spots (vs pale to medium green, concolorous or very occasionally

with obscure paler green circles in A.brachyphyllus and A. niahensis ).

The male flowers not fused into longitudinal rows, the subentire to

weakly bilobed stigma and echinate spadix appendix resemble those of A.

niahensis, although the echinate texture is markedly more pronounced in

A. juliae. Amorphophallus juliae is readily distinguished from A. niahensis

by the opening of the lower part of spathe strongly recurving to form a

conspicuous collar ca lcm wide during anthesis (margins not or only

minutely recurved in A. niahensis) and spathe limb margins recurving very

markedly.

Amorphophallus juliae is most readily distinguished from A.

brachyphyllus by the male flowers not fused into longitudinal rows, the densely

arranged pistils, the proportionately shorter spathe limb (comprising ca 1/3

or less of the entire spathe vs % or more of entire spathe), and the spathe

limb reflexing at male anthesis (vs. spathe limb erect throughout anthesis),

and subentire to weakly trilobed stigma (versus deeply bi-trilobed).

Etymology: Named for Julia anak Sang, a forest botanist from the Forestry

Research Department, Sarawak Forestry Corporation, Kuching, and co-

collector of the type specimen.

264 Gard. Bull. Singapore 61 (2) 2010

Other specimen seen: MALAYSIA. Sarawak, Bintulu/Kapit Division border,

Belaga District, Bukit Merirai, trail from Sungai Bekuyat to Gua Naga, 11

July 2005, P. Leong, R. Kiew, S. Julia et al. PL 270, (SAR, SING).

5. Amorphophallus niahensis P.C. Boyce & Hett., sp. nov.

Ad A. brachyphyllus spadix appendice echinatis, floribus masculinus nec in

serialis longitudinaliter tectis, stigmate sessile subintegris; spathae anthesin

feminibus reflexis, non erectis; ab A. juliae petiolorum foliis non pallide

rubro et maculatis, spathae lamina marginem nec valde revolutis differt. —

Type: Sarawak, Miri Division, Niah Suai District, Gunung Subis, Gua Niah

N.P., along plank walk to Niah caves, 21 Aug 2002, Julaihi L et al. S 89309

(Holotypus, SAR). Fig. 5.

Medium-sized, rather robust, aseasonally dormant geophytic herb to | m tall.

Tuber depressed globose, with irregular raised areas, up to 12 cm diam., 6 cm

high, surface pale brown, interior white. Leaf solitary, petiole proportionately

short compared to the lamina diameter; petiole up to 60 cm long, ca 2.5 cm

diameter at base, moderately turgid, cylindrical, smooth, uniform pale-green

- with a few obscure paler spots present in some individuals, subtended by 2-3

marcescent cataphylls, these extending ca 1/10 the length of the petiole;

lamina moderately dissected, up to 1 m diam., thinly coriaceous, adaxial

surface mid-green, abaxial surface paler, rachises naked except for terminal

leaftets long-decurrent, narrowly canaliculate, mid-green; /eaflets elliptic-

lanceolate, 2-15 x 1.5-6 cm, terminal-most petiolulate, petiolule 1-2.5 cm

long, narrowly canaliculate adaxially, lowermost pair of leaflets symmetrical,

all others asymmetrical and obliquely inserted on petiolule, apex shortly

acuminate; 3-9 pairs of primary lateral veins, these slightly impressed

adaxially, abaxially slightly raised; venation forming distinct submarginal

veins; interprimary veins less well-defined, secondary veins forming a weak

reticulum. Inflorescence solitary, occasionally two or rarely three together,

flowering before emergence of foliage leaves but in any one colony mature

plants at all stages of growth (emerging leaves to ripe infructescences)

present simultaneously; peduncle and lower part of spathe encased in sub-

fleshy cataphylls, peduncle cylindrical, up to 17 cm long x5-6 mm diam., pale

green; cataphylls several per inflorescence; elongate-ovate, 3-16 cm x 1-4

cm, pale greenish white, sub fleshy at anthesis, soon withering and decaying,

drying mid brown. Spathe broadly oblong-ovate, narrowly funnel-form, up

to 9 cm long x 3 cm wide (fresh), pressed material up to 5 cm wide; spathe

limb accounting for ca 1/3 or less of the spathe length, spathe mouth margins

hardly recurved at anthesis and spathe limb margins planate; interior white,

smooth, somewhat glossy, exterior very pale greenish white to white; lower

spathe convolute and much inflated at anthesis, 2-7 cm long, interior muricate-

A Review of the White-flowered Amorphophallus Species in Sarawak 265

Figure 5. Amorphophallus niahensis P.C.Boyce & Hett. A. Flowering plants in habitat, note

the echinate spadix appendix and recurved spathe limb; B. Inflorescence with spathe artifi-

cially opened. Note the warty lower spathe interior and the male flowers not in fused longi-

tudinal rows; C. Mature infructescence; D. Mature leaf. Note the complex division. [Images

A-D © Peter Boyce].

266 Gard. Bull. Singapore 61 (2) 2010

verrucate, deep reddish purple, exterior pale greenish white, sometimes

slightly pink-tinged. Spadix exceeding spathe, 9-12 cm long. Appendix up to

9 cm long, elongate-fusiform, ca 9 mm diam. (fresh), ca 4 mm diam, (dried),

white, moderately echinate, producing a sharp smell of rotten fish at anthesis.

Flowers unisexual; male flower zone cylindrical, up to 3 cm long,6 mm diam.,

cream; stamens ca | mm long, ca 0.5 mm broad across, pores paired, partially

fused or solitary, pollen dark yellow. Female flower zone shortly cylindrical,

contiguous with the male zone, ca 4.5-7 mm x ca 7 mm diam.; pistils few,

densely arranged; ovaries compressed globose, ca 1.5 x 1.5 mm, dark purple,

mostly bilocular; stigma sessile, three lobed. Infructescence with up to 35

berries, pedunculate up to 22 cm long, 6 mm diam. at base, 9 mm diam. at

apex, with blackish dark brown V-shaped scar from the marcescent spathe,

basally with remains of the cataphylls. Fruit ellipsoid 15-16 x 8-10 mm, apex

rounded, with blackish stigma remnants, when ripe deep orange, one seeded.

Seeds ellipsoid, 12-14 mm x 7-8.2 mm wide, testa smooth, thin, yellowish

green, seed copiously starchy, embryo small.

Distribution: East Malaysia, Sarawak, so far recorded only from Niah

. National Park.

Ecology: Limestone forest, growing shady areas in humus-filled fissures and

holes in limestone, often on limestone emergent in swampy areas, ca 45 m

asl.

Notes: Amorphophallus niahensis is most similar to A. brachyphyllus,

especially in the rather short petiole compared to the lamina diameter, but

is readily distinguished by the conspicuously echinate spadix appendix,

the male flowers not arranged in longitudinal lines, the concolourous deep

purple pistils, and the spathe limb reflexing at female anthesis.

Amorphophallus niahensis approaches A. juliae in the echinate

spadix appendix morphology (although the echinate texture is markedly

more pronounced in A. juliae), but is readily distinguished by the pale green

(not pink) petiole lacking any markings (vs reddish-brown spotted) and the

spathe limb margins not or only slightly recurving at anthesis.

Curious to note is that in nature the inflorescences very often produce

two morphologically normal spathes, set at 180° to one another. Similar

observations have been made with A. eburneus and A. brachyphyllus.

Etymology: The species is named for originating from Gua Niah, to which

it is endemic.

A Review of the White-flowered Amorphophallus Species in Sarawak 267

Other specimens seen: MALAYSIA. Sarawak, Miri Division: Niah Suai

District, Niah National Park, Gunung Subis, outside Great cave, 24 Apr

1972, J.A.R. Anderson § 31903 (SAR); Niah N.P., trail to Great cave, 03° 49’

09.9”; 113° 46’ 52.3”, 13 Oct 2005, PC. Boyce, Jeland ak Kisai & Jipom ak

Tisai AM-101 (SAR); Niah Suai District, Niah National Park, Madu Trail,

03°48’ 57.9”; 113° 4618.3”, 13 Jul 2006, PC. Boyce et al. AM-107 (SAR);

Subis, Gua Niah N.P., below W mouth of Great cave, 22 Aug 2002, K. Pearce

et al. S 89487 (SAR); Batu Niah. 113 46 E. 3 49 N, Vogel 970616 (L, cult.

Hortus Botanicus, Leiden, sub. Hetterscheid H.AM.895).

References

Bogner, J. 1989. A new Amorphophallus (Araceae) from Sarawak.

Willdenowia 18: 441-443.

Bogner, J.,S.J. Mayo and M. Sivadasan. 1985 (1986). New species and changing

concepts in Amorphophallus. Aroideana 8(1): 15-25.

Bogner, J.and W.L.A. Hetterscheid. 1992. Notes on the genus Amorphophallus

(Araceae) — 1. Three new species from tropical Asia. Blumea 36: 467-475.

Boyce, P.C. and S.Y. Wong. 2008. Studies on Homalomeneae (Araceae) of

Borneo I: Four new species and speculation on informal species group in

Sarawak. Gardens’ Bulletin Singaore 60: 1-29.

Hay, A. 1998. The genus Alocasia (Araceae-Colocasieae) in West Malesia

and Sulawesi. Gardens’ Bulletin Singapore 50: 221-334

Hay, A. 2000. Alocasia nebula. Botanical Magazine, n.s. 17(1): 14-18, pl. 381.

Hay, A. and Yuzammi. 2000. Schismatoglottideae (Araceae) in Malesia I -

Schismatoglottis. Telopea 9(A): 1-177.

Hetterscheid, W.L.A. 1994. Notes on the genus Amorphophallus (Araceae)

— 2. New species from tropical Asia. Blumea 39: 237-281.

Hetterscheid, W.L.A. and R.W.J.M. van der Ham. 2001. Notes on the genus

Amorphophallus (Araceae) — 11. New and obsolete species from East

Malaysia and continental Southeast Asia. Blumea 46: 253-282.

268 Gard. Bull. Singapore 61 (2) 2010

Ipor,1I.B.,CheksumT. and P.C. Boyce.2004.A new species of Amorphophallus

(Araceae: Thomsonieae) from Sarawak, Borneo. Gardens’ Bulletin

Singapore 56: 153-159.

Ipor, I.B., Cheksum T., A.Simon, K. Meekiong and A. Faud. 2007. A new

species of Amorphophallus (Araceae: Thomsonieae) from Sarawak.

Folia Malaysiana 8(1): 1-10.

Mayo, S.J. and E. Widjaja. 1982. Amorphophallus lambii. Curtis's Botanical

Magazine 184(2): 61-64, tab. 852.

Gardens’ Bulletin Singapore 61 (2): 269-317. 2010 269

Studies on Homalomeneae (Araceae) of Borneo II: The

Homalomena of Nanga Sumpa (Batang Ai) — Novel and

Pre-existing Taxa, and Notes on Iban Usages

PC. BOYCE , S.Y.WONG’ AND FASIHUDDINB.A.>

‘Visiting Scientist, Pusat Pengajian Sains Kajihayat

(School of Biological Sciences), Universiti Sains Malaysia

1180 USM, Pulau Pinang, Malaysia

: Email: phymatarum@gmail.com

- Department of Plant Science & Envionmental Ecology, Faculty of Resource

Science & Technology, Universiti Malaysia Sarawak, 94300 Kota Samarahan,

’ Sarawak, Malaysia

‘Deputy Dean (Postgraduate & Research), Faculty of Resource

Science & Technology, Universiti Malaysia Sarawak,

94300 Kota Samarahan Sarawak, Malaysia

Abstract

Fieldwork targeting indigenous Homalomena at Nanga Sumpa, part of the

Batang Ai drainage system (Sri Aman Divison, Sarawak) revealed 14 species

of which six are novel and herewith described: Homalomena atrox P.C.Boyce,

S.Y.Wong & Fasihuddin, H. clandestina P.C.Boyce, S.Y.Wong & Fasihuddin,

H. hanneae P.C.Boyce, S.Y.Wong & Fasihuddin, H. sengkenyang P.C.Boyce,

S.Y.Wong & Fasihuddin, H. symplocarpiifolia P.C.Boyce, S.Y.Wong &

Fasihuddin, and H. vivens P.C.Boyce, S.Y.Wong & Fasihuddin. Of the six

Homalomena species present at Batang Ai for which there are pre-existing

names, two, H. borneensis Ridl. & H. humilis (Jack) Hook.f., are species

complexes still awaiting a full taxonomic and systematic investigation, and

are treated here as morpho-taxa to which we apply the earliest applicable

epithet. The remaining four species with available names have only recently

been described: H. geniculata M.Hotta (1967); H. vagans P.C.Boyce (1994),

and H. josefii P.C.Boyce & S.Y.Wong, and H. pseudogeniculata P.C.Boyce &

S.Y.Wong (2008). Additionally, two further species located during fieldwork,

that while unarguably novel based on their vegetative morphology, were

not located as fertile plants and have yet to flower in cultivation; they are

here treated as sp. nov. A & B. Of the 14 species present at Batang Ai, five

have significance in the ethnobotany of the indigenous Iban people of the

Ai drainage, and of these five, four are novel. A key to the Homalomena in

the Batang Ai drainage area is given, and all species are illustrated.

270 Gard. Bull. Singapore 61 (2) 2010

Introduction

As noted in the first of these papers (Boyce & Wong, 2008), Homalomena

is the most abundant, speciose and conversely least well understood

mesophytic aroid genus in tropical Asia. It is also now becoming the focus

for pharmaceutical studies by virtue of the prevalence of uses amongst

Malaysian indigenous peoples mainly due to the presence of aromatic

compounds in almost all tissues.

That a taxonomic study is urgently required is no better exemplified

than by the work of Hanna Christensen (Christensen, 2000) in which five

Homalomena species are highlighted as having moderate to significant

importance as plants utilized by the indigenous Iban community of Nanga

Sumpa, Batang Ai, yet four of the five species are scientifically novel. These

are among the novelties dealt with here in this preliminary account of

Homalomena on the Ai drainage system.

Five of the species here newly described, namely, H. clandestina

P.C.Boyce, S.Y.Wong & Fasihuddin, H. hanneae P.C.Boyce, S.Y.Wong

& Fasihuddin, H. sengkenyang P.C.Boyce, S.Y.Wong & Fasihuddin, H.

- symplocarpiifolia P.C.Boyce,S.Y.Wong & Fasihuddin & H. vivens P.C.Boyce,

S.Y.Wong & Fasihuddin, belong in the informal Cyrtocladon supergroup,

and one, H. atrox P.C.Boyce & S.Y.Wong, in the Chamaecladon supergroup

(see Boyce & Wong, 2008).

Usage among Iban indigenous people at Nanga Sumpa

Christensen (2000) listed 6 Homalomena utilized by the Iban community

of Nanga Sumpa, although only one, H. borneensis (reported under the

synonym H. ovata in Christensen, 2000) is identified to species. Four of

the Homalomena utilized at Nanga Sumpa are known collectively by the

Iban as “kemuyang”; a fifth (here described as H. clandestina) is called

“subung tilan” , while a sixth species occurs wild with populations near the

longhouse artificially managed, here described as H. sengkenyang, is called

“sengkenyang bakung”.

The uses of the species known as kemuyang are rather generic (see

below under each relative species), while suwbung tilan is used soley to flavour

dishes such as fish. More interesting is the usages of the semi-managed

Homalomena known as sengkenyang bakung, whichis utilized as a protector

of rice plants. Plants utilized for sengkenyang are enormously important to

the Iban as they are believed to protect the rice plants against malevolent

spirits and prevent the beneficial rice spirits from leaving the farm. Very

few plants have the potential to be appointed to the status of sengkenyang.

The Homalomena of Nanga Sumpa Di

This species is here described as Homalomena sengkenyang (see below).

Key to the Homalomena of the Batang Ai drainage

faecal lamina base* truncate, cordate, or Sagittate ..2.....:ccscsiecessessecaaceszeaceeeee 2

Pee ai Aina DASE ACULE 10 CECULTENE -................-cscnseacsesseseedscabanseantactndasasdacte 6

NR eA AS SAMA eine cca anc dane oonnscsaneqnasscnssuadseaesscacshasensnecndasiecaideatens 3

ee ASC IGUMCALC OF COLGALG 22.25.02. <cecessncsiescaseoneeteceasonesoas senavecooeduseseeveseeacs 8

3. Pistils not associated with interpistillar staminodes ............ 12. H. vivens

3. Pistils associated with interpistillar staminodes ..................:cesscescesseesseneees 4

4. Spadix producing copious amber resin droplets at anthesis; pistils

RAN inde Saccnt sence vaneeeonessevansancs-ses.s Os Ld. annede

4. Spadix not producing resin droplets at anthesis; pistils various colours

I eID ONL a one ean sas Sagan ontee stone scvassaveucBiceNeddbaenees 5

5. Medium plants not exceeding 50 cm tall and often only half this height;

Pee DEP hE PTCCH: SHIPMACMLITE ...: :....0.002025....---c-00s000- 9. H. sengkenyang

5. Large plants often exceeding | m tall; pistils cream; stigma trisulcate

Peete co a aE Sat ock ocgesavtvsazecvaceranrisbideacsnce 7. H. josefii

6. Leaf lamina oblong, venation deeply impressed adaxially; spathe with a

conspicuous dorsal median ridge; female and male flower zones equal in

PAIL CE COULOUOUS 6-3. 222.2-022 0 s2eceecevssteeeeceetaacet-s 10. H. symplocarpifolia

6. Leaf lamina ovate to very weakly ovato-sagittate; venation not impressed;

spathe without a ridge; female flower zone exceeding male zone in

Rtas errr Pere eI SI obi svn dassbwn tats taa unseen REINS ff

7. Lamina abaxially glaucous; male flower zone more or less contiguou with

the female flower zone, female flower zone stoutly fusiform; pistils densely

arranged , not ascending, with stigmas coherent................2. H. borneensis

7. Lamina not glaucous abaxially; male flower zone separated from the

female flower zone by a brief naked interstice; female flower zone weakly

fusiform; pistils somewhat lax, ascending, stigmas not coherent ..................

eee cee eee de ee Perea EN ee eS, Clandestina

8. Leaves in a strictly distichous arrangement .................2. 4. H. geniculata

che LEE ESTA ESTOS 1 re T0177 ea 9

22, Gard. Bull. Singapore 61 (2) 2010

9. Diminutive plants, microscopically puberulent, with a velvety or

scintillating quality ..22<.s6.2.eecccteaseteeraee scenester eee 10

9. Larger plants, without a velvety or scintillating quality ................ccseee 11

10. Leaf lamina oblong-lanceolate to oblanceolate laminae, long-decurrent;

Spadix.sessile; top-most flowers!Steiile-.- ee eee 1. H. atrox

10. Leaf lamina oblong-ovato-sagittate, base shallowly cordate; spadix

Stipitate, fertile to the {ip zc ce eee ee 6. H. humilis

—

. Facultative rheophytes, lamina | pendent on the petiole, lanceolate-

elliptic 11-23.5 x 1.5-6 cm, infructescence with peduncle declinate and

Spathie ascend ing: .:)25. reser tee eee eee ee eee 11. H. vagans

. Terrestrial on slopes, lamina spreading, oblongo-lanceolate, sometimes

oblongo-elliptic to ovate, 18-33 cm long x 6-15 cm Wide «0.0... ceeeeeeeeeeees

wean dvi ug vest be awdieoa he aeehe tmetan ee cck cask eee mere 8. H. pseudogeniculata

—

*The leaf shape is based on leaves of mature, preferably flowering, shoots;

juvenile Homalomena are virtually impossible to identify to species by

leaf shape alone.

Taxonomic part

1. Homalomena atrox P.C.Boyce, S.Y.Wong & Fasihuddin, sp. nov.

Ab aliis specibus flumenicolis Borneensibus folliis et petiolis scintillans et

spadice sessile cum antheris terminalis sterilis differt. — Typus: Malaysia,

Sarawak, Sri Aman Division, Lubok Antu, Batang Ai, Nanga Sumpa,

Rumah Gumbang, Sungai Delok, 01° 12’ 16.2”; 112° 03’ 26.0”, 24 May 2008,

P.C.Boyce et al. AR-2375 (holo, SAR, + spirit). Fig. 1.

Diminutive, almost odourless evergreen microscopically pubescent herbs

to ca 10 cm tall. Stem pleionanthic, erect to decumbent, ca 5 mm thick, dull

red, internodes to ca 3 mm long. Leaves up to ca 6 together; petiole very

shallowly channelled adaxially, rounded abaxially, sub-erect, up to 5 cm long,

with a weak articulation ca 1/3 way along, microscopically asperate, matte dull

reddish, drying dark brown; petiolar sheath to ca 3 cm long, over 2 of petiole

length, sheath long-persistent, lower clasping part with undulate-bullate

margins; lamina oblong-lanceolate to oblanceolate, 8-12 cm long x 2-3 cm

wide, thinly and rather softly, glossy mid-green adaxially (fresh), drying pale

olive green, abaxially pale and slightly refractive (fresh), drying pale brown,

base cuneate-decurrent, posterior lobes absent, lamina tip acute, thence

apiculate for ca 3.5 mm; midrib prominently rounded-raised abaxially, (fresh

and dry), same colour as lamina, adaxially flush with or very slightly sunken

The Homalomena of Nanga Sumpa 273

into lamina (fresh and dry), ca 1.5 mm wide, with ca 5 primary lateral veins on

each side, diverging at 20°- 30° from the midrib, adaxially impressed (fresh),

flush with lamina when dry, abaxially slightly raised (fresh and dry), joining a

near the margin; interprimary veins ca %4 width of the primary lateral veins,

alternating irregularly with primaries, posterior lobes each with 3-4 primary

lateral veins; secondary venation rather obscure, striate; tertiary venation not

visible, all veins running into a thickened intermarginal vein, this particularly

conspicuous at the leaf tip, drying paler than the lamina. Inflorescences 1-2

together, erect at anthesis, later declinate; peduncle to ca 3 cm long x ca 1 mm

diam., matte dull red. Spathe 1-1.2 cm long, not constricted, matte dull red

externally, shiny greenish red internally, with a terminal short mucro (ca 0.5

mm long), spathe opening at anthesis by inflation and thence a broad slit;

lower spathe and spathe limb not obviously differentiated. Spadix 1-1.2 cm

long x 0.30-0.35 cm diam., the uppermost portion sterile by the presence

of 2-4 sterile male flowers, sessile; female flower zone 0.5 mm long; pistils

somewhat distant, broadly ovoid, ca 1 mm tall x 0.8-0.9 mm diam. greenish

whitish, stigma sessile to subsessile, disk-like, 0.3-0.4 mm diam.; each pistil

with a single staminode situated on ventral side of the flower relative to

the base of the spadix; interpistillar staminodes irregularly clavate, ca 0.2

mm long, white; suprapistillar interstice zone very short, less than 0.1 mm,

naked, pale pink; male flower zone 10-12 mm long, apex acute; male flowers

broadly dumbbell shaped, each consisting of two stamens, stamens rounded,

ca 0.5 mm tall, 0.6-0.8 mm long x ca 0.4 mm wide, ivory-white, anther thecae

opening by a broad terminal slit. Infructescence declinate, dull red, peduncle

matte dark red. Fruits and seeds not observed.

Distribution: Borneo. Sarawak, Sri Aman Division — endemic to riverine shales

of the Batang Ai drainage system.

Habitat: Riverine forest on red soils overlying shale, plants exclusively on

exposed shales, 80-150 m asl.

Notes: Homalomena atrox is immediately recognizable by the thinly soft-

leathery, oblong-lanceolate to oblanceolate laminae with decurrent bases,

and sessile spadix with the terminal portion sterile by the presence of 2-4

sterile flowers. In overall habit is vaguely reminiscent of H. paucinervia Rid.

(sandstones, Matang area) but is readily distinguished by the much softer

textured leaves not slightly glaucous abaxially.

Homalomena atrox is often sympatric with riverside populations of

H. humilis sensu lat., but always grows nearer the water. It appears to be a

facultative rheophyte, this view is further supported by the stenophyllous

habit.

274 Gard. Bull. Singapore 61 (2) 2010

Uses: No recorded uses among the Iban communities at Batang Ai.

Iban name: None recorded.

Etymology: The specific epithet is from the Latin atrox — terrible — in

gently chiding allusion to the taxonomic complexity of the Chamaecladon

supergroup to which this remarkably distinctive new species belongs.

Other specimens seen: SARAWAK. Sri Aman Division: Lubok Antu, Batang

Ai, Nanga Sumpa, Rumah Gumbang, Sungai Delok, 01° 11° 40.8”; 112° 04

04.2”, 28 Jul 2004, P.C.Boyce et al. AR-1115 (SAR); Lubok Antu, Batang

Ai, Nanga Sumpa, Sungai Pedali, 01° 11’ 58.9”; 112° 03’ 27.0”, 25 May 2008,

P.C.Boyce et al. AR-2389 (SAR).

2. Homalomena borneensis Ridl., J. Straits Branch Roy. Asiat. Soc. 44: 173

Sree? — Typus: Malaysia, Sarawak, Kuching, Ridley s.n. (holo, SING). Fig.

ae Homalomena ovata Engl. non (Schott) Hook.f., Fl. Brit. Ind. 6: 536

(1893), nom. illeg., based on Malaysia, Sarawak, Kuching Division, Matang,

Beccari p.b. 1780.

Medium, rather robust strongly aromatic (mango-resin) evergreen glabrous

herbs to ca 85 cm tall. Stem pleionanthic, erect, up to ca 3 cm thick, pale

green, internodes to ca 1.5 cm long. Leaves up to ca 15 together, ca 5 per

module; petiole weakly D-shaped in cross section, erect, 15-45 cm long,

obscurely pulvinate ca 2/; along length, pulvinus ca 2 cm long; petiole bases

clasping, pale green, sometimes suffused reddish for the basal 4, matte,

drying medium brown, petiolar sheath ca 9-20 cm long, ca 1/3 of petiole length,

equal, decurrent at apex, margin erect when fresh, margins membranous,

pale green, persistent, occasionally slightly scarious on the oldest leaves,

these oldest sheathes often red; lamina ovate to very weakly ovato-sagittate,

10-22 cm long x 9-15 cm wide, thinly leathery, pale green adaxially, drying

medium olive-brown, abaxially slightly glaucous(fresh), drying glaucous

pale brown, base subtruncate to very shallowly cordate, posterior lobes

absent or very short, straight, if present the n ovato-triangular, 0.5-3 cm

long, tip acute, acuminate for ca 1 cm thence tubular-apiculate for ca 2-3

mm; midrib raised abaxially (fresh and dry), glaucous green when fresh,

adaxially sunken slightly into lamina, ca 3 mm wide, with ca 7-10 primary

lateral veins on each side, diverging at 30°-70° from the midrib, adaxially

impressed (fresh), flush with lamina when dry, abaxially slightly raised

(fresh and dry), distal-most veins curved slightly towards the apex when

near the margin, interprimary veins ca 4 width of the primary lateral veins,

The Homalomena of Nanga Sumpa ATS

alternating with primaries; secondary venation very obscure, striate; tertiary

venation not visible, all veins running into a slightly thickened intermarginal

vein, this often reddish-flushed. Inflorescences up to 6 together, erect and

smelling of anise at anthesis, thence sinuous-declinate, each subtended by

prophyll, to ca 10 cm long; peduncle to ca 15 cm long x ca 3.5 mm diam., pale

green. Spathe 9-11 cm long, tightly furled prior to anthesis, lower spathe

inflating at female anthesis, spathe limb loosening at female anthesis, thence

inflating and then opening wide, entire spathe white at anthesis; lower

spathe ovoid-ellipsoid, equalling spathe limb, 5-6 cm long, constricted at

the junction of the spathe limb, the constriction coinciding with junction

of the male and female flower zones; spathe limb ovato-triangular, ca 5-6

cm long x ca. 3.5 cm wide (at male anthesis), spathe limb margins recurving

at male anthesis, apex mucronate to ca 5 mm long. Spadix subequalling

the spathe, ca 9-10.5 cm long, stipitate; stipe ca 4 mm long x 3 mm diam.,

cylindrical, and inserted obliquely on peduncle, white, female flower zone

ca 4.5 cm long x ca 1 cm wide, ca % length of spadix, stoutly fusiform;

pistils ca 1.3 mm x 0.75 mm, densely arranged, globose-cylindrical, white;

stigma globose-capitate, slightly exceeding the ovary and coherent with its

neighbours, staining deep brown in alcohol, each pistil associated with one

interpistillar staminode; interpistillar staminodes clavate on a stout stipe ca

0.4 mm diameter, equalling or slightly overtopping the associated female

flower waxy white; suprapistillar interstice zone + absent: male flower zone

more-or-less contiguous with female flower zone but markedly narrower,

to ca 5 cm long, ca 2 length of spadix, separated from female zone by the

prominently truncate upper part of the latter, with the lowest portion of

male flower zone comprised of sterile male flowers intergrading into a single

row of staminodes intermixed, this zone separated from the remainder of

the male flower zone by a distinct constriction; male flowers ca 3 mm x 2mm

trapezoid, comprising 3 —5 truncate stamens, each overtopped by a large, flat

connective. Infructescence declinate, spathe entirely persistent, pale green,

somewhat glaucous, ripening mid-green, less often deep red, peduncle matte

medium green. Fruits and seeds not observed.

Distribution: Borneo. Sarawak, widespread west of the Rejang valley. This

species is recorded from scattered localities in Kalimantan Barat, Kalimantan

Tengah & west Kalimantan Timur.

Habitat: Ever moist to slightly seasonal evergreen forest on sandstone or

shale-derived clays with deep leaf litter. 30-600 m asl.

Notes: The leaf laminae almost always lacking posterior lobes, and pale

green, rather matte adaxially and moderately to conspicuously glaucous

276 Gard. Bull. Singapore 61 (2) 2010

abaxially are diagnostic throughout Sarawak. The spathe limb shorter than

the lower part of the spathe is shared with H. borneensis, H. clandestina,

along with numerous undescribed species.

At Batang Ai H. borneensis may be confused with H. clandestina, with

which it grows intermixed. Homalomena borneensis may be distinguished

by the male flower zone being contiguous with the female flower zone (not

separated by a naked interstice); the female flower zone being far more

stoutly fusiform, with much more densely arranged pistils that are not

ascending, and with coherent stigmas.

Sterile plants are immediately recognizable by the leaf lamina abaxially

glaucous with the primary lateral venation much less pronounced.

Homalomena ovata Engl. non . (Schott) Hook.f., a nom. illeg., based

on a specimen from Matang, Kuching, is incontrovertibly the same as H.

borneensis.

Uses: Used as an antidote to snakebites, scorpion stings, etc. The bitten area

is kept in the smoke of the burning stems (reported as rhizomes) as long as

the patient can stand it. The leaves are used in post-partem petangas,a herbal

sauna. The rhizomes are also used to treat sick chickens by keeping them in

the smoke from burning ‘rhizomes’ of this and other species. Traditionally

burning rhizomes were also used to keep monkeys away from fields. The

aromatic rhizome is used in perfume mixtures. The ripe fruit is edible, with

a sourish taste.

[ban name: Kemuyang.

Etymology: “Borneo + ensis”, referring to origins on the Sunda shelf island of

Borneo.

Other specimens seen:SARAWAK. Kuching Division: Bau, Kampung Segong,

Sungai Adis, 11 Mar 2004, P.C.Boyce & Jeland ak Kisai AR-5.4 (SAR); Bau,

Kampung Segong, Ulu Sungai Adis, Sungai Bronand, 4 May 2004, Jeland ak

Kisai & Jipom ak Tisai AR-45 (SAR); Matang, Maha Mariamman Temple,

trail to Indian Temple, 2 Mar 2004, P.C.Boyce & Jeland ak Kisai AR-227

(SAR); Bau, Kampung Jugan, 26 Mar 2004, P.C.Boyce & Jeland ak Kisai

AR-275 (SAR); Bau, Rieng Opui, 28 May 2004, Jeland ak Kisai & Jipom

ak Tisai AR-406 (SAR); Bau, Kampung Segong, Ulu Sungai Adis, 9 May

2004, Jeland ak Kisai & Jipom ak Tisai AR-411.1 (SAR); Padawan, Subang,

7 Dec 2004, M.Gibernau AR-830 (SAR); Krokong, Kampung Tringgus, 01°

15’ 40.2”; 110° 05’ 35.9”, 19 Feb: 2005, P:C Boyce eval, AiK-99F (SAR)? Bau,

Kampung Segong, 01° 32’ 00.9”; 110° 08’ 58.8”, 10 Aug-05, P.C.Boyce et al.

AR-1330 (SAR); Bau, Gunung Juita,01° 23’ 48.7”; 110° 08° 07.2”, 28 Oct 2005,

The Homalomena of Nanga Sumpa DT

P.C.Boyce et al. AR-1497 (SAR); Padawan, Kampung Tubih, Labak Payang,

25 Jan 2006, Simon Kutuh ak Paru AR-1674 (SAR); Matang, Kubah National

Park, Waterfall Trail, 01°35’ 40.2”; 110° 10°45.9”, 28 July 2007, P.C. Boyce etal.

AR-2/12 (SAR); Bau, Bongo Range, trail to Tegora Mine, 01° 19° 41.5”; 110°

09° 19.0”, 8 Sep 2007, P.C.Boyce et al. AR-2190 (SAR). Samarahan Division:

Serian, Gunung Ampungan, 01° 09’ 08.2”; 110° 37° 21.2”, 11 Nov 2003,

P.C.Boyce & Jeland ak Kisai AR-5.2 (SAR), AR-158 (SAR); Serian, Pichin,

Umon Murut, Tiab Belanting, 01° 08° 03.7”; 110° 27° 00.3”, 20 Aug 2004,

Simon Kutuh ak Paru AR-653 (SAR); AR-654 (SAR); Serian, Pichin, Bung

Biringan, 28 Oct 2004, Simon Kutuh ak Paru A R-736 (SAR); Serian, Pichin,

Gunung Kedadum, Sugun Kerang, 13 Nov 2004, Simon Kutuh ak Paru AR-

743 (SAR); Serian, Pichin, Utak Dibung, 8 Mar 05, Simon Kutuh ak Paru

AR-1022 (SAR); Serian, Pichin, Sungai Abu Sijo, 19 Mar 2005, Simon Kutuh

ak Paru AR-1110 (SAR); Serian, Pichin, Labak Singong, 8 Mar 2005, Simon

Kutuh ak Paru AR-1023 (SAR); Kuap, Pangkalan Kuap, 01° 26° 16.7”; 110°

22° 18.9”, 25 Oct 2005, P.C.Boyce et al. AR-1486 (SAR). Sri Aman Division:

Lubok Antu, Batang Ai, Nanga Sumpa, 01° 12’ 02.3”; 112° 03° 09.3”, 27

Jul 2004, P.C.Boyce et al. AR-544 (SAR); AR-546 (SAR); Serian, Pichin,

Kampung Bidak, 6 Mar 2006, Simon Kutuh ak Paru AR-1718 (SAR); Serian,

Pichin, Pakan Jilayau, 1 Apr 2006, Simon Kutuh ak Paru AR-1758 (SAR);

AR-1800 (SAR); Serian, Pichin, Sungai Jarak, 25 Apr 2006, Simon Kutuh

ak Paru AR-1805 (SAR). Sarikei: Pakan, Binatang, Ulu Binatang, 02° 02’

30.1”; 111° 43’ 34.4”, 6 Dec 2005, P.C.Boyce et al. AR-1571 (SAR); AR-1572

(SAR); Lubok Antu, Batang Ai, Nanga Sumpa, Rumah Gumbang, Sungai

Delok, 01° 12’ 16.2”; 112° 03’ 26.0”, 24 May 2008, P.C.Boyce et al. AR-2361

(SAR); Lubok Antu, Batang Ai, Nanga Sumpa, Sungai Pedali, 01° 11° 58.9”;

112° 03’ 27.0”, 25 May 2008, P.C.Boyce et al. AR-2386 (SAR).

3. Homalomena clandestina P.C.Boyce, S.Y.Wong & Fasihuddin, sp. nov.

Ab H. borneensis sed foliis foliis subtus non glaucis, venis lateralibus primariis

abaxialiter plus prominentis, inflorescentia femina ab inforescentia mascula

interstitio neutro separatus, pistillis sublaxis ascendens et stigmate non

confluentibus distinguenda. —Typus: Malaysia, Sarawak, Sri Aman Division,

Lubok Antu, Batang Ai, Nanga Sumpa, Sungai Pedali, 01° 11° 58.9”; 112° 03°

27.0”, 25 May 2008, P.C.Boyce et al. AR-2385 (holo, SAR, + spirit). Fig. 3.

Medium, strongly aromatic (bitter lime (limau-purut) peel) evergreen

glabrous herbs to ca 75 cm tall. Stem pleionanthic, erect to ascending, ca

2.5 cm thick, medium green, internodes to ca | cm long. Leaves up to ca 10

together, ca 3-5 per module; petiole terete, erect, 30-32 cm long, obscurely

pulvinate ca ?/; along length, pulvinus ca 2 cm long; petiole bases clasping,

petioles medium green, sometimes suffused reddish for the basal 4,

278 Gard. Bull. Singapore 61 (2) 2010

semi-glossy, drying dark brown; petiolar sheath ca 11-13 cm long, ca '/4 of

petiole length, equal, decurrent at apex, margin erect when fresh, margins

persistent; lamina narrowly weakly ovato-sagittate, 16-20 cm long x 9-11

cm wide, thinly leathery, glossy dark green adaxially (fresh), drying medium

olive-brown, abaxially pale green (fresh), drying medium brown, base

shallowly cordate, posterior lobes short, straight, ovato-triangular, 1-3 cm

long, apex acute, acuminate for ca 2 cm, thence tubular-apiculate for ca

2-3 mm; midrib raised abaxially (fresh and dry), green when fresh, drying

reddish brown, adaxially sunken slightly into lamina, ca 2 mm wide, with

ca 7-11 primary lateral veins on each side, diverging at 30°-80° from the

midrib, adaxially impressed (fresh), flush with lamina when dry, abaxially

slightly raised (fresh and dry), distal-most veins curved slightly towards the

apex when near the margin, interprimary veins ca ¥2 width of the primary

lateral veins, alternating irregularly with primaries; secondary venation very

obscure, striate; tertiary venation not visible, all veins running into a slightly

thickened intermarginal vein. Inflorescences up to 4 together, erect and

smelling of anise at anthesis, thence declinate, each subtended by prophyll,

to ca 14 cm long that soon degrades into fibres, peduncle to ca 18 cm long

x ca 2.5 mm diam., medium green. Spathe 9-10 cm long, tightly furled prior

to anthesis, lower spathe inflating at female anthesis, spathe limb loosening

at female anthesis, thence inflating and then opening wide, entire spathe

white at anthesis; lower spathe ovoid-ellipsoid, 5-6 cm long, moderately

constricted at the junction of the spathe limb, the constriction coinciding

with junction of the male and female flower zones; spathe limb narrowly

ovato-triangular, ca 5-6.5 cm long x ca. 2 cm wide (at male anthesis), limb

margins recurving at male anthesis, apex mucronate to ca 5 mm long.

Spadix subequalling the spathe, ca 8.5-9.5 cm long, stipitate; stipe ca 3 mm

long x 2 mm diam., cylindrical, and inserted obliquely on peduncle, white;

female flower zone ca 4 cm long x ca 1 cm wide, ca 2 length of spadix,

weakly fusiform; pistils ca 1.3 mm x 0.75 mm, somewhat laxly arranged,

globose-cylindric, directed upwards, white; stigma globose-capitate, slightly

exceeding the ovary, dirty grey in nature, staining paler brown than ovary in

alcohol, each pistil associated with one interpistillar staminode; interpistillar

staminodes clavate on a very slender stipe ca 0.4 mm diameter, equalling or

slightly overtopping the associated female flower, waxy white, suprapistillar

interstice zone to ca 2 mm long x 2 mm wide, lower part with scattered

pistils, these seemingly sterile (ovary slender cylindrical) and intermixed

with a single row, and a few scattered staminodes each comprising a single

or rarely 2 anthers, upper part of interstice naked; male flower zone to ca 5

cm long, ca 2/3 length of spadix, separated from naked part of interstice by

a prominently truncate lower part, with the lowest portion comprised of

fertile male flowers intergrading into a single row of staminodes intermixed

The Homalomena of Nanga Sumpa 279

with one to several pistils, lowermost flowers fertile male, this lower part

separated from the remainder by a weak constriction; male flowers ca 3 mm

x 2 mm trapezoid, comprising 3 — 5 truncate stamens, each overtopped by

a large, flat connective. Infructescence declinate, reddish, peduncle matte

dark green. Fruits and seeds not observed.

Distribution: Borneo. Sarawak, Sri Aman Division — endemic & known only

from the type.

Habitat: Terrestrial on shales in evermoist forest, 80 m asl.

Notes: Sterile plants resemble the widespread and abundant H. borneensis but

are readily identifiable by the abaxial venation much more prominent, and

the leaf laminae not abaxially glaucous. The spadix with a naked interstice,

longer (ca’/; spadix length) male flower zone truncate proximally with the

lowermost male flowers intermixed with pistillodes, and the weakly fusiform

female flower zone with somewhat laxly arranged pistils are diagnostic.

Uses: The sour-tasting leaves of Homalomena clandestina are utilized at as

flavouring, especially for fish.

Iban name: Subung tilan (Christensen, 2002: 283)

Etymology: From the Latin, cladestinus , hidden, in allusion to the fact that

the authors initially overlooked this species due to its overall similarity to

sympatric H. borneensis.

Other specimen seen: SARAWAK. Sri Aman Division: Lubok Antu, Batang

Ai, Nanga Sumpa, Sungai Pedali, 01° 11’ 58.9”; 112° 03’ 27.0”, 7 Apr 2005,

P.C.Boyce et al. AR-1160 (SAR).

4. Homalomena geniculata M.Hotta, Acta Phytotax. Geobot. 22: 155 (1967).

—Typus: Malaysia, Sarawak, Bintulu Division, vicinity of Minah camp, Hotta

15779 (holo, KYO!). Fig. 4.

Small, aromatic (mango/resin) evergreen glabrous solitary, rarely weakly

clustering herbs to ca 35 cm tall. Stem pleionanthic, initially erect but soon

spreading-decumbent with the active shoot tip weakly ascending, ca 1.5 cm

thick, medium green, internodes to ca 1 cm long. Leaves up to ca 20 together,

ca 3-5 per module but exact number obscured by the tightly distichous leaf

bases; petiole terete, spreading, 5-22 cm long, strongly pulvinate at the lamina

280 Gard. Bull. Singapore 61 (2) 2010

insertion, pulvinus ca 1-1.5 cm long; petiole bases distichous-clasping,

petioles medium green, sometimes suffused brownish-reddish basal, matte,

petiolar sheath ca 2-12 cm long, ca 2 of petiole length, decurrent at apex,

margin erect when fresh, persistent; lamina narrowly oblanceolate, 10-30 cm

long x 2.5-7 cm wide, chartaceous, matte medium green adaxially (fresh),

drying olive-brown, abaxially glaucous (fresh), drying glaucous-brownish,

base acute, tip acute, acuminate for ca 2 cm thence tubular-apiculate for

ca 3 mm; midrib raised abaxially (fresh and dry), adaxially sunken slightly

into lamina, ca 1.5 mm wide, with ca. 5-11 primary lateral veins on each side,

diverging at 30°-70° from the midrib, adaxially impressed (fresh), flush with

lamina when dry, abaxially slightly raised (fresh and dry), distal-most veins

curved slightly towards the apex when near the margin, interprimary veins

ca ¥2 width of the primary lateral veins, alternating regularly with primaries;

secondary venation obscure, striate; tertiary venation not visible, all veins

running into a slightly thickened intermarginal vein. Inflorescences solitary,

erect and smelling powerfully of anise at anthesis, declinate beneath the

leaf fan soon after anthesis and there remaining until fruit dispersal, thence

decomposing, subtended by a prophyll to ca 5 cm long, peduncle to7-12 cm

long x ca2mm diam.,medium green. Spathe 7-8.5 cm long, tightly furled prior

to anthesis, lower spathe inflating at female anthesis, spathe limb loosening

at female anthesis, thence inflating and then opening wide, entire spathe

white at anthesis; lower spathe ovoid-ellipsoid, 2.5-3 cm long, constricted at

the junction of the spathe limb, the constriction coinciding with junction of

the male and female flower zones; spathe limb narrowly ovato-triangular, ca

2-3 cm long x ca. 2 cm wide (at male anthesis), apex mucronate to ca 1.5 cm

long. Spadix subequalling the spathe, ca 6-7 cm long, stipitate; stipe ca 3 mm

long x 2 mm diam., cylindrical, inserted obliquely on peduncle, pale green;

female flower zone ca 1.5-2.5 cm long x ca 0.5 cm wide, ca '/3 length of spadix,

fusiform; pistils ca 1.3 mm x 0.75 mm, without interpistillar staminodes, very

densely arranged, cylindrical-globose; stigma discoid, slightly exceeding the

ovary and coherent with adjacent stigmas; suprapistillar interstice zone to

ca 4mm long x 1.5 mm wide, entirely clothed with staminodes comprising

2 anthers and these intergrading with fertile stamens; male flower zone to

ca 2.5-2.8 cm long, ca 2 length of spadix, the lowest portion comprised of

fertile male flowers intergrading into a few rows of staminodes intermixed;

male flowers ca 3 mm x 2 mm trapezoid, comprising 2-4 truncate stamens,

each overtopped by a large, flat connective. Infructescence erect above the

leaf fan, spathe entirely persistent, reddish at fruit maturity, peduncle matte

dark green. Fruits pale green, exposed by the circumscissile splitting and

tearing of the spathe; weakly fruit-smelling. Seeds ovoid, ca 1.5 mm long,

dark-brown, longitudinally sulcate.

The Homalomena of Nanga Sumpa 281

Distribution: Borneo. Sarawak, only known with certainty north and east of

the Batang Lupar (but see below).

Habitat: Terrestrial on soils overlying shales & sandstones, usually on step to

vertical banks with the plants projecting horizontally, 80-430 m asl.

Notes: The tightly distichous leaf arrangement, the habit on vertical or near

vertical earth banks and the petioles prominently pulvinate at junction

of the lamina make H. geniculata immediately recognizable. However, it

appears that there are at least two species involved. As defined here true

H. geniculata has leaf narrowly oblong leaf laminae that are pronouncedly

glaucous abaxially occurs east of the Batang Lupar (‘Lupar Divide’) whereas

plants with the leaves broadly oblongo-lanceolate and abaxially pale green

and occurring in NW Sarawak (specifically on the Matang Range, Kuching

Division) are referable to H. crassinervia Ridl. The status of geniculata-

like plants from Kuching (excluding Matang) and Samarahan Divisions

requires clarification. If these plants all prove to be conspecific, the name H.

crassinervia has priority.

Homalomena geniculata is not utilized at Batang Ai, and no record

exists for its utilization elsewhere in Sarawak.

Uses: None recorded.

Iban name: None recorded.

Etymology: Latin, geniculatus,‘kneed’ in allusion to the conspicuous swelling

(geniculum, or more accurately pulvinus) at the junction of the petiole and

lamina.

Other specimens seen: SARAWAK. Sri Aman Division: Pantu, Gunung

Gaharu, 01° 02’ 39.5”; 110° 53’ 18.3”, 8 Aug 2004, P.C.Boyce et al. AR-

644 (SAR); Lubok Antu, Batang Ai, Nanga Sumpa, Wong Ensalai, 01° 11

51.0”; 112° 03’ 39.9”, 26 May 2008, P.C.Boyce et al. AR-2424 (SAR). Kapit

Division: Nanga Gaat, Rejang Wood Concession, Sungai Piat, 01° 38° 09.1”;

113° 24 09.9”, 14 Oct. 2003, P.C.Boyce & Jeland ak Kisai AR-112.1 (SAR);

Nanga Gaat, Reyang Wood Concession, stream below Camp Gahada, 01°

41’ 49.4”; 113° 26’ 16.3”, 15 Oct 2003, P.C.Boyce & Jeland ak Kisai AR-

112.2 (SAR); Nanga Gaat, Rejang Wood Concession, km 65 road to Camp

Gahada, 01° 42’ 01.1”; 113° 31’ 14.8”, 12 May 2004, P.C.Boyce et al. AR-

319, AR-335 & AR-365 (SAR); Rejang Wood Concession, km 3.5 after heli-

logging camp on road to Camp Gahada, Sungai Bereng, 01° 45’ 36.0”; 113°

27 54.7”, 15 Dec 2004, P.C.Boyce et al. AR-903 (SAR); Belaga, Belaga road,

282 Gard. Bull. Singapore 61 (2) 2010

02° 43° 45.8”; 113° 45° 37.1”, 12 Oct 2005, P.C.Boyce et al. AR-1424 (SAR).

Bintulu Division: Bukit Satiam, 02° 58° 47.6”; 112° 56’ 37.5”, 10 Aug 2004,

P.C.Boyce & Jeland ak Kisai AR-579 (SAR); Bukit Satiam, 02° 59° 10.0”;

112° 55’ 42.8”, 14 Jul 2006, P.C.Boyce et al. AR-1905 & 15 Jul 2006, AR-1907

(SAR); Tatau, Bukit Sarang, Ulu Sungai Sarang, 13 Jul 2006, Skornickova

144 sub. AR-2028 (SAR). Limbang Division: Mulu, Long Lama, Mulu N.P.,

Sungai Empangau, tributary from Sungai Mendalam, 04° 13’ 41.6”; 114° 52’

50.5”, 30 Sep 2007, P.C.Boyce et al. AR-2250 (SAR).

5. Homalomena hanneae P.C.Boyce, S.Y.Wong & Fasihuddin, sp. nov.

Ab altis speciebus Homalomenae combination pistillis armeniacus et spadice

resinfer per anthesin mascula unica est. -Typus: Malaysia, Sarawak, Sri Aman

Division, Lubok Antu, Batang Ai, Nanga Sumpa, Rumah Gumbang, Sungai

Delok, 01° 12’ 16.2”; 112° 03’ 26.0”, 24 May 2008, P.C.Boyce et al. AR-2360

(holo, SAR,+ spirit). Fig. 5.

Medium, strongly aromatic (mango resin) evergreen glabrous herbs to ca

50 cm tall. Stem pleionanthic, erect to ascending, ca 3 cm thick, dull red,

- internodes to ca 1 cm long. Leaves up to ca 10 together, ca 3 per module;

petiole sub-terete, erect, 30-35 cm long, obscurely pulvinate ca 2/3 along

length, pulvinus ca 2 cm long; petiole bases clasping, petioles medium

green, suffused deep red or red-brown for the basal 4, semi-glossy, drying

dark brown, petiolar sheath ca 6-10 cm long, ca '/s of petiole length, equal,

decurrent at apex, margin erect when fresh, soon degrading, reddish to

scarlet; lamina narrowly to rather broadly ovato-sagittate, 16-22 cm long

x 9-16 cm wide, thinly leathery, glossy dark green adaxially (fresh), drying

medium brown, abaxially pale green (fresh), drying medium brown, base

hastato-cordate to cordate, posterior lobes short, straight, ovate to ovato-

triangular, 3-5 cm long, tip acute, acuminate for ca 2 cm thence tubular-

apiculate for ca 2-3 mm; midrib raised abaxially (fresh and dry), green when

fresh, drying reddish brown, adaxially sunken slightly into lamina, ca 3 mm

wide, with ca 7-11 primary lateral veins on each side, diverging at 30°-80°

from the midrib, adaxially impressed (fresh), flush with lamina when dry,

abaxially slightly raised (fresh and dry), distal-most veins curved slightly

towards the apex when near the margin, interprimary veins ca ¥2 width of

the primary lateral veins, alternating irregularly with primaries; secondary

venation obscure, striate; tertiary venation not visible, all veins running into

a slightly thickened intermarginal vein. Inflorescences up to 7 together,

with additional synflorescences often arising after the next foliage leaf,

thus plants with 20 or more inflorescences at various developmental stages

not uncommon, erect and smelling powerfully of anise at anthesis, thence

declinate, each subtended by prophyll, to ca 11 cm long; peduncle to ca 20

The Homalomena of Nanga Sumpa 283

cm long x ca 4 mm diam., pale creamy green. Spathe 10-14 cm long, tightly

furled prior to anthesis, lower spathe inflating at female anthesis, spathe limb

loosening at female anthesis, thence inflating and then opening wide, entire

spathe white at anthesis; closing post anthesis and then turning pink and

finally deep red; lower spathe ovoid-ellipsoid, about equalling the spathe

limb in length, 5-7 cm long, moderately constricted at the junction of the

spathe limb, the constriction coinciding with junction of the male and female

flower zones; spathe limb ovato-triangular, ca 5-7 cm long x ca 2.5 cm wide

(at male anthesis), limb margins very slightly recurving at male anthesis,

apex mucronate to ca 3 mm long. Spadix slightly exceeding the spathe, ca

8.5-9.5 cm long, briefly stipitate; stipe ca 3 mm long x 2 mm diam., gibbose-

cylindrical, and inserted slightly obliquely on peduncle, pale yellow, female

flower zone ca 5 cm long x ca 1.5 cm wide, slightly less than ca % length of

spadix, stoutly fusiform; pistils ca 1.8 mm x 0.9 mm, densely arranged, stoutly

cylindrical, very pale apricot, each associated with a large staminode; stigma

globose-capitate, slightly exceeding the ovary, very pale apricot, staining paler

brown than ovary in alcohol, each pistil associated with one interpistillar

staminode; interpistillar staminodes stoutly clavate on a slender stipe ca

0.4 mm diameter, equalling or slightly overtopping the associated female

flower, waxy white; suprapistillar interstice zone almost absent, at most to

ca 0.5 mm long, naked; male flower zone to ca 8 cm long, just exceeding ca

Y% length of spadix, lowest portion comprised of a few sterile male flowers,

producing large numerous amber-coloured resin droplets during late female

and at onset of male anthesis; male flowers moderately well defined from

one another, ca 3 mm x 2 mm trapezoid, comprising 3-5 truncate stamens,

each overtopped by a large, flat connective, white Infructescence declinate

to pendent, spathe entirely persistent, matte deep red, peduncle deep green

with darker green, short striae, matte. Fruits and seeds not observed.

Distribution: Borneo. Sarawak, Sri Aman Division — endemic to the Batang

Ai drainage system.

Habitat: Terrestrial in deep leaf litter on red soils over shales, often on step

banks above the river, 80-200 m asl.

Notes: Homalomena hanneae belongs to the large and taxonomically

complex Cyrtocladon supergroup. Species have usually cordate leaf laminae

and inflorescences with a distinct constriction between the lower spathe

and spathe limb. The numerous species separate on subtle combinations

of characters in the spathe and spadix that may or may not relate to

morphological differences, and vice versa, linked to often highly localized

distributions.

284 Gard. Bull. Singapore 61 (2) 2010

At Batang Ai H. hanneae is immediately identifiable by apricot-

coloured pistils and the presence of copious amber-coloured resin droplets

on the spadix at the onset of male anthesis. These droplets mix with the pollen

to form a sticky paste that adheres readily to visiting insects, notably beetles

of Scarabaeidae, and Chrysomelidae (Dercetina sp.) and is seemingly linked

to pollination strategy. Similar resin-pollen pastes have been observed in

Neotropical Philodendron (cf. Gottsberger and Amaral, 1984; Grayum, 1990;

Mayo, 1991).

Uses: Used as an antidote to snakebites, scorpion stings, etc. The bitten area

is kept in the smoke of the burning stems (reported as rhizomes) as long as

the patient can stand it. The leaves are used in post-partem petangas,a herbal

sauna. The rhizomes are also used to treat sick chickens by keeping them in

the smoke from burning ‘rhizomes’ of this and other species. Traditionally

burning rhizomes were also used to keep insect pest away from fields. The

aromatic rhizome is used in perfume mixtures. The ripe fruit is edible, with

a sourish taste.

Iban name: Kemuyang.

Etymology: Named for Hanne Christensen whose meticulous work on the

ethnobotany of the indigenous peoples of Sarawak drew attention to the

many new species yet to be described.

Other specimens seen: SARAWAK. Sri Aman Division: Lubok Antu,

Batang Ai, Nanga Sumpa, Rumah Gumbang, Sungai Delok, 01° 11° 58.9”;

112° 03’ 27.0”, 28 Jul 2004, P.C.Boyce et al. AR-560 (SAR); Lubok Antu,

Batang Ai, Nanga Sumpa, Wong Enseluai, 01° 11’ 00.9”; 112° 04 20.8”, 6

Apr 2005, P.C.Boyce et al. AR-1137 (SAR); Lubok Antu, Batang Ai, Nanga

Sumpa, Sungai Pedali, 01° 11’ 58.9”; 112° 03’ 27.0”, 25 May 2008, P.C.Boyce

et al. AR-2382 & AR-2406 (SAR, + spirit); Lubok Antu, Batang Ai, Nanga

Sumpa, Wong Ensalai, 01° 11° 51.0”; 112° 03’ 39.9”, 26 May 2008, P.C.Boyce

et al. AR-2408 (SAR, + spirit).

6. Homalomena humilis (Jack) Hook.f., Fl. Brit. India 6:533 (1893).—Typus:

Malaysia, Penang, Wallich (holo, K-W). Fig. 6.

Syn: Calla humilis Jack, Malayan Misc. 1(1): 22 (1820) — Chamaecladon

humile (Jack) Miq., Fl. Ned. Ind. 3: 213 (1856).

Very small to small, very weakly aromatic (resin-like) peel, clumping,

evergreen, microscopically puberulent herbs to ca 20cm tall, but usually much

less tall. Stem pleionanthic, erect to creeping, occasionally pendent under its

The Homalomena of Nanga Sumpa 285

own weight, ca 4 mm thick, medium green to deep maroon, internodes to ca

4 mm long. Leaves up to ca 15 together, ca 5-7 per module; petiole sub-terete

in cross section, erect to spreading, 5-15 cm long, obscurely pulvinate ca 2/3

along length, pulvinus ca 1 cm long: petiole bases weakly clasping, petioles

bright to dark green, brownish pink, or deep maroon, drying dark brown;

petiolar sheath ca 2-4 cm long, ca '/; of petiole length, equal, decurrent at

apex, margin erect when fresh, margins persistent; lamina ovato-sagittate,

2-16 cm long x 1-9 cm wide, softly thinly leathery, scintillating bright to dark

green, brownish pink or deep maroon adaxially (fresh), drying medium

olive-brown, abaxially paler (fresh), drying medium brown, base shallowly

cordate, posterior lobes short, straight, ovato-triangular 1-3 cm long, tip

acute, acuminate for ca 2 mm thence tubular-apiculate for ca 2-3 mm; midrib

raised abaxially (fresh and dry), same colour as lamina when fresh, drying

reddish brown, adaxially sunken slightly into lamina, ca 2 mm wide, with ca

4-7 primary lateral veins on each side, these diverging at 30°-80° from the

midrib, adaxially impressed (fresh), flush with lamina when dry, abaxially

slightly raised (fresh and dry), distal-most veins curved slightly towards the

apex when near the margin, interprimary veins ca 2 width of the primary

lateral veins, alternating irregularly with primaries; secondary venation very

obscure, striate; all veins running into a slightly thickened intermarginal

vein. Inflorescences up to 10 together, erect and smelling slightly sour at

anthesis, thence declinate, each subtended by small prophyll, to ca 2 cm long;

peduncle to ca 4 cm long x ca 1 mm diam., medium green, brownish pink,

or deep maroon. Spathe ca 1 cm long, tapering-cylindrical, tightly furled

prior to anthesis, gaping at female anthesis, then closing, /ower spathe not

differentiated from spathe limb, spathe narrowly ovato-triangular, ca lcm

long x ca.5 mm wide (at male anthesis), margins recurving at male anthesis,

apex mucronate to ca | mm long. Spadix exceeding the spathe at anthesis,

ca 1.2 cm long, stipitate; stipe ca 0.5-2 mm long x 0.5 mm diam., cylindrical,

and inserted obliquely on peduncle, pale green or pink, female flower zone

ca 4mm long x ca 1 cm wide, ca '/3 length of spadix, cylindrical; pistils ca 0.5

mm x 0.4 mm, loosely arranged, cylindrical-globose, pale green, brownish

pink m, or pink; stigma globose-capitate, less than ovary diam., each pistil

associated with one interpistillar staminode; interpistillar staminodes

clavate on a slender stipe ca 0.1 mm diameter, shorter the associated female

flower, suprapistillar interstice zone absent; male flower zone to ca 7 mm

long, ca 2/3 length of spadix, with the lowest portion comprised of a very

few sterile male flowers; male flowers ca 0.3 mm x | mm, dumbbell-shaped,

comprising 1-2 truncate stamens, overtopped by a larger rounded connective.

Infructescence declinate, spathe entirely persistent, reddish, peduncle matte

dark green, brownish pink, or deep maroon. Fruits oblong-ovoid, pale green

or tinged pinkish, ca 2 x 1 mm. Seeds not observed.

286 Gard. Bull. Singapore 61 (2) 2010

Distribution: As defined here distributed from southern Peninsular Thailand

throughout Peninsular Malaysia, and thence into Borneo, but see note

below.

Habitat: Lowland to upper hill evergreen perhumid to everwet forest on

a variety of substrates including limestone. Sometimes on bare limestone

under forest. 10-700 m asl.

Notes: The taxonomy of H. humilis is chaotic and thus the full distribution is not

clear. The small stature of the plants, their natural variability, and the small

and fleeting nature of the inflorescences, coupled with over-dependence

on dried specimens, has compounded an extremely unwieldy infraspecific

classification, without a key and with labyrinthine synonymy proposed by

Furtado [Gard. Bull. Straits Settlem. 10 (1939): 199], and including taxa at

the leveis of variety, subvariety and forma (some illegitimate). The synonymy

given above intentionally omits all synonyms previously cited for H. humilis

in, e.g., Hay et al. (1995), Govaerts & Frodin (2002), and Govaerts et al.

(2009). The humilis complex requires an exhaustive study throughout its

range.

Uses: No recorded uses among the Iban communities at Batang Aj.

Iban name: None recorded.

Etymology: Latin, humilis - low or low-growing, refers to the growth habit.

Other specimens seen: SARAWAK. Kuching Division: Bau, Gunung Juita,

01° 23’ 48.7”; 110° 08° 07.2”, C.C.Lee AR-4.1 (SAR); Bau, Krokong, Gua

Peri-peri, 01° 22’ 51.9”; 110° 07’ 09.3”, 29' Oct 2003; P.C.Boyce &Yelandak

Kisai AR-4.3(SAR); Lundu, Gunung Gading, 01° 42’; 109° 50°, 3 Mar 2004,

P.C.Boyce & Jeland ak Kisai AR-4.4 (SAR); Matang, Maha Mariamman

Temple trail to Indian Temple, 2 Mar 2004 P.C.Boyce & Jeland ak Kisai AR-

4.5 (SAR); Padawan, Bukit Manok, 01° 12’; 110° 18’, 18 Mar 2004, P.C. Boyce,

Jeland ak Kisai & A.Shafreena AR-4.6 (SAR); AR-4.7 (SAR); Bau, Gunung

Ti-Ton, 01° 23’; 110° 07’, 17 Mar 2004, P.C.Boyce & Jeland ak Kisai AR-

4.8 (SAR); Bau, Jambusan, Kampung Seromah, 2 Apr 2004, P.C.Boyce &

Jeland ak Kisai AR-4.9 (SAR); Bau, Gunung Singai, 18 Apr 2004, P.C. Boyce

& Jeland ak Kisai AR-4.10 (SAR); Serian, 13th mile Kuching - Serian road,

Bukit Quap, 01° 23’ 44.6”; 110° 21° 10.6”, 1 Sep 2004, P.C.Boyce & Jeland

ak Kisai AR-669 (SAR); Bau, Gunung Juita, 01° 23’ 48.7”; 110° 08° 07.2”, 18

Jun 2005, P.C.Boyce & Jeland ak Kisai A R-1238 (SAR); Padawan, Kampung

Sadir, 2 Feb 2006, P.C.Boyce & Simon Kutuh ak Paru AR-1698 (SAR); Bau,

The Homalomena of Nanga Sumpa 287

Kampung Stenggang, Sungai Topah, 25 Mar 2006, P.C.Boyce & Jipom ak

Tisai AR-1747 (SAR); AR-1748 (SAR); Lundu, Gunung Gadin, trail to

Waterfall Trail above Batu Apek, 01° 41° 48.2”; 109° 50° 20.5”, 14 Dec 2006,

P.C. Boyce et al. AR-2063 (SAR); Matang, Kubah National Park, Waterfall

Trail, 01°35’ 40.2”; 110° 10° 45.9”, 8 Jul 2007, P.C.Boyce et al. AR-2109

(SAR). Samarahan Division: Serian, Pichin, Enteng Sabung, 28 Feb 2005,

P.C.Boyce & Simon Kutuh ak Paru AR-1009 (SAR); AR-1010 (SAR); AR-

1011 (SAR); Serian, Pichin, Bubung, Sungai Bomo, 23 Apr 2005, P.C.Boyce

& Simon Kutuh ak Paru AR-1176 (SAR);5 May 2005, P.C.Boyce & Simon

Kutuh ak Paru AR-1186 (SAR); Serian, Pichin, Jalan Tahang Nodoi, 10 Jul

2005, P.C.Boyce & Simon Kutuh ak Paru AR-1263 (SAR) AR-1264 (SAR):

Serian, Pichin, Sungai Umpu Sabai, 12 Jul 2005, P.C.Boyce & Simon Kutuh

ak Paru AR-1265 (SAR); Serian, Pichin, Labu Sungai Tiyab, 26 Jul 2005,

P.C.Boyce & Simon Kutuh ak Paru AR-1294 (SAR); AR-1297 (SAR):

Serian, Pichin, Gunung KedadumBubung Darud, 6 Aug 2005, P.C.Boyce &

Simon Kutuh ak Paru AR-1312 (SAR); Serian, Pichin, Ampan Balah, Dawu,

6 Aug 2005, P.C.Boyce & Simon Kutuh ak Paru AR-1323 (SAR); Serian,

Pichin, Sungai Kerasik, 15 Sep 2005, P.C.Boyce & Simon Kutuh ak Paru

AR-1382 (SAR); Serian, Pichin, Darud Tirimun, Tiab Lawak, 20 Oct 2005,

P.C.Boyce & Simon Kutuh ak Paru AR-1493 (SAR). Sri Aman Divison:

Lubok Antu, Batang Ai, 01° 13’ 18.0”; 112° 03° 21.2”, 28 Jul 2004, P.C. Boyce

& Jipom ak Tisai AR-538 (SAR); Lubok Antu, Batang Ai, Nanga Sumpa,

Rumah Gumbang, Sungai Delok, 01° 11° 40.8”; 112° 04 04.2”, 28 Jul 2004,

P.C.Boyce et al. AR-563 (SAR); AR-564 (SAR); 24 May 2008, P.C.Boyce et

al. AR-2372 (SAR); Lubok Antu, Batang Ai, Nanga Sumpa, Sungai Pedali,

01° 11° 58.9”; 112° 03’ 27.0” 25 May 2008, P.C.Boyce et al. AR-2401(SAR).

Bintulu Division: Bukit Merairi, 02° 46° 26.9%; 113° 39° 19.8”, SJulia &

L.Chan-24 A R-1288 (SAR, SING); Bintulu, Bukit Satiam, 02° 59° 13.3”; 112°

55° 57.5”, 14 Jul 2006 P.C.Boyce et al. AR-1891 (SAR); Tatau, Bukit Sarang,

Grand Perfect field station, path to Gua Lubang, Batu Rusa L-45, 12 Jul

2006, J. Leong-Skorniékova 106 (AR-2008) (SAR, SING). Sarikei Division:

Pakan, Berasok, 01° 54’ 30.8”; 111° 38’ 59.1”, 06 Dec 2005, P.C.Boyce et al.

AR-1567(SAR). Miri Divison: Niah Suai, Niah N.P., Madu Trail, 03°48? 57.9”;

113° 46° 18.3”, 13 Jul 2006, P.C.Boyce et al. AR-1586 (SAR); Marudi, Long

Lama, Mulu N.P, trail from Clearwater Cave, 04° 03° 49.2”; 114° 49° 51.7”,

8 Aug 2006, P.C.Boyce et al. AR-1961 (SAR); Marudi, Long Lama, Mulu

N.P., trail to Long Lansat, Sungai Licat, 04° 00° 03.5”; 114° 48° 49.8", 9 Aug

2006, P.C.Boyce et al. AR-1978 (SAR). Limbang Division: Limbang, Nanga

Medamit, Mulu N.P., Sungai Empangau, tributary from Sungai Mendalam,

04° 13’ 41.6”; 114° 52’ 50.5”, 30 Sep 2007, P.C.Boyce et al. AR-2245 (SAR);

AR-2246 (SAR); Limbang, Nanga Medamit, Mulu N.P., Sungai Abun Kiri,

tributary from Sungai Terikan, from the back of Mentawai Research Station,

288 Gard. Bull. Singapore 61 (2) 2010

04° 14 07.4”; 114° 52’ 27.6”, 2 Oct 2007, P.C.Boyce et al. AR-2295 (SAR);

Limbang, Nanga Medamit, Mulu N.P., Melinau Gorge, 3 Oct 2007, P.C.Boyce

et al. AR-2315 (SAR). INDONESIA. Kalimantan Barat: Gunung Saran, 22

Nov 1999, C.C.Lee AR-4.2 (SAR).

7. Homalomena josefii P.C.Boyce & S.Y.Wong, Gard. Bull. Singapore 60(1):

13 (2008). — Typus: Malaysia, Sarawak, Bintulu Division: Bukit Satiam, 02°

59° 10.0”; 112° 55° 42.8”, 14 Jul 2006, P.C.Boyce et al. AR-1894 (holo, SAR,

+ spirit). Fig. 7.

Medium to robust strongly aromatic (ginger/resin) evergreen glabrous herbs

to ca 125 cm tall. Stem pleionanthic, erect to ascending, ca 5 cm thick, dark

red to green, internodes to ca 1 cm long. Leaves up to ca 15 together, ca 5-7

per module; petiole terete, erect to decumbent, 50-70 cm long, petiole bases

clasping, eventually falling to leave a conspicuous lunate scar, petioles matte

dark reddish to matte green, dark reddish forms with longitudinal ridges, green

forms always with pinkish red bases, drying dark brown; petiolar sheath ca 16-

21 cm long, ca % to '/; of petiole length, equal, sometimes unequal, broader

side rounded at apex, narrower side, weakly decurrent at apex, margin always

convolute when fresh, sometimes wide open with broader petiolar sheath,

sheath initially long-persistent with the marginal 1.5 mm soon drying paler,

eventually the whole sheath marcescent; lamina broadly ovato-sagittate, 25-45

cm long x 18-32 cm wide, thinly leathery, glossy dark to pale green adaxially

(fresh), drying pale olive green, abaxially matte medium green (fresh), drying

pale brown, base cordate, posterior lobes spreading, subtriangular 7-9 cm

long, tip obtuse, short-acuminate for ca 1 cm thence stiffly apiculate for ca 2-7

mm; midrib raised abaxially (fresh and dry), green when fresh, drying reddish

brown, adaxially flush with lamina, ca 1.5 mm wide, with 6-9 primary lateral

veins on each side, diverging at 50°-90° from the midrib, adaxially impressed

(fresh), flush with lamina when dry, abaxially slightly raised (fresh and dry),

curved sharply towards the apex when near the margin, interprimary veins

ca ¥2 width of the primary lateral veins, alternating irregularly with primaries,

posterior lobes each with 3-4 primary lateral veins; secondary venation rather

obscure, striate; tertiary venation not visible, all veins running into a thickened

intermarginal vein, often red when fresh, this particularly conspicuous at the

leaf tip and there drying paler than the lamina. Inflorescences 1-7 together,

erect at anthesis, later declinate, each subtended by prophyll, to ca 9 cm long,

followed by cataphyll, ca 2-8 cm long; peduncle to ca 15 cm long x ca 5 mm

diam., matte deep red. Spathe 6.5-15.3 cm long, tightly furled prior to anthesis,

lower spathe inflating at female anthesis, spathe limb loosening at female

anthesis, thence inflating and then opening wide, lower spathe pale green,

stained deep red at insertion of peduncle, flushed pink above, spathe limb

The Homalomena of Nanga Sumpa 289

white at anthesis, with apex and mucro shading to dark pink during anthesis;

lower spathe ovoid-ellipsoid, 2.5-6.5 cm long, moderately constricted at the

junction of the spathe limb, the constriction coinciding with the lower-most

fertile male flowers; spathe limb narrowly to broadly elliptic, ca 3.3-8.5 cm

long x 2.3 cm wide (at male anthesis), prominently keeled along the dorsal

midline, limb margins with the middle ca 2/3 reflexing slightly at male anthesis,

apex mucronate to ca 3.5 mm long. Spadix equalling the spathe, ca 6-15.3

cm long, elongate cylindrical-fusiform, narrowing in the lower male zone

coinciding with the constriction of the spathe and there intergrading with

staminodes, stipitate; stipe ca 3.5 mm long x 5 mm diam., shortly fusiform, with

a few staminodes present at the insertion of peduncle, similar to interpistillar

staminodes; female flower zone ca 2 cm long x ca 1.2 cm wide, ca '/3 length

of spadix, weakly fusiform; pistils ca 1.3 mm x 0.75 mm, densely arranged,

globose-cylindric; stigma as broad or slightly exceeding the ovary, raised

and weakly trisulcate, staining pale brown in alcohol, extending beyond the

ovary as a translucent collar, lowermost flowers each mostly associated with

two or more interpistillar staminodes, and seemingly sterile; interpistillar

staminodes truncate on a very slender stipe ca 0.3 mm diameter equalling

or slightly overtopping the associated female flower, a few pistillodes at the

base of interstice, these of similar size to pistils; suprapistillar interstice zone,

to ca 1cm long x 5 mm wide, sometimes wider than female zone; suprapistillar

staminodes each comprising a single anther; male flower zone to ca 4 cm long,

ca ¥2 length of spadix, separated from interstice by weakly constricted lower

part of male zone, clothed with fertile male flowers intergrading into a single

row of staminodes., distal- and proximal-most flowers apparently sterile; male

flowers ca 3 mm x 2 mm trapezoid, comprising 3-5 truncate stamens, each

overtopped by a large, flat connective, terminal-most flowers sterile and spadix

often topped with a vestigial naked appendix. Infructescence declinate, spathe

entirely persistent, pale green stained reddish pink, sometimes whole reddish

pink, peduncle matte dark red, matte. Fruits and seeds not observed.

Distribution: Borneo. In Sarawak endemic to Bintulu & Sr Aman

Divisions.

Habitat: Terrestrial on shales and seasonally inundated alluvium, 7-200 m

asl.

Notes: This is a new Division record, hitherto known only from Bintulu

Division. Homalomena josefii is rare at Batang Ai, with only single small

population so far located. It is by far the largest Homalomena species

present, often exceeding Im tall and with a stem up to 5 cm thick. The

spathe is keeled along the dorsal median line much as in the much smaller

290 Gard. Bull. Singapore 61 (2) 2010

H. symplocarpifolia described elsewhere in this paper, aside from the much

greater size and ovato-sagittate leaves, H. josefii also differs by the weakly

trisulcate stigma.

Uses: No recorded uses among the Iban communities at Batang Ai.

Iban name: None recorded.

Etymology: Named for Dr Josef Bogner (formerly of Botanischer Garten

Miinchen, Germany), one of the foremost experts on the aroids and perhaps

the only person to have seen all currently recognized aroid genera in the

field.

Other specimens seen: SARAWAK. Sri Aman Division: Lubok Antu, Batang

Ai, Nanga Sumpa, Rumah Gumbang, Sungai Delok, 01° 11° 58.9”; 112° 03°

27.0”, 28 July 2004, P.C.Boyce et al. AR-559 (SAR). Bintulu Divison: Bukit

Satiam, 02° 59’ 13.3”; 112° 55’ 57.5”, 14 Jul. 2006, P.C.Boyce et al. AR-1900

(SAR, + spirit); Bukit Satiam, 02° 59° 07.4”; 112° 55’ 47.0”, 15 Jul. 2006,

P.C.Boyce et al. AR-1908 (SAR, + spirit); Bintulu, road to Kampung Jepak,

ca 3.3 km after bridge over Batang Kemena en route to Sibu from Bintulu,

03° 08" 32.3”; 113° 03? 24.3”,,.15 Jul. 2006; P:@Boyee eral -AR9 TiS AiRee

spirit).

8. Homalomena pseudogeniculata P.C.Boyce & S.Y.Wong, Gard. Bull.

Singapore 60(1): 17 (2008). — Typus: Malaysia, Sarawak, Sarikei Division,

Ulu Sarikei, 01° 55’ 05.4”; 111° 29’ 35.8”, 7 Dec 2005, P.C.Boyce et al. AR-

1583 (holo, SAR). Fig. 8.

Medium to moderately robust, strongly aromatic (pine resin) evergreen

glabrous herbs to ca 50 cm tall. Stem pleionanthic, decumbent with apex

erect, frequently creeping for several metres and branching laterally while

still continuing a physionogmically unbranched primary axis, stem green,

internodes to ca | cm long. Leaves ca 8-12 together, ca 5 per module, each

module subtended by prophyll, up to ca 12 cm long; petiole terete, erect

to decumbent, up to ca 30 cm long, pulvinate apically, distance ca 3-13 cm

from lamina base, roots penetrating petiole bases, petioles matte green

when fresh, drying light brown; petiolar sheath to ca 14 cm long, ca 2 of

petiole length, sheath convolute, initially persistent, eventually the whole

sheath marcescent; lamina oblongo-lanceolate, sometimes oblongo-elliptic

to ovate, 18-33 cm long x 6-15 cm wide, rather thinly coriaceous, matte

mid-green adaxially , sometime the mid-rib paler (fresh), drying pale olive

green, abaxially matte pale green, sometimes with conspicuous pellucid

The Homalomena of Nanga Sumpa 291

dots when fresh, very occasionally red (fresh), drying pale brown, base

decurrent to truncate, tip obtuse, acuminate for ca 2 cm thence apiculate

to ca 9 mm; midrib raised abaxially (fresh and dry), drying straw-coloured,

adaxially flush with lamina, but slightly channelled towards the leaf base, ca

2.5 mm wide, with 6-9 primary lateral veins on each side, diverging at 45°-

55° from the midrib, adaxially impressed (fresh), flush with lamina when

dry, abaxially slightly raised (fresh and dry), curved towards the apex when

near the margin, interprimary veins ca 2 width of the primary lateral veins,

alternating irregularly with primaries, secondary venation rather obscure,

striate; tertiary venation not visible, all veins drying in intermittent raised

and flush strips especially when near to leaf margin, all veins running into

intermarginal vein. Inflorescences 1-5 together, erect, each subtended by

a prophyll up to ca 6.2 cm long: peduncle to ca 12-15 cm long x 1.5-1.6

cm diam., yellowish green. Spathe ca 10.6 cm long, tightly furled prior to

anthesis, loosening at female anthesis and yet further at male anthesis, lower

spathe yellowish green to white at maturity, spathe limb white prior to and at

anthesis, with apex and mucro pale green at anthesis; lower spathe narrowly

ellipsoid, ca 3.5 cm long, weakly constricted at the junction of the spathe limb,

the constriction coinciding with the lower-most fertile male flowers; spathe

limb narrowly lanceolate, ca 7 cm long, mucronate to ca 7 mm long. Spadix

shorter than the spathe, ca 8.3 cm long, stipitate; stipe ca 4.5 mm long, weakly

dorso-ventrally flattened, obliquely inserted on peduncle; female flower zone

ca 2.2 cm long x 6.5 mm wide, ca % length of spadix, weakly fusiform; pistils

densely arranged, ca 1.3 mm diam. x | mm tall, round-cylindrical, lower-

most pistils ca twice the size of fertile females, stigma exceeding the ovary,

coherent to adjacent stigma, slightly raised, staminodes absent at the base

of insertion, pistils with no associated interpistillar staminode, suprapistillar

interstice zone wider than the other zones; staminodes truncate, ca 1.5 mm

wide, slightly overtopping the pistils, male flower zone ca 4.8 cm long x

5.2 mm wide, ca % length of spadix, cylindrical, tapering to a sharp end,

narrowing in the lower part coinciding with the constriction of the spathe,

distal- and proximal-most flowers apparently sterile: male flowers ca 3 mm

x 1.6 mm trapezoid comprising (3) 4-7 truncate stamens overtopped by a

large connective, seemingly fertile to the tip. Infructescence declinate, spathe

entirely persistent, lower spathe dark red, limb green, peduncle green. Fruits

and seeds not observed.

Distribution: A Bornean endemic: Sarawak, Kuching, Sarikei, Sri Aman,

Kapit and Miri Divisions; and Brunei.

Habitat: Terrestrial mostly under full shade in deep soil on various substrates,

frequently on shales, rarely on granite. 62-600 m asl.

292 Gard. Bull. Singapore 61 (2) 2010

Notes: Homalomena pseudogeniculata is distinctive by its pulvinate petioles

and remarkable decumbent-creeping stem giving rise to short leafy side

shoots while maintaining a primary axis. Plants frequently occur growing

down steep forested slopes giving the impression of several individual plants

in a row but on investigation revealing a single creeping stem/rhizome with

numerous short lateral branches.

The other currently recognized species with geniculate (pulvinate)

petioles is the Sumateran endemic H. elegantula A.Hay, which differs from

H. pseudogeniculata, among other characters, by hapaxanthic shoots, overall

much smaller and less robust habit, and smaller (1 cm long) spathes with

only a very weak constriction between the limb and lower part.

Uses: No recorded uses among the Iban communities at Batang Ai.

Iban name: None recorded.

Etymology: The specific epithet is coined from the superficial similarity of

the leaves of this species to H. geniculata — hence pseudo — false.

Other specimens seen: SARAWAK. Kuching Division: Lundu, Gunung

Gading, trail to Waterfall, trail above Batu Apek, 01° 41° 48.2”; 109° 50’ 20.5”,

14 Dec 2006, P.C. Boyce et al. AR-2064 (SAR). Kapit Division: Nanga Gaat,

Rejang Wood Concession, stream below Camp Gahada, 01° 41° 49.4”; 113°

26° 16.3”, 15 Oct 2003, P.C.Boyce & Jeland ak Kisai AR-141.1 (SAR); Nanga

Gaat, Rejang Wood Concession, km 65 road to Camp Gahada, 01° 42’ 01.1”;

113° 31’ 14.8”, 12 May 2004, P.C.Boyce et al. AR-363 (SAR); Nanga Gaat,

Rejang Wood Concession, km 55 road to Camp Gahada, 01° 44 44.5”; 113°

28° 32.3”, 13 May 2004, P.C.Boyce et al. AR-385 (SAR); Nanga Gaat, Rejang

Wood Concession, trail to water catchment behind main camp, 01° 53’ 00.2”;

113° 26° 53.9”, 14 Dec 2004, P.C.Boyce et al. AR-882 (SAR); Nanga Gaat,

Rejang Wood Concession, km 65 road to Camp Gahada,01° 41° 59.7"; 113° 31

13.7”, 16 Dec 2004, P.C.Boyce et al. AR-907 (SAR); Pelagus, Pelagus Rapids,

Woodpecker Trail, 02° 11’ 15.1”; 113° 03’ 29.01”, 14 Mar. 2005, P.C.Boyce et

al. AR-1034 (SAR); Belaga, Belaga road, 02° 43’ 45.8”; 113° 45’ 37.1”, 12 Oct

2005, P.C.Boyce et al. AR-1455 (SAR); Belaga, Belaga road, 02° 42’ 55.9”;

113° 45’ 29.3”, 12 Oct 2005, P.C.Boyce et al. AR-1457 (SAR); Belaga, Belaga

road, 02° 42’ 55.9”; 113° 45’ 29.3”, 12 Oct 2005, P.C.Boyce et al. AR-1461

(SAR + spirit); Belaga, km 10 Bakun, Bintulu-Miri road junction, 02° 50°

51.7”; 114° 01’ 57.6”, 11 Oct 2005, P.C.Boyce et al. AR-1481 (SAR, + spirit).

Miri Division: Mulu, Long Lama, Mulu National Park, trail to Gunung Mulu

Summit, 04° 02’ 18.7”; 114° 49° 44.2”, 7 Aug 2006, P.C.Boyce et al. AR-1955

(SAR); Mulu National Park, trail to Long Lansat, Sungai Licat, 04° 00° 03.5”;

The Homalomena of Nanga Sumpa 293

114° 48° 49.8”, 9 Aug 2006, P.C.Boyce et al. AR-1985 (SAR); Miri, Marudi,

Sungai Silat Basin, Sungai Palutan, 02° 49.59’; 115° 00.30’, 25 Mar 2003, Lim

S.P. §.90424 (SAR). Limbang Division: Limbang, Nanga Medamit, Mulu

National Park, trail from Camp 5 to Kuala Terikan, 04° 12’ 58.0”; 114° 53’

20.1”, 29 Sep 2007, P.C.Boyce et al. AR-2237 (SAR). BRUNEI. Temburong

District: Sungai Temburong at Kuala Belalong, banks of Sungai Belalong. 4°

32’, 225° 9’, 24 Jun 1989, P.C. Boyce 431 (BRUN, K, L).

9. Homalomena sengkenyang P.C.Boyce, S.Y.Wong & Fasihuddin, sp.

nov.

Per pistillis viride clarus Homalomena sengkenyang mirabilis est. — Typus:

Malaysia. Sarawak, Sri Aman, Lubok Antu, Batang Ai, Nanga Sumpa,

Rumah Gumbang, 01° 12’ 16.2”; 112° 03’ 26.0”, 24 May 2008, P.C.Boyce et

al. AR-2362 (holo, SAR + spirit). Fig. 9.

Medium, strongly aromatic (juniperus/mango resin) evergreen glabrous

weakly clumping herbs to ca 50 cm tall. Stem pleionanthic, erect to

ascending, ca 2.5 cm thick, pale green, internodes to ca 2 cm long. Leaves

up to ca 7 together, ca 3-4 per module; petiole terete, erect, 25-30 cm long,

obscurely pulvinate ca 7/3 along length, pulvinus ca 2.5 cm long, petiole bases

clasping, petioles medium matte green, suffused reddish from the pulvinus

to the lamina insertion in moist plants, slightlyglossy, drying brown; petiolar

sheath ca 19-13 cm long, ca '/s of petiole length, slightly unequal, decurrent at

apex, margin erect when fresh, margins soon scarious, brown, remainder of

the sheathremaining medium green; lamina narrowly ovato-sagittate, 10-22

cm long x 5-13 cm wide, thinly leathery, glossy dark green adaxially (fresh)

with scattered darker glands (extrafloral nectaries?), drying medium olive-

brown, abaxially pale green (fresh), drying medium brown, base shallowly

and narrowly, cordate, posterior lobes short, straight, ovato-triangular 1-3

cm long, tip acute, acuminate for ca 2 cm thence tubular-apiculate for ca

3-5 mm; midrib raised abaxially (fresh and dry), green when fresh, drying

reddish brown, adaxially sunken slightly into lamina, ca 2 mm wide, with

ca 7-9 primary lateral veins on each side, diverging at 30°-80° from the

midrib, adaxially deeply impressed, giving a weakly quilted effect (fresh),

almost flush with lamina when dry, abaxially slightly raised (fresh and dry),

distal-most veins curved slightly towards the apex when near the margin,

interprimary veins ca ¥2 width of the primary lateral veins, alternating

irregularly with primaries; secondary venation very obscure, striate; tertiary

venation not visible, all veins running into a slightly thickened intermarginal

vein. Inflorescences up to 3 together, erect and smelling powerfully of anise

at anthesis, thence declinate, each subtended by prophyll, to ca 14 cm long

soon marcescent, peduncle to ca 13 cm long x ca 3 mm diam., matte pinkish

294 Gard. Bull. Singapore 61 (2) 2010

brown. Spathe 9-10 cm long, tightly furled prior to anthesis, lower spathe

inflating at female anthesis, spathe limb loosening at female anthesis, thence

inflating and then opening wide, entire spathe white at anthesis; lower spathe

ovoid-ellipsoid, 3-3.5 cm long, constricted at the junction of the spathe limb,

the constriction coinciding with junction of the male and female flower zones;

spathe limb narrowly ovato-triangular, ca 7-6.5 cm long x ca. 2.5 cm wide

(at male anthesis), apex mucronate to ca 4 mm long. Spadix subequalling

the spathe, ca 9-9.5 cm long, stipitate; stipe ca 3 mm long x 2 mm diam.,

weakly obconical, glossy pale green; female flower zone ca 3 cm long x ca

1 cm wide, ca '/ length of spadix, fusiform; pistils ca 1.5 mm x 0.75 mm,

densely arranged, globose, bright green; stigma capitate, slightly exceeding

the ovary, pale green, staining brown and darker than ovary in alcohol, each

pistil associated with one interpistillar staminode; interpistillar staminodes

clavate on a slender stipe ca 1 mm diameter, overtopping the associated

female flower, waxy white, suprapistillar interstice zone to ca 1 cm long x 1|

cm wide, contiguous with and equalling diam. of female flower zone, covered

with large scattered staminodes, these intergrading to the lowermost fertile

flowers of the male flower zone; male flower zone to ca 5 cm long, ca '/3

length of spadix; male flowers ca 3 mm x 3 mm trapezoid, comprising 3-5

truncate stamens, each overtopped by a large, flat connective. Infructescence

declinate, spathe entirely persistent, pinkish to reddish, peduncle matte dark

green flushed and stained dull red and with paler conspicuous longitudinal

striate, these particularly prominent on the distal-most portion. Fruits and

seeds not observed.

Distribution: Borneo. Sarawak: Sri Aman Division — endemic to the Batang

Ai drainage system.

Habitat: Terrestrial in deep leaf litter on red soils over shales, often on step

banks above the river, 80-165 m asl.

Notes: A common species at Batang Ai with extensive colonies in the forest

and also in areas close to the longhouses, in which areas the plants are

maintained artificially.

Homalomena sengkenyang is immediately identifiable by the bright

green pistils, a character hitherto never reported for Homalonena.

Uses: This is by far the most important aroid utilized at Batang Ai, with the

status of sengkenyang bakung, plants of which are utilized as a protector

of rice plants. Plants utilized for sengkenyang are enormously important to

the Iban, as they are believed to protect the rice plants against malevolent

spirits and prevent the beneficial rice spirits from leaving the farm. Very few

The Homalomena of Nanga Sumpa 295

plants have the potential to be appointed to the status of sengkenyang.

Iban name: Sengkenyang bakung.

Etymology: The species epithet is derived from the Iban sengkenyang.

Other specimens seen: SARAWAK. Sri Aman Divison: Lubok Antu, Batang

Ai, Nanga Sumpa, 01° 12’ 02.3”; 112° 03’ 09.3”, 27 Jul 2004, P.C. Boyce et al.

AR-547 (SAR); Lubok Antu, Batang Ai, Nanga Sumpa, 01° 12’ 07.6”; 112°

02° 51.2”, 27 Jul 2004, P.C. Boyce et al. AR-548 (SAR); Lubok Antu, Batang

Ai, Nanga Sumpa, Sungai Pedali, 01° 11° 58.9”; 112° 03’ 27.0”, 25 May 2008,

P.C. Boyce et al. AR-2381, AR-2387, AR-2388 & AR-2403 (SAR,+ spirit).

10. Homalomena symplocarpiifolia P.C.Boyce, S.Y.Wong & Fasihuddin,

Sp. nov.

Species notabilis ab combinatio foliis oblongo et spathe exterior cum carina

longitidinalis dorsalis. Ab spathe inferior cum lamina spathae equalis H.

borneensis simillima est sed folis oblongo et inflorescntiae mascula et feminae

eadem diametero differt. —-Typus: Malaysia. Sarawak, Sri Aman, Lubok Antu,

Batang Ai, Nanga Sumpa, Wong Ensalai, 01° 11° 51.0”; 112° 03° 39.9”, 26

May 2008, P.C.Boyce et al. AR-2411 (holo, SAR, + spirit). Fig. 10.

Robust, very strongly aromatic (pinene) evergreen glabrous herbs to ca

50 cm tall. Stem pleionanthic, erect, ca 3 cm thick, glossy medium green,

internodes to ca 1 cm long. Leaves up to ca 5 together, ca 3-5 per module;

petiole weakly D-shaped in cross section, spreading, 20-30 cm long, stoutly

pulvinate ca ?/; along length, pulvinus ca 1.5 cm long; petiole bases loosely

clasping, medium green, semi-glossy, drying almost black; petiolar sheath

ca 8-10 cm long, less than '/s of petiole length, equal, weakly auriculate at

apex, margin strongly incurved, persistent, open; lamina oblong, 10-20 cm

long x 9-12 cm wide, leathery, glossy medium green adaxially (fresh), drying

medium olive-brown, abaxially pale (fresh), drying medium brown, base

very shallowly cordate, posterior lobes almost absent, , tip blunt, very briefly

tubular-apiculate for ca 2-3 mm; midrib raised abaxially (fresh and dry), green

when fresh, drying reddish brown, adaxially sunken slightly into lamina, ca 4

mm wide, with ca 8-10 primary lateral veins on each side, these diverging at

30°-80° from the midrib, adaxially impressed (fresh), abaxially slightly raised

(fresh and dry), distal-most veins curved slightly towards the apex when

near the margin, interprimary veins ca 2 width of the primary lateral veins,

alternating with primaries; secondary venation very obscure, striate; tertiary

venation not visible, all veins running into a slightly thickened intermarginal

296 Gard. Bull. Singapore 61 (2) 2010

vein. Inflorescences up to 4 together, erect and smelling of overripe banana/

anise at anthesis, thence declinate, each subtended by somewhat persistent

prophyll, to ca 10 cm long, peduncle to ca 10 cm long x ca 3 mm diam.,

glossy green. Spathe 7-9 cm long, tightly furled prior to anthesis, lower

spathe inflating at female anthesis, spathe limb loosening at female anthesis,

thence inflating and then opening moderately wide, entire spathe white at

anthesis; lower spathe narrowly ovoid-ellipsoid, 4-5 cm long, the dorsal side

often with a pronounced ridge, moderately constricted at the junction of

the spathe limb, the constriction coinciding with junction of the male and

female flower zones; spathe limb narrowly ovato-triangular, ca 3-4 cm long

x ca 2 cm wide (at male anthesis), limb margins incurved at male anthesis,

apex mucronate to ca 3 mm long. Spadix subequalling the spathe, ca 8.5-9

cm long, stipitate; stipe ca 5 mm long x 4 mm diam., cylindrical, and inserted

obliquely on peduncle and spadix, female flower zone ca 2.4 cm long x ca

1 cm wide, somewhat isodiametric, ca 1/3 length of spadix, weakly conical

cylindrical, basally oblique; pistils ca 1 mm x 0.75 mm, densely arranged,

globose-cylindrical; stigma globose-capitate, slightly exceeding the ovary,

greyish at anthesis, staining deep brown in alcohol, most pistils associated

- with one interpistillar staminode; interpistillar staminodes stoutly clavate on

a slender stipe ca 0.6 mm diameter, equalling the associated female flower;

suprapistillar interstice zone contiguous with female and male flower zones

and almost indistinguishable from male flower zone, to ca 5 mm long x | cm

wide, covered with scattered staminodes each comprising a single or rarely

3anthers; male flower zone to ca 5 cm long, ca 2/3 length of spadix, the lowest

portion comprised of sterile male flowers intergrading with staminodes; male

flowers ca 3 x 2 mm regularly trapezoid, comprising 3-4 truncate stamens,

each overtopped by a large, flat connective. Infructescence declinate, pale

pinkish red, peduncle glossy medium green. Fruits and seeds not observed.

Distribution: Borneo. Sarawak: Sri Aman Division — endemic to the Batang

Ai drainage system.

Habitat: Terrestrial in deep litter on red soils on seasonally inundated

alluvium and shale mud banks mostly in riverine forest. 100 m asl.

Notes: A highly distinctive species by virtue of the almost oblong leaves and

conspicuously longitudinally dorsally ridged lower spathe. The lower spathe

exceeding the spathe limb is a character shared with H. borneensis, from which

H. symplocarpifolia is readily distinguished by leaf shape.

Uses: Same as in Homalomena hanneae as an antidote to snakebites, scorpion

stings, etc.

The Homalomena of Nanga Sumpa 297

Iban name: Kemuyang.

Etymology: The trivial epithet is based on the lamina shape which in outline

and general, but not detailed, venation much resembles that of Symplocarpus

foetida Salisb. (Araceae: Orontoideae).

11. Homalomena vagans P.C.Boyce, Kew Bull. 49(4): 799 (1994). — Typus:

Brunei. Temburong: Batu Apoi Forest Reserve, Sungai Belalong, steep bank

near river, 4° 33’ N, 115° 9° E, 14 Mar 1992, Poulsen & Motte 273 (holo, AAU:

iso, BRUN, K). Fig. 11.

Relatively small, creeping, strongly aromatic (pinene) facultatively

rheophytic herbs, 20-45 cm tall. Stem pleionanthic, epigeal-creeping, 10-20

cm x 6-8 mm, lower part leafless with age, upper part obscured by densely

overlapping leaf bases, rooting along length and attaching strongly to

substrate, roots emerging through the splitting leaf sheath. Leaves many

together, ca 5 per module; petiole slender, subterete, canaliculate basally,

11-19 cm x 2-6 mm, base moderately expanded, pulvinate ca ?/;along length,

pulvinus marked, ca2 cm long: petiolar sheath 4.5-7 cm long, persistent,

open, the margins incurved; lamina 11-23.5 x 1.5-6 cm, pendent on the

petiole, lanceolate-elliptic, leathery, base decurrent, apex long acuminate,

lamina (living) dark green adaxially, yellow green abaxially, lamina (dry)

dull grey-green adaxially, pale brownish green abaxially: midrib prominent

abaxially, rounded, flush to somewhat impressed adaxially; primary lateral

venation slightly impressed adaxially, prominent abaxially, all other venation

+ obscure. Inflorescences several together, each sympodium subtended by

a persistent linear-lanceolate prophyll 2-4 cm x 2-5 mm, inflorescences

smelling sickly-sweet at anthesis; peduncle 7-8 cm x ca 1.5 mm: spathe ca

6 cm x 9 mm: lower spathe ca 2 cm , then constricted and distally tapering,

yellow-green, later pure white: spadix 3-5 cm x ca 5 mm (lower part) to 3

mm (upper part), stipitate: stipe 3 x 1.5 mm, terete, white; female flower

zone 1-2 x ca 5 mm, cylindric, usually with a few stamens interspersed

with the first few pistils, and with each pistil with an associated staminode;

pistils 1 x ca 0.75 mm, ovary obpyriform, pale green; stigma circular, ca half

ovary diam., ca 0.5 mm diam., stigma surface slightly papillate and wet at

anthesis: interpistillar staminodes clavate, truncate, slightly shorter than the

pistil, 0.75-1 x ca 0.5 mm; suprapistillar interstice zone narrower than female

flower zone, clothed with a few rows of sterile male flowers, white: male

flower zone 2-3 cm x ca long, narrowly cylindric; male flowers ca 0.75 x 0.75

mm, polygonal, densely arranged, white. Infructescence peduncle declinate

with the spathe erect at maturity, 8-11 cm x ca 2 mm. Fruit + globose, 1.5-3

298 Gard. Bull. Singapore 61 (2) 2010

x ca 1.5 mm, slightly truncate apically, pale to mid-green, stigma remaining

darker. Seed ellipsoid, ca 0.75 x 0.5 mm, mid-brown.

Distribution: Malaysia (Sarawak) & Brunei.

Habitat: Facultative rheophyte on sandstones and shales in lowland

evergreen perhumid or everwet forest, 60-450 m asl.

Notes: Homalomena vagans forms extensive colonies along river bank just

above the high-water mark and is readily identifiable by the long-creeping

rhizome-like stem and dull-green, rather coriaceous, lanceolate leaves

pendent on the petioles.

Boyce (1997) reported the species to be rheophytic but further

observations seem to show that this is in error, and that H. vagans is at most

only facultatively rheophytic.

There are several facultatively rheophytic, lanceolate-leaved riverside

Homlaomena in Sarawak, including H. /ancea Ridl. (Matang), and a number

of undescribed species.

Uses: No recorded uses among the Iban communities at Batang Ai.

[ban name: None recorded.

Etymology: Named from Latin vagans, wandering, in allusion to the long-

creeping rhizome-like stem, a feature that at the time of publication was

considered unusual in the genus.

Other specimens seen: SARAWAK. Kuching Division: Bau, Kampung Jugan,

Sungai Merah, 27 Apr 2004, P.C.Boyce & Jeland ak Kisai AR-33 (SAR);

Bako, trail to N.P., ca 3 hours by foot from park boundary, 13 Feb 2004,

P.C.Boyce & Jeland ak Kisai AR-219 (SAR); Bau, Kampung Apar, 17 Mar

2006, P.C.Boyce & Jipom ak Tisai AR-1735 (SAR). Sri Aman Division:

Lubok Antu, Batang Ai, Nanga Sumpa, Rumah Gumbang, Sungai Delok,

01° 11° 40.8”; 112° 04 04.2”, 28 Jul 2004, P.C.Boyce et al. AR-553 (SAR);

Pantu, Gunung Gaharu, 01° 02° 39.5”; 110° 53’ 18.3”, 8 Aug 2004, P.C.Boyce

et al. AR-645 (SAR); Lubok Antu, Batang Ai, Nanga Sumpa, Sungai Pedali,

01° 11’ 58.9”; 112° 03’ 27.0”, 25 May 2008, P.C.Boyce et al. AR-2390 (SAR).

Sarikei Division: Sarikei, Sungai Lepong, 01° 57° 12.9”; 111° 30° 34.9”, 8

Dec 2005, P.C.Boyce et al. AR-1602 (SAR); Maradong, Sungai Matob,

01° 52’ 06.1”; 111° 55’ 30.7”, 8: Dec 2005 P.C.Boyce et al. AR-1616 (SAR).

Limbang Division: Limbang, Nanga Medamit, Mulu N.P., Sungai Abun

Kanan, tributary from Sungai Terikan, from the back of Mentawai Research

The Homalomena of Nanga Swnpa 299

Station. 04° 14° 08.7”; 114° 52° 17.3”, 1 Oct 2007. P.C.Boyce et al. AR-2272

(SAR): Limbang. Nanga Medamit. Mulu N.P.. Sungai Abun Kiri, tributary

from Sungai Terikan, from the back of Mentawai Research Station. 04° 14

07.47: 114° 52° 27.6°,2 Oct 2007 P.C.Boyce et al. AR-2297 (SAR). BRUNEI.

Temburong: along the Sungai Temburong and Sungai Belalong, near their

junction, 4° 30°, 115° 10°, 2 Oct 1958, Jacobs 5615 (BRUN, G, L!, K!, US):

Batu Apoi Forest Reserve. Sungai Belalong. steep bank near river, 4° 33’.

115° 9°, 14 Mar 1992. Poulsen & Motte 47 (AAU! BRUN! K, spirit!).

12. Homalomena vivens P.C.Boyce. S.Y.Wong & Fasihuddin. sp. nov.

Ab allii Homalomenae foliis sagittato Batang Aiensis similis H. hanneae

praesertium, sed ab allii staminodiis inter pistilla nullis distinguida. Folii

laminorum produnde viride, nervis lateralibus primariis impressus et petioli

vagina long persistens uniga est. — Typus: Malaysia. Sarawak. Sri Aman.

Lubok Antu, Batang Ai, Nanga Sumpa. Sungai Pedali,01° 11° 58.97: 112° 03°

27.0°.25 May 2008, P.C.Boyce et al. AR-2402 (holo SAR + spirit). Fig. 12.

Medium, aromatic (camphor) evergreen glabrous herbs to ca 45 cm tall.

Stem pleionanthic, erect. ca 2 cm thick, dark green, internodes to ca 1.5 cm

long. Leaves up to ca 8 together. ca 3-5 per module: petioles dark green.

suffused reddish for the apical %, semi-glossy, drying dark brown, terete,

erect, 25-35 cm long, obscurely pulvinate ca 7/3 along length. pulvinus ca 2 cm

long, petiole bases clasping: petiolar sheath ca 12-15 cm long, ca 1/3 of petiole

length, equal. decurrent at apex. margin erect when fresh. margins persistent

and remaining bright green: lamina narrowly sagittate, 16-20 cm long x 9-11

cm wide, thinly and rather softly leathery. semi-glossy dark green adaxially

(fresh). drying olive-brown, abaxially paler green (fresh), drying medium

brown, base cordate-sagitatte, posterior lobes, straight, ovato-triangular 3-5

cm long. tip acute, acuminate for ca 1 cm thence tubular-apiculate for ca

24 mm: midrib raised abaxially (fresh and dry). green when fresh, drying

reddish brown, adaxially sunken slightly into lamina, ca 2 mm wide, with

ca 7-11 primary lateral veins on each side. diverging at 30°-80° from the

midrib, adaxially impressed (fresh), flush with lamina when dry. abaxially

slightly raised (fresh and dry), distal-most veins curved slightly towards the

apex when near the margin, interprimary veins ca % width of the primary

lateral veins, alternating irregularly with primaries: secondary venation very

obscure, striate: tertiary venation not visible, all veins running into a slightly

thickened intermarginal vein. Inflorescences up to 6 together. erect and

smelling powerfully of anise at anthesis. thence declinate. each subtended

by prophyll. to ca 5 cm long: peduncle to ca 16 cm long x ca 3.5 mm diam..,

semi-glossy pale pink to somewhat reddish. Spathe 10-13 cm long, tightly

furled prior to anthesis, lower spathe inflating at female anthesis, spathe

300 Gard. Bull. Singapore 61 (2) 2010

limb loosening at female anthesis, thence inflating and then opening wide,

entire spathe white at anthesis; lower spathe ovoid-ellipsoid, 3-5 cm long,

constricted at the junction of the spathe limb, the constriction coinciding with

junction of the male and female flower zones; spathe limb ovato-triangular,

ca 5-8.5 cm long x ca. 3 cm wide (at male anthesis), limb margins recurving at

male anthesis, apex mucronate to ca 5 mm long. Spadix exceeding the spathe

at anthesis, ca 10-14 cm long, stipitate; stipe ca 5 mm long x 2 mm diam.,

gibbose-cylindrical, glossy white; female flower zone ca 5 cm long x ca 1 cm

wide, just over 2/3length of spadix, weakly fusiform; pistils ca 1.3 x 0.75 mm,

very densely arranged, cylindric-globose; stigma capitate, slightly exceeding

the ovary, coherent with its neighbours, staining paler brown than ovary

in alcohol, interpistillar staminodes absent; suprapistillar interstice zone to

ca 6 mm long x 9 mm wide, completely covered with sterile male flowers,

lower part with coherent with female flower zone, upper part of interstice

contiguous with male flower zone; male flower zone to ca 7 cm long, over 2

length of spadix, separated interstice by a weak constriction; male flowers ca

3mm x 3 mm trapezoid, comprising 4-5 truncate stamens, each overtopped

by a large, very slightly convex connective, white. Infructescence declinate,

deep scarlet, peduncle matte dark green or slightly reddish. Fruits and seeds

not observed.

Distribution: Borneo. Sarawak: Sri Aman Division — endemic to the Batang

Ai drainage system.

Habitat: Terrestrial in deep litter over red on shale mud banks in riverine

forest. 80-200 m asl.

Notes: Homalomena vivens resembles several of the sagittate-leaved

Homalomena at Batang Aj, particularly H. hanneae, but is readily identifiable

from all by lacking interpistillar staminodes. The narrowly sagittate glossy

deep green leaf lamina, with the primary veins adaxially impressed, and the

petiolar sheaths bright green and long-persistent are diagnostic.

Etymology: From the Latin for living (vivens) to highlight the bright green,

long persistent petiolar sheathes that are a distinctive feature of this plant.

Other specimens seen: SARAWAK. Sri Aman Division: Lubok Antu, Batang

Ai, Nanga Sumpa, Rumah Gumbang, Sungai Delok, 01° 11° 58.9”; 112° 03°

27.0”, 28 Jul 2004, P.C.Boyce et al. AR-555 (SAR); Lubok Antu, Batang

Ai, Nanga Sumpa, Wong Enseluai, 01° 11’ 00.9”; 112° 04 20.8”, 6 Apr 2005,

P.C.Boyce et al. AR-1145 (SAR); Lubok Antu, Batang Ai, Nanga Sumpa,

Rumah Gumbang, Sungai Delok, 01° 12’ 16.2”; 112° 03’ 26.0”, 24 May 2008,

The Homalomena of Nanga Sumpa 301

P.C.Boyce et al. AR-2363 (SAR,+ spirit); Sri Aman, Lubok Antu, Batang

Ai, Nanga Sumpa, Sungai Pedali, 01° 11° 58.9”; 112° 03’ 27.0”, 25 May 2008,

P.C.Boyce et al. AR-2384 (SAR, + spirit).

Inadequately known species

13. Homalomena sp. A. Fig. 13A.

Medium, very strongly aromatic (mango peel resin) evergreen glabrous herbs

to ca 30 cm tall. Stem pleionanthic, erect to decumbent-ascending, ca 2 cm

thick, very dark green, internodes to ca 2.5 cm long. Leaves up to ca 8 together,

ca 3-4 per module; petiole terete, erect, 20-30 cm long, very obscurely pulvinate

ca 7/3 along length, pulvinus ca 2 cm long, petiole bases clasping; petioles dark

green, glossy, drying dark brown; petiolar sheath ca 10-15 cm long, ca % petiole

length, equal, decurrent at apex, margin erect when fresh, margins persistent,

later degrading into papery wings and then shedding; lamina spreading,

elliptic, 10-15 cm long x 6-11 cm wide, stiffly chartaceous, semi-glossy dark

green adaxially (fresh), drying olive-brown, abaxially much paler green (fresh),

drying medium brown, base broadly cuneate, tip acute, acuminate for ca 1 cm

thence tubular-apiculate for ca 2 mm; midrib raised strongly abaxially (fresh

and dry), green when fresh, adaxially sunken slightly into lamina, ca 3 mm

wide, with ca 9-14 primary lateral veins on each side, these diverging at 70°-

80° from the midrib, adaxially impressed (fresh), abaxially prominently raised

(fresh and dry), distal-most veins curved slightly towards the apex when near

the margin, interprimary veins almost the same width as primary lateral veins,

alternating with primaries and occasionally arching to meet them; secondary

venation semi-obscure, striate; tertiary venation not visible, all veins running

into a slightly thickened intermarginal vein. Inflorescences, infructescences,

fruits and seeds not observed.

Distribution: Borneo. Sarawak: Sri Aman Division, Batang Ai drainage.

Habitat: Terrestrial on shales in evermoist gallery forest, 100 m asl.

Notes: Although as yet found only sterile and as yet to flower in cultivation

the leaf morphology of this species is so distinctive we have no hesitation is

assigning this to an undescribed species.

Specimen seen. SARAWAK. Sri Aman Division: Lubok Antu, Batang Aji,

Lubok Antu, Batang Ai, Nanga Sumpa, Wong Ensulai, 01° 11° 51”; 112° 03°

39.9”, 26 May 2008, P.C.Boyce et al. AR-2414 (SAR).

302 Gard. Bull. Singapore 61 (2) 2010

14. Homalomena sp. B. Fig. 13B.

Small, strongly aromatic (citrus peel) evergreen glabrous herbs to ca 25 cm

tall. Stem pleionanthic, decumbent-ascending, ca 1.5 cm thick, medium green,

internodes to ca 2.5 cm long. Leaves up to ca 7together, ca 3-4 per module;

petioles pale purple, glossy, drying medium brown, terete, erect, 10-15 cm

long, obscurely pulvinate ca ?/; along length, pulvinus ca 1.5 cm long, petiole

bases weakly clasping; petiolar sheath ca 5-10 cm long, ca %2 petiole length,

equal, decurrent at apex, margin erect when fresh, margins persistent, open;

lamina spreading, oblong, 12-18 cm long x 7-110 cm wide, thinly leathery,

glossy deep green adaxially (fresh), drying olive-brown, abaxially glossy

maroon (fresh), drying medium brown, base broadly cuneate, tip obtuse,

acuminate for ca | cm thence tubular-apiculate for ca 2 mm; midrib raised

abaxially (fresh and dry), adaxially slightly sunken slightly into lamina, ca 3

mm wide, with ca 7-10 primary lateral veins on each side, these diverging at

50°-70° from the midrib, adaxially impressed (fresh), abaxially raised (fresh

and dry), distal-most veins curved slightly towards the apex when near the

margin, interprimary veins much less prominent than primary lateral veins,

- alternating with primaries; secondary venation semi-obscure, striate; tertiary

venation not visible, all veins running into a slightly thickened intermarginal

vein. Inflorescences, infructescences, fruits and seeds not observed.

Distribution: Borneo. Sarawak: Sri Aman Division, Batang Ai drainage.

Habitat: Terrestrial on sandstone-derived clays in evermoist gallery forest, 100

m asl.

Notes: This species resembles the widespread H. insignis N.E.Br. from west of

the Lupar, but differs by the leaf lamina adaxially glossy and abaxially shiny

deep maroon (vs. leaf lamina semi-glossy and abaxially glaucous).

Specimens seen. SARAWAK. Sri Aman Division: Lubok Antu, Batang Aj,

Nanga Sumpa, Rumah Gumbang, Sungai Delok, 01° 11° 40.8”; 112° 04

04.2”, 28 Jul 2004, P.C.Boyce et al. AR-562 (SAR); Lubok Antu, Batang Aj,

Nanga Sumpa, Wong Enseluai, 01° 11’ 51.0”; 112° 03° 39.9”, 26 May 2008,

P.C.Boyce et al. AR-2407 (SAR).

The Homalomena of Nanga Sumpa

Figure 1. Homalomena atrox P.C.Boyce, S.Y.Wong & Fasihuddin. A. Flowering shoot, note

the scintillating quality of the petioles and abaxial surface of the lamina; B. Inflorescence at

female anthesis with spathe partially artificially removed. All photos from P.C.Boyce et al.

AR-2375 (SAR).

Gard. Bull. Singapore 61 (2) 2010

Y ied

Beary,

Rite tel

: "

; B. Inflorescence just prior to

. Inflorescence at late female

anded spathe limb; the insects are

Figure 2. Homalomena borneensis Ridl. A. Flowering plant

female anthesis, note the entire spathe still tightly furled; ¢

anthesis, note the inflated lower spathe and fully e

Colocasiomyia (Diptera: Drosophilidae); D. Spadix at female anthesis (spathe artificially

removed), note the dense pistils and coherent stigmas. A-B photos from P.C.Boyce et al. AR-

544 (SAR); C-D photos from P.C.Boyce et al. AR-546 (SAR).

Aes

The Homalomena of Nanga Sumpa 305

Figure 3. Homalomena clandestina P.C.Boyce, S.Y.Wong & Fasihuddin. A & B. Inflorescence

at early male anthesis, note the damage to the male flower zone resulting from beetles

chewing the spadix; C. Spadix at female anthesis (spathe artificially removed), note naked

interstice between the male and female flower zones, the lax, ascending pistils, and separate

stigmas; D. Details of the female flower zone and lowermost part of the male flower zone; E.

Post-anthesis spadix, note the beetle-damaged male flower zone, and that the interpistillar

staminodes are now missing, having been eaten. All photos from P.C.Boyce et al. AR-2385

(SAR).

Gard. Bull. Singapore 61 (2) 2010

Figure 4. Homalomena geniculata M.Hotta. A. Plant in habitat; B. Inflorescence at female

anthesis; C. Spadix at female anthesis, spathe artificially removed; note that there are no

interpistillar staminodes; D. Detail of distichous leaf arrangement. All photos from P.C.Boyce

et al. AR-2424 (SAR).

The Homalomena of Nanga Sumpa 307

Figure 5. Homalomena hanneae P.C.Boyce, S.Y.Wong & Fasihuddin. A. Flowering plant; B.

Inflorescence at early anthesis, note amber-coloured resin droplets on the spadix; C. Detail of

post-anthesis female flower zone and lowermost part of the male flower zone; the black lines

are fly larvae; D. Spadix at female anthesis, spathe artificially opened; note the resin droplets

of the spadix and also the resin exuded from the cut spathe: E-F. Post antheis spadix showing

extensive beetle damage. All photos from P.C.Boyce et al. AR-2360 (SAR).

Gard. Bull. Singapore 61 (2) 2010

Figure 6. Homalomena humilis (Jack) Hook.f. A. Flowering plant, red expression; B. Plant

with inflorescences at various stages of development, the sequence running from right

(youngest) to left (oldest); C. Detail of inflorescence at female anthesis; note that spadix is

fertile to the tip. A photo from P.C.Boyce & Jipom ak Tisai AR-538 (SAR); B photo from

P.C.Boyce et al. AR-563 (SAR), and C photo from P.C.Boyce et al. AR-2401 (SAR).

The Homalomena of Nanga Sumpa

Figure 7. Homalomena josefii P.C.Boyce & S.Y.Wong. A. Plant; B. Petioles of green form

with inflorescences and infructescences; C. Emerging inflorescence bud, with declinate

infructescences behind; D. Inflorescence at late anthesis. All photos from P.C.Boyce et al.

AR-1894 (SAR).

Gard. Bull. Singapore 61 (2) 2010

Figure 8. Homalomena pseudogeniculata P.C.Boyce & S.Y.Wong. A. Plant in habitat; B.

Emerging inflorescence showing the distinctive mucro; C. Two infructescences. All photos

from P.C.Boyce et al. AR-1583 (SAR).

The Homalomena of Nanga Sumpa

Figure 9. Homalomena sengkenyang P.C.Boyce, S.Y.Wong & Fasihuddin. A. Plant in habitat;

B. Detail of petioles showing scarious petiolar sheath margins; C. Spz spathe artificially

removed to show the diagnostic green pistils; D. Developing infructescences. All photos from

P.C.Boyce etal. AR-2362 (SAR).

Gard. Bull. Singapore 61 (2) 2010

Figure 10. Homalomena symplocarpifolia P.C.Boyce, S.Y.Wong & Fasihuddin. A. Plant in

habitat, note the oblong leaves; B. Inflorescence at late male anthesis; C. Spadix, spathe

artificially removed to show the contiguous male and female flower zones; the lowermost

flowers of the male zone are sterile; D. Detail of female flower zone. All photos from

P.C.Boyce et al. AR-2411 (SAR).

The Homalomena of Nanga Sumpa

Figure 11. Homalomena vagans P.C.Boyce. A. Plant in habitat: B. Detail of the older part of

the creeping stems; C. Spadix, spathe artificially removed; D. Detail of female flower zone,

note the proportionately small stigmas: E. Infructescences with the diagnostic declinate

peduncle and erect spathe. All photos from P.C.Boyce et al. AR-2380 (SAR).

Gard. Bull. Singapore 61 (2) 2010

Figure 12. Homalomena vivens P.C.Boyce, S.Y.Wong & Fasihuddin. A. Flowering plant in

habitat; B-C. Inflorescence at male anthesis; C. Spadix, spathe artificially removed, note the

female flower zone lacks interpistillar staminodes. All photos from P.C.Boyce et al. AR-2402

(SAR).

The Homalomena of Nanga Sumpa

Figure 13. Homalomena sp. A & B. A. Plant of H. sp. A in habitat, note the pronounced

abaxial venation and somewhat ascending stem; B-C. Plants of H. sp. B in habitat. A photo

from P.C.Boyce et al. AR-2414 (SAR); B & C from P.C.Boyce et al. AR-2407

316 Gard. Bull. Singapore 61 (2) 2010

Acknowledgements

The collaboration and support of the Sarawak Forestry Department,

notably Dr Mohd. Shahbudin Sabki, Dr Azahari Bin Omar, L.C.J. Julaihi,

and Lucy Chong, and the Sarawak Biodiversity Centre, in particular Dr

Rita Manurong, & Dr Charlie Yeo Tiong Chia, is gratefully acknowledged.

This study is funded by MOSTI E-Science: 05-01-09-SF0006, under Sarawak

Forestry Department Research Permit No. NPW.907.4.2(1)-101 & Park

Permit No. 58/20076.

References

Hotta, M. 1967. Notes on Bornean plants, II]. Acta Phytotaxonomica

Geobotanica 22: 153 — 162.

Boyce, P.C. and Wong, S. Y. 2008. Studies on Homalomeneae (Araceae) of

Borneo I: Four new species and speculation on informal species group in

Sarawak. Gardens’ Bulletin Singapore 60(1): 1-29.

Boyce, P.C. 1994. New species of Araceae from Brunei. Kew Bulletin 49(4):

793 — 801.

Christensen, H. 2002. Ethnobotany of the Iban & the Kelabit. Forest

Department Sarawak, Malaysia, NEPCon Denmark & University of

Aarhus, Denmark. pp 384.

Furtado, C.X. 1939. Notes on some Indo-Malaysian Homalomena species.

Gardens Bulletin Straits Settlements 10: 183 — 238.

Gottsberger, G. and A. Amaral. 1984. Pollination strategies in Brazilian

Philodendron species. Berichte der Deutschen Botanischen Gesellschaft

97(3-4): 391-410.

Govaerts, R., Bogner, J., J. Boos, P.C.Boyce, B. Cosgriff,T. Croat, E. Goncalves,

M.H. Grayum, A. Hay, W.L.A. Hetterscheid, S. Ittenbach, E. Landolt, S.J.

Mayo, J. Murata, V.D. Nguyen, C.M. Sakuragui, Y. Singh, S. Thompson,

and G.-H. Zhu. 2009. World Checklist of Araceae. The Board of Trustees

of the Royal Botanic Gardens, Kew. Published on the Internet; http://

www.kew.org/wesp/ accessed 9 April 2009; 02:15 GMT.

The Homalomena of Nanga Sumpa S17

Govaerts, R. and D.G. Frodin, with J. Bogner, P.C. Boyce et al. 2002. World

checklist and bibliography of Araceae (and Acoraceae). x1, 560pp. Kew:

Royal Botanic Gardens.

Grayum, M.H. 1990. Evolution and phylogeny of the Araceae. Annals of

Missouri Botanical Garden 77: 628-697.

Hay, A., J. Bogner, P.-C. Boyce, W.L.A. Hetterscheid, N. Jacobsen and J.

Murata. 1995. Checklist and Botanical Bibliography of the Aroids of

Malesia, Australia, and the tropical western Pacific. Blumea, Suppl. 8:

1-210.

Hotta, M. 1967. Notes on Bornean plants, II]. Acta Phytotaxonomica

Geobotanica 22: 153-162.

Mayo, SJ. 1991. A revision of Philodendron subgenus Meconostigma

(Araceae). Kew Bulletin 46(4): 601-681.

Gardens’ Bulletin Singapore 61 (2): 319-326. 2010 319

Lost for a Century: Rediscovery of the Endemic Ridley’s

Jewel Orchid, Zeuxine exilis Ridl., on Christmas Island,

Indian Ocean

P.T. GREEN ', J. CLAUSSEN’* AND D.J.O’DOWD !

‘Department of Botany, La Trobe University, Bundoora, Victoria 3086 Australia

*Christmas Island National Park, Indian Ocean 6798 Australia

Corresponding author: p.green@latrobe.edu.au

Abstract

In a botanical expedition to Christmas Island in 1904, Sir Henry Ridley, the

first director of the Singapore Botanic Garden, discovered and thereafter

described an endemic ground orchid, Zeuxine exilis Ridl. Botanical

expeditions and surveys over the century since the original discovery failed

to relocate Z. exilis.We report here the rediscovery of Z. exilis in rainforest in

the western section of the island, and in Ridley’s honour, propose *Ridley’s

jewel orchid” as its common name. The distribution and conservation status

of Z. exilis remains to be determined. Despite a century of ecological insults

to this unique oceanic island, primarily through phosphate mining and the

introduction of invasive species, the rediscovery of this endemic orchid

renews hope that the imperiled biodiversity on this island is resilient and

can be conserved.

Introduction

More than a century ago, Sir Henry N. Ridley, founding director of the

Singapore Botanic Gardens, visited Christmas Island on two occasions,

first for just 10 hours in 1890 (Ridley, 1891) and again in 1904, for about

five weeks (Ridley, 1905a). This oceanic island, 360 km south of Java in the

northeastern Indian Ocean (10°29° S, 105°38° E) is covered in tall tropical

rainforest where it has not been cleared for phosphate mining. Ridley’s

expeditions yielded important collections that served as the basis for his

comprehensive treatment of the island’s flora (Ridley, 1905b). He described

several new species as endemic to the island, and one of these, Zeuxine exilis

Ridl. (Orchidaceae), has not been collected since, and was presumed extinct.

Here we report on the rediscovery of this endemic orchid, 105 years after its

first and only collection.

320 Gard. Bull. Singapore 61 (2) 2010

The rediscovery and identification of Zeuxine exilis

Specimens of an unidentified ground orchid were collected on 17 Nov 2009

from rainforest in the western area of the island. An initial identification as

Z. exilis was made by reference to Du Puy et al. (1993) and by comparison

with digital images of herbarium specimens from the Singapore Botanic

Gardens (Fig. 1A) and the Royal Botanic Gardens, Kew. An authoritative

determination was provided by Dr. M. Clements, orchid specialist at the

Centre for Plant Biodiversity Research, Australian National Herbarium

(CANB) Canberra, Australia. Given the undetermined conservation status

of Z. exilis, we do not report specific localities. Specimens are currently

being lodged with the Australian National Herbarium.

Although most individuals of Z. exilis observed in the field were

senescing (see below), we located several fresh plants (Fig. 1B-F). The

attributes observed are consistent with Ridley’s initial description (Ridley,

1905b, p. 236, presented below with SI units):

“Whole plant 30-45 cm tall, succulent, rhizome shortly creeping, roots

fleshy. Leaves lanceolate, acute, light green, 2.5-7.5 cm long, 1.2-2 cm

wide, glabrous, shortly petioled; sheaths papery, 1.2 cm long. Stem

white-hairy, peduncle 7.5-15 cm long, raceme many-flowered, 2.5-7.5

cm long. Bracts lanceolate, acuminate, hairy. Sepals reddish, hairy,

lanceolate, acute, 3 mm long. Petals thin, white, adnate to the upper

sepal. Lip base saccate with broad wings, then narrowed, limb broadly

bilobed, lobes broad oblong, divaricate edges crenulate, processes in

the base of the lip slender, subulate, curled, whole lip white with a cen-

tral yellow bar. Column short, anther lanceolate, beak up-curved, dull

red, pollinia elongate, pyriform, disc large, oblong. Rostellum lobes

linear, acuminate. No accessory processes. Capsule pubescent, elliptic,

6 mm long. Centre of the island, among ferns, not rare. Endemic.”

Given its all too brief history, Z. exilis has never enjoyed a common

name. We name this species as “Ridley’s jewel orchid” in honour of his

important contributions to botany on Christmas Island, and his discovery

and description of Z. exilis in particular.

Natural history of Zeuxine exilis

The genus Zeuxine comprises approximately 70 species, mostly distributed

in the tropics from Africa through southern and southeast Asia to Australia

and the Pacific (Ormerod et al., 2003). Species in the genus vary in their life

histories from annual to perennial. On Christmas Island, Ridley collected

Lost for a Century: Rediscovery of the Endemic Ridley’s Jewel Orchid 321

Z. exilis in full flower in Oct 1904, whereas we found mostly senescent plants

in Nov 2009. This suggests that Z. exilis appears aboveground for relatively

brief periods. It is unclear whether plants die off completely or just back

to rhizomes. Reproductive systems in Zeuxine vary from outcrossing in

Z. gracilis to apomixis in Z. strateumatica (Sun and Wong, 2001). Zeuxine

exilis may be apomictic given that all flowers appear to set fruit. We found

individuals in both heavily shaded rainforest understorey and in disturbed,

higher-light environments next to roads. Clearly, more information on the

life history, breeding system, and population biology of Z. exilis is needed.

Furthermore, Z. exilis is morphologically similar Z. gracilis and their

taxonomical differences need clarification.

Discussion

Rugged terrain and limited time forced the earliest naturalists visiting

Christmas Island, J. J. Lister in 1887 (Lister, 1888) and H. N. Ridley in

1890 (Ridley, 1891), to confine their activities to the area immediately

surrounding Flying Fish Cove, the only safe anchorage. The upper plateau

was not surveyed by a naturalist until C.W. Andrews stayed on the island

for 10 months in 1897/98 (Andrews, 1900). Although he collected several

orchid species, he did not find Z. exilis. Ridley made his second collecting

trip to the island in October 1904, this time for about five weeks. He made

several collecting forays in the northeast of the island (Ridley, 1905a), but

also made an expedition of several days duration in which he traversed the

island across the upper plateau from northeast to southwest (Fig. 2). It was

on this trip that Ridley collected a species of Zeuxine, and later formally

described it as an endemic species, Z. exilis,in a major work, The Botany of

Christmas Island (1905b).

Ridley’s specimens from that day were eventually sent to the

Singapore Botanic Gardens (SING 0046902, see Fig. 1A), the Royal Botanic

Gardens at Kew (K00079379), and the Natural History Museum, London

(BM00007629). Both the SING and BM specimens are labeled holotypes,

while the Kew specimens are labeled lectotypes. Apparently, there is a need

to lectotypify the species to resolve the designation of the type status. The

specimen collection labels on both the Singapore Botanic Gardens and

Kew Gardens’ sheets have the collection date and location simply stated as

“Murray Hill Track, Oct 1904”. In 1904, this track extended a considerable

distance, some 15 km, from Flying Fish Cove in the northeast, to Murray

Hill and beyond in the west.

We can now pin down the exact collection date and narrow considerably

the area in which Ridley discovered Zeuxine exilis. In addition to the

formal description of his collections (Ridley, 1905b), he gave an almost daily

322 Gard. Bull. Singapore 61 (2) 2010

description of his collecting activities (Ridley, 1905a), including the day he

discovered Z. exilis. He describes finding this plant on 18 Oct, 1904, close to

where he and the party spent the night of 18/19 October, southwest of Ross’

‘old encampment’. Ridley did not provide a map to show even approximately

where this location may have been, and since then, formal descriptions of

the orchids of Christmas Island (Wood, 1982; Du Puy et al., 1993) have been

forced to report Ridley’s original, imprecise locality. However, many years

after the 1904 expedition, Ridley’s fellow expeditioner, Dr Karl Hanitsch,

the first curator of the Raffles Museum, prepared handwritten notes for a

lecture he gave to the Ashmolean Natural History Society of Oxfordshire

on 15 May, 1923 (Hanitsch, 1923). This unpublished manuscript contains

a map of Christmas Island marked with the exploring party’s route and

campsites (Fig. 2). The coastline and contours in this map have obviously

been copied from the foldout map in Andrews’ (1900) monograph, and we

presume that Hanitsch drew on his original field notes from 1904 to plot the

route. Knowing the exact date that Ridley collected Z. exilis, and where the

party was on that day from Hanitsch’s map, we can now narrow the type

locality to somewhere in the vicinity of present-day Aldrich Hill, or perhaps

~ to its west.

Zeuxine exilis has not been collected since, even though the island

has been re-surveyed several times. Comprehensive collections were made

by D.A. Powell and H’ng Kim Chey in the 1980s and by B.A. Mitchell in

1984 (Mitchell, 1985). These collections have been lodged at Kew and the

Australian National Herbarium in Canberra, respectively. D.J. Du Puy vis-

ited the island twice in 1987 in preparation for his treatment of the flora of

Christmas Island (Du Puy, 1993). Here Du Puy listed Z. exilis as ‘appar-

ently extinct’ (p. 13), and ‘possibly endangered’ (p. 521). Previously, in an

unpublished report that focused on native and endemic plants with limited

distributions, he had listed Z. exilis as ‘extinct’ (Du Puy, 1988). Since then

two other attempts have been made to locate this orchid, both without suc-

cess (Holmes and Holmes, 2002; R. de Kok, Royal Botanic Gardens Kew,

pers. comm., 2009). Even though Z. exilis has been considered extinct by

botanists, it has no official status as such under the Environment Protection

and Biodiversity Conservation Act (1999) in Australia.

Ridley’s original account of Z. exilis described it as ‘not rare... Why

then has it taken more than a century for it to be seen again? We believe

that Ridley’s jewel orchid was not present in the western area of the island

where it was rediscovered until the last few years. We conducted intensive

seedling surveys across this area over 10 years from the late 1980s (Green et

al., 1997, 2008) and never encountered it. These surveys included the tran-

sition from dry to wet season (Oct-Dec) when Z. exilis should be evident.

Similarly, inappropriately timed surveys cannot explain the failure of Du

Lost for a Century: Rediscovery of the Endemic Ridley’s Jewel Orchid JLe

ke }

f |

‘¢)

] 3)

ens

, ‘|

- 4a ,

Phi]

» ~~

_ we! d

PX E- Fe

4 Lf

<— sil :

Figure 1. Zeuxine exilis Ridl. A. Holotype of Zeuxine exilis Ridl. (SING 0046902; scale bar

is 10 cm): B. Z. exilis in situ in rainforest, 17 Nov, 2009 (bar is 5 cm); C. Lanceolate leaf (bar

is 4 cm); D. Creeping rhizome (bar is 3 cm); E. Flowers (bar is 3 mm); F. A raceme showing

flowers, with developing and dehiscent capsules (bar is 3 cm).

324 Gard. Bull. Singapore 61 (2) 2010

Sperece Rint

Figure 2. Hanitsch’s original hand-drawn map of the 1904 expedition to Christmas Island

(see Hanitsch, 1923). Expedition route is a red dashed line; camping locations with dates

indicated by red crossed circles. Arrow indicates campsite on 18/19 Oct 1904 near the site

where Ridley collected Zeuxine exilis.

Puy (1988), Holmes and Holmes (2002), or the team from Kew in 2005 (R.

de Kok, pers. comm.) to relocate the orchid. We can only note that the reap-

pearance of the orchid coincides with major changes in rainforest under-

story structure as a result of widespread invasion by the yellow crazy ant

(Anoplolepis gracilipes) and its impacts (O’Dowd et al., 2003). Until the

distribution and abundance of Ridley’s jewel orchid is determined across

the island, the reasons for its unexpected but welcome reappearance, 105

years after Ridley’s discovery, remain enigmatic.

Lost for a Century: Rediscovery of the Endemic Ridley’s Jewel Orchid 325

Acknowledgements

We thank S. Brecknell and D. Rogers of Oxford University Museum for

the map from Hanitsch’s unpublished report, and the Singapore Botanic

Gardens for permission to use the image of holotype specimen of Z. exilis

kept at SING herbarium. R. Barnes provided one of the photographs in

Figure 1.

References

Andrews, C.W. 1900. A Monograph of Christmas Island. British Museum

(Natural History), London.

Du Puy, D.J. 1988. Mapping of Christmas Island native and endemic plants

with limited distributions. Unpublished report to the Australian National

Parks and Wildlife Service, Canberra, A.C.T., Australia.

Du Puy, D.J. 1993. Christmas Island, pp. 1-30. In: Flora of Australia, vol. 50,

Oceanic Islands. Australian Government Publishing Service, Canberra,

A.C.T., Australia.

Du Puy, D.J., D.L. Jones, and E. Edgar. 1993. Orchidaceae, pp. 516-528.

In: Flora of Australia, vol. 50, Oceanic Islands. Australian Government

Publishing Service, Canberra, A.C.T., Australia.

Environment Protection and Biodiversity Conservation Act. 1999. Australian

Government, Department of Environment, Water, Heritage and the Arts.

Canberra, A.C.T., Australia. http://www.environment.gov.au/epbc/about/

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by land crabs in rain forest on a remote oceanic island. Ecology 78: 2474-

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Green, P.T., D.J. O’Dowd, and P.S. Lake. 2008. Recruitment dynamics in

a rainforest seedling community: context-independent impact of a

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Hanitsch, K.R. 1923. A visit to Christmas Island. Unpublished manuscript.

Oxford University Museum archives, Oxford.

326 Gard. Bull. Singapore 61 (2) 2010

Holmes, J. and G. Holmes. 2002. Conservation status of the flora of Christmas

Island, Indian Ocean. Unpublished report to Parks Australia North,

Christmas Island, Indian Ocean.

Lister, J.J. 1888. On the natural history of Christmas Island, in the Indian

Ocean. Proceedings of the Zoological Society of London 1888: 512-564.

Mitchell, B. A. 1985. A vegetation survey of Christmas Island. Unpublished

report for the Australian National Parks and Wildlife Service. 40 pp.,

including Appendix II. List of collections made on Christmas Island

for the National Botanic Gardens Herbarium during June/July and

November/December 1984.

O’Dowd, D.J., P.T. Green, and PS. Lake. 2003. Invasional ‘meltdown’ on an

oceanic island. Ecology Letters 9: 812-817.

Ormerod, P., P.J. Cribb, and A.M. Pridgeon. 2003. Zeuxine, pp. 151-153. In:

A.M. Pridgeon, P.J. Cribb, M.W. Chase and F.N. Rasmussen (eds.). Genera

Orchidacearum: vol. 3 (Vanilloideae and Orchidoideae, part 2). Oxford

University Press, Oxford.

Ridley, H.N. 1891. A day at Christmas Island. Journal of the Straits Branch,

Royal Asiatic Society 23: 123-139.

Ridley, H.N. 1905a. An expedition to Christmas Island. Journal of the Straits

Branch, Royal Asiatic Society 45: 137-155.

Ridley, H.N. 1905b. The botany of Christmas Island. Journal of the Straits

Branch, Royal Asiatic Society 45: 156-271.

Sun, M. and K.C. Wong. 2001. Genetic structure of three orchid species

with contrasting breeding systems using RAPD and allozyme markers.

American Journal of Botany 88: 2180-2188.

Wood, J.J. 1982. The orchids of Christmas Island. The Orchadian 7: 142-146.

Gardens’ Bulletin Singapore 61 (2): 327-333. 2010 227

New Species of Hoya (Apocynaceae) from Brunei

and the Philippines

R. KLOPPENBURG ' AND S.V. SIAR *

' 6427 N. Fruit Ave., Fresno, CA 93711, USA

* Institute of Plant Breeding, College of Agriculture, University of the

Philippines Los Banos, College, Laguna, The Philippines.

Abstract

Two new species of Hoya collected from Brunei and the Philippines are

described and illustrated with coloured photographs of the flower details.

Introduction

Wild and cultivated species of Hoya in Borneo and the Philippines have been

documented and studied in recent years (Kloppenburg, 1991; Kloppenburg

and Wayman, 1993; Kloppenburg and Siar, 2009; Wennstrom and Stenman,

2008). Several species of Philippine and Bornean Hoya have also been used

in molecular study to shed light on their phylogenetic position in the genus

(Wanntorp et al. 2006). A recent examination of live plants grown in private

gardens and nurseries has revealed two new species of Hoya taken into

cultivation from the wild. They are described and illustrated below.

1. Hoya yapianum Kloppenburg, sp. nov. Section Otostemma

A Hoya sipitangensis Kloppenburg & Wiberg similes, sed hic sp. nova

sepalorae apex obtusam non acutam; pedecilli glabras non puberulas;

coronae apex interiora acutum non spathulatum; folia magnior; pollinaria ca

1/3 breves, differt. - Holotypus: BRUNEI. Tutong District, Kuala Belait, 21

Jan 2008, Wang Luan-Keng BN090122 (SING). Figs. 1 & 2.

A twining herb. Stem long slender, glabrous. Leaves lanceolate, variable in

size, 4.4-8.5 cm long and 1.5-3 cm at the widest part near the middle, apiculate,

nerves not conspicuous, pinnately veined, majour veins extending from

base to near apex. Petiole terete, 1-1.8 cm long. Inflorescence umbelliform,

consisting of 15-30 flowers. Pedicels variable in length, curved, glabrous with

crystalline surface, longest ca 1.5 cm and 0.10 cm in diameter, slight enlarged

at the calyx. Calyx small, whitish, and calyx lobes (sepals) do not reach the

corolla sinuses. Outer surface of calyx lobes granulose, thickened centrally

slightly, edges membranous, 0.08 x 0.08 cm, glabrous inside, apex obtuse,

328 Gard. Bull. Singapore 61 (2) 2010

Figure 1. Hoya yapianum Kloppenburg. A. Plant habit (x1); B & C. Inflorescences (B: x1.5;

©) x2).

narrow ligules present. Sepals very little overlapped at base. Corolla whitish,

thick, tightly revolute and outer surface villous-pubescent except for the

area under the corona. Outer surface of corolla glabrous, about 0.51 cm

long and 0.21 cm wide. The corona reddish in color, corona crown has the

center raised with outer lobes long and narrow, with very thin translucent

edges. The inner lobe is raised with an acute apex that does not cover the

membranous anthers. Central column pronounced, 0.07 cm long with a 0.10

cm opening, surface glabrous but finely crystalline. Skirt lobes rounded with

a central groove leading to the anther winged outer apex. The outer scale

membranous, dorsally keeled from apex to apex. The keel about mid way

has a hump as it extends to the outer apical area. Scale edges are sharp.

Stylar crown short, ovaries 2, cone shaped, glabrous, ca 0.11 cm long and the

base pair ca 0.06 cm broad. Anthers membranous, triangular; pollinia two

in pollinarium, oblong, 0.25 mm long and 0.12 mm wide; translator 0.10 mm

long and 0.04 mm wide. Caudicles clearly distinct, cone shaped. Retinaculum

0.06 mm long with extensions 0.04 mm. Fruit not seen.

This new species is similar to Hoya sipitangensis but different as the

apex of the sepals are obtuse not acute and the pedicels are glabrous not

puberulent; inner apex of the corona is acute not spathulate, the foliage is

Figure 2. Hoya yapianum Kloppenburg, details of inflorescence. A. Pedicel enlarged (x15),

showing also calyx and ovary of pistil; B. Top inside view of the calyx enlarged (x15). The

sepals are very little overlapped at the base; C. View of ventral side of a flower showing

corolla enlarged (x8). Corolla is tightly revolute and outer surface is villous-pubescent

except for the area under the corona; D. Outer surface of the corolla enlarged (8) with

one lobe extended, note this surface is glabrous; E. Dorsal surface of corolla (x8). There is a

slight pentagonal dip in the corolla center under the corona that is glabrous; F. View of the

o>)

Gard. Bull. Singapore 61 (2) 2010

flower with corona attached (x8). The corona is reddish in color and the corolla is off white.

Crown has center raised with outer lobes long and narrow, with very thin translucent edges.

The inner lobe is raised with an acute apex that does not cover the membranous anthers; G.

Corona enlarged (x16). The species of Otostemma Section has a very pronounced central

column, surface is glabrous but finely crystalline. Note the thin translucent outer coronal

lobe at the left extreme; H. Corona dorsal view enlarged (x16). The inner lobes are thin and

acute, not covering nor exceeding the inner anther apices; I. Individual scale side view (x16),

note the membranous outer scale lobe; J. Flower center showing the inner apex of two lobes

(x32). The retinaculum of a pollinarium is visible in the central lower left; K. Area between

two adjacent coronal outer lobes (x32), In the center is the anther wing not clearly in focus.

Note the thinness of the outer coronal lobes and the central keel. All coronal surfaces are

glabrous; L. Above two pollinaria the arrow on scale pointer is 0.1 mm. long (x60).

larger and the pollinia are about 1/3 shorter.

The new species was collected from Brunei by Ms.Wang Luan-Keng

and grown to flower by Mr. K. F. Yap on 22 January 2009 in Singapore.

Microphotographs and species data taken from materials sent by K. F. Yap.

Note: In size the pollinarium of the new species is close to Hoya revoluta

Wight, but the outer lobe of the corona of the new species is not divaricate

2. Hoya benitotanii Kloppenburg, sp nov. Section Acanthostemma

A Hoya gigantanganensis Kloppenburg similes, sed folia ovata-lanceolata

apiculatis, basi obtusis vs. linear-elliptic et basi acuminata; 9 x 4.cm vs. 11-16

x 2.4-6.2 cm; calycis lobus rotundatis 0.15 = 0.18 cm longis vs triangularis

et 0.21 x 0.17 cm longis; corolla aliquantump parvum differt. — Holotypus:

Kloppenburg 2010-6 (UC) hic designatus, ex hort. by Ted Green, Kaaawa,

Hawaii. Figs. 3 & 4.

A long twining herb. Stem slender, long, glabrous. Leaves ovate lanceolate

with an obtuse base, variable in size, 9 x 14 cm long, nerves 3-5, pinnately

branched, majour veins extending from base to near apex. Petiole terete.

Inflorescence umbelliform, consisting of 12-15 flowers. Pedicels pale violet

or rose in color, terete, curved, 2 cm long, 0.10 cm in diameter, extremely

glabrous and shiny. Calyx small, sepals very rounded, ciliate, finely reticulate-

veined, 0.15 cm long and 0.18 cm wide with nearly 1/2 overlap; ligules at base

white, small. Corolla revolute, outer surface glabrous, inside surface finely

pubescent with whitish shiny stellate hairs; lobes cut more than half way

and the apex acute. Corona glabrous, inner lobes spatulate, not long, leaving

much more of the anthers exposed. The bilobes with flattened upper surfaces,

which are longitudinally sulcate, extending beyond outer lobe apices, and the

outer lobe apex drops off sharply as in the other species of Hoya. Anthers

5, membranous, triangular, covering staminal crown, and exceeding inner

lobes. Below the lobes are channeled, but only a short distance toward the

ane.

New Species of Hoya from Brunei and the Philippines S51

é S} y t

Figure 3. Hoya benitotanii Kloppenburg. A. Plant habit (0.5): B & C. Inflorescence (B: «1.5:

Cax225):

center where there is a prominent thickened column 0.12 cm in diam (inside)

and 0.13 cm (outside) and 0.12 cm tall. Central collar is 0.03 cm in diameter

(opening) thickened to 0.02 cm and slightly raised. Pollinarium with two

long pollinia, 0.57 mm long and 0.14 mm wide; translators prominent, 0.10

mm long and 0.02 mm wide; caudicles prominent; the retinaculum has a

broad head and body with long well developed extensions. Fruit not seen.

Similar to Hoya gigantanganensis Kloppenburg but the leaves are

ovate lanceolate with an obtuse base versus linear-elliptic with a cuneate

leaf base; the calyx lobes are round not triangular, the corolla is slightly

smaller. In addition the pollinium are also shorter in this new species.

The garden clone originated from a 1997 collection of David Bicknell

taken from the wild in Matutinao in Badian, Cebu, The Philippines, and

grown to flower by Ted Green in Hawaii in the Spring of 2003. The original

plant was collected from forest at about 50 ft elevation in a coastal mountain

a few meters from the seashore. With the report of this new species, the

Philippine Hoya has now 69 species, many of which are local endemics.

oS)

io)

ie)

Gard. Bull. Singapore 61 (2) 2010

Figure 4. Hoya benitotaniti Kloppenburg, details of inflorescence: A. Top view of the calyx

(x8). These sepals are very rounded as opposed to the triangular sepals found on H.

gigantanganensis; B. Side view of the pedicel (x8), which is actually pale violet or rose in color;

C. Side view of flower with the corona removed showing the revolute corolla and the ovaries

(x8); D. & E. Inner (D) and outer (E) surfaces of the corolla (x8). Inner surface is pubescent

(D), outer surface is glabrous (E); F. View of the flower (x8). The crown is very similar to that

of Hoya gigantanganensis, except that the inner lobes are not as long, leaving much more of

the anthers exposed; G. & H. Upper (G) and lower (F) views of the corona (x8). Note the

inner lobes flare out toward the apex and do not exceed much from the anthers in the center; I.

Pollinarium enlarged («150). The pollinia are very long, translators and caudicles prominent;

the retinaculum has a broad head and body with long well developed extensions.

oS)

io)

New Species of Hoya from Brunei and the Philippines

References

Kloppenburg, R.D. 1991. Philippine Hoya Species. Orca Publisher Co.

Klopppenburg, R.D. and A. Wayman. 1993. The Hoya Handbook: A Guide

for the Grower and Collection.

Kloppenbur, R D.S.V. Siar. 2009. Additional four new species of Hoya R.Br.

(Apocynaceae) from the Philippines. Asia life Sciences 18(1): 139-154.

Wanntorp, L., A. Kocyan and S.S. Renner. 2006. Wax plants disentangled: a

phylogeny of Hoya (Marsdenieae, Apocynaceae) inferred from nuclear

and chloroplast DNA sequences. Molecular Phylogenetics and Evolution

39: 722-733.

Wennstrom, A. and K. Stenman. 2008. The Genus Hoya — Species &

Cultivation. Botanova. 144 pp.

Gardens’ Bulletin Singapore 61 (2): 335-341. 2010 335

Pecteilis rawatii (Orchidaceae), a New Species from India

P KUMAR AND JE VELDKAMP>

"Department of Habitat Ecology, Wildlife Institute of India, Post Box # 18,

Dehradun - 248001, Uttarakhand, India.

: Email: pankajsahani@rediffmail.com

~ National Herbarium of The Netherlands, Leiden University, Post Box 9514,

2300 RA, Leiden, The Netherlands.

Abstract

Pecteilis rawatii, sp. nov. (Orchidaceae) is described and compared to its

nearest ally, P. triflora.

Introduction

The genus Pecteilis Rafin. (Orchidaceae) is represented by 5 to 12 species

distributed from India to South China and Malesia. In India there are 4

species (Misra, 2007; Govaerts ef al., 2008). It is similar to Habenaria Willd.

and Peristylus Blume on the basis of the separate stigmatic lobes, but

not freely extending from the column as in Habenaria, nor adnate to the

hypochile as in Peristylus. The rostellum is characteristic in forming a broad

band above the stigmatic lobes.

While studying the orchids of Chotanagpur (Kumar ef al., 2007), a

species that turned out to be Pecteilis triflora (D.Don) T. Tang & F.T. Wang was

collected (P. Kumar 051062, WII). Pecteilis triflora was originally described

based on a Wallich’s collection from Nepal (Original material: Wallich 7035,

A,K! IDC microfiche 7394; *CAL, L!), but is known to be distributed also in

India (Chotanagpur plateau and Uttarakhand) and Bangladesh (Govaerts

et al., 2009).

During our research we came across a publication by Hooker f.

(1895), who, based on an illustration by Mr. G.C. Dass, described what he

thought was an aberrant form of Habenaria triflora D. Don (the basionym

of P. triflora). He noted that it was a plant of unknown origin, “presumably

flowered in the Calcutta Botanic Garden” (Fig. 1) and remarked that it

represented a many-flowered, gigantic state of H. triflora, mentioning some

other differences as well. He did not describe it as new as he thought that

the best course would be to refer it doubtfully to H. triflora.

This publication was largely overlooked until Pradhan (1976, 1979)

described Platanthera triflora (D. Don.) Pradhan var. multiflora Pradhan

based on the report. Unfortunately this trinomial was invalidly published as

336 Gard. Bull. Singapore 61 (2) 2010

the diagnosis was in English, and not in Latin as is required after 1 January

1935. We now agree that it is a taxon different from Pecteilis triflora, and

after having seen living material of P. triflora, it became clear that this is not

a mere variety, but a quite distinct species (Table 1).

Pecteilis rawatii P. Kumar & Veldk., sp. nov.

Syn. Platanthera triflora (D. Don.) Pradhan var. multiflora Pradhan, Indian

Orchids: Guide Identif. & Cult. 1:57 (1976), inval., anglice; ibid. 2:680 (1976),

sine descr.

Pecteilis triflorae (D. Don) T: Tang & ET: Wang similis, foliis bracteatisque

fasciculatis, floribus plus quam 3, labii lobis lateralibus apice rotundato,

margine integra, lobo medio lobis lateralibus longiore, columna latiore

differt. Pecteilis triflora folia breacteaeque distantiter dispersae, flores haud

plus quam 3, labii lobi laterales apice acuto, margine crenulata, lobo medio

lobis lateralibus multo breviore, columna quadrata gaudet. — Typus: Hooker

ff, Ann. Roy. Bot. Gard. (Calcutta) 5: t. 99.1895. Figs. 1 & 3.

- Plant terrestrial, herbaceous, erect with stout stem. Leaves many, green,

glabrous, single veined, densely placed in the lower half on the plant, base

sheathing, blade ovate, acute, margins entire. Leaves smaller and narrower

upwards, gradually decreasing in size, becoming bract-like. Inflorescence

terminal, short and dense, erect. Floral bracts green, glabrous, many, single

veined, base slightly sheathing the ovary, ovate, acute, margins entire, longer

than ovary. Flowers 9, pure white, densely placed. Sepals subequal in size,

midvein longitudinally ridged on the outer side, with two veins on either

side. Dorsal sepal ovate, apex obtuse, slightly notched. Lateral sepals broadly

ovate, apex obtuse. Petals linear, 1-veined, slightly constricted in the lower

half. Labellum trilobed, partition of lobes extending till the centre of the

lower half of labellum; lateral lobes broadly ovate, gradually constricting

towards the base, with a transverse basal callus, entire, apex rounded;

midlobe linear, distinctly longer than the lateral lobes, 1-veined, apex acute;

spur with a wide opening in the hypochile, gradually tapering and then again

broadening towards the apex, much longer than ovary. Column concave;

rostellum forming a broad transverse band above the broad, slightly concave

stigmatic lobes. Anther locules obpyriform, diverging towards the apex of

the column. Pollinia oblong, apex acute, caudicle thin, viscidium present

(Fig. 1):

Distribution. India, possibly once cultivated in Calcutta, original provenance

unknown.

Pecteilis rawatii (Orchidaceae), a New Species from India 337

Figure 1. Pecteilis rawatii P. Kumar & Veldk. A. Plant; B. Flower (front view); C. Flower

(dorsal view); D. Dissected flower and column: E. Labellum; F. Pollinia; (illustration redrawn

from Hooker f., 1895 by P. Kumar).

338 Gard. Bull. Singapore 61 (2) 2010

: i) ‘S=

11 mm

Figure 2. Pecteilis triflora (D. Don) T. Tang & F.T. Wang. A. Plant; B. Flower; C. Dissected

flower (floral bract, dorsal sepal, lateral petal, lateral sepal and labellum); D. Column; E.

Pollinia; F. Single pollen (based on P. Kumar 051062, WII). (Drawn by P. Kumar).

Pecteilis rawatii (Orchidaceae), a New Species from India 39

gé

Figure 3. Pecteilis triflora (D. Don) T. Tang & F.T. Wang (A-E). A. Habit; B. Close up of flower;

C. Dissected flower; D. Column: E. Pollinia. Pecteilis rawatii P. Kumar & Veldk. (F-G). F.

Habit; G. Close-up of flower [A-E: based on P. Kumar 051062, WII, photos by P. Kumar; F &

G, courtesy of Singapore Botanic Gardens].

340

Gard. Bull. Singapore 61 (2) 2010

Etymology. Named after Prof. Dr. Gopal Singh Rawat, Wildlife Institute of

India (WII), who is one of the leading phytotaxonomists and ecologists of

India and a source of inspiration for his students including the first author

and colleagues.

Table 1. Differences between P. rawatii (Fig. 1) and. P. triflora (Fig. 2).

Character Pecteilis rawatii Pecteilis triflora

states

Leaves Densely placed, 5 or 6, apex | Scattered on the stem, many, apex

acute but not sharply pointed. sharply pointed _acute.

Inflorescence | ca 9 flowered 1-3 flowered

Flower Sepals and _ petals widely | Sepals and petals forming a hood,

spread, each with two lateral | each with three lateral veins on each

veins on each side of the | side of the midrib. Base of lateral

midrib. Base of lateral sepals | sepals truncate.

oblique.

Labellum Midlobe much longer than the | Midlobe (much) shorter than the

side lobes. Margins of lateral | lateral lobes. Margins of lateral

lobes entire and apex rounded. | lobes crenulate and apex acute.

Base of midlobe broadly | Base of midlobe elongated and

cuneate. shortly attenuate.

Column Column broader towards the | Column more or less square

upper side.

Acknowledgements

The authors wish to thank the Mr. P.R. Sinha, Director (WII); Dr. G.S. Rawat

(WII); Dr. C. Sathish Kumar, Coordinator, All India coordinated research

project on the Taxonomy of Orchids; Dr. J.R. Bhatt, Director, GOI-MOEF;

Dr. D. Scherberich (LYJB); Mr. P. Boyce, Kuching, Sarawak, Malaysia; Dr.

P. Cribb, Dr. J.J. Wood and Renata Borosova (K) for translations from the

Latin, provision of digital images of type materials and valuable suggestions.

The authors are also thankful to Dr. Jana Leong-Skorniékova, Singapore

Botanic Gardens (SING), for her valuable suggestions and also for providing

the images of colour plates from Hooker (1895).

sae = e = = = ‘2

Pecteilis rawatii (Orchidaceae), a New Species from India 341

References

Don, D. 1825. Prodromus florae nepalensis: 25. J. Gale, London.

Govaerts, R., M.A. Campacci, D.H. Baptista, P. Cribb, A. George, K. Kreutz

and J. Wood. 2009. World Checklist of Orchidaceae. The Board of Trustees

of the Royal Botanic Gardens, Kew. Published on the Internet; http://

www.kew.org/wcsp/ accessed on 10 June 2009; 14:10 IST.

Hooker, J.D. 1890. Orchidaceae, p. 142. In: The Flora of British India 6. L.

Reeve & Co, Brook nr Ashford.

Hooker, J.D. 1895. A century of Indian orchids. Annals of the Royal Botanic

Gardens (Calcutta) 5: 66, t. 99.

Kumar, P., Jalal, J.S. and Rawat, G.S. 2007. Orchidaceae, Chotanagpur, State

of Jharkhand, India, checklist 3, 4: 297-304.

Misra, S. 2007. Orchids of India, a glimpse. Bishen Singh Mahendra Pal

Singh. Dehradun

Pradhan, U.C. 1976. Indian orchids. Guide to identification and culture 1:57.

U.C. Pradhan, Kalimpong.

Pradhan, U.C. 1979. Indian orchids. Guide to identification and culture 2: 680.

U.C. Pradhan, Kalimpong.

Gardens’ Bulletin Singapore 61 (2): 343-357. 2010 343

Land Use, Land Cover Change and Conservation in the

Dipterocarp Rain Forest Area of Southern Yunnan, China

J. LIANG '” ,H. ZHU ' AND Y.-X. MA !

Xishuangbanna Tropical Botanical Garden, Chinese Academy of Science,

88 Xuefu Road, Kunming 650223, China

5 Department of Life Science, Huaihua University, Huaihua 418008, China

Author for correspondence: zhuh@xtbg.ac.cn

Abstract

Based on Landsat TM/ETM images from 1988, 2003 and field data of 2006,

land uses and land cover changes were researched over 18 years in the

dipterocarp rain forest area in Southern Yunnan of China. The expansion

of rubber plantations has resulted in a dramatic decrease in natural forest

cover, especially the tropical seasonal rain forest at lower elevation. In 1988,

rubber plantations covered 765.06 ha which increased to 2,294.07 ha in 2003,

with an annual rate of change at 13.32%.The pace of change increased after

2003, with a change of 213.69 ha per annum. The tropical seasonal rain

forest has decreased by 111.35 ha per annum since 1988 in the study area.

Arable lands increased during 1988 and 2003 but declined rapidly from

2003 to 2006 due to expansion of rubber plantations and the construction

of reservoirs. There was an increase in water bodies from 2003 to 2006 as

well as construction areas. Market prices, policies, increasing population,

and the unregulated pursuit of commerce and trade, at times at the cost

of the environment were the main driving forces of change. We suggest

that local government takes strong action to regulate further expansion of

rubber plantations and creates conditions for sustainable and harmonious

development of economy, society and natural resources in biodiversity rich

region of Southern Yunnan.

Introduction

With the increasing concern about global climate change and biodiversity

conservation, study of land uses and land cover changes (LUCC) has become

an extremely important and hot topic (Li, 1996). LUCC is one of the most

important human alterations affecting the surface of the earth (Lambin er

al., 2001). It directly impacts biodiversity (Sala et al., 2000), contributing

to local extinction and regional climate changes (Chase et al., 1999), and

causing land degradation by altering ecosystem services and livelihood

support systems (Vitousek et al., 1997).

344 Gard. Bull. Singapore 61 (2) 2010

There have been dramatic land uses and land cover changes in some

places in China in recent years. By using remote sensing (RS) and geographic

information system (GIS), Chinese scholars have carried out many studies

on land use and land cover change detections and their impacts on the

environments (Gao et al., 2002; Li et al., 2003, 2007; Liu et al., 2006).

Southern Yunnan bordering Laos and Myanmar,is in a biogeographical

transition zone between tropical SE Asia and temperate East Asia, and

is one of the most biodiversity-rich regions in China (Zhu, 1997). The

dipterocarp rain forest is the most species-rich plant community in southern

Yunnan (Zhu, 2000). It is dominated by the species Parashorea chinensis

(Dipterocarpaceae), a first grade protected plant species at national level.

However, during the last few decades, forest cover has dramatically decreased

from 60% to 27% (Zhang and Cao, 1995; Liu et al., 2005). Currently, forests

remain primarily in nature reserves and state forests, and land areas outside

nature reserves, which were previously forested, have largely been converted

into rubber plantations. There is an essential and urgent research need to

survey the land uses and land cover changes in tropical region in China in

order to collect and update scientific data and to provide suggestions on

_ how to improve the land use policy in a way that balances economic needs

with biodiversity conservation. Our work here focuses on a dipterocarp rain

forest area in Southern Yunnan and presents findings for a discussion on the

land uses and land cover changes. It also shows the underlying driving forces

in the local tropical rain forest and, thereby, provides a scientific basis for

biodiversity conservation and management of the forest in situ.

The Study Area

The study area is located in the administrative region of Xishuangbanna,

which includes three counties (Menghai, Jinghong and Mengla), and borders

on Laos to the South and Myanmar to the Southwest (Fig. 1). The topography

of Xishuangbanna consists of alternating hills and valleys, with elevations

ranging from 2,430 m above sea level (asl) in the North and 480 m asl in

the South. About 95% of the region is mountainous and hilly. The Mekong

River cuts through Xishuangbanna, and the region contributes more than 20

important tributaries, resulting in many river valleys and small basins (Cao

and Zhang, 1997). This region has basically a tropical monsoon climate. In

the lower hills and valleys, the annual mean temperature is about 20° C, and

frost has never been recorded. The annual precipitation is about 1,500 mm,

of which more than 80% falls during the rainy season, which starts in May

and lasts until the end of October. The dry season from November to April

is characterized by little rain, but there is always heavy fog and dew, which

can compensate for lack of rain. Thus the tropical rain forest in the region

occurs only at lower elevations. It was also revealed that the occurrence

Land Cover Change and Conservation in the Dipterocarp Rain Forest Area of Southern Yunnan, China 345

of the tropical rain forest in southern Yunnan is more influenced by local

habitats and microclimates than the regional climate (Zhu, 2004, 2008)

ae ee ed

Jinghong

s Menghai a Ss

Mengla

Study area |

Figure 1. Xishuangbanna region and the location of study site.

The dipterocarp rain forest in Xishuangbanna is located mainly in

the South of Mengla County (Zhu, 2000). Our study site (21°33'~21°38' N,

101°32'~101°41'E) covers the whole distribution of dipterocarp rain forests

encompassing an area of 125 km . The altitude varies from 650 m to 1,600 m

asl. The dipterocarp rain forest occurs in the area mainly below at 900 m asl,

and other vegetation types such as tropical montane evergreen broad-leaf

forest occurs in the mountains (Zhu et al., 2005).

Materials and Methods

Data Sources

We obtained a Landsat Thematic Mapper (TM) image (2 February 1988-

#130/45) and a Landsat Enhanced Thematic Mapper (ETM) image (7 March

2003-#130/45) of the study area. Both images were acquired during the dry

season. Four land use maps (scale = 1:25000) developed by the Xishuangbanna

Department of Land and Resource Management (Xishuangbanna Land

Use Status Map, 1991) were used as references for the classification of the

TM image. Four topographic maps (scale = 1:25,000) published by the China

State Bureau of Surveying and Mapping were used to correct the images of

1988 and 2003 and to draw the land use map of 2006.

346 Gard. Bull. Singapore 61 (2) 2010

1988

2003

2006

[_] Arable lands

feeael Rubber plantations N

GE Construction area

HB Water area / N

Tropical seasonal rain forest

Tropical montane evergreen broad-leaved forest

4 8 12 Kilometers

Figure 2. Land use in the distribution area of dipterocarp rain forest in Xishuangbanna, SW

China in 1988, 2003 and 2006.

Land Cover Change and Conservation in the Dipterocarp Rain Forest Area of Southern Yunnan, China 347

Data processing

Based on the land use classes developed by the National Agricultural

Zoning Committee (1984), as well as characteristics of the images of study

area, land uses were categorized into 5 classes (Table 1): arable lands, rubber

plantations, forests, developed/constructed areas and water bodies. Due

to low resolution of images, it was difficult to classify the land use types

in detail; hence, the forest was simply divided into tropical seasonal rain

forest and tropical montane evergreen broad-leaf forest based on elevation.

Shrublands were incorporated into forests, grasslands and sugar cane fields

were incorporated into arable lands.

The TM and ETM images had already undergone radiant correction,

so we only had to do the geometric correction. Both images were rectified to

the topographic maps and the rectification errors were <0.5 pixels.

After the correction, the images were classified using the supervised

maximum likelihood classification method. For each land use type, at least

10 training areas were used to reflect the variation within a land use due to

topography and slope effects.

The classified images were then transformed by using clump,elimination

and filter options in Erdas Imagine (Version 8.7, Leica Geosystems). Then

with the ArcView GIS (Version 3.3, Environmental Systems Research

Institute, Redlands, USA) and ArcGIS (Version 8.3, Environmental Systems

Research Institute, Redlands, USA), the transformed images were converted

to land use maps for analysis.

The paper topographic map was firstly magnified and used in the field

to draw the outline of the land use map of 2006 by using contour, topography

and other information from the topographic map. The current land use

cover was determined at the field site and the locations were determined by

using a global positioning system (GPS). The GPS points were also used to

prepare the land use map of 2006 and to assess the accuracy of the land use

maps of 1988 and 2003. The field works lasted two weeks and covered the

whole study area. The working map was scanned and digitized, and finally

was made into a land use map of 2006 using ArcGIS.

The accuracy of the classification of both images of 1988 and 2003 was

verified by the points obtained in the field in 2006. All land use maps (1988,

2003, 2006) were rectified to Albers Conical Equal Area projection system,

with Beijing 54 datum and Krosovsky ellipsoid. By using ArcView GIS, the

database of the land uses and land cover changes of 1988, 2003 and 2006

was determined, which was used to analyze land use and land cover changes

during the research period.

348 Gard. Bull. Singapore 61 (2) 2010

Table 1. Land use classes used in our image classification.

Land use class General description

Arable lands (AL) Shifting cultivation or permanent agriculture (e.g.

paddy rice, dry lands, fallow lands, slash and

burn)

Rubber plantations (RP) Forested areas with rubber trees clearly planted in

rows and deciduous during the dry season,

having a homogeneous canopy

Tropical seasonal rain forest Forested areas with a canopy cover more than

(TSRF) 30% and below 900 m above sea level.

Tropical montane evergreen Forested areas with a canopy cover more than

broad-leaf forest (TMEBF) 30% and above 900 m above sea level.

Construction areas (CA) Land covered by constructions, including urban

areas, residential areas, factory and traffic areas

Water areas (WA) Natural terrestrial water, including river, pool,

lake and reservoir

Notes: The construction areas in the study site are dispersed and too small

to be distinguished from the images, so we just made a simple estimate of the

increased area from 2003 to 2006. There are many rivers in the study area,

but most of them are also too small to be distinguished from the images, so

we only considered the charge of Nanla River, the biggest river in the study

area.

Data analysis

The annual change rate index was used to reflect the rate of land use and land

cover change (Wang et al., 1999), and it may be expressed by the following

equation:

OG, 0,

<= ! ee

In this expression, K was the annual rate of change of one land use

class, U. and U» were the initial and the final area values of the research

Land Cover Change and Conservation in the Dipterocarp Rain Forest Area of Southern Yunnan, China 349

period respectively, and T was the time (year) of the period.

The contribution rate of LUCC was used to reflect the source of

existing land use class, and it could be expressed as follows:

Bi = 7 xl00%

In this expression, B,,; was the contribution rate of land use type / to j

from the initial time a to the ending time b of the research period, U;;; was

the area that land use type / turned to j, and U,;was the area of land use type

jin time b.

In order to analyze the change of the major land use type (Zhu et al.,

2003), there is another index—the intensity index of LUCC.

pee a). ese 1000 %,

U t &

In this expression, K is the change intensity index of the land use class /,

Uis the area of the whole study site, U,,,, U,,;is the initial and the final area of j of

the research period respectively, and 7 is the duration time (year) of the period.

Results

Area changes in land cover

Changes in land cover in the study area from 1988 to 2006 are shown in Fig.

2 and summarized in Table 2. The most obvious change was the decrease

in forest cover and an increase in rubber plantations. In 1988, the area of

rubber plantations was 765.06 ha, which increased to 2,294.07 ha in 2003,

with an annual rate of change at 13.32%. However, the expansion of rubber

plantation has further accelerated since 2003, increasing 213.69 ha per

annum. The forested area declined rapidly from 1988 to 2006, especially

the tropical seasonal rain forest at lower elevation, at a decreasing rate of

111.35 ha per annum. The forest at higher elevation also declined rapidly

during 2003 and 2006. The arable lands increased during 1988 and 2003 but

declined rapidly from 2003 to 2006. Water body in the study site did not

change before 2003, but abruptly increased after 2003 due to the building

of a big reservoir-Dashaba Reservior on Nanla River. The open-up area for

settlement also increased with the building of the reservoir.

Transitions in land cover

Table 3 and Table 4 are contribution matrixes showing resources of existing

350 Gard. Bull. Singapore 61 (2) 2010

Table 2. Area changes of land use classes of the study area in different time period.

AL RP TSRF TMEBF WA

Area ( ha)

1988 389.51 765.05 4653.50 6709.90 40.16

2003 509.59 2294.07 3202.47 6511.83 40.16

2006 403.68 2935.13 2649.23 6236.41 333.67

% of area coverage

1988 3.10 6.09 37.06 53.43 0.32

2003 4.06 18.27 25.50 51.85 0.32

2006 3.21 PE} Sy) 21.10 49.66 2.66

Change area (_ ha)

1988 — 2003 120.08 1529.02 -1451.03 -198.07 0.00

2003 — 2006 -105.91 641.06 -553.24 -275.42 293.51

Annual change area (_ ha)

1988-2003 8.01 101.93 -96.74 -13.20 0.00

2003-2006 -35.30 213.69 -184.41 -91.81 -

Annual change rate (%)

1988-2003 2.06 13.32 -2.08 -0.20 0.00

2003-2006 -6.93 9.31 -5.76 -1.41 -

Notes: The increase of water area is due to the construction of the reservoir and it is

a once-off activity, so we don’t calculate its annual change area and annual change

rate. [AL: arable lands; RP: rubber plantations; TSRF: tropical seasonal rain forest;

TMEBF: tropical montane evergreen broad-leaf forest; WA: water area.|

Table 3. Contribution matrix of land-cover categories in the study area between 1988 and

2003 (%).

2003

1988

Arable lands — Rubber plantations _—- Water area TSRF TMEBF

Arable lands 28.49 9.66 0.00 0.71 0.00

Rubber plantations 31.23 24.78 0.00 1.10 0.08

Water area 0.00 0.00 100.00 0.00 0.00

TSRF 35.17 58.00 0.00 98.19 0.01

TMEBF 5.11 7.56 0.00 0.00 99.91

(5%)

Land Cover Change and Conservation in the Dipterocarp Rain Forest Area of Southern Yunnan, China

Table 4. Contribution matrix of land-cover categories of study area between 2003 and 2006

(%).

2006

2003

Arable lands Rubber plantations Water area CA TSRF TMEBF

Arable lands 24.23 8.71 34.19 fA 0.71 0.08

Rubber plantations 47.23 60.75 39.40 24.23 5.88 0.66

Water area 0.00 0.01 12.22 0.52 0.00 0.00

TSRF 11.52 21.78 14.19 4.12 93.41 0.00

TMEBF 17.02 8.75 0.00 0.00 0.00 99.26

land cover classes of 2003 and 2006. Except for the water area. all other

land use categories contributed to increasing rubber plantations. Increase in

rubber plantations took place in the tropical seasonal rain forest. The water

body area increased between 2003 and 2006 at the expense of arable lands

as well as rubber plantations. The cleared settlement area also increased

with the completion of the reservoir, mainly at the expense of arable lands

and rubber plantations. A large part of tropical mountain evergreen broad-

leaf forest converted to agricultural use between 2003 and 2006 was the

result of sugar cane planting.

Change intensity in land cover

Fig. 3 shows the change intensity index of land use categories from 1988 to

2006. The intensity of change between 2003 and 2006 is much higher than

that between 1988 and 2003. Analyzing the change intensity of these two

periods respectively, we found that rubber plantation was the major land

cover category of LUCC between 1988 and 2003, with a change intensity

index of 8.12%o, followed by the tropical seasonal rain forest, with a change

intensity index of -7.70%o. Rubber plantation was still the major land-cover

category of LUCC after 2003, with a change intensity index of 17.02%o,

followed by tropical seasonal rain forest, water bodies, tropical montane

evergreen broad-leaf forest, and arable lands.

Discussion

This study investigated the changes in land use and land cover of dipterocarp

rain forest area in Xishuangbanna from 1988 to 2006. The results showed a

dramatic change in land use and land cover in the study area. The most

obvious change was a decrease in forest cover and an increase in rubber

plantations. From 1988 to 2003, rubber plantations increased at a rate of

35) Gard. Bull. Singapore 61 (2) 2010

20.00

D0 1988 — 2003

15.00

B 2003 — 2006

8 10.00

ZB 5.00

= 0.00

Ss -5.00

-10.00

-15.00

20.00 AL RP TSRF TMEBF WA

Figure 3. Comparison of land use intensity (using the permillage to describe) of study area

from 1988 to 2006. [AL: arable lands; RP: rubber plantations; TSRF: tropical seasonal rain

~ forest; TMEBF: tropical montane evergreen broad-leaf forest; WA: water area.]

101.93 ha per annum, between 2003 and 2006 this rate increased to 213.69

ha per annum. In contrary, the area of tropical seasonal rain forest and

tropical mountain evergreen broad-leaf forest declined continually over

time, especially the tropical seasonal rain forest at a rate of 111.35 ha per

annum. In recent years, tropical mountain evergreen broad-leaf forest has

declined faster as a result of rubber plantations expanding also to higher

elevations in the region. Arable lands also decreased rapidly between 2003

and 2006 from the expansion of rubber plantations and the building of

Dashaba Reservoir on Nanla River. The normal water level of the reservoir

was about 680 m asl. After the construction of reservoir, some arable lands

were submerged and others were replaced by buildings of hydropower

station, house, hotel, parking sites, recreation ground, etc., which resulted in

an increase of constructed area from 2003 to 2006.

The research also showed that almost all land use categories, in

particular the tropical seasonal rain forest at low elevation, contributed

to the increase of the rubber plantations. The change intensity of LUCC

also became larger than before and the remaining forests, including the

dipterocarp rain forest, are threatened by the expanding rubber plantations.

Increasing population is the basic driving force at LUCC, with the

increasing demand for food, housing and public facilities, which resulted in

the expansion of rubber plantations, arable lands and construction areas.

2&7

Land Cover Change and Conservation in the Dipterocarp Rain Forest Area of Southern Yunnan, China 259)

Bubeng Village

1050 -

1000 -

= ee Be

& 900 -

850 -

800

Nee Poe ic SG AS OD Did

SH DS DMP MMMA MA

ne A) SS WS? 5) SS) OS) oS AS

Time

Jingpiao Village

Population

750 +

7 aaa

ar ey es 8 2! AG! Me aes aE Rp

PD BY DB? DM? MD HM A

PoP SD SAD DP DP PP aD of

Time

Figure 4. The changes of population of Bubeng and Jingpiao Village in recent years.

Gard. Bull. Singapore 61 (2) 2010

The population of the study area increased rapidly during the past 18 years.

Figure 4 shows the populations of Bubeng and Jingpiao Village of the study

area from 1988 to 2002.

The Household Responsibility Policy provided land to individual

households and the local people have the rights to manage their own lands.

With the development of the market economy in China, the increasing

market price has a profound effect on land use. The price of natural rubber

is constantly rising, which increases people’s enthusiasm for rubber planting.

As a result, rubber plantations became the major income source for local

people in the study region.

The dipterocarp rain forest is the most species-rich community in

southern Yunnan and is dominated by the species Parashorea chinensis

(Dipterocarpaceae), a first grade protected plant species at national level.

Similar tropical rain forests occur also at ca 27° 31° N in northeastern India

(Proctor et al., 1998) and Burma (Kingdon-Ward, 1945), but these places still

have basically tropical wet climates due to their geographical location and

lower elevations.

The tropical rain forest in southern Yunnan occurs really at the

_ northern climatic limits of tropical rain forest and is unique in biogeography

and biodiversity conservation. The forest is at present strongly protected

by the government policy. However, over the past 18 years, there has been

a dramatic land uses and land cover changes in its area of distribution. In

our research, we found that almost all forests and shrubs outside the nature

reserves were converted to rubber plantations. In some places, even the

nature reserves were nibbled away by rubber plantations. The dipterocarp

rain forests have become quite fragmented and exposed to other external

impacting factors. These include changes in the micro-climate of the habitat

and the physicochemical property of the soil, which would consequently

change the characteristics of the plant communities and their inherent

biodiversity (Liu et al., 2001, Sophia et al., 2000, Zhu et al., 2004).

We suggest that local government takes strong action to control further

expansion of rubber plantation and creates a sustainable and harmonious

development of the local economy, the society and natural resources in the

region of biodiversity rich dipterocarp forest area in southern Yunnan.

Acknowledgements

This project was funded by the National Natural Science Foundation of

China (30570128, 30770158), also by the grant 2007 OFA 91660-5. We thank

M. Ma who provided invaluable assistance with data collection and H.

Wang, S.-S. Zhou who assisted with fieldwork. We also offer thanks to H.-

M. Li, W.-J. Liu, Y.-J. Zhang and L.-P. Yang for their helpful comments on

Land Cover Change and Conservation in the Dipterocarp Rain Forest Area of Southern Yunnan, China 395

the manuscript. We are very grateful to Mr. Hasan Moinuddin from GMS

Environment Operations Center for his help in English improvements on

the manuscript.

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ioe)

Ne)

Siliquamomum oreodoxa (Zingiberaceae): a New Species

from Southern Vietnam

N.S. LY .S. HUL’ AND J. LEONG-SKORNICKOVA’

‘The VNM Herbarium, Institute of Tropical Biology, 85 Tran Quoc Toan Road,

District 3, Hochiminh City, Vietnam. E-mail: sahucultural96@yahoo.com

"Département de Systématique et Evolution, Herbier (P)—Plantes vasculaires, 57

; Rue Cuvier, 75231 Paris Cedex 05, France. E-mail: hul@mnhn.fr

The Herbarium, Singapore Botanic Gardens, 1 Cluny Road, 259569 Singapore.

Email: jana_skornickova@seznam.cz

Abstract

The second species of Siliquamomum (Zingiberaceae), S. oreodoxa N.S.Ly

& Skornick.,is described as new and illustrated. The two species in the genus

are compared and a key is provided for their identification.

Introduction

The genus Siliquamomum was described by Baillon (1895) based on a

collection made by Benedict Balansa in Lankok valley of Ba Vi Mountain,

Tonkin (North Vietnam). The only representative of the genus known till

date, S. tonkinense Baill., has also been reported from SE Yunnan, China

(Wu and Larsen, 2000; Gao et al., 2005) where it is found in dense forests 1n

mountain valleys at 600-800 m elevation.

During an expedition in southern Vietnam in June 2008, the first

author found a Siliquamomum species growing quite abundantly at ca 1500

m in a moist and shady area of Bidoup Nui Ba National Park. The type

material of S. tonkinense (Balansa 4218, Nov 1887, 4 sheets, P) was located

and examined and the original description by Gagnepain (1908) studied.

The material collected by us is different in several significant characters

and a new species is therefore described below including a colour plate. In

addition, a key to the two species of Siliquamomumi is provided.

Siliquamomum oreodoxa N.S. Ly & Skornick., sp. nov.

Siliquamomo tonkinens!i similis, surculo foliaceo multifolio (foliis 9-13 contra

3), petiolo breviore (ad 2 cm contra 2.5-9 cm longum), laminis minoribus (12-

18 x 3-4.5 cm contra 25-42 x 7-12 cm), inflorescentia breviore sed densiore

(ad 9 cm longa floribus 8-25 contra 13-15 cm longum floribus ca 12) differt.

— Typus: Vietnam, Lam Dong Province, Lac Duong District, Bidoup Nui

360 Gard. Bull. Singapore 61 (2) 2010

Ba National Park, Cong Troi station, along the road between Yang Ly and

Hon Giao station, alt. 1449 m asl., 12° 10’ 41.9” N, 108° 43’ 26.4” E, 14 Jun

2008 (flowering), Hul & Ly Ngoc Sam 3583 (holo, VNM; iso, E, P, SING with

flowers in alcohol). Figs. 1 & 2.

Terrestrial herb forming loose clumps. Rhizome subterranean, branched, ca

0.8-1.2 cm in diam., creamy white externally and internally, slightly aromatic;

scales papery, brown, triangular scales, ca 1-2 cm long. Leafy shoots ca 3-5 cm

apart, 0.9 m long, leafless for about 15-30 cm from the base, with 9-13 leaves

per shoot. Leafless sheaths 3-5, yellowish green when young, turning dark

brown-green with age, glabrous with dark brown margins. Ligule ca 2-3 mm

long, bilobed, dark brown, becoming papery and brittle with age, glabrous.

Petiole 1-2 cm long, green, glabrous. Lamina elliptic to narrowly ovate, (9.5-)

12-18 x (2.5-) 3-4.5 cm, dark green and glossy above, slightly lighter beneath,

apex acuminate, base cuneate, margin entire. Inflorescence terminal,peduncle

enclosed by leaf sheaths of the pseudostem, terminated by a lax pendulous

thyrse with 8-25 flowers, first (lowermost) and second bract at the base of

thyrse boat-shaped, 4.5-7 x 1.8-2.5 cm, light green, glabrous, sometimes

- with a small lamina at the apex, enclosing the budding inflorescence, soon

caducous, leaving semicircular scars on the axis, fertile bracts narrowly ovate

to elliptic, the bract subtending the lowermost cincinnus the biggest, ca 25-30

x 6-8 mm, those subtending upper cincinni usually minute, bluntly triangular

ca 1-8 x 1-1.5 mm, whitish green, turning papery brown, soon dehiscent

(best seen in young inflorescences with unopened buds), axis of thyrse to

9 cm long (measured from the lowermost caducous bract to the top of the

axis), to 2 mm diam., light green, glabrous, cincinni 1-3 flowered at the base

of the axis, single flowered at the top, bracteoles absent. Pedicel 3-6 mm, ca

1 mm diam., light green, glabrous. Flower ca 5.5 cm long, calyx tubular 18-25

mm long, 6-7 mm diam. at broadest, unilaterally shit ca 9-11 mm, translucent

white sometimes with a slight pink tinge near margin, glabrous, 3-toothed

at apex, teeth 3-5 mm long, 4 mm wide at base, becoming brownish and

papery with age. Floral tube 19-21 mm long, white, glabrous at base, sparsely

hairy at apex, dorsal corolla lobe elliptic-oblong, concave, 22-27 x 8-10 mm,

translucent white, glabrous, lateral corolla lobes oblong to obovate-oblong,

concave, 20-24 x 6-8 mm, translucent white, glabrous. Lateral staminodes

connate to labellum in basal third, oblong-obovate, apex rounded, 19-21 x

11-13 mm, white, glabrous, margin slightly undulate. Labellum obovate, 22-

26 mm long, 18-20 mm wide at broadest, apex obtuse to retuse, white at

periphery with dark green patch with yellow margins in the centre, adaxially

hairy along the midline and green patch, hairs ca | mm long, white, glabrous

abaxially. Stamen 15-20 mm long, filament 6-8 x 2-2.5 mm, white, sparsely

hairy at base. Anther 9-12 x 3-3.5 mm, attached to filament at 160°, white

Siliquaamomum oreodoxa (Zingiberaceae): a New Species from Southern Vietnam 361

7 8 9

! 1 2 3 4 5 6

1 2 3

(iil ated talstalateLadasslatstate silstal

Figure 1. Siliguamomum oreodoxa. A. Inflorescence; B. Plant habit; C. Ligules; D. Close-up

of flower; E. Sterile caducous bracts enclosing budding inflorescence; F. Rhizome. (Based on

Hul & Ly Ngoc Sam 3583: Photo by N.S. Ly)

9) : .

362 Gard. Bull. Singapore 61 (2) 2010

Ee ee ee

| a HIT] Pega

9 110 i

1 2 5 6

STEVE

Figure 2. Siliquamomum oreodoxa. A. Flower detail and dissection; B. Close-up of ovary

dissection; C. Close-up of central part of labellum with hairs along the midline; D. Close-up of

anther. (Based on Hul & Ly Ngoc Sam 3583; Photo by N.S. Ly).

or light cream, with glandular hairs on connective, apex emarginate, anther

thecae 9-11 mm long, white or light cream, dehiscing longitudinally for its

entire length, glabrous, each with a minute, tooth-shaped, ca 1 mm long

crest. Ovary narrowly cylindrical, 12-14 mm long, ca 1.2 mm diam. at base,

up to 2 mm at apex, light yellow-green, glabrous, trilocular, placentation

axile. Style 33-36 mm long, white, glabrous, stigma ca 1.2 mm diam., club-

shaped, top of club ciliate, ostiole ciliate, sub-apical, transverse elliptic, ca |

mm wide, facing forwards to upwards. Epigynous glands two, 2-3 mm long,

ca | mm diam, with blunt apex, cream to very light brown. Fruit not seen. All

measurements based on living and spirit material of the type.

Phenology. Flowering June to July.

Siliquamomum oreodoxa (Zingiberaceae): a New Species from Southern Vietnam 363

Habitat and ecology: Growing in moist and shady understorey of coniferous

and broadleaved mixed forest in Bidoup Nui Ba National Park at 1500-

1800 m. This forest type covers most of the Langbiang Plateau (or Lam

Vien Plateau) of Lam Dong Province. The prevailing climate in this area

is tropical montane with an annual rainfall of 1775 mm and an average

temperature of 18:C.

Etymology: Greek oreo, “pertaining to mountains” and doxa, “glory”. We

chose the epithet oreodoxa because this beautiful species thrives in the

mountains. The word oreodoxus (-a, -um) has often been regarded in botany

as an adjective, but in classical Greek it was regarded to be a noun, which,

when used as an epithet, cannot be declined.

IUCN assessment: In spite of several surveys of Bidoup Nui Ba National Park

and adjacent areas, Siliquamomum oreodoxa is only known from the type

locality situated immediately adjacent to a camp occupied by members of

a minorities ethnic group who have established agricultural fields and built

numerous roads to facilitate access to the area. These conditions present a

clear threat to the single known population of S. oreodoxa. Based on the

IUCN Red List criteria (IUCN 2001), we propose a provisional conservation

status of Critically Endangered (CR Blab(i1)+2ab(iii)); D).

Notes: The characteristic features of the genus Siliqguamomum, based on its

previously sole species S. tonkinense, include a leafy shoot with few leaves,

lax terminal raceme with flowers borne singly, minute bracts, bracteoles

absent, petaloid lateral staminodes, each anther theca with an apical linear-

acuminate crest and a trilocular ovary developing into narrowly fusiform

capsule (e.g., Larsen et al., 1998). Hul & Ly Ngoc Sam 3583 conforms with

all these characters except the higher number of leaves and the fact that the

flowers are produced in cincinni of 1—3 flowers. Even though we have not

seen ripe fruits, the ovary is conspicuously narrowly cylindrical and identical-

looking with ovary of S. tonkinense. It is therefore justified to describe

the new taxon in Siliquamomum and update some of the circumscriptive

characters of the genus, 1.e., the higher number of leaves and the presence

of cincinni.

Siliquamomum oreodoxa differs from S. tonkinense by several

characters (Table 1), the most important by its leafy shoot with many leaves

(9-13 vs.3),a shorter petiole (to 2 cm vs. 2.5-9 cm), a smaller lamina (12-18 x

3-4.5 cm vs. 25-42 x 7-12 cm), and a shorter and denser inflorescence (up to

9 cm long with 8-25 flowers vs. 13-15 cm long with ca 12 flowers).

The flowers of Si/iguamomum oreodoxa are mostly white, but the

labellum has a dark green patch with a yellow border in the centre. The

364 Gard. Bull. Singapore 61 (2) 2010

midline and the green—yellow coloured patch are shortly hairy. The original

description of S. tonkinense states merely that the flower is yellowish white

without any further details but from Gagnepain’s drawing attached to the

type, it appears that the labellum is glabrous. Based on this comparison, a

key to the two species was constructed and is provided below.

Colour pictures of S. tonkinense from Yunnan (Gao et al., 2005)

show yellowish white flowers, and a labellum with a yellow patch in the

centre ornamented with dark green lines. Some of the measurements of

S. tonkinense from Yunnan differ from those made from type material of

Vietnamese origin. Only a comparison of living material re-collected from

type locality in Vietnam can shed light on the question, whether the Chinese

material is indeed conspecific.

The genus Siliquamomum has been traditionally placed in the former

tribe Hedychieae (Burtt and Smith, 1972;Smith, 1981; Larsen er al., 1998; now

Zingibereae sensu Kress et al., 2002) on the account of its petaloid lateral

staminodes. The plane of the distichy as observed by us on S. oreodoxa is

transverse and the pseudostems are evergreen. Both of these features are

typical for Alpinioideae, while genera in Zingiberoideae have parallel plane

_ of distichy and most of the species have capacity of going into dormancy

during the dry season in monsoonal climates (Kress et al.,2002). The presence

of few caducous sterile bracts enclosing the budding inflorescence, is yet

another character common in some genera of Alpinioideae. The most recent

comprehensive study that dealt with the phylogeny of the whole family

Zingiberaceae (Kress et al.,2002) was unable to place Siligquamomum in any

of the two tribes, Riedelieae or Alpinieae, of Alpinioideae. The study noted

that the long silique-like fruits of Si/iguamomum are similar to those found

in the Riedelieae but it differs by the presence of a well developed labellum

and lateral staminodes, as well as the absence of the extrafloral nectaries on

the leaf midribs. Siliquamomum was therefore left as an uncertainly placed

genus in Alpinioideae. A later study focusing on molecular phylogeny of

Alpinia (Kress et al., 2005) placed Siliquamomum as sister to the rest of the

Alpinieae.

With asecond species of Siliquamomum collected and described, future

sequences may assist in confirming where the genus should be placed and

what morphological characters are important for the generic delimitation.

Key to Siliquamomum

1. Pseudostem with ca 3 leaves; petiole to 9 cm long; lamina 25-42 x 7-12 cm

RO ee ra ria Cet Scone onkoe Siliquamomum tonkinense

1. Pseudostem with 9-13 leaves; petiole to 2 cm long, lamina 9.5-18 x 2.5-4.5 cm

PT eT es tir Mira) Retr ycruicerasodadoon votes Siliquamomum oreodoxa

Siliquaamomum oreodoxa (Zingiberaceae): a New Species from Southern Vietnam 365

Table 1. A comparison of characters of S. oreodoxa and S. tonkinense (based on Baillon, 1895;

Gagnepain, 1908; our measurements of type herbarium material marked by *. Characters of

S. tonkinense from Chinese material stated in Wu & Larsen, 2000, and Gao et al., 2005 are

not included).

Characters S. oreodoxa S. tonkinense

Leafy shoot to 0.9 m long, at least 0.6 m high,

9-13 leaves per shoot 3 leaves per shoot

Lamina elliptic to narrowly ovate, ovate-lanceolate,

(9.5-) 12-18 x (2.5-) 3-4.5 cm 25-42* x 7-12* cm

Inflorescence to 9 cm long, with 8-25 flowers 13-15 cm long, with ca 12 flowers

—

Pedicel 3-6 mm long 5-9 mm long

18-25 mm long ca 25 mm long *

Lateral staminodes ca 19-21 mm long ca 8-9 mm long

Labellum obovate, 22-26 mm long, 18-20 orbicular, ca 12 mm in diam., flowers

mm at broadest point, adaxially yellowish white

hairy along the midline with green

and yellow patch, hairs ca 1 mm

long, white, abaxially glabrous

ca 9-12 mm long ca 11-12 mm long

club-shaped funnel shaped

cylindric, trilocular elliptic, unilocular at apex, 3 locular at

base

Acknowledgements

We thank the management of the Institute of Tropical Biology and

specifically Dr. Luu Hong Truong, the curator of VNM, for providing

research facilities. The French Government Project, Sud-Expert-Plantes

(SEP n°228: Improving Services and Capacity of the National Herbarium

of the Institute of Tropical Biology) is acknowledged for sponsorship of

the field trip, and the director and staff of Bidoup Nui Ba National Park

for granting permits to conduct it. We thank the curators of P and SING

for letting us examine specimens in their care and the Asian Zingiberaceae

Information Centre (AZIC) at Singapore Botanic Gardens for providing

protologues, related references and images of type specimens. The first

366 Gard. Bull. Singapore 61 (2) 2010

author thanks Dr. M. Newman (E) for introducing him to Zingiberaceae

taxonomy, to Mr. Truong Quang Tam and Mr. Nguyen Tran Vy for helpful

discussions and overall support. The Russell E. Train Education for Nature

(EFN) Program of World Wildlife Fund supported the first author’s training

on Zingiberaceae at Singapore Botanic Gardens. We are grateful to Dr. J.F.

Veldkamp (L) for translating the diagnosis into Latin and helpful discussion

on Greek etymology, Dr. A. Poulsen (E) and Dr. M. Newman (EB) for helpful

comments on the manuscript.

References

Baillon, M.H. 1895. Une Musacée-Zingibérée a fruit siliquiforme. Bulletin

Mensuel de la Société Linnéenne de Paris 2: 1193-1194.

Burtt, B.L. and R.M. Smith. 1972. Tentative keys to the subfamilies, tribes

and genera of Zingiberaceae. Notes from the Royal Botanic Gardens

Edinburgh 31: 171-176.

- Gagnepain, F. 1908. Zingibéracées. Pp: 25-121 in: Lecomte, H. (ed.), Flore

Générale de l Indo-Chine, vol. 6. Masson & Co., Paris.

Gao, J.-Y., Y.-M. Xia, J.-Y. Huang and Q.-J. Li. 2005. The Zingiberaceae of

China. Scientific Publisher, Beijing.

IUCN. 2001. IUCN Red List Categories and Criteria, Version 3.1. IUCN

Species Survival Commission. Gland, Switzerland and Cambridge, UK.

Kress, W.J., L.M. Prince and K.J. Williams. 2002. The phylogeny and a new

classification of the gingers (Zingiberaceae): evidence from molecular

data. American Journal of Botany 89 (11): 1682-1696.

Kress, W.J.,A-Z. Lui,M. Newman and Q.-J. Li.2005.The molecular phylogeny

of Alpinia (Zingiberaceae): a complex and polyphyletic genus of gingers.

American Journal of Botany 92 (1): 167-178.

Larsen, K., J.M. Lock, H. Maas and P.J.M. Maas. 1998. Zingiberaceae, pp.

474-495. In: Kubitzki, K. (ed.) The Families and Genera of Vascular Plant,

vol. 4. Springer-Verlag, Berlin.

Pham, H.H. 2003. Cay cé Viét Nam, An illustrated Flora of Vietnam. Vol. 3:

432-461. Youth Publication, Hochiminh City.

367

Siliquaamomum oreodoxa (Zingiberaceae): a New Species from Southern Vietnam

Smith, R.M. 1981. Synoptic keys to the genera of Zingiberaceae pro parte.

Royal Botanic Garden Edinburgh, Departmental Publication series 2:

1-28.

Wu, T.-L. and K. Larsen. 2000. Zingiberaceae. In: Wu, C.-Y. and Raven, P.H.

(eds.) Flora of China. Vol. 24: 322-377. Science Press, Beijing.

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Gardens’ Bulletin Singapore 61 (2): 369-378. 2010 369

Three New Species of Wrightia

(Apocynaceae: Apocynoideae) from Thailand

D. J. MIDDLETON

Royal Botanic Garden Edinburgh, 20A Inverleith Row, Edinburgh, EH4 1ET,

Scotland, U.K.

Abstract

Three new species of Wrightia from Thailand are described: Wrightia

karaketii D.J.Middleton, Wrightia tokiae D.J.Middleton and Wrightia

poomae D.J.Middleton.

Introduction

The genus Wrightia R.Br. is reasonably well known due to a revision of the

entire genus by Ngan (1965), coupled with more recent regional revisions

for China (Li et al., 1996), Thailand (Middleton, 1999), Malesia (Middleton,

2007a), Peninsular Malaysia (Middleton, 2010), and Cambodia, Laos and

Vietnam (Middleton, in press). This literature has meant that undescribed

species have been relatively easy to discern and several have been published

in recent years (Middleton and Santisuk, 2001; Middleton 2005a, 2007b). The

pattern that has emerged is of a number of widespread species, mostly not

growing on limestone, along with several local endemic species, mostly on

limestone. Many of the more recent discoveries are these limestone endemics

as collecting efforts in a number of countries, particularly in Thailand, have

intensified. A similar pattern of widespread non-limestone species and

fairly narrowly endemic limestone species can be seen in Alstonia R.Br.

(Sidiyasa, 1998; Middleton, 2005b). Recent collecting in northern Thailand

has brought to light three undescribed species of Wrightia, two of them from

limestone, the other from dry evergreen forest.

Ngan suggested that Wrightia could be divided into four sections.

Livshultz et al. (2007) sampled six species of the genus in their molecular

phylogeny of Apocynaceae subfamily Apocynoideae. The two species of

Wrightia sect. Scleranthera (Pichon) Ngan which were sampled, namely W.

dubia (Sims) Spreng. and W. coccinea (Roxb. ex Hornem.) Sims, are nested

within Wrightia sect. Wrightia. Wrightia sirikitiae D.J.Middleton & Santisuk,

which was not included in Ngan’s system (Ngan, 1965), but would key out to

Wrightia sect. Wallida A.DC., is also nested within Wrightia sect. Wrightia.

No species from Wrightia sect. Balfouria (R.Br.) Ngan were included. This

370 Gard. Bull. Singapore 61 (2) 2010

section, confined to Africa and Australia, was defined on the basis of the

corona lobes being coherent, only attached to the corolla at the very base and

forming a ring around the stamens. Of the three new species described here,

Wrightia tokiae would key out in Wrightia sect. Balfouria in Ngan (1965),

but, given the findings of Livshultz et al. (2007), the sectional arrangement

should be abandoned until further research is done.

Conservation assessments have been applied to each taxon using the

IUCN guidelines (IUCN, 2001; IUCN Standards and Petitions Working

Group, 2008).

Wrightia karaketii D.J.Middleton, sp. nov.

Corolla rotata. Corona annulum continuum ad corollam praeter fimbrias

marginales adnata. Ovaria apocarpa. Fructus bini.— Typus: Thailand, Chiang

Mai, Chiang Dao, Ban Arunothai, Kio Phawok, 13 May 2007, Pooma,

Karaket, Pattharahirantricin & Sirimongkol 6732 (holotype, BKF; isotype,

E). Figs. 1-3.

Tree 5-10 m tall, 15 cm dbh; bark pale brown, lenticellate; twigs puberulent

_ when young with short hooked hairs, soon glabrescent. Leaves opposite;

petiole 9-10 mm long, densly puberulent with short hooked hairs; blade

elliptic, 9-21 x 5.6-11.8 cm, 1.2-1.9 times as long as wide, apex emarginate-

apiculate to rounded-apiculate or shortly acuminate, base cuneate, mid

green above, slightly paler beneath, densely puberulent beneath, sparsely so

above but more densely so on venation, venation weakly brochidodromous,

15-19 pairs of secondary veins, these occasionally forking, tertiary venation

alternate percurrent. Inflorescence a short terminal cyme, few-flowered,

puberulent; peduncle 8-23 mm long; pedicels ca 4.5 mm long. Sepals ovate,

3.7-5.5 x 2.5-2.8 mm, apex obtuse, puberulent outside, greenish, with two

broad colleters per sepal at base inside, these bifid at the apex. Corolla bright

red, greenish at base outside, rotate; tube ca 2 mm long, minutely puberulent

outside; lobes overlapping to the left in bud, obovate, 17-20 x 12-15 mm, apex

rounded, papillose inside and outside, minutely puberulent at base outside.

Corona bright red, a continuous ring likely to be composed of antepetalous

and alternipetalous lobes, adnate to the corolla thereby joining the bases of

the corolla lobes together, 4-5 mm long opposite the corolla lobes, slightly

shorter between the corolla lobes where it is 2.2-3.2 mm long, glabrous,

margin shortly fimbriate, the fimbriae free from the corolla, fimbriae 1.2-

1.5 mm long. Stamens 5, attached in a ring to the style head, inserted at the

mouth of the corolla; filaments ca 1.5 mm long, puberulent; anthers yellow,

narrowly triangular, base sagittate,ca 6.5 x 2mm, shortly puberulent dorsally.

Gynoecium of two apocarpous ovaries united into a common style; ovaries

ca 2.5 mm high, glabrous; style + style head ca 6 mm long. Fruit of paired

ae: ! 1 371

Three New Species of Wrightia (Apocynaceae: Apocynoideae) from Thailand J

Figure 1. Distribution of Wrightia karaketii D.J.Middleton ( —). Wrightia tokiae D.J.Middleton

(4) and Wrightia poomae D.J.Middleton ( ® ).

Figure 2. Wrightia karaketii D.J.Middleton - flower (x1.5). Photo of Pooma et al. 6732 by

Rachun Pooma.

372 Gard. Bull. Singapore 61 (2) 2010

follicles, these pendulous and slighly diverging or occasionally twisting

around each other but still free, often rejoining at the tips, 29.5-40 cm long,

7-9 mm wide, brown, densely cream lenticellate, glabrous. Seeds linear, ca

17.5 x 2.2 mm, coma pointing towards base of fruit, ca 31 mm long.

Additional collection studied: THAILAND. Chiang Mai, Chiang Dao District,

Kio Phawok border checkpoint, 750 m altitude, 21 Sep 2008, Middleton,

Karaket, Triboun, Kawatkul & Meeboonya 4541 (BK, BKF, E, K).

Distribution: Only known from Kio Phawok in Thailand, very close to the

border so is also likely to occur in Burma.

Habitat. On karst limestone rocks in mixed deciduous forest at 750 m

altitude.

IUCN conservation assessment: DD. Under IUCN guidelines, as this species

is only known from the type locality and there is no information on possible

threats, the category of Data Deficient should be given (IUCN Standards

~ and Petitions Working Group, 2008).

Etymology: This species is named after Mr Preecha Karaket, one of the

collectors of both known specimens and an excellent photographer.

Notes: This species is highly distinctive with its corona in a ring almost

entirely adnate to the corolla and its apocarpous ovaries which give rise to

paired fruits. Its affinities are probably with Wrightia lanceolata Kerr and W.

siamensis D.J.Middleton. It differs from both species in its larger leaves, the

shorter corolla tube, the slightly longer corolla lobes and the more fimbriate

corona.

Wrightia tokiae D.J.Middleton, sp. nov.

Corolla rotata. Corona annulum continuum e corolla ad basin exceptum

libera. Ovaria apocarpa. Fructus bini.—Typus: Thailand, Tak, Maesot, Phawo

spirit house on the Tak-Maesot road, 700 m altitude, 24 May 2008, Pooma,

Karaket, Pattharahirantricin & Saengrit 6906 (holotype, BKF; isotypes, A,

AAU, BKF,E,K,L,SING). Figs. 1, 4, 5.

Tree 15 m tall; bark pale brown, with fine lenticels; twigs densely short

puberulent with short hooked hairs, eventually glabrescent. Leaves

opposite; petiole 4-6 mm long, puberulent; blade ovate to elliptic, 5.3-21 x

3.4-8 cm, 1.4-2.6 times as long as wide, apex acuminate, base rounded to

Three New Species of Wrightia (Apocynaceae: Apocynoideae) from Thailand

Figure 3. Wrightia karaketii D.J.Middleton - fruit («0.25). Photo of Middleton et al. 4541 by

Preecha Karaket.

Figure 4. Wrightia tokiae D.J.Middleton - flower (0.15). Photo of Pooma et al. 6906 by

Preecha Karaket.

374 Gard. Bull. Singapore 61 (2) 2010

acute, densely puberulent beneath, papillose above, sparsely puberulent on

midrib, venation brochidodromous, 13-18 pairs of secondary veins, tertiary

venation alternate percurrent. Inflorescence a terminal cyme, few-flowered,

densely puberulent; peduncle 6-10.5 mm long; pedicels 3-5 mm long; flowers

slightly fragrant. Sepals pale green, ovate, 3.5-4.2 x 2.4-4.8 mm, apex obtuse

to rounded, puberulent; colleters absent. Corolla yellowish green inside,

paler outside, fallen corolla reddish, rotate; tube 2.5 mm long, minutely

papillose outside; lobes overlapping to the left in bud, elliptic, ca 13 x 11

mm, apex rounded, margins reflexed, papillose inside, minutely so outside.

Corona yellow, a continuous cup-like ring, slightly narrower at the top than

in the middle, adnate to corolla only at very base, ca 5 mm long, glabrous

inside, minutely papillose outside, margin dentate. Stamens 5, attached in a

ring to the style head, inserted at the mouth of the corolla; filaments 1 mm

long, puberulent; anthers yellow, narrowly triangular, base sagittate, ca 7 x

1.8 mm, shortly puberulent dorsally. Gynoecium of two apocarpous ovaries

united into a common style; ovaries ca 1.5 mm high, glabrous; style + style

head ca 5.5 mm long. Fruit of two closely associated but not fused parallel

follicles joined at the apex, ca 13.5 cm long, each follicle ca 1.3 cm wide, dark

- brown, pale brown lenticellate except on inner surfaces. Seed not seen.

Additional collection studied: THAILAND. Tak, Maesot, behind Phawo

spirit house, 24 August 2008, Karaket 3 (BKF, E).

Distribution: Only known from the type locality.

Habitat: In dry evergreen forest on limestone hills.

IUCN conservation assessment: DD. Under IUCN guidelines, as this species

is only known from the type locality and there is no information on possible

threats, the category of Data Deficient should be given (IUCN Standards

and Petitions Working Group, 2008).

Etymology: This species is named after Ms Nannapat Pattharahirantricin,

more commonly known as Tok, one of the collectors of the type.

Note: This species would appear to fall into Wrightia sect. Balfouria in

the sectional system proposed by Ngan (1965). This section is otherwise

only known from Africa and Australia. However, as noted above, the

sectional arrangement in the genus is in need of revision. In flower it is

perhaps most similar to Wrightia coccinea, but that species does not have

its corona in a ring and the ovaries are syncarpous. There does not appear

to be any sepaline colleters in Wrightia tokiae, which, although similar in

Three New Species of Wrightia (Apocynaceae: Apocynoideae) from Thailand

E £ |

Figure 5. Wrightia tokiae D.J.Middleton - fruit (x0.6). Photo of Karaket 3 by Preecha

Karaket.

Figure 6. Wrightia poomae D.J.Middleton - flower (x1.5). Photo of Pooma 6973 by Preecha

Karaket.

376 Gard. Bull. Singapore 61 (2) 2010

Wrightia poomae, is highly unusual in the genus. The fruit of this species

appears to be syncarpous as in Wrightia coccinea, but the follicles, although

very closely associated (see Fig. 5) are not fused together.

Wrightia poomae D.J.Middleton, sp. nov.

Corolla rotata. Corona e lobis antepetalis alternipetalis et alternantibus

glabris composita. Ovaria apocarpa. Fructus ignoti. — Typus: Thailand, Tak,

Umphang, roadside from Huai Nam Khao Forest Protection Unit to summit

of Mae Chan Tha Forest Protection Unit, 700 m altitude, Pooma, Karaket,

Pattharahirantricin & Saengrit 6973 (holotype BKF; isotype A, AAU, BKF,

E, K, L, SING). Figs. 1, 6.

Tree ca 4 m tall; bark greyish-green; twigs sparsely puberulent when young,

soon glabrescent. Leaves opposite; petiole 2-5 mm long, sparsely puberulent;

blade ovate to elliptic, 2.3-12 x 1.5-5.6 cm, 1.8-2.1 times as long as wide,

apex acuminate to subcaudate, base obtuse to acute, dull green above,

pale green beneath, minutely papillose and very sparsely puberulent on

. venation above and beneath, venation weakly brochidodromous, 8-12 pairs

of secondary veins, tertiary venation alternate percurrent. Inflorescence a

terminal cyme, few-flowered, sparsely puberulent; peduncle 4-10 mm long;

pedicels 2.5-7 mm long; flowers fragrant. Sepals pale green, ovate, 4.2-4.5

x 5.8-6.4 mm, apex rounded, sparsely puberulent, ciliate; colleters absent.

Corolla pale yellowish-orange, turning pale red with age, rotate; tube ca 2.7

mm long, minutely papillose outside; lobes overlapping to the left in bud,

elliptic, ca 19 x 13 mm, apex truncate, margins somewhat reflexed, minutely

papillose inside, barely so outside. Corona pale orange, of three distinct

parts: antepetalous, alternipetalous and alternating lobes, the antepetalous

and alternipetalous lobes glabrous inside, minutely papillose outside, the

alternating lobes glabrous; antepetalous lobes ca 6.5 mm long, ca 1/3 width

of and adnate to corolla lobes for approximately half their length, fimbriate

to 1/3 of their length; alternipetalous lobes in same plane as antepetalous,

adnate to antepetalous lobes at base, ca 5 mm long, bifid; alternating lobes

between the antepetalous and alternipetalous at the very base (but somewhat

irregular in number up to 10), simple, ca 1.7 mm long. Stamens 5, attached

in a ring to the style head, inserted at the mouth of the corolla; filaments ca

1.5 mm long, puberulent; anthers yellow, narrowly triangular, base sagittate,

ca 9 x 2.5 mm, shortly puberulent dorsally. Gynoecium of two apocarpous

ovaries united into a common style; ovaries ca 2.2 mm high, glabrous; style

+ style head ca 7 mm long. Fruit and seed not known.

Distribution: Only known from the type collection.

Three New Species of Wrightia (Apocynaceae: Apocynoideae) from Thailand 377

Habitat: In dry evergreen forest at 700 m altitude.

IUCN conservation assessment: DD. Under IUCN guidelines, as this species

is only known from the type locality and there is no information on possible

threats, the category of Data Deficient should be given (IUCN Standards

and Petitions Working Group, 2008).

Etymology: This species is named after Dr Rachun Pooma, one of the

collectors of the type.

Note: This species appears superficially similar to Wrightia pubescens

R.Br., which is common in Thailand, but that species normally has only

two whorls of corona lobes, the corona is pubescent inside and the ovaries

are syncarpous. It is most similar to Wrightia kwantungensis Tsiang from

China and northern Vietnam, which also has three whorls of corona, but

that species has smaller sepals with an acute apex and colleters at the base

inside, and a much longer corolla tube. There does not appear to be any

sepaline colleters in Wrightia poomae, which, although similar in Wrightia

tokiae, is highly unusual in the genus.

Acknowledgements

My most grateful thanks go to Rachun Pooma, Nannapat Pattharahirantricin

and Preecha Karaket for bringing these plants to my attention, and, along

with them, to U. Kawatkul, R. Meeboonya S. Sirimongkol, S. Saengrit and P.

Triboun for collecting the material of these new taxa. In addition, Rachun

Pooma provided valuable advice on the manuscript. I should also like to

thank R. Mill for the Latin descriptions.

References

IUCN. 2001. IUCN Red List Categories and Criteria: Version 3.1. 'UCN

Species Survival Commission. IUCN, Gland, Switzerland and Cambridge

IUCN Standards and Petitions Working Group. 2008. Guidelines for Using

the IUCN Red List Categories and Criteria. Version 7.0. Prepared by the

Standards and Petitions Working Group of the IUCN SSC Biodiversity

Assessments Sub-Committee in August 2008.

Li, P.T., A.J.M. Leeuwenberg and D.J. Middleton. 1996. Apocynaceae in

Flora of China 16: 143-188.

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Livshultz, T., D.J. Middleton, M.E. Endress, and J. Williams. 2007. Phylogeny

of Apocynoideae and the APSA clade. Annals of Missouri Botanical

Garden 94: 323-361

Middleton, D.J. 1999. Apocynaceae. Flora of Thailand 7(A): 1-152.

Middleton, D.J. 2005a. A revision of Wrightia (Apocynaceae: Apocynoideae)

in Malesia. Harvard Papers in Botany 10: 161-182.

Middleton, DJ. 2005b. A new species of Alstonia (Apocynoideae:

Rauvolfioideae) from Vietnam. Harvard Papers in Botany 10: 63-65.

Middleton, D.J. 2007a. Apocynaceae, subfamilies Rauvolfioideae and

Apocynoideae. Flora Malesiana 18: 1-471.

Middleton, D.J. 2007b. A new species of Wrightia (Apocynaceae:

Apocynoideae) from Thailand. Thai Forest Bulletin 35: 80-85.

- Middleton, D.J. 2010. Apocynaceae, subfamilies Rauvolfioideae and

Apocynoideae. Flora of Peninsular Malaysia, in press.

Middleton, D.J. (in press). Apocynaceae, subfamilies Rauvolfioideae and

Apocynoideae. In: Flore du Cambodge, du Laos et du Vietnam, in press.

Middleton, D.J. and T. Santisuk. 2001. A new species of Wrightia

(Apocynaceae: Apocynoideae) from Thailand. Thai Forest Bulletin 29:

1-10.

Ngan, P. T. (1965). A revision of the genus Wrightia (Apocynaceae). Annals

of Missouri Botanical Garden 52: 114-175.

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(Apocynaceae). Blumea Supplement 11: 1-230.

Gardens’ Bulletin Singapore 61 (2): 379-388. 2010 379

Two Basidiomycetes New to Indonesia, Prerygellus

armeniacus and Rimbachia leucobryi

O. MIETTINEN’ AND N. HERNAWATI-

Finnish Museum of Natural History, Botanical Museum (H)

Box 7, FI-00014 University of Helsinki, Finland

‘ Email: otto.miettinen@helsinki.fi

~The Herbarium (ANDA), Universitas Andalas

Padang, Sumatra Barat, Indonesia 25163

Abstract

Rimbachia leucobryi, a small pleurotoid agaric, is described as new to

science. The species grew on living stems of Leucobryum sanctum in lowland

rainforest. It is characterised by lamellate hymenophore, hymenial cystidia

and small, ellipsoid spores. Prerygellus armeniacus, a bright-coloured

stipitate stereoid species, is reported as new to Indonesia. The two species

are illustrated.

Introduction

Tropical Southeast Asian mycota is one of the most poorly known globally.

Only a few mycologists work in the area while deforestation rates in the

mycologically most diverse rainforest areas hit all time high. To answer the

call to catalogue the vanishing mycological diversity, Herbarium of Andalas

University (ANDA, Padang, Indonesia) and Botanical Museum, University

of Helsinki (H, Finland) have established cooperation, during which

basidiomycete diversity in Sumatra is recorded and described. From the

collections made so far, two minute species new to Indonesia are described

and reported below.

Materials and methods

The basic mounting medium used in microscopic studies was Cotton Blue

(CB), but also Melzer’s reagent (IKI) and 5% KOH. Spore and other

measurements were made and illustrations were drawn in CB. Entry CB+

means cyanophily, CB(+) weak but distinct cyanophilous reaction, CB-

acyanophily; IKI— means neither amyloid nor dextrinoid reaction; KOH-—

means that hyphae were left almost unchanged. Measurements were done

using x1000 magnification and phase contrast illumination; eyepiece scale

380 Gard. Bull. Singapore 61 (2) 2010

bar with l-um-grid was used, and dimensions were estimated subjectively

with an accuracy of 0.1 um (see Miettinen et al., 2006 for further detail).

The following symbols are used for spore measurements: L = mean

length, W = mean width, Q = L/W, i.e. average length divided by average

width, Q’ = length/width ratio of individual spores, n = number of spores

measured from given number of specimens, for instance 90/3 means 90

spores measured from 3 specimens. In presenting the variation of spore size

and Q’, the whole range is given in parentheses. The 90% range excluding

the extreme 5% of values from both ends is given without parentheses. In

case the 5% limit falls between two values, the one further from the median

was chosen to represent the 5% tail. Whenever the figures within and outside

parentheses are identical, parentheses are omitted.

Species descriptions

Pterygellus armeniacus Corner

A monograph of Cantharelloid fungi (1966) 168.

Basidiocarp centrally to more rarely laterally stipitate, cap irregular, venose,

~ upwards directed, often divided from the base to several separate spathulate

lobes, particularly when young, stipe 10-15 mm long, cap 5 mm in diameter, 0.2

mm thick. Stipe and upper surface covered with rough, upward-pointing tufts,

brownish orange, hymenium paler than rest of the basidiocarp, consistently

tough but brittle when dry. Hymenial surface consists of branching, blunt

veins. Cap margin irregularly sharp with saw-like pattern, following vein

patterns of the cap. Hyphal system monomitic. Hyphae homogenous

throughout the basidiocarp, slightly thick-walled, clamps absent, CB— but

inclusion stained in CB, IKI— but cells with golden yellow content, (2.8-) 3.2-

4.3 (-6) um in cap trama, (3.2-) 4-5.4(-7) um in stipe context. Hyphal tufts

of the upper and stipe surface formed by parallel, tightly arranged hyphae,

similar to normal generative hyphae albeit narrower, (2.8-) 3-4.3 (-4.7) um

(measured from the stipe). Hyphae forming the cap surface not different

from the tramal hyphae, but slightly projecting, blunt, occasionally slightly

swollen hyphal ends present. Cystidia absent. Hymenium. Basidia with a

tapering base, narrowly clavate, occasionally sinuous, often with visible,

granular inclusion, 25-65 x 6.5-8.6 um, with (2-)3-5 sterigmata, length 4-7

um. Basidiospores broadly ellipsoid to subglobose, CB-, IKI-, slightly thick-

walled, smooth, 7.5-9.2 (-9.8) x (5.9-) 6.2-7.8 (-7.9) um, L=8.50 um, W=6.92

um, Q’=1.1-1.3(-1.5), Q=1.23, n=40, apiculus prominent, about 0.8 x 0.8 um.

Inclusion granular, without guttules. In some slightly dehydrated spores

the spore wall seems to have minute ornamentation in the form of small

granules, not unlike the inclusion. Figs. 1-3.

= 2 . x J

Two Basidiomycetes New to Indonesia, Pterygellus armeniacus and Rimbachia leucobryi 381

Specimens studied: INDONESIA. Sumatera Barat, Padang, Limau Manis,

Andalas University Biology research forest, S 0° 54.61, E 100° 28.36’,

alt. 430 m, on the ground in a steep slope of natural forest dominated by

Dipterocarpaceae and Fagaceae, 13.VII.2008, O. Miettinen 13004, field no

NOM292 (ANDA, H).

Habitat and ecology: The species grew on the ground without any evident

connection to decomposing substrate. It is probably ectomycorrhizal,

although a root-rotting habit can not be ruled out. Ectomycorrhizal fungi

are not uncommon in this forest type dominated with two well-known

ectomycorrhizal tree families, Dipterocarpaceae and Fagaceae. Prerygellus

is traditionally included in Cantharellales, which includes numerous

ectomycorrhizal fungi. Verbeken and Walleyn (1999) report Prerygellus

polymorphus to be ectomycorrhizal with Euphorbiaceae.

Distribution: The type locality of this species is in Singapore, and the new

find extends the distribution to Sumatra.

Notes: Our specimen fits well with Corner’s (1966) description with the

exception of the number of sterigmata in each basidium, (3-) 4-6 according to

Corner. We studied over 60 basidia, and could not spot any 6-steritmate (Fig.

3). Peculiarly 5-sterigmate basidia were almost as common as 4-sterigmate,

and 3-sterigmate were not rare either.

Rimbachia leucobryi Miettinen, sp. nov.

Basidioma pileatum, lamellatum, estipitatum, 2-5 mm latum, muscicola.

Systema hypharum monomiticum, hyphis fibulatis, inflatis, cystidiis

hymenialibus subulatis, basidiosporae ellipsoideae, 5-6 x 3.8-4.8 um. —Typus:

INDONESIA, Riau, Kabupaten Kampar, Balung, Teratak Baru, N 0° 8.6’,

E 100° 49.4’, alt. 170-250 m, on living stems of Leucobryum sanctum (Brid.)

Hampe growing on a large, fallen log in a logged-over primary rain forest,

24.XI11.2006, O. Miettinen 11267 (holotype, ANDA; isotypes, H, SING).

Figs. 4-5.

Basidiocarp pileate, laterally attached or slightly pendant, without stipe,

pure white when fresh, drying cream-coloured, 2-5 mm wide when fresh,

1-2 mm when dry, often curving inwards upon drying. Consistency rather

fragile. Hymenophore consists of forked lamellae or folds, same colour

as the upper surface. Hyphal system monomitic. Hyphae thin- to slightly

thick-walled, clamps present in nearly all septa, CB-, IKI-. Tramal (and

contextual) tissue relatively dense, hyphae inflated, subparallel, (3-)3.8-6.5(-

15) um in diameter. Hyphae at upper surface not differentiated. Sparse,

382 Gard. Bull. Singapore 61 (2) 2010

loose, inconspicuous hyaline, shiny granules or encrustation present in

all parts of the basidiocarp in CB; in IKI scanty golden encrustation seen

along hyphae. Cystidia thin-walled, projecting above hymenium, subulate,

occasionally branched, arising from subhymenium just as other hymenial

cells, 20-44 x 4.4-5.8um. Hymenium. Basidia cylindrical to clavate, with (2-)

4 sterigmata, 18-23.5 x 5-6.8 um, sterigmata up to 3.5 um long. Basidiospores

ellipsoid to subglobose, often pip-shaped, thin- to very slightly thick-walled,

smooth, CB-, IKI-, (4.8-) 5-6(-6.1) x 3.8-4.8(-5.0), L=5.36, W=4.28, Q’=1.1-

1.4, Q=1.25, n=50, with a prominent apiculus, typically 1.5 x 1.5 um.

Habitat and ecology: Evidently the species has a close connection to its

bryophyte host. Whether the relationship is parasitic, saprobic or symbiotic,

we can not judge at present. All the host stems and the moss colony in

general seem to be in good condition.

Distribution: The only find comes from lowland rainforest in central Sumatra.

The host species, L. sanctum, has a wide distribution in tropical Asia from

India and China to Papua New Guinea and is found throughout this area in

natural forests, being common in at least insular Southeast Asia (Enroth, 1990;

Yamaguchi, 1993). Other similar moss-inhabiting fungi are not species specific.

Thus the species could potentially be found anywhere in tropical Asia.

Etymology: Named after the host plant species, Leucobryum sanctum

Notes: In search of the name for the present species, all species validly

combined in the genera Arrhenia, Cheimonophyllum, Cyphallostereum,

Leptoglossum, Mniopetalum, Pleuromycenula, Pseudocratarellus, Pterygellus,

Rhodoarrhenia, and Rimbachia were compared using Mycobank (2009) and

literature. The most suitable genus according to the key of Singer (1986)

seems to be either Pleuwromycenula or Mniopetalum, both included by

Redhead (1984) in Rimbachia.

The current species fits best in Mniopetalum or Rimbachia being

astipitate, pleurally attached, small, pale-coloured, moss-inhabiting and

gilled, with clamped, inflated tramal hyphae, inamyloid, hyaline spores and

lacking differentiated pileipellis. The hymenial hyphoid cystidia are the only

deviating character.

DNA-based analysis would be needed to sort out relationships

between the minute-sized agarics that lack clear spore and _ cystidial

characters. No DNA sequence data exist currently in Genbank for the

type - or any other - species of Rimbachia, Flabellimycena, Mniopetalum, or

Pleuromycenula. Redhead et al. (2002a, b) did not include any members of

these genera in their DNA-based phylogenetic analysis. For these reasons

Two Basidiomycetes New to Indonesia, Pterygellus armeniacus and Rimbachia leucobryi 383

we follow Redhead’s (1984) broad concept of Rimbachia here, inclusive of

Mniopetalum.

The genus Cheimonophyllum includes small pleurotoid species with

cheilocystidia warranting generic comparison. The three species currently

included in the genus are all lignicolous. The type species, Cheimonophyllum

candidissimum (Berk. & M.A. Curtis) Singer, has non-inflated hyphae of

rather small diameter in contrast to inflated hyphae in Rimbachia (based

on literature and studied Finnish material in H). The cystidia in the type

species are long, branching and restricted to the gill edge, and not simple

and hymenial (pleurocystidia) as in R. lewcobryi. Considering the ecology,

hyphal structure and even cystidial characters, Rimbachia seems a more

natural placement for the new species.

At the species level, Rimbachia bryophila (Pers.) Redhead is

externally and ecologically very similar to R. Jewcobryi. However, according

to Kuyper (1995) and Senn-Irlet and Moreau (2003) based on European

material, its spores are clearly larger, 6-8.5 x 5-6.5 um. Redhead (1984:878)

reported slightly smaller spores for American material, 5-7 x 4.5-7 um,

but these are still larger than those of R. leucobryi. Same applies to

Leptoglossum subbryophilum Singer (spores 6.5-8.5 x 5.5-8 um, Singer, 1931:

521), Mniopetalum flabelliforme M. Zang and Mniopetalum miniatum M.

Zang (spores 7-10 x 5-6 um for both, see Zang, 1986).

Rimbachia furfuracea (Petch) Redhead and Mniopetalum distinctum

Horak (considered a synonym of the first-mentioned by Redhead, 1984)

are also quite similar. We rely on the detailed descriptions of Corner (1966:

76), Horak (1980: 100), and Pegler (1986: 95). Basidiocarps of Rimbachia

furfuracea are spathulate and bear a short lateral stipe; the basidiocarps

are terrestrial or growing on mosses, 5-25 mm in size, the gills are thick,

blunt, shallow and frequently anastomosing. The spore size is 5-6.5 x 3.2-4,

L=6, W=3.5 (Pegler, 1986), or 4.5-6 x 3-4 (Horak, 1980, for Mniopetalum

distinctum), hyphae 3-15 um wide and inflated. The microscopic characters

agree with R. /eucobryi, except that no cystidia are mentioned. The ecology

and macroscopic characters (shallow gills and larger basidiocarps, stipe) are

clearly different from R. leucobryi.

Rimbachia cyphelloides Lloyd (=Rhodoarrhenia cyphelloides (Lloyd)

Singer) from Brazil has a reticulate hymenium and a short stipe. Its spores

are also larger, 8 x 6 um (Lloyd 1918: 802) or 5.5 - 7-4 x 3.5-5.4 um (Singer,

1963: 143). Rimbachia ellipsoidea (Singer) Redhead from Mexico produces

small, white basidiocarps with a venose hymenium, growing on mosses. The

original description (Singer, 1973: 27) states that the spores measure 6.2-7.5

x 3.5-4.5 um, clearly longer than in R. /eucobryi. Redhead (1984) reports

that the type is in bad condition. None of the above-mentioned species is

reported to bear hymenial cystidia as does R. leucobryi.

9QQ

384 Gard. Bull. Singapore 61 (2) 2010

Figure 1. Pterygellus armeniacus (Miettinen 13004), photographed when fresh.

Two Basidiomycetes New to Indonesia, Pterygellus armeniacus and Rimbachia leucobryi 385

5 pm

10 um

Figure 2. Pterygellus armeniacus (Miettinen 13004). Normal spores (a), a dehydrated,

“ornamented” spore (b), and hymenium (c).

op]

2 2 4 5

sterigmata/basidium

Figure 3. Number of stergimata per basidium in Prerygellus armeniacus (based on Miettinen

13004). Total number of basidia included is 61.

386 Gard. Bull. Singapore 61 (2) 2010

Figure 4. Rimbachia leucobryi, based on fresh type material.

+ O0d00GHG

OS NY:

Figure 5. Rimbachia leucobryi, holotype: spores (a) and hymenial cells (b).

Two Basidiomycetes New to Indonesia, Pterygellus armeniacus and Rimbachia leucobryi 387

Acknowledgements

We express our gratitude to Dr. Arbinis Arbain, Prof. Dr. Syamsurdi, Dr.

Rusdji Tamin, all our friends at the Herbarium of Andalas University, and

Prof. P. Uotila (H) for providing generous assistance to this project. We also

thank Dr. J. Enroth (H) for identifying the bryophytes, Prof. T. Ahti (H) for

revising the Latin description, and Dr. T. Niemela (H) for his comments on

the manuscript.

References

Corner E.J.H. 1966. Monograp of cantharelloid fungi. Annals of Botany

Memoir 2: 1-255, + 5 pls.

Enroth, J. 1990. Bryophyte flora of the Huon Peninsula, Papua New Guinea

36. Leucobryaceae (Musci). Acta Botanica Fennica 139: 65-120.

Horak E. 1980. Indian Boletales and Agaricales revisions and new taxa.

Sydowia 33: 88-110.

Kuyper, T.W. 1995. Rimbachia, pp. 134-135. In: Bas, C., Kyper, T.W.,

Noordeloos, M.E. & Vellinga, E.C. (eds). Flora Agaricina Neerlandica 3.

Tricholomataceae.

Lloyd, C.G. 1918. Mycological notes 56: 798-812.

Mycobank. 2009. http://www.mycobank.org/mycotaxo.aspx [23 March 2009]

Miettinen, O.,T. Niemela and W. Spirin. 2006. Northern Antrodiella species:

the identity of A. semisupina, and type studies of related taxa. Mycotaxon

96: 211-239.

Pegler, D.N. 1986. Agaric flora of Sri Lanka. Kew Bulletin, Additional Series

12: 1-519.

Redhead, S.A. 1984. Arrhenia and Rimbachia expanded generic

concept, and reevaluation of Léptoglossum with emphasis on

muscicolous Noth American taxa. Canadian Journal of Botany 62:

865-892

388 Gard. Bull. Singapore 61 (2) 2010

Redhead, S.A., F. Lutzoni, JM. Moncalvo and R. Vilgalys. 2002a. Phylogeny

of agarics: partial systematics solutions for bryophilous omphalinoid

agarics outside of the Agaricales (euagarics). Mycotaxon 82: 151-168.

Redhead, S.A., F. Lutzoni, JM. Moncalvo and R. Vilgalys. 2002b. Phylogeny

of agarics: partial systematics solutions of core omphalinoid genera in

the Agaricales (euagarics). Mycotaxon 83: 19-57.

Senn-Irlet, B. and P.-A. Moreau. 2003. Notes on three Rimbachia species

from the Alps. Czech Mycology 54: 145-154.

Singer, R. 1931. Pilze aus dem Kaukasus 2. Ein Beitrag zur Flora Swanetiens

und einiger angrenzender Taler. Beihefte zum Botanischen Centralblatt

48: 513-542.

Singer, R. 1963. New genera of fungi 13. Rhodoarrhenia. Sydowia 17: 142-

145.

- Singer, R 1973. Diagnoses fungorum novorum Agaricalium 3. Beihefte zur

Sydowia 7: 1-106.

Singer, R. 1986. The Agaricales in modern taxonomy, 4th revised edition.

Koenigstein, Koelz Scientific Books. 981 pp + 88 pls.

Verbeken, A. and N. Walleyn. 1999. Is Pterygellus mycorrhizal with a

euphorbia? Mycologist 13: 37.

Yamaguchi, T. 1993. A revision of the genus Leucobryum in Asia. Journal of

Hattori Botanical Laboratory 73:\-123.

Zang, M. 1986. New or interesting species of the genus Mniopetalum from

the Eastern Himalayas. Mycotaxon 26: 297-307.

Gardens’ Bulletin Singapore 61 (2): 389-400. 2010 389

Ten New Records of Mosses from Doi Inthanon National

Park in Thailand

Y. NATHI ,.B.C. TAN AND T.SEELANAN |

Plants of Thailand Research Unit. Department of Botany, Faculty of Science.

, Chulalongkorn University, Bangkok 10330, Thailand

- The Herbarium, Singapore Botanic Gardens, | Cluny Road, Singapore 259569:

also Affiliate staff at Department of Biological Sciences,

National University of Singapore Singapore 119260

Abstract

Ten species of mosses collected from Doi Inthanon National Park are

reported newly for the flora of Thailand. Of these, Rhizomnium and

Oligotrichum are two new moss generic records for the country. The report

includes notes on ecology, morphology, taxonomy, and distribution of the

new species records.

Introduction

Thailand is located centrally in continental SE Asia. The country encompasses

a total land area of 513,115 km . The elevation ranges from sea level to 2,565

m (Doi Inthanon). Because of its geographical position, the flora is rich in

temperate Himalayan and Chinese elements in the north, and in tropical

Malesian moss taxa to the south.

The moss flora of Thailand has been studied intermittently since the

first westerner, J. Schmidt, collected moss specimens from Koh Chang in

1899 and 1900 (see Brotherus, 1901). Dixon (1932) published the first moss

checklist for Thailand based on the large collections of A.F.G. Kerr. When

Tixier (1971) published a summary of moss taxa for Thailand, the flora

consisted of 500 species. In his paper, Tixier analyzed the floristic affinity

of the moss flora of Thailand, which showed nearly an equal percentage

of species sharing with the Indian subcontinent, Indochina and Malesia.

From 1950 on, bryological activities in Thailand have intensified with

the participation and publication of bryologists from The Netherlands (see

Touw, 1968), France (Tixier, 1971, 1972), Japan (Horikawa and Ando, 1964),

Singapore (Tan and Iwatsuki, 1993, Tan and Tran Ninh, 1998; Tan et al., 2006),

USA (He, 1998), and Denmark (Larsen, 1979, 1992). A recent listing of

the bryoflora prepared by Sornsamran and Thaithong (1995) reported 644

species of mosses based on publications from 1900-1979 (see Tan, 1998).

390 Gard. Bull. Singapore 61 (2) 2010

More recent researches on Thai mosses include a number of local workers,

such as, Chantanaorrapint ef al. (2004), Koronochalert (2006), Manachit

(2006), and Wongkuna er al. (2009). Finally, an updated checklist of Thai

mosses prepared electronically and housed at Missouri Botanical Garden

(see http://www.mobot.org/MOBOT/moss/Thailand/welcome.shtml) listed

620 species and 31 subspecific taxa in 190 genera and 52 families.

In this paper, we report 10 new moss records for the Thai flora

collected from Doi Inthanon National Park, Chiang Mai Province. All the

new records are collected from montane evergreen forests at an elevation

above 2000 m. Of these, Rhizomnium and Oligotrichum are two new generic

records for the country’s moss flora. The voucher specimens are deposited,

as indicated separately under each species entry, at the Herbarium of

Chualongkorn University (BCU) and the Herbarium at Singapore Botanic

Gardens (SING).

New Records of Thai Moss Flora

Family Fissidentaceae

_ 1. Fissidens obscurus Mitt. Fig. 1.

This is a medium sized plant that is dark green in color. It was

collected on wet rock and sandy soil. Fissidens obscurus is similar to

Fissidens polypodioides Hedw., but easily recognize by having abundant

tomentose rhizoids on the underside of stems. The lack of hyaline nodule on

the stem, the smooth leaf cells, the absence of leaf limbidium and the obtuse

to obtusely acute leaf apex, in combination, identify this species.

Fissidens obscurus has a widely scattered distribution in China, Japan,

Nepal, India, and now in northern Thailand.

Specimens studied: Y. Nathi 914, 1015, 1024 (BCU).

Family Hypnaceae

2. Glossadelphus prostratus (Dozy & Molk.) M.Flesich.

Syn. Taxiphyllum prostratum (Dozy & Molk.) W.R.Buck

The plants were collected from Kew Mae Pan on Doi Inthanon,

forming a mat of intertwined, fine and elongate branches. Leaves

are small, about 0.25-0.4 mm long, varying from ovate with obtuse

apex on branchlets and near the distal ends of branches, to ovate

with short acuminate apex on primary and secondary branches. Leaf

margins are serrate and the elongate laminal cells are often prorulose.

Glossadelphus prostratus is known already from China, Laos,

Vietnam, Indonesia (Java, Lombok, Irian Jaya), Papua New Guinea

and several Pacific Islands. It is not surprising to find it in Thailand.

Specimen studied: Y. Nathi 945 (BCU, SING).

Ten New Records of Mosses from Doi Inthanon National Park in Thailand 391

Figure 1. Fissidens obscurus. a-c. Leaves; d. Cells at leaf margin; e- f. Cross section of leaf.

Family Meteoriaceae

3. Meteorium subpolytrichum (Besch.) Broth.

This epiphytic moss species is large, dark-green and densely branched.

Leaves are imbricate to appressed on branches, but the overall morphology

of the branch foliation is not as strongly julaceous as in Meteorium buchananii

(Brid.) Broth. The ovate-oblong to ligulate leaves have rounded to truncate

apices, auriculate and undulate bases, and crenulate leaf margins; costae

reach ca 2/3 of the leaf length.

Meteorium subpolytrichum is distributed in the Himalayas, China,

Japan, the Philippines and is here reported from northern Thailand.

Specimens studied: Y. Nathi 137, 262 (BCU).

Family Mniaceae

4. Rhizomnium striatulum (Mitt.) T.J.Kop. — Fig. 2.

The two specimens of Rhizomnium striatulum were found along a

shaded stream, growing on rocks on Doi Inthanon. The plants are small

and their stems are dark to reddish, 0.8-1.0 cm long. Leaves are elliptic to

obovate in outline, slightly contorted when dry and widely spreading when

moist. Leaf borders are entire, strong and dark brown in color, comprising

392 Gard. Bull. Singapore 61 (2) 2010

of 2-3 rows of linear and elongate-rectangular cells. The leaf tip ends in an

apiculus and the costa ends near the leaf apex.

Rhizomnium striatulum is a new species record and also a new generic

record for the flora of Thailand. Distribution of this species is from Eastern

Russia, Korea, Japan, China, Taiwan, Himalayas to northern Thailand.

Specimens studied: Y. Nathi 1016, 1051 (BCU).

4 ry

A TSS

Se :

Figure 2. Rhizomnium striatulum. a. Plant habit; b -c. Leaves; d. Leaf tip; e. Leaf cells.

Family Polytrichaceae

5. Oligotrichum aligerum Mitt. Fig. 3.

This is a moss collected from the roadside on way to the summit of

Doi Inthanon. The specimens were found growing mixed with species of

Pogonatum. The plants are small and the stems are rigid and short, about

0.5-0.8 cm high. The leaves are oblong to oval in shape, covered with slightly

wavy rows of ventral and dorsal lamellae, each lamella is 4-8 cells high. The

Ten New Records of Mosses from Doi Inthanon National Park in Thailand 393

broadly acute to acute leaf apices and the presence of several low dorsal

lamellae identify this species from the other members of the genus in

continental SE Asia. Oligotrichum aligerum is a new generic and species

record for Thailand.

Distribution of Oligotrichum aligerum ranges from North and

Central Americas, Russian Siberia, Japan, Korea, China, Taiwan, northern

Philippines, Nepal, India to northern Thailand.

Specimens studied: Y. Nathi 29, 196, 955 (BCU).

Figure 3. Oligotrichum aligerum. a-c. Plant habits; d-f. Leaves; h. Leaf cells; i. Leaf cell at

base; j-k. Cross section of leaves.

Family Pottiaceae

6. Didymodon maschalogena (Renauld & Cardot) Broth. Fig. 4.

Syn. Didymodon michiganensis (Steere) K.Saito

Didymodon maschalogena grows along the roadside on Doi Inthanon

and is easy to recognize because of the presence of abundant, uniquely dark

and round gemmae in leaf axils. The plants are small, about 1-1.5 cm high.

394 Gard. Bull. Singapore 61 (2) 2010

Leaves are dense and close to stem when dry, but spread out when wet. Leaf

shape is characteristically ovate and abruptly acuminate at apex. Its leaf

margins are narrowly recurved in the lower part. Laminal cells are mostly

round to hexagonal, slightly bulging and smooth, and arranged in observable

longitudinal rows.

Didymodon maschalogena is known in North America as D.

michiganensis. (Frahm et al., 1996; Jiménez et al., 2004). It is also known

from Africa, Mexico, Japan, India, Sri Lanka, Himalayas, China and the

Philippines. Its presence in Thailand represents a local range extension. A

worldwide distribution map of this species was presented by Jiménez et al.

(2004).

Specimens studied: Y. Nathi 49, 411, 577, 986 (BCU).

(>)

O80

Figure 4. Didymodon maschalogena. a. Plant habit; b-d. Leaves; e. Leaf cells; f. Leaf tip; g.

Gemmae

oS)

Ten New Records of Mosses from Doi Inthanon National Park in Thailand

Family Sematophyllaceae

7. Clastobryopsis brevinervis M.Fleisch.

This is an epiphyte on branches in montane mossy forest near summit

of Doi Inthanon. The plant is small, about 1 cm long. It is recognized by its

single costa-like leaf appearance. Actually one of the two costae is short. As

a typical member of the genus, the leaf base is decurrent and the alar cells

are made up of a mixture of quadrate and rectangular, more or less thick-

walled cells. Leaves of this species are lanceolate, acuminate, plicate, and the

leaf margins are somewhat revolute.

Clastobryopsis brevinervis is distributed in China, Japan, Indonesia

(Java, lesser Sunda), Malaysia (Sabah), the Philippines and Papua New

Guinea. It is new to Thailand.

Specimens studied: ¥. Nathi 150, 627, 1071 (BCU).

8a. Clastobryopsis planula (Mitt.) M.Fleisch. var.planula Fig. 5.

This mat-forming species is commonly found in open sites of forest near

the summit of Doi Inthanon. It grows on trunk and branches of trees. Stems

are densely branched and produce an enlarged and complanate terminal

with many propaguliferous gemmae found inside the leaf axil. Leaves are

broadly ovate and ovate-lanceolate, double-costate, with acuminate tip, and

the leaf margins are slightly reflexed. Leaf bases are decurrent and the alar

cells are quadrate to rectangular in shape, often colored.

Clastobryopsis planula is known from China, Japan, India (Sikkim),

Nepal, Indonesia, the Philippines, and is new to Thailand.

Specimens studied: Y. Nathi 258, 328, 486, 838, 924, 959 (BCU).

8b. Clastobryopsis planula var. delicata (M.Fleisch.) B.C.Tan & Y_Jia

Although treated as a synonym of Clastobryopsis planula by many

workers (cf. Tan and Jia, 1999), in Doi Inthanon National Park, this variety

is distinctly small and more delicate in appearance. Its leaf outline is also

narrower than the typical variety. It is also an epiphyte found on branches

in upper montane forest.

Specimens studied: Y. Nathi 55, 349, 525, 653, 664 (BCU).

9. Clastobryopsis robusta (Broth.) M.Fleisch.

This rather widespread tropical Malesian species grown on branches

of a tree at Doi Inthanon and appears to be rare in occurrence locally. The

two specimens represent undoubtedly juvenile plants. Stems are creeping

and irregularly branched. Stem leaves are ovate-lanceolate to oblong-

lanceolate in shape. The broadly ovate and somewhat plicate branch leaves,

coupled with acuminate apex, broadly decurrent leaf bases, and narrowly

recurved margins with teeth above, identify this species from its congeners.

396 Gard. Bull. Singapore 61 (2) 2010

Figure 5. Clastobryopsis planula. a. Propaguliferus branch; b. Branch; c-d. Propaguliferus

leaves; e-h. Branch leaves; i. Alar cells.

The leaf alar cells, like all members of the genus, are typically numerous,

brownish, rectangular to quadrate in shape, and not inflated.

Clastobryopsis robusta is distributed in Japan, China (Taiwan),

Philippines, Borneo, Java and Papua New Guinea. It is newly found in

Thailand.

Specimens studied: Y. Nathi 627, 1072 (BCU).

10. Warburgiella bistrumosa (Miill.Hal.) M.Fleisch. Fig. 6.

The spcimens were collect from Kew Mae Pan Nature Trail at Doi

Inthanon National Park. The small and slender plants are about 0.5 cm

long, and irregularly branched. Leaves are narrowly lanceolate, gradually

long acuminate, concave and mostly falcate. Leaf cells are smooth at base

and develop a papilla on lumen of many cells in upper half of the leaf.

Occasionally one or two leaf cells can be observed to have two papillae.

Alar cells are big, thin-walled and inflated.

oct

Ten New Records of Mosses jrom Doi Inthanon Nasional Park in Thailand 397

This species was treated as Trichosteleum bistrumosum (Mill.

Hal.) A-Jaeger (see Bartram. 1939) because of its unipapillose leaf cell

character. Since no species of Trichostelum in the region have such strongly

falcate leaves, we preferred to follow Fleischer (1904-1923) in placing it in

Warburgiella. We also think that the leaf morphology of the species indicates

a possible link to Radulina.

Warburgiella bistrumosa, an endemic species in the Philippine, is now

found in Thailand.

Specimens studied: Y. Nathi 109, 207, 683, 1037 (BCU).

Figure 6. Warburgiella bistrumosa. a. Branch: b.-d_ Leaves: e-£ Alar cells: g Leaf cells.

398 Gard. Bull. Singapore 61 (2) 2010

Acknowledgments

The first author thanked the SING Herbarium at the Singapore Botanic

Gardens for the warm reception given her during the visit in 2008, and

to the Head of Doi Inthanon National Park and park rangers for their

generous helps during the field works. This project was supported by the

Thai government budget 2006 under the Research Program on Conservation

and Utilization of Biodiversity and the Center of Excellence in Biodiversity,

Faculty of Science, Chulalongkorn University (CEB_M_34 2007), the

Graduate School of Chulalongkorn University, and Chulalongkorn

University Centenary Academic Development Project given to the first and

the last authors.

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Tan, B.C., K. Wongkuna, S. Manachit and K. Santanachote. 2006. New

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Bryology 27: 95-100.

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Touw, A. 1968. Miscellaneous notes on Thai mosses. Natural History Bulletin

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Gardens’ Bulletin Singapore 61 (2): 401-435. 2010 401

Epifoliar Fungi of Singapore

DER. REYNOLDS

University of California Herbarium

Berkeley, California USA 94720

Abstract

This article is a discussion of the known epifoliar fungi from Singapore

found in several worldwide Herbaria, and those collected by the author.

Fifty species in 15 genera are reviewed and annotated. An additional twenty

new combinations in Setameliola are proposed to contain Meliola species

with an ascocarp bearing straight setae. Critical attention is given to the

species concepts in Meliola and Polychaeton.

Introduction

Ecologically, the epifoliar fungi are stratified in the tropical canopy

(Reynolds, 1972; Gilbert et al., 2007) with a prevalence in the understory

where abundant hosts support a greater number and diversity of fungi.

Environmental conditions affecting spatial distributions are sites in the dark

canopy and openness.

Epifoliar fungi are specialized as nutritional guilds found only on the

surface of living plants, particularly the leaves (Gilbert et al., 2007). Several,

principally, Ascomycete lineages that together comprise a polyphyletic

group, have evolved into this habitat as four nutritional guilds: saprobes,

plant parasites, fungal parasites, and lichenized species. With the exception

of lichenized species, all share common adaptive morphological traits

including a dark, melanoid pigmentation and reproduction by ascospores

and mitospores. For the non-lichenized taxa, the habits and morphological

traits are convergent plesiosynapsies that distinguish groups of fungi at

lower clade levels. In contrast, lichenized fungi have acquired the epifoliar

habit at higher clade levels equivalent to genus or species (Liicking, 2002).

The Singapore lichenized species have been reviewed by Sipman (2003a,

2003b, 2007).

Thaung (2006) observed that one distribution pattern of epifoliar fungi

is comparable with the phytogeography of the Indo-Malayan Biogeograhic

Realm with tropical rainforests, (mongabay.com/0102.htm). He suggested

that the radiation pattern indicates an Indo-Himalayan origin with dispersal

southward into SEAsia, including Singapore.

402 Gard. Bull. Singapore 61 (2) 2010

The information on currently known epifoliar fungi from Singapore is

obtained from several worldwide Herbaria, and those collected by the author

in Singapore in 2007. The epifoliar fungi from Singapore are historically

known from the Fungi Malayana exsiccatum assembled by C.F. Baker from

1916-1920. Numbers 401- 500 of Century V were collected in Singapore

out of the total 600 specimens. Twenty-two Singapore species of epifoliar

ascomycetes were identified; 12 of them were described as new by Saccardo

(1918). The exsiccatum specimens are currently known to be curated in

Herbaria BPI, CUP, FH, PC,S and UC (Pfister, 1985; Holmgren & Holmgren,

1998). Lim (1975) published a list of 13 epifoliar species from Singapore; the

cited collections are currently unavailable for reexamination.

The list of taxa is as follows —

1. Asteridiella McAlpine, Proc. Linn. Soc. New South Wales 187 (1897) 38.

Ascospores 4 septate, brown; asci evanescent; ascocarp superficial, globose,

glabrous;mycelium superficial, brown, septate, branched, asetose, hyphopodia

alternate.

1.1. Asteridiella umirayensis (Yates) Hansford, Sydowia 10 (1957) 51.

-Meliola umirayensis Yates, Philipp. Jour. Sci. Bot. 13 (1918) 370;

-Irenia umirayensis (Yates) Hansford, Proc. Linn. Soc. London 157 (1946)

70:

-Irenia bakeriana Hansford, Proce. Linn. Soc. London 157 (1946) 169.

Ascospores 4 septate, oblong to ellipsoid, 36-45 x 14-16 um. Ascocarp

scattered, to 180um diameter, surface cells conoid projecting to 15um.

Mycelium to 3mm diameter, hyphopodia alternate, ampulliform.

Specimen reported: C.F. Baker Fungi Malayana 455, determined as Meliola

reticulate Karsten & Roumeguere; BPI 696686-87, Flora of Singapore 5796,

Coll. JJ. Chupp, 2 Jul 1920.

Illustration: Hansford, 1963.

Distribution: Java, Philippines, Singapore.

2. Asterina Léveillé, Ann. Ecole Natl. Agric. Montpellier (1845) 59.

Ascospores brown, 1 septate, ellipsoid-oblong; Asci _ fissitunicate,

aparaphysate; Ascocarp dimidiate, circular, radiate, astomate; Mycelium

brown, with hyphopodia or node cells.

Epifoliar Fungi of Singapore 403

2.1. Asterina erebia Sydow, Ann. Mycol. 25 (1927) 59, sensu Stevens & Ryan,

Ill. Biol.

Monogr. (1939) 70. Fig. A.

Ascospores 26-32 x 12-16 um. Ascus 55-65 um. Ascocarp 150-300 um:

hyphopodia alternate, 2-celled.

Specimens reported: Singapore, Bukit Timah Nature Reserve, Oct 2007,

DRR 15, 16, 17, 18.

Distribution: Singapore.

Figure A. Asterina erebia Sydow. Mycelium with hyphopodia and ascocarps [from DRR Sin-

gapore 16; line = 20 um].

2.2. Asterina singaporensis Sydow, Ann. Mycol. 18 (1920) 159. Fig. B.

Ascospores 16-20 x 6-8 um. Asci 30-40 um. Ascocarp 80-110 um diameter.

Colonies epiphyllous, 2-5 mm diameter, hyphopodia dispersed rarely

opposite, 10-15 um long.

Specimen reported: C.F. Baker Fungi Malayana 401. Holotype at UC.

Sydow and Sydow (1917) noted that this specimen was distributed as

Asterina trachycarpa Sydow (see also Saccardo, 1918).

Distribution: Singapore.

404 Gard. Bull. Singapore 61 (2) 2010

Figure B. Asterina singaporensis Sydow. The dimidiate ascocarp is formed beneath

_ hyphopodate hyphae [from C.F Baker Fungi Malayana 301; line = 10 um].

3. Ciferriusia Batista in Batista & Ciferri, Sydowia Beih. 3 (1962) 17.

Ascospore 3 transseptate. Asci fissitunicate, paraphysate. Ascocarp

developed beneath a mycelia pellicle, without setae. Mycelium hyaline to

subhyaline, septate.

3.1. Ciferriusia orientalis Batista & Costa in Batista & Ciferri, Sydowia

Beih. 31962) 17:

Ascocarp globose-depressed, with a mycelical shield, 90-180 um diameter,

ostiolate. Asci 40-60

um. Ascospores clavate, 3-transversely septate, hyaline, 15-20 x 4-5 um.

Specimen collected: Singapore Botanical Garden, Oct 2007, DRR 4.

Illustration: Batista & Ciferri, 1962: Fig. 11.

Distribution: Philippines, Singapore.

Note: Arx and Miiller (1975) considered Ciferriusia to be a synomym of

Yatesula H. Sydow & P. Sydow, which has muriform ascospores.

4. Ellisiodothis Theissen, Ann. Mycol. 12 (1914) 73.

Ascospores hyaline, unicellular. Ascus fissitunicate, pseudoparaphysate, 45-

50 x 16 um; Ascocarp hypostomatic, superficial, dimidiate, nonradiate, dark.

Epifoliar Fungi of Singapore 405

This genus is closely related to Microdothella Sydow and Myiocopron

Spegazzini. It was suggested as a synonym of the latter taxon by A. Sivanesan

(see Kirk, 2008).

4.1. Ellisiodothis grammatophylli (Saccardo) Sydow & Sydow, Ann. Mycol.

18 (1920) 185.

-Microthyrium grammatophylii Saccardo, Bull. Orto Bot. Regia Univ. Napoli

6 (1918) 49.

-Ellisiodothis grammatophylli Sydow, Ann. Mycol. 18 (1920) 134.

Ascospores 22 x 6 um. Ascus 45-50 x 16 um. Ascocarp 325-450 um.

Specimen reported: C.F. Baker Fungi Malayana 467. Holotype at PAD and

isotype, UC.

Distribution: Singapore.

5. Lembosia Léveillé, Ann. Sci. Nat. 3 (1845) 58.

Ascospores 1 septate, brown. Asci spherical, fissitunicate, paraphysate.

Ascocarp long or linear forming, X or Y forming, with a longitudinal slit.

Mycelium superficial, hyphae with lateral or intercalary hyphopodia,

hypostroma or innate hyphae absent

5.1. Lembosia hormosiana Saccardo, Bull. Orto Bot. Regis Univ. Napoli 6

(1918) 52. Ascospores 17-16 x 15-6.5 um. Ascus 35-40 x 18-20 um. Ascocarp

235-400 x 90 um.

Specimen reported: C.F. Baker Fungi Malayana 443. Holotype at PAD and

isotype, UC.

Distribution: Singapore.

5.2. Lembosia pandani (Rostafinski) Theissen, Ann. Mycol. 11 (1913) 457.

-Asterina pandani Rostafinski, Bot. Tidsskr. 24 (1902) 361.

Ascospores 28-38 x 24-26 um. Ascus 28-30 x 24-26um. Ascocarp 350-850

um.

Specimen reported: C.F Baker Fungi Malayana 444. Holotype at FH and

isotype, SING. Distribution: Philippines, Singapore.

6. Lembosina Theissen, Ann. Mycol. 11 (1913) 437.

Ascospores brown, l-septate. Asci fissitunicate, paraphysate. Ascocarp

superficial, elongated or linear, textura radiate, opening by a longitudinal slit.

406 Gard. Bull. Singapore 61 (2) 2010

Mycelium subcuticular or intraepidermal, crustose stroma present. Hyphae,

septate, without hyphopodia, forming hypostroma between the epidermis

and cuticle.

6.1. Lembosina heptapleuri (Saccardo) vy. Arx in Arx & Miiller, Studies in

Mycol. 9 (1962) 123.

-Lembosia heptapleuri Saccardo, Bull. Orto Bot. Regis Univ. Napoli 6 (1918)

325

-Echidnodes heptapleuri (Saccardo) H. Sydow & P. Sydow, Ann. Mycol. 18

(1921) 185;

Ascospores 16 x 5 um. Ascus 40 x 12 um. Ascocarp 300-1000 x 100-120 um.

Specimen reported: C.F. Baker Fungi Malayana 442, as Lembosia heptapleuri.

Holotype at PAD and isotype, UC.

Distribution: Singapore.

7. Meliola Fries emend. Mibey & Hawksworth, Mycol. Pap. 174 (1997) 23.

_ - Amphitrichum Nees ex Sprenger, Pl. Crypt. Trop. (1820) 46 pro parte sensu

Hansford (1961).

- Asteridium Saccardo, Syll. Fung. | (1882) 49.

- Courturea Castagne In Fries, Summ. Veg. Scand. (1846) 407.

- Meliola Fries emend. Bornet, Ann. Sci. Nat III: 16 (1851) 267.

- Meliola Fries, Syst. Orb. Veg. (1825) 111.

- Mycothecium Kunze ex Fries, Syst. Mycol. 3 (1829) 232.

- Sphaeria Fries, Syst. Orb. Veg. (1823) 513, pro parte sensu Hansford

(1961).

Ascospores darkly pigmented, 3-4 transseptate; Ascus unitunicate, uniformly

thin-walled; Ascocarp globose, glabrous. Mycelium superficial, comprised of

darkly pigmented, septate, setose hyphae; capitate hyphopodia alternate or

unilateral, opposite or mixed opposite and alternate. Asexual reproduction,

when present, by single phialides producing unicellular, hyaline mitospores.

Meliola was first described by Fries (1825) with a single species and currently

includes 2957 taxa (Kirk, 2008). The genus was one of five genera recognized

in the Meliolineae by Hansford (1961). This lineage was recognized as the

Meliolaceae by Dennis (1968, 1970), Stevens (1927, 1928), and Stevenson

(1975), and Meliolales by Alexopoulos er al. (1996) and Miiller and Arx

(1973). Luttrell (1989) placed Meliola in the Meliolales of the Pyrenomycetes,

which was sustained in a phylogeny discovered by Saenz and Taylor (1999).

A careful and detailed monographic study of 1814 taxa in the genus

was undertaken by Hansford (1961). Nonetheless, a more than double

Epijoliar Fungi of Singapore 407

proliferation of names has been inadvertently manifested with Hansford’s

unaccountable arrangement of the species according to the families of the

host plants on which they occurred. Subsequent new taxa have been described

because of the compartmentalization introduced with the associated

vascular plant family as a plesiomorphic character. A comparison of species

descriptions that disregards this character demonstrates the taxonomic

redundancy. This artificial classification is underscored in the Hansford

(1961) monograph by some species listed as occurring on members of more

than one vascular plant family and by the recognition of species in which the

host is unknown. The Hansford (1961) approach also implies a co-evolution

of Meliola with angiosperms at the family level, which is not supported by

distribution patterns on a phylogenetic clade for the latter.

The consequence of Hansford’s Meliola species concept is a high

potential for a redescription of the same fungus several times. An example

is the excessive number of proliferation of uncritically described species by

subsequent workers (Hosagoudar 1994, 1996, 2000: Hosagoudar et al., 1997:

Hosagoudar and Agarwal, 2008: Hu, 1996, 1999: and Mibey and Hawksworth

1997: Rodriguez 2006). This report does noi recognize the associated vascular

plant host as a plesiomorphic character.

The species concept of Meliola recognized here does not recognize

the associated vascular plant host as a plesiomorphic character, although

the host range appears to be limited for some clades (Stevens, 1926; Ciferni,

1954: Rodriguez, 2001). The consequence is a description of the same fungus

several times because of the host affiliation.

A useful device for an assessment of a Meliola species was introduced

by Beeli (1920). A numerical code was derived from a set of eight character

states with from 3 to 6 variables. The Beeli Formula was revised by Stevens

(1927, 1928). Hansford (1961) and Farr (1971). Some authors (Mibey and

Hawksworth, 1977) criticize the Beeli Formula as not representing all

features and thus was said to be diagnostically lacking. The occurrence of

the same Beeli Formula for a group of species was dismissed as insignificant

because of the supposed value of other features that would purportedly

separate species. These include finite distinctions such as small variations in

ascospore size and hyphal or hyphopodial cell size. Most of the features they

(Mibey and Hawksworth, 1977) would utilize in the species concept for

critical separation of taxa are highly variable and thus unreliable in species

recognition, except as a promulgation.

I believe that Beeli recognized the core characters that comprise a

species concept for Meliola. The continuous character delimitations reflect

natural size ranges. Furthermore I regard all putative taxa with a similar

Beeli Formula to represent the same or a group of closely related species.

Phylogenetically, a large number of overlapping names have been proposed

408 Gard. Bull. Singapore 61 (2) 2010

for the termini of a monophyletic lineage with morphological expression

better represented by the Beeli Fomula than the minutial variations of these

same characters as exemplified by Mibey and Hawksworth (1997).

This predominately tropical genus first appeared in the fossil record

in the Eocene (Dilcher, 1965), at a time in the Cretaceous when associated

angiosperm hosts were diversifying from a Triassic origin (Taylor et al.,2009).

The fossilized, melanoid ascocarp walls of this taxon and other foliicolous

ascomycetes are well preserved on compressed leaf cuticles. A lineage older

than the Eocene is suggested by the remarkable similarity of specimens to

extant species, although the centrum was rarely preserved.

7.1. Meliola aethiops Saccardo, Bull. Orto Bot. Regis Univ. Napoli 6 (1918)

41.

-Meliola aglaina Hansford, Sydowia 9, (1955) 59.

-Meliola ekebergiae Hansford, J. Linn. Soc. Bot. 51 (1937) 274.

-Meliola kauaiensis Stevens, Bernice P. Bishop Mus. Bull. 19 (1925) 39.

-Meliola mataybae Stevens, Ann. Mycol. 26 (1928) 228;

-Meliola capensis (Klatchbrenner & Curtis) Theissen var mataybae (Stevens)

~ Hansford,

Sydowia Beih. 2 (1961) 447.

-Meliola nicaraguensis Spegazzini, Bol. Acad. Nac. Cien. Cordoba 26 (1926)

378.

-Meliola opposite Sydow var africana Hansford, Sydowia Beih. 2 (1961)

411.

Ascospore 32-45 x 11um.Ascocarp to 180 um diameter. Mycelium to 3 mm

diameter;

capitate hyphodia opposite or alternate, ampulliform; mycelial setae grouped

around ascocarp, straight, simple, 280 um.

Beeli Formula: 31123221

Specimen reported: C.F: Baker Fungi Malayana 449, determined as Meliola

aethiops. Holotype at SING (#33777). A Singapore specimen was cited as

Meliola aethiops by Lim (1975).

Illustration: Hansford, 1963.

Distribution: Philippines, Singapore.

7.2. Meliola citricola Sydow, Ann. Mycol. 15 (1917) 183.

[IMI Descriptions of Fungi and Bacteria 2006, 168, Sheet 1672].

Epifoliar Fungi of Singapore 409

-Meliola bussei Mibey in Mibey & Hawksworth, Mycol. Pap. 174 (1997) 32.

-Meliola khayae Hansford var minor Hansford & Deighton Mycol. Pap. 23

(1948) 43.

-Meliola ostryodemidis Hansford & Deighton. Mycol. 3 (1948) 34.

-Meliola ostryodemidis Hansford & Deighton var /eptoderridis Hansford &

Deighton 1948. Mycol. Pap. 23 (1948) 35.

-Meliola stizolobii Hanstord & Deighton var desmodii-salicifolii Hansford

& Deighton, Mycol. Pap. CMI 23 (1948) 31.

Ascospores 30-48 um x 9-19 um. Ascocarp to 140-170 um diameter. Mycelium

to 5 mm diameter; capitate hyphopodia alternate, opposite, straight to

antrorse. Mycelial setae scattered or grouped around ascocarp,160-800 um.

Briania phialides (Reynolds, 1989b) sometimes present.

Beeli Formula: 31334223

Specimens reported: Hansford (1961) cites a Singapore collection, Chupp

5949, which is curated in SING as #38557 and #96896; they are determined

as Meliola. An addition specimen, Baker 1627, is also cited (Type: Herb. PBS

23747).

Illustrations: Stevens, 1928; Hansford, 1963; Lim, 1975; Hosgoudar & Agarwal,

2008.

Distribution: Borneo, Philippines, Singapore, Sumatra, Taiwan.

7.3. Meliola cylindropoda Doidge, Trans. Roy. Soc. South Africa 8 (1920)

138.

-[renina mangostana (Saccardo) Stevens, Ann. Mycol. 25 (1927) 457.

-Meliola araliicola Yamamoto, Trans. Nat. Hist. Soc. Formosa 31 (1941) 224.

-Meliola euchrestiae Yamamoto, Trans. Nat. Hist. Soc. Formosa 31 (1941)

226.

-Meliola eugeniae-jamboloidis Hansford var paulensis Hansford, Sydowia 9

G1955))' 62.

-Meliola garcinae Yates var mangostana (Saccardo) Hansford, Proc. Linn.

Soc. London 160 (1948) 120.

-Meliola littoralis Sydow, Bothalia 2 (1928) 462.

-Meliola mangostana Saccardo, Bull. Ort. Bot. Napoli 6 (1921) 42.

-Meliola plumbaginis Hansford & Stevens, J. Linnean Soc. London 51 (1937)

280.

-Meliola tecleae Hansford var toddaliae-asiaticae Hansford, Proc. Linn. Soc.

London 153 (1941) 11.

410 Gard. Bull. Singapore 61 (2) 2010

Ascospores 38-44 um x 15-17 um. Ascocarp to 240 um diameter. Mycelium

to 6 mm diameter, capitate hyphopodia alternate, straight or bent. Mycelial

setae scattered, to 1110 um.

Beeli Formula: 31114233

Specimen reported: C.F. Baker Fungi Malayana 453.This is the type specimen

for M. mangostana (Herb. SING 33779).

Illustration: Hansford, 1963.

Distribution: Philippines, Singapore, Taiwan.

7.4. Meliola fagraeae Sydow & P. Sydow, Ann. Mycol. 12 (1914) 549.

- Meliola mannavanensis Hosagoudar, C.K. Biju, Baram & Crane. Mycotaxon

76 (2000) 302.

-Meliola rajamalaensis Hosagoudar, C. K. Biju & Abraham. Nova Hedwigia

80 (2005) 493.

Ascospores 41-55 um x 15-19 um. Ascocarp to 225 um diameter. Mycelium

to 3mm diameter, capitate hyphopodia alternate, straight. Mycelial setae

numerous, scattered, straight or flexuous, to 360 um.

Beeli Formula: 31215232

Specimen reported: BO 5354, reported in Hansford 1961:525.

Illustration: Hansford, 1963.

Distribution: Philippines, Singapore, Sumatra

7.5. Meliola garciniae Yates, Philipp. Journ. Sci. Bot. 13 (1918) 369.

-Meliola dognyensis Huguenin, Rev. Mycol. 34 (1969) 52.

-Meliola kisubiensis Hansford, var. peleicola Hansford, Sydowia 9 (1955) 43.

-Meliola kydia Saccardo Bull. Orto Bot. Regis Univ. Napoli 6 (1918) 13.

-Meliola mauritiana Hansford, Sydowia 9 (1955) 20.

-Meliola megalochaeta Sydow, Philipp. Journ. Sci. 21 (1932) 135.

-Meliola teke Hansford, Proc. Linnean Soc. Lond. 157 (1946) 180.

Ascospores 41-59 x 15-25 um. Ascocarp 180-360 um diameter. Mycelium (3)

5-10 mm diameter; capitate hyphopodia opposite or alternate; mycelia setae

simple, entire, acute to obtuse or dentate, 325-1600 um.

Beeli Formula: 31115334

Epifoliar Fungi of Singapore 41 1

Specimen reported: C.F. Baker Fungi Malayana 450, determined as Meliola

kydia. Holotype at PAD and isotype, UC.

Illustration: Hansford, 1963.

Distribution: Amboina, Borneo, Java, Philippines, Singapore, Vietnam.

7.6. Meliola heterodonta Sydow, Ann. Mycol. 14 (1916) 357.

-Meliola columbiensis Hansford, Sydowia 9 (1955) 61.

-Meliola justiciae Hansford, Journ. Linn. Soc. Lond. 31 (1938) 541.

-Meliola micropoda Hansford, Proc.Linn. Soc. Lond. 121 (1946) 121.

-Meliola millettiae-chrosyphillae Deighton, Sydowia 7 (1951) 7.

-Meliola nephelii Saccardo, Bull. Orto Bot. Regis Univ. Napoli 6 (1918) 42.

-Meliola newbouldiae Hansford & Deighton, Mycol. Pap. 23 (1948) 66.

-Meliola oncinotidis Doidge, Bothalia 4 (1948) 851.

-Meliola securidacae Hansford, Sydowia 10 (1957) 88.

-Meliola securidacicola Hansford, The Meliolineae, a monograph. Sydowia

Beih. 2 (1961) 82.

-Meliola stizolobii Hansford & Deighton, Mycol. Pap. 23 (1948) 31.

-Meliola tounatae Stevens, Ann. Mycol. 26 (1948) 204.

Ascospores 28-40 x 11-24 um. Ascocarp to 190 um: Mycelium 8 mm diameter;

mycelia setae simple, entire, acute to obtuse, or dentate, scattered or grouped

around perithecium, to 960 um: capitate hyphopodia opposite or alternate,

subanthrose to straight. Hughesia phialides sometimes present.

Beeli Formula: 31133223.

Specimens reported: Baker Fungi Malayana 252 (holotype, PBS 4031); also

Baker Fungi Malayana 454 (holotype of Meliola nephelii, SING 33781).

Illustrations: Hansford, 1963; Hosagoudar & Agarwal, 2008.

Distribution: Java, Philippines, Singapore.

7.7. Meliola inocarpi Stevens, Ann. Mycol. 26 (1928) 232.

Ascospore 43-52 x 17-20 um. Ascocarp 190 um diameter. Mycelium 2-8

mm diameter, setae to 900 um; capitate hyphopodia opposite to alternate,

straight or bent.

Beeli Formula: 31124323

4] 2 Gard. Bull. Singapore 61 (2) 2010

Specimen reported: C.F. Baker Fungi Malayana 459, determined as Meliola

sp. Holotype at ILL, and isotype, UC.

Illustration: Hansford, 1963. Distribution: Singapore

7.8. Meliola litseae Sydow, Ann. Mycol. 15 (1917) 366.

-Meliola golaensis Deighton, Sydowia 11 (1958) 105.

-Meliola litseae Yates, Philip. Journal Sci. C. Botany 12 (1918) 366.

-Meliola micheliae Hansford, Proc. Linn. Soc. London 158 (1947) 34.

-Meliola singaporensis Hansford, Proceedings of the Linnean Society,

London 157 (1946) 17.

Ascospores 4 septate, 42-60 x 16-25 um. Ascocarp to 225 um. Mycelium

amphigenous, thin to velvety, | to7 mm diam. Hyphae substraight to undulate

or flexuous, capitate hyphopodia alternate, antrorse to ampulliform, mycelia

setae scattered or grouped around ascocarp, entire, obtuse or dentate, 600-

1000 um.

Beeli Formula: 31114323

Specimens reported: C.F Baker Fungi Malayana 457, determined as Meliola

sp. C.F Baker Fungi Malayana 480, type of M. singaporensis (holotype,

SING).

Illustrations: Hansford, 1963; Hosagoudar & Agarwal, 2008.

Distribution: Philippines, Singapore, Taiwan.

7.9. Meliola mangiferae Earle, Bull. New York Bot. Gard. 3 (1905) 307.

-Meliola helicae Yamamoto, Trans. Nat. Hist. Soc. Formosa 31 (1941) 54.

-Meliola naisophyllae Hansford & Deighton, Mycol. Paper IMI 23 (1948)

ish

-Meliola palmicola Winter var coperniciae Spegazzini, Anal. Mus. Nac. Bueos

Aires 32 (1924) 384.

-Meliola subdentata Patouillard, Journal de Bot. (1897) 347.

-Meliola subdentata Patouillard var microspora Hansford & Deighton,

Mycol. Paper IMI 23 (1948) 72.

-Meliola taityuensis Yamamoto, Trans.Nat. Hist. Soc. Formosa 31 (1941) 131.

-Meliola trichoscyphae Hansford & Deighton, Sydowia 10 (1957) 94.

Ascospores4 septate, 40-59 x 12-25 um. Ascocarp 180-350 um. Mycelium 3 to

10mm diameter, hyphopodia alternate or opposite, ampulliform to antrorse,

mycelia setae scattered or grouped around ascocarp, simple, entire, acute to

obtuse or dentate 810 4m. Hughesia phialides sometimes present.

e 6

Epifoliar Fungi of Singapore 413

Beeli Formula: 31315333

Specimen reported: C.F. Baker Fungi Malayana 452, determined as Meliola

mangiferae. Holotype, NY; SING

33779, annotated as “type”.

Illustrations: Hansford, 1963; Hosagoudar & Agarwal, 2008.

Distribution: Sierra Leone, Australia, Singapore, Taiwan, Philippines,

Indonesia, Jamaica, Porto Rico, Venezuela.

7.10. Meliola panici Earle, Muehlenbergia 1 (1901) 12.

-Meliola africana Hansford, Sydowia 10 (1957) 62.

-Meliola allophylorum Mibey in Mibey & Hawksworth, Mycol. Pap. 174

(1977) 29.

-Meliola alyxiae Stevens, Bernice P. Bishop Mus. Bull. 19 (1925) 30.

-Meliola ambigua Patouillard & Guillard var caseariaecola Ciferri,Mycopath.

Mycol. Appl. 7 (1954) 93.

-Meliola aristolochiae Stevens & Tehon, Mycologia 18 (1926) 4.

-Meliola banarae Stevens, Ann. Mycol. 26 (1928) 249.

-Meliola beilschmiediae Yomomoto, Trans. Nat. Hist. Soc. Taiwan 31 (1941)

a2.

-Meliola berliniae Hansford & Deighton, Mycol. Pap. (1948) 23.

-Meliola camellicola Yamamoto, Trans. Nat. Hist. Soc. Taiwan 31 (1941) 53.

-Meliola canarii Sydow, Ann. Mycol. 12 (1914) 550.

-Meliola caseariae-arboreae Hansford var. guatemalensis Hansford, Sydowia

Beth: 1(1957) 103.

-Meliola caseariae-guanensis Hansford, Sydowia 9 (1955) 11.

-Meliola castaneifoliae Mibey in Mibey & Hawksworth, Mycol. Pap. 174

(1977). 37.

-Meliola castanha Theissen, Broteria 12 (1914) 24.

-Meliola chasaliae Mibey in Mibey & Hawksworth, Mycol. Pap. 174 (1977)

38.

-Meliola cumbrensis Hansford, Sydowia 10 (1957) 69.

-Meliola cunoniae Hansford, Sydowia 9 (1955) 13.

-Meliola dactylipoda Sydow, Bothalia 2 (1928) 460.

-Meliola dasiana Mibey in Mibey & Hawksworth, Mycol. Pap.174 (1977)

41.

-Meliola daviesii Hansford var longiseta Hosagoudar, Mycopath. Mycol.

Appl. 44 (2006) 44.

-Meliola desmodii-laxiflori Deighton var crotalariae Deighton, Sydowia 11

(1958) 41.

414 Gard. Bull. Singapore 61 (2) 2010

-Meliola eryeibes Hansford, Reinwardtia 3 (1954) 81.

-Meliola erythrinae Sydow, Ann. Mycol. 15 (1917) 185.

-Meliola erythrinae Sydow var. psophocarpi Hansford, Sydowia 10 (1957) 70.

-Meliola excoecariae Doidge, Trans. Roy. Soc. South Africa 8 (1920) 139.

-Meliola ficium Yates var ugandensis Hansford, Sydowia 10 (1957) 72.

-Meliola forsteroniae (Stevens) Hansford, Proc. Linn. Soc. Lond. 160 (1948)

1920.

-Meliola garryae Hansford, Sydowia 9 (1955) 17.

-Meliola gnetii Hansford, Reinwardtia 3 (1954) 85.

-Meliola gregoriana Stevens, Bernice P. Bishop Mus. Bull. 19 (1925) 39.

-Meliola grewiicola Hansford, Sydowia 11 (1958) 56.

-Meliola ichnocarpii Hansford & Thirumalachar., Farlowia 3 (1948) 295.

-Meliola ichnocarpicola Hansford, Sydowia Beih. 2 (1961) 559

-Meliola knowltoniae Doidge, Bothalia 1 (1924) 308.

-Meliola landolphiae-floridae Hansford, Jour. Linn. Soc. Bot. 51 (1938) 542.

-Meliola landolphiicola Hansford, Proc. Linn. Soc. Lond. 157 (1945) 23.

-Meliola leptochaeta Sydow, Ann. Mycol. 15 (1917) 187.

-Meliola lianchangensis Jiang, Acta Mycol. Sin. 14 (1995) 2.

-Meliola litseae Graff, Mem. Torrey Bot. Club 17 (1917) 61.

~ -Meliola litseae Sydow var. rotundipodia Hansford, Reinwardtia 3 (1954) 88.

-Meliola logiseta H6hnel, Akad.Wiss. Wien, Sitzungsber. Math.-Naturwiss.

KI. Abt. 1:116 (1907) 100

-Meliola lucumae Stevens, Illinois Biol. Monogr. 2 (1916) 49.

-Meliola mombasana Mibey in Mibey & Hawksworth, Mycol. Pap.174 (1977)

55)

-Meliola ochrocarpii Thite & Patil, Geophytology 13 (1983) 125.

-Meliola pachystellae Mibey in Mibey & Hawksworth, Mycol. Pap.174

(1977) 62

-Meliola palaquiicola Hansford, Sydowia 11 (1958) 58.

-Meliola pandanii Sydow, Ann. Mycol. 26 (1928) 89.

-Meliola panici Earle var. vetiveriae Hansford & Deighton in Hansford,

Sydowia 10 (1958) 82.

-Meliola pinnatae Miber in Mibey & Hawksworth, Mycol. Pap.174 (1977) 66.

-Meliola plectroniae Hansford, Sydowia 9 (1955) 72.

-Meliola pseudocapensis Hansford, Sydowia 9 (1955) 23.

-Meliola ruiacearum Mibey in Mibey & Hawksworth, Mycol. Pap. 174 (1977)

70.

-Meliola sauropicola Yates, Philipp. Journ. Sci. Bot. 12 (1917) 368.

-Meliola semecarpii Sydow, Ann. Mycol. 21 (1923) 95.

-Meliola semecarpicola Hansford, Sydowia 11 (1958) 58.

-Meliola serjaniae Stevens, Illinois Biol. Monogr. 2 (1916) 44

-Meliola tabernaemontanae Spegazzini var forsteroniae Stevens, Illinois Biol.

Epijoliar Fungi of Singapore 415

Monogr. 2 (1916) 50.

-Meliola terecitensis Hansford, Sydowia Beih. 1 (1957) 117.

-Meliola tijucensis Hansford, Sydowia 9 (1955) 77.

-Meliola torricelliae Kar & Maity. Nytt Mag. Bot. 17 (1970) 81.

-Meliola trichiliicola Spegazzini, Anal. Mus. Nac. Buenos Aires 32 (1924)

366.

-Meliola uncariicola Deighton, Sydowia 11 (1958) 42.

-Meliola venezuelana Orejuela, Mycologia 36 (1944) 437.

-Meliola vicina Sydow, Ann. Mycol. 21 (1923) 95.

Ascospores 4-septate, 17-66 x 11-28 um. Ascocarp to 250 um diameter.

Colony diameter to 20 mm, thin to dense, velvety: Mycelium with setae,

scattered or grouped around ascocarp, simple, acute. to 1000 um: capitate

hyphopodia alternate to unilateral or opposite, antrorse.

Beeli Formula: 31114223

Specimen reported: C.F. Baker Flora Malayana 456, determined as Meliola

sp. Listed as M. panici by Hansford (1961): 745. Holotype at NY: isotype.

UC.

Illustrations: Hansford, 1963; Hosagoudar & Agarwal, 2008.

Distribution: Borneo, Java, Malaya, Philippines, Singapore, Taiwan.

7.11. Meliola psidii Fries, Linneae 5 (1830) 549.

-Amazonia gniomae Doidge, Bothalia 1 (1924) 204.

-Meliola aethiops Saccardo var cassia Rao, Mycopath. Mycol. Appl. 33 (1967)

163.

-Meliola alchorneae Stevens & Tehon, Mycologia 12 (1926) 21.

-Meliola allyphyli Doidge var pervillei Mibey & Hawksworth, Mycol. Pap.

174 (1997) 28.

-Meliola amadelpha Sydow, Leafi. Philipp. Bot. 9 (1925) 3114.

-Meliola anodendri K. Sawada & Yamamoto, Spec. Publ. Coll. Agri. Nat.

Taiwan Univ. 8 (1959) 28.

-Meliola asclepiadacearum Hansford var brasiliensis Hansford, Sydowia 11

(1958) 52.

-Meliola banarae Stevens var aculeatae Ciferri, Mycopathologia 7 (1954) 99.

-Meliola bicornis var galactiae Stevens, Ill. Biol. Monogr. 2 (1916) 65.

-Meliola biparasitica Ciferri, Ann. Mycol. 36 (1938) 205.

-Meliola borbonicae Mibey in Mibey & Hawksworth, Mycol. Pap. 174 (1997)

als

-Meliola bridiicola Hansford, Proc. Linn. Soc. Lond. 157 (1946) 175

416 Gard. Bull. Singapore 61 (2) 2010

-Meliola bryae Hansford, Sydowia 10 (1957) 65

-Meliola bwaniana Mibey in Mibey & Hawksworth, Mycol. Pap. 174 (1997)

33:

-Meliola canthii Hansford var leonensis Hansford & Deighton, Mycol. Pap.

23 (1948) 59.

-Meliola clavulata Winter var jamaicensis Hansford, Sydowia 11 (1957) 54.

-Meliola cnestidis Doidge, Bothalia 2 (1928) 453.

-Meliola cochlospermifolii Batista apud Batista, Atlas Inst. Micol. Univ.

Recife 1 (1960) 34.

-Meliola corazoyensis Hansford, Sydowia Beih. 2 (1961) 375.

-Meliola dactylipoda Sydow var. jamaicensis Hansford, Sydowia 10 (1957) 68.

-Meliola daviesii Hansford, Proc. Linn. Soc. Lond. 157 (1946) 176.

-Meliola dipholidis Stevens , Ill. Biol. Monogr. 2 (1916) 44.

-Meliola erioglossi Hansford, Sydowia Beth. 1 (1957) 107.

-Meliola feretiae Hansford, Sydowia 10 (1957) 71.

-Meliola ferruginiae Mibey in Mibey & Hawksworth, Mycol. Pap. 174 (1997)

45.

-Meliola funtumiae Beeli, Bull. Jard. Bot. Etat. 7 (1920) 95.

_ -Meliola galactiae (Stevens) Hansford, Sydowia Beih. (1961) 287.

-Meliola gardneriae Hansford & Thirumalachar, Farlowia 3 (1948) 293.

-Meliola goianensis Batista & Maia apud Batista, Atlas Inst. Micol. Univ.

Recife 2 (1965) 265.

-Meliola goniomae Doidge, Bothalia 2 (1928) 461.

-Meliola guareae Spegazzini, Ann. Mus. Nac. Buenos Aires 23 (1913) 42.

-Meliola hendrickxiana Hansford, Proc. Linn. Soc. Lond. 159 (1947) 25.

-Meliolahenyri Hosagoudar var oldenlandiae Hosagoudar, Biju & Abraham,

Nova Hedw. 80 (2005) 486.

-Meliola heyneae Hansford & Thirumalachar, Farlowia 3 (1948) 294.

-Meliola hippomaneae Stevens, Ann. Mycol. 28 (1928) 284.

-Meliola horrida Ellis & Everhart in Smith, Bull. Univ. lowa 2 (1893) 396.

-Meliola hoyae Saccardo, Atti Accad. Sci. Ven. 10 (1917) 60.

-Meliola hughesiana Hansford, Sydowia 10 (1957) 76.

-Meliola hypselodelphidis Hughes, Myco. Pap. 48 (1952) 51.

-Meliola jasmine Hansford & Stevens in Hansford, Journ. Linn. Soc. Bot.

(1937) 273:

-Meliola lasiacidis Toro in Chardon & Toro, Monograph Univ. Porto Rico B

21934) 121%

-Meliola lictorea Ciferni, Ann. Mycol. 36 (1938) 214.

-Meliola lobellicola Farr, Canad. J. Bot. 47 (1969) 372.

-Meliola longispora (Gaillard) Stevens, Ann. Mycol. 26 (1928) 244.

-Meliola malacotricha Spegazzini var. longispora Gaillard, Le Genre Meliola

(1892) 82.

Epifoliar Fungi of Singapore 4 l i.

-Meliola microthea Sydow, Ann. Mycol. 37 (1939) 331.

-Meliola mitragynicola Deighton var leonensis (Hansford & Deighton)

Deighton, Sydowia 11 (1958) 111.

-Meliola mitragynicola Deighton, Sydowia 11 (1958) 110.

-Meliola ocoteicola Stevens, Ill. Biol. Monogr. 2 (1916) 45.

-Meliola oldenlandiae Hansford & Stevens var indica Hosagoudar, Biju &

Abraham, Nova Hedw. 80 (2005)

492.

-Meliola oligopoda Sydow, Ann. Mycol. 21 (1923) 89.

-Meliola opuntiae Hansford, Sydowia 10 (1957) 81.

-Meliola ormocarpi Hansford & Deighton, Mycol. Pap. 23 (1948) 30.

-Meliola osmanthi Sydow var. hawaiiensis Hansford, Sydowia 9 (1955) 44.

-Meliola panici Earle var lasiacidis (Toro) Hansford, Sydowia Beih. (1961)

747.

-Meliola panici Earle var. olyrae Hansford, Sydowia 10 (1957) 82.

-Meliola parenchymatica Gaillard, Le Genre Meliola (1892) 180

-Meliola pentaphylacis Song & Hu in Song & Hugh, Journ. Trop. Subtrop.

ot.15\(1997) 37.

-Meliola petrospermicola Stevens & Roldan, Philipp. Journ. Sci. 56 (1935) 69.

-Meliola phoebes Hansford, Proc. Linn. Soc. London 160 (1948) 136.

-Meliola physostigmatis Hansford & Deighton, Mycol. Pap. 23 (1948) 33.

-Meliola psychotriae Hansford, Sydowia Beih. 2 (1961) 375.

-Meliola psychotriae Earle var moreliae Hansford & Deighton, Sydowia

Beil: 2 (1961) 597.

-Meliola randiae-aculeate Hansford, Sydowia 9 (1955)74.

-Meliola ranganathii Hansford, Proc. Linn. Soc. London 157 (1946) 185.

-Meliola samydae Ciferri, Ann. Mycol. 29 (1931) 285.

-Meliola saurAuiae Sydow, Ann. Mycol. 37 (1939) 232.

-Meliola scabriseta Hansford & Deighton var. brasiliensis Hansford,Sydowia

91955). 47.

-Meliola scaevolae Sydow, Ann. Mycol. 12 (1914) 551.

-Meliola secamonis Hansford, Proc. Linn. Soc. Lond. 156 (1944) 39.

-Meliola silventvalleyensis Hosagoudar, Journ. Mycopath. Mycol. Appl.

Research 45 (2006) 3.

-Meliola straussiae Hansford, Sydowia Beih. 1 (1957) 117.

-Meliola strophanthi Hansford non Doidge, Journ. Linn. Soc. Bot. 51 (1937)

282

-Meliola strophanthicola Hansford, Sydowia Beih. 61 (1961) 551.

-Meliola taitensis Mabey & Cannon, Crypto. Mycol. 20 (1999) 277.

-Meliola thungergiicola Hansford & Deighton, Mycol. Pap. 23 (1948) 68.

-Meliola trichostroma (Kunze) Toro, Journ. Depart. Agric. Univ. Porto Rico

36 (1952) 62.

418 Gard. Bull. Singapore 61 (2) 2010

-Meliola yuanjiangensis Jiang, Acta Mycol. Sinica 8 (1989) 177.

-Meliola zamboangensis Hansford, Sydowia Beith. 1 (1957) 119.

-Sphaeria trichostroma Kunze in Weigelt’s Exsiccatum (1827). (see Pfister,

Mycotaxon 23: (1985) 1-139.

Ascospores 33-48 x 11-21 um. Ascocarp 140-200 um diameter. Mycelium

to 15 mm; mycelial setae scattered or grouped around ascocarp, simple,

sometimes dentate, 270-600 um; capitate hyphodia alternate.

Beeli Formula: 31114222

Specimen reported: A specimen of A. Chupp 5948 (SING 38558) at K is cited

by Hansford

(1961) from Singapore. Holotype at S.

Illustrations: Hansford, 1963; Housgoudar & Agarwal, 2008.

Distribution: Borneo, Burma, Malaya, Philippines, Singapore, Taiwan.

_ 7.12. Meliola sp.

Specimen reported: C.F. Baker Fungi Malayana 458. UC, BPI 692188-89.

Distribution: Singapore

Note: The Baker collection is not a Meliola; the material appears to be sterile.

8. Meliolina H. Sydow & P. Sydow, Ann. Mycol.12 (1914) 553.

Ascospores 3-septate, constricted, darkly pigmented, often with thin-

walled pale bands. Ascus extenditunicate, paraphyses evanescent. Ascocarp

superficial on external mycelium, globose, glabrous, astomatic, often setose.

Mycelium brown, superficial, exhyphopodiate, forming stomopodia, setose,

furcated.

8.1.Meliolina malacensis (Saccardo) Trotter in Saccardo, Sylloge Fungorum

24 (1926) 360.

-Meliola malacensis Saccardo, Bull. Orto Bot. Regia Univ. Napoli 6 (1918) 43.

Ascospores with midconstriction, 35-45 x 12-15 um. Ascus oblong-clavate,

80-90 um. Ascocarp 250-300 um; setae simple, acute, 300-350 x 5-5.5 um. The

associated mitosporic state, Briania (Reynolds, 1989), was not found.

Specimen reported: C.F. Baker Fungi Malayana 451, determined as Meliola

malacensis Saccardo.

Epifoliar Fungi of Singapore 419

Distribution: Singapore

Note: This taxon was recognized by Hansford (1961) as a Meliolina and

excluded from Meliola. This species was not accepted in the treatment of

Meliolina by Hughes (1993). Meliola and Meliolina are morphologically

different as well as distant in phylogeny. These two members of the

Pezizomycotina were phylogenetically inferred by Saenz and Taylor

(1999) to be on separate clades. Meliola is a taxon in the Sordariomycetes,

Meliomycetidae, Meliolales with a unitunicate ascus, as predicted by Luttrell

(1989), and Meliolina is in the Dothidiomycetes, Dothidomycetidae, with an

extenditunicate modification of the fissitunicate ascus (Reynolds, 1989a).

8.2. Meliolina cladotricha (Léveillé) H. Sydow & P. Sydow, Ann. Mycol. 17

(1914) 553h.

-Meliola octospora Cooke, Grevillea 11 (1882) 71.

-Meliolina octospora (Cooke) Hoéhnel, Akad.Wiss. Wien, Sitzungsber.,

Math.-Naturwiss. K1. Abt. 118 (1909) 813.

-Meliolina octospora Hohnel, Akad.Wiss. Wien Sitzungsber. Math.-

Naturwiss. KI]. Abt 1. 128 (1919) 557.

Ascospores with hyaline band in terminal cells near septum, 45-57 x 17-

20 um. Ascus with abundant paraphyses, 100-120 um. Ascocarp spherical,

400um diameter, setae 145-186 um.

Specimen reported: A Flora of Singapore collection (SING 96897) made

by an unknown collector in 1899 was labeled as Meliola octospora Cooke.

There are no spores.

Note: Hughes (1993) argues for the acceptance of M. octospora Hohnel

based on his interpretation of Hohnel’s (1909, 1919) observations and

supposed prior misuse by Stevens (1927), Hansford (1954), Yamamoto

(1957) and Sivanesan (1984). We find Hughes’ argument to be suspicious

in that the attempt to affirm the observations of von Hohnel was based on

his inadequate material now curated in Herbarium FH and assumptions

made from the observations of Yamamoto (1957) from Taiwan, but without

surviving specimens.

9. Micropeltella Montagne, Ann. Sci. Nat. Bot., sér 2, 17 (1842) 325.

Ascospores hyaline, 2-8 septate, clavate. Asci clavate, cylindrical, fissitunicate,

aparaphysate. Ascocarp orbicular, dimidiate-scutulate, margin pelluculoae,

blue to green-black, textura meandriforme, plechymatous; ostiole distinct,

round. Mycelium absent at ascocarp maturity.

420 Gard. Bull. Singapore 61 (2) 2010

Batista (1959) noted that the species of this taxon were removed from

Micropeltis because of a lack of paraphyses, based on the examination of

Singapore specimens cited here. The type specimen is Montagne 1134, Herb.

9.1 Micropeltella marginata (Montagne) Batista, Inst. Micol. Univ. Recife

Publicagao 56 (1959) 160.

-Micropeltella albo-marginata (Spegazzini) Batista, Inst. Micol. Univ. Recife

Publicacao 56 (1959) 153; -Micropeltis albo-marginata Spegazzini, Bol. Acad.

Nac. Ci. Argent. 11 (1889) 572.

-Micropeltella orchidearum (P. Hennings) Batista, Inst. Micol. Univ. Recife

Publicagao 56 (1959) 163.

-Micropeltis albo-ostiolata P. Hennings, Hedwigia 47 (1908) 268.

-Micropeltis ekmanii Petrak & Ciferri, Ann. Mycol. 30 (1930) 205.

-Micropeltis marginata Montagne, Ann. Sci. Nat. (Paris) 4, 3 (1855) 133.

-Micropeltis orchidearum P. Hennings In: A. Engler & K. Prantl, Die

naturlichen Pflanzenfamilien 23 (1897) 286.

Ascospore 87-125 x 12-15 um. Ascus fissitunicate. Ascocarp dimidiate,

- plectenchymatic, blue- black, ostiolate, orbicular, 535-700 um diameter.

Specimens reported: C.F. Baker Fungi Malayana 460; see also SING 36128,

36129. CF Baker

Malayana #460 is determined as Micropeltis marginata Montagne

Illustration: Batista, 1959: 88, Fig. 18.

Distribution: Singapore.

10. Micropeltis Montagne Ann. Sci. Nat. Bot. sér 2, 17 (1842) 122.

Ascus paraphysate, fissitunicate. Ascospores hyaline, 2-many septate, clavate,

oblong to cylindrical. Ascocarp dimidiate-scutate, parenchymatic, circular,

ostiolate, green-black. Mycelium absent.

10.1. Micropeltis trimera Saccardo, Bull. Orto Bot. Regis Univ. Napoli 6

(1918)ySil:

Ascospores fusoid, 2-3 septate, 35 x 8um. Ascocarp margin pelliculose, black,

510-610 um diameter.

Specimen reported: C.F: Baker Fungi Malayana 461, holotype in BP and

isotype, UC. The UC specimen has an ascocarp measuring 350-421 tm in

diameter; the immature asci measured 100 um in length; the ascospores are

immature.

Epifoliar Fungi of Singapore 421

Illustration: Batista 1959: 146, Fig. 60.

Distribution: Singapore.

11. Polychaeton (Persoon) Léveillé

-Fumago {subgenus] Polychaeton Persoon, Mycologica europea. Sectio

prima. Erlangae (1822)

-Morfea Roze Bull. Soc. Bot. France 14: (1868) 15.

-Morfea (G. Arnaud) Ciferri & Batista (1963) 140.

-Subgenus Morfea G. Arnaud (1913) 280.

Lectctype: Polychaeton quercinum (Persoon) O. Kuntze (1891) =Fumago

quercina Persoon, Mycologica europea. Sectio prima. Erlangae (1822). Type:

Herb. L.

Mitospores formed in an expanded basal or midway centrum within a wide

stalk. Stalk single, or branched with a columnate extension with a fringed

ostiole. Mitospores unicellular, hyaline.

11.1. Polychaeton artocarpi (Batista, Nascimento & Ciferri) Khodap,

Rostaniha 7 (2006) 79.

-Microxiphium artocarpi Batista, Nascimento & Ciferri in Batista & Ciferri,

Quaderno 31 (1963)114.

-Microxiphium footi (Berkeley & Desmaziéres) Spegazzini 1918.

-Microxiphium jafarnizamie Manocharachar (1979) Nova Hedwigia 63:185.

-Microxiphium jambosae Batista apud Batista & Ciferri, Quaderno 31

(1963) 132.

-Microxiphium leptospermi Fisher, Proc. Royal Soc. Victoria, N.S.W. 45

(1963) 191.

-Microxiphium obtusulum Saccardo Phil. J. Sci. (1921) 19: 602.

-Microxiphium secundum Batista & Ciferri, Quaderno 31 (1963) 136.

Mitospores measuring 2 x 5 um.

Note: Lim (1975) listed three Singapore collections of Microxiphium

artocarpi, M. leptospermi,and M. secundum.

11.2. Polychaeton brasiliense (Batista in Batista & Ciferri), comb. nov.

Basionym: Microxiphium brasiliense Batista in Batista & Ciferri, Quaderno

31 (1963) 119.

-Microxiphium alanqii Agarwal & Sharma, J. Indian Bot. Soc. 53(1-2) (1974)

fap

-Microxiphium ciliolatum (Saccardo) Ciferri & Batista apud Batista &

422 Gard. Bull. Singapore 61 (2) 2010

Ciferri, Quaderno 31 (1963) 121.

-Microxiphium coffeanum Batista & Matta, Quaderno 31 (1963) 122.

-Microxiphium inspersum Batista & Ciferri, Quaderno 31 (1963) 132.

-Microxiphium spathodeae Batista & Matta apud Batista & Ciferri,Quaderno

31(1963)137-

-Microxiphium unedonis (Marie & Saccardo) Batista & Ciferri, Quaderno

31° G963) 139:

-Polychaeton bassiae Manocharachary, Kunwar, Sarath & Nagamani in

Manocharachary, Kunwar, Babu & Nagamani. 2003.

Mitospores measuring 2 x 3.5 um.

Note: Lim (1975) listed two Singapore collections of Microxiphiumcoffeanum

and M. spathodeae.

11.3. Potychaeton tenellum (Saccardo), comb. nov.

Basionym: Microxiphium tenellum Saccardo, Bull. del’orto Bot. della R.

Univ. Napoli 24 (1918) 5.

-Microxiphium aciculiforme Ciferri, Batista & Nascimento in Batista &

_ Ciferri, Quaderno 31 (1963) 110.

-Microxiphium atmosphaericum Batista, Bezerra & Garnier in Batista &

Ciferri, Quaderno 31 (1963) 117.

-Microxiphium columnatum Batista, Ciferri & Nascomento in Batista &

Ciferri, Quaderno 31 (1963) 123.

-Microxiphium cylindricum Batista & Ciferri, Quaderno 31 (1963) 127.

-Microxiphium dubium Saccardo, Ann. Mycol. 13 (1915)127.

-Microxiphium philippinensis Ciferri & Batista in Batista & Ciferri,

Quaderno 31 (1963) 135.

-Microxiphium virde Batista & Ciferri, Quaderno 31 (1963) 141.

-Polychaeton pinicola Batista, Nascimento & Ciferri apud Batista & Ciferri,

Quaderno 31 (1963) 135.

-Polychaeton purpuraefaciens (J.F.H. Beyma) Reynolds & Gilbert, Aust.

Syst. Bot. 18(3) (2005) 275.

Mitospores measuring 1.5 x 5 um.

Specimens reported: C.F. Baker Fungi Malayana 468, holotype at BP; isotype,

UC; Singapore, Bukit Timah Reserve, Oct 2007, DRR Singapore 20 (SING,

UC). Lim (1975) also listed two Singapore collections of this species as

Microxiphium aciculiforme, M. columnatum.

Illustrations: Beyma, 1931; Batista & Ciferri, 1963; Ciferri et al., 1956.

Distribution: Indonesia, Philippines, Singapore, Taiwan, Thailand, Vietnam.

ae mrt 423

Epifoliar Fungi of Singapore yar)

Note: Forty six species have been described as Microxiphium

(=Microxyphium) (Harvey) ex Berkeley & Desmazieres in Thiimen (1879).

Most of these are synonyms of Polychaeton.

Yamamoto (1955) noted a “Microxyphium (Certatopycnidium) type”

in the “imperfect stage,” described as, “A peridium is semispherical...”

Reynolds and Gilbert (2005) found that Microxiphium is based on two

mitosporic forms. The mycelium forms a setose pellicle and mitosporic

reproduction is twofold: 1. a rosette of hialidic cells found in the mycelium

or as a cortex of hyphae surrounding the mycelia setae (Microxiphium);

2. subpellicular areas producing triradiate, septate mitospores (Bisbyopeltis

Phoebesii).”

The pycnidium of Microxiphium sensu Batista & Ciferri (1963b) is the

source of a few characters that are used to distinguish 28 taxa. It is described

as cylindric. A fimbriated appearance is given to the apex by hyaline hyphal

extensions beyond the darkly pigmented cells forming the apical dispersal

column. The taxa recognized range in a continuous progression from an

attenuated form to one with asomewhat attenuated lower area that continues

as an extended neck formed around a canal through which the mitospores

are dispersed from the basal conidiogenous centrum. A continuum of

character variability can be found in an examination of the types and other

collections of Microxiphium spp. in Herbarium URM and in pure culture

isolates derived from a single mitospore (unpublished research).

Polychaeton forms a mitosporic centrum within a wide stalk, located

from the central to the upper part. Conidioxyphium Batista and Ciferri was

found to be indistinguishable microscopically (Singh er a/.,2006) with cluster

analysis, principal components analysis, and discriminate analysis (Faull er

al., 2002). Arnaud (1913) considered these structures to be spermagonia.

Beyma (1931) gives details of development in culture. The conidiogenous

hyphae of the centrum are sympodially branched, forming phialides with

collarettes. The stalks may be single or branched; the branching often

arises from mitospores germinating directly on the stalk. A columnate

extension beyond the centrum often has a fringed ostiole. The mitospores

are unicellular and hyaline. Observations (Reynolds, 1978) made on the

mitosporic structures of Scorias spongiosa and collections from tropical

climates indicate that the pycnidia of Polychaeton are variable in length

and branching patterns. The fruit body stalk that subtends the mitosporic

centrum is characteristically as wide as the centrum. The mitospore

morphology is stable and produced within an internal pycnidial centrum.

The hyaline, unicellular spore is exuded from the centrum along a canal

424 Gard. Bull. Singapore 61 (2) 2010

in a columnate neck. A droplet of water soluble matrix contain multiple

conidia forms under humid conditions. The mitospores flow down the

fruit body onto a substrate; dilution of the spore containing drop initiates

germination. Typically the mitospore divides to form a hyphal initial and

the melanoid pigmentation accumulates in the cell walls. Measurements of

the mitospores as they are exuded from the apex of the fruit body comprise

primary taxonomic characters.

The name Morfea has been utilized for both a concept of mitosporic

and ascosporic monomorphic and pleomorphic species by Roze (1868) and

Batista and Ciferri, (1963a). The name was clarified as a mitosporic nomen

and affirmed as Polychaeton (Hughes, 1976; Punithalingam, 1981; Reynolds,

1998).

12. Schizothyrium Desmazieres, Ann. Sci. Nat. Bot. sér 3, Bot. 11 (1840)

360.

Ascospores |-septate, hyaline; Ascus extenditunicate; ascocarp dimidiate,

dark, with radial margin; Mycelium inconspicuous.

12.1. Schizothyrium longispora (Patouillard) Arx, in Miller & Arx, Die

Gattungen der didymosporen Pyrenomycetes (1962) 202.

-Dictyopeltis lucumae Batista, Anais IV. Congresso Nac. Soc. Bot. Brazil

i953) 07:

-Eremotheca philippensis Sydow, Ann. Mycol.15 (1917) 235,

-Microthyriella enaequalis Batista, Mycopath. Mycol. Appl. 5 (1951) 171.

-Microthyriella guianensis Stevens & Manter, Bot. Gaz. 79 (1925) 290.

-Microthyriella macrospora Hohnel, Akad.Wiss. Wien, Sitzungsber. Math.-

Naturwiss. KI. Abt. I, Biologie 127 (1918) 630.

-Microthyrium browneanum Saccardo, Bull. Orto Bot. Regis Univ. Napoli 6

(1918) 50.

-Microthyrium lomgisporum Patouillard, Bull. Soc. Mycol. France 20 (1888)

118.

Ascospores 35-60 x 12-17 um. Asci 60-75 x 40-50 um. Ascocarp superficial,

roundish, 400-1200 um diameter, dark brown.

Specimen reported: C.F. Baker Fungi Malayana 465. Holotype at UC.

Distribution: Singapore.

13. Setameliola, gen. nov

=Meliola sensu Mibey & Hawksworth (1997)

Sporidia 3-4, septata, fusca. Asci con 2-4 spori, aparaphysati. Ascocarp in

Epifoliar Fungi of Singapore 425

mycelio maculiformi superficiali, globosa, astoma, setosus. Mycelium ex

hyphis brunneis, septatis vel hyphis formans catillus, cum hyphopodia, setosus.

- Typus species: Setameliola argentina (Spegazzini) D.R.Reynolds

Diagnosis: This taxon is distinguished by the presence of setae on the

perithecium as well as the mycelium. Meliola Fries emended Bornet (1851)

sensu Hansford (1961) includes species that have glaborous as well as setose

ascocarp. The Mibey and Hawksworth (1997) Meliola emendment provides

for a perithecium “devoid of setae or appendages.” Accordingly, Setameliola

includes the 23 species of Meliola sensu lato with setose ascocarp. The

mycelial setum is the character distinction of Meliola and Setameliola from

Trenopsis (setose ascocarp), Appendiculella (laviform ascocarp appendages)

and Asteridiella (glabrous ascocarp). Two of the 24 species of Setameliola

(S. araucariae and S. agathidis) form unusual thalloid plates (Ellis, 1974;

Hansford, 1961; Katumoto & Hosagoudar, 1989).

13.1 Setameliola argentina (Spegazzini), comb. nov.

Basionym: Meliola argentina Spegazzini, Ann. Soc. Cient. Argentina 9 (1880)

177. [Holotype: Speggazini 510, S].

-[reniopsis martiniana (Gaillard) Stevens, Ann. Mycol. 25 (1927) 437

-Meliola argentina Spegazzini var hawaiiensis Hansford, Sydowia 11 (1958)

Sik:

-Meliola argentina Spegazzini var africana Hansford, Sydowia 9 (1955) 9.

-Meliola argentina Spegazzini var leeuwenii Hansford, Sydowia 10 (1957) 63.

Type: BO 5607.

-Meliola circinans Earle, Bull. New York Bot. Gard. 3 (1905) 304.

-Meliola circinans Earle var rhynchosporae Hansford (1904) 304.

-Meliola intricata Sydow, Philipp. Journ. Sci. Bot. (1913) 268. Type: PBS

TAS2

-Meliola juddiana Stevens, Bernice B. Bishop Mus. Bull. 19 (1925) 32.

-Meliola martiniana Gaillard, Le Genre Meliola (1892) 68.

-Meliola setulifera (Spegazzini) Stevens, Ann. Mycol. 25 (1927) 285. Type:

Rabenhorst and Winter. Fungi Europaei et Extraeruopaei 3852 (Herb. F).

Ascospores oblong, obtuse, 4-septate, 37-47 x 15-18 um. Ascocarp to 190

um diameter, with setae straight to acute, 90 x 8 um. Mycelium epiphyllous,

dense, velvety; hyphae sub-straight to undulate, branching opposite; capitate

hyphopodia alternate to opposite, head cell straight or bent, stalk cell

cuneate; mycelial setae, simple with acute apex, entire, 400 um length.

Illustrations: Hansford, 1963; Stevens, 1928: 381, Plate [V-40.

Distribution: Borneo, Philippines, Singapore, Taiwan

426 Gard. Bull. Singapore 61 (2) 2010

The following recombinations are made on the basis of the setose ascocarp:

13.2. Setameliola agathidis (Ellis), comb. nov.

Basionym: Meliola agathidis Ellis, Trans. Brit. Mycol. Soc. 63 (1974) 96. Type:

IMI 73853.

13.3.Setameliola apiculata (Hansford), comb. nov.

Basionym: Meliola apiculata Hansford, Proc. Linn. Soc. London 160 (1948)

137) Type: Herb ILL:

13.4. Setameliola araucariae (Ellis), comb. nov.

Basionym: Meliola araucariae Ellis, Trans. Brit. Mycol. Soc. (1974) 93. Type:

IMI 170578.

13.5. Setameliola artocarpiicola (Stevens ex Hansford), comb. nov.

Basionym: Meliola artocarpticola Stevens ex Hansford, Sydowia Beih. 11

(1958) 52. Type: PBS 36433.

13.6.Setameliola bayamonensis Tehon var guettardae (Ciferri), comb. nov.

Basionym: Meliola bayamonensis Tehon var guettardae Ciferri, Mycopath.

Mycol. Appl. 7(1954) 98. Type: Herb. S.

13.7. Setameliola brinkii (Hansford), comb. nov.

Basionym: Meliola brinkii Hansford, Reinwardtia 3 (1954) 96. Type: BO

123i;

13.8. Setameliola canariicola (Stevens ex Hansford), comb. nov.

Basionym: Meliola canariicola Stevens ex Hansford, Sydowia Beih. | (1957)

102. Type: PBS 34009.

13.9. Setameliola canthii Hansford var aristata (Hansford), comb. nov.

Basionym: Meliola canthii Hansford var aristata Hansford, Proc. Linn. Soc.

Lond. 51 (1945) 22. Type: Herb. IMI.

-Meliola woodiana Saccardo var aristata Hansford, Journ. Linn. Soc. Bot.51

(1937) 284.

13.10. Setameliola circinans (Earle), comb. nov.

Basionym: Meliola circinans Earle, Bull. New York Bot. Gard. 3 (1905) 304.

Type: Heller 6347, NY.

13.11.Setameliola circinans Earle var rhynchosporae (Hansford) comb. nov.

Basionym: Meliola circinans Carle var rhychosporae Hansford, Sydowia 9

Epifoliar Fungi of Singapore 427

(1955) 13. Type: Nash 1803, FLS.

13.12. Setameliola coriae (Hueguenin), comb. nov.

Basionym: Meliola coriae Hueguenin, Rev. Mycol. (Paris) 34 (1969) 30. Type:

Herb. P.

13.13. Setameliola cyperi (Patouillard in Gaillard), comb. nov.

Basionym: Meliola cyperi Patouillard in Gaillard, Le Genre Meliola (1892)

70 bype: Herb: FH.

13.14. Setameliola fusispora (Yamamoto), comb. nov.

Basionym: Meliola fusispora Yamamoto, Trans. Nat. Hist. Mus. Taiwan 31

(1941) 219. Type: Herb. TNS.

13.15. Setameliola kaduae (Stevens), comb. nov.

Basionym: Meliola kaduae Stevens, Bernice B. Bishop Mus. Bull. 19 (1925)

SUstype: Herb. ILL.

13.16. Setameliola martiniana (Gaillard), comb. nov.

Basionym: Meliola martiniana Gaillard, Le Genre Meliola (1892) 68. Type:

Rabenhorst’s Fungi Europea 3852 (Herb. F).

-[renopsis martiniana (Gaillard) Stevens, Ann. Mycol. 25 (1927) 437.

-Meliola perseae Stevens forma setulifera Spegazzini. Bol. Acad. Nac. Cien.

Cordoba 26 (1923) 380.

-Meliola setulifera Stevens, Ann. Mycol. 26 (1928) 285.

13.17. Setameliola microtricha (Sydow in Sydow & Sydow), comb. nov.

Basionym: Meliola microtricha Sydow in Sydow & Sydow, Ann. Mycol.18

(1920) 157. Type: Herb. UC.

The Singapore record for this taxon is CF Baker Fungi Malayana 490

determined as Tetrachia singularis Saccardo (1918). The taxon is cited

by Hansford (1961) as the type under Meliola microtricha H. Sydow & P.

Sydow [see Annales Mycologia 18 (1920) 157]. The Saccardo description is

of a sporodochial fungus and only conidia are described. Sydow & Sydow

(1920) regarded Tetrachia singularis as a synonym of Spegazzinia meliolae

Zimmerman. Damon (1953) considered that Spegazzinia tessartha (B & C)

Saccardo comprised a monotypic genus and cited 7? singularis as asynomym

of Isthmospora trichophila (Atkinson) Damon. Hughes (1953) considered

T. singularis to be a synonym of Trichothrium.

The UC specimens 490 and 49/ (determined as T. singularis, n.

gen. et n. sp.) from the C.F. Baker Fungi Malayana exsiccatum exhibit

428 Gard. Bull. Singapore 61 (2) 2010

two epifoliar fungi. The isthmospores of 7) singularis or T: asterophorum

sensu Hughes (1953) similar to those of Spegazzinia, are found in both

specimens. Tetrachia singularis is a hyperparasite of a second fungus

present that agrees with the Hansford (1961) description of Setameliola

(=Meliola) microtricha. No thyriothecia of JT. singularis were seen.

13.18. Setameliola paratrophidis (Hansford), comb. nov.

Basionym: Meliola paratrophidis Hansford, Reinwardtia 3 (1954) 96. Type:

Herb. BO.

13.19.Setameliola pradosiae (Batista in Batista, Nascimento & Maia), comb.

nov.

Basionym: Meliola pradosiae Batista in Batista, Nascimento & Maia, Inst.

Micol. Publicacao 25 (1956) 8. Type: Herb. URM.

13.20. Setameliola pseudomori (Hansford), comb. nov.

Basionym: Meliola pseudomori Hansford, Proc. Linn. Soc. New South Wales

(1954) 65. Type Herb. IMI.

13.21. Setameliola sakahensis (Yamamoto), comb. nov.

Basionym: Meliola sakahensis Yamamoto, Trans. Nat. Hist. Soc. Taiwan 30

(1940) 421. Type: Herb. BPI.

14. Trichomerium Spegazzini emend. Reynolds, Mycotaxon 14 (1982) 190.

-Capnobatista Ciferri & Leal in Batista,A.C. & R. Ciferri, Saccardoa 2 (1963)

WD:

-Chaetopotius Batista, Mycopath. Mycol. Mycol. Appl. 5 (1951) 151. Pro

parte

-Triposporiopsis Yamamoto, Special Publ. Kasai Shuppan Institute Minato-

ku Tokyo (1955) 55.

Ascospores hyaline, fusiform, 18-32 x 5-10 um. Hymenium periphysate. Asci

fissitunicate, obclavate.

Ascocarp dark brown, setose, ostiolate 80-230 um. Mycelium of superficial,

branching, septate hyphae, not setose nor hyphopodiate

14.1 Trichomerium grandisporum (Ellis & Martin in Ellis & Everhart) Batista

& Ciferri, Saccardoa 2 (1963) 210 (see Reynolds, 1982, for synonyms).

Ascospores hyaline, fusoid, granulate, 4-6 septate, 18-38 x 5-10 um. Asci

aparaphysate, fissitunicate. Ascocarp ostiolate, 100-170 x 88-170 um; Setae

62.5-170 um in length. Mycelium of superficial, branching, septate hyphae,

brown to blackish, not setose nor hyphopodiate.

Epifoliar Fungi of Singapore 429

Specimens reported: Singapore, Bukit Timah Reserve. Oct 2007. DRR 1, 5, 6,

14; 20:22. (SING, UC).

Illustration: Batista & Ciferri 1963a: figs 75-96.

Distribution: Pantropical

Note: A specimen identified by F. von Hohnel as Aithaloderma setosum is

represented by a microscope slide preparation (/MI/ 63263, F). The sparse

data indicate that the collections were made in Singapore in 1956. An

unclear understanding of Aithaloderma species is manifested by different

interpretations of the ascocarp. In one type, the setose ascocarp is globose

similar to that of Trichomerium. Other species have been attributed with

a shield over the ascocarp that is characteristic of Chaetothyrium. We find

that the Aithaloderma examined by von Hohnel should be assigned to

Trichomerum grandisporum.

Acknowledgements

I thank the Singapore Botanical Garden for a Research Fellowship in

October of 2007; Dr. M.M. Thang, Dr. D. Pfister and Dr. Paul Kirk made

helpful suggestions.

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Gardens’ Bulletin Singapore 61 (2): 437-481. 2010 437

A Conspectus of the Lichens (Lichenized Fungi) of

Singapore

H.J.M. SIPMAN

Free University of Berlin

Botanical Garden and Museum

Konigin-Luise-Str. 6-8

D-14195 Berlin, Germany

Abstract

A total of 296 species of lichenized fungi are reported from Singapore and

presented in an annotated list with local distributional information. It is

based on herbarium and literature study and the fieldwork done in the year

2000. Unidentified samples suggest the figure to be an underestimation,

while some of the listed species may have become extinct. Lists of synonyms

and collectors are added.

Introduction

Tropical conurbations, the world’s most fast-growing habitat, have turned

out to harbour significant numbers of lichenized fungi. Aptroot and Seaward

(1999) and Aptroot and Sipman (2001) report no less than 308 species for

the city of Hongkong. Singapore seems particularly suitable for a study of

lichens in an urbanized tropical area because it has received regular attention

from botanists during its development from primary lowland forest with

small settlements ca 1800 to extensive plantations a century later and to the

present urbanized area with large built-up high rise areas interspersed by

parks and secondary forest. During 1800-1964 scattered lichen collections

were made by visiting and resident general botanists, e.g., E. Almquist, O.

Beccari, T.R. Chipp, Kiah, A.M. Lemaitre, A.C. Maingay, H. Moller. Their

collections have been investigated and published by, e.g., Krempelhuber

(1875, 1877), Nylander and Crombie (1884) and Miiller Argoviensis (1893).

From 1964-1992 three lichen experts visited the area and made herbarium

vouchers, Aptroot, Degelius, and Tibell (see list of collectors below). In

November 2000 the author made a lichen inventory in collaboration with

Dr. B.C. Tan (NUS/SING) and Prof. D.H. Murphy (Singapore), and took

samples from 18 study sites spread over Singapore Island and on some of

the smaller off shore islands. Presented below is an evaluation of the present

day lichen flora of Singapore. A comparison with temperate urban areas

438 Gard. Bull. Singapore 61 (2) 2010

and a discussion of probable changes in the island’s lichen flora through the

centuries are presented in another paper (Sipman, in press).

Material and methods

The evaluated records originated mainly from fieldwork conducted in 2000

by the author in collaboration with Dr. B.C. Tan, Prof. D.H. Murphy and

Ms Farida then at the National University of Singapore. This yielded 962

specimens of lichenized fungi and 1,126 records when taking into account

mixed specimens and field observations. Where possible the names of the

phorophytes of lichens collected were noted, and in the Botanical Garden

also the tree ID number. The specimens are deposited in the herbaria of

B and SING and the data of the B set are available in the web database -

http://www.bgbm.org/scripts/ASP/lichcol/query.asp. For a list of the visited

localities and habitats see Table 1. In addition, the lichenological literature

was searched for lichen records from Singapore, and relevant specimens

were also borrowed from the herbaria SING and UPS, and the private

collections of A. Aptroot, P. Diederich and F. Schumm. Additional collections

~ were found notably via the herbarium database of UPS (http://www-hotel2.

uu.se:8888/cgi-bin/wwwdrive.fytotek/beginner). All indicated specimens

were examined by the author, unless otherwise stated. The specimens were

investigated in the usual way by stereomicroscope and photomicroscope,

and selected specimens were analysed by TLC (Orange et al., 2001).

Table 1. List of localities of the fieldwork by the author in 2000 with codes used in the species

list (in bold), collection numbers and habitat information.

1 - Sungei Buloh Nature Park, on N-coast, opposite Johor Baru. At E-margin of

reserve. Mangrove forest ca 10 m tall. Elev. ca 1 m. Coord. 1° 27’ N, 103° 44°

E, 4 Nov 2000 - H. Sipman & D.H. Murphy 45441-45514.

1. Away from the coast. On Allophylus cobbe branchlet. la. On Rhizophora

apiculata trunk/Foliicolous in undergrowth, on Brownlowia tersa leaves/on

Xylocarpus granatum leaves/on Asplenium nidus leaves/On Rhizophora

apiculata trunk. 1b. On Xylocarpus granatum trunk. le. Away from the

coast. On Excoecaria agallocha trunk. Id. Away from the coast. On Hibiscus

tiliaceus branches. le. At the coast. On Sonneratia alba trunk. If. On Avicennia

officinalis trunk (Nr. 45477 on A. alba in splash zone). 1g. On Bruguiera

cylindrica trunk. 1h. On Avicennia alba trunk. li. On Excoecaria agallocha

trunk. 1j. Land-side. On Excoecaria agallocha trunk.

2 - Campus of National University of Singapore, around Kent Ridge, parkland

with scattered buildings and roads. Elev. ca 50 m Coord. 1° 18’ N, 103° 45.5’ E,

5 Nov 2000 - H. Sipman 45515-45637.

2. On thin palm stem (Ptychosperma macarthurii)/On trunk of leguminose

A Conspectus of the Lichens (Lichenized Fungi) of Singapore 439

tree/Kent Ridge Road. On weathered siliceous stone/On low concrete ridge

along road, shady, on top/On soil of steep bank of path, open, over weathered

stone/Kent Ridge Road. On branches of Erythrina crista-galli shrub. 2a. On

small tree of Bauhinia purpurea, on trunk ca 20 cm diam. 2b. On ca 20-30

cm diam. trunk of small Cassia fistula tree. 2e. On ca 10-30 cm diam. Acacia

auriculiformis trunk. 2d. On ca 50-100 cm diam. Samanea saman trunk. 2e.

Kent Ridge Road. On Peltophorum pterocarpum trunk. 2f. Kent Ridge

Road. On Khaya sp.(Meliaceae) trunk ca 40 cm diam. 2g. Kent Ridge Road.

On branches of Callistemon sp. treelet/On Roystonea regia trunk/On Cassia?

trunk/On trunk.

3 - Singapore Botanic Gardens, parkland with scattered trees and shrubs. Elev. ca 50

m Coord. 1° 18’ N, 103° 48° E, 7-9 Nov and 10 Nov 2000 - H. Sipman 45638-

45814, 45836-45855.

3. On trunk./(45814) On lava stone in half shade of shrub in succulent

garden./Epiphyte./On concrete roadside/On soil on ant heap in lawn. 3a. On

50 cm diam. Peltophyllum pterocarpum (nr. 5500) trunk. 3b. On 25 cm diam.

Archontophoenix alexandrae (nr. 10151) trunk. 3e. On 15 cm diam. Phoenix

rupicola (nr. 5315) trunk. 3d. On 15 cm diam. Atalantia monophylla (nr.5322)

trunk. 3e. On 50 cm diam. Tectona grandis (nr. 5325) trunk. 3f. On 150 cm

diam. short Calophyllum inophyllum (nr. 5323) trunk. 3g. On ca 25 cm diam.

Lepisanthes rubiginosa (nr. 5301) trunk. 3h. On ca 20 cm diam. Azadirachta

indica (nr. 5313) trunk. 3i. On ca 60 cm diam. Hevea brasiliensis (nr. 5305)

trunk. 3j. On ca 30 cm diam. Jacaranda obtusifolia ssp. rhombifolia (nr. 5498,

planted 1968) trunk. 3k. On ca 30 cm diam. Podocarpus falcatus (nr. 5503)

trunk. 31. On ca 40 cm diam. Podocarpus neriifolius (nr. 5513) trunk. 3m.

On ca 20-40 cm diam. tree trunks and branches of Juniperus chinensis (nr.

5488, 5489, 5379). 3n. On ca 35 cm diam. Podocarpus rumphii trunk (nr. 5286

planted 1942, 5287). 30. On ca 30 cm diam. slanting Majidea zanguebarica

(nr. 5497, planted 1955) trunk. 3p. On ca 30 cm diam. Libocedrus macrolepis

var. formosana (nr. 5385) trunk. 3g. On ca 30 cm diam. Podocarpus gracilior

(nr. 5389, planted 1932) trunk. 3r. On ca 30 cm diam. Swietenia macrophylla

(nr. 5983) trunk. 3s. On ca 20 cm diam. 10 m tall palm stem of Scheelea

insignis (nr. 5054). 3t. On ca 40 cm diam. Mangifera caesia (nr. 5062) trunk.

3u. On ca 40 cm diam. Podocarpus neriifolius trunk. 3y. On ca 25 cm diam.

branchy trunk of small Pithecellobium dulce (Nr. B-72). 3w. On ca 100 cm

diam. Fagraea fragrans trunk. 3x. On ca 100 cm diam. Tetrapleura thonningii?

(near Nr. 6002) trunk. 3y. On ca 100 cm diam. Michelia alba (Nr. 6516) trunk.

On ca 100 cm diam. Michelia alba (Nr. 6516) trunk. 3z. On ca 40 cm diam.

Cassia fistulosa (Nr. 09354A) trunk. 3aa. On ca 70 cm diam. Peltophorum

pterocarpum trunk. 3ab. On trunks and branches of Plumeria. 3ac. On ca

100 cm diam. Carapa guianensis (Nr. 6007) trunk, between the butresses. 3ad.

On ca 20 cm diam. Podocarpus sp. trunk. 3ae. On ca 80 cm diam. Samanea

saman (Nr. 5968) trunk. 3af. On ca 35 cm diam. Quercus bambusaefolia (Nr.

B/136/36/6, planted 1935) trunk. 3ag. On ca 30 cm diam. Artocarpus altilis

(Nr. j/184/93/897) trunk. 3ah. On ca. 100 cm diam. trunk, between buttresses.

3ai. On ca. 20-30 cm diam. Maniltoa browneoides trunk. 3aj. On ca 70 cm

440)

Gard. Bull. Singapore 61 (2) 2010

diam. 15 m tall palm stem of Roystonea oleracea near herbarium entrance.

3ak. On ca 40 cm diam. Heritiera alata (Nr. 09332G) trunk, on buttresses.

3al. On ca. 30 cm diam. trunk. 3am. On ca 25 cm diam. palm stem of Phoenix

sylvestris (Nr. K/00/7094). 3an. On ca 20 cm diam. stem of tall palm. 3ao0. On

ca 15 cm diam. young Shorea curtisii (Nr. K99/95/4990A ) trunk. 3ap. On base

of palm stem of Phoenix loureirii. 3ar. On ca 60 cm diam. Tamarindus indicus

(Nr. XH 26) trunk. 3as. On ca 80 cm diam. trunk. 3at. On ca 25 cm diam.

Shorea fa. apuetiana (Nr. XH 64) trunk. 3au. On ca 20 cm diam. branch of

Eugenia brasiliensis shrub (Nr. 19970843 A 2). 3av. On Plumeria dwarf trees.

3ax. On ca 80 cm diam. Fagraea fragrans (Nr. XH-11) trunk./On ca 80 cm

diam. Fagraea fragrans (Nr. XH-11) trunk. 3ay. On ca 60 cm diam. Myristica

lowiana (Nr. XH-79) trunk. 3az. On ca 100 cm diam. Pterocarpus indicus (Nr.

XH-26) trunk. 3ba. On ca 80 cm diam. trunk.

4 - Singapore Botanic Gardens, Rainforest Reserve. Elev. ca 50 m Coord. 1° 18’ N,

103° 48° E - 7-9 Nov 2000 - H. Sipman 45815-45835.

On lower part of Castilla elastica trunk 50 cm diam./On lower part of Dyera

costulata trunk 100 cm diam./On lower part of small tree trunk./Foliicolous

in undergrowth.

5 - SE side of MacRitchie Reservoir. Elev. ca 50 m Coord. 1° 21’ N, 103° 50’ E - 12

Nov 2000 - H. Sipman & B.C. Tan 45856-45944.

5a. Parkland with scattered trees and shrubs. On ca 120 cm diam. Peltophorum

pterocarpum trunk. 5b. Parkland with scattered trees and shrubs. On ca 100

cm diam. Mangifera trunk. 5¢e. Parkland with scattered trees and shrubs. On

ca 40 cm diam. tree trunk of leguminose. 5d. Parkland with scattered trees

and shrubs. On ca 70 cm diam. Calophyllum inophyllum trunk. 5e. Secondary

forest with primary forest remnants. On 10 cm diam. Calophyllum trunk at

forest margin, within reach from the soil/On 10 cm diam. tree trunk at forest

margin, within reach from the soil/On tree trunk, within reach from the soil/

On loamy bank of path at forest margin on lake shore/Secondary forest with

primary forest remnants. On Pandanus leaves in undergrowth/On leaves in

undergrowth. 5f. Secondary forest with primary forest remnants. On tree

trunk in gap (Macaranga macrophylla 10-15 cm diam.), within reach from

the soil.

6 - Bukit Timah Nature Reserve, slightly disturbed primary forest. Elev. ca 50-80 m

Coord. 1° 21’ N, 103° 41’ E - 13, 15,24 Nov 2000 - H. Sipman & D.H. Murphy

45945-45985, H. Sipman & Farida 46076-46114, H. Sipman & B.C. Tan 46379-

46383.

6. SE-side, Taban valley. On trunks within reach from the ground/On leaves

in undergrowth/On Pandanus leaves in undergrowth/On Streblus elongatus

leaves in undergrowth/E-side, along Cave Path and Rock Path. On 20 cm

diam. Adinandra dumosa trunk within reach from the ground/On Macaranga

triloba trunk within reach from the ground/On tree trunk within reach from

the ground/E-side, Rock Path. Slightly disturbed primary forest. On ca 50

cm diam. trunk of fallen tree. 6a. E-side, along Cave Path and Rock Path. On

leaves in undergrowth.

7 - Southern islands: Lazaro Island. Secondary scrub and beach forest of Terminalia

A Conspectus of the Lichens (Lichenized Fungi) of Singapore 44]

and Casuarina. Elev. ca 1 m Coord. 1° 13’ N, 103° 51.5’ E - 14 Nov 2000 - H.

Sipman & B.C. Tan 45986-46048.

7. On trunk in secondary scrub/On siliceous stone on clearing in beach forest/

On loamy soil of termite heap in road bank/On loamy soil of road bank/On

rusty iron in clearing/On humid, shady concrete. 7a. On Terminalia catappa

trunk, among buttresses./On Terminalia catappa trunk. 7b. On Casuarina

equisetiolia trunk.

8 - Southern islands: Kusu Island. Cleared beach forest with scattered Casuarina,

Terminalia and planted trees. Elev. ca 1 m Coord. 1° 13’ N, 103° 52’ E- 14 Nov

2000 - H. Sipman & B.C. Tan 46049-46075.

On Casuarina equisetifolia trunk within reach from the ground/On trunk

within reach from the ground/On trunk within reach from the ground, in

fissure/On Terminalia catappa trunk within reach from the ground.

10 - Nee Soon, SE of Upper Selatar Reservoir. Freshwater swamp forest. Elev. ca

20 m Coord. 1° 23.5’ N, 103° 48.5’ E - 16 Nov 2000 - H. Sipman, D.H. Murphy

& B.C. Tan 46115-46201.

10. On trunks within reach from the ground/On twig within reach from

the ground. 10a. On trunk and canopy of fallen tree. 10b. On leaves in

undergrowth.

11 - Nature trail N of Lower Peirce Reservoir. Low secondary forest with Nepenthes.

Elev. ca 20 m Coord. 1° 23’ N, 103° 49’ E - 16 Nov. 2000 - H. Sipman, D.H.

Murphy & B.C. Tan 46202-46217.

On trunks within reach from the ground.

12 - Lower Peirce Reservoir Park. Elev. ca 20 m Coord. 1° 23’ N, 103° 49’ E - 16 Nov

2000 - H. Sipman, D.H. Murphy & B.C. Tan 46218-46219. On Roystonea palm

stem along road.

13 - Fort Canning Park, in town center. Elev. ca 30 m Coord. 1° 17.5’ N, 103° 51’ E -

18 Nov 2000 - H. Sipman 46220-46202.

13. On trunks in park on hilltop. 13a. On Plumeria trunks in park on hilltop.

14 - Pulau Ubin Island, from ferry to Kampong Melayu. Abandoned gardens. Elev.

ca 5 m Coord. 1° 24’ N, 103° 58’ E - 19 Nov 2000 - H. Sipman & B.C. Tan

46263-46361.

14. On granite cliff at border of garden/On loamy bank of road/On trunk

of fruit tree (Nephelium rambutan)/On trunk/On termite heap/On roots of

betelnut palm/On trunk. 14a. On durian (Durio sp.) trunk. 14b. On trunk of

fruit tree (Lansium domesticum). 14e. On coconut stem in plantation. 14d.

On trunk of fruit tree (Psidium guava). 14e. On trunk of rubber tree (Hevea

brasiliensis) in abandoned plantation. 14f. On leaves of lower branches of

fruit trees.

15 - Labrador Park, at coast SW of city center. Elev. ca 10 m Coord. 1° 16’ N, 103°

48° E - 23 Nov 2000 - H. Sipman & D.H. Murphy 46362-46378. Secondary

forest on hilltop at the coast. On Eugenia grandis trunk within reach from

the ground/On leaves in undergrowth/On trunk within reach from the

ground/On Millettia atropurpurea trunk within reach from the ground/On

hana near the ground/On Barringtonia indica trunk/On Tabebuia trunk/On

Casuarina trunk./Scattered trees on lawns at the coast. On tree trunk./On

442 Gard. Bull. Singapore 61 (2) 2010

coastal conglomerate rock, sheltered, about 1 m above highwater level./On

underside of overhanging, ca 10 cm diam. Hibiscus tiliaceus trunk at sheltered

coast, about 2 m above highwater level.

16 - Bukit Timah Nature Reserve, N-side, abandoned quarry. Elev. ca 100 m Coord.

1° 21° N, 103° 41’ E - 24 Nov 2000 - H. Sipman & B.C. Tan 46384-46397.

Shaded, humid rockface at base of cliff.

17 - Bukit Brown Chinese Cemetery, S of MacRitchie Reservoir. Grassy vegetation

with scattered trees and grave tombs. Elev. ca 30 m Coord. 1° 21’ N, 103° 49°

E - 25 Nov 2000 - H. Sipman & B.C. Tan 46388a-46391.

On tree stump in secondary forest/On leaves of undergrowth in secondary

forest/On brick of tomb/On granite of tomb/On fallen, dry branch.

18 - Sembawan Park, on N-coast. Elev. ca 2 m Coord. 1° 28’ N, 103° 51’ E - 25 Nov

2000 - H. Sipman & B.C. Tan 46392-46412. Grassland with scattered trees and

shrubs. On trunk within reach from the ground.

Results

The accepted lichen taxa are presented in the following list. In addition

lists are given of synonyms used in past publications relating to Singapore

lichens, rejected records, and the collectors.

Alphabetical list of the lichen species reported from Singapore with comments

For each species herbarium vouchers and literature references are given.

Abbreviations: S = Sipman; SF = Sipman & Farida;SM = Sipman & Murphy;

SMT = Sipman, Murphy & Tan; ST = Sipman & Tan; obs.: field observation by

the author, without voucher; accompanying species in herbarium specimens

deposited under another name are indicated as “in [collector + number]

{Herbarium abbreviations] (= [name or taxon under which it is deposited])”.

Locality codes are given in brackets; they correspond to table 1. Pictures

of many species are available in the website http://www.bgbm.fu-berlin.de/

sipman/Zschackia/Singa/genuslist.htm

Amandinea diorista (Nyl.) Marbach — (13) S 46228 [B, SING].

Amandinea efflorescens (Miill.Arg.) Marbach — (2b) § 45523 [SING]; (2d)

S 45557 [B]; (2f) S 45615 [B]; (2g) S 45626 [B], 45637 [B], 45636 [SING] c.

apoth.; (3) obs.; (7b) ST 46005 [SING; (8) ST 46052 [B, SING]; (13) obs.;

(14) obs.; (15) obs.; corticola prope Singapore, ca 1890, Maingay 158 |Miiller

Arg., 1893: 129, type description; Marbach, 2000: 61]; Fort Canning Park, on

tree in park, 1989, Aptroot 25954 [B, Hb. Aptroot; Kalb and Elix, 1998: 468 as

Buellia; Marbach 2000: 63]; crossing Oxley Road/Oxley Rise, roadside trees,

1994, Diederich 12223 |Hb. Diederich].

A Conspectus of the Lichens (Lichenized Fungi) of Singapore 443

Anisomeridium biforme (Borrer) R.C.Harris — Kusu Island, 1989, Aptroot

25972 [Hb. Aptroot].

Anisomeridium foliicola R.Sant. & Tibell — (10b) SMT 46172 [SING].

Anisomeridium subprostans (Nyl.) R.C.Harris — (31) S 45663 [SING]; (3ae)

S 45778 |B, SING]; (3ba) S 45854 [B, SING]; Botanic Gardens, forest area,

1989, Aptroot 25962 [B, Hb. Aptroot].

Anisomeridium terminatum (Nyl.) R.C.Harris — (3ab) S 45745 [B, SING].

Anisomeridium throwerae R.C.Harris — (1}) SM 45509 [SING], in SM

45514 |B, SING] (= Pyxine cocoes); (2a) S 45516 [B, SING]; (2c) S 45546 [B,

SING]; (2d) S 45576 [B, SING]; (Se) ST 45942 [SING]; (6a) SF 46100 [B]: (7)

ST 46039 [SING]; (13) S 46245 [SING]; (14d) ST 46299 [B, SING]; (14f) ST

46353 [SING]; (18) ST 46411 [B, SING].

Here all Anisomeridium specimens are included with long-beaked,

seta-like pycnidia. The conidia are of two types, suggesting that more than

one species may be on hand: 9-12 x 4-5 um with rounded ends, 1-septate

(45516, 46411) and 8 x 4 um, cubic, simple (45942, 46299). The conidia are

lacking in many specimens.

Arthonia catenatula Ny|.— (1a) SM 45444 [B, SING]: (1b) SM 45446 [B,

SING], in SM 45452 [B, SING] (= Graphis caesiella); (1c) SM 45457 [SING];

(le) SM 45474 [B, SING]; (1f) SM 45483 [B, SING]; (1g) obs., in SM 45489

[SING] (= Graphis caesiella); (Ah) SM 45494 [SING]; (1i) SM 45498 [SING]:

(1j) SM 45511 [B, SING]; (2) S 45520 [SING]; (2b) in S 45526 [SING] (=

Cryptothecia cf. subnidulans); (2c) § 45542 [B, SING]; (2d) S 45559 [B, SING],

S 45575 [B, SING]; (2e) S 45589 [SING]; (2f) S 45604 [SING]; (2g) in S 45631

[B] (= Amandinea efflorescens), in S 45635 [SING] (parasite), in S 45637 [B,

SING] (= Lecanora helva); (3k) obs., in S 45668 [B, SING] (= Enterographa

pallidella); (3v) S 45723 [SING]; (3y) obs., in S 45732 [SING] (= Graphis cf.

caesiella); (3ab) obs., in S 45750 [B, SING] (= Arthonia cinnabarina); (3ac)

obs., in S 45773 [B] (= Porina tetracerae); (3ad) obs., in S 45775 [B, SING]

(= Phaeographis sp.); (3at) S 45839 [SING]; (3av) obs., in S 45842 [SING]

(= Trypethelium tropicum); (5a) ST 45863 [SING]; (7) obs.; (7a) ST 45999

[SING]; (7b) ST 46006 [B, SING]; (8) obs.; (13) obs., S 46246 [SING]; (14)

obs.; (14c) ST 46285 [SING], ST 46285a [B, SING]; (15) SM 46375 [SING];

(18) in ST 46409 [B, SING] (= Arthonia sp. E); crossing Oxley Road/Oxley

Rise, roadside trees, 1994, Diederich 12230 |Hb. Diederich].

The ascocarps are usually stellate with narrow radii, but conspicuously

rounded ascocarps occur occasionally (45575, 45723, 46246, 46285a).

Deviating ascospores (attenuated towards both ends, resembling A. sp.

A, but larger) were observed in 46409. TLC: tr. atranorin, confluentic acid

(45511, 45522, 45559, 45575). The ascospore size (usually about 30 x 12 um)

deviates from the protologue and the identification is provisional.

Arthonia cinnabarina (DC.) Wallr.— (1b) SM 45447 [SING]; (2a) S 45518

444 Gard. Bull. Singapore 61 (2) 2010

[B, SING]; (3ab) S 45750 [B, SING]; (18) ST 46412 [B, SING].

Arthonia complanata Fée — Vega expedition, corticola, 1879, Almquist

[Nylander, 1891: 22].

Arthonia subbessalis Nyl. — Vega expedition, corticola, 1879, Almquist

[Nylander, 1891: 23, type description].

Arthonia trilocularis Miill.Arg.— (6a) SF 46107 [SING]; (10b) SMT 46169

[SING], SMT 46170 [B, SING] cf.; (14f) ST 46358 [B, SING] cf.

The material is often in poor condition and the identification

provisional.

Arthonia sp. A — (2b) S 45528 [B, SING], S 45537 [B, SING]; (2e) S 45598

[B, SING]; (7a) ST 45999a [B, SING]; (18) ST 46410 [B, SING].

Superficially like A. catenatula, but apothecia pale brownish and

ascospores attenuated at both ends, 20-26 x 6-7 um, 5-7-septate.

Arthonia sp. B — (3d) S 45648 [B, SING]; (3ac) S 45771 [B, SING]? (no

spores); (3ah) S$ 45790 [B, SING]? (no spores); (Sa) ST 45860 [SING]? (no

spores); (14c) ST 46285b [B, SING]; (18) ST 46408 [B, SING].

Ascocarps brown to black, rounded or shallowly lobed, often slightly

brownish-pruinose; ascospores 12-15 x 4.5-6 um, 1-2-septate, one terminal

— cell swollen.

Arthonia sp. C — (2d) S 45576a [SING], S 45580 [B, SING]; (31) S 45673 [B,

SING]; (3w) S 45726 [B, SING]; (Sf) ST 45931 [SING]; (14) ST 46321 [B,

SING]; (14e) ST 46314 [B, SING]; (15) SM 46366 [B, SING].

Ascocarps black, rounded, small; ascospores 6-11 x 3 um, 1-septate,

with one swollen terminal cell.

Arthonia sp. D — (1c) SM 45458 [B, SING]?; (3a) S 45641 [SING]? (no

spores); (3v) S 45722 [SING]? (no spores); (3y) S 45730 [SING]; (3aa) S

45740 [SING]? (no spores); (3az) S 45850 [B, SING]; (3ba) S 45853 [B,

SING]; (Sa) ST 45858 [B, SING]; (18) ST 46407 [B, SING].

Ascocarps black, rounded, immersed when on soft, large-celled

bark; ascospores 10-16 x 3 um, 2-septate, with one swollen terminal cell;

epithecium dark-brown.

Arthonia sp. E — (18) ST 46409 [B, SING].

Similar to A. catenatula, but ascocarps with prominent white margin.

Arthotheliumsp.—(3i)S45662[SING];(3ab)$45748[B];(S5d)S 745878 [SING].

Ascocarps small, lobed, blackish; ascospores regularly muriform, 40 x

16 um, ca 10 x 4 locules.

Astrothelium ochrothelizum (Nyl.) Mill.Arg. — Vega expedition, corticola,

1879, Almquist |Nylander, 1891: 26 as Trypethelium|.

Astrothelium subfuscum Kremp.-— Ad cortices, Beccari 256 [Krempelhuber,

1875: 64, type description].

Bacidia rubellovirens (Nyl.) Zahlbr. — Vega expedition, corticola, 1879,

Almquist |Nylander, 1891:21 as Lecidea, type description].

A Conspectus of the Lichens (Lichenized Fungi) of Singapore 445

Bacidia sp. — (Se) ST 45922 |B, SING].

Apothecia pale brown; ascospores unripe, becoming acicular?

Bacidina aff. arnoldiana (K6rb.) V.Wirth & Vézda— (2g) § 45627 [B, SING].

Pycnidia present only; conidia ca 60 x 0.5 um, curved.

Bacidina sp.? — (18) ST 46397 [SING].

A poor specimen probably belonging to this genus, but not conspecific

with the preceding species.

Bactrospora metabola (Nyl.) Egea & Torrente — (5c) ST 45867 [B, SING].

Bactrospora myriadea (Fée) Egea & Torrente — (1b) obs.,in SM 45447

[SING] (= Arthonia cinnabarina); (1c) in SM 45456 [SING] (= Chrysothrix

xanthina); (1f) SM 45485 [B, SING]; (11) SM 45496 [B, SING]; (2b) S$ 45536

[SING]; (2f) S 45607 [B, SING]; (3) S 45657 [SING]; (7) ST 46046 [B].

Badimia sp.? — (17a) ST 46388b [SING].

The material is very scarce and was not examined microscopically.

Biatora (s.1.) sp. — (2) S 45585 [B, SING]; (3) obs.

Material with biatorine apothecia and simple, hyaline ascospores, of

unclear affinity.

Biatorella (s.1.) sp. — (2) S 45581 [B, SING].

Material with biatorine apothecia, polysporous asci and simple,

hyaline ascospores, of unclear affinity.

Buellia sp. A — (17) ST 46389 [SING].

Single thallus, on brick.

Buellia sp. B -- (7) ST 46013 [SING].

Single thallus, on granitic boulder.

Byssoloma leucoblepharum (Ny1.) Vain.—(10b) SMT 46191 [B, SING]; 1882,

on Cantley 48 (Memecylon cantleyi) [K, not seen; Santesson, 1952: 487].

Byssoloma tricholomum (Mont.) Zahlbr. — Ad folia coriacea, Beccari 269a

[Krempelhuber, 1875: 60 as Lecanora epiphylla; Santesson, 1952: 483].

Calenia aspidota (Vain.) Vézda — (14f) ST 46355 [B, SING].

Calicium hyperelloides Ny\.— (3s) S 45715 [SING]; (14c) ST 46288 [B, SING];

(15) SM 46368 |B, SING]; Sentosa, along the southern shore, outskirts of

forest along the beach, on trunk, 1980, Tibell 8865 [UPS L-057807].

Calopadia subcoerulescens ( Vain.) Vézda — (10b) SMT 46186 [SING].

Calopadia cf. vermiculifera (Vain.) Sérus.? — (10b) SMT 46173 [B, SING].

Poor specimen, identification uncertain.

Calopadia sp. — (14f) ST 46354 [B, SING].

Poor specimen with muriform ascospores 1|-2/ascus.

Caloplaca sp. A — (14) ST 46266 [SING].

Single, saxicolous thallus.

Caloplaca sp. B — Kusu Island, 1989, Aptroot 25969 [Hb. Aptroot].

The apothecia have a grey thalloid margin and orange disc.

Carbacanthographis candidata (Nyl.) Staiger & Kalb — Vega expedition,

446 Gard. Bull. Singapore 61 (2) 2010

corticola, 1879, Almquist [Nylander, 1891: 23 and Redinger, 1936: 50 as

Graphis singaporina, type description; Staiger and Kalb, 1999: 124].

Carbacanthographis marcescens (Fée) Staiger & Kalb — (6) SM 45963 [B,

SING].

Catarraphia dictyoplaca (Mont. & v.d. Bosch) A.Massal.— (10) SMT 46124

[B, SING].

Catillaria (s.1.) sp. — (11) SMT 46202 [B, SING].

Specimen with biatorine apothecia and hyaline, uniseptate ascospores,

of uncertain affinity.

Chapsa indica A.Massal. — (13) S 46258 [B, SING].

TLC: none.

Chapsa platycarpella (Vain.) A.Frisch — (Se) ST 45899 [B, SING]; (10) SMT

46133 [B, SING]; (11) SMT 46208 [SING].

Chiodecton leptospermum Miill.Arg. — (3ab) S 45770 [B, SING]; (3an) S

45810 [SING].

Chiodecton natalense Nyl.— (3v) S 45721 [B, SING]; (3af) S 45781 [SING];

(Sa) ST 45862 [SING]; (5c) ST 45869 [SING].

Chroodiscus australiensis Vézda et Lumbsch — (10b) SMT 46193 [B,

SING].

Chroodiscus mirificus (Kremp.) R.Sant.— (10b) SMT 46199 [B, SING]; Nee

Soon Forest Reserve, tropical forest remnant, on leaves of Aglaea trinervis,

1980, Tibell 8813 [UPS L-057770]; ibidem, on leaves of Calamus scipionum,

1980, Tibell 8815 [UPS L-057772].

Chroodiscus cf. mirificus (Kremp.) R.Sant.— (6a) SF 46103 [B, SING]; (10b)

SMT 46199a [B, SING].

The material deviates from SMT 46199 because the rounded

schizidia develop on the thallus, not at the margin, and are thickened in

the centre. In the absence of ascocarps the classification is provisional.

Chrysothrix xanthina (Vain.) Kalb — (1c) SM 45456 [SING]; (1g) in SM

45490 [SING] (= Herpothallon granulare); (2b) S 45525 [SING]; (2d) S$

45574 |B, SING]; (2e) in S 45598 [B, SING] (= Arthonia sp. A); (2f) S 45603

[B, SING]; (2g) in S 45630 [SING] (= Lecanora helva), in S 45635 [SING] (=

parasite), in § 45624 [SING] (= Pyrrhospora quernea); (7) obs.; (8) obs.; (13)

obs.; (14) obs.; (15) obs.; (18) obs.; on mango tree along road, Pulau Tekong,

2000, B. C. Tan [Hb. Tan]; crossing Oxley Road/Oxley Rise, roadside trees,

1994, Diederich 12222 [Hb. Diederich].

Cladonia subradiata (Vain.) Sandst. — (3w) S 45725 [B, SING].

TLC: fumarprotocetraric acid.

Coccocarpia erythroxyli (Sprengel) Swinsc. & Krog — sine loc. [BM, L,

not seen; Arvidsson 1982: 62]; Vega expedition, corticola, 1879, Almquist

[Nylander, 1891: 18 as Coccocarpia ciliolata].

Coccocarpia palmicola (Spreng.) L.Arvidss. & D.J.Gallow. — (14) ST 4635/

A Conspectus of the Lichens (Lichenized Fungi) of Singapore 447

[B, SING]; sine loc. [US, not seen; Arvidsson, 1982: 76]; St. John’s Island,

1989, Aptroot 26000 [Hb. Aptroot]; Botanical Garden, on Cupressus sp.,

1980, Tibell 8889b [UPS L-057820, not seen].

Coccocarpia pellita (Ach.) Miill.Arg. — (3a) S 45645 [B, SING]; (14c) ST

46291 [B, SING]; sine loc. [W, not seen; Arvidsson, 1982: 79].

Coccocarpia rottleri (Ach.) L. Arvidss. — (14e) ST 46301 [B, SING].

Coenogonium confervoides Ny|.— Vega expedition, corticola, 1879, Almquist

[Nylander, 1891: 20].

Coenogonium dilucidum (Kremp.) Kalb & Liicking — (6a) SF 46/02 [B,

SING]; (10b) SMT 46184 [B, SING]; (14f) ST 46359 [B, SING].

Coenogonium epiphyllum Vain. — (2d) S 45572 |B, SING] cf.; (6a) SF 46106

[B, SING], 46/08 [B, SING].

Coenogonium luteum (Dicks.) Kalb & Liicking — Vega expedition, corticola,

1879, Almquist [Nylander, 1891: 21 as Gyalecta lutea).

Coenogonium subluteum (Rehm) Kalb & Liicking — (10b) SMT 461/82 [B,

SING].

Collema actinoptychum Nyl. — 1897, Moller [S, TUR, not seen; Degelius,

1974: 125].

Collema leptaleum Tuck. var. biliosum (Mont.) Degel. — University area,

roadside trees, often abundant, 1964, Degelius As-533, 535, 559 [UPS-Hb.

Degelius, not seen; Degelius, 1974: 108].

Collema rugosum Kremp. — Singapore Botanic Gardens, 189?, Ridley A.

54 [SING]; Economic Gardens, Bombax, 1919, unknown collector, 5416,

pp. |K, UPS not seen; Degelius, 1974: 153]; Singapore Botanical Garden,

on Cupressus sp., 1980, Tibell 8887 [UPS L-057817, not seen]; university

area, roadside trees, locally abundant, 1964, Degelius As-489, 495 [UPS-Hb.

Degelius; Degelius, 1974: 153].

Cratiria lauricassiae (Fée) Marbach — Singapore Botanic Gardens, 1959,

Burkill 2178 [SING].

Cresponea flava (Vain.) Egea & Torrente — (2d) S 45549 [B, SING]; (2g) S

45632 |B, SING], in S 45630 [SING] (= Lecanora helva), in S 45635 [SING]

(= parasite); (3r) S 45713 [SING]; (3z) S 45736 [B, SING]; (3aa) S$ 45741

[B]; (Sb) obs., in ST 45864 [B, SING] (= Graphis insulana); (7a) obs., in ST

45999 [SING] (= Arthonia catenatula); (8) ST 46055 [B, SING]; (13) S$ 46243

[SING]; (13a) S 46224 [SING]; Fort Canning Park, on tree in park, 1989,

Aptroot [Hb. Aptroot]; Kusu Island off the coast, on tree along the coast,

1989, Aptroot [Herb. Aptroot]; St. John’s Island, 1989, Aptroot 25995 |Herb.

Aptroot, det. Egea 1992]; Singapore Botanical Garden, on Cedrela toana,

1983, Tibell [UPS; Egea and Torrente, 1993: 316]; crossing Oxley Road/Oxley

Rise, roadside trees, 1994, Diederich 12229 [Hb. Diederich].

Cresponea proximata (Nyl.) Egea & Torrente — (14) ST 46325 [B, SING],

46348 [SING].

448 Gard. Bull. Singapore 61 (2) 2010

Crocynia pyxinoides Nyl. — (3m) S 45682 [B, SING], in S 45678 [SING] (=

Porina tetracerae).

TLC: atranorin, stictic acid, pannarin?, indet. substances (45682).

Cryptothecia aleurella (Nyl.) Makhija & Patwardhan — Vega expedition,

corticola, 1879, Almquist [Nylander, 1891: 22 as Arthonia, type description].

Cryptothecia candida (Kremp.) R.Sant. — (10) SMT 46143 [B, SING],

46145 [B, SING]; (10b) SMT 46197 [B, SING]; Nee Soon Forest Reserve,

tropical forest remnant, on leaves of Aglaea trinervis, 1980, Tibell 8814 [UPS

L-057771, not seen].

TLC: 2’-0-methylanziaic acid or similar spot (46143, 46145, 46197).

Cryptothecia irregularis Liicking et al.— (10b) SMT 46196 [B, SING].

Cryptothecia lunulata (Zahlbr.) Makhija & Patwardhan -- (1j) SM 45507

[B, SING]; (2a) S 45515 [SING]; (2c) S 45540 [B, SING]; (3q) in S 45709

[B] (= Enterographa pallidella); (Se) ST 45886 [B, SING], 45977 [B, SING],

45927 |B, SING]; (Sf) ST 45934 [B, SING]; Benjamin Lee n.c. [SING].

TLC: confluentic and/or barbatic? acid (45540, 45886, 45917, 45934).

Cryptothecia obtecta Makhija & Patwardh. — (5f) ST 45935 [B, SING]; (6)

SF 46090 [SING]; (10) SMT 46128 [B, SING], 46/30 [B, SING], 46/44 [B,

SING]; (14) ST 46331 [SING].

TLC:confluentic, gyrophoric? acids (45935, 46128, 46130, 46144, 46425).

Cryptothecia scripta Thor — (1d) SM 45468 [SING]; (3a) S 45643 [B, SING];

(31) S 45664 [SING]; (3m) S 45683 [B, SING]; (3ab) S 45763 [B, SING]; (3ar)

S 45836 [B, SING]; (Sa) ST 45859 [SING]; (Sc) ST 45868 [B, SING]; (13) in

S 46240 [B] (= Diorygma rufopruinosum); (14) ST 46318 [SING]; (14e) ST

46310 [B, SING]; (17) ST 46391 [SING]; Singapore Botanic Gardens, 1959,

in Burkill 2177 [SING] (= Dirinaria picta) vs., 1+ blue, C+ red; juv. asci.

TLC: gyrophoric/hiascic acid complex (45643, 45683, 45763, 45836,

45868, 46310).

Cryptothecia cf. subnidulans Stitt. — (2b) S 45526 [SING]; (6) SM 45971

[B, SING], SF 46097 [B, SING]; (14) ST 46323 [SING]; Benjamin Lee [B,

SING].

TLC: gyrophoric acid “ape, ~G597, 4o09T)) Wiceusins):

Cryptothecia sp. A — (13) S 46238 [B, SING]; (14) ST 46319 [B, SING]; (18)

ST 46394 [SING].

TLC: barbatic acid (46319). Thick, sorediate thalli of unclear affinity.

Cryptothecia sp. B — (1f) SM 45484 [B, SING]; (3a) in S 45643 [B, SING] (=

Cryptothecia scripta); (30) S 45692 [SING]; (5d) ST 45877 [SING]; (Sf) ST

45936 [B, SING]; (7) ST 46009 [SING]; (13) S 46230 [SING]; (14a) ST 46273

[B, SING]; (14d) ST 46300 [B, SING]; (15) obs.

The material is chemically variable and probably includes more than

one species. It is sorediate and lacks ascocarps and is therefore of unclear

affinity.

A Conspectus of the Lichens (Lichenized Fungi) of Singapore 449

Cryptothelium sp. — (10a) SMT 46159 [B, SING], 46164 [B, SING].

Ascospores 4/ascus, muriform, 60-80 x 20 um.

Dichosporidium boschianum (Mont.) Thor — (6) SM 45949 [SING], 45962

[B, SING]: Bukit Timah, 1959, Burkill 2139 [SING]; Botanical Garden,

Jungle, on Castillea elastica, Tibell 8872, 1980 [UPS]; Bukit Timah, 1950,

Lemaitre [H, not seen; Thor, 1990: 67].

Diorygma hieroglyphicum (Pers.) Staiger & Kalb — (1f) SM 45478 [SING];

(3ab) S 45757 [B].

TLC: stictic, cryptostictic acids (45478).

Diorygma pruinosum (Eschw.) Kalb, Staiger & Elix — (3ag) S 45786 [SING]:

(13) S 46236 [B]: (14a) ST 46270 [B]: (14d) ST 46298 [SING].

TLC: protocetraric acid (45789, 46298).

Diorygma reniforme (Fée) Kalb, Staiger & Elix — (3m) S 45685 [B, SING];

(13) S 46257 [B, SING].

TLC: salazinic acid (45685, 46257). Ascocarps absent.

Diorygma rufopruinosum (A.W.Archer) Kalb, Staiger & Elix — (2d) S$

45554 [B, SING], 45577 [B, SING]; (2f) S 45612 [SING]; (3m) S 45681 [B];

(3p) S 45705 [B, SING]; (3ai) S 45793 [SING], 45794 [SING]; (3au) S 45840

[B, SING]; (7) ST 46007 [SING], 46026 [B, SING]; (13) S 46240 [B]: (15) SM

46372 |B, SING], 46373 [B, SING] (c. apoth.).

TLC: norstictic, connorstictic, protocetraric acids (45554, 45577,

45840, 46240, 46372, 46373). The Singapore material of this species deviates

by the thallus, which is densely covered by very irregular warts turning into

pustules, which burst and become sorediate. It is a common “sterile crust” in

Singapore and ascocarps are rare.

Dirinaria applanata (Fée) Awasthi — University area, on trees, 19464,

Degelius As-491 [Degelius in UPS]: Awasthi, 1975:82; 1879, Almquist [S, not

seen], Harmand [M, not seen; Awasthi, 1975:82]; Vega expedition, corticola,

1879, Almquist [Nylander, 1891: 18 as Physcia picta f. sorediata]; university

area, roadside trees, 1964, Degelius As-491 [UPS L-099232, not seen].

The reports may be erroneous, see note under D. picta.

Dirinaria caesiopicta (Ny|.) Awasthi — On bark, 1879, Almquist [H-Nyl

31822, not seen; Awasthi, 1975: 96].

Dirinaria confluens (Fr.) Awasthi — Botanical Garden, ad corticem palmae,

1949, Lemaitre [SING 031008; det. Awasthi 1966].

Dirinaria consimilis (Stirton) Awasthi — On stones, 1949, Lemaitre [H, not

seen; Awasthi, 1975: 93].

Dirinaria naggarana (Kremp.) Awasthi— Botanical Garden, 1949, Lemaitre

[H, SING, not seen; Awasthi, 1975: 62].

Dirinaria picta (Sw.) Clem. & Shear — (1c) SM 45454 [B, SING], 45455 [B,

SING]; (1d) SM 45464 [B, SING]: (le) in SM 45476 [SING] (= Pyrenula

ochraceoflava); (11) in SM 45499 [B] (= Pyrenula ochraceoflava); (1j) SM

450 Gard. Bull. Singapore 61 (2) 2010

45503 |B, SING], 45513 [SING]; (2a) in S$ 45519 [B] (= Graphis caesiella);

(2b) S 45535 [B, SING]; (2d) S 45557 [B, SING], 45555 [B, SING], 45579

[SING]; (2f) S 45616 [B], 45617 [B, SING]; (2g) S 45628 [B, SING], 45629

[SING], 45633 [B, SING], 45634 [SING], in 45624 [SING] (= Pyrrhospora

quernea); (3) obs.; (3a) in § 45643 [B, SING] (= Cryptothecia scripta); (3m)

in S 45679 [B] (= Graphis insulana); (3aa) S 45737 [SING]; (3aj) S 45802

[B], 45802a [B]; (Sd) ST 45871 [SING]; (7) ST 46021 [SING]; (7a) ST 45993

[SING], 45994 [B]; (8) ST 46064 [B, SING], 46065 [SING], 46066 [B, SING],

46067 [B, SING], 46069 [SING], obs.; (12) SMT 46219 [B, SING] c. ap.; (13)

S 46260 [SING], 4626/ [B, SING]; (14) ST 46322 [SING]; (14c) ST 46287 [B,

SING]; (15) obs. (2x); (18) obs.; Botanic Gardens, 1959, Burkill 2177 [SING];

Pulau Terkukor, 1960, in Lemaitre [SING] (= Pyxine farinosa); Singapore

Botanical Garden, on palm trunk, 1949, Lemaitre [SING 031010; Awasthi,

1975: 75]; Singapore Botanical Garden, 1991, Gams [Hb. Aptroot]; on trees,

1861-1865, Maingay [Nylander and Crombie, 1884: 52 as Physcia]; Pulau

Tekong, on rubber tree, 2000, B.C. Tan [Hb. Tan]; Pulau Tekong, on Acacia

tree at seashore, 2000, B.C. Tan |Hb. Tan]; Pulau Tekong, on mango tree along

road, 2000, B. C. Tan [Hb. Tan]; crossing Oxley Road/Oxley Rise, roadside

— trees, 1994, Diederich 12228 [Hb. Diederich]; Sentosa, close to the ferry, on

trunk of tree, 1980, Tibell 8853 [UPS L-057795, not seen]; Fort Canning Park,

1989, Aptroot 25952 |Hb. Aptroot]; Botanical Garden, ad ramas arborum,

1949, Lemaitre [SING 500374; Awasthi, 1975: 82 as D. applanata|.

This sorediate species tends to form clones. Thus, it is not uncommon

to find two separate populations differing in lobe size on a single trunk.

These may give the impression of separate species, in particular D. picta

together with D. applanata (Fée) D.D.Awasthi. However, populations

from adjacent trunks are often intermediate and the large material

available did not allow a clear subdivision in two species. TLC: atranorin,

sekikaic acid, terpenoids (45455, 45503, 45551, 45616, 45628, 45633,

45802, 46261, 46287, 46219); atranorin, divaricatic acid, terpenoids

(45464, 45535, 4555, 45617, 45802a, 45994, 46064, 46066, 46067). No

morphological differences seem to exist between the two chemotypes.

eeaee afzelii (Ach.) A.Massal. — (30) S 45697 [SING]; (3ai) S 45795

[B]; (Se) ST 45900 [B, SING]; (10a) SMT 46161 [SING]; Mandai Road,

1920, pee 5801 [SING]; Botanic Gardens, 1920, in Kiah 5846 [SING]

(= Phaeographina?); Botanic Gardens, 1920, in Noor 5663 [SING] (=

Trypethelium sp.); On bark of trees, 1861-1865, Maingay [Nylander and

Crombie, 1884: 57 as Graphis|; Vega expedition, corticola, 1879, Almquist

[Nylander, 1891: 23]; Beccari 240 [M, not seen; Redinger, 1936: 54;

Krempelhuber, 1875: 61]; Beccari 244 [M, not seen; Redinger, 1936: 55 and

Krempelhuber, 1875: 61 as Graphis atro-alba].

According to Redinger (1936) Graphis atro-alba is a damaged stage

A Comspectus of the Lichems (Lichemized Fumgi)) of Singapore 451

of D. afzelii.

Echinoplaca pellicula (Mill_-Arg.) R-Sant.— (4) S 45835 [B. SING}: (6a) SF

46099 |B. SING]. 46112 [B. SING]: (10b) SMT 46177 [B. SING]. 46178 [B.

SING]: (17a) ST 46388a [B. SING].

Echinoplaca sp. — (5e) ST 45943 [B. SING}]?: (10b) SMT 46166 [SING]: Bot-.

Gardens, 1919. Chipp 4915 [SING].

The material lacks apothecia but contains hygrophores. More then

one species may be involved.

Endocarpon pallidum Ach. — (2) S 45583 |B. SING]. 45584 [SING]: (7)

obs.

Enterographa anguinella (Ny\.) Redinger — (le) SM 45475 [B. SING:

Sparnius. 2004: 27].

Enterographa angustissima ( Vain.) R-Sant. — (Se) ST 45938 [B. SING]:

(10b) SMT 46176 [SING].

Enterographa divergens (Miull_Arg.) Redinger — (7a) ST 45989 [B. SING:

Sparnius. 2004: 37].

Enterographa pallidella (Ny\.) Redinger — (1a) SM 45445 [SING]: (1b) SM

45451 |B]: (ih) SM 45495 [B. SING]: (3k) S 45668 [B. SING]: (3q) S 45709

[B: Sparrius, 2004: 50]: (14b) ST 46278 [B. SING]: (15) SM 46378 [SING].

Enterographa subserialis (Ny1.) Redinger — (3a) S 45638 [B: Sparrius, 2004:

61].

Enterographa tropica Sparrius — (30) S 4569] [B: Sparrius. 2004: 62]: (4)

S 45829 |B: Sparrius. 2004: 62]: (6) ST 46383 [B holotype. SING isotype:

Sparnus. 2004: 61. 62]; (11) SMT 46203 [SING].

Enterographa sp. — (15) SM 46374 {[B. SING).

This material has lecanoroid apothecia and reminds E. anguinella.

im particular the morph named E. lecanoroides R.C.Harris. However, it

lacks psoromic acid and contains an unidentified depsid staying low in the

standard solvent systems.

Eremothecella palmulacea (Mill-Arg.) Sérusiaux — (10b) SMT 46168 [B.

SING].

Eschatogonia? sp. A — (4) S 45817 [B. SING}: (14e) ST 46315 [B. SING} c.

ap.

The genus identification is provisional and based on the squamules

with a glossy, corticate lower side.

Eschatogonia? sp. B — (5<e) ST 45898 [B. SING}.

The genus identification is provisional and based on the squamules

with a glossy, corticate lower side.

Eugeniella micrommata (Kremp.) Licking. Sérus. & Kalb — (6a) SF 4610]

[B. SING]: (10b) SMT 46189 [B. SING]: Nee Soon Forest Reserve. tropical

forest remnant. on leaves of Calamus scipionum, 1980. Tibell 8816 [UPS

L-057773].-

452 Gard. Bull. Singapore 61 (2) 2010

Fellhanera bouteillei (Desm.) Vézda — (6) SM 45984 [SING]; (14f) ST 46356

[B, SING].

Fellhanera sp. A — (2d) S 45578 [SING]; (3m) S 45676 [B, SING]; (3al) S$

45807 |B, SING]; (3am) S 45808 [B, SING].

This species has conspicuous, dark-brown, beaked pycnidia and brown

apothecia with three-septate ascosporesca 10-12 x2.5-4um.Conidia3 x 1.5 um.

Fellhanera sp. B — (1a) SM 45441c [SING].

Foliicolous, grey, granular thallus with pale yellow-brown, small

apothecia; ascospores uniseptate, 14 x 5 um.

Fellhanera sp. C — (2d) S 45571 [B, SING]; (3g) S 45658 [B, SING] (c. pycn.);

(6) SM 45957 [B, SING]; (7) ST 46012 [B, SING], 46030 [B, SING]; (4) S

45818 [B, SING].

Apothecia pale brown with an often prominent, whitish margin;

ascospores 3-septate, 10-12 x 3 um. Pycnidia like small, young apothecia;

conidia 3-4 x 1.5 um.

Fellhanera sp. D — (10b) SMT 46171 [B, SING].

Apothecia tiny, dark-brown, dense; ascospores 3-septate, 10 x 3 um.

Fellhanera sp. E — (3) S 45855 |B, SING]; (7) ST 46015 [B, SING], 46016 [B,

SING]; (7a) ST 45986 [B, SING]; (13) S 46256 [SING]?; (14) ST 46336 [B,

SING], 46337 [B, SING]; (14c) ST 46292 [B, SING]; (14e) ST 46312 [SING];

(16) ST 46387 [SING]; 1989, Aptroot [Hb. Aptroot].

Apothecia pale brown to brown;ascospores 3-septate, 13-20(-24) x4um.

Fellhanera sp. F — (14) ST 46317 [B, SING].

Muscicolous; apothecia black, soon convex; ascospores 3-septate, 16

x5) tinal,

Fissurina cf. dumastii Fée — (Se) ST 45907 [B, SING]; (6) SF 46077 [B,

SING] (no spores), 46098 [B, SING] (no spores); (7) ST 46041 [B, SING]

(thin septa); (11) SMT 46205 [B, SING] (no spores), 46206 [B, SING] (thin

septa); (14) ST 46339 [B] (thin septa).

TLC: none (45907, 46041, 46077, 46098, 46205, 46206). The

identifications in this genus are provisional.

Fissurina cf. incrustans Fée — (3m) S 45677 [B, SING]; (30) S 45696 [SING];

(4) § 45820 [B, SING], 45828 [B, SING]; (6) SF 46093 [B, SING], 46095 [B,

SING], 46096 [SING]; (14e) ST 46305 [B, SING], 46309 [B, SING].

TLC: stictic acid (45677, 45820, 45828, 46096, 46305, 46309).

Fissurina cf. radiata Mont. — (6) SM 45973 [B, SING]; (6) SF 46089 [B,

SING] (no spores); (10) SMT 46122 [B, SING].

TLC: none (45973, 46122).

Fissurina sp. A — (1d) SM 45472 |B, SING].

TLC: none; ascospores muriform, ca 4/ascus, 18 x 7 um.

Glyphis cicatricosa Ach. — (2b) in § 45525 [SING] (= Chrysothrix xanthina);

(3ag) S 45785 [SING]; on Cocos palms, 1861-1865, Maingay [Nylander and

A Conspectus of the Lichens (Lichenized Fungi) of Singapore 453

Crombie, 1884: 59]; corticola, Maingay [Nylander, 1891: 25]; Beccari 239 [M,

not seen; Redinger, 1936: 98; Krempelhuber, 1875: 62].

Glyphis scyphulifera (Ach.) Staiger — (13a) S 46220 [B, SING].

Graphis assimilis Nyl. — Vega expedition, corticola, 1879, Almquist

[Nylander, 1891: 23].

Graphis caesiella Vain. — (1) in SM 45473 [SING] (= Phaeographis sp. C);

(1b) SM 45452 [B, SING]; (1d) SM 45466 [B, SING]; (1f) obs., in SM 45485

[B, SING] (= Bactrospora myriadea); (1g) SM 45489 [SING]; (2a) S 45519

[B]; (2b) S 45527 [SING]; (2c) in S 45539 [B, SING] (= ster. crust.); (2e) obs.,

in S 45587 [B, SING] (= Phaeographis intricans); (2f) S 45609 [B, SING];

(2g) S 45625 [B, SING], in S 45635 [SING] (= parasite); (30) S 45689 [SING];

(3y) S 45732 [SING] cf. (large ascospores); (3ab) obs., 45758 [B]; (7) obs., ST

46027 [B, SING]; (7a) ST 45995 [B, SING]; (8) ST 46074 [B, SING]; (13) obs.,

S 46233 [SING]; (13a) obs., in S 46227 [SING] (= Pyxine cocoes); (14) in ST

46352 [B] (= Graphis insulana); (14a) ST 46275 [B, SING] (small); (14c) ST

46286 |B, SING]; (15) obs.; (18) ST 46395 [B, SING].

TLC: norstictic, tr. connorstictic acids (45625, 46286, 46395); (tr.)

norstictic, salazinic acids (45466, 45519, 45609 (with galbinic acid), 46074);

salazinic acid (46027, 46275); none (45995). Salazinic acid-containing

specimens would fit Graphis bakeri Vain. (Licking et al., 2008). However,

the salazinic acid is usually accompanied by traces of norstictic acid not

mentioned by Liicking et al. (2008), the chemical variation is not correlated

with any morphological differences and an intermediate specimen containing

about equal amounts of norstictic and salazinic acids was observed. Therefore

all material is considered to be a single species. Only a specimen containing

stictic acid but otherwise rather similar in morphology is treated here as a

separate species, G. dendrogramma. Its ascocarps seem less pruinose and

more radiately branched.

Graphis cf. cleistoblephara Ny|.— (7) ST 46023 [B, SING].

TLC: norstictic acid. The specimen deviates by the smaller ascospores,

ca 35 x 12 um, ca 4 /ascus.

Graphis confluens (Mont.) Nyl.— Vega expedition, corticola, 1879, Almquist

[Nylander, 1891: 24.

This record may need reinvestigation to establish its current taxonomic

position.

Graphis dendrogramma Ny|.— (2d) S 45564 |B, SING].

TLC: stictic acid with traces of cryptostictic and ?constictic acid. See

comment under G. caesiella.

Graphis glaucescens Fée — (2d) S 45562 |B, SING], 45563 [B, SING]; (3m) S$

45686 [B, SING]: (3ab) S 45761 [SING]; (5d) ST 45872 [B] (large ascospores);

(Se) ST 45908 [SING]; (6) SM 45975 [B, SING]; SF 46087 [B, SING].

TLC: indet. terpenoid? (45562, 45563, 45686, 45761, 45908, 45975,

454 Gard. Bull. Singapore 61 (2) 2010

46087).

Graphis insulana (Miill.Arg.) Liicking & Sipman — (1b) SM 45448 [B,

SING]; (1c) SM 45459 [SING]; (1i) SM 45497 [B, SING]; (2d) S 45560 [B,

SING], 45565 [B, SING]; (2g) in $ 4563/1 [B] (= Amandinea efflorescens),;

(3a) S 45642 [SING]; (31) S 45665 [B, SING]; (3m) S 45679 [B]; (3y) in S$

45732 [SING] (= Graphis caesiella) (degenerated); (3ae) S 45777 [SING];

(Sb) ST 45864 [B, SING]; (7) ST 46042 [SING]; (8) ST 46071 [SING]; (13) S

46237 [SING]; (14) ST 46352 [B]; (14e) ST 46307 [B, SING]; (15) obs.; (18)

obs.; Jungis Banactas, 1920, Ridout 5657 [SING]; Sentosa, along the southern

shore, on trunk, 1980, Tibell 8858 [UPS L-057800]; nahe der Philippinischen

Botschaft, 1999, Schumm & Schwarz [Hb. Schumm 5849].

TLC: norstictic acid with or without traces of connorstictic and

galbinic? acid (45448, 45497, 456065, 45777, 45864, 46352); indet. high spot

(45560, 45565); none (45679, 46042, 46071, 46307). The species is easily

recognizable by the thick thalline margin of the lirellae and the finely

inspersed hymenium with large, muriform, single ascospores. Its chemistry

is somewhat variable because norstictic acid is sometimes accompanied

by other substances, and occasionally it is not observed on the TLC plates.

The excipulum is usually only laterally carbonized, but may be thinly

carbonized below. It has been confused with G. hiascens (Fée) A.W.Archer.

Graphis librata C.Knight — (1d) SM 45462 [SING]; (11) SMT 4621/5 [B,

SING] (thin lirellae).

TLC: norstictic acid (45462, 46215).

Graphis rustica Krempelh.— Ad cortices, Beccari 258 [M, not seen; Redinger,

1936: 49; Krempelhuber, 1875: 61, type description].

Graphis scripta Ach. — On bark of trees, atypical, 1861-1865, Maingay

[Nylander and Crombie, 1884: 57]; Vega expedition, corticola, 1879, Almquist

[Nylander, 1891: 23]; species dubia vel excludenda [Redinger, 1936: 118].

These records may need reinvestigation to establish their current

taxonomic position.

Graphis scripta Ach. var. serpentina Ach. — On cocos-nut and palms, 1861-

1865, Maingay [Nylander and Crombie, 1884: 57].

This record may need reinvestigation to establish its current taxonomic

position.

Graphis_ striatula Ach. — Vega expedition, corticola, 1879, Almquist

[Nylander, 1891: 23].

Graphis tenella Ach. — (10a) SMT 46156 [B, SING]; (18) ST 46392 [B,

SING].

TLC: none (46/56).

Graphis vestitoides (Fink) Staiger — (2d) S 4556/7 [B, SING].

TE@znone:

Gyalideopsis vainioi Kalb & Vézda — (13) S 46251 [B, SING].

A Conspectus of the Lichens (Lichenized Fungi) of Singapore 45

Gyalideopsis sp. — (3ab) S 45752 [B]; (4) S 45834 [SING]; (14e) ST 46302

[B, SING].

The material is insufficiently developed for certain identification.

More than one species may be involved.

Haematomma rufidulum (Fée) A.Massal. — on trees, 1861-1865, Maingay

67 [BM, not seen; Staiger and Kalb, 1995: 162]; on trees, 1861-1865, Maingay

[Nylander and Crombie, 1884: 53 as Lecanora punicea]; Vega expedition,

corticola, 1879, Almquist [Nylander, 1891: 19 as Lecanora punicea].

It is unclear if the Maingay specimen investigated by Staiger is the

same as that investigated by Nylander and thus belongs to H. rufidulum.

For the time being all literature reports are considered to belong to the

same species. However, many Haematomma species look very similar and

occur in similar habitats, so that the presence of more than one species in

Singapore is not unlikely.

Herpothallon granulare (Sipman) Aptroot & Liicking — (1g) SM 45490

[SING]; (2d) S 45566 [SING]; (3f) S 45654 [SING]; (31) S 45671 [B, SING];

(3as) S 45838 [B holotype, SING isotype); (3az) S 45851 [B, SING]; (7) ST

46031 [B, SING], 46036 [SING].

See also Sipman (2003).

Lepraria? sp. A (thin yellowish) — (2f) S 45600 [B, SING]; (7a) ST 45988 [B,

SING]; (13) obs.; (14c) ST 46280 [B, SING]; (18) obs.; crossing Oxley Road/

Oxley Rise, roadside trees, 1994, Diederich 12226 |Hb. Diederich].

TLC: usnic acid, zeorin (45600, 45988, 46280). Thallus finely sorediate,

therefore generic affinity uncertain.

Lepraria sp. B — (3w) S 45727 [B, SING].

TLC: indet. substances.

Leptogium cyanescens (Rabenh.) K6rb.— (17) ST 46388 [B, SING]; Botanical

Garden, on Cupressus sp., 1980, Tibell 8888 [UPS L-057818].

The species is understood here in a wide sense, to accommodate thin-

lobed Leptogium species with phyllidia.

Leptogium cochleatum (Dicks.) P.M.Jorg. & PJames — New Tiew, Ama

Kong, 1949, Lemaitre [SING].

Leptogium marginellum (Sw.) S.F.Gray — Botanical Garden, root of

Swietenia macrophylla, 1964, Degelius As-486 [UPS L-102634].

Leptogium phyllocarpum (Pers.) Nyl.— Botanical Garden, root of Swietenia

_ macrophylla, 1964, Degelius As-485 [UPS L-102652].

Leucodecton occultum (Eschw.) A.Frisch — (1}) SM 45504 [B, SING]; (3ab)

S 45754 [B, SING].

TLC: norstictic, stictic, tr. cryptostictic acids (45504, 45754).

Lithothelium illotum (Ny1|.) Aptroot — (1d) SM 45471 [B, SING].

Lopadium sophodinum (Nyl.) Zahlbr. — Vega expedition, corticola, 1879,

Almquist [Nylander, 1891: 21 as Lecidea, type description].

This record needs reinvestigation to establish its current taxonomic

position.

Malcolmiella cf. olivaceolurida (Vain.) INED. — (4) S 45815 [SING]; (10)

SMT 46126 |B, SING], 46/47 [B, SING].

Ascospores 7-10 x 4-5 um. The description of Lecidea olivaceorufa

Vain. fits, but an investigation of type material is desirable for certainty.

Therefore no formal new combination is made.

Malcolmiella sp. A — (10) SMT 46126a [B, SING].

Malcolmiella sp. B — (6) SF 46097 [B, SING].

Material sorediate, without ascocarps, therefore identification

provisional.

Malcolmiella sp. C — (6) SF 46086 |B, SING]; (14) ST 46329 [B, SING]; on

treefork in disturbed forest, 2000, B. C. Tan [SING].

Mazosia phyllosema (Nyl.) Zahlbr. — (Se) ST 45937 [SING] vs.; (6a) SF

46105 [SING]; (10b) SMT 46185 [B, SING]; (14f) ST 46361 [B, SING].

Melanotrema aff. meiospermum (Hale) A.Frisch — (14a) ST 46276 [B,

SING].

A Conspectus of the Lichens (Lichenized Fungi) of Singapore 457

TLC: none. The material deviates from the description by Frisch and

Kalb (2006) by the convex columella.

Micarea cf. leprosula (Th.Fr.) Coppins & A.Fletcher — (3ax) in S 45844 [B,

SING] (in Micarea sp.), S 45845 [B, SING] (ster.).

Micarea cf. prasina Fr. — (2c) S 45545 [B, SING] c. apoth.

Micarea sp. A — (2c) S 45544 [B, SING]; (3n) S 45688 [B, SING]; (3w) S

45724 [B, SING].

Thalius granular, apothecia black.

Micarea sp. B — (3ax) S 45844 [B, SING].

Apothecia brown, without spores.

Micarea sp. C — (8) ST 46051 [B, SING]; (15) SM 46367 [B, SING].

Thallus finely granular, green and apothecia black.

Microtheliopsis uleana Mill.Arg. — (10b) SMT 46187 [B, SING].

Monoblastia pellucida Aptroot — (13) S 46249 [B, SING]: (14) ST 46326 [B,

SING].

Mycoporum eschweileri (Mill.Arg.) R.C.Harris — (2f) § 45614 [B, SING]?:

(3w) S 45728 [B, SING]?; (3ah) S 45788 [B, SING]?; (3an) S 45809 [B,

SING]?; (3az) S 45852 [B, SING]; (Se) ST 45925 [B, SING]?; (6) SM 45979

[SING]?; (7a) ST 46002 [SING]?; (8) ST 46063 [B, SING]: (15) SM 46369 [B,

SING]?; Kusu Island, 1989, Aptroot 25977, 25978 [Hb. Aptroot]; St. John’s

Island, 1989, Aptroot 25998 [Hb. Aptroot].

The material is often poorly developed and the identification

provisional. Ascospores uniseptate, 15-18(-22) x 3-4 um. Nr. 46369 is

pycnidiate; conidia 3 x 1 um, curved.

Mycoporum sp.? — (Se) ST 45879 [B, SING].

Myeloconis erumpens P.M.McCarthy & Elix — (6) SM 45948 [B, SING]: (10)

SMT 46123 [B, SING].

Myriotrema albocinctum Hale — Bukit Timah Nature Reserve, 1989, Aptroot

25988 [Hb. Aptroot].

Myriotrema glaucophaenum (Kremp.) Hale — (Se) ST 45897 [B, SING],

45910 [SING], 45926 [B]; (6) SM 45945 [B, SING].

TLC: psoromic acid (459/0).

Myriotrema microporellum (Nyl.) Hale — (Se) ST 45973 [B, SING].

TLC: hypoprotocetraric acid.

Myriotrema subconforme (Nyl.) Hale — (3b) S 45646 [SING]; (3c) S 45647

[SING]; (3m) S 45680 [B, SING]; (3p) S 45703 [B]; (3t) S 45716 [B, SING];

(3ah) S 45789 [SING]; (3ay) S 45849 [B, SING]; (4) S 45824 [SING]; Bukit

Timah nature reserve, 1989, Aptroot 25989 [Hb. Aptroot !].

TLC: none (45646, 45647, 45789, 45824, 45913).

Ocellularia cavata ( Ach.) Mill.Arg.—on the trunks of dead trees, 1861-1865,

Maingay [Nylander and Crombie, 1884: 53 as Thelotrema]; Vega expedition,

corticola, 1879, Almquist [Nylander, 1891: 19 as Thelotremal].

458 Gard. Bull. Singapore 61 (2) 2010

These records need reinvestigation to establish their current taxonomic

position.

Ocellularia crocea (Kremp.) Overeem & D.Overeem — (3n) S 45687 [B,

SING]; (3u) S 45778 [B, SING]; (3an) obs., in S 45810 [SING] (= Chiodecton

leptospermum); (3ay) S 45847 [SING]; (4) obs.; (Sa) ST 45857 [B, SING];

(Sd) ST 45875 [B, SING]; (Se) ST 45882 [B, SING], 45894 [SING], 45901

[SING], 45905 [B], 45918 [B, SING]; (6) ST 46382 [B, SING], SM 45953 [B],

45966 [SING], 45980 [SING], SF 46083 [B, SING]; (10) SMT 46118 [SING],

46127 [B], 46142 [B], 46146 [SING]; (10a) SMT 46163 [B, SING], 46165 [B,

SING]; (11) SMT 46214 [SING]; (14) ST 46330 [B]; (14e) ST 46313 [SING];

(15) SM 46363 [SING]; ad cortices, Beccari 261 |Krempelhuber, 1875: 60 as

Ascidium]; Sentosa, along the southern shore, on trunk, 1980, Tibell 8859

[UPS L-057801]; Bukit Timah nature reserve, 1989, Aptroot 25991 [Hb.

Aptroot].

Morphologically very similar to O. papillata, and hard to separate

when the characteristic, yellow pigment is scarce. TLC: traces (pigment)

(45687, 45718, 45857, 45918, 46083, 46127, 46165).

Ocellularia dolichotata (Nyl.) Zahlbr. — (6) SM 45972 [SING]; (10a) SMT

_ 46152 [B, SING]; Vega expedition, corticola, 1879, Almquist [H-Nyl. 22748

lectotype, S isolectotype, not seen; Nylander, 1891: 19 as Thelotrema; Hale,

1981: 303].

TLC: none (45972).

Ocellularia feigei Sipman — (6) SM 45964 |B holotype, SING isotype]; (10)

SMT 46121 [B}.

For details see Sipman (2003).

Ocellularia interponenda (Nyl.) Hale — (6) ST 46379 [SING]; (10) SMT

46129 [B, SING], 46140 [B, SING]; Vega expedition, corticola, 1879, Almquist

[ Nylander, 1891: 20 as Ascidium, type description]; Nu Soon Forest Reserve,

in tropical forest remnant, on trunk, 1980, Tibell 8834 [UPS L-057782].

Ocellularia nylanderiana Hale — Vega expedition, corticola, 1879, Almquist

[Nylander, 1891: 20 as Ascidium majorinum var. longius, type description].

Ocellularia orthomastia (Kremp.) Zahlbr. — (Se) ST 45912 [B, SING];

ad corticem, Beccari 247 [Krempelhuber, 1875: 60 as Ascidium, type

description].

TLC: none (45912).

Ocellularia papillata (Leight.) Zahlbr. — (3m) S 45675 [SING] (vs., poor

material); (4) S 45827 [SING], 45827 [B, SING]; (Se) ST 45903 [B, SING],

45915a [B, SING]; (Sf) ST 45928 [B, SING]; (6) ST 46381 [B, SING], SM

45951 [SING], 45955 [SING], 45958 [B, SING], 45969 [B], 45974 [SING], SF

46085 [B, SING], 46094 [SING]; (10) SMT 46132 [SING], 46137 [B], 46139

[SING]; (10a) SMT 46158 [B, SING], 46160 [B, SING] cf.; (11) SMT 46209

[SING]; (14) ST 46346 [B, SING]; (15) SM 46365 [SING]; Nu Soon Forest

A Conspectus of the Lichens (Lichenized Fungi) of Singapore 459

Reserve, in tropical forest remnant, on trunk of Lithocarpon, 1980, Tibell

8810 [UPS L-057767]; ibidem, on bark of Macaranga, 1980, Tibell 8827 [UPS

L-057777].

Very similar to O. crocea, and differring by the absence of pigment.

Since the pigment concentration seems variable, the delimitation between

the two species is not always clear. TLC: none (45827, 459/5a, 45928, 45969,

46085, 46137, 46346).

Ocellularia tanii Sipman — (5e) ST 45923 [B, SING]: (6) SF 46081] [B, SING],

46084 |B, SING]; (10a) SMT 46151 [B holotypus, SING isotypus].

For details see Sipman (2003).

Ocellularia terebrata (Ach.) Mill.Arg. — Vega expedition, corticola, 1879,

Almquist |Nylander, 1891:19 as Thelotrema terebratum typicum];ad corticem,

Beccari 255 [Krempelhuber, 1875: 60 as Thelotrema olivaceum].

Ocellularia terebrata (Ach.) Miill.Arg. f. swbeminescens (Nyl.) Zahlbr. —

Vega expedition, corticola, 1879, Almquist [Nylander, 1891:19 as Thelotrema

terebratum f. subeminescens].

This record needs reinvestigation to establish its current taxonomic

position.

Ocellularia triglyphica (Kremp.) Zahlbr. — Ad cortices, Beccari 256

[Krempelhuber, 1875: 60 as Ascidium].

Ocellularia xanthostromiza (Ny!.) Zahlbr. — Vega expedition, corticola,

1879, Almquist [Nylander, 1891: 20 as Ascidium, type description].

This record needs reinvestigation to establish its current taxonomic

position, it may be conspecific with O. crocea.

Ocellularia sp. A — (3p) S 45706 [B, SING}.

Ascospores muriform, 1/ascus. TLC: none.

Ocellularia sp. B — (Se) ST 45890 [B, SING].

Like O. papillata, but without columella. TLC: none.

Opegrapha graphidiza Ny|.— (8) ST 46072 |B, SING}.

Identification kindly provided by D. Ertz (Meise), for more details see

his forthcoming publication in Bibliotheca Lichenologica.

Opegrapha irosina Vain. — (4) S 45832 [B}.

Identification kindly provided by D. Ertz (Meise), for more details see

his forthcoming publication in Bibliotheca Lichenologica.

Opegrapha medusulina Ny|.— (15) SM 46376 [B, SING].

Identification kindly provided by D. Ertz (Meise), for more details see

his forthcoming publication in Bibliotheca Lichenologica.

Opegrapha subrimulosa Ny}. — (2a) S 45517 [B, SING]; (2d) S 45568 [B].

Identification kindly provided by D. Ertz (Meise), for more details see

his forthcoming publication in Bibliotheca Lichenologica.

Opegrapha subvulgata Ny\.— (1d) SM 45463 [B].

Identification kindly provided by D. Ertz (Meise), for more details see

460 Gard. Bull. Singapore 61 (2) 2010

his forthcoming publication in Bibliotheca Lichenologica.

Opegrapha varia Pers. — (11) SMT 46217 [B, SING].

Identification kindly provided by D. Ertz (Meise), for more details see

his forthcoming publication in Bibliotheca Lichenologica.

Opegrapha vegae R.Sant.— (Se) ST 45941 [SING]; (6) SM 45982 [B, SING],

45983 [B, SING], 45985 [B, SING]; (6a) SF 46104 [B, SING]; (10b) SMT

46174 [B, SING]; Vega expedition, foliicola, 1879, Almquist 37a [S, UPS,

not seen; Nylander, 1891: 22 as O. phyllobia; Santesson, 1952: 100]; 1879,

Almquist |UPS L-025868 isotypus, not seen].

Nr. 46174 deviates by the presence of 7-septate ascospores.

Pallidogramme chrysenteron (Mont.) Staiger, Kalb & Liicking — (14a)

ST 46274 [B, SING]; on bark of old trees, without ascospores, 1861-1865,

Maingay [Nylander and Crombie, 1884: 58 as Graphis chrysentera? |.

TLC: stictic acid (46274).

Parmeliella pannosa (Sw.) Mill.Arg. — Botanical Garden, on Cupressus,

1980, Tibell 88S89a [UPS L-057819]; Vega expedition, corticola, 1879, Almquist

[Nylander, 1891: 19 as Pannaria]; ad lignum (vel corticem) putridum, Beccari

234a [Krempelhuber, 1875: 60 as Pannarial.

Parmotrema gardneri (C.W.Dodge) Sérus. — (30) S$ 45701 [B, SING]; (14c)

ST 46282 |B, SING]; Botanic Gardens, 1949, Lemaitre [SING]; Botanical

Garden, on Peltophorum ferrugineum, 1980, Tibell 8879 [UPS L-057814];

ibidem, on Cupressus sp., 1980, Tibell 8891 [UPS L-057822].

TLC: atranorin, protocetraric acid (45701, 46282).

Parmotrema praesorediosum (Nyl.) Hale — (3ab) in S 45766 [SING] (=

Parmotrema tinctorum); (3aj) S 45804 [B, SING]; (12) SMT 46218 [SING];

(13) S 46262 [SING]; (14) ST 46324 [B]; (18) ST 46401 [B, SING]; Vega

expedition, corticola, 1879, Almquist [H-Nyl 35547 holotype, S isotype,

not seen; Nylander, 1891: 18 as Parmelia|; Government House Domain, on

telephone line insulator, 1959, Burkill 2130 [SING 500253].

TLC: atranorin, 3 fatty acids (45804, 46218, 46262, 46324, 46401).

Parmotrema saccatilobum (Tay|.) Hale — (3m) S 45684 [SING]; (3aj) S 45801

[B, SING]; (13) in S 46242 [SING] (= Trypethelium subeluteriae); (14c) ST

46290 [B, SING]; Elix, 1994: 156; Louwhoff and Elix, 1999: 111; on trunk in

rubber plantation, 2000, B.C. Tan [SING].

TLC: atranorin, protocetraric acid (45684, 45801, 46290).

Parmotrema tinctorum (Despr.ex Nyl.) Hale — (30) $ 45702 [B, SING]; (3ab)

obs., S 45766 [SING]; (3aj) S 45803 [B, SING]; (14) obs.; Singapore Botanic

Gardens, 1949, Lemaitre [SING]; Singapore Botanic Gardens, 1959, Burkill

2171 [SING]; on rocks, 1861-1865, Maingay [Nylander and Crombie, 1884: 51

as Parmelia; Nylander, 1891: 18 as Parmelia|; Vega expedition, corticola, 1879,

Almquist [Nylander, 1891: 18 as Parmelia|; Singapore Botanical Garden,

on Cedrela glaziovii (Nr. D6), 1959, Burkill 2176 [SING 500375]; ibidem,

A Conspectus of the Lichens (Lichenized Fungi) of Singapore 461

on Araucaria cunninghamii, 1980, Tibell 8874 [UPS L-057809]; ibidem, on

Peltophorum ferrugineum, 1980, Tibell 8876 [UPS L-057811]; ibidem, on

Harpullia zanguebarica, 1980, Tibell S882 [UPS L-057815]; ibidem, Jungle,

on Sterculia rubiginosa, 1980, Tibell 8870 [UPS L-058096].

TLC: atranorin, lecanoric acid (45702, 45766, 45803).

Phaeographina alutacea Zahlbr.— In horto botanico, Schiffner 3053 |W, not

seen; Redinger, 1936: 94].

This record needs reinvestigation to establish its current taxonomic

position.

Phaeographina flexans (Nyl.) INED. —Vega expedition, corticola, 1879,

Almquist |H-Nyl. 7975, not seen; Nylander, 1891: 24 as Lecanactis, type

description; Egea and Torrente, 1994: 183 as Graphidaceae].

This record needs reinvestigation to establish its current taxonomic

position. The species is poorly known and no formal recombination is

presented here without an investigation of the type.

Phaeographina prosiliens (Mont. & v.d.Bosch) Miill.Arg. — (10a) SMT

46155 [B, SING].

TLC: none.

Phaeographina subrigida (Ny|.) Zahlbr. — On Jack-fruit trees, 1861-1865,

Maingay [H, not seen; Nylander and Crombie, 1884: 58 as Graphis, type

description; Redinger, 1936: 98]; Vega expedition, corticola, 1879, Almquist

[Nylander, 1891: 24 as Lecanactis}.

These records need reinvestigation to establish their current taxonomic

position.

Phaeographis caesioradians (Leight.) A.W.Archer — (2) S 4562/a [SING];

(2e) S 45597 [B, SING]; (30) S 45700 [SING]; (3ag) S 45784 [SING]; (3ai) S$

45797 |B]; (Se) ST 45881 [SING]; (7) ST 46029 [B, SING], 46038 [B, SING],

in ST 46007 [SING] (= Diorygma rufopruinosum)? (no spores); (7a) ST

45998 [SING]: (8) ST 46075 [B, SING]; (13) S 46235 [SING]; (14a) ST 46268

[B, SING], 46269 [B, SING]; (14c) in ST 46283 [B, SING] (= Phaeographis

intricans); (15) in SM 46370 [B, SING] (= Lecanora helva).

TLC: none (45597, 45621a, 45700, 45797, 45881, 46029, 46075, 46269).

Externally similar to Phaeographis sp. C, but with muriform ascospores.

Phaeographis circumscripta (Kremp.) Zahlbr. — Pulo Pusang, ad cortices,

Beccari 243b [M, not seen; Krempelhuber, 1875: 62 as Graphis (Fissurina);

Redinger, 1936: 78].

This record needs reinvestigation to establish its current taxonomic

position.

Phaeographis dendroides (Leight.) Miill.Arg. — (3ab) S 45759 [B, SING].

TLC: stictic acid.

Phaeographis diversula (Nyl.) Zahlbr. — Vega expedition, corticola, 1879,

Almquist [Nylander, 1891: 24 as Lecanactis diversa}.

462 Gard. Bull. Singapore 61 (2) 2010

Lecanactis diversas not synonymized here with Graphina obtrita (Fée)

Mill.Arg. as indicated by Zahlbruckner (1923: 416), because its ascospores

seem in conflict with this. Instead the synonymisation of “Graphis diversa

Kremp.” in Redinger (1936: 89) is followed. The record needs reinvestigation

to establish its current taxonomic position.

Phaeographis intricans (Nyl.) Staiger — (1) in SM 45473 [SING] (=

Phaeographis sp. C); (1g) SM 45487 [B, SING]; (1j) SM 45508 [SING];

(2) S 45622 [SING], S 45623 [B, SING]; (2b) obs., in S 45531 [B, SING] (=

Trypethelium eluteriae); (2d) obs.,in S 45556 [SING] (= Pyxine cocoes); (2e)

S 45587 [B, SING], 45594 [B, SING]; (2f) S 45602 [B, SING], 45607 [B, SING];

(2g) in S§ 45630 [SING] (= Lecanora helva); (30) obs., S 45698 [SING]; (3p)

S 45704 [SING]; (3ab) S 45756 [SING]; (3a0) S 45812 [B]; (Se) ST 45880 [B,

SING]; (6) SM 45967 [SING]; (7) obs., ST 46011 [B, SING], 46079 [SING],

46032 [B, SING], 46034 [B], 46037 [B, SING]; (7a) ST 45996 [B, SING]; (8) ST

46073 |B, SING]; (13) in S 46260 [SING] (= Dirinaria picta); (14) obs.; (14a)

ST 46271 |B, SING]; (14c) ST 46283 [B, SING]; (15) obs.; (18) obs.; Sentosa,

along the southern shore, on trunk,1980, Tibell 8857 [UPS L-057799]; Kusu

Island, 1989, Aptroot 25980, 25982 [Hb. Aptroot].

TLC: none (45487, 45587, 45602, 46073).The material is very variable

and the identification provisional. Noteworthy is the presence of specimens

with guttulate hymenium: in 45473, 45508, 45623, 45704, 45996.

Phaeographis maeandrata (Kremp.) Zahlbr. — Vega expedition, corticola,

1879, Almquist [Nylander, 1891: 24 as Lecanactis serpentosa].

This record needs reinvestigation to establish its current taxonomic

position.

Phaeographis punctiformis var. nylanderi (Vain.) Redgr.? — (2e) S 45593 [B,

SING], 45599 [B, SING]; (2g) in S 45630 [SING] (= Lecanora helva); (8) ST

46050 [B, SING]; (13) S 46234 [SING]; (13a) S 46225 [SING]; Sentosa, along

the southern shore, on trunk, 1980, Tibell 8858 pp [UPS L-057800, Graphis

insulana}.

TLC: norstictic acid (45593, 46050, 46234). The material deviates

clearly from P. punctiformis s. str. and the identification is provisional.

Phaeographis ramigera Redgr. — Beccari 268 [M, not seen; Redinger, 1936:

male

This record needs reinvestigation to establish its current taxonomic

position.

Phaeographis scalpturata (Ach.) Staiger — (3ag) S 45783 [SING]; (3ai) S$

45791 [B, SING]; (Se) ST 45887 [B, SING]; (10a) SMT 46157 [B, SING];

Sentosa, along the southern shore, on ae 1980, Tibell 8856 [UPS

L-057798].

TLC: none (45791, 45887, 46157). The identification of 45783 is

uncertain in the absence of ascospores.

A Conspectus of the Lichens (Lichenized Fungi) of Singapore 463

14 : § / Sa]

Phaeographis sp. A — (3ak) S 45805 [B, SING].

Ascospores submuriform, ca 25 x 10 um: TLC. norstictic acid.

Phaeographis sp. B — Benjamin Lee {SING}.

Phaeographis sp. C — (1) SM 45473 [SING]: (2) S 45621 [B, SING]: (2f) S

45608 |B, SING]; (3ad) S 45775 [B, SING].

Externally similar to Phaeographis caesioradians, but with transversely

septate ascospores. TLC: none (45608, 45621, 45775).

Phaeographis sp. D — (14e) ST 46306 [B, SING].

Ascospores bacillar, ca 4/ ascus, 30-50 x 10-12 um; TLC: none.

Phyllopsora parvifolia (Pers.) Mill.Arg.—ad lignum putridum, Beccari 233,

234 |Krempelhuber, 1875: 60 as Lecidea parvifolia var. fibrillifera].

Phyllopsora? sp. — (14) ST 46328 [B, SING].

TLC: stictic acid. A squamulose lichen deviating from Phyllopsora by

the absence of prothallus and from Eschatogonia by the absence of a lower

cortex; in the absence of apothecia its affinities are unclear.

Physcia cf. tribacioides Ny|. — (3ab) S 45768 [B, SING]; (3aj) S 45800 [B,

SING].

Physcia sp. — (3d) S 45653 [B, SING]: (3x) S 45729 [B, SING]; (14) ST 46264

[B, SING], 46320 [B, SING], 46332 [B, SING]: (18) ST 46400 [B, SING]; Nu

Soon forest reserve, in tropical forest remnant, on trunk, 1980, Tibell 8845

[UPS L-057788]; St. John’s Island, 1989, Aptroot 26003 [Hb. Aptroot].

A small-lobed species with marginal soralia and without lower cortex.

It is apparently widespread and common in cultivated areas in the Malesian

area and specimens are available in B from Indonesia and the Philippines.

Physma byrsaeum (Ach.) Tuck. — Vega expedition, corticola, 1879, Almquist

[Nylander, 1891: 18 as Dichodium byrsinum]; Singapore Botanical Garden,

treestem, 1920, Kiah 6038 [SING].

Platygramme flexuosa (Ny|.) INED.-— (7) ST 46045 [SING]; (7a) ST 45997

[B, SING]: (8) ST 46070 [B, SING]: Sentosa, along the southern shore, on tree

trunk, 1980, Tibell 8855 [UPS]; Vega expedition, corticola, 1879, Almquist

[Nylander, 1891: 25 as Lecanactis; Redinger, 1936: 98 as Phaeographina].

TLC: none (45997, 46070). No formal combination is presented

without investigation of the type.

Platythecium sp.? — Beccari 235 [M, not seen: Krempelhuber, 1875: 61 as

Graphis grammitis var. seductilis, type description; Redinger, 1936: 62] as

Graphis grammitis var. seductilis.

This record needs reinvestigation to establish its current taxonomic

position.

Platythecium sp. — (6) SM 45968 {B, SING].

TLC: stictic acid.

Pocsia mucronata P.M.McCarthy — (la) SM 45441b [B, SING], 4544/e [B.

SING].

464 Gard. Bull. Singapore 61 (2) 2010

Polyblastiopsis augescens (Nyl.) Zahlbr. — On bark of trees, 1861-1865,

Maingay [Nylander and Crombie, 1884: 60-61 as Verrucaria, type description;

Nylander, 1891: 25 as Verrucaria].

Porina atropunctata Liicking & Vézda — (10b) SMT 46190a |B, SING],

46190b [B, SING].

Porina cf. canthicarpa P.M.McCarthy — (10b) SMT 46180 [B, SING].

Algal cells rectangular, in closed, radiating plates; ascospores 3-septate,

20'x 2-5 [mt

Porina chlorotica (Ach.) Mill.Arg. — (16) ST 46386 [B, SING].

On granitic rock.

Porina epiphylla (Fée) Fée — (10b) SMT 46190 [B, SING], 46190c [B,

SING].

Porina internigrans (Nyl.) Mill.Arg. — (Se) ST 45920 [B, SING]; (6) SM

45960 |B, SING]; (10) SMT 46134 [B, SING]; (10a) SMT 46148 [B, SING].

Porina mirabilis Liicking & Vézda — (10b) SMT 46167 [SING] cf., 46195

[SING].

Porina monocarpa (Kremp.) Schilling — ad folia coriacea, Beccari 269e

[Krempelhuber, 1875: 63 as Verrucaria,|; Beccari 269b [M, holotype, G, not

seen; Krempelhuber, 1875: 63 as Verrucaria; Santesson, 1952: 256].

Porina semecarpi Vain. — (10b) SMT 46181] [B, SING].

Porina tetracerae (Malme) R.Sant. — (1d) in SM 45460 [SING] (= Pyrenula

sp.), SM 45467 [B, SING]; (1j) SM 45512 [B, SING]; (3m) S 45678 [SING];

(3ac) in S 45773 [B] (= Porina cf. tetracerae); (4) S 45816 [SING], in S 45822

[B, SING] (= Thelotrema sp. B), S 45830 [B, SING]; (Se) ST 45919 [B, SING];

(6) SM 45961 [B, SING]; (6a) SF 46113 [SING]; (10) SMT 46138 [B, SING];

(10b) SMT 46179 [B, SING].

While most specimens are isidiate and without ascocarps, the frequency

of ascocarps and isidia is very variable and there seemed no reason to

separate isidiate from non-isidiate specimens as proposed by some other

authors.

Porina cf. tetracerae (Malme) R.Sant. — (3m) in § 45678 [SING] (= Porina

tetracerae); (3ac) S 45773 [B]; (4) S 45825 [B, SING]; (13) S 46252 [B, SING];

(14) ST 46334 [B, SING].

These specimens deviate by the absence of cortex on the propagules,

which remain usually short and granular. No specimen with ascocarps

was seen, so that the classification is tentative and based on superficial

resemblance.

Porina tetramera Vain. — (17a) ST 46388c [B, SING].

Porina virescens (Kremp.) Miill.Arg. — (10b) SMT 46192 [B, SING].

Porocyphus sp.? — (2) S 45582 [B, SING].

A gelatinous algal cover on concrete, which may be lichenized because

a few ascocarps seem present.

A Conspectus of the Lichens (Lichenized Fungi) of Singapore 46

Psorotichia sp.? — (2) in S 45584 [SING] (= Endocarpon pallidum).

Pyrenula anomala (Ach.) Vain. — on trunks of trees, 1861-1865, Maingay

[Nylander and Crombie, 1884: 61 as Trypethelium platystomum|; ad cortices,

Beccari 236b [Krempelhuber, 1875: 63 as Trypethelium platystomum]; Vega

expedition, corticola, 1879, Almquist [Nylander, 1891: 25 as Verrucaria

aggregata].

Pyrenula aspistea (Ach.) Ach. — (5e) ST 45904 [B, SING]: (5f) ST 45930

[B. SING]: (6) SM 45978 [SING]. SF 46092 [B, SING]: (11) in SMT 46210

[B, SING] (= Pyrenula santensis) (no spores), SMT 46212 [B. SING]: Bukit

Timah nature reserve, 1989, Aptroot 25992 [Hb. Aptroot].

Pyrenula concatervans (Ny\.) R.C.Harris — (1a) SM 45443 [B, SING]: (11)

SM 45502 [B, SING]; (3a) S 45640 [B, SING]: (8) ST 46059 [SING]: (13a) S

46222 [B].

The ascospores are variable, from 4-loculate and 6-loculate to

submuriform, 6 x 1-2 locules, and muriform, 8 x 2-3 locules, and range in size

from 18 x 8 to 25 x 10 um. However, they are much smaller than the related

species with muriform ascospores. P. macularis (Zahlbr.) R.C.Harris.

Pyrenula mamillana ( Ach.) Trev. — (14) ST 4634] [B, SING].

Pyrenula nitidula (Bres.) R.C.Harris — (2f) S 45610 [B, SING].

Pyrenula ochraceoflava (Ny|.) R.C.Harris — (la) SM 45442 [B, SING]:

(1c) in SM 45456 [SING] (= Chrysothrix xanthina): (le) SM 45476 [SING]:

(1g) SM 45492 [SING]: (11) SM 45499 [B]: (1j) SM 45506 [SING]. 45510 [B.

SING]; (2a) in S 45519 [B] (= Graphis caesiella); (2d) S 45570 [B, SING]:

(2g) in S 45624 [SING] (Pyrrhospora quernea), in S 45631 [B] (Amandinea

efflorescens), in S 45635 [SING] (parasite): (3) obs.: (3a) in S 45642 [SING]

(= Graphis insulana); (3ab) S 45742 [B], 45743 [SING]: (3ag) S 45782

[SING]: (3av) S 458417 [SING]: (5c) ST 45870 [SING]: (7) obs., ST 46044 [B].

46047 [SING]: (7a) ST 46000 [B]: (8) ST 46056 [SING], 46057 [B]: (13) obs.:

(13a) S 46221 [SING]: (14) obs., ST 46342 [B], 46343 [SING]: (14c) ST 46295

[SING]: (15) obs., SM 46377 [B, SING]; Fort Canning Park, 1989, Aptroot

25958 [Hb. Aptroot]; Sentosa, along the southern shore, outskirts of forest

along the beach, on trunk, 1980, Tibell 8862 [UPS L-057804]: Fort Canning

Park, 1989, Aptroot 25960 [Hb. Aptroot].

This species is conspicuous by the presence of yellow to red

anthraquinone pigments. However, their concentration is very variable,

perhaps depending on vitality of the thallus and shading. Some specimens

seem to lack pigments completely, while in others it is restricted to a reddish

pigment on the ascocarps. Both look very different from the modal form

with yellowish thallus and perithecia but agree in anatomical characters.

TLC: 2 anthraquinones (45499, 46000): 4 anthraquinones (45570).

Pyrenula santensis (Ny|.) Mill.Arg.? — (11) SMT 46210 [B, SING].

The specimen lacks ascospores.

466 Gard. Bull. Singapore 61 (2) 2010

Pyrenula santensis var. murina INED. — ad cortices, Beccari 257, 265

[Krempelhuber, 1875: 63 as Verrucaria santensis Tuck. var. murina].

This record needs reinvestigation to establish its current taxonomic

position.

Pyrrhospora quernea (Dicks.) Korb. — (2f) S 4561/8 [B, SING]; (2g) S 45624

[SING], in S 45637 [B, SING] (= Lecanora helva); (7a) ST 45987 [B, SING];

(7b) ST 46004 [B, SING]; (8) ST 46049 [B, SING]; (13) S 46248 [B, SING];

(14) obs.; (15) obs.; (18) ST obs.

TLC:thiophanicacid,tr.arthothelin (45618, 45987, 46004, 46049, 46248).

Surprisingly this material is morphologically and chemically identical with a

species known until recently only from temperate and mediterranean climate

areas, where it is not uncommon. Apothecia, desirable for confirmation of

the taxonomic position, were not found. It is also reported from Hong Kong

by Aptroot and Seaward (1999).

Pyxine cocoes (Sw.) Nyl. — (1j) SM 45514 [B, SING]; (2d) S 45556 [SING];

(13) S 46259 [B, SING]; (13a) S 46227 [SING]; (15) obs.; Fort Canning Park,

1989, Aptroot 25950 [Hb. Aptroot]; Sentosa, close to the ferry, on trunk of

broad-leved tree, 1980, Tibell 8846, 8847 [UPS L-057789, L-057790]; ibidem,

on trunk of Tamarindum indica, 1980, Tibell 8849 [UPS L-057792].

Pyxine farinosa Kashiw. — (3a) in S 45643 [B, SING] (= Cryptothecia scripta),

S 45644 [B]: (3aj) S 45798 [SING]; (8) ST 46068 [B, SING]; (14c) ST 46281

[SING]; Pulau Terkukor, 1960, Burkill 2566 [SING]; 1949, Lemaitre [SING];

St. John’s Island, 1989, Aptroot 26001 [Hb. Aptroot]; Kusu Island, 1989,

Aptroot 25965 [Hb. Aptroot].

Ramonia microspora Vézda — (14) ST 46350 [SING].

Rinodina cinereovirescens (Harm.) Zahlbr. — (14) ST 46265 [B, SING].

Rinodina oxidata (A.Massal.) A.Massal.— (14) ST 46267 [B, SING].

TLC: atranorin, gyrophoric acid.

Sarcographa concisa (Kremp.) Zahlbr. — ad cortices, Beccari 262 [M, not

seen; Redinger, 1936:108; Krempelhuber, 1875: 31,61 as Graphis concisa].

This record needs reinvestigation to establish its current taxonomic

position.

Sarcographa heteroclita (Mont.) Zahlbr. — (3q) S 45712 [B, SING]; (3af) S

45780 [SING]; (Se) ST 45892 [SING]; (10) SMT 46115 [B, SING]; (11) SMT

46213 [B, SING].

Two specimens have pycnidia and lack ascocarps, 45780 and 45712;

conidia 5-6 x 0.5 um, slightly curved. TLC: stictic, tr. hypostictic acid (45892,

46115); stictic acid (46213).

Sarcographa labyrinthica (Ach.) Miill.Arg. — (Se) ST 45896 [SING], 459/71

[B]; (10) SMT 46117 [B, SING]; (14e) ST 46304 [SING]; Bukit Timah, 1959,

Burkill 2172 [SING]; on Betel-palms, 1861-1865, Maingay [Nylander and

Crombie, 1884: 59 as Glyphis|; Vega expedition, corticola, 1879, Almquist

A Conspectus of the Lichens (Lichenized Fungi) of Singapore 467

[Nylander, 1891: 25 as Glyphis; Redinger, 1936: 107]; Pulau Pinang, ad

corticem, Beccari 241 [Krempelhuber, 1875: 62 as Glyphis labyrinthica

var. insulata|; Nee Soon forest reserve, tropical forest remnant, on bark of

Macaranga, 1980, Tibell 8826 [UPS L-057776].

TLC: stictic, tr. cryptostictic acids (45896, 46304).

Sarcographa lactea Miill.Arg. — Beccari 241 |M, not seen; Redinger 1936:

109].

This record needs reinvestigation to establish its current taxonomic

position.

Sarcographa protracta (Kremp.) Zahlbr. — Pulo Pinang, ad cortices,

Beccari 242 {M, not seen; Redinger, 1936: 110; Krempelhuber, 1875: 63 as

Acanthoglyphis, type description].

This record needs reinvestigation to establish its current taxonomic

position. Probably this is not a Singapore record.

Sarcographa ramificans (Kremp.) Staiger — (Se) ST 45914 [B, SING]; (10a)

SMT 46162 |B, SING]; ad cortices, Beccari 268 [Krempelhuber, 1875: 61

as Graphis|; Nu Soon Forest Reserve, tropical forest remnant, on trunk

of Macaranga, 1980, Tibell 8807, 8825 [UPS L-057765, L-057775]; Vega

expedition, corticola, 1879, Almquist [H, not seen; Nylander, 1891: 25 as

Glyphis duodenaria, type description; Redinger, 1936: 112, as S. heteroclita

var. duodenaria}.

TLC: stictic acid (46/62).

Sarcographa tricosa (Ach.) Miill.Arg.—on Betel-palms, 1861-1865, Maingay

[Nylander and Crombie, 1884: 58 as Medusula]; Vega expedition, corticola,

1879, Almquist [Nylander, 1891: 25 as Medusula].

Sarcographina glyphiza (Nyl.) K.Singh & Awasthi— (30) S$ 45695 [B, SING];

(3v) S 45720 [SING]; (3y) S 45734 [B]; (3ab) S 45760 [B, SING]; (3ai) S 45796

[SING]; (4) S 45837 [B]; (6) SF 46088 [SING]; (14c) ST 46284 [B]; (14e)

ST 46303 [SING]; Singapore Botanical Garden, on Harpullia zanguebarica,

1980, Tibell 8883 [UPS L-057816]; ad corticem Arecae, Maingay [H, not

seen; Redinger, 1936: 113 as Sarcographina gyrizans]; on Betel-palms, 1861-

1865, Maingay [Nylander and Crombie, 1884: 59 as Glyphis circumplexa,

type description; Nylander, 1891: 25 as Glyphis circumflexa].

TLC: stictic, tr. cryptostictic acids (45695, 45760, 46088). This species is

not treated by Staiger (2002), and its generic position in the new systematic

arrangement of Graphidaceae seems unsettled.

Sclerophyton dendrizans (Ny}.) Zahlbr. — Vega expedition, corticola, 1879,

Almquist [Nylander, 1891: 23 as Chiodecton, type description; Redinger,

1936: 117].

This record needs reinvestigation to establish its current taxonomic

position. Sparrius (2004) suggests its affinity to Phaeographis but did not

investigate the type.

468 Gard. Bull. Singapore 61 (2) 2010

Septotrapelia triseptata (Hepp) Aptroot — Economic Gardens, 1920,

Hippance 5935 [SING]; Reformatory Road, 1920, Chipp 6191 [SING].

Sporopodium sp. — (10b) SMT 46198 [B, SING]; (14f) ST 46360 [B, SING];

(17a) ST 46388d |B, SING]; Nee Soon forest reserve, tropical forest remnant,

on leaves of Nephelium lappaceum, 1980, Tibell 8803 [UPS L-057764].

The specimens have campylidia and no apothecia, and cannot be

identified with certainty at species level. The may belong to more than one

species.

Stegobolus berkeleyanus Mont. -— (6) SM 45947 [B, SING].

Stegobolus croceoporus (Hale) A.Frisch — (10) SMT 46131 [SING].

Strigula concreta (Fée) R.Sant. — (Se) ST 45944 [B, SING]; (6a) SF 46111

[SING]; (14f) ST 46357 [B, SING].

Strigula ys. nemathora Mont. — (6a) SF 46109 [B]; (10b) SMT 46201

[SING].

The identification is tentative because the specimens are in poor

condition.

Strigula nitidula Mont. — (1a) SM 45441f |B, SING]; (6a) SF 46/10 [B,

SING]; (10b) SMT 46200 [B, SING].

The identification of 46//0 is tentative because it contains only

pycnidia.

Strigula orbicularis Fr.— (1a) in SM 45441f [B, SING] (= Strigula nitidula);

(15) SM 46364 [B, SING].

Strigula smaragdula Fr. — (1a) SM 45441d [B, SING]; 1879, Almquist 38 [S,

not seen; Santesson, 1952: 169 as S. elegans].

Strigula sp.? — Vega expedition, foliicola, 1879, Almquist [Nylander, 1891: 26

as Strigula actinoplaca}.

This record needs reinvestigation to establish its current taxonomic

position.

Syncesia cf. glyphisoides (Fée) Tehler — (14) ST 46345 [B, SING].

TLC: lichexanthone, traces.

Thalloloma sp. A — (6) SM 45959 [B, SING].

TLC: indet. substance.

Thelidium quinqueseptatum (Nyl.) Arnold — Vega expedition, corticola,

1879, Almquist [Nylander, 1891: 26 as Verrucaria, type description.

Thelotrema aggregatum (Hale) Hale — Nee Soon Forest Reserve, tropical

forest remnant, on bark of Gonystylis, 1980, Tibell 8820 [UPS L-057774].

Thelotrema diplotrema Nyl.— (Se) ST 45902 [B, SING].

TLC: none.

Thelotrema isidiophorum (Kremp.) Zahlbr. — ad cortices, Beccari 264

[Krempelhuber, 1875: 43, 60 as Ascidium; Krempelhuber, 1877: 134 as

Ascidium|.

Thelotrema santessonii Hale — (10) SMT 46120 [B, SING].

A Conspectus of the Lichens (Lichenized Fungi) of Singapore 469

TLC: stictic acid.

Thelotrema sp. A — Nee Soon Forest Reserve, tropical forest remnant, on

bark of Garcinia, 1980, Tibell 8809 [UPS L-057766].

Thelotrema sp. B — (4) S 45822 [B, SING].

Ascospores muriform, hyaline, 8/ascus, ca. 60 x 20 um, I-negative.

Thysanothecium scutellatum (Fr.) D.J. Gallow. — (3p) S 45707 [B, SING];

(3ap) S 45813 [B, SING]; (14c) ST 46293 [B, SING].

TLC: divaricatic acid (45813, 45707).

Trapelia cf. involuta (Tayl.) Hertel — (3) S 45814 [SING].

Tricharia sp. — (Se) in ST 45942 [SING] (= Anisomeridium throwerae): (10b)

SMT 46188 [SING]; ad folia coriacea, Beccari 232a, 269d [Krempelhuber,

1875: 64 as T: orbicularis, type description; Santesson, 1952: 379].

The new collections lack apothecia.

Trypethelium deformis Makhija & Patwardhan — 1861-1865, Maingay 165

[BM holotype, not seen; Makhija and Patwardhan, 1992: 240].

Trypethelium eluteriae Spreng. — (2b) S 45531 [B, SING]; (2e) S 45588

[SING]; (2f) S 45606 [B, SING]; (2g) in S 45630 [SING] (= Lecanora helva);

(3) obs.; (7) ST 46020 [SING], 46022 [SING]; (14) ST 46338 [SING]; (14a)

ST 46272 [B]; (18) in ST 46405 [SING] (= Trypethelium subeluteriae); Fort

Canning Park, 1989, Aptroot 25957 [Hb. Aptroot]; nahe der Philippinischen

Botschaft, 1999, Schumm & Schwarz |Hb. Schumm 5850].

The colour of 46272 suggests that it has a deviating pigment

composition.

Trypethelium epileucodes Ny}\. — (3) S 45655 [SING]; (3g) S 45659 [B];

(30) S 45693 [B]; (3aa) S 45739 [SING]; (Se) ST 45885 [B, SING], 45924

[SING]; (8) ST 46061 [SING]; (10) SMT 46116 [SING]; (10a) SMT 46150

[B, SING], 46/54 [SING]; Bukit Timah nature reserve, 1989, Aptroot 25993

[Hb. Aptroot].

Trypethelium cf. platystomum Mont. — (2b) S 45533 [B, SING]; (2e) S 45595

[SING]; (2f) S 45605 [B]; (8) ST 46060 [SING]; (13) S 46242 [SING]; (14) ST

46344 |B]: (14e) ST 46311 [B]; (18) ST 46405 [SING].

Trypethelium stramineum Kremp.—ad corticem, Beccari 246 |Krempelhuber,

1875: 63].

This record needs reinvestigation to establish its current taxonomic

position.

Trypethelium straminicolor Ny|.— Vega expedition, corticola, 1879, Almquist

[Nylander, 1891: 26].

This record needs reinvestigation to establish its current taxonomic

position.

Trypethelium tropicum (Ach.) Miill.Arg. — (1b) in SM 45452 [B, SING] (=

Graphis caesiella); (1g) SM 45488 [B, SING]; (2b) S 45522 [B, SING]; (2e) in

S 45595 [SING] (= Trypethelium subeluteriae) (no spores); (3k) in S 45668

470 Gard. Bull. Singapore 61 (2) 2010

[B, SING] (= Enterographa pallidella);(3q) S 45711 |B, SING]; (3av) S 45842

[SING]; (Se) in ST 45904 [B, SING] (Pyrenula aspistea) (no spores); (8) in

ST 46060 [SING] (= Trypethelium subeluteriae) (no spores); (13) obs.; (13a)

S 46223 [SING]; (18) ST 46406 [B, SING]; Fort Canning Park, 1989, Aptroot

25956 [Hb. Aptroot]; on wild Lime-trees, 1861-1865, Maingay [Nylander and

Crombie, 1884: 60 as Verrucaria; Nylander, 1891: 25 as Verrucaria]; nahe der

Philippinischen Botschaft, 1999, Schumm & Schwarz (Hb. Schumm 6689);

Botanical Garden, on trunk of Pangium edule, 1964, Degelius As-497 [UPS

L-056602]; crossing Oxley Road/Oxley Rise, roadside trees, 1994, Diederich

12224 [Hb. Aptroot].

Trypethelium variolosum Ach. — (2b) S 45532 |B, SING], 45538 [B, SING];

(3) S 45656 [B]? (no spores; c. pycn.); (3q) S 45710 [B, SING]; (3ab) S 45746

[SING], 45747 [B]; (3ad) S 45774 [SING]; (3ah) S 45787 [SING]; (Se) ST

45888 |B]; (13) S 46241 [B]; (14c) ST 46294 [SING]; (18) ST 46403 [B], 46404

[B, SING]; Vega expedition, corticola, 1879, Almquist |Nylander, 1891: 26 as

T. ochroleucum|.

Trypethelium sp. A — Sentosa, along the southern shore, on trunk, 1980,

Tibell 8860 [UPS L-057802].

Tylophoron sp. — (1h) SM 45493 [B, SING].

The specimen lacks ascocarps and contains thalloconidia.

Verrucaria mundula PM.McCarthy — (11) SM 45500 [B, SING].

Vezdaea sp.? — (2) S 45586 [B, SING].

The specimen has immature ascocarps without spores and the

identification is therefore uncertain.

A Conspectus of the Lichens (Lichenized Fungi) of Singapore 47]

Figure 1. Some of the commonest lichens in Singapore. A. Arthonia catenatula (SM 45498): B.

Dirinaria picta (SM 45454): C. Pyrenula ochraceoflava (S 45499): D. Phaeographis intricans

(ST 46283): E. Chrysothrix xanthina (S 45603): F. Graphis insulana (S 45679). Scale for all:

bar = 1 mm.

472 Gard. Bull. Singapore 61 (2) 2010

Figure 2. A-C. Forest-inhabiting lichens. A. Ocellularia crocea (ST 45905); B. Myriotrema

subconforme (S 45646); C. Porina tetracerae (ST 45919). D. Chroodiscus australiensis, a

foliicolous lichen (SMT 46193). E-F. Two lichen species recently described from Singapore.

E. Lepraria usnica (S 45651); F. Herpothallon granulare (SM 45490). Scale for all: bar = 1

mm.

A Conspectus of the Lichens (Lichenized Fungi) of Singapore 473

hae |

We BER

2S Nye

Figure 3. Diversity of Graphidaceae in Singapore. A. Diorygma rufopruinosum (SM 46373);

B. Dyplolabia afzelii (S 45697); C. Fissurina cf. dumastii (S 46456); D. Glyphis cicatricosa (S

45785); E. Glyphis scyphulifera (S 46220); F. Graphis glaucescens (S 45562); G. Graphis libra-

ta (SM 45462); H. Pallidogramme chrysenteron (ST 46274); 1. Phaeographina prosiliens (SM

46155); J. Phaeographis caesioradians (ST 46029): K. Phaeographis punctiformis var. nylan-

deri (S 46234): L. Sarcographa heteroclita (SMT 46115). Scale for all pictures: bar = 1 mm.

474 Gard. Bull. Singapore 61 (2) 2010

Synonyms used in published records

Acanthoglyphis protracta Kremp. = Sarcographa protracta.

Arthonia aleurella Nyl. = Cryptothecia aleurella.

Arthothelium aleurellum (Nyl.) Zahlbr. = Cryptothecia aleurella.

Ascidium croceum Kremp. = Ocellularia crocea.

Ascidium interponendum Ny\l. = Ocellularia interponenda.

Ascidium isidiophorum Kremp. = Thelotrema isidiophorum.

Ascidium majorinum Nyl. var. longius Nyl. = Ocellularia nylanderiana.

Ascidium orthomastium Kremp. = Ocellularia orthomastia.

Ascidium triglyphicum Kremp. = Ocellularia triglyphica.

Ascidium xanthostromizum Nyl. = Ocellularia xanthostromiza.

Buellia efflorescens Mill. Arg. = Amandinea efflorescens.

Chiodecton dendrizans Nyl. = Sclerophyton dendrizans.

Coccocarpia ciliolata Mont. = Coccocarpia erythroxyli.

Cryptothecia granularis Sipman = Herpothallon granulare.

Dichodium byrsinum (Ach.) Nyl. = Physma byrsaeum.

Graphis afzelii Ach. = Dyplolabia afzelii.

Glyphis circumflexa Nyl. (misspelling for “circumplexa”) = Sarcographina

glyphiza.

Glyphis circumplexa Nyl. = Sarcographina glyphiza.

Glyphis duodenaria Ny\. = Sarcographa ramificans.

Glyphis labyrinthica Ach. = Sarcographa labyrinthica.

Glyphis labyrinthica var. insulata Kremp. = Sarcographa labyrinthica.

Graphis atroalba Kremp. = Dyplolabia afzelii.

Graphis chrysentera Mont. = Pallidogramme chrysentera.

Graphis circumscripta Kremp. = Phaeographis circumscripta.

Graphis concisa Kremp. = Sarcographa concisa.

Graphis ramificans Kremp. = Sarcographa ramificans.

Graphis singaporina Nyl. = Carbacanthographis candidata.

Graphis subrigida Nyl. = Phaeographina subrigida.

Gyalecta lutea Dicks. = Coenogonium luteum.

Lecanactis diversa Nyl. = Phaeographis diversula.

Lecanactis flexans Nyl. = Phaeographina flexans.

Lecanactis flexuosa Nyl. = Platygramme flexuosa.

Lecanactis serpentosa Nyl. = Phaeographis maeandrata.

Lecanactis subrigida (Nyl.) Nyl. = Phaeographina subrigida.

Lecanora epiphylla Auct., non Fée = Byssoloma tricholomum.

Lecanora punicea Ach. = Haematomma sp.

Lecidea parvifolia Pers. var. fibrillifera Nyl. = Phyllopsora parvifolia?

Lecidea rubello-virens Ny\. = Bacidia rubellovirens.

Lecidea sophodina Nyl. = Lopadium sophodinum.

A Conspectus of the Lichens (Lichenized Fungi) of Singapore 475

Medusula tricosa (Ach.) = Sarcographa tricosa.

Opegrapha phyllobia auct., non Nyl. = Opegrapha vegae.

Pannaria pannosa (Sw.) Nyl. = Parmeliella pannosa.

Parmelia praesorediosa Nyl\. = Parmotrema praesorediosum.

Parmelia tinctorum Despr. = Parmotrema tinctorum.

Phaeographina flexuosa (Nyl.) Mill.Arg. = Platygramme flexuosa.

Physcia picta (Sw.) Nyl. = Dirinaria picta.

Physcia picta (Sw.) f. sorediata (Hepp) = Dirinaria applanata.

Sarcographa heteroclita (Mont.) Zahlbr. var. duodenaria (Nyl.) Redgr. =

Sarcographa ramificans.

Sarcographina gyrizans (Leight.) Miill.Arg. = Sarcographina glyphiza.

Strigula actinoplaca Nyl. = Strigula sp.?

Strigula elegans (Fée) Mill.Arg. = Strigula smaragdula.

Thelotrema cavatum Ach. = Ocellularia cavata.

Thelotrema dolichotatum Nyl. = Ocellularia dolichotata.

Thelotrema olivaceum Mont. = Ocellularia terebrata.

Thelotrema terebratum Ach. = Ocellularia terebrata.

Thelotrema terebratum Ach. f. subeminescens Nyl. = Ocellularia terebrata f.

subeminescens.

Tricharia orbicularis Kremp. = T. sp.

Trypethelium ochroleucum Eschw. = Trypethelium variolosum.

Trypethelium ochrothelizum Nyl. = Astrothelium ochrothelizum.

Trypethelium platystomum Mont. = Pyrenula anomala.

Verrucaria aggregata Fée = Pyrenula anomala.

Verrucaria augescens Nyl. = Polyblastiopsis augescens.

Verrucaria monocarpa Kremp. = Porina monocarpa.

Verrucaria quinque-septata Nyl. = Thelidium quinqueseptatum.

Verrucaria santensis Tuck. var. murina Kremp. = Pyrenula sp.

Verrucaria tropica Ach. = Trypethelium tropicum.

Rejected records

Collema flaccidum Ach. — Vega expedition, corticola, 1879, Almquist

[Nylander, 1891: 17]. Erroneous record (Degelius, 1974), perhaps mistake

for C. leptaleum.

Parmelia dilatata Vain. — Johnson A-109 [US, not seen; Hale, 1965: 247]. This

record needs reinvestigation, it was given in a study (Hale, 1965) where

the similar Parmotrema gardneri was not recognized and most probably

it represents that species.

Parmelia perforata Ach. — Vega expedition, corticola, 1879, Almquist

[Nylander, 1891: 18]. This record needs reinvestigation to establish

its current taxonomic position, the species name has been much

misapplied.

476 Gard. Bull. Singapore 61 (2) 2010

List of collectors of Singapore lichens

Ernst ALMQUIST: Vega expedition; 1879; collections in S, H-Nyl.

André & Mariette APTROOT: Nrs. 25950-26005?, 21-31 June 1989;

collections in Herb. Aptroot.

O. BECCARI: collections in M.

H. M. BURKILL: Nrs. 2139, 2171-2180?, 2566; dd. 1959, 1960; collections in

SING.

T. R. CHIPP: Nr. 4915, 5801, 6191; dd. 1919-1920; collections in SING.

Gunnar DEGELIUS: Nrs. As 485-497?; 25-26 Mar. 1964; collections in

URS:

Paul DIEDERICH: 1994; in Hb. Diederich.

FARIDA: NUS student collaborator with H. Sipman.

HARMAND (?): collections in M.

JOHNSON: Nr. A-109; collection in US.

KIAH: Nr. 5846, 6038; 1920; collection in SING.

Aino M. LEMAITRE (“Lamaitre”): 1949; collections in H, SING.

Benjamin LEE: 2000; collections in SING.

A. C. MAINGAY: 1861-1865; collections purchased by Sir Joseph Hooker,

now in BM?, pp. in H-Nyl.?

H. MOLLER: collections in S, TUR.

D. H. MURPHY: collaborator with H. Sipman.

M. NOOR: Nr. 5663; 1920; collection in SING.

H. N. RIDLEY: Nr. A 54; 189x; collections in SING.

G. RIDORET: Nr. 5657; 1920; collection in SING.

Felix SCHUMM: 199: Hb. Schumm

Uwe SCHWARZ: collaborator with F Schumm.

Harrie J. M. SIPMAN: Nr. 45661-46412; 4-25 Nov. 2000; collections in B,

SING.

Benito C. TAN: 2000; collections in SING; collaborator with H. Sipman.

Leif TIBELL: Nrs. 8800-8872? , 13925; 16-18 Sept. 1980,5 Oct. 1983;collections

in UPS.

Discussion

The list of lichen species reported for Singapore contains 296 taxa. A

discussion of the significance of this biodiversity is given in Sipman (2009).

However, the figure is only preliminary. On one hand it contains some old

reports, which need verification and may turn out to be synonyms of other

listed species. On the other hand there are several dozens of samples are left

unidentified and may well represent another 25 species. Many are vegetatively

reproducing thalli without ascocarps to help establish their affinity. Some

A Conspectus of the Lichens (Lichenized Fungi) of Singapore 477

very characteristic and frequent types have recently been described as new

species (Sipman, 2003): Lepraria usnica, Cryptothecia granularis (ct. Table

3), but many less frequent forms without clear affinities are still awaiting

treatment. Also some of the better sites in Singapore could be rewarding

for additional investigation, e.g., the Nee Soon swamp. Moreover, at least 84

species are identified with certainty to genus level only. This suggests that

the currently documented lichen diversity of Singapore amounts to over 300

species, perhaps even 325. Remarkably 68 taxa were found only before 1985.

In view of the much more intense sampling during the author’s field work

in Singapore, it is unlikely that so many species have been overlooked. This

leaves the suggestion that many may have become extinct in the meantime

(for discussion see Sipman in prep.), so that the currently present figure may

be only 250 species.

For an impression of the distribution of the lichens within Singapore,

the species numbers per locality are compared in Table 5. It shows that the

Botanical Garden is the most important habitat for lichens in Singapore. The

importance of botanic gardens for lichen diversity is a common feature in

urbanized areas, and was observed also in, e.g., Berlin (Sipman and Aptroot,

2008). Also the ecologically rather stable and varied landscape of Pulau

Ubin appears favorable for lichens. Among the areas with primary forest

relics, the Nee Soon swamp 1s clearly better than Bukit Timah. Notably poor

in lichens are the rainforest remnant in the Botanical Garden, the Bukit

Brown Cemetery, and the abandoned quarry at Bukit Timah. Cemeteries

and abandoned quarries are often sites with high lichen diversity, but

apparently not in Singapore.

When looking at the numbers of species found only once (Table 5, third

column), the sequence changes slightly. Nee Soon Swamp comes on the first

place. This depends mainly on the many foliicolous lichens (specialized for

growing on leaves) that have been found only here. Also the NUS campus

ranks higher, probably because more time was involved in the study of its

lichen flora than elsewhere, so that more rare or inconspicuous species were

found.

Table 5. Number of lichen species and of unique finds per locality.

Singapore Botanical Garden 81 10

Pulau Ubin (island E of Singapore) 74 12

Nee Soon (nature reserve with swamp forest) 61 7]

MacRitchie Reservoir (secondary forest) Si 7

Bukit Timah (primary forest relic) Sil 4

NUS university campus 48 9

Sungei Buloh (nature reserve) 41 5

Fort Canning Park (city park) 4] 4

478 Gard. Bull. Singapore 61 (2) 2010

Lazaro Island (small island S of Singapore) 33 3)

Labrador Park (at the coast, W-side) Di 2

Sembawan Park (near the coast, E-side) 25 D,

Kusu Island (small island S of Singapore) 24 2

Lower Peirce Reservoir (secondary forest) 16 ]

Botanical Garden - Rainforest (primary forest relic) 15 2

Bukit Brown Chinese Cemetery 8 3

Bukit Timah quarry 3 1

Most lichens were observed on free standing trees, mostly in parks.

Folicolous lichens were observed mostly in Nee Soon, and less commonly

in other forest remnants. Saxicolous lichens, so common in the cold climatic

zones of the world, are very uncommon in Singapore. Only occasionally a few

thalli were found on hard, crystalline stone, on anthropogenic occurrences.

Concrete, now so common as constructing material, is usually devoid of

lichens; these were found only occasionally in half-shade situations, perhaps

depending on a certain degree of weathering. It could be speculated that

increased weathering and arrival of new immigrant species will cause an

increase in the number of saxicolous species.

Special attention was paid to the relation with phorophytes (supporting

trees). However, the high number of different tree species involved makes

any correlation insignificant. Nevertheless the information is presented in the

species list to make it available for monitoring of future changes in the lichen

growth. With this purpose also the tree numbers in the Singapore Botanical

Garden have been added. The observations confirm the great significance

of SBG for the maintenance of biological diversity in Singapore. Not only it

houses a higher number of species than any other site, for many interesting

species it appears to be the only site, like Chiodecton leptospermum, Crocynia

pyxinoides, Enterographa subserialis, and Jamesiella perlucida.

Acknowledgements

The generous support provided by Dr. B.C. Tan (at NUS then) is to be

mentioned first here. He suggested the project, kindly arranged many

details of the study visit, including the required financial resources, and

provided research facilities. P-M. Jorgensen kindly allowed publishing his

identification of Lecidopyrenopsis, D. Ertz his Opegrapha identifications

(all except O. vegae). For loan of specimens, A. Aptroot, P. Diederich, F.

Schumm and the curators of SING and UPS are gratefully acknowledged.

Prof. C.H. Murphy and Ms Farida kindly provided fieldwork support and

pleasant company. The National Parks Board and Singapore Botanic

Gardens helpfully provided the necessary permits for the fieldwork and the

A Conspectus of the Lichens (Lichenized Fungi) of Singapore 479

Raffles Museum for Biodiversity Research (RMBR) enabled the project by

giving the financial support.

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Gardens’ Bulletin Singapore 61 (2): 483-522. 2010 483

A Checklist of Merremia (Convolvulaceae) in Australasia

and the Pacific

G.W. STAPLES >

‘The Herbarium, Singapore Botanic Gardens

1 Cluny Road, Singapore 259569

Formerly at Herbarium Pacificum, Bishop Museum

Honolulu Hawai‘i 96817, U.S.A

Abstract

A checklist of Merremia species is presented for Australasia and the

Pacific. In total, 49 species are enumerated here. Total numbers of species

are estimated by region and for the genus as a whole. Comments on the

systematics and taxonomy of the genus are reported and several problematic

species complexes are identified. An index of numbered specimens examined

is included to facilitate curation and identification of herbarium specimens.

Introduction

The generic name Merremia was published as a nomen nudum by Dennstedt

(1818: 12, 23, 34) and several modern floras attribute the valid publication

of the name to Hallier (1893). Validation was provided substantially earlier,

however, by Endlicher (1841) and both Index Nominum Genericorum and

the current International Code of Botanical Nomenclature note this fact,

as well as the status of nomen conservandum for Merremia. The genus has

been recognized at times, and merged into /pomoea at other times, ever

since the concept was first established. Following the publication of Hallier’s

(1893) classification for the family, in which he noted a pollen difference

between /pomoea (spinulose pollen grains) and Merremia (non-spinulose

grains) that was given considerable importance in recognizing subfamilial

and tribal groupings, the genus Merremia gradually gained wider acceptance

and by the mid-twentieth century was almost universally accepted in floras

world-wide.

Recent molecular analysis (Stefanovic ef al., 2002, 2003) strongly

indicates that Merremia is a polyphyletic genus, a fact that was already

suggested by a comprehensive survey of pollen morphology (Ferguson er

al., 1977). In a family where one, or at most two, pollen types are typical for

a genus, the five pollen types documented for Merremia are indicative that

the genus is a mixed assemblage of species.

484 Gard. Bull. Singapore 61 (2) 2010

Merremia has long proven to be difficult to characterize as a genus,

with no single morphological character that can be used to recognize it.

O’Donell (1941) proposed infrageneric groupings based on his revision

of the American species; these sections were adopted by Van Ooststroom

and Hoogland (1953) for Malesia, with the addition of one new section,

Wavula. Later, two species with non-spinose, pantoporate pollen grains were

segregated from Merremia as the new genus Xenostegia (Austin & Staples,

1980); although this genus has not been universally accepted, particularly

in African floras, the molecular data supports the segregation and indicates

that Xenostegia is sister group to the genus Hewittia (Stefanovic et al., 2002).

A thorough revisionary study is needed to elucidate just how many genera

ought to be recognized from the polyphyletic Merremia and its closest allies

(Operculina, Hewittia, Remirema, Decalobanthus) once their relationships

are better understood. Until that is possible, an enumeration of the Asian

species heretofore included in Merremia will provide a useful reference as

well as a starting point for further study. And to put the Asian species in

context, the species recognized elsewhere in the world must be considered.

The number of species of Merremia has been variously estimated in

the botanical literature at ca 70 (Mabberley, 1997; Staples and Brummitt,

2007) or ca 80 (Van Ooststroom and Hoogland, 1953; Verdcourt, 1963; Airy

Shaw, 1980; Fang and Staples, 1995; Deroin, 2001; Meeuse and Welman, 2000;

inter alia). However, these estimates have always seemed too low and it was

decided to tabulate the species world-wide to see how many are currently

recognized. Tabulation of the accepted names used in recent literature (see

Appendix for list of literature consulted) via an Excel® spreadsheet allowed

for geographic distribution to be compared and synonyms to be reconciled.

Once redundancy in the list was eliminated, there are 27 unique species in

the Americas, 31 species in Africa (including Madagascar), just 3 species

in Australia, and 40 species in Asia/Malesia/Pacific. This brings the actual

number of species to just over 100, which reveals there has been a sizable

underestimate (by at least 20%) of the species richness in Merremia.

Checklist of Asian Merremia Species

The Asian and Pacific species are enumerated here — 49 in total — in

alphabetical order by species name. Only selected synonyms relevant for

the Asia/Malesia/Pacific regions are included. Excluded names and selected

synonyms are listed at the end, to assist in cross-referencing to accepted

names.

It has not been possible to produce a key for identification of all Asian

species at the present level of knowledge, largely because several species

are known only from the type gathering, or from few collections, that are

A Checklist of Merremia (Convolvulaceae) in Australasia and the Pacific 485

not available for study. Problem groups have been discussed in the Notes

sections; two principal complexes of intergrading taxa bear mentioning.

1. The Bornean endemics comprising MM. crassifolia/elmeri/gracilis/

korthalsiana

Van Ooststroom (1939), and later Van Ooststroom and Hoogland

(1953), were able to clearly distinguish several species endemic to Borneo,

based on the small number of collections available for study at that time. In

the intervening decades, many more collections have been made, and the

distinctions between species do not seem nearly so clear cut now. Certainly

there are specimens in herbaria that are intermediate in their characters,

and hence can’t be identified precisely, though they do seem close to one

or another of the species in this complex. Revisionary study is needed to

elucidate the true relationships in this group.

2. The M. bambusetorum/kingii/tonkinensis/umbellata group

O’Donell (1941), in tropical America, and Van Ooststroom (1939), in

Malesia, were able to sharply delineate M. umbellata from all other species

of the genus that they studied; based on several characters they assigned M.

umbellata to sect. Xanthips. However, on the continental Asian mainland,

there are a number of additional species, all clearly referable to sect.

Xanthips, and some of these named entities intergrade with M. umbellata.

To date, attempts to resolve the intergradation at the floristic level have

met with mixed success (Songkhla and Khunwasi, 1993); what is needed is

a regional study of the entire complex, with careful comparison of all type

specimens. For now, some observations and comments have been made in

the Notes section to guide development of such a revisionary study.

Merremia Dennst. ex Endl., Gen. PI. 1: 1403. 1841, nom. cons.

1. Merremia aegyptia (L.) Urban, Symb. Antillanae 4: 505. 1910.

Ipomoea aegyptia L., Sp. Pl. 162. 1753. — Type: Herb. Linn. 218.35 (LINN);

lectotype chosen by D.F. Austin, Fl. Ecuador 15: 84. 1982.

Syn.: Convolvulus pentaphyllus L.,nom. illeg.:1. pentaphylla Jacq.;Spiranthera

pentaphylla (Jacq.) Bojer; Batatas pentaphylla (Jacq.) Choisy; Merremia

pentaphylla (Jacq.) Hallier f.; C: nemorosus Roem. & Schultes; C. hirsutus

Roxb., nom. illeg.; C. munitus Wall. ex Wight.

Distribution: Pakistan, India, Vietnam, Australia, Hawaiian Islands (Kaua‘1,

O‘ahu, Moloka‘i, Lana‘i, Maui, Kaho*olawe, Hawai‘i). Native to tropical

America and now widespread around the world as a weed.

486 Gard. Bull. Singapore 61 (2) 2010

Because Convolvulus pentaphyllus L. is an illegitimate name the first

time the epithet pentaphyllus is taken up in another genus, in this case by

Jacquin (1788), provides the first valid publication for it and all subsequent

combinations of the epithet pentaphyllus are to be based on Ipomea

pentaphylla Jacq.

Notes: Fosberg & Sachet (1977) reported this species from Micronesia based

on a single collection, D. Anderson 227, with duplicates in several herbaria.

However, the BISH duplicate of Anderson 227 is Ipomoea pes-tigridis L.

I have seen no authentic material of M. aegyptia from the Pacific Basin

aside from the Hawaiian Islands, where it is well established (Wagner et al.,

1999).

2. Merremia aniseiifolia Ooststr., Blumea 12: 363. 1964.

Type: Indonesia, Irian Jaya, Kebar Valley, Andjai, VW. Moll BW9511

(holotype, L!; iso LAE, n.v.).

Distribution: New Guinea.

Notes: Known only from the type collection until quite recently, when a single

additional collection was made. When Van Ooststroom described this species

he commented that it resembles M. tridentata, a species now removed to the

genus Xenostegia (see Excluded Species). It would be worthwhile examining

the pollen of M. aniseiifolia to determine whether the resemblance is purely

superficial, or whether this might be a third species of Xenostegia. It is a pity

that most of the duplicates of the following collection have no flowers; only

the Kew sheet had a single flower

Specimen examined: INDONESIA. Irian Jaya: NE Kepala Burung, between

Sungai Apriri and Sungai Aremi, 11 May 1994, M.J. Sands et al. 6544 (BISH!,

K!, KEP!, SING!).

3. Merremia bambusetorum Kerr, Bull. Misc. Inform. 1941: 18. 1941.

Type: Thailand [SE], Trat, Khao Kuap, Kerr 17704 (holotype, K!; isotype,

BK!).

Distribution: Myanmar, Thailand, Vietnam.

Notes: Distinguishing M. bambusetorum from M. kingii and some forms

of M. umbellata is quite difficult. It is possible that the holotype of M.

bambusetorum is conspecific with one or more of the syntypes for M.

kingii, which may be a mixed concept. Only careful comparison of all type

A Checklist of Merremia (Convolvulaceae) in Australasia and the Pacific 487

specimens concerned will resolve the ambiguity in species concepts. It is

possible that M. caloxantha belongs with this complex as well.

That said, this complex of three intergrading entities can immediately

be recognized among Asian Merremia by the presence at the petiole base of

tiny paired auricles that look very much like stipules. These can be hidden in

the typically dense hair-cloth of M. umbellata, but are usually quite evident

for M. bambusetorum and M. kingii.

4. Merremia bimbim (Gagnep.) Ooststr., Blumea 3: 343. 1939.

Ipomoea bimbim Gagnep., Not. Syst. (Paris) 3: 140. 1915. — Type: Vietnam,

Hanoi Prov., near Vo-xa, Bon 2700 (holotype, P, n.v.).

Distribution: Vietnam.

Notes: I have not seen the type specimen for this name nor any other

collections of it. The species appears to be distinct and remains enigmatic.

5. Merremia boisiana (Gagnep.) Ooststr., Blumea 3: 343. 1939.

Ipomoea boisiana Gagnep., Notul. Syst. (Paris) 3: 141.1915.—Types: Vietnam,

montagnes de Cai-kinh, Bois 138 (syntype, P!); mont Cha-pa, Lecomte &

Finet s.n. (syntype, P!). Laos, vers Ken-trap, Spire 1049 (syntype, P!): du

Mékong a Hue, Harmand 1827 (syntype, P!). See notes.

Syn.: 1. boisiana var. fulvopilosa Gagnep., I. boisiana var. rufopilosa Gagnep..,

Merremia boisiana (Gagnep.) Ooststr. var. rufopilosa (Gagnep.) C.Y. Wu.

Distribution: China, Laos, Vietnam, Indonesia (Sumatra).

Notes. Floras for Malesia recognize two varieties: var. boisiana and var.

sumatrana Ooststr., and floras for China and SE Asia recognize var. boisiana

and var. fulvopilosa (Gagnep.) Ooststr. See Van Ooststroom and Hoogland

(1953: 450) for a key to the Malesian varieties and Fang and Staples (1995:

299) for a key to the Chinese varieties.

In transferring the epithet to Merremia, Van Ooststroom (1939b:

344) excluded a fifth syntype cited by Gagnepain in the protologue, Beccari

3594 from Borneo, and referred it to his new species Merremia crassinervia

Ooststr. Chinese specimens differ from Malesian ones in several characters

and it is possible that they are a different species. No taxonomic changes are

initiated here until revisionary study clarifies relationships.

6. Merremia borneensis Merr., Univ. Calif. Publ. Bot. 15: 260. 1929.

Type: Malaysia, Sabah, Elphinstone province, Tawao, A.D.E. Elmer 20990

(isotypes, A!, BO, F!, G!, GH!,K, MO(x 2)!, PR, NY!, SING!, U).

488 Gard. Bull. Singapore 61 (2) 2010

Distribution: Malaysia (Peninsula, Sabah, Sarawak), Brunei, Indonesia

(Kalimantan).

Notes: In herbaria, several other Bornean species are often confused with,

and misidentified as, M. borneensis. The basal petiole attachment, medium-

large flowers, and puckered, rugose leaf surface in dried material makes it

distinctive.

7. Merremia bracteata P.S. Bacon, Bot. J. Linn. Soc. 84: 259. 1982.

Type: Solomon Islands, Kolombangara, near Poitete forestry camp, PS.

Bacon 2 (holotype, K!; isotypes, BSIP, L!).

Distribution: Solomon Islands. Known from only two collections; apparently

endemic.

Note: The general facies of this species comes very near to Operculina

ventricosa, known from the Micronesian islands and the Caribbean (Staples,

2007). The similarity may be only superficial but requires investigation. A

few South Pacific insular collections seen (New Caledonia, Vanuatu [Santo,

Vaté]) appear intermediate between these two species bearing large, foliose,

persistent inflorescence bracts.

Specimens examined: SOLOMON ISLANDS (Santa Cruz group). Vanikoro:

secondary forest near Government House, Peou, 27 Mar 1963, 7:S. Whitmore

BSIP lors (1 ').

8. Merremia caloxantha (Diels) Staples & R.C. Fang, Novon 5: 109. 1995.

Ipomoea caloxantha Diels, Notes Roy. Bot. Gard. Edinburgh 5: 203. 1912.

— Type: China, Yunnan: in valley of Mekong River, at crossing of Teng Yueh-

Talifu road, G. Forrest 1111 (holotype, E!).

Distribution: China, Laos, Vietnam.

Notes: An enigmatic species known, for most of the twentieth century, only

from the type collection. A recent specimen from Laos has been tentatively

identified as conspecific. Although the corollas on these SE Asian specimens

are smaller than those on the holotype sheet, in all other characters (leaf

shape and indumentum, calyx characters, corolla pubescence), they are a

good match with the type specimen. The following collections document the

distribution for M. caloxantha.

A Checks: of Merremiz (Commolvelacese) in Amsralasic and the Pacific 489

Specimens examined: CHINA. Yunnan: Nujiang Lisu Auton. Prefect.. Lushui

Xian, road from Man Hui village to Shanjiang. 3 Oct 1997. Gaoli Gongshan

Exped. 1997-9928 (E!): Szemao [=Simao]. A. Henry 1344] (E!. K!): valley

of Mekong at the crossing of Teng Yueh—Dalifu road. Sep 1905. G. Forrest

71111 (E!). PDR LAO. Khammouan Province: along tracks S of Ban Mak

Phueang. 31 Oct 2005, M_E Newman et al. LAO-659 (E!). VIETNAM. Ho

Yung Shan and vicinity. Tien-yen_ 13 Oct-22 Nov 1940. W_-T. Tsang 30645

(Bt BO! Ct El L!).

Type: Fin. Puna, Jun 1860. Seemann 324 (holotype, K!: isotype. BM!).

Distribution: Fiji, apparently endemic.

Notes: As suggested previously (Staples, 2009) M. calyculata may prove to

be no more than a variant of the widespread and variable M. pacifica. One

stenle specimen, for which the vernacular name. “wabula”. was recorded

by the collector. may belong here. Van Ooststroom (1939a: 265) recorded

wavula as the vernacular name for M. calyculata and veliyawa for M. pacifica.

It remains to be seen whether the difference in vernacular names reflects a

genuine taxonomic distinction. Further collecting and field study of Fijian

plants is needed to resolve whether one species or two is present.

Specimen examined: FIJ1. Bequ: Rukua village. 21 July 1972. M. Weiner MW-

72-F-124 (BISH!).

Convolvulus cissoides Lam.. Tab. Ency. 1: 462. 1793. — Ipomoea cissoides

(Lam.) Gniseb. — Type: French Guiana, Cayenne, Leblond sn. (holotype.

P-LA. n_v.: IDC microfiche 6207. 462: I. 4!).

Distribution: India (West Bengal). Sn Lanka, Papua New Guinea, and

Thailand.

Note: Native to tropical America and increasingly naturalized in the Old

World tropics.

11. Merremia clemensiana Ooststr.. Blumea 3: 350. 1939.

Type: Malaysia. Sarawak, Kapit. upper Rejang River. J. & M.S Clemens

21133 (holotype, BO!: isotypes. A!. B!, BISH!, MO!. NY(x2)!. P!. SAR!).

490 Gard. Bull. Singapore 61 (2) 2010

Distribution: Malaysia (Sarawak, Sabah?).

Note: Merremia clemensiana is expected to be more widespread on Borneo

than the few available collections would indicate. See the comments under

M. gracilis.

12. Merremia cordata C.Y. Wu & R.C. Fang, Fl. Reipubl. Popularis Sin.

64(1): 163. 1979.

Type: China, Yunnan: Likiang, Cheli, R.C. Ching 21660 (holotype, KUN, n.v.;

iso A!).

Distribution: China (Sichuan, Yunnan).

Notes: The following collections have been examined and agree well with

the description for the species.

Specimens examined: China. Yunnan: Ta-pin-tze, 2 Sep 1887, J.M. Delavay

3073 (P(x3)!); Yung Ning, May 1933, McLaren’s collector 204 (K); sine loco

or date, G. Forrest HI2T (KK):

13. Merremia crassinervia Ooststr., Blumea 3: 350. 1939.

Type: Malaysia, Sarawak, Saribas, Paku, Haviland & Hose 3523E (holotype,

L!),3523K (isotype, K!), 3523A (isotype, SAR!).

Distribution: Malaysia (Sarawak), Indonesia (Kalimantan).

Notes: Van Ooststroom described this species from material that lacked

mature flowers, thus the shape of the corolla remains imperfectly known.

The inflorescence architecture and leaf venation characters he noted seem

distinct but further comparative study is needed to correctly associate all

stages in the life history.

14. Merremia davenportii (F. Muell.) Hallier f., Bot. Jahrb. Syst. 16: 552.

1893.

Ipomoea davenportii F. Muell., Fragm. Phytogr. Austral. 6. 1868. — Type:

Australia, [Northern Territory] hills of Davenport’s Range, J.M. Stuart s.n.

(holotype, MEL?, n.v.).

Distribution: Australia (Western Australia, Northern Territory).

Specimen examined: AUSTRALIA. Western Australia: near Hedland,

Pippingarra Station, 20 Jun 1962, R.D. Royce 7492 (PERTH!).

A Checklist of Merremia (Convolvulaceae) in Australasia and the Pacific 49]

15. Merremia dichotoma Ooststr., Blumea 3: 311. 1939.

Type: Indonesia, south middle Timor, Kolbano, S coast, M.E. Walsh 375

(holotype, BO!; isotype, BO (x2)!, L!).

Distribution: Malesia (Timor).

Notes: Apparently rather rare; just two recent collections have come to light

since the type gathering was made. The species bears an uncanny similarity

to M. hainanensis (q.V.).

Specimens examined: INDONESIA. Timor: Berg Sanbet, 21 Apr 1971, CW.

Kooy 794 (L!), Luid Belu, 28 Jun 1970, C.W. Kooy 832 (L!).

16a. Merremia dissecta (Jacq.) Hallier f. var. dissecta, Bot. Jahrb. Syst. 16:

592.1893.

Convolvulus dissectus Jacq., Obs. Bot. 2: 4. tab. 28. 1767; Ipomoea dissecta

(Jacq.) Pers.; Operculina dissecta (Jacq.) House. — Type: Jacquin plate #28

in protologue (lectotype, designated by Austin, Florida Scientist 42: 219.

1979).

Syn. /. sinuata Ortega

Distribution: Pakistan, India, Sri Lanka, Myanmar (reported), China,

Indonesia (cultivated Java, Sumatra), Philippines (Mindanao), Australia

(Northern Territory, Queensland, Western Australia), French Polynesia

(Tahiti), Papua New Guinea, Solomon Islands (Guadalcanal), Samoa

(Upolu), Tonga (Niuafo’ou), Seychelles (Praslin); native in tropical and

subtropical America and now virtually ubiquitous as a tropical weed.

16b. Merremia dissecta var. edentata (Meisn.) O’Donell, Lilloa 6: 502.

1941.

Ipomoea sinuata Ortega var. edentata Meisn. in Martius, Fl. Brasil. 7: 285.

1869. — Type: Brazil. Bahia: prope Ilheos, Riedel 131] (isosyntype NY!); Rio

de Janeiro: prope Copacabana, Riedel s.n. (n.v.).

Syn. M. fulva (Bertol.) Manitz

Distribution: Brazil, Bolivia, Paraguay, Argentina; naturalized in French

Polynesia (Society Islands: Mo‘orea).

Notes: There is no consensus about the rank at which this taxon should be

recognized. Manitz (1983: 180) has raised it to specific rank and taken up the

name M. fulva for it but without description or discussion of the characters

used to justify this action. Austin & Staples (unpublished data) consider it

492 Gard. Bull. Singapore 61 (2) 2010

to be a variety of the widespread and variable M. dissecta, and that rank

is accepted here. It has been treated this way in a recent checklist for the

Argentine flora (Zuloaga and Morrone 1999: 564).

How this temperate South American variety of a widespread species

came to be naturalized on a small Pacific island is unknown (Staples, 2009).

The earliest Polynesian collection dates from 1931 (M.L. Grant 5359). Plants

have been collected at various sites around Mo‘orea ever since, the most

recent dating from 1999 (Murdoch M014).

17. Merremia eberhardtii (Gagnep.) T.N. Nguyen in Averyanov et al., Mater.

Fl. Rast. Ostrov. Vetnama 43. 1988.

Ipomoea eberhardtii Gagnep., Not. Syst. (Paris) 3: 145. 1915. — Type:

Vietnam, Thua-thien province, near Long-co, Eberhardt 1708 (isotypes P,

n.v., photos!)

Distribution: Vietnam. Endemic and very seldom collected, apparently. Only

one collection seen aside from the type.

Specimen examined: Vietnam. Hue and vicinity, 1927,J. & M.S. Clemens 4091

CAYBIMEINGIOKY Pl IN),

18. Merremia elmeri Merr., Univ. Calif. Publ. Bot. 15: 261. 1929.

Type: Malaysia, Sabah: Elphinstone province, Tawao, A.D.E. Elmer 20396

(isotypes, B!, BO!, G!, L!, MO!, NY!, SING!).

Distribution: Malaysia (Sabah), Indonesia (Kalimantan).

Notes: Two varieties are recognized — var. elmeri and var. glaberrima Ooststr.,

both found on Borneo. See Van Ooststroom and Hoogland (1953: 453) for

distinguishing features of the varieties.

19. Merremia emarginata (Burm. f.) Hallier f., Bot. Jahrb. Syst. 16: 552.

1893.

Evolvulus emarginatus Burm. f., Fl. Ind. 77. 1768. — Type: Indonesia, Java,

Kleinhof 85 (lectotype, G-Burman!, chosen by Staples and Jacquemoud,

Candollea 60: 449. 2005).

Syn.: Convolvulus excisus Zipp.; C. reniformis Roxb.; Ipomoea reniformis

(Roxb.) Choisy; Lepistemon reniformis (Roxb.) Hassk.

Distribution: Nepal, India, Sri Lanka, Bangladesh, Myanmar, China, Thailand,

Malaysia, Indonesia (Java), Philippines; also in Africa.

A Checklist of Merremia (Convolvulaceae) in Australasia and the Pacific 493

20. Merremia gemella (Burm. f.) Hallier f., Bot. Jahrb. Syst. 16: 552. 1893.

Convolvulus gemellus Burm. f., Fl. Ind. 46. 1768.—/. gemella (Burm. f.) Roth.

— Type: Indonesia, Java, collector unknown (lectotype, G-Burman!, chosen

by Staples and Jacquemoud, Candollea 60: 448. 2005).

Syn.: Ipomoea cymosa var. radicans (Blume) Miq.; I. polyantha Miq.; I.

radicans Bume non Choisy.

Distribution: India, Sri Lanka, Myanmar, Thailand, Vietnam, Taiwan,

Malaysia, Indonesia (Java, Seram, Irian Jaya), Philippines, Papua New

Guinea, Australia, Micronesia (Mariana Islands: Guam).

Notes: Malesian floras recognize two varieties — var. gemella is widespread

throughout Malesia, Asia, and the Pacific; and var. splendens Ooststr.

is confined to a small area in Papua New Guinea. See Van Ooststroom

and Hoogland (1953: 441) for a key to the Malesian varieties. An earlier

lectotypification for this species (Van Ooststroom, 1939) had to be set aside

because the specimen chosen was not part of the original material available

to Burman: see Staples and Jacquemoud (2005) for details.

21. Merremia gracilis Campbell & Argent, Notes Roy. Bot. Gard. Edinburgh

45(2): 345-348. 1988.

Type: Malaysia, Sabah: Lahad Datu distr., Ulu Segama, Danum Valley Field

Centre, Argent & Campbell 411854 (holotype, SAN!; isotypes, A!, E!, K!,

L!)

Distribution: Malaysia (Sabah), Indonesia (Kalimantan).

Notes: Merremia gracilis was described and compared by its authors with the

species assigned to sect. Wavula (e.g., M. similis, M. pacifica, M. calyculata);

on that basis it seemed distinctive. However, my own examination of the

type gathering revealed it is much more similar to species of sect. Hailale

(e.g., M. clemensiana, M. crassinervia, M. korthalsiana). It appears that the

name M. gracilis has been applied to plants from Sabah, where they are

widespread and abundant, whereas plants from Sarawak and Brunei have

been otherwise named. Revisionary study may show that these names refer

to one, or more, variable species distributed across the whole of Borneo.

22. Merremia hainanensis H. S. Kiu, Fl. Hainan 3: 587. 1974.

Type. China. Hainan Province: Wanning County, H.F How 71693 (holotype,

where?, n.v.;iso A!).

494 Gard. Bull. Singapore 61 (2) 2010

Distribution: China (Hainan), Vietnam.

Notes: The protologue does not state where the holotype specimen is

deposited and I have failed to locate it. Kiu Hua-shing compared this new

species from Hainan to M. dichotoma and the excellent figure accompanying

the protologue demonstrates that the two are uncannily similar. When I

saw a duplicate of How 71693 in A, I first called it M. dichotoma without

hesitation, not knowing that it was an isotype for Kiu’s species. Ultimately,

these two species may prove to be conspecific, though their type localities

are far apart. A single specimen was discovered in the Paris herbarium that

indicates M. hainanensis also occurs in nearby Vietnam.

Specimen examined: Vietnam. Can Tho, route de Can Tho a Long xuyén, 2

Nov 1967, Vu Van Cuong 690 (P!).

23. Merremia hederacea (Burm. f.) Hallier f., Bot. Jahrb. Syst. 18: 118. 1893.

Evolvulus hederaceus Burm. f., Fl. Ind. 77. 1768. — Type: Indonesia, Java,

Pryon s.n. (lectotype, G-Burman!, chosen by Van Ooststroom, Blumea 3:

306. 1939).

Syn.: Convolvulus acetosellifolius Desr.; C. chryseides (Ker Gawl.) Spreng.; C.

dentatus Vahl; C. flavus Willd.; C. lapathifolius Spreng.; [pomoea acetosellifolia

(Desr.) Choisy; /. chryseides Ker Gawl.; I. dentata (Vahl) Roem. & Schultes;

I. subtriflora Zoll. & Moritzi; Lepistemon glaber Hand.-Mazz.; L. muricatum

Spanoghe; Merremia chryseides (Ker Gawl.) Hallier f.; M. convolvulacea

Dennst. ex Hallier f.

Distribution: Pakistan, Nepal, India, Sri Lanka, Bangladesh, Myanmar,

Thailand, China, Taiwan, Japan (Ryukyu, Ogasawara), Laos, Cambodia,

Vietnam, Malaysia (Sarawak, Sabah), Singapore, Indonesia (Sumatra,

Kalimantan), Philippines, Australia (Northern Territory, Queensland,

Western Australia), Papua New Guinea, Micronesia (Caroline Islands:

Yap; Mariana Islands: Guam), French Polynesia (Society Islands: Tahiti);

and widespread in tropical Africa, Madagascar, Indian Ocean islands

(Réunion, Mauritius); introduced in the Americas and naturalized around

the Caribbean (Cuba, Lesser Antilles, Colombia, Trinidad and Tobago).

24. Merremia hirta (L.) Merr., Philipp. J. Sci. 7: 244. 1912.

Convolvulus hirtus L., Sp. Pl. 1: 159. 1753. — Type: India, Osbeck 11 in Herb.

Linn. 218.56 (lectotype, LINN!, chosen by Merrill, Philipp. J. Sci., sect. C, 7:

245.1912).

Syn.: C. caespitosus Roxb.; C. reptans L., p.p.; ? Ipomoea hepaticaefolia

Blanco; /. linifolia B1L.; 1. philippinensis Choisy; Lepistemon decurrens Hand.-

A Checklist of Merremia (Convolvulaceae) in Australasia and the Pacific 495

Mazz.: Merremia caespitosa (Roxb.) Hallier f.: M. decurrens (Hand.-Mazz.)

H. S. Kiu: Skinneria caespitosa (Roxb.) Choisy.

Distribution: India, Bangladesh, Myanmar, Thailand, China, Taiwan,

Laos, Vietnam, Malaysia, Singapore, Indonesia (Sumatra, Irian Jaya),

Philippines, Australia (Queensland), Papua New Guinea, Solomon Islands

(Guadalcanal).

Notes: Two varieties are recognized — var. hirta is widespread: var. retusa

Ooststr. is found in the Philippines (Luzon). See Van Ooststroom and

Hoogland (1953: 442) for a key to the Malesian varieties.

25. Merremia hungaiensis (Lingelsh. & Borza) R.C. Fang, Fl. Reipubl.

Popularis Sin. 64(1): 76. 1979.

Ipomoea hungaiensis Lingelsh. & Borza, Repert Spec. Nov. Regni Veg. 13:

389. 1914. — Type: China. Yunnan: Talifu [Tali town], bei Tschian-t’ou-shao

vor Hun-gai, 19 Aug 1913, H.W. Limpricht 928 (holotype, WRSL, n.v.).

Syn. Ipomoea wilsonii Gagnep. (1915), nom. illeg., non House (1907):

Merremia wilsonii Verdc.; I. hungaiensis vat. linifolia C.C. Huang.

Distribution: China (Guizhou, Yunnan).

Notes: Chinese floras recognize two varieties — var. hungaiensis and var.

linifolia (C.C. Huang) R.C. Fang. See Fang and Staples (1995: 297) for a key

to the varieties.

26. Merremia incisa (R. Br.) Hallier f., Meded. Rijks Herb. Leiden 1: 21.

1923 [t.p. 1910].

Ipomoea incisa R. Br., Prodr. 486. 1810.—Type: Australia, locality not stated,

R. Brown s.n. (holotype, BM, n.v.).

Distribution: Australia. Endemic.

Notes. A presumed new species of Merremia reported from the Northern

Territory of Australia (Elliot and Jones 1993: 389) is actually M. incisa (B.

Johnson, pers. comm.,4 Feb 2009).

27. Merremia kingii (Prain) Kerr, Fl. Siam. En. 3(2):5. 1954.

Ipomoea kingii Prain, J. As. Soc. Bengal 1, 13, pt. 2: 110. 1894. — Types:

specimens from India, Bhutan, and Burma were cited (all CAL, n.v.).

Syn.: Ipomoea cymosa Roem. & Schultes var. macra C.B. Clarke: M.

umbellata subsp. macra (C.B. Clarke) P.J. Parmar.

496 Gard. Bull. Singapore 61 (2) 2010

Distribution: Bhutan, India [West Bengal (Sikkim), Assam, Arunachal

Pradesh, Nagaland, Meghalaya], Myanmar, Thailand.

Notes: This is one of the most difficult species to define among all Asian

Merremia. Prain based his new species /. kingii on at least 14 collections,

all in CAL, which would be syntypes under today’s rules of nomenclature.

He sent duplicates for some of these collections to Kew, where O. Stapf

compared them with specimens cited by Clarke in the Flora of British India

Convolvulaceae account. Based on Stapf’s findings, Prain subsumed Clarke’s

variety, /. cymosa var. macra, and included Clarke’s syntypes for that name

in his list of collections cited under /. kingii, making these collections types

for both names.

The lack of a clear-cut type specimen for this name makes it impossible

to establish a species concept. Specimens called M. kingii in herbaria

intergrade with both M. bambusetorum and M. umbellata, among others.

Careful study of the full series of Prain’s material in Calcutta, and choice of

a lectotype from the many specimens he included, will determine how this

name should be applied. A mini revision of the species complex including

kingit/bambusetorum/umbellata and others is much needed.

28. Merremia korthalsiana Ooststr., Kew Bull. 1938: 175. 1938.

Type: Indonesia. Borneo: [Kalimantan], Doesoen, Korthals 237 (holotype

L!, 2 sheets).

Distribution: Malaysia (Sarawak, Sabah), Indonesia (Kalimantan), Brunet.

29. Merremia mammosa (Lour.) Hallier f., Teysmannia 7: 164. 1897.

Convolvulus mammosus Lour., Fl. Cochin. 108. 1790; I. mammosa (Lour.)

Choisy. —Type: No specimen traced.

Syn.: Ipomoea gomezii C.B. Clarke, in part, as to flowers.

Distribution: India (West Bengal, Assam, Andaman Islands), Myanmar,

Thailand, Vietnam, Indonesia (cultivated and escaped — Java, Ball,

Moluccas).

Notes: The application of this name has stabilised since Van Ooststroom

(1939b, and in Van Ooststroom & Hoogland 1953) explicated the species

concept and applied it to a distinctive Malesian species with large flowers,

perhaps the biggest in the genus, and fascicled, swollen roots used as food

and medicine. Previously, Merrill (1935) pointed out that there is no type

specimen extant for this Loureiro name and the Rumphius plate (Herbar.

Amboin. 5: 370, pl. 131. 1750) — cited by Loureiro as a synonym — may be a

A Checklist of Merremia (Convolvulaceae) in Australasia and the Pacific 497

mixed concept. Merrill initially (1917: 442) stated that this plate depicts the

roots of a Dioscorea with the above-ground parts of some convolvulaceous

plant and later (Merrill 1935: 325) decided that the above-ground parts

referred to the sweet potato. Merrill accordingly reduced C. mammosa Lour.

to synonymy with /pomoea batatas. However, it seems that the Rumphius

plate agrees very well with the plant from SE Asia that has latterly been

called Merremia mammosa.

Clarke (1883:211) created a mixed concept when he named /. gomezii:

the flowers, based on a Burmese collection, W. Gomez s.n. (K!, 2 sheets),

are M. mammosa; the fruits, based on Kurz specimens from the Andaman

Islands, appear to belong to Operculina riedeliana.

30. Merremia pacifica Ooststr., Blumea 3: 263. 1939.

Type: Fiji. Vanua Levu: Mbua, A.C. Smith 1690 (holotype, L!; isotypes, BISH!,

MUNY!, P!).

Distribution: Fiji (Naitasiri, Ovalau, Vanua Levu), Papua New Guinea

(Bougainville, New Britain, Rossel), Solomon Islands (Guadalcanal,

Vanikoro, Kolombangara, New Georgia, Buka, Faro, Savo, San Cristoval),

Vanuatu (Maewo), Micronesia (Caroline Islands — Ponape).

Notes: A full description, illustration, and greatly expanded Pacific

distribution for this species appears in Staples (2009).

31. Merremia palmata Hallier f., Bot. Jahrb. Syst. 18: 112. 1893.

Type: Namibia. Ameib, Belck 52 (lectotype, BRA, n.v.; iso, JE, n.v.).

Distribution: Pakistan; otherwise known from tropical and southern Africa.

Notes: Austin & Ghazanfar (1979: 56) tentatively took up this name for two

collections from Pakistan that B. Verdcourt identified as M. palmata. 1 have

seen no material.

32. Merremia peltata (L.) Merr., Interpr. Rumphius Herb. Amboin. 441.

2917.

Convolvulus peltatus L.,Sp. P1.2:1194. 1753;. peltata (L.) Choisy; Operculina

peltata (L.) Hallier f. — Type: Icon in: Rumphius, Herb. Amboin. 6: pl. 159.

1750 [lectotype, chosen by Merrill, op. cit. 31, 441.1917].

Syn.: ?Chironia capsularis Blanco; ?C. lanosanthera Blanco; ?I[pomoea

menispermacea Domin; I. nymphaeifolia Blume; Merremia nymphaeifolia

(Blume) Hallier f., nom. illeg.

498 Gard. Bull. Singapore 61 (2) 2010

Distribution: India (Andaman Islands), Thailand, Malaysia (Peninsula,

Sarawak, Sabah), Singapore, Brunei, Indonesia (Aru arch., Java, Kalimantan,

Key Islands, Selebes, Sulawesi), Philippines (Mindanao, Sulu arch.), Papua

New Guinea, Australia (Queensland), Solomon Islands (Guadalcanal, San

Cristobal), Vanuatu (Banks group, Efate, Erromanga, Santo), Fiji (Bua,

Kandavu, Naitasiri, Vanua Balavu, Vanua Levu, Viti Levu), Wallis & Futuna,

New Caledonia, Cook Islands (Rarotonga), French Polynesia (Austral

Islands — Raivavae, Rurutu; Society Islands — Huahine, Maupiti, Moorea,

Raiatea, Tahaa, Tahiti), Niue, Samoa (Sava‘i, Tutuila, Upolu), Tonga (Eua,

Tafahi, Vavau), Federated States of Micronesia (Kosrae, Ponape), Belau

(=Palau); also in tropical east Africa, Madagascar, and throughout the

Indian Ocean islands.

Notes: Merremia peltata is immediately recognizable for several features: the

lianoid habit with large flowers; broadly campanulate corollas either bright

golden yellow or white; the peltate leaf attachment (though small leaves just

below an inflorescence may be basally attached); and the anthers with large

tufts of hairs. The vines are rampant growers that quickly take advantage of

any disturbance such as forest cutting or land clearing for agriculture; the

species is now considered invasive on several Pacific islands, despite being

native in Oceania. The only specimen seen from Belau states that it is an

introduced plant, grown at the Aimireek Experiment Station.

33. Merremia poranoides (C.B. Clarke) Hallier f., Bull. Herb. Boiss. 5: 375.

1897.

Ipomoea poranoides C.B. Clarke in Hook., Fl. Brit. India 4: 208. 1883.—Type:

India. Sikkim: Darjeeling, CB. Clarke 9189A (lectotype, K!, designated by

Staples, Edinb. J. Bot. 60: 91. 2006).

Syn.: Ipomoea courchetii Gagnep.; I. longipedunculata C.Y. Wu (1965), non

Hemsl. (1882); Merremia longipedunculata R.C. Fang.

Distribution: ?Bhutan, India (West Bengal, Nagaland), China (Guizhou,

Yunnan), Thailand, Vietnam.

Notes. The synonymy and typification for M. poranoides were clarified by

Staples (2006); soon thereafter it was documented in Thailand (Staples and

Traiperm, 2008). The species enjoys a wide geographic range but is evidently

not common anywhere.

34. Merremia pulchra Ooststr., Blumea 3: 348. 1939.

Type: Brunei Darussalam. Brunei Muara District: Muara, Brunei Bay, Apr

1896, G. K. Gns 14 (holotype SING!). |

A Checklist of Merremia (Convolvulaceae) in Australasia and the Pacific 499

Distribution: Brunei, Malaysia (Sarawak), ?Philippines.

Notes: Evidently Van Ooststroom erred in stating the type collection came

from Sarawak, because the locality given on the label —“*Muara, Brunei bay”

— corresponds to modern Brunei. The name of the collector is ambiguous: it

appears to be G.K. Gns, just as Van Ooststroom rendered it, but this name is

not identifiable with anyone known to have collected in the Malesian region

(Van Steenis-Kruseman, 1950, 1958, 1974). Although M. pulchra is reported

to occur in the Philippines (Coode et al., 1996: 61), and this is quite possible,

I have seen no authentic material from there. The large flowers of a vivid

yellow color make this species worthy of introduction to horticulture.

35. Merremia quinata (R. Br.) Ooststr., J. Arnold Arbor. 29: 417. 1948.

Ipomoea quinata R. Br., Prodr. 486. 1810; Convolvulus quinatus (R. Br.)

Spreng. — Type: Australia. Northern Territory: Arnhem Bay, Mallinson’s

Island, R. Brown sub J.J. Bennett 2755 (holotype, BM!; probable isotypes,

BK !)).

Syn.: /. hirsuta R. Br.; 1. pentadactylis Choisy.

Distribution: Myanmar, Thailand, Vietnam, China (Hainan), Taiwan,

Philippines, Papua New Guinea, Australia (Queensland, Northern Territory).

Surely more widespread in Malesia than the collections seen to date would

indicate.

Notes: Van Ooststroom (1953: 447) commented that continental Asian

specimens might not be conspecific with the specimens he examined from

Australia, Papua New Guinea, and the offshore islets between. There is great

variation in indumentum density, from glabrous to copiously hirsute with stiff

yellowish hairs, but otherwise, the morphology is remarkably consistent. The

peculiar crest on the seed apex is noteworthy and suggests seed characters

might be taxonomically informative if studied comprehensively. Like some

other Asian species (e.g., M. subsessilis, M. verruculosa) the corolla tube has

a slight curve in life that gives the flower a weakly zygomorphic aspect.

36. Merremia quinquefolia (L.) Hallier f., Bot. Jahrb. Syst. 16: 552. 1893.

Ipomoea quinquefolia L., Sp. Pl. 162. 1753; Convolvulus quinquefolius (L.)

L. — Type: Icon in: Plukenet, Phytographia plate 167, fig. 6. 1692 (lectotype

designated by D.F. Austin, Ann. Missouri Bot. Gard. 62: 182. 1975).

Distribution: Indonesia (Java), Australia (Queensland), Papua New Guinea,

French Polynesia (Society Islands: Tahiti), Nauru, Fiji (Labasa, Lautoka,

Vanua Levu), Tonga (Foa); native in tropical America, where it is widespread

500 Gard. Bull. Singapore 61 (2) 2010

and abundant, and also naturalized in Africa (Lebrun and Stork 1997, and

references cited therein).

Notes. I have seen a single old specimen (Jalmy s.n.) from Indochina,

without locality or date. Gagnepain and Courchet (1915: 239) listed this

specimen under their mixed concept “/pomoea pentaphylla”, and gave the

provenance as “Cochin-chine”, which is now Vietnam. The distributional

record for Vietnam is based on this Talmy collection.

37. Merremia rajasthanensis Bhandari, J. Bombay Nat. Hist. Soc. 84: 645.

1988.

Type: India. Rajasthan: Jodhpur District, Sardarsamand, 28 Aug 1975,

Bhandari 1976 (holotype, K, n.v.; isotype, JAC, n.v.).

Distribution: India; endemic to Rajasthan.

Notes: The author, when naming this species, stated that it “is closely allied

to Merremia quinquefolia (Linn.) Hall. f. ... However, it appears to be related

to the African M. palmata Hall. f. and M. verecunda Rendle.” and went on

to note, “The stem of this species is typically winged like that of African M.

pterygocaulos (Choisy) Hall. f’’ The similarity with these African species

has already been pointed out by Austin (1979: 56-57), citing B. Verdcourt,

who examined a duplicate in Kew of Bhandari 1971 (from the type locality)

and considered it to be M. palmata, though perhaps differing sufficiently

from the African populations of that species to be worthy of subspecific

rank.

I have not examined the type collection nor the other material cited

in the protologue but based on the detailed plate included there this species

appears to me to be conspecific with one of these lobed-leaved African

species; it is not remotely similar to the neotropical M. quinquefolia, which

presents an entirely different facies. Given the well-known biogeographic

affinity between eastern tropical Africa and the western Indian subcontinent,

it would be entirely possible for M. rajasthanensis to be no more than a

disjunct population of one of the widespread, variable African species.

38. Merremia rhynchorrhiza (Dalzell) Hallier f., Bot. Jahrb. Syst. 16: 552.

1893.

Ipomoea rhynchorrhiza Dalzell, J. Bot.3:179.1851.—Type: India.“in montibus

Syhadree, prope Tulkut-ghat, fl. Aug. et Sept.” Dalzell s.n. (holotype, K,

hve)

Distribution: India, western peninsula.

A Checklist of Merremia (Convotvulaceae) in Australasia and the Pacific 501

Notes: Writing more than a century ago Cooke (1905: 239) remarked

concerning M. rhynchorrhiza “The tubers are eaten and greedily sought for

by the natives, which accounts for the rarity of the plant. The leaves are also

used as a vegetable.” Perhaps this is why not a single recent collection has

been seen: the species should certainly be investigated in terms of IUCN

conservation status and threats to its survival.

Specimen examined: India. “Malabar. Concan, etc.. regio trop.” Stocks, Law

& Co, sn.(F!, UPS!, S!):

39. Merremia semisagittata (Griseb. ex Peter) Dandy in Andrews, Flw. PI.

Sudan 3: 123. 1956.

Ipomoea semisagittata Griseb. ex Peter in Engl. & Prantl, Pflanzenfam. 4

(3a): 31. 1891. —Type: “Mesopotamia”, no specimen cited.

Distribution: Saudi Arabia (Alfarhan and Thomas, 2001). otherwise known

from dryland African above the tropical belt (Sebsebe, 2001).

Notes. The nomenclatural status of several Ipomoea names published by

Peter (1891) is unsatisfactory because he did not cite specimens that could

be considered types under the ICBN. A thorough and comprehensive study

of the Peter names is needed to sort out the taxonomic concepts involved

and to neotypify the names. if it can be proven that there are no specimens

extant that could be eligible as original material for them.

I have seen no Asian material of this species.

Convolvulus sibiricus L., Mant. Pl. Alt. 203. 1771: Ipomoea sibirica (L.) Pers.

— Type: without locality, Herb. Linn. 218.5 (LINN!, lectotype, designated by

Staples in Taxon 55: 1022. 2006).

Distribution: India (Himalayas), China, Mongolia, NE Russia.

Notes: Chinese floras recognize 5 varieties: var. sibirica, vat. macrosperma

C.C. Huang, var. vesiculosa C.Y. Wu, var. trichosperma C.C. Huang, and var.

Jiuhuaensis B.A. Shen & X-L. Liu. See Fang and Staples (1995: 295) for a

key to the varieties. As pointed out years ago by Ferguson et al. (1977: 768)

the pollen of M. sibirica is unique in the genus Merremia. While the true

affinities of the species await disclosure, the pollen discordance is suggestive

that M. sibirica does not belong with any of the other species groups lumped

under Merremia.

Merremia sibirica was introduced to horticulture in Britain and Europe

502 Gard. Bull. Singapore 61 (2) 2010

in the nineteenth century but apparently does not survive in cultivation. The

flowers are small but borne in masses they are attractive; the repeatedly

monochasial cymes result in a distinctive candelabrum-shaped inflorescence

when well developed.

41. Merremia similis Elmer, Leafl. Philip. Bot. 1: 335. 1908.

Type: Philippines. Leyte, Palo, Elmer 7341 (isotypes, BO, n.v., E!, G!, K!,

NY!)

Syn.: ? Convolvulus distillatorius Blanco, ?Merremia distillatoria (Blanco)

Merr., p.p.

Distribution: Taiwan (Hengchun Peninsula), Philippines (Leyte, Luzon,

Mindanao, Panay, Samar, Sibuyan).

Notes:The reports of pinkish to red corollas in M. similis require confirmation;

this would be most unusual in a genus where white or yellow corollas are

the norm.

42. Merremia steenisii Ooststr., Blumea 20: 127. 1972.

Type: Papua New Guinea, Sepik District, Wewak-Angoram area, road from

Yangoru to Kworo, in grassland, | Sep 1959, Robbins 2316 (holotype, CANB,

n.v., photo!).

Distribution: New Guinea.

Notes: Based on a single unicate specimen; M. steenisii has not been

recollected since the type gathering was made in 1959. In describing it, Van

Ooststroom (1972) stated that M. steenisii is closely allied to M. aniseiifolia.

If that is the case, then the pollen of M. steenisii should also be examined, to

see whether the similarity it bears to Xenostegia spp. is superficial or whether

it should be transferred to that genus.

43. Merremia subsessilis (Courchet & Gagnep.) T.N. Nguyen, Vasc. Plts.

Syn. Vietnam. Flora 1: 183. 1990.— Ipomoea subsessilis Courchet & Gagnep.,

Not. Syst. (Paris) 3: 148. 1915. — Types: Vietnam. Lam, Mouret 189 (syntype,

P!); Ouonbi, Balansa 811 (syntype P!), same loc., Balansa 8/2 (syntype, P!);

baie de Fi-tsi-long dans Vile Verte, Balansa s.n. (syntype, P, n.v.); Quang-yen,

d’Alleizette 187 (syntype, P!).

Misapplied: M. collina sensu Songkhla & Khunwasi (1993).

Distribution: Thailand, Vietnam, Laos.

A Checklist of Merremia (Convolvulaceae) in Australasia and the Pacific 50

Notes: Although the protologue lists a third Balansa collection without

number that would be a syntype, I did not find this in Paris. There is, however,

a specimen with identical locality data numbered 813 and this may be the

third Balansa collection cited.

The foliose bract clasping the peduncle is distinctive in this species,

as is the slightly curved corolla tube, which gives the living flower a weakly

zygomorphic appearance.

44. Merremia thorelii (Gagnep.) Staples, Thai Forest Bull. (Bot.) 36: 98.

2008.

— Ipomoea thorelii Gagnep., Not. Syst. 3: 148. 1915. — Type: Vietnam, Ti-tinh,

Thorel s.n. [syntype, P (x2)!]; Laos, Khong, Thorel s.n. (syntype. P, n.v.).

Syn. M. collina S.Y. Liu, Guihaia 4: 199. 1984.

Distribution: China (Guangxi), Thailand, Laos, Vietnam.

45. Merremia tuberosa (L.) Rendle, Fl. Trop. Africa 4(2): 104. 1905.

— Ipomoea tuberosa L., Sp. Pl. 160. 1753; Operculina tuberosa (L.) Meissn.

— Type: Jamaica, Herb. Linn. 219.4 (LINN!, lectotype designated by D.F.

Austin, Ann. Missouri Bot. Garden 62: 182. 1975).

Syn.: ?Convolvulus paniculatus Blanco; Ipomoea tuberosa var. oligantha

Hassk.

Distribution: Pakistan, Sri Lanka, Myanmar, Thailand, Malaysia (Kuala

Lumpur, Sabah), Taiwan, Australia, French Polynesia (Society Islands:

Tahiti), Hawaiian Islands (Kaua‘i, O°’ahu, Maui, Hawai‘i), Micronesia

(Guam), Easter Island. Native in tropical America and widely dispersed

through cultivation.

Notes: Merremia tuberosa has been introduced throughout the tropics as

an ornamental. The vines are grown for the large, golden yellow flowers

as well as the papery, indehiscent fruits (wood rose), which are used in a

variety of handicrafts. It is surely more widespread in tropical Asia than the

distribution above would indicate. Merremia tuberosa frequently naturalizes

where it is introduced, especially in damp to wet sites with plenty of sun and

the rampant vines soon blanket roofs, trees, and powerlines.

46. Merremia umbellata (L.) Hallier f., Bot. Jahrb. Syst. 16: 552. 1893.

— Convolvulus umbellatus L., Sp. Pl. 1: 155. 1753; 1. umbellata (L.) G. Meyer,

non L. (1759). — Type: Icon in Plukenet, Phytographia plate 167, fig. 1. 1692

(lectotype, designated by D.F. Austin, Florida Sci. 42: 221. 1979).

Syn.: C. cymosus Desr., Ipomoea cymosa (Desr.) Roem. & Schultes, /.

504 Gard. Bull. Singapore 61 (2) 2010

modesta Choisy; I. pilosa Houtt.; 1. sepiaria Zoll. & Moritzi, non Koenig; /.

tonkinensis Gagnep., Merremia tonkinensis (Gagnep.) T.N. Nguyen.

Distribution: Nepal, India, Sri Lanka, Bangladesh, China, Myanmar, Thailand,

Cambodia, Laos, Vietnam, Malaysia, Singapore, Indonesia, Philippines,

Papua New Guinea, Australia, New Caledonia, Hawaiian Islands (O‘ahu),

French Polynesia (Society Islands: Raiatea, Tahiti); also widespread in

tropical Africa (but absent from Madagascar), the Indian Ocean islands, the

Americas.

Notes: Van Ooststroom (1939b, in Van Ooststroom and Hoogland, 1953)

recognized two subspecies of M. umbellata in the Malesian material he

studied: subsp. orientalis (Hallier f.) Ooststr., comprised plants native in

the Old World, whereas subsp. wmbellata comprised plants native in the

neotropics and introduced in Malesia (see 1953 reference for characters used

to distinguish the two subspecies). For many years I followed this taxonomy

implicitly, but as more material became available, both from Malesia and

continental Asia, the distinctions have blurred and many specimens could

_ not be satisfactorily placed in either taxon. Latterly I have abandoned all

attempts to recognize subspecies.

The difficulty is compounded because the small-flowered plants

typically called M. umbellata intergrade with larger flowered Asian plants

that have been named as M. bambusetorum and M. kingii. Indochinese

material called M. tonkinensis also belongs in this complex and while these

specimens seemed distinctive on first appraisal, they were later found to be

virtually identic with Bornean material that agrees almost exactly in terms

of small, oblong-lanceolate leaves, dense whitish indumentum, and small

flowers with white corollas. Clearly, this whole complex requires intensive

study to sort out the taxonomy. The most distinctive feature of all plants in

this complex is the presence of paired stipule-like appendages at the petiole-

stem junction; this feature was first pointed out by Songkhla and Khunwasi

(1993) and I have since found these appendages consistently in all specimens

checked; that they have never previously been reported is remarkable. On

densely hairy specimens the tiny appendages may be hidden among the

trichomes, but careful examination reveals that they are present.

47. Merremia verruculosa S.Y. Liu, Bull. Bot. Res., Harbin 7(2): 133. 1987.

Type: China. Guangxi: Yongning Xian, SY. Liu & S.J. Wei 1355 (holotype,

GXCM, n.v.; iso, KUN, n.¥v. ).

Distribution: China, Thailand, Laos.

A Checklist of Merremia (Convolvulaceae) in Australasia and the Pacific 50

Notes: The excellent figure accompanying the protologue (p. 136) provides

a clear visual image for this species; the curious warty outgrowths on the

abaxial side of the outer sepals are distinctive and this species is not easily

mistaken for anything else. As in M. subsessilis, the corolla tube in life has a

slight curvature, giving the flowers of M. verruculosa a weakly zygomorphic

appearance.

48. Merremia vitifolia (Burm. f.) Hallier f., Bot. Jahrb. Syst.16: 552. 1893.

— Convolvulus vitifolius Burm. f., Fl. Ind. 45. t. 18, f. 1. 1768: Ipomoea vitifolia

(Burm. f.) Blume — Type: Indonesia. Java, Garzin s.n. (G-Burman!, lectotype

designated by Staples and Jacquemoud, Candollea 60: 449. 2005).

Syn.: C. angularis Burm. f.; 1. vitifolia var. angularis (Burm. f.) Choisy.

Distribution: Nepal, Bhutan, India, Sri Lanka, Bangladesh, Myanmar,

Thailand, Laos, Vietnam, China (Hainan, Yunnan), Taiwan, Malaysia

(Sabah), Indonesia (Java, Maluku, Selebes, Sumatra, Timor), Philippines

(Balabac, Luzon, Palawan, Paragua).

Notes: A species easily recognized; corolla color varies from pure bright

yellow, to yellow with a wine-red base of the throat, through paler shades of

yellow to almost white.

49. Merremia yunnanensis (Courchet & Gagnep.) R. C. Fang, Fl. Reipubl.

Popularis Sin. 64(1): 74. 1979. — Ipomoea yunnanensis Courchet & Gagnep..,

Notul. Syst. (Paris) 3: 151. 1915.—Types: China. Yunnan: Ta-pin-siou, Ducloux

4454 (syntype, P!), Lou-pou, prés Tong-tchouan, Ducloux 6398 (syntype,

P!), col de Piou-sé, Delavay 3184 (syntype, P!), pres de Ta-pin-tzé, Delavay

3931 (syntype, P!); Western China, without province or locality: Wilson 4183

(syntype, P!, iso, A!).

Syn.: 1. yunnanensis var. uniflora C.Y. Wu.

Distribution: China (Guizhou, Sichuan, Yunnan). Endemic.

Notes: Chinese floras recognize three varieties: var. yunnanensis, vat.

glabrescens (C.Y. Wu) R.C. Fang, and var. pallescens (C.Y. Wu) R.C. Fang.

See Fang and Staples (1995: 296) for a key to the varieties.

Undescribed species

There are published reports of undescribed species of Merremia, for

example from Brunei (Coode et al., 1996: 61), as well as new species yet

to be described for Australia (R. Johnson, pers. comm. 4 Feb 2009). I have

506 Gard. Bull. Singapore 61 (2) 2010

seen no specimens for either of these, but the reports indicate there are

still novelties awaiting discovery. Indeed, given the vast tracts of continental

tropical and subtropical Asia that remain severely under-collected — Laos,

Cambodia, Myanmar, and eastern Tibet among them, a region here shown

to be a center of species richness for Merremia — it is quite likely that the

number of species will increase in future.

Excluded Species & Synonyms

Merremia angustifolia (Jacq.) Hallier f. = Xenostegia tridentata (L.) D.F.

Austin & Staples

Merremuia boisiana var. rufopilosa (Gagnep.) C.Y. Wu = Merremia boisiana

var. fulvopilosa

Merremia caespitosa (Roxb.) Hallier f. = M. hirta

Merremia chryseides (Ker-Gawl.) Hallier f. = M. hederacea

Merremia collina S.Y. Liu = M. thorelii

Merremia convolvulacea Dennst. ex Hallier f. = M. hederacea

Merremia crispatula Prain = Operculina petaloidea (Choisy) Ooststr.

Merremia decurrens (Hand.-Mazz.) H.S. Kiu = M. hirta

Merremia distillatoria (Blanco) Merr. = M. similis

Merremia gangetica (L.) Cufod. = Cocculus hirsutus (L.) Diels; vide Van

Ooststroom (1934: 245).

Merremia hastata Hallier f. = Xenostegia tridentata (L.) D.F. Austin &

Staples; vide Van Ooststroom (1972b: 939) for nomenclatural notes

on the Merremia name.

Merremia longipedunculata R.C. Fang = M. poranoides

Merremia medium (L.) Hallier f. = Xenostegia medium (L.) D.F. Austin &

Staples

Merremia nymphaeifolia (Blume) Hallier f. = M. peltata

Merremia pentaphylla (Jacq.) Haller f. = M. aegyptia

Merremia petaloidea (Choisy) Burkill = Operculina petaloidea (Choisy)

Ooststr.

Merremia platypeltis Prain = Operculina riedeliana (Oliv.) Ooststr.

Merremia riedeliana (Oliv.) Hallier f. = Operculina riedeliana (Oliv.)

Ooststr.

Merremia tridentata (L.) Hallier f. = Xenostegia tridentata (L.) D.F. Austin

& Staples

Merremia tridentata subsp. genuina Ooststr. = Xenostegia tridentata (L.) D.F.

Austin & Staples

Merremia tridentata subsp. hastata Ooststr. = Xenostegia tridentata (L.) D.F.

Austin & Staples

Merremia triquetra (Vahl) Roberty = Operculina turpethum (L.) S. Manso

A Checklist of Merremia (Convolvulaceae) in Australasia and the Pacific 507

Merremia turpethum (L.) Rendle = Operculina turpethum (L.) S. Manso

Merremia wilsonii Verdc. = M. hungaiensis

Table 1. Number of total species by region, unique species and infraspecific taxa of Merremia

in different regions of the world.

Region total # species # unique # infraspecific taxa

in region species

Americas 29 ; Pr, as

Africa & Madagascar 39 3]

Australia 13 rae ; —

Asia & Pacific et ens (99 40. , nebo

TOTAL Se |e

Appendix: data sources for Table 1, grouped by region:

Americas (North, Middle, South, and West Indies)

The primary source used is an unpublished manuscript prepared for Flora

Neotropica (Austin and Staples, circa 1979), which expands on and updates

the revision by O’Donell (1941). A number of new neotropical species

have been described in recent years (McDonald, 1987, 2008; Valencia and

Martinez, 1995) that have been incorporated into the working list.

Arrica (including Madagascar)

The primary source for the African tropics is the Convolvulaceae listing

from Lebrun and Storck (1997) with additional recent floras consulted for

South Africa (Meeuse and Welman, 2001), Madagascar (Deroin, 2001),

Ethiopia (Sebsebe, 2006), and Somalia (Thulin, 2006).

AUSTRALIA

Names were compiled from diverse historical literature for Australia, and

updated against two floras that cite Australia as part of the distribution (Van

Ooststroom and Hoogland, 1953; Fang and Staples, 1995). Additional names

were found in Elliot and Jones (1993). Bob Johnson (pers. comm., June

2007) kindly reviewed the working list of names that resulted and provided

the current taxonomy to be adopted for the forthcoming Flora of Australia

account.

ASIA AND PACIFIC

For Asia the primary flora used as the foundation of the species list was

508 Gard. Bull. Singapore 61 (2) 2010

Malesia (Van Ooststroom and Hoogland, 1953, and subsequent addenda

and corrigenda) with updates and additions from the floras for Saudi

Arabia (Alfarhan and Thomas, 2001), Pakistan (Austin and Ghazanfar,

1979), Sri Lanka (Austin, 1980), India (Clarke, 1883; Cooke, 1905; Gamble,

1923; Saldanha and Nicolson, 1976; Babu, 1977; Bhandari, 1988; Panigrahi

and Murti, 1989), Bhutan (Mill, 1999), Bangladesh (Khan, 1985), Myanmar

(Kress et al., 2003), Thailand (Songkhla and Khunwasi, 1993; Staples et al.,

2005; Staples and Traiperm, 2008), China (Fang and Staples, 1995), Taiwan

(Staples and Yang, 1998), Vietnam (Nguyen, 1990), Brunei (Coode et al.,

1996), Peninsular Malaysia and Singapore (Turner, 1995).

For the Pacific region additional references consulted included

floras for Micronesia (Fosberg and Sachet, 1977), the Marquesas (Sachet,

1975), New Caledonia (Heine, 1984), and Staples (2009). Specimens I have

examined in the past two decades have been used to correct and expand the

distributions compiled from the literature.

Index of numbered collections examined

The following alphabetical list includes numbered herbarium specimens I

have examined from Asia and the Pacific. Numbers correspond to the list

below, which is identical with the numbering used in the text. Type specimens

have been indicated with (T).

1. Merremia aegyptia 16b. Merremia dissecta var. edentata

2. Merremia aniseiifolia 17. Merremia eberhardtii

3. Merremia bambusetorum 18. Merremia elmeri

4. Merremia bimbim — var. elmeri

5. Merremia boisiana — var. glaberrima

— var. boisiana 19. Merremia emarginata

— var. fulvopilosa 20. Merremia gemella

— var. sumatrana — var. gemella

6. Merremia borneensis — var. splendens

7. Merremia bracteata 21. Merremia gracilis

8. Merremia caloxantha 22. Merremia hainanensis

9. Merremia calyculata 23. Merremia hederacea

10. Merremia cissoides 24. Merremia hirta

11. Merremia clemensiana 25. Merremia hungaiensis

12. Merremia cordata — var. hungaiensis

13. Merremia crassinervia — var. linifolia

14. Merremia davenportii 26. Merremia incisa

15. Merremia dichotoma 27. Merremia kingii

16a. Merremia dissecta var. dissecta 28. Merremia korthalsiana

A Checklist of Merremia (Convolvulaceae) in Australasia and the Pacific 509

29. Merremia mammosa 41. Merremia similis

30. Merremia pacifica 42. Merremia steenisii

31. Merremia palmata 43. Merremia subsessilis

32. Merremia peltata 44. Merremia thorelii

33. Merremia poranoides 45. Merremia tuberosa

34. Merremia pulchra 46. Merremia umbellata

35. Merremia quinata subsp. orientalis

36. Merremia quinquefolia — subsp. umbellata

37. Merremia rajasthanensis 47. Merremia verruculosa

38. Merremia rhynchorrhiza 48. Merremia vitifolia

39. Merremia semisagittata 49. Merremia yunnanensis

40. Merremia sibirica

— var. sibirica

— var. macrosperma

— var. vesiculosa

— var. trichosperma

— var. jiuhuaensis

1984 Sino-American Botanical Expedition 1323 (25).

Aban Gibot SAN 76999 (46): SAN 90220 (21): SAN 95362 (32): SAN 96805 (21):

SAN 96627 (46). — Abang Mohtar S. 56140 (46).— Abang Mohtar er al. S. 56133 (6).

— Abbe, L.B. 9373 (48): 9374 (46). — Aldén, B. et al. 1798 (25).— Alston, A.H.G. 14781

(5).—Ambol, A. AA-45 (32).— Ambriansyah & Arifin AA-190 (32).— Amdjiah 918

(18).— Amin et al. SAN 121515 (32): SAN 117823 (32).— Amin, G. et al. SAN 67448

(21); SAN 117823 (32); SAN 121515 (32): SAN 121576 (18): SAN 96557 (21): SAN

96729 (21).—Amor, E.A. 48 (32); 66 (30); 92 (30).—Amphon 68 (48).— Anderson, E.F.

4285 et al. (6). — Anderson, J.A.R. & I. Paie S. 28489 (34): S. 16009 (46). — Anthony,

Y.T. A-699 (46); A-775 (32). — Arbainsyah er al. AA-3135 (23).— Argent, G. & Iqbar

9964 (48): 9965 (46); 9980 (32); 9986 (5). — Argent, G. & E. Campbell 411854 (T)

(21). — Argent, G. et al. 93119 (32): 108280 (32); 1987171 (21).—Arifin, Z. BRF-1812

(21). — Arishah FM-09583 (46). — Arnott, D. 2288 (23). — Ashton, P.S. S. 19005 (34):

S. 19795 (18). — Asik Mantor SAN 120221 (46); SAN 136121 (21). — Austin A-1212

(46). — Australian National University ANU 1715 (16). — Averyanov, L. et al. VH

1380 (5): VH 1063 (5); VH 830 (3): VH 1012 (46). — Awang Yakup S. 8280 (34).

B.R. E-48 (23). — B.G.O. staff QSBG 2427 (29): QSBG 5376 (23): QSBG 8386

(23); QSBG 8565 (48); QSBG 8788 (48); 17 (48): 3556 (32). — B.S. & S.P. 1519 (46).

—Bacani, E. FB-16477 (20); FB-16490 (24): FB-16656 (20). — Backer, C.A. 21115

(16); 33473 (29). — Bacon, P. PB-1 (30): PB-2 (T)(7). — Balakrishnan, N. NBK-1153

(46). — Balansa, B. 811 (T)(43): 812 (T)(43): 813 (T)(43): 3535 (46); 3536 (23); 3537

(46): 3538 (46): 4459 (T)(33).— Balgooy, M.M.J. van 1735 (32); 1768 (46).— Banyeng,

L. et al. S. 57355 (32). — Bartlett, H.H. 8159 (32). — Baré, D. 114 (32). — Bauman-

510 Gard. Bull. Singapore 61 (2) 2010

Bodenheim, M. 11317 (32). — Beauvais, J. 704 (T) (25); 1205 (T)(25). — Beccari, O.

665 (32); 3954 (T)(13). — Bermejos, J. BS-265 (32). — Bernardi, L. 14636 (16a). —

Beusekom, C.F. van 12535 (24).— Beusekom, C.F. van & C. Charoenphol 1687 (46);

2017 (24). — Beusekom, C.F. van & C. Phengklai 2649 (24).— Beusekom, C.F. van

et al. 3825 (35); 3953 (19); 4002 (23); 4287 (43); 4473 (48); 4474 (46); 4705 (46). —

Bhargava, N. 3375 (32). — Blake, S.T. 16701 (16); 23196 (16); 23286 (23). — Blewett,

J.B.J. B-19 (21); B-20 (21). — Bloembergen, S. 4612 (32). — Bodinier, E. & Ducloux

272 (T)(25); Bodinier, E. & L. Martin 1806(T), in part (49); in part (25). — Boeea,

Rahmat Si 1330 (32); 1532 (23); 1645 (32); 2268 (46); 3778 (23). — Bois, D. 138 (T)(5).

— Bon, l’abbé 2425 (T)(46); 2813 (46); 4801 (5); 5216 (T)(46). — Boom, B.K. 25770

(40). — Bor, N.L. 6294 (27); 6689 (33); 6774 (27); 16699 (29). — Borden, T.E. FB-2714

(20). -Bourdy, G. 675 (30). — Boyland, D.E. & J.G. Gilliett 367 (16). — Bragg, K. 22

(48); 217 (48). — Brass, L.J. 545 (46); 2712 (32); 7578 (T)(20); 7646 (20); 7778 (20);

27240 (24); 27264 (32); 28540 (30); 29365 (20). — Brass, L.J. & C.T. White 145 (32).

— Bray, P.S. FRI 11786 (46). — Brook, Miss W.M.A. 8281 (18); 8828 (23). — Brooke,

W. 10802 (23).— Brown, R. [Bennett catalogue no.] 2755 (T)(35); 2756 (35). — Bryan

Jr., E.H. 584 (32). — Bunchuai, K. 4 (46). — Biinnemeyer, W. 12477 (48). — Bureau of

Science BS 483 (48).— Burgess, P.F. FRI 9222 (32); FRI 9253, in part (32), in part (6);

FRI 10161 (6); SAN 25166 (18). — Burkill, 1-H. Mo-2829 (16). — Burley, J.S. NGS-651

(32); NGS-666 (46). — Bush, W. 32725 (46); 32727 (46). — Byrnes, N. 2469 (20).

C.S.S. 259 (32). — Cabalion, P. 316 (32). - Campbell, E.J.F EC-34 (6). — Carr, C.E.

12936 (20); 12319 (20); 12632 (32); 12850 (46). — Carter, B.J. 564 (23). — Cavalerie,

J. 3640 (33). — Chai, P.C.K. S. 34126 (13). — Chang, C.-E. 1986 (20); 2467 (41); 6760

(41). — Chantaranothai, P. et al. 927 (48). — Chantharanothai, P. & Parnell, J. 90/75

(43). -Charoenphol, C. et al. 4309 (27). — Cheeseman, L. 16 K (32). — Cheke, AS.

T-9 (48). — Cheng, S.K. CL 560 (3). — Chevalier, A. [legit A.Fleury] 30239 (44). —

Ching, R.-C. 7344 (T)(23); 7451 (46); 7557 (46); 7775 (24); 8251 (46); 20823 (25);

21076 (25); 21660 (T)(12). — Chippendale, G. NT-4393 (1); NT-4393 (1). — Chitr 166

(3). — Christophersen, E. 23 (32); 576 (32). — Christophersen, E. & Hume, E.P. 1897

(32). — Chun, N.-K. 40259 (46). — Chun, W.-Y. 7632 (23). — Chung, C.-S. 2581 (46). —

Clarke, C.B. 9189A (T)(33); 9680 (T)(33); 9956 (T)(33); 12384A (T)(27); 24775A

(27): 40909A (33): 40909D (33): 40914A (27).— Clarkson, J.R. 4466 (35).— Clarkson,

J.R. & Neldner, V.J. 9437 (23). — Clarkson, J.R. & Simon, B.K. 6975 (20). — Clemens,

J. & Clemens, M.S. 3238 (5); 4091 (17); 5075 (46); 21133 (T)(11); 21134 (13); 21527

(28). — Clemens, M.S. 256 (32); 785 (20). — Collett, C.B. 488 (T)(27). — Comanor,

P.L. 703 (10); 803 (46). — Comins, R.B. 204 (30). — Conover, J.T. 512 (23); 575 (23). —

Coode, M.J.E. et al. MC-7958 (32). — Cooray, R.G. 70032308 P, in part (20), in part

(23). — Craven, L.A. 4393 (20). — Crosby, C.S. 131 (32). — Cuadra, A. A 2233 (32). —

Cuming 1030 (19). — Cunningham, A. 110 (T)(20). — Curran, H.M. FB8873 (20).

Dahlstrom, E. 500 (24).—d’Alleizette 100 (T)(43); 187 (T)(43).— Darbyshire, P.J. 651

(16): 688 (20); 821 (20).— Daud, Z. et al. FM-2882 (45).— Day, A.L. 575 (32).— Dayang

Awa, A.L. & Ilias, P. S. 47474 (18). — Dee 952 (24); 1014 (43). —- Degener, O. & Bush,

W. 32727 (46). — Degener, O. & Degener, I. 36052 (36). — Delavay, J.M. 2761 (T)(25);

3073 (12): 3184 (T)(49): 3931 (T)(49).— Din 160 (48). — Diwol Sundaling SAN 97036

A Checklist of Merremia (Convolvulaceae) in Australasia and the Pacific S11

(21). -— Doan Khao & Viét-Trung 4606 (43). — Drake, E. 5 (32). — Ducloux, F. 4454

(T)(49): 6398 (T)(49): 7148 (49): 7360 (25). — Ducloux, F. [legit S. Ten] 1419 (49). —

Diirbye 36 (10); 411 (16). — Duthie, J.F. 8346 (19).

Eames, A.J. 125 (16). — Eberhardt 1440 (46): 1708 (T)(17): 1994 (46). — Edanio, G.E.

PNH 15617 (41). — Edgeworth, M.P. 423 (40). —- Elmer, A.D.E. 5577 (20): 7341 (T)

(41); 8920 (46): 10836 (32); 12061 (41): 12603 (46); 12880 (48); 15602 (41): 15605 (32);

18477 (20); 20260 (6); 20329 (23): 20396 (T)(18); 20990 (T)(6). — Enggoh, A. 10530

(32). — Esquirol, J. 1065 (T)(25).

Falanruw, M.V.C. MVCF-5392 (32). — Falconer 347 (48). — Feng, K.-M. 2252 (49):

2716 (49). — Fernandes, J. 938 (46). — Fidilis Krispinus SAN 82187 (32): SAN 139370

(46). — Flecker, H. 3258 (36); 4016 (32).— Fleury, A. [sub Chevalier no.] 30239 (44). —

Florence, J. 3620 (32); 3698 (32); 4831 (32); 5023 (32); 6130 (32); 7717 (23); 9291 (45);

10377 (46); 10703 (16): 11474 (36). — Florence, J. & Sykes, W. 11240 (32). — Florence,

J. & Tahuaitu 11487 (32); 11760 (32). — Fliigel, H.J. FI-T 66 (46): FI-T 67 (48); FI-T

114 (23): 7006 (48): 7007 (46); 7008 (46). — Forbes, H.O. 2651 (48).— Forrest, G. 10346

(49): 10930 (49): 11121 (12): 11314 (40); 11676 (49); 12474 (49); 14596 (25); 14970 (40);

28952 (40); 30547 (40); 71111 (T)(8).— Forster, R.H. 2716 (48). — Fosberg, F.R. 26572

(32); 31231 (20); 31240 (23); 58373, in part (30), in part (32); 58765 (36); 60938 (16b):;

61187 (32); 62118 (36); 62726 (46); 63526 (32); 63680 (32); 64918 (32); 65021 (46). —

Fosberg, F.R. & Florence, J. 62640 (46). — Fosberg, FR. et al. 51137 (19). — Franck,

C.W. 404 (24). — Froideville, C.M. de 591 (24). — Fuentes, F. 2 (45). — Fujikawa, K. &

Nomachi, A. 35807 (46).— Fukuoka, N. T-7337 (48): T-7338 (46). — Fukuoka, N. & Ito,

M. T-34720 (24). — Fukuoka, N. & Koyama, H. T-62040 (24); L-65143 (43).— Fung, H.

A 456 (23).— Fung, H. [Lingnan Univ. accession] A-360 (23).

Galathea Expedition 1203 (46).— Galore, M. NGF 17564 (35).— Gamble, J.S. 3408A

(T)(27); 3409A (T)(27); 10580 (T)(27). — Gaoligong Shan Exped. 1997-8540 (25);

1997-9074 (40); 1997-9928 (8). — Garrett, H.B.G. 250 (23): 251 (20): 270 (46). — Gates,

F.C. 7126 (20): 7216 (48). — Gaudichaud 24 (1). — Geesink, R. & Maxwell, J.F. 7805

(20). — Geesink, R. et al. 7718 (46); 8905 (21). - Geh & Samsuri, A. GS Y-403 (11);

GSY-1005 (34). — George, M. et al. SAN 123735 (23). — Gillison, AN. NGF 22228

(16).—Gns, G.K. 14 (T)(34).— Grant, M.L.5359 (16b): 4271 (32).—Green, PS. RSNH-

1316 (32). — Greenwood, W. 199 (36): 199A (36).— Greuter, W. 24604 (20). — Griffith

East India Co. Herbarium 5862 (48): 5865 (T)(27): 5871 (24); 8565 (46). — Guam

Experiment Station 18 (20).— Guppy, H.B. 197 (30).

Hallier, H. C.164a (29); C.164b (29); C.164c (29); C.164d (29); D.206a (46); 1844

(18): 3006 (28); 3370 (T)(18). — Hamid, H.A. 48228 (34). — Hamid, R. FM 8424

(46). — Hamilton, C. & Congdon, G. 12 (46). — Hansen, B. & Smitinand, T. 11873

(46). — Hansen, C. 606 (11); 1409 (18). — Hardial, S. & Samsun, A. HS-1025 (32).

— Harmand 142 (5); 1827 (T)(5). — Hartley, T.G. 10209 (35). — Haviland, G.D. 3055

(23). — Haviland, G.D. & Hose, C. 3523A (T)(13); 3523E (T)(13); 3523K (T)(13). —

Hellwig, F. 189 (35). — Henderson, M.R. SF-22911 (20). — Henderson, R.J. H-2422

(46). — Henry, A. 1946 (24): 9786 (46): 10174 (48): 10276 (25); 10987 (49); 13441 in

S12 Gard. Bull. Singapore 61 (2) 2010

part (8), in part 12). — Henty, E.E. NGF 10681 (35); NGF 14335 (35). — Herbarium

Trip 605 (29): 639 (3); 669 (43): 687 (43): 693 (44); 1229 (44). — Herbst, D. 749 (45). —

Herbst, D. & Spence, G. 5027 (45); 5245 (45); 5575 c (45). — Herre, A.W. 305 (20): 317

(20). — Hira Lal er al. 31919 (19). — Hoff, M. 4264 (32). - Hoogland, R.D. & Craven,

L.A. 10327 (20). — Hoover, S. et al. 5016 (46). — Hosaka, E. Y. 3301 (32). — Hose, C.

221 (32).— Hosseus, C.C. 162 (35): 175 (23): 373 (46); 466 (46). — How, F.-C. 70817 (5):

71596 (46); 71693 (T)(22): 71860 (5).— How, F-C. & Chun, N.-K. 70028 (48). — Hunt,

P.F. 2172 (30). — Hyland, B. 3111 (32).

Ilias, P. & Jegong S 51849 (28).— Insani, A.R. SAN 12 (20).

Jacobs, M. 9239 (32); 9683 (32). —Jafu, S.M.H. 1616 (23). —Jaikrasane, P. 173 (10); 252

(10). — James, D.M. S 33437 (32). — Jarvie, J. & Ruskandi, A. 5025 (21). — Jayasuriya,

A.H.M. et al. 1494 (46). — Jayasuriya, M. 347 (46). — Jayasuriya, M. et al. 639 (20). —

Jiang, F. 7710 (5). — Jibrin, S. JS 24 (32); JS 214 (32). — Juan, G.E. 188 (29): 446 (29).

—Julaihi et al. S. 78918 (13). — Jusimin, D. JD 285 (6).

K.K. 127 (27); 1584 (27). — Kadir A-540 (32): A-670 (23): A-2034 (6); A-2734 (6).

— Kajewski, S.F. 1404 (32). — Kanehira, R. 297 (32). — Kansuntisukmongkol, K. 127

(27): 1584 (27). — Kartawinata, K. 960 (32). — Kato, M. et al. 828 (46): C-7941 (20).

— Kaudern, W. 375 (32). — Keenan, J. et al. 1834 (29). - Kemppainen, B. 139 (20). —

Kenneally, K.F. 9751 (16). — Kenneally, K.F. & Hyland, B.P.M. 10475 (46). — Kerr,

A.F.G. 1609B (23): 2256 (35); 4014 (46): 4769 (46): 4825 (35); 9741 (24): 10427 (3):

13461 (24): 13569 (46): 13605 (24): 13935 (20): 15681 (32); 16268 A (46): 16581 (32):

17112 (32); 17704 (T)(3); 18138 (46); 19817 (24). — Kessler, P. et al. Berau-259 (6).

— Kessler, P.K. et al. PK-2303 (46). — Khunwasi, C. 1 (3); 29 (46); 38 (46); 40 (3); 42

(43): 43 (32). — Kiew, R. RK-345 (18): RK-1281 (46); RK-1282 (46); RK-3739 (46).

— King 55457 (32). — King, G. 180 (T)(27). — Kingdon Ward, F. 5282 (49); 9003 (23);

11240 (46): 18209 (27). — Kjellberg, G. 700 (32); 1372 (32); 4006 (48). — Kleinhof 85

(T)(19). — Ko, S.-P. 51633 (46): 52209 (46). — Kobbi, W.H. FB-6599 (32). — Kodoh

Tarodop SAN 83685 (6).— Koizumi, M. & Lalo MK-774 (21). — Kondo, Y. & Edano,

G. PNH 38864 (32): PNH 38930 (46). — Konta., F. et al. 4279 (27): 4321 (48): 4488 (23).

— Koonkhunthod, N. er al. 324 (48). — Kooy, C.W. 794 (15); 832 (15). — Koroivebau,

D. DA-10496 (36): DA-11503 (32); DA-13194 (30). — Korthals 237 (T)(28): 241 (32).

— Kostermans, A.J.G.H. 82 (32); 4809 (21); 10035 (6); 27984 (46). — Koyama, H. er

al. T-30084 (3): T-30131 (3): T-30771 (23); T-30924 (35): T-31008 (43); T-31079 (46):

T-32308 (48): T-32961 (46): T-33019 (46): T-48942 (27): T-49060 (23). — Koyama, T. et

al. 15313 (48); 15314 (46): 15315 (46): 15316 (46): 15446 (46): 15543 (27). — Krukoff,

B.A. 4434 (32).— Kukkonen, I. 10871 (36).— Kulip, J. & Gon, G.S. SAN 137093 (6). -

Kunstler, H. 1193 (24). — Kuntze, O. 4890 (32); 4900? (46); 5263 (24); 5471 (20):5785

(20); 5970 (46): 6184 (24): 6597 (24). — Kurz, S. 1915 (29).

La Saviniere, E. de 1167 (20). — Lai et al. S. 67718 (11). — Lai, J. et al. LJ-317 (28). -

Larsen, K. 8267 (23). — Larsen, K. & Larsen, S.S. 32706 (46); 40289 (44); 47251 (48).

— Larsen, K. et al. 31875 (29): 44458 (46): 44737 (46). — Lau, S.-K. 155 (5); 364 (35):

578 (46): 771 (23); 1033 (48); 1058 (46): 1124 (19); 1151 (23); 3193 (48); 6315 (5);

A Oheckine of Mexsoams (Conroivadacese) m Amsmrdlasia and the Pacific

24913 (23)- 25134 (23).— Lazandes. M_ 3905 (36)- 8864 (23). — Leach. BJ. BSIP 7855

(32). — Ledermann_ C. 6723a (32): 8116 (32): 13233 (30).— Lee. S et al. WC-78 (24).

— Lei C-L 614 (5); 865 (5).— Leland_B. ef al. 16 (46).— Leong, P. et al. SING2005-26

(24).— Levine. C_O. 950 (23): 1246 (23)- 1831 (46)- 1907 (24): 1929 (23).— Lépme.I. 39

(32). — Laang_ H_-Y. 61780 (5)- 63849 (46)- 63937 (46): 65362 (46): 66110 (46): 66401

(23): 66420 (46); 65352 (48). — Lampricht. H.W. 928 (1)(25). — Lingnan University

jcollector H_ Fung] A-360 (23).— Lister, LL. 136 (46).— Lomudmm.T_ LT-282 (32): LT

323 (32).— Lorence. L_ LL-33 (6).— Lorains_ 1. A_ 4234 (5): 4723 (3); 12281 (5): 12945

(46); 13054 (24): 16126 (16).— Lundqvist. N. 8888 (46).— Lwin U Thein 498 (46).

MG Gale-? 5266 (48).— MG Kyaw San 13038 (46): 13046 (48).— MacGregor. RW.

2574 (46). — Madam, L.- SAN 63531 (21). — Madulid. DA. e al. PPI-3138 (2). —

Mahani FM-242? (46). — Mahmood. S. 64221 (46). — Maidum, L. ML-?17 (32). —

Majawat_ G. et al. SAN 135601 (21).— Mamit_ LD. S. 33437 (32).— Mamsus et al SAN

117156 (21). — Martin. L_ 1806 (25).— Matus. K- RK-18 (6): AU-36 (28).— Maxwell.

LE 01-405 (44)- 01-478 (44)- 01-592 (43)- 71-730 (43): 76-387 (47): 85-1054 (20): 83

1087 (46)- 86-1 (46)- 87-170 (23)- 88-128 (48): 88-160 (46)- 88-308 (48): 89-1469 (23):

90-36 (48): 90-315 (46)- 91-1114 (23)- 94-1272 (46): 95-108 (48): 96-218 (27): 97-372

(48): 98-1089 (29). — McClure. FA. 1744 (46): 3090 (5): 8223 (24): COC-8223 (24):

LU-13121 (46).— McDonald. LA. & Ismail NGS-4141 (48). — McGregor. R-C. 153

(32); BS-531 (45).— McKee. HLS. 1633 (30): 6465 (46): 29260 (32): 31250 (46): 35717

(G2); RSNH-24261 (32).— McLaren's Collector 24 (12).— McMillen. [R_ 147 (48):

155 (48): 170 B (46). — Meebold, A. 5158 (46): 5878 (27)-14214 (46); 26554 (32). —

Meger, W. SAN 122495 (48)- SAN 43309 (28).— Memill_ E_D. 447 (46)- 644 (20)- 698

(32). 733 (48)- 738 (41): 3398 (23): BS 2048 (46): BS 3419 (23): BS 3615 (T)(24). —

Memill_ E_D. ‘Species Blancoanae” 357 (19): 679 (24): 953 (32).— Meyer. R- FB 2502

(20).— Middleton, D_I_ et al. 2261 (46)- 2366 (46)- 2338 (23): 2773 (3).— Millar. AN &

Vandenberg. |. NGF 48518 (30).— Millspaugh. C_F. 2812 (23): 2824 (20).— Moll. VW.

BW 9511 (T) (2).— Moore. LW.211 (32).— Morat_ P. 6925 (32): 7078 (32).— Morrison.

C. 181 (30). — Moulton. LC. SF 6721 (28). — Mouret 189 (T)(43). — Mukerjee. SK_

1548 (46).— Munting. F_ S. 56351 (32).— Murata. G_ & Phengklai_ C T-50398 (35)-

T50425 (35). — Murata, G. 2 al. B-3023 (18): T-41732 (43): T-51003 (46): T-51271

(43). — Murata. I er al. 22110 (46)- 41585 (46): 41645 (48): 41700 (46). — Murdock.

AM 14 (16b).— Murray. E_ 26 (16b).— Must J. 1040 (35): 1115 (20).

Naktsi BSIP 7324 (30).— Nanakorn. W. et al. $499 (43).— Nangkat. N NN-148 (34).

— Nelson, P. 105 (23): 527 (45).— Newman. M_ e7 al. 1410 (46): LAO-355 (48): LAO-

659 (8)- LAO-796 (24)- LAO-958 (23). — Ng. ESP. FRI 27377 (46): FRI 27379 (32).

— Nielsen. LC. ef al. 1623 (43). — Niyomdham. C. 1856 (24): 4904 (3): 3297 (27). —

Niyomdham. C. ef al. 2791 (32)- 5967 (46).— Nooteboom. H_P. NB-6071 (32).— Nor

Ezzawanis & Rafidah. AR. FRI 54630 (46). — Noreen. ML ez al. SAN 147163 (32).

—Nyman_ E_ 760 (24): 1048 (46).

O'Reilly. K. North Queensland Nat. Club 3543 (36). — Orchard, AE. 4654 (24). —

Osbeck 11 (T)(24).

514 Gard. Bull. Singapore 61 (2) 2010

Paisooksantivatana, Y. Y571-81 (27). — Pan, M. SAN 83697 (45). — Panatkool, M.

207 (35); 441 (35). — Panigrahi, G. 21672 (29). — Panoff, F. 60 (30). — Parham, J.W.

& Parham, M.E. 16958 (32). — Parker, R.N. 3080 (46); 3189 (48). — Parkinson, C.E.

4331 (23).— Patrick SAN 117671 (23). — Petelot 2526 (46); 2533 (43). — Phengklai, C.

12313 (43); 12848 (46). — Phengklai, C. et al. 3096 (48); 3359 (23); 3426 (35); 3511 (24);

3603 (23); 3616 (44); 3634 (24); 11008 (27); 11250 (23); 12953 (46). — Philipson, W.R.

10414 (32).— Phusomsaeng, S. 24 (46); 368 (32); 369 (48). — Phusomsaeng, S. et al. 125

(24). — Pierre, L. 13 (46); 1013 (23); 1482 (23).— Pleyte, D.R. 101 (46).— Ploenchit 427

(46); 1354 (48); 1505 (3). — Po Chin 4216 (24). — Po Khant 2306 (46). — Po Khant &

D.R. 1106 (23). — Polak, A.M. 1056 (46). — Pongamornkul, W. 501 (27). — Pooma, R.

1006 (48). — Pooma, R. et al. 2299 (44); 2449 (47); 2531 (29); 2731 (43); 6529 (32). —

Poster et al. SAN 139888 (6). — Posthumus, W. 1072 (23); 2085 (18). — Pradit 671 (43).

— Puasa FMS 36752 (32). — Pullen, R. 6642 (36). — Put, N. 523 (3); 2565 (23); 4286?

(23). — Puudyaa, P. 86 (20); 784 (23).

Rahim, A.b.d. SAN 55912 (46). — Ramawoorthy, T.P. HFP-1309 (46). — Ramos, M.

BS 1988 (20); BS 22467 (48): BS 24088 (23).— Ramos, M. & Edano, G. BS 31391 (41);

BS 44610 (24). — Ramos, M. & Pasgasio, J. BS 34758 (41). — Rankin, M.O. 2518 (1).

— Rantai et al. S 68859 (32). — Rechinger, K. & Rechinger, L. 1853 (16). — Richards,

P.W. 2219 (28); 5631 (34). — Ridley, H.N. SF 11956 (46). — Ridsdale, C.E. et al. 2131

(21). Riedel 131 (T)(16b).— Robbins, R.G. 2316 (T)(42). — Robinson, C.B. BS 1352

(23); BS 1353 (48); 1441 (46); BS 9619 (19); BS 9717 (48). — Rock, J.F. 1928 (48); 5318

(49): 10482 (49). — Rodin, R.J. 815 (23). — Romero, E.M. & Chavez, R.V. PPI-29274

(16). — Rostado, E. 13 (6).— Royce, R.D. 7492 (14). — Royen, P. van NGF 15068 (23).

— Ryding, O. 681 (24); 805 (23); 808 (43); 882 (46).

Sachet, M.-H. & Maclet, JN. 1047 (32). — Said, I.M. et al. BRUN 15849 (34). -

Saldanha, C.J. 12464 (46); 16199 (46); 16351 (46); 16421 (46). — Salsedo, C.A. 181

(32).-— Sam, Y.Y. & Mustapa, D. FRI 47157 (6). — Samsuri, A. et al. 240 (32); KJ-18

(32). — Sands, M.J.S. et al. 6544 (2). — Sangkhachand, B. 1518 (46). — Sangkhachand,

B. & Phusomsaeng, S. 1519 (46). — Sangkhachand, B. et al. 1034 (32). — Santisuk, T.

6897 (48): 8617 (23). — Santisuk, T. et al. 438 (48). — Sarif, M. FM 945 (46). — Sasirat, S.

109 (23): 155 (20). — Sato, T. et al. 43 (46). — Scarth-Johnson 1521A (35). — Schneider,

C. 215 (25); 2243 (25); 3833 (25). — Schodde, R. 2481 (32); 2563 (24); 2628 (36); 2664

(20). — Schodde, R. & Craven, L.A. 3967 (30). — Schomburgk, R. herbarium [legit

Schultz] 851 (20). — Seemann, B. 324 (T)(9). — Serre, A. 695 (40). — Setchell, W.A. &

Parks, H.E. 148 (16a).—Shaik Mokim 251 (48).—Shawan,T. SAN 119531 (6).— Shea,

G. SAN 75641 (21). — Shimizu, T. et al. T-18157 (3); T-19201 (33) — Shiwiwat, N. &

Nimanong, B. 35 (48). — Sidiyasa, K. BRF 1958 (21); BRF 1959 (18). — Sidiyasa, K. et

al. 1153 (13); 1155 (32); 1219 (46). — Sigin, G. et al. SAN 68600 (6). — Silva, J.M. de 91

(23). — Sinclair, J. 8910 (46); 10548 (34); 10667 (23).— Sinclair, J. & Kiah bin Salleh SF

39934 (6).—Sirirugsa, P. 862 (24).—Smith, A.C. 191 (32); 1690 (T)(30); 6634 (32); 9211

(32). — Smith, H. 989 (40). — Smith, H.M. 41 (16b); 66 (46). — Smith, L.S. 10210 (32);

11678 (32). — Smitinand, T. 3145 (20); 3690 (20); 4441 (19); 4450 (23); 5521 (27); 8431

(43); 11218 (35). — Smitinand, T. & Ploenchit 99 (46). — Soepadmo, E. 1458 (32). —

Songkhla, B. na 581 (46); 609 (46); KTJ-7 (27); KTJ-32 (27). —S@rensen, T. et al. 1224

A Checklist of Merremia (Convolvulaceae) in Australasia and the Pacific 51

(48).— Specht, R-L. 923 (35): 1064 (16).— Spire. Dr. 1049 (T)(5).— Squires, R.W. 183

(46): 221 (48): 832 (46). — Staples, G. 428 (20). — Staples, G. & Promdej. C. 231 (24):

241 (24): 244 (23): 246 (23): 255 (24): 260 (3): 268 (3). — Staples. G. & Ueachirakan,

W. 185 (48): 186 (24): 195 (24): 202 (32): 211 (48). — Staples. G. & Wathaniyakun. T.

286 (35): 291 (35): 297 (29): 310 (24): 314 (23):316 (19).— Staples, G. & Wongprasert.

T. 135 (3); 142 (3): 144 (3): 165 (3): 168 (46): 173 (24): 175 (24): 225 (24): 328 (23):

332 (24): 339 (24): 340 (20): 345 (44): 347 (23): 348 (24): 350 (46): 352 (46): 357 (24):

358 (3): 365 (43): 371 (35): 372 (43): 394 (46): 396 (48): 403 (43). — Staples, G. et al.

317 (35): 411 (23). — Steenis. CCG. van 2291 (46). — Stevens. PF. et al. 551 (21). —

Stone. B.C. 4722 (23): 5447 (32): 8342 (46): 14332 (48): 15957 (46). — Straatman, R.

BMF-210 (10). — Strachey. R. & Winterbottom, JE. 3 (40). — Streimann, H. NGF

47799 (24). — Stremann, H. & Kairo. A. NGF 44040 (36). — Suddee. S. 158 (47): 212

(44).— Sumbing Jimpin SAN 103523 (18): SAN 103673 (21): SAN 128455 (32): SAN

136703 (21): SAN 139340 (48).— Sumithraarachchi. D.B. & Sumithraarachchi, D. 836

(23).— Supapol, J. 261 (48): 262 (46). — Suvarnakoses, P. 2206 (46). — Suwannaratana,

S. 16 (35).— Symington, CF. FMS 35753 (32).

Taam, Y.-W. 1894 (46). — Takamatsu. M. 828 (32): 921 (32). — Takeuchi. W. 6601 (20):

6726 (32). — Taleon S.H. 101 (6); 55275 (6). — Ten. Pater S. 133 (49): 318 (49): 1419

{under Ducloux number] (49). — Teng. S.-W. 90925 (33). — Tepnarin, P. 349 (46). —

Teruya, Z. 1900 (32).—Teysmann 105 (36). — Th. S. et al. 281 (27). — Thaew 111 (46).

— Thaithong. O. 185 (3).— Thakur Rup Chand 4496 (27). — Thorel 91 (24); 340 (23).

— Toppin, S. 2547 (48). — Topping. D.L. BS 1971 (41). — Tsai, H.-T. 53655 (25): 56218

(T)(49): 60836 (5).—Tsang & Fung 18008 (46).—Tsang. W.-T_ 513 (5): 855 (46): 26848

(27); 27070 (27): 27409 (5): 29207 (5): 29610 (27): 30052 (5): 30070 (27): 30075 (5):

30211 (5): 30453 (23): 30487 (27): 30645 (8). — Tsi. Zhanhuo 92-003 (46). — Tsiang.

Y. 1133 (23); 1487 (23). —Tsinglan. E-H. 83 (20). — Tso, C.-L. 21216 (23): 21816 (46):

23014 (23): 23059 (48).

UNESCO [Kostermans c.s.] 82 (32). — Uong Sing Po [FCU herbarium] 12531 (23):

12580 (23).

Van Niel, J.P. 3769 (34).— Van Ooststroom. S_J. 12535 (24).— Vidal. J.-E. 670 (27):5739

(47); 5813 (23).— Vu Van Cuong 690 (22).

Wallich herbarium catalogue 1343/1 (T)(46): 1343/3 (48): 1348 (48): 1354/1 (1):

1354/2 (T)(1); 1354/4 (1); 1354/b (T)(1): 1377 (16): 1385 (20): 1393 (24): 1398 (T)

(19): 1423/1 (10); 1423/2 (10). — Wallich herbarium catalogue [legit F. de Silva] 1357

(24). — Wallich herbarium catalogue [legit Roxburgh] 1398 (T)(19). — Walsh. M_E.

375 (T)(15). — Wang. C. 32353 (24): 34047 (46): 34104 (46): 35072 (46): 36194 (46).

— Wang. C.-W. 69365 (25): 71521 (25): 80171 (T)(33). — Wang, H. 41417 (25): 41474

(25). — Warburg. O. 10718 (20): 21261 (20): 21263 (32): 21264 (24): 21269 (32). —

Ward, J.V. 103 (32). — Waterhouse, B.M. BMW-5766 (32). — Waterhouse, J.H-L. 13

(30):41B (30).— Weiner, M.A. MW-72-F-124 (9).— Wen. TS. 562 (25).—Wenzel. CA.

640 (32): 1675 (41). — Whistler. A. 6075 (36): 6225 (32): W-2362 (16): W-2374 (32):

W-3118A (32): W-5177 (32). — White, CT. BSIP 123 (30). — White, O.E. 74 (19): 266

516 Gard. Bull. Singapore 61 (2) 2010

(23).— Whitmore, T.C. 6293 (32); BSIP 1613 (7); BSIP 4420 (30). — Whitmore, T.C. &

Sidiyasa, K. TC W-3477 (32). — Wicke (sp.?) herb. 1981 (23). — Wight, R. herbarium

catalogue 1968 (1); 2288 (23); 2374 (23); 2385 (48); Kew distribution no.1984 (46).

— Wilde, W.J.J.O. de & Wilde-Duyfjes, B.E.E. de 19591 (5). — Williams, K. et al. 1474

(48). — Williams, L. 17269 (24). — Williams, R.S. 284 (20); 353 (20); 356 (46); 2033

(20); 2424 (32); 2732 (46); 3000 (24). — Wilson, E.H. 2762 (5); 4183 (T)(49); 4184 (T)

(25). — Womersley, J.S. BSIP 1003 (24); NGF 43702 (36). — Wong, C.C.Y. 470 (23).

— Wongprasert, T. et al. 032-36 (48). — Wood, G.H.S. SAN A4250 (11); SAN 15472

G2).

Yaacob, A. 70568 (46). — Yalin, S. YS 15 (6).— Yates, H.S. 975 (5).— Yen, D.E. 802 (30):

X-73 (32). — Yu, P.C. S. 61358 (6); S. 61408 (46). — Yili, P.C. et al. S.53776 (18); S. 71784

(34). — Yuncker, T.G. 9850 (32); 16001 (16). — Yusuf, R. & Hussain, A. 114 (45). — Yui,

T-P 1177 (25) 2727225); (676. (235) 37810149),

Zimmermann, R. 34 (23); 83 (20); 89 (46). — Zollinger 119 (46). — Zwichey, A.L. 158

G2):

Acknowledgments

This material is based upon work supported by the US National Science

Foundation under Grant No. 0212762. I began the research at Herbarium

Pacificum (BISH); all the staff there are gratefully acknowledged. The

project was brought to a conclusion at Singapore Botanic Garden; I am

grateful to the SBG administration and the National Parks Board for

support. I thank Robert Johnson (BRI) for generously sharing information

about the Australian taxa. Reviews of the manuscript by R. Johnson and D.

F. Austin (ARIZ) are gratefully acknowledged.

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Gardens’ Bulletin Singapore 61 (2): 523-526. 2010 523

Syzygium subscandens (Myrtaceae), a New Species from

Sumatra

P WIDODO ’.T. CHIKMAWATI AND A. HARTANA >

‘Sekolah Pascasarjana Institut Pertanian Bogor, Kampus Dramaga Bogor 16680

* Fakultas Biologi Universitas Jenderal Soedirman.

J] Dr. Soeparno 63 Purwokerto, 53122 INDONESIA.

Email: pwidodo@unsoed.ac.id

* Departemen Biologi, Institut Pertanian Bogor, Kampus Dramaga Bogor 16680

INDONESIA.

Abstract

Anewspecies of Syzygium Gaertn. from Sumatra is described and illustrated:

Syzygium subscandens Widodo. Morphologically the new species 1s similar

to S. garciniifolium, but differs in being a small tree and in having longer

terminal and axillary panicles.

Introduction

Sumatra is one of the largest islands in Indonesia and is one of the richest

in biodiversity (Whitten, 1999), however, it is also one of the least known

flora in the country. Land conversions, the development of industry, illegal

logging, and forest fires have created severe pressure on the biodiversity of

Sumatra.

Species of Syzygium are key components of Southeast Asian and

Sumatran lowland forests. They provide nectar, pollen, and fruits as food for

a wide range of insects, birds and mammals (Parnell er al., 2007). They are

also of economic importance to humans as sources of fruits, timber, tannin

and medicine (Heywood et al., 2007).

The genus Syzygium is largest in the family Myrtaceae, comprising ca

1200 species (Craven et al., 2006), or approximately 1040 species (Govaerts

et al., 2008). The current concept of Syzygium includes species with and

without an intercotyledonary inclusion; the inflorescences are either solitary

or in clusters, either on old trunk, axillary, or terminal; and the calyx may be

either calyptrate or free (Craven et al. 2006).

The new species was identified during preparation of a revision of

Syzygium in Sumatra. To make sure that the new species has not been

named and reported anywhere else outside of Sumatra, we compared it with

specimens of morphologically similar species from the surrounding islands

kept in BO, L and K.

524 Gard. Bull. Singapore 61 (2) 2010

The new species of Syzygium is an important discovery for Sumatra

because it creates awareness and appreciation of the island’s tree biodiversity.

The new information can be used for the development of an effective

conservation strategy.

Syzygium subscandens Widodo, sp. nov.

Frutices subscandentes. Ramuli teretes, luteo-brunnei pallidi. Folia coriacea,

obovata, basietapice obtusa. Inflorescentiae terminales etlaterales, paniculatae;

floribus cruciformibus stellatis, hypanthio obconico-urceolati; corollis albo-

roseis. -Typus: Indonesia. Sumatra WC, west side of Mt. Merapi, 2,300 m

alt., 22 Jun 1953, JV Brossum W 2201 (holotype, BO!; iso, L.). Fig. 1.

Treelets, climbing on tree. Twigs terete, smooth, yellowish. Leaves obovate,

14-18 x 7.5-10.6 cm, drying yellowish brown, lower surface paler; base obtuse,

apex obtuse; petiole thick, not swollen, 1-1.2 cm long; midrib narrowly

furrowed on the upper surface, rounded, raised below; lateral veins 5-8 pairs,

straight or curved, 2-2.8 cm apart, at an angle of 45°-70°; oil dots none or

few; intramarginal vein looped, faint, 5-10 mm from margin. Inflorescence

a panicle, up to 9 cm long, to 9 cm wide, terminal and axillary; peduncle 2-9

mm; ultimate inflorescence axis ca 5 mm long. Flower buds turbinate, white;

pseudostipe sometimes none, hypanthium smooth, obconic to urceolate.

Sepals 4, free, triangular, margin hyaline, 5-8 mm long and wide, pink. Petals

4, free, whitish pink, 4-8 mm long and wide, orbicular or slightly triangular, a

few gland dots per petal. Stamens deciduous, not seen. Style not seen. Fruits

globose ca 5 mm (immature). Seeds solitary, globose, ca 3 mm diameter.

Habitat: Primary forest, climbing on trees.

Distribution: Known only from Mt. Merapi in Sumatra.

Notes: S. subscandens looks like S. garciniifolium. It differs in its habit, being

a small tree or shrub climbing on trees, and in its longer terminal and axillary

panicles. It is characterised also by the thick leathery leaves with very faint

looped intramarginal veins.

Acknowledgements

We would like to thank Dr. Eko B. Walujo (BO), Prof. E. F. Simets (L) ad Dr.

D. Mabberley (K), Herbarium Bogoriense for the permission to examine the

collections and for the use of facilities. Prof. Dr. Mien A. Rifai, Herbarium

Bogoriense, and Mr. Lyn Craven, CSIRO, Canberra, Australia are thanked

for advice, references, and discussion. Prof. Dr. John Parnell, Trinity College,

525

Syzygium subscandens (Myrtaceae), a New Species from Sumatra

Figure 1. Syzygium subscandens. A. Leaty twig with inflorescence; B. Flower; C. Young Fruit.

A,B and C from JV Brossum W No. 2201 (BO). Drawn by Subari.

526 Gard. Bull. Singapore 61 (2) 2010

Dublin, Ireland, is appreciated for references. Subari is also thanked for

drawing the figures. The Directorate General of Higher Education of the

Republic of Indonesia is acknowledged for providing scholarships via IPB

(Bogor Agricultural University).

References

Craven, L.A, E. Biffin and P.S. Ashton. 2006. Acmena, Acmenosperma,

Cleistocalyx, Piliocalyx and Waterhousea formally transferred to

Syzygium (Myrtaceae). Blumea 51: 131.

Govaerts, R., M. Sobral, P.S. Ashton, F. Barrie, B.K. Holst, L.L. Landrum, K.

Matsumoto, FF. Mazine, E.Nic Lughadha, C. Proenga, L.H. Soares-Silva,

P.G. Wilson and E. Lucas. 2008. World Checklist of Myrtaceae. The Board

of Trustees of the Royal Botanic Gardens, Kew.

Heywood V.H., R.K. Brummitt, A. Culham and O. Seberg. 2007. Flowering

plant families of the world. Royal Botanic Gardens, Kew.

Parnell, J.A.N.,L.A.Craven and E. Biffin. 2007. Matters of scale: dealing with

one of the largest genera of angiosperms. In: Hodkinson, T.R. and J.A.N.

Parnell (eds.). Reconstructing the tree of life: taxonomy and systematics

of species rich taxa. Systematic Association Special Volume Series 72: 251-

TT:

Whitten, T. 1999. The Ecology of Sumatra. Tuttle Publishing.

Gardens’ Bulletin Singapore 61 (2): 527-540. 2010 ST

Studies on Schismatoglottideae (Araceae) of Borneo XII:

Three New Species of Schismatoglottis in the Multiflora

Group

S. Y. WONG

Department of Plant Science and Environmental Ecology

Faculty of Resource Science and Technology

Universiti Malaysia Sarawak, 94300 Samarahan, Sarawak, Malaysia

E-mail: sywong@frst.unimas.my

Abstract

Fieldwork since 2002 has revealed three novel taxa of Schismatoglottis in

Sarawak, Malaysian Borneo. Based on the presence of a free ligular portion

to the petiolar sheath, these new taxa belong to the Multiflora Group sensu

Hay and Yuxammi (2000). Here, these species, Schismatoglottis clausula

S.Y.Wong, S. dulosa S.Y.Wong, and S. jitinae S.Y.Wong are described, and

illustrated, and a key to the Multiflora Group in Sarawak is presented.

Introduction

The last revision of Schismatoglottis was that of Hay & Yuzammi (2000), at

which time Schismatoglottis in Malesiastood at 94species,of which 63 occurred

in Borneo. Since then, a further 7 taxa have been published: Schismatoglottis

hayana Bogner & P.C.Boyce, S. jelandii P.C.Boyce & S.Y.Wong, S. jipomii

P.C.Boyce & S.Y.Wong, S. maelii P.C.Boyce & S.Y.Wong, S. ardenii A.Hay, S.

tahubangensis A.Hay & Herscovitch, and S. bulbifera H.Okada, H.Tyukaya

& Y.Mori (see Bogner and Boyce, 2009; Boyce and Wong, 2006; Hay, 2002;

Hay and Herscovitch, 2003; Okada et al., 1999, and Wong and Boyce, 2008).

These taxa are all endemic to Borneo. Here, a further three novel Bornean

endemic taxa, Schismatoglottis clausula S.Y.Wong, S. dulosa S.Y.Wong, and

S. jitinae S.Y.Wong are described. These taxa belong to the Multiflora Group

sensu Hay and Yuzammi (2000).

Multiflora Group

Based on fieldwork observations, and of plants in cultivation in the Botanical

Research Centre, Semenggoh (BRC), Schismatoglottis pudenda A.Hay

does not belongs to the Multiflora Group. The foliage leaves of S. pudenda

528 Gard. Bull. Singapore 61 (2) 2010

alternate with cataphylls, and have the petiolar sheath very much reduced,

with the role of protecting the emerging leaf/shoot/inflorescence taken over

by the subtending cataphyll, therefore, this species belongs to the Tecturata

Group. A separate paper (Boyce and Wong, in prep.) will address this and

other changes in the Tecturata Group. Based on molecular work on two

plastid regions on the Schisamatoglottid Alliance (Tribe Schismatoglottideae

+ Cryptocoryneae + Philonotieae) (Wong, ef al., in press), Schismatoglottis

josefit A.Hay and S. sarikeensis (Bogner & Hotta) Bogner & Hay do not

belong in Schismatoglottis, but rather in one of the ‘satellite’ genera, possibly

in a redefined Hottarum Bogner & Nicolson. Therefore, S. josefii and S.

sarikeensis are here excluded, These changes mean that currently there are

thirteen species in the Multiflora Group in Sarawak.

Key to Schismatoglottis Multiflora Group in Sarawak

1. Petiolar sheath fully attached to the petiole, either long or extremely

abbreviated (sometimes with a short apical ligulate portion) ...............

seb lage hay el as pee ee RR ee Schismatoglottis excl. Multiflora Group

1. Petiolar sheath attached to the petiole only at the base, the remainder

free-ligulate and comprising at least half of the entire sheath...

Eee es PE A to 2. Schismatoglottis Multiflora Group

2. stem elongated, erect, sometimes ascemdimpence...- scene ee 3

2. Stem condensed, sometimes more-or-less creeping but never erect ..... 4

3. Plant medium; petiolar sheath marcescent; petiole asperulate ; lamina

softly leathery, matte at least adaxially; peduncle stout and short, not

elongated; lower spathe succulent; female zone almost as wide as long;

appendix present. Never cocOpiyile ccc... ee ee eee S. jitinae

3. Plant small; petiolar sheath persistent; petiole glabrous; lamina thinly

leathery, glossy at least adaxially; peduncle slender, elongated during

anthesis; lower spathe softly coriaceous; female zone longer than wide;

appendix absent, Always mheop lytic yeesccscenccceers oecee ee ree ee S. erecta

4, Lamina corrugated adaxially; primary and interprimary veins prominently

raised adaxially;spathe limb green at anthesis; inflorescence strongly fruity-

smelling at anthesis «2.08 eee eee S. maelii

4. Lamina never corrugated; primary and interprimary veins smooth

adaxially; spathe limb always white at anthesis; inflorescences usually

smelling powerfully eSterie at AmtMesisss-2.ctesesereesectesccenscate ee sus encnsee eee 5

: = En : E a9

Studies on Schismatoglottideae (Araceae) of Borneo XII 529

5. Appendix absent or very reduced (comprising just a few terminal

PLETE TEE ceceece benedeni eee 6

pe = SDP ELE CEPEE STD Ds Ea nee ee 10

G2 OE SISTED SSL ea eee en eee 7

2B LEE REE DT AU] TG sit CA ee 8

7. Lamina abaxially not glaucous, base obtuse to slightly decurrent,secondary

and tertiary venation obscure: pore at edge of stamen, punctiform (c. 0.15

mm) with protruding tissue from innermost surface, connective elevated.

male zone remained white basally and stained brown distally in alcohol.

Lithophytic on limestones, Bau, Kuching Division ................. S. bauensis

7. Lamina abaxially glaucous, leaf base always decurrent, secondary

venation prominent, tertiary venation sometimes pronouncedly

tessellate; pore at inner surface of stamen, oblong (ca 0.3 mm), connective

flat, male zone completely stained brown in alcohol. Terrestrial in deep

soil on limestones and sandstones, Padawan, Kuching Division and Serian,

5 EES DS Se ee S. confinis

8. Spathe inflated at anthesis; spadix with a blunt end ..... 9

8. Spathe unrolling slightly at anthesis but not inflated: spadix with a

WL) SURE SE eR ee S. multiflora

9. Leaf adaxially very glossy: primary lateral veins are flush adaxially: spathe

caducous, pink at anthesis. Rejang Valley .............0.....-+- S. roseospatha

9. Leaf adaxially slightly glossy: primary lateral veins are sunken adaxially:

spathe Marcescent, white at anthesis. Sematan ..............000...0.. S. hayana

10. Plant large, up to ca 1 m tall; lamina to ca 40 cm long: placenta 1 per ovary:

_ oT ES DES ee S. monoplacenta

10. Plant medium, less than 60 cm tall, lamina less than 30 cm long: placenta

more than | per ovary: West Sarawak, never on limestone .................. 11

2 LAE TE DYES Se So SE Sa OE ne RSE 12

Sarat ae INNA eRe FS ah td tee ee 13

12. Spathe limb disarticulating into numerous circumferential rings along the

length, and then collapses downwards: appendix thicker than the male

zone: sterile interstice wider than the distal portion of female zone, and

ie en hs MMs Pree ees SESS eA NaS ei iE te S. clausula

12. Spathe limb remaining erect marcescent in one piece:appendix contiguous

from the male zone: sterile interstice contiguous with the distal portion

530 Gard. Bull. Singapore 61 (2) 2010

13.Spadix usually ca 8 cm long, witha short sterile interstice of sterile stamens

between the fertile zones; stamens and appendical staminodes large, ca

1. mm-actoss; Matamg 222 eeecce eee eee eee ee S. mayoana

13. Spadix usually ca 4.5 cm long, with the fertile zones contiguous; stamens

and appendical staminodes small, ca 0.3 mm across; Bako ...........cceceeeeees

Schismatoglottis clausula S.Y.Wong, sp. nov.

Ab Schismatoglottis mayoana (in sylvis mons Matangae restrictus) spathae

laminorum tempore anthesis clausus, lamina senescens per disarticulatis

circumferentialis numerosis, mox collabens inde marcescens remanens

postanthesin; staminis parva (0.2-0.3 mm), cum margini thecae elevates

connectivo antherae superantis, fovea latus profunde cingens differt — Typus:

Malaysia, Sarawak, Sarikei, Ulu Sarikei, 01° 55’ 05.4”; 111° 29’ 35.8”, 7 Dec

2005, P.C. Boyce et al. AR-1582 (holo, SAR). Fig. 1.

Herbs to ca 30 cm tall. Stem pleionanthic, condensed, more or less creeping,

ca 2 cm diam., green. Leaves ca 5-7 together; petiole terete, 18-26 cm long,

densely scabrid, green, sheathing only at the extreme base, the sheath

extended into a bicarinate narrowly lanceolate free ligular portion to ca 10

cm long, drying red-brown; laminae lanceolate to oblongo-ovate, ca 15-25 cm

long x 4-9 cm wide, matte dark green adaxially, abaxially paler, drying brown,

base obtuse, sometimes slightly decurrent, tip acute and acuminate for up to

ca 2 cm and with a cylindric mucro to 2 cm long; midrib grooved adaxially

and abaxially very prominent (dry) with 14-16 primary lateral veins on each

side alternating with fine interprimaries and diverging at ca 60°; secondary

venation fine and very dense, adaxially obscure, mostly arising from the

midrib; tertiary venation obscure. Inflorescences up to 2 together, pendent,

ca 8-10 cm long, subtended by cataphylls resembling the ligular leaf sheaths;

peduncle less than the length of the free ligules. Spathe 8-10 cm long; lower

spathe obliquely inserted from peduncle, ca 2.4 cm long, thickly coriaceous,

green, differentiated from the limb by a weak constriction corresponding

to the distal portion of the female zone; spathe limb 5-6 cm long, white,

oblongo-lanceolate, apiculate for 3 mm, barely open during anthesis, senesce

by disarticulating into numerous circumferential rings along the length, and

this then collapsing downwards, and remaining marcescent post anthesis.

Spadix sessile, 5.5-6.0 cm long, subcylindric; female zone 2.2-2.4 cm long x

0.5 cm proximally (0.4 cm distally) diam., adnate to the spathe in the lower

4, somewhat conic in the free part; pistils many and crowded, more or less

ie |

Studies on Schismatoglottideae (Araceae) of Borneo XII 2)) l

Figure 1. Schismatoglottis clausula $.Y.Wong. A. A pair of inflorescences. B & C. Inflores-

cence at female anthesis. D. Spathe limb senescense by disarticulating into numerous circum-

ferential rings along the length; E. female zone (spathe removed artificially); F. Sterile interstice,

male zone and appendix at female anthesis; G. Spadix at the end of male anthesis; H & I. Spathe

limb marcescent.

Dis Gard. Bull. Singapore 61 (2) 2010

globose, ca 0.4 mm diam., green when fresh; stigma sessile, button-like, ca

0.3 mm diam., turn orange post female anthesis; interpistillar staminodes

confined to a basal row along the spathe/spadix adnation, shortly stalked,

clavate, round-topped, ca 0.5 mm wide; sterile interstice present, above

the spathe constriction, ca 0.3-0.5 cm long x 0.6 cm diam., wider than the

distal portion of the female zone and the male zone, crowded, irregularly

polygonal, flat-topped, 0.5 — 0.6 mm diam., white prior to anthesis, greyish

orange post anthesis; male zone cylindrical, 1.1 cm long x 0.5 cm diam.;

stamens crowded, small, ca 0.2-0.3 mm diam., truncate, dumbbell-shaped,

white when fresh, turn orange post male anthesis, connective narrow; thecae

with slightly elevated rims (overtopping the connective) surrounding a deep

broad pit-like pore; appendix ca 2 cm long, slightly wider than the top of

the male zone, in the distal half tapering to an acute point; staminodes of

appendix flat-topped, irregularly polygonal, small, ca 0.3 mm diam., white

when fresh, turn yellow post male anthesis. Infructescence unknown.

Distribution: Malaysia, endemic to Sarawak, known only from the type

locality.

Habitat: Lithophytic in shade of primary lowland dipterocarp forest on

shales, 60 m asl.

Etymology: The specific epithet is derived from the Latin clausus meaning

closed, referring to the spathe limb barely opening at anthesis.

Notes: Schismatoglottis clausula is most similar to Schismatoglottis mayoana

Bogner & M.Hotta but is readily distinguished by the senescence mechanics

of the spathe limb. The spathe limb senesces by disarticulating into numerous

circumferential rings along the length, and then collapses downwards, while

remaining marcescent post anthesis. The stamen is small (0.2-0.3 mm), with

thecae rims slightly elevated and overtopping the connective, these rims each

surrounding a deep, broad, pit-like pore. The interstice staminodes become

markedly elevated prior to male anthesis. This latter character has so far only

been observed in S. mayoana.

Schismatoglottis dulosa S.Y.Wong, sp. nov.

Ab Schismatoglottis multiflora similis, combinatio appendice spadicis,

inflorescentis erectis et spathae laminorum marcescenti remanens, non

reflexis postanthesis, distinguitur — Typus: Malaysia, Sarawak, Kuching, Bau,

Kampung Jugan, 01° 28’ 46.4”; 110° 05’ 08.5”, 26 Mar 2004, P.-C. Boyce &

Jeland ak Kisai AR-279 (holo, SAR). Fig. 2.

Studies on Schismatoglottideae (Araceae) of Borneo XII BS i

Herbs to ca 35 cm tall. Stem pleionanthic, condensed, more or less creeping,

ca 2 cm diam. Leaves ca 8-12 together; petiole terete, 20-30 cm long, always

tinged reddish towards the base, glabrous, sheathing only at the extreme

base, the sheath extended into a bicarinate narrowly lanceolate free ligular

portion to ca 12 cm long, drying red—brown; laminae broadly lanceolate,

softly coriaceous, ca 15-25 cm long x 4-9 cm wide, dull dark green adaxially,

abaxially paler and glaucous when fresh, drying brown, base always obtuse,

tip acute and acuminate for 2 cm and with a cylindric mucro to 2 cm long:

midrib adaxially grooved and abaxially very prominent (dry) with 14-16

primary lateral veins on each side alternating with fine interprimaries and

diverging at ca 60°;secondary venation fine and very dense, adaxially obscure,

mostly arising from the midrib; tertiary venation tessellate but obscure.

Inflorescences up to 3 together, up to ca 13 cm long, emerging and remaining

erect throughout anthesis, subtended by cataphylls resembling the ligular

sheath extensions; peduncle less than the length of the free ligules. Spathe

ca 13 cm long, lower spathe obliquely inserted from peduncle, ca 4 cm long,

thickly coriaceous, pale green at base, whitish towards the distal portion,

differentiated from the limb by a weak constriction corresponding with the

top portion of the female zone; spathe limb ca 9 cm long, white, oblongo-

lanceolate, apiculate for 5 mm, forming a hook, remaining marcescent, not

reflexing, turning brownish red. Spadix sessile, ca 10 cm long, subcylindric;

female zone ca 4 cm long x 1 cm proximally (0.6 cm distally) diam., adnate

to the spathe in the lower 2,somewhat conic in the free part: pistils crowded,

more or less globose, ca 0.5 mm diam., yellowish green when fresh; stigma

sessile, overtopping the ovary, pappilate, ca 0.6 mm diam.; interpistillar

staminodes present, scattering, shortly stalked, clavate, round—topped, ca

0.4 mm wide, white when fresh; sterile interstice present, monometric with

female and male zone, cylindrical, ca 0.5 cm long x 0.6 cm diam., white

when fresh; male zone cylindric, ca 2.3 cm long x 0.6 cm diam.; stamens laxly

arranged, truncate, more or less rectangular, ca 0.7 mm long x 0.5 mm wide,

pale orange when fresh, with the connective wide and about the height of the

thecae, the thecae at the short ends, each with one pore: appendix contiguous

from the male zone, ca 4 cm long, in the distal half tapering to an acute

point; staminodes of appendix flat-topped, cylindrical, white when fresh,

turn yellow post male anthesis, ca 1.0 mm diam. Infructescence unknown.

Distribution: Malaysia, endemic to Sarawak, known only from the type

locality.

Habitat: Lithophytic in shade of lowland forest on limestone, 50 m asl.

Gard. Bull. Singapore 61 (2) 2010

‘

Figure 2. Schismatoglottis dulosa $.Y.Wong. Adaxial (A) and abaxial (B) side of lamina; C.

Petiole with marcescent free petiolar sheath; D. An inflorescence at male anthesis with two

emerging inflorescences; E & F. Female zone at female anthesis; G. Male zone; H. Spathe

limb marcescent; I. Spadix at male anthesis.

Studies on Schismatoglottideae (Araceae) of Borneo XII 535

Etymology: The specific epithet is derived from the Latin du/lon meaning

enslaving, referring to the spathe limb remaining marcescent and enclosing

the spadix post anthesis.

Notes: Schismatoglottis dulosa is vegetatively most similar to Schismatoglottis

multiflora Ridl.except an appendixis presentin S. dulosa(absentin S. multiflora),

and the inflorescence remains erect in S. dulosa (declinate by flexing of the

peduncle in S. multiflora). Schismatoglottis dulosa is further distinguished by

the spathe limb marcescent, not reflexing and falling caducous post anthesis

as occurs in S. multiflora.

Schismatoglottis jitinae S.Y.Wong, sp. nov.

Inter alti specibus Schismatoglottidorum grex Multiflorae per combinatio caulis

elongatis, erectis ascendentis, cum inflorescentia femina et inforescentia mascula

in partem conubialiter intra spatham inferiorem contentis cum constricta

spathae supra media Schismatoglottis jitinae proprius. Ab S. erectus ligulis

vaginae petiolaris marcescentis et laminis foliorum lenis coriacea, impolitus,

abaxaliter pro totam longitidinem, et basali (adaxaliter) roseus, pedunculo

valde brevis et crassus, erectus, differ; spathae fructiferorum succulentus differt.

— Typus: Malaysia, Sarawak, Kapit, Pelagus Rapids, 02° 11° 15.1”; 113° 03°

29.01”, 14 Mar 2005, P.C. Boyce et al. AR-1038 (holo, SAR). Fig. 3.

Herbs to ca 40 cm tall. Stem pleionanthic, elongated, erect, sometimes

ascending, ca 2-2.5 cm diam.. Leaves ca 12-15 together; petiole short, 10-15 cm

long, terete, always D-shaped towards the base, asperulate, pink in new leaf,

sometimes remain dark red at base, puberulent, sheathing only at the extreme

base, the sheath extended into a bicarinate narrowly lanceolate free ligular

portion to ca 12 cm long, drying red—brown; laminae lanceolate to obovate,

ca 18-25 cm long x 4-9 cm wide, softly leathery, matte green adaxially, paler

and glaucous abaxially, base decurrent, always unequal, the tip acute then

acuminate for 2 cm and with a cylindric mucro to 2 cm long; midrib adaxially

grooved, always pink during innovation at the base (adaxially) and at the

whole length (abaxially), primary lateral veins very prominent abaxially, 14-

16 on each side alternating with fine interprimaries and diverging at ca 60°;

secondary venation fine and very dense, adaxially obscure, mostly arising

from the midrib; tertiary venation obscure. Inflorescences solitary, up to ca 8

cm long, emerging erect, remains throughout anthesis, peduncle short, ca 2.5

cm, stout, subtended by cataphylls resembling the ligular sheath extensions,

always pink. Spathe ca 5.0 cm long; lower spathe, obliquely inserted from

peduncle, ca 2.3 cm long, succulent, pale green at base, whitish towards

the distal, differentiated from the limb by a weak constriction coinciding

with the male zone; spathe limb ca 2.7 cm long, oblongo-lanceolate, white,

Gard. Bull. Singapore 61 (2) 2010

Figure 3. Schismatoglottis jitinae S.Y.Wong. A. Whole plant; B. Emerging inflorescence; C.

Inflorescence at female anthesis; D. Spathe limb caducous in small pieces; E. Female zone

(spathe removed artificially); F. Spadix (spathe removed artificially).

Studies on Schismatoglottideae (Araceae) of Borneo XII 537

apiculate for 1 mm, caducous, fragmenting into small pieces. Spadix sessile,

ca 4.5 cm long, subcylindric; female zone ca 1.3 cm long x 1.2 cm proximally

(0.7 cm distally) diam., adnate to the spathe in the lower 2/3, somewhat conic

in the free part; pistils crowded, more or less globose, yellowish green when

fresh, ca 0.5-0.7 mm diam.; stigma sessile, button-like, pappilate, overtopping

ovary; interpistillar staminodes confined to the basal of female zone, a few,

shortly stalked, clavate, round-topped, ca 0.5 mm wide, white when fresh;

sterile interstice present, ca 0.4 mm long x 0.7 cm wide, monometric with

the female and male zone, white when fresh; male zone cylindric, ca 1.4 cm

long x 0.6 cm diam.; stamens packed, truncate, dumbbell shaped, ca 0.5 mm

long x 0.3 mm wide, white when fresh, with the connective wide and taller

than the thecae, the thecae at the short ends, each with one pore; appendix

contiguous from the male zone, ca 1.7 cm long, in the distal half tapering

to a blunt point; staminodes of appendix flat-topped, irregularly polygonal,

small, white when fresh, ca 0.3 mm diam. Infructescence unknown.

Distribution: Malaysia, endemic to Sarawak, Kapit Division.

Habitat: Mesophytic in steep gallery forest on shales, 50-430 m asl.

Etymology: The specific epithet is for Madam Jitin ak. Ojek, one of the

nursery staff at Malesiana Tropicals Sdn. Bhd., and who formerly assisted

with the maintenance of the Araceae research collection.

Notes: Schismatoglottis jitinae is distinctive by its elongated and erect aerial

stem, the leaf laminae basally attenuate and usually unequal, and softly leathery,

with the midribs of the innovations longitudinally bright pink abaxially and

basally adaxially. Cataphylles are also always pink.

In having erect, elongated stems, S. jitinae approaches S. erects M.Hotta,

from which it differs by the short and stout peduncle not elongating during

anthesis, the succulent lower spathe and the spathe constriction coinciding

with the middle part of the male zone.

Other specimens seen: MALAYSIA. Sarawak: Kapit Division, Nanga Gaat,

Rejang Wood Concession, Sungai Bereng, 01° 45’ 36.0”; 113° 27° 54.7”, 15

Dec 2004, P.-C. Boyce et al. AR-891 (SAR); Pelagus, Pelagus rapids, 02° 11°

15.1”; 113° 03’ 29.01”, 14 Mar 2005, P.-C. Boyce et al. AR-1039 (SAR); Kapit,

Taman Rekreasi Sebabai, 01° 56’ 45.6”; 112° 54’ 16.8”, 19 Apr 2006, P.-C.

Boyce et al. AR-1785 (SAR).

538 Gard. Bull. Singapore 61 (2) 2010

Acknowledgements

This study is funded by the Ministry of Higher Education, Malaysia

Fundamental Research Grant Scheme No. FRGS/01(04)/609/2006(42) under

Sarawak Forestry Department Research Permit Nos. NPW.907.4.2(1)-101

& NCCD.907.4(1V)-41 & Park Permit Nos. 58/2007 & 37/2009. The

collaboration and support of the Sarawak Forestry Department, the Forest

Research Centre (Kuching), The Semenggoh Botanical Research Centre,

and the Sarawak Biodiversity Centre are gratefully acknowledged. Thanks

are due to Peter Boyce for translating the Latin diagnoses.

References

Bogner, J.and P.C. Boyce. 2009. Studies on the Schismatoglottideae (Araceae)

of Borneo VI: A new Schismatoglottis species from Sarawak, Malaysian

Borneo. Gardens’ Bulletin Singapore 60 (2): 1-9.

Boyce, P.C. and Wong, S. Y. 2006. Studies on Schismatoglottideae (Araceae)

of Borneo I: A trio of new Schismatoglottis from Sarawak, Borneo.

Gardens’ Bulletin Singapore 58(1): 7-18.

Hay, A. 2002. A new Bornean species of Schismatoglottis (Araceae).

Aroideana 25: 67-69.

Hay,A.and C. Herscovitch. 2003 A new species of Schismatoglottis (Araceae)

from Sabah. Gardens’ Bulletin Singapore 55: 27-30.

Hay, A. and Yuzammi 2000. Schismatoglottideae (Araceae) in Malesia I -

Schismatoglottis. Telopea 9 (A): 1-177.

Okada, H., H. Tsukaya and Y. Mori. 1999. A new species of Schismatoglottis

(Schismatoglottidinae, Araceae) from West Kalimantan and observations

of its peculiar bulbil development. Systematic Botany 24 (1): 62-68.

Wong, S.Y. and P.C. Boyce, 2008. Studies on Schismatoglottideae (Araceae)

of Borneo II: Schismatoglottis confinis, a Putative Sister Taxon to

Schismatoglottis bauensis from Sarawak, Malaysian Borneo. Gardens’

Bulletin Singapore 60(1): 155-163.

Wong S.Y., P.C. Boyce, A. Sofiman, Othman and C.L. Pin. (2010). Molecular

phylogeny of tribe Schismatoglottideae based on two plastid markers

Studies on Schismatoglottideae (Araceae) of Borneo XII 539

and recognition of a new tribe, Philonotieae, from the neotropics. Jaxon

59: 117-124.

Gardens’ Bulletin Singapore 61 (2): 541-548. 2010 541

Studies on Schismatoglottideae (Araceae) of Borneo X.

Pichinia, a New Genus from Sarawak, Malaysian Borneo

S.Y.WONG ” AND PC. BOYCE’

‘Department of Plant Science and Environmental Ecology, Faculty of Resource

Science and Technology, Universiti Malaysia Sarawak, 94300 Samarahan,

Sarawak, Malaysia.

‘ Corresponding author e-mail: sywong@frst.unimas.my

_ School of Biological Sciences, Universiti Sains Malaysia, 11800 Penang, Malaysia

“Forest Herbarium (BKF), The Office of Forest and Plant Conservation Research,

National Park, Wildlife and Plant Conservation Department,

61 Phahonyothin Road Chatuchak, Bangkok, 10900 Thailand

Abstract

Pichinia S.Y. Wong & P.C. Boyce is described as a new genus from Sarawak,

Malaysian Borneo with one species, Pichinia disticha S.Y. Wong & P.C.

Boyce. This genus is, so far, known only from the type locality, Pichin,

Serian, Bahagian Kuching, Sarawak. The genus is illustated and a key to the

Schismatoglottideae is presented.

Introduction

Tribe Schismatoglottideae comprises Schismatoglottis Zoll. & Moritzi (with

probably in excess of 150 species), and six mono- to oligospecific ‘satellite’

genera: Aridarum Ridl., Bakoa P.C. Boyce & S.Y. Wong, Bucephalandra

Schott, Phymatarum M.Hotta, Piptospatha N.E. Br. and Schottarum P.C.

Boyce & S.Y. Wong (Bogner and Hay, 2000; Boyce and Wong, 2008). The

overwhelming majority of Schismatoglottis species, and all the satellite

genera except Piptospatha, are endemic to Borneo. Initially, the new species

described here was tentatively placed as a sp. nov. in Schismatoglottis,

although the unique shoot architecture and basal placentation are anomalous.

However, subsequently molecular analyses based on two plastid markers

by the first author (Wong et al., in review) revealed that the species to be

well-supported as a taxon distinct from to Schismatoglottis sensu Hay &

Yuzammi, the latter being a grossly polyphyletic assemblage. We are here

describing this taxon as a new genus, Pichinia S.Y. Wong & P.C. Boyce, basal

to Schismatoglottis, plus the satellite genera with the exception of Schottarum

P.C. Boyce & S.Y. Wong.

542

Gard. Bull. Singapore 61 (2) 2010

Key to genera of Schismatoglottideae and their principle subgeneric

divisions in Borneo, Jawa and Nusa Tenggara

Cn GN

Wings of petiolar sheath fully or almost fully attached to the petiole;

seeds never. with a micropylar appendage -.--..-<. ee 2

Wings of petiolar sheath extended into a free ligular portion; seeds

sometimes with micropylamappemdage mr ciccccee eee eee 3)

Inflorescences on very slender peduncles, nodding at anthesis, peduncle at

spathe insertion flexing 180° from vertical axis. Infructescences narrowly

campanulate, noddime, Plamts/Of poGSols 22.--...--2.s--<seeceeeee sere Hestia

Inflorescences erect to nodding at anthesis, if nodding, then either

peduncle massive, and peduncle at spathe insertion at most 45° from

vertical axis. Infructescences fusiform with a constricted orifice, if

campanulate, then thick-walled and erect, never nodding. Plants of

Vanious substrates Dut ME Ver On POUSOIS =e eee ee 4

Modules monoeuphyllous, congested in a distichous arrangement; ligular

SHC ath Persistent 2. cecc.ceccerare cece cence tees eee eee Pichinia

Modules polyeuphyllous and leaves never distichous; ligular sheath

When presemt mManrCeSCeml 22. -cceieccesecs sense nes coe reneeeceeers eens ees eee 3)

Spathe limb persistent into fruiting. Petiolar sheath usually fully

deciduous; spadix interstice always present, invariably at least partly

naked .:.An0G eRe a Ee ee ees Apoballis

Spathe limb deciduous during anthesis, or marcescent. Petiolar sheath

persistent or marcescent;interstice where present always fully clothed with

sterile MOWEES 2.2 eee ee 12 Schismatoglottis

Spathe mot: constricted: yx0rce Aedes fan eee cee oct ee rten reece areas teeta eee 6

Spathe comstricted) c.:5......8.. Jcastbes Se easeseeeen etoenn nee toteeoec op eee tev eeee ete -cemeeeeneeeme 11

Thecae of anther never with horn- or needle-like projections ............... 7

Thecae of anther each with a horn- or needle-like projection, although

these sometimes visible only after female anthesis «0.0.0.0... ce eeeeeeeee 8

Spadix almost completely adnate to spathe; male flowers mostly sterile

with a narrow zone of fertile flowers exposed by the spathe opening;

peduncle declinate during fruit maturation but twisting to become

semi-erect at fruit maturity; spathe persistent into fruiting, and then at

fruit maturity swiftly drying, reflexing and opening basally by tearing at

Studies on Schismatoglottideae (Araceae) of Borneo X 543

peduncle insertion to expose fruits but remaining distally convolute and

while in this situation clasping the spadix. Seeds with blunt micropyle ......

Ine eee. 2 gE 5 288s va ndlsceasl ducucead eateabicbhtaseneal vivian Mee Bakoa

7. Spadix either entirely free or only part of the female flower zone which is

adnate to spathe; male flowers all fertile; peduncle erect (and then spathe

limb caducous) or declinate (and spathe persistent) throughout the fruit

dispersal; spathe limb either caducous early in anthesis or persistent until

fruit maturity and then falling, still fresh to reveal entire spadix and ripe

fruits. Seeds with a pronounced, hooked, micropylar appendage ..............

ee Na Eeinseln cies serine nace onechssnsensennssondeu’ Piptospatha

8. Thecae with needle-like projection extending only after female anthesis;

projection tipped with a weakly peltate ovate-triangular flap. Appendix

MAREE SS ALO NPIS EO ACS oe tess ches coc vcerstnsabenoddoonsenedebarensebecee Schottarum

8. Thecae with a horn- or needle-like projection present prior to female

anthesis; with the projection pointed and never associated with a terminal

flap. Appendix, where present, composed of staminodes ............e eee )

9. Sterile interstice of spadix with flattened scale-like staminodes; anthers not

Se ReRO AR fe see a, BF. as, sd occencanscdedasenendivencens Bucephalandra

9. Sterile interstice absent or with truncate staminodes; anthers nearly

always with the top excavated (except A. INCAVALUM) .......ceceeeseeseeeeee 10

iesnecae at cach end of the anther (seen from aDOVE) .......:....c...2ss.seseacersesees

Ria A ET 6 cd sonani vee Aridarum Sect. Aridarum

10. Thecae adjacent on one side of the anther (seen from above) .............06

ar Ns oo oooh csevaesdvonssanevaces Aridarum Sect. Caulescentia

11.Thecae of anther without horn- or needle-like projections; ovules on

parietal placenta; seeds without a micropylar appendage ............. eee

ee ee Ey eGo casa stedanevecvese Schismatoglottis Multiflora Group

11. Thecae of anther, each with horn- or needle-like projections; ovules on

basal placanta;seeds with along, hooked micropylar appendage ................

Sea eg eS ec Sass Cavcet viv dan dsdonaonttcsleeaxeSavaxst Phymatarum

ae MRM MA EN ANNU eee IE soso suene cohen sweet dsnnsnv se vos tick dnesnsuaidnatecasenwetdscnnnce 13

IPP Stem Bap ax amie «...sc.0c2cccccence0sss 40s Schismatoglottis Calyptrata Group

13. Petiole sheathing only at extreme base; each foliage leaf alternating with

ZV G1 0) 1h | eee Schismatoglottis Tecturata Group

13.Petiole usually sheathing for at least a third of its length (rarely less);

foliage leaves not alternating with cataphylls:........08.c.c.c.ceccsssseedoncces 14

544 Gard. Bull. Singapore 61 (2) 2010

14. Inflorescence erect; spathe limb irregularly crumbling and breaking away

ator after male anthesis; smalltomedium plants .:2-s-ersee.soeeeseeeeee ee

Se ape rR ero drier ce ascnbeaGocsc Schismatoglottis Asperata Group

14. Inflorescence nodding; spathe limb clasping the spadix and more or

less marcescent after anthesis, finally falling with spent parts of spadix;

MASSIVE Pachy Cause Schismatoglottis Corneri Group

Pichinia S.Y. Wong & P.C. Boyce, gen. nov.

Herba lithophytica, caulis porrecto vel decumbens. Folii plura, in modulis

monophyllorum distichus congestis, petioli vagina long persistens.

Inflorescentia 1, erecta sub anthesis; pedunculus quam petiolo valde brevior;

spatha erecta leniter constricta ad medium; spadix ad spatham basaliter

adnatus, parte femina densiflora, parte mascula densiflora, staminodia ad

basim et apicem habens, baccae carnosae densiter dispositae in spathae

fructiferorum anguste-campanulatum persistens; flores masculi 2-andricis;

flores feminei ovarium ovoideum, 2-loculare, ovulum plura ad basim loculi

insertum. — Typus: Pichinia disticha S.Y. Wong & P.C. Boyce, sp. nov.

Lithophytic mesophytes. Stem creeping to erect. Leaves many together,

distichously arranged, each module with prophyll, cataphyll (both long

persistent, as long as petiolar sheath) and single foliage leaf; petiole D-shaped,

puberulent, petiolar sheath with a long- persistent free ligular portion; lamina

oblonceolate, orthotropic to petiole, basally cuneate, apex acuminate with

tubular mucro, this short,and soon marcescent and brown, softly chartaceous,

adaxially matter pale olive-green, abaxially glaucous; primary lateral veins

prominent, pinnate, adaxially sunken, abaxially prominent, secondary

venation pinnate, running parallel to primary veins, tertiary venation obscure.

Inflorescence solitary, erect; peduncle terete, shorter than petiole; spathe up

to 8 cm long; lower spathe ellipsoid, strongly oblique at insertion of spadix/

peduncle, sometimes with a ventral triangular gap formed during female

anthesis and then closed prior to male anthesis, inflating at female anthesis

but remaining constricted at top, lower spathe orifice gaping slightly at male

anthesis once spathe limb shed; constriction between upper and lower spathe

weakly defined; spathe limb caducous in a single piece; spadix often slightly

exceeding spathe at anthesis, or at least equalling, conico-cylindric; female

flower zone cylindrical, attenuate distally; pistils ellipsoid-cylindrical, white;

stigma overtopping ovary, translucent white; ovary incompletely 2-locular,

placenta basal, ovules several, micropylar appendage absent; interpistillar

staminodes absent; interstice present, wider than female flower zone, with

pistillodes proximally, these smaller than pistil, and staminodes distally,

these very similar to stamens; male flower zone short; stamens irregular,

Studies on Schismatoglottideae (Araceae) of Borneo X 54

crowded, whitish yellow; anthers 2 per flower, oblong to somewhat irregular

in shape; pores apical, pollen extruded in strings; appendix present, conic,

distally, staminodes crowded, irregular. Infructescence with lower spathe,

narrowly campanulate, persistent; fruits berry, crowded, oblongo-globose;

seeds ellipsoid, weakly longitudinally striate.

Distribution: Malaysia, Sarawak, Samarahan, Serian, Pichin.

Habitat: Perhumid evergreen forest on limestone, 100-250 m asl.

Notes: The shoot architecture of P disticha is unique in_ the

Schismatoglottideae. Although plants are polyphyllous, the individual

modules are monoeuphyllous, comprising a prophyll, a cataphyll, both long-

persistent, and a foliage leaf (euphyll). This basic model is similar to the

module architecture of Schimatoglottis tecturata (Schismatoglottis Tecturata

Group). However, unlike the Tecturata Group, the petiolar sheath in

Pichinia is elongated in the form of a greatly lengthened persistent ligule,

which would seem to have some protective role associated with the emerging

shoots (as compared with the Tecturata Group where the petiolar sheath is

greatly reduced to a minute ridge at the petiole base and the protective role

of the sheath is homeotically taken by the cataphylls.

The distichous arrangement appears to favour litter-trapping

ability, the plants growing horizontally out from vertical or near-vertical

rock surfaces. The leaf posture is typically with the lamina orthotropic to

the petiole in nature. Similar gross morphology and associated ecology is

found in some Homalomena species, notably H. geniculata M. Hotta and H.

crassinervia Ridl.

Occasionally the spathe has a triangular-shaped opening ventrally

during female anthesis and which closes prior to the onset of male anthesis;

it is not observed elsewhere in the tribe. Nothing is currently known about

the pollination ecology of Pichinia.

Etymology: This genus is named after the type location, Pichin, Serian,

Samarahan, Sarawak, Malaysia. This place is remarkable for the forested

limestones that have been well preserved by the local community.

Pichinia disticha S.Y. Wong & P.C. Boyce, sp. nov.

Ab alii Schismatoglottidorum folii plura in modulis monophyllii distichus

congestis et petioli vagina long persistens differt. Ad generis certeris

Schismatogottideae (speciebus Schismatoglotti et Hestii excludens) spatha

erecta leniter constricta ad medium baccae carnosae densiter dispositae in

spathe erecta anguste-campanulatus persistens, seminae sine appendice

546 Gard. Bull. Singapore 61 (2) 2010

microphylorum et ovulum plura ad basim loculi distinguitur. — Typus:

Malaysia, Sarawak, Samarahan, Serian, Pichin, Gunung Kedadum, Sugun

Karang, 01° 06° 17.6”; 110° 29° 04.5”, 29 Jun 2006. P.C. Boyce, Simon Kutuh

ak Paru & Wong Sin Yeng AR-1860 (holotypus, SAR, + spirit). Fig. 1.

Mesophytic lithophyte up to 40 cm tall. Stem creeping to erect, up to 10 x

2 cm diam. Leaves many, distichously arranged, each module with prophyll,

a cataphyll (both long persistent), up to 7 cm long, and a single foliage leaf;

petiole D-shaped, puberulent, up to 12 x 1 cm diam., petiolar sheath with

a long persistent free ligular portion up to 7 cm long; lamina oblonceolate,

up to 25 x 9 cm, softly chartaceous, base ovate to cuneate, apex acuminate,

with a short tubular mucro, up to 2 mm long, this soon marcescent, brown;

lamina matte pale olive-green , abaxially greyish glaucous; primary lateral

venation pinnate, up to 11 per side, prominent abaxially, sunken adaxially,

interprimary venation barely distinguished from primaries; secondary

venation pinnate, running parallel with primary veins, tertiary venation

obscure. Inflorescence solitary, erect, smelling weakly esteric at anthesis;

peduncle terete, pale green, up to 8 cm long; spathe up to 8 cm long; lower

spathe mid green with darker longitudinal veins, ca 3 cm long, strongly

oblique at insertion, with a ventral triangular gap sometimes forming during

female anthesis and closing prior to male anthesis, barely constricted in

between upper and lower spathe, spathe limb broadly triangular, ca 5 x 3

cm, white, caducous in a single piece, in interior surface at first shiny, then

degrading into raised scales prior to falling; spadix exceeding or at least

equalling spathe, ca 7 cm x 8 mm, conoid-cylindrical; female flower zone

cylindrical, attenuate distally, ca 3 cm long; ovary ellipsoid-cylindrical, pale

green; stigma overtopping ovary, translucent white; interpistillodes absent;

interstice present, wider than female flower zone, with pistillodes proximally,

these smaller than pistil, light orange, and staminodes distally, these similar

to stamens, whitish yellow; male flower zone short, ca | cm long; stamens

irregular, crowded, whitish yellow, ca 2 mm diam., pores apical, pollen

extruded in strings; appendix ca_ 1.5 cm long, tapering-conic, appendiccal

staminodes crowded, irregular, pale whitish yellow.

Distribution: Known only from the type locality and nearby forested

limestone hills.

Habitat: Lithophytic mesophytes on exposed limestone or earth banks in

perhumid limestone forest, 100-250 m alt.

Etymology: The specific epithet is for the distichous arrangement of the

leaf.

Studies on Schismatoglottideae (Araceae) of Borneo X 547

Figure 1. Pichinia disticha S.Y.Wong & P.C.Boyce. A. Plant in habitat, note the distichous leat

arrangement; B. Plant in habitat showing the leaf lamina glaucous abaxially and persistent

free-ligular petiolar sheaths; C. Cultivated plant showing detail of the leaf arrangement. The

white arrow indicates a petiolar sheath; the red arrow indicates a prophyll; D. Inflorescence

at onset of male anthesis, note the spathe limb caducous in a single piece and the pollen

extruding in strings; E. Inflorescence very late female anthesis, note gaping orifice to the

lower spathe; F. Spadix at early male anthesis with spathe artificially removed; G. Young

infructescences showing the open orifice to the persistent lower spathe; the spathes limbs

in both examples have failed to fall after being shed due to dry air. All photos based on

P.C.Boyce et al. AR-1860 (SAR).

548 Gard. Bull. Singapore 61 (2) 2010

Other specimens seen: MALAYSIA. Sarawak, Samarahan Divison: Serian,

Pichin, Tubih, Tahang Sipukam, 01° 07° 16.6”; 110° 26’ 51.2”, 26 Jul 2005, P.-C.

Boyce et al. AR-1402 (SAR); Pichin, Sugun Serabu, 10 Oct 2005, P.-C. Boyce

& Simon Kutuh ak Paru AR 1476, (SAR); Pichin, Gunung Kedadum, Sugun

Karang, 01° 06° 17.6”; 110° 29° 04.5”, 7 Apr 2006, P.C. Boyce, Simon Kutuh

ak Paru & Wong Sin Yeng AR-1761 (SAR).

Acknowledgements

This study is funded by the Ministry of Higher Education, Malaysia by

fundamental research grant scheme No. FRGS/01(04)/609/2006(42) under

Sarawak Forestry Department Research Permit No. NPW.907.4.2(1)-101 &

Park Permit No. 58/20076. The collaboration and support of the Sarawak

Forestry Department, the Forest Research Centre (Kuching), and the

Sarawak Biodiversity Centre, are gratefully acknowledged.

References

- Bogner, J. and A. Hay. 2000. Schismatoglottideae (Araceae) in Malesia 2:

Aridarum, Bucephalandra, Phymatarum and Piptospatha. Telopea 9(1):

179-222.

Boyce, P.C. and S.Y. Wong. 2008. Studies on Schismatoglottideae (Araceae)

of Borneo VII: Schottarum and Bakoa, two new genera from Sarawak,

Malaysian Borneo. Botanical Studies 49: 393-404.

Gardens’ Bulletin Singapore 61 (2): 549. 2010 549

Book Review: Tan, Benito C. and Ho Boon-Chuan. 2008. A Guide to

the Mosses of Singapore. Photographs by Lim Yao-Hui. Science Centre

Singapore. 149 pp. Price: SGD$ 5.50.

Finally it is here! After many months of waiting, I have received my copy of

the book pent by post from a good friend in Singapore. It arrived yesterday

and I have not stopped until I read it all. It is a great complement to my

other bryology books and the first to cover tropical mosses. Co-authored by

the world renown bryologist Dr. B.C. Tan, it expertly covers the species of

mosses found in Singapore. And yet it is written in simple flowing English

and accompanied by a wealth of clear colour photographs which will be

enjoyed by both newcomers to moss biology and more seasoned bryologists

alike. The first few pages are in fact dedicated to newcomers in moss biology

with a description on morphology, life cycles and general information about

these tiny green plants.

Over 150 species of indigenous mosses have been recorded in Singapore

of which two are endemic to the island at present. The book contains a

checklist of all these species with useful comments on habitat, distribution

and survival status. As can be expected of a small book, not all of these

mosses are described in detail. However over 56% of the book is dedicated

to portraits of selected species. In this section the more common species are

described in detail, accompanied by photographs of habit and morphology,

which would allow the amateur bryologist to make an identification even in

the field. Selected rare and unusual species are also described here in detail

to make this a good cross section of the species to be found in the region. To

make this chapter even more enjoyable the descriptions include the various

uses of these mosses. This means that enthusiasts from the aquarium hobby,

the bonsai hobby, and traditional medicine have been offered a good fodder

for their interests.

The habitat and locations in Singapore have been also been listed

for each species, which will help amateur bryologists find them easily. They

will also find the keys to the families and genera of Singapore mosses in the

book useful. If needed, there is a short description on how to collect and

preserve mosses for later identification or study by specialists working at the

herbarium and plant museum overseas. Finally the glossary complements

the other sections by serving as a quick reference to the technical terms

used in the book.

The Guide to the Mosses of Singapore serves as both a wonderful

armchair read as well as a practical field guide. My only problem with it is

that it has aggravated my desire to come over to do moss “hunting” in the

island country called Singapore.

Stephan Mifsud

Malta, Europe

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Gardens’ Bulletin Singapore 61 (2): 551-552. 2010 >

Book Review: Heide-Jorgensen, H S. 2008. Parasitic Flowering Plants. Brill

NV, Leiden. 438 pp., numerous color photographs. ISBN 978 90 04 16750 6.

Price: € 99/ US $147.

It is a pleasure to review such a beautiful book as this. The author has clearly

crafted a labor of love, based on decades of study and research, followed by

years of effort to write, design, and illustrate it. The book brings together an

enormous amount of information about the biology, physiology, taxonomy,

life history, and systematics (classification) of all flowering plants that are

wholly or partly parasitic by nature. Their global distribution is mapped by

family, which is a very handy feature.

In total, the author estimates that there are approximately 4,490

species of parasitic plants known, of which ca 4,100 species are hemiparasites

(partly self-sustaining through photosynthesis) and only 390 species are

holoparasites (entirely dependent on a host plant for survival). This total

represents less than 1% of the estimated number of flowering plant species

on earth. While the book does not present any figures for how many species

are covered, an impressive diversity of them has been described and

illustrated.

Virtually every page of text is enlivened by color photographs, most

are of good to high quality, photomicrographs, schematic diagrams, and line

art. The list of photo acknowledgments extends to 4 full pages; the amount of

effort the author must have expended to track down and obtain permission

to use the hundreds of color images in the book can scarcely be imagined.

Only someone passionate about parasitic plants could have seen through to

the end the task of searching for, and obtaining permission to use, so many

fine color images, with such admirable results. Readers interested to know

more about a particular image will find the full contact details provided

for the photographers, a welcome touch. I note, in passing, that among the

photographers is Singaporean Joseph T. K. Lai, and readers can expect to

find photos in the book that were taken in Singapore.

Quibbles: there are a few. The running head on every one of the 438

pages states the title of the book, Parasitic Flowering Plants, but the reader

has no idea in which of the 8 chapters or 43 subchapters he/she is in on any of

those pages. In an information-dense and copiously illustrated tome such as

this, finding one’s way around within the book can be tricky, especially given

the number of cases where illustrations cited in the text appear elsewhere

in the book. It would have been far more user-friendly to use the running

heads to indicate the chapters and subchapters to ease internal navigation.

Like many books that are edited by the author, there are some quirks

and oddities of a very minor nature that creep into the text. Another pair of

eyes is always beneficial in catching these little gremlins and removing them.

352. Gard. Bull. Singapore 61 (2) 2010

For example, the state of New Jersey (in the USA) has consistently been

rendered as “New Yersey” everywhere it occurs in text and photo captions.

On the whole, however, typos are remarkably few and the text reads very

well.

The price is off-putting: like most books published by scientific and

technical publishers, the price tag will deter many people from buying

this magnificent book. It is to be hoped that Brill will produce a soft cover

edition at a more economical price. Libraries, scientific institutes, and the

serious student of parasitic plants will buy the hardcover edition, but the

general public probably will not, purely for cost consideration. And that is

a pity, because there is so much information contained in the book that will

fascinate, educate, and delight anyone with an interest in the natural world.

It is to be hoped that Singaporeans, and others in Southeast Asia generally,

will splurge and indulge themselves by buying this book — it has much to

offer. And all of the parasitic flowering plants native in our part of the world

— Rafflesia, Striga, Balanophora, even a species of Lepionurus — are to be

found within its covers, which brings the information in this book very close

to home.

I can wholeheartedly offer my sincere congratulations to the author

and publisher for bringing this comprehensive and beautiful work into

existence; I believe it will be the standard of excellence on the subject of

parasitic plants for decades to come.

George Staples

Senior Researcher

The Herbarium

Singapore Botanic Gardens

Gardens’ Bulletin Singapore 61 (2): 553-554. 2010 553

Book review: Fernando E.S., Min Hwan Suh, Jaeho Lee and Don Koo Lee.

2008. Forest Formations of the Philippines. ASEAN-Korea Environmental

Cooperation Unit (AKECU), Korea, 232 pp. ISBN 978-89-92239-40-0

93530.

This is a beautiful pictorial guidebook produced under the joint research

program of ASEAN-Korea_ Environmental Cooperation — project

(AKECOP). The book illustrates the various types of forest formations

found in the Philippines today with minimal text but heavily elucidated

with hundreds of colour images of the forest structures and representative

plant species. According to the book, the aim is, “to help increase awareness,

understanding, and appreciation of the rich biodiversity of the Philippine

forests, and promote their conservation and sustainable use.” Indeed,

considering that the country today has less than 10% of the original forest

cover, the publication is a timely reminder and serves to educate the young

generation of Filipinos and people from the region what the country looked

like once upon a time with its extensive and diverse forest formations.

The book starts with a short and succinct introduction that includes a

terse description of the physical geography, geological origin and vegetation

history of the island groups. It shows in a tabulated form the 12 types of

natural primary forest vegetation still found in the country and their

differences in soil water requirement, location, soil type, and elevation range.

The 12 forest types described and illustrated in the book follow the forest

classification proposed by Whitmore in 1984 in his book entitled, * Tropical

Rain Forests of the Far East” (Oxford Press). They are: (1) Tropical lowland

evergreen rain forest, (2) Tropical lower montane rain forest, (3) Tropical

upper montane rain forest, (4) Tropical subalpine forest, (5) Forest over

limestone, (6) Forest over ultramafic rocks, (7) Beach forest, (8) Mangrove

forest, (9) Peat swamp forest, (10) Fresh water swamp forest, (11) Tropical

semi-evergreen rain forest, and (12) Tropical moist deciduous forest. The

book concludes with a short glossary of technical terms, the Literature

Cited, an Acknowledgements and an Index.

Separate illustrative maps showing the limited range and disjunctive

location of the limestone forest formation, the ultramafic forest formation,

peat swamp forest formation and the fresh water swamp forest formation in

the Philippines are provided. It would be more useful and educational had

the authors of the book also provided up-to-date distribution maps for all

other forest formations today.

No doubt, this pictorial book, with its many colourful photos, has

made the reading and understanding of the usually dry subject matter of

forest vegetation types easy and enjoyable.

What struck me most are the three chapters reporting on the ultramafic

Gard. Bull. Singapore 61 (2) 2010

forest, peat swamp forest and fresh water swamp forest formations in the

Philippines. My own knowledge about them in the Philippines has been

enriched after reading the book. To my knowledge, this book is the first

to document in writing and show with photos the existence of peat swamp

forest in the Philippine archipelago! I can see from the photos presented that

the peat swamp forest discovered recently in southern Philippines does have

a similar plant community structure and harbors the same dominant plant

species found in the peat swamp forest in Borneo. As these last mentioned

three types of forest formations are also not well known to many in the

neighboring countries, the book will be useful for any library in the region

to possess.

My main “complaint” about the book is the uneven quality of

sharpness of the colored photos printed in the book ranging from good to

excellent. A cursory read of the book will reveal some photos that appear

dull in color separation and not very clear in image outline, while others are

printed in brilliant, colourful texture to show details in good contrast. Some

of the well composed and reproduced photos of individual plant species

seen in the book that merit a special mention are Monophyllaea merrilliana

on p. 93, Xanthostemon fruticosus on p. 102, Bruguiera gymnorhiza on p.

143, B. sexangula on p. 144, Nypa fruticans on p. 160, Medinilla teysmanii

on p. 179 and Symphorema luzonicum on p. 210. As several of the plant

species photographed and shown in the book are the uncommon, rare and

endemic plant taxa in Philippines, the book has an added value in providing

a pictorial aid to the identification of these little known species.

My other criticism against the book is the very small print of the photo

credits of the photographers in the acknowledgement.

I like to recommend this pictorial book to everyone who has an interest

in knowing and learning about the different types of forest formation in the

region, not just in the Philippines. It is a great pleasure and deep intellectual

satisfaction to read this book and appreciate the beauty of the plant world,

as well as to imbibe the rich information of the types of forest formation

conveyed by the book. In fact, I will recommend the book to every school

and university, research institutions and government offices across the

country, and in the region too, as an important reference.

The authors are to be congratulated for a pictorial book well prepared

to create public awareness on an important topic of conservation today —

the tropical rainforest vegetation.

B.C. Tan

The Herbarium

Singapore Botanic Garden

INSTRUCTIONS TO AUTHORS

The Gardens’ Bulletin publishes original research findings and reviews of progress in the fields of plant

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Authors should look at the layout of articles recently published in the journal to ensure that submitted manuscripts

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references. Manuscripts may be submitted in electronic form (PC-compatible WORD) together with a hardcopy

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given below the title. A short running title should also be provided. Lengthy papers must have contents listed at

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Abstract: An abstract of 100 to 200 words should be provided. It should comprehensively summarise the

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Literature citation: Citation in the text should take the form: King and Gamble (1886), or (King and Gamble,

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2000a, 2000b, etc. When papers are by three or more authors they should be cited as King er al. (1886) in the

text, but with all the authors’ names given in the reference section. All references must be placed in alphabetic

order according to the family name of the first author. The journal title must be given in full, as in the following

examples:

Stone, B.C. 1994. Additional notes on the genus Glycosmis (Rutaceae). Gardens’ Bulletin Singapore 46:

113-119.

Kress, W.J.,L.M. Prince, and K. J. Williams. 2002. The phylogeny and a new classification of the gingers

(Zingiberaceae): evidence from molecular data. American Journal of Botany 89:1682-1696.

References to books and monographs should be cited according to the following form:

Ridley. H.N. 1930. The Dispersal of Plants Throughout the World. L. Reeve, Ashford, U.K.

For literature citations in taxonomic papers, the main standards are Stafleu & Cowan, Taxonomic Literature,

ed. 2, Regnum Vegetabile, Utrecht, for abbreviated names of books, and Botanico-Periodicum-Huntianum

(B-P-H), Pittsburgh for abbreviated names of periodicals. The following style is required:

Medinilla alternifolia Blume, Mus. Bot. Ludg.-Bat. 1:1 (1849) 19.

Sterculia acuminatissima Merr., Philipp. J. Sci. 21 (1922) 524.

Offprints: Authors will be given 50 offprints gratis. Additional capies must be ordered and paid for in advance

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