The Gardens’ Bulletin Singapore Vol. 68(1) 2016

The Gardens’ Bulletin

Singapore

ISSN 0374-7859

Singapore Botanic Gardens

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 life of the Southeast Asian-Pacific region.

V

Dr David J. Middleton Dr Jana Leong-Skomickova

(Editor-in-Chief) (Managing Editor)

Felicia Tay Yee Wen Low Chiistina Soh

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

Editorial Advisory Board

Professor Sir Peter Crane

Yale University

US. A.

Dr Rogier P.J. de Kok

Honoraty Research Associate

Singapore Botanic Gardens

Dr W. John Kress

National Museum of Natural History

Smithsonian Institution

U.S.A.

Dr Mark Hughes

Royal Botanic Garden Edinburgh

UK.

Dr Kiat W. Tan

Gardens By The Bay

Singapore

Dr Nigel P. Taylor

Singapore Botanic Gardens

National Parks Board, Singapore

Dr Ian M. Turner

Royal Botanic Gardens Kew

UK.

Dr Jan-Frits Veldkamp

National Biodiversity Center

The Netherlands

Dr Jun Wen

National Museum of Natural Histoty

Smithsonian Institution

U.S.A.

Professor Nianhe Xia

South China Institute of Botany

PR. China

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

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

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

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

rps 16 intron and //7?L(UAA)-F(GAA) intergenic spacer. Nordic J. Bot. 20: 257-269.

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

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

of Fiji 143-193.

References to web-based resources should include either a doi (digital object identifier) specification

or full URL mentioning also the date it was accessed. Use of DNA sequences from GenBank should be

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

Style of nomenclatural summaries. The following style is required:

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

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

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

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

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Front cover picture: Aeschynanthus speciosus (Photo by J. Leong-Skomickova)

The Gardens’ Bulletin

Singapore

VOL. 68(1) 2016 ISSN 0374-7859

CONTENTS

D.J. Middleton

A revision of Aeschynanthus (Gesneriaceae)

in Singapore and Peninsular Malaysia 1

I.M. Turner

Notes on the Annonaceae of the Malay Peninsula 65

K.M. Wong, A.K. Muhammad Ariffin & A.A. Joffre

Novitates Bruneienses, 5. Polyalthia watui (Annonaceae),

a new tree species from Brunei, Borneo 71

D. Girmansyah

Three new species of Begonia (Begoniaceae)

from Sumbawa Island, Indonesia 77

D. Sulistiarini, D. Arifiani & Y. Santika

New records of Orchidaceae from Bali, Indonesia 87

R. Kiew

Olea luzonica (Oleaceae), a new name

for O. obovata from the Philippines 97

T. Phutthai & M. Hughes

A new species and a new record in Begonia sect.

Platycentrum (Begoniaceae) from Thailand

99

J.D. Mood, A.G. Hussain & J.F. Veldkamp

The resurrection of Boesenbergia albosanguinea (Zingiberaceae)

with a new record for Peninsular Thailand

109

J.D. Mood, H.D. Tran, J.F. Veldkamp & L.M. Prince

Boesenbergia siphonantha (Zingiberaceae),

a new record for Thailand and Vietnam

with notes on the molecular phylogeny 125

F. Wen, T.V. Do, X. Hong, S. Maciejewski & Y.G. Wei

Boeica ornithocephalantha (Gesneriaceae),

a new species from northern Vietnam 139

M. Moller, K. Nishii, H. J. Atkins, H.H. Kong, M. Kang,

Y.G. Wei, F. Wen, X. Hong & D.J. Middleton

An expansion of the genus Deinostigma (Gesneriaceae) 145

J.F. Veldkamp

The valid publication of Monolophus (Zingiberaceae) revisited 173

Date of publication: 20 May 2016

National Parks Board

Singapore Botanic Gardens

1 Cluny Road

Singapore 259569

Printed by Oxford Graphic Printers Pte Ltd

Gardens’ Bulletin Singapore 68(1): 1-63. 2016

doi: 10.3850/S2382581216000016

1

A revision of Aeschynanthus (Gesneriaceae)

in Singapore and Peninsular Malaysia

DJ. Middleton

Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, Singapore 259569

david_middleton@nparks ,gov. sg

ABSTRACT. The genus Aeschynanthus Jack is revised for Singapore and Peninsular Malaysia.

Four species for Singapore and fourteen species for Peninsular Malaysia are recognised, keys

to the species are given, all names are typified, and detailed descriptions of all species are

provided. Conservation assessments are provided for all species. Eleven names are lectotypified

here and one epitype is designated.

Keywords. Conservation assessments, Didymocarpoideae, identification key, lectotypifications

Introduction

Aeschynanthus Jack is a large and variable genus with around 160 species from Sri

Lanka and India through southern China and Southeast Asia to New Guinea and

the Solomon Islands (Weber, 2004; Middleton, 2007). The last complete account of

the genus was by Clarke (1883) who included 64 species. More recently, regional

revisions have been published for China (Wang et ah, 1998), Thailand (Middleton,

2007), Cambodia, Laos and Vietnam (Middleton, 2009), and India (Bhattacharyya &

Goel, 2015). Checklists have been published for Singapore (Turner, 1993; Chong et

ah, 2009), Peninsular Malaysia (Turner, 1 997), Myamnar (Kress et al., 2003), Sulawesi

(Mendum & Atkins, 2003), and Sumatra (Tjitrosoedirdjo et al., 2009). Several of these

checklists are now rather out-of-date due to the discovery of new species and/or due to

the synonymisation of names.

A general background to research on Aeschynanthus was given in Middleton

(2007). For Singapore and Peninsular Malaysia the last comprehensive treatment

was by Ridley (1923), in which 14 species of Aeschynanthus for the Malay Peninsula

were recognised. The checklists for Singapore (Turner, 1993; Chong et ah, 2009) both

included four native species; the checklist for Peninsular Malaysia (Turner, 1997)

included 17 species (one with two varieties).

In preparation for a revision of Gesneriaceae for the Flora of Peninsular

Malaysia the genus has been revised for Peninsular Malaysia and for neighbouring

Singapore. Fourteen species are recognised. All fourteen species are in Peninsular

Malaysia and four of these are also in Singapore. One of the species in Singapore

is presumed extinct there and the other three are considered Critically Endangered

(Chong et al., 2009; Williams, 2014).

2

Gard. Bull. Singapore 68(1) 2016

An infrageneric classification is not followed in this paper pending a more

thorough investigation of the genus using molecular sequence data to compare the

morphological characters. A preliminary study by Denduangboripant et al. (2001)

found that a phylogeny based on ITS data did not reveal monophyletic clades that

corresponded to the existing sections.

All provisional lUCN Conservation Assessments given here, calculated using

the methodology of lUCN (2012), are for the species throughout their range rather

than only for Peninsular Malaysia and/or Singapore. Additional comments are given

under each species when the local situation differs from the global.

Morphological Characters

A more detailed discussion of morphological characters is given in Middleton (2007)

and references given therein. Here is a brief discussion of the range of characters found

in the species in Singapore and Peninsular Malaysia.

All species are epiphytes or occasionally lithophytes. Species such as

Aeschynanthus speciosus Hook., A fulgens Wall, ex R.Br. and A. rhododendron Ridl.

are large with robust stems that arch or hang due to their own weight. Others such as

Aeschynanthus aJbidus (Blume) Steud., A. angustifolius (Blume) Steud., A. fecundus

P. Woods, A. longicaulis Wall, ex R.Br., A. longiflorus (Blume) A.DC., A. obconicus

C.B. Clarke in A.DC. & C.DC. and A. wallichii R.Br. are generally more delicate

and are more pendulous, although usually not loosely hanging (as, for example, is

found in A. gracilis Parish ex C.B.Clarke from Thailand to NE India). Aeschynanthus

dischidiodes (Ridl.) D.J.Middleton has often been reported as growing from ants’ nests

in trees. Aeschynanthus pulcher (Blume) G.Don, A. radicans Jack and probably A.

volubilis Jack either have the habit of the more delicate species previously mentioned,

sometimes with long pendent stems, or creep over tree trunks and branches or rocks,

rooting at the nodes. The erect habit of species such as Aeschynanthus andersonii

C.B.Clarke and A. humilis Hemsl. from Thailand is not found in Singapore or

Peninsular Malaysia.

The leaves of most species are opposite but are always in whorls of three or

more m Aeschynanthus speciosus and A. angustifolius. In Aeschynanthus angustifolius

the leaf blades are extraordinarily variable in shape (see discussion under that species).

In the other species the leaves are generally ovate to elliptic and not particularly

variable in shape and size. In most species the margins are entire but they are distinctly

toothed in Aeschynanthus dischidioides, sometimes slightly so in A. angustifolius, and

strongly undulate in A. speciosus. All species are shortly petiolate and the blades are

mostly coriaceous, more rarely thinner and softer.

The structure of the inflorescence is discussed in Middleton (2007) and references

therein. In Singapore and Peninsular Malaysia all species either lack a peduncle or the

peduncle is so short as to appear absent. The flowers are, therefore, either axillary and

solitary, axillary and appearing fasciculate, or subterminal and clustered. In all species

in Singapore and Peninsular Malaysia the bracts are fairly small, simple and linear.

The calyx consists of five lobes free to the base {Aeschynanthus angustifolius.

Aeschynanthus in Singapore and Peninsular Malaysia

3

A. disch idiocies, A.fecundus, A. longicaulis, A. longiflorus and A. speciosus), or a very

short tube with five long lobes (A. albidus), or of a iiaiTow or somewhat widening tube

that is around half or more of the length of the calyx with either obvious or obscure

lobes at margin {A.fulgens, A. pulcher, A. radicans,A. rhododendron and^. volubilis),

or of an open cup-shaped calyx with obscure lobes at the margin (A. obconicus and^.

wallichii). The colour of the calyx can either be quite fixed within a species (it always

appears to be red in A. obconicus for example) or very variable within a species (dark

purple or red to green in A. pulcher for example). The calyx shape and size is very

variable in a number of species.

The corolla is zygomoiphic, tubular, and weakly to quite strongly curved. The

limb is 2-lipped with the upper lip 2-lobed and the lower lip 3-lobed. Visitation by

animals has not been observed in most species but pollination is assumed to be by

birds for most, possibly all, species (see Middleton (2007) for further discussion). The

corolla is red, orange, yellow or green, or a combination of these colours, in the species

in Singapore and Peninsular Malaysia. Flower colour is highly diagnostic for most

species although is often very subjectively recorded on herbarium specimen labels

when it is described at all.

All Aeschynanthus species are strongly protandrous with the stamens withering

as the style elongates and the stigma enlarges. There are four stamens in two pairs with

the anthers of each pair fused at their tips. In Singapore and Peninsular Malaysia no

species have all four anthers fiised together as is found in Aeschynanthus chiritoides

C.B.Clarke from Vietnam to NE India. The gynoecium consists of the stipe, the

ovary, the style and the stigma. The stipe is short in all species but rather longer in

Aeschynanthus rhododendron. The dimensions given in the descriptions below reflect

the measurements made on specimens to hand. It should be borne in mind though that

the absolute and relative lengths of the parts of the gynoecium are enormously variable

depending on the age of the flower.

The fruit is a long and narrow capsule in all species. Dehiscence is loculicidal.

The basal portion of the capsule, the stipe of the unfertilised gynoecium, lacks seeds

and is generally short except in Aeschynanthus rhododendron where it fonns an

obvious naiTow stalk.

The seeds have been the principal source of characters for earlier infrageneric

classifications of Aeschynanthus (see Middleton (2007) and references cited therein).

The seeds of Aeschynanthus species consist of the seed grain, one apical appendage

and one or more hilar appendages. Seeds with two hilar appendages are not found

anywhere in Malesia (Mendum et al., 2001). The apical appendage points towards the

base of the capsule. In Singapore and Peninsular Malaysia there is one hilar appendage

in Aeschynanthus angustifolius, A. fulgens, A. longiflorus, A. obconicus, A. pulcher,

A. radicans, A. rhododendron, A. speciosus, A. volubilis and A. wallichii and three or

more hilar appendages in A. albidus, A. dischidioides, A. fecundus and^^. longicaulis.

Of those with only one hilar appendage, the appendage is long and filifoim in all

species except Aeschynanthus rhododendron where it is short and stout. In a number

of species there is a curious cluster of inflated cells, termed bubble cells, at the hilar

end of the seed. These are found in A. obconicus, A. pulcher, A. radicans, A. volubilis

and A. wallichii.

4

Card. Bull. Singapore 68(1) 2016

Materials studied

Herbarium material was studied from the following herbaria: A, AAU, BISH, BKF,

BM, C, CGE, E, FI, G, G-DC, K, KEP, KLU, K-W, L, LAE, M, MEL, MICH, NY, P,

PSU, SING, SINU, TI, U, UKMB, US (herbarium codes from Thiers (continuously

updated)). All specimens cited have been seen unless otherwise indicated with nv. A

single standardised name is given for those collectors whose name appears in more

than one fonu, including when abbreviated to initials, on different collections.

The dimensions given in the descriptions are for dried material for vegetative

characters and rehydrated or fresh material for floral characters. Dimensions given

closely resemble those given in Middleton (2007) for some of the taxa due to the

paucity of material collected since and because Malaysian material was used in the

Thai descriptions when the Thai material was insufficient (as it was for several taxa).

The vegetation types given in the Habitat and Ecology sections below follow

notes on the specimens and observations in the field using the vegetation classification

of Saw (2010).

Aeschynanthus Jack

Trans. Linn. Soc. London 14: 42 (1823); C.B.Clarke inA.DC. & C.DC., Monogr. Phan.

5(1): 18 (1883); Ridley, FI. Malay Penins. 2: 496 (1923); Wang, FI. Reipubl. Popularis

Sin. 69: 498 (1990); Middleton, Edinburgh J. Botany 64: 368 (2007); Middleton,

Edinburgh J. Botany 66: 393 (2009). - TYPE: Aeschynanthus volubilis Jack.

Trichosporum D.Don, Edinburgh Philos. J. 7: 82 (1822), nom. rej; Blume, Bijdr.

FI. Ned. Ind. (1826). - TYPE: Trichosporum paniflorum D.Don {= Aeschynanthus

parviflorus (D.Don) Spreng.), lectotype designated by Middleton (2007).

Rheitrophyllum Hassk., Flora 25 (2): beibl. 56 (1842). - TYPE: Rheitrophyllum

subverticillatum Hassk. (= Aeschynanthus angustifolius (Blume) Steud.).

Oxychlamys Schltr., Bot. Jahrb. Syst. 58: 286 (1923). - TYPE: Oxychlamys pullei

Schltr. {= Aeschynanthus oxychlamys Mendum)

Euthamnus Schltr., Bot. Jahrb. Syst. 58: 284 (1923). - TYPE: Euthamnus papuanus

Schltr. {= Aeschynanthus papuanus (Schltr.) B.L.Burtt)

Micraeschynanthus Ridl., FI. Malay Penin. 5 : 324 ( 1 925). - TYPE: Micr aeschynanthus

dischidioides Ridl. (= Aeschynanthus dischidioides (Ridl.) D. J.Middleton)

Epiphytic herbs or subshrubs with erect, arching or pendulous stems, these sometimes

rooting along their lengths when in contact with a suitable substrate. Leaves opposite or

verticillate, pedicellate; blades coriaceous to distinctly fleshy, more rarely herbaceous.

Aeschynanthus in Singapore and Peninsular Malaysia

5

simple, margins entire to weakly crenate or weakly dentate, sometimes somewhat

undulate, venation pinnate but more often than not obscure. Inflorescence an axillary

few-flowered cyme, or flowers solitaiy in the axils of leaves, or a pseudoterminal

cluster. Flowers strongly protandrous. Calyx of 5 sepals, these free or variously fused

into a tube for part or most of length, when fused the whole tubular or cup-shaped.

Corolla zygomorphic, tubular, widening towards lobes, curved to various degrees,

sometimes distinctly inflated at the base, glabrous to variously pubescent outside and

inside; with 5 lobes, these consisting of a 2-lobed upper lip, 2 lateral lobes and a lower

lobe; very variable in colour but most frequently red, orange, yellow or green (or

combination of these) and then often with other darker or lighter patterning. Stamens

4, in 2 pairs, attached to the inside of the corolla tube and occupying the space in

the upper curve of the flowers, included or exserted from corolla tube when mature;

vestigial staminode present; anthers of each pair attached by their apices (occasionally

all 4 attached together outside this region). Disk present, annular to dentate. Pistil

developing as filaments wither and reflex downwards and also occupying the space in

the upper curve of the corolla tube, consisting of a sterile stipe at the base, the fertile

ovary section, the style and the peltate stigma; ovules many, anatropous. Fruit a long

naiTow capsule which opens loculicidally by two valves. Seeds many, tiny, with short

to long appendages at both ends.

About 160 species from India and southern China through Southeast Asia and Malesia

to the Solomon Islands. Fourteen species in Peninsular Malaysia, four species in

Singapore (of which one is considered to be nationally extinct).

Key to Aeschynanthus species recorded from Singapore

1 a. Corolla predominantly green or yellowish, inside with coarse multicellular hairs;

seeds with many hairs at one end \.A. albidus

lb. Corolla red, inside without coarse multicellular hairs; seeds with only 1 hair at

each end 2

2a. Calyx < 7 mm long, in a wide and shallow cup or saucer, much wider at apex than

at base 14. wallichii

2b. Calyx >10 mm long, tube mostly parallel to corolla tube, apex not much wider

than base 3

3 a. Ovary, stipe and style densely pubescent; leaves pubescent beneath

10. A. radicans

3b. Ovary with sessile glands, only stipe and style pubescent; leaves usually glabrous,

more rarely sparsely pubescent beneath 9. A. pulcher

6

Gard. Bull. Singapore 68(1) 2016

Key to Aeschynanthus species recorded from Peninsular Malaysia

1 a. Leaves in whorls of 3 or more 2

lb. Leaves opposite 3

2a. Corolla predominantly green to yellow, 18-25 mm long; leaves extremely

variable from ovate to linear, 1.7-43 times as long as wide

2. A. angustifolius

2b. Corolla predominantly red and orange, 54-1 1 8 mm long; leaves ovate to elliptic,

never linear, 2.3-6. 1 times as long as wide 12.4. speciosus

3a. Inside of corolla tube with dense multicellular hairs towards base, often in tufts;

leaves sometimes conspicuously variegated; corolla at least in part green, yellow

or yellowish green; seeds with > 2 hairs at hilar end 4

3b. Inside of corolla tube without dense multicellular hairs towards base; leaves not

conspicuously variegated; corolla entirely red or orange, red/orange and yellow,

or a combination of colours but never green; seeds with 1 hair at hilar end 7

4a. Calyx fused into a short tube at base 1.4. albidus

4b. Calyx lobes free to base 5

5a. Leaves not variegated, 1-2.4 times as long as wide, margin often distinctly dentate

3.4. dischidioides

5b. Leaves variegated, 1.7-9 times as long as wide, margin entire or very minutely

and obscurely dentate 6

6a. Corolla predominantly green, 20.5-31 mm long; stamens exserted from corolla

tube 7.4. longicaulis

6b. Corolla red in upper third and on lobes, 14.5-19 mm long; stamens not exserted

from corolla tube 4. A.fecundus

Idi. Calyx lobes free to base 6. 4. longiflorus

7b. Calyx fused into a tube for part or most of length 8

8a. Calyx in a wide cup, very much wider at apex than at base, lobes often barely

discemable but if obvious then much wider than long; seeds with bubble cells ....

9

8b. Calyx tube clasping corolla tube or only gently ftaring from base, lobes usually

easily discemable, either wider than long or longer than wide; seeds with or

without bubble cells 10

9a. Calyx red, 8-19 mm long 8. 4. obconicus

9b. Calyx green, 2. 5-6. 5 mm long 14. 4. wallichii

Aeschynanthus in Singapore and Peninsular Malaysia

7

10a. Calyx lobes usually longer than wide, apex acute to acuminate (tips of lobes

sometimes rounded); seeds without bubble cells 11

10b. Calyx lobes usually wider than long, apex rounded; seeds with bubble cells ... 12

lla. Stamens not or barely exserted from corolla tube; corolla lobes spreading or

reflexed 11.^. rhododendron

llb. Stamens strongly exserted from corolla tube; corolla lobes not reflexed,

occasionally slightly spreading 5. A. fulgens

12a. Corolla 19.5-27 mm long 13. ^4. volubilis

12b. Corolla 42-66 mm long 13

13a. Ovary, stipe and style densely pubescent; leaves sparsely to densely pubemlent

beneath 1 0 . ^4. radicans

1 3b. Ovary with sessile glands, only stipe and style pubescent; leaves usually glabrous,

more rarely sparsely puberulent beneath 9. A. pulcher

Aeschynanthus albidus (Blume) Steud., Nomencl. Bot. ed. 2 1: 32 (1840); A.DC.,

Prod. 9: 262 (1845); Bakhuizen van den Brink, Blumea 6: 395 (1950); Backer &

Bakliuizen van den Brink, FI. Java 2: 523 (1965); Burtt & Woods, Notes Roy. Bot.

Card. Edinburgh 33: 479 (1975); Turner, Card. Bull. Singapore 47: 243 (1997

[‘1995’]); Lok & Tan, Nat. Singapore 1: 5 (2008). - Bignonia albida Blume, Verh.

Batav. Genootsch. Kunsten 9: 195 (1823). - Trichosporum albidum (Blume) Nees,

Flora 8: 144 (1825). - Lysionotus albidus (Blume) Blume, Bijdr. FI. Ned. Ind. 765

(1826). - Aeschynanthus purpurascens Hassk., Cat. Hort. Bot. Bogor. 154 (1844),

nom. illegit; Hooker, Bot. Mag. 72: T.4236 (1846); Miquel, FI. Ned. Ind. 2: 717 (1858);

C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 37 (1883); Ridley, FI. Mai. Pen.

2: 497 (1923); Henderson, Malay. Wild. FI. Dicot. 340 (1959); Turner, Gard. Bull.

Singapore 45: 92 (1993). - TYPE: Indonesia, Java, Jawa Barat, Gunung Salak, Blume,

C.L. s.n. (lectotype L [L0003309], designated here). (Fig. 1, 2)

1 Aeschynanthus atropurpureus Van Houtte, Hort. Vanhoutt. 1(2): 42 (1846); Miquel,

FI. Ned. Ind. 2: 718 (1858). - TYPE: Not known. No original material known but

Miquel’s later description is speciflc enough to identify the taxon and it has long been

placed in synonymy of Aeschynanthus albidus by other authors.

Aeschynanthus discolor T.yiooxQ, Paxton’s FI. Gard. 3: 55 (1852). - TYPE: Not found.

Synonymy based on detailed description given in protologue.

Aeschynanthus motleyi C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 20 (1883);

Ridley, J. Linn. Soc. Bot. 32: 500 (1896); Ridley, J. Straits Branch Roy. Asiat. Soc.

44: 12 (1905); Ridley, J. Asiat Soc. Bengal, Pt. 2, Nat. Hist 74(2): 732 (1909). -

Trichosporum motleyi (C.B.Clarke) Kuntze, Revis. Gen. PI. 478 (1891). - TYPE:

8

Gard. Bull. Singapore 68(1) 2016

Indonesia, Kalimantan, Kalimantan Selatan, Banjarmasin, Motley, J. 916 (lectotype K

[K000831891], effectively designated by Burtt & Woods (1975)).

Aeschynanthus motleyi van sumatrensis C.B. Clarke in A.DC. & C.DC., Monogr.

Phan. 5(1): 20 (1883). - TYPE: Indonesia, Sumatra, Sumatera Barat, Padang, Ayer

Mancior, Beccari, O. 823 (lectotype K [K000831892], designated here; isolectotypes

BM [BM000537118], FI [FI013073], K [K000831893], L [L0281671], MEL).

Aeschynanthus fraserianus KraenzL, J. Linn. Soc. Bot. 37: 284 (1906). - Trichosporum

fraserianum (KraenzL) Merr., J. Straits Branch Roy. Asiat. Soc. special number:

530 (1921). - TYPE: Indonesia, Kalimantan, Marisinsing, Fraser 268 (holotype K

[K000831890]).

Epiphyte with stems upright, arching or pendulous; stems green flushed purple,

glabrous. Leaves opposite; petiole 3-13 mm long, purple, glabrous; blade slightly fleshy

or coriaceous, obovate, elliptic or ovate, above green and sometimes with paler green

or yellow mottling, beneath green with purple-red mottling or completely purple-red,

2.3-13.7 X 1. 1-5.5 cm, 1. 5-9.4 times as long as wide, apex acuminate, base rounded

to cuneate, glabrous above and beneath, margin vei'y weakly crenate, c, 3 pairs of

secondary veins, obscure or weakly visible, tertiary venation obscure. Inflorescence

subterminal or axillary, 1-5-flowered; peduncle c. 1 mm long; pedicels 7-10 mm long,

glabrous. Calyx with a short tube at base, lobes free, tube clasping corolla tube at base,

often slightly narrower at apex, green, green flushed with purple or red or entirely

reddish, glabrous or sparsely eglandular pubemlent, total length 14-30 mm long; tube

2-8.5 mm long which is 14-39% of total length, 3-6.5 mm wide at top of tube; lobes

linear or naiTOwly triangular, slightly spreading or erect, 10.5-22 x 1-3.5 mm, apex

acuminate. Corolla 1 6-29 mm long, tube broad at base, slightly curved, externally

green or greenish yellow, lobes green or green with faint purple central lines, internally

green, lobes green with red speckling or with dark purple chevrons and lines; upper

lobes orbicular, not spreading or reflexed, 2.2-3. 1 x 2. 6-3. 5 mm, sinus 2.5-3 nmi deep,

apex rounded; lateral lobes orbicular, slightly spreading or not, 2. 1-2.9 x 2. 9-4. 5 mm,

apex rounded; lower lobe ovate or orbicular, slightly spreading, 2. 7-3. 5 x 2. 5-2, 8 mm,

apex rounded; glabrous except for ciliate lobes, inside with five tufts of multicellular

hairs near base, sometimes with some additional multicellular hairs higher up, sessile

glands present at top of tube. Stamens long exserted, fused in 2 pairs; filaments cream

or green, glandular pubescent, anthers pink; anterior filaments inserted at 9-14.5 mm

from corolla base which is 48-53% of corolla length, filaments 21.5-23.5 mm long,

anthers 2. 7-3.2 x 0.8-1 .3 mm; posterior filaments inserted at 9.5-14 mm from corolla

base which is 50-58% of corolla length, filaments 19-20.5 mm long, anthers 1.2-2. 3

X 0. 7-1.1 mm; staminode 0.7-1. 6 mm long. Disk 1.2-1. 5 mm high, 5-crenate. Pistil

16-26 mm long; stipe c. 2-3 mm long, with sessile glands; ovary 7-12 mm long, with

sessile glands; style cream or green, 8-12 mm long, glandular puberulent; stigma pale

red, 2 mm across. Capsule 11.5-40 cm long, 2.5^ mm wide. Seed grain 1.5 - 2.4 x

0.4-0. 5 mm, warty, bubble cells absent; apical appendage a filiform hair, 13-21 mm

Aeschynanthus in Singapore and Peninsular Malaysia

9

Fig. 1. A. Aeschynanthus albidus (Blume) Steud. B. Aeschynanthus fecundus P. Woods. C.

Aeschynanthus fulgens Wall, ex R.Br. (Photos: A, Saw Leng Guan; B-C, David Middleton)

long; hilar appendages of many (30-40) filiform hairs, 14-19 mm long; appendages

papillose.

Distribution. Peninsular Malaysia (Johor, Kelantan, Pahang, Perak, Selangor,

10

Gard. Bull. Singapore 68(1) 2016

Fig. 2. Distribution of Aeschynanthus albidus (Blume) Steud. in Singapore and Peninsular

Malaysia (•).

Terengganu), Singapore, Sumatra, Java, Borneo.

Habitat and ecology. In lowland mixed dipterocarp forest, sometimes by streams, or in

lower montane forest at 20-1440 m altitude (to 1520 m on Mt Kinabalu).

Provisional lUCN conservation assessment. Least Concern (LC). This species is

widespread and locally fairly common. In Peninsular Malaysia it has been collected

over a fairly wide area in the last 20 years. In Singapore it is listed as nationally Critically

Endangered by Chong et al. (2009). Although it was fonnerly more widespread it is

now known in Singapore only in Nee Soon swamp forest.

Singaporean and Peninsular Malaysian specimens examined. PENINSULAR MALAYSIA:

Johor: Gunong [Gunung] Ledang VJR, 19 Aug 1974, Kochummen, K.M. FRI 16796 (K, KEP,

SING); Lenggor Forest Reserve, Jong, K. 9017 (KLU); Batu Pahat, Nov 1891, Nongchi 8

Aeschynanthus in Singapore and Peninsular Malaysia

11

(SING); Kota Tinggi, 1 Oct 1929, Teruya, Z. 951 (SING); Bekok, Sungei Bekok, 7 Mar 1971,

Heaslett, E.A. s.n. (SING); Kelantan: Kuala Aring, 7 Sep 1899, Yapp, R.H. 161 (CGE, K);

Sungai Keteh, 6 Feb 1924, Md Nur & Foxworthy, FW 11963 (SING); Gua Musang, Lojing

FR, 860 m, 27 Apr 201 1, Mohd Hairul, M.A., Sid Munirah, M. Y. & Mohd Nazri, A. FRI72324

(KEP); Pahang: Jalan Bentong bt. 18, 13 Sep 1973, MK & AR 1321 (L, UKMB); Fraser’s Hill,

700 m, 10 Mar 1995, Chin, S.C. et al 4533 (SING); Pulau Manis, Jul 1 891, Ridley, H.N. 2151

(SING); Sabai Estate near Bentong, 26 Jan 1958, Shah, M. 155 (K, L, SING); Kuala Lipis, 22

Nov 1 924, Burkill, H.M. & Md Haniff 1 5750 (SING); Pulau Tioinan, Kpg. Juara, 371 m, 25 Apr

2012, Saw, L.G. & Mohd Hairul, M.A. FRI48326 (E, KEP); Perak: Gopeng, 150-240 m, Jun

1883, Kunstler, H. 4463 (NY, SING); Dipang, Mar 1885, Scortechini, B. 1815 (SING); Kurau,

Wray, L. 4245 (SING); Taiping, Gunung Hijau, 1440 m, 19 Mar 2007, Julius, A. FR153313

(E, KEP); Taiping, Bukit Larut, Jun 1893, Ridley, H.N. s.n. (SING); Kuala Kangsar, Bubu

FR, Gunung Bubu, 12 Mai* 2010, Julius, A., Coode, M.J.E. & Angan, A. FR157691 (KEP);

Selangor: Genting Highlands, Ulu Gombak, 8 Apr 1921, Hume, H.L. 9596 (SING); ibidem, 16

Apr 1921, Hume, HE. 9772 (SING); 17 mile Ulu Gombak, 26 Oct 1937, MdNur SFN34254

(SING); Terengganu: Gunong [Gunung] Lawit, Old trail along Sungai Kamiah, 300-610 m, 1

Apr 1970, Davidson, C. 1292 (L).

SINGAPORE: s.I, 1894, Ridley, HN. s.n. (SING); Seletar, 1894, Ridley, H.N. 6244 (SING);

ibidem, 3 Sep 1889, Ridley, H.N s.n. (SING); Bukit Mandai, Mar 1890, Ridley, H.N. s.n.

(SING); Chan Chu Kang, Jul 1891, Ridley, H.N. s.n. (SING); ibidem, 1892, Ridley, HN. s.n.

(SING); Teban, 17 May 1891, Goodenough, J.S. s.n. (SING); Krangi [Kranji], 1894, Ridley,

H.N. s.n. (SING); Nee Soon Swamp Forest, 22 Nov 2007, Ng, HH. & Lok, A.F.S. L. s.n. (SINU);

Nee Soon Swamp Forest, 25 Mar 2015, Lua, H.K. & Ibrahim, H. S1NG201 5-084 (SING).

Notes. A very variable species, particularly in the size and relative dimensions of the

calyx.

Burtt & Woods (1975) gave the type as “Type: cult. Hort. Bogor (L?)”.

However, in the proto logue two collections are mentioned, one from “Tjoo” and one

from “Salak”, There is no mention of cultivated material from Bogor so the material

designated as type by Burtt & Woods (1975) cannot be considered original material. A

Blume collection from Salak is, therefore, here chosen as lectotype.

This species has generally been called Aeschynanthus purpurascens or, less

frequently, A. motley i^ in the literature from the Malay Peninsula. Aeschynanthus

purpurascens is an illegitimate name as Aeschynanthus albidus and several other

earlier combinations are included m synonymy. Aeschynanthus motley was described

from Borneo but the Peninsular Malaysia, Borneo, Sumatra and Java material cannot

be distinguished.

2. Aeschynanthus angustifolius (Blume) Steud., NomencL Bot. ed. 2 1: 32 (1840);

A.DC., Prod. 9: 262 (1845); Miquel, FI. Ned. Ind. 2: 716 (1858); C.B.Clarke inA.DC.

& C.DC., Monogr. Phan. 5(1): 38 (1883); Bakhuizen van den Brink, Blumea 6: 395

(1950); Backer & Bakliuizen van den Brink, FI. Java 2: 524 (1965); Woods, Kew Mag.

8(1): 22 (1991); Turner, Gard. Bull. Singapore 47(1): 243 (1997 [A995']). - Bignonia

angustifolia Blume, Catalogus 82 (1823). - Trichosporum angustifolium (Blume)

Nees, Flora 8: 144 (1825). - Lysionotus angustifolius (Blume) Blume, Bijdr. FI. Ned.

12

Gard. Bull. Singapore 68(1) 2016

Ind. 765 (1826). - TYPE: Indonesia, Java, Jawa Barat, Bogor, Reinwardt, C.G.C. s.n.

(lectotype L, designated here). (Fig. 3, 4)

Rheitrophyllum subverticillatum Hassk., Flora 25(2): beibl. 56 (1842). - TYPE: Java,

Bogor Botanic Garden, 21 August 1840, Hasskarl s.n. (not traced). Although the type

has not been traced the description of the leaves in the protologue is suitably diagnostic

to be certain that it is this taxon.

Aeschynanthus tetraquetrus C.B. Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 38

(1883). - Trichosporum tetraquetrum (C.B. Clarke) Kuntze, Revis. Gen. PI. 478 (1 891).

-TYPE: Indonesia, Sumatra, Sumatera Barat, Gunung Singgalan, 1700 m, Beccari, O.

s.n. (lectotype FI [F1013076], designated here; isolectotypes FI [FI013075], K).

Aeschynanthus stenophyllus Ridl., J. Asiat. Soc. Bengal 74 (2): 733 (1909); Ridley, FI.

Mai. Pen. 2: 498 (1923). - Trichosporum stenophyllum (Ridl.) Merr., Contr. Arnold

Arbor. 8: 152 (1934). - TYPE: Peninsular Malaysia, Perak, Goping [Gopeng], Kinta,

150-240 m, August 1883, Kings Collector 4738 (lectotype SING [SING0035634],

designated by Woods (1991); isolectotype K).

Trichosporum ternifolium Merr., Contr. Arnold Arbor. 8: 153 (1934). - TYPE:

Indonesia, Sumatra, in ravines between Baboeli and Paekas, 1200 m, 9 January

1932, Bangham, W.N. & Bangham, C.M. 781 (holotype A [A00261780]; isotype NY

[NY003 13032]).

Epiphyte with pendulous or arching stems; stems glabrous. Leaves verticillate in

whorls of 3-7, rarely opposite; petiole 1-6 mm long, glabrous; blade coriaceous or

slightly fleshy, ovate or elliptic to linear, 1-8.6 x 0.15-1.7 cm, 1.7^3 times as long as

wide (see note below), apex obtuse to acuminate, base cuneate, mid green above, paler

beneath, not variegated, glabrous above and beneath, margin weakly and distantly

dentate or entire, secondaiy veins obscure, tertiary veins obscure. Flowers solitary

in the axils of leaves, bracts minute; pedicels 9.5-10 mm long, glabrous. Calyx of

separate lobes free to base, green, glabrous except with very few hairs at tips of lobes;

lobes nan*owly triangular, 1. 5-4.2 x o. 7-0.9 mm, apex acute, glabrous. Corolla 18-25

mm long, tube fairly narrow, slightly wider at base, fairly straight, curved at apex,

externally yellowish to yellowish green to green, margins of lobes reddish brown,

internally yellowish green, reddish brown on margins of lobes and speckled on inner

lobes; upper lobes oblong, not spreading or reflexed, 2.2-3. 8 x 2.2-3. 2 mm, sinus 2.6-

2.8 mm deep, apex rounded; lateral lobes oblong, not spreading or reflexed, 3. 5-4. 3 x

3-3.2 mm, apex rounded; lower lobe oblong, slightly spreading, 3.5-6 x 2.5-4 mm,

apex rounded; tube and outside of lobes minutely glandular puberulent, lobes ciliate,

inside of lobes and tube with scattered short glandular hairs. Stamens shortly exserted,

fused in 2 pairs; filaments glandular pubescent; anterior filaments inserted at 11-15

mm from corolla base which is 61-65% of corolla length, filaments 10-12 mm long,

anthers 1.2-1. 5 x 0.8-0. 9 mm; posterior filaments inserted at 12-16 mm from corolla

Aeschynanthus in Singapore and Peninsular Malaysia

13

Fig. 3. Aeschynanthus angustifolius (Blume) Steud. A. Habit with yellow-flowered form. B.

Habit with green-flowered form. C. Flower disseetion with three ealyx lobes removed. D.

Flowers from the front showing stamens in two pairs and developing style and stigma in the

flower on the left and a more developed style and the stamens beginning to wither and reflex in

V

the flower on the right. (Photos: A, David Middleton; B-D, Jana Leong-Skornickova)

14

Gard. Bull. Singapore 68(1) 2016

Fig. 4. Distribution of Aeschynanthus angustifoHus (Blume) Steud in Peninsular Malaysia (•).

base which is 67-73% of corolla length, filaments 8.5-9 mm long, anthers 0.8-1. 2

X 0. 6-0.7 mm; staminode c. 0.5 mm long. Disk 1.2-1. 5 mm high, margin 5-crenate.

Pistil 28-30 mm long, pale green; stipe 5-6 mm long, with sparse sessile glands; ovary

8-9 nmi long, with sparse sessile glands; style c. 15 mm long, with short glandular

hairs. Capsule 13-30 cm long, 2.4-3. 3 mm wide. Seed grain 1.6-1. 9 x 0.3 mm, bubble

cells absent; apical appendage a filiform hair 44-50 mm long; hilar appendage a single

filiform hair, 28-30 mm long.

Distribution. Peninsular Malaysia (Johor, Pahang, Terengganu), Sumatra, Java, Borneo.

Habitat and ecology. Primary or secondary lowland to lower montane forest. In

Peninsular Malaysia recorded from 150-1390 m altitude and in Sumatra recorded up

to 1700 m.

Aeschynanthus in Singapore and Peninsular Malaysia

15

Provisional lUCN conservation assessment. Data Deficient (DD). The ECO and AOO

for this species using the existing collections would suggest an assessment of Least

Concern. However, all of the collections I have seen of this species collected in the

last 20 years have been made only in Borneo over a range that would suggest Near

Threatened. 1 have seen no collections from Peninsular Malaysia made since 1930 and

its current status there is uncertain. It has never been collected in Singapore.

Additional Peninsular Malaysian specimens examined. PENINSULAR MALAYSIA: Johor:

Sedenak, Nov 1909, Md Nur s.n. (SING); ibidem, 1909, Ridley, H.N. s.n. (SING); Pahang:

Cameron Highlands, 1390 m, 1 Apr 1930, Henderson, M.R. s.n. (NY); Terengganu: Kuala

Berang, 14 May 1925, Holttum, R.E. 15312 (K, SING).

Notes. Leaf characters are extremely variable within this species even though most

material does show linear leaves > 7 times as long as wide. Wide variation occurs even

within individuals as was previously commented upon by Woods (1991). For example,

Poulsen et al. 1566 (KEP) from Long Maga, Ulu Padas, Sabah, has leaves from 1 .7-30

times as long as wide on a single branch. In this case there appear to have been two

distinct phases of growth with older, shorter and wider leaves on the older part of the

branch and then a new flush of growth with linear leaves on the younger part of the

branch.

This species has been described as several different species from the different

parts of its range due to differences in corolla colour and leaf shape but there are no

clear discontinuities in these characters across its range.

3. Aeschynanthus dischidioides (Ridl.) D.J.Middleton, Edinburgh J. Bot. 64: 425

(2007). - Micraeschynanthus dischidioides Ridl., FI. Malay Penin. 5: 325 (1925).

- TYPE: Peninsular Malaysia, Pahang, Gunung Tahan, 1670 m, Ridley, H.N. 16122

(lectotype K, designated by Middleton (2007); isolectotype SING [SING0089714]).

(Fig. 5, 6)

Aeschynanthus myrmecophilus P. Woods, Notes Roy. Bot. Gard. Edinburgh 33: 483

(1975); Turner, Gard. Bull. Singapore 47(1): 243 (1997 [‘1995’]). - TYPE: Peninsular

Malaysia, Pahang, Cameron Highlands, Robinson’s Falls, 1500 m, 16 April 1968,

Woods, P.J.B. d/d(holotype E [E00062778]).

Aeschynanthus hildebrandii auct. non Hemsl. ex Hook.f : Ridley, J. Linn. Soc, Bot.

32: 502 (1896); Ridley, J. Straits Branch Roy. Asiat. Soc. 44: 15 (1905); Ridley, J.

Asiat. Soc. Bengal 74(2): 734 (1909); Ridley, FI. Mai. Pen. 2: 499 (1923).

Epiphyte, often rooted in ants nests, with stems hanging; stems sparsely puberulent

or glabrous. Leaves opposite (although sometimes appearing whorled due to vei 7

short intemodes); petiole 1-3 mm long, purple, glabrous; blade thickly coriaceous,

green edged with puiple above, not variegated, green to reddish puiple beneath, ovate

16

Gard. Bull. Singapore 68(1) 2016

or orbicular, 0. 7-2.4 cm long, 0. 6-2.6 cm wide, 1-2.4 times as long as wide, apex

rounded to acuminate, base rounded or subcordate, glabrous above and beneath, not

punctate beneath, margin dentate or entire, undulate or not, secondary veins obscure,

tertiary veins obscure. Inflorescence axillary or terminal, 1-6-flowered; pedicels 8.5-

16 mm long, green or puiple, with sparse and minute hairs, smaller ones more or less

papillae, hairs (papillae) green or red in life. Calyx of separate lobes free to base,

green, with sparse and minute hairs; lobes narrowly ovate or narrowly triangular, erect,

2-6.5 X 0.6-1. 6 mm, apex acute. Corolla 1 1-25 mm long, tube broad for all of length,

slightly to strongly curved, outside of tube yellow to yellow-orange for most of length

and then abruptly turning to dark orange or red on top of tube and lobes, sometimes

only on lobes, inside of tube yellow, lobes dark red to orange-red at margin and yellow

at base with dark bands on lower 3 lobes, outside sparsely puberulent, denser on

ciliate lobes, inside with a ring of robust multicellular hairs in lower half; upper lobes

oblong, not spreading or reflexed, conspicuously shorter than neighbouring lateral

lobes, 2.6^ X 2. 5^. 8 mm, sinus 3.2-4 .2 mm deep, apex rounded; lateral lobes ovate

or oblong, not spreading or reflexed, 3.6-6 x 2. 9-5. 5 mm, apex rounded, truncate or

retuse; lower lobe oblong, not spreading or reflexed, 4.9-6 x 3-5.8 mm, apex rounded

to truncate. Stamens clearly exserted, fused in 2 pairs; filaments various shades of

green to red, posterior pair glandular hairy at apex and papillose at base, anterior pair

only papillose, anthers grey; anterior filaments inserted at 8-1 1 mm from corolla base

which is 44-62% of corolla length, filaments 12-16 mm long, anthers 1.4-2 x 0.6-1. 2

mm; posterior filaments inserted at 9.5-11.5 mm from corolla base which is 48-69%

of corolla length, filaments 10.5-13 mm long, anthers 1. 1-1.6 x 0.5-1 mm; staminode

0.7-1 mm long. Disk 1 .2-1 .5 mm high, 5-dentate. Pistil 21-24 mm long; stipe 2.5-3. 5

mm long, glabrous or with sparse sessile glands; ovary green, 6-6.5 mm long, with

sessile glands, these sometimes sparse; style 13-14 mm long, glandular pubescent.

Capsule 8-24 cm long, 2.7-3 mm wide. Seed grain 1.2-1 .6 x 0. 3-0.4 mm, warty,

bubble cells absent; apical appendage a filifonn hair,. 10-50 mm long; hilar appendage

of 3-6 filifonn hairs, 17-30 mm long.

Distribution. Endemic in Peninsular Malaysia (Pahang, Perak, Selangor).

Habitat and ecology. In lower montane forest at 1100-2000 m altitude, often growing

in ant gardens associated with Lecanopteris or Dischidia.

Provisional lUCN consen’ation assessment. Vulnerable (VU Blab(iii)). The known

EOO is around 15,000 km^ (but see note below) and the locations where it occurs are

disturbed by fanning, plantations and tourism.

Additional Peninsular Malaysian specimens examined. PENINSULAR MALAYSIA:

Pahang: Cameron Highlands, 1390 m, 1 Apr 1930, Henderson, M.R. s.n. (NY, SING); ibidem,

9 Mar 1947, Smith, J.W. 63690 (KEP, SRJG); ibidem, 14 Oct 1929, Symington, C.F. 20938

(KEP); ibidem, 1976, Anthony, S. SA230 (KEP); Cameron Highlands, Along route no. 7 to G.

Beremban, 1500 m, 3 Mar 1994, Perumal, B., Gan, C.L., Shahril, K.Z., Angan, A. & Bedul

FRI41631 (KEP, KLU); Cameron Highlands, Gedung FR, Near Sg. Ichat tea plantation, 1365

Aeschynanthus in Singapore and Peninsular Malaysia

17

Fig. 5. Aeschynanthus dischidioides (Ridl.) D.J.Middleton. A. Habit. B. Flower dissection

showing multicellular hairs in corolla tube and undeveloped style after two calyx lobes

removed. C. Flower from below with stamens fused in two pairs. D. flower, side view. (Photos:

A, David Middleton; B-D, Jana Leong-Skomickova)

18

Gard. Bull. Singapore 68(1) 2016

Fig. 6. Distribution of Aeschynanthus dischidioides (Ridl.) DJ.Middleton in Peninsular

Malaysia (•).

m, 19 Feb 2008, Syahida-Emiza & Chew, M.Y. FRI57277 (KEP); Cameron Highlands, G. Jasar,

9 Aug 2008, Rosdi, M. & Phoon, S.N. et al. FRI59S72 (KEP); Cameron Highlands, Gunung

Batu Berinchang, Kpg. Melayu Brinchang, 22 Feb 2012, Imin, K. & Syahida-Emiza FRI76065

(KEP); Cameron Highlands, Gunung Batu Berinchang, 2000 m, 20 Jun 1975, Van Balgooy,

M.M.J. 2658 (E, L); Cameron Highlands, Boh Tea Estate, 29 Apr 1937, Md Nur SFN32980

(SING); ibidem, 28 Jul 1991, Kiew, R. RK3241 (SING); Cameron Highlands, Tanah Rata,

18 Aug 1986, Weber. A. s.n. (KEP); ibidem, 1440 m, 9 Sep 1956, Burkill H.M HMB868

(SING); Cameron Highlands, Tanah Rata, Parit Waterfall, 1457 m, 26 Sep 2012, Imin, K, et al.

FRI77536 (E, KEP, SING); Cameron Highlands, Tanah Rata, Jin Tenkolok, 1489 m, 23 Feb

2012, Imin, K. & Syahida-Emiza FRI76073 (KEP); Cameron Highlands, Robinson’s Falls, 5

Apr 2006, Kiew, R. RK5307 (SING); Cameron Highlands, en route from Robinson Waterfall

to Boh tea plantation, 600-1100 m, 29 Aug 1990, Okada, H., Darnaedi, D., Akiyama, H.,

Kawahara, T & Watano, Y. 7 Ddt) (TI); Taman Negara, 1 100-1530 m, 18 Aug \990, Darnaedi,

D., Akiyama, H., Kawahara, T. & Khairuddin, K 566 (TI); Taman Negara, Kem Bonsai,

From Kem Bonsai to Kem Belumut, 1985 m, 8 May 2008, Yao, T.L. FRI65302 (E, KEP);

Aeschynanthus in Singapore and Peninsular Malaysia

19

Taman Negara, Gunung Tahan, 1820ni, 1 Sep 1928, Holttum, R.E. s.n. (SING); ibidem, 7 Jul

1905, Wray, L. & Robinson, H.C. 5483 (BM, K, SING); ibidem, Jul 1911, Ridley, H.N. s.n.

(SING); Gunung Tahan, Foot of Gunung Gedong, 1520 m, 3 Sep 1928, Holttum, R.E. 20772

(SING); Gunung Tahan, Skeats Hill, Jul 1911, Ridley, H.N. s.n. (SING); Fraser’s Hill, Pine tree

hill, 1450 m, 19 Apr 1955, Purseglove, J.W. P4222 (K, SING); ibidem, 1450 m, 3 Sep 1966,

Burkill, H.M. HMB4221 (SING); Padang Camp, 1670m, 14 Sep 1937, Corner, E.J.H. s.n. (L);

Kluang Tabang, 1900, Barnes, E. s.n. (SING); Perak: Gunong [Gunung] Jasar, 17 Apr 1968,

Woods, PJ.B. 634 (E); Taiping, 1210m, 14Feb 1917, Md Haniffd Md Niir 2469 (K, SING);

ibidem, 1240 m, 14 Feb 1917, Md Haniff & MdNur 2347 (SING); Taiping, Bukit Larut, Jun

1893, Ridley. H.N. & Curtis, C. 7365 (SING); Kuala Kangsar, Bubu FR, Gunung Bubu, 1675

m, 20 Dec 2006, Kamarul Hisham, M., Kiieh, H.E, Lim, C.L. & Yao, T.L. FRI52094 (KEP);

Selangor: Ulu Kali, 1600 m, 2 Mar 1996, Van Balgooy M.M.J. 7149 (L).

Notes. This is currently the only species of Aeschynanthus considered to be endemic

to Peninsular Malaysia. However, considering that most of its distribution closely

mirrors that of Aeschynanthus rhododendron, suggesting similar habitat requirements,

it may also occur in the far south of Thailand and in Sumatra where A. rhododendron

has been collected.

The combination in Aeschynanthus was published by Middleton (2007)

when the monotypic Micraeschynanthus dischidiodes was synonymised with

Aeschynanthus myrmecophilus . Middleton (2007) concluded that the type material

of Micraeschynanthus dischidiodes has only immature and aben*ant flowers but that

in all other characters it matches Aeschynanthus myrmecophilus, thereby requiring

synonymisation and a new combination in Aeschynanthus.

4. Aeschynanthus fecundus P. Woods, Notes Roy. Bot. Gard. Edinburgh 33: 482

(1975); Turner, Gard. Bull. Singapore 47: 243 (1997 [‘1995’]); Burtt, Thai Forest Bull.,

Bot. 29: 83 (2001); Smitinand, Thai PI. Names ed. 2, 14 (2001); Middleton, Edinburgh

J. Bot. 64: 379 (2007). - Aeschynanthus pafMflorus Ridl., J. Fed. Malay States Mus.

4: 48 (1909), nom. illeg. - Aeschynanthus breviflorus Ridl., FI. Mai. Pen. 2: 497

(1923), nom. illeg. - TYPE: Peninsular Malaysia, Pahang, Telom, November 1900,

Ridley H.N. 13599 (holotype SING [SING0089822]; isotypes BM [BM000537160],

K[K000190151]). (Fig. 1,7)

Epiphyte with erect, arching or pendulous stems; stems green, flushed purplish or

reddish brown, glabrous. Leaves opposite; petiole green, 2-4 mm long, glabrous;

blade slightly fleshy, green with paler green or yellow variegation above, pale green

beneath, elliptic, 2. 1-8.6 x 0.5-3. 7 cm, 1 .7-5.3 times as long as wide, apex acuminate,

base rounded to cuneate, glabrous above and beneath, not punctate beneath, margin

with few minute red-tipped teeth but appearing more or less entire, secondary veins

obscure, tertiary venation obscure. Inflorescences axillary, 1-2 -flowered, peduncle

absent; pedicels green, 4-7 mm long, sparsely minutely papillose or glabrous. Calyx

of separate lobes free to base, yellow-green or green, tips red, glabrous or with sparse

sessile glands; lobes linear to narrowly ovate, slightly spreading or erect, 2.3-13.5

20

Gard. Bull. Singapore 68(1) 2016

Fig. 7. Distribution of Aeschynanthus fecundus P. Woods (•) and Aeschynanthus fulgens Wall,

ex R.Br. (T) in Peninsular Malaysia

X 0.6-1 .5 mm, apex acute or acuminate. Corolla 14.5-19 mm long, externally tube

yellowish green or yellow in lower two-thirds, brownish red higher, lobes dark red or

brownish red, internally tube light yellowish, lobes pale brownish red, tube fairly broad

at base; upper lobes oblong or elliptic, not spreading or reflexed, 1 .1-2.3 x 1 .2-2.3 mm,

sinus 1.4-2. 6 mm deep, apex rounded; lateral lobes ovate, not spreading or reflexed,

1.1-2. 5 X 1.6-2. 9 mm, apex rounded; lower lobe elliptic, not spreading or reflexed,

1.3-2. 3 X 1-2.6 mm, apex rounded; glabrous or with few sessile glands around top

outside, with irregular tufts of multicellular hairs near base and glandular papillose

on ventral surface near throat inside. Stamens not exserted, fused in 2 pairs, filaments

yellow or greenish yellow, with glandular hairs, anthers yellow to grey, pollen cream;

anterior filaments inserted at 7-8.6 mm from corolla base which is 45-48% of corolla

length, filaments 7-9 mm long, anthers 1-1.3 x 0.6 mm; posterior filaments inserted at

8-12 mm from corolla base which is 55-63% of corolla length, filaments c. 6 mm long.

Aeschynanthus in Singapore and Peninsular Malaysia

21

anthers 0.7-0. 9 x 0. 5-0.6 mm; staminode 0.8-2 mm long. Disk 1 mm high, a simple

annular ring. Pistil 17-19 mm long; stipe c. 1.5 mm long, with few sessile glands;

ovary green, 4. 5-5. 5 mm long, with sessile glands; style green or yellowish green,

11-12 mm long, glandular pubeseent; stigma pink or purple. Capsule 3.6-1 2 cm long,

1.9-2. 8 mm wide. Seed grain 2-2.5 x 0.2-0. 3 mm, only weakly warty, bubble cells

absent; apical appendage a filiform hair, 19-28 mm long; hilar appendage of 11-18

filiform hairs, 12-17 mm long, appendages papillose.

Distribution. Western and southern Thailand, Peninsular Malaysia (Kedah).

Habitat and ecology’. In lowland and hill dipterocarp forest, sometimes on bamboo.

Altitude information is not available for the species in Peninsular Malaysia but in

Thailand it occurs up to 700 m altitude.

Provisional lUCN conservation assessment. Least Concern (LC). Even though this

species has been collected only once in recent years in Peninsular Malaysia, and only

twice ever, in Thailand it has a wide distribution and has been collected fairly often.

Additional Peninsular Malaysian specimens examined. PENINSULAR MALAYSIA: Kedah:

Baling, Gunung Inas Forest Reserve, Bukit Palong, 2 Nov 2007, Imin, K. et al. FRI58593 (K,

KEP, SING).

Notes. This species is difficult to tell apart from Aeschynanthus longicaulis when in

fruit. In Thailand this species was once found in fruit with the seeds already geraiinating

before the seeds had dispersed. It is not known if tliis was an aberration or coimnon

occurrence.

S. Aeschynanthus fulgensNfdiW. ex R.Br., Cyrtandreae 115 (1839); Steudel, Nomencl.

Bot. ed. 2, 1: 32 (1840); Brown in Bennett, PL Jav. Rar. 115 (1840); A.DC., Prod. 9:

261 (1845); C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 21 (1883); Clarke

in Hooker, FI. Brit. Ind. 4: 338 (1884); Burtt, Thai Forest Bull., Bot. 29: 83 (2001);

Smitinand, Thai PI. Names ed. 2, 14 (2001); Kress et ah, Checkl. Myanmar 261

(2003); Pooma, Threatened PL Thailand 70 (2005); Middleton, Edinburgh J. Bot. 64:

381 (2007); Middleton, Edinburgh J. Bot. 66: 414 (2009). — Trichosporum fulgens

(Wall, ex R.Br.) Kuntze, Revis. Gen. PL 477 (1891). - TYPE: Bunna, Tenasserim

Division, Tavoy, Gomez, W. in Wallich 797 (lectotype K-W, designated by Middleton

(2007); isolectotypes BM [BM000883874], CGE, E [E00259870, E00259871], G

[G00370743], G-DC, K [K000831871, K000831872]). (Fig. 1, 7)

Aeschynanthus evrardii Pellegr., Bull. Soc. Bot. France 72: 824 (1926 [1925]);

Lecomte, FI. Indo-Chine 4: 499 (1930); Pham Hoang Ho, Cayco Vietnam ed. 3, 3 (1):

4 (1993); Newman et al., Checkl. Vase. PL Lao PDR 146 (2007). - TYPE: Vietnam,

Lam Dong, Ang Kroet, 26 October 1920, Evrard, F. 358 (lectotype P [P00492372],

designated by Middleton (2007); isolectotypes P [P00606317, P00606318]).

22

Gard. Bull. Singapore 68(1) 2016

Aeschynanthus stenosiphonius W.T.Wang, Bull. Bot. Res., Harbin 3 (4): 49 (1983);

Burtt, Thai Forest Bull., Bot. 29: 84 (2001); Smitinand, Thai PI. Names ed. 2, 15

(2001). - TYPE: Laos, Vieng Pa Pow Tha Kaw, 2 November 1921, Hayata, B. s.n.

(holotype TI; isotype TI).

Hoya pseudolanceolata Costantin, FI. Indo-Chine 4: 132 (1912). - TYPE: Laos,

Sayabouri, Pak Lay, Spire 1490 (lectotype P [P00639825], designated here).

Aeschynanthus macranthus auct. non (Merr.) Pellegr.: Pellegrin, FI. Indo-Chine 4: 498

(1930) pro parte; Barnett, FI. Siam. 3 (3): 201 (1962); Burtt, Thai Forest Bull., Bot.

29: 84 (2001).

Epiphyte with stems arching and pendulous or upright, green or dull olive-green,

sometimes with purple mottling, glabrous. Leaves opposite; petiole 3-20 mm long,

glabrous; blade coriaceous to fleshy, elliptic, oblong or ovate, mid to dark green above,

paler green beneath, not variegated, 3.3-17 x 1-5.2 cm, 1.7-7. 5 times as long as wide,

apex acuminate, base cuneate to rounded, glabrous above and beneath, margin entire,

4-8 pairs of secondary veins, obscure to clearly visible, tertiary venation obscure.

Inflorescences subtenninal or axillary, 3-16-flowered, peduncle absent; bracts deep

maroon, ovate or linear, 2.1-11 mm long; pedicels 3.5-24 mm long, green, glandular

puberulent or glabrous. Calyx with a tube at base, lobes free, tube very variable in

width from nan'ow and clasping corolla tube to quite wide and not clasping corolla,

yellowish, greenish or brownish, sometimes with reddish lobes, glabrous except for

ciliate lobes, or few hairs only on very tips of lobes, to glandular pubescent all over,

total length (7-)l 0.7-26 mm long; tube (2.4-)6-18.5 mm long which is (34-)50-94%

of total length, 3.5-14 nun wide at top of tube; lobes triangular, slightly spreading or

erect, l-5(-l 1 .5) x (1 .2-) 1 .6-6.5 mm, apex acute, sometimes ultimately with a rounded

tip, to acuminate. Corolla (41-)49-73 nun long, tube nan'ow at base, externally bright

red to darker red or orange red, sometimes yellowish at base, usually with darker lines

on tube, red or orange-red with a black central line on lobes, sometimes yellowish at

very base, internally yellowish to pale red in tube, lobes orange-red or red with a dark

red or black centi'al line or arrow and pale orange to cream at base; upper lobes slightly

falcate, mostly not reflexed or spreading, sometimes slightly spreading but not fully

reflexed, 3. 2-7. 5 x 3.3-7 mm, sinus 3.5-1 0.5 mm deep, apex obtuse or rounded; lateral

lobes deltoid, not spreading or reflexed, 2.8-8 x 5.2-9. 5 mm, apex rounded or obtuse;

lower lobe oblong or elliptic, reflexed, 5.1-14 x 3-6 mm, apex obtuse or rounded;

sparsely to densely glandular puberulent outside or only at top of tube and on lobes

or only ciliate on lobes, minutely and sparsely to very sparsely glandular puberulent

internally, this sometimes denser towards the upper half, sessile glands present at top

of tube. Stamens long exserted, fused in 2 pairs or rarely all 4 attached; filaments

mostly darker in the upper half and ranging Irom reddish to purplish and then to white

at base, sparsely glandular pubescent to glabrous or with few sessile glands in lower

part, anthers dark maroon, purple, yellow, white and purple, or greyish purple; anterior

filaments inserted at 26^8 mm from corolla base which is 50-72% of corolla length.

Aeschynanthus in Singapore and Peninsular Malaysia

23

filaments 25-50 mm long, anthers 3-5.6 x 0.9-2. 3 mm; posterior filaments inserted

at 32-50 mm from corolla base which is 58-82% of corolla length, filaments 25^2

mm long, anthers 2.5-5 x 1-2.1 mm; staminode 0.7-11 mm long. Pollen ochre or

yellow. Disk 0.5-2. 5 mm high, a simple annular ring, 5-dentate, 5-crenate or strongly

5-lobed. Pistil 15.5-80 mm long; stipe 3-26 imn long, glabrous; ovary green or cream,

8-38 mm long, glabrous or with sessile glands; style cream or white, 4.5-41 mm

long, glandular pubescent to glabrous. Capsule 15.7-42 cm long, 2-5 mm wide. Seed

grain 0.8-2 x 0.2-0. 5 mm, warty, bubble cells absent; apical appendage a filiform hair,

16-34 mm long; liilar appendage a single filifonn hair, 13.5-35 mm long; appendages

papillose.

Distribution. Myanmar, Thailand, Cambodia, Laos, Vietnam, Peninsular Malaysia

(Kedah). In Peninsular Malaysia only recorded from Gunung Jerai in Kedah.

Habitat and ecology. Across its distribution it has been recorded in a wide variety of

habitats including primary or disturbed evergreen, mixed deciduous or mossy forest at

10-1500 m altitude. The ecology and altitude for this species on Gunung Jerai was not

recorded on the specimens.

Provisional lUCN conservation assessment. Least Concern (LC). This species has a

widespread distribution and is common in parts of its range, particularly in Thailand.

In Peninsular Malaysia it has only been collected a few times and at only one locality,

Gunung Jerai. Its current status there needs to be clarified.

Peninsular Malaysian specimens examined. PENINSULAR MALAYSIA: Kedah:

Gunung Jerai, 16 Oct 1976, Stone, B.C.M. 12758 (KLU); ibidem, 18 Nov 1977, Lo,

Y.N. & Mahmud 183 (KLU); ibidem, 23 Nov 1999, Kiew, R. RK4862 (KEP, SING).

Notes. This is a widespread and variable species but in Peninsular Malaysia has only

been collected on Gunung Jerai. The description above includes the species from

throughout its range as the very few Malaysian specimens are inadequate to write a full

description. The material seen from Malaysia has a long and narrow calyx tube with

small lobes but elsewhere in the distribution of Aeschynanthus fulgens this species is

known to be very variable, especially in this character.

6. Aeschynanthus longicaulis Wall, ex R.Br., Cyrtandreae 116 (1839); Steudel,

Nomencl. Bot. ed. 2, 1 : 32 (1840); Brown in Bennett, PI, Jav. Rar. 116 (1840); A.DC.,

Prod. 9: 262 (1845); Miquel, FI. Ned. Ind. 2: 719 (1858); C.B.Clarke in A.DC. &

C.DC., Monogr. Phan. 5(1): 19 (1883); C.B.Clarke in A.DC. & C.DC., Monogr. Phan.

5(1): 19 (1883); Barnett, El. Siam. 3 (3): 201 (1962); Burtt & Woods, Notes Roy.

Bot. Gard. Edinburgh 33: 481 {\91S),pro parte; Chin, Gard. Bull. Singapore 32: 147

(1979); Li, Acta Bot. Yunnan. 5 (1): 36 (1983); Wang, FI. Reipubl. Popularis Sin. 69:

526 (1990); Turner, Gard. Bull. Singapore 47: 243 (1997 [‘1995’]); Wang, Pan, Li,

24

Gard. Bull. Singapore 68(1) 2016

Weitzman & Skog, FI. China 18: 385 (1998); Burtt, Thai Forest Bull., Bot. 29: 83

(2001), pro parte; Smitinand, Thai PI. Names ed. 2, 14 (2001); Kress et ah, Checkl.

Myamiiar 261 (2003). - Trichosporum longicaule (Wall, ex R.Br.) Kuntze, Revis.

Gen. PI. 478 (1891). - TYPE: Burma, Chappedong, Wallich, N. 888 (lectotype K

[K000096741], designated by Middleton (2007); isolectotypes BM [BM000883859],

G-DC, K-W). (Fig. 8, 9)

Aeschynanthus griffithii R.Br., Cyrtandreae 115 (1839); Brown in Bennett, PI. Jav.

Rar. 115 (1840); A.DC., Prod. 9: 261 (1845); Miquel, FI. Ned. Ind. 2: 719 (1858);

C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 24 (1883); Clarke in Hooker, FI.

Brit. Ind. 4: 339 (1884); Kress et ah, Checkl. Myanmar 261 (2003). - Trichosporum

grijfithii (R.Br.) Kuntze, Revis. Gen. PI. 477 (1891). - TYPE: Burma, Tenasserim

Division, Tavoy, Griffith, W. s.n. (holotype BM [BM000883870]).

Aeschynanthus zebrinus Van Houtte, Hort. Vanhoutt. 1(2): 42 (1846). - TYPE: Not

located.

Aeschynanthus marmoratus TMoore, Paxton’s FI. Gard. 3: 56 (1852); C.B.Clarke in

A.DC. & C.DC., Monogr. Phan. 5(1): 38 (1883); Ridley, J. Linn. Soc. Bot. 32: 500

(1896); Ridley, J. Straits Branch Roy. Asiat. Soc. 44: 13 (1905); Ridley, J. Asiat. Soc.

Bengal 74 (2): 732 (1909); Ridley, FI. Mai. Pen. 2: 498 (1923); Barnett, FI. Siam. 3 (3):

202 (1962). - Trichosporum marmoratum (TMoore) TMoore ex Kuntze, Revis. Gen.

PI. 478 (1891). - TYPE: Cultivated specimen, October 1851, Moore, T s.n. (holotype

K[K000831883]).

Epiphyte with arching and pendulous branches; stems grey-brown or gi'ey, glabrous.

Leaves opposite; petiole 2-14 mm long, glabrous; blade fleshy, background mid to

dark green above but variegated with much paler venation, same but paler beneath,

elliptic, 1.9-12.2 x 0.3^ cm, 2.2-9 times as long as wide, apex caudate to acuminate,

base cuneate to acute, margin with few minute teeth but appearing more or less entire,

glabrous above and beneath, 5-6 pairs of secondary veins, weakly visible, tertiary

venation obscure. Inflorescences terminal, peduncle absent; pedicels 9.5-11 mm

long, green siifhised purple, glabrous. Calyx of separate lobes free to base, orange,

purple or green flushed maroon-purple, glabrous or with sparse glandular hairs; lobes

narrowly elliptic to linear, erect, 8-1 8 x 0. 8-2.4 mm, apex acute to acuminate. Corolla

20.5- 31 mm long, externally tube green, yellowish green or green with flushes of

orange, pui*ple or brown, lobes purple or maroon with an orange or yellow margin or

all orange, internally tube dark red, maroon or green, lobes maroon, purple or red with

a yellow or orange margin; upper lobes ovate or orbicular, not spreading or reflexed,

2. 5- 3. 8 X 2.4-4. 5 mm, sinus 2.2-4. 5 mm deep, apex rounded; lateral lobes orbicular

or oblique ovate, not spreading or reflexed, 2-3.9 x 3. 3-4. 8 mm, rounded; lower lobe

ovate or squarish, not spreading or reflexed, 2. 5-5. 8 x 3. 6^.2 mm, apex rounded,

outside glabrous except for ciliate lobes, to densely glandular puberulent, inside with

five dense tufts of multicellular hairs near base. Stamens long exserted; filaments

Aeschynanthus in Singapore and Peninsular Malaysia

25

Fig. 8. A. Aeschynanthus longicaulis Wall, ex R.Br. B. Aeschynanthus longiflorus (Blume)

A.DC. C. Aeschynanthus obconicus C.B. Clarke. (Photos: A, Preecha Karaket; B, Jana Leong-

Skornickova; C, David Middleton)

green or yellow, with glandular hairs; anthers pale brown or grey; anterior filaments

inserted at 12-19.5 mm from corolla base which is 48-52% of corolla length, anthers

2. 6-3.2 X 1-1.2 mm; posterior filaments inserted at 12.5-21 mm from corolla base

26

Gard. Bull. Singapore 68(1) 2016

Fig. 9. Distribution of Aeschynanthus longicaulis Wall, ex R.Br. in Peninsular Malaysia (•).

which is 50-55% of corolla length, filaments 23-27.5 mm long, anthers 1.8-2. 8 x

0.7-1. 3 mm; staminode 0.9 mm long; pollen greenish grey. Disk 0.9-1 .3 mm high,

a simple annular ring. Pistil 14-A5 mm long; stipe 1.5^ mm long, glabrous; ovary

5.5-12 mm long, glabrous or with sessile glands; style yellow or green flushed purple

towards apex, 7-3 1 mm long, glandular pubescent; stigma purple or yellow. Capsule

5.4-37 cm long, 1. 8-3.2 mm wide. Seed grain 1. 6-2.2 x 0. 1-0.6 mm, warty, bubble

cells absent; apical appendage a filifomi hair, c. 22.5 mm long; hilar appendages of

many (13-25) filifonn hairs, 10-25 mm long; appendages papillose.

Distribution. Eastern Myanmar, western and southern Thailand, Peninsular Malaysia

(Kedah, Kelantan, Penang, Perak, Perlis).

Habitat and ecology^. In primary and secondary lowland forest, reaching to upper

dipterocarp forest, sometimes on limestone, at 360-1150 m in Peninsular Malaysia.

Aeschynanthus in Singapore and Peninsular Malaysia

27

Most Malaysian material does not have good habitat and altitude notes but in Thailand

it has also been collected in rubber plantations.

Provisional lUCN conservation assessment. Least Concern (LC). This species has a

wide distribution and has been recorded from a variety of habitats. However, it should

be noted that it has been collected rather infrequently in Peninsular Malaysia in recent

years and only once since 2000 so its status should be monitored.

Peninsular Malaysian specimens examined. PENINSULAR MALAYSIA: Kedah: Langkawi,

Gunung Raya, 880 m, 13 Sep 1988, Tay, E.P 125 (SING); Weng Road near Baling, 2 Nov

1929, Best, G.A. 21255 (SING); Kelantan: Jeli, Upper Sungai Pergau, 25 Sep 1986, Latiff,

A. et al ALM1813 (PSU, UKMB); Penang: Government Hill, 760 m, Sep 1889, Curtis, C.

2142 (SING); ibidem, Mar 1889, Ridley, H.N. 1700 (BM, SING); Penang Hill, 760 m, 8

Sep 1982, Kochummen, K.M. FR129332 (K, KEP, L); Perak: Gunong [Gunung] Chabang,

Scortechini, B. 35 (SING); Ulu Temengor, Jul 1909, Ridley, H.N. 14280 (BM, SING); Grilc, 19

Jun 1924, Burkill, H.M. & Md Haniff 12542 (SING); Taiping, Bukit Earut, 1150 m, 12 May

2009, Yao, T.L. FRT57961 (KEP); ibidem, Heng, HP. et al. HPIO (SINU); Perils: Kaki Bukit,

1962, Cultivated C4877 (E); Bukit Bintang Forest Reserve, 360 m, 25 Apr 1962, Burtt, B.L. &

Woods, P.J.B. B1739{E).

Notes. This species is difficult to distinguish from Aeschynanthus fecundus when not

in flower. The name has also been applied widely to species from other parts of Asia

that are now recognised as distinct taxa such as Aeschynanthus memhranifolius and

A. sinolongicalyx W.T.Wang. These species all share characteristic variegated leaves.

Flowering material is needed to assess whether Aeschynanthus longicaulis occurs

further north i nto northern Thailand than previously suspected or whether other related

species rather have the wider distribution.

The name Aeschynanthus marmoratus, originally described from cultivated

material, was long applied to the material from Peninsular Malaysia but the type

material is indistinguishable from that of Aeschynanthus longicaulis.

7. Aeschynanthus longifloriis (Blume) A.DC., Prod. 9: 262 (1845); Hooker, Bot.

Mag. 73: t.4328 (1847); Zollinger, Syst. Verz. 3: 57 (1855); Miquel, FI. Ned. Ind. 2:

717 (1858); C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 32 (1883); Ridley,

J. Linn. Soc. Bot. 32: 499 (1896); Ridley, J. Straits Branch Roy. Asiat. Soc. 44: 14

(1905); Ridley, J. Asiat. Soc. Bengal 74 (2): 734 (1909); Ridley, FI. Mai. Pen. 2: 498

(1923); Backer & Bakhuizen van den Brink, FI. Java 2: 524 (1965); Turner, Gard.

Bull. Singapore 47: 243 (1997 [‘1995’]); Burtt, Thai Forest Bull., Bot. 29: 83 (2001);

Smitinand, Thai PI. Names ed. 2, 15 (2001); Middleton, Edinburgh J. Bot. 64: 400

(2007). -Lysionotus longifloms Blume, Bijdr. FI. Ned. Ind. 766 ( 1 826). - Trichosporum

longiflorum (Blume) Kuntze, Revis. Gen. PI. 478 (1891); Merrill, Contr. Arnold Arbor.

8:151 (1934). - TYPE: Indonesia, Java, Blume, C.L. 247 (lectotype L [Leiden number

903,307-102], designated by Middleton (2007); isolectotypes L [2 sheets]). (Fig 8, 10)

28

Gard. Bull. Singapore 68(1) 2016

Aeschynanthus perakensis Ridl., J. Linn. Soc. 32: 499 (1896); Ridley, J. Straits Branch

Roy. Asiat. Soc. 44: 15 (1905); Ridley, J. Asiat. Soc. Bengal 74 (2): 734 (1909);

Ridley, FI. Mai. Pen. 2: 498 (1923); Henderson, Malay. Wild. FI. Dicot. 340 (1959);

Turner, Gard. Bull. Singapore 47: 243 (1997 [‘1995’]). - TYPE: Peninsular Malaysia,

Perak, Taiping, Bukit Larut, 1891, Ridley, H.N. s.n. (lectotype SING [SING0035528],

designated by Middleton (2007)).

Epiphyte with arching stems; stems glabrous. Leaves opposite; petiole 3-16 mm long,

glabrous; blade slightly fleshy or coriaceous, dark to mid green above, pale green

beneath, ovate or elliptic, not variegated, 3.7-17.5 x 1.6-6. 7 cm, 1. 5-6.1 times as

long as wide, apex acuminate to caudate, base rounded to cuneate, glabrous above and

beneath, margin entire, secondary veins obscure to weakly visible, 3-8 pairs, tertiary

venation obscure. Inflorescences terminal, 2-5-flowered, peduncle absent, bracts

linear, 3-6 mm long; pedicels 1 1-16 mm long, brownish green or purplish, glabrous.

Calyx of separate lobes free to base, often differing quite substantially in length even

within a single flower, green to dark red or purplish, with sparse glandular hairs or

glabrous, sometimes with ciliate lobes or with just a few hairs only on very tips, 4.8-

23 mm long; lobes linear, narrowly triangular, narrowly ovate or oblong, erect, 4.8-23

X 1-2.2 mm, apex obtuse to acuminate. Corolla 43-90 mm long, tube nanow at base;

externally tube bright red or dark red, lobes bright red or dark red, internally tube

cream-coloured, lobes bright red on upper 2 lobes and bright red at margin with a

darker W-shaped line on lower 3 lobes; upper lobes orbicular, oblong or ovate, not

spreading or reflexed, 3.2-8. 1 x 3.2-5 mm, sinus 3-5.8 imn deep, apex rounded;

lateral lobes oblique ovate or deltoid, not spreading or reflexed, 3.2-7. 5 x 5.5-9 mm,

apex rounded; lower lobe ovate, not spreading or reflexed, 6-12 x 4-6.2 mm, apex

rounded; outside glabrous to sparsely glandular puberulent, usually with ciliate lobes,

inside with sparse glandular hairs throughout, sometimes very sparsely so, and sessile

glands below lobes. Stamens long exserted, fused in 2 pairs; filaments reddish, purple

or pink, with small glandular hairs, anthers pale pink; anterior filaments inserted at

49-60 mm from corolla base which is 72-77% of corolla length, filaments 29-50 mm

long, anthers 3.3-5 x 1-2 mm; posterior filaments inserted at 50-64 rmii from corolla

base which is 77-81% of corolla length, filaments 23-37 mm long, anthers 2.4-3. 8 x

0.9-2 mm; staniinode 0.7-5 mm long. Disk 1-1 .5 mm high, 5-crenate. Pistil 45-83

nun long; stipe 13-20 mm long, glabrous to sparsely minutely papillose; ovary 19-29

mm long, minutely papillose puberulent or with few sessile glands; style puiple or

purplish pink, 13-34 mm long, glandular pubescent to papillose, sometimes sparsely

so. Capsule 21-57 cm long, 3.5-4 mm wide. Seed grain 1-1.8 x 0. 3-0.4 mm, warty,

bubble cells absent; apical appendage a filifonn hair, 1 5-24 mm long; hilar appendage

a single filiform hair, 1 5-24 nun long; appendages papillose.

Distribution. Peninsular Thailand, Peninsular Malaysia (Kelantan, Negeri Sembilan,

Pahang, Penang, Perak, Selangor, Terengganu), Sumatra, Java, Borneo.

Aeschynanthus in Singapore and Peninsular Malaysia

29

Fig. 10. Distribution of Aeschynanthus longiflorus (Blume) A.DC. in Peninsular Malaysia (•).

Habitat and ecology. This species is recorded most commonly in lower montane

forest up to 1535 m altitude and occasionally from lowland or hill dipterocarp forest.

However, in Peninsular Malaysia it is also recorded from as low as 90 m altitude.

Provisional lUCN conservation assessment. Least Concern (LC). This species is

widely distributed in the region.

Additional Penitisular Malaysian specimens examined. PENINSULAR MALAYSIA:

Kelantan: Jeli, Upper Sungai Pergau, 25 Sep 1986, Latiff, A. etal ALM1789 (PSU, UKMB);

Kuala Krai, Stong Tengah FR, Sungai Kenerong, 564 m, 4 Apr 2009, Rosdi, M., Phoon, S.N.,

Ong, PT & Imin, K. FR166272 (KEP); Negeri Senibilan: Jelebu, Bkt Tangga, G. Telapak

Burok, 9 Apr 2008, Phoon, S.N. & Imin, K. FRI60648 (KEP); Pahang: Tahan River, 1520 m,

1891, Ridley H.N. 2167 (K, SING); Fraser’s Hill, Nov 1976, Keng, H. et al. 4755 (SINU);

ibidem, 1210 m, 18 Apr 1955, Purseglove, J.W. P4164 (E, K, L, SING); ibidem, 1210-1320

m, 16-30 Sep 1922, Burkill, KM. & Holttum, R.E. 7888 (K, SING); ibidem, 1500 m, 16 Apr

30

Gard. Bull. Singapore 68(1) 2016

1962, Burtt B.L & Woods, P.J.B. B1644 (E, SING); ibidem, 13 Mar 1986, Anthony, S. SA481

(KEP); ibidem, 1270 m, 4 Oct 1961, Burkill, H.M. HMB2812 (SING); ibidem, 1210 m, 20 Mar

1929, Holttum, R.E. 21528 (SING); ibidem, 25 Oct 1959, Mitchell s.n. (SING); ibidem, 27

Aug 1991, Kiew, R. RK3293 (SING); Fraser’s Hill, Rompin Trail, 1218 m, 26 Oct 2010, Julius,

A. FRI73605 (K, KEP, SING); Fraser’s Hill, Bishop’s Trail, 1275 m, 31 Mar 2007, Kiew, R.,

Lindsay, S. & Middleton, DJ. FR157497 (KEP); Fraser’s Hill, Raub, 1260 m, 21 Mai' 1994,

Perumal, B., Gan, C.L., Shahril, K.Z., Angan, A. & Bediil FRI41512 (KEP); Fraser’s Hill,

Tras Valley, 1090 m, 27 Sep 1922, Burkill, HM, 8859 (SING); ibidem, 1150 m, 25 Mar 1929,

Holttum, R.E. s.n. (SING); 3rd Mile Genting Highlands Road, 910 m, 2 Oct 1 970, Kochummen,

K.M. FR116215 (KEP); Cameron Highlands, 48th Mile Blue Valley, 23 Mar 1963, Mohd.

Kassim bin Rajah 488 (KLU); Cameron Highlands, Gunung Berembun. Nov 1908, Ridley,

H.N. 13603 (SING); Cameron Highlands, Nov 1940, Quaife s.n. (SING); Telom, Nov 1908,

Ridley, H.N. s.n. (SING); Cameron Highlands, G. Siku FR, 1430 m, 21 Jan 201 0, Mohd Hairul,

M.A., Ong, P.T., Siti Munirah, M.Y. cS: Kiieh, H.L. FRI69946 (KEP); Cameron Highlands, G.

Siku FR, Beside the road to G. Silcu FR, 1535 m, 25 Oct 2007, Imin, K. & Kueh, H.L. et al.

FR158561 (KEP); Cameron Highlands, Robinson’s Falls, 24 Mar 2007, Julius, A., Middleton,

D.J. & Lindsay, S. et al. FRI57479 (KEP); Penang: unknown s.n. (E); Perak: Gunung Korbu,

1210-1300 m, 11 Mar 1913, Robinson, H.C. s.n. (K); Taiping, 910-1210 m, Nov 1885, Kings

Collector 8314 (K); Taiping, Bukit Larut, 790 m, 18 Sep 1949, Sinclair, J. & Kiah SFN38800

(E, K, SING); ibidem, 1270 m, 25 Nov 1980, Keng, H. et al. 85 (SINU); ibidem, Dec 1902,

Ridley, H.N. 11447 (K, SING); ibidem, 1060-1210 m, Dec 1884, Kings Collector 7022 (K,

SING); ibidem, 18 Sep 1949, Sinclair, J. 6198 (E); ihidQm, Anderson, J.W 95 (SING); ibidem,

910 m, 29 Oct 1968, Smith, G. 443 (KLU); ibidem, 1290-1 330 m, 8 Mar 1939, Spare, G.H. 2114

(SING); Caulfield’s Hill, Wray, L. 625 (K, SING); Gunung Bintang, Bukit Kuala Bintang, 18

Apr 1928, Md Haniff 21088 (SING); Taiping, Gunung Hijau, Sep 1889, Curtis, C. s.n. (SING);

ibidem, 1300 m, 20 Oct 1992, Saw, L.G. FRI37667 (KEP); Selangor: Genting Highlands,

Gunung Bunga Buah, 1090 m, 5 May 1999, Chiia, L.S.L. FRI 40785 (KEP); Semangkok Pass,

1909, Ridley, H.N. s.n. (SING); Gombak, Gunung Bungah Buah, Trail to Gunung Bunga Buah,

1250 m, 31 Oct 2010, Yao, T.L., Kiew, R. & Wen, J. FR 165495 (E, KEP); Hulu Langat, Gn.

Nuang, Above Pacat Camp, 1418 m, 12 Dec 2013, Yao, T.L., Imin, K, Mohd Nazri, A. & Kueh,

H.L. FR177335 (KEP); Terengganu: Ulu Brang, Gunong [Gunung] Padang, Near Kampong

Lallang, 90 m, 16 Sep 1969, Whitmore, T.C. FRI12601 (K, KEP, L, SING).

Notes. This species is most similar to Aeschynanthus speciosus, particularly those

forms with redder flowers, but is easily distinguished by the opposite leaves of A.

longiflorus and the whorled leaves of A. speciosus.

The calyx pubescence character used by Ridley (1923) to distinguish

Aeschynanthus perakensis from A. longiflorus does not hold up to scrutiny and this

character is very variable in the species. As noted by Middleton (2007) the type of

Aeschynanthus perakensis is cited by Ridley as ‘Perak, on Gunong Hijan; Thaiping

Hills, at an altitude of 5,000 feet’. No Ridley material has been found which has this

locality information on the label and only one specimen has been found which could

possibly be original material. This specimen is labelled ‘Maxwell’s Hill’, now known

as Bukit Larut of which Gunung Hijau (the modem name for Gunong Hijan) is a part.

This specimen was lectotypified by Middleton (2007). A second Ridley specimen from

the area cannot be counted as original material as Ridley explicitly says he has seen no

fmiting material and the specimen has fmits.

Aeschynanthus in Singapore and Peninsular Malaysia

31

8 . Aeschynanthus obconicus C.B. Clarke in A.DC. & C.DC., Monogr. Phan. 5(1):

50 (1883); Clarke in Hooker, FI. Brit. Ind. 4: 343 (1884); Ridley, J. Linn. Soc.

Bot. 32: 501 (1896); Ridley, J. Asiat. Soc. Bengal 74 (2): 737 (1909); Ridley, FI.

Mai. Pen. 2: 500 (1923); Turner, Card. Bull. Singapore 47: 243 (1997 [‘1995’]). -

Trichosporum obconicum (C.B. Clarke) Kuntze, Revis. Gen. PI. 478 (1891). - TYPE:

Peninsular Malaysia, Selangor, Klang, 26 December 1878, Beccari, O. 36 (liolotype

FI [FIO 13085]). (Fig. 8, 11)

Epiphytic, stems mostly hanging, green, mostly glabrous, occasionally sparsely

pubemlent only around nodes. Leaves opposite; petiole 3-10 mm long, glabrous or

rarely sparsely puberulent; blade coriaceous, ovate or elliptic, 2.5-10.7 x 0.9-6 cm,

1. 5-3.1 times as long as wide, apex acuminate or acute, base cuneate to rounded, mid

green above, paler beneath, not variegated, glabrous above and beneath, occasionally

with few marginal hairs at margin base, margin entire, sometimes undulate, 4-7 pairs of

secondary veins, weakly visible or obscure, tertiary venation obscure. Inflorescences

axillai 7 or subterminal, 1-6-flowered; peduncles 4 mm long; bracts elliptic, 1.3-4 mm

long; pedicels 6-12 mm long, dark red, sparsely to densely eglandular puberulent.

Calyx with a tube and short and broad lobes or barely any lobes at all, the whole

broadly cup-shaped, dark red or dark red at base and becoming bright red higher,

sometimes purplish brown, outside densely to sparsely pubescent with red hairs, these

varying degrees of eglandular and glandular, very rarely glabrous, inside sparsely

pubescent to glabrous, total length 8-19 nmi long; tube 7-15 mm long which is 74-

94% of total length, 15-27 mm wide at top of tube; lobes semicircular or a weak

curve of the upper rim, erect, spreading or recurved, 0. 7-6.5 mm long, 9-16 mm wide,

apices rounded. Corolla 23-35 mm long, tube broad at base, externally bright to dark

red, internally cream, lobes externally bright red with cream at base of lower 3 lobes,

internally red with cream and dark red markings on lower 3 lobes; upper lobes ovate,

oblong or slightly falcate, spreading, 4.2-10 x 1 J-6 mm, sinus between lobes 3-5 mm

deep, apices rounded; lateral lobes ovate to oblong, spreading to reflexed, 4.2-12.5 x

4.2-9. 8 mm, apices rounded; lower lobe ovate or elliptic, spreading, 5-11.4 x 5. 6-6. 7

mm, apex rounded; outside densely pubescent or rarely glabrous except for ciliate

lobes, inside with scattered short glandular hairs with robust base, sessile glands at

lobe sinuses. Stamens not or only slightly exerted, fused in 2 pairs, filaments cream

to bright red, papillose, anthers cream or yellow; anterior filaments inserted at 9-15

mm from corolla base which is 32-58% of corolla length, filaments 15-1 8.5 mm long,

anthers 2.4-2. 7 x 1.2-1. 5 mm; posterior filaments inserted at 12-17 mm from corolla

base which is 43-63% of corolla length, filaments 12.8-16 mm long, anthers 1.6-2. 5

X 0.9-1 .6 mm; staminode c. 1 mm long. Disk 0.5-1 mm high, a simple aimular ring.

Pistil 18.5-36.5 mm long; stipe 6-13 mm long, with few sessile glands or glabrous;

ovary cream to pale green, 9.5-26 mm long, with sessile glands; style 2-7.5 mm long,

glandular pubescent, sometimes these sparse. Capsules 1 8-41 cm long, 2.8 mm wide.

Seed grain 0.8 mm x 0.3 mm, papillose, bubble cells present at base of hilar appendage;

apical appendage a filiform hair, 12.5 mm long; hilar appendage a single filiform hair,

10 mm long; appendages not papillose.

32

Gard. Bull. Singapore 68(1) 2016

Fig. 11. Distribution of Aeschynanthus obconiciis C.B. Clarke in Peninsular Malaysia (•).

Distribution. Peninsular Malaysia (Kedah, Kelantan, Kuala Lumpur, Pahang, Perak,

Selangor, Terengganu), Sumatra, Borneo.

Habitat and ecology. Epiphytic in lowland dipterocarp forest, often on trees by rivers,

occasionally in lower montane forest, at 0-1300 m altitude.

Provisional lUCN conservation assessment. Least Concern (LC). This species is

widespread and occurs in a variety of forest types.

Additional Peninsular Malaysian specimens examined. PENINSULAR MALAYSIA:

unlaiown loc., 21 Nov 1932, unknown collector FMS 17 068 (K, KEP); unknown state, Belau

Tajor, Wray> L. 1772 (SING); Kedah: Gunong [Gunung] Bintang, 8 Apr 1968, Sidekbin Kiah

8.277 (L, SING); Kelantan: Kpg. Biliai, Sg. Kerchit, 13 Feb 2003, Kiew, R. RK5239 (SING);

Kuala Krai, Stong Tengah FR, Sg. Renerong, 4 Apr 2009, Imin, K„ Phoon, S.N., Ong, P.T &

Rosdi, M. etal. FRI68188 (KEP); Kuala Lumpur: Feb 1890, Curtis, C. s.n. (SING); Pahang:

Aeschynanthus in Singapore and Peninsular Malaysia

33

Sungai Telom, 450 m, 24 Aug 1930, Kiah 23943 (NY, SING); Puku, Kuala Teku, 150 m, 21

Feb 1921, Seimund, E. 459 (SING); Kuala Terla Forest Reserve, Trail from Kg. Sg. Jarik, 540

m, 23 Mar 2007, Julius, A. FRI57451 (KEP, SING); Gunung Tahan, Teku, 21 Juii 1922, Md

Haniff & Md Nur 8046 (SING); Taman Negara, Sg. Tahan, Kem Teku, 283 m, 12 May 2008,

Lim, C.L. & Kiieh, H.L. FRI64886 (KEP); Perak: unlmown loc., Jan 1886, Kings Collector

10179 (SING); Gunong [Gunung] Inas, Sira Rimau, 27 Dec 1899, Yapp, R.H. 547 (CGE);

Earut Matang, Bkt. Larut, Sungai Bt. Tugoh, 72 m, 28 Jul 201 1, Imin, K. & Asmarayani, H,S.

FRI76004 (KEP); Taiping, Bukit Larut, 610 m, 1901, Curtis, C. s,n. (SING); Taiping, Dec

1902, Ridley, H.N. s.n. (SING); ibidem, 25 Aug 1971, Kockummen, KM. FRI16427 (KEP);

ibidem. 450 m, Sep 1889, Curtis, C. s.n. (SING); ibidem, Jul 1881, King’s Collector 2012

(SING); ibidem, 1000-1300 m, Dec 1882, Kings Collector 3641 (SING); ibidem, Jul 1881,

Kmg’s Collector 2049 (SING); ibidem, 9 Mar 1924, Burkill, H.M. & Md Hanijf 12798 (SING);

ibidem, 4 Mar 1965, Flardial, S. & Samsuri Ahmad 290 (SING); ibidem, Mai’ 1882, King’s

Collector 2849 (SING); ibidem, Mai’ 1892, Ridley, H.N. 2905 (BM, SING); ibidem, 76a-790

m, 16 Sep 1949, Sinclair, J. & Kiah SFN38805 (E, SING); ibidem, 610 m, 25 Aug 1971,

Whitmore, T.C. FR120392 (K, KEP, L, SING); Gunong [Gunung] Bubu, 600 m, 18 Aug 1966,

DingHou 657 (K, KEP, L, SING); ibidem, 610 m, 16 Aug 1966, Chew, W.-L. CWL1213 (K,

KEP, SING); Kampong Sekam, 5 10 m, 19 Oct 1982, Ave, W. 165 (K, L); Dinding, 1888, Curtis,

C. 1388 (K, SING); Kinta, Kinta Dam, Trail to G. Korbu, 450 m, 10 Jun 2010, Imin, K. et al.

FR171615 (K, KEP); Gunung Korbu, 610 m, 20 Mar 1913, Robinson, H.C. s.n. (K); Kuala

Dipang Forest Reserve, 30-210 m, 19 Feb 1976, Sidek bin Kiah SK515 (C, KEP, SING); Sg.

Siput, Chior Forest Resei've, Sungai Legap, 180 m, 9 Oct 1967, Ng, F.S.P. FRI5813 (KEP);

Chenderiang, Gunong [Gunung] Bujang Melaka, Sungai Rias, 610 m, 12 Feb 1975, Shah,

M., Ahmad & Mahmud MS3407 (C, KEP, SING); Bujang Melaka, Aug 1898, Curtis, C 3335

(SING); Batu Kurau, Curtis, C. 2990 (SING); Pondok Tanjong Forest Reserve, 14 Feb 1939,

Spare, G.H. SFAid22d (SING); Sungei Merbau, 10 Dec 1929, Symington, C.F. 27 JPtl (SING);

Bleinja [?Belanjar], Wray, L. 149 (SING); Scortechini, B. 46a (SING); Ijok Forest Reserve,

II Feb 1939, Aziz Budin 48644 (KEP [2 sheets]); Dinding, Curtis, C. s.n. (SING); Tapah,

Nov 1908, Ridley. H.N. 14063 (SING); Bujang Melaka, Sep 1898, Ridley, H.N. s.n. (SING);

Dinding, 1892, Curtis, C. s.n. (SING); Gunung Bintang, Bukit Kuala Bintang, 17 Apr 1928, Md

Haniff 21074 (SING); Bintang Hijau FR, Sg. Tebing Tinggi, 317 m, 26 Aug 2009, Chan, Y.M.,

Norzamli, A, & Norazmi, A. FRI70603 (KEP); Selangor: Petaling, 5 Jul 1 889, Ridley, H.N. s.n.

(SING); Kanching Reserve, 17 Sep 1925, Strugnell, E.J. 10515 (K); Kanching River, 250 m, 3

Oct 1974, Van Balgooy, M.M.J. 2118 (E, L, NY); Templer National Park, Bukit Perangin, 1110

m, Cultivated C7650 (E); Gombak, Bukit Lagong FR, Sg. Kepong, 392 m, 6 Sep 2007, Chan,

Y.M. FR149298 (KEP); Bukit Lagong Forest Reserve, 300 m, 24 Sep 1980, Kochummen, K.M.

FRI29126 (K, KEP, L, SING); Genting Highlands, Gombak F.R., 580 m, 23 Nov 1939, Wong,

Y.K KEP93272 (K); Ridge above Gombak Road, 4 Oct 1970, Stone, B.C.M. 9566 (KLU);

Sungei Buloh, Aug 1908, Ridley, H.N. 13372 (BM, SING); ibidem, 80 m, 4 Dec 1960, Poore,

M.E.D. 1064 (KLU); Kanching, Sg. Bangkap, 10 Oct 1974, Stone, B.C.M. & Badaruddin

12068 (BKF, KLU); Genting Sempah ridge, 800 m, 21 May 1982, Stone, B.C.M. 15167 (KLU);

ibidem, 760 m, 6 Mar 1973, Stone, B.C.M. 11091 (KLU); ibidem, 850 m, 18 Aug 1974, Stone,

B.C.M, s.n. (KLU); Terengganu: Ulu Sungei Setiu, 28 Apr 1986, Kiew, R. RK2262 (KEP);

Hulu Terengganu, Tembat Forest Reserve, 364 m, 1 Apr 2009, Mohd. Hairul, M.A. FR160925

(KEP, SING); Besut, Sungai Kemia, 210 m, 29 Aug 1996, Chua, L.S.L. FRI26700 (KEP).

Notes. This is one of the most readily identified species oY Aeschynanthus in Peninsular

Malaysia due to its wide cup-shaped red calyx. It is most similar to Aeschynanthus

34

Gard. Bull. Singapore 68(1) 2016

tricolor Hook, from Borneo but that species mostly has a shorter corolla and has dark

lines extending down the corolla tube (absent in A. obconicus). It is also similar to

Aeschynanthus wallichii but that species has a smaller, more open and green calyx.

9. Aeschynanthus pulcher (Blume) G.Don, Gen. Syst. 4: 656 (1838); A.DC., Prod. 9:

262 (1845); Zollinger, Syst. Verz. 3: 56 (1855); Miquel, FI. Ned. Ind. 2: 721 (1858);

C.B. Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 43 (1 883); Middleton, Edinburgh

J. Bot. 64: 412 (2007); Middleton, Edinburgh J. Bot. 66: 437 (2009). - Trichosporum

pulchrum Blume, Bijdr. FI. Ned. Ind. 764 (1826). - TYPE: Indonesia, Java, Blume,

C.L. s.n. (lectotype L [Leiden number 903,307-167], designated by Middleton (2007)).

(Fig. 12, 13)

Aeschynanthus pai^ifolius R.Br., Cyrtandreae 115 (1839); Brown in Bennett, PL Jav.

Rar. 115 (1840); A.DC., Prod. 9: 262 (1845); Miquel, FI. Ned. Ind. 2: 720 (1858);

C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 42 (1883); Ridley, FI. Mai. Pen. 2:

500 (1 923); Henderson, Malay. Wild. FI. Dicot. 342 (1959); Chin, Gard. Bull. Singapore

32: 147 (1979); Stone, Fed. Mus. J. 26 (1): 98 (1981); Turner, Gard. Bull. Singapore

45: 92 (1993); Turner, Gard. Bull. Singapore 47: 243 (1997 [‘1995’]); Smitinand, Thai

PI. Names ed. 2, 15 (2001). - Trichosporum panifolium (R.Br.) Kuntze, Revis. Gen.

PI. 478 (1891). - TYPE: Indonesia, Sumatra, Pulau Bangka, 1813, Horsfield, T s.n.

(lectotype BM [BM000847144], designated by Middleton (2007)).

Aeschynanthus boschianus de Vriese, Ami. Soc. Agric. Bot. Gand 1: 403 (1845); Van

Houtte, Hort. Vanhoutt. 1 (2): 31 (1846). - Trichosporum boschianum (de Vriese)

Kuntze, Revis. Gen. PI. All (1891); Merrill, Contr. Arnold Arbor. 8: 151 (1934). -

TYPE: Cultivated plant grown by Jacob-Makoy in Belgium from plant collected on

Mt Gede in .lava (untraced).

Aeschynanthus lobbianus Hook., Bot. Mag. 72: t.4260 (1846); Miquel, FI. Ned. Ind. 2:

721 (1858); C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 44 (1883); Clarke in

Hooker, FI. Brit. Ind. 4: 343 (1884); Ridley, J. Linn. Soc. Bot. 32: 501 (1896); Ridley,

J. Straits Branch Roy. Asiat. Soc. 44: 16 (1905); Ridley, J. Asiat. Soc. Bengal 74 (2):

735 (1909). - Trichosporum lobbianum (Hook.) Kuntze, Revis. Gen. PI. 478 (1891). -

TYPE: Indonesia, Java, Lobb, T. s.n. (holotype K).

Aeschynanthus javanicus Rollinson ex Hook., Bot. Mag. 76: t.4503 (1850); Miquel,

FI. Ned. Ind. 2: 721 (1858); C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 44

(1883). - Trichosporum javanicum (Hook.) Kuntze, Revis. Gen. PI. All (1891). -

TYPE: Indonesia, Java, unknown s.n. (holotype K).

Aeschynanthus neesii Zoll. & Moritzi in H.Zollinger, Syst. Verz. 3: 56 (1855), nom.

nud. - Based on Zollinger, H. 1546 (P).

Aeschynanthus in Singapore and Peninsular Malaysia

35

Fig. 12. A-D. Aeschynanthus pulcher (Blume) G.Don showing the variation in flower

morphology and eolour, particularly in the calyx. E. Aeschynanthus radicans Jack. F.

Aeschynanthus wallichii R.Br. (Photos: A, David Middleton; B-D, Jana Leong-Skomickova;

E, Preecha Karaket; F, Ali Ibrahim)

36

Gard. Bull. Singapore 68(1) 2016

Fig. 13. Distribution of Aeschynanthus pulcher (Blume) G.Don in Singapore and Peninsular

Malaysia (•).

Aeschynanthus lampongus Miq., FI. Ned. Ind., Eerste Bijv. 563 (1861); C.B.Clarke in

A.DC. & C.DC., Monogr. Phan, 5(1): 42 (1883). - Trichosporum lampongum (Miq.)

Burkill, Kew Bull. 1935: 319 (1935). - TYPE: Indonesia, Sumatra, Lampung, Radja-

bassa, 910 m, Teijsmann, J.E. HB4482 (lectotype U, designated by Middleton (2009);

isolectotype L).

Aeschynanthus beccarii C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 47 (1883).

- Trichosporum beccarii (C.B.Clarke) Kuntze, Revis. Gen. PI. All (1891). - TYPE:

Indonesia, Sumatra, Sumatera Barat, Padang, Beccari, O. 796 (lectotype K, designated

by Middleton (2009); isolectotypes BM [BM000537111], FI n.v., K, MEL).

Aeschynanthus zollingeri C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 44

(1883). - Trichosporum zollingeri (C.B.Clarke) Kuntze, Revis. Gen. PI. 478 (1891). -

Aeschynanthus in Singapore and Peninsular Malaysia

37

TYPE: Indonesia, Java, Zollinger, H. 1512 (lectotype BM [BMOOl 125703], designated

by Middleton (2009); isolectotypes B [presumably destroyed], P [P00492368]).

Aeschynanthus lanceolatus RidL, J. Bot. 62: 299 (1924); Ridley, FI. Malay Penin.

5: 324 (1925); Turner, Card. Bull. Singapore 47: 243 (1997 [‘1995’]). - TYPE:

Peninsular Malaysia, Pahang, Fraser’s Hill, 1210-1320 m, 16 October 1922, Burkill,

H.M. & Holttum, R.E. 8418 (holot 3 ^e K).

Aeschynanthus lampongus war.paj^ifolius RidL, FI. Malay Penin. 5 : 324 ( 1 925); Turner,

Card. Bull. Singapore 47: 243 (1997 [‘1995’]). - TYPE: Peninsular Malaysia, Pahang,

Puku, Kuala Teku, 20 December 1920, Seimund, E. s.n. (lectotype K, designated by

Middleton (2007); isolectotype SING).

Lithophytic or epiphytic, hanging or creeping, stems dark purple to green, sparsely

puberulent to glabrous. Leaves opposite; petiole 1-3(6) nmi long, sparsely puberulent

or glabrous; blade coriaceous or slightly fleshy, elliptic or ovate, dark green to purplish

above and beneath, not variegated, 0.9-5. 9 x 0.25-2.9 cm, 1.3-5. 6 times as long as

wide, apex rounded to acuminate, base subcordate to cuneate, glabrous above, glabrous

to very sparsely puberulent all over beneath, margin entire, secondai^ veins weakly

visible or obscure, c. 3 pairs, tertiary venation obscure. Inflorescences subterminal

or axillary, 1-2 -flowered; peduncle 0-12.5 mm long; bracts elliptic or ovate, 3-6.5 x

2-5.5 mm; pedicels 6-13 mm long, green, sparsely puberulent to densely long hairy.

Calyx with a tube for most of length and free lobes, dark red to purplish or purplish

brown, mostly sparsely eglandular puberulent, more rarely densely puberulent,

glabrous or with a few hairs only on veiy tips, 1 1 .5-29 mm long; tube 10-25 mm long,

89-96% of total length, 6-2 1 mm wide at top of tube; lobes triangular, semicircular

or merely a weak curve of the upper rim, slightly spreading or erect, 1-5 x 4-7 mm,

apex rounded or obtuse. Corolla 42-66 mm long, inflated at base, externally tube

dark or bright red, often white at extreme base inside calyx tube, lobes bright red,

internally lobes bright red with cream and dark markings on lower 3 lobes; upper

lobes oblong or ovate, spreading or not, 5.7-13 x 1. 9-6.5 mm, sinus 2.2^. 6 mm deep,

apex rounded; lateral lobes deltoid or ovate, spreading or not, 6.1-11,5 x 6.5-10.5

mm, apex rounded; lower lobe elliptic or ovate, spreading or not, 7.6-13 x 6.5-8 mm,

apex rounded; outside sparsely to densely puberulent, inside glabrous or with scattered

papillae or small glandular hairs which are denser in the upper half, sessile glands

below lobes. Stamens reaching end of corolla or slightly exserted, fused in 2 pairs;

filaments red, glabrous or with very few sessile glands or papillose; anterior filaments

inserted at 29^7 mm from corolla base which is 54-68% of corolla length, filaments

21-27 mm long, anthers 2.4-3. 3 x 0.8-1. 6 mm; posterior filaments inserted at 33.5-

41 mm from corolla base which is 60-70% of corolla length, filaments 17.5-21 mm

long, anthers 2. 3-2. 9 x 0.8-1. 9 mm; staminode 0.5-1. 6 mm long. Disk 0.8-1. 5 mm

high, strongly 5-lobed, 5-crenate or a simple annular ring. Pistil 37-58 mm long; stipe

16-27 mm long, puberulent; ovary pale yellow, 14-30 imn long, minutely papillose

or with sessile glands; style pale yellow, 4.5-14 mm long, densely pubescent. Capsule

38

Gard. Bull. Singapore 68(1) 2016

18-40 cm long, 2-A mm wide. Seed grain 0. 6-0.9 x 0.2-0.3 mm, papillose, bubble

cells present at base of hilar appendage; apical appendage a filiform hair, 7-9.5 mm

long; hilar appendage a single filiform hair, 6-9 mm long; appendages not papillose.

Distribution. Extreme south of Thailand, southern Vietnam, Peninsular Malaysia

(Johor, Kedah, Kelantan, Melaka, Negeri Sembilan, Pahang, Penang, Perak, Perils,

Selangor, Terengganu), Singapore, Sumatra, Java, ?Borneo. See also note below.

Habitat and ecology. Epiphytic or clambering over rocks, most common in lower

montane forest, sometimes in lowland, hill dipterocarp or upper montane forest, in

peat swamp forest or in forest on quartzite or sandstone, at 0-2100 m altitude.

Provisional lUCN consen’ation assessment. Least Concern (LC). This species is

widespread and locally common. In Singapore it was long thought to be nationally

extinct but has been rediscovered in Nee Soon Swamp (Williams, 2014).

Additional Singaporean and Peninsular Malaysian specimens examined. PENINSULAR

MALAYSIA: unknown loc., Jan 1882, Kunstler, H. 2636 (NY); Johor: unknown loc., May

1924, Franck, C.W. 209 (C); ibidem, 6 Jun 1965, Jumali K2077 (SINU); Pontian, Pengkalan

Raja, 25 Jun 1939, Corner, E.J.H. & Henderson, M.R. SFN36611 (K, P), SFN36612 (SING);

western tip of Gunong [Gunung] Panti West, 560 m, 1 8 Jul 1981, Maxwell, J.F 81-166 (AAU,

KLU, SING); Gunong [Gunung] Panti, 1 Aug 1960, unknown s.n. (SINU); ibidem, 480 m, 4

Feb 1981, Collenette, l.S. 2253 (E); ibidem, 520 m, 5 Jul 1970, Samsuri Ahmad S.311 (BKF, C,

G, K, KLU, L [2 sheets], LAE, SING); ibidem, 460 m, 31 Jul 2008, Yao, T.L., Lim, C.L., Rosdi,

M. &Ayau, K. FR165 387 (KEP); Kuala Sedili Road, 26 May 1961, Biirkill, H.M. HMB2665 (K,

SING); Kota Tinggi, Kuala Sedili New Road, 23 Jun 1959, Kadim & Noon M. 135 (E, K, L,

LAE, SING); Mawai-Sedili Road, 8 Feb 1961, CheM>, W.-L, CWL224 (C, E, K, L, SING); 8th

mile Mawai-Kuala Sedili New Road, 8 Feb 1961, Sinclair, J. 10568 (E, L, SING); Mersing,

Lombong Batu, 6 Jun 1965, Keng, H. & Jumali K2077 (L); Kulai FR, Gunung Pulai, 29 Jun

1987, Zainudin, A. et al. AZ2438 (UKMB); Penggaram, Nov 1900, Ridley, H.N. s.n. (SING);

Batu Pahat, 1892, Nongchi s.n. (SING); Tanjong Kupang, 1894, Ridley, H.N. s.n. (SING);

Tanah Runto, 14 Feb 1890, Goodenough, J.S. s.n. (SING); ibidem, 22 Apr 1890, Ridley, H.N.

s.n. (SING); Batu Pahat, Nov 1900, Ridley, H.N. s.n. (SING); Tempayang River, Apr 1909,

Ridley, H.N s.n, (SING); Kulai Hutan Simpan, Gunung Pulai, 24 Apr 1922, Md Nur & Kiah

7783 (SfNG); Jambu Larang, Oct 1892, Fielding, J. s.n. (SING); Sungei Tukong, 0 m, 20 Dec

193 1, Spare, G.H. 991 (SING); Segamat, Gunung Chabang Tiga, Gunung Chabang Tiga South

Peak, 935 m, 25 Aug 2007, Chew, M.Y & Rosdi, M. FR155583 (KEP); Gunong [Gunung]

Bekok, 18 Aug 1968, Jumali & Heaslett, E.A. KJ732 (SINU); Gunung Ledang, 7 Dec 2010,

Yeo, C.K. s.n. (SINU); ibidem, 24 Feb 1980, Zainal Mustafa ZM64 (UKMB); ibidem, 26 Apr

1972, Year III students FSC312 (KLU); Gunung Ledang, Above Bukit Besar, 12 Jun 1892,

Ridley, H.N. s.n. (SING); Kedah; unknown loc,, 24 Apr 1868, Maingay, A.C. K.D.1218 (K);

Langkawi, Gunong [Gunung] Raya, Sep 1890, Curtis, C. 2503 (SING); Kedah Peak, 1060 m,

Dec 1915, Robinson, H.C, & Kloss, C.B. 5997 (K, SING); ibidem, 910 m, 2 Dec 1915,

Robinson, H.C. & Kloss, C.B. 6049 (K, SING); ibidem, 1000-1400 m, 21 Jan 1983, Davis

69431 (E); ibidem, 1893, Ridley, H.N. s.n. (SING); ibidem, 910-1210 m, 3 Jun 1971, Keng, H.

etal. 67 (SING, SINU); ibidem, Aug 1893, Ridley, H.N. 5513 (BM, SING); Gunung Jerai, 850

m, 12 Feb 1961, Yong KEP99330 (K, KEP); ibidem, 1200 m, 14 Sep 1979, Rao & et al. 167

Aeschynanthus in Singapore and Peninsular Malaysia

39

(L); ibidem, 8 Nov 1962, Samsuri Ahmad 309 (SING); ibidem, 1121 m, 9 Feb 2010, Imin, K.,

Nor Ezzawanis, A.T., Hovenkamp, P. & Angan, A. FRI66493 (KEP); ibidem, 3 Sep 1995, Choo,

J.P.S., Chua, D.W.S., Lian, S.H.L., Lim, PY. & Wong, J.Y. PYL2 (SINU); ibidem, 1200 m, 14

Sep 1979, Rao & et ai 167 (SINU); ibidem, 850 m, 15 May 1969, Smith, G. 513 (KLU);

ibidem, 1060 m, 10 Jul 1977, Lo, YN. & Mahmud 83 (KLU); ibidem, 1150 m, 13 May 1969,

Stone, B.C.M. & Mahmud 8500 (KLU); Kelantan: S. Lebir, Sungai Terang, 8 Jul 1935,

Henderson, M.R. SFN29643 (K, SING); Gua Musang, 3 Aug 1962, UNESCO limestone

expedition 239 (SING); Gua Musang*, Gua Batu Boh, 300 m, 12 Aug 1971, Chin, S.C, 1441 (L);

Kuala Krai, Gunung Stong Utara Forest Reserve, VJR Gunung Stong Utara, 180 m, 19 Jun

2001, Sam, Y-Y FR146602 (KEP); Dabong, Gunung Setong, 15 May 1988, Kie\\>, R. RK2727

(SING); Melaka: unknown loc., 7 Oct 1865, Maingay, A.C. K.D.1219 (K); Ayer Panas, Nov

1893, Ridley, H.N 1578 (BM, SING); Jus, Oct 1893, Goodenough, J.S 1711 (SING); Jasin,

Bkt. Senggeh FR, Batu Lebah, Kg. Bkt Senggeh, 175 m, 28 Aug 2007, Chan, K,Y FR149293

(KEP); Negeri Sembilan: Serting Ulu Forest Reserve, 4 Jun 1992, Saw, LG. & Mustafa, D.

FRI37514 (K, KEP); Jelebu, Gunung Telapa Buruk, 1000 m, 21 Apr 1988, Kiew, R. RK2641

(KEP); Rembau, Gunung Datuk, 400 m, 10 Jul 1979, Asmah 22 (UKMB); Tampin, Tampin

Forest Reserve, Hutan Lipur G. Tampin, 351 m, 5 May 2009, Chan, K.Y FR164737 (KEP,

SING); Jelebu, Bkt Tangga, G. Telapak Burok, 1038 m, 1 1 Apr 2008, Rosdi, M. & Rqfidah, A.R.

FR159837 (KEP); Pahang: Genting Highlands, 1150 m, 11 Jun 1978, Stone, B.C.M. 13764

(KLU); ibidem, 25 Nov 1984, Kiew, R. RK1548 (KEP); ibidem, 1450 m, 14 Mar 1982, Stone,

B.C.M. 15134 (KLU); ibidem, 15 Nov 1966, Stone, B.C.M. 6543 (G, KLU, L); ibidem, 910 m,

9 May 1972, Kochummen, K.M. FRI16516 (KEP); Genting Highlands, Gunung Ulu Kali, 1600

m, 9 Apr 1978, Ma.xweU, J.F 78-90 (L); Genting Sempah ridge, Nov 1970, Stone, B.C.M. s.n.

(KLU); Kuantan, Bukit Berserah Forest Reserve, 3 Sep 1933, Mahmood KEP 172 16 (K, KEP,

L); Kwantan [Kuantan], Sep 1889, Dtirnford, L. s.n. (SING); Pekan, Jul 1917, Evans, J.H.N.

s.n. (K); ibidem, 1 Apr 1890, Haviland, G.D. s.n. (SING); ibidem, 30 Nov 1924, Burkill, H.M.

& Md Haniff 17258 (SING); Bukit Cheras, 150 m, 11 Oct 1931, Henderson, M.R. 25065

(SING); Rompin, 19 May 1919, Gerb 3257 (SING); ibidem, 15 Nov 1929, Mahammud 17178

(SING); Gunung Mengkuang, 29 Mar 1959, unknown KEP78844 (KEP); Temerluh, Krau GR,

Trail to Batu Bergambar From Batu Gajali, 550 m, 4 Jun 2000, Cliua, L.S.L., Damahuri, S.,

Ayau, K. & Ramli, P. FR145633 (KEP); Gunung Senyum, 29 Jul 1929, Henderson, M.R. s.n.

(SING); Fraser’s Hill, 1210 m, 18 Apr 1955, Purseglove, J.W P4182 (K, L, LAE, SING);

ibidem, 4 Oct 1961, Burkill H.M. HMB2811 (K, L, LAE, SING); ibidem, 1300 m, 19 Apr

1955, Purseglove, J.W. P4202 (E, K, L, SING); ibidem, 1210 m, 24 Sep 1959, Shah, M. &

Noon M. MS607 (E, K, SING); ibidem, 1 6 Jul 2002, Bramley, G. & Sam, Y.-Y. GB27 {E, KEP);

ibidem, 450 m, 16 Apr 1962, Burtt, B.L. & Woods, P.J.B. B1643 (E); ibidem, 22 Dec 1979,

Bremer, B. & Bremer, K. 1808 (KLU, L); ibidem, 1240-1270 m, 3 Oct 1987, Worthington, R.D.

13326 (L); ibidem, 16 Dec 1979, Kiew, R. RK832 (KEP); ibidem, 21 Nov 1980, Kiew, R.

RKI002 (KEP); ibidem, 13 Feb 1986, KieM’ R. RK2160 (KEP); ibidem, 13 Mar 1986, Anthony,

S. SA482 (KEP); ibidem, 10 Nov 1987, Anthony, S. SA897 (KEP); ibidem, 1030 m, Kassim, M.

550 (UKMB); ibidem, 25 Oct 1979, Zainudin, A. AZ40 (UKMB); ibidem, 31 Oct 1991, Kiew,

R. RK3356 (SING); ibidem, 1210 m, 13 Jul 1957, Hawkins, A.S.M. s.n. (SING); ibidem, 1210

m, 25 Aug 1923, Henderson, M.R. FMS11263 (SING); ibidem, 14 Jun 1930, Kalong 22423

(SING); ibidem, 13 Nov 1981, Keng, H. et al. 86 (SING); ibidem, 21 Jun 2006, Phoon, S.N,

Kamarudin S., Ktieh, H.L. & Rafidah, A.R. FR15I576 (KEP); ibidem, Mendum, M. s.n. (E);

ibidem, 26 Aug 1991, Kiew, R. RK3264 (SING); ibidem, 1240 m, 25 Aug 1959, Burkill H.M.

HMB1996 (SING); ibidem, 1210-1320 m, 16-30 Sep 1922, Burkill, H.M. & Holttum, R.E. s.n.

(SING); ibidem, 21 Nov 1977, Keng, H. et al. 97 (SINU); ibidem, 5 Nov 1973, Keng, H. et al.

40

Gard. Bull. Singapore 68(1) 2016

54 (SINU); ibidem, 28 Oct 1974, Keng, H. etal. 93 (SEMU); ibidem, 13 Nov 1981, Keng, H. et

al. CTV86 (SINU); ibidem, Dec 1970, Mahmud bin Sidek s.n. (KLU); ibidem, Sep 1940,

Addison, G.H. s.n. (SING); ibidem, 1 120 m, 29 Aug 1923, MdNur 11138 (SING); ibidem, 1210

m, 29 Aug 1923, Henderson, M.R. FMS11453 (SING); Fraser’s Hill, Jeriau Road, 1060-1210

m, 20 Aug 1960, Burkill, H.M., Shah, M. & Noor, M. HMB2430 (SING); ibidem, 1090 m, 17

Jun 1972, Stone, B.C.M. 10798 (KLU); Fraser’s Hill, Mager Trail, 1 390 m, 3 May 2007, Chew,

M.Y, Imin, K., Kiew, R. & Nooteboom, H.P. FR153648 (E, KEP, SING); Fraser’s Hill, Jalan

High Pines, 1200 m, 13 Nov 2006, Phoon, S.N. FRI51987 (KEP, KLU, SING); Fraser’s Hill,

Pine tree hill, 1450 m, 19 Sep 1922, Burkill, H.M. & Holttum, R.E. 8549 (SING); Fraser’s Hill,

Richmond, 27 Oct 2010, Imin, K., Utteridge, T.M.A. & Julius, A. FRI71855 (KEP); Fraser’s

Hill, Nuri Trail, 3 Dec 1991, Kiew, R. & Anthony, S. RK3414 (KEP); Fraser’s Hill, Gunong

[Gunung] Peninjau, 1300 m, 26 Aug 1959, Burkill, H.M. HMB2038 (K, SING); Cameron

Highlands, 1450 m, 1 Apr 1930, Henderson, M.R. s.n. (NY, SING); ibidem, 1916, Anthony, S.

SA233 (KEP); ibidem, 1450 m, 18 Jan 1924, Henderson, M.R. 11713 (SING); ibidem, 10 Jul

1979, Everard, B. & Young, D. 94 (K); Cameron Highlands, Tanah Rata, 1450 m, 23 Nov 1925,

Henderson, M.R. SFN17921 (K, SING); Cameron Highlands, Sungai Palas Estate, 1520 m, 8

Sep 1956, Burkill, H.M. HMB856 (K, L, SING); Cameron Highlands, Sungai Ichat, 19 Feb

2008, Rqfidah, A.R., Imin, K. & JJmmul Nazrah, A.R. FRI55631 (KEP, SING); Cameron

Highlands, 1 Ian to Sg. Bisik Waterfall from Ringglet road, 1201 m, 19 Nov 2009, Imin, K,

Ong, P. T., Kueh, H.L. & Paiman, N. FRI66402 (KEP); Rompin, Endau-Rompin State Park,

Trail to Gunung Keriong, 260 m, 20 Aug 2002, Sam, Y.-Y, Apok, K. & Markandan, M. FR14 7150

(KEP); Cameron Highlands, Sungai Ichat, 12 Aug 1931, Jaemat 25181 (KEP); Cameron

Highlands, G. Siku FR, Trail to G. Siku, 1545 m, 21 May 2007, Rosdi, M., Imin, K. & Ummul

Nazrah, A.R. et al. FRI58753 (KEP); Cameron Highlands, Gunung Siku Forest Reserve, 1535

m, 25 Oct 2007, Imin, K. et al. FRI58557 (KEP, SING); Cameron Highlands, Path to Taman

Sedia, 6 Apr 1934, Symington, C.F 36073 (KEP); Cameron Highlands, Jin Kamunting 44th

mile, 1520 m, 21 Jul 1965, Khoo, R. & Ng. S.M. 074 (KLU); Penang: Government Hill, Jul

1894, Curtis, C. s.n. (SING); Perak: unknown loc., Scortechini, B. 36a (SING); Ulu Kinta, 620

m, 19 Apr 1961, Castle-Smith, PM. 15 (K); Gunung Korbu, 610 m, 24 Mar 1913, Robinson,

H.C. s.n. (K); Waterfall north of Tana Rata acc. no. 19680649 vouchered as , Cultivated C7431

(E); Kuala Kangsar, Bubu FR, Gunung Bubu, 938 m, 10 Mar 2010, Julius, A., Coode, M.J.E.

& Angan, A. FRI57716 (KEP); Kuala Kangsar, 850 m, 8 Apr 1995, Chua, L.S.L. FR139082

(KEP); Sunga Ryah, Scortechini, B. 39 (SING); Gunung Batu Puteh, Wray, L. 873 (SING);

Bujang Melaka, 1898, Ridley, H.N. s.n. (SING); Taiping, 910m, 12 Feb 1917, MdHaniff &Md

Nur2314 (K, SING); ibidem. Mar 191 1, Anderson, J. W. 156 (SING); ibidem, Dec 1902, Ridley,

H.N. s.n. (SING); ibidem, Nov 1952, Carter s.n. (SING); Taiping, Gunung Hijau, 14 Jul 1906,

Ernst, A. 1189 (L); ibidem, 8 Oct 1899, Fox, W. s.n. (SING); Taiping, Bukit Lai'ut, Oct 1908,

Long, FR. 14 (K); ibidem, 24 Nov 1 999, Kiew, R. RK4869 (SING); ibidem, 9 Mar 1 939, Spare,

G.H. 2139 (SING); ibidem, 1130-1240 m, 9 Sep 1949, Sinclair, J. SFN38609 (BM, E, K, L,

SING); ibidem, Wray, L. 655 (K, SING); ibidem, 20 Oct 1992, Kamarudin S. FR134629 (K);

ibidem, Birch Hill, 1330 m, 9 Sep 1977, Lewis, G.P. 251 (K, KEP); ibidem, 1060-1210 m, 3

Dec 1965, Shah, M. & Sidek MSI 064 (E, L, SING); ibidem, Sep, Wray, L. 3217 (SING);

ibidem, 1210 m, 22 Nov 1980, Keng, H. et al. 27 (SINU); ibidem, Aug 1967, unknown K6502

(SINU); ibidem, 4 Sep 1995, Heng, H.P et al. HP 12 (SINU); ibidem, 1090 m, 2 Mar 1924,

Burkill, H.M. & Md Haniff 1297 1 (SING); ibidem, 28 Feb 1924, Burkill, H.M. & MdHaniff

12833 (SING); ibidem, 1210-1360 m, 6 Mar 1939, Spare, G.H. 2018 (SING); ibidem. Mar

1884, Scortechini, B. 291 (SING); ibidem, 1000 m, 29 Oct 1969, Everett, B. FRI13553 (KEP);

ibidem, 16 Jan 1994, Anthonysamy, S. SA1155 (KEP); ibidem, 880-1060 m, 15 May 1992,

Aeschynanthus in Singapore and Peninsular Malaysia

41

Rahimatsah Amat N12 (KEP); ibidem, 1060 m, 23 Oct 1962, Merton, L.F.H. 004194 (KLU);

ibidem, 1080 m, 20 Mar 2007, Julius, A. FRJ54942 (KEP, SING); ibidem, Sep 1889, Curtis, C.

s.n. (SING); ibidem, 25 Oct 1972, Hons. Students I (SING); ibidem, 910 m, 29 Oct 1968,

Smith, G. 448 (KLU); Perils: Bukit Bintang Forest Reserve, 7 May 1 988, Latiff, A. & Zainudin,

A. ALM2764 (UKMB); Selangor: Klang, Bukit Changgang, 1 Oct 1937, Md Nur SFN33963

(SING); Klang Gates Rridge, 4 Oct 1981, Kiew, R. RK1081 (KEP); Chemara, 3 Jan 1959,

Carrick, J. 607 (KLU); Telok Forest Reserve, 2 1 Feb 1969, Kochummen, KM. FR12654 (KEP);

ibidem, 18 Jul 1963, unknown 4636 (KLU); Sungei Buloh, 30 Nov 1889, Goodenough, J.S. s.n.

(SING); ibidem, 1890, Goodenough, J.S. s.n. (SING); Gunong [Gunung] Bunga Bua, 28 May

1966, Ng, F.S.P. FRI1136 (KEP, SING); Semangkok Pass, Feb 1904, Napier, W. s.n. (SING);

Tanjong Karang, Sungei Tinggi, 30 m, 27 Sep 1975, Samsiiri Ahmad SA1125 (SING); Hulu

Langat, Gn. Nuang, Above PacatCamp, 1416 m, 12 Nov 201 3. Yao, T.L., Imin, K., Mohd Nazri,

A. & Kueh, H.L. FRI77328 (KEP); Gombak, Klang Gates Ridge, Quartz Ridge, 270 m, 6 Apr

2007, Syahida-Emiza & Angan, A. FR155108 (KEP); Kanching, Bukit Talcun, 300^00 m, 19

Mar 1972, Chin, S.C. 1812 (KLU); ibidem, 300 m, 11 Jul 1965, Stone. B.C.M. 5889 (KLU);

Ulu Besut, Bukit Tangga, 610 m, 20 Jul 1984, Shah, M. & Mahmud MS4978 (E, SING); Kuala

Selangor, Sg. Kai'ang FR, 26 m, 1 Jul 2013, Lim, C.L. FR173033 (KEP); Kuala Selangor,

Sungai Tinggi, 14 Oct 1937, MdNur SFN34079 (BM, K, L, MICH, P, SING); Kanching, Bukit

Takun, 1 50 m, 3 Nov 1 93 7. Md Nur SFN34400 (K, MICH, SING, US); Telok Forest Reserve,

Mar 1965, Whitmore, T.C. FR1227 (K, KEP, L, SING); Genting Highlands, Gunung Bunga

Buah, 10 May 1991, Tan, W.K. et al. TWK12 (SING); Genting Highlands, Old trail to Gunung

Bunga Buah, 1095 m, 18 Jan 2008, Chan, Y.M. & Kiew, R. et al. FR160504 (KEP); Genting

Highlands, Ulu Gombak, 910 m, 14 Jun 1973, Mohd Shah & MohdAli MS 3046 (C, SING);

Terengganu: Ulu Besut, Bukit Tangga, 910 m, 18 Jul 1984, Shah, M. & Mahmud MS4895 (E,

KEP, SING); Bundi, Feb 1904, Rostado 64 (SING); Hulu Terengganu, Sg. Cicir, 4 Aug 2007,

Jutta, M. & Kueh, H.L. FR159583 (KEP).

SINGAPORE: Changi, 10 Dec 1889, Ridley, H.N. 2706 (K); MacRitchie Reservoir, 4 Dec

1948, Sinclair, J. 5376 (E); Bukit Timah, 18 May 1964, Hardial, S. & Samsuri Ahmad 18

(C, L, LAE, SING); Ki'angi, 30 Nov 1899, Ridley, H.N. s.n. (L); Krangi, 1891, Goodenough,

J.S. 2705 (SING); Krangi, 29 Nov 1889, Goodenough, J.S. 2706 (BM, SING); Tuas, 1 May

1891, Goodenough, J.S. s.n. (SING); Chan Chu Kang, May 1889, Corporal s.n. (SING); Bukit

Timah, 7 Jun 1974, Noor, M, MN.1913 (SING); Krangi, 10 Dec 1889, Goodenough, J.S. s.n.

(SING); Tengeh Reservoir, 4 Aug 1890, Goodenough, J.S. s.n. (SING); Krangi, 2 Nov 1889,

Goodenough, J.S. s.n. (SING); Krangi, 7 Dec 1889, Ridley, H.N. s.n. (SING); Ki'angi, 8 Apr

1890, Goodenough, J.S. s.n. (BM); Krangi, 1909, Ridley, H.N. s.n. (BM); Nee Soon Swamp

Forest, 8 Aug 2010, Gw’ee, A.T. S1NG201 0-443 (SING); Nee Soon Swamp Forest, 1 7 Feb 2009,

Staples, G., Leong, P., Chew, PT. & Ibrahim, A. et al. SING2009-157 (KEP, SING); Nee Soon

Swamp Forest, 1 7 Apr 2012, Leong, P, Yam, T. W, Liew, D. & Rodda, M. et al. S1NG2012-165

(SING); Tengeh Reservoir, 1891, Ridley, H.N. 2710 (SING); Mandai Road, Aug 1920, Burkill,

I.H. 6102 (SING); Nee Soon Swamp Forest, 8 May 2013, Ibrahim, H., Lua, H.K. & Saijfudin,

S. S1NG201 3-093 (SING); Nee Soon Swamp Forest, 8 May 2013, Ibrahim, H. & Lua, H.K.

SING-2013-092 (SING).

Notes. This is a very widespread and variable species in both leaf and flower characters.

Although it already has many synonyms further investigation is required as to

whether a number of other species in Borneo should also be included in synonymy. In

Peninsular Malaysia and Singapore this species has most commonly gone by the name

Aeschynanthus parvifolius but was previously synonymised in Middleton (2007) as it

42

Gard. Bull. Singapore 68(1) 2016

falls within the range of variation of the species. Likewise, Aeschynanthus lanceolatus

is not distinct from this variable species.

There are a small number of specimens with pubescent leaves, reminiscent of

Aeschynanthus radicans but with the ovary pubescence of^. pule her. Further studies

should be undertaken to see if there is any possibility of hybridisation between these

two similar species.

10 . Aeschynanthus radicans Jack, Trans. Linn. Soc. London 14: 43 (1823); Brown,

Cyrtandreae 115 (1 839); Steudel, Nomencl. Bot. ed. 2, 1: 32 (1840); Brown in Bennett,

PI. Jav. Rar. 115 (1840); A.DC., Prod. 9: 262 (1 845); Zollinger, Syst. Verz. 3: 56 (1855);

Miquel, FI. Ned. Ind. 2; 720 (1858); C.B.Clarke in A.DC. & C.DC., Monogr. Phan.

5(1): 41 (1883); Clarke in Hooker, FI. Brit. Ind, 4: 343 (1884); Ridley, J. Linn. Soc.

Bot. 32: 501 (1896); Ridley, J. Asiat. Soc. Bengal 74 (2): 736 (1909); Ridley, FI. Mai.

Pen. 2: 500 (1923); Bakhuizen van den Brink, Blumea 6: 395 (1950); Merrill, J. Arnold

Arb. 33: 214 (1952); Barnett, FI. Siam. 3 (3): 202 (1962); Backer & Bakliuizen van

den Brink, FI. Java 2: 524 (1965); Chin, Gard. Bull. Singapore 32: 148 (1979); Turner,

Gard. Bull. Singapore 45: 92 (1993); Turner, Gard. Bull. Singapore 47: 244 (1997

[‘1995’]); Buidt, Thai Forest Bull., Bot. 29: 84 (2001); Smitinand, Thai PI. Names ed.

2, 15 (2001); Middleton, Edinburgh J. Bot. 64: 414 (2007). - Trichosponim radicans

(Jack) Nees, Flora 8: 144 (1825); Memll, Contr. Arnold Arbor. 8: 152 (1934). - TYPE:

Indonesia, Sumatra, Lampung, Gunung Rati Telanggaran, 400 m, 14 November 1921,

Iboet 57 (neotype L, designated by Middleton (2007)). (Fig. 12, 14)

Trichosporum ovatum D.Don ex C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1):

41 (1883), nom. nud.

Aeschynanthus radicans var. robustior C.B.Clarke in A.DC. & C.DC., Monogr. Phan.

5(1): 41 (1883). - TYPE: Kalimantan, Kalimantan Selatan, Banjarmasin, Motley, J.

715 (lectotype K, designated here).

Aeschynanthus radicans var. lanuginosa Ridl., J. Asiat. Soc. Bengal 74 (2): 736

(1909); Turner, Gard. Bull. Singapore 47: 243 (1997 [‘1995’]). - TYPE: Peninsular

Malaysia, Perak, Scortechini, B. 330 (not traced). This variety is placed in synonymy

based on the fact that the characters used to distinguish it are continuously variable in

the species.

Epiphytic and hanging to lithoph 3 Tic and creeping or even epiphytic and climbing by

adventitious roots on the stem; stems sparsely to densely puberulent or with longer

hairs. Leaves opposite; petiole 1^.5 mm long, sparsely to densely puberulent; blade

slightly fleshy, ovate, mostly green above and beneath, not marbled, orbicular or

elliptic, 1-5 X 0.8-2. 6 cm, 1.1-2. 6 times as long as wide, apex rounded and apiculate

or acute to acuminate, base subcordate to obtuse; glabrous to sparsely puberulent all

over above, sparsely to densely puberulent all over beneath, margin entire; secondary

Aeschynanthus in Singapore and Peninsular Malaysia

43

veins obscure, tertiary venation obscure. Inflorescences axillary, generally 1 -flowered;

peduncle 0-3 nun long, bracts elliptic to ovate, 5-6 mm long; pedicels 7-14 mm long,

sparsely to densely puberulent or with longer hairs. Calyx with a tube for most of

length and free lobes, rarely slightly zygomorphic, purple, green with red veins, or

green, puberulent, sometimes with quite long hairs, 19.5-26 mm long; tube 13-22 mm

long which is 65-91% of total length, 6.5-9 mm wide at top of tube; lobes narrowly

triangular, ovate or oblong, erect, 2-8 x 2.6-5 mm, apex rounded. Corolla 47.5-58

mm long, inflated at base, externally tube bright red, lobes bright red, internally tube

yellowish, lobes red with yellowish at base and darker red markings on lower 3 lobes;

upper lobes oblong or ovate, slightly spreading or not, 6.5-10 x 2^.4 mm, sinus 2.4-

6.7 mm deep, apex rounded; lateral lobes ovate or deltoid, slightly spreading or not,

7-10 X 6.2-7. 5 mm, apex rounded; lower lobe elliptic or oblong, slightly spreading or

not, 6.5-10.7 X 5. 6-7. 7 mm, apex rounded, outside densely puberulent, inside glabrous

44

Gard. Bull. Singapore 68(1) 2016

or with sessile glands and short stalked glandular hairs throughout, sessile glands

below lobes. Stamens reaching to end of upper corolla lobes or slightly exserted, fused

in 2 pairs, filaments with very few glandular hairs or sessile glands; anterior filaments

inserted at 27-36 mm from corolla base which is 52-63% of corolla length, filaments

22-27 mm long, anthers 2. 1-3.4 x 1 .5-1 .9 mm; posterior filaments inserted at 31-38.5

mm from corolla base which is 59-70% of corolla length, filaments 15.5-21 mm long,

anthers 2.1-3 x 1.2-1. 7 imn; staminode 0.5-5 mm long. Disk 1-8 mm high, a simple

annular ring. Pistil 45.5-61 mm long; stipe 18-28 mm long, densely puberulent,

often with a mix of glandular and eglandular hairs; ovary 14-28 mm long, densely

puberulent, often with a mix of glandular and eglandular hairs; style 6-15 mm long,

densely pubemlent, often with a mix of glandular and eglandular hairs. Capsule 19-35

mm long, 2,5-3 mm wide. Seed grain 0.8-0. 9 x 0.15-0.3 mm, papillose, bubble cells

present at base of hilar appendage; apical appendage a filiform hair, 7-8 mm long;

hilar appendage a single filiform hair, 6-8 mm long; appendages not papillose.

Distribution. Southern Thailand, Peninsular Malaysia (Johor, Kedah, Kelantan,

Melaka, Negeri Sembilan, Pahang, Penang, Perak, Selangor, Terengganu), Singapore

(but cun'ently considered to be nationally extinct), Sumatra, Borneo.

Habitat and ecology. Epiphytic or clambering over rocks in lowland or hill dipterocarp

forest, often by streams, occasionally in swamp forest, at 0-1000 m altitude.

Provisional lUCN conseryation assessment. Least Concern (LC). This species is

widespread and locally common although there are rather few recent collections of it

from Malaysia. The last collection made of this species in Singapore was in 1931 on

Bukit Timah. It has since been listed as nationally extinct by Chong et al. (2009). At

one timQ Aeschynanthus pulcher was also listed as nationally extinct in Singapore and

has since been rediscovered (Williams, 2014). Aeschynanthus radicans may also still

occur in Singapore in small numbers in the forest canopy or may come back in from

Peninsular Malaysia or Sumatra.

Additional Singaporean and Peninsular Malaysian specimens examined. PENINSULAR

MALAYSIA: Johor: Kota Tinggi, 20 Aug 1954, Sinclair, J. 8151 (E); ibidem, 20 Aug 1954,

Sinclair, J. SFN40358 (SENG); Bekok, Endau State Park, Sungai Selai, 200 m, 16 Aug 2002,

Sam, Y.-Y FRI47104 (KEP); Bekok, Endau State Park, Sungai Selai, 200 in, 16 Aug 2002, Sam,

Y-Y FRI47104 (KEP); Kedah: Weng, 3 Aug 1985, Bogner 1699 (M); Giinung Inas Recreation

Park, 29 May 2001, Sam, Y-Y. FR146549 (KEP); Baling, Gunung Inas Forest Reserve, Bukit

Iboi, 2 Nov 2007, Imin, K., Kiieli, H.L & Phoon, S.N. et al FRI58594 (KEP, SING); Kelantan:

Gua Panjang, 7 Aug 1962, UNESCO limestone expedition 440 A (K, L, SING); Gua Musang,

Kuala Beds, 8 Oct 1985, Latiff,A. & Zainiidin, A. ALM1033 (L); Sg. Ketil, May 1991, Davison,

G. W.H. s.n. (KEP); Jeli, Kuala Yong, 22 Sep 1 986, Latiff, A., Zainudin, A. & Miran, S. ALM1644

(UKMB); Kelumpor, 3 Feb 1923, Md Haniffd MdNur 10390 (SING); Kuala Krai, Stong FR,

8 Aug 2009, Julius, A. & Imin, K. FRI56237 (KEP); Melaka: unknown loc., Cuming, H. 2387

(K); ibidem, Griffith, W. s.n. (K); ibidem, 20 Aug 1916, Burkill, LH. 2159 (SING); Sungei

Rambei River, Jun 1889, Derry, R. 205 (SING); ibidem, Oct 1889, Deny, R. 305 (SING);

Aeschynanthus in Singapore and Peninsular Malaysia

45

Selandar, Oct 1893, Goodenough, J.S. 1518 (SING); Negeri Sembilan: Gunung Angsi, 7 Sep

1937, Franck, C. W. 1150 (C); Selam Forest Reserve, 27 Nov 1922, Holttum, R.E. 9715 (SING);

Bukit Siitu, 1 Nov 1 885. Alvins, M. K 1942 (SING); Jelebu, Berembun FR, Jeram Toi, 446 m, 8

Apr 2008, Yao, T.L., Angan, A. & Norazmi, A. FRI57913 (KEP); Pahang: Tahan River, 1 50 in,

8 Sep 1937, Corner, E.J.H. s.n. (L); Sungei Takan, Aug 1891, Ridley, H.K s.n. (SING); Puku,

Kuala Teku, 21 Dec 1920, Seimimd, E. 390 (SING); Sungai Sat, 18 Jul 1929, Henderson, M.R.

SFN21928 (BM, NY, SING); Jerantut, Tekam Forest Reserve, 3 Oct 2002, Sam, Y.-Y. FRI44492

(KEP); Tasek Bera, 1 6 Mar 1 939, Mashall 35847 (KEP); Jerantut, Tekam Forest Reserve, 3 Oct

2002, Sam, Y.-Y. FRI44492 (KEP); Penang: Government Hill, Jul 1894, Curtis, C. s.n. (SING);

Penang Hill, 500 m, 14 Aug 1986, Weber, A. s.n. (KEP); Beside Hill railway, 14 Aug 1925,

Flippance, F s.n. (SING); Perak: Changkat Mentri, Sep 1918, Kloss, C.B. 6476 (K); Kuala

Kangsar, Bubu Forest Reserve, Sungei Kenas, 410 m, 22 Oct 1992, Saw, L.G. FR137684 (KEP);

Sunkai, 24 Feb 1987, Anthony, S. SA740 (KEP); Perak River, 14 Jan 1963, Allen, B.M. 4852A

(SING); Sungai Ryah, Scortechini, B. 37 (SING); Temengor, 1909, Ridley’, H.N. 14281 (SING);

ibidem, Jul 1909, Ridley, H.N. 14282 (BM); Taiping, Bukit Larut, Sep 1908, Long, FR. 13 (K);

ibidem, Keng, H, etal. I (SINU); Selangor: Ulu Langat, 19 Aug 1959, Gadoh anak Umbai for

A.H. Millard KL1680 (K, KEP, L, SING); ibidem, 23 Aug 1958, Gadoh anak Umbaifor A.H.

Millard KL7 81 (KEP); Genting Sempah, 21 Oct 1921, Hume, H.L. 9228 (SING); Semenyih,

23 Jul 1921, Hume, H.L. 8253 (SING); Genting Highlands, Ulu Gombak, 10 Oct 1921, Hume,

H.L. 8943 (SING); Sungei Buloh, Oct 1899, Goodenough, J.S. s.n. (SING); Terengganu:

Bukit Besar, 760 m, 25 Aug 1901, Annandale, N. & Robinson, H.C. s.n. (CGE, K); ibidem,

910 m, 3 May 1899, Gwynne-Vaughan, D.T. 410 (CGE); Hulu Terengganu, Sg. Cicir, 4 Aug

2007, Jiitta, M. & Kueh, H.L. FR159582 (KEP); Hulu Terengganu, Tembat Forest Reserve, 5

Apr 201 0, Mohd Hairul, M.A., Ong, P.T.,0 ’Byrne, P. & Mohd Nazri, A. et al. FR170936 (KEP);

ibidem, 30 Jul 2009, Rosdi, M., Kamarul Hisham, M. & Angan, A. FRI66336 (KEP).

SINGAPORE: Bukit Timah, 12 Sep 1948, Sinclair, J. s.n. (P); ibidem, 26 Feb 1931, Md Nur

SFN24637 (NY, SING); ibidem, 1891, Ridley, H.N. 2704 (K); ibidem, 12 Sep 1948, Sinclair,

J. 5090 (E); ibidem, 1891, Ridley, H.N. 2704 (SING); ibidem, 1900, Ridley, H.N. s.n. (BM).

Notes. Aeschynanthus radicans is rather similar to rare forais of A. pulcher with hairs

on the leaves but can be distinguished from it by the hairs on the ovary (glands only

in A. pulcher).

11. Aeschynanthus rhododendron Ridh, J. Linn. Soc. 32: 500 (1896); Ridley, J.

Straits Branch Roy. Asiat. Soc. 44: 15 (1905); Ridley, J. Asiat. Soc. Bengal 74 (2):

735 (1909); Ridley, FI. Mai. Pen, 2: 499 (1923); Turner, Gard. Bull. Singapore 47: 244

(1997 [‘1995’]); Middleton, Edinburgh J. Bot. 64: 418 (2007). - TYPE: Peninsular

Malaysia, Perak, Taiping, Gunung Hijau, 1520 m, 1892, Ridley, H.N. s.n. (lectotype

SING [SING0035553], designated by Middleton (2007)). (Fig. 15, 16)

Aeschynanthus longicalyx Ridl., J. Straits Branch Roy. Asiat. Soc. 44: 16 (1905);

Ridley, J. Asiat. Soc. Bengal 74 (2): 735 (1909); Ridley, FI. Mai. Pen. 2: 499 (1923);

Henderson, Malay. Wild. FI. Dicot. 341 (1959); Turner, Gard, Bull. Singapore 47: 243

(1997 [‘ 1 995’]). - TYPE: Malaysia, Selangor, Semangkok Pass, February 1 904, Burn-

Murdoch s.n. (lectotype SING [SING0035404], designated by Middleton (2007);

isolectotype K).

46

Gard. Bull. Singapore 68(1) 2016

Aeschynanthus longicalyx var. superbus Ridl., J. Fed. Malay States Mus. 5: 43

(1914); Stone, Fed. Mus. J. 26 (1): 98 (1981); Middleton, Edinburgh J. Bot 64: 422

(2007); Middleton, Edinburgh J. Bot. 66: 440 (2009). - TYPE: Peninsular Malaysia,

Selangor, Gunung Mengkuang Lebah, 1520 m, 4 February 1913, Robinson, H.C. s.n.

(lectotype SING [STNG0035408], designated by Middleton (2007); isolectotypes BM

[BM000537105], K [K000831889]).

Epiphyte with erect and arching stems; branchlets glabrous. Leaves opposite; petiole

3- 12 mm long, glabrous; blade coriaceous to fleshy, mid to dark green above, paler

beneath, not marbled, ovate or elliptic, 2.1-13 x 0.9-5. 9 cm, 1.3-5. 8 times as long as

wide, apex acuminate, base cuneate to rounded, glabrous above and beneath, punctate

or not beneath, margin entire, secondary veins obscure to clearly visible, 3-5 pairs,

tertiary venation obscure or laxly reticulate. Inflorescences axillary or subterminal

with 1-2 flowers; peduncle absent; bracts linear, c. 2 mm long; pedicels 8-19 mm

long, glabrous. Calyx with a tube and free lobes, rarely slightly zygomoiphic, red,

sometimes with some green at base, glabrous or with a few hairs only on very tips

of lobes, 16-65 mm long; tube 11-37 mm long which is 45-88% of total length,

7.5-27 mm wide at top of tube; lobes triangular or naiTowly triangular, erect, 2.3-32 x

1.8-11 mm, apex acuminate or acute. Corolla 54-102 mm long, tube curved, narrow

at base, externally bright red on tube and lobes, internally red with darker markings

on lower 3 lobes and pale orange at base of lobes and in tube; upper lobes oblong,

slightly spreading to reflexed, 9-22 x 8.5-12 mm, sinus 5-11 mm deep, apex rounded;

lateral lobes ovate or orbicular, reflexed, 7-18 x 7-16 mm, apex rounded; lower

lobe oblong or obovate, spreading or reflexed, 9-21 x 8.5-12 mm, apex rounded;

glabrous to sparsely eglandular puberulent outside, sometimes only around top, inside

with scattered glandular hairs throughout except at base and becoming more dense in

throat, sessile glands inside tube present. Stamens not exserted or only vei*y slightly

exserted beyond upper lobes, fused in 2 pairs; filaments bright red, with glandular

hairs; anterior filaments inserted in tube at 40-50.5 mm from corolla base which is

48-59% of corolla length, filaments 32-41 mm long, anthers 3.6-5 x 1.4-2. 5 mm;

posterior filaments inserted in tube at 51-56 mm from corolla base which is 62-69%

of corolla length, filaments 20-25 mm long, anthers 3-4.5 x 1.1-2. 3 mm; staminode

0.7^ mm long. Disk 0.8-1. 5 nmi high, a simple annular ring or 5-crenate. Pistil 56-

84 mm long; stipe 1 8-33 mm long, glabrous; ovary 20-50 mm long, glabrous; style

7-22 imn long, densely glandular and eglandular pubescent. Capsule 11-22 cm long,

4- 6 mm wide, with a long stipe. Seed grain 0.8-1. 5 x 0.2-0.4 mm, papillose, bubble

cells absent; apical appendage short and stout, 0.7-1. 2 mm long; hilar appendage a

single stout appendage, 0.7-1. 1 mm long; appendages not papillose.

Distribution. Extreme south of Thailand, Peninsular Malaysia (Kelantan, Pahang,

Perak, Selangor), Sumatra.

Habitat and ecology. Recorded from (90-) 1 200-2 1 00 m altitude. Almost all collections

of this species have been made in upper montane forest at over 1200 m altitude or in

Aeschynanthus in Singapore and Peninsular Malaysia

47

Fig. 15. Aeschynanthus rhododendron Ridl. A. Flower with larger-lobed calyx from southern

part of range. B. Flower with smaller-lobed calyx from northern part of range. C. Habit showing

the stamens beginning to wither and reflex in the flower on the right. D. Flower from the front.

— — —

E. Flower dissection. (Photos: A-B, David Middleton; C-E, Jana Leong-Skornickova)

mossy forest at even higher altitudes. However, there is a colleetion, Mohd Shah &

Sidek MS1073 (E, SING) recorded from much lower altitude, 300^00 feet (90-120

m) at Bukit Larut. This may be a mistake.

48

Gard. Bull. Singapore 68(1) 2016

Fig. 16. Distribution of Aeschynanthus rhododendron Ridl. in Peninsular Malaysia (•).

Provisional lUCN conservation assessment. Least Concern (LC). This species is

widespread and locally common.

Additional Peninsular Malaysian specimens examined. PENINSULAR MALAYSIA:

Kelantan: Kuala Krai, G. Tera, Permatang C, 11 Feb 2007, Yao, T.L., Kiew, R., Chew, M.Y. &

Kamarudin S. FRI55845 (KEP); Pahang; Genting Highlands, 25 Nov 1984, Kiew, R. RK1546

(KEP); ibidem, 15 Feb 1987, Worthington, R.D. 12463 (L, NY); ibidem, 1520 m, 14 Jun 1967,

Whitmore, T.C. FR13887 (KEP, L, SING); Genting Highlands, Trail to Bt. Tunggul, 1500 m, 18

Jan 1994, Perumal, B., Gan, C.L., Angan, A. & Bedul FRI41661 (KEP); Genting Highlands,

Gunung Ulu Kali, 25 Mar 2008, Phoon, S.N. & O Byrne, P. et al. FR 1605 5 7 (K, KEP); ibidem,

9 May 1991, Tan, W.K. et al. TWKl (SING); ibidem, 1670 m, 19 Sep 1967, Dransfield, J. s.n.

(KEU); ibidem, 1500 m, 21 Dec 1982, Stone, B.C.M. 15366 (KLU); ibidem, 1767 m, 25 Mar

2008, Phoon, S.N. FRI60496 (K, KEP); ibidem, 1700 m, 24 Sep 1998, Kamarudin S. FRI33755

(KEP); ibidem, 1750 m, 12 Oct 1974, Van Balgooy, M.M.J. 2154 (E, L); ibidem, 1670 m, 18

Mar 1979, Stone, B.C.M. 14045 (KLU, L); ibidem, 1700 m, 4 Jun 1977, Siew Wei Hoe 47 (L);

Aeschynanthus in Singapore and Peninsular Malaysia

49

ibidem, 2 Mar 1996, Van Balgooy, M.M.J. 7133 (K, KEP, L); ibidem, 14 Mar 1982, Stone,

B.C.M. 15071 (BISH); ibidem, 1756 m, 22 Mar 2006, Syahida-Emiza FRI51460 (KEP, SING);

ibidem, 16 Feb 2007, Wilkie, R, Julius, A., Nor Ezzawanis, A.T. & Imin, K. FR152906 (KEP);

Cameron Highlands, 1976, Anthony, S. SA231 (KEP); ibidem, 30 Mar 1932, Mead, J.P 27945

(SING); ibidem, 9 Mar 1947, Wyatt-Smith, J. KEP 56945 (K, KEP, L); ibidem, 1820-2000 m,

Aug 1975, Keng, H. et ai K8010 (SINU); ibidem. Batten Pooll, A.H. s.n. (SING); ibidem, 2120

m, 1 Feb 1962, Togashi, M. 622111 (Tl); ibidem, 2000 m, 5 Mar 1970, Togashi, M. s.n. (Tl);

ibidem, 1820-2000 m, 25-31 Aug 1975, Rao & et ai K8010 (AAU, L); Cameron Highlands,

Ulu Bertam Forest Reserve, 1450 m, 9 Nov 1959, Abbe, E.C., Kadim bin Passim, Mansor bin

Omar & Abbe, L.B. 9119 (L, NY); Cameron Highlands, Gunung Batu Berinchang, Tarred road

to microwave station descending 11 km to Brinchang, 2000 m, 7 May 2007. Yao, T.L.,

Nooteboom, H.P. &Angan,A. FRI55898{KEPy, Cameron Highlands, Gunung Batu Berinchang,

1620 m, 20 Jan 2010, Mohd Hairul, M.A., Ong, P T, Siti Muni rah, M. Y. & Kueh, H.L. FR169938

(KEP); ibidem, 1950 m, 20 Jan 2011, Imin, K., Mohd Nazri, A. & Julius, A. et al. FR171954

(KEP); ibidem, 9 Apr 1987, Saw, L.G. FR134368 (KEP); ibidem, 22 Aug 1995, Tam Shell May

TSM2 (KEP); ibidem, 1997 m, 11 Feb 2010, Saw, L.G., Damahuri, S. & Norzamli, A. FRI48241

(BKF, KEP); ibidem, 2020 m, 30 Aug 1970, Chin, S,C. 233 (KLU); ibidem, 1980 m, 22 Mar

2007, Julius, A. FR156020 (KEP, SING); ibidem, 5 Nov 1960, Poore, M.E.D. 466 (KLU);

ibidem, 30 Jul 1967, Stone, B.C.M. 7218 (KLU); ibidem, 29 Jul 2007, Low, Y.W. LYW140

(KLU); ibidem, 1920 m, 9 Dec 1961, Abdul Samat bin Abdullah 68 (KLU); ibidem, 2024 m, 26

Oct 2007, Imin, K. et al. FRI 58570 (K, KEP, SING); ibidem, 1945 m, 21 Feb 2008, Syahida-

Emiza FR157293 (K, KEP, SING); ibidem, 1910 m, 10 Jun 1961, Castle-Smith, P.M. 42 (K);

ibidem. 17 Mar 1996, Van Balgooy, M.M.J. 7228 (KEP, L); ibidem, 1910 m, 18 Apr 1968,

Woods, P.J.B. 680 (E); ibidem, 1870 m, 26 Jul 2002, Bramley, G. & Neale, S. GB31 (E, K,

KEP); ibidem, 2000 m, 20 Jun 1975, Van Balgooy, M.M.J. 2663 (E, L); ibidem, 13 Sep 1985,

Latiff, A., Zainudin, A. & Miran, S. ALM970 (L); ibidem, 1880 m, 12 Aug 1986, Wong, K.M.

FR135244 (K, KEP, L, SING); ibidem, 2020 m, 4 Nov 1958, Sinclair, J. 9949 (E, SING);

ibidem, 1970 m, 11 Oct 1963, Chew, W.-L. CWL922 (K, L, SING); ibidem, 2020 m, 7 Mar

1960, Hawkins, A.S.M. 6 (SING); ibidem, 2 Apr 1989, Latiff, A. & Zainudin, A. ALM3121

(UICMB); ibidem, 1520 m, 15 Feb 1975, Mohd Shah MS 3446 (SING); ibidem, 8 Sep 1970,

Whitmore, T.C. FR115450 (K, KEP, L, SING); ibidem, 1820 m, 9 Apr 1930, Henderson, M.R.

SFN23534 (NY, SING); ibidem, 2020 m, 8 Apr 1960, Abbe, L.B., Abbe, E.C., Kadim bin Passim

& Mansor bin Omar 9767 (K, L, NY); Gunung Benom, 14 Nov 2009, Mohd Hairul, M.A. &

Mohd Nazri, A. et al. FRI69892 (KEP); Cameron Highlands, Gunung Berembun, 1750 m, 5

Apr 1988, Sabari, D. FRI32715 (KEP); ibidem, 28 Jun 1988, Samsuri Ahmad SA96 (SINU);

ibidem, 9 Aug 2008, Rosdi, M. & Plioon, S.N et al. FRI59867 (KEP); ibidem, Nov 1908,

Ridley, H.N 13600 (BM, K, SING); ibidem, 1820 m, 25 Nov 1925, Henderson, M.R. 17988

(SING); ibidem, 4 Apr 1930, Henderson, M.R. s.n. (SING); ibidem, 1820 m, 10 Sep 1970,

Whitmore, PC. FRH5495 (K, KEP, L, SING); ibidem, 1670 m, 3 Oct 1963, Chew, W.-L.

CWL772 (C, G, K, L, SING); Cameron Highlands, Trail to G. Berembun, 1670 m, 1 Mar 1968,

Ng, F.S.P. FRI5931 (K, KEP, L); Cameron Highlands, near Parit Falls, Mar 1952, .lohnston,

A.M. 76 (SING); Cameron HigWands, Break Pressure Tank Hill, 1480 m, 3 1 Aug 1956, Burkill,

H.M. HMB760 (K, L, LAE, SING); Cameron Highlands, Tanah Rata, 1400-1500 m, 12 Jan

1983, Davis 69248A (E [2 sheets], SING); ibidem, 1440 m, 29 Aug 1956, Burkill, H.M.

HMB730 (K, L, SING); Cameron Highlands, Tanah Rata, Parit Waterfall, 28 Aug 1 990, Okada,

K, Darnaedi, D., Akiyama, H, Kawahara, T. & Watano, Y. 1027 (Tl); Cameron Highlands,

Robinson’s Falls, 1430 m, 24 Mar 2007, Julius, A., Middleton, D.J. & Lindsay, S. et al.

FRI57483 (KEP); ibidem, 1400-1840 m, 20 Mar 1992, Klackenberg, J. & Lundin, R. 689 (L);

50

Gard. Bull. Singapore 68(1) 2016

ibidem, 9 May 1964, Mahmud bin Sidek 4816 (KLU); ibidem, 14 Apr 1968, Woods,

Black, M. & Wycherley, R 615 (E, KEP); Cameron Highlands, Tanah Rata, 1440 m, 29 Aug

1970, Chin, S.C. 165 (KLU); Cameron Highlands, Brinchang Mist Forest, 1670 m, 20 Feb

1962, Poore, M.E,D, 1028 (KLU); Gunung Benom, 31 Jul 1925, unknown s,n. (SING); ibidem,

28 Jul 1925, Federated Malay States Museum Coll s.n. (K); Gunung Mengkuang, 1520 m, 28

Mar 1959, Wyatt-Smith, J. 78808 (KEP); Gunung Tahan, Tangga Duabelas, 1530 m, 24 Mar

1987, Kiew\ R. RK2430 (KEP); Above Genting Sempah, 1440 m, 23 Jul 1967, Stone, B.C.M.

7194 (KLU, L); Fraser’s Hill, 15 Nov 1977, Keng, H. et al. 410 (SINU); ibidem, 1450 m, 27

Aug 1923, Henderson, M.R. 11365 (SING); Fraser’s Hill, Pine tree hill, 1450 m, 19 Sep 1922,

Burkill, HM. & Holttum, R.E. 8531 (SING); ibidem, 1450 m, 27 Aug 1923, Md Nur 11056

(SING); Gunong [Gunung] Gedong, 3 Sep 1928, Holttum, R.E. 20769 (SING); Gunung Tahan,

1911, Ridley, H.N. 16090 (K, SING); Perak; unknown loc., Scortechini. B. s.n. (SING);

Caulfield’s Hill, 1210 m, Apr 1884, Scortechini, B. 388 (SING); ibidem, Aug 1885, Wray, L.

656 (K, SING); Taiping, Scortechini, B. 468b (SING); ibidem, 12 Nov 1889, Curtis, C. s.n.

(SING); Taiping, Gunung Hijau, 1530 m, 7 Jun 1983, Stone, B.C.M. 15497 (KLU); ibidem, 7

Mar 1939, Spare, G.H. 2047 (SING); ibidem, Sep 1889, Curtis, C. s.n. (SING); ibidem, 1448

m, 19 Mar 2007, Julius, A., Middleton, D.J. & Lindsay, S. et al. FRI53302 (KEP, SING);

ibidem, 1448 m, 14 Jul 2006, Kamarul Hisham, M. FR152054 (KEP); ibidem, 1440 m, 5 Mar

1924, Henderson, M.R. 11829 (SING); ibidem, 1360 m, 14 Feb 1907, Md Haniff & Md Nur

2456 (SING); ibidem, 8 Oct 1899, Fox, W. s.n. (SING); ibidem, 1210 m, Dec 1887, Curtis, C.

1311 (K, SING); ibidem, 1210-1360 m, 4 Dec 1965, Mohd Shah & Sidek MSll 10 (SING);

ibidem, 1520 m, 1892, Ridley, H.N. s.n. (SING); ibidem, 1380 m, 11 Sep 1949, Sinclair, J. &

Kiah SFN38670 (BM, E, K, L, P, SING); Sungei Pelus, 1520 m, 1889, Wray, L. s.n. (SING);

Gunong [Gunung] Inas, 1180-1360 m, 9 Dec 1899, Yapp, R.H. 440 (CGE^ R); Gunung Batu

Puteh, 1920 m, 26 Feb 1994, Perumal, B., Gan, C.L., Shahril, K.Z. & Angan, A. FRI41602

(KEP, KLU); Gunong [Gunung] Jasar, 23 Aug 1977, Ng, FS.P FR127146 (KEP); Gunong

[Gunung] Jasar, 17 Apr 1968, Woods. P.J.B. 632 (E); Gunung Korbu, 1670 m, 10 Mar 1913,

Robinson, HC. s.n. (K); Taiping, Bukit Larut, Bukit B intang Hijau, 1451 m, 1 Mar 2011,

Wilkie, P., Siti Munirah, M.Y., Mohd. Hairul, M.A. & Nazre FRI75005 (E, KEP); ibidem, 16

Aug 1986, Weber, A. s.n. (KEP); ibidem, 90-120 m, 9 Dec 1965, Mohd Shah & Sidek MS 1073

(E, SING); ibidem, 1270 m, 20 Oct 1988, Saw, LG. FRI36398 (KEP); ibidem, 27 Jun 1984,

Keng, H. et al. D-li5 (SINU); Selangor; Semangkok Pass, Feb 1904, Burn-Murdoch s.n. (K,

SING); Gunung Mengkuang Lebah, 1520 m, 16 Jan 1913, Robinson, H.C. s.n. (K); ibidem,

1520 m, 17 Jan 1913, Robinson, H.C. s.n. (K); ibidem, 1520 m, 4 Feb 1913, Robinson, H.C. s.n.

(BM, K, SING); Bukit Fraser, Jul 1899, Hose, G. 46 (SING); Gunong [Gunung] Bunga Buah,

1360m acc. no. 19680624 vouchered as Cultivated C7315 (E, KEP); Ulu Langat, Feb 1912,

Kloss, C.B. s.n. (K); Hulu Langat, Ulu Langat Forest Reserve, 1440 m, 25 Jan 2003, Sam, Y.-Y.

FR147208 (KEP); Ulu Langat, Gunung Nuang, 12 May 1940, Symington, C.F 51762 (KEP);

Gunong [Gunung] Nuang, 15 Aug 1974, Lee, D. W. s.n. (KLU); Ulu Selangor, Gunung Moyang,

3 Nov 1940, Symington, C.F. 56711 (KEP).

Notes. This species is rather variable in the shape of the calyx. The type of the species

is from Gunung Hijau in the Taiping Hills where the calyx is generally naiTow and

the lobes small. This is the fonn also found in Thailand. In the Genting Highlands,

Fraser’s Hill and Gunung Benom the calyx tube flares gently from the base so that the

mouth is wider and the lobes are much larger. This is the plant fonnerly recognised

as Aeschynanthus longicalyx. The largest calyx seen is from Gunung Ulu Kali in the

Genting Highlands. Material from the Cameron Highlands is rather intermediate.

Aeschynanthus in Singapore and Peninsular Malaysia

51

12 . Aeschynanthus speciosus Hook., Bot. Mag. 73: t.4320 (1847); Miquel, FI. Ned.

Ind. 2: 718 (1858); Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 33 (1883); Ridley,

J. Linn. Soc. Bot. 32: 499 (1896); Ridley, J. Straits Branch Roy. Asiat. Soc. 44: 14

(1905); Ridley, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74(2): 733 (1909); Ridley, FI.

Mai. Pen. 2: 498 (1923); Turner, Gard. Bull. Singapore 47(1): 244 (1997 [H995’]);

Burtt, Thai Forest Bull., Bot. 29: 84 (2001); Smitinand, Thai PI. Names, ed. 2: 15

(2001); Middleton, Edinburgh J. Bot. 64: 420 (2007). - Trichosporum speciosum

(Hook.) Kuntze Revis. Gen. PL: 478 (1891). - TYPE: Java, Jawa Barat, Mt Asapan,

Lobb, T. s.n. (lectotype K, designated by Middleton (2007)). (Fig. 17, 18)

Aeschynanthus aucklandiae Low, Sarawak: 386 (1848). - TYPE: Borneo, East

Malaysia, Sarawak, Low, H. s.n. (holotype CGE).

Epiphyte with erect, arching and pendulous stems; branches glabrous. Leaves in whorls

of 3-6; petiole generally wide and flat, sometimes slightly winged, 2-12 mm long,

glabrous; blade coriaceous or slightly fleshy, mid to dark green above, paler beneath,

not marbled, elliptic or ovate, 3.8-15.3 x 1.2-5. 5 cm, 2.3-6. 1 times as long as wide,

apex acuminate to caudate, base cuneate to rounded, glabrous above and beneath,

not punctate beneath, margin dentate to entire, often strongly undulate, secondary

veins obscine to weakly visible, c. 7-8 pairs, tertiary venation obscure. Inflorescences

tenninal with 4-12 flowers; peduncle absent; bracts linear, 5-13 mm long; pedicels

7-14.5 mm long, glabrous or sparsely eglandular puberulent. Calyx of separate lobes

free to the base, pale green, yellowish or dark reddish brown and then often flushed with

one of the other colours, glabrous to densely eglandular or glandular pubemlent; lobes

nan'owly triangular or linear, erect, 6.5-26.3 x 1-2.6 mm, apex acute or acuminate.

Corolla 54-118 mm long, tube naiTOw at base, externally yellow, yellowish green or

orangish on basal half to two-thirds of tube and orange-red to bright red above, more

rarely bright red all over, lobes orange-red to bright red, internally light yellowish

in tube and red or orangish on upper 2 lobes and red or orangish at the margin with

darker V- or W-shaped markings on lower 3 lobes and yellowish at the base; upper

lobes oblong, not spreading or reflexed, 4.5-11 x 3.6-8 mm, sinus 4-8.2 mm deep,

apex rounded; lateral lobes oblong, deltoid or ovate, spreading, 5.7-14 x 5.7-13.5

mm, apex rounded; lower lobe oblong or elliptic, spreading or reflexed, 7-15.5 x

4.5- 12 mm, apex retuse to rounded; outside glabrous, slightly papillose or sparsely

glandular puberulent, sometimes only on ciliate lobes, inside with sparse sessile

glands. Stamens long exserted, fused in 2 pairs; filaments bright red or white, with

glandular hairs, anthers grey, pale brown or purple-black; anterior filaments inserted at

50-96 mm from corolla base which is 67-81% of corolla length, filaments 29-46 mm

long, anthers 3.9-7 x 1.3-2. 3 mm; posterior filaments inserted at 53-100 mm from

corolla base which is 72-85% of corolla length, filaments 21-38 mm long, anthers

2.6- 5 X 1. 2-2.1 mm; staminode 0.3-4 mm long. Disk 1.2-2. 5 mm high, 5-dentate

or 5-crenate. Pistil 83-1 30 mm long; stipe 20-30 mm long, with few sessile glands

or glabrous; ovary 27-44 mm long, with sessile glands, these sometimes very few;

style yellow or green, 14-65 mm long, glandular pubescent, especially in upper half

52

Gard. Bull. Singapore 68(1) 2016

Capsule 20-45 cm long, 2. 3-3. 6 mm wide. Seed grain 0.9-1. 3 x 0.3-0.4 mm, warty,

bubble cells absent; apical appendage a filifonn hair, 1 5-22 mm long; hilar appendage

a single filiform hair, 15.5-23 mm long; appendages papillose.

Distribution. Southern Thailand, Peninsular Malaysia (Pahang, Perak, Selangor),

Sumatra, Java, Borneo.

Habitat and ecology. In hill dipterocarp forest habitats at 240-760 m altitude. It has

been recorded to 1900 m altitude in Sumatra and is likely to also occur at higher

altitudes in Peninsular Malaysia.

Provisional lUCN conseiyation assessment. Least Concern (LC). This species is

remarkably infrequently collected despite having very showy flowers such that it is

readily observed in the forest for collection. It is likely to be relatively rare over most

of its range but both the EOO and AOO are much higher than those suggesting a threat

categoiy would be merited.

Peninsular Malaysian specimens examined. PENINSULAR MALAYSIA: Pahang: Bentong

Sg., Naming, 760 m, 18 Jan 1959, unknown KEP93108 (KEP); Rompin, Endau-Rompin State

Park, Ulii Kinchin, Sg. Kinchin, Norain, Raihana & Rosjana 07 (SINU); Perak: Near Ulu

Selama, 8 Jan 1900, Yapp, R.H. 610 (CGE); Bruseh near Bidor, 240 m, 27 .lun 1904, Napier,

W. s.n. (SING); Selangor: Ulu Langat, Gadoh, U. KL1319 (KEP, SING); Serendah Ridge, 300

m, 18 Jul 1957, Hawkins, A.S.M. s.n. (SING); Bukit Elam, Jan 1891, Kelsall, H. s.n. (SING).

Notes. It is possible that Aeschynanthus pseudohybridus Mendum from Borneo is a

synonym of this species but further study is necessary. It differs in the more uniformly

red flowers compared to the orange and yellow flowers of A. speciosus.

13. Aeschynanthus volubilis Jack, Trans. Linn. Soc. London 14: 42 (1823); Brown,

Cyrtandreae 115 (1839); Steudel, Nomencl. Bot. ed. 2, 1: 32 (1840); Brown in Bennett,

PI. Jav. Rar. 115 (1840); A.DC., Prod. 9: 262 (1845); Miquel, FI. Ned. Ind. 2: 719

(1858); C.B. Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 46 (1883); Balchuizen

van den Brinlc, Blumea 6: 395 (1950); Merrill, J. Arnold Arb. 33: 214 (1952); Backer

& Bakhuizen van den Brink, FI. Java 2: 524 (1965); Sinha, El. Gt. Nicobar Isl. 329

(1999). - Trichosporum volubile (Jack) Nees, Flora 8: 144 (1825). - TYPE: Plate II,

fig. 3 in Jack, Transactions of the Linnean Society of London vol. 14, unnumbered

page between pp. 44 and 45 (1823), lectotype designated here. Epitype: Sumatra, East

Coast [Sumatera Utara], Asahan, Silo Maradja, June 1927, Bartlett 8695 (epitype NY,

designated here; isoepitype MICH). (Fig. 19)

Aeschynanthus obovatus C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 47

(1883). - Trichosporum obovatum (C.B.Clarke) Kuntze, Revis. Gen. PI. 478 (1891). -

TYPE: Kalimantan, Kalimantan Selatan, Banjarmasin, Motley, J. 1158 (holotype K).

Aeschynanthus in Singapore and Peninsular Malaysia

53

Fig. 17. Aeschynanthus speciosus Hook. A. Habit. B. Flowers, side view. The flower on the

right is older with a much longer style and the stamens beginning to reflex (and with one

pair becoming detached). C. Flower dissection with 2 calyx lobes removed to view nectary.

(Photos: Jana Leong-Skomickova)

Aeschynanthus obovatus var. pallidus C.B. Clarke in A.DC. & C.DC., Monogr.

Phan. 5(1): 48 (1883). - TYPE: Java, Zippelius, A. Leiden number 52 (leetotype L

[L0281661], designated here; isoleetotypes L [L0281673, L0281674]).

Aeschynanthus hoseanus KraenzL, J. Linn. Soe. Bot. 37: 284 (1906). - Trichosporum

hoseanum (KraenzL) Merr., J. Straits Branch Roy. Asiat. Soe. special number: 530

54

Gard. Bull. Singapore 68(1) 2016

Fig. 18. Distribution of Aeschynanthus speciosus Hook, in Peninsular Malaysia (•).

(1921). - TYPE: Borneo, East Malaysia, Sarawak, Saribas, 24 Nov 1893, Haviland,

G.D. & Hose, C. 3528 (holotype K; isotype BM [BM000797673]).

Aeschynanthus flippancei Ridl., Bull. Misc. Info. Kew. 1926: 473 (1926); Turner,

Gard. Bull. Singapore 47: 243 (1997 [‘1995’]). - TYPE: Cultivated in Penang

Botanic Gardens but originally from Malaysia, Penang, Balik Pulau, Flippance, F s.n.

(lectotype K, designated here; isolectotype SING [SING0035633]).

Epiphyte with erect, arching or pendulous stems; stem glabrous. Leaves opposite;

petiole 3-14 mm long, glabrous or sparsely puberulent; blade coriaceous, mid to dark

green above, paler beneath, elliptic or ovate, 1.8-5. 7 x 0. 7-3.2 cm, 1.6-2. 7 times as

long as wide, apex obtuse to acuminate, base rounded to cuneate, glabrous above and

beneath, not punctate beneath, not variegated, margin entire, 3-5 pairs of secondary

veins, weakly visible or obscure, tertiary venation obscure. Inflorescence subterminal

Aeschynanthus in Singapore and Peninsular Malaysia

55

or axillary, 3^-flowered; peduncle 3-5 mm long; pedicels 7-12 mm long, glabrous.

Calyx with a tube for most of length and free lobes, tube fairly broad and gently flaring

from base, 10-16.5 mm long; tube 8.8-14 mm long, 83-90% of total length, c. 6 imn

wide at top of tube; lobes triangular or semicircular, slightly spreading or erect, 1-2.5

X 2.3-5 mm, apex rounded to acute, rarely acuminate. Corolla 19.5-27 mm long, tube

slightly curved or almost straight, broad at base; upper lobes oblong, not spreading or

reflexed, 3. 2^.5 x 2.5 mm, sinus 1.5-2 mm deep, apices rounded; lateral lobes deltoid,

reflexed, 3-5.2 x 3. 8^.4 mm, apices rounded; lower lobe oblong, spreading, 4.2-5. 5

X 3.2-4 mm, apex rounded; outside glabrous except for ciliate lobes, inside with sparse

glandular hairs in upper part of tube and more densely so on inside of lobes except near

margin. Stamens long exserted, fused in 2 pairs; filaments with sparse short stalked

glands throughout; anterior filaments inserted at 14-17 mm from corolla base which

is 52-63% of corolla length, filaments 16-18 mm long, anthers 2.4 x l mm; posterior

filaments inserted at 16-19 mm from corolla base which is 48-67% of corolla length,

filaments 12.5-13 mm long, anthers 2-2.2 x 1-1.6 mm; staminode 0.4-1. 3 mm long.

Disk 1-1.2 mm high, a simple annular I'ing or 5-crenate. Pistil 18-32 mm long; stipe

1.7- 8 mm long, glabrous; ovary 10-16 mm long, with very few sessile glands and

appearing glabrous; style 4-8 mm long, glandular pubescent. Capsule 10-28 cm long,

2.8- 3 mm wide. Seed grain 0.5-0. 6 x 0. 1-0.2 mm wide, smooth, bubble cells present

at base of hilar end; apical appendage a filiform hair, 4-8.5 mm long; hilar appendage

a single filiform hair, 7.5-9 mm long.

Distribution. Peninsular Malaysia (Penang), Sumatra (but see note below).

Habitat and ecology. None of the collections I have seen of this species from Malaysia

or elsewhere include habitat data or altitudinal range.

Provisional lUCN conservation assessment. Data Deficient (DD). The limits of the

species and its distribution are unclear.

Additional Peninsular Malaysian specimens examined. PENINSULAR MALAYSIA: Penang:

BalikPulau, Flippance, F s.n. (K, SING); Government Hill, May 1921, Md Haniff s.n. (SING).

Notes. Aeschynanthus volubilis is the type species of the genus. It is known in

Peninsular Malaysia only from a small number of specimens formerly identified as

A eschynanyth us flippancei.

There are a number of species and specimens from Borneo and eastern Malesia

which appear similar to Aeschynanthus volubilis but require further study to ascertain

whether they are synonyms and/or extensions of the distribution of the species.

14 . Aeschynanthus wallichii R.Br., Cyrtandreae 116 (1839); Brown in Bennett, PI.

Jav. Rar. 116 (1840); de Candolle, Prod. 9: 263 (1845); Miquel, FI. Ned. Ind. 2: 722

(1858); C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5(1): 48 (1883); Clarke in

56

Gard. Bull. Singapore 68(1) 2016

Hooker, FI. Brit. Ind. 4: 343 (1884); Ridley, J. Linn. Soc. Bot. 32: 502 (1896); Ridley,

J. Asiat. Soc. Bengal 74 (2): 736 (1909); Ridley, FI. Mai. Pen. 2: 500 (1923); Turner,

Gard. Bull. Singapore 45: 92 (1993); Turner, Gard. Bull. Singapore 47: 244 (1997

[‘1995’]). - Aeschynanthus mdicans Wall., Num. List no. 798 (1829), nom. nud. -

TYPE: Singapore, 1822, Wallich, N. 798 (lectotype K-W [K001111917], designated

here; isolectotypes CGE, K [K000831885]). (Fig. 12, 19)

Aeschynanthus brevicalyx Miq., FI. Ned. Ind. 2: 720 (1858); C.B. Clarke in A.DC. &

C.DC., Monogr. Phan. 5(1): 49 (1883). - Trichosporum brevicalyx (Miq.) Kuntze,

Revis. Gen. PI. 477 (1891). - TYPE: Indonesia, Sumatra, Sikilieh, Teijsmann, J.E.

HB1194 (lectotype U, designated here; isolectotype L).

Epiphytic; stems glabrous. Leaves opposite; petiole 5-12 mm long, glabrous; blade

coriaceous or slightly fleshy, ovate or elliptic, 3.1-10.5 x 0.8-5 cm, 1.7-3. 8 times as

long as wide, apex acuminate, base cuneate to subcordate, not marbled, glabrous above

and beneath, not variegated, margin glabrous, entire, 4-5 pairs of secondary veins,

only weakly visible or obscure, tertiary venation obscure. Inflorescence subterminal

or axillary, 3-6-flowered; peduncles to 2.5 mm long, subglabrous to densely eglandular

puberulent; pedicels 4.2-9 mm long, green, subglabrous to densely eglandular

puberulent. Calyx fused into a wide open cup with lobes semicircular or only as curves

on rim, green or yellow-green, sparsely to densely eglandular pubemlent, sparsely

pubescent inside, 2. 5-6. 5 mm long; tube 1. 9-5.5 mm long which is 70-90% of total

length, tube 4-7 mm wide at apex; lobes semicircular or a weak curve of the rim,

spreading or erect, 0.6-1 .2 x 3. 5-5. 5 nmi, apices rounded. Corolla 19.5-27 mm long,

tube slightly curved or more or less sti'aight, externally bright or dark red, base of tube

not gibbose, outside sparsely to densely glandular to eglandular puberulent, inside

with short stiff upward pointing hairs near base or throughout, sessile glands at inside

sinuses of lobes; upper lobes orbicular or oblong, not spreading or reflexed, 3. 8-6. 6 x

3^.5 mm, sinus 2.2-3. 1 mm deep, apices rounded; lateral lobes ovate or deltoid, not

spreading or reflexed, 5-8 x 4.2-6. 5 mm, apices rounded; lower lobe ovate or orbicular,

not spreading or reflexed, 4.7-7 x 4.7-9 mm, apex rounded. Stamens slightly or not

exserted, flised in 2 pairs, filaments papillose; anterior filaments inserted at 10-12.7

mm from corolla base which is 48-51% of corolla length, filaments 14-15 mm long,

anthers 1 .9-2.2 x 0.9-1 mm; posterior filaments inserted at 1 1 .5-14.5 mm from corolla

base which is 55-58% of corolla length, filaments 10-11 mm long, anthers 1.7-2. 1 x

0.9-1 mm; staminodes 1-1.2 mm long. Disk c. 0.9 mm high, a simple annular ring

or 5-crenate. Pistil 16.5-22 mm long; stipe 1.5-4 mm long, glabrous or with few

sessile glands; ovary 12.5-14.7 mm long, with sessile glands; style 2. 5-3. 3 mm long,

glandular pubescent; stigma circular. Capsule 18-37 cm long, c. 4 mm wide. Seed

grain 0.8-1 .2 x 0.2-0.3 mm, papillose, bubble cells present at base of hilar appendage;

apical appendage a filiform hair, c. 9 mm long; hilar appendage a single filiform hair,

c. 10 mm long; appendages not papillose.

Distribution. Peninsular Malaysia (Johor, Pahang, Perak, Terengganu), Singapore,

Sumatra.

Aeschynanthus in Singapore and Peninsular Malaysia

57

Fig. 19. Distribution of Aeschynanthus volubilis Jack (▼) in Peninsular Malaysia and

Aeschynanthus wallichii R.Br. (•) in Singapore and Peninsular Malaysia.

Habitat and ecology. In lowland dipterocarp forest, or in peat swamp forest at 0-760

m altitude.

Provisional lUCN consen^ation assessment. Least Concern (LC). This species is

relatively infrequently collected but has an EOO and AOO considerably above levels

considered to be concerning. In Singapore it is considered to be Critically Endangered

and is now confined to Nee Soon Swamp.

Additional Singaporean and Peninsular Malaysian specimens examined. PENINSULAR

MALAYSIA: Johor: 5 miles S of Labis FR, 17 Apr 1966, Whitmore, T.C. KEP207 (KEP);

Mawai-Sedili Road, 24 Jan 1961, Chew, W.-L. CWL222 (C, E, K, L, SING); Kuala Sedili

fresh water swamp forest, 6 m, 26 May 1961, Burkill, H.M. HMB2668 (SING); Kota Tinggi,

Kuala Sedili New Road, 23 Jun 1959, Kadim & Noor, M. 138 (SING); ibidem, 23 Jun 1959,

Kadim & Noor, M. 131 (SING); ibidem, 25 Sep 1959, Shah, M. & Noor, M. MS813 (E, L,

58

Gard. Bull. Singapore 68(1) 2016

SING); ibidem, 23 Jun 1959, Shah, M. 138 (K); Kota Tinggi, Gunung Panti West, 550 m,

19 Jul 1981, Maxwell, J.F. 81-182 (L, SING); Gunung Panti, 520 m, 5 Jul 1970, Samsuri

Ahmad SA318 (SING); ibidem, 4 Mar 1987, Kiew, R. RK2398 (SING); ibidem, 470 m, 24 Aug

1986, Wong, K.M. s.n, (KEP); ibidem, 22 Jan 1994, Kiew, R. RK3775 (KEP); ibidem, 4 Mar

1987, Kiew, R. RK2398 (KEP); ibidem, 460 m, 31 Jul 2008, Yao, T.L., Lim, C.L., Rosdi, M.

6 Ayau, K. FR165386 (KEP); ibidem, 480 m, 5 Feb 1981, Collenette, IS. 2254 (E); Pontian,

Pengkalan Raja, 28 Jun 1939, Henderson, M.R. SFN36656 (K, SING); Gunung Ledang, Lobb,

T 111 (K); Alor Bukit, 22 Nov 1966, Hardial, S. 537 (K); Sungai Kayu, 12 Oct 1936, Kiah

SFN32054 (BM, K, LAE, SING); Alor Bukit, 22 Nov 1966, Hardial, S. 537 (SING); Endau,

Sg. Sempanong, 4 Feb 1986, Kiew, R. RK2112 (SING); Ulu Batu Pahat, Kampong Simpai,

1892, Lake & Kelsall, H. s.n. (SING); Sebrau [?Tebrau], Aug 1908, Ridley, HN. s.n. (SING);

Tempayang River, 1908, Ridley, H.N. s.n. (BM, SING); Tanjong Kupang, Jun 1890, Ridley,

HN. s.n. (SING); Nov-Dec 1935—1935, Vesterdal, A. 306 (C); Kluang, 20 Nov 1922, Holttum,

R.E. 9489 (SING); ibidem, 7 Mar 2000, Morgany, T, Tan, H.T.W. & Loo, A.H.B. M260 (SINU);

Pahang: Pekan, Pekan Forest Reserve, Sungai Bebar, 21 Apr 2005, Zainon, A.S. FRI50105

(KEP); Rompin, Endau-Rompin State Park, Trail to Gunung Keriong, 20 Aug 2002, Sam, Y.-

Y. FR147144 (KEP); Perak: Chenderiang, Gunong [Gunung] Bujang Melaka, 120 m, 10 Aug

1959, Mrs. Allen & Kadim 436 (SING); Terengganu: Ulu Brang, 760 m, Jul 1937, Moysey,

L. & Kiah SFN33624 (LAE, SING); Sekayu, 7 May 1988, Kiew, R. RK2696 (KEP); Sakayu,

29 Aug 1986, Anthony, S. SA639 (KEP); Dungun, Pasir Raja Forest Reserve, Hutan Lipur

Cemerung, 18 Oct 2002, Sam, Y.-Y. FR147158 (KEP); ibidem, 18 Mar 1998, Anonymous s.n.

(SING); Ulu Brang, Terengganu Tambahan FR, Sg. Chib, 360 m, 26 Mar 201 3, Imin, K., Ong,

P.T.& Kiieh, HL. FRL77847 (KEP).

SINGAPORE: unknown loc., 1822, Wallich, N. 798 (CGE, K-W); ibidem, unknown s.n.

(SING); Seletar, Gan, J.T.W.M., Hardie, J.A. & Khng, Y.W.K. 1012 (SINU); Seletar, 16 May

1992, Gan, J.T.W.M., Hardie, J.A. & Khng, Y.W.K. 1037 (SINU); Seletar, 2 Oct 1948, Sinclair,

J. SFN38248 (SING); Seletar, 18 Sep 1948, Sinclair, J. SFN38246 (E, NY, P, SING); Jurong

Road 15tb mile, 19 Oct 1932, Corner, E.J.H. SFN26034 (K, SING); Nee Soon Swamp Forest,

26 Jan 1995, Karim, N.A. et al. NK206 (SING); ibidem, 5 m, 19 Mar 1982, Maxwell, J.F. 82-

80 (SING); Chan Chu Kang, Jun 1892, Ridley, H.N. s.n. (MEL); Chan Chu Kang, Jan 1892,

Ridley, H.N s.n. (SING); ibidem, 4 Apr 1890, Goodenough, J.S. s.n. (SING); Krangi [Kranji],

7 Dec 1889, Ridley, H.N. s.n, (SING); ibidem, 1894, Mat s.n. (SING); Mandai Road, 19 Feb

1931, Spare, G.H. F905 (K); Bukit Mandai, Sep 1890, Goodenough, J.S. s.n. (SING).

Notes. This species is most similar to Aeschynanthus obconicus but the flowers, and

in particular the calyx, are smaller. The calyx is also green (vs red) and more saucer-

shaped than cup-shaped.

Further study necessary

Rafidah, A.R. & Nor Ezzawanis, A.T. FRL64274 (KEP) from G. Telapak Burok in

Berembun Forest Reserve, Negeri Sembilan, Malaysia, has small opposite leaves, 0.9-

2.1 X 0.6-1. 1 cm, and seeds with one long appendage at each end. This combination

of characters is otherwise unknown in Peninsular Malaysia and without flowers the

identity of this plant remains a mystery. Aeschynanthus longiflorus, which has the

same fruit and seed type, is known from the same locality but it is not known to

Aeschynanthus in Singapore and Peninsular Malaysia

59

have leaves anywhere near as small. FRI64274 is rather reminiscent of Aeschynanthus

fruticosus from Sumatra in leaf shape and seed type but that species always has most

leaves in whorls of three or more.

ACKNOWLEDGEMENTS: I thank Jana Leong-Skomickova (SfNG) for handling the review

process of this paper and, particularly, for taking many photographs and compiling the plates

for me. I thank Ali Ibrahim (SfNG), Preecha Karaket (BKF) and Saw Leng Guan (KEP) for

additional photos. I also thanlc Ruth Kiew (KEP) for her assistance in locating a number of

Malaysian place names; Martin Pullan (E) for both enabling the use of and assistance with the

Padme database; and the Singapore Botanic Gardens’ Herbarium curatorial staff for arranging

loans. Ruth Kiew (KEP) and Gemma Bramley (K) are thanked for their useful coimuents on

the manuscript.

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Clarke, C.B. (1 883). Cyrtandreae. In: De Candolle, A. & De Candolle, C. (eds) Monographiae

phanerogamanim, vol. 5. Pp. 1-303, 32 pi. Paris: Masson.

Denduangboripant, J., Mendum, M. & Cronk, Q.C.B. (2001). Evolution in Aeschynanthus

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J. Bot. 60: 299-304.

Mendum, M., Lassnig, P., Weber, A. & Christie, F. (2001). Testa and seed appendage

morphology in Aeschynanthus (Gesneriaceae): phytogeographical patterns and

taxonomic implications. Bot. J. Linn. Soc. 135: 195-213.

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Bot. 64: 363-429.

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Vietnam. Edinburgh J. Bot. 66: 391^46.

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(Gesneriaceae, Pp. 495-547)

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of Sumatra, Indonesia. Blumea 54: 278-279.

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Appendix 1. Index of Exsiccatae. The number in parentheses refers to the number of the taxon

in the text.

Abbe, E.C. et ah 9119 (11); Abbe, L.B. et al. 9767 (11); Abdul Samat bin Abdullah 68 (11);

Addison, G.H. s.n. (9); Allen, B.M. 4852A (10); Alvins, MJ^. 1942 (10); Anderson,

J.W. 95 (7), 156 (9); Annandale, N. & Robinson, H.C. s.n. {\Qi); Anonymous s.n. (14);

Anthony, S. SA2S0O>),SA231 (11),&4255 {9),SA481 {1),SA482 {9),SA639 {\A),SA740

( 1 0) , SA897 (9); Anthonysamy, S. SA1155 (9); Asmah 22 (9); Ave, W. 165 (8); Aziz Budin

48644 (8).

Barnes, E. s.n. (3); Batten Pooll, A.H. s.n. (11); Beccari, O. 36 (8); Best, G.A. 21255 (6);

Bogner 1699 (10); Brantley, G. & Neale, S. GB31 (11); Brantley, G. & Sam, Y.-Y. GB27

(9); Bremer, B. & Bremer. K, 1808 (9); Burkill, H.M. HMB730 (11), HMB760 (11),

HMB856 (9), HMB868 (3), HMB1996 (9), HMB2038 (9), HMB2665 (9), HMB2668

(14), HMB2811 (9), HMB2812 (7), HMB4221 (3), 8859 (7); Burkill, H.M. & Holttum,

R.E. s.n. (9), 7888 (7), 8418 (9), 8531 (11), 8549 (9); Burkill, H.M. & Md Haniff 12542

(6), 12798 {S), 12833 (9), 12971 (9), 15750{\), 17258 {9y, Burkill, H.M. etal. HMB2430

(9); Burkill, l.H. 2159 (10), 6102 (9); Burn-Murdoch s.n. (11); Burtt, B.L. B1739 (6);

Burn, B.L & Woods, P.J.B. B1644 (7), B1739 (6), B1643 (9).

Garrick, J. 607 (9); Carter s.n. (9); Castle-Smith, PM. 15 (9), 42 (11); Chan, K.Y. FRI49293

(9), FR164737 (9); Chan, Y.M. FRI49298 (8); Chan, Y.M. & Kiew, R. et al. FRI60504

(9); Chan, Y.M. etal. FR170603 (8); Chew, M.Y. & Rosdi, M. FRI55583 (9); Chew, M.Y.

et al. FR153648 (9); Chew, W.-L. CWL222 (14), CWL224 (9), CWL772 (11), CWL922

(11) , CWL1213 (8); Chin, S.C. 165 (11), 233 (11), 1441 (9), 1812 (9); Chin, S.C. et

ah 4533 (1); Choo, J.PS. et ah PYL2 (9); Chua, L.S.L FRI26700 (8), FR139082 (9),

FRI40785 (7); Chua, L.S.L. etal. FR145633 (9); Collenette, l.S. 2253 (9), 2254 (14);

Corner, E.J.H. s.n. (10), SFN26034 (14), s.n. (3); Corner, E.J.H. & Henderson, M.R.

SFN36611 (9), SFN36612 (9); Corporal s.n. (9); Cultivated C4877 (6), C7315 (11),

C7431 (9), C7650 (8); Cuming, H 2387 (10); Curtis, C. s.n. (11), 1311 (11), 1388 (8),

2142 (6), 2503 (9), 2990 (8), 3335 (8).

Darnaedi, D. et ah 566 (3); Davidson, C. 1292 (1); Davis 69248A (11), 69431 (9); Davison,

G.W.H. s.n. (10); Deny, R. 205 (10), 305 (10); Ding Hou 657 (8); Dransfield, J. s.n.

(11); Durnford, L. s.n. (9).

Ernst, A. 1189 (9); Evans, J.H.N. s.n. (9); Everard, B. & Young, D. 94 (9); Everett, B. FRI13553 (9).

Federated Malay States Museum Coll s.n. (11); Fielding, J. s.n. (9); Flippance, F. s.n. (13);

Fox, W. s.n. (11); Franck, C.W 209 (9), 1150 (10).

Aeschynanthus in Singapore and Peninsular Malaysia

61

Gadoh anak Umbai for A.H. Millard KL781 (10), KL1680 (10); Gadoh anak Umbai KL1319

(12); Gan, J.T.WM. et al. 1012 (14), 1037 (14); Gerb 3257 (9); Goodenough, J.S. s.n.

(9), 7575 (10), 7777 {9\2705 (9), 2706 (9); Griffith, W. s.n. (10); Gwee, A.T. SING2010-

443 (9); Gwynne- Vaughan, D.T. 410 {\0i).

Hardial, S. 537 (14); Hardial, S. & Samsuri Ahmad 18 (9), 290 (8); Haviland, G.D. s.n. (9);

Hawkins, A.S.M. s.n. (12), 6 (11); Heaslett, E.A. s.n. (1); Henderson, M.R. s.n. (2),

s.n. (11), s.n. (9), FMS11263 (9), 11365 (11), FMS11453 (9), 11713 (9), 11829 (11),

SFN17921 (9), 17988 (11), SFN21928 {\01SFN23534 (11), 25065 (9), SFN29643 (9),

SFN36656 (14); Heng, HP. et al. HPIO (6), HP 12 (9); Holttum, R.E. s.n. (7), 9489 (14),

9715 (10), 15312 (2), 20769 (11), 20772 (3), 21528 (7); Hons. Students 1 (9); Hose, G.

46 (11); Hume, HE. 8253 (10), 8943 (10), 9228 (10), 9596 (1), 9772 (1).

Ibrahim, H. & Lua, H.K. SING-2013-092 (9); Ibrahim, H, et al SING201 3-093 (9); Imin, K.

6 Asmarayani, H.S. FRI76004 (8); Imin, K. & Kueh, HE. et al FRI58561 (7); Imin,

K. & Syahida Emiza, S. FRI76065 (3), FRI76073 (3); Imin, K. et al. FRI58557 (9),

FRI58570 (11), FRI58593 (4), FRI58594 (10), FRI66402 (9), FRI66493 (9), FRI68188

(8) , FRI71615 (8), FR171855 (9), FRI71954 (1 1), FRI77536 (3), FR177847 (14).

Jaemat 25181 (9); Johnston, A.M.76 {\\ ); Jong, K. 901 7(1); Julius, A. FRI53313 ( 1 ), FRI54942

(9) , FR156020 (11 ), FRI57451 (8), FR173605 (7); Julius, A. & Imin, K. FRI56237 (10);

Julius, A. et al FRI53302 (11), FRI57479 (7), FRI57483 (11), FRI57691 (1), FRI57716

(9); Jumali K2077 (9); Jumali & Heaslett, E.A. KJ732 (9); Jutta, M. & Kueh, H.E.

FRI59582 (10), FRI59583 (9).

Kadim &. Noor; M. 131 (14), 135 (9), 138 (14); Kalong 22423 (9); Kamarudin S. FRI34629

(9), FRI33755 (11); Kamarid Hisham, M. FRI52054 (11); Kamarul Hisham, M. et al.

FRI52094 (3); Karim, N.A. et al NK206 (14); Kassim, M. 550 (9); KelsaU, H. s.n.

(12); Keng, H. & Jumali K2077 (9); Keng, H. et al K8010 (11), CTV86 (9), D-5 (11),

7 (10), 27 (9), 54 (9), 67 (9), 85 (7), 86 (9), 93 (9), 97 (9), 410 (11), 4755 (7); Khoo, R.

& Ng, S.M. 074 (9); Kiah SFN32054 (14), 23943 (8); Kiey^\ R. RK832 (9), RK1002 (9),

RK1081 (9), RK1546 {\\ ), RK1548 (9), RK2112 (14), RK2160 (9), RK2262 (8), RK2398

(14), RK2398 (14), RK2430 (11), RK2641 (9), RK2696 (14), RK2727 (9), RK3241 (3),

RK3264 (9), RK3293 (7), RK3356 (9), RK3775 (14), RK4862 (5), RK4869 (9), RK5239

(8), RK5307 (3); Kmw R. & Anthony, S. RK3414 (9); Kiew, R. et al FRI57497 (7);

King’s Collector 2012 (8), 2049 (8), 2849 (8), 3641 (8), 4738 (2), 7022 (7), 8314 (7),

10179 (8); Klackenberg, J. & Eundin, R. 689 (11); Kloss, C.B. s.n. (11), 6476 (10);

Kochummen, KM. FRI2654 (9), FRI16215 (7), FRI16427 (8), FRI16516 (9), FRI16796

(1), FRI29126 (8), FRI29332 (6); Kunstler, H. 2636 (9), 4463 (1).

Eake & Kelsall, H. s.n. (14); Eatiff, A. 347 (1); Eatijf, A. & Zainudin, A, AEM1033 (10),

AEM2764 (9), AEM3121 (11); Eatiff A. et al. AEM970 (11), AEM1644 (10), AEM1789

(7), AEM1813 (6); Eee, D.W. s.n. (11); Eeong, P S1NG201 0-809 (9); Eeong, P et al

SING2012-165 (9); Eewis, G.P 251 (9); Eirn, C.E. FRI73033 (9); Eim, C.E. & Kueh,

H.E. FRI64886 (8); Eo, Y.N & Mahmud 83 (9), 183 (5); Eobb, T. Ill (14); Eong, F.R.

13 (10), 14 (9); Eow, Y.W. EYW140 (11); Eua, H.K. & Ibrahim, H. SING20I5-084 (1).

Mahammud 17178 (9); Mahmood KEP17216 (9); Mahmud bin Sidek s.n. (9), 4816 (11);

Maingay, A.C. K.D.1219 (9), K.D.1218 (9); Mashall 35847 (10); Mat s.n. (14); Maxwell,

J.F 78-90 (9), 81-166 (9), 81-182 (14), 52-571 (14); Md Hanijf s.n. {U), 21074 (8),

21088 (7); Md Ha niff & Md Nur 2314 (9), 2347 (3), 2456 (11), 2469 (3), 8046 (8),

10390 (10); Md Nur s.n. (2), 11056 (11), 11138 (9), SFN24637 (10), SFN32980 (3),

SFN33963 (9), SFN34079 (9), SFN34254 (1), SFN34400 (9); Md Nur & Foxworthy,

FW. 11963 (1); Md Nur & Kiah 7783 (9); Mead, J.P 27945 (11); Mendurn, M. s.n.

62

Gard. Bull. Singapore 68(1) 2016

(9); Merton, L.F.H. 004194 (9); Mitchell s.n. (7); MK & AR 1321 (1); Mohd Hairul,

M.A. & Mohd Nazri, A. et al. FR169892 (11); Mohd Hairul, M.A. et al. FRI69938 (11),

FR169946 (7), FRI70936 (10), FR172324 (1); Mohd Shah MS 3446 (11); Mohd Shah &

MohdAli MS 3046 (9); Mohd Shah & SidekMS 1073 (11), MSI 110 (11); Mohd. Hairul,

M.A. FR160925 (8); Mohd. Kassim bin Rajab 488 (7); Morgany, T. et al. M260 (14);

Moysey, L. & Kiah SFN33624 (14); Mrs. Allen & Kadini 436 (14).

Napier, W. s.n. (12); Ng, F.S.P. FR11136 (9), FR15813 (8), FR15931 (11), FR127146 (11); Ng,

H.H & Lok, A.FS.L. s.n. (1); Nongchi s.n. (9), 8 (1); Noor, M. MN.1913 (9); Norain,

Raihana & Rosjana 07 (12).

Okada, H. et al. 1027 (11), 1050 (3).

Perumal, B. et al. FR141512 (7), FR141602 (11), FRI41631 (3), FRI41661 (11); Phoon, S.N.

FRI51987 (9), FRI60496 (11); Phoon, S.N. & Imin, K. FR160648 (7); Phoon, S.N. &

O’Byme, P. et al. FR160557 (11); Phoon, S.N. et al. FR151576 (9); Poore, M.E.D. 466

(11), 1028 (11), 1064 (8); Purseglove, J. W. P4202 (9), P4182 (9), P4164 (7), P4222 (3).

Quaife s.n. (7).

Rafidah, A.R. & Nor Ezzawanis, A.T FRI64274 (1); Rafidah, A.R. et al. FRI55631 (9);

Rahimatsah Amat N1 2 (9); Rao & et al. K8010 (11), 167 (9); Ridley, H.N. s.n. (1), 1578

(9) , 1700 (6), 2151 (1), 2167 (7), 2704 (10), 2706 (9), 2710 {9), 2905 (8), 5513 (9), 6244

(I) , 11447 {J\ 13372 75599(4), 13600{\\\ 13603 (7), 14063 (8), 14280(6), 14281

(10) , 14282 (10), 16090 (11), 16122 (3); Ridley, H.N. & Curtis, C. 7365 (3); Robinson,

H.C. s.n. (7); Robinson, H.C. & Kloss, C.B. 5997(9), 6049(9); Rosdi, M. & Phoon, S.N.

et al. FRI59867 (11), FR159872 (3); Rosdi, M. & Rafidah, A.R. FRI5983 7 (9); Rosdi, M.

et al. FR158753 (9), FR166272 (1), FR166336 (10); Rostado 64 (9).

Sabari, D. FRI32715 (1 1); Sam, Y.-Y. FRI44492 (10), FRI46549 (10), FRJ46602 (9), FRI47104

(10), FRI47144 (14), FRI47158 (14), FRI47208 (11); Sam, Y.-Y. et al. FRI47150 (9);

Samsuri Ahmad SA96 (11), 309 (9), S.311 (9), SA318 (14), SA1125 (9); Saw, EG.

FR134368 (11), FR136398 (11), FR137667 (1), FR137684 (10); Saw, LG. cfe Mohd

Haind, M.A. FRI48326 (1); Saw, L.G. & Mustafa, D. FRI37514 (9); Saw, L.G. et al.

FR148241 (\\);Scortechini, B. 36a(9), 46a(^), s.n. (11), 468b(]\), 55 (6), 57 (10), 59

(9) , 291 (9), 330 (10), 388 (11), 1815 (1); Seimund, E. s.n. (9), 390 (10), 459 (8); Shah,

M. 138 (14), 155 (1 ); Shah, M. & Mahmud MS4895 (9), MS4978 (9); Shah, M. & Noor,

M. MS607 (9), MS813 (14); Shah, M. & SidekMS1064 (9); Shah, M. et al. MS3407 (8);

Sidek bin Kiah S.277 (8), SK515 (8); Siew Wei Hoe 47 (W); Sinclair, J. s.n. (10), 5090

(10) , 5376 (9), 6198 (7), 8151 (10), 9949 (11), 10568 (9), SFN38246 (14), SFN38248

( 1 4), SFN38609 (9), SFN40358 (10); Sinclair, J. & Kiah SEN 38670 ( 1 1 ), SFN38800 (7),

SFN38805 (8); Smith, G. 443 (7), 448 (9), 513 (9); Smith, J. W. 63690 (3); Spare. G.H

F905 (14), 99/ (9), 2018 (9), 2047 (U ), 2 114 (7), 2139 (9), SFN36226 (8); Staples, G. et

al SING2009-157 (9); Stone, B.C.M. s.n. (8), 5889 (9), 6543 (9), 7194 (1 1), 7218 (11),

9566 (8), 10798 (9), 11091 (8), 12758 (5), 13764 (9), 14045 (11), 15071 (11), 15134

(9), 15167 (8), 15366 (1 1), 75797 (1 1); Stone, B.C.M. & Badaruddin 12068 (8); Stone,

B.C.M. & Mahmud 8500 (9); Sti'ugnell, E.J. 10515 (8); Syahida Emiza, S. FR151460

(II) , FRI57293 (11); Syahida Emiza, S. &Angan, A. FRI55108 (9); Syahida Emiza, S.

& Chew, M.Y.FR1572 77 (3); Symington, C.F 20938 (3), 21390 (8), 36073 (9), 51762

(11) , 56711 (11).

Tam Shell May TSM2 (11); Tan, W.K. et al. TWKl (11), TWK12 (9); Tay, E.P 125 (6); Teruya,

Z. 951 (1); Togashi, M. s.n. (11), 622111 (11).

UNESCO limestone expedition 440A (10), 239 (9); unknown FMS17068 (8), KEP78844 (9),

KEP93108 (12), s.n. (11), K6502 (9), s.n. (7), 4636 (9).

Aeschynanthus in Singapore and Peninsular Malaysia

63

Van Balgooy, MM.J. 2118 (8), 2154 (11), 2658 (3), 2663 (11), 7133 (11), 7149 (3), 7228 (11).

Vesterdal, A. 306 (14).

Wallich, N. 798 (14); Weber, A. s.n. (11); Whitmore, T.C. KEP207 (14), FRI227 (9), FRI3887

(11), FR112601 (7), FRI15450 (11), FRI15495 (11), FR120392 (8); Wilkie, R et al

FR152906 (11), FR175005 (11); Wong, KM. FRI35244 (11), s.n. (14); Wong, Y.K.

KEP93272 (8); Woods, P.J.B. 616 (3), 632 (11), 634 (3), 680 (11); Woods, P.J.B. et al.

615 (11); Worthington, R.D. 12463 (11), 13326 (9); Wray, L. s.n. ^1), 149 (8), 625 (7),

655 (9), 656(11), 873 (9), 1636 (ii), 1772 (S), 3217 (9), 4245 (1); Wray, L. &Robinson,

H.C. 5483 (3); Wy’att-Smith, J. KEP 56945 (11), 78808 (11).

Yao, T.L.FR165302(3),FR157961 (6); Yao, T.L. etal. FRI5 5845 {\\),FRI5 5898 {\\),FRI579 13

(10), FRI65386 (14), FRI65387 (9), FRI65495 (7), FRI77328 (9), FRI77335 (7); Yapp,

R.H. 161 (1), 440 (11), 547 (8), 610 (12); Year III students FSC312 (9); Yeo, C.K. s.n.

(9); Yong KEP99330 (9).

Zainal Mustafa ZM64 (9); Zainon, A.S. FRI50105 (14); Zainudin, A. AZ40 (9); Zainudin, A. et

al. AZ2438 (9).

Gardens’ Bulletin Singapore 68(1): 65-69. 2016

doi: 10.3850/S2382581216000028

65

Notes on the Annonaceae of the Malay Peninsula

LM. Turner

Research Associate, Royal Botanic Gardens Kew,

Richmond, Surrey TW9 3AE, U.K.

j_trop_ecol@yahoo.co.uk

ABSTRACT. Notes concerning the three genera Alphonsea Hook.f. & Thomson, Artabotrys

R.Br. ex Kew Gawl. and Stelechocarpus Hook.f. & Thomson (Annonaceae) for the Malay

Peninsula are presented. Alphonsea kingii J.Sinclair is accepted as an endemic species and a

second-stage lectotypification is presented for Alphonsea cylindrica King. Seven species of

Artabottys are lectotypified. The recently described genus Winitia Chaowasku is reduced to

Stelechocarpus and a new combination is made for Winitia expansa Chaowasku.

Keywords. Alphonsea, Artabotrys, new combination, Stelechocarpus, typification, Winitia

Alphonsea

Alphonsea kingii

Sinclair (1955) described Alphonsea kingii J.Sinclair as a new species in his account

of the Annonaceae of the Malay Peninsula. It had earlier been confused with a species

of Xylopia L. by King (1892, 1 893). Kessler (1995), in revising the genus Alphonsea

Hook.f. & Thomson, excluded A. kingii, suggesting that it belonged in Mitrephora

Hook.f. & Thomson. Kessler did not report seeing any specimens; his decision was

apparently based on King’s plate (King 1893: t. 189 A). Tn studying the genus in

Peninsular Malaysia it became necessary to reconsider Alphonsea kingii. I was kindly

sent photographs of the type material from the Central National Herbarium of the

Botanical Survey of India (CAL). The second specimen cited by Sinclair (1955), a

Scortechini collection from Perak, could not be located in SING (D.J. Middleton

pers. comm.). However among material on loan from the herbarium of the Forest

Research Institute of Malaysia (KEP), I found another specimen of the species, which

had previously been detemiined as Alphonsea elliptica Hook.f. & Thomson. Study of

the images and specimen gave no indication that Sinclair was wrong in his assignment

of the species to Alphonsea. Flowers are still unknown, but in vegetative and fruit

characters the specimens are congruent Wiih Alphonsea, with leaves similar to those of

A. malayana P.J.A.Kessler, A. johorensis J.Sinclair and A. elliptica. Notable features

of the species include the rather dense reticulations of the lamina venation, and

particularly the verrucose fruits that dry blackish. It seems to be a species of limestone

outcrops - a habitat affinity not previously reported in MdildydiW Alphonsea. I conclude

that Alphonsea kingii is a good species that should be listed in the Annonaceae flora of

Peninsular Malaysia.

66

Gard. Bull. Singapore 68(1) 2016

Alphonsea kingii J. Sinclair, Gard. Bull. Singapore 14: 386 (1955). - TYPE: Peninsular

Malaysia, Perak, Kinta, top of limestone hills near G.M. [possibly Gunung Mesua, R.

Kiew pers. comm,], January 1885, King’s Collector [Kunstler, H.H.] 7097 (lectotype

CAL [CAL0000025047], designated here; isolectotypes CAL [CAL000025046], DD,

K (reported by Sinclair (1955: 386) but not seen lately)).

Additional specimen studied. PENINSULAR MALAYSIA: Pahang: Raub, Bukit Serdam, 20

Jun 1971, Chin, S.C. 1073 (KEP [138213]).

A second-stage lectotj pification

Kessler (1995: 88) reported the holotype of Alphonsea cylindrica King to be located

in the Central Herbarium of the Botanical Survey of India (CAL), without seeing the

specimen. As there are duplicates of this collection in other herbaria, Kessler (1995)

effectively lectotypified the name to this specimen. However, enquiries addressed to

the Botanical Survey of India were very kindly answered and I was informed that

there were two specimens under the type number in CAL. Therefore, here I propose

a second-stage typification to restrict Kessler’s choice of lectotype to the better of the

two sheets in CAL.

Alphonsea cylindrica King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 61(2): 127 (1892). -

TYPE: Peninsular Malaysia, Perak, Ulu Bubong, July 1 886, King 's Collector [Kunstler,

H.H.] 10633 (lectotype CAL [CAL0000004699], designated by Kessler (1995) at the

first stage and here at the second stage; isolectotypes A, BM [x2 mounted on same

sheet], BO, CAL [CAL0000004700], DD, G, K [K000574904], L, WU).

Artabotrys

A number of names among Artabotrys species from Peninsular Malaysia require

lecto typification. These are dealt with here.

Artabotrys crassifolius Hook.f & Thomson, El. Brit. Ind. 1: 54 (1872). - TYPE:

Peninsular Malaysia, Malacca, Griffith, W. s.n. [HEIC (Herbarium of the East India

Company) no. 426] (lectotype K, designated here).

Remaining syntype, Bunua, Martaban, Brandis, D. (n.v.).

Artabotrys lowianiis Scort. ex King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 61(2):

34 (1892). - TYPE: Peninsular Malaysia, Perak, Scortechini, B. 2012 (lectotype

K [K000381019], designated here; isolectotypes BM [BM000898096], CAL

[CAL0000004298, CAL0000004297, CAL0000004299], SING [SING009624I]).

Artabotrys oblongiis King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 61(2): 33 (1892). -

TYPE: Peninsular Malaysia, Perak, Lamt, August 1884, King’s Collector [Kunstler,

H.H.] 6524 (lectotype K [K000381015], designated here; isolectotypes BM

[BM000898111], CAL [CAL0000004314, CAL0000004313]).

Notes on Malay Peninsula Annonaceae

67

Artabotrys oxycarpiis King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 61(2): 34 (1892). —

TYPE: Peninsular Malaysia, Perak, November 1 883, Kings Collector 5150 (lectotype

K [K000381016], designated here; isolectotypes BM [BM000898086], CAL, DD).

Remaining syntypes. Peninsular Malaysia, Larut, February 1884, King’s Collector

5605 (BM [BM000898087], CAL, K [K000381017]); s. loc., Wray L 328b (CAL, K

[K000381018], SING [STNG00096242]).

Artabotrys pleurocarpus Maingay ex Hook.f. & Thomson, FI. Brit. India 1 : 54 (1872).

- TYPE: Peninsular Malaysia, Malacca, 6 February 1868, Maingay, A.C. 3261 [Kew

distrib. no. 34] (lectotype K [K000381010], designated here explicitly excluding

material in attached packet which represents another gathering).

Remaining syntypes. Peninsular Malaysia, Malacca, 14 April 1868, Maingay, A.C.

3261A [Kew distrib. no. 34] (K [K000381009]).

Artabotrys scortechinU King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 61(2): 32 (1892).

- TYPE: Peninsular Malaysia, Perak, Scortechini, B. s.n. (lectotype K [K000381008],

designated here).

Remaining syntypes. Peninsular Malaysia, Perak, Scortechini, B. 488 (BM

[BM000898101], CAL [CAL0000004270, CAL0000004271, CAL0000004278],

SING [SING0096243]).

Artabotrys wrayi King, J. Asiat, Soc. Bengal, Pt. 2, Nat. Hist. 61(2): 37 (1892). -

TYPE: Peninsular Malaysia, Perak, Larut, November 1882, King’s Collector 3615

(lectotype K [K000381023], designated here; isolectotypes BM, CAL).

Remaining syntypes. Peninsular Malaysia, Perak, Wray, L. 4006 (SING

[SING0096239]); ibidem, Wray, L. 2663 (SING [SING0096240]).

Stelechocarpusl Winitia

In a recent analysis of the genera Stelechocarpus Hook.f. & Thomson and Sager aea

Dalzell based on morphological and molecular data, Chaowasku & Van der Ham

(201 3 ') demonstrated the presence of three groups. These corresponded to two strongly

supported clades; one consisting of the Sageraea species sampled, and the other to

Stelechocarpus cauliflorus (Schefif.) R.E.Fr, plus another species. Stelechocarpus

burahol (Blume) Hook.f. & Thomson, the type species of Stelechocarpus, formed the

third, less strongly supported clade as sister to the S. cauliforus clade. As Chaowasku &

Van der Ham noted, there were three taxonomic options for reflecting this phylogenetic

relationship. Firstly all the species could be included in an enlarged Sageraea, secondly

the status quo could be maintained (i.e. the two genera Sageraea and Stelechocarpus),

' A corrigendum published later (Syst. Biodivers. 11(4): 537 (2013)) added L.W. Chatrou to the list of

authors of the paper, but to avoid eonfusion the paper is referred to as first published.

68

Gard. Bull. Singapore 68(1) 2016

or finally the three main clades could be given generic rank. Chaowasku & Van der

Ham favoured the last option and described the genus Winitia Chaowasku to include

Winitia cauliflom (Scheff.) Chaowasku transfeired from Stelechocarpus and the

new species from Peninsular Thailand, Winitia expansa. This leaves Stelechocarpus

burahol on its own in Stelechocarpus.

I do not have any criticisms to make of the phylogenetic analysis but I do think

the recognition of Winitia is a backward step for taxonomy. As a family, the Annonaceae

can be characterised by its large number of genera. Some 1 09 are cuiTently recognised,

with 42 in the Asia-Pacific region alone (Couvreur et al., 2012). This is problematic for

non-specialists. Increasing the number of recognised genera is not helpful and single-

species genera (such as Stelechocarpus sensu Chaowasku & Van der Ham) have low

information content. It seems to me that maintaining the status quo in tenns of genera

is a better option than splitting Stelechocarpus. The phylogenetic relationship could

be recognised by infrageneric taxa. The added advantage of this arrangement is that

Stelechocarpus is easy to recognise - for instance, the raised midrib on the upper

surface of the leaves is a reliable vegetative character (Sinclair, 1955; Van Heusden,

1995) with few confusable taxa. Stelechocarpus s.s. and Winitia can be separated on

various characters, including flower colour, relative size and spatial distribution of male

and female flowers, stamen number, stigma fomi and pollen moi*phology (Chaowasku

& Van der Ham, 2013), but this ignores the similarities including the raised midrib,

monoecy, convex male torus and general gestalt. The recognition of Winitia, therefore,

represents the splitting of a well-defined and distinctive genus into two less easily

distinguished entities. The two Stelechocarpus clades could readily be considered as

infrageneric taxa, perhaps subgenera, but 1 refrain from doing so here.

In order to allow the maintenance of Stelechocarpus in its broader sense, I make

a new combination for Winitia expansa in Stelechocarpus.

Stelechocarpus expansus (Chaowasku) I.M. Turner, comb. nov. - Winitia expansa

Chaowasku in Chaowasku & Van der Ham, Syst. Biodivers. 11: 203 (2013).

ACKNOWLEDGEMENTS. I thank Dr T.K. Paul (CAL) for providing images and information

on specimens. David Johnson and an anonymous reviewer provided constructive criticisms

that helped to improve the paper.

References

Chaowasku, T. & Van der Ham, R.W.J.M. (2013). Integrative systematics supports the estab-

lishment of Winitia, a new genus of Aimonaceae (Mahneoideae, Miliuseae) allied to

Stelechocarpus and Sageraea. Syst. Biodivers. 11(2): 195—207.

Couvreur, T.L.P., Maas, P.J.M., Meinke, S., Johnson, D.M. & Kessler, P.J.A. (2012). Keys to

the genera of Annonaceae. Bot. J. Linn. Soc. 169: 74-83.

Kessler, P.J.A. (1995). Studies on the tribe Saccopetaleae (Annonaceae), IV. Revision of the

gQnus Alphonsea Hook.f. & Thomson. Bot. Jahrb. Syst. 118: 81-112.

Notes on Malay Peninsula Annonaceae

69

King, G. (1892). Materials for a flora of the Malayan Peninsula, no. 4. J. Asiat. Soc. Bengal,

Pt. 2, Nat. Hist. 61: 1-130.

King, G. (1893). The Annonaceae of British India. Ann. Roy. Bot. Gard. Calcutta 4: 1-169.

Sinclair, J. (1955). A revision of the Malayan Annonaceae. Gard. Bull. Singapore 14: 149-516.

Van Heusden, E.C.H. (1995). Revision of the Southeast Asian genus Stelechocarpus

(Annonaceae). Blumea 40: 429^38.

Gardens’ Bulletin Singapore 68(1): 71-76. 2016

doi: 10.3850/S238258121600003X

71

Novitates Bruneienses, 5. Polyalthia watui (Annonaceae),

a new tree species from Brunei, Borneo

K.M. Wong^, A.K. Muhammad Ariffitf & A.A. Joffre^

^Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, Singapore 259569

wkin2000@.ginail .com

^Brunei National Herbarium, Forest^ Department,

Ministry of Primary Resources and Tourism, Jalan Menteri Besar,

Berakas, BB3910 Brunei Darussalam

ABSTRACT. A new species, Polyalthia watui, is described. It resembles Polyalthia bullata

King in having bullate leaves with a cordate base, but is distinguished by its flowers with

shorter pedicels, longer sepals and petals, subsessile monocarps with long stiff erect hairs; and

broad-obovate leaves with fewer pairs of secondary veins. The new species is only known from

Brunei’s Belait and Tutong districts and is very likely endemic to the Belait geosyncline that

includes these areas.

Keywords. Annonaceae, biodiversity, Brunei, endemic species, Polyalthia, taxonomy

Introduction

The Annonaceae are a predominantly pantropical lowland forest family of trees,

shrubs and lianas, estimated to include some 2440 species (Chatrou et ah, 2012) in

122 genera (Rainer & Chatrou, 2014). Polyalthia Blume, a common Indo-Malesian

genus in this family, was, until recently, a polyphyletic genus with some 170 accepted

species, harbouring a morphologically heterogeneous assemblage of species. The

reorganisation of the genus, coirelating with molecular phylogenetic analyses, now

recognises several distinct genera, including Maasia (Mols et al., 2008) Fenerivia

Diels (Saunders et al., 2011), Monoon Miq. (Xue et al., 2012), Huberantha Chaowasku

(Chaowasku et al., 2015), and Polyalthia s.s., as well as species that fall within

Marsypopetalum Scheff. (Xue et al., 2011).

The taxonomically pruned version oi Polyalthia is estimated to include some 80

species (Turner et al., 2014), with c. 30 species in Borneo (Turner, 2009, 2010; Turner

et al., 2014). Various features in combination serve to distinguish the genus from

others in Borneo: tree habit; hairs simple when present; non-glaucous leaves with non-

prominent midrib on leaf upper surface and brochidodromous venation (with looping

secondary veins); valvate perianth whorls with 3 sepals; apically flat and non-saccate

petals in two similar whorls; non-veiTucose non-clawed inner petals that are not tightly

appressed over the reproductive parts; more than 3 carpels with 2-6 oxailes per carpel;

and commonly 1-2 seeds per carpel, each seed with a shallow circumferential groove

(Turner et al., 2014).

72

Gard. Bull. Singapore 68(1) 2016

In recent years, a distinctive species of Polyalthia with strongly bullate leaves has

been collected in Brunei’s Belait and Tiitong districts, and during recent fieldwork there

in 201 5 under the auspices of the Botanical Surrey of Brunei Darussalam progi'amme

organised by the Brunei Forestry Department and Singapore’s National Parks Board

(see Joffi'e et ah, 2015), it was possible to further study this species in the field (Fig.

1). This unnamed species resembles several other species in individual characters but

in the combination of characters it possesses it most resembles, and would appear to

be most closely related to, Polyalthia bullata King (King, 1892; Sinclair, 1955; Turner

et ah, 2014). It also has similarities to Polyalthia endertii D.M. Johnson (Johnson &

Murray, 1999). The new species is, however, clearly distinguished by characteristics

of the flowers, fruits and leaves (Table 1). We are pleased to name this after Watu

Awok of the Brunei National Herbarium’s field team for his enthusiastic support and

excellent company during field suiweys, and for diligently searching out individuals of

this extraordinary Polyalthia in the field.

Herbarium acronyms used follow Tliiers (continuously updated). Conservation

assessments follow the methodology of lUCN (2012). Dimensions in the description

are for dried material. As shrinkage when dried can be quite pronounced, the dimensions

from spirit material are also given for some flower parts.

Polyalthia watui K.M.Wong, sp. nov.

The new species resembles Polyalthia bullata King in having bullate leaves with a

cordate base, but differs in its flowers with shorter (8-9 mm long) pedicels, 13-14 mm

long ovate sepals, 24-31 mm long and 5-9 mm wide oblanceolate petals; monocarps

with only 1-2 mm long stipes and stiff erect hairs mostly 3-5 mm long; and broad-

obovate leaves with 16-18 pairs of secondary veins. In contrast, Polyalthia bullata has

flowers with 10-25 mm long pedicels, 5-7 mm long lanceolate sepals, 21-25 mm long

and 2-2.5 mm wide linear petals; monocarps with c. 5 mm long stipes and short hairs

less than 1 mm long; and nan*owly lanceolate to oblong-lanceolate leaves with 25-40

pairs of secondary veins. - TYPE; Brunei, Tutong, Lamunin, Bukit Sulang, Lamunin

Forestry office, behind ex Plantation Nursery, 17 Febmary 2005, flowers, Muhammad

Ariffin BRUN 21189 (holotype BRUN; isotypes K, SAN, SING). (Fig. 1, Table 1)

Treelet 1-4 m high; stem axes proximally vertical, distally curving over to a near-

horizontal orientation, the older trunk developing from a sympodium of such proximal

portions and attaining 1-3.5 cm diameter at the base; leafy branches near-horizontal.

Young twigs densely covered with medium brown suberect 2-3 mm long hairs. Leaves

distichously arranged throughout, broad-obovate, 20-25 cm long, 9-12 cm wide, apex

acute to obtuse or rounded, base markedly cordate, auriculate, the basiscopic auricle

conspicuously larger than the acroscopic auricle, the auricles overlapping the twig,

chartaceous to thin-coriaceous, bullate, above glabrous except for the densely brown

short-hairy midrib, below with dense brown suberect 1.5-2 mm long hairs on the

midrib and sparse brown suberect 0.5-1 mm long hairs on the secondary veins, midrib

sunken above and prominent below, secondary veins 16-18 pairs, fonning bold loops

near the margin, sunken above and prominent below, tertiary and higher-order venation

subscalariform to reticulate, sunken above and prominent below, medium green and

New Polyalthia from Brunei

73

Fig. 1. Polyalthia watui K.M.Wong. A. Mature treelet displayed by Watu Awok of the Brunei

National Herbarium. B. Strongly bullate leaf supper surfaces. C. A flower from the type

specimen BRUN 21189 showing sepals (only one out of 3 indicated, s), 3 outer petals (op) and

3 inner petals (ip). D. Monocarps covered in white erect long hairs and basally subtended by

the 3 persistent sepals. (Photos: A, B, D: K.M. Wong; C: Muhammad Ariffin)

slightly reflective above, pale grey-green and matt beneath, subsessile, petioles 4-8

mm long, stout. Flowers solitary, axillary to extra-axillary, deflexed to a pendulous

position, pedicels 8-9 mm long (to 12 mm long in spirit material). Sepals 3, broad-

ovate, apex acute, 13-14 mm long, 5-6 mm wide (29 mm long, 15 mm wide in spirit

74

Gard. Bull. Singapore 68(1) 2016

Table 1. Comparison of Polyalthia bullata King, P. endertii D.M. Johnson and the new speeies.

(Measurements are from dried herbarium material.)

Character

Polyalthia bullata

Polyalthia endertii

Polyalthia watui

Flower pedicels

10-25 mm long

c. 20 mm long

8-9 mm long

Flower sepals:

shape, size

Lanceolate, 5-7 mm

long, 1.5-2 mm

wide

Lanceolate-ovate,

20-21 mm long, 11 mm

wide

Ovate, 13-14 mm

long, 5-6 mm wide

Petals: shape, size

Linear, 21-25 mm

long, 2-2.5 mm

wide

Oblanceolate, c. 27 mm

long, 7 mm wide

Oblanceolate to strap-

shaped, 24-31 mm

long, 5-9 mm wide

Persistent sepals of

fruit

5-7 imn long, 3^

mm wide

(Unknown)

20-29 mm long,

10-15 mm wide

Monocarp stipes

c. 5 mm long

(Unknown)

1-2 mm long

Monocarp surface

Puberulous, hairs

less than 1 mm long

(Unknown)

Covered in stiff erect

hairs 3-5 mm long

Leaf shape

Narrowly lanceolate

to oblong-lanceolate

Oblanceolate, slightly

panduriform

Broad obovate

Leaf dunensions

25-50 X 0.5-12 cm

16.8-32 X 4.8-8 cm

20-25 X 9-12 cm

Leaf lateral veins

25^0 pairs

20-27 pairs

16-18 pairs

Leaf surface

Conspicuously

bullate

Generally plane, not

bullate

Conspicuously

bullate

material), coriaceous, sparsely pubescent on both sides, creamy yellow. Petals in 2

series of 3, oblanceolate to strap-like, apices acute, coriaceous, outer petals 29-31

mm long, 8-9 imn wide (45 mm long, 10 mm wide in spirit material), with 3-5 bold

longitudinal ridges on the imier surface, inner petals 24-26 mm long, 5-6 mm wide

(42 mm long, 6 mm wide in spirit material), with 1-3 bold longitudinal ridges on

the inner surface, all densely covered with pale hairs mostly less than 1 mm long on

both surfaces, creamy yellow in live materiah Stamens numerous, connectives slightly

convex, apices of anther connectives bright yellow (measurements not taken). Carpels

15 seen in a single flower, elliptic-subglobose, densely pubescent, stigma sub-capitate,

puberulous. Monocarps subglobose, 5-7 mm diameter (10-15 mm diameter when

fresh), pale green to yellowish green, densely covered with stiffly erect pale 3-5 mm

long hairs, 2-12 together (very rarely solitary) embraced by the 3 persistent ovate

sepals 20-29 mm long, 10-15 mm wide (25-30 mm long, 20-25 mm wide when

fresh); stipes 1-2 mm long. Seeds 2, plano-convex, with a shallow circumferential

groove near the edge of the plane surface; endospenu glassy, ruminate by thin laminar

intrusions of the seed coat.

New Polyalthia from Brunei

75

Additional specimens examined. BRUNEI: Belait: Labi, Labi Hills Forest Reserve,

Compartment 49, Ulu Sg. Rampayoh, 20 May 2009, fruits, Azlan BRUN 22629 (BRUN, K,

SAN, SENG); Tutong: Rambai, Ladan Hills Forest Reserve, Benutan Dam catchment forest,

tributary of Sungai Benutan, 7 Jun 2015, fruits, Wong & Watu WKM 3425 (BRUN, SING),

Ladan Hills Forest Reserve, Nyamokning West, southeast of LP 230, 4”24.45’N 114‘’48.36’E,

35 m asl, riverine mixed dipterocarp forest, 6 Jun 1996, fruits, Joffre BRUN 17529 (BRUN).

Distribution. So far known only from Brunei, in the Belait and Tutong districts, in

small populations. As discussed by Joffre et al. (2015), the Belait syncline covering

both these districts is a geo-ecologically well-defined enclave that is likely to harbour

its own endemic taxa. Given that the adjacent parts of Sarawak and nearby southwest

Sabah have been generally well-collected and this species has not been documented

in those tenitories, it is not likely to have a wider distribution than in these districts.

The related Polyalthia bullata is known from Peninsular Malaysia and all territories in

Borneo, including in adjacent Sarawak as well as Brunei (Wong WKM 1710 (BRUN,

K, SING)).

Habitat. Understorey of mixed dipterocarp forest on sandy clays and alluvium criss-

crossed by shallow streams feeding tributaries of the Belait and Tutong rivers. The

more widely distributed Polyalthia bullata also occurs in lowland forest but on

hillsides and river terraces that are not as low-lying and moist as the habitat of watui.

Provisional lUCN conservation assessment. Polyalthia watui is so far documented

only from undisturbed lowland primary forest in the Labi Hills Forest Reserve, Belait

district, and several pockets in the Ladan Hills Forest Reserve, Tutong distiict. The

lUCN status proposed here is Least Concern (LC) as the species is protected in forest

reserves with no imminent threats. The Brunei Govenmient officially announced in

2014 that logging will be phased out in all forest reserves (Brunei Times, 2014).

ACKNOWLEDGEMENTS. Our botanical surveys in Bmnei Darussalam were sponsored by

the Brunei Forestry Department of the Ministry of Prhnary Resources and Tourism, and the

National Parks Board, Singapore through the Singapore Botanic Gardens. Y.W. Low assisted in

the composition of Fig. 1. Dr David M. Johnson (Ohio Wesleyan University) and an anonymous

reviewer provided useful comments.

References

Bmnei Times (2014). Logging no longer allowed in forest reserves: Ministry of Industry

and Primary Resources. 23 March 2014. http://bt.com.bn/news-national/2014/03/23/

logging-no-longer-allowed-forest-reserves-mipr (accessed on 13 Feb. 2015).

Chaowasku, T, Johnson, D.M., Van der Ham, R.W.J.M. & Chatrou, L.W (2015). Huberantha,

a replacement name for Hubera (Annonaceae: Malmeoideae: Miliuseae). Kew Bull.

70: 23.

76

Gard. Bull. Singapore 68(1) 2016

Chatrou, L.W., Pirie, M.D., Erkens, R.HJ., Couvreiir, T.L.R, Neubig, K.M., Abbott, R.J.,

Mols, J.B., Maas, J.W., Saunders, R.M.K. & Chase, M.W. (2012). A new higher-

level classification of the pantropical floweiing plant family Annonaceae informed by

molecular phylogenetics. Bot. J. Linn. Soc, 1 69: 5^0.

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

and UK, Cambridge: lUCN.

Joffre, A. A., Ariffin, M.A.K., Low, Y.W. & Wong, K.M. (2015). Novitates Bmneienses, 1.

A background to the botanical survey of Brunei Damssalam, and a new species of

Jarandersonia (Malvaceae). Card. Bull. Singapore 61 {\)\ 51-60.

Johnson, D.M. & Muinay, N.A. (1999). Four new species of Polyalthia (Annonaceae) from

Borneo and their relationship to Polyalthia insignis. Contr Univ. Michigan Herb. 22:

95-104.

King, G. (1892) Materials for a Flora of the Malay Peninsula. No. 4. J. Asiat. Soc. Bengal, Pt.

2, Nat. Hist. 61(1): 1-130.

Mols, J.B., Kessler, P.J.A., Rogstad, S.H. & Saunders, R.M.K. (2008). Reassignment of six

Polyalthia species to the new genus Maasia (Annonaceae): molecular and morphological

congruence. Syst. Bot. 33: 490-494.

Rainer, H. & Chatrou, L.W. (2014). AnnonBase: world species list of Annonaceae, ver. Jan.

2014. http://www.sp2000.org and http://www.annonaceae.org (accessed on 14 Oct.

2015).

Saunders, R.M.K., Su, Y.C.F. & Xue, B. (2011). Phylogenetic affinities of Polyalthia species

(Annonaceae) with columellar-sulcate pollen: Enlarging the Madagascan endemic

goxwxs Fenerivia. Taxon 60\ 1407-1416.

Sinclair, J. (1955). A revision of the Malayan Annonaceae. Gard. Bull, Singapore 14: 149-518.

Thiers, B. (continuously updated). Index Herbariorum: A global directoiy of public herbaria

and associated staff. New York Botanical Garden’s Virtual Herbarium. http://sweetgum.

nybg.org/ih/ (accessed on 30 Jul. 2015).

Turner, I.M. (2009). New species and nomenclatural combinations m Polyalthia, Meiogyne and

Mitrella (Annonaceae) from Borneo. Malayan Nat. J. 61: 267-276.

Turner, I.M. (20 1 0). New species of Polyalthia (Annonaceae) from Borneo and their relationship

to Polyalthia cauliflora. Nordic J. Bot. 28: 267-279.

Turner, I.M., Weerasooriya, A.D., Saunders, R.M.K. & Ganesan, S.K. (2014). Annonaceae.

In: Soepadmo, E., Saw, L.G., Chung, R.C.K. & Kiew, R. (eds) Tree Flora of Sabah

and Sarawak, vol. 8. Kepong, Malaysia: Sabah Forestry Department, Forest Research

Institute Malaysia & Sarawak Forestry Department.

Xue,B., Su, Y.C.F., Mols, J.B., Kessler, P.J. A. & Saunders, R.M.K. (2011). Further fragmentation

of the polyphyletic genus Polyalthia (Annonaceae): molecular phylogenetic support for

a broader delimitation of Marsypopetalurn. Syst. Biodivers. 9: 17-26.

Xue, B,, Su, Y.C.F., Thomas, D.C. & Saunders, R.M.K. (2012). Pruning the polyphyletic genus

Polyalthia (Annonaceae) and resurrecting the genus Monoon. Taxon 61: 1021-1039.

Gardens’ Bulletin Singapore 68(1): 77-86. 2016

doi: 10.3850/S2382581216000041

77

Three new species of Begonia (Begoniaceae)

from Sumbawa Island, Indonesia

D. Girmansyah

Herbarium Bogoriense, Research Center for Biology,

Indonesian Institute of Sciences, Jl. Raya Jakarta-Bogor Km 46,

Cibinong, 16911, Bogor, West Java, Indonesia

deden_bo@yahoo .com

ABSTRACT. Three new species of Begonia L., Begonia sumbawaensis Girm., Begonia

brangbosangensis Girm. and Begonia jaranpusangensis Girm., are described from Sumbawa,

Indonesia. All species are illustrated and identification keys and distribution maps are provided.

Keywords. Begonia sect. Petermannia, Begonia sect. Reichenheimia, Betunin, endemic

Introduction

Begonia L. (Linnaeus, 1753), is currently estimated to have around 1803 species

distributed throughout tropical and subtropical Asia, Africa and America (Hughes

et al., 2015b). In Southeast Asia, Indonesia is a centre of diversity for Begonia and

many new species are still being discovered (Ginnansyah & Susanti, 2015; Hughes

et al., 2015a; Undaharta et ah, 2015). CuiTently, 231 accepted species of Begonia

are known from hidonesia, distributed from Sumatra to Papua. In Sumbawa (West

Nusa Tenggara), only three species have been recorded to date {Begonia bimaensis

Undaharta & Ardaka (Undaharta et ah, 2015), B. multangula Blume (Hughes et ah,

2015b) and B. muricata Blume (Hughes, 2008). Some specimens previously attributed

to Begonia muricata are included below in the new species B. sumbawaensis.

Sumbawa is situated between Lombok to the west and Flores to the east.

Botanical exploratory efforts on Sumbawa are summarised in Van Steenis-Kruseman

(1950). Recent botanical explorations in March 2004 and July 2005 conducted by

Herbarium Bogoriense (RCB-LIPI) and sponsored by the New England Tropical

Conservatory, USA resulted in three new species of Begonia being collected. The new

species were collected from the Mt Batu Pasak complex, the Mt Ngengas complex and

the Jaran Pusang complex (Fig.l).

Three new species. Begonia sumbawaensis Ginn., Begonia brangbosangensis

Ginu. dindBegonia jaranpusangensis Girm., are described here. Begonia sumbawaensis

is placed in Begonia sect. Reichenheimia (Klotzsch) A.DC. (De Candolle, 1859)

as it exhibits the characters typical of the section: rhizomatous stems, protandrous

inflorescences, and three locular ovaries with entire placentae (Doorenbos et ah,

1998). Begonia jaranpusangensis and B. brangbosangensis are placed in Begonia

sect. Petermannia (Klotzsch) A.DC. (De Candolle, 1859) as they exhibit characters

78

Gard. Bull. Singapore 68(1) 2016

O

00

o

b

116WE 118WE

116 WE 118"6'0"E

Fig 1. Distribution of Begonia sumbawaensis Girm. (triangles), B. brangbosangensis Girm.

(circles) and B. jaranpusangensis Girm. (square).

typical of the section: upright stems, protogynous inflorescences, and three locular

ovaries with branched placentas (Doorenbos et ah, 1998).

Key to the species of Begonia from Sumbawa

la. Rhizomatous or tuberous herb; leaves ovate to broadly ovate 2

lb. Erect herb; leaves ovate to oblong 3

2a. Base tuberous; leaves lobed, upper surface with long hairs B. bimaensis

2b. Base rhizomatous; leaves entire, upper surface glabrescent B. sumbawaensis

3a. Leaves broadly ovate or orbicular, margin shallowly lobed; fruit fleshy and

indehiscent B. multangula

3b. Leaves elliptic, oblong, ovate or narrowly ovate, margin entire; fruit dry and

dehiscent 4

Three new Begonia species from Indonesia

79

4a. Leaves ovate to elliptic; petiole hairy B. jaranpusangensis

4b. Leaves narrowly ovate to oblong; petiole glabrous B. brangbosangensis

Begonia sumbawaensis Girm., sp. nov. § Reichenheimia

Similar to Begonia lugrae Ardhaka & Undaharta ( Ardi et al., 20 1 3) but differs in having

tomentose petioles (not glabrous), stipule narrowly triangular (semicircular in Begonia

lugrae), female flower tepals four (three in Begonia lugrae) and stamens 48 (90 in

Begonia lugrae), - TYPE: Indonesia, West Nusa Tenggara, West Sumbawa, Betunin

Hill, 10 July 2005, Deden Girmansyah 461 (holotype BO; isotypes BO, E). (Fig. 2)

Perennial, creeping, rhizomatous monoecious herb, up to 30 cm tall. Stems

rhizomatous, rooting at the nodes, intemodes very close, 5-10 mm long; stipules

nan*owly triangular, with dense hairs along the dorsal vein, 9 x 4.5 mm, with a hair-

like appendage at the apex covered with additional dense hairs, semi persistent. Leaves

alternate; petioles 10-25 cm long, tomentose, reddish; lamina basifixed, 10-30x 9-29

cm (in live material), asymmetric, suborbicular to broadly ovate, base cordate, lobes

overlapping, apex acute to attenuate, margin sub-entire, with minute teeth at the

end of the veins, adaxial surface glabrescent to sparsely hairy, light green; abaxial

surface shiny red with scattered short hairs, venation palmate, primary veins 7-8.

Inflorescence cymose, with 6-10 flowers, axillary, protandrous, bisexual; peduncle

10-30 cm long, diameter 2-4 mm, puberulous; bracts sub-orbicular, red, c. 3.3 x 2.5

mm, margin slightly fimbriate, persistent. Male flowers: pedicels c. 2 cm long, haiiy;

tepals 4, white to reddish, two outer tepals sub-orbicular, 1.4-2 x 1.2-1. 5 cm, base

slightly cordate, margin entire, apex rounded or slightly undulate, glabrous; two inner

tepals narrowly obovate, white, 1.2-1. 5 x 0.8-1 cm, glabrous; androecium symmetric,

the cluster globose; stamens yellow, c. 48 in number, subequal; filaments 1-1 .5 mm

long, fused at the base; anther obovoid, c. 1 mm long, dehiscing through unilateral

slits as long as anther. Female flowers: pedicels 12-18 mm long, puberoulus; tepals

4, unequal, two outer tepals orbicular to broadly ovate, c. 11 x 14 mm, margin entire,

apex rounded, two inner tepals narrowly obovate, c. 10x5 mm; ovary sub-globose,

5-7 X 2-3 mm, locules 3, placentation axile, entire, wings 3, unequal, obtuse at the

apex, the widest point at the middle of the ovary, 5 mm wide, glabrous; stigmas 3,

yellowish green, Y-shaped, 3 mm long. Fruits with pedicels 2.2-3 cm long; capsule

ellipsoid, 6-10 x 4-8 mm (excluding wings), dehiscent, splitting between the locules

and wings, wings 3, subequal, wings 7-10 mm at the widest point, thinly fibrous.

Seeds barrel-shaped, 0.35-0.37 mm long, collar cells more than a half of seed length.

Distribution. Endemic to West Sumbawa, West Nusa Tenggara, Indonesia.

Habitat Terrestrial herb on steep slopes in tropical forest between 1300-1600 m alt.

Additional specimens examined. West Nusa Tenggara: West Sumbawa, Batu Linting, 15 Jul

2005, Deden Girmansyah 506 (BO); West Sumbawa, Mt Batu Pasak, 18 July 2005, Deden

Gard. Bull. Singapore 68(1) 2016

■4i>a»>ie

1015

Fig. 2. Begonia sumbawaensis Girm. A. Habit. B. Female flower. C. Style. D. Male flower.

E. Stamen. F, Fruit in cross section. G. Fruit. H. Stipule. I. Seed. J. Bract. Drawn by A.

Kusumawati.

Three new Begonia species from Indonesia

81

Girmansyah 535 (BO); West Sumbawa, Olat Tebelah Hill, 9 Jul 2005, Deden Girmansyah 429

(BO); West Sumbawa, Upper Batu Dulang, Brang Bosang River Slope of Batu Linting, 1 5 Jul

2005, HW 12644 (BO); ZW Sumbawa, De Voogd 1634 (BO); West Sumbawa, Mt Batulanteh,

22 Apr 1961, Kostermans 18400 (A, K); West Sumbawa, September 1879, Coifs 299 (L [3

sheets]).

Notes. The epithet is derived from the name of Sumbawa Island. With its creeping habit

Begonia sumbawaensis is typical for Begonia sect. Reichenheimia. It is a very attractive

species with bright red petioles and leaf undersides, and with the inflorescence being

longer than the petioles and bearing many flowers. As such the plant has potential for

ornamental use. This species is found on steep slopes and hence was very difficult to

collect.

Begonia brangbosangensis Girm., sp. nov. ^Petermannia

Begonia brangbosangensis differs from B. lombokensis Girm. (Girmansyah, 2009)

in having the female flower solitary in the leaf axils (in pairs in B. lombokensis),

number of stamens 45 (24 in B. lombokensis), and the female flower pedicel 0.5-0. 8

cm long (1.3 cm long in B. lombokensis). - TYPE: Indonesia, West Nusa Tenggara,

West Sumbawa, Brang Bosang river, upper Batu Dulang Village, 18 July 2005, Deden

Girmansyah 528 (holotype BO; isotypes BO, E). (Fig. 3)

Perenial, erect, monoecious herb, 1 m tall. Stems erect, intemodes 2-10 cm long,

diameter 5-7 mm; stipules narrowly lanceolate, c. 10 x 4.5 mm, pale green, glabrous,

persistent. Leaves alternate; petioles 2-5 cm long, glabrous, green or reddish; lamina

nan'owly ovate to oblong, basifixed, asymmetric, 9-20 x 2.5-6 cm, base subcordate,

margin distantly serrulate, apex attenuate, adaxial surface green, glabrous, abaxial

suface green; venation pinnate, primary veins 3-4 pairs along the midrib; impressed

above, prominent beneath, pale green to burgundy beneath, glabrous. Inflorescence

racemose, bisexual, 3-7 cm long, erect, male flowers distal, female flowers basal;

peduncle 1-3 cm; bracts bright green, elliptic, 9x3 mm, margin entire, caducous.

Male flowers: pedicels c. 1 cm long, glabrous; tepals 2, white to pink, broadly elliptic,

8-1 0 X 6-9 mm, margin entire, glabrous; androecium c. 0.4 mm long, stamens c. 45 in

number, filaments 1-1.5 mm long, fused at the base, anther pale yellow c. 1 mm long,

obovoid, delnscing through unilateral slits c. 'A as long as anther. Female flowers:

pedicels 5-10 mm long, green, glabrous; tepals 2, white, obovate, base subcordate,

apex obtuse to rounded, equal, glabrous, c. 12 x 10mm; ovai*y pale green to reddish

brown, subglobose to ovoid, 8-1 5 x 8-1 3 mm (wings included), capsule ovoid, locules

3, placentation axillary, placental branches 2 per locule; wings 3, unequal, rounded at

the apex, 3-5 mm wide; styles 3, yellow, c. 6 mm long, free to the base, bifurcating with

pale yellow stigma. Fruits with pedicel 1 5 mm long, capsule ellipsoid, 8-20 x 5-9 mm

(excluding wings), dehiscent, splitting between the locules and wings, glabrous, wing

shape as for ovary, wings 8-10 mm wide at the widest point. Seeds numerous, brown,

ellipsoid, 0.34-0.36 mm long, collar cells more than a half of seed length.

82

Gard. Bull. Singapore 68(1) 2016

10 mm

6 mm

A

Fig. 3. Begonia brangbosangensis Gimi. A. Habit. B. Male flower. C. Stamen. D. Female

flower. E. Style. F. Stipule. G. Seed. H. Fruit in cross section. 1. Fruit. J. Ovary. Drawn by A.

Kusumawati.

Three new Begonia species from Indonesia

83

Distribution. Endemic to West Sumbawa, Sumbawa, West Nusa Tenggara, Indonesia.

Habitat. Terrestrial on the forest floor on wet and moist substrates especially along

stream margins and river banks at 1300-1700 m altitude.

Additional specimens examined. West Nusa Tenggara: West Sumbawa, between Brangbosang

river and Mt Batulinting, 15 Jul 2005, Deden Girmansyah 509 (BO); West Sumbawa, Upper

Tepal Traditional Village, slope of MtNgengas, top of Batu Tunin, East Moimtain Forest, 10

Jul 2005, Deden Girmansyah 463 (BO); West Sumbawa, upper Tepal Traditional village, Olah

Tebelat, Pagenong, slope of Mt Ngengas, 9 Jul 2005, Deden Girmansyah 432 (BO); West

Sumbawa, Mt Batulanteh, trail from Batu Dulang to Pusu, 19 Apr 1961, Kostemans 18347 (A,

BO,K); West Sumbawa, Mt Batulanteh, 22 Apr 1961 , Kostermans 18396 (A, BO, K).

Notes. The epithet is derived from the name of the Brang Bosang River in West

Sumbawa.

Begonia jaranpusangensis Ginn., sp. nov. § Petermannia

Begonia jaranpusangensis is similar to B.padangensis Irmsch. in habit and in

having hairy petioles, but differs in having two tepals in the male flowers (four in B.

padangensis), two tepals in the female flowers (five in B. padangensis), and female

flower pedicels 0.5-0.7 cm (10-15 mm in B. padangensis). - TYPE: Indonesia,

West Nusa Tenggara, West Sumbawa, Plampang, between Tana Silo to slope of Mt

JaranPusang, 23 July 2005, Deden Girmansyah 552 (holotype BO; isotypes BO, E).

(Fig. 4)

Perenial, erect, monoecious herb, 1 m tall. Stem green to reddish green, hirsute, 3-10

mm diameter when fresh, intemodes 2-10 cm long; stipules narrowly lanceolate, green,

puberulent, 15-25 x 4-8 mm, margin entire, apex acuminate, ending with a short hair,

1- 2 mm long, persistent. Leaves alternate; distant; petioles red, pilose, 1-5 cm long,

diameter 2-3 mm, grooved above; lamina basifixed, ovate to elliptic, 1 1-18 x 5-9 cm,

asymmetric, base rounded on the broad side and cuneate on the nan*ow side, broad side

4-6.5 cm wide, narrow side 1-3 cm wide, margin denticulate, apex acute to attenuate;

venation pinnate, primary veins 5-6 pairs, impressed above, prominent beneath.

Inflorescences racemose, protogynous, bisexual; male inflorescence monochasial

at base, sympodial at apex, peduncle 1.5-2. 5 cm long, erect, paniculate; female

inflorescence a simple dichasium with a pair of flowers, peduncle 2.5-3 cm long;

bracts bright green, narrowly elliptic, c. 15 x 5 mm, persistent. Male flowers: pedicels

2- 2.5 cm long, white to pinkish, glabrous; tepals 2, suborbicular to broadly ovate ,

white to pink, 12 x 9-10 mm, margin entire, apex rounded, glabrous; androecium c. 5

mm long, stamens c. 34 in number, filaments 1-1 .5 mm long, fused at the base, anthers

pale yellow 1.5-2 mm long, obovoid, dehiscing through unilateral slits c. ’A as long

as anther. Female flowers: pedicels green, puberulent, 5-8 mm long; tepals 5, white,

naiTowly elliptic to lanceolate, glabrous, 12-1 3 x 6-8 mm; ovary pale green, triangular

in outline, 10x4 mm (wings included), wings 3, subequal, rounded at the apex, the

Three new Begonia species from Indonesia

85

widest point 5 mm wide; styles 3, yellow, 4-5 mm long. Fruits with pedicel c.l5 mm,

capsule c. 2 X 1.5 cm (excluding wings), dehiscent, splitting between the locules and

wings. Seeds numerous, brown, broadly ellipsoid to ellipsoid, c. 0.31-0.35 mm long.

Distribution. Endemic to Mt Jaranpusang, West Sumbawa district, Sumbawa, West

Nusa Tenggara, Indonesia.

Habitat. Terrestrial herb, found on the slope of Mt Jaranpusang in rather dry conditions,

at c. 615 m altitude.

Additional specimens examined. West Nusa Tenggara, West Sumbawa, Plampang, between

Tana Silo to slope of Mt Jaranpusang, 23 Jul 2005, Deden Girmansyah 461 (BO).

Notes. The species epithet is derived from the name of Mt Jaran Pusang in West

Sumbawa. Mount Jaran Pusang is separate from the mountains in the central Mt Pasak

complex and has a drier climate. The base of the mountain is covered by bamboo

gardens and the habitat is noticeably dry in appearance. The peak of Mt Jaran Pusang

is rocky and there are few trees of any size; the floristic composition is also rather

different compared with the flora from elsewhere in the middle of West Sumbawa.

Begonia jaranpusangensis was collected from the middle slopes of Mt Jaran Pusang

and is only known from this site.

ACKNOWLEDGEMENTS. I am grateful to the Director of Herbarium Bogoriense for making

available the specimens to study. Dr Harry Wiriadinata and Mr Scott Hoover who encouraged

me to collect Begonia specimens during their 2004 and 2005 expeditions to West Sumbawa

sponsored by the New England Tropical Conservatory (NETC), and Dr Mark Hughes from the

Royal Botanic Garden Edinburgh for cormnents on an earlier version of the manuscript and for

his assistance with the English. 1 would like to thank Wahyudi Santoso and Anne Kusumawati

for their excellent line drawings.

References

Ardi, W.H., Ardhaka, I.M., Hughes, M., Undaharta, N.K.E., Girmansyah, D. & Hidayat, S.

(2013). Two new species of Begonia (Begoniaceae) from Bali and Lombok. Gard. Bull.

Singapore 65(2): 135—142.

De Candolle, A.L.P.P. (1859). Memoire sur la famille des Begoniacees. Ann. des Sci. Nat., Bot.

11:93-149.

Doorenbos, J., Sosef, M.S.M. & De Wilde, J.J.F.E. (1998). The sections of Begonia including

descriptions, keys and species lists (Studies in Begoniacoao VI). Wageningen Agric.

Univ. Pap. 98: 1-266.

Ginnansyah, D. (2009). A taxonomic study of Bali and Lombok Begonia (Begoniaceae).

Reinwardtia 12(5): 419^34.

Ginnansyah, D. & Susanti, R. (2015). Two new species of Begonia (Begoniaceae) from

Bomoo. Kew Bull. 70(2)-19: 1-7.

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Gard. Bull. Singapore 68(1) 2016

Hughes, M. (2008). An Annotated Checklist of Southeast Asian Begonia, xii + 164 p. UK:

Royal Botanic Garden Edinburgh.

Hughes, M., Girmansyah, D. & Ardi, W. (2015a). Further discoveries in the ever-expanding

genus Begonia (Begoniaceae): Fifteen new species from Sumatra. Eur. J. Taxon. 167:

1 ^0 http://m.europeanjoumaloftaxonomy.eu/index.php/ejt/article/view/289 (accessed

on 28 Dec. 2015).

Hughes, M., Moonlight, P., Jara, A. & Pullan, M. (2015b). Begonia Resource Centre. Royal

Botanic Garden Edinburgh http://elmer.rbge.org.ulUbegonia/ (accessed on 22 Dec.

2015).

Linnaeus, C. (1753). Species plantanm: exhibentes plantas rite cognitas, ad genera i^elatas,

cum differentiis specificis, norninibus tiivialibus, synonyntis selectis, locis natalibus,

secundum systema sexiiale digestas. Holmiae: Laurentii Salvii.

Undahaita, N.K.E., I Made Ardaka, Agung Kumiawan & Bayu Adjie (2015). Begonia

bimaensis, a new species of Begonia from Sumbawa Island, Indonesia. GarB. Bull.

Singapore 67(1): 95-99.

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

cyclopedia of botanical exploration in Malaysia and a guide to concerned literature up

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

Gardens’ Bulletin Singapore 68(1): 87-95. 2016

doi: 10.3850/S2382581216000053

87

New records of Orchidaceae from Bali, Indonesia

D. Sulistiarini, D. Arifiani & Y. Santika

Herbarium Bogoriense, Research Center for Biology,

Indonesian Institute of Sciences, Jl. Raya Jakarta-Bogor KM 46,

Cibinong, Bogor, Indonesia

dsulistiarmi@yahoo.com

ABSTRACT. During plant inventory work in the forested areas of Mt Mesehe and Mt Merbuk,

Jembrana district, Bali, twenty orchid species were collected. Among the species collected,

five were confirmed as new records for Bali. The five newly recorded species are Calanthe

zollingeri Rchb.f , Plocoglottis plicata (Roxb.) Ormerod, Podochilus setptylHfolius (Blume)

Lindl., Styloglossum speciosum (Blume) T.Yukawa & RJ.Cribb, and Vanilla albida Blume.

Descriptions of taxa and images are provided.

Keywords. Calanthe, Mt Mesehe, new records, orchids, Plocoglottis, Podochilus, Styloglossum,

Vanilla

Introduction

Bali is famous for its beautiful landscapes and beaches, for which the island has

become the number one tourist destination in Indonesia. In contrast, the plant diversity

of Bali is not well known. There are few floristic publications about Bali, including

little infonnation on the orchid species of the island.

Recent classifications of the Orchidaceae recognise five subfamilies, namely

Apostasioideae, Vanilloideae, Cypripedioideae, Orchidoideae and Epidendroideae,

with about 27,800 species in total (Stevens, 2015). Schuiteman et al. (2014) have

reported that in Malesia there are about 6000 species of which 200 are found in the

Lesser Sunda Islands (Bali, Nusa Tenggara Barat and Nusa Tenggara Timur). For Bali

alone, 146 species of orchids have been recorded (Ginnansyah et al., 2013). Comber

(1990) suggested that Java had 731 species, while Sumatra had a higher number of

1118 species (Comber, 2001). However, more recently, Schuiteman et al. (2014) have

increased the number of orchid species known from Java and Sumatra to 769 and 1 126

species, respectively. Therefore, it is to be expected that more species are to be found

in Bali than those recorded by Ginnansyah et al. (2013).

Most herbarium specimens deposited in the Herbarium Bogoriense (BO) were

collected in the central and eastern parts of Bali, while fewer specimens are from the

western parts of the island. Therefore, an attempt was made recently to explore plant

diversity in the western parts, especially in .lembrana district in the forested areas of

Mt Merbuk (1386 m asl.) and Mt Mesehe (1300 m ask). These are the two highest

mountains in Jembrana district.

88

Gard. Bull. Singapore 68(1) 2016

Jembrana district, which covers 841.8 km^ or 14.93% of Bali, is situated at

8°03'40"-8°50'48"S and 114°25'53"-114°42'40"E. The topography is gently rolling

country to hilly, with a fine-grained soil texture in which the grey-brown alluvial and

regosol types are dominant. Jembrana district has two seasons, i.e., the rainy and dry

season. The heavy rainfall occurs from December to March and the dry season usually

starts from April or May. The average rainfall in 2012 was 1663 mm/year with 112

rainy days per year on average (Anonymous, 2014).

In the Mt Mesehe area, the Yeh Mesehe waterfall (at c. 980 m asl.) is an important

water source for the villages in the region. In the Mt Merbuk area, big rivers, such as

Tukad Aya, Titis and Ijo Gading, are important for maintaining forest conditions. In

Jembrana, 39. 17% of the land area consists of protected forests which in turn preserves

the water catchment. The forests in the Mt Mesehe area are in a good condition from

350 m asl upwards with cultivated areas are only below this altitude. The villagers grow

cacao, coffee, banana and coconut. From 900 m asl upwards the forest is very moist

and foggy, the litter is rather thick to about 1 0 cm, and available trails to explore the

area are veiy steep. Cyathea sp.. Rhododendron sp. and Vaccinium sp. are commonly

found at this elevation. On Mt Merbuk wild palms are common at around 500 m asl,

whereas at 600 to 760 m altitude Dipterocarpus sp., Podocarpus sp., Fagraea sp. and

Pavetta sp. are common. Along the Titis river, there are, amongst others, Nephrolepis

biserrata (Sw.) Schott, Asplenium nidus L., Dry^opteris sp., Selaginella sp., bryophytes.

Begonia sp. and some orchids. The rattans Daemonorops melanochaetes Blume and

Calamus javensis Blume, along with climbers such as Freycinetia javanica Blume and

members of the Vitaceae are also commonly found.

Methods

A plant inventory was conducted in the Jembrana district in the vicinity of Mt Mesehe

in May 2013 and in the Mt Merbuk forest area in April 2014. The collection of orchids

followed the methodology of Van Balgooy (1987). The collections were photographed

when flowering and then dried as herbarium specimens. Some spirit samples of flowers

were also prepared using a mix of glycerine 40%, alcohol 96% and distilled water at

a ratio of 1:70:29 (Rugayah et al., 2004). Sterile specimens were also collected for a

pennanent record of all orchid species in Bali. All measurements in the descriptions

are based on dried specimens.

Five of the orchid species found are new records as they are not included in

the recently published checklist of the Flora of Bali (Girmansyah et al., 2013) and

their occurrence in Bali is not noted in eMonocot (2015), Schuiteman et al. (2014),

Comber (1990, 2001), Seidenfaden & Wood (1992) and Wood et al. (1993). Many

orchid species are reported to be distributed in the Lesser Sunda Islands (LSI) without

further elaboration on which specific island(s) they can be found. In this paper we do

not include those species as new records.

New orchid records from Bali

89

Results and Discussion

The exploration of Mt Mesehe and Mt Merbuk has resulted in many new orchid

specimens of 20 species (Table 1) for Herbarium Bogoriense (BO). Previously only

about sixty orchid species have been recorded from Bali but among the 20 species

newly collected we were able to confirm that five species are new records. Four of

these species belong to the subfamily Epidendroideae, i.e. Calanthe zollingeri Rchb.f ,

PlocogJottis plicata (Roxb.) Onuerod and Podochilus serpyUifoUus (Blume) LindL,

Styloglossum speciosum (Blume) T.Yukawa & P.J.Cribb. The remaining species is

Vanilla albida Blume belonging to the subfamily Vanilloideae, Epidendroideae includes

78% of the species of Orchidaceae and is characterised by the presence of incumbent

anthers which bend forward during column elongation (Stevens, 2015; Freudenstein

& Chase, 2014). In Indonesia, Styloglossum speciosum and Plocoglottis plicata were

previously known from Java, Maluku, Sulawesi, Kalimantan and Sumatra. Calanthe

zollingeri, Podochilus seipyllifolius and Vanilla albida were previously known from

Java, Kalimantan and Sumatra. These species are relatively common, yet they have

not been recorded from Bali before.

Apart from the five newly recorded species, there are another eleven that have

previously been recorded from the Lesser Sunda Islands but further study is needed to

ascertain whether those records included Bali.

Description of taxa

Calanthe zollingeri Rchb.f, Bonplandia 5: 38 (1857); Comber, Orchids Java: 93

(1990). - TYPE: Java, Banyuwangi, Leg. ign. 2858 (holo W, n.v.). (Fig. 1)

Pseudobulbs close together, bearing 4-5 leaves. Leaves 23-40 cm long, 8-9 cm wide,

ovate, acuminate, petiole 10-12 cm long. Inflorescences 40-60 cm tall, rachis c. 6 cm

long, bearing 10-20 flowers, not all open at one time. Flowers white, about 2.5 cm

wide; sepals and petals about 1 cm long, oblong; lip white and then turning yellow to

orange, unlobed, circular, deeply split, outer margin incurved.

Distribution. Peninsular Malaysia, Sumatra, Java, Bali, Borneo.

Habitat. Lowland forest along river, also in primary forest.

Specimens examined. INDONESIA. Bali: Jembrana district: Mendoyo subdistrict, Batu

Agung village, Panca Seining, forest surrounding Mt Sekar, 550 m asl., 1 7 May 2013, N. Utami

2032 (BO); Jembrana subdistrict, Dauh Warn village, Mundulc Warn, forest surrounding Mt

Merbuk, 520 m asl., 23 Apr 2014, Y. Santika 749 (BO); Jembrana subdistrict, Batu Agung

village, Palungan Batu, Tuntuna, along Tukad Aya river, forest surrounding Mt Merbuk, 200 m

asl., 23 Apr 2014, S. Sunarti 755 (BO).

90

Card. Bull. Singapore 68(1) 2016

Plocoglottis plicata (Roxb.) Ormerod in Comber, Orchids Sumatra: 293 (2001).

- Limodorum plicatum Roxb., FI. Ind. ed. 3: 465 (1832). - Plocoglottis acuminata

Blume, Ann. Mus. Bot. Lugduno-Batavum 1: 46 (1849). - TYPE: Java, Blume s.n. (L,

n.v.). (Fig. 2)

Pseudobulbs well-spaced along rhizome, 4-6 cm long, 0.2-0.5 cm wide, cylindrical,

usually covered by leaf sheath. Rliizome creeping, branched, intemodes 1.5-2. 5 cm

long, 0.3-0.4 cm thick. Leaf 1, ovate, 20-25 cm long, 7-8.5 cm wide, acuminate,

petiole 4-7 cm long. Inflorescences from base of young pseudobulb, 30^0 cm long,

bearing several flowers. Flowers opening widely, yellow with red markings, 3 cm

wide; sepals and petals narrowly lanceolate, acute; lip in outline almost square.

Distribution. Sumatra, Java, Bali, Borneo, Moluccas, Sulawesi, Philippines.

Habitat. Lowland forest along river; primary forest, sometimes growing on rocks.

Specimens examined. INDONESIA. Bali: Jembrana district: Mendoyo subdistrict, Poh

Santen village, Pasatan, forest suiTounding Mt Mesehe, 400 m ash, 18 May 2013, D. Arifiani

1365 (BO); Mendoyo subdistrict, Batu Agung village, Panca Seming, Yeh Mesehe protected

forest, 860 m ash, 17 May 2013, K Santika 548 (BO); Jembrana subdistrict, Batu Agung village,

Palungan Batu, Pangkung Palir, along Tukad Aya river, forest surrounding Mt Merbuk, 280 m

ash, 23 Apr 2014. D. Arifiani 1416 (BO).

Podochilus serpyllifolius (Blume) Lindh, J. Proc. Linn. Soc., Bot. 3: 37 (1858);

Comber, Orchids Java: 202 (1990). - Cryptoglottis serpyllifolia Blume, Bijdr. Fh Ned.

Ind.: 296 (1825). - TYPE: ?Indonesia, T. Lobb 155 (AMES, n.v.). (Fig. 3)

Pseudobulbs absent. Stems hard, creeping, hanging, branching. Leaves very small,

in two rows along the stem, about 1 mm apart, ovate, 3-4 mm long, c. 1 mm wide,

sessile, acute. Inflorescences usually tenninal, bearing up to 4 flowers. Flowers not

opening widely, about 4.5 mm long and 1 .5 mm wide, white with a pink-purple spot in

the middle of the petal and lip.

Distribution. Sumatra, Java, Bali, Borneo.

Habitat. Primary forest, growing in moss on rocks.

Specimen examined. INDONESIA. Bali: Jembrana district: Mendoyo subdistrict, Batu

Agung village, Panca Seming, Yeh Mesehe protected forest, 935 m ash, 17 May 2013, Y.

Santika 570 (BO).

New orchid records from Bali

91

Fig. 1. A-B. Calanthe zollingeri Rchb.f. (Photos: Yessi Santika)

Fig. 2. Plocoglottis plicata (Roxb.) Ormerod. A. Habit. B. Detail of the flower. (Photos: Deby

Ariflani)

92

Gard. Bull. Singapore 68(1) 2016

Styloglossum speciosum (Blume) T.Yiikawa & PJ.Cribb, Bull. Natl. Mus. Nat. Sci.,

Ser. B, 40(4): 150 (2014). - Calanthe speciosa (Blume) Lindl., Gen. Sp. Orchid. PL:

250 (1833); Comber, Orchids Java: 93 (1990); Seidenfaden & Wood, Orchids Penins.

Malaysia Singapore: 175 (1992). - Amblyglottis speciosa Blume, Bijdr. FI. Ned. Ind.:

371 (1825). - TYPE: Java, Blume s.n. (L, n.v.). (Fig. 4)

Pseudobulbs close together, with 4-7 leaves, hidden by leaf sheaths. Leaves 40-50

cm long, 8-10 cm wide, oblong, nerves prominent, acute, leaf sheaths overlapping.

Inflorescences 45-80 cm long; sterile bracts 5, boat-shaped, 2-6 cm long; bracts

caducous, absent during flowering. Flowers 3 cm wide, petals and midlobe of lip

yellowish, sepals golden yellow; sepals and petals ovate, acuminate; lip three-lobed,

with two basal rounded calli.

Distribution. Southeastern and southern China, Japan (R 5 aikyu), Taiwan, Vietnam,

Thailand, Peninsular Malaysia, Sumatra, Java, Bali, Borneo, Sulawesi, Moluccas,

Philippines.

Habitat. Primary forest.

Specimen examined. INDONESIA. Bali: Jembrana district: Mendoyo subdistrict, Batu

Agung village, Panca Seining, Yeh Mesehe protected forest, 700 m asL, 17 May 2013, Y.

Santika 542 (BO).

Vanilla albida Blume, Cat. Gew. Buitenz.: 100 (1823); Comber, Orchids Java: 76

(1990); Seidenfaden & Wood, Orchids Penins. Malaysia Singapore: 127 (1992);

Comber, Orchids Sumatra: 130 (2001). - TYPE: Java, Blume s.n. (holo L, n.v.).

Pseudobulbs absent. Stems climbing, with intemodes 10-11 cm long, leaves alternate,

aerial roots opposite leaves. Leaves lanceolate, 15-18 cm long, 3-4 cm wide, acute,

petiole 1-1.5 cm long. Inflorescences axillary, bearing many flowers, floral bracts

obtuse. Flowers open only on one day, at first widely open and then half-closing;

sepals and petals pale yellow or yellowish green; lip white, but violet inside near the

base, recurved, trumpet-like.

Distribution. Andaman Islands, Nicobar Islands, Sumatra, Java, Bali, Borneo.

Habitat. Lowland forest.

Specimen examined. INDONESIA. Bali: Jembrana district: Mendoyo subdistrict, Poh

Santen village, Pasatan, forest surrounding Mt Mesehe, 360 m asL, 18 May 2013, D. Arifiani

1386 (BO).

New orchid records from Bali

93

Fig. 3. Podochilus serpyllifolius (Blume) Lindl. A. Habit. B. Detail of the flower. (Photos:

Yessi Santika)

Fig. 4. Styloglossum speciosum (Blume) T. Yukawa & P. J. Cribb. A. Habit. B. Detail of the

infloreseence with flower in front view (inset C). (Photos: Yessi Santika)

94

Gard. Bull. Singapore 68(1) 2016

Table 1. Orchid species collected from Mt Mesehe and Mt Merbuk. (** = newly recorded

species from Bali, names with cf. are based on sterile specimens).

No.

Species name

Notes

1

Appendicula comuta Blume

Mt Mesehe, moist forest, alt. 500-1500 m.

Epiphyte.

2

Bulbophyllum lobbii Lindl.

Mt Mesehe, moist forest, alt. 900-1000 m.

Epiphyte.

3

Calanthe cf. mcisuca (D.Don) Lindl.

Mt Merbuk, shady primary forest, alt. 400-

1 600 m, Ten-estriaf

4

Calanthe flava (Blume) C.Morren

Mt Merbuk, primary forest, alt. 700-2200 m.

Terrestrial.

5

Calanthe zollingeri Rchb.f. **

Mt Mesehe and Mt Merbuk, lowland forest

along river, also in primary forest, alt. 200-

550 m. Terrestrial.

6

Ceratostylis brevibrachiata J.J.Sm.

Mt Mesehe, moist forest, alt. 800-1 000 m.

Epiphyte.

7

Dendrobium cf montanum J.J.Sm.

Mt Mesehe, moss forest, alt. 900-1000 m.

Epiphyte.

8

Dendrobium plicatile Lindl.

Mt Mesehe, lowland forest, alt. 500-800 m.

Epiphyte.

9

Liparis condylobulbon Rchb.f

Mt Mesehe and Mt Merbuk, lowland forest

in open area, alt. 500-800 m. Epiphyte.

10

Mycaranthes oblitterata Blume

Mt Mesehe, lowland forest, alt. 400-800 m.

Epiphyte.

11

Nervilia concolor (Blume) Schltr.

Mt Merbuk, lowland forest in open area, alt.

400-600 m. Ten'estrial.

12

Phaius pauciflorus (Blume) Blume

Mt Mesehe and Mt Merbuk, lowland forest

in open area, alt. 400-700 m. Terrestrial.

13

Pinalia cf. midtiflom (Blume)

Kuntze

Mt Mesehe, lowland forest in open area, alt.

600-800 m. Epiphyte.

14

Plocoglottis plicata (Roxb.) Onnerod

**

Mt Mesehe and Mt Merbuk, lowland forest

or primary forest, sometimes growing on

rocks, alt. 200-900 m. Terrestrial.

15

Podochilus serpyllifolius (Blume)

Lindl. **

Mt Mesehe, primary forest, alt. 900-1000 m.

Epiphyte.

16

Spathoglottis plicata Blume

Mt Merbuk, lowland forest in open area, alt.

400-600 m. Terrestrial.

17

Styloglossum speciosum (Blume)

T. Yukawa & RJ.Cribb **

Mt Mesehe, primary forest, alt. 700-850 m.

Terrestrial.

18

Trichotosia ferox Blume

Mt Mesehe, primary moist forest, alt. 800-

950 m. Epiphyte.

19

Tropidia angidosa (Lindl.) Blume

Mt Merbuk, lowland forest, alt. 400-600 m.

Terrestrial.

20

Vanilla albida Blume **

Mt Mesehe, lowland forest, alt. 300^00 m.

Terrestrial becoming climber.

New orchid records from Bali

95

ACKNOWLEDGEMENTS. This study was financially supported by the Research Center for

Biology-Indonesian Institute of Sciences (DIPA 2013 and 2014). We are grateful to the Head of

the Dinas Kehutanan Jembrana, to the staff for their valuable support, and to the people of Batu

Agung village for their assistance during field work. Two anonymous reviewers are thanked for

their valuable and constructive suggestions.

References

Anonymous (2014). Profil Kabupaten Jembrana 2014. 227 p. Bali: Badan Perencanaan dan

Pembangunan Daerah dan Penanaman Modal, Pemerintah Kabupaten Jembrana, Bali.

Comber, J.B. (1990). Orchids of Java. UK: Royal Botanic Gardens, Kew.

Comber, J.B. (2001). Orchids of Sumatra. Kota Kinabalu: Natural History Publications

(Borneo) & UK: Royal Botanic Gardens, Kew.

eMonocot (2015). An online resource for monocot plants, http://e-monocot.org. (accessed on

Jim. 2015).

Freudenstein, J.V. & Chase, M.W. (2014). Phylogenetic relationships in Epidendroideae

(Orchidaceae), one of the great flowering plant radiations: progressive specialization

and diversification. Bot. 115(4): 665-681.

Girmansyah, D., Santika, Y., Retnowati, A., Wardani, W, Haerida, L, Widjaja, E.A & Van

Balgooy, M.M.J. (2013). Flora of Bali: An Annotated Checklist. Jakarta: Research

Center for Biology-LlPl & Yayasan Pustaka Obor Indonesia.

Rugayah, Retnowati, A., Windadri, F.I. & Hidayat, A. (2004). Pengumpulan data taksonomi. In:

Rugayah, E.A. Widjaja & Praptiwi (eds) Pedoman Pengumpulan Data Keanekaragaman

Flora. Pp. 5^2. Jakarta: Pusat Penelitian Biologi— Lembaga Ilmu Pengetahuan

Indonesia.

Schuiteman, A., De Vogel, E.F., Vermeulen, J.J., Kessler, P. & Vogel, A. (2014). Malesian

Orchid Genera Illustrated. http://’www.nationaalherbarium.nl/pubs/orchidweb/

Malesianorchidgenera.htm. (accessed on Oct. 2014).

Seidenfaden, G. & Wood, J.J. (1992). The Orchids of Peninsular Malaysia and Singapore.

Fredensborg: Olsen & Olsen. UK: Royal Botanic Gardens, Kew & Singapore: Singapore

Botanic Gardens.

Stevens, P.F. (2015). Angiospenn Phylogeny Website, version 12, July 2012 [and more or

less continuously updated since]. http://www.mobot.org/MOBOT/research/APweb/

(accessed on Jul. 2015).

Van Balgooy, M.M.J. (1987). Collecting. In: De Vogel, E.F. (ed) Manual of Herbarium

Taxonomy, Theory and Practice. Pp. 14-19. Jakarta: UNESCO.

Wood, J.J., Beaman, R.S. & Beaman, J.H. (1993). The Plants of Mount Kinabalu, 2. Orchids.

UK: Royal Botanic Gardens, Kew.

Gardens’ Bulletin Singapore 68(1): 97-98. 2016

doi: 10.3850/S2382581216000065

97

Olea luzonica (Oleaceae), a new name

for O. obovata from the Philippines

R. Kiew

Forest Research Institute Malaysia,

52109 Kepong, Selangor, Malaysia

ruth@frim .gov.my

ABSTRACT. A new name, Olea luzonica Kiew, is proposed to replace the later homonym Olea

obovata (Merr.) Kiew.

Keywords. Illegitimate, Linociera obovata

Introduction

The Philippine species Olea obovata (Merr.) Kiew has an illegitimate name because

Olea obovata Baker was earlier published for a species from Mauritius (Baker, 1 877:

219). This older name came to light when Hong-Wa et al. (2014) revised Noronhia

for Mauritius. Their Noronhia obovata (Baker) Hong-Wa & Callm. is based on Olea

obovata Baker and its type. This, therefore, necessitates a new name for the Philippine

Olea and it is here proposed to name it Olea luzonica Kiew because it is known only

from that island.

Taxonomy

Olea luzonica Kiew, nom. nov. -Linociera obovata Merr. [in Elmer, Leafl. Philip. Bot.

5: 1652 (1913) nom. mid.] Philip. J. Sci. (Bot.) 10: 338 (1915); Merrill, Enum. Philip.

PI. 3: 304 (1923). - Olea obovata (Merr.) Kiew, Gard. Bull. Singapore 51 : 94 (1999),

non Baker (1877); Green, Kew Bull. 57: 116 (2002). - TYPE: Philippines, Luzon,

Laguna Province, San Antonio, September 1912, Ramos Bur. Sci. 15014 (lectotype K,

designated by Kiew (1999)).

Small tree, glabrous except minutely pubemlent on branchlets and inflorescences.

Twigs grey to whitish, stout, terete, glabrous. Leaves: petioles 1-1.5 cm; lamina very

thickly coriaceous, obovate, 4-7 x 2^.5 cm, pale when dry, upper surface glossy, dull

beneath, base narrowed and more or less decurrent, margins entire, recurved, apex

broadly rounded, sometimes very shortly and broadly acuminate, sometimes even

slightly retuse; lateral veins indistinct, often subobsolete, c. 8 on each side of midrib.

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Gard. Bull. Singapore 68(1) 2016

Inflorescences axillary, sessile, densely many-flowered cymose panicles, branched

from base, brown when dry, 2-5 cm long, the branches spreading and 2-3 cm wide.

Flowers (mature female flowers not seen) 4-merous, buds ellipsoid or obovoid, c. 2

mm long, pedicels 1-3 mm. Male flowers', calyx 0. 5-0.7 mm long, lobes irregularly

broadly triangular, 0.3-0. 5 imn long, apex acute; corolla thickish, tube 0.5 mm long,

lobes involute, 1 mm long; filaments 0.2 mm long, anthers ellipsoid, 0.8 mm long,

with a very small terminal appendage. Drupes ellipsoid 6-7 x 4.5-5 mm.

Distribution. Endemic in the Philippines, Luzon (Ilocos Norte, Laguna, Nueva Ecija

and Tayabas Provinces).

Ecology. In mossy forest at about 2000 m. From herbarium specimens, it apparently

has the habit of many trees of mossy forest as judged from its much branched, bushy

canopy with branches of upstanding, thick leaves with recurved margins at the tip of

the twigs.

Additional specimens examined. PHILIPPINES: Luzon: Ilocas Norte, Mt. Palimlim Ramos

Bur. Sci. 33262 (L); Nueva Ecija, Mt. Umingan Ramos & Edano Bur Sci 26328 (K); Tayabas,

Mt. Alzapan Ramos & Edano Bur Sci. 45667 (K), 45716 (K).

Notes. Olea luzonica is a very distinctive species in its small, very thick, obovate

leaves with a rounded apex and decurrent base, obscure lateral veins, recurved margin

and dense pyramidal inflorescences. It was first listed by Elmer (1913) as Linociera

obovata, but lacking a Latin diagnosis the name was a nomen nudum. Merrill (1915)

validated this name by providing a description and the required diagnosis. In 1999,

Kiew recognised it was an Olea species and transferred it to Olea.

ACKNOWLEDGEMENTS. I am indebted to Jose Ignacio de Juano Clavero for pointing out

the illegitimate status of Olea obovata (Merr.) Kiew.

References

Baker, J.G. (1877). Flora of Mauritius and the Seycelles. London: L. Reeve & Co.

Elmer, A.D.E. (1913). Philippine L/woczera. Leaf. Philipp. Bot. 5: 1651-1657.

Hong-Wa, C., Callmander, M.W. & Baider, C. (2014). Taxonomy and conseiwation of the genus

Noronhia Thouars (Oleaceae) in Mauritius. Candollea 69: 157-163.

Kiew, R. (1999). Reappraisal of Olea species in Malesia. Gard. Bull. Singapore 51: 85-98.

Merrill, E.D. (1915). New or noteworthy Philippine plants XII. Philipp. J. Sci., C. 10: 287-349.

Gardens’ Bulletin Singapore 68(1): 99-107. 2016

doi: 10.3850/S2382581216000077

99

A new species and a new record in Begonia sect.

Platycentrum (Begoniaceae) from Thailand

T. Phutthai^ & M. Hughes^

‘Conservation Biology Program, Division of Biological and Natural

Resources Sciences, Mahidol University (Kanchanaburi campus),

Sai-Yok, Kanchanaburi 71150, Thailand

thamarat65@hotmail.com

^Royal Botanic Garden Edinburgh, 20A Inverleith Row,

Edinburgh, EH3 5LR, Scotland, U.K.

ABSTRACT. Begonia khaophanomensis Phutthai & M. Hughes is described from a collection

made on granitic rock in Krabi Province in Peninsular Thailand. It is endemic to the type

locality of Khao Phanom Bencha National Park, and is assessed as Least Concern using

the lUCN criteria. Begonia macrotoma Innsch. is noted as a new record for Thailand from

collections in Chiangmai, Nakhon Ratchasima and Nakhon Nayok provinces; the species

is also found in China, Myanmar, NE India, Nepal and Vietnam. Both species belong to

Begonia sect. Platycentrum. A key to the species of Begonia sect. Platycentrum in Thailand

is provided.

Keywords. Begonia khaophanomensis. Begonia macrotoma, China, Myanmar, NE India,

Nepal, Peninsular Thailand, Vietnam

Introduction

The genus Begonia L. is one of the largest angiospenn genera, comprising around

1803 species (Hughes et al,, 2015). Thailand has 54 accepted species of Begonia,

which are found in habitats from near sea level to high mountain summits, with nearly

half of them occurring on karst limestone (Hughes, 2008; Phutthai et al., 2009, 2012,

2014).

Begonia sect. Platycentrum (Klotzsch) A.DC. comprises 126 species (Hughes

et al., 2015) and is widely distributed in Asia, ranging from India to the Himalayas,

Indo-China, China, Taiwan, and Malesia (Doorenbos et al., 1998; Shui et al., 2002;

Ding et al., 2014). In Thailand, six species are reported in the section to date: Begonia

cathcartii Hook.f., B. hatacoa Buch.-Ham. ex D.Don, B.palmata D.Don, B. siamensis

Gagnep, B. smithiae Geddes and B. venusta King. They are all confined to montane

forests above 1000 m in altitude (Hughes, 2008; Phutthai et al., 2009). However,

based on molecular data (Thomas et al., 2011), the placement of Begonia smithiae in

Begonia sect. Platycentrum appears to be eiToneous and is a phylogenetically rather

isolated species closer to Begonia sect. Parvibegonia A.DC. (Doorenbos et al., 1998).

100

Gard. Bull. Singapore 68(1) 2016

A new species and a new record in Begonia sect. Platycentrum are presented

here. They are characteristic of the section in having rhizomatous stems, 2-locular

ovaries with branched placentas, and anthers with extended connectives. Begonia

khaophanomensis Phutthai & M.Hughes is a rare and narrowly endemic species

which is known only from the type locality in Krabi Province. It grows in granitic

rock crevices at altitudes around 1200 m in Khao Phanom Bencha National Park. We

also report Begonia macrotoma Irmsch. as a new record for the flora of Thailand from

Chiangmai, Naklion Ratchasima and Nakhon Nayok provinces. Terminology in the

descriptions follows Beentje (2010).

Key to the species of Begonia sect. Platycentrum in Thailand

la. Leaf lamina ovate to lanceolate, without lobes 2

lb. Leaf palmatifid, with some degree of lobing 5

2a. Stems and petioles glabrous or subglabrous 3

2b. Stems and petioles with a distinct indumentum 4

3a. Leaf lamina ovate, tepals white B. venusta

3b. Leaf lamina lanceolate, tepals with red stripes on the outer surface B. hatacoa

4a. Stem and petiole puberulent, leaves glabrous above 5. siamensis

4b. Stem and petiole hirsute, leaves echinate above B. cathcartii

5a. Stem and petioles reddish brown sericeous B. palmata

5b. Stem and petioles glabrous or white hirsute or pilose 6

6a. Stem with red striations, leaf lamina deeply palmatifid, bulbils often present,

ovary glabrous B. macrotoma

6b. Stem unifoiTn green, leaf shallowly palmatifid, bulbils absent, ovary white hirsute . .

B. khaophanomensis

Begonia khaophanomensis Phutthai & M.Hughes, sp. nov. § Platycentrum

Begonia khaophanomensis is similar to B. palmata in habit and leaf shape, but differs

in having leaves occuiring in opposite pairs at the apex of the plant (not alternate),

with white short stiff hairs on the petioles and leaves (not brown sericeous), and a

white ovary with white short stiff hairs (not pinkish green and subglabrous). - TYPE:

Thailand, Krabi, Kliao Phanom Distiict, Kliao Phanom Bencha National Park, trail

from Ban San to top of Khao Phanom Bencha, on rock along trail in lower montane

forest, 8°17'02"N 98°56T7"E, 1200 m, 19 June 2006, Williams, K., Pooma, R.,

Poopath, M., Chamchamroon, V. & Seasin, S. 1943 (holotype E; isotypes BKF). (Fig.

1 , 2 )

A new Begonia from Thailand

101

Fig. 1. Begonia khaophanomensis Phutthai & M.Hughes. A. Habit. B. Staminate and pistillate

flowers. From the type K. Williams et al. 1943. (Photos: M. Poopath)

N.O.OaZ NmO.OoBL N.O.OoSl N„0,0oSL N„0.0o6 N„0,0c9

102

Gard. Bull. Singapore 68(1) 2016

Vietnam

Myanmar

Thailand

Cambodia

Vietnam

Kilometers

Malaysia

Indonesia

Sources: Esri.'lJSGS NOAA

Fig. 2. Distribution of Begonia khaophanomensis Phutthai & M.Hughes (triangle) and B.

macrotoma Irmsch. (dot).

6“0'0''N eWN 12°0’0"N 15°0'0"N 18°0’0'’N

A new Begonia from Thailand

103

Monoecious herb, c. 60 cm tall. Rhizome elongated with numerous fibrous roots,

glabrous. Stems erect, terete, pale green. Leaves 6-8 per plant (occurring as an

opposite pair subtending the inflorescence), lamina basifixed; petiole 7-12 cm long,

with dense white short stiff hairs (hirsute); leaf blade membranaceous, asymmetric,

chartaceous when dry, base deeply cordate, adaxial surface densely minutely hirsute,

dull dark green, abaxial surface paler, densely hirsute especially on veins, 8-10 x

5-13 cm, shallowly palmatifid with 6-8 lobes, margin seiTate-fimbriate with long

stiff hairs, venation palmate, midrib and lateral veins concolourous with lamina, 5-7

veins, slightly impressed above and prominent beneath, apex acuminate. Stipules

prominent, pale green, ovate-lanceolate, densely hairy, 5-15 x 3-5 mm, apex acute,

margin entire. Inflorescences terminal or axillary, bisexual, 8-10 cm long, cymose

with c. 3 staminate flowers and 2 pistillate flowers, protandrous; peduncle c. 8 cm

long, terete, pale green with dense short stiff hairs. Bracts caducous, lanceolate, pale

green 3-5 x 2-4 mm, apex acute, margin entire. Staminate flower, pedicels white, c.

10 mm long; tepals 4, white; outer 2 broadly elliptic, c. 15 x 20 mm, apex rounded,

margin entire, base rounded, outer surface with dense short scabrid hairs, inner surface

glabrous; inner 2 obovate, c. 12 x lo mm, apex obtuse, margin entire, base obtuse,

outer surface with sparse short scabrid hairs, inner surface glabrous; androecium

actinomorphic, globose, stamens numerous, yellow, filaments united at the base, c. 3

mm long, anthers oblong, c. 2 mm long, dehiscing by longitudinal slits, comiective

extended, rounded. Pistillate flowers', pedicels white, c. 10 mm long; tepals 5, white,

subequal; densely scabrid adaxially, 1 8-20 x 20 mm; styles 2, fused at base, crescent-

shaped, stigmatic band twice spirally twisted; ovary white, densely minutely scabrid,

with 3 unequal wings, 2-locular, placenta bilamellate. Fruits dehiscent between the

two smaller wings, reddish green, drying pale brown, capsule obovate, c. 14 x g

mm, abaxial wing oblong or obliquely triangular, densely minutely scabrid, c. 40 mm

long, 2 lateral wings c. 4 mm long, cucullate. Seeds numerous, brown, barrel-shaped,

0.4-0.5 mm long.

Distribution. Endemic to Peninsular Thailand (known from only the type locality).

Habitat and ecology’. On granitic rock crevices by streams in hill evergreen forest and

seasonally dry evergreen forest at c. 1200 m. Flowering and fruiting June to October.

Provisional lUCN category. Least Concern (LC). This species is only known from

the type locality in the Khao Phanom Bencha National Park which currently is well

protected and likely covers the entire natural range of the species. Hence, although

rare, the threat criteria for Critically Endangered or Endangered are not met. Begonia

khaophanomensis potentially qualifies as Vulnerable under VUD2, although with the

lack of a plausible future threat we consider Least Concern to be the most appropriate

category as long as Khao Phanom Bencha remains well managed.

104

Gard. Bull. Singapore 68(1) 2016

Notes. The opposite leaves and stiff hairs indicate an alliance to Begonia areolata Miq.

from Sumatra and Java which differs in having leaves which are minutely bullate and

with red hairs, and in having petals which are elongate-elliptic in the male flowers (not

broadly elliptic). The specific epithet refers to the district in which the type material

was collected, Khao Phanom.

Begonia rnacrotoma Irmsch., Notes Roy. Bot. Gard. Edinburgh 21; 41 (1951); Gu et

ah, FI. China 13: 185 (2007); Hughes, Amiot. Checklist of Southeast Asian Begonia

11 (2008). - TYPE: China, Yunnan, 29 June 1938, Yu, TT 17778 (holotype KUN, n.v.;

isotype E). (Fig. 2, 3)

Monoecious herb, 60-90 cm tall. Rhizome elongated with numerous fibrous roots,

glabrous, c. 20 mm diam. Stems erect, terete, pale green with numerous dark red

striations. Leaves 3-6 per plant, lamina basifixed; petiole 10-20(-30) cm long, pilose;

leaf blade membranaceous, asymmetric, chartaceous when dry, base slightly cordate,

adaxial surface densely scabrid, dull dark green, abaxial surface paler, sparsely scabrid,

glossy; 12-30 x 7-28 cm, palmatifid to deeply palmatifid with 3-8 lobes, margin

sinuate or denticulate, venation palmate, midrib and lateral veins concolourous with

lamina, 5-7 veins, slightly impressed above and prominent beneath, apex acuminate;

bulbils and plantlets fonn at the sinus of the lamuia. Stipules caducous, dark red,

ovate, 2-3 x 1.8 cm, apex cuspidate, margin entire. Inflorescences terminal, bisexual,

20-40 cm long, compound cyme, branching 1-2 times with 3-1 2 staminate flowers

and 2-8 pistillate flowers per branch, protandrous; peduncle 15-25 cm long, terete,

pale green with dense brown villous hairs. Bracts caducous, ovate, dark red, 3-10 x

4-10 mm, apex acute, margin entire. Staminate flowers: pedicels pale pink, 10-20

mm long; tepals 4, white or pale pink; outer 2 orbicular, 20-25 mm diameter, apex

rounded, margin entire, base rounded, outer surface magenta hirsute at the centre,

inner surface glabrous; inner 2 obovate, 15-20 x 5-10 rmii, apex obtuse, margin entire,

base obtuse, glabrous on both sides; androecium actinomoi*phic, globose, stamens

numerous, yellow, filaments united at base, c. 2 mm long, anthers obovate, c. 2.5

mm long, dehiscing by lateral slits, connective extended, rounded. Pistillate flowers:

pedicels pale pink, 10-20 mm long; tepals (?3-)5, white or pale pink, unequal; outer

2 orbicular, outer surface pubescent, 18-20 x 20-25 mm, inner 1-3 obovate, c. 20 x

8-12 mm, outer surface pubescent; styles 2, crescent-shaped, fused at base, stigmatic

band twice spirally twisted; ovary pale pinlc, minutely colliculate, with 3 unequal

wings, 2-locular, placenta bilamellate. Fruits dehiscent dehiscent between the two

smaller wings, pale green, drying pale brown, capsule obovate, c. 14x8 mm, abaxial

wing oblong with a rounded tip, 35-40 mm long, 2 lateral wings c. 5 mm long, curved

along the ovary. Seeds numerous, brown, ban*el-shaped. 0.5-0. 6 mm long.

Distribution. In Northern Thailand, in Chiangmai province, and in Southeastern

Thailand, in Nakhon Ratcahsima and Nakhon Nayok Provinces. Further distribution

in China, Myanmar, NE India, Nepal and Vietnam.

A new Begonia from Thailand

105

Fig. 3. Begonia macrotoma Irmsch. A. Leaf shape and bulbils/plantlets at the sinus of the leaf

lamina. B. Rhizome. C. Pistillate flowers. D. Inflorescenee. E. Staminate flowers. F. Magenta

hirsute hairs on the outer tepals. A & C from Middleton et al. 4515 in cultivation at the Royal

Botanic Garden Edinburgh, accession number 20082075; B, D-F from Phutthai 145. (Photos:

A & C: M. Hughes; B, D-F: T. Phutthai)

106

Gard. Bull. Singapore 68(1) 2016

Habitat and ecology. In granitic rock crevices by streams in hill evergreen forest and

seasonally dry evergreen forest at around 600 m. Flowering May to November; fmiting

November to January.

Provisional lUCN category. Least Concern (LC). The species has a wide distribution

in evergreen and seasonal montane forests.

Specimens examined. THAILAND: Chiang Mai: Doi Inthanon National Park, 25 Jan 1990,

Hoover, JV.S. 727 (K [3 sheets]); Doi Inthanon National Park, Wachirithan Waterfall, 19 Sep

2008, Middleton, D.J. el al 4515 (BK, BKF, E); ibidem, 24 Nov 2007, Phutthai, T. 145 (PSU,

BKF); Nakhon Ratchasima: Khao Yai National Park, Orchid Waterfall, 22 Oct 1969, van

Beusekom, C. F. & Charoenpol, C. 1834 (AAU [2 sheets], BKF, E [2 sheets], L); ibidem,

31 Oct 1970, Charoenpol, C. et al 4349 (AAU, BICF); Kliao Yai National Park, Haew Sai

Waterfall. 20 Nov 1 984, McAllanA. 9 (K [3 sheets]); Nakhon Nayok: Khao Yai National Park,

Haew Suwat Waterfall, 21 Dec 2006, Peng, C-I. & Somprasong, W. 21017 (HAST [2 sheets]).

Notes. Begonia macrotoma is a widespread species. The red striations on the stem and

the bulbils which appear at the sinus of the large (up to 30 cm long) palmatifid leaves

are distinctive. The speeies was originally described from a single specimen, which

according to the protologue has female flowers with three tepals. However all female

material seen by the authors has flve tepals; in all other aspects the specimens agree.

ACKNOWLEDGEMENTS. The authors would like to thank the curators and staff of the

following herbaria: AAU, ABD, BK, BKF, BM, C, HAST, K, E, L, PSU and SING. The Royal

Botanic Garden Edinburgh is supported by the Scottish Government’s Rural and Environment

Research and Analysis Directorate. Field studies of Begonia in Thailand were made possible by

Dr Somran Suddee and the staff of the Forest Herbarium (BKF), Department of National Parks,

Wildlife and Plant Conservation, Ministry of Natural Resources and Environment. This work

was supported by the Newton Fund Researcher Links programme, part of the UK’s official

development assistance programme administered by the British Council. We are very grateful

to two anonymous referees for correcting errors in the manuscript.

References

Beentje, H.J. (2010). The Kew Plant Glossary: An Illustrated Dictionary of Plant Terms. UK:

Royal Botanic Gardens, Kew.

Ding, B., Nakamura, K., Kono, Y, Ho, M.J., Peng, C.-I. (2014). Begonia jinyunensis

(Begoniaceae, section Platy centrum), a new palmately compound leaved species from

Chongqing, China. Bot. Stud. 55: 62.

Doorenbos, J., Sosef, M.S.M. & De Wilde, J.J.F.E. (1998). The sections of Begonia, including

descriptions, keys and species lists (Studies in Begoniaceae VI). Wageningen Agric.

Univ. Pap. 98(2): 1-266.

Hughes, M. (2008). An Annotated Checklist of Southeast Asian Begonia, xii+1 64 p. UK: Royal

Botanic Garden Edinburgh.

A new Begonia from Thailand

107

Hughes, M., Moonlight, P.W., Jara, A. & Pullan, M. (2015). Begonia Resource Centre, http://

padme.rbge.org.uk/begonia (accessed on 26 Aug. 2015).

Phutthai, T., Sands, M. & Sridith, K. (2009). Field surveys of natural populations of Begonia L.

in Thailand. Thai Forest Bull, Bot, Special Issue: 186-196.

Phutthai, T., Hughes, M. & Sridith, K. (2012). A new species of Begonia (Begoniaceae) from

Peninsular Thailand. Edinburgh J. Bot. 69: 287-292.

Phutthai, T., Hughes, M. & Sridith, K. (2014). Begonia kanburiensis (sect. Diploclinium,

Begoniaceae), a new species from Thailand. Thai Forest Bull, Bot. 42: 43^7.

Shui, Y.-M., Peng, C.-I. & Wu, C.-Y. (2002). Synopsis of the Chinese species of Begonia

(Begoniaceae), with a reappraisal of sectional delimitation. Bot. Bull Acad. Sin. 43:

313-327.

Thomas, D.C., Hughes, M., Phutthai, T., Rajbhandary, S., Rubite, R., Ardi, W.H. & Richardson,

J.E. (2011). A non-coding plastid DNA phylogeny of Asian Begonia (Begoniaceae):

evidence for morphological homoplasy and sectional polyphyly. Mol. Phylogenet. Evol.

60: 428^44.

Gardens’ Bulletin Singapore 68(1): 109-124. 2016

doi: 10.3850/S2382581216000089

109

The resurrection of Boesenbergia albosanguinea

(Zingiberaceae) with a new record

for Peninsular Thailand

J.D. Mood^ A.G. Hussain^ & J.R Veldkamp^

Ry on Arboretum, University of Hawaii,

3860 Manoa Road, Honolulu, Hi 96822, U.S A.

boesenbergia@.gmail.com

^Herbwalk Consultancy, Batu 8, Jalan Ayer Hangat,

07000 Langkawi, Kedali Darul Ainan, Malaysia

-''Naturalis Biodiversity Center, RO. Box 9517,

2300 RA Leiden, The Netherlands

ABSTRACT. Boesenbergia albosanguinea (Ridl. ) Loes. (Zingiberaceae) is resurrected from the

synonymy of B. prainiana (King ex Baker) Schltr. It is redescribed in detail with comparative

tables, figures, and illustrations. This species is a new record for Thailand.

Keywords. Boesenbergia prainiana, Langkawi Islands, limestone, Malaysia, Penang

Introduction

During fieldwork in Satun Province for the Flora of Thailand Project in 2012, a

Boesenbergia species [J. Mood & P Vatcharakorn M3291 (BKF)] was found growing

on a limestone outcrop near the sea (Fig. 1 ). Although it was superficially similar to

other large Boesenbergia species in the peninsula, such as B. trangensis K.Larsen and

B. plicata (Ridl.) Holttum, its distinct inflorescence and unusual habitat placed it as an

unknown species. In a follow-up review of all Boesenbergia published from Thailand

and Peninsular Malaysia, only one species appeared to be similar, B. albosanguinea

(Ridl.) Loes. Curiously, that species had been made a synonym of B. prainiana (King

ex Baker) Schltr. by Holttum (1950). As this latter species had been collected on several

occasions by the first author, it was well-known and could be described as a small to

medium sized, 1-2 leaf ginger found growing in wet, shady forest areas on sandstone

or quartz-derived soils. It is not known to occur near the sea or on limestone. In order

to sort this issue of synonymy and try to positively identify the Satun collection, an in-

depth investigation of the taxonomic history of these two species was initiated.

Taxonomic History

In September 1 894 a ginger plant cultivated at the Botanic Gardens Penang flowered

for the first time and was illustrated in excellent detail (Fig. 2). The drawing was

no

Gard. Bull. Singapore 68(1) 2016

annotated by Charles Curtis (1853-1928) as “12-18 in. high, sepals white, linear,

incurved; upper petal white with pink tinge, lower large with dark red margins,

stamens/anthers two celled, Perak, 1894, Nat. size.” Five years later Ridley (1899)

published this ginger as Gastrochilus albosanguineus ['' albo-sanguineus'] with a brief

description. No mention was made of the artwork or specimen. During or soon after this

publication, Curtis added more annotations to the 1894 drawing, ""Gastrochilus albo-

sanguineus Ridl., collected by F.A. Wooldridge on Maxwell’s Flill, Perak, flowered in

the botanical garden of Penang, Sept. 1 894” and with a separate pencil note “drawing

of the type collection”. This latter note refers to a specimen at Kew (K) labelled as the

holotype. Both the drawing and this specimen are annotated with a stylized “A”.

In 1902 several more living plants identified as Gastrochilus albosanguineus

were brought to the gardens by Curtis from Langkawi, from which specimens were

taken (SINGO 155 198, STNGO 155248, SINGO 155266) and a watercolour drawn by M.

Hussain (Fig. 3). Five years later, Langkawi was added to the distributional record of

G. albosanguineus based on this material (Ridley, 1907).

Although Kuntze (1891) replaced Gastrochilus Wall, with Boesenbergia

Kuntze, as Gastrochilus Wall, is a later homonym of Gastrochilus D.Don (Don, 1 825),

some botanists of the day, including Ridley and Curtis, did not accept this change

and continued to use Gastrochilus. It was not until Loesener (1930) that Gastrochilus

albosanguineus was eventually moved to Boesenbergia albosanguinea (Ridl.) Loes.

Holttum (1950) later reviewed the Boesenbergia of Peninsular Malaysia. He decided

thdiX Boesenbergia albosanguinea was the same as B. prainiana and discussed a number

of reasons why Ridley’s taxon is just “re-describing” B. prainiana. Among them were

minor inconsistencies in plant/stem height of the type as compared to the drawing,

calling the longer stem length (noted in the protologue) as “maybe due to conditions

of cultivation.” His conclusion was that “There is no other clear distinction from B.

prainiana.’’" It should be noted that in this discourse, Holttum erroneously uses ""B.

albo-marginata"" tliroughout (rather than B. albosanguinea) to include the holotype

which he cites as “Maxwell’s Hill, per Wooldridge, cult. Penang (type of G albo-

marginata)."" There is no doubt to which species Holttum is referring (based on the

references) and this error is to be regarded as a slip of the pen and not a nomen novum.

Boesenbergia prainiana was first described as Kaempferia prainiana King

ex Baker [Kings collector = Kunstler 726 (CAL, K, SING)] from Goping, Perak,

Malaysia (Baker, 1890). Ridley (1899) transferred it to Gastrochilus Wall, and later

Schlechter (1913) to Boesenbeigia. Holttum’s (1950) description added considerable

detail to the earlier protologue, as did the several publications by Ridley (1899, 1907,

1924).

Materials and Methods

Soon after the collection of J. Mood & P. Vatcharakorn M3 291, an investigation was

begun to study the taxonomic history, specimens, illustrations and photographs from

Malaysia and Thailand of Boesenbergia albosanguinea and B. prainiana. In August

Resurrection of Boesenbergia albosanguinea

111

Fig. 1. Boesenbergia albosanguinea (Ridl.) Loes. A. Mature plants on limestone. B.

Inflorescence and flower. C. Dissected inflorescence with flower. D. Anther with staminodal

cup. E. Flower dissected. F. Rhizome with roots. From J. Mood & P Vatcharakorn M3 291.

(Photos: J. Mood)

2013 another visit to Satun Province, Thailand was undertaken to reassess the coastal,

limestone habitat and look for more populations of B. aff albosanguinea. In addition,

BK and BKF herbaria were visited to study and photograph specimens, as was SING

herbarium later in March 2014. With considerable information in hand, a final fact-

finding trip was made to Langkawi Islands, Malaysia in August 2015 to locate, study

112

Gard. Bull. Singapore 68(1) 2016

W A / •

J! ■

^(,.ljti£^X^ ij ^.^uv (TU

Fig. 2. Boesenbergia albosanguinea (Ridl.) Loes. Ink line drawing by an unknown artist (1894)

at Penang Botanic Gardens. Reproduced with permission of the Singapore Botanic Gardens.

Resurrection of Boesenbergia albosanguinea

113

Ctjjrtnau^

'Aa»m mf

//" ,

O^0t£*»£u P.

.t4*

y. 9- o a.

Fig. 3. Boesenbergia albosanguinea (Ridl.) Loes. Ink line drawing with watercolour by Md.

Hussain at Penang Botanic Gardens (1902). Reproduced with permission of the Singapore

Botanic Gardens.

114

Gard. Bull. Singapore 68(1) 2016

and document any Boesenbei^ia aff. albosanguinea populations which might remain,

as over 100 years had passed since the last known collections. At the same time, any

other Boesenbergia species encountered were documented, with special emphasis on

finding B. prainiana and, if so, in what habitat.

Results

The isolectotype of Boesenbergia prainiana in Singapore Botanic Gardens

(SING0044027) is a two-leafed plant, c. 30 cm tall with erect leaves and a very narrow,

cylindrical inflorescence c. 20 cm. long. The plant is tufted with only a c. 3 cm stem

made up of a bladeless sheath and two leaf sheaths. No flowers are present. Across

Peninsular Malaysia and adjoining Peninsular Thailand, considerable variation can

be seen in plant height, leaf size/shape and inflorescence length/shape. Unmentioned

in any known description is that this species is evergreen and as such has a growth

habit distinct from deciduous Boesenbergia species such as B. plicata. With one to

two thick leaves per element, Boesenbergia prainiana is similar to B. minor (Baker)

Kuntze, another evergreen ginger from Perak. Since evergreen Boesenbergia species

can grow throughout the year, plant elements do not require a large foliar area for

perennation and are often unifoliate with the inflorescence coming directly off the

rhizome, clasped by a bladeless sheath and one leaf sheath (Fig. 4). Due to unknown

factors, occasionally up to four leaves can occur [Corner 302Q9 (SING)]. The leaf

number aside, stem length is c. 3 cm in material measured from Malaysia and Thailand.

To further address Holttum’s synonymy, comparisons of both living fertile

plants and herbarium specimens of the two species were made. The most obvious

difference is vegetative, as Boesenbergia albosanguinea has a distinct stem made up

of tightly appressed leaf sheaths with 4-6 (rarely 2) thin-textured leaves, while B.

prainiana has a very short stem and 1 or 2 (rarely 3) thick-textured leaves per element.

The inflorescence on the latter is elongate and cylindrical or sometimes slightly

flattened with a short peduncle covered by the sheaths. The many dull green, red-

dotted bracts (up to 22) are loosely appressed and appear puffy. As the inflorescence

matures, the bracts often open outward. In comparison, Boesenbergia albosanguinea

has a lanceolate inflorescence with fewer (up to 16), firm, tightly appressed, shiny

green bracts. Both species have distichous bracts with the rachis fully covered on both

sides.

In a comparative floral test, fresh flowers of Boesenbergia aff. albosanguinea

{J. Mood & P. Vatcharakorn M3291) and of B. prainiana from a Peninsular Malaysian

collection [J. Mood 3395 (BKT)] were compared to the 1894 drawing of the former

(Fig. 2) and a watercolour of the latter (Fig. 4). The artwork and living samples were

easily matched and distinguishable by species. The same flowers were then pressed

and dried to compare with those on the holotype of Boesenbergia albosanguinea

(Fig. 5f). The flower from the Thai collection matched in shape and size (Fig. 5d),

whereas the flower of B. prainiana was smaller with a shorter floral tube and smaller,

vertically oriented lateral staminodes which barely covered the labellum (Fig. 5e). In

Resurrection of Boesenbergia albosanguinea

115

Fig. 4. Boesenbergia prainiana (King ex Baker) Schltr. Ink line drawing with watercolour by

Juraimi at Singapore Botanic Gardens (1948). Reproduced with permission of the Singapore

Botanic Gardens.

116

Gard. Bull. Singapore 68(1) 2016

Boesenbergia albosanguinea, the lateral staminodes initially hide the long anther, so

that it cannot be seen from the top looking down. This appearance is due to the long

staminodal length, broad width and full overlap of the margins. Only later in the floral

development do the broad staminodal apices slightly curl toward the corolla lobes,

opening the entrance and making the anther tip slightly visible from above. Even then,

the result is a nairow, tubular entrance for pollinators, something mentioned by Ridley

(1899) and seen in the drawings (Figs. 2, 3, 7). Conversely, in Boesenbergia prainiana

the lateral staminodes are shorter, naiTower, acute at the apices, and only overlap at

their bases. Upon opening of the flower, this morphology allows for over half the anther

length to be seen from the top, providing a broad, rounded, top entrance for pollinators.

The top of the anther and connective is glabrous and shiny with glandular hairs only

along the thecae margins. This is an important diagnostic character state as compared

with Boesenbergia albosanguinea which is completely covered with glandular hairs

on both anther and connective. Thus the basic floral differences between these two

species rest in the length of the floral tube, labellum shape/length, lateral staminode

shape/length and anther length/vestiture (Table 1).

Table 1. Comparison of Boesenbergia albosanguinea (Ridl.) Loes. and B. prainiana (King ex

Baker) Schltr. from living plants.

Character

B. albosanguinea (Thailand)

B. prainiana (Malaysia)

Plant height

To c. 60 cm, not tufted

To c. 40 cm, tufted

No. of leaves

(2-)4-6

1-2 (-3)

Leaf longevity

Deciduous

Evergreen

Petiole length

2-11 cm

2-15 cm

Lamina

c. 38 X 15 cm

c. 24 X 7 cm

Infloreseence

To c. 20 X 2.5 cm, lanceolate to

fusifonn, flattened

15-23 X 2 cm, oblong to

cylindrical, slightly flattened

Bracts

Shiny green, tightly appressed

Dull green, red-dotted, puffy,

later loosely imbricating

Floral tube

c. 3.7 cm long

c. 2.5 cm long

Lateral staminodes

Rotund, c. 20 X 12 mm

Obovate, c. 7 x 2 mm

Labellum

Saccate, c. 3 x 1.7 cm

Saccate, c. 2 x 1.2 cm

Labellum apex

Bilobed, margin frilled

Entire, margin plain

Labellum colour

White with dark red pattern

White with a dark red pattern

Anther

c. 13 mm long, adaxial glandular

hairs throughout

c. 7 mm long, glandular hairs

on the margins of thecae only

Flowering

July to mid-October

May-November

Ecology

Limestone outcrops, seashore

Evergreen forest, mountains

Elevation

5-20 m

nomially >100 m

Resurrection of Boesenbergia albosanguinea

117

Fig. 5. Species comparison. A. Boesenbergia prainiana (Baker) Schltr. (left), B. albosanguinea

(Ridl.) Loes. (right). B. B. prainiana inflorescence and flower. C. Boesenbergia prainiana,

(left), B. albosanguinea (right). D. Pressed and dried flower of B. albosanguinea {M3291).

E. Pressed and dried flower of B. prainiana {M2 172). F. Pressed and dried flower of B.

albosanguinea (holotype). (Photos: J. Mood)

When Ridley (1899) published Boesenbergia albosanguinea, he classified the

known Boesenbergia species into three “groups” based on the position and source of

118

Gard. Bull. Singapore 68(1) 2016

the inflorescence: Acrcinthi (spike borne on the top of a leafy stem), Mesanthi (centre

of the leaf tuft), and Exanthi (outside the leaf tuft). Boesenbergia albosanguinea,

along with the type of the genus, B. puJcherrima (Wall.) Kuntze, was placed in the

first group, while B. prainiana was placed in Exanthi with species now considered

as Scaphochlamys Baker. Holttum made no mention of this classification which,

although now rather outdated, has some diagnostic importance. The character

state of having a significant stem vs little or none at all is quite important. Clearly

Boesenbergia albosanguinea is caulescent (stemmed) as can be seen in the drawings

(Figs. 2, 3), although on the holotype, which is a poor specimen, it is less distinctive

due to separation of the leaves and lack of inclusion of the full stem (Holttum, 1950).

If the 1 894 drawing is compared to this type (supposedly drawn from the same plant),

it is initially hard to see distinctive similarities except in the leaves. If this drawing is

compared to the isolectotype of Boesenbergia prainiana at SING, the caulescent vs

acaulescent (tufted) character state is very evident. If the inflorescences of the types of

both species are compared with the 1 894 drawing (Fig. 2), they are very similar, but

differ in length and number of bracts.

The historic materials aside, if well-prepared and fully documented herbarium

specimens of the two species are compared, two additional identifying clues are the

date of collection and location/ecology. Boesenbergia prainiana, being evergreen,

occurs primarily in wet, evergreen forest on sandstone or quartz derived soils while

B. albosanguinea is deciduous, goes dormant for c. 5 months of the year and has only

been found near the sea on shaded, limestone outcrops. Consequently, the latter cannot

be collected November to March, whereas the former can be collected year-around.

It was also stated that the original collection area of B. albosanguinea was Maxwell’s

Hill (c. 1000 m), far inland from the sea. The type locality of Boesenbergia prainiana

is also in Perak. Cun'ently there are no other Boesenbergia species known from the

Thaiping Hills area that have a morphology similar to B. prainiana, with the exception

of Boesenbergia minor.

So how is this incongruence of habitat (mountain vs coastal) explained if

Boesenbergia albosanguinea is synonymous with B. prainiana as Holttum stated?

Firstly, the holotype of Boesenbergia albosanguinea was not collected in the field

on Maxwell’s Hill, but was from a cultivated plant in the botanic gardens of Penang.

Supposedly this plant was originally given to the gardens by Theodore A. Wooldridge,

an orchid collector who employed local people to make collections. There are many

avenues for eiTor, starting from the time of collection to months later when it flowered

in Penang, e.g., labelling, accession records, etc. At least four years prior, Curtis had

already collected at least one specimen in Langkawi [Curtis 2677 (SING)]. Given

Curtis’s love of horticulture and previous employment from 1 878-1884 as a collector

for the horticultural company Veitch & Son, England, he most likely always collected

living plants for the gardens. Consequently, it seems that when the drawing was made

and then the holotype processed from the same plant, the resultant components were

not taken from Wooldridge’s donated plant, but from a different plant collected in

Langkawi by Curtis. As Ridley was writing his description prior to publication (Ridley,

1899), it is quite likely he saw living plants at the Penang Botanic Gardens as he stated

Resurrection of Boesenbergia albosanguinea

119

Fig. 6. Boesenbergia albosanguinea (Ridl.) Loes. A. A.G. Hussain with wild plants on

limestone at Pulau Langgun, Langkawi. B. Mature plant population. C. Flowering plant. D.

Flower in profile. E. Flower showing anther. (Photos: A.G. Hussain)

“an exceedingly similar if not identical plant occurs also in Langkawi {Curtis 2677)''

an observation difficult to deduce from herbarium specimens alone. Note: Curtis made

eight explorations to Langkawi Islands 1888-1902 (http://www.nationaalherbarium.

nl/FMCollectors/C/CurtisC .htm) .

As a final comparison of materials, in 2015 the second author photographed

(Fig. 5 & 6) and collected plants of Boesenbergia albosanguinea [Ghani s.n. (KEP)]

on Palau Langgun, a small, limestone island off the northeast coast of Langkawi Island.

Based on AGH’s many years of local collecting, this is the only location where this

120

Gard. Bull. Singapore 68(1) 2016

species has been found in recent times. Further, the only other Boesenbergia species

known to occur on Langkawi and the surrounding islands are B. curtisii (Baker) Schltr.

and B. plicata.

Taxonomy

=Boesenbergia albosanguinea (Ridl.) Loes. in Engler & Prantl, Nat. Pflanzenfam., ed.

2, 15a: 566 (1930). - Gastrochiliis albosanguinea Ridl., J. Straits Branch Roy. Asiat.

Soc. 32: 109 (1899) {''albo-sanguinea''). - Type: Specimens from a plant cultivated

at Penang Botanical Gardens, Sep 1894, "'Wooldridge'' [but more likely Curtis] s.n.

(holotype K! [K0()0255404]). (Fig. 1-3, 5-7; Tab. 1, 2).

Boesenbergia prainiana auct. non (Baker) Schltr.: Holttum, Gard. Bull Singapore 13:

111 (1950).

Deciduous, perennial herb up to 60 cm, tightly clumping; rhizome with multiple

elements developing linearly and vertically, c. 3 cm tall, 1 cm diam., externally

yellowish white, internally with two concentric rings, both dark yellow, new rhizome

elements produced at some nodes; roots fleshy to 22 cm long, 4 mm diam., white,

short root hairs along the full length, swollen and fusifonn at the tips, these c. 5 x

1 cm, white, surface smooth, root hairs along the full length; fibrous roots few from

the rhizome to 27 cm long. Stems five to numerous, up to 7 cm long, base ovoid, c.

2 cm diam., without bladeless sheaths (cataphylls), with (rarely 2) 4-6 lealy sheaths

6-16 cm long, dark red, changing to green towards the ligule, surface finely veined,

glabrous. Leaves (rarely 2) 4-6, lower leaves in pairs, nearly opposite; petiole 2-11

cm, longer on upper leaves, green, glabrous; ligule bilobed, each lobe triangular, 2-15

mm long, shortest on upper leaves, translucent, glabrous, from the ligule base a red,

hyaline margin extends down the leaf sheath 2-7 cm long; lamina elliptic to ovate, 15

X 9 cm (lower) to 38 x 15 cm (upper), base rounded, apex acuminate, adaxially veins

raised, medium shiny green, glabrous, abaxially convex between the veins, light green,

glabrous. Inflorescence tenninal, clasped between the sheaths, peduncle short, c. 8 mm

diam.; spike broadly lanceolate to slightly fusifonn, 15-20 long, 2 cm wide, c. 1 cm

thick, exserted for half of its length, ovate in cross-section, dull green with a reddish

centre in the lower one-third. Bracts distichous, overlapping equally on both sides

of the rachis, cymbifomi, lanceolate when opened and flattened, lowest bract sterile,

9 cm long, fertile bracts open to the base, c. 3 x 3 cm, distally decreasing slightly in

size, green in the centre, light red on a wide margin, glabrous; bracteole lanceolate,

curved, open to the base, on the opposite side of the floral tube fi'om the bract, margins

overlap and fully enclose the floral tube, 4.2 cm long, 4 mm diam. at the base, white,

glabrous. Flowers 16-18, one flower per bract, c. 6 cm. long, flowering sequence

basipetalous (from the apex downward), labellum oriented 90° to the bract apex, all

facing the same direction and downward. Calyx tubular, c. 1.1 cm long, translucent

white, glabrous, apex undulate, truncate. Floral tube c. 3.7 cm long, c. 4 mm diam.

Resurrection of Boesenbergia albosanguinea

121

Fig. 7. Boesenbergia albosanguinea (Ridl.) Loes. Ink line drawing with watercolour by Linda

Ann Vorobik (2015). Based on J. Mood & P Vatcharakorn M3291.

122

Gard. Bull. Singapore 68(1) 2016

at base, white, glabrous; dorsal corolla lobe broadly elliptical, cymbiform, c. 25 x 14

mm, apex cucullate, white, glabrous, ventral corolla lobes elliptical, c. 2.5 x l.o cm,

apex cucullate, white, glabrous; androecial cup oriented 90° to the floral tube, c. 8 x 6

mm wide, throat with long hairs. Labellum deeply saccate, orbicular (natural shape),

c. 3 X 1.7 cm wide, c. 3 cm wide (flattened), throat light red, maculate, lip streaked

with dark red to the apex, lobe margins violet with age, glabrous and iridescent on

both sides, apex emarginate, 2-3 nun, overall white; lateral staminodes obovate, c.

2.2 X 1.3 cm, apex slightly reflexed, white, sparsely pubescent, iridescent. Stamen c.

13 imn long, filament c. 3 x 2 mm, white, dorsal, with glandular hairs, thecae c. 11 x

2 mm (each), white, dehiscent full length, connective white, dorsally with glandular

hairs, no anther crest, pollen white. Ovary trilocular, elongate, c. 5 x 2 mm, green,

glabrous; style filiform, c. 6 cm, white, stigma triangular, white, ostiole oval with “v”

notch, glabrous; epigymous glands aculeate, c. 6 mm long, yellow. Fruit not seen.

[Measurements based on living, cultivated material of J. Mood & P. Vatcharakorn

M329J from Thailand].

Distribution. Malaysia: Langkawi, Pulau Langgun; Thailand: Satun Province.

Ecology. Found on limestone outcrops in shaded habitats in close proximity to the sea.

Phenology. Observations in Satun Province and Langkawi indicate that flowering

normally occurs from July to mid-October. Flowers open in the morning and close the

following day.

Etymology. Named for the white and blood-red colour of the labellum.

Specimens examined. MALAYSIA: Kedah: Langkawi, Pulau Langgun, limestone outcrop,

shoreline forest, 10 m asl, 15 Sep 2015, Ghani s.n. (KTP); no exact location, Sep 1890, Curtis,

C. 2677 (SING); no exact locations, cultivated at Penang Botanical Gardens, no dates (SING

[SING0155198, S1NG0155248, SlNGOl 55266]).

THAILAND: Satun: La Ngu, Ko Kabeng, 80 m, Sep 1 999, Phengklai, C. 12105 (BKF), 12106

(BKF); La Ngu, Mu Ko Phetra N.P., 70 m, 10 Sep 2008, Middleton, D.J. et al. 4436 (BKF,

E); La Ngu, Mu Ko Phetra N.P., 30 m, 20 Sep 2010, Middleton, D.J. et al. 5492 (E); near Ko

Phetra, 10 m, evergreen scmb forest on limestone, 06°51.162’N 99°45.78rE, 1 Aug 2012,

Mood, J. & Vatcharakorn, P. M3291 (BKF); near Langu, 54 m, evergreen scrub forest on

limestone, 06°51 .162’N 99°45.781’E, 4 Aug 2013, Mood, J. & Vatcharakorn, P. M3386 (BKF).

Notes. This species as it appears in Satun Province is upright to slightly deeumbent

with thick, multiple stems in a clump, quite similar to the vegetative habit of

Boesenhergia trangensis and B. plicata. The leaves are also plicate, but tend to be

slightly smaller in size with a rounded leaf base. When fertile, it is easily identified by

the very symmetrical, lanceolate inflorescence that is narrow at the base, wider in the

centre and slowly tapered to the apex. The bracts are symmetrical and tightly overlap

on both sides of the rachis. The leaf sheath margins and lower bracts are reddish which

gives the appearance of a broad, red streak on the stem and partially up the centre of

Resurrection of Boesenbergia albosanguinea

123

the inflorescence. The inflorescence protrudes for about half its length out of the leaf

sheaths and maintains a mostly vertical stance. The flowers are pure white with a light

red, narrowed, maculate pattern in the tliroat, and a lip lightly streaked with dark red

especially on the apex margin. Ridley’s comment on the narrow opening for pollination

is due to the large, overlapping lateral staminodes which cover roughly two-thirds of

the labellum length, forming a tube. In the Langkawi populations, the plants tend

to be generally less robust, shorter in height with narrower, shorter leaves. The leaf

sheaths are red but the colour does not extend onto the rachis. The inflorescences are

more cylindrical, less flattened with slightly longer, nan'ower bracts which sometimes

deflex slightly away from the rachis on some plants. Flower shape and colour are

nearly identical to the Thai populations, albeit slightly smaller.

Boesenbergia prainiana (King ex Baker) Schltr., Repert. Spec. Nov. Regni Veg. 12:

316 (1913). - Kaempferia prainiana Baker in Hook.f,, FI. Brit. India 6: 220 (1890).

- Gastrochilus prainianus (Baker) Ridl., J. Straits Branch Roy. Asiat. Soc. 32: 115.

(1899); Ridl., Mat. FI. Malay Penins. 2: 18 (1907); Ridl., FI. Malay Penins. 4: 247

(1924). - Type: Malay Peninsula, Perak, Gopeng, Kinta Valley, King’s Collector

(Kunstler) 726 [cited as “226” by Ridley (1899, 1907)] (lectotype K [K000255402!],

designated here; isolectotypes CAL [CAL0000001005!], K [K000255403!], SING

[SING0044027!]). (Fig. 4; Table 1)

Conclusions

It is concluded that Boesenbergia albosanguinea is distinct from B. prainiana. The

Langkawi collections by Curtis labelled as Boesenbergia albosanguinea are confirmed,

as are those from Satun Province, the latter constituting a new record for Thailand.

The disparity of Wooldridge’s type locality in the mountains of Perak as

compared with the documented coastal collections in Langkawi and Satun Province

cannot be explained by any known historical references such as accession books or

associated Wooldridge memorabilia (pers. com. Singapore Botanic Gardens Library).

The facts remain that Boesenbergia albosanguinea has never again been recorded from

the Thaiping Hills and, ecologically, it is unlikely that species otherwise only known

from coastal limestone sites would also be found in a distinctly different, mountainous

environment. Therefore, the only logical explanation for this inconsistency is that the

plant at Penang used for the holotype, protologue and illustration was not the same

plant Wooldridge collected. More likely it was one of Curtis’s plants collected from

Langkawi between 1890 and 1893.

ACKNOWLEDGEMENTS. We would like to thank the staff of SING and BKF for continued

assistance; Laura Green (K) for specimen assistance; Eric Sinnaya (Langkawi, Malaysia) for

field assistance; The Field Museum, Chicago, USA (F) for use of laboratory facilities; and

Linda Ann Vorobik (Berkeley, USA) for the watercolour artwork.

124

Gard. Bull. Singapore 68(1) 2016

References

Baker, J.G. (1890). Scitamineae. In: Hooker, J.D. (ed) The Flora of British India 6: 198-264.

London: L. Reeve & Co.

Don, D. (1825). Prodromus Florae Nepalensis. Pp. 23-37. London: Gale.

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

106-117.

Kuntze, O. (1891). Revisio Generum Plantarum 2: 683-698. Felix: Leipzig.

Loesener, T. (1930). Zingiberaceae. In: Engler, A. & Prantl, K. (eds) Die Naturlichen

Pflanzenfamilien, ed. 2, 15a: 541-640. Leipzig:Engelmann .

Ridley, H.N. (1899). The Scitamineae of the Malay Peninsula. J. Straits Branch Roy. Asiat.

Soc. 32: 85-184.

Ridley, H.N. (1907). Materials for a Flora of the Malayan Peninsula 2: 1-63. Singapore:

Methodist Publishing House.

Ridley, H.N. (1924). Flora of the Malay Peninsula 4: 233-285. London: L. Reeve & Co., Ltd.

Schlechter, R. (1913). Die Gattungen Gastrochilus Don. und Gastrochilus Wall. Feddes Repert.

Spec. Nov. Regni Veg. 12: 315-317.

Gardens’ Bulletin Singapore 68(1): 125-137. 2016

doi: 10.3850/S2382581216000090

125

Boesenbergia siphonantha (Zingiberaceae),

a new record for Thailand and Vietnam

with notes on the molecular phylogeny

J.D. Mood^, H.D. Tran^, J.F. Veldkamp^ & L.M. Prince^

^Lyon Arboretum, University of Hawaii, 3860 Manoa Road,

Honolulu, Hi 96822, U.S.A.

boesenbergia@ginail.com

^National University, University of Science,

Ho Chi Minh City, Vietnam

^Naturalis Biodiversity Center, RO. Box 9517,

2300 RA Leiden, The Netherlands

"^The Field Museum, Department of Botany, 1400 S Lake Shore Dr.,

Chicago, IL 60605, U.S.A.

ABSTRACT. Boesenbergia siphonantha (King ex Baker) M.Sabu, Prasanthk. & Skomick.

(Zingiberaceae) is newly recorded from Thailand and Vietnam. Its nomenclatural history,

taxonomy, and molecular phylogeny are discussed. Figures, a comparative table and a

watercolour illustration are provided.

Keywords. Andaman Islands, Gastrochilus, Kaempferia

Introduction

During field work in 201 0 for the Flora of Thailand Project, an unusual Boesenbergia

was discovered growing on and around limestone outcrops under deciduous forest in

Kanchanaburi Province [J. Mood & R Chalermglin M2 05 6 (BKF)]. The plant with four,

deeply plaited laminae on long petioles was highlighted by a terminal inflorescence

and a single flower with a shiny lavender lip and red throat. Except for the colouration,

the floral appearance was reiTiiniscent of Boesenbergia longiflora (Wall.) Kuntze,

due in part to the long floral tube and saccate labellum. On further examination of

other plants in the Kanchanaburi population, numerous radical inflorescences were

observed. Except as an abnormal occurrence, bi-positional flowering, defined here as

radical and terminal inflorescences on the same element, is quite rare in Zingiberaceae.

However, it is not without precedent as Boesenbergia prainiana (Baker) Schltr., B.

tenuispicata K.Larsen, and B. trangensis K.Larsen consistently exhibit this habit.

Bi-positional flowering was later observed in two other populations, a second site in

Kanchanaburi [J. Mood & P Vatcharakorn 3081 (BKF)] and in Tak Province [/. Mood

& R Vatcharakorn 3366 (BKF)].

126

Gard. Bull. Singapore 68(1) 2016

In 2008, preceding the Thai collections, the second author discovered an

unidentified Boesenbergia species at Cat Tien National Park, Dong Nai Province,

Vietnam [Tran et ah 125 (SING, VNM)] during MSc research at the Vietnam National

\ -s

University, Ho Chi Minh City (Tran, 2009). Later, photos of this plant were sent to JDM

for possible identification, along with a leaf tissue sample for addition to the authors’

ongoing molecular phylogenetic research on Boesenbergia. Although the taxon could

not be identified immediately, the molecular results showed that it grouped in the same

clade as the three aforementioned Thai collections. Later, moiphological comparisons

also indicated close similarity except for plant stature with the Thai collections being

taller (Table 1). During further study, Boesenbergia specimens at AAU, BK, BKF, C,

CAL, CMU, CMUB, E, K, L, P, SING, QBG, VNM and W were surveyed. The search

produced three unidentified, morphologically similar collections from Thailand and

one from northern Myanmar, Surprisingly, an Andaman Islands species, Boesenbergia

siphonantha (King ex Baker) M.Sabu, Prasanthk. & Skomick., was also vei^ similar.

Boesenbergia siphonantha is considered to be a rare, insular endemic from

the evergreen and semi-deciduous forests of the Andaman Islands, India. It was

described by Baker (1890) as Kaempferia siphonantha based on Kings Collector 572

(K). This and three similar specimens deposited in CAL were collected by Kunstler

on 21 July 1884 in the Andaman Islands during exploration for George King, then

Director, Royal Botanic Garden, Calcutta. It was named to describe the flower’s very

long floral tube. For reasons unknown. Baker did not associate this new taxon with

Boesenbergia longiflora which also had radical inflorescences and flowers with a

long floral tube. Perhaps his statement in the Gardeners' Chronicle (Baker, 1894: 34)

shows his sentiment: ''Gastrochilus [Boesenbergia (Kuntze, 1891)] is hardly worth

separating as a genus Ifom Kaempferia''. Consequently he classified it as a Kaempferia

L. along with six others which had little similarity to this new species. Of these, only

Kaempferia parvula King ex Baker and K. elegans Wall, have withstood the test of

time and remain in Kaempferia.

Considering this information, the Thai and Vietnamese specimens were compared

to the lectotype, protologue (Baker, 1890), and later descriptions of Boesenbergia

siphonantha (Sabu et al., 2004; Kumar et al., 2010), all of which compared favourably.

To further verify the tentative identification, a complete set of photos of the Thai

v

collections was sent to J. Leong-Skomickova (SING) who had firsthand knowledge of

this species. Although some differences were noted and discussed, it was eventually

agreed that the non-Andaman collections were representative of Boesenbergia

siphonantha sensti lato. Unfoitunately, at the time, this conclusion could not be

supported phylogenetically since a tissue sample of Boesenbergia siphonantha from

the Andaman Islands was not available. Recent molecular research by Aishwarya et

al. (2015), which included Boesenbergia siphonantha, later allowed for inclusion of

partial data here. The molecular results support the decision to identify the Thai and

Vietnamese taxa as Boesenbergia siphonantha.

Boesenbergia siphonantha, new record for Thailand and Vietnam

127

Materials and Methods

Field research. In 2008, HDT collected a voucher with silica-dried leaf tissue from a

single population of Boesenbergia siphonantha in Vietnam. Between 2010-2015, JDM

collected B. siphonantha vouchers with silica-dried tissue samples from three different

populations in Thailand. All of the populations were thoroughly studied, documented,

and photographed. Ex situ nursery cultivation was accomplished in Thailand with

several living collections, providing additional information.

Molecular phylogeny. The above tissue samples were used in this analysis. Additional

sequences providing the scaffold were taken from the authors’ ongoing research.

Boesenbergia siphonantha Sabu 73-1 is not available in GenBank but a partial tmK

sequence, con*esponding to the matK genetic fingerprint region, was generously

provided by M. Sabu (University of Calicut, India). ITS data were not available for

that sample. A complete list of samples and GenBank accession numbers are provided

in Appendix 1 .

DNA extraction and analytical methods follow Mood et al. (2013) and will

only be summarised here. The entire plastid trnK region was amplified. Amplification

was done in two parts, the first using IF and 1235R pruners and the second using mlF

and 2R. The nuclear ribosomal ITS (nrlTS) region was amplified using the 18S-F

and 26S-R primers (Prince, 2010). Data were collected on an ABI Genetic Analyzer

(ThermoFisher Scientific) and trace files for each specimen were edited in Sequencher

v4.9 (Gene Codes Corporation, Ann Arbor, Michigan, USA). Consensus sequences

were exported from Sequencher for manual alignment in Se-al (Rambaut, 1996). Areas

of ambiguous alignment were identified and analyses were conducted both including

and excluding these data. Maximum parsimony analyses were run in PAUP* (version

4.0b 10; Swofford, 2002) for each genomic data partition, firstly independently and

later in combination. Branch and bound search methods were conducted in each case

using “farthest” stepwise addition, saving a maximum of 100,000 trees. Branch support

was also estimated using Branch and Bound parsimony bootstrap (BS) in PAUP*.

Results

The ITS analyses produced eight shortest trees based on 97 potentially phylogenetically

informative characters (PPIC) with a consistency index of 0.6703 and a retention

index of 0.7980 (excluding uninformative characters). The trnK analyses, with only 35

PPIC, generated 21 shortest trees (consistency index=0.7400, retention index=0.8750).

The combined partition matrix yielded 132 PPIC and generated 18 shortest trees

(consistency index=0.6598, retention index=0.5231) when the sample Sabu 73-1

(partial tmK sequence only) was excluded from the analyses.

The ITS partition included four samples of Boesenbergia siphonantha {M2056,

M3081, M3 366, and T123), while the trnK partition included those already mentioned,

plus a partial sequence from Sabu 73-1. Analyses of all data partitions, alone or in

128

Gard. Bull. Singapore 68(1) 2016

combination, group the samples of Boesenbergia siphoncmtha together in a strongly

supported monophyletic clade (100% bootstrap support for ITS, 98% for trnK, and

98% for the combined analyses). Exclusion of the partial trnK sequence for Sabu 73-1

resulted in a clade bootstrap value of 100%. The sample Sabu 73-1 is identical to the

other four samples for the region sequenced. Characters supporting the monophyly

include a large number of autoapomorphic substitutions distributed across the coding

and non-coding regions (3 from the ITSl, 5 from the ITS2, 1 from the 5' trnK-matK

intergenic spacer, and 4 from the matK coding region). A unique 3 base pair insertion

was also observed in the 5 'trnK-matK intergenic spacer.

For simplicity, only a single phylogeny is shown here; a phylogram of one of

the most parsimonious trees from the combined data analyses where gaps are treated

as missing data, and the sample Sabu 73-1 was excluded (Fig. 1). Bootstrap values

are provided above the branch. The ITS analyses further resolved a moderately

supported (86% BS) relationship between the Vietnam sample (T123) and one sample

from Thailand (M3366). The closest infeiTed relatives from the sampling utilised

here were members of the Boesenbergia longiflora clade, but that relationship was

only weakly supported (BS 68%). This contrasts with results of the trnK analyses,

where Boesenbergia albomaculata S.Q.Tong and B. pulcherrima (Wall.) Kuntze were

indicated as closest relatives, again with only weak support (74%).

78

100

98

100

10 changes

96 i-^ B. kerrii

100

B. longiflora

B. collinsii

B. hamiltonii

B. maxwellii

B. kingii

100

B. siphonantha M3366

^ — B, siphonantha T123

B. siphonantha M2056

B. siphonantha M3081

B. pulcherrima

B. albomaculata

B. plicata plicata

B. tenuispicata

B. clivalis

B. rotunda

B. ochroleuca

Kaempferia sp. M2043

Kaempferia parviflora M3087

Fig. 1. Combined analysis of Boesenbergia siphonantha (King ex Baker) M.Sabu, Prasanthk.

& Skomick. using ITS and trnK sequence data. Bootstrap values are provided above the branch

for those with > 50% support.

Boesenbergia siphonantha, new record for Thailand and Vietnam

129

Taxonomy

V

Boesenbergia siphonantha (King ex Baker) M.Sabu, Prasanthk. & Skomick., Rheedea

14: 55 (2004). — Kaempferia siphonantha King ex Baker in Hook.f., FI. Brit. India 6:

222 (1890), — TYPE: India, Andaman Islands, 1884, King’s Collector 372 (lectotype

CAL [CAL0000000916]; isolectotypes CAL [CAL0000000912, CAL0000000913], K

[K000640517]), first step designation by Sabu et al. (2004), second step here. (Figs.

2 - 6 )

Deciduous^ perennial herb 20-60 cm tall, tightly clumping. Rhizome with multiple

elements, development in various directions forming a clonal mat, subteiTanean,

globular, 1 .2-1 .5 cm diam., cream-yellow inside, faintly aromatic, bitter; primary roots

few, fleshy, conical, elongate, to 12 cm long, 1 cm. diam. at the top, white, surface

smooth, short root hairs full length, the terminus swollen, spherical to obovoid, 1-2.5

cm long, 0. 8-2.2 cm diam., white, short root hairs the full length, many fibrous roots

from the rhizome to c. 17 cm long. Stems c. 5 cm long, c. 1 .1 cm diam., oval in cross

section, leafless sheaths 1-2, deeply corrugate, green or purple-red. Leaves 4-7 per

stem; leaf sheaths 10-21 cm long, finely ribbed with white blotches, many scattered

hairs, dark green, margin brown; ligule bilobed, lobes linear to rounded, 1-6 mm long,

light green or translucent turning pale brown, glabrous or with few hairs; petiole 3-12

cm, ribbed, green, glabrous; lamina oblong-lanceolate or ovate 10-26 x 4-12 cm,

base cordate, sometimes oblique, apex acute or acute-acuminate, veins 3-7 mm apart,

adaxially dark green, glabrous, abaxially lighter green or purple, glabrous or slightly

pubescent. Inflorescence radical and tenninal, the radical inflorescence attached to

a rhizome near the stem base, the terminal inflorescence between the leaf sheaths,

sometimes shortly exserted, flowering sequence basipetalous; peduncle 1-3 cm long,

narrow, slender, branched or unbranched; spike cylindrical, 4-6 cm long, c. 0.4 cm

diam., base white, apex dark purple-brown, ribbed; bracts 2-8, distichous, ovate-

lanceolate to linear, 3.5-5 x 0.5-1 cm, white, green or green with maroon tinge at

the apex, glabrous, overlapping equally on both sides of the rachis; bracteole tubular,

1-2.9 cm long, c. 0.1 cm diam., glabrous or pubescent, white or cream, translucent,

apex 2 or 3-toothed, laterally split 1-3.6 mm. Flowers 2-8 per inflorescence, one per

bract, 7- 1 2 cm long, erect; calyx tubular, c. 6 x 1,5 mm, translucent white or white with

tiny maroon dots, apex tri-dentate, c. 3 mm longitudinal slit; floral tube cylindrical,

6.5-9 cm long, c. 2 mm diam., white, glabrous; dorsal corolla lobe triangular to

lanceolate, cymbifonn, 1.2-1. 3 x 0.4-0.6 cm, sometimes mucronate, cream, glabrous;

lateral corolla lobes linear, c. 1 .3 x 0.4 cm, cream, glabrous, veins translucent, margin

translucent, all lobes deflexed 1 80° away from the labellum at maturity; androecial

tube 4-6 mm long, c. 4 mm wide, throat with few hairs; labellum deeply saccate,

ovate to obovate (in live plants), 2 x 0.5-1 .7 cm, cream-white, yellowing with age,

throat and medial line bright red, maculate, colour broadening outward, changing

in last third to dark pink, maturing dark purple or distal half violet with reddish

bands toward the centre, externally covered in short glandular hairs, margin slightly

undulate, sometimes crumpled, apex irregular, emarginate; lateral staminodes obovate

130

Gard. Bull. Singapore 68(1) 2016

Fig. 2. Boesenbergia siphonantha (King ex Baker) M.Sabu, Prasanthk. & Skomick. A.

Flowering plant. B. Flower. C. Flower disseetion. D. Seed with aril. (Photos: J. Leong-

Skomickova of an ex situ plant from the Andaman Islands)

to oblong, 0.8-3 x 0.4-1 cm, closely appressed with the labellum towards the base,

cream-white, dorsally covered with short, glandular hairs or glabrous, apex revolute.

Stamen c. 7 mm long; filament 1-4 mm long, white, glabrous; anther elliptical, pale

yellow, darker towards the tip, glabrous, thecae 5-6 x 3-3.7 mm (each) at opening,

5-6 X 4^.7 mm at maturity, rotating away from labellum 90°, dehiscent full length.

Boesenbergia siphonantha, new record for Thailand and Vietnam

131

Fig. 3. Boesenbergia siphonantha (King ex Baker) M.Sabu, Prasanthk. & Skomick. from

Kanehanaburi, Thailand. A. Plants in situ. B. Flower on a radieal infloreseenee. C. First day

flower. D. New flower on a terminal infloreseenee. From J. Mood & R Chalermglin M2056.

(Photos: J. Mood)

white, light brown at senescence, connective white, anther crest absent, pollen white.

Ovary trilocular, 3-6 x 2 mm, placentation axile, white, glabrous, shiny; style filiform,

to 7.5 cm long, white; stigma white, ostiole ciliate; epigynous glands linear, two, 4-4.5

mm long, yellowish or pale cream, apex acute. Fruit a capsule, elliptical, c. 1 cm

long, trilocular, white with red spots throughout, darker on the interior, dehiscence

132

Gard. Bull. Singapore 68(1) 2016

loculicidal, valves rolling outward into coils; seed ellipsoidal, brown, hirsute, apex

yellowish, funiculus present, yellowish, aril medusa-form, yellow-white, translucent.

(Note: This description is based on the protologue and collections from Thailand and

Vietnam.)

Distribution & Ecology. Andaman Islands: Moist, deciduous and inland evergreen

forests on humus rich soil, 5^5 m elevation. Thailand: On and around limestone

outcrops in deciduous, secondary forest with bamboo, medium to heavy shade at c.

70-800 m elevation. Vietnam: Deciduous Lagersti^oemia L. forest on lateritic rocks.

Phenology. See Table 1 .

Additional specimens examined. THAILAND: Mae Hong Son: Baan Bo Khai, 800 m, 12 Sep

1999, Srisanga, P. & Pujf, C. 1067 (QBG); Baan Bo Khai, 800 m, 12 Sep 1999, Srisanga, P &

Puff, C. 1069 (QBG); Tak: N. of Ban Ta Song Yang, 1 7°32.691 ’ N 97°56.291 ’£,180 m, 28 Jul

Table. 1. Comparison of Boesenbergia siphonantha (King ex Baker) M.Sabu, Prasanthk. &

Skomick. from three geographical areas

Character

India

(Sabu et al. 2004)

Thailand

{M2056)

Vietnam

{Tran 123)

Plant height

20-30 cm

50-60 cm

c. 30 cm

Petiole length

3-6 cm

10-12 cm

4-6 cm

Lamina dimensions

10-15 X 4-7 cm

23-26 X 10-12 cm

12-15 X 8 cm

Inflorescence (radical)

Branched

Unbranched

Inflorescence (termmal)

Unbranched

Flowers per inflorescence

4-8

2-5

2 or 3

Flower length

10.5-12 cm

7-10.5 cm

c. 9.5 cm

Labellum

c. 20 X 5-17 mm

c. 21 X 11 mm

c. 20 X 14 mm

Flowers per day

Several

One

Floral display position

Above the lamina

Below the lamina

Intermediate

Phenology

F ebruary-August

July-August

Boesenbergia siphonantha, new record for Thailand and Vietnam

133

Fig. 4. Boesenbergia siphonantha (King ex Baker) M.Sabu, Prasanthk. & Skomick. from

Kanehanaburi, Thailand. A. Inflorescence opened. B. Flower with calyx, bracteole and bract

removed. C. Stamen with staminodal cup at base. D. Senesced flower with thecae rotated 180°.

E. Rhizome with storage roots. From/. Mood & P. Chalermglin M2 05 6. (Photos: J. Mood)

2013, Mood, J. & Vatcharakorn, P. 3366 (BKF). Kanehanaburi: Sai Yok, Wangkamen, 200

m, 29 Jan 2000, Phengklai et al. 14098 (BKF); W of Chong Sadao 20 km, 14° 12.626’ N, 99°

02.925’ E, 70 m, 21 Aug, 2011, Mood, J. & Vatcharakorn, P 3081 (BKF).

VIETNAM: Dong Nai: Cat Tien National Park, August 2008, Tran et al. 123.

Note: AHuk, A. s.n. collection from northern Myanmar, Jul 1892 (CAL) could also be this

species.

134

Gard. Bull. Singapore 68(1) 2016

Fig. 5. Boesenbergia siphonantha (King ex Baker) M.Sabu, Prasanthk. & Skomick. from Dong

Nai Province, Vietnam. A. Mature plant with terminal inflorescence. B. Mature plant with

radical inflorescence. C. First day flower. D. Dissected inflorescence and flower (Scale in cm).

From Tran et al. 123. (Photos: H.D. Tran)

Discussion

The Andaman Islands were physically connected to the Southeast Asian mainland

along the Arakan Yoma ridge in southwestern Myanmar at the end of the Pleistocene

Boesenbergia siphonantha, new record for Thailand and Vietnam

135

Fig. 6. Boesenbergia siphonantha (King ex Baker) M.Sabu, Prasanthk. & Skomick. Ink line

drawing with watercolour by Linda Ann Vorobik (2015).

136

Gard. Bull. Singapore 68(1) 2016

c. 10,000 years ago (www.worldwildlife.org/ecoregions/im0101). This land bridge

occun‘ed due to ocean levels being c. 100 m lower than at present. Although the

duration of the connection is unknown, it was long enough to allow dispersal of

species to and from the islands, to the point that the present-day flora of the Andamans

has 72% of its species in common with the mainland. Most of the remaining species

are also found in the adjacent Nicobar Islands which have a greater species similarity

to Peninsular Malaysia and Sumatra, (www.woiidwildlife.org/ecoregions/im0101). It

is not known which direction Boesenbergia siphonantha spread during this era of low

sea levels but, based on its close phylogenetic relationship to the B. longiflora clade

of six continental species, most likely its origin is also continental. The moiphological

variations observed among the countiy populations show considerable plasticity in

vegetative form, floristic habit and ecology.

Vegetatively, these differences are not unlike other widely distributed

Boesenbergia species such as B. kingii Mood & L.M. Prince. Floristically, the Andaman

populations are very showy with multiple flowers displayed well above the leaves. In

contrast, the continental plants nomially only produce one flower at a time, hidden

down amongst the leaves. This

significant variation suggests very different pollination strategies which are yet to

be investigated. Finally, although the ecology among country populations varies

considerably, it has been shown from ex situ cultivation experiments in India using

Andaman plants (Kumar et al., 2010) and in Thailand with Thai plants (pers. obs.), that

this species is quite adaptable to a variety of environmental regimes.

ACKNOWLEDGEMENTS. We thank the staff at BKF, QBG, and SING for assistance; Piya

Chalennglin (TISTR), Pramote Triboun (TISTR), and Poonsak Vatcharakom (Thailand) for

field assistance; The Field Museum, Chicago, USA (F) for use of laboratory facilities; Jana

Leong-Skoraickova (SING) for photography of B. siphonantha in the Andaman Islands; and

Linda Ann Vorobik (Berkeley, USA) for the watercolour and plate composition.

References

Aishwarya, K., Vinitha, M.R., Thomas, G. & Sabu, M. (2015). Anew species of Boesenbergia

and rediscovery of 5. rotunda (Zingiberaceae) from hidia. Phytotaxa 197: 186-196.

Baker, J.G. (1890). Scitaminae. In: Hooker, J.D. (ed) The Flora of British India 6: 220. London:

L. Reeve & Co.

Baker, J.G. (1894). New or noteworthy plants: Gastrochilus albo-lutens. Gard. Chron., ser. 3,

16: 34.

Kumar, P.K.M., Sabu, M., Thomas, V.P., Prasanth, A.V. & Mohanan, K.V. (2010). A study of

Island Purple Ginger [Boesenbergia siphonantha (Baker) M.Sabu et al.]- A potential

ornamental ginger of the tropics. Ind. J Bot. Res. 6: 165-170.

Kuntze, O. (1891). Revisio Generum Plantanim 2: 682-698. Leipzig: Felix.

Mood, J.D., Prince, L.M., Veldkamp, J.F. & Dey, S. (2013). The history and identity of

Boesenbergia longiflora (Zingiberaceae) and descriptions of five related new taxa.

Gard. Bull. Singapore 65: 47-95.

Boesenbergia siphonantha, new record for Thailand and Vietnam

137

Prince, L.M. (20 1 0). Phylogenetic relationships and species delimitation in Canna (Cannaceae).

In: Seberg, O., Petersen, G., Barfod, A.S. & Davis, J.I. (eds) Diversity, Phylogeny, and

Evolution in the Monocotyledons, Pp. 307-331. Aarhus: Aarhus University Press.

Rambaut, A, (1996). Se-Al (v2.0all) Sequence Aligtment Editor. http.7/tree.bio. ed.ac.uk/

software/seal/ (accessed on 26 Nov. 2012).

Sabu, M., Prasanthkumar, M.G., Skomickova, J. & Jayasree, S. (2004). Transfer of Kaempferia

siphonantha Baker to Boesenbergia Kuntze (Zingiberaceae). Rheedea 14: 55-59.

Swofford, D.L. (2002). PAUP*: Phylogenetic Analysis Using Parsimony (*and other methods),

vers. 4.0b 10. Sunderland, Massachusetts, USA: Sinauer Associates, Inc.

Tran, H.D. (2009). The Diversity and Ecology of Zingiberaceae Martinov in Southeast

^

Vietnam. Unpublished M.Sc. Thesis. 112 pp. Ho Chi Minh City: Vietnam National

University.

Appendix 1 : GenBank accession numbers for Zingiberaceae used in this study. Identification:

plant sample number [for newly sequenced samples] (collector and voucher number, herbarium),

ITS GenBank number/tniK GenBank number.

Boesenbergia albomaciilata S.Q. Tong: M11C83 {Murata, J. et al. 20050595K, MBK)

KU 1 593 17/KU 159408. Boesenbergia clivalis (Ridl.) Schltr.: M11C127 (Lim, C.K. s.n., UPM)

KU159398/KU 159409. Boesenbergia collinsii Mood & L.M.Prince: Mood, J. 12PJ71 (BK)

JX99275 1/JX9928 12. Boesenbergia kerrii Mood, L.M.Prince & Triboun: Mood, J. & Triboun,

P. 12PI 70 (BK) JX992756/JX99281 7. Boesenbergia hamiltoniiMood, S. Dey, & L.M. Prince:

DeyNU53 (CAL) JX992754/JX992815. Boesenbergia kingii Mood & L.M.Prince: Mood, J.

& Vatcharakorn, P. 12P173 (BK) c2-JX992789, c3-JX992790, c4-JX992791, cJX992792,

c8-JX992793/JX992829. Boesenbergia maxwellii Mood, L.M.Prince & Triboun: Mood, J.

& Triboun, P 12P172 (BK) JX992800, JX992833. Boesenbergia ochroleuca (Ridl.) Schltr.:

M3116 {Mood, J. & Vatcharakorn, P. 3116, BfCF) KU159399/KU159410. Boesenbergia plicata

van plicata (Ridl.) Holttum: Mood, J. cfe Vatcharakorn, P 3177 (BKF) JX992807/JX992840.

Boesenbergia pulcherrima (Wall.) Kuntze: Mood, J. 08P276 (BKF) JX992748/JX992809.

Boesenbergia rotunda (L.) Mansf: M11P27 {Mood, J. 11P27, BKF) KU159400/KU15941 1.

Boesenbergia siphonantha (King ex Baker) M.Sabu, Prasanthk. & Skornick.: M2 05 6 {Mood,

J.& Chalermglin, P. 2056, BKF) KU 1 5940 1/KU 1594 12. Boesenbergia siphonantha (King

ex Baker) M.Sabu, Prasantlik. & Skornick.: M3081 {Mood, J.& Vatcharakorn, P. 3081, BKF)

KU159402/KU159413. Boesenbergia siphonantha (King ex Baker) M.Sabu, Prasanthk.

& Skornick.: M3366 {Mood, J. & Vatcharakorn, P. 3366, BKF) KU159403/KU 159414.

Boesenbergia siphonantha (King ex Baker) M.Sabu, Prasanthk. & Skornick.: Tran & al. 123

(SING, VNM) KU159404/KU159415. Boesenbergia siphonantha (King ex Baker) M.Sabu,

Prasanthk. & Skornick.: Sabu, M. 7i-/(CALI) Not registered in GenBank. Boesenbergia

temiispicata K. Larsen: M3292 {Mood, J. 3292, BKF) KU159405/KU159416. Kaempferia sp. :

M2043 {Mood, J. 2043, BKF) KU159395/KU 159406. Kaempferia parviflora Wall, ex Baker:

M3087 {Mood, J. 3087, BKF) KU159396/KU1 59407.

Gardens’ Bulletin Singapore 68(1): 139-144. 2016

doi: 10.3850/S2382581216000107

139

Boeica ornithocephalantha (Gesneriaceae),

a new species from northern Vietnam

F. Wen' T.V. Do^, X. Hong'*, S. Maciejewski^-’ & Y.G. *

^Guangxi Key Laboratoiy of Plant Conservation and Restoration Ecology

in Karst TeiTaiii, Guangxi Institute of Botany, Guangxi Zhuang

Autonomous Region and the Chinese Academy of Sciences,

CN-541006, Guilin, Guangxi, China

wenfang760608@139.cora

^Gesneriad Conservation Center of China (GCCC),

CN-541006, Guilin, Guangxi, China

^Department of Biology, Vietnam National Museum of Nature,

Vietnam Academy of Science and Technology, 1 8 Hoang Quoc Viet,

Cau Giay, Ha Noi, Vietnam

dovantruong_bttn@yalioo.cora

"^The Key Laboratory of Conseivation and Employment of Biological

Resources of Anhui, Herbarium, College of Life Sciences,

Anhui Nonnal University, Wuhu, CN-241000, Anhui, China

hongxinl989@vip.qq.cora

^The Gesneriad Society, Inc. 1122 East Pike Street, PMB 637,

Seattle, WA 98122-3916, USA

tecml949@gmail.com

Corresponding author: weiyigang@aliyun.com

ABSTRACT. A new species, Boeica ornithocephalantha F.Wen, Y.G. Wei & T.V.Do

(Gesneriaceae), is described from the northern part of Vietnam. This new species is most

similar to Boeica ferniginea Drake, but clearly differs from the latter by the unusual corolla. A

provisional lUCN conservation assessment is provided.

Keywords. Boeica, Gesneriaceae, lUCN conservation assessments, Vietnam

Introduction

The genus Boeica Clarke (Gesneriaceae) was first published by Clarke (1874). It

is a small genus in subfamily Didymocarpoideae, tribe Trichosporeae, subtribe

Leptoboeinae of the Gesneriaceae (Weber et ah, 2013). Currently, there are about

twelve species of Boeica from China, Myanmar, Bhutan, northern India and the

northern part of Vietnam (Wang et ah, 1 990, 1 998; Li & Wang, 2004; Weber, 2004), the

species described most recently being B. glandulosa B.L.Burtt (Burtt, 2001). Boeica

species usually grow in shaded and damp locations with the different species having

variable flowering and fhiiting seasons, thereby often making it a challenge to collect

specimens for identification. Except for Boeica guileana Burtt (Burtt, 1977), which is

140

Gard. Bull. Singapore 68(1) 2016

a rosette plant without distinct stems, the remaining species of Boeica are caulescent,

including the newly described species below.

During a field investigation of the flora of northern Vietnam we collected an

unknown species of Boeica which required further investigation. Plants were grown

at the nursery of Guilin Botanical Garden and the Gesneriad Conservation Center

of China (GCCC) from seeds collected in the Pu Hu Nature Reserve, Quan H6a

district, Thanh H6a province, northern Vietnam. When the plants flowered, the floral

moiphology provided further confinnation that the collection was a new species of

Boeica. Michael Moller, a Gesneriaceae researcher at the Royal Botanic Garden

Edinburgh, also supported this conclusion. We describe and illustrate the proposed

new species here and provide a provisional lUCN conservation assessment following

the guidelines in lUCN (2012).

Taxonomy

Boeica ornithocephalantha F.Wen, T.V.Do & Y.G.Wei, sp. nov.

Most similar to Boeica fermginea Drake but B. ornithocephalantha is characterised

by a shorter petiole (1-1.5 cm long vs 3-5 cm long in Boeica ferruginea), adaxial leaf

blade densely glandular and non-glandular pubescent (adaxial ly appressed puberulent,

abaxially sparsely woolly to glabrescent, woolly to villous along veins in Boeica

ferruginea), peduncle 1 2-20 cm long and sparsely glandular pubescent (5-8 cm and

villous in Boeica ferruginea), bracts 3 (2 in Boeica ferruginea), corolla in lateral view

reminiscent of the head of a bird and outside glabrous (not reminiscent of a bird and

outside sparsely pubescent in Boeica ferruginea), corolla tube 1-1.2 mm long (c. 2

mm long in Boeica ferruginea), and disc inconspicuous or nearly none (circular, c. 0.2

mm high in Boeica ferruginea). - TYPE: Vietnam, Thanh Hoa province, Quan H6a

district, Quan Hoa town, Pu Hu Nature Reserve, alt. 900 m, 18 July 2014, flowering.

Wen Fang WF1407 18-01 (holotype VNMN; isotype IBK). (Figs. 1 & 2)

Herbs, stoloniferous. Stems 8-18 cm long, unbranched or rarely branched, densely

yellowish-brown woolly to villous or pilose when young, mostly glabrescent when

mature, rarely glabrous; rhizomes short, 2-4 cm long, c. 2 mm in diameter. Leaves

alternate, intemodes c. 1 cm long, usually 6-8 leaves clustered at the top of stem;

petiole 1-1.5 cm long, densely brown villous; blades ovate-elliptic to oblong, 10-

18 X 7-10 cm, mature leaf blades usually ascending, apex obtuse to rounded, base

attenuate, margin in*egularly serrate, densely glandular and non-glandular pubescent

on the upper surface, upper surface distinctly bullate, lateral veins 10-13 on each

side, impressed adaxially and prominent abaxially, densely yellowish-brown woolly

on the veins of the lower surface, the rest yellowish-brown villous. Inflorescence

axillary, a dichasium, 12- to 35-flowered or more; peduncle 12-20 cm long, c. 1.2 mm

in diameter, sparsely purplish brown and white erect glandular pubescent, the hairs

varying in length; bracts 3, the larger two narrowly lanceolate, c. 5 mm long, c. 1 mm

in diameter at base, apex acute, margin entire, the remaining one linear, c. 1.5 mm

Boeica ornithocephalantha, a new species from Vietnam

141

Fig. 1. Boeica ornithocephalantha F.Wen, Y.G.Wei & T.V.Do. A. Top view of plant. B. Upward

view of plant. C. Cyme with open flowers. D. Front view of cyme. (Photos: Fang Wen)

long, 0.2-0. 3 mm in diameter at base, apex acuminate, margin entire, outer surface

sparsely purplish brown glandular puberulent, inside nearly glabrous; pedicels 1 .5-4.5

cm long, 1-1.2 mm in diameter, sparsely purplish brown and white erect glandular

pubescent. Calyx divided to the base, 5-lobed, apex green, the rest pinkish purple,

lobes linear lanceolate, 4-5 mm long, c. 0.9 mm in diameter at base, margin entire,

outside reddish purple glandular puberulent, inside nearly glabrous; Flowers slightly

fragrant. Corolla small, 4-6 mm long, c. 5 mm in diameter, outside glabrous, initially

shallowly campanulate, limb inconspicuously 2-lipped, two larger lobes of abaxial

lip indexed from the middle and covering the anthers; corolla tube extremely short,

1-1.2 mm long; adaxial lip 2-lobed at least to middle of lip, lobes nearly equal, 3-4.5

mm long, c. 2 mm in diameter at base, wider but mostly shorter than abaxial lip lobes,

external margin of lobes white, the centre of lobes green, internal margin of lobes

white to pale pink, the centre dark purplish red; abaxial lip 3-lobed, all white, lateral

lobes larger, broadly lanceolate to oblong, apex obtuse, c. 3.5 mm long, c. 1.5 mm in

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Gard. Bull. Singapore 68(1) 2016

Fig. 2. Boeica ornithocephalantha F. Wen, Y.G.Wei & T.V.Do. A. Obliquely view of corollas.

B. Inside of corolla. C. Outside of corolla. D. Pistil and calyx. E. Capsule. (Photos: A, E: Fang

Wen; B-D: Hua-Fei Cen)

diameter at base, the central lobe lanceolate, apex acuminate and indexed, 3.5-4 mm

long, 1.2-1. 5 mm in diameter at base. Stamens 4, free; filaments white, 0.5-0. 8 mm

long, glabrous, adnate to the base of corolla; anthers dark purplish red, cordate, c. 0.5

mm long, 0.4-0.6 mm in diameter at base, dehiscing poricidally; staminode 1, c. 0.1

mm long, dark purple to black, adnate to the base of corolla. Disc inconspicuous or

nearly absent. Ovary ovoid, 1.5-2 mm long, c. 1 mm in diameter, glandular puberulent;

style 5-7 mm long, mostly glabrous but glandular puberulent at the base. Stigma 1,

apex truncate, pale green. Capsule linear, nearly glabrous, 1.5-2 cm long.

Distribution. Only known from Pu Hu Nature Reserve, near En Village, Quan H6a

town, Quan H6a district, Thanh Hoa province, Vietnam.

Habitat. Primarily growing on shaded slopes in valleys above streams, often locally

common, at low altitude. It occurs on rocks covered with humus under evergreen

forest, not prone to human disturbance.

Boeica ornithocephalantha, a new species from Vietnam

143

Etymology. The scientific name is due to the perception by the authors thatthe lateral view

of the mature corolla looks like the head of a bird. The epithet, ''ornithocephalantha'',

is composed of the Greek elements ‘omitho-’ for bird, ‘cephal-’ for head and ‘-antha’

for flower.

Provisional lUCN Conservation Assessment. Vulnerable VU B2ab(ii,iii). This species

is only known from very few collections and details on the size of the population in Pu

Hu Nature Reserve are unknown. We acknowledge that the type locality is protected

from exploitation and development by being in a Nature Reserve, thereby guaranteeing

the plants protected status. Based on our observations during the field work cuiTently

the plants appear to be locally abundant. However, the known habitat is also disturbed

by intentional and unintentional human activity such as for path building. According to

the lUCN red list categories and criteria, the species should be considered Vulnerable

(lUCN, 2012).

Notes. Boeica species are characterised by an indumentum of shiny bristly brown

hairs, a corolla with a short tube and flat limb, four free stamens, and poricidal or

transverse anthers that are coherent at the tips (Pellegrin, 1926; Burtt, 1977; Li &

Wang, 2004). Among the 12 known species, some have a distinctly zygomoiphic

corolla, a very short corolla tube, a spreading corolla limb, and four obvious stamens

(here connivent), traits that Burtt (1977) considered to be ‘primitive’. The new species

possesses some of the above-mentioned morphological traits.

ACKNOWLEDGEMENTS. We thank Dr Michael Moller of the Royal Botanic Garden

Edinburgh for the recognition of this species, and Hua-Fei Cen for the photos. We also

acknowledge the Vietnam Ministry of Agriculture and Rural Development and the Vietnam

National Museum of Nature for granting the collecting permissions. This study was

financially supported by the Guangxi Natural Science Foundation (201 3GXNSFAA01 9071,

2013GXNSFBA019078 & 20I5GXNSFBB139004), Guangxi Forest S & T Projects (Gui Lin

Ke Zi [2014] 27), International S and T Cooperation Projects of Guangxi (Guikehe 1347004-4)

& Guilin (20130412) and the Science Research Foundation of the Guangxi Institute of Botany

(Guizhiye 11005).

References

Burtt, B.L. (1977). Studies in the Gesneriaceae of the old world. XLI. Notes on Boeica and

Didissandra. Notes Roy. Bot. Card. Edinburgh 35: 369-374.

Burtt, B.L. (2001). Flora of Thailand: annotated checklist of Gesneriaceae. Thai Eorest Bull,

Bol 29: 81-109.

Clarke, C.B. (1874). Commelynaceae et Cyrtandraceae Bengalenses. London: Spink & Co.

lUCN (20 1 2). lUCN Red List Categories and Criteria: Version 3. 1. 2nd ed. - Gland, Switzerland

and Cambridge, UK: lUCN,

Li, Z.Y. & Wang, Y.Z. (2004). Boeica. In: Li, Z.Y. & Wang, YZ. (eds) Plants of Gesneriaceae

in China. Pp. 109-112. Zhengzhou: Henan Science and Technology Publishing House.

144

Gard. Bull. Singapore 68(1) 2016

Pellegrin, F. (1926). Les Gesneracees-Cyrtandrees d’Indo-Chine. Bull Soc. Bot. France 73(3):

412^29.

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 1 8: 244-401 . Beijing, China: Science Press and

St. Louis, Missouri, USA: Missouri Botanical Garden Press.

Wang, W.T., Pan, K.Y., Zhang, Z.Y. & Li, Z.Y (1990). Gesneriaceae. In: Wang, W.T. (ed) Flora

ReipiibUcae Popularis Sinicae 69: 125-581. Beijing: Science Press.

Weber, A. (2004). Gesneriaceae. In: Kubitzki, K. & Kadereit, J.W. (eds) The Families and

Genera of Vascular Plants, vol. 7. Dicotyledons. Lamiales (except Acanthaceae inch

Avicenniaceae). Heidelberg: Springer.

Weber, A., Clark, J.L. & Moller, M. (2013). A new formal classification of Gesneriaceae.

Selbyana 31(2): 68-94.

Gardens’ Bulletin Singapore 68(1): 145-172. 2016

doi: 10.3850/S2382581216000119

145

An expansion of the genus Deinostigma (Gesneriaceae)

M. Moller', K. Nishii", HJ. Atkins', H.H. Kong^ M. Kang^

Y.G. WeP, F. Wen', X. Hong‘'& D.J. Middleton'

^Royal Botanic Garden Edinburgh^ 20 A Inverleith Row,

Edinburgh EH35LR, Scotland, U.K.

^Key Laboratory of Plant Resources Conservation

and Sustainable Utilization, South China Botanical Garden,

Chinese Academy of Sciences, Guangzhou 510650, China

^Guangxi Key Laboratoiy of Plant Conservation and Restoration

Ecology in Karst Terrain, Guangxi Institute of Botany,

Guilin 541006, China

"^College of Life Sciences, Anliui Normal University,

Wuhu 241000, China

^Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, Singapore 259569

david_niiddleton@nparks . go v. sg

ABSTRACT. Based on molecular, morphological and cytological studies the previously

monotypic genus Deinostigma W.T.Wang & Z.Y.Li has been expanded to include several

species previously ascribed to Primulma Hance. Deinostigma now comprises seven species,

including one previously placed in synonymy. The new combinations Deinostigma cicatricosa

(W.T.Wang) D.J.Middleton & Mich.Moller, Deinostigma cycnostyla (B.L.Burtt) D.J.Middleton

& H.J.Atkins, Deinostigma cyrtocarpa (D.Fang & L.Zeng) Mich.Moller & H.J.Atkins,

Deinostigma eberhardtii (Pellegr.) D.J.Middleton & H.J.Atkins, Deinostigma minutihamata

(D.Wood) D.J.Middleton & H.J.Atkins and Deinostigma tamiana (B.L.Burtt) D.J.Middleton

& H.J.Atkins are made. Deinostigma eberhardtii is lectotypified. The genus is defined by

a combination of an alternate leaf arrangement, hooked hairs on many plant parts, flowers

with the pedicel inserted at an angle and off-centre on the receptacle, and, where known, a

somatic cliromosome number (2n) of < 36. This new circumscription of the genus expands its

distribution from Vietnam into South China.

Keywords. Molecular phylogeny, ovary morphology, Primulina, taxonomy

Introduction

The genus Deinostigma W.T.Wang & Z.Y.Li in the Gesneriaceae currently has only

one species, D. poilanei (Pellegr.) W.T.Wang & Z.Y.Li from the southern Annamite

range of Vietnam. It was erected as a genus to accommodate the species first described

as Hemihoea poilanei Pellegr. This species was placed in Hemiboea C.B. Clarke due

to its bilocular ovary and the perception that one of the locales was sterile, a trait of

the species in Hemiboea. Wang & Li (1992) argued that it did not belong in Hemiboea

due to having alternate leaves, free bracts, inside of corolla glabrous, filaments broader

146

Gard. Bull. Singapore 68(1) 2016

in the upper half, anthers hairy, anther locules divaricate with apices confluent, disc

absent, and stigma bifid. In further arguments they noted how it differed from other

Asian genera and concluded that a new genus was necessary to accommodate it. Since

then, the status and relationships of Deinostigma poilanei have remained obscure due

to being known from only very few specimens and because relatively little research

has been done on the Gesneriaceae of Vietnam.

During work at the Museum National d’Histoire Naturelle in Paris for an as-yet-

unpublished checklist of the Gesneriaceae of Cambodia, Laos and Vietnam by DJM, it

became clear that Deinostigma poilanei was similar to a number of species from Vietnam

and China that were moved into Primulina Hance from Chirita Buch.-Ham. ex D.Don

by Weber et al. (2011), namely P. cycnostyla (B.L.Burtt) Mich.Moller & A.Weber,

P. cyrtocarpa (D.Fang & L.Zeng) Mich.Moller & A.Weber, P. eberhardtii (Pellegr.)

Mich.Moller & A.Weber, P. minutihamata (D.Wood) Mich.Moller & A.Weber and P.

tamiana (B.L.Burtt) Mich.Moller & A.Weber. None of these Deinostigma-hke species

were included in the molecular phylogenetic analyses by Weber et al. (2011) but were

moved into Primulina based on their previous inclusion in Chirita sect. Gibbosaccus

C.B. Clarke by Wood (1974). All species of the hitherto Chirita sect. Gibbosaccus that

were included in the molecular analyses formed a single highly supported clade with

Primulina tabacum Hance, the type of Primulina (Weber et al., 2011).

Deinostigma poilanei and the species of Primulina similar to it share a number

of moiphological traits that are not found in other species of Primulina. These are

alternate leaf an*angement (opposite or verticillate in the remaining Pnwu/mu species),

hooked hairs on many plant parts but particularly on the pedicels, and flowers with

the pedicel inserted at an angle and off-centre on the receptacle. In several species

the fruit is strongly curved. The alternate leaf arrangement is not particularly clear in

the species with a congested rosette of basal leaves but is nevertheless observable by

only a single leaf emerging from the apical meristem at a time. Differences also exist

in the basic chromosome numbers, with Primulina having exclusively x = 18 (> 100

species counted out of > 150 described) (Moller & Pullan, 2015 onwards) while one

Deinostigma-hke species, P tamiana, has been counted with x = 16 (Christie et al.,

2012). In other features the plants are indeed similar to Primulina. It should also be

noted that these Deinostigma-hke, Primulina species occur on a variety of substrates

but not on limestone and that the majority of the remaining Primulina species occur

on limestone substrates.

In Wang et al. (1998) Chirita cicatricosa W.T.Wang is placed in synonymy

of C. minutihamata D.Wood (= Primulina minutihamata), typified by material from

China and Vietnam respectively. We found that the material from China has longer,

more slender and more falcate fruits and somewhat larger flowers than material fi"om

Vietnam. Coupled with their large disjunct distribution, and pending a revision of the

species, we suspect that they may represent distinct species and treat them thus here

(see also below).

The aim of this study was to investigate the relationship between Deinostigma

and Primulina and clarify the status of the Deinostigma-likQ Primulina species.

An expansion oi Deinostigma

147

Materials and Methods

Plant materials

Herbarium specimens and living collections of Deinostigma poilanei and the

Deinostigma-\\kQ Primulina species were studied in the Royal Botanic Garden

Edinburgh and the Museum National d’Histoire Naturelle in Paris.

Materials for moiphological and phylogenetic analyses were collected in

Vietnam {Deinostigma poilanei, Primulina tamiana) and China {Primulina cyrtocarpa,

Chirita cicatricosa) (Table 1). Data for the phylogenetic analyses were downloaded

from GenBank (including one additional sample of Primulina minutihamata from

China which we are treating here as Chirita cicatricosa). These basically represented

a reduced matrix of Old World Gesneriaceae as presented in Middleton et al. (2015).

Altogether 132 samples (129 species) were included, covering all 32 genera currently

recognised in the subtribe Didymocarpinae of tribe Trichosporeae (Moller et al., 2009,

2011, 2014; Weber et al., 2013; Middleton et al., 2014a, 2015). The sampling included

20 samples of 19 species oi Primulina, including the type, P. tabacum (Table 1). The

phylogenetic trees were rooted on samples of Microchirita (C.B. Clarke) Yin Z.Wang

(Moller et al., 2009, 2011).

DNA extraction, PCR and phylogenetic analysis

Sequences of the nuclear ribosomal internal transcribed spacers (ITS) and the plastid

trnL-V intron-spacer (^mL-F) for four samples, Chirita cicatricosa, Deinostigma

poilanei, Primulina cyrtocarpa, and P. tamiana, were obtained. The extraction of

genomic DNA was carried out using a CTAB procedure (Doyle & Doyle, 1 987, 1 990),

The PCR amplification of ITS and trnL-¥ were perfomied using primers ‘5P’

(5’-GGA AGG AGA AGT CGT AAC AAG G-3’) and ‘8P’ (5’-CAC GCT TCT CCA

GAC TAC A-3’) (Moller & Cronk, 1997) and ‘c’ (5-CGA AAT CGG TAG ACG CTA

CG-3’) and ‘f (5’-ATT TGA ACT GGT GAC ACG AG-3’) (Taberlet et al., 1991),

respectively, run on a Biorad TlOO™ Thermal Cycler (Kernel Hempstead, UK). The

10 pL reactions contained 1 pL lOx NH^ reaction buffer (Bioline, UK), 1 pL dNTPs

(2 inM), 0.3 pL MgCl, (50 inM), 0.4 pL of each primer (10 pM), 5.6 pL ddH^O, 0.1

pL Biotaq polymerase (5U/pL) (Bioline, UK) and 1 .2 pL DNA template. The PCR

thermocycle profile for ITS started with an initial denaturation for 3 min at 94°C,

followed by 30 cycles of 1 min at 94°C, 1 min at 55°C and 1.5 min at 72°C, finished

with a final extension step for 5 min at 72°C. For trnL-¥ it was: initial denaturation for

4 min at 94°C, followed by 30 cycles of 45 s at 94°C, 45 s at 55°C and 3 min at 72°C,

with a final extension step for 10 min at 72°C. PCR products were run on 1% agarose

gels to check for amplification success and quality. PCR amplified fragments were

purified using ExoSAP-IT (Affymetrix, UK) following the manufacturer’s protocol,

and sequenced using the dideoxy chain-termination method. Sequencing samples were

prepared using the BigDye Terminator v3.1 Cycle Sequencing Kit (Thermo Fisher

Scientific, UK) following the manufacturer's recommendations, and sequencing was

earned out by the Edinburgh Genomics sequencing service (University of Edinburgh,

148

Gard. Bull. Singapore 68(1) 2016

UK). Editing and assemblage of sequencing results were perfonned using the programs

Sequencher 4.5 (Gene Codes Coip, Ann Arbor, USA). The newly acquired sequences

were added to the reduced matrix of Middleton et al. (2015) and the matrices realigned

manually. They were subsequently submitted to GenBank (Table 1).

Since the combinability of the ITS and trnL-F matrices, tested with the ILD test

(Farris, 1995a, 1995b), implemented as PHT in PAUP* 4.0al46 (Swofford, 2002),

did not indicate incongruent phylogenetic signals (P = 0.58), the two matrices were

analysed together. The reconstruction of phylogenetic trees by maximum parsimony

(MP) and Bayesian inference (Bl), including the calculation of bootstrap (BS) and

posterior probability (PP) branch support values, were carried out as previously

described (Moller et al., 2009, 2011; Weber et al., 2011; Middleton et al., 2015): the MP

analysis on the combined data was carried out in PAUP* v.4.0bl0 (Swofford, 2002),

on unweighted and unordered characters. Alignment gaps were treated as missing data.

Starting trees were found by parsimony ratchet (Nixon, 1999), in PAUPRat (Sikes &

Lewis, 2001) and PAUP*, and the saved trees further optimised in PAUP*, with both

TBR and Multrees on. Statistical branch support was obtained from 10,000 heuristic

bootstrap replicates each starting with a random addition tree, optimized with TBR on

and Multrees off in PAUP* (Moller & al., 2009, 2011).

The BI analyses were run in MrBayes v.3.1.2 (Huelsenbeck & Ronquist,

2001; Huelsenbeck et al., 2007). Models and parameter prior settings were obtained

independently for the trnL-F, the ITS spacers and 5.8S sequences using MrModeltest

v.2.3 (Nylander, 2004), and were GTR + I + G for trnL-F and the ITS spacers, and

SYM + I + G for the 5.8S gene, respectively, as suggested by the Akaike Information

Criterion (AIC; Akaike, 1974). Five million generations were run in two independent

analyses each with four Markov chain Monte Carlo (MCMC) chains. One tree was

sampled every 1000^ generation (= 5000 trees), and the first 250 trees (5%) discarded

as burn-in, detemiined after plotting the generations against the Log(n) likelihoods

and a majority mle consensus tree constmct with the ‘sumt’ command and posterior

probabilities (PP) obtained in MrBayes.

Cytology

The chromosome numbers for Deinostigma poilanei {R. Rybkovd HB 222) and

Primulina cyrtocarpa {M.Mdller &Y.G. Wei MMO 06-908) (Table 1), were determined

from root tips as previously described (Jong & Moller, 2000; Christie et al., 2012).

For Chirita cicatricosa {ZuUn 131585, China, Guangxi, Shangsi), leaf cuttings were

cultivated in pots at the South Chma Botanical Garden, Chinese Academy of Sciences.

Actively growing root tips were harvested and pretreated in the dark with 0.1%

colchicine and 0,1% 8-Hydroxyquinoline (1:1) at room temperature for 3 hours and

then fixed in 3 : 1 absolute ethanol : glacial acetic acid at 4°C for 2 hours. They were

macerated in a mixture of 1 : 1 IM HCl and 45% acetic acid at 37°C in a water bath

for 45 min, and stained and squashed in 1% aceto-orcein.

An expansion oi Deinostigma

149

Results and Discussion

Phylogeny

The MP analysis resulted in 204 most parsimonious trees with a length of 4123 steps,

Cl of 0.3772 and RI of 0.6797. For the B1 mns, the average standard deviation of

split frequencies was 0.003949 indicating a strong convergence of the two runs.

Furthemiore, a close correlation of the PP values between the two runs was observed,

confirming the reproducibility of the runs.

In both the MP (Fig. 1) and BI (Fig. 2) phylogenetic trees, Primulina s.s.

(excluding Chirita cicatricosa, P cyrtocarpa, andP. tamiana) fonns a highly supported

clade (BS = 100%; PP= 1), and, most importantly, is the sister clade to Petrocodon

with maximum branch support (BS = 1 00%; PP = 1 ). Deinostigma poilanei, together

with Chirita cicatricosa, Primulina cyrtocarpa andP tamiana, fonns a separate clade

(BS = 73%; PP = 0.88), which is sister to the monotypic Metapetrocosmea W.T.Wang

with maxumim branch support (BS = 100%; PP = 1). Although most of the genera

are in highly supported clades, as in previous studies (Mdller et al., 2009, 2011), the

backbone of the phylogenetic tree is not resolved. However, since both Primulina s.s.

and the Deinostigma clade fonn strongly supported sister relationships to other genera

respectively, it is unlikely that the two genera are closely related. Consequently, we

transfer the Deinostigma-Mks Primulina species to Deinostigma and refer to them thus

hereafter.

Morphology

All species are herbs. They are all probably perennial and, judging from the numerous

leaf scars on the stems of some species, they are likely to be deciduous in the dry season.

In particular, Deinostigma cycnostyla has a thick stem with numerous congested leaf

scars. The other species have a relatively thin stem and species such as Deinostigma

minutihamata are mostly procumbent.

All species have an alternate leaf arrangement although this is not so obvious

in Deinostigma cycnostyla where the leaves are very congested. The alternate leaf

arrangement is in contrast to Primulina s.s. where the leaves are opposite or in whorls,

usually of three. The leaf blade is ovate to elliptic with a weakly {Deinostigma tamiana)

to strongly (D. cyrtocarpa) crenate or dentate margin. In all species the length of the

petiole is very variable within an individual, but compared to most other Asian genera,

it is generally long in proportion to the length of the leaf blade, and often longer than

the blade.

The inflorescences arise Ifom the axils of the upper leaves, are borne on

long peduncles, and are dichasia with the typical paired-flower arrangement of the

Gesneriaceae. They are few-flowered in most species to many-flowered in Deinostigma

eberhardtii. In most species the inflorescence is fairly lax but is rather congested in the

terminal branches in Deinostigma cycnostyla. There are hooked hairs on the pedicels,

which are not found in Primulina s.s.

The corolla is white, blue or variations on purple (or combinations of these),

infiindibuliform, with the lower lip 3-lobed and the upper lip 2-lobed. From the colour

and morphology bee pollination is likely.

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Gard. Bull. Singapore 68(1) 2016

Microchirita

Codonoboea

Henckelia

Billolivia

Aeschynanthus

Metapetrocosmea peltata

Deinostigma tamiana

Deinostigma poilanei

Deinostigma cyrtocarpa

Deinostigma cicatricosa_KN173

Deinostigma cicatricosaJCWB

Agalmyla

Pseudochirita

Loxostigma

Allostigma guangxiense

Chayamaritia

Petrocosmea

Oreocharis

Conandron ramondioides

Ridleyandra

Hexatheca fuiva

Cyrtandra

Didymostigma

Cathayanthe biflora

Liebigia barbata

Allocheilos guangxiensis

Gyrocheilos

Didymocarpus

Raphiocarpus sinicus

Anna

Briggsiopsis dela vayi

Glabrefla

Lysionotus

Hemiboea

Petrocodon

Primulina gemelia

Primulina dryas

Primulina pinnata

Primulina glandulosa

Primulina Timbrisepala

Primulina giand.var.yangshuo.

Primulina hochiensis

Primulina tabacum

Primulina renifolia

Primulina luochengensis

Primulina ligulifoimls

Primulina lon^angensis

Primulina linearifolia

Primulina minutimaculata

Primulina wentsaii

Primulina ophlopogoides

Fig. 1. Maximum parsimony strict consensus tree based on eombined ITS and trnL-¥ sequenee

data. Clades representing entire genera are collapsed. Numbers along the branches are bootstrap

values. Asterisks denote branehes reeeiving < 50% support.

An expansion oi Deinostigma

151

0.78

Microchirita

Codonoboea

Henckeiia

Didymostigma

Billolivia

Conandron ramondioides

Ridleyandra

Metapetrocosmeapeltata

0.88

Deinostigma tamiana

Deinostigma poilanei

Deinostigma cynocarpa

Deinostigma cicatricosa_KN173

Deinostigma cicatricosa_XWB

Hexatheca fulva

0.95

0.69

Cyrtandra

Oreocharis

Aeschynanthus

Agalmyla

Cathayanthebiflora

Atlocheilos guangxiensis

Gyrocheilos

Liebigia barbata

Didymocarpus

Pseudochirita

Loxostigma

Atlostigma guangxiense

Chayamaritia

Petrocosmea

Briggsiopsis dela vayi

Glabrella

Hemiboea

Anna

Raphiocarpus sinicus

Lysionotus

Petrocodon

Primuiina gemeila

Primulina dryas

Primuiina pinnata

Primulina glandulosa

Primuiina fimbrisepala

Primulina giand.var.yangshuo.

Primulina hochiensis

Primulina renifolia

Primulina luochengensis

Primulina liguliformis

Primulina tabacum

Primulina longgangensis

Primulina linearifolia

Primulina minutimaculata

Primulina wentsaii

Primulina ophiopogoides

Fig. 2. Bayesian inference majority rule consensus tree based on combined ITS and trnL-¥

sequence data. Clades representing entire genera are collapsed. Numbers along the branches

are posterior probability values. Asterisks denote branches with < 0.5 posterior probability.

152

Gard. Bull. Singapore 68(1) 2016

There are two fertile stamens and three staminodes (the central one sometimes

obscure). The filaments are geniculate around the middle and glandular or hairy

distally. The anthers are haiiy or glandular in all species.

The ovary of Deinostigina poilanei is small, c. 5 mm long, at an oblique or

sometimes almost a right angle to the pedicel and slightly curved upwards, similar to

D. cycnostylci (c. 2.4 mm long), D. eherhardtii (c. 4.5 mm long), D. tamiana (c. 5.5

mm long) and D. cyrtocarpa (7-9 mm long). At maturity, the capsules in Deinostigma

poilanei, D. tamiana and D. cyrtocarpa are 1.5-2 cm long and in D. eberhardtii

2. 5-2. 9 cm long. In these species, the capsules are falcate and inserted at an angle to

the pedicel to be earned ± horizontally (not known in Deinostigma cycnostyda).

In Deinostigma cicatricosa and D. minutihamata, the ovary is much larger, c.

12-18 mm long, and only at a slight angle in relation to the pedicel. In Deinostigma

cicatricosa, the mature capsule is 4.5-5 cm long, has a slight curve and is slender, while

in D. minutihamata, the capsule remains straight, 2. 3-3. 5 cm long, and is broader. In

all Deinostigma species where it is known tlie capsule dehiscence is loculicidal.

The internal stmeture of the ovaries could not be studied in all species. In the

species studied the mtemal stmeture varies considerably. In Deinostigma poilanei

it is bilocular for its entire length, with axile recurved bifid placentation (Fig. 3A). In

Deinostigma tamiana and D. cicatricosa only the basal part is bilocular while in the

middle and distal parts the carpels are not fiised and the ovary is unilocular (Fig. 3B,

C). We surveyed other species previously within the circumscription of Chirita, and

now included in other genera (i.e., HenckeUa dielsii (Borza) D.J.Middleton & Mich.

Moller, Microchirita prostrata J.M.Li & Z.Xia, Primulina hochiensis (C.C.Huang &

X. X.Chen) Mich.Moller & A. Weber and Primulina liguliformis (W.T.Wang) Mich.

Moller & A. Weber), and found them to possess a unilocular ovary for their entire length

(Fig. 3E-G). The exception was Primulina dry’as (Dunn) Mich.Moller & A.Weber, the

ovary of which is bilocular throughout but with the abaxial locule being sterile (Fig. 3D).

Chromosome numbers

The chromosome number of Deinostigma poilanei and D. cicatricosa was determined

as 2n = 32, that of D. cyrtocarpa as 2n = c. 30 (Fig. 4). For all species, the cliromosomes

within each complement did not differ significantly in size and were c. 0.8- 1.2 pm in

length. Deinostigma tamiana was counted previously as 2n = 32 (Christie et al., 2012).

The counts for all other Primulina species to date (139 counts for 118 species

and 4 varieties, Moller & Pullan, 2015 onwards) were unifonnly 2n = 36, except for a

tetraploid count of 2n = 72 for one accession of P longgangensis (W.T.Wang) Y.Liu &

Y. Z.Wang (Christie et al., 2012) and one count of 2n = 28 for the same species (Cao et

al., 2003), the latter likely erroneous since two other counts of P. longgangensis showed

2n = 32 (Liu et al., 2012; Kang et al., 2014). Overall, the chromosome size of species

of Primulina s.s. is slightly larger than those with 2n = c. 30 and 32 chromosomes,

being mostly up to 1.6 pm long, with the exception of P. aff balansae (Drake) Mich.

Moller & A.Weber with chromosomes up to 2.0 pm long. Overall, the chromosome

numbers are in full congruence with the phylogeny, separating Primulina s.s. with 2n

= 36 from those with 2n = c. 30 and 2n = 32 chromosomes.

An expansion oi Deinostigma

153

A)

Deinostigma

poilanei

B)

Deinostigma

tamiana

C)

Deinostigma

cicatricosa

D)

Primulina

dryas

E)

Primulina

liguliformis

F)

Henckelia

d/e/s/7

G)

Microchirita

prostrata

near base middle near tip

Fig. 3. Photographic sections taken near the base, the middle and the tip of ovaries of diverse

species belonging to Deinostigma W.T.Wang & Z.Y.Li (A-C), species previously included in

Chirita Bueh.-Ham. (D-F) Microchirita prostrata J.M.Li & Z.Xia (G). (Photos: M. Moller)

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Gard. Bull. Singapore 68(1) 2016

• • ^ •

B

♦

r *\

» ;

■\

Fig. 4. Chromosome spread preparations of three Deinostigma W.T. Wang & Z.Y.Li species. A.

Deinostigma poilanei (Pellegr.) W.T. Wang & Z.Y.Li prometaphase with 2n=32 chromosomes.

B. Deinostigma cicatricosa (W.T. Wang) D.J. Middleton & Mich.Moller metaphase with 2n =

32 chromosomes. C. Deinostigma cyrtocarpa (D.Fang & L.Zeng) Mich.Moller & H.J.Atkins

prometaphase with 2n = c. 30 chromosomes. Scale bar: 10pm. (Photos: A, C: M. Moller; B:

Hui-Min Li)

Geography

The Annamite range of mountains that straddles the border between Vietnam and

Laos, and extends into NE Cambodia and further south in Vietnam to just north of Ho

Chi Minh City, is known to be an area of high biodiversity and home to many endemic

species of plants and animals (Averyanov et al., 2003; WWF, 2015). The Annamites

are particularly interesting for Gesneriaceae as the genus Billolivia DJ.Middleton,

now with seven species (all of which were also described as new), was recently

described from the southern end of this range (Middleton et al., 2014a, 2014b; Vu et

al., 2015), Deinostigma, however, occurs across almost the full length of the mountain

range and into southern China, a distribution pattern not otherwise observed in other

Gesneriaceae genera except for those which are generally much more widespread (e.g.

Aeschynanthus Jack, Didymocarpus Wall., Rhynchotechum Blume).

Conclusion

In summary, phylogeny, cytology and morphology support a separation of Chirita

cicatricosa (previously included in synonymy oi Primulina minutihamata), Primulina

cyrtocarpa and P tamiana from Primulina and their integration into an expanded

genus Deinostigma with a distribution range covering Southern China and Vietnam.

Primulina cycnostyla, P eberhardtii and P minutihamata must also be moved into

Deinostigma based on their close morphological similarities to species which have

been included in the phylogeny. The genus includes species with leaves in alternate

arrangement with distinct nodes (except congested inD. cycnostyla), bilocular ovaries,

at least at the base, and a chromosome number smaller than 2n = 36, these characters

distinguishing Deinostigma from Primulina s.s. Metapetrocosmea, though monotypic

and sister to Deinostigma in the phylogeny, is excluded from Deinostigma due to its

many differences, such as the unilocular ovary, epipetry, free anthers, capitate stigma,

and the capsule globose and straight in relation to the pedicel.

An expansion oi Deinostigma

155

Taxonomy

We do not attempt a revision here of the speeies now included in Deinostigma but

note that a revision is needed. The only major change to the existing taxonomy that we

make is to resurrect Chirita cicatricosa from synonymy of Primulma minutihamata

and make combinations in Deinostigma for both (see below). All type material cited

below has been seen by the authors.

Deinostigma W.T.Wang & Z.Y.Li, Acta Phytotax. Sin. 30(4): 356 (1992). - TYPE:

Deinostigma poilanei (Pellegr.) W.T.Wang & Z.Y.Li

Chirita subsect. Cicatricosae W.T.Wang, Bull. Bot. Res., Harbin 1(4): 69 (1981). -

TYPE: Chirita cicatricosa W.T.Wang

Perennial herbs, caulescent with distinct or short stem. Leaves simple, alternate,

sometimes congested into a rosette, petiolate; blade slightly peltate or not, margin

crenate or dentate, secondary veins pinnate. Inflorescences axillary, cymose; pedicels

with hooked hairs. Flowers with the pedicel inserted at an angle and off-centre on the

receptacle. Calyx lobes divided to base, elliptic, those on ventral side slightly longer

and wider. Corolla white, purple, blue or combinations thereof, infundibulifomi,

lower lip 3-lobed, upper lip 2-lobed, lobe apices rounded. Fertile stamens 2, filaments

slightly curved, anthers adnate face to face, hairy or with glands, staminodes 3 but

with centi'al staminode sometimes obscure. Nectary 5 -crenate or apparently lacking.

Ovary fusiform, bilocular throughout or at least near base, then unilocular from middle

to apex; stigma of only lower lip developing, broad, flat and weakly 2-lobed. Fruit

straight to strongly falcate, oblique in relation to the pedicel, dehiscing loculicidally;

many-seeded, seeds unappendaged.

Distribution. Seven species in southern China and Vietnam. Some of the Vietnamese

material was collected close to the border with Laos and it may also occur in that

country.

Deinostigma cicatricosa (W.T.Wang) D.J.Middleton & Mich.Moller, comb. nov.

- Chirita cicatricosa W.T.Wang, Bull. Bot. Res., Harbin 1(4): 69 (1981). - TYPE:

China, Guangxi, Dongxing, Banba Commune, Renbei, 3 October 1976, Fang, D. etal

1525 (holotype GXMI [GXMI050619]). (Fig. 5A-D)

Distribution. China (Guangxi).

Notes. Wood (1974) included a specimen from Southern Guangxi (although he cited

it as being from ‘Kwangtung’ (Guangdong)) in his new species Chirita minutihamata,

along with material, including the type, from Vietnam. Wang (1981) later described

Chirita cicatricosa from Guangxi without explicitly including the Chinese material

156

Gard. Bull. Singapore 68(1) 2016

of C. minutihamata or referring to that species. Wang (1985), Wang et al. (1990) and

Wang et al. (1998) placed Chirita cicatricosa in synonymy of C. minutihamata and

cited localities corresponding to the type locality of C. cicatricosa and the material

cited by Wood (1974). We acknowledge that the material from China and the material

from Vietnam are indeed very similar but the differences between them, particularly in

the longer, slender, more falcate fruits and the somewhat larger flowers of the Chinese

material, along with the widely disjunct distribution, lead us to suspect that they may

be recognised as distinct species when the group is revised. We therefore provide the

combination in anticipation of this.

Deinostigma cycnostyla (B.L.Burtt) D.J.Middleton & H.J.Atkins, comb. nov.

- Chirita cycnostyla B.L.Burtt, Notes Roy. Bot. Gard. Edinburgh 23: 96 (1960). -

Primulina cycnostyla (B.L.Burtt) Mich.Mdller & A.Weber, Taxon 60: 781 (2011). -

TYPE: Vietnam, Da Nang, BaNa, 1000-1500 m, 27 February 1939, Poilane, E. 29123

(holotypeP[P00602510]).

Distribution. Only known from the Ba Na Hills of central Vietnam.

Notes. This species has not been included in molecular phylogenetic analyses but has

been transferred to Deinostigma based on its morphological similarity to Deinostigma

poilanei.

Deinostigma cyrtocarpa (D.Fang & L.Zeng) Mich.Mdller & H.J.Atkins, comb. nov.

- Chirita cyrtocarpa D.Fang & L.Zeng, Acta Phytotax. Sin. 31(5): 468 (1993). -

Primulina cyrtocarpa (D.Fang & L.Zeng) Mich.Mdller & A.Weber, Taxon 60: 781

(2011). - TYPE: China, Guangxi, Hezhou City, 130-140 m, 16 June 1991, Zhou L.S.

& Zeng L. 1263 (holotype GXMI [GXMI050608]). (Fig. 5E-H)

Distribution. NE Guangxi (Hezhou).

Deinostigma eberhardtii (Pellegr.) D.J.Middleton & H.J.Atkins, comb. nov. - Chirita

eberhardtii Pellegr., Bull. Soc. Bot. France 73: 418 (1926). - Primulina eberhardtii

(Pellegr.) Mich.Mdller & A.Weber, Taxon 60: 782 (2011). - TYPE: Vietnam, Thua

Thien-Hue, Baika, Eberhardt 2466 (lectotype P [P00602512], designated here;

isolectotype VNM).

Distribution. Vietnam (Thua Thien-Hue and Da Nang).

Notes. This species has not been included in molecular phylogenetic analyses but has

been transferred to Deinostigma based on its morphological similarity to Deinostigma

poilanei.

An expansion oi Deinostigma

157

Fig. 5. Deinostigma cicatricosa (W.T.Wang) D.J.Middleton & Mich.Moller (A-D) and D.

cyrtocarpa (D.Fang & L.Zeng) Mich.Moller & H.J.Atkins (E-H). A. Habit and mature fruits.

B. Flower cut open. C. Flower top view. D. Flower front view. E. Habit. F. Mature fruits. G.

Flowering branch. H. Flowers front view. (Photos: A, B, E, F: M. Moller; C, D, G, H: Yi-Gang

Wei)

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Gard. Bull. Singapore 68(1) 2016

Deinostigma minutihamata (D.Wood) DJ.Middleton & HJ. Atkins, comb. nov. -

Chirita minutihamata D.Wood, Notes Roy. Bot. Gard. Edinburgh 31: 370 (1972). -

Primulina minutihamata (D.Wood) Mich.Moller & A.Weber, Taxon 60: 783 (2011).

- TYPE: Vietnam, Kon Turn, Ngok Pa Not, 2300 m, 12 December 1946, Poilane, E.

35803 (holotype P [P00602518]; isotype P [P00602519]).

Distribution. Vietnam (Kon Turn).

Notes. See notes under Deinostigma cicatricosa. This species has not been included in

molecular phylogenetic analyses but has been transferred to Deinostigma based on its

morphological similarity to D. cicatricosa and D. poilanei.

Deinostigma poilanei (Pellegr.) W.T.Wang & Z.Y.Li, Acta Phytotax. Sin. 30(4): 357

(1992). - Hemiboea poilanei Pellegr., Bull. Soc. Bot. France 73: 421 (1926). - TYPE:

Vietnam, Khanh Hoa, Nha Trang, 300 m, 29 May 1922, Poilane, E. 3846 (holotype P

[P00606338]; isotype P [P00634330]). (Fig. 6A-E)

Distribution. Vietnam (Klianh Hoa, Thua Thien-Hue and Da Nang).

Deinostigma tamiana (B.L.Burtt) D.J.Middleton & H.J.Atkins, comb. nov. - Chirita

tamiana B.L.Burtt, Gloxinian 49(4): 20 (1999). - Primulina tamiana (B.L.Burtt)

Mich.Moller & A.Weber, Taxon 60: 785 (2011). - TYPE: Originally collected as

Soviet-Vietnam Expedition 0/114 from Vietnam, Vinh Phuc, Tam Dao National

Park, cultivated in RBGE under accession number 1998 1743* A, vouchered for the

herbarium as CULTE 15738 (holotype E [E00269898]). (Fig. 6F-J)

Distribution. Vietnam (Vinh Phuc, Tam Dao NP).

ACKNOWLEDGEMENTS. We are grateful to R. Rybkova for providing plant material of

Deinostigma poilanei; the horticultural staff of the Royal Botanic Garden Edinburgh (RBGE)

for the cultivation of research material, pailicularly S. Barber and A. Ensoll; S. Barber for

photographs of Deinostigma poilanei; and the Science Technical Services at RBGE for

access to laboratory facilities and their support of the molecular and morphological work. We

acknowledge financial support from the Percy Sladen Memorial Fund (2006) and the RBGE

Foreign Travel fund (2006; 2007), and the Guangxi Natural Science Foundation (2015GXN

SFBB 1 39004/20 15GXNSFBA 139 105) for fieldwork, and the Museum Nationale d’Histoire

Naturelle for funding DJM’s visit. The research of DJM in Singapore Botanic Gardens is

supported by the National Parks Board, Singapore. The Royal Botanic Garden Edinburgh is

supported by the Rural and Environment Science and Analytical Services division (RESAS) in

the Scottish Government.

An expansion oi Deinostigma

159

Fig. 6. Deinostigma poilanei (Pellegr.) W.T.Wang & Z.Y.Li (A-E) andZ). tamiana (B.L.Burtt)

D.J.Middleton & H.J.Atkins (F-J). A. Habit. B. Flower side view. C. Flower front view.

D. Immature fruit. E. Mature fruit. F. Habit. G. Flower side view. H. Flower front view. I.

Immature fruit. J. Mature fruit. (Photos: A, B, C, F-J: M.Moller; D, E: Sadie Barber)

160

Gard. Bull. Singapore 68(1) 2016

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Gardens’ Bulletin Singapore 68(1): 173-174. 2016

doi: 10.3850/S2382581216000120

173

The valid publication of Monolophus

(Zingiberaceae) revisited

J.R Veldkamp

Naturalis Biodiversity Center, RO. Box 9517,

2300 RA Leiden, The Netherlands

jef.veldkamp@naturalis.nl

ABSTRACT. The nomenclatural history of Monolophus Delafosse, Guill. & J.Kuhn is

discussed. The combinations Monolophus picheansoonthonii (Phokham & Prasarn) Veldk. and

Monolophus pubescens (Picheans. & Phokham) Veldk. are made.

Keywords. Kaempferia, new combination, nomenclature, Wallich

Introduction

Mood et al. (2014) argued that Monolophus (Zingiberaceae) was validated by Wallich

(1832), having first been mentioned by Wallich (1830: April) in the second issue of

the Plantae Asiaticae Rariores. However, the name was validated earlier. There is

a review of Wallich’s book by Guillemin (1830: July) in the Bulletin ties Sciences

Naturelles et de Geologie. The French text for Kaempferia elegans Wall, is a more

or less summarised translation of that by Wallich (1830) and thus Monolophus is not

validated in this work: “suffirait peut-etre pour eloinger cette plante du Kaempferia.. .

en former un genre sous le nom de Monolophus'".

On page 3 1 of the Index in the next volume of Bulletin des Sciences Naturelles

et de Geologie, published in October 1830 (fide printers mark on p. [1] of the issue)

Monolophus is mentioned without an author and with the comment “Genre nouveau de

Scitaminees” [New genus of the Scitaminae]. This may be regarded as an acceptance

and validation of the name here. Wallich as the original author was not mentioned.

The journal was edited by [G.] Delafosse [mineralogist], [J.B.A.] Guillemin

[botanist] and [J.] Kuhn [zoologist] and the name is, therefore, to be cited as:

Monolophus Delafosse, Guill. & J.Kuhn, Bull. Sci. Nat. Geol. 23, Index: 31 (October

1830).

Mood et al. (2014) have shown that Kaempferia elegans is not a suitable

candidate for the lectotype of Monolophus. Yet, Picheansoonthon (2016) erroneously

maintained it as such. He also, mistakenly, thought that the new combinations in

Monolophus by Mood et al. are illegitimate, which is against Art. 55.1 of the ICN

(McNeill et al., 2012).

A proposal to conserve Caulokaempferia K.Larsen by Inthararapichai et al.

(2014) has not yet been considered by the appropriate committee. Since the publication

174

Gard. Bull. Singapore 68(1) 2016

of Mood et al. (2014) two new species of Caulokaempferia have been published. These

are here transferred to Monolophus.

Monolophus picheansoonthonii (Phokham & Prasam) Veldk., comb. nov. -

Caulokaempferia picheansoonthonii Phokliam & Prasam in Phokham et ah, J. Jap.

Bot. 90: 155, t. 1,2 (2015). - TYPE: South-western Thailand, Kanchanaburi, Sankhla

Buri Distr., Thung Yai Narasuan Wildlife Sanctuary, Mt Kulu Huamee, near Ro

Kee stream, 15‘’12'97.4'TS[ 98“29'58.2"E. CP [Chayan Picheansoonthon] 160814-1

(holotype BK).

Monolophus pubescens (Picheans. & Phokham) Veldk., comb. nov. - Caulokaempferia

pubescens Picheans. & Phokliam in Phokliam et al., Taiwania 60: 77, t. 1,2. (2015). -

TYPE: Northern Thailand, Mae Hong Son, Mueang Mae Hong Son Distr., CP [Chayan

Picheansoonthon^ 180713-8 (holotype BK).

ACKNOWLEDGEMENTS. Dr K. Gandhi (A), Mr R. Govaerts (K), an anonymous reviewer,

and Dr D.J. Middleton (SING) are much thanked for their critical comments and constructive

help.

References

Delafosse, G., Guillemin, J.B.A. & Kuhn, J. (eds) (1830). Index. Bull. Sci. Nat. Geol. 23: 25, 31.

Guillemin, J.B.A. (1830). # 43. Plantae Asiaticae Rariores. Bull Sci. Nat. Geol. 22: 68-69, 76.

Intharapichai, K., Phokham, B., Wongsuwan, P. & Picheansoonthon, C. (2014). The genus

Caulokaempferia (Zingiberaceae) in Vietnam. J. Jap. Bot. 89: 129-136.

McNeill, J., Bame, F.R., Buck, W.R., Demoulin, V., Greuter, W., Hawksworth, D.L., Herendeen,

P.S., Knapp, S., Marhold, K., Prado, J., Prud’hormne van Reine, W.F., Smith, G.F.,

Wiersema, J.H. & Turland, N.J. (2012) International code of Nomenclature. Regnum

Veg. 154.

Mood, J.D., Veldkamp, J.F., Dey, S. & Prince, L.M. (2014). Nomenclatural changes in

Zingiberaceae: Caulokaempferia is a superfluous name fox Monolophus and Jirawongsea

is reduced to Boesenbeigia. Gard. Bull. Singapore 66(2): 215-231.

Phokham, B., Intharapichai, K., Wongsuwan, P. & Picheansoonthon, C. (201 5). Caulokaempferia

pubescens (Zingiberaceae) - a new species from northern Thailand. Taiwania 60: 77-80.

Phokham, B., Prasam, S., Sangnark, S. & Wongsuwan, P, (2015). A new species of

Caulokaempferia (Zingiberaceae) in western Thailand. J. Jap. Bot. 90: 153-157.

Picheansoonthon, C. (2016). Nomenclatural controversy between Caulokaempferia and

Monolophus (Zingiberaceae). Fol. Malaysiana 17: 67 — 74.

Wallich, N. (1830). Plantae Asiaticae Rariores 1 : 24, t. 27. London: Treuttel & Wiirz.

Wallich, N. (1832). A Numerical List of Dried Specimens of Plants in the East India Company s

Museum, collected under the superintendence of Dr. Wallich, of the Company’s Botanic

Garden at Calcutta (The Wallich Catalogue). # 6591-6593.