THE GARDENS’ BULLETIN, SINGAPORE

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conservation, with emphasis on the plant life of the Southeast Asian-Pacific region.

Dr David J. Middleton Dr Jana Leong-Skomickova

(Editor-in-Chief) (Managing Editor)

S. Lee Y.W. Low Christina Soh

(Graphics Editor) (Copy Editor) (Business Manager)

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Professor Sir Peter Crane

Yale University

USA.

Dr Rogier P.J. de Kok

Royal Botanic Gardens, Kew

UK.

Dr W. John Kress

National Museum of Natwal History

Smithsonian Institution

U.S.A.

Dr Mark Hughes

Royal Botanic Garden Edinburgh

UK.

Dr Kiat W. Tan

Gardens By The Bay

Singapore

Dr Nigel P. Taylor

Singapore Botanic Gardens

National Parks Board, Singapore

Dr Ian M. Turner

Royal Botanic Gardens Kew

U.K

Dr Jan-Frits Veldkamp

National Biodiversity Center

The Netherlands

Dr Jun Wen

National Museum of Natural History

Smithsonian Institution

U.S.A.

Professor Nianhe Xia

South China Institute of Botany

PR. China

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References listed at the end. There, works mentioned in the text are listed alphabetically as follows:

Dallwitz, M.J., Paine, T.A. & Zurcher, E.J, (1999). User’s Guide to the DEETA Editor, http://

biodiversity.uno.edu/delta/ (accessed on 2 Aug. 2010).

Persson, C. (2000). Phylogeny of Gardenieae (Rubiaceae) based on chloroplast DNA sequences from the

rps 16 nitron and P‘«L(UAA)-F(GAA) intergenic spacer. Nordic J. Bot. 20: 257-269.

Ridley, H.N. (1930). The Dispersal of Plants Throughout the World. Ashford, U.K.: L. Reeve.

Smith, A.C. & Daiwin, S.P. (1988). Rubiaceae. In: Smith, A.C. (ed) Flora Viliensis Nova, A New Flora

q/T/// 4: 143-193.

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acknowledged and the studies for which the sequences were generated should be cited.

Style of nomenclatural summaries. The following style is required:

Ornithoboea arachnoidea (Diels) Craib, Notes Roy. Bot. Gard. Edinburgh 11: 251 (1920); Burtt, Notes

Roy, Bot. Gard. Edinburgh 22: 294 (1958).

Ornithoboea parishii C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 148 (1883).

If authors include full bibliographic data for these works in the list of references at the end of the paper,

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Front cover picture: Hanguana rubinea (Photo by Jana Leong-Skomickova)

The Gardens’ Bulletin

Singapore

VOL. 67(1) 2015 ISSN 0374-7859

CONTENTS

J. Leong-§kornickova & P.C. Boyce

Hanguana in Singapore demystified: an overview

with descriptions of three new species and a new record 1

M. Rodda, D. Lee & S. Mishak

Dischidia acutifolia (Apocynaceae, Asclepiadoideae)

a new record for the Singapore flora 29

I.M. Turner

Yvom Ancistwcladus to Tristaniopsis via Tetramerista

- the convoluted history of a Wallich collection and its impact

on the native flora of Singapore 35

S. Lindsay & D.J. Middleton

The clarification and synonymisation of two taxa of Vittaria

from Peninsular Malaysia and a new combination

in Haplopteris (Pteridaceae subfam. Vittarioideae) 39

A.R. Rafidah, S. Syahida-Emiza, R. Kiew & K. Imin

Bothriospermum (Boraginaceae), a new generic record

for the Flora of Peninsular Malaysia 45

A. A. Joffre, A.K. Muhammad Ariffin, Y.W. Low & K.M. Wong

Novitates Bmneienses, 1 . A background to the botanical survey of Brunei

Darussalam, and a new species of Jarandersonia (Malvaceae) 51

Y.W. Low, A.A. Joffre & A.K. Muhammad Ariffin

Novitates Bmneienses, 2. A remarkable new species oi Begonia

sect. Petermannia (Begoniaceae) from Bmnei Damssalam

K. M. Wong & Y.W. Low

Novitates Bruneienses, 3. Eight new woody plants

in the Brunei flora, including five new species 69

S. Rahayu, U. Meve & M. Rodda

Hoya undulata (Apocynaceae, Asclepiadoideae), a new myrmecophytic

species from Borneo, and typification of H. darwinii 85

N.K.E. Undaharta, I Made Ardaka, Agung Kurniawan & Bayu Adjie

Begon ia bimaensis, a new species of Begonia

from Sumbawa Island, Indonesia 95

C. Puglisi, S. Suddee, P. Triboun & D.J. Middleton

Anew species of Paraboea (Gesneriaceae) from Thailand 101

H. Kurzweil & Saw Lwin

New orchid records for Myanmar, including

the first record of the genus Stereosandra 107

M.F. Newman

A new species of Zingiber (Zingiberaceae) from Lao P.D.R 123

L. Bai, J. Leong-Skornickova & N.H. Xia

Taxonomic studies on Zingiber (Zingiberaceae) in China I: Zingiber kerrii

and the synonymy of Z. menghaiense and Z. stipitatum 129

C. Deori, N. Odyuo & A.A. Mao

Pennilabium labanyaeanum (Orchidaceae),

a new species from Meghalaya, Northeast India 143

K. Thoiba, A.K. Pradeep & C.N. Sunil

Tripogon bimucronatus (Poaceae: Chloridoideae: Tripogoninae),

a new species from India I5I

K. Bransgrove & D.J. Middleton

A revision of Epithema (Gesneriaceae) 159

B.R. Maslin

Synoptic overview of Acacia sensu lato (Leguminosae: Mimosoideae)

in East and Southeast Asia

231

BOOK REVIEW

Flowering Plants of the Western Ghats, Volume 1 (Dicots)

T.S. Nayar, A. Rasiya Beegam & M. Sibi. 2014 & Flowering Plants

of the Western Ghats, Volume 2 (Monocots). T.S. Nayar, M. Sibi

& A. Rasiya Beegam. 2014. (J. Leong-Skomickova)

Date of publication: 4 June 2015

National Parks Board

Singapore Botanic Gardens

1 Cluny Road

Singapore 259569

251

Printed by Oxford Graphic Printers Pte Ltd

Gardens’ Bulletin Singapore 67(1): 1-28. 2015

doi: 10.3850/S2382581215000010

Hanguana in Singapore demystified: an overview with

descriptions of three new species and a new record

J. Leong-Skomickova^ & P.C. Boyce^

^Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, 259569, Singapore

jana_skomickova@nparks.gov.sg

^Institute of Biodiversity and Environmental Conservation (IBEC),

Universiti Malaysia Sarawak, 94300 Samarahan, Sarawak, Malaysia

phymatarum@gmail.com

ABSTRACT. The genus Hanguana (Hanguanaceae - Commelinales) was until recently

believed to be represented in Singapore by a single species, Hanguana malayana. Recent

extensive surveys, coupled with the detailed study of fresh and herbarium material, however,

suggests the presence of six species. In addition to the recently described Hanguana neglecta,

three additional species, H. corneri, H. rubinea and H. triangulata, are described here as new to

science. Of the tliree newly described species, Hanguana rubinea and//, triangulata are native,

while H. corneri is believed to have been introduced from Peninsular Malaysia to Singapore

and planted in Bukit Timah Nature Reserve in the 1930s. Hanguana nitens is recorded for

the first time fi*om Singapore and is also certainly native. Wliile a large helophytic species of

Hanguana, currently interpreted to be Hanguana malayana, is cultivated as an ornamental

plant in Singapore, no native populations have been observed by us in the wild and neither do

any herbarium records exist to suggest that this species was ever native to Singapore. Colour

plates and a key to all Hanguana species are provided. Notes on habitats and preliminary lUCN

assessments are also included for all native species to better facilitate conservation efforts of

Hanguanaceae in Singapore. The existence and taxonomic potential of scale structures sheathing

the inner staminodes in female flowers, named here as staminodial scales, are highlighted here

for the first time. The nnportance of seed characters is also discussed.

Keywords. Bukit Timah Nature Reserve, Central Catchment Nature Reserve, Hanguana

corneri, H. malayana, H. neglecta, H. nitens, H. rubinea, H. triangulata, seeds, staminodial

scales

Introduction

Hanguanaceae (Commelinales) is a small monogeneric family distributed in S and

SE Asia, with the highest diversity in the Sunda region. The family is perhaps the

most understudied family of monocots (Siti Nurfazilah et al, 2011), deserving much

closer attention in the region. Currently 1 1 species are recognised of which eight were

described in the last five years (Siti Nurfazilah et al., 2010, 201 1; Mohd Fahini et al.,

2012; Niissalo et al., 2014). We, however, estimate that the number of species is likely

to exceed 50 as future exploration and detailed studies, particularly of mature fruiting

material, progress.

Gard. Bull. Singapore 67(1) 2015

While Singapore has the best-known flora in SE Asia owing to the small size of

the country, coupled with long-tenn botanical exploration (Niissalo et ah, 2014), the

state of Icnowledge is still far from complete. Numerous new records (e.g,, Low et ah,

2014; Rodda et ah, 2012), as well as descriptions of new taxa (Sugumaran & Wong,

2014; Leong-Skomickova et ah, 2014), including Hanguana neglecta (Niissalo et ah,

2014), have recently been published from Singapore.

All existing herbarium records of Hanguana at SfNG, collected in Singapore

between 1885 and 2009 (but excluding our recent collections) and amounting to

20 specimens in all, were previously misidentified as Hanguana malayana (Jack.)

Men*., a name much misapplied across Asia. As such, Hanguana malayana is the only

species listed in works dealing with the Singapore flora (Keng et ah, 1998; Chong et

ah, 2009). During extensive fieldwork since 201 0 by the first author, it was observed

that several Hanguana species are present in Singapore's forests rather than just the

single species previously suggested. Further fieldwork, coupled with the expertise of

the second author who has been working on the Hanguanaceae of the Sunda region

since 2010, revealed that at least six species are present in Singapore, although only

four are considered native. These include Hanguana neglecta recently described from

Singapore (Niissalo et ah, 2014), H. nitens a species recently described from Johor,

Peninsular Malaysia (Siti Nurfazilah et ah, 2010), and two new species described

below as H. mbinea and H. triangulata. There is no confirmed record of Hanguana

malayana ifom the wild, be it by direct observation, or through the existence of even

a single herbarium specimen from Singapore. Hanguana corneri, while described

here as new to science, has been introduced to Bukit Timah Nature Reserve, and is not

considered a Singaporean native.

The difficulty of studying species of Hanguana from herbarium material alone,

and the importance of infmctescence structure, stigma, fruits, seeds and indumentum

for delimitation of taxa, has been highlighted before by Siti Nurfazilah et ah (2010,

2011). The importance of the morphology of the stigma cannot be overstated as it

preserves well in herbarium material. The position of the stigma in relation to

fruit, however, still requires further study. We have observed that in some species

(e.g., Hanguana mbinea and H. triangulata^ both described below), the position

of the stigma on ripe fruit (oblique or central) is influenced by the number of seeds

developing in the berry. Hanguana berries are trilocular, with a single ovule in each

locule, of which one, two or all three can develop. The actual number of seeds which

eventually develops presumably depends on pollen availability, but also on species

strategy. While the number of seeds which develop per berry is more or less stable in

some species (e.g., always single mH. neglecta), it can vary in others (e.g., one or two

in H. nitens, H. mbinea and H. triangulata', one, two or thi'ee in H. malayana). While

seeds were previously acknowledged as a potential source of characters for Hanguana

taxonomy, the topic has not been discussed in detail. Descriptions or illustrations of

seeds are imfoitimately not available for many of the recently described species. Our

current study shows that seeds are extremely informative and this topic is, therefore,

discussed below. We also highlight the existence of previously overlooked structures

occurring in female flowers.

Hanguana in Singapore

The taxonomic importance of staminodial scales and seeds

Tillich (1996), Tillich & Sill (1999) and Rudall et al. (1999) made major contributions

to our understanding of the floral and seed structure of Hanguana. This then served as

a basis for recent descriptions and was further improved with the recent work of Siti

Nurfazilah et al. (2010, 2011) and Mohd Falimi et al. (2012).

Hanguana flowers are rather unifonn across the entire genus and seem to offer

few characters helpful in species delimitation. Male flowering material is rarely found

in the field (and, therefore, is also scarce in herbaria). The male flowers/inflorescences

are short-lived in comparison to female flowers, which progress to fruiting stage (a

process lasting several months) while retaining all floral parts practically unchanged or

slightly enlarged (e.g., tepals, staminodes, stigma). As fruits offer additional characters,

particularly the colour of the ripe fruit and seed morphology, Hanguana species are

most often described from fmiting material, a trend certain to be continued.

While examining fresh fruiting material of various species from Singapore,

the presence of scale-like stmctures sheathing the base of the three inner staminodes

was noticed. None of the previous publications dealing with Hanguana mentions

their existence (e.g. Airy Shaw, 1980; Tillich, 1996; Bayer et al., 1998; Tillich & Sill,

1999; Siti Nurfazilah et al., 2010, 2011), with the exception of Rudall et al. (1999)

who noticed them but misinterpreted them for inner staminodes. Further detailed

examinations of fresh as well as herbarium material, however, confirmed that six

staminodes are present in each flower (three outer and three umer) and that the three

scales are positioned at the base of the three imier staminodes, sheathing them partially

of fiilly. These scales differ in shape and in the presence of a hyaline margin (Fig. 1,

3B, 5B, 7B, 9B, 12B). They preserve well in herbarium specimens and are, therefore,

of good taxonomic potential, at least in certain species. While the exact origin and

function of these scales is not yet clear and requires further study, it has been suggested

(H.-J. Tillich, pers. comm.) that these might be a basal outgrowth of the staminodes,

possibly exuding a mucilage or a particular scent to attract a pollinator. We propose

here to use the term stammodial scales for these structures. Although staminodial

scales have been observed by us in all species we have so far examined from living

and herbarium material, it is not yet clear if these are indeed present in all Hanguana

species.

The seeds of Hanguana are unique in the monocots (H.-J. Tillich, pers. comm.).

They are more or less bowl-shaped with the hilum positioned at the base of the bowl,

the cavity being filled up by a placental tissue. Although the importance of seed size, the

varying depth of cavity and the degree of incurving of the rim were briefly mentioned

by Siti Nurfazilah et al. (2011), descriptions of seeds were unfortunately not provided

by Siti Nurfazilah et al. (2010) for most of the recently described species {Hanguana

exultans, H. pantinensis, H podzol icola, H stenopoda) or the large helophytic species

currently interpreted as Hanguana malayana.

From our recent work it is obvious that the morphological range of the seed

shape, and therefore their taxonomic potential, is considerable. The recently described

Hanguana neglecta (Niissalo et al., 2014) exhibits seeds which are almost round to

Gard. Bull. Singapore 67(1) 2015

Fig. 1. Detail of staminodial scales at the base of inner staminodes. A. Hanguana triangulata

Skomick. & P.C.Boyce. Note the presence of a hyaline margin on the staminodial scale. B.

Hanguana malayana (Jack.) Merr. Note the staminodial scale has no hyaline margin and is

composed of lobes. (Photos: Jana Leong-Skomickova)

Hanguana in Singapore

ovoid with a wedge-shaped ostiole. Similar seeds are also observed in Hanguana corneri

which is described below. The seeds of Hanguana rubinea and H. triangulata, also

described below, although both bowl-shaped, exhibit differently shaped appendages

along the proximal part of the rim. As previously mentioned, Hanguana species are

mostly collected in the fruiting stage, and fruits are often present in ample numbers

on the herbarium collections. Seeds, including small structures like appendages,

preserve well in dried material, as long as the fruits are collected fully mature or close

to maturity. It is possible to study even old material, from which the seeds can be

carefully extricated by removing the layer of fruit pulp after the berries are soaked

overnight in water at room temperature. In this study, seeds from specimens as old

as 1 889 collected by H.N. Ridley, were successfully extracted from fruits, providing

valuable confimiation of the determinations of Hanguana triangula ta and H. neglecta.

Hanguana in Singapore

The following account presents an identification key and colour plates to all six Han-

guana species so far known to be present in Singapore, including introduced and

cultivated species. Full descriptions are provided for the new taxa, while recently

described species are provided with notes to address any additions or deviations from

original or recent descriptions, these being clearly referenced. Based on sterile living

material we have observed in the field, we suspect the existence of at least one addi-

tional species, although field studies are needed to resolve its identity.

The tenninology used in the descriptions follows Beentje (2012) and the most

recent Hanguana works cited above. The type material of the species described from

ripe fruiting material also includes spirit material of fruits as well as extracted seeds.

Preliminary lUCN conservation assessments followed Davison et al. (2008) and

lUCN guidelines (lUCN, 2012) for local and global assessments respectively. Geo-

CAT (Bachman et al. 2011) was used to calculate EOO.

The native species are treated first in this account, followed by introduced and

cultivated species.

Key to Hanguana in Singapore

la. Large stoloniferous colonial herbs 2

lb. Solitary or clumping herbs lacking stolons 3

2a. Leaves stiffly erect with acute apex; lamina more or less flat or weakly irregularly

corrugate, semi-matt green; staminodial scales composed of lobes without hyaline

margin; stigma lobes large, flat, coimate at base, forming a bluntly triangular to

clover-leaf shape, almost obscuring the apex of ovary

H. malayana (sensu Siti Nurfazilah et al. 2010)

Card. Bull. Singapore 67(1) 2015

2b. Leaves weakly arching, with long-attenuate apex; lamina prominently corrugate,

shiny green with visible pattern of lighter and darker green (best observed on

young and medium aged leaves); staminodial scales entire with hyaline margin;

stigma lobes small, erect, separate, tear-shaped to obovate

H. nitens

3 a. Leaves green on both sides 4

3b. Leaves dark emerald-green above and dark red-purple underneath ....//. corneri

4a. Large herbs over 1 m in height; leaves arching; ripe fruits cream-white or ruby-

red; seeds bowl-shaped, more or less hemispherical 5

4b. Medium sized herbs not exceeding 0. 8 m in height; leaves spreading (not arching);

ripe fruits black; seeds y4 globose to ovoid with wedge-shaped opening

H. neglecta

5a. Lamina almost flat, abaxially with evenly distributed silky indumentum; ripe

fruits ruby-red; stigma lobes connate basally (sometimes imperfectly), with round

apices, foiming bluntly triangular structure; seed appendage acute

H, nibinea

5b. Lamina more or less conxigate, abaxially with unevenly distributed flocculose

indumentum; ripe fruits cream-white; stigma lobes with shaiply acute apices,

connate basally (always perfectly), fomiing sharply triangular structure; seed

appendage bluntly bilobed H. triangulata

Species native to Singapore

Hanguana neglecta Skomick. & Niissalo, Phytotaxa 188(1): 15 (2014). - TYPE:

Singapore, Bukit Timah Nature Reserve, slopes on lower end of Taban Loop along the

stream, 28 May 2014, Leong-Skornickovd, J. & Thame, A, JLS-2793 (holotype SING

(including spirit material); isotypes E, KEP, K). (Fig. 2, 3)

Ecology and distribution. In Singapore, Hanguana neglecta occurs on slopes in the

lowland primary tropical forest of Bukit Timah Nature Reserve, with increased density

closer to the stream. More than a century old herbarium records show that this species

also occun'ed in the Nee Soon and MacRitchie areas. Since it was described, we have

been able to re-locate a small population in the MacRitchie area {JLS-3099, deposited

at SING), but there are still no recent sightings of this species from the Nee Soon area.

The three currently known records from Peninsular Malaysia are confined to Johor

state, and are 56, 107 and 120 years old respectively.

Provisional lUCN conservation assessment. Following the criteria for national and

global conservation assessments (Davison et ah, 2008; lUCN, 2012), Hanguana

Hanguana in Singapore

Fig. 2. Hanguana neglecta Skomick. & Niissalo. A. Habit. B. Base of the plant showing semi-

aseending leafless stem (inset: detail of fruits, note the stigma composed of three obovate lobes

connate at base). C. Infructescence with fruits in various stages of ripeness (immature green to

glossy black mature fruits). From type JLS-2793. (Photos: Jana Leong-Skornickova)

Gard. Bull. Singapore 67(1) 2015

Fig. 3. Hanguana neglecta Skornick. & Niissalo. A. Detail of tepals (side view). B. Detail

of inner tepals, staminodes and staminodial scales. C. Detail of stigma on young fruit. D.

Longitudinal section of fruit, which is always single-seeded. E. Seed (top view). F. Seed (side

view). From type JLS-2793. (Photos: Jana Leong-Skomickova)

neglecta should be considered Endangered (EN D) in Singapore and at least Vulnerable

(VU Blab(iii)) globally (for details see Niissalo et ah, 2014).

Notes. The first collection of Hanguana neglecta on Bukit Timah dates back to 1885

and has been re-collected from the same location several times since. It was Tillich

& Sill (1999) who first realised that this species is not Hanguana malayana. Without

adequate material to hand, they did not pursue a description, but clearly labelled this

species as Hanguana ‘Singapur’ in their morphological study. A complete description

of Hanguana neglecta and a list of specimens examined are provided in Niissalo et al.

(2014).

Hanguana nitens Siti Nurfazilah, Mohd Fahmi, Sofiman Othman & P.C. Boyce,

Willdenowia 40: 207 (2010). - TYPE: Peninsular Malaysia, Johor Bahru, Mersing,

Hutan Simpanan Lenggor, 2°1572.7"N, 103°4376.7"E, 55 m, 18 April 2010, Siti

Nurfazilah bt Abdul Rahman, Boyce, P.C. & Ooi Im Hin HA-48 (holotype KEP; not

yet deposited as of 5 September 2014). (Fig. 4, 5)

Ecology and distribution. Lowland humid swamp forests, in blackwater mires or slow

moving streams. So far known to occur in Peninsular Malaysia and Singapore.

Hanguana in Singapore

Fig. 4. Hanguana nitens Siti Nurfazilah, Mohd Fahmi, Sofiman Othman & P.C.Boyce. A.

Habit. B. Detail of ripe fruits (inset upper left: male flower buds; inset lower left: young

unripe fruits); note the separate, erect, pointed stigma lobes. C. Infructescence. From native

population at MacRitchie sector. Central Catchment Nature Reserve, JLS-3029. (Photos: Jana

Leong- Skomicko va)

Gard. Bull. Singapore 67(1) 2015

Fig. 5. Hanguana nitens Siti Nurfazilah, Mohd Fahmi, Sofiman Othman & P.C.Boyce. A.

Detail of tepals (side view). B. Detail of inner tepal, staminode and staminodial seale. C. Detail

of stigma on young fruit. D. Cross seetion of fruit showing single seed and two empty locules.

E. Longitudinal section of two-seeded fruit. F. Detail of stigma on fully ripe fruit. G. Seed (top

view). H. Seed (side lateral view); note there are no appendages along the incurved rim. I. Seed

viewed from bottom. From native population at MacRitchie sector, Central Catchment Nature

Reserve, JLS-3029. (Photos: Jana Leong-Skornickova)

Provisional lUCN conservation assessment. Although when first described Hanguana

nitens was known to occur in three locations in Johor, a revision of herbarium material

at SING and KEP suggests that this species is more widespread, with specimen records

from Pahang, Selangor and Perak. Based on an EOO, which is larger than 20,000 km^’

this species can be provisionally placed into category of Least Concern (LC) following

the lUCN criteria (lUCN, 2012). Future explorations should, however, focus on an

assessment of population sizes and potential decline in extent and/or quality of habitat

Hanguana in Singapore

and adjust accordingly with more accurate data. In Singapore, Hanguana nitens is

so far known from a single locality in the Central Catchment Nature Reserve with

an area of occupancy (AGO) of c. 200 m^ and a predicted continuing decline in

quality of habitat. The colony of about 150 full-sized shoots consists of both male and

female plants, although considering the stoloniferous colonial nature of Hanguana

nitens, it almost certainly represents only a few individuals (genotypes), and as such

the species should be considered Critically Endangered (CR D) at the national level

(Davison et ah, 2008). Conservation should focus on regular harvesting of ripe fruits,

re-introduction into suitable habitats, and establishment and propagation of material to

saturate potential horticultural interest.

Additional specimens examined. SINGAPORE: Bukit Panjang: 1901, Ridley, H.N.1397

(SING); ibidem, 1906, Ridley, H.N. s.n. (SING); Bukit Timah Road: 19 Dec 1900, Ridley,

H.N. sji. (SING); Central Catchment Nature Reserve: MacRitchie sector, 21 May 2014,

Leong-Skornickovd, J. & Thame, A. JLS-3028 (SING); ibidem, 1 Jul 2014, Leong-Skornickovci,

J. & Thame, J. JLS-3029 (SING); Changi: 4 Oct 1890, Ridley, H.N. s.n. (SING); Chua Chu

Kang [Choa Chu Kang]: 1905, Ridley, H.N. s.n. (SING); With no precise locality: Feb 1837,

Gaudichaud, M. 112 (P, 3x).

PENINSULAR MALAYSIA: Johor: sine dat, Best 8290 (SING); Batu Pahat, Ayer Hitam

F. R., 21 Nov 1965, Ng KEP 100008 (KEP); Ponitan, Pengkalan Raja, sine dat., Alvins 72

(SING); ibidem, 28 Jun 1939, Ngadiman 36651 (SING); Kota Tinggi, Nam Heng estate, 20

Mar 1926, Teniya 640 (SING). Pahang: Kuantan, Sg. Baging F. R., 6 Oct 1989, Khairuddin

FRI 35418 (KEP, 2 sheets). Selangor: Petaling, Batu Tiga, Feb 1904, Ridley, H.N. s.n. (SING,

2 sheets). Kuala Lumpor, 1890, [? Ridley] s.n. (SING); Klang, Changgang, 3 Oct 1937, Mohd

Nurs.n. (SING). Perak: Hilir Perak, Hutan Melintang F.R., 13 Sep 1967, Ng FRI 5698 (KEP);

Taiping, BatuTogoh, Jun 1888, Wray 2128 (SING).

Note: Specimens of Hanguana nitens from Johor cited by Siti Nurfazilah et al. (2010) have not

yet been deposited at KEP as of 5 September 2014.

Notes. The original description of Hanguana nitens as given by Siti Nurfazilah et al.

(2010) largely agrees with our observation of the Singapore population, although the

plants in Singapore are overall more robust, reaching 2 m in height. The largest leaves

reach up to 2.2 m in length with leaf blades to 115 x 1 3.5 cm. Female inflorescences/

infiiictescences are also more robust with the peduncle and rachis reaching 1.1 m

and the median branches on the lowermost levels reaching up to 27 cm in length.

Male inflorescences were not included in the original description of the species (Siti

Nurfazilah et al., 2010). We observ^ed young male inflorescences in May, but with as

yet unopened buds. The structure of the inflorescences is similar to the female plants,

but generally much more slender. Flower buds appear in dense clusters of four to

seven. The tepals are bright green with a bronze tinge externally (Fig. 4B - inset upper

left).

Hanguana nitens certainly has ornamental potential with its beautiful glossy

corrugate leaves which, especially in young leaves, exhibit a pattern of mid-green and

dark green (Fig. 4). Its landscape usage may be similar to Hanguana malayana (as

cuiTently applied) but, as observed in the field, H. nitens is better suited to partially

shaded areas where the largest individuals occur, compared to H. malayana, which

thrives best in full sun.

Gard. Bull. Singapore 67(1) 2015

Hanguana mbinea Skomick. & P.C. Boyce, sp. nov.

Close to Hangtiana pantinensis in fruits ripening to ruby-red, but differing by a

more compact infructescence composed of shorter, stiffer partial inflorescences

attached almost peipendicularly to rachis, median branches to 11 cm with lateral

branches progi'essively shorter, and basally connate stigma lobes (compared to partial

infructescences composed of slender markedly erect branches to 14 cm long, median

and lateral branches of almost same length, and free stigma lobes). Hanguana mbinea

is also similar to H. triangulata (described below), but differs by fruits ripening to

ruby-red and stigma lobes connate basally (sometimes imperfectly) with free round

apices forming bluntly triangular structure (vs ripe fruits cream-white and stigma lobes

with sharply acute apices, connate at base, forming a sharply equilaterally triangular

structure in H. tiiangulata). - TYPE: Singapore, Central Catchment Nature Reserve,

forest around Upper Seletar Reservoir, 19 August 2014, Leong-Skornickovd, J. &

Thame, A. JLS-3037 (holotype SING; isotypes E, KEP). (Fig. 6, 7)

Herbaceous^ dioecious mesophyte to 1.5 m tall; stem terete, to 2.5 cm in diam.,

basally semi-ascending, with age becoming leafless at base, terminally ascending

with crown of up to 25 leaves; stolons absent. Leaves to 165 cm long, spreading

then arching, bases imbricate with hyaline margins (young leaves), turning erose-

marcescent with age; pseudopetiole 50-85 cm long, c. 13 mm wide, accounting

for 1/3-1 /2 of entire leaf length, roundly channelled with sharp margins; leaf blade

65-95 X 12-16 cm, narrowly elliptic, base attenuate, tip long and narrowly attenuate

with apicule 1 5-20 mm, leathery, adaxially mid to dark green, sparsely hairy (silky

appressed hairs evenly distributed; less visible in older leaves), abaxially lighter

green when ifesh, sparsely with silky appresed hairs evenly distributed (denser than

on upper surface); midrib weakly impressed, of the same colour as the rest of the

lamina adaxially, round-raised, mid green, almost glabrous and shiny abaxially.

Male inflorescences not observed, female inflorescences erect at anthesis, of same

structure and dimensions as infructescence. Infructescence erect, comprising up to

8 partial, whorled, altemate-secund, thyrsoid infructescences plus a tenninal spike;

partial infructescences spreading almost perpendicularly to rachis; peduncle and

rachis together up to 50 cm tall, dark purple-brown when fresh, conspicuously pale

brown-grey flocculose, visible portion of peduncle up to 20 cm long; one sterile bract

per peduncle, foliaceous, persistent, narrowly ovate with a basal claw, 88 (inch 8

cm long claw) x 14.5-11.5 cm; bract subtending partial infructescences similar to

sterile bracts, the bract supporting most basal partial infructescence c. 38 x 9 cm,

diminishing in size distally along the infructescence and fully reduced in uppermost

partial infructescences; partial infructescences each comprising up to 11 branches at

basal levels (occasionally two branches connate at base), fewer towards the apex of

the inflorescence, branches arising simultaneously from the axil of the subtending

bract, lateral branches progressively shorter in length (outermost lateral branches c.

y4 of the median branch), median branches at basal levels usually further branched

7-11 cm long, 3-4 mm in diam. Female flowers scattered, always solitary, sessile, all

with an associated minute bract and bracteole; perianth composed of 6 tepals in two

Hanguana in Singapore

Fig. 6. Hanguana rubinea Skomick. & P.C.Boyce. A. Habit. B. Detail of rachis with prominent

dense floeeulose indumentum, whieh is easily rubbed off (inset: detail of fruits and stigma). C.

Infructescenee. From type (Photos: Jana Leong-Skornickova)

Gard. Bull. Singapore 67(1) 2015

Fig. 7. Hanguana rubinea Skomick. & P.C.Boyce. A. Detail of tepals tightly clasping the base

of the fruit. B. Detail of inner tepals, staminodes and staminodial scales. C. Detail of stigma. D.

Cross section of fruit showing single seed and two empty locules. E. Longitudinal section of

fruit. F. Fruit in top view, after removal of apical half of the pulp, showing two seeds and one

empty locule; note the acute appendage at the top of the seed. G. Seed (top view). H. Seed (side

lateral view). I. Seed in semi-lateral view showing an acute appendage. From type JLS-3037.

(Photos: Jana Leong-Skomickova)

whorls tightly clasping ovary/fruit in fresh material, all tepals with prominent bulbous

thickening at base (more prominent in outer whorl), light green with more or less dense

minute red-brown speckles, margin c. 0.2 mm wide, hyaline translucent white; outer

tepals semi-eircular, 1.5-2. 5 mm long, c. 2.5 mm broad, connate at base, sparsely

araehnoid; inner tepals almost semi-circular, 3-3.5 mm long, 4^.5 mm broad at base,

free to base, almost glabrous (occasionally sparsely arachnoid); staminodes 6, in two

whorls, pale green to eream white, triangular, outer staminodes, 0.5 mm long, 0.5 mm

Hanguana in Singapore

broad at base, inner staminodes longer, c. 1 mm long, 0.5 mm at base, each basally

sheathed with a broad narrow scale (often shallowly bilobed), c. 0.5 mm long, and c.

2 mm broad, brown with translucent margin; ovary green, ovoid, glossy green; stigma

3-lobed, each lobe 1.2-1. 4 mm long (fruiting material), broadly ovate (-to bluntly

trullate) with round apex, lobes connate basally (sometimes imperfectly) with free

apices forming bluntly triangular structure (c. 2,5 mm in diam, in fruiting material)

with points of connation seen as grooves, green (flowering stage), matte dark brown

(fruiting stage). Ripe fruit globose, 9-10 imn diam., dark pink-red externally, pulp 1-2

mm thick, cream- white, fairly hard, exuding yellow juice when disturbed, ripenmg

from bright green through cream- white to dark pink-red; seeds 1 -2 per fruit, c. 5 x 4

mm, brown, bowl-shaped with slightly incurved margins, with a triangular appendage

positioned on the distal part of the rim, c. 5 x 4 mm, deeply excavated, cavity filled

with placental tissue.

Etymology. The specific epithet is derived from the ruby-red colour of the ripe fruits.

Ecology and distribution. Growing on the slopes of, or in the proximity of, small/

seasonal streams in primary or partially disturbed primary lowland forest. So far

endemic to Singapore.

Provisional lUCN conservation assessment. Based on our recent collections and on

reliably identified fertile herbarium records from the past 30 years, Hanguana ruhinea

occurs in four locations in Singapore (Bukit Timah, Mandai, MacRitchie and Seletar).

The extent of occun*ence (EOO) is c. 12 km-, the habitat is fragmented and, based on

our observations, the number of adult individuals is fewer than 250, with fewer than

50 individuals in each sub-population. With impending and proposed developments

in some of the existing locations, and serious damage caused by wild boars observed

in two locations, further decline of the populations and further fragmentation of the

habitat is foreseen. Hanguana rubinea should be therefore considered as Critically

Endangered (CR C.2) locally (Davison et al., 2008), and because it is so far endemic

to Singapore, also globally CR Blab(iii,v); C2a(i)) (lUCN, 2012).

Additional specimens examined. SINGAPORE: Bukit Timah Nature Reserve: upper end of

Taban Loop, 28 May 2014, Leong-Skornickovd, J. & Thame, A. JLS-2791 (SING); Central

Catchment Nature Reserve: sector 53 [MacRitchie] 29 Apr 1992, Yong et al. NSR 598

(SING); MacRitchie, off Shinto Trail, 1 Jiil 2014, Leong-Skornickovd, J. & Thame, A. JLS-

3027 (SING); MacRitchie, off Lomie Trail, 14 Aug 2014, Leong-Skornickovd, J, & Thame, A.

JLS-3030 (SING); Chan Chu Kang: 14 Apr 1890, Goodenough, J.S. s.n. (SING); Mandai:

Mandai Forest, 6 Jan 2009, Gwee, A.T SING 2009-09 (SING); Mandai forest, 19 Aug 2014,

Leong-Skornickovd, J. & Thame, A. JLS-3034 (SING; sterile); Ponggol: 1905, Ridley, H.N.

s.n. (SING).

Note: The following two specimens might also represent ifr/ngwana rubinea, but the specimens

deviate slightly from typical H. rubinea and are cited here with caution: Mandai Road, 28 Jul

1929, Corner, E.J.Hs.n. (SING); Seletar, 29 Mar 1889, Ridley, H.N 170 (SING).

Gard. Bull. Singapore 67(1) 2015

Notes. In late fmiting stage, Hanguana rubinea is easy to recognise by its pretty

ruby-red fruits. In early fruiting stages, when the ovaries are still creamy white, it

can potentially be mistaken for Hanguana tnangulata with which it shares a similar

infructescence stmcture with partial infmctescence branches attached almost

perpendicularly to the rachis. However, the shape of the stigma is unmistakeable in the

latter species as the three stigmatic lobes with acutely sharp apices fonn an equilateral

triangle (compare Fig. 7C and 9C). Further differences are in the leaves and the seeds.

The leaf blades of Hanguana rubinea are almost flat, or only weakly corrugate, with

an evenly distributed abaxial silky indumentum, compared to H triangulata which has

corrugated leaves and the indumentum is more visible and flocculose. While the seeds

of both species are bowl-shaped, Hanguana rubinea has an acute appendage along

the rim (Fig. 7G-1) compared to a blunt appendage composed of two sub-lobes in H

triangulata (Fig. 9G-1).

Hanguana triangulata Skomick. & P.C.Boyce, sp. nov.

Similar to Hanguana exultans by fmits ripening cream-white to pale yellow, but

differing by a more compact infructescence composed of shorter, stiffer partial

inflorescences attached almost perpendicularly to rachis and stigma lobes with sharply

acute apices, connate at base, fonning an equilateral triangular structure (vs partial

infructescence somewhat erect, attached at c. 45° to rhachis and stigma composed

of three free lobes). - TYPE: Singapore, Bukit Timah Nature Reserve, slopes around

Taban Loop, 28 May 2014, Leong-Skornickovd, J. & Thame, A. JLS-2789 (holotype

SING). (Fig. 8, 9)

Herbaceous, dioecious mesophyte to c. 1 .6 m tall; stem terete, to 2.5 cm in diam., basally

semi-ascending, with age becoming leafless, terminally ascending with crown of up 20

leaves; stolons absent. Leaves to 150 cm long, spreading then arching; bases imbricate,

margins hyaline (young leaves), turning erose-marcescent with age; pseudopetiole

40-50 cm long, c. 12 mm wide, accounting for c. 1/3 of entire leaf length, roundly

channelled with sharp margins, sparsely softly flocculose; leaf blade 90-106 x 12-13

cm, nan*owly elliptic, base attenuate, tip long and narrowly attenuate with apicule

to 5 mm, leathery, adaxially dark green, sparsely hairy (silky appressed hairs evenly

distributed; less or not visible in older leaves), abaxially mid-green, somewhat shiny,

covered with silky flocculose hairs (unevenly distributed, much denser than on upper

surface); midrib weakly impressed, almost of the same colour as the rest of the lamina,

sparsely flocculose (especially towards the base) adaxially, round-raised, mid green,

sparsely flocculose and shiny abaxially. Female and male inflorescences not observed,

although, based on observations of infructescence architecture, almost certainly erect

at anthesis. Infructescence erect, comprising up to 6 partial, whorled, alteraate-secund,

thyrsoid iiifructescences plus a terminal spike; partial infructescences spreading

almost perpendicularly to rachis; peduncle and rachis together up to 70 cm tall, green

when fresh, conspicuously pale brown-grey flocculose, visible portion of peduncle

up to 40 cm long; sterile bracts two per peduncle, foliaceous, narrowly ovate with a

Hanguana in Singapore

Fig. 8. Hanguana triangulata Skornick. & P.C.Boyce. A. Habit. B. Detail of fruits and the

typieal sharply triangulate stigma. C. Inflorescence. From type 759. (Photos: Jana Leong-

Skornickova)

Gard. Bull. Singapore 67(1) 2015

Fig. 9. Hanguana triangulata Skomick. & P.C.Boyce. A. Detail of tepals (side view). B. detail

of inner tepals, staminodes and staminodial scales. C. Detail of stigma. D. Cross section of

fruit showing single seed and two empty locules. E. Longitudinal section of fruit. F. Fruit in

basal view, after removal of tepals, showing three staminodial scales still attached. G. Seed

(top view). H. Seed (side lateral view). I. Seed (semi-lateral view showing a blunt appendage

composed of two sub-lobes. From type JLS-2789. (Photos: Jana Leong-Skomickova)

basal claw, persistent, 36-71 x 6.5-11.5 cm; bract subtending partial infructescences

similar to sterile bracts, the bract supporting most basal partial infmctescence 20

X 3 cm, diminishing in size distally along the infmctescence and fully reduced in

uppermost partial infructescences; partial infructescences each comprising up to 10

branches at basal levels (fewer towards the apex of the inflorescence), branches arising

simultaneously from the axil of the subtending bract, lateral branches progressively

shorter in length (outermost lateral branches 2/3- 1/2 of the median branch), median

branch 6-8 cm long, c. 3 mm in diam. Female flowers scattered, always solitary.

Hanguana in Singapore

sessile, all with an associated minute bracteole; perianth composed of 6 tepals

in two whorls tightly clasping ovary/fruit in fresh material, outer 3 tepals broadly

triangular with round apex, all tepals with prominent bulbous thickening at base (more

prominent in outer whorl), light green, margin c. 0.2 mm wide, hyaline translucent

white; outer tepals semi-circular, c. 2 mm long, c. 2.8 mm broad, connate at base,

sparsely arachnoid; inner tepals almost semi-circular, c. 3 mm long, c. 4 mm broad,

free to base, almost glabrous (occasionally sparsely arachnoid); staminodes 6, in two

whorls, cream-white, triangular to naiTowly triangular, outer staminodes minute, c.

0.3 mm long, 0.2 mm broad at base, inner staminodes larger, c. 1 mm long, 0.3 mm at

base, each basal ly sheathed with semi-circular staminodial scale, c. 0.8 mm long, and

c. 1.3 mm broad, brown with irregular (usually bilobed) translucent margin; stigma

3-lobed, lobes connate basally, each lobe c. 1.5 mm long (fruiting material), trullate

with shaiply acute apex, lobes perfectly connate basally forming equilateral triangle

(c. 2.5 mm in diam. in fruiting material), raised, matte dark brown in late fruiting stage.

Ripe fruit cream externally, pulp c, 2 mm thick, hard, cream-white, exuding yellow

juice when disturbed, globose, 9-10 mm diam., ripening from bright green to cream-

white; seeds usually two per fruit, c. 5-6 x 4.5-5 mm, brown, broadly boat-shaped to

bowl-shaped, deeply excavated, with a blunt appendage composed of two sub-lobes

positioned on the distal part of the rim, cavity filled with placental tissue.

Etymology. The specific epithet refers to the sharply triangular shape of the stigma, a

character which is conspicuous even in herbarium material.

Ecology and distribution. Hanguana triangulata is an undergrowth species growing

in the proximity of streams in lowland evergreen forest. So far endemic to Singapore.

Provisional lUCN conservation assessment. Only two locations in Singapore with

fewer than 20 individuals in all (AOO 0.5 km2) were observed during recent fieldwork.

There are signs of severe damage caused by wild boars in some adult individuals,

thereby warranting a status of Critically Endangered (CR D) at the national and, given

its endemic status, also global levels (Davison et al., 2008; lUCN, 2012). Immediate

conservation efforts to ensure the survival of this species have focused on the harvesting

of ripe fruits, optimising the protoeol for ex situ cultivation and multiplication, and re-

introductions to suitable habitats.

Additional specimens examined. SINGAPORE: BukitTimah: Ridley, H.N. 9531 (SING);

Bukit Timah Nature Reseive, slopes around Taban Loop, 12 Aug 2014, Leong-Skornickovd, J.

& Thame, A. JLS-2792 (SING); ibidem, slopes around Tiup Tiup path, 14 Sep 2014 , Leong-

Skornickovd, J. J LS- 3 04 6 (SXHGy, Central Catchment Nature Reserve: forests around Upper

Seletar Reservoh, 19 Aug 2014, Leong-Skornickovd, J. & Thame, A. JLS-3036 (SING, sterile);

Kranji: 5 Aug 1889, Goodenoiigh, J.S. s.n. (SING); Pulaii Ubin: Aug 1898, Ridley, H.N. s.n.

(SING).

Notes. Hanguana triangulata is currently known to occur only in the Bukit Timah

Nature Reserve (confirmed from fruiting material) and in the Central Catchment

Gard. Bull. Singapore 67(1) 2015

Nature Reserve in forests around Upper Seletar Reservoir (identification based on

sterile material only). Historical herbarium records suggesting its past presence in

Kranji and Pulau Ubin are both over a century old. The sharply triangular stigma

is easily observable in dried material, although the sharp tips may cuiTe somewhat

downwards. Seeds extracted from material as old as 1 889 match well to seeds extracted

from recently collected material in Bukit Timah Nature Reserve (both exhibiting the

blunt weakly bilobed appendage, see Fig. 9G-I).

Introduced and naturalised species

Hanguana corneri Skomick. & RC.Boyce, sp. nov.

Unique amongst cuiTently known Hanguana species by its leaves which are shiny

emerald-green above and deep purple and silvery flocculose beneath. - TYPE:

Peninsular Malaysia, Johor, Sungai Berassau, Mawai-Jemulang Road, 7 February

1935, Corner, E.J.H.s.n. (holotype SING!). (Fig. 10)

Herbaceous, dioecious mesophyte to c. 0.8 m tall; stem terete, to 2 cm in diam.,

basally semi-ascending, with age becoming leafless at base, terminally with crown

of 15-20 leaves; stolons absent. Leaves to 85 cm long, spreading then arching, bases

imbricate with hyaline margins (young leaves), turning erose-marcescent with age;

pseudopetiole 10-20 cm long, accounting for !4-‘/3 of entire leaf length, roundly

channelled with sharp margins; leaf blade 54-66 x 10-14.5 cm, narrowly elliptic

with undulate margins, base attenuate, tip nan'owly acute with apicule 2-3 mm, softly

leathery, adaxially dark green, with glossy lustre, abaxially purple red, with dense soft

appressed flocculose indumentum; midrib weakly impressed, mid-green (much lighter

than the rest of the lamina) adaxially, round-raised, light pink-brown, flocculose

abaxially. Female inflorescence not observed, although, based on obseiwations of

infmctescence architecture, almost certainly erect at anthesis. Male inflorescence

erect at anthesis, comprising 6 partial inflorescences and a terminal spike; peduncle

and rachis c. 30 cm long; peduncle c. 6 cm long, no sterile foliaceous bract observed;

bract subtending lowest partial inflorescence foliar, naiTOwly ovate, c. 19 cm long

(width obscured in type specimen), with basal claw c. 2 cm, distally diminishing in size

to narrowly triangular bracts, fully reduced at uppermost partial inflorescences. Male

flowers scattered, in groups of (2-)3-5; perianth composed of 6 tepals in two whorls,

all tepals ovate, greenish (according to a note on type specimen), outer tepals c. 1 mm

long, inner tepals c. 1 .5 mm long (measured from dried flower buds). Female flowers

scattered, in pairs or solitary, sessile, all with an associated minute bract and bracteole;

perianth composed of 6 tepals in two whorls tightly clasping fruit (in dry material),

all tepals with prominent bulbous thickening at base (more prominent in outer whorl);

outer tepals semi-circular, c. 1 mm long, c. 1.5 mm broad, connate at base; inner

tepals almost semi-circular, c. 2 mm long, 2-2.5 mm broad, free to base; staminodes

and staminodial scales not observed; stigma 3-lobed, each lobe c. 7 mm long (dried

fruiting material), ovate with blunt apex, lobes perfectly connate basally with free

Hanguana in Singapore

Fig. 10. Hanguana corneri Skornick. & P.C.Boyce. A. Habit. B. Base of the plant showing

semi-aseending leafless stem. C. Detail of lamina abaxially, showing the unique dark purple-

red eoloration and dense cover of silvery flocculose hair. From plant growing at Bukit Timah

Nature Reserve, JLS-2790. (Photos: Jana Leong-Skomickova)

Gard. Bull. Singapore 67(1) 2015

apices forming bluntly triangular structure with raised centre (c. 1.1 mm in diam., in

dried fruiting material), matte dark brown in late fruiting stage. Infmctescence erect,

comprising up to 5 partial altemate-secund, thyrsoid infructescences plus a teraiinal

spike; partial infructescences semi-erect, at an angle c. 40°-50“ to rachis; peduncle

and rachis together up to 45 cm tall; peduncle up to 10 cm long, one sterile bract per

peduncle, foliaceous, persistent, nari'owly ovate with basal claw, c. 35 (inch 5 cm claw)

X c. 6 cm; bract subtending partial infructescences similar to sterile bract, distally

diminishing in size to nan'owly ti'iangular bracts, fully reduced in uppermost partial

inflorescences; partial infructescences each comprising up to 5 branches at basal levels,

fewer towards the apex of the inflorescence, branches arising simultaneously from the

axil of the subtending bract, lateral branches progressively shorter in length (outemiost

lateral branches c. % of the median branch), both median and lateral branches usually

further branched, median branch to 10 cm long, c. 2.5 mm in diam. Fruit (described

from old dried fruit) globose, c. 4 mm in diam.; seeds brown, c. 3-3.5 mm in diam.,

34 globose to ellipsoid, ostiole wedge-shaped accounting for c. % of the seeds, deeply

excavated, cavity filled with placental tissue.

Etymology. We name this species for E.J.H. Comer (1906-1996), a botanist and

mycologist who first collected this beautiful species. He was a giant among tropical

Asian botanists and, in particular, for ‘Malayan’ botany.

Ecology and distribution. Native to Peninsular Malaysia (Johor) and introduced

to Singapore. According to the notes on Comer’s specimen, this species occurs in

swampy forests with flowering occuming in Febmary. Populations of what is almost

certainly the same species, although not occurring in swampy areas, are known from

SW Sarawak (Boyce, pers. obs.), notably in the Penrissen Range, where plants occur

intermixed with another two as-yet undescribed Hanguana species a few metres from

a precipitous escarpment marking the border with Kalimantan Barat.

Provisional lUCN conservation assessment: Owing to a lack of data on the distribution

and population sizes of this species in Peninsular Malaysia, and the uncertainty as to

whether this species also extends to Borneo, we propose that this species be treated as

Data Deficient (DD).

Additional specimens examined. SINGAPORE: Bukit Timah Nature Reserve: slopes around

Taban Loop, 28 May 2014, Leong-Skornickovd, J. & Thame, A. JLS-2790 (SING).

Notes. A single plant of this conspicuous species has been discovered in the Taban

Loop area of Bukit Timah Nature Reserve (BTNR). A revision of material from

Peninsular Malaysia at SING revealed the existence of a male flowering specimen

of this species, collected by E.J.H, Comer in 1935 from Sungai Berassau (Peninsular

Malaysia, Johor). Although it is almost certain that living plants of this species were

brought to Singapore from a field trip conducted by Comer, who at that time worked

in Singapore Botanic Gardens (1929-1945), it is not clear how they ended up on Bukit

Hanguana in Singapore

Timah. The first author has observed, as part of earlier work on native Zingiberales,

that several Scaphochlamys species, all of them certainly not native to Singapore, also

occurred around the same area. Some of these Scaphochlamys species were described

as new to science by R.E. Holttum from herbarium material collected by Comer.

Holttum (1950) explicitly stated that he based his descriptions solely on the herbarium

material and that he had never seen these species alive, making any connection of

Holttum to the Bukit Timah plantings unlikely. According to John Dransfield and Ruth

Kiew, who both pursued their PhD studies under Comer’s supervision and knew him

well, it is also highly unlikely that it was Comer himself who planted these on Bukit

Timah (J. Dransfield, R. Kiew, pers. comm.). While there are no official records, we

tend to agree with a suggestion by Dransfield, that living plant material was most

likely brought back to Singapore Botanic Gardens by Comer’s field assistants Kiah

and Md. Nur. When and by whom the material was planted on Bukit Timah, however,

remains unclear.

Several plants of Hanguana corneri are cultivated in the living collections at

the Forest Research Institute, Malaysia. These collections, unfortunately, lack exact

provenance record, although they are also likely to be from Johor (Saw Leng Guan,

pers. comm.).

The above description is based on cuirently available material consisting of

sterile living plants m the Bukit Timah Nature Reserve, a male dried specimen (the

type), and old infmctescences observed in the living collections of the Forest Research

Institute, Malaysia. While the description is incomplete, we feel that the species is

so distinct and recognisable, even in herbarium specimens, that it wamants fomial

description. The description can be further improved with additional material gathered

in the future.

Hanguana malayana (Jack) Merr., Philipp. J. Sci., C, 10: 3 (1915). - Veratrum

malayanum Jack, Malayan Misc. 1(1): 25 (1820). - Veratronia malayana (Jack) Miq.,

FI. Ned. Ind. 3: 553. (1859). - Susum malayanum (Jack) Planch, ex Hook, f, FI. Brit.

India 6: 391 (1892). - TYPE: Malaysia, Pulau Pinang [“Poeloe Pinang”], (not

traced). (Fig. 11, 12)

Hanguana anthelminthica (Blume ex Roem. & Schult.) Masam., Enum. Phan. Bom.:

81. (1942). — Susum anthelminthicum Blume ex Roem. & Schult., Syst. Veg. 7(2):

1493 (1830). — Susum malayanum f. aquatica Backer, Handb. FI. Java 3: 3 (1924).

— Hanguana malayana subsp. anthelminthica (Blume ex Roem. & Schult.) Backer,

Bekn. FI. Java 10(212): 2 (1949). — Hanguana malayana var. anthelminthica (Blume

ex Roem. & Schult.) Bakli., Blumea 6: 399 (1950). - TYPE: Indonesia, Java “in

paludibus circa Buitenzorg”, Blume s.n. (lectotype L! [L21 07638, L21 07639; both

sheets cross-referenced as being part of single gathering], here designated; isotype BO

n.v. (as per Siti Nurfazilah et al., 2010).

Hanguana aquatica Kaneh., Trans. Nat. Hist. Soc. Formosa 25: 8 (1935). - TYPE:

Caroline Islands, Palau, Almonogni, Babeldaob, 13 April 1938, S. Hatusima 4866

(holotype FU!).

Gard. Bull. Singapore 67(1) 2015

Fig. 11. Hanguana malayana (Jack.) Merr. A. Habit. B. Branches with ripe fruits (inset: close-

up detail of fruits and stigma). C. Infrueteseenee. From plants eultivated at Singapore Botanie

Gardens, JLS-3033. (Photos: Jana Leong-Skornickova)

Hanguana in Singapore

Fig. 12. Hanguana malayana (Jack.) Merr. A. Detail of tepals clasping base of ripe fruit (side

view). B. Detail of inner tepals, staminodes and staminodial scales. C. Detail of stigma. D.

Cross section of fruit showing three well-developed seeds. E. Longitudinal section of fruit. F.

Detail of tepals clasping base of ovary (side view). G. Seed (top view). H. Seed (bottom view).

I. Seed (lateral view). From plants cultivated at Singapore Botanic Gardens JLS-3033. (Photos:

Jana Leong-Skornickova)

Notes: The identity of Hanguana malayana has been the subjeet of mueh debate, as

the original material by Jack is still missing and Jack’s original description, based

on material from Pinang [Pulau Penang], although fairly long, does not offer many

characters which could be considered species specific. The name has been uncritically

applied to almost all forest species in Malaysia and Singapore since Backer’s treatment

of Hanguana for Flora Malesiana (Backer, 1951). The most recent attempt to clarify

the identity and circumscription of Hanguana malayana is that of Siti Nurfazilah et al.

(2010), who proposed that the name H. malayana should be applied to a widespread

Gard. Bull. Singapore 67(1) 2015

large, helophytic, stoloniferous species. As this is the species which occurs in cultivation

in Singapore (Fig. 11, 12), this current application is followed here with caution.

Hanguana malayana (sensu Siti Nurfazilah et aL, 2010) is freely available in

Singapore’s nurseries and is occasionally used in landscaping. All herbarium records

originating from Singapore, and previously identified as Hanguana malayana, at the K

and SING herbaria (no specimens from Singapore exist at KEP), turned out to belong

to the four species above treated. Our field explorations have so far not confirmed the

presence of any populations which could be considered to be wild, although populations

of Hanguana malayana have been planted in various locations in Singapore where

they thrive and may naturalise easily. There is also no confirmed herbarium record

of Hanguana malayana from areas close to Singapore. A revision of all Hanguana

herbarium sheets at K, KEP, P and SING revealed that there are no collections of H.

malayana from Johor. All confinned collections in Peninsular Malaysia were collected

only in Perak, Selangor, Terengganu, and a single historical collection of Griffith 6014

(K), collected in Malacca [Melaka], which is about 200 km from Singapore. We have

located two collections of Hanguana malayana from Sumatra (Indonesia), of which

Yates 2281 (P, 4x) is from Medan (c. 600 km from Singapore), while Zollinger’s undated

collection is without precise locality. Considering the above, there is, therefore, no

reliable evidence, historical or recent, which would support the theory that Hanguana

malayana was ever native to Singapore.

Conclusion

Recent discoveries of six native species new to science in heavily urbanised Singapore

(including Hanguana rubinea and //. triangulata described above; H, neglecta -

Niissalo et ah, 2014; Zingiber singapurense Skomick. - Leong-Skomickova et ah,

2014; two Utania species - Sugumaran & Wong, 2014), serves to remind us of several

important points. Firstly, that although Singapore is the most densely collected country

in SE Asia and its primary vegetation has been severely disturbed, the process of

documenting and fully understanding its rich biodiversity is incomplete. Secondly,

that while herbarium-based taxonomy is sufficient in some plant groups, satisfactory

progress on others without observation of fertile material in the field is almost

impossible (for example in Zingiberaceae and Hanguanaceae). And lastly, that such

field-based knowledge can lead to the discovery of previously overlooked characters,

in this case the staminodial scales and the seed stmcture, which can then be applied to

the identification of existing herbarium collections.

A poor understanding of the genus Hanguana in Singapore and the misapplication

of the name Hanguana malayana, which was previously treated as Vulnerable in

the Singapore Red Data Book (Davison et al., 2008), obviously has implications

for conservation of all four native species. These four species are all currently

considered to be Endangered or Critically Endangered locally, and those endemic to

Singapore also globally {H. neglecta ENATJ, H. nitens CR/LC, H. rubinea CR/CR

and H. triangulata CR/CR). Further conservation work, including the observation of

Hanguana in Singapore

pollinators and dispersal agents and the DNA barcoding of native species is planned.

Also, now that the hidden diversity of Hanguana in Singapore has been recognised, in

situ and ex situ conservation and propagation efforts for this interesting yet extremely

neglected plant family in Singapore is underway. Due to the lack of any records

suggesting that the large stoloniferous helophytic species, currently called Hanguana

malayana, was ever native to Singapore, we recommend that this species be treated as

not native to Singapore.

ACKNOWLEDGEMENTS. The first author thanks the National Parks Board, Singapore.

Our sincere thanks are due to numerous NParks staff and volunteers, who provided us with

continuous support in our taxonomic and conservation research on the Singapore flora, in

particular: Desmond Lee, Mishak Sunari (Sungei Buloh Wetland Reserve, field support),

Derek Liew, Chew Pin Ting (Central Catchment Nature Reserve, field support), Ang Wee Fong

(Horticulture and Community Gardening Division), Aung Thame, Sunia Teo and Michael Leong

(Singapore Botanic Gardens, field support and propagation of native species for conservation

purposes). We thank Prof Hans-Juergen Tillich for fruitflil discussions and suggestions on the

possible origin and function of staminodial scales, and confinnation of the identity of Hanguana

bogneri, and Dr Saw Leng Guan from the Forest Research Institute, Malaysia for granting

access to FRIM’s living collections of Hanguana , which helped us improve the description

of the imperfectly known Hanguana corneri. We also thank Eva-Nicole Fisher (volunteer at

Singapore Botanic Gardens) for her help with a translation of the research paper by Tillich &

Sill (1999) which was originally published in German, and to the anonymous reviewers for

useful comments to improve this manuscript.

References

Airy Shaw, H.K. (1980). Anew species of Hanguana from Borneo. Kew Bull. 5(4): 819-821.

Bachman, S., Moat, J., Hill, A.W., de la Torre, J. & Scott, B. (2011 ). Supporting Red List threat

assessments with GeoCAT: Geospatial Conservation Assessment Tool. In: Smith, V. &

Penev, L. (eds) e-lnfrastmctures for data publishing in biodiversity science. ZooKeys

150: 117-126. (Version BETA)*

Backer, C.A. (1951). Hanguana. In: Steenis, C.G.G.J. van (ed) Flora Malesiana, ser. 1, 4(3):

248-250.

Bayer, C., Appel, O. & Rudall, P.J. (1998). Hanguanaceae. In: Kubitzki, K. (ed) The Families

and Genera of Vascular Plants, vol. IV. Pp. 223-225. Berlin-Heidelberg: Springer-

Verlag.

Beentje, H. (2012). The Kew Plant Glossary, an illustrated dictionary of plant terms (revised

edition). Royal Botanic Gardens, Kew: Kew Publishing.

Chong, K.Y., Tan, H.T.W. & Corlett, R.T. (2009). A Checklist of the Total Vascular Plant Flora

of Singapore: Native, naturalised and cultivated species. Singapore: Raffles Museum of

Biodiversity Research, National University of Singapore.

Davison, G.W.H., Ng, P.K.L. & Ho, H.C. (eds) (2008) The Singapore Red Data Book:

Threatened plants and animals of Singapore, 2nd ed. Singapore: The Nature Society

(Singapore).

Gard. Bull. Singapore 67(1) 2015

Holttum, R.E. (1950). The Zingiberaceae of the Malay Peninsula. Gard. Bull Singapore 13:1-

249.

lUCN (2012). lUCN Red List Categories and Criteria: Version 3.1. Second edition. Switzerland,

Gland and UK, Cambridge: TUCN.

Keng, H., Chin, S.C. & Tan, H.T.W. (1998). The Concise Flora of Singapore, volume IT.

Monocotyledons. Singapore: Singapore University Press.

Leong-Skomickova, J., Thame, A. & Chew, P.T. (2014). Notes on Singapore native Zingiberales

I: A new species of Zingiber and notes on the identities of two further Zingiber taxa.

Card. Bull. Singapore 66(2): 153-167.

Low, Y.W., Leong, P.K.F., Tee, S.P., Rajesh Singh, Tay, M.L.C. & Wong, K.M. (2014).

Margaritaria (Phyllanthaceae), a new generic record for the Singapore flora. Gard.

Bull. Singapore 66(1): 47-56.

Mohd Faluui, A.B., Alimad Sofiman, O. & Boyce, PC. (2012). Studies on Hanguana

(Commelinales: Hanguanaceae) for Sunda TV: Hanguana major clarified, a new species

from Sarawak, Malaysian Borneo, and description of male reproductive organs in H.

bakoensis. Ke\v Bull. 67: 205-211.

Niissalo, M., Wijedasa, L., Boyce, RC. & Leong-Skomickova, J. (2014). Hanguana neglecta

(Hanguanaceae): a new plant species from a heavily collected and visited reserve in

Singapore. 188(1): 14-20.

Rodda, M., Tran, H.D., Chiew, P.T., Liew, D. & Leong-Skomickova, J. (2012). The rediscovery

of Dischidia hirsuta (Apocynaceae, Asclepiadoideae) in Singapore. Gard. Bull.

Singapore 64(2): 293-299.

Rudall, PJ., Stevenson, D.W. & Linder, H.P. (1999). Stmcture and systematics of Hanguana, a

monocotyledon of uncertain affinity. Austral Syst. Bot. 12: 311-330.

Siti Nurfazilah, A.R., Mohd Fahini, A.B., Ahmad Sofiman, O. & Boyce, PC. (201 0). Studies on

Hanguana (Commelinales-Hanguanaceae) for Sunda II: Five new forest species, a new

species record from Peninsular Malaysia, and a redefinition of the taxonomic limits of

Hanguana malayana. Willdenowia 40: 205-219.

Siti Nurfazilah, A.R., Ahmad Sofiman, O. & Boyce, PC. (2011). Studies on Hanguana

(Commelinales-Hanguanaceae) for Sunda I: Hanguana bakoensis, a new forest species

from Sarawak, Malaysian Borneo, and notes on critical morphologies for elucidating

Hanguana tAxomvny. Acta Phytotax. Geobot. 61(3): 139-143.

Sugumaran, M. & Wong, K.M. (2014). Studies in Malesian Gentianaceae, VI. A revision of

Utania in the Malay Peninsula with two new species. PI. Ecol Evol. 147(2): 213-223.

Tillich, H.-J. (1996). Seeds and seedlings in Hanguanaceae and Flagellariaceae

(Monocotyledons). Sendinera 3: 187-197.

Tillich, H.-J. & Sill, E. (1999). Systematische studien zur morphologic und anatomic von

Hanguana Blume (Hanguanacae) und Flagellaria L. (Flagellariaceae). Mit der

Beschriebung einer neuen Art, Hanguana bogneri spec. nov. Sendtnera 6: 215-238.

Gardens’ Bulletin Singapore 67(1): 29-33. 2015

doi: 10.3850/S238258 12 15000022

Dischidia acutifolia (Apocynaceae, Asclepiadoideae)

a new record for the Singapore flora

M. Rodda\ D. Lee^ & S. Mishak^

^Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, 259569 Singapore

rodda.michele@gmai] .com

^Conservation Division, Central Nature Reserve, National Parks Board,

1 Cluny Road, 259569 Singapore

ABSTRACT. We report the discovery of Dischidia acutifolia Maingay ex Hook.f., a new

record for the Singapore flora. The species is only known from a single locality in Sungei

Buloh Wetland Reserve and is therefore considered Critically Endangered in Singapore. A

lectotype for the name is selected.

Keywords. lUCN Red Lists, Sungei Buloh Wetland Reserve

Introduction

In Singapore there are ten native species of Dischidia R.Br. (Chong et al., 2009;

Rodda et ah, 2012). Five of these are considered nationally extinct and one, Dischidia

singaporensis Ridh, is considered a globally extinct species. A general introduction

to the genus Dischidia can be found in various recent local treatments (Rintz, 1980;

Li et ah, 1995; Forster et ah, 1996; Jagtap & Singh, 1999; Livshultz et ah, 2005)

while information on the local Singapore species is provided in Rodda et ah (2012). In

September 2013 Desmond Lee and Mishak Shunari found an unidentified Dischidia

growing at the base of the tmnk of Talipariti tiliaceum (L.) Fryxell in Sungei Buloh

Wetland Reserve. A small sterile cutting was brought to the Singapore Herbarium

for identification. The cutting was incorporated into the Singapore Botanic Gardens’

Research Living Collections where it rooted and soon after started flowering before

producing any new growth. It was then identified as Dischidia acutifolia, a species

rather common from Thailand to the Moluccas (Rintz, 1980). The species is not in

the most recent checklist of Singapore’s vascular plant flora (Chong et ah, 2009) and

no specimens collected in Singapore have previously been deposited in SING. We

therefore here report it as a new record for the Singapore flora.

We can speculate that the Singapore plant may originate from a recent

colonisation from seeds wind-dispersed from the coastal mangroves in Johore

(Malaysia). However it is equally likely that the species, an epiphyte often growing

high up in the tree canopy and bearing inconspicuous flowers, may have always been

part of the Singapore flora but was never previously collected.

Gard. Bull. Singapore 67(1) 2015

Conservation considerations

The species is only known from a single collection and is, therefore, to be considered

Critically Endangered in Singapore, according to guidelines given in The Singapore

Red Data Book (Davison, 2008). The original plant observed in 2013 was negatively

affected by the drought in early 2014 (Ziegler et ah, 2014), and the recovery has been

slow (Lee and Mishak, personal observation). In contrast, the clone is very vigorous in

cultivation and is being mass propagated at the Singapore Botanic Gardens (SBG ace.

no. 20132480), Stock will soon be released for reintroduction trials.

The morphological description below is based on the specimen collected in

Singapore. Measurements from cultivated materials are indicated in parentheses when

they are significantly different from wild material.

Taxonomy

Dischidia aciitifolia Maingay ex Hook.f, FI. Brit. India 4(10): 51 (1883). - TYPE:

Peninsular Malaysia, Malacca, 30 Oct 1 865 or 6, Maingay, A.C. 1960 {Kew distribution

number 1122) (lectotype K, designated here [K0009 11032]). (Fig. 1)

Plants succulent, herbaceous, glabrous; epiphytic or hemi-epiphytic vines growing

loosely rooted on the host tree stems, clinging by adventitious roots, rarely pendulous.

Latex white. Roots on mature plants adventitious, produced at the node, along the

intemodes and just below each node, where they are sometimes paired. Leafy stems

terete, up to 2.5 mm diameter, dark green; intemodes (2-)5-20(-25) cm long. Leaves

opposite, petiolate; petiole cylindrical, slightly flattened above, 4-7 mm long, 1.5-2

mm in diameter; lamina dark green, underneath lighter green with darker midrib and

margin, fleshy, stiff (less so in cultivation), lanceolate, 4-7 x 1.5^ cm (to 10 x 5

cm in cultivation); apex acute-apiculate, base (round) acute, margin entire, midrib

and secondary veins slightly raised on adaxial surface, secondary veins 3-6 pairs,

branching at 35-60° from midrib. Inflorescences usually bearing a single open flower

at a time (up to 5 in cultivation) and 2-A developing buds; peduncle extra-axillary or

apparently axillary when borne on very short shoots, persistent, rachis one or two per

peduncle, bearing scars of previous flowerings 2-5 mm long, 2-2.5 nmi in diameter

(to 2 cm long in cultivation); pedicels 0. 6-0.8 mm long, c. 0.6 mm in diameter.

Calyx c. 2 mm in diameter, green, lobes (round) ovate 0.6-0. 8 x 0.5-0. 8 mm, apex

round, sparsely ciliate. Calycine colleter 1 or 2 at each calyx lobe sinus, ovate, with a

round apex, 100-120 x 80-100 pm. Corolla succulent, urceolate, 3.5^ x 2-2.5 mm,

basally lighter yellow, progressively fading into light pink or white at the lobe apices,

externally glabrous, corolla throat with one ring of retrorse hairs; lobes triangular,

0.5-0. 7 X c. 0.6 mm, apex acute. Corolline corona absent. Staminal corona lobes

1.5-1. 7 mm high, c. 1.5 mm in diameter, composed of five light yellow appendages

shaped like an inverted anchor with a stipe and lunate apex. Style head conical, hidden

by the anther appendages. Pollinarium erect, 550-600 x 500-600 pm with two oblong

Dischidia acutifolia a new record for Singapore

Fig. 1. Dischidia acutifolia in Singapore. A. The original plant as eollected by Lee and Mishak

in Sungei Buloh Wetland Reserve in 2013. B. First flowering in cultivation at SBG. C. Dissected

flower. The single ring of hairs lining the corolla throat, the yellow corona appendages and the

anther appendages covering the stigma head can be observed. D. Inflorescence with five open

flowers; usually in wild-collected plants the inflorescences have only one open flower at a

time. E. Pollinarium with twin pollinia. B-E Based on living plants cultivated at the Singapore

Botanic Gardens (SBG ace. no. 20132480). (Photos: M. Rodda)

Gard. Bull. Singapore 67(1) 2015

pollinia, 230-250 x 90-100 pm, apex and base rounded to obtuse, retinaculum 170-

190 X 60-70 pm, caudicles elongated, laterally crested, 250-270 pm long. Ovaty bi-

carpellate, bottle-shaped, 1-1.5 mm long, each carpel c. 0.7 mm in basal diameter.

Fruits and seeds not seen.

Singapore specimens examined. Sungei Buloh Wetland Reserve, Lee, D. & Shunari, M. s.n.,

eultivated at the Singapore Botanie Gardens, vouehered on 30 Get 2014 as Rodda, M. MR898

(SING, K, KEP).

Notes. The description of Dischidia acutifolia by Hooker (1883) was based on the

collection Maingay {Kew Distrib. 1122). The herbarium of a particular specimen

was not indicated. In Kew there is a specimen labelled as Kew distribution number

1122 which is also labelled with the Maingay numbering series (Maingay 1960). An

exhaustive search for duplicates has not been earned out but since the K specimen is

not labelled as a unicate it is possible that duplicates may be found in other herbaria

holding Maingay materials (B, BM, CAL, CGE-B, G, L [Steenis-Kruseman, 1950]).

Therefore a lectotype needs to be selected. The K specimen is well preserved and

matches the description of Dischidia acutifolia and is therefore selected as lectotype

for the name.

ACKNOWLEDGEMENTS. We thank National Parks Board, Singapore, for funding our

research, and Tanya Livshultz and one anonymous referee for their constructive comments on

the manuscript.

References

Chong, K.Y., Tan, H.T.W. & Corlett, R.T. (2009). A Checklist of the Total Vascular Plant Flora

of Singapore: Native, naturalised and cultivated species. 273 p. Singapore: Raffles

Museum of Biodiversity Research, National University of Singapore.

Davison, G.W.H. (2008). The Red List categories. In: Davison, G.W.H., Ng, P.K.L. & Ho, H.C.

(eds) The Singapore Red Data Book: Threatened plants and animals of Singapore. 2nd

ed. Pp. 1-4. Singapore: Nature Society (Singapore).

Forster, P.I., Liddle, D.J. & Nicholas, A. (1996). Asclepiadaceae. In: Orchard, A.E. (exec, ed)

Flora of Australia, vol. 28 (Gentianales). Pp. 197-283. Melbourne: CSIRO.

Hooker, J.D. (1883). Dischidia. In: Hooker, J.D. (ed) The Flora of British India 4: 49-52.

London: L. Reeve & Co.

Jagtap, A.P. & Singh, N.P. (1999). Asclepiadaceae. Fascicles of Flora of India. Fascicle 24.

Calcutta: Botanical Survey of India.

Li, P.T., Gilbert, M.G. & Stevens, W.D. (1995). Asclepiadaceae. In: Wu, Z. & Raven, P.H.

(eds) Flora of China 16 (Gentianaceae through Boraginaceae). Pp. 189-270. St. Louis:

Missouri Botanic Garden.

Livshultz, T, Tran, T.B., Bounphanmy, S & Schott, D. (2005). Dischidia (Apocynaceae,

Asclepiadoideae) in Laos and Vietnam. Bliimea 50: 113-134.

Rintz, R.E. (1980). The Peninsular Malayan species of Dischidia (Asclepiadaceae). Blumea

26: 81-126.

Dischidia acutifolia a new record for Singapore

Rodda, M., Tran, H.D., Chiew, RT., Liew, D. & Leong-Skomickova, J. (2012). The rediscovery

of Dischidia hirsuta (Apocynaceae, Asclepiadoideae) in Singapore. Gard. Bull.

Singapore 64: 293-299.

Steenis-Kniseman, M.J. van (1950). Malaysian plant collectors and collections: Being a

cyclopaedia of botanical exploration in Malaysia and a guide to the concerned literature

up to the year 1950. Flora Malesiana, Ser. 1,1: 5-605. Jakarta: R Noordhoff

Ziegler, A.D., Terry, J.R, Oliver, G.J.H., Daniel Friess, A., Chuah, C.J., Chow, W.T.L. &

Wasson, R.J. (2014). Increasing Singapore‘s resilience to drought. Hydrolog. Process.

28: 4543-4548.

Gardens’ Bulletin Singapore 67(1): 35-37. 2015

doi: 10.3850/S238258 12 15000034

From Ancistrocladus to Tristaniopsis via Tetramerista - the

convoluted history of a Wallich collection and its impact

on the native flora of Singapore

LM. Turner

Research Associate, Royal Botanic Gardens Kew,

Richmond, Surrey, UK

j_trop_ecol@yahoo.co.uk

ABSTRACT. Ancistrocladus sagittatus Wall, ex Planch. (Ancistrocladaceae) is the name of a

plant species solely based on a sterile collection made by Nathaniel Wallich in Singapore in

1822. Since 1875, this name has been treated as a variety or synonym oi Tetramerista glabra

Miq. (Tetrameristaceae). However, the original material is actually referable to Tristaniopsis

(Myrtaceae), and Ancistrocladus sagittatus is here lectotypified and reported as a synonym

of Tristaniopsis merguensis (Griff) Peter G.Wilson & J.T.Waterh. for the first time. The new

determination of the Wallich collection means that the record of Tetramerista glabra in the

native flora of Singapore requires reassessment. It is concluded that Tetramerista glabra, the

genus Tetramerista and the family Tetrameristaceae should all be excluded from inventories of

the native flora of Singapore.

Keywords. Ancistrocladus, flora, Singapore, synonymy, Tetramerista, Tristaniopsis, typification

Introduction

Nathaniel Wallich (1786-1854) was a Danish surgeon who studied botany in

Copenhagen. In 1 807, he arrived at Serampore in India to work as a medical doctor.

By 1814 he had officially joined the British East India Company, still working in the

medical service, and in 1 8 1 5 he was appointed Superintendent of the Botanic Garden

in Calcutta. Wallich undertook various expeditions to collect specimens. This included

a trip from Calcutta to Penang and Singapore in 1822. Of the many collections made

by Wallich and his assistants on this trip, one from Singapore, of apparently only sterile

material, was Ancistrocladus sagittatus by Wallich. The name was included as

number 1055 in Wallich ’s monumental listing of the East India Company Herbarium

known as the Numerical List (Wallich, 1828-1849) or Wallich’s Catalogue. As with

the vast majority of plant names in the Numerical List, Ancistrocladus sagittatus was

a nomen nudum in the absence of a description or reference to a published one. An

exception to this rule was actually provided by the generic name Ancistrocladus.

This was validly published in the Numerical List as a replacement name for Wormia

Vahl, a later homonym of Wormia Rottb. (Dilleniaceae). The top set of the East India

Company collections was maintained as a closed collection, often refeiTed to as the

Wallich Herbarium, firstly at the Limiean Society of London and later, from 1913, at

Gard. Bull. Singapore 67(1) 2015

the Royal Botanic Gardens Kew (K-W). Other material was distributed by Wallich

under the list numbers to many different individuals, including William Hooker, and

institutions, though it should not be assumed that all specimens with the same number

represent true duplicates.

Ancistrocladus sagittatus was first validated by the French botanist Jules Emile

Planchon in a paper on Ancistrocladus (Planchon, 1849). Planchon based the species

solely on Wallich ’s collection, but did not state which specimens he had seen. Planchon

worked as assistant to William Hooker at Kew in the period 1 844-1 848 and, for several

other Ancistrocladus species in his revision, he mdicated that he had seen specimens

in Hooker’s personal herbarium. I therefore designate a specimen distributed under

the East India Company number 1055, now in the general herbarium of the Royal

Botanic Gardens Kew and formerly in Hooker’s personal herbarium, as the lectotype

of Ancistrocladus sagittatus Wall, ex Planch.

Alfred William Bennett seems to have been the first to realise thsit Ancistrocladus

sagittatus did not belong in Ancistrocladus. In the account of the Ochnaceae for the

Flora of British India, Bennett (1875) treated Ancistrocladus sagittatus as a variety

of Tetramerista glabra Miq. (Tetrameristaceae, fonnerly placed in the Oclmaceae or

Theaceae). A casual encounter with this piqued my interest because, if this reduction

were taxonomically correct, then Ancistrocladus sagittatus, the older of the two

names, should provide the correct epithet at the rank of species. The treatment of

Ancistrocladus sagittatus as a synonym of Tetramerista glabra has been repeated in

subsequent revisions of Ancistrocladus (Steenis, 1948; Gereau, 1997; Taylor et al.,

2005) and Tetramerista (Lim, 2010).

The online catalogue of the Kew Herbarium made it easy to discover that the

recent detenninations of the Wallich specimen in the general herbarium were under

the myrtaceous genus Tristaniopsis (formerly Tristania) rather than Tetramerista. The

two genera bear superficially similar foliage. However, Tetramerista lacks a clear

intramarginal nerve (Keng, 1989), which is evident in the Wallich specimens at Kew,

leading me to agree with the placement in Tristaniopsis. Following the treatment of

Tristaniopsis by Ashton (2005, 2011), I therefore comi&QX Ancistrocladus sagittatus to

be a new synonym of Tristaniopsis merguensis Griff.

Tristaniopsis merguensis (Griff.) Peter G.Wilson & J.T.Waterh., Austral. J. Bot. 30:

439 (1982). - Tristania merguensis Griff, Account Bot. Coll. Cantor 18 (1844-1845).

TYPE: Burma, Mergui, W. Griffith 235 (isotypes K [barcode nos. K000793713,

K000793713]).

Ancistrocladus sagittatus Wall. [Numer. List no. 1055 (1829), nom. nud.] ex Planch.,

Ann. Sci. Nat., Bot. ser. 3, 13: 319 (1849), synon. nov. - Tetramerista glabra Miq. var.

sagittata (Wall, ex Planch.) A. W.Benn. in Hook.f , FI. Brit. India 1: 526 (1875). TYPE:

Singapore, 1822, N. Wallich s.n. [EIC 1055] (lectotype K [barcode no. K000793725],

designated here; isolectotypes K-W [barcode nos. KOOl 110861, K001110862]).

Tristanopsis merguensis in Singapore

As well as dealing with the correct placement of the namQAncistrocladus sagittatus, this

finding also brings into question the occun'ence of Tetramerista glabra in Singapore.

The species has been listed in the Singapore flora (Ridley, 1900; Keng, 1990) based

solely on the Wallich record which we now know to be an error of identification. The

work of Comer (1978) on the swamp forests of Johore and Singapore supports the

absence of Tetramerista glabra from Singapore. He found the species in the tme peat

swamp relict at Pontian in Johore, but not in the freshwater swamps in the Sedili forests

of Johore or Singapore. 1 therefore consider that there is no record of Tetramerista

glabra from Singapore and the species, genus and family all have to be removed from

listings of the native flora until evidence to the contrary is forthcoming.

References

Ashton, P.S. (2005). New Tristaniopsis Peter G. Wilson & J.T.Waterh. (Myrtaceae) from

Borneo. Gard. Bull. Singapore 57: 269-278.

Ashton, PS. (2011). Myrtaceae. In: Soepadmo, E., Saw, L.G., Chung, R.C.K. & Kiew, R.

(eds) Tree Flora of Sabah and Sarawak 7: 87-330. Kepong: Forest Research Institute

Malaysia.

Bennett, A.W. (1875). Ochnaceae. In: Hooker, J.D. (ed) Flora of British India 1: 523-527.

London: L. Reeve & Co.

Comer, E.J.H. (1978). The freshwater swamp-forest of South Johore and Singapore. Gard.

Bull. Singapore, Suppl. I: 1-266.

Gereau, R.E. (1997). Typification of names in Ancistrocladus Wallich (Ancistrocladaceae).

Novon 7: 242-245.

Keng, H. (1989). Tetrameristaceae. In: Ng, F.S.P. (ed) Tree Flora of Malaya 4: 470^71.

Petaling Jaya: Longman Malaysia Sdn. Bhd.

Keng, H. (1990). The Concise Flora of Singapore: Gymnosperms and Dicotyledons. Singapore:

Singapore University Press.

Lim, C.L. (2010). Tetrameristaceae. In: Kiew, R., Chung, R.C.K., Saw, L.G., Soepadmo, E. &

Boyce, PC. (eds) Flora of Peninsular Malaysia, Series II Seed Plants (Malayan Forest

Records 49) 1 : 265-269. Kepong: Forest Research Institute Malaysia.

Planchon, J.-E. (1849). Essai monographique d’une nouvelle famille de plantes propose sous le

nom d’Ancistrocladees. Ann. Sci, Nat., Bot. sen 3, 13: 316-320.

Ridley, H.N. (1900). The flora of Singapore. J. Straits Branch Roy. Asiat. Soc. 33: 27-196.

Steenis, C.G.G.J. van (1948). Ancistrocladaceae. In: Steenis, C.G.G.J. van (ed) Flora

Malesiana, Series 1, 4: 8-10. Jakarta: Noordhoff-Kolff N.V.

Taylor, C.M., Gereau, R.E. & Walters, G.M. (2005). Revision of Ancistrocladus Wall.

(Ancistrocladaceae). Missouri Bot. Gard. 93: 360-399.

Wallich, N. (1828-1849). A Numerical List of Dried Specimens of Plants in the East India

Company’s Museum, Collected Under the Superintendence of Dr. Wallich of the

Company’s Botanic Garden at Calcutta. London: Honourable East India Company.

Gardens’ Bulletin Singapore 67(1): 39-43. 2015

doi: 10.3850/S238258 12 15000046

The clarification and synonymisation of two taxa of

Vittaria from Peninsular Malaysia and a new combination

in Haplopteris (Pteridaceae subfam. Vittarioideae)

S. Lindsay^ & D J. Middleton^

'Gardens by the Bay, 18 Marina Gardens Drive,

Singapore 018953

stuart.lindsay@gardensbythebay.com.sg

^Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, Singapore 259569

david_middleton@nparks .gov. sg

ABSTRACT. A variety and a species of Vittaria in Peninsular Malaysia are synonymised and

the new combination Haplopteris sessilifrons (Miyamoto & H.Ohba) S. Linds, is made.

Keywords. Adiantaceae, Haplopteris sessilifrons, Vittaria ensiformis var. latifolia, Vittaria

sessilifrons, Vittariaceae

Introduction

As research towards an account of Adiantaceae for the Flora of Peninsular Malaysia

is progressing, papers are being published to clarify the correct generic placement of

a number of species (Lindsay, 2010; Lindsay & Chen, 2014). Wider discussions on

the current delimitation of the Asian genera formerly placed in the family Vittariaceae

(most often treated as part of Pteridaceae subfam. Vittarioideae but to be included in

Adiantaceae for the Flora of Peninsular Malaysia) can be found in Crane (1998),

Lindsay (2004), Ruhfel et al. (2008) and Lindsay & Chen (2014).

This current paper concerns a species described by Miyamoto & Ohba (1992),

Vittaria sessilifrons Miyamoto & H.Ohba, and a variety written about by Holttum

(1955), Vittaria ensiformis var. latifolia,

Miyamoto & Ohba (1992) described Vittaria sessilifrons based on several of

their own collections from Pahang in Peninsular Malaysia. We had been unable to

locate any of the isotypes or paratypes, said to be in a number of herbaria, until it

was discovered that none of the type material had been distributed. This has now

been rectified and SING herbarium received an isotype and two paratypes in 2014.

Examination of this material has revealed it to be the same taxon previously written

about as Vittaria ensiformis var. latifolia by Holttum (1955). Unfortunately, Holttum’s

variety was not validly published due to the lack of a Latin diagnosis or description

(Art. 39.1; McNeill et al. 2012).

We had earlier delayed making a decision on what to do with the material

referable to Vittaria ensiformis var. latifolia, due to its apparent intermediate position

Gard. Bull. Singapore 67(1) 2015

between Haplopteris ensiformis (Sw.) E.H.Crane and H. elongata (Sw.) E.H.Crane

(formerly Vittaria ensiformis Sw. and V. elongata Sw.). Until the additional three

specimens were received from Japan, we had seen only five specimens that could

be referred to this taxon (three annotated by Holttum plus two others: Mohd Shah &

Samsuri MS3822 (SING) and R. Kiew RK1989 (KEP)). We remained unsure as to

whether it blun'ed the distinction between Haplopteris ensiformis and H. elongata,

whether it should be described as a variety of one or the other (but leaning towards H.

elongata), or whether it should be described as a new species (before we discovered

that it was already described as Vittaria sessilifrons). Holttum (1955) made it a variety

of Vittaria ensiformis, based on its sessile fronds, but noted its similarities to V.

elongata. Miyamoto & Ohba (1992), however, compared their new species to Vittaria

elongata and V scolopendrina (Bory) Sclikuhr ex Thwaites & Hook, (now Haplopteris

scolopendrina (Bory) C.Presl) although noted it ‘differs greatly’ from the latter.

With the availability of more material, now covering a broader geographical

distribution in Peninsular Malaysia, it has become clear that the characters which

define this taxon are quite stable and that Miyamoto & Ohba (1992) were quite correct

to describe it at the rank of species. It is similar to Haplopteris ensiformis in having

sessile fronds and sori in deep marginal or submarginal grooves but differs in having

a distinct costa above (at least in the lower half of the frond) and a wider and thinner

lamina in which the lateral veins are visible (at least with transmitted light). It is similar

to Haplopteris elongata in the width and texture of the lamina, the visibility of the

venation and the position and structure of the sori, but differs in having sessile fronds

and costae that are more strongly raised above (the costae of H. elongata are usually

distinct but flat or hardly raised above). It is similar to Haplopteris scolopendrina in

the texture of the lamina and the visibility of the venation but differs in having much

narrower and shorter fronds and sori in deep marginal or submarginal grooves (the sori

of H. scolopendrina are in shallow and broad submarginal grooves).

Based on the arguments to be found in Crane (1998) and Lindsay (2004), Vittaria

sessilifrons now requires a combination in Haplopteris which is here provided.

New combination

Haplopteris sessilifrons (Miyamoto & H.Ohba) S.Linds., comb. nov. - Vittaria

sessilifrons Miyamoto & H.Ohba, Acta Phytotax, Geobot. 43(1): 33, f. 2 (1992). -

TYPE: Peninsular Malaysia, Pahang, Gunung Tahan, Sungai Luis, alt. 700 m, 19

March 1990, Ohba, H. d Miyamoto, F 900536 (holotype Tl!; isotypes FRI, L, US,

SING!).

Vittaria ensiformis var. latifolia Holttum, Rev. FI. Malaya 2: 614 (1955 [‘1954’]),

nom. inval.

Epiphytic or lithopyhtic. Rhizome short-creeping, usually obscured by a mass of

extremely hairy roots, c. 1-2 mm diameter (in the dry state), bearing fronds close

New combination in Haplopteris

together, densely scaly. Scales linear, stiff, straight, gradually narrowing from a

cordate base towards a long-tailed and ultimately filifonn apex, up to c. 6 mm long and

c. 0.7 mm wide (at base), clathrate, sparsely toothed at margin particularly near base

and somewhat bicoloured, the walls of the inner cells being thicker and darker than

those of the outer cells (most noticeable in the section of the tail that is two cells wide;

the common wall in the middle is thick and black while the two marginal walls are

thinner and somewhat red). Fronds pendulous, simple, linear, 9^0 cm long, 0.45-1 .3

cm wide, gradually nan*owing towards both ends, the apex narrowly acute, the base

sessile. Lamina coriaceous, glabrous, the margins revolute almost the entire length

of the frond, midrib and margins moderately to strongly raised on the upper surface

particularly in the lower half of the frond, the midrib usually becoming flatter towards

the frond apex but the margins remaining raised throughout, midrib flat and indistinct

below. Lateral veins more-or-less visible from above (or with transmitted light),

indistinct below, simple, very oblique, parallel, joined by a continuous submarginal

vein. Sori very long, linear, uninterrupted, appearing to be marginal (but actually arising

from the submarginal vein), in a deep, two-lipped, groove in the revolute frond margin

(therefore only visible from below), the two lips more-or-less equal. Sporangia with

annuli composed of 16-18 cells. Soral paraphyses copious, their stalks multicellular,

filiform, mostly colourless; their apical cells large, funnelform, twice as long as wide

or longer, reddish-brown. Stint ctures resembling sporangial stalks (but without

sporangia attached) more numerous than paraphyses. Spores bilateral, bean-shaped,

monolete, pale, translucent, smooth.

Distribution. Peninsular Malaysia and, possibly, Borneo. We have seen specimens from

Johor, Negeri Sembilan, Pahang and Terengganu (close to the border with Kelantan)

in Peninsular Malaysia. Holttum (1955) lists Perak too. We have not verified material

from Borneo listed as Vittaria ensiformis var. latifolia in online herbarium specimen

databases.

Ecology. Information is scant. However, three collections are described as epiphytic

on mossy tree-trunks near streams in forests, one as epiphytic 3 m from the ground

above a river, and one as growing on rocks. The lowest recorded altitude is 200 m and

the highest is 762 m.

Provisional lUCN consen^ation assessment. Data Deficient (DD). Although not often

collected, this species is widespread in Peninsular Malaysia which would suggest an

assessment of Least Concern. However, of the collections known, several are more

than 100 years old and none have been collected since 1990.

Additional specimens examined. PENINSULAR MALAYSIA: Johor: Kota Tinggi, [locality

illegible], 1910 [month illegible], Ridley, H.N. s.n. (SING [SING0033264]); Mount Austin,

[1901?; date illegible], Ridley, H.N. s.n. (SING [SlNG008566i]); Sungai Salat, Ulu Endau,

Sep 1985, Kiew, R. RKJ989 (KEP). Negeri Sembilan: Gunung Angsi, alt., 2500 ft. [762 m], 22

Nov 1923., M/r. Md SFN11617 (SING). Pahang: Gunung Tahan, Sungai Juram, alt. 200 m, 11

Gard. Bull. Singapore 67(1) 2015

Mar 1990, Ohba, H. & Miyamoto, F 900120 (SING, TI); Smigai Juram to Sungai Luis, alt. 700

m, 13 Mar 1990, Ohba, H. & Miyamoto, F. 900208 (SING, Tl). Terengganu: Gunung Ayam

via Ulu Besut, alt. 2,400 ft. [732 m], 5 Mar 1976, Shah, Mohd & Samsuri MS3822 (SING).

Notes. The holotype and SING isotype are clearly labelled as having been collected on

19 March 1990; the date in the protologue of 13 March 1990 is a printing error.

In his key to the Malayan species of Vittaria, Holttum (1955: 608) says of

Vittaria ensiformis var. latifolia that the “midrib [is] never strongly raised on [the]

upper surface”. This statement is at odds with the protologue of Vittaria sessilifrons

which says “costa and margin raised on upper side” and illustrates both as being rather

strongly raised (see Miyamoto & Ohba, 1992, Fig. 2E & G). We have examined three

specimens annotated by Holttum and, in each, the costa and margins are moderately to

strongly raised on the upper surface of the basal half of the frond.

Miyamoto & Ohba (1992) state that Vittaria sessilifrons differs greatly from V

scolopendrina in having a raised costa. This implies that Flaplopteris scolopendrina,

using the cuiTent name, does not have a raised costa when in reality it does. The costa

of Haplopteris scolopendrina is, in fact, very strongly raised on the upper surface of

the frond (even more so than in Flaplopteris sessilifrons) for almost the entire length

of the frond.

The illustration of the paraphyses in the protologue of Vittaria sessilifrons shows

them with ovoid or clavate heads but we have not seen any paraphyses of this sort in

the type material or in any other material. Instead, all material that we have examined

has paraphyses with funnelform apical cells, a shape that is typical for Haplopteris. As

well as these paraphyses, all of the specimens of Haplopteris sessilifrons have a very

large number of structures that look like sporangial stalks without sporangia attached.

It is unclear whether these are an artefact of dried specimens or if, as we suspect, these

develop without ever producing sporangia. Similar structures are sometimes found in

both Haplopteris elongata and H. ensiformis but never so densely.

ACKNOWLEDGEMENTS. We thank Tetsuo Ohi-Toma (TI), Akiko Shimizu (TI), Futoshi

Miyamoto (Tokyo University of Agriculture) and Serena Lee (SING) for arranging the transfer

of type material from TI to SING. We also thank Jana Leong-Skomickova for her valuable

comments and for guiding this paper through the review process.

References

Crane, E.H. (1998 [‘1997’]). A revised circumscription of the genera of the fern family

Vittariaceae. Syst. Bot. 22(3): 509-517.

Holttum, R.E. (1955 [‘1954’]). A Revised Flora of Malaya, edition 1, vol. 2 Ferns of Malaya.

Government Printing Office, Singapore.

Lindsay, S. (2004). Considerations for a revision of the fern family Vittariaceae for Flora

Malesiana. Telopea 10(1): 99-112.

New combination in Haplopteris

Lindsay, S. (2010). New combinations m Haplopteris (Adiantaceae) for the Flora of Peninsular

Malaysia. Gard. Bidl. Singapore 62(1): 119-120.

Lindsay, S. & Chen, C.W. (2014). Three new combinations m Haplopteris (Pteridaceae subfam.

Vittarioideae). Gard. Bull. Singapore 66(2): 169-171.

McNeill, J., Barrie, F.R., Buck, W.R., Demoulin, V., Greuter, W., Flawksworth, D.L., Herendeen,

P.S., Knapp, S., Marhold, K., Prado, J., Prud’homme van Reine, W.F., Smith, G.F.,

Wiersema, J.H. & Turland, N.J. (2012). International Code of Nomenclature for algae,

fungi and plants (Melbourne Code). Regnum Vegetabile 154. 205 pp. Konigstein: Koeltz

Scientific Books.

Miyamoto, F. & Ohba, H. (1992). Two new ferns from Gunung Tahan, Malay Peninsula. Acta

Phytotax. Geobot. 43(1): 31-35.

Ruhfel, B., Lindsay, S. & Davis, C.C. (2008). Phylogenetic placement of Rheopteris and the

polyphyly of Monogramma (Pteridaceae s.l.): Evidence from rbcL sequence data. Syst.

Bot. 33(1): 37^3.

Gardens’ Bulletin Singapore 67(1): 45-49. 2015

doi: 10.3850/S2382581215000058

Bothriospermum (Boraginaceae), a new generic record for

the Flora of Peninsular Malaysia

A.R. Rafidah, S. Syahida-Emiza, R. Kiew & K. Imin

Forest Biodiversity Division, Forest Research Institute Malaysia (FRIM),

52109 Kepong, Selangor, Malaysia

rafidahar@fi'im.gov.my

ABSTRACT. Bothriospermum zeylanicum (J.Jacq.) Druce (Boraginaceae), a species native

to Asia, has recently become naturalised in Cameron Highlands, Pahang, Malaysia. A

description of the genus and species together with colour photographs are provided. It is the

only representative of Boraginaceae sensu stricto (i.e. subfam. Boraginoideae) in Peninsular

Malaysia.

Keywords. Boraginaceae, Bothriospermum, naturalised weed, new genus record, taxonomy,

Malaysia

Introduction

In 2007 a fieldtrip to Cameron Highlands, Pahang, Penmsular Malaysia led to the

discovery of a species of Boraginaceae that was tentatively identified as a Myosotis, a

genus not otherwise recorded from Peninsular Malaysia. Realising it was something

new, a second fieldtrip was organised in April 2014 to obtain more material and to take

photographs so that it could be correctly identified. It proved to be Bothriospermum

zeylanicum, which is a new record for the genus and species for Peninsular Malaysia;

the genus is not recorded in either the Flora Malesiana revision of the Boraginaceae

(Riedl, 1997), nor in the checklist by Turner (1997). New weeds, often of temperate

species, are regularly introduced into the Cameron Highlands, where extensive

vegetable and flower farms have replaced the lower montane forest. For example,

Kiew (2009) recently reported 17 new records of naturalised weed species, of which

ten were from Cameron Highlands. Interestingly, Bothriospermum zeylanicum occurs

at a lower altitude (c. 1200 m) than most of these other weeds. A ftdl morphological

description of the genus and species, together with colour photographs, is provided

here.

Turner (1997), in his catalogue of Peninsular Malaysian plants, listed under

Boraginaceae 16 species in 9 genera: Argusia (1 species), Carmona (1 species),

Coldenia (1 species), Cordia (4 species), Ehretia (4 species), Heliotropium (1

species), Pteleocarpa (1 species), Rotula (1 species) and Tournefortia (2 species).

Recent molecular work by Refulio-Rodriguez & Olmstead (2014) separated Cordia

and Coldenia into Cordiaceae; Ehretia, which now includes Carmona and Rotula

(Gottschling et ah, 2014), into Ehretiaceae; 'md Argusia, Heliotropium and Tournefortia

Gard. Bull. Singapore 67(1) 2015

into Heliotropiaceae. Pteleocarpa is now placed in Gelsemiaceae (Struwe et al.,

2014). In other words, all the genera listed by Turner under Boraginaceae have been

assigned to other families meaning that now the Boraginaceae s.s. is represented in

Malaysia only by this new record, Bothriospermum zeylanicum. Even if the narrow

family circumscription adopted by Refulio-Rodriguez & Olmstead (2014) is not

followed and one prefers to recognise Cordiaceae, Ehretiaceae and Heliotropiaceae

as subfamilies Cordioideae, Ehretioideae and Heliotropoideae (e.g. Engler & Prantl,

1897; APG III, 2009), it remains true that Bothriospermum zeylanicum is the only

Malaysian representative of the type subfamily Boragmoideae.

Taxonomic treatment

Bothriospermum Bunge, Enum. PI. China 47 (1833); Clarke, FI. Brit. India 4: 167

(1885); Backer & Bakhuizen /, FI. Java 2: 463 (1965); Zhu et al., FI. China 16: 418

(1995); Riedl, FI. Malesiana ser. I, Spemiat. 13: 63 (1997); Hsiao & Liu, FI. Taiwan 4:

387 (1998). - TYPE: Bothriospermum chinense Bunge.

Annual or biennial herbs, hispid or pubescent. Stems erect or prostrate. Leaves spirally

arranged; lamina elliptic, ovate or oblanceolate, margin entire. Flowers extra-axillary,

small, solitaiy but often fonning a leafy raceme in distal part of the branches, distinctly

pedicellate; calyx 5-lobed, the lobes divided to the base, narrowly lanceolate, slightly

enlarged in fruit; corolla blue or white, rotate, tube short, lobes 5, spreading, obtuse,

5 small scale-like appendages present in the throat. Stamens included in the corolla,

filaments very short, anthers 5, ovoid. Ovary deeply 4-lobed, style short, persistent,

stigma small, capitate; gynobase flat. Fruits-, nutlets 4, veiy small, erect, ellipsoid or

subglobose attached to the flat or nearly flat receptacle, surface usually sculptured,

ventral side with a large aperture surrounded by prominent margin.

Distribution. About 5 species, distributed in Afghanistan, Kazakhstan, Kyrgyzstan,

Tajikistan, Turkmenistan, Uzbekistan, Russia, Pakistan, India, China (its centre

of distribution), Japan, Korea, Taiwan, Vietnam and the Philippines; naturalised in

Indonesia (Java).

Bothriospermum zeylanicum (J.Jacq.) Druce, Bot. Exch. Club Brit. Isles 4: 610

(1917); Zhu et al., FI. China 16: 419 (1995); Riedl, FI. ser. I, Speraiat. 13: 63 (1997);

Hsiao & Liu, FI. Taiwan 4: 388 (1998). -Anchusa zeylanica J.Jacq., Eel. PI. Rar. 1(3):

47, t. 29 (1812). - TYPE: Plate t. 29 in Jacquin, Eel. PI. Rar. 1(3) (1812). (Fig. 1 & 2)

Anchusa tenella Homem., Hort. Bot. Hafii. 1: 176 - Bothriospermum tenellum

(Homem.) Fisch. & C.A.Mey., Index Sem. Hort. Petrop. I: 23 (1835); Clarke, FI. Brit.

India 4: 168 (1885); Backer & Bakliuizen, f, FI. Java 2: 463 (1965). - TYPE: Herb.

Vahl s.n. ‘Hab. in China’ (holotype C n.v.).

Bothriospermum, a new generic record for Peninsular Malaysia

Fig. 1. Bothriospermum zeylanicum (JJacq.) Druce. A-B. Habit. C. Arrangement of flowers

and fruits. D. Nutlet. E. Leaf F. Flower. Scale bars: A, B, C (5 cm); D (1 mm); E (2 mm); F (5

mm). (Photos: A.R. Rafidah)

Gard. Bull. Singapore 67(1) 2015

Fig. 2. Habitat of Bothriospermum zeylanicum (JJacq.) Druce. (Photo: A.R. Rafidah)

Short-lived herbs with prostrate stems. Stems slender, densely pubescent throughout,

much branched, up to 50 cm long. Leaves: petiole short, c. 5 mm long in lower leaves,

lacking in the upper; lowermost laminas 20-30 x 6-10 mm, decreasing in size to 4-10

X 2-4 mm in the upper ones, elliptic to lanceolate, margin slightly undulate, densely

strigose on both surfaces, base cuneate, midrib impressed above, prominent beneath,

apex acute. Pedicels c. 2 mm long. Flowers: calyx green, densely hairy outside;

corolla white or very pale purple (in Peninsular Malaysia), c. 2 mm long, lobes 5,

longer than the tube, c. 2 mm long, apex broadly rounded, throat appendages white,

trapeziform, emarginate, c. 0.2 mm. Stamens 5, included, inserted at the middle of

corolla tube, filaments short, anther yellowish turning brown. Ovary green, style less

than 1 mm long, terete. Nutlets green turning brown, ellipsoid, to 1 mm long, aperture

longitudinally elliptic, surface warty; calyx persistent, c. 3 mm long.

Distribution. As for the genus, in Peninsular Malaysia only from Cameron Highlands,

Pahang.

Ecology. Growing in waste land on sandy soil or in open fields or near rivers, at

c. 1200 m altitude.

Bothriospermum, a new generic record for Peninsular Malaysia

Specimens examined. PENINSULAR MALAYSIA: Pahang: Cameron Highlands, Belati

Estate, 4°27'N 101°28'E, 8 Oct 2007, Kiew et al, FRI70433 (KEP), Kampung Terla, 4°31'N

I01°23'E, 8 Oct 2007, Kiew et al, FRI70473 (KEP), Habu, 4°26'N I0I°23'E, 23 Apr 2014,

Rqfidah etal, FR175968 (KEP), Habu, 4°26'N I01°23'E, 23 A^x2(dU,Rafidah etal, FRI75969

(KEP).

ACKNOWLEDGEMENTS. This research was carried out as part of the Documentation and

Inventory Flora Malaysia project (Flora of Peninsular Malaysia) funded by the Ministry of

Science, Teclmology and Innovation at the Forest Research Institute Malaysia (FRIM), Kepong.

We are grateful to the curators and staff of the herbaria at BO, K and KEP for permission to

examine specimens in their care and for providing information and images. We thank Dr Saw

Leng Guan and Dr Richard Chung for their helpful comments and two anonymous reviewers

for their valuable comments on the manuscript.

References

APG III. (2009). An update of the Angiospenn Phylogeny Group classification for the orders

and families of flowering plants: APGIII. Bot. J. Linn. Soc. 161: 105-121.

Engler, H.G.A, & Prantl, K. (1897). Die NatiirJichen Pflanzenfamilien. Leipzig: W. Engelmann.

Gottschling, M., Luebert, F., Hilger, H.H. & Miller, J.S. (2014). Molecular delimitations in the

Ehretiaceae ( Boraginales). Mo/^c. Phylogenet. Evol. 72: 1-6.

Kiew, R. (2009). Additions to the weed flora of Peninsular Malaysia. Malayan Nat. J. 61:

133-142.

Refulio-Rodriguez, N.F. & Olmstead, R.G. (2014). Phylogeny of Lamiidae. Amer. J. Bot.

101(2): 287-299.

Riedl, H. (1997). Boraginaceae. Flora Malesiana, Ser. 1, 13; 43-144.

Struwe, L., Soza, V.L, Sugumaran, M. & Olmstead, R.G. (2014). Gelsemiaceae (Gentianales)

expanded to include tire enigmatic Asian genus Pteleocarpa. Bot. J. Linn. Soc. 175:

482-496.

Turner, I.M. (1997 [‘1995’]). A catalogue of the vascular plants of Malaya. Card. Bull.

Singapore 41: 1-757.

Gardens’ Bulletin Singapore 67(1): 51-60. 2015

doi: 10.3850/S238258121500006X

Novitates Bruneienses, 1. A background to the botanical

survey of Brunei Darussalam, and a new species of

Jarandersonia (Malvaceae)

A. A. Joffre^, A.K. Muhammad Ariffin^, Y.W. Low^ & K.M. Wong^

^Brunei National Herbarium, Forestry Department,

Ministry of Industry and Primary Resources, Jalan Menteri Besar,

Berakas, BB3910 Brunei Darussalam

^Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, 259569 Singapore

wkm2000@gmail.com

ABSTRACT. A brief introduction to research and botanical documentation of the Brunei

flora, and the collaborative programme for a continuing botanical survey of the country, is

given. An outline of the key biogeographical features of the Brunei area supports the premise

that distinct geo-ecological enclaves occur which are special units of species richness, within

which a signiflcant level of undocumented plant diversity still exists. Jarandersonia yahyantha

K.M. Wong, Joffre, Ariffin & Y.W.Low (Malvaceae), a new tree species so far only known in

Brunei, is described.

Keywords. Biological hotspot, biogeography, Borneo, Malesia, new species. Southeast Asia

Introduction: The floristic richness of Borneo and Brunei Darussalam

This new series supports a recently established programme to further the botanical

exploration of Brunei Darussalam, in northwestern Borneo, the third-largest island in

the world and biologically the richest landmass of the Sundaland region (MacKinnon

& MacKinnon, 1986; Ashton, 1989). The Malesian region as a whole (Malay

Peninsula and Malay Archipelago, including the area from the Philippines through to

New Guinea) has a diverse flora estimated to include 42,000 seed plant species (Roos,

1993). Myers et al. (2000) considered Sundaland, which includes the Malay Peninsula,

Sumatra, Java and Borneo, one of tropical Southeast Asia’s four biodiversity hotspots.

Estimates of the Bornean vascular flora have ranged from 9000 (Merrill, 1921)

to 15,000 species (Merrill, 1950; Raes et al., 2009). However, although recent work

completed by Beaman & Anderson (2004) enumerated more than 5000 vascular plant

species in Kinabalu Park (Sabah) alone, further work has suggested that many more

novelties were to be expected there (Chen et al., 2014), as well as generally with

the Tree Flora of Sabah and Sarawak (e.g., Soepadmo & Wong, 1995 and further

volumes). As such, Merrill’s upper limit for Bomeo, or a tally in excess of that, is to

be expected. On the other hand, Brunei, with its land area of 5765 knP being less than

a mere 1% of the whole of Bomeo, was estimated to have close to 5000 species of

Gard. Bull. Singapore 67(1) 2015

seed plants, including 2000 species of tree (Wong, 1997). A first intensive checklisting

of the flora based on organised field collecting over some seven years, 1989—1995,

much of it an active collaboration between the Brunei Forestry Department and the

Royal Botanic Gardens, Kew (Coode et ah, 1996), accounted for some 3500 species

of indigenous seed plants, including 1900 tree species (Wong, 1999). Apart from the

Dipterocarpaceae, which has been well treated by Ashton (1964, 1 982) and is generally

well collected, this checklist remains a first-stage investigation of the flora because (1)

many families and genera still had significant numbers of undetennined specimens;

(2) recent critical revisions were not available then for many genera; and (3) in the

ensuing period the Brunei Herbarium continued to gather interesting taxa that had not

been captured by earlier collecting effort. Thus it can be appreciated that the Brunei

flora is a very rich one.

In 2013, the Government of His Majesty The Sultan of Brunei Damssalam

and the National Parks Board, Singapore, entered into a collaborative programme for

a new phase of botanical survey. This seeks to intensify fieldwork for biodiversity

exploration and documentation, and to focus on the botanical inventory of Brunei,

including taxonomic identifications by specialists from our own and outside institutions

when possible.

Key biogeographical features of the Brunei area

In the same year as this new collaboration began, a review of the flowering plants

endemic to Brunei was published (Henrot et al., 20 1 3), listing 65 species of angiospemis

hitherto known only from Brunei, and suggesting that, as with a number of other species

generally occurring only in the Brunei area (which includes adjacent parts of SW Sabah

or NE Sarawak), some of the so-called endemics could subsequently be found to be

not so restricted. The latter category is exemplified by the aroids Homalomena scutata

S.Y.Wong & P.C. Boyce, occuning in both the Belait and Temburong districts of

Brunei but found also in the Limbang and Mulu areas in adjacent NE Sarawak (Wong

& Boyce, 2014), and H. cowleyae RC, Boyce & S.Y.Wong, known in the Temburong

and Mulu areas (Boyce & Wong, 2014),

There are at least two biogeographic scenarios why this work on endemic

plants is significant. One is that we could indeed expect there would be true Brunei

endemics, as botanical experience has shown for, especially, herb species, including

many Begonia or gesneriad taxa, as well as smaller plants in general. Henrot et al.

(2013) found that most of the documented endemic taxa were smaller plants in the

Begoniaceae (15 taxa), Araceae (8 taxa), Gesneriaceae (7 taxa), Orchidaceae (5 taxa),

Zingiberaceae (5 taxa), and palms (Arecaceae: 5 taxa), so it could be true that many

large flowering plants have better dispersal than small ones. Although it should be

borne in mind that, thus far, the collecting and research effort has concentrated much

on trees, rather than herbs, other reasons, including recent evolution, may restrict

plant distribution. A slender bamboo, Temhurongia simplex S.Dransf. & K.M.Wong,

is the only known representative of this peculiar genus (Dransfield & Wong, 1996)

A new Jarandersonia from Brunei Darussalam

that is still entirely restricted to Brunei’s Temburong valley, even after two decades

of continuing botanical collecting in Borneo. There is a suite of similarly restricted

Begonia and other species. Some others are borderline cases: for instance, the palm

Livistona exigua Dransf, documented from Bmnei’s Batu Patam ridge, still has not

been documented from adjacent parts in Sarawak.

Wong (1997, 1 999) discussed how the Brunei area includes its own geographical

and ecological enclaves within the geologically distinct (but relatively youthful) and

larger northwest Borneo area, taking into consideration how its main area is hemmed

in by either highlands (such as around inland Temburong, Tutong and Belait) or

extensive swamp systems (as in the lower Belait and Baram drainage complex to its

southwest) (Fig. 1). The western side of Brunei is largely the remains of an ancient

syncline, now drained by the Belait and Tutong rivers, whereas the eastern Temburong

district is mostly the drainage area of one major river system, the Temburong (Ashton,

1958). This, together with precipitous topogi*aphy marked by steeply incised valleys

in the hard sandstones of inner Temburong, or in the uppermost reaches of the Belait

and Tutong, can conceivably bring about population isolation that engenders highly

localised endemism (Coyne & Orr, 2004). Even in the Belait district, where topographic

relief is generally lower than in the Temburong, hilly pockets isolated by swamps or

river systems, or highly specific habitats such as moist sandstone bluffs, may harbour

narrowly endemic or habitat-specialist taxa, such as Homalomena spp. documented

only from the Teraja area of Belait (Wong & Boyce, 2011). Many of the flowering

plants endemic to Brunei Darussalam are either very rare or undercollected: 15 are

known from a single collection only and 83% are known from three sites or fewer.

The other significant dimension is that Bmnei has maintained maximal levels

of forest cover within the country (Bryan et ah, 2013). Borneo’s forest cover declined

from 71% of the island (mid-1980s) to 54% by 2000, an alarming rate of deforestation

(Stibig & Malingreau, 2003). With continuing rainforest degradation in the region,

Brunei Darussalam could continue to serve as a critical refuge for Bornean plants

impacted by such processes. Henrot et al. (2013) discussed the increasing importance

of ‘anthropogenic endemics’ in refuge in Brunei fi*om the wider Brunei area or beyond:

it is a scenario we would not wish to see exacerbated, but the significance of conserved

resources in Brunei becomes obvious. In upping the ante, the Brunei Ministry of

Industry and Primary Resources phased out loggmg in all Forest Reserves (previously

some had been classified for timber production) in 2014 (Brunei Times, 2014).

Continuing efforts

All this underscores how important it is now for authorities in Borneo to redouble

efforts into biodiversity inventory and conservation planning, in order that as many

specially diverse or unique areas as possible can be recognised in time and be

brought under effective management. For our part, it brings us back to the special

collaboration now in place between the Brunei National Herbarium (acronym BRUN;

Thiers, continuously updated) and the Singapore Herbarium (SING), and their partner

institutions, for continuing the Botanical Survey of Bmnei Damssalam.

Gard. Bull. Singapore 67(1) 2015

Fig. 1. The Brunei area: Key geological, tectonic and topographical features.

Uncoloured'. Pliocene - Miocene Sediments, in places overlain by Pleistocene or more recent

deposits. Yellow. Miocene - Oligocene Sediments. Brown: Rajang Group Deepwater Sediments

(Late Cretaceous - Eocene). Anticlines: Al. Belait anticline. A2. Jerudong anticline. A3.

Danau anticline. A4. Tangga anticline. A5. Mulu Uplift. Synclines: SI. Liku-Badas syncline.

S2. Belait syncline. S3. Limbang syncline. S4. Labu syncline. S5. Tangga syncline. Simple

line with arrows: Approximate boundary of synclinal basins within Brunei. PS. Peat body

or peatswamp vegetation over low-lying ground. Map prepared by Alsa Moo & K.M. Wong,

based on information from Liechti (1960), Wilford (1960) and Tate (2002).

In this series, we document new botanical findings in Brunei advanced through

our research or that of our collaborators: the new taxa discovered or diagnosed, the

new records of plants not previously documented in Brunei, and new and noteworthy

observations. It is our hope, given the indications from past and ongoing studies,

that the Brunei area continues to yield new and interesting plant taxa, and that such

a series will help focus due attention on Brunei’s floristic richness. As the present

paper shows, both newer (post Checklist, Coode et ah, 1996) as well as older (Ashton)

collections from Brunei are still important for our continuing studies, attesting to the

value of a good botanical archive in helping to attain further or newer biodiversity

A new Jarandersonia from Brunei Darussalam

documentation. All this is permitted only by sustained botanical effort and resources

over the longer time frame, by necessity trans-generational in scope.

A new species of Jarandersonia (Malvaceae)

Jarandersonia yahyantha K.M. Wong, Joffre, Ariffin & Y. W.Low, sp. nov.

This new species is most similar to Jarandersonia pannfolia Kosterm. in having fruits

with slender spines that are tuberculate and which bear tufted-hairy indumentum, but

differs in its subcordate to cordate (not cuneate to rounded) leaf bases and fruit spines

of 25-35 mm (not 10-15 mm) long. - TYPE: Brunei, Belait, Sg Liang, Andulau Forest

Reserve, Compartment 18, Labi Road, 10 April 2014, Ariffin, Jangarun & Raiizaidi

BRUN 24174 (holotype BRUN; isotypes A, K, KEP, L, SAN, SAR, SING). (Fig. 2)

Medium sized tree 10-25 m tall, trunk slightly fluted; bark smooth, pale grey brown.

Leaves broad-elliptic, 4.5-17(-22.7) cm long, 3.5-8(-11.5) cm wide; base subcordate

to cordate; apex acute to rounded to obtuse to slightly emarginate; midrib flat to slightly

sunken on upper surface, prominent on lower surface, secondary veins 6-8 pairs,

making angular loops near the leaf margin, the basal 1-2 pairs making a smaller angle

with the midrib than other secondaries (so resembling basal veins in a 3- to 5 -nerved

leaf base, but not reaching a fifth of the leaf length), flat to very slightly prominent

on upper side, prominent on lower side; glabrous on entire upper surface, with dense,

overlapping stellate scales completely occluding the lower surface; petioles 7-15 mm

long, 1.5-2 mm diameter, densely stellate lepidote. Inflorescences axillary, borne in

distal leaf axils, on young leafless imiovations or more proximal twig portions that

have lost their leaves, paniculate, to (l,5-)4-9,5 cm long, bearing (2-)4-9 flowers;

bracts ovate, acute, 2-4 mm long, 1.5-2 mm broad; flower pedicels (3-)5-9 mm long

(7-11 mm long in fresh material), scaly; calyx tube c. 2.5^ mm long and 2. 5-3. 5 mm

diameter (3^.5 mm long and 5-6 mm diameter in fresh material), scaly outside (scale

margins subentire-erose to short-fimbriate), lobes triangular, 2. 5-3. 5 mm long and

2. 5-3. 5 mm wide at base (6-7 mm long and 3-4 mm wide at base in fresh material),

scaly outside; petals 5, obovate, (5. 5-) 7-9 mm long and 3-6 nnm wide (16-18 nmi

long and 8-12 mm wide in fresh material), plane to slightly incurved, creamy white;

stamens many, filaments 3-7 mm long (7-1 5 mm long in fresh material), pale greenish

yellow, anthers knobby, c. 0.3 mm long, pale yellow; ovary subglobose, 5-lobed when

dried, 1-1.5 mm diameter (2-3 mm diameter in fresh material), pale greenish yellow,

short-hairy. Infructescence 3.5-13 cm long. Fruit subglobose, 6-10 mm diameter,

3 -locular, outside covered with slender soft spines 25-35 mm long, spines tuberculate

and bearing tufted hairs; seeds l(-2), ovoid-elipsoid, 17 mm long, 4 mm wide, dark

brown, smooth.

Distribution. The new species is apparently endemic to Brunei Darussalam. Of the

other species in the genus, only Jarandersonia parvifolia, which has a widespread

distribution in Sarawak, has been recorded in Brunei (Jaamat & Tachun FMS 39640,

KEP) in the Labi Forest Reserve.

Gard. Bull. Singapore 67(1) 2015

Fig. 2. Jarandersonia yahyantha K.M.Wong, Jof&e, Arififin & Y.W.Low. A. Tree in flower. B.

Flowering braneh. C. Fallen fiaiits (some germinating) and leaves. A & B from Ariffin et al. BRUN

24087; C from Ariffin et al. BRUN 24174. (Photos: A-B: Muhammad Ariffln, C: K.M. Wong)

A new Jarandersonia from Brunei Darussalam

Provisional lUCN conservation assessment. The provisional lUCN conservation

status for Jarandersonia yahyantha, assessed with the aid of GeoCAT (Bachman et ah,

2011), is Critically Endangered (CR B2ab(iii); D) (lUCN, 2012). The ‘B2’ designation

results from an area of occupancy (AGO) estimated to be less than 10 km^ (about 8

km2 for J. yahyantha); ‘a’ is due to a fragmented distribution as it occurs in only two

populations, namely at Bukit Jerudong and Andulau Forest Reserve; ‘b(iii)’ is due

to reductions in the area and quality of its habitat as J. yahyantha is so far known

only in the Compartments 7 and 18 of the Andulau Forest Reserve, which has some

disturbance along part of their boundary by adjacent road development, and on Bukit

Jerudong near Kampung Peninjau, which is previously disturbed forest; ‘D’ is due to

the small population size of only four mature individuals known so far.

Etymology’. The new species honours Pehin Orang Kaya Seri Utama Dato Seri Setia

Awang Haji Yahya bin Begawan Mudim Dato Paduka Haji Bakar, Brunei Darussalam’s

Minister of Industry and Primary Resources, for his interest in conservation and

leadership in ceasing all timber production from Bmnei’s forest reseiwes.

Additional specimens examined. BRUNEI. Belait: Simgai Liang, Andulau Forest Reserve,

Compartment 1 8, Labi Road, 200-300 m from roadside, 28 May 2014, Ariffin, Watu & Khairul

BRUN 24112 (A, BRUN, K, KEP, L, SAN, SAR, SING), 24112 (BRUN, K, SAN, SING).

Brunei-Muara: Jerudong, Kampung Peninjau, Bukit Jerudong, 11 Mar 2014, Ariffin, Watu,

Azlan & Khairul BRUN 24087 (BRUN, K, L, SAN, SAR, SING).

Notes. There are six known species of this genus (Tan et al., 201 1; Chung et al., 2012),

established by Kostermans (1960), in which the fruits are typically covered with

spines bearing setose hairs. Of these, the new species here is most like Jarandersonia

pentaceoides R.C.K.Chung & H.S.Tan, J. paimfolia and J. rinoreoides Kosterm. in

having slender soft fruit spines (the other species, J. clemensiae (Burret) Kostenn., J.

pursglovei (Kosterm.) Kosterm. and J. spinulosa Kosterm., have short stout and stiff

fruit spines.) The new species here differs from Jarandersonia pentaceoides in having

short basal secondary veins that do not reach even a fifth of the length of the leaf (in

that species the basal secondary veins are conspicuous in reaching halfway or more

along the leaf length) and fruit spines that are 25-35 mm long (the fruit spines in that

species are only 10-20 mm long). It differs from Jarandersonia rinoreoides in having

subcordate to cordate (not typically cuneate) leaf bases and fruit spines that are 25-35

mm (not 6-28 mm) long and short-tuberculate (rather than non-tuberculate); and from

J. parxAfolia as mentioned above in the diagnosis. Leaf size is not a particularly reliable

character because the flowering or fruiting material taken for herbarium specimens

mostly occurs on higher branches bearing smaller leaves, although larger leaf blades

may also be occasionally found on such branches.

Note that Tan et al. (201 1 ) and Chung et al. (2012) refer to the tuberculate spines

as “short-branched”. Also, the regularly polygonal fine vein areoles on the lower leaf

surface of Jarandersonia pentaceoides, which they described as “distinct honey comb-

like quaternary veins”, is not restricted to that species, being also found on occasion

in J. rinoreoides (e.g., Dewol SAN 99462 in SING); this feature is, however, absent

Gard. Bull. Singapore 67(1) 2015

from the leaves of the new species described here, in which the lower leaf surface is

completely occluded by dense, overlapping scales.

ACKNOWLEDGEMENTS. We would like to express our gratitude to Pehin Orang Kaya Seri

Utama Dato Seri Setia Awang Haji Yahya bin Begawan Mudim Dato Paduka Haji Bakar, the

Minister of Industry and Primary Resources, Brunei Darussalam for his interest and continuing

support of the botanical survey in Brunei. Special thanks are due to Hajah Nomiah Suria Hayati

binti Pehin Jamil (Permanent Secretary to the Ministry of Industry and Primary Resources),

Hajah Hasnah binti Haji Ibrahim (Deputy Permanent Secretary), Haji Saidin Salleh (Director

of Forestry), and Hajah Jamilah binti Haji Abdul Jalil (Acting Deputy Director of Forestry), as

well as Mahmud bin Haji Yussof, Chief Executive Officer of Brunei’s Heart of Borneo Centre.

We are likewise grateful for the mterest of past and present High Commissioners of Singapore

to Bmnei Damssalam, HE Joseph Koh and HE Jaya Ratnam, for facilitating the start of this

collaboration, which is jointly funded by both the Brunei Forestry Department and the National

Parks Board, Singapore. On the Singapore side, we thank the Ministry of National Development

and National Parks Board (NParks), in particular Kenneth Er, CEO of NParks; Dr Leong Chee

Chiew, Deputy CEO of NParks; Dr Nigel Taylor, Director of the Singapore Botanic Gardens;

and Dr David Middleton, Deputy Director and Keeper of the Singapore Herbarium (SING).

Many others have been most supportive and helpful to our programme: Roslinah binti Haji

Mohsin and Noralinda binti Haji Ibrahim (Forestiy Department, Brunei), Dr Kamariah Abu

Salim (Universiti Brunei Damssalam); Serena Lee, Paul K.F. Leong, Paul Pamsuraman Athen

and Alsa Moo (SING); Watu bin Awok, Jangamn anak Eri, Azlan bin Pandai, Muhanunad

Waffiuddin bin Zainol Ariffin, Mediana binti Kampong and Rauzaidi bin Mureh (BRUN). Not

least, the comments and suggestions of reviewers and the editor Dr David Middleton have

helped improve the text of this paper.

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Malaysia: Forest Research Institute Malaysia, Sabah Forestry Dept. & Sarawak Forestry

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Stibig, H.J. & Malingreau, J.P. (2003). Forest Cover of Insular Southeast Asia Mapped from

Recent Satellite Images of Coarse Spatial Resolution. Ambio 32 (7): 469^75.

Tan, H.S., Chung, R.C.K. & Soepadmo, E. (2011). A synopsis of Jarandersonia (Malvaceae -

Brownlowioideae). In: Wong, K.M., Leong-Skomickova, J., Lee, S. & Low, Y.W, (eds)

Proceedings of the 8th Flora Malesiana Symposium, Singapore, 23-27 August 2010.

Card. Bull. Singapore 63(1 & 2): 137-1 44.

Tate, R.B. (2002). The Geology^ of Borneo Island. CD-Rom. Kuala Lumpur: Geological Society

of Malaysia.

Gard. Bull. Singapore 67(1) 2015

Thiers, B. (continuously updated). Index Herbariorum: A global directory of public herbaria

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nybg.org/ih/ (accessed on 31 May 2014).

Wilford, G.L. (1960). Geological Map of Brunei and Adjacent Parts of Sarawak. Scale

1:125,000. Sheets I and II. Kuala Lumpur: Geological Survey Department, Federation

of Malaya.

Wong, K.M. (1997). In Brunei Forests. An Introduction to the Plant Life of Brunei Darussalam,

revised edition. 127 p. Kota Kinabalu: Natural History Publications & Brunei

Darussalam: Forestry Department.

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Trees and Forests of Brunei Darussalam. Pp. 53—73. Bandar Sri Begawan: University

of Brunei Darussalam, Forestry Department, Brunei 8l Brunei Shell Petroleum Co. Sdn.

Bhd.

Wong, S.Y. & Boyce, P.C. (2011 ). Studies on Homalomeneae (Araceae) of Borneo X: Two new

Homalomena species from Brunei, and a new infonual species complex for supergroup

Cyrtocladon. Webbia 66{2)\ 133-141,

Wong, S.Y. & Boyce, P.C. (2014). Studies on Homalomeneae (Araceae) of Borneo XVIII:

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44: 279-285.

Gardens’ Bulletin Singapore 67(1): 61-68. 2015

doi: 10.3850/S2382581215000071

Novitates Bruneienses, 2. A remarkable new species of

Begonia sect. Petermannia (Begoniaceae)

from Brunei Darussalam

Y.W. Low\ A. A. Joffre^ & A.K. Muhammad Ariffin^

'Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, 259569 Singapore

low_y ee_wen@nparks. go V. sg

^Bioinei National Herbarium, Forestry Department,

Ministry of Industry and Primary Resources, Jalan Menteri Besar,

Berakas, BB3910 Bmnei Damssalam

ABSTRACT. Begonia jamilahana Y.W.Low, Jofffe & Ariffin is described and illustrated

here as a new species based on a collection from Ladan Hills Forest Reserve, Tutong, Brunei

Damssalam. This new taxon is closely related to B. conniegeriae S. Julia & Kiew and B.

papyraptera Sands, but differs in a suite of morphological characters. Begonia jamilahana is

placed in Begonia section Petermannia, the largest of all the sections recognised.

Keywords. Begonia jamilahana, Begoniaceae, Borneo

Introduction

Begonia is a well-known genus, largely due to its popularity in horticulture. They

are mostly cultivated for their attractive foliage, as well as showy flowers (Holttum

& Enoch, 1995; Tebbit, 2005). At present, there are about 1500 species distributed

throughout the tropics, except in Australia, with many more taxa yet to be discovered

as individual species are commonly known to be confined to very small geographical

areas (Tebbit, 2005; Heywood et ah, 2007; Hughes, 2008).

Memll (1921), in his bibliographic enumeration, listed 26 species of Begonia

for Borneo, most of which were endemic species with the exception of B. repens

Blume. The checklist by Hughes (2008) enumerated 95 species for Borneo, all of

which are strictly restricted to the island. For Brunei, Coode et al. (1996) recorded 16

species, namely. Begonia awongii Sands, B. bahakensis Sands, B. baramensis Merr.,

B. bnmeiana Sands, B. chlorandra Sands, B. cyanescens Sands, B. eutricha Sands, B.

fuscisetosa Sands, B. hexaptera Sands, B. laccophora Sands, B. leucochlora Sands, B.

leucotricha Sands, B. papyraptera Sands, B. sibutensis Sands, B. stenogyna Sands and

B. tern burongen sis Sands. Four subspecies were recognised for Begonia bruneiana,

namely B. brimeiarm subsp. bruneiana, B. bruneiana subsp. angustifolia Sands, B.

bruneiana subsp. labiensis Sands and 5. bi^uneiana subsp. I'etakensis Sands. All of

the Brunei begonias were newly described by Sands (1996), with the exception of

Begonia baramensis Merr.

Gard. Bull. Singapore 67(1) 2015

A recent collection from the Ladan Hills Forest Reserve reveals yet another

novelty for Brunei from Begonia section Petermannia (Klotzsch) A.DC. as delimited

by Doorenbos et al. (1998). It has the following defining characters of the section,

namely (i) 2-tepaled male flowers, (ii) anthers with unilaterally positioned slits, (iii)

5-tepaled female flowers, (iv) inflorescence protogynous, with 1-2-flowered female

inflorescences, (v) trilocular ovai*y with axile placentation and bilamellate placentae,

and (vi) fruits with equal wings. The Ladan Hills novelty is closely related to Begonia

papyraptera, but differs in having (i) a smooth and terete stem in live material, (ii)

scantily scattered translucent hispid hairs on both sides of the leaf lamina, (iii) a mature

upper leaf surface that is dark green with red markings on vein axil nodes nearer to

the base of the leaf, (iv) a pale green lower leaf surface, and (v) base of capsule wings

cuneate. In contrast. Begonia papyraptera has (i) a ribbed and angular stem in live

material, (ii) a leaf lamina that is glabrous on both sides, (iii) a leaf upper surface that

is plain dark green with conspicuous white spots, (iv) a maroon lower leaf surface, and

(v) base of capsule wings rounded. Apart from Begonia papyraptera^ the Ladan Hills

taxon is also closely related to Begonia conniegeriae S. Julia & Kiew, but differs in the

characters enumerated in Table 1 . Hence, the Ladan Hills taxon is described here as

new.

New species

Begonia jamilahana Y.W.Low, Joffre & Ariffin, sp. iiov. (sect. Petermannia)

Similar to Begonia papyraptera Sands but differs in having a smooth and terete stem,

leaf blade scantily covered with franslucent hispid hairs on both sides, upper mature

leaf surface dark green with red markings on vein axil nodes nearer to the base of

the leaf, lower leaf surface pale green, and base of capsule wings cuneate. - TYPE:

Brunei, Tutong District, Lamunin, Ladan Hills Forest Reserve, Compartment 1, 58

m, 4°42’16”N 114°44’12”E, lowland mixed dipterocarp forest on yellow sandy clay

soils, steep slope, 15 Dec 2010, P. Azlan, A.K. Muhcl Ariffin, A. Watii & M. Rauzaidi

BRUN 23373 (holotype BRUN (including spirit material); isotypes K, SING). (Fig. 1,

2 )

Cane-like begonia, c. 40 cm tall, most parts of the plant glabrous except for the upper

and lower surface of lamina scantily covered with hispid translucent hairs neatly

arranged in between veins. Stems pale green, 0.8-1. 5 cm thick, very succulent, terete,

internodes 2.5-9 cm long, laxly branched, nodes red, not swollen. Stipules dark red,

narrowly lanceolate, c. 1-2 cm long, c. 0.3-0. 5 cm wide at the base, keeled, margin

entire, apex acute, caducous. Leaves alternate, oblique, distant, held horizontally;

petioles light green with both ends red, 5-20.5 cm long, terete; lamma dark green with

red markings on vein axil nodes nearer to the base of the leaf and iridescent above, pale

green beneath, young leaves very rarely with small white spots above, fading in older

leaves, asymmetric, elliptic to broadly elliptic, 12-19 x 11.5-14 cm, broad side 8-10.5

A new Begonia from Brunei Darussalam

cm wide, base cordate, basal lobes 3.5-7 cm, margin dentate, apex acuminate, acumen

1.5-2 cm long; venation palmate-pinnate, 7-9 veins, each branching dichotomously

2-3 times, vein nodes red near junction of petiole and gradually fading to pale green

towards the margin, flat and prominent on the upper surface, raised and prominent

on the lower surface. Inflorescences terminal, protogynous; female inflorescences

(l-)2 -flowered, positioned one node below the male inflorescences or solitary,

peduncles absent; male inflorescences composed up to 4 cymose partial inflorescences,

each monochasium with 3-6 flowers, peduncles 7-15 mm long. Bracts dark red, ovate,

keeled, c. 0.9-1. 1 cm long, c. 0.8 cm wide, margin entire, persistent. Male flowers:

pedicel white to pale pink, 7-9 mm long; tepals 2, pale pink with red flushes, ovate-

elliptic, 7-9 X 4-5 mm, margin entire, apex rounded; stamens c. 39, cluster conical,

stalked; filaments c. 0.8 nun long; anthers yellow, oblong, c. 1 mm long, apex slightly

emarginate. Female flowers: pedicel pale green, 4-6 mm long; ovary pale green, 1.3-2

X 0.3-0.4 cm, wings 3, equal, locules 3, placentae 2 per locule, placentation axile;

tepals 5, pink, outer 4 tepals ovate, c. 1 8-20 x 8-1 1 mm, innermost tepal oblanceolate,

c. 10 X 8 mm, margin entire, apex rounded; styles 3, deeply Y-shaped, c. 6 mm long,

divided c. 2 mm from the base; stigma yellow, papillose fonning a continuous twisted

band. Capsules (l-)2, pendent, 2.7-30 x 1,5-1 .7 cm, pale green when young, brown

when mature, locules 3, wings 3, equal, leathery when fresh, thinly fibrous when dry,

slightly expanded, tip rounded, dehiscing between locule and wing; pedicel c. 4-6 mm

long.

Additional specimens examined. BORNEO: Brunei: Tutong District, Lamunin, Ladan Hills

Forest Reserve, Ladan Hill, 20 Nov 2014, A.A. Joffre et al. BRUN 24029 (BRUN, SING).

Distribution and habitat. Begonia jamilahana is so far known only from the Ladan

Hills Forest Reserve, where it was documented on steep earth banks and adjacent

slopes in the mixed dipterocarp forest on yellow sandy clay soils.

Etymology. This species is named for Hajah Jamilah binti Haji Abdul Jalil, acting

Deputy Director of the Brunei Forestry Department, upon her retirement in 2015, as

a token of appreciation for her excellent service rendered to the Forestry Department,

as well as her unwavering support for the Brunei-Singapore botanical exploration

programme.

Provisional lUCN conservation assessment. Begonia jamilahana is so far known only

from a single population at Compartment 1, Ladan Hills Forest Reserve. The lUCN

status proposed here is Least Concern (LC) as the species is protected in a forest

reserve with no discernible threats. The Brunei Ministry of Industry and Primary

Resources officially announced in 2014 that logging will be phased out in all forest

reserves (Bmnei Times, 2014).

Notes. Begonia papyraptera Sands was described in Coode et al. (1996), based on a

single collection from the helipad Landing Point (LP) 286, Batu Apoi Forest Reserve,

Gard. Bull. Singapore 67(1) 2015

Table 1. Comparison of morphological characteristics, habitat and distribution among Begonia

conniegeriae S.Julia & Kiew, B. jamilahana Y.W.Low, Joffre & Ariffin and B. papyraptera

Sands.

Begonia conniegeriae

Begonia jamilahana

Begonia papyraptera

Stem (in life),

texture

Smooth and terete

Ribbed and angular

Stem (in life),

colour

Reddish or red brown

Pale green with red

nodes

Pale green

Stem nodes (in

life), swollen

Yes

Upper lamina

surface,

pubescence

Sparsely covered with

translucent erect hairs

Scantily covered with

translucent erect hairs

Glabrous

Upper lamina

surface (in life),

colour and

markings

Dark green

Dark green with red

markings on vein axil

nodes near tire leaf

base

Dark green with

white spots

Lower leaf

surface,

pubescence

Sparsely covered with

translucent erect hairs

Scantily covered with

translucent erect hairs

Glabrous

Lower leaf

surface (in life),

colour

Pale green

Maroon

Capsule wings,

shape proportion

Unequal

Equal

Capsule wing

base, shape

Cuneate

Rounded

Habitat, substrate

Limestone

Yellow sandy clay

soils

Unknown

Distribution

Lobang Cina, Gunung

Mulu National Park,

Marudi District,

Sarawak, Malaysia

Ladan Hills Forest

Reseiwe, Tutong

district, Brunei

Damssalam

Batu Apoi Forest

Reserve, Temburong

district, Brunei

Darussalam

A new Begonia from Brunei Darussalam

Fig. 1. Begonia jamilahana Y.W.Low, Joffre & Arififin. A. Habit. B. Close-up of upper leaf

surface. C. Detail of stem showing red nodes. D. Close-up of male portion of inflorescence.

E. Close-up of female flowers. F. Detail of styles showing twisted papillose stigmas. G. Cross

section of an immature fruit. H. Immature fruit (side view). I. Detail of a stipule. A & B from

P. Azlan et al. BRUN 23373, C-J from ex-situ collection at BRUN. (Photos: A & B, A.K.

Muhammad Ariffin; C-J, Y. W. Low)

Gard. Bull. Singapore 67(1) 2015

Fig. 2. Distribution of Begonia conniegeriae S. Julia & Kiew, Begonia jamilahana Y.W.Low,

Joffre & Ariffin, and Begonia papyraptera Sands.

Temburong district {Johns et al. RJ7422), without a detailed description but with a

brief Latin diagnosis as follows:

“a B. congesta Ridley alls fructus latioribus, caulibus plusminusve glabris

(non ‘hirtis’), staminibus 15 vel plus (non 12), tepalis staminibus multo

longioribus (non brevioribus); a B. leucochlora (vide supra) fructus apice

rotundato (non alis acutis et fructus apice truncate), petiolis longioribus

differt.”

A total of three duplicates were gathered, with two sheets indicated as holotype in K

(Sheet 1 of 2: K000761105, and Sheet 2 of 2: K000761106); while the third sheet,

which is an isotype, is lodged in BRUN (herbaria acronyms follow Thiers, continuously

updated). For this study, the two holotype sheets were examined through the Southeast

A new Begonia from Brunei Darussalam

Asian Begonia database (Hughes & Pullan, 2007), as well as the JSTOR® Global Plants

website (Global Plants, continuously updated). Detailed examination was carried out

on the isotype preserved in BRUN. In addition, a living plant of Begonia papyraptera

was discovered recently by the second author in cultivation in the ex-situ collection of

begonias at the Kuala Belalong Field Study Centre (KBFSC). This cultivated specimen

in KBFSC matches the type materials represented in K and BRUN and, therefore,

provided us with further insights into the species for this study.

Begonia conniegeriae S. Julia & Kiew was described in Sang et al. (2013). We

have been unable to examine the materials as the isotype indicated in the article as having

been distributed to SING had not arrived by 13 March 201 5 (Serena Lee (SING), pers.

comm.). Images of the type materials were also not available for examination either in

the Southeast Asian Begonia database (Hughes & Pullan, 2007) or the JSTOR® Global

Plants website (Global Plants, continuously updated). The morphological characters of

Begonia conniegeriae used in this study are extracted from Sang et al. (2013).

ACKNOWLEDGEMENTS. We are grateful to Haji Saidin bin Salleh, the Director of Forestry

Department, Brunei Darussalam for providing us with field support and assistance for this study.

Special thanks to Hajah Jamilah binti Haji Abdul Jalil and Noralinda binti Haji Ibrahim for

their encouragement and continuous support and assistance. Dr Ruth Kiew (KEP) kindly made

available translated Latin diagnoses by Dr Mark Coode (K) of all the Begonia species described

by Martin Sands (K) published in Coode et al. (1996) for this study. Various staff members of

the Brunei National Herbarium (BRUN) have kindly helped: Mediana Kampong and Muhd.

Wafiuddin Zainol Arifhn assisted with herbarium materials and database; Watu Awok, Azlan

Pandai, Jangarun Eri and Rauzaidi Mureh provided excellent assistance in the field, as well

as maintaining the ex-situ reseai'ch collection. Research funding for YWL is provided by the

National Parks Board, Singapore and the Singapore Botanic Gardens. Christina Soh (Library of

Botany and Horticulture, Singapore Botanic Gardens) kindly helped with missing references.

We are grateful to the two reviewers, Dr Mark Hughes (E) and Dr Daniel C. Thomas (SING),

for their constructive comments in improving this manuscript. This research collaboration is

conducted under the MoU programme "'’The Botanical Sw’vey of Brunei Darussalam'’'' between

the Brunei Forestry Department, Ministry of Industry and Primary Resources and the National

Parks Board, Singapore.

References

Brunei Times (2014). Logging no longer allowed in forest reserves: Ministry of Industry

and Primary Resources. 23 March 2014. http://bt.com.bn/news-national/2014/03/23/

logging-no-longer-allowed-forest-reserves-mipr (accessed on 13 Feb. 2015).

Coode, M.J.E., Dransfield, J., Fonuan, L.L., Kirkup, D.W. & Said, l.M. (1996). A Checklist of

the Flowering Plants and Gymnosperms of Brunei Darussalam, xvii + All p. Brunei

Darussalam: Ministry of Industry and Primary Resources. (Begoniaceae, p. 38-41)

Doorenbos, J., Sosef, M.S.M. & de Wilde, J.J.F.E. (1998). The Sections of Begonia: including

descriptions, keys and species lists. 266 p. The Netherlands: Wageningen Agricultural

University.

Gard. Bull. Singapore 67(1) 2015

Global Plants (continuously updated). JSTOR® Global Plants. ITHAKA. http://plants.jstor.

org/ (accessed on 13 Mar, 2015).

Heywood, V.H., Brummitt, R.K., Culhani, A. & Seberg, O. (2007). Flowering Plant Families

of the World. 424 p. UK: Royal Botanic Gardens, Kew. {Begoniaceae, p. 58-59)

Holttum, R.E. & Enoch, I. (1995). Gardening in the Tropics, viii + 384 p. Singapore: Times

Edition.

Hughes, M. (2008). An Annotated Checklist of Southeast Asian Begonia, xi + 164 p. UK: Royal

Botanic Garden Edinburgh.

Hughes, M. & Pullan, M. (2007). Southeast Asian Begonia Database. Royal Botanic Garden

Edinburgh. http://elnier.rbge.org.uk/Begonia/ (accessed on 10 Dec. 2014).

Merrill, E.D. (1921). A bibliographic enumeration of Bornean plants. J. Straits Branch Roy.

Asiat. Soc. Special No. Singapore: Fraser & Neave Ltd. {Begoniaceae, p. 414-416)

Sands, M.J.S. (1996). Begoniaceae: Begonia. In: Coode, M.J.E., Dransfield, J., Fonnan, L.L.,

Kirkup, D. W. & Said, I.M. (eds) A Checklist of the Flowering Plants and Gymnosperms

of Brunei Damssalam. Brunei Darussalam: Ministry of Industry and Primary Resources.

{Begoniaceae, p. 38-41)

Sang, J., Kiew, R. & Geri, C. (2013). Revision of Begonia (Begoniaceae) from the Melinau

limestone in Gunung Mulu National Park and Gunung Buda National Park, Sarawak,

Borneo, including thirteen new species. Phytotaxa 99(1): 1-34.

Tebbit, M.C. (2005). Begonias: Cultivation, Identification and Natural History. Ill p. Portland:

Timber Press.

Thiers, B. (continuously updated). Index Herbariorum: A global directory of public herbaria

and associated staff. New York Botanical Garden’s Virtual Herbarium. http://sweetgum.

nybg.org/ih/ (accessed on 6 Mar. 201 5).

Gardens’ Bulletin Singapore 67(1): 69-84. 2015

doi: 10.3850/S2382581215000083

Novitates Bruneienses, 3. Eight new woody plants in the

Brunei flora, including flve new species

K.M. Wong & Y.W. Low

Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, 259569 Singapore

wkin2000@gmai 1 .com

ABSTRACT. Five new species of seed plants are described: Alangium kayimiga K.M. Wong

(Cornaceae), Melastoma ariffinii K.M.Wong, Melastoma ashtonii K.M. Wong, Melastoma

jojfrei K.M.Wong & Y.W.Low (Melastomataceae), and Lasianthus jangarimii Y.W.Low

(Rubiaceae). New plant records for the ongoing Brunei floristic inventory include Planchonea

valida (Blunie) Blume (Lecythidaceae), Melastoma velutinosum Ridl. (Melastomataceae), and

Gardenia costiilata Ridl. (Rubiaceae).

Keywords. Borneo, Cornaceae, Lecythidaceae, Malesia, Melastomataceae, new records, new

species, Rubiaceae, Southeast Asia

Introduction

In this third installment of the series Novitates Bmneienses, we report our findings

of new species in four plant families. These include a new Alangium species

(Cornaceae), tliree new species and a new record in Melastoma (Melastomataceae), a

new Lasianthus species and a new Gardenia record (Rubiaceae), and a new record of

Planchonea (Lecythidaceae). Herbarium acronyms used follow Thiers (continuously

updated). All conservation assessments follow the methodology of lUCN (2012). For

the new species the assessments are global, for the new records we provide national

assessments for Brunei. All specimens cited have been seen by the authors.

Five new species

CORNACEAE

Alangium kayuniga K.M.Wong, sp. nov.

This new species is distinctive among the tree species of Alangium with a pinnately

veined leaf base, differing from A. havilandii Bloemb. in its subtruncate calyx tube

(the calyx tube in A. havilandii has distinct triangular teeth) and from A. javanicum

(Blume) Wangerin in its thicker leaves that dry medium brown, sparse and reticulate

tertiary venation and conspicuously asymmetric leaf base {A. javanicum has thinner

leaves typically di 7 ing olive brown to purple brown, regularly conspicuous scalarifonn

tertiary venation and typically symmetric leaf base). The new species also differs by its

Card. Bull. Singapore 67(1) 2015

glabrous calyx and fruit from both Alangium havilandii and A. javanicum, which have

a velvety hairy calyx and fruit. - TYPE: Brunei, Belait, Sungai Liang, Andulau Forest

Reserve, trail entrance to Compartment 7, mixed dipterocarp forest, 21 March 2007,

YusopBRUN 22161 (holotype BRUN; isotype SING). (Fig. 1)

Small tree 10-12 m tall, trunk 8 cm diameter. Leaves elliptic, 6.5-17 cm long, 2. 7-6. 6

cm wide, base cuneate, asymmetric, apex cuspidate (cusp 5-1 2 mm long), coriaceous,

glabrous, drying medium brown; midrib sunken on upper surface, prominent on lower

surface, secondary veins 7-9 pairs, tertiary venation sparse and reticulate; petioles 4-7

mm long, 1-1 .5 mm diameter, slightly channelled on upper side. Flowers not known.

Infructescence a short cyme, peduncle 3-6 mm long, with l-3(^) faiits. Fruits on

short pedicels 4-5 mm long, 1-1.5 mm diameter; ovoid-ellipsoid, slightly compressed

with 10 longitudinal ridges and grooves, 1.8-2. 2 cm long, 1.2-1. 4 cm wide; glabrous,

crowned by the persistent calyx limb and disc; calyx tube subtruncate, 1 .5-2 mm high,

glabrous. Seeds one per fruit, ellipsoid-compressed, 1.2-1 .3 cm long, 0. 8-0.9 cm

wide, 2-3 mm thick, testa smooth.

Etymology. ‘Kayu’ refers to tree or wood in Brunei Malay; this species is named for

Mohd. Niga bin Abdullah Nangkat, formerly Senior Forestry Assistant with the Brunei

National Herbarium and now retired, who led many collecting trips in Brunei forests.

Distribution. This would seem to be a very rare and narrow endemic in Brunei as it is

only known from the type collection.

Provisional lUCN conseivation assessment. Data Deficient (DD) as it has only been

collected once and no information is available on its distribution or population size.

Further field observations would enable a better understanding of its conservation

status in Bmnei.

Additional specimens examined. The species is only known from the type.

Notes. The slightly compressed ovoid-ellipsoid fruit, with longitudinal ridges and

grooves and crowned by the persistent calyx limb and disc, is characteristic oi Alangium

(Bloembergen, 1935, 1939). The most recent revision of Bornean Alangium was by

Berhaman (1995), in which ten species were recognised for the island. Of these, two

are climbers and another six have 3-5-veined leaf bases, characters not found in our

new species. Instead, it more closely resembles Alangium havilandii (with asymmetric

leaf bases and a swamp habitat) and A. javanicum (with symmetric leaf bases and a

mixed dipterocarp forest provenance). However, the glabrous calyx and fruit surface

oi Alangium kayuniga is consistent for both younger and more mature fruit, whereas in

A. havilandii and A. javanica the velvety nature of the calyx and fi'uit surface is evident

from young stages and persistent through to mature fruit.

Woody novelties in Brunei flora

Fig. 1. Alangium kayuniga K.M.Wong, fruiting branch. (Photo: Muhammad Ariffin)

MELASTOMATACEAE

Melastoma arijjinii K.M.Wong, sp.nov.

This new species resembles Melastoma stenophyllum Merr. in having linear leaves and

broad-triangular hypanthium scales that are less than three times as long as their width,

but differs in its hypanthium scales that are subulate and basally inflated with subentire

to sparsely denticulate margin, and which are more sparsely set and intermixed with

minute similar scales (in contrast to the irregularly short-fringed flat hypanthium scales

that are of one general size and closely overlapping in M. stenophyllum). - TYPE:

Tutong, Ramba, Ulu Tutong, down valley to SW of helicopter pad LP 239, 4°25'N

114°50'E, 150-200 m asl, 8 May 1992, Johns, Niga, Shanang & Han 7547 (holotype

BRUN; isotype K). (Fig. 2A-B)

Melastoma polyanthum var. linearifolium Bakh.f., Meded. Bot. Mus. Herb. Rijks

Univ. Utrecht 91: 69 (1943). - TYPE: Borneo, Gunong Narik, Kelam, May 1894,

Molengraaff B3460 (lectotype L, designated here).

Gard. Bull. Singapore 67(1) 2015

Fig. 2. Hypanthium scale types and detailed view of upper leaf surfaces in Melastoma ariffinii

(A, B), M ashtonii (C, D) and M joffrei (E, F). A & B from Othman et al. S 56078; C & D from

Ashton BRUN 5629; E & F from Wong & Joffre WKM 3207 (all SING). (Photomicrographs:

Y.W. Low)

Rheophytic bush 1-1.5 m high or more. Branch internodes covered with dense,

appressed, ovate-lanceolate, subentire to slightly short-serrate scales. Leaves with

petioles 0.4-2 cm long, c. 0.1 cm diameter; blades linear, 3.2-10 cm long, 0.4-1. 5 cm

wide, the 3 longitudinal veins sunken on the upper surface and prominent on the lower

surface, covered by a mixture of larger lanceolate appressed scales (c. 1 mm long)

and smaller appressed lanceolate scales (<0.3 mm long), lamina on the upper surface

Woody novelties in Brunei flora

in dried material with abundant conspicuous elongate spicule-like patterns fomied by

rows of pale crystalliferous cells, glabrous or occasionally with micro-hairs less than

0.1 mm long (Fig. 2B), on the lower surface with scabrid scales with emergent scabrid

tips to 0.1 mm long. Flowers 1-3 in short cymes, peduncle 3-5 nun long, branches 1-2

mm long only; pedicels 4-6 mm long; hypanthium (Fig. 2 A) 5-7 imn long, 4-6 mm

diameter, green then magenta, with sparsely airanged scales, the scales subulate and

basally inflated, to 2 mm long, with entire to sparsely denticulate margins, intermixed

with minute similar scales mostly 0.5-1 nun long or shorter, lobes 5, broad-triangular,

5-6 mm long, alternating with inter-sepalar protuberances 1-2 nuu long; petals

obovate, c. 2.5 cm long, 1.8 cm broad, mauve; stamens not seen. Fruits c. 9 mm long,

8 mm diameter, green to reddish brown.

Etymology. The species is named after Muhammad Ariffin A. Kalat, experienced forest

botanist at the Brunei National Herbarium.

Distribution. This is apparently the common rheophytic Mela stoma species in

Clearwater streams of the Northwest Borneo region, including Brunei, Sarawak and

W Kalimantan.

Provisional lUCN conservation assessment. Least Concern (LC) as the species is

common and widespread. Besides that, the habitat in Brunei is also protected in Forest

Reserves and a National Park.

Additional specimens examined. BRUNEI: Temburong: First big waterfall on R. Temburong,

c. 500 ft, 6 Nov 1959, Ashton BRUN 759, (BRUN, K, SfNG). MALAYSIA: Sarawak: 5th

Division, Ulu Lawas, Kota Forest Reserve, on bank of Sg. Kota, 21 Oct 1971, Chai & Ilias S

31110 (A, K, L, SAN, SAR, SING); Kapit Division, Balleh, Ulu Mengiong, Sg. Entejum, 27

Oct 1988, Othman, Rantai & Jugah S 56078, (K, KEP, L, SAR, SING); Kapit Division, Sg.

Belaga at upper rapids, 12 Apr 1963, Ashton S 18242 (K, L, SAR, SING).

Notes. Meyer (200 1 ) had apparently not studied specimens of this rheophytic Melastoma

taxon in Borneo; none bears his annotation or are mentioned or indexed in his revision.

He placed Melastoma polyanthum var. linearifolium Bakh.f. in the synonymy of M.

malabathricum, but the variety has not hitherto been lectotypified. The syntypes are

MolengraaffB3460 from Mount Kelam in W Kalimantan (L) (which is our M. ariffinii

here) and Seimund s.n. from Kwala Teku in the Malay Peninsula (K: K000867833)

(which is a nan‘ow-leafed rheophytic foiua of M. malabathricum). Ridley’s Melastoma

polyanthum var. angustifolium (Ridley 1922: 765, as ‘angustifolia’) was applied

to this Malay Peninsula taxon and requires lectotypification (to be addressed in a

separate review of the Malayan species), but in any case is different from our new

species here. Here we lectotypify Melastoma polyanthum var. linearifolium with the

Bornean specimen, as Bakhuizen’s work addressed mainly the Netherlands East Indies

(Bakhuizen van den Brink Jr., 1943).

The Sumatran rheophyte Melastoma stenophyllum was also reduced by Meyer

(2001) to synonymy under M. malabathricum L. although it is, in our view, quite

Gard. Bull. Singapore 67(1) 2015

distinct: not only because the leaves are stenophyllous in an extreme manner, with

blade lengths 9-11 times the width (much more linear than narrow variations of the

typically elliptic leaves in M. malabathricum which has blade lengths 5-7(-8) times

the width), but also the flowers are typically solitary (compared to cymes of several to

7-9 flowers iiiM malabathricum).

Steenis (1981) confused ariffinii withM borneense (Cogn.) Bakh.f.,

which is typified by a collection from West Kalimantan: De Vriese 168 (L). The De

Vriese collection has leaves just L3-2.2 cm wide, but still clearly elliptic in shape,

and not sufficiently linear as in the truly rheophytic taxa; Melastoma ariffinii here

has leaves that are linear and only 0.4-1. 5 cm wide (length 8-10 times the width).

Also, the De Vriese specimen has 2-7 flowers in a cyme, and a hypanthium with

short-fringed, flat ovate-triangular scales, whereas Melastoma ariffinii has only 1-3

flowers and hypanthium scales that are subentire to remotely denticulate, and subulate

inflated structures. Lastly, the upper leaf surface has scabrid hairs 0.3-0. 5 mm long in

Melastoma borneense but subglabrous with only occasionally scabrous micro-hairs

less than 0. 1 mm long in M. ariffinii, so that the latter’s upper leaf surface is not scabrid

to the touch. We agree with Meyer (2001) that Melastoma borneense is synonymous

withM malabathricum', the two taxa have the same hypanthium scales and leaf upper

surface characters, among others.

Melastoma ashtonii K.M.Wong, sp.nov.

This new species resembles Melastoma stenophyllum Men*, in having linear leaves,

but differs in its hypanthium scales that are basally thickened and deeply incised into

linear segments (Fig. 2C) (in contrast to the fringed flat ovate-triangular scales on the

hypanthium in M. stenophyllum). - TYPE: Brunei, Belait, Sg. Ingei, rocky river bank,

21 January 1959, Ashton BRUN 5629 (holotype BRUN; isotypes K, L, SAR, SING).

(Fig. 2C-D)

Rlieophytic bush 1-2 m high, stems sometimes gnarled. Branch internodes covered

with dense, appressed, ovate-lanceolate, subentire to slightly short-serrate scales.

Leaves with petioles 0.5-1 .4 cm long, c. 0. 1 cm diameter; blades linear, 3.5-9 cm long,

0.4-1 cm wide, the 3 longitudinal veins sunken on the upper surface and prominent

on the lower surface, covered by a mixture of larger lanceolate scales 0.5- 1.2 mm

long and smaller lanceolate scales 0.2-0. 3 mm long, lamina on the upper surface in

dried material with abundant conspicuous pale elongate rows of crystalliferous cells

fomiing spicule-like patterns and quite glabrous (Fig. 2D), on the lower surface with

scabrid scales with 0.2-0. 3 mm emergent scabrid tips. Flowers 1-3 in short sessile

cymes, branches 1-2 mm long only; pedicels \-A mm long; hypanthium 10-11 mm

long, 7-8 mm diameter, green then crimson, moderately to densely scaly, the scales

with an entire thickened base 0. 1-0.2 mm high and linear segments 0.6-1.4(-2) mm

long, lobes 5, narrowly triangular, 8-10 mm long, alternating with inter-sepalar lobes

4-5 mm long; petals obovate, c. 3.3 cm long, c. 2 cm wide, mauve; short stamens

with a filament c. 7-8 mm long and anthers c. 5 mm long and 2.5-3 mm wide, with

Woody novelties in Brunei flora

apiculate tips, long anthers only one seen, filament unknown, anther c. 12 mm long.

Fruits (young) c. 8 mm long, 4 mm diameter, green to reddish brown.

Etymology. The species is named after Professor Peter Shaw Ashton, Brunei’s first

Forest Botanist who collected the type on his first assignment to the tropics, and who

later taught at Aberdeen and Haiward Universities.

Distribution. Known only from the Sungai Ingei area in Brunei.

Provisional lUCN conset'vation assessment. Least Concern (LC) as the habitat of the

species is well protected in the Sungai Ingei Conservation Area, which is also now

included within the designated Heart of Borneo area within Brunei.

Additional specimens examined. BRUNEI: Belait: Falls just upstream from Batu Melintang, 4

Jan 1989, Wong WKM 680 (BRUN, K, SING).

Notes. See comments under Melastoma ariffinii above. The new rheophytic species

here, Melastoma ashtonii, is also clearly different from M. malahathricum by its

hypanthium scale type: basally thickened scales that are deeply incised into linear

segments, often resembling miniature rakes, instead of the iiregularly short-serrate to

shallowly laciniate flat narrowly triangular-lanceolate scales as m M. malabathricum

(Fig. 3). Also, in both Melastoma malabathricum and M. stenophyllum, the pale

spicule-like patterns formed by rows of crystalliferous cells embedded in the upper leaf

surface are distally continuous with short scabrid hairs, but the upper laminar surface in

M. ashtonii is quite glabrous. In addition, Melastoma ashtonii has rather conspicuous

inter-sepalar lobes 4-5 mm long, compared to the much less conspicuous ones that

occasionally occur in M. malabathricum, absent altogether inM. stenophyllum.

Steenis (1981: 283) had already noticed the potential novelty of this taxon,

which he enumerated as “Melastoma sp. (nov. ?)” in his Rheophytes of the World.

Melastoma joffrei K.M.Wong & Y.W.Low, sp.nov.

This species is unique among Melastoma species with isomorphic stamens by

its hypanthium covered by small triangular scales of different sizes (Fig. 2E) (not

hairlike bristles as in M. cynoides Sm. and M. moluccanum Blume, nor penicillate

emergences as in M. montanum (Lauterb.) K.Meyer), very small bracts that do not

enclose the hypanthium (unlike in M. montanum), and scabrid upper leaf surfaces with

a mixture of longer and tiny coarse suberect to upcurved hairs (Fig. 2F) (not stidgose to

appressed pilose upper leaf surfaces as in the other species). - TYPE: Brunei, Tutong

district, Rambai, Ladan Hills Forest Reserve, on ridge NE of campsite beside Sungai

Buing, along old logging track, 13 October 2012, Wong & Joffre WKM 3207 (holotype

BRUN; isotype SING). (Fig. 2E-F, 4)

Treelet (single-stemmed) or shrub (with several stems from the base) to 1.5 m high.

Branch internodes eoarse with dense, spreading to erect small triangular scales.

Gard. Bull. Singapore 67(1) 2015

Fig. 3. Consistently narrowly lanceolate, serrate-laciniate and densely overlapping hypanthium

scales, and scabrid-hairy upper leaf surfaces in Melastoma malabathricum from the Malay

Peninsula (A, B), Sabah (C, D) and Sarawak (E, F). A & B from Jagoe s.n.,C&.T) from Kadir

A 2855; E & F from Moulton 6723 (all SING). (Photomicrographs: Y.W. Eow)

Leaves with petioles 0.6-2 cm long, with a mixture of scattered to dense suberect short

scales and scabrid hairs; blades ovate-elliptic, 5.5-12.5 cm long, 1.5-4. 5 cm wide,

longitudinal nerves 3-5 (-7), sunken on upper surface, raised on lower surface and

bearing a mixture of suberect short scales and scabrid hairs, transverse veins between

the nerves scalariform, slightly sunken on upper surface, raised on lower surface;

Woody novelties in Brunei flora

Fig. 4. Melastoma joffrei K.M.Wong & Y.W.Low. Open flowers showing isomorphic stamens.

Inset: Detail of upper leaf surface showing different hair types contributing to a sandpapery

roughness. All from Wong & Joffre WKM 3207. (Photos: K.M. Wong)

upper laminar surface with a mixture of longer and tiny coarse suberect to upcurved

hairs, scabrid-sandpapery to the touch; lower laminar surface similarly scabrid.

Inflorescence a compact terminal cyme of (7-)9-13 flowers, main cyme branches

3-11 mm long, bracts tiny, broad triangular to semicircular, c. 1 mm long and wide.

Flowers small, only c. 2 cm across; hypanthium urceolate, 4.5-5 mm long, 4.5-5 mm

wide, surface scales short triangular, appressed, margins minutely denticulate, not or

only slightly imbricate and not entirely obscuring the hypanthium surface, green; calyx

lobes 5, broadly triangular, 1.5-2. 5 mm long, 1-2 mm wide; petals 5, obovate, 6-7.5

mm long (8-9 mm in fresh material), 4-5.5 mm wide (5-6 mm in fresh material),

the apex subtruncate to rounded, margins short-ciliate, pale purple-pink; stamens

10, isomorphic, filaments 3-4 mm long (3 .4-4.5 mm in fresh material), white, each

apically bearing two knoblike pale yellow appendages at the insertion of the anther,

anthers erect, 2-2.5 mm long (2.5-3 mm in fresh material), pale creamy yellow, apex

rounded, opening by apical pores; style slender-cylindric, 4. 5-5. 5 mm long (5.5-7

mm in fresh material), with a pale green base, then pink for the most part, apically

pale yellow; stigma rounded to subtruncate, 4-5-lobed, pale green. Fruit urceolate to

subglobose, 5.5-6 mm long, 5.5-6 mm wide, dehiscing irregularly transversely.

Gard. Bull. Singapore 67(1) 2015

Etymology. The species is named for our colleague Joffre bin Haji Ali Ahmad, Forest

Botanist at the Brunei National Herbarium and experienced and long-time collector of

Brunei plants.

Additional specimens examined. BRUNEI: Tutong; Rambai, Ladan Hills Forest Reserve, Sg.

Buing, disturbed mixed dipterocarp forest, on ridge, 29 Oct 2013, Ariffin, Low, Y.W. & Azlan

BRUN 23229 (BRUN, SING). MALAYSIA. Sarawak: Tatau district, Anap, Muput Kanan,

Ulu Naoung, 6 m asl, 14 Oct 1963, Ashton S 19569 (A, BO, K, KEP, L, SAN, SAR, SING).

Distribution. Known only from the type locality in Brunei and another collection in

Sarawak’s Tatau district.

Provisional lUCN conservation assessment. Data Deficient (DD) as it has been only

twice collected with one of the collections made over 50 years ago. Field observations

are badly needed for a better understanding of its conservation status.

Notes. In Meyer’s revision of Melastoma (Meyer, 2001), there is a small group of

three Melastoma species with isomorphic stamens, M. cynoides, M. moluccanum and

M. montanum, which he recognises for the more seasonal parts of insular SE Asia and

New Guinea; the rest of the species in the genus have fiowers with dimorphic stamens.

Melastoma joffrei is Borneo’s first known tom a species with isomorphic stamens.

RUBIACEAE

Lasianthus jangamnii Y.W.Low, sp. nov.

This new species resembles Lasianthus linearifolius H.Zhu in having linear leaves but

differs by having triangular stipules, bullate leaves with a thin-papery and crispy texture,

a revolute margin and brochidodromous venation (with 17-40 pairs of secondary

veins fonning a distinct marginal vein), and a calyx with shorter tube (c, 0.4 mm long)

and shorter lobes (c. 0.4-0.9 mm long). In contrast, Lasianthus linearifolius has awl-

shaped (subulate) stipules, leaves with a plane surface and leaf margin, subcoriaceous

texture, craspedodromous venation (with 10-16 pairs of secondary veins), and a

calyx with longer tube (c. 1 mm long) and longer lobes (c. 1 mm long). - TYPE:

Brunei, Temburong, Amo, Sungai Temburong and Sungai Machang junction, ridge,

120-250 m alt., 18 September 1990, Puff et al. 900818-1/1 f mature fruits (holotype

BRUN; isotypes K, SAR, SING 2 sheets). (Fig. 5)

Understorey treelet c. 2 m high, stem c. 3^ mm wide. Branches solitary, slender,

terete, c. 1-2 mm wide, sparsely puberulent to subglabrous. Stipules narrowly

triangular, c. 1-3 mm long, sparsely puberulent. Petiole 4— 6(-7) mm long, 0.8-1. 2

mm wide, sparsely puberulent. lamina linear, (8.5-)l 1-13.7(-18.4) cm long,

(0.8-)l-1.5(-2.2) cm wide, bullate, thin-papery and crispy; leaf base cuneate; leaf

apex long-caudate; leaf margin entire to slightly repand, revolute; midrib prominent

Woody novelties in Brunei flora

Fig. 5. Lasianthus jangarunii Y.W.Low. A. Branches with conspicuously corrugated leaves.

B. Close-up of a flower bud and an immature fiaiit with persistent calyx lobes visible. C. Habit (Ulu

Temburong National Park). All from Low et al. LYW 629. (Photos: A-B: K.M. Wong, C: Y.W. Low)

Gard. Bull. Singapore 67(1) 2015

and glabrous on both sides; secondary veins 17^0 pairs, making an angle of 85-90°

with the midrib and joining to fonn a distinct marginal vein, inconspicuous to slightly

raised and glabrous on upper side, prominent and glabrous on lower side; tertiary

venation inconspicuous to slightly raised on upper side, inconspicuous on lower side.

Inflorescences sessile; bracts absent. Flowers fascicular, sessile; calyx campanulate,

tube c. 0.4 mm long, outside surface puberulent; marginal lobes 4(-5), triangular,

c. 0.4-0.9 mm high; corolla hypocrateriform, white; tube c. 1 mm long, outside

puberulent; lobes 4(-5), triangular, c. 1.2 mm long, c. 1 mm wide, valvate, outside

puberulent. Drupes subglobose, c. 2-3 mm long, 2.5-3 mm wide, 4(-5)-ridged when

dry, puberulous, maturing black {Kirkup et al. 898); calyx persistent at fruit apex, with

4(-5) small triangular calyx lobes. Pyrenes 4(-5), thick-walled, each with a single

seed.

Etymology. This species is named for Jangarun anak Eri, knowledgeable field assistant

and tree climber attached to the Brunei National Herbarium (BRUN), who has assisted

numerous botanists with fieldwork.

Distribution and habitat. Lasianthus jangarunii is endemic to northwest Borneo

(Brunei: Temburong district, and Sarawak: Marudi district), where it grows on clay-

rich sedimentary soils as an understorey treelet on steep slopes of mixed dipterocarp

forest.

Provisional lUCN conservation assessment. Least Concern (LC) as the species is

protected in the Ulu Temburong National Park in Brunei and, although only collected

twice, in Sarawak it is recorded from Pulong Tau National Park where the habitat is

also protected.

Additional specimens examined. BRUNEI: Temburong: Amo, Kuala Belalong (‘Kuala

Temburong’ on label), hill, 21 Jul 1988, Wong WKM 253, leafy branch (BRUN, K, SING),

Batu Apoi Forest Reserve, on slope along Sungai Engkiang, 20 Nov 1991, Hansen 1592, young

fruits (BRUN); Amo, Ulu Belalong, LP 382, ridge top, 22 Jan 1994, Kirkup et al. 898, young

fruits (BRUN); Amo, Ulu Temburong National Park, hill behind Lubok Umar, ridge, 18 May

2014, Low et al. LYW 629, flower buds (BRUN, K, L, SAN, SING). MALAYSIA. Sarawak:

Marudi, Pulong Tau National Park (western part), Ulu Sungai Baong, on steep slope, 93 1 m

asl, 10 May 2007, Sang et al S 98058, young ifuits (K, KEP, L, SAN, SAR, SING); ibidem,

trail along the ridges to Bukit Tenidan, 1 1 May 2007, Muliati et al. S 97911, fruits (KEP, SAR).

Notes. Davis in Coode et al. (1996) identified this species as a taxonomically distinct

entity of Lasianthus from Temburong, which was listed only as Lasianthus “sp. 7”.

Zhu et al. (2012) evidently overlooked Coode et al. (1996) and this species while

preparing for the revision of Malesian Lasianthus; he stated that only “a selection

of the Malesian collections in the herbaria BKF, KEP, MO, and SING” had been

consulted. A recent collection from Ulu Temburong National Park {Low et al. LYW

629) matched Lasianthus “sp. 7” of Davis (Coode et al., 1 996), and is distinct from all

other Lasianthus taxa recognised by Zhu et al. (2012). This is so far the only known

Lasianthus species in Malesia with strongly bullate and linear leaves.

Woody novelties in Brunei flora

Table 1. Comparison of morphological characteristics and distribution between Lasianthus

jangarimii Y.W.Low and L. linearifolius H.Zhu.

Lasianthus jangamnii Lasianthus Uneanfolius

Stipules triangular

Leaves bullate

Leaf texture thin-papery and erispy

Leaf margin revolute

Secondary veins 17-30 pairs

Loop-veined secondary venation

(broehidodromous)

Calyx tube c. 0.4 mm long

Calyx lobes c. 0.4-0. 9 mm long

Restricted to northwest Borneo (Brunei:

Temburong district, and Sarawak: Marudi

district) on clay-rich sedimentary soils

Stipules awl-shaped (subulate)

Leaves smooth

Leaf texture subcoriaceous

Leaf margin plane

Seeondary veins 10-16 pairs

Secondary venation emerging

from the midrib running towards

and terminating at the leaf margin

(eraspedodromou s)

Calyx tube c. 1 nun long

Calyx lobes e. 1 mm long

Restrieted to Mount Kinabalu (Malaysia:

Sabah) on ultramafic soil

Lasianthus jangarunii differs fromZ. linearifolius, which also has linear leaves,

by its stipules, leaf texture and venation, and calyx dimensions (see diagnosis above).

The character-states are compared in Table 1 .

Apart from that, Lasianthus jangarunii is restricted to northwest Borneo

(Brunei: Temburong district, and Sarawak: Marudi district) on clay-rich sedimentary

soils, whereas L. linearifolius is endemic to Mount Kinabalu (Sabah) growing on

ultramafic soil.

Lasianthus has some consistent characters that are easily recognised, when

adequate material permits, such as paired axillary inflorescences and pyreniferous

blue fruits (as in the type L. cyanocarpus Jack, but there are some species with white,

red or black fruits). In addition, Lasianthus spp. are more typically treelets in the

forest understorey with only solitary (not paired) primary branches developing along

the vertical stem. This branching feature was not noted in Zhu et al. (2012) but is

nevertheless important and represented in more recent collections of the genus; a good

example was illusfrated for Lasianthus pedicellatus H.Zhu in Zhu et al. (2012: 72, Fig.

30).

Gard. Bull. Singapore 67(1) 2015

New plant records for Brunei

LECYTHIDACEAE

Planchonea valida (Blume) Blume

BRUNEI. Temburong: Amo, Kuala Belalong, Field Study Centre, 24 Apr 1998, Joffre BRUN

19016 {mum).

This is a new tree genus and species record for Brunei, collected after the publication

of Coode et al. (1996).

Provisional lUCN Conservation Assessment. Data Deficient (DD) for Brunei as the

species is only known from a single collection from Kuala Belalong. Further field

observations are needed for a better understanding of its conservation status in Brunei.

MELASTOMATACEAE

Melastoma velutinosum Ridl.

BRUNEI: Belait: Labi, Labi Forest Reserve, Compt 49, Ulu Sungai Rampayoh, 19 May 2009,

Yusop BRUN 22636 (BREfN); Melilas, Ulu Ingei hotsprings, 4°08’N, 114‘’43’E, 20 m asl, 7

Mar 1 996, Joffre BRUN 1 7295 (BRUN, SING). Tutong: Rambai, Ladan Hills Forest Reserve,

Bukit Bedawan, northwest of LP 263, 4‘’29'33"N 1 14''48'52"E, 250 m asl, 28 Mar 1997, Joffre

BRUN 18147 (BRUN, SING).

This species is known for the Malay Peninsula and Borneo (Sabah and Sarawak)

(Meyer, 2001). The several collections here record it for Brunei for the first time.

It is a treelet (single-stemmed) or slmib (with several stems from the base) to 3

m high. Distinctive features include the branch interaodes that are hirsute with dense,

spreading to curved long bristles; ovate-elliptic leaves with 3-5(-7) longitudinal nerves

and an upper lamina surface that is appressed-pilose (hairs to 0.5 mm long), slightly

velvety to the touch; a compact terminal cyme of 3-7 small flowers (only 2 cm across

in dried material); and a hypanthium densely covered by slender bristles 3-5 mm long.

Provisional lUCN conservation assessment. Least Concern (LC) for Brunei as the

species is common. This species occurs throughout northwest Borneo.

RUBIACEAE

Gardenia costulata Ridl.

BRUNEI: Belait: Andulau, without date, Ashton A2865 (BRUN). Temburong: Bukit Patoi,

without date, Ashton 3970 (BRUN).

Woody novelties in Brunei flora

Gardenia costulata Ridl. was first described by Ridley (1934) based on a single

collection (Beccari 1986, K, BM) from Sarawak. While reviewing Sundaland

Gardenia, Low (2010) enumerated additional materials of G. costulata, including

a collection for Sabah (Madius SAN 50094) and Kalunantan {Hallier B1285), most

of which had been erroneously identified as G. pterocaJyx Valeton in many herbaria.

Gardenia costulata has truncate stipules with a revolute margin, and thin-coriaceous

leaves with an acuminate to long-cuspidate apex and a pubescent lower leaf surface.

These vegetative features adequately pemiit its distinction from G. pterocalyx, which

also has tmncate stipules but with a plane margin, and thicker leaves with a typically

rounded apex and a glabrous lower leaf surface. Although the two Brunei specimens at

the Brunei Herbarium were without flower or fruit, it was possible to match these two

specimens to G. costulata based on the distinctive stipule character, the shape of the

leaf apex, the leaf texture, and the presence of pubescence. Apart from moiphological

characters, the kerangas vegetation at Bukit Patoi in Brunei where Ashton 3970 was

collected also matches the habitat known for other specimens of G. costulata.

Gardenia costulata is restricted to Borneo, and so far has only been recorded

from kerangas forest. The habitat of Ashton A2865 from Andulau is not known,

although the main habitat type recorded for Andulau is Mixed Dipterocarp Forest

(Ashton, 1964).

Provisional lUCN conservation assessment. Data Deficient (DD) for Brunei as the

species is only known from two localities, naiuely Andulau (Belait District) and

Bukit Patoi (Temburong District). Field observations are badly needed for a better

understanding of its conservation status in Brunei.

ACKNOWLEDGEMENTS. The Ministry of Industry and Primary Resources and the Forestry

Department, Brunei Darussalam, and the Singapore Botanic Gardens, National Parks Board,

Singapore, provided the opportunity and fiuiding for this work. For special field coordination

and assistance, we thank Noralinda binti Haji Ibrahim, Roslinah bind Haji Mohsin (Forestiy

Department), Joffre Haji Ali Ahmad, Muhd. Ariffin A. Kalat, Watu bin Awok, Jangarun anak

Eri, Azlan bin Pandai, Muhammad Waffiuddin bin Zainol Ariffin and Rauzaidi bin Mureh

(BRUN); Serena Lee, Paul K.F. Leong and Paul Parusuraman Athen (SING). We are grateful

to the keepers and curators of the BO, BRUN, L, SAN, SAR and SfNG herbaria for facilitating

study of specimens in their care.

References

Ashton, P.S. (1964) Ecological Studies in the Mixed Dipterocarp Forests of Brunei State.

Oxford Forestry Memoirs 25. Oxford: Clarendon Press.

Bakhuizen van den Brink Jr., R.C. (1943). A contribution to the knowledge of the

Melastomataceae occurring in the Malay Archipelago especially in the Netherlands East

Indies. Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 91: 1-391.

Berhaman, A. (1995). Alangiaceae. In: Soepadmo, E. & Wong, K.M. (eds) Tree Flora of Sabah

and Sarawak, vol. 1. Pp. 5-14. Malaysia: Forest Research Institute Malaysia, Sabah

Forestry Dept. & Sarawak Forestry Dept.

Gard. Bull. Singapore 67(1) 2015

Bloembergen, S. (1935). The genus Alangium in the Netherlands Indies. Blumea 1: 241-294.

Bloembergen, S. (1939). A revision of the genus Alangium. Bull Jard. Bot. Buitenzorg 3, 16:

139-235.

Coode, M.J.E., Dransfield, J., Fonnan, L.L., Khkup, D.W. & Idris, M.S. (1996). A Checklist

of the Flowering Plants and Gymnospenns of Brunei Darussalam. All p. Brunei

Darussalam: Ministry of Industry and Primary Resources.

lUCN (2012). lUCN Red List Categories and Criteria: Version 3.1. 2nd ed. Switzerland, Gland

and UK, Cambridge: lUCN.

Low, Y.W. (2010). Taxonomy and Biogeography of Sundaland Gardenia (Rubiaceae). M. Sc.

thesis. Kuala Lumpur: University of Malaya.

Meyer, K. (2001). Revision of the Southeast Asian genus Melastoma (Melastomataceae).

Blumea 46: 351-398.

Ridley, H.N. ( 1 922). The Flora of the Malay Peninsula, Vol. 1. London: L. Reeve and Company.

Ridley, H.N. (1934). New Malayan Rubiaceae. J. Bot. 72: 274—275.

Steenis, C.G.G.J. van (1981). Rheophytes of the World. 407 p. The Netherlands: Sijthoff &

Noordhoff

Thiers, B. (continuously updated). Index Herbariorum: A global directory of public herbaria

and associated staff New York Botanical Garden’s Virtual Herbarium. http://sweetgum.

nybg.org/ih/ (accessed on 31 May 2014).

Zhu, H., Roos, M.C. & Ridsdale, C.E. (2012). A taxonomic revision of the Malesian species of

Lasianthus (Rubiaceae). Blumea 57: 1-102.

Gardens’ Bulletin Singapore 67(1): 85-94. 2015

doi: 10.3850/S2382581215000101

Hoya undulata (Apocynaceae, Asclepiadoideae),

a new myrmecophytic species from Borneo,

and typification of H. darwinii

S. Rahayu\ U. Meve^ & M. Rodda^

'Bogor Botanic Gardens, Indonesian Institute of Sciences,

Bogor, Indonesia

srirahayukrb@yahoo.com

^Department of Plant Systematics, University of Bayreuth,

95440 Bayreuth, Germany

^Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, 259569 Singapore

rodda.michele@gmai 1 .com

ABSTRACT. In this paper we document the discovery of a new Hoya species from Borneo,

H. undulata S.Rahayu & Rodda. The leaves of the new species form domatia, specialised

structures harbouring ants, previously only observed in Hoya mitrata Kerr and H. darwinii

Loher. The corona is unique among Bornean Hoya species as its lobes develop two lateral

hooked appendages, also present in H. griffithii Hook.f, a species from mainland Asia. A

lectotype for Hoya darwinii is also selected.

Keywords. Ant plant, domatia, heath forest, Hoya mitrata, Indonesia, myrmecophyte. West

Kalimantan

Introduction

Hoya R.Br. is a poorly known tropical genus for which a modern revision is lacking.

Extensive field investigations and accurate documentation of new species are essential

before a generic revision can be compiled (Rodda & Ercole, 2014). In Borneo,

Merrill (1921) listed only eight Hoya species and more recently Forster et al. (1998)

described the diversity of the genus in Borneo as ‘unknown’. Recently, new taxa have

been described based on collections from Sabah (e.g. Lamb et al., 2014; Rodda et

al., 2013) and to a lesser extent, Sarawak (Rodda & Simonsson, 2011a, b; Rodda &

Simonsson Juhonewe, 2013a). The higher number of novelties described from Sabah

is directly linked to efforts by the staff at Kipandi Park (Kampung Kipandi, Moyog,

Sabah) to cultivate wild-collected sterile specimens until they bloom so that they can

be identified (Lamb et al., 2014), an approach successfully adopted by Rintz (1978) in

Peninsular Malaysia. Among the species described in Lamb et al. (2014) only Hoya

sammannaniana A.Lamb et al. is also found in Kalimantan. Two further species, Hoya

rintzii Rodda & Simonsson and Hoya beccarii Rodda & Simonsson can be found

throughout Borneo and are widespread elsewhere in SE Asia (Rodda & Simonsson

Gard. Bull. Singapore 67(1) 2015

Jiionewe, 2013b; Rodda et al., 2014). The diversity of Hoya in Indonesian Borneo is

least known due to the scarcity of recent collections; the only information on Hoya

diversity in Central Kalimantan can be found in Rahayu (2006) who lists only nine

species. Lamb et al. (20 14) suggested that 60 to 70 Hoya species may occur in Sabah

alone and it is becoming apparent that the diversity of Hoya of Borneo is expected to

be comparable to or exceeding that of the Philippines with 104 species (Aurigue et al.,

2013) or of New Guinea with 74 species (Forster, 1996).

During a recent expedition in West Kalimantan Mr Sulaiman Hasim discovered

a new Hoya species that we formally describe and illustrate here. It is compared with

the vegetatively similar Hoya darwini and H mitrata. The latter was lectotypified in

Rodda (2012) while the former is here lectotypified.

Hoya undulata S. Rahayu & Rodda sp. nov.

Similar to Hoya mitrata Kerr and H. darwinii Loher when sterile because it has

specialised convex leaves foiming multileaved domatia. Easily separated when fertile

because the flowers have a rotate corolla formed by almost completely free corolla

lobes, while Hoya mitrata and H. danvinii have reflexed corollas with a tube almost as

long as the lobes. - TYPE: Indonesia, West Kalimantan, Putussibau, 300-380 m, on a

slope above a stream, February 2014, Sulaiman Hasim s.n. (holotype BO). (Fig. 1, 2)

Epiphytic climber with white latex in all vegetative parts, glabrous. Stems slender,

intemodes very variable in length, from 0.5 cm in the parts ofthe vine forming domatia, to

20 cm long in climbing stems, 3-5 mm diameter, green with darker spots; adventitious

root sparsely produced along the stem. Leaves petiolate; petiole broader than stem,

0.5-2. 5 cm long, 3-5 mm in diameter, light green to purple; lamina dimorphic, the first

type ovate, obovate or oblanceolate, usually almost flat, located along climbing stems,

10-20 X 5-10 cm, the second type almost round to elliptic, usually convex and forming

domatia with 4-10 leaves each 3-7 (-10) x (2-)4-5 cm, base cordate with numerous

minute basal colleters fonning a convex line on the adaxial surface at the base of the

lamina, apex acute or acuminate, margins minutely undulate but appearing minutely

dentate in the proximal half of the lamina, sometimes entire along the distal part of the

lamina; venation pinnate, secondary veins 4-5 pairs, diverging at c. 70®, anastomosing,

tertiary venation reticulate; dark green above with grey to purple spots, below lighter

green, occasionally all purple, midrib depressed above, raised below, light green,

sometimes purple. Inflorescence positively geotropic, pseudo-umbellate, convex,

2-5-flowered(-12 flowered in cultivated material); peduncle positively geotropic

or ageotropic, l-2(-5) cm long, 3-5 mm in diameter, green with dark purple spots,

glabrous. Flowers unscented, lasting 4-5 days in cultivation; pedicel 2. 5^.5 cm long,

1.5-2 mm in diameter, white-light green with red spots, glabrous. Calyx lobes ovate,

2-2.5 X 1.5-2 mm, light pink to purple, glabrous with ciliate translucent margins,

basal colleters one in each calyx lobe sinus, 0.3-0. 5 x c. 0.3 mm, ovate to triangular

with a round tip. Corolla rotate, concave, with corolla lobe tips inflexed, 2,5-4 cm

in diameter, 4-5 cm when flattened; corolla lobes almost free, ovate to oblanceolate.

New Hoya from Borneo

Fig. 1. Hoya undulata S.Raha)^ & Rodda (from M. Rodda MR650, SING) A. Buds. B.

Inflorescence, top view. C. Inflorescence from underneath. D. Domatia; E. Leaf, abaxial, with

a magnification of the minutely undulate margin. F. Leaf, adaxial, with a magnification of the

basal colleters. (Photos: A, B, E, F, Michele Rodda; D, Surisa Somadee)

Gard. Bull. Singapore 67(1) 2015

Fig. 2. Hoya undulata S.Rahayu & Rodda (from M. Rodda MR650, SING). A, B, C. Flower.

D. Corolla after removing the corona. E. Corona, top view. F. Corona, from underneath. G.

Corona, side view. H. Calyx and ovaries. I. Pollinarium with twin pollinia, with a magnification

of the much reduced sterile edge. (Photos: Michele Rodda)

New Hoya from Borneo

2-2.5 X 1-1.5 cm, white with purple red dots outside, cream-light pink inside, margin

recurved predominantly in the proximal half of the lobe, apex acute, curved toward the

adaxial surface, pilose inside with glabrous apex, glabrous outside. Corona staminal,

12-13.5 mm in diameter, 5-7 mm high; lobes spreading, when observed from above

clavate, 5. 5-6. 5 x 2.5-3 mm, carinate above, below sulcate with revolute margins,

outer process concave, inner apex rounded, upcurved, abruptly narrowing into a terete

linear inner process with upcurved acuminate tip; two lateral hooked appendages at

the junction between outer and inner process. Anthers ovate to almost round, c. 1.5 x

1.5 mm, with apical round membranaceous appendage just exceeding the style-head

apex. Pollinia oblong, 900-1000 x 350^00 pm, narrowing towards the base, apex

round, sterile edge much reduced, limited to the upper outer part of the pollinium;

corpusciilum rhomboid with acute tips, 500-600 x 200-250 pm; caudicle broadly

triangular, hyaline, unwinged, 250-300 x 250-300 pm at the widest. Style-head 5

angled in cross section, with 5 lobes alternating with the stamens, style-head apex

round, 1 .5 mm long, c. 3 mm broad at the base. Ovaries 2, conical with round tip,

1.5-2 mm long, c. 1 mm wide at the base, light green, red at the base and at the apex.

Fruit and seed not seen.

Distribution. Only known from the type locality in West Kalimantan, close to

Putussibau. A second unlocalised collection is widely available in cultivation.

Etymolog)?. The specific epithet refers to the characteristic undulate leaf margin.

Habitat and ecology. Observed in lowland heath forest at 300-380 m above sea level

on a slope above a stream, epiphytic on small tree trunks about 1.5 m above ground,

growing in about 80% humidity and 50% sunlight. (Sulaiman Hasim pers. comm.).

According to the epiphytic zonation by .Johansson (1975), the species grows in zone

B. The zonation of epiphytes is mainly correlated with their light, nutiients and water

requirement. Some species are restricted to strongly illuminated sites, some to shady

sites, while some avoid both strong light and deep shade and yet others have a wide

range of tolerance (Benzing, 1990). Usually, occupying the B zone means that the

species is adapted to moist shady habitats. The leaves of Hoya undulata can adapt to

high light levels, but they have been observed to develop a purple colour in intense

sunlight. At the type locality, the plants were rooting in ant nests and, on casual

observation, the leaf domatia harboured ants.

Provisional lUCN conservation assessment. Known from only one locality, the

preliminaiy conservation status of Hoya undulata is Data Deficient (DD, lUCN 2014).

Ex situ collections are present in Bogor Botanic Gardens (from the type locality) and

in Singapore Botanic Gardens (SBG acc. no. 20132428).

Notes. Two species, Hoya mitrata, from Thailand, Peninsular Malaysia, Sumatra,

Borneo, Sulawesi and Java, and H. darwinii, endemic to the Philippines, both

commonly associated with ants (Kleijn & Donkelaar, 2001), are similar to H. undulata.

Gard. Bull. Singapore 67(1) 2015

Fig. 3. Hoya mitrata Kerr (Photographed at Gunung Panti, Johor, Malaysia) A. Inflorescence.

B. A small domatium. Hoya darwinii Loher (M Rodda MR421, SING). C. Inflorescence. D.

Domatia. E. Leaves not forming domatia. (Photos: Michele Rodda)

In vegetative morphology, the three speeies have two types of leaves, the first with a

fiat or slightly eonvex lamina, occurring at widely spaced nodes on long climbing

stems, the other convex, occurring on stems with shorter intemodes, and these forming

domatia harbouring ant colonies. The first type of leaf is oblong-lanceolate in Hoya

New Hoya from Borneo

mitrata and broadly lanceolate in H. danvinii, both with an entire edge, while in H.

undulata it can be ovate, obovate or oblanceolate, with a minutely undulate margin

which may appear minutely dentate in the proximal half of the lamina, sometimes

entire along the distal part of the lamina. In Hoya danvinii the domatia are globose,

4-7 cm in diameter and fonned by 2-6 convex leaves with an inrolled edge (Fig.

3D). The domatia of Hoya mitrata are instead cabbage-shaped, usually larger, fonned

by tightly clustered broadly obovate convex leaves without inrolled edges (Fig. 3B)

(Weissflog et al., 1999; Kleijn & Donkelaar, 2001). In its natural environment, the

domatia of Hoya undulata are generally looser than those of//, mitrata or//, dai^vinii,

formed by 4-10, round to elliptic convex leaves 3-7 (-1 0) cm long, while in cultivation

domatia rarely develop and are usually formed by larger leaves (Fig. ID),

The inflorescences of Hoya darwinii and H. mitrata are flat to slightly convex,

negatively geotropic, bearing 10-15 flowers (Fig. 3 A, C), the corolla has a tube almost

as long as the lobes, and the lobes are reflexed; the outer process of the corona lobes

is swollen, erect. The inflorescences of Hoya undulata are also flat to slightly convex

but instead positively geotropic (Fig, lA-C), bear 2-5 flowers (exceptionally 12 in

cultivation); the corolla is concave, with almost free lobes, the corona lobes outer

process is concave, spreading with an upcurved apex.

The pollinia of Hoya darwnnii and H mitrata do not have a sterile edge, a

character that led Kloppenburg (1994) to accommodate both species in the new section

Rudimentalia Kloppenb. The pollinaria of Hoya undulata are overall similar to those

of //. mitrata and //. darwinii but its pollinia instead have a sterile edge, albeit much

reduced, limited to the upper outer side of the pollinium (Fig. 21).

A striking similarity can be observed between the corona of Hoya undulata and

H. griffithii from mainland Asia. The corona lobes are clavate in both species, with two

lateral hooked appendages at the junction between the outer and the inner processes.

These appendages have not been documented in any other Hoya species.

A key to separate Hoya danvinii, H. mitrata and H. undulata is provided below.

Other specimens examined. Unlocalised, 30 March 2014, Rodda, M. MR650, collected from

material cultivated at the Singapore Botanic Gardens, acc. no. 20132428 (SING).

Key to Hoya species forming domatia

la. Inflorescence negatively geotropic with 10-15 flowers, outer process of corona

lobes swollen, erect 2

lb. Inflorescence positively geotropic, with 2-5 (-12) flowers, outer process of corona

lobes concave, spreading with upcurved apex Hoya undulata

2a. Domatia globose, formed by concave leaves with inrolled edges

Hoya darwinii

2b. Domatia cabbage-shaped, formed by vaulted leaves without inrolled edges

Hoya mitrata

Gard. Bull. Singapore 67(1) 2015

Lectotypification of Hoya darwinii

Hoya darwinii Loher, Gard. Chron. 47: 66 (1910). - TYPE: Philippines, Luzon, Rizal

Prov., September 1909, Loher, A. 14574 (lectotype UC [UC243291], designated here).

When Loher described darwinii he did not cite any specimens as belonging to the

taxon. He only indicated that the species was collected in Luzon and that it bloomed

in March and April. Loher collected in Luzon over a long period of time, from 1889

to 1915. His specimens collected before 1906 were deposited at K, with substantial

sets of duplicates in PNH, US, M, CAL; his collections from 1908 to 1915 (numbers

12000 to 15170) were deposited in PNH (Merrill, 1925), with a first duplicate set sent

to M and additional specimens to UC and A (orchids only) (Steenis-Kruseman, 1950).

We have examined Hoya specimens at K, and made enquires to M, UC, PNH and US

about the presence of any relevant Loher material. At K we located numerous early

Loher specimens, none identifiable as Hoya daminir, at M we found post- 1908 Loher

specimens, once again not belonging to H, danvinii. No Hoya darwinii specimens

collected by Loher could be found at PNH or US. However, three specimens of Hoya

danvinii were found at UC: two duplicates of Loher 14990, collected in Luzon, Rizal

Prov., in 1915, one of which is also labelled 395 (this label should be removed

as it likely refers to a different Hoya species with yellow flowers, while H darwinii

generally has pink flowers), and Loher 14574, collected in Luzon in September 1909

(Fig. 4). The latter is the only available specimen collected before the publication date

of Hoya danvinii and most probably belongs to the original material on which this

taxon is based. Thus, it is the only specimen available for lectotypification. However,

it must be noted that it does not fully match Loher ’s description as the flowering

period was indicated as March-April. If new evidence were to be found to indicate

this specimen is not original material, and in the absence of the discovery of alternative

original material, then the lectotypification designated here would count as an effective

neotypification under Art. 9.9 of the ICN (McNeill et al., 2012).

ACKNOWLEDGEMENTS. We would like to thank Mr Sulaiman Hasim who provided material

of the new species for study and cultivation in Bogor Botanic Gardens; Yoyo Budiman for his

assistance in transferring the material, Teguh (BBG) for assistance in pressing the materials and

Surisa Somadee, who provided photographs oi Hoya undidata. We also thank the curators of

BCU, BK, BKF, BM, BO, BRUN, FI, K, KEP, L, LAE, M, P, PNH, SAN, SAR, SNP, SING,

UC, US and UPM herbaria for allowing access and/or for providing high quality images of

herbarium specimens, an anonymous reviewer and Dr David Middleton, Editor of Gardens’

Bulletin Singapore, for suggesting improvements to this manuscript.

New Hoya from Borneo

References

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New Species from Samar and Biliran Islands, Philippines. J. Nat. Stud. 12 (1): 12-17.

Benzing, D.H. ( 1 990). Vascular epiphytes: general biology and related biota. Cambridge Univ.

Press, Cambridge..

Forster, P.l. ( 1 996). A checklist of the Asclepiadaceae of Papua. Sci. New’ Guinea 22(1 ): 1 5-22.

Forster, PL, Liddle, D.J., & Liddle, I.M. (1998). Diversity in the genus Hoya (Asclepiadaceae-

Marsdenieae). Aloe 35: 44^8,

Johansson, D.R. (1975). Ecology of epiphytic orchids in West African rain forests. Amer.

Orchid Soc. Bull. 44: 125-136.

Kleijn, D. & van Donkelaar, R. (2001). Notes on the taxonomy and ecology of the genus Hoya

{Asclepiadaceae) in Central Sulawesi. Bluniea 46: 457-483.

Kloppenburg, R.D. (1994). Hoya sections. Fraterna 1994(1&2): I-XXIV.

lUCN Standards and Petitions Subcommittee (2014). Guidelines for Using the lUCN Red

List Categories and Criteria. Version 10.1. http://www.iucnredlist.org/documents/

RedListGuidelines.pdf (accessed on 20 Jan. 2014).

Lamb, A., Gavrus, A., Emoi, B. & Gokusing, L. (2014). The hoyas of Sabah, a commentary

with seven new species and a new subspecies. Sandakania 1 9: 1-89.

McNeill, J., Buck, W.R., Demoulin, V., Greuter, W., Hawkworth, D.L., Herendeen, P.S., Knapp,

S., Marhold, K., Prado, J., Pmd’homme van Reine, W.F., Smith, G.F., Wiersema, .T.H. &

Turland, N.J. (2012). International Code of Nomenclature for algae, fungi, and plants

(Melbourne Code). Regnum Vegetabile 154. 205 p. Konigstein: Koeltz Scientific Books.

Merrill, E.D. (1921). A bibliographic enumeration of Bornean plants. J. Straits Branch Roy.

Asiat. Soc., Special Number. Singapore: Fraser & Neave Ltd.

Merrill, E.D. (1925). New species of Philippine plants collected by A. Loher. Philipp. J. Sci.

27: 21-59.

Rahayu, S. (2006). Keanekai'agaman jenis Hoya di Hutan Lindung Bukit Batikap, Pegunungan

Muler, Kalimantan Tengah. Biodiversitas 7(2): 139-142.

Rintz, R.E. (1978). The Peninsular Malaysian species of Hoya {Asclepiadaceae). Malayan Nat.

J. 30: 467-522.

Rodda, M. (2012). (2074) Proposal to conserve the name Hoya mitrata against H. wallichiana

{Apocynaceae, Asclepiadoideae). Taxon 61(3): 685-686.

Rodda, M. & Ercole, E. (2014). Hoya papaschonii (Apocynaceae: Asclepiadoideae), a new

species from southern Thailand with a peculiar corona. Phytotaxa 175(2): 097-106.

Rodda, M. & Simonsson, N. (2011a). Hoya devogelii {Apocynaceae: Asclepiadoideae), a new

species from Sarawak, Borneo. Webbia 66(1): 33-38.

Rodda, M, & Simonsson, N. (2011b). Hoya medinillifolia (Apocynaceae: Asclepiadoideae), a

new species from lowland forest of Sarawak, Borneo. Webbia 66(2): 149-154.

Rodda, M. & Simonsson Juhonewe,N. (20 13a). Hoya nuttiana (Apocynaceae: Asclepiadoideae),

a new species from Sarawak, Malaysian Borneo. Phytotaxa 140 (1): 56-60.

Rodda, M. & Simonsson Juhonewe, N. (2013b). The taxonomy of Hoya micrantha and Hoya

revoluta (Apocynaceae, Asclepiadoideae). Webbia 68(1): 7-16.

Rodda, M., Simonsson Juhonewe, N. & Ercole, E. (2013). Hoya coiymbosa (Apocynaceae:

Asclepiadoideae), a new unusual species from Sabah, Borneo and its systematic position

based on phylogenetic analyses. Syst. Bot. 38 (4): 1125-1131.

Gard. Bull. Singapore 67(1) 2015

Rodda, M., Simonsson Juhonewe, N. & Rahayu, S. (2014). Taxonomic revision of the Hoya

mindorensis complex (Apocynaceae:-Asclepiadoideae). Webbia 69(1): 39^7.

Steenis-Kruseman, M.J. van (1950). Malaysian plant collectors and collections: Being a

cyclopaedia of botanical exploration in Malaysia and a guide to the concerned literature

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Weissflog, A., Moog, J., Federle, W., Wernr, M., Hashim, R. & Maschwitz, U. (1999). Hoya

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Gardens’ Bulletin Singapore 67(1): 95-99. 2015

doi: 10.3850/S2382581215000101

Begonia bimaensis, a new species of Begonia from

Sumbawa Island, Indonesia

N.K.E. Undaharta^, I Made Ardaka, Agung Kumiawan & Bayu Adjie

Bali Botanic Garden - Indonesian Institute of Sciences

Candilaining, Baturiti, Tabanan 82191, Indonesia

'nika002@lipi.go.id

ABSTRACT. A new species of Begonia sect. Reichenheimea, B. bimaensis Undaharta &

Ardaka, is described from Mt Muria in Binia District, Sumbawa Island, Indonesia. The species

is a narrow endemic, currently known from only one population, and has been assessed as

Vulnerable using lUCN criteria.

Keywords. Begonia, Sumbawa, new species

Introduction

The pantropical genus Begonia L. is one of the largest genera of flowering plants,

comprising more than 1700 species (Hughes, 2008 and the many species described

since then). Southeast Asia is a hotspot of species diversity for Begonia (Hughes,

2008), with Indonesia being particularly rich in species and with many species still

being discovered (Hughes, 2008; Thomas & Hughes, 2008; Girmansyah, 2009, 2012;

Girmansyah et ah, 2009; Hughes et ah, 2009; Thomas et ah, 2009, 2011; Ardi &

Hughes, 2010; Wiriadinata, 2011; Ardi et ah, 2013, 2014; Ardi, 2014; Lin & Peng,

2014). From these works the current estimate for the total number of Begonia species

in Indonesia is 213 species.

There are only a few Begonia species recorded for Nusa Tenggara Barat (West

Nusa Tenggara), and currently only two species have been reliably recorded from

the largest island, Sumbawa {B. multangula Blume - Hughes & Pullan, 2007; and B.

muricata Blume - Hughes, 2008). Examination of herbarium specimens shows that

several endemic species remain to be described from the region. The lack of records

for widely distributed species, such as Begonia longifolia Blume, may also reflect the

poor documentation of the Begonia flora of Sumbawa, since we have seen herbarium

material of this species from the sun'ounding islands of Bali, Lombok, Sumba and

Timor.

During a fieldtrip to Sumbawa Island, a small population of an unknown Begonia

from Mount Maria was found. Two living specimens were brought to the Bali Botanic

Garden where they were cultivated. As these specimens could not be identified, the

new species Begonia bimaensis is described here. Begonia bimaensis is placed in

Begonia section Reichenheimia (Klotzsch) A.DC. as it exhibits the characters typical

of the section: rhizomatous or tuberous stems, protandrous inflorescences, and three

locular ovaries with entire placentae (Doorenbos et ah, 1998).

Card. Bull. Singapore 67(1) 2015

All available Begonia specimens from the Herbarium of Bali Botanic Garden,

BO, E, K, L and SfNG have been consulted and hence it must be assumed, at least until

more intensive collecting in Sumbawa reveals otherwise, that the species described

here has a very restricted range.

Species description

Begonia bimaensis Undaharta & Ardaka sp. nov. § Reichenheimia

Similar to Begonia sendangensis Ardi but differs in the dense white, long hairs on

the adaxial surface of leaves (versus glabrous), female flower with four tepals (versus

three) and the ovary with 3 wings, reddish, equal or subequal, locules 3, placentation

axillary. Ovary characters for Begonia sendangensis, pinkish, equal, rounded at base,

placentae axile. - TYPE: Cultivated at Bali Botanical Garden from vegetative material

collected in the wild from Indonesia, West Nusa Tenggara, Sumbawa Island, Bima,

Wawo, Ntori, Mt. Maria, 08°29'55"S 118°52'41"E, 525 m elev., 4 October 2010.

Cultivated material vouchered and selected as type material on 11 December 2012

under I Made Suja HK 1276 (holotype Herbarium Bali Botanic Garden; isotype BO).

(Fig 1)

Perennial, tuberous, monoecious small herb, less than 10 cm tall. Stems very reduced,

an elongated tuber 1.5-1 . 8 cm long, diameter 3-4 mm, intemodes c. 1 mm long;

stipules ovate to elliptic, c. 3 x 1-2 mm, pale green, apex projecting up to c. 6 mm long,

persistent. Leaves alternate; petioles 3. 5-7. 5 cm long, red, sparsely covered with long

pilose hairs c. 2-5 mm long; lamina basifixed, 4-6.5 x 2.5-5 cm, ovate to broadly ovate,

asymmetric, base cordate, lobes not or rarely overlapping, apex acuminate, margin

shallowly to moderately lobed, ciliate with a fringe of hairs; adaxial surface green to

reddish green, densely covered with white pilose hairs, abaxial surface red, glabrous;

venation palmate, red, primary veins 5-6, adaxially green to brownish, abaxially

pale red. Inflorescence a simple dichasial cyme, few-flowered, axillary, protandrous,

bisexual; peduncle 3-6 cm long, pink, glabrous; bracts minute, sub-orbicular, margin

slightly fimbriate, deciduous. Male flowers: pedicels 15-18 mm long; tepals 4, pink,

glabrous, two outer tepals ovate to suborbicular, 8-14 x 8-11 mm, apex rounded, two

inner tepals narrowly obovate, 5-12 x 4-6 mm, tip slightly obtuse, androecium yellow,

symmetric, globose; stamens c. 40, filaments fused at base into short column c. 1 mm

long, anthers c. 0.5 mm long, obovate, dehiscing through lateral slits more than half

the length of the anther, apex slightly retuse. Female flowers: pedicels 2.5^ mm long,

bracteoles present, with hairs, ovate, c. 2 mm long, persistent; tepals 4, pink, glabrous,

two outer tepals suborbicular, c. 9.5-12 x 7.5-12.5 mm; glabrous, two inner tepals

elliptic, c. 8-10 x 3.5-5 mm; ovary 3^ x 4 mm (excluding wings), ellipsoid, glabrous,

greenish white, locules 3, placentation axile, wings 3, equal or sub equal, reddish green

triangular, widest point at the middle of the ovary, c. 1 .5 mm long; stigma 3, U-shaped,

stigmatic surface twisted. Fruit: pedicel 2-5 mm long, capsule ovoid, 4-5 x 5-6.5 mm

(excluding wings), dehiscent, splitting along the wing attachments, wing shape as for

Begonia bimaensis, a new species from Indonesia

Fig. 1. Begonia bimaensis from cultivated material in Bali Botanie Garden. A-B. Habit; C.

Leaf, adaxial surfaee; D. Leaf, abaxial surfaee; E. Inflorescence; F. Female flower; G. Female

flower; H. Male flower; I. Bracts; J. Ovary transverse seetion; K. Tubers; Scale bars: 1 cm.

(Photos: Gede Wawan Setiadi)

Card. Bull. Singapore 67(1) 2015

the ovary, wings widest subapically, 8 mm. Seeds numerous, brown, widely ellipsoid

or ellipsoid, c. 0.35 mm long, 0.2 wide.

Distribution. Endemic to Mt Muria, Bima district, Sumbawa, West Nusa Tenggara, Indonesia.

Habitat. This species grows on steep rocky cliffs in shade, at c. 525 m altitude.

Preliminaty lUCN conservation assessment. Begonia bimaensis is probably a narrowly

endemic species, restricted to Mt Muria which is not fonnally protected. As the species is

known only from a single population with a small number of individuals, a provisonal lUCN

category of CR D is appropriate (lUCN, 2012).

Notes. The epithet 'bimaensis' refers to the Bima district from where the type material

was collected. Begonia bimaensis is unusual in Begonia section Reichenheimia in

Indonesia on account of its tuberous habit, which is similar to Begonia sendangensis, a

species recently described from the neighbouring island of Lombok ( Ardi et. al, 2014).

However, apart from the presence of tubers, the two species are morphologically

dissimilar and Begonia bimaensis can easily be distinguished by the dense white long

hairs on the upper surface of the leaves. The number of tepals in the female flower

and the shape of the ovary wings provide further differences; The female flowers of

Begonia bimaensis have four tepals whereas the female flowers of B. sendangensis

have three tepals; in B. bimaensis, the shape of the wings in the ovaiy is very distinctive,

being triangular and mostly cuneate at apex (versus rounded at base and truncate at the

apex), reddish.

ACKNOWLEDGEMENTS. The authors would like to express their gratitude to Nyoman

Sudiatna for collecting the living specimen (during a fieldtrip supported by the Research

Incentive for Researchers and Engineers No. 053/m/Kp/II/2010 to AK), and to the curators

of the Herbarium of Bali Botanic Garden, BO, E, K, L and SING for allowing us access to

herbarium material. We also thank Mark Hughes (Royal Botanic Garden Edinburgh) for editing

and correcting the English and Wisnu Ardi (Bogor Botanic Gardens) for his assistance.

References

Ardi, W.H. (2014). Two new species of Begonia (Begoniaceae) from Sulawesi, Indonesia.

Edinburgh J. Bot, 71: 259-268.

Ardi, WH. & Hughes, M. (2010). Begonia droopiae Ardi (Begoniaceae). A new species of

Begonia from West Sumatra. Card. Bull. Singapore 62: 1 7-22.

Ardi, W.H., Ardhaka, I.M., Hughes, M., Undaharta, N.K.E., Ginnansyah, D. & Hidayat, S.

(201 3). Two new species of Begonia (Begoniaceae) from Bali and Lombok. Card. Bull.

Singapore 65: 135-142.

Begonia bimaensis, a new species from Indonesia

Ardi, W.H., Kusuma, Y.W.C., Lewis, C.E., Risna, R.R., Wiiiadinata, H., Abdo, M.E. & Thomas,

D.C. (2014). Studies on Begonia (Begoniaceae) of the Molucca Islands I: Two new

species from Halmahera, Indonesia, and updated description of Begonia holosericea.

Reinwardtia 14: 19-26.

Lin, C.-W. & Peng, C.-I. (2014). Begonia natunaensis (Sect. Reichenheimia, Begoniaceae), a

new species from Natuna Island, Indonesia. Taiwania 59: 368-373.

Doorenbos, J., Sosef, M.S.M. & de Wilde, J.J.F.E. (1998). The sections of Begonia including

descriptions, keys and species lists (Studies mBegoniacomW). Agric. Univ. Wageningen

Pap. 98: 1-266.

Girmansyah, D. (2009). A taxonomic study of Bali and Lombok Begonia {BegoniacoeiQ).

Reinwardtia 12: 419-434.

Ginnansyah, D. (2012). Begonia ranaiensis (Begoniaceae), a new species from Mt Ranai,

Natuna Island, Indonesia. Kew Bull. 68: 1^.

Girmansyah, D., Wiriadinata, H. & Thomas, D.C. (2009). Two new species and one new

subspecies of Begonia (Begoniaceae) from Southeast Sulawesi, Sulawesi, Indonesia.

Reinwardtia 13: 69-74.

Hughes, M. (2008). An Annotated Checklist of Southeast Asian Begonia. Edinburgh: Royal

Botanic Garden Edinburgh.

Hughes, M. & Pullan, M. (2007). Southeast Asian Begonia Database. Royal Botanic Garden

Edinburgh, http ://elmer,rbge.org.uk/5ego«/n!/ (accessed on 18 Aug. 2014).

Hughes, M., Girmansyah, D., Ardi, W.H. & Nurainas (2009). Seven new species of Begonia

from Sumatra. Card. Bull. Singapore 61: 29-44.

lUCN (20 1 2). lUCN Red List Categories and Criteria: Version 3. 1 . Second Edition. Switzerland,

Gland and UK, Cambridge: lUCN.

Thomas, D.C. & Hughes, M. (2008). Begonia varipeltata (Begoniaceae): Anew peltate species

from Sulawesi, Indonesia. Edinburgh J. Bot. 65: 369-374.

Thomas, D.C., Ardi, W.H. & Hughes, M. (2011). Nine new species of Begonia (Begoniaceae)

from South and West Sulawesi, Indonesia. Edinbutgh J. Bot. 68: 225-255.

Thomas, D.C, Ardi, W.H., Hartutiningsili & Hughes, M. (2009). Two new species of Begonia

(Begoniaceae) from South Sulawesi, Indonesia. Edinburgh J. Bot. 66: 229-238.

Wiriadinata, H. (2011). A new species of Begonia (Begoniaceae) from Sagea Lagoon, Weda

Bay, Halmahera Island, North Moluccas Indonesia. Reinwardtia 13: 263-270.

Gardens’ Bulletin Singapore 67(1): 101-106. 2015

doi: 10.3850/S2382581215000113

101

A new species of Paraboea (Gesneriaceae)

from Thailand

C. Puglisi^, S. Suddee^, R Triboun^ & DJ. Middleton^

'Royal Botanic Garden Edinburgh, 20A Inverleith Row,

Edinburgh EH3 SLR, Scotland, U.K.

c.puglisi@rbge.ac.uk

^The Forest Herbarium, Department of National Parks,

Wildlife and Plant Conservation,

Bangkok, 1 0900, Thailand

^Thailand Institute of Scientific and Technological Research,

Technopolis, Klong 5, Klong Luang,

Pathumthani 12120, Thailand

"'Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, Singapore 259569

ABSTRACT. A new species, Paraboea maculata C.Puglisi, is described. It is compared to

similar species and the habitat on granite rocks, unusual for the genus, is highlighted.

Keywords. Gesneriaceae, Paraboea, Thailand

Introduction

A Paraboea (C.B. Clarke) Ridl. specimen, collected in fruit in Kliao Khitchakut

National Park in Chanthaburi, Thailand, in 2012, was recognised at the time as likely

to be an undescribed species but was not published due to lack of flowering material.

It has now flowered in the Royal Botanic Garden Edinburgh and is here described as

a species new to science.

Paraboea was revised by Xu et al. (2008) but since then the genus has been

considerably enlarged through the inclusion of Trisepalum C.B. Clarke and Phylloboea

Benth. (Puglisi et al., 2011) and by the description of many more species (Chen et al.,

2008; Kiew, 2010, 2012; Chen et al., 2012; Triboun & Middleton, 2012; Xu et al.,

2012; Triboun, 2013; Wen et al., 2013). There are now over 130 species in the genus

with many species being very locally endemic (Xu et al., 2008) which brings with it

conservation concerns (Xu et al., 2008; Triboun & Middleton, 2012). The centre of

diversity of the genus is in Thailand with over 75 species but it is likely that there are

many undiscovered species in neighbouring countries.

Paraboea species are lithophytes, mostly on limestone substrates. A small

number of species are recorded from other substrates, such as Paraboea elegans

(Ridl.) B.L.Burtt (southern Thailand, Peninsular Malaysia) and P. gratiiticola Z.R.Xu

(southern central Vietnam) from granite, and P. pubicorolla Z.R.Xu & B.L.Burtt and

102

Card. Bull. Singapore 67(1) 2015

Paraboea lavandulodora Triboun from sandstone. In Paraboea these are a small

number of exceptions to the otherwise overwhelming preference for limestone. In

Thailand this results in a very uneven distribution of the large number of species,

with very few in the relatively limestone-poor areas of southeastern and northeastern

Thailand.

In Xu et al. (2008) only one collection of 3 . Paraboea species was reported from

the southeastern province of Chanthaburi. No further collections from Chanthaburi

were reported by Triboun & Middleton (2012) and Triboun (2013). Therefore, when a

Paraboea species was collected from granite rocks in Kliao Khitchakut National Park

in Chanthaburi in 2012, it was only the second known collection of any species of the

genus from the province and only the third collection from the whole southeastern

region (the third collection being from the island of Ko Chang). The Khao Khitchakut

plant has a temiinal inflorescence with leaf-like bracts towards the base of the peduncle,

characters which place the species in the Paraboea martinii group (Xu et ah, 2008).

When only known from the fruiting collection, it could already be determined that the

Kliao Kliitchakut plant was not one of the described species in that alliance nor one of

the other species recorded from the southeastern region by Xu et al. (2008). With the

flowering material now available this is confirmed and the species is here described.

Paraboea maculata C.Puglisi, sp. nov.

Differs from other Paraboea species in the Paraboea martinii group, i.e. those with

opposite leaves, campanulate corolla, twisted fruit and a terminal inflorescence, by the

combination of a predominantly white and laterally compressed corolla, the purple

spots at the base of the tube, the sticky glandular secretion on the bracts and calyx,

and the winged petioles. - TYPE: Thailand, Chanthaburi, Khao Khitchakut, Khao

Khitchakut National Park, Kliao Phra Bhat, 1 2°50' 1 4"N 1 02° 1 0'3 "E, 900 m, 27 August

2012, fr., Middleton, D.J., Karaket, P, Suddee, S. & Triboun, P 5675 (holotype E;

isotypes BK, BKF). (Fig. 1)

Lithophytic, caulescent herb to 60 cm high. Stem short and erect, with an indumentum

of sessile and stalked glands, and sparse multicellular, eglandular hairs. Leaves

opposite, congested; petioles winged, 2-9 cm long, those of a pair fonning a distinct

auricle across the node; lamina lanceolate to ovate, 8-19 x 4-10 cm, about twice as

long as wide, surface rugose, apex acute, base rounded and then shortly attenuate

onto the wing of the petiole, margin irregularly crenate, indumentum arachnoid and

deciduous above, with white glands and a pale, thin, interwoven, semi-deciduous

indumentum below, secondai*y veins 10-13 pairs, tertiary veins reticulate, venation

prominent below. Inflorescence a terminal panicle, 25-30 cm long, occasionally

flanked by subtenninal cymes, with at least 4 orders of branching, axes deep red-purple

to bright brown, appearing chestnut brown in herbarium specimens, with a diffuse

arachnoid indumentum or glabrous, bearing a series of sessile, leaf-like, opposite and

decussate pairs of bracts of a progressively decreasing size, with intemodes 5-15 cm

long; uppermost bracts 0.4-1 cm long, 0.3-0. 6 cm wide at the base, sessile, joined at

A new Paraboea from Thailand

103

Fig. 1. Paraboea maculata C.Puglisi. A. Inflorescence. B. Winged petiole. C. Corolla, front

view. D. Corolla and calyx, side view. (Photos: A, C, D: Lynsey Wilson; B: Sadie Barber)

104

Card. Bull. Singapore 67(1) 2015

the base or nearly so, deltoid, concave at the base, proximally red or brown, turning

green towards the tip, densely covered in sessile glands and sticky on both sides but

particularly so on the outer, which is glossy due to an exudate; pedicels 1-1,6 cm long,

glabrous. Calyx with lobes divided to the base, resembling terminal bracts in colour,

indumentum and glossiness; lobes 3-5 mm long, c. 1 mm wide, linear to nan'owly

lanceolate, apex broadly acute. Corolla strongly zygomorphic, campanulate, white

with dark purple-red markings inside towards the base of the tube and around the

staminode insertions, visible from the outer side, laterally and above as a pink shade;

tube laterally compressed with the exception of two lateral bulges miming along its

length, ventrally with 3 narrower bulges, separated by two depressions, running to the

base of the central lower lobe; covered in minute glandular hairs outside making the

corolla sticky but without the glossy appearance of the bracts and calyx; tube 9-1 3 mm

long, slightly oblique, mouth compressed; upper lobes 4-5 x 9-1 1 mm, elliptic; lateral

lobes of the lower lip elliptic, apex obtuse, 5-8 x 5-6 mm; central lobe 6-10 x 4-9 mm

(4 mm at the base widening to 9 mm), slightly folded around the central of the three

bulges running along the tube, apex obtuse, margin undulate. Stamens arising from

the base of the corolla; filaments 0.8-1 cm long, purple both proximally and distally,

with a white to bright yellow knee in the middle, glabrous in the purple areas, densely

covered in glandular hairs on the knee; anthers coherent, 1-2 x 3-3.5 mm, white, with

the connectives and tips of the thecae tinged with purple; thecae strongly divergent;

staminodes 3, white, the lateral ones 1-1.5 imn long, arising 1.5-2 mm above the base

of the tube, with scattered glandular hairs along the filament, the central staminodes c.

0.5 mm long, arising c. 2.5 mm above the base of the tube, glabrous. Disc annular, pale

yellow, c. 0.5 mm long. Gynoecium 11-12 mm long, glabrous; ovary 5-7 mm long,

indistinctly developing into a c. 5 mm long style; style enantiostylous, apically bent,

stigma reduced. Fruit (immature) a strongly twisted capsule, glabrous, green, 2^ cm

long, 1-1.5 mm diameter. Seeds numerous, ellipsoid, slightly compressed, 0.5-0. 7 x

0.15-0.2 mm.

Distribution. Only known from the type locality

Ecology. Evergreen forest, on granite bedrock in deep shade.

Etymology. The epithet refers to the purple markings inside the corolla tube.

Provisional lUCN conservation assessment. Data Deficient (DD). This species is

currently only known from one locality within a National Park. Although there is

some disturbance due to tourism at this one site, it is uncertain how extensive these

possible threats are. There is less than 400 km- of suitable habitat (forest of over 800

m altitude) in the immediate vicinity of the collection locality, which would suggest

an assessment of Endangered based on a restricted Extent of Occurrence if there were

corresponding threats. Similar forest types occur in neighbouring parts of Cambodia,

where the threat levels are even less well known. Even if these forests are included,

the EOO remains less than 2000 km^, still within the boundaries of Endangered based

A new Paraboea from Thailand

105

on EOO. However, a clearer assessment of the populations, the distribution and the

threats would first have to be made before an accurate assessment could be proposed.

Additional specimens examined. THAILAND: Chantanaburi: Khao Khitchakut, Kliao

Khitchakut National Park, Khao Phra Bhat, Middleton, D.J. et al. 5675, cultivated as RBGE

20121417, vouchered 25 Jun 2014 as Puglisi, C. CP2 506 14/1 (E); ibidem, vouchered 15 July

2014 as Atkins, H.J. 18 (SING).

Aote.?.This new species is unusual in being one of the very few species in the genus

to occur on a substrate other than limestone. It quite clearly belongs in the Paraboea

martinii group (Xu et al., 2008), which is characterised by opposite and decussate

leaves, tenninal inflorescence, reduced leaf-like bracts at the base of the peduncle, a

large campanulate corolla, distinct limb, filaments with a bearded knee, and twisted

capsules. Other species in this group include Paraboea doitungensis Triboun &

D.J.Middleton, P. glutinosa (Hand.-Mazz.) K.Y.Pan, P. paramartinii Z.R.Xu &

B.L.Burtt and P. thorelii (Pellegr.) B.L.Burtt. It differs from Paraboea doitungensis

in the shorter calyx lobes, the flower colour, the laterally compressed corolla opening,

the lateral swellings, and the longer fruit; it differs from Paraboea glutinosa in the

broader leaves and the laterally compressed corolla opening; it differs from Paraboea

martinii in the winged petiole, the lack of a matted or glandular indumentum on the

floral axis and the compressed corolla; it differs fi-om Paraboea paramartinii in the

shorter petioles, the broader petiole wings and the more deeply crenate leaf margin;

and it differs from Paraboea thorelii in the broader leaves, lack of glandular hairs on

the inflorescence axes, laterally compressed corolla opening, calyx indumentum, and

the pubescence on the upper leaf surface. We have also compared it to the material

from Cambodia and Vietnam identified provisionally as Paraboea cf. glutinosa by Xu

et al. (2008). It is not the same species as Poilane 28717 from Cambodia, which has

a different leaf indumentum, smaller upper bracts and calyx (but which is otherwise

very similar to Paraboea maculata in shape and stickiness), very much larger lower

bracts, and longer inflorescence intemodes. It is also not the same species as Poilane

16562 from Vietnam, which is an altogether hairier plant than Paraboea maculata,

with bigger bracts and no visible glands.

ACKNOWLEDGEMENTS. We thank the Director and staff of Khao Khitchakut National Park

for the opportunity to study the plants of the Park; Gunnar Ovstebo and Sadie Barber of the

Royal Botanic Garden Edinburgh for their subsequent care and cultivation of the plants; and

Lynsey Wilson and Sadie Barber for the photos.

References

Chen, W.H., Moller, M., Shui, Y.M. & Zhang, M.D. (2008). A new species of Paraboea

(Gesneriaceae) from a karst cave in Guangxi, China, and observations on variations in

flower and inflorescence architecture. Bot. J. Linn. Soc. 158: 681-688.

106

Card. Bull. Singapore 67(1) 2015

Chen, W.H., Moller, M., Zhang, M.D. & Shui, Y.M. (2012). Paraboea hekouensis and R

manhoaensis, two new species of Gesneriaceae from Ch\m..Ann. Bot. Fenn. 49; 179-

187.

Kiew, R. (2010). Two new species of Paraboea (Gesneriaceae) from Peninsular Malaysia and

Thailand. Edinburgh J. Bot. 67: 209-217.

Kiew, R. (2012). Paraboea (Gesneriaceae) in Sabah, Borneo, including a new Malayan

Nat. J. 64; 1-8.

Puglisi, C., Middleton, D.J., Triboun, P. & M5ller, M. (2011). New insights into the relationships

between Paraboea, Trisepalum and Phylloboea (Gesneriaceae) and their taxonomic

consequences. Taxon 60: 1693-1702.

Triboun, P. (2013). Paraboea middletonii (Gesneriaceae), a new species from Thailand. Thai

For. Bull (Bot.) A\: 45-41 .

Triboun, P. & Middleton, D.J. (2012). Twenty new species of Paraboea (Gesneriaceae) from

Thailand. Card. Bull. Singapore 64: 333-370.

Wen, F., Hong, X., Chen, L.Y., Zhou, S.B. & Wei, Y.G. (2013). A new species of Paraboea

(Gesneriaceae) from a karst limestone hill in southwestern Guangdong, China.

Phytotaxa 131; 1-8.

Xu, W.B., Huang, Y.S., Wei, G.F., Tan, W.N. & Liu, Y. (2012). Paraboea angustifolia

(Gesneriaceae): a new species from limestone areas in northern Guangxi, China.

Phytotaxa 62: 39-43.

Xu, Z., Burtt, B.L., Skog, L.E. & Middleton, D.J. (2008). Arevision ofParaboea (Gesneriaceae).

Edinbuj^h J. Bot. 65: 161-347.

Gardens’ Bulletin Singapore 67(1): 107-122. 2015

doi: 10.3850/S2382581215000125

107

New orchid records for Myanmar, including

the first record of the genus Stereosandra

H. KurzweiP & Saw Lwin^

^Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, Singapore 259569

h ubert_kurz\veil @nparks . gov. sg

^Myanmar Floriculturist Association, Yangon, Myanmar

myanmarorchid@gmail.com

ABSTRACT. Nine orchid species, BulhophyUum capnophyton J.J.Verm. et al., Dendrobium

macrostachyum Lindk, D. praecinctiim Rchb.T, Habenaria reniformis (D.Don) Hook.f.,

Liparis distcms C,B. Clarke, L. resupinata Ridk, Micropera thailandica (Seidenf. & Smitinand)

Garay, Oberonia obcordata Lindl. and Stereosandra javanica Blume, are reported as new for

Myanmar. Of particular interest is the Stereosandra as it is the first record of this genus in the

country.

Keywords. Myanmar, Orchidaceae, new distribution records

Introduction

The orchid flora of Myanmar is very rich but as yet poorly known (a brief overview

of the current state of orchid floristics in the country is given in Ormerod & Sathish

Kumar, 2003, 2008), and a complete inventory of all species of Orchidaceae found in

the country is not yet available. Our ongoing studies resulted in nine additions to the

orchid flora of Myanmar, namely BulhophyUum capnophyton J.J.Verm. et al. (syn.

Trias nana Seidenf.), Dendrobium macrostachyum Lindl., D. praecinctiim Rchb.f.,

Habenaria reniformis (D.Don) Hook.f., Liparis distans C.B. Clarke, L. resupinata

Ridk, Micropera thailandica (Seidenf. & Smitinand) Garay, Oberonia obcordata

Lindl. and Stereosandra javanica Blume, all of which are new records for Myanmar

of species found in adjacent countries. Stereosandra is recorded from Myanmar for

the first time.

Brief descriptions, various notes and illustrations are provided for all species.

Detailed locality data below the level of townships are not given here, in order not to

reveal localities which may endanger their populations through over-collecting. The

morphological descriptions are partly based on the specimens from Myanmar and

have been partly extracted from previously published descriptions.

Enumeration of new records

1 . Bulbophyllum capnophyton J.J.Verm., Schuit. & de Vogel, Phytotaxa 166: 110

(2014). - Trias nana Seidenf., Bot. Tidsskr. 71: 24 (1976); Seidenf., Opera Bot. 89:

108

Card. Bull. Singapore 67(1) 2015

164, fig. 106 (1986). - TYPE: Peninsular Thailand, Nam Yong, Seidenfaden, G. &

Smitinand, T GT 6096 (holotype C). (Fig. lA)

Bulbophyllum sp., Seidenf. & Smitinand, Orch. Thailand (Prelim. List): 388, fig. 294

(1961). Based on A.F.G. Kerr 0361 (BK) from Thailand.

Herb to 2 cm tall, glabrous. Pseudobulbs spaced along a creeping rhizome at intervals

of 10-14 mm, globular, somewhat flattened, 5-8 mm high, 6-10 mm in diameter,

1 -leaved. sessile, fleshy, duplicate, oblong-lanceolate, acute, 10-21 mm long.

Inflorescences 1 -flowered, scape 1-2 mm long, pedicel and ovary c. 4 mm long,

bract 1-2 mm long, hyaline, cup-shaped. Flowers light yellow, the lip deeper yellow.

Sepals spreading, oblong-elliptic, acute, 5-veined, 7-8 x 2.3-3 mm. Petals narrowly

ovate, acute, 1 -veined, 2.5-3 x c. 1.5 mm; lip oblong, obtuse, 2. 3-2. 8 x c. 1 mm,

with forward curved auricles at the base; column c. 1 .7 mm long (excluding its anther

process), stelids insignificant, anther process fleshy, oblong, somewhat spoon-shaped,

c. 1.6 mm long, apex emarginate.

Distribution. Peninsular regions of Thailand and Myanmar.

Ecology and phenology. Very little habitat infomiation on this species is available.

The specimen from Myanmar was growing as an epiphyte at an elevation of about 400

m. In neighbouring Thailand the specimen Kerr 0361 (BK) was collected in savanna

(Seidenfaden & Smitinand, 1961 : 388). In the distribution area of the species flowering

has been recorded in November and January.

Material examined. MYANMAR. Taninthayi Region: Yebyu Township, 22 Nov 2012, Saw

Lwin, Pan Khet Khet &. law Oo Wai TNRO 162 (SING, SING [spirit], herbarium of Taninthayi

Nature Reseive Education Centre).

Notes. The specimen from Myanmar, Lwin et al. TNRO 162, matches Seidenfaden’s

type very well in its stixicture (Seidenfaden, 1976a). However, it differs in the slightly

larger size of the leaf and flowers: leaf 20-21 mm long (as opposed to 10-14 mm in the

type); sepal c. 8 mm long (as opposed to c. 7 mm in the type); petal c. 3 mm long (as

opposed to 2. 5-2,8 mm in the type); lip c. 2.8 mm long (as opposed to 2. 3-2. 5 mm in

the type). The epithet nanum was not available in Bulbophyllum when Trias nana was

moved into it due to Bulbophyllum nanum De Wild.

2. Dendrobium macrostachyum Lindl., Gen. Sp. Orchid. PI. 78 (1830). - Callista

macrostachya (Lindl.) Kuntze, Revis. Gen. PI. 2: 655 (1891). - TYPE: Sri Lanka,

1829, Macrae, J. 77 (holotype K-LINDL! [K000364613]). (Fig. 2A)

Dendrobium stuartii F.M.Bailey, Proc. Roy. Soc. Queensland 1: 12 (1884);

Vaddhanaphuti, Wild Orch. Thailand, ed. 4, 125, inch colour photo (2005); Seidenf. &

New records of Myanmar orchids

109

Fig. 1. A. Anther of Bulbophylliim capnophyton JJ.Verm., Schuit. & de Vogel. From Lwin

et al TNRO 162, B. Column of Micropera thailandica (Seidenf & Smitinand) Garay. From

Lwin et al. TNRO 61. C. Inflorescence of Ohemnia obcordata Lindl. Tmm Kurzv^feil & Twin

KL 2664. D. Lip of Oberonia obcordata Lindl. From Kurzweil & twin KL 2664. E. Habit of

Stereosandra javanica Blume. from Twin et al. TNRO 5. F. Flower of Stereosandra javanica

Blume. From Lwin et al. TNRO 5. G. Lip of Stereosandra javanica Blume. From Lwin et al.

TNRO 5. Scale bars: A-D, F-G = 1 mm, E = 20 mm. (Drawn by Joshua Yang)

Card. Bull Singapore 67(1) 2015

Fig. 2. A. Dendrobium macrostachyum Lindl. Vvom Lwin MPO 050. B. Dendrobium praecinctum

Rchb.f. ¥rom Lwin SL 49. C. Habenaria reniformis (D. Don) Hook.f. Vvom Lwin et al. MPO 020.

D-E. Liparis distans C.B. Clarke. From Kurzweil & Lwin KL 2460 (Photos: Saw Lwin)

New records of Myanmar orchids

111

JJ.Wood, Orchids Penins. Malaysia Singapore 389, fig. 175, pi. 26d (1992). - Callista

stuartii (F.M.Bailey) Kiintze, Revis. Gen. PL 2: 655 (1891). TYPE: Australia, drawing

by R.D.Fitzgerald in Austral Orch. 2(3): t. 6 (1888) (neotype designated by Clements,

1989).

Dendrohium tetrodon auct. non Rchb.f. ex Lindl.: Seidenf., Opera Bot. 83: 34, fig. 13

(1985).

Herb with slender pseudobulbous and sometimes almost wiry, pendulous stems to

70 cm long, glabrous except for the lip. Stems slightly zig-zagged, intemodes 2-2.5

cm long, entirely covered by white membranous sheaths, leafy throughout. Leaves

distichous, sessile, spreading, ovate to oblong-lanceolate, acute or acuminate, 4-8 x

0.9-2. 5 cm. Inflorescences to 2.5 cm long, with l-3(^) flowers; bracts oblong-ovate,

acute, 2. 5-2. 6 x 1 .5-1 .8 mm. Flowers 20-25 mm across, sepals and petals spreading

or recurved; whitish to greenish yellow, lip with red-brown or purple veins on the

side lobes, ageing to yellow; median sepal oblong-lanceolate, acute, 13-15 x 3,5^

mm; lateral sepals similar but often slightly longer; mentum c. 6 mm long, narrowly

conical; petals elliptic-lanceolate, acute or obtuse, as wide as the sepals, 13-1 5 x 3.5^

mm; lip quadrately-ovate and very obscurely 3-lobed, 17-18 x c. 12 mm, unspurred,

hairy particularly on the margin and midrib, basal part inrolled around the column,

callus in the basal lip portion of three diverging ridges; column 3.5^ mm long.

Distribution. Widespread in tropical Asia from Sri Lanka, India and the Himalayas to

eastern Malesia; also found in northern Australia.

Ecology and phenology. The specimen from Myanmar was growing as an epiphyte

on medium-sized road-side trees at an elevation of 1150 m. Elsewhere the species is

found in wet and dry low-altitude and montane forest to 1220 m (Jayaweera, 1981;

Seidenfaden & Wood, 1992). In the distribution area of the species flowering has been

recorded in January and February.

Material examined. MYANMAR. Mandalay Region: Kyaukpadaung Township, Jan 2010,

Saw Lwin MPO 050 (SING, SING [spirit]). Very uncommon in this locality, growing in a small

population of about 20 individuals.

Notes. A widespread species with rather small flowers in bunches of mostly 1-3 on

leafless stem. The taxonomy of this species was previously disputed (Christenson

& Wood, 2003), in particular its relationship to Dendrohium aphyilum (Roxb.)

C.E.C.Fisch., but this has now been clarified (Schuiteman, 2011).

3. Dendrohium praecinctum Rchb.f., Gard. Chron., n.s., 7: 750 (1877); N.Pearce &

RJ.Cribb, FI. Bhutan 3(3): 416, pi. 23 [top right] (2002); Jin et al., Acta Bot. Yunnan.

32: 332 (2010). - Callista praecincta (Rchb.f.) Kuntze, Revis. Gen. PI. 2: 655 (1891).

TYPE: Unlocalised, cult. Veitch s.n. (holotype W). (Fig. 2B)

112

Card. Bull. Singapore 67(1) 2015

Dendrobium pauciflorum King & Pantl., J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 64: 332

(1895); Seidenf., Opera Bot. 83: 97, fig. 57, pi. XIIc {19S5), - Dendrobium sikkimense

A.D.Hawkes & A.H.Heller, Lloydia20: 124 (1957), nom. illeg. -TYPE: India, Sikkim,

July 1895, Pantling, R. 172 (holotype K! [K000943939]).

Herb, pendent stems to 70 cm long, but often much shorter. Stems leafy in the distal

half, branching from nodes in the upper part of the stem, intemodes 1 .4-4 cm long.

Leaves distichous, to 12, lanceolate or linear-lanceolate, acute, sessile, 3-9 x 0.7-1. 2

cm. Inflorescences axillary, arising from apical nodes of the leafless stem, with 1-5

flowers; floral bracts ovate, obtuse, c. 2 mm long. Flowers to 20 mm long, not opening

widely; yellow to whitish yellow, sepals and petals with prominent purple marginal

areas, midlobe and side lobes of lip with red spots; pedicel and ovary 5-10 mm long;

median sepal lanceolate or lanceolate-ovate, obtuse, 3-10 x up to 4 mm; lateral

sepals ovate, weakly falcate, keeled, 4-10 x 2-4 mm; petals lanceolate-ovate, usually

ciliolate, 4-10 mm long and up to 3 mm wide; lip rather deeply 3-lobed, somewhat

clawed, 4-12 x 3-8 mm, hispid-fimbriate on the margins; lateral lobes narrow, tooth-

like; mid-lobe oblong, flat; disk with three prominent, thickened veins; column broad,

with a foot 2-7 mm long.

Distribution. Northeastern India and eastern Himalayas to Myanmar, Thailand and

southwestern China (Yunnan).

Ecology and phenology. Habitat information of the specimen from Myanmar has not

been recorded. In Sikkim and Bhutan the plants are epiphytic in evergreen broad-

leaved forest at elevations ranging from 1000 to 1330 m, and rarely up to 1850 m

(Pearce & Cribb, 2002). In the distribution area of the species flowering has been

recorded in June and July.

Material examined. MYANMAR. Shan State: Ho Pone Township, 2011, Saw Lwin SL 49

(herbarium of the Myanmar Floriculturist Association).

Notes. Easily recognised by the prominent purple edges on the pale yellow flowers

and the rather deeply 3-lobed lip which is strongly hairy to fringed on the margins.

As far as we are aware no vouchered record of the occurrence of Dendrobium

praecintum in Myanmar has previously been made. However, a photograph of an

unidentified Dendrobium species from Putao in northern Kachin State was published

by Nyan Tun (2014: 254) and was later refeiTed to this species by Paul Onnerod (pers.

comm.). Interestingly, the disnibutioii ‘Myamnar’ was also listed in a publication

newly reporting Dendrobium praecinctum in Yunnan, China (Jin et al., 2010), but no

specimen from Myanmar was cited and therefore this listing appears to be based on a

sight-record only.

New records of Myanmar orchids

113

4. Habenaria reniformis (D.Don) Hook.f., FI, Brit. India 6: 152 (1890); Seidenf.,

Dansk Bot. Ark. 31(3): 132, fig. 85 (1977); S.C.Chen & P.J.Cribb in Z.Y.Wu et al.

(eds), FI. China 25: 149 (2009); Kurzweil, FI. Thailand 12(1): 133, fig. 76 (2011). -

Listera reniformis D.Don, Prodr. FI. Nepal.: 28 (1825). - Neottia reniformis (D.Don)

Spreng., Syst. Veg. 3: 707 (1826). - TYPE: Nepal, 1821, Wallich, N. 7067 (lectotype

K-W!, designated by Seidenfaden (1977)). (Fig. 2C)

Habenaria clovisii Gagnep., Bull. Soc. Bot. France 78: 68 (1931). -TYPE: Cambodia,

Stung-streng, 1866-1868, Thorel, C. 2143 (syntypes P! [P00439699, P00439700]);

Vietnam, near Dalat, 17 Oct 1924, Evrard, F 1480 (syntype P! [P00439698]).

Herb to 20 cm tall, glabrous. Tubers globose to oblong, to 1 cm in diameter, fleshy.

Stems slender, wiry, to 1 mm in diameter. Leaves 2-4, basal and adpressed to the

substrate, orbicular, elliptic or ovate, acute or obtuse, 1.4-4 x 1-2.7 cm. Inflorescences

lax, with 3-7(-10) flowers; with or without a spreading amplexicaul leafy bract near

the base; sterile bracts 4-6(-9), mostly erect, lanceolate, acuminate, 0.5-1. 8 cm long;

rachis 3-8 cm long; floral bracts lanceolate or ovate-lanceolate, acuminate, 7-10 x

1.5- 2 mm, much shorter than the ovary. Flowers 6.5-9 mm in diameter, resupinate;

green, greenish-white, yellow-green or brownish; ovary twisted mamly in its basal

portion, cylindric-fusiform, indistinguishable from the pedicel, 13-18 mm long;

median sepal erect, concave, ovate or ovate-oblong, obtuse, 3-veined, 3.5-4(-5) x

1.5- 3 mm; lateral sepals spreading or reflexed, obliquely ovate-lanceolate, subacute

or acute, 3-veined, 3.5-4(-6) x 1.7-3. 4 mm; petals fonning a hood with the median

sepal, falcately elliptic-lanceolate or linear-triangular, subacute or obtuse, 1 -veined,

3^.5 X 0.8-1. 6 mm, sometimes widened at the base; lip 4.5-10 mm long, either

unlobed and linear, or 3-lobed with filifonn side lobes to 29 x 0. 1-0.3 mm; spur either

present or absent, if present saccate or cylindric and to 7.6 mm long; column 1-2.3

mm long; anther canals insignificant; stigmas subcylindric or clavate, 1 .5-2 mm long.

Capsule fusiform-elliptic, 9.5-10 x 2.2-3 mm.

Distribution. India and Nepal to Indochina and southern China; also recorded from

northern Sumatra.

Ecology and phenology. No ecological information on the specimens from Myanmar is

available except that the species grows terrestrially. Elsewhere found in grassy places

in forests, in bamboo thicket and scrubland at elevations ranging from 1 50 to 825 m,

sometimes also in cracks of limestone rocks (Chen et al., 2009; Kurzweil, 2011). In

the distribution area of the species flowering has been recorded in August, October and

November.

Material examined. MYANMAR. Mandalay Region: Kyaukpadaung Township, 26 Oct 2009,

Saw Lwin, Pan KhetKhet & Sandar Maung MPO 020 (SING). Growing in a small population.

114

Card. Bull. Singapore 67(1) 2015

Notes. Habenaria reniformis is characterised by small greenish or brownish flowers

with sepals mostly less than 4.5 mm long. The lip shape is variable, ranging from entire

to 3-lobed (Banerji & Pradhan, 1984: 36; Chen et al., 2010: 193; Baretto et al, 2011:

260), and the spur can be absent or cylindrieal and up to 7.6 mm long (Seidenfaden,

1977: 133; Kurzweil, 2011: 133; Baretto et al., 2011: 260). The rather robust anther

and the long protruding stigmas are characteristic of the species (Seidenfaden, 1977).

In the collection from Myanmar, Lwin et al. MPO 020, the petals are basally

widened to more than double their width above. The petals and the lip are basally

united with the column. The lips of the two plants in this collection are mostly unlobed.

The only exception is one flower which has an asymmetric 3-lobed lip and appears to

be abnonually developed: one side lobe is linear-filiforai and 4 mm long, and the other

one semicircular and only 0.5 mm long.

Habenaria reniformis is very close to H. hnmidicola Rolfe and H. poilanei

Gagnep., both of which differ by having larger leaves, consistently deeply 3-lobed lips

and longer spurs. The Habenaria diphylla (Nimmo) Dalzell group is also very close.

The relationships among these species are not fully resolved and it is not yet clear if all

of these taxa can be mamtained as separate species. As already noted by Seidenfaden

(1977), further research is needed to reach a final position on their status.

5. Liparis distans C.B. Clarke, J. Linn. Soc., Bot. 25: 71 (1889); Seidenf , Dansk Bot.

Ark. 31(1): 76, fig. 50 (1976); N.Pearce & P.J.Cribb, FI. Bhutan 3(3): 212 (2002);

S.C.Chen et al. in Z.Y.Wu et al. (eds), FI. China 25: 225 (2009). - Leptorkis distans

(C.B.Clarke) Kuntze, Revis. Gen. PL 2: 671 (1891). - Stichorkis distans (C.B. Clarke)

Marg., Szlach. & Kulak, Acta Soc. Bot. Poloniae 77: 38 (2008). - TYPE: India,

Nagaland, Kohima, 21 Oct 1885, Clarke, C.B. 41071 (syntype K! [K000387820],

CAL); Kohima, Clarke, C.B. 41099 (syntype K, not found); Kohima, 21 Oct 1885,

Clarke, C.B. 41105 (syntype K! [K000387819]); Kohima, 21 Oct 1885, Clarke, C.B.

41574 (syntype K! [K0003 87821]). (Fig. 2D-E)

Liparis macrantha Hook.f, Hooker’s Icon. PI. 19: t. 1854 (1889). - TYPE: India,

Nagaland, Prain, D. 44 (lectotype K, designated by Seidenfaden (1976b)).

Liparis yunnanens is Rolfe, J. Linn. Soc., Bot. 36: 8 (1903). - TYPE: China, Yunnan,

Henry, A. 10485 (holotype K! [K000364665]).

Liparis oxyphylla Schltr., Repert. Spec. Nov. Regni Veg. Beih. 4: 63 (1919). - TYPE:

China, Yunnan, Henry, A. 12593 (lectotype K! [K0003 64666], designated here;

isolectotype E! [E00286229]).

Herb mostly 20-40 cm tall, pseudobulbous, glabrous. Pseudobulbs clustered,

subcylindric or narrowly ovoid, 2-6(-9.5) cm long, slightly compressed. Leaves

2, distichous, shortly and indistinctly petiolate, articulate, spreading, nan'owly

oblanceolate, acute or acuminate, 15-35 x 1-2.8 cm. Inflorescences arcuate or

New records of Myanmar orchids

115

pendulous, 15-39 cm long, lax, with up to 12 distant flowers; peduncle somewhat

compressed, narrowly winged, with 2-3 sterile bracts; floral bracts narrowly triangular,

6-12 mm long. Flowers medium-sized, olive-green to dull yellow; pedicel and ovary

14-22 mm long; sepals narrowly lanceolate, obtuse, mostly 10-16 x 3-5 mm, margins

often revolute; petals linear-filifoiin, obtuse, 10-16 x c. 2 mm; lip ovate, obovate or

elliptic, 10-14 x 10-11 mm, basal part a short claw, margin irregularly denticulate,

rounded or obtuse at the apex, with a 2-lobed callus at the base, unspurred; column

arcuate, 5-6 mm long, narrowly winged in the upper part. Capsule ellipsoid or

narrowly obovoid-oblong, 12-18 x 6-7 mm.

Distribution. Widespread from northeastern India and the eastern Himalayas to

Myanmar, Indochina and southwestern and southern China. Also in the Philippines.

Ecology and phenology. The specimen from Myanmar was growing as an epiph34e in

primary mountain forest. Elsewhere the species is lithophytic or epiphytic in various

forest types at elevations up to 2400 m (Chen et al., 2009). In Vietnam often found in

limestone regions (Averyanov, 2013). In the distribution area of the species flowering

has been recorded between September and February.

Material examined. MYANMAR. Kachin State: Border region of Putao and Nogmung

Townships, 7 Mar 2007 (fruiting at the time of collection; flowering in cultivation), Kurzweil,

H. & Saw Lwin KL 2460 (herbarium of the Myanmar Floriculturist Association).

Notes. Distinct with its large flowers in lax inflorescences and the wide lip with

denticulate margins.

6.LiparisresupinataRid\., J. Linn. Soc., Bot. 22: 290 (1886); Seidenf., DanskBot. Ark.

31(1): 88, fig. 60 (1976); N.Pearce & P.J.Cribb, FI. Bhutan 3(3): 209, pi. 7 [top right]

(2002); S.C.Chen et al. in Z.Y.Wu et al. (eds), FI. China 25: 228 (2009). - Leptorkis

resupinata (Ridl.) Kuntze, Revis, Gen. PI. 2: 671 (1891). - Platystyliparis resupinata

(Ridl.) Marg., Richardiana 7: 39 (2007). - TYPE: India, Darjeeling, 1844, Griffith, W.

s.n. (syntype K-LINDL! [K000873787]); India, Kliasia, sine collector 986 (syntype

K-LINDL! [K000873786]); India, Khasia, Griffith, W s.n. (syntype K-LlNDL!

[K000873788]); India, Sikkim, 5000 ft [1524 m], Hooker, J.D. 90 (syntype K-LINDL!

[K000873785]); India, Kliasia, Lobb, T. 122 (syntype K-LINDL! [K000873784]);

India, Kliasia, Hooker, J.D. & Thompson, T. 90 (syntype K-LINDL! [K000873783]).

(Fig. 3A-B)

Liparis ridleyi Hook.f, Hooker’s Icon. PI. 19: t. 1887 (1889). - Leptorkis ridleyi

(Hook.f) Kuntze, Revis. Gen. PI. 2: 671 (1891). - TYPE: India, Sikkim, 3000-5000

ft [91 4-1 524 ni], Hooker, J.D. 89 (syntype K-LINDL! [K000873782]); India, Sikkim,

Hooker, J.D. s.n. (syntype K-LINDL! [K000387829]); Griffith s collector, Kew Distr.

5081 (syntype K); India, Darjeeling, 5500 ft [1676 m], 11 Mar 1875, Clarke, C.B.

116

Card. Bull Singapore 67(1) 2015

Fig. 3. A-B. Liparis resupinata Ridl. From Lasi BawkNaw BW 32. C. Micropera thailandica

(Seidenf. & Smitinand) Garay. FromLwm et al. TNRO 61 (Photos: A, H. Kurzweil; B-C, Saw

Lwin)

27235 (syntype K-LINDL! [K000387827]; India, Darjeeling, 5000 ft, [1524 m], 2 Mar

1876, Clarke, C.B. 27088 (syntype K-LINDL! [K000387828]).

Herb 12-28 cm tall, with densely arranged pseudobulbous stems. Pseudobulbs 1.5-5

X 0.3-0. 6 cm, made up of a few nodes, with leaves in the upper half Leaves 2-A,

papery, subsessile, narrowly oblong to linear-lanceolate, acute to acuminate, 6-13

X 0.4-1. 2 cm, articulate. Inflorescences arching or pendent, lax, 10-50-flowered;

peduncle slightly zig-zagged, 5-8 cm long, with several sterile bracts 0.5-0. 9 cm

long; rachis 8-18 cm long; floral bracts lanceolate, acute, concave, 3-5 x 1-1.5 mm.

Flowers to 2 mm wide and 5 mm high when seen from the front; in the specimen

from Myanmar pale greenish orange-yellow with orange on the basal part of the lip

including its auricles, in other parts of the distribution area reported as pale green,

greenish yellow, golden brownish or creamy; pedicel and ovary 5-7 mm long; sepals

subequal, spreading, oblong or elliptic-oblong, obtuse or acute, 1 -veined, 3-4 x 1-1.8

mm; petals reflexed downwards, narrowly linear, obtuse, 2-3.5 x c. 0.3 mm; lip knee-

New records of Myanmar orchids

117

like, bent with the lower portion erect and parallel to the column and the distal portion

spreading forwards, broadly elliptic-oblong or broadly ovate-oblong, 2.5-3 mm long,

unspun*ed, base auriculate and with a bilobed callus, apex obtuse, margins incurved,

forming a pouch; column erect, 1-2.8 mm long, with two suborbicular wings, each

with a prominent pendulous filifonn appendage. Capsule obovoid-oblong, c. 5 x 2-3

mm.

Distribution. India and Himalayas to Indochina and southwestern China (Xizang and

Yunnan). Also recorded in the Gaoligongshan Mountains in western Yunnan Province

(Jin et ah, 2009), which is very close to where our specimen from Myanmar was

collected.

Ecology and phenology. The specimen from Myanmar was growing as an epiphyte

in degraded mixed broad-leaved forest with much bamboo and moss-covered trees

at an elevation of 2100 m. Elsewhere the species is found in evergreen broad-leaved

and coniferous forest from 1500 to 2500 m (Pearce & Cribb, 2002; Chen et ah, 2009;

Averyanov, 2013). In the distribution area of the species flowering has been recorded

from October to March.

Material examined. MYANMAR. Kachin State: Waingmaw Township, 10 Mar 2011, Stephen

Lasi Bawk Naw BW 32 (herbarium of the Myamnar Floriculturist Association). Locally

connnon.

Notes. One of the epiphytic Liparis species with few-leaved pseudobulbs. Well-

characterised by the shape of the small flowers that have a broadly oblong lip with

an obtuse and pouched apex and the column with characteristic pendulous filiform

appendages on the wings.

7. Micropera thailandica (Seidenf. & Smitinand) Garay, Bot. Mus. Leafl. 23: 187

(1972); Seidenf., Opera Bot. 95: 126, fig. 72 (1988); Vaddhanaphuti, Wild Orch.

Thailand, ed. 4, 186, inch colour photo (2005). - Camarotis thailandica Seidenf &

Smitinand, Orch. Thailand (Prelim. List) 712, fig. 529 (1965). - TYPE: Thailand,

Satun Province, Feb 1961, Seidenfaden, G.& Smitinand, T. GT 4118 (holotype C).

(Fig. IB, 3C)

Herb with monopodial growth. Stems to 30 cm long, glabrous, covered by the dark,

rugose bases of the leaves. Leaves about 1 0, distichous, spaced at intervals of about

2 cm, flat, conduplicate, oblong-linear, apex unequally bilobed with rounded lobes,

8-15 X 1-1,5 cm, fleshy. Inflorescences 17-23 cm long, several-flowered; peduncle

10-15 cm long, with 2-3 sheathing sterile bracts 0. 3-0.4 cm long; rachis 8-10 cm

long, unbranched; bracts broadly triangular, obtuse, 2-2.5 x c. 4 mm, fleshy. Flowers

yellow or light orange-yellow, lip base sometimes white, sepals sometimes tipped with

brown, column white or cream; median sepal elliptic-ovate, apex broadly rounded.

118

Card. Bull. Singapore 67(1) 2015

to 9 X 3.5 mm, concave, fleshy; lateral sepals similar but somewhat shorter and wider,

adnate to the column-foot; petals elliptic-lanceolate, acute, to 8 x 3.3 mm; lip 7 mm

long, deeply boat-shaped with a conical spur 4-5 mm long in its distal part, obscurely

3-lobed, very fleshy; midlobe triangular, c. 2 x 2-2.5 mm; side lobes inconspicuous,

broadly triangular, edges minutely serrate; callus a triangular tongue at the base of

the midlobe, split at the end, below this a rounded callus at the backwall of the spur

(terminology after Seidenfaden, 1988); spur with a longitudinal septum; column c. 6

mm long, venti'ally with a sudden horizontal edge with two short lateral horns below

the stigma, above which the column narrows to about one half of its width below,

higher up widening again under the anther and the rostellum, rostellum beak c. 2 mm

long, not or only slightly twisted.

Distribution. Myanmar, Thailand and Vietnam.

Ecology and phenology. The specimen from Myanmar was found in primary evergreen

forest at about 220 m, growing as an epiphyte on tall trees. In Thailand recorded at

elevations of between 700 and 1000 m (Seidenfaden, 1988). In the distribution area of

the species flowering has been recorded in February and March.

Material examined. MYANMAR. Taniiithayi Region: Yebyu Township, 712 ft [217 m], 25

Mar 2012, Saw Lwin, Pan Khet Khet & Zaw Oo Wai TNRO 61 (SING, SING [spirit], herbarium

of the Myanmar Floriculturist Association). Locally common.

Notes. This species is unmistakable in its lip and column structure. It is known from

several collections in Thailand and is also found in Vietnam; in addition it is here

newly reported for Myanmar. Seidenfaden & Smitinand (1965) compared it with the

Malaysian Camarotis adnata (Ridl.) Holttum (now included in Micropera fuscolutea

(Lindl.) Garay), which is superficially similar, but this species differs in details of its

lip and column (Seidenfaden & Wood, 1992).

8 . Oberonia obcordata Lindl., Fol. Orchid. 8: 7 (1859); Seidenf, Dansk Bot. Ark.

33(1): 30, fig. 19 (1978); N.Pearce & P.J.Cribb, FI. Bhutan 3(3): 230 (2002); S.C.Chen

et al. in Z.Y.Wu et al. (eds), FI. China 25: 240 (2009). - Malaxis obcordata (Lindl.)

Rchb.f. in W.G.Walpers, Ann. Bot. Syst. 6: 216 (1861). - Iridorkis obcordata

(Lindl.) Kuntze, Revis. Gen. PL 2: 669 (1891). - TYPE: India, Sikkim, Hooker, J.D.

& Thompson, T. 112 (syntype K-LINDL! [K000974237]); India, Darjeeling, 1844,

Griffith, W. s.n. (syntype K-LINDL! [K000974238]); India, Kliasia, Griffith, W. s.n.

(syntype K-LINDL! [K000974231]). (Fig. IC-D, 4A)

Oberonia treutleri Hook.f, Hooker’s Icon. PI. 18: t. 1786 (1888). - Iridorkis treutleri

(Hook.f.) Kuntze, Revis. Gen. PI. 2: 669 ( 1 89 1 ). - TYPE: India, Sikldm, 6000 ft [ 1 829

m], 1 Dec 1874, Treutler, W.J. 1151 (holotypeK! [K000387726]).

New records of Myanmar orchids

119

Fig. 4. A. Fruiting specimens of Oberonia obcordata Lindl. From Lasi Bawk Naw BW 8. B.

Fruiting specimen of Stereosandra javanica Blume in its natural habitat in southern Myanmar.

From Lwin et al. TNRO 5 (Photos: A, H. Kurzweil; B, Saw Lwin)

Herb to 9 cm tall, with leafy stems to 4 cm tall, glabrous. Leaves (3-)5-8, distichous,

equitant, bilaterally eompressed, fleshy, linear-oblong, slightly falcate, acute to

aeuminate, mostly 1.5-5 x 0. 5-0.9 cm, base not articulate. Inflorescences terminal,

densely or subdensely many-flowered, 4. 5-7. 5 cm long; peduncle connate to lower

part of uppermost leaf and therefore appearing to arise from its blade, with few to

many erect sterile braets about 0.5 cm long; rachis slender, 1 .5-6 em long; floral braets

verticillate, lanceolate or ovate-lanceolate, apex acuminate-caudate, 2-A mm long,

much longer than the flowers, margin slightly erose. Flowers 1-1.5 mm in diameter;

reddish brown or red; pedicel and ovary 0.5-1. 5 mm long; median sepal ovate, acute,

to 1.1 X 0.6 mm; lateral sepals broadly ovate, obtuse, to 1.1 x l mm; petals linear-

oblong, subaeute, to 0.9 x 0.5 mm; lip 3-lobed, not spurred, to 1.6 mm long; side

lobes divergent, ovate-oblong, obtuse, to 0.5 mm long; midlobe obcordate or reniform,

emarginate, to 1 x 0.8 mm, with a prominent basal callus; column to 0.5 mm long.

Capsule ovoid, to 5 x 4 mm.

Distribution. Himalayas, northeastern India, Myanmar and southwestern China to

Thailand.

Ecology and phenology. The specimens from Myanmar were found in forest at various

elevations (see below). Elsewhere either epiphytic or lithophytic, usually at elevations

120

Card. Bull. Singapore 67(1) 2015

up to 3000 m (Pearce & Cribb, 2002; Chen et al., 2009). In the distribution area of

the species flowering has been recorded between July and December and in March.

Flowering and fruiting plants can sometimes be found at the same time.

Material examined. MYANMAR. Kachin State: Putao Township, undisturbed evergreen

forest, 1 500 m, 2 1 Mar 2009, Kurzweil, H. & Saw Lwin KL 2664 (SING); Waingmaw Township,

degraded mixed forest, 2200-2300 m, 9 Mar 2011, Stephen Lasi Baw’k Naw BW 8 (RAF,

herbarium of the Myanmar Floriculturist Association). Locally common in both localities.

Notes. Very distinct due to the inflorescence which is connate to the uppermost leaf,

the long and whorled floral bracts, and the small flowers with their large lip side lobes

and the basal lip callus.

9. Stereosandra javanica Blume, Mus. Bot. 2: 176 (1856); Seidenf , Dansk Bot. Ark.

32(2): 173, fig. 107 (1978); Hedge & Rao, Indian J. Forest. 10(3): 196 (1987); Seidenf.

& J.J. Wood, Orchids Penins. Malaysia Singapore 143, fig. 60 (1992); N.Pearce &

P.J.Cribb, FI. Bhutan 3(3): 582 (2002); S.C.Chen et al. in Z.Y.Wu et al. (eds), FI. China

25: 207 (2009); H.A.Pedersen, FI. Thailand 12(2): 632, fig. 342 (2014). - TYPE:

Western Java, Bantam Province, Kuhl, H. & van Hasselt, J.C.A. s.n. (? holotype L!).

(Fig. lE-G, 4B)

Stereosandra pendula Ki'aenzL, Bot. Tidsskr. 24: 11 (1901).- TYPE: Thailand, South-

Eastern, Ko Chang, Schmidt, J. s.n. (not known to exist; see Seidenfaden, 1978: 173).

Herb terrestrial, holomycotrophic, 15-40 cm tall, glabrous. Corm ellipsoid or ovoid,

1. 5-4.4 X 0.6-2.4 cm. Stems leaf-less, slender to robust, 1^ mm in diameter; with 2-7

erect and sheathing sterile bracts 1.1-1. 8 cm long. Inflorescences laxly 3- to many-

flowered; rachis 3-19 cm long; floral bracts lanceolate, acuminate, 3.5-11.6 x 1. 2-4.3

mm, mostly shorter than the ovary. Flowers resupinate; white or cream with purple

markings and purple-tipped sepals, petals and lip; pedicel and ovary distinct from

each other, ovary ellipsoid, 3-8.2 mm long, pedicel 1.5-5 mm long; sepals lanceolate,

acute to acuminate, 8-10.1 x 1. 6-2.2 mm; petals similar, linear-lanceolate, acute to

acuminate, 7-9.2 x 2.2-2. 6 mm; lip ovate-lanceolate, obtuse to subacuminate, entire to

irregularly undulate-crenate, 5-8 x 2. 3-3. 5 mm, concave; with two ellipsoid wart-like

calli at the base; column incurved, 3^.2 mm long. Capsule ellipsoid to subspherical,

5-13 X 3-7 mm.

Distribution. Widespread from the eastern Himalayas through the whole of tropical

and subtropical Asia to New Guinea and the islands of the southwestern Pacific Ocean

(east to Samoa),

Ecology and phenology. The specimens from Myanmar were found in bamboo-rich

hill evergreen forest near a stream at an elevation of 1 80 m. Elsewhere the species is

New records of Myanmar orchids

121

found in primary evergreen forest from 100 to 1500 m (Seidenfaden & Wood, 1992;

Chen et aL, 2009; Pedersen, 2014). In the distribution area of the species flowering

occurs mostly between March and June but has also been recorded between December

and February. At the locality in southern Myanmar both flowering and fruiting plants

were seen at the same time.

Material examined. MYANMAR. Taninthayi Region: Yebyu Township, 23 Mar 2012, Saw

Lwin, Pan KhetKhet & Zaw Oo Wai TNRO 5 (SING, SING [spirit]). Locally common.

Notes. This is a new generic record for the country. Given its overall distribution

(tropical Asia from Thailand and Peninsular Malaysia eastwards to New Guinea;

Pridgeon et al., 2005 : 528) the occurrence of Stereosandra in Myanmar is not surprising.

In neighbouring Thailand Stereosandra javanica is also found in areas adjacent to the

border with Myanmar. Known localities in southern and southwestern Thailand are the

provinces of Ranong and Phetchaburi, both bordering Myanmar’s Taninthayi Region

where our specimen was collected. We suggest that Stereosandra javanica may be

more widespread in the southern part of Myanmar, but has probably been overlooked

because of its inconspicuous appearance.

ACKNOWLEDGEMENTS. Fieldwork on Mount Popa, in Kachin State, and in Taninthayi

Nature Reserve was made possible by the Myanmar Forest Department and we would like to

thank the officials concerned for providing the necessary pennits. We would like to acknowledge

Htoo Foundation for providing research funding for the orchid surveys on Mount Popa. Thanks

are also due to Stephen Lasi Bawk Naw, Brang Aung and local foresters for their assistance in

the field. We would like to thank Paul Onnerod for advice on the newly reported species, Jaap

J. Vermeulen for providing infonuation on BuJbophyllum capnophyton, and Jonathan Gregson

for checking herbarium material at BM. Thanks are also due to .loshua Yang for making the line

drawing, and the fu'st author would like to acknowledge the National Parks Board in Singapore

for finaneial support.

References

Averyanov, L. (2013). The orchids of Vietnam illustrated survey. Part 4. Turczaninowia 16(1):

5-163.

Baneiji, M.L. & Pradhan, P. (1984). The Orchids of Nepal Himalaya. Lieehtenstein, Vaduz: J.

Cramer, A.R. Gantner Verlag Kommanditgesellschaft.

Baretto, G., Cribb, P.J. & Gale, S. (2011). The Wild Orchids of Hong Kong. Malaysia, Kota

Kinabalu: Natural Histoiy Publications (Borneo) & China, Hong Kong: Kadoorie Farm

and Botanic Garden.

Chen, S.-C., Liu, Z.J., Zhu, G.H., Lang, K.-Y, Tsi, Z.-H., Luo, Y.B., Jin, X.H., Cribb, P.J.,

Wood, J.J., Gale, S.W., Ormerod, P., Veniieulen, J.J., Wood, H.P., Clayton, D. & Bell,

A. (2009). Orchidaceae. In: Wu, Z.Y., Raven, P.H. & Hong, D.Y. (eds) Flora of China,

vol. 25: 1-570. China, Beijing: Science Press & USA, St. Louis: Missouri Botanical

Garden Press.

122

Card. Bull. Singapore 67(1) 2015

Chen, S.-C., Liu, Z.J., Zhu, G.H., Lang, K.-Y., Tsi, Z.-H., Luo, Y.B., Jin, X.H., Cribb, RJ.,

Wood, JJ., Gale, S.W., Ormerod, R, Vermeulen, J.J., Wood, H.R, Clayton, D. & Bell,

A. (2010). Orchidaceae. In: Wu, Z.Y, Raven, RH. & Hong, D.Y (eds) Flora of China,

vol. 25, illustrations: 1-666. China, Beijing: Science Rress & USA, St. Louis: Missouri

Botanical Garden Rress.

Christenson, E. & Wood, H. (2003). The mystery of Limodorum aphyllum - Resolving lingering

taxonomic problems concerning two dendrobiums. Orchids (West Palm Beach) 72:

774-777.

Clements, M.A. (1989). Catalogue of Australian Orchidaceae. Austral. Orchid Res. 1: 1-160.

Jayaweera, D.M.A. (1981). Orchidaceae. In: Dassanayake, M.D. & Fosberg, F.R. (eds) A

revised Handbook to the Flora of Ceylon, vol. 2. India, New Delhi: Amerind Rublishing

Co.

Jin, X.-H., Zhao, X.-D. & Shi, X.-C. (2009). Native Orchids from Gaoligongshan Mountains,

China. China, Beijing: Science Rress.

Jin, X.-H., Dai, Z.-Q., Liu, Q.-Y. & Ju, X.-Y. (2010). Miscellaneous taxonomic notes on

Orchidaceae from China. Acta Bot. Yunnan. 32: 331-333.

Kurzweil, H. (2011). Habenaria. In: Santisuk, T. & Larsen, K. (eds) Flora of Thailand, vol.

12(1): 79-149. Thailand, Bangkok: The Forest Herbarium, Department of National

Parks, Wildlife and Riant Conservation.

Nyan Tun (2014). Wild Orchids of Myanmar. Myamnar, Yangon: Daw Tin Tin Aye (Green Leaf

Publishing House).

Ormerod, P. & Sathish Kumar, C. (2003). Orchidaceous additions to the flora of Burma

(Myanmar). Rheedea 13: 43-50.

Ormerod, P. & Sathish Kumar, C. (2008). Orchidaceous additions to the flora of Myanmar 2.

Rheedea 18: 75-80.

Pearce, N.R. & Cribb, RJ. (2002). Flora of Bhutan: Including a record of plants from Sikkim

and Darjeeling, vol. 3(3), The Orchids of Bhutan. UK: Royal Botanic Garden Edinburgh

& Bhutan: Royal Government of Bhutan.

Redersen, H.^. (2014). Stereosandra. In: Santisuk, T. & Balslev, H. (eds) Flora of Thailand,

vol. 12(2), pp. 631-633. Thailand, Bangkok: The Forest Herbarium, Department of

National Parks, Wildlife and Plant Conservation.

Pridgeon, A.M., Cribb, P.J., Chase, M.W. & Rasmussen, F.N. (2005). Genera Orchidacearum,

vol. 4 (Epidendroideae, part 1). UK: Oxford University Press.

Schuiteman, A. (2011). The strange case of Dendrobium aphyllum. Orchid Rev. 119: 104-110.

Seidenfaden, G. (1976a). Conhibutions to the orchid flora of Thailand VII. Bot. Tidsski: 71:

1-30.

Seidenfaden, G. (1976b). Orchid genera in Thailand IV. Liparis L.C.Rich. Dansk Bot. Ark.

31(1): 1-105.

Seidenfaden, G. (1977). Orchid genera in Thailand V. Orchidoideae. Dansk Bot. Ark. 31(3):

1-149.

Seidenfaden, G. (1978). Orchid genera in Thailand VI. Neottioideae Lindl. Dansk Bot. Ark.

32(2): 1-195.

Seidenfaden, G. (1988). Orchid genera in Thailand XIV. Fifty-nine vandoid genera. Opera Bot.

95: 1-398.

Seidenfaden, G. & Smitinand, T. (1961). The Orchids of Thailand - A preliminary list. Part III.

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Denmark, Fredensborg: Olsen & Olsen.

Gardens’ Bulletin Singapore 67(1): 123-127. 2015

doi: 10.3850/S2382581215000137

123

A new species of Zingiber (Zingiberaceae)

from Lao P.D.R.

M.R Newman

Royal Botanic Garden Edinburgh, 20A Inverleith Row,

Edinburgh EH3 5LR, Scotland

m.newman@rbge.ac.uk

ABSTRACT. Zingiber nitens M.F.Newman from Lao RD.R. is described and illustrated.

Keywords. lUCN conservation assessment, Lao RD.R., new species, Zingiber

Introduction

Rhizomes of a wild ginger collected in 2011 by Dr Vichith Lamxay of the National

University of Laos and his colleagues were donated to the Royal Botanic Garden

Edinburgh for cultivation. As the material they collected was without flowers but had

the remains of a temiinal inflorescence, the rhizomes were labelled Alpinia. When

the plant flowered in the living collection at Edinburgh in 2012, it was immediately

apparent that this was a new species of Zingiber.

Zingiber Mill, is among the larger genera of Zingiberaceae with 144 species

(Govaerts et ah, 2015) distributed from Sri Lanka and India to southern China, Japan

and SE Asia. Schumann (1904) classified the genus into four sections, two of which,

Zingiber section Dymczewiczia (Horan.) Benth. and Z. section Pleuranthesis Benth.,

contained species with terminal inflorescences. What Schumann did not observe was

that some species in Zingiber section Zingiber and Z. section Ciyptanthiiim Horan.,

such as Zingiber barbatum Wall., Z. gramineum Noronha ex Blume, and Z. junceum

Gagnep., could produce inflorescences both radically and tenuinally (Triboun et al.,

2014). Schumann (1904) placed Zingiber barbatum in Zingiber section Cryptanthium

Horan, and Z. gramineum in Zingiber section Zingiber (which he called Zingiber

section Lampugium Horan., nom. inval.); Gagnepain (1906) placed Z. junceum in

Zingiber section Zingiber (again as Zingiber section Lampugium).

Now, there are at least eight species of Zingiber known to produce terminal

inflorescences. They are found from the eastern Himalaya (Z. capitatum Roxb. and

Z. clarkei King ex Baker in India and Sikkim) to Papua New Guinea (Z. brevifolium

K.Schum.). Zingiber confine Miq. is from southern China, Z. pellitum and Z.

rufopilosum Gagnep. are found in Continental SE Asia, Z. gramineum is Irom Java,

and Z. marginatum Roxb. is a poorly known species of unknown origin,

Theerakulpisut et al. (2012) estimated the phylogeny of 23 species of Zingiber

using ITS sequences and concluded that Zingiber sections Zingiber, Dymczewiczia

and Pleuranthesis were individually weakly supported but together formed a well-

supported clade sister to a well-supported clade consisting of Z. section Cryptanthium.

124

Card. Bull. Singapore 67(1) 2015

The following description and illustration were made from a full-grown, living

plant and the lUCN assessment uses the criteria given in lUCN version 3.1 (2012).

Zingiber nitens M.F.Newman, sp. nov.

Belongs to Zingiber section Dymczewiczia (Horan.) Benth. because the inflorescences

are produced tenninally on the leafy shoots; similar to Zingiber capitatum Roxb. in

its slender habit with narrow leaves and tenninal inflorescences but differs from it by

its glabrous, glossy, dark green bracts wliich remain green even at fruiting (not green

bracts which turn red at fruiting, and are sparsely to densely villose at the margins). -

TYPE: Originally a living collection from Lao P.D.R., Bolildiamxai prov., Khamkeut

district. Ban Thongpe, Nakai-Nam Theun NPA, Lao- Vietnam Border Protected Area,

18°1L54" N 104°35'52.9"E, 573 m altitude, primary evergreen forest along river,

1 August 2011, Lamxay, K, Lanorsavanh, S., Souvannakoummai, K, & Somphone

VL2188, grown on as cultivated material at RBGE acc. no. 201 11 043 A, vouchered

and selected as type as Newman, M.F. 2647 (holotype L, inch spirit). (Fig 1, 2)

Clump-forming herb 0.65-1 m tall. Rhizome c. 1 cm in diameter. Leafy shoots

composed of c. 12 leaves, leaf sheaths dark brownish green, especially lower ones,

coarsely white villose, hairs pointing in all directions; pseudostem c. 1 cm in diameter;

ligule 3-5 mm long with truncate apex, translucent green when young, soon becoming

light brown, especially at margin, coarsely white pubescent; swollen petiole light

green. Leaf blade narrowly elliptic, 18-25 x 2-3 cm, glabrous and shiny adaxially,

sparsely white villose abaxially, especially on midrib, base rounded, apex very long

acute. Inflorescence a terminal, erect thyi'se, 9-13 x 1.3-2 cm, cylindrical to fusifonn,

composed of 9-1 5 bracts, wrapped round and obscuring rhachis. Bracts subtending to

3-4 flowers, somewhat obovate, c. 35 x 33 mm, glossy, dark green, glabrous, surface

with minute oil glands, apex acute, margin translucent. First flower of cincinnus

ebracteolate, subsequent flowers with a boat-shaped bracteole, open to base, c. 20

X 6.5 mm, translucent at base, green at apex, glabrous. Flower exserted from its

bract; 4 cm long, calyx 13-16 x c. 4 mm, tubular, inflated, translucent, glabrous,

split halfway down one side, apex with 2 short, blunt teeth; floral tube 30-3 1 mm

long, widenmg slightly towards apex, white at base, pale yellow at apex, lobes pale

yellow, glabrous, dorsal corolla lobe triangular, 19-21 x c. 6 mm, margins slightly

inrolled, lateral corolla lobes 18-20 x 5-5.5 mm; lateral staminodes triangular, 2.5-6

mm long, almost free from labellum, pale yellow, sometimes with a few red dots;

labellum elliptic, c. 18 x 1 1-12.5 mm, dark maroon with small yellow dots and yellow

patch in throat, margins deflexed, apex bifid for 3 mm. Stamen: filament 2.5-3 x 3

mm long, yellow; anther c. 1 1 x 4 mm, connective tissue yellow, thecae dehiscing by

longitudinal slits; anther crest 9-10 mm long, wrapped around stigma, curved down

into cleft of lip, dark maroon. Style white, glabrous, stigma white, round, scarcely

wider than style, ostiole with ring of straight cilia; epigynous glands 2, subulate, c. 3.6

mm long; ovary cylindrical, c. 3.5 x 2.5 mm, glabrous, incompletely trilocular at base,

the three placentas forming a very short axis bearing 10-14 ovules, unilocular above.

New Zingiber from Lao P.D.R.

125

Fig. 1. Zingibernitens M.F.Newman. A. Habit. B. Ligule. C. Inflorescence. D. Cincinnus, bract

removed. E. Bract. F. Bracteoles, showing size range. G. Calyx. H. Floral tube, corolla lobes,

labellum and lateral staminodes, abaxial and adaxial views. I. Stamen, style, stigma and dorsal

corolla lobe. J. Detail showing position of style in corolla tube. K. Upper style and stigma,

adaxial and lateral views. L. Ovary (cross-sectioned in apical part). Scale bars: A= 10 cm, B,

J— L = 5 mm, C— I = 2 cm. Drawn by Claire Banks from RBGE living accession 201 1 1043 A.

126

Card. Bull Singapore 67(1) 2015

Fig. 2. Zingiber nitens M.F.Newman, from RBGE living accession 20 11 1043 A. (Photo: Mark

Newman)

placentas petering out on inside walls not far from base. Infmctescence c. 10 cm long,

mature fruits unknown.

Phenology. The wild collection, Lamxay et al. VL2188, had finished flowering in

August but detailed observations on phenology will require further collections.

Distribution & ecology. Zingiber nitens is only known from the type locality, where it

grows in primary evergreen forest along a river at c. 570 m altitude.

Provisional lUCN conservation assessment. Data Deficient (DD). Zingiber nitens

is a perennial, terrestrial herb which may be expected to support a certain amount

of disturbance. The area in which it was found has a degree of legal protection and

there is no evidence that the species is being harvested or otherwise targeted. Until the

extent of the wild population is better known, it is impossible to give a more precise

assessment.

New Zingiber from Lao P.D.R.

127

Etymology. The epithet “nitens”, Latin for shining, refers to the shiny braets of the

inflorescence.

Notes. One of the vegetative characters of Zingiber, and many other Zingiberoideae,

is the presence of an abscission layer at the base of the pseudostem which allows the

plant to shed its pseudostems for the duration of the dormant season, whether that is

the dry season in the monsoon tropics, or the winter in more northerly areas. Zingiber

nitens possesses an abscission layer but, in cultivation in Edinburgh, it does not shed

all its pseudostems. Instead, the laminae become rather chlorotic looking in the winter

but persist. It is not known what happens in wild populations.

All parts of the living plant give off a foetid smell when bruised, and the flowers also

smell foetid.

ACKNOWLDEGEMENTS. Claire Banks is gratefully acknowledged for the line drawing

used in Fig. 1.

References

Gagnepain, F. (1906). Zingiberacees nouvelles de I’herbier du Museum. Bull. Soc. Bot. France

53: 132-150.

lUCN (2012). lUCN Red List Categories and Criteria: v. 3.1. 2nd ed. Switzerland, Gland &

UK, Cambridge: lUCN.

Schumann, K.M. (1904). Zingiberaceae. In: Engler, H.G.A. (ed) Das Pflanzenreich: regni

vegetabilis conspectus, IV, 46, Heft 20. Leipzig: W. Engelmann.

Theerakulpisut, P., Triboun, P., Mahakham, W., Maensri, D., Khampila, J. & Chantaranothai, P.

(2012). Phylogeny of the genus Zingiber (Zingiberaceae) based on nuclear ITS sequence

data. Kew Bull. 67(3) 389-395.

Triboun, P., Larsen, K. & Chantaranothai, P. (2014). Akey to the genus Zingiber (Zingiberaceae)

in Thailand with descriptions of 1 0 new taxa. Thai J. Bot. 6(1): 53-77.

Govaerts, R., Newman, M. & Lock, J.M. (2015). World Checklist of Zingiberaceae. Royal

Botanic Gardens, Kew. http://apps.kew.org/wcsp/ (accessed on 11 Mar. 2015).

Gardens’ Bulletin Singapore 67(1): 129-142. 2015

doi: 10.3850/S2382581215000149

129

Taxonomic studies on Zingiber (Zingiberaceae) in China I:

Zingiber kerrii and the synonymy of Z. menghaiense

and Z. stipitatum

L. J. Leong-Skoraickova^ & N.H. Xia^

^Key Laboratory of Plant Resources Conservation & Sustainable Utilization,

South China Botanical Garden, Chinese Academy of Sciences,

Guangzhou, 510650 People’s Republic of China

nhxia@scib.ac.cn

"University of Chinese Academy of Sciences,

Beijing, 100049 People’s Republic of China

bailin@scib.ac.cn

^Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, 259569 Singapore

j ana_skomicko va@nparks .gov. sg

ABSTRACT. A general introduction to Zingiber is provided, along with an outline of the

materials and methods used in this paper and which are to be used in our subsequent taxonomic

studies on Zingiber in China. Based on an investigation of the protologues, and on comparisons

of both the type specimens and living materials collected from the type localities and their

immediate vicinities. Zingiber stipitatum S.Q.Tong and Z. menghaiense S.Q.Tong are here

reduced to synonymy under Z. kerrii Craib. A previous lectotypification of Zingiber kerrii is

discussed and amended. The existence of mixed collections among the isotypes of Zingiber

stipitatum is highlighted and a proposal to exclude the vegetative parts belonging to Z.

neotruncatum from the type material is made. A description and a colour plate of Zingiber

kerrii based on a collection from China are provided.

Keywords. India, Myanmar, synonym, Thailand

Introduction

Zingiber Mill. (Zingiberaceae, Zingibereae) is an economically important genus, best

known for the ginger of commerce. Zingiber officwaJe (L.) Roscoe. The genus is

widely distributed in tropical to warm-temperate Asia (Wu & Larsen, 2000). According

to The Plant List (2013), 244 names have been published in this genus, corresponding

approximately to 100-150 species (Theilade, 1999; Wu & Larsen, 2000; Kishor &

Leong-Skomickova, 2013). Strongly supported as monophyletic (Kress et al., 2002),

this genus is easily recognised among Zingiberaceae either by the flower structure (the

horn-shaped anther crest embracing the upper part of the style) or by the vegetative

character (the existence of a swollen part of the petiole, widely known as a pulvinus).

The currently accepted infrageneric classification of Zingiber recognises four

sections, based on the nature and position of the inflorescence: (1) sect. Zingiber,

130

Card. Bull. Singapore 67(1) 2015

having a spike on an erect and, usually, long peduncle; (2) sect. Cryptanthium Horan.,

characterised by radical inflorescences composed of a spike appearing at ground level

with a, usually, short procumbent peduncle; (3) sect. Pleuranthesis Benth., with spikes

breaking though the leaf sheaths laterally; (4) sect. Dymczewiczia Benth., with tenuinal

inflorescences. A recent molecular study based on a single marker and limited material

indicates that Zingiber sect. Dymczewiczia and Zingiber sect. Pleuranthesis are not

well segregated from Zingiber sect. Zingiber (Theerakulpisut et ah, 2012). As more

studies involving broader sampling are certainly needed before a new classification

can be formally proposed, the traditional sectional treatment is utilised in our current

work.

The last comprehensive taxonomic revisions of the genus are more than a

century old (Horaninow, 1862; Schumann, 1904), covering only 23 and 55 species

respectively. Several regional studies have been made in S & SE Asia in the meantime,

e.g. Ridley (1909), Valeton (1918), Loesener (1930), Holttum(1950), Smith (1988a,b),

Theilade (1996), Theilade (1999), Sabu (2003), Sabu (2006) and Triboun (2006).

Poor or missing type material, difficulties in the preservation of important

floral characters on herbarium specimens through traditional drying methods, as

well as variability of certain morphological characters coupled with hybridisation

and polyploidy in some genera, make Zingiberaceae taxonomically one of the most

challenging plants groups (Larsen, 1980; Theilade, 1999; Skomickova & Sabu,

2005; Leong- Skomickova et al., 2010; Ardiyani et al., 2011). Our current poor

understanding, particularly of the larger ginger genera, is further exacerbated by the

fact that many previous workers limited their area of study to a single country, rather

than working monographically, leading to repeated descriptions of the same taxa. A

number of taxonomists have written that the ideal approach to the taxonomic study

of gingers is to work with living fertile material or material well-preserved in spirit

(Smith, 1988a; Theilade, 1999; Skomickova & Sabu, 2005; Leong-Skomickova et

al., 2010). This may involve revisiting and collecting plants from the type localities

in order to take precise notes aided by good photo-documentation, and to preserve at

least fertile bracts, bracteoles and flowers (or better still the entire inflorescence) in

spirit, including having several flowers preserved separately to avoid damage (Burtt

& Smith, 1976). A thorough review of the pertinent literature of the species across its

phyto-geograph ical range and good communication between taxonomists working on

gingers is also crucial.

The genus Zingiber in China

Fifty-three names have been associated with Zingiber taxa in China, of which 46 are

based on types from China. The most recent taxonomic treatment of Zingiber is that

of Wu & Larsen (2000) in the Flora of China. In this work, 42 species are recognised,

with 34 being endemic to China. The genus is mostly restricted to the subti'opical

zone of China, i.e. south of the Qinling-Huaihe Line (Zhang & Tang, 1991). While

the account by Wu & Larsen (2000) serves as a useful overview, the work was based

Zingiber kerrii and the synon3Tny of Z. menghaiense and Z. stipitatum

131

mostly on the study of imperfectly preserved herbarium material, of which most

are not accompanied by spirit material. This resulted in short and often incomplete

descriptions lacking basic rhizome and floral characters, and in doubtful synonymies.

About a quarter of Chinese Zingiber species are known only from the type collections

and their identities are poorly understood. There are also a number of presumably

well-known species to which a certain name has been applied for an extended period

of time, sometimes simultaneously to more than one species, and yet an understanding

of what these species really are is not straightforward. These issues can only be

um-avelled with detailed taxonomic work.

While a thorough revision of Zingiber in China is far from complete, accounts

for various regional floras in neighbouring countries are in progress. It is therefore

useful to disseminate our existing results which include clarifications of the identities

of various taxa, the establishment of new synonyms, and the descriptions of new taxa.

The present paper, clarifying the identities of two species from Yunnan, Zingiber

menghaiense S.Q.Tong and Z. stipitatum S.Q.Tong, is the first in the series. It,

therefore, includes a more detailed introduction to the genus as outlined above, as well

as an explanation of the materials and methods applied in our studies, the details of

which will not be repeated in subsequent papers.

Material and methods

Our work on a revision of Zingiber in China was initiated in 201 1 . The protologues of

all published names along with all other pertinent literature on the genus were collated

and reviewed. Searches were made in the relevant herbaria for the original material

or type material identified from the protologues. All Chinese specimens of Zingiber

from China were examined at CDBI, EMA, GXMG, GXMI, HGAS, HITBC (in older

literature often referred as YNTBI), IBK, IBSC, KUN, LBG, PE, SYS. Additional

material from China and neighbouring countries (particularly Thailand, Laos and

Vietnam) was mostly accessed as hi-resolution digital images from the following

herbaria: AAU, BK, BKF, BM, C, E, G, HAST, K, KFRI, L, P, SING, TAI, TAIF, US,

W, WU. Since 2012 the first author has conducted extensive fieldwork to re-collect

fertile material from type localities or their vicinities. Spirit collections, consisting

of fertile bracts, bracteoles and single flowers, were made in the field for further

study. Extensive photographic documentation was also made following the protocols

established by Leong-Skomickova et al. (e.g. 2014a, 2014b). Rhizomes were collected

and brought for planting to the greenhouse in South China Botanic Garden for further

observation. The terminology in general follows Beentje (2010) and the recent works

of Kishor & Leong-Skomickova (2013) and Leong-Skornickova et al. (2014a). While

some previous works (e.g. Theilade, 1999; Wu & Larsen, 2000) treated the labellum as

a tri-lobed stmcture composed of a mid- lobe and two side lobes (implymg the absence

of staminodes in the genus Zingiber), this approach has not been supported by any

moi*phological study and is not in accord with our current knowledge of ginger flower

stmcture. In Zingiberaceae, of the six stamens, only the median posterior stamen

132

Card. Bull. Singapore 67(1) 2015

of the inner whorl is fertile, while the remaining two are sterile and comiate into a

labellum. Of the outer whorl, one of the stamens is fully reduced, while the remaining

two appear as sterile staminodes (ranging in shape from large and petaloid to small

teeth-like stmctures), flanking the stamen or adnate to the labellum (e.g. Larsen et ah,

1998). Petaloid staminodes strongly adnate to labella occur in other Zingiberaceae

genera, e.g. Siliquamomum or Siphonochilus. We therefore agree with the conclusion

of various authors (e.g. Ridley, 1899; Holttum, 1950; Smith, 1988a; Larsen et al.,

1998) that the side lobes in Zingiber are staminodes, which in some species are free to

the base and in others are well developed and might be either fully or partially connate

to the labellum. In a few species they may be almost fully reduced or even missing.

The way we have measured the labellum and lateral staminodes is shown in Fig. 1.

The degree of comiation is also an important character which should be included in

descriptions.

The identities of Zingiber menghaiense and Z. stipitatum

During initial herbarium work it was noticed that the holotypes of Zingiber menghaiense

S.Q.Tong and Z. stipitatum S.Q.Tong appear to belong to the same taxon (Fig. 2 A-C).

At the same time it was also noticed that two of the tliree existing isotypes of Zingiber

stipitatum consisted of mixed collections (Fig. 2C). Furthermore, two specimens

originally identified as Zingiber stipitatum and collected from the vicinity of the type

locality of Z. menghaiense {Pei, S.J. 11344 in KUN^ were redetennined as Z. kerrii

Craib by Dr Pramote Triboun in 2002. This prompted us to investigate the complex of

these three names in greater detail.

Zingiber menghaiense (Tong, 1987) was described from two collections from

Xishuangbanna in south Yunnan, China, Tong, S.Q. & Li, A.M. 32860 from Menghai

Xian and Tong, S.Q. & Li, A.M. 32902, Irom Jinghong city, with the HITBC specimen

of the former collection being designated as the holotype (Fig. 2 A). When first

described, it was inexplicably compared to Zingiber zerumbet (L.) Roscoe ex Sm., a

very different species with incurved bracts, rather than to other more similar species

with appressed bracts.

In the same article, Tong (1987) described Zingiber stipitatum based on a

single collection, Tong, S.Q. & Liao, C.J. 24836 from Ruili Shi, Dehong Zhou, in

the southwest of Yunnan (holotype HITBC48857, Fig. 2B). In the protologue it was

compared to Zingiber menghaiense. Both species have a narrow, lanceolate, glabrous

lamina of similar size and shape, a long erect peduncle, an oblong or narrowly ovoid

spike, pale green bracts with a purple red margin, a whitish labellum, and lateral

staminodes with red patches on both sides of the bases. Zingiber stipitatum was said

to differ by the following characters: (1) the ligules 6 imn long, emarginate (versus 6

mm long, bilobed in Z. menghaiense)', (2) the labellum orbicular with emarginate apex

and petiolate [meaning it has an attenuate base or is clawed] (versus labellum narrowly

obovate, apex bifid in Z. menghaiensey, (3) the entire lateral staminodes which are

shortly acuminate at the apices (versus bifid at apices in Z menghaiense).

Zingiber kerrii and the synonymy of Z. menghaiense and Z. stipitatum

133

Fig. 1. Schematic outlines of selected types of labellum (pink) and lateral staminodes (blue)

oeeurring in the genus Zingiber. From left to right, lateral staminodes nearly free from labellum,

lateral staminodes eonnate to labellum by basal %, lateral staminodes absent, (a) length of

labellum, (b) width of labellum, (e) length of lateral staminodes, (d) width of lateral staminodes.

A comparison of the holotypes of the two species reveals that they are quite

similar to each other. Also, the holotype of Zingiber stipitatum does not match the

protologue as the ligules are c. 3 mm long and clearly 2-lobed. Though shorter, the

ligules of Zingiber stipitatum are also very similar to those of Z. menghaiense in their

nearly coriaceous texture, being slightly pubescent, with a prominent raised vein on

the base and the membranous margin. As pointed out by Triboun (2006), the length of

ligules in Zingiber varies according to the position of leaves on the pseudostem and

those in the middle part of the shoot are usually longer than the others. Three isotypes

of Zingiber stipitatum were located at HITBC and KUN herbaria (HITBC49032,

KUN0833208 and KUN0833209). Further examination of these sheets revealed

that while the isotype at HITBC is of the same taxon as the holotype, the plant parts

mounted on the other two isotypes at KUN (KUN0833208, KUN0833209 (Fig. 2C))

belong to two different species. The ligules on the shoots are emarginate and c. 6 mm

long, consistent with the description and the painting in the protologue, while they

conflict with the protologue and the holotype in the lower lamina surface, sheaths

and ligules being densely villous (they are glabrous in the protologue and holotype).

These two shoots are easy to recognise as Zingiber neotruncatum T.L. Wu, K.Larsen &

Turland, which also occurs in Ruili as mentioned in its protologue (Tong, 1987), while

the inflorescences are certainly of Z. stipitatum (the inflorescence of Z. neotruncatum

has shorter peduncles and is more or less bent and without a dark brown tinge on the

tips of the bracts). As confirmed by the first author in the held. Zingiber neotruncatum

is quite common at the type locality of Z. stipitatum. It appears that the description

of Zingiber stipitatum in the protologue is based on these two different species. The

character of the ligules being emarginate, the only vegetative character used by Tong

(1987) to distinguish Zingiber stipitatum from Z. menghaiense, is derived from the

leafy shoots of Z. neotruncatum present in the two isotypes. After their exclusion, all

the remaining original materials show no critical differences to the type specimens

of Zingiber menghaiense. Further observation of floral characters was made through

134

Card. Bull Singapore 67(1) 2015

Fig. 2. Zingiber kerrii Craib. A. Holotype of Z. menghaiense S.Q.Tong {Tong, S.Q. & Li, AM.

32860, HITBC Acc. No. 048849). B. Holotype of Z. stipitatum S.Q.Tong {Tong, S.Q. & Liao,

C.J. 24836, HITBC Acc. No. 049032). C. One of the two mixed isotypes of Z. stipitatum

S.Q.Tong; the inflorescence belongs to Z. stipitatum, the leafy shoot belongs to Z. neotruncatum

{Tong, S.Q. & C. J. Liao, C.J. 24836, KUN Acc. No. 0833209; barcode 1219333). D. Lectotype

of Zingiber kerrii Craib {Kerr, A.F.G. 1290; barcode K000255235); reproduced with the kind

permission of the Director and the Board of Trustees, Royal Botanic Gardens, Kew.

Zingiber kerrii and the synon3Tny of Z. menghaiense and Z. stipitatum

135

the field work of the first author in 2012 and 2013. More than ten individuals were

found at the type locality of Zingiber menghaiense, with the flowers examined having

entire lateral staminodes (as opposed to staminodes with bifid apices as stated in the

proto logue) but all other characters matched the protologue and the type specimens well.

According to our observation of several other Zingiber species (i.e. Z. longyangjiang

Z.Y.Zhu), the phenomenon of lateral staminodes being bifid to various degrees occurs

occasionally in populations with otherwise entire staminodes, and is therefore not to

be considered a reliable character to distinguish two species. The shape of the labellum

also varies within and between populations and we have observed varying degrees of

narrowing at the bases and notches at the apices which covers the shapes of labella as

depicted in the line drawings of Zingiber stipitatum and Z menghaiense published in

the protologues. At the type locality of Zingiber stipitatum, which is 360 km away (as

the crow flies) from the type locality of Z. menghaiense, only Z. neotruncatum was

found. However, plants matching the description of Zingiber stipitatum were found

within 20 km of the type locality and these are identical to those from the type locality

of Z. menghaiense. It is, therefore, concluded here that Zingiber stipitatum is the same

taxon as Z. menghaiense.

The identity of Zingiber kerrii Craib

Zingiber kerrii Craib was described in 1912 from a collection from Northern Thailand

(Chiang Mai Kerr 1290, Fig. 2D) (Craib, 1912). It was included in the revisions of

Zingiber in Thailand by Theilade (1999) and Triboun (2006) and both works included

photos of the inflorescence and flower. The species was also reported to occur in Shan

state in Myanmar (Kress et al., 2003) and in Manipur state in India (Thongam et al.,

2013). Compared to the Thai collections (Theilade, 1999; Triboun, 2006), the Indian

collection is slightly different by having the labellum and lateral staminodes creamy

white throughout. Our collection of Zingiber menghaiense, approximately 400 km

away from the type locality of Z. kerrii, shows more resemblance to Z. kerrii than the

Indian record, especially in the coloration of the labellum and lateral staminodes. Even

though the laminas are only c. 2.5 cm wide in the type material of Zingiber kerrii,

and described as 2.3 cm wide by Triboun (2006), they can reach up to 5 cm wide in

some other specimens in Thailand, similar to the measurements reported from India

(Thongam et al., 2013). Chinese collections consistently have laminas 3.5^ cm wide.

In conclusion, we agree with Triboun that Zingiber menghaiense (and also Z. stipitatum

as noted above) is the same taxon as Z. kerrii. Therefore, Zingiber menghaiense and Z.

stipitatum are reduced to synonymy under Z. kerrii. Colour plates, including a flower

dissection of Z. kerrii from Yunnan, are provided (Fig. 3, Fig. 4).

Theilade (1999), in her revision, indicated the presence of syntypes of Kerr

1290 at BK, BM, E and K and proposed the lectotypification of Zingiber kerrii with

a specimen deposited at K. However, the Kew herbarium has two specimens of Kerr

1290, both of which contain both a leafy shoot and an inflorescence, and as there

is no indication that these two specimens are part of a single preparation, a further

136

Card. Bull Singapore 67(1) 2015

Fig. 3. Zingiber kerrii Craib. A. Young shoots (inset: leaf sheath and ligule). B. Inflorescences

(left at anthesis, right in fruit). C. Old rhizome with root tubers and section of young rhizome

and root tubers. D. From left: Bract, bracteole, mature capsule (dehisced) and seeds enclosed

in arils. From Bai, L. 13080301 and Bai, L. 12091402, from the type locality of Zingiber

menghaiense. (Photos: L. Bai)

second step lectotypification is needed. In conformity with the ICN (Arts. 8.3, 9.17)

we designate here one of these two specimens, which also contains a flower dissection

(barcode K000255235), as the lectotype.

Zingiber kerrii and the synonymy of Z. menghaiense and Z. stipitatum

137

Fig. 4. Zingiber kerrii Craib. A. Flower (front view). B. Flower (semi-side view). C. Flower

dissection (from left to right: bract, bracteole, single flower in side view, dorsal corolla lobe,

two lateral corolla lobes, labellum with lateral staminodes partially basally connate, stamen

with upper part of style in the groove between two anther thecae. From the type locality of

Zingiber menghaiense, Bai, L. 13080301. (Photos: L. Bai)

Zingiber kerrii Craib, Bull. Misc. Inform. Kew. 10: 403 (1912); Loesener in Nat.

Pflanzenfam.lSa: 588 (1930). - TYPE; Thailand, Chiang Mai, Doi Sootep, in

evergreen jungle, 660 m., 24 July 1910, Kerr, A.F.G. 1290 (lectotype K! [K000255235],

138

Card. Bull. Singapore 67(1) 2015

designated here; isolectot 5 q)es BK n.v., BM! [BM000858181], E! [E00097850], K!

[K000255234], P! [P00450941]).

Zingiber rnenghaiense S.Q.Tong, Acta Phytotax. Sin. 25 (2): 145-146, pi. 1, f. 2.

(1987); S.Q.Tong in FI. Yunnan 8: 533 (1997); T.L.Wu & K.Larsen in FI. China. 24:

326 (2000), synon. nov. - TYPE: China, Yunnan Province, Xishuangbanna Daizu

Zizhizhou, Menghai Xian, Menghai Zhen, under the forest on the roadside, 1 200 m, 2

Jul 1982, Tong, S.Q. &. Li, AM. 32(5(56^(holotypeHITBC! [catalogue number 048849];

isotype KUN! [catalogue number 0833203]).

Zingiber stipitatum S.Q.Tong, Acta Phy to tax. Sin. 25 (2): 146-147, pi. 1, f. 3. (1987);

S.Q.Tong, in FI. Yunnan 8:532(1 997); T.L.Wu & K.Larsen in FI. China. 24: 326 (2000),

synon. nov. - TYPE: China, Yunnan Province, Dehong Daizu Jingpozu Zizhizhou,

Ruili Shi, Mengxiu Xiang, on the way from Mengxiu Cun to Daoba Zhai, 1200 m, 25

Jul 1983, Tong, S.Q. & Liao, C.J. 24536 (holotype HITBC! [catalogue number 48857];

isotypes FllTBC! [catalogue number 49032], KUN! ([catalogue number 0833208],

pro parte, excluding the leafy shoot), KUN! ([catalogue number 0833209], pro parte,

excluding the leafy shoot).

Feremiial rhizomatous herb 1.2-1.6(-2.5) m tall, glaucous throughout (although not

obviously so when old). Rhizomes fleshy, densely branched, 10-20 mm in diameter,

light brown externally, internally purple-pink when young, orange-yellow when

old; root tubers ovate to fusifonn, c. 2 x 1 cm, externally whitish brown, internally

pale grey. Leafy shoots to 15 in a tuft, with up to 44 leaves when flowering, less

than from the base leafless; leaf sheaths longitudinally striate, tubular at base; ligule

l-4(-6) mm long, bilobed, apices obtuse triangular, base green, turning blackish and

coriaceous when old, with obvious raised veins, upper part hyaline, membranaceous,

densely pubescent, glabrescent when old; petiole to 9 mm long, consisting of pulvinus

only, sparsely pubescent; lamina linear to naiTowly-ovate, 15-34 x 1.8-3(^.5)

cm, glaucous and glabrous on both side, base obtuse to attenuate, apex acuminate.

Inflorescences 1-3, radical; peduncles\2-20 cm long, erect, with pink scales on the

base; spikes fusiform or cylindrical, apices acute, 10-15 x 5-6 cm, fertile bracts each

subtending one flower, broadly obovate or broadly spathulate, slightly longer than

the floral tube, green with purple red apices, 3^.5 x 2-A cm, apices rounded, whole

spikes turning scarlet when fruiting; bracteoles naiTOwly ovate, apex acute, c. 30 x 6

mm, semi-translucent, sparsely pubescent externally, glabrous internally. Flowers 5-6

cm long; calyx tubular, membranaceous, c. 1 cm long, unilaterally split to 4 mm, apex

slightly dentate or nearly truncate, sparsely pubescent outside, glabrous inside, semi-

transhicent;y?ora/ tube creamy white, c. 3 cm long; dorsal corolla lobe narrowly ovate,

c. 18x9 mm, cream- white, concave, glabrous, apex mucronate; lateral corolla lobes

narrowly ovate, c. 18 x 7 mm, creamy, glabrous; labellum obovate with conspicuous

attenuate base, c. 26 x 12 mm, creamy, with purple red patch on the base, apex

emarginate or bifid, margin revolute; lateral staminodes narrowly obovate, nan'owly

oblong or narrowly triangular, c. 18 x 4 mm, basal 'A, or even less, connate to labellum.

Zingiber kerrii and the synon3Tny of Z. menghaiense and Z. stipitatum

139

creamy with two purple-red patches on the base, apex rounded to attenuate. Stamen

c. 17 mm \ong; filament reduced to less than 1 mm, anther c. 11 rmii long, comiective

tissue pale yellow, anther thecae 11 mm long, dehiscing throughout entire length,

pollen pale yellow, anther crest c. 6 mm long, pale yellow. Style filiform, white,

glabrous; stigma funnel form, white, ostiole ciliate. Ovary cylindrical, pale yellow,

trilocular, central placentation, with c. 20 ovules in each locule, c, 5 x 3 mm, densely

pubescent; epigynous glands two, pale yellow, c. 4 mm long, 0.3 mm in diameter, apex

blunt. Fruits obovoid to nearly sphaerical or bluntly trigonous capsules, c. 22 x ] g

mm, sparsely villous, greenish-cream with red tinge externally, red internally; seeds

obovoid, c. 5 X 4 mm, dark brown, glossy, with obvious white hilum on the base; aril

white, sac-like, apex with irregular margin, covering most of the seed.

Distribution. Zingiber kerrii is distributed in India, China, Myanmar and Thailand.

(Fig. 5). According to our photographic records, the species is also present in Laos

(Leong-Skomickova, unpublished).

Ecology. In China, Zingiber kerrii occurs in grassy areas on the forest edge, in bamboo

forest and evergreen broad-leaf forest at altitudes of 700-1300 m.

Provisional lUCN conservation assessment. Least Concern (LC). This species has

a large extent of occurrence (around 230,000 km2) and is Icnown from more than

10 localities of which some are protected. We, therefore, provisionally propose this

species to be Least Concern (LC) according to the latest lUCN criteria (2012).

Additional specimens examined. CHINA; Yunnan: Dehong Daizu & Jingpozu Zizhizhou,

Lianghe Xian, Mengyang Zhen, Dangliang Cun, 26 Sep 2012, Bai, L. 12092603 (IBSC);

ibidem, 17 Aug 2013, Bai, L. 13081701 (IBSC); Ruili Shi, Mengliu Xiang, Mangling Cun,

Moli, 21 Sep 2012, Bai, L. 12092103 (IBSC); Xishuangbanna Daizu Zizhizhou, Jinghong Shi,

Jinuo Xiang, Longpa cun, Zhou, S.S. 3212 (HITBC); Menghai Xian, Menghai Zhen, Manduan

Cun, 14 Sep 2012, Bai, L. 12091402 (IBSC); ibidem, 3 Aug 2013, Bai, L. 13080301 (IBSC);

Mengla Xian, on the road from Mengxing Xiang to Mania Zhen, 1 100 m, 21 Oct 2005, Zhou,

S.S. 3069 (HITBC); Mengla Xian, Ymm Xiang, 700 m, 9 Nov 1959, Pei, S.J. 59-11344 (two

sheets in KUN); Zhenyue Xian (Mengla Xian), 1200 m, Nov 1936, Wang, C. W. 80620 (PE).

THAILAND: Chiang Mai: Chom Thong District, Vachi Ratham Waterfall on Doi Intanon,

650-800 m, 15 Sep 1995, Larsen, K. et ai. 46498 (AAU); Doi Sutep, 1250 m, 30 Jul

1968, Sorenseri, T. et al. 3948 (AAU); Doi Sutep, 1120 m, in 1958, Sorensen,!, et al. 4539

(AAU); Inthanon National Park, 850 m, 22 Jul 1988, Phengklai, C. etal. 6757 (BKF); Mae Sa,

Rock Tower Mountain, 18 Sep 1995, Larsen, K. et al. 46636, (AAU); Pong Pho 12 km of Doi

Chieng Dao, 1200 m, 30 Jul 1968, K. Larsen et al. 2874 (E, AAU, BKF); Pliitsanulok: Chat

Trakan District, Phu Soi Dao National Park, 20 Jul 2006, Poulsen, A.D. & Suksathan, P. 2401

(E); Chaiyaphum: Thep Sathit District, 29 Aug 2001, Pooma, R. et al. 2940 (BKF).

Notes. Zingiber kerrii is similar to Z. laoticum Gagnep. in the entire plant being glaucous

externally (particularly prominent in young shoots, becoming inconspicuous when

older), bracts pale gi*een with pink-red margins and rhizomes and pseudostems purple-

red internally. Zingiber laoticum differs from Z. kerrii by the broader label lum which

140

Card. Bull Singapore 67(1) 2015

Fig. 5. Distribution of Zingiber kerrii Craib, based on herbarium material and literature records.

The type localities of the three names are indicated in red. (a) Zingiber kerrii Craib, (b) Z.

menghaiense S.Q.Tong, (c) Z. stipitatum S.Q.Tong.

is richly mottled in dark purple-brown, and longer (e. 15 mm), entire, membranous

ligules.

ACKNOWLEDGEMENTS. We thank Dr Yunfei Deng for critical reading and providing advice

for the paper, the curators of the herbaria CDBI, EMA, GXMG, GXMI, HGAS, HITBC, IBK,

IB SC, KUN, LBG, PE and SYS for their help with the specimens for our studies, the curators

of AAU, BK, BKF, BM, C, E, G, HAST, K, KFRI, L, P, SING, TAI, TAIF, US, W, WU for

the high-resolution images of specimens. The first author is grateful to the Singapore Botanic

Gardens for granting research fellowships to support the study at SING. The research of the first

and third authors was supported by the National Natural Science Foundation of China (Grant

no. 31170185), the research of the second author is supported by the National Parks Board

Singapore and the Grant Academy of the Czech Republic (GACR grant no. P506/14/13541S).

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Card. Bull. Singapore 67(1) 2015

Theilade, I. (1996). Revision of the genus Zingiber in Peninsular Malaysia. Gard. Bull.

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record for India from Manipur. Taiwania 58(4): 291-294.

Triboun, P. (2006). Biogeography and Biodiversity of the Genus Zingiber in Thailand. PhD

Thesis, Khon Kaen University, Khon Kaen (unpublished).

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Gardens’ Bulletin Singapore 67(1): 143-149. 2015

doi: 10.3850/S2382581215000150

143

Pennilabium labanyaeanum (Orchidaceae), a new species

from Meghalaya, Northeast India

C. Deori, N. Odyuo & A. A. Mao

Botanical Survey of India, Eastern Regional Centre,

Sliillong-793003, Meghalaya, India

drchayadeoribsi@gmail.com

nodyuo@yahoo.com

aamao2008@gmail.com

ABSTRACT. Anew orchid species, Pennilabium labanyaeanum C.Deori, N.Od5aio & A. A.Mao,

is described and illustrated from Meghalaya, India. A key to the species of Pennilabium in India

is provided.

Keywords. India, Meghalaya, new species, Orchidaceae, Pennilabium

Introduction

The genus Pennilabium JJ.Sm. (Orchidaceae) (penni- = feathered, -labium = lip) is a

genus of 1 6 species of epiphytes in the tribe Vandeae, subtribe Aeridinae. It is distributed

from northeastern India and southern China to the Philippines and Sulawesi, with

centres of diversity in Peninsular Malaysia and Borneo (Seidenfaden, 1988; O ’Byrne,

2000; O’Byme et ah, 2009; Schuiteman, 2013; Govaerts, 2014; Pridgeon et al., 2014).

The plants are small twig or branch epiphytes, the flowers are ephemeral and open in

succession, and the lip is spurred with small or large and often fimbriate or marginally

toothed side lobes, often with a fleshy mid-lobe. The column lacks a foot and contains

two pollinia on a long strap-like stipe; the rostellum is nonnally beaked (Pridgeon et

ah, 2014).

In India two Pennilabium species are known, both of which are confined to

the Northeast: namely, P. prohoscidcum A.S.Rao & J.Joseph from Meghalaya and

P. struthio Carr from Arunachal Pradesh. During a botanical tour to Laitkyrhong,

5 km from Smith, East Kliasi Hills District, Meghalaya, in July 2014, the first two

authors discovered a small Pennilabium with pure white flowers. After critical study

of specimens and the literature (Holttum, 1957; Rao & Joseph, 1969; Garay, 1972;

Comber, 1990; Rao, 1996), and comparison to the protologue and spirit material

of the type specimen of the morphologically most similar taxon, Pennilabium naja

P.O’Byme (O’Byme, 2000), this plant was found to be morphologically different from

all other species. Our plant from Meghalaya is therefore described as the new species

Pennilabium labanyaeanum C.Deori, N. Odyuo & A. A.Mao. Schuiteman (in Pridgeon

et al., 2014: 230) points out that ‘the generic delimitation of the genera closely allied

to Pteroceras, including Pennilabium, needs further study, and Pennilabium is only

tentatively accepted here’. Kocyan & Chase (in Pridgeon et al., 2014: 230) pointed

144

Card. Bull. Singapore 67(1) 2015

that only two Pennilabium species were sampled in their molecular study, and said ‘it

would appear that some changes to the taxonomy of this clade will be required once

greater sampling of the species has been accomplished.’

Pennilabium labanyaeanum C.Deori, N.Od 5 uio & A.A.Mao, sp.nov.

Pennilabium labanyaeanum is similar to P. naja in having reduced sidelobes but

differs in the flower being only half open; the sepals and petals pure white; the sepals

obtuse, keeled dorsally and 5-veined; the lip base forming a narrow hollow channel

with minute hairs; the lip epichile imlobed with erose margins, without a prominent

raised callus but with an area of yellow powdery substance widening towards the

lip apex; and the short white spur (see Table 1, Fig. 2). - TYPE; India, Meghalaya,

Laitkyrhong, 5 km from Smith, East Khasi Hills, 1753 m, 25°26'53.2"N 91°52'48.6"E,

23 July 2014, Deori, C. & Odyuo, N. 131601 (holotype CAL [as 131 601 A]; isotypes

ASSAM [as 131601 B-E]), (Fig.l, 2)

Small monopodial epiphytic herbs, 3-5 cm tall, glabrous. Roots 5-30 cm long, 2-4

mm in diameter, several from base of stem below leaves, white with green tips. Stems

1-2.5 X 0. 3-0.5 cm, very short, compressed with a few leaves close together. Leaves

1.5-6 X 0.5-1. 5 cm, distichous, elliptic or broadly lanceolate, more or less fleshy,

often twisted at the base, leathery, green, basally sheathing, unequally bilobed at apex,

apices subacute to obtuse. Inflorescence 2.5-6 cm long, unbranched, lateral, emerging

through sheath opposite leaf, erect or parallel to the leaves, producing 2-3 flowers

in succession; peduncle 2-3.5 cm long, slender, cylindrical, base covered with two

sheaths; sheathing sterile bracts below rachis two, c. 2 x 4.5 mm, somewhat rectangular

in shape, lemon green; rachis 5-10 mm long, complanate-cylindrical, ridged with one

to three distichously arranged flowers opening in succession; floral bract 2-2.5 x c.

1.5 mm, persistent, ovate-triangular, obtuse, lemon green. Flowers 12-14 mm long

from the tip of the dorsal sepal to the tip of the spur, 8-10 rmii broad, ephemeral,

half open, formed in succession, sepals and petals pure white with yellow markings

on the epichile of the lip. Pedicel with ovary 1-2.6 cm long, lemon green, 3 -ridged.

Sepals oblong with narrow bases, obtuse, 5-veined, on the dorsal side with keels which

apically project as small apicules; dorsal sepal 7-10 x 3-4 mm; lateral sepals 8-9.5 x

3.5^ mm. Petals 7.5-9 x 2.5-3 mm (base nan-ow), oblong with narrow base, obtuse

with 3 main veins, lateral veins branched. Lip spuiTed, 1 1.5-12.5 mm long (including

the spur), 4-5 rmu wide at the centre and 3 mm at the apex, unlobed, suborbicular or

ovate, lateral margins of lip erose; lip base with a naiTow hollow chamiel with minute

hairs within; epichile fleshy, solid, thick (1 mm in diameter), with an area of yellow

powdery substance widening towards the lip apex. Spur 5-7 mm long from junction

of lip base and the column, white, slender, tip rounded, 1.5 mm wide, aseptate and

ecallose; longitudmal section of the spur shows sparse minute hair-like structure or

raised edges at mouth towards the centre. Column 2-2.5 x c. 2 mm, without a foot,

rounded, without stelidia; stigmatic cavity very broadly ovate; rostellum not seen.

Anther 1-1.5 x 1.5-2 mm, operculum c. 0.2 x 0.5 mm, truncate, entire. Pollinia two.

Pennilabium labanyaeanum, a new orchid from India

145

Fig. 1. Pennilabium labanyaeanum C.Deori, N.Odyiio & A. A. Mao. A-C. Habit. D.

Inflorescence. E. Flower bud. F. Flower (side view). G. Peduncle sheath. H. Floral bract. I-J.

Flower (front & side views). K. Floral perigon with spurred lip, L. Column with pedicel and

ovary. M. Dorsal sepal (dorsal view). N. Lateral sepal (dorsal view). O. Lip (front view). P. Lip

front view (highlighting the narrow hollow channel with minute hairs at the base). Q. Spurred

lip (side view) with column, pedicel and ovaiy. R. Longitudinal section of spur. S-T. Anther

(front views). U. Pollinia with stipe. V. Stipe. W. Fruit. (Drawn by: C. Deori)

146

Card. Bull Singapore 67(1) 2015

Pennilabium labanyaeanum, a new orchid from India

147

c. 1 mm long (including stipe), 0.5-0. 7 mm in diameter, globose, entire; stipe 0.7-1

mm long, widening near the attaclunent of the pollinia, with a small disc. Fruits erect,

slender, 3-5.2 cm long which is more than twice the flower length, 3-ridged.

Distribution and habitat. Pennilabium labanyaeanum was found growing epiphytically

on moss-covered branches of trees. The species was flourishing in a steep subtropical

hill forest along a stream side at 1753 m altitude near Laitkyrhong, 5 km from Smith,

East Kliasi Hills District, Meghalaya, India. It was associated with other orchid species

in the genera Dendrobium Sw. and Eria Lindl. s.l., amongst others.

Phenology. Flowering and fruiting: July-August.

Etymology. The speciflc epithet is given in honour of the first author’s mother (L.

Labanya Deori, 1945-1993).

Provisional lUCN consen’ation assessment. Data Deficient (DD). Pennilabium

labanyaeanum is so far known only from a single locality in the East Khasi Hills District

in Meghalaya. According to O’Byme (2009), all species in the genus Pennilabium are

uncommon or rare and occur in small colonies that can be widely separated from each

other. If this is also the case for Pennilabium labanyaeanum, the new species might

be more widespread in Meghalaya than is currently known and further exploration is

necessaiy before a threat status can be given.

Notes. Pennilabium labanyaeanum is distinct in the genus due to its unlobed

suborbicular lip. The new species is similar to Pennilabium naja from Sulawesi

but differs as discussed above (see also Table 1, Fig. 2). As already pointed out by

Garay (1972) there are two types of lip stmcture in Pennilabium: 1 . Lateral lobes well

developed and midlobe reduced to a small fleshy lobe; 2. Lateral lobes much reduced,

possibly represented only by small auricle-like lobes, and midlobe very prominent.

Fig. 2. A-S. Pennilabium labanyaeanum C.Deori, N.Odyuo & A.A.Mao. A-A2. View of the

type locality. B-B2. Pennilabium labanyaeanum in the wild. C-C2. Habit. D. Inflorescence.

E. Floral bract. F. Peduncle sheath. G & Gl. Flower, front & side views. H. Fruit. 1. Dorsal

sepal, dorsal view. J. Lateral sepal, dorsal view. K. Flower bud, side view. L. Lip, front view.

LI. Lip, front view (highlighting the narrow hollow channel with minute hairs at the base).

M. Floral perigon with spurred lip. N. Spurred lip(side view) with column, pedicel and ovary.

O. Column, close view. Ol. Column with pedicel & ovary. P & PI. Anther, front views. Q.

Longitudinal section of spur. R. Pollinia with stipe. S. Stipe. Pennilabium naja P.O’ Byrne

(images taken from the spirit sample of the holotype O 'Byrne SUL107 at SING) T. Flower, side

view. U. Floral perigon with spurred lip. V. Lip, front view. W. Anther, front view. X. Pollinia

with stipe. (Photos: C. Deori & N. Odyuo)

148

Card. Bull. Singapore 67(1) 2015

Table 1. Comparison of Pennilabium naja and P. labanyaeamim.

Pennilabium naja

Pennilabium labanyaeanum

The flowers are fully open with yellow

sepals and petals, about 12 mm wide.

The flowers are not fully open with

pure white sepals and petals, 8-10 mm

wide.

Sepals spreading with recurved tips,

oblong obtuse, 3 -veined, not keeled.

Sepals not spreading, broadly oblong

obtuse, 5 -veined, keeled dorsally

at the apex.

Lip spurred, fleshy, tongue shaped,

white with lateral wings at the base

of the spur aperture; side lobes not

distinct; midlobe ovate in outline,

recurved in centre, epichile obtuse,

lamina concave with rolled under

margins, fleshy, iTigulose, raised into a

broad low ridge along median line from

base of callus to just before lip apex.

Lip spurred, fleshy, sub-orbicular, white

with no lateral wings; sidelobe absent;

midlobe not recurved at the centre,

epichile unlobed with erose margins,

without a prominent raised callus

but with an area of yellow powdery

substance widening towards the lip

apex.

Spur yellow, 9-10 mm long, slender.

Spur white, 5-7 mm long.

fleshy and solid. Most of the species in the genus fall into the first category. Two of

the three species found in India, Pennilabium proboscidcum and P. struthio, have well

developed side lobes, but the lip of the new species R labanyaeanum lacks side lobes

altogether, thereby separating it from all other species in the genus.

A key to the species of Pennilabium in India

la. Lip sidelobes present, large and fimbriate 2

lb. Lip sidelobes absent P. labanyaeanum

2a. Sepals and petals purple dotted, creamy white; lip sidelobes at least twice as

long as broad at the truncate apex, which has erose-fimbriate edges

P. proboscidcum

2b. Sepals and petals yellowish with dark red; lip sidelobes not twice as long as

broad near the truncate apex, which has a rim of dense, branching lacinia

P. struthio

Pennilabium labanyaeanum, a new orchid from India

149

ACKNOWLEDGEMENTS. The authors are thankful to Dr P. Singh, Director, Botanical Survey

of India for encouragement and facilities. The authors are thankful to Dr Andre Schuiteman,

Senior Researcher, Herbarium, Royal Botanic Gardens, Kew, for his comments in support

of the novelty of the species described here. We are also grateful to Guat Bee from OSSEA

(Orchid Society of Southeast Asia) for sending the protologue of Pennilabium naja and to

Bazilah Ibrahim of the Herbarium at Singapore Botanic Gardens, for sending spirit material of

the holotype of Pennilabium naja for study. We are also thankful to Rudan Kliakhongor and

Iboklang Kharhongor for assisting us during the field trip.

References

Comber, J.B. (1990). Orchids of Java. UK: Royal Botanic Gardens, Kew.

Garay, L. (1972). On the systematics of the monopodial orchids I. Bot. Mus. Leaf. 23(2):

149-212.

Govaerts, R., Bemet, R, Kratochvil, K., Gerlach, G., Can-, G., Alrich, R, Pridgeon, A.M., Pfahl,

J., Campacci, M.A., Holland Baptista, D., Tigges, H., Shaw, J., Cribb, R, George, A.,

Kreuz, K. & Wood, J.J, (2014). World Checklist of Orchidaceae. Royal Botanic Gardens

Kew. http://apps.kew.org/wcsp/home.do (25 Jul. 2014).

Holttum, R.E. (1957). Flora of Malaya (Orchidaceae), vol.l. Singapore: Government Printing

Office.

O’Byme, RO. (2000). Three new orchid species from Sulawesi. Malayan Orchid Rev. 34: 64.

O’Byme, RO., Phoon, S.N. & Ong. RT. (2009). Pennilabium armariv. A new species from

Peninsular Malaysia. Malesian Orchid J. 3: 113-116.

Pridgeon, A.M., Cribb, P.J., Chase, M.W. & Rasmussen, F.N. (2014). Genera Orchidacearum,

vol. 6 (Epidendroideae, part 4). UK: Oxford University Press.

Rao, A.S. & Joseph. J, (1969). Pennilabium proboscidcum A.S. Rao & Joseph — Anew orchid

species from K. & J. Hills, Assam with incidental first record of the genus from India.

Bull. Bot. Surv India 10: 231.

Rao, A.N. (1996). Five new records of orchids from Arunachal Pradesh, India. J. Econ. Taxon.

Bot. 20(3): 708.

Schuiteman, A. (2013). Avery odd thing: Pennilabium hewittii (Orchidaceae). KeM> Bull. 68:

679-681.

Seidenfaden, G. (1988). Orchid genera in Thailand XIV. Fifty-nine Vandoid Genera. Opera

Bot. 95: 330-332.

Gardens’ Bulletin Singapore 67(1): 151-157. 2015

doi: 10.3850/S2382581215000162

151

Tripogon bimucronatus (Poaceae: Chloridoideae:

Tripogoninae), a new species from India

K. Thoiba^, A.K. Pradeep^ & C.N. SuniP

^Department of Botany, University of Calicut,

Kerala, 673635, India

akpradeep 1 @rediflFmail.com

^Department of Botany, S.N.M. College, Maliankara,

Kerala, 683516, India

ABSTRACT. Tripogon bimucronatus Tlioiba & Sunil, a new species of grass from the

Nelliyampathy Hills, Palakkad District of Kerala, South India is described and illustrated. It

is moiphologically similar to Tripogon vellarianus Pradeep but differs by having adaxially

scabrid, linear-acuminate leaves; ligules a fringe of hairs to 2.5 nun long; spikelets with 6-8

closely packed florets; lower glume asynuuetrical; upper glume awned; lemma awn scabrid;

palea elliptic-lanceolate, apex bimucronate and notched at the centre.

Keywords. Tripogon, Poaceae, new species, Kerala, South India

Introduction

The genus Tripogon Roem. & Schult. belongs to subtribe Tripogoninae Stapf, tribe

Cynodonteae Dumort, subfamily Chloridoideae Kunth ex Beilschm. in the family

Poaceae Barnhart (Peterson et al., 20 1 0, 20 14; Soreng et al., 20 1 2). The genus comprises

44 species distributed in Africa, America, Australia, Temperate and Tropical Asia

(Clayton et ah, 2006 onwards). It is known to have 22 species in India, which includes

six recently described species (Murugesan & Balasubramaniam, 2008; Newmaster et

al., 2008; Kabeer et al., 2009; Chorghe et al., 2013; Thoiba & Pradeep, 2014).

During an exploration along the Western Ghats of South India, an interesting

specimen of Tripogon was collected from the hill slopes of the Nelliyampathy, Palakkad

District of Kerala, South India. Critical study revealed it to be quite distinct from the

remaining species of the genus and hence is described here as a new species, bringing

the total number of Tripogon species for India to 23. It is closely allied to Tripogon

vellarianus Pradeep (Pradeep & Sunil, 1999) but can easily be distinguished (Table 1).

Tripogon bimucronatus Thoiba & Sunil, sp. nov.

It is very similar to Tripogon vellarianus Pradeep but differs in having adaxially scabrid,

linear-acuminate, ribbed leaves, ligules being hairy at the ends of leaf sheaths, with

the hairs reaching up to 2.5 mm; spikelets with 6-8 closely packed flowers; the lower

glume being asymmetrical, and the apex of upper glume awned; lermna 1 -awned,

scabrid, straight or geniculate; palea elliptic-lanceolate, apex bimucronate and notched

152

Card. Bull. Singapore 67(1) 2015

at the centre. - TYPE: India, Kerala, Palakkad District, Nelliyampathy hill top, 1200

m elevation, 16 November 2010, Sunil, C.N. 4477 (holotype SING; isotypes BRIT,

CALI,K). (Fig, 1,2)

Tufted perennial herb. Culms 30-60 cm tall, erect; nodes glabrous. Leaf blades

30-60 cm long, 0.4-0.6 cm wide, linear, hispid towards base, apex acute-acuminate,

scabrid adaxially, margins involute. Sheaths closely involute to the culm, almost rigid,

sparsely hairy; ligtiles a fringe of hairs 2-2.5 mm long. Racemes 30^0 cm long,

spikelets arranged along both sides, 45-65 spikelets per raceme; peduncle glabrous,

terete, 10-15 cm long; rachis stout, minutely scabenilose, triangular. Spikelets linear-

lanceolate, 1 .3-1.8 cm long, 2-2.5 mm wide, 6-8-flowered; callus bearded. Rachilla not

persistent, 1-2 mm long, slightly scabrid. Lower glume 3-3.5 x 0.5-1 mm, lanceolate,

asymmetrical, notched on one-side below the middle, chartaceous, 1 -nerved, apex

acuminate. Upper glume 5-6 x 0.7-1 mm, elliptic-lanceolate, prominentely 3-nerved,

glabrous, apex with awns 0.5-1 mm long. Lemma 5-5.5 x 1 .5-2 mm (excluding awn),

3-nerved, l-awned, the awn arising between lateral lobes, median awn 5-5.5 mm long,

scabrid, straight or geniculate, lateral awns absent. Palea 4-5 x 1-2 rmu, hyaline,

narrowly elliptic, keeled and winged, keels minutely puberulous, apex bimucronate,

notched at the centre. Lodicules 2, c. 0.25 mm, quadrate, apex coarsely 3-toothed.

Stamens 3, anthers 1-1.5 mm long, oblong, filaments 0.5-0.75 mm long, slender,

glabrous. Ovary 0.25-0.5 mm long, obovate; style 2, slender, hyaline, 1 mm long;

stigma feathery, 1-1.5 mm long. Caryopsis 1. 1-1.3 x 0.4-0. 5 mm, naiTowly oblong-

lanceolate, obtusely trigonous, pale brown, obtuse at apex, longitudinally ribbed

abaxially.

Distrihution & Ecology^ Tripogon bimucronatus usually grows in the granitic grassy

slopes of Nelliyampathy Hills at an elevation of 1000-1200 m. It is found growing in

association with Arthraxon castratus (Griff.) V. Naray. ex Bor, Garnotia elata (Am.

ex Miq.) Janowsky., Cyanotis papilionacea (Bunn.f.) Schult. & Schult.f, Swertia

angustifolia Buch.-Ham. ex D. Don., Isachne bourneorum C.E.C.Fisch., Tripogon

wightii Hook.f., Tephrosia pulchella Hook.f, Cymbopogon commutatus (Steud.) Stapf,

Osbeckia spp., Parasopubia delphiniifolia (L.) H.-P.Hofm. & Eb, Fisch., Chrysopogon

nodulibarbis (Hochst. ex Steud.) Henrard, Sopubia trifida Buch.-Ham. ex D.Don,

Eulalia trispicata (Schult.) Henrard, and various moss species. It also occurs along

Meenuliyan Para Hills in Idukki District, Kerala at an elevation of 800 m.

Phenology^. It flowers from early September to October. Fmit set is low with the seeds

ripening by late October-November.

Etymology. The epithet ‘bimucronatus’ refers to the bimucronate palea apex.

Additional specimen examined. INDIA: Kerala: Idukki Dt., Meenuliyan Para, Vannappuram,

800 m elevation, 28 Nov 2014, Sunil 4910 (SNM College Herbarium, Kerala).

Tripogon bimucronatus, a new grass from India

153

Table 1. Comparison between Tripogon vellarianus Pradeep and Tripogon bimucronatus

Thoiba & Sunil.

Tripogon vellarianus

Tripogon bimucronatus

Leaf blades 8-15 mm wide, glabrous

on both surfaces

Leaf blades 5-8 mm wide, ribbed,

slightly scabrid adaxially

Leaf sheaths closely clasping, very

rigid; ligules membranous

Leaf sheath closely involute, almost

rigid; ligules ciliate, 2-2.5 mm long

Spike] ets distantly arranged,

8-10-flowered

Spikelets closely packed,

6-8-flowered

Lower glumes 4-5 x l .25 mm,

symmetrical, unlobed, acuminate at

the apex

Lower glumes 3-5 x 1mm,

asymmetrical, slightly 1-lobed on

one-side, acute-acuminate at the apex

Upper glumes 5-7.5 x 1.5 mm, acute

at apex, unawned

Upper glumes 5-6 x 1mm, a small

awn between the sinus at apex, awns

0.5-1 mm long

Lemmas 5-10 x 2 mm (excluding

awn), lateral awns absent or up to

5mm long, median awn 3 mm long,

glabrous, straight

Lemma 5-5.5 x 1 -2mm (excluding

awn),l-awned in between the lobes,

awn 5-5.5 mm long, scabrid, straight

or geniculate

Paleas c. 4 x 2 mm, elliptic, acute at

apex

Paleas 4-5 x 1-1.2 mm, elliptic-

lanceolate, bimucronate at apex with a

notch at the centre

Key to the species of Tripogon in India

(with lemma entire or 2-cleft at apex)

la. Lowest lemmas empty, dissimilar to other lemmas T. wardii

lb. Lowest lemmas bisexual, similar to all other lemmas 2

2a. Culms thickened below by the persistent leaf sheaths; leaf blades equitant, rigid,

apex pungent T. pungens

2b. Culms not thickened below by the persistent leaf sheaths; leaf blades not equitant,

not rigid or pungent 3

Tripogon bimucronatus, a new grass from India

155

3a. Central awns of the lemmas flexuous, capillary, several times as long as lemmas

T. capillatus

3b. Central awns of the lemmas straight or curved, not more than twice as long as the

lemmas 4

4a. Rachilla intemodes 1 mm long; apices of lemmas 1-awned 5

4b. Rachilla intemodes 2 mm long; apices of lemmas 3-awned T. copei

5a. Culms 15-25 cm high; leaf blades 5-20 cm long 6

5b. Culms 30-90 cm high; leaf blades 30-60 cm long 7

6a. Upper glumes 5.5-7 mm long; awns of the lemmas 6-8 mm long, straight or

curved T. wightii

6b. Upper glumes 8-9.5 mm long; awns of the lemmas 10.5-12 mm long, always

straight T. velliangiriensis

1 a. Leaf blades and culms glaucous, involute, filifomi; ligules very short and ciliate ....

T. jacquemontii

7b. Leaf blades and culms green, flat, sometimes rolled; ligules obsolete 8

8a. Leaf blades 3-6 mm wide; racemes 7-25 cm long; median awns shorter than

lemmas T. lishoae

8b. Leafblades 8-15 mm wide; racemes 30^0 cm long; median awn equal to or longer

than lemmas 9

9a. Ligules membranous, a fringe of hairs at the ends of leaf sheath; paleas subulate at

apex T. malabaricus

9b. Ligules ciliate, membranous or absent; paleas not subulate at apex 10

10a. Upper glumes 3-4 mm long; lemmas 2. 5-3. 5 m i n long 11

10b. Upper glumes 4-7.5 mm long; lemmas 4-10 mm long 12

lla. Ligules glabrous; lemma base glabrous, median awns as long as or shorter than the

lemmas; inflorescence 20-45 cm long; spikelets with 5-8 florets

T. sivarajanii

llb. Ligules ciliate at apex; lemma base bearded; median awns 1.5 times longer than

lemma; inflorescence 15-20 cm long; spikelets with 9-10 florets ... T. tirumalae

12a. Ligules a fringe of hairs; upper glumes awned at sinus; apex of paleas bimucronate

T. bimucronatus

12b. Ligules glabrous; upper glumes acuminate at apex; apex of paleas acute

T. vellarianus

156

Card. Bull. Singapore 67(1) 2015

Fig. 2. Distribution map of Tripogon vellarianus Pradeep (■) and Tripogon bimucronatus

Thoiba & Sunil (•) in Kerala

ACKNOWLEDGEMENTS. The authors are thankful to Dr K.H. Amitha Bachan, Western

Ghats Hombill Foundation (WGHF), Thrissur, Kerala for help during collection trips to the

Nelliyampathy hills, Palakkad District of Kerala, and to the anonymous reviewers for useful

suggestions that improved the text. The last author is thankful to the University Grants

Commission, New Delhi for financial assistance.

References

Chorghe, A., Rasingam, L., Prasanna, P.V. & Rao, M.S. (2013). Tripogon tirumalae (Poaceae),

a new species from the Seshachalam Hills of Andhra Pradesh, India. Phytotaxa 131:

17-22.

Clayton, W.D., Vorontsova, M.S., Harman, K.T. & Williamson, H. (2006 onwards). GrassBase-

the online world grass flora, http://www.kew.org/data/grass.db.html. (accessed on 10

Aug. 2013).

Tripogon bimucronatus, a new grass from India

157

Kabeer, K.A.A., Nair, V.J. & Muithy, G.V.S. (2009). Tripogon borii - a grass species new to

science from India. Bull. Bot. Siirw India 50: 1 15-118.

Murugesan, M. & Balasubramaniam, V. (2008). Tripogon velliangiriensis (Poaceae) - a new

species from Tamil Nadu, India. Indian J, Forest. 31: 109-111.

Newmaster, S.G., Balasubramaniam, V., Murugesan, M. & Ragupathy, S. (2008). Tripogon

cope (Poaceae: Chloridoideae), a new species supported by morphometric analysis and

a synopsis of Tripogon in India. Syst. Bot. 33: 695-701.

Peterson, P.M., Romaschenko, K., & Jhonson, G. (2010). A classification of the Chloridoideae

(Poaceae) based on multi-gene phylogenetic trees. Molec. Phylogenet. Evol. 55: 580-

598.

Peterson, P.M., Romaschenko, K. & Herrera Arrieta, Y. (2014). A molecular phylogeny and

classification of the Cteniinae, Farragininae, Gouiniinae, Gymnopogoninae, Perotidinae,

and Trichoneurinae (Poaceae: Chloridoideae: Cynodonteae). Taxon 63:275-286. http://

dx.doi.org/1 0. 1 2705/632.35

Pradeep, A.K. & Sunil, C.N. (1999). Two new species of Tripogon (Poaceae) from India. Sida

18: 809-814.

Soreng, R.J., Davidse, G., Peterson, P.M., Zuloaga, F.O., Judziewicz, E.J., Filgueiras, T.S.,

Mon'one, O., & Romaschenko, K. (2012). A world-wide phylogenetic classification of

Poaceae (Graminae). http://www.tropicos.org/projectwebportal.aspx?pagename=Classi

ficationNWG&projectid=10 (accessed on 10 Aug. 2013).

Thoiba, K. & Pradeep, A.K. (2014). Tripogon malabarica (Poaceae: Chloridoideae:

Tripogoninae), A new species from India, Kerala. J. Bot. Res. Inst. Texas 8: 525-529.

Gardens’ Bulletin Singapore 67(1): 159-229. 2015

doi: 10.3850/S2382581215000174

159

A revision of Epithema (Gesneriaceae)

K. Bransgrove^’^ & DJ. Middleton^

'Royal Botanic Garden Edinburgh, 20A Inverleith Row,

Edinburgh, EH3 SLR, Scotland, U.K.

^Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, Singapore 259569

david_niiddleton@nparks.gov.sg

^Current address: Australian Tropical Herbarium,

James Cook University, Cairns Campus, Building E2,

RO. Box 6811, Cairns, Queensland 4870, Australia

ABSTRACT. The genus Epithema Blume (Gesneriaceae) is revised. A key is given, all

species are described and several are illustrated. Twenty species are recognised and two

new combinations are made, Epithema philippinum (Hilliard & B.L.Burtt) Bransgrove and

Epithema pusillum (C.B. Clarke) Bransgrove. Provisional lUCN conservation assessments are

given for all species.

Keywords. Africa, Epithema, Gesneriaceae, Southeast Asia, taxonomic revision

Introduction

Epithema Blume is a genus of lithophytic herbs in central tropical Africa (Cameroon,

Central African Republic, Democratic Republic of Congo, Equatorial Guinea, Guinea,

Ivory Coast, Liberia, SieiTa Leone and Uganda), South Asia (Nepal, India, Sri Lanka),

Southeast Asia (southern China, Myanmar, Thailand, Vietnam, Cambodia, Laos,

Taiwan), Malesia (Philippines, Malaysia, Indonesia) and the Solomon Islands.

The genus was first described by Blume (1826) with a single species, Epithema

saxatile Blume. Aikinia R.Br. (in Wallich, 1832) was published shortly thereafter for a

different species {Aildnia brunonis) but it was soon synonymised under Epithema by

Decaisne (1834). Bentham (1835) and Spanoghe (1841) added further combinations or

new taxa to Epithema but Don (1838) continued the use of Aikinia, placing Epithema

in synonymy and moving Epithema carnosiim to Aikinia. Later authors, however,

correctly published treatments under the name Epithema. Major publications on the

genus mclude two complete treatments of Epithema (Candolle, 1845; Clarke, 1883).

Candolle (1845) included five species in his account and Clarke (1883) six species, of

which five were from Asia and one from Africa. The account by Clarke (1883) also

included eight new varieties and one new varietal combination. After 1883 there were

occasional additional regional accounts and the descriptions of new taxa (Henriques,

1892; Chevalier, 1912; Merrill, 1916; Ohwi, 1943; Burtt, 1958; Kiew, 1985; Ying,

1992; Li & Kao, 1998; Wang et al., 1998) and a set of brief but valid publications

of nine new taxa and combinations by Hilliard & Burtt (1997) in anticipation of a

revision of the genus which, sadly, never materialised.

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Card. Bull. Singapore 67(1) 2015

The classification of the Gesneriaceae into subfamilies, tribes etc. has been

rather unsettled (Weber et ah, 2013) with the result that Epithema has been placed in a

number of different subfamilies, tribes and series. Burtt & Wiehler (1995) and Weber

et al. (2013) both placed Epithema in the tribe Epithemateae (Meisn.) C.B.Clarke of

the subfamily Didymocarpoideae (called Cyrtandroideae by Burtt & Wiehler (1995)).

Weber et al. (2013) further placed it in its own subtribe, Epitheminae. Mayer et al.

(2003), in a molecular phylogenetic study of the group, found that the Epithematoid

Gesneriaceae are sister to the Didymocarpoid Gesneriaceae and that Epithema is

monophyletic and sister to a clade with a single species each of Loxonia Jack and

Stauranthera Benth. The study only included four species of Epithema {E. tenue, E.

saxatile, E, membranaceum and E. taiwanense [= E. ceylanicum]) and, therefore,

provides little information about the interrelatedness of the species within the genus.

In flower, Epithema is not easily confused with other genera although some

specimens have been previously mistaken for MonophyUaea R.Br. due to a superficial

resemblance to some species. The two genera can be separated by inflorescence

arrangement, sepal shape and aestivation, placentation, the number of locules in the

ovary, seed shape and hair type.

The species of Epithema are not readily distinguished from each other. Clarke

(1883) suggested that the then known five Asian species could possibly be recognised

as only one. Many are similar and there are few good characters to separate the species

as both vegetative and reproductive characters can be variable. In particular, the species

from mainland Asia are very similar and difficult to separate.

The gender of Epithema is neuter. Clarke (1883), however, described several

new varieties with feminine endings. These have been corrected to neuter in the

revision below.

Morphological Characters

Epithema is primarily lithophytic. In eastern and southeastern Asia it is found on

limestone and, to a lesser extent, on granite and sandstone. It appears to be restricted

to limestone in Borneo. In Africa, India and non-peninsular Myamnar it is found on

volcanic rocks and soils. It requires shaded conditions with high humidity and is

frequently found near or in cave entrances, on rocks on the forest floor, or on rocks

beside, or in, shallow streams and rivers. Information from specimen labels indicates

Epithema species will occasionally grow on fallen trees and is often collected from

road sides and road side embankments.

All species of Epithema are small herbs. Epithema, like other didymocarpoid

Gesneriaceae, is anisocotyledonous with the cotyledons withering early. The lowermost

true leaf, however, is solitary and looks rather like the macrocotyledon found in other

genera such as MonophyUaea (Weber, 2004).

The majority of species are caulescent but a few species have a very short

stem with leaves arising more or less at ground level, therefore appearing acaulescent

(Epithema dolichopodum, E. Jongipetiolatum, E, philippinum, E, rennellense, E.

strigosum and some specimens of E. benthamii, E. ceylanicum and E. tenue). One

Revision of Epithema

161

caulescent species, however, often exhibits a more sprawling rather than an upright

habit {E. sarawakense) and one is unifoliate in most of its distribution (E. horsfieldii).

The hairs on all plant parts are simple, unbranched, straight or hooked, have 3-5

cells and are from 0.02 mm to (rarely) 1.5 mm long. Hooked hairs are usually shorter

than straight hairs. The type of hairs (pubescent, strigose, setose, hispid or villous),

density (glabrous to dense), colour (hyaline to brown) and length are quite variable.

The combination of these characters is useM in some species but in others it is often

too variable to be of any taxonomic use. It is, therefore, included in the descriptions

only where it is taxonomically useful. Where ‘minute hooked pubescence’ is included

in descriptions it denotes hooked hairs of a maximum of 0.1 mm long.

The leaves of Epithema are thinly to thickly membranous and are usually

narrowly to broadly ovate, more rarely oblong to linear. The apex of the leaves is

rounded to acute, the margin entire to serrate, dentate or crenate (or variations thereof)

and the bases cordate to truncate to occasionally attenuate. The upper surface of

the leaves is green to dark or black-green and the lower surface is usually lighter in

colour than the upper surface and may dry whitish green to grey-green. Specimens of

Epithema saxatile and E. ceylanicum, and possibly others, show some green and silver

variegation on the upper surface of the leaves. The variegation has only been seen in a

small number of specimens from Thailand and Peninsular Malaysia. The indumentum

of the upper surface is variable in composition and the hairs are up to 1.5 mm long. The

indumentum of the lower surface is similar to that of the upper surface but the hairs

are usually sparser, shorter and finer. The indumentum of the leaves is occasionally

taxonomically useful and is only included in the descriptions in those cases.

The leaf venation is cladodromous with the secondary veins mostly strongly

ascending. There are often weaker intersecondaries and the tertiary venation is

mostly fairly obscure but, when visible, laxly alternate percurrent. The number and

arrangement of secondary veins is variable, between 4 and 1 5 and alternate to opposite

(or both witliin one leaf), and not taxonomically important.

Leaf arrangement is variable. Caulescent species can have opposite or solitary

leaves that in turn may be petiolate or sessile. A combination of all leaf an*angements

may be found on one plant or within a population. Most acaulescent species have

unpaired leaves. When present, solitary upper leaves are nearly always petiolate and

opposite one or more peduncles.

Weber (2004) described the infiorescence of Epithema as “reduced to a single

bract/inflorescence unif ’ which is derived from a “more elaborate thyrse, with several

bract/inflorescence units in alternate position”. The appearance is of a small, multi-

flowered, scorpioid cyme. The flowers are an*anged into rows, two to thi*ee across.

The indumentum of the peduncle is glabrous to hispid with hairs to 1 .25 mm long.

The flowers are collectively subtended by one bract which is green and frequently

cucullate, partially or entirely enclosing the inflorescence, but can be very small. The

uppermost margm of the bract is usually dentate but can be entire to dentate. If the

margin of the bract is lobed, the margin of the lobes may be entire, dentate or crenate.

The indumentum of the lower surface of the bract is of a short, hooked pubescence

and/or strigose to setose, straight and/or hooked hairs to 1 mm long. The indumentum

162

Card. Bull. Singapore 67(1) 2015

of the upper surface is usually of one hair type only, towards the margin and to 0.6

mm long.

The flowers mature sequentially from the base of the inflorescence towards the

apex. The apex is always curved over to more or less horizontal and is often covered

by the bract which subtends the entire inflorescence. The cells at the base of the

inflorescence continue to grow so that the inflorescence changes position as flowers

mature and the open flower is presented between the horizontal and around 45*^. The

number and size of the flowers in the inflorescence and the size and shape of the

bract varies between and sometimes within species but the size of the bract is a useful

character for the recognition of many species.

The pedicel is short, mostly up to only 5 mm long, and can elongate considerably

at maturity but only rarely to as long as 7.5 mm. The calyx consists of flve sepals that

are fused from the base of the calyx to half or two-thirds the length of the sepals. It

is actinomorphic, persistent and expands as it matures, providing a receptacle for the

operculate capsule that sits in the centre of the calyx. The young calyx is generally

oblong to obovate and becomes more campanulate at maturity. The lobes are linear to

triangular, often becoming broader and acuminate in maturity. The shape of the calyx

does not vary significantly between species.

The indumentum of the pedicel and calyx is similar to that on the bract and in

many species is the same on both structures. Hairs on the pedicel are to 1 mm long, to

1.5 mm long on the outside of the calyx and to 0.5 mm long on the inside of the calyx.

If present, the hairs on the inside of the calyx are usually strigose and found only on the

lobes of the calyx. The combination of type and densities of the hair on the outside of

the calyx is much more variable than on the inside of the calyx. Despite the variability

it can be a useftil character for species differentiation (e.g. E. tenerum and E. steemsii).

The corolla consists of a tube and five free corolla lobes in two lips. The tube

accounts for approximately two thirds to three quarters of the length of the corolla.

The smaller upper lip is of two lobes and the lower of three lobes. The upper lip is

sometimes much smaller than the lower. The colour of the corolla is variable. Specimen

labels indicate that the flowers may be white or pink, but usually the tube is white or

of unstated colour and the lobes are blue to violet. The flowers of many species have

irregular darker (purple) marks at the base of the lobes or in the upper throat.

In most species there are villous hairs on the inside of the eorolla tube and there

may be a minute pubescenee on the outside of the lobes. The villous hairs are often in a

dense band around the perimeter of the corolla tube and are usually placed somewhere

between half way and two-thirds of the way up the tube and below the stamens. The

tube is rarely glabrous or pubescent externally with short hairs inside. The presence

and type of hairs inside the corolla is a useful character for separating some species

of Epithema, while the hairs on the outside of the corolla lobes are not. Hairs inside

the corolla can be difficult to see in rehydrated material. The hairs on the outside of

the corolla can be difficult to observe and are not necessarily persistent. They are not

included in the species deseriptions.

The androecium consists of two fertile stamens and two staminodes. The

staminodes are a filament with an arrow- or serpent’s-head-shaped end. They are often

Revision of Epithema

163

slightly shorter than the filaments of the fertile stamens. The stamens and staminodes

are glabrous. The third, medial, staminode present in many Gesneriaceae is not seen

in Epithema. The anthers are apically coherent, the thecae are divaricate and apically

confluent. The stamens and staminodes are inserted on the top of a more or less square

flap of androecial tissue which is in turn inserted on the wall of the corolla. The flap

of tissue is up to 2.5 mm wide and high. The stamens are inserted on either end of the

flap and the staminodes to the inside of each of them. The upper edge of the flap of

tissue may be rounded leaving the staminodes at a slightly higher plane than the fertile

stamens, up to 0.5 mm higher. The head of the staminode is frequently adjacent to, but

not joined to, the back of the anther or the very top of the stamen filament. The style

grows through the space between the flap and the two coherent anthers. The androecial

tissue is normally inserted about half way to two-thirds of the way up the wall of

the corolla tube, but can be placed higher in the tube. In'espective of the position on

the corolla tube, the placement of the stamens in relation to the staminodes does not

change.

The nectary is comprised of one, two or three discrete lobes. In many species

there are two lobes, one on each side of the ovary. Sometimes the nectary lobe(s)

completely enclose the ovary, while in others the nectary is apparently absent. The

composition of the nectary is variable within and between species and is not a good

taxonomic character.

The ovary is superior, spherical to cylindrical and has one locule. The shape is

variable and is not taxonomically useful. The placentation is parietal with two ‘stalked’

placentae, one on each side of the ovary. The placentophores are fused in their lower

half to the ovaiy wall while the upper half supporting the placenta is free. This allows

the formation of a dehiscent operculum in fruit. The ovules are produced from, and

connected to, the placenta at the head of the placentophore by individual funicles.

The upper portion of the ovary is usually pubescent and the lower portion is glabrous.

Like the remainder of the plant, the indumentum consists of straight and/or hooked

hairs. The hairs are not always entirely persistent, but density, length and shape of the

hairs on the ovary are taxonomically useful. Unfortunately, the hairs are short (usually

0. 1-0.3 mm long) and not easily observed and, therefore, are of limited use for field

identification. The stigma is bilobed and the surface is covered in minute protrusions

or papillae of c. 0. 1 mm long. The style is slender and is inserted into the ovary in

the centre of the upper margin of the ovary. The style may be glabrous to densely

pubescent, the hairs being from 0.04-0.3 mm long and placed at the base of the style.

Rarely are they up to mid-way on the style.

The fi*uit of Epithema is a circumscissile or operculate capsule which is oblong,

obovate or sub-spherical in shape. The indumentum on the operculum is the same as

on the ovary although it may be sparser. After fertilisation the style withers and breaks

off leaving a small portion 0.3-2. 3 mm long. This remnant is persistent and seen on

the mature operculum. As the fruit matures, movement of the inflorescence brings the

fruit into a vertical position, the calyx lobes grow and open widely outwards, leaving

a basin-like structure in which sits the mature fruit. At maturity, the operculum is

lost, leaving the seeds open to the elements. The hair on the operculum is a useful

164

Card. Bull. Singapore 67(1) 2015

taxonomic character, while fmit shape, size and length of the style remnant are not.

A large number of seeds is produced per capsule. The seeds are narrowly to

broadly ovoid, rarely sub-cylindrical to slightly sigmoid in shape, with acute and/or

constricted ends. They are not appendaged and they separate entirely from the funicle

by which they comiect to the placenta. They vary from light to dark brown and the

ends are darker than the main body of the seed. Colour can vai*y between capsules of

the same inflorescence and is certainly associated with stages of maturity, becoming

darker with age. Some variation in colour might also be an artefact of drying.

The surface of the seed coat of all species of Epithema is arranged into more or

less parallel lines that, when viewed under magnification, are darker than the remainder

of the seed coat. The lines are straight to spiralled in the direction of the long axis of

the seed and are between 0.02 and 0.05 mm apart. These longitudinal lines may fork

and cross-walls are often present between the longitudinal lines. The density of the

cross-walls and the forking does vai 7 , giving some seeds a sub-reticulate look. Older

descriptions place importance on the angle of the longitudinal lines, but angle of the

pattern varies considerably even between seeds from one capsule or different capsules

of one inflorescence. Scamiing electron microscope (SEM) photographs of the seed

of a small number of species of Epithema indicate that the surface of the seed is not

taxonomically useful (Michael Moeller and Frieda Christie, pers. comm.). Further

examination of the seed by SEM was not conducted in this revision. In only a few

species is the length of the seed useful.

Splash cup dispersal is the primary means of seed dispersal in Epithema. The

splash cups are fornied from the persistent calyx and seeds are dispersed directly from

the capsule, usually by raindrops. Splash dispersal is usually only effective over short

distances but it is also possible that seed may be transported longer distances on the fur

or the feet of animals, or in streams.

Materials and Methods

Approximately 1400 specimens from the following herbaria have been examined: A,

AAU, ABD, BK, BKF, BM, BO, C, CAL, CALI, CMU, COl, E, G, GH, K, KEP, KLU,

KYO, L, MSC, MO, NY, P, PDA, PNH, PSU, QBG, S, SAR, SING, TI, UPM, US and

WAG (Thiers, continuously updated). These specimens were collected between the

1820s and 2014 from Cameroon, Central African Republic, Democratic Republic of

Congo, Equatorial Guinea, Guinea, Ivory Coast, Liberia, Sierra Leone and Uganda, Sri

Lanka, India, Nepal, China, Taiwan, Myanmar, Thailand, Laos, Cambodia, Vietnam,

Malaysia, Singapore, Philippines, Indonesia, Timor Leste and the Solomon Islands.

All specimens cited have been seen unless othemise indicated. All measurements for

vegetative characters, peduncles, bracts, pedicels and calyces are from dried specimen

material. Measurements of other reproductive structures are taken from rehydrated

material. Observations on herbarium material have been augmented by observations

in the field (in Malaysia and Thailand) by both authors.

There are relatively few collections from areas such as eastern Thailand, Laos,

Revision of Epithema

165

Cambodia, Vietnam, Sulawesi, Maluku, The Sunda Islands, Timor and Timor Leste,

New Guinea and the many small islands of south-eastern Asia. In some of these areas

Epithema species may genuinely be absent or scarce due to the lack of ideal habitats

(e.g. absence of limestone) but collecting densities are known to be low in several of

these areas (Johns, 1995; Middleton, 2003; Newman et ah, 2007). Peninsular Malaysia

and other areas of Thailand are relatively well collected.

Provisional conservation assessments are made using the methodology proposed

by lUCN (2001).

The species are presented in alphabetical order as little data is available on

relationships between the species. The dots on the maps represent the distributions of

the specimens cited in this revision.

The flowering and fruiting times given are based on the data from herbarium

specimens only and do not preclude flowering and fruiting at other times of year.

Constraints

Flowers in an Epithema inflorescence mature sequentially within the inflorescence with

only one or few flowers fully expanded at any one time. In addition, the corolla is not

persistent. It is, therefore, common for each specimen or sheet of specimens to contain

no corolla material. This means that a thorough evaluation of infraspecific variability

in corolla, stigma, style and stamen characters was limited by the availability of

flowering material. When only immature flower buds were present they were dissected

to gather character states for corolla and ovary pubescence. It is acknowledged that

character states could change during the maturation process but we still found these

bud characters useful.

For a number of species there are very few available specimens. This limits our

understanding of the variability and distribution of the species.

Plant description, expedition date and/or geogi'aphy data was limited or absent

from many labels of the older specimens. For species with only small numbers of older

specimens, distributions below country level are unattainable and transient character

states such as corolla colour remain unknown.

Epithema Blume

Bijdr. 737 (1826); C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1); 176 (1883);

C.B.Clarke in Hook.f., FI. Brit Ind. 4; 369 (1884); King & Gamble, J. Asiat. Soc,

Bengal, Pt. 2, Nat Hist. 74(2): 783 (1909); Ridt, FI. Malay Penins. 2: 539 (1923);

Gamble, FI. Madras 2: 991 (1924); Pellegr., FI. Indo-Chine 4: 558 (1930); Kanjilal et

at. Ft Assam 3; 399 (1939); BameU, Ft Siam. 3(3): 205 (1962); Backer & Bakh.f ,

FI. Java 2: 527 (1965); W.L.Theob. & Grupe in Dassan. & Fosberg, Revis. Handb.

FI. Ceylon 3: 102 (1981); Pham-Hoang Ho, Cayco Vietnam ed. 3, 3(1): 24 (1993);

W.T.Wang et at, FI. China 18: 400 (1998); B.L.Burtt, Thai For. Bull. (Bot) 29: 93

166

Card. Bull. Singapore 67(1) 2015

(2001); Hilliard in Grierson & D.G.Long, FL Bhutan 2(3): 1328 (2001); A. Weber in

Kubitzki, Fam. Gen. Vase. PI. 7: 129 (2004). - TYPE: Epithema saxatile Blume.

Aikinia R.Br. in Wall., PI. Asiat. Rar. 3: 65 (1832). - TYPE: Aikinia bnmonis Wall. (=

Epithema brunonis (Wall.) Decne.).

Carpocalymna Zipp., Alg. Konst- Lett.-Bode 1: 297 (1829), nom. nud.

Herb, caulescent or acaulescent, occasionally only one leaf developing. Leaves', lowest

leaf petiolate and solitary, upper leaves petiolate or sessile, opposite or alternate; blades

thinly to thickly membranous, usually ovate to cordate, more rarely elliptic to orbicular,

if asymmetrical then the wider side up to 1 .75 times as wide as the nan'ower side, apex

rounded to acute, base mostly cordate or sub-auriculate to obtuse, inserted evenly or

unevenly on petiole, margin entire to crenate, serrate or (bi-)dentate, venation pinnate,

upper surface pale to black-green, sometimes variegated, lower surface light to mid-

or olive-green or purplish. Inflorescences l-15(-many) per plant; peduncles usually

originating from the leaf axils, occasionally from the petiole and/or the midrib of the

blade; singular bract subtending each inflorescence, cucullate and enclosing the entire

inflorescence to reduced, margin entire to dentate. Calyx cylindrical to campanulate,

consisting of a tube and 5 lobes, with an embedded gland towards the apex of each

lobe. Corolla tube usually white, lobes pale pink to blue or purple, connnonly with

darker markings on either lip; tube cylindrical to narrowly fluted, occasionally slightly

constricted at the apex; lobes with margins entire to fimbriate. Stamens and staminodes

inserted on top of flap of androecial tissue which is inserted in corolla tube; fertile

stamens 2; anthers coherent at thecae tips or along entire thecae; staminodes 2, third

staminode absent. Nectary apparently absent or one- to three-lobed, margin entire to

undulate. Ovary cylindrical to spherical, glabrous to densely pubescent, unilocular,

placentation parietal; style short; stigma bi-lobed, papillate, glabrous. Fruit (sub-)

cylindrical to (sub-)spherical; operculum circumscissile or irregularly dehiscing at

maturity, indumentum as ovary; surrounded by persistent calyx. Seed usually naiTowly

ovoid to broadly ovoid, ends acute and/or constricted, light to dark brown with darker

ends, pattern straight to spiralled, walls of pattern usually splitting and merging and

with cross- walls, occasionally walls of pattern appear thickened and rigid.

Twenty species from central tropical Africa, India, Sri Lanka, Nepal, southern China

and through Southeast Asia and Malesia to the Solomon Islands.

Key to Epithema species

la. First leaf arising more or less at ground level or from a very short stem 2

lb. First leaf arising above a definite stem 10

2a. Indumentum on ovary/operculum straight or primarily straight 3

2b. Indumentum on ovary/operculum hooked or primarily hooked, operculum

rarely glabrous 8

Revision of Epithema

167

3a. Ovary/operculum indumentum mostly villous with hairs to 1.5 mm long;

corolla > 9 mm long; Africa 20. E. tenue p.p.

3b. Ovary/operculum indumentum generally not villous, < 1 mm long (usually

much less); corolla variable; Asia 4

4a. Hairs on ovary/operculum both straight and hooked; Solomon Islands

14. E. rennellense p.p.

4b. Hairs on ovary/ operculum straight only; not Solomon Islands 5

5a. Leaf margin entire or only minutely toothed, more rarely with larger and

irregular teeth; Eastern Indonesia 1. E. longipetiolatum p.p.

5b. Leaf margin distinctly toothed; widespread in Malesia 6

6a. Ovary/operculum hairs to 0.6(-l) mm long; upper leaf surface with only

strigose to villous hairs with at least some hairs > 1 mm long; Philippines and

eastern Malesia 1. benthamii p.p.

6b. Ovary/operculum hairs to 0.3 mm long; upper leaf surface sometimes

with minute hair covering as well as longer hairs but all hairs < 1 mm long;

Philippines and Sumatra 7

7a. Leaves strongly crenate or with large, wide teeth with rounded apices; bracts >

10 mm long; Sumatra 18. E. strigosum

7b. Leaves dentate/serrate, teeth narrow with pointed apices; bracts <10 mm long;

Philippines 12. E. philippinum

8a. Hairs on ovary/operculum hooked or hooked and straight; corolla 10.4-15 mm

long; Solomon Islands 15. rennellense p.p.

8b. Hairs on ovary/operculum only hooked or operculum rarely glabrous; corolla

5-8.5 mm long; not Solomon Islands 9

9a. Inflorescences 1-5 per plant, usually large and not in fascicles; peduncles

(2-)9-27 cm long; bracts 10-25 x 6-14 mm; operculum rarely glabrous;

Sabah 4. E. dolichopodum

9b. Inflorescences 1 to many, small, often different lengths, in fascicles; peduncles

up to 10(-17) cm long; bracts 3-16(-20) x 3-10 mm; operculum with hooked

hairs; continental Asia, Taiwan, Philippines

3. E. ceylanicum (rare acaulescent forai)

10a. Ovary/operculum glabrous 11

10b. Ovary/operculum pubescent or at least with few hairs 14

lla. Plants with only 1 leaf, occasionally with 2 leaves, these never opposite;

inflorescences arising from leaf blade midrib or junction of petiole and midrib

5. E. horsfieldii p.p.

llb. Plants usually with more than 2 leaves, leaves alternate or opposite;

inflorescences in the leaf axils or opposite a solitary leaf 12

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Card. Bull. Singapore 67(1) 2015

12a. Upper leaves sessile 9. E. madulidii p.p.

12b. Upper leaves petiolate 13

13a. Leaves all or primarily alternate; 1-5 peduncles arising in sequence from

base of petiole; bracts large (10-32 x 6-12 mm) and usually enclosing

inflorescence; plants with strigose or setose hairs, inner calyx glabrous or with

sparse straight strigose hairs; Malesia 15. E. sarawakense p.p.

13b. Leaves opposite; peduncles arising from leaf axils; bracts small (4-8 x 4-9

mm) and not enclosing inflorescence, plants glabrous to sparsely pubescent,

inner calyx glabrous; southern India E. pusillum

14a. Plants with only 1 leaf, occasionally with 2 leaves, these never opposite;

inflorescences arising from leaf blade midrib or junction of petiole and midrib,

ovary hair hooked 5. E. horsfieldii p.p.

14b. Plants usually with more than 2 leaves, upper leaves usually opposite;

inflorescences Irom base of petioles or in the axils of upper petiolate or sessile

leaves, rarely at base of midrib in the rare case leaves are alternate, 1-5

peduncles arising in sequence from base of petiole 15

15a. Upper leaves sessile 16

15b. Upper leaves petiolate 19

16a. Indumentum on ovary/operculum stiuight or primarily straight; Africa

20. E. tenue p.p.

16b. Indumentum on ovary/operculum hooked or primarily hooked; not Africa

17a. Corolla glabrous inside; Philippines 9. E. madulidii p.p.

17b. Corolla pubescent inside 18

18a. Bract enclosing inflorescence, 13-35 x 8-12 mm 16. saxatile p.p.

18b. Bract not enclosing inflorescence, 4-16 x 4-9 mm 3. E. ceylanicum p.p.

19a. Indumentum on ovary/operculum straight or primarily straight 20

19b. Indumentum on ovary/operculum hooked or primarily hooked 24

20a. Indumentum on ovary to 1.5 mm long; Africa 2Q. E. tenue p.p.

20b. Indumentum on ovary < I mm long; Asia 21

21a. Bracts cucullate, (2-)5-23 x ( 3 -) 4 -ii mm; Philippines, eastern Malesia ... 22

21b. Bracts not cucullate, 1-13 x 2-9 mm; continental Asia and Peninsular

Malaysia 23

Revision of Epithema

169

22a. Leaves mostly not opposite, only occasionally so and then often of markedly

different sizes; leaf margin entire or only minutely toothed, more rarely with

larger and irregular teeth 1. E. longipetiolatum p.p.

22b. Upper leaves usually opposite, only rarely upper leaves solitary; leaf margin

distinctly and usually regularly toothed E. benthamii p.p.

23a. Bract small, rarely sub-cucullate or trullate, not enclosing inflorescence, 1-7

X 2-7 mm, margin often deeply lobed with 3 lobes; plants often with dense

white hairs on most plant parts; corolla 4.2-5 .4 mm long; Peninsular Malaysia

11.^. pandbracteatum p.p.

23b. Bract small or sub-cucullate, enclosing a small portion of the inflorescence,

1-13 X 3-9 mm, margin entire to dentate; plants sub-glabrous to pubescent;

corolla 5. 5-7. 5 mm long; Myanmar, China, India, Laos, Nepal, Thailand

2. E. carnosum

24a. Corolla glabrous inside 6. E. involucratum

24b. Corolla with hair (villous) inside 25

25a. Leaves all or primarily alternate; 1-5 peduncles arising in sequence from

base of petiole; bracts large (10-32 x 6-12 mm) and usually enclosing

inflorescence; Malesia 15. E. sarawakense p.p.

25b. Leaves primarily opposite; number and position of peduncles variable; bracts

small to large and enclosing inflorescence or not; Continental Asia and Malesia

26a. Corolla >10 mm long 8. E. longitubum

26b. Corolla <10 mm long 27

27a. Peduncles arising along petiole, often in fascicles, often many and of very

different lengths 3. E. ceylanicum p.p.

27b. Peduncles primarily arising from axils of leaves, usually singly or few and then

of not remarkably different length 28

28a. Bract only rarely sub-cucullate or trullate but not enclosing the inforescence,

1-11 X 2-12 mm 29

28b. Bract sub-cucullate to cucullate enclosing a portion or all of the inflorescence,

(3-)6-23 X 3-12 mm 31

29a. Lower to mid-style pubescent; calyx without hispid white hairs; delicate

growth forni 10. E. mej7tbranaceum

29b. Style glabrous or with only a few hairs; calyx indumentum variable but with

hispid white hairs; often with a robust growth form 30

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Card. Bull. Singapore 67(1) 2015

30a. Plants with dense white hairs on most plant parts; Peninsular Malaysia

ll. E. pai^ibracteatum p.p.

30b. Plants not densely pubescent; not in Peninsular Malaysia

3. ceylanicum (rare form with few inflorescences and smaller bracts)

31a. Bract usually enclosing only a portion of the inflorescence 32

31b. Bract usually enclosing most or all of inflorescence; corolla 6. 5-9. 5 mm long

32a. Plant 5-14(-20) cm; leaves often broadly lanceolate to narrowly ovate; corolla

5.2-6 mm long; Sulawesi 19. E. tenerum

32b. Plant 4-40 cm; leaves usually cordate, ovate or elliptic; corolla 5-8.5 mm

long; not in Sulawesi 3. E. ceylanicum p.p.

33a. Calyx with strigose hairs only outside, hairs 0.5-0. 8 mm long; Aceh

17. E. steenisii

33b. Calyx often glabrous, if hairs present these < 0.5 mm long; Myanmar,

Thailand, Malaysia, Sumatra, Java, Borneo, Sulawesi \6. E. saxatile p.p.

l.Epithema benthamii C.B. Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 1 80 (1 883).

- TYPE: Philippines, Albay, 1841, Cuming, H. 1265 (lectotype K [K000438696],

designated here; isolectotype BM, K [K000438697]). (Fig. 1, 2)

Epithema brunonis var. scabridum C.B.Clarke in A.DC. & C.DC., Monogr. Phan.

5(1): 178 (1883). - TYPE: Indonesia, Sulawesi, Gorontalo, Riedel s.n. (holotype K).

Epithema calcicola Ohwi, Bot. Mag., Tokyo 57: 125 (1943). - TYPE: Indonesia,

West Papua, Waren, 60 miles south of Manokwari, 26 March 1940, Kanehira, R. &

Hatusima, S. 13143 (lectotype FU, image seen, designated here; isolectotype BO).

Herb 5-30 cm high, caulescent, occasionally appearing acaulescent with leaves at

near ground level, sparsely to densely strigose and/or setose with hairs to 0.8 mm

long, the lower bract, pedicel and the outside of the calyx also minutely pubescent with

hairs to 0.2 mm long; stem 1-4 mm wide with 2-3 nodes, intemodes 0.5-6. 5 cm long.

Leaves strongly membranous, upper leaves petiolate and opposite, infrequently with a

solitary upper leaf; petiole of lowest leaf 2-1 3.5 cm long, upper petioles 0.2-6 cm long;

blade of lowest leaf 4.3-14.5 x 2.2-7. 5 cm; blades of upper leaves 2-12 x 1-6 cm,

all leaves linear, narrowly ovate, or almost oblong, occasionally ovate or rhomboid,

usually symmetrical, apex acute to rounded, base sub-cordate, rounded to truncate,

occasionally sub-auriculate, usually inserted evenly on the petiole, margin usually

in-egularly serrate or biserrate; upper surface mid-green to black-green, sparsely to

densely weakly strigose to villous, without minute hairs interspersed, straight hairs to

1 .5 mm long, hooked hairs 0.5 mm long; lower surface generally lighter in colour than

Revision of Epithema

171

Fig. 1. Epithema benthamii C.B. Clarke. A. Habit. B. Leaf. C. Infloreseenee. D. Calyx. E.

Flower opened out. F, Stigma lateral view. G. Fruit showing seeds, placenta and operculum. H.

Seeds. Scale bars: A = 6 cm, B = 3 cm, C = 5 mm, D = 3 mm, E = 4 mm, F = 1 rmu, G = 2 mm,

H = 0.5 mm. Drawn by Claire Banks from Wenzel 437 (A, C-H) and Coode 5313 (B).

172

Card. Bull. Singapore 67(1) 2015

Fig. 2. Distribution of Epithema benthamii C.B. Clarke (•).

upper surface. Inflorescences l^(-6) per plant; peduncles 2.5-16 cm long, arising

from leaf axils or, rarely, from the base of the uppermost petioles; bracts sub-cucullate

or cucullate, enclosing a proportion of the inflorescence, occasionally reduced, (2-)7-

23 X (3-) 5-1 1 mm, margin usually dentate with large teeth but entire if reduced, lower

surface with long strigose hairs 0.6-1 mm long, upper surface glabrous to pubescent

at upper margin with hairs to 0.5 mm long; pedicels 1 .5^ mm long. Calyx 2.4-6 mm

long; tube 1.7-3 x 1.5-2. 5 mm; lobes 1.7-3 x 0.8-1. 5 mm; glabrous inside, rarely

with few straight hairs to 0. 1 mm long in lobes. Corolla often white, but also blue to

purple, 5-10 nmi long; tube (4-)5-6(-8) x 1.3-2. 5 mm; lobes 1-2 mm long. Stamens

1.4-1. 5 mm long; filaments 1 mm long; anthers 0.4-0.5 mm long; staminodes 0.8-0. 9

mm long. Nectary apparently absent or of two lobes, 0.8-1. 4 x 0.6-1. 1 mm, margin

entire or occasionally notched. Ovary sub-spherical to spherical, 0.9-1 x 0.9 mm,

densely pubescent to pilose, hairs straight or primarily straight with some hooked

hairs (hooked hairs are shorter and often most easily visible on the fruit operculum),

0.1-0.6(-l) mm long, on very upper portion of ovary; style 2. 9-4.2 imn long, usually

glabrous, occasionally 1 or 2 hooked hairs on base of style, 0.2 mm long; stigma 0.4

mm wide. Fruit cylindrical, ovate or obovate, 2, 1-2.3 x 1.7-1. 9 mm; operculum 0.6

mm long, pubescence as on ovary. Seed narrowly ovoid to ovoid, 0. 3-0.5 x 0. 1-0.2

mm, medium brown, pattern spiralled and reasonably even.

Distribution. Indonesia (Sulawesi, Maluku (Bum, Seram), West Papua), Philippines.

Habitat and ecology. Lithophytic on both limestone and other rocks, but also

occasionally terrestrial. Frequently found on stones and boulders in or beside streams

Revision of Epithema

173

and rivers, but also in rock crevices in humid, shady, if not wet, places. Collected from

0-1600 m altitude.

Provisional lUCN conservation assessment. Least Concern (LC). This species is

coimnon and very widespread. However, as it frequently occurs on karst limestone,

which is often commercially exploited, its status should be monitored.

Additional specimens examined: INDONESIA: Maluku: Burn, NW Bum, Bara, Waeduna

River, 30 Nov 1984, Nooteboom, H.P 5357 (L); Bum, Nov 1984, Sukendar SK740 (BO);

Maluku, Bum, Ehoe, 27 Sep 1921, Toxopeus, F.J. 358 (BO); Bum, Waeha, 12 Nov 1984, van

Balgooy, M.M.J. 4712 (E, L); Halmahera, Kampong Goal, Idjan 175 (BO); ibidem, 14 Sep

1951, Pleyte, D.R. 150 (BO, L); Seram, Tehoru, Saunulu, Murkele Ridge, 19 Feb 1985, Kato,

M. etal. C.6908 (L); Seram, Roho, 21 Nov 1917, Kornassi 482 (BO). Papua: Chaban, 28 Feb

1940, Kanehira, R. & Hatiisima, S. 11848 (A, K). Papua Barat: Raja Ampat, Misool, Near

Waima, 26 Sep 1948, Pleyte, D.R. 1053 (BO); Vogelkop Peninsula, Aitinjoe, 11 May 1958,

Versteegh, C. BW7405 (L); Biak to Parieri, 11 Sep 1966, Kostermans, A.J.G.H. & Soegeng

914 (BO, L); Biak, Kampong Saba, 31 May 1959, de Wilde, JJ.F.E. 1155 (L); Biak, 29 Aug

1915, Feuilletau de Bruyn, W.K.H. 335 (BO); Manokwari, Arfak Mts, Minjambau, 22 May

1962, Versteegh, C. BW 12694 (BO, L); Manokwari, Arfak Mts, Mupi Dessa, 16 Apr 1995,

Sands, M.J.S. 6841 (E, K); Manokwari, Arfak Plains, 25 Apr 1994, Sands, MJ.S. 6337 (E,

K); ibidem, 24 Apr 1994, Sands, M.J.S. 6318 (K); Manokwari, Kebar Valley, Andjai Airstrip,

Sobor Hills, 15 Nov 1954, van Royen, P. 5063 (A, E, L); Manokwari, Kebar Valley, Andjai

Village, 5 Nov 1 954, van Royen, P. 4946 (A, K, E); Manokwari, Mt Krabo, 3 Nov 1 960, Foster,

C. BW10773 (A, L); Manokwari, Warmare, Mokwam Trail, 10 Apr 1994, Sands, M.J.S. 6141

(K); Sorong, Warsamson Valley, 17 Aug 1961, Schram, FA.W BW 13259 (L); Wandammen

Peninsula, Wondiwoi Mountains, 28 Feb 1962, Foster, C. BW13665 (L); ibidem, 24 Feb 1962,

Schram, FA.W. BW 10631 (L). Sulawesi: s.l., 1871, RiedeH.w (P). Sulawesi Tengah: Gunung

Nokilalald, 5 May 1975, Meijer, W. 10045 (BO, L, US); Donggala, c. 57 km north of Palu on

Palu-Oti Road, 3 Jun 1979, van Balgooy, M.M.J. 3600 (L). Gorontalo: 9 Apr 2002, Mendum,

M. et al. 43 (E); Pinogu to Tulabolu, 15 Apr 2002, Mendum, M. et al. 113 (BO, E); Gunimg

Boliohutu, Mendum, M. et al. 180 (BO). Sulawesi Tenggara: Kesali-Porema, 23 Oct 1929,

Fjellbetg, G.F. 2616 (BO, S). Sulawesi Utara: G. Klabat, Pinili, 31 Oct 1973, Dransfield, J.

3869 (BO); Sangihe Island, Tamako, Kentuliang, 23 Nov 1998, Hicks, D. 151 (K).

PHILIPPINES: Agusan el Norte: Cabadbaran, Mt Urdaneta, Aug 1912, Elmer, A.D.E. 13470

(BO, E, G, GH, K, L, NY, P, US). Bohol: Aug 1923-Oct 1923, Ramos, M. BS43313 (BO, G,

K, P, US). Bukidnon: Jul 1913-Aug 1913, Escritor, L. BS21452 (US); Mt Galintan, Ramos,

M. & Edaho, G. BS48897 (NY). Cagayan: Penablanca, Mar 1929-May 1929, Ramos, M.

BS76791 (GH, SING). Davao del Sur: Sep 1911-Oct 1911, Weber, C.M. 1440 (E, NY, P).

Davao Oriental: Baguan River, 8 Mar 1949, Edam, G. 758 (A, PNH); ibidem, 8 Mar 1949,

Edaho, G. 746 (PNH); Davao, Mati, Mar 1927-Apr 1927, Ramos, M. & Edaho, G. BS49227

(NY); Davao, Mount Bilbogan, 1949, Edaho, G. 1965 (A, L, PNH). Davao del Norte: Mt

McKinley, 14 Aug \946, Edaho, G. 720 (PNH). Ifugao: Feb \9U, McGregor, R.C. BS19951

(US). Isabela: Barangay San Jose, 7 Mar 1997, Argent, G.C.G. et al. 29038 (E). Laguna:

Los Banos, Apr 1906, Elmer. A.D.E. 8318 (BO, E, G, K, NY); ibidem, 18 Dec 1903, Hallier,

H.G. 4128 (L); ibidem, 7 Mar 1906, Merrill, E.D. 5106 (BO, K, NY, US); ibidem, 14 Jan

1905, Williams, R.S. 2046 (NY); Mt Makiling, Jun 1917-Jul 1917, Elmer, A.D.E. 18283 (BO,

GH, K, US); ibidem, Jun 1917-Jul 1917, Elmer, A.D.E. 17551 (BO, GH); ibidem, Jan 1913,

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Card. Bull. Singapore 67(1) 2015

Robinson, C.B. BS17216 (US). Lanao del Sur: Lake Lanao, Cajnp Keitliley, Sep 1906-Oct

1906, Clemens, M.S. 724 (US). Leyte: Wenzel, C.A. 231 (G, US); ibidem, 28 Jun 1913, Wenzel,

C.A. 437 (GH); Mt Abucayan, Feb 1923, Edano, G. BS41748 \g, K, US). Negros Oriental:

Cuemos de Negros, Guinsuan Creek, 6 Jim 1948, Edano, G. 435a (A); ibidem, 6 Jun 1948,

Edano, G. 435 (PNH); Dumaguete, Cuernos Mts, Apr 1908, Elmer, A.D.E. 9698 (BO, E, G, L,

NY, US); Lake Balinsasayo, 1 6 Sep 1 948, Edano, G. 418 (A, PNH). Oriental Mindoro: Subaan

River, 14 Apr 1986, Coode, M.JE. 5313 (K, L). Palawan: Mt. Kabangaan, Apr 1929, Edano,

G. s.n. (GH). Quezon: Mt Cristobel, 14 Dec 1996, Reynoso & Majaducon PPI27265 (K).

Rizal: Morong, 25 Mar 1893, Loher, A. 4231 (K); Mt Angilog, Feb 1923, Lopez, G. BS42049

(BO, L, P, US); ibidem, Feb 1923, Lopez, G. BS42031 (US); Mt. Lumutan, Apr 1923, Ramos,

M. 42176 (BO). Samar: Matuguinao, Baruz, 20 Jan 1952, Gachalian, F. 147 (A, PNH); Mt

Purog, 24 Dec 1951, Edano, G. 34 78 (A); Mt Sarawag, 3 Dec 1951, Edano, G. 307 OA (PNH).

Sorsogon: Irosin, Mt Bulusan, Apr 1916, Elmer, A.D.E. 15877 (BO, C, G, GH, K, L, NY, P,

US); ibidem, 1915, Elmer, A.D.E. 15626 (BO, GH, NY). Zamboanga: Nov 1911-Dec 1911,

Merrill, E.D. 8222 (US); Sax River, 18 Feb 1905, Williams, R.S. 2401 (NY).

Notes. There are two informal forms of Epithema benthamii. The typical form is found

throughout the distribution of the species, while a finer, more thinly membranous and

more elongate fonn is found on Bum Island (Maluku) and Biak Island off Papua (e.g.

Sands 6318). These plants are similar to Epithema longipetiolatum, but still have the

typical leaf form and the characteristic long, straight or primarily straight, hairs on the

ovary and operculum of E. benthamii. The distributions of Epithema benthamii and E.

longipetiolatum overlap on Bum Island and Biak Island. See further discussion under

Epithema longipetiolatum. The indumentum of the leaf is useful for identification of

Epithema benthamii in that there are no minute hairs interspersed between the larger

hairs. The strigose hairs are often white, clearly visible and distinctive on the surface

of the dried leaves.

The specimen Kanehira and Hatusima 11848 (A) is unusual in that the peduncle

is branched and racemosely bears many complete inflorescences.

In the type specimen of Epithema brunonis var. scabrida the leaves are

somewhat dentate rather than entire as stated in the protologue, the indumentum is

of the same form as Epithema benthamii, albeit slightly denser, and the hairs on the

ovary/operculum are long and straight as in E. benthamii rather than the hooked hairs

of E. brunonis. It is quite clearly a synonym of Epithema benthamii.

The only apparent difference between Epithema calcicola and E. benthamii is

in the nan*ower somewhat reflexed bracts of E. calcicola. This species was known

only from the type collection and we conclude it is only a rather unusual specimen

of Epithema benthamii based on all other characters and that it occurs within the

distiibution of this species.

2. Epithema carnosum Benth., Scroph. Ind. 57 (1835); Barnett, FI. Siam. 3(3):

205 (1962), p.p.; Wang et ah, FI. China 18: 400 (1998); B.L.Burtt, Thai For. Bull.

(Bot.) 29: 93 (2001). - TYPE: Nepal, Nag Argute, 1832, Wallich, N. s.n. (lectotype K

[K000438688], designated here). (Fig. 3)

Revision of Epithema

175

Fig. 3. Distribution of Epithema carnosum Benth. (■), Epithema ceylanicum Gardner

(A) md Epithema piisillum (C.B. Clarke) Bransgrove (+).

Herb 6-38 cm high, caulescent; stem 1.5-5 mm wide with 2-3 nodes, intemodes

3-1 5 cm long, usually sub-glabrous to strigose or pubescent, rarely densely so. Leaves

thinly membranous to membranous, lower leaf petiolate, upper leaves petiolate or

sessile; petioles of lowest leaf 5-7.5 cm long, upper petioles 0-7 cm long; blades of

the lowest leaf 6.5-17 x 6-12.5 cm, upper leaves 2.5-12.5 x 2.3-6 cm, sub-spherical,

cordate or often ovate, symmetrical or barely asymmetrical, apex acute to broadly

acute, occasionally rounded, base sub-cordate to truncate, inserted evenly on petiole

or not, margin often sub-entire, but also serrate, dentate or crenate-sinuate (especially

upper leaves); hairs on upper surface to 1 .2 mm long; lower surface whitish-green or

paler than upper surface. Inflorescences 1-12 per plant; peduncles 0.5-7 cm long,

arising from leaf axils, from the petioles or from the leaf midribs, primarily from

the upper leaves; bracts small or sub-cucullate, enclosing a small portion of the

inflorescence, 1-13 x 3-9 mm, margin roughly entire with few dentate teeth to more

dentate; pedicels 1-5 mm long. Calyx light or whitish-green, 3. 3-5. 4 mm long; tube

2-3.3 X 1.5-2. 3 mm; lobes 1. 3-2.1 x 0.7—1 mm; indumentum outside a minute hooked

pubescence under strigose hairs 0.25-0.5 imn long. Corolla blue to dark puiple, some

with purple spots on the lower lip, 5. 5-7. 5 mm long; tube (3-)4-5.5 x (i_)i.5_2 mm

long; lobes 1-2.3 mm long; glabrous outside, villous band of hair inside. Stamen

filaments 0.6-0.8 mm long; anther dimensions unknown; staminodes 0.6-0. 8 mm

long. ISectary absent, of one lobe almost encircling ovary, or of two lobes, undulate,

each end a distinctly different height from the other, 0.6-1 x 1 .1-1.8 mm. Ovary sub-

spherical, 0.5-0.83 X 0.4-0.75 mm, pubescent or pilose, medium to high density, hairs

straight or primarily straight, hyaline, usually 0. 1-0.25 mm long, occasionally to 0.4

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Card. Bull. Singapore 67(1) 2015

mm long, on upper area of ovary; style c. 4 mm long, glabrous to pubescent, hairs

sparse to high density, to one third or half way up style, hairs hooked, hyaline, to

0.1 mm long; stigma c. 0.5 mm wide, Fmit sub-cylindrical 2. 1-2.5 x 1.7-2. 5 mm;

operculum c. 0.6 mm long, indumentum as ovary. Seed oval to fusiform, ends acute to

constricted, 0.4-0. 6 x 0. 1-0.2 mm, medium brown, pattern often straight to more or

less straight, but also to spiralled.

Distribution. India, Nepal, China, Myanmar, Thailand, Laos, Vietnam.

Habitat and ecology. Lithophytic. Found in humid, shaded places, often near water and

in rocky crevices at 0-2120 m altitude. Flowering and fruiting April, July-October.

Provisional lUCN conseryation assessment. Least Concern (LC). This species is

very widespread. However, as it frequently occurs on karst limestone, which is often

commercially exploited, its status should be monitored.

Additional specimens examined. INDIA: Andra Pradesh: Chintapalli, 31 Aug 1966, Subba

Rao, G. V. 28210 (E); Godavari, Rampa Country, Devarakonda, 6 Oct 1920, Narayanaswami,

M.A. 478 (CAL); Visakhapatnam, Araku, 24 Aug 1960, Balakrishnan, N.P. 10815 (BSI, CAL,

E). Meghalaya: Khasia, 28 Oct 1872, Clarke, C.B. 19076 (US); Khasia, Cherrapunjee, 26

Aug 1850, Hooker, J.D. & Thomson, T. 2161 (K); Khasia, Sohra, 22 Oct 1871, Clarke, C.B.

15665 (L); Mont. Khasia, Hooker, J.D. & Thomson, T. s.n. (C, CAL, E, GH, L, NY, P). Orissa:

Koraput, 8 Oct 1 950, Mooney, H.F 4045 (K); Koraput, Pottangi, 1 0 Jul 1 950, Mooney, H.F. 3853

(K) ; ibidem, 13 Jul 1950, Mooney, H.F. 3891 (K); Koraput, TuriaKonda, 10 Oct 1950, Mooney,

H.F. 4092 (K); Sikkim, 1878, King, G. s.n. (CAL); ibidem, Thomson, T. s.n. (BO, L); ibidem,

Unknown 3847 (K); Sikkim, Mongpo, 5 Oct 1884, Clarke, C.B. 36192 (G, K). Uttaranchal:

Kumaon, 25 Sep 1 884, Duthie, J.F. 2990 (K). West Bengal: Darjeeling, Unknown 12200 (BO);

Darjeeling, Mungpoo, 23 Sep 1875, Clarke, C.B. 24805 (K); Darjeeling, Rishap, 2 Aug 1870,

Clarke, C.B. 12300 (CAL, L); Darjeeling, Rishap, Rishap Ravine, 4 Sep 1869, Clarke, C.B.

8997 (L).

CHINA: Heou-hay tse, 7 Jul 1907, Esquirol, J. 711 (E). Guangxi: Tan-Ngar, 12 Jul 1928,

Clung, R.C. 6421 (A). Yunnan: Shweli-Salwin Divide, 1917-1919, Forrest, G. 15699 (E, K);

Simao, Henry, A. 13497 (K, US); ibidem, Henry, A. 12280 (K); ibidem, Hemy, .4. 12705 (K).

MYANMAR: Pegu, Scott 3 (L); Mandalay, Merktila, Taungbaw-yo, 22 Oct 1936, Smith, H.C.

16273 (K); Mandalay, Myingyan, Pope Hill, 4 Aug 1909, Lace, J.H. 4892 (E, K).

THAILAND: Chiang Mai: Chiang Dao, Doi Cliiang Dao, 22 Jul 1989, Maxwell, J.F. 89-920

(L) ; ibidem, 27 Sep 1971, Murata, G. et al. 15146 p.p. (C, K, KYO); ibidem, 27 Sep 1971,

Murata, G. et al. T.15156 (KYO); ibidem, 19 Oct 1926, Put 410 (K); ibidem, 26 Oct 1979,

Shimizu, T. et al T.20909 (KYO). Mae Hong Son: Muang, Doi Mae Sakut, 23 Sep 1995,

Nanakorn, W. et al. 4664 (E). Nan: Sapan Waterfall, 7 Sep 1995, Larsen, K. et al. 46161

(AAU). Tak: Nam Tok Phacharoen National Park, 23 Apr 2004, Poorna, R. et al. 4580 (BKF).

LAOS: Sayabourj^: Phiang, Nam Phon, 1 Sep 1999, Maxwell, J.F. 99-278 (A, L).

VIETNAM: Bac Kan: Na Ri District, Liem Thuy Municipality, Na Bo Village, 15 Jul 2004,

Atha, D. 4785 (NY); Ha Giang: Pho Bang, Pho Cao, 12 Aug 1977, Phuong, V.X. 311 (HN).

Notes. Epithema carnosum is widespread across tropical Asia with the exception of

Malesia. It is similar to some of the forms of Epithema ceylanicum and was originally

Revision of Epithema

111

separated from it (as E. dentatum) by E. carnosim having crenate-sinuate leaves vs. E.

dentatum having small teeth on the leaf margin that are sometimes subacute or doubly

seiTate. Also Epithema dentatum was said to have shorter petioles on the upper leaves.

These characters cannot consistently be used to separate Epithema carnosum from

the now more broadly defined E. ceylanicum which includes E. dentatum. It is better

separated from Epithema ceylanicum by the hairs on the ovary and operculum. They

are straight in E. carnosum and hooked or primarily so in E. ceylanicum. In nearly

all specimens from India, Nepal and Myanmar there is no variation in this character,

but on specimens from China, Thailand and Vietnam a proportion of the hairs on

the ovary/operculum of Epithema carnosum may be hooked. In all cases where the

majority of the hairs are straight, the specimens are assigned to E. carnosum. This is

also essentially tine of Epithema ceylanicum where the hairs on almost all specimens

from India and Myanmar have only hooked hairs on the ovary and operculum but

some specimens from north and north-west Thailand also have some straight hairs

on the ovai^/operculum. In all cases where the majority of the hairs are hooked, the

specimens are assigned to E. ceylanicum.

Epithema carnosum occasionally produces peduncles from the petioles of the

leaves of the tenninal leaf pair. Peduncles arising from the midrib of these leaves has

only been observed in specimens of Epithema carnosum from Khasia in Northeast

India.

3. Epithema ceylanicum Gardner, Calcutta J. Nat. Hist 6: 492 (1846). - Epithema

carnosum var. ceylanicum (Gardner) C.B. Clarke in A.DC. & C.DC. Monogr. Phan.

5(1): 178 (1883). - TYPE: Sri Lanka, Gardner, G. 606 (lectotype K, designated here).

(Fig. 3-5)

Epithema carnosum var. hispidum C.B. Clarke in A.DC. & C.DC., Monogr. Phan.

5(1): 178 (1883), - Epithema dentatum subsp. hispidum (C.B. Clarke) Hilliard &

B. L.Burtt, Edinburgh J. Bot. 54: 112 (1997). - TYPE: India, Tamil Nadu, Western

Ghats, Courtallum, August 1 835, Wight, R. 2350 (lectotype K, designated here)

Epithema carnosum var. dentatum C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1):

111 (1883), - Epithema dentatum (C.B.Clarke) Hilliard & B.L.Burtt, Edinburgh J.

Bot. 54: 111 (1997). - TYPE: Bunua, Mon, Moulmein, Farm Cave Rocks, Parish,

C. S.P 63 (lectotype K, designated by Hilliard & Burtt (1997)).

Epithema brunonis veiv. fasciculatum C.B.Clarke in A.DC. & C.DC., Monogr. Phan.

5(1): 180 (1883). - Epithema taiwanense vm. fasciculatum (C.B.Clarke) Z.Yu Li

& M.T.Kao, FI. Taiwan ed. 2, 4: 697 (1998). - TYPE: Philippines, Tayabas, 1841,

Cuming, H. 823 (lectotype K [K000438698], designated here; isolectotype BM,

G-DC, K, L, P).

178

Card. Bull Singapore 67(1) 2015

Fig. 4. Epithema ceylanicum Gardner. Leaves and inflorescenees. (Photo: Preeeha Karaket)

Fig. 5. Epithema ceylanicum Gardner. Flowers. (Photo: Preeeha Karaket)

Revision of Epithema

179

Epithema taiwanense S.S.Ying, Coloured 111. FI. Taiwan 4: 795 (1992); Li & Kao, FI.

Taiwan ed. 2, 4: 697 (1998); Wang et al., FI. China 1 8: 400 (1998). - TYPE: Taiwan,

Chia-yi, Kuanyin waterfalls, 200-300 m, 25 June 1 992, Yirjg, S.S. s.n. (holotype NTUF

n.v.).

Herb 4^0 cm high, caulescent or with very short primary intemode and appearing

acaulescent; stem 1-10 mm wide, intemodes 0.5-15 cm long, sub-glabrous, sparsely

strigose or sparsely pubescent, hairs 0.5-1 imn long. Leaves thinly to strongly

membranous, lowest leaf always petiolate, upper leaves sessile to petiolate, sometimes

with a solitary upper, sessile leaf per plant; petiole of lowest leaf (0.2-)l-12 cm long,

sub-glabrous to densely pubescent, hairs to 0.7 mm long, petioles of upper leaves 0-2

cm long, sub-glabrous, minutely pubescent or with longer hairs of medium to high

density, to 1 mm long; blades of lowest leaves 3-20 x 1.8-20 cm, upper or solitary

leaves ( 1 -) 1 .5-1 1 x 0.8-7 cm, lower leaves cordate to ovate, upper leaves cordate, ovate

or elliptic, occasionally almost oblong or sub-spherical, symmetrical or asymmetrical,

apex acute to rounded, base of lower leaf cordate to truncate, base of upper leaves

obtuse, rounded or truncate, occasionally sub-cordate, inserted evenly on petiole or

not, where inserted unevenly one side is often cordate, the other truncate, margin sub-

entire to dentate, bidentate or seirate; upper surface pale- to mid-green, with sparse

straight and/or hooked hairs to 0.9 mm long and sparsely to densely strigose, hairs

0.25-1 .5 mm long; lower surface paler green or purplish in colour, sub-glabrous to

strigose, hairs sparse to medium density, sometimes with hooked and straight hairs on

veins to 0.3 mm long. Inflorescences 1-6 (to many) per plant; peduncles arising from

the leaf axil, the petiole or the base of the midrib, usually from the upper leaves, often

rather fasciculate, (0rt-)l-10(-17) cm long, often of very variable lengths in a single

cluster, sub-glabrous to densely setose, hairs to 1.1 mm long; bracts (sub-)cucullate

but enclosing a small proportion of the inflorescence if at all, 3-16(-20) x 3_9(_i0)

mm, 0.6-2(-3.4) times as long as wide, margin entire to partially or entirely dentate;

lower surface glabrous to strigose, hairs sparse to medium density, hairs 0.25-0.8 mm

long, with or without shorter layer of hair 0. 1-0.4 mm long; upper surface glabrous

to sparsely strigose, hairs to 0.25 nun long, hairs on upper half of bract or towards

margins; pedicels 0.5-6 mm long, densely pubescent with fine minute hairs and more

sparsely with larger strigose hairs to 1 mm long. Calyx 2.1-8 mm long; tube 0.8-6 x

1-2.9 mm; lobes 1-2.5 x 0.7-2. 1 mm; sub-glabrous or sparsely to densely pubescent

outside with fine, minute straight and hooked hairs to 0.25 mm long, with or without

sparse, strigose hairs, to 0.4 mm long; glabrous or subglabrous inside, sometimes with

strigose hafts on inside of lobes. Corolla whitish or blue, blue-violet or pink, tube

white, 5-8.5 mm long; tube 3.2-6.3 x 1-3 mm; lobes 1-3 x 1-1,3 mm long, margins

often slightly fimbriate; glabrous to sparsely pubescent outside on lobes, hairs straight;

dense band of villous hair inside, one-third to three-quarters way up the tube from the

base. Stamens 1. 4-2.1 nun long; filaments 1-1.7 mm long; anthers 0.4-0. 5 mm long;

staminodes 1-1.7 mm long. Nectary apparently absent or one or two discrete lobes,

may almost encircle ovary or not, margin entire or sub-entire, undulate or notched,

0.8-1. 8 X 1-1.8 mm. Ovary sub-spherical to cylindrical, 0.7-1. 5 x 0.9-1. 5 mm,

180

Card. Bull. Singapore 67(1) 2015

sparsely to densely pubescent on the upper half or uppermost part of the ovary, hairs

hooked or primarily hooked, hyaline, to 0.15 mm long; style 2.5-5 mm long, glabrous

or sparsely pubescent in lower half, hairs hooked; stigma to 0.5-0. 8 mm wide. Fruit

spherical to cylindrical, 1.5-5 x 1-3 mm; operculum 0.4-0. 8 mm long, sparsely to

densely pubescent as on ovary. Seed sub-cylindrical to ovoid, 0.4-0. 7 x 0. 1-0.2 mm,

pattem often straight or almost straight, but also to spiralled, regular, walls clearly

defined, often appearing thickened and rigid.

Distribution. India (including Andaman Islands), Sri Lanka, Taiwan, Myanmar,

Thailand, Cambodia, Vietnam, Philippines.

Habitat and ecology. Lithophytic. Found on rocks and rock faces in wet, humid, shady

places, occasionally on wet tree trunks at 0-2000 m altitude.

Provisional lUCN conset^ation assessment. Least Concern (LC). This species is

very widespread. However, as it frequently occurs on karst limestone, which is often

commercially exploited, its status should be monitored.

Additional specimens examined. INDIA: Andaman and Nicobar Islands: North Andaman,

Saddle Peak National Park, 10 Feb 2000, Sumathi, R. 17895 (K). Karnataka: Chorla

Ghats, Chorla, Stocks s.n. (K); Coorg, Mercara, Jersey Falls, 12 Sep 1934, Barnes, E. 928

(K); Mysore, Hassan, Bisle ghat, 14 Aug 1971, Ramamoorthy, T.P. HFP2007 (US); Mysore,

Hassan, Shiradi ghat, 7 Aug 1969, Saldanha, C.J. 14426 (US); Mysore, Kenchankumri State

Forest, 15 Aug 1971, Ramamoorthy, T.P. HFP2035 (K, US); Mysore, Shimoga, Hulical ghat,

24 Aug 1963, Raghavan, R.S. 90185 (E). Kerala: Idikki District, Sabarimalai slopes, 26 Sep

1972, Sharma, B.D. 42032 (E); Kochi (Cochin), Parambikulam, 25 Sep 1935, Barnes, E. 1300

(K); Kochi (Cochin), Parambikulam Hills, Dec 1934, Barnes, E. 1056 (K); Palghat, Silent

Valley, 21 Aug 1966, Vajvavelu, E. 27558 (L). Madhya Pradesh: Bastar, Upper Kanger Valley,

26 Aug 1959, Subramanyam, K. 8644 (E). Maharastra: Concan, Hooker, J.D. & Thomson,

T. s.n. (P); ibidem. Stocks, Law & co. s.n. (C, L). Orissa: Dhenkanal State, Goyalpathar ghati

valley, 16 Sep 1942, Mooney, H.F 2067 (K); Dhenkanal State, Kapilas Hill, Mooney, HE 2754

(K) ; Dhenkanal State, Saptasajya, 20 Sep 1993, Dhal, N.K. & Rout, N.C, 8122 (E); Ganjam,

Budhakliol Pahad, 14 Aug 1971, Brahmam, M, 2776 (E). Tamil Nadu: Banliyar, ?Wight s.n.

(L) ; ibidem, ? Wight s.n. (L); Madurai, Way, 14 Oct 1959, Subramanyam, K. 8975 (E); Mont.

Nilghiri & Kurg, Thomson, G. 121 (K); ibidem, Thomson, G. s.n. (E, G); Vilangadu Forest, 27

Aug 1985, Pradeep, A.K. 5850 (CAL); Western Ghats, Courtallum, W. C. 2680 (E); ibidem,

Wight, R. 556 (K); Western Ghats, Courtallum, Swamiar Falls, 2 Oct 1975, Nan; K.K.N. 1354

(CALI). West Bengal: Kolcata, Kabini tributory falls, 17 Aug 1964, Ellis, J.L. 20452 (E).

SRI LANKA: s.l., 1891, Deschamps s.n. (G); Mrs. General Walker 140 (K); Mrs. General

Walker s.n. (K); Thwaites, GH.K. 2844 (BO, G, K, P); 1 836, Wight, R. 1/162 (K). Uva: Badulla

District, Between Mesitale and Dunhinda Falls, 26 Jan 1982, Namba, T & Mikage, M. 82-

413 (KYO); Badulla District, Dunhinda Falls, 8 Jan 1983, Lucas 1025 (US). Central: Matale

District, Wiltshire Forest, 2 Jul 1974, Sumithraarachchi, D.B. DBS388 (K); Deltotte, Nov

1888, Unknown s.n. (SING); Kandy, Laksapana, 19 Sep \912, Jayasuriya, A.H.M. 843 (US);

Peradeniya, 10 Sep 1934, Unknown 753 (L). Sabaragamuwa: Kegalle District, Dolosbage,

Windsor Forest, 17 Nov 1978, Grey-Wilson, C. & Grey-Wilson, C.M. 3016 {K. US); Kegalle

District, Kadaganawa, 1 Sep 1969, Grupe, D.A. 200 (US); ibidem, 22 Aug 1968, Theobald,

Revision of Epithema

181

W.L. & Gmpe, D.A. 2404 (US); ibidem, 30 Jul 1968, Theobald, W.L. & Grupe, D.A. 2368 (US);

Ratnapura, Katussagala Hill, 5 Dec 1976, Faden, R.B. & Faden, A.J. 76/494 (US); Ratnapura,

Pannil Kanda, 5 Jul 1975, Waas, S. 1407 (K); Ratnapura, Rakwana, 7 Dec 1977, Fosberg, FR.

57278 (K, US).

TAIWAN: Takow, Ape’s Hill, Hemy, A. 1916 (E, K, NY, US); Kaohsiung Hsien, Shuangchi,

Meilong, 31 Oct 1988, Hsieh, C.F et al. s.n. (E); Kaohsiung Hsien, Takangshan, 14 Nov 1988,

Yang, K.C. 3465 (E).

MYANMAR; Mon: Moulmein, Jul 1880, Brandis, D. 676 (K); ibidem, Lobb, T 387 (K);

ibidem, McKee, H.S. 1958 (P); ibidem, Parish, C.S.P. 133 (K); Moulmein, Kyauk Ta Lon, Jul

1958, McKee, H.S. 6317 {K, P).

THAILAND: Bueng Kan: Mueang, Chaiyaphon, Singhanat Ban Phot Temple, 26 Aug 2001,

Pooma, R. et al 2697 (BKF). Chiang Mai: Chiang Dao, Pooma, R. 1059 (BKF); ibidem, 16

Jul 1958, Smitinand, T. 47/P (BKF); ibidem, 15 Aug 1963, Smitinand, T. & Sleumer, H.O. 1006

(BKF); Chiang Dao, Doi Chiang Dao, 27 Sep 1971, Murata, G. et cd. 15146 p.p. (L); ibidem,

17 Jul 1958, Sorensen, T et al. 4235 (A, BKF, C, E); Chiang Dao, Doi Chiang Dao, Between

Pong Pho and Khun Klong, 3 1 Jul 1 968, Larsen, K. et al. 2923 (AAU); Doi Sutep, 1 0 Oct 87,

Maxwell, J.F. 87-1154 (BKF, L); ibidem, 12 Sep 1958, Sorensen, T. et al. 4936 (ABD, C); Mae

Ram Subdistrict, Doi Sutep-Pui National Park, Mae Ram Stream, 10 Sep 1992, Palee, P. 73

(A, CMU, GH, L); Fang, Doi Ang Khang, 21 Sep 2008, Middleton, D.J. et al. 4544 (E); Mae

Dang, Mawn Ngaw, Doi Mawn Ngaw, 12 Aug 2002, Palee, P. 544 (L). Chumphon: Sapli, 8

Sep 1927, Put 1015 (ABD, K). Kanchanaburi: 15 Aug 1968, Nimanong, B. & Phusomsaeng,

S. 288 (BKF); ibidem, Suvanakoses, P. 2060 (BKF); Kanburi, Baw Re, 21 Jul 1926, Put 219

(K); Sai Yok, Sai Yok National Park, Lam Tam Keio, 10 Aug 1982, Shimizu, T. et al. T.28519

(BKF, SING); Sangklilaburi, 28 Jul 1968, Prayad 1422 (BKF); Thong Pha Phum, 4 Jul 1973,

Maxwell J.F. 73-107 (AAU, BK); Wang Pho, 26 Oct 1969, Kasem 641 (BKF); Khon Kaen,

Chum Phae, Phu Pha Man national park, Tham Pha Puang, 1 Aug 2011, Nonsaengsri, M. et

al. 7965 (QBG); Khon Kaen, Nanong Turn, Ban Na Chan, 23 Aug 2011, Norsaengsri, M. et

al. 8048 (QBG); Khon Kaen, PhaNok Khao, 9 Sep 1963, Smitinand, T & Sleumer, H.O. 1109

(K, L, SING). Lampang: Doi Pang La, Huay Tak, 25 Sep 1967, Shimizu, T. et al. T. 10789

(BKF); Muang Bahn, Chae Son, Jae Sawn Stream, 22 Aug 1995, Maxwell, J.F. 95-536 (BKT,

CMU, L); Ngao, Near Tham Pha Thai, 24 Sep 2008, Middleton, D.J. et al. 4579 (E); Loei,

Phu Kradung, 15 Oct 1967, Prayad 1034 (BKF). Loei: Phu Ruea District, Phu Luang, 16

Sep 1966, Phusomsaeng, S. & Bunchuai, K. 42 (BKF, L). Mae Hong Son: Muang, Doi Pui,

23 Sep 1995, Larsen, K. et al. 46861 (AAU); Mueang, Tliam Pla-Namtok Pha Suea National

Park, Pha Daeng Cave, 22 Aug 2013, Norsaengsn, M. 10568 (QBG). Nakhon Nayok: Nang

Rong Falls, 4 Aug 1992, Larsen, K. et al. 43771 (P). Nan: Tham Pha Toop Forest Park, 2 Sep

1999, Middleton, D.J. 149 (A, BKF, E, K); Muang, Tham Phatup Forest Park, 16 Aug 2012,

Middleton, D.J. et al. 5613 (E, SING); Muang Tam PaaToop, 13 Sep 1995, Nanakorn, W. 4240

(QBG); Tham Pha Tok, 25 Jul 1992, Larsen, K. et al 43589 (AAU); ibidem, 13 Sep 1995,

Larsen, K. et al 46426 (AAU). Phayao: Doi Luang National Park, Champa Thong Waterfall,

9 Aug 1997, Petrmitr, O. 70 (L). Prachuap Khiri Khan: Bang Saphan, Khao Maa Rong, 5 Sep

2008, Middleton, D.J. et al. 4284 (E), Phrae: Nam Pu, 23 Sep 1 9 1 2, Van Pruk, L. 350 (BK, K).

SaKaeow: KhaoChakan, 17 Oct Staples, G.W. etal. 1409 {E, SING). Saraburi: Kaeng

Khoi, Amata-khuha Cave, 20 Aug 2001 , Pooma, R. et al 2100 (BKF); Kaeng Khoi, Tham Pra

Photisat, 7 Oct 1979, Shimizu, T. et al. T. 194 13 (KYO, L); Muak Lek, 10 Nov 1924, Marcan,

A. 1869 (ABD, K); ibidem, 3 Sep 1928, Put 1853 (K); Phraphutthabat, Wat Khaowong, 18 Sep

2004, Pooma, R. et al. 4824 (E), Surat Thani: Kanchanadit, 31 Jul 1927, Kerr, A.F.G. 13053

(K); Koh Samui, Kao Noi, 28 Jun 1966, Sakol 1150 (BKF). Tak: Mae Sot, Doi Muser, 20 Aug

182

Card. Bull. Singapore 67(1) 2015

1961, Chermsirivathana, C. 1 (BKF); Mae Sot, Khao Pha Wo, 23 Jul 1973, Murata, G. et al.

T.16922 (AAU, BKF, KYO, L); ibidem, 12 Oct 1979, Shimizu, T. et al. T.18519 (KYO); Mae

Sot, Wat Tham Inthanin, 11 Sep 2009, Middleton, D.J. & Triboun, P 4846 (E). Tak: Umphang,

Takhobi Cave, 18 Oct 2014, Middleton, D.J. et al. 5768 (BKF, E, SING). Uthai Thani: Lan

Sale, Huppatat Non Flunting Area, 14 Oct 2014, Middleton, D.J. et al. 5687 (BKF, E, SING).

CAMBODIA: Kampot: 20 Sep 1903, Geoffray, M. 95 (P).

VIETNAM: Lang Son: hmg\mc,Eberhardt 3363 (P).

PHILIPPINES: Apayao: May 1917, Fenix, E. BS28106 (US). Benguet: Mt Trail, Oct 1929,

Quisumbing, E. BS78105 (NY); Twin Peaks, 8 Sep 1904, Williams, R.S. 937 (NY). Bukidnon:

Tanculan, Jul 1916, Fenix, E. BS26049 (K, NY, US). Bulacan: 8 Sep 1935, Bartlett, H.H.

14744 (GH), Ilocos Norte: Bangui, Nov 1923, McGregor, R.C. BS43566 (K); Burgos, Jul

1918, Ramos, M. BS32892 (GH, K, US). Laguna: Los Banos, Loher, A. 1555 (K). Oriental

Mindoro: Mansalay, Mt Yagaw, Nov 1952-Dec 1952, Sulit, M.D. & Conklin, H.C. 16913 (L,

PNH). Pampanga: Mt Arayat, Sep 1905, Merrill, E.D. 4216 (K). Rizal: Aug 1911, Ramos, M.

BS13624 (BO, G, K, L, P, US). Rizal: Antipolo, Jul 1917, Ramos. M. & Edaho, G. BS29466

(BO, P, US); Kay Ungulan, 18 Aug 1935, Bartlett, H.H. 15380 (GH); Montalban, 11 Aug 1935,

Bartlett, H.H. 14487 (GH, PNH); ibidem, 3 Aug 1990, Loher, A. 1556 (K); ibidem, Dec 1904,

Loher, A. 6669 (K); ibidem, Dec 1904, Loher, A. 6668 (K, US); ibidem, 26 Sep 1890, Loher, A.

1554 (K, P, US); ibidem, Nov 1909, Merrill, E.D. 52 (G, US); ibidem, 2 Nov 1909, Robinson,

C.B. BS9534 (NY); ibidem, 2 Nov 1909, Robinson, C.B. BS9525 (L, US); Morong, Aug 1906,

Ramos, M. BS1435 (NY, US).

Notes. Epithema ceylanicum is a widespread and variable species. The protologue of

Epithema ceylanicum states the plants are hispidly pilose, have a persistent style and

have five distinct nectaries. Only a proportion of the specimens from Sri Lanka are

densely pubescent, only the basal section of the style is persistent and we have seen no

more than two nectaries. As a variety Clarke (1883) separated Epithema carnosum var.

ceylanicum (as ""zeylanica'") from E. carnosum var. carnosum on its larger calyx and

seeds that are twice as large. However, neither of these characters holds up to closer

scrutiny. Instead, it is easily distinguished from Epithema carnosum by the hooked

rather than straight hairs on the operculum.

Rarely, it can be difficult to differentiate Epithema ceylanicum fromE’. saxatile,

particularly as they both have hooked hairs on the ovary/operculum. They are most

easily distinguished in the size of the bract which in Epithema saxatile usually encloses

the inflorescence.

Epithema taiwanense was previously distinguished by the presence of many

short, often fasciculate, peduncles with very small inflorescences and quite consistently

rounded or rounded-acute leaf apices. However, these characters are within the overall

range of variation of the very variable Epithema ceylanicum and we have, therefore,

decided to synonymise the names and recognise a single species.

Some specimens bear a resemblance to Epithema involucratum but they can be

distinguished by the presence of hair in the corolla and the usually numerous, small

inflorescences among other characters. One of the specimens from the Philippines,

Fenix BS28106 (US), somewhat resembles Epithema benthamii in leaf shape but is,

again, easily separated by having hooked hairs on the ovary/operculum rather than the

straight hairs of E. benthamii.

Revision of Epithema

183

One collection from the Philippines, R.S. Williams 937 (NY(x2)), is rather

distinct from most of the material of this species in the Philippines in its acaulescent

habit and sessile upper leaves. It also has large bracts. None of these characters is

unique in the species but the combmation is unusual. Further collections are necessary

to see whether this is distinct from Epithema ceylanicum.

Some collections of this species have strongly variegated leaves but this

character does not appear to have any taxonomic significance.

4. Epithema dolichopodum Hilliard & B.L.Burtt, Edinburgh J. Bot. 54: 112 (1997).

- TYPE: Malaysia, Sabah, Lahad Datu, Bukit Tempadong, 100-150 m, 11 June 1984,

Beaman, J.H. 10068 (holotype E; isotypes GH, K, L, MSC, NY, US). (Fig. 6)

Herb 7-28 cm tall, acaulescent with first leaves arising at ground level, sometimes

with 1-2 additional elongated intemodes, indumentum of vegetative organs sparsely

to densely strigose and/or setose, hairs to 0.6 mm long; stem 2-6 mm wide with

0-2 nodes, intemodes to 8 cm long. Leaves membranous to sh'ongly membranous,

petiolate, solitary, opposite one or more peduncles; petioles (1.5-)5-37 cm; blades

5- 24 X 4-15.5 cm, cordate, broadly ovate or occasionally sub-spherical, symmetrical

to barely asymmetrical, apex usually rounded, occasionally acute, base cordate or

sub-truncate, occasionally sub-auriculate, inserted evenly on petiole or not, margin

sub-entire to dentate and bidentate. Inflorescences 1-5 per plant; peduncles (2-)9-27

cm long, originating from the base of the plant or opposite solitary leaves; bracts

pale green or green, cucullate but not always enclosing entire inflorescence, 10-25 x

6- 14 mm, 1.2-2. 5 times as long as wide, margin dentate; pedicels 2. 1^.6 mm long,

sub-glabrous to densely pubescent, hairs minute and hooked. Calyx lobes purple or

purple only at tips, 3. 8-5. 6 mm long; tube 2-4 x l .5-3 mm, lobes triangular, 1 .3-2.7 x

0.8-2 mm; minutely densely pubescent outside, hairs white; glabrous to very sparsely

pubescent inside, hairs minute, in tips of lobes. Corolla tube white, lobes blue, with or

without piiiple markings on lobes, 5-8.4 mm long; tube cylindrical, 3. 3-6. 3 x 1.3-3

mm, lobes 1 ,7-2. 1 mm long, margin slightly fimbriate to fimbriate; band of villous hair

inside. Stamen filaments 1 mm long; anther characters unknown; staminodes 1 mm

long. Nectary of two discrete lobes, 0.4-1. 5 x 0.8-1 .7 mm, margin undulate. Ovary

sub-cylindrical to sub-spherical, 0.8-1. 3 x 0.6-1 .3 mm, minutely pubescent on upper

portion of ovary, medium to high density, hairs hooked, hyaline or white, 0.04-0.1 mm

long; style 3.7-4. 1 mm long, glabrous or rarely with minute, hooked hairs; stigma c.

0.5 mm wide. Fruit cylindrical to obovate, 2.3-3 x 2. 1-2.5 mm; operculum 0.6-0. 8

mm long, glabrous or as on ovary. Seed narrowly ovoid to ovoid, 0.3-0. 6 x 0. 1-0.2,

dark brown, pattern partially spiralled or spiralled, regular.

Distribution. Malaysia (Sabah), Philippines (Palawan).

Habitat and ecology. Lithophytic on limestone. Found on boulders, outcrops, cave

entrance walls in humid and shaded places at 90-150 m altitude. Flowering and

fruiting in June, September-November.

184

Card. Bull. Singapore 67(1) 2015

Fig. 6. Distribution of Epithema dolichopodum Hilliard & B.L.Burtt (A), Epithema horsfieldii

(R.Br.) DC. (•), Epithema madulidii Hilliard & B.L.Burtt (★), Epithema philippinum (Hilliard

& B.L.Burtt) Bransgrove (+) and Epithema tenerum (C.B. Clarke) Hilliard & B.L.Burtt (■).

Provisional lUCN conservation assessment. Least Concern (LC). Although this species

is not especially widespread, its distribution is not restricted enough to fall into the

Vulnerable category under the B criteria if there were accompanying threats. However,

as it fi'equently occurs on karst limestone, which is often commercially exploited, its

status should be monitored.

Additional specimens examined. MALAYSIA: Borneo: Sabah: Hutan Simpan Madai,

15 Sep 1976, Tamura, M. & Hotta, M. 690 (KYO); Kudat, Pulau Balembangan, Oct 1994,

Anthonysamy, S. etal. SB7 (E); Kudat, Pulau Balembangan, Kampung Sina, 9 Apr 1977, Stone,

B.C. & Anderson, E.F SAN86719 (KLU); Kunak, Madai Hill, 9 Jun 1996, Lim, S.P et al.

LSP681 (SING); ibidem, 9 Jun 1996, Lim, S.P. et al. LSP675 (SING); Lahad Datu, Segama

River, Batu Sarang, 25 Aug 1999, Kiew, R. RK4759 (SING); Madai Baturong Forest Reserve,

Madai Caves, 14 Nov 1968, Kokawa, S. & Hotta, M. 1125 (KYO, L); ibidem, 14 Nov 1968,

Kokawa, S. & Hotta, M. 1073 (KYO); Segarong, Kiew, R. et al. RK4321 (K, SENG); ibidem,

Symington, C.F. & Agama, J. 9361 (K, SING); Sukau, Bod Tai, 14 Sep 1996, Kiew, R. & Lim,

S.P. RK4138 (K, SING); Tawao, Oct 1922-Mar 1923, Elmer, A.D.E. 20569 (G, GH, K, L,

SING); Tawau District, Mount Wullersdorf Forest Reserve, Teck Guan Quarry, Nadiah, I. et

al. SAN149136 (SING).

PHILIPPINES: Palawan: Mt Kabangaan, Apr 1929, Edano, G. BS77711 (GH); Mt

Mantalingajan, Apr 1929, Edano, G. BS77570 (GH, SING).

Notes. This species is most easily confused with other acaulescent species such as

Epithema strigosum, E. longipetio latum and^”. rennellense. One of the differentiating

features of Epithema dolichopodum are the very long peduncles which are usually

Revision of Epithema

185

(l-)2(-3) times the height of the rest of the plant. In Epithema rennellense, E.

longipetiolatum and E. strigosum the peduncles are shorter to barely longer than the

height of the plant. Other differences are discussed in the notes section of the other

species.

The hairs on the ovary and operculum are hooked and minute, the only variation

being the density of the hairs. While it is infrequent, the lower style may be pubescent

with hooked hairs. Fruit with glabrous and pubescent styles were observed from the

same plant.

While the upper surface of the leaves can be densely strigose, the surface of the

leaves and the stem may appear glabrous due to the small size of the hairs.

The collections available for examination are from relatively few sites, four of

which were in close proximity to each other.

To-date, the only other species of Epithema found in Sabah are E. saxatile and

E. sarawakense.

S. Epithema horsfieldii (R.Br.) A.DC,, Prod. 9: 279 (1845). -Aikinia horsfieldii R.Br.,

PL Asiat. Rar. 3: 66 (1 932). - TYPE: Indonesia, Java, 1 802-1818, Horsfield, T 85, also

labelled Cyrtandn 3 (lectotype BM [BMOO 1125214], designated here; isolectotype K

(x2)). (Fig. 6)

Epithema difformis Span., Linnaea 15: 331 (1841). - Carpocalymna monophylla

Zipp. ex Span., Linnaea 15: 331 (1841), nom. inval. - TYPE: Indonesia, Timor, 1841,

Spanoghe, J.B. 57 (lectotype P, designated here; isolectotype L).

Epithema horsfieldii var. epiphyllum Hilliard & B.L.Burtt, Edinburgh J. Bot. 54: 112

(1997). - TYPE: Indonesia, Western Sumbawa, Batudulang and Sampar Olat Ranges,

Mt Batulanteh, April 1961, Kostermans, A.J.G.H. 18839 (holotype L; isotypes BO

(image seen), K).

Herb 3.2-22 cm high, caulescent (but can appear as just one large leaf with the stem

mistaken for a petiole), unifoliate or rarely with two leaves; stem 0.5-3. 8 mm wide.

Leaves sessile (but see above), membranous to strongly membranous; blade 3.2-17.5

X 2.4-13 cm, usually cordate or sub-cordate but also to oblong, symmetrical or not,

apex broadly acute to rounded, base cordate to truncate, inserted evenly on petiole,

margin entire to dentate, bidentate, sen'ate or crenate, arrangement of veins variable

but the lower secondary veins often appearing to arise from around the base of the

peduncles, often widely spaced, occasionally running down sides of midrib; upper

surface pale green to green; lower surface light green or purplish. Inflorescences 1-1 5

per plant; peduncles 0.3-13.3 cm long, originating from the midrib at the base of the

blade, or from the midrib up to 2.3 cm from the base; bracts cucullate, sub-cucullate

or small and only enclosing a small proportion of the inflorescence, 2-22 x 3-12 mm,

margin variable, irregularly dentate to dentate, occasionally lobed; pedicels 0.7-5 mm

long. Calyx 2. 5-5. 7 mm long; tube 1-4.4 x 0.9-2. 7 mm; lobes 0.6-5 x 0.4-2. 3 mm;

186

Card. Bull. Singapore 67(1) 2015

glabrous inside or rarely sub-glabrous with hairs to 0.1 mm long. Corolla blue, lilac,

lilac-blue, violet-purple and white in proximal half of the two upper lobes or violet

only at tips, 6.5-12 mm long; tube cylindrical 4-9 x 0.9-2 mm; lobes 1.5^.2 x ]

mm, margin undulate to slightly fimbriate; band of villous hair inside at one third to

half way up the tube, occasionally the band is incomplete or the hair sparse. Stamens

0.7-1. 9 mm long; filaments 0,6-1. 7 mm long; anthers 0.3-0. 6 mm long; staminodes

0.4-1 mm long. Nectary apparently absent or one or two lobes, sometimes encircling

the ovary, margin entire or undulate and of variable height, 0.2-1. 3 x 0. 1-1.5 mm.

Ovary sub-cylindrical to sub-spherical, 0.7-1. 6 x 0.5-1. 7 mm, sparsely to densely

pubescent, hairs hooked to 0.1 mm long, on upper half of ovary or the uppermost

portion of the ovary below the base of the style; style 0.5-5. 8 mm long, glabrous to

pubescent at the base, hairs hooked, 0.04-0.2 mm long; stigma 0.3-0.6 mm wide.

Fruit obovate-cylindrical, shortly cylindrical or sub-spherical, 1.3-3 x 1-3 mm long;

operculum 0.5-1 mm long, glabrous or pubescent in upper parts, hairs medium to

high density, hooked, white, 0.04-0.13 mm long. Seed narrowly ovoid to ovoid,

infrequently slightly sigmoid, ends acute or constricted, 0.3-0. 6 x 0. 1-0.2 mm, light

to dark brown, pattern straight to spiralled and more or less regular.

Distribution. Indonesia (Java, Lesser Sunda Islands, Sulawesi), Timor Leste.

Habitat and ecology^. Primarily lithophytic on limestone hills, rocks and cliffs, usually

in shaded to heavily shaded areas. Also found in roadside cuttings. Collected at 40-

900 m altitude. Flowering and fruiting January-May.

Provisional lUCN consei^ation assessment. Data Deficient (DD). Although this

species is widespread and would normally be considered to be Least Concern there

are surprisingly few recent collections from much of its range, particularly from Java.

It is unclear whether this is due to a lack of collecting activity or whether it really has

declined over much of its range. This will need to be investigated before a realistic

conservation assessment can be made.

Additional specimens examined. INDONESIA: Bali: Singaraja, Gitgit Waterfall, 5 Apr 1936,

van Steenis, C.G.G.J. 7788 (BO); Gunung Kelatakan, 2 Aug 1918, Maier Sarip, R. 162

(BO). Java: s.l., 8 May 1931, Clason, E.W. C99 (L); Zollinger, H. 2002 (S). Jawa Timur:

Surabaya, Mantup, Coster, C. s.n. (BO); Gunung Wilis, 1914, Backer, C.A.B. 11469 (BO);

Besuki, Gunung Ringit, Kampong Agoeng, 8 Mar 1940, Buwalda, P. 7542 (BO); Besuki, Jang

Plateau, Djeloewang Ravine, 18 Jul 1938-19 Jul 1938, van Steenis, C.G.G.J. 11079 (BO);

Besuki, Puger, 28 Feb 1940, Buwalda, P. 7225 (BO, L); Nusa Baron, 21 Feb 1845, Zollinger,

H. 2612 (BO); Pasuman, 1 8 Mar 1929, Backer, C.A.B. 36621 (L); Ponorogo to Soemoroto, Jan

1918, Beiimee, J.G.B. 1328 (BO); Soekapoera, Gorge of Oemboelan, 12 Apr 1925, Jeswiet,

J. 670 (WAG). Jawa Tengah: Gunung Lawoe, 1913, Backer, C.A.B. 6607 (L); ibidem, 1913,

Backer, C.A.B. 6687 (BO); Yogyakarta, Gunung Kidoel, 1912, Backer, C.A.B. 2789 (BO, L);

Yogyakarta, Unknown (ex Herb. Lugd. Batav.) s.n. (L); Yogyakarta, Prambanan, Feb 1912,

Ridley, H.N. s.n. (K). Jawa Barat: Pameungpeuk, Goenoeng Karikil, Jan 1933, Jacobson, E.

199 (BO). Nusa Tenggara Barat: Lombok, Sambelia, Gunung Rinjani, 17 Feb 1998, de Wilde,

Revision of Epithema

187

W.JJ.O. & de Wilde-Duyjjes, B.E.E. 21940 (E, L); ibidem, 2 May \96\, Kosterincms, AJ.G.H.

18629 (K, L); Western Sumbawa, Batudulang, Mt Batulante, 10 Apr 1961, Kostermans,

A.J.G.H. 18053B (K); Western Sumbawa, Olat seli, Pemek, 17 May 1961, Kuswata 213 (BO);

Western Sumbawa, Sumbawa Besar, 29 Mar 2004, Hoover, W.S, etal. Dedenl 51 (US); Western

Sumbawa, Sumbawa Besar, Semongkat Atas, 26 Mai* 2004, Hoover, W.S. et al. HW11417 (BO,

US); ibidem, 1 May 1961, Kuswata 105 (BO, L); ibidem, 2 May 1961, Kuswata 121 (K, L).

Nusa Tenggara Timur: Flores, Loeters, JJ. 1625 (L); Flores, Lempe to Waewako, 26 Mar

1974, Schmutz, E. 3623 (L); Flores, Waewako, 21 Feb 1978, Schmutz, E. 4027 (L); Western

Flores, Manggarai, 18 Feb 1979, Schmutz, E. 4345 (L); ibidem, 6 Feb 1981, Schmutz, E. 4732

(L); Timor, Naukae, 29 Jan 1971, Kooy, C.W. 789 (L); Timor, Nikiniki, 20 Feb 1966, Kooy

C.W. 403 (L). Sulawesi Selatan: Bulukumba, Malino, 2 Apr 1922, Bunnemeijer, H.A.B. 10713

(BO, L); South ibidem, 8 Apr 1921, Bunnemeijer, H.A.B. 10890 (BO); Bulukumba, Malino,

Bulutana, 10 Feb 2000, Mendum, M. et al. 0034 (E, SING); Maros, 23 Jan 1998, de Wilde,

W.J.J.O. & de Wilde-Duyfjes, B.E.E. 21918 (E, L).

Notes. Hilliard & Burtt( 1997) descrihedthQ variety Epithema horsfieldiiYar. epiphyllum

which they distinguished from the type variety by the position of the inflorescences. In

Epithema horsfieldii var. horsfieldii they usually arise from the base of the leaf blade

and in E. horsfieldii var. epiphyllum tfoin the midrib, up to 2.3 cm from the base of

the blade. Occasionally the inflorescences of the plants placed in Epithema horsfieldii

var. horsfieldii also arise from the lower midrib but these specimens are more densely

villous inside the corolla than the material placed in E. horsfieldii var. epiphyllum. In

addition the material assigned to Epithema horsfieldii var. horsfieldii is generally more

pubescent than E. horsfieldii var. epiphyllum and has larger and more cucullate bracts.

However, specimens with the morphology of the type variety were found disjunctly in

Java and Timor and that of Epithema horsfieldii var. epiphyllum in Lombok, Sumbawa

and Flores. This would be biogeographically unlikely. The material tfom Sulawesi

is rather different again as much of the material has inflorescences arising in a slight

groove at the base of the leaf blade which is not found in the two existing varieties

whereas other specimens are more similar to the type variety. What is clear is that the

patterns of variation require considerably more study and that if varieties are to be

reeognised they should only be so after more collecting from tliroughout the range of

the species (but particularly in Timor) and after a more thorough understanding of the

variation.

6. Epithema involucratum (Roxb.) B.L.Burtt, Notes Roy. Bot. Gard. Edinburgh 22:

308 (1958). - Gratiola involucrata Roxb., [Hort. Bengal. 80 (1810), nom. nud.] FI.

Indica ed. 1, 138 (1820). - Epithema roxburghii A.DC., Prod. 9: 279. (1845), nom.

superfl. - TYPE: “Moluccas”, Roxburgh s.n. (lectotype PDA, designated here). (Fig.

Aikinia brunonis Wall., PI. Asiat. Rar. 3: 66 (1832); G.Don, Gen. Hist. 4: 665 (1838). -

Epithema brunonis (Wall.) Decne., Nouv. Ann. Mus. Hist. Nat. 3: 404 (1834); Decne.,

Herbarii Timorensis Descriptio 76 (1835); A.DC., Prodr. 9: 279 (1845); Clarke in

188

Card. Bull. Singapore 67(1) 2015

Fig. 7. Distribution of Epithema involucratum (Roxb.) B.L.Burtt (■), Epithema longipetiolatum

(Merr.) Hilliard & B.L.Burtt (A), Epithema longitubum Hilliard & B.L.Burtt (•).

A.DC. & C.DC., Monogr. Phan. 5(1): 179 (1883). - TYPE: Indonesia, East Nusa

Tenggara, Timor, Coepang, Apr 1803, Brown, R. s.n. (lectotype BM, designated here).

Epithema brunonis var. violaceum C.B. Clarke in A.DC. & C.DC., Monogr. Phan. 5(1):

179 {ISS3). - Epithema violaceum Span., Linnaea 15: 330 (1841), nom. mid. - TYPE:

Timor Leste, Spanoghe 58 (holotype P; isotype L).

Herb 4.5-20 cm high, caulescent, indumentum strigose to hispid, hairs sparse to

medium density, to 0.3 mm long on the stem, petioles, peduncles, lower bract, and

the outside of the calyx, to 0.8 mm long on the upper and lower surface of the leaves,

indumentum of lower bract, pedicels and the outside of the calyx also of minute hairs;

stem 1-3 mm wide with 2 to 3 nodes, intemodes 1 .3-3 cm long. Leaves membranous

and petiolate, upper leaves opposite; petioles 0.2^. 5 cm long; blades 1.3-6 x 1-5.4

cm, ovate or cordate, rarely almost oblong symmetrical or barely asymmetrical, apex

rounded to truncate, more rarely broadly acute, base cordate or truncate, inserted evenly

on petiole or not, margin entire or with few, minute teeth, undulate. Inflorescences up

to 5 per plant, tenninal, in lower leaf axils or on base of petioles; peduncles 1 .3-5.5

cm long; bracts cucullate but not always enclosing entire inflorescence, 7-17 x 4-10

mm, margin variable, almost entire to dentate, upper surface sub-glabrous, hairs to 0.2

mm long, placed primarily towards margins of the bract; pedicels 0.9-3. 5 mm long,

indumentum may include strigose hairs to 0.5 mm long. Calyx 2. 9^.4 mm long, tube

1.7-2. 5 X 1-2.5 mm, lobes 1-2.1 x 0.6-1. 9 mm; glabrous inside. Corolla light blue

or bluish to dark violet, tube may be white, 9-11 miu long; tube 5-6 x 1.5-2 mm;

lobes 4-5 X 3-A mm, almost as long as the tube; glabrous outside and inside. Stamen

Revision of Epithema

189

filaments 1 .2 mm long; anther dimensions unknown; staminodes 0.7 mm long. Nectary

of two discrete lobes, 0.6 x 0.6 mm. Ovary cylindrical or sub-cylindrical, 0.8-1 x 0.7

mm, pubescent, hairs dense, hooked, white, to 0. 1 mm long on upper portion of ovary;

style 3.5 mm long, glabrous or sub-glabrous, hairs, when present, hooked and on lower

style; stigma to 0.5 mm wide. Fruit sub-cylindrical to sub-spherical, 1.7-1. 9 x 1.7

mm; operculum 0 . 6 - 0. 8 mm long, sparsely to densely pubescent, hairs hooked, to 0.1

mm long, on entire operculum. Seed fusiform to broadly oval, 0.3-0. 5 x 0. 1-0.2 mm;

pattern straight to spiralled and even.

Distribution. Indonesia (Sulawesi, Timor, Maluku), Timor Leste.

Habitat and ecology. In shaded areas. Highly likely to be lithophytic on limestone and

found in humid places and/or near water.

Provisional lUCN conservation assessment. Data Deficient (DD). The known

localities of this species suggest a fairly widespread distribution. However, it has not

been collected often, many of the collections are fairly old and the exact localities of

some of these older specimens are not known. We, therefore, assign it the status of

Data Deficient until more infomiation becomes available.

Additional specimens examined. INDONESIA: Maluku: Babar Islands, Pulau Weten, 28 Feb

1956, van Borssum Waalkes, J. 3025 (BO, L). Nusa Tenggara Timur: Sumba, Laora, 10 Nov

1925, Iboet 221 (BO); Timor, Spanoghe, J.B. 226 (L); ibidem, Unknown (probably Spanoghe)

226 (L). Sulawesi Tenggarah: Pulau Butung, Baoe Baoe, 20 Feb 1929, Kjellberg, G.K. 314

(BO, S); ibidem, 20 Feb 1929, Kjellberg, G.K. 315 (S).

Notes. key macroscopic features distinguishing Epithema involucratum from

other species are the cordate to ovate leaves with clearly rounded leaf tips and that

it is one of the few species that is glabrous inside the corolla. Epithema longitubum

and some specimens of E. ceylanicum are similar to E. involucratum. The Epithema

ceylanicum specimens can be distinguished from E. involucratum by the presence

of an indumentum inside the corolla, smaller inflorescences and shorter corolla

length. Epithema longitubum can be distinguished by the often longer corolla length

and presence of an indumentum in the corolla. While there were not many corollas

available to examine, it would seem that the lobes of E. involucratum are quite large

in comparison to other species.

Van Borssum Waalkes 3025 (L) from Maluku is rather intennediate between

Epithema involucratum and E. longitubum. It is included in Epithema involucratum

on the basis of its leaf shape, absence of hair in the corolla and the lack of an upper,

solitary leaf It does have medium to high density, strigose hairs on the inner side of the

calyx lobes which is unusual for either species. Its inclusion in Epithema involucratum

significantly extends the known distribution of E. involucratum, as it would also have

done for E. longitubum if it were included there.

Kjellberg 3 14 (S) from Sulawesi also extends the range of Epithema involucratum

although, while most of its characters are typical for E. involucratum, it has hooked

hairs at medium density on the ovary which is unusual for this species.

190

Card. Bull. Singapore 67(1) 2015

Epithema brunonis var. violaceum is included in synonymy of E. involucratum.

Clarke ( 1 883) used depth of flower colour, length of the corolla and the leaf pubescence

to delineate it from the type variety. The flower colour and leaf pubescence is not

different from that seen in other E. involucratum specimens and the corolla length,

from material that Clarke himself examined, is 9 mm not 12 mm as Clarke stated when

differentiating the variety. The key differences are nominal; the shape of the base of

a couple of leaves differ slightly and the peduncles can be up to 15 mm longer than

in typical E. involucratum. Epithema brunonis var. violaceum, however, is within the

range of variation for^. involucratum.

In the protologue of Aikinia brunonis there is no indication of a collector of

the specimen used to make the description, only that it is based on a collection from

Coepang (now called Kupang) in Timor which was flowering and fruiting in April

1803. There is a Robert Brown collection from Coepang collected in April 1803 in the

BM. This specimen is lectotypified here.

7. Epithema longipetiolatum (Merr.) Hilliard & B.L.Burtt, Edinburgh J. Bot. 54: 112

(1997). - Epithema brunonis var. longipetiolatum Mem, Philipp. Joum. Sci., C. 11:

313 (191 6). - TYPE: Indonesia, Maluku, Ambon, Halong, 50-100 m, 26 September

1913, Robinson, C.B. 1727 (lectotype L, designated here; isolectotypes BO, K, US).

(Fig. 7)

Herb 5-30 cm high, acaulescent or caulescent; stem 2-3 mm wide with two nodes,

intemodes 7-10 cm long. Leaves membranous, petiolate, usually solitaiy but upper

leaves may be opposite, opposite leaves are occasionally unequal in size; petioles (0-

)0.5-12(-l 8) cm; blade 3-13.6 x 2.4-10.2 cm, cordate to broadly ovate or occasionally

sub-lanceolate, symmetrical or asymmetrical, apex acute, base sub-cordate or sub-

auriculate to cuneate, inserted evenly on petiole or distinctly offset, margin almost

entire to semulate or denticulate, veins often widely spaced; upper surface glabrous or

sparsely to densely strigose, hairs to 0.4 mm long, with or without a minute hooked

pubescence; lower surface finely pubescent, at medium to high density, surface often

appearing fuzzy. Inflorescences 1-6 per plant; peduncles 0.2-6 cm long, terminal or

in the axils of upper leaves; bracts cucullate but not enclosing the entire inflorescence,

5-12 X 4-9 mm, margin somewhat dentate to dentate and serrate, upper surface

sub-glabrous to sparsely strigose, hairs straight, to 0.2 mm long, denser towards the

margin; pedicels 1-4.2 nun long. Calyx 2. 7-5. 6 mm long; tube 1.5-5 x 1.5-3. 8 mm,

lobes 1.2-2. 7 x 0.6-1. 7 mm; minutely pubescent (often finely) outside, occasionally

strigose, at medium to high density, all hairs hooked or primarily hooked and to 0.25

mm long, hairs on inside of lobe tips to 0.13 mm, occasionally sub-glabrous. Corolla

tube white, lobes lilac or pale lilac, lobes with darker coloured but variable markings,

5.5-10.2 mm long; tube cylindrical, 4.2-6.4 x 1.5-1. 9 rmu, lobes 1.2-3. 8 rmu long;

with a band of villous hairs inside around middle, up to 1 mm wide. Stamens 2. 1-3.1

mm long; filaments 1.5-2. 3 mm long; anthers 0.6-0. 8 mm long; staminodes 1.2-2. 3

mm long. Nectary of two discrete lobes, margin entire or undulate, 0. 5-1.1 x 0.7-1. 7

Revision of Epithema

191

mm. Ovary sub-cylindrical or sub-spherical, 1-1.5 x 0.7-1. 3 mm, densely pilose,

hairs usually straight or occasionally straight and hooked, placed on upper portion of

ovai*y, 0.04-0.08(-0.25) mm long; style 3. 1^,4 mm long, wider at base to 0.33 nuu,

style glabrous or with very few hairs at base, these straight, to 0.17 mm long; stigma

0.4 mm wide. Fruit obovate to sub-cylindrical, 2-A x 2-3.5 mm; operculum 0.6-1 .9

mm, finely setose or pilose, hairs medium to high density, as on ovary. Seed narrowly

to broadly ovoid, 0.3-0. 5 x 0. 1-0.2 mm, light to medium brown, pattern straight to

partially spiralled, often with thick, clearly defined walls.

Distribution. Indonesia (Ambon, Bum, Seram, Am, Sulawesi, Papua).

Habitat and ecology. Lithophytic on limestone. In humid, shaded places, often on

rocks in stream beds at 0-1770 m altitude. Flowering and fmiting throughout the year.

Provisional lUCN conseiyation assessment. Least Concern (LC). This species

is widespread. However, as it fi'equently occurs on karst limestone, which is often

conuuercially exploited, its status should be monitored.

Additional specimens examined. INDONESIA; Maluku: Am Islands, Pulau Kobroor,

Namadoeboele, 1 0 Jun 1938, Biiwalda, P. 5202 (BO); Am Islands, Pulau Wokam, Dosinamalaoe,

1 Jun 1938, Buwalda, P. 5110 (A, BO, K, L); Bum, NW Bum, Bara, 5 Dec 1984, van Balgooy,

M.M.J. 5066 (BO, L); Buai, NW Bum, Bara, Waeduna River, Nooteboom, H.P. 5217 (L);

ibidem, 25 Nov 1984, van Balgooy, M.M.J. 4889 (E, L); Ambon, Boerlage, J.G. 528 (BO);

ibidem, Teijsmann, J.E. s.n. (BO); Seram, 9 Jan 1938, Eyma, P.J. 25 74 (BO, L); Seram,

Kampong Kwaos, Gunung Toenlean, 8 Sep 1938, Buwalda, P. 6006 (K, L); Seram, Kampong

Selagor, Gunung Selagor, 26 Aug 1938, Buwalda, P 5748 (BO, L); Seram, Kecamatan Seram

Utara, Goa Pohon Damar to Gunung Roihelu, 24 Jan 1985, Kato, M. et al. C.5232 (L); Seram,

Kecamatan Seram Utara, Goa Pohon Damar to Sawai, 27 Jan 1985, Kato, M. et al. C.5728

(L); Seram, Pileana, 28 Oct 1937, Eyma, PJ. 1826 (A, BO, K, L, SING); Seram, 4 Feb 1918,

Kornassi 967 (BO, L); Seram, Manusela National Park, Gunung Binaia, 26 Aug 1987, Argent,

G.C.G. C8754a (E); Seram, Tehoru, Hatumete Pass, 29 Nov 1917, Kornassi 621 (BO, L);

Seram, Tehom, Saunulu, Murkele Ridge, 21 Jul 1986, Kato, M. etal. C, 11 441 (L); ibidem, 21

Feb 1985, Kato, M. et al. C.5144 (L); Seram, Taniwel, Between Buria and Wae Mala, 3 Feb

1985, Kato, M. et al. C.6007 (A, L). Papua Barat: Kabupaten Raja Ampat, Waigeo Island,

Tuluk Mayalipit, Warsamdin, 1 May 2008, Okada, H. & Tsukaya, H. OT-48 (BO); Biak, Aet

& Idjan 942 (BO, L); Biak, Gunung Wawah, 22 Jul 1939, Aet & Idjan 32 (A, BO, K, L); Fak-

Fak, Fak-Fak River, 23 Feb 1962, Vink, W. BW12141 (L). Sulawesi Tengah: Soroako, Mt.

Wawonsem, 2 Jul 1 979, Hennipman, E. 610g (A, E, L).

Notes. The specimens of Epithema longipetiolatum with leaves arising from ground

level could be confused with E. rennellense and E. strigosum although there is no

overlap in distribution. Leaves arising from ground level are always present in E.

rennellense and E. strigosum and opposite leaves are never seen in these species. The

leaf margin of Epithema strigosum has large wide teeth (entire to weakly dentate in E.

longipetiolatum). Epithema rennellense has straight and hooked hairs on the ovary and

operculum (usually only straight in E. longipetiolatum).

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Card. Bull. Singapore 67(1) 2015

‘Loiigipetiolatiim’ refers as much to the length of the secondary intemode as

to the petioles, giving the plant an elongate appearance. For example, in Kornassi

967 (L), the secondai*y internodes are a third of the total plant height and equal to the

primary intemode and the upper petiole and blade combined.

Epithema longipetiolatum may have solitary leaves only, opposite leaves only,

or opposite and solitaiy leaves. This does not coirelate with geography and both types

can be found within a population. For example, duplicates of the type collection have

only opposite leaves or only solitary leaves while Argent C8754a (E) from Seram

includes plants that have only opposite leaves or both opposite and solitary leaves.

Some specimens of Epithema longipetiolatum are similar to specimens of E.

henthamii. For example, one duplicate of Eyma 1826 (L), from Seram, has a similar

leaf shape and margin to E. benthamU. We have seen no specimens of Epithema

benthamii from Seram, but E. longipetiolatum and E. benthamU are found in the same

locality in the Waeduna River district on Burn Island. The ovary and operculum hair

of specimens of Epithema longipetiolatum found on Burn Island often have a mixture

of long (to 0.25 mm), almost villous hairs that are straight or straight and hooked like

E. benthamii, in addition to the expected, short hairs. This contrasts with the type

specimen of Epithema longipetiolatum from Ambon, theF^. longipetiolatum specimens

from Seram, Wokam Island and the Aet & Idjan 942 (L) specimen from Biak in West

Papua, which have minute, pilose, straight hairs (rarely with a few hooked hairs) on

the ovary and operculum. It is possible some hybridisation occurs between Epithema

longipetiolatum and E. benthamii in these localities.

The minute hooked pubescence on the pedicels and outer calyx is quite

distinctive, especially for the specimens from Seram. For most specimens of Epithema

longipetiolatum, the indumentum of the upper leaf surface is of long, strigose hairs and

looks somewhat like the indumentum of the upper surface of the leaves of E. benthamii.

Specimens from Bum, however, have a dense, uniform and short indumentum, giving

the leaves a fuzzy look.

8. Epithema longitubum Hilliard & B.L.Burtt, Edinburgh J. Bot. 54: 112 (1997). -

TYPE: Timor, Nasinutan, 11 March 1939, Bloembergen, S. 3458 (holotype L; isotype

BO (image seen)). (Fig. 7)

Herb 7.5-20 cm high, with a distinct (but sometimes short) stem before first leaf; stem

0.6-2. 5 mm wide with two nodes, intemodes 1.5-7 cm long. Leaves membranous,

petiolate, upper leaves opposite or with an uppermost leaf solitary opposite a peduncle;

petiole of lowest leaf 4-6 cm long, petioles of upper leaves 0.5-5. 5 cm; blade 1 .7-11

X 1-7.8 cm, sub-orbicular to broadly ovate or somewhat oblong, symmetrical or not, if

not one side up to 1 .3 times wider than the other, apex rounded to acute, base shallowly

auriculate or cordate to rounded, inserted evenly on the petiole or not, margin almost

entire to sermlate or denticulate, venation sub-opposite towards base, usually alternate

towards leaf tip, widely spaced; upper surface with a longer indumentum of hairs to

1 mm long, with or without an additional shorter pubescence of hooked hairs; lower

Revision of Epithema

193

surface drying pale, indumentum as upper surface but less dense. Inflorescences 2-10

per plant; peduncles 0.5-8 cm long, tenninal or in leaf axils; bracts diamond-shaped

or trullate, cucullate but only enclosing a small portion of the inflorescence, 2-9 x

3-6 nmi, margin roughly entire with occasional dentation or with some crenate and

dentate lobes, indumentum of lower surface variable with hairs to 0.5 mm long; upper

surface sub-glabrous, sparsely and minutely pubescent or sparsely strigose; pedicels

0. 75^.2 mm long. Calyx 2.9-5.2(-7) mm long; tube 1. 4-4.2 x 1.3-2. 5 mm; lobes

1-2.9 X 0.4-1 .3 mm; minutely and sparsely to densely pubescent outside and with

or without larger hairs to 0.7 mm long at medium to high density throughout calyx

or largely on lobes; glabrous to sparsely and minutely pubescent inside on upper part

of lobes. Corolla blue-purple, c. 10-17 mm long; tube cylindrical, 8-12 x 1.3-2 mm,

up to 7 times as long as wide; lobes 2-5 mm long; with band of villous hair inside

in lower part. Stamens 1.8-2. 3 mm long; filaments 1-1.5 mm long; anthers 0.8 mm

long; staminodes 0.6-1 .4 mm long. Nectaiy of one or two lobes, partly to entirely

encircling ovary, 0. 5-0.9 x 1.3-1 .5 mm, margin undulate and of varying height. Ovary

sub-cylindrical or broadly ovoid with a truncate base, 0. 9-1.1 x 0.4— 0.6 mm, densely

pubescent on upper portion, hairs hooked; style 8.8-10 mm long, glabrous, rarely

with few hooked hairs at base. Fruit sub-spherical to cylindrical, 1.3-2. 5 x 1.7-2

mm; operculum 0. 5-0.6 mm, pubescent tliroughout but denser at top, hairs hooked, to

0.08 mm long. Seed ovoid to broadly ovoid, 0.3-0. 5 x 0. 1-0.2 mm, pattern partially

spiralled to spiralled, regular.

Distribution. Indonesia (Timor, Flores).

Habitat and ecology. Lithophytic on limestone in humid, shady places near or beside

streams and cave entrances at 900-1200 m altitude. Flowering and fruiting March and

May.

Provisional lUCN consen’ation assessment. Data Deficient (DD). The most recent

collection of this species is from 1974 and the collection density on both Flores and

Timor is too low to have an accurate idea of the curtent distribution.

Additional specimens examined. INDONESIA: Nusa Tenggara Timur: Flores, 25 Mar 1972,

Verheijen, J.A.J. 3070 (L); ibidem, Mar 1974, Verheijen, J.A.J. 3323 (L); Flores, Ruteng-Mano,

16 May 1973, Schmutz, E.3208 (L); Flores, Ruteng-Reo, 7 Mar 1968, Schmiitz, E.2127 (L);

Timor, 30 Jan 1929, Walsh, M.E. 62 (BO); Timor, Nasinutan, 17 Mar 1939, Bloembergen, S.

3496 (BO, L).

Notes. This species is most similar to Epithema involucratum but the inflorescences,

bract and corollas are usually much larger. Unfortunately the locality information on

specimens of both Epithema longitubum and E. involucratum is poor and, therefore, it

is as yet unclear whether these two species co-occur or not without further field work.

Although nectary characters have not proven useful in Epithema, and relatively

few specimens have been examined for this species, it is interesting to note that the

194

Card. Bull. Singapore 67(1) 2015

specimens from Flores have two nectary lobes and the specimen from Timor has one.

The sampling density, however, is currently too low to make any firm conclusions.

9. Epithema madulidii Hilliard & B.L.Burtt, Edinburgh J. Bot. 54: 112 (1997). -

TYPE: Philippines, Coron Island, Banol, 30 Sep 1993, Madulid, D.A., Agoo, Reynoso

& Puentes, R. 11563 (holotype PNH; isotype E). (Fig. 6)

Herb 13.2-50 cm high, caulescent; stem 1^.2 mm wide with two nodes, intemodes

1.5-11 cm long. Leaves membranous or strongly membranous, lower leaf petiolate,

upper leaves sessile; lower petioles 1-10 cm long, sparsely to densely hairy, often

with two distinct lengths of hair, hairs white, to 1.1 mm long; lower leaf blade 9-13 x

4.2-7.2 cm, upper leaves 2-4.5 x 1.7-3. 5 cm, lower leaf cordate to ovate, base cordate

to rounded, upper leaves ovate to elliptic, all leaves usually asymmetrical, one side

1. 1-1.8 times wider than the other, apex broadly acute to rarely rounded, base cordate

to obtuse, margin sub-entire to sinuate to minutely dentate, inserted evenly on petiole

or not; upper surface sparsely to densely hairy, often with two distinct lengths of hair,

hairs to 0.4 mm long; lower surface similar, hairs to 0.5 mm long, indumentum on veins

often white and to 0.8 mm long. Inflorescences 1-3 per plant, tenninal; peduncles

1.4-8 cm long; bracts whitish at base, upper half violet, cucullate, partially enclosing

inflorescence, 10-25 x 6-15 mm, margin dentate to irregularly dentate, lower surface

sub-glabrous or sparsely strigose, upper surface glabrous; pedicels 1^.2 mm long.

Calyx 6.3-7 imn long; tube 3-5.2 x 2. 1-3.8 nmi, often drying a lighter colour than the

lobes; lobes 1.5-2. 9 x 1,1-3. 8 mm, outside glabrous or with sparse hairs primarily on

lobes, hairs to 0.6 mm long; sub-glabrous or pubescent inside, hairs straight, to 0.1 mm

long, mostly only in lobes, occasionally in the tube. Corolla violet at tips, 5-10.4 mm

long; tube cylindrical, 3. 3-6. 5 x 1 .5-1 .8 mm; lobes 1 .7-4.5 mm long; sub-glabrous to

pubescent on lobes outside, hairs hooked or straight, glabrous inside. Stamens 1.3-1. 5

mm long; filaments 0.6-0. 7 mm long; anthers 0.7-0. 8 mm long; staminodes 0.6-0.75

mm. Nectary of one lobe, encircling half to two-thirds of the ovary, 0.5-0. 9 mm high,

margin entire or almost entire. Ovary sub-spherical, 0.7-1. 1 x 0.5-0. 8 mm, densely

pubescent on upper portion of ovaiy, hairs hooked, white, to 0.04 mm long; style 6.3

mm long, glabrous or occasionally with few straight hairs. Fruit obovate to obovate-

cylindrical, 2.6^ x 2-2.7 imu; operculum 0.6-0. 9 mm, sub-glabrous to densely

pubescent, hairs straight and hooked, to 0.13 mm long. Seed ovoid or with one side

flattened, 0. 5-0.6 x 0. 1-0.2 mm, medium to dark brown, pattern more or less straight

to spiralled, regular, lines of pattern thickened.

Distribution. Philippines (Coron Island).

Habitat and ecology. Found in shallow soil in limestone rock crevices in humid,

shaded areas at c. 50-550 m.

Provisional lUCN conservation assessment. Data Deficient (DD). This species is only

known from two eollections from Coron Island and has both an EOO and AOO that

Revision of Epithema

195

would put it in the Critically Endangered category. However, this limestone island and

neighbouring islands are not well collected and the impact of tourism on the habitat is

not clear.

Additional specimen examined. PHILIPPINES: Palawan: Coron Island, Sep 1922, Lopez, G.

BS41343 (BO, K, L, P, US).

Notes. This species is known from two collections from Coron Island collected in 1922

and 1993. Due to the poor condition of the specimens, particularly the Lopez BS41343

collection, some information such as leaf blade, base and tip shape is assessed from

very few leaves.

The calyx of this species is quite large and the tube dries a very light brown

colour in comparison to the lobes although this is based on a very small sample. The

corolla tube is glabrous, or appears so, and, in combination with the light colour and

large size of the calyx, is useful for identification from dried specimens. It is similar to

the sessile-leaved form of Epithema saxatile but can be distinguished by the complete

lack of hair inside the corolla and the presence of straight hairs on the operculum. In

addition the calyx is larger (6.3-7 mm versus 3.3-5 mm) and there are hairs on the

inside of the calyx lobes (lacking in Epithema saxatile).

10. Epithema membranaceum (King) Kiew, Malayan Naturalist J. 38(3): 36 (1985);

B.L.Burtt, Thai For. Bull. (Bot.) 29: 93 (2001). - Argostemma membranaceum King,

J. Asiat. Soc. Bengal, Pt. 2, 72: 145 (1903); Ridley, FI. Malay Penins. 2: 24 (1923),

- TYPE: Malaysia, Penang, Muka Head, Curtis 955 (lectotype SING, designated by

Kiew (1985)). (Fig. 8, 9)

Herb 10-30 cm high, caulescent, indumentum of minute hooked pubescence and

strigose hairs, usually sub-glabrous or sparse; stem 1.5 mm wide, can be red-brown in

colour, with 2-3 nodes, intemodes 0.6-12 cm long. Leaves often finely membranous,

petiolate, upper leaves opposite, infrequently upper leaves solitary or sub-opposite;

petioles 0.8-14.5 cm long; blade 3.6-11.5 x 3-10.5 cm, broadly ovate or cordate,

occasionally sub-orbicular or almost oblong, symmetrical or barely asymmetrical,

apex acute to rounded, upper portion of leaf sometimes curved, base sub-auriculate

to truncate, inserted evenly on petiole, margin entire to seiTate, crenate or with large,

rounded dentation; upper surface glabrous to sparsely strigose (occasionally to medium

density), hairs to 0.6 mm long; lower surface glabrous, sub-glabrous or covered

in dense, minute hooked pubescence, giving a fuzzy appearance. Inflorescences

(l-)3-9 per plant; peduncles 0.5-17 cm long, primarily from terminal leaf axils,

rarely from the base of the petioles; bracts usually small, obovate, ovate or elliptic,

not enclosing inflorescence, if sub-cucullate then only enclosing a small proportion

of the inflorescence, 2-11 x 3-12 mm, margm entire or with limited dentation,

lower surface sub-glabrous, upper surface glabrous or with sparse hair towards bract

margin; pedicels 1-3.5 mm long, densely and minutely pubescent, with or without

few additional straight, strigose hairs to 0.3 mm long. Calyx 2. 1-5.2 mm long; tube

196

Card. Bull. Singapore 67(1) 2015

Fig. 8. Epithema membranaceum (King) Kiew A. Habit. B. Inflorescence. C. Calyx. D.

Calyx indumentum detail. E. Flower opened out. F. Stigma lateral view. G. Fruit showing

seeds, placenta and operculum. H. Operculum indumentum. I. Seeds. Scale bars: A = 4

cm, B = 5 mm, C = 3 mm, D, H, I = 0.5 mm, E = 4 mm, F = 1 mm, G = 2 mm. Drawn by

Claire Banks from Larsen et al 42245 (A-D, G-I) and Larsen et al. 42312 (E, F).

Revision of Epithema

197

Fig. 9. Distribution of Epithema membranaceum (King) Kiew (•) Epithema sarawakense

Hilliard & B.L.Burtt ( A).

1.25-3.5 X 1.3-2. 3 mm; lobes 0.6-3. 1 x 0.75-2.5 mm; indumentum outside as on

pedicel, sub-glabrous inside, a few hairs in lobes. Corolla colour variable, white, pale

pink, pale violet, pale blue or purple or whitish-purple with purple markings, 3.5-7

mm long; tube 2.5-5 x 2 mm, lobes 1-1.9 mm long; pubescent outside, hairs hooked

and straight to 0.1 mm long, with a band of villous hair inside. Stamen filaments

0.75-0.8 mm long; anther characters unknown; staminodes 0.6 mm long. Nectaiy of

one lobe almost or entirely encircling ovary, 0.75-1 x 2.2 nun. Ovary spherical or sub-

spherical, 0.75-1 X 0.6-1 mm, pubescent, hairs medium to high density, fine, hooked,

rarely with one or two straight hairs, hyaline, on upper portion of ovary; style 2-3.8

mm long, occasionally glabrous, pubescent on lower to mid-style, hairs hooked, to

0.13 imn long; stigma c. 0.4 mm wide. Fruit obovate to cylindrical, 2-3 x 2-3 mm;

operculum 1 mm long, pubescence as for ovary. Seeds narrowly ovoid to ovoid, 0.3-

0.5 X 0. 1-0.2 mm, dark brown, pattern straight to spiralled, regular.

Distribution. Thailand, Peninsular Malaysia, Indonesia (Sumatra).

Habitat and ecology. Primarily on limestone, occasionally granite, in shaded areas of

evergreen forests near waterfalls or streams. Found in lowland dipterocarp forests in

Malaysia at 30-415 m altitude. Flowering and fruiting Febmary, April-December.

Provisional lUCN consefyation assessment. Least Concern (LC). This species is

common and widespread although, as it mostly occurs on limestone and limestone

habitats which are often threatened, its status should be monitored.

198

Card. Bull. Singapore 67(1) 2015

Additional specimens examined. THAILAND: Chachoeiigsao: Kliao Tak Groep, 6 Nov 1993,

Larsen, K. etal. 44271 (AAU), Nakhon Si Thammarat: Klia Noni, Khuan Thong, Khao Krod

Cave, 20 Dec 2006, Pooma, R. et al. 6486 (E); Nop Phitum, Khao Luang National Park, Khao

Luang, 25 Oct 1991, Larsen, K. et al. 42584 (AALf); ibidem, 25 Oct 1991, Larsen, K. et al.

42579 (AAU). Narathiwat: 21 Dec 1968, Phusomsaeng, S. etal. 19 (BKF, E); Bacho National

Park, 11 Oct 1991, Larsen, K. et al. 42212 (AAU, P); Sungai Padi, Chatvarin Falls, 18 Oct

1970-19 Oct 1970, Charoenphol, C. et al 3976 (AAU, E, K, L); Pattani, Kliao Chai Son, 23

Dec 2006, Pooma, R. et al 6578 (E); Pattani, Sai Khao Waterfall, 9 Oct 1991, Larsen, K. et

al. 42245 (AAU); ibidem, 20 Dec 1972, Santisuk, T. & B.N. 451 (BKF). Phatthalung: Khao

Chai Son, Road to Than Nam Yen, 23 Dec 2006, Pooma, R. et al. 6579 (BKF); Kao Ok Thalu,

21 Apr 1928, Kerr, A.F.G. 15775 (K). Songkhla: Kliao Chang Lon, 24 Jul 1928, Kerr, A.F.G.

15893 (K); Saba Yoi, Thani Ru Nok Sak (Cave), 21 Oct 1991 , Larsen, K. et al. 42471 (AAU,

BKF); Tam ta Lord, 25 Nov 1990, Larsen, K. et al. 41713 (AAU). Surat Thani: Khan Thuli,

6 Sep 1931, Put 4118 (K). Yala: Bannang Sata, 26 Nov 1961, Suvanakoses, P. 1727 (BKF, K,

L, P); Nam Tok Than To Falls, 10 Dec 1966, Sangkhacliand, B. 1405 (BKF); Tahndo, Tahndo

Falls Forest Park, 1 2 Nov 1 986, Maxw>ell, J.F 86-893 (L, PSU); Than To, Bang Lang National

Park, 17 Jul 2004, Pooma, R. et al. 4298 (BKF).

MALAYSIA: Peninsular Malaysia: Kedah: Baling, 24 Apr 1987, Stone, B.C. et al. 870424-

5/1 (KLU); Kelantan: Gua Musang, 11 Aug 1971, Chin, S.C. 1404 (KLU); ibidem, 1 Nov

1 967, Shimizu, T. & Stone, B.C. 74427 (KYO); Kuala Beds, 15 Feb 2003, Kiew, R. RK5254 (E,

SING). Pahang: Bentung, Bukit Chintamani, 3 Oct 1931., Henderson, M.R. 25010 (BO, SING);

Gua Tipus, 10 Aug 1929, Henderson, M.R. 22580 (SING); ibidem, 15 Oct 1927, Henderson,

M. R. 19406 (BO); Gunung Senyum, 30 Jul 1929, Henderson, M.R. 22382 (BO, SING); ibidem,

28 Nov 1984, Kiew, R. RK1590A (UPM); Kota Gelanggi, 28 Nov 1984, Kiew, R. RK1584

(UPM); Merapoh, Gua Layang, 13 Aug 1971, Chin, S.C. 1514 (E, K); Taman Negara, Batu

Luas, 21 Oct 1982, Kmv, R. RK1231 (UPM); Taman Negara, Batu Luas, Gua Daun, 1 Oct

1984, Kiew, R. RK1347 (UPM); Taman Negara, Gua Luas, Wilkie, P. FRI52899 (E). Penang:

1822, Wallich, N. 9080 (K). Perak: Grik, Temenggor Dam, 9 Sep 1993, Turner, I.M. & Yong

158 (SING); Grik, Temenggor Dam, Sungei Singor, 8 Nov 1993, Saw, L.G. FR139945 (K).

INDONESIA: Sumatera Utara: Asahan, 12 May 1927-21 May 1927, Bartlett, HH. 7740

(US).

Notes. This species is usually quite delicate and distinctive in its vegetative form.

Usually the lower and middle petioles are 0.7-1. 5 times the length of the leaves

which are all cordate-ovate with sub-cordate to sub-auriculate leaf bases. The petiole/

leaf length ratio decreases up the plant as in all petiolate species of Epithema, but

even the uppemiost petioles often appear long in comparison to other species. The

curved asyimnetric upper leaves of some plants are also seen in Epithema steensii and

occasionally in E. tenerum. While peduncles usually originate in the axils of temiinal

leaves, a small number of specimens also have peduncles arising from the base of the

petioles of the terminal leaves.

Epithema membranaceum can be similar to E. parvibracteatum and more rarely

to E. saxatile. The similarity to Epithema parvibracteatum is discussed under that

species. Label data suggests that Epithema membranaceum co-occurs with E. saxatile

on Gunung Senyum and at Batu Luas, both in Pahang, Peninsular Malaysia, Epithema

membranaceum can be separated from E. saxatile by the size of the bract, pubescence

on the style and shorter hairs on the ovary/operculum.

Revision of Epithema

199

11 . Epithema paimbracteatum Hilliard & B.L.Burtt, Edinburgh J. Bot. 54: 112 (1997).

- TYPE: Malaysia, Selangor, Batu Caves, 60 m, 3 November 1953, Sinclair, J. 7821

(holotype E). (Fig. 10)

Herb 6.5-19.6 cm high, caulescent, indumentum of strigose and setose to hispid hairs,

to 0. 8-1.1 mm long on stem, petioles, upper leaf surfaces and peduncles; stem 0.8-

4.4 mm wide with two nodes, internodes 1.2-3. 5 cm long. Leaves membranous or

strongly membranous, petiolate; petiole of lowest leaf 0.8-5 cm long, upper petioles

0.4-2 cm long, densely hairy; blade of the lowest leaf 2.6-1 1.3 x 1.8-8 cm, upper leaf

blades 2. 3-8. 5 x 1.3-6 cm, leaves frequently asymmetrical, one side up to 1.4 times

wider than the other, apex acute to broadly acute, base shallowly auriculate to obtuse,

inserted evenly on petiole, margin sub-entire to dentate, bidentate or serrate; upper

surface often with two clearly different lengths of hair at medium to high density;

lower surface drying white-green, longest hairs primarily on veins and at margins.

Inflorescences 1-5 per plant; peduncles (0.2-)0.5-10.8 cm long, terminal and in leaf

axils, hispid; bracts small, rarely sub-cucullate or trullate, 1-7 x 2-7 mm, margin often

deeply lobed with 3 lobes, but also dentate; lower surface densely hairy with hairs to

0.6 mm long, upper surface with primarily straight hairs to 0.6 mm long, mostly denser

and longer towards margins; pedicels 0. 8-4.2 mm long, sparsely to densely hairy, hairs

hooked and to 0.2 mm long, also with or without longer straight hairs to 0.6 mm long.

Calyx 3. 5-4. 3 mm; tube 1.5-2. 9 x \ .1-2.1 mm, lobes 0.8 -2.9 x 0.6-1. 9 nun; densely

hispid throughout or mainly on lobes outside, hairs to 0.7 mm long; pubescent inside,

hairs straight, on lobes. Corolla white, 4.2-5.4 mm long; tube cylindrical, 2.9-3. 8 x

1-1.5 mm; lobes 0.8-1. 3 mm long, margin entire; with few hooked hairs inside or

with ring of villous hair. Stamens 1. 1-1.6 mm long; filaments 0.6-1 mm long; anthers

0.5-0. 6 mm long; staminodes 0.75-1 mm long. Nectary 1-3 discrete lobes, if tlnee

the third lobe is small, margin almost entire or undulate, 0.6-1. 3 x 0.3-1 .9 mm. Ovary

spherical or sub-cylindrical, 1-1.3 x 0.9-1 mm, sub-glabrous to densely pubescent

or pilose, hairs straight and/or hooked, dense on upper ovary, to 0.2 nuu long; style

1. 9-3.2 mm long, glabrous or with few straight and/or hooked hairs; stigma c. 0.3 mm

wide. Frw/f obovate to cylindrical to sub-spherical, 1,5-2. 8 x 1.9-2, 5 mm; operculum

0.4-0. 9 mm long, indumentum as on ovary, hairs may be densest at the very top of the

ovary. Seed narrowly to broadly ovoid, 0.3-0. 5 x 0. 1-0.2 mm, light brown to medium

brown, pattern almost straight to spiralled, reasonably regular.

Distribution. Peninsular Malaysia.

Habitat and ecology. On limestone or other rocks, in shade, often in crevices of

limestone, at 60-1 50 m altitude. Flowering and fruiting February, May, July-November.

Provisional lUCN conservation assessment. Least Concern (EC). This species is fairly

widespread in Peninsular Malaysia although, as it mostly occurs on limestone and

limestone habitats which are often threatened, its status should be monitored.

200

Card. Bull. Singapore 67(1) 2015

Fig. 10. Distribution of Epithema parvibracteatum Hilliard & B.L.Burtt (•), Epithema steenisii

Hilliard & B.L.Burtt (A) and Epithema strigosum (C.B. Clarke) Hilliard & B.L.Burtt (■).

Additional specimens examined. MALAYSIA: Peninsular Malaysia: Kelantan: UluKelantan,

Bertram, 30 Jul 1962, Unknown 147 (K, L), Pahang: Kota Gelanggi, Aug 1891, Ridley, H.N.

s.n. (SING); Kuantan, Kuantan-Keniaman Road, 19 Jul 1979, Kochummen, K.M. FR126221

(L); Taman Negara, Batu Subuh, 8 Oct 1984, Kmy R. RK1498 (UPM), Perak: Hulu Perak,

Lenggong, Kpg. Gua Badak, 28 Oct 2008, Imin, K. FRI63208 (SING); Kamuning, Feb 1904,

Ridley, H.N. 11883 (K). Selangor: Kanching, Bukit Anak Taloin, 4 Oct 1969, Stone, B.C. 8830

(KLU); ibidem, 29 May 1970, Stone, B.C. & Mahmud 8400 (KLU). Selangor: Kanching,

Bukit Takun, 27 Sep 1970, Chin, S.C. 392 (KLU); ibidem, 24 Oct 1967, Shimizu, T. & Stone,

B.C. 13739 (KYO).

Notes. The distinguishing characters for this species are a small bract and the dense

and long hair covering, particularly on the peduncles, pedicels and calyx. In addition,

the corolla is quite small, only 4.2-5. 4 mm long. There is considerable variation in the

ovary/operculum hair. This variation was observed both within a single inflorescence

and between specimens. Variation in the type of hair found on the inside of the corolla

is unusual and, apart from Epithema parvibracteum, is only seen in E. horsfieldii and

E. rennellense.

The larger plants are rather similar to Epithema membranaceum and some

also to E. saxatile. The specimens of Epithema parvibracteatum, however, have an

extremely reduced bract and a hispid indumentum on the inflorescence.

12. Epithema philippinum (Hilliard & B.L.Burtt) Bransgrove, stat. nov. - Epithema

strigosum philippinum Hilliard & B.L.Burtt, Edinburgh J. Bot. 54: 113 (1997).

Revision of Epithema

201

- TYPE: Philippines, Mindanao, Surigao, Mt Kabatuan, 83 m, 18 March 1949,

Mendoza, D.R. & Convocar, RR 297 [PNH10434] (holotype PNH; isotype A). (Fig. 6)

Herb 6-13 cm high, usually acaulescent with leaves arising near gi'ound level; stem

1-2.5 mm wide with 0-1 nodes, intemode if present 1.4-2. 3 cm long; stem densely

pubescent, hairs to 0.5 mm long. Leaves strongly membranous, petiolate, with one or

more inflorescences opposite each solitary leaf; petioles 2-20 cm long; blade 3-11

X 2.4-4.4 cm, cordate, symmetrical, apex and tip acute, base cordate to truncate,

inserted evenly on petiole, margin dentate and/or bidentate or crenate; upper surface

weakly to strongly strigose, sometimes with setose hairs, hairs straight, hooked or

a combination of both, to 0.7 mm long; lower surface pubescent, hairs to 0.4 mm

long. Inflorescences 1-2 per plant; peduncles 1.6-10.5 cm long, originating opposite

solitary leaves, strigose and setose, haii's dense, to 0.6 mm long; bracts cucullate but

not enclosing entire inflorescence, 3-8 x 3-8 mm, margin dentate; lower surface

strigose, sometimes with two distinct lengths of hair or layers of hair visible, hairs to

0.4 mm long; upper surface strigose, on entire surface but denser towards margins,

hairs to 0.3 mm long; pedicels 1.2-3. 8 mm long, pubescent or hispid, hairs primarily

straight, to 0.4 mm long. Calyx cylindrical, 3. 5-5. 5 mm long; tube 2.3-3 x 1.5-3 mm;

lobes 1.2-2. 5 x 0.8-1. 3 mm; outside with a minute hooked pubescence and longer,

primarily straight hairs covering the entire calyx or only on lobes, straight hairs 0.2-

0.4 mm long, hooked hairs 0.13-0.3 mm long; hairs on lobes inside. Corolla bluish,

5-6 mm long; tube cylindrical, 3. 7-4.2 x 1.2-3 mm; lobes 1.5-2 mm long; band of

villous hair inside in upper tube. Stamens 1-1.5 mm long; filaments c. 0.7-1 mm

long; anthers c. 0.5 mm long; staminodes 0.7-1 mm long. Nectary of two discrete

lobes nearly encircling ovary, 0.8-1 .5 x 0.7-1. 5 mm. Ovary sub-cylindrical or sub-

spherical, 0.7-1 .3 X 0.7-1 .2 mm, upper part of ovary densely pubescent to pilose, hairs

straight, to 0.1 mm long; style 3. 7-4.6 mm long, glabrous or with few straight hairs at

base; stigma c. 0.4 mm wide. Fruit sub-cylindrical or sub-spherical, 1.9-2. 8 x 2-2.5

mm; operculum 0.6-0. 9 imn, indumentum as on ovaiy. Seed narrowly ovoid to ovoid,

0.4-0. 5 X 0. 1-0.2 mm, medium to dark brown, pattern straight to spiralled, regular,

infrequently splitting and merging.

Distribution. Philippines (Mindanao).

Habitat and ecology. Lithophytic, probably on limestone in humid, shady areas.

Recorded at 83 m altitude. Flowering and fruiting March.

Rrovisional lUCN conservation assessment. Data Deficient (DD). This species is only

known from two quite old collections and the current status of the species is unknown.

The area where these collections were made is still largely forested so the population

may still be in good condition.

Additional specimen examined. PHILIPPINES: Mindanao: Agusan: Jabonga, Kitsarao, 14

Mar 1949, Mendoza, D.R. & Convocar, RR. 226 [PNH10394] (PNH).

202

Card. Bull. Singapore 67(1) 2015

Notes. Epithema philippinum is most similar to E. stiigosum. See discussion under

Epithema strigosum regarding the differences between E. philippinum and E. strigosum.

Plants of Epithema philippinum that only have one leaf opposite one or more

peduncles could also be confused with E. horsfieldii but is most easily distinguished

by the straight hairs on the ovary and operculum in E. philippinum (hooked in E.

horsfieldii).

13. Epithema pusillum (C.B. Clarke) Bransgrove, stat. nov. - Epithema carnosum

var. pusillum C.B. Clarke, Monogr. Phan. 5(1): 178 (1883). - Epithema dentatum

ym. pusillum (C.B.Clarke) Hilliard & B.L.Burtt, Edinburgh J. Bot. 54: 112 (1997). -

TYPE: India, Bombay, Stocks s.n. (holotype K). (Fig. 3).

Herb 4.5-12 cm high, caulescent, indumentum of straight and hooked hairs; stem 1 mm

wide with two nodes 1.2^ cm apart and 1-8 cm to the first node, stem sub-glabrous

to sparsely pubescent, 0.2 mm long. Leaves thinly membranous, petiolate; petiole of

lowest leaf 1-2 cm long, petioles of upper leaves 0.1-1. 8 cm long, pubescent, hairs

to 0.2 mm long; blade of the lowest leaf 4.3-6 x 1 .8^.5 cm, upper leaves 2-6.5 x

1 .2^.8 cm, lowest leaf broadly ovate, upper leaves ovate, all symmetrical, apex acute

to rounded, base sub-cordate to truncate, inserted evenly on petiole, margin serrate;

upper surface weakly strigose, hairs sparse, hyaline and/or white, 0.4 nun long; lower

surface sub-glabrous to sparsely pubescent with primarily hooked hairs to 0,5 mm

long. Inflorescences 1-3 per plant; peduncles 1-2.5 cm long, usually originating from

axils of tenninal leaves, indumentum sparse, strigose, straight hairs to 0.3 mm long, or

with a sparse to dense minute hooked pubescence; bracts small and sub-cucullate to

cucullate but not enclosing entire inflorescence, 4-8 x 4-9 mm, margin sub-entire with

occasional dentation; lower surface glabrous or sub-glabrous; upper surface glabrous;

pedicels 1.9-2. 5 mm long, glabrous or with a minute, sparse pubescence. Calyx 2.1-

4.6 mm long; tube 0.8-2. 5 x 1.7-2. 5; lobes 1-2,1 x 0.7-2. 1 mm; usually glabrous or

with a minute hooked pubescence, sometimes with additional straight and/or hooked

hairs to 0.33 mm long outside; glabrous inside. Corolla white, c. 6 mm long; tube c.

4.6 mm long; lobes c. 1.4 mm long; glabrous outside, with villous band of hair inside

in upper half Stamen and staminode characters unknown. Nectary of one lobe, almost

encircling the ovary, c. 0.8 x 0.8-1. 8 mm, margin entire or sub-entire. Ovary sub-

cylindrical to sub-spherical, 0.5-1 x 0.5-1 mm, glabrous; style c. 4 mm long; stigma

c. 0.5 mm wide. Fruit sub-cylindrical, 1.5-2. 3 x 2. 1-2.3 mm; operculum c. 0.6 mm,

glabrous. Seed elliptic to ftisiform, 0.4-0. 5 x 0. 1-0.2 mm, pattern partially spiralled

to spiralled.

Distribution. India (Maharashtra).

Habitat and ecology. Lithophjdic and epiphytic (on at least Dysoxylum sp.) in humid,

probably shaded, places. Recorded from 739 m altitude. Flowering and fruiting

September.

Revision of Epithema

203

Provisional lUCN conservation assessment. Data Deficient (DD). This species is only

known from tliree collections, none of which were collected recently. The area where

these collections were made has been heavily deforested and a survey of the status of

this species is, therefore, vital.

Additional specimens examined. INDIA: Maharashtra: Ratnagiri, Amboli Ghat, Temple Point,

3 Sep 196^, Kulkami, B.G. 108673 (E); Savantwadi, 30 Sep 197^, Almeida, S.M. s.n. (E).

Notes. Epithema pusillum is easily separated from the species under which it has

been included as a variety, E. dentatum (now E. ceylanicum) and E. carnosum, by its

glabrous ovary. Epithema pusillum is only known from central, western peninsular

India. The specimen from Amboli Ghat is not as fine as the other two specimens seen

here, but the ovary and the fruit, however, are consistently glabrous.

14. Epithema rennellense Hilliard & B.L.Burtt, Edinburgh J. Bot. 54: 112 (1997). -

TYPE: Solomon Islands, Rennell Island, 18 August 1962, Dissing, H. 2738 (holotype

E; isotypes E, C).

Herb 12-27 cm high, often with an acaulescent appearance with first leaves arising

near ground level, indumentum of short pubescent hairs and longer strigose or hispid

hairs, the latter up to 0.8 mm long on petioles, upper leaf surfaces, peduncles, pedicels

and outer calyces; stem 2-4.4 mm wide with one or two nodes, intemodes 3.5-11.7

cm long. Leaves membranous to strongly membranous, petiolate, leaves solitary;

petiole of lowest leaf 7-13 cm long, upper petioles 2.5-11.5 cm long; blade of lowest

leaf 6-11.5 x 4.2-9. 3 cm, upper leaf blades 4-14 x 3.4-10 cm, all blades cordate

to oblong cordate or sub-orbicular, symmetrical or not, if asymmetrical, one side up

to 1 .4 times wider than the other, apex acute to rounded, base shallowly auriculate

to rounded, inserted evenly on petiole or slightly offset, margin dentate, bidentate

or serrate; lower surface light green or drying white-green, often finely pubescent

or hispidulous. Inflorescences 1-2 per plant; peduncles 2-21 cm long, originating

opposite single leaves or occasionally in leaf axil; bracts sub-cucullate to cucullate,

occasionally small, enclosing the entire inflorescence or portion thereof, 3-13 x 4-10

mm, margin entire with occasional crenate or dentate lobes to dentate; upper surface

sub-glabrous to sparsely pubescent, hairs to 0.2 mm long, usually on upper portion of

bract; pedicels 1 .3-5 mm long. Calyx light green, 4. 2-7. 5 mm long, tube (1 .7-)3-4.2 x

(1.5-)2-3 mm, lobes 1.5-5 x 1-1.5 mm; glabrous to sparsely strigose on lobes inside,

hairs straight or sti'aight and hooked, to 0.13 mm long. Corolla white, 10.4-15 mm

long; tube 6-11 x 2-3.3 mm; lobes 3-4.6 mm long, margin uneven/undulate; glabrous

or sub-glabrous or with a band of villous hair inside. Stamens 1. 2-3.1 mm long;

filaments 0.7-2. 1 mm long; anthers 0.5-1 mm long; staminodes 0.8-1. 5 mm long.

Nectary variable, one lobe (not entire) encircling ovary or with two discrete lobes

not encircling ovary, 0.6-1 .6 x 0.7-3. 5 mm, margin entire to undulate. Ovary 1-1 .6 x

0.8-1. 6 mm, sub-glabrous or pubescent at sparse to medium density on upper portion

204

Card. Bull. Singapore 67(1) 2015

of ovary, hairs hooked or both straight and hooked, 0.04-0.3 mm long; style to 8.8

mm long, glabrous or with few hairs at base, hairs straight and hooked; stigma c. 0.5

mm wide. Fri//r obovate, cylindrical or sub-cylindrical, 2.5^ x 1-3.3 mm; operculum

0.7-1. 3 iimi, indumentum as ovary, but often on top of operculum or densest on top of

operculum. Seed nan'owly ovoid to ovoid, 0.4-0. 8 x 0. 1-0.2 mm, light to dark brown,

pattem straight to spiralled, reasonably regular, walls somewhat thickened and rigid.

Distribution. Solomon Islands (Rennell Island).

Habitat and ecology. Growing on well-drained hillsides over limestone (Rennell

Island is a limestone, coral atoll) at 20-90 m altitude.

Provisional lUCN conservation assessment. Least Concern (LC). Although this species

has not often been collected, and it is endemic to an island that is only 660 km2, the

island is still largely forested and there is little evidence that the the population is under

serious threat.

Additional specimens examined. SOLOMON ISLANDS: s.l. 1894, Officers of H.M.S.

“Penguin ” (K). Rennell Island: 21 Aug 1962, Dissing, H. 2798 (C); 6 May 1968, Sirute’e, B.

etal. BSIP9621 (K, L, SING); Hutuna, 25 Mar 1965, Wolff, T. 3004 (C); Matangi Area, 1 7 May

1969, Gafiii, I.H. & collectors BSIP14759 (K, L, SING); Nuipani, 14 Mar 1965, Wolff, T. 3025

(C); Tuhungganggo, 24 May 1969, Gafui, I.H. & collectors BSIP 14763 (K, L, SING).

Notes. This species is similar to Epithema longipetiolatum and E. strigosum in the

leaves arising at ground level. Epithema rennellense is also somewhat similar to E.

dolichopodum but differs in peduncle length/plant height ratio, leaf shape and leaf

margin, the length of the calyx and the corolla, the ovary /operculum hair and the size

of the seed. The variation in the shape of the hair on both the ovary and the operculum,

between hooked and both straight and hooked, can be seen on different fruit from the

one plant. The variation in corolla hair is unusual and is seen in few other species,

none of which are acaulescent species. Epithema rennellense has the largest seed of all

species of Epithema and, along with E. longitubum, the largest flowers.

15. Epithema sarawakense Hilliard & B.L.Burtt, Edinburgh J. Bot. 54: 113 (1997). -

TYPE: Malaysia, Sarawak, Fifth Division, Sungai Medalam, Gunung Buda, 20 June

1975, Burtt, B.L. 8336 (holotype E; isotype SAR). (Fig. 9)

Herb 25^0 cm high, sprawling or caulescent, indumentum of pubescent or strigose

and setose hairs, to 0.7 mm long on the lower leaf surface and calyx, to 1 mm long

on the upper leaf surface; stem 1.2-5 mm wide with 2 to 4 nodes, intemodes 1-8 cm

apart long. Leaves membranous to strongly membranous, petiolate, primarily solitary,

rarely with one or two sets of opposite leaves per plant; petioles 1-7 cm long; blade

2.2-17 X 1.4-13.2 cm, often cordate, but can be sub-orbicular, elliptic or sub-oblong.

Revision of Epithema

205

symmetrical or not, if asymmetrical, one side up to 1.5 times wider than the other, apex

acute to rounded, occasionally almost truncate, base usually (sub-)cordate but also to

truncate, inserted evenly on petiole or not, margin sub-entire to dentate or bidentate;

upper surface sparsely to densely strigose or villous, occasionally sub-glabrous;

lower surface may dry grey-green, glabrous to villous. Inflorescences 3-11 per plant;

peduncles 0.5-16 cm long, 1-5 arising in sequence opposite solitary leaves or rarely

in the axils of upper opposite leaves; bracts cucullate, usually completely enclosing

inflorescence, 10-32 x 6-12 mm, margin dentate or irregularly dentate; lower surface

with hairs to 0.5 mm long; upper surface often glabrous; pedicels 1.3^. 6 mm long,

often glabrous. Calyx 3. 3-6. 6 mm long; tube 1.5-3. 5 x 1-3.1 mm, lobes 1.5-3. 8 x

0.6-2. 3 mm; frequently sub-glabrous to finely pubescent outside (short, hooked hairs),

occasionally with sparse strigose hairs; inside glabrous or with few straight hairs, 0.1

mm long, in tips of lobes. Corolla white and pale blue to purple, pink, or mauve, may

have darker markings on the lobes, 8.7-11 mm long; tube 6-7 x 1.5-2. 5 mm, lobes

2. 7^. 2 X 1. 5-2.1 mm, margin slightly fimbriate. Stamens 3 mm long; filaments 1.5

mm long; anthers 1.5 mm long; staminodes 1.5-1. 8 mm long. Nectary a single lobe,

partially to completely encircling ovary, 1-1.3 x 1.2-2. 5 mm, margin almost entire to

undulate, occasionally almost dividing into smaller lobes. Ovary sub-cylindrical, 0.8-

1 X 0.7-0.9 mm, usually glabrous, occasionally ovary pubescent with hooked hairs on

upper ovary, hairs to 0.1 mm long; style c. 6 mm long, slightly wider at base, glabrous;

stigma c. 0.4 mm wide. Fruit obovate-cylindrical, 2-3 x 1 mm; operculum glabrous,

occasionally sub-glabrous, indumentum as on ovary. Seed narrowly ovoid to ovoid or

somewhat sigmoid, 0.4-0.5 x 0. 1-0.2 mm, light brown to dark brown, pattern straight

or almost so, occasionally partially spiralled to spiralled.

Distribution. Malaysia (Sabah, Sarawak), Indonesia (Kalimantan, Sumatra: Pulau

Enggano).

Habitat and ecology. Lithophytic on limestone (but substrate not always reported).

Humid, shady areas, beside streams, limestone outcrops, cave entrances. Occasionally

found on wood or on soil over rock. At 30-1250 m altitude. Flowering and fmiting

year round.

Provisional lUCN conservation assessment. Least Concern (LC). This species is

widespread with many of the collections having been made in the protected areas of

Gunung Mulu National Park and Niah National Park.

Additional specimens examined. MALAYSIA; Borneo: Sabah: Tambunan, Crocker Range,

Tambunan/Ranau Road, 5 Mar 1995, Sugau, J. JBS96 (E). Sarawak: 1st Division, Jambusan,

Gunung Batu , 6 Oct 1 977, Martin^ P.J. S. 39273 (E, K, L); Baram District, Jun 1 894, Haviland,

G.D. & Hose, C. 3525 R (K); Baram District, Batu Gading, 5 Jan 1965, van Niel, J.P. 3554

(L); Baram District, Gunung Mulu National Park, Bukit Binarat, 28 Apr 1 985, Moktar, A. et al.

S. 49467 (E, K); Baram District, Gunung Mulu National Park, Deer Cave, 16 i 1978, Hansen,

C. 17 (C, E, SAR); ibidem, 16 Jan 1978, Hansen, C. 17 (C, E); ibidem, 2 Oct 2007, Julia, S. et

206

Card. Bull. Singapore 67(1) 2015

al. S.99303 (E); ibidem, 5 May 1978, Kiew, R. RK525 (E, UPM); ibidem, 1 6 Jan 1978, Nielsen,

I. 17 (E); ibidem, 23 Jul 1987, Prirnack, R.B. S. 42 402 (K, L); Baram District, Gunung Mulu

National Park, Gua Rusa, 30 Oct 1977, Argent, G.C.G. & Kerby, R. 621 (E); Baram District,

Gunung Mulu National Park, 22 Mar 1964, Hotta, M. 15281 (E, KYO, L); ibidem, 14 Mar

1964, Hotta, M. 14402 (E, KYO); ibidem, 19 Mar 1990, Yil PC. & Abu Talib S.58618 (SAR);

Miri, Mulu National Park, en route from HQ to Deer Cave, 20 Dec 1 999, Irnaichi, R. et al. 30

(SAR); Miri, Mulu National Park, Royal Mulu Resort, 6 Oct 2001, Julaihi S. 86868 (SAR);

Miri, Mulu National Park, Simons Cave, 27 Apr 1997, Haegens, RM.A.P. & Klazenga, N. 515

(SAR); Baram District, Gunung Mulu National Park, Melinau Gorge, 23 Jun 1962, Burtt, B.L.

& Woods, P.J.B. B. 2230 (E); ibidem, 20 Feb 1978, Nielsen, 1. 416 (E); Baram District Gunung

Mulu National Park, Pala River, 14 Nov 1977, Argent, G.C.G. & Collins, M. 733 (E); Baram

District, Ulu Melinau, Aug 1958, Ashton, PS. A. 339 (K); ibidem, Aug 1958, Ashton, PS. A.

340 (K); Baram District, Ulu Sg. Tutoh, Batu Kalulong, 13 Apr 1997, Julaihi et al. S. 76938

(SAR); Batu Niah, Nov 1 932, Synge, PM. S.565 (K); ibidem, Nov 1 932, Synge, PM. S.634 (K);

Batu Niah, Gunung Subis, 28 Nov 1966, Anderson, J.A.R. et al. S. 26090 (A, E, K, L, SING);

ibidem, Jan 1961, Mohidin S.21608 (K); ibidem, 19 Aug 2002, Sabli, J. et al. S. 89048 (SAR);

Batu Niah, Niah Cave Park, 4 Jun 1962, Burtt, B.L. & Woods, P.J.B. B.2003 (E); ibidem, 3

Jun 1962, Chew, W.-L. CWL.293 (K, L, SAR, SING); ibidem, 23 Dec 1999, Irnaichi, R. et al.

38 (SAR); ibidem, 20 Aug 2006, Prieditis, N s.n. (E); ibidem, 20 Aug 2006, Prieditis, N. s.n.

(E); ibidem, 18 Apr 1978, Stone, B.C. 13730 (K, KLU, L); Belaga District, Bukit Merirai,

Gua Pak Danum, 12 Jul 2005, Leong, P. et al. PL320 (SING); Belaga District, Ulu Merirai, 9

Jul 2005, Leong, P. et al. PL215 (SING); Belaga District Ulu Merirai, Gua Tiang, 6 Jul 2005,

Julia, S. et al. S.93376 (SING); Bintulu, Ulu Sg. Kakus, Bukit Sarang, 14 Mar 1965, Anderson,

J. A.R. S. 2095 7 (A, K); Binmlu, Ulu Sg. Kakus, Bukit Sarang, Batu Anyi, 5 Oct 2004, Julia,

S. et al. S. 9475 7 (SAR, SING); Buseau, 1890, Haviland, G.D. s.n. (K); Kapit, Melmau, 16

Jul 1961, Anderson, J.A.R. & Keng, H. KlOO (SAR); Kapit, Ulu Melmau, Gunung Api, 1971,

Anderson, J.A.R. S.31770 (E, K, L, SING); ibidem, 10 Sep 1970, Chai, P. S.30372 (A, BO, K,

L, SING); ibidem, 1970, Lehmann, P.F. PPL. 596/S, 30372 (E); ibidem, 7 Sep 1970, Lehmann,

PE S.30091 (E, K, SAR); ibidem, 29 Mar 1990, Yii, PC. & Abu Talib S.58841 (SAR); ibidem,

29 Mar 1990, Yii, PC. & Abu Talib S.58841 (SAR); Lobang Rusa, 9 Jun 1975, Burtt, B.L. B.

8227 (E, SAR); Lobang Rusa, Sungei Melinau Paku, 9 Feb 1966, Chew, W.-L. CWLlOl 7 (A, K,

L, SING); Marudi District The Pinnacles, Gunung Api, 17 Jun 1995, Beaman, J.H. 11727 (K);

Miri, Gunong Mulu National park. Hidden Valley, 5 Apr Argent, G.C.G. et al. 905b (E);

Niah, 7 Oct 1954, Ahmad I (SING); Niah National Park, 12 Dec 1981, Rogstad, S.H. 728 (A).

INDONESIA: Sumatra: Bengkulu: Pulau Enggano, Boea-boea, 11 Jun 1936, Lutjeharms,

W.J. 4598 (L); Pulau Enggano, Malakoni, 26 Jun 1936, Liltjehaims, W.J. 5112 (BO, GH, K, L,

P). Kalimantan: Peningin, Jo/zm 1522 (BO).

Notes. Epithema sarawakense is usually quite distinctive with its near complete lack

of opposite leaves and one to five infloresences arising from the base of the solitary

petiole. Epithema sarawakense is often densely hairy, giving the dried leaves a grey-

green colour. There are two forms, however, one of which is glabrous or sub-glabrous.

The glabrous form is found almost exclusively in Gunung Mulu National Park in the

environs of Deer Cave and the base of Gunung Mulu. While there are a small number

of collections of the pubescent fonn from near Deer Cave, it has primarily been

collected from Gunung Api, Gunung Buda, Gunung Benarat, along Sungai Melinau

and in the Melinau Gorge.

Revision of Epithema

207

There are a small number of specimens that show some similarity to Epithema

saxatile. These specimens have some opposite leaves but, in most cases, there are

always solitary leaves with peduncles arising opposite the solitary leaf on the plant.

These opposite leaves are unequal in size while Epithema saxatile has even-sized

opposite leaves. In addition, the range of the corolla length of Epithema saraM^akense

is 8-11 mm while that of E. saxatile is 6. 5-9. 5 mm and the upper o vary/ operculum

of E. sarawakense is glabrous or sub-glabrous (hairs hooked) while the operculum

of E. saxatile is always at least sparsely pubescent (hairs hooked). The specimens of

Epithema sarawakense that are similar to E. saxatile are not segregated geographically

from more typical E. sarawakense.

A small number of specimens from Pulau Enggano in Sumatra are included in

Epithema sarawakense. While the distribution is difficult to explain, there are currently

no characters to separate the specimens from the remainder of the species as found in

Sarawak. They have shorter stems than is usual for Epithema sarawakense, thereby

resembling E. strigosum, but they differ from E. strigosum in that the operculum is

glabrous to sparsely pubescent with hooked hairs rather than dense and with straight

hairs as in E. strigosum.

16 . Epithema saxatile Blume, Bijdr. 738 (1826); Ridley, FI. Malay Penins. 2: 539

(1923); B.L.Burtt, Thai For. Bull. (Bot.) 29: 94 (2001). - TYPE: Indonesia, Java,

Blume, C.L. 1029 (lectotype L [L0003232], designated here; probable isolectotype U

[without number]). (Fig. 11, 12)

Epithema carnosum auct. non Benth.: Barnett, FI. Siam. 3(3): 205 (1962), p.p.

Herb 6-40 cm high, caulescent; stem 2-5 mm wide with 2-5 nodes, intemodes

1.2-10.5 cm long, usually glabrous or sub-glabrous. Leaves membranous to strongly

membranous, petiolate or with sessile upper leaves, rarely with one or two solitary

upper leaves; petiole of lowest leaf 2.5-12 cm long, upper petioles 0-2.5 cm long;

blade oflowest leaf 7-20 x 6.5-12.5 cm, upper leaves 4.5-1 3.5 x 3-8.5 cm, lowest leaf

sub-spherical to broadly ovate, upper leaves variable, often narrowly ovate to almost

oblong, but also cordate, broadly ovate or elliptic, symmetrical or not, if asymmetrical,

one side up to 1.4 times wider than the other, apex acute to broadly acute, rarely

rounded, base cordate to truncate, inserted evenly on petiole to distinctly offset (up to

2 cm), margin sub-entire to dentate, bidentate or serrate (often all or degrees of these

on one plant); upper surface green or dark-green, indumentum occasionally to 1.1 mm

long; lower surface light green, whitish or occasionally purplish. Inflorescences 1-10

per plant; peduncles 2-17 cm long, indumentum occasionally hispid, to 0.7 mm long;

bracts cucullate, usually enclosing the entire inflorescence, 13-35 x 8-12 mm, margin

usually dentate, occasionally sub-entire with one or two teeth or lobes; lower surface

usually sparsely to densely strigose, hairs straight or primarily straight, 0. 3-0.5 mm

long; upper surface glabrous or sub-glabrous, hairs towards margins of bract; pedicels

1.1^.2 mm long. Calyx 3.3-5 mm long; tube 1.6-3. 3 x 1. 5-2.1 mm; lobes 1.5-2. 7 x

208

Card. Bull Singapore 67(1) 2015

Fig. 11. Epithema saxatile Blume. Flowers and cucullate bract. (Photo: David Middleton)

Fig. 12. Distribution of Epithema saxatile Blume (•).

Revision of Epithema

209

0.6-0. 9 nmi, often glabrous at maturity, hairs to 0.5 mm long, glabrous inside. Corolla

white, pink, blue, purple, or purple and white, frequently with highly variable purple

markings on the upper lip, 6. 5-9.5 mm long; tube 5-7 x 1.5-2. 5 mm; lobes 1-3 x

1.7-2. 3 mm; band of villous hair up to 1 mm wide inside. Stamen filaments 1. 3-2.1

mm long; anthers c. 0.6 mm long; staminodes 1. 3-2.1 mm long. Nectary apparently

absent or with one or two lobes partially to entirely encircling ovary, margin undulate,

0.7-1. 3 X 0.6-2 mm. Ovary sub-cylindrical or sub-obovate, 0.6-1. 5 x 0.5-1 mm, very

sparsely to densely pubescent, hairs hooked, on the upper or very upper portion of

ovary, hairs 0.04-0.63 nnu long; style 2.9-5 mm, glabrous or sparsely pubescent on

the lower style; stigma 0.3 mm wide. Fruit cylindrical or obovate, 1.9-2. 6 x 1.7-2. 1

mm; operculum 0.4-0. 7 mm long, pubescence as ovary. Seed narrowly ovoid to ovoid,

0.3-0. 5 X 0. 1-0.2 mm, pattern stright to spiralled, reasonably regular.

Distribution. Myanmar, Thailand, Malaysia (Peninsular Malaysia, Sarawak, Sabah),

Indonesia (Sumatra, Java, Kalimantan, Sulawesi).

Habitat and ecology. Shaded, humid areas, usually on or amongst limestone rocks or

in limestone soil, near, by or in streams or rivers. Also found near or on the walls of

cave mouth entrances. At 0-910 m altitude. Flowering and fruiting hugely variable.

Provisional lUCN conservation assessment. Least Concern (LC). This species is

common and widespread.

Additional specimens examined. MYANMAR: Tenasserim, 1877, Gallatly, G. 1035 (SING);

Tenasserim, Tavoy, Paungdaw, Aug 1 96\, Keenan, J. et al. 798 (E).

THAILAND: Chanthaburi: Nam Tok Phliu National Park, Nam Tok Phliu Waterfall, 30

Aug 1969, Maxwell, J.F s.n. (AAU); Priu Waterfall, 12 Oct 1965, Chermsirivathana, C. 413

(BKF). Chiang Mai: Doi Sutep, 4 Aug 1912, Kerr, A.F.G. 2655 (K); Me Ta Chang, 17 Oct

1922, Ken; A.F.G. 6381 (ABD, E, K). Chon Buri: Bahn Beung District, Ang Chang Nam, 1

Sep 1975, Maxwell, J.F. 75-958 (AAU, BKF, L); Pong NamRawn, 3 Sep 1956, Smitinand, T.

3489 (BKF, E); Sriricha, Chundaten Falls, 1 7 Aug 1 974, Maxwell, J.F. 74-814 (AAU, BK, L),

Chumphon: Bang Son, 11 Sep 1927, Put 1051 (ABD, K). Kanchanaburi: Between Huay

Ban Kao and Kritee, 4 Jul 1973, Geesink, R. & Phengklai, C. 6082 (BKF, E, L); ibidem, 12 Jul

1973, Geesink, R. & Phengklai, C. 6231 (AALT, BKF, C, E, K, L, P). Kanchanaburi: Hindato,

24 Jul 1946, Kosterrnans, A.J.G.H. 1346 (BO, A, L, P); Dongyai, 14 Aug 1971, Phengklai, C.

2933 (BKF); Kliaibuing, CP. BS& 5A 2997 (BKF, E, L); Kin Sayok, 13 Jul \946,Kostermans,

A.J.G.H. 1103 (A, BO, L, P, SING, US); Lai Wo, Toong Yai Naresuan Wildlife Reserve, Ban

Saneh Pawng, 12 Aug 1993, Maxwell, J.F. 93-886 (CMU); Rintin, 31 Jul 1946, Kosterrnans,

A.J.G.H. 1401 (A, BO, L, P, SING, US); Si Sawat, Hua Lum Kao Ngoo, 5 Jul 1 973, Sutheeson,

S. 2489 (BKF); Tawng Pa Poom, Huay Ban Kliao, 12 Jul 1973, Maxwell, J.F 73-285 (AAU).

Khainphaeng Phet: Khlong Lan National Park, 21 Aug 1995, Parnell, J. et al. 95-335 (K).

Krabi: 29 Mar 1930, Kerr, A.F.G. 19385 (K); Ao Luk, 21 Jun 2006, Williams, K. et al. 2033

(A); Nai Chong, 21 Oct 1979, Sutheeson, S. 5004 (BKF). Mae Hong Son: Muang, Nahng

Rawng Falls, 16 Sep 1972, Maxwell, J.F. 72-362 (L). Nakhon Nayok: Nang Rong Falls, 13

Aug 1968, Larsen, K. et al. 3363 (AAU, BKF, C, E, K, L, P, SING). Nakhon Si Thammarat:

Thung Song District, Khao Tham Long, 31 Aug 1982, Shimizu, T. et al. T. 28985 (BKF); Nop

210

Card. Bull. Singapore 67(1) 2015

Phitum, Khao Luang National Park, Krung Ching Falls, 24 Sep 2010, Middleton, D.J. et al.

5524 (E); ibidem, 12 Feb 2005, Williams, K. et al. 1400 (E). Phangnga: Khao Lak, 21 Sep

1963, Smitinand, T & Sleumer, H.O. 1207 (BKF, L). Phangnga: Muang, Suan Somdet, 8 Dec

1999, Wongprasert, T, 9912-30 (BKF); Muang, Tham Pha Phueng, 15 Sep 2010, Middleton,

D.J. etal. 5425 (E); Panga, Sep 1894, Curtis, C. s.n. (SING); ibidem, Aug 1893, Curtis, C. s.n.

(SING); Pulau Tebun, 29 Dec 1918, Nur, M. 3585 (K, SING); Talcuapah, 14 Jul 1972, Larsen,

K. et ai 30967 (AAU, E); Tham Tong Lang, Shimizu, T, et al. Z29151 (BKF). Phatthalung:

Khao Pu-Khao Ya National Park , 24 Sep 1986, Maxwell, J.F. 86-723 (A, BKF, L, PSU).

Phitsanulok: N. Phitsanulok, Thung Salaeng Luang National Park, 22 Jul 1966, Larsen, K.

et al. 722 (AAU, BKF, L); ibidem, 25 Jul 1973, Murata, G. et al. T.17092 (AAU, BKF, KYO,

L); ibidem, Phusomsaeng, S. et al. 83 (BKF); ibidem, 30 Sep 1967, Tagawa, M. et al. T. 11 259

(KYO); Thung Salaeng Luang National Park, Kaeng Sopa Waterfall, 22 Oct 1984, Murata, G.

et al. 38544 (BKF); ibidem, 22 Oct 1984, Murata, G. et al. T. 38467 (BKF); Thung Salaeng

Luang National Park, Kang So Pa Waterfall, 17 Sep 1990, Chantharanothai, P. et al. 90/303

(K); Thung Salaeng Luang National Park, Poi Waterfall, 22 Oct 1984, Murata, G. etal. T.38587

(BKF). Saraburi: Khao Muak Lek, 2 Oct 1963, Bin Rajab, M.K. 714 (KLU). Songkhla:

Boripath Waterfall, 1990, Larsen, K. et al. 41243 (AAU); ibidem, 16 Aug 1984, Maxwell, J.F.

84-55 (A, PSU); Boriphat Falls National Park, 9 Nov 1990, Larsen, K. et al. 41243 (P). Surat

Thani: Ban Kawp Kiep, 5 Aug 1927, Kerr, A.F.G. 13183 (ABD, E, K). Surat Thani: Klilong

Phanom National Park, Middleton, D.J. et al. 4345 (BKF, E); Phanom, Khao Sok National

Park, 6 Sep 2008, Middleton, D.J. et al. 4330 (E); Phanom, Khao Sok National Park, Chong

Lorn, 12 Dec 1979, Shimizu, T. et al. T.27103 (KYO); ibidem, 12 Dec 1979, Shimizu, T. et al.

T.27111 (BKF, KYO); ibidem, 12 Dec 1979, Shimizu, T. etal. T.27100 (KYO); Phanom, Khao

Sok National Park, Sii Ru Cave, 27 Feb 2006, Middleton, D.J. et al. 4055 (BKF, E); Sawng Pi

Nawng, 20 Mar 1927, Kerr, A.F.G. 12385 (K). Satun: Khuan Don, Thale Ban National Park,

Along cliff face from Ton Din Cave, 9 Sep 2010, Middleton, D.J. et al. 5350 (E); Thale Ban

National Park, 27 Aug 1995, Larsen, K. et al. 46055 (AAU); Thung Wa, Than Plew Waterfall,

1 0 Sep 2010, Middleton, D.J. et al 5368 (E). Trang: Ampoe Kao Kao, Kao Chom Lem, 2 Aug

1929, Rabil 318 (K). Trang: Ban Nam Phrai, 17 Dec 1979, Shimizu, T. et al T.27551 (BKF,

KYO, L); ibidem, 17 Dec 1979, Shimizu, T. etal T.27571 (KYO); Chawng, 15 Mar 1959,

Smitinand, T. & Abbe, E.C. 6149 (BKF, K); Huay Yot, Wat Tham Iso, Middleton, D.J. et al.

4428 (E, SING); Lamphura, 15 Nov 1990, Larsen, K. et al. 41413 (AAU, BKF); ibidem, 16

Nov 1990, Larsen, K. et al. 41450 (AAU); Trang, Nam Tai, 1 1 Oct 1970, Charoenphol, C. et

al. 3662 (AAU, E); ibidem, 1 1 Oct 1970, Charoenphol, C. et al. 3661 (AAU, BKF); Trat, Koh

Chang, 7 Sep 1992, Niyomdham, C. 3276 (BKF). Yala: Betong, 27 Aug 1923, Kerr, A.F.G.

7687 (K).

LAOS: Chanipassak: Thorel, C. s.n. (P); ibidem, Thorel, C. s.n. (P).

VIETNAM: Luang, Aug 1 868, Pierre, L. 4542 (P). Dong Nai: Bien Hoa, Cochinchine, Pierre,

J.B.L. 4541 (P); Gia Lai: Dak Doa, Poilane, E. 18108 (P).

MALAYSIA: Sabah: 24 Feb 1985, Lamb, A.L. 8/85 (E); Interior Zone, Batu Punggol, Apr 1987,

Venneulen, J.J. 1188 (L); Kinabatangan District, Kori Timber Camp, 7 Nov 1948, Cuadra, A.

A. 2168 (A, K, KEP, SAN, SING); Kinabatangan District, Northern Tabin Wildlife Reserve

(Near Tabin River), 16 Oct 2000, Poulsen, A.D. et al. 1659 (E); Lahad Datu, Bukit Baturong, 7

Jul 2000, Kiew, R. RK5040 (SING); Lahad Datu, Tabin Wildlife Reserve, 24 Oct 2000, Kiew, R.

RK5120 (SING); ibidem, 25 Oct 2000, Pius, G. etal. SANl 43649 (E, K); Madai Baturong Forest

Reserve, Baturong Hill, 12 Jim 1996, Lim, S.P. et al. LSP738 (SING); Sandakan, Gomantong,

13 Feb 1960, Meijer, W. SAN20756 (A, K, L); Sandakan, Gomantong, Gomantong Caves, 31

Oct 1968, Kokawa, S. & Hotta, M. 551 (KYO); ibidem, 31 Oct 1968, Kokawa, S. & Hotta,

Revision of Epithema

211

M. 557 (KYO); ibidem, 31 Oct 1968, Kokawa, S. & Hotta, M. 565 (KYO); ibidem, 25 Sep

1968-26 Sep 1968, Ogata, K. 10975 (KYO, L); Sandakan, Gomantong, Gomantong Hill, 20

Sep 1970, Banggilon, K. SAN66579 (K, L); ibidem, 28 Apr 1996, Lirn, S.P et ai 580 (SING);

ibidem, 26 Apr 84, Sands, MJ.S. & Young, R.G.N. 3936 (E, K); Sandakan, Hutan Simpan

Gomanton, Between Gomantong Cave and Bukit Dulong Lambu, 10 Sep 1976, Taniura, M.

& Hotta, M. 602 (KYO); Sandakan, Tabin River, 16 Oct 2000, Venneulen, J.J. 2037 (SING);

Sukau, Panggi, 18 Sep 1996, Kiew. R. & Urn, S.R RK4211 (K, SING); Tawao, Oct 1922-Mar

1923, Elmer, A.D.E. 20569 (G, GH, K, SING). Sarawak: 1st Division, 28 May 1975, Biirtt,

B.L. B.8207 (E); 1st Division, Ban, 24 Apr 1955, Brooke, W.M.A. 9872 (L); 1st Division, Ban

District, Burtt, B.L. & Woods, P.J.B. B. 1898 (E); 1st Division, Ban District, Bukit Jebong, 7 Aug

1970, Lehmann, PF PFL364 (E); 1st Division, Bau District, Bukit Kapor, 22 May 1962, Burtt,

B.L. & Woods, P.J.B. B.1887 (E); 1st Division, Bau District, Fairy Cave, 9 Feb ]999, Jemree,

S. et al. S. 82056 (SAR); 1st Division, Bau District, Seburan Mine, 24 May 1962, Burtt, B.L. &

Woods, P.J.B. B.1925 (E); 1st Division, Bau Limestone Hills, Bukit Boring, 14 Nov 1985, Yii

et ai S. 503 5 3 (K, L); 1st Division, Bau, Guniing Stulang, 16 Oct 200\, Meekiong, K. SBC778

(SING); 1st Division, Bulit Majing, Tebakang, 16 Dec 1988, Kessler 226 (L); 1st Division,

Gunung Berloban, 19 Apr 1984, Yii & Othman S.46225 (K, L); 1st Division, Jambusan, Poak

Road, 16 Dec 1989, Frodin, D.G. & Bin Ismawi, O. 2077 (K); 1st Division, Padawan, Bukit

Megetang, 4 Mar 1969, Wright, E. & Chai, P S. 27 4 5 7 (E, K); 1st Division, Padawan, Gunung

Regu, 6 May 1975, Burtt, B.L. B.8089 (E); 1st Division, Taiton, Yii et al. S. 5 1 207 (L, SAR);

Bidi, Jul 1893, Ridley, H.N. s.n. (SING); Bidi Cave, Oct 1929, Clemens, J. & Clemens, M.S.

7434 (NY); ibidem, 22 Oct 1929, Clemens, J. & Clemens, M.S. 7627 (K); Kuching, Idzurni, H.

(Sc Togashi, M. s.n. (TI); ibidem, 29 Oct 1975-31 Oct \91 5, Idzurni, H. (Sc Togashi, M. s.n. (TI);

Kuching to Padawan, 30 Sep 1981, Croat, T.B. 53161 (SAR); Kuching, Daerah Kecil Siburan,

Kampong Mambong, 12 Dec 1999, Imaichi, R. et al. 2 (SAR); Kuching, Pemissen, Gunung

Braang, 8 Dec 2000, Jemree, S. etal. S. 84036 (SAR); Kuching, Penrissen, Mambong, Gunung

Bar, 19 Feb 1993, Lai & Rantai et al. S. 66009 (K, SAR); Kuching, Tiang Bekap, Mt. Mentawa,

1 1 Mar 1 967, CheM’, W.-L. CWL. 1288 (A, AAU, K, L, SING); Kuching, Tiang Bekap, Padawan

Road, 3 Apr 1960, Anderson, J.A.R. 12349 (K, L, SING); Samarahan , Sedan District, Lobang

Mawang, Bukit Selabor, 26 Sep 1968, Ilias bin Pale S. 28047 (K, L, SING); ibidem, 26 Sep

1968, Ilias bin Paie S. 28051 (K); Tambusan, Sep 1904, Ridley, H.N. s.n. (SING); Mongkos,

Gunong Selebur, 20 Apr 1999, Jamree et al S. 82239 (SAR); Sedan Distinct, Gunung Niyat,

Ulu Sg. Majat, 27 Feb 2002, Jemree, S. & Enjah, A. S.85584 (SAR). Peninsular Malaysia:

Kedah: Baling, 5 Aug 1941, Naven, J.C. 38038 (SING); Kota Setar, G. Keriang, 24 Jul 2009,

Rosdi, M, FRI66329 (SING); Pulau Langkawi, Gn. Raya Forest Reserve, 5 Oct 2010, Chan,

Y.M. FRI70632 (SING); Pulau Langkawi, Selat Panchor, 23 Nov 1934, Henderson, M.R.

29079 (K, SING); Langkawi, 20 Nov 1941, Corner, E.J.H. s.n. (SING); ibidem, 20 Nov 1941,

Corner, E.J.H. s.n. (SING); ibidem, Nov 1889, Curtis, C. 2107^.^ (SING); ibidem, Curtis, C.

2107a (SING); Langkawi, Gunung Raya, 22 Aug \9'^^, Anthony samy, S. SA959 (SING); Pulau

Langkawi, Pulau Dayang Buting, 23 Aug 1925, Holttum, R.E. 15125 (BO); Langkawi, Pulau

Langgun, 4 Nov 1971, Chin, S.C. 1759 (KLU). Kelantan: Bukit Mangga, 20 Feb 2003, Kie^^

R. RK5276 (SING); Gua Maka, 19 May 1990, Kiew, R. (S Anthonysamy, S. RK3020 (SING);

Kuala Aring, 1 Sep 1992-11 Sep 1992, Hamid H18 (SING). Pahang: Bukit Cheras, 10 Oct

1901, Henderson, M.R. 25205 (NY, SING); ibidem, 26 Nov 1984, Kiew, R. RK1566 (UPM);

Gunung Jebak Puyuh, Kiew, R. RK2145 (UPM); Gunung Senyum, 22 Aug 1986, Anthonysamy,

S. SA515 (UPM); Kampong Sri Jaya, Bukit Batu, 28 Oct 1986, Kiew, R. RK2369 (SING);

Kuala Lipis, 25 Aug 2008, Mohd Hairul, M.A. FRI60060 (SING); Kuantan, Bt Cheras, 25

Aug 1986, Anthonysamy, S. SA558 (UPM); Kuantan, Bukit Punching, 26 Nov 1984, Kiew’, R.

212

Card. Bull. Singapore 67(1) 2015

RK1570 (UPM); Maran, Jengka Forest Reserve, Hutan Lipur Jebak Puyuh area, 14 Oct 2008,

Mohd Hairul, M.A. FRI60066 (SING); Taman Negara, Batu Kepayang, 3 Oct 1984, Kiew, R.

RK1423 (UPM); Taman Negara, Batu Luas, 1 Oct 1984, Kiew, R. RK1349 (UPM); Temerluh,

Gn. Jebak Puyuh, Gn. Senyum, 9 Feb 2011, Kamarul Hisham, M, FRI67230 (SING). Perak:

Jan 1885, Kings Collector 7046 (SING); ibidem, Nov 1880, Kiinstler, H. 983 (SING); ibidem,

1890, Ridley, H.N. s.n. (SING); Batu Kurau, Dec 1884, Scortechini 1580 (SING); Chemar,

Perak Tong Temple, 23 Oct 1958, Sinclair, J. 9846 (E, L, SING, US); Grik, Flutan Simpan

Kemerong limestone area, 18 Nov 2011, Mohd Hairul, M.A. FRI 54071 (SING); Ipoh, 1 Dec

1966, Ng, FS.R FRI1795 (L, SING); Ipoh, Gunung Rapat, 2 Mar 1959, Allen, B.M. 4272

(SING); ibidem, 14 Jan 1983, Davis 69304 (E); Kampong Jahang, Gunung Kanda, 5 May

1962, Biirtt, B.L. & Woods, P.J.B. B.1820 (E); Larut, Apr 1884, King’s Collector 5872 (SING);

Larut & Matang, Lata Puteh, 22 Nov 2008, Yao, T.L. FRI65393 (SING); Tambun, Rotan Segar,

29 Nov 1960, Allen, B.M. 4654 (SING). Perils: Besih Hangat, 17 Nov 1929, Henderson,

M.R. 22875 (SING); Bukit Bintang Forest Resei've, Bukit Bintang, 9 Aug 1986, Weber, A.

860809-1/1 (E); Mata Ayer F.R., Biildt Rongkit, Kiew, R. RK3712 (KEP); Wang Kelian, 28 Jun

1993, Kiew, R. RK3653 (SING). Selangor: Kuala Lumpur, Jul 1890, Curtis, C. s.n. (SING);

Batu Caves, 24 Aug 1908, Ridley, H.N. 13380 (K, SING); ibidem, 4 Sep 1966, Burkill, H.M.

HMB4223 (SING); ibidem, 10 Jul 1906, Ernst, A. 1118 (L); ibidem, 23 Jan 1966, Hardial, S. &

Sidek474 (C, K, L, SING); ibidem, 9 Jun 1999, Kmv, R. RK4711 (SING); ibidem, 15 Sep 1968,

Kokawa, S. 6381 (KYO); ibidem, Dec 1898, Ridley, H.N. 8217 (SING); ibidem, Dec 1891,

Ridley, H.N. s.n. (SING); ibidem, 30 Oct 1967, Shimizu, T. & Fukuoka, N. M. 14 143 (AAU,

K, KYO, L, SING); Genting Highlands, Ulu Gombak, 26 Oct 1937, Nur, M. 34253 (SING).

Terengganu: Batu Biwa, 22 Oct 1986, Kiew, R. RK2287 (SING).

INDONESIA: Java: s.l., 1857-1861, de Vriese, W.H. s.n. (L); Unknown 2648 (L); Gunung

Buning (Gunung Bunder), 19 Dec 1893, Schiffner, V.F. 2567 (A, BO, K, L); Gombak, Sempor,

17 Apr 1936, Brinkman, R. 672 (BO). Jawa Timur: s.l., Coert, J.H. 1093 (L); Kangean Island,

Ardjasa, 1920, Backer, C.A.B. 26969 (BO); Kangean Island, Batoe Poetih, 28 Mar 1919,

Backer, C.A.B, 27805 (BO); Kangean Island, Tambajangan, 22 Mar 1919, Backer, C.A.B. 27410

(BO); Soekapoera, 3 Jun 1927, Backer, C.A.B. s.n. (BO); Tretes, Gunung Ai'djoeno, May 1919,

Bremekamp, C.E.B. s.n. (BO); Zuidergebergte, 27 Mar 1927, Backer, C.A.B. & Posthumus,

O. s.n. (BO); Madioen, Ngebel, 16 Mar 1898, Koorders, S.H. 29801 (K, L); Pogal, Mousset

450 (BO, L). Jawa Tengah: Lebak Barang, 12 Jan 1918, Backer, C.A.B. 23287 (BO). Jawa

Barat: Pasir Masigit, Feb 1933, Jacobson, E. 213 (BO); Papandajan, Mar 1930, van der Pijl,

L. 205 (BO); Bandung, Zollinger, H. 2002 (G, GH, P, S); Bogor, 1909, Backer, C.A.B. s.n. (L);

Bogor, Gunung Tjibodas , Feb 1912, Backer, C.A.B. 2480 (L); Cibodas, 1910, Backer, C.A.B.

32476 (BO); ibidem, 1917, Backer, C.A.B. 22091 (BO); ibidem, 1917, Backer, C.A.B. 22091

(BO); ibidem, 18 Mar 1928, van Steenis, C.G.G.J. 616 (BO); Tjiampea, Gunung Tjibodas,

15 Feb 1924, Bakhuizen van den Brink, R.C. 3568 (BO, L); ibidem, 30 Jan 1921, Bakhuizen

van den Brink, R.C. 5153 (BO, L); Klappa Noenggal, 1912, Backer, C.A.B. 5848 (BO, L);

Padelarang, Kampong Sempang, Coert, J.H. 625 (L); Res. Preanger, Tjidadap, Tjibeber, 23

Feb 1917, Bakhuizen 2608 (L); Tjampea, Burck, W. & de Monchy, B.J. s.n. (BO); ibidem,

9 Feb 1913, Koorders, S.H. 40438 (BO); ibidem, de Monchy, B.J. s.n. (L); ibidem, 4 May

1895, Hallier, H.G. 91 (L); ibidem, 25 Sep 1904, Hochreutiner, B.P.G. 1916 (G); UJung Kulon

National Park, Tjilintang, 25 Apr 1963, Wirawan, N. 45 (A, BO, K, L, SING); Palimena, Mont.

Prope, Junghuhn, F.W. 3 (K, L). Sumatra: s.l., Yates, H.S. 1939 (NY). Aceh: Gajolanden, 25

Feb 1937, van Steenis, C.G.G.J. 9267 (BO, K, L); Gaju and Alas Lands, Pendeng to Bivouac

Aer Putih Waterfall, 17 Feb 1937, van Steenis, C.G.G.J. 8879 (BO, L); Gunung Leuser Nature

Reserve, Ketambe Research Station, Gunung Giring, 17 Mar 2008, Wilkie, P. etal. PW764 (E);

Revision of Epithema

213

Gunung Leaser Nature Reserve, Ketambe, Mt Ketambe, 24 May 1972, de Wilde, W.J.J.O. & de

Wilde- Duyjjes, B.E.E. 12353 (BO, K, L); Ketambe, 17 Mar 2008, Sumadijaya, A. AX360 (BO);

Kloet Nature Reserve, South Kloet, Pucuk Lembang, 10 Jul 1985, de Wilde, W.J.J.O, d de

Wilde- Duyjjes, B.E.E. 19894 (BO, L); Pulau Weh, Sabaug, 31 Dec 1933, van Steenis, C.G.G.J.

5734 (BO). Lampung: Gunung Rati, 24 Nov 1921, Unknown 208 (BO, L); Muaradua, 13 Apr

1929, de Voogd, C.N.A. 357 (BO, L). Riau: Anambas Islands, Siantan Island, Terempak, 6 Apr

1928, Henderson, M.R. 20277 (K, SING). Sumatera Barat: Jorong Gasang, Lake Maninjau,

18 Jun 2{)\\ , Puglisi, C. & Hughes, M. CP61 (BO); Padang, Indarung, Kanipong Putih, 16 Feb

1981, Hotta, M. & Okada, H. 247 (KYO); Pajakumbuh, Halaban, 2 Feb 1958, Maradji 482

(L, SING). Sumatera Selatan: Aer Telanai, 11 Nov 1929, Van Steenis, C.G.G.J. 3925 (BO).

Sumatera Utara: Pulau Tello, Nov 1924, Boden-Kloss, C. s.n. (SING); Batu Islands, Pualu

Telo, 23 Jan 1 897, Unknown 654 (BO); Upper Langkat, 6 Nov 1 938, Lorzing, J.A. 1 7359 (BO);

Wampu, 1918-1919, Lesger 359 (BO). Borneo: Kalimantan Timur: West Koetai, Kombeng,

22 Nov 1925, Endert, F.H. 5143 (BO, L); West Koetai, Lahren, 29 Jun 1925, Endert, F.H. 1762

(BO, L); Tanah Grogot, Batu Kajang, Desa Kasungai, 27 Nov 1979, Ma ’wef, A. AM.250 (BO,

K) ; Berau, Sungai Kelai, Gunung Njapa, 18 Oct 1963, Kostemians, A.J.G.H. 21338 (BO, G, K,

L) ; Berau, Tanjung Redeb, Gmiung Buntung, 28 Nov 198 1-1 Sep 1981, Kato, M. et al. B. 11 733

(BO, KYO); Kenangan, 12 Aug 1979, Dransfield, J. 4402 (BO). Kalimantan Selatan: 1908,

Winkler, H. 2670 (BO, G, K, L); Gunung Serempaka, 26 Nov 1971, Dransfield, J. & Saerudin,

D. 2314 (BO, L); Djaro Dam, 11 Nov 1971, Kuswata 718 (BO); Muara Uja, 21 Nov 1971,

Dransfield, J. & Saerudin, D. 2268 (BO, K, L). Sulawesi Selatan: Bantimurong National

Park, 20 Feb 1938, Buwalda, P 3761 (BO), Sulawesi Tengah: G. Batoe, 1913, Rachmat 439

(BO); Luwuk, 9 Oct 1989, Coode, M.J.E. 5855 (K); Soroako-Wasuponda Road, 7 Jul 1979,

van Balgooy, M.M.J. 3922 (L). Sulawesi Tenggarah; Pohara, Muara Sampara, 14 Apr 1929,

Kjellberg, G.K. 1299 (BO, S); Pulau Butung, Jismal Camp, 12 Nov 1989, Coode, M.J.E.

6206B (K); Pulau Butung, 17 Feb 1929, Kjellberg, G.K. 195 (S). Sulawesi Utara: Bogani

Nani Waitabone National Park, 24 May 2002, Uji, T. 4595 (BO).

Notes. Epithema saxatile is one of the most widely distributed species of Epithema.

At first it appears that there are a few different forms of E. saxatile, one with petiolate

upper leaves, one with sessile upper leaves and one with occasional, solitary upper

leaves, but all of these can be found within a single population.

Some collections of this species have strongly variegated leaves but this

character does not appear to have any taxonomic significance.

17. Epithema steenisii Hilliard & B.L.Burtt, Edinburgh J. Bot. 54: 113(1 997). - TYPE:

Indonesia, Sumatra, Aceh, Gaju & Alas Lands, Gadjah to Pendeng, 16 February 1937,

van Steenis, C.G.G.J. 8825 (holotype L; isotypes BO (image seen), K, SING). (Fig.

10 )

Herb 3-1 5 cm high, caulescent, indumentum of strigose, setose or hispid hairs; stem

1-2 mm wide with 2-4 nodes, intemodes 1-3 cm long. Leaves strongly membranous,

petiolate, occasionally with more than one solitary lower leaf; petioles 0.2^. 5 cm

long; blade of lowest leaf 3.2-9 x 1 .7-4 cm, upper leaves 1 .5-9 x 0.7-4. 7 cm, more or

less lanceolate, elliptic, or naiTowly ovate, upper portion sometimes curved and upper

214

Card. Bull. Singapore 67(1) 2015

halves asymmetric, if the blade is asymmetrical, one side up to 1.5 times wider than

the other, apex acute to more rarely broadly acute, rarely rounded, base sub-truncate,

more rarely cuneate to rounded, inserted evenly on petiole or not, margin variable,

weakly serrate to bidentate; upper surface densely strigose to villous, hairs 0.75-1

mm long; lower surface densely pubescent or strigose and setose, hairs to 1 . 1 mm

long. Inflorescences 2^ per plant; peduncles 0.5-10.5 cm long, arising from leaf

axils; bracts cucullate, often completely enclosing inflorescence, 9-23 x 5-12 mm,

margin dentate, teeth small, lower surface strigose, hairs to 0.6 mm long, upper surface

glabrous or sub-glabrous; pedicels 1.5-3. 6 mm long, often with only strigose hairs

to 0.5 mm long, but sometimes also with additional minute hairs. Calyx 3.5-5 mm

long; tube 2-3 x 1 .2-2.3 mm, 1-2 times as long as wide, lobes 1. 5-2.7 x 0.6-1. 5 mm;

outside sparsely to densely strigose, hairs straight, occasionally hooked, 0.5-0.8 mm;

glabrous inside. Corolla white or pale lilac to lilac, 6.7-9 mm long; tube 4-6.5 x 1-1.5

mm; lobes 2.5 x 1 mm. Stamen filaments c. 1.2 mm long; anther characters unknown;

staminodes c. 1.2 mm long. Nectary of two discrete lobes, c. 0.8 mm high. Ovary

sub-cylindrical, c. 0.9 x 0.8 mm, sub-glabrous to densely pubescent, hairs hooked,

0.05 mm long and placed on upper portion of ovary; style 3. 7-4. 6 mm long, glabrous

or sub-glabrous, hairs at base of style. Fruit obovate to cylindrical, 1.4— 2.3 x 1.7-

2.3 mm; operculum 0.5-0. 8 mm, sub-glabrous to sparsely pubescent, as ovai*y. Seed

sub-cylindrical or narrowly to broadly ovoid, the ends often only slightly constricted,

0.3-0. 6 X 0. 1-0.3 mm, dark brown, pattern straight to spiralled and reasonably regular.

Distribution. Indonesia (Aceh).

Habitat and ecology. Shaded areas, on and among wet (mossy) rocks and in wet sand

by streams and rivers at 200-600 m altitude. Flowering and fruiting February, March,

May, June, August.

Provisional lUCN consei^ation assessment. Vulnerable VU D2. This species is only

known from the Indonesian province of Aceh in northern Sumatra and many of the

populations are from Gunung Leuser National Park. The species is only known to

occur below 600 m altitude and Gunung Leuser has been subjected to continued

deforestation despite being a “protected” area (Barber et al., 2002).

Additional specimens examined. INDONESIA: Aceh: Between Lau Simerah and Lau

Penanggajan, 20 Mar 1954, Alston, A.H.G. 14535 (A); Gaju & Alas Lands, Pendeng-Bivouac

Aer Putih Waterfall, 17 Feb 1937, van Steenis, C.G.G.J. 8879A (BO, L); Gunung Leuser Nature

Reseiwe, Camp Simpang, 18 Aug 1972, de Wilde, W.J.J.O. & de Wilde- Duyfles, B.E.E. 14353

(BO, K, L, US); Gunung Leuser Nature Reserve, Lau Alas Valley, Ketambe, 21 May 1972, de

Wilde, W.J.J.O. & de Wilde- Duyfjes, B.E.E. 12238 (BO, L); Ketambe, Alas River, 9 Jun 1979,

de Wilde, W.J.J.O. & de Wilde-Duyjjes, B.E.E. 18042 (BO, K, L).

Notes. This species is very similar to Epithema tenerum from Sulawesi but can be

distinguished by the difference in the indumentum on the outside of the ealyx. In

Revision of Epithema

215

addition the cucullate bract usually encloses the inflorescence in Epithema steenisii

whereas this is variable in E. tenerum. See further discussion in the notes section of

Epithema tenerum.

18. Epithema strigosum (C.B. Clarke) Hilliard & B.L.Burtt, Edinburgh J. Bot. 54: 113

(1997). - Epithema brunonis var. strigosum C.B.Clarke in A.DC. & C.DC., Monogr.

Phan. 5(1): 179. (1883). - TYPE: Indonesia, Sumatra, Gunung Singgalan, Korthals,

RW. 692 (lectotype L, designated by Hilliard & Burtt (1997) [see also note below]).

(Fig. 10)

Herb 6-20 cm high, acaulescent or with first leaves arising from a very short stem;

stem 2-4 nun wide with 0-1 nodes, intemodes 0-1.5 cm long. Leaves membranous to

strongly membranous, petiolate, one lower solitai^ leaf and one upper, or several leaves

in short-stemmed tuft; petioles 0.5-21 cm long; blade 1-19 x 0.5-15 cm, cordate to

elongate ovate, symmetiical or not, apex acute, base cordate, sub-auriculate or rounded,

inserted evenly on petiole or not, margin crenate or dentate, teeth wide at base and

mostly with rounded tips; both surfaces with hairs to 0.8 mm long. Inflorescences 1^

per plant; peduncles 6.5-23.5 cm long, originating opposite solitary leaves or at base

of petioles; bracts cucullate, completely enclosing the inflorescence, 14-18 x 8-15

mm, margin entire to dentate, lower surface setose and strigose, hairs medium density,

to 0.4 mm long, upper surface glabrous to strigose, hairs to medium density, primarily

straight or straight and hooked, to 0.4 mm long; pedicels 1 .7-5(-7.5) mm long, finely

densely pubescent, hairs hooked, also with or without sparse straight hairs 0. 1-0.4 mm

long. Calyx 3. 1-5.2 mm long; tube 1. 7-4.2 x 1.5-2. 7 mm, lobes L5-2.3 x 0.8-1. 7

mm; sparsely setose and strigose outside, hairs primarily hooked or both straight and

hooked, white, to 0.4 mm long, covering calyx; inside with hairs throughout to only on

lobes, hairs straight or both straight and hooked, to 0.13 mm long. Corolla white, 4.5-

8.3 mm long; tube 3.7-5 x 1.5 mm; lobes 0.8-3. 3 mm long; margin slightly fimbriate;

a partial or complete band of villous hairs in upper half inside. Stamens 2. 1-2.4 mm

long; filaments 1 .5-1 .8 mm long; anthers c. 0.6 mm long; staminodes 1 .2-1 .7 mm long.

Nectary of one or two lobes, 0. 5-0.3 mm long. Ovary 0.8-1 x 0.7-0. 8 mm, densely

pilose, hairs straight, white, 0. 1-0.3 mm long, placed on upper portion of ovary; style

3. 5^. 6 mm long, glabrous or sparsely pubescent at base of style, hairs straight; stigma

c. 0.5 mm wide. Fruit cylindrical to spherical, 1 .5-1 .9 x 1 ,1-2.3 mm; operculum c. 0.6

mm long, pubescence as on ovary. Seed ovoid, sometimes somewhat curved, 0.4-0.5

X c. 0. 1 mm, light to medium brown, pattern almost straight to spiralled, regular.

Distribution. Indonesia (Sumatra).

Habitat and ecology. Lithophytic. Found in shaded, humid areas beside rivers and

streams and along paths at 100-1000 m altitude. Flowering and fruiting January-

March, August.

216

Card. Bull. Singapore 67(1) 2015

Provisional lUCN conservation assessment. Endangered (EN Blab(iii)). This species

has a restricted distribution in West Sumatra in areas subject to continuing degradation

of natural habitats.

Additional specimens examined. INDONESIA: Sumatera Barat: KabupatenPadangPariaman,

Sipisang, 9 Jan 1995, Okada, H. et al. 1343 (BO); Gunung Singgalan, Singalang, Korthals,

P.W s.n. (L); Mt Sago, 9 Mar 1957, Meijer, W 5637 (L); Padang, Indarung, K. Putih, 16 Feb

1981, Hotta, M, Okada, H. & Kohyama, T. 319 (KYO); Padang, Indai’ung, Ladang Padi, K.

Putih, 8 Aug 1984, Hotta, M. & Okada, H. 1087 (KYO); ibidem, 14 Aug 1981, Hotta, M. &

Okada, H. 516 (KYO); Padang, Ladang Padi, 28 Aug 1984, Hotta, M., Okada, H. & Kohyama,

T. 689 (KYO, L); Padang, Ulu Gadut, 1 0 Jan 1 983, Hotta, M., Okada, H. & Ito, M. 402 (KYO);

Padang, Ulu Gadut, Sungai Gadut Gadang, 2 Aug 1984, Hotta, M., Okada, H. & Kohyama, T

324 (KYO); ibidem, 2 Aug 1984, Hotta, M., Okada, H. & Kohyama, T 339 (KYO).

Notes. Epithema strigosum is most similar to E. philippinum but differs from E.

philippinum in the larger calyx, the lack of a minute pubescence on the outside of

the calyx and the larger hairs only being sparse, the pubescence inside the calyx is

less dense, the leaf margin is crenate rather than dentate or serrate, the corolla can be

longer (4.5-8. 3 imn long rather than 5-6 mm long) and is reportedly white rather than

blue. Epithema strigosum is only found in Sumatra and E. philippinum only in the

Philippines. All specimens of Epithema strigosum are from an area of limestone east

of Padang.

Korthals s.n. (L0796748) is possibly a duplicate of Korthals 692, the type

collection, but has no field number, only a herbarium number of 113 which is also on

the lectotype. The plants are very similar and could be the same collection.

19. Epithema tenerum (C.B. Clarke) Hilliard & B.L.Burtt, Edinburgh J. Bot. 54: 113

(1997). - Epithema brunonis var. tenerum C.B. Clarke in A.DC. & C.DC., Monogr.

Phan. 5(1): 180 (1883). - TYPE: Indonesia, Sulawesi, South Sulawesi, Maros,

Zollinger, H. 1172 (lectotype G, designated here; isolectotypes L, P). (Fig. 6, 13)

Herb 5-14(-20) cm high, caulescent; stem 0.8-2 mm wide with 2-3 nodes, intemodes

0.5-5(-l 0) cm long. Leaves thinly membranous to membranous, petiolate, upper leaves

opposite or sub-opposite; petiole of lowest leaf 1-5 cm long, upper petioles 0.2-5 cm

long. Blade of the lowest leaf 2.3-12 x 1.4-8 cm, upper leaves 1.9-10 x 0.9-5 cm,

leaves usually broadly lanceolate to narrowly ovate or elliptic, but occasionally broadly

ovate or obovate to broadly obovate, symmetrical or not, if asymmetrical one side up

to 1 .4 times wider than the other, apex acute to broadly acute, base sub-cordate, sub-

tmncate or obtuse, inserted evenly on petiole or not, margin entire to weakly serrate;

upper surface primarily strigose, hairs straight, medium density, to 0.7 mm long; lower

surface may dry grey-green, sub-glabrous to pubescent, hairs medium density, to 0.4

mm long, veins raised with distinct, sparsely to densely pubescent, primarily strigose

hairs, to 0.6 mm long. Inflorescences 1-6 per plant; peduncles 0.3-7(-18) cm long,

originating from the terminal and lower axils; bracts small, sub-cucullate or cucullate.

Revision of Epithema

111

Fig. 13. Epithema tenerum (C.B. Clarke) Hilliard & B.L.Burtt. A. Habit. B. Underside of leaf.

C. Inflorescenee. D. Calyx. E. Flower opened out. F. Stigma lateral view. G. Fruit showing

seeds, placenta and operculum. H. Seeds. Scale bars; A = 4 cm; B = 5 mm; C, E = 3 mm; D, G

= 2 mm; F = 1 mm; H = 0.5 imn. Drawn by Claire Banks from Zollinger 1172 (A), Zollinger

1064 (B), P.C. Thomas and W H. Aroli 09-74 (C-G), Jaag 1600 (H).

218

Card. Bull. Singapore 67(1) 2015

6-16 X 3-7 mm, imdiilate to lobed with occasional dentation or dentate; lower surface

strigose, hairs straight, to medium density and 0.5 mm long; upper surface glabrous

or sub-glabrous; pedicels 0.8-3, 3 mm long, finely pubescent hairs to 0.13 mm long,

also with or without straight, strigose hairs to 0.5 mm long. Calyx occasionally tinged

purple, 1.7-3 .4 mm long; tube 0.9-2. 5 x 0.9-2. 5 mm; lobes 0.6-1. 5 x 0.6-1. 9 mm;

sub-glabrous, strigose or setose outside, also with minute, hooked hairs, hairs medium

density, straight hairs to 0.5 mm long, hooked hairs to 0.3 mm long; glabrous inside.

Corolla blue, pale-blue or pale purple, 5.2-6 mm long, tube 4-4.8 x 0.8-1. 3 mm;

lobes 0.9-1. 8 x 1-2.1 mm, margins entire to slightly crenate, dense band of villous

hair inside. Stamen filaments c. 1.5 mm long; anther characters unknown; staminodes

c.1.25 mm long. Nectary usually of two discrete lobes (occasionally one), margin

entire, 1 x 0.75-1.5 mm. Ovary sub-cylindrical, 0. 8-0.9 x 0.6-0.75 mm, with hooked

hair on upper half of ovary, hairs to densely pubescent, c. 0.1 mm long; style c. 3.3

mm long, glabrous; stigma up to c. 0.5 mm wide. Fruit sub-spherical, 1. 6-2.1 x 1.9-

2.2 mm, operculum 0.6-0. 8 mm long, indumentum as ovary. Seed narrowly ovoid to

ovoid, 0.4-0. 5 x 0. 1-0.2 mm, light to medium brown, pattern sub-erect to spiralled,

reasonably even.

Distribution. Indonesia (Sulawesi).

Habitat and ecology. Lithophytic, often found directly on limestone or on rocks of

unknown type at 20-750 m altitude. Usually beside streams and waterfalls but also on

road sides and on rock walls.

Provisional lUCN conservation assessment. Vulnerable (VU D2). This species is only

known from a small number of localities and these mostly karst limestone, a habitat

which is frequently exploited for cement and tourism.

Additional specimens examined. INDONESIA: Sulawesi Utara: Bolaang Mongondow,

Dumoga Bone National Park, Edwards Camp, 26 Mar 1985, de Vogel, E.F & Vermeulen, J.J.

6759 (L); Maros, Zollinger, H. 1064 (A, P). Sulawesi Selatan: Bantimurung, 16 Jun 1938,

Jaag, 0.1600 (L); ibidem, 20 May 1929, Rant, A. 40 (BO); ibidem, 20 Feb 1938, Buwalda,

P 3671 (L); ibidem, 20 Feb 1938, Biiwalda, P. 3677 (BO, L); ibidem, 13 Apr 1975, Meijer,

W.9133 (BO, L, US); Bantimurung, Bantimurung Waterfall, 15 Apr 2009, Thomas, D. &Ardi,

W.H. 09-74 (E); Pankadjene, 8 May 1931, Teruya, Z. 1848 (SING). Sulawesi Tenggara:

Kesali-Porema, 23 Oct \929, Kjellherg, G.K. 2618 (BO, S).

Notes. This species is most similar to Epithema steenisii from Aceh, Sumatra. It differs

mainly in the type of calyx hair, the density of the leaf hair and the length of the

corolla. On the calyx there are small, hooked hairs in addition to the longer, straight

hairs which are present in both Epithema tenerum and E. steenisii. In addition, the

longer hairs are generally shorter in Epithema tenerum than in E. steenisii (to 0.5 mm

vs. to 0.8 mm long). The hair on the upper surface of the leaves is usually much less

dense in Epithema tenerum than in E. steenisii and the corolla tube is 4—4.8 nmi and

4-6.5 mm respectively.

Revision of Epithema

219

Epithema tenerum usually follows the general Epithema pattern of having one

larger, lower leaf and one or more pairs of opposite, upper leaves. Some upper leaf

pairs are actually sub-opposite and this is most clearly seen in the larger specimens

(eg. Teruya 1848 (SING)).

The species has been collected from few localities in Sulawesi, principally from

Bantimurung in the south-west of the island. The specimens from northern Sulawesi,

De Vogel and Vermeulen 6759 (L) and, in particular, Kjellberg 2618 (S), are generally

larger than those from Bantimumng. The multi-specimen sheet from Pankadjene,

Teruya 1848 (SING), in southern Sulawesi, however, has plants spamiing the entire

size range.

20 . Epithema tenue C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 181 (1883). -

TYPE: Equatorial Guinea, Bioko, 1863, Mann, G. 2345 (lectotype K, designated here;

isolectotype P). (Fig. 14, 15)

Epithema thomense Henriq., Bol. Soc. Brot. 10: 145 (1892). - TYPE: Sao Tome and

Principe, Sao Tome, May 1888, Quintas, F 1272 (lectotype COI, designated here;

isolectotype K).

Epithema graniticolum A.Chev., Bull. Soc. Bot. France 58 (Mem. 8d.): 189 (1912).

- TYPE: Guinea, Montagne de Boola, Pays de Guerzes, 1048 m, 16 March 1909,

Chevalier, AJ.B. 20924 (lectotype P, designated here).

Herb 5-40 cm higli, acaulescent or caulescent, occasionally the plant is of only one leaf;

stem 1-5 mm wide with 1-3 nodes, intemodes 1-16.5 cm long. Leaves membranous

or strongly membranous, lower leaves petiolate, upper leaves petiolate and/or sessile

or occasionally with a solitary, upper leaf opposite peduncles; petiole of lowest leaf

2-10 cm long, upper petioles 0.2-8 cm long; blade of lowest leaf 6.5-24 x 4.5-19

cm, upper leaves 2-10 x 1. 5-8,4 cm, the lowest leaf very large and almost oblong to

broadly ovate, ovate or cordate, upper leaves ovate to broadly elliptic or spherical,

symmetrical or not, apex often rounded but also acute, base cordate or sub-cordate,

inserted evenly on petiole or base of upper leaves with one side truncate or obtuse

and the other cordate, margin variable, often sub-entire or minutely serrate or dentate,

but also serrate, dentate or crenate (usually the upper leaves); upper surface weakly

strigose or strigose, hairs to medium density, to 1.5 mm long; lower surface strigose

to medium density, 0. 5-0.8 mm long, and pubescent with minute, hooked hairs to

0.1 mm long. Inflorescences 1-6 per plant; peduncles 0.5-14 cm long, originating

from the base of the stem, opposite the solitary lower leaf or from the tenninal axil;

bracts frequently very large, cucullate and enclosing the inflorescence but may be

small and enclosing a small proportion of the inflorescence, 4-40 x 5-14 mm, margin

dentate or entire; lower surface weakly strigose or strigose, hairs 0.2-0. 8 mm long;

upper surface glabrous or sub-glabrous, hairs straight and concentrated towards

margins; pedicels 0.8^ mm long, weakly strigose, hairs to medium density and 0.8

Fig. 14. Epithema tenue C.B. Clarke. A. Habit. B. Inflorescence. C. Calyx. D. Flower opened

out. E. Stigma lateral view. F. Fruit showing seeds, placenta and operculum. G. Seeds. Scale

bars: A = 1 0 cm; B = 5 mm; C, F = 3 mm; D = 6 mm; E = 1 mm; G = 0.5 mm. Drawn by Claire

Banks from Letouzy 7722 (A, B), Letouzy 13973 (D, E) and Sita 2886 (C, F, G).

Revision of Epithema

221

Fig. 15. Distribution of Epithema tenue C.B. Clarke (•).

mm long, also hooked hairs to medium density and 0.1 mm long. Calyx 2-8 x 1.3-2

mm; tube 1.3-4 mm long; lobes 0.8-3. 8 mm long; with long hairs usually villous, or

occasionally to glabrous or sub-glabrous in maturity, hairs to high density, especially

in young flowers, hyaline, 0.6-1 .5 mm long; glabrous inside. Corolla blue-purple, tube

white, 9.6-10,5 mm long; tube 6. 6-7. 5 x 2 mm; lobes c. 3 mm long; a band of villous

hairs inside, hairs at half to two-thirds the way up the tube. Stamens 2-2.5 mm long;

filaments 1.5-2 mm long; anthers c. 0.5 mm long; staminodes 1.5-2 mm long. Nectary

usually apparently absent, but also of one or two lobes, sometimes reduced, 0.6-1 x

0.8-1. 3 mm, margin undulate or entire. Ovary ovoid, spherical or cylindrical, 0.8-1. 5

X 0.6-1. 3 mm, weakly hirsute to villous with hairs straight, 0.5-1. 5 mm long, with or

without additional hooked hairs to 0. 1 mm long (often seen most easily on the fruit

operculum); style 6 mm long, glabrous or one or two hairs at base, hair to 0.08 mm

long; stigma c. 0.5 mm wide. Frw/7 sub-cylindrical to sub-spherical, 1.5-3 x 1 .5-3 mm;

operculum 0.5-1 mm long, indumentum as on ovary. Seed nan’owly ovoid to ovoid,

0.4-0, 5 X 0. 1-0.2 mm, medium brown, pattern sub-erect or erect, reasonably even.

Distribution. Guinea, Sierra Leone, Liberia, Ivory Coast, Nigeria, Cameroon,

Equatorial Guinea, Gabon, Sao Tome and Principe, Democratic Republic of Congo,

Uganda, South Sudan.

Habitat and ecology^ Primarily lithophytic, often on granite at 180-1100 m altitude.

Found in humid areas, typically in shade, usually on or near rocky areas, in or beside

streams, waterfalls or cave entrances. Also found beside paths and highways. Flowering

and fruiting year round.

222

Card. Bull. Singapore 67(1) 2015

Provisional lUCN conservation assessment. Least Concern (LC). This is a widespread

species not known to be under any immediate tlireats.

Additional specimens examined. GUINEA: Kindia, Jacqiies-Felix, H. s.n. (P).

SIERRA LEONE: Between Kumbonla and Seredu, 1 5 Nov 1 965, Morton, J.K. SL2530 (E, K);

Loma Mountains, Oct 1944, Jaeger, P. 227 (K).

LIBERIA: Kolahun, Vahon, 7 Nov 1947, Baldwin, J.T. 10215 (K, US).

IVORY COAST: Chevalier, A.J.B. 21418 (K, P); Chevalier, A.J.B. 21506 (P); Banco, 11 Apr

1979, Knedl, M. 721 (G); Gueoule, 3 Feb \ 9S4, Hepper, F.N. & Maley, J. 8073 (K); Haut Nuon,

Chevalier, A.J.B. 21141 (P); Haut Sassandra, Droupleu to Zoanle, 5 May 1909, Chevalier,

A.J.B, 21457 (P); Man, Mt Tankoui, 4 Jul 1966, Ake Assi, L. 9043 (G); ibidem, Aug 1978,

Knedl, M. 9/365 (G); Tiapleu, 25 Sep 1955, Unknown 3296 (P).

NIGERIA: Gongola, Mambilla, River Nwum Forest Reseiwe, 30 Jun 1976, Chapman, J.D.

4505 (K); Sarduna, Gembu, Kamatan Forest Reserve, 14 Dec 68, Daramola, B.O. FHI62313

(K).

CAMEROON: Bamenda Province, Bamenda District, Metschum Falls, 26 Aug 1952, Savory,

H.J. UCI299 (K); Bamenda Province, Wum, Nbika’s By-Pass, 28 Jun 1951, Ujor, E. FHI29260

(K); Banyo, Tako-Atta Manga, 5 Jul 1967, Letouzey, R. 8732 (P); Collines a 5km au SW

d’Ebianeme-Yong pres Nyabessan, 10 Apr 1968, Letouzey R. 9238 (P); Ga Village, Pagan

Hill, 29 Aug 1 966, Letouzey, R. 7722 (COI, K, P); LolodorL 1 5 Jun 1918, Annet, E. 33 7 (K, P);

Muyuka, Munyenge, 29 May 1 976, Letouzey, R. 15020 (P); Muyuka, Munyenge, Mt Cameroun,

29 May 1976, Letouzey, R. 15026 (K, P); Nkambe, Vallen Forest, 12 Nov 1974, Letouzey, R.

13198 (P); Northern Cameroon, Munkep, Essu-Munkep Trail, 7 Jul 1975, Letouzey, R. 13973

(P); South West Province, Etinde, Upper Boando, Upper Boando Village, 1 Dec 1993, Cable,

S. 283 (K); South West Province, Fako, Buea, Likombe, 2 Sep 1992, Nkeng, P. 90 (K); South

West Province, Kupe-Muanenguba Division, Kupe Village, 27 May 1996, Etuge, M. 1973 (K);

ibidem, 10 Jul 1996, Kenfack, D. 259 (K); ibidem, 15 Jul 1996, Zapfack, L, 929 (K); South

West Province, Kupe-Muanenguba Division, Ngomboku, 10 Dec 1999, Mackinder, B.A. 303

(K); South West Province, Kupe-Muanenguba Division, Nyale, 17 Nov 1988, Cheek, M.R.

et al. 9647 (K); South West Province, Kupe-Muanenguba Division, Nyasoso, 24 Jun 1996,

Cable, S, 3256 (K, WAG, YA); ibidem, 19 Dec 1993, Cable, S. & Ajebe, E.F 644 (K); ibidem,

2 Jun 1996, Etuge, M. 2057 (K); ibidem, 19 Oct 1995, Sidwell, K. et al. 305 (K); South West

Province, Kupe-Muanenguba Division, Nyasoso, Mount Kupe, 7 Feb 1996, Cable, S. 3532 (K,

YA); ibidem, 7 Oct 1998, Harris, D. 5815 (E); ibidem, 23 Oct 1995, Sebsebe Demissew 4984

(K); ibidem, 7 Jul 1992, Wheatley, J.I. 4117 (K); ibidem, 2 Jun 1996, Zapfack, L. 597 (K);

South West Province, Meme, Nyasoso, 23 Jun 1996, Cable, S. 3208 (K); South West Province,

Meme, Nyasoso, Mt Kupe, 5 Aug 1993, Balding, S. & Sivell, D. 37 (K); ibidem, 19 Sep 1992,

Cable, S. 19 (K).

EQUATORIAL GUINEA: Bioko, 16 Jan 1947, Guinea, E. 1408 (K); Bioko, Basile, 30 Aug

1990, Carvalho 4464 (K); ibidem, 29 Jul 1986, Carvalho 2130 (K).

GABON: 13 Mar 1925, Le Te.stu, M.G. 5449 (P); Lastoursville, 1929-1931, Le Testu, M.G.

7018 (K); Libreville, 2 Feb 1 899, Klaine, R.P 151 7 (K); ibidem, Klaine, R.P. 2666 (P); Ndende,

22 Nov 1983, de Wilde-Dujfjes, B.E.E. et al. 745 (K).

SAO TOME AND PRINCIPE: St Thome, Henriques, J.A. 5 (K); ibidem, Henriques, J.A. 12

(K).

DEMOCRATIC REPUBLIC OF CONGO: Meya, Camp O.R.S.T.O.M., 12 Feb 1969, Sita, P.

2886 (P).

Revision of Epithema

223

UGANDA: 1906, Bagshawe, A.W.G. 1282 (US); Bundibugyo, Sempaya Hot Springs, 14 Sep

1997, Lye, K.A. & Katende, A.B. LYE22885 (K); Bwamba, Balanga, 22 Nov 1935, Thomas,

A.S. ThJ529 (K); Bwamba, Bulanga, 30 Sep 1932, Thomas, A,S. Th.729 (K); Lake Albert

Edward, 1906-1907, Bagshawe, A. W.G. 1376 (US).

SOUTH SUDAN: Imatong Mountains, Talanga, 26 Nov 1980, Friis, I. & Vollesen, K.B. 462

(C, K).

Notes. There are two forms of Epithema tenue, an acaulescent and a caulescent form.

The acaulescent fonn is found in the Central African Republic, Equatorial Guinea

(Bioko), Cameroon and Gabon while the caulescent fonn is found in the Ivory Coast,

Nigeria, Sudan, Uganda, Liberia, Guinea, Sierra Leone, Cameroon and Gabon. In

only Cameroon and Gabon are there collections of both fonns. The acaulescent form

has, primarily, big bracts and inflorescences and calyces which are pubescent with

long hairs. The size of the inflorescence is more variable in the caulescent form, but

can be still large and pubescent. The duplicates of Harris 5815 (E) from Cameroon

show the acaulescent and caulescent forms can be found within one population. In this

collection the acaulescent specimens are small and membranous and have small bracts

and inflorescences but still have the characteristic long, straight hair on the ovary/

operculum.

Some of the caulescent specimens could be mistaken for Epithema carnosum.

They differ IroniE. carnosum by the size of the bract and calyx and the dense indumentum

on the outside of the calyx. Some, however, have small bracts, inflorescences and

calyces, and the indumentum is sparse. These can still be separated using the length of

the hair on the operculum. While this hair is straight in both species, in Epithema tenue

it is from 0.5-1 .5 mm long and in E. carnosum it is from 0.08-0.25 mm long. There

are a small number of specimens that are not readily differentiated from Epithema

carnosum as the indumentum on the ovary is short. While it is sparse, the indumentum

of the calyx of these specimens is that of E. tenue, being hyaline, villous and long

instead the hyaline to white, short and more strigose hair of E. carnosum. As these

two species occur on different continents confusion between the two should not be a

problem.

Excluded Species

Epithema triandrum (Blanco) Fem.-Vill., Nov App. 150. (1880). - Ophiorrhiza

triandra Blanco, FI. Filip., ed. 2 [F.M. Blanco] 65 (1845). = Ophiorrhiza oblongifolia

DC. (see Merrill, 1918).

ACKNOWLEDGEMENTS. We thank the Sibbald Trust for providing the funding for this

revision. We thanlc the directors and curatorial staff of the herbaria that loaned material or

hosted visits. Martin Pullan is gratefully acknowledged for his help with the Padme taxonomic

database. Claire Banks is thanked for the beautiful illustrations and Serena Lee is thanked

224

Card. Bull. Singapore 67(1) 2015

for her work on the maps. Junji Matsumura (FU) is thanked for providing images of the type

of Epithema calcicola and Abdulrokhman Kartonegoro (BO) for images of isotypes of E.

horsfieldii van epiphyllum, E. longitubum and E. steenisii. The staff of the Forest Herbarium

Bangkok and Pramote Triboun are gratefully aeknowledged for enabling and assisting field

work in Thailand for the second author. Hamiah Atkins and Ruth Kiew are thanked for their

thorough reviews of the manuscript.

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Appendix 1. Index of exsiccatae.

Epithema benthamii C.B, Clarke (1); Epithema carnosum Benth. (2); Epithema ceylanicum

Gardner (3); Epithema dolichopodum Hilliard & B.L.Burtt (4); Epithema horsfieldii (R.Br.)

A.DC. (5); Epithema involucratum (Roxb.) B.L.Burtt (6); Epithema longipetiolatum

(Merr.) Hilliard & B.L.Burtt (7); Epithema longitubum Hilliard & B.L.Burtt (8); Epithema

madulidii Hilliard & B.L.Burtt (9); Epithema membranaceum (King) Kiew (10); Epithema

parvibracteatum Hilliard & B.L.Buitt (11); Epithema philippinum (Hilliard & B.L.Burtt)

Bransgrove (12); Epithema pusillum (C.B. Clarke) Bransgrove (13); Epithema rennellense

Hilliard & B.L.Burtt (14); Epithema sarawakense Hilliard & B.L.Burtt (15); Epithema saxatile

Blume (16); Epithema steenisii Hilliard & B.L.Burtt (17); Epithema strigosum (C.B. Clarke)

Hilliard & B.L.Burtt ( 1 8); Epithema tenerum (C.B. Clarke) Hilliard & B.L.Burtt (19); Epithema

tenue C.B.Clarke in A.DC. & C.DC. (20).

Aet & Idjan 32 (7), 942 (7); Ahmad 1 (15); Ake Assi, L, 9043 (20); Allen, B.M. 4272 (16),

4654 (16); Almeida, S.M. s.n. (13); Alston, A.H.G. 14535 (17); Anderson, J.A.R. et al.

S.26090 (15); Anderson, J.A.R. 12349 (16), S.20957 (15), S.31770 (15); Anderson,

J.A.R. & Keng, H. KlOO {15)-, Annet, E. 337 (20); Anthonysamy, S. SB7 (4); SA515 (16),

SA558 (16); SA959 (16); Aigent, G.C.G. et al. 905b (15), 29038 (1); Argent, G.C.G.

C8754a (7); Argent, G.C.G. & Collins, M. 733 (15); Argent, G.C.G. & Kerby, R. 621

(15) ; Ashton, RS. A.339 {\5),A.340 {\5)\Atha, D. 4785 (2).

Backer, C.A.B. s.n. (16), 2480 (16), 2789 (5), 5848 (16), 6607 (5), 6687 (5), 11469 (5), 22091

(16) , 23287 (16), 26969 (16), 27410 (16), 27805 (16), 32476 (16), 36621 (5); Backer,

C. A.B. & Posthumus, O. s.n. (16); Bagshawe, A.W.G. 1282 (20), 1376 (20); Bakhuizen

van den Brink, R.C. 2608 (16), 3568 (16), 5153 (16); Balalaishnan, N.P. 10815 (2);

Balding, S. & Sivell, D. 37 (20); Baldwin, JET. 10215 (20); Banggilon, K. SAN66579

(16); Barnes, E. 928 (3), 1056 (3), 1300 (3); Bartlett, H.H. 7740 (1 0), 14487 (3), 14744

(3), 15380 (3); Beaman, J.H. 10068 (4), 11727 (15); Beumee, J.G.B. 1328 (5); Bin

Rajab, M.K. 714 (16); Bloembergen, S. 3458 (8), 3496 (8); Blume, C.L. s.n. (16), 1029

(16); Boden-Kloss, C. s.n. (16); Boerlage, J.G. 528 (7); Brahmam, M. 2776 (3); Brandis,

D. 676 (3); Bremekamp, C.E.B. s.n. (16); Brinkman, R. 672 {\6y, Brooke, W.M.A. 9872

(16); Brown, R. s.n. (6); Biinnemeijer, H.A.B. 10713 (5), 10890 (5); Burck, W. & de

Monchy, B.J. s.n. (16); Burkill, H.M. HMB4223 (16); Burtt, B.L. B.8089 (16), B.8207

(16), B.8227 (15), B.8336 (15); Burtt, B.L. & Woods, P.J.B, B.1820 (16), B.1887 (16),

B.1898 (16), B.1925{\6\ B.2003{\5\ B.2230 {\5)\ Buwalda, P. 3671 (19), 3677(19),

3761 (16), 5110 (7), 5202 (7), 5748 (7), 6006 (7), 7225 (5), 7542 (5).

Cable, S. 19 (20), 283 (20), 3208 (20), 3256 (20), 3532 (20); Cable, S. & AJebe, E.F 644

(20); Carvalho 2130 (20), 4464 (20); Chai, P. S.30372 (15); Chan, Y.M. FRI70632 (16);

Chantharanothai, P. et cd. 90/303 (16); Chapman, J.D. 4505 (20); Charoenphol, C. et

226

Card. Bull. Singapore 67(1) 2015

al. 3661 (16), 3662 (16), 3976 (10); Cheek, M.R. et al. 9647 (20); Chennsirivathana,

C. 1 (3), 413 (16); ChevaHer, A.J.B. 20924 (20), 21141 (20), 21418 (20), 21457 (20),

21506 (20); Chew, W-L. CWL.293 (15), CWLJ017 (15), CWL.1288 (16); Chin, S.C.

392 (11), 1404 (10), 1514 (10), 1759 (16); Clung, R.C 6421 (2); Clarke, C.B. 8997 (2),

12300 (2), 15665 (2), 19076 (2), 24805 (2), 36192 (2); Clason, E. W. C99 (5); Clemens,

J. & Clemens^ M.S. 7434 (16), 7627(16); Clemens, M.S. 724 (1); Coert, J.H. 625 (16),

1093 (16); Coode, MJ.E. 5313 (1), 5855 (16), 6206B (16); Corner, EJ,H s.m (16);

Coster SM. (5); CP, BS & BN 2997 (16); Croat, T.B. 53161 (16); Ctiadra, A. A.2168

(16); Cuming, H. 823 (3), 1265 (1); Curtis, C. s.n. (16), 955 (10), 2107 {16), 2107a (16).

Daramola, B.O. FHI62313 (20); Davis 69304 (16); de Monchy, B.J. s.n. (16); de Vogel, E.F.

& Vermeulen, J.J. 6759 (19); de Voogd, C.NA. 357 (16); de Vriese, W.H, s.n. (16); de

mide, J.J.FE. 1155 (1); de Wilde, W.J.J.O. & de Wilde- Duyfjes, B.E.E. 12238 (17),

12353 (16), 14353 (17), 18042 (17), 19894 (16), 21918 (5), 21940 (5); de Wilde-Dujfjes,

B.E.E. et al. 745 (20); Deschamps s.n. (3); Dhal, N.K. & Rout, N.C. 8122 (3); Dissing,

FI. 2738 (14), 2798 (14); Dransfield, J. 3869 (1), 4402 (16); Dransfield, J. & Saerudin,

D. 2268 (16), 2314 (16); Duthie, J.F. 2990 (2).

Eberhardt 3363 (3); Edaho, G. s.n. (1), 418 (1), 435 (1), 435a (1), 720 (1), 746 (1), 755 (1),

7965 (1), 3070A (1), 3478 (1), BS41748{\), BS77711 (4), BS77570 (4); Ellis, J.L. 20452

{3)\ Elmer A.D.E. 5575(1), 9695(1), 75476(1), 75626(1), 75577(1), 77557 (1), 75255

(1) , 20569 (16); Endert, FH. 1762 (16), 5143 (16); Ernst, A. 1118 (16); Escritor, L.

5527452 (1); Esquirol, J. 711 (2); Etuge, M. 1973 (20), 2057 (20); Eyma, P.J. 1826 (7),

2574 (7).

Faden, R.B. & Faden, A.J. 76/494 (3); Fenix, E. BS26049 (3), BS28106 (3); Feuilletau de

Bruyn, W.K.H. 335 (1); Forrest, G. 15699 (2); Fosberg, FR. 57278 (3); Frodin, D.G. &

Bin Ismawi, O. 2077 (16).

Gachalian, F. 147 (1); Gafui, I.H. & collectors BSIP14759 (14), BSIP14763 (14); Gallatly,

G. 1035 (16); Gardner, G. 606 (3); Geesink, R. cS: Phengklai, C. 6082 (16), 6257 (16);

Geqffray, M. 95 (3); Grey-Wilson, C. & Grey-Wilson, C.M. 3016 (3); Grupe, D.A. 200

(3); Guinea, E. 1408 (20).

Haegens, R.M.A.P. & Klazenga, N. 515 {\5y, Hallier, H.G. 91 (16), 4725 {\);Flamid H18 (16);

Hansen, C. 17 (15); Hardial, S. & Sidek 474 (16); Harris, D. 5815 (20); Haviland,

G.D. s.n. (15); Haviland, G.D. & Hose, C. 3525 R (15); Henderson, M.R. 19406 (10),

20277 (16), 22552 (10), 22580 (10), 22875 (16), 25010 (10), 25205 (16), 29079 (16);

Hennipman, E. 610g (7); Henriques, J.A. 5 (20), 12 (20); Heniy, A. 1916 (3), 12280

(2) , 12705 (2), 75497 (2); Hepper, FN. & Maley, J. 8073 (20); Hicks, D. 151 (1);

Hochreutiner B.P.G. 7976 (16); Holttum, R.E. 15125 (16); Hooker, J.D. & Thomson,

T. s.n. (2), 2161 (2); Hoover, W.S. et al. HW11417 (5), Deden 757 (5); Horsfield, T.

Cyrtandra 3 (5), 55 (5); Hotta, M. et al. 313 (18), 579 (18), 524 (1 8), 559 (1 8), 402 (18),

659 (18); Hotta, M. 14402 (15), 75257 (15); Hotta, M. & Okada, H. 247 (16), 576 (18),

1087 (18); Hsieh, C.F et al. s.n. (3).

Iboet 221 (6); Idjan 175 (1); Idzumi. H. & Togashi, M. s.n. (16); Ilias bin Paie S. 28047 (16),

5.25657 (16); Imaichi, R. etal. 2 (16), 30 (15), 55 (15); Imin, K. FR163208 (11).

Jaag, O. 1600 (19); Jacques-Felix, H. s.n. (20); Jacobson, E. 199 (5), 275 (16); Jaeger, P. 227

(20); Jaheri 1522 (15); Jamree et al. S. 82239 (1 6); Jayasuriya, A.H.M. 843 {3);Jemree,

5. & Enjah, A. S. 85584 (16); Jemree, S. et al. S. 82056 (16), S. 84036 (16); Jeswiet, J.

670 (5); Julaihi S. 86868 (15); Julaihi et al. S. 76938 (15); Julia, 5. et al. S. 93376 (15),

5.94757 (15), 5.99565 (15); Junghulm, F.W. 3 (16).

Revision of Epithema

227

Kamarul Hisham, M. FRI67230 (16); Kanehira, R. & Hatusima, S. 11848 (1), 13143 (1);

Kasem 641 (3); Kato, M. et al. C.5728 (7), C.5232 (7), C.5144 (7), C.6007 (7), C.6908

(1), C.11441 (7), BJ1733 (16); Keenan, J. et aL 798 (16); Kenfack, D. 259 (20); Kerr,

A.F.G. 2655 (16), 6381 (16), 7657(16), 12385 (16), 13053 (3), 13183 (16), 15775 (10),

15893 (10), 19385 (16); Kessler 226 (16); Kie\v, R. et al. RK4321 (4); Kiew, R. RK525

(15) , RK1231 (10), RK1347 (10), RK1349 (16), RK1423 (16), RK1498 (11), RK1566

(16) , RK1570 (16), RK1584 (10), RK1590A (10), RK2145 (16), RK2287 (16), RK2369

(16), RK3653 (16), RK3712 (16), RK4711 (16), RK4759 (4), RK5040 (16), RK512()

(16X RK5254 (10), RK5276 (16); Kiew, R. & Anthonysamy, S. RK3020 (16); Kiexv, R.

& Lim, S.F. RK4138 (4), RK4211 (16); King, G. s.n. (2); King’s Collector 5872 (16),

7046 (16); Kjellberg, G.K 195 (16), 314 (6), 315 (6), 1299 (16), 2616 (1), 2618 (19);

Klaine, R.P. 1517 (20), 2666 (20); Knedl, M. 9/365 (20), 721 (20); Kochummen, K.M.

FRI26221 (11); Kokawa, S. 6381 (16); Kokawa, S. & Hotta, M. 551 (16), 557 (16), 565

(16), 1073 (4), 1125 (4); Koorders, S.H. 29801 (16), 40438 (16); Kooy, C.W. 403 (5),

789 (5); Kornassi 482 (1), 621 (7), 967 (7); Korthals, P.W s.n. (18), 692 (18); Koster,

C. BW10773 {\\ BW13665 (1); Kostemians, AJ.G.H. 1103 (16), 7546(16), 1401 (16),

18053B (5), 18629 (5), 18839 (5), 21338 (16); Kostermans, A.J.G.H. & Soegeng 914

(1); Kulkarni, B.G. 108673 (13); Kunstler, H. 983 (16); Kuswata 105 (5), 121 (5), 213

(5), 775 (16).

Lace, J.H. 4892 (2); Lai & Rantai et al. S. 66009 (16); Lamb, A.L. 8/85 (16); Larsen, K. et al.

722 (16), 2923 (3), 3363 (16), 30967 (16), 41243 (16), 41413 (16), 41450 (16), 41713

(10), 42245 (10), 42312 (10), 42471 (10), 42579 (10), 42584 (10), 43589 (3), 43771

(3), 44271 (10), 46055 (16), 46161 (2), 46426 (3), 46861 (3); Le Testu, M.G. 5449 (20),

7018 (20); Lehmann, PF. S.30091 (15), PFL364 (16), PFL596/S, 30372 (15), S.30091

(15) ; Leong, P. et al. PL215 (15), PL320 (15); Lesger 359 (16); Letouzey R. 7722 (20),

8732 (20), 9238 (20), 13198 (20), 13973 (20), 15020 (20), 15026 (20); Lim, S.P et al.

580 (16), LSP675 (4), LSP681 (4), LSP738 (16); Lobb, T. 387 (3); Loeters, J.J. 1625 (5);

Loher, A, 1554 (3), 1555 (3), 7556 (3), 4231 (I ), 6665 (3), 6669 (3); Lopez, G. BS42031

(1), BS42049 (1), BS41343 (9); Lorzing, J.A. 17359 (16); Lucas 1025 (3); Lutjeharms,

W.J. 4598 (15), 5112 (15); Lye, KA. & Katende, A.B. LYE22885 (20).

Mackinder, B.A. 303 (20); Madulid, D.A. et al. 11563 (9); Maier Sarip, R. 162 (5); Mann, G.

2345 (20); Maradji 482 (16); Marcan, A. 1869 (3); Ma’roef, A. AM. 250 (16); Martin,

PJ. S.39273 (15); Maxwell, J.F s.n. (16), 72-362 (16), 73-107 (3), 73-285 (16), 74-814

(16) , 75-958 (1 6), 84-55 (1 6), 86-723 ( 1 6), 86-893 (10), 87-1154 (3), 89-920 (2), 93-886

(16), 95-536 (3), 99-278 (2); McGregor, R.C. BS19951 (1), BS43566 (3); McKee, H.S.

1958 (3), 6577 (3); Meekiong, K. SBC778 (1 6); Meijer; W. 5657 (1 8), 9133 (19), 10045

(1), SAN20756 (16); Mendoza, D.R, & Convocar, P P. 226 (12), 297 (12); Mendum,

M. et al. 34 (5), 45 (1), 775 (1 ), 180 (1 ); Merrill, E.D. 52 (3), 4216 (3), 5106 (1 ), 8222

(1); Middleton, D.J. et al. 4055 (16), 4284 (3), 4330 (16), 4545 (16), 4428 (16), 4544

(3), 4579 (3), 5350 (1 6), 5565 (1 6), 5425 (1 6), 5524 (1 6), 5675 (3), 5657 (3), 5765 (3);

Middleton, D.J. & Triboun, P. 4846 (3); Mohd Hairul, M.A. FRI54071 (16), FR160060

(16), FR160066 (16); Mohidin S.21608 (15); Moktar, A. et al. S.49467 (15); Mooney,

H.F 2067 (3), 2754 (3), 5555 (2), 3891 (2), 4045 (2), 4092 (2); Morton, J.K. SL2530

(20); Mousset 450 {\6); Mrs. General Walker s.n. (3), 140 (3); Mwra/a, G. etal. T. 15 146

p.p. (3), T.15146 p.p. (2), T.15156 (2), T.16922 (3), T.17092 (16), T.38467 (16), 55544

(16), 7:55557(16).

Nadiah, 1. et al. SAN149136 (4); Nair, K.K.N. 1354 (3); Namba, T. & Mikage, M. 82-413

228

Card. Bull. Singapore 67(1) 2015

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Gardens’ Bulletin Singapore 67(1): 231-250. 2015

doi: 10.3850/S2382581215000186

231

Synoptic overview of Acacia sensu lato (Leguminosae:

Mimosoideae) in East and Southeast Asia

B.R. Maslin

Western Australian Herbarium, Department of Parks and Wildlife,

Locked Bag 104, Bentley Deliveiy Centre, Western Australia 6983

Baice.IVIaslin@dpaw.wa.gov.au

Honorary Research Associate

Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, Singapore 259569

ABSTRACT. Recent research shows that the formerly broadly circumscribed, pantropical genus

Acacia Mill. (Leguminosae: Mimosoideae) is polyphyletic and should be treated as comprising

at least five genera, namely. Acacia Mill, sensu stricto, Acaciella Britton & Rose, Mariosousa

Seigler & Ebinger, Senegalia Raf. and Vachellia Wight & Arn. The indigenous flora of Acacia

sensu lato in East and Southeast Asia comprise 52 species: 32 species (38 taxa) of Senegalia, 12

species of Acacia sensu stricto and eight species of Vachellia. These species are listed and their

geographic distributions given, showing that Acacia sensu lato is unevenly distributed across

the region, with centres of species-richness in Indonesia, Myanmar and Thailand. A summary

of the classification histoiy of Acacia sensu lato is provided and nomenclatural impacts of the

recent retypification of Acacia briefly discussed.

Keywords. Classification, nomenclature, phylogeny, Senegalia, Vachellia

Introduction

Molecular and other evidence has shown that the fonnerly broadly circumscribed,

pantropical genus Acacia Mill. (Leguminosae: Mimosoideae) is polyphyletic and

should be treated as comprising at least five genera, namely, Acacia Mill, sensu stricto,

Acaciella Britton & Rose, Mariosousa Seigler & Ebinger, Senegalia Raf. and Vachellia

Wight & Arn. (Miller & Seigler, 2012). Collectively, these five genera are called

Acacia sensu lato in the discussion below. The name Acacia is now conserved with a

new type (McNeill & Turland, 2011), an action which has had global nomenclatural

repercussions. AJthough increasingly understood and adopted, the new classification

for Acacia sensu lato has not yet been universally adopted, despite the fact that names

congment with the new classification are now available for most currently accepted

taxa.

The indigenous species of Acacia sensu lato in East and Southeast Asia comprise

52 species, mostly belonging to Senegalia (see Appendix 1), These species are unevenly

distributed across the region with centres of species-richness in Indonesia, Myanmar

and Thailand (see Appendix 2). The tenu East Asia here refers to China and Taiwan

where species of Acacia sensu lato occur both naturally and as introductions. The term

232

Card. Bull. Singapore 67(1) 2015

Southeast Asia includes the countries from Papua New Guinea through Indonesia,

Philippines, Brunei, Vietnam, Laos, Cambodia (Kampuchea), Singapore, Malaysia,

Thailand to Myanmar (Burma). Although New Guinea is geographically part of

Australasia, it is included here for convenience and because the island is included in

the largest Flora project in Southeast Asia, namely. Flora Malesiana. The species of

Acacia sensu lato of this whole region are currently under review (Maslin, in prep.),

a work that benefits greatly from the excellent taxonomic foundation provided by the

late Ivan Nielsen for Southeast Asian taxa (Nielsen, 1980, 1981, 1985a, 1985b, 1992)

and the treatments by Sun & Chen (1990) and Wu & Nielsen (2010) for taxa from

China.

The aims of this communication are to summarise the rationale underlying the

new classification and updated nomenclature of Acacia sensu lato and to show how

this structure and these names apply to the indigenous species of East and Southeast

Asia. This will provide an introduction to the above mentioned taxonomic review

and will facilitate implementation of the new classification and nomenclature for the

region.

Phylogeny and classification of Acacia sensu lato

As defined until relatively recently, the genus Acacia comprised a very large group of

about 1350 species distiHbuted throughout tropical and warm temperate areas of the

world, occurring on all continents except Antarctica (web. ref 1). However, during

the past decade the genus has undergone fragmentation, a process that has been driven

largely by evidence derived from molecular phylogenetic studies.

According to Bentham (1840), during the 80 years following its original

description by Miller (1754) Acacia had become an “unwieldy, ill-defined, and

comparatively unnatural assemblage of plants”. Bentham (1842) remedied this situation

by restricting the name Acacia (today’s Acacia sensu lato) to Mimosoid plants having

indefinite, free stamens, a definition that has persisted to modem times. In a series of

subsequent publications, culminating in his 1 875 magnum opus, Bentham did much to

clarify not only the definition of Acacia but also its internal classification (Bentham,

1875). During the ensuing 60 years 15 new genera were described but these were

ultimately treated as congeneric with Acacia sensu lato. A discussion of this generic

history is outlined in Maslin et al. (web ref. 1).

In 1986 Pedley published a reassessment of the classification of Acacia,

dividing the genus into three, namely. Acacia sensu stricto, Senegalia diwd Racosperma

(DC.) Mart. These genera corresponded to three subgenera of Acacia sensu lato that

had previously been recognised by Vassal (1972), namely. Acacia subgenera Acacia,

Aculeiferum Vassal and Heterophyllum Vassal (=subgen. Phyllodineae (DC.) Seringe)

respectively. At the time, Pedley ’s classification was not widely adopted for reasons

that are outlined in Maslin (1989) and Maslin et al. (web ref. 1). One of these reasons

was that the evidence presented was considered inconclusive and/or incomplete.

Synoptic overview oi Acacia sensu lato

233

especially considering the significant impact to global nomenclatural that would flow

from such a division.

However, since 1986 a number of broad-based, comparative studies (particularly

using chloroplast and nuclear DNA) provided evidence that enabled more robust

and infoinied decisions to be made concerning the phytogeny and classification of

Acacia (see publications listed in Miller & Seigler, 2012). These studies included not

only species of Acacia sensu lato and Faidherbia A.Chev. (which together comprise

tribe Acacieae) but importantly, also included representatives from the related tribes

Ingeae and Mimoseae. As summarised in Miller & Seigler (2012) this genetic data

has consistently demonstrated that Acacia sensu lato is polyphyletic and comprises at

least five monophyletic groups, which each warrant recognition as a distinct genus.

These genera are shown in Table 1 and comprise (1) two small New World endemic

genera, Acaciella (resurrected by Rico Arce & Bachman, 2006) and Mariosousa

(new genus described in Seigler et al., 2006), (2) two relatively large pan-tropical

genera, Senegalia (resun-ected by Pedley, 1986) and Vachellia (which contains the

original type species of Acacia, A. nilotica (L.) Willd. ex Del.), and (3) the enormous,

predominantly Australian genus, Acacia sensu stricto (syn. Racosperma). Vachellia

is taxonomically well-removed from the other genera of Acacia sensu lato, being

nested within a paraphyletic tribe Mimoseae; as noted by Miller & Seigler (2012)

there are two, well-supported subclades within Vachellia, one of African species and

the other American, to which the type of Vachellia, V farnesiana (L.) Wight & Am.,

belongs. The other four genera are in a paraphyletic grade with genera of tribe Ingeae

(fide Miller & Seigler, 2012). It is this five-genus classification fox Acacia sensu lato

that is generally accepted today. In addition to the phylogenetic evidence based on

nucleotide data, a range of morphological, biochemical, palynological and other data

help support, define and characterise the five genera (see Pedley, 1986; Chappill &

Maslin, 1995; Maslin et al., 2003).

Notwithstanding the above it is possible that further generic segregates will

be recognised within uicacia sensu lato, particularly as more taxa are added to the

genetic datasets. Miller & Seigler (2012) have already suggested a possible new

genus in South America, segregated fi*om Senegalia, to accommodate species allied

to S. skleroxyla (Tussac) Seigler & Ebinger. While major generic realignments are not

anticipated for East and Southeast Asia (but see note below concerning Delaportea

Gagnep.), it is noted that scarcely any species from the region has been included in

existing comparative genetic studies of Acacia sensu lato

Nomenclature of Acacia sensu lato

In July 2011 the Nomenclature Section of the XVII International Botanical Congress

(IBC) in Melbourne, Australia, voted with a clear majority (68%) to accept the Vienna

Code (McNeill et al., 2007) that lists Acacia Mill, with a conserved type (McNeill &

Turland, 2011). This action ended a long debate that began following the publication

234

Card. Bull. Singapore 67(1) 2015

of a formal proposal by Orchard & Maslin (2003) to replace the original type of

Acacia, the African species A. scorpioides (L.) W.F. Wight (=A. nilotica), with a new

type, the Australian species, A. penninei^is Sieber ex DC. A major consequence of the

Melbourne IBC decision is that the ndmQ Acacia now applies to the large, predominantly

Australian group that was formerly called Acacia subgen, Phyllodineae; this group is

referred to here as Acacia sens. str. and the name Racosperma is a synonym of it.

Additionally, the name Vachellia is the correct name for the smaller, pan-tropical group

that was fonnerly called Acacia subgen. Acacia. Further infoiination concerning this

nomenclatural issue is provided on the Worldwide Wattle website (web ref 2).

Combinations for almost all the currently recognised species of Acacia sensu

lato now referable to Acaciella, Mariosousa, Senegal ia and Vachellia have been made,

with only 13 species (10 from Madagascar and three from Africa) outstanding (web

ref 3). The WorldWideWattle website (web ref 4) provides a list of names under

the new classification, and where appropriate, each name is cross-referenced to its

analogue in Acacia when that group was treated as a single genus.

After the Vienna IBC, the ‘new’ generic names replacing Acacia sensu lato had

begun to appear in a range of publications and web databases, and the trend continues

today; however, there seems to have been a lesser uptake of these names by herbaria. As

summarised by Maslin (201 1 ), even prior to the Melbourne IBC the name Vachellia had

been adopted, in lieu of Acacia, in many publications including Flora treatments, field

guides, scientific research papers and books. One particularly important publication

to adopt the new nomenclature early was Mabberley s Plant-book 3^^* ed. (Mabberley,

2008), which is a primaiy reference source for the correct names of vascular plant

genera and families of the world. The new nomenclature has now been adopted in

some large web databases such as the National Center for Biotechnology Infomiation

(web ref. 5) and will be incorporated into the Catalogue of Life database and available

online by mid 2015 (Y. Roskov, pers. comm.). Others web resources such as Tropicos

(web ref 6) and The Plant List (web ref 7) currently adopt a half-way approach by

listing many names as accepted under both Acacia and Vachellia/Senegalia. Some

important legume-centric databases on the web still maintain the old nomenclature,

e.g. the online version of Legumes of the World (web ref 8) and the International

Legume Database & Information Service (ILDIS) (web ref. 9). Furthennore, the

online version of Index Nominum Genericorum (web ref. 10) still lists A. scorpioides

as the type of Acacia. There are of course many possible reasons why the uptake of the

new generic nomenclature has been slow, erratic or has not occurred at all. Resource

and time limitations are two of the more obvious reasons, but some databases may

simply not have the capacity to be changed (ILDIS is presently in this latter category).

In East and Southeast Asia the new names have begun to appear on the web

with Senegalia being listed for both the Philippines (web ref. 11) and for the Hengduan

Mountains region of south-central China (web ref 12). Also, the new generic

nomenclature has been adopted in the checklist of plants of China that is expected

to be soon published and in the checklist of the plants of India that is cun'ently in

preparation and which is expected to appear online during 2015, and published in

hard copy about a year thereafter. Australia has adopted the new nomenclature in its

Synoptic overview oi Acacia sensu lato

235

national names database (web ref. 13) and Australian herbaria have by and large done

likewise with their specimen records (web ref. 14). However, this was a relatively

easy achievement for Australia because so few name and specimen records had to be

changed, with just 1 1 indigenous species being affected (see Table 1).

It is recognised that it often takes a long time for taxonomic and nomenclatural

changes to be accepted and implemented. For herbaria the adoption of such changes

can be especially troublesome not only because of the large amount of work and

resources needed to redetermine specimens and (often) modify database records,

but also because of the disruption caused by having to re-aiTange specimen storage

systems. In the case of Acacia sensu lato these problems are compounded due to

its very many species and global distribution (thus many herbaria are aftected), and

also because of the drawn-out debate over the application of the name Acacia itself

While some people may still wish that change had not occun'ed (a sentiment that

is understandable) the fact remains that the international botanical nomenclatural

community has voted in favour of Acacia retypification. It is therefore appropriate that

the new nomenclature for Acacia sensu lato be adopted universally so that stability can

be provided for this unportant group of legumes. There is now a particular imperative

to achieve this nomenclatural stability because the global legume community is

discussing ways to produce a new phylogenetic classification for the Leguminosae,

one that will incorporate morphological data hito the phytogeny (Bruneau et af, 2013).

One of the basic starting points for this project will be the development of a core list of

taxa; Acacia sensu lato, whose species constitute about a third of the total number for

subfamily Mimosoideae, will be a significant component of that list.

The indigenous species of Acacia sensu lato in East and Southeast Asia

Current evidence shows Acacia sensu lato to be represented in the East and Southeast

Asia by 52 native species accoimiiodated in three genera, namely. Acacia sens,

str., Senegalia and Vachellia, The species are listed, together with their geographic

distributions, in Appendices 1 and 2. However, it should be noted that some modification

to these data can be expected to occur within the context of the review of Acacia sensu

lato for East and Southeast that is currently in progress (Maslin, in prep.).

Acacia sensu stricto

Acacia sensu stricto (1073 species in total) is largely restricted to Australia where

1063 species occur (Table 1), making it the largest genus of vascular plants on that

continent. Only 19 taxa (representing 18 species) occur naturally outside Australia

(Brown et al, 2012), 12 in Asia and seven on islands of the Pacific.

All 12 Asian species are geographically restricted, one occurring in East Asia

{Acacia confusa Merr. from Taiwan and the Philippines) and 1 1 in Southeast Asia.

There are four endemic taxa, three in the south of the region, where Acacia sp. (Wetar)

236

Card. Bull. Singapore 67(1) 2015

and A. wetarensis Pedley are restricted to the small Indonesian island of Wetar and A.

peregrin alls M.W. McDonald & Maslin which is confined to New Guinea, and one in

the northeast (the above mentioned A. confusa). All eight species that extend beyond

the region occur also in Australia. There are no indigenous species of Acacia sensu

stricto in mainland China but a few are cultivated there (fide Wu & Nielsen, 2010).

The indigenous species of Acacia sensu stricto in East and Southeast Asia are

shrubs or trees with phyllodinous foliage (bipinnate foliage occurs on a few introduced

species in Asia). Like species of Vachellia they are generally found in drier habitats

than those of Senegalia.

Some taxa of this genus foiin a significant component of the plantation forestry

industry in parts of Southeast Asia, most notably in Indonesia and Vietnam, where

Acacia crassicarpa A.Cuim. ex Benth., A. mangium Willd. and A. auriculiformis

A.Cunn. ex Benth. x mangium are the main ones grown for pulp and solid wood

products (see Griffin et al., 2011 for overview).

Senegalia

Senegalia (201 species in total) has a pantropical distribution and is represented by 45

species in Asia as a whole (19 in India), 68 in Africa, 102 in the Americas and 2 taxa

in Australia (see Table 1 ).

There are 38 indigenous taxa of Senegalia (representing 32 species) in East

and Southeast Asia, making it by far the largest group of Acacia sensu lato within the

region. Thirteen taxa occur in East Asia with four, Senegalia delevayi (Franch.) Maslin

et al. (van delavayi and van kiinmingensis (C.Chen & H.Sun) Maslin et al.), S. teniana

(Harms) Maslin et al. and S. yunnanensis (Franch.) Maslin et al., endemic in China.

Thirty three taxa occur in Southeast Asia with almost half of them endemic to the sub-

region, namely, Senegalia borneensis (I.C.Nielsen) Maslin et al., S. comosa (Gagnep.)

Maslin et al., S. donnaiensis (Gagnep.) Maslin et al., S. kekapur (I.C.Nielsen) Maslin

et al., S. kostermansii (I.C.Nielsen) Maslin et al., S, meeboldii (Craib) Maslin et al.,

S. merrillii (I.C.Nielsen) Maslin et al., S. pala\\>anensis (I.C.Nielsen) Maslin et al.,

S. pluricapitata (Steud. ex Benth.) Maslin et al., S. pluriglandulosa (Verde.) Maslin

et al., S. pseiidointsia (Miq.) Maslin et al., S. sulitii (I.C.Nielsen) Maslin et al., S.

tawitaMdensis (I.C.Nielsen) Maslin et al., S. thailandica (I.C.Nielsen) Maslin et al. and

S, verheijenii (I.C.Nielsen) Maslin et al. These endemic taxa are scattered generally

tliroughout the sub-region, occurring in all countries except Brunei (which has no

indigenous Acacia sensu lato recorded). Indonesia has the highest concentration of sub-

regional endemics with nine species recorded, but only Senegalia kostermansii and S.

verheijensii are restricted to that country. Almost all of the 16 species of Senegalia that

extend beyond the region range westward to India and nearby countries, with 10 also

ranging north to China. Only one taxon, Senegalia pennata subsp. kerrii (I.C.Nielsen)

Maslin et al., extends to Australia, which has a very poor representation of indigenous

species of Senegalia (just two taxa, the endemic S, albizioides (Pedley) Pedley and S,

pennata subsp. kerrii).

Synoptic overview oi Acacia sensu lato

237

Species of Senegalia are characterised by having bipinnate leaves and cauline

prickles. Most of the indigenous species of East and Southeast Asian are woody

lianes with only the widespread Senegalia catechu (L.f.) P.J.H.Hurter & Mabb., S.

chundra (Roxb. ex Rottler) Maslin and S. ferruginea (DC. ) Pedley (all of which have

prickles at the nodes) and the geographically restricted, endemic S. kostermansii

from Indonesia and S. teniana from China (both of which have scattered prickles)

seemingly obligate shrubs or trees which lack scandent branches. Most species have

broad, flat, ±chartaceous and straight pods but in Senegalia thailandica the pods

are slightly inflated and tightly curled. These species can be arranged in four sub-

groups (defined principally by prickle distinbution and leaflet characteristics) centred

on Senegalia pennata, S. caesia, S. andamanica (l.C.Nielsen) Maslin et al and S.

catechu respectively. The one species with very different caipological features is

the widespread and variable Senegalia rugata (Lam.) Britton & Rose (syn. Acacia

concinna (Willd.) DC.). In this species the pods are smooth, thick and fleshy when

fresh but they dry characteristically wrinkled, blackish and with a very hard texture.

This species name has about 10 heterotypic synonyms and fiirther study is needed to

reassess the taxonomic status of Senegalia rugata. Senegalia albizioides (restricted to

Australia) is currently the only recognised close relative of S. rugata.

Vachellia

Vachellia (163 species in total) has a pantropical distribution and is represented by

30 species in Asia as a whole (11 in India), 84 in Africa, 57 in the Americas and 8 in

Australia (see Table 1).

There are just eight indigenous species of Vachellia in Southeast Asia (none

occurs in East Asia), making it the least well-represented of the three genera within

the region. There are five endemic species, all of which occur m northern areas, three

restiicted to Myanmar {Vachellia inopinata (Prain) Maslin et al., V. kingii (Prain)

Maslin et al. and V. myaingii (Lace) Maslin et al.: all these species are poorly known

and require study to reassess their taxonomic status), one to Thailand {V siamensis

(Craib) Maslin et al.) and one widespread {V. harmandiana (Pierre) Maslin et al.

which occurs in Cambodia, Laos, Thailand, Vietnam). The three species that extend

beyond the region, namely, Vachellia nilotica subsp. indica (Benth.) Kyal. & Boatwr.,

V leucophloea (Roxb.) Maslin et al. and V tomentosa (Rottler) Maslin et al., range

westward to India with V. nilotica subsp. indica extending to the Middle East.

Excluding Vachellia harmandiana and V leucophloea, the only indigenous species of

Vachellia that is widespread in Southeast Asia is V. tomentosa.

Gagnepain (1911, 1952) and Craib (1927) described species under new generic

names, Delaportea and Nimiria Prain ex Craib respectively. At present these species

are treated as conspecific with Vachellia harmandiana, V. leucophloea and V. siamensis.

They appear to fonn a natural group that is recognised by an unusual combination of

characters, namely, stipules spinescent (characteristic of Vachellia) and heads arranged

in open, terminal panicles (characteristic of many species of Senegalia). Further field,

238

Card. Bull. Singapore 67(1) 2015

morphological and genetic study is needed to reassess the taxonomic status of this

group and to determine if it is appropriate to resurrect the genus Delaportea (with

Nimiria as a synonym),

Vachellia farnesiana (L.) Wight & Am. has been recorded as an introduction

(sometimes naturalised) in a number of countries of East and Southeast Asia (see

Nielsen, 1981, 1985b, 1992; Wu & Nielsen, 2010). It is a native of tropical America

and comprises three varieties (Seigler & Ebinger, 2005). Although it is not known

what varieties occur in Asia it is most likely the typical one (D. Seigler, pers. comm.).

The indigenous species of Vachellia in Southeast Asia are shrubs or trees

characterised by having bipinnate foliage subtended by spiny stipules. Like species of

Acacia sens. str. they are generally found in drier sites than those of Senegalia.

ACKNOWLEDGEMENTS: Prof Phan Ke Loc (Vietnam National Herbarium, Hanoi) is

thanked for discussions that provided the motivation for me to prepare this communication.

The following colleagues are thanked for providing constructive comments on the manuscript:

Prof David Mabberley (Sydney), Dr Kevin Thiele (W.A. Herbarium, Perth) and Dr Boon-

Chuan Ho (Singapore Botanic Gardens). The following herbaria are thanked for allowing

me access to their collections: Herbarium Bogoriense, Cibinong, Indonesia (BO); Bangkok

Herbarium, Chatuchak, Thailand (BK); The Forest Herbarium Bangkok, Chatuchak, Thailand

(BKF ); Chiang Mai University Herbarium, Thailand (CMU); Hanoi Herbarium, National Center

for Natural Sciences and Technology, Hanoi, Vietnam (HN); Vietnam National Herbarium,

Hanoi University, Hanoi, Vietnam (HNU); Forest Research Institute Malaysia Herbarium,

Kuala Lumpur, Malaysia (KEP); Kunming Institute of Botany, Chinese Academy of Sciences

Herbarium, Kumning, China (KUN); Singapore Botanic Gardens Herbarium, Singapore

(SING); Taiwan Forestry Institute, Taipei (TAIF); Institute of Tropical Biology Herbarium, Ho

Chi Minh City, Vietnam (VNM).

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241

2. Acacia Name Issue. http://www.worldwidewattle.com/infogalleiy/nameissLie/index.php.

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Appendix 1. Indigenous taxa of Acacia sensu lato currently recognised for East and Southeast

Asia.

See Introduction for definition of the terms East and Southeast Asia. Only selected synonyms

are given in this list.

ACACIA Mill., Gard. Diet. Abr., ed. 4, [25] (1754), nom. cons. TYPE: Acacia penninervis

Sieber ex DC. {typ. cons., vide Regnum Veg. 146: App. Ill, 286. 2006).

Racosperma (DC.) Mart., Hort. Reg. Monac. Semin. 4 (1835). LECTOTYPE: Racosperma

pennineiA^e (Sieber ex DC.) Pedley, Bot. J. Linn. Soc. 92: 239 (1986). Based on Acacia sect.

Phyllodineae DC., Prodr. 2: 448 (1825).

242

Card. Bull. Singapore 67(1) 2015

Acacia aur iculif or mis A.Cmm. ex Benth., London J. Bot. 1: 377 (1842).

Distribution. Southeast Asia: Indonesia (Moluccas), Papua New Guinea. Australia: Northern

Territory & Queensland.

Introduced/cultivated/naturalised. Americas (Florida, Panama), Africa, Bangladesh, Brunei,

China, India (inch Andaman Is.), Indonesia, Laos, Malaysia, Myanmar, Nepal, Pakistan,

Philippines, Singapore, Sri Lanka, Vietnam.

Acacia confusa Men*., J. Sci. (Bot.) 5: 27 (1910).

Distribution. East Asia: Taiwan. Southeast Asia: Philippines.

Cultivated/introduced. China, Hawaiian Islands, India, Indonesia, Japan (southern Ryuk5m

Islands), Mauritius, Malaysia, Northern Marianas, Seychelles, Vietnam.

Acacia crassicarpa A.Cunn. ex Benth., London J. Bot. 1: 379 (1842).

Distribution. Southeast Asia: Papua New Guinea. Australia: Queensland.

Cultivated. Australia (Northern Territory), Vietnam.

Acacia leptocarpa A.Cunn. ex Benth., London J. Bot. 1: 376 (1842).

Distribution. Southeast Asia: Papua New Guinea. Australia: Northern Territory, Queensland,

Western Australia.

Acacia mangium Willd., Sp. PL, ed. 4, 4(2): 1053 (1806).

Distribution. Southeast Asia: Indonesia (New Guinea, Moluccas), Papua New Guinea.

Australia: Queensland.

Introduced/naturaUsed. Americas (Nicaragua), Australia (Northern Temtory), Bangladesh,

Brunei, China, Central America (Nicaragua), hidia, Philippines, Singapore, Vietnam.

Acacia oraria F.Muell., Fragm. 11: 66 (1 879).

Distribution. Southeast Asia: Indonesia (Flores, Timor). Australia: Queensland.

Acacia peregrinalis M.W.McDonald & Maslin, Nuytsia 14(3): 455 (2002).

Distribution. Southeast Asia: Indonesia (New Guinea), Papua New Guinea.

Acacia pubirhachisVQdlQy, Contr. Queensland Herb. 15: 15 (1974).

Distribution. Southeast Asia: Papua New Guinea. Australia: Queensland.

Acacia simsii A.Cunn. ex Benth., London J. Bot. 1: 368 (1842).

Distribution. Southeast Asia: Indonesia (New Guinea), Papua New Guinea. Australia:

Northern Territoiy, Queensland.

Acacia spirorbis subsp. solandri (Benth.) Pedley, Austrobaileya 3: 216 (1990).

Distribution. Southeast Asia: Papua New Guinea. Australia: Queensland.

Acacia wetarensis ?Qd\Qy, Contr. Queensland Herb. 18: 18 (1975).

Distribution. Southeast Asia: Indonesia (Wetar).

Acacia sp. (Wetar).

Distribution. Southeast Asia: Indonesia (Wetar).

Note. This entity is noted by Nielsen (1992: 59) [inder Acacia leptocarpa. It appears close to

the glabrous variant of A. elachantha M.W.McDonald & Maslin (McDonald & Maslin, 1997),

and preliminary studies suggest that it may represent a distinct taxon.

Synoptic overview oi Acacia sensu lato

243

SENEGALIA Raf., Sylva Tell. 1 19 (1838). LECTOTYPE: Mimosa Senegal L. {vide Britton &

Rose, N. Amer. FI. 23: 106. 1928) {Senegalia Senegal (L.) Britton [Senegalia triacantha Raf.,

nom. illeg.])

Senegalia andamanica (l.C.Nielsen) Maslin, Seigler & Ebinger, Blumea 58: 40 (2013). -

Acacia andamanica l.C.Nielsen, Adansonia ser 2, 19: 354 (1980).

Distribution. Southeast Asia: Thailand. South Asia: Andaman Islands, Nicobar Islands.

Senegalia borneensis (l.C.Nielsen) Maslin, Seigler & Ebinger, Blumea 58: 40 (2013). -Acacia

borneensis l.C.Nielsen, Opera Bot. 81: 20, fig. 9 (7-13) (1985).

Distribution. Southeast Asia: Indonesia (Kalimantan), Malaysia (Sabah).

Senegalia caesia (L.) Maslin, Seigler & Ebinger, Blumea 58: 40 (2013). - Mimosa caesia E.,

Sp. PI. 1: 522 (1753). -Acacia caesia (L.) Willd., Sp. PI., ed. 4, 4(2): 1090 (1806).

Distribution. East Asia: China (Guangdong, Hainan, Sichuan, Yunnan), Taiwan. Southeast

Asia: Cambodia, Laos, Myamnar, Thailand, Vietnam. South Asia: Bangladesh, Bhutan, India,

Sri Lanka.

Note. Although Acacia caesia van subnuda (Craib) l.C.Nielsen is recognised by Nielsen

(1981 : 53) and Sanjappa (1 992: 37), it is regarded as conspecific with .^4. caesia by Chakrabarty

& Gangopadhyay (1996: 604) and Wu & Nielsen (2010: 57). Maslin et al. (2013) did not

recognise the variety in Senegalia. Further study is required to reassess the taxonomic status

of this variety.

Senegalia catechu (L.f ) P.J.H.Hurter & Mabb. in Mabberley’s Plant-book ed. 3, 1021 (2008).

-Mimosa catechu L.f, Suppl. PI. 439 (1782 [1781 publ. Apr. 1782]). - Acacia catechu (L.f.)

Willd., Sp. PI., ed. 4, 4(2): 1079 (1806).

Distribution. East Asia: China (Yunnan). Southeast Asia: Myanmar, Thailand (uncertain, fide

Lock & Heald 1994: 33). South Asia: Bangladesh, Bhutan, India, Nepal, Pakistan, Sri Lanka.

Cultivated/introduced. China (Fujian, Guangdong, Guangxi, Hainan, Yunnan, Zhejiang),

Indonesia (Java), Japan (southern Ryukyu Islands), Mauritius, Philippines, Taiwan, Vietnam.

Senegalia chiindra (Roxb. ex Rottler) Maslin, Nuytsia 22(6): 466 (2012). - Mimosa chundra

Roxb. ex Rottler, Neue Schriften Ges. Naturf. Freunde Berlin 4: 207 (1803). - Acacia chundra

(Roxb. ex Rottler) Willd., Sp. PI. ed. 4, 4(2): 1078 (1806).

Acacia sundra (Roxb.) DC., Prodr. 2: 458 (1825). - Mimosa siindra Roxb., PI. Corom. 3: 19,

tab. 225 (1811).

Distribution. Southeast Asia: Myamnar. South Asia: India, Sri Lanka.

Introduced/naturalised. Australia: Northern Territory, Reunion Island.

Note. Despite their similar-sounding epithets. Mimosa chundra andM sundra were independent

descriptions based on different types,2i4fe Maslin et al. (2013: 466). The complex nomenclatural

history involving these names is discussed by Kshirsagar (web ref. 15).

Senegalia comosa (Gagnep.) Maslin, Seigler & Ebinger, Blumea 58: 40 (2013). - Acacia

comosa Gagnep., Notul. Syst. (Paris) 2: 113 (1911).

Distribution. Southeast Asia: Cambodia, Laos, Thailand, Vietnam.

Senegalia delavayi (Franch.) Maslin, Seigler & Ebinger, Blumea 58: 40 (2013). - Acacia

delavayi Franch., PI. Delavay. 194 (1890).

Note. Two varieties recognised; both endemic to China.

244

Card. Bull. Singapore 67(1) 2015

Senegalia delavayi (Fraiich.) Maslin, Seigler & Ebinger van delavayi - Acacia delavayi

Franch. van delavayi established by publication of A. delavayi van kunmingensis C.Chen &

H. Sun, Acta Bot. Yunnan. 12: 262 (1990).

Distribution. East Asia: China (Yunnan).

Senegalia delavayi van kunmingensis (C.Chen & H.Sun) Maslin, Seigler & Ebinger, Blumea

58: 40 (2013). - Acacia delavayi van kunmingensis C.Chen & H.Sun, Acta Bot. Yunnan. 12:

262 (1990).

Distribution. East Asia: China (Guizhou, Yunnan).

Senegalia donnaiensis (Gagnep.) Maslin, Seigler & Ebinger, Blumea 58: 40 (2013). - Acacia

donnaiensis Gagnep., Not. Syst. (Paris) 2: 114 (1911).

Distribution. Southeast Asia: Cambodia, Indonesia (Kalimantan), Malaysia (Sabah), Vietnam.

Senegalia ferntginea (DC.) Pedley, Bot. J. Linn. Soc. 92: 250 (1986). - Acacia ferruginea

DC., Prodr. 2: 458 (1825).

Distribution. Southeast Asia: Myanmar. South Asia: India, Sri Lanka.

Senegalia gageana (Craib) Maslin, Seigler & Ebinger, Blumea 58: 40 (2013). - Acacia

gageana Craib, Bull. Misc. Inform. Kew 1915: 409 (1915).

Distribution. Southeast Asia: Myanmar. South Asia: Bangladesh, Bhutan, India, Nepal,

Pakistan.

Senegalia intsia (L.) Maslin, Seigler & Ebinger, Blumea 58: 40 (2013). -Mimosa intsia L., Sp.

PI. 1: 522 (1753). -Acacia intsia (L.) Willd., Sp. PL, ed. 4, 4(2): 1091 (1806).

Distribution. Southeast Asia: Myanmar. South Asia: Bangladesh, India, Nepal, Sri Lanka.

Senegalia kekapur (I.C.Nielsen) Maslin, Seigler & Ebinger, Blumea 58: 40 (2013). - Acacia

kekapur I.C.Nielsen, Opera Bot. 81: 13, fig. 5 (1-8) (1985).

Distribution. Southeast Asia: Indonesia (Java, Sumatra), Singapore.

Note. Nielsen (1985a) noted that plants from NE Sumatra and Singapore may be taxonomically

different from S. kekapur sens. typ. from Java and southern Sumatra; this matter is currently

under investigation.

Senegalia kostermansii (I.C.Nielsen) Maslin, Seigler & Ebinger, Blumea 58: 40 (2013). -

Acacia kostermansii I.C.Nielsen, Opera Bot. 81:15 (1985).

Distribution. Southeast Asia: Indonesia (Alor Island, Sumbawa, Flores).

Senegalia meeboldii (Craib) Maslin, Seigler & Ebinger, Blumea 58: 40 (2013). - Acacia

meeboldii Craib, Bull. Misc. Infonn. 1928: 66 (1928).

Distribution. Southeast Asia: Myanmar, Thailand.

Senegalia megaladena (Desv.) Maslin, Seigler & Ebinger, Blumea 58: 41 (2013). - Acacia

megaladena Desv., J. Bot. Agric. 3: 69 (1814).

Note. Three varieties recognised.

Senegalia megaladena (Desv.) Maslin, Seigler & Ebinger var. megaladena - Acacia

megaladena Desv. var. megaladena established by publication of 4. megalodena var. garrettii

I. C. Nielsen and var. indo-chinensis I.C. Nielsen, Adansonia ser. 2, 19: 351 (1980).

Distribution. East Asia: China (Guangxi, Yunnan). Southeast Asia: Indonesia (Java), Laos,

Myanmar, Thailand, Vietnam. South Asia: Bangladesh, India, Nepal.

Synoptic overview oi Acacia sensu lato

245

Senegalia megaladena var. indochinensis (I.C.Nielsen) Maslin, Seigler & Ebinger, Blumea

58: 41 (2013). - Acacia megaladena var. indochinensis I.C.Nielsen, Adansonia ser. 2, 19: 351

(1980).

Distribution. Southeast Asia: Cambodia, Indonesia (Java, but probably introduced according

to Nielsen 1985a: 26), Laos, ?Malaysia (recorded for northern Peninsular Malaysia by Nielsen

1985a: 26 & 1992: 51, but not listed in Turner 1997: 290), Thailand, Vietnam.

Senegalia megaladena var. garrettii (I.C.Nielsen) Maslin, Seigler & Ebinger, Blumea 58: 41

(20X2)). - Acacia megaladena var. garrettii I.C.Nielsen, Adansonia sen 2, 19: 351 (1980).

Distribution. East Asia: China (Guangxi, Yunnan). Southeast Asia: Thailand.

Senegalia merrillii (I.C.Nielsen) Maslin, Seigler & Ebinger, Blumea 58: 41 (2013). - Acacia

merrillii I.C.Nielsen, Opera Bot. 81: 9, fig. 2 (1985).

Distribution. Southeast Asia: Indonesia (Moluccas, Sulawesi), Philippines.

Senegalia palawanensis (I.C.Nielsen) Maslin, Seigler & Ebinger, Blumea 58: 41 (2013). -

Acacia palawanensis I.C.Nielsen, Opera Bot. 81: 16, fig. 6 (7-11) (1985).

Distribution. Southeast Asia: Philippines.

Senegalia pennata (L.) Maslin, Nuytsia 22(6): 466 (2012). -Mimosa pennata L., Sp. PI. 1 : 522

(1753). -Acacia pennata (L.) Willd., Sp. PI, ed. 4, 4(2): 1090 (1806).

Note. Four subspecies recognised.

Senegalia pennata subsp. pennata. - Acacia pennata (L.) Willd. subsp. pennata established

by publication of A. pennata subspp. hainanensis (Hayata) I.C. Nielsen, insuavis (Lace)

I.C.Nielsen and kerri I.C.Nielsen, Adansonia ser. 2, 19(3): 352-353 (1980).

Distribution. Southeast Asia: Myanmar, Thailand (probably introduced). South Asia:

Bangladesh, Bhutan, India, Nepal, Sri Lanka.

Senegalia pennata subsp. hainanensis (Hayata) Maslin, Seigler & Ebinger, Blumea 58: 41

(2013). — Acacia hainanensis Hayata, Ic. PI. Fonuos. 3: 86 (1913). - Acacia pennata subsp.

hainanensis (Hayata) I.C. Nielsen, Adansonia ser. 2, 19(3): 352 (1980).

Distribution. East Asia: China (Fujian, Guangdong, Guangxi, Hainan, Yunnan). Southeast

Asia: My amnar, Vietnam. South Asia: India.

Senegalia pennata subsp. insuavis (Lace) Maslin, Seigler & Ebinger, Blumea 58: 41 (2013). -

Acacia insuavis Lace, Bull. Misc. Infonn. 1915: 401 (1915). -Acacia pennata subsp. insuavis

(Lace) I.C.Nielsen, Adansonia ser. 2, 19(3): 353 (1980).

Distribution. Southeast Asia: Cambodia, Laos, Myanmar. South Asia: India.

Introduced/cultivated. USA (Florida), Australia (Northern Territory, Queensland), Singapore,

Thailand.

Note. Pedley (2014) treated this entity as S. insuavis (Lace) Pedley; however, its taxonomic

status is currently under investigation by the present author.

Senegalia pennata subsp. ken ii (I.C.Nielsen) Maslin, Nuytsia 22(6): 467 (2012).

Acacia pennata subsp. kerrii I.C.Nielsen, Adansonia ser. 2, 19(3): 353 (1980).

Distribution. Australia: Queensland. East Asia: China (Yunnan). Southeast Asia: Cambodia,

East Timor, Indonesia (Java, Sulawesi, Flores, Komodo, Lombok, Sumbawa), Laos, Malaysia

(Peninsular Malaysia), Myanmar, Thailand, Vietnam. South Asia: Bhutan, India, Nepal, Sri

Lanka.

246

Card. Bull. Singapore 67(1) 2015

Senegalia pluricapitata (Steud. ex Benth.) Maslin, Seigler & Ebinger, Blumea 58: 41 (2013).

Acacia pluricapitata Steud. ex Benth., London. J. Bot. 1:516 (1842).

Distribution. Southeast Asia: Indonesia (Sumatra, Java, Kalimantan), Malaysia (Peninsular

Malaysia, Sabah), ?Myanmar, Philippines, Thailand, Vietnam.

Senegalia pUiriglandulosa (Verde.) Maslin, Seigler & Ebinger, Blumea 58: 41 (2013). -

Acacia phiriglandulosa Verde., Kew Bull. 32: 472 (1978).

Distribution. Southeast Asia: Indonesia (West Papua), Papua New Guinea, Philippines.

Senegalia pruinescens (Kurz) Maslin, Seigler & Ebinger, Blumea 58: 41 (2013). - Acacia

pruinescens Kurz, J. Asiat. Soc Bengal, Pt. 2, Nat. Hist. 45: 296, 298 (1877).

Distribution. East Asia: China (Guangxi, Yunnan). Southeast Asia: Myanmar, Vietnam.

South Asia: India.

Senegalia pseudointsia (Miq.) Maslin, Seigler & Ebinger, Blumea 58: 41 (2013). - Acacia

pseudointsia Miq., FI. Ned. Ind. 1: 12 (1855), as ’"Pseudo-Intsia.

Distribution. Southeast Asia: Indonesia (Java, Sumatra), Malaysia (Peninsular Malaysia,

Sabah, Sarawak), Thailand.

Senegalia rugata (Lam.) Britton & Rose, N. Amer. FI. 23(2): 120 (1928). - Mimosa rugata

Lam., Encycl. 1: 20 (1783).

Acacia concinna (Willd.) DC., Prodr. 2: 464 (1 825). - Mimosa concinna Willd., Sp. PI., ed. 4,

4(2): 1039(1806).

Distribution. East Asia: China (Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hunan,

Jiangxi, Yunnan). Southeast Asia: Cambodia, Indonesia (Ambon, Java, Kai Is., Sumba, Flores,

Moluccas, Sulawesi, Sumatra), Laos, Malaysia (Peninsular Malaysia), Myamnar, Papua

New Guinea, Philippines, Thailand, Vietnam. South Asia: Bangladesh, Bhutan, India (inch

Andaman Is.), Nepal.

Introduced. Australia (Queensland), Japan (Okinawa), Madagascar, Mauritius, Reunion Island.

Note. A variable species in need of critical revision.

Senegalia sulitii (I.C.Nielsen) Maslin, Seigler & Ebinger, Blumea 58: 42 (2013). - Acacia

sulitii I.C.Nielsen, Opera Bot. 81: 24, fig. 10 (6-10) (1985).

Distribution. Southeast Asia: Indonesia (Sulawesi), Philippines.

Senegalia tawitawiensis (I.C.Nielsen) Maslin, Seigler & Ebinger, Blumea 58: 42 (2013). -

Acacia tawitawiensis I.C.Nielsen, Opera Bot. 81: 22, fig. 10 (1-5) (1985).

Distribution. Southeast Asia: Philippines.

Senegalia teniana (Hanns) Maslin, Seigler & Ebmger, Blumea 58: 42 (2013). -Acacia teniana

Harms, Repert. Spec. Nov. Regni Veg. 17: 133 (1921),

Distribution. East Asia: China (Yunnan. Sichuan),

Senegalia thailandica (I.C.Nielsen) Maslin, Seigler & Ebinger, Blumea 58: 42 (2013). -

Acacia thailandica I.C.Nielsen, Adansonia ser. 2, 19: 356, pi. 1 (1980).

Distribution. Southeast Asia: Cambodia, Thailand.

Senegalia tonkinensis (I.C.Nielsen) Maslin, Seigler & Ebinger, Blumea 58: 42 (2013). -

Acacia tonkinensis I.C.Nielsen, Adansonia ser. 2, 19: 358, pi. 2 (1980).

Distribution. East Asia: China (Yunnan). Southeast Asia: Laos, Thailand, Vietnam.

Synoptic overview oi Acacia sensu lato

247

Senegalia torta (Roxb.) Maslin, Seigler & Ebinger, Blumea 58: 42 (2013). - Mimosa torta

Roxb., FI. Ind., ed. 2, 2: 566 (1832). -Acacia torta (Roxb.) Craib, Kew Bull. 1915: 410 (1915).

Distribution. Southeast Asia: ?Thailand. South Asia: India, Pakistan.

Note. It is unlikely that this species occurs in Southeast Asia. The Thailand records by Nielsen

(1985b: 167) are probably Senegalia tonkinensis (fide Srisanga & Sasirat, 2000).

Senegalia verheijenii (I.C.Nielsen) Maslin, Seigler & Ebinger, Blumea 58: 42 (2013). -Acacia

verheijenii I.C.Nielsen, Opera Bot. 81: 16, fig. 6 (1-6) (1985).

Distribution. Southeast Asia: hidonesia (Flores).

Senegalia vietnamensis (I.C.Nielsen) Maslin, Seigler & Ebinger, Blumea 58: 42 (2013). -

Acacia vietnamensis I.C.Nielsen, Adansonia ser. 2, 19: 360, pi. 3 (1980).

Distribution. Southeast Asia: Laos, Vietnam. (Erroneously recorded for China by Wu &

Nielsen 2010: 57).

Senegalia yunnanensis (Franch.) Maslin, Seigler & Ebinger, Blumea 58: 42 (2013). - Acacia

yunnanensis Franch., PI. Delavay. 193 (1890).

Distribution. East Asia: China (Yunnan, Sichuan).

VACHELLIA Wight & Am., Prodr. 272 (1834). TYPE: Vachellia farnesiana (L.) Wight &

Arn.

Delaportea Thorel ex Gagnep., Notul. Syst. (Paris) 2: 117 (1911). TYPE: Delaportea armata

Thorel ex Gagnep. (= Vachellia harmandiana (Pierre) Maslin, Seigler & Ebinger).

Nimiria Prain ex Craib, Bull. Misc. Infonn. 1927: 393 (1927). TYPE: Nimiria siamensis Craib

(= Vachellia siamensis (Craib) Maslin, Seigler & Ebinger).

Vachellia harmandiana (Pierre) Maslin, Seigler & Ebinger, Blumea 58: 42 (2013). -

Pithecolobium harmandianum Pierre, FI. Forest. Cochinch. 5: tab. 394A (1899). - Acacia

harmandiana (Pierre) Gagnep., Not. Syst. (Paris) 2: 115 (1911). - Delaportea armata Thorel

ex Gagnep., Not. Syst. (Paris) 2: 118 (1911).

Distribution. Southeast Asia: Cambodia, Laos, Thailand, Vietnam.

Vachellia inopinata (Prain) Maslin, Seigler & Ebinger, Blumea 58: 42 (2013). - Acacia

inopinata Prain, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 66: 507 (1897). - Nimaria inopinata

(Prain) Craib, Bull. Misc. Infonn. Kew 1927: 393 (1927).

Distribution. Southeast Asia: Myamnar.

Vachellia kingii (Prain) Maslin, Seigler & Ebinger, Blumea 58: 42 (2013). - Acacia kingii

Prain, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 66: 506 (1897).

Distribution. Southeast Asia: Myamnar.

Vachellia leiicophloea (Roxb.) Maslin, Seigler & Ebinger, Blumea 58: 42 (2013). - Mimosa

leucophloea Roxb., PI. Coromandel 2: 27, tab. 150 (1800). - Acacia leucophloea (Roxb.)

Willd., Sp. PI., ed. 4, 4(2): 1083 (1806).

Delaportea microphylla Gagnep., Bull. Soc. Bot. Fr. 99: 46 (1952).

Delaportea ferox Gagnep., Bull. Soc. Bot. Fr. 99: 47 (1952).

248

Card. Bull. Singapore 67(1) 2015

Distribution. Southeast Asia: East Timor, Indonesia (Java, Bali, Timor), Laos, Malaysia

(Peninsular Malaysia), Myanmar, Thailand, Vietnam. South Asia: India, Pakistan, Sri Lanka.

Introduced. Mauritius, Trinidad.

Note. Maslin et al. (20 1 3 : 42-43) recognise two varieties, VacheUia leucophloea var. leucophloea

and var. microcephcda (Kurz) Maslin, Seigler & Ebinger (recorded only for Myamnar). The

taxonomic status of VacheUia leucophloea var. microcephala, however, needs to be reviewed.

VacheUia myaingii (Lace) Maslin, Seigler & Ebinger, Blumea 58: 43 (2013). -Acacia myaingii

Lace, Bull. Misc, Infonn. 1915: 114 (1915), as “Myaingii”.

Distribution. Southeast Asia: Myanmar.

VacheUia nilotica (L.) P.J.H. Hurter & Mabb. subsp. indica (Benth.) Kyal. & Boatwr., Bot. J.

Linn. Soc. 172: 515 (2013). -Acacia arabica var. indica Benth., London J. Bot. 1: 500 (1842).

-Acacia nilotica subsp. indica (Benth.) Brenan, Kew Bull. 12: 84 (1957).

Distribution. Southeast Asia: Myanmar. South Asia: Bangladesh, India, Nepal, Pakistan.

West Asia: Iran, North Yemen, Oman, Qatar, South Yemen.

Introduced/naturalised. Africa, America, Australia (Northern Ten'itoiy, Queensland, South

Australia, Western Australia), India (Andaman Is,), Indonesia (Timor), Sri Lanka.

VacheUia siamensis (Craib) Maslin, Seigler & Ebinger, Blumea 58: 43 (2013). - Nimiria

siamensis Craib, Bull. Misc. Inform. 1927: 393 (1 927). -Acacia craibii I.C.Nielsen, Adansonia

ser.2, 19(3): 344 (1980).

Distribution. Southeast Asia: Thailand.

VacheUia tomentosa (Rottler) Maslin, Seigler & Ebinger, Blumea 58: 43 (2013). - Mimosa

tomentosa Rottler, Ges. Naturf. Freunde Berlin Neue Schriften 4: 208 (1803). - Acacia

tomentosa Willd. Sp. PL, ed. 4, 4(2): 1087 (1806).

Distribution. Southeast Asia: East Timor, Indonesia (Java, Sumba, Sumbawa, Sulawesi),

Myanmar, Thailand, Vietnam. South Asia: Bangladesh, India, Sri Lanka.

Appendix 2. Indigenous taxa of Acacia sensu lato in East and Southeast Asia, listed by

country of occurrence.

See Introduction for definition of the tenns East and Southeast Asia. E = endemic (additional

to the 1 5 countiy-specific endemics shown below there are 1 0 species of Senegalia that are

endemic to Southeast Asia which occur in more than one country, see text above).

East Asia

China

10 species

(13 taxa)

Senegalia caesia, S. catechu, S. delavayi (var. delavayi

& var. kunmingensis) [both E], S. megaladena (var.

megaladena & var. garrettii), S. pennata (subsp. hainanensis

& subsp. kerrii), S. pniinescens, S. rugata, S. teniana [E], S.

tonkinensis, S. yunnanensis [E].

Taiwan

2 species

Acacia confusa. Senegalia caesia.

Southeast Asia

Brunei

No indigenous species recorded.

Synoptic overview oi Acacia sensu lato

249

Cambodia

8 species

(9 taxa)

Senegalia caesia, S. comosa, S. donnaiensis, S. megaladena

van indochinensis, S. pennata (subsp. insuavis. & subsp.

kerrii), S. rugata, S. thailandica. Vachellia harmandiana.

East Timor

3 species

Senegalia pennata subsp. kerrii. Vachellia leucophloea, V.

tomentosa.

Indonesia

21 species

(22 taxa)

Acacia auriculiformis, A. mangium, A. oraria, A.

peregrinalis,A. wetarensis [E],4. sp. (Wetar) [E], Senegalia

borneensis, S. donnaiensis, S. kekapur, S. kostermansii

[E], S. megaladena (van megaladena & var. indochinensis

- probably introduced, see note above), 5*. merrillii, S.

pennata subsp. kerrii, S. pliiricapitata, S. pluriglandulosa,

S. pseudointsia, S. rugata, S. sulitii, S. verheijensii [E].

Vachellia leucophloea, V tomentosa.

Laos

9 species

(11 taxa)

Senegalia caesia, S. comosa, S. megaladena (var.

indochinensis & var. megaladena), S. pennata (subsp.

insuavis. & subsp. kerrii), S. rugata, S. tonkinensis, S.

vietnamensis. Vachellia harmandiana, V leucophloea.

Malaysia

8 species

Senegalia borneensis, S. donnaiensis, S. megaladena var.

indochinensis (uncertain, see note above), S. pennata subsp.

kerrii, S. pluricapitata, S. pseudointsia, S. rugata. Vachellia

leucophloea.

Myanmar

18 species

(21 taxa)

Senegalia caesia, S. catechu, S. chundra, S. ferruginea,

S. gageana, S. intsia, S. rneeboldii, S. megaladena var.

megaladena, S. pennata (subsp. hainanensis, subsp.

in.suavis, subsp. kerrii & subsp. pennata), S. pluricapitata

(uncertain), S. pruinescens, S. rugata. Vachellia inopinata

[E], V. kingii [E], V. leucophloea, V. myaingii [E], V. nilotica

subsp. indica, V. tomentosa.

Papua New

Guinea

10 species

Acacia auriculiformis, A. crassicarpa, A. leptocarpa.

Acacia mangium, A. peregrinalis, A. pubirhachis, A. simsii,

A. spirorbis subsp. solandri. Senegalia pluriglandulosa, S.

rugata.

Philippines

8 species

Acacia confusa. Senegalia merrillii, S. palawanensis [E], S.

pluricapitata, S. pluriglandulosa, S. rugata, S. sulitii [E], S.

tawitawiensis [E].

Singapore

1 species

Senegalia kekapur.

Thailand

17 species

(20 taxa)

Senegalia andamanica, S. caesia, S. catechu (uncertain), S.

comosa, S. rneeboldii, S. megaladena (var. indochinensis,

var. megaladena & var. garrettii), S. pennata (subsp. kerrii

& subsp. pennata), S. pluricapitata, S. pseudointsia, S.

rugata, S. thailandica, S. tonkinensis, S. torta (unlikely,

see note above). Vachellia harmandiana, V. leucophloea, V.

siamensis [E], V. tomentosa.

Vietnam

13 species

(15 taxa)

Senegalia caesia, S. comosa, S. donnaiensis, S. megaladena

(van indochinensis, van megaladena), S. pennata (subsp.

hainanensis and subsp. kerrii), S. pluricapitata, S.

pruinescens, S. rugata, S. tonkinensis, S. vietnamensis.

Vachellia harmandiana, V. leucophloea, V. tomentosa.

250

Card. Bull. Singapore 67(1) 2015

Table 1. Classification schemes for Acacia sensu lato showing species numbers and major

areas of occurrence.

Column 3 (in bold) gives the current classification and nomenclature of Acacia sensu lato.

Species numbers are sourced from the Worldwide Wattle website (web ref 4) plus 1 9 Australian

species of Acacia sensu stricto published in recent years but not yet posted to that site (see

Maslin, 2014a-d; Maslin & Bairett, 2014; Kodela, 2015); these numbers refer to accepted,

indigenous species only (not including infraspecific, infomial or hybrid formulae taxa).

Pre- Vienna IBC names

(Acacia treated as a single

genus with A. nilotica the

type)

Post- Vienna IBC names; Acacia sensu lato treated as five genera

With Acacia

nilotica the

type

With

Acacia

penninenns

the tj pe

Species numbers and distribution

Americas Africa*

Asia^

Australia

/Pacific

Total

Acacia

VachelUa

843

1633

subgenus Acacia

subgenus Aculeiferum

section Spiciflorae

Senegalia

102

68^

201'*

section Filicinae

Acaciella

Acacia coidteri group

Mariosoiisa

Mariosousa

subgenus Phyllodineae

Racosperma

Acacia

1064®

1067

^Including Madagascar and Mascarenes. -New Guinea to the Middle East. ^Including three

species in Madagascar and one in Africa for which combinations are not yet available in

VachelUa. including seven species in Madagascar and two in Africa for which combinations

are not yet available in Senegalia. ^Acacia heterophylla was formerly recognised for Reunion

Island but was recently shown by Le Roux et al. (2014) to be conspecific with the Hawaiian

species A. koa and to have colonised the Mascarene archipelago dkectly from the Hawaiian

Islands < 1 .4 million yr ago. ^1 057 species in Australia, seven in the Pacific region.

Gardens’ Bulletin Singapore 67(1): 251-252. 2015

doi: 10.3850/S2382581215000198

251

BOOK REVIEW: Flowering Plants of the Western Ghats, Volume 1 (Dicots)

T,S, NayaVy A. Rasiya Beegam & M. Sibi. 2014 & Flowering Plants of the

Western Ghats, Volume 2 (Monocots). T.S. NayaVy M, Sibi & A, Rasiya

Beegam. 2014.

Thimvananthapuram: Jawaharlal Nehru Tropical Botanic Garden and Research Institute. 26 x

16.5 cm, hard cover, 1683 pp (Vols. 1. & 2). ISBN 978-81-920098-2-7 (Vol. 1) & ISBN 978-

81-920098-3-4 (Vol. 2). Price for the set of two volumes INR 3500 / US$ 200.

The mountain range of the Western Ghats,

whieh extends along the western coast of

the Indian subcontinent and exceeds 1600

km in length, is a well-known biodiversity

hotspot and a UNESCO World Heritage

Site. It is home to more than 8000 taxa

of flowering plants, of which about 15%

are endemic to the area. The Western

Ghats run through six states (from north

to south: Gujarat, Maharashtra, Goa,

Karnataka, Kerala and Tamil Nadu) and

although there are published checklists and/or Floras for each of these states, there

has not hitherto been a work focusing on the entire range. As such, this large and

comprehensive checklist of Flowering Plants is a very welcome addition to the

literature on Indian plants.

The checklist is published in two volumes. Volume 1 starts with a short

introduction to the Western Ghats, including basic information on the geography of

the region, rainfall, climate, major rivers, and forest types, together with a schematic

map. In the paragraph on methodology, the authors describe how they acquired and

verifled the information. The main body of Volume 1 is dedicated to an account of

Dicot families (arranged alphabetically) and genera (also arranged alphabetically, but

with cultivated taxa placed at the end of each family). Volume 2 deals with Monocots

in the same manner, but also includes two Addenda: additional taxa to be included

while the work was in production, and a list of species of doubtful occurrence. There

is also a list of references, and indices to scientiflc and local names. The index of local

names consists of over 13,000 entries and covers six languages spoken across the area:

Gujarati, Hindi, Kannada, Malayalam, Marathi and Tamil. The entire work is almost

1700 pages long.

The checklist is well-formatted, making full use of various fonts to provide

additional information (e.g. when the Western Ghats are highlighted in bold, it means

it is endemic to the area; and when the reference number is in italics it means that

the plant name is treated there as a synonym). The User’s Key explains well how the

entries are constructed and referenced, as well as explaining the formatting details,

making the work easy to use.

Each taxon entry starts with its accepted name, followed by any synonyms, its

habit and the availability of a description and illustration of the taxon. Information on

252

Card. Bull. Singapore 67(1) 2015

its complete native distribution is followed by its distribution and phenology within

the Western Ghats. These are followed by the threat category, economic importance

and a list of local names. Considering that this is just a checklist, it is surprisingly

well referenced. The list of references contains full citations to 2936 numbered works

with these numbers referenced in each entry to justify the information provided and to

allow the reader to follow up on details if needed.

A random check on families with which I am familiar shows that the authors

have strived to gather the latest literature to ensure that the names used are indeed, in

the vast majority of cases, up-to-date. However, I did spot some unusual placements

of genera, e.g. Cheilocostus and Costus of the family Costacaeae are placed in the

Zingiberaceae. The family Costaceae has been unambiguously recognised for at

least the last 50 years and has been reflected in all major recent works and databases,

including those used by the authors for verifications, so their family concept comes as

a bit of a surprise.

As a potential end user I regret that such a work as this, during an era of rapid

botanical exploration and publication, will inevitably become outdated in a few years,

and was not rather designed as an online project. Not only would that have allowed

various targeted searches to be performed, but the authors could also more dynamically

keep the data updated and respond to feedback. It may also perhaps have formed the

kernel of an expansion of the project into fully-fledged Flora of the Western Ghats,

which could also be illustrated. It would be lovely to see ferns and gymnosperms also

included.

Considering the extent of this work, the authors must no doubt be congratulated

on the final product, which must have required much perseverance and the ability to

cope with extremely large numbers of names, synonyms and references.

Jana Leong-Skornickova

Singapore Botanic Gardens