THE GARDENS’ BULLETIN SINGAPORE

The Gardens’ Bulletin Singapore is a peer-reviewed journal publishing original papers and

reviews on a wide range of subjects: plant taxonomy (including revisions), phytogeography,

floristics, morphology, anatomy, as well as horticulture and related fields, such as ecology and

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

Dr David J. Middleton Dr Jana Leong-Skornickova

(Editor-in-Chief) (Managing Editor)

Felicia Tay Dr Daniel C. Thomas Christina Soh

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

Editorial Advisory Board

Professor Sir Peter Crane

Yale University

U.S.A.

Dr Rogier P.J. de Kok

Royal Botanic Gardens, Kew

U.K.

Dr W. John Kress

National Museum of Natural History

Smithsonian Institution

U.S.A.

Dr Mark Hughes

Royal Botanic Garden Edinburgh

U.K.

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

P.R. China

Published twice yearly by the National Parks Board, Singapore, the annual subscription for the journal

is Singapore $100.00 including postage. Overseas subscribers should make payment in the form of

bank draft or international money order in Singapore currency, payable to National Parks Board. Please

forward payment to “Accounts Receivable Section, National Parks Board, Headquarters, Singapore

Botanic Gardens, 1 Cluny Road, Singapore 259569”.

Instructions for contributing authors

(more details in http://www.sbg.org.sg/research/instructions-for-authors.pdf)

Manuscripts should be sent by e-mail to <gardbullsing@gmail.com> or posted to

The Editor, Gardens’ Bulletin Singapore, Singapore Botanic Gardens, 1 Cluny Road,

Singapore 259569.

The language used is English. Prior to submission, it may be advisable to have manuscripts checked by

someone fluent or proficient in that language. Contributions must not have been published or be under

consideration elsewhere. There are no page charges. Submitted material cannot be returned.

Authors will receive a pdf reproduction of their contribution, gratis.

Cover letter, submission and copyright. In your cover letter or message, state clearly you are submitting

your manuscript (state title) for publication in the Gardens’ Bulletin Singapore. By submitting the

manuscript(s), authors agree that upon acceptance, copyright of the entire work becomes the property

of the Singapore Botanic Gardens, National Parks Board, Singapore. Authors must ensure all required

permission has been obtained in writing to publish any material they present; proof may be required.

Manuscripts are best submitted in electronic form (Microsoft Word 97-2003 preferred). Use Times New

Roman, 10-point font, 1.5-line spacing. Do not send original artwork unless requested by the Editor.

Number all pages serially including the title and abstract on the first.

(continued on inside back cover)

The Gardens ’ Bulletin

Singapore

VOL. 69(2) 2017 ISSN 0374-7859

CONTENTS

M.A. Niissalo & J. Leong-Skornickova

Hangnanapodzolicola (Hanguanaceae),

a new record for Singapore.157

R.P.J. de Kok

Two new records of Litsea (Lauraceae) from Singapore

and the lectotypification of twenty-two names

from several Lauraceae genera.167

M. Dancak, M. Hrones, R.S. Sukri, F. Metali & A.A. Joffre

Novitates Bruneienses, 9. A synopsis of Epirixanthes (Polygalaceae)

in Brunei Darussalam and notes on species elsewhere.179

M. Ardiyani, M.F. Newman & A.D. Poulsen

Anew species of Zingiber (Zingiberaceae)

east of Wallace’s Line.189

G.G. Hambali, S. Sunarti & Y.W. Low

Syzygium jiewhoei (Myrtaceae), a new endemic tree

from Western New Guinea, Indonesia.201

C. Puglisi & D.J. Middleton

A revision of Microchirita (Gesneriaceae) in Thailand.211

M. Rodda & U. Meve

Ceropegia laotica (Apocynaceae, Asclepiadoideae):

the first new species of Ceropegia described from Laos

285

W.H. Chen, D.J. Middleton, H.Q. Nguyen, H.T. Nguyen,

L.V. Averyanov, R.Z. Chen, K.S. Nguyen, M. Moller & Y.M. Shui

Two new species of Oreocharis (Gesneriaceae)

from Northwest Vietnam.295

S. Li, Z.B. Xin, X. Hong, L.F. Fu & F. Wen

Primulina wuae (Gesneriaceae), a new species

from southern China.307

Date of publication: 20 November 2017

National Parks Board

Singapore Botanic Gardens

1 Cluny Road

Singapore 259569

Printed by Oxford Graphic Printers Pte Ltd

Gardens' Bulletin Singapore 69(2): 157-165. 2017

doi: 10.26492/gbs69(2).2017-01

157

Hanguana podzolicola (Hanguanaceae),

a new record for Singapore

M.A. Niissalo 1 & J. Leong-Skornickova 2

Molecular Biology & Micropropagation, Research & Conservation Branch,

Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, 259569, Singapore

matti. niis salo @ gmail .com

2 Herbarium, Research & Conservation Branch,

Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, 259569, Singapore

j ana_skornicko va@nparks. gov. sg

ABSTRACT. Hanguana podzolicola (Hanguanaceae) is newly recorded for Singapore.

Detailed colour plates are given alongside the main characters for distinguishing this species

from the two most similar species in Singapore, Hanguana rubinea and H. triangulata. The

seeds of Hanguana podzolicola are described for the first time. A local conservation assessment

is given. As no original material of Hanguana podzolicola could be traced, a neotype is

designated here. Following a recent clarification of several historical Hanguana names it is

noted here that the correct name for the large helophytic stoloniferous species often cultivated

in Singapore under the name Hanguana malayana is Hanguana anthelminthic a. An updated

key to Hanguana species in Singapore is provided.

Keywords. Central Catchment Nature Reserve, Hanguana anthelminthic a, H. malayana, H.

pantiensis, IUCN conservation assessment, Johor, neotypification, Peninsular Malaysia

Introduction

In a recent overview of Hanguana Blume in Singapore, Leong-Skornickova & Boyce

(2015) recognised four native, one introduced, and one cultivated species. Since the

publication of this paper, fieldwork on Hanguanaceae has continued and a survey of

all populations of Hanguana, along with a systematic survey of the primary forest

habitats of Zingiberales in Singapore, has been undertaken (Niissalo et al., 2017).

During these surveys we found a population of a species that has not been previously

reported from Singapore.

We have compared our material to all seven Hanguana species so far described

from Peninsular Malaysia (Jack, 1820; Siti Nurfazilah et al., 2010; Leong-Skornickova

& Kiew, 2016). The plants in Singapore are similar to two species described from Johor,

Hanguana podzolicola Siti Nurfazilah et al. and H. pantiensis Siti Nurfazilah et al., in

their leaf characters, small fruits and strongly obliquely positioned stigmas, although,

of these two, the spreading, long inflorescence branches of the Singaporean plant

158

Card. Bull. Singapore 69(2) 2017

better fit H. podzolicola. We have not seen individuals with particularly pronounced

aerial stems in Singapore, but based on our observations in Singapore and Peninsular

Malaysia, we believe that this character has little taxonomic value, as it is regularly

seen in many taxa and is likely an indication of an old individual. Siti Nurfazilah et al.

(2010) only reported female plants and, likewise in Singapore, only female plants have

been collected. We provide a full description, including the previously undescribed

seeds, and photographic illustrations of the species based on the Singapore population

to aid future taxonomic work.

The type specimens and all paratypes for the five names published in Siti

Nurfazilah et al. (2010) were never deposited in KEP. The whereabouts of these

materials, including those of Hanguana podzolicola , remain unknown. There are no

morphologically well-matched specimens of Hanguana podzolicola from the type

locality or nearby areas in any of the herbaria we have examined (E, K, KEP, L, P,

SING). A single sheet of Hanguana podzolicola was found at USM (. Mohd Fahmi Bin

Abu Bakar et al 59; Sofiman Othman, pers. comm), but this collection is not mentioned

in the protologue and it is not from the type locality. The specimen consists of a young

female inflorescence, but it has no leaves or fruits and therefore does not allow for

unambiguous identification of the species. As the population in Singapore appears to

be morphologically inseparable from the plants from the type locality illustrated by

Siti Nurfazilah et al. (2010), we designate a neotype from a fully ripe female specimen

collected in Singapore.

We consider the two specimens, Corner s.n. from Mandai Road and Ridley

170 from Seletar, previously cited with caution as Hanguana rubinea Skomick. &

P.C.Boyce by Leong-Skornickova & Boyce (2015), to rather be H. podzolicola as

they match the living material seen in all aspects, most prominently in the large size

of the female inflorescences. The ripe fruit on Comer s.n. also matches Hanguana

podzolicola in the strongly obliquely positioned stigmas and seed structure.

The vegetative parts of this species, especially the corrugated leaves and fairly

prominent flocculose indumentum, are very similar to those of Hanguana triangulata

Skornick. & B.C.Boyce. Our preliminary results from genetic analyses of Singapore’s

Hanguana populations (Niissalo et al., manuscript in preparation) suggest that the

sterile specimen from Upper Seletar originally cited by Leong-Skorniclcova & Boyce

(2015) under H. triangulata ( Leong-Skornickova, J. & Thame, A. JLS-3036), is

genetically part of the same population as our collections of fertile H. podzolicola.

We therefore correct its identification to Hanguana podzolicola here. The only

surviving populations of Hanguana triangulata are therefore in Bukit Timah Nature

Reserve and, as confirmed by the preliminary results of our analyses, all plants with

corrugated leaves in the northern parts of the Central Catchment Nature Reserve

are H. podzolicola. In the fruiting stage these two taxa are not easily confused as

Hanguana podzolicola has much longer and more slender inflorescence branches (to

c. 30 cm), smaller pink fruits (c. 5-7 mm in diam.) that turn translucent green-dull

pink to green-brown as they ripen, and strongly obliquely positioned and rounded

stigma lobes (compared to short and almost perpendicular branches to 8 cm long,

larger cream-white fruits 9-10 mm in diam., a stigma which is terminal or slightly

Hanguana podzolicola, a new record for Singapore

159

oblique, and stigma lobes which are connate at base and with sharp apices forming

an equilateral triangle in H. triangulata). In addition the seed has a single, broadly

and bluntly acute appendage (compared to the seed appendage bluntly bilobed in H.

triangulata). Hanguana podzolicola is also similar in stature and inflorescence details

to H. rubinea, but differs from it by its prominently corrugated leaves and abaxially

more thickly flocculose indumentum (compared to the a lm ost flat lamina with a

sparse indumentum in H. rubinea). They also differ in inflorescence branch length,

fruit, and seed details (in Hanguana rubinea branches are up to 11 cm long, fruits are

larger, 9-10 mm in diam., and turn ruby-red as they ripen, stigma is terminal or only

slightly oblique, stigma lobes are similar to H. podzolicola , except larger, and the seed

appendage is triangular). Hanguana podzolicola has the largest female inflorescences

of any native Hanguana in Singapore.

With the recent work on Hanguana in Singapore (Niissalo et al., 2014; Leong-

Skomickova & Boyce, 2015), the current paper brings the number of native Hanguana

species in Singapore to five. Hanguana malayana (Jack) Merr., the name provisionally

applied to the cultivated massive helophyte by Leong-Skornickova & Boyce (2015;

see Fig. 1 & 11-12), is here updated to Hanguana anthelminthic a (Blume ex Schult.

& Schult.f.) Masam. to reflect a recent clarification of the old names of Hanguana

(Leong-Skornickova & Niissalo, 2017). Hanguana malayana is a solitary forest

species that is so far only known from Penang and has never been collected from

Singapore, whereas Hanguana anthelminthica is the widespread massive helophyte

that is also widely cultivated in the tropics, including in Singapore. An updated key to

all Hanguana species in Singapore is provided.

Hanguana podzolicola Siti Nurfazilah et al. - TYPE: Singapore, Mandai Road, 24

Aug 2015, Leong-Skornickova, J. HAN-76 (neotype SING [mounted over 2 sheets

and including fruits preserved in spirit as part of a single specimen], here designated).

(Fig. 1, 2)

Herbaceous, dioecious mesophyte to c. 1.6 m tall; stem terete, to 3 cm in diam.,

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

of up to 20 leaves; stolons absent. Leaves to 170 cm long, spreading then arching,

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

pseudopetiole 60-70 cm long, c. 10-14 mm wide, accounting for 1/3—1/2 of entire

leaf length, roundly channelled with sharp margins, strigose; leaf blade 80-105 x

14-17 cm, narrowly elliptic, base attenuate, tip long, narrowly attenuate with apicule

c. 4 mm, leathery, irregularly corrugated, adaxially mid to dark green, sparsely hairy

(silky appressed hair; falling off in older leaves), abaxially lighter green when fresh,

prominently flocculose (falling off in older leaves); midrib weakly impressed, of the

same colour as the rest of the lamina adaxially, round-raised, lighter green, almost

glabrous and shiny abaxially. Male inflorescences not observed, female inflorescences

erect at anthesis. Female inflorescence/infructescence erect, comprising up to 9

partial, whorled, alternate-secund, thyrsoid infructescences plus a terminal spike;

160

Card. Bull. Singapore 69(2) 2017

Fig. 1. Hanguana podzolicola Siti Nurfazilah et al. A. Lower side of the lamina showing

flocculose indumentum. B. Upper side of the lamina. C. Habit (inset: detail of the petiole). D.

Detail of young fruits (photographed on 29 Apr 2015) E. Detail of ripe fruits (photographed

from the same individual on 24 Aug 2015). F. Infructescence. From Leong-Skornickovd, J.

HAN-76. (Photos: Jana Leong-Skornickova)

Hanguana podzolicola, a new record for Singapore

161

Fig. 2. Hanguana podzolicola Siti Nurfazilah et al. A. Side view detail of fruit attached to a

branch showing 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, showing asymmetrically

placed stigma. G. Seed (side view, appendage facing camera). H. Seed (side view, appendage

to the right). I. Seed (top view, appendage to the left). From Leong-Skornickova, J. HAN-76.

(Photos: A-F: Jana Leong-Skornickova; G-I: Matti A. Niissalo)

partial inflorescences spreading almost perpendicularly to rachis (lowermost 2

ascending); peduncle and rachis together up to 90 cm tall, green when fresh,

conspicuously pale brown-grey flocculose, visible portion of peduncle up to 30 cm

long; one sterile bract per peduncle, foliaceous, persistent, narrowly ovate with a basal

claw, c. 95 (inch 30 cm long claw/pseudopetiole) x c. 11 cm; bract subtending partial

inflorescences similar to sterile bracts, diminishing in size and becoming narrowly

triangular distally along the infructescence, the bract supporting basal-most partial

162

Card. Bull. Singapore 69(2) 2017

inflorescences c. 50 x 10 cm (incl. 6 cm claw), fully reduced in uppermost partial

inflorescences; partial inflorescences each comprising up to 17 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 1/2- 2/3 of

the median branch), median branches at basal levels usually further branched, 20-30

cm long (10-25 cm long in upper levels). Female flowers scattered, solitary or in

pairs, sessile, all with an associated minute bract and bracteole; perianth composed

of 6 tepals in two whorls tightly clasping ovary/fruit in fresh material, all tepals

with prominent bulbous thickening at base (more prominent in outer whorl), light

green, margin c. 0.2-0.3 mm wide, hyaline translucent white; outer tepals broadly

ovate, 2.5-3 mm long, c. 2.8-3 mm broad, connate at base (only 0.3 mm), sparsely

arachnoid; inner tepals almost circular, c. 3 mm long, 3-3.2 mm broad, free to base,

almost glabrous; staminodes 6, in two whorls, pale green to cream white, triangular,

outer staminodes , 0.3-0.4 mm long, 0.2-0.3 mm broad at base, inner staminodes

longer, narrowly triangular, c. 1 mm long, 0.2-0.3 mm at base, each basally sheathed

with a broad narrow scale (often shallowly bilobed), c. 0.8-0.9 mm long, and c. 1.2

mm broad, brown with translucent margin, stigma 3-lobed, c. 1.5 mm in diam., each

lobe 0.8-1 mm long (fruiting material), broadly ovate with round apex, lobes connate

basally, with points of connation seen as grooves, matte dark brown (fruiting stage);

position of stigma in ripe fruits strongly oblique (only a single seed ever develops).

Ripe fruit with pink blush externally (ripe fruit appears dirty green-pink or green-

brown as the dark seed colour is visible through the increasingly translucent pulp),

globose, 5-7 mm diam.; pulp 0.7-1 mm thick, pale yellow, fairly hard, ripening from

cream-white with bright pink blush, to dirty pale yellow with dull pink tinge; seeds

1 per fruit (2 seeds in a single fruit have not been observed), c. 5 x 4 mm, brown,

deeply bowl-shaped with slightly incurved margins, with a small broadly bluntly acute

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

Distribution. Both the historic and recent collections of this species in Singapore

are from the northern parts of the Central Catchment Nature Reserve or locations

immediately adjacent to it. The species is now only found in the Nee Soon Freshwater

Swamp-forest (NSFS). One historic collection locality, Seletar, has likely been

deforested, but the locality name was used for a larger swamp-forest complex that

was once connected to NSFS (O’Dempsey & Chew, 2013). In Peninsular Malaysia

the species has been recorded from two localities in Johor, Hutan Simpanan Lenggor,

Mersing and Hutan Lipur Panti, Kota Tinggi (Siti Nurfazilah et al., 2010).

Ecology & phenology. In Singapore Hanguana podzolicola is strictly a swamp-forest

species, occurring in locations with permanently wet or flooded soil in NSFS. The

only other species recorded there, Hanguana rubinea, occurs in drier locations. The

two are only occasionally sympatric at the edges of the swamp forest (Upper Seletar

and southern end of NSFS). Based on historical collections and our observations, the

species flowers early in the year (c. March) and fruits ripen by July-August.

Hanguana podzolicola, a new record for Singapore

163

Provisional IUCN conservation assessment. The known Extent of Occurrence of

Hanguana podzolicola is c. 500 km 2 and the species is currently only known from

three localities, with an estimated Area of Occupancy of c. 10 km 2 . Two of the known

localities are in protected areas, but the third is highly susceptible to forest loss. The

species should therefore be considered to be Endangered globally, EN Blab(iii)

(IUCN, 2012). In Singapore there are c. 50 stems, all solitary, and the majority of them

are found within a small area (c. 1/2 ha) of particularly wet, low-lying forest, with

only very few or single stems elsewhere in NSFS. Given the small number of stems,

small total area of occupancy and the highly clustered distribution, the species should

be considered Critically Endangered (CR C2) nationally using the slightly amended

national categories of Davison et al. (2008).

Additional specimens examined. SINGAPORE: Seletar: 29 Mar 1889, Ridley, H.N. 170

(SING, female flowers). Central Catchment Nature Reserve: Mandai Road, 28 Jul 1929,

Corner, E.J.H. s.n. (SING, fully ripe infructescence); forests around Upper Seletar Reservoir,

19 Aug 2014, Leong-Skornickovd, J. & Thame, A. JLS-3036 (SING, sterile); Mandai Road, 29

Apr 2015, Leong-Skornickovd, J. & Niissalo, M. HAN-75 (SING, leaf and old infructescence).

Notes: The fruit colour develops in an unusual way in this species, and the fruits

illustrated in Siti Nurfazilah et al. (2010) are not yet fully ripe. In the fruits we have

observed, the fruits have nearly reached their mature size by late April. At this point

the fruits have a bright pink coloration, but they remain unripe and the seeds remain

soft. The fruits ripen very slowly and become more translucent over time. As the seeds

ripen dark brown, the fruits appear dirty greenish-dull pink or even green-brown when

the seeds are fully developed. This ripening stage lasts for about five months after the

fruits have their final size (Fig. 1), bringing the total flowering and fruiting process to

last 7-8 months.

Singapore plants overall agree closely with measurements from Malaysia (Siti

Nurfazilah et al., 2010), but the petioles we measured are proportionally longer (1/3 of

leaf length in Malaysia, up to 1/2 of leaf length in Singapore), and the outer whorls of

petals are also slightly larger in Singapore than in the Malaysian plants (1 x 1.8 mm in

Malaysia, 2.5-3 x 2.8-3 mm in Singapore). However, when comparing photographs

of the fruits of both Malaysian and Singapore collections, the proportional dimensions

of outer and inner tepals compared to diameter of the fruits appears to be identical.

Leong-Skomickova & Boyce (2015) discussed the taxonomic importance of

oblique stigmas in Hanguana and, as they mention, oblique stigmas can occur in

many Hanguana species when only a single seed develops inside the fruit. However,

in Hanguana podzolicola (and apparently H. pantiensis) the stigma is very strongly

oblique, even closer to the base of the fruit than the apex, and such a strongly oblique

stigma is an unmistakable character of taxonomic significance. It is also well preserved

in dry material. The species is also characterised by the thick, flocculose indumentum,

but this character is often lost on old herbarium specimens and also on the older leaves

of live plants and therefore unreliable for herbarium-based taxonomy.

164

Card. Bull. Singapore 69(2) 2017

Updated 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

a hyaline margin; stigma lobes large, flat, connate at base, forming a bluntly

triangular to clover-leaf shape, almost obscuring the apex of ovary.

. H. anthelminthic a

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

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

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

stigma lobes small, erect, separate, teardrop-shaped to obovate. H. nitens

3a. Leaves green on both sides.4

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

4a. Large herbs over 1 m in height; leaves arching; ripe fruits cream-white, dull pink

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 3/4 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. H. rubinea

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

flocculose indumentum, ripe fruits cream-white or dull pink.6

6a. Ripe fruits cream-white, 9-10 mm in diam.; stigma lobes with sharply acute

apices, forming sharply triangular structure; seed appendage bluntly bilobed.

. H. triangulata

6b. Ripe fruits dull pink, 5-7 mm in diam.; stigma lobes connate basally (sometimes

imperfectly), with round apices, forming bluntly triangular structure; seed

appendage single, broadly bluntly acute. H. podzolicola

ACKNOWLEDGEMENTS. Our research is supported by the National Parks Board,

Singapore. We thank our colleagues Aung Thame and Derek Liew, as well as the volunteers

Michael Leong and Lahiru Wijedasa, for their help during our fieldwork in Singapore. We

thank Dr Gillian Khew for supervision and Khoo-Woon Mui Hwang for support with ongoing

molecular work on Hanguana populations. We also thank the curators of herbaria (E, K, KEP,

L, P, SING and USM) for allowing us access to specimens in their care and/or making high-

resolution images accessible. We thank Sofiman Othman for his help with checking whether

Hanguana podzolicola, a new record for Singapore

165

Hanguana podzolicola specimens are present at USM and for providing us with photographs

of two collections. Dr David Middleton and the reviewers are thanked for linguistic help and

constructive comments on this manuscript.

References

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).

IUCN (2012). IUCN Red List Categories and Criteria: Version 3.1, 2 nd ed. Gland, Switzerland

and Cambridge, UK: IUCN.

Jack, W. (1820). Descriptions of Malayan plants No 2. In: Jack W., Malayan Miscellanies, vol.

1, no 5, pp. 1—49. Bencoolen: Sumatran Mission Press.

Leong-Skomickova, J. & Boyce, PC. (2015). Hanguana in Singapore demystified: an overview

with descriptions of three new species and a new record. Gard. Bull. Singapore 67:

1-28.

Leong-Skornickova, J. & Kiew, R. (2016). Hanguana fraseriana (Hanguanaceae), a new

species from Peninsular Malaysia. Gard. Bull. Singapore 68: 209-214.

Leong-Skomickova, J. & Niissalo, M.A. (2017). Identity and typification of Hanguana

malayana and H. anthelminthica, and notes on other early names in Hanguanaceae.

Plant Syst. Evol. 303:1213—1223.

Niissalo, M.A., Wijedasa, L.S., Boyce, PC. & Leong-Skornickova, J. (2014). Hanguana

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

reserve in Singapore. Phytotaxa 188: 14-20.

Niissalo, M.A., Leong-Skornickova, J., Khew, G.S. & Webb, E.L. (2017). Very small relict

populations suggest high extinction debt of gingers in primary forest fragments of a

tropical city. Am. J. Bot. 104: 182-189.

O’Dempsey, T. & Chew, PT. (2013). The freshwater swamp forests of Sungei Seletar

catchment: a status report. In: Ming, L.T. & Chew, H.H. (eds) Proceedings of Nature

Society, Singapore’s Conference on ‘Nature Conservation for a Sustainable Singapore’

- 16th October 2011, pp. 121-166. Singapore: The Nature Society (Singapore).

SitiNurfazilah, A.R., Mohd Fahmi, A.B., Ahmad Sofiman, O. & Boyce, PC. (2010). 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.

Gardens' Bulletin Singapore 69(2): 167-177. 2017

doi: 10.26492/gbs69(2).2017-02

167

Two new records of Litsea (Lauraceae) from Singapore

and the lectotypification of twenty-two names

from several Lauraceae genera

R.PJ. de Kok

Honorary Research Associate, Singapore Botanic Gardens,

National Parks Board, 1 Cluny Road, 259569 Singapore

dekokrogier@gmail.com

ABSTRACT. Two species of Litsea (Lauraceae) are recorded for Singapore for the first time

C Litsea spathacea Gamble and L. tomentosa Blume). Both species are known only from 19th

century specimens and must be considered nationally extinct in Singapore. Descriptions and notes

on distribution, conservation status and ecology are given. In addition, twenty-two Lauraceae

names are lectotypified: Beilschmiedia curtisii Gamble, B. perakensis Gamble; Cinnamomum

subavenium Miq.; Cryptocarya argentea Gamble, C. infectoria (Blume) Miq., C. tomentosa

Blume; Lindera lucida (Blume) Boerl., L. malaccensis Hook.f.; Litsea accedens (Blume) Boerl.,

L. amara Blume var. attenuata Gamble, L. gracilis Gamble, L. gracilipes Hook.f., L. griffithii

Gamble, L. lanceolata (Blume) Kosterm., L. machilifolia Gamble, L. machilifolia Gamble var.

angustifolia Gamble, L. megacarpa Gamble, L. pustulata Gamble, L. sarawacensis Gamble, L.

singaporensis Gamble, L. spathacea Gamble, L. umbellata (Lour.) Merr.

Keywords. Beilschmiedia, Cinnamomum, Cryptocarya, Lindera, Litsea, Singapore

Introduction

Recently several genera of Lauraceae have been revised for the Flora of Singapore

(Beilschmiedia Nees, Cinnamomum Schaeff., Cryptocarya R.Br., Lindera Thunb. and

Litsea Lam.). The Lauraceae (Laurel Family) are of major ecological and economic

importance in Southeast Asia as they comprise a major part of many forests in the

region. In particular, species of the genus Cinnamomum are important as sources

of various spices and several species have been introduced into Singapore for this

purpose. In Singapore, there are two species of Beilschmiedia (de Kok, 2016a), five of

Cinnamomum (de Kok, in prep.), seven of Cryptocarya (de Kok, 2015, 2016b), one

of Lindera and eighteen of Litsea (de Kok, in prep.). These will be enumerated in the

forthcoming Flora of Singapore. No species is endemic to Singapore. Unlike in the

Lamiaceae (de Kok et ah, 2016), it is relatively easy to determine if a species is native

or naturalised in Singapore. Two species are recorded from Singapore for the first

time: Litsea tomentosa Blume, and L. spathacea Gamble. Both species are common

in the lowland forests of Peninsular Malaysia and their occurrence in Singapore is

therefore not a surprise.

168

Card. Bull. Singapore 69(2) 2017

The revisions of Lauraceae from Singapore are based on herbarium collections

made in Singapore and now housed at the Natural History Museum, London (BM), the

Royal Botanic Gardens, Kew (K and K-W), the Forest Research Institute of Malaysia

(KEP) and the Singapore Botanic Gardens (SING). Type material not available at

those institutions was studied online via JSTOR (https://plants.jstor.org/plants) and

accessed in July 2017. In addition to the new records for Litsea, twenty-two names

from all of the genera revised need to be lectotypified. Barcode information is given

for all specimens where available.

New species records for Singapore

Litsea spathacea Gamble, Bull. Misc. Inform. Kew 358 (1910); Ridley, FI. Malay

Penins. 3: 120 (1924); Kostermans, Bibliogr. Lauracearam 881 (1964); Kochummen

in Ng, Tree FI. Malaya 4: 165 (1989). - TYPE: [Peninsular Malaysia] Perak, Sungei

Larut, July 1988, L. Wray 2286 (lectotype K [K000797118], designated here;

isolectotype SING [SING0229222]).

Tree or shrub 4.5-15 m tall; dbh 7-30 cm. Twigs slender to stout 2.7-9 mm thick,

grey (brown), angled to round in cross-section, densely hairy, glabrescent; hairs

appressed, yellowish; terminal leaf bud 2-6.3 mm long, lanceolate to ovate, apex acute,

velutinous. Leaves alternate. Leaf blade elliptic to oblanceolate or obovate, 11-29 x

4-12 cm, apex (broadly) acute, base cuneate, margins straight, blade membranous

to thinly leathery, secondary veins 9-14 pairs, curving near margin and sometimes

brochidodromous, tertiary veins scalariform-reticulate; upper surface glossy, dark

green, glabrous with a few hairs on midrib, midrib sunken, secondary veins raised,

tertiary veins distinct; lower surface green with yellowish midrib, glaucous, glabrous

to sparsely hairy, always some hairs present on midrib, midrib and secondary veins

raised, tertiary nerves distinct. Petiole slender to slightly swollen, half-terete, 10-25

mm long, sparsely hairy. Inflorescence 1—4 cm long, formed of clusters of umbels

in axils of leaves or along branchlets, umbels 6-7.5 mm in diam., pale yellow-green

or greenish white; bracts 5, usually imbricate, (sub)orbicular, concave, 5-6.5 x 5-6

mm, velutinous outside. Male flowers white, 6-8 in each umbel; tepals 5-6, elliptic-

lanceolate, subequal, 1.5-4 x 0.8-1 mm, hairy; stamens 8-14, unequal; filaments 1-5

mm long, a few hairs present; anthers 0.5-1.5 mm long, yellow. Female flowers 3-6

in each umbel; tepals 5-6, elliptic-lanceolate, subequal, 2-3 x 0.9-3 mm, sparsely

hairy; staminodes 7-12, linear, 0.5-3 mm long, glabrous; ovary ovoid, 0.5-1 mm

in diam., glabrous; style 1-2 mm long; stigma peltate. Infructescence of 1-3 fruits.

Fruits globose to elliptic 15-20 x 11—14 mm, smooth, glabrous, red or purplish when

mature; cupule shallow, 13-15 mm in diam., 6-7.2 mm high, glabrous, margin entire

or undulate, surface smooth; fruiting pedicel 5-10 mm long, slightly thickened, 1.7-

2.6 mm thick.

New records and typifications for Lauraceae in Singapore

169

Distribution. Peninsular Malaysia and Singapore. In Singapore, it is only reported

from two specimens both without a locality or date (Cantley’s Collector s.n. (SING);

Unknown s.n. (BM)).

Ecology. In the rest of its range growing in lowland and hill forests, sometimes along

rivers, from 90-975 m altitude. Flowering all year round; fruiting from February to

March.

Conservation. Least Concern (LC) globally, this species is widespread and has been

collected over a wide area in the last 20 years (see also IUCN, 2017). In Singapore,

it has been collected only twice, in the late 19th or early 20th century, and must,

therefore, be considered to be nationally extinct.

Notes. There are eleven different gatherings, which are usually duplicated in several

herbaria, available for lectotypification for this name. As Gamble was working at K

when he wrote his paper (Gamble, 1910), the K specimen of the gathering: L. Wray

2286 (K000797118) is designated here as the lectotype.

As mentioned above, it is only known from Singapore by two specimens,

one at SING, collected by Cantley’s Collector , without a collection number, while a

specimen with no collector’s name and number is housed at BM. Cantley worked in

Peninsular Malaysia and Singapore in the 1880’s. During this time, he had a collector

(M.V. Alvins) who also made collections for him from the same area, Alvins’ labelling

is said to be inadequate (only giving the state for localities) and sometimes incorrect

(Van Steenis-Kruseman, 1950: 100). The label on the BM collection is from the early

20th century. It seems likely, therefore, that both these specimens where collected in

the period between 1880 and 1900, with a question mark about the accuracy of the

locality of Cantley’s Collector (M.V. Alvins). The species is common in the lowlands

of Peninsular Malaysia, but is not known from open vegetation. It is, therefore, not

surprising that it used to grow in Singapore and that it now no longer does. However,

the species is difficult to recognise as it is morphologically similar to the very common

Litsea umbellata (Lour.) Merr. and could, therefore, have been overlooked in surveys.

Litsea tomentosa Blume, Bijdr. FI. Ned. Ind. 11: 566 (1826); Gamble, J. Asiat. Soc.

Bengal 75: 132 (1912); Ridley, FI. Malay Penins. 3: 115 (1924); Backer & Bakhuizen,

FI. Java (Spermatoph.) 1: 125 (1963); Kostermans, Bibliogr. Lauracearum 886 (1964);

Kochummen in Ng, Tree FI. Malaya 4: 165 (1989); Ngernsaengsaruay et al., Thai

Forest Bull., Bot. 33: 90 (2011). - TYPE: [Indonesia] Java, C.L. Blume s.n. (lectotype

K [K000815822], designated by Ngernsaengsaruay et al. (2011: 90)).

Tree 6—40 m tall; dbh 7—40 cm, buttresses extending out to 60 cm; bark smooth,

lenticellate, greyish or light brown to black, inner bark red brown, wood yellow.

Twigs slender to stout 3.8-12 mm thick, round or angular in cross-section, velutinous,

glabrescent; hairs appressed to patent, yellowish; terminal leaf bud 5.5-8.3 mm

170

Card. Bull. Singapore 69(2) 2017

long, ovate, apex acuminate, velutinous. Leaves spiral, crowded toward the apices

of the branchlets. Leaf blade elliptic to oblanceolate, 8—40 x 4-16 cm, apex acute to

obtuse, base cuneate to rarely rounded, sometimes asymmetric, margins straight, blade

chartaceous, secondary veins 11-16 pairs, curving and sometimes brochidodromous,

tertiary veins scalariform-finely reticulate; upper surface green, glabrous with densely

hairy midrib and secondary veins, midrib raised to sunken, secondary veins shallowly

sunken, tertiary veins faint; lower surface glaucous, sparsely hairy to densely so on

major veins, midrib and secondary veins raised, tertiary veins distinct. Petiole half-

terete, 10-55 mm long, velutinous, glabrescent; hairs appressed to patent, yellowish.

Inflorescence 1.5-3 cm long, formed of clusters of umbels along branchlets, velutinous,

umbels 12-20 mm in diam.; bracts 4-5, suborbicular or broadly ovate, concave, 5-6 x

4-5 mm, velutinous outside. Male flowers 5-6 in each umbel; tepals 9-12, lanceolate,

unequal, 4-6 x 1.5-2 mm, sparsely hairy; stamens 24-30, unequal; filaments 2-5 mm

long, sparsely hairy; anthers 1-1.5 mm long. Female flowers 5-6 in each umbel, tepals

8—12, lanceolate, 2-3.1 x 1-1.3 mm, densely hairy; staminodes 26-30, linear, 1.5-

3.8 mm long, sparsely hairy; ovary ovoid, 1-1.5 mm in diam., glabrous; style 1.5-2

mm long; stigma peltate. Fruits (sub)globose, 1.6-2.2 x 2-2.1 cm, smooth, glabrous,

glossy, red when mature; cupule shallow, 11-15 x 1.5-3 mm, sparsely hairy, margin

entire, surface warty; fruiting pedicels 7-9.6 mm long, not swollen, 4.3-5.8 mm thick,

sparsely hairy.

Vernacular names. Medang taudok (Malay).

Distribution. Peninsular Thailand, Malaysia (Peninsular and Borneo), Singapore,

Indonesia (Borneo, Sumatra and Java) and the Philippines. In Singapore, it has been

collected only once, at Tanglin (H.N. Ridley s.n.. Upper Tanglin, behind barracks,

1893).

Ecology. Throughout its range in open areas in disturbed lowland forests, between

0-750 m altitude. Flowering from February to June; fruiting from September to

December.

Conservation. Least Concern (LC) globally, this species is widespread and has been

collected over a wide area in the last 20 years. In Singapore, it has only been collected

once, in the late 19th century, and must therefore be considered to be nationally extinct.

Notes. This species has not previously been recorded for Singapore, where it is only

known from a single specimen, collected by H.N. Ridley in 1893, from upper Tanglin,

behind the barracks. The species is common in the lowlands of Peninsular Malaysia

and on the islands of the Sunda shelf and can occur in open areas as well as in disturbed

forests. It is, therefore, not surprising that it used to grow in Singapore, but given that it

can adapt to open vegetation, it is perhaps surprising that it no longer does. The species

is very distinctive with its broad hairy leaves which are crowded near the apex of the

young twigs. It is unlikely that it has been overlooked in botanical surveys.

New records and typifications for Lauraceae in Singapore

171

Lectotypifications

Beilschmiedia curtisii Gamble, Bull. Misc. Inform. Kew 148 (1910). - TYPE:

Singapore, Garden Jungle, July 1896, H.N. Ridley 8075 (lectotype K [K001083364],

designated here; isolectotype SING [SING 0219777]).

At least two gatherings are mentioned in the original description of Beilschmiedia

curtisii Gamble (Gamble, 1910: 148): C. Curtis 1015 and H.N. Ridley 8075. As Curtis

numbered species rather than gatherings, there is no guarantee that all Curtis 1015

specimens are from the same collection event, despite all having the same locality:

Government Hill. The Curtis collections are, therefore, not selected for lectotypification.

As Gamble was working at Kew at the time he described this species, the K specimen

of H.N. Ridley 8075 is designated here as the lectotype.

Beilschmiedia perakensis Gamble, Bull. Misc. Inform. Kew 149 (1910). - TYPE:

[Peninsular Malaysia] Perak, King’s Collector 10026 (lectotype K [K000768677],

designated here; isolectotypes K [K001098151, K001098153]).

Several gatherings are mentioned in the original description of Beilschmiedia

perakensis Gamble (Gamble, 1910: 149): B. Scortechini s.n. and King’s Collector

8489, 10026 and 10432, which are all very similar. Of these, one of the K specimens

of King’s Collector 10026 [K000768677] is designated here as the lectotype.

Cinnamomum subavenium Miq., FI. Ned. Ind. 1: 90 (1858). - TYPE: [Indonesia]

Sumatra, Solok, J.E. Teijsmann s.n. (lectotype U [U0002677], designated here;

isolectotypes BO, U [U0002678]).

Only one gathering is mentioned in the original description of Cinnamomum

subavenium Miq. (Miquel, 1858: 90): Teijsmann, Sumatra, Solok ‘Madang koelit

manis’. At U, two sheets collected by Teijsmann have this locality and the same local

name and sheet number, HB1037. One specimen has a few flowers and a very young

fruit, while the other is sterile. The fertile specimen, U0002677, is therefore designated

here as the lectotype.

Cryptocarya argentea Gamble, Bull. Misc. Inform. Kew 144 (1910). - Cryptocarya

kurzii var. argentea (Gamble) Airy Shaw, Bull. Misc. Inform. Kew 535 (1939). -

TYPE: [Peninsular Malaysia] Perak, Aug 1885, King’s Collector 7966 (lectotype K

[K001084384], designated here; isolectotypes BM [BM001124600], K [K001084383],

P [P00745434], SING [SING0059162]).

172

Card. Bull. Singapore 69(2) 2017

Only one gathering was mentioned in the original description of Cryptocarya argentea

Gamble (Gamble, 1910: 144-145): King’s Collector 7966. As Gamble was working

at Kew at the time he wrote this article, one of the two K specimens is selected as the

lectotype. As sheet K001084384 has more immature fruits than K001084383, it is

designated here as the lectotype.

Cryptocarya infectoria (Blume) Miq., FI. Ned. Ind. 1: 924 (1858). - Cylicodaphne

infectoria Blume, Mus. Bot. 2: 11 (1856). - TYPE: In Archipelago Indico, F.A.C.

Waitz s.n. (lectotype L [L0036159] designated here; isolectotypes L [L0036157,

L0036158]).

In the original description, Blume (1856) mentioned only the F.A.C. Waitz gathering, of

which there are three sheets available at L for lectotypification. One sheet, L0036159,

has one whole immature fruit and a further dissected immature one, the second sheet,

L0036158, has only an immature fruit, and the last sheet, L0036157, is sterile. The first

specimen is designated here as the lectotype.

Cryptocarya tomentosa Blume, Mus. Bot. 1(21): 335 (1851). - TYPE: [Indonesia]

Java, Patuhae, C.L. Blume s.n. (lectotype L [L0036276] designated here; isolectotype

L [L0036277]).

In the original decription, Blume (1851) mentioned two different gatherings: one from

the mountain Patuhae on Java and one from South Borneo. In L, two specimens of

the former can be identified, L0036276 and L0036277, and one possible one from

Borneo, L0036282. The specimen L0036276 has an old (but not original) label clearly

stating that it was collected at Patuhae, while the second specimen, L0036277, is

much smaller than the first and its label is more recent. The Borneo specimen has no

collection data. The specimen from Java with the older label is designated here as the

lectotype [L0036276].

Lindera lucida (Blume) Boerl., Handl. FI. Ned. Ind. 3: 147 (1900). - Litsea lucida

Blume, Bijdr. FI. Ned. Ind. 11: 562 (1826). - TYPE: [Indonesia] Java, C.L. Blume s.n.

(lectotype L [L0036501], designated here).

In the original description, Blume (1826) only mentioned that the type specimen came

from Java. Several gatherings from Java are present at L which Blume saw and which

could be used for lectotypification (L0036495, L0036496, L0036497, L0036498,

L0036499, L0036500, L0036501, L0036502). One ofthese is almost sterile, L0036495,

while the others have several inflorescences with flowers. Sheet L0036501 bears an

original handwritten note with the local name, which is also mentioned in the original

description and so this specimen is designated here as the lectotype.

New records and typifications for Lauraceae in Singapore

173

Lindera malaccensis Hoolc.f., FI. Brit. India [J.D. Hooker] 5: 183 (1886). - TYPE:

‘Peninsular Malaysia and Birma’, W. Griffith [Kew Distribution no.] 4297 (lectotype

K [K000815598], designated here).

Several gatherings are mentioned in the original description of Lindera malaccensis

Hook.f. (Hooker, 1886: 183): W. Griffith [Kew Distribution no. 4297]; A.C. Maingay

1988 [Kew Distribution no. 1257]; A.C. Maingay 3095 [Kew Distribution no. 1257];

A.C. Maingay 1781 [Kew Distribution no. 1257]; A.C. Maingay, [Kew Distribution

no. 1272]. As Hooker worked at Kew, the K specimens are most suitable for

lectotypification. Of these, only the Griffith specimen [Kew Distribution no. 4297] has

the characteristic card with flower dissections, notes and signature of Gamble and it is

designated here as the lectotype.

Litsea accedens (Blume) Boerl., Handl. FI. Ned. Ind. 3: 145 (1900). - Tetranthera

accedens Blume, Mus. Bot. 1(24): 383 (1851). - TYPE: [Indonesia, Sulawesi]

Celebes, Tondano, July 1840, E.A. Forsten s.n. (lectotype L [L0036569], designated

here; isolectotypes L [L0036568, L0036570]).

In the orginal description of Tetranthera accedens Blume (Blume, 1851) no mention

is made of any collectors or numbers but two localities are given: Borneo and Celebes.

It appears that he was referring to the following gatherings: Korthals s.n. from Borneo

[U0002782] and Forsten s.n. from Tondano (now in Sulawesi Utara), collected in July

1840 [L0036568, L0036569, L0036570]. Most of these collections are sterile. The one

Forsten gathering with flowers, L0036569, is designated here as the lectotype.

Litsea amara Blume var. attenuata Gamble, J. Asiat. Soc. Bengal 75: 142 (1912). -

TYPE: [Peninsular Malaysia] Malacca, Batu Tiga, 1892, R. Derry 990 (lectotype K

[K000797034], designated here; isolectotypes BM, SING [SING0229216]).

There are ten gatherings mentioned in the original description of Litsea amara Blume

var. attenuata Gamble (Gamble, 1912: 142): H.N. Ridley 2265, 9473, 13781; King’s

Collector 4614, 8750, 10142; Burn-Murdoch 4; A.C. Maingay 1266, 1278; R. Derry

990. These are all very similar to one another and as Gamble was working at K the

time, the K specimen of R. Derry 990 is designated here as the lectotype.

Litsea gracilis Gamble, Bull. Misc. Inform. Kew 317 (1910). - TYPE: [Peninsular

Malaysia] Perak, Ulu Temango, July 1909, H.N. Ridley 14603 (lectotype

K [K000797038], designated here; isolectotypes K [K000797039], SING

[SING0055927]).

174

Card. Bull. Singapore 69(2) 2017

There are two sheets at K of the one gathering cited in the original description of Litsea

gracilis Gamble (Gamble, 1910: 317). The sheet bearing a card with notes, dissections

and the signature of Gamble, K000797038, is designated here as the lectotype.

Litsea gracilipes Hook.f., FI. Brit. India [J.D. Hooker] 5: 159 (1886). - TYPE:

[Peninsular Malaysia] Malacca, W. Griffith [Kew Distribution no.] 4311 (lectotype K

[K000797135], designated here; isolectotypes K [K000797136], P [P00745274])

In the original description of this name, only one gathering, with male flowers, was

cited: W. Griffith [Kew Distribution no. 4311] from Malacca (Hooker, 1886: 159-160).

In K, two specimens of this gathering are present, K000797136 and K000797135, and

the one with the dissection of male flowers and notes on the sheet, K000797135, is

designated here as the lectotype.

Litseagriffithii Gamble, J. Asiat. Soc. Bengal 75: 187-188 (1912). -TYPE: Singapore,

Reservoir Woods, 1893, H.N. Ridley 4823 (lectotype K, designated here; isolectotype

SING [SING0043342]).

In the original description of this name, many Peninsular Malaysian, Singaporean and

Sumatran gatherings are mentioned: Burn-Murdock 282 (Ridley 14284), Forbes 1762,

W. Griffith [4285], King’s Collector 5865, 8573, King & Hullett s.n., Maingay 12521,

Ridley 4823, 5569, and Wray 2757 (Gamble, 1912: 187-188). The gathering from the

Reservoir Woods in Singapore, Ridley 4823, housed at K, is designated here as the

lectotype.

Litsea lanceolata (Blume) Kosterm., Reinwardtia 7: 348 (1968). -Aperula lanceolata

Blume, Mus. Bot. 1(27): 367 (1851). - TYPE: [Indonesia] Java, C.L. Blume s.n.

(lectotype U [U0002822], designated here; isolectotype U [U0002823]).

In the original publication of this name, Blume (1851: 367) gave the collection data

only ‘in Java \ There are two possible sheets available at U for lectotypification,

one only has a single sterile leaf [U0002823] and the other has a twig with several

leaves and a few small inflorescences [U0002822], The latter is designated here as the

lectotype.

Litsea machilifolia Gamble, Bull. Misc. Inform. Kew 320 (1910); Ng, Gard. Bull.

Singapore 57: 234 (2005). - TYPE: [Peninsular Malaysia] Penang, MoniotRoad, June

1886, C. Curtis 795 (lectotype SING [SING 0068773], first step designated by Ng

(2005), second step lectotypification designated here).

New records and typifications for Lauraceae in Singapore

175

In their paper dealing with the species of Litsea from Thailand, Ngernsaengsaraay et al.

(2011: 70) selected Ridley 4706 at K as the lectotype of L. machilifolia. However, six

years before, Ng (2005: 238), in his paper dealing with the species from Borneo, had

already selected a different specimen (Penang, Curtis 795 at SING) as the lectotype

for this name. There are, however, two sheets at SING with this number and locality:

Penang, Moniot Road, April 1886, Curtis 795 [SING 0055932] with flowers, and

Penang, Moniot Road, July 1886, Curtis 795 [SING 0068773] with fruits. As only one

of these can serve as the type, and as fruits are more important than flowers in species

recognition within Litsea , the fruiting specimen is designated here in a second step

lectotypification (see McNeill et al., 2012: Art. 9.17, Ex. 12).

Litsea machilifolia Gamble var. angustifolia Gamble, J. Asiat. Soc. Bengal 75: 172

(1912). - TYPE: [Peninsular Malaysia] Perak, Larut, Nov. 1884, King ’s Collector 6815

(lectotype K designated here; isolectotypes SING [SING0229214, SING0229215]).

In his description of this variety, Gamble (1912: 172) mentioned t hir teen very s im ilar

separate gatherings, for which there are often duplicates in several other herbaria. As

Gamble was working at K when he wrote his paper, the K specimen of Kings Collector

6815 is designated here as the lectotype.

Litsea megacarpa Gamble, Bull. Misc. Inform. Kew 364 (1910). - TYPE: Singapore,

at Toas, 1894, H.N. Ridley 6456 (lectotype K, designated here; isolectotypes BM,

SING [SING0013205])

Twenty very similar gatherings, for which there are often duplicates in several herbaria,

were cited by Gamble when he published the name Litsea megacarpa (Gamble, 1910:

364-365): Curtis 2511; Wray 2337, 2730; Scortechini 265, 270, 589; King’s Collector

3697, 6076, 6124, 6155, 6237, 6409, 6567, 6613, 6689, 6774, 6866, 7204; Ridley 6455

and 7621. As Gamble was working at K when he wrote this paper, the K specimen

of the gathering from Singapore, Toas, 1894, Ridley 6456 is designated here as the

lectotype.

Litsea pustulata Gamble, Bull. Misc. Inform. Kew 359 (1910). - TYPE: [Peninsular

Malaysia] Perak, Larut, Nov. 1883, King’s Collector5140 (lectotype K [K0000797089],

designated here; isolectotype K).

Several morphologically very similar gatherings were mentioned in the original

description of Litsea pustulata by Gamble (1910: 359-360): King’s Collector 2544,

3418 and 5140. The specimen with the most fruits of King’s Collector 5140 housed at

K is designated here as the lectotype.

176

Card. Bull. Singapore 69(2) 2017

Litseasarawacensis Gamble, Bull. Misc. Inform. Kew365 (1910). -TYPE: Singapore,

Reservoir Woods, July 1893, H.N. Ridley 4S7 7 (lectotype K [K000797140], designated

here; isolectotypes B, SING [SING 0011250]).

In the original description of Litsea sarawacensis Gamble (Gamble, 1910: 365-366),

several morphological very similar gatherings were cited: Ridley 4817; Beccari 905,

1475, 1708, 1796; Havilland 3077 and 3646. The K sheet of the gathering from

Singapore, Ridley 4817, is designated here as the lectotype.

Litsea singaporensis Gamble, Bull. Misc. Inform. Kew 358 (1910). - TYPE:

Singapore, Garden Jungle, Nov. 1892, H.N. Ridley 4135 (lectotype K, designated here;

isolectotypes BM, SING [SING0000630]).

Numerous gatherings were mentioned in the original description of Litsea singaporensis

by Gamble (1910: 358): H.N. Ridley 2118, 3372, 3894, 4135, 4826, 5065, 5736, 5738,

9075 and R. W. Hullett 5738. I was unable to locate two of these (H.N. Ridley 2118 and

R.W. Hullett 5738). As Gamble was working at Kew at the time he wrote this article,

the K specimen of H.N. Ridley 4135 is designated here as the lectotype.

Litsea umbellata (Lour.) Merr., Philipp. J. Sci. 14: 242 (1919). -Hexanthus umbellatus

Lour., FI. Coc hin ch 196 (1790). - TYPE: Cochinchina, Loureiro s.n. (lectotype BM

[BM000951060], designated here; isolectotype BM [BM000951061]).

Two sheets of type material of Hexanthus umbellatus are extant at BM. One has flowers,

leaves and twigs [BM000951060] and the other has several loose leaves, a twig and

some flowers [BM000951061]. The first sheet is designated here as the lectotype.

ACKNOWLEDGEMENTS. The author is grateful to the curators of the BM, GH, K, KEP,

L, P, U and SING herbaria for access to the specimens used in the present study. I am very

grateful to Helen Hopkins for checking my English. This research received support from the

SYNTHESYS Project (http://www.synthesys.info/), which is financed by the European Union

Research Infrastructure Action under the FP7 “Capacities Program”. This work was also

supported by a Visiting Research Fellowship at FRIM and two Research Fellowships at the

Singapore Botanic Gardens, which are gratefully acknowledged.

References

Blume, C.L. (1826). Bijdragen tot de Flora van Nederlansch Indie, ll e stuk. Batavia: Ter

Lands Drukkerij.

Blume, C.L. (1851). Museum Botanicum Lugduno-Batavum, vol. 1. Leiden: E.J. Brill.

Blume, C.L. (1856). Museum Botanicum Lugduno-Batavum, vol. 2. Leiden: E.J. Brill.

New records and typifications for Lauraceae in Singapore

111

De Kok, R.P.J. (2015). Cryptocarya nitens (Lauraceae), a new species record for Singapore.

Gard. Bull. Singapore 67: 253-260.

De Kok, R.P.J. (2016a). A revision of Beilschmiedia (.Lauraceae ) of Peninsular Malaysia.

Blumea 61: 147—164.

De Kok, R.P.J. (2016b). A revision of Cryptocarya R.Br. (Lauraceae) of Peninsular Malaysia.

Kew Bull. 71(7): 1—16.

De Kok, R., Sengun, S. & Bramley G.L.C. (2016). Two new records for the Lamiaceae of

Singapore. Gard. Bull. Singapore 67: 189-200.

Gamble, J.S. (1910). New Lauraceae from the Malayan region I. Bull. Misc. Inform. Kew:

142-153.

Gamble, J.S. (1912). Materials for a Flora of the Malayan Peninsula no 22. J. Asiat. Soc.

Bengal, Pt. 2, Nat. Hist. 75: 1-204.

Hooker, J.D. (1886). Flora British India, vol. 5. London: L. Reeve.

IUCN (2017). The IUCN Red List of Threatened Species, version 2017-1. http://www.

iucnredlist.org. Accessed 1 Jul. 2017.

McNeill, J., Buck, W.R., Demoulin, V., Greuter, W., Hawkworth, 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, vol. 154. Konigstein: Koeltz Scientific Books.

Miquel, F.A.W. (1858). Flora van Nederlandsch Indie, vol. 1(1), part 6. Amsterdam: C.G. Van

der Post and Utrecht: C. Van der Post Jr.

Ng, F.S.P. (2005). Taxonomic Notes on Bornean Litsea, Lindera, Neolitsea and Iteadaphne

(Lauraceae). Gard. Bull. Singapore 57: 217-246.

Ngernsaengsaruay, C., Middleton, D.J. & Chayamarit, K. (2011). A revision of the genus Litsea

Lam. (Lauraceae) in Thailand. Thai Forest Bull, Bot. 39: 40-119.

Van Steenis-Kruseman, M.J. (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. In: van Steenis, C.G.G.J. (ed.) Flora Malesiana, ser. 1, vol. 1, pp.

3-639. Jakarta: Noordhoff-Kolff N.V.

Gardens' Bulletin Singapore 69(2): 179-187. 2017

doi: 10.26492/gbs69(2).2017-03

179

Novitates Bruneienses, 9. A synopsis of Epirixanthes

(Polygalaceae) in Brunei Darussalam

and notes on species elsewhere

M. Dancak 1 , M. Hrones 2 , R.S. Sukri 3 , F. Metali 3 & A.A. Joffre 4

department of Ecology & Environmental Sciences, Faculty of Science, Palacky University

Slechtitelu 27, CZ-78371 Olomouc, Czech Republic

martin. dancak@upol. cz

department of Botany, Faculty of Science, Palacky University

Slechtitelu 27, CZ-78371 Olomouc, Czech Republic

Environmental and Life Sciences Programme, Faculty of Science, Universiti Brunei

Darussalam, Jalan Tungku Link, BE 1410, Brunei Darussalam

4 Brunei National Herbarium, Forestry Department, Ministry of Primary Resources and

Tourism, Jalan Menteri Besar, Berakas, BB3910 Brunei Darussalam

ABSTRACT. The genus Epirixanthes Blume is revised for Brunei Darussalam. Four

species are recognised for the country: Epirixanthes cylindrica Blume, E. elongata Blume,

E. kinabaluensis T.Wendt and E. papuana J.J.Sm., with the two latter species being newly

recorded for the Brunei flora. A single collection from Brunei that was formerly identified as

Epirixanthes pallida T.Wendt is now confirmed as E. papuana. A revised key for the genus is

included.

Keywords. Distribution, herbs, Malesia, mycoheterotrophic plants, north-western Borneo,

taxonomy, understorey

Introduction

Epirixanthes Blume is a species-poor genus of holomycoheterotrophic herbaceous

plants from the family Polygalaceae which inhabit the understorey of tropical

rainforests (Van der Meijden, 1988; Merckx et al., 2013). It is sister to the autotrophic

genus Salomonia Lour, with which it shares several synapomorphies such as spilce-like

terminal inflorescence and three antesepalous stamen primordia (Van der Meijden,

1988; Mennes et al., 2015). Members of the genus are generally tiny plants with

reduced bract-like leaves and dense spike-like inflorescences (Van der Meijden, 1988).

Due to the reduction of vegetative organs as a result of mycoheterotrophy, the set of

morphological characters useful for species determination is very limited and includes

mostly reproductive organs (i.e., bracts, bracteoles, sepals and fruits). Currently, only

seven species of Epirixanthes are recognised worldwide, and all of them are endemic

to Indo-Malesia (Van der Meijden, 1988; Pendry, 2010; Tsukaya et al., 2016). The

centre of its taxonomic diversity lies in Borneo where six of the species co-occur, with

one of them apparently endemic to the island (Fig. 1). In the Checklist of the Flowering

180

Card. Bull. Singapore 69(2) 2017

Fig. 1 . Global distribution and diversity of Epirixanthes with number of species for each region

and number of endemic species in parentheses.

Plants and Gymnosperms of Brunei Darussalam (Coode et al., 1996), three species of

Epirixanthes are reported from the country, namely E. cylindrica Blume, E. elongata

Blume and E. pallida T.Wendt. During our recent ecological research activities in

Kuala Belalong (Ulu Temburong National Park, Temburong district), we frequently

encountered Epirixanthes plants in the lowland dipterocarp forest understorey.

However, some of the observed plants did not match with any of the three species

reported from Brunei Darussalam by Coode et al. (1996), indicating they possibly

belong to unrecorded species. We therefore decided to clarify the taxonomic status of

these plants and present a revision of Epirixanthes in the country.

Material and methods

We examined specimens deposited in BRUN (Brunei National Herbarium) as well

as our field collections from Brunei Darussalam deposited in OL (Herbarium of the

Department of Botany at Palacky University in Olomouc). Available duplicates kept

in K (Royal Botanic Gardens Kew) and AAU (Science Museums, Aarhus University)

Epirixanthes in Brunei Darussalam

181

were also seen. The identification of each specimen was checked and the identifications

updated when necessary. All cited specimens have been seen. Bruneian localities from

herbarium labels are sorted below according the current administrative districts of

Brunei Darussalam.

Results

Four species of Epirixanthes were found to occur in Brunei Darussalam, namely E.

cylindrica, E. elongata, E. kinabaluensis T.Wendt and E. papuana J.J.Sm. The two

former species were previously known from the country while the two latter species

are new additions to the country’s flora. Epirixanthes pallida should be excluded from

the Brunei checklist as all three duplicates (deposited in BRUN, K, and AAU) of the

single collection ( Poulsen 3) cited in Coode et al. (1996) consist of misidentified plants

of E. papuana. All four species co-occur in the Towland Mixed Dipterocarp forest of

the Ulu Temburong National Park in the Temburong district, which further highlights

the floristic and conservation significance of this area.

The Epirixanthes species of Brunei Darussalam

1. Epirixanthes cylindrica Blume, Cat. Gew. Buitenzorg 82 (1823). (Fig. 2A)

Global distribution. Its range includes Myanmar, Sumatra, Java, Borneo and New

Guinea (Van der Meijden, 1988).

Distribution in Brunei Darussalam. It is documented only from the Temburong district

in a few localities in Kuala Belalong - Bukit Belalong area. Its known elevational

range in Brunei Darussalam is from c. 100 m a.s.l. up to 850 m a.s.l.

Specimens examined. BRUNEI DARUSSALAM: Temburong: Amo, Ulu Belalong, L.P. 382,

north-facing slopes below campsite, 18 Jan 1994, Coode 7845 (BRUN); Amo, Bukit Belalong,

hill dipterocarp forest on Setap Shales ridgetop, 24 Feb 1992, Dransfield 1246 (BRUN); Amo,

Bukit Belalong, hill dipterocarp forest steep slope near valley bottom, 25 Feb 1992, Dransfield

1266 (BRUN); Bukit Belalong, east ridge, 20 Jul 1989, de Vogel 8989 (BRUN); Batu Apoi

Forest Reserve, ridge between K. Belalong and Bt. Belalong, Jalan Tengga, damp, flat area

below Bukit Belalong, 21 Mar 1992, Poulsen 304 (AAU, BRUN); Kuala Belalong, Batu Apoi

Forest Reserve, upstream the Sungai Belalong river from the Kuala Belalong Field Studies

Centre, along path, 10 Nov 1991, Hansen 1540 (BRUN); Kuala Belalong, E ridge of Sungai

Belalong, ca. 0.9 km ESE from its confluence with Sungai Temburong, ecological plot 1,13

Jan 2014, Dancdk 2014/161 (OL); ibidem, 30 Jan 2016, Dancdk 2016/367 (OL).

Notes. This species is distinguished by a rather short and thick inflorescence. The apex

of the inflorescence is covered by imbricate bracts. As most of the specimens come

from middle altitudes this species seems to be rare in true lowlands.

182

Card. Bull. Singapore 69(2) 2017

Fig. 2. Epirixanthes species of Brunei Darussalam. A. Epirixanthes cylindrica Blume. B. E.

elongata Blume. C. E. kinabaluensis T.Wendt. D. E. papuana J.J.Sm. E. E. pallida T.Wendt.

A-D, all from Kuala Belalong, Brunei Darussalam; E from Kelabit Highlands, Sarawak. (Pho¬

tos: A: Ondrej Popelka, B, C: Michal Hrones, D, E: Michal Sochor)

2. Epirixanthes elongata Blume, Cat. Gew. Buitenzorg 82 (1823). (Fig. 2B)

Global distribution. The species range extends from eastern India and southern China

to the Moluccas (Van der Meijden, 1988; Chen et al., 2008).

Distribution in Brunei Darussalam. It is documented from various locations across

the Temburong district. Outside Temburong, it has only been collected from a single

locality around Labi in the Belait district. Its known elevational range in Brunei

Darussalam is from c. 40 m a.s.l. up to 420 m a.s.l.

Epirixanthes in Brunei Darussalam

183

Specimens examined. BRUNEI DARUSSALAM: Belait: Labi, Sungai Rampayoh, ca. 3.5

km above road towards Waterfall No. 2, 9 Jan 1994, Coode 7784 (BRUN). Temburong:

Batu Apoi Lorest Reserve, ridge W of Kuala Belalong Lield Studies Centre, in Danish Plot,

mixed dipterocarp forest, 1991, Poulsen 224 (AAU, BRUN); Kuala Belalong, E ridge of

Sungai Belalong, ca. 1.5 km SE from its confluence with Sungai Temburong, depression NW

of ecological plot 2, 13 Feb 2015, Hedl & Chudomelova RH132015 (OL); Kuala Belalong,

E ridge of Sungai Belalong, ca. 0.9 km ESE from its confluence with Sungai Temburong,

ecological plot 1,13 Jan 2014, Dancdk 2014/6 (OL); ibidem, 13 Jan 2014, Dancdk 2014/125

(OL); ibidem, 13 Jan 2014, Dancdk 2014/160 (OL); ibidem, 13 Jan 2014, Dancdk 2014/180

(OL); ibidem, 4 Feb 2015, Hrones & Kobrlovd 702015 (OL); Sungai Temburong at Kuala

Belalong, 23 Jun 1989, Dransfield 1007 (BRUN); Apan, ridge to the north of the river, 13 Jul

1993, Sands 5786 (BRUN); Amo, southeast of LP 297 Bkt. Lutut, 6 Apr 2004, Ariffin et al.

BRUN20797 (BRUN); Labu, Peradayan F. R., 5 Feb 2002, Ariffin et al BRUN 19913 (BRUN).

Notes. This is the most common species of Epirixanthes. It is locally abundant

elsewhere in Borneo, and is presumably also common in Brunei Darussalam. In Kuala

Belalong it is almost ubiquitous and by far the most abundant species of Epirixanthes.

This species is easily recognised among Bruneian Epirixanthes as its bracts are shed

well before the flowers open and the inflorescence is very long and narrow. One of

the specimens studied ( Hrones & Kobrlovd 702015) is a very pale-coloured plant

conspicuously different from the typical brownish-purple plants of E. elongata. Such

pale ivory individuals are known to occur within populations of E. elongata and they

were described from West Kalimantan as E. elongata f. alba Tsukaya & H.Okada

(Tsukaya & Okada, 2012).

3. Epirixanthes kinabaluensis T.Wendt, FI. Males., Ser. 1, Spermat. 10(3): 491 (1988).

(Fig. 2C)

Global distribution. The species is found in Sumatra and Borneo (Van der Meijden,

1988).

Distribution in Brunei Darussalam. It is known only from two locations in the

immediate vicinity of the Kuala Belalong Field Studies Centre in the Temburong

district. Its known elevational range in Brunei Darussalam is very narrow as it is

recorded only from altitudes around 100 m a.s.l., even though its type locality, the

slopes of Mt. Kinabalu in Sabah, lies at around 900-1200 m a.s.l. We have observed

the species in the Kelabit Highlands of Sarawak up to c. 1300 m a.s.l.

Specimens examined. BRUNEI DARUSSALAM: Temburong: Kuala Belalong, Sungai Esu

valley, at its confluence with Sungai Belalong, 9 Jan 2014, Dancdk 2014/162 (OL); ibidem, 18

Jan 2014, Dancdk 2014/343 (OL); Kuala Belalong, Sungai Esu, clayey bank near its confluence

with Sungai Belalong, 21 Jan 2017, Dancdk 2017/46 (BRUN); Kuala Belalong, Earthwatch

ecological plot ca. 0.3 km W from the Kuala Belalong Field Studies Centre, 28 Jan 2016,

Dancdk 2016/292 (OL).

184

Card. Bull. Singapore 69(2) 2017

Notes. This species is the most robust of all Bruneian Epirixanthes, although E.

elongata is usually taller. It has a rather thick inflorescence with long bracts which

sometimes persist on the axis of the inflorescence after the fruits are shed.

4 .Epirixanthespapuana J.J.Sm., Repert. Spec. Nov. Regni Veg. 10: 486 (June 1912).

(Fig. 2D)

Global distribution. It is distributed throughout Malesia from Sumatra to the Solomon

Islands (Van der Meijden, 1988).

Distribution in Brunei Darussalam. It is known only from a few locations immediately

surrounding the Kuala Belalong Field Studies Centre in Temburong district and one

locality in the Bukit Sawat area in Belait district. However, its actual distribution in the

country could be much more extensive. Like most of the other species of Epirixanthes ,

this species is likely to be overlooked by collectors and therefore poorly represented in

herbaria. Its known elevational range in Brunei Darussalam is rather narrow, reaching

from c. 20 m a.s.l. up to 180 m a.s.l. However, the species might potentially also

occur at higher elevations because the type collection from New Guinea comes from

altitudes from 500 m a.s.l. up to 1800 m a.s.l.

Specimens examined. BRUNEI DARUSSALAM: Belait: Labi, Bukit Sawat, Sungai Malayan,

10 Jun 2003, Ariffin et al. BRUN20353 (BRUN). Temburong: Batu Apoi Forest Reserve, ridge

W of Kuala Belalong Field Studies Centre, mixed dipterocarp forest, 18 Mar 1991, Poulsen

3 (AAU, BRUN, K); Kuala Belalong, in Sungai Mata Ikan gorge near its confluence with

Sungai Belalong, 26 Jan 2015, Hrones & Kobrlova 662015 (OL); ibidem, 21 Jan 2017, Dancdk

2017/55 (BRUN); Kuala Belalong, Sungai Mata Ikan valley ca. 0.5 km WNW from Kuala

Belalong Field Studies Centre, 28 Jan 2016, Dancdk, M. 2016/291 (OL); Kuala Belalong,

Sungai Bald valley near its confluence with Sungai Temburong, 18 Jan 2014, Dancdk 2014/342

(OL); Kuala Belalong, Sungai Esu, clayey bank near its confluence with Sungai Belalong, 21

Jan 2017, Dancdk 2017/47 (BRUN).

Notes. This species is distinguished by its paler colour (creamy brownish) and the

rather short and narrow inflorescence with patent bracts that are tumed-up at their

apices.

Notes on other species of Epirixanthes

Epirixanthes pallida T.Wendt, FI. Males., Ser. 1, Spermat. 10(3): 492 (1988). (Fig.

2E)

Global distribution. The species is found in Borneo and Sulawesi (Van der Meijden,

1988).

Epirixanthes in Brunei Darussalam

185

Notes. The species was included in the Brunei Checklist (Coode et al., 1996) based

on a single collection (Poulsen 3) originally identified as Epirixanthes pallida but

which has been reidentified as E. papuana. Therefore Epirixanthes pallida should

be excluded from the flora of Brunei Darussalam. Epirixanthes pallida is readily

distinguished from the Bruneian species by large and very early caducous whitish

or pinkish bracts. It seems that Epirixanthes pallida is a highland species and thus

probably does not grow in lowland rainforests. Its known altitudinal range is from

c. 600 m a.s.l. at the type locality up to c. 1200 m a.s.l. in the Kelabit Highlands of

Sarawak. As its type locality (slopes of Gunung Api in Sarawak, Malaysia) is just a

few kilometres from Brunei’s borders, it might potentially occur at higher altitudes in

the Temburong district. It is, however, unlikely that Epirixanthes pallida occurs in the

Tutong and Belait districts because of the generally lower terrain in these areas. Even

though the type locality is on a limestone bedrock, we have observed the species in the

Kelabit Highlands in Sarawak at several localities on sandstone bedrock.

Epirixanthes confusa Tsukaya et al., Phytotaxa 266(2): 147 (2016).

Global distribution. It is endemic to Borneo (Tsukaya et al., 2016).

Notes. This species is a recent addition to the genus and was described from Imbak

Canyon in Sabah, Malaysia (Tsukaya et al., 2016). It undoubtedly belongs to the group

of species with free sepals and wide fruits which also includes Epirixanthes elongata,

E. pallida and E. comp res sa Pendry, but differs by its long-persistent bracts. So far it

is known only from the type locality.

Epirixanthes compressa Pendry, Thai Forest Bull., Bot. 38: 184 (2010).

Global distribution. It is endemic to Thailand (Pendry, 2010).

Notes. This biogeographically remarkable species is narrowly distributed in south¬

eastern Thailand (Chanthaburi Province) and is, therefore, the only member of the

genus which does not occur in Malesia. Tike the previous species, it belongs to the

group of Epirixanthes with free sepals and wide fruits.

Key to the species of Epirixanthes

The following key is adapted from Van der Meijden (1988), Pendry (2010) and Tsukaya

et al. (2016).

la. Sepals free; fruit wider than long.2

lb. Sepals connate for V^-A; fruit longer than wide or equal.5

186

Card. Bull. Singapore 69(2) 2017

2a. Bracts caducous before the flowers open.3

2b. Bracts persistent at least until the fruits mature.4

3a. Both leaves and bracts hairy with glandular hairs; inflorescence apex ± pointed,

not fully covered by bracts; bracts narrowly triangular, c. 1.2 mm long and 0.5 mm

wide, brown with central purple stripe. E. elongata

3b. Both leaves and bracts glabrous; inflorescence apex ± rounded, completely

covered in imbricate bracts; bracts ovate, 2-3 mm long and 1-1.5 mm wide,

whitish or pinkish. E. pallida

4a. Bracts glabrous, some of them persistent after the fruits have fallen; inflorescence

elongate, 2-6 cm long; corolla caducous; endemic to Sabah. E. confusa

4b. Bracts minutely ciliate, all of them shed with fruits; inflorescence shortly

cylindrical, up to 2.5 cm long; corolla persistent; endemic to Thailand.

. E. compressa

5a. Each flower subtended by a bract and pair of subulate bracteoles; bracts ± patent

during anthesis, linear with sharply upturned obtuse apex; fruit longer than wide,

much shorter than sepals. E. papuana

5b. Each flower subtended only by a bract, bracteoles absent; bracts ± erect during

anthesis, with straight acute or slightly curved apex; fruit as long as wide, as long

as sepals or longer.6

6a. Bracts lanceolate, 2-2.5 mm long and 0.5-0.6 mm wide, brown with purple

central stripe, usually persistent after the fruits have fallen; inflorescence apex not

completely covered by bracts; fruit as long as sepals. E. kinabaluensis

6b. Bracts ovate, 1.5-2 mm long and 0.8-1.2 mm wide, brown with basal purple

patch, usually shed with fruits; inflorescence apex completely covered by

imbricate bracts; fruit longer than sepals. E. cylindrica

Conservation status of Epirixanthes

The forests of Borneo, including Brunei, Sabah and Sarawak, harbour the highest

diversity of Epirixanthes. Six species have so far been reported from this island

(Van der Meijden, 1988; Coode et al., 1996; Tsukaya et al., 2016). Similar to other

mycoheterotrophic plants, all species of Epirixanthes are closely tied to primary or

very lightly logged forests and as such are threatened by continuing deforestation.

However, published data on their distribution and ecology are very scarce and most

of the species are poorly represented in herbaria. For these reasons, we propose that

all species of Epirixanthes in this study be evaluated as data deficient (DD, IUCN

Standards and Petitions Subcommittee, 2016) which emphasises the need for further

study and sampling.

Epirixanthes in Brunei Darussalam

187

ACKNOWLEDGEMENTS. We are very grateful to the staff of the Kuala Belalong F ield Studies

Centre and the Brunei National Herbarium for their service and support. We are indebted to

Salwana Jaafar, Hazimah Din and a group of Universiti Brunei Darussalam students for field

assistance and logistical arrangements. We also thank Radim Hedl, Marketa Chudomelova,

Michal Sochor, Lucie Kobrlova and Ondrej Popelka for their field assistance. We thank

Universiti Brunei Darussalam and the Brunei Forestry Department for permission to conduct

research at KBFSC and in the Ulu Temburong National Park respectively, and the Biodiversity

Research and Innovation Centre (BioRIC) for granting our export permit. Researchers from

Palacky University were supported by project no. CZ. 1.07/2.2.00/28.0149.

References

Chen, S.K., Ma, H.-Y. & Parnell J.A.N. (2008). Polygalaceae. In: Wu, Z.Y., Raven, PH. &

Hong, D.Y. (eds) Flora of China, vol. 11, pp. 139-159. Beijing and St. Louis: Science

Press.

Coode, M.J.E., Dransfield, J., Forman, L.L., Kirkup, D.W. & Idris, M.S. (1996). A Checklist

of the Flowering Plants and Gymnosperms of Brunei Darussalam . Brunei Darussalam:

Ministry of Industry and Primary Resources.

IUCN Standards and Petitions Subcommittee (2016). Guidelines for Using the IUCN

Red List Categories and Criteria, ver. 12. http://www.iucnredlist.org/documents/

RedListGuidelines.pdf.

Mennes, C.B., Moerland, M.S., Rath, M., Smets, E.F., Merckx, V.S.F.T. (2015). Evolution of

mycoheterotrophy in Polygalaceae: The case of Epirixanthes. Am. J. Bot. 102: 598-608.

Merckx, V.S.F.T., Freudenstein, J.V., Kissling, J., Christenhusz, M.J.M., Stotler, R.E., Crandall-

Stotler, B., Wickett, N., Rudall, P.J., Maas-van de Kamer, H. & Maas, P.J.M. (2013).

Taxonomy and classification. In: Merckx, V.S.F.T. (ed.) Mycoheterotrophy: The biology

of plants living on fungi, pp. 19-102. New York: Springer.

Pendry, C.A. (2010). Epirixanthes compressa Pendry, a new mycoheterotrophic species of

Polygalaceae from Thailand. Thai Forest Bull, Bot. 38: 184-186.

Tsukaya, H. & Okada, H. (2012). A color variation of Epirixanthes species (Polygalaceae)

found in West Kalimantan, Borneo, Indonesia. Acta Phytotax. Geobot. 62: 95-97.

Tsukaya, H., Suleiman, M. & Okada, H. (2016). Anew species of Epirixanthes (Polygalaceaea)

from Imbak Canyon, Sabah, Borneo. Phytotaxa 266: 146-150.

Van der Meijden, R. (1988). Polygalaceae. In: van Steenis, C.G.G.J. & de Wilde, W.J.J.O.

(eds) Flora Malesiana, ser. I, vol. 10(1), pp. 455-539. Dordrecht, Boston and London:

Kluwer Academic Publishers.

Gardens' Bulletin Singapore 69(2): 189-199. 2017

doi: 10.26492/gbs69(2).2017-04

189

A new species of Zingiber (Zingiberaceae) east of

Wallace’s Line

M. Ardiyani 1 , M.F. Newman 2 & A.D. Poulsen 2

herbarium Bogoriense, Research Center for Biology, Indonesian Institute of Sciences,

Jl. Raya Bogor Km. 46, Cibinong 16912, Indonesia

marlina.ardiyani@lipi.go.id

2 Royal Botanic Garden Edinburgh, 20A Inverleith Row,

Edinburgh EH3 5LR, Scotland, UK

ABSTRACT. Zingiber Mill, is distributed from India to the Pacific but only a few species

are known from east of Wallace’s Line, whereas the area to the west is rich in species. A

recent collection from limestone at Bantimurung, South Sulawesi, Indonesia represents a new

eastern species, Zingiber ultralimitale Ardiyani & A.D.Poulsen, which is described, illustrated,

and barcoded using three of the four barcoding loci (rbcL, trnR-psbA and ITS). Placement of

this species using morphological evidence is ambiguous but a combination of evidence from

morphology, pollen anatomy and molecular analysis indicates that it belongs to Zingiber sect.

Zingiber.

Keywords . Bantimurung, DNA barcode, Indonesia, limestone, Sulawesi, Wallacea

Introduction

Species of Zingiber Mill, occur from India in the west, through Malesia to the western

Pacific, and those east of Wallace’s Line are poorly known. There are about 180 species

of Zingiber in the world (Zingiberaceae Resource Centre, 2017). Of the 45 species and

six varieties of Zingiber occurring in Indonesia west of Wallace’s Line (Ardiyani et al.,

unpublished), four cross Wallace’s Line, all of them cultivated (Z officinale Roscoe, Z.

montanum (J.Koenig) A.Dietr., Z. odoriferum Blume, Z. zerumbet (L.) Sm.), indicating

that their occurrence to the east may be anthropogenic.

Zingiber is currently classified into four sections based on the habit of the

inflorescence (Baker, 1894; Schumann, 1904; see Table 1). Most Zingiber species

occurring in Malesia belong to Zingiber sect. Zingiber.

Theilade et al. (1993) studied the pollen morphology of a range of species and

found that it was not congruent with the division of the genus into sections. Their study

showed that in Zingiber sect. Zingiber and Zingiber sect. Dymczewiczia (Horan.)

Benth. & Hook, the pollen grains are spherical with cerebroid sculpturing, whereas in

Zingiber sect. Cryptanthium Horan, they are ellipsoid with spiro-striate sculpturing.

For this reason, Theilade et al. suggested that Zingiber sect. Dymczewiczia should be

included in Zingiber sect. Zingiber.

190

Card. Bull. Singapore 69(2) 2017

The molecular study of Theerakulpisut et al. (2012) supports the classification

of Zingiber into four sections although Zingiber sect. Zingiber and Zingiber sect.

Dymczewiczia are only weakly supported. This weak support does not, therefore,

necessarily refute Theilade’s conclusion that Zingiber sect. Dymczewiczia should be

included in Zingiber sect. Zingiber. These two sections are more closely related to each

other than to Zingiber sect. Cryptanthium and Zingiber sect. Pleuranthesis Benth. &

Hook. (Theerakulpisut et al., 2012).

Plants of an unidentified Zingiber were collected from limestone at Bantimurung,

South Sulawesi in 2009. When they were first encountered, several individuals could

be seen from the road but they were not easy to reach. At first the sterile plants were

thought to be a species of Globba L. different from the only other species known to

occur east of Wallace’s Line, G. marantina L. As this would be a biogeographically

exciting discovery, we decided to try to collect sterile plants for cultivation. Despite

some difficulty because the roots were deeply anchored in cracks in the limestone,

we managed to collect 10 plantlets that were exported to Java, half of which were

deposited in Bogor Botanic Gardens and the other half in the Royal Botanic Garden

Edinburgh (accession numbers 20092015-20092019). Some of the plants flowered in

Edinburgh, enabling us to conclude that it is an undescribed species which we here

name Zingiber ultralimitale, and to make a full description including floral details and

pollen morphology. In addition to describing this species formally, we investigate the

sectional placement of this geographically unusual find using palynological data and

molecular systematic analyses.

Table 1. Sections of Zingiber with their characteristics and geographical distribution (Baker

1894; Schumann 1904; Theerakulpisut et al. 2012).

Section

Geography

Inflorescence

Pollen morphology

Zingiber

India, China, Indo-

China, Thailand,

Peninsular Malaysia,

Sumatra, Borneo, Java,

Cultivated

Borne on radical, erect

peduncle or borne

apically on a leafy stem

Spherical with cerebroid

exine sculpturing

Cryptanthium

India, Thailand, China,

Indo-China

Borne on a radical,

procumbent peduncle

Ellipsoidal with spiro-striate

sculpturing

Dymczewiczia

India, China, Indo-

China, Thailand, Java,

Papua New Guinea

Borne apically on a

leafy stem

Spherical with cerebroid

exine sculpturing

Pleuranthesis

China (Yunnan),

Vietnam

Arising from the side of

the leafy stem

Spherical with cerebroid

exine sculpturing

Zingiber ultralimitale, a new species from Sulawesi

191

Material and methods

Detailed studies of the morphology were made using living collections grown in the

greenhouse of the Royal Botanic Garden Edinburgh. Measurements were made using

a ruler and a calibrated eye piece under a dissecting microscope.

For the DNA barcoding, three barcoding regions were successfully sequenced,

namely rbcL, trnH-psbA and the nuclear Internal Transcribed Spacer (ITS). The

fourth barcoding region, matK could not be sequenced. Extraction, amplification and

sequencing followed Kress et al. (2002) and Kress & Erickson (2007). The material

sequenced was from Poulsen et al. 2767. GenBank accession numbers for the three

barcoding regions are summarised in Table 2. A phylogenetic analysis based on ITS

sequences was performed using additional sequences from NCBI GenBank after

BLAST was done. GenBank has very few records of rbcL and trnH-psbA so it was

not possible to run an analysis using these regions to address the affiliation of Zingiber

ultralimitale. Kaempferiaparviflora Wall, ex Baker and K. elegans (Wall.) Baker were

chosen as the outgroup following Theerakulpisut et al. (2012) (Table 3). Sequence

alignment and Maximum Likelihood analysis were conducted using MEGA version 6

(Tamura et al., 2013).

Table 2. Voucher information and Genbank accession numbers.

Species

Gene region

Genbank accession

number

Material

(Herbarium location)

Zingiber ultralimitale

rbcL

KU891637

Poulsen et al. 2767

trnH-psbA

KU891638

(BO, E)

ITS

KU891639

For the palynological study, material was collected from a living plant growing

at the Royal Botanic Garden Edinburgh (ace. no 20092015 A) and fixed in FAA for

at least 24 hours. Mature but unopened flowers were selected. For SEM, anthers were

dissected from the flowers, put into tiny chambers and dehydrated for 45 minutes

in 50%, 70%, 95%, and 100% ethanol, ten minutes in acetone (two changes of five

minutes each). Critical point drying was carried out in an Emitech K850 dryer and

samples were sputter-coated with platinum and examined using a Zeiss Supra 55VP

SEM.

Results and discussion

Most of the Zingiber species of western Malesia have tightly imbricate bracts, long or

very short peduncles, and lateral staminodes joined to the labellum, although Zingiber

192

Card. Bull. Singapore 69(2) 2017

Table 3. Species of Zingiber and outgroups obtained from NCBI GenBank.

Species

Distribution/Section

GenBank Acc.

No. (ITS)

Zingiber (ingroup)

Z. barbatum Wall.

Thailand/Zingiber

DQ064578

Z bradleyanum Craib

Thailand/Cryptanthium

DQ064579

Z capitatum Roxb.

India, Himalaya/Zingiber

KM983532

Z capitatum

India, Himalaya/Zingiber

KM983536

Z. citriodorum Theilade & Mood

Thailand/Zingiber

DQ064591

Z coloratum N.E.Br.

B orneo/Zingiber

AF414498

Z corallinum Hance

Thailand, China/Zingiber

AL254460

Z corallinum

Thailand, China/Zingiber

DQ064587

Z. ellipticum (S.Q.Tong & Y.M.Xia)

Q.G.Wu & T.L.Wu

Yunnan- China/Pleuranthe si s

AL478799

Z. fragile S.Q.Tong

Thailand/Cryptanthium

DQ064581

Z. gramineum Noronha ex Blume

Indochina, Java,

Sumatra/Zingiber

DQ064577

Z. gramineum

Indochina, Java,

Sumatra/Zingiber

AL478800

Z. isanense Triboun & K.Larsen

Thailand/Zingiber

DQ064586

Z. junceum Gagnep.

Thailand/Zingiber

DQ064588

Z. junceum

Thailand/Zingiber

AY424774

Z. longipedunculatum Ridl.

Borneo/Zingiber

AB097254

Z. montanum (J.Koenig) Link ex

A.Dietr.

Cultivated/Zingiber

KJ872221

Z montanum

Cultivated/Zingiber

DQ064585

Z. neesanum (J.Graham)

Ramamoorthy

India, Indochina/Zingiber

KJ872228

Z neesanum

India, Indochina/Zingiber

KJ872226

Z. neotruncatum T.L.Wu, K.Larsen &

Turland

India, China/Zingiber

DQ064589

Z. newmanii Theilade & Mood

Thailand/Zingiber

DQ064575

Z. ocloriferum Blume

Cultivated/Zingiber

KL304561

Z. ocloriferum

Cultivated/Zingiber

KL304562

Z. officinale Roscoe

Cultivated/Zingiber

DQ064590

Z orbiculatum S.Q.Tong

Thailand/Cryptanthium

DQ064573

Z ottensii Valeton

Cultivated/Zingiber

DQ064582

Z ottensii

Cultivated/Zingiber

AJ388298

Z. parishii Hook.f.

Thailand/Zingiber

KF304567

Z. parishii

Thailand/Zingiber

DQ064576

Z. pellitum Gagnep.

Thailand/Dymczewiczia

DQ064574

Z purpureum Roscoe (syn. of Z.

montanum )

Cultivated/Zingiber

HM236153

Z. rubens Roxb.

Thailand/Cryptanthium

DQ064580

Z. spectabile Griff.

Sumatra, Malay

Peninsula/Zingiber

AL414499

Z. sulphureum Burkill ex Theilade

Sumatra, Malay

Peninsula/Zingiber

AL478801

Z. wrayi Prain ex Ridl.

Sumatra, Malay Peninsula,

Thailand/Zingiber

AL478802

Z. wrayii

Sumatra, Malay Peninsula,

Thailand/Zingiber

DQ064583

Zingiber ultralimitale, a new species from Sulawesi

193

Table 3. Continued.

Species

Distribution/Section

GenBank Acc.

No. (ITS)

Z. zerumbet (L.) Roscoe ex Sm.

Cultivated/Zingiber

KC582863

Z. zerumbet

Cultivated/Zingiber

DQ064584

Z. ultralimitale Ardiyani &

A.D.Poulsen

Sulawesi/Zingiber

KU891639

Z. ultralimitale

Kaempferia (outgroup)

Sulawesi/Zingiber

KU891639

Kaempferia elegans (Wall.) Baker

China, Indochina, Malesia

AY424764

K. parviflora Wall, ex Baker

India, Indochina

DQ064592

engganoense Ardiyani from Enggano Island, Sumatra (Ardiyani, 2015), has a long

peduncle with tightly imbricate bracts and lateral staminodes free from the labellum.

Zingiber ultralimitale , by contrast, has loosely imbricate bracts and relatively large

yellow lateral staminodes which are free from the labellum. Loosely imbricate bracts

are found in Zingiber sect. Cryptanthium west of Wallace’s Line in Asia. Some species

of Zingiber sect. Cryptanthium which resemble Z. ultralimitale are Z. nanlingensis Lin

Chen et al. from China with the same arrangement of inflorescence and yellow flowers

but staminodes not free from the labellum; Zingiber liangshanense Z. Y.Zhu, also from

China, with free staminodes but a purple labellum; and Z. chrysanthum Roscoe from

Nepal with bright red corolla lobes. Zingiber ultralimitale is morphologically distinct

from the other species in Zingiber sect. Zingiber by having the bract arrangement of

Zingiber sect. Cryptanthium.

A phylogenetic analysis of sequence data from the Internal Transcribed Spacer

(ITS) results in an unresolved backbone in the phylogenetic tree of Zingiber. Zingiber

ultralimitale is well supported as belonging to Zingiber sect. Zingiber (Lig. 1).

The pollen grains of Zingiber ultralimitale are spherical with cerebroid

sculpturing (Lig. 2) which, according to Theilade et al. (1993), is characteristic of

Zingiber sect. Zingiber. Lurthermore, the molecular results also place Zingiber

ultralimitale in Zingiber sect. Zingiber (Lig. 1). Taking all the evidence together,

therefore, we place the species in Zingiber sect. Zingiber.

Taxonomy

Zingiber ultralimitale Ardiyani & A.D.Poulsen, sp. nov.

This species is distinct from all others by the following combination of characters:

narrow long loose green bracts, bright yellow flowers, and large free yellow lateral

staminodes.—TYPE: Indonesia, originally collected from South Sulawesi Province,

Bantimurung NP, along Jalan Poros Maros-Soppeng, and cultivated as RBGE

20091017*A, flowering material vouchered on 12 June 2013 as Newman, M.F. 2552

(holotype BO; isotype E). (Fig. 2, 3)

194

Card. Bull. Singapore 69(2) 2017

gg I Zingiber ultrolimitaie ADP2767 1

Zingiber ultrolimitaie ADP2767 2

Zingiber zer umbel KC 582863

new spates

Zingiber zer umbel DQ064584

— Zingibercoloraium AMI4498

— L— Zingiber iongipedunculotum A8097254

- Zingiber wrayi AF478802

- Zingiber wrayi DQQ64533

—- Zingiber newmanii DQ064575

-- Zingiber sulphureum AF4788G1

Zingiber ottensii A J388298

Zingiber ottensii DQ064582

Zingiber spectabile AF414499

Zingiber parishii Kf 304567

- Zingiber parishii DQ064 5 76

Zingiber citrkxhrum DQ064591

WCT

Zingiberjunieum DQ064588

— Zingiberneotruncatum DQ064589

Zingiber gramineum AF47880G

- Zingiber gramineum DQ064577

Zingiber junceum AY424774

— Zingiber corallsmjm AF 254460

— Zingiber coraitim/m DQ064 587

-- Zingiber barbotum DQ064578

- Zingiber rsanense DQ064586

p- Zingiber capitawm KM9835 36

Zingiber capita turn KM983 532

|- Zingiber neesanum KJ872228

' - Zingiber neesanum KJ872 226

p Zingiber odorifetum KF3G4562

- Zingiber odoriferum KF304561

Zingiber monlanum KJS72221

Zingiber purpureum HM236153

sal—

Zingiber monlanum DQ064585

- Zingiber offidnoie DQ064 590

Zingiber peifitum DQ064574 I Section Dymczewiczia

Zingiber ellipticum A F4 78799

— Zingiber bradleyanum DQ064579

100

- Zingiber arbiculatum DQ064573

I-- Zingiber fragile DQ064581

I— Zingiber rubens DQ0645B0

- Kaempferio parviflora DQ064592

(Section Pleuranthesis

Section Cryptanthium

Kpempferia ehgans AY424764

0.01

I- 4

Fig. 1. Phylogenetic tree of Zingiber based on Maximum Likelihood analysis using the Internal

Transcribed Spacers data (ITS). Bootstrap values (based on 1000 replicates) are presented

above the branches. Only Bootstrap values higher than 50% are shown.

Section Zingiber

Zingiber ultralimitale, a new species from Sulawesi

195

Lithophytic herb. Rhizome branching profusely, 5-8 mm across, brown externally,

yellowish internally, tasting like Zingiber officinale but very bitter; roots with irregular

ellipsoid tubers to 2.5 x 1.5 cm, white in cross-section, tasting like potato. Leafy shoots

in a loose clump of up to five pseudostems, 2—4 cm apart, to 40-75 cm tall, with 9—12(—

20) leaves. Base of leafy shoot to 9 mm diam., white to yellowish brown. Sheaths light

green to mid-green, ± finely hairy especially near margin and apex. Ligule to 8 mm

long, membranous, emarginate at least for half its length, with scattered, appressed

hairs, pale green. Petiole 2-9 mm long, channelled above, appressed villose abaxially,

especially in lower half. Lamina ovate to narrowly ovate, 9-18 x 3.5-5.5 cm, slightly

plicate, yellowish green and glabrous above, pale green and sparsely appressed villose

beneath especially near midrib, base rounded when fresh, cuneate to truncate when

dried, apex long acuminate. Inflorescence erect, to 16 cm long, arising close to the base

of leafy shoot, with 9-12 flowers, one flower open at a time in late morning. Peduncle

3-6 cm long with 4 peduncular bracts, upper one to 4.5 cm long, being the longest,

with scattered fine hairs. Spike to 10.5 cm long. Bracts laxly arranged at an angle

to the rachis, c. 5 mm apart exposing the mid-green rachis, each subtending a single

flower, lowermost bract c. 4.5 x 1.7 cm, translucent pale green, slightly darker towards

apex, with visible cross-veins and hyaline margins, apex of lower bracts acuminate,

Fig. 2. Pollen of Zingiber ultralimitale (acc. no 20092015 A). SEM made by Zou Pu.

196

Card. Bull. Singapore 69(2) 2017

Fig. 3. Zingiber ultralimitale Ardiyani & A.D.Poulsen. A. Habit on limestone boulders at

Bantimurung. B. Rhizome, including roots with tubers. C. Leafy shoot and inflorescence. D.

Sheath, ligule, petiole and base of lamina. E. Spike with single flower, front view. F. Spike with

single flower, lateral view. G. Bract. H. Bracteole and flower. I. Calyx. J. Ovary and corolla

tube. K. Dorsal corolla lobe, ventral view. L. Labellum and lateral corolla lobes, ventral view.

M. Corolla tube, stamen and stigma. N. Ovary and epigynous glands. A from Poulsen et al.

2767 ; B from Poulsen & Yeats 2989 ; C-N from Poulsen & Yeats 2984. (Photos: A.D. Poulsen)

Zingiber ultralimitale, a new species from Sulawesi

197

upper bracts with apex recurved, subulate appendages 6-8 mm long (reminiscent of

a reduced lamina), finely ciliate especially near margin and apex, glabrous inside.

Bracteoles ± as long as bracts, longer than calyx, to 4.6 x 1.3 cm (when flattened), split

to base, margins supervolute, translucent cream at base becoming green towards apex,

with scattered hairs near margin and along veins, glabrous inside. Calyx c. 2.6 cm

long, tubular with one incision in upper half, membranous, translucent cream, minutely

three-toothed at the apex, pubescent at apex. Corolla tube c. 3.5 cm long, white at base,

becoming pale yellow apically; dorsal corolla lobe narrowly ovate, slightly hollow, c.

3.4 x 1.2 cm, pale orange-yellow; lateral corolla lobes narrowly ovate, c. 2.5 x 0.7

cm, adnate c. 3 mm to base of labellum, pale orange-yellow. Labellum obovate, c.

3.5 x 3-3.5 cm, central lobe c. 3.5 x 2-2.5 cm, bifid, incision c. 8 mm, the two halves

slightly divergent and recurved, glabrous throughout, pale orange-yellow. Lateral

staminodes (side lobes) petaloid, c. 2 x 0.8 cm, paler than labellum, and free from

it. Stamen c. 3 cm long, filament distinct, c. 0.3 cm long, pale orange-yellow, anther

c. 1.5 cm, crest 1.3-1.6 cm long (when stretched), wrapped around style, incurved,

pale orange. Anther thecae open along their entire length, ± white. Style pale yellow.

Stigma tubular with a downwards-facing, ciliate ostiole which holds a droplet, held

by anther crest; epigynous glands 2, subulate, 4 mm long, pale yellow. Ovary c. 4 x 3

mm, trilocular, sometimes incompletely so, with axile placentation, ovules c. 25-30,

pale yellow-green. Fruit not seen.

Distribution. Only known from the type locality in South Sulawesi.

Etymology. The specific epithet ultralimitale means ‘on the other side of the border’,

referring to the occurrence of this species east of Wallace’s Line.

Ecology and habitat. Limestone cliffs and boulders in forest, lowlands at c. 300 m.

During the first year of cultivation in Edinburgh, it was discovered that the species has

a dormancy period during which it survives entirely underground.

Phenology. Flowering in May-June in cultivation. Our field assistant, Firdaus, checked

the populations at the type locality in June 2016 and found no leafy shoots. From this

we infer that the onset of the dry season had already resulted in the plant becoming

dormant.

Conservation status. Vulnerable VU Blab(iii) (IUCN, 2001). Abundant at one locality

along the road inside a national park but the extent of occurrence is estimated as

<20,000 km 2 . It is known from <10 locations and there is a decline in extent and

quality of the limestone habitats in Sulawesi.

Additional specimens examined. INDONESIA: South Sulawesi: Bantimurung National Park,

along Jalan Poros Maros-Soppeng, 5°2'32.4"S 119°44'9.4"E, 340 m, 23 January 2009, A.D.

Poulsen et al. 2767 (BO, E); Bantimurung National Park, along Jalan Poros Maros-Soppeng,

5°2'32.4"S 119°44'9.4"E, 340 m, 23 January 2009, living collection of Poulsen, A.D., Marlina

198

Card. Bull. Singapore 69(2) 2017

Ardiyani & Firdaus 2767 cultivated as RBGE 20091015*A, vouchered 12 June 2013 as

Newman, M.F. 2551, flowering (BO, E); ibidem, cultivated as 20091017*A, vouchered 28 May

2015 as Poulsen & Yeats 2984 , flowering (E); ibidem, cultivated as 20091017*A, vouchered 5

Feb 2016 as Poulsen & Yeats 2989, dormant (E); ibidem, cultivated as 20091017*A, vouchered

9 June 2016 as Yeats 16, flowering (E, SING); ibidem, cultivated as 20091019*A, vouchered

21 June 2016 as Yeats 17, flowering (E, CEB).

Notes. The rhizome branches and forms large clumps in cultivation, which explains

why it was almost impossible to remove it from the cracks in the limestone boulders

of its natural habitat. The leafy shoots collected in the field were much longer than

those of cultivated plants (75 vs. 40 cm) and had up to 20 leaves per shoot (vs. up to

12) which were narrower. In 2015, only one of the plants in the glasshouses produced

an inflorescence, which contained a total of 12 flowers usually opening late morning.

Only one flower opened per day but sometimes there would be a day without flowers

in between.

ACKNOWLEDGEMENTS. The Davis Expedition Fund, Augustinus, and Blaxall Valentine

Awards sponsored the expedition in Indonesia in 2009 during which the plant was discovered.

The Sibbald Trust funded A.D. Poulsen as a Davis Research Fellow. The State Ministry of

Research and Technology (RISTEK) issued the necessary permit. We are indebted to Helen

Yeats, RBGE for keeping the plants alive, thereby enabling us to study the flowers and describe

the species, and for making additional collections for herbaria. The Royal Botanic Garden

Edinburgh (RBGE) is supported by the Scottish Government’s Rural and Environmental

Science and Analytical Services Division. We thank J.F. Veldkamp and Philip Oswald for help

in Latinising the epithet, Zou Pu for letting us include her pollen photos, Firdaus for travelling

to the type locality to check the phenology, and Ida Theilade, Pramote Triboun, Bai Lin, John

Mood and Jana Leong-Skornickova for useful discussions. We also thank Susila and Hani

Kharismantari for helping us in the Molecular Lab, Herbarium Bogoriense, Research Center

for Biology, LIPI. Last but not least, the first author would like to thank the Sibbald Trust of the

Royal Botanic Garden Edinburgh which allowed her to examine the herbarium specimens of

Zingiber at E and K after the 10th Flora Malesiana Symposium.

References

Ardiyani, M. (2015). A new species of Zingiber (Zingiberaceae) from Enggano Island,

Indonesia. Reinwardtia 14(2): 307-310.

Baker, J.G. (1894). Scitamineae. In: Hooker, J.D. (ed.) Flora of British India, vol. 6, pp. 198—

264. London: L Reeve & Co.

IUCN (2001). IUCN Red List Categories and Criteria'. Version 3.1. Gland, Switzerland and

Cambridge, UK: IUCN.

Kress, W.J. & Erickson, D.L. (2007). A two-locus global DNA barcode for land plants: The

coding rbcL gene complements the non-coding trnW-psbA spacer region. PLoS ONE

2: e508.

Kress, W.J., Prince, L.M. & Williams, K.J. (2002). The phylogeny and a new classification of

the gingers (Zingiberaceae): evidence from molecular data. Am. J. Bot. 89: 1682-1696.

Zingiber ultralimitale, a new species from Sulawesi

199

Schumann, K. (1904). Zingiberaceae. In: Engler, H.G.A. (ed.) Das Pflanzenreich, IV(46), fas.

20, pp. 1-458. Leipzig: W. Engelmann.

Tamura, K., Stecher, G., Peterson, D., Filipski, A. & Kumar, S. (2013). MEGA6: Molecular

Evolutionary Genetics Analysis version 6.0. Molec. Biol. Evol. 30: 2725-2729.

Theerakulpisut, P., Triboun, R, Mahakham, W., Maensiri, D., Khampila, J. & Chantaranothai,

P. (2012). Phylogeny of the genus Zingiber (Zingiberaceae) based on nuclear ITS

sequence data. Kew Bull. 67: 389-395.

Theilade, I., Maersk-Mpller, M.L., Theilade, J. & Larsen, K. (1993). Pollen morphology and

structure of Zingiber (Zingiberaceae). Grana 32: 338-342.

Zingiberaceae Resource Centre (2017). Royal Botanic Garden Edinburgh, http://padme.rbge.

org.uk/ZRC/welcome. Accessed 3 Jul. 2017.

Gardens' Bulletin Singapore 69(2): 201-210. 2017

doi: 10.26492/gbs69(2).2017-05

201

Syzygium jiewhoei (Myrtaceae), a new endemic tree

from Western New Guinea, Indonesia

G.G. Hambali 12 , S. Sunarti 3 & Y.W. Low 4 - 5 - 6

‘Taman Buah Mekarsari, Cileungsi, Bogor, Indonesia

2 Yatazawa R & D Gardens, Bogor, Indonesia

gregham2010 @ yahoo.com

3 Herbarium Bogoriense, Research Center for Biology, Indonesian Institute of Science,

Jl. Raya Jakarta-Bogor KM 46, Cibinong, 16911, Bogor, Indonesia

4 Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, 259569 Singapore

Comparative Plant and Fungal Biology, Royal Botanic Gardens Kew,

Richmond, Surrey TW9 3AE, UK

6 School of Biological Sciences, University of Aberdeen,

AB24 3UU Aberdeen, UK

ABSTRACT. Syzygium jiewhoei Hambali, Sunarti & Y.W.Low, a new species from Western

New Guinea, Indonesia, is described and illustrated. It is closely related to Syzygium

recurvovenosum (Lauterb.) Diels but differs in a range of vegetative and reproductive

morphological characteristics.

Keywords. East Malesia, Papua Province, Sahul shelf, Syzygium recurvovenosum

Introduction

New Guinea, the largest tropical island in the world, is located in the Malesian region

with an area of approximately 800,000 km 2 . Politically, it is divided into two roughly

equal halves: to the east is Papua New Guinea and to the west is Indonesian New

Guinea, comprising the provinces of Papua and West Papua. Biogeographically, New

Guinea is part of the Sahul shelf, that also includes Australia and Tasmania, and these

regions share similar floristic elements that are distinct from those on the Sunda shelf

(Gressitt, 1982; Pieters, 1982). Hence, the phytogeography of the Malesian region

has been a focus of research by tropical botanists (Van Steenis, 1950; Van Welzen et

al., 2011; Crayn et al., 2015). A comprehensive Flora of New Guinea is still lacking.

However, efforts by various botanical institutions have led to checklists, guides and

monographs of selected plant groups, such as the orchids (Schuiteman & de Vogel,

2001, 2002, 2005, 2006, 2008; Schuiteman et al., 2010), palms (Heatubun, 2002;

Baker & Dransfield, 2006; Heatubun et al., 2012), and alpine plants (Van Royen,

1979a, 1979b, 1982, 1983; Johns et al., 2006), etc.

Syzygium Gaertn. is one of the largest genera in the world, with about 1200-

1800 species occurring principally in the Old World (Frodin, 2004; Parnell et al.,

2007; Syzygium Working Group, 2016). A recent analysis reveals that Syzygium is

202

Card. Bull. Singapore 69(2) 2017

the genus with the largest number of tree species, about 1069 species in all (Beech et

al., 2017). Syzygium diversity in New Guinea is expected to be extremely rich, with

species occurring from the lowlands to the highlands (Craven, 2006), but identification

of Syzygium species has always been difficult due to a lack of good morphological

diagnostic characters, resulting in an accumulation of unidentified materials in many

herbaria (Hartley & Perry, 1973; Craven, 2006; Craven & Biffin, 2010).

Among the earliest botanists to examine New Guinean Syzygium was Carl

Lauterbach (Van Steenis-Kruseman, 1950). He described many new species, including

some under segregate genera that are now considered synonyms of Syzygium , such as

Aphanomyrtus Miq. and Jambosa Adans. (Lauterbach, 1910, 1912). Later, Friedrich

Diels (see Van Steenis-Kruseman, 1950) described many more Syzygium species for

New Guinea, some of them under Jambosa (Diels, 1922; Diels et al., 1929). Merrill &

Perry (1939a, 1942a, 1942b) meticulously examined the enormous volume of material

of New Guinean Syzygium gathered during the Archbold Expeditions. It was Merrill

& Perry (1939b), in a treatment of Syzygium in Borneo, who synonymised Jambosa ,

and also a few other segregate genera in the Syzygium alliance, under Syzygium. Later,

Hartley & Perry (1973) attempted to revise the by-then much larger Syzygium diversity

of Papuasia. They recognised 138 species and also listed 69 species of unknown

status or which were to be excluded from Papuasia, citing lack of materials for close

examination while preparing the account. A recent enumeration in the World Checklist

of Myrtaceae recorded 195 species of Syzygium for New Guinea (Govaerts et al.,

2017).

In 1993, while assisting Freeport to establish a plant nursery at Timika under

the Incubator Project, the first author stumbled upon several attractive juvenile plants

of an unidentified Syzygium species with exceptionally long pendulous leaves in the

forested area of Kuala Kencana. Ten years later, the first author visited Timika again

and this provided an opportunity to procure seeds and seedlings of the peculiar Kuala

Kencana Syzygium species for cultivation. From these materials, one of the plants was

successfully established and is growing vigorously in the garden of Mr Tan Jiew Hoe

in Singapore. The tree in Singapore eventually produced flowers and fruits, providing

us with sufficient materials to examine for this study. The unidentified Kuala Kencana

Syzygium is closely related to Syzygium recurvovenosum (Lauterb.) Diels but is distinct

based on the suite of morphological characters listed in Table 1. It is concluded that

the attractive Kuala Kencana Syzygium is an undescribed novelty from Western New

Guinea and is named here as Syzygium jiewhoei Hambali, Sunarti & Y.W.Low.

Material and methods

A review of all New Guinean Syzygium species was conducted, based on herbarium

specimens preserved in BO, K and SING (herbarium acronyms follow Thiers, 2017,

continuously updated). Conventional methods employed in herbarium taxonomy were

applied in this study. All measurements were taken from dried herbarium specimens.

Photographic documentation is of the living specimen growing in Mr Tan Jiew Hoe’s

Syzygium jiewhoei, a new species from Western New Guinea

203

Table 1 . Comparison of morphological characteristics between Syzygium jiewhoei Hambali,

Sunarti & Y.W.Low and S. recurvovenosum (Lauterb.) Diels.

Syzygium jiewhoei

Syzygium recurvovenosum

Young twigs

Prominently quadrangular

and winged

Prominently quadrangular

Lamina

Slightly bullate

Unknown

Secondary veins

90-100 pairs

Up to 55 pairs

Inflorescence

c. 14-16 cm long

Up to 9 cm long

Peduncle

c. 13-15 mm wide

Up to 3.5 mm wide

Style

c. 8-18 mm long

c. 4 mm long

Fruits

Ovoid to broad-ellipsoid, 4-6 cm

long, maturing salmon-pink

Pyriform, 2.3 cm long,

red (immature)

garden in Singapore, from which the material for the herbarium was taken. Type

materials of Syzygium species at BO, K and SING were examined, including type

images of all Malesian Syzygium species available on JSTOR® Global Plants website

(http://plants.jstor.org). Provisional conservation assessments are made using the

methodology proposed by IUCN (2012).

Taxonomy

Syzygium jiewhoei Hambali, Sunarti & Y.W.Low, sp. nov.

Similar to Syzygium recurvovenosum (Lauterb.) Diels but differs in having 90-100

pairs of secondary veins (vs up to 55 pairs of secondary veins in S. recurvovenosum :),

14-16 cm long inflorescences with 13-15 mm wide peduncles (vs up to 9 cm long and

c. 3.5 mm wide in S. recurvovenoswri), and 8-18 mm long styles (vs 4 mm long in S.

recurvovenosum). - TYPE: Native to Indonesia, Western New Guinea, Papua, Timika,

Kuala Kencana, living collection cultivated in Mr Tan Jiew Hoe’s garden in Singapore,

vouchered on 3 July 2016 as Hambali, G.G. s.n. (holotype BO; isotype SING). (Fig.

1 , 2 )

Tree up to 9 m tall, diameter at breast height c. 20 cm, without buttresses. Bark papery,

peeling, greyish brown. Branchlets glabrescent, green, prominently quadrangular

and winged when young, becoming terete on older branches. Leaves purple when

young, opposite and decussate on erect shoots, becoming secondarily distichous on

lateral branches; blades narrowly elliptic to broadly linear, 60-105 x 14-25 cm, with

hardly visible minute oil glands; margin entire; base cordate, apex long-acuminate;

midrib and secondary veins on lower surface prominent, midrib flat and secondary

veins sunken on upper surface; secondary veins 90-100 pairs, rather abruptly curving

out from the midrib, intramarginal vein c. 2 mm from the leaf margin; glabrous on

204

Card. Bull. Singapore 69(2) 2017

Fig. 1 . Syzygium jiewhoei Hambali, Sunarti & Y.W.Low. A. Young leaves. B. Cauliflorous habit

with many inflorescences at various stages. C. Close-up of inflorescences showing flowers

at anthesis. D. Close-up of infructescence. All from type Hambali, G.G. s.n. (Photos: G.G.

Hambali)

Syzygium jiewhoei, a new species from Western New Guinea

205

Fig. 2. Syzygium jiewhoei Hambali, Sunarti & Y.W.Low. A. Young leafy branch showing

winged and angular stem. B. Close-up of cordate leaf base and numerous almost parallel

secondary veins arranged very closely together. C. An immature inflorescence. D. Close-up

of a mature flower showing the numerous stamens. E. Close-up of a longitudinal section of

a mature flower depicting the arrangement of stamens along the rim of the hypanthium. F. A

mature fruit. G. Longitudinal section of a mature fruit showing echinate testa intrusions into the

cotyledons. Drawn by Subari from holotype, Hambali, G.G. s.n.( BO).

206

Card. Bull. Singapore 69(2) 2017

both sides; upper surface green, drying dark brown, lower surface pale green, drying

brown; petiole c. 7-10 mm long, 6-13 mm wide, leaves appearing subsessile due

to length of blades. Inflorescences developing on the stem (cauliflorous), compound

cymes 14-16 cm long with up to c. 250 flowers; peduncle dark brown, 3.5-6.5 cm

long, 13-15 mm wide, branching to 4 orders, bearing flowers usually in triads at their

tips; bracteoles rudimentary, caducous. Flowers bisexual, white, 18-24 mm long,

hypanthium turbinate, 12-14 mm long, 3—4 mm wide at the top; sepals rudimentary;

petals roundish, 3^1 mm across, pale green, caducous at anthesis. Stamens numerous,

up to 76; filaments 6-10 mm, white; anthers c. 0.3 mm long, dirty white. Pistil

persistent, green; style 8-18 mm long; ovary inferior. Fruits 4-6 x 4.5-5 cm, ovoid

to broad-ellipsoid, slightly grooved longitudinally, salmon-pink and sour when fully

ripe. Seeds 3.5 x 2.8 cm.

Etymology. We are pleased to name this handsome tree, with foliage very much

resembling that of Anthurium veitchii Mast. (Araceae), after Mr Tan Jiew Hoe, a

benefactor of science who has a great interest in natural history, particularly in the

fields of botany and horticulture (see Kurzweil & Lwin, 2014; Kiew et al., 2015;

Leong-Skomickova & Newman, 2015; Lamb & Rodda, 2016).

Distribution and habitat. Syzygium jiewhoei is so far known only from the lowland

forests around the vicinity of Timika, Papua Province, Indonesian New Guinea.

However, the species has now been introduced for cultivation as an ornamental tree in

Bogor (Java, Indonesia) and Singapore (Fig. 3).

Provisional IUCN conservation assessment. Data Deficient DD, following the

guidelines in IUCN (2012). Syzygium jiewhoei is known only from Kuala Kencana,

Timika, Papua Province, Indonesian New Guinea. As the distribution of this species

is inadequately known, we propose a status of Data Deficient (DD). As for all species

given this status, reassessment wifi be required as more data is gathered to document

the flora of New Guinea. One current initiative is the Tropical Important Plant Areas-

Indonesian New Guinea programme (TIPAs Indonesian New Guinea), led by the Royal

Botanic Gardens, Kew in collaboration with Universitas Papua (UNIPA), Manokwari,

to identify important areas for plant conservation with an emphasis on the Bird’s Head

Peninsula, West Papua Province based on a set of criteria proposed by a team of plant

conservationists (Darbyshire et al., 2017; Royal Botanic Gardens Kew, 2017).

Notes. Syzygium jiewhoei is a majestic free-flowering tree with a somewhat broad

conical crown, which makes it a highly desirable ornamental tree. At maturity it

produces showy inflorescences on its trunk. Flowers of Syzygium jiewhoei produce

copious nectar for up to three days after anthesis, thereby attracting large numbers of

the Asian honey bee (Apis cerana ) and stingless bee ( Trigona laeviceps ). The mature

salmon-pink fruits of Syzygium jiewhoei range from ovoid to broad-ellipsoid and

somewhat oblong.

Syzygium jiewhoei, a new species from Western New Guinea

207

Fig. 3. Distribution of Syzygium jiewhoei Hambali, Sunarti & Y.W.Low in New Guinea.

It’s worth noting that there is a solitary tree of Syzygium jiewhoei growing beside

the road at the back of PT Freeport’s Environmental Department building in Kuala

Kencana. It was previously erroneously identified as Syzygium versteegii (Lauterb.)

Merr. & L.M.Perry. Syzygium versteegii differs from Syzygium jiewhoei in having

terete branchlets and oblong leaves with only 15 pairs of secondary veins.

ACKNOWLEDGEMENTS. We would like to express our gratitude to Mr Tan Jiew Hoe for

permission to collect flowering and fruiting materials of the new species from his garden for this

study. GGH would like to thank Ms Siti Hutami Endang Adiningsih, Director of Taman Buah

Mekarsari, for permission to perform this taxonomic research and write-up the manuscript

and to Dr Wong Khoon Meng for his kindness in making some corrections and suggestions

in the early stage of the manuscript preparation. SS is especially grateful to Dr Joeni Setijo

Rahajoe (Keeper of BO) and Dr Rugayah for their continuous support and encouragement

with the on-going Syzygium and Myrtaceae research conducted in BO. YWL is grateful to

the National Parks Board, Singapore, through Singapore Botanic Gardens, for his research

opportunities. YWL is also grateful to Prof. Dr David Burslem (University of Aberdeen), Dr

David Middleton (Singapore Botanic Gardens) and Dr Eve Lucas (Royal Botanic Gardens

Kew) for their guidance, support and encouragement with his on-going PhD research on

Southeast Asian Syzygium , with a special focus on New Guinea. Finally, we are extremely

grateful to the Keepers of BO, K and SING for permission to access materials under their care.

Mr Subari (BO) prepared the line drawing and Ms Serena Lee (SING) helped with the map

used here in this study.

208

Card. Bull. Singapore 69(2) 2017

References

Baker, W.J. & Dransfield, J. (2006). Field Guide to the Palms of New Guinea. London: Royal

Botanic Gardens, Kew.

Beech, E., Rivers, M., Oldfield, S. & Smith, PR (2017). GlobalTreeSearch: The first complete

global database of tree species and country distributions. J. Sustainable For. 36(5):

454-489.

Craven, L.A. (2006). Myrtaceae of Papua. In: Marshall, A.J. & Beehler, B.M. (eds) The Ecology

of Papua, vol. 1, pp. 429-434. Hong Kong: Periplus Editions (HK) Ltd.

Craven, L.A. & Biffin, E. (2010). An infrageneric classification of Syzygium (Myrtaceae).

Blumea 55: 94-99.

Crayn, D.M., Costion, C. & Harrington, M.G. (2015). The Sahul-Sunda floristic exchange:

dated molecular phylogenies document Cenozoic intercontinental dispersal dynamics.

J. Biogeogr. 42(1): 11-24.

Darbyshire, I., Anderson, S., Asatryan, A., Byfield, A., Cheek, M., Clubbe, C., Ghrabi, Z.,

Harris, T., Heatubun, C.D., Kalema, J., Magassouba, S., McCarthy, B., Milliken, W.,

de Montmollin, B., Lughadha, E.N., Onana, J.-M., Saidou, D., Sarbu, A., Shrestha, K.

& Radford, E.A. (2017). Important Plant Areas: revised selection criteria for a global

approach to plant conservation. Biodivers. Conserv. 26(8): 1767-1800.

Diels, L. (1922). Die Myrtaceen von Papuasien. Bot. Jahrb. Syst. 57: 356-426.

Diels, L., Markgraf, R., Mansfeld, R., Schmidt, O.C., Gilg, E. & Markgraf, T. (1929).

Descriptions of new species collected in British Papua by L.J. Brass. J. Arnold Arbor.

10(2): 75-84.

Frodin, D.G. (2004). History and concepts of big plant genera. Taxon 53(3): 753-776.

Govaerts, R., Sobral, M., Ashton, P.S., Barrie, F., Holst, B.K., Landrum, L.L., Matsumoto, K.,

Fernanda Mazine, F., Nic Lughadha, E., Proenca, C., Soares-Silva, L.H., Wilson, P.G.

& Lucas, E. (2017). World Checklist of Myrtaceae. Royal Botanic Gardens, Kew. http://

apps.kew.org/wcsp/. Accessed 29 Aug. 2017.

Gressitt, J.L. (1982). General introduction. In: Gressitt, J.L. (ed.) Biogeography and Ecology of

New Guinea, vol. 1, pp. 3-13. The Hague: Dr W. Junk Publishers.

Hartley, T.G. & Perry, L.M. (1973). A provisional key and enumeration of species of Syzygium

(Myrtaceae) from Papuasia. J. Arnold Arbor 54(2): 160-227.

Heatubun, C.D. (2002). A monograph of Sommieria (Arecaceae). Kew Bull. 57: 599-611.

Heatubun, C.D., Dransfield, J., Flynn, T., Tjitrosoedirdjo, S.S., Mogea, J.P & Baker, W.J.

(2012). A monograph of the betel nut palms (, Areca: Arecaceae) of East Malesia. Bot. J.

Linn. Soc. 168(2): 147-173.

IUCN (2012). IUCN Red List Categories and Criteria: Version 3.1, 2 nd ed. Gland, Switzerland

and Cambridge, UK: IUCN.

Johns, R.J., Edwards, P.J., Utteridge, T.M.A. & Hopkins, H.C.F. (2006). A Guide to the Alpine

and Subalpine Flora of Mount Jaya. London: Royal Botanic Gardens, Kew.

Kiew, R., Sang, J., Repin, R. & Joffre, A.A. (2015). A Guide to Begonias of Borneo. Kota

Kinabalu: Natural History Publications (Borneo).

Kurzweil, H. & Lwin, S. (2014). A Guide to Orchids of Myanmar Kota Kinabalu: Natural

History Publications (Borneo).

Lamb, A. & Rodda, M. (2016). A Guide to Hoyas of Borneo. Kota Kinabalu: Natural History

Publications (Borneo).

Lauterbach, C. (1910). Myrtaceae. In: Lorentz, H.A. (ed.) L’Expedition scientifique

Neerlandaise a la Nouvelle-Guinee. Nova Guinea 8(1): 319-323.

Syzygium jiewhoei, a new species from Western New Guinea

209

Lauterbach, C. (1912). Myrtaceae. In: Lorentz, H.A. (ed.) Uitkomsten der Nederlandsche

Nieuw-Guinea-Expeditie. Nova Guinea 8(2): 849-855.

V _

Leong-Skornickova, J. & Newman, M.F. (2015). Gingers of Cambodia, Laos and Vietnam.

Singapore: National Parks Board, Singapore Botanic Gardens.

Merrill, E.D. & Perry, L.M. (1939a). Plantae Papuanae Archboldianae, I. J. Arnold Arbor.

20(3): 324-345.

Merrill, E.D. & Perry, L.M. (1939b). The Myrtaceous genus Syzygium in Borneo. Mem. Amer.

Acad. Arts 18(3): 135-202.

Merrill, E.D. & Perry, L.M. (1942a). Plantae Papuanae Archboldianae, IX. J. Arnold Arbor.

23(2): 233-265.

Merrill, E.D. & Perry, L.M. (1942b). Plantae Papuanae Archboldianae, IX. J. Arnold Arbor.

23(3): 267-297.

Parnell, J.A.N., Craven, L.A. & Biffin, E. (2007). Matters of scale: Dealing with one of the largest

genera of angiosperms. In: Hodkinson, T.R. & Parnell, J.A.N. (eds) Reconstructing the

Tree of Life: Taxonomy and Systematics of Species Rich Taxa, pp. 251-273. Boca Raton,

Florida: CRC Press.

Pieters, PE. (1982). Geology of New Guinea. In: Gressitt, J.L. (ed.) Biogeography and Ecology

of New Guinea, vol. 1, Monographiae Biologicae, vol. 42, pp. 15-38. The Hague: Dr

W. Junk Publishers.

Royal Botanic Gardens Kew (2017). Tropical Important Plant Are as-Indonesian New

Guinea (TIPAs-Indonesian New Guinea). https://www.kew.org/science/projects/tipas-

indonesian-new-guinea. Accessed 21 Sep. 2017.

Schuiteman, A. & de Vogel, E.F. (2001). Flora Malesiana: Orchids of New Guinea, vol. 1:

Illustrated Checklist and Genera. CD-ROM. Expert-Center for Taxonomic Identification.

Schuiteman, A. & de Vogel, E.F. (2002). Flora Malesiana: Orchids of New Guinea, vol. 2:

Dendrobium and Allied Genera. CD-ROM. Expert-Center for Taxonomic Identification.

Schuiteman, A. & de Vogel, E.F. (2005). Flora Malesiana: Orchids of New Guinea, vol. 3:

Genera Acanthephippium to Hymenorchis (excluding Dendrobiinae s.L). CD-ROM.

Expert-Center for Taxonomic Identification.

Schuiteman, A. & de Vogel, E.F. (2006). Flora Malesiana: Orchids of New Guinea, vol. 4:

Genera Kuhlhasseltia to Ophioglossella (excluding Dendrobiinae s.L). CD-ROM.

Expert-Center for Taxonomic Identification.

Schuiteman, A. & de Vogel, E.F. (2008). Flora Malesiana: Orchids of New Guinea, vol. 5:

Genera Pachystoma to Zeuxine (excluding Dendrobiinae s.L). CD-ROM. Expert-

Center for Taxonomic Identification.

Schuiteman, A., Vermuelen, J.J. & de Vogel, E.F. (2010). Flora Malesiana: Orchids of

New Guinea, vol. 6: Genus Bulbophyllum. CD-ROM. Expert-Center for Taxonomic

Identification.

Syzygium Working Group (2016). Syzygium (Myrtaceae): Monographing a taxonomic giant

via 22 coordinated regional revisions. PeerJ Preprints 4: el930vl.

Thiers, B. (2017, 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 24 Aug. 2017.

Van Royen, P. (1979a). The Alpine Flora of New Guinea, vol. 1, General Part. Vaduz: J. Cramer.

Van Royen, P. (1979b). The Alpine Flora of New Guinea, vol. 2, Taxonomic Part: Cupressaceae

to Poaceae. Vaduz: J. Cramer.

Van Royen, P. (1982). The Alpine Flora of New Guinea, vol. 3, Taxonomic Part: Winteraceae

to Polygonaceae. Vaduz: J. Cramer.

210

Card. Bull. Singapore 69(2) 2017

Van Royen, R (1983). The Alpine Flora of New Guinea, vol. 4, Taxonomic Part: Casuarinaceae

to Asteraceae. Vaduz: J. Cramer.

Van Steenis, C.G.G.J. (1950). The delimitation of Malaysia and its main plant geographical

divisions. In: Flora Malesiana, ser. I, vol. 1, pp. lxx-lxxv. Jakarta: Noordhoff-Kolff

N.V.

Van Steenis-Kmseman, M.J. (1950). Malaysian plant collectors and collections: being a

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

up to year 1950. In: van Steenis, C.G.G.J. (ed.) Flora Malesiana, ser. I, vol. 1, pp.

3-639. Jakarta: Noordhoff-Kolff N.V.

Van Welzen, PC., Parnell, J.A.N. & Slik, J.W.F. (2011). Wallace's Line and plant distributions:

two or three phytogeographical areas and where to group Java? Biol. J. Linn. Soc.

103(3): 531-545.

Gardens' Bulletin Singapore 69(2): 211-284. 2017

doi: 10.26492/gbs69(2).2017-06

211

A revision of Microchirita (Gesneriaceae) in Thailand

C. Puglisi 1 ’ 2 & D.J. Middleton 1

Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, 259569 Singapore

david_middleton@nparks.gov.sg

2 Royal Botanic Garden Edinburgh, 20A Inverleith Row,

Edinburgh, EH3 5LR, UK

ABSTRACT. Microchirita (C.B.Clarke) Yin Z.Wang (Gesneriaceae: Didymocarpoideae) in

Thailand is revised and 29 species are recognised, two of which have three varieties each.

Eight new species are described, Microchirita albocyanea C.Puglisi, Microchirita glandulosa

C.Puglisi, Microchirita hypocrateriformis C.Puglisi, Microchirita limbata C.Puglisi,

Microchirita luteola C.Puglisi, Microchirita tadphoensis C.Puglisi, Microchirita tetsanae

C.Puglisi, Microchirita thailandica C.Puglisi; three new varieties are described, Microchirita

involucrata var. gigantiflora C.Puglisi, Microchirita mollissima var. glabra C.Puglisi,

Microchirita mollissima var. glandulophylla C.Puglisi; and one name is combined at a new

rank, Microchirita involucrata var. capitis (Craib) C.Puglisi. Two lectotypifications are made,

one of which is a second step lectotypification. A key to all taxa is given, all taxa are described,

and many are illustrated.

Keywords. Chirita, Didymocarpoideae, Flora of Thailand, Gesneriads, new species,

taxonomy

Introduction

Generic delimitation in Asian Gesneriaceae has been the focus of much recent research

and has led to considerable change (Wei et al., 2010; Moller et al., 2011, 2014, 2016;

Puglisi et al., 2011, 2016; Wang et al., 2011; Weber et al., 2011a, 2011b; Middleton

& Moller, 2012; Middleton et al., 2014, 2015). This research, much of it based on

the results of DNA sequence data, has resulted in an overall reduction in the number

of genera but also the description of new genera to accommodate new species which

were found to belong to previously unknown lineages (Middleton & Triboun, 2012;

Middleton et al., 2015), and the splitting up of genera found to be polyphyletic (Weber

et al. 2011a; Moller et al., 2014). Although there are still many questions to address, a

relative degree of stability in the delimitation of Asian Gesneriaceae genera has been

reached.

The genus Chirita Buch.-Ham. is one of the genera that was split up as a

consequence of molecular phylogenetic research which found the genus to be

polyphyletic (Wang et al., 2011; Weber, 2011a). Chirita was characterised by the

presence of a “chiritoid” stigma (a two-lipped stigma with the upper lip reduced and the

lower bilobed) but was otherwise extremely morphologically diverse. The genera into

212

Card. Bull. Singapore 69(2) 2017

which it has been divided are Damrongia Kerr ex Craib, Henckelia Spreng. (including

Chirita itself), Liebigia Endl., Microchirita (C.B.Clarke) Yin Z.Wang and Primulina

Hance (Weber et al. 2011a). Each of these genera is much more morphologically

coherent than was Chirita. Primulina continues to grow in the number of species and

is in need of revision, Liebigia has been revised as a section of Chirita (Hilliard, 2004),

Damrongia has been revised for Thailand, its centre of diversity (Puglisi & Middleton,

2017), Henckelia is being investigated by a number of research groups (Moller, pers.

comm.; Sirimongkol, pers. comm.), and Microchirita is the subject of this paper.

Microchirita is found in India, Myanmar, southern China, Thailand, Vietnam,

Laos, Cambodia, Peninsular Malaysia, Sumatra, Java and Borneo, almost exclusively

in limestone habitats. As part of his revision of Chirita, Wood (1974) included 18

species in Chirita sect. Microchirita C.B.Clarke. Of these, Chirita elata Ridl. has

since been removed to the genus Codonoboea Ridl. (Rafidah et al., 2011) and eleven

additional species have been described (Punekar & Lakshminarasimhan, 2009;

Middleton & Triboun, 2013; Rafidah & Haron, 2013; Puglisi et al., 2016). This brings

the total to 28 species of which 22 have been recorded for Thailand (Wood, 1974;

Burtt, 2001; Middleton & Triboun, 2013; Puglisi et al., 2016), one mistakenly ( Chirita

caerulea R.Br. by Wood, 1974), leaving 21 species. This clearly makes Thailand the

centre of diversity of the genus with 3 A of the known species in the genus found there.

The lectotype of the genus is Microchirita hamosa (R.Br.) Yin Z.Wang.

The most characteristic morphological feature of many species of Microchirita

is the cristate inflorescence consisting of a single row of flower pairs (Fig. 1A). The

young flowers develop against the base of the lamina. Subsequently the pedicels

straighten to an upright position when the flowers reach maturity, and bend backwards

towards the main stem at the fruiting stage. The series of flower pairs appears as a

crest along the petiole, and often there is tissue melding together the peduncles at the

base. Wood (1974) interpreted this crest as a single inflorescence whose peduncle is

fused with the petiole. Weber (1975) instead supported the idea of a system of multiple

inflorescences generated by an enlarged meristem which is displaced from the axil onto

the petiole. No attempt was made to characterise the inflorescence in this study, and

for merely practical reasons we have chosen to refer to a “cristate inflorescence” when

applicable, without implying the acceptance of Wood’s theory over Weber’s. Some

species (e.g. Microchirita involucrata (Craib) Yin Z.Wang) have inflorescences which

do not appear cristate. They consist of one or few well-developed peduncles arising

from the axillary end of the petiole, each topped by paired bracts (free or fused), and

culminating in a subumbellate cluster of pedicels (Fig. IB).

The species of Microchirita are annual or short-lived. The stems are often fleshy

and green, tinged with purple-brown. Another characteristic feature of Microchirita is

the leaf arrangement: the basal leaf, the macrocotyledon, is single, although sometimes

the paired leaf, the much reduced microcotyledon, persists. The subsequent leaves are

opposite. This general structure may not be apparent in Microchirita mollissima (Ridl.)

A.Weber & D.J.Middleton and most Malaysian species (Rafidah, 2017). In these the

internodes are shorter and the leaves more crowded, thereby obscuring the phyllotaxis.

Also, the inflorescences develop at the axillary end and are less dense. Some species

A revision of Microchirita in Thailand

213

Fig. 1. Inflorescence types. A. Cristate inflorescence of Microchirita bimaculata (D.Wood)

A.Weber & D.J.Middleton. From Middleton, D.J. et al 4514. B. Bracteate inflorescence of

Microchirita rupestris (Ridl.) A.Weber & Rafidah. From Puglisi, C. et al. CP409. (Photos: A,

D.J. Middleton; B, P. Karaket)

214

Card. Bull. Singapore 69(2) 2017

may flower when only the macrocotyledon has developed and can be mistakenly

interpreted as unifoliate, a common state in Microchirita hamosa for example.

Generally this is very variable, even within a single population, with a mixture of

plants flowering at different sizes and stages of development. The macrocotyledon can

be much larger than the paired leaves, leading to broad ranges of leaf size and number

of secondary veins.

Much variation can be observed in the corolla of Microchirita. The tube is

generally narrow at the base, often bent downwards, and widens more or less abruptly,

into a campanulate, funnel-shaped or tubular upper corolla. The colour of the corolla

varies substantially from pure white to a lilac-purple-blue palette, or to a pale yellow-

yellow-dark orange range. The colour is rarely uniform throughout the corolla. Some

species (e.g. Microchirita albocyanea C.Puglisi) have lobes of a different shade from

the tube; most species (e.g. M. tubulosa (Craib) A.Weber & D.J.Middleton) have a

ventral yellow line running from the base of the ventral lobe to the throat; many species

(e.g. M. bimaculata (D.Wood) A.Weber & D.J.Middleton) have lines or spots either at

the base of the lateral lobes or at the base of the upper lip; finally, in few taxa (e.g. M.

rupestris (Ridl.) A.Weber & Rafidah), the venation of the petals is visible in the form

of fine purple lines running along the lobes and anastomosing terminally. The colour

and patterning of the corolla are taxonomically significant. Some species appear to

have a purple/blue form and a white equivalent (e.g. M. hypocrateriformis C.Puglisi).

Where this variation was not substantiated by allopatry or further morphological

differentiation, no taxonomic distinction is made.

Anthers of Microchirita species can be free, coherent face-to-face, or apically

joined by a ligament. Furthermore, anthers can be glabrous or bear an indumentum,

especially dorsally at the filament insertion. The indumentum is usually of long,

somewhat woolly, eglandular hairs, but in some species additional, shorter and straight

hairs are seen.

In this account, Microchirita in Thailand is revised and 29 species, two of which

have three varieties each, are recognised. Of these, eight species and three varieties are

newly described.

Material and methods

This revision is based on a study of the specimens from the herbaria A, AAU, ABD,

BK, BKF, BM, CMU, CMUB, E, K, KEP, L, P, PSU, QBG, SING, US (Thiers et

al., 2017, continuously updated), and Mahidol University, Kanchanaburi Campus

(MUKA). All specimens have been seen unless otherwise noted. Measurements of

all but the floral parts were taken from dry specimens. Flower measurements are from

rehydrated or pickled flowers. When no additional material of the recently published

species (Middleton & Triboun, 2013; Puglisi et al., 2016) was obtained, we give

measurements from the original descriptions. Measurements of the flowers were taken

with a microruler and should be considered accurate to 0.05 mm.

Conservation assessments follow the guidelines laid out in IUCN (2012).

A revision of Microchirita in Thailand

215

The authors have seen more than half of the species in this revision in the

field. Field characters have been instrumental in gaining a better understanding of

the species to aid in species delimitation. In particular it becomes quite obvious in the

field that plants flowering as early as the macrocotyledon stage are completely mixed

into populations of much larger plants and that previous taxonomic distinctions based

on plant size are untenable. We have not seen living plants of some of the species

in the Northeast of Thailand but have been greatly aided by photographs taken by

collectors. Many species of Microchirita are rather similar when pressed and dried

and we urge collectors to ensure good collection notes, especially in corolla colour and

colour variation characters, and to take photographs of the living plants. The species

are presented in alphabetical order.

Taxonomic treatment

Microchirita (C.B.Clarke) Yin Z.Wang, J. Syst. Evol. 49: 59 (2011). - Chirita sect.

Microchirita C.B.Clarke in Candolle & Candolle, Monogr. Phan. 5: 127 (1883). -

Roettlera sect. Microchirita (C.B.Clarke) Fritsch in Engler & Prantl, Nat. Pflanzenfam.

IV/3b: 148 (1895 ). - Didymocarpus sect. Microchirita (C.B.Clarke) Chun, Sunyatsenia

6: 290 (1946). - TYPE: Microchirita hamosa (R.Br.) Yin Z.Wang, lectotype designated

by Burtt (1954: 196).

Annual or semiperennial herbs, often with succulent stems. First leaf (macrocotyledon)

often persistent and single (not paired), second cotyledon rudimentary or suppressed;

all other leaves in opposite pairs, sessile or petiolate. Inflorescences axillary or

epiphyllous, often crested (see Introduction above); peduncles normally developed

in axillary inflorescences, reduced and sometimes fused with each other in crested

inflorescences; bracts present in axillary inflorescences, divided or somewhat fused;

flowers 5-merous. Calyx lobes divided almost to base or short tube present. Corolla of

a narrow lower tube, a wider upper tube and a bilabiate limb, upper lip 2-lobed, lower

lip 3-lobed, variable in shape and colour. Stamens 2, filaments straight or bent, anthers

often densely hairy and joined by a ligature; staminodes 3. Disk present, often annular,

occasionally partial, or absent. Ovary unilocular with parietal placentation, mostly

fusiform; style distinct; stigma chiritoid (see Introduction above); ovules many. Fruit

a narrow, bivalved capsule. Seeds many.

Key to Microchirita species in Thailand

la. Inflorescence with distinct bracts in inflorescence, bracts free or fused together..

. 2

lb. Inflorescence without distinct bracts in inflorescence.6

2a. Glandular hairs present on pedicels and/or peduncles.3

2b. Glandular hairs not present on pedicels and/or peduncles.4

216

Card. Bull. Singapore 69(2) 2017

3a. Bracts fused at the base; leaves with dense short glandular hairs and long

eglandularhairs. 6.M. glandulosa

3b. Bracts free; leaves with only eglandular indumentum.11. M. involucrata

4a. Corolla 32-45 mm long, lobes orange.16. M marcanii

4b. Corolla 16.5-33 mm long, lobes white to purple or blue.5

5a. Leaves sessile to shortly petiolate, < 2.5 cm long; leaf blades 2.2-7.2 times as

long as wide, with a silvery white “soft” indumentum; stems with indistinct

internodes.18 .M. mollissima

5b. Leaves with petioles 2-10 cm long; leaf blades 1.5-2.7 times as long as wide,

with a hispid “rough” indumentum; stems with well-defined internodes.

.22. M. rupestris

6a. Glandular hairs present on pedicels and/or peduncles in addition to any

eglandular indumentum.7

6b. Glandular hairs not present on pedicels and/or peduncles in addition to any

eglandular indumentum.10

7a. Corolla lobes pale yellow; calyx lobes 14-17 mm long.15. M. luteola

7b. Corolla lobes blue or purple; calyx lobes 4.5-15 mm long.8

8a. Corolla with two spots inside the lateral lobes; calyx lobes 13-15 mm long.

.3. M. aratriformis

8b. Corolla without spots inside the lateral lobes; calyx lobes 4.5-8.5 mm long.9

9a. Corolla tubular, widening abruptly; upper lobes dark purple at base.

.26. M. thailandica

9b. Corolla with a narrow lower tube which broadens into a campanulate upper

tube; upper lobes without a dark patch.14. M. limbata

10a. Anthers glabrous.11

10b. Anthers hairy.15

11a. Corolla >18 mm long, with a patch of elongated glands internally between the

upper lobes.18. M. mollissima

lib. Corolla <15 mm long, without a patch of elongated glands between the upper

lobes (but sometimes with shorter and more evenly distributed glands on the

upper lobes).12

12a. Corolla with a broadly campanulate upper tube, lobes very pale lilac.

.13. M. lilacina

12b. Corolla with an upper tube which is not campanulate, lobes (at least the lower)

pure white.13

A revision of Microchirita in Thailand

217

13a. Corolla personate, upper lip greenish white and much smaller than lower lip.

.20. M. personata

13b. Corolla not personate, upper lip pure white and of similar size to lower lip .... 14

14a. Corolla curved downwards and trumpet-shaped; fruit 2-5 cm long, sparsely

hairy.1. M. albiflora

14b. Corolla not curved downwards and tubular; fruit 1.2-2 cm long, densely hairy

.7. M. hamosa

15a. Corolla with obvious dark markings either at the base of the lateral lobes, at the

base of the upper lobes, or deeper inside the throat.16

15b. Corolla without obvious dark marking and fairly uniform in colour.27

16a. Calyx with glandular hairs outside.9. M. huppatatensis

16b. Calyx without glandular hairs outside.17

17a. Corolla lobes lilac, violet, purple or blue.18

17b. Corolla lobes white, yellow or orange.20

18a. Elongated glands present inside sinus of upper corolla lobes; anthers coherent

but not apically joined by a connective [Peninsular Thailand].28. M. viola

18b. Elongated glands absent inside sinus of upper corolla lobes; anthers apically

joined by a connective [SE, N and NE Thailand].19

19a. Corolla tube gradually flaring, not abruptly widening, lobes spreading,

predominantly whitish, pale yellow or blue [N & NE Thailand]... 25. M. tetsanae

19b. Corolla tube of a distinct narrow lower tube and abruptly widening into a

campanulate upper tube, lobes not spreading, predominantly dark purple

[Chanthaburi].21. M. purpurea

20a. Corolla bright yellow with an obvious dark ring inside the throat.

.19. A/, oculata

20b. Corolla lobes white, yellow or orange without a dark ring in throat.21

21a. Corolla 32-45 mm long, lobes orange.16. M. marcanii

21b. Corolla 12-30 mm long, lobes white or pale to bright yellow.22

22a. Corolla lobes predominantly bright yellow.23

22b. Corolla lobes predominantly white to very pale yellow or cream.24

23a. Ventral corolla lobe 8-10 x 6.5-10 mm; filaments geniculate or twisted so as to

appear divided into two halves.5. M. elphinstonia

23b. Ventral corolla lobe 2.3-5.2 x 2.5-7.5 in in : filaments straight and uniform.

.4. M. bimaculata

218

Card. Bull. Singapore 69(2) 2017

24a. Corolla with dark spots at the base of the upper lobes.25. M. tetsanae

24b. Corolla with dark spots at the base of the lateral lobes.25

25a. Corolla ventral lobe < 5 mm long, overall corolla length 12-18 mm long, lobes

white.12. M. karaketii

25b. Corolla ventral lobe > 5 mm long, overall corolla length 17-25 mm long, lobes

white or yellow-white.26

26a. Corolla with yellow-white lobes and yellowish throat in addition to the ventral

yellow line and lateral markings; anthers with a strongly dimorphic indumentum

.29. M. woodii

26b. Corolla with pure white lobes and white in throat in addition to the ventral

yellow line and lateral markings; anthers with only one type of hairs.

.27. M. tubulosa

27a. Anther indumentum of two distinct hair types.28

27b. Anther indumentum of a single hair type.30

28a. Filaments glabrous. 2.M. albocyanea

28b. Filaments eglandular hairy.29

29a. Calyx 7.5-13 x 1.2-1.7 mm, apex acute; corolla yellow.17. M. micromusa

29b. Calyx 4.5-8 x 1-1.2 mm, apex acuminate; corolla white or purple-blue.

.25. M. tetsanae

30a. Corolla pale yellow.24. M tadphoensis

30b. Corolla white, lilac, purple or blue.31

31a. Corolla with a ventral yellow or orange stripe.32

31b. Corolla without a yellow ventral stripe.33

32a. Corolla with a narrow tube abruptly opening up into spreading lobes, corolla

white or blue, with a fringe of glandular hairs at the base of the upper lobes

[Chaiyaphum, Loei].10. M. hypocrateriformis

32b. Corolla broadly campanulate, white to pale lilac except at very base of tube,

with short stalked glands not forming an obvious fringe [Kanchanaburi].

.8. M. hemratii

33a. Corolla 18-33 mm long; leaves softly tomentose [Peninsular Thailand].

.18. M. mollissima

33b. Corolla 10-13 mm long; leaves sparsely eglandular hairy [Northern Thailand]

.23. M. suddeei

A revision of Microchirita in Thailand

219

1. Microchirita albiflora D.J.Middleton & Triboun, Thai Forest Bull., Bot. 41: 19

(2013). - TYPE: Thailand, Chiang Rai, Mae Fa Tuang District, Road to Doi Tung

Royal Residence, 1000 m alt., 23 September 2008, Middleton, D.J., Karaket, R,

Triboun, P., Kawatkul, U. & Meeboonya, R. 4567 (holotype BKF; isotypes BK, E

[E00629491], K, P [P00966762], QBG, SING [SING0229831]). (Fig. 2A-C)

Caulescent herb to 50 cm tall, with internodes to c. 10 cm, unbranched. Stems fleshy,

glabrescent or with very sparse short eglandular hairs, pale green or sometimes purple-

brown at the base. Leaves opposite, except for the basal leaf; petioles 0.2-1.7 cm

long, basal leaf often sessile, glabrescent; blades mid-green above, pale green beneath,

ovate, 2.6-23 x 1.7-11 cm, 1.5-2.2 times as long as wide, base cordate, apex acute to

shortly acuminate, eglandular tomentose above, hispid beneath, margin entire, 7-12

pairs of secondary veins, tertiary venation laxly reticulate. Inflorescences cristate,

peduncles reduced or emerging to 1-3 mm long, fused with each other, glabrous, bracts

absent; pedicels 5-10 mm long, glabrescent or sparsely eglandular hairy. Calyx green,

bilabiate; tube 0.5-3.7 mm dorsally, c. 0.2 mm ventrally, lobes narrowly lanceolate

to ligulate, upper lobes 5-7.8 mm long, lower lobes 4-7.8 x 0.5-1.2 mm, sparsely

eglandular pubescent especially on midrib and tip, glabrous inside, apex acuminate.

Corolla 7-15 mm long, white, sometimes with a ventral yellow line, tube curved

downwards, trumpet-shaped, lobes not spreading, outside glabrous basally, eglandular

tomentose above, inside glabrous, with a glandular patch below the upper lobes; tube

6.5-13 mm long, equal dorsally, laterally and ventrally; lobes broadly orbicular, upper

lobes 1.6-1.9 x 2.2-3.1 mm, lateral lobes 1.6-2.7 x 2-3 mm, lower lobe 1.4-2 x 2.6

mm. Stamens arising 2.7-4.7 mm above the corolla base, filaments straight, minutely

glandular (glabrous in protologue), 2.1-3.1 mm long, 0.2-0.5 mm wide; anthers

glabrous, 0.6-1.1 x 1-2 mm, free or coherent by a weak ligature, thecae divergent;

lateral staminodes c. 0.6 mm long, arising near the corolla base, central staminode c.

0.4 mm long, arising near the corolla base. Disk partial, ventral, lobed, 0.3-0.7 mm.

Pistil c. 12 mm long; ovary 4.7-6 mm long, c. 0.6 mm diameter, glabrous or papillose

in lower 2/3, densely pubescent in upper 1/3, many ovules; style c. 4 mm long, densely

pubescent; stigma deeply bilobed, 0.7-1 mm long. Fruit green, 2-5 cm long, 1-1.7

mm diameter, with sessile glands at base, then becoming sparsely eglandular hairy,

curved. Seeds dark brown, elliptic, 0.6-0.7 x 0.3-0.4 mm.

Distribution. Northern Thailand. India, (Myanmar?), Laos.

Habitat. On rocks in mixed forest.

Provisional IUCN conservation assessment. Near Threatened NT. In Middleton &

Triboun (2013) this species was given a conservation status of Endangered. Since then

specimens have also been identified from NE India and Myanmar. From the known

localities, the AOO is still within the threshold for an Endangered assessment but large

swathes of territory within the large EOO, particularly in Myanmar, have not been

adequately explored to base an assessment on this. As limestone areas throughout the

220

Card. Bull. Singapore 69(2) 2017

region are threatened by mining and tourism an updated assessment of Near Threatened

is appropriate.

Additional specimens examined. THAILAND: Chiang Mai: Mae Taeng, Sop Berng, Mawk Fa

Falls, 650 m, 18 Aug 1990, Maxwell, J.F. 90-887 (A, CMU, L). Chiang Rai: Mae Fa Luang,

along main road, 500 m N of Phratamnak, Doi Tung Development Project offices, 23 Sep 2010,

van de Bult, M. 1076 (BKF); Mae Sai, Wat Tham Pha Jom, 500 m, 24 Sep 2008, Middleton,

D.J. et al. 4577 (BKF, E, P, QBG, SING); Mae Sai, Pong Ngam, Ban Tham, Tham Pla, 400

m, 28 Oct 2005, Maxwell, J.F. 05-607 (A, BKF, CMUB); Mae Sai, Pong Ngam, San Pa Sak,

Wat Tham Boom, 525 m, 7 Sep 2006, Maxwell, J.F. 06-646 (A, CMUB, QBG); Mae Sai, Pong

Ngam, Wat Tham Phum, 410 m, 2 Nov 2010, Norsaengsri, M. & Tathana, N. 7228 (BKF,

QBG); Mae Sai, Tahm Ban Santisuk, 550 m, 23 Sep 1989, Maxwell, J.F. 89-1120 (A, CMU, E).

Notes. This northern Thai species is recognisable by the white corolla and the glabrous

anthers. It is most easily confused with Microchirita hamosa, from which it differs

in the longer and more markedly broadened upper portion of the corolla (shorter and

tubular in M. hamosa ), and in the much longer and narrower capsule, which is also

very sparsely hairy (densely hairy in M. hamosa ). It differs from Microchirita suddeei

D.J.Middleton & Triboun in the glabrous anthers and in being pure white (not pale

lilac). The floral measurements reported were largely extracted from Middleton &

Triboun (2013) as not much new material was available for dissection.

2. Microchirita albocyanea C.Puglisi, sp. nov.

Most similar to Microchirita limbata C.Puglisi in the overall shape of the corolla and

in colour, but differs in not having a glandular indumentum and in the much longer

corolla and larger calyx. - TYPE: Thailand, Loei, Pha Khao, Wat Phu Phang, 447 m,

5 November 2014, Tetsana, N. et al. 876 (holotype BKF; isotype SING). (Fig. 2D-F)

Caulescent herb to 30 cm tall, internodes 3-5 cm. Stem succulent, sparsely eglandular

hairy; not branching. Leaves opposite, apart from the basal leaf; petioles 0.1-1 cm

long, sparsely eglandular hairy; blades mid green above, paler beneath, ovate to

elliptic, 3.5-14.2 x 1.5-8 cm, 1.5-3 times as long as wide, base shortly attenuate

to obtuse, apex acute to acuminate, eglandular pubescent above and beneath, margin

finely serrulate, 4-13 pairs of secondary veins, venation sunken above and raised

beneath in fresh material, flat in dry specimens, tertiary venation seldom visible in

dry specimens. Inflorescences cristate, 2-15-flowered; peduncles reduced and shortly

fused with each other, eglandular pubescent; bracts absent; pedicels pale green, 1-18

mm long, eglandular pubescent. Calyx greenish white, succulent at the base, tube c.

0.5 mm long, lobes narrowly lanceolate, c. 10-14 x 1.6-2.5 mm, apex acuminate and

thickened, margin entire, with eglandular hairs outside especially along the midrib and

the margin, glabrous inside. Corolla 21-25 mm long, tube white, throat white with a

yellow patch at the base of the filaments, lobes purple-blue, tube narrow at the base,

then becoming wider, not curved or only slightly so, lobes almost equal, glabrous at the

base, then becoming progressively hairier, eglandular hairy outside, glabrous inside;

A revision of Microchirita in Thailand

221

Fig. 2. Microchirita albiflora D.J.Middleton & Triboun. A. Habit. B. Front view of flower.

C. Side view of the flower. All from Middleton, D.J. et al. 4567. Microchirita albocyanea

C.Puglisi. D. Habit. E. Side view of the flower. F. Front view of the flower. All from Tetsana,

N. et al. 876. (Photos: A, B, P. Karaket; C, D.J. Middleton; D-F, N. Tetsana)

222

Card. Bull. Singapore 69(2) 2017

tube 17-19 mm long dorsally, 18-22 mm ventrally, 16-20 mm laterally between lips;

lobes elliptic, upper lobes 2.5-3.7 x 5-6.5 mm, lateral lobes 4.5-5 x 5-7.5 mm, ventral

lobe 4.5-5 x 4.5-7 mm. Stamens arising 8.5-9 mm above the corolla base; filaments

geniculate, yellow, glabrous, 2-3 mm long, 0.5-0.6 mm wide; anthers white, with a

woolly orange indumentum dorsally and at base, and short appressed orange hairs

around the apex, c. 1.5-2 x 3 mm, apically coherent and joined by a short ligature,

thecae divergent; lateral staminodes 1.5-3 mm long, arising 6-7 mm above the corolla

base, central staminode 0.3-0.5 mm long, arising c. 6 mm above the corolla base. Disk

annular, margin entire or cleft dorsally, 0.1-0.4 mm high. Pistil (immature) 15-17

mm long; ovary 8-9 mm long, 1.2-1.7 mm diameter, glabrous around the base, then

densely eglandular hairy; style c. 4 mm long, eglandular hairy, more sparsely towards

the stigma; stigma bilobed, lobes elliptic, 0.5-2 x 0.5 mm. Fruit green, 2.3-5 cm long,

1-1.7 mm diameter, eglandular hairy, straight or slightly curved. Seeds light brown,

narrowly elliptic, tri-tetrangular, mucronate, 0.2-0.5 x 0.1-0.5 mm.

Distribution. Northeastern Thailand (Loei).

Habitat. On limestone, in shade.

Provisional IUCN conservation assessment. Critically Endangered CR Blab(iii).

Although only known from a single collection the plant was collected from a limestone

outcrop which has a total potential area suitable for the growth of Microchirita species

of around 14 km 2 and which is not in a protected area. There has been development

within the limestone range and agricultural land encroaches right to the base of the

outcrop with the potential for changes in microclimate.

Notes. The epithet refers to the colour of the corolla tube and lobes.

3. Microchirita aratriformis (D.Wood) A.Weber & D.J.Middleton, Taxon 60: 778

(2011). - Chirita aratriformis D.Wood, Notes Roy. Bot. Gard. Edinburgh 31: 367

(1972); Wood, Notes Roy. Bot. Gard. Edinburgh 33: 197 (1974); Burtt, Thai Forest

Bull., Bot. 29: 87 (2001). - TYPE: [Vietnam], Tonkin, Langson, Khanmoi, Eberhardt

3332 (holotype P [P00602506]). (Fig. 3A-B)

Caulescent herb to 60 cm tall, branching, internodes to c. 7 cm. Stem green,

slightly fleshy, with a mixed indumentum of glandular and eglandular hairs. Leaves

opposite except for the basal leaf; petioles 0.5-3 cm long, eglandular hairy; blades

green above, paler beneath, lanceolate to elliptic, 4.8-10(-13+) x 2.5-10 cm, 1.6-

2.1 times as long as wide, base acute to shortly acute or subcordate, apex acute to

acuminate, sparsely eglandular hispid above and beneath, margin entire, 8-14+ pairs

of secondary veins (basal leaf not seen), tertiary venation not visible. Inflorescences

cristate, 4-10-flowered; peduncles 1-17 mm long, fused with each other, eglandular

hairy; bracts absent; pedicels 15—20(—31 reported in literature) mm long, glandular

A revision of Microchirita in Thailand

223

Fig. 3. Microchirita aratriformis (D.Wood) A.Weber & D.J.Middleton. A. Side view of the

flower. B. Front view of flower. All from Tetsana, N. et al. 871. Microchirita hamosa (R.Br.)

Yin Z.Wang. C. Cristate inflorescence and fruits. D. Habit. E. Side view of the flower. F. Front

view of flower. C from Middleton, D.J. et al. 4519; D from Middleton, D.J. et al 4522 ; E, F

from Middleton, D.J. et al. 5016. (Photos: A, B, N. Tetsana; C, D, D.J. Middleton; E, F, P.

Karaket)

224

Card. Bull. Singapore 69(2) 2017

and eglandular hairy. Calyx green, lobes divided almost to base, lanceolate, 13-15 x

1.2-1.5 mm, apex acuminate, glandular and eglandular hairy outside, especially along

margins and midrib, glabrescent inside. Corolla c. 23(-28 reported in literature) mm

long, white at base, turning purple above, lobes purple inside and outside, throat with

a yellow ventral line and lateral dark patches, tube slightly curved, broadening into

a pouched upper corolla, indumentum outside sparsely eglandular hairy, densely so

by the lobes, inside glandular hairy. Stamens with hairy anthers. Disk not seen. Pistil

enantiostylous, white to pale purple, ovary densely hairy. Fruit 2.1-9.5 cm long, 1.2-

1.8 mm diameter, densely eglandular hairy, straight or slightly curved. Seeds brown,

elliptic, 0.6-0.7 x 0.2-0.3 mm.

Distribution. Northeastern Thailand (Khon Kaen, Loei). Vietnam.

Habitat. On limestone rock in shade.

Provisional IUCN conservation assessment. Endangered EN B2ab(iii). This species is

quite widespread but only known from rather few collections in Thailand and Vietnam.

It is likely to also occur in Laos but the distribution is fragmented and the likelihood of

the AAO being >500 km 2 , even with new localities being discovered, is rather remote.

Throughout the region limestone habitats are threatened with mining, tourism and

land use changes. The assessment is not altered by changes in the status of the dubious

specimens below.

Additional specimens examined. THAILAND: Loei: Nong Hin, Num Sow, Phu Pha Lorn

Forest Park, 435 m, 5 Nov 2014, Tetsana, N. et al. 871 (BKF). Khon Kaen: Phuphaman NP,

300 m, 24 Aug 2011, Norsaengsri, M. et al. 8086 (QBG).

Dubious specimens. THAILAND: Loei: near Phu Kradung, Pha Nok Khao, 250-350 m, 6 Nov

1970, Charoenphol, C. et al. 4585 (AAU, E, L). Khon Kaen: Pha Nok Khao, 600-700 m, 9

Sep 1963, Smitinand, T. et al. 1129 (E, L).

Notes. Microchirita aratriformis is a poorly known species. The placement of any Thai

specimens into it has to be considered tentative and based on the presence of axillary

shoots, cristate inflorescence, pedicels with mixed indumentum types, ventral yellow

line, and an approximate match of calyx and corolla size. The flowering material

available for this study was extremely limited, therefore the only thorough description

of the inner flower parts remains that of Wood (1972, 1974). The specimens cited as

‘dubious’ above are those placed in this species by Burtt (2001) which are not good

collections and we cannot be entirely sure they belong here.

4 . Microchirita bimaculata (D.Wood) A.Weber & D.J.Middleton, Taxon 60: 778

(2011). - Chirita bimaculata D.Wood, Notes Roy. Bot. Gard. Edinburgh 31: 368

(1972); Wood, Notes Roy. Bot. Gard. Edinburgh 33: 196 (1974); Burtt, Thai Forest

A revision of Microchirita in Thailand

225

Bull., Bot. 29: 87 (2001). - TYPE: Thailand, Maeklang Falls, c. 50 km Northwest of

Chiang Mai, c. 430 m, 3 November 1967, Burtt, B.L. 5611 (holotype E [E00155280]).

(Fig. 4)

Caulescent herb to 50 cm tall, internodes 2-10 cm. Stems succulent, green or brown,

glabrous or glabrescent, unbranched. Leaves opposite, apart from the basal leaf;

petioles 0.2-1.5(-2.5) cm long, eglandular hairy; blades mid green above, pale green

beneath, lanceolate or ovate, 3.2-16.6(-30+) x 1.4—9.5(—19) cm (measures in bracket

refer to estimate measurements of incomplete basal leaves), 1.4-2.8 times as long

as wide, base cordate, rounded or seemingly shortly attenuate, apex acuminate or

acute, eglandular hairy above and beneath, not ciliate along the margin, margin entire,

8-15 pairs of secondary veins, venation slightly sunken above and raised beneath in

fresh material, flat in dry specimens, tertiary venation laxly reticulate. Inflorescences

cristate, peduncles reduced or emerging to 15 mm long and fused together; bracts

absent; pedicels green to reddish brown, 5-20 mm long, eglandular hairy. Calyx pale

to mid green, regular, succulent at base, tube 0.3-1 mm, lobes narrowly lanceolate

to elliptic, 5-10 x 0.7-1.8 mm, apex acute, margin irregularly entire, occasionally

toothed, hairy on midrib, margin and tip outside, glabrescent or glandular inside.

Corolla c. 21 mm long, bright yellow with a reddish brown patch ventrally inside,

sometimes split into two lateral patches, throat greenish yellow, narrow part of the

tube curved, then widening, upper lobes spreading, base of tube glabrous, broader part

sparsely eglandular hairy, lower lobes papillose, upper lobes and throat with scattered

glandular hairs; tube 10.5-18 mm long dorsally, 12-19 mm ventrally, 10.5-15 mm

laterally between lips; lobes elliptic, apices rounded, upper lobes 1-4 x 2.5-5.1 mm,

lateral lobes 2.5-5 x 3.3-6 mm, ventral lobe 2.3-5.2 x 2.5-7.5 mm. Stamens arising

5-11 mm above the corolla base; filaments straight, pale green, glabrous, 2-2.6 mm

long, c. 0.2-0.6 mm wide; anthers hairy dorsally, 1.1-2 x 0.6-1.1 mm, apically joined

by a connective, thecae subparallel; lateral staminodes 0.6-1.5 mm long, arising 3-6.5

mm above the corolla base, central staminode 0.4-0.5 mm long, arising 7-8 mm

above the corolla base. Disk annular or cleft dorsally, margin entire, 0.2-0.9 mm high.

Pistil 11.5-15 mm long; ovary 2-6 mm long, papillose at the base, apically eglandular

hairy; style 5-7.5 mm long, pubescent, eglandular in the bottom half, becoming

predominantly glandular in the top half; stigma glabrous, 0.5-1.1 mm long. Fruit

green, 1.5-5 cm long, 1.2-2 mm diameter, glabrous with sparse pubescence at the tip

(where the persistent style starts), usually curved at maturity, occasionally straight.

Seeds brown, narrowly elliptic, 0.3-0.4 x c. 0.1 mm.

Distribution. Northern and Northeastern Thailand. Laos, India. Likely also to be in

Myanmar.

Habitat. On limestone in deciduous forest.

Provisional IUCN conser\>ation assessment. Least Concern LC. This species is known

over a wide area and from many localities within Thailand. Its distribution in Myanmar

226

Card. Bull. Singapore 69(2) 2017

Fig. 4. Microchirita bimaculata (D.Wood) A.Weber & D.J.Middleton. A. Unifoliate plants. B.

Caulescent plants. C. Detail of a unifoliate plant. D. Front view of the flower. E. Fruits. F. Side

view of the flower. A, C from Suddee, S. et al. 4970\ B from Middleton, D.J. et al. 4514\ D from

Middleton, D.J. et al. 4520 ; E, F from Middleton, D.J. et al. 4479. (Photos: A, C, S. Suddee;

B, D-F, P. Karaket)

A revision of Microchirita in Thailand

227

is currently unknown but is likely to occur there. As limestone habitats are threatened

throughout the region the status of this species should be monitored.

Additional specimens examined. THAILAND: Buengkan: Bung Khla, Phu Wua Wildlife

Sanctuary, Tham Foon Waterfall, 167 m, 21 Oct 2015, Suddee, S. et al. 4970 (BKF, SING).

Chiang Mai: Cultivated at RBGE from Burtt 5611, C5927, (E); Chiang Dao, Doi Chiang Dao

Wildlife Sanctuary, 28 Aug 1935, Garrett, H.B.G. 1280 (K); ibidem, 13 Oct 1926, Put, N. 306

(ABD, BM, K); ibidem, Bunchuai, K. 260 (BKF); ibidem, 1100-1800 m, 13 Sep 1967, Tagawa,

N. et al. T-9916 (BKF); ibidem, 28 Aug 1935, Garrett, H.B.G. 1002 (E, K [2 sheets], L, P);

ibidem, 1100-1800 m, 13 Sep 1967, Tagawa, N. et al. T-9915 (BKF); ibidem, 9 Nov 1962,

Smitinand, T. et al. 7734 (BK); Chiang Dao, Doi Chiang Dao Wildlife Sanctuary, Ban Chiang

Dao, 25 Aug 1990, Maxwell, J.F. 90-902 (A, CMU, L); Doi Chiengdao, Ban Tham, 500 m,

14 Aug 1963, Smitinand, T. & Sleumer, H.O. 1003 (BKF); Wat Chiang Dao, c. 450 m, 24 Sep

1971, Murata, G. et al. T-14857 (BKF, L); Khao Chiang Dao, 30 Oct 1963, Bunchuai, K. 1294

(BKF); Chiang Dao, Doi Chiang Dao Wildlife Sanctuary, trail to Tam Chiang Dao, 500 m, 20

Sep 2008, Middleton, D.J. et al. 4520 (BKF, E, P, SING); Foot of Doi Chiang Dao, 500-600

m, 11 Sep 1967, Tagawa, N. et al. T-9785 (BKF); Doi Inthanon, Mae Pan Fall, c. 1400 m, Oct

1979, Santisuk, T. s.n. (BKF); Chom Tong, Doi Inthanon National Park, Wachirathan Waterfall,

710 m, 19 Sep 2008, Middleton, D.J. et al. 4514 (BKF, E); Doi Suthep, 10 Oct 1987, Maxwell,

J.F. 87-1158 (CMU, L); ibidem, 8 Sep 1958, S0rensen, T. et al. 4829 (ABD); ibidem, 1500 ft,

14 Nov 1909, Kerr, A.F.G. 897 (BM, K, L, P); ibidem, 2400 ft, 25 Sep 1910, Kerr, A.F.G. 1422

(BM, K, L, P); ibidem, 1000 m, 5 Sep 1958, S0rensen, T. et al. 4751 (ABD, BKF); Summit

of Doi Su Thep, 14 Oct 2003, Mattapha, S. 425 (BKF); Mueang Chiang Mai, Doi Suthep-Doi

Pui National Park, Montatarhn waterfalls, 1 Nov 2005, Palee, P. 882 (A); Doi Suthep-Doi Pui

National Park, above Huay Dteung Tau Lake, 825-875 m, 28 Aug 1990, Maxwell, J.F. 90-918

(L); East side of Doi Sutep, Kohntatahn Falls, 600 m, 4 Nov 1987, Maxwell, J.F. 87-1366

(BKF, CMU, L); Muang Chiang Mai, Doi Suthep-Doi Pui National Park, San Gou, 1030 m,

17 Sep 2008, Middleton, D.J. et al. 4479 (BKF, E, P); Hang Dong, Ban Pong, Tham Takkatan

(Grasshopper Cave), above Nam Sum (Mong), 850-930 m, 3 Sep 2003, Maxwell, J.F. 03-270

(BKF, L); Mae Rim, Pong Yeang, Pong Taa Hoen, 1000 m, 8 Sep 1995, Nanakorn, W. 4160

(E, QBG); Mae Rim, Pong Yeang, c. 1100 m, 16 Aug 1994, Nanakorn, W. et al. 1348 (E,

QBG); Mae Taeng, Doi Pah Dae, 9 Aug 1996, Maxwell, J.F. 96-1063 (A, L); Mae Dtang, Geut

Chang, Pad Dahn village, Doi Saing Liang, 1025 m, 20 Sep 1997, Maxwell, J.F. 97-1011 (A,

BKF, L); Mae Taeng, Ban Keud, c. 520 m, 25 Aug 1994, Nanakorn, W. et al. 1458 (QBG);

Mueang Chiang Mai, 12 Sep 1982, Fusai, P. 7 (CMUB); Inthanon, Mae Paam Falls, 1000 m,

5 Oct 1999, Suksathan, P. 2336 (QBG); Mae Wang, Mae Win, Wang Pah Boon, 24 Aug 2004,

Palee, P. 695 (A). Lampang: Chae Horn, Jae Sawn National Park, east side, 525 m, 25 Aug

1995, Maxwell, J.F. 95-556 (BKF, L); Chae Horn, Chae Son National Park, Tad Rung, 527 m,

2 Sep 2009, Norsaengsri, M., et al. 6042 (QBG); Jae Son National Park, 300 m, 8 Nov 1999,

Suksathan, P. 1809 (E, QBG); Doi Pang La, Huay Tak, 400 m, 25 Sep 1967, Shimizu, T. et al.

T-10790 (BKF); en route from Pang La to Huay Tak, 350 m, 24 Sep 1967, Shimizu, T. et al.

T-10724 (BKF). Lamphun: Mae Tah, Doi Kuhn Dahn National Park, South side, Doi Hoa

Chang, 1225 m, 4 Sep 1994, Maxwell, J.F. 94-981 (BKF, L). Mae Hong Son: Mae La Noi,

road from Mae Sariang to Mae Hong Son, 560 m, 20 Oct 2014, Middleton, D.J. et al. 5800

(BKF, E). Phitsanulok: Thung Salaeng Luang National Park, Kaeng Sopa Waterfall, 375-410

m, Murata, G. et al. T-38537 (BKF); ibidem, 375—410 m, Murata, G. et al. T-38516 (BKF);

ibidem, 17 Sep 1990, Chantharanothai, P. et al. 90/298 (K); Thung Salang Luang, Pinnin, S. et

al. 113 (BKF, E); Thung Salang Luang, S.P. [Pinnin, S.?] et al. 84 (BKF); Tung Salaeng Luang

228

Card. Bull. Singapore 69(2) 2017

National Park, Kang So Pa Waterfall, 350 m, 11 Oct 1979, Shimizu, T. et al. T-18365 (BKF).

Tak: Mae Ramad, Ban Nam Ok-roo, 15 Sep 2005, Pooma, R. 5687 (BKF); Tha Song Yang,

road to Mae Sariang, 832 m, 19 Oct 2014, Middleton, D.J. et al. 5790 (BKF, E); Tha Song

Yang, Tham Ook Ru Temple, 100 m, 6 Sep 2013, Phaosrichai, P. & Wongnak, M. 21 (QBG).

Notes. This species is broadly distributed across the North of Thailand. It is recognisable

by the bright yellow corolla with lateral dark spots inside the throat, the often curved

fruit, steep secondary veins, and the dull orange-silver hue of the abaxial side of the

dry leaves.

5 . Microchirita elphinstonia (Craib) A.Weber & D.J.Middleton, Taxon 60: 778

(2011). - Chirita elphinstonia Craib, Bull. Misc. Inform. Kew 149 (1932); Barnett, FI.

Siam. 3: 224 (1962); Wood, Notes Roy. Bot. Gard. Edinburgh 33: 195 (1974); Burtt,

Thai Forest Bull., Bot. 29: 87 (2001). - TYPE: Cult, in Hort. Aberdeen from seeds

of Marcan 2561, coll. Thailand, Krabin, Ban Keng, 30 m, limestone hill (lectotype

K [K000545615], designated by Wood (1974: 195); isolectotypes A [00054810],

ABD, B [B100277728], BK, BR [692324], C [C10012723], CHR [CHR213604], E

[E00155273], L, LE [LE00017212] P [P00602511], SING, US [00126000]).

Caulescent herb to 50 cm tall, internodes 3-10 cm long. Stems succulent, glabrous,

branching from the petioles. Leaves opposite, apart from the basal leaf, but sometimes

2 single basal leaves are present; petioles 0.5-1.4 cm long, sparsely eglandular hairy;

blades ovate, 1-10.5 x 2-6.3 cm (paired leaves, basal leaf not measured), 1.5-1.8 times

as long as wide, base shortly attenuate to seemingly cordate, apex acute to shortly

acuminate, with short eglandular hairs above and very sparse hairs beneath, minutely

ciliate along the margin, margin entire, 4-21+ pairs of secondary veins. Inflorescences

cristate, can be congested and many-flowered; peduncles 0.2-5 mm long, fused with

each other, glabrous and curved; bracts absent; pedicels 8-12 mm long, glabrescent or

sparsely hairy, curved. Calyx pale green, actinomorphic, tube 0.5-1 mm, lobes elliptic,

8-15 x 1.5-1.9 mm, apex narrowly acute, margin entire, sometimes with indistinct

small teeth near the apex, eglandular hairy outside, especially on tip and margin, inside

with sessile glands. Corolla 20-30 mm long, bright yellow, with lateral dark spots in

throat, tube narrow, curved, then progressively widening, lobes spreading, eglandular

hairy outside, lobes papillose, ventral lobe prominent; tube 17-21 mm long dorsally,

17-20 mm ventrally, 17-23 mm laterally between lips; lobes elliptic, apices obtuse

to rounded, margin irregular, upper lobes 2-4.5 x 5-6 mm, lateral lobes 5-7 x 6-7.5

mm, ventral lobe 8-10 x 6.5-10 mm. Stamens arising 9.5-12 mm above the corolla

base; filaments geniculate or twisted in the middle, pale in the lower half, darker

above, glabrescent or sparsely hairy below the knee, glandular above, 4-5 mm long,

0.3-0.5 mm wide; anthers yellow, with long eglandular hairs dorsally, 2.5 x 1.7-2.5

mm, apically joined by a connective, thecae divergent; staminodes not seen. Disk

partial, 0.4-0.5 mm high, or complete and 0.6-0.8 mm long. Pistil 18.5-20 mm long;

ovary 5-8 mm long, 0.7-1.2 mm diameter, glabrous at the base, hairy apically; style

A revision of Microchirita in Thailand

229

7-10 mm long, hairy, becoming glabrescent towards apex; stigma glabrous, 0.5-0.6

mm long. Fruit straight or curved, 4-6.5 cm long, sparsely hairy. Seeds dark brown,

elliptic, 0.4-0.5 x 0.2-0.3 mm.

Distribution. Northeastern to Southeastern Thailand. Cambodia.

Habitat. On limestone in evergreen forest.

Provisional IUCN conservation assessment. Endangered EN B2ab(iii). Although

this species has a large EOO, it is known from rather few collections and has a

currently known AOO well within the range for an Endangered status, coupled with

a fragmented distribution and the many threats to limestone habitats throughout the

region from mining, tourism and land use changes. Within the EOO there are rather

limited opportunities for the AOO to increase beyond the threshold for Endangered

due to the lack of suitable habitats.

Additional specimens examined. Unknown origin: Cultivated at RBGE from material grown

at University Botanic Garden, Utrecht, ace. 800333, P12, Nov 1980, C13665 (E); Cultivated

at RBGE from material grown at University Botanic Garden, Utrecht, ace. 800335, P12, Nov

1980, C13667 (E); Cult Cantonspark, Mennega, E.A. 61-70 (E); Entry s.n. (K [2 sheets]));

Cultivated in Hort. Aberdeen, 1931, “Kerr” 337 (L); Cultivated in Hort. Aberdeen, 1931,

“Kerr” 338 (L). THAILAND: Khon Kaen: Phu Pha Man, Tadyai Waterfalls, 18 Nov 2011,

Lakoet, C. 237 (QBG). Loei: Phu Luang Wildlife Reserve, Nam Tok to Pa Paw trail, 29 Sep

1990, Chantaranothai, P. et al. 90/439 (BKF, K). Nong Khai: Ban Phu Ngam, Singhanat Ban

Phot temple, 150 m, 26 Aug 2001, Pooma, R. et al. 2686 (BKF [2 sheets]). Prachin Buri:

Krabin, Kao Sungto, c. 50 m, 10 Nov 1930, Marcan, A. 2561 (ABD [2 sheets], BK, BM [2

sheets], K [2 sheets], E); Cultivated at Smithsonian Institution from Marcan 2561, accession

78-139, Skog, L.E. & Staton-Hodapp, S. 5333 (US). Sa Kaeo: Khao Chakan Temple, 88 m, 17

Oct 2010, Staples, G. et al. 1407 (BKF, E, K, SING). Sisaket: Khun Han, Sam Rongkiat Falls

Arboretum, 200 m, 22 Dec 2005, Pooma, R. et al. 6094 (BKF, E).

Notes. Microchirita elphinstonia is a yellow-flowered species recognisable by the

filament, which is somewhat divided in two halves (it can be geniculated or more

commonly twisted), and appears darker in the upper part than in the lower in dry

specimens. The leaves are ovate and the secondary veins can be numerous and very

close to each other, although this is not a consistent character, e.g. in the type material.

6. Microchirita glandulosa C.Puglisi, sp. nov.

Similar to Microchirita involucrata (Craib) Yin Z.Wang and M. rupestris (Ridl.)

A. Weber & Rafidah) in having bracteate inflorescences. Differs from both in the bracts

being fused only at the base (i.e. not divided as in Microchirita involucrata and not

fused into a cup as in M. rupestris ), in the dimorphic indumentum of sparse, long

eglandular hairs and dense short glandular hairs on the leaf (eglandular indumentum

in M. involucrata and M. rupestris ), and in the tripartite calyx. It differs further from

230

Card. Bull. Singapore 69(2) 2017

Microchirita involucrata in the serrate margin of the bracts and from M. rupestris in

the much smaller size of the bracts. - TYPE: Thailand, Nan, Song Kwaw, Sakoen,

Khao Tham Plakang, 750 m, 3 September 2006, Watthana, S. 2126 (holotype QBG;

isotype CMU).

Herb to 50 cm tall. Stems fleshy, sparsely glandular hairy. Leaves opposite, with the

exception of the single basal leaf; petioles 0.1-2 cm long, glandular hairy and with

sparse long eglandular hairs; blades lanceolate, 2-8.5 x 1-6.5 cm, 1.3-2.2 times as long

as wide, base cordate or obtuse, apex acute, with a dimorphic indumentum of dense and

short glandular hairs and longer and thicker eglandular hairs, margin delicately serrate,

8-10 pairs of secondary veins in the opposite leaves, tertiary venation inconspicuous.

Inflorescence 1-3-flowered, arising from the axil end of the petiole; peduncles 2-8

mm long, glandular hairy; bracts fused at the base and enclosing the pedicels, 0.6-1.1

x 0.3-0.5 mm, sessile or shortly petiolate, margin and indmentum matching those

of the leaves; pedicels 0.8-1.2 cm long, covered in minute glandular hairs. Calyx

zygomorphic, tube 1.5 mm dorsally, 1 mm laterally and 0.5 mm ventrally, lobes linear

to oblanceolate, 4.5-7.5 x 0.8-2.2 mm, minutely toothed towards the apex, apex acute,

densely glandular hairy outside and inside. Corolla 15-18 mm long, reportedly white,

tube with a narrow basal portion, slightly curved downwards, then progressively

broadening, externally with a minute glandular indumentum; tube c. 12.5 mm long

dorsally, 15-16 mm ventrally, 13 mm laterally between lips; upper lobes elliptic, c.

1.5 x 2.5 mm, lateral and lower lobes seemingly small but not measured. Stamens

arising c. 7 mm above the corolla base; filaments glabrous, c. 3.5 mm long, S-shaped;

anthers with a dense indumentum at the insertion, c. 1.3 x 2.5 mm, apically joined by

a connective, thecae subparallel; lateral staminodes c. 0.5 mm long, arising c. 6.5 mm

above the corolla base, central staminode c. 0.7 mm long, arising c. 6.5 mm above

corolla base. Disk annular, 0.4-0.5 mm high. Pistil 12-14 mm long; ovary 5-6 mm

long, c. 1.5 mm diameter, with minute glandular hairs throughout its length, except

for the lowermost half millimetre, which is glabrous; style c. 6 mm long, covered in

the same indumentum as the ovary; stigma lobes c. 1.5 mm long, glabrous. Fruit and

seeds not seen.

Distribution. Northern Thailand (Nan).

Habitat. On exposed limestone.

Provisional IUCN conservation assessment. Data Deficient DD. This species is

only known from a single collection from a National Park. There is extensive but

underexplored limestone in the region and more information is needed on the

distribution of the species and on potential threats before a satisfactory conservation

assessment can be given.

Notes. The epithet refers to the conspicuous presence of a glandular indumentum.

A revision of Microchirita in Thailand

231

7. Microchirita hamosa (R.Br.) Yin Z.Wang, J. Syst. Evol. 49: 60 (2011). - Chirita

hamosa R.Br., Cyrtandreae 117 (1839); Barnett, FI. Siam. 3: 224 (1962); Wood, Notes

Roy. Bot. Gard. Edinburgh 33: 191 (1974); Burtt, Thai Forest Bull., Bot. 29: 87 (2001).

- TYPE (conserved - see Middleton & Puglisi, 2015): Thailand, Tak, Umphang,

Umphang [Doi Hua Mot] Wildlife Sanctuary, 915 m, 17 October 2014, Middleton,

D.J., Hemrat, C., Karaket, P., Puglisi, C. & Suddee, S. 5762 (holotype E; isotypes

BKF, SING). (Fig. 3C-F)

Caulescent herb to 25 cm tall; internodes 3-6 cm, although more often than not

appearing as a unifoliate plant. Stems fleshy, with long eglandular hairs, pale green

or sometimes tinged with purple-brown. Leaves opposite, except for the single basal

cotyledon; petioles 0.1-0.5 cm long, densely hairy; blades green above, pale green

beneath, ovate to lanceolate, 2-13.3 x 1.1-9.3 cm, 1.2- 2.1 times as long as wide,

base of macrocotyledon/basal leaf cordate, cauline leaves shortly attenuate, apex

acute, densely eglandular tomentose above and beneath, margin entire or subentire,

6-12 pairs of secondary veins, venation slightly sunken above and raised beneath in

fresh material, tertiary venation laxly reticulate. Inflorescences cristate, few to many

flowered; peduncles to 4 mm long, not fused with each other; bracts absent; pedicels

pale green, 3-20 mm long, all axes with long eglandular hairs. Calyx green, regular, tube

0.1-0.3 mm, lobes narrowly lanceolate, 3-4.5 x 0.4-0.7 mm, eglandular hairy outside,

glabrous inside except at the tip, apex acuminate, margin entire. Corolla 8.5-13 mm,

white with a yellow stripe ventrally in tube, tube tubular, not curved downwards, lobes

not spreading, outside eglandular hairy, glabrescent, becoming densely eglandular

hairy towards the lobes, inside densely glandular, especially on lobes; tube 8-9 mm

long, equal dorsally, laterally and ventrally; upper lobes 0.7 x 1 mm, lateral lobes

1.1 x 1.1 mm, lower lobe 1 x 0.7 mm. Stamens arising c. 3.9 mm above the corolla

base; filaments white or pale yellow, straight, glabrous, c. 1.5 mm long, 0.1 mm wide;

anthers white or pale yellow, glabrous, 0.6-1.1 x 1-2 mm, free, thecae divergent;

staminodes not seen. Disk a single ventral lobe, 0.1 mm. Pistil c. 16 mm long; ovary c.

14 mm long, c. 0.7 mm diameter, glabrous or papillose in lower 1/3, densely pubescent

in upper 2/3, many ovules; style c. 3 mm long, densely pubescent; stigma shallowly

bilobed, c. 0.2 mm long. Style and stigma green to white. Fruit green, 1.2-2 cm long,

1-1.2 mm diameter, glabrous at the base, then becoming densely eglandular hairy,

straight or slightly curved. Seeds dark brown, elliptic, 0.2-0.3 x 0.1-0.2 mm.

Distribution. Northern and Northeastern Thailand. India, China, Myanmar, Laos,

Vietnam.

Habitat. On limestone rocks or sandy soil in dry dipterocarp forest.

Provisional IUCN conservation assessment. Least Concern LC. This species is

common and widespread although could potentially be impacted by removal of

limestone habitats for cement.

232

Card. Bull. Singapore 69(2) 2017

Additional specimens examined. Unknown locality: Cultivated in Cantonspark, Utrecht (NL),

19 Aug 1964, Mennega, accession 62-1556 (E). Kerr, A.F.G. 2174 (E). THAILAND: Bueng

Kan: Bungkhla, Phu Wua Wildlife Sanctuary, 200 m, 27 Aug 2001, Pooma, R. et al. 2796

(BKF). Chiang Mai: Chom Tong, Mae Soi Valley, Pa Peung area, 675 m, 1 Oct 1991, Maxwell,

J.F. 91-816 (E, L, P); Chiang Dao, road to WiangHaeng, 700 m, 21 Sep 2008, Middleton, D.J.

et al. 4540 (BKF, E); Chiang Dao, Ban Pak Chiang, 31 Oct 1963, Adisai 586 (BK); Chiang

Dao, Doi Chiang Dao Wildlife Sanctuary, 500 m, 20 Sep 2008, Middleton, D.J. et al. 4519

(BKF, E, P); Chiang Dao, Doi Chiang Dao Animal Sanctuary, north side of Doi Luang, 975

m, 11 Oct 1995, Maxwell, J.F. 95-901 (BKF, CMU, L); Chiang Dao, Doi Chiang Dao Wildlife

Sanctuary, Tam Pha Phlong, 600 m, 20 Sep 2008, Middleton, D.J. et al. 4522 (BKF, E); Chiang

Dao, Tham Chiang Dao, 9 Sep 1999, Srisanga, P. et al. 1020 (QBG); Chiang Dao, Kio Phawok

border checkpoint, 30 Sep 2009, Middleton, D.J. et al. 5016 (BKF, E, P, SING); Doi Chiang

Dao, Pa Blawng Cave area, 550 m, 21 Oct 1989, Maxwell, J.F. 89-1283 (L). Mae Teng, Pa Pae,

Mok Fah Waterfalls, 620 m, 13 Sep 1999, Srisanga, P. & Puff, C. 1070 (BKF, E, QBG); Mae

On, Mae On Cave, 500 m, 7 Sep 2011, Pooma, R. et al. 7796 (BKF, E [2 sheets]); Sangaapang,

Muang Awn Cave, 525 m, 12 Oct 1989, Maxwell, J.F. 89-1217 (A, CMUB, L); San Kamphaeng,

Sahagon, Doi Muang Awn, 15 Sep 1998, Palee, P. 407 (CMUB); Mae Ping Rapids, 26 Nov

1920, Kerr, A.F.G. s.n. (BM); Payap, Sop Aep, Hennipman, E. 3346 (L). Chiang Rai: Doi Tam

Yup, c. 390 m, 14 Sep 1924, Garrett, H.B.G. 198 (ABD [2 sheets], BK, BM, K [2 sheets]).

Kamphaeng Phet: Khlong Lan, Khlong Lan National Park, Khong Lan Waterfalls, 200 m, 4

Nov 2010, Pooma, R. et al. 7469 (BKF). Lamphun: Mae Tah, Doi Khun Dahn National Park,

Daht Muey Falls, 800 m, 23 Oct 1993, Maxwell, J.F. 93-1269 (A, BKF, CMU); Mae Tah, Doi

Khun Dtan, Daht Muey Waterfall, 775 m, 30 Sep 2001, Palee, P. 497 (A, BKF, CMU); Mae

Tah, Doi Khun Dahn National Park, Tah Goo station area, 650 m, 28 Oct 1994, Maxwell, J.F.

94-1130 (A, BKF, CMU, L). Loei: Mueang Loei, Phu Pha Lorn Temple, 402 m, 9 Sep 2014,

Tetsana, N. et al. 797 (BKF, SING). Mae Hong Son: 600 m, 10 Sep 1974, Larsen, K. &

Larsen, S. 34337 (AAU); Mae La Noi, road from Mae Sariang to Mae Hong Son, 560 m, 20

Oct 2014, Middleton, D.J. et al. 5801 (BKF, E); Pangmapha, Tahm Lawt, Nam Lawng River,

675 m, 7 Aug 1999, Maxwell, J.F. 99-111 (CMU, E, L); Bang Ma Pah, Ban Jah bo, Mae La Nah

Cave, 950 m, 30 Sep 2003, Palee, P. 632 (A, CMUB); Bahng Ma Pah, Tahm Lawt, Doi Pah

Baw, above Muang Pam village, 750 m, 10 Nov 2004, Palee, P. 771 (A, CMUB); Muang Mae

Hong Son, Pha Suea Waterfall National Park, Tham Pla, 225 m, 21 Oct 2014, Middleton, D.J.

et al. 5808 (BKF, E); Doi Pui, 800 m, 23 Sep 1995, Larsen K. et al. 46859 (AAU). Nan: Bo

Kluea, Doi Phu Kha National Park, 1210 m, 15 Aug 2012, Middleton, D.J. et al. 5607 (BKF,

E); Song Khwae, Tham Sakoen National Park, 700 m, 14 Aug 2012, Middleton, D.J. et al.

5598 (BKF, E, P, SING); Song Kwae, Tham Sakoen National Park, Pha Than Waterfalls, 16 Jul

2012, La-ongsri, W. et al. 2417 (QBG); Song Kwae, Tham Sakoen National Park, Tham Hai

Tak, 689 m, 27 Jul 2011, La-ongsri, W. et al. 1830 (QBG); Pua, Doi Phu Kha National Park,

1250 m, 1 Sep 2000, Srisanga, P. 1572 (E, QBG). Sakhon Nakon: Pha Nang Moen, Phu Phan

National Park, 31 Jul 1999, Newman, M.F. 947 (BKF, E).

Notes. This species is recognisable by the small, white flower with a tubular corolla,

and by the relatively short and densely hairy fruit. It is widespread across the north

of Thailand and in the neighbouring countries. It frequently flowers when only a

unifoliate plant.

Microchirita hamosa is conserved with a conserved type (proposed by

Middleton & Puglisi, 2015; supported by Nomenclature Committee for Vascular

Plants in Applequist, 2016).

A revision of Microchirita in Thailand

233

Wood (1974) listed a number of synonyms for Chirita hamosa, all from India.

As noted by Middleton & Triboun (2013), several different species were included in

Wood’s concept of Chirita hamosa so we refrain from listing any of these synonyms

pending a thorough revision of the genus in India.

8. Microchirita hemratii C.Puglisi, Kew Bull. 71(l)-2: 4 (2016). - TYPE: Thailand,

Tak, Mae Sot distr., Wat Tham Inthanin, 660 m, 18 October 2014, Middleton, D.J.,

Hemrat, C., Karaket, P., Puglisi, C. & Suddee S. 5775 (holotype BKF; isotypes E

[E00663027], SING). (Fig. 5A-B)

Herb to 50 cm tall. Stems fleshy, dark red or green at base, otherwise pale green,

glabrous or sparsely hairy. Leaves opposite, with the exception of the single basal leaf,

fleshy (very thin when dry); petioles 0.2-1.5 cm long, glabrous; blades pale green

above, paler beneath, lanceolate, 3.75-14 x 2.6-8.5 cm, 1.3-1.7 times as long as wide,

base obtuse to cordate, apex acute to acuminate, sparsely and minutely tomentose

above and beneath, above with additional scattered eglandular hairs, margin entire to

obscurely serrulate, 3-8 pairs of secondary veins in the opposite leaves, 7-13 in the

basal leaf, venation raised beneath in fresh material, tertiary venation almost invisible.

Inflorescence 1-8-flowered, cristate; peduncles reduced; bracts absent; pedicels green,

0.5-1.3 cm long, sparsely eglandular hairy. Calyx green, actinomorphic, tube 0.2-

0.5 mm long, lobes lanceolate, 3-9 x 0.5-1 mm, inconsistently with some serration

towards the apex, apex acute, outside sparsely eglandular hairy at base, hairier distally,

inside with an indumentum of sparse sessile glands. Corolla 10-15 mm long, narrow

basal portion greenish white, rest of tube white, with a yellow stripe ventrally inside,

lobes pale lilac, tube with a narrow basal portion, then broadening into a funnel-shaped

distal portion, this curved down, pubescent outside except at base, inside glabrous,

the upper lobes densely covered in stalked glands, the lower central lobe sparsely so,

the lower lateral lobes glabrous; tube 9-12 mm long dorsally, 10-11 mm ventrally,

8.5-9 mm laterally between lips, upper lobes broadly elliptic, 2.5-3 x 4^1.7 mm,

lower lateral lobes elliptic, 2-4 x 4-5 mm, lower central lobe rounded, 3-4 x 4-5 mm.

Stamens arising 4-6.5 mm above the corolla base, filaments glabrous, 3-3.5 mm

long, straight; anthers with a dense indumentum of long hairs dorsally, c. 1 x 0.7-0.8

mm, apically joined by a connective, thecae divergent; lateral staminodes c. 2 mm

long, arising 3-5 mm above the corolla base, central staminode c. 1 mm long, arising

c. 4 mm above corolla base. Disk absent. Pistil 8-12 mm long; ovary 3-5 mm long,

papillose, with pubescence restricted to the apex; style 5-6 mm long, apically curved,

hairy; stigma with lobes c. 0.5 mm long, c. 0.8 mm wide, narrowly elliptic, glabrous.

Fruit green, 2-6 cm long, 0.9-1.1 mm diam., glabrous at the base, sparsely hairy

distally, straight or curved. Seeds light brown with reddish apices, elliptic, 0.5-0.6 x

0.2-0.3 mm.

Distribution. Northern Thailand (Tak).

234

Card. Bull. Singapore 69(2) 2017

Habitat. On limestone in deciduous forest.

ProvisionalIUCNconservation assessment. Endangered EN B1 ab(iii,iv) + B2ab(iii,iv).

The known EOO of this species would qualify it for Critically Endangered but the

collecting localities are in a limestone range much of which has so far not been

explored and where it is also likely to occur. Even if it were to occur throughout this

range its EOO would still qualify it as Endangered. Parts of this range, including some

of the known localities, are outside protected areas and subject to disturbance from

visitors, particularly at the religious sites.

Additional specimens examined. THAILAND: Tak: Mae Sot, Tham Inthanin Temple, 640 m,

17 Oct 2013, Pooma, R. et al. 7851 (BKF, E); ibidem, 650 m, 5 Nov 2010, Pooma, R. et al.

7522 (BKF); ibidem, 500 m, 11 Sep 2009, Middleton, D.J. & Triboun, P. 4849 (BK, E); Maesot,

Phra Wo, Phra Wo Spirit House, 700 m, 5 Nov 2010, Pooma, R. et al. 7507 (BKF); Mae Sot,

Mae Kah Soh, Po Tip Tawng Cave, 300 m, 20 Aug 1994, Maxwell, J.F. 94-905 (A, BKF, L).

Notes. This species is recognisable by the pale lilac corolla with a yellow ventral line,

the broad mouth, and the dense anther indumentum. It is currently known only from

the Mae Sot district of Tak province.

9. Microchirita huppatatensis C.Puglisi, Kew Bull. 71(l)-2: 2 (2016). - TYPE:

Thailand, Uthai Thani, Lan Sak, Huppatat Non Hunting Area, 122 m, 14 October

2014, Middleton, D.J., Hemrat, C., Karaket, P, Puglisi, C., Suddee, S. 5689 (holotype

BKF). (Fig. 5C-D)

Herb to 40 cm tall. Stems fleshy, red at base and around the basal nodes, otherwise pale

green, pubescent. Leaves opposite, with the exception of the single basal leaf; petioles

0.5-1.2 cm long, pubescent; blades pale green above, paler beneath, lanceolate or

elliptic, 5-12.8 x 1.6-6.1 cm (undamaged basal leaf not seen), 2-3 times as long as

wide, base acute to obtuse, apex acuminate, minutely tomentose above and beneath,

margin entire, 5-9 pairs of secondary veins in the opposite leaves, at least 13 in the

basal leaf, venation raised beneath in fresh material, tertiary venation barely visible

and loosely reticulate. Inflorescence cristate, 4-15-flowered; peduncles reduced,

rarely emerging and fused together at the base; bracts absent; pedicels pale green,

0.2-2 cm long, pubescent. Calyx pale green, actinomorphic, tube 0.3-0.5 mm long,

lobes narrowly lanceolate, 8-9 x 0.7-1.1 mm, apex acute, outside hairy throughout,

with mixed glandular and eglandular hairs, inside with a sparse minute glandular

indumentum. Corolla 15-20 mm long, base of tube greenish white, rest of tube white,

ventral pouch of tube marked by a yellow, raised and papillose line running throughout

the inner surface of the tube, with a purple-brown streak on either side of it, tube with

a narrow basal portion, curved downwards, then abruptly broadening into a ventral

pouch, upper lobes reflexed, lower lateral lobes spreading, the central lobe straight,

limb externally sparsely hairy, with stalked glands at the base; tube 9-10 mm long

A revision of Microchirita in Thailand

235

Fig. 5. Microchirita hemratii C.Puglisi. A. Front view of flower. B. Side view of the flower. All

from Middleton, D.J. et al. 5775. Microchirita huppatatensis C.Puglisi. C. Detail of the flower.

D. Habit. All from Middleton, D.J. et al. 5689. (Photos: P. Karaket)

dorsally, 10-14 mm ventrally, c. 10 mm laterally between lips, upper lobes elliptic,

3-3.5 x 3.5^1 mm, lower lateral lobes elliptic, 5-6 x 4-5 mm, lower central lobe

rounded, 6-8 x 5-8 mm. Stamens arising 4-6 mm above the corolla base, filaments

yellow, minutely glandular, 1-1.5 mm long, with a thicker base and two geniculations;

anthers yellow, connective white, with a small patch of short hairs dorsally, 1.5-2 x

1.2-1.5 mm, held at a right angle, apically joined by a connective, thecae parallel; all

three staminodes much reduced, less than 0.5 mm long, arising 5-6 mm above the

corolla base. Disk an annular ring, slightly lobed, 1-1.3 mm high, whitish at base and

darker along the edge. Pistil c. 5 mm long; ovary 2.5-3.5 mm long, glabrous in the

basal 2/3, apically eglandular hairy; style 0.5-0.7 mm, glabrous; stigma with lobes

0.5-0.7 mm long, 0.4-0.6 mm wide, elliptic. Immature fruit pale green, c. 1.5 cm long,

2-3 mm diam., pubescent throughout its length, straight. Seeds not seen.

236

Card. Bull. Singapore 69(2) 2017

Distribution. Northern Thailand (Tak).

Habitat. On limestone in secondary forest.

Provisional IUCN conservation assessment. Critically Endangered CR Blab(iii,iv) +

B2ab(iii,iv). This species is only known from the type collection growing in a mixed

population with Microchirita personata. The limestone range there is only about 12

km 2 in total and is subject to disturbance from tourism. There are no collections from

the nearby Khao Pha Ra, and the area is surrounded by cultivated land.

Notes. Microchirita huppatatensis is most similar to M. woodii D.J.Middleton &

Triboun in the pattern of the corolla colour, but differs in the shorter corolla tube,

acuminate leaves and the overall corolla shape. The description and measurements

reported here match the original description (Puglisi et al., 2016), as no new specimens

have been collected since.

10. Microchirita hypocrateriformis C.Puglisi, sp. nov.

Differs from all other species of Microchirita (C.B.Clarke) Yin Z.Wang in the

combination of long, narrow corolla tube, abruptly opening into spreading limb, in the

long lower corolla lobe, and in the presence of a fringe of glandular indumentum at the

base of the upper lip. - TYPE: Thailand, Chaiyaphum, Khon Sarn, Wat Tham Huang

Po, 400 m, 19 October 2015, Suddee, S., Keiwbang, W., Hemrat, C. 4967 (holotype

BKF; isotype SING). (Fig. 6)

Herb to 50 cm tall with elongated stem runners. Habit caulescent, internodes 4-10 cm.

Stems succulent, with sparse eglandular indumentum; branches sometimes arising from

the petiole of the basal leaf. Leaves opposite, apart from the basal leaf; petioles 3-7 mm

long, sparsely eglandular hairy; blades pale green above, paler beneath, lanceolate to

elliptic, 3.5-22.5 x 2.4-17.5 cm, 1.1-2.4 times as long as wide, base shortly attenuate

to cordate, apex acute to acuminate, sparsely eglandular tomentose above and beneath,

ciliate along the margin, margin entire or sparsely and minutely denticulate, 5-17

pairs of secondary veins in the opposite leaves, venation slightly sunken above and

raised beneath in fresh material, flat in dry specimens, tertiary venation inconspicuous.

Inflorescences cristate, peduncles reduced or to 6 mm long, not fused with each other,

bracts absent; pedicels pale green, 0.5-2.5 cm long, pubescent. Calyx green, the two

lips completely divided, ventral tube c. 1 mm long, dorsal tube c. 0.3 mm long, lobes

ligulate to lanceolate, 12-15 x 1.7-3 mm, apex acute and slightly thickened, margin

entire or with 1-few minute teeth, succulent at the base, eglandular hairy on both

sides, outside with a dimorphic indumentum of fine and large eglandular hairs, inside

hairy only towards the tip. Corolla 20-25 mm long, tube white outside, limb white

or dark violet-blue with a bright yellow/orange stripe inside tube ventrally, which

can almost circle the throat, tube narrow, more or less curved, abruptly opening up

into spreading lobes, tube glabrous, lobes eglandular pubescent outside, inside with a

A revision of Microchirita in Thailand

237

Fig. 6. Microchirita hypocrateriformis C.Puglisi. A. Unifoliate plants. B. Caulescent plant.

C. Front view of a white flower. D. Side view of the flower. E. Front view of a blue flower. F.

Blue-flowered caulescent plant. A-D from Tetsana, N. et al. 888 ; E, F from Tetsana, N. et al.

834. (Photos: N. Tetsana)

238

Card. Bull. Singapore 69(2) 2017

fringe of fine translucent glandular hairs and glands at the base of the upper lobes, and

glands and short hairs over the yellow ventral stripe; tube 15-17 mm long dorsally,

17-21 mm ventrally, 15-19 mm laterally between lips; upper lobes broadly elliptic,

4.2-5 x 6-7 mm, lateral lobes broadly ovate, 4-9 x 8-9 mm, ventral lobe ovate, 6-9

x 9-12 mm. Stamens arising 1-1.25 cm above the corolla base, filaments straight,

white, glabrous, 2-2.5 mm long, 0.4-0.7 mm wide; anthers white or pale yellow, with

an abundant indumentum of coloured hairs (blue in blue-flowered specimens, brown

in white-flowered specimens) above and dorsally, 2.3 x 1.3 mm, apically joined by a

connective, thecae more or less parallel; lateral staminodes 0.6-1.1 mm long, arising

c. 4.5-8.5 mm above the corolla base, central staminode 0.1-0.7 mm long, arising

5-8.5 mm above the corolla base. Disk annular, slightly lobed, 0.4-1 mm high. Pistil

c. 23 mm long; ovary 0.6-1.4 cm long, c. 1 mm diameter, glabrous in the basal 0.5-3

mm, then densely eglandular hairy; style c. 0.9 cm, sparsely eglandular hairy; stigma

with a weakly bilobed lower lip, densely hairy, 1-1.6 mm long. Fruit green, brown

when ripe, 2-3.7 cm long, c. 0.6 mm diameter, eglandular hairy, straight. Seeds brown,

narrowly elliptic, 0.3-0.4 x c. 0.1 mm.

Distribution. Eastern and Northeastern Thailand.

Habitat. On limestone in mixed deciduous forest.

Provisional IUCN conservation assessment. Endangered EN Blab(iii) + B2ab(iii).

The known EOO is < 500 km 2 and the known AOO is only 12 km 2 . Not all of the

known localities are in protected areas and those that are not are subject to disturbance

from visitors.

Additional specimens examined. THAILAND: Chaiyaphum: Khon Sarn, Wat Huang Po, 378

m, 7 Nov 2014, Tetsana, N. et al. 888 (BKF, SING); Khon Sarn, Wat Pa Thum Thep Nimit Doi

Kitchakoot, 484 m, 12 Sep 2014, Tetsana, N. et al. 834 (BKF, SING). Loei: Nong Hin, Suan Sa

Wan, PhaNgam Forest Park, 662 m, 11 Sep 2014, Tetsana, N. et al. 824 (BKF, SING).

Notes. The epithet refers to the narrow corolla tube. Most of the specimens observed

have pure white corollas, a yellow ventral stripe, and brown anther indumentum. The

specimen Tetsana, N. et al 834, instead, has a blue-purple corolla, with similarly

coloured anther indumentum and a yellow ventral stripe. The morphology of the white

and blue forms are otherwise identical, and no intermediate colour variants have been

observed.

The collection from Loei, Tetsana et al. 824, has a longer calyx than the material

from Loei. Further collections are needed to see whether the Loei material differs

taxonomically from Microchirita hypocrateriformis.

11. Microchirita involucrata (Craib) YinZ.Wang, J. Syst. Evol. 49: 60 (2011); Rafidah,

Gard. Bull. Singapore 69: 15 (2017). - Chirita involucrata Craib, Gard. Chron., Ser.

3, 83: 140 (1928); Barnett, FI. Siam. 3: 223 (1962); Wood, Notes Roy. Bot. Gard.

A revision of Microchirita in Thailand

239

Edinburgh 33: 199 (1974); Burtt, Thai Forest Bull., Bot. 29: 88 (2001). - TYPE:

Cult. Hort. Bot. Aberdeen from seeds collected by Kerr ( Kerr 11172), Thailand, Kaw

Tao [Surat Thani, Kao Tao], 30/12/1926 (lectotype ABD [specimen with appended

protologue], designated by Puglisi in Rafidah (2017); isolectotypes ABD [2 sheets]).

(Fig. 7)

Chirita involucellata Craib, Gard. Chron., Ser. 3, 83: 140 (1928). - TYPE: Cult. Hort.

Bot. Aberd., 17/11/1927, from seeds collected by Kerr, Thailand, [Chumphon], Tako,

8/2/1927 (holotype ABD).

Notes. This species is easily recognisable from the inflorescence, which has two

bracts that are not fused across the axis (compare to Microchirita rupestris where the

bracts are fused together). The flower is narrowly funnel shaped, variously coloured

in shades of purple, and the fruit is narrow, long and more or less curved. In Thailand

three varieties can be distinguished.

Key to the varieties of Microchirita involucrata

la. Bracts 16-45 mm long; corolla 18-21 mm long.2

lb. Bracts to 10 mm long; corolla c. 35 mm long.

. He. M. involucrata var. gigant flora

2a. Anthers hairy.11a . M. involucrata var. involucrata

2b. Anthers glabrous.lib. M. involucrata var. capitis

11a. Microchirita involucrata var. involucrata

Lithophytic or terrestrial, succulent herb to 2 m tall. Stem fleshy, very sparsely hairy

(mixed glandular and eglandular hairs), green or purple-reddish, especially around

the nodes and the younger parts, branching, with internodes 1-7 cm. Leaves opposite,

except for the single basal leaf; petioles 1.5-6 cm long, eglandular hairy; blades mid

to dark green above, greenish white beneath, ovate or lanceolate, 4.1-11 x 2.6-6.8 cm,

1.2-1.9 times as long as wide, base shortly attenuate to rounded, often unequal, apex

more or less broadly acute, eglandular tomentose above, much less so beneath, margin

entire or obscurely denticulate, 5-8 pairs of secondary veins in the opposite leaves,

venation slightly sunken above and raised beneath in fresh material, tertiary venation

inconspicuous. Inflorescences arising from the petiole, close to the axil, consisting of

a main pedunculate inflorescence, which can be more or less compound, and a bud at

its base, axes light green; peduncles sparsely glandular and eglandular hairy, 2-35 mm

long; bracts paired, free, sessile or shortly petiolate, petiole predominantly glandular

hairy, lamina with mixed glandular and eglandular hairs above and beneath, entire to

wealdy serrate, ovate, 1 . 6 ^ 1 . 5 x 1 . 1-2.2 cm; pedicels sparsely hairy, mixed glandular

and eglandular, predominantly glandular, 1-2.5 cm. Calyx light green to maroon,

240

Card. Bull. Singapore 69(2) 2017

Fig. 7. Microchirita involucrata (Craib) Yin Z.Wang var. involucrata. A. Habit and fruit. B.

Front view of the flower. C. Side view of the flower. D. Lateral view. All from Middleton, D.J.

et al. 5391. (Photos: P. Karaket)

tube 0.5-3 mm long, lobes lanceolate, overlapping, 4-7 x 0.7-1.4 mm, apex acute,

predominantly glandular hairy on the outer side, inside glabrescent or with sparse,

fine glandular hairs. Corolla 18-21 mm long, bluish-violet, dark purple at mouth and

dorsally by the anthers, with two purple stripes laterally along the filaments, lobes violet

or light blue, flower occasionally reported to be much paler to off white especially at

mouth, tube narrow at base, straight to bent, gradually broadening, glandular pubescent

outside, especially dorsally between the upper lobes and ventrally below the central

lobe, internally with sparse eglandular hairs on the upper lip and centrally in the lower

lip, tube glabrous inside, lobes not reflexed; tube 12.5-16.5 mm long dorsally, 15-19.5

mm ventrally, 12.5-17 mm laterally between lips; upper lobes broadly elliptic, 2.3-4.5

x 3.1-5 mm, lateral lobes elliptic, 2.2-4.5 x 2.6-7 mm, lower lobe elliptic, 2-5 x 2.9-9

mm. Stamens arising 8-10 mm above the corolla base; filaments white to purple, bent

twice, or almost straight, sometimes swollen in the middle, glabrous, 3.3-4.6 mm

A revision of Microchirita in Thailand

241

long, 0.4-0.7 mm wide, with a small projection at the anther’s insertion; anthers white,

hairy dorsally and on the upper and lower surfaces, hairs sparse to abundant, 1.7-2.5 x

1.3-1.9 mm, apically weakly joined by a thin connective, free in old specimens, thecae

slightly divergent, anthers whitish, apices slightly acuminate; lateral staminodes 0.7-3

mm long, to 0.1 mm wide, arising 8.5-10 mm above the corolla base, hairy at the tip,

central staminode 0.3-0.7 mm long, arising 8-10 mm above the corolla base, often

inconspicuous. Disk annular (some specimens appear to have a partial disk, probably

due to breakage?), margin irregular to lobed, 0.2-0.6 mm high. Pistil 12-20 mm long;

ovary 6.5-10 mm long, glabrous, many ovules; style green, 0.4-0.7 cm, glandular

pubescent; stigma white, broadly bilobed, lobes 0.5-2.5 mm long, 0.4-0.7 mm wide,

elliptic, often glandular hairy outside. Fruit light green, turning brown at maturity, to

10 cm long, glabrous or with persistent style hairs distally, straight or slightly curved.

Seeds brown, broadly elliptic, c. 0.4 x 0.2 mm.

Distribution. Thailand. Vietnam, Cambodia, Malaysia.

Habitat. Lithophyte on different types of rocks, in shaded areas in evergreen forest.

Provisional IUCN conservation assessment. Least Concern LC. This variety is

common and widespread.

Additional specimens examined. Unknown locality: Smith, E. 71 (BK); Cult. 11172 coll.

7/11/1929 (ABD [2 sheets]), Cultivated at RBGE from material grown at Botanische Anstalt

University, Basel, acc. 801771, P12, C13666 (E); Cult Aberdeen “150”, no data (ABD), Cult

Aberdeen “180” (ABD); Near Langkawi, 15 Jan 1916, Annandale, N. 1850 (SING); Kaw Koh

Suwan near Langkawi, 14 Jan 1916, Annandale, N. 1833 (SING); Koh Si Kah, 17 Jan 1916,

Annandale, N. 1589 (SING); Koh Si Kah, 17 Jan 1916, Annandale, N. 1590 (SING).

THAILAND: Ayutthaya: s.l., 8 Jan 2496 [1953], Piyakarnchown, T. 11 (BK). Chonburi: Si

Racha, Chan Ta Then Falls, 300^100 m, 17 Nov 1974 Maxwell, J.F. 74-988 (BK); Sattahip, Si

Chon Tiam, Nong Nooch BG, 6 Sep 2004, Palee, P. 715 (CMUB); Sriracha, 14 Nov 1926, Put,

N. 444 (ABD, BK, BM, K); Sriracha, Nong Nam Kheo, 240 m, 15 Nov 1926, Collins, D.J.

1277 (K); Hoop Bon, Sriracha forest, c. 400 ft, 25 Oct 1927, Collins, D.J. 1694 (K); Si Racha,

Kow Kieo, 200 m, 20 Oct 1975, Maxwell, J.F. 75-1064 (E). Chumphon: 24 Mar 1971, Bogner

434 (E); Cultivated RBGE from Bogner 434, C8252 (E); Sawi, Khao Thalu, 31 Dec 1999,

Wongprasert, T. 9912-04 (BKF); ibid., 5 Dec 1999, Wongprasert, T. 9912-14 (SING); Lamae,

Tham Khao Phu, 25 m, 7 Feb 2005, Williams, K. et al. 1259 (A, BKF, E); Mueang Chumphon,

Wat Tham Khao Khun Krating, 70 m, 12 Jun 2006, Williams, K. et al. 1666 (A, BKF, E); Thun

Tako, Ban Khao Talu, 4 Dec 2002, Koonkhunthod, N. et al. 308 (BKF); Sawi, Khao Thalu, 5

Dec 1999, Wongprasert, T. 9912-20 (BKF); Pa Thieo, Thale Sap, Wat Tham Thale Sap, 50 m,

17 Dec 2006, Pooma, R. et al. 6372 (BKF); Sawi, Khao Khai, Tham Thip Prida San Chang

Len, 70 m, 26 Dec 2006, Pooma, R. et al. 6680 (A, BKF); Bang Son, 10 Sep 1927, Put 1042

(K); Bang Son, 10 Sep 1927, Put, N. 1035 (BM, K); Sapli, 9 Sep 1927, Put, N. 1021 (BK, BM,

K); cultivated in Edinburgh from seeds from Bogner 424, Nov 1971, C8251-71-0977 (E [2

sheets]); Luang Suan, 23 Mar 1971, Bogner 424 (E); Kao Muang, c. 20 m, 11 Jan 1927, Kerr,

A.F.G. 11368 (BK, BM, K). Krabi: Cultivated in Penang BG, flowered in Aug 1897, Kasoom,

Nov 1896, Curtis 3221 (SING [2 sheets]); Khao Phra Bang Wildlife Reserve, 50 m, 5 Oct 1996,

242

Card. Bull. Singapore 69(2) 2017

Bcirfod, A. et al. 43828 (BKF, K); Tham Suea, 50-250 m, 24 Oct 1991, Larsen, K. et al. 42542

(BKF, P); Bang Kram Forest Reserve, 14 Jul 1992, Larsen, K. et al. 43405 (BKF); Khao Phra

Bang Wildlife Reserve, Sa Morakote (Emerald pool), 75 m, 1 Oct 2005, Maxwell, J.F. 05-508

(A, BKF); Khao Panom Bencha National Park, Tham Phung, 250 m, 13 Oct 2002, Palee, P. 559

(CMUB). Nakhon Ratchsima: Lat Bua Kao, Oct 1916, Kloss, C.B. s.n. (K). Nakhon Si

Thammarat: Lan Saka, Khao Luang National Park, Wat Khiriwong, 90 m, 31 Feb 1966,

Hansen, B. & Smitinand, T. 12172 (BKF, SING); Lan Saka, Ka Rom Waterfall, 15 Dec 1972,

Santisuk, T. & Nimanong, B. 293 (BKF, SING); Khiriwong, Suvarnakoses, P. 1809 (BKF);

Thung Song, 28 Aug 1967, Shimizu, T. & Nalampoon, A. T-8255 (BKF); Thung Song, Ban Thi

Wang, Wat Tham Phra Kru, 55 m, 11 Sep 2010, Middleton, D.J. et al. 5391 (BKF, E);

Nopphitam, Tham Lot Cave, 80 m, 13 Feb 2005, Williams, K. et al. 1457 (A, BKF, E); Khao

Luang National Park, 25 Oct 1991, Larsen, K. et al. 42581 (BKF); Si Chon, Si Chon Si Khit,

Waterfall NP, 21 Dec 2006, Pooma, R. et al. 6526 (BKF, E); Khanom, Mu Ko Thale Tai

National Park, Nam Tei Waterfall, 130 m, 18 Feb 2004, Middleton, D.J. et al. 3207 (A, BKF);

Khiriwong, 31 Jan 1966, Suthasorn, S. 879 (BK); Lan Saka, Khao Luang National Park, Wat

Khiriwong, 100-700 m, 17 Jan 1966, Tagawa, M. et al. T-4546 (BKF, E, K); Lan Saka, c. 100

m, 25 Apr 1928, Kerr, A.F.G. 15391 (ABD, BK, K, P); Cultivated in Aberdeen, 14 Nov 1928,

from Kerr 15391 (ABD). Narathiwat: s.L, 100 m, 5 Feb 1961, Smitinand, T. 7119 (BKF, E,

SING); Budo-Sungai Padi National Park, Pajo Falls, 250 m, 27 Dec 1999, W ongprasert, T.

9912-81 (BKF); Bacho, 20 Dec 1968, S.P 16 (BKF, SING); Bacho, 15 Dec 1968, Sangkhachand,

P. 1572 (BK). Pattani: Sai Khao Waterfall, 15 Nov 1968, Smitinand, T. s.n. (BKF); Sai Kao

Waterfall, 100-150 m, 9 Oct 1991, Larsen, K. et al. 42247 (BKF); Sai Kao Waterfall, 50-200

m, 20 Dec 1972, Santisuk, T. & Nimanong, B. 446 (BKF, SING); Koke Po: Sai Khao Falls

Forest Park, 24 Jan 1985, Maxwell, J.F. 85-100 (BKF). Phang Nga: Khao Lak National Park,

21 Nov 2004, Chamchumroon, V. 2116 (BKF, PSU); Mueang Phang Nga, Suan Somdet, 8 Dec

1999, Wongprasert, T. s.n. (BKF, SING); Mueang Phang Nga, Sri Nakharin Park, 18 Nov 2014,

Suddee, S. et al. 4813 (BKF); Takua Pah, Khao Lak-Lam Ru National Park, 6 Nov 2014,

Suddee, S. et al. 4789 (BKF). Phatthalung: Khao Chai Sohn, Pratudong Cave, 50 m, 12 Jan

1987, Maxwell, J.F. 87-23 (BKF, CMU); Khuan Khanum, MakokNuea, Wat Phu Khao Thong,

30 m, 24 Dec 2006, Pooma, R. et al. 6615 (BKF, E); Ban Phot, 100-150 m, 20 Dec 1979,

Shimizu, T. et al. T-27742 (BKF, L); Khao Chai Sohn, 30 m, 23 Dec 2006, Pooma, R. et al. 6580

(BKF, E); Kao Oktalu, c. 200 m, 21 Apr 1928, Kerr, A.F.G. 15349 (BK, K); Cultivated from

Kerr, A.F.G. 15349, coll. 6/11/1928 (ABD [2 sheets]); Kao Hua Tek, 50 m, 2 May 1930, Kerr,

A.F.G. 19454 (K); Khao Oktalu, 120 m, 21 Apr 1938, Kerr, A.F.G. 15776 (K). Phetchaburi:

s.l., c. 10 m, 7 Nov 1926, Kerr, A.F.G. 11064 (ABD, BM, K); Cha Am, Khao Nang Phanthurat

Forest Park, 50 m, 15 Dec 2006, Pooma, R. et al. 6322 (BKF, E). Phuket: Thalang, Khao

Pateaw Wildlife Sanctuary, Ton Sai Waterfall, 13 Dec 1979, Shimizu, T. et al. T-27153 (BKF).

Prachuap Khiri Khan: s.l., 27 Nov 1966, Sangkhachand, B. 1107 (E); Bangsaphan, Phong

Prasat subdistrict, Khao Ma Rong cave, 2 Oct 2001, Pooma, R. et al. 3063 (BKF); Pran Buri,

Khao Sam Roi Yot, 26 Oct 1973, Suthasorn, S. 2722 (BK); Hui Yang Fall, 6 Oct 1930, Put, N.

3234 (ABD [2 sheets], BM [2 sheets], BK, K [3 sheets]); Kui Buri, Khao Daeng, 22 Oct 1973,

Suthasorn, S. 2676 (BK); Hui Yang Fall, 24 Oct 1964, Chermsirivathana, C. 175 (BK [2

sheets]); Kui Buri, Khao Daeng, 9 Nov 1964, Adisai 926 (BK); Pran Buri, Sam Roi Yot, Turn

Pa Ya Na Khon, 30 Aug 2006, Trihoun, P. 3640 (E); Khao Loom Muak, 200 m, 5 Sep 2008,

Middleton, D.J. et al. 4277 (E). Ranong: Kra Buri, Phra Khayang Cave, 2 Nov 2007,

Pattharahirantricin, N. 166 (BKF). Ratchaburi: Tham Chompol Arboretum, 24 Nov 1987,

Smitinand, T. & Santisuk, T. s.n. (BKF); Cultivated in Bangkok from a plant collected in

Ratchaburi, 14 Oct 1924, Kerr, A.F.G. 9321 (BM, K); Cultivated in Bangkok from plant

A revision of Microchirita in Thailand

243

collected in Ratchaburi, 25 Oct 1924, Kerr, A.F.G. 9341 (BM). Satun: Tarutao National Park,

Adang Island, Oct 1979, Congdon, G. 57 (E). Songkhla: s.l., 21 Dec 1978, Hamilton &

Congdon, G. 101 (BKF); 0-10 m, 8 Apr 1928, Kerr, A.F.G. 15101 (BK, BM, K); Tam Ta Lord,

100 m, 25 Nov 1990, Larsen, K. et al. 41711 (BKF); ibidem, 100 m, 25 Nov 1990, Larsen, K.

et al. 41714 (BKF, P); 10 km NE of Boriphat Falls, 28 Aug 1983, Eddie, W.M.M. s.n. (BKF);

Rattapoom, Khao Rak Kiat, 50 m, 9 Dec 1986, Maxwell, J.F. 86-1035 (BKF, CMU); Boriphat

Falls National Park, 20 Dec 1984, Maxwell, J.F. 84-538 (BKF); ibidem, 120-150 m, 18 Dec

1979, Shimizu, T. et al. T-27611 (BKF, L); ibidem, 120-150 m, 18 Dec 1979, Shimizu, T. et al.

T-27591 (BKF, L); Boripath waterfall, 9 Nov 1990, Larsen, K. et al. 41239 (BKF); Rattaphum,

21 Dec 1965, Youngboonkird, U. 263 (BK); Sabahoy, Bahoi, Ban Ranuea, 200 m, 1 Nov 1998,

Maknoi, C. 26 (QBG); Kao Changlan, c. 50 m, 24 Jul 1928, Kerr, A.F.G. 15894 (BM, K). Surat

Thani: s.l., 5 Dec 1975, Phraphat, D. 103 (BKF); Tai Rom Yen National Park, 100 m, 18 Dec

2006, Pooma, R. et al. 6409 (BKF, E); Tai Rom Yen National Park, Tham Khamin, 100 m, 18

Dec 2006, Pooma, R. et al. 6419 (A, BKF); Viphavadee, Ta Kuk Tai, Viphavadee Waterfall,

Ban Wan Phak Wan 16 Feb 2012, Sirimongkol, S. et al. 290 (BKF); Ko Samui, Namuang Falls,

50 m, 4 Feb 1987, Maxwell, J.F. 87-145 (BKF, CMU); Kaw Samui, c. 100 m, 10 Apr 1927,

Kerr, A.F.G. 12591 (BM, K); Kaw Samui, c. 100 m, 8 Apr 1927, Kerr, A.F.G. s.n. (BM); Koh

Samui, 50-100 m, 3 Dec 1974, Geesink, R., Hiepko, P. & Phengklai, C. 7738 (L); Khao Sok,

26 Dec 1976, Santisuk, T. 856 (BKF); Kaw Ngua Talam, 7 Apr 1927, Kerr, A.F.G. 12935 (K);

Kaw Tao, c. 30 m, 30 Dec 1926, Kerr, A.F.G. 11172 (ABD, BM, K); Kaw Tao, c. 10 m, 14 Apr

1927, Kerr, A.F.G. 11172A (ABD, BM, K); Cult, from Kerr, A.F.G. 11172 coll. 7/11/1929 (ABD

[2 sheets]); Cult, from Kerr, A.F.G. 11172 coll. 6/11/1928 (ABD [2 sheets]); Cult, from Kerr,

A.F.G. 11172 no coll, date (ABD [2 sheets]); Ban Kawp Kep, c. 100 m, 5 Aug 1927, Kerr,

A.F.G. 13179 (ABD, BK, BM, K); Ban Khan Thuli, 7 Sep 1931, Put, N. 4130 (BM, K); Kau

Hoa Kwai, Tassateng, 3 Jan 1935, Seidenfaden 2134 (SING). Trang: Huay Yot, Wat Tham Iso,

130 m, 6 Mar 2006, Middleton, D.J. et al. 4099 (A, BKF, E); ibidem, 101 m, 9 Sep 2008,

Middleton, D.J. et al. 4423 (BKF, E); Khao Pina, 16 Nov 1990, Larsen, K. et al. 41475 (BKF);

Na Yong, Na Muen Si, Wat Hua Khao, Santikhunakorn ladder, 60 m, 24 Dec 2006, Pooma, R.

et al. 6598 (BKF, E); Khao Pina, 150 m, 23 Oct 1991, Larsen, K. et al. 42505 (BKF); Khao

Chong, 5 Mar 1969, Sangkhachand, P. 1758 (BK). Yala: s.l., 31 Jan 1931, Put, N. 3685 (ABD

[2 sheets], BM, K); Wat Tam, 50 m, 10 Dec 1972, Santisuk, T. & Nimanong, B. 367 (BKF (3),

SING); Than To, Bang Lang National Park, 80 m, 8 Feb 2004, Middleton, D.J. et al. 2784 (A,

BKF); Bannang Sata, Khuean Bang Lang, Ban Santi, 150 m, 28 Oct 2005, Poopath, M. 403

(BKF, E); Bannang Sata, Khuean Bang Lang, Ban Santi, 150 m, 1 Sep 2005, Poopath, M. 350

(BKF, E); Than To, Bang Lang National Park, 200 m, 8 Dec 2004, Palee, P. 681 (A, E).

lib. Microchirita involucrata var. capitis (Craib) C.Puglisi, stat. nov. - Chirita

capitis Craib, Bull. Misc. Inform. Kew 173 (1930); Barnett, FI. Siam. 3: 223 (1962). -

TYPE: “Described from plants raised in Aberdeen from seed collected in Bangkok by

Dr. A.F.G. Kerr. Flowered in November 1928”, the specimen is labelled Kerr “179”,

collected on 6/11/1928, cultivated in Aberdeen from Bangkok seed (lectotype ABD,

designated by Wood (1974: 199); isolectotype K [K000545592]).

Distribution. Central and Southwestern Thailand.

Habitat. On rocks and walls.

244

Card. Bull. Singapore 69(2) 2017

Provisional IUCN conservation assessment. Data Deficient DD. The known EOO and

AOO for this variety would qualify it as Endangered if there were associated threats.

However, most of the collections are old and distributions from the areas in Central

Thailand cannot be verified.

Additional specimens examined. Unknown locality: Cult Aberdeen, 9 Nov 1929, Kerr, A.F.G.

“241 ” (ABD [2 sheets]). THAILAND: Ayutthaya: s.l., 21 Dec 1936, Kerr, A.F.G. 19891 (BK,

BM, K). Bangkok: under 5 m, 3 Jan 1923, Kerr, A.F.G. 6717 (ABD, K, BM); 29 Nov 1925,

Kerr, A.F.G. s.n. (ABD, BK, BM, K); 16 Oct 1919, Kerr, A.F.G. 3820 (ABD, BM, K); 9 Oct

1920, Kerr, A.F.G. 3820A (ABD, BM, K); 5 Dec 1920, Marcan, A. 482 (BM, K); Wat Jum,

6 Oct 1921, Kerr, A.F.G. s.n. (BM). Lop Buri: Ayuthia, 22 Oct 1926, Smith, H.M. s.n. (BM).

Prachuap Khiri Khan: Khao Sam Roi Yot National Park, 3 Dec 1928, Put, N. 2521 (K); Pran

Buri, Sam Roi Yot, 7 May 1974, Larsen, K. & Larsen, S.S. 33698 (AAU).

Notes. This variety was first described by Craib (1930) as a species but was included

in synonymy of Microchirita involucrata by Wood (1974). It is here reinstated at the

rank of variety. The only distinguishing feature is the lack of an indumentum on the

anthers. Given the otherwise identical morphology and the overlapping distribution, a

varietal status is preferred for this taxon.

11c. Microchirita involucrata var. gigantiflora C.Puglisi, var. nov.

This variety differs from Microchirita involucrata var. involucrata in the longer

flower (c. 35 mm vs. c. 20 mm) and the much smaller bracts (< 1 cm long in var.

gigantiflora , > 1.6 cm in var. involucrata). - TYPE: Thailand, Nakhon Si Thammarat,

Kha Nom, Khuan Thong, Khao Wang Thong Cave, 50 m, 20 December 2006,

Pooma, R., Phattarahirankanok, K. & Sirimongkol, S. 6472 (holotype BKF; isotype

E [E00311111]).

Distribution. Peninsular Thailand (Nakhon Si Thammarat).

Habitat. On limestone in evergreen forest.

Provisional IUCN conservation assessment. Data Deficient DD. This variety is only

known from the type collection and its distribution and potential threats to it are

unknown.

Notes. The floral characters match Microchirita involucrata var. involucrata, including

key features such as the glabrous ovary and the loosely coherent anthers, except that

the corolla parts are proportionately larger.

12. Microchirita karaketii D.J.Middleton & Triboun, Thai Forest Bull., Bot. 41:

17 (2013). - TYPE: Thailand, Chiang Mai, Chiang Dao District, Doi Chiang Dao

Wildlife Sanctuary, Tam Pak Piang, 530 m alt., 20 September 2008, Middleton, D.J.,

A revision of Microchirita in Thailand

245

Karaket, R, Triboun, P., Kawatkul, U. & Meeboonya, R. 4526 (holotype BKF; isotypes

E [E00629480], P [P00966764], QBG). (Fig. 8)

Herb to 60 cm tall with elongated stem runners. Habit caulescent, internodes c. 8

cm (only one seen). Stems succulent, green, sometimes tinged with reddish, glabrous

or glabrescent; branches sometimes arising from the petioles. Leaves opposite, apart

from the basal leaf; petioles 0.5-2 cm long, green, sparsely eglandular hispid; blades

mid to dark green above, whitish green beneath, ovate, 5.1-25 x 3.5-19 cm, 1.6-2.6

times as long as wide, base cordate, apex acuminate, sparsely eglandular hairy above

and beneath, ciliate along the margin, margin entire, 7-15 pairs of secondary veins,

steeply ascending, venation sunken above and raised beneath in fresh material, flat in

dry specimens, tertiary venation laxly reticulate. Inflorescences cristate, peduncles

reduced or emerging to 10 mm long and sometimes fused together; bracts absent;

pedicels pale green, 4.5-10 mm long, glabrescent or very sparsely hairy. Calyx green,

bilabiate, lower lobes completely divided, upper lobes joined for 1-5 mm, lobes

narrowly lanceolate, 1-6 x 0.8-1.6 mm, apex acuminate and slightly thickened,

margin entire, sparsely hairy especially at the tip. Corolla 12-18 mm long, white

with a yellow stripe ventrally inside and violet patches at its sides, lobes white, tube

narrow, curved, progressively widening, upper lobes spreading, eglandular hairy

outside, glabrous inside; tube 9-13 mm long dorsally, 9.3-14 mm ventrally, 7.8-14

mm laterally between lips; lobes broadly orbicular to elliptic, apices rounded, upper

lobes 2.7-3.5 x 2.8^1.7 mm, lateral lobes 2.8-5 x 4-5.8 mm, ventral lobe 3-4.7 x

3.8-5.5 mm. Stamens arising 3.3-6.5 mm above the corolla base, filaments straight,

glabrous, 2.8-3.5 mm long, c. 0.3 mm wide; anthers anthers white or pale yellow, with

a sparse long indumentum by the insertion, 1.4-2 x 0.8-1.2 mm, apically joined by

a connective, thecae divergent; staminodes arising c. 3 mm above corolla base, c. 0.3

mm long. Disk absent or a ventral half ring 0.6-0.7 mm high. Pistil c. 15 mm long;

ovary 3.5-5 mm long, c. 1 mm diameter, papillose or with sparse hairs in the top half;

style 6-7 mm long, pubescent; stigma chiritoid, c. 1.2 mm long. Immature fruit green,

to 5 cm long, c. 1.5 mm diameter, glabrous, straight or slightly curved. Seeds brown,

narrowly elliptic, acuminate, 0.4-0.5 x c. 0.2 mm.

Distribution. Northern Thailand. Myanmar.

Habitat. On limestone in mixed deciduous forest.

Provisional IUCN conservation assessment. Vulnerable VU Blab(iii). The EOO is

around 8,000 km 2 , it is known only from fewer than 10 populations, and some of the

known localities are not in protected areas and subject to disturbance. The assessment

will have to be reviewed once the distribution in Myanmar is better known.

Additional specimens examined. THAILAND: Chiang Mai: Chiang Dao, road to Wiang

Haeng, 610 m, 21 Sep 2008, Middleton, D.J. et al. 4536 (BKF, E); Chiang Dao, Daan Pha

Woak, 740 m, 20 Aug 1999, Watthana, S. et al. 559 (QBG); Chiang Dao, Kio Phawok border

246

Card. Bull. Singapore 69(2) 2017

Fig. 8. Microchirita karaketii D.J.Middleton & Triboun. A. Fruit. B. Habit. C. Side view of

the flower. D. Front view of the flower. A, C, D from Middleton, D.J. et al. 4526\ B from

Middleton, D.J. et al. 4536. (Photos: A-C, D.J. Middleton; D, P. Karaket)

checkpoint, 750 m, 30 Sep 2009, Middleton, D.J. et al. 5017 (E). Mae Hong Son: Phang

Mapha, Viewpoint, 867 m, 12 Sep 2013, Lakoet, C. 400 (QBG).

Notes. The description provided is a slightly updated version of the original description

by Middleton & Triboun (2013). Microchirita karaketii is recognisable from its colour

pattern (white corolla, with a ventral yellow line and lateral violet markings). Other

species with a similar colour pattern are Microchirita huppatatensis and M. tubulosa,

which, however, differ greatly in the shape of the corolla and the anther indumentum.

Microchirita karaketii is otherwise remarkably similar to M. bimaculata, which is

bright yellow with dark brownish markings. Although the two species are sympatric

in northern Thailand, no intermediate colour forms have been observed. Microchirita

karaketii has flowers smaller than M. bimaculata , and its dry leaves do not acquire the

yellowish hue observed in M. bimaculata.

A revision of Microchirita in Thailand

247

13. Microchirita lilacina C.Puglisi, Kew Bull. 71(l)-2: 5 (2016). - TYPE: Thailand,

Tak, Umphang, 504 m, 15 October 2014, Middleton, D.J., Hemrat, C., Karaket, P,

Puglisi, C. & Suddee, S. 5704 (holotype BKF; isotypes AAU, E [E00663028], K,

QBG, SING). (Fig. 9)

Herb to 40 cm tall. Stems fleshy, red at base, otherwise green, with pale green hairs,

which are lost in dried specimens. Leaves opposite, with the exception of the single

one or two basal leaves, fleshy (very thin when dry); petioles 0.3-2.5 cm long,

glabrous; blades pale to mid green above, paler beneath, ovate, 2.5-18 x 1.4-13 cm,

1.3-1.8 times as long as wide, base cordate, apex acuminate or rarely acute, densely

tomentose above, sparsely so beneath, margin entire, 6-10 pairs of secondary veins in

the opposite leaves, 10-15 in the basal leaf, venation raised beneath in fresh material,

tertiary venation finely and irregularly reticulate, visible only beneath. Inflorescence

epiphyllous, 2-15-flowered, cristate; peduncles reduced or shortly emerging and fused

together; bracts absent; pedicels green, 0.2-1.5 cm long, glabrous or sparsely hairy.

Calyx pale green, bilabiate, tube 0.5-1.5 mm long, lobes narrowly lanceolate, 6.5-7

x 0.8-1.2 mm, with the central upper lobe (alternate to the upper corolla lobes) c.

1 mm shorter than the other lobes, membranous along the margins, apex narrowly

obtuse, outside with scattered eglandular hairs along the midrib, inside with a minute

indumentum of sessile glands. Corolla 8-12 mm long, markedly variable even within

populations, tube whitish pale lilac, with a yellow stripe ventrally, lobes very pale

lilac, tube with a narrow basal portion 3-4 mm long, c. 2.5 mm diam., then abruptly

broadening into a campanulate tube, pubescent with very fine eglandular hairs outside

except at base, inside glabrous, the upper lobes with sparse stalked glands, especially

abundant towards the centre of the upper lip, the lower minutely papillose; tube 11-13

mm long dorsally, 13-14 mm ventrally, 10.5-12.5 mm laterally between lips, upper

lobes elliptic, 3-3.5 x c. 5 mm, lower lateral lobes elliptic, 3-3.5 x 5-6.5 mm, lower

central lobe rounded, 4-4.5 x 4.5-5 mm. Stamens arising 4.5-5 mm above the corolla

base, filaments glabrous, 3.5-4.5 mm long, slightly twisted; anthers cream-white

with a dark spot beside the attachment of the filament, glabrous, 1.7-3 x 0.6-1.5 mm,

apically joined by a connective, thecae divergent and apiculate; lateral staminodes

1.5-3 mm long, arising 3-3.7 mm above the corolla base, central staminode c. 1.7 mm

long, arising c. 2.8 mm above corolla base. Disk a ventral lobe, 0.3-0.4 mm high, with

irregular margin. Pistil 12-16 mm long; ovary 6-11 mm long, with scattered glands

for 1/2-2/3 and then distally with fewer glands and a dense eglandular pubescence;

style 5-6 mm long, curved downwards, hairy as the upper part of the ovary; stigma

with pronounced lobes, lobes 1-1.4 mm long, 0.6 - 0.8 mm wide, narrowly elliptic,

glabrous outside, papillose inside. Fruit green, 2-6 cm long, 0.8-1 mm diam., glabrous

at the base, sometimes with indumentum terminally, straight or curved. Seeds brown,

elliptic, 0.4-0.5 x c. 0.2 mm.

Distribution. Southwestern and Northern Thailand (Kanchanaburi, Tak).

Habitat. On limestone.

248

Card. Bull. Singapore 69(2) 2017

Fig. 9. Microchirita lilacina C.Puglisi. A. Unifoliate habit. B. Caulescent habit. C. Cristate

inflorescence, fruit and side view of the flower. D. Front view of the flower. A from Middleton,

D.J. et al. 5704", B-D from Middleton, D.J. et al. 5699. (Photos: P. Karaket)

Provisional IUCN conservation assessment. Vulnerable VU Blab(iii). Puglisi et al.

(2016) assessed this species as Endangered but since that publication several new

localities have been discovered which have expanded the EOO to around <20,000

km 2 . This new assessment recognises that the threats to the habitat of the species still

exist but that its wider distribution reduces the overall threat.

Additional specimens examined. THAILAND: Kanchanaburi: Sangkhla Buri, Tham Sukho,

180 m, 17 Dec 2009, Pooma, R. et al. 7433 (BKF); Sangkhla Buri, c. 120 m, 29 Aug 1999,

Chayamarit, K. etal. 1831 (BKF [2 sheets]). Mae Hong Son: Muang Mae Hong Son, Mae Surin

National Park, 840 m, 21 Oct 2014, Middleton, D.J. et al. 5820 (E, SING). Tak: Umphang, Pha

Phueang, 990 m, 8 Oct 2014, Suddee, S. et al. 4759 (BKF); Umphang, Doi Hua Mot, Suddee,

S. et al. 3312 (BKF); Umphang, Umphang-Maesot road, 500 m, 19 Oct 2013, Pooma, R. et al.

7894 (BKF); Umphang Wildlife Sanctuary, Doi Hua Mot area, 900 m, 18 Oct 2013, Pooma,

A revision of Microchirita in Thailand

249

R. et al. 7886 (BKF, E); Umphang-Maesod road, 500 m, 24 Apr 2004, Pooma, R. et al. 4643

(BKF); Umphang, Ban Wa Khrue Kho, 543 m, 15 Oct 2014, Middleton, D.J. et al. 5699 (AAU,

BKF, E, K, QBG, SING); Umphang, Doi Hua Mot Wildlife Sanctuary, 915 m, 17 Oct 2014,

Middleton, D.J. et al. 5763 (AAU, E, K, QBG, SING); Tah Song Yang District, Tham Usu, 500

m, 10 Sep 2009, Middleton, D.J. & Triboun, P. 4842 (BKF, E, P).

Note. This species is recognisable by the pale lilac corolla and the glabrous anthers.

14. Microchirita limbata C.Puglisi, sp. nov.

Species characterised by the tubular corolla with white tube and blue lobes, and by

the widespread glandular indumentum. It is most similar to Microchirita albocyanea

C.Puglisi in the overall shape and colour of the corolla, but differs in the smaller

flowers and in having a glandular indumentum. - TYPE: Thailand, Chaiyaphum,

Khon San, Wat Tham Huang Po, 443 m, 19 October 2015, Suddee, S., Keiwbang, W.,

Hemrat, C. 4968 (holotype BKF; isotype SING). (Fig. 10)

Herb to 80 cm tall. Habit caulescent, internodes 1-10 cm. Stems succulent, glandular

hairy; branches sometimes arising from the petiole of the basal leaf. Leaves opposite,

apart from the basal leaf; petioles 0.2-1.7 cm long, glandular hairy; blades mid green

above, much paler beneath, lanceolate to elliptic, 3.1-13 x 0.6-5.2 cm, 1.6-5.2 times

as long as wide, base shortly attenuate to acute, apex acute to acuminate, eglandular

pubescent above and beneath, ciliate along the margin with glandular and eglandular

hairs, margin entire or obscurely denticulate, 4-14 pairs of secondary veins, venation

slightly sunken above and raised beneath in fresh material, flat in dry specimens, tertiary

venation seldom visible in dry specimens. Inflorescences cristate, l-7(-12)-flowered;

peduncles reduced or 1-7 mm long, fused with each other, glandular pubescent;

bracts absent; pedicels pale green to green, 0.2-1.2 cm long, glandular pubescent.

Calyx pale green, succulent at the base, tube c. 0.3 mm long, lobes lanceolate, 4.5-8

x 0.7-1.5 mm, apex acuminate and thickened, margin entire, with mixed glandular

and eglandular hairs on both sides, sparse inside, margin ciliate. Corolla 14-16 mm,

tube white, throat white with a yellow patch at the base of the filaments, lobes purple-

blue, tube narrow at the base, then becoming campanulate, only slightly curved, lobes

almost equal, eglandular hairy outside except at base, densely glandular inside; tube

11-12.5 mm long dorsally, 11-12 mm ventrally, 11.5-13.5 mm laterally between lips;

lobes obtuse (broadly elliptic becoming triangular with a broad apex) with irregular

margin, upper lobes 2-3 x 3.7-4.8 mm, lateral lobes 2.5-4 x 3.5-6.5 mm, ventral

lobe 3.5-4 x 2.7-7 mm. Stamens arising 4-5 mm above the corolla base, filaments

geniculate near the base, green-yellow, glabrous, 2-3 mm long, 0.3-0.4 mm wide;

anthers white, with white to brown hairs throughout except for the apices, 1.9-2.5

x 2.5-2.6 mm, apically coherent but not joined by a connective, or at most with a

very weak and short ligature, thecae parallel or slightly divergent; lateral staminodes

0.3-0.9 mm long, arising 3-3.8 mm above the corolla base, central staminode 0.2-0.3

mm long, arising 3-3.6 mm above the corolla base. Disk annular, margin irregular,

250

Card. Bull. Singapore 69(2) 2017

Fig. 10. Microchirita limbata C.Puglisi. A. Habit. B. Front view of the flower. C. Fruits. D.

Lateral view of the flower. A, D from Tetsana, N. et al. 883; B, C from Suddee, S. et al. 4968.

(Photos: A, D, N. Tetsana; B, C, S. Suddee)

0.1-0.2 mm high. Pistil c. 10.5 mm long; ovary 3.5-5 mm long, c. 1.1 mm diameter,

glabrous at the base, then densely tomentose with glandular and eglandular hairs; style

and stigma with the same indumentum observed on the ovary; style c. 4.5-5.5 mm;

stigma bilobed, lobes elliptic, 0.5-1 mm. Fruit green, 1-4.5 cm long, 0.7-1.2 mm

diameter, densely glandular hairy, straight or curved. Seeds brown, narrowly elliptic,

mucronate, 0.3-0.4 x 0.1-0.2 mm.

Distribution. Northeastern and Eastern Thailand.

Habitat. On limestone in mixed deciduous forest.

Provisional IUCN conservation assessment. Endangered EN Blab(iii) + B2ab(iii).

The EOO and AOO are within the threshold for the Endangered status and not all of

A revision of Microchirita in Thailand

251

the localities are in protected areas with the associated threats to limestone habitats

that entails.

Additional specimens examined. THAILAND: Khon Kaen: Sum Pak Nam, Phu Pha Man

National Park, Chum Phae, 377 m, 21 Oct 2007, Norsangsri, M. & Lakoet, C. 2789 (E, QBG);

Chum Phae, Phu Pha Man National Park, Tham Phaphung section, 25 Oct 2007, Norsangsri,

M. & Lakoet, C. 2811 (QBG). Loei: Phu Kra Dueng, 504 m, 6 Nov 2014, Tetsana, N. et al.

883 (BKF, SING); Phukradung National Park, Sohme Krae area, 1000 m, 4 Oct 2003, Palee,

P. 636 (CMUB).

Notes. The epithet refers to the contrast between the colour of the corolla tube and the

lobes. The specimens from Khon Kaen have denser inflorescences (up to 12 flowers)

and a denser indumentum, but are otherwise alike.

15. Microchirita luteola C.Puglisi, sp. nov.

Similar to Microchirita tubulosa (Craib) A.Weber & D.J.Middleton but differs in not

having spots inside the lateral corolla lobes, having an entire disk (usually dorsally cleft

in M. tubulosa), glandular indumentum on the stems (vs. eglandular), and acuminate

calyx lobes (vs. usually acute, more rarely slightly acuminate). It is also similar to

Microchirita marcanii in the shape of the corolla, but differs in the mixed eglandular

and glandular indumentum on many plant parts (eglandular only in M. marcanii

(Craib) A.Weber & D.J.Middleton) and the corolla colour pattern (light yellow corolla

with a yellow stripe vs. orange corolla with lateral purple spots). Finally, it differs from

Microchirita elphinstonia (Craib) A.Weber & D.J.Middleton in having a glandular

indumentum and in the larger and much paler yellow corolla. - TYPE: Loei, Nong

Hin, Suan Sa Wan, Pha Ngam Forest Park, 662 m, 11 September 2014, Tetsana, N. et

al. 829 (holotype BKF; isotype SING). (Fig. 11A-C)

Caulescent herb to 50 cm tall, internodes 2.5-10 cm. Stems, petioles, pedicels and

peduncles succulent, green and often tinged with reddish-brown, glandular and

eglandular hispid; branches sometimes arising from the petioles of the basal leaf as

well as the opposite leaves. Leaves opposite, apart from the basal leaf; petioles 0.1-1.5

cm long; blades pale to mid green above, whitish green beneath, ovate, elliptic or

obovate, 3.8-8 x 1.5-5.8 cm, 1.2-3.1 times as long as wide, base shortly attenuate to

cordate, apex acute to acuminate, eglandular strigose above, hispid beneath, ciliate

along the margin with eglandular hairs, margin entire or obscurely denticulate,

7-17+ (leaf incomplete) pairs of secondary veins, venation slightly sunken above

and raised beneath in fresh material, flat in dry specimens, tertiary venation visible

in dry specimens. Inflorescences cristate, 1^1-flowered; peduncles 4-10 mm long,

shortly fused with each other; bracts absent; pedicels pale green to reddish brown,

3-25 mm long. Calyx reddish brown at the base, green apically, succulent at the base,

tube c. 1.6 mm long, lobes imbricate at base, lanceolate, 14-17 x 3-3.5 mm, apex

acuminate, margin entire, outside with short, patent, mixed glandular and eglandular

252

Card. Bull. Singapore 69(2) 2017

hairs, inside glabrous except at the tip, margin ciliate at the tip. Corolla c. 35 mm long,

tube white at the base, turning pale yellow distally, base of ventral lobe with a yellow

stripe, lobes pale yellow at the base, paler towards the margin, tube narrow at the base,

curved basally and gradually broadening, mouth broad and slightly gibbous laterally

and ventrally, base of the tube, upper lobes and ventral lobe glabrous, the rest of the

corolla externally covered in short mixed glandular and eglandular hairs, internally

with sparse glandular hairs, forming a denser patch at the base of the upper lobes, lobes

slightly papillose; tube c. 23 mm long dorsally, c. 27.5 mm ventrally, c. 24 mm laterally

between lips; lobes broadly elliptic, upper lobes c. 5.5 x 10 mm, lateral lobes c. 6 x

8.5 mm, ventral lobe c. 8.5 x 10.5 mm. Stamens arising c. 17 mm above the corolla

base, filaments strongly geniculate, with a basal, thinner segment c. 2 x 0.7 mm, and

an apical, thicker segment c. 4.5 x 1.6 mm, pale yellow, glabrous; anthers white, with

white hairs dorsally, c. 3.5 x 2.5 mm, apically joined by a connective, thecae parallel or

slightly divergent; lateral staminodes c. 0.7 mm long, arising c. 9 mm above the corolla

base, central staminode c. 0.5 mm long, arising c. 9 mm above the corolla base. Disk

annular, margin entire, 1 mm high, c. 0.5 mm thick (exceptionally thick). Pistil c. 29

mm long; ovary c. 10 mm long, c. 1.6 mm diameter, lower 1/3 glabrous, then densely

tomentose with eglandular and perhaps glandular hairs; style c. 18 mm long, with the

same indumentum as ovary; stigma bilobed, lobes elliptic, 1 mm. Fruit green, 7-10.5

cm long, 0.5-2.5 mm diameter, densely hairy, straight or slightly curved. Seeds light

brown, narrowly elliptic, with irregular surface, 0.3 x 0.1 mm.

Distribution. Northeastern Thailand.

Habitat. On limestone in evergreen forest.

Provisional IUCN conservation assessment. Endangered EN B2ab(iii). This species

has an AOO of around 20 km 2 and is only known from a small number of collections,

not all of which are in protected areas.

Additional specimens examined. THAILAND: Kalasin: Somdat, Pu Parn, 24 Oct 1975,

Sutharson, S. & Sangkhachand, P. 3450 (BKF). Loei: Nong Hin, Ban Puen Phue, c. 640 m, 22

Sep 2007, Thitimetharoch, T. et al. 797 (BKF); Phu Luang, 22 Aug 1966, Phusomsaeng, S. &

Bunchuai, K. 11 (BKF, K, L); Phu Krading, Sam Khae, c. 1050 m, 16 Oct 1954, Smitinand, T.

2026 (BKF).

Notes. This species is named after its pale yellow corolla.

16. Microchirita marcanii (Craib) A.Weber & D.J.Middleton, Taxon 60: 778 (2011).

- Chirita marcanii Craib, Bull. Misc. Inform. Kew 171 (1926); Barnett, FI. Siam.

3: 226 (1962); Wood, Notes Roy. Bot. Gard. Edinburgh 33: 193 (1974); Burtt, Thai

Forest Bull., Bot. 29: 88 (2001). - TYPE: Thailand, Saraburi, Muak Lek, c. 250 m,

10 November 1924, Marcan 1872 (lectotype ABD, designated by Wood (1974: 193);

isolectotypes K (2)). (Fig. 11D-F)

A revision of Microchirita in Thailand

253

Fig. 11. Microchirita luteola C.Puglisi. A. Habit. B. Front view of the flower. C. Side view

of the flower. All from Tetsana, N. et al. 829. Microchirita marcanii (Craib) A.Weber &

D.J.Middleton. D. Flowers. E. Side view of the flower. F. Front view of the flower. All from

RBG Edinburgh accession number 20121420. (Photos: A-C, N. Tetsana; D-F, D.J. Middleton)

254

Card. Bull. Singapore 69(2) 2017

Caulescent herb to 60 cm tall, internodes 4-13 cm long, branches sometimes arising

from the petiole of the basal cotyledon. Stem fleshy, pale green with purplish base,

sparsely eglandular hairy. Leaves opposite, apart from the basal leaf; petioles pale

green, 7-14 mm long; blades mid green above, paler beneath, ovate, 6.2-24 x 3.2-12.6

cm, 1.7-2 times as long as wide, base cordate, apex acute to obtuse, eglandular hairy

above and beneath, margin ciliate, margin entire to obscurely toothed, 8-15 pairs of

secondary veins, venation sunken above and raised beneath in fresh material, flat in dry

specimens, tertiary venation laxly reticulate. Inflorescences cristate, 10-16-flowered,

all floral axes sparsely to densely eglandular hairy; peduncles 2-13 mm long, shortly

fused with each other; bracts absent, except for first inflorescence arising from the

petiole of each leaf, observed only in cultivated material; pedicels pale green to reddish

brown, 3-26 mm long. Calyx actinomorphic, green, lobes succulent along the midrib,

tube 1.3-3 mm, lobes imbricate, narrowly ovate, 7-18 x 2.9-3.6 mm, apex acute,

margin entire, outside with eglandular hairs along the midrib, tip and margin, inside

with sessile glands. Corolla 32-45 mm long, tube pale green at the base, then becoming

dark orange, throat dark red with lateral purple spots, tube narrow at base, curved,

then becoming broad, upper and lateral lobes spreading, tube glabrous at the base,

then with mixed glandular (especially ventrally) and eglandular (especially dorsally)

hairs, lobes papillose inside, the rest of the inside of the corolla glabrescent or sparsely

glandular; tube 26-30 mm long dorsally, 29-32 mm ventrally, 22-27 mm between the

lips; lobes with broadly obtuse apex and a minutely crenate margin, upper lobes 6-6.5

x 10 mm, lateral lobes 8 x 12-13 mm, ventral lobe 10-11 x 12 mm. Stamens arising

1.6-1.9 cm above the corolla base (the point where the corolla broadens), filaments

slightly to strongly geniculate, 4.5-7 x 1 mm, pale yellow, glabrous, anthers white,

hairy dorsally, 3-3.5 x 1.5-2.7 mm, apically joined by a connective, thecae divergent;

lateral staminodes 0.5-2 mm long, arising 0.8-1.25 cm above the corolla base, central

staminode 0.3-1 mm long, arising 0.9-1.35 cm above the corolla base. Disk annular,

margin slightly lobed, 0.8-1 mm high. Pistil 27 mm long; ovary c. 1 cm long, 1.7

mm diameter, glabrous in the bottom 1/3—1/2, densely hairy above; style slender, 1-2

cm long, hairy; stigma bilobed, lobes c. 0.9-1 mm, papillose/plumose. Fruit green,

4.5-6.2 cm long, 3.5^1 mm diameter, glabrous at the base, eglandular hairy apically,

slightly curved. Seeds very dark brown, oval, 0.7-0.8 x 0.2-0.4 mm.

Distribution. Central and Eastern Thailand.

Habitat. On limestone.

Provisional 1UCN conservation assessment. Vulnerable VU Blab(iii) + B2ab(iii). This

species has an EOO and an AOO within the threshold to be considered endangered but

is known from 5-10 localities, thereby making the Vulnerable status more appropriate.

Additional specimens seen. Unknown province: 22 Oct. 1929, Kerr, A.F.G. (ABD); no label

data (ABD); Cult. Hort. Bot. Reg. Kew., Nov 1926 “Bot. Mag. t. 9244” (K [2 sheets]); Cult.

Hort. Bot. Reg. Kew. A.D. 196? (K); Cultivated, doubtfully from Mueang Ngao, Lampang,

A revision of Microchirita in Thailand

255

Kerr A.F.G. 20571 (ABD). THAILAND: Nakhon Nayok: Nang Rawng, c. 100 m, 24 Nov

1937, Smitinand, T. 3881 (BKF). Nakhon Ratchasima: Cultivated at RBG Edinburgh from

Middleton, D.J. 5641, accession number 20121420, 21 Nov 2012, Middleton, D.J. 5288 (E,

SING [2 sheets]); Pak Chong, Wat Tham Phrom Machan Thammaram, 480 m, 22 Aug 2012,

Middleton, D.J. et al. 5641 (BKF, E); Pak Chong, Khao Yai National Park, 19 Sep 2002,

Charoenchai, P. & Poompuang, S. 329 (CMUB); Khao Yai National Park, Nam Dohk Lek

Falls, 20 Oct 2003, Palee, P. 646 (E); Khao Yai National Park, 8 Aug 1979, Shimizu, T. et al.

9481 (BM, L). Saraburi: Ban Nawng Bua, 1 Oct 1927, Put, N. 1116 (ABD mixed specimen,

see also M. tubulosa ); Muak Lek, Khao Hin Fon, Pa Chai Badan, 19 Nov 2011, Suparinyo,

P.P. s.n. (BKF [2 sheets]); Muak Lek, 3 Sep 1928, Put, N. 1854 (K); Cult. Hort. Bot. Reg. Kew

from Marcan 1872, 11 Nov 1925, 685.25 (K); Khao Sawng Phi Nawng, c. 400 m, 4 Oct 1963,

Smitinand, T. & Sleumer, H.O. 1348 (BKF, E, K, L).

Notes. This species is easily recognised by the large, orange corolla. It is most similar to

Microchirita tubulosa, which however has a white flower with purple lateral markings

and a ventral yellow line.

In the protologue, Craib (1926) cited the wild collection Marcan 1872 and

a cultivated plant grown in Aberdeen from seeds of this plant and noted that the

description was largely based on the cultivated plant. This would make the cultivated

plant a better choice for lectotypification but we have been unable to find any material

unequivocally vouchered from this living collection in Aberdeen before publication of

the protologue.

17. Microchirita micromusa (B.L.Burtt) A.Weber & D.J.Middleton, Taxon 60: 778

(2011). - Chirita micromusa B.L.Burtt, J. Roy. Hort. Soc. 85: 28 (1960); Barnett, FI.

Siam. 3: 226 (1962); Wood, Notes Roy. Bot. Gard. Edinburgh 33: 194 (1974); Burtt,

Thai Forest Bull., Bot. 29: 88 (2001). - TYPE: Thailand, Nakhon Nayok, cult, in

Montreal Botanic Garden from seeds collected by Raymond & Smitinand, Raymond,

M. ref. 106-59 (holotype E [E00155279]).

Caulescent herb to 30 cm tall, internodes 4-7 cm long. Stems succulent, glabrescent

or sparsely eglandular hairy, branching from the petiole of the basal leaf. Leaves

opposite apart from the basal leaf; petioles to 3 cm long, glabrescent; blades lanceolate

to elliptic, 7-25 x 4.1-12 cm, 1.7-2.5 times as long as wide, base shortly attenuate to

subcordate, apex acute to broadly acuminate, with eglandular hairs above and sparse

hairs beneath, margin entire or minutely serrulate, 8-18+ pairs of secondary veins.

Inflorescences cristate, 4-10-flowered (20 according to the protologue), floral axes

sparsely eglandular hairy to glabrescent; peduncles short and fused with each other;

bracts absent; pedicels straight or curved, 7-16 mm long. Calyx actinomorphic, lobes

divided to base, lanceolate, 7.5-13 x 1.2-1.7 mm apex acute, margin entire, with

minute glands on both sides, and a sparse eglandular indumentum outside on margin

and midrib. Corolla 20-26 mm long, yellow with an orange patch ventrally, narrow at

base then gradually broadening, with eglandular hairs along the upper part of the tube

and glandular hairs inside by the mouth; tube c. 16.5 mm long dorsally, c. 17.5 mm

256

Card. Bull. Singapore 69(2) 2017

laterally, c. 20 mm ventrally; lobes elliptic with rounded apex, upper lobes 3.5—4.5 x

6 mm, lateral lobes 5-6 x 6 mm, lower lobe 5-6 x 5.5 mm. Stamens arising c. 11 mm

above the corolla base, filaments straight, sparsely eglandular hairy, c. 3 mm long, c.

0.3 mm diameter; anthers pale, with a dimorphic indumentum of long purple hairs

pointing upwards and short straight white hairs pointing down, arising from the base

of the anther, anthers c. 1.5 x 1.5 mm, apically joined by a connective, thecae slightly

divergent; staminodes 2, 0.4 mm long, arising 7.5 mm above the corolla base. Disk

annular, slightly lobed, 0.4-0.6 mm tall. Pistil c. 15 mm long; ovary c. 6 mm long,

glabrous or finely glandular at base, eglandular hairy in the upper 1/3; style curved,

c. 6 mm long, hairy; stigma lower lip elongate, 1.2-1.5 mm long, 1 mm wide. Fruit

straight or occasionally slightly curved, c. 5-6 cm long, narrow, with sparse glands

along most its length and eglandular hairs terminally. Seeds not seen.

Distribution. Central and eastern Thailand. Possibly also occurring in Nakhon

Ratchasima (photos from Sukonthip Sirimongkol seen).

Habitat. “On sandy soil in deciduous forest”. Habitat data are missing from most

collections of this species but it is noteworthy that the collection localities appear to

be from areas that do not have karst limestone or a limestone bedrock. This makes it

unusual in the genus.

Provisional IUCN conservation assessment. Data Deficient DD. There are rather few

collections of this species and many of these are more than 40 years old. Further

collections are needed in the Khao Yai and Sisaket areas to set a conservation

assessment.

Additional specimens seen. Unknown province: Cultivated at Cornell University, Ithaca,

originally from Montreal Botanical Garden, 23 Oct 1958, Moore, J.R. 7685 (E); Cult. RBGE,

received from Smitinand, C3797 (E); Cult. RBGE, received from Frankfurt BG in 1964, Sep

1964, C4327 (E). THAILAND: Nakhon Nayok: Suvatabundha, K., s.n. (BK); Muang, Nahng

Rawng falls [Nang Rong Waterfall], 16 Sep 1972, Maxwell, J.F. 72-355 (AAU, BK); ibidem, c.

100 m, 22 Oct 1960, Smitinand, T. 6984 (A, E, K). Sisaket: Kantalak, Panomdongrak Wildlife

Sanctuary, Samrong Kiat Waterfall, 300 m, 21 Oct 2003, Palee, P. 647 (CMU).

Notes. Microchirita micromusa is similar to M. elphinstonia, from which it differs in

the orange patch in the corolla and in the clearly dimorphic anther indumentum. Burtt

named it “micromusa” because M. Raymond described the fruits as “a miniature hand

of bananas” (Burtt, 1960). However, short, curved fruits are rather more characteristic

of another yellow-flowered species, Microchirita bimaculata, than of M. micromusa.

There has been some confusion between the two species, resulting in many specimens

of M. bimaculata from Northern Thailand being misidentified as M. micromusa.

Microchirita micromusa is distributed in Central and Eastern Thailand and is not

known to overlap with the distribution of M. bimaculata.

A revision of Microchirita in Thailand

257

18. Microchirita mollissima (Ridl.) A.Weber & D.J.Middleton, Taxon 60: 778 (2011).

- Chirita mollissima Ridl., J. Linn. Soc., Bot. 32: 517 (1896); Barnett, FI. Siam. 3:

226 (1962); Wood, Notes Roy. Bot. Gard. Edinburgh 33: 188 (1974); Burtt, Thai

Forest Bull., Bot. 29: 88 (2001). - TYPE: Siam [Thailand], Pungah [Phangnga], July

1893, Curtis 2944 (lectotype SING [SING0117733] designated by Wood (1974: 188),

isolectotypeK [K000450489], SING [SING0117731] [SING0117732] [SING0117734]

[SING0117735]). (Fig. 12A-C)

Notes. Microchirita mollissima is easily recognised by its elongated and sessile

or subsessile leaves with a dense white indumentum, and by the nearly axillary

inflorescences. Three varieties can be distinguished in Thailand.

Key to the varieties of Microchirita mollissima

la. Anthers hairy, with a small projection at the filament insertion.2

lb. Anthers glabrous, without a projection at the filament insertion.

.18b. M. mollissima var. glabra

2a. Leaves, calyx and corolla with eglandular indumentum only.

. 18a . M. mollissima var. mollissima

2b. Leaves, calyx and corolla with glandular indumentum.

. 18c. M. mollissima var. glandulophylla

18a. Microchirita mollissima var. mollissima

Caulescent perennial herb to 50 cm tall with elongated stem runners, internodes to c.

2 cm. Stems not fleshy, densely hairy. Leaves opposite, condensed at the stem apex;

petioles inconspicuous or to 2.5 cm long, densely eglandular hairy; blades pale to

dark green above with silvery white soft hairs, grey-green, flushed purple beneath,

lanceolate, 3.5-28 x 0.7-9 cm, 2.2-7.2 times as long as wide, base acute to attenuate,

often unequal, apex acute to acuminate, densely eglandular tomentose above and

beneath, margin entire to irregularly denticulate, 6-14 pairs of secondary veins

in the opposite leaves, venation slightly sunken above and raised beneath in fresh

material, tertiary venation inconspicuous. Inflorescences cristate, peduncles 10-60

mm, not fused to each other, densely eglandular hairy, the first inflorescence is rarely

compound and bracteate. Calyx green, tube c. 0.3 mm long, lobes lanceolate, 5-14 x

1.5^1 mm, apex acute, eglandular hairy on both sides, with additional sparse, minute

glands inside. Corolla 18-33 mm long, blue, pale blue, purple, pink or white, tube

creamy white or greenish white, lobes very pale lilac, dorsal fringe of yellow/orange

glandular, papillose hairs inside, narrow at base (7-11 mm, c. 2.2 mm diameter), then

slightly curved downwards and gradually broadening, lobes not reflexed, eglandular

pubescent outside except at base, especially dorsally between the upper lobes and

ventrally below the central lobe; tube 14-26 mm long dorsally, 20-36 mm ventrally,

258

Card. Bull. Singapore 69(2) 2017

14-28 mm laterally between lips; upper lobes broadly elliptic, 4-7 x 7-11 mm, lower

lateral lobes elliptic, 3-12.5 x 4-13 mm, lower central lobe round-elliptic, 4.5-10.5

x 6.5-17 mm. Stamens arising 7.5-16 mm above the corolla base; filaments white or

pale yellow, glabrous, 7.5-9 mm long, 0.8-1.3 mm wide; anthers white or pale yellow,

hairy dorsally, 3.5-4.5 x 2-2.5 mm, apically coherent but not joined by a connective,

thecae divergent; lateral staminodes 3.5-4.5 mm long, 0.2 mm wide, arising c. 7.5-16

mm above the corolla base, central staminode 0.6 mm long, arising c. 8 mm above

the corolla base. Disk annular, margin irregular, dark, 0.4-0.5 mm high. Pistil 13

(immature)-28 mm long, eglandular hirsute throughout; ovary 0.7-1.3 cm long, c. 1

mm diameter; style c. 1.2 cm; stigma broadly bilobed, lobes 1.1-2 mm long, 0.9-1.3

mm wide, elliptic. Fruit brown when ripe, 3-7 cm long, 1-1.5 mm diameter, densely

eglandular hairy, straight. Seeds not seen.

Distribution. Peninsular Thailand.

Habitat. On limestone, in shade.

Provisional IUCN conservation assessment. Near Threatened NT. This variety has an

EOO and an AOO within the limits to be considered endangered; it is, however, known

from many localities.

Additional specimens examined. Unknown locality: 1930, Kerr, A.F.G. 240 (L). THAILAND:

Krabi: Nai Chong, Khap Thong Thai, 30 m, 19 Jan 1966, Hansen, B. & Smitinand, T. 11997

(BKF, E, K, SING); Aoluk, Ban Naiyuan Khaek, c. 20 m, 5 Sep 1982, Shimizu, T. etal. T-29113

(BKF); Mueang Krabi, Thambon Krabi Noi, Wat Tham Seua, 85 m, 11 Sep 2008, Middleton,

D.J. et al. 4443 (BKF, E); Cult. RBGE from Middleton, D.J. et al. 4443, Middleton, D.J. 5208

(E); Khao Phanom Bencha National Park, c. 100 m, 13 Sep 1983, Smitinand, T. s.n. (BKF);

between Krabi and Panom Bencha, 0 m, 24 Oct 1991, Larsen, K. et al. 42534 (AAU, BKF);

Muang, near Panom Bencha National Park, Tham Khao Pheung, 60 m, 17 Jun 2006, Williams,

K. et al. 1826 (BKF); Khao Kopi, c. 50 m, 14 Aug 1965, Smitinand, T. 8929 (BKF); Ao Luk,

8 Oct 1970, Charoenphol, C. et al. 3444 (AAU, E); Plai Pray a, Wat Ban Hian, 24 Sep 2010,

Triboun, P. & Sonsupab, B. 4583 (E). Phang Nga: Pungah, 6 Dec 1918, Hanijf & Nur 3880

(SING); Muang, Tham Khao Ngoom, 20-50 m, 7 Sep 1982, Shimizu, T. et al. T-29191 (BKF);

Mueng, Suan Somdej, 11 Dec 1986, Phengklai, C. & Smitinand, T. 6015 (BKF); Tupput, Tham

Saeng Tham Nimit Dharma office, 60 m, 23 Feb 2001, Chayamarit, K. et al. 2655 (BKF);

Mueang, Thamnamphood, c. 50 m, 1 Jan 2000, Wongprasert, T. 001-11 (BKF); Mueang, Sra

Nang Manora Forest Park, 224 m, 17 Nov 2014, Suddee, S. et al. 4800 (BKF); Khao Phang

Nga, 200 m, 27 Feb 1929, Kerr, A.F.G. 17265 (K); Muang Phangnga, Sra Nang Manora Forest

Park, 16 Sep 2010, Middleton, D.J. et al. 5449 (E); Kan Bow Koranee cascade, 9 May 1973,

Geesink, R. & Santisuk, T. 5292 (L). Surat Thani: Phanom, Khlong Phanom National Park,

200 m, 7 Sep 2008, Middleton, D.J. et al. 4361 (BKF, E).

18b. Microchirita mollissima var. glabra C.Puglisi, var. nov.

Differs from the type variety in the lack of an indumentum dorsally on the anthers

and the small projection of the filament at the insertion. - TYPE: Thailand, Krabi,

A revision of Microchirita in Thailand

259

Turn Pra Sat Na Ra Ki Ling, c. 50 m, 29 October 2006, Triboun, P. 3671 (holotype E

[E00428772]).

Distribution. Thailand (Krabi).

Habitat. On limestone.

Provisional IUCN conservation assessment. Data Deficient DD. This variety is only

known from the type collection and its distribution is too poorly known.

Note. The corolla of the type specimen is reported to be pale pink.

18c. Microchirita mollissima var. glandulophylla C.Puglisi, var. nov.

Differs from the type variety by the glandular indumentum on the calyx and corolla,

and the mixed glandular and eglandular indumentum on the leaves. - TYPE: Thailand,

Phang Nga, Ban Tham Thong Lang, c. 50 m, 25 January 1969, Smitinand, T. & Scheller,

R. 10642 (holotype BKF).

Distribution. Thailand (Phang Nga).

Habitat. On limestone rocks.

Provisional IUCN conservation assessment. Endangered EN Blab(iii) + B2ab(iii).

This species has an EOO and AOO of < 20 km 2 and is only known from a small

number of collections from fewer than 5 locations, not all of which are in protected

areas.

Additional specimens examined. THAILAND: Phang Nga: Thap Put, Khao Sam’kob, c. 80 m,

24 Aug 1967, Shimizu, T. et al. T-7903 (BKF); Tham Thong Lang, 50 m, Jun 1971, Cultivated

RBGE, accession number 691872, vouchered as C8246 (E); Muang Phangnga, 29 Sep 2006,

Triboun, P. 3665 (E).

19. Microchirita oculata (Craib) A.Weber & D.J.Middleton, Taxon 60: 778 (2011). -

Chirita oculata Craib, Bull. Misc. Inform. Kew. 174 (1930); Barnett, FI. Siam. 3: 226

(1962); Wood, Notes Roy. Bot. Gard. Edinburgh 33: 194 (1974); Burtt, Thai Forest

Bull., Bot. 29: 88 (2001). - TYPE: “described from a plant which flowered in Aberdeen

in Jul 1928. It was raised from seed of Kerr 9750 which was collected on Kao Sakan,

6 November 1928” (lectotype ABD, designated by Wood (1974: 194); isolectotype E

[EOO 155281]). (Fig. 12D-F)

Caulescent herb to 40 cm tall, internodes 1-11 cm. Stems succulent, green, eglandular

tomentose; branches sometimes arising from the petioles of the basal leaf. Leaves

opposite, apart from the basal leaf; petioles 0.2-1.7 cm long, green, sparsely eglandular

260

Card. Bull. Singapore 69(2) 2017

Fig. 12. Microchirita mollissima (Ridl.) A.Weber & D.J.Middleton var. mollissima. A. Front

view of the flower. B. Habit. C. Side view of the flower. A, B from Middleton, D.J. et al.

5449', C from Middleton, D.J. 5208. Microchirita oculata (Craib) A.Weber & D.J.Middleton.

D. Habit. E. Side view of the flower. F. Front view of the flower. All from Middleton, D.J. et al.

5581. (Photos: A, B, T. Phutthai; C, D.J. Middleton; D-F, P. Triboun)

A revision of Microchirita in Thailand

261

hairy; blades green above, paler beneath, ovate or lanceolate, 4-16 x 1.7-10 cm,

1.4- 2.4 times as long as wide, base obtuse to subcordate, apex acute to acuminate,

dimorphic eglandular indumentum above, more sparsely so beneath, ciliate along the

margin, margin entire to obscurely toothed, 5-14 pairs of secondary veins, steeply

ascending, venation slightly sunken above and raised beneath in fresh material, flat

in dry specimens, tertiary venation lax. Inflorescences cristate, all axes with a dense

indumentum of white eglandular hairs; peduncles reduced or to 2 mm long, not fused

with each other; bracts absent; pedicels pale green, 1-6 mm long. Calyx whitish green,

actinomorphic, tube 0.5 mm, lobes triangular, 11.5-13 x 1.4-1.6 mm wide, apex

acuminate, margin entire, densely eglandular hairy outside, inside hairy only towards

the tip. Corolla 16-20 mm long, bright yellow, with a dark reddish brown ring at the

throat, tube narrow, straight or curved, abruptly widening, lobes spreading, the ventral

prominent, eglandular hairy outside, papillose inside, especially on the lobes, and with

larger glands at the base of the upper lobes; tube 14-17 mm long dorsally, 15-18 mm

ventrally, 14-16 mm laterally between lips; lobes broadly orbicular and imbricate,

apices obtuse to rounded, margin obscurely crenulate, upper lobes 2.5-4.5 x 4.5-

6.5 mm, lateral lobes 4-5 x 5-9 mm, ventral lobe 4-8 x 6-11 mm. Stamens arising

9-10.5 mm above the corolla base; filaments straight, glabrescent or sparsely hairy,

1.5- 2 mm long, 0.2 mm wide; anthers white, with an indumentum of thick, probably

glandular, dark hairs dorsally, 1.3-2 x 1-1.1 mm, apically joined by a connective,

thecae divergent, the apices pointing towards the base of the corolla; staminodes all

inconspicuous, or only the lateral staminodes present, lateral staminodes 1-9 mm long,

arising 2.1-7 mm above the corolla base. Disk a small ventral lobe up to a 3/4 ring,

0.4-0.5 mm high. Pistil c. 12 mm long; ovary c. 5 mm long, 0.7-0.8 mm diameter,

glabrous at the base, densely covered in white hairs apically; style c. 5.5 mm long,

hairy, becoming glabrescent towards the top; stigma glabrous or papillose-plumose, 1

mm long. Fruit 3-4 cm long, slightly curved. Seeds not seen.

Distribution. Eastern Thailand (Sa Kaeo).

Habitat. On limestone.

Provisional IUCN conservation assessment. Endangered EN Blab(iii). This species

has a very small EOO and AOO, being only known from the Khao Chakan District of

Sa Kaeo Province. None of the known localities are in protected areas and are subject

to high human disturbance.

Additional specimens examined. THAILAND: Sa Kaeo: Khao Chakan, Wat Ratanakiri, 100

m, 21 Aug 2012, Middleton, D.J. et al. 5581 (BKF, E, P); Khao Chakan Temple, 88 m, 17 Oct

2010, Staples, G. et al. 1408 (BKF, E, SING); Krabin, Kao Sakan, c. 50 m, 24 Dec 1924, Kerr,

A.F.G. 9750 (ABD, BK, BM, K [2 sheets]).

Notes. This plant is easily recognised by its corolla colour pattern: yellow, with a dark

reddish brown ring at the throat. All other species of Microchirita which have a yellow

or orange corolla have smaller lateral spots at each side of the ventral corolla lobe.

262

Card. Bull. Singapore 69(2) 2017

Another characteristic feature of Microchirita oculata is the dense white indumentum

on the pedicels and calices. Floral dissection of the material available suggest this

species to be strongly protandrous.

20. Microchiritapersonata C.Puglisi, Kew Bull. 71(l)-2: 1 (2016). - TYPE: Thailand,

Uthai Thani, Lan Sak, Huppatat Non Hunting Area, 122 m, 14 October 2014, Middleton,

D.J., Hemrat, C., Karaket, P., Puglisi, C. & Suddee, S. 5688 (holotype BKF; isotypes

AAU, E [E00663026], K, SING). (Fig. 13A-C)

Herb to 35 cm tall. Stems fleshy, pale green, glabrous or sparsely hairy. Leaves

opposite, with the exception of the single basal leaf, fleshy (very thin when dry);

petioles 0.1-0.5 cm long, sparsely hairy; blades pale green above, pale grey-green

beneath, lanceolate, 3.4^10 x 1.7-29 cm, 1.3-2.2 times as long as wide, base usually

rounded obtuse, sometimes slightly cordate or slightly decurrent, apex acuminate to

acute, minutely tomentose above and beneath, margin entire, 6-11 pairs of secondary

veins in the opposite leaves, 8-14 pairs in the basal leaf, venation raised beneath in

fresh material, tertiary venation barely visible and loosely reticulate. Inflorescence

epiphyllous, many-flowered (10 to more than 20 flowers), cristate; peduncles reduced;

bracts absent; pedicels pale green, 5-13 mm long, pubescent. Calyx green, slightly

bilabiate, tube 0.4-0.7 mm long, lobes lanceolate, 4-7 x 0.4-0.6 mm, membranous

along the margins, apex more or less narrowly obtuse, outside eglandular hairy

throughout, inside only towards the apex. Corolla 7-10 mm long, tube greenish white,

upper lip pale green, lower lip white, with a densely papillose yellow marking at base,

tube narrowly tubular, mouth personate due to the raised ventral lip, upper lip strongly

reflexed upwards, tube glabrous at the base, limb densely pubescent outside, densely

papillose inside; tube 4-8 mm long dorsally, 6-9 mm ventrally, 4-7 mm laterally

between lips; upper lobes small and elliptic, 0.7-0.8 x 1-1.3 mm, lower lateral lobes

elliptic, 1.5-1.8 x 2.2-2.3 mm, lower central lobe elliptic, 1.2-1.3 x 1.8-3 mm.

Stamens arising 2.7-3.7 mm above the corolla base, filaments pale green, glabrous,

0.5-1.2 mm long; anthers white, glabrous, 1.1-1.7 x 0.3-0.5 mm, apically joined by

a connective, thecae slightly divergent; lateral staminodes absent or to 0.2 mm long,

arising c. 1.7 mm above the corolla base, central staminode extremely reduced or

absent. Disk a ventral lobe, 0.6-1 mm high. Pistil 2.2-2.3 mm long; ovary urceolate,

c. 1.5 mm, glabrous or sparsely glandular at base, apically eglandular hairy; style c.

0.6 mm, hairy; stigma broadly bilobed, lobes 0.1-0.2 mm long, rounded. Fruit green,

1-3 cm long, 1-2 mm diam., hairy almost exclusively in the terminal half, straight or

curved. Seeds pale brown, elliptic, 0.4-0.5 x 0.2-0.3 mm.

Distribution. Northern Thailand (Uthai Thani).

Habitat. On limestone in secondary forest.

Provisional IUCN conservation assessment. Critically Endangered CR Blab(iii,iv) +

A revision of Microchirita in Thailand

263

B2ab(iii,iv). This species is only known from the type collection in the Huppatat Non

Hunting Area, where only one, small population was observed. The limestone range

there is only about 12 km 2 in total and is subject to disturbance from tourism. There are

no collections from the nearby Khao Pha Ra, and the area is surrounded by cultivated

land.

Additional specimens examined. THAILAND: Uthai Thani: Lan Sak, Huppatat Non Hunting

Area, 19 Sep 2015, Tanming, W. 901 (QBG).

Notes. This species can be recognised by the small, personate corolla and the glabrous

anthers. The measurements reported are largely based on the original publication in

Puglisi et al. (2016), integrated with the newly acquired material.

21. Microchirita purpurea D.J.Middleton & Triboun, Thai Forest Bull., Bot. 41: 14

(2013). - TYPE: Thailand, Chanthaburi, Kaeng Hang Maeo, Khao Chamao National

Park, Wat Khao Wong Kot, 30 m alt., 27 August 2012, Middleton, D.J., Karaket,

P., Suddee, S. & Triboun, P. 5681 (holotype BKF; isotypes A [00435722], BK, E

[E00626983], K, P [P00966760], QBG, SING [SING0229833]). (Fig. 13D-F)

‘Chirita sp. nov.?’ in Barnett, FI. Siam. 3: 228 (1962).

Caulescent herb 25-100 cm tall with elongated stem runners, internodes 2-10 cm.

Stems succulent, green with reddish parts, with a sparse hispid eglandular indumentum;

branches sometimes arising from the petioles. Leaves opposite, with the exception of

the basal leaf; petioles 2-9.5 cm long, reddish, sparsely eglandular hairy; blades pale

green above and beneath, ovate or obovate, 4.3-20 x 2.7-9.5 cm, 1.3-1.8 times as

long as wide, base cordate, apex acuminate, sparsely hairy above and beneath with a

dimorphic indumentum of longer and shorter eglandular hairs ciliate along the margin,

margin entire, 9-23 pairs of secondary veins, steeply ascending, venation slightly

sunken above and raised beneath in fresh material, flat in dry specimens, tertiary

venation inconspicuous. Inflorescences cristate, peduncles reduced or emerging

and fused together; peduncle absent or to 5 mm long; bracts absent; pedicels pale

green or tinged with reddish purple, 5-30 mm long, densely pubescent. Calyx green,

actinomorphic, completely divided to base, lobes narrowly ovate, 7-9.5 x 1.5-1.9

mm, apex acuminate, pubescent. Corolla 24-30 mm long, purple, slightly paler

on tube and lobes, darker in the throat, tube narrow at base, curved, then widening

abruptly into a campanulate upper tube, lobes not spreading, purple, eglandular hairy

outside, glabrous inside, except for a patch of glandular hairs dorsally in the tube by

the anthers; tube c. 19-19.5 mm ventrally; lobes broadly orbicular, apices rounded,

upper lobes 4x6 mm, lateral lobes 6 x 9.5 mm, ventral lobe c. 5-8.5 x 13-15 mm.

Stamens arising from the point where the tube widens, c. 9 mm above the corolla base,

filaments upright, glabrous, c. 1.5 mm long, c. 0.2 mm wide; anthers white or pale

yellow, attached at a right angle from the filament, with hairs by the filament insertion.

264

Card. Bull. Singapore 69(2) 2017

Fig. 13. Microchirita personata C.Puglisi. A. Inflorescence. B. Habit. C. Detail of the flower

from above. All from Middleton, D.J. et al. 5688. Microchirita purpurea D.J.Middleton &

Triboun. D. Habit. E. Lateral view of the flower. F. Front view of the flower. All from Middleton,

D.J. et al. 5681. (Photos: P. Karaket)

A revision of Microchirita in Thailand

265

c. 2.5 x 1.5 mm, apically joined by a connective, thecae parallel; lateral staminodes

c. 1 mm long, arising near the corolla base, central staminode c. 0.7 mm long, arising

near the corolla base. Disk annular, lobed, 1 mm high. Pistil c. 12 mm long; ovary 5

mm long, c. 0.5 mm diameter, glabrous; style 7 mm long, glabrous; stigma chiritoid,

sparsely glandular, c. 1.2 mm long. Fruit green when immature, c. 4 cm long, c. 2 mm

diameter, glabrous, curved. Seeds not seen.

Distribution. Southeastern Thailand. Vietnam.

Habitat. On limestone.

Provisional IUCN conservation assessment. Endangered EN B2ab(iii). This species

is common at the type locality and can be observed occurring high up on rather

inaccessible cliffs. At the site where it was collected there is only about 1 km 2 of

suitable habitat and the base of the outcrop and the surroundings are degraded by

human activity. The only other locality where it has been collected is just over 400

km to the SE from the type locality in southern Vietnam. Available limestone sites

inbetween are rather few.

Additional specimens examined. THAILAND: Rayong: 25 Oct 1960, Chandraprasong, K.

69 (BK); Temple near Khao Chamao, 20 Sep 2000, Watthana, S. & Wongnak, M. 991 (QBG).

Notes. This species can be recognised by the dark purple colour of the corolla, the

abruptly campanulate corolla, and the small upper two lobes and the broad lower

three lobes of the corolla. The description is largely based on the original publication

(Middleton & Triboun, 2013), with some additional data.

22. Microchirita rupestris (Ridl.) A.Weber & Rafidah, Taxon 60: 779 (2011); Rafidah,

Gard. Bull. Singapore 69: 18 (2017). - Chirita rupestris Ridl., J. Straits Branch Roy.

Asiat. Soc. 44: 59 (1905); Barnett, FI. Siam. 3: 227 (1962); Wood, Notes Roy. Bot.

Gard. Edinburgh 33: 201 (1974); Burtt, Thai Forest Bull., Bot. 29: 89 (2001). - TYPE:

Malaysia, Kedah, Langkawi, on damp rocks, sea level, November 1889, Curtis

2120 (lectotype SING [SING0042989], designated by Puglisi in Rafidah (2017: 18);

isolectoype SING [SING0042990]). (Fig. 14)

Chirita glasgovii Ridl., J. Straits Branch Roy. Asiat. Soc. 44: 60 (1905). - TYPE:

Malaysia, Perak, Waterloo Estate, December 1897, Robertson-Glasgow s.n. (holotype

SING [SING0042988]).

Chirita kerrii Craib, Bull. Misc. Inform. Kew 129 (1914); Barnett, FI. Siam. 3: 225

(1962). - TYPE: Thailand, Mae Ping Rapids, Fa Man, in crevices of damp rock, 180 m,

Kerr 2194 (lectotype K [K000545603], designated by Wood (1974: 200), isolectotype

BM [BM000997770]).

266

Card. Bull. Singapore 69(2) 2017

ChiritageoffrayiPellcgr. inLecomte,Fl. Indo-Chine4: 529(1930).-TYPE: Cambodia,

Kampot, Kabal-Romeas, 18 October 1903, Geoffray 123 (lectotype P [P00602516],

designated by Wood (1974: 199)).

Chirita hamosa var. unifolia auct. non C.B.Clarke: Barnett, FI. Siam. 3: 225 (1962).

Chirita caerulea auct. non R.Br.: Wood, Notes Roy. Bot. Gard. Edinburgh 33: 200

(1974), p.p.; Burtt, Thai Forest Bull., Bot. 29: 87 (2001), p.p.

Terrestrial or lithophytic caulescent herb to 50 cm tall, not branching. Stems fleshy,

eglandular hairy, green or purple-brown, with white and patent hairs, internodes to

c. 13 cm. Leaves opposite, except for the single basal cotyledon; petioles 2-10 cm

long, eglandular hairy; blades pale to dark green above with hispid eglandular hairs,

pale green beneath, with the same type of indumentum, elliptic, ovate or lanceolate,

4.2-14.5 x 1.8-7.5 cm, 1.5-2.7 times as long as wide, base acute to obtuse or almost

cordate, often unequal, apex acute to acuminate, margin entire to serrate, 7-16 pairs of

secondary veins, venation slightly sunken above and raised beneath in fresh material,

tertiary venation often inconspicuous. Leaves and bracts mid green above, whitish

green beneath. Inflorescences arising from the petiole, at the axil or in close proximity,

consisting of one to three main pedunculate inflorescences, each very compressed and

thus appearing subumbellate and bearing 1-10 flowers; peduncle 2-140 mm long,

eglandular hairy; bracts sessile, partly or completely fused in a cup, 5-50 mm long,

with patent white eglandular hairs on both sides, palmate venation, margin serrate

and when partly fused, with acute apex; pedicels green, 0.1-15 mm long, eglandular

pubescent. Calyx yellow-green, bilabiate with the lower lobes shorter than the upper,

tube 0.3-0.5 mm long, lobes narrowly lanceolate to triangular, 7-8 x 1.1-1.2 mm, apex

narrowly acute, eglandular hairy on the outer side, sometimes only at the tip and along

the margin, inside glabrous. Corolla 16.5-22 mm long, with tube white and lobes

white to violet, with thin dark stripes and a yellow marking ventrally in the throat,

tube narrow at base (5.5-6 mm), then abruptly opening into spreading lobes, tube

glabrous, limb densely eglandular hairy outside, internally with dense glandular hairs

in the throat, especially on the upper lip and on the ventral lobe, further in with shorter

glandular hairs; tube c. 13-13.5 mm long dorsally, 12-16.5 mm ventrally, 11.5-13.5

cm laterally between lips; upper lobes broadly elliptic, 4.5-5.5 x 4.5-6 mm, lateral

lobes elliptic, 4-5.5 x 5-6 mm, lower lobe elliptic, 4.5-6.5 x 5.5-7 mm. Stamens

arising 6-9 mm above the corolla base, slightly bent, glabrous, 3.5-4 mm long, 0.4-

0.5 mm wide; anthers pale yellow, hairy dorsally and partially beneath, 2-2.1 x 1-1.3

mm, free, thecae divergent; lateral staminodes 0.7-2.5 mm long, arising 5.5-7.5 mm

above the corolla base, glabrous, central staminode 0.2-0.7 mm long, arising 6-8

mm above the corolla base. Disk a partial ring interrupted dorsally, lobed, 0.3-0.5

mm. Pistil 13.5-16.5 mm long; ovary 7.5-11 mm long, glabrous for 2-3 mm at base,

otherwise densely eglandular hairy; style white, 4.5 mm, more sparsely hairy than the

ovary; stigma white, elongated and flexed downwards, eglandular hairy, lobes 1.5-1.8

x 0.4-0.5 mm, elliptic. Fruit green, 1.1-4 cm long, c. 1.5 mm diameter, glabrous at the

base, otherwise sparsely eglandular hairy, straight or slightly curved. Seeds not seen.

A revision of Microchirita in Thailand

267

Fig. 14. Microchirita rupestris (Ridl.) A.Weber & Rafidah. A. Unifoliate habit. B. Caulescent

habit. C. Side view of the flower. D. Detail of the corolla. A, B from Middleton, D.J. et al.

5721; C from Middleton, D.J. et al. 4836; D from Middleton, D.J. et al. 5204. (Photos: A, B, P.

Karaket; C, D, D.J. Middleton)

Distribution. Widespread in Thailand. Cambodia, Vietnam, Malaysia.

Habitat. On rocks in shady areas.

Provisional IUCN conservation assessment. Least Concern LC. This species is

common and widespread.

Additional specimens examined. THAILAND: Ayutthaya: s.l., c. 30 m, 21 Nov 1970,

Smitinand, T, et al. 11377 (BKF, E). Chiang Mai: 400 m, 23 Nov 1920, Kerr, A.F.G. s.n.

(BM); Sangampang, Muang Awn Cave, 525 m, 12 Oct 1989, Maxwell, J.F. 89-1221 (CMU,

E); Mae On, 500 m, 7 Sep 2011, Pooma, R. et al. 7797 (BKF, E); Mae On, Doi Lohn, 13 Apr

2005, Palee, P. 799 (CMUB); Mae Ping Rapids 150 m, 17 Dec 1913, Kerr, A.F.G. 3044 (BM,

268

Card. Bull. Singapore 69(2) 2017

K); Keng Um Kong, Mae Ping Rapids, 250 m, 24 Nov 1920, Kerr, A.F.G. 4651 (BM, K, BK).

Kanchanaburi: Ban Wangpho, 15 Oct 1967, Chermsirivathana, C. 791 (BK); Thong Pha

Phum Pha Tad National Park, 130 m, 6 Sep 2007, Suddee, S. et al 3268 (BKF); Thong Pha

Phum, 22 Aug 2006, Triboun, P. 3635 (E); ibidem, 130 m, 15 Sep 2006, Suddee, S. et al. 2854

(BKF); ibidem, 240 m, 29 Nov 1982, Koyama, H. et al T-30467 (BKF); Thong Pha Phum,

Wat Tham Mong Kala, 5 Oct 2015, Puglisi, C. et al CP409 (SING); Thong Pha Phum, Wat

Tha Khanun, 170 m, 28 Oct 2009, Middleton, D.J. & Triboun, P 5204 (BKF, E); Thong Pha

Phum, Cha Lae, Ban Kroeng Krawia, 20 Sep 2011, Sirimongkol, S. 226 (BKF, SING), S. 227

(BKF); Erawan Waterfall, c. 150 m, 10 Oct 1971 ,Murata, G. etal T-16141 (BKF, L); Mahidol

University Campus, 4 Oct 2015, Puglisi, C. et al CP402 (BKF); Sai Yok, Mahidol University

Campus, 275 m, 13 Sep 2005, Maxwell, J.F. 05-489 (BKF, CMUB); Srisawat, Erawan National

Park, 100-300 m, 3 Nov 1979, Shimizu, T. et al T-21532 (BKF); Erawan National Park, 400

m, 18 Nov 1971, van Beusekom, C.F. et al 3843 (L); Sai Yok, Khao Yen, 7 Oct 2015, Puglisi,

C. et al CP430 (BKF, E, MUKA, SING). Lampang: Ngao, near Tham Pha Thai, 520 m,

24 Sep 2014, Middleton, D.J. et al. 4585 (BKF, E). Satun: Tarutao Island, 30 m, Oct 1979,

Congdon 9 (E); Tha Lea Ban National Park, 250 m, 8 Dec 1986, Niyomdham, C. 1297 (BKF,

E, K, P); Khuan Doan Khao Mot Oaeng, 456 m, 5 Sep 2013, Suddee, S. et al 4574 (BKF); La-

ngu, Koh Kabeng, under 80 m, Sep 1999, Phengklai, C. 12152 (BKF); Cult. Aberdeen from a

Kerr collection in Satul, 27 Nov 1927, “785” (ABD); Cult. Aberdeen from a Kerr collection in

Satul, 6 Nov 1927, “785” (ABD). Tak: Umphang, Umphang Wildlife Sanctuary, 860 m, 16 Oct

2014, Middleton, D.J. et al 5721 (BKF, E); Tali Song Yang, Tham Usu, c. 500 m, 10 Sep 2009,

Middleton, D.J. & Triboun, P. 4836 (BKF, E, P); Umphang, Umphang Wildlife Sanctuary, 868

m, 16 Oct 2014, Middleton, D.J. et al 5711 (BKF, E). Trang: Huay Yot, Wat Tham Iso, 102 m,

9 Sep 2008, Middleton, D.J. et al. 4424 (BKF, E); Nam Tai, 11 Oct 1970, Charoenphol, C. et

al. 3628 (AAU, BKF, E, K, L); Khao Pma, 150 m, 23 Oct 1991, Larsen, K. et al. 42504 (BKF);

Palien, Lipang, Ban Tahkao, 50 m, 21 Nov 1986, Maxwell, J.F. 86-957 (CMU, L). Trat: Klong

Kloi, Kaw Chang, 30 m, 30 Sep 1924, Kerr, A.F.G. 9260 (ABD [4 sheets], E, K).

Notes. Thai specimens from south of the Isthmus of Kra have leaves with longer

petioles, more strongly serrate margin and more acuminate apex, and inflorescences

less hairy on pedicels and bracts. Wood (1974) attributed some Thai specimens to

Microchirita caerulea (R.Br.) Yin Z.Wang, an Indonesian species with a bracteate

inflorescence. Microchirita caerulea has glandular hairs intermixed with eglandular

hairs in the stem. The bracts are fused on one side only and have glandular and

eglandular hairs along the margin. The peduncles have a mixed indumentum, too, and

the pedicels are glandular. Microchirita rupestris does not have any glandular hairs.

Furthermore, the leaf base is cordate in Microchirita caerulea and only very rarely so

in M. rupestris.

23. Microchirita suddeei D.J.Middleton & Triboun, Thai Forest Bull., Bot. 41: 18

(2013). - TYPE: Thailand, Phrae, Rong Kwang District, Tham Pha Nang Khoi,

210 m alt., 17 August 2012, Middleton, D.J., Karaket, R, Suddee, S. & Triboun,

P. 5618 (holotype BKF; isotypes E [E00629451], P [P00966761], QBG, SING

[SING0229832]). (Fig. 15A-C)

A revision of Microchirita in Thailand

269

Caulescent herb to 40 cm tall, internodes 4-13 cm. Stems succulent, pale green or red

at base, sometimes branching from basal petiole, with sparse sessile glands. Leaves

opposite, apart from the basal leaf; petioles 0.3-2.2 cm long, very sparsely eglandular

hairy; blades mid green above, paler beneath, ovate, 4.2-35 x 1.3-18.8 cm, 1.3-3.3

times as long as wide, base rounded to cordate, apex acuminate, sparsely eglandular

hairy above and beneath, sparsely ciliate along the margin, margin entire, 6-22 pairs of

secondary veins, venation slightly sunken above and raised beneath in fresh material,

flat in dry specimens, tertiary venation laxly reticulate. Inflorescences cristate;

peduncles fused, 2.5-5 mm long, fused together, glabrous; bracts absent; pedicels

green, 4.4-11 mm long, sparsely hairy. Calyx green, bilabiate, the two lips completely

divided, ventral tube c. 0.1 mm long, dorsal tube to c. 3.5-5 mm long, lobes ligulate,

dorsal lobes 7.4-8 mm long, lower lobes 3-5 mm long, apex acute, margin entire,

irregular towards the tip, sparsely hairy outside, hairier at the tip, sparsely glandular

inside. Corolla 10-12 mm long, tube white, lobes pale lilac, tube narrow, straight or

just slightly curved, lobes not always spreading, eglandular hairy outside, glabrous

inside; tube c. 8 mm long dorsally, c. 10 mm long ventrally; lobes broadly orbicular,

apices rounded or obtuse, upper lobes c. 2.6 x 3.7 mm, lateral lobes c. 2.4 x 4 mm,

ventral lobe c. 1.8-3 x 4 mm. Stamens arising 3.2-4 mm above the corolla base,

filaments slightly curved, glabrous, 2.2-2.5 mm long, c. 0.3-0.4 mm wide; anthers

with an abundant indumentum at the sides and dorsally, 2.2-2.5 x 1.1-1.2 mm, apically

joined by a weak connective. Disk a small ventral lobe, 0.1-0.6 mm high. Pistil c. 10

mm long; ovary 5.5-6.5 mm long, papillose in the bottom half, densely pubescent

above; style white, 3.3^1 mm long, densely pubescent; stigma white, c. 1 mm long.

Immature fruit 2-3 cm long, c. 1 mm diameter, glabrous, straight. Seeds not seen.

Distribution. Northern Thailand.

Habitat. On limestone in evergreen or mixed deciduous and bamboo forest.

Provisional IUCN conservation assessment. Vulnerable VU Blab(iii) + B2ab(iii).

The EOO is almost 10,000 km 2 , well within the boundary for Vulnerable. However,

the AOO is within the boundary for Endangered, not all of the localities are within

protected areas, and limestone sites are degraded throughout the region. Since there

are five known localities and there are likely to be more, an assessment of Vulnerable

is more appropriate than Endangered.

Additional specimens examined. THAILAND: Lampang: Ngao, near Tham Pha Thai, 520

m, 24 Sep 2008, Middleton, D.J. et al. 4580 (BKF, E, P); Chae Horn, Ban Sa village, 15 Oct

2006, Palee, P. 1027 (A, CMUB). Phrae: Long, Doi Pha Klong National Park, Bah Gah Lahn

Hill, 26 Oct 2005, Palee, P. 846 (A). Tak: Tha Song Yang, Doi Noi, 242 m, 6 Sep 2013,

Phaosrichai, P. & Wongnak, M. 23 (QBG).

Notes. The floral characters reported are taken from the original description of

Microchirita suddeei (Middleton & Triboun, 2013), as no new material has been

270

Card. Bull. Singapore 69(2) 2017

identified during this study. This species can be recognised by the hairy anthers, the

pale lilac corolla and the absence of a ventral line and lateral spots in the corolla. It is

most similar to Microchirita lilacina, which has a broader mouth and glabrous anthers,

and to M. albiflora, which is completely white and lacks sessile glands on the stem.

24. Microchirita tadphoensis C.Puglisi, sp. nov.

Most similar to Microchirita hamosa (R.Br.) Yin Z.Wang in the delicate habit and to

M. bimaculata (D.Wood) A.Weber & D.J.Middleton in the shape of the corolla. Differs

in having a shortly campanulate pale yellow corolla (white in Microchirita hamosa )

with a ventral darker yellow marking but no lateral spots (spots always present in M.

bimaculata). - TYPE: Thailand, Nakhon Phanom, Ban Phaeng, Phu Langka National

Park, Tad Pho Waterfall, 224 m, 23 October 2015, Suddee, S., Keiwbang, W. & Hemrat,

C. 4980 (holotype BKF; isotype SING). (Fig. 15D-F)

Herb to 20 cm tall, stem sparsely aglandular hairy, not branched, internodes c. 3 cm

long. Leaves opposite, apart from the basal leaf; petioles 1-4 mm long, sparsely

eglandular hairy; blade ovate to elliptic, 1.7-4.2 x 1.2-3.2 (a partial much larger basal

leaf was seen that would be beyond these measurements if complete), 1.3-1.5 times as

long as wide, base subcordate, apex acute to broadly acute, eglandular hairy on both

surfaces, margin entire and ciliate, 7-9+ pairs of secondary veins, flat and scarcely

visible on both sides. Inflorescence cristate, 1-5-flowered, floral axes eglandular

hairy; bracts absent; pedicels 0.5-1.5 cm, straight or curved. Calyx pale green, slightly

zygomorphic, lobes almost free, lanceolate-elliptic, 3-3.3 x 0.5-0.9 mm, lower lobes

slightly larger than the upper, apex acute, margin entire, eglandular hairy outside,

glabrous inside. Corolla c. 12 mm long, white outside, inside pale yellow with a yellow

patch ventrally, base of tube narrow and curved, then gradually broadening, sparsely

eglandular hairy outside, inside glabrous basally and with sessile glands apically and

on the lobes; tube c. 9.3 mm dorsally, c. 9.7 mm ventrally and c. 8 mm between the

lips; upper lobes c. 2 x 2.5 mm, lateral lobes c. 1.3 x 3 mm, ventral lobe c. 2 x 3 mm.

Stamens arising c. 4.6 mm above the corolla base; filaments c. 2 mm long, 0.2 mm

diamenter, straight, glabrous at the base, with sessile glands in the top half; anthers c.

1.2 x 0.5 mm, probably ligate, with a small patch of straight hairs by the attachment,

thecae strongly divergent; staminodes not seen. Disk absent. Pistil c. 10 mm long;

ovary c. 5 mm long, c. 0.7 mm diameter, glabrous in the basal 1/3 to Vi, then eglandular

hairy; style c. 3.5 mm, delicate, eglandular hairy; stigma with lobes c. 0.7 x 0.5 mm,

somewhat plumose inside. Fruit eglandular hairy, more or less curved, 1.7-3.4 cm

long, 0.7-1.2 cm diameter. Seeds dark brown to black, broadly elliptic and furrowed,

c. 0.5 x 0.2 mm.

Distribution. Northeastern Thailand.

Habitat. On sandstone in dry evergreen forest.

A revision of Microchirita in Thailand

271

Fig. 15. Microchirita suddeei D.J.Middleton & Triboun. A. Habit. B. Inflorescences. C. Detail

of the corolla. All from Middleton, D.J. et al. 5618. Microchirita tadphoensis C.Puglisi. D.

Lateral view of the corolla. E. Front view of the corolla. F. Fruits. All from Suddee, S. et al.

4980. (Photos: A-C, P. Karaket; D-F, S. Suddee)

272

Card. Bull. Singapore 69(2) 2017

Provisional IUCN conservation assessment. Data Deficient DD. This species is only

known from the type collection, which is within a protected area.

Notes. The epithet refers to the type locality.

25. Microchirita tetsanae C.Puglisi, sp. nov.

Species characterised by the presence of a dimorphic indumentum on the anthers and

sparse hairs on the filaments, and by a little projection at the anther insertion. It is

most similar to Microchirita thailandica C.Puglisi, but differs in the upper lobes not

being imbricate with the lower, and in the filament projection. - TYPE: Thailand,

Phetchabun, Mueang Phetchabun, Wat Tham Nam Bang, 130 m, 13 September 2014,

Tetsana, N. et al. 855 (holotype BKF; isotype SING). (Fig. 16A-B)

Caulescent herb to 50 cm tall with elongated stem runners, internodes to 3.5-12.5

cm. Stems succulent, with some eglandular indumentum; branches to 30 cm long,

sometimes arising from the petioles. Leaves opposite, apart from the basal leaf; petioles

0.1-2.5 cm long, eglandular hairy; blades mid green above, paler beneath, lanceolate

to elliptic, 3-16 x 2.2-10 cm, 1.3-2.1 times as long as wide, base subcordate to

cordate, apex acute to acuminate, finely pubescent above and beneath, ciliate along the

margin, margin entire or sparsely and minutely denticulate, 6-17 pairs of secondary

veins, venation slightly sunken above and raised beneath in fresh material, flat in dry

specimens, tertiary venation inconspicuous. Inflorescences cristate, 1-5-flowered;

peduncles reduced or 3-8 mm long, fused with each another, pubescent; bracts absent;

pedicels pale green, 0.1-2.5 cm long, pubescent. Calyx pale green, succulent at the

base, tube 0.3-0.5 mm long, lobes imbricate, narrowly lanceolate, 4.5-8 x 1-1.2 mm,

apex acuminate and thickened, margin entire, outer indumentum of long eglandular

hairs along the midrib, margin eglandular ciliate, inner indumentum of eglandular

hairs at the tip and sparse sessile glands. Corolla 17-26 mm, tube white to pale blue,

lobes white or purple-blue, the base of the upper lobes white or dark blue, lower

lip pale blue or white at the base, with a bright yellow stripe ventrally, tube narrow,

strongly curved, lower lip expanded, upper lip slightly reflexed, glabrous at the base,

then finely eglandular hairy, the base of the upper lobes and of the ventral lobe with a

dense glandular indumentum, the lobes sparsely glandular hairy; tube 10-13 mm long

dorsally, 13-20 mm ventrally, 10-14.5 mm laterally between lips; upper lobes elliptic,

2-6 x 2-7 mm, lateral lobes obtuse, 4-6 x 5-8.5 mm, ventral lobe elliptic, 4.5-7.5 x

6-12 mm. Stamens arising 6-10.5 mm above the corolla base; filaments straight, with

a little projection dorsally at the base of the anther, pale yellow, sparsely and minutely

hairy, 2.5-3 mm long, 0.4-0.6 mm wide; anthers white, with an indumentum of long

coloured hairs growing on the outer end and short white hairs growing on the inner end,

1.7-2.5 x 1.5 mm, apically joined by a connective, thecae parallel; lateral staminodes

0.7-1 mm long, arising c. 1.5-7 mm above the corolla base, central staminode 0.2-0.5

mm long, arising 3-7.5 mm above the corolla base. Disk annular, margin subentire,

0.3-0.7 mm high. Pistil c. 15-20 mm long; ovary 6-6.5 mm long, c. 1 mm diameter,

A revision of Microchirita in Thailand

273

sparsely eglandular hairy or glabrescent in the bottom half, hairier towards the apex;

style arising at almost a right angle to the ovary, 8.5-9.5 mm, eglandular hairy; stigma

c. 0.7 mm long, glabrescent, bilobed, lobes rounded, 0.3-0.5 mm. Fruit and seeds not

seen.

Distribution. Northern and Northeastern Thailand.

Habitat. On limestone in dry evergreen forest.

Provisional IUCN conser\>ation assessment. Vulnerable Blab(iii). This species is only

known from a few collections and has an EOO of < 6,000 km 2 . Not all of the localities

are within protected areas. Its AOO might suggest a status of Endangered but much

of the potential distribution has not been adequently explored to be confident that its

AOO is within the limit.

Additional specimens examined. THAILAND: Khon Kaen: Choonpa [Chum Phae], 23 Nov

1963, Pradit 738 (BK). Loei: Phu Luang Wildlife Reserve, Nam Tok to Pa Paw trail, 29 Sep

1990, Chantaranothai, P. et al. 90/439 (BKF, K). Phitsanulok: Noen Maprang, Bahn Mung,

Thung Salaeng Luang National Park, 85 m, 31 Aug 2008, Sreepoowiang, K. K 52 (SING);

Noen Maprang, Tham Pha Tha Phon Non-Hunting Area, 9 Sep 2011, Norsaengsri, M. &

Insea, R. 8188 (QBG); Noen Maprang, Tham Pha Tha Phon Wildlife Sanctuary, 17 Nov 2010,

Maknoi, C. 3973 (QBG).

Notes. This species is named after Dr Naiyana Tetsana from the Forest Herbarium

Bangkok, who collected and shared a number of specimens and photos of the new

species described in this article. Microchirita tetsanae has individuals with purple-

blue and cream-white flowers, sympatric in Phitsanulok. No morphological characters

support a possible recognition of these as different taxa and they are treated here only

as colour forms of the same species. It is also reported from Uttaradit (photos from

Pranee Nangnam seen).

26. Microchirita thailandica C.Puglisi, sp. nov.

Species most similar to Microchirita tetsanae C.Puglisi in the colour pattern of the

corolla, differing in having a narrower tube which widens abruptly (gradually widening

in M. tetsanae ), all corolla lobes imbricate (vs. lateral lobes not imbricate with the

upper in M. tetsanae ), a shorter ventral tube, and in not having a projection at the

anther insertion. - TYPE: Thailand, Chaiyaphum, Phak Dee Chumphon, Wat Thum

Wua Daeng, 460 m, 8 November 2014, Tetsana, N. et al. 904 (holotype BKF; isotype

SING). (Fig. 16C-E)

Caulescent herb 10-50 cm tall, internodes 3-6 cm. Stems succulent, with sparse

eglandular indumentum, branching from the basal petiole. Leaves opposite, apart

from the basal leaf; petioles 0.3-0.8 cm long, sparsely eglandular hairy; blades mid

274

Card. Bull. Singapore 69(2) 2017

Fig. 16. Microchirita tetsanae C.Puglisi. A. Habit. B. Front view of the flower. All from

Tetsana, N. et al. 855. Microchirita thailandica C.Puglisi. C. Habit. D. Front view of the flower.

E. Lateral view of the flower. All from Tetsana, N. et al 904. (Photos: N. Tetsana)

A revision of Microchirita in Thailand

275

green above, ovate to lanceolate, 2.8-11 x 1.5-6.8 cm (basal leaf likely to have been

larger but not seen), 1.6-1.9 times as long as wide, base truncate to shortly attenuate,

apex acute to acuminate, eglandular hairy above and beneath, margin ciliate, margin

entire, 7-8 pairs of secondary veins (basal leaf not seen), venation slightly sunken

on the adaxial side. Inflorescence cristate, many-flowered, floral axes hispid with

mixed eglandular and glandular hairs; peduncles extremely reduced and fused with

each other; bracts absent; pedicels 2-15 mm long. Calyx pale green, c. 9.5 mm long,

tube 5-7 mm long, lobes narrowly lanceolate, c. 8.5 x 1.5 mm, apex narrowly acute

to acuminate, lobes not imbricate, with sparse eglandular hairs outside and an even

sparser indumentum inside, mixed with sessile glands. Corolla c. 23 mm long, tube

purple, lobes purple-blue, mouth pale blue, with a yellow ventral marking and a dark

purple patch at the base of the upper lobes, tube narrow, bent downwards, abruptly

widening, outer side hairy throughout, inside densely covered in eglandular hairs in

the mouth and dense sessile glands over the ventral yellow spot; tube c. 13.5 mm

dorsally, c. 14 mm ventrally and laterally; upper lobes c. 5.5 x 8.5 mm, lateral lobes c.

5.5 x 8 mm, ventral lobe c. 7 x 8 mm. Stamens arising c. 9 mm above the corolla base,

filaments straight, c. 2 mm long, c. 0.5 mm diameter, without any projection; anthers

ligate, with an indumentum of long brown hairs growing on the outer end and short

white hairs growing on the inner end, c. 1.1 x 1.3 mm; lateral staminodes c. 1 mm

long, arising c. 8 mm above the corolla base, central staminode c. 0.8 mm long, arising

c. 9 mm above the corolla base. Disk annular, c. 0.9 mm. Pistil c. 13 mm long; ovary c.

8 mm long, c. 1 mm diameter, glabrous in the basal 1/3, eglandular tomentose above;

style enantiostylous, c. 5 mm, eglandular hairy; stigma 0.8 mm long, densly glandular

hairy, shallowly bilobed, lobes rounded, c. 0.4 mm. Fruit straight, sparsely eglandular

hairy, 3-5 cm. Seeds c. 0.4 x 0.2 mm, brown, elliptic.

Distribution. Eastern Thailand.

Habitat. On limestone.

Provisional IUCN conservation assessment. Data Deficient (DD). This species is only

known from the type locality and too little is known of its distribution or possible

threats (except that the single collection was not made in a protected area).

Notes. This species has been named after the country to which it is endemic.

27. Microchirita tubulosa (Craib) A.Weber & D.J.Middleton, Taxon 60: 79 (2011).

- Chirita tubulosa Craib, Bull. Misc. Inform. Kew 173 (1922); Barnett, FI. Siam.

3: 227 (1962); Wood, Notes Roy. Bot. Gard. Edinburgh 33: 190 (1974); Burtt, Thai

Forest Bull., Bot. 29: 89 (2001). - TYPE: “described from plants grown from seed

collected by Kerr. Flowered in October 1921” (lectotype ABD, first step designated

by Wood (1974: 190), second sted designated here [the specimen which matches the

image deposited inE, barcode E00155288]; isolectotypes ABD, E). (Fig. 17)

276

Card. Bull. Singapore 69(2) 2017

Chirita barbata auct. non Sprague: Barnett, FI. Siam. 3: 222 (1962).

Caulescent herb to 50 cm tall, internodes 1.5-9 cm long, sometimes branches arising

from the petioles. Stem fleshy, green or flushed red-brown, eglandular hairy. Leaves

opposite, apart from the basal leaf; petioles green or purple-brown, 2-20 mm long;

blades pale to dull green above, paler beneath, lanceolate, elliptic or ovate, 3.5-36 x

1.4-20 cm, 1.7-3.9 times as long as wide, base obtuse to subcordate to rarely somewhat

attenuate, apex acute to acuminate, eglandular hispid above and beneath, margin

ciliate, margin entire to obscurely toothed, midrib green or red-brown, 5-14 pairs

of secondary veins, venation more or less sunken above and raised beneath in fresh

material, flat in dry specimens, tertiary venation lax and inconspicuous. Inflorescences

cristate, 1-8-flowered; peduncles 2-10 mm long, sometimes fused with each other;

bracts absent; pedicels green, 0.5-2 cm long, eglandular hairy. All floral axes with

sparse to dense eglandular hairs. Calyx bilabiate, green, lobes succulent along the

midrib, lower tube 0.6-0.8 mm, lateral tube 0.1-0.6 mm, upper tube c. 1.2 mm, lobes

imbricate, lanceolate, 11-17 x 2-3 mm (the dorsal wider than the lateral, which in turn

are wider than the ventral), apex acute to slightly acuminate, margin entire, eglandular

hairy outside on margin and midrib, glabrous inside except tip. Corolla 17-25 mm

long, white outside on tube and lobes, creamy yellow in throat, with a yellow ventral

stripe and lateral purple-brown stripes or spots, tube narrow, usually curved, becoming

pouched, with upper and lateral lobes spreading, ventral lobe prominent and forming

a sinus at the base where it connects with the lateral lobe, tube glabrous at the base,

outside densely and minutely eglandular and glandular hairy, glabrous inside, with

sessile or stalked glands all over the throat but especially dense under the upper lobes;

tube 12-20 mm long dorsally, 16-22 mm ventrally, 15-20 mm between the lips; lobes

with obtuse apex, upper lobes 2—4.5 x 5-5.5 mm, lateral lobes 4.5-6 x 6.5-10 mm,

ventral lobe 5.5-7.5 x c. 7 mm. Stamens arising 1.1-1.2 cm above the corolla base,

filaments bent at base, 4-7 x 0.7-1.2 mm, yellow, glabrous; anthers white with white

hairs at the top and dorsally, 3-3.3 x 2-2.5 mm, apically joined by a connective, thecae

divergent, apiculate; staminodes absent or 2,0.3-0.4 mm long, arising 6-7.5 mm above

the corolla base. Disk annular or more often dorsally cleft, margin slightly lobed, 0.5-1

mm high. Pistil 13-20 mm long; ovary 0.9-1.2 cm long, c. 1.1 mm diameter, glabrous

to minutely glandular in the bottom 1/3-1/2, densely eglandular hairy above and with

some glandular hairs between ovary and style; style 0.2-0.5 mm diameter, eglandular

hairy except at the very top, 0.8-1.4 cm long; stigma broadly bilobed, lobes 1-1.3 mm,

hairy. Fruit green, 2.5-4 cm long, 1.7-2.6 mm diameter, sparsely eglandular hairy,

straight or slightly curved. Seeds dark brown, elliptic, acuminate, c. 0.4 x 0.2 mm.

Distribution. Widespread in Central, Eastern and Southeastern Thailand.

Habitat. On limestone in deciduous forest.

Provisional IUCN conservation assessment. Near Threatened NT. The EOO is >

45,000 km 2 , well beyond the threshold for a Vulnerable status. The AOO, however, is

A revision of Microchirita in Thailand

277

Fig. 17. Microchirita tubulosa (Craib) A.Weber & D.J.Middleton. A. Inflorescence and side

view of the flower; B. Lateral view of the flower. C. Front view of a mature flower. D. Front

view of a young flower. A, C from Middleton, D.J. et al. 4809; B, D from Tetsana, N. et al 800.

(Photos: A, C, D.J. Middleton; B, D, N. Tetsana)

<75 km 2 , which would be within the threshold for an Endangered status except that

the number of locations is too high for Endangered or Vulnerable. Nevertheless, many

of the collection localities for this species are not in protected areas and the regions

in which this species occurs are also where mining of limestone is particularly active.

Additional specimens examined. Unknown locality: “145” coll. 9 Nov 1927 (ABD [2 sheets]);

Cultivated at RBG Kew, 5 Nov 1923, 418-23 (K); various specimens with no data or barcode

(ABD). THAILAND: Ayutthaya: Kanam Pasak, Keng Koi, 50 m, 9 Dec 1923, Kerr, A.F.G.

7961 (ABD, BK, K). Chonburi: Sriracha, Koh Seechang Isl., 7 Nov 1969, van Beusekom,

C.F. & Smitinand, T. 2039 (AAU, BKF, E, L, P); Siracha, Si Chang Island, 25 m, 10 Sep 1993,

278

Card. Bull. Singapore 69(2) 2017

Maxwell, J.F. 93-1040 (A, BKF, CMUB, L). Loei: Na Dueng, Thum Pha-ya temple, 433 m, 9

Sep 2014, Tetsana, N. et al. 800 (SING). Lop Buri: 22 Oct 1926, Smith, H.M. s.n. (BK, K);

Khao Wongkot temple, 35 m, 25 Apr 2004, Pooma, R. et al. 4657 (BKF); Tha Wting, Khao

Samo Khon, 50 m, 14 Sep 2008, Phonsena, P. et al 6245 (BK, BKF); Cultivated in Bangkok,

originally from Lop Buri, 28 Sep 1930, Kerr, A.F.G. 19720 (ABD [2 sheets], K [2 sheets], E).

Nakhon Sawan: Kao Sung, near Paknampo, 60 m, 9 Dec 1913, Kerr, A.F.G. 3019 (BM, K);

Takhli, Khao Tham Phet, 50-100 m, 27 Sep 2003, Phonsena, P. et al. 4006 (BKF); Takli, Tahm

Pet Tahm Tong Forest Park, 140 m, 15 Oct 2006, van de Bull, M. 913 (BKF, CMUB); Ban Takli,

22 Oct 1929, Put, N. “224 ” (ABD); Cultivated in Aberdeen, 1931, “Kerr” 224 (ABD, L, US);

Ban Phot Phisai, Wat Thep Satha Phon, 50 m, 8 Sep 2009, Middleton, D.J. & Triboun, P. 4809

(BKF, E, P); Banphot Phisai, Ban Daen, Khao Huai Lung, 100 m, 5 Sep 2011, Pooma, R. et al.

7761 (BKF); Banphot Phisai, Ban Daen, 100 m, 5 Sep 2011, Pooma, R. et al. 7761A (BKF);

Kao Sang, Ban Den, c. 100 m, 8 Nov 1920, Kerr, A.F.G. 4542 (ABD, BK, BM, K); Cultivated

at RBGE from Kerr 4542, 1921 (E). Saraburi: Chalerm Phrakiet, Na Phralan, Tham Sriwilai

Temple, 100 m, 9 Sep 2006, Pooma, R. & Phattarahirankanok, K. 6313 (AAU, BKF, E, SING,

QBG); Na Pra Larn, Khao Talu, 100^100 m, 6 Oct 1979, Shimizu, T. et al. T-17989 (BKF); Na

Pra Larn, Khao Talu, 100-400 m, 6 Oct 1979, Shimizu, T. et al. T-17988 (BKF); Na Pra Lam,

Khao Talu, 100-400 m, 6 Oct 1979, Shimizu, T. et al. T-18023 (BKF, L); Turn Sri Ni Lai, 11

Sep 2006, Triboun, P. 3649 (E); Kaeng Khoi, Wat Thamcharoen, 8 Sep 2006, Pooma, R. et al.

6279 (AAU, BKF, E, KEP, QBG); Kaeng Khoi, Tharn Pra Photisat, 100-200 m, 7 Oct 1979,

Shimizu, T. et al. T-19417 (BKF, L); Kaeng Khoi, Tham Pra Photisat, 7 Oct 1979, Shimizu, T.

et al. T-19464 (BKF); Htn Lap, c. 100 m, 3 Sep 1924, Kerr, A.F.G. 9127 (BK, BM, K); Hin

Lap, 19 Aug 1929, Put, N. 2410 (ABD, BK, K); Tham Pha Phothisat, Hin Lap, 100 m, 26 Sep

1979, Smitinand, T. s.n (BKF); Kang Koi, 11 Oct 1926, Lakshnakara, M.C. 287 (ABD, BK, K);

Phraphuttabat, Wat Khaowong, 18 Sep 2004, Pooma, R. et al. 4820 (BKF); BanNawng Bua, 1

Oct 1927, Put, N. 1116 ( ABD*, BK, K *an ABD specimen is a mixed collection of Microchirita

tubulosa and M. marcanii ); Phu Khae, Chong Khao Khad, Bencha Khiri Temple, 20 m, Suddee,

S. et al. 3341 (BKF); Muak Lek, 300 m, 2 Oct 1963, Smitinand, T. & Sleumer, H.O. 1317 (BKF,

E, K, L, SING); Muak Lek, 300 m, 11 Nov 1934, Marcan, A. 1885 (ABD, K).

Notes. Several specimens in ABD have no label data and could be original material.

Microchirita tubulosa is characterised by a white corolla with a ventral yellow stripe

and lateral purple markings (blotches or stripes; if stripes, sometimes multiple on each

side). It is most similar to Microchirita marcanii and M. luteola (see discussion under

each), but differs in the corolla colour pattern and the smaller flower.

28. Microchirita viola (Ridl.) A.Weber & Rafidah, Taxon 60: 779 (2011); Rafidah,

Gard. Bull. Singapore 69: 26 (2017). - Chirita viola Ridl., J. Linn. Soc., Bot. 32: 516

(1896); Barnett, FI. Siam. 3: 228 (1962); Wood, Notes Roy. Bot. Gard. Edinburgh

33: 190 (1974); Burtt, Thai Forest Bull., Bot. 29: 89 (2001). - Didymocarpus viola

(Ridl.) Williams, Bull. Herb. Boiss. Ser. 2, 5: 434 (1905). - TYPE: Malaysia,

Peninsular Malaysia, Kedah, Langkawi, September 1890, Curtis 2570 (lectotype

SING [SING0042993], designated by Rafidah (2017: 26); isolectotypes SING

[SING0042994, SING0042995]). (Fig. 18A-B)

A revision of Microchirita in Thailand

279

Caulescent perennial herb to 40 cm tall, internodes 1-10 cm, occasionally branching

from the basal portion of the basal petiole. Stems purple or green, strigose, more densely

so at the nodes. Leaves opposite; petioles 2-10 mm long, c. 1.5 mm diameter, strigose;

blades elliptic to lanceolate, green above, pale green beneath, 2.1-15 x 1.6-8.4 cm,

1.1-2.3 times as long as wide, base shortly attenuate to rounded to subcordate, apex

acute, eglandular tomentose above, hispid beneath, ciliate along the margin, margin

entire (can be serrate in some Malaysian specimens), 6-11 pairs of secondary veins

in the opposite leaves, venation slightly sunken above and raised beneath in fresh

material, tertiary venation lax and seldom visible. Inflorescence cristate, 1-6-flowered;

peduncles reduced; bracts absent; pedicels 7-20 mm long, eglandular hairy. Calyx

green, succulent at the base, lobes almost free to base, narrowly lanceolate, c. 6-10 x c.

1 mm, apex narrowly acute, outside eglandular hairy, inside with sparse hairs becoming

denser at the tip. Corolla 17-21 mm long, tube yellowish white inside, lobes blue-

purple with with dark blue lines, tube narrow at base, bent downwards, then gradually

expanding, with sessile glands inside, elongated glands at the base of the upper lobes

and very minute glandular hairs on the lower lip; tube 13 mm long dorsally, 16 mm

ventrally, 12 mm laterally between lips; upper lobes c. 2.5 x 4 mm, lateral lobes, c.

3.5 x 4.3 mm, lower lobe c. 5 x 4.5 mm. Stamens arising c. 8.5 mm above the corolla

base; filaments straight, glabrous, c. 3 mm long, c. 0.3 mm wide; anthers sparsely

hairy dorsally, c. 2 x 1 mm, apically coherent but not joined by a connective, thecae

divergent; lateral staminodes c. 1.2 mm long, central staminode not seen. Disk annular,

margin slightly lobate, c. 0.3 mm high. Pistil c. 14 mm long, densely eglandular hairy

throughout except at the base of the ovary; ovary c. 5.5 mm long, c. 0.6 mm diameter;

style c. 6.5 mm; stigma deeply bilobed, lobes c. 0.9 x 0.4 mm, shortly plumose inside.

Fruit 1^1.5 cm long, 0.9-1 mm diameter, densely eglandular hairy, straight or slightly

curved. Seeds brown, broadly elliptic, 0.4-0.5 x 0.2-0.3 mm.

Distribution. Peninsular Thailand. Malaysia.

Habitat. On limestone in evergreen forest.

Provisional IUCN conservation assessment. Vulnerable VU Blab(iii). This species

occurs on Langkawi in Malaysia and has also previously been collected on the

mainland in Kedah. Rafidah (2017) reports that no recent collections have been made

from the Kedah mainland and suggests a conservation assessment of Near Threatened

as the species is reported to be common and protected on Langkawi. In Thailand,

however, most of the known localities are not in protected areas and are subject to high

disturbance. The EOO and number of collection localities, coupled with these threats,

suggest an assessment of Vulnerable.

Additional specimens examined. THAILAND: Krabi: Ao Luek, c. 50 m, 17 Nov 1959,

Smitinand, T. & Abbe, E.C. 6151 (BKF, K (2)); Muang, Krabi Noi, Wat Tham Suea, 25 m, 17

Sep 2010, Middleton, D.J. et al. 5467 (E); Kasoom, Nov 1896, Curtis 3221 (K). Phangnga:

s.L, 28 Feb 1929, Kerr 17563 (K); Mueang Phangnga, Sri Nakharin Park, 1 m, 18 Nov 2014,

280

Card. Bull. Singapore 69(2) 2017

Suddee, S. et al. 4807 (BKF); Thap Put, Kee Ree Wong Temple, 24 Oct 2004, Saway & Rob.

11138 (QBG).

Notes. Microchirita viola is represented by rather few specimens in Thailand. A more

comprehensive description of this species is provided by Rafidah (2017).

29. Microchirita woodii D.J.Middleton & Triboun, Thai Forest Bull., Bot. 41: 15

(2013). - TYPE: Thailand, Nan, Muang Nan, Tham Pha Tup Forest Park, trail to Phra

Cave, 300 m alt., 16 August 2012, Middleton, D.J., Karaket, P., Suddee, S. & Triboun,

P. 5612 (holotype BKF; isotypes BK, E [00629450], P [P00966763]). (Fig. 18C-F)

Caulescent herb to 50 cm tall with elongated stem runners, internodes 2-15 cm. Stems

succulent, green, glabrescent or with a sparse eglandular indumentum. Leaves opposite,

apart from the basal leaf; petioles 0.3-1 cm long, green, sparsely eglandular hairy;

blades pale green above and beneath, ovate to elliptic, 2.7-30 x 1.8-21.5 cm, 1.2-2.2

times as long as wide, base cordate, apex acute, sparsely eglandular hairy above and

beneath, ciliate along the margin, margin entire with sparse obscure teeth, 8-15 pairs

of secondary veins, venation sunken above and raised beneath in fresh material, flat

in dry specimens, tertiary venation laxly reticulate. Inflorescences cristate, peduncles

reduced or emerging to 5 mm long but not fused together, bracts absent; pedicels

pale green, 7-20 mm long, pubescent. Calyx green, actinomorphic, lobes completely

divided, narrowly ovate, 5.7-10.5 x 0.9-1 mm, apex acuminate and slightly thickened,

margin entire, glabrescent or sparsely hairy, hairier at tip, glabrous inside. Corolla

18-22 mm long, tube very pale green, lobes yellow-white, throat yellow with a darker

yellow ventral patch and dark brown spots to its sides, tube narrow, curved, then

widening gradually, lobes somewhat spreading, eglandular hairy outside, inside with

glandular hairs at the base of the upper lobes and dorsally in the tube; tube 12.5-15.5

mm long dorsally, 15.9-19.5 mm ventrally, c. 15 mm laterally between lips; lobes

broadly orbicular to ovate, apices rounded, upper lobes 3-4.2 x 4-6.6 mm, lateral

lobes 4.5-5.7 x 5.5-7 mm, ventral lobe 5-7.5 x 6.2-8 mm. Stamens arising 9-11.2

mm above the corolla base, filaments straight, glabrous, 2-2.5 mm long, 0.4 mm wide;

anthers white, with a dimorphic indumentum of long brown hairs above dorsally and

short white hairs below, 1.9-2.2 x 1.3-1.8 mm, apically joined by a connective, thecae

more or less parallel; staminodes reduced, c. 0.2 mm long. Disk annular, 0.4-0.8 mm

high. Pistil c. 12 mm long; ovary 5.5-7.5 mm long, c. 1.2 mm diameter, papillose at the

base, pubescent above; style 6-7.5 mm long, pubescent; stigma c. 0.8 mm long. Fruit

green, to 7 cm long, 0.8-1.2 mm diameter, eglandular hairy or glabrescent, straight or

slightly curved. Seeds maroon, narrowly elliptic, acuminate 0.3-0.5 x 0.1-0.2 mm.

Distribution. Northern Thailand (Nan).

Habitat. On limestone in evergreen forest.

Provisional IUCN conservation assessment. Data Deficient (DD). Microchirita woodii

A revision of Microchirita in Thailand

281

Fig. 18. Microchirita viola (Ridl.) A. Weber & Rafidah. A. Lateral view of the flower. B. Front

view of the flower. All from Middleton, D.J. et al. 5467. Microchirita woodii D.J.Middleton &

Triboun. C. Habit. D. Front view of the flower. E. Inflorescence. F. Side view of the flower. All

from Middleton, D.J. et al. 5612. (Photos: A, B, D.J. Middleton; C-F, P. Karaket)

282

Card. Bull. Singapore 69(2) 2017

is only known from Tham Pha Tup Forest Park which is less than 1 km 2 in size and

subject to disturbance from tourism. However, there have been far fewer collections

made in other forested areas to the north and until it can be assessed whether this

species also occurs there, its conservation status is uncertain.

Additional specimens examined. THAILAND: Nan: Tham Pha Toop Forest Park, 230 m, 2 Sep

1999, Middleton, D.J. 148 (BKF); Tham Pa Tok, 350 m, 13 Sep 1995, Larsen, K. et al. (AAU,

BKF, SING); Mueang Nan, Tham Pha Toop, 400 m, 18 Sep 1999, Srisanga, P. & Puff, C. 1110

(BKF, E, QBG); Muang, Taam Paa Toop, c. 300 m, 13 Sep 1995, Nanakorn, W. et al. 4236 (E,

QBG).

Notes. This species is recognisable by the colour pattern of the corolla which is pale

yellow-white with dark reddish-brown spots, and the dimorphic anther indumentum.

The other pale-coloured species with markings are M. huppatatensis, from which

M. woodii differs in the slightly compressed corolla mouth and the dimorphic anther

indumentum, and M. karaketii, which has violet markings and differs in the shape of

the corolla (more slender). In central and eastern Thailand, M. tubulosa has a similar

colour pattern, but its corolla is much larger and a very different shape.

Incertae sedis

The following specimens could not be determined to species for a variety of reasons,

either because the material was sterile or otherwise too poor to identify, or because

the specimen may be of an undescribed taxon but the material inadequate to describe.

Khon Kaen: Pa Potiyan, 17 Dec 1963, Youngboonkird, U. 129 (BK); Phuphaman NP,

300 m, 24 Aug 2011, Norsaengsri, M. et al. 8086 (QBG).

Loei: Phu Kradueng, Maholan Cave, 370 m, 17 Dec 1982, Koyama, H. et al. T-31171

(BKF).

Mukdahan: Mueang, Phu Pha Thoep National Park, 175 m, 25 Oct 2015, Suddee, S.

et al. 4989 (BKF).

Sakon Nakhon: Phu Pha Man National Park, Laan Sao La to Pha Nang Maen, 14 Oct

1990, Chantharanothai, P. & Parnell, J. 90/704 (K).

Saraburi: Phraphutthabath, Khunkhol Tham Makak, 50 m, 20 Nov 2005, Pooma, R.

et al. 5719 (E); Phraphutthabath, 22 Sep 1999, Wongprasert, T. s.n. (BKF).

ACKNOWLEDGEMENTS. The authors wish to thank the curators of the herbaria for the loans

of specimens; Mr Preecha Karaket, Dr Somran Suddee, Dr Pramote Triboun, Dr Thammarat

_ V

Phutthai and Dr Naiyana Tetsana for the photographs; Dr Jana Leong-Skornickova for help

with the figures; the staff of the Forest Herbarium Bangkok (BKF) for their assistance in the

field; the Davis Fund for supporting field work; and the curators and staff of the Royal Botanic

A revision of Microchirita in Thailand

283

Garden Edinburgh, Singapore Botanic Gardens and the Forest Herbarium Bangkok for other

help. This study was funded by the Sibbald Trust of the Royal Botanic Garden Edinburgh (UK),

the Garden City Fund (Singapore), the Carlsberg Foundation (Denmark) and the National Parks

Board (Singapore).

References

Applequist, W.F. (2016). Report of the Nomenclature Committee for Vascular Plants: 68.

Taxon 65: 1153-1165.

Burtt, B.F. (1954). Studies in the Gesneriaceae of the Old World II. Notes Roy. Bot. Gard.

Edinburgh 21: 193-208.

Burtt, B.F. (1960). Anew Chirita from Thailand. J. Roy. Hort. Soc. 85: 27-28.

Burtt, B.F. (2001). Flora of Thailand: Annotated checklist of Gesneriaceae. Thai Forest Bull,

Bot. 29: 81-109.

Craib, W.G. (1926). Contributions to the Flora of Siam. Additamentum XVIII. Bull. Misc.

Inform. Kew: 154-174.

Craib, W.G. (1930). Contributions to the Flora of Siam. Additamentum XXVII. Bull. Misc.

Inform. Kew: 161-174.

Hilliard, O.M. (2004). A revision of Chirita sect. Liebigia (Gesneriaceae). Edinburgh J. Bot.

60: 361-387.

IUCN (2012). IUCN Red List Categories and Criteria: Version 3.1, 2 nd ed. Gland, Switzerland

and Cambridge, UK: IUCN.

Middleton, D.J. & Moller, M. (2012). Tribounia, a new genus of Gesneriaceae from Thailand.

Taxon 61: 1286-1295.

Middleton, D.J. & Puglisi, C. (2015). Proposal to conserve the name Chirita hamosa

(Microchirita hamosa) (Gesneriaceae) with a conserved type. Taxon 64: 1328-1329.

Middleton, D.J. & Triboun, P. (2012). Somrania, a new genus of Gesneriaceae from Thailand.

Thai Forest Bull., Bot. 40: 9-13.

Middleton, D.J. & Triboun, P. (2013). New species of Microchirita (Gesneriaceae) from

Thailand. Thai Forest Bull, Bot. 41: 13-22.

Middleton, D.J., Atkins, H., Truong, F.H., Nishii, K. & Moller, M. (2014). Billolivia, a new

genus of Gesneriaceae from Vietnam with five new species. Phytotaxa 161: 241-269.

Middleton, D.J., Nishii, K., Puglisi, C., Forrest, F. & Moller, M. (2015). Chayamaritia

(Gesneriaceae: Didymocarpoideae), a new genus from Southeast Asia. PI Syst. Evol

301(7): 1947-1966.

Moller, M., Forrest, A., Wei, Y.G. & Weber, A. (2011). A molecular phylogenetic assessment

of the advanced Asiatic and Malesian didymocarpoid Gesneriaceae with focus on

nonmonophyletic and monotypic genera. PI Syst. Evol. 292(3-4): 223-248.

Moller, M., Chen, W.H., Shui, Y.M., Atkins, H.J. & Middleton, D.J. (2014). A new genus of

Gesneriaceae in China and the transfer of Briggsia species to other genera. Gard. Bull

Singapore 66(2): 195-205.

Moller, M., Nishii, K., Atkins, H.J., Kong, H.H., Kang, M., Wei, Y.G., Wen, F., Hong, X. &

Middleton, D.J. (2016). An expansion of the genus Deinostigma (Gesneriaceae). Gard.

Bull Singapore 68(1): 145-172.

Puglisi, C. & Middleton, D.J. (2017). A revision of Damrongia (Gesneriaceae) in Thailand.

Thai Forest Bull., Bot. 45: 79-93.

284

Card. Bull. Singapore 69(2) 2017

Puglisi, C., Middleton, D.J., Triboun, P. & Moller, M. (2011). New insights into the relationships

between Paraboea, Trisepalum and Phylloboea (Gesneriaceae) and their taxonomic

consequences. Taxon 60: 1693-1702.

Puglisi, C., Yao, T.L., Milne, R., Moller, M., & Middleton, D.J. (2016). Generic recircumscription

in the Loxocarpinae (Gesneriaceae), as inferred by phylogenetic and morphological

data. Taxon 65: 277-292.

Punekar, S.A. & Lakshminarasimhan, P. (2009). Chirita sahyadriensis (Gesneriaceae), a new

species from Western Ghats, India. Folia Malaysiana 10: 17-22.

Rafidah, A.R. (2017). Taxonomy and conservation status of Microchirita (Gesneriaceae) in

Peninsular Malaysia. Gard. Bull. Singapore 69: 1-31.

Rafidah, A.R. & Haron, N.W. (2013). Microchirita ruthiae sp. nov. (Gesneriaceae) from

Peninsular Malaysia. Nord. J. Bot. 31: 612-615.

Rafidah, A.R., Kiew, R. & Haron, N.W. 2011. To which genus does the enigmatic Peninsular

Malaysia Chirita data (Gesneriaceae) belong? Blumea 56: 18-20.

Thiers, B. (2017, 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 12 July 2017.

Wang, Y.-Z., Mao, R.-B., Liu, Y., Li, J.-M., Dong, Y., Li, Z.-Y. & Smith, J. (2011). Phylogenetic

reconstruction of Chirita and allies (Gesneriaceae) with taxonomic treatments. J. Syst.

Evol. 49: 50-64.

Weber, A. (1975). The cristate inflorescence of Chirita sect. Microchirita (Gesneriaceae). Notes

Roy. Bot. Gard. Edinburgh 34: 221-230.

Weber, A., Middleton, D.J., Forrest, A., Kiew, R., Lim, C.L., Rafidah, A., Triboun, P, Wei, Y.G.,

Yao, T.L., Sontag, S. & Moller, M. (2011a). Molecular systematics and remodelling of

Chirita and associated genera (Gesneriaceae). Taxon 60: 767-790.

Weber, A., Wei, Y.G., Puglisi, C., Mayer, V. & Moller, M. (2011b). A new definition of the

genus Petrocodon (Gesneriaceae). Phytotaxa 23: 49-67.

Wei, Y.G., Fang, W., Chen, W.H., Shui, Y.M. & Moller, M. (2010). Litostigma, a new genus from

China: A morphological link between basal and derived didymocarpoid Gesneriaceae.

Edinburgh J. Bot. 67: 161-184.

Wood, D. (1972). Studies in the Gesneriaceae of the Old World XXXV: new species and

combination in Chirita. Notes Roy. Bot. Gard. Edinburgh 31: 367-371.

Wood, D. (1974). A revision of Chirita (Gesneriaceae). Notes Roy. Bot. Gard. Edinburgh 33:

123-205.

Gardens' Bulletin Singapore 69(2): 285-293. 2017

doi: 10.26492/gbs69(2).2017-07

285

Ceropegia laotica (Apocynaceae, Asclepiadoideae):

the first new species of Ceropegia described from Laos

M. Rodda 1 & U. Meve 2

herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, 259569 Singapore

rodda.michele@gmail.com

department of Plant Systematics, University of Bayreuth,

95440 Bayreuth, Germany

ulrich.meve @ uni-bayreuth.de

ABSTRACT. A new species, Ceropegia laotica Rodda & Meve, from the Bolaven Plateau in

southern Laos, is described and illustrated. It shows a twining habit, linear-lanceolate leaves

and possesses small root tubers. The new species is compared with Ceropegia cochleata

Kidyoo from Thailand.

Keywords. Ceropegia cochleata, Ceropegia digitiformis, Ceropegieae, Indochinese Peninsula,

Southeast Asia

Introduction

In the traditional, morphology-based circumscription of Ceropegia L. the genus includes

only taxa possessing pitfall flowers that trap small visiting flies for 24 hours in funnel-

shaped, basally inflated corollas. Molecular-based studies have shown, however, that

the phylogeny of the terminal branch of the tribe Ceropegieae, the Stapeliinae, to which

Ceropegia belongs, is complex. The other genera of Stapeliinae (. Brachystelma R.Br.

and the stapeliads) are all nested and scattered through the Ceropegia tree, making

Ceropegia multiply paraphyletic (Meve & Liede-Schumann, 2007; Surveswaran et

al., 2009; Bruyns et al., 2015). These studies have documented that complex pitfall

flowers - always thought to represent the peak of evolution in the Ceropegieae

- have been independently lost several times during evolution to be replaced by

open, radiate flowers. However, in order to prevent severe taxonomic instability in

an extremely species-rich group (Stapeliinae comprises around 700 species), Meve

& Liede-Schumann (2007) suggested that paraphyly in Ceropegia / Stapeliinae be

accepted and that the “Ceropegia pitfall flower” be retained as a diagnostic character

for Ceropegia. However, recently, Bruyns et al. (2017) proposed about 400 new

names and combinations in Ceropegia in order to make it monophyletic. As long as

the driving forces of the complex evolution in this group are not at least minimally

understood, we prefer to retain the traditional classification, keeping Brachystelma and

the stapeliad genera separate. There are around 210 “true” Ceropegia species of small

erect or twining herbs which occur widely in the Old World tropics and subtropics of

286

Card. Bull. Singapore 69(2) 2017

Southeast Asia, India, Arabia, Africa, Madagascar, reaching east to New Guinea and

Northern Australia and west to the Canary Islands (with two species). The diversity

within Ceropegia, its wide distribution, and the attractive flowers were and are

fascinating. Many floristic, often geographically focused works, have been produced

for places such as India (Hooker, 1883; Ansari, 1984), China (Li et al., 1995), Arabia

(Bruyns, 1988), Madgascar (Meve & Liede, 1994), Southern Africa (Dyer, 1980),

and Tropical East Africa (Masinde, 2012). Only one complete revision of the genus

has been undertaken (Huber, 1957), although a wide-ranging treatment (lacking the

non-succulent taxa) was published more recently by Meve (2002). China (Li et al.,

1995) and India (Ansari, 1984) are by far the diversity hotspots in Asia. Without doubt

Asia is the centre of origin of Ceropegia (Bruyns et al., 2015; Meve et al., 2017). In

Southeast Asia, however, the number of taxa is considerably lower. Myanmar has nine

species, according to the last compilation by Kress et al. (2003) and the inclusion of

Ceropegia farrokhii McCann in C. kachinensis Prain by Singh et al. (2015), and in

Vietnam just three species are recorded (Costantin, 1912; Vietnam Plant Data Center:

http://www.botanyvn.com). However, recent research on Ceropegia in Thailand has

led to the discovery of numerous new species bringing the count for the country to

ten (Kerr, 1951; Boonjaras & Thaithong, 2003; Meve, 2009; Kidyoo, 2014a, 2014b,

2015a, 2015b; Kidyoo & Paliyavuth, 2017). Finally, in Laos, only one unidentified

Ceropegia species has so far been recorded (Newman et al., 2007). Recent fieldwork

in southern Laos by the first author has now led to the discovery of an unknown species

of Ceropegia that is here described and illustrated as a new taxon.

Taxonomy

Ceropegia laotica Rodda & Meve, sp. nov.

Similar to Ceropegia cochleata Kidyoo in trailing to twining annual growth habit,

presence of small tubers, leaves more or less lanceolate and small, and in habitat

preferences: sandstone rocks with sparse soil. Ceropegia laotica can be separated

from C. cochleata because the former has corolla lobes broadly ovate and folded

while the latter has corolla lobes long-tailed and twisted. - TYPE: Laos, Champasak

Province, Paksong, Dong Hua Sao NBCA, Bolaven Plateau, Ban Hnongluang Wat,

15°07'57"N 10°20'57"E, 1258 m, 23 October 2015, Rodda, M., Tan, L., Guglielmone,

L., Peereman, J.M.E., Souvannaphou, S. & Soulimoungichoun, O. MR1418 (holotype

SING; isotypes FOF, Herbarium of Pha Tad Ke Botanic Garden, TO). (Fig. 1, 2)

Perennial climbing herb with 1 or 2 stems from the base, twining, with clear sap in all

parts. Rootstock tuberous; tubers subglobose, slightly apically compressed, smooth,

medium brown, 2-3 cm in diameter, with many additional, straight, fleshy roots.

Stems rarely branched, terete, glabrous, to 2 m long, 1.5-2.5 mm in diameter, greenish

red to maroon red; intemodes (3—)5—10(—15) cm long. Leaves spreading, petiolate,

petiole terete, slightly channelled above, 5-7 x 1-1.5 mm in diameter, pubescent

only above, trichomes curved, 100-200 pm long; lamina linear(-lanceolate), (4-)5-

New Ceropegia from Laos

287

Fig. 1 . Ceropegia laotica Rodda & Meve. A. Single flower. B. Corolla, longitudinal section

with corona removed. C. Corona, side view. D. Corona, top view. E. Corona, from underneath.

F. Pollinarium. A-C & F based on M. Rodda et al. MR1418, D, E based on M. Rodda et al.

MR1802. (Photos: M. Rodda)

288

Card. Bull. Singapore 69(2) 2017

Fig. 2. Ceropegia laotica Rodda & Meve. A. Fruiting branch. B. Plant in situ, climbing over

bamboo. C. The exposed tuber and roots after removing a thin layer of leaf litter. (Photos: M.

Rodda)

New Ceropegia from Laos

289

10(—12) x 0.5-1.2 cm, slightly fleshy but flexible, base attenuate, apex acute, pale

green above, paler underneath, sparsely pubescent above, underneath pubescent along

main vein only, trichomes curved, 100-200 pm long, older leaves glabrous; venation

pinnate, lateral veins obscure; colleters 1 or 2 at base of lamina, globose, 200-300

pm in diameter. Inflorescences 1-2 flowered, with only one flower open at a time,

peduncles extra-axillary, one per node, 0.5-2 x 1.5-2 mm in diameter, reddish green,

glabrous; pedicels erect, 8—15(—25 when fruiting) x c. 1.5 mm in diameter, reddish

green to whitish, glabrous or very sparsely pubescent; bracts 1 or 2 at base of pedicel,

lanceolate, 1-2 mm long, reddish, apex acute. Calyx lobes narrowly lanceolate, 4-5

x 0.7-1 mm, glabrous, apex acuminate, reddish, glabrous; with one colleter at each

sinus, conical, 400-500 x 150-200 pm, acute. Corolla 35—45 mm in total length, tube

upright, straight or slightly curved, glabrous, with a bulbous base, a cylindrical mid

portion and funnel shaped apex; tube (20-)23-27 mm long, base 5-7 mm in diameter,

cylindrical part 2.5-3.5 mm in diameter; base white and densely spotted red outside,

inside white and striped red; cylindrical part red with paler dots outside, dark red and

striped maroon inside; apex red and spotted green outside, cream and striped green

inside; corolla lobes broadly ovate, folded, (12-) 15-17 x 7-8.5 mm, basal part cream-

yellow, ciliate, apex acute, connate at tips, green and pubescent and ciliate inside,

maroon red and glabrous outside. Gynostegium stipitate, stipe 0.4-1.6 x 0.9-1.1 mm

in diameter, white, glabrous. Gynostegial corona of staminal and interstaminal parts,

interstaminal lobes joined to form a shallow cup, 4.5-5 mm in diameter, purple with

a white base; free lobes deeply bifid with the tips of adjacent lobules touching each

other, ascending-erect, small triangular, c. 1.5 x 1 mm, maroon red, inside pilose

with spreading translucent-white trichomes c. 0.5 mm long, staminal corona lobes

connivent-erect, linear, terete, 1—1.2 x 0.25-0.35 mm, pinkish white. Pollinaria:

pollinium broadly ovoid, yellow, 350^100 x 350^100 pm, caudicles c. 200 pm long,

corpusculum clavate, reddish brown, 250-300 x 90-110 pm. Ovary narrowly conical,

2.1-2.3 x 0.5-0.7 mm wide at the base, apex truncate, glabrous. Mericarps held erect,

linear, slightly curved, c. 11.5 x 3 mm (immature), reddish green. Seeds not seen.

Distribution. Laos, Champasak Province, in the vicinity of Ban Hnongluang Wat.

Habitat and ecology. Ceropegia laotica is a perennial tuberous herb found in a single

population where all individuals were rooted in very thin soils (3-6 cm thick) over

large sandstone boulders partially exposed. The plants were growing among grasses

and climbing over gingers, bamboo and Ficus spp. at the edge between open grassland

vegetation and short stunted forest (Fig. 3).

Provisional IUCN conservation assessment. Endangered EN Blab(iii) + B2ab(iii)

using the guildelines from the IUCN Standards and Petitions Subcommittee (2017).

Ceropegia laotica is known only from the type locality. A thorough survey of the

Bolaven Plateau is lacking and we have little information on similar habitats in

neighbouring Thailand and Vietnam for a conclusive conservation assessment.

If, however, we estimate the EOO as the entire Bolaven Plateau (< 5000 km 2 ) and

290

Card. Bull. Singapore 69(2) 2017

Fig. 3. The habitat of Ceropegia laotica Rodda & Meve in the vicinity of Ban Hnongluang Wat,

Laos. Open grassland vegetation on very thin soils with exposed bedrock. (Photo: M. Rodda)

consider that only a small part of it has suitable rocky habitat for the species (AOO <

500 km 2 ) and also take into account that the area is heavily affected by habitat loss due

to land conversion to agriculture, mining and dams (Delang et al., 2013), a provisional

conservation status can be assessed as endangered pending more information which

might increase the threat status further.

Additional specimens examined. Ceropegia laotica. Cultivated plant grown on from Laos,

Champasak Province, Paksong, Dong Hua Sao NBCA, Bolaven Plateau, Ban Hnongluang Wat,

15°07'57"N 10°20'57"E, 1258 m, 23 October 2015, Rodda, M. et al. MR1418, vouchered at

Singapore Botanic Gardens on 16 May 2017 as Rodda, M. MR1802 (SING).

Notes. The type of Ceropegia thorelii Constantin was collected from the border region

of Cambodia and Laos. It was treated by Huber (1957) as Ceropegia macrantha Wight

var. thorelii (Constantin) H.Huber and is a lowland species only reported from St(e)

ung Treng (Cambodia, Mekong River). The type specimen in P ( Thorel 2148 ) is a fairly

sturdy plant with elliptical leaves, pedunculate inflorescences and a corolla which is

coarsely brownish spotted. Hence, Huber (1957) placed it in Ceropegia macrantha

despite its hairy peduncles and pedicels, as did Meve (2002) who included C. thorelii

as a full synonym of C. macrantha. Interestingly, a later collection made at Steung

Treng (cf. Maxwell, 2009: fig. 14) has a corolla superficially comparable to the type

of C. thorelii , but the stems and leaves are densely pilose, the leaves seem to be rather

New Ceropegia from Laos

291

small and, most importantly, the basal inflation of the corolla tube (ostiolum) is basally

constricted. This latter character is significant and of high taxonomic value in, for

example, African Ceropegia. Therefore, Maxwell's (2009) determination as C. thorelii

cannot be confirmed here. Instead, although the corolla of Ceropegia cochleata Kidyoo,

with its long-tailed, twisted lobes, does not show much similarity with the relatively

short and obtuse corolla lobes of the new species, Ceropegia cochleata and C. laotica

are vegetatively, ecologically and geographically much closer. They share the trailing

to twining (when support is available) habit and growth form; they have wiry stems

and spreading, fairly small, more or less lanceolate leaves; they have similar habitat

preferences as they both occur on sandstone rocks with sparse soil and vegetation

cover dominated by grasses and mosses (Kidyoo, 2015b); both species develop small

tubers; and the above-ground parts do not survive the dry season.

There are also a number of similarities in habit, ecology and flower morphology

between Ceropegia laotica and C. digitiformis Kidyoo. The latter species is found in

Thailand in the Phu Wua Wildlife Sanctuary and Phu Langka National Park, c. 400 km

northwest of the type locality of C. laotica. However, although Ceropegia digitiformis is

also deciduous, the aerial stems survive the dry season because they are fleshy (Kidyoo

& Paliyavuth, 2017), and not wiry as in C. laotica. Also the leaves are laminar, elliptic,

ovate to rounded in Ceropegia digitiformis (and not rather linear and fleshy as in C.

laotica ) - displaying a close relationship to the Indian C. thwaitesii Hook. Kidyoo &

Paliyavuth (2017) distinguish Ceropegia digitiformis from C. thwaitesii only on small

deviations in corona morphology. In addition, Ceropegia digitiformis is unlikely to be

a sister species of C. laotica because of the pollinarium morphology. The shape of the

pollinium in Ceropegia digitiformis is ovoid with the germination crest subapical (cf.

Kidyoo & Paliyavuth, 2107: figs. IF and 2E), whereas the pollinia in C. laotica (and

C. cochleata ) are broadly ovate to (sub)rectangular with the pellucid margin along the

inner side of the pollinium (Fig. 2f). On the other hand, Ceropegia digitiformis and C.

laotica share the character of a stipitate corona although the interstaminal lobules in

C. laotica are not linear-lanceolate as in C. digitiformis, but rather triangular and much

more similar to those of C. cochleata, including in indumentum.

To conclude, Ceropegia laotica, C. cochleata and C. digitiformis all inhabit

rocky, montane habitats on sandstone with sparse vegetation in the central Indochinese

Peninsula. They share a number of morphological similarities but can each be

distinguished by unique morphological characters. In the important habit, leaf and

corona/gynostegial characters, we regard Ceropegia cochleata as presumably most

closely related to our new species.

ACKNOWLEDGEMENTS. This study is part of an on-going research project on the

systematics of Asian Apocynaceae. Financial support was received from the National Parks

Board (Singapore) which sponsored numerous field trips to Laos and herbarium study trips

to Asian and European institutions. The curators of the E, P and SING herbaria are thanked

for allowing access and/or for allowing loans or high quality images of herbarium specimens.

We thank two anonymous reviewers and Dr David Middleton, Editor of Gardens’ Bulletin

Singapore, for suggesting improvements to this manuscript.

292

Card. Bull. Singapore 69(2) 2017

References

Ansari, M.Y. (1984). Fascicles of Flora of India 16. Asclepiadaceae - Genus Ceropegia.

Calcutta: Botanical Survey of India.

Boonjaras, T. & Thaithong, O. (2003). Ceropegia hirsuta (Asclepiadaceae), a new record for

Thailand. Thai Forest Bull., Bot. 31: 1-6.

Bruyns, P. V. (1988). Studies in the flora of Arabia XXIV: The genus Ceropegia in Arabia. Notes

Roy. Bot. Gard. Edinburgh 45: 287—326.

Bruyns, P.V., Klak, C., & Hanacek, P. (2015). Recent radiation of Brachystelma and Ceropegia

(Apocynaceae) across the Old World against a background of climatic change. Molec.

Phylogenet. Evol. 90: 49—66.

Bruyns, P.V., Klak, C. & Hanacek, P. (2017). A revised, phylogenetically-based concept of

Ceropegia (Apocynaceae). S. African J. Bot. 112: 399^-36.

Costantin, J.N. (1912). Asclepiadacees. In: Lecomte, H. (ed.) Flore generate de Vhidochine,

vol. 4, pp. 1-154. Paris: Masson & Co.

Delang, C.O., Toro, M. & Charlet-Phommachanh, M. (2013). Coffee, mines and dams: conflicts

over land in the Bolaven Plateau, southern Lao PDR. Geogr. J. (London) 179: 150-164.

Dyer, R.A. (1980). Asclepiadaceae (Brachystelma, Ceropegia, Riocreuxia). In: Leistner, O.A.

(ed.) Flora of Southern Africa, vol. 27, part 4. Rotterdam: A. A. Balkema.

Hooker, J.D. (1883). Asclepiadeae In: Hooker, J.D. (ed.) The Flora of British India, vol. 4, pp.

1-78. London: L. Reeve.

Huber, H. (1957). Revision der Gattung Ceropegia. Mem. Soc. Brot. 12: 1-203.

IUCN Standards and Petitions Subcommittee (2017). Guidelines for Using the IUCN Red List

Categories and Criteria, ver. 13.

http://www.iucnredlist.org/documents/RedListGuidelines.pdf. Accessed 4 Sep. 2017.

Kerr, A.F.G. (1951). Ceropegia L. In: Pendleton, R.L. (ed.) Florae Siamensis Enumeratio, vol.

3, pp. 49-51. Bangkok: Bangkok Times Press.

Kidyoo, M. (2014a). Ceropegia suddeei sp. nov. (Apocynaceae, Asclepiadoideae) from

northeastern Thailand. Nord. J. Bot. 32: 569-574.

Kidyoo, M. (2014b). Two new species of Ceropegia (Apocynaceae, Asclepiadoideae) from

eastern Thailand. Phytotaxa 162: 91-98.

Kidyoo, M. (2015a). Ceropegia tribounii (Apocynaceae, Asclepiadoideae), a new species from

western Thailand. Phytotaxa 205: 59-64.

Kidyoo, M. (2015b). Ceropegia cochleata sp. nov. (Apocynaceae, Asclepiadoideae) from

Thailand. Nord. J. Bot. 33: 668-672.

Kidyoo, M., & Paliyavuth, C. (2017). Ceropegia digitiformis sp. nov. (Apocynaceae,

Asclepiadoideae) from northeastern Thailand. Taiwania 62(1): 24-28.

Kress, W. J., DeFilipps, R.A., Farr, E. & Yin Yin Kyi, D. (2003). A checklist of the trees, shrubs,

herbs and climbers of Myanmar. Washington: Smithsonian Institution.

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

(eds) Flora of China, vol. 16, pp. 189-270. Beijing: Science Press & St. Louis: Missouri

Botanical Garden.

Masinde, S. (2012). Ceropegia L. In: Beentje, H.J. (ed.) Flora of Tropical East Africa,

Apocynaceae II, pp. 115-530. Rotterdam: A.A. Balkema.

Maxwell, J.F. (2009). Vegetation and Vascular Flora of the Mekong River, Kratie and Steung

Treng Provinces, Cambodia. Maejo Int. J. Sci. Techn. 3(1): 143-211.

Meve, U. (2002). Ceropegia. In: Albers, F. & Meve, U. (eds) Illustrated Handbook of Succulent

Plants: Asclepiadaceae, pp. 63-106. Berlin: Springer.

New Ceropegia from Laos

293

Meve, U. (2009). Ceropegia thailandica (Asclepiadoideae-Ceropegieae), a spectacular new

Thai species. Bradley a 27: 161-164.

Meve, U. & Liede, S. (1994). A conspectus of Ceropegia L. (Asclepiadaceae) in Madagascar,

and the establishment of C. sect. Dimorpha. Phyton (Horn) 34: 131-141.

Meve, U. & Liede-Schumann, S. (2007). Ceropegia (Apocynaceae, Ceropegieae, Stapeliinae):

paraphyletic, but still taxonomically sound. Ann. Missouri Bot. Gard. 94: 392-406.

Meve, U., Heiduk, A. & Liede-Schumann, S. (2017). Origin and early evolution of Ceropegieae

(Apocynaceae-Asclepiadoideae). Syst. Biodivers. 15: 143-155.

Newman, M., Ketphanh, S., Svengsuksa, B., Thomas, P., Sengdala, K., Lamxay, V.V. &

Armstrong, K. (2007). Checklist of the Vascular Plants of Lao PDR. Edinburgh: Royal

Botanic Garden Edinburgh.

Singh, R.K., Garg, A. & Singh, P.K. (2015). Lectotypification and anew synonym of Ceropegia

kachinensis (Apocynaceae). Phytotaxa 197: 215-221.

Surveswaran, S., Kamble, M.Y., Yadav, S.R., & Sun, M. (2009). Molecular phylogeny of

Ceropegia (Asclepiadoideae, Apocynaceae) from Indian Western Ghats. PI. Syst. Evol.

281: 51-63.

Gardens’ Bulletin Singapore 69(2): 295-305. 2017

doi: 10.26492/gbs69(2).2017-08

295

Two new species of Oreocharis (Gesneriaceae)

from Northwest Vietnam

W.H. Chen 1 , D J. Middleton 2 , H.Q. Nguyen 3 , H.T. Nguyen 3 , L.V. Averyanov 4 , R.Z.

Chen 1 , K.S. Nguyen 5 , M. Moller 6 & Y.M. Shui 1

‘Key Laboratory for Plant Diversity and Biogeography of East Asia,

Kunming Institute of Botany, Chinese Academy of Sciences,

132 Lanhei Road, Kunming 650201, Yunnan, China

y mshui @ mail .kib .ac.cn

2 Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, 259569 Singapore

3 Center for Plant Conservation of Vietnam (CPC),

Vietnam Union of Science and Technology Associations,

25/32 Lane 191, Lac Long Qua Road, Hanoi, Vietnam

4 Botanical Institute of the Russian Academy of Science,

Prof. Popov Street, 2, RU-197376, St Petersburg, Russia

institute of Ecology and Biological Resources,

Vietnam Academy of Science and Technology,

18 Hoang Quoc Viet Road, Hanoi, Vietnam

6 Royal Botanic Garden Edinburgh, 20A Inverleith Row,

Edinburgh EH3 5LR, UK

ABSTRACT. Two new species of Oreocharis (Gesneriaceae), O. argyrophylla W.H.Chen,

H.Q.Nguyen & Y.M.Shui and O. blepharophylla W.H.Chen, H.Q.Nguyen & Y.M.Shui, from

the Xuan Nha nature reserve, Van Ho district, Son La province, in northwestern Vietnam are

described. They are compared to their most similar species and diagnostic characteristics are

provided.

Keywords . China, corolla colour, Indo-China Peninsula, Thailand

Introduction

The genus Oreocharis Benth. in the Gesneriaceae has recently been greatly enlarged

by the inclusion of a number of other genera (Moller et al., 2011). Most of the

approximately 113 species are found in China, with a few species also in north-eastern

India, northern Myanmar, northern Thailand and northern Indo-China (Moller et

al., 2011, 2015, 2016; W.H. Chen et al., 2014, 2016; Cai et al., 2015; Yang et al.,

2015, 2017; Wei et al., 2016; R.Z. Chen et al., 2017; Do et al., 2017; Li et al., 2017).

A recent botanical exploration in north-western Vietnam near the border to Laos

resulted in collections of two unknown Oreocharis species. After consultation of the

literature (Pellegrin, 1930; Wang et al., 1990, 1998; Ho, 2000; Li & Wang, 2004) and

examination of type specimens, as well as comparison to other species in the genus,

296

Card. Bull. Singapore 69(2) 2017

we propose two new species: Oreocharis argyrophylla W.H.Chen, H.Q.Nguyen &

Y.M.Shui (Fig. 1, 2) and O. blepharophylla W.H.Chen, H.Q.Nguyen & Y.M.Shui (Fig.

3, 4). The former species is most similar to Oreocharis argyreia Chun ex K.Y.Pan and

O. hirsuta Barnett, and the latter to O. magnidens Chun ex K.Y.Pan and O. curvituba

J.J.Wei & W.B.Xu, but differ from these species in the characters listed in Tables 1 and

2. The two new species occur in close proximity to each other in the Van Ho district in

Son La province in north-western Vietnam.

Below, CPC refers to the Herbarium of the Center for Plant Conservation, Vietnam

Union of Science and Technology Associations, Hanoi.

Oreocharis argyrophylla W.H.Chen, H.Q.Nguyen & Y.M.Shui, sp. nov.

The new species is most similar in leaf characteristics to Oreocharis argyreia Chun

ex K.Y.Pan but differs from it in having yellow flowers (vs purple), corolla narrowly

infundibuliform (vs tubiform) and 2.5-3.1 cm long (vs 2-2.3 cm), and coherent anthers

(vs free). In flower characteristics it is also similar to Oreocharis hirsuta Barnett in

its yellow and tubular corolla but differs from it in its deeply bilabiate corolla limb

(vs shallowly bilabiate) and coherent anthers (vs free). - TYPE: Vietnam, Son La

province, Van Ho district, Tan Xuan municipality, Cot Moc village, territory of Xuan

Nha nature reserve, eastern slopes of Pha Luong Mountain, at an elevation of 1000-

1400 m, 20°40'33.3"N 104°39'00.3"E, 15 November 2013, in flower, Averyanov, L.,

Hiep, N.T., Khang, N.S., Thang, N.D. & Qui, L.D. CPC 7175 (holotype KUN; isotypes

CPC, LE). (Fig. 1, 2)

Perennial herb with very short stem. Leaves numerous, rosulate, petiolate; petiole

4-9 cm, densely appressed pubescent; blade narrowly obovate to broadly lanceolate,

12-18 x 4.5-6.5 cm, both surfaces densely pubescent with appressed silvery hairs

especially adaxially, base narrowly cuneate, apex acute to short acuminate, margin

serrulate above middle of leaf blade, lateral veins 6-8 on each side of midrib, distinct.

Inflorescences axillary, 3^1-branched, cymes 3-many-flowered; peduncle 8-12 cm

long, pubescent with appressed hairs; bracts 2, linear-lanceolate, 10-13 x 1-2 mm,

apex tapering, margin entire, pubescent and villous abaxially, glabrous adaxially;

pedicels 0.7-1.8 cm long, white pubescent. Calyx of 5 lobes free to base; lobes

equal, linear, 6-8 x c. 1 mm, margin entire, pubescent with appressed hairs abaxially,

glabrous adaxially. Corolla yellow, tubular, apically infundibuliform, 2.5-3.1 cm long,

outside glandular pubescent, glabrous inside; tube gradually slightly ampliate from

the middle, 1.6-2.2 cm long, 2.8-3 mm in diam. at base, 6-6.2 mm in diam. and not

constricted at the throat; limb 2-lipped; adaxial lip 2-lobed, dissected from near base

of the adaxial lip, lobes ovate, apices rounded or obtuse, 5.8-6 x 4.1-4.3 mm; abaxial

lip 3-lobed, lobes ovate, apices rounded or obtuse, subequal, median lobe slightly

larger, 7.4-7.5 x 6.1-6.2 mm, lateral lobes 7-7.1 x 4.4^1.7 mm. Stamens 4, anthers

coherent in pairs, included, adaxial stamens 8-9 mm long, adnate to corolla tube

1.5-1.6 cm from base, abaxial stamens 1.4-1.5 cm long, adnate to corolla tube 5-5.5

mm from base; filaments glabrous; anthers basifixed, subglobular, 2-locular, dehiscing

Two new species of Oreocharis in Vietnam

297

Fig. 1 . Oreocharis argyrophylla W.H.Chen, H.Q.Nguyen & Y.M.Shui. A. Habitat. B. Habit

(scale bar 4 cm). C. Inflorescences (scale bar 3 cm). D. Flowers (scale bar 2 cm). (Photos: S.K.

Nguyen)

longitudinally; staminode 1.4-1.5 mm long, adnate to corolla tube 4-5 mm from base.

Disc ring-like, c. 1.2 mm long, shallowly 5-lobed. Pistil 1.5-1.6 cm long, glabrous;

ovary fusiform, 1.1-1.4 cm long, glabrous, unilocular; style glabrous, 4-5 mm long;

stigma 1, peltate. Capsule straight, fusiform, loculicidally dehiscent, 3.3-4 cm long,

with persistent style 5-6 mm long. Seeds fusiform, 5-6 x c. 2 mm.

Etymology. The specific epithet refers to the silvery leaf indumentum.

Ecology, distribution and phenology. The species grows as a lithophytic herb on large

moss covered boulders in remnants of primary and secondary broad-leaved evergreen

298

Card. Bull. Singapore 69(2) 2017

10 cm

1 cm

Fig. 2. Oreocharis argyrophylla W.H.Chen, H.Q.Nguyen & Y.M.Shui. A. Flowering plant. B.

Dissected corolla showing two pairs of stamens cohering at the anther tips. C. Pedicel, calyx,

disc and pistil (mature). Drawn by Ling Wang from the holotype Averyanov, L. et al. CPC 7175.

Two new species of Oreocharis in Vietnam

299

Table 1. Characteristics distinguishing Oreocharis argyrophylla, O. argyreia and O. hirsuta.

Characters

O. argyrophylla

O. argyreia

O. hirsuta

Leaf hairs

Densely appressed

silvery pubescent

Densely appressed

silvery pubescent

Sparsely spreading

hirsute

Corolla

Yellow, narrowly

infundibuliform, 2.5-

3.1 cm

Blue-purple to purplish

white, nearly cylindrical,

2-2.3 cm

Yellow, nearly

cylindrical, 2-2.9 cm

Corolla tube

1.7-2.2 cm long, 2.8-

3 mm in diam. at base,

6-2.8 mm in diam. at

throat

1.5-2 cm long, almost

0.5-0.6 cm in diam. from

base to top

1.6-1.9 cm long, 4-5

mm in diam. from base

to top

Corolla limb

Distinctly 2-lipped,

lips unequal;

Slightly 2-lipped, lips

subequal;

Weakly 2-lipped, lips

unequal.

Adaxial lip

Lobes oblong, 5.8-6 x

4.1-4.3 mm

Lobes suborbicular, 2-

3 mm in diam.

Lobes ovate, 7.6-8 x c.

5 mm

Abaxial lip

Lobes oblong, subequal,

middle 7.4-7.5 x 6.1-6.2

mm, lateral lobes 7-7.1

x 4.4-4.7 mm

Lobes suborbicular,

equal, 2-3 mm in diam

Lobes ovate, subequal,

middle 7.6-8 x

6.7-7 mm, lateral lobes

7.2-7.7 x 6-6.1 mm.

Anthers

Coherent in 2 pairs

Not coherent

humid forests on very steep mountain slopes composed of shale and sandstone at

elevations of 1000-1850 m. This species is only known from the Xuan Nha nature

reserve, Son La province of north-western Vietnam, where it is common. Flowering in

September-December and fruiting in October-December.

Provisional IUCN conservation status. Endangered EN Blab(ii,iii,v) + B2ab(ii,iii,v),

following IUCN (2012, 2016) guidelines. This is based on an EOO of < 5000 km 2

and an AOO of < 500 km 2 , being known from fewer than five populations, and with

evidence of a reduction in the area of occupancy, a decline in habitat quality, and a

decline in the number of individuals due to deforestation, largely for agriculture and

due to fire.

Additional specimens examined. VIETNAM. Son La: Van Ho district, Chieng Xuan municipality,

Co Hong village, territory of Xuan Nha nature reserve, Pha Luong Mountain, 13 Nov 2013, in

fruit, Averyanov, L. et al. CPC 7020 (CPC, KUN, LE); Moc Chau Distr., Chieng Son comm.,

Pha Luong vill., Pha Luong Mountains, 1400-1500 m a.s.L, 20°41'33.2"N 104°37'37.0"E, 22

Sep 2016, Averyanov, L. et al. CPC 7939 (CPC, LE); Moc Chau Distr., Chieng Son comm.,

Pha Luong vill., Pha Luong Mountain, 20°40'23.0"N 104°37'52.0"E, 1750-1850 m a.s.L, 23

Sep 2016, Averyanov, L. et al. CPC 7993 (CPC, LE); Moc Chau Distr., Chieng Son comm., Pha

Luong vill., Pha Luong Mountain system, 20°4L30.9"N 104°38'08.9"E, 1425 m a.s.L, 25 Sep

2016, Averyanov, L. et al. CPC 8026 (CPC, E); Van Ho Distr., Tan Xuan comm., A Lay village,

around point 20°40'46.2"N 104°39'49.6"E, approx. 1500 m a.s.L, 1 Oct 2016, Averyanov, L. et

al. CPC 8173 { CPC, PE).

300

Card. Bull. Singapore 69(2) 2017

Notes. The new species is similar to Oreocharis argyreia in its leaf characteristics

(silvery indumentum with densely appressed hairs), but can be distinguished easily by

its yellow corolla which is also of a different size and shape. The differences between

the two species are detailed in Table 1. In Indo-China, Oreocharis argyrophylla is also

quite similar to O. hirsuta from northern Thailand in the yellow and tubular corolla,

but differs in the densely appressed pubescence on the leaves and the cuneate leaf

base (hirsute hairs and obtuse to rounded base in O. hirsuta), more unequal upper and

lower corolla lips (subequal lips in O. hirsuta ), and anthers coherent in pairs (free in

O. hirsuta).

Oreocharis blepharophylla W.H.Chen, H.Q.Nguyen & Y.M.Shui, sp. nov.

This species is similar to Oreocharis magnidens Chun ex K. Y.Pan in leaf characteristics,

but differs from it in the linear bracts and calyx lobes (vs ovate in O. magnidens),

a slightly 2-lipped corolla (vs strongly 2-lipped), a cylindrical corolla tube without

inflated base (vs a corolla tube with inflated base) and coherent anthers (vs free anthers).

In habit characteristics it is also similar to Oreocharis curvituba J.J.Wei & W.B.Xu but

differs from it mainly in having narrowly obovate corolla lobes with purple stripes at

base (vs narrowly oblong without purple stripes in O. curvituba). - TYPE: Vietnam,

Son La province, Van Ho district, Chieng Xuan municipality, Co Hong village, territory

of Xuan Nha nature reserve, Pha Luong Mountain, at an elevation of 1200-1400 m,

20°4T40.5"N 104°39'24.7"E, 13 November 2013, in flower, Averyanov, L., Hiep, N.T.,

Khang, N.S., Thang, N.D. & Qui, L.D. CPC 7019 (holotype KUN; isotypes CPC, LE).

(Fig. 3, 4)

Perennial herb with very short stem. Leaves rosulate, petiolate; petiole 1.1-2.9 cm

long, densely spreading villous; blade narrowly obovate to elliptic, 3.5-6 x 2-3.8 cm,

adaxially with tuberculate setae, abaxially with sparsely strigose hairs along veins,

base cuneate, margin crenulate and ciliate, apex short acuminate to obtuse, lateral veins

5-6 on each side of midrib, indistinct adaxially and distinct abaxially. Inflorescences

axillary, with few flowers; peduncles 5-6 cm long, spreading villous; bracts 2, linear,

2.4-3 x 0.4-0.5 mm, adaxially glabrous and abaxially villous, margin entire; pedicels

8-9 mm, white villous. Calyx of 5 lobes free to base, lobes equal, linear, 6-7 x 1-1.2

mm, glabrous adaxially, villous abaxially, margin entire. Corolla purple with white

limb, 2-2.4 cm long, outside pubescent, inside glabrous; tube cylindrical, purple,

1.1-1.2 cm long, 2.1-2.2 mm in diam., not constricted at throat; limb 2-lipped, white

with purple stripes at the base; adaxial lip 2-lobed, dissected from near base, lobes

broadly obovate, 6-8 x 4-5 mm, apices rounded or slightly retuse; abaxial lip 3-lobed,

lobes subequal, obovate, 9-12 x 5-7 mm, broadening from the base, apices rounded

or slightly retuse. Stamens 4, anthers coherent in 2 pairs; adaxial stamens 1.5-6

mm long, adnate to corolla tube 2.5-3 cm from base, abaxial stamens 2.6-3.1 mm

long, adnate to corolla tube 5-6 mm from base; filaments glabrous, slender; anthers

basifixed, subglobular, 2-locular, dehiscing longitudinally; staminode c. 1.2 mm long,

adnate to corolla tube from the base. Disc ring-like, 1.2-1.4 mm in diam., shallowly

Two new species of Oreocharis in Vietnam

301

Fig. 3. Oreocharis blepharophylla W.H.Chen, H.Q.Nguyen & Y.M.Shui A. Habit (scale bar 1

cm). B. Flower (scale bar 6 mm). C. Villous leaves (scale bar 2 cm). D. Habitat (scale bar 6

cm). (Photos: S.K. Nguyen)

5-lobed. Pistil 4-5 cm long, glabrous; ovary narrowly fusiform, glabrous, 1.5-1.6 mm

long, unilocular; style glabrous, 2.6-2.8 cm long; stigma 1, peltate. Capsule straight,

fusiform, loculicidally dehiscent, 0.8-1 cm long, with persistent style 2.4-2.9 mm

long. Seeds fusiform, 0.5-0.6 x c. 0.2 mm.

Etymology. The specific epithet refers to the long-ciliate leaf margin.

Ecology, distribution and phenology. The new species grows as a lithophytic herb

in primary coniferous and mixed forests with Pinus cernua Aver, et al. on very

steep slopes along a ridge composed of brown sandstone (Averyanov et al., 2014) at

elevations of 1200-1850 m. It is endemic to the Xuan Nha nature reserve in Son La

province of north-western Vietnam. Flowering is October-November and fruiting is

December-January.

302

Card. Bull. Singapore 69(2) 2017

1 cm

Fig. 4. Oreocharis blepharophylla W.H.Chen, H.Q.Nguyen & Y.M.Shui A. Flowering plant.

B. Corolla. C. Cut-open corolla showing two pairs of stamens cohering at the anther tips. D.

Calyx, disc and pistil (immature). E. Capsule dehisced loculicidally on one side. Drawn by

Ling Wang from the holotype Averyanov, L. et al. CPC 7019.

Two new species of Oreocharis in Vietnam

303

Table 2. Characteristics distinguishing Oreocharis blepharophylla, O. magnidens and O.

curvituba.

Characters

O. blepharophylla

O. magnidens

O. curvituba

Petiole

Densely spreading

villous

Densely appressed

silvery pubescent

Sparsely spreading

hirsute

Leaf lateral veins

Adaxially with

tuberculate setae, lateral

veins 5-6 on each side

Adaxially with

tuberculate setae, lateral

veins 8-9 on each side

Adaxially with strongly

bullate setae, lateral

veins 4-7 on each side

Peduncle

5-3 cm, spreading

villous

7-20 cm, densely

glandular pubescent

3-15 cm, brown

pubescent, glandular

puberulent apically

Bracts

Linear, 2.4-3 x 0.4-0.5

mm, and abaxially

villous

Ovate, 5-6 x 1.5-3.5

mm, abaxially brown

sericeous lanate

Lanceolate-linear, c.

3 mm long, glandular

villous outside

Calyx

Lobes linear, 6-7 x

1-1.2 mm

Lobes ovate, 2.5-3 x

1-1.5 mm

Lobes ovate-triangular,

2-2.2 x 0.7-1 mm

Corolla

2-2.4 cm long

0.75-1.2 cm long

1.1-1.3 cm long

Corolla tube

Not constricted at throat,

straight, purple

Narrowed at throat,

inflated at base, straight,

white

Not constricted at throat,

slightly curved, purple

Corolla limb

Limb slightly 2-lipped,

lobes white with purple

stripes

Limb distinctly 2-lipped,

lobes white with purple

tips

Limb distinctly 2-lipped,

lobes white or purplish

without tips

Adaxial lip

Lobes obovate or

oblong, 6-8 x 4-5 mm

Lobes narrowly oblong to

lanceolate, 5-6 x 1.5-2

Upper lips narrowly

oblong, ca. 3.8-4 x

1.2-1.4 mm

Abaxial lip

Lobes obovate or

oblong, 9-12 x 5-7 mm,

apex rounded or obtuse

Lobes narrowly oblong to

lanceolate, 5-9 x 1-1.5

mm, apex acuminate

Lower lips narrowly

oblong, 4.2-4.8 x

1.3-1.5 mm, apex

slightly retuse

Anthers

Coherent in 2 pairs

Not coherent

Provisional IUCN conservation status. Endangered EN Blab(ii,iii,v) + B2ab(ii,iii,v),

following IUCN (2012, 2016) guidelines. This is based on an EOO of < 5000 km 2

and an AOO of < 500 km 2 , being known from fewer than five populations, and with

evidence of a reduction in the area of occupancy, a decline in habitat quality, and a

decline in the number of individuals due to deforestation, largely for agriculture and

due to fire.

Additional specimens examined. VIETNAM. Son La: Moc Chau Distr., Chieng Son comm., Pha

Luong village, PhaLuong Mountain summit, 1750-1850 m a.s.l. 20°40'23.0"N 104°37'52.0"E,

23 Sep 2016, Averyanov, L. et al. CPC 8000 (CPC, LE); Van Ho Distr., Tan Xuan comm., A

Lay village, 20°40'46.2"N 104°39'49.6"E, 1500 m a.s.l., 1 Oct 2016, Averyanov, L. et al. CPC

8171 (CPC, LE).

304

Card. Bull. Singapore 69(2) 2017

Notes. This new species can be distinguished from Oreocharis magnidens by the purple

base of the white corolla lobes (vs white base of purple corolla lobes), corolla tube

throat not constricted (vs constricted throat), and coherent anthers (vs free anthers).

It is also similar to Oreocharis curvituba but differs mainly in the colour pattern and

shape of the corolla (Wei et al., 2016). Table 2 details the differences between these

species.

ACKNOWLEDGMENTS. This work was supported by projects of The National Natural

Science Foundation of China (grant no. 31470306, 31000258); Key Laboratory of Plant

Diversity and Biogeography of East Asia, Kunming Institute of Botany, the Chinese Academy

of Sciences (grant no. 2014CB954100); a US National Geographic Society project entitled

“Exploration of primary woods along constructed highway Hanoi-Ho Chi Minh for their

sustainable conservation in limits of Ha Tinh and Nghe An provinces of central Vietnam” (grant

no. #9129-12); the Mohamed bin Zayed Species Conservation Fund “Conservation assessment

of endangered Lao-Vietnamese stenoendemic-Pmn^ cernua, Pinaceae” (grant 152511753);

and the National Parks Board Singapore. RBGE is funded by the Rural and Environment

Science and Analytical Services division (RESAS) in the Scottish Government.

References

Averyanov, L.V., Nguyen, T.H., Nguyen, K.S, Van, T.P., Lamxay, V., Bounphanmy,

S., Lorphengsy, S., Phan, L.K., Lanorsavanh, S. & Chantthavongsa, K. (2014).

Gymnosperms of Laos. Nordic J. Bot. 32: 768-805.

Cai, L., Chen, R.Z., Yin, Z.J., Zhang, G.X., Chen, W.H. & Shui, Y.M. (2015). Tremacron

hongheense, a new species of Gesneriaceae from Southeastern Yunnan, China. PL

Diversity Resources 37(6): 733-736.

Chen, R.Z., Chen, W.H., Wei, Y.G., Wen, F., Yu, X.L. & Shui, Y.M. (2017). Oreocharis

crispata, a new species of Oreocharis (Gesneriaceae) from Guangxi, China. Phytotaxa

311: 195-199.

Chen, W.H., Shui, Y.M., Yang, J.B., Wang, H., Nishii, K., Wen, F., Zhang, Z.R. & Moller, M.

(2014). Taxonomic status, phylogenetic affinities and genetic diversity of a presumed

extinct genus, Paraisometrum W.T. Wang (Gesneriaceae) from the karst regions of

Southwest China. PLoS ONE 9: el07967.

Chen, W.H., Chen, R.Z. & Moller, M. (2016). Oreocharis ninglangensis, a showy new species

of Gesneriaceae from northwestern Yunnan in China. Phytotaxa 261: 282-286.

Do, T.V., Wei, Y.G. & Wen, F. (2017). Oreocharis caobangensis (Gesneriaceae), a new species

from Cao Bang Province, northern Vietnam. Phytotaxa 302: 65-70.

Ho, PH. (2000). Gesneriaceae. In: Ho, PH. (ed.) An Illustrated Flora of Vietnam , vol. 3, pp.

12-29. Ho Chi Minh: Youth Publishing House.

IUCN (2012). IUCN Red List Categories and Criteria: Version 3.1, 2nd ed. Gland, Switzerland

and Cambridge, UK: IUCN.

IUCN Standards and Petitions Subcommittee (2016). Guidelines for Using the IUCN Red List

Categories and Criteria, ver. 12.

http://www.iucnredlist.org/documents/RedListGuidelines.pdf.

Two new species of Oreocharis in Vietnam

305

Li, Z.Y. & Wang, Y.Z. (2004). Plants of Gesneriaceae in China. Zhengzhou: Henan Science &

Technology Publishing House.

Li, J.M., Wang, T. & Zhang, Y. (2017). Oreocharis zhenpingensis (Gesneriaceae), a new species

from Shaanxi, China. Phytotaxa 307: 292-296.

Moller, M. (2015). Transfer of Tremacron hongheense to Oreocharis (Gesneriaceae). Phytotaxa

239(3): 295-296.

Moller, M., Middleton, D.J., Nishii, K., Wei, Y.G., Sontag, S. & Weber, A. (2011). A new

delineation for Oreocharis incorporating an additional ten genera of Chinese

Gesneriaceae. Phytotaxa 23: 1-36.

Moller, M., Wei, Y.G., Wen, F., Clark, J.L. & Weber, A. (2016). You win some you lose some:

updated generic delineations and classification of Gesneriaceae—implications for the

family in China. Guihaia 36: 44-60.

Pellegrin, F. (1930). Gesneriaceae. In: Lecomte, H. (ed.) Flore generate de L’Indo-Chine, vol.

4, pp. 487-565. Paris: Masson & Cie.

Wang, W.T., Pan, K.Y. & Li, Z.Y. (1990). Gesneriaceae. In: Wang, W.T. (ed.) Flora Reipublicae

Popularis Sinicae , vol. 69, pp. 190-203. Beijing: Science Press.

Wang, W.T., Pan, K.Y., Li, Z.Y., Weitzman, A.L. & Skog, L.E. (1998). Gesneriaceae. In: Wu,

Z.Y. & Raven, P.H. (eds) Flora of China, vol. 18, pp. 268-272. Beijing: Science Press;

and St. Louis: Missouri Botanical Garden Press.

Wei, J.J., Xiong, G.C. & Zou, C.Y. (2016). Oreocharis curvituba, a new species of Gesneriaceae

from northeastern Guangxi, China. Phytotaxa 280: 190-194.

Yang, L.H., Zhou, J.G., Xu, P, Chen, Z.T., Lu, Y.H. & Kang, M. (2015). Oreocharis

pilosopetiolata, a new species of Gesneriaceae from southeastern Guangdong, China.

Phytotaxa 239: 287-292.

Yang, L.H., Huang, J.Z., Deng, F.D. & Kang, M. (2017). Oreocharis imiflora, a new species of

Gesneriaceae from Guangdong, China. Phytotaxa 295: 292-296.

Gardens’ Bulletin Singapore 69(2): 307-313. 2017

doi: 10.26492/gbs69(2).2017-09

307

Primulina wuae (Gesneriaceae), a new species

from southern China

S. Li 1 ’ 2 , Z.B. Xin 1 - 2 , X. Hong 3 , L.F. Fu 12 & F. Wen 1 - 2

'Guangxi Key Laboratory of Plant Conservation and Restoration Ecology

in Karst Terrain, Guangxi Institute of Botany, Guangxi Zhuang

Autonomous Region and the Chinese Academy of Sciences,

CN-541006, Guilin, Guangxi, China

2 Gesneriad Conservation Center of China (GCCC), CN-541006,

Guilin, Guangxi, China

wenfang760608@ 139.com

3 School of Resources and Environmental Engineering,

Anhui University, CN-230601, Hefei, Anhui, China

ABSTRACT. Anew species, Primulina wuae F.Wen & L.F.Fu (Gesneriaceae), is described from

the southern part of China. This new species is most similar to Primulina pseudoroseoalba Jian

Li et al., P. roseoalba (W.T.Wang) Mich.Moller & A.Weber, P. subrhomboidea (W.T.Wang)

Yin Z.Wang and P beiliuensis B.Pan & S.X.Huang var .fimbribracteata F.Wen & B.D.Lai, but

differs from these in characters such as the size and indumentum of the bracts, the indumentum

of the pedicels and anthers, the length of the pistils etc. A provisional conservation assessment

is also provided.

Keywords. Critically endangered, IUCN conservation assessment, limestone flora, new taxon,

Primulina pseudoroseoalba

Introduction

Until recently Primulina Hance was a monotypic genus in the family Gesneriaceae.

Based on molecular phylogenetic and morphological studies, all species of Chiritopsis

W.T.Wang, two species of Wentsaiboea D.Fang & D.H.Qin, and almost all species of

Chirita sect. Gibbosaccus C.B.Clarke have since been incorporated into a dramatically

expanded Primulina (Wang et al., 2011; Weber et al., 2011). Now the genus comprises

more than 176 species, including many recently described species (Moller et al., 2016;

IPNI, 2017).

W hil e conducting field investigations of the limestone flora of northern Guangxi

(China), we found an unknown species of Primulina that required further study. As

well as herbarium specimens, live plants and seeds were collected and grown at the

nursery of Guilin Botanical Garden and the Gesneriad Conservation Center of China.

After comparison to other Primulina species in local Floras and monographs (e.g.

Wang et al., 1990, 1998; Li & Wang, 2004; Wei et al. 2010), comparison to the many

new larger-flowered species described in recent years, especially those from Guangxi

(e.g. Zhu et al., 2014; Huang et al., 2015; Ning et al., 2015; Liu et al., 2016; Luo et al.,

308

Card. Bull. Singapore 69(2) 2017

2016; Wen et al., 2016), and comparison to specimens of Primulina in herbaria (e.g.

IBK, IBSC, KUN, PE), we have confirmed that it is indeed a new species of Primulina

that is here described and illustrated. A provisional IUCN conservation assessment

following the guidelines in IUCN Standards and Petitions Subcommittee (2016) is

also provided.

Taxonomy

Primulina wuae F.Wen & L.F.Fu sp. nov.

This new taxon is morphologically similar to Primulinapseudoroseoalba Jian Fi et al.,

P. roseoalba (W.T.Wang) Mich.Moller & A. Weber, P. subrhomboidea (W.T.Wang) Yin

Z.Wang and P. beiliuensis B.Pan & S.X.Huang var .fimbribracteata F.Wen & B.D.Fai,

but differs from these in the following characters (taxa listed in same order in each

comparison to avoid repetition): bract size (2.1-2.5 x 0.85-1.1 cm in P. wuae vs 1.5-2

x 0.8-1.2 cm in P pseudoroseoalba, vs 1.1-1.3 x 0.25-0.3 cm in P. roseoalba, vs

0.4-0.6 x 0.2-0.3 cm in P. subrhomboidea, and vs 2.8^- x 2.5-3 cm in P. beiliuensis

vwc. fimbribracteata)', bract indumentum (outside sparsely erect pubescent and densely

puberulent, inside puberulent vs outside densely puberulent, inside glabrous, vs outside

strigose, inside glabrous, vs outside puberulent and pilose, inside glabrous, and vs

outside sparsely purple-pubescent, inside glabrous); pedicel indumentum (pubescent

vs puberulent and glandular puberulent, vs puberulent and glandular puberulent, vs

densely pubescent and pilose, and vs densely glandular-pubescent); anther indumentum

(glabrous vs bearded, vs sparsely puberulent, vs villous, and vs densely white lanate),

and pistil length (3-3.5 cm long vs 2.2-2.5 cm long, vs c. 2.8 cm long, vs c. 3.3 cm

long, and vs 8^1 cm long). - TYPE: China, Guangxi Zhuangzu Autonomous Region,

Quanzhou County, Shitang town, alt. 900 m, 18 July 2014, flowering, Wu Xiang-Hong

& Wen Fang WF140718-01 (holotype IBK; isotype IBK). (Fig. 1)

Perennial herb. Stem axis extremely short with indistinct internodes. Leaves 4-8, all

basal, opposite, petioles 1.5-2.5 x 0.35-0.5 cm; leaf blade ovate to ovate-oblong,

left-right slightly asymmetric, but not falcate, 6.5-9.5 x 4.8-8 cm, chartaceous

when dried, adaxially and abaxially appressed eglandular-puberulent, base cuneate

to broadly cuneate, margin entire, repand to undulate, apex acute, obtuse to nearly

rounded; lateral veins 3^4- on each side of midrib, adaxially impressed, abaxially

conspicuously prominent. Inflorescence axillary, a dichasium, 1 or 2 inflorescences on

each plant, (l-)2^4-flowered or more ;peduncle 5.5-1 cm long, eglandular-puberulent

and eglandular-pubescent; bracts 2, free and opposite, narrowly cordate to rhombic,

2.1-2.5 x 0.85-1.1 cm, outside sparsely erect pubescent and densely puberulent,

inside puberulent, margin entire and usually revolute, apex acute; pedicels 1.1-1.6

cm long, pubescent. Calyx 5-sect from base; segments equal, narrowly lanceolate to

lanceolate, 9-10 x c. 3.5 mm, outside puberulent and glandular puberulent, inside

nearly glabrous, margin entire, apex acute. Corolla 35^-0 mm long, dark pink to

purple, the throat pale purple, the upper part of the interior of the corolla tube with

Primulina wuae, a new species from China

309

Fig. 1 . Primulina wuae F.Wen & L.F.Fu. A. Habitat. B. Habit in flower. C. The plant in flower

in cultivation. D. Top view of corolla and calyx. E. Lateral view of corolla, calyx and cyme.

F. Frontal view of corolla with the detail of stigma in inset. G. Opened corolla showing

stamens and staminodes. H. Bracts adaxial surface on the left, abaxial surface on the right. I.

Inflorescence. J. Pistil without calyx. K. Adaxial surface of sepals. A-B, D-K from the type

collection, C from cultivation. (Photos: F. Wen)

310

Card. Bull. Singapore 69(2) 2017

two dark brown flaps, the surface of the flaps glandular-puberulent, outside glandular-

pubescent, inside glandular-puberulent; tube tubular, 24-26 mm long, orifice c. 18

mm in diameter; limb distinctly 2-lipped, adaxial lip 2-partite for more than half of

length, lobes slightly oblique to obviously oblique, triangular to ovate, adaxial lobes

7-8 x 8-8.5 mm; abaxial lip 3-partite to half or slightly more than half of length,

lateral lobes obliquely ovate to oblong, 12-14 x 6-8 mm, the central one oblong,

c. 12 x 6 mm, all five lobes with 4-5 dark purple stripes which sometimes connect

and become somewhat reticulate. Stamens 2, adnate to corolla tube c. 12 mm above

the base; filaments white to pale purple, geniculate near the base, c. 11 mm long,

glabrous; anthers fused by their entire adaxial surfaces, oblong, c. 3 x 1 mm, glabrous

dorsally; staminodes 2, translucent to white, linear, apex capitate, glabrous, c. 7 mm

long, adnate to corolla 10-12 mm above base. Disc purplish brown, annular, glabrous,

1.3-1.5 mm high. Pistil 30-35 mm long; ovary cylindrical, 16-18 mm long, 1.8-2 mm

in diameter, densely puberulent and glandular puberulent; style 15-17 mm long, c. 1

mm in diameter, densely glandular puberulent. Stigma translucent to white and pale

purple, cuneate, chiritoid, 2-lobed, 2.2-2.5 mm long. Capsule linear, slightly upward-

curved, 5-6 cm long, surface densely puberulent when young.

Distribution. Only known from the type locality, near Shijiaopen Village, Shitang

Town, Quanzhou County, Guangxi Zhuangzu Autonomous Region, China.

Habitat. Primary forest on shaded slopes and vertical cliffs in limestone valleys and

hills, rare, at low altitude, c. 177 m. It occurs in crevices in limestone rocks under

evergreen forest and is rather prone to human disturbance.

Etymology. The new species is named after Ms Xiang-Hong Wu, who first discovered

and collected this rare species and who accompanied us on a number of subsequent

field expeditions in Quanzhou, Guangxi.

Provisional IUCN Conservation Assessment. Critically Endangered CR B2ab(iii,v)

according to the IUCN red list criteria (IUCN Standards and Petitions Subcommittee

2016). We have visited the type locality of this new species many times and have

found not more than 400 individuals which are scattered in a fairly narrow area at the

top of a cliff. They grow in a popular scenic spot near the village of Shitang town, but

most of the plants are protected from tourists by their elevated location.

Vernacular name. The Chinese pronunciation of Primulina wuae is

‘Wu Shi Bao Chun Jfi Tai\

Notes. Primulina wuae has a pair of relatively large bracts. Large bracts also occur in

Primulina beiliuensis B.Pan & S.X.Huang (Pan et al., 2013), P. eburnea (Hance) Yin

Z.Wang (Hance, 1883; Wang et al., 2011), P. lutea (Yan Liu & Y.G.Wei) Mich.Moller

& A.Weber (Liu & Wei, 2004; Weber et al., 2011), P. lunglinensis (W.T.Wang) Mich.

Moller & A.Weber (Wang, 1981; Weber et al., 2011), P. lungzhouensis (W.T.Wang)

Table 1. Primulina wuae F.Wen & L.F.Fu compared to closely related species.

Primulina wuae, a new species from China

311

> s

•2 §

« &

•2 £

P 2 *'

a; ^

-a

•s

-a

®N*

0 )

—

2 S

§ &

X 3

.3 Oh

* X

X 05

3 05

J^ »

"3 .3

3 *

O 3

* 3

P 3

o oJ

o> P

Oh ,5

3 3

CD 3

O ^

X m

r- P

<U g

S 3

!-h

3 &

2 X

>• .3

2 %

c/5 3

_r 3

Oh „

05 X

j *,2

I | §

3 I £

X3 W) 3

3 CD Oh

>> 5

~~! 3

> -.,

o *

r, 3 - C

<D X 3

[3 3

CS t 3

Oh 3

X 3

bfl

X *

bi)

3 D

s I

x &

r-~

£? §

73 ■*-»

w £

2 «

0 ) 3

2 -2

O o

ip ?o

X >.

Jg -S

2 ) 2

aj 3

3? ^

1 «

X 3

7j | .

•oi 3

CD 05

73 -+j

S -c

O bO

Jh • -H

J-H

"S c3 fe

^ 3 I

2 Oh bfl

Jo x

p D

-4-> r—;

o 1

S"S

. 2 " 6 b

?! ..

Jo q cd

X V X

X O

bD c3

X >

00 O

5-h ^

2? 2

Oh <D

cd X5

g |

>< 3

c 3

'S *3->

C/5 00

a O

a-) .73

X Oh

g> c

-4—»

C3 ^

3 o

o X

°f S

l C o

m Oh

c 3

O ^

^ c

Ch p

u s

m a>

7 3

a> Oh

r§

c 3

F 5b

b£

r^H

CO Oh

3 3

x x

c c

c3 cd

7b 'tb

<D (D

g> C

§ ^

3 <p

<D O

X X

3 3

in Oh X

^ O

*05

rd*

5-h

•*-*

bi)

rs .

5-h

O <N

V X

<L>

Hi—*

c 3

_d ^

D 0 )

„ <D

% 3

i—c 05

S) 3

3 °

3 _r

•a

o o

jo x

^ X

P o

2 J.

3 X

£ o

‘Qh

c 3

-*—»

CD 05

5 g

Oh bX)

a->

c 3

31)

' 05

r5 ^

£ c5

75 v

OJ X

.9 m

a->

■4—»

c 3

'■e

■4—»

c 3

^ 05

3 2

a 3

a> r5

73 bo

^ •

(D ^

>, <D

r—P 05

<D C3

05 OJ

^3 ^

Oh 73

CD cb

■S3

is S

•S

5-h

<D <D

X X

O =5

Oh Oh

a M

° o

(N X

I o

in cb

-H* 3o

3 <D

<D C/5

X O

bh 'Bh

l-s

s -

7|-

r^H

O r-.

Oh ^

S-H

g 1 !

O ^

in x

7 M

-H 3)

CD 9

x S

Oh =

^ ^H

cb ^

3 3

3 a

£ «

o 3

00 X

CO J

CD 3)

. CD

>> X>

— o

bi)

■3 od

O 3

^ Vi

£ •£

^ s"

m S

7 &

oi ^

in *2 g

oo 3 g

^ 3 3

r- Oh Qh

3 o

5-h

7 *

_H CZ1

^3 3

oo -§

O Oh X Oh 3

2 b

bfl

3 (D

bfl X

X vi

3 «

3 __

•4—» X

o o

JD ^CD

3 3

5-h 5-h

CD CD

XXX

o o

Oh Oh

7l-

3 b

05 Cg

3 o

° ^

. . CD

bi) 73

73 *05

C £

o o o

CD CD CD

2 2 2

—I CD (D <D

I X X X

0 3 3 3

—< Oh Oh Oh

3 3

x 3

o 73

o 3

3 ^

3 00

£ x

1 P §

cj s

^ 2

O CD

7 3

ON Oh

*05

CD 05

2 §

g |

o 3

Oh bi)

O ^

Oh C

OD QD

.P 05

05 CD

3 -§

O Oh

D ”

2 3

22 'P

.£ 3

.. x

3 «

D J

3 O

D «

Oh 3

D 2

3 2

05 CD

3 g

O Oh

3 .s

D x

X 3

2 ”2

D 3

3 C

Oh °

3 2

1 2

3 gj

Oh X

“ 3

D Oh

2 D

Vi "D

3 ' Vi

O .£

3 .S

D x

X 3

2 x

D 3

3 C

>. 3

3 2

1 2

3 QJ

Oh X

“ 3

D Oh

2 D

vi X

■hh .3

3 Vi

O 2

fl -3

D 3

D D

D 3

3 «

2 ?

7 3

a 73

i3 o

bi) c3

OD bO

*05 73

3 *05

o .a

J 3

X ^

^ o

T3 P

c/)

5-h CD

73 DD

Oh O

CO Oh

^CD

5-h

‘Oh

ChC ^

o a

Oh D

3 O

D X

D 3

cz) CD

x is

3 2

Oh 3

2 ? 3

D 3

(ZJ

dh bo

3 -

vi aj

D 2

5-h

fd*

5-h

S 3

s/)

•4-J

H—»

5-h

H-H

, , -4-^

O a

73 '

3 3

^ £

S §

o S

Vh O

2 d £

fd*

2 §

o Xi

i J

i—i X

X 3

3 X

u .£

E 2

< 2

55 .

bi)

'So o

O ^

CD X

Q Oh

bi)

312

Card. Bull. Singapore 69(2) 2017

Mich.Moller & A.Weber (Wang, 1981; Weber et al., 2011), P. maguanensis (Z.Yu

Li et al.) Mich.Moller & A.Weber (Xu et al., 2008; Weber et al., 2011), and others.

Although Primulina wuae shares this very striking large bract character with these

species, overall it more closely resembles P. beiliuensis var. fimbribracteata F.Wen

& B.D.Lai (Lai & Wen, 2015). The corolla of many Primulina species is generally

rather featureless except for a few distinct features such as yellow ridges ventrally

in the corolla tube (also present in other Gesneriaceae genera) and coloured ‘flaps’

dorsally in the tube that hold the style in place (Weber, 2004), as in P. wuae. The new

species also looks rather similar to Primulina pseudoroseoalba (Li et al., 2014) which

has the same leaf shape but differs in corolla characters. Primulina wuae lacks corolla

ridges but has ‘flaps’, whereas both ridges and flaps are lacking in P. pseudoroseoalba.

However, flaps and a similar corolla colouration are present in other species such

as Primulina subrhomboidea and P. roseoalba. Primulina wuae is compared to the

morphologically similar taxa P. pseudoroseoalba, P. roseoalba, P. subrhomboidea and

P. beiliuensis var. fimbribracteata in Table 1.

ACKNOWLEDGEMENTS. This study was financially supported by the Foundation of

Key Laboratory of Plant Resources Conservation and Sustainable Utilization, South China

Botanical Garden, Chinese Academy of Sciences; the Foundation of Guangxi Key Laboratory

of Plant Conservation and Restoration Ecology in Karst Terrain (16-A-01-01, 16-A-01-02,

16-B-01-01); the Natural Science Foundation of Guangxi (2015GXNSFBB139004); Guangxi

science and technology project (Guike AB16380053); the Open Project of Beijing Engineering

Research Center of Rural Landscape Planning and Design (kf2017061); the STS Program of

the Chinese Academy of Sciences (Grant No. KFJ-3W-Nol). We also thank the anonymous

reviewers’ valuable suggestions for improvements to this paper.

References

Hance, H.F. (1883). New Chinese Cyrtandreae. J. Bot. 21: 165-170.

Huang, S.L., Fu, L.F., Li, J.J., Ge, Y.Z., Ma, W., Wen. F. & Zhou, S.B. (2015). Primulina

duanensis sp. nov. (Gesneriaceae) from Guangxi, China. Nord. J. Bot. 33: 209-213.

IPNI (2017). The International Plant Names Index, http://www.ipni.org. Accessed 1st June

2017.

IUCN Standards and Petitions Subcommittee (2016). Guidelines for Using the IUCN Red List

Categories and Criteria, ver. 12.

http://www.iucnredlist.org/documents/RedListGuidelines.pdf.

Lai, B.D. & Wen, F. (2015). Primulina beiliuensis war. fimbribracteata (Gesneriaceae), a new

variety in a limestone cave from northern Guangdong, China. Taiwania 60(1): 43—48.

Li, Z.Y. & Wang, Y.Z. (2004). Boeica. In: Li, Z.Y. & Wang, Y.Z. Plants of Gesneriaceae in

China, pp. 109-112. Zhengzhou: Henan Science and Technology Publishing House.

Li, J., Hong, X., Wen, F. & Yan, L.J. (2014). Primulina pseudoroseoalba (Gesneriaceae), a new

species from a karst cave in Guangxi, China. Ann. Bot. Fenn. 51: 86-89.

Liu, Y. & Wei, Y.G. (2004). Chirita lutea Yan Liu & Y.G. Wei, a new species of Gesneriaceae

from Guangxi, China. J. Wuhan Bot. Res. 22(5): 391-393.

Primulina wuae, a new species from China

313

Liu, Y., Su, L.Y., Zheng, K.W., Huang, S.L., Tian, F. & Wen, F. (2016 ). Primulina melanofilamenta

sp. nov. (Gesneriaceae) from Guangxi, China. Nord. J. Bot. 34: 38-42.

Luo, W.H., Pan, B. & Wen, F. (2016). Primulina gigantea (Gesneriaceae), a new species from

Guangxi, China. Ann. Bot. Fenn. 53: 426-430.

Moller, M., Wei, Y.G., Wen, F., Clark, J.L. & Weber, A. (2016). You win some, you lose some:

updated generic delineations and classification of Gesneriaceae - implications for the

family in China. Guihaia 36: 44-60.

Ning, Z.L., Li, D.M., Pan, B. & Kang, M. (2015). Primulina rubribracteata , a new species of

Gesneriaceae from southern Hunan, China. Phytotaxa 239(1): 55-64.

Pan, B., Wen, F., Zhao, B., Deng, T., Xu, W.B. & Huang, S.X. (2013). Primulina beiliuensis

B. Pan & S.X. Huang, a new species of Gesneriaceae from limestone areas in Guangxi,

China. Guihaia 33(5): 591-598.

Wang, W.T. (1981) Notulae De Gesneriaceis Sinensibus (II). Bull. Bot. Res., Harbin 1: 35-75.

Wang, W.T., Pan, K.Y., Zhang, Z.Y. & Li, Z.Y. (1990). Gesneriaceae. In: Wang, W.T. (ed.)

Flora Reipublicae Popularis Sinicae, vol. 69. Beijing: Science Press.

Wang, W.T., Pan, K.Y., Li, Z.Y, Weitzman, A.L. & Skog, L.E. (1998). Gesneriaceae. In: Wu,

Z.Y. & Raven, PH. (eds) Flora of China , vol. 18. Beijing, China: Science Press and St.

Louis, Missouri, USA: Missouri Botanical Garden Press.

Wang, Y.Z., Mao, R.B., Liu, Y., Li, J.M., Dong, Y., Li, Z.Y. & Smith, J.F. (2011). Phylogenetic

reconstruction of Chirita and allies (Gesneriaceae) with taxonomic treatments. J. Syst.

Evol. 49(1): 50-64.

Weber, A. (2004). Gesneriaceae. In: Kadereit, J.W. (ed.) Flowering Plants. Dicotyledons , vol.

7. Berlin, Germany: Springer.

Weber, A., Middleton, D.J., Forrest, A., Kiew, R., Lim, C.L., Rafidah, A.R., Sontag, S., Triboun,

P, Wei, Y.G., Yao, T.L. & Moller, M. (2011). Molecular systematics and remodelling of

Chirita and associated genera (Gesneriaceae). Taxon 60: 767-790.

Wei, Y.G., Wen, F., Moller, M., Monro, A., Zhang, Q., Gao, Q., Mou, H.F., Zhong, S.H. &

Cui, C. (2010). Gesneriaceae of South China. Nanning, China: Guangxi Science and

Technology Publishing House.

Wen, F., Lai, B.D., Zhao, Z.G., Wang, B.M & Wei, Y.G. (2016). Primulina linearicalyx

(Gesneriaceae), a new species from Guangxi, China. Phytotaxa 269(1): 41-46.

Xu, H., Li, Z.Y. & Jiang, H. (2008). A new species of Chirita (Gesneriaceae) from Yunnan,

China. Bot. J. Linn. Soc. 158: 269-273.

Zhu, X.X., Wen, F. & Sun, H. (2014). Primulina glandaceistriata (Gesneriaceae), a new species

from Guangxi, China. Phytotaxa 188(1): 49-54.

The Gardens’ Bulletin

Singapore

VOL. 69 2017

ISSN 0374-7859

VOLUME 69

No. 1 pages 1-156 (printed version published 24 May 2017 )

No. 2 pages 157-313 (printed version published 20 November 2017)

THE GARDENS’ BULLETIN SINGAPORE

The Gardens’ Bulletin Singapore is a peer-reviewed journal publishing original papers and

reviews on a wide range of subjects: plant taxonomy (including revisions), phytogeography,

floristics, morphology, anatomy, as well as horticulture and related fields, such as ecology and

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

Dr David J. Middleton Dr Jana Leong-Skornickova

(Editor-in-Chief) (Managing Editor)

Felicia Tay Dr Daniel C. Christina Soh

(Graphics Editor) Thomas (Business Manager)

(Copy Editor)

Editorial Advisory Board

Professor Sir Peter Crane

Yale University

U.S.A.

Dr Rogier P.J. de Kok

Royal Botanic Gardens, Kew

U.K.

Dr W. John Kress

National Museum of Natural History

Smithsonian Institution

U.S.A.

Dr Mark Hughes

Royal Botanic Garden Edinburgh

U.K.

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

Published twice yearly by the National Parks Board, Singapore, the annual subscription for the journal

is Singapore $100.00 including postage. Overseas subscribers should make payment in the form of

bank draft or international money order in Singapore currency, payable to National Parks Board. Please

forward payment to “Accounts Receivable Section, National Parks Board, Headquarters, Singapore

Botanic Gardens, 1 Cluny Road, Singapore 259569”.

National Parks Board

Singapore Botanic Gardens

1 Cluny Road

Singapore 259569

Printed by Oxford Graphic Printers Pte Ltd

CONTENTS OF VOLUME 69 (2017)

VOL. 69(1), May 2017

A.R. Rafidah

Taxonomy and conservation status of Microchirita

(Gesneriaceae) in Peninsular Malaysia.1

M. Rodda

Index of names and types of Hoya

(Apocynaceae: Asclepiadoideae) of Borneo.33

Y.W. Low, A.K. Muhammad Ariffin, A.A. Joffre & D. Duratul Ain

Novitates Bruneienses, 8. Macrosolen brunsing (Loranthaceae),

a new hemiparasitic shrub from Brunei Darussalam.67

I Putu Gede P. Damayanto & E.A. Widjaja

A noteworthy Dendrocalamus (Poaceae: Bambusoideae)

from Sumatra, Indonesia.75

I. Haerida

Liverworts of Bali, Indonesia, with new records to the island.81

C.-W. Lin, D.C. Thomas, W.H. Ardi & C.-I Peng

Begonia ignita (sect. Petermannia, Begoniaceae), a new species

with orange flowers from Sulawesi, Indonesia.89

N. Simonsson Juhonewe & M. Rodda

Contribution to a revision of Hoya (Apocynaceae: Asclepiadoideae)

of Papuasia. Part I: ten new species, one new subspecies

and one new combination.97

S. Lindsay, P.H. Hovenkamp & D. J. Middleton

New combinations and typifications in Aglaomorpha (Polypodiaceae).149

VOL. 69 (2), November 2017

M.A. Niissalo & J. Leong-Skornickova

Hanguana podzolicola (Hanguanaceae),

a new record for Singapore.157

R.P. J. de Kok

Two new records of Lit sea (Lauraceae) from Singapore

and the lectotypification of twenty-two names

from several Lauraceae genera.167