THE GARDENS’ BULLETIN, SINGAPORE

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

Dr. K.M. Wong Dr. Jana Leong-Skorni¢ékova

(Editor-in-Chief) (Managing Editor)

SalEee Y.W. Low Christina Soh

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Yale University

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Royal Botanic Gardens, Kew

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National Museum of Natural History

Smithsonian Institution

ESCA

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Royal Botanic Garden Edinburgh

U.K.

Dr. Kiat W. Tan

Gardens By The Bay

National Parks Board, Singapore

Dr. Nigel P. Taylor

Singapore Botanic Gardens

National Parks Board, Singapore

Dr. I.M. Turner

Singapore Botanic Gardens

National Parks Board, Singapore

Dr. J.F. Veldkamp

Nationaal Herbarium Nederland

The Netherlands

Dr. Jun Wen

National Museum of Natural History

Smithsonian Institution

U.S.A.

Professor Nianhe Xia

South China Institute of Botany

PR. China

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(continued on inside back cover)

ARNOLD ARDG Ree FEB 09 oe

Y) LIBRARY

—

The Gardens’ Bulletin

Singapore

VOL. 64(2) 2012 ISSN 0374-7859

Centenary Issue 2012

CONTENTS

Peter C. Boyce, Zulhazman Hamzah and Sofiman Othman

Studies on Monstereae (Araceae) of Peninsular Malaysia IV: The enigmatic

a MAOPHOSG COrnervTetOUNd Alter 75 VAIS ...............0....0000eeseenexseeeeeadeeseseseness

C. Pramod, A.K. Pradeep and J.F. Veldkamp

Coelachne madayensis (Poaceae: Pooideae: Isachneae), a new species from

RE Teed AP Gh 1 cA a FiO ors Ne ek aiden cans susan a wuss cas aa aR Soa ua et bexecTouREcaaeee

M. Rodda, H.D Tran, P.T. Chiew, D. Liew and J. Leong-Skorni¢kova

The rediscovery of Dischidia hirsuta (Apocynaceae, Asclepiadoideae)

oo) LE ERYET OS ac EE ee eee eee

M. Sugumaran and K.M. Wong

Studies in Malesian Gentianaceae I: Fagraea sensu lato—complex genus or

semeraleeneray Amolecular phylogenetic study .........cc1.c..0..cscsessseseerosscssesseeoes

Pramote Triboun and David J. Middleton

Twenty new species of Paraboea (Gesneriaceae) from Thailand ................00.

I.M. Turner

Annonaceae of Borneo: a review of the climbing species ...............::cccccesseeeeeeeees

K.M. Wong and M. Sugumaran

Studies in Malesian Gentianaceae II: A taxonomic framework

for the Fagraea complex, including the new genus Limahlania ...........c:cccee

K.M. Wong and M. Sugumaran

Studies in Malesian Gentianaceae III: Cvrtophyllum reapplied

MOUSE aS HACA [FAST OMS AUTANC Sy a Fe crake angie ee see kc cn eee eacaestadoosesacdssonenesstaneessee

wel

eae ey)

293

eo Oil

333

sero

K.M. Wong

Studies in Malesian Gentianaceae IV: A revision of Picrophloeus .......ccccccccccceeeees Sl

Zulhazman Hamzah, Peter C. Boyce and Mashhor Mansor

Studies on Homalomeneae (Araceae) of Peninsular Malaysia IV:

Homalomena stongensis, a remarkable new species endemic

to Gunung Stong; Kelamtam’ ...2.....5.5.cee eset ee eee eee ee 523

BOOK REVIEWS

Etlingera of Sulawesi. A.D. Poulsen. 2012 (J. Leong-Skorniékova) ..........c:ce0eeeee 529

Flora of Hong Kong. Volume 4. N.H. Xia, K.W. Cheung, K.S. Pang

& KL. Yip(eds)- 2011 (Ro IMs Woe) eee eae ee eee cee ce eee eee 533

Private Lives. L.K. Wang, D.C.J. Yeo, K.K.P. Lim & S.K.Y. Lum (eds).

2012 (K.M. Wong), «:.:.cccesseesch ed SeeGeneee ties eee eee ee ee 1S ))

Date of publication: 12 December 2012

National Parks Board

Singapore Botanic Gardens

1 Cluny Road

Singapore 259569

Printed by Oxford Graphic Printers Pte Ltd

Gardens’ Bulletin Singapore 64(2): 281-288. 2012 281

Studies on Monstereae (Araceae)

of Peninsular Malaysia IV:

The enigmatic Rhaphidophora corneri refound

after 75 years

Peter C. Boyce'’, Zulhazman Hamzah? and Sofiman Othman!

"School of Biological Sciences, Universiti Sains Malaysia,

11800 USM, Pulau Pinang, Malaysia

*phymatarum@googlemail.com

*Faculty of Earth Science, Universiti Malaysia Kelantan,

Jeli Campus, Locked Bag 100,

17600 Jeli, Kelantan, Malaysia

ABSTRACT. Rhaphidophora corneri P.C.Boyce, a highly distinctive but hitherto poorly

known species described from fragmentary material collected by E.J.H. Corner late in 1935

from Kemaman (Terengganu state in Peninsular Malaysia) has recently been refound in

neighbouring Kelantan. A much-expanded species description is provided, along with new

information pertaining to its ecology. A reinterpretation of possible relationships with other

Rhaphidophora species is offered in light of these novel data. Photographs depicting newly

observed vegetative morphology are provided.

Keywords. Monstereae, Peninsular Malaysia, Raphidophora

Introduction

The taxonomic revision of Rhaphidophora for Peninsular Malaysia and Singapore

by Boyce (1999) included the description of a highly distinctive new species from

Kemaman (Terengganu state) based on somewhat fragmentary material collected

by E.J.H. Corner in November 1935. Despite the paucity of material (Fig. 1) Boyce

noted at the time that “Rhaphidophora corneri is a remarkable species, unique in the

genus by the manner of inflorescence production, small leaves with dense reticulate

veins, and large perforations...”, although the nature of the single specimen left many

questions open.

During a preliminary survey of the aroids of Taman Negara Kuala Koh,

Kelantan, in June 2010, a small population of R. corneri was located. A second more

detailed survey by the second author in the following July revealed the population

to be larger. These plants represent only the second ever collection, and enable

expansion of the species description presented by Boyce, resolving several aspects of

the plant’s morphology. Furthermore, these additional data permit insight into possible

relationships of R. corneri with other Rhaphidophora species.

282 Gard. Bull. Singapore 64(2) 2012

Rhaphidophora corneri P.C.Boyce, Gard. Bull. Singapore 51 (1999) 183-256. TYPE:

Malaysia, Kemaman, Terengganu, Ulu Kajang, 13 Nov 1935, E.J.H. Corner SFN

30441 (holo SING!). (Fig. 1-3)

Small, rather slender, homeophyllous, leptocaul, liane up to 2.7 m height, with shoots

extending along the ground for up to 2.4 m; seedling stage and pre-adult plants

unknown; adult shoot architecture comprised of a much elongated, free or weakly

clinging, physiognomically monopodial, leafy stem; stems smooth, without prophyll,

cataphyll and petiolar sheath fibre, the internodes 1—14 x 0.5—0.7 cm long, with up to

31 internodes per module; branches arising almost perpendicular to main axis, up to

44 cm long, junction with main axis with a prominent corky leaf scar; roots one per

node, adherent, stout, corky; /eaves glossy medium green, erect or spreading on adult

shoots, cataphylls and prophylls membranaceous, soon drying and falling; petiole

narrowly canaliculate, pulvinate, 5—14 « 0.5—1.6 cm, smooth, apical and basal pulvini

prominent; petiolar sheath obscure except for basal-most portion, extending to the

apical pulvinus, sheath of newest leaf degrading into a very few feeble fibres, sheath

soon falling to leave a proportionally wide, corky scar basally on the petiole; blade

oblong-lanceolate, slightly oblique, 15-27 x 8—14 cm, sub-coriaceous, entire or with

1-4 perforations on the wider side, these large, ovate to rhomboid or trapezoid, often

extending almost to the margin, base obtuse to weakly cordate, apex acute to acuminate

with a slightly prominent apiculate tubule; mid-rib prominently raised abaxially,

slightly raised adaxially; primary venation pinnate but distal-most veins becoming

weakly reticulate and not reaching leaf margin, raised abaxially, raised (but weakly so)

adaxially, 34 per side, alternate; interprimaries weakly reticulate to sometimes sub-

parallel to primaries, slightly raised abaxially, weakly raised adaxially, often forming a

weak reticulum; secondary venation reticulate, slightly raised abaxially, weakly raised

adaxially; tertiary venation prominently reticulate. Inflorescence solitary, terminating

a much-abbreviated specialised axis arising from the axil of a foliage leaf, with

several inflorescences arising from sequential or semi-sequential leaf axils; peduncle

terete, 1—1.3 x 0.2—0.25 cm; spathe canoe-shaped, thick, stout-beaked, c. 2.7—2.9 x

0.7—0.8 cm, soon caducous; spadix cylindrical, sessile, inserted slightly obliquely on

peduncle, c. 2 x 0.4 cm; stvlar region well developed, rounded-rhombohexagonal, c. 3

< 2 mm, convex to truncate, smoothly rounded; stigma impressed, irregularly elliptic,

longitudinally oriented, c. | x 0.5 mm; stamens elongating at anthesis; fi/aments lorate,

c. 2.5 x 1 mm; anthers ellipsoid, bluntly rostrate terminally, c. 1.5 x 0.6 mm; mature

infructescence not observed.

Distribution. Peninsular Malaysia: Terengganu (Kemaman) and Kelantan (Kuala

Koh).

Habitat. Perhumid lowland mixed dipterocarp forest on ridgetops and flat open areas,

80—100 m asl.

= 70>

Rhaphidophora corneri rediscovered 283

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Fig. 1. Rhaphidophora corneri P.C.Boyce. Holotype: E.J.H.Corner SFN 30441 (SING).

Gard. Bull. Singapore 64(2) 2012

Fig. 2. Rhaphidophora corneri P.C.Boyce. A. Terrestrial population. B. Climbing axis. C.

Adult leaf blade, adaxial view. (Photos: Zulhazman H.)

; We

Rhaphidophora corneri rediscovered 285

Fig. 3. Rhaphidophora corneri P.C.Boyce. A. Adult shoot, leaves in adaxial view; note the single

root arising from each node. B. Adult shoot, abaxial view. The lateral branches arise almost

perpendicularly to the main axis. C. Adult leaf blade, abaxial view. (Photos: Zulhazman H.)

286 Gard. Bull. Singapore 64(2) 2012

Other specimen examined: Malaysia, Kelantan, Kuala Koh National Park, 4°52’5.37”N

102°26°31.2"E, 1 June 2010, Zulhazman H.UMK00031 (Herbarium Universiti

Malaysia Kelantan, Faculty of Earth Science, UMK).

Notes. Boyce (1999) speculated that the manner of inflorescence production in R.

corneri, as then understood—several inflorescences together, each subtended by a

cataphyll—indicated a close relationship with the New Guinean R. /edermannii Engl.

& K.Krause and R. versteegii Engl. & K.Krause (Spathacea Group: see Boyce 2000,

2001a, b). However, re-examination of the single flowering shoot on the holotype

of R. corneri (SING), coupled with insights provided by the hitherto unknown

vegetative morphologies presented here, suggest that R. corneri is markedly more

similar to Peninsular Malaysian R. tetrasperma Hook.f., and through this probably

related to morphologically congruent taxa such as R. pertusa (Ridl.) Schott and R.

nicolsonii P.C.Boyce. The taxa together form a distinctive and hitherto unrecognised

morphotaxon—the Pertusa Group.

The holotype collection of R. corneri is, as remarked, somewhat fragmentary.

The single flowering shoot bears four inflorescences; three post-anthesis and one

in bud, and the entire inflorescence-bearing portion is leafless, with the last foliage

leaf on the shoot situated some distance behind the first inflorescence, and that

with a somewhat reduced blade. Along the inflorescence-bearing portion there are

numerous conspicuous scars, with one scar associated with each of the post-anthesis

inflorescences. Initial interpretation of the inflorescence-bearing shoot was that each

inflorescence had been subtended by a prophyll/cataphyll, the scars being the result

of this being shed, and that the inflorescence in bud marked the tip of a still-extending

flowering axis. Such shoot morphology, albeit with the axis between each inflorescence

considerably reduced, is found in the Rhaphidophora Spathacea Group (Boyce 2001b),

while remarkably similar inflorescence architecture (but there certainly independently

derived) occurs in Pothos L. (Araceae: Potheae), notably in P. insignis Engl. (Boyce &

Poulsen 1994; Boyce & Hay 1997, 2001).

At the time of the first examination (late 1997), the age and fragility of the

holotype material, its unique nature, and nomenclatural importance, precluded further

(destructive) examination. More detailed examination has now been enabled, aided

by digital enhancement, and we are now of the belief that the probable origin of the

scars subtending each of the inflorescences is result of the loss of foliage leaves rather

than prophylls or cataphylls and that the flowering shoot is better interpreted as having

originally comprised solitary inflorescences, each terminating a much-abbreviated

specialised axis arising from the axil of a foliage leaf. This is exactly the manner in

which R. tetrasperma produces inflorescences (see Fig. 4).

Vegetative characteristics revealed by this rediscovery support the new

interpretation presented here. Rhaphidophora corneri, together with all species in

the Pertusa Group, has green, smooth stems, and bears foliage leaves at all except

the first few congested nodes of a new module. Virtually every node produces one

or occasionally two roots that may either remain short and function as climbing

287

Rhaphidophora corneri rediscovered

ROYAL BOTANIC GARDENS KEW

MIDEA ANAM

K000291788

SOY |OMUOD 10JOD HYCO

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ys¢ HERBARIUM MUS; PERSE |

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; Species £ ‘ se

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‘

| Taocality Type /

Collected by “Rovd. Father Soortechini ;

} { é a Vee ha

FLORA GF BEITIKH INDIA. Ya & psy)

se ey Samed og MTD HOOKER ,

Fig. 4. Rhaphidophora tetrasperma Hook.f. Holotype: Fr B. Scortechini 169b (K).

288 Gard. Bull. Singapore 64(2) 2012

roots, or extend to the ground as feeder roots. Leaf blades in the group are always

irregularly perforated and/or irregularly pinnate. All species have one or more primary

axes which remain sterile and produce lateral shoots, along which are fertile nodes

(Stone’s Model—see Hallé & Oldeman 1970; Hallé, Oldeman & Tomlinson 1978).

Inflorescences are born on very short lateral shoots arising from the axils of foliage

leaves towards the ends of moderately elongated, lateral shoots.

ACKNOWLEDGEMENTS. The authors would like to acknowledge the Department of Wildlife

and National Parks, Peninsular Malaysia (PERHILITAN) for allowing them to conduct the

study in the Kuala Koh National Park. Special thanks to Mr. Nik Yuszrin Yusof, Ms. Naziah

Zaid and Ms. Norzielawati Salleh for their kind assistance in our work. The first author’s

fieldwork was supported by USM Grant No: 1001/ JNC/ AUPRMO0O1. The second author’s

project is funded by Universiti Malaysia Kelantan through short-term research grant R/SGJP/

A03.00/00279A/001/2009/000021 via the Faculty of Agro Industry & Natural Resources.

References

Boyce, P.C. (1999) The genus Rhaphidophora Hassk. (Araceae-Monsteroideae-Monstereae) in

Peninsular Malaysia and Singapore. Gard. Bull. Singapore 51: 183-256.

Boyce, P.C. (2000) The genus Rhaphidophora Hassk. (Araceae-Monsteroideae-Monstereae)

in the southern and western Indonesian archipelago. Gard. Bull. Singapore 52: 101-183.

Boyce, P.C. (2001a) The genus Rhaphidophora Hassk. (Araceae-Monsteroideae-Monstereae)

in Borneo. Gard. Bull. Singapore 53: 19-74.

Boyce, P.C. (2001b) The genus Rhaphidophora Hassk. (Araceae-Monsteroideae-Monstereae)

in New Guinea, Australia and the tropical western Pacific. Gard. Bull. Singapore 53:

75-183.

Boyce, P.C. & Hay, A. (1997) Diversity in shoot architecture in Pothos (Araceae: Pothoideae):

Observations towards a new infrageneric classification. In: J. Dransfield et al. (eds) Plant

Diversity in Malesia III, pp. 51-58.

Boyce, P.C. & Hay, A. (2001) A taxonomic revision of Araceae tribe Potheae (Pothos,

Pothoidium and Pedicellarum) for Malesia, Australia and the tropical Western Pacific.

Telopea 9(3): 449-571.

Boyce, P.C & Poulsen, A.D. (1994) Notes on Pothos insignis (Araceae: Pothoideae). Kew Bull.

49(3): 523-538.

Hallé, F. & Oldeman, R.A.A. (1970) Essai sur l’Architecture et la Croissance des Arbres

Tropicaux. 178 pp. Paris: Masson et Cie.

Hallé, F., Oldeman, R.A.A. & Tomlinson, P.B. (1978) Tropical Trees and Forests; An

Architectural Analysis. 441 pp. Heidelberg: Springer-Verlag.

Gardens’ Bulletin Singapore 64(2): 289-292. 2012 289

Coelachne madayensis (Poaceae: Pooideae: Isachneae),

a new species from Kerala, India

C. Pramod’, A.K. Pradeep’ and J.F. Veldkamp*

‘Department of Botany, Govt. Brennen College,

Darmadam, Thalassery, Kerala, India 670 106

cpramod4@gmail.com (corresponding author)

> Department of Botany, University of Calicut,

Kerala, India 673 635

akpradeep 1 (@rediffmail.com

> Naturalis Biodiversity Center,

National Herbarium of The Netherlands, Leiden University,

P.O. Box 9514, 2300 RA Leiden, The Netherlands

jan-frits.veldkamp(@naturalis.nl

ABSTRACT. A new species of Coelachne (Poaceae: Pooideae: Isachneae) from Madayippara

in the Kannur District of Northern Kerala, India is described and illustrated. A key for the

identification of the Indian congeners is provided.

Keywords. Coelachne, India, Kerala, Poaceae

Introduction

The genus Coelachne R.Br. (Poaceae: Pooideae: Isachneae) is represented by 10

species distributed in the tropics and subtropics of the Old World (Mabberley, 2008;

Clayton et al. 2006+). In India the genus is represented by 3 species, of which one

species and one variety are endemic (Prakash & Jain 1984). In Kerala there are 2

species, viz. Coelachne infirma Buse [syn. C. simpliciuscula (Wight & Arn. ex Steud.)

Munro ex Benth.] and C. perpusilla (Arn. ex Steud.) Thwaites, which usually occur in

wet places (Sreekumar & Nair 1991).

During a floristic survey in July 2011 of the lateritic plateau habitats of

Northern Kerala, India, the first two authors noticed a population of a very small species

of Coelachne in the shallow seasonal pools and nearby wet areas. The specimens

turned out not to belong to any described species of Coe/achne, though they showed

some similarity with the Indian species C. minuta Bor in their general habit, size,

and organization of the spikelets. However, they differed in their non-tufted habit,

absence of a ligule, ovate leafblades, flagleaves without a blade, racemes with up to

15 spikelets, lower florets with 0-nerved glumes and paleas, and 3 stamens. Therefore

they are here described as a new species.

290 Gard. Bull. Singapore 64(2) 2012

Coelachne madayensis Pramod & Pradeep, sp. nov.

Coelachnae minutae similis in habitu generali magnitudineque spicularum structura,

sed habitu non caespitoso, foliorum laminis ovatis sine ligula, ultimis lamina carentibus,

racemis paucispiculatis, glumis paleisque avenis, staminibus 3 differt. TYPE: India,

South India, Kannur District, Madayippara, alt. 36 m, 12°1.792’N 75°15.246’E, 25

July 2011, Pramod & Pradeep CU126681 (holo K; iso BRIT, CALI, SING). (Fig. 1)

Erect aquatic herb, 3-8 cm high, rooting at the lower nodes. Culm canaliculate,

internodes 3—5 mm long, glabrous, nodes glabrous. Sheaths up to 6 mm long, as long

as or shorter than the internodes, laxly clasping the culm, distinctly striate, sparsely

hispid (especially on the midrib); Jigule absent; blades ovate, 0.8—1.2 x 0.5—0.6

mm, 7—15-nerved, glabrous above, puberulous below, base cordate, margin at base

ciliate, apex acute; flagleaf sheaths 1-3 cm long, glabrous, margins hyaline, apex

pointed, blade absent. Panicles 3—6 cm long; branches up to 8 mm long, with gland-

like swellings at base. Spikelets up to 15, 1—3 per branch, elliptic, 1-1.5 x 0.5—0.7

mm, purplish. Glumes ovate, purplish, margins hyaline; lower glume c. 0.75 x 0.5

mm, purplish; upper glume, 0.9 <x 0.5 mm. Lower lemma lanceolate, 1.5 x 0.5 mm,

0-nerved, purplish, margins hyaline, margins and upper surface pilose. Palea boat-

shaped, c. 1.25 x 0.8 mm * c. 0.5 mm, deep purplish, margins hyaline, 0-nerved,

upper surface pilose. Stamens 3; anthers 1.25 x 0.5 mm, greenish white. Style forked,

hyaline, stigma plumose. Upper floret tufted cottony hairy at base, c. 1.5 x 0.5 mm,

purplish. Lemma boat-shaped, c. 1.2 x 0.5 mm, 0-nerved, pilose, purplish, margins

hyaline. Palea slightly shorter than lemma, boat-shaped, c. 1 < 0.5 mm, purplish,

margins hyaline. Caryopses ellipsoid, c. 1 x 0.5 mm, deep purplish, glabrous, shiny.

Ecology. The species is found growing in shallow seasonal pools and nearby wet

areas along with, e.g., Eriocaulon spp., Fimbristylis sp., Geissaspis cristata Wight &

Arn. var. tenella (Benth.) M.R.Almeida, /sachne veldkampii K.G.Bhatt & Nagendran,

Rhamphicarpa longiflora Benth., Rotala malabarica Pradeep, K.T.Joseph & Sivar., R.

malampuzhensis R.V.Nair ex C.D.K.Cook, and Utricularia spp. Usually the panicle

alone is seen emerging from the water.

Etymology. The new species is named after its type locality, Madayippara in the Kannur

District of Kerala, India.

Key to the Indian taxa of Coelachne

(Prakash & Jain, 1984, modified)

la: Culmsverecty Spikelets: 11/5) tin Vomiey2 2 essere fester ee ee ee D

b. Culms-‘ascending. Spikelets‘over 125 mong yes eee 5

Anew Coelachne from India

291

er tit

MHI eee es

CARED ne brs oor

Fig. 1. Coelachne madayensis Pramod & Pradeep. A. Habit. B. Spikelet. C. Lower glume. D.

Upper glume. E. Lower lemma. F. Lower palea. G. Upper lemma. H. Upper palea. I. Pistil. J.

Caryopsis. Scale bars: A= 1 cm; B= 1 mm; C-J = 0.5 mm. (Drawn by C. Pramod)

Gard. Bull. Singapore 64(2) 2012

2a. Culms not tufted. Sheaths laxly clasping the culm. Ligule absent. Blades ovate.

Flagleaf bladeless. Panicle usually with less than 15 spikelets. Lower glume c. 0.75

mm long; upper glume c. 0.9 mm long, 0-nerved. Palea 0-nerved. Stamens 3 ........

asiasesa.aeaavaeevGadvan Davee tetebed acd dud olgdudanalted sate had eae eee eae ene C. madayensis

b. Culms tufted. Sheaths firmly clasping the culm. Ligule present, silky hairy. Blades

lanceolate. Flagleaf with a blade. Panicle with more than 15 spikelets. Lower

glume c. 0.25 mm long; upper glume c. 0.5 mm long, 2-nerved. Palea 2-nerved.

StaMMGMS)2> 25... ssuckucscselsiceastecacesesciiavester ten teconells tote ee tenet eeeee cee eRe ee eR Renee ree C. minuta

3a. Panicles few-flowered, very lax. Spikelets 2.5—4 mm long. Lemmas densely

bearded at bases. ssi. iwendinn aes eta oe ee eae oe se eee eee ~

b. Panicles many-flowered, mostly contracted and spiciform. Spikelets 1.5—2.5 mm

long: Lemmas glabrous at base 24.2 ce-s ee ee ee ee C. infirma

4a. Culms slender. Blades flat, linear-lanceolate. Spikelets 3.54.0 mm long. Lower

glume 1.7—2.0 mm long, 1—3-nerved; upper glume 2.3—2.5 mm long. Upper lemma

and palea not keeled, 0- (or 1-)nerved, glabrous or sparsely hairy in the lower half.

Anthers15=1.7 mm longa ee ee C. perpusilla var. nilagirica

b. Culms capillary. Blades involute, filiform. Spikelets 2.5-3.0 mm long. Lower

glume 1—1.5 mm long, 3—7-nerved; upper glume 1.5—2.0 mm long. Upper lemma

and palea 2-keeled, 2-nerved, hairy along the keels and all over the dorsal surface.

Anthers’0:8=l(E 125) mma\lon geese eee eee C. perpusilla var. perpusilla

ACKNOWLEDGEMENTS. The authors wish to thank Prof. M. Sabu, University of Calicut,

Prof. S.R. Yadav, Kolhapur, Mr. Vivek C. Paulose, Botanical Survey of India, Coimbatore

and Dr. Muhamed Jafer Palot, Zoological Survey of India, Calicut for various assistance and

valuable suggestions. The first author thanks the University Grants Commission, Govt. of

India for the financial assistance in the form of a minor research project (MRP(S)-592/09-10/

KLKAOI1/UGC-SWRO).

References

Clayton, W.D., Vorontsova, M.S., Harman, K.T. & Williamson, H. (2006+) GrassBase - The

online world grass flora. http://www.kew.org/data/grasses-db.html accessed 8 March

2012:

Mabberley, D.J. (2008) The Plant-Book: A Portable Dictionary of Vascular Plants. Cambridge:

Cambridge University Press.

Prakash, V. & Jain, S.K. (1984) Poaceae: tribe Isachneae. Fascicles of Flora of India 14: 3—7,

t. 1-13. Howrah: Botanical Survey of India.

Sreekumar P.V. & Nair, V.J. (1991) Flora of Kerala—Grasses. Pp. 414-416. Calcutta: Botanical

Survey of India.

Gardens’ Bulletin Singapore 64(2): 293-299. 2012 293

The rediscovery of Dischidia hirsuta

(Apocynaceae, Asclepiadoideae) in Singapore

M. Rodda'-?, H.D Tran’, P.T. Chiew’,

D. Liew? and J. Leong-Skorniékova'

‘Singapore Botanic Gardens,1 Cluny Road, 259569 Singapore

*Conservation Division, Central Nature Reserve,

National Parks Board, | Cluny Road, 259569 Singapore

‘rodda.michele@gmail.com

ABSTRACT. This paper seeks to document the rediscovery of Dischidia hirsuta (Bl.) Decne.

in Singapore. The taxon was last collected from Singapore in 1903 and later considered

nationally extinct. Botanical investigation of Nee Soon Swamp Forest in May 2011 resulted

in the rediscovery of a single specimen of D. hirsuta, now considered critically endangered.

Keywords. Dischidia, UCN Red Lists, Nee Soon Swamp Forest, Singapore

Introduction

The genus Dischidia R.Br. occurs from India, across SE Asia, to the east Pacific region

and Australia. The genus has not been recently revised but it has been the subject of

modern floristic treatments (Rintz 1980; Li et al. 1995; Forster et al. 1996; Jagtap &

Singh 1999; Livshultz et al. 2005). Dischidia species are epiphytic and lithophytic

climbers with long and slender stems rooting at the nodes and along internodes. The

leaves are usually opposite, entire, ovate to elliptic but can alternatively develop into

three distinct types: shell-like leaves, usually tightly appressed to the host plant bark,

pitchers, and terete leaves (observed in Dischidia bengalensis Colebr. only). Shell-like

leaves and pitcher leaves accommodate ants that establish a symbiotic interaction with

the plant (Janzen 1974).

Inflorescences are spirally-elongating racemes producing flowers more-or-

less regularly over a variable amount of time, sometimes up to years. The flowers

are pentamerous, with a small calyx and a typically urceolate corolla more-or-less

pubescent internally. All species have a staminal corona composed of five hyaline

anchor-shaped appendages alternating with the stamens. In some species, a corolline

corona also occurs together with the staminal corona and is formed by five lobes

attached to the inner surface of the corolla. The pollen is carried in pollinia lacking

sterile edges, attached to a central retinaculum by two often broad and cristate

caudicles. Fruit follicles are variable, from terete to reniform or triangular in cross

section (Rintz 1980). The phylogenetic position of Dischidia with respect to Hoya is

unresolved (Wanntorp et al. 2006a, b; Wanntorp et al. 2011) but they are considered to

be separate on a morphological basis (Wanntorp & Kunze 2009).

294 Gard. Bull. Singapore 64(2) 2012

In Singapore, Ridley (1900) recognised seven species at first, but later eight

(Ridley 1923). Rintz (1980) made a critical revision of Dischidia for the Malay Peninsula

(now including Peninsular Malaysia and Singapore) and recognised seven species

occurring in Singapore. Currently nine species are listed as native in the Checklist of

Singapore Flora (Chong et al. 2009) but five of these are considered nationally extinct.

Among these, D. singaporensis Ridl., is considered a globally extinct species.

Distribution and variability of D. hirsuta

Dischidia hirsuta derived its specific epithet from the characteristic pubescence of the

young leaves and stems. It was first described as Leptostemma hirsuta Bl. (1826) and

transferred to the genus Dischidia by Decaisne (1844). It is a rather common species

occurring continuously from Burma across to New Guinea (Rintz 1980). It occupies a

narrow altitudinal range and can be found growing from lowland to hill forests below

100 m asl. (Rintz 1980). It is usually found on small branches high in the tree canopy

where it is exposed to nearly full sun. For this reason, despite its showy flowers, it

is difficult to notice unless it is found on recently fallen branches, for example, after

storms.

The species 1s very variable in vegetative and flower morphology depending

on the provenance and on the growing condition. Flower colour can range from pale

yellow with white corolla lobes to burgundy red, and corolla shape can be from broadly

urceolate with a nearly flat base to narrowly ovoid (Fig. 1). Leaf shape can vary from

lanceolate to orbicular, flat with visible veins or thick, succulent, with an obscure

venation, and leaf size ranges from 0.5 x 0.5 cm to 2.5 x 1.5 cm. Leaf colour varies from

dark green to light green-yellow with red or maroon spots (Fig. 1). Lighter-coloured,

thicker leaves are typical of specimens growing in direct sunlight but even this may

vary from one locality to another (Rodda, pers. obs.). In mature parts of the plant the

pubescence typical of juvenile leaves is less conspicuous and often completely absent.

While different collections have been described under different names over the years,

D. hirsuta is currently considered a single species with a large intraspecific variation.

Such a high morphological variation has not yet been studied in detail and future

studies could lead to resurrection of some of the six species names that are currently

considered to be synonymous with D. hirsuta (Rintz 1980).

Dischidia hirsuta in Singapore

From herbarium records it appears that D. hirsuta was rather widespread in Singapore

at the end of the 19th century. It was recorded from natural areas such as Kranji

and Seletar, but also along the more populated Nassim Road. The last collection

(in Woodlands) dates back to 1903 and therefore the species had been considered

presumably extinct (Tan et al. 2008; Davison 2008) until a recent botanical survey

of Nee Soon Swamp Forest we conducted in May 2011. It was found growing on

905

Fig. 1. Morphological variation of Dischidia hirsuta across its distribution area. For each accession

a leaf (adaxial surface) and a flower (side view) is presented. A. Borneo, Sabah, 1989, Ridsdale

s.n. (Hortus Botanicus Leiden number 910235). B. Thailand, 24 Aug 2000, de Wilde 143909

(20031259). C. Indonesia, Irian Jaya, Feb 1998, Yumte s.n. (981096). D. s. loc, s. coll., (970839).

E. Singapore, Upper Seletar track, 3 May 2011, Rodda, Leong-Skornickova, Tran, Liew & Chew

MR11-008 (SING). F. Philippines, 21 Jan 1993, Ridsdale & Pancho s.n. (930049). G. Borneo,

Sarawak, 1998, Vogel, Schuiteman & Roelfsema s.n. (980056). H. s. loc. 21 Jan 1993, Ridsdale

s.n. (930045). I. Indonesia, Sulawesi, Tauecchio ETS94031 (20000247). All pictures from living

specimens cultivated in the Hortus Botanicus Leiden unless otherwise stated. (Photos: M. Rodda)

296 Gard. Bull. Singapore 64(2) 2012

fallen branches on the forest floor in deep shade in secondary forest, but the branches

had grown in the canopy where they were exposed to nearly full sun. We could not

locate any additional specimen on neighbouring trees. The species can be observed

in similar habitats in Johore, Malaysia, where D. hirsuta is a common epiphyte in

lowland freshwater swamps (Rodda, pers. obs.).

Conservation considerations

Since the discovery of a single specimen of D. hirsuta and its precarious state, the

species is more suitably considered Critically Endangered in Singapore, according

to guidelines given in The Singapore Red Data Book (Davison 2008). It is plausible

that further specimens of D. hirsuta and other nationally extinct epiphytic species

could still persist in the canopy of suitable trees in the relatively under-collected Nee

Soon Swamp Forest. This habitat should therefore be further explored and preserved.

Cuttings for propagation were taken from the specimen of D. hirsuta in Nee Soon

Swamp Forest. Currently about 50 plants have been raised at the Singapore Botanic

Gardens (SBG Acc. No. 20111110) to create enough stock for conservation work,

including possible re-introduction trial planting.

Further collections and detailed studies both at the population level and across

the distribution range are needed to verify the extent of morphological as well as

genetic variation in order to understand whether D. hirsuta is a species complex, which

may then require to be recognised as different taxonomic units. From a conservation

perspective, the geographic and genetic origin of propagation materials to be potentially

used for reintroduction is therefore of essential importance to understand together with

its taxonomy. This then helps to avoid genetic contamination of the natural populations

of the taxon, in particular in Singapore, where natural populations could be highly

restricted and ecological repercussions potentially impactful.

Description of the Singapore taxon

The following morphological description is based on specimens collected in Singapore

(Fig: 2):

Plants succulent, herbaceous; epiphytic vines growing tightly appressed to the

host tree stems, clinging by adventitious roots, rarely pendulous, usually preferring to

colonise small canopy branches up to 5 cm diameter. Latex white. Roots on mature

plants adventitious, produced just below each node and along the internodes. Leafy

stems terete, up to 2.5 mm diameter, dark green, hirsute when young; internodes 1—2

cm long. Leaves opposite, petiolate; petiole cylindrical, 1.5—2.5 mm long, c. 1 mm in

diameter, sparsely pubescent; lamina dark green, underneath lighter green with darker

spots, coriaceous, fleshy, (orbicular) broadly ovate (lanceolate), 1-2.5 =< 0.6—1.5

cm, hirsute on both surfaces when young; apex acute-apiculate, base round-truncate

(shortly attenuate), margin entire, main vein evident on adaxial surface, marked by a

dark green line on abaxial surface, secondary veins 3-4 pairs, branching at 60—80(—

90)° from main vein. Inflorescences usually bearing a single open flower at a time;

peduncle extra-axillary, perennial, bearing scars of previous flowerings 2—5 mm long,

c. | mm in diameter, pubescent; pedicels 2-3 mm long, c. 0.7 mm in diameter, glabrous

= 297

Fig. 2. Dischidia hirsuta in Singapore. A. Flowering branches bearing single-flowered

inflorescences. B. Single-flowered inflorescence. C. Dissected flower. The double ring of hairs

lining the corolla tube, the hyaline appendages sorrounding the staminal corona and the central

anther appendages can be observed. Based on Rodda, Leong-Skornickova, Tran, Liew & Chew

MR11-008 (SING). (Photos: M. Rodda (A, C) and J. Leong-Skorniékova (B)).

(sparsely hirsute). Calyx c. 3 mm in diameter, red-purple, lobes ovate 1—1.5 < 0.6—1

mm, apex acute to round, glabrous. Corolla succulent, urceolate, c. 7 x 4 mm, basally

dark red, progressively fading into light pink at the lobe apices, externally glabrous,

298 Gard. Bull. Singapore 64(2) 2012

corolla throat with two rings of hairs, the first just below the corolla lobe sinus, the

second at the same high as the anther appendages; lobes nearly entirely fused, glabrous,

with free ovate tips up to 2 mm long. Corolline corona absent. Staminal corona 3 mm

high, 3 mm in diameter, basally pubescent, composed of five hyaline appendages with

a truncate apex and two downcurved lobes. Stigma head prominent, convex, hidden

by the anther appendages. Pollinarium erect (measuring c.) 1000 « 600 tum with two

oblong pollinia, 600 x 200 um, apex and base rounded to obtuse, retinaculum 420 x

120 um, caudicles broad, laterally crested, 400 um long. Ovary bi-carpellate, bottle

shaped, 1.5 mm long, each carpel c. 0.7 mm in basal diameter. Fruits and seeds not

seen.

Singapore specimens examined. Tanglin, Nassim Road, 1901, H.N. Ridley 14165 (SING,

barcode SING0003915): Seletar, Nov? 1889, H.N. Ridley 2730 (SING, barcode SING0003916):

Kranji, 26 Feb 1889, H.N. Ridley s.n. (SING barcode SING0003914); Seletar, 10 Mar 1890,

J.S. Goodenough s.n. (SING barcode SING0003919); Woodlands, Jun 1903, H.N. Ridley s.n.

(SING barcode SING0003917); Upper Seletar track, 3 May 2011, Rodda, Leong-Skornickova,

Tran, Liew & Chew MR1I-008 (SING, spirit barcode SINGO126958).

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

research, in particular the Urban Ecology RF programme, which, through the project on

Enhancement of Biodiversity Research for Gingers, enabled extensive surveys across Singapore

leading to this unexpected discovery. We would also like to acknowledge Art Vogel and Ed de

Vogel for allowing access to the extensive living Dischidia collection in the Hortus Botanicus

Leiden.

References

Blume, C. (1826) Asclepiadeae. Bijdr. Fl. Ned. Ind. 16: 1048-1066.

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

Singapore: Native, Naturalised and Cultivated Species. Raffles Museum of Biodiversity

Research, National University of Singapore, Singapore. 273 pp. Uploaded 12 November

2009. http://rmbr.nus.edu.sg/raffles museum_pub/flora_of singapore_tce.pdf.

Decaisne, J. (1844). Dischidia. In: De Candolle, A. P., Prodr. 8: 631—633.

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

(eds) The Singapore Red Data Book: Threatened Plants and Animals of Singapore. 2nd

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

Forster, P.I., Liddle, D.J. & Nicholas, A. (1996) Asclepiadaceae. In: Robertson, R. et al. (eds)

Flora of Australia 28. Gentianales: 197-283. Melbourne: CSIRO.

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

Calcutta: Botanical Survey of India.

Janzen, D.H. (1974) Epiphytic myrmecophytes in Sarawak: mutualism through the feeding of

plants by ants. Biotropica 6: 237-259.

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

Flora of China 16. Gentianaceae through Boraginaceae: 189-270. St. Louis: Missouri

Botanic Garden.

Dischidia hirsuta in Singapore 299

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

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

Ridley, H.N. (1900) Flora of Singapore. J. Straits Branch Roy. Asiat. Soc. 33: 27-196.

Ridley, H.N. (1923) Hoya. Flora of the Malay Peninsula. 2: 402—410. London: Reeve & Co.

Rintz, R.E. (1980) The Peninsular Malayan species of Dischidia (Asclepiadaceae). Blumea 26:

81-126.

Tan, H.T.W., Tan, K., Ali Ibrahim, Chew, P.T., Chua, K.S., Duistermaat, H., Ganesan, S.K.,

Goh, M.W.K., Gwee, A.T., Kiew, R., Lee, S.M.L., Leong, P., Lim, J., Lok, A-F.S.L., Loo,

A.H.B., Lum, S.K.Y., Morgany, T., Saifuddin Suran, Sim, S., Samsuri Ahmad, Wee, Y.C.,

Yap, K.F., Yeo, C.K. & Yong, J.W.H. (2008) Checklists of Threatened Species—Seed

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

2nd ed. Singapore: The Nature Society (Singapore). Pp. 213-244.

Wanntorp, L., Kocyan, A., van Donkelaar, R. & Renner, S.S. (2006a) Towards a monophyletic

Hoya (Marsdenieae, Apocynaceae): Inferences from the chloroplast trnL region and the

rbcL-atpB spacer. Syst. Bot. 31(3): 586-596.

Wanntorp, L., Kocyan, A. & Renner. S.S. (2006b). Wax plants disentangled: A phylogeny of

Hoya (Marsdenieae, Apocynaceae) inferred from nuclear and chloroplast DNA sequences.

Molec. Phylogen. Evol. 39: 722-733.

Wanntorp, L. & Kunze, H. (2009) Identifying synapomorphies in the flowers of Hoya and

Dischidia—towards phylogenetic understanding. Jnt. J. Pl. Sci. 170(3): 331-342.

Wanntorp, L., Gotthardt, K. & Muellner, A.N. (2011) Revisiting the Wax Plants (Hova,

Marsdenieae, Apocynaceae) phylogenetic tree using the chloroplast markers matK gene

and psbA-trnH intergenic spacer. Jaxon 60(1):4—14.

Gardens’ Bulletin Singapore 64(2): 301-332. 2012 301

Studies in Malesian Gentianaceae I:

Fagraea sensu lato—complex genus or several genera?

A molecular phylogenetic study

M. Sugumaran' and K.M. Wong?

'Rimba I]mu Botanic Garden,

Institute of Biological Sciences, University of Malaya,

50603 Kuala Lumpur, Malaysia

maran(@um.edu.my

Singapore Botanic Gardens,

1 Cluny Road, Singapore 259569

wkm2000@gmail.com

ABSTRACT. Phylogenetic studies of Fagraea s.l. based on maximum parsimony and

Bayesian analyses of gene sequences for the nuclear ITS region and a number of chloroplast

regions (¢rnL intron, t7nL—-F spacer and two partial sequence regions of ndhF) were carried out.

Separate experiments with an ingroup of 29 taxa of Fagraea s.|. (8 from section Cyrtophyllum,

16 from section Fagraea and 5 from section Racemosae; all new sequences) were made with

individual gene-region and combined data sets; and with 43 taxa using only an ITS data set that

included published gene sequences of other recently revised, well-established genera of the

same tribe (Potalieae). Reasonably consistent clade composition was obtained with all analyses:

two clades could be equated to sections Fagraea and Racemosae, another two (Elliptica

and Gigantea clades) are different portions of the section Cyrtophyllum, and the solitary F.

crenulata resolved basal to the Fagraea clade in the chloroplast gene analyses but was a distinct

lineage in a polytomy with the Fagraea, Racemosa and Gigantea clades in the ITS analyses.

The equivalence of these clades and the F- crenulata lineage to other monophyletic groups

represented by established genera in the expanded-ITS analysis, as well as considerations of

potential morphological synapomorphies for these individual entities, suggest that Fagraea s.1.

is too morphologically and phylogenetically divergent to be considered a single genus.

Keywords. Cyrtophyllum, Fagraea, generic circumscription, Gentianaceae, Malesia, molecular

phylogenetic analyses, morphology, Potalieae, Racemosae, synapomorphic characters

Introduction

Fagraea Thunb. and s.|. are paleotropical, with a distribution from Sri Lanka and India,

through tropical South East Asia, reaching east to Polynesia (Struwe et al. 2002). The

genus (or group) is centred in Malesia, where over 70 species are present (Struwe et al.

2002), and around SO species are distributed within the Malay Peninsula and Borneo

(Leenhouts 1962; Wong & Sugau 1996). Species of Fagraea s.|. represent a variety

of life forms. They are tall canopy trees, smaller understory trees reaching only a few

meters in height, shrubs, or are epiphytes and hemi-epiphytes. They occur from sea

level to about 3000 m in very moist montane conditions, mostly in forest gaps, forest

edges, rocky outcrops, along stream beds in wet tropical forests, and less commonly in

302 Gard. Bull. Singapore 64(2) 2012

mesic forests, mangrove swamps and savannas (Motley 2004).

With a large diversity in habit and form, Fagraea s.1. includes species that are

both conspicuous and ecologically important in natural landscapes. The widespread

F. fragrans Roxb. is a common pioneer on sandy sites, a frequent secondary forest

species in the lowlands, and can persist in mature forest. In the Malay Peninsula, F.

racemosa Jack ex Wall. is also a common secondary forest species and the large-

leafed F. auriculata Jack with long-tubed flowers is often conspicuous in coastal

sandy sites and on quartz ridges in the lowlands. Throughout the lowland and lower

montane forests of the Malay Peninsula and Borneo, the frequent presence of Fagraea

epiphytes or hemi-epiphytes is detected by fallen corollas on the ground at different

times during the year. Burkill (1936), Macmillan (1991), Motley (2004), Perry (1980),

Quisumbing (1978) and Watson (1935) describe how a number of species are used

mainly for wood, medicinal purposes and horticulture.

In the past, Fagraea s.1. was placed within the Loganiaceae (e.g., Leeuwenberg

& Leenhouts 1980; Leenhouts 1962) but this classification is now controversial.

Subsequent analyses of morphological data (Struwe et al. 1994; Struwe & Albert 1997)

and molecular characteristics (Struwe et al. 2002; Downie & Palmer 1992; Olmstead

et al. 1993) have demonstrated that Fagraea s.]. and its tribe Potalieae are better

placed in the Gentianaceae. There is also support for this from some phytochemical

evidence (Jensen 1992; Jensen & Schripsema 2002). Potalieae members have diverse

flower merosity (e.g., 3-merous in the genus Pycnosphaera Gilg, up to 16-merous in

the genus Anthocleista Afzel. ex R.Br.) and habit (trees, lianas, shrubs, scramblers

and herbs) compared to most other tribes within Gentianaceae. The distribution of

Potalieae is strictly tropical, mostly found around the equatorial regions (Struwe et al.

2002).

Molecular analyses have consistently shown that Anthocleista, Fagraea and

Potalia Aublet form a monophyletic group that justifies inclusion in the same subtribe

(Potaliinae) within Gentianaceae (Struwe et al. 2002; Molina & Struwe 2009). Whereas

Fagraea s.\. is a large Indo-Pacific group, Pofalia is found in the Central and South

American region with nine recorded species and Anthocleista is only found in tropical

Africa and Madagascar with 14 known species (Struwe et al. 2002). The only two clear

morphological (non-synapomorphous) characters that make the subtribe Potaliinae

aberrant in Gentianaceae are the occurrence of fleshy berries and occasional large tree

habit in some taxa (Molina & Struwe 2009). Other members of the Gentianaceae have

dry and capsular fruits and relatively smaller size; and there are trees in other gentian

genera, but typically much less than 10 m high (Struwe et al. 2002).

Within Fagraea, three subgeneric groups have been recognised, considered

as the sections Cyrtophyllum (Reinw.) Blume, Fagraea and Racemosae Benth.

(Leenhouts 1962). This classification was adopted by Wong & Sugau (1996), who

delimited the sections by inflorescence form and branching, seed form, the nature of

fruit epidermis, axillary scale characters and stigma form.

In their review for Borneo, Wong & Sugau (1996) enumerated 42 species,

including 20 that they newly described. This was a big contrast to the previous work by

Leenhouts (1962, 1984) for Malesia, where he only enumerated 15 species for Borneo.

1S)

Molecular phylogeny of the Fagraea complex 30

The species concepts of Leenhouts were considered too broad by Conn & Brown

(1993) and Wong & Sugau (1996). An example stated by Wong & Sugau (1996) is

that Leenhouts (1962) had only one species accepted in the whole section Racemosae

for Malesia, having reduced many previously described species to synonymy. Wong

& Sugau (1996) found many of such taxa morphologically distinct and possible to

key out using both vegetative and flower and fruit characters, and resurrected many

corresponding names from synonymy.

Scope of the present study

The bizarre contrast between the results obtained by Leenhouts (1962) and Wong

& Sugau (1996), the demonstration of what appears to be likely impractical species

concepts in Leenhouts (1962) by both Conn & Brown (1993) and Wong & Sugau

(1996), and the lack of any recent detailed reviews for Malesia other than Borneo,

precipitated the present inquiry. Just as species concepts were apparently too broad, so

the inclusion of morphologically very different sections in the same genus was perhaps

also questionable.

At the fundamental level, the distinctness of the subgeneric groups recognised

as Fagraea sections Cyrtophyllum, Fagraea and Racemosae require further

consideration. This is in view of somewhat conspicuous characters distinguishing

them, as summarised by Wong & Sugau (1996), and very preliminary possibilities

for generic distinction suggested by limited molecular evidence discussed by Struwe

& Albert (1997). This problem is investigated using a molecular analysis of as many

taxa as possible from the Malay Peninsula and Borneo, and augmented by molecular

results for other taxa in the same subtribe and tribe from other geographical regions.

The extents of morphological distinction among the so-called sections are also re-

examined.

Finally, the merits and intricacies of classifying the several morphologically

distinct groups of species within Fagraea s.l. as either sections or distinct genera

will be discussed. From this, recommendations would be made for an appropriate

classification of the group.

Sectional classification of Fagraea s.1.

The recognition of infrageneric groups within Fagraea has a convoluted history.

Blume (1838) erected Fagraea section Cyrtophyllum, which included the two species

Cyrtophyllum peregrinum Reinw. (this is in fact a synonym of the earlier published and

better known F. fragrans) and C. speciosum Blume (synonymous with Picrophloeus

javanensis Blume, to which also some authors have applied the name F elliptica

Roxb.). Blume had evidently done this because the many-branched cymes with much

smaller flowers in these taxa were rather different from the sparsely branched cymes

with larger flowers of F- ceilanica Thunb., the type species of the genus.

304 Gard. Bull. Singapore 64(2) 2012

Later, Blume (1850) included two of his species, -) kimangu Blume and

F, picrophloea Blume (both synonyms of his P. javanensis) in his Fagraea section

Eufagraea, with a number of other species that had a very different inflorescence form,

where cyme-like clusters of flowers were borne along an elongate main inflorescence

axis that superficially resembled a complex raceme (described by Blume as “cymis

in racemum terminalem longissimum’). In the same paper, Blume (1850) placed F:

ceilanica (spelled as F- zeylanica) and associated species in “section Fagraea verae”’.

Bentham (1856) appears to be the first person who clearly differentiated

three groups within Fagraea, as section Parviflorae Benth. (= Blume’s section

Cyrtophyllum), section Racemosae (whichis that group with raceme-like inflorescences

in Blume’s section Ewfagraea), and section Corymbosae Benth. (= Blume’s section

Fagraea verae). Subsequent authors also provided infrageneric names. Miquel (1857)

employed subgenus Cyrtophyllum and subgenus Eufagraea, in place of Blume’s

sections Cyrtophyllum and Fagraea verae, respectively. Solereder (1892), on the other

hand, named section Pseudoracemosae and section Pseudocorymbosae for Bentham’s

Racemosae and Corymbosae, respectively.

In his revision for the Flora Malesiana, Leenhouts (1962) recognised three

infrageneric groupings that he called sections Fagraea (containing the type and by far

the largest number of species), Cyrtophyllum (as designated by Blume) and Racemosae

(as proposed by Bentham). Leenhouts (1962) gave very few characters for sectional

distinction, some of which were somewhat inconsistent (see Table 1).

Wong & Sugau (1996) also used these sections as circumscribed by Leenhouts

(1962) for their account of Fagraea in Borneo, but added more distinguishing

characters and provided a key to sections. They used such characters as fruit size, ease

of epidermis detachment on drying, seed form, inflorescence branching, characters

of the petiole base (including whether scale-like structures develop at the leaf axils),

stigma form and the extent of stamen and style exsertness in open flowers.

Alternative taxonomic interpretations

Several species placed in Fagraea s.l. by Leenhouts (1962) and other authors have

been the basis of other generic names. Although Cammerloher (1923) considered

Cyrtophyllum a synonym of Fagraea, Ridley (1923) had distinguished the two

genera, using the former name for C. /anceolatum (Wall.) DC., C. peregrinum and C.

giganteum (Ridl.) Ridl. The equivalents for these names in Fagraea are F: lanceolata

Wall. (a synonym of F) wallichiana Benth.), F) peregrina Blume (a synonym of F:

fragrans), and F: gigantea Ridl., respectively.

Leenhouts (1962) suggested that Cyrtophyllum and Picrophloeus Blume are

synonyms of Fagraea section Cyrtophyllum. He also placed the names Utania G.Don,

Kuhlia Reinw. and Kentia Steud. (the latter two illegitimate as a later homonym and a

name lacking a description, respectively) as synonyms of Fagraea section Racemosae.

Molecular phylogeny of the Fagraea complex

Table 1. Diagnostic characteristics of three sections in Fagraea s.]. according to Leenhouts (1962).

Character / Section

leaves

stipules

inflorescence form

inflorescence

position

flower size

corolla form

stamens, style

fruit shape, size

sect. Fagraea

auriculate or not

(not explicitly stated)

corymbose, dichasial.

glomerulous or with

solitary flowers;

“usually with a pair

of strong branches

in the upper leaf

axils, therefore the

inflorescence as a

whole mostly sessile”

always terminal

(not explicitly stated)

hardly or not exserted

(not explicitly stated)

sect. Cyrtophyllum

not auriculate

(not explicitly stated)

corymbose

(not explicitly

stated, but including

terminal and axillary

types in different

taxa)

small

tubular

far exserted (except

in F. umbellifiora)

globular, small

sect. Racemosae

not auriculate

connate in an ochrea

racemiform, with a

number of decussate

pairs of small cymes

always terminal

(not explicitly stated)

not or only slightly

exserted

(not explicitly stated)

Materials and methods

Herbarium materials and studies. Study specimens were loaned from, or examined

at, the following herbaria: Royal Botanic Gardens, Kew (K), Forest Research Institute

Malaysia, Kepong (KEP), Nationaal Herbarium Nederland (L), Forest Research

Centre, Sabah (SAN), Singapore Botanic Gardens (SING) and the University of

Malaya (KLU). A handlens (x20 magnification) and binocular microscope (40

magnification) were used to examine material.

Field collections and processing. Additional material collected yielded specimens

that were oven-dried at 55°C for 24 weeks: and flowers, fruits and leafy branches

preserved in AWG solution (70% alcohol + 28% distilled water + 2% glycerol) for

306 Gard. Bull. Singapore 64(2) 2012

study. Leaf material of 30 taxa of Fagraea collected from various locations were

preserved in silica gel for subsequent DNA extraction (Table 2).

DNA extraction, gene regions and primers. Total DNA extraction from silica-dried

or fresh leaf tissue samples followed the CTAB (cetyltrimethylammonium bromide)

method of Doyle & Doyle (1987), or made use of DNeasy plant kits (Qiagen)

following the manufacturer’s protocol (Qiagen 2003-2009). Gene regions chosen for

the phylogenetic analyses were: ITS (Internal Transcribed Spacer) (nuclear) and the

trnL intron, t7nL—F spacer and ndhF region (chloroplast). These regions have been

widely used to study phylogenetic relationships at the family level and lower (Chassot

et al. 2001; Davis et al. 2001; Gielly & Taberlet 1996; Ranker et al. 2003; Struwe et

al. 2002; Thiv et al. 1999; Hagen & Kadereit 2001; Yuan & Kiipfer 1995, 1997; Yuan

et al. 1996, 2003). The whole ITS region (ITS1, 5.88 rDNA & ITS2) was amplified

with universal primers: ITS 1 (forward) and ITS 4 (reverse), following White et al.

(1990). For the trnL intron and trnL-F regions, universal primer sets (C & D and E &

F, respectively) given by Taberlet et al. (1991) were used. Two new primer sets for the

ndhF gene sequences were constructed at conserved regions following alignment of

ndh¥ sequences for three taxa obtained from the GenBank: Fagraea sp. (AJ 235830),

Anthocleista grandiflora (AJ 235829) and Potalia resinifera (AJ 235831). Each of

these two sets of primers (GBI Fwd & GBI Rev and GB2 Fwd & GB2 Rev) was

estimated to amplify about 900-1000 bp of the ndhF gene. Sequences of all primers

used in this study are listed in Appendix A.

Polymerase Chain Reaction (PCR). PCR for all the regions were performed in

a Whatman Biometra T Gradient or Perkin Elmer GeneAmp PCR System 9600

thermocycler. A total reaction mix of 50 ul was used, containing 25 ul of PCR master

mix (2x Go Taq* Green Master Mix), 2 ul (SO mM) each of forward and reverse

primer, I|—3 ul of DNA template and RNAse free water topped up to a final volume of

50 ul. Parameters for PCR amplification were: | cycle of 3 min at 94 °C, linked to 30

cycles of 10s at 94°C, 20s at 55°C, 90 s at 72°C, followed by 4 min at 72°C to complete

primer extension. PCR-amplified samples were electrophoresed on agarose gel and

samples that had a clear single desired band were selected for sequencing. Purification

of PCR products used the QIlAquick PCR purification kit (Qiagen) following the

manufacturer’s protocol. When multiple bands of different sizes were observed in

some cases, a gradient PCR (with different annealing temperatures) was carried out

to find the optimal temperature that amplified a single desired band. The sample with

such band was then selected for purification and sequencing. Occasionally, samples

still produced multiple bands after PCR despite optimising the annealing temperature.

For these, the PCR product was electrophoresed on agarose gel (1.2—1.5 %) for about

60 to 90 min. The target band was then excised and purified using a Qiaquick Gel

extraction Kit (Qiagen) following the manufacture’s protocol. This product was then

sequenced.

Molecular phylogeny of the Fagraea complex 307

Table 2. Voucher specimens and GenBank accession numbers of sequences for Fagraea s.1.

taxa representing the different “sections” collected for the present study.

Fagraea sp. A, F- sp. B and F. sp. C are new Malay Peninsula taxa diagnosed in the present

study, whereas two other unidentified Bornean taxa were labeled as F: sp 1 and F. sp 2. Fagraea

elliptica is represented by two accessions from Peninsular Malaysia and Borneo. Sections: C

(Cyrtophyllum), F (Fagraea), R (Racemosae). Vouchers with ‘SAN’ numbers were deposited

in the Forest Research Centre Sandakan, Sabah (SAN); all other vouchers were deposited in

the University of Malaya herbarium (KLU). The two partial non-overlapping ndhF sequences

are represented with two GenBank accession numbers.

GenBank accessions for

oe peetionea! Neucher. specimen (ITS) — (trnL—F) — (ndhF: 2 partial sequences)

F. belukar e Postar & Ahmad SAN 147987 JX283355—JX217749-JX283385 & JX283414

F. caudata Cc Low LYW 213 JX283356—-JX217750—JX283386 & JX283415

F: collina Low LY W 260 JX283357—JX217751—JX283387 & JX283416

F elliptica | Low LY W 358 JX283358—-JX217752—JX283388 & JX283417

F. elliptica 2 Postar et al. SAN 147993 JX283359-JX217753—JX283389 & JX283418

F. fragrans Sugumaran SM 212 JX283360—JX217754-JX283390 & JX283419

F. gigantea Sugumaran SM 193 JX283361—JX217755—JX283391 & JX283420

F. wallichiana

F. auriculata

F. carnosa

F. crassifolia

F. crenulata

F. curtisii

F. gardenioides

F. imperialis

F. oblonga

F. renae

F. resinosa

F. ridleyi

F. splendens

Low LY W 206

Sugumaran SM 240

Lee DLKP 30

Low LYW 244

Sugumaran SM 246

Low LYW243

Sugumaran SM 170

Sugumaran SM 238

Sugumaran SM 165

Sugumaran SM 177

Postar et al. SAN 147998

Low LYW 227

Zahid ZMS 42

JX283362—JX217756—JX283392 & JX283421

JX283363—-JX217757—JX283393 & JX283422

JX283364—JX217758—JX283394 & JX283423

JX283365—JX217759-JX283395 & JX283424

JX283366—-JX217760—JX283396 & JX283425

JX283367—JX217761—JX283397 & JX283426

JX283368—JX217762—JX283398 & JX283427

JX283369-JX217763-JX283399 & JX283428

JX283370—-JX217764—JX283400 & JX283429

JX283371—JX217765—JX283401 & JX283430

JX283372—-JX217766—JX283402 & JX283431

JX283373-JX217767—JX283403 & JX283432

JX283374-JX217768—-JX283404 & JX283433

F. stonei Wong et al. SAN 147989 JX283375—JX217769-JX283405 & JX283434

F sp. A Low LYW 138 JX283376—-JX217770—-JX283406 & JX283435

F. sp. B Sugumaran SM164 JX283377-JX217771—JX283407 & JX283436

F. sp. 1 Postar et al. SAN 149702 JX283378—-JX217772—-JX283408 & JX283437

F. cuspidata

F. racemosa

F. spicata

F. volubilis

Busps ©

F. sp. 2

rofl tzof. oll lof Ny of rofl Mas|P ool Meol Saolitash oof tool Teale tleols tool) tesla gest Glog tool leo tool fey ie) (oe) (my @) .(@)

Seligi & Lingkong SAN 145303

Sugumaran SM 248

Postar & Ahmad SAN 147985

Sugumaran SM 206

Sugumaran SM 201

Postar SAN 149705

JX283379-JX217773-JX283409 & JX283438

JX283380—JX217774—JX283410 & JX283439

JX283381—-JX217775—JX283411 & JX283440

JX283382

JX283383—-JX217777—-JX283412 & JX283441

JX283384—-JX217778-JX283413 & JX283442

308 Gard. Bull. Singapore 64(2) 2012

Sequencing PCR products and data authentication. PCR products were sequenced

on an Applied Biosystems 3730xl1 DNA Analyser with BigDye*Terminator ver. 3.1

Sequencing Kit with: 25 cycles of 96°C for 10 s, 50°C for 5 s and 60°C for 4 min, at

rapid thermal ramp for 1°C/s. Sequencing in both directions used the same primers

as for PCR. All sequence data were manually checked by eye with the corresponding

electrophoregrams. Ambiguous base pairs (those with multiple peaks) were cross-

checked with the complementary sequence data (1.e., forward sequence was cross-

checked with reverse sequence and vice versa). This could be done because forward

and reverse strands that were sequenced had an approximate overlap of about 80%.

Samples that had very noisy sequence data were not used in the analyses.

Sequence alignment. Sequence data were initially aligned using ClustalX 2.0.10

(Larkin et al. 2007). Aligned sequences were then manually adjusted using the software

Bioedit (Hall 1999). The boundaries of the ITS, trnL intron, trnL—F spacer and ndhF

were determined by comparison with the published outgroup sequence, Anthocleista

grandiflora, that was also used in the analyses (Table 3). The ¢rnL intron and trnL—F

spacer sequences were aligned and combined into a single data matrix (hereafter

referred to as the “trnL—F data set”). Numerous single and multibase insertions or

deletions (indels) were introduced for the ITS and ¢rnL-F data sets but no indels were

needed for aligning the ndhF data set. The aligned data was then saved in a Nexus file

format and phylogenetic analyses were performed using PAUP* version 4.0b4a for

Macintosh (Swofford 2001) and MrBayes v.3.1 (Huelsenbeck & Ronquist 2001).

Phylogenetic assessments: scope and experimental design. Two phylogenetic

assessments were conducted. The first assessment included analyses with individual

data sets (ITS, tmL—F or ndhF) and a combined data set (ITS + trnL—F + ndhF).

This assessment aims to evaluate if (a) reasonable support can be found for any

monophyletic groups forming within the ingroup; (b) if such clades correspond to

the sections of Fagraea s.|. circumscribed by existing classification; and (c) 1f these

clades are consistent (or congruent) among the topologies representing the different

gene regions. The combined data analysis was performed to evaluate whether clades

were better resolved in comparison with single-gene analyses. This assessment was

performed with 29 taxa of Fagraea s.1. (8 from section Cyrtophyllum, 16 from section

Fagraea and 5 from section Racemosae) and one taxon as outgroup (A. grandiflora).

The 29 DNA sequences of Fagraea s.1. from all four gene regions included in these

analyses are new (details in Table 2). The sequences for 4. grandiflora were obtained

from GenBank (Table 3).

The second assessment was done with only the ITS data set where sampling

was expanded to 45 taxa. This assessment was done to evaluate whether clades formed

among so-called Fagraea species within the ingroup can be reasonably circumscribed

as distinct genera. The selected ingroup taxa are all from Potalieae and include the

well recognised genera Anthocleista and Potalia (in the same subtribe, Potaliinae,

as Fagraea) as well as Lisianthius (subtribe Lisianthiinae) (Weaver 1972; Sytsma

1988). The two outgroup taxa belong to two other distantly related tribes within the

Molecular phylogeny of the Fagraea complex 309

Table 3. List of species for which GenBank accessions were included for this study, including

tribal and subtribal classification in Gentianaceae (Struwe et al. 2002) and voucher information.

Herbarium acronyms: E — Royal Botanic Garden Edinburgh, Edinburgh, Scotland, UK; HNWP

— Northwest Plateau Institute of Biology, Xining, Qinghai, China; F — Field Museum of Natural

History, Chicago, Illinois, USA; NEU — Université de Neuchatel, Neuchatel, Switzerland; VY

— New York Botanic Garden, Bronx, New York, USA; UPS — Uppsala University, Uppsala,

Sweden; US— Smithsonian Institutution, Washington, District of Columbia, USA.

GenBank accession

Taxa Current tribe—subtribe Voucher (Herbarium) :

(and gene region)

Anthocleista : - :

; ; Potalia—Potaliinae Woulhauser PBZT (NEU) AJ489863 (ITS)

amplexicaulis

AJ489864 (ITS)

Callmander s.n. (NEU) AJ490190 (trnL

Anthocleista Baie Panes Callmander s.n. (VEU) (UAA) intron)

grandiflora aca Callmander s.n. (VEU) AY251777

Exacum affine

Fagraea

berteroana

F. ceilanica

F. elliptica

F. macroscypha

F. salticola

Gentiana algida

Lisianthius

brevidentatus

Lisianthius

cuspidatus

Lisianthius

laxiflorus

Potalia amara

Potalia elegans

Exaceae

Potalia—Potaliinae

Gentianeae—Gentianinae

Potalia—Lisianthiinae

Potalia—Potaliinae

Bremer 3098 (UPS)

Miller et al. 6201 (£)

L. Struwe 1219 (NY)

L. Struwe 1300 (NY)

Takeuchi 7122 (NY)

Beaman et al. 8867 (US)

Pullen 326 (US)

Liu 1257 (HNWP)

Ortiz 1664 (F)

Lewis 895 (F)

Struwe & Specht 1153 (VY)

S. Mori 24123 (NY)

P. Berry 7434 (NY)

(trnL-F spacer)

AJ235829 (ndhF)

AJ489879 (ITS)

DQ449918 (ITS)

FJ23257 (ITS)

FJ232579 (ITS)

FJ232573 (ITS)

FJ232571 (ITS)

DQ398659 (ITS)

FJ32569 (ITS)

FJ32567 (ITS)

FJ232552 (ITS)

DQ449919 (ITS)

DQ449920 (ITS)

Potalia resinifera _ Potalia—Potaliinae B. Stahl 1872 (NY) DQ449921 (ITS)

Gentianaceae, namely, Exaceae (Exacum affine) and Gentianeae (Gentiana algida)

(Struwe et al. 2002). The ITS region was chosen because it was relatively more

informative than the other two data sets; moreover, many more ITS sequences were

available in GenBank compared to trnL—F (intron + spacer) and ndhF sequences. The

ingroup included 35 Fagraea taxa, two Anthocleista taxa, three Lisianthius taxa and

three Potalia taxa. Of the Fagraea sequences used in this assessment, 30 are new

310 Gard. Bull. Singapore 64(2) 2012

sequences, including F. volubilis (which was excluded from the first assessment as

sequencing results for the ndhF region was very poor) (Table 2). All other sequences,

including five sequences of Fagraea and the outgroup sequences, were obtained from

GenBank (Table 3).

Parsimony analyses in PAUP* were performed with heuristic searches where

all characters were unordered and unweighted (‘Fitch parsimony’; Fitch 1971).

All gaps in the sequence were treated as missing data. Starting trees were obtained

using simple stepwise addition sequences, with one tree held at each step with tree

bisection—reconnection (TBR) branch swapping algorithm, MULTREES option in

effect, accelerated transformation (ACCTRAN), branches with zero length collapsed

and topological constraints not enforced. Separate and combined analyses of the three

data sets (ITS, trnL—F and ndhF) were performed and strict consensus trees were

generated. The bootstrap method (Felsenstein 1985) was used to estimate robustness

of the various clades revealed in the consensus tree. Bootstrap values were estimated

from 1000 replicates of full-heuristic searches using simple addition sequence and

TBR branch swapping with a set ‘MAXTREES’ limit of 10000 trees per bootstrap

replicate. Branches less than 50% value were collapsed. The consistency index (CI)

(Kluge & Farris 1969), the retention index (RI) and rescaled consistency index (RC)

(Farris 1989) were also calculated using PAUP* as measures of character fit to the

phylogenetic trees.

For Bayesian inference analyses, an appropriate evolutionary model was

selected using MrModeltest 2.2 (Nylander 2004) together with PAUP* (Swofford

2001). The programme (MrModeltest 2.2) specifically tests the 24 models available

common to PAUP* and MrBayes (Huelsenbeck & Ronquist 2001) with a given data

set. The Akaike Information Criterion tests selected the General Time Reversible model

with gamma distribution of rates for all individual data sets (ITS, t*nL—F, ndhF and

ITS-expanded). Two independent runs of 1.5 million generations were performed each

with four MCMC (Markov Chain Monte Carlo) chains. One tree was sampled every

500 generations and the first 750 trees (burn-in=750) were excluded from the analyses

which amounts to 25% of trees sampled. A 95% credible set of trees was generated by

including all trees with the highest posterior probabilities until the cumulative posterior

probabilities (PP) was 95% (Huelsenback et al. 2001). A 50% majority rule consensus

of these generated trees was used to estimate the PP of each clade. PP values above

p=0.95 were considered to be statistically significant (Huelsenback & Ronquist 2001;

Larget & Simon 1999; Lewis 2001; Rannala & Yang 1996; Kauff & Lutzoni 2002).

The incongruence length difference test (ILD; Farris et al. 1994, 1995) as the

partition homogeneity test (PHT) implemented in PAUP* was employed to test the

null hypothesis that the three data sets (ITS, t*nL—F and ndhF) were homogeneous

with respect to phylogenetic information. PHT was performed with 100 replicates of

heuristic searches, Maxtree=1000 and TBR branch swapping. It has been suggested

that PHT p values more than 0.01 indicate that the data sets are congruent and if

combined will either improve or will not affect the accuracy of the phylogenetic

information (Cunningham et al. 1998).

Molecular phylogeny of the Fagraea complex 311

Results

Sequence variation

The aligned data matrix of the ITS data set with 30 taxa had 639 characters. Out

of these, 89 characters were variable but parsimoniously uninformative whereas 64

characters (10% of total characters) were phylogenetically informative. The unaligned

length of ITS sequences among the 30 taxa of Fagraea (Table 2) used in this study

varied from 622 to 631 bp. These data resulted in uncorrected pairwise sequence

divergences ranging from 0 (F. sp 1 vs. F. gardenioides, F. curtisii vs. F. crassifolia)

to 13.9% (F. oblonga vs. F. sp. 2).

The individual trnL—F sequences (frnL intron + trnL-F spacer) in Fagraea

used in this study ranged from 652 to 836 bp. The length varied mainly at several

A-T rich regions where alignment could not be readily done due to repeated motifs.

These highly variable regions which accounted for 156 bp were excluded from the

analyses. The final aligned t#7nL—F data matrix had a total of 811 characters where 60

characters were variable and parsimoniously uninformative, and 49 characters (6% of

total characters) were phylogenetically informative. These data resulted in uncorrected

pairwise sequence divergences ranging from 0 (F auriculata vs. F- imperialis, F-

gigantea vs. F- caudata, F- wallichiana vs. F- caudata, F- wallichiana vs. F- gigantea,

F- spicata vs. F- cuspidata, F- sp. C vs. F- cuspidata and F: spicata vs. F: sp. C) to 3.5%

(F. stonei vs. F. fragrans).

The two new primer pairs that were used to amplify two partial sequence

regions of ndhF produced 1832 bp in total. The primer pair “GB1” produced 899 bp

whereas the “GB2” pair produced 933 bp. Based on the ndhF sequence of A. grandiflora

(GenBank Acc. AJ235829), the 899 bp sequence data corresponds with positions 161 to

1059 whereas the 933 bp sequence data corresponds with positions 1182 to 2114. The

ndhF sequences were easily aligned and the total 1832 characters had 114 characters

that were variable and parsimoniously uninformative whereas 81 characters (4.4%

of total characters) were informative. These data resulted in uncorrected pairwise

sequence divergences ranging from 0 (F- racemosa vs. F- sp. C, F- cuspidata vs. F. sp.

C, F. sp. 2 vs. F sp. C, F. splendens and F. sp. B and F- imperialis vs. F: auriculata) to

2% (F. resinosa vs. F- sp. C, F- resinosa vs. F: racemosa, F- resinosa vs. F: spicata, F-

resinosa VS. F. sp. 2).

The combined data set of ITS, #mL—F and ndhF resulted in 3282 characters.

The data matrix had 263 variable and parsimoniously uninformative characters

whereas 194 characters (5.9% of total characters) were informative.

The aligned matrix of the expanded ITS data set (45 taxa) had 697 characters

where 131 characters were variable and parsimoniously uninformative, and 185

characters (26.5% of total characters) were phylogenetically informative. These data

resulted in uncorrected pairwise sequence divergences ranging from 0 (F. ridlevi vs.

F. gardenioides, F: sp. 1 vs. F. gardenioides, F- sp. 1 vs. F- ridleyi, F. curtisii vs. F-

crassifolia and F. macroscypha vs. F. auriculata) to 14% (F. sp. 2 vs. F. imperialis).

312 Gard. Bull. Singapore 64(2) 2012

Phylogenetic analyses

Data set characteristics are presented in Table 4. In all the analyses, parsimony and

Bayesian tree topologies were generally congruent and bootstrap (BS)-supported

nodes mostly also had high support of posterior probabilities (PP). The trees presented

were obtained from maximum parsimony analyses and congruent branch support

values obtained from the Bayesian analyses (PP) are stated.

Analyses of the ITS data set

Parsimony analyses of the 30-taxon ITS data set resulted in 3520 trees with a tree

length of 211 (strict consensus shown in Fig. 1), CI (Consistency Index) of 0.8152 and

RI (Retention Index) of 0.8691. Four monophyletic groups can be recognised, referred

to as the Fagraea, Racemosa, Gigantea and Elliptica clades. In these analyses, all

included taxa representing Fagraea section Fagraea were represented on the Fagraea

clade except F. crenulata, which is placed in a tetrachotomy with three other clades.

Within the Fagraea clade, two monophyletic sister groups are formed where one group

is smaller, consisting of three taxa, and the rest form a bigger group with 12 taxa. The

3-taxon group is well-supported (BS = 100%; PP = 1.00) with F- resinosa sister to the

branch with F) auriculata and F- imperialis. Within the 12-taxon group, there are two

smaller well-supported subgroups (F: crassifolia and F. curtisii; F- splendens and F. sp.

B) as well as other species whose relationships are unresolved.

Species included in the study representing section Racemosae (Racemosa

clade) form a strongly supported monophyletic group (BS = 100%; PP = 1.00). Within

the Racemosa clade, F’ sp 2 is sister to an equivocal clade of only moderate support

(BS = 69%) with four unresolved taxa. Section Cyrtophyllum is paraphyletic with

the eight representative taxa split into two distinct monophyletic groups. The groups

comprising four taxa each are referred to as the Elliptica and Gigantea clades. The

Elliptica clade is sister to the rest of Fagraea s.|. and well-supported with BS (91%)

and moderately supported with PP (0.82) values. Within the Elliptica clade, F) belukar

is sister to a clade with moderate BS (65%) but strong PP (0.99) support containing

the three other taxa. The Gigantea clade is well supported (BS = 81%; PP = 1.00) and

also well resolved where F: fragrans is the most basal taxon. The branches within this

clade received moderate BS and good PP support.

Analyses of the trnL—F data set

The resulting 7224 most parsimonious trees with the #7nL—F data set were 135 steps

long (strict consensus shown in Fig. 2) with a CI of 0.8519 and RI of 0.9206. The

clades formed are similar to those observed for the ITS tree; however, the interspecific

relationships within each clade were less resolved with the trnL-F data set. Again, as in

the ITS tree, F crenulata was outside of the Fagraea clade. Fagraea crenulata is basal

to both the Elliptica and Fagraea clades together and the relationship is well supported

(BS = 82%; PP = 1.00). The rest of the species representing F) section Fagraea (the

Fagraea clade) form a strongly supported clade (BS = 99%; PP = 1.00). Within the

Fagraea clade, F’ sp. A and F. gardenioides were unresolved, F. crassifolia and F:

curtisii form a strongly supported group (BS = 100%; PP = 1.00) and the rest also form

Molecular phylogeny of the Fagraea complex 313

Table 4. Characteristics of the parsimony-based analyses with individual and combined data sets.

trnL—F ndhF ITS

Characteristics ITS (trnL intron + (2 combined ITS +tmL-F (expanded

trnL-F spacer) _ partial regions) + ndhF data set)

Number of taxa 30 30 30 30 45

Total characters 639 811 1832 3282 697

Constant characters 486 702 1637 2825 381

escniony 64 49 81 194 185

informative characters

Variable characters 89 60 114 263 131

Most parsimonious 3520 7204 1 4 145

trees

Tree length 211 135 242 591 582

Consistency Index, CI

(values excluding 0.82 0.85 0.83 0.83 0.73 (0.65

uninformative sites in (0.66) (0.73) (0.68) (0.68) Bie eee

parentheses)

Retention Index, RI 0.87 0.92 0.91 0.90 0.81

ee tatty 97 0.78 0.76 0.74 0.59

Index, RC

a well-supported group (BS = 84%; PP = 1.00). In this latter group, F. ridlevi is placed

in a trichotomy with two well-supported subclades. These subclades have three and

seven taxa, respectively. The 3-taxon subclade comprising of F. resinosa, F- imperialis

and F. auriculata were well resolved where F- resinosa was sister to the other two.

In the 7-taxon subclade, F’ sp. B and F- splendens formed a strongly supported group

while the rest were unresolved.

Section Cyrtophyllum ts paraphyletic, with four out of eight species represented

forming the Elliptica clade and three other species forming the Gigantea clade. The

Elliptica clade was basal to the Fagraea clade and the relationship was only moderately

supported with BS (60%) but well supported with PP (0.99) values. The Elliptica clade

itself was only weakly supported with 51% BS and a moderate PP support (0.93). The

relationship within the Elliptica clade was unresolved.

The Gigantea and Racemosa clades plus F. fragrans formed a well-supported

group (BS = 97%; PP = 1.00). The Gigantea clade received strong branch support (BS =

97%; PP = 1.00) but was internally unresolved. The remaining taxon, F. fragrans, was

placed in a trichotomy with the Racemosa and Gigantea clades. The Racemosa clade

included all the five representative taxa of section Racemosae and was monophyletic

with moderate BS (67%) and good PP (0.96) support. The F. cuspidata and F. spicata

subgroup within the Racemosa clade received moderate BS (66%) and good PP (0.99)

support; the other three taxa were not resolved.

314

1.00

0.98

100

1.00

100

1.00

0.93

0.99

Gard. Bull. Singapore 64(2) 2012

F. oblonga

F. gardenioides

F. ridleyi

F. crassifolia

F. curtisii

F. sp. 1

F. splendens

F.sp.B

F. carnosa

F. stonei

F.sp.A

F. renae

F. auriculata

F. imperialis

F. resinosa

F. racemosa

F. sp. c

F. cuspidata

F. spicata

Fispyz

F. fragrans

F. gigantea

F. wallichiana

F. caudata

F. crenulata

F. belukar

F. elliptica PM

F. elliptica B

F. collina

A. grandiflora

Fagraea

clade

Racemosa

clade

Gigantea

clade

Elliptica

clade

Fig. 1. Strict consensus of 3520 equally parsimonious trees based on the ITS sequence data. The

numbers above and below the branches denote Bootstrap and Bayesian Posterior Probability

values, respectively. Length (L) = 211; consistency index (CI) = 0.8152; retention index (RI) =

0.8691. A. = Anthocleista; F: = Fagraea. Different accessions of F: elliptica are indicated (PM

= Peninsular Malaysia, B = Borneo).

Molecular phylogeny of the Fagraea complex 315

F. resinosa

98 * . -

1.00 67 F. imperialis

0.84 F. auriculata

1.00 F. splendens

+. F. sp. 1

: Sei | Ficamosa

| 1.00

+——————— F. oblonga Fagraea

clade

+ F renae

ae, ee Tw) F. stonei

1.00 _ F ridleyi

F. gardenioides

F. curtisii

F. crassifolia

F.sp.A

F. elliptica PM

F. elliptica B

Elliptica

F. collina clade

F. belukar

F. crenulata

F. caudata

F. gigantea Gigantea

clade

F. wallichiana

F. cuspidata

F. spicata

F. sp. 2 Racemosa

F. sp. C ede

F. racemosa

Sana IF PEUPEINE

A. grandiflora

Fig. 2. Strict consensus of 7224 equally parsimonious trees based on the trnL—F sequence

data. The numbers above and below the branches denote Bootstrap and Bayesian Posterior

Probability values, respectively. Length (L) = 135; consistency index (CI) = 0.8519; retention

index (RI) = 0.9206. A. = Anthocleista; F. = Fagraea. Different accessions of F. elliptica are

indicated (PM = Peninsular Malaysia, B = Borneo).

316 Gard. Bull. Singapore 64(2) 2012

Analyses of the ndhF data set

Parsimony analysis of the ndhF data set produced a single most parsimonious tree with

tree length of 242 (Fig. 3), a CI of 0.8347 and RI of 0.9109. The four major clades

common to the previous two analyses were also observed here and all had good BS

and PP support. The F’ section Fagraea (Fagraea clade + F. crenulata), was a well-

supported monophyletic group, with F) crenulata basal within the clade. There were

two well-supported sister groups, with four and eleven taxa, respectively. The 4-taxon

group was well resolved with good branch support where F’ gardenioides was basal.

The 11-taxon group has two well supported subgroups: F. sp. B plus F. splendens; and

a subgroup with F. resinosa, F- imperialis and F- auriculata. These groups were also

observed in the analyses with the ITS and #7nL-F data sets. The relationships among

the other taxa in this clade were unresolved.

Section Cyrtophyllum was paraphyletic where four out of eight taxa formed

the Elliptica clade, while a further three taxa grouped to form the Gigantea clade. The

Elliptica clade was sister to the Fagraea clade, both forming a group with good branch

support (BS = 75%; PP 0.99). The Elliptica clade itself was well supported (BS = 97%;

PP = 1.00) and the interspecies relationships within the clade were well resolved with

moderate branch support.

The Gigantea clade was well supported and was sister to the Racemosa clade.

However, the interspecies relationships within the Gigantea clade were not resolved.

The Racemosa clade was monophyletic and received good branch support (BS = 100%;

PP = 1.00). Within the Racemosa clade, F. sp. C, F’ racemosa and F. sp 2 formed

a group with moderate BS (66%) but good PP (0.98) support. The remaining two

species were unresolved. Fagraea fragrans resolved as a basal taxon for the Gigantea

+ Racemosa clades.

Analyses of the combined ITS, trnL—F and ndhF data sets

The result of the ILD test was not significant (p=0.03), indicating that the null

hypothesis of data set homogeneity could not be rejected. The three data sets were thus

combined into a single matrix and parsimony and Bayesian analyses were performed.

Parsimony searches on the combined data set produced four trees having a tree length

of 591 (strict consensus shown in Fig. 4) with a CI of 0.8274 and RI of 0.8979. The

combined data set tree was most congruent to the tree resulting from the ndhF data

set. Fagraea section Fagraea (Fagraea clade + F) crenulata) was monophyletic and

strongly supported (BS = 96%; PP = 1.00) with F) crenulata as the basal taxon for

the clade. Fagraea crenulata is then sister to two well-supported groups, one smaller

comprising of four taxa and another bigger with eleven taxa. These groups were also

observed in the ndhF tree but the internal resolutions were slightly better in this tree.

The 4-taxon group was well resolved and received good branch support. The 11-taxon

group was divided into two subclades with one unresolved taxon (F° ridleyi). The F-

resinosa, F: imperialis and F: auriculata clade was well resolved with good branch

support. Fagraea resinosa was sister to the latter two and this clade was also observed

in all the individual data set analyses. The interspecies relationships among the taxa

in the remaining group were not resolved except for two taxa, F splendens and F. sp. B,

Molecular phylogeny of the Fagraea complex 317

ae F. auriculata

100 1.00 F. imperialis

F. resinosa

F sp.B

1.00 F. splendens

—_________—- F. carmosa

ee ae

1.00 | F. renae

Fagraea

F.sp.1 clade

F. ridleyi

100 | F. oblonga

1.00

HH F stonei

a F. curtisii

99 96 1.00 F. crassifolia

98 SEY

F. gardenioides

75 ~~ F crenulata

a F. belukar

58 «(9.93 F elliptica B Elliptica

0.93 clade

_______________ F_ eliptica PM

F. collina

Fsp.G¢

F. racemosa

400 F. sp. 2 Racemosa

_ 1.00 + F. spicata PEE

63 | —————— F. cuspidata

F. gigantea

F. caudata Gigantea

clade

1.00

F. wallichiana

F. fragrans

A. grandiflora

Fig. 3. Single most parsimonious tree based on the ndhF sequence data. The numbers above and

below the branches denote Bootstrap and Bayesian Posterior Probability values, respectively.

Length (L) = 242; consistency index (CI) = 0.8347; retention index (RI) = 0.9109. A. =

Anthocleista; F. = Fagraea. Different accessions of F. elliptica are indicated (PM = Peninsular

Malaysia, B = Borneo).

318 Gard. Bull. Singapore 64(2) 2012

that formed a well-supported subclade. This subclade was also found in the all the

individual data set analyses.

The section Cyrtophyllum was paraphyletic and resolved into two distinct

clades (Elliptica and Gigantea) as also observed in all individual data set analyses.

The Elliptica clade was sister to the Fagraea clade with good support (BS = 82%; PP

= (0.95). The Elliptica clade was itself strongly supported (BS = 100%; PP = 1.00),

wherein F; collina was sister to a clade with weak support (BS = 59%; PP = 0.69) that

contained the remaining three taxa.

The Gigantea and Racemosa clades were sisters to each other with good branch

support (BS = 99%; PP = 1.00). The Gigantea clade was internally well resolved, with

F- fragrans basal to the remaining three taxa. This relationship received moderate BS

support (65%) but good PP support (0.96). The Racemosa clade was well supported

(BS = 100%; PP = 1.00), with F: sp 2 sister to a weakly formed clade (BS = 58%).

Within this clade, F) cuspidata and F- spicata formed a moderately supported subclade

(BS = 64%; PP = 0.94).

Analyses of the expanded ITS data set

Parsimony analyses of the expanded ITS data set produced a total of 2145 most

parsimonious trees having tree length of 582 with a CI of 0.7337 and RI of 0.8089

(strict consensus shown in Fig. 5). As seen in the individual and combined data set

results, Fagraea s.|. was divided into four major clades. Fagraea crenulata was placed

in a tetrachotomy with the Fagraea, Racemosa and Gigantea clades. A similar pattern

was also seen in the 30-taxon ITS tree (Fig. 1).

Within the Fagraea clade, F. berteroana and F: salticola form a well-supported

subclade (BS = 92%; PP = 0.99). These two taxa are sister to a subclade with two

other monophyletic groups, where one is larger with 13 taxa and the other is a smaller

4-taxon clade. The 13-taxon clade is moderately supported with BS = 67% but well

supported in the Bayesian analyses (PP = 0.98). Within this 13-taxon clade there are

two smaller subclades (F) splendens and F: sp. B; F-. crassifolia and F- curtisii) with

good BS and PP support, as well as several other species whose relationships are

unresolved. The 4-taxon clade is well supported with both BS and PP values. Within

this clade, F. resinosa is sister to a moderately supported (BS = 62%; PP = 0.91)

subclade.

The Racemosa clade which represents all the taxa selected from section

Racemosae, is monophyletic and received good support (BS = 100%; PP = 1.00).

Within this clade, F’ sp 2 was sister to the rest of the taxa in a weakly formed clade (BS

=71%).

As with all other individual data set analyses, section Cyrtophyllum is

paraphyletic and was split into two distinct monophyletic groups, viz., the Elliptica

and Gigantea clades. The Gigantea clade was moderately supported with BS = 78%

but well-supported in the Bayesian analyses (PP = 1.00). The clade was internally well

resolved with moderate BS and strong PP support. The widespread F: fragrans was

placed as the basal taxon in the Gigantea clade.

Molecular phylogeny of the Fagraea complex 319

F. oblonga

F. sp. 1

i) — F. splendens

a +00 1__ F. sp.B

F. carnosa

[+ FF. stone

a EINES

1.00

F. ndleyi Fagraea

’ clade

| a F. auriculata

7s _ 100 | 1.00 F. impenalis

: 1.00

F. resinosa

F. gardenioides

96 99 F oli

70071 =e 1 00 es crassifolia

O58 > |) a F. curtisii

1.00 |

F.sp.A

0.95 F. crenulata

F. belukar

Ee Oe F elliptica B a

0.69 sb ee Elliptica

100 | F. elliptica PM clade

F. collina

a EE RERTESE

58 }-—_— Fsp.C

| | ~

100 64 F. cuspidata Racemosa

99 Saas FG E

1.00

F. fragrans

0.96 F. gigantea i

i 100 | Gigantea

100 |" gj F. wallichiana clade

cs eee F. caudata

A. grandiflora

Fig. 4. Strict consensus of four equally parsimonious trees based on the combined ITS, t#mL—F

and ndhF sequence data. The numbers above and below the branches denote Bootstrap and

Bayesian Posterior Probability values, respectively. Length (L) = 591; consistency index (CI)

= 0.8274; retention index (RI) = 0.8979. 4. = Anthocleista; F. = Fagraea. Different accessions

of F. elliptica are indicated (PM = Peninsular Malaysia, B = Borneo).

320 Gard. Bull. Singapore 64(2) 2012

The position of the Elliptica clade, sister to the rest of Fagraea s.1., was well

supported (BS = 98%; PP = 0.99). The Elliptica clade itself received moderate branch

support (BS = 77%; PP = 0.81) and within this clade, F) belukar was sister to the

rest. The three accessions of F. e/liptica from Peninsular Malaysia, Borneo and New

Guinea along with F) collina were clustered together, receiving moderate supported

with BS (72% ) but good support with PP (0.99).

The other three genera represented in the ingroup, Anthocleista, Potalia and

Lisianthius, were each monophyletic. Anthocleista and Potalia both formed a group

sister to Fagraea s.l. with good support (BS = 97%; PP = 1.00), whereas Lisianthius

was sister to Anthocleista, Potalia and Fagraea s.\. in turn, also with good support (BS

= 90%; PP = 0.96).

Implications of the molecular phylogenetic analyses

Clade correspondence to named taxonomic sections

Regardless of which gene regions were employed in the study, including the

individual and combined data set analyses, as well as the different methods used (MP

and Bayesian), the results show Fagraea s.|. segregating into four reasonably well-

supported monophyletic groups. Among these, the Fagraea clade and the Racemosa

clade closely correspond to the sectional classification, 1.e., Fagraea section Fagraea

and F; section Racemosae, respectively.

The position of the bizarre F) crenu/ata is interesting. It was unresolved in

a polytomy in both the ITS and the ITS—expanded data sets. It also failed to resolve

with the Fagraea clade in the analyses of the #7nL—F data set. Only the analyses with

the ndhF and the combined data set supported the placement of F. crenulata as a basal

taxon to the Fagraea clade.

Taxa sampled as Fagraea section Racemosae were shown to be monophyletic

in all the analyses. The clade received strong BS and PP support in analyses with all

the data sets except in the #7mL-F data set where it received moderate support for BS

(67%).

Fagraea section Cyrtophyllum as defined by Leenhouts (1962) was

paraphyletic and resolved as two distinct clades, 1.e., the Elliptica and Gigantea clades,

in all the analysed data sets. The inclusion of F’ fragrans within the Gigantea clade

was shown in the analyses with the ITS, ITS-expanded and combined data sets. In the

wnL-F data set analyses, the position of F. fragrans was unclear and in the ndhF data

set analysis, F) fragrans was sister to the Gigantea + Racemosa clades.

Thus, there was large but incomplete correspondence between well-formed

monophyletic groups in the series of analyses conducted with the various molecular

data sets and the existing taxonomic “sections” of Fagraea s.l. Whereas sections

Fagraea and Racemosae appear well-defined, the section Cyrtophyllum appears to be

an artificial grouping of two natural groups, and F) crenulata appears to be somewhat

isolated within Fagraea s.1.

321

Molecular phylogeny of the Fagraea complex

F. oblonga

F. gardenioides

0.98

0.99 1.00

1.00

0.96 82

0.94

0.96

0.81

100

1.00

0.99

1.00

100

1.00

mn (9) te le lee ay ae asl pay eS at jal ant al gat Saal fel Til aah fa) cael Gel Sol aul ah Geb Se ant ant au Sut an Sat fel el an oa Sel fel Ral Ga al

ndleyi

crassifolia

curtisii

sp. 1

splendens

sp.B

camosa

stonei

sp.A

ceilanica *

renae

resinosa

macroscypha *

aunculata

impenalis

salticola *

berteroana *

sp. 2

volubilis

spicata

cuspidata

racemosa

sp. C

fragrans

gigantea

wallichiana

caudata

crenulata

belukar

collina

elliptica PM

elliptica B

elliptica NG *

amplexicaulis *

grandiflora *

elegans *

amare *

resinifera *

laxiflorus *

cuspidatus *

brevidentatus *

algida *

Fagraea

clade

Racemosa

clade

Gigantea

clade

Elliptica

clade

Anthocleista

clade

Potalia

clade

Lisianthius

clade

OUTGROUP

affine *

Fig. 5. Strict consensus of 2145 equally parsimonious trees based on the expanded ITS sequence

data. The numbers above and below the branches denote Bootstrap and Bayesian Posterior

Probability values, respectively. Length (L) = 582; consistency index (CI) = 0.7337; retention

index (RI) = 0.8089. 4. = Anthocleista; E.= Exacum; F. = Fagraea; G.=Gentiana; L. = Lisianthius;

P. = Potalia. Different accessions of F: elliptica are indicated (PM = Peninsular Malaysia, B

= Borneo, NG = New Guinea). An asterisk after a name indicates the taxon’s sequence data

obtained from GenBank (see Methods and Materials).

322 Gard. Bull. Singapore 64(2) 2012

Potentially synapomorphic morphological characters for clades

An attempt to identify unique, non-molecular characters defining monophyletic groups

(likely synapomorphies in cladistic terms) was made. This is shown in summary form

in Table 5. This attempt might also be viewed against the pre-existing notion that

Fagraea s.|. was not particularly well distinguished as a natural group with a large

defining character suite. Prevost (1978), surveying modularity in growth architecture

among tropical woody plants, remarked that Fagraea s.l. was quite polymorphic

in the sense that it included many examples of architectural models. The examples

she provided included F) crenulata, F: fragrans and F) racemosa s.\. (she cited an

Australian-New Guinea provenance for this), with the models of Fagerlind, Aubréville

and Roux, respectively. We have surveyed multiple species for the tree architectural

character and find distinct forms representing each of the identified lineages or clades

in our phylogenetic study.

From the present survey, it can be appreciated that the section Racemosae appears to

have the most number of unique character-states among the sections compared within

Fagraea s.\. It is thus highly distinct by morphological characteristics from the other

two sections and from F) crenulata. Fagraea crenulata appears to diverge from all other

Fagraea s.|. in a number of characteristics, including Fagerlind’s architectural model

with modular branch construction (as mentioned by Hallé et al. 1978 and Prevost 1978),

and the presence of stem/branch prickles and a serrulate leaf margin. Likewise, two

groups appear morphologically well distinguished within the section Cyrtophyllum—

one with Scarrone’s architectural model, with orthotropic branch complexes and

terminal flowering; the other with Aubréville’s architectural model, with branches that

extend plagiotropically by apposition and axillary flowering. Likely synapomorphies

were identifiable for the Fagraea clade, the Racemosa clade and the Gigantea clade, but

not for the Elliptica clade, which resolved most basally in the ITS analyses (Fig. | & 5).

THE FAGRAEA CLADE (Fagraea s.s., excluding F) crenulata) — The growth habits

of members in this group are erect, scrambling, climbing or scandent shrubs or smallish

trees, which are also facultatively hemi-epiphytes. All the other distinguished groups

as well as F. crenulata are free-standing trees and do not have scrambling, climbing

or hemi-epiphytic habit. Anthocleista and Potalia are free-standing trees, with a few

climbers found in the former genus (Struwe et al. 2001; Struwe & Albert 2004).

Members of Fagraea s.s. produce copious amounts of creamy-pale to yellowish latex

in the fruit pericarp (visible especially when fresh fruits are cut or bruised). The other

groups as well as F. crenulata either have small amounts of translucent gummy latex or

(like the Racemosae clade, Anthocleista and Potalia) do not produce latex at all. The

seed shape in Fagraea s.s. is ellipsoid-rounded (similar to the condition in Anthocleistus

and Potalia: Struwe et al. 2001; Struwe & Albert 2004), compared to polygonal in all the

other three groups of Fagraea s.1. as well as F. crenulata.

FAGRAEA CRENULATA — The Fagerlind’s tree architectural model is found only in F.

crenulata within Fagraea s.1. The model applies to trees that have a monopodial trunk

Molecular phylogeny of the Fagraea complex 323

with episodic growth as well as branching tiers. The branch modules are terminated

by an inflorescence. The other groups have Scarrone’s, Aubréville’s or Roux’s models.

Perhaps the character that makes this species bizarre among Fagraea s.1. is the presence

of prickles on the stems and branches. Prickles are completely absent in all other species

within Fagraea s.1. and the Potaliinae. Another aberrant character which unmistakably

distinguishes this species is the serrulate leaf margin. All other species in Fagraea s.1.,

Anthocleista and Potalia (Struwe et al. 2001; Struwe & Albert 2004) have entire leaf

margins. In terms of morphology, F: crenulata is arguably the most enigmatic species in

Fagraea s.1.

Notwithstanding, Fagraea s.s. and F. crenulata do share a number of characters

(Table 5), although most of these are not exclusive to them and can be found in other

groups or genera. A possible link is that the petiolar sheaths of a leaf pair in both Fagraea

s.s. and F crenulata do not fully fuse. The slight fusion at the extreme edges of the

pairing sheaths do not form a consistent cup-like structure (ochrea) around the stem as

in Anthocleista and Potalia (Struwe et al. 2001; Struwe & Albert 2004; Struwe, pers.

comm.) and other taxa of Fagraea s.1.

THE RACEMOSA CLADE — Roux’s architectural model applies to all members of

the Racemosa clade. Species with this growth model have a monopodial orthotropic

stem / trunk with continuous growth. In comparison, all other taxa in Fagraea s.1.

have episodic stem / trunk growth. The branches on the stem/trunk in the Racemosa

clade are opposite and decussate but leaf arrangement on the branches is secondarily

distichous; in all the other groups of Fagraea s.1., the leaves are opposite-decussate

in arrangement. The branches in the Racemosa clade are plagiotropic, ending with a

terminal inflorescence, whereas the branches in other groups are orthotropic complexes

or plagiotropic by apposition. The terminal buds of vegetative shoots in the Racemosa

clade are not conspicuously covered with any resinous substances whereas all the

others (including Anthocleista and Potalia: Struwe, pers. comm.) have creamy to

yellowish resin covering the shoot apices.

Among Racemosa clade members, the inflorescence is generally a pendulous

elongate panicle with cymose branching where the branching pairs are condensed and

distinctly shorter than the rachis. In comparison, the inflorescences in the other groups

bear only a solitary flower or are branched cymes in which the longest basal branches

are nearly as long as the rachis. The mature fruit colour in the Racemosa clade are

generally pale to dark brown, whereas in the other groups they are yellow-orange to

red-scarlet or creamy pale grey-green to white. The fruits in the Racemosa clade (and

also Anthocleista and Potalia) do not exude any conspicuous latex, whereas fruits in

all the other Fagraea s.]. groups produce a gummy latex. Also, fruits in the Racemosa

clade (and also in Anthocleista and Potalia: Struwe, pers. comm.) in both fresh and

dried specimens, have a rather intact fruitwall epidermis that does not easily come off;

on the other hand, in all the other Fagraea s.]. groups, the epidermis separates easily

as a thin, tough and translucent peel. The mature fruit wall in dried herbarium samples

is firm and retains its rounded structure (as in Anthocleista and Potalia: Struwe, pers.

comm.), whereas in other groups the fruit wall breaks down and crumples as it dries.

Gard. Bull. Singapore 64(2) 2012

324

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326 Gard. Bull. Singapore 64(2) 2012

THE GIGANTEA CLADE — The general architecture of members of the Gigantea

clade follows Aubréville’s model in which the monopodial main trunk shows episodic

growth with opposite-decussate phyllotaxis, and the branches extend plagiotropically

by apposition (this is sometimes referred to as Terminalian branching) (Hallé et al.

1978; Prevost 1978). The other Malesian groups in Fagraea s.|. follow Scarrone’s,

Fagerlind’s or Roux’s models. The position of the inflorescence in the Gigantea clade

is axillary, whereas in all other groups of Fagraea s.|. and its subtribe (including

Anthocleista and Potalia: Struwe et al. 2001; Struwe & Albert 2004), the inflorescence

is terminal.

THE ELLIPTICA CLADE — The Gigantea and Elliptica clades have several

morphological similarities that distinguish them from other taxa in Fagraea s.1. The

exsertness of the filament from the corolla tube is very prominent within these two

clades, typically more than 70% of the total length of the filament, whereas in the

others, the filament is not exserted at all or only slightly to moderately so. Also, style

exsertness in these two clades is typically more than 40% of the total length of the pistil,

whereas in the other groups it is either not or only slightly exserted (refer to Appendix

B for a detailed comparison of filament and style exsertness among the taxa). The

structure of the stigma in these two clades is knoblike or capitate and the base of the

stigma does not expand conspicuously, as in Anthocleista and Potalia (Struwe, pers.

comm.). In Fagraea s.s., F: crenulata and the Racemosa clade, the stigma structure is

peltate due to the base of the stigma expanding conspicuously into a circular plate-like

rim. Further, the fruits in the Gigantea and Elliptica clades turn yellow-orange to red-

scarlet upon maturity whereas in the other two clades as well as F. crenulata, the fruits

ripen creamy grey-green to white or dark brown. The fruits are also generally smaller

in these two clades compared to the others. Given the distinctness of the Gigantea and

Elliptica clades as expressed in the topologies resulting from the molecular analyses

(Fig. 1-5), any apparently shared character-states they have must be considered

homoplasious in nature (1.e., similarity not due to common ancestry). Furthermore,

inflorescence position is consistently terminal in the Elliptica clade, and axillary in the

Gigantea clade.

Fagraea: complex genus or several genera?

The analysis with the expanded ITS data set (Fig. 5) shows that the monophyletic

groups are comparable in distinctness to several recently reviewed or revised ingroup

genera, 1.e., Anthocleista, Potalia and Lisianthius (Struwe et al. 2002; Struwe & Albert

2004; Weaver 1972; Sytsma 1988), which are well-established genera of the same

tribe. From the systematic and taxonomic points of view, therefore, the respective

clades are best recognised as separate genera because of their phylogenetic resolution

as monophyletic groups or isolated lineages, and their equivalence to other well-

established genera of the same tribe.

A survey of possible morphological markers also suggests that Fagraea s.1.

Molecular phylogeny of the Fagraea complex 327

contains morphologically well-distinguished groups of taxa with clear-cut boundaries.

It was possible to ascribe potential morphological synapomorphies for the four

monophyletic groups in Fagraea s.|. and for F- crenulata.

In summary, Fagraea s.|. is morphologically too divergent to be considered as

a single genus and even the sectional classification available (Leenhouts 1962) appears

to be incompletely circumscribed. The proposed concepts here for recognising the main

lineages as genera are clearly applicable to Sundaland species, which represent the

major geographical core of the complex, as these were well-represented in the present

analyses. Notwithstanding an increased clarity now available for sorting out Fagraea

s.l., various remarks on remaining problems and suggestions for future work may be

made. The two species, F. berteroana and F. salticola (New Guinea and the south-

west Pacific islands) included in the expanded ITS analysis, were resolved within the

Fagraea clade (= Fagraea s.s.). These species are among several from east Malesia that

form a group morphologically distinguished from others in Fagraea s.s. by a fleshy

ring on the inside of the corolla tube, at the insertion level of the stamens (Leenhouts

1962; Struwe et al. 2002). Better taxon sampling that includes more species from other

parts of Malesia and the south-west Pacific islands in future phylogenetic analyses

may give better resolution and confidence about clade relationships for this group.

The potential inclusion of taxa from beyond the so-called boundaries of Malesia, such

as F- schlechteri Gilg & Gilg-Ben. from New Caledonia, which Prevost (1978) has

observed to have the modular growth model of Koriba (so far not encountered within

the Malesian Fagraea complex), also promises greater insight. Also, the exploration

of further gene regions in similar studies should be interesting.

ACKNOWLEDGEMENTS. This paper is based in part on the doctoral research of the senior

author (MS) at the University of Malaya, supervised by and augmented with research findings

of the second author (KMW). The University of Malaya provided financial support for the

fieldwork and phylogenetic study through Fundamental Research Grant FS 264/2007C. We

thank Dr. Chandran Somasundram for kindly allowing use of equipment and space in his

laboratory; Y.W. Low and Zulkapli Ibrahim for specially assisting with field work and material

for DNA extraction; and N. Doraisamy, R. Sujatha, D.K.P. Lee and W.L. Goh for assistance

in the lab. The Keepers and Curators of the BO, K, KEP, KLU, L, SAN and SING herbaria

are thanked for loans of, and permission to study, specimens. Much thanks are also due to

the following for their kind assistance rendered: Assoc. Prof. Normawati Haron and Dr. K.T.

Yong (KLU); Dr. J.T. Pereira, Dr. R.C. Ong, Dr. R. Nilus, M. Postar, S. Ahmad and J. Jamirus

(SAN); Joffre Hj Ali Ahmad, Muhd Ariffin (BRUN); Dr. A.S. Kamariah (University of Brunei

Darussalam); Dr. R.C.K. Chung (KEP); Dr. Benjamin van Ee and Dr. Wenheng Zhang (Harvard

University Herbaria); Dr. B.J. Conn (NSW). Research for, and preparation of, this work have

specially benefitted through advice and comments from Prof. Rofina Yasmin and Prof. A.L.

Lim (University of Malaya): Prof. Lena Struwe (Rutgers University), who also provided plant

materials and literature; Prof. Nianhe Xia (South China Institute of Botany); Dr. Koichi Kamiya

(Ehime University); Prof. Charles Davis (Harvard University Herbaria); and Prof. Christian

Puff (University of Vienna). MS wishes to thank Dr. R. Shanti for her patience and support; Dr.

M. Vijayndra, who patiently proofread most of his dissertation; and many friends, colleagues

and associates at the University of Malaya’s Rimba Ilmu Botanic Garden. We are grateful for

328 Gard. Bull. Singapore 64(2) 2012

the support of Prof. Mohd Sofian Azirun, Dean of Science, and Prof. Rosli Hashim, Head of

the Institute of Biological Sciences, University of Malaya, as well as the previous Deans, Prof.

Hj Mohamed Abdul Majid and Prof. Amru Nasrulhaq Boyce.

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Appendix A. Sequences of primers used in PCR for amplifying the ITS, mL intron, #7mL—F

spacer and ndhF regions.

PCR Primer Sequences (5’ — 3”) Approximate size Source

amplified names of amplification in

regions PCR

ITS ITS 1 TCC GTA GGT 700-750 bp White et al.

GAA CCT GCG G 1990

TEC TCE GET TAL

ail TGA TAT GC

tn intron *€- CGA AAT CGG 300-400 bp Taberlet et al.

TAG ACG CTA CG 199]

al D3 GGG GAT AGA

GGG ACT TGA AC

tnL—-F Bl es GGT TCA AGT 300400 bp Taberlet et al.

spacer CECEACLAUC CE 199]

gl aks ATT TGA ACT GGT

GAC ACG AG

ndhF GB | CTT TCATTC CAC — 900-1000 bp This study

Fwd TE CAG TIGeT

TAT AGG GTG AAT

GB 1 AGC CAA GAA

Rev GEE

GB 2 AAA GCC AAA 900-1000 bp This study

Fwd ATAIGG TLE TTA

TGGG

GB 2 AAA TAA ATA GAA

Rev GAA AAT ATA AGA

AGA AAT GCG

332 Gard. Bull. Singapore 64(2) 2012

Appendix B. Style and filament exsertness from the corolla tube in selected species of Fagraea

s.l., representing sections Cyrtophyllum, Fagraea, and Racemosae.

The three blocks of species from top to bottom in the Table, correspond to Fagraea sections

Cyrtophyllum, Fagraea and Racemosae, respectively. CA = Cyrtophyllum (Axillary

Inflorescence), CT = Cyrtophyllum (Terminal Inflorescence), F. = Fagraea, R. = Racemosae.

mid secon in -«Covollatube Stylelength scesion protision Filament on protrusion

Baca length (mm) (mm) icatim) (%) length (mm) Ga) (%)

F. fragrans - CA (46-8 — (14)18-22 ee 57-64 See ne _ 76-80

F. gigantea - CA 7-8 (12-)18-22 =(5-)12-14 42-64 13-15 12-13 87-92

F wallichiana-CA (12-)20-25 yaa 22-25(-30) 55-65 (27-)30-38 =e € 73-74

F. elliptica - CT 3-5 (6) —7-9 34 44-50 (4-)7-8 (4-)7-8 100

F. auriculata - F 60-82 70-90 8-10 11-12 45-60 15-23 33-38

F: carnosa - F 106-140 123-126 0-17 0-13 12-15 5 33441

F. crassifolia - F 26-30 28-30 0-2 0-7 — ~ -

F. crenulata - F 15-18 15—20(—23) 0-S 0-22 12-14 7-8 57-58

F. curtisti - F 35-55 c. 60 CS) c. 8.3 32-40 c. 10 C29

F, gardenioides - F 40-53 50-55 2-10 4-18 20-27 12-17 60-63

F. imperialis - F 90-160 90-115 0-5 0-4 (55-)80-90 — (5-)27-40 9-44

F, littoralis - F 25-32 32-35 3-7 9-20 23-25 12-13 Crow,

F- oblonga - F 21-36 22-30 1-6 5-20 18-22 c.8 36-44

F. renae - F 23-34 28-32 0-5 0-16 20-26 10-14 50-54

F. ridleyi - F 32-37 40-45 c. 8 18-20 25-28 c. 10 3640

F. splendens - F 25-37 40-45 8-15 20-33 20-25 7-8 32-35

F. tubulosa - F 75-93 85-88 5-10 6-12 15-18 c. 10 56-67

F sp. B-F 26-36 35-42 6-9 14-26 20-24 6-10 30-42

Fi sp.D-F 43-60 65-70 10-22 15-31 40-50 17-20 40-43

Fisp.E-F 22-30 ~ ~ - 16-20 5-6 30-31

F. maingayi- R 16—25 12-23 0 0 19-23 2-6 11-26

F- peninsularis - R c. 14 17-19 3-5 18-26 6-7 I-—2 17-28

F. racemosa - R 10-22 15-20(-25) 3-5 20 11-17(-20) 6-8 40-55

F. volubilis - R 15-25 17-20 0-2 0-10 11-13 0-3 0-23

F sp.C-R 21-26 22-25 0-1 0-4 10-15 0 0

F sp.G-R 20-25 20-28 0-3 0-11 15-16 45 27-31

19>)

ivy)

Gardens’ Bulletin Singapore 64(2): 333-370. 2012

Twenty new species of Paraboea (Gesneriaceae) from Thailand

Pramote Triboun' and David J. Middleton?

"Bangkok Herbarium, Plant Variety Protection Division.

Department of Agriculture, Chatuchak, Bangkok 10900, Thailand

pramtrib@ gmail.com (corresponding author)

"Royal Botanic Garden Edinburgh,

20A Inverleith Row, Edinburgh, EH3 SLR, Scotland. U.K.

d. middleton@rbge.ac.uk

ABSTRACT. Twenty new species of Paraboea are described from Thailand: Paraboea

arachnoidea Triboun, Paraboea axillaris Triboun, Paraboea bhumiboliana Triboun &

Chuchan, Paraboea doitungensis Triboun & D-.J.Middleton, Paraboea eburnea Triboun,

Paraboea insularis Triboun, Paraboea lavandulodora Triboun, Paraboea monticola Triboun

& D.J.Middleton, Paraboea nana Triboun & Dongkumfu, Paraboea nobilis Triboun &

D.J.Middleton, Paraboea peninsularis Triboun & D.J.Middleton, Paraboea phanomensis

Triboun & D.J.Middleton, Paraboea quercifolia Triboun, Paraboea rosea Triboun, Paraboea

sangwaniae Tziboun, Paraboea siamensis Triboun, Paraboea takensis Triboun, Paraboea

tenuicalyx Triboun, Paraboea vachareea Triboun & Sonsupab and Paraboea xylocaulis

Triboun. Full descriptions and conservation assessments are provided for all taxa.

Keywords. Gesneriaceae. [UCN conservation assessments, Paraboea, Thailand.

Introduction

Paraboea (C.B.Clarke) Ridl. is one of the larger genera in the Gesneriaceae (Weber,

2004) and has recently been revised by Xu et al. (2008) who recognised 89 species

and five varieties. Since the publication of the revision three more species have been

described (Chen et al., 2008: Kiew, 2010). In addition the genera Phy//oboea Benth. and

Trisepalum C.B.Clarke have been synonymised into Paraboea (Puglisi et al., 2011),

following the conservation of Paraboea against these other two names (Middleton et

al., 2010). This broader genus concept has resulted in about 107 currently accepted

species.

The Gesneriaceae are currently under revision for the Flora of Thailand. This

project has involved intensive field work all over Thailand and has already resulted

in a number of new discoveries (Middleton & Triboun, 2010; Triboun & Middleton,

2010). It has also yielded many collections that were not available to Burtt (1984), for

his synopsis of Trisepalum, nor to Xu et al. (2008), for their revision of Paraboea. In

the revision by Xu et al. (2008) very many taxa were shown to be very locally endemic

and consequently more recent collections of previously unexplored or underexplored

limestone areas has revealed many taxa that are new to science. All of these are described

in Paraboea but of these the following have the characters that would previously have

qualified them as belonging to Trisepalum: Paraboea axillaris, P. bhumiboliana, P.

lavandulodora, P. nana, P. nobilis, P. peninsularis, P. phanomensis, P. sangwaniae,

334 Gard. Bull. Singapore 64(2) 2012

P. siamensis, P. takensis and P. xylocaulis. These make up a large proportion of the

newly described species partly because Trisepalum has received considerably less

research attention than Paraboea in recent years. Limestone areas in Thailand still

require considerably more exploration. We estimate that about 60% of karst limestone

in Thailand has been unexplored and of the remaining 40% about half has had only the

minimum of collecting possible at the site.

Xu et al. (2008) noted the desirability of providing IUCN conservation

assessments for all species of Paraboea under IUCN criteria IUCN, 2001), especially

given the prevalence of Paraboea species in karst limestone habitats, a habitat that is

under pressure throughout Southeast Asia, particularly due to exploitation for cement

(see Clements et al., 2006). Since the publication of Xu et al. (2008) the more intensive

field work in limestone areas of Thailand has indeed confirmed that most of the

species thought to be rather locally endemic are indeed so. We, therefore, make some

assumptions in order to provide these provisional IUCN conservation assessments:

1. we assume that species known only from the type or very few geographically

restricted specimens really are confined to the area in which they were collected; 2. we

assume that when the plant is known only from the type collection or from very few

collections which were all collected from the same site its extent of occurrence (see

IUCN, 2001 for definitions) is < 100 km? and area of occupancy is less than 10 km’.

The Thai names adopted below have been coined by Pramote Triboun. The

flowering and fruiting periods listed below are mostly estimated by comparing them to

the phenology of similar and better known species and extrapolating from the fertile

condition of the specimens when collected.

Paraboea arachnoidea Triboun, sp. nov. (Fig. 1A)

Paraboeae roseo Triboun magnitudine ubique magna, foliis rosulatis, planta pro

parte maxima pilis arachnoideis dense obtecta et floribus densis similis, sed pilis

arachnoideis in foliis magis densis, foliis ellipticis, pedunculis longioribus, corollis

violaceis et capsulis brevioribus differt. TYPUS: P. Triboun, B. Sonsupab & P.

Yothakaew 4325, 2 September 2009, Thailand, Krabi, Mu Koh Lanta, Koh Lapu Le

(holo BK; iso E).

Lithophytic perennial herb to subshrub. Stem erect, 15—40 cm high, c. 1.3 cm in diam.,

hard, woody, cracked and rough, the upper parts covered with white arachnoid hairs, the

remains of old petioles and scars remaining in the lower parts, woody root stock stout.

Leaves 11—25, arranged in a dense rosette at the top of the stem; petiole 2-6 cm long,

c. 7 mm in diam., densely covered with white arachnoid hairs; blade subcoriaceous,

elliptic, 8.5—20 x 3—5S.5 cm, apex obtuse, base cuneate or slightly attenuate, margin

crenate, secondary veins 9-11 on each side of midrib, tertiary veins reticulate,

densely covered with short multicellular glandular hairs but these somewhat obscured

by dense white arachnoid hairs on both sides. Inflorescences 6—10, subterminal,

compound pleiochasia, loosely congested 4—S branches at each node, 4-8 orders of

branching in each branch, 1—S terminal orders mostly compound dichasia; peduncle

Twenty new Thai Paraboea 555

18-26 cm long, c. 4 mm in diam., densely covered with white arachnoid hairs; bracts

elliptic or lanceolate, 1.2—1.8 < 0.3—-0.7 cm, the upper ones strongly reduced, densely

covered with white arachnoid hairs, flowers paired; pedicels 3-4 mm, covered with

white woolly hairs. Calyx green, fused for less than 0.5 mm at base, 5-lobed; lobes

lanceolate, c. 1.2 = c. 0.6 mm, apex acute, slightly covered with white hairs. Corolla

violet-blue, with a short tube and spreading lobes; tube 1.5—2.5 mm long; 2 posterior

lobes obovate or orbicular, 3—3.8 x c. 2.5 mm, apex obtuse, inner margins overlapping

each other, 3 anterior lobes ovate, 2.5—3.5 x c. 2 mm, the 2 lateral ones asymmetrical,

the middle one smaller than the lateral ones, apex rounded. Stamens 2, inserted near

base of corolla tube; filament creamy white, c. 2 mm long, geniculate; anthers yellow,

c. 1.5 x c. 2.5 mm; staminodes strongly reduced. Ovary ovoid, c. 2 mm long; style

white or light mauve, 3-4 mm long; stigma capitate, white or light mauve. Capsules

cylindric, 0.8—1 cm long, c. 1.5 mm in diam., twisted, glabrous.

Distribution. Endemic to Thailand. So far known only from the type locality.

Ecology. Open sun or in shade of dry evergreen forest on steep limestone cliff, alt.

10-200 m.

Phenology. Flowering and fruiting in July to October.

Vernacular. Cha Rue Si Yai Mang Mum (2197 LeasayN).

Etymology. The specific epithet refers to the dense arachnoid hair covering.

Proposed IUCN conservation assessment. Endangered (EN D). This species is only

known from the type locality which is in a legally protected area. Although the exact

size of the population is uncertain we estimate that it is between 50 and 250 individuals.

Notes. Paraboea arachnoidea is most similar to another new species Paraboea rosea

Triboun in its large overall size, the leaves in a rosette, the dense covering of arachnoid

hairs on most parts and the dense flowers, but it differs in the denser covering of

arachnoid hairs on the leaves, elliptic leaves, longer peduncles (18-26 cm long in P.

arachnoidea, 10-15 cm in P. rosea), violet corollas and shorter capsules (0.8—1 cm in

P. arachnoidea, 1—1.9 cm in P. rosea).

Paraboea axillaris Triboun, sp. nov. (Fig. 1B)

Paraboeae glandulosae (B.L.Burtt) C.Puglisi bracteis globosis, calvce distincte

spectabili et floribus albis similis sed foliorum pubescentia, floribus paucioribus et

capsulis longis tenuibus cylindricis differt. TYPUS: P. Triboun 3608, 14 July 2006,

Thailand, Tak, Umphang, Palata (holo BK; iso E).

Lithophytic perennial herb. Stem erect, 20-70 cm high, rarely branched, densely

covered with brown arachnoid hairs, woody root stock short and finely striped. Leaves

336 Gard. Bull. Singapore 64(2) 2012

8—12, the lower ones immediately above the root stock congested, opposite decussate,

the upper ones on upright stems loosely opposite; petiole 1-6 cm long, the upper ones

short, sulcate near base, densely covered with brown arachnoid hairs; blade papery,

ovate, elliptic or orbicular, 5-16 =< 5—8.5 cm, apex obtuse, base cuneate, margin

crenate, covered with multicellular glandular hairs above, densely covered with

brown woolly hairs beneath, secondary veins 8—9 on each side of midrib, tertiary veins

reticulate, venation prominent on both sides. Inflorescences 3—8, axillary, compound

monochasia, the lower ones sometimes subtended by reduced leaves; peduncle 3—5.5

cm long, 1—1.5 mm in diam., glabrous or lightly covered with brown woolly hairs;

bracts shiny green, globose, c. | cm, apex rounded or obtuse, glabrous; flowers paired,

2-5; pedicels 0.3—1 cm long, unequal, glabrous. Calyx cotyliform, light green, c. 1

cm long, 5-lobed, imbricate at base, upper 3 lobes fused at base; each lobe ovate or

globose, c. 3 x c. 2 mm, apex rounded, glabrous, 2 lower lobes free, ovate, c. 2.5 x c. 2

mm, apex rounded, glabrous. Corolla white, campanulate; tube 0.8—1 cm long, slightly

pale green at base; lobes globose or widely ovate, 4—6.5 x 6.5—8.5 mm, apex rounded.

Stamens 2; filaments white, c. 3 mm long, geniculate; anthers brownish yellow, c. 2

c. 4 mm; staminodes 3, narrowly linear, c. 1.5 mm long. Ovary ovoid, c. 3 mm long,

glabrous; style white, c. 5 mm long; stigma linguiform, 5—7 mm long, grooved on the

dorsal side. Capsules cylindric, 1.8—2.2 = c. 0.25 cm, twisted, glabrous.

Distribution. Endemic to Thailand. So far known only from the type locality.

Ecology. On limestone rocks in dry evergreen or mixed bamboo deciduous forest, alt.

600—900 m, in the same habitat as Paraboea takensis and P. vachareeae.

Phenology. Flowering and fruiting in June to October.

Vernacular. Khao Tok Rue Si (Y1IMANANIS).

Etymology. The specific epithet refers to the distinct axillary inflorescences.

This species is known only from the type locality. It is in a legally protected area but

is subject to burning from adjacent agricultural practices. The extent of occurrence 1s

< 100 km’.

Notes. Paraboea axillaris 1s similar to Paraboea glandulosa (B.L.Burtt) C.Puglisi

in its globose bracts, distinctly showy calyx and white flowers, but it differs in the

pubescence on the leaves, the fewer flowers (2—5 in P. axillaris, 6-12 in P. glandulosa)

and the long slender cylindric capsules (1.8—2.2 cm in P. axillaris, 2—2.7 cm in P.

glandulosa).

Twenty new Thai Paraboea 357

Paraboea bhumiboliana Triboun & Chuchan, sp. nov. (Fig. 1C—D)

Paraboeae takensi Triboun magnitudine ubique modesta, ramis tenuibus, floris

magnitudine et calycibus gemmis juvenibusque cum pilis albis arachnoideis tectis

similis, sed ramis multis, inflorescentiis brevibus, pedunculo brevi, bracteis minoribus

et corolla pallide purpurea ad pallide malvina ad basin lobi inferiore cum macula flava

orbiculari notata recedit. TYPUS: P. Triboun, T. Chuchan, S. Pintasean, C. Intasan &

J. Sittikan 3980, 1 September 2007, Thailand, Lamphun, Li, Bhumibol dam, Mae Ping

Rapid (holo BK; iso BKF, E).

Lithophytic perennial herb to undershrub. Stem erect or pendulous, 20-60 cm high,

with 10—30(—S0) branches, 10-40 cm long, slender, 2-5 mm in diam., all parts covered

with dense white to pale brown hairs, lower part woody, straight or bent, 15—20 cm

long, 1.2—1.7 cm in diam., bark brownish, root stock cracked and corky with parallel

grooves. Leaves 5—14, opposite decussate, crowded towards the apex of each branchlet

when in flower; petiole 3—7 mm long, winged, base amplexicaul and connected to the

one opposite; blade coriaceous, elliptic, oblong, or ovate, 2.5—-6 « 1.5—2.5 cm, apex

obtuse, base cuneate or obtuse (the young ones attenuate), margin slightly crenate,

secondary veins 6-8 on each side of midrib, prominent on both sides, densely covered

with multicellular glandular hairs above, densely covered with white woolly hairs

beneath, these extending beyond margin. Inflorescences 9-20, with a solitary flower

or a short cyme of 2—5 flowers, terminal or subterminal, 1—2 flowers opening at a time,

young bud densely covered with white arachnoid hairs; peduncle very short, less than 5

mm long; bracts leaf-like, lanceolate, 9-12 x 3-4 mm wide, apex obtuse or mucronate,

appressed and close to the lateral sepals; pedicel very short. Calyx cotyliform, 6—8

mm long, 5-lobed, base slightly imbricate, upper 3 lobes fused at base; each lobe

triangular, 1—1.5 mm long, apex reflexed and hooded, acute, 2 lower lobes free, elliptic

or oblong, 3—6 x 3—3.5 mm, apex acute to acuminate, all 5 lobes green and covered

with white arachnoid hairs. Corolla campanulate, c. 2 cm across; tube 1.4—1.8 cm long,

limb bilabiate, lower lobe 3-lobed, upper lobe 2-lobed, not symmetrical, base of lobes

slightly imbricate; lobes suborbicular, 0.8—1 cm long, 1—1.3 cm wide, apex rounded,

pale purple to pale mauve with yellow circular patch at base of lower lobe. Stamens 2;

filament white, flattened, somewhat parallel, upper part wider than lower part, 5.5—7

x c. 1.5 mm, fused to corolla tube near base: anthers 1.5—2.5 x 4-5 mm, creamy white

or pale brown, mauve at juncture with filament; staminodes 3, narrowly linear, |.8—2

mm long. Ovary 3—5 mm long, pubescent; style 6-7 mm long, sparsely hairy; stigma

linguiform, 4-5 mm long, apex split into 2 lobes. Capsules ellipsoid, 1—1.5 « 0.3—0.5

cm, twisted, covered with dense arachnoid hairs, calyx persistent.

Distribution. Endemic to Thailand. So far known only from the type locality.

Ecology. On limestone cliffs and slopes above the river in dry evergreen to mixed

deciduous forests.

Gard. Bull. Singapore 64(2) 2012

Fig. 1. A. Paraboea arachnoidea Triboun. B. Paraboea axillaris Triboun. C—D. Paraboea

bhumiboliana Triboun & Chuchan. (Photos: P. Triboun)

Twenty new Thai Paraboea 339

Phenology. Flowering and fruiting in July to December.

Vernacular. Bhumibolin (niiwauns), name given by His Majesty King Bhumibol of

Thailand.

Etymology. The specific epithet refers to the collection locality in the Bhumibol Dam

area in Lamphun and Tak Provinces in Thailand. The name also honours His Majesty

King Bhumibol of Thailand who has taken a keen interest in the conservation of forests

and the relationship between people and the environment. This species is dedicated to

him on the occasion of his 84" Birthday.

Proposed IUCN conservation assessment. Least Concern. Although this species is

only known from the type locality it is on both sides of the boundary of two well

legally protected areas and there are a few subpopulations.

Notes. Paraboea bhumiboliana is similar to the new species Paraboea takensis

Triboun in its globose bracts, distinctly showy calyx and white flowers, but it differs

in the pubescence on the leaves and often fewer flowers (2—5 in P. bhumiboliana, 2-8

in P. takensis).

Paraboea doitungensis Triboun & D.J.Middleton, sp. nov. (Fig. 2, Fig. 3A—B)

Paraboeae paramartinii Z.R.Xu & B.L.Burtt in dichasio amplissimo composito

terminali vel subteminali et capsulis longis cylindricis similis, sed calycibus

spectabilibus, pedicellis brevioribus et bracteis floralibus majoribus obovatis recedit.

TYPUS: P. Triboun & B. Sonsupab 4041, 26 July 2008, Thailand, Chiang Rai, Mae

Fah Luang District, Doi Tung (holo BK; iso E).

Lithophytic perennial herb. Stem erect, 10-30 cm high, more rarely up to 70 cm,

woody root stock stout. Leaves 4-10, opposite decussate and congested at the apex

of the stem; petiole narrowly winged, 6—12 cm long; blade papery, ovate, obovate

or elliptic, 27-40 « 10.5—12.5 cm, apex acute, base narrowly attenuate onto petiole,

margin indistinctly crenate, secondary veins 13—15 on each side of midrib, tertiary

veins reticulate, venation obscure above and prominent beneath, mid green and

glabrous above, lightly covered with brownish woolly hairs underneath. Inflorescences

1-2, terminal or subterminal compound dichasia, 34(—5) orders of branching in each

compound dichasium, some congested with 34 branches; peduncle dark red or green,

8-35 cm long, 1.5—2 mm in diam., the upper order branching with axes (1.5—)10(—15)

cm long; bracts cordiform or linear, the upper ones strongly reduced; flowers paired;

pedicels 0.6—1.5 cm long. Calyx green, brown at tips, fused at base for c. 1 mm,

5-lobed; lobes narrowly cylindric, (4.5—)7—7.5 x 0.5—1 mm, apex round, glabrous.

Corolla pink, campanulate; tube 7-9 mm long, very pale pink, tinted yellow at base;

lobes orbicular or widely ovate, c. 5 x 6-7 mm, apex rounded. Stamens 2, inserted

near base of corolla tube; filament creamy white, geniculate, 7-10 mm long, c. 1.5 mm

340 Gard. Bull. Singapore 64(2) 2012

Fig. 2. Paraboea doitungensis Triboun & D.J.Middleton. A. Habit. B. Flower, lateral view. C.

flower, front view showing enantiostyly. D. Calyx opened out. E. Corolla opened out. F. Ovary

and style. G. Twisted fruit. Scale bars: A= 5 cm; B-F = 0.5 cm; G= 1.5 cm. (Drawn by Claire

Banks)

Twenty new Thai Paraboea 341

wide; anthers creamy yellow, c. 2 x c. 4 mm. Staminodes 2—3, white or tinted pink

at tip, cylindric, c. 2 mm long, medial staminode not apparent. Disc annular. Ovary

ellipsoid, 4-6 mm long, c. | mm wide, green; style enantiostylous, white or pale green,

3-4 mm long; stigma capitate with glandular hairs. Capsules narrowly cylindric, 5—6.8

cm long, c. 1.8 mm wide, twisted, glabrous.

Additional specimens examined: THAILAND. Chiang Rai: Doi Tam Tu Pu, c. 520 m, 5

October 1924, H.B.G. Garrett 204 (K-2 sheets); Mae Fa Luang District, Doi Tung, 1365 m alt.,

20°19.6'N 99°50.0’E, 23 September 2008, D.J. Middleton, P. Karaket, P. Triboun, U. Kawatkul

& R. Meeboonya 4555 (BK, BKF, E); Mae Fa Luang District, summit of Doi Tung, 1510 m

altitude, 20°19.6'N 99°50.0’E, 23 September 2008, D.J. Middleton, P. Karaket, P. Triboun, U.

Kawatkul & R. Meeboonya 4576 (BK, BKF, E).

Distribution. Endemic to Thailand. So far known only from the type locality.

Ecology. Montane forest on summit of limestone hill, alt. 1100-1450 m.

Phenology. Flowering and fruiting in July to November.

Vernacular. Cha Rue Si Doi Tung (7191#AAAS).

Etymology. The specific epithet refers to the collection locality in Doi Tung, Chiang

Rai Province in Thailand.

Proposed IUCN conservation assessment. Endangered (EN Blab(iii,v)). This species

is only known from Doi Tung and has an Extent of Occurrence of < 100 km? but is

known from two distinct populations. The area is not protected and there has been a

reduction in the quality of the habitat through large scale tourism and a reduction in

the number of mature individuals.

Notes. Xu et al. (2008) included the specimen Garrett 204 in Paraboea paramartinii

Z.R.Xu & B.L.Burtt but also suggested that further collecting could reveal that this is

an undescribed species. New collections have indeed confirmed that this is a distinct

and undescribed species. It is most similar to Paraboea paramartinii, with which it

shares the characters of a large terminal or subterminal and glabrous inflorescence,

papery leaves and long cylindrical capsules, and to P. martinii (H.Lév.) B.L.Burtt and

P. glutinosa (Hand.-Mazz.) K.Y.Pan, with which it shares the characters of a terminal

or subterminal compound dichasium and long slender cylindrical twisted capsules.

However it can be distinguished from these by the showy calyx.

Paraboea eburnea Triboun, sp. nov. (Fig. 3C)

Paraboeae variopilae Z.R.Xu & B.L.Burtt inflorescentiis, floribus et fructibus cum

pilis glandulis multicellularibus omnino obtectis similis, sed ramis inflorescentiae

342 Gard. Bull. Singapore 64(2) 2012

compositis monochasialibus, foliis et floribus majoribus differt. TYPUS: P. Triboun,

B. Sonsupab & P. Yothakaew 3835, 12 September 2007, Thailand, Ranong, Kraburi,

Thum Pra Khayang (holo BK; iso E).

Lithophytic perennial herb. Stem erect, 50-80 cm high, 7-8 mm in diam., greyish

brown, woody, 4-6(—10) branches, hanging, young parts covered with rusty brown

hairs, woody root stock stout. Leaves 10-20, loosely opposite decussate; sessile; blade

papery, oblong, obovate or oblanceolate, 18—23 x 6—7.8 cm, apex obtuse, base attenuate,

margin crenate or serrate, secondary veins 8—10 on each side of midrib, tertiary veins

reticulate, venation prominent on both sides, sparsely covered in short glandular hairs

above, densely covered with brownish woolly hairs beneath. Inflorescences 3—10 (2-4

on each branch), subterminal dichasia with 1(—2) orders of branching, each branch a

compound monochasium (rarely dichasial), whole inflorescence densely covered with

whitish multicellular glandular hairs; peduncle c. 4 cm long, c. 1.5 mm in diam.; bracts

strongly reduced or absent; flowers paired; pedicels 7—9 mm long, covered by whitish

multicellular glandular hairs. Calyx green, fused at base for less than | mm, 5-lobed;

lobes lanceolate, c. 3 x 1 mm, apex obtuse, covered by whitish multicellular glandular

hairs. Corolla white, with a short tube and spreading lobes; 2 posterior lobes ovate, c.

7 x c. 5 mm, apex obtuse, 3 anterior lobes oblong to obovate, 9-10 x 6—7 mm, apex

obtuse or truncate, the middle lobe longer than the laterals. Stamens 2, inserted near

the base of corolla tube; filament white, c. 3 mm long, geniculate; anthers yellow, c.

2.5 x 3.5-4 mm; staminodes 3, white, c. 2.5 mm. Ovary white, ellipsoid, c. 3.5 mm

long, covered with short whitish multicellular glandular hairs; style white, c. 4.5 mm

long, glabrous; stigma capitate, white. Capsules narrowly cylindric, 2.2—2.9 x c. 0.2

cm wide, twisted, covered with whitish multicellular glandular hairs.

Distribution. Endemic to Thailand. So far known only from the type locality.

Ecology. On rock in shade in evergreen forest on an isolated limestone hill, surrounded

by mangrove forest dominated by Nypa fruticans, alt. 50-100 m.

Phenology. Flowering and fruiting in late July to November.

Vernacular. Cha Rue Si Pra Kha Yang (21914 W3Z9HWA).

Etymology. The specific epithet refers to the white flowers.

Proposed IUCN conservation assessment. Critically Endangered (CR D). This

species is only known from one small population at the type locality. The population

is estimated to contain fewer than 50 individuals and the locality currently has no

protected status.

Notes: Paraboea eburnea is similar to Paraboea variopila Z.R.Xu & B.L.Burtt in the

complete covering of multicellular glandular hairs on the inflorescences, flowers and

Twenty new Thai Paraboea

Fig. 3. A-B. Paraboea doitungensis Triboun & D.J. Middleton. C. Paraboea eburnea Triboun.

D. Paraboea insularis Triboun. (Photos: P. Triboun)

344 Gard. Bull. Singapore 64(2) 2012

fruits, but it differs in the compound inflorescence branches being monochasial, the

larger leaves (18-23 cm long in P. eburnea, 5—17 cm long in P. variopila) and larger

flowers (upper lobes c.7 mm long in P. eburnea, c. 3.5 mm long in P. variopila).

Paraboea insularis Triboun, sp. nov. (Fig. 3D)

Paraboeae suffruticosae (Ridl.) B.L.Burtt in habitu et statura, foliis ellipticis usque

lanceolatis, et florum fructuumque magnitudinum similis, sed partibus inflorescentiarum

ultimis dense congestis et corollis violascenti-caeruleis differt. TY PUS: P. Triboun, B.

Sonsupab & P. Yothakaew 4587, 25 September 2010, Thailand, Phangnga, limestone

island border of Phang-gna-Krabi bay (holo BK; iso E).

Lithophytic perennial herb to subshrub. Stem erect, 7-20 cm high, solitary or with 2-4

twigs, 3-6 mm in diam., grey, deeply grooved and corky, root stock stout and rough.

Leaves 8—16, arranged in a whorl or slightly opposite decussate, densely congested

at the apex; petiole yellowish brown, 0.5—2 cm long, sulcate, densely covered with

short brown woolly hairs; blade coriaceous, elliptic to lanceolate, 5-8 « 1.42.6 cm,

apex acute, base cuneate, margin slightly crenate, glaucous, densely covered with

muiticellular glandular hairs above, densely covered with light brown woolly hairs

beneath, secondary veins 14—17 oneach side of midrib, prominent beneath, tertiary veins

reticulate, obscure on both sides. Inflorescences 1—3, subterminal, mostly compound

dichasium, rarely pleiochasial at first node, 3-4 orders of branching; peduncle green

or brown, 5—9 cm long, c. 1.5 mm in diam., densely covered with short light brown

hairs; bracts leafy, elliptic, 1-2.5 x 0.3—0.7 cm, apex obtuse, densely covered with

light brown arachnoid hairs; the first order branch axis long and slender, 3—6 cm long,

c. 0.5 mm in diam., densely covered with short light brown hairs, subtended by leafy

bracts and without a flower at the branching point, the upper orders densely congested,

less than | cm long; flowers paired; pedicels 0.3—1 cm long, densely covered with light

brown hairs. Calyx light brown, fused at base for c. | mm, 5-lobed; lobes narrowly

linear, 2—3.5 = c. 0.5 mm, apex obtuse, densely covered with light brown woolly hairs.

Corolla violet-blue, with a short tube and 2-lipped spreading limb, tube 3—S mm long,

the lower lip much longer; 2 posterior lobes orbicular, c. 3 x c. 3 mm, apex rounded,

3 anterior lobes unequal, 2 lateral ones obovate, 3—3.5 x c. 3.5 mm, the middle one

orbicular, c. 3.5 * c. 3 mm, apex rounded. Stamens 2; inserted near base of corolla

tube; filament white, c. 3 mm long, geniculate; anthers yellow, c. 2 < c. 3.5 mm;

staminodes 2, c. 1.5 mm long. Ovary creamy white, ellipsoid, c. 3 mm long, glabrous;

style white, c. 3 mm long; stigma capitate, white. Capsules ellipsoid, 6-8 x 1.5—2 mm,

twisted, glabrous.

Additional specimens examined: THAILAND. Krabi: Koh Hin Bai, 31 October 2006, P.

Triboun 3673 (BK, E).

Distribution. Endemic to Thailand. So far known only from the type locality.

Twenty new Thai Paraboea 345

Ecology. \n open sun on limestone cliffs, alt. 10-150 m.

Phenology. Flowering and fruiting in July to November.

Vernacular. Sud Sa Khon (qaqa).

Etymology. The specific epithet refers to the collection locality on limestone islands in

Phangnga-Krabi Bay in Thailand.

Proposed IUCN conservation assessment. VU D1. Although this species is only known

from the type locality this is in a legally protected area. It is difficult to estimate the

population size as the plants are on inaccessible cliffs but is likely to be between 250

and 1000 mature individuals.

Notes. Paraboea insularis 1s similar to Paraboea suffruticosa (Ridl.) B.L.Burtt in its

habit and overall size, elliptic to lanceolate leaves, flower and fruit sizes, but it differs in

the densely congested ultimate parts of the inflorescences and the violet-blue corollas.

Paraboea lavandulodora Triboun, sp. nov. (Fig. 4A)

Paraboeae glabrescenti (Barnett) C.Puglisi floribus binatis in monochasiis compositis

densis aggregatis, bracteis globosis et corollis dilute caeruleis similis, sed foliis

sessilibus, inflorescentiis brevibus et partibus plurimis cum trichomatibus glandulis

viscidis tectis differt. TYPUS: P. Triboun 4433, vouchered from material flowering at

BK on 11 April 2010, originally collected in the wild, Thailand, Bung Kan, Phu Tok

(holo BK).

Lithophytic perennial herb. Stem erect, 16-45 cm high, 24 mm in diam., brown.

Leaves 8-15, shortly opposite decussate; sessile; blade coriaceous, elliptic, lanceolate

or obovate, 5—7 « 2-4 cm, apex acute, base broadly attenuate, margin serrate, covered

with short multicellular glandular hairs on both sides, sticky, also covered with light

brown arachnoid hairs, secondary veins 6—8(—10), prominent on both sides, tertiary

veins reticulate. Inflorescences 2—3, subterminal or axillary; compound monochasia,

rather dense; peduncle c. 9 cm long, 2—2.3 mm in diam., covered with multicellular

glandular hairs, bracts green, globose, 0.8—1.1 = c. 1 cm, apex rounded or shallowly

lobed, base slightly plicate, covered with multicellular glandular hairs; paired flowers

3—5; pedicels c. 7 mm long, unequal, green, covered with multicellular hairs. Calyx

cotyliform, 5-lobed, upper 3 lobes fused at base; each lobe linear to elliptic, 7—7.5

x c. 3.5 mm, apex obtuse, 2 lower lobes fused together at the base, free from the

upper ones, lobes lanceolate to elliptic, c. 8.5 < c. 3.5 mm, apex obtuse, covered by

short glandular hairs. Corolla light blue, campanulate; tube 8—9 mm long; lobes widely

ovate, 5.5—6 x 9-9.5 mm, apex rounded. Stamens 2, inserted near base of corolla tube;

filament white, c. 4.5 x c. | mm, flattened, geniculate; anthers creamy yellow, c. 2 x

346 Gard. Bull. Singapore 64(2) 2012

c. 4.5 mm; staminodes 3, white, linear, c. 1.5 mm long. Disc annular, c. 0.3 mm high.

Ovary light green, ellipsoid, c. 3 x c. 1.8 mm, glabrous; style 5—5.5 mm long; stigma

white, linguiform, c. 3 x c. | mm, apex bilobed. Capsule not seen.

Distribution. Endemic to Thailand. So far known only from the type locality.

Ecology. On sandstone cliffs in shade of mixed deciduous forest.

Phenology. Flowering in May to July (data only from cultivation).

Vernacular. Cha Rue Si Bai Hom (mis LuUVAN).

Etymology. The specific epithet refers to the mild lavender-like scent, particularly

noticeable in the leaves of fresh plants.

Proposed IUCN conservation assessment. Near Threatened. This species is only known

from the type locality which is in the grounds of a temple. This means that it has no

formal protected status but is unlikely to be under any immediate threat. However, we

are unsure of the population size and the total distribution of the species 1s restricted.

It requires regular monitoring of its status to see if it requires a higher threat status.

Notes. Paraboea lavandulodora is similar to Paraboea glabrescens (Barnett) C.Puglisi

in its paired flowers in dense compound monochasia, globose bracts and the light blue

corollas, but it differs in the sessile leaves, short inflorescences (peduncle c. 9 cm long

in P. lavandulodora, \0—15 cm in P. glabrescens) and having most parts covered with

sticky glandular trichomes.

Paraboea monticola Triboun & D.J. Middleton, sp. nov. (Fig. 4B—C)

Paraboeae regulari (Ridl.) Ridl. magnitudine ubique parva et fructibus rectis similis,

sed foliis tenuibus chartaceisque, petiolis multo longioribus et tenuibus, floribus

fructibusque multis differt. TYPUS: P. Triboun 3662, 25 September 2006, Thailand,

Phangnga, Tai Toy (holo BK; iso E).

Lithophytic perennial herb or subshrub. Stem erect, 10-35 cm high, solitary or in

clumps of 2—6 shoots, c. 4 mm in diam., light brown or grey, woody root stock slender

and striped. Leaves 6—15, arranged densely in whorls or alternate and congested at the

apex; petiole slender, (1.5—)4(—7) cm long, the ones lower on the plant longer, sulcate,

densely covered with short brown hairs; blade papery, obovate, oblanceolate, ovate or

elliptic, 4-10.5 x 2.5—5.5 cm, apex acute, base oblique, cuneate or rounded, margin

serrate or cuneate, glabrous above, densely covered with short hairs and furfuraceous

beneath; secondary veins 7-10 on each side of midrib, tertiary veins finely reticulate,

prominent on both sides. Inflorescences 5—15, axillary or subterminal, compound

Twenty new Thai Paraboea 347

dichasia, 24 orders of branching; peduncle thin and slender, 1-4 cm long, c. 0.5

mm in diam., furfuraceous; bracts reduced, linear, 1—3.5 = c. 0.3 mm, the upper ones

strongly reduced, apex acute; flowers paired, pedicels thin and slender, 0.5—1.5 cm,

furfuraceous. Calyx greenish brown; fused at base for c. 0.5 mm long, 5-lobed; lobes

linear, c. 2 mm long, apex obtuse, furfuraceous. Corolla white, with a short tube and

spreading lobes; tube c. 1.5 mm long; lobes ovate to obovate, 6—9 x c. 6.5 mm, apex

rounded. Stamens 2, inserted near base of corolla tube; filament white, c. 2.5 mm,

geniculate; anthers yellow, c. 2 x 2 mm; staminodes strongly reduced, c. 1.5 mm,

white. Ovary light green, ellipsoid, c. 1.3 x 0.7 mm; style white, c. 2.5 mm; stigma

capitate, white with green at tip. Capsules cylindric, 0.8-1 x c. 1.5 mm, straight,

glabrous or furfuraceous.

Additional specimens examined: THAILAND. Surat Thani: Phanom District, Khlong

Phanom National Park, trail from Park headquarters, 200 m altitude, 8°52'4'N 98°40'6"E,

7 September 2008, D.J. Middleton, P. Triboun, V. Chamchumroon, S. Saengrit & R. Simma

4363 (E): Phanom District, Khao Sok National Park, 100-200 m altitude, 12 December 1979,

T. Shimizu, H. Toyokuni, H. Koyama, T. Yahara & C. Nivomdham T-27069 (L). Phangnga:

Muang Phangnga, 25 September 2006, P. Triboun 3662 (E); Pulau Panji, 2 December 1928,

Haniff & Nur 4013 (K).

Distribution. Endemic to Thailand. Known only from Phangnga and Surat Thani

Provinces.

Ecology. On shaded limestone rocks in evergreen forest, alt. 20-200 m.

Phenology. Flowering and fruiting in late June to October.

Vernacular. Cha Pho Ta Rue Si (wawaargré).

Etymology. The specific epithet refers to its collection localities in low but rugged hilly

areas in Phangnga and Surat Thani Provinces in Thailand.

Proposed IUCN conservation assessment. Vulnerable (VU Blab(ili,v)). This species

is relatively widespread over two provinces and known from several populations,

some of which are in legally protected areas and others which are not. The Extent of

Occurence is < 20,000 km? and those populations not in protected areas are in decline

due to habitat disturbance.

Notes. Haniff & Nur 4013 (K) was included in Paraboea regularis (Ridl.) Ridl. by

Xu et al. (2008). Although Paraboea monticola is similar to Paraboea regularis in its

small stature and straight fruits it differs in having thin papery leaves, much longer and

slender petioles and many flowers and fruits. With the removal of Haniff & Nur 4013

from Paraboea regularis this species is now not known from Thailand.

348 Gard. Bull. Singapore 64(2) 2012

Paraboea nana Triboun & Dongkumfu, sp. nov. (Fig. 4D)

Paraboeae birmanicae (Craib) C. Puglisi magnitudine ubique parva, petiolis oppositis

conjungentibus trans nodum auriculam formantibusque et floribus parvis similis, sed

floribus in inflorescentia paucioribus, pedicellis robustis et corollis dilute caeruleis

recedit. TY PUS: P. Triboun & W. Dongkumfu 4035, 18 July 2008, Thailand, Lamphun,

Li District, Mae Ping National Park, Kor Noi Falls (holo BK; iso BKF, E).

Lithophytic perennial herb or subshrub. Stem erect, 10-25 cm high, few branches

with many shoots, glaucous, woody root stock slender. Leaves 8-14, loosely opposite

decussate; petiole brown, 1—1.3 cm long, forming an auricle with the opposite petiole

at the node, sparsely covered with glandular hairs; blade coriaceous, ovate or elliptic,

3.2-5.3 x 1.42.4 cm, apex acute or obtuse, base cuneate, margin slightly crenate,

densely covered with multicellular glandular hairs above, densely covered with brown

wooly hairs beneath, secondary veins 3—5 on each side of midrib, prominent beneath.

Inflorescence 1—2 flowers only, monochasia, subterminal and axillary; bract linear, c.

lcm = 1.5—2 mm, light brown covered with white arachnoid hairs; pedicel robust,

2.2—3 cm long (up to 4 cm when fruiting), unequal, light brown covered with white

arachnoid hairs. Calyx cotyliform, 5—6 mm long, 5-lobed, slightly imbricate at base,

upper 3 lobes fused at base; each lobe ovate to obovate, c. 5 x 5—5.5 mm, apex obtuse,

2 lower lobes free, ovate to oblong, c. 4 x c. 5 mm, apex obtuse, covered with light

brown arachnoid hairs. Corolla light blue, campanulate; tube light blue to nearly white,

8—9 mm long; lobes broadly ovate, c. 5 x 4.5—7 mm, apex rounded or obtuse. Stamens

2, filament white, c. 3 mm long, geniculate; anthers creamy yellow, c. 3 x c. 2 mm;

staminodes 2, 0.5—0.6 mm long. Ovary subglobose, 1.5—2 mm long; style white, c. 3.5

mm long, covered with glandular hairs; stigma linguiform, c. 1.5 mm, white. Capsules

ovoid, 1—1.4 cm long, twisted, covered with white arachnoid hairs, surrounded by

persistent calyx.

Distribution. Endemic to Thailand. So far known only from the type locality.

Ecology. On limestone cliffs in dry evergreen to mixed dipterocarp forests, alt. 600—

900 m.

Phenology. Flowering and fruiting in late June to October.

Vernacular. Cha Rue Si Noi (2191swAL).

Etymology. The specific epithet refers to the very small stature of the plant and the

particularly small flowers amongst those plants previously referred to Trisepalum.

Proposed IUCN conservation assessment. Least Concern. Although this species is

only known from the type locality it is on both sides of the boundary of two legally

protected areas and there are a few subpopulations.

Twenty new Thai Paraboea

Fig. 4. A. Paraboea lavandulodora Triboun. B—C. Paraboea monticola Triboun & D.J.

Middleton. D. Paraboea nana Triboun & Dongkumfu. (Photos: P. Triboun)

350 Gard. Bull. Singapore 64(2) 2012

Notes. Paraboea nana is similar to Paraboea birmanica (Craib) C.Puglisi 1n its small

overall size, the opposite petioles joining and forming an auricle across the node and

the small flowers, but it differs in the fewer flowers in an inflorescence (1—2 in P. nana,

4-10 in P. birmanica), the robust pedicels and the light blue corollas.

Paraboea nobilis Triboun & D.J. Middleton, sp. nov. (Fig. SA)

Paraboeae phanomensi Triboun & D.J.Middleton pedunculo pedicelloque tenui,

foliis lanceolatis et magnitudine ubique similis sed pedicellis longioribus, floribus

paucioribus et majoribus, lobis corollae atriore caeruleis recedit. TYPUS: D.J.

Middleton, P. Triboun, V. Chamchumroon, S. Saengrit & R. Simma 4312, 6 September

2008, Thailand, Surat Thani, Ban Thakhun subdistrict, Khao Sok National Park (holo

BK; iso BKF, E, K, KEP).

Lithophytic perennial herb or small shrub. Stem erect, 25—50 cm high, branches many,

c..20 cm long, c. 6.5 mm in diam., woody root stock robust and rough. Leaves many,

10-15 on each shoot, arranged rather densely opposite decussate; petiole 1.5—2 cm

long, sulcate, covered with light brown wooly hairs; blade coriaceous, lanceolate or

oblong, 6—9 x 1.4—2.6 cm, apex obtuse or acute, base cuneate, margin indistinctly

crenate, loosely covered with white arachnoid hairs above, glaucous and densely

covered with light brown wooly hairs beneath, secondary veins 10—13 on each side

of midrib, tertiary venation not visible, prominent on both sides. Inflorescences 4—5,

subterminal or axillary, mostly a single flower or rarely | pair of monochasia; bracts

brown, coriaceous, globose, 1.3—1.5 cm, hooded, lightly rugose, apex rounded or

acute, a protruding bulge present around the middle to base on the outside, outside

light brown and densely covered with wooly hairs, inside green and glabrous; pedicel

and peduncle 3—5.5 cm long, c. 1.3 mm in diam., densely covered with woolly hairs.

Calyx cotyliform, tube 5—6 mm long, 3-lobed; lobes obovate, dorsal lobe c. 1.2 x c.

0.9 cm, lateral lobes c. | =< c. 0.6 cm, apex acute, margin fimbriate, densely covered

with light brown woolly hairs. Corolla campanulate; tube white, c. 1.5 cm long; lobes

violet-blue, widely globose or widely ovate, 1.2—1.5 x 1.3—1.5 cm, apex rounded,

margin plicate and fringed. Stamens 2; filaments white, c. 5 mm long, geniculate;

anthers yellowish brown, c. 4 < c. 6 mm; staminodes 3, linear, 2—2.5 mm long. Ovary

ovoid, c. 5 x c. 3 mm, covered with white arachnoid hairs; style white, c. 7 mm long;

stigma linguiform, c. 8 mm long,white. Capsules woody, ellipsoid, c. 2.7 x c. 0.7

cm, twisted, glaucous and loosely covered with white arachnoid hairs, surrounded by

persistent calyx.

Distribution. Endemic to Thailand. So far known only from the type locality.

Ecology. On limestone rocks in evergreen forest, alt. 100-300 m.

Phenology. Flowering and fruiting in May to October.

Twenty new Thai Paraboea 351

Vernacular. Sri Ra Cha Pra Pha (@39%9¢/1).

Etymology. The specific epithet refers to its large and spectacular flowers, coupled

with an attractive leaf shape, all in pleasant proportion to the size of the plant.

Proposed IUCN conservation assessment. Endangered (EN D). This species is only

known from the type locality which is in a legally protected area. Although, the exact

size of the population is uncertain we estimate that it is between 50 and 250 individuals.

Notes. Paraboea nobilis is most similar to another new species, Paraboea phanomensis

Triboun & D.J.Middleton, in its slender peduncle and pedicel, lanceolate leaves and

overall size, but it differs in longer pedicels (3—5S.5 cm in P. nobilis, 2-6 mm in P.

phanomensis), fewer (1—2 in P. nobilis, 2-5 in P. phanomensis) and larger flowers

(corolla tube 1.5 cm in P. nobilis, \—1.3 cm in P. phanomensis) and darker blue corolla

lobes.

Paraboea peninsularis Triboun & D.J. Middleton, sp. nov. (Fig. 5B—C)

Paraboaeae barnettiae C. Puglisi in habitu et statura, foliis lanceolatis, inflorescentiis

longis et capsulis tortis cylindricis similis sed foliorum apicibus acutis, bractearum

apicibus rotundis et corollis albis differt. TYPUS: P. Triboun & P. Yothakaew 4300, 31

July 2009, Thailand, Krabi, Hat Nop Parat Thara (holo BK; iso E).

Lithophytic perennial herb to small shrub. Stem erect, 20-65 cm high, 10-20

branches, 10—20 cm long, c. 2 mm in diam., woody root stock thick and rough. Leaves

many (10—20 on each shoot), loosely opposite decussate; petiole 0.6—1.5 cm long,

sulcate, densely covered with short brown hairs; blade coriaceous, lanceolate, 6.5—10

x 2.3-3 cm, apex acute, base cuneate, margin indistinctly crenate, secondary veins

7-10 on each side of midrib, prominent beneath, above shiny green and glabrous,

densely covered with brownish woolly hairs beneath. Inflorescences 1—2, subterminal,

compound monochasia, c. 6 cm long; peduncle 5—7 cm long, c. 2 mm in diam., square

in cross-section, densely covered with light brown woolly hairs; bracts globose,

1.4-1.7 cm, apex rounded, hooded, outside densely covered with light brown woolly

hairs, inside green and glabrous; flowers paired c. 6; pedicels 2-8 mm long, unequal,

densely covered with light brown woolly hairs. Calyx cotyliform c. 1 cm long, 3-lobed,

almostly free; lobes globose, c. 1.3 cm, apex rounded or obtuse and fimbriate, densely

covered with light brown woolly hairs. Corolla white, campanulate; tube c. 1 cm long;

lobes orbicular, c. 0.6 x 0.9—1.1 cm, apex rounded. Stamens 2; filament creamy white,

4-5 mm long, geniculate; anthers yellow, c. 2 x c. 4 mm; staminodes 2, linear, c. 2.5

mm long. Ovary ovoid, c. 4.5 * c. 2 mm, densely covered with woolly hairs; style

white, c. 6.5 mm long; stigma linguiform, 3.5—4 mm long, white. Capsules cylindric,

c. 2 x c. 0.5 cm, twisted, densely covered with light brown woolly hairs.

352 Gard. Bull. Singapore 64(2) 2012

Additional specimens examined: THAILAND. Krabi: Muang Krabi District, Ko Phi Phi

National Park, small limestone island on Nop Parat Thara Beach, alt. 5 m, D./. Middleton, P.

Triboun, V. Chamchumroon, S. Saengrit & R. Simma 4449 (BK, BKF, E).

Distribution. Endemic to Thailand. So far known only from the type locality.

Ecology. On rocks in dry evergreen forest on limestone island near shore, alt. 20-100 m.

Phenology. Flowering and fruiting in June to October.

Vernacular. Sin Sa Mut (@waayns).

Etymology. The specific epithet refers to Peninsular Thailand in which Krabi Province

is to be found.

Proposed IUCN conservation assessment. Endangered (EN D). This species is only

known from the type locality which 1s in a legally protected area. Although, the exact

size of the population is uncertain we estimate that it is between 50 and 250 individuals.

Notes. Paraboea peninsularis 1s similar to Paraboea barnettiae C.Puglisi in its habit

and overall size, lanceolate leaves, long inflorescences and twisted cylindric capsules,

but it differs in having acute leaf apices, rounded bract apices and white corollas.

This species was referred to as Trisepalum sp. nov. 2 in Puglisi et al. (2011).

Paraboea phanomensis Triboun & D.J. Middleton, sp. nov. (Fig. 5D, Fig. 6)

Paraboeae albidae (Barnett) C.Puglisi magnitudine ubique magna, bracteis globosis

et foliorum forma similis, sed foliorum pagina superiore glabra, bracteis cum apice

acuto, inflorescentiis cum axibus ultimis dense fasciculatis et capsulis minoribus

differt. TYPUS: D.J. Middleton, P. Triboun, V. Chamchumroon, S. Saengrit & R.

Simma 4365, 7 September 2008, Thailand, Phanom District, Khlong Phanom National

Park, Trail from Park headquarters (holo BK; iso BKF, E, K, KEP, L).

Lithophytic pernennial herb to small shrub. Stem erect, 25—60 cm high, branches

many, 20-40 cm long, 3—3.5 mm in diam., woody root stock robust and finely striate.

Leaves many, 10—16 on each shoot, arranged densely opposite decussate; petiole

1.42 cm long, sulcate, covered with light brown woolly hairs; blade coriaceous,

lanceolate, 4.5—-8(—12) <x 2—2.8(—3.7) cm, apex acute or subacute, base cuneate,

margin crenate, glabrous or loosely covered with white arachnoid hairs above,

glaucous and densely covered with light brown woolly hairs beneath, secondary

veins 10-13 on each side of midrib, prominent beneath. Inflorescences 4-8,

subterminal or axillary, compound monochasia, axis 1.5—3 cm long, 1—2 flowers

at each node; peduncle slender, 3—6 cm long, 0.8—1 mm in diam., glaucous, hairs

Twenty new Thai Paraboea

Fig. 5. A. Paraboea nobilis Triboun & D.J. Middleton. B—C. Paraboea peninsularis Triboun

& D.J. Middleton. D. Paraboea phanomensis Triboun & D.J. Middleton. (Photos: P. Triboun)

354 Gard. Bull. Singapore 64(2) 2012

sparse; bracts globose, 0.8—1 = c. 0.8 cm, hooded, apex sharply acute, outside light

brown and densely covered with white woolly hairs, inside green and glabrous;

pedicels 2-6 mm, unequal, densely covered with white or light brown woolly hairs.

Calyx cotyliform, 3-lobed, almostly free; posterior lobe oblong to elliptic, 6—8.5 x

34.5 mm, apex sharply acute, margin fimbriate, 2 interior lobes linear to lanceolate,

c. 7 x c. 2 mm, apex acute, margin fimbriate. Corolla campanulate; tube white or

pale pink, (7—)1—1.3 cm long; lobes light blue, pink or pale blue, broadly globose or

broadly ovate, (0.6—)0.8—0.9 « 0.9-1.3 cm, apex rounded, margin fringed. Stamens

2; filaments creamy white, c. 5.5 x c. 2 mm, geniculate, flattened; anthers c. 2.5 x c.

4 mm, yellow; staminodes 2-3, linear, c. | mm long, white, medial staminode not

apparent. Ovary ovoid, c. 5 mm, densely covered with brown woolly hairs; style

white, 6-8 mm long; stigma linguiform, 4-6 mm long, white. Capsules woody,

cylindric to ellipsoid, 2—2.7 x 0.3—0.45 cm, twisted, glaucous and densely covered

with brown woolly hairs, surrounded by persistent calyx.

Distribution. Endemic to Thailand. So far known only from the type locality.

Ecology. On limestone rocks at edge of evergreen forest, alt. 100-300 m.

Phenology. Flowering and fruiting in May to October.

Vernacular. Sri Su Rat (qa1Q9).

Etymology. The specific epithet refers to the collection locality in Khlong Phanom

National Park, Surat Thani province in Thailand.

Proposed IUCN conservation assessment. Least Concern. This species is known

from several populations over a wide area. Some of these populations are not in

protected areas and monitoring is necessary to assess whether this species may

qualify for a threat category in the future.

Notes. Paraboea phanomensis 1s similar to Paraboea albida (Barnett) C.Puglisi in

its large overall size, globose bracts and leaf shape, but it differs in the glabrous

upper leaf surface, the acute apex to the bracts, inflorescences with densely clustered

ultimate axes and smaller capsules (2—2.7 cm long in P. phanomensis, 2.3-3 cm long

in P. albida).

Several new species fromarange of families have been described from Khlong Phanom

National Park in recent years (e.g. Lindsay & Middleton, 2004; Phuphathanapong,

2006; Lindsay et al., 2008; Inthachub et al., 2009) even though this is a relatively

small National Park. It is fairly accessible and many of these species may eventually

also be found in the more difficult terrain of the neighbouring Khao Sok National

Park.

Twenty new Thai Paraboea 355

Fig. 6. Paraboea phanomensis Triboun & D.J.Middleton. A. Habit. B. Flower, lateral view. C.

Flower, lateral view cross section. D. Corolla opening. E. Calyx opened out. F. Corolla opened

out. G. Ovary and style front and lateral views. H. Twisted fruit. Scale bars: A= 5 cm; B—G =

0.5 em; H = 1.5 cm. (Drawn by Claire Banks)

356 Gard. Bull. Singapore 64(2) 2012

Paraboea quercifolia Triboun, sp. nov. (Fig. 7A)

Ab aliis speciebus generis combinatione characterum sequentium: caulis brevis

tenuisque lignosus ad apicem cum foliis congestis, inflorescentiae pilis glandulis

minutis viscidis dense obtectae, calyx 5-partitus cum lobis anguste ovatis et corollae

violaceo-caeruleae. TYPUS: P. Triboun & P. Yothakaew 4296, 18 June 2009, Thailand,

Lopburi, Sri Samrong-Chibadan, Khao Tumbol, Wat Tham Wiweg Sri Sawat (holo

BK; iso E).

Lithophytic perennial herb, solitary or in clumps, stem 5—10 cm high with 2—3 shoots,

woody rootstock cracked and rough. Leaves 10-20, arranged in a dense rosette at the

top of the stem; sessile; blade lanceolate, elliptic or oblong, 3—6.5 cm = 2.5—3.5 cm,

the lower ones larger than the upper ones, apex acute or obtuse, margin crenate but

entire near the base, green above, light brown beneath, secondary veins 4—5 on each

side of midrib, obscure above, prominent beneath, both sides with dense glandular

hairs and a sparse covering of arachnoid hairs. Inflorescences 1—4, subterminal, 2—4

times branched in each compound dichasium; peduncle red, 6-15 cm long, 2—3 mm in

diam., covered with dense red multicellular glandular hairs, tips of hairs with yellow

sticky substance; bracts linear, the upper ones strongly reduced; bracteoles reduced or

absence; flowers paired; pedicels up to 3 cm long, those of a pair unequal in length.

Calyx greenish brown, c. 5 mm long, fused at base for c. 1 mm, 5-lobed; lobes narrowly

ovate, covered with dense tiny glandular hairs. Corolla violet-blue, with a short tube

and spreading lobes; tube c. 4.5 mm long; 2 posterior lobes ovate or obovate, c. 5 x

c. 5.5 mm, apex rounded; 3 anterior lobes orbicular to ovate, c. 7 x c. 6 mm, apex

rounded, the middle one hooded at apex. Stamens 2, inserted near base of corolla tube;

filament mauve, geniculate, c. 3 mm long, c. 0.8 mm wide; anthers yellow, c. 3 x c. 3

mm. Staminodes 2, white, + triangular, 1-2 mm long, medial staminode not apparent.

Disc annular. Ovary ovate, c. 2 mm long, green, covered with tiny glandular hairs.

Style mauve, 3-4 mm long. Stigma capitate, yellowish brown. Capsules narrowly

cylindric, 0.8—1 x c. 0.1 cm, twisted.

Distribution. Endemic to Thailand. So far known only from the type locality.

Ecology. Mixed deciduous forest on summit of limestone hill, alt. 100-300 m.

Phenology. Flowering and fruiting in late May to August.

Vernacular. Cha Rue Si Bai Ko (w1Miisluna).

Etymology. The specific epithet refers to the oak-like, deeply crenate leaf margin.

Proposed IUCN conservation assessment. Critically Endangered (CR Blab(iti,v)).

This species is only known from the type locality which is in the grounds of a temple

surrounded by farmland and prone to fire damage.

Twenty new Thai Paraboea S57,

Notes. Paraboea quercifolia differs from all other species in the genus by the

combination of a short and slender woody stem with congested leaves at top of stem,

the inflorescences densely covered in minute sticky glandular hairs, the calyx 5-partite

with narrowly ovate lobes and violet-blue corollas.

Paraboea rosea Triboun, sp. nov. (Fig. 7B)

Paraboeae arachnoideae Triboun magnitudine ubique magna, foliis rosulatis, partibus

plantae plurimis pilis arachnoideis dense obtectis et floribus densis similis, sed in foliis

cum pilis arachnoideis minus densis, foliis lanceolatis, pedunculis brevioribus, corollis

rubescentibus et capsulis longioribus recedit. TYPUS: P. Triboun, P. Blenchitra & B.

Sonsupab 3844, 14 September 2007, Thailand, Krabi, Mu Koh Lanta, Koh Talabeng

(holo BK; iso E).

Lithophytic perennial herb to subshrub. Stem erect, 10-40 cm high, 1—1.8 cm in

diam., hard and woody, robust, grooved, cracked and rough, densely covered with

petiole scars, covered with light whitish brown arachnoid hairs, woody root stock

stout. Leaves 10—15, arranged in a dense rosette at the top of the stem; petiole 3—5.5

cm long, 5—7 mm in diam., densely covered with very light whitish brown arachnoid

hairs; blade sub-coriaceous, lanceolate, 8-16 x 3—6.5 cm, apex acute or obtuse, base

crenate or slightly attenuate, margin crenate, secondary veins 7—9 on each side of

midrib, tertiary veins reticulate, densely covered with short multicellular glandular

hairs but somewhat obscured by white arachnoid hairs above, densely covered with

whitish brown arachnoid hairs beneath. Inflorescences 1—3, subterminal, compound

pleiochasia, loosely congested 4-6 branches at each node, 3—6 orders of branching in

each branch, 1—2 terminal orders mostly compound dichasia; peduncle 10—15 cm long,

c. 2.5 mm in diam, sulcate, covered with white arachnoid hairs; bracts lanceolate, c.

1 x c. 0.2 cm, covered with white arachnoid hairs, flowers paired; pedicels 3—9 mm,

covered with brown woolly hairs. Calyx green, red tinged at tips of lobes, fused for

less than 0.5 mm at base, 5-lobed; lobes lanceolate, 1.5—1.8 * c. 0.7 mm, apex acute,

sparsely covered with white hairs. Corolla pinkish red, with a short tube and spreading

lobes; tube c. 2.5 mm; 2 posterior lobes ovate or orbicular, c. 3.5 * c. 2.5 mm, apex

rounded, 3 anterior lobes obovate, c. 3 x c. 2 mm, the 2 lateral ones asymmetrical,

apex rounded or obtuse. Stamens 2, inserted near base of corolla tube, surrounded by

red markings; filament white, c. 2.5 mm, geniculate; anthers yellow, c. 2.5 x c. 2 mm:

staminodes 2, strongly reduced. Ovary ovoid, c. 2 mm: style white to light green, c. 3

mm; stigma capitate, white or light brown. Capsules cylindric, 1—1.9 cm, 2—2.5 mm

in diam., twisted.

Distribution. Endemic to Thailand. So far known only from the type locality.

Ecology. Open sun or in shade on steep limestone cliffs, alt. 10-200 m.

358 Gard. Bull. Singapore 64(2) 2012

Phenology. Flowering and fruiting in July to October.

Vernacular. Dao Pra Dab Pha (avauszaue).

Etymology. The specific epithet refers to the pinkish red colour of the corollas.

Proposed IUCN conservation assessment. Vulnerable (VU D1). This species is only

known from the type locality which is in a legally protected area. Although, the exact

size of the population is uncertain we estimate that it is < 1000 individuals.

Notes. Paraboea rosea is most similar to another new species, Paraboea arachnoidea

Triboun, in its large overall size, the leaves in a rosette, the dense covering of arachnoid

hairs on most parts and the dense flowers, but it differs in a less dense covering of

arachnoid hairs on the leaves, lanceolate leaves, shorter peduncles (10—15 cm in P

rosea, 18-26 cm long in P. arachnoidea), pinkish red corollas and longer capsules

(1-1.9 cm in P. rosea, 0.8—1 em in P arachnoidea).

Paraboea sangwaniae Triboun, sp. nov. (Fig. 7C—D)

Paraboeae axillari Triboun habitu haud frutescenti, foliorum in superficiebus ambabus

cum venis secundariis tertiariisque prominentibus, inflorescentiis axillaribus et

subterminalibus et capsula glabra similis, sed pedunculis robustis, corollis purpureis

usque malvinis cum tubo albo ad pallide marronino et in tubo venis pallide viridibus,

et capsulis majoribus differt. TYPUS: P. Triboun, P. Blenchitra & B. Sonsupab 4039,

26 July 2008, Thailand, Chiang Rai, Mae Fa Luang, Doi Tung (holo BK; iso E).

Lithophytic perennial herb. Stem erect or trailing, 20-60 high, lower part 6-8 mm in

diam, with old and dry persistent leaves, bark densely covered with brown woolly hairs

giving a felt-like appearance. Leaves 8-12, opposite decussate, sometimes congested

towards the apex; petiole 1.5—5 cm long, base amplexicaul, covered with woolly hairs;

blade subcoriaceous, ovate or obovate, 7.5—12 « 4.7—5.5 cm, apex acute, base attenuate

or oblique, margin crenate, secondary veins 8—10 on each side of midrib, tertiary veins

reticulate, venation prominent on both sides; green with some red tinges, covered

with multicellular glandular hairs above, covered with white woolly hairs beneath.

Inflorescences 1—6, compound monochasia, subterminal or axillary; peduncle 4-8 cm

long, c. 2.2 mm in diam., densely covered with white arachnoid hairs; second order

branching c. 4 cm long, flowers 2—6, paired; bracts orbicular, c. 1.9 x c. 2.5 cm, apex

acute, green and maroon, shiny above, glabrous; pedicel 1.2—1.7 cm long, c. 3.5 mm

in diam., covered with brown woolly hairs. Calyx appressed to corolla tube, fused at

base for 6-9 mm long, and then divided into lobes, the upper three lobes further fused

and then divided, base of lobes slightly imbricate; lobes lanceolate, apex acute, upper

3 lobes c. 5 mm long, 4-5 mm wide; lower 2 lobes free, c. 1.2 x c. 0.4 cm; all 5 lobes

green with maroon patches, glabrous. Corolla campanulate, c. 2.3 cm long; tube 2.6—

Fig. 7. A. Paraboea quercifolia Triboun. B. Paraboea rosea Triboun. C—D. Paraboea

sangwaniae Triboun. (Photos: P. Triboun)

360 Gard. Bull. Singapore 64(2) 2012

2.9 cm long, white to pale maroon with pale green veins, limb bilabiate, lower lobe

3-lobed, upper lobe 2-lobed, not symmetrical and not equal; lobes circular to widely

ovate, |—1.2 x 1.2—2 cm, apex rounded to obtuse, pale purple to mauve. Stamens 2;

filaments creamy white, flattened, somewhat parallel, 5-6 mm long, c. | mm in diam.,

fused to corolla tube near base; anthers semicircular in shape, c. 5 mm, creamy yellow

and turning brown when very mature; staminodes 3, inserted at base of corolla tube,

spathulate, 3.54 x c. 0.5 mm, white with yellow tip. Ovary ovoid to ellipsoid, 4-5 x

2.3-2.7 mm; style 4-10 x 1—1.5 mm, pale green, glabrous; stigma linguiform, 4.5—6.5

x ].5—1.8 mm wide, white, apex split into 2 lobes. Capsules ellipsoid, 2.5—3.5 x c. 0.4

cm, twisted, glabrous, calyx persistent.

Additional specimen examined: THAILAND. Chiang Rai: Mae Fa Luang District, Summit of

Doi Tung, 20°19.6'N 99°50.0’E, alt. 1510 m, 23 September 2008, D.J. Middleton, P. Karaket,

P. Triboun, U. Kawatkul & R. Meeboonya 4572 (BK, BKF, E).

Distribution. Endemic to Thailand. So far known only from the type locality.

Ecology. On open limestone summit, alt. 1000-1500 m.

Phenology. Flowering and fruiting in July to December.

Vernacular. Nagarinthra (UA3UNF1), name given by His Majesty King Bhumibol of

Thailand.

Etymology. The specific epithet honours Princess Sangwan (1900-1995), the late

Princess Mother of King Ananda Mahidol (Rama VIII) and King Bhumibol Adulyadej

(Rama IX), under whose initiative the conservation and reforestation of the collection

locality at Doi Tung in Chiang Rai in Thailand was established.

Proposed IUCN conservation assessment. Critically Endangered (CR Blab(ili,v)).

This species is only known from Doi Tung, has an Extent of Occurrence of < 100 km?

and is known from only one population. The area is not protected and there has been

a reduction in the quality of the habitat through large scale tourism which has also led

to a reduction in the number of mature individuals.

Notes: Paraboea sangwaniae is most similar to the new species Paraboea axillaris

Triboun in its non shrubby habit, the secondary and tertiary veins prominent both

abaxially and adaxially, the axillary and subterminal inflorescences and the glabrous

capsule, but it differs in having robust peduncles, purple to mauve corollas with a

white to pale maroon tube and pale green veins in the tube, and larger capsules (2.5—

3.5 cm in P. sangwaniae, 1.8—2.2 cm in P. axillaris).

This species was referred to as Trisepalum sp. nov. | in Puglisi et al. (2011).

Twenty new Thai Paraboea 361

Paraboea siamensis Triboun, sp. nov. (Fig. SA—B)

Paraboeae acauli (Barnett) C.Puglisi habitu perenni suffruticoso, phyllotaxe ad instar

rosulae, foliorum laminis ellipticis, pilis arachnoideis densis et flore magno similis sed

foliis minoribus, bracteis calycibusque majoribus ac magis spectabilibus et corolla

caerulea differt. TYPUS: P. Triboun, B. Sonsupab & P. Yothakaew 4565, 7 September

2010, Thailand, Tak, Umphang, Doi Hua Mot (holo BK; iso BKF, E).

Lithophytic perennial herb to subshrub; each plant with 1—5 branches, lower part of

each branch with persistent old and decayed leaves; rootstock long, rough. Leaves

10—25, arranged in a dense rosette at the top of the stem: sessile or petiole very short;

blade lanceolate to elliptic, S—12 cm = 1.5—3.5 cm, the lower ones larger than the upper

ones, apex acute, base attenuate and winged, amplexicaul to petiole and node, margin

slightly crenate, both sides covered with white arachnoid hairs, more densely so

beneath, secondary veins 5—8 on each side of midrib. Inflorescences 1-4, subterminal

or axillary near shoot apex, a compound dichasium with | order of branching; peduncle

5—9 cm long, 2—2.5 mm in diam., covered with white arachnoid hairs; axis 1.2—1.5

cm long, covered by white arachnoid hairs; bracts reddish mauve in the upper half,

broadly ovate, c. 2.5 cm long, apex rounded, margin erose, sparsely covered with

white arachnoid hairs, uppermost bracts 1.8—2 cm long, c. 1.7 cm wide; flowers paired;

pedicels c. 4 mm, covered with white arachnoid hairs. Calyx green, tube 4-5 mm long,

c. 5 mm in diam., 3-lobed; lobes obovate, ovate or oblong, 1—1.4 x 0.7—1 cm, margin

erose, sparsely covered with white hairs, each lobe closed and slightly imbricated.

Corolla blue, campanulate; tube 1.2—1.8 cm long; posterior and anterior lobes nearly

same shape and size, broadly ovate, orbicular or obovate, 6-8 =< 0.9-1.1 cm, apex

rounded or obtuse. Stamens 2, inserted near base of corolla tube; filaments white,

geniculate, c. 5 mm long; anthers yellow, reniforme, c. 3 x c. 6 mm; staminodes 2, 2—3

mm long, medial staminode not apparent. Disc annular, c. 2 mm high. Ovary ovoid,

c. 3.5 mm long, green, glabrous; style 1.3—1.5 cm long, white to light green; stigma

minute, obcordately bilobed, yellowish brown. Capsules narrowly ovate, c. 1 cm long,

slightly twisted, calyx persistent.

Additional specimens examined: Thailand, Tak, Um Phang, Doi Hua Mot, alt. 900 m, 23 April

2004, R. Pooma, K. Phattarahirankanok, S. Sirimongkul & M. Poopath 4606 (BKF, E, L).

Distribution. Endemic to Thailand. So far known only from the type locality.

Ecology. Among rock on open limestone hill.

Phenology. Flowering and fruiting in July to November.

Vernacular. Sri Siam (ASag1N).

Etymology. The specific epithet refers to the collection locality in Thailand, Siam

being the former name for Thailand.

362 Gard. Bull. Singapore 64(2) 2012

Proposed IUCN conservation assessment. Critically Endangered (CR Blab(iii,v), D).

This species is only known from a very small population of < 50 individuals. It is in a

legally protected area but the but site is prone to fire and is impacted by tourism.

Notes. Paraboea siamensis is unusual because although it is quite clearly part of

the Trisepalum group it has a minutely capitate stigma rather than the linguiform

stigma usually found in species traditionally placed in Trisepalum. It is most similar

to Paraboea acaulis (Barnett) C.Puglisi in its perennial suffruticose habit, rosette-

like phyllotaxy, elliptic leaf blades, the dense covering of arachnoid hairs and the

large flower, but it differs in the smaller leaves (S—12 cm long in P. siamensis, 6—14

in P. acaulis), more showy and larger bracts (1.8—2 cm in P. siamensis, c.1.2 cm

in P. xylocaulis) and calyces (1.4—-1.9 cm long in P. siamensis, 1—1.2 cm long in P.

xyvlocaulis), and the blue corolla.

Paraboea takensis Triboun, sp. nov. (Fig. 8C—D)

Paraboeae strobilaceo (Barnett) C.Puglisi magnitudine modica, foliis chartaceis,

venatione prominenti et floribus albis similis sed foliis haud scabridis, bracteis ovatis

vel naviculiformibus et floribus paucioribus recedit. TY PUS: P. Triboun 3606, 14 July

2006, Thailand, Tak, Umphang, Doi Hua Mot (holo BK; iso E).

Lithophytic perennial herb. Stem erect, 20—SO cm high, c. 2.5 mm in diam., brownish

grey, with 1—2 young shoots at the base of stem; woody root stock slender and finely

striped. Leaves 8-15, loosely opposite decussate; upper pairs sessile, lower ones with

petiole 12.5 cm long, glaucous, densely covered with white hairs; blade papery,

lanceolate, elliptic or ovate, 3-10 x 1.4—3.3 cm, apex acute, base cuneate or attenuate,

margin slightly crenate, slightly glaucous, covered with short glandular trichomes

above, covered with brown woolly hairs beneath, secondary veins (6—)10—12 on each

side of midrib, tertiary veins reticulate, venation prominent beneath. Inflorescences

2—5, subterminal, a single flower or in compound monochasia, with 1—2 orders of

branching; peduncle 2.5—2.8 cm long, densely covered with white arachnoid hairs,

bracts ovate or naviculiform, 1.2—-1.5 <x 8—1.1 cm, apex obtuse or acute, outside

glaucous and sparsely covered with arachnoid hairs, inside green; pedicels to c. 1.2

cm, unequal, some nearly sessile, densely covered with white hairs. Calyx cotyliform,

3-lobed; lobes ovate or lanceolate, upper 3 lobes fused at base, c. 1.2 x 0.4—0.5 cm,

the 2 lateral lobes covering the middle lobe, 2 lower lobes c. 1.2 x c. 0.6 cm, free.

Corolla white, campanulate, tube ¢. 1.8 cm long, lobes broadly obovate or broadly

ovate, 0.8—1.1 < 1.3—1.5 cm, apex rounded. Stamens 2; filaments creamy white, c. 3.5

mm long, geniculate; anthers light brown, c. 4.5 x c. 4.8 mm; staminodes 3, linear, c.

2.5 mm. Ovary ovoid, c. 3 mm, glabrous; style white, c. 7 mm long; stigma linguiform,

c. 1.2 cm long, white. Capsules not seen.

Distribution. Endemic to Thailand. So far known only from the type locality.

Twenty new Thai Paraboea

Fig. 8. A-B. Paraboea siamensis Triboun. C—D. Paraboea takensis Triboun. (Photos: P.

Triboun)

364 Gard. Bull. Singapore 64(2) 2012

Ecology. On open limestone rock in mixed deciduous oak forest, alt. 600-1000 m.

Phenology. Flowering and fruiting in late June to October.

Vernacular. Khao Tog Yai (aIAaNluA).

Etymology. The specific epithet refers to the locality in Tak province in Thailand.

Proposed IUCN conservation assessment. Critically Endangered (CR Blab(iii,v), D).

This species is only known from a very small population of < 50 individuals. It is in a

legally protected area but the but site is prone to fire and is impacted by tourism.

Notes. Paraboea takensis is similar to Paraboea strobilaceum (Barnett) C.Puglisi in

its moderate size, papery leaves, prominent venation and white flowers, but it differs

in its non scabrid leaves, ovate or naviculiform bracts and fewer flowers (2—8 in P

takensis, 4-10 in P. strobilaceum).

Paraboea tenuicalyx Triboun, sp. nov. (Fig. 9A)

Paraboeae harrovianae (Craib) Z.R.Xu in inflorescentiis pleiochasiis compositis, lobis

calycis minimis linearibusque et corollis dilute malvinis similis sed apicibus foliorum

acutis, pedunculis longioribus, inflorescentiis (pedunculi, bracteae, pedicelli et

calyces) sine pilis albis arachnoideis, et inflorescentiis magis laxis recedit. TYPUS: P.

Triboun 3837, 28 August 2007, Thailand, Kanchanaburi, Tung Yai Naresuan Wildlife

Reserve, Lai Wo Subdistrict, Ban Saneh Pawng area (Karen village) (holo BK; 1so E).

Lithophytic perennial herb. Stem erect, 8—18 cm high, solitary, 4-6 mm in diam, densely

covered with greyish brown woolly hairs, root stock short and robust, sometimes

young shoots sprouting from root stock. Leaves 4-8, opposite decussate, slightly

congested at the apex of the stem; petiole (2—)4—8(—12) cm long, 1.5—2.5 mm in diam.,

densely covered with brown woolly hairs; blade coriaceous, lanceolate or ovate, 8-17

x 3.5—6.5 cm, apex acute, base cuneate or obtuse, margin indistinctly crenate, glaucous

beneath, secondary veins 8-11 on each side of midrib, obscure above, prominent

beneath; glabrous above, densely covered with short light brown woolly hairs beneath.

Inflorescences 1—2, subterminal, compound pleiochasia, 4-6 branches at each node,

each branch with 4—5 orders of branching; peduncle 30-60 cm long, 3-4.5 mm in

diam., densely covered with light brown woolly hairs; bracts linear, 1—2.5 = 0.1—0.5

cm, apex acute, the upper ones strongly reduced; flowers paired; pedicels slender,

3—7 mm long, glabrous. Calyx very small, fused at base for 1—1.5 mm, 5-lobed; lobes

narrowly linear, 1—1.5 x 0.2—0.3 mm, apex obtuse, glabrous. Corolla light mauve, with

a short tube and spreading lobes; lobes widely ovate, orbicular or obovate, 2—4 x 2.54

mm, apex rounded or obtuse. Stamens 2; filament light green, slightly geniculate, c. 2

mm long; anthers yellow, reniforme, c. 1.8 < c. 2 mm; staminodes 2, c. 0.8 mm long,

median staminode not apparent. Ovary ovoid, c. 2.5 mm long, light green, glabrous;

Twenty new Thai Paraboea 365

style c. 2.5 mm long, white; stigma capitate, white. Capsules slender, narrowly

cylindric, 2—2.5 x c. 0.1 cm, twisted, glabrous.

Additional specimens examined: THAILAND. Kanchanaburi: Kin Sayok, about 120 km NW

of Kanchanaburi, alt. 100-150 m, 4 August 1946, 4. Kostermans 1451 (US); Thong Pha Poom,

4 July 1973, J.F Maxwell 73-103 (AAU); Sangklaburi, Toong Yai Naresuan Wildlife Reserve,

Lai Wo subdistrict, Ban Saneh Pawng area (Karen village), 300 m, 11 October 1993, LF

Maxwell 93-1223 (L); between Huay ban kao and Kritee, alt. 500 m, 4 July 1973, R. Geesink

& C. Phengkhlai 6084 (AAU); near Neeckey, near Wangka, alt. 150 m., G. den Hoed 580 (K).

Distribution. Endemic to Thailand. So far known only from Kanchanaburi Province.

Ecology. On limestone rocks in dry evergreen to mixed deciduous forest, alt. 300—S00 m.

Phenology. Flowering and fruiting in late June to November.

Vernacular. Khao Tok Phra Nares (aTAMANWSZULSA ).

Etymology. The specific epithet refers to the minute calyces.

Proposed IUCN conservation assessment. Vulnerable (VU Blab(iil,v)). This species

is quite widespread within Kanchanaburi and most populations are in legally protected

areas. However, there are fewer than 10 populations and some of these populations are

prone to fire damage from neighbouring farming practices leading to a nett erosion in

the quality of the habitat and the number of mature individuals.

Notes. Several of the paratypes listed above were included in Paraboea harroviana

(Craib) Z.R.Xu var. harroviana by Xu et al. (2008) although they did note that the

material from Kanchanaburi was morphologically distinct. With additional material

and field observations we conclude that these populations form a distinct species

differing from Paraboea harroviana in leaf apex shape, longer peduncles (30—60 cm

in P. tenuicalyx, 10-20 cm in P. harroviana), laxer inflorescences, and the lack of

white arachnoid hairs on all inflorescence parts.

Paraboea vachareea Triboun & Sonsupab, sp. nov. (Fig. 9B)

Paraboeae harrovianae (Craib) Z.R.Xu magnitudine ubique modica, dichasio

composito congesto et corollis dilute caeruleis similis, sed foliis lanceolatis cum

apicibus acutis et inflorescentiis minoribus differt. TYPUS: P. Triboun, B. Sonsupab

& P. Yothakaew 4566, 8 September 2010, Thailand, Tak, Umphang, Palata (holo BK;

iso E).

Lithophytic perennial herb. Stem erect, 10-25 cm high, 3-4 mm in diam., with a few

twigs and branches 2—5 cm long, 2—5 mm in diam., woody root stock slender and

366 Gard. Bull. Singapore 64(2) 2012

finely striped. Leaves 9-18, densely opposite decussate; petiole brown, 3—8 cm long,

covered with woolly brown hairs; blade coriaceous, lanceolate, 5—11 cm = 3—5.5 cm,

apex acute, base cordate or cuneate, margin crenate, shiny green above, covered by

short dark woolly brown hairs beneath, secondary veins 5—7 on each side of midrib.

Inflorescences 1—5, subterminal, c. 3 orders of branching in each compound pleiochasia,

some congested with 4-8 branches on each order; peduncle (1.5—)10—25 cm long,

covered by woolly brown hairs; bracts linear, 0.6—1.5 cm long, the upper ones strongly

reduced; flowers paired; pedicels 6-7 mm, glabrous. Calyx brownish green, fused less

than | mm at base, 5-lobed; lobes narrowly linear, c. 2.5 mm long, glabrous. Corolla

light blue, campanulate; tube 0.5—1 cm long; 2 posterior lobes obovate, 4-5 = c. 2.5 mm,

apex rounded; 3 anterior lobes obovate, c. 5—5.5 « 3 mm, apex rounded, the middle one

largest. Stamens 2, inserted near base of corolla tube; filament light green, c. 2 mm long,

geniculate; anthers white, c. 2 < c. 3.5 mm. Staminodes 3, narrowly linear, 1|—2 mm long.

Ovary ovoid, c. 3 x c. 1.5 mm long; style white, c. 3 mm long; stigma capitate, white.

Capsules narrowly cylindric, twisted.

Distribution. Endemic to Thailand. So far known only from the type locality.

Ecology. On limestone rock in dry evergreen to mixed bamboo deciduous forest, alt.

600-900 m.

Phenology. Flowering and fruiting in August to November.

Vernacular. Va Cha Ree Phun (J#3w39au).

Etymology. The specific epithet honours Mrs. Vacharee Prachasaisoradej, a senior

botanist at the Bangkok Herbarium (BK), who encouraged the first author in the field of

plant taxonomy.

Proposed IUCN conservation assessment. Critically Endangered (CR Blab(it1,v), D).

This species is known from only one small population of fewer than 50 individuals.

Although it is in a legally protected area the population is prone to fire damage from

neighbouring farming practices.

Notes. The type of inflorescence in this species is found in many species of Paraboea,

particularly in Thailand. It is most similar to Paraboea harroviana (Craib) Z.R.Xu in its

size and the light blue corollas. It can be distinguished by its lanceolate leaves with an

acute apex and its dense inflorescences.

Paraboea xylocaulis Triboun, sp. nov. (Fig. 9C—D)

Paraboeae siamensi Triboun planta magna robusta, foliis lanceolatis, calyce spectabili

et corollis pallide purpureis similis sed calycibus bracteisque minoribus et foliis,

pedunculis pedicellisque cum pilis albis arachnoideis obtectis differt. TYPUS: P.

Twenty new Thai Paraboea 367

Triboun, B. Sonsupab & P. Yothakaew 4585, 25 September 2010, Thailand, Border of

Krabi-Phangnga bay, Pa Koh (holo BK; iso BKF, E).

Lithophytic perennial herb or small shrub. Stem woody, up to 70 cm high, with many

twigs and branches, 15—20 cm long, 5—1.5 cm in diam., hanging from cliff but then with

upright shoots, bark greyish brown, cracked and rough, lower part of each branch with

persistent old and decayed leaves; root stock woody, robust, cracked and rough. Leaves

8-15, densely opposite decussate or 3-whorled, congested at the apex; appearing sessile

but actually a winged petiole 1.2—5 x 0.25—0.5 cm, covered with light brown arachnoid

hairs and brownish orange hairs along the midrib; blade coriaceous, lanceolate, 2.7—7.5

x 1.42.5 cm, apex acute, base attenuate, margin indistinctly crenate, densely covered

with multicellular glandular hairs above, densely covered with brown woolly hairs

beneath, secondary veins 8—10 on each side of midrib. Inflorescences 6—20 (2—6 on each

branch), subterminal, in | pair of monochasia, rarely compound monochasia; peduncle

c. | cm long, covered with brownish orange woolly hairs; bracts brownish green,

globose, c. 1.2 x c. 1 em, hooded, apex rounded, glabrous; flowers paired; pedicels 3—9

mm long, unequal, glabrous. Calyx shiny green with red margin, 3-lobed, almostly free;

lobes elliptic to oblong, 1—1.2 cm long, the posterior lobe 7-8 mm wide, apex obtuse or

shallowly 2—3-lobed, the inferior lobes 5—S.5 mm wide, apex obtuse, glabrous. Corolla

campanulate; tube light purple, c. 1.2 cm long; lobes purple, orbicular or broadly ovate,

0.8-0.9 x 0.9-1.2 cm, apex rounded. Stamens 2, inserted near the base of corolla tube;

filament white, c. 6 mm long geniculate; anthers white or creamy white, c. 3 x c. 3 mm;

staminodes 3, white, linear, 0.5—2 mm, the middle one smaller than the lateral ones.

Ovary creamy yellow, oblong to ovoid, c. 3 mm long; style white, c. 1 cm long, covered

with glandular hairs; stigma linguiform, c. 5 mm, white. Capsule ovoid, c. 1.5 c. 0.85

cm, green tinged red, glabrous, surrounded by persistent calyx.

Additional specimens examined: THAILAND. Krabi: Koh Hin Bai, 31 October 2006, P. Triboun

3674 (BK, E).

Distribution. Endemic to Thailand. So far known only from the type locality.

Ecology. On steep cliffs on limestone islands, alt. 30-150 m.

Phenology. Flowering and fruiting in August to November.

Vernacular. Cha Ha Nu Man (21WiWN1u).

Etymology. The specific epithet refers to the distinct woody stem of this plant; xy/o (Gk.)

= woody, caulis = stem.

Proposed IUCN conservation assessment. Critically Endangered (CR D). Although this

species is only known from the type locality this is in a legally protected area. However,

the population contains fewer than 50 individuals.

Gard. Bull. Singapore 64(2) 2012

Fig. 9. A. Paraboea tenuicalyx Triboun. B. Paraboea vachareea Triboun & Sonsupab. C—D.

Paraboea xylocaulis Triboun. (Photos: P. Triboun)

Twenty new Thai Paraboea 369

Notes. Paraboea xylocaulis is most similar to the new species Paraboea siamensis

Triboun in being a large and robust plant, the lanceolate leaves, showy calyx and blue

corollas, but it differs in the smaller calyces (1—1.2 cm long in P. xylocaulis, 1.4-1.9 cm

long in P. siamensis) and bracts (c. 1.2 cm in P. xylocaulis, 1.8—2 cm in P. siamensis)

and the covering of white arachnoid hairs on the leaves, peduncles and pedicels.

This species was referred to as Trisepalum sp. nov. 3 in Puglisi et al. (2011).

ACKNOWLEDGEMENTS. We thank the many additional collectors who helped us with

the field work: V. Chamchumroon, P. Karaket, U. Kawatkul, S. Lindsay, R. Meeboonya, T.

Phutthai, S. Saengrit, R. Simma, S. Suddee, P. Suksathan, P. Yothakaew, B. Sonsupab, P.

Inthachup, P. Rukkid, P. Piriyavinit, P. & P. Wanichananan, T. Chuchan and W. Dongkumfu.

We thank the Royal Botanic Garden Edinburgh (E), The Forest Herbarium Bangkok (BKF),

The Bangkok Herbarium (BK), Queen Sirikit Botanical Garden (QBG), Prince of Songkhla

University (PSU) and the Biodiversity Research and Training Program (BRT) for logistical

and/or financial support for the field work. We also thank the Royal Society of Edinburgh, the

Royal Botanic Garden Edinburgh Members Small Project Fund and the Sibbald Trust for funds

for the first author to travel to Edinburgh. We also thank the following herbaria which have

loaned herbarium material or hosted a visit for this work: A, AAU, BK, BKF, C, E, K, KKU,

L, NY, P. PSU, QBG, SING, US, W. We thank Dr. Robert Mill (E) for translating the diagnoses

into Latin and Claire Banks for the illustrations.

References

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concepts for Boea and its allies. Notes Roy. Bot. Gard. Edinburgh 41: 401-452.

Chen, W.H., Mdller, M., Shui, Y.M. & Zhang, M.D. (2008) A new species of Paraboea

(Gesneriaceae) from a karst cave in Guangxi, China, and observations on variations in

flower and inflorescence architecture. Bor. J. Linn. Soc. 158: 681-688.

Clements, R., Sodhi, N.S., Schilthuizen, M. & Ng, P.K.L. (2006) Limestone karsts of Southeast

Asia: imperiled arks of biodiversity. BioScience 56: 733-742.

Inthachub, P., Vajrodaya, S. & Duyfjes, B.E.E. (2009) Review of the genus Stichoneuron

(Stemonaceae). Edinburgh J. Bot. 66: 213-228.

IUCN (2001) IUCN Red List Categories and Criteria, Version 3.1. Gland and Cambridge:

IUCN Species Survival Commission.

Kiew, R. (2010) Two new species of Paraboea (Gesneriaceae) from Peninsular Malaysia and

Thailand. Edinburgh J. Bot. 67: 209-217.

Lindsay, S. & Middleton, D.J. (2004) Adiantum phanomensis (Adiantaceae), a new fern species

from Peninsular Thailand. Harvard Pap. Bot. 8: 137-140.

Lindsay, S., Middleton, D.J. & Suddee, S. (2008) Two new species of ferns from Thailand. Thai

Forest Bull., Bot. 36: 46-51.

Middleton, D.J., Puglisi, C., Triboun, P. & MGller, M. (2010) Proposal to conserve Paraboea

against Phylloboea and Trisepalum (Gesneriaceae). Taxon 59: 1603.

Middleton, D.J. & Triboun, P. (2010) Two new species of Petrocosmea (Gesneriaceae) from

Thailand. Thai Forest Bull., Bot. 38: 42-47.

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Phuphathanapong, L. (2006) New taxa of Aristolochia (Aristolochiaceae) from Thailand. Thai

Forest Bull., Bot. 34: 179-194.

Puglisi, C., Middleton, D.J., Triboun, P. & M6ller, M. (2011) New insights into the relationships

between Paraboea, Trisepalum and Phylloboea (Gesneriaceae) and taxonomic

consequences. 7axon 60: 1693-1702.

Triboun, P. & Middleton, D.J. (2010) A new species of Damrongia (Gesneriaceae) from

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genera of vascular plants, volume 7. Flowering plants: Dicodyledons; Lamiales (except

Acanthaceae including Avicenniaceae). Pp. 63-158. Berlin & Heidelberg: Springer.

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Gardens’ Bulletin Singapore 64(2): 371-479. 2012 371

Annonaceae of Borneo: a review of the climbing species

I.M. Turner

Research Associate, Royal Botanic Gardens Kew

Research Associate, Singapore Botanic Gardens

Present address:

Fairfield, Pett Level Road, Winchelsea Beach, East Sussex TN36 4ND, U.K.

Turner187@pbtinternet.com

ABSTRACT. The climbing species of the Annonaceae native to Borneo are reviewed. Eight

genera of lianas are represented: Artabotrys (17 spp.), Desmos (4 spp.), Fissistigma (15 spp.),

Friesodielsia (9 spp.), Mitrella (3 spp.), Pyramidanthe (1 sp.), Sphaerocoryne (1 sp.) and

Uvaria (19 spp.). The species are described. Synonymy, typifications and keys for identification

are included. No nomenclatural novelties are presented in this account.

Keywords. Brunei Darussalam, Indonesia, Malaysia, Malesia, lianas, systematics, taxonomy

Introduction

The Annonaceae are essentially a pantropical family and an importent element in the

flora of the lowland tropical forest. The family is well represented in Borneo, which is

hardly surprising given that it is a large equatorial island with a warm and wet climate

and was, until recently, covered in vast tracts of tropical rain forest. Most species

of Annonaceae are free-standing trees, treelets or shrubs, but a substantial minority

are woody climbers (lianas) or scandent shrubs. It is this element of the family as

represented in Borneo that is the subject of the present paper.

The annonaceous climbers of Borneo merit attention for several reasons.

Firstly, there has been relatively little work on them by taxonomists in recent decades.

Secondly, they are particularly important in terms of representation amongst the liana

flora. While the Annonaceae are pantropical, climbing species are almost confined

to the Old World. Gentry (1991) reports only six species from two genera in the

Neotropics, compared to a rich diversity in the Palaeotropics e.g. 41 species in West

Africa (Jongkind & Hawthorne 2005), 49 species recognised here for Borneo. While

published inventories of lianas in the forests of Borneo are rare and none is extensive,

the data suggest that the Annonaceae are typically among the top three families in terms

of representation by species or numbers of individuals (Putz & Chai 1987; Appanah et

al. 1993: DeWalt et al. 2006), contesting top spot with the Leguminosae and Palmae.

The climbing Annonaceae of Borneo

The present revision is greatly facilitated by the strong tendency of life form to be shared

across all members of a genus within the Annonaceae. Sinclair (1955) emphasised the

SI Gard. Bull. Singapore 64(2) 2012

taxonomic value of knowing if a plant is a climber or not in the Annonaceae of the

Malay Peninsula. Among the species in Borneo, the split of genera in to lianas or

trees/treelets appears completely sound. The only possible doubt concerns Desmos

and Sphaerocoryne. In cultivation, members of both genera will grow as shrubs if

given no support. A number of Sphaerocoryne collections I have seen from Borneo are

reportedly from trees, but this also happens for other genera of strict climbers such as

Uvaria, so confusion between climber and mechanical host may be the explanation.

Alternatively some species may be found as both climbers and free-standing plants in

nature. In Borneo, the species of Dasymaschalon are treelets, but there is at least one

climbing species in Thailand (Wang et al. 2009).

I recognise eight genera in the current revision. This is fewer than the 11

Sinclair (1955) recognised in his revision for the Malay Peninsula. This difference 1s

explained by the reduction of genera into Uvaria. Cvathostemma, Ellipeia and others

have been considered untenable after molecular phylogenies became available (Zhou

et al. 2009). Notably, the only Asia-Pacific genus of climbing Annonaceae not recorded

from Borneo is the poorly known New Guinea endemic Schefferomitra.

Molecular phylogenies make it clear that the climbing habit has arisen

independently several times in the Annonaceae. Among the Bornean representatives,

two lines are present. All the genera, except Artabotrys, are members of tribe Uvariae

of subfamily Annonoideae (Chatrou et al. 2012), which consists mostly of lianas and

represents a majority of the Old World climbing Annonaceae species. Artabotrys 1s

a member of the same subfamily, but forms tribe Xylopieae with the non-climbing

Xylopia. Artabotrys differs from the other Bornean climber genera in having specialised

structures to assist climbing. Recurved hooks are formed from the inflorescence axes.

These act as grapples, securing attachment to the supporting plants. Members of the

other genera climb by means of twining and coiling stems, a response apparently

elicited by the presence of a support, as growth appears relatively straight otherwise.

Phyllotaxy also differs between the two climber clades; Artabotrys typically has

spirally arranged leaves whereas the other genera show a distichous leaf arrangement

(Johnson 2003).

Identification

Climbing Annonaceae material from Borneo 1s relatively easily identified as such.

Even sterile material should be identifiable to family given the fairly clear characters

of twig and leaf available.

Annonaceae (description applies to the climbing genera of Borneo)

Large woody climbers never truly herbaceous, unarmed except for Artabotrys. Twigs

with fibrous bark, often striate or latticed, medullary rays broadening outwards

generally obvious to the naked-eye as a star-shaped pattern in cross-section, pith

Climbing Annonaceae of Borneo 373

often septate. Indumentum hairs simple; stellate or caespitose in Uvaria. Leaves

simple, entire, generally alternate and distichous, rarely spiral, membranous to

coriaceous, often glaucous, pinnately nerved, with lateral veins looping within the

lamina margin, though often obscurely, stipules absent. Inflorescences axillary or

terminal, sometimes appearing leaf-opposed or supra-axillary through growth of the

uppermost axillary vegetative shoot or the inflorescence axis being partly adnate to

the vegetative shoot. Flowers bisexual, trimerous, rarely dimerous, solitary, paired

or in few—many-flowered fascicles, regular and actinomorphic, sessile or stalked,

often nodding or pendulous, fleshy and often rather brittle, typically bracteate, often

fragrant; development often open without an obvious bud stage (not so in some Uvaria

spp.). Sepals (2—)3(-4), hypogynous, valvate or imbricate, free to entirely connate.

Petals mostly green, white or yellow, sometimes red or purple, hypogynous, usually

2 whorls of (2—)3(-4), valvate or imbricate, free or connate at the base, often apically

coherent, usually alternating with the sepals. Stamens hypogynous, numerous, spirally

arranged, closely packed on the torus, sometimes staminodes present, filaments absent

or short, anthers linear, generally opening by longitudinal slits, connective mostly

with a truncate dilated apex, but sometimes produced to a cone, point or tongue-like

projection, filaments short and free. Carpels several to many, free or slightly connate

at the base, ovules 1—many, basal or lateral; styles free or united, mostly short and

terete, stigma capitate, oblong or variously folded. Fruits several to many fleshy or

somewhat woody indehiscent, sessile or stipitate monocarps, monocarps cylindrical

(moniliform in Desmos), ellipsoidal, globose or irregular, with 1—many seeds in 1—2

rows, with seeds transverse or longitudinal to long axis of monocarp. Seeds sometimes

with a circumferential groove, sometimes pitted, endosperm abundant, ruminate, hard,

oily, often divided almost to the axis into a series of horizontal plates; embryo straight,

minute.

Key to genera of climbing Annonaceae in Borneo

MPC ARABS ONES EID x cnr tne he fe 8Ot «des ce suaen nanen sensed bacunsntasedceeeesoustnssuteaseteess Uvaria

SOMES Cell Ae AIT SA SOME me Sepa tahe-nscccoce Si ccacese cca uueiastetlentoeneauhshennvonesbaraaataasexaeracnicaars yi

Pm RNIN SCENMEG AXIS MOOKEC. x c0ccc-<cec¢eeacvacvencesdctsccesesivuccueansotovorassnvessosse Artabotrys

PMB TORESCEMGSPA NTS MOUMOOK CG: <n ca5-5.s20csqccecsessesoovncwtoennseescsalosnessosascrteiorscsveeessacs 3

3a. Petal whorls of similar shape and size with petals at least 2 cm long, monocarps

(ETCOART ITF GUTTA § xcye ects hee be Uae Sn EAP CU RRR ESS REPS OPTS RECN SD rin en Desmos

b. Petals whorls of dissimilar shape and size, or if similar then less than 2 cm

Mee eeamO TNO Sekt [9S MO alan OM MTL OI aoe anc d esc en ea en encomctebireowtolon noid vewsesecnnctesseesseeans a

Adee etlawnOors Of sitailar Shape and SIZE «.....2.2:...s0-.c0sceeoccevsreoeecees Sphaerocoryne

Baer beralynoris dissmailar in Shape and/OL SIZE ...c.-scc-cscc.cexpsteesoceneeneesdesesensenscee> 5

374 Gard. Bull. Singapore 64(2) 2012

5a. Leaves with a pair of glands in the margin of the lamina base, one on

each side of the petiole, apex of anther connective more or less flat .......

Friesodielsia

6a. Leaves with lateral nerves distinct, interlateral veins absent, tertiary venation

scalariform, inflorescences generally branched and multi-flowered, though

some species have |- or 2-flowered inflorescences, calyx connate near the base

6) 0h Ane ear aes nen remdn herric ih. UA reen eigacassiarransristandaoa aces! LOOSE DG!

b. Leaves with lateral nerves indistinct, or 1f distinct with clear interlateral nerves,

tertiary venation not scalariform, inflorescences usually 1- or 2-flowered, calyx

mostly or entirely comnate....<-.2..c2--cstene econ ee eeee secrete eee eee eee ee 7

7a. Calyx united into a shallow circular or weakly three-pointed cup..............:0008

wk sb fst de shes ccue ene Sovaee gaee soutew sabe Rate PSS ENESE CE SOE ee ee Pyramidanthe

~b. Calyx united into a distinctly three-pointed Cup...............:cscesceseeeeeees Mitrella

An index to the specimens identified in this study is given in the Apendix.

ARTABOTRYS R.Br.

(Greek, artane, artao = that by which something is hung up,

botrys = cluster, bunch of grapes; the hooked inflorescences)

Bot. Reg. 5 (1820) t. 423. Sinclair, Gard. Bull. Singapore 14 (1955) 246. Nurainas,

Floribunda 2(5) (2004) 117-127. Turner, Folia Malaysiana 10 (2009) 59. TYPE:

Artabotrys odoratissimus R.Br., nom. illegit. (= Annona hexapetala L.f., = Artabotrys

hexapetalus (L.f.) Bhandari)

Ropalopetalum Griff., Not. Pl. Asiat. 4 (1854) 716. TYPE: Ropalopetalum uniflorum

Griff.

Woody climbers. Twigs glabrous or hairy. Leaves chartaceous to coriaceous, midrib

sometimes raised above, lamina often decurrent to petiole. Inflorescences extra-axillary,

borne on thick woody peduncles that form reflexed, often laterally compressed, hooks

that assist with climbing, fascicled or single-flowered. Flowers bisexual, white or

yellow, often fragrant, sepals 3 valvate, free or united at the base, petals 6, valvate in

two whorls, subequal, clawed with concave bases connivent, closely adpressed over

reproductive organs, inner whorl often remaining attached by their bases when it falls,

torus flat or concave, stamens many, truncate, apex dilated, carpels numerous, ovules

2, basal. Monocarps few to many, cylindrical or ellipsoidal, often sessile. Seeds 2,

erect, collateral.

Climbing Annonaceae of Borneo 375

Distribution and diversity: 100 or more species in the Old World tropics from Africa

to Australia. 17 species recorded from Borneo.

Notes. The persistent inflorescence hooks make Artabotrys a relatively easy genus to

recognise.

Key to Artabotrys species

la. Petal blades of both whorls less than 2 mm wide, monocarps per pedicel

typically 3 or fewer (monocarps of A. sarawakensis are unknown but carpel

Mumia IMCS HS 10, DG EUG) 2 .<05.2220-50025sbu caps otic cht <2cbecenact Lata ethene tase 2

b. Petals blades of one or both whorls at least 2 mm wide, more than three

ULULEGE FES PSs (LE 2e [Bee Ree pene oee aetna on a eee eee eee 5

ae seedicehol flower longer than 16 mM .............::..:dsescseeececennes 4 4. sarawakensis

PeeCoIcc Ge NoOWwer Up £0 16 mina TOP: <.2c...22.2.2..cccescces-seneasesontevenstetade-eneeencseees 3

3a. Petals drying black with sparse pale hairs, monocarps more than 2 cm wide

NUR NDR ea fe So Neha wo RG caida cpser ecko arene vr oeea ata 4. veldkampii

b. Petals drying brown often with dense pale tomentum, monocarps drying less

LT LU OVE. Soe nal goed 9 ee eer rh ba REP RARER L ot EERE SRE IE ON IW =

4a. Pedicel longer than flower, outer petals with relatively little distinction between

claw and blade when viewed abaxially, blade relatively flat, monocarps 10

mm or more in diameter, longitudinally ridged and beaked .........0...00.. eee

vo cS acts Ste oc en A 4. gracilis

b. Pedicel shorter than flower, outer petals with clear distinction between claw

and filiform blade, monocarps less than 10 mm in diameter, smooth with

OU DINL ST ST SL 2 0 eee a 4. suaveolens

Saesidaes oLanner petals 2.1mm wide Or Iess. .......-<....2<.-sce<cusesesccnteceiset-auensedesosese 6

Pemebindes of inter peialssmore than 2 Mm WIdE::..;..2..:2...-.2vsee-becsecdasennenssbessbes 8

6a. Flower pedicel less than 5 mm long, petals covered with red-brown hairs .....

ngs HE, SE eae ne eo nr en EE ae 4. sumatranus

b. Flower pedicels more than 5 mm long, petals densely covered with pale hairs

ep ES GA SEE ROMP Pe eee RR ne SR eee 7

7a. Pedicel shorter than flower, monocarps with apex rounded ................eee

Re ON ae aca chiar «va onan Bese As Uaewswnubone nnd. uaovaesaliacedeanan) A. roseus

b. Pedicel longer than flower, monocarps with apex beaked ............. 4. costatus

376

10a.

16a.

Gard. Bull. Singapore 64(2) 2012

Pedicel villose, monocarps flat-topped, generally angular ..........0.... ee 9

Pedicel glabrous or tomentose but not villose, monocarps ellipsoidal and

beaked(flat-topped in A; pandanicar pus) -eecnecc cee 1]

Leaves typically coriaceous, sepals broader than long, monocarps glabrous

siiib U4 sas wach lotic mutgegtt dt ata catroete a aeeetraat tetas eee Rect eae eae eee A. lanuginosus

Leaves typically chartaceous, sepals longer than broad, monocarps shortly

tomentose; -atleastiitear base -aeeretse cesta eee eee eect eee ee 10

Outer petals lanceolate, more than 2 cm long, monocarps with angled sides

extendingvover thewhole length ac-.cee errata eee eee A. hirtipes

Outer petals ovate, less than 2 cm long, monocarps with angled sides near

thie DASE * omy Meee see ekarerens eee eee erence ee eee A. kinabaluensis

Blades of at least one petal whorl more than 10 mm wide ..................ccesees 1D

Blades of both petal whorls less than 10 mm wide ....................ssseseeeene LA

Petal blades ovate, monocarps flat-topped and angled ....... A. pandanicarpus

Petal blades of at least one whorl lanceolate, monocarps ellipsoidal, markedly

bealkced, mot angled sco vie reson coteeecaes nee eee cena eee mene ee eee 13

Flower pedicel less than 10 mm long, monocarps to 2.5 cm long, 1 cm

diameter beak veryihatdsatte/ shamp ies. seseasssce teres ree eee ee A. polygynus

Flower pedicel more than 20 mm long, monocarps to 4 cm long, 2 cm

diameter, beakenot hard and Sinan cen costae cet eacer nee ene A. macropodus

Foliage, flowers and fruits often with yellow powdery spots, monocarps

PUSUA OLIN sec fae Aosrseageete lac cect e cm aa roee omen A. atractocarpus

Foliage, flowers and fruits not yellow spotted, monocarps sessile ................ 15

Petals densely red-brown tomentose, monocarps drying dull brown

PRE ee ener eee ero so Me te eoonbon acct shcnon con A. venustus

Petals densely to sparsely pale or brown tomentose, monocarps drying shiny

SiO OtH LO Wats oso Seesciereurere see secon stemenenoesneavoceeeusen ate sexs aces ee: eeeereee eee eee 16

Petals relatively thick and fleshy, little difference in shape of blades between

whorls; monocarps longer than DrOad/ies....cacie cater sheen eases A. maingavi

Petals relatively thin, blades of inner whorl narrower than outer whorl,

monocarps broadenth anil Om Cys screenees ee eee eee 4. ochropetalus

Climbing Annonaceae of Borneo 377

1. Artabotrys atractocarpus |.M.Turner

(Greek, atractos = spindle-shaped, karpos = fruit)

Folia Malaysiana 10 (2009) 79. TYPE: Borneo, Sarawak, Miri Division, Sungai

Tutoh, Melana Protection Forest, Bukit Pelamau, 20 April 1997, R.M.A.P. Haegens

& N. Klazenga 438 (holotype: L[<2] (barcode nos. L 0196939, L0196940); isotypes:

K, SAR).

Large woody climber. Foliage, flowers and fruits in dry specimens often with yellow

powdery spots and splashes (a feature of several Borneo Annonaceae species including

Polyalthia rumphii (Blume ex Hensch.) Merr. and Artabotrys gracilis). Twigs drying

brown, sometimes quite dark, smooth and shiny but tending to become longitudinally

wrinkled or latticed, glabrous or with a few scattered erect short pale hairs, sometimes

with small pale lenticels. Leaves chartaceous, drying brown or grey-brown above,

brown beneath typically with the midrib and lateral nerves contrasting a darker brown,

midrib more or less flesh above in dry leaves, prominent beneath, lateral nerves very

slightly raised above, prominent beneath, glabrous except for scattered pale adpressed

hairs along midrib and laterals below, often with abundant irregular minutely pimply

pustules on lower lamina, lamina ovate to elliptic, more rarely obovate, 6-17 « 2.5—7

cm, base rounded, truncate or obtuse, ultimately narrowly decurrent to petiole with

lamina edge continuing in to the raised edge of a groove along the top of the petiole,

apex acuminate, acumen rather narrow and sharp, lateral veins 10—13 pairs, looping

very distinctly within the margin, tertiary venation reticulate; petiole 4-7 mm long,

0.5—2 mm thick. Inflorescence hooks variably adpressed pale hairy, single-flowered.

Flowering pedicel 822 mm long, c. 1 mm thick, widening distally, drying brown or

blackish, shiny, longitudinally wrinkled, sometimes bumpy, sepals thick, triangular,

c. 3 x 4mm, apex acute, reflexing, base slightly connate, scattered pale hairs outside,

glabrous within, petals green, coriaceous, with clear distinction between claw and

blade, blade of outer petals lanceolate, c. 26 x 6-7 mm, more or less flat, though

sometimes with a faintly raised central ridge adaxially, drying red-brown, dense short

pale tomentum soon lost except near base of petal, often pustulated, inside of claw

drying dark brown, shiny, glabrous, inner petals coherent at margins of claw, with

linear blade c. 26 x 3 mm, incurving with tips crossing, flat or slightly grooved with

transition to claw pentagonal in cross section, drying red-brown, often pustulate,

short pale tomentose near base, stamens many, c. 2 mm long, connective flat-topped,

minutely erect hairy, carpels c. 8, c. 1.5 mm long, glabrous. Fruiting pedicel 15 mm

long, 2 mm thick, monocarps c. 6, fusiform, to 30 mm long including 10-mm beak, 13

mm across, drying black and shiny with a series of laterally flattened ridges or flanges

running longitudinally from base to apex, the two most pronounced on opposite sides

extending to form the laterally compressed beak of the fruit, between these there are

several less-pronounced longitudinal ridges, some reduced to a series of raised warts,

stipe to 15 mm long. Seeds 2.

378 Gard. Bull. Singapore 64(2) 2012

Distribution. Endemic to Borneo. Collected from eastern Sarawak and East Kalimantan.

Ecology. Lowland forest.

Notes. When I published this as a new species, the diagnosis was based on the stipitate

monocarps in comparison to the sessile monocarps of other Malesian Artabotrys

species. I have subsequently seen Nengah Wirawan 439 collected from Java that has

stipitate monocarps and appears to be A. sumatranus. Forbes 3242 from Sumatra also

has stipitate monocarps. The monocarps from these specimens have a rounded apex

and not the pronounced beak of A. atractocarpus.

Vegetatively the frequent yellow powdering leads to confusion with Artabotrys

gracilis King but the flowers of A. atractocarpus are far larger.

2. Artabotrys costatus King

(Latin, costatus = ribbed, the distinct lateral veins)

J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 61(2) (1892) 37. Ridley, Sarawak Mus. J. 1(3)

(1913) 78. Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921) 267. Masamune,

Enum. Phan. Born. (1942) 280. Sinclair, Gard. Bull. Singapore 14 (1955) 255. Beaman

et al., Pl. Mt. Kinabalu 4 (2001) 81. TYPE: Peninsular Malaysia, Perak, Ulu Bubong

500-800’, King’s Collector [H.H. Kunstler] 4297 (lectotype, designated by Turner

(2009a), K (barcode no. K000381022); isolectotypes: BM, BO, CAL, K, SING[*2]).

Large woody climber. Twigs dark brown, smooth, some red-brown hairs on youngest

parts. Leaves chartaceous, drying (pale) brown, midrib flush to slightly raised above,

prominent below, secondary veins faintly raised above, but lamina often a little bullate

giving impression of sunken veins above, sparse scattered hairs on lamina and along

midrib in youngish leaves, leaves oblong elliptic, 9-26 x 3—9 cm, base obtuse, slightly

decurrent, apex acuminate, lateral veins distinct, 11—13 pairs, petiole to 8 mm long, 2

mm thick. Inflorescence hooks recurved bearing many flowers. Pedicel to 3 cm long,

very slender, c. 0.5 mm wide when dry, more or less glabrous, sepals broadly triangular

2 mm long, 3 mm across, petals yellow, densely covered with pale tomentum, outer

petals clawed but with little distinction between claw and blade visible from outside,

blade ovate oblong, 7 x 3 mm, inner petals clawed, blade concave, 6 x 2 mm, stamens

many, carpels 3—8. Fruiting pedicel to 3 cm long, 3 mm thick, monocarps to 7 or more,

ripening red, cylindrical, 3 1.5 cm, drying black, strongly wrinkled, sessile, often

beaked. Seeds | or 2, 20 x 10 x 6 mm, pale brown, longitudinally grooved.

Distribution. Malay Peninsula, Borneo. Recorded from Brunei, Sabah and Sarawak.

Ecology. Lowland and montane forest to 1500 m.

Climbing Annonaceae of Borneo 379

3. Artabotrys gracilis King

(Latin, thin, slender)

J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 61(2) (1892) 35. Ridley, Sarawak Mus. J. 1(3)

(1913) 77. Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921) 267. Masamune,

Enum. Phan. Born. (1942) 280. Sinclair, Gard. Bull. Singapore 14 (1955) 258.

Kessler & van Heusden, Rheedea 3 (1993) 54. Bygrave in Coode et al., Checkl. FI.

Pl. Gymnosp. Brunei (1996) 12 as ‘cf. gracilis’. TYPE: Peninsular Malaysia, Perak,

King s Collector {H.H. Kunstler] 4987 (lectotype, designated by Nurainas (2004), BO;

isolectotypes: BM, CAL, K, L, SING).

Woody climber to at least 25 m long. Twigs smooth, sometimes faintly latticed, drying

brown, glabrous. Leaves chartaceous, drying brown, generally darker above, often with

powdery splashes of white or yellow reminiscent of Polyalthia rumphii, ovate-elliptic,

5—15 x 2-6 cm, base obtuse, slightly decurrent, apex acuminate, almost caudate, midrib

slightly sunken above, prominent below, lateral veins c. 8 pairs, arching forwards and

looping within the margin. Petioles 4-7 mm long, 0.5—1 mm thick. Inflorescences

hooks recurved and laterally compressed. Flowers pedicels 10—16 mm long, c. 0.5

mm thick, thickening distally, glabrous, sepals free, broadly ovate, 2 x 3 mm, petals

pink when young, yellow when older, when dry densely covered in pale adpressed

hairs, often with powdery white or yellow spots, outer whorl clawed, blade triangular,

5 x 1.5 mm, a broad-based triangle in cross section with the flat face inwards, claw

broadly ovate, c. 2 x 3 mm, adpressed pale hairy externally, glabrous and concave

within with a distinct raised upper rim, inner whorl clawed, blade terete, triquetrous or

flattened, 2-3 mm long by | mm wide, claw c. 3 x 3 mm, glabrous except for a narrow

central inverted triangle externally, glabrous within, stamens many, c. | mm long,

apex truncate, carpels 3. Fruits pedicels 10—16 mm long, c. 1 mm thick, monocarps 1

or 2, ellipsoidal, 15—22 x 10-15 mm, glabrous, drying black or brown sometimes with

many pale brown pustules, drying slightly ridged longitudinally, shortly beaked, stipe

to c. 3 mm long. Seeds 1-2, 10 x 8 x 4mm.

Distribution. Malay Peninsula, Sumatra and Borneo. In Borneo collected from Brunei,

Kalimantan, Sabah and Sarawak.

Ecology. Lowland forest to 800 m.

4. Artabotrys hirtipes Ridl.

(Latin, hirti = hairy, pes = foot, the hairy pedicel)

Bull. Misc. Inform. Kew (1912) 383. Sarawak Mus. J. 1 (1913) 77. Merrill, J. Straits

Branch Roy. Asiat. Soc. Spec. No. (1921) 267. Masamune, Enum. Phan. Born.

380 Gard. Bull. Singapore 64(2) 2012

(1942) 280. TYPE: Borneo, Sarawak, Rejang Kapit, G.D. Haviland 2326 (lectotype

designated by Turner (2009a), K (barcode no. K000691269)).

Artabotrys trichopetalus Merr., J. Straits Branch Roy. Asiat. Soc. 85 (1922) 176.

Merrill, Univ. Calif. Publ. Bot. 15 (1929) 71. Masamune, Enum. Phan. Born. (1942)

281. TYPE: Borneo, Sabah, Sandakan, September-December 1920, M. Ramos 1465

(lectotype, designated by Turner (2009a), K (barcode no. K000691311); isolectotypes:

A[2], BM, L, US).

Artabotrys cinnamomeus Diels, Notizbl. Bot. Gart. Berlin-Dahlem 11 (1931) 84.

TYPE: Borneo, Sarawak, September 1865, O. Beccari PB. 531 (holotype: FI-B;

isotype: B(fragment)).

Artabotrys sp. 1, Bygrave in Coode et al., Checkl. Fl. Pl. Gymnosp. Brunei (1996) 13.

Large woody climber. Twigs pale when older, latticed or longitudinally wrinkled,

bark rather papery, brown tomentum on the youngest twigs. Leaves chartaceous to

subcoriaceous, drying brown, generally darker above, glabrous above, scattered long

adpressed hairs along midrib beneath, midrib flush above, prominent beneath, narrowly

elliptic-obovate to oblanceolate, 10-32 = 3.5—6 cm, lateral veins c. 8 pairs, arching

well forward, petiole drying black, 6-10 mm long, 2—3 mm thick. Inflorescence hooks

recurved, bearing many flowers. Pedicel 12—15 mm long, densely rusty or pale brown

villose, sepals triangular, c. 10 mm long, 5 mm across, apex acute, villose outside,

glabrous inside, to 2.5 cm long, basal bract foliose to 3 cm long, medial bract to 2

cm long, petals relatively thin, densely covered with pale hairs, outer petals blade

lanceolate, 25 mm long, 9 mm wide, inner petals similar length, slightly narrower

c. 6 mm wide, stamens many, c. 2 mm long, carpels many, c. 1.5 mm long, densely

pale hairy. Monocarps many in large groups to 6 cm diameter, pyriform, to 4 x 2.5

cm, drying brown, angled, flat-topped, central apiculus, covered with very short, pale

tomentum. Seeds 2, 22 x 15 x 7 mm, pale brown, smooth.

Distribution. Endemic to Borneo. Collected from Brunei, Kalimantan, Sabah and

Sarawak.

Ecology. Lowland forests.

Notes. Artabotrys hirtipes is close to two other species from Borneo, A. lanuginosus

and A. kinabaluensis. Vegetatively A. hirtipes 1s closer to A. kinabaluensis which

tends to have less coriaceous leaves and less hairy foliage than A. /anuginosus. The

three species are most easily separated on the size and shape of the sepals and outer

petals. Artabotrys lanuginosus has particularly broad sepals, which unlike the other

two species are broader than long. In A. hirtipes the outer petals are lanceolate and

longer than the more ovate outer petals of A. kinabaluensis. In fruit the species can

also be distinguished. The monocarps of A. /anuginosus are glabrous whereas the other

Climbing Annonaceae of Borneo 381

two species are at least partially shortly pale tomentose. Artabotrys hirtipes and A.

lanuginosus typically have angled sides running up the length of the sessile fruitlets.

In A. kinabaluensis the base of the monocarp may have angled faces but the angles do

not extend up the length of the fruitlet which remains more or less a flattened circle in

cross section in the distal portions.

See notes under A. pandanicarpus for differences from that species.

5. Artabotrys kinabaluensis |.M.Turner

(of Mt Kinabalu)

Malayan Nat. J. 62: (2010) 360. TYPE: Borneo, Sabah, Mt Kinabalu, Tenompok,

7 March 1932, J. Clemens & M.S. Clemens 28697 (holotype: BM (barcode no.

BM000895993): isotypes: A, K, SING).

Artabotrys aff. hirtipes, Beaman et al., Pl. Mt. Kinabalu 4 (2001) 81.

Large woody climber. Twigs drying pale brown to pale grey brown, latticed with

raised pale lenticels, youngest twigs brown with adpressed long straight pale hairs,

sometimes quite dense. Leaves chartaceous, drying brown, generally darker above,

midrib sometimes a paler yellow-brown below, glabrous above, adpressed hairs along

lamina below, sometimes scattered on lower lamina, midrib in dry leaves flush above,

prominent beneath, lateral veins slightly raised above, raised below, lamina elliptic

to obovate, 7—15.5 x 3—6 cm, apex shortly acuminate, base obtuse, lateral veins 9-11

pairs, arching forward and looping distinctly within the margin; petiole 4-6 mm long,

1-2 mm thick, often drying blackish with adpressed hairs. Inflorescence axes bearing

several flowers. Flower pedicels 10-15 mm long, c. 1.5 mm thick, densely brown

hairy, sepals ovate-lanceolate, c. 9 x 6 mm, densely covered with long and straight

brown hairs, inside drying black, glabrous except near margins where hairs are finer

and sparser than externally, outer petals broadly ovate 16 x 11 mm, densely covered in

straight brown hairs except for near the base inside which is glabrous and surrounded

by a raised rim, inner petals clawed, blade broadly ovate 11 x 10 mm, densely brown

hairy, stamens many c. 2 mm long. Fruiting pedicel 12-15 mm long, c. 5 mm thick,

monocarps to 15 or more, sessile to 4 x 3 cm, ellipsoidal, slightly compressed laterally,

drying slight sunken between the seeds, apex often quite flat-topped and ultimately

apiculate, base obtuse, sometimes slightly angled with flat faces, drying black or very

dark brown, minutely warty, with scattered short brown hairs densest near the base,

much denser when young. Seeds 2, c. 3 x 1 cm.

Distribution. Endemic to Borneo where it is only known from the Mount Kinabalu

area of Sabah.

Ecology. Montane forest from 1500 to 2100 m.

382 Gard. Bull. Singapore 64(2) 2012

Notes. See notes under A. hirtipes for distinction of A. kinabaluensis from A. hirtipes

and A. lanuginosus.

6. Artabotrys lanuginosus Boer].

(Latin, woolly, downy)

Cat. Pl. Phan. (1899) 23. Boerlage, Icon. Bogor 1 (1899) 121 525(ps 159) eas

‘lanuginosa’. Ridley, Sarawak Mus. J. 1(3) (1913) 78. Merrill, J. Straits Branch Roy.

Asiat. Soc. Spec. No. (1921) 267. Masamune, Enum. Phan. Born. (1942) 280. TYPE:

Borneo, Penigin, 1896-1897, Jaheri 320 (lectotype, designated by Turner (2009a), BO

(sheet no. BO-1339062)).

Artabotrys macranthus Holth., Blumea 5 (1942) 180. Kessler & van Heusden, Rheedea

3 (1993) 54, as ‘macrantha’. TYPE: Sulawesi, Sangi and Talaud Islands, Karakelang,

Pasir Malap, E. of Lobo, 14 May 1926, H.J. Lam 3003 (holotype: L (barcode no. L

0037895); isotypes: BO, L[ <2], K).

Artabotrys sp. 2, Bygrave in Coode et al., Checkl. Fl. Pl. Gymnosp. Brunei (1996) 13.

Large woody climber. Twigs covered with dense red-brown tomentum, becoming

glabrous with age when brown, latticed, often with raised, dark lenticels giving

a rough feel. Leaves coriaceous to chartaceous, drying chestnut brown, sometimes

slightly paler beneath, midrib slightly raised to slightly sunken above in dry leaves,

prominent beneath, lateral veins flush to slightly sunken above, prominent beneath,

lamina glabrous above, hairy below, hairs long (to 2 mm), more or less straight, red-

brown becoming pale with age, scattered over lower lamina, denser and more clearly

adpressed on midrib below, lamina elliptic or ovate-elliptic, 16-21 x 7—10 cm, apex

shortly acuminate, base obtuse, lateral nerves 11-14 pairs, angled forward and arching,

looping obscurely within margin, clearly visible from both surfaces, tertiary venation

rather lax, only visible from below. Petiole drying brown, horizontally wrinkled, c. 3 =

7-10 mm, tomentose. Inflorescence hooks recured, bearing many flowers. Flowering

pedicel to 15 mm long, 2 mm thick, densely covered with long pale hairs, sepals

drying brown, broader than long, 6 x 10 mm, densely pale hairy outside, much more

sparsely hairy within, petals green becoming yellow, petal whorls similar in shape,

ovate, 25 x 15 mm, apex blunt; densely covered on both surfaces with pale brown

hairs, outer petals glabrous near base inside which is surrounded by a raised rim, inner

petals clawed with a thick, pointed rim over top of claw; stamens many, carpels many.

Fruiting pedicel not seen, monocarps ellipsoidal, slightly laterally compressed, c. 3 x 2

x 1.5 cm, sessile or subsessile, apex rather flat-topped, with angles ridges extendin the

length of the monocarp, drying dark brown with paler patches where in contact with

neighbours, glabrous, surface minutely warty and somewhat shiny. Seeds 2.

Distribution. ?Sumatra, Borneo and Sulawesi. In Borneo known from a few scattered

Climbing Annonaceae of Borneo 383

collections from Brunei, Kalimantan and Sabah. S 60776 from Limbang in Sarawak

may also belong here.

Ecology. Lowland forest.

Notes. See notes under A. hirtipes for distinction of A. /anuginosus from A. hirtipes

and A. kinabaluensis.

7. Artabotrys macropodus |.M.Turner

(Greek, makro = big, podus = footed, based; the large pedicel of the flower)

Folia Malaysiana 10 (2009) 77. TYPE: Borneo, Sabah, Ranau District, about 3 miles

NW of Kampung Pinanantai, 9 May 1973, G. Shea & Aban SAN 76878 (holotype: K

(barcode no. K000581385); isotypes: L, SAN, SING).

Large woody climber. Twigs brown, often with tiny raised lenticels, youngest parts

dark brown, shiny, more or less glabrous but for a few adpressed hairs around nodes.

Leaves chartaceous to coriaceous, drying various shades of brown, main nerves more

consistent dark brown above and below, contrasting with lamina when pale, midrib in

dry leaf flush to slightly raised above, prominent below, lateral nerves slightly raised

above, prominent below, leaves more or less glabrous except sometimes for some sparse

pale long adpressed hairs along midrib below, lamina oblong-ovate or oblong-elliptic,

more rarely oblong-obovate, 10-31 =x 4—13 cm, base acute, obtuse or rounded, apex

shortly acuminate, lateral nerves 8—14 pairs, looping distinctly within margin, tertiary

venation reticulate, distinct from both surfaces in dry leaves. Petioles 7-12 mm long,

2—5 mm thick. Inflorescence hooked, single-flowered. Flower pedicel 2—3 cm long,

0.5—1.0 mm thick, pale adpressed hairs, sepals triangular 5—8 x 4-5 mm, apex acute,

externally with long pale hairs, inside glabrous, outer petals thick, coriaceous, ovate

with only slight constriction between claw and blade, 20-22 < 10-13 mm, densely

adpressed pale hairy externally, particularly near the base, inside similarly tomentose

except for the excavation at the base which is glabrous, inner petals cohering at their

margins with gap between, blades thick, flat, ovate-lanceolate, 10-14 x 5—10 mm,

densely pale hairy on both surfaces, stamens many, c. 2 mm long, connective apex

convex, carpels many. Fruiting pedicel 2.5—-3.5 cm long, 2—5 mm thick, monocarps

5—20 or more, ripening red, ellipsoidal c. 4 x 2 cm, drying brown, apparently markedly

wrinkled if collected fully ripe, more or less glabrous except for a few short hairs

near base, unripe fruit drying smooth with faint longitudinal lines, surface minutely

warty under magnification, apex acuminate with a distinct beak, subsessile, stipe c.

3 mm long. Seeds 2, ellipsoidal with one flattened face, c. 26 x 14 x 8 mm, with a

longitudinal groove.

Distribution. Endemic to Borneo. In Borneo collected from Kalimantan, Sabah and

Sarawak.

384 Gard. Bull. Singapore 64(2) 2012

Ecology. Lowland forest.

Notes. Among Bornean species 4. macropodus is not easily confused. Artabotrys

costatus King has similar long pedicels, but they are more slender and the petals are

much narrower and the monocarps are less broad. Artabotrys polygynus has broad

petals and numerous monocarps but the leaves dry a characteristic very dark brown, the

petals are thicker but not so broad as 4A. macropodus and the monocarps are narrower

and have a characteristic very hard, sharp point when dry.

8. Artabotrys maingayi Hook.f. & Thomson

(Alexander C. Maingay (1836-1869), English doctor and prison administrator in

colonial service)

FI. Brit. India 1 (1872) 55. Sinclair, Gard. Bull. Singapore 14 (1955) 259. Turner, Folia

Malaysiana 10 (2009) 62. TYPE: Peninsular Malaysia, Malacca, 1867-1868, A.C.

Maingay 2617 [Kew distrib. no. 34] (holotype: K[x2] (barcode nos. K000381024,

K000381029); isotypes: BM, CGE).

Artabotrys havilandii Ridl., Bull. Misc. Inform. Kew 1912 (1912) 382. Ridley, Sarawak

Mus. J. 1(3) (1913) 77. Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921)

267. Masamune, Enum. Phan. Born. (1942) 280. Sinclair, Sarawak Mus. J. 5 (1951)

598. Bygrave in Coode et al., Checkl. Fl. Pl. Gymnosp. Brunei (1996) 12. TYPE:

Borneo, Sarawak, near Kuching, 9 September 1892, G.D. Haviland 1629 (lectotype,

designated by Turner (2009a), K (barcode no. K000691275); isolectotype: SAR).

Large woody climber. Twigs drying dark and smooth, more or less glabrous. Leaves

chartaceous to thinly coriaceous, drying grey-brown, ovate-elliptic to oblong ovate,

7-13 =< 3-5 cm, base obtuse to rounded, slightly decurrent, apex acuminate, midrib

slightly raised above, prominent beneath. Inflorescence hooks recurved, drying black,

glabrous. Pedicel 7-10 mm long, drying black, longitudinal wrinkled, some scattered

pale hairs, sepals ovate, 4 x 4 mm, scattering of pale hairs outside, petals relatively

thick and fleshy, densely covered with dense, pale adpressed hairs, outer petals ovate

lanceolate, 20 x 6-7 mm, blade 15 x 6—7 mm, inner petals clawed, blades shaped like

rabbits’ ears, 15 « 4-5 mm, stamens many, carpels many. Fruiting pedicel c. 10 x 4

mm, monocarps 2-5, ellipsoidal to 4.5 = 3.5 cm, ripening orange-yellow, drying black,

glabrous, thick-walled (to 4 mm), apex beaked, subsessile, stipe c. 3 mm long, 5 mm

thick. Seeds 2, plano-convex, c. 2.5 x 1.5 x 1 cm.

Distribution. Malay Peninsula and Borneo. In Borneo from Brunei and Sarawak.

Earlier reports of the species from Kalimantan (Turner 2009a), were based on

misidentification of A. veldkampii specimens.

Ecology. Lowland forest.

Climbing Annonaceae of Borneo 385

Notes. Refer to A. veldkampii for notes on confusion of fruiting specimens with that

species.

9. Artabotrys ochropetalus |.M.Turner

(Greek, ochros = pale yellow, petalon = petal)

Folia Malaysiana 10 (2009) 63. TYPE: Borneo, Sabah, Tawau Division, Lahad Datu

District, Ulu Sungai Segama, Danum Valley Research Centre, 28 February 1985,

Argent et al. SAN 108284 (holotype: K (barcode no. K000581905); isotypes: K, KEP,

L, SAN).

Artabotrys roseus auct. non Boerl., Merrill, Univ. Calif. Publ. Bot. 15 (1929) 71.

Kessler & van Heusden, Rheedea 3 (1993) 55.

Large woody climber. Twigs drying black or brown, smooth, more or less glabrous,

sometimes with sparse pale brown adpressed hairs on youngest parts. Leaves

chartaceous, drying brown, generally darker below, midrib flush to slightly raised

above, prominent beneath, lamina elliptic to elliptic lanceolate, 7-20 x 2.5—6.5 cm, base

acute to obtuse, slightly decurrent, apex acuminate, lateral veins 10—13 pairs, looping

clearly within margin; petiole 10 mm long, 2 mm thick. Inflorescence hooks recurved,

multi-flowered. Pedicels 6-11 mm long, with pale adpressed hairs, sepals triangular,

apex acute, 3 x 3 mm, with sparse pale hairs; petals yellow, drying brown with short

pale pubescence, outer petals with slight constriction between claw and blade, blade

lanceolate 21-30 x 4-5 mm, pale pubescent to glabrescent on both surfaces, claw

c. 2 mm long, 3 mm wide, pubescent externally, excavated, glabrous within; inner

petals with blade 3—S mm wide with a central ridge adaxially, pale pubescent on both

surfaces, claw c. 2 x 2 mm, slight ridge externally, pubescent externally, excavated,

glabrous within, stamens many, c. 1.5 mm long, apex truncate, carpels c. 6, ovoid,

glabrous but with long hairs among carpels. Fruiting pedicel c. 10 mm long, 6 mm

thick, monocarps to 6, ripening yellow, ellipsoidal, wider than long, to 3 x 4.5 = 2.5

cm, longitudinal ridges, glabrous, drying brown, beaked. Seeds 2, 14 « 10 x 8 mm,

smooth, brown, pale hilum.

Distribution. Endemic to Borneo where it has been collected from Brunei, Kalimantan,

Sabah and Sarawak, but is particularly commonly collected in Sabah and East

Kalimantan.

Ecology. Lowland forest.

Notes. The name Artabotrys roseus was widely misapplied to this species which is one

of the more commonly collected taxa.

386 Gard. Bull. Singapore 64(2) 2012

10. Artabotrys pandanicarpus |.M.Turner

(Greek, carpos = fruit, the fruits like those of Pandanus)

Folia Malaysiana 10 (2009) 83. TYPE: Borneo, Sarawak, near Kuching, 7 October

1892, G. D. Haviland 1775 [Garai leg.| (holotype: K (barcode no. K000581386)).

Artabotrys blumei auct. non. Hook.f. & Thomson, Ridley, Sarawak Mus. J. 1(3) (1913)

78. Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921) 266. Masamune, Enum.

Phan. Born. (1942) 279.

Large woody climber. Twigs drying brown with youngest parts almost black, occasional

pale hairs soon lost. Leaves thinly coriaceous, drying dark brown and shiny above,

brown and dull beneath, midrib flush to slightly raised above, prominent beneath,

lateral nerves flush to slightly raised above, raised beneath, leaf glabrous except for

a few adpressed pale hairs on midrib below, lamina ovate or elliptic, 9-14.5 « 4—7

cm, base obtuse to rounded, ultimately slightly decurrent to petiole, apex acuminate,

lateral veins, 10-12 pairs, looping within margin, tertiary venation areolate; petiole

6-7 mm long, 1-2 mm thick. Inflorescence hooks, recurved, laterally compressed,

adpressed pale hairy when young, glabrous with age. Flowering pedicel unknown

(no specimens with attached flowers available), sepals relatively thin, triangular, c.

6 x 5 mm, apex acute, base slightly connate, densely adpressed pale hairy externally,

glabrous within, petals thick and coriaceous, clawed, outer petals with ovate blade, c.

18 x 13 mm, flat, apex blunt, densely covered with very short pale tomentum, claw

c. 4 x 5 mm, excavated, glabrous within with raised rim around excavation, blade of

inner petals ovate to almost sagittate, c. 14 x 12 mm, apex rounded, densely short

pale tomentose on both surfaces, claw c. 5 x 3 mm, glabrous within (reproductive

structures unknown). Monocarps many, at least 10, sessile, ellipsoidal, to 3.5 « 3 cm,

angular with generally 4-5 longitudinal ridges, apex with a distinct 4—S-sided flat-

bottomed depression, thick-walled (3—S5 mm), drying dark brown, smooth, covered

with dense pale, very short, tomentum. Seeds, 1, c. 2.5 * 1.5 * 1 cm.

Distribution. Endemic to Borneo, where it is known from two or three collections from

Sarawak (type and a Haviland collection with the same number which may or may not

be a true duplicate from Kuching, S 28025 from Bintulu).

Ecology. Lowland forest.

Notes. The species of the A. hirtipes group (see notes under A. hirtipes) have sessile,

angular, flat-topped monocarps similar to those of A. pandanicarpus. However the A.

hirtipes group members have leaves with much more distinct secondary venation and

floral parts covered with long hairs and relatively thin petals, whereas 4. pandanicarpus

has short to very short hairs on the flowers and thick fleshy petals.

Climbing Annonaceae of Borneo 387

11. Artabotrys polygynus Mig.

(Greek, poly = many, gyne = female; the many carpels)

Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 41. Ridley, Sarawak Mus. J. 1(3) (1913) 77.

Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921) 267. Masamune, Enum.

Phan. Born. (1942) 280 as ‘polygonus’. Bygrave in Coode et al., Checkl. FI. PI.

Gymnosp. Brunei (1996) 12, as ‘cf. polygynus’. Beaman et al., Pl. Mt. Kinabalu 4

(2001) 82. TYPE: Borneo, Mt Pamatton, P W. Korthals s.n. (lectotype, designated by

Turner 2011, U (barcode no. U0000239)).

Artabotrys borneensis Merr., J. Straits Branch Roy. Astat. Soc. 85 (1922) 175. Merrill,

Univ. Calif. Publ. Bot. 15 (1929) 71. Masamune, Enum. Phan. Born. (1942) 280.

Bygrave in Coode et al., Checkl. Fl. Pl. Gymnosp. Brunei (1996) 12. TYPE: Borneo,

Sabah, Sandakan, September—December 1920, VM. Ramos 1366 (lectotype, designated

by Turner (2009a), BM (barcode no. BM000898081); isolectotypes: A, B, BO, K, L,

PAU):

Artabotrys hexagonolobus Priyanti, Floribunda 2(6) (2004) 161. TYPE: Borneo,

Kalimantan, East Kalimantan, Berau, 9 October 1997, P.J.A. Kessler et al. B867

(holotype: BO; isotypes: K, L, WAN).

Large woody climber. Twigs drying dark brown or black sometimes paler brown, often

noticeably latticed, rusty adpressed tomentose on young parts, sometimes glabrescent.

Leaves chartaceous, drying dark brown often greyish above, midrib flush to slightly

raised above, prominent below, sometimes with pale, relatively long adpressed hairs

along midrib and lamina beneath, otherwise glabrous, elliptic to obovate, base acute

to obtuse, decurrent, apex apiculate to acuminate, 5—16 x 2—6 cm, lateral veins visible

from both sides in dry leaves, 8—9 pairs, petiole 3-5 mm long. Inflorescence hooks

recurved, single-flowered. Pedicel 7-8 mm long, dilating distally, covered with red-

brown hairs, densely brown hairy, sepals triangular 5-8 mm long, 5—6 mm across,

apex acute; petals yellowish, drying dark brown with pale adpressed hairs sparser on

blades, outer petals thick fleshy, clawed, blade lanceolate 30 mm long, 12-15 mm

wide, inner petals, blade ovate lanceolate, 20 mm long, 8-10 mm wide, stamens many,

2-3 mm long, carpels many. Monocarps 8 or more, irregularly cylindrical, spindle-

shaped, sessile, apex sharply beaked, 2—2.5 cm long, | cm diameter, drying black or

brown. Seeds 2, 15 x 7 x 4 mm, brown with longitudinal groove.

Distribution. Endemic to Borneo. In Borneo it has been recorded from Brunei,

Kalimantan, Sabah and Sarawak.

Ecology. Lowland and hili forest to 1500 m.

388 Gard. Bull. Singapore 64(2) 2012

12. Artabotrys roseus Boer].

(Latin, rosy, pink)

Icon. Bogor. | (1899) 122, t. 53 (p. 161), as ‘rosea’. Ridley, Sarawak Mus. J. 1(3)

(1913) 77. Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921) 267. Merrill,

Univ. Calif. Publ. Bot. 15 (1929) 71. Masamune, Enum. Phan. Born. (1942) 280.

Kessler & van Heusden, Rheedea 3 (1993) 55. TYPE: Borneo, Sarawak, near Kuching,

12 October 1894, G.D. Haviland s.n. (holotype: BO (sheet no. BO-1344062).

Artabotrys pleianthus Diels, Notizbl. Bot. Gart. Berlin-Dahlem 11 (1931) 84. TYPE:

Sarawak, September 1865, O. Beccari PB. 554 (holotype: FI-B; isotypes: B, K, M).

Large woody climber. Twigs drying brown, grey or blackish, latticed, often with raised

lenticels giving a very rough feel, short brown tomentum on youngest parts. Leaves

chartaceous to coriaceous, often drying grey-green or yellowy brown with yellowish

venation, midrib distinctly raised above, prominent beneath, more or less glabrous,

ovate elliptic to obovate, 2.5—12 « 1.2-4.5 cm, base obtuse to rounded, lamina slightly

decurrent, apex apiculate to acuminate, lateral veins 9-10 pairs, clearly visible from both

surfaces in dry leaves. Inflorescence hooks recurved, laterally compressed, sometimes

with adpressed brown hairs, many-flowered. Pedicel 8-20 mm long, tomentose; sepals

broadly ovate, 4 mm long by 3 mm wide, apex acute, slightly connate at base, outer

petals triangular with little external distinction between blade and claw, 15 mm long,

4 mm wide, densely short pale or brown hairy on both surfaces except glabrous basal

excavation inside; inner petals clawed, blade linear c. 1 mm wide, densely tomentose

on both surfaces; stamens many, carpels c. 15. Fruiting pedicel to 2 cm long, drying

2 mm thick, monocarps to 8 or more, globose to ellipsoidal, to 3.5 x 2 cm, apex

rounded not beaked, drying black or brown, smooth, glabrous, not sessile, stipe short

but distinct, 2-6 mm long, 3—4 mm thick. Seeds 2, c. 19 x 13 x 8 mm.

Distribution. Endemic to Borneo. In Borneo collected from Brunei, Kalimantan,

Sabah and Sarawak.

Ecology. Lowland forest including heath forest.

Notes. Similar to A. havilandii but A. roseus can be distinguished by its thinner petals,

linear inner petals and unconstricted outer ones.

13. Artabotrys sarawakensis |.M.Turner

(of Sarawak)

Folia Malaysiana 10 (2009) 70. TYPE: Borneo, Sarawak, Baram District, 26 October

1894, C. Hose 302 (holotype: K (barcode no. K000581384); isotypes: BM, CGE).

Climbing Annonaceae of Borneo 389

Large woody climber. Twigs brown to dark brown, smooth or with faint longitudinal

wrinkling or latticing. Leaves drying brown, often dark brown, rather shiny above,

glabrous, midrib and lateral nerves raised on both surfaces in dry leaves, lamina

elliptic to oblong elliptic, 6.5—-11.5 x 2.5-4.5 cm, base acute to obtuse, decurrent,

apex acuminate, venation distinct, particularly from above, lateral nerves 8—9 pairs,

looping distinctly within margin. Inflorescences several flowered. Flower pedicel 17—

23 mm long, c. 0.5 mm thick, drying black, longitudinally wrinkled, glabrous, sepals

triangular 3 x 3 mm, petals yellow, clawed, densely covered in pale straw-coloured

hairs on all surfaces including very short tomentum on interior excavations, blades of

two whorls similar, triquetrous, sometimes grooved adaxially, 12-15 mm long, | mm

across, stamens c. 20, c. | mm long, carpels c. 3, glabrous. Fruits unknown.

Distribution. Endemic to Borneo where it is known from Brunei and Sarawak.

Ecology. Lowland forest including heath forest.

Notes. Similar in flower form to Artabotrys suaveolens in having the blades of both

petal whorls relatively narrow and not flat in cross section but the flowers are distinctly

larger than A. suaveolens and the pedicels are much longer. The ‘corniculatus’ form

of A. suaveolens that occurs in Sabah (described by Merrill as Artabotrys trigynus)

has larger flowers and longer pedicels than typical A. suaveolens but it also has large,

thick sepals not found in 4. sarawakensis. Also the petals of A. sarawakensis are

more densely and regularly tomentose — in 4. swaveolens the petal blades are typically

glabrescent giving a brown appearance in dried material.

Artabotrys costatatus has flowers with long pedicels, but the outer petals have

flat blades and the strongly ribbed leaves are unlike those of A. sarawakensis.

14. Artabotrys suaveolens (Blume) Blume

(Latin, fragrant, sweetly smelling)

Fl. Javae Anonaceae (1830) 62, t. 30. Ridley, Sarawak Mus. J. 1(3) (1913) 76, Merrill,

J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921) 267. Merrill, Enum. Philipp. FI.

Pl. 2 (1923) 173. Merrill, Univ. Calif. Publ. Bot. 15 (1929) 67. Airy Shaw, Bull. Misc.

Inform. Kew 1939 (1939) 289. Masamune, Enum. Phan. Born. (1942) 280. Sinclair,

Gard. Bull. Singapore 14 (1955) 256. Kessler & van Heusden, Rheedea 3 (1993) 55.

Bygrave in Coode et al., Checkl. Fl. Pl. Gymnosp. Brunei (1996) 12. Beaman et al., PI.

Mt. Kinabalu 4 (2001) 82. Unona suaveolens Blume, Bijdr. (1825) 17. TYPE: Java, in

sylvis montium Salak, Seribu etc

Unona corniculata Blanco, FI. Filip. (1837) 469. — Artabotrys corniculatus (Blanco)

Merr., Sp. Blancoanae (1918) 150 as ‘corniculata’. TYPE: Philippines, Luzon, Laguna

Province, 15 March 1913, M. Ramos Species Blancoanae No. 298 (neotype, designated

by Turner (2009a), US; isoneotypes: BM, K, L).

390 Gard. Bull. Singapore 64(2) 2012

Artabotrys parviflorus Migq., Fl. Ned. Ind., Eertse Bijv. (1861) 375. — Artabotrys

suaveolens var. parviflorus (Mig.) Mig., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 43.

TYPE: Sumatra, Lampong Privince, near Kebang, Anon. s.n. [Herb. Bogor. no. 4317]

(lectotype, designated by Turner (2011b), U (barcode no. U 0000238)).

Artabotrys rolfei S. Vidal, Rev. Pl. Vasc. Filip. (1886) 39. TYPE: Philippines, Luzon,

Prov. Albay, 1841, H. Cuming 1099 (lectotype, designated by Turner (2009a), K

(barcode no. K000691330); isolectotypes: BM, CGE, MA, MEL).

Artabotrys monogynus Merr., Philipp. J. Sci., C. 14 (1919) 383. Merrill, Enum. Philipp.

Fl. Pl. 2 (1923) 173. TYPE: Philippines, Luzon, Zambales Province, Mt Canaynayan,

Castillejos, December 1916, G. Edano Bur. Sci. 26826 (lectotype, designated by

Turner (2009a), K (barcode no. K000691328); isolectotypes: A, US).

Artabotrys trigynus Merr., J. Straits Branch Roy. Asiat. Soc. 85 (1922) 177. TYPE:

Borneo, Sabah, Sandakan, September-December 1920, 1 Ramos 1178 (lectotype,

designated by Turner (2009a), K; isolectotypes: A, BM, GH, L, P, US).

Woody climber to 25 m long of more. Twigs smooth, drying brown, typically glabrous

with a few scattered pale hairs on youngest parts, but some collections have dense

brown tomentum on the twigs. Leaves chartaceous to coriaceous, drying light brown,

sometimes greenish, ovate-elliptic, 6-12 x 2.5—5 cm, apex acuminate, base obtuse,

slightly decurrent; midrib slightly raised above particularly near the base where the

broad, flat-topped midrib is just proud of the lamina in the dry leaf, prominent beneath,

generally glabrous, sometimes with adpressed hairs on lamina and along midrib below,

lateral veins 9-12 pairs, looping well within the margin, easier to see from above.

Petioles 3—S mm long, less than | mm thick when dry. Inflorescence hooks recurved,

laterally compressed, side branches to 20 mm long. Flowers sweet smelling, pedicels

5—8 mm, widening distally, glabrous or brown tomentose, basal bracts lanceolate to

4 mm long; sepals free, triangular 1-3 mm wide by 1-4 mm long, apex acute; petals

pink or red turning pale creamy yellow, clawed, blades filiform (terete, triquetrous or

channelled), 8-11 mm long, less than | mm wide, claw covered with short dense pale

adpressed hairs, blade less densely tomentose, stamens many, truncate, c. | mm long;

carpels c. 7. Fruits pedicels S—8 mm long, monocarps | or 2, ellipsoidal, 12—15 x 7—9

mm, smooth, drying dark brown or black, unbeaked, sessile. Seeds 1—2, ellipsoidal, 10

x 6 x 5 mm, one face flattened if seeds paired, longitudinally grooved.

Distribution. Indochina to the Philippines. In Borneo widespread and apparently

common as indicated by the many collections, particularly from Sabah.

Ecology. Lowland and montane forests up to 1220 m, including heath, peat swamp and

limestone forests.

Notes. As 1s to be expected from a wide-ranging species, 4. suaveolens shows some

Climbing Annonaceae of Borneo 39]

variation in form. A few collections from Sabah were described as A. trigynus. They

are much the same as 4. corniculatus (Blanco) Merr. from the Philippines, which in

turn may simply represent the upper end of the flower size range for A. suaveolens. I

have reduced much of this variation to synonymy of A. suaveolens, but do recognise

a set of distinctive collections from Borneo as 4. sarawakensis. See this for notes on

how it differs from 4. suaveolens.

15. Artabotrys sumatranus Mig.

(of Sumatra)

Fl. Ned. Ind., Eertse Biv. (1861) 375. Ridley, Sarawak Mus. J. 1(3) (1913) 77. Merrill,

J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921) 267. Masamune, Enum. Phan.

Born. (1942) 281. Kessler & van Heusden, Rheedea 3 (1993) 57. TYPE: Sumatra,

Lampongs, near Tiga-nennin, s. dat., JE. Teijsmann s.n. [Herb. Bogor. 4382] (holotype:

U (barcode no. U 0000242)).

Large woody climbers. Twigs drying brown or dark brown with fine longitudinal

wrinkles or latticing, sparse pale or brown hairs. Leaves chartaceous, drying rather pale

brown or grey-brown, glabrous except for spare long adpressed hairs on midrib below,

midrib more or less flush above, prominent beneath, lateral nerves very slightly raised

on both surfaces, lamina elliptic to obovate, 7.5—9 = 2.5—3.5 cm, apex acuminate, base

acute, lateral nerves 10—12 pairs, looping within margin; petiole 4-5 mm long, c. 1

mm thick. Inflorescence hooks bearing several short-branched axes densely packed

with many flowers. Flower pedicel 2-4 mm long, c. 0.5 mm thick, drying dark brown

with sparse red-brown hairs, sepals ovate, 3 x 2.5 mm, drying dark brown, minutely

wrinkled and warty outside, inside smoother, petals yellow, drying brown, outer petals

clawed, to 15 mm long, blade linear lanceolate, 11 x 2 mm, red-brown hairy on both

surfaces, claw broadly ovate, 2.5—3 = 3.5 mm, red-brown hairy externally, internally

with a slightly raised upper rim, more or less glabrous, verrucose, inner petals similar

length to outer petals, blade linear, chanelled in upper portion, 11-12 mm long, c.

1 mm wide, red-brown hairy, claw c. 3 x 2 mm, hairy outside, glabrous, verrucose

within, rather thick in upper portion, blade with peltate attachment to claw, stamens

many, c. | mm long, apex acute, carpels 5—8, c. | mm long. Fruits unknown.

Distribution. Sumatra, Java and Borneo. In Borneo known with certainty only from

Kalimantan with three unnumbered collections made by P.W. Korthals and Kostermans

6820.

Ecology. Lowland forest.

Notes. As noted under A. atractocarpus, it seems that A. sumatranus produces

stipitate, but not fusiform, monocarps. I refrain from including these in the description

as no fruiting specimens are known from Borneo. While the flowering specimens

392 Gard. Bull. Singapore 64(2) 2012

from Borneo seem a good match to the type of A. swmatranus from (not surprisingly)

Sumatra, I have not had access to enough specimens to confirm that the fruiting

specimens are truly A. swmatranus.

16. Artabotrys veldkampii |.M.Turner

(J.F. Veldkamp (1941—), Dutch botanist)

Folia Malaysiana 10 (2009) 73. TYPE: Borneo, Central Kalimantan, Bukit Raya,

Batu Badinging, KCT 47 km, 7 February 1983, JF. Veldkamp 8567 (holotype: L[2]

(barcode nos. L 0180936, L 0180937); isotype: US).

Large woody climber. Twigs drying brown to black, generally smooth with some

shallow latticing, more or less glabrous except for a few pale hairs on young twigs

near nodes. Leaves chartaceous to pergamentaceous, drying brown or grey-brown

with midrib and laterals below a darker shade, midrib and laterals above slightly

raised, midrib prominent below, laterals slightly raised, glabrous except for adpressed

pale or brown hairs on midrib below, scattered on lateral nerves, lamina elliptic or

oblong-elliptic, 9.5—17 x 3.5—6.5 cm, apex acuminate, base obtuse ultimately slightly

decurrent, lateral nerves 11—13 pairs, looping distinctly within the margin, tertiary

venation reticulate, distinct form both surfaces in dry leaves; petioles 10-12 mm long,

1.5—2 mm thick. Inflorescence hooks with adpressed pale or brown hairs, flowering

repeatedly, distichously, from persistent, short straight side branches to 10 mm

long. Flower pedicel 5-6 mm, | mm thick, widening distally, drying black, shiny,

longitudinally wrinkled, with sparse brown hairs, sepals fleshy, ovate c. 2 x 2.5 mm,

drying black, wrinkled outside with scattered hairs, inside smoother and glabrous,

outer petals clawed, very narrowly ovate-lanceolate, blade 12—21 « 1—-1.5 mm, broadly

triquetrous, drying black with adpressed pale tomentum, densest towards base, claw c.

2 x 3 mm, excavation glabrous within, outside with adpressed pale tomentum, inner

petals more distinctly filiform than outer, blade 13—25 = 1 mm, terete or quadrangular

in cross section, drying black with adpressed pale tomentum, claw c. 3 x 2 mm, with

two longitudinal ridges externally, dense very short pale tomentum outside, inside

centrally glabrous with short pale tomentum marginally, stamens many c. | mm long,

apex of connective flat-topped, carpels c. 5, c. | mm long, glabrous. Fruiting pedicel

8-10 mm long, to 5 mm thick, monocarps 1-2, globose but flattened apically and

laterally compressed, to 3 cm from base to apex, to 3.5 cm at greatest width and 2.5

cm perpendicular to that, so wider then long, thick-walled, drying black, glabrous and

relatively smooth to the naked eye, sometimes with slight longitudinal ridges, under

magnification the surface appears black, shiny and wrinkled. Seeds 2, more or less

hemispherical c. 17 mm diameter, 12 mm high, with circumferential groove, drying

dull brown.

Climbing Annonaceae of Borneo 393

Distribution. Endemic to Borneo where it has been collected from Kalimantan, Sabah

and Sarawak.

Ecology. Lowland forest.

Notes. Artabotrys veldkampii has filiform petals like A. suaveolens but they are longer

and slightly wider. The fruits are, however, much larger than the pea-sized monocarps

of A. suaveolens. Fruiting A. veldkampii is likely to be confused with A. ochropetalus,

but the monocarps of A. veldkampii dry black, shiny and minutely wrinkled and

truly sessile whereas A. ochropetalus fruits dry brown, not shiny, and smoother and

generally not truly sessile. The fruits of A. maingayi are also very similar, drying black,

but they are typically longer than wide and have a more pronounced apex and stalk

than A. veldkampii. The slender petals of A. veldkampii readily distinguish it from A.

ochropetalus and A. maingayi.

A group of specimens from Sabah may belong here (SAN 41074, SAN 41075,

SAN 147972) —they have more distinct lateral veins than A. veldkampii generally does,

and the petals are broader.

17. Artabotrys venustus King

(Latin, beautiful, graceful)

J. Asiat. Soc. Bengai, Pt. 2, Nat. Hist. 61(2) (1892) 32. Sinclair, Gard. Bull. Singapore

14 (1955) 252. TYPE: Peninsular Malaysia, Perak, August 1884, Kings Collector

[H.H. Kunstler] 6499 (lectotype, designated by Nurainas (2004), BO; isotypes CAL,

K, L, SING).

Large woody climber. Twigs dark, longitudinally wrinkled, drying pale yellow-

brown when young. Leaves drying grey olivaceous, midrib slightly prominent above,

glabrous, elliptic, 9-15.5 x 4—7 cm, base obtuse to acute, apex acuminate, lateral veins

c. 10 pairs, petioles 5-7 mm long. Inflorescence hooks tightly coiled, few flowered.

Pedicel 10-25 mm long, | mm thick, sepals refiexing at anthesis, triangular, 3—5 mm

long, 2—3 mm wide, sparse adpressed red-brown hair, petals yellow, coriaceous, dense

rusty tomentose, outer petals flat, with no constriction externally at claw, blade ovate

c. 20 x 7 mm, inner similar length to outer, slightly narrower, stamens many, carpels

c. 10. Monocarps ellipsoidal, 4.5 x 2—3.5 cm, drying brown, prominently and sharply

beaked, more or less sessile. Seeds 2, c. 2.5 * 1.3 cm.

Distribution. Malay Peninsula, Sumatra, Borneo. In Borneo only known from a few

collections from Sarawak.

Ecology. Lowland forest.

394 Gard. Bull. Singapore 64(2) 2012

DESMOS Lout.

(Greek, desmos = a bond, halter or fetter;

the moniliform monocarps are reminiscent of a chain)

Fl. Cochinch. (1790) 352 as ‘Desmis’. Sinclair, Gard. Bull. Singapore 14 (1955)

261. Ban, Bot. Zhurn. 59 (1974) 1766. Turner, Folia malaysiana 10 (2009) 59.

Unona [unranked] Desmos (Lour.) Dunal, Monogr. Anonac. (1817) 97, 110. Unona

section Desmos (Lour.) DC., Syst. Nat. 1 (1817) 485, 493. LECTOTYPE: Desmos

cochinchinensis Lour. (designated by Safford 1912).

Woody climbers or scandent shrubs. Twigs glabrous or hairy. Leaves chartaceous to

coriaceous, often with paired glands in lamina margin on each side of the petiole.

Inflorescences 1- or 2-flowered, leaf-opposed or (supra-)axillary. Flowers bisexual,

sepals 3, valvate more-or-less connate at the base, petals valvate in two whorls of 3,

subequal, stamens numerous, connective truncate, carpels numerous, ovules several,

jateral. Fruits monocarps many, moniliform. Seeds few to many.

Distribution and diversity. 25—30 species from India and Sri Lanka to south China and

the Philippines. Four species recorded from Borneo.

Notes. Specimens are quite readily identified as belonging to Desmos, particularly

if it is considered in the narrow sense with Dasymaschalon as a separate genus. The

flower form with relatively undifferentiated petal whorls of quite large, free, petals;

and the moniliform monocarps, easily distinguish Desmos from other genera. Sterile

specimens might be confused with Friesodielsia, as both have paired glands in the

leaves, though these are less distinct in herbarium material in the Desmos species

compared to most species of Friesodielsia.

Key to Desmos species

la. Leaves withveins obscure snot @laucOusic ..c.cres:oscsasesstnr sce Chee hae eee ee

b; Leaves with veins distinct; ‘often elaucous.cpeeeosss scene nee eee

2a. Pedicels more than 3 cm long; opposite leaves .........<.21.2....- +0660 MaGUMnS

b. _Pedicels less tham 2.cmilongevinileat axalls: a5c1kss.ctsmcwa een cape nie mae meant

Sac 6 eaves ClaDrous:atssc seve teocccees Seca cc ue tae eee eect See aeee eee eee D. chinensis

b. Leaves tomentose: ssxics.ceoseccces secant ccasertae anne eta eee a et

1. Desmos acutus (Teijsm. & Binn.) 1.M.Turner

(Latin, acute, pointed)

Malayan Nat. J. 62 (2010) 368. — Uvaria acuta Teijsm. & Binn., Natuurk. Tijdschr.

Climbing Annonaceae of Borneo 395

Ned.-Indié 4: (1853) 398. — Unona acuta (Teijsm. & Binn.) Zoll., Linnaea 29 (1858)

320. — Habzelia acuta (Teijsm. & Binn.) Migq., Fl. Ned. Ind. 1 (1858) 37. TYPE: Java,

Anon. s.n. (lectotype, designated by Turner (2010a), L (barcode no. L 0186510)).

Desmos teysmannii (Boerl.) Merr., Philipp. J. Sci., C. 10 (1915) 235. Merrill, J. Straits

Branch Roy. Asiat. Soc. Spec. No. (1921) 256. Masamune, Enum. Phan. Born. (1942)

282. Sinclair, Gard. Bull. Singapore 14 (1955) 264. Bygrave in Coode et al., Check.

Fl. Pl. Gymnosp. Brunei (1996) 14. Beaman et al., Pl. Mt. Kinabalu 4 (2001) 83. —

Unona teysmannii Boerl., Icon. Bogor. 1 (1899) 103. Ridley, Sarawak Mus. J. 1(3)

(1913) 79. TYPE: Borneo, Sungai Landak, JE. Teijsmann s.n. [Bogor distribution no.

185] (holotype: L (barcode no. L 0037925); isotypes: B, K, US).

Scandent shrub or large woody climber to 15 m long at least. Twigs slender, dark,

lenticellate, minutely pubescent when young. Leaves membranous, glabrous except

for some hairs on midrib beneath, sometimes glaucous beneath, drying pale brown

above, ovate or oblong-elliptic to oblong-lanceolate, S—13 x 2—6 cm, base rounded to

acute, apex acute; midrib immersed above, prominent below, lateral veins 11—13 pairs,

very fine and faint on both surfaces, tertiary venation reticulate. Petioles S—7 mm long.

Inflorescences opposite leaves, single-flowered. Flowers pendent, fragrant, pedicels

long and slender, 3—6 cm, less than 0.5 mm thick; sepals triangular, 5-10 mm long,

apex acute, reflexed at anthesis; petals greenish-yellow, thin, puberulous, lanceolate,

4-7 cm long, 10-18 mm wide at widest, apex obtuse; stamens many; carpels many.

Fruits pedicels 3-6 cm long, monocarps moniliform, to 20 or more, to 5 cm long,

stipe 1-2 cm long. Seeds 2-6, globose or ellopsoidal, c. 7 x 6 mm, drying pale brown,

smooth.

Distribution. Malay Peninsula, Java, ?Bali and Borneo. In Borneo from Kalimantan,

Sabah and Sarawak.

Ecology. Lowland forest.

Notes. Fruiting material can be confused with that of Sphaerocoryne affinis, but the

persistent calyx and decurrent veins of the latter identify it as Sphaerocoryne.

2. Desmos chinensis Lourt.

(of China)

Fl. Cochinch. (1790) 352. Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921)

255. Masamune, Enum. Phan. Born. (1942) 282. Sinclair, Gard. Bull. Singapore 14

(1955) 266. Beaman et al., Pl. Mt. Kinabalu 4 (2001) 83. — Unona chinensis (Lour.)

DC., Syst. Nat. 1 (1817) 495. TYPE: China, Canton, s. dat., Loureiro s.n. (lectotype,

designated by Ban (1974a), BM).

396 Gard. Bull. Singapore 64(2) 2012

Uvaria monilifera Gaertn., Fruct. Sem. Pl. 2 (1790) 156, t. 114. TYPE: Gaertner’s

plate (Gaertner 1790: t. 114) (lectotype designated by Turner (2011b)). Java, East Java,

Besuki, Nusa Barung Island, between Teluk Tjambak and Teluk Kandangan, 8°30’S,

113°20’E, 11 May 1957, M. Jacobs 4729 (epitype, designated by Turner (2011b), K;

isoepitypes: A, BO, L, LAE, NY, PNH, SING).

Unona discolor Vahl, Symb. Bot. 2 (1791) 63, t. 36. Ridley, Sarawak Mus. J. 1(3)

(1913) 79. TYPE: East Indies, J.G. Kdnig s.n. (holotype: C).

[| Uvaria undulata Roxb., Hort. Bengal. (1814) 43, nom. nud.|

Unona lessertiana Dunal, Monogr. Anonac. (1817) 107, t. 26. TYPE: Uvaria uncata

Vahl in herb. Delessert.

Unona biglandulosa Blume, Biydr. (1825) 16. TYPE: Java, Tjikao, s. dat., J.C.A.

van Hasselt s.n. [Herb. Blume no. 1206] (lectotype, designated by Turner (2011b), L

(barcode no. L 0186343)).

Unona discolor var. bracteata Blume, Fl. Javae Anonaceae (1830) 37, t. 15. TYPE:

Java, in Provincia Krawang.

Unona undulata Wall., Pl. As. Rar. 3 (1832) 42, t. 265, non Unona undulata (P. Beauv.)

Dunal (1817). — Uvaria undulata Walp., Repert. Bot. Syst. | (1842) 77. TYPE: India,

cultivated in Hort. Bot. Cale., Anon. s.n. [EIC 6423B] (lectotype, designated by Turner

(2011b), K-W; isolectotypes: CAL, K).

Uvaria cordifolia Roxb., [Hort. Bengal. (1814) 43, nom. nud.| Fl. Ind. 2 (1832) 662.

TYPE: India, Eastern India, Wo Roxburgh 3663 (lectotype, designated by Turner

(2011b), BM).

Unona amherstiana A.DC., Mem. Soc. Phys. Geneév. 5 (1832) 204. TYPE: Burma,

Amherst, 14 July 1827, N. Wallich s.n. [EIC 6424] (lectotype, designated by Turner

(201 1a), K-W; 1solectotype: BM).

| Unona roxburghiana Wall., Numer. List (1832) no. 6423, nom. nud. |

Unona discolor var. latifolia Hook.f. & Thomson, FI. Ind. (1855) 133. TYPE: Peninsular

Malaysia, Malacca, s. dat., W. Griffith s.n. (lectotype, designated by Turner (2011b), K).

Unona discolor var. laevigata Hook.f. & Thomson, FI. Ind. (1855) 133. — Desmos

chinensis var. laevigatus (Hook.f. & Thomson) Debika Mitra, Bull. Bot. Survey India

35 (1997) 117, as ‘laevigata’. TYPE: India, cultivated in Hort. Bot. Calc., Anon. s.n.

[EIC 6423B] (lectotype, designated by Mitra (1997), CAL; isolectotypes: CGE, K,

K-W).

Climbing Annonaceae of Borneo 397

Unona discolor var. pubiflora Hook.f. & Thomson, FI. Ind. (1855) 133. TYPE: India.

Unona discolor var. pubescens Hook.f. & Thomson, FI. Ind. (1855) 133. Type: India,

Sikkim, s. dat., Anon. [J.D. Hooker] /43 (lectotype, designated by Turner (2011b), K

(barcode no. K000739190)).

Unona discolor vat. parviflora Miq., Fl. Ned. Ind., Eerste Bijv. 3 (1861) 376. TYPE:

Sumatra, Ins. Pulu Pisang prope Padang, Teijsmann.

Unona discolor var. angustipetala Boerl., Cat. Pl. Phan. (1899) 17. TYPE: Java, cult.

in Hort. Bot. Bogor sub XI.A.33a, Anon. s.n. [Bogor. distrib. no. 168) (lectotype,

designated Turner (2011b), L (barcode no. L 0186415): isolectotype: K).

Unona discolor var. brevifolia Teijsm. & Binn. ex Boerl., Cat. Pl. Phan. (1899) 17.

— Desmos chinensis var. brevifolius (Teijsm. & Binn. ex Boerl.) Ban, Bot. Zhurn. 59

(1974) 1773. TYPE: Java, cult. in Hort. Bogor., Anon. s.n. (Bogor. distrib. no. 170 B)

(lectotype, designated by Ban (1974a), US).

Unona discolor var. macropetala Teijsm. & Binn. ex Boerl., Cat. Pl. Phan. (1899) 16.

— Desmos chinensis var. macropetalus (Teijsm. & Binn. ex Boerl.) Ban, Bot. Zhurn.

59 (1974) 1774. TYPE: Java, cult. in Hort. Bogor., Anon. s.n. [Bogor. distrib. no. 174

A] (lectotype, designated by Ban (1974a), US).

Unona discolor var. neglecta Boerl., Cat. Pl. Phan. (1899) 17. TYPE: Java, cultivated

in Hort. Bot. Bogor. sub XI.A.17, Anon. s.n. [Bogor. distrib. no. 168] (lectotype,

designated by Turner (2011b), L (barcode no. L 0181967)).

Unona discolor var. siamensis Scheff. ex Boerl., Cat. Pl. Phan. (1899) 17. TYPE:

Java, cultivated in Hort. Bot. Bogor. sub XI.A.40a, Anon. s.n. [Bogor. distrib. no. 182]

(lectotype, designated by Turner (2011b), L (barcode no. (L 0181979)).

Artabotrys esquirolii Lév., Fl. Kouy-Tcheou (1914-1915) 29. TYPE: China, Kouy-

Tcheou [Gui Zhou], 17 April 1910, J.H. Esquirol 2039 (lectotype, designated by

Turner (2011b), E (barcode no. E00181425)).

Desmos cochinchinensis auct. non Lour.: Merrill, Enum. Philipp. Fl. Pl. 2 (1923) 158.

Scandent shrub or large climber to 40 m long. Twigs slender, striate, dark, slightly

pubescent at the tips. Leaves thinly chartaceous, glabrous, glaucous beneath, oblong,

6-13 x 3-6 cm, base rounded, sometimes emarginate, apex acute to acuminate; midrib

immersed above. prominent below, sparsely hairy below, lateral veins 8—10 pairs,

feint above, prominent below. Petioles 5 mm long, slender. Inflorescences opposite the

leaves, single-flowered. Flowers pendulous, pedicels 3—6 cm long, less than 0.5 mm

thick, slender, glabrous or puberulous; sepals ovate-lanceolate, 5-10 mm long, apex

398 Gard. Bull. Singapore 64(2) 2012

acute, puberulous to glabrous; petals greenish yellow, clawed, narrowly lanceolate,

4-8 cm long, glabrous or sparsely pubescent; stamens numerous, c. | mm long, carpels

many. Fruits pedicel to 6 cm long, c. 20 moniliform monocarps, 34 cm long, glabrous

or minutely pubescent, terminal section apiculate, stipes 10-14 mm long. Seeds 2-6,

globose or ellipsoidal, c. 6-7 x 5 mm, drying light brown, smooth.

Distribution. Widespread from India to the Philippines. In Borneo, widespread,

commonly collected in Sabah, occasional from Sarawak and Kalimantan.

Ecology. Lowland forests. May be cultivated in gardens for its fragrant blooms.

Notes. As a widespread species with a long history of cultivation, it 1s not surprising

that Desmos chinensis has a long list of synonyms including many infraspecific taxa.

The specimens from the wild in Borneo appear relatively uniform and fall well within

the range of morphological variation of D. chinensis.

3. Desmos dumosus (Roxb.) Saff.

(Latin, bushy)

Bull. Torrey Bot. Club 39 (1912) 506. Sinclair, Sarawak Mus. J. 5 (1951) 599. Sinciair,

Gard. Bull. Singapore 14 (1955) 268. Bygrave in Coode et al., Check. Fl. Pl. Gymnosp.

Brunei (1996) 14. — Unona dumosa Roxb., Fl. Ind. 2 (1832) 670. TYPE: Roxburgh

Icon. 2294 (lectotype, designated by Turner (2011b), K). India, cult. in Hort. Bot. Calc.,

s. dat. Anon. s.n. [EIC 6429B] (epitype, designated by Turner (2011b), K-W [x2];

isoepitypes: BM, CGE, K).

Unona subbiglandulosa Miq., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 11. Ridley, Sarawak

Mus. J. 1(3) (1913) 78. — Desmos subbiglandulosus (Miq.) Merr., Philipp. J. Sci.,

C. 10 (1915) 235. Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921) 256.

Masamune, Enum. Phan. Born. (1942) 282. TYPE: Borneo, Martapoera, P W. Korthals

s.n. (lectotype, designated by Turner (2011b), L (barcode no. LO182002)).

Oxymitra monilifera Merr., Univ. Calif. Publ. Bot. 15 (1929) 73. Masamune, Enum.

Phan. Born. (1942) 292. — Dasymaschalon moniliferum (Merr.) P.T. Li, Acta Phytotax.

Sin. 14(1) (1976) 104. TYPE: Borneo, Sabah, near Tawao, October 1922—March 1923,

A.D.E. Elmer 20525 (\ectotype, designated by Turner (2011b), MO (barcode no. MO-

176168); isolectotypes: A, BISH, BM, BO, C, CM, DS, K, L, M, MICH, MO, NY, P,

PH, S; SING; VU).

Scandent shrub. Twigs relatively stout, tomentose when young, striate with numerous

lenticels. Leaves membranous, variably pubescent, drying pale brown or grey with

midrib and veins dark brown, ovate to oblong-ovate, 6-17 x 4—7 cm, base subcuneate to

rounded and emarginate, apex obtuse to acute; midrib sunken above, prominent below,

Climbing Annonaceae of Borneo 399

sparsely pubescent, lateral veins 10—12 pairs, prominent beneath, tertiary venation

scalariform. Petioles 7-20 mm long, rusty tomentose. Inflorescences subopposite leaves,

single-flowered. Flowers pedicels c. 2 cm long, tomentose, sepals ovate-lanceolate,

5—10 mm long, apex acute, almost glabrous; petals greenish yellow, clawed, narrowly

lanceolate, 4-8 cm long, densely pubescent; stamens numerous, c. | mm long, carpels

many. Fruits pedicels 24 cm long, monocarps 30 or more, moniliform, 2-4 cm long,

stipes 1—1.5 cm long, minutely pubescent, terminal segment usually beaked. Seeds 2-5,

globose or ellipsoidal, c. 6-7 x 5—6 mm, drying light brown, smooth.

Distribution. Widespread from India to Borneo. In Borneo recorded from Brunei,

Kalimantan, Sabah and Sarawak.

Ecology. Lowland forest.

4. Desmos dunalii (Wall. ex Hook.f. & Thomson) Saff.

(M.F. Dunal (1789-1856) French botanist who published a monograph of the

Annonaceae in 1817)

Bull. Torrey Bot. Club 39 (1912) 506. Sinclair, Gard. Bull. Singapore 14 (1955) 263. —

Unona dunalii Wall. ex Hook.f. & Thomson, FI. Ind. 1 (1855) 131. TYPE: Peninsular

Malaysia, Penang, G. Porter s.n. [EIC 6425] (lectotype, designated by Turner (2011b),

K (barcode no. K000691358): isotypes: CAL, CGE, GZU, K, K-W).

Liana to 30 m long. Twigs slender, glabrous with pale lenticels. Leaves subcoriaceous,

glabrous except for some hairs on midrib beneath, sometimes glaucous beneath,

drying pale brown above, oblong-elliptic to oblong-oblanceolate, 6-13 <x 3-6 cm,

base rounded to acute, apex acute; midrib flush to slightly sunken above, prominent

below, lateral veins 11-13 pairs, fine and faint on both surfaces, tertiary venation

reticulate. Petioles S~7 mm long. Inflorescences axillary or terminal, flowers single

or occasionally in pairs. Flowers fragrant, pedicel 8-10 mm long with tiny adpressed

hairs, sepals reflexed, broadly ovate, 5—7 mm long, puberulous; petals greenish yellow,

drying brown, narrowly oblong-lanceolate, 25-32 mm long, sometimes covered with

short adpressed hairs; stamens numerous, carpels many. Fruits pedicel to 15 mm long,

monocarps moniliform, numerous, glabrous, 13-35 mm long. Seeds 1—5, globose or

ellipsoidal, c. 8 x 5 mm, drying brown, smooth.

Distribution. Malay Peninsula and Borneo. In Borneo collected from Brunei, Sabah

and Sarawak.

Ecology. Lowland forest.

Notes. Care must be taken not to include fruiting Sphaerocoryne affinis with this or D.

acutus, though Sphaerocoryne does not have moniliform monocarps.

400 Gard. Bull. Singapore 64(2) 2012

Sap

FISSISTIGMA Griff.

(Latin, fissum = to split, divide, separate, stigma = stigma)

Notul. Pl. Asiat. (Posthum. Pap.) 4 (1854) 706. Sinclair, Gard. Bull. Singapore 14

(1955) 349. Irawan, Floribunda 2(7) (2005) 173. Turner, Nord. J. Bot. 27 (2009) 362—

369. TYPE: Fissistigma scandens Griff.

Woody climbers typically tomentose with simple hairs. Leaves with scalariform

tertiary venation. Inflorescences leaf-opposed, or axillary, sometimes appearing

terminal, fascicles of few to many fowers. Flowers bisexual, buds conical, sepals 3,

valvate, more or less connate at base, petals valvate, coriaceous, two whorls of 3, inner

slightly smaller, coriaceous, triquetrous, concave at base, stamens many, connectives

with slightly prolonged apex, obtuse or apiculate, carpels many, pubescent, ovules 2 or

more, biseriate. Fruits cylindrical, ellipsoidal or globose, thick-walled, stipitate. Seeds

several in two rows, smooth and shiny.

Distribution and diversity: Some 60 species or so. India and Sri Lanka, Indochina,

China and through Malesia to Australia. 15 species recorded from Borneo.

Notes. Merrill (1919) included Mitrella and Pyramidanthe in Fissistigma. Based on

molecular analyses, Bygrave (2000) supported this generic reduction. However other

authors (Sinclair 1955; Ban 1974b; Kessler 1993; Jessup 2000; Irawan 2005) have

maintained Fissistigma in the narrower sense. I follow the latter position because

Fissistigma is quite readily distinguished from Mitrella and Pyramidanthe on

morphological grounds, at least as far as Borneo material is concerned, and there has

been no recent peer-reviewed publication supporting unification.

Key to Fissistigma species

la. Midrib of dry leaf with a raised longitudinal ridge on upper surface, monocarps

drying rugullose ses ccdvanecdeisseaeel dso ee F. rugosum

b. Midrib of leaf without a raised longitudinal ridge, monocarps drying relatively

smooth (F° /atifolium with irregular ridges, F’ Jongipetalum verruculose) .....

2a. Leaves more than 13 cm wide, petiole 6 mm thick or more, pedicel of flower

atileast2 -Sicmilong.,3: min thicken eee eee ee F. crassicaule

b. Leaves less than 13 cm wide, petiole generally 3 mm or less in diameter (up

to 5 mm in F: kingii), pedicel of flower generally 2 mm or less in diameter, if

thicker then pedicel less tham 2-5 cml Lomein. ascents renter eee eee ee ee 3

Climbing Annonaceae of Borneo 40]

6a.

Ta.

8a.

9a.

10a.

ia:

ower lamina elabrous' (to naked: eyeior touch) 2.20. 2.c.02.542 secs kesncessecescoceseotes 4

LIGETI AG) Ng kya ee ea ey 2 7!

eaves with 18 of more paits of latetal VEINS ...0:2.06:csc0cesceceaeo-csesnesee F. elmeri

Beavestwitns| 2.06 fewer pairs of lateral VeIns,,....2....24.....2.0.d0s0deeeseeesicenelocsecee 5

Leaf with midrib above glabrous, flowers drying dark brown externally,

medial bract absent or very close (3 mm or less) to the base of the pedicel,

monocarps more or less glabrous, drying dark brown ................. F. bygravei

Leaves with fringe of hairs on midrib above, flowers drying pale or golden

brown externally, medial bract more than 3 mm from the base of the

pedicel, monocarps covered in short dense tomentum drying pale to mid-br

Hairs on midrib above more or less erect, outer petals more than 2 cm long,

monocarps verruculose, covered with dense short brown curly hairs ............

ON PMMIRC DES 5285 2 Aon op Sn sdsnduicssnsuishveaitddadae casbceu saSeeoutensddes F- longipetalum

Hairs on midrib above prostrate or decumbent, outer petals less than 2 cm

long, monocarps smooth, covered with straight adpressed golden brown hairs

ee E ea ERIN CT enn SIE FEY a3, LaN stews Suse Sh RapeBouheosteMeseesawoes F- fulgens

Leaf lower lamina when viewed under a lens (« 10) with more or less straight

RSG E ORGS GUND CNG ooo csc tcnon cocnsseqenecuacets vanceuceterccsscess «uacpaccertuncuvesesth 8

Leaf lower lamina when viewed under a lens (* 10) with curled hairs ......... 11

Leaves typically drying silvery grey above, medial bract and sepals 5 mm or

TGS: KATIE os a2 nak cs en en F) manubriatum

Leaves typically not drying silvery grey above, medial bract and sepals to 4

RATAN ABE IA ES Owe etn ca 8s See se 25h tS sud Raivnancusaenatee tbeddnepbecsssieas dedi davesassenanuhe 9

Hairs on lower lamina more or less erect, flowers in few-flowered panicles

RR A re Ce 5th ub ae aches 2h sas ews dinnidasacinswabus sts cetesuasate F, paniculatum

Hairs on lower lamina decumbent to adpressed, flowers solitary in leaf axils ..

ae RN Mn eM ey. Te Lock Sa seca bat asp ekdea seth tote mate sate ace eotbceaeteneeb ova 10

Flower pedicel very slender (c. 0.5 mm thick when dry), monocarps to 1.5

eimeiametemistip|es to:425 cma 1OM ec. 4.1.24 see cnsestessscsvocsvoncesseneee F. kinabaluense

Pedicel c. 1 mm thick when dry, monocarps c. 2.5 cm diameter, stipes to 2.5

CUNT CLO ati ads ER a a a Ee SE Se eee eee F. carrii

Tertiary venation clearly visible under magnification (* 10) ......... eee I

Tertiary venation not clearly visible under magnification or obscured by dense

ROWSE TACIT) cole Cr ne Aa ar os Ee REPRE Ro ce Soe ee een et 14

402 Gard. Bull. Singapore 64(2) 2012

12a. Many orders of venation on lower lamina clearly visible under magnification

(x 10) and distinctly raised beneath, monocarps globose, stipes typically

longer than'seed=bearing portion Of mOnOCakpSie. snes tee ee eee 13

b. Tertiary venation on lower lamina clearly visible under magnification but

higher orders of venation obscure, monocarps ellipsoidal, stipes shorter than

Seed-beanihe, POnilOnOl MONOCALS acre e eee ee ee F) brevistipitatum

13a. Leaves drying brown above with lateral veins sunken, inner petals more or

less glabrous, monocarps drying with irregular ridges or wrinkles ...............

si SSE A ERT ee TE F: latifolium

b. Leaves drying grey-green above with lateral veins flush, inner petals covered

with a short grey tomentum externally and in upper portion of the inner

surface, monocarps drying more or less smooth ................::ceeseeeeeeee: F- kingie

14a. Tertiary venation of lower lamina obscured by dense brown tomentum, medial

bract 44mm or miore Ome C.c.rcccesccsestsceacesteceeso eee eee F. multivenium

b. Tertiary venation of lower lamina scarcely visible under magnification, not

obscured by tomentum, medial bract absent or to | mm long .... -) montanum

1. Fissistigma brevistipitatum |1.M.Turner

(Latin, brevi= short, stipitatus = provided with a stalk; the short stipes to the monocarps)

Nordic J. Bot. 27 (2009) 363. TYPE: Borneo, Sabah, Interior Division, Nabawan

District, Sungai Millian, 7 November 1986, Swmbing Jimpin SAN 118571 (holotype:

K (barcode no. K000580891); isotypes: A, L, SAN).

Fissistigma sp. 4: Beaman et al., Pl. Mt. Kinabalu 4 (2001) 86.

Large woody climber. Twigs generally drying black with abundant raised pale lenticels,

sometimes brown and striate, young parts densely red-brown tomentose. Leaves

chartaceous to subcoriaceous, expanding leaves very densely red-brown hairy below

and on nerves above, pale hairs on lamina above, mature leaves drying pale brown or

grey-brown above with lateral nerves distinctly paler, brown below with midrib and

nerves a darker shade, ultimately glabrous above except for hairs on midrib which

can be restricted to a few near the petiole attachment, below with red-brown curly

hairs, dense on nerves, abundant on lamina, lamina oblong elliptic to oblong obovate,

4-8.5 « 1.5-4 cm, base obtuse to rounded, apex obtuse to rounded, lateral veins 7—11

pairs, arching forward, looping obscurely, tertiary venation visible from below. Petiole

7— 11 mm long, 1—2 mm thick. Inflorescence terminal or leaf-opposed, few-flowered

panicle. Pedicel to 15 mm long, 2 mm thick, densely red-brown hairy, medial bract

acute, to 2 mm long, densely hairy outside, sepals triangular, c. 3 x 3 mm, outer petals

15 x 5 mm, dense red-brown hairs externally, inside covered with very short pale

brown wool, inner petals ovate lanceolate c. 11 x 4 mm, dense very short brown hairs

Climbing Annonaceae of Borneo 403

outside, inside also in upper portion, excavation glabrous, verrucose, stamens many,

carpels numerous, densely red-brown hairy. Fruiting pedicel 12-15 mm long, 24 mm

thick, monocarps to 5 or more, ellipsoidal, 3—3.5 x 2.5 cm, apex broadly nippled.

drying densely, very shortly red-brown woolly, stipe, shorter than seed-bearing portion

of the monocarp, 1—1.5 cm long, 4 mm thick. Seeds c. 12 in 2 rows, smooth, brown,

c.13x9x4mm.

Distribution. Endemic to Borneo where it has been collected quite widely in Sabah and

once from Sarawak.

Ecology. Lowland and hill forest to 1200 m.

2. Fissistigma bygravei |.M.Turner,

(Paul C. Bygrave (1970—), British Annonaceae taxonomist and nurseryman)

Nordic J. Bot. 27 (2009) 366. TYPE: Borneo, Sabah, West Coast Division, Penampang

District, 5® mile path from Kpg. Babgon to Ulu Terian, 15 October 1969, RF. Cockburn

SAN 635500 (holotype: K (barcode no. K000580483); isotypes: L, SAN, SAR, SING).

Large woody climber. Twigs pale brown, striate, youngest parts brown, sometimes with

very short dark brown hairs. Leaves chartaceous, drying dark grey brown above, dark or

pale purply brown below, midrib and laterals flush above in dry leaves, midrib prominent

below, laterals very slightly raised below, leaves glabrous above, below glabrous to the

naked eye but very short adpressed golden brown hairs visible under magnification,

leaves elliptic, ovate or oblong obovate, 3.5—-15 < 1.5—7 cm, base obtuse to rounded,

apex very slightly notched, blunt or shortly acuminate, lateral veins, 9-14 pairs, arching

forward and looping obscurely, tertiary venation visible from both surfaces in dry leaves.

Petiole 4-12 x 1-2 mm. Inflorescences terminal or leaf-opposed, many-flowered panicle.

Pedicel 7-17 mm long, c. 1 mm thick, with red-brown or dark brown short adpressed

hairs, medial bract sometimes absent, otherwise within 3 mm of base of pedical, sepals

ovate, c. 1 x 2 mm, somewhat reflexed, drying black with short adpressed brown hairs,

outer petals coriaceous, ovate lanceolate, 12-17 x 4 mm, drying blackish or dark brown

with short adpressed brown or red-brown hairs outside, inside and on edges with short

pale woolly hairs, inner petals ovate lanceolate 8 x 3 mm, base concave, short woolly pale

brown externally and on upper portion internally, concavity glabrous, stamens many, c. |

mm long, carpels many. Fruiting pedicel 16-18 mm long, 3 mm thick, monocarps to 12

or more, ellipsoidal, to 2 cm long, apex with a broad short beak, glabrous, drying black

or dark brown, stipe c. 10 mm long, 3 mm thick. Seeds several.

Distribution. Endemic to Borneo where it has been collected from Kalimantan, Sabah

and Sarawak.

404 Gard. Bull. Singapore 64(2) 2012

Ecology. Lowland rain forest to 680 m.

Notes. Among Bornean species, F) bygravei is most similar to F. longipetalum.

Fissistigma bygravei differs from F) longipetalum in having leaves glabrous above,

medial bract absent or very close (3 mm or less) to the base of the flower pedicel, outer

petals drying dark brown and monocarps drying dark brown, smooth and more or less

glabrous.

3. Fissistigma carrii 1.M.Turner

(Cedric Erroll Carr (1892—1936), New Zealand-born rubber planter and orchidologist)

Nordic J. Bot. 27 (2009) 367. TYPE: Borneo, Sabah, Mt Kinabalu, path to Ranau,

c. 4800 ft, 15 Apr 1933, CE. Carr SFN 27006 (holotype: SING (barcode no. SING

0108643)).

Fissistigma sp. 2: Beaman et al., Pl. Mt. Kinabalu 4 (2001) 85.

Large woody climber. Twigs drying dark grey with shallow longitidinal wrinkles,

youngest parts covered with decumbent golden brown hairs. Leaves chartaceous to

subcoriaceous, drying pale to dark brown above, brown to pale brown beneath, midrib

immersed to slightly sunken above, prominent beneath, lateral veins immersed above

raised beneath, glabrous above except for hairs on midrib, below sparsely to densely

tomentose, with more or less straight, decumbent pale brown hairs lying parallel

to lateral nerves, lower lamina surface realtively smooth, not granular or wrinkled,

elliptic, ovate to ovate-lanceolate, 3.5-10 =x 1.5—3 cm, base obtuse to rounded, apex

obtuse, acute or acuminate, lateral nerves 12—20 pairs, distinct from both surfaces in

dry leaves, tertiary venation sometimes distinct from below. Petiole 6-13 mm long,

1-2 mm thick. Inflorescences axillary, single-flowered. Flower pedicel 6—9 mm long,

1 mm thick, densely golden brown hairy, sepals triangular, c. 2 * 2 mm, brown hairy

outside, more or less glabrous within, outer petals ovate-lanceolate c. 11 x 4, outside

brown hairy, inside with very short curly brown hairs near margins otherwise more or

less glabrous, drying black, minutely papillate, inner petals ovate-lanceolate, c. 9 x 3

mm, drying black, glabrous except for scattering of short brown hairs coinciding with

the gaps between the outer petals, drying verrucose inside, stamens many, carpel many,

hairy. Fruiting pedicel 7-15 x 3-4 mm, monocarps to 16 or more, globose, c. 2.5 cm

diameter, drying dark brown, smooth, sparsely covered with adpressed golden brown

hairs, densest near apex, apex rounded sometimes with very short, broad apiculus,

stipe 1—2.5 cm long, c. 2 mm thick. Seeds 10-12, flat oblong but with one side more

curved, c. 16 x 9 x 3 mm, dark brown, smooth and shiny.

Distribution. Endemic to Borneo. Only known from around Mt Kinabalu in Sabah.

Ecology. Montane forest at 1200-1800 m.

Climbing Annonaceae of Borneo 405

Notes. Fissistigma carrii is similar to F. kinabaluense, which, as its name suggests.

was also described from Mt Kinabalu. Fissistigma carrii differs in not having the long

slender flower pedicels of F kinabaluense, or the clear central ridge and radiating

nerves visible on the outer petals. The monocarps of F. carrii are larger (2.5 vs 1.5 cm

diameter) than those of F- kinabaluense.

4. Fissistigma crassicaule |.M.Turner

(Latin, crassus = thick, caulis = stem)

Nordic J. Bot. 27 (2009) 365. TYPE: Borneo, Sarawak, Kuching Division, Kuching

District, Stampin, 5 miles south of Kuching, 6 January 1966, J.A.R. Anderson & P.

Chai § 22770 (holotype: K[<2] (barcode nos. K000580484, K000580485): isotypes:

L, SAR).

Large woody climber. Twigs densely red-brown hairy, at least when young. Leaves

chartaceous to coriaceous, drying dark brown to dark grey-brown above, brown or

greeny brown below, glabrous above except for dense fringe on midrib and some

laterals, below densely golden brown or pale brown hairy on nerves, sparser on lamina,

lamina surface smooth, oblong-elliptic, 22-32 x 13-16 cm, base obtuse, truncate to

very slightly cordate, apex rounded to slightly notched, lateral veins 16—17 pairs.

Petiole to 3 cm long, 6 mm thick. Inflorescence terminal, many-flowered panicle.

Pedicel 2.5—3 cm long, c. 3 mm thick, densely brown hairy, medial bract ovate c. 4 x

3 mm, brown hairy outside, sepals ovate, 5 x 4 mm, brown hairy outside, outer petals

ovate lanceolate, 15 x 8-10 mm, densely brown hairy outside, inside glabrous, drying

dark brown, except for edges and narrow margin where pale brown adpressed woolly,

inner petals ovate acuminate, c. 12 x 8 mm, apex acute, almost glabrous except for

pale hairs near apex externally, drying dark brown, pimply outside, verrucose inside,

stamens many, red, drying pale, carpels numerous, hairy. Fruiting pedicel c. 4.5 cm

long, 6 mm thick, densely brown hairy, monocarps to 20 or more, globose, c. 1.5 cm

diameter, densely golden brown hairy, hairs long, straight, more or less erect, stipe to

2 cm long, 3 mm thick.

Distribution. Endemic to Borneo. Known from two collections (type, Beaman 11834)

from disturbed lowland kerangas (heath) forest in western Sarawak.

Notes. In its general robustness Fissistigma crassicaule is only approached among

the Bornean species by F. rugosum. However F. rugosum is readily distinguished

by a characteristic raised longitudinal ridge along the top side of the midrib in

dry leaves, its almost glabrous foliage, relatively indistinct tertiary venation and

monocarps drying rugose. Otherwise leaves more than 13 cm wide, petioles 6 mm

or more thick and flower pedicel 25 mm long and 3 mm thick or more distinguish

F. crassicaule from all the other species native to Borneo. Fissistigma crassicaule is

essentially a giant version of Fissistigma latifolium (Dunal) Merr. (at least as far as

406 Gard. Bull. Singapore 64(2) 2012

Borneo material referred to F- Jatifolium is concerned), and the type of F. crassicaule

has been previously determined as F- Jatifolium. However F: crassicaule lacks the

ridges on the dry monocarps characteristic of F. /atifolium and the surface of the lower

lamina is smooth in F- crassicaule lacking the granular or cobwebbed appearance of F:

latifolium. One can speculate whether F. crassicaule represents a local species derived

from F: /atifolium in the heath forest of Sarawak.

5. Fissistigma elmeri Mert.

(A.D.E. Elmer, 1870-1942, American plant collector and botanist)

Univ. Calif. Publ. Bot. 15 (1929) 72. Masamune, Enum. Phan. Born. (1942) 284. TYPE:

Borneo, Sabah, Tawao, October 1922-March 1923, 4.D.E. Elmer 20881 (lectotype,

designated by Turner (2011b), UC (sheet no. 289981); isolectotypes: B[ <2], BISH,

BM[x2], BO, C, CM, DS, GH, K, L, M, MICH, MO, NY, P[<2], PH, S, SING, US[<2]).

Large woody climber. Twigs drying black or dark chestnut brown with shallow

latticing, redder when young, striate, very youngest parts with pale hairs, soon lost.

Leaves chartaceous to subcoriaceous, drying pale grey brown to dark brown above, or a

mixture of shades, quite shiny, more uniform mid-brown below with darker midrib and

laterals, midrib slightly sunken above in dry leaves, prominent beneath, laterals flush

above, prominent beneath, short hairs on midrib above, but to the naked eye appearing

more or less glabrous, sparse very short adpressed hairs on lamina and nerves beneath,

oblong oblanceolate or oblong elliptic, 9-19 « 3.5—9 cm, base broadly cuneate with a

round or truncate termination, apex emarginate, obtuse, apiculate or acuminate, acumen

rather sharply pointed, lateral veins 18—22 pairs, angled forward, more or less parallel,

looping obscurely, short but distinct intersecondary veins present, tertiary venation

scalariform, ultimate reticulations areolate giving a characteristic matt appearance to

lamina below. Petiole 8-14 mm long, 2 mm thick. Inflorescence terminal or axillary.

Pedicels 10-20 mm long, | mm thick, drying red-brown, slightly wrinkled, with pale

hairs, medial bract ovate, c. 2 x 1 mm, base truncate, sepals coriaceous, triangular, c.

2—3 « 2-3 mm, slightly connate at the base, densely pale brown hairy outside, glabrous

inside; outer petals thick, coriaceous, ovate lanceolate, 12-35 x 4-6 mm, outside with

central longitudinal ridge, covered in adpressed pale brown hairs, inside very short

adpressed pale hairs, sparse on papillate portion near base; inner petals ovate lanceolate,

8-10 x 3 mm, base deeply concave, externally with central ridge, covered with dense,

very short adpressed pale hairs, inside sparsely hairy in upper portion, more or less

glabrous basally, stamens many, I—1.5 mm long, connective apex pointed, curved

inwards, carpels pale hairy, stigmas drying black with long pale hairs. Fruiting material

scarce, monocarps globose, c. 2 cm diameter, stipitate, densely red-brown adpressed

tomentose.

Distribution. Endemic to Borneo, where it has been collected quite widely from

Kalimantan, Sabah and Sarawak.

Climbing Annonaceae of Borneo 407

Ecology. Lowland forest to 600 m.

6. Fissistigma fulgens (Hook.f. & Thomson) Merr.

(Latin, shiny, bright-coloured)

Philipp. J. Sci., C. 15 (1919) 131. Sinclair, Gard. Bull. Singapore 14 (1955) 353.

Bygrave in Coode et al., Check]. Fl. Pl. Gymnosp. Brunei (1996) 15. Beaman et al., PI.

Mt. Kinabalu 4 (2001) 84. Irawan, Floribunda 2(7) (2005) 178. — Melodorum fulgens

Hook.f. & Thomson, Fl. Ind. (1855) 120. Ridley, Sarawak Mus. J. 1(3) (1913) 90.

Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921) 261. Masamune, Enum.

Phan. Born. (1942) 287. TYPE: Peninsular Malaysia, Malacca, s. dat., W. Griffith s.n.

(lectotype, designated by Turner (2011b), K (barcode no. K000574633)).

[Uvaria fulgens Wall., Numer. List (1832) no. 6482, nom. nud.]

[Myristica finlaysoniana Wall., Numer. List (1832) no. 6793), nom. nud.|

Magnolia ferruginea P. Parm., Bull. Sci. France Belg. 27 (1896) 203, 263; non M.

ferruginea Hort. ex W. Wats. (1889). TYPE: India (?wrongly localised), s.dat., TS.

Ralph s.n. (holotype: P (barcode no. P01964102)).

Melodorum parviflorum var. angustifolium Boerl., Icon. Bogor. 1 (1899) 134. Ridley,

Sarawak Mus. J. 1(3) (1913) 90. Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No.

(1921) 262. TYPE: Borneo, Sarawak, near Kuching, 17 October 1894, G.D. Haviland

& C. Hose 416L (holotype: BO (sheet no. BO-1349056); isotype: CGE).

Large woody climber. Twigs drying dark brown or dark grey, young parts with dense

red-brown to pale brown adpressed hairs giving a rough feel to finger running proximally

down twig. Leaves chartaceous, drying dark brown to almost black above, often with

patches or wash of a paler grey-brown, rich brown below with veins and midrib a

slightly redder hue, midrib slightly sunken above in dry leaves, prominent beneath,

laterals flush above, prominent beneath, tiny golden or red-brown tightly adpressed

hairs on lamina and nerves below, hairs on lamina running more or less parallel to

the lateral nerves, scarcely visible to naked eye except as a reflective metallic sheen,

above pale hairs on nerves, lamina ovate lanceolate, 4.5—18 < 1—S cm, base obtuse to

rounded, apex acute to acuminate, lateral veins 7—11 pairs, arching forward, looping

obscurely, tertiary venation obscure to the naked eye; petiole 6-11 mm long, 1-2

mm thick. Inflorescence terminal or axillary. Pedicels 7—9 mm long, c. 1 mm thick,

widening distally, densely adpressed straw brown or golden brown hairy, medial bract

ovate, 3 x 2 mm, apex acute, sepals triangular, c. 3 x 3 mm, densely hairy outside,

glabrous inside except for scattered hairs near the apex; outer petals coriaceous, ovate

lanceolate, 10 x 5-6 mm, apex acute, externally densely covered in adpressed straw

brown hairs, similar inside except for a glabrous papillate portion near the base; inner

408 Gard. Bull. Singapore 64(2) 2012

petals ovate lanceolate, 7-8 x 4—5 mm, base deeply concave, drying dark brown except

for scattering of adpressed pale hairs near the apex externally, stamens many, 1—2 mm

long, connective apex acute, curved inwards, carpels many. Fruiting pedicel c. 15 mm

long, 4 mm thick, monocarps to 15 or more, ellipsoidal, c. 2.5 x 2 cm, densely golden

brown adpressed hairy, apex rounded, drying relatively smooth, stipe to 20 x 3 mm.

Seeds many, drying dark brown, smooth, shiny, 11-14 « 8—9 x 4 mm.

Distribution. Malay Peninsula, Sumatra, Borneo and the Philippines. In Borneo

widespread with most collections from Sabah, but also Brunei, Kalimantan and

Sarawak.

Ecology. Lowland forest.

7. Fissistigma kinabaluense (Stapf) Merr.

(of Mt Kinabalu)

Philipp. J. Sci., C. 15 (1919) 132. Beaman et al., Pl. Mt. Kinabalu 4 (2001) 84. Irawan,

Floribunda 2(7) (2005) 180. — Melodorum kinabaluense Stapf, Trans. Linn. Soc. Ser.

II 4 (1894) 130. Ridley, Sarawak Mus. J. 1(3) (1913) 90. Merrill, J. Straits Branch

Roy. Asiat. Soc. Spec. No. (1921) 261. Masamune, Enum. Phan. Born. (1942) 287.

TYPE: Borneo, Sabah, Mt Kinabalu, Penokok alt. 3000 ft, G.D. Haviland 1310 [e.h.e.]

(holotype: K (barcode no. K000574681); isotypes: SAR[*2], SING).

Large woody climber. Twigs drying dark grey or dark brown, striate or latticed,

when young with a covering of erect pale hairs, densest on youngest parts. Leaves

chartaceous, drying dark brown or grey brown above, or a mixture of shades, more

uniform brown below with redder brown midrib and laterals, midrib and lateral more

or less flush above, prominent beneath, lower lamina covered with decumbent pale

hairs angled in direction of the laterals, giving a slightly rough feel when rubbed in

the opposite direction, lamina ovate, ovate lanceolate or ovate oblong, 5—12 x 2—S cm,

base rounded or obtuse, apex obtuse to acuminate, lateral veins 12—15 pairs, angled

forward, more or less parallel, looping obscurely, tertiary venation scalariform; petiole

6-8 mm long, 1—2 mm thick. Inflorescence axillary. Pedicels 24 cm long, c. 0.5 mm

thick, densely pale brown hairy, medial bract c. 2 mm long, sepals triangular, c. 4 x 2

mm, apex acute, hairy outside, glabrous inside, outer petals ovate, 13—14 x 3-11 mm,

outside with central longitudinal ridge and radiating nerves, covered in semi-adpressed

pale hairs, inside covered with very short pale woolly tomentum; inner petals ovate, c.

8 x 5 mm, externally with central ridge, covered with pale tomentum on both surfaces,

stamens many, c. | mm long, carpels. Fruiting pedicel 2—3.5 cm « 2 mm, monocarps

to 20 or more, irregularly globose, c. 1.5 cm diameter, apex minutely apiculate, drying

blackish, scurfy grey, stipe to 4.5 cm long, 1 mm thick. Seeds several, dark brown

shiny, c. 10 x 8-9 x 4-5 mm.

Climbing Annonaceae of Borneo 409

Distribution. Endemic to Borneo, where is restricted to Sabah; mostly, but not solely,

to Mt Kinabalu.

Ecology. Hill and montane forest to 900 m.

Notes. See notes under F: carrii for distinction from that species.

8. Fissistigma kingii (Boerl.) Burkill

(Sir George King (1840-1909), British botanist, Superintendent of Royal Botanic

Garden, Calcutta, and first director of the Botanical Survey of India)

Bull. Misc. Inform. 1935 (1935) 317. Masamune, Enum. Phan. Born. (1942) 284.

Sinclair, Gard. Bull. Singapore 14 (1955) 358. Kessler & van Heusden, Rheedea 3

(1993) 63. Irawan, Floribunda 2(7) (2005) 180. Beaman et al., Pl. Mt. Kinabalu 4

(2001) 84. — Melodorum kingii Boerl., Icon. Bogor. 1 (1899) 134. TYPE: Peninsular

Malaysia, Perak, King’s Collector [H.H. Kunstler] 4070 (lectotype, designated by

Irawan (2005), BO (sheet no. BO-1372487); isotypes: CAL, K).

Melodorum parviflorum auct. non Scheff., King, J. As. Soc. Bengal 51 (1892) 107.

Melodorum rubiginosum auct. non (A. DC.) Hook.f. & Thomson. Ridley, Sarawak

Mus. J. 1 (1913) 91, p.p. Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921)

262, p.p. Masamune, Enum. Phan. Born. (1942) 288, p.p.

Melodorum fagifolium Ridl., Bull. Misc. Inform. 1912 (1912) 386. Ridley, Sarawak

Mus. J. 1(3) (1913) 91. Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921) 261.

Masamune, Enum. Phan. Born. (1942) 287. — Fissistigma fagifolium (Ridl.) Merr.,

Philipp. J. Sci., C. 15 (1919) 131. — Fissistigma kingii (Boerl.) Burkill var. fagifolium

(Ridl.) Airy Shaw, Bull. Misc. Inform. Kew 1939 (1939) 238. Masamune, Enum.

Phan. Born. (1942) 284. TYPE: Borneo, Sarawak, Entagut River, December 1894,

C. Hose 397 (lectotype, designated by Turner (2011b), K (barcode no. K000574676);

isolectotypes: A, B, BM, CGE, K, L, P).

Fissistigma kingii var. grandiflorum Airy Shaw, Bull. Misc. Inform. Kew 1939 (1939)

287. Masamune, Enum. Phan. Born. (1942) 284. TYPE: Borneo, Sarawak, near

Kuching, 13 November 1894, G.D. Haviland & C. Hose 3336K (holotype: K (barcode

no. K000574680); isotypes: SAR, SING).

Fissistigma kingii var. multinerve Airy Shaw, Bull. Misc. Inform. Kew 1939 (1939)

287. Masamune, Enum. Phan. Born. (1942) 284. TYPE: Borneo, Sarawak, Fourth

Division, Mt Dulit (Ulu Tinjar) near Long Kapa, 10 August 1932, RW. Richards 1192

(lectotype, designated by Turner (2011b), K (barcode no. K000574679); isolectotypes:

K, SING).

410 Gard. Bull. Singapore 64(2) 2012

Large woody climber. Twigs drying brown, dark to pale, sometimes with pale lenticels,

latticed or striate, youngest parts with short crisped chocolate-brown hairs, almost

velutinous. Leaves chartaceous to coriaceous, typically drying pale grey-green or grey-

brown above, light brown below with darker midrib and veins, midrib and laterals more

or less flush or very slightly sunken above, prominent beneath, upper lamina with short

brown curled hairs in youngest leaves, soon lost except along midrib, lower lamina

with curled chocolate-brown hairs, generally dense on the veins, sometimes longer

straighter hairs present, lower lamina with pale granular appearance under tomentum,

lamina ovate lanceolate to oblong obovate, 6-30 x 2—14.5 cm, base truncate, rounded

or broadly obtuse, apex obtuse, rounded, slightly acuminate or slightly emarginate,

lateral veins 7—23 pairs, looping obscurely, tertiary venation visible from below;

petiole 10-20 mm long, 2—6 mm thick. Inflorescence axillary or terminal. Pedicels

5—20 mm long, 1-3 mm thick, densely brown woolly, medial bract c. 2 x 1 mm long,

apex acute, sepals slightly connate at the base, triangular, 2 x 2-3 mm, apex obtuse,

brown woolly outside, outer petals coriaceous, ovate lanceolate, 10—22(—40) x 3-6

mm, outside with brown woolly tomentum, inside with very short pale grey woolly

tomentum on upper parts, lower glabrous and papillate; inner petals ovate lanceolate,

8-19 x 3-5 mm, base excavate, externally with pale grey short woolly tomentum,

inside woolly also in upper portion, lower part glabrous and papillate, stamens many,

c. | mm long, carpels many. Fruiting pedicel 12-30 mm =x 2-4 mm, monocarps to

20 or more, globose, 2—2.5 cm diameter, apex rounded or minutely apiculate, drying

dark brown, dense brown woolly tomentum, stipe to 3.5 cm long, 2 mm thick. Seeds

several, dark brown shiny, 12—14 x 9-11 x 4-5 mm.

Distribution. Malay Peninsula and Borneo. In Borneo recorded from Brunei,

Kalimantan, Sabah and Sarawak.

Ecology. Lowland forest at 300-900 m.

Notes. This is a variable species and there has been formal recognition of a number

of entities at varietal rank. However the varieties appear to represent fairly arbitrary

divisions of the variation, particularly in terms of number of lateral vein pairs and

flower size. I therefore reduced the varieties to synonymy.

9. Fissistigma latifolium (Dunal) Mert.

(Latin, /ati- = broad, wide; folius = leaved)

Philipp. J. Sci., C. 15 (1919) 132. Merrill, Univ. Calif. Publ. Bot. 15 (1929) 72. Sinclair,

Gard. Bull. Singapore 14 (1955) 359. Kessler & van Heusden, Rheedea 3 (1993) 64.

Irawan, Floribunda 2(7) (2005) 181. Beaman et al., Pl. Mt. Kinabalu 4 (2001) 85.

Unona latifolia Dunal, Monogr. Anonac. (1817) 115. — Uvaria latifolia (Dunal)

Blume, Fl. Jav. Anon. (1828) 37, t. 15. — Melodorum latifolium (Dunal) Hook.f. &

Thomson, FI. Ind. (1855) 177. Ridley, Sarawak Mus. J. 1(3) (1913) 90. Merrill, J.

Climbing Annonaceae of Borneo 411

Straits Branch Roy. Asiat. Soc. Spec. No. (1921) 262. Masamune, Enum. Phan. Born.

(1942) 285. TYPE: Based entirely on Cananga sylvestris III latifolia, Rumphius, Herb.

Amb. 2 (1741) 198.

Uvaria longifolia Blume, Bijdr. (1825) 13, non Uvaria longifolia Sonn. (1782). TYPE:

Java, Anon. s.n. (lectotype, designated by Turner (2011b), L (ex Herb. Blume et Herb.

Groningen) (barcode no. L 0186878)).

Annona rufa C. Presl, Rel. Haenk. 2 (1830) 75. — Melodorum rufum (C. Presl) Merr.,

Philipp. J. Sci., C. 3 (1908) 223. — Fissistigma rufum (C. Presl) Merr., Philipp. J. Sci.,

C. 15 (1919) 132. Merrill, Enum. Philipp. Fl. Pl. 2 (1923) 174. TYPE: Philippines,

Luzon, T.-P.X. Haenke s.n. (holotype: PR[<2] (sheet no. 360883): isotype: HAL).

Melodorum molissimum Miq., F\. Ned. Ind., Eerste Bijv. 3 (1861) 374. TYPE: Sumatra,

Lampongs, J/.E. Teijsmann s.n. [Herb. Bogor. 4252] (holotype: U; isotype: BO).

Melodorum borneense Miq., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 36. Ridley, Sarawak

Mus. J. 1(3) (1913) 88. Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921)

261. Masamune, Enum. Phan. Born. (1942) 287. Fissistigma borneense (Miq.)

Merr., Philipp. J. Sci., C. 15 (1919) 131. Kessler & van Heusden, Rheedea 3 (1993)

63. Irawan, Floribunda 2(7) (2005) 176. TYPE: Borneo, P W. Korthals s.n. (lectotype,

designated by Turner (2011b), L (barcode no. L0O186730)).

Melodorum clementis Merr., Philipp. J. Sci., C. 3 (1908) 136. TYPE: Philippines, Lake

Lamao, Camp Keithley, July 1907, M.S. Clemens s.n. (lectotype, designated by Turner

(2011b), US (barcode no. 00098807); isolectotypes: B, M).

Large woody climber. Twigs relatively smooth, persistently hairy with dense red-

brown tomentum on young twigs, generally paler with age. Leaves chartaceous to

coriaceous, drying dark brown to grey brown above, brown below with main veins

a rather redder shade, midrib flush above, laterals very slightly sunken above in dry

leaves, midrib prominent, rather broad and fiat, beneath, laterals raised beneath,

glabrous above, below densely red-brown hairy on the main veins, lamina and minor

veins with long fine, not completely straight, hairs, lamina surface granular, papillate

or cob-webbed, lamina ovate oblong to elliptic oblong, 8—28 =< 3—10 cm, base slightly

cordate, truncate, rounded or broadly obtuse, apex notched, broadly obtuse, or very

shortly acuminate. Inflorescence axillary or terminal, few- to many-flowered panicles.

Pedicel 1—2.5 cm long, 1-2 mm thick, dense short red-brown hairy, medial bract ovate,

2-5 x 1-3 mm, hairy outside, sepals triangular 2.5-4 x 3-4 mm, hairy outside, outer

petals coriaceous, ovate lanceolate, 9-15 x 5—7 mm, densely red-brown hairy outside,

inside densely short brown woolly on edges and near margins, centrally and basally

generally, glabrous, black or dark brown, slightly papillate or verrucose, inner petals

much thinner than outer petals, ovate lanceolate, 7-8 x 3 mm, more or less glabrous

except for scattering of brown woolly hairs near apex externally, drying black or dark

412 Gard. Bull. Singapore 64(2) 2012

brown, papillate outside, smoother within, stamens many, drying white, apex acute,

carpels many. Fruiting pedicel c. 2 cm long, 3 mm thick, monocarps to 20 or more,

globose, to 2.5 cm diameter, with irregular raised blunt-topped, ridges, surface densely

brown hairy, stipe to 4 cm long, 2 mm thick. Seeds several, semi-circular or ellipsoid

in outline with one or two flat or convex faces, c. 12 x 8-10 x 4 mm, drying dark

brown, smooth.

Distribution. Widespread from Indo-China to the Moluccas. In Borneo recorded from

Brunei, Kalimantan, Sabah and Sarawak.

Ecology. Lowland and hill forest to 1500 m, including peat swamps.

Notes. The name Fissistigma latifolium is entirely based on Rumphius’s Cananga

sylvestris III latifolia. There is no original material or illustration available and I have

not seen any specimen of Fissistigma latifolium collected from Ambon that would be

a potential neotype. I have seen a specimen from nearby Ceram that is fairly close to

F: latifolium.

Two collections from Sabah (SAN 35802, SAN 59263) are more like the

Philippines form of F /atifolium than the typical Borneo form. A couple of specimens

from Mount Kinabalu (Clemens & Clemens 40781, SAN 86075) seem to be a narrow-

leaved form of F. /atifolium, but there are no fruiting specimens available to confirm

this placement.

10. Fissistigma longipetalum (Ridl.) Merr.

(Latin, /ongi = long, petala = petal)

Philipp. J. Sei., C. 15 (1919) 133. Irawan, Floribunda 2(7) (2005) 187. — Melodorum

longipetalum Ridl., Bull. Misc. Inform. 1912 (1912) 387. Ridley, Sarawak Mus. J. 1(3)

(1913) 92. Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921) 262. Masamune,

Enum. Phan. Born. (1942) 287. TYPE: Borneo, Sarawak, Kuching, 11 January 1893,

G.D. Haviland 2102 |Ngaiang leg.| (holotype: K (barcode no. K000574672); isotypes:

BM, SAR, SING).

Large woody climber. Twigs drying black or dark brown, longitudinally wrinkled,

young parts covered with golden brown woolly tomentum. Leaves glaucous below,

thinly chartaceous, drying pale grey brown to dark brown above, or a mixture of

shades, pale or mid-brown below, midrib and laterals flush to slightly sunken above in

dry leaves, prominent beneath, fringe of erect pale hairs on midrib above, adpressed

golden brown hairs on lamina and nerves beneath, lamina ovate to ovate lanceolate,

4-8 x 1.5—3 cm, base obtuse to rounded, apex acute to acuminate, more rarely obtuse,

lateral veins 9-12 pairs, arching forward, more or less parallel, looping obscurely,

tertiary venation scarcely visible to the naked eye; petiole 6-10 mm long, 1 mm thick.

Inflorescence axillary. Pedicels 7-20 mm long, | mm thick, golden brown woolly,

Climbing Annonaceae of Borneo 413

sepals triangular, c. 3.5 « 2 mm, red-brown or golden brown hairy outside, glabrous

inside; outer petals coriaceous, ovate lanceolate, 25-30 x 4 mm, outside with slight

central longitudinal ridge, covered in adpressed golden brown hairs, inside very short

pale curly hairs, sparse on papillate portion near base; inner petals ovate lanceolate,

c. 12 x 4 mm, base deeply concave, externally covered with dense, very short brown

curly hairs, inside more or less glabrous, stamens many, carpels many. Fruiting pedicel

12 x 3 mm, with short brown hairs, monocarps to 5, globose or ellipsoidal, to 2.5

x 2 cm, apex rounded to broad short beak, drying brown with irregular bumps and

wrinkles, densely covered with short brown hairs, stipe 10-20 mm long, c. 4 mm thick.

Seeds c. 5, lunar c. 12—13 x 8 x 5 mm, drying brown, shiny.

Distribution. Endemic to Borneo where it is known from Kalimantan and Sarawak.

Ecology. Lowland forest to 600 m.

11. Fissistigma manubriatum (Hook.f. & Thomson) Merr.

(Latin, manubriatum = handle, heft)

Philipp. J. Sci., C. 15 (1919) 134. Sinclair, Gard. Bull. Singapore 14 (1955) 355.

Kessler & van Heusden, Rheedea 3 (1993) 60. Irawan, Floribunda 2(7) (2005) 184. —

Melodorum manubriatum Hook.f. & Thomson, Fl. Ind. (1855) 118. Ridley, Sarawak

Mus. J. 1(3) (1913) 88. Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921) 262.

Masamune, Enum. Phan. Born. (1942) 287. TYPE: Peninsular Malaysia, Penang, s.

dat., G. Porter s.n. [EIC 6456A] (lectotype, designated by Turner (2011b), K (barcode

no. K000574630); isolectotypes: BM, K-W).

[Uvaria manubriata Wall., Numer. List (1832) no. 6456, nom. nud.].

Melodorum korthalsii Mig., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 37. Ridley, Sarawak

Mus. J. 1(3) (1913) 88. — Fissistigma korthalsii (Miq.) Merr., Philipp. J. Sci., C.

15 (1919) 132. Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921) 261.

Masamune, Enum. Phan. Born. (1942) 287. TYPE: Borneo, Martapoera, s. dat., PW.

Korthals s.n. (lectotype, designated by Turner (2011b), L (barcode no. L 0182131)).

Melodorum bancanum Scheff., Natuurk. Tijdschr. Ned.-Indie 31 (1869) 343. TYPE:

Type: Bangka, prope Muntok et Blienjoe detexit Teysmann.

Large woody climber. Twigs drying black or dark brown with shallow longitudinal

fissures, brown or red-brown woolly tomentum, dense on young parts, generally

persistent. Leaves chartaceous to subcoriaceous, drying silvery grey or dark grey-

brown, shiny, above, golden brown beneath, midrib and lateral more or less flush

above, prominent beneath, lower lamina and nervature covered with dense, long fine

tomentum giving a furry feel with hairs generally running towards margin parallel to

414 Gard. Bull. Singapore 64(2) 2012

the lateral nerves, sometimes relatively sparse, but always present, above glabrous

except for fringe of red-brown or pale hairs along midrib, lamina lanceolate to

narrowly oblong obovate, S—20 x 1.5—6.5 cm, base rounded to obtuse, apex acute

or more rarely acuminate, lateral veins 9—14 pairs, tertiary venation obscure below,

visible from above; petiole 8-10 mm long, 1—2 mm thick. Inflorescence axillary or

terminal. Pedicels 10-15 mm long, 2 mm thick, densely golden brown woolly hairy,

medial bract clasping, ovate, c. 6 5 mm, densely hairy outside, glabrous, black,

inside; sepals triangular or ovate, 6-8 « 5—7 mm, apex broadly acute, densely hairy

outside, glabrous inside, outer petals coriaceous, ovate lanceolate, 15—20 x 6—9 mm,

apex blunt, outside woolly hairy, inside with very short pale woolly tomentum on

upper portions otherwise glabrous and dark brown; inner petals ovate lanceolate, c. 12

x 5 mm, apex acute, base concave, externally with short greyish wool, brown pimply

inside, stamens many, carpels. Fruiting pedicel 10-15 mm =x 4—5 mm, monocarps to 20

or more, globose to ovoid, c. 2 x 2.5 cm diameter, densely woolly brown hairy, apex

rounded to broadly nippled, stipe 1—5.5 cm long, 2—3 mm thick. Seeds c. 10—12, brown

shiny, c. 11 x 7 x 3 mm.

Distribution. Malay Peninsula, ?Sumatra and Borneo. In Borneo widespread, occurring

in Kalimantan, Sabah and Sarawak.

Ecology. Lowland forest.

12. Fissistigma montanum |.M.Turner

(Latin, pertaining to mountains, montane)

Nordic J. Bot. 27 (2009) 364. TYPE: Borneo, Brunei, Pagon Ridge, 5000 ft, April

1958, P.S. Ashton BRUN 2343 (holotype: K (barcode no. K000580482); isotypes:

BRUN, L).

Fissistigma sp. 1: Bygrave in Coode et al., Check]. Fl. Pl. Gymnosp. Brunei (1996) 15.

Fissistigma sp. 1: Beaman et al., Pl. Mt. Kinabalu 4 (2001) 85.

Large woody climber. Twigs drying dark, black, grey or brown, often with abundant, pale

rather irregular lenticels, youngest parts with dense curly brown tomentum, becoming

paler with age. Leaves chartaceous to coriaceous, drying dark brown, brown or grey

brown above, brown or reddish brown below, midrib flush to slightly sunken above,

prominent beneath, laterals immersed above, raised beneath, leaves glabrous above

except for fringe of pale or brown hairs along midrib, below brown hairy, often dense,

hairs very curly, lower lamina surface relatively smooth, not granular or papillose,

lamina elliptic, ovate or obovate, 4-11 x 24.5 cm, base truncate, rounded, obtuse

or acute, apex notched to very shortly acuminate, lateral veins 10—20 pairs, arching

forward and looping obscurely, tertiary venation obscure from both surfaces in dry

Climbing Annonaceae of Borneo 415

leaves, petiole 8-14 mm long, 1—2 mm thick. Inflorescence terminal or leaf-opposed,

1- to few-flowered panicle. Pedicel 1—2.5 cm long, c. | mm thick, covered with dense

brown curly hairs, medial bract tiny, sometimes absent, sepals broadly triangular, 24

x 34 mm, outer petals coriaceous but relatively thin, ovate, 15 x 7, externally densely

short red-brown hairy, inside with very short pale brown hairs on edges and upper and

marginal areas, otherwise glabrous, minutely papillate, drying brown, inner petals, 13

x 5 mm, dense pale brown hairy outside, inside relatively glabrous, drying dark brown

or black, stamens many, carpels many, brown hairy. Fruiting pedicel 1—2 cm long, c. 4

mm thick, monocarps to 25 or more, globose, to 2 cm in diameter, covered with dense,

very short, red-brown woolly tomentum, stipe to 2 cm long, 3 mm thick. Seeds several.

Distribution. Endemic to Borneo. Recorded from Sarawak, Sabah and Brunei.

Ecology. Hill and montane forest at 1000—2600 m.

13. Fissistigma multivenium (Diels) 1.M.Turmer

(Latin, mu/ti = many, vena = vein)

Nordic J. Bot. 27 (2009) 363. — Melodorum multivenium Diels, Notizbl. Bot. Gart.

Berlin-Dahlem 11 (1931) 85. TYPE: Borneo, Sarawak. Bellaga, O. Beccari PB 3789

(holotype: FI-B; isotypes: B(fragment), K).

Fissistigma kingii var. tomentosum Airy Shaw, Bull. Misc. Inform. Kew 1939 (1939)

288. Masamune, Enum. Phan. Born. (1942) 284, Sinclair, Sarawak Mus. J. 5 (1951)

600. Beaman et al., Pl. Mt. Kinabalu 4 (2001) 85. TYPE: Borneo, Sarawak, Bellaga.

O. Beccari P.B. 3789 (holotype: K (barcode no. K000574683); isotype: FI-B).

Large woody climber. Twigs drying black with shallow longitudinal wrinkles or

latticing, with brown woolly tomentum, dense on young parts, generally persistent.

Leaves chartaceous to subcoriaceous, drying grey brown, dark brown or grey-green

above, brown beneath, midrib slightly sunken above in dry leaves, prominent beneath,

lateral nerves more or less flush above. prominent beneath, glabrous above except

for some very short brown or pale hairs along midrib, lamina below with brown

woolly hairy, usually dense enough to give a soft feel, lamina oblong obovate, 10.5—

20 =x 4.5-8 cm, base obtuse, truncate, rounded or slightly cordate, apex apiculate or

acuminate, lateral veins 21—23 pairs, angles forward almost parallel and straight,

looping obscurely, tertiary venation, despite tomentum, usually visible from below,

faint above; petiole 15-17 mm long, 3 mm thick. Inflorescence axillary or terminal.

Pedicels 13-36 mm long, densely brown woolly hairy, medial bract ovate, 4-6 x 34

mm, apex obtuse, densely hairy outside, glabrous, inside: sepals slightly connate at

base, broadly ovate, 3-5 x 5-7 mm, apex obtuse, densely hairy outside, glabrous

within, outer petals coriaceous, ovate lanceolate, 12-15 =< 5-8 mm, apex blunt,

outside woolly hairy, inside with very short pale grey woolly tomentum near margins,

416 Gard. Bull. Singapore 64(2) 2012

otherwise glabrous and black, papillate near the base; inner petals ovate lanceolate,

10-13 x 2-6 mm, base excavate, externally with short greyish woolly patches, brown

verrucose inside, stamens many, carpels many. Fruiting pedicel 2-3 cm = 2—3 mm,

monocarps to 15 or more, globose, often rather irregular, c. 1 x 1.5 cm diameter,

apex rounded or apiculate, stipe 2-4 cm long, 1—2 mm thick. Seeds 1—several, brown

smooth, shiny, 8—11 x 7-8 x 3 mm.

Distribution. Endemic to Borneo, occurring in Kalimantan, Sabah and Sarawak.

Ecology. Lowland forest.

14. Fissistigma paniculatum (Ridl.) Merr.

(Latin, bearing panicles)

Philipp. J. Sci., C. 15 (1919) 134. — Melodorum paniculatum Ridl., Bull. Misc. Inform.

1912 (1912) 386. Ridley, Sarawak Mus. J. 1(3) (1913) 90. Merrill, J. Straits Branch

Roy. Asiat. Soc. Spec. No. (1921) 262. Masamune, Enum. Phan. Born. (1942) 287.

Bygrave in Coode et al., Check]. Fl. Pl. Gymnosp. Brunei (1996) 15. TYPE: Borneo,

Sarawak, Kuching, G.D. Haviland 1845 (holotype: K (barcode no. K000574667);

isotype: K).

Large woody climber. Twigs drying dull dark grey, shallow longitudinal wrinkles,

youngest parts densely red-brown hairy. Leaves chartaceous to subcoriaceous, drying

dark brown or grey brown above, brown to light brown below with midrib and laterals

darker, midrib flush to very slightly sunken above in dry leaves, prominent beneath,

laterals flush above, prominent beneath; lower lamina with long red-brown hairs under

which lamina surface pale and granular or minutely wrinkled, young leaves hairy

above but soon lost except for pale or brown hairs on midrib, lamina elliptic or oblong

elliptic, 2.5—-13 x 2—5.5 cm, base broadly obtuse to rounded, apex rounded, obtuse,

apiculate or acuminate, lateral veins 8-15 pairs, arching forward, looping obscurely,

tertiary venation scalariform, clearer from below in dry leaves, petiole 6—12 mm long,

c. 2 mm thick. Inflorescences axillary or terminal. Pedicel 1-3 cm long, c. 1 mm

thick when dry, densely red-brown hairy, medial bract triangular to ovate, 2—3 x 1.5—2

mm. apex acute, red woolly outside, glabrous within, sepals triangular or ovate, 2—3 x

2.5—3 mm, apex acute, hairy outside, glabrous within, outer petals coriaceous, ovate,

7-10 x 4-6 mm, red-brown hairy outside, inside with short pale brown or whitish

hairs densest on edges and near margins, more or less glabrous and drying black or

dark brown in central portion, rather variable in surface texture, sometimes papillate

or rugulose, inner petals excavate, ovate or ovate lanceolate, 6-7 x 2—4 mm, very

short pale hairs on upper part inside and out, otherwise glabrous, drying black or dark

brown, inside rather bumpy, stamens many, c. | mm long, carpels many hairy. Fruiting

pedicel to 2 cm x 3 mm, monocarps c. 6, globose or ellipsoidal, 2 x 1.5—2 cm, apex

rounded, drying wrinkled, red-brown woolly hairy, stipe to 5.5 cm long by 2 mm thick.

Climbing Annonaceae of Borneo 417

Distribution. Endemic to Borneo where it has been recorded from Brunei. Kalimantan.

Sabah, Sarawak.

Ecology. Lowland and hill forest to 900 m including heath and peatswamp forest.

15. Fissistigma rugosum J.Sinclair

(Latin, wrinkled or folded)

Sarawak Mus. J. 5 (1951) 600. TYPE: Borneo, Sarawak, Kuching, s. dat., GD.

Haviland s.n. (holotype: SAR).

Cyathocalyx borneensis Boerl., Icon. Bogor. (1899) t. 56. Ridley, Sarawak Mus. J. 1

(1913) 76. Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921) 266. Masamune,

Enum. Phan. Born. (1942) 281. TYPE: Borneo, Sebalouw, JE. Teijsmann s.n. [Herb.

Bogor. no. 10838] (holotype: BO (BO-1346836): isotypes: K, L).

Fissistigma sp., Merrill, Univ. Calif. Publ. Bot. 15 (1929) 72.

Melodorum rubiginosum auct. non (A. DC.) Hook.f. & Thomson, Ridley, Sarawak

Mus. J. 1 (1913) 91, p.p. Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921)

262, p.p. Masamune, Enum. Phan. Born. (1942) 288, p.p.

Large woody climber. Twigs drying somewhat irregular in cross section, smooth,

brown or blackish, young parts with dense short erect brown tomentum, relatively

persistent. Leaves chartaceous or coriaceous, drying dark brown, medium brown or

pale grey-brown above, brown, grey-brown or pale grey-brown below with midrib and

laterals generally a darker shade, midrib above with a prominent central ridge even

if midrib is slightly channelled in dry leaf, laterals also very slightly raised above in

dry leaves, midrib and laterals prominent beneath, lamina glabrous above except for

scattered pale hairs in young leaves, below lamina with dense, very short adpressed

tomentum with a sparser cover of longer pale brown hairs which are denser on the

nerves, lamina ovate to oblong ovate, 13.5-37 x 7-20 cm, base rounded to truncate,

apex broadly obtuse to shortly acuminate, lateral veins 15—28 pairs, arching forward,

more or less parallel, looping relatively more distinctly than other Fissistigma species

in Borneo, tertiary venation subscalariform, distinct from both surfaces in dry leaves.

petiole 15—25 x 2-5 mm. Inflorescences axillary or terminal. Pedicel 7-20 mm long, 2

mm thick, densely brown adpressed hairy, medial bract tiny, broadly triangular c. 1 x

1.5 mm, hairy outside, sepals broadly triangular, c. 1.5 x 2 mm, hairy outside, glabrous

within, outer petals c. 8 x 4 mm, covered with adpressed brown hairs externally, short

woolly hairs cover upper portion internally, inner petals c. 5 x 2.5 mm, concave, dense

very short adpressed hairs outside, glabrous within, stamens 1—1.5 mm long, carpels

many. Fruiting pedicel 20-22 mm long, c. 3 mm thick, monocarps ripening bright

yellow, to 4 or more, globose, c. 2.5 cm diameter, drying brown, rugose, covered with

418 Gard. Bull. Singapore 64(2) 2012

very dense, very short brown hairs, apex rounded to broadly bluntly apiculate, stipe

to 12 mm long, 5 mm thick. Seeds several, drying dark brown, shiny, smooth, c. 15 =

9 x 4-5 mm.

Distribution. Endemic to Borneo where it is known from Brunei, Kalimantan, Sarawak.

Ecology. Lowland forest to 400 m.

FRIESODIELSIA Steen.

(K.R.E. Fries (1876-1966), Swedish botanist;

F.L.E. Diels (1874-1945), German botanist; both Annonaceae specialists)

Bull. Jard. Bot. Buitenzorg, sér. 3. 17 (1948) 458. van Steenis, Blumea 12 (1964) 353.

Polyalthia section Oxymitra Blume, Fl. Javae Anonac. (1830) 71. Oxymitra (Blume)

Hook.f. & Thomson, FI. Ind. (1855) 145, non Oxymitra Bisch. ex Lindenb. (1829).

Sinclair, Gard. Bull. Singapore 14 (1955) 447. TYPE: Polyalthia cuneiformis (Blume)

Blume (= Guatteria cuneiformis Blume, = Oxymitra cuneiformis (Blume) Hook.f. &

Thomson, = Friesodielsia cuneiformis (Blume) Steen.).

Woody climbers. Leaves often glaucous beneath, with paired glands in the edge of the

lamina near the insertion of the petiole, tertiary venation scalariform. Inflorescences

leaf-opposed or supra-axillary, single-flowered. Flowers bisexual, buds conical, sepals

3, valvate, more or less connate at base, petals valvate, two whorls of 3, inner much

shorter than outer, outer petals spreading at maturity, inner petals cohering about

reproductive structures, stamens many, connectives with truncate prolonged apex,

carpels many, ovules 1—5, lateral. Fruits cylindrical, ellipsoidal or globose, thin-walled,

nipple-tipped, stipitate. Seeds 1—few, in cross-section quartered by the longitudinal

walls of the endosperm.

Distribution and diversity: Some 50—60 species. Tropical Africa and Asia, though the

African species may belong in a separate genus. Nine species recorded from Sabah

and Sarawak.

Key to Friesodielsia species

la. Leaves typically more than 20 cm long, base of lamina obtuse slightly decurrent

to petiole, petiole 15 mm or more long, outer petals thin and flat, monocarps

more than 3: ci lOMG./eceraeegses decane eae ee F) formosa

b. Leaves generally smaller than 20 cm long, lamina base often rounded or

auriculate, petiole less than 15 mm long, outer petals thick and generally

triangular or rhomboidal in cross section, monocarps less than 3 cm long ...... 2

Climbing Annonaceae of Borneo 419

2a. Leaves with tightly adpressed hairs beneath, often giving a golden sheen to the

UEC 2 se ncin 5 hae de SE Rc F biglandulosa

b. Leaves hairy or not beneath, but not with a golden sheen ...................0......3

3a. Sepals with ridges radiating externally from base ............0.ccccccccsssescesseeeeeeees “

Bee Sepals without visible ridges externally .........5..55c0ccccceceus sessoecbessstsiecesverse D

4a. Leaves chartaceous, drying dark brown above, glaucous beneath, monocarps

SSC Scile aSep CMO MiiiN IONS. se) a. dices) ssadeoucis acd ineiaadhioaReeeecadieecs F: affinis

b. Leaves coriaceous, drying mid-brown above, beneath hairy brown overlying

glaucous lamina, monocarps stipitate, stipe more than 3 mm long ...... F. excisa

aECAVES TOL EKECEUING S.5-% 325 CMs. ccvenc ieccsevecsececessetsenvestnenesantene soe F- ovalifolia

Be ecavesitypically bieper thaw S:5 * 35 Ci x. s2i.csci.502.ccedeecssacdsoccvecetaddsasectueseneses 6

6a. Leaves glaucous beneath, though this may be obscured by brown pubescence

“ssc ss nee sleuo19 20 iE @alse eRe SEU ISS Ra nei a Ac Na a i

Pe cAVeSNOL SlANCOUS, DENGAN... ...2.....cc.02¢s0s0dscaccesoeaevwucessaceedvassonteeee F. acuminata

7a. Lamina hairy beneath, or if glabrous then under x15 or greater magnification

Ech 2 SAeULL SIF SYS SE RING Sa em ee ee ae 8

b. Lamina glabrous beneath and under magnification relatively smooth and not

TASES IF a 2 RA ae ee ee ee ee eo F. glauca

8a. Leaves typically more than 6 cm wide, flower pedicel to 15 mm long, monocarps

drying dark brown with rusty pubescence ................::cssceseeeeeeeeees F. grandifolia

b. Leaves not exceeding 6 cm wide, flower pedicel 20 mm or more long,

monocarps drying warm brown with rusty pubescence .............. F, borneensis

1. Friesodielsia affinis (Hook.f. & Thomson) D.Das

(Latin, neighbouring, akin to)

Bull. Bot. Survey India 5 (1963) 93. — Oxymitra affinis Hook.f. & Thomson, FI. Brit.

India 1 (1872) 70. Sinclair, Gard. Bull. Singapore 14 (1955) 450. TYPE: Peninsular

Malaysia, Malacca, 12 April 1867, 4.C. Maingay 1851 [Kew distribution no. 59]

(lectotype, designated by Turner (2009), K (barcode no. K000691787)).

Fissistigma magnisepalum Irawan, Floribunda 2(7) (2005) 184, as ‘magnisepala’.

TYPE: Borneo, East Kalimantan, Long Iram Subdistrict, Maruwai, Block Lampunut,

19 March 1999, P. Kessler 2621] (holotype: BO[x2] (sheet nos BO-1318961, BO-

1318960), isotypes: K, L, WAN).

420 Gard. Bull. Singapore 64(2) 2012

Large woody climber. Twigs drying black, latticed, youngest parts densely red-brown

or dark brown hairy. Leaves chartaceous, glaucous beneath, drying dark brown above,

grey-brown beneath with red-brown midrib and laterals, expanding leaves densely

hairy, soon lost above except for fringe on midrib and main nerves, below hairy on

lamina, giving a furry feel, and densely hairy on midrib and laterals, hairs generally

slightly curied or kinked, lamina oblong elliptic to oblong obovate, 5—19(—26) x

2.5—8.5(—10) cm, base rounded to slightly cordate, apex broadly truncate apiculate to

acuminate, midrib and lateral nerves slightly sunken above in dry leaves, prominent

beneath, lateral nerves 7—11 pairs, tertiary venation scalariform, distinct from below,

less clear from above. Petiole 5-6 mm long, 2-3 mm thick. Inflorescences single-

flowered, extra-axillary. Flowers, pedicel 5-8 mm long, | mm thick, densely red-brown

hairy, medial bract ovate to 8 mm long, 5 mm wide, base sometimes cordate (n.b. bracts

and sepals may enlarge after anthesis), sepals free, ovate, c. 6 x 6 mm, base truncate,

apex obtuse to rounded, red-brown hairs inside and out, nerves visible as raised ridges

externally, outer petals coriaceous, ovate lanceolate, 2—5 cm long, excavated at base,

inner petals shorter with a very short claw, stamens many, carpels many. Monocarps

glaucous, 10 or more, drying brown with a greyish bloom, ellipsoidal, c. 10 x 8 mm,

apiculate, covered with red-brown hairs when young, becoming glabrescent, stipe c. 3

mm long, 1-2 mm thick. Seed 1, ellipsoidal, c. 10 <x 7 mm, longitudinal groove.

Distribution. Malay Peninsula and Borneo. In Borneo widespread but little collected

with specimens from Kalimantan, Sabah and Sarawak.

Ecology. Lowland and hill forest to 800 m.

2. Friesodielsia biglandulosa (Blume) Steen.

(Latin, bi = two, glandulosa = bearing glands; pair of glands at leaf base)

Blumea 12 (1964) 358. Kessler & van Heusden, Rheedea 3 (1993) 66. Bygrave in

Coode et al., Checkl. Fl. Pl. Gymnosp. Brunei (1996) 15. Beaman et al., Pl. Mt.

Kinabalu 4 (2001) 86. — Guatteria biglandulosa Blume, Fl. Javae Anonaceae (1828)

102, t. 51. — Monoon biglandulosum (Blume) Mig., Ann. Mus. Bot. Lugd.-Bat. 2

(1865) 19. — Oxymitra biglandulosa (Blume) Scheff., Natuurk. Tijdschr. Ned.-Indié

31 (1870) 341. Ridley, Sarawak Mus. J. 1(3) (1913) 88. Merrill, J. Straits Branch

Roy. Asiat. Soc. Spec. No. (1921) 258. Merrill, Univ. Calif. Publ. Bot. 15 (1929) 73.

Airy Shaw, Bull. Misc. Inform. Kew 1939 (1939) 288. Masamune, Enum. Phan. Born.

(1942) 291. Sinclair, Gard. Bull. Singapore 14 (1955) 459. — Polyalthia biglandulosa

(Blume) Hook.f. & Thomson, Fl. Brit. Ind. 1 (1872) 65. — Richella biglandulosa

(Blume) R.E. Fr. in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, xvii a. I. (1959) 139.

TYPE: Java, in montibus provinciae Bantam.

Large woody climber. Twigs drying black or dark grey, striate or latticed, youngest

parts with adpressed golden brown hairs, soon lost. Leaves chartaceous to coriaceous,

Climbing Annonaceae of Borneo 421

drying dark brown above, rarely grey-brown or pale brown, paler brown below with

darker midrib and lateral veins, midrib slightly sunken above in dry leaves, prominent

beneath, lateral veins very slightly raised above, raised beneath, lamina beneath with

tiny, tightly adpressed, yellow-brown hairs, sometimes sufficiently dense to give a

golden or coppery sheen to leaf underside, sometimes very sparse but can be found

with a lens, lamina elliptic or oblong elliptic to obovate, 5—26 x 2—11 cm, base cuneate,

obtuse, truncate to slightly rounded, glands visible as darkened patches in lamina edge

near petiole attachment; lateral veins 7-13 pairs, arching forward, looping indistinctly,

tertiary venation scalariform, distinct from both surfaces in dry leaves. Petiole 5—13

mm long, 1-3 mm in diameter. Inflorescences of solitary flowers, extra-axillary or on

short twiggy, irregularly branched shoots with few flowers. Pedicel 5-10 mm long, c.

1 mm thick, drying dark brown, adpressed golden brown hairs, medial bract to 3 = 1

mm, sepals free, ovate lanceolate, 5 x 1 mm, adpressed coppery brown hairs, outer

petals green, lanceolate, to 30 x 6 mm, base spoon-shaped, central ridge externally just

visible, covered with dense adpressed pale hairs externally, glabrous internally, inner

petals cream, coherent at edges, ovate lanceolate, 10 x 34 mm, apex acute, some

adpressed hairs externally, central ridge evident, stamens numerous, c. | mm long,

top of connective flat to slightly convex, carpels many, pubescent. Fruiting pedicel

to 20 mm long, 2 mm thick, monocarps to 20, ripening yellow, ellipsoidal, 15—20 x

9 mm (rarely 2-seeded monocarps cylindrical to 30 mm long, with slight constriction

between seeds in dry state) drying black, minutely bumpy, apex rounded, apiculate,

stipe to 20 mm long, | mm thick, some adpressed brown hairs particularly near apex

and on stipe. Seeds 1, rarely 2, ellipsoidal, c. 15—18 « 7-8 mm, drying brown, smooth

with a longitudinal groove.

Distribution. Malay Peninsula, ?Sumatra and Borneo. In Borneo collected from

Brunei, Kalimantan, Sabah and Sarawak.

Ecology. Lowland to montane forest to 900 m, including heath and peat swamp forest.

3. Friesodielsia borneensis (Migq.) Steen.

(of Borneo)

Blumea 12 (1964) 358. Kessler & van Heusden, Rheedea 3 (1993) 66. Beaman et al.,

Pl. Mt. Kinabalu 4 (2001) 86. — Oxymitra borneensis Miq., Ann. Mus. Bot. Lugd.-Bat.

2 (1865) 30. Ridley, Sarawak Mus. J. 1(3) (1913) 88. Merrill, J. Straits Branch Roy.

Asiat. Soc. Spec. No. (1921) 258. Masamune, Enum. Phan. Born. (1942) 291. Sinclair,

Sarawak Mus. J. 5 (1951) 606. TYPE: Borneo, Martapoera, s.dat., PR W. Korthals s.n.

(lectotype, designated by Turner (2011b), L (barcode no. L 0037935)).

Oxymitra oxyphylla Mig., Ann. Mus. Bot. Lugd. Bat. 2 (1865) 29. Ridley, Sarawak

Mus. J. 1(3) (1913) 89. Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921)

259. Masamune, Enum. Phan. Born. (1942) 292. — Friesodielsia oxyphylla (Miq.)

422 Gard. Bull. Singapore 64(2) 2012

Steen., Blumea 12 (1964) 361. TYPE: Borneo, Dano-Bahang, s.dat., PR W. Korthals s.n.

(lectotype, designated by Turner (2011b), L (barcode no. L 0187213)).

Oxymitra beccarii Diels, Notizbl. Bot. Gart. Berlin-Dahlem 11 (1931) 85. —

Friesodielsia beccarii (Diels) Steen., Blumea 12 (1964) 358. TYPE: Borneo, Sarawak,

Bintulu, September 1867, O. Beccari PB. 3742 (holotype: FI-B[<2](Erb. Becc. nos.

305 and 305A); isotypes: B, M).

Large woody climber. Twigs drying dark grey to black, striate or latticed, rusty

tomentose when young. Leaves glaucous beneath, chartaceous, drying dark brown

above, grey-brown or red-brown below, obovate to oblong obovate, 3.5—16 2—6 cm,

base rounded to slightly auriculate, apex obtuse to shortly acuminate, midrib slightly

sunken above in dry leaves, prominent beneath, lateral veins 9—11 pairs, flush above,

prominent beneath. Petiole 3-10 mm long, 1-2 mm thick. Inflorescences single-

flowered, extra-axillary. Flowering pedicel, 2—3 cm long, very slender, sepals c. 6 x 5

mm, hairy, outer petals coriaceous, flat, ovate lanceolate, to 6 cm long, c. 6 mm wide,

base excavated, hairy externally, central ridge, glabrous internally, inner petals c. 8

x 4 mm, markedly acuminate, stamens many, carpels many. Fruiting pedicel 3—8.5

cm long, monocarps to 20 or more, globose to ellipsoidal, 8-12 x 7 mm, drying red-

or mauve-brown with rusty pubsecence, sometimes more or less glabrous, minutely

apiculate, stipe 6-13 mm long, more than | mm thick. Seeds 1, ellipsoidal c. 7-11 =

6 mm.

Distribution. Malay Peninsula, Sumatra and Borneo. In Borneo this species has been

recorded from Brunei, Kalimantan, Sabah and Sarawak.

Ecology. Lowland forest.

Notes. Material of this species outside Borneo is referred to var. suwmatrana.

4. Friesodielsia excisa (Miq.) Steen.

(Latin, cut out)

Blumea 12 (1964) 359. — Oxymitra excisa Miq., Ann. Mus. Bot. Lugd. Bat. 2 (1865)

32. Ridley, Sarawak Mus. J. 1(3) (1913) 89. Merrill, J. Straits Branch Roy. Asiat. Soc.

Spec. No. (1921) 259. Masamune, Enum. Phan. Born. (1942) 291. Sinclair, Gard.

Bull. Singapore 14 (1955)452. — Richella excisa (Mig.) R.E. Fr. in Engl. & Prantl,

Nat. Pflanzenfam., ed. 2, xvii a. II. (1959) 171. TYPE: Sumatra, PW. Korthals s.n.

(lectotype, designated by Turner (2009), L (barcode no. L0O182297)).

Fissistigma cordifolium \rawan, Floribunda 2(7) (2005) 176. TYPE: Borneo, West

Kalimantan, Gunung Palung Nature Reserve, 21 June 1986, van Balgooy & van Setten

5559 (holotype: BO (sheet no. BO-1372431); isotype: L).

Climbing Annonaceae of Borneo 423

Large woody climber. Twigs drying black, striate or latticed, persistently tomentose

with short, dense, brown or red-brown hairs. Leaves chartaceous to coriaceous, densely

brown hairy below covering a glaucous lamina, drying mid-brown above, generally

darker brown below because of the tomentum, lamina oblong to obovate, or even

obdeltoid, 9-36 < 5—15 cm, base rounded to slightly cordate, apex truncate or even

emarginate to obtuse acuminate, midrib and lateral nerves sunken above in dry leaves,

prominent beneath, 10—13 pairs of laterals, tertiary venation scalariform, readily

discernable from both surfaces, many orders of venation visible below, all slightly

raised, short pale hairs on lamina above lost with age but persisting on midrib and

laterals, dense hairs below, rather hooked or curled. Petiole 5-10 mm long, 34 mm

thick. Inflorescences single-flowered, extra-axillary. Flower, pedicel 5-12 mm long,

1-2 mm thick, densely brown hairy, medial bract ovate 5—7 x 6 mm, apex acute, sepals

ovate, c. 6 x 6 mm, apparently enlarging after corolla drops, apex obtuse to rounded,

red-brown hairs, veins raised externally, outer petals coriaceous, ovate lanceolate,

14-16 x 7-8 mm; inner petals coriaceous, ovate lanceolate, c. 12 x 6 mm, apex acute,

stamens many, carpels many. Monocarps ripening red, to 20 or more, ellipsoidal, 15

x 9 mm, or more rarely if two-seeded to 20 mm long, drying dark brown, glabrous

except for near apex and on stipe, stipe to 9 mm long, 2 mm thick. Seed | or rarely 2,

ellipsoidal, c. 13 x 8 mm.

Distribution. Malay Peninsula, Sumatra and Borneo. In Borneo recorded from Brunei,

Kalimantan, Sabah and Sarawak.

Ecology. Lowland forest to 500 m, including peat swamps.

5. Friesodielsia formosa |.M.Turner

(Latin, handsome, beautiful)

Edinburgh J. Bot. 66 (2009) 366. TYPE: Borneo, Central Kalimantan, headwaters of S.

Kahayan, 5 km north-west of Tumbang Sian logging camp, 26 April 1988, J.S. Burley

et al. 832 (holotype: K (barcode no. K000580218); isotypes: E, L, SAR, SING).

Large woody climber. Twigs drying black or rather dirty brown, striate or latticed,

with raised brown lenticels, darker and smoother when young, more or less glabrous.

Leaves chartaceous, glaucous beneath, glabrous, drying dark brown or rather patchy

dull brown above, red-purple brown beneath, sometimes with a greyish wash, with

midrib and laterals dark brown or black, lamina oblong elliptic to oblong obovate, 17—

33 x 6—10 cm, base obtuse, slightly decurrent to petiole but not auriculate, apex obtuse

to slightly acuminate, midrib sunken above and prominent beneath, lateral veins c.

15 pairs, immersed above, prominent beneath, tertiary venation distinct from both

surfaces in dry leaves, more so from above, less distinctly scalariform than other species

in Borneo. Petiole 15—22 mm long, 2-3 mm thick, drying dark, shallowly furrowed.

Inflorescences axillary or supra-axillary, few-flowered or solitary. Flowers fragrant,

424 Gard. Bull. Singapore 64(2) 2012

pedicel 15-40 mm long, c. 0.5 mm thick, drying black, striate, tiny medial bract c. |

mm long, sepals free, broadly ovate, 6-10 « 8 mm, tending to reflex at anthesis, apex

blunt, drying black or dark brown, outer petals yellow or cream, reflexing at anthesis,

thin, ovate lanceolate, 40-50 x 15-20 mm, drying very dark brown, sparse covering

of tiny brown adpressed hairs on both surfaces, pale spots externally, inner petals

ovate, 10-11 x 8 mm, apex acute, drying black with sparse very tiny brown adpressed

hairs, stamens many, c. 2 mm long, carpels c. 6-10, c. 2 mm long, stigmas globose,

hirsute. Fruiting pedicel to 4.5 cm long, 2 mm thick, monocarps 1-7, ripening orange-

red, stipe green, ellipsoidal 4-5 = 1.5—2 cm, drying dark brown, deeply, irregularly

wrinkled, wrinkles with rounded not sharp-edged margins, glabrous, apiculus scarcely

discernible, stipe drying black, 2—3 cm long. Seed 1, c. 3 x 1.5 cm.

Distribution. Endemic to Borneo where it is known from Kalimantan and Sarawak.

Ecology. Lowland forest to 200 m, including on limestone.

6. Friesodielsia glauca (Hook.f. & Thomson) Steen.

(Greek, g/aukos = bluish grey or bluish green; with the powdery bloom as on grapes)

Blumea 12 (1964) 359. — Oxymitra glauca Hook.f. & Thomson, FI. Ind. 1 (1855)

146. Ridley, Sarawak Mus. J. 1(3) (1913) 88. Merrill, J. Straits Branch Roy. Asiat. Soc.

Spec. No. (1921) 259. Mermrill, Enum. Philipp. Fl. Pl. 2 (1923) 176. Masamune, Enum.

Phan. Born. (1942) 291. Sinclair, Gard. Bull. Singapore 14 (1955) 460. — Richella

glauca (Hook.f. & Thomson) R.E.Fr. in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, xvii

a. IT. (1959) 139. TYPE: Peninsular Malaysia, Prince of Wales Island [Penang]. s. dat.,

Anon. [W.E. Phillips] s.n. (holotype: K (barcode no. K000691773)).

Oxymitra diadena Mig., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 31. Ridley, Sarawak Mus.

J. 1(3) (1913) 89 as ‘diadema’. Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No.

(1921) 258, as ‘diadema’. Masamune, Enum. Phan. Born. (1942) 291, as ‘diadema’.

— Friesodielsia diadena (Miq.) Steen., Blumea 12 (1964) 358. TYPE: Borneo, Mt

Sakoembang, PW. Korthals s.n. (lectotype, designated by Turner (2011b), L (barcode

no. L 0182284)).

Oxymitra linderifolia Ridl., Bull. Misc. Inform. Kew 1912 (1912) 385. Ridley,

Sarawak Mus. J. 1(3) (1913) 89. Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No.

(1921) 259. Masamune, Enum. Phan. Born. (1942) 292. — Friesodielsia linderifolia

(Ridl.) Steen., Blumea 12 (1964) 360. TYPE: Sarawak, Kuching, 19 April 1893, G.D.

Haviland 2333 (lectotype, designated by Turner (2012), K (barcode no. K000691760);

isolectotypes: BM, K, SAR, SING).

Oxymitra argentea J.Sinclair, Gard. Bull. Singapore 14 (1955) 461. Bygrave in Coode

et al., Checkl. Fl. Pl. Gymnosp. Brunei (1996) 15. — Friesodielsia argentea (J.

Climbing Annonaceae of Borneo 425

Sinclair) Steen., Blumea 12 (1964) 358. Beaman et al., Pl. Mt. Kinabalu 4 (2001) 86.

TYPE: Peninsular Malaysia, Trengganu, Bukit Kajang, 14 November 1935, E.J.H.

Corner SFN 30457 (holotype: SING (barcode no. SING 0048676); isotype: K).

Large woody climber. Twigs drying black, striate or latticed, when young smooth

and brown, variably hairy, sometimes densely brown tomentose. Leaves chartaceous,

glaucous below, drying dark brown above, grey-brown beneath, lamina glabrous or

glabrescent, hairs more frequent on veins and midrib below, lamina obovate or oblong

obovate to lanceolate, 4-21 < 1.5—8 cm, base obtuse, rounded or truncate, apex broadly

and rather shortly acuminate, midrib and lateral nerves flush to slightly sunken above

in dry leaves, prominent beneath, laterals 7—13 pairs. Petiole S—9 mm long, 0.5—2 mm

thick. Inflorescences, single or few-flowered, extra-axillary. Flower, pedicel 15—35

mm long, less than 1 mm thick, widening distally, very short dense brown pubescence,

medial bract ovate c. 3—5 mm long, apex acute, sepals free, ovate, 4-6 x 2-4 mm, apex

acute, hairy; outer petals coriaceous, ovate lanceolate, 20-40 x 4-7 mm, excavated at

base internally, adpressed brown hairs externally, inner petals ovate acuminate 10—12

x 2-3 mm, central ridge externally, stamens many, carpels many. Fruiting pedicel 6-60

mm long, c. 1 mm thick; monocarps 20-25 or more, ripening red, globose or ellipsoidal

8-11 x 7-8 mm, apiculate, drying black or rusty brown, red-brown pubescent when

young, sometimes persisting, stipe 4-20 mm long, c. | mm thick. Seeds 1,ellipsoidal,

c. 7-10 x 6-7 mm.

Distribution. Malay Peninsula, Sumatra and Borneo. In Borneo collected from Brunei,

Kalimantan, Sabah and Sarawak.

Ecology. Lowland forest.

Notes. There is considerable variation in the material here recognised under the name

Friesodielsia glauca, both within Borneo and across its range. Some of the entities

included appear quite distinct but intermediate forms occur. Friesodielsia linderifolia

is close to typical F- glauca and is certainly a synonym of it.

7. Friesodielsia grandifolia (Merr.) 1.M.Turner, Edinburgh J. Bot. 66 (2009) 369.

Oxymitra grandifolia Merr., J. Straits Branch Roy. Asiat. Soc. 85 (1922) 179. Merrill,

Univ. Calif. Publ. Bot. 15 (1929) 72. Masamune, Enum. Phan. Born. (1942) 291. TYPE:

Borneo, Sabah, Sandakan, September—December 1920, M. Ramos 1910 (lectotype,

designated by Turner (2009b), BO (sheet no. BO-13593592)).

Oxymitra latifolia auct. non Hook.f. & Thomson, Sinclair, Sarawak Mus. J. 5 (1951)

607.

Friesodielsia latifolia auct. non (Hook.f. & Thomson) Steen.: Beaman et al., Pl. Mt.

Kinabalu 4 (2001) 86.

426 Gard. Bull. Singapore 64(2) 2012

Large woody climber. Twigs drying dark grey, younger twigs woolly hairy. Leaves

chartaceous, glaucous or brown hairy below, drying dark brown, occasionally mid-

brown, above, brown or grey-brown below, lamina oblong, oblong obovate or

obovate, 12-30 « 4-14 cm, base clightly cordate to rounded, apex rounded, obtuse

to acuminate, midrib slightly sunken above in dry leaves, raised below, lateral nerves

12-16 pairs, flush to slightly sunken above, raised beneath, tertiary venation clear

from both surfaces unless hidden by tomentum below. Petiole 5-10 mm long, 24 mm

thick. Inflorescence single-flowered, extra-axillary. Flower, pedicel 15—20 mm long, 1

mm thick, rusty tomentose, sepals free, triangular S—8 x 5—7 mm, apex blunt to acute,

rusty hairy, outer petals coriaceous, flat, oblong lanceolate, c. 32-45 x 8-10 mm, base

excavated, externally with brown hairs, densest near base, central ridge discernible but

not prominent, internally glabrous, minutely papillate inside excavation, inner petals

coherent, ovate acuminate, c. 8-10 x 4-6 mm, glabrous except for a few scattered

hairs externally with a slight central ridge, minutely papillate inside, stamens many,

carpels many. Fruiting pedicel 10-20 mm long, 1-2 mm thick, monocarps to 25 or

more, globose to ellipsoidal, 12—13 x 8 mm, to 17 mm long in two-seeded monocarps,

drying black or dark brown with rusty hairs particularly dense near apex and on stipe,

stipe to 13 mm long. Seeds, 1| or rarely 2, ellipsoidal, c. 10-12 x 6-7 mm.

Distribution. Endemic to Borneo where it is recorded from Kalimantan, Sabah and

Sarawak.

Ecology. Lowland forest, also on limestone.

8. Friesodielsia korthalsiana (Migq.) Steen.

(Pieter W. Korthals (1807-1892), Dutch botanist who collected widely in Malesia)

Blumea 12 (1964) 360. — Oxymitra korthalsiana Migq., Ann. Mus. Bot. Lugd.-Bat. 2

(1865) 29. Ridley, Sarawak Mus. J. 1(3) (1913) 89. Merrill, J. Straits Branch Roy. Asiat.

Soc. Spec. No. (1921) 259. Merrill, Univ. Calif. Publ. Bot. 15 (1929) 73. Masamune,

Enum. Phan. Born. (1942) 292. TYPE: Borneo, Banjoewiran, PW. Korthals s.n.

(lectotype, designated by Turner (2011b), L (barcode L 0182331)).

Oxymitra acuminata Merr., J. Straits Branch Roy. Asiat. Soc. 85 (1922) 180.

Masamune, Enum. Phan. Born. (1942) 291.— Friesodielsia acuminata (Merr.) Steen.,

Blumea 12 (1964) 357. Beaman et al., Pl. Mt. Kinabalu 4 (2001) 86. TYPE: Borneo,

Sabah, Sandakan, September-December 1920, Ramos 1567 (lectotype, designated

by Turner (2011b), BO (sheet no. BO-1349528); isolectotypes: BM, GH).

Large woody climber. Twigs black, striate or latticed, youngest with short brown

woolly tomentum, soon lost. Leaves chartaceous, drying brown, generally dark or

rich brown, above, cinnamon brown beneath, ferruginous hairy above when very

young, soon hairs confined to midrib and laterals above, more or less glabrous below,

Climbing Annonaceae of Borneo 427

lamina oblong obovate, 6—16.5(—25) x 2—5.5(—9) cm, base truncate or rounded to

slightly cordate, apex obtuse to acuminate, sometimes markedly so, midrib and lateral

nerves slightly sunken above in dry leaves, prominent beneath, 8—13 lateral pairs,

tertiary and higher-order venation distinct from both surfaces in dry leaves. Petiole

5—7 mm long, 1—2 mm thick. Inflorescence single-flowered, extra-axillary. Flower,

pedicel 1—6.5 cm long, slender, c. 0.5 mm thick, medial bract linear-lanceolate 2—3

mm long, | mm wide, hairy adaxially, sepals ovate, 4-7 x 3 mm, apex sometimes

acuminate, outer petals coriaceous, lanceolate, c. 36 * 6 mm, flat externally except for

a faint central longitudinal ridge, adpressed brown hairs, inside with central ridge and

excavated base, more or less glabrous, inner petals, ovate lanceolate, 10 x 4 mm, apex

acuminate, glabrous, faint central longitudinal ridge, stamens many, carpels many, red

hairy. Fruiting pedicel to 12 cm long, monocarps 10 or more, ellipsoidal to cylindrical,

11-17 x 7 mm, apiculate, drying black or brown with rusty pubescence, stipe 10—16

mm, less than | mm thick. Seed 1, ellipsoidal c. 10-15 « 5-6 mm.

Distribution. Endemic to Borneo, where it has been collected from Kalimantan and

Sabah.

Ecology. Lowland forest.

Notes. A number of collections from montane forest at 1000-1500 m on Mt Kinabalu

(Sabah) (Clemens & Clemens 3766, 26205, 30534, 30866, 34022, Matamin Rumutom

279) have narrow leaves and long pedicels, but may belong here.

9. Friesodielsia ovalifolia (Ridl.) 1.M.Turner

(Latin, oval-leaved)

Blumea 55 (2010) 118. — Melodorum ovalifolium Ridl., Bull. Misc. Inform. Kew

1912 (1912) 387. Ridley, Sarawak Mus. J. 1(3) (1913) 92. Merrill, J. Straits Branch

Roy. Asiat. Soc. Spec. No. (1921) 262. Masamune, Enum. Phan. Born. (1942) 287. —

Fissistigma ovalifolium (Ridl.) Merr., Philipp. J. Sci., C. 15 (1919) 134 — Oxymitra

ovalifolia (Ridl.) J. Sinclair, Sarawak Mus. J. 5 (1951) 607. — Richella ovalifolia

(Ridl.) Steen., Blumea 12 (1964) 357. TYPE: Borneo, Sarawak, near Kuching, 17

May 1894, G.D. Haviland & C. Hose 3151 (\ectotype, designated by Turner (2010b),

K (barcode no. K000574639)).

Large woody climber. Twigs drying dark grey or blackish, striate or latticed, youngest

parts densely brown woolly hairy. Leaves chartaceous, drying dark brown above, brown

or grey with brown veins below, scattered pale hairs on young leaves above, soon lost

except for along midrib, similar below but brown hairs more persistent, lamina elliptic

or elliptic oblong to slightly obovate, 2.5—8.5 = 1.5—3.5 cm, base rounded, truncate

or slightly cordate, apex emarginate, obtuse or shortly acuminate, midrib sunken

above, prominent beneath, lateral veins 7-8 pairs, flush above, prominent beneath,

428 Gard. Bull. Singapore 64(2) 2012

tertiary venation distinct from both surfaces. Petiole 4-5 mm long, to | mm thick.

Inflorescence single-flowered, extra-axillary. Pedicel 15-30 mm long, very slender,

c. 0.4 mm thick, tiny medial bract, red-brown woolly hairy; sepals free, triangular, c.

3 x 3 mm, apex acute, red-brown woolly hairy, outer petals yellow, coriaceous, c. 2

cm long, 5 mm wide, brown woolly hairy outside with slight external central ridge,

base excavated internally, glabrous inside; inner petals glabrous, ovate lanceolate, 6 =

2 mm, apex sharply acuminate; stamens many, carpels many. Fruiting pedicel c. 3 cm

long, | mm thick, monocarps to 10 or more, ellipsoidal, 10-12 x 7 mm, brown hairy,

apiculate, stipe 12-15 mm long. Seed 1, ellipsoidal, c. 9-11 x 5—6 mm.

Distribution. Endemic to Borneo where it appears restricted to the Kuching area of

Sarawak (Haviland & Hose 3141, 3151; Hewitt A.7.13, S 27795), though Church et al.

8/8 from Central Kalimantan may be this.

Ecology. Lowland forest.

MITRELLA Mia.

(diminutive of Greek, mitra = mitre, cap)

Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 38. Sinclair, Gard. Bull. Singapore 14 (1955)

364. Polyalthia section Kentia Blume, Fl. Javae Anonac. (1830) 71. Polvalthia section

Schnittspahnia Rehb., Deut. Bot. Herb.-Buch., nom. superfl. Melodorum section

Kentia (Blume) Hook.f. & Thomson, FI. Ind. (1855) 122. Unona section Kentia

(Blume) Baill., Hist. Pl. 1 (1868) 213. TYPE: Polyalthia kentii (Blume) Blume (=

Unona kentii Blume, = Mitrella kentii (Blume) Mig.)

Woody climbers. Leaves tertiary venation reticulate. Inflorescences axillary or terminal,

solitary or more rarely paired. Flowers bisexual, sepals 3, valvate, connate or not,

petals valvate, two whorls of 3, inner much shorter than outer petals, stamens many,

connectives with truncate or obtuse apex, carpels 6—15, ovules |—many, biseriate.

Fruits ellipsoidal or globose, stipitate. Seeds 1—many, smooth, shiny, sometimes pitted.

Distribution and diversity: About six species. Thailand and Malay Peninsula through

to New Guinea and Australia. Three species in Borneo.

Notes. | find it difficult to see meaningful distinctions between Mitrella and

Pyramidanthe. Mitrella dielsii appears to bridge the gap in morphological terms

between Mitrella kentii and M. clementis on the one hand, and Pyramidanthe

prismatica on the other. However I refrain from making a formal reduction feeling

a wider morphological survey and molecular anlayses are required to undertake this

step. However I include P. prismatica in the key below. I do not follow Ban (1974b) in

including Sphaerocoryne in Mitrella.

Cisnbme Amnomacca: of Bomeo 429

Key to Mitrella and Pyramidanthe species

la. Leaves when dry with secondary and intersecondary nerves clearly visible from

below, outer petals ovate, dark brown velvety outside. monocarps strongly

Te et = | ce M. dielsii

b. Leaves when dry with indistinct secondary veins from below, outer petals ovate

lanceolate, or if ovate not dark brown velvety outside, monocarps smooth

weakly muricate, not dark brown velvety -

2a. Outer petals ovate lanceolate, calyx cup with mm more or less flat or with

very broadly obtuse points, mner petals more or less glabrous, monocarps

cylindrical or ellipsoidal, more than 3 cm long, seeds smooth ..... P_ prismatica

b. Outer petals ovate or. if ovate lanceolate. calyx cup with relatively sharp

points, inner petals hairy, at least on edges, monocarps globose or PRE:

not exceeding 1.5 cm long. seeds pitted .. = Borie: ree ;

3a. Outer petals ovate, inner petals covered with hairs on the outside ...... M. kentii

b. Outer petals ovate lanceolate, inner petals with hairs confined to the edges and

USE OE sv oot SYS a eee M. clementis

1. Mitrella clementis (Merz.) 1.M-Tumer

(Latin, clementis (clemens in genitive case) = of gentility, mildness, mercy:

commemorative of Joseph Clemens (1862—1936), a Methodist minister, originally

from England. who served with the US Amny, and his American wife Mary Strong

Clemens (1873—1968), who together collected plants professionally).

Malayan Nat. J. 61 (2009) 273. — Fissistigma clementis Merr., J. Straits Branch Roy.

Asiat. Soc. 85 (1922) 178. Memill, Univ. Calif. Publ. Bot. 15 (1929) 71. Masamune.

Enum. Phan. Bom. (1942) 284. TYPE: Bomeo. Sabah, Sandakan and vicinity,

September—December 1920, M. Ramos 1474 (lectotype. designated by Tumer (2009c),

K (barcode no. K000574737); isolectotypes: A[*2], BM. L, US).

Large woody climber. Twigs drying black or dark brown, stmate or latticed_ very

youngest parts with some sparse red-brown hairs. Leaves chartaceous. drying grey-

brown above, matt middle brown to pale grey-brown beneath, sparse small adpressed

hairs on lamina below, glabrous above. midnb slightly sunken above in dry leaves.

prominent beneath, laterals flush on both surfaces, lamina elliptic to oblong lanceolate.

3.S-11 = 1.5-3.5 cm, base obtuse, apex acuminate, 10—12 pairs of laterals. arching

forward, looping obscurely, reticulations relatively indistinct. Petiole 3-12 mm long.

0.5-1 mm thick. Inflorescences single-flowered or more rarely im pairs, axillary.

Flower, pedicel 5-7 mm long. c. 1 mm thick, with golden brown adpressed hairs.

sepals connate, c. 2 x 3 mm, apiculate, outer petals, ovate lanceolate, 15-25 = 6

mm outside covered with short brown or pale hairs. inside excavated. very short

430 Gard. Bull. Singapore 64(2) 2012

pale woolly hairs, excavation glabrous; inner petals ovate lanceolate, c. 5 x 2.5 mm,

glabrous externally except for a very short pale wool along the margins and near the

apex, stamens many, carpels many. Fruiting pedicel to 10 mm long, 1—2 mm thick,

calyx persistent, monocarps to 8 or more, globose, ellipsoidal or rather irregular, 8—12

x 6-10 mm, drying brown sometimes minutely wrinkled, stipe, generally shorter than

the seed-bearing portion of the monocarps, 3—4 mm long, c | mm thick. Seeds c. 4,

discoid or hemispherical, c. 6-7 mm diameter, drying dark brown, shiny, pitted.

Distribution. Endemic to Borneo where it has been collected from Brunei, Kalimantan,

Sabah and once from Sarawak (S 52509).

Ecology. Lowland forest, once from 1800 m in Sabah.

Notes. This species has been consistently confused with M. kentii. The flowers of the

two species differ markedly in the shape of the outer petals. Mitrella kentii generally

has more coriaceous leaves, but some collections have narrow, chartaceous leaves

almost indistinguishable from M. clementis. The fruits are also very similar though

M. clementis seems to have somewhat larger, but relatively shorter-stiped, monocarps

with generally more seeds.

2. Mitrella dielsii J.Sinclair

(F.L.E. Diels (1874-1945), German botanist)

Gard. Bull. Singapore 15 (1956) 14. Bygrave in Coode et al., Check]. Fl. Pl. Gymnosp.

Brunei (1996) 18. — Melodorum beccarii Diels, Notizbl. Bot. Gart. Berlin-Dahlem

11 (1931) 85, non Melodorum beccarii Scheff. (1885). TYPE: Borneo, Sarawak,

near Sungai Igan, October 1867, O. Beccari PB. 3899 (holotype: FI-B; isotype:

B(fragment)).

Large woody climber. Twigs drying very dark brown, uniform in colour, longitudinally

striate, very youngest parts with dense, very short, curly dark red-brown tomentum.

Leaves coriaceous, drying dark brown shiny above, matt brown beneath with darker

redder-brown midrib and laterals, lamina hairy below, usually sufficiently so to give

a felty feel, hairs curly, brown or red-brown, densest along nerves, expanding leaves

with pale hairs above, soon lost, midrib slightly sunken above in dry leaves, prominent

below, laterals flush above, raised slightly beneath, lamina ovate, ovate lanceolate or

oblong elliptic, 5-19 x 2.5~7 cm, base obtuse, rounded or truncate, apex acuminate,

lateral veins 10-15 pairs, arching forward, looping indistinctly, reticulations visible

from both surfaces in dry leaves. Petiole 12—16 mm long, c. 2 mm thick. Inflorescences

single-flowered, axillary. Pedicel 5-6 mm long, c. 2 mm thick, densely covered with

curly brown hairs, sepals connate, broadly triangular, 2—3 = 5 mm, hairy outside, outer

petals coriaceous, ovate, 16-19 x 9-12 mm, basally excavate on the inside, externally

dark brown velutinous, internally covered with very short pale grey woolly hairs

Climbing Annonaceae of Borneo 431

except the glabrous excavation, inner petals concave, ovate, 5 x 3-4 mm, drying dark

brown, more or less glabrous, stamens many, c. 1.5 mm long, apex of connective acute,

carpels c. 10, c. 2 mm long, drying black, glabrous, stigma very short. Fruiting pedicel

to 10 mm long, 6 mm thick, calyx persistent, monocarps to 9 or more, ellipsoidal, c.

3.5 x 2.5 cm, strongly muricate, densely covered with very short curly dark brown

hairs, stipe to 7 mm long, 4 mm thick. Seeds to 12 or more, 2 rows, drying smooth,

brown, flat, 11-13 x 8 x 24 mm.

Distribution. Endemic to Borneo where it is known from Brunei and Sarawak.

Ecology. Apparently restricted to peatswamp forest.

3. Mitrella kentii (Blume) Miq.

(William Kent, 1779-1827, Dutch gardener, first curator of the Botanic Garden in

Buitenzorg, Java)

Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 39. Merrill, J. Straits Branch Roy. Asiat. Soc.

Spec. No. (1921) 260. Masamune, Enum. Phan. Born. (1942) 289. Sinclair, Gard.

Bull. Singapore 14 (1955) 365. Kessler & van Heusden, Rheedea 3 (1993) 71.

Bygrave in Coode et al., Checkl. Fl. Pl. Gymnosp. Brunei (1996) 18. Beaman et al.,

Pl. Mt. Kinabalu 4 (2001) 91. Unona kentii Blume, Biyjdr. (1825) 16. — Polyalthia

kentii (Blume) Blume, Fl. Javae Anonac. (1830) 77. — Melodorum kentii (Blume)

Hook.f. & Thomson, FI. Ind. | (1855) 116. Ridley, Sarawak Mus. J. 1(3) (1913) 93.

— Fissistigma kentii (Blume) Merr., Philipp. J. Sci., C. 15 (1919) 132. TYPE: Java,

Anon. s.n. (lectotype, designated by Turner (2011b), L (ex herb. Blume) (barcode no.

L 0183051)).

[Uvaria elegans Wall., Numer. List (1832) no. 6474, nom. nud.]

Uvaria mabiformis Griff., Notul. 4 (1854) 709. — Fissistigma mabiforme (Griff.)

Merr., Philipp. J. Sci., C. 15 (1919) 133. TYPE: Peninsular Malaysia, Malacca, Aloor

Gajah, Verupha s.n. [Kew Distib. no. 389] (lectotype, designated by Sinclair (1955),

K(barcode no. K000574743)).

Melodorum elegans Hook.f. & Thomson, FI. Ind. 1 (1855) 122. — Fissistigma elegans

(Hook.f. & Thomson) Merr., Philipp. J. Sci., C. 15 (1919) 131. TYPE: Peninsular

Malaysia, Penang, 1822, Anon. [N. Wallich] s.n. [EIC 6474A] (lectotype, designated

by Turner (2011b), K ex herb. Hook. (barcode no. K000574739); isolectotypes: C,

CALYCGE, E, GZU, K, K-W, L; NY, PH).

Melodorum pisocarpum Hook.f. & Thomson, FI. Ind. 1 (1855) 123. TYPE: Peninsular

Malaysia, Malacca, s. dat., W. Griffith s.n. (lectotype, designated by Turner (2011b), K

ex herb. Hook. (barcode no. K000574741)).

432 Gard. Bull. Singapore 64(2) 2012

Orophea borneensis Mig., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 26. Ridley, Sarawak

Mus. J. 1(3) (1913) 87. Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921)

265. Masamune, Enum. Phan. Born. (1942) 290. TYPE: Borneo, W.H. de Vriese s.n.

(lectotype, designated by Kefiler (1988), L (barcode no. L 0038052; isolectotype: L).

Large woody climber. Twigs drying blackish, dark brown or dark grey, sometimes with

shallow longitudinal wrinkles or latticing, youngest parts with red-brown or pale hairs.

Leaves chartaceous to subcoriaceous, drying pale grey brown to almost black above,

uniform matt brown, midrib generally rather pale brown, glabrous above, below with

tiny adpressed, red-brown or pale hairs, hairy on lamina and along nerves, midrib very

slightly sunken below, raised beneath, lateral nerves flush above, very slightly raised

beneath, lamina elliptic. ovate or ovate lanceolate, S—13 = 2.5—6 cm, base obtuse,

apex shortly acuminate, lateral nerves 9-13 pairs, arching forward, looping obscurely,

reticulations visible from both surfaces in dry leaves. Petiole 8-16 mm long, 1-2 mm

thick. Inflorescences 1—3-flowered, axillary. Pedicel 5—7 mm long, c. 1 mm thick,

widening distally, drying dark brown with decumbent red-brown or brown hairs,

medial bract absent, sepals connate, c. 2 x 3 mm, forming shallow triangular dish,

externally red-brown hairy, glabrous within, outer petals coriaceous, ovate to elliptic,

9-11 x 5—9 mm long, externally very densely covered with very short red-brown hairs,

inside very short pale grey woolly except lower part of excavation, inner petals concave

ovate, c. 5 x 2.5 mm, pale grey woolly outside, glabrous within except for long pale

hairs from upper part, stamens many, I—1.5 mm long, carpels many, glabrous. Fruiting

pedicel 7-12 mm long, 1.4—2 mm thick, calyx persistent, monocarps to 12, globose or

irregular, c. 7 mm diameter, drying brown, relatively smooth, glabrous, apex typically

rounded, stipe 4-7 mm long, c. | mm thick. Seeds two or more, hemispherical, pitted,

drying brown, c. 5 mm diameter.

Distribution. Malay Peninsula, Java and Borneo. In Borneo recorded from Brunei,

Kalimantan, Sabah and Sarawak.

Ecology. Lowland and heath forest to 300 m.

Notes. See M. clementis for distinctions with that species and Sphaerocoryne affinis

likewise.

PYRAMIDANTHE Mica.

(Greek, pyramis, pyramidos = pyramid, anthos = flower)

Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 39. Sinclair, Gard. Bull. Singapore 14 (1955) 362.

Unona section Pyramidanthe (Migq.) Baill., Hist. Pl. 1 (1868) 213. Melodorum section

Pyramidanthe (Miq.) Kurz, J. As. Soc. Bengal 43 (1870) 56. TYPE: Pyramidanthe

rufa Miq.

Climbing Annonaceae of Borneo 433

Woody climbers. Leaves coriaceous, lateral nerves distinct, tertiary venation laxly

reticulate. Flowers axillary. Sepals 3, valvate, united in a flat disc, petals in 2 whorls

of 3, outer triquetrous, much longer than inner, stamesn with a truncate, dilated apex,

carpels 5—6. Monocarps stipitate, tuberculate. Seeds in 2 rows, shiny.

Distribution and diversity: One species, Pyramidanthe prismatica (Hook.f. &

Thomson) Merr., found in Malay Peninsula, Sumatra and Borneo.

Notes. See comments under Mitrel/a regarding status of the genus and for a key

including P. prismatica.

1. Pyramidanthe prismatica (Hook.f. & Thomson) Merr.

(Latin, having several longitudinal angles and intermediate flat surfaces)

J. Straits Branch Roy. Asiat. Soc., Spec. No. (1921) 262. Masamune, Enum. Phan.

Born. (1942) 296. Sinclair, Gard. Bull. Singapore 14 (1955) 362. Bygrave in Coode

et al., Checkl. Fl. Pl. Gymnosp. Brunei (1996) 22. Beaman et al., Pl. Mt. Kinabalu

4 (2001) 97. Melodorum prismaticum Hook.f. & Thomson, FI. Ind. (1855) 121.

Ridley, Sarawak Mus. J. 1(3) (1913) 88. — Fissistigma prismaticum (Hook.f. &

Thomson) Merr., Philipp. J. Sci., C. 15 (1919) 135. TYPE: Peninsular Malaysia,

Penang, August 1822, N. Wallich s.n. [EIC 6455] (lectotype, designated by Turner

(2011b), K-W; isolectotypes: BM, CGE).

[Uvaria prismatica Wall., Numer. List (1832) no. 6455, nom. nud.]

Oxymitra bassiifolia Teijsm. & Binn., Natuurk. Tijdschr. Ned.-Indie 25 (1863) 419, as

‘bassiaefolia’. TYPE: Bangka, near Planyas, s.dat., J.E. Teijsmann s.n. [Herb. Bogor.

no. 17645] (lectotype, designated by Turner (2011b), BO (sheet no. BO-1408032);

isolectotype: BO (sheet no. BO-1408033)).

Pyramidanthe rufa Mig., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 39. TYPE: Borneo

australis in Martapoera; Sumatra occidentalis: Korthals

Melodorum cylindricum Maingay ex Hook.f. & Thomson, FI. Brit. Ind. 1 (1872) 80.

Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921) 261. Masamune, Enum.

Phan. Born. (1942) 284. — Fissistigma cylindricum (Maingay ex Hook.f. & Thomson)

Merr., Philipp. J. Sci., C. 15 (1919) 131. Merrill, Univ. Calif. Publ. Bot. 15 (1929) 71.

TYPE: Peninsular Malaysia, Malacca, 1865-1866, 4.C. Maingay 1507 [Kew Distrib.

no. 78] (holotype: K (barcode no. K000574661); isotype: CAL).

Melodorum maingayi Hook.f. & Thomson, FI. Brit. Ind. 1 (1872) 80. Ridley, Sarawak

Mus. J. 1(3) (1913) 90. Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921)

262. Masamune, Enum. Phan. Born. (1942) 287. — Fissistigma maingayi (Hook.f.

434 Gard. Bull. Singapore 64(2) 2012

& Thomson) Merr., Philipp. J. Sci., C. 15 (1919) 133. TYPE: Peninsular Malaysia,

Penang, s.dat., 4.C. Maingay s.n. [Kew Distrib. no. 108] (holotype: K (barcode no.

K000574660)).

Pyramidanthe rufa var. parvifolia Boerl., Icon. Bogor. 1 (1899) 131, t. 44. —

Pyramidanthe prismatica vat. parvifolia (Boerl.) Merr., J. Straits Branch Roy. Asiat.

Soc., Spec. No. (1921) 263. Masamune, Enum. Phan. Born. (1942) 297. TYPE: Borneo,

Sarawak, nr Kuching, 1892, G.D. Haviland 421 (lectotype, designated by Turner

(2011b), BO (sheet no. BO-134059)).

Melodorum rigidum Ridl., Bull. Misc. Inform. 1912 (1912) 386. Ridley, Sarawak

Mus. J. 1(3) (1913) 91. Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921)

262. Masamune, Enum. Phan. Born. (1942) 288. — Fissistigma rigidum (Ridl.) Merr.,

Philipp. J. Sci., C. 15 (1919) 135. Sinclair, Sarawak Mus. J. 5 (1951) 600. TYPE:

Borneo, Sarawak, near Kuching, G.D. Haviland 421 [Garai leg.] [d.k.m.a.] (lectotype,

designated by Turner (2012), K (barcode no. K000574656)).

Large woody climber. Twigs drying black or dark brown, with shallow longitudinal

wrinkling or latticing, sometimes with raised brown lenticels, youngest parts with dark

brown adpressed tomentum. Leaves chartaceous to coriaceous, drying dark brown to

almost black above, brown to pale brown below with darker midrib and lateral veins,

midrib slightly sunken above in dry leaves, prominent beneath, lateral veins very

slightly raised on both surfaces, lamina glabrous above, variably hairy below from tiny

adpressed to dense erect hairs, lamina oblong elliptic to obovate, 8—23.5 = 3.5—8 cm,

base obtuse to rounded, apex acuminate, lateral veins 10—14 pairs, arching forward,

looping indistinctly. Petiole 9-20 mm long, 2—3 mm in diameter. Inflorescences axillary.

Pedicel 7-10 mm long, c. 1.5 mm thick, widening distally, red-brown adpressed hairy,

tiny medial bract near the base of the pedicel, sepals connate, c. 2 x 5 mm, spreading,

forming an almost circular salver under the corolla, externally brown or red-brown hairy,

outer petals ovate lanceolate, to 40 x 8 mm, outside covered with dense short golden

brown hairs, inside very short pale grey woolly except for lower part of excavation

which is glabrous, inner petals ovate lanceolate, c. 7 x 4 mm, glabrous except for a

few pale hairs near apex externally, drying brown, minutely wrinkled externally, more

deeply wrinkled internally, stamens many, 1—1.5 mm long, connective apex polygonal,

carpels many c. 2.5 mm long, sparsely hairy, stigma frilly. Fruiting pedicel to 10 mm

long, 5 mm thick, calyx persistent, monocarps to 20 or more, often fewer, globose,

ellipsoidal or cylindrical, 3.5—4.5 x 1.5—2.5 cm, drying relatively smooth to muricate,

sometimes covered with short golden brown tomentum, stipe 8-10 < 2-4 mm. Seeds

many, 2 rows, drying smooth, shiny dark brown, c. 10-12 = 6 x 2-4.

Distribution. Malay Peninsula, Sumatra and Borneo. In Borneo widespread, recorded

from Brunei, Kalimantan, Sabah and Sarawak.

Ecology. Lowland forest.

Climbing Annonaceae of Borneo 435

SPHAEROCORYNE (Scheff. ex Boerl.) Scheff. ex Ridl.

(Greek, sphaera = ball, globe, sphere, coryne = club)

J. Straits Branch Roy. Asiat. Soc. 75 (1917) 8. Polyalthia section Sphaerocoryne

Scheff. ex Boerl., [Cat. Pl. Phan. 1 (1899) 26, nomen] Icon. Bogor. 1 (1899) 196.

TYPE: Polyalthia siamensis Boerl. (= Sphaerocoryne siamensis (Boerl.) Ridl.).

Unona subgenus Mesnya Pierre, Fl. Forest. Cochinch. (1881) t. 17. TYPE: Unona

mesnyi Pierre, nom. illegit. (= Polyalthia aberrans Maingay ex Hook.f. & Thomson).

Woody climbers or scandent shrubs, sometimes erect shrubs. Twigs glabrous or hairy.

Leaves chartaceous to coriaceous. Inflorescences |- or 2-flowered, terminal or axillary.

Flowers bisexual, sepals 3, valvate more-or-less connate at the base, persisting in

fruit, petals valvate in two whorls of 4, inner only slightly smaller than outer, stamens

numerous, connective truncate, carpels numerous, ovules 1—2. Fruits monocarps many,

stipitate. Seeds 1-2.

Distribution and diversity: Indo-China to the Philippines. Three species of which one

species, Sphaerocoryne affinis, recorded from Borneo.

1. Sphaerocoryne affinis (Teijsm. & Binn.) Ridl.

(Latin, neighbouring, akin to)

J. Straits Branch Roy. Asiat. Soc. 75 (1917) 8. — Polyalthia affinis Teijsm. & Binn.,

Natuurk. Tijdschr. Ned.-Indié 27 (1864) 37. TYPE: Java, cult. in Hort. Bot. Bogor.,

s.dat., Anon. s.n. [Bogor. distrib. no. 113] (neotype, designated by Turner (2011b), L

(barcode no. L0197456)).

Monoon submitratum Miq., Ann. Mus. Lugd. Bat. 2 (1865) 16. — Polyalthia

submitrata (Migq.) Ridl., Sarawak Mus. J. 1(3) (1913) 82. Merrill, J. Straits Branch

Roy. Asiat. Soc. Spec. No. (1921) 257. Masamune, Enum. Phan. Born. (1942) 295.

TYPE: Borneo, Moeara Karrau, PW. Korthals s.n. (lectotype, designated by Turner

(2011b), L (barcode no. L 0184462)).

Polyalthia aberrans Maingay ex Hook.f. & Thomson, Fl. Brit. Ind. 1 (1872) 67. —

Unona mesnyi Pierre, Fl. Forest. Cochinch. (1881) t. 17, nom. superfl. — Popowia

mesnyi Craib, Bull. Misc. Inform. Kew 1914 (1914) 5, nom. superfl. — Mitrella

mesnyi (Craib) Ban, Bot. Zhurn. 59(2) (1974) 244, nom. superfl. — Popowia aberrans

(Maingay ex Hook.f. & Thomson) Pierre ex Finet & Gagnep., Bull. Soc. Bot. Fr. Mem.

4(2) (1906) 109. — Sphaerocoryne aberrans (Maingay ex Hook.f. & Thomson) Ridl.,

J. Straits Branch Roy. Asiat. Soc. 75 (1917) 8. — Melodorum aberrans (Maingay ex

Hook.f. & Thomson) J. Sinclair, Gard. Bull. Singapore 14 (1953) 41. Sinclair, Gard.

Bull. Singapore 14 (1955) 332. — Mitrella aberrans (Maing. ex Hook.f. & Thomson)

436 Gard. Bull. Singapore 64(2) 2012

Ban, Bot. Zhurn. 59(2) (1974) 244. TYPE: Peninsular Malaysia, Malacca, 28 April

1868, 4.C. Maingay 3141 [Kew distrib. no. 110] (lectotype, designated by Turner

(2011b), K (barcode no. K000380043)).

Melodorum clavipes Hance, J. Bot. 15 (1877) 328. — Sphaerocoryne clavipes (Hance)

Craib, Bull. Misc. Inform. Kew 1922 (1922) 168. TYPE: Cambodia, s.dat., L. Pierre

19770 (holotype: BM (barcode no. BM000554028)).

Polyalthia siamensis Boerl., Cat. Pl. Phan. (1899) 26. — Sphaerocoryne siamensis

(Boerl.) Ridl., J. Straits Branch Roy. Asiat. Soc. 75 (1917) 8. TYPE: Java, cultivated

in Hort. Bot. Bogor. sub XI.A.71 and XI.A.41.

Dasymaschalon scandens Merr., Philipp. J. Sci., C. 10 (1915) 238. Merrill, Enum.

Philipp. Fl. Pl. 2 (1923) 175. TYPE: Philippines, Palawan, Taytay, May 1913, E.D.

Merrill 9277 (lectotype, designated by Turner (2011b), K (barcode no. K0006911800)).

Tree or scandent shrub. Twigs drying brown, shallowly striate or latticed, glabrous but

for a few pale hairs on the youngest parts. Leaves chartaceous, glaucous beneath, drying

brown or grey-brown above, sometimes shiny, paler and duller beneath with midrib

a contrasting red-brown, glabrous above, below with a scattering of pale adpressed

hairs, sometimes almost glabrous, midrib flush above in dry leaves, prominent beneath,

lateral nerves slightly raised on both surfaces, lamina elliptic, 4-14.5 « 1.5—S cm, base

acute, apex acuminate, lateral veins 14—18 pairs, tertiary venation clearly visible from

both surfaces. Petioles 4-8 mm long, c. | mm thick. Inflorescences single-flowered,

axillary. Flowers pedicel 15-16 mm long, c. | mm thick, widening distally, drying

brown, striate, more or less glabrous, medial bract c. 1.5 x 1 mm, with sparse pale

hairs, perianth sometimes 4-merous, sepals slightly connate at base, broadly ovate, c.

2 x 4 mm, drying black, sparse, short adpressed pale hairs outside, glabrous within,

petals rather thick, outer petals broadly ovate, c. 7 x 6 mm, covered with short dense

pale hairs, glabrous near base inside, inner petals c. 6 x 5 mm, hairy outside, more or

less glabrous within, stamens many, carpels many, I|—1.5 mm long, white hairs near

apex. Fruits pedicel 2—2.5 cm long, c. | mm thick, widening distally, calyx persistent,

monocarps to 25 or more, ellipsoidal, typically drying 8—9 x 5—6 mm, 2-seeded fruits

to 13 mm long, minutely beaked, drying dark red-brown, minutely pimpled, glabrous

except for a few short pale hairs near the apex, stipe very slender, 8—14 mm long, c.

0.5 mm thick. Seeds 1, rarely 2, ellipsoidal, c. 8 x 6 mm, drying smooth, pale brown.

Distribution. Indo-China, Thailand, Malay Peninsula, ?Java, Borneo and_ the

Philippines. In Borneo recorded from Sabah and once from Kalimantan (type of

Polyalthia submitrata).

Ecology. Lowland forests to 200 m altitude.

Notes. This species has generally been overlooked among material from Borneo.

Climbing Annonaceae of Borneo 437

Mostly collected in fruit, specimens have generally been confused among a group of

superficially similar small-leaved species including Mitrella kentii, Demos acutus or

D. dunalii, and Uvaria micrantha. Mitrella kentii and M. clementis have adpressed

hairs on the lower lamina unlike S. affinis which has a very sparse indumentum. The

stipes of the monocarps also appear less slender in the Mitrella species. The Desmos

species have moniliform monocarps, but in some poor specimens care must be taken

to identify fruits with more than one seed. Uvaria micrantha has stellate pubescence

but the hairs are small and may be difficult to see in the field. The confusable Desmos

and Uvaria species do not have the calyx persisting in fruit like Sphaerocoryne (and

Mitrella).

UVARIA L.

(Latin, wva = cluster, cluster or bunch of grapes; alluding to clustered fruits)

Sp. Pl. (1753) 536. Sinclair, Gard. Bull. Singapore 14 (1955) 199. Zhou et al., Syst.

Biodivers. 7 (2009) 249-258. Zhou et al., Bot. J. Linn. Soc. 163 (2010) 33-43.

LECTOTYPE: Uvaria zeylanica L. (designated by Hutchinson (1923)).

Marenteria Noronha ex du Petit-Thouars, Gen. Nova Madag. (1806) 18. TYPE: non

designatus.

Cyathostemma Griff., Notul. Pl. Asiat. 4 (1854) 707. Utteridge, Blumea 45 (2000)

377. TYPE: Cyathostemma viridiflorum Griff.

Ellipeia Hook.f. & Thomson, Fl. Ind. (1855) 104. Sinclair, Gard. Bull. Singapore 14

(1955) 230. TYPE: Ellipeia cuneifolia Hook.f. & Thomson.

Anomianthus Zoll., Linnaea 29 (1858) 324. TYPE: Anomianthus heterocarpus (Blume)

Zoll.

Tetrapetalum Mig., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 23. TYPE: Tetrapetalum

volubile Miq.

Rauwenhoffia Scheff., Ann. Jard. Buitenz. 2 (1885) 21. TYPE: Rawwenhoffia siamensis

Scheff.

Uva Kuntze, Rev. Gen. 1 (1891) 7, nom. superfl.

Uvariella Ridl., Fl. Malay. Penins. 1 (1922) 22. TYPE: Uvariella leptopoda (King)

Ridl.

Ellipeiopsis R.E. Fr., Ark. Bot. ser. 2. 3 (1955) 41. TYPE: Ellipeiopsis ferruginea

(Buch.-Ham. ex Hook.f. & Thomson) R.E. Fr.

438 Gard. Bull. Singapore 64(2) 2012

Dasoclema J. Sinclair, Gard. Bull. Singapore 14 (1955) 273. TYPE: Dasoclema

siamensis (Craib) J. Sinclair.

Balonga Le Thomas, Adansonia sér. 2, 8 (1968) 106. TYPE: Balonga buchhoizii (Eng

& Diels) Le Thomas.

Woody climbers with stellate or caespitose hairs. Twigs glabrous or stellate hairy.

Leaves chartaceous to coriaceous. Inflorescences fascicles or solitary flowers, terminal,

leaf-opposed, rarely cauliflorous. Flowers bisexual, trimerous, rarely dimerous, sepals

valvate, often connate, petals imbricate in two whorls, sometimes connate at the base,

stamens numerous, outer sometimes sterile, truncate, carpels few to many, ovules

few to many, uni- or biseriate. Fruits few to many stipitate, subglobose to ellipsoidal

monocarps. Seeds few to many, in 2 rows.

Distribution and diversity: More than 100 species across the Old World tropics from

‘Africa to Australia. 19 species recorded from Borneo.

Notes. For both nomenclatural and taxonomic reasons, Utteridge (2000) reduced

Tetrapetalum to Uvaria. Recent molecular phylogenies have provided the critical

weight for a number of other satellite genera to gravitate into an enlarged Uvaria (Zhou

et al. 2009, 2010). Most relevant to the student of Bornean Annonaceae, Cyathostemma

and Ellipeia have been reduced to Uvaria, unfortunately with many epithet changes.

Key to Uvaria species

la. Plants cauliflorous with inflorescences arising from main stem ................ 2

b.. Plants moticauwlitlorous csc fiecce accceeee eee eee ee ee 3

2a. Calyx entirely enclosing flower in bud, petals reflexing at anthesis and more

than 1 cm long, nionocarps drying brownie. sete eee U. monticola

b. Calyx not entirely enclosing flower in bud, petals not reflexing at anthesis,

less than 1 cm long, monocarps drying black .................csscceseeees U. griffithii

3a. Inflorescences, generally arising on branches behind leaves, congested

cymes of 4—20)(Gub)sessillediowiens sc... eeere eee eee U. excelsa

b. Inflorescences, generally arising among leaves, not congested cymes,

flowers distinctly pedicellate: xcs .cc.ces trae tenes eee areata 4

4a. Inner petals distinctly shorter than outer petals, monocarps smooth, stipitate

with lateral attachment of Stipe: amd apiCulits yes cgectnees-ecnene tae cesce seers se eeeeee 5

b. Petal whorls equal or subequal in length, monocarps sessile or stipitate with

more or less basal attachment, apiculus terminal or absent .................:000 6

Climbing Annonaceae of Borneo 439

Sa. Lateral nerves looping obscurely, lower lamina not distinctly rough to the

louchmonocatps less than 2 em longs. s..2cc kc. ceces. U. cuneifolia

b. Lateral nerves looping distinctly within the margin of the leaf, lower lamina

distinctly rough to the touch, monocarps more than 2 cm long .. U. schefferi

pasemlenvinedcaves Brow mM Deneathies....c....c6s0.1c.0k eds bacco seebiees U. lanuginosa

ieee isiyane leaves nob. brown DEMEAE 0. ....).6. Asics ccectsecdtcscncectestcee eveenesseee: 7

P aaa eaves SCaDrouS ON UPPEF SUILACE <...........:cc..sccceseseccccsecseessonondeedees U. javana

PERE CAVES TIOUSEADROUS SAD OVE \<..o0.se.ccse2s0acse0cancieeeetsstssnsassvotevaceshvsesstesubovessseneeys 8

See IPS CMSA COVELEH WIE HAlfS ......:.5.<s.<.0c22<<-s2-c0+--0502<seance eneesesnezeceseeveeese 9

IPS PL ADFOUS OF WAN SPATSC NAILS 22. .<.c.<c..0scd0ceese-voseesiesecdabsvtssossessconsevee 1]

9a. Twigs and leaves pilose, with long (1—3 mm) erect, straight red brown hairs

nec aches SAGAS ELSE Se SES eee U. hirsuta

10a. Flowers more than 5 cm across, monocarps cylindrical with longitudinal

PAS ere eae ea eies 8s vise Bat oe on reves hesaiedanteucse eee. CG Brandifiora

b. Flowers less than 3 cm across, monocarps globose ............ U. curvistipitata

lla. Flowers dimerous, monocarps stipitate, with stipe shorter than seed-bearing

portion, cylindrical, rough to the touch but without longitudinal ridges .....

suson dgnsechs tages oe be Spe Cee SSeS U. borneensis

b. Flowers trimerous, monocarps (sub)sessile or if stipitate then globose or

ellipsoidal, or 1f cylindrical then with stipe longer than seed-bearing portion

SieAnine IONE TUGIHAl MOSES,» Hen Cue. cathe. ACen ete ed ace 12

12a. Petals more than 2 cm long, erect at anthesis, monocarps (sub)sessile and

muricate, with a distinct lateral apiculus .................::::cceeeeeeeee U. verrucosa

b. Petals spreading or relexing at anthesis, or if not spreading or reflexing then

less than 1 cm long, monocarps distinctly stipitate and not muricate, or if

sessile and muricate (U. beccarii) then without a lateral apiculus ......... 13

13a. Petals spreading at anthesis and at least 10 mm long, monocarps muricate,

or if not muricate cylindrical and drying black, or globose—ellipsoidal with

Sofi oecr eam pa Ken eRe hcl Ae INES Sm cenece ede ehie, OR eaves sbi eaets she ake 14

b. Petals not spreading and less than 10 mm long, monocarps not muricate, if

monocarps cylindrical then drying brown, or globose and ellipsoidal with

Ven SMendeistipec.0.s mimiin diameters. /2i215..800: tassel eel ieee G7

14a. Petals white or yellow, monocarps sessile and muricate ............ U. beccarii

b. Petals red, monocarps distinctly stipitate, muricate Or NOt .............: eee 15

440 Gard. Bull. Singapore 64(2) 2012

15a. Calyx entirely covering flower in bud, petals connate at base, monocarps

cylindrical, smooth, drying blacks: pee. see ere eee eee U. concava

b. Calyx not entirely covering flower in bud, petals free at base, monocarps

globose torvellipsoidal} drying brownies ecee cee eee eee 16

l6a. Leaves with lateral nerves looping indistinctly in dry leaves, pedicel of

flower 10 mm or less in length, monocarps muricate............. U. lobbiana

b. Leaves with lateral nerves looping distinctly in dry leaves, pedicel of flower

15 mm long or more, monocarps not muricate ....................... U. littoralis

17a. Leaves typically less than 4.5 cm wide, stipe of monocarp very slender (c.

OLS mind Alani.) 25. Sy ciao cecdeooa ae oece oo ee et ees ace eee EE EO MEPDLEGR CRITI

b. Leaves typically more than 4.5 cm wide, stipe of monocarp relatively thick

(2 PIM OF MOLE) ake ares ese etake eee eee ee eee 18

18a. Inflorescence subtended by a persistent foliose bract, monocarps with stipe

shorter thanseed=beanine portions eer cee eee ene U. argentea

b. Inflorescence bract not foliose, monocarps with stipe longer than seed-

Dearie POFWOM s.cca.cc-ceceussncteec essere tee eee ee eet ae ORL EIICTIIES

1. Uvaria argentea Blume

(Latin, silvery)

Fl. Javae Anonaceae (1830) 24. Ridley, Sarawak Mus. J. 1(3) (1913) 73. Merrill, J. Straits

Branch Roy. Asiat. Soc. Spec. No. (1921) 253. Masamune, Enum. Phan. Born. (1942)

297. — Uva argentea (Blume) Kuntze, Rev. Gen. Pl. 1 (1891) 7. — Cyathostemma

argenteum (Blume) J. Sinclair, Sarawak Mus. J. 5 (1951) 599. Sinclair, Gard. Bull.

Singapore 14 (1955) 220. Utteridge, Blumea 45 (2000) 382. TYPE: Java, J.C.A. van

Hasselt s.n. (holotype: L).

Uvaria bracteata Roxb., Fl. Ind. (1832) 660. — Uva bracteata (Roxb.) Kuntze, Rev.

Gen. Pl. 1 (1891) 7. TYPE: Roxb. Icon. 2290 (lectotype, designated by Utteridge

(2000), K).

Uvaria gomeziana A. DC., Mem. Soc. Phys. Genév. 5 (1832) 203. TYPE: Burma,

Tavoy, 8 September 1827, W. Gomez 197 [N. Wallich 1279, EIC 6459] (lectotype,

designated by Turner (2011a), K-W).

[ Cyathostemma nitidum Bakh.f., Blumea 12 (1963) 61, nom. inval. |

Woody climber to 5 m long. Twigs densely pubescent when young, becoming glabrous

with age. Leaves chartaceous to subcoriaceous, glabrous except for midrib above,

Climbing Annonaceae of Borneo 44]

oblong-lanceolate, 10-15 = 4.5—6 cm, base cuneate or rounded, apex acute, lateral

nerves 12-14 pairs. Petioles 3—5 mm long, c. 1 mm thick. Inflorescences, 1-4 flowered,

opposite leaves, basal bract foliose, orbicular, 7-9 x 7-9 mm, sepals coriaceous, broadly

ovate, 2 x 3 mm, apex obtuse, densely pubescent, petals broadly ovate, 5 mm long,

stamens many, carpels many. Monocarps oblong-ellipsoid, c. 4 x 3 cm, apex rounded,

drying slightly constricted between seeds, tuberculate with longitudinal ridges, stipe to

2.5 cm long, c. 5 mm thick. Seeds more than 6 in 2 rows.

Distribution. Bangladesh and Burma to Java and Borneo. In Borneo sparsely collected

from Kalimantan, Sabah and Sarawak.

Ecology. Lowland forest.

2. Uvaria beccarii Attan., 1.M.Turner & R.M.K.Saunders

(Odoardo Beccari (1843-1920), Italian naturalist and author of Ne//e Foreste di Borneo)

Novon 21 (2011b) 161, Fig. 1. TYPE: Borneo, Sarawak, Mt Matang, 1863-1865, O.

Beccari P.B. 1648 (holotype: K; isotypes: BO, FI-B).

Large woody climber. Twigs dark grey or grey-brown, striate or latticed, young twigs

densely brown stellate hairy. Leaves chartaceous to subcoriaceous drying dark brown,

brown or grey-brown above, dark brown to brown below, midrib slightly sunken above

in dry leaves, prominent beneath, lateral nerves more or less flush above, prominent

beneath, glabrous above except for hairs on midrib above, densely stellate hairy

on nerves below, with scattered hairs on lower lamina. lamina elliptic to narrowly

oblanceolate, 6-14 x 2—5.5 cm, base obtuse, truncate or rounded, apex acuminate,

lateral nerves 11—14 pairs, looping obscurely well within the margin, tertiary venation

areolate, visible from below, indistinct from above. Petiole 3-5 mm long, 1—2 mm thick.

Inflorescence subopposite leaves, single-flowered. Flower pedicel 12—22 mm long, |

mm thick, densely brown stellate hairy, basal bract lanceolate c. 16 x 2 mm, brown

hairy abaxially, sparsely hairy adaxially, medial bract ovate, c. 3 x 2 mm, hairy outside,

glabrous within, calyx completely covering flower bud, sepals at anthesis triangular, c.

6 x 6m, drying dark brown, outside with brown hairs densest near the base, inside with

small pale hairs, petals white or pale yellow, reflexing at anthesis, oblong-ovate c. 10 =

5 mm, apex blunt, densely pale brown hairy on both surfaces, stamens many truncate,

carpels many hairy. Fruiting pedicel c. 2.5 cm long, 4 mm thick, drying striate, brown

stellate hairy, monocarps to 15 or more, sessile, pyriform, muricate, often with flat

oblique faces basally, apex rounded, to 3.5 x 2.5 cm, drying brown, covered with short

dense golden brown tomentum. Seeds several, ellipsoidal with one or two flattened

faces, 12 x 6-8 x 3-6 mm, drying smooth, brown, shiny.

Distribution. Endemic to Borneo, where it has been collected from Brunei, Sabah and

Sarawak.

442 Gard. Bull. Singapore 64(2) 2012

Ecology. Lowland dipterocarp forests (primary and disturbed), near streams and

freshwater swamps; 30-720 m.

Notes. Beccari appears to have recognised that the his collection represented a new

species and distributed the material under the name ‘Uvaria vallombrosana’, the

proposed epithet deriving from the name of the house, ‘Vallombrosa’, he had built on

Mt. Matang in Sarawak.

Fruiting material of U. beccarii 1s morphologically similar to that of

U. verrucosa, although the two species are unlikely to be confused if flowering material

is available. Uvaria verrucosa fruits are borne on shorter peduncles and pedicels, and

the monocarps are smaller and subglobose with an apiculate apex, and have fewer

seeds.

3. Uvaria borneensis (Merr.) Utteridge

‘(of Borneo)

Blumea 45 (2000) 393, as ‘borneense’. — Tetrapetalum borneense Merr., Univ.

Calif. Publ. Bot. 15 (1929) 64. Masamune, Enum. Phan. Born. (1942) 297. Sinclair,

Sarawak Mus. J. 5 (1951) 608. TYPE: Borneo, Sabah, Tawau, October 1922—March

1923, A.D.E. Elmer 21211 (lectotype, designated by Utteridge (2000), K (barcode no.

K000691249); isolectotypes: A, B, BISH, BM, BO, C, CM, DS, IBSL, L, MICH, MO,

NYA Eas UsZ):

Uvaria lambirensis (K. Momose) Utteridge, Blumea 45 (2000) 393, as ‘lambirense’.

— Tetrapetalum lambirense K. Momose, Blumea 43 (1998) 117. TYPE: Borneo,

Sarawak, Miri, Lambir Hills National Park, 1 April 1996, K. Momose 5069 (holotype:

KYO; isotypes: BM, BO, K, L).

Large woody climber. Young twigs rusty stellate hairy. Leaves chartaceous to

coriaceous, hairy on midrib and veins above when young, ovate to ovate lanceolate,

7.5-27 =< 3-12 cm. base rounded to truncate, apex acuminate. Inflorescence supra-

axillary or behind leaves, pedunculate to 10 mm, usually with | or 2 flowers. Pedicel

to 3 cm, often much shorter, densely stellate tomentose. Often drying with longitudinal

wrinkles. Clasping medial bract c. 3 mm long; sepals 2, orbicular, 8-10 mm across, 4

mm long; petals 4, green, membranous, concave, oblong ovate, 13 x 10 mm; stamens

many, carpels many. Monocarps to 30 or more, ellipsoidal, c. 3 x 2 cm, drying brown,

minutely wrinkled, densely covered with very short hairs, apex blunt, stipe to 2 cm

long. Seeds many, flattened, c. 15 = 10 x 3 mm, smooth, shiny.

Distribution. Endemic to Borneo. Known from Sarawak, Sabah and Kalimantan.

Ecology. Lowland forest.

Climbing Annonaceae of Borneo 443

4. Uvaria clementis (Merr.) Attan., 1.M.Turner & R.M.K.Saunders

(Latin, clementis (clemens in genitive case) = of gentility, mildness, mercy;

commemorative of Joseph Clemens (1862-1936), a Methodist minister, originally

from England, who served with the US Army, and his American wife Mary Strong

Clemens (1873-1968), who collected plants professionally).

Novon 21 (2011) 166. — Artabotrys clementis Merr., J. Straits Branch Roy. Asiat.

Soc. 85 (1922) 174. Masamune, Enum. Phan. Born. (1942) 280. TYPE: Borneo,

Sabah, Sandakan and vicinity, September-December 1920, . Ramos 1667 (lectotype,

designated by Attanayake et al. (2011), K; isolectotype: A).

Uvaria parviflora Hook.f. & Thomson, FI. Ind. 1 (1855) 103, non U. parviflora A. Rich.

(1831), nec U. parviflora (Michx.) Torr. & A. Gray (1838). — Uva parviflora Kuntze,

Rev. Gen. Pl. 1 (1891) 8. — Cyathostemma hookeri King, J. Asiat. Soc. Bengal, Pt.

2, Nat. Hist. 61(2) (1892) 10, nom. superfi. Ridley, Sarawak Mus. J. 1(3) (1913) 73.

Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921) 263. Masamune, Enum.

Phan. Born. (1942) 297. Sinclair, Gard. Bull. Singapore 14 (1955) 223. Utteridge,

Blumea 45 (2000) 386. Beaman et al., Pl. Mt. Kinabalu 4 (2001) 83. TYPE: Peninsular

Malaysia, Prince of Wales Island (Penang), W-E. Phillips s.n. (lectotype, designated by

Attanayake et al. (2011), K (barcode no. K 000615957)).

Large woody climber. Twigs becoming glabrous with age. Leaves drying brown or

grey-brown, coriaceous, sparsely pubescent below, oblong to obovate, 12—20 = 5.5—8

cm, base cuneate, apex acute to acuminate, lateral nerves 12—17 pairs. Petiole 4-7 mm

long, 1-3 mm thick. Inflorescence opposite leaves, 3- or 4-flowered. Pedicel to 12 mm

long, minutely verruculose, basal and medial bracts minute, sepals ovate 2.5—3 « 2.5—5

mm, petals, elliptic to ovate, 4-6 x 2-3 mm, stamens many, carpels many. Monocarps

to 20 or more, globose or ellipsoidal, 3-4 cm long, c. 2—2.5 cm diameter, glabrous,

drying dark red-brown, apex rounded, stipes 2—5 cm long. Seeds up to 8.

Distribution. Malay Peninsula and Borneo. In Borneo collected from Sarawak, Sabah

and Kalimantan.

Ecology. Lowland forest.

Notes. Fruiting specimens can be difficult to distinguish from those of U. concava but

the monocarps of U. concava dry black whereas those of U. clementis are a reddish

brown in the dry state.

5. Uvaria concava Teijsm. & Binn.

(Latin, concave, curved inwards, hollowed out)

Natuurk. Tjdschr. Ned.-Indié 3 (1852) 331. Teijsmann & Binnendijk, Ned. Kruidk.

444 Gard. Bull. Singapore 64(2) 2012

Arch. 3 (1855) 406. — Uva concava (Teijsm. & Binn.) Kuntze, Rev. Gen. PI. 1 (1891)

8. TYPE: Java, cult. in Hort. Bot. Bogor., S. Binnendijk s.n. (Herb. Bogor. 17623]

(lectotype, designated by Turner (2011b), BO (sheet no. BO-1824857)).

Uvaria lurida Hook.f. & Thomson, Fl. Ind. 1 (1855) 101. — Uva lurida (Hook.f. &

Thomson) Kuntze, Rev. Gen. Pl. 1 (1891) 8. TYPE: India, Assam, Khasia, 2000 feet,

2 October 1850, J.D. Hooker & T. Thomson 2468 (lectotype, designated by Turner

(2011b), K (barcode no. K000380703)).

?Uvaria sessiliflora Rchb.f. & Zoll., Linnaea 29 (1858) 306, 312. TYPE: Java, Hort.

Bot. Bogor, H. Zollinger 1412 (isotype: P (barcode no. PO0260015) (n.v.)).

Uvaria stellata Merr., Publ. Bur. Sci. Gov. Lab. 29 (1905) 14. TYPE: Philippines,

Luzon, Province of Benguet, Twin Peaks, May 1904, 4.D.E Elmer 6322 (lectotype,

designated by Turner (2011b), K (barcode no. K000691146); isolectotypes: NY, P,

US).

Unona leytensis Elmer, Leafl. Philipp. Bot. 5 (1913) 1744. — Uvaria leytensis (Elmer)

Merr., Philipp. J. Sci., C. 10 (1915) 230. Merrill, Enum. Philipp. Fl. Pl. 2 (1923) 155.

TYPE: Philippines, Mindanao, Province of Agusan, Cadadbaran (Mt Urdanete),

September 1912, 4.D.E. Elmer 13880 (lectotype, designated by Turner (2011b), L

(barcode no. LO190815); isolectotypes: BM, BO, CAL, GH, K, L, NA(ex MOAR), NY,

PUAUS)):

Uvaria nudistellata Elmer, Leafl. Philipp. Bot. 5 (1913) 1746. Merrill, Enum. Philipp.

Fl. Pl. 2 (1923) 156. TYPE: Philippines, Palawan, Puerto Princesa (Mt Pulgar), April

1911, A.D.E. Elmer 13015 (lectotype, designated by Turner (2011b), L (barcode no.

L0190867); isolectotypes: A, BISH, BM, CAL, K, L, MO, NSW, NY, U, US, Z).

Uvaria sympetala Merr., Univ. Calif. Publ. Bot. 15 (1929) 63. Masamune, Enum. Phan.

Born. (1942) 298. TYPE: Borneo, Sabah, near Tawao, October 1922—March 1923,

A.D.E. Elmer 21090 (lectotype, designated by Turner (2011b), UC (sheet no. 289957);

isolectotypes: A, BISH, BM[2], BO, C, CM, DS, GH, K, L, M, MICH, MO, NY, P, PH,

5; SING, U; UC, Z):

Large woody climber. Foliage largely glabrous. Twigs dark brown or grey-brown,

latticed. Leaves drying olive green-grey, elliptic to lanceolate, 12-20 < 6-7 cm, base

cuneate to rounded, apex acute to acuminate, reticulations very distinct from above.

Petioles 5-10 x 1—2 mm. Inflorescence terminal, flowers solitary. Pedicel to 2 cm,

covered with very short adpressed brown stellate hairs, sepals connate, entirely covering

flower bud, splitting, petals red-brown, connate at base, obovate c. 15 mm long, concave,

stamens many, carpels many. Monocarps many (40 or more), drying black, cylindrical,

4 x 1.5 cm, glabrous except for a scattering of very tiny brown stellate hairs, stipe to 7

cm long. Seeds many.

Climbing Annonaceae of Borneo 44

Distribution. India to Australia including the Malay Peninsula, Sumatra, Java, Borneo

and the Philippines. In Borneo only recorded from Sabah.

Ecology. Lowland forest.

Notes. The monocarps drying black distinguish fruiting specimens from the otherwise

similar U. clementis that has monocarps drying a reddish brown.

6. Uvaria cuneifolia (Hook.f. & Thomson) L.L.Zhou & al.

(Latin, cuneatus = wedge-shaped, folius = leaf)

Syst. Biodivers. 7 (2009) 255. — Ellipeia cuneifolia Hook.f. & Thomson, FI. Ind. 1

(1855) 104, Ridley, Sarawak Mus. J. 1(3) (1913) 76. Mermill, J. Straits Branch Roy.

Asiat. Soc. Spec. No. (1921) 254. Masamune, Enum. Phan. Born. (1942) 283. Sinclair,

Gard. Bull. Singapore 14 (1955)230. TYPE: Peninsular Malaysia, near Malacca, W.

Griffith s.n. (\ectotype, designated by Tumer (2011b), K (barcode no. K000382203)).

Ellipeia gilva Miq., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 9. — Uvaria gilva (Mig.) L.L.

Zhou & al., Syst. Biodivers. 7 (2009) 255. TYPE: Borneo, W-H. de Vriese s.n. (lectotype.

designated by Turner (2011b), L (barcode no. L 0484299)).

Uvaria laha Miq., F1. Ned. Ind., Eertse Bijv. (1861) 369. — Ellipeia laha (Mig.) Mig..

Ann. Mus. Lugd.-Bat. 2 (1865) 10. Mermill, J. Straits Branch Roy. Asiat. Soc. Spec. No.

(1921) 254. Masamune, Enum. Phan. Born. (1942) 283. TYPE: Sumatra, West Sumatra,

Palembang near Tubuan, Ogau-ulu, JE. Teijsmann s.n. [Herb. Bogor. 3811] (holotype:

U (barcode no. U 0000268); isotype: BO (sheet no. BO-1349058)).

Large woody climber. Twigs longitudinally wrinkled or latticed, drying dark brown,

young twigs densely short stellate hairy, glabrescent with age. Leaves chartaceous, drying

dark above uniform brown beneath, or various shades of brown, lamina above with

uniform but sparse covering of pale stellate hairs, lost with age, dense brown tomentum

on midrib and nerves, beneath dense covering of stellate red-brown hairs, giving a felty

feel, or almost glabrous except for a few hairs along the midrib, leaves ovate to obovate

to more typically oblanceolate, 5-25 x 2-10 cm, base rounded, apex acuminate, lateral

veins 15-20 pairs, arching forward, looping obscurely within the margin. Petioles 4-5

mm long, to 3 mm thick. Inflorescences terminal, many-flowered. Flowers pedicel 11-14

mm long with short dense red-brown stellate tomentum, basal bracteole narrowly ovate

to 8 mm long, clasping medial bract to 34 mm long, sepals broadly triangular to ovate,

4 mm long, 4-5 mm across, apex obtuse, rusty stellate outside; petals creamy yellow,

outer petals broadly ovate, 17 x 11 mm, apex obtuse, inner petals triangular, 4 mm long,

5 mm wide; stamens numerous, carpels many. Fruits pedicel to 20 mm long, 2 mm thick,

monocarps to 20 or more, compressed ovoid, 15 x 8-10 x 6 mm, drying pale brown,

shortly tomentose, minutely bumpy, apiculus lateral, stipe to 15 mm long. Seeds 1.

446 Gard. Bull. Singapore 64(2) 2012

Distribution. Malay Peninsula, Sumatra and Borneo. In Borneo collected from Sabah

and Sarawak, and probably Kalimantan.

Exology. Lowland forest.

7. Uvaria curvistipitata Attanayake & al.

(Latin, curvi- = curved, stipitatus = with a stipe or short stalk; curved stipes to the

monocarps)

Novon 21 (2011) 164, fig. 3. TYPE: Borneo, Sabah, Labuk and Sugut, west side of

Bukit Doji pass from Telupid to Ulu Karamuak, 25 October 1968, S. Kokawa & M.

Hotta 435 (holotype: SAN; isotypes: KYO, L).

Large woody climber. Twigs drying dark brown or black, latticed, younger parts with

dense, quite coarse, brown hairs. Leaves chartaceous to coriaceous, drying brown or

grey brown above, often rather patchy, brown beneath, midrib slightly sunken above

in dry leaves, prominent beneath, lateral nerves more or less flush above, prominent

beneath, variably hairy above, sometimes restricted to veins, densely hairy beneath,

lamina (oblong-)elliptic to typically obovate, 8-25 x 4-13 cm, base truncate, rounded

or auriculate, apex shortly acuminate, lateral nerves 13-17 pairs, arching forward,

looping obscurely within margin, tertiary venation visible from below, obscure from

above. Petiole 5-8 mm long, 2—5 mm thick, densely hairy. Inflorescence subopposite

leaves, several-flowered, peduncle c. | cm long. Flowering pedicel c. 8 mm long, 2.5

mm thick, densely hairy, medial bract very broadly elliptic, 3 x 8 mm, with short brown

hairs, sepals broadly triangular, c. 5 x 7 mm, connate at base, hairy externally, petals,

whorls similar, ovate, c. 8 x 8 mm, covered on both surfaces with short brown hairs,

stamens many, carpels many. Fruiting pedicel c. 10 mm long, 4 mm thick, monocarps

to 25 or more, globose to ellipsoidal, 1.5—2 x 1.5 cm, apex rounded, sometimes with

some longitudinal ridges, densely brown hairy, stipe to 3.5 cm long, 3 mm thick. Seeds

c. 6 in 2 rows, ellipsoidal, flattened on one or two faces, 11—12 x 7-8 x 4-6 mm, drying

brown, smooth, shiny, glabrous.

Distribution. Endemic to Borneo. Collected from Kalimantan and Sabah.

Ecology. Lowland forest to 200 m.

8. Uvaria excelsa (Hook.f. & Thomson) King

(Latin, lofty, high)

J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 61(2) (1893) 22. Ridley, Sarawak Mus. J.

1(3) (1913) 74. Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921) 253.

Masamune, Enum. Phan. Born. (1942) 297. — Mitrephora excelsa Hook.f. &

Climbing Annonaceae of Borneo 447

Thomson, Fl. Ind. 1 (1855) 114. Ridley, Sarawak Mus. J. 1(3) (1913) 86. Merrill, J.

Straits Branch Roy. Asiat. Soc. Spec. No. (1921) 263. Masamune, Enum. Phan. Born.

(1942) 289. — Kinginda excelsa (Hook.f. & Thomson) Kuntze, Rev. Gen. Pl. (1891)

7. — Cyathostemma excelsum (Hook.f. & Thomson) J. Sinclair, Gard. Bull. Singapore

14 (1955)226. Kessler & van Heusden, Rheedea 3 (1993) 60. Bygrave in Coode et

al., Checkl. Fl. Pl. Gymnosp. Brunei (1996) 14. Utteridge, Blumea 45 (2000) 384.

Beaman et al., Pl. Mt. Kinabalu 4 (2001) 83. TYPE: Peninsular Malaysia, Penang, G.

Porter s.n. [EIC 6477] (holotype: K (barcode no. K000691369): isotypes: CGE, K,

K-W, WU).

[Uvaria excelsa Wall., Numer. List (1832) no. 6477, nom. nud. |

Tetrapetalum volubile Miq., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 1. Ridley, Sarawak

Mus. J. 1(3) (1913) 73. Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921)

254. Masamune, Enum. Phan. Born. (1942) 297. TYPE: Borneo, 1857, W.H. de Vriese

s.n. (holotype: L[<3] (barcode nos. L0037889, L0037900, L0037901); isotype: U).

Uvaria confertiflora Merr., Univ. Calif. Publ. Bot. 15 (1929) 61. Masamune, Enum.

Phan. Born. (1942) 297. TYPE: Borneo, Sabah, Tawau, October 1922—March 1923,

A.D.E. Elmer 21081 (lectotype, designated by Utteridge (2000), L (barcode no. L

0037902); isolectotypes: A, BISH, BM[x2], BO, C, CM, DS, K, MICH, MO, NY, P,

Pi. S, SING, U; UC, Z).

Large woody climber. Twigs pubescent becoming glabrous with age. Leaves coriaceous,

glabrous above except on midrib, below variably pubescent from extremely densely

so, to almost glabrous, becoming less hairy with age, but apparently a lot of variability

between plants or populations in hairiness; lamina oblong-obovate 13—22 « 6-12 cm,

base cordate, apex acute or acuminate, lateral nerves 8-11 pairs. Petiole 7-12 mm

long, 1-3 mm thick. Inflorescence clustered cymes of 4-20 flowers, on twigs behind

leaves or supra-axillary. Pedicels 0-5 mm long, bracts coriaceous, pubescent, ovate,

sepals 2 or 3, connate at base, coriaceous, suborbicular 2-4 = 3—5.5 mm, densely

pubescent, petals 4 or 6, broadly ovate, 5-6 « 5—6 mm, apex obtuse, stamens many,

carpels many. Monocarps to 20 or more, ripening yellow orange, densely pubescent,

globose, 2—2.5 cm diameter, tuberculate, stipes to 3 cm long. Seeds c. 12 in two rows,

semi-circular in outline with two flat faces and one convex, c. 14-16 * 8-10 x 5 mm,

drying brown.

Distribution. Malay Peninsula, Sumatra and Borneo. In Borneo recorded from Brunei,

Kalimantan, Sabah and Sarawak.

Ecology. Lowland forest.

Notes. Sometimes confused with Polyalthia chrysotricha Ridl., though this is a tree

not a climber.

448 Gard. Bull. Singapore 64(2) 2012

9. Uvaria grandiflora Roxb. ex Hornem.

(Latin, grandis = large, florus = flower, with large flowers)

Hort. Hafn., Suppl. (1819) 141. Kessler & van Heusden, Rheedea 3 (1993) 85. Beaman

et al., Pl. Mt. Kinabalu 4 (2001) 98. TYPE: India, cultivated in the Calcutta Botanic

Garden, April 1818, Anon. s.n. (lectotype, designated by Turner (2011b), C).

[Uvaria grandiflora Roxb., Hort. Bengal. (1814) 43, nom. nud.]

Unona grandiflora DC., Prodr. 1 (1824) 91. — Uva grandiflora (DC.) Kuntze, Rev.

Gen. Pl. 1 (1891) 8. TYPE: India, Bengal, 1821, Leschenault de la Tour s.n. (holotype:

G-DC (barcode no. G00201447)).

Uvaria purpurea Blume, Biydr. (1825) 11. Ridley, Sarawak Mus. J. 1(3) (1913) 75.

Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921) 254. Merrill, Enum.

‘Philipp. Fl. Pl. 2 (1923) 156. Masamune, Enum. Phan. Born. (1942) 298. TYPE: In

sylvis humilioribus Javae Insulae. Floret Nov Dec, nomen kadjang.

Uvaria platypetala Champ. ex Benth., Hooker’s J. Bot. Kew Gard. Misc. 3 (1851)

257. TYPE: Hong Kong, East Point, low jungle near the Buddhist Temple, East Point,

1850, J.G. Champion 38 (holotype: K (barcode no. K000380694)).

Uvaria rhodantha Hance ex Walp., Ann. Syst. Bot. 2 (1851) 19. TYPE: Hong Kong,

April 1852, H.-F) Hance 933 (neotype, designated by Turner (2011b), BM; isoneotype:

CGE):

? Uvaria purpurea vat. subbiflora Migq., Fl. Ned. Ind., Eerste Biyv. 3 (1861) 368. TYPE:

Sumatra austr. secus flumen Tarabangi (Teijsmann).

? Uvaria flava Teijsm. & Binn., Natuurk. Tijdschr. Ned.-Indié 25 (1863) 419. — Uvaria

purpurea vat. flava (Teijsm. & Binn.) Scheff., Natuurk. Tijdschr. Ned.-Indié 31 (1869)

4. TYPE: Bangka, Teijsmann.

? Uvaria purpurea vat. alba Scheff., Natuurk. Tijdschr. Ned.-Indié 31 (1869) 4. TYPE:

Sumatra, Priaman, Diepenhorst.

Uvaria purpurea var. tuberculata King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 61(2)

(1892) 18. — Uvaria grandiflora Roxb. ex Hornem. var. tuberculata (King) J. Sinclair,

Gard. Bull. Singapore 14 (1955) 203. TYPE: Peninsular Malaysia, Perak, August

1883, King’ Collector [H.H. Kunstler] 4786 (lectotype, designated by Turner (2011b),

K (barcode no. K0003809129): isolectotype: CAL).

Uvaria rubra C.B. Rob., Bull. Torrey Bot. Club 35 (1908) 68. Merrill, Enum. Philipp.

Fl. Pl. 2 (1923) 156. TYPE: Philippines, Mindanao, Prov. Davao, Santa Cruz, 29 June

Climbing Annonaceae of Borneo 449

1905, R.S. Williams 3042 (holotype: NY(barcode no. 00059991): isotypes: NY[*2]).

?Uvaria cardinalis Elmer, Leafi. Philipp. Bot. 5 (1913) 1748, as ‘cardinales’. Merrill,

Enum. Philipp. Fl. Pl. 2 (1923) 156. TYPE: Philippines, Luzon, Sorzogon, 4.D.E.

Elmer 7317 (holotype: PNH (destroyed)).

Large woody climber. Twigs golden hairy when young, becoming glabrous with age.

Leaves typically hirsute, chartaceous, obovate to (ob)lanceolate, 9-30 x 3-15 cm, base

rounded to cordate, apex acuminate. Petiole 4-7 mm long, 2—3 mm thick. Inflorescence

subopposite leaves, usually solitary, occasionally more than one flower. Pedicel to 2.5

cm long, basal bract foliose to 3 cm long, medial bract to 2 cm long, calyx entirely

enclosing flower in bud, splitting into three orbicular sepals, 2 cm across, petals fleshy,

red-purple (white or yellow reported from outside Borneo), ovate to obovate, 30 = 15

mm, stamens many, carpels many. Monocarps to 30 or more, cylindrical 5 x 1.5 cm,

longitudinally ridged, covered in dense adpressed hairs, stipe to 1 cm. Seeds to 20 or

more in two rows, more or less semi-circular, c. 10 x 5 x 3 mm, drying brown.

Distribution. From Burma and China, through Indochina to Java and the Philippines.

In Borneo collected from Kalimantan, Sabah and Sarawak (only known from Gunung

Subis).

Ecology. Lowland forest.

10. Uvaria griffithii L.L.Zhou & al.

(William Griffith (1810-1845), English surgeon and botanist)

Syst. Biodivers. 7 (2009) 255. — Cyathostemma viridiflorum Griff., Not. Pl. Asiat. 4

(1854) 707. Sinclair, Gard. Bull. Singapore 14 (1955) 221. Kessler & van Heusden,

Rheedea 3 (1993) 62. Utteridge, Blumea 45 (2000) 390. Beaman et al., Pl. Mt. Kinabalu

4 (2001) 83. TYPE: Peninsular Malaysia, Malacca, W. Griffith s.n. [Kew Distribution

no. 432] (lectotype, designated by Utteridge (2000), K (barcode no. K000582099)).

Cyathostemma scortechinii King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 61(2)

(1893) 9. — Cyathostemma viridiflorum var. scortechinii (King) Ridl., Fl. Malay

Penins. 1 (1922) 27. TYPE: Peninsular Malaysia, Perak, Gopeng, King’s Collector

[H.H. Kunstler] 5857 (lectotype, designated by Utteridge (2000), K (barcode no.

K000380130); isolectotypes: BM, CAL, SING).

Large woody climber. Twigs slightly hairy when young, otherwise glabrous. Leaves

typically drying grey above, grey-brown or brown below, coriaceous, glabrous except

for midrib above, oblong-elliptic to oblong-lanceolate, 12—17 « 5.5—8 cm, base rounded,

apex acute. Petiole 6-12 mm long, c. 2 mm thick. Inflorescence cauliflorous, many-

flowered cymes, peduncles pendulous, 2.5—8 cm long, densely pubescent. Pedicel 6—9

450 Gard. Bull. Singapore 64(2) 2012

mm long, bracts coriaceous, long-persistent, 34 mm long, sepals broadly ovate, 3 x

3—5 mm, connate at base, petals broadly ovate, greenish yellow, 5—6 x 4-5 mm, apex

acute, stamens many, carpels many. Monocarps to 10 or more, cylindrical-ellipsoidal,

drying black, stipes to 2 cm. Seeds 7—10 in 2 rows, semi-circular, c. 10 x 5 x 3 mm.

Distribution. Malay Peninsula, Sumatra and Borneo. In Borneo, once from Sarawak

and a few collections from Kalimantan.

Ecology. Lowland forest.

11. Uvaria hirsuta Jack

(Latin, hairy)

Malay Misc. | (1820) 46. Sinclair, Sarawak Mus. J. 5 (1951) 608. Sinclair, Gard.

Bull. Singapore 14 (1955) 203. — Uvaria blumeana Steud., Nomencl. 2 (1841)

737, nom. superfl. — Uva hirsuta (Jack) Kuntze, Rev. Gen. Pl. 1 (1891) 8. TYPE:

Peninsular Malaysia, Penang, 1822, N. Wallich s.n. [EIC 6458B] (neotype, designated

by Turner (2011b), K-W (barcode no. K000442828); isoneotypes: ?BM, ?K (barcode

no. K000739195).

Uvaria velutina Roxb. ex Blume, Bijdr. (1825) 13, non U. velutina DC. in Dunal

(1817). — Uvaria hirsuta Blume, Fl. Javae Anonac. (1830) 22, t. 5, non U. hirsuta

Jack (1820), nec U. hirsuta Vell. (1829). TYPE: Java, West Java, Kuripan, C.L. Blume

s.n (holotype: L (sheet no. 898.63-384)).

Uvaria trichomalla Blume, F1. Java. Anonac. (1830) 42, t. 18. TYPE: Java, West Java,

Tjikao, Kuhl & van Hasselt 1837 (holotype: L).

Guatteria pilosa Roxb. ex G. Don, Gen. Hist. 1 (1831) 100. — Uvaria pilosa (Roxb. ex

G. Don) Roxb., Fl. Ind. 2 (1832) 665. TYPE: ?Moluccas, W. Roxburgh s.n. [EIC Herb.

6458A] (lectotype, designated by Veldkamp (2011), K-W (barcode no. K000442826)).

Large woody climber. Twigs pale grey-brown, latticed, covered with 1-3 mm long,

more or less straight, red-brown hairs. Leaves characteous to subcoriaceous, upper

surface slightly rough to the touch, pilose below, drying brown, grey-brown or blackish

brown above, brown below, midrib and laterals more or less flush to the surface above,

prominent beneath, lamina elliptic, oblong-elliptic, obovate or oblong obovate, 5—24 x

3—8 cm, apex shortly or bluntly acuminate but very tip often a distinct slender apiculus,

base truncate, obtuse, rounded or auriculate, lateral veins 9—13 pairs, arching forward

and looping obscurely within the margin, tertiary venation indistinct, particularly

from above. Petiole 3—6 mm long, 1-3 mm thick. Inflorescences among leaves, supra-

axillary, generally single-flowered. Flowering pedicel, 2—3 cm long, red-brown pilose,

medial bract lanceolate 4-10 « 1.5—3 mm, red-brown pilose on both surfaces, denser

Climbing Annonaceae of Borneo 451

outside, petals dark red, ovate c. 10 x 8 mm, stamens many, carpels many. Fruiting

pedicel to 2.5 cm long, 2 mm thick, monocarps to 8, globose, ellipsoidal or cylindrical,

2-5 cm long, c. 1.5 cm diameter, densely covered with red-brown hairs, stipe 23.5 cm

long, c. 2 mm thick. Seeds 12 or more in 2 rows, with two flat faces and one convex,

6-8 x 5 x 4 mm, drying brown, smooth.

Distribution. Burma, Thailand, Malay Peninsula, Java and Borneo. In Borneo from

Sarawak.

Ecology. Lowland forest.

12. Uvaria javana Dunal

(of Java)

Monogr. Anonac. (1817) 91, pl. 14. Merr., Univ. Calif. Publ. Bot. 15 (1929) 62.

Masamune, Enum. Phan. Born. (1942) 297. Sinclair, Gard. Bull. Singapore 14 (1955)

216. TYPE: Hab. in Java Lahaie (v.s.h. Deless.).

[Uvaria ochroleuca Zoll., Linnaea 29 (1858) 304, 307, nom. inval. (cf. ICBN Art.

34.1(b))]

?Uvaria javana var blumei Boerl., Cat. Pl. Phan. (1899) 13. TYPE: Java, cult. in Hort.

Bot. Bogor. sub XI.A.44.

Uvaria larep auct. non Mig.:Merrill (1929: 62). Masamune (1942) 298.

Large woody climber. Twigs drying dark brown to black, smooth, longitudinally

wrinkled or latticed, youngest parts covered in dense erect brown stellate pubescence.

Leaves chartaceous to subcoriaceous, typically with a scabrous feel above, though

some specimens are relatively smooth, drying grey-brown or grey above, brown or

pale brown below with venation a darker shade, midrib very slightly sunken above

in dry leaves, prominent below, lateral nerves more or less flush above slightly

raised beneath, typically pubescent with a dense fringe of hairs on midrib above with

scattered stellate hairs on upper lamina, below densely brown stellate hairy on nerves,

lower lamina with dense pale stellate hairs giving a felty feel, otherwise pubescence

relatively sparse, lamina elliptic, oblong-elliptic or obovate, 6—16.5 x 3—9.5 cm, base

obtuse, truncate or rounded, apex (rarely emarginate) obtuse to shortly acuminate,

lateral nerves 7-13 pairs, looping indistinctly within the margin; tertiary venation

generally difficult to see unaided. Petiole 3-7 mm long, 1-3 mm thick, densely hairy.

Inflorescences supra-axillary or on twigs behind leaves, single-flowered. Flowering

pedicel 6-30 mm long, densely brown stellate hairy, medial bract ovate 3-4 x 3 mm,

stellate hairy abaxially, glabrous adaxially, sepals triangular, c. 4 x 4 mm, apex obtuse

to rounded, hairy outside, glabrous within, petals yellow, reflexing right back to pedicel

452 Gard. Bull. Singapore 64(2) 2012

at anthesis, whorls similar, ovate-oblong 10—12 x 5—6 mm, apex obtuse, densely hairy

on both surfaces, drying pale brown, stamens many 1.5—2 mm long, connective apex

tongue-like, carpels many, c. 2 mm long, hairy, stigma drying black. Fruiting pedicel

1.5—2 cm long, 5-6 mm thick, monocarps to 35 or more, ellipsoidal typically with

slight concavity on one long side and convexity on the other, to 4.5 x 3 cm, drying

brown with dense, very short pale brown tomentum, markedly rugulose particularly

in immature fruit, apex rounded, stipe, typically shorter than seed-bearing portion

of monocarp, 0.5—2 cm long, c. 4 mm thick. Seeds many, in two rows, ellipsoidal,

flattened, 10-12 x 7-8 x 4 mm, dark brown, smooth, shiny.

Distribution. Malay Peninsula, Java and Borneo. In Borneo collected from Kalimantan,

Sabah and Sarawak.

Ecology. Lowland forest.

13. Uvaria lanuginosa Ridl.

(Latin, woolly, downy)

Bull. Misc. Inform. Kew 1912 (1912) 382. Ridley, Sarawak Mus. J. 1(3) (1913) 75.

Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921) 253. Masamune, Enum.

Phan. Born. (1942) 298. Bygrave in Coode et al., Checkl. Fl. Pl. Gymnosp. Brunei

(1996) 23. Beaman et al., Pl. Mt. Kinabalu 4 (2001) 99. TYPE: Borneo, Sarawak, near

Kuching, 13 November 1894, G.D. Haviland & C. Hose 3334, 13 November 1894

(holotype, K (barcode no. K000380301); isotypes, BM, SAR).

Large woody climber. Leaves red-brown woolly floccose above and below when

young, mature leaves coriaceous becoming glabrous above, very densely brown

stellate hairy below, felty to the touch, ovate to obovate 7-13 x 3—6 cm, base rounded,

apex acuminate. Inflorescence axillary, supra-axillary or below leaves on twigs, few

flowered. Pedicels to 1.5 cm long, sepals, broadly triangular, 7 x 7 mm, apex blunt,

petals yellow ovate, 10-15 x 5—10 mm, stamens many, carpels many. Fruits unknown.

Distribution. Endemic to Borneo. Collected from Brunei and Sarawak.

Ecology. Lowland forest.

14. Uvaria littoralis (Blume) Blume

(Latin, of the seashore)

Fl. Javae Anonac. (1830) 26. Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No.

(1921) 253. Merr., Univ. Calif. Publ. Bot. 15 (1929) 61. Masamune, Enum. Phan.

Born. (1942) 298. — Unona littoralis Blume, Bijdr. (1825) 16. — Uvaria gamopetala

Climbing Annonaceae of Borneo 453

Zoll., Linnaea 29 (1858) 304, 310, nom. superfl. TYPE: in Bataviae locis stagnosis.

Floret Febr. Martio.

[Uvaria macrophylla Roxb., Hort. Bengal. (1814) 43, nom. nud]

Guatteria cordata Dunal, Monogr. Anonac. (1817) 129, t. 30. — Uva cordata (Dunal)

Kuntze, Rev. Gen. PI. 1 (1891) 8. — Uvaria cordata (Dunal) Alston, Handb. Fl. Ceylon

6(suppl.) (1931) 4, non U. cordata Schumach. & Thonn. (1827). Sinclair, Gard. Bull.

Singapore 14 (1955) 207. TYPE: Uvaria zeylanica (herb. Deless.), Hab in Java (v.s.h.

Deless.)

Uvaria ovalifolia Blume, Fl. Javae Anonaceae (1830) 27, tab. 8. Ridley, Sarawak Mus.

J. 1(3) (1913) 75. Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921) 253.

Bygrave in Coode et al., Checkl. Fl. Pl. Gymnosp. Brunei (1996) 23. Beaman et al., PI.

Mt. Kinabalu 4 (2001) 99. TYPE: van Hasselt cataractam Ramu prov. Bantam.

Uvaria macrophylla Roxb. ex Wall., Pl. As. Rariores 2 (1830) 22, pl. 122. TYPE:

India, cultivated in Calcutta Botanic Garden, N. Wallich s.n. [EIC 6487A], lectotype,

designated by Turner (2011b), K-W (barcode no. K000442833); isolectotypes: BM,

CGE).

Uvaria rufescens A.DC., Mem. Soc. Phys. Genev. 5 (1832) 202. TYPE: Burma,

Prome, 1826, N. Wallich 130 [EIC 6487B] (lectotype, designated by Turner (201 1a),

K-W (barcode no. K000442832)).

Unona camphorata Blanco, FI. Filip. (1837) 468. TYPE: Philippines, [Alabat Island,

December 1916], Anon. s.n. [Merrill: Species Blancoanae No. 1057] (neotype,

designated by Turner (2011b), K; isoneotypes: L, P, US (barcode no. 00688554)).

Uvaria sorzogonensis C. Presl, Rel. Haenk. 2 (1835) 76. Merrill, Enum. Philipp. FI.

Pl. 2 (1923) 157. TYPE: Philippines, Luzon, Sorzogon, 7-P.X. Haenke s.n. (holotype:

PR[*2] (sheet nos. 212920A and 212920B).

?Uvaria acrantha Mig., Fl. Ned. Ind., Eerste Bijv. 3 (1861) 368. — Uvaria ovalifolia

Blume var. acrantha (Mig.) Mig., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 7. TYPE:

Sumatra austr. in prov Lampongs prope Kebang, Teijsmann.

?Uvaria macrophylla var. glabrior Miq., Fl. Ned. Ind., Eerste Bijv. 3 (1861) 368.

TYPE: Bangka, Teijsmann.

?Uvaria ovalifolia var. borneensis Miq., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 7. Kessler

& van Heusden, Rheedea 3 (1993) 86. TYPE: Borneo australis Korthals.

?Uvaria ovalifolia var. normalis Mig., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 7. —

454 Gard. Bull. Singapore 64(2) 2012

Uvaria littoralis var. miquelii Boerl., Cat. Pl. Phan. (1899) 14, nom. superfl. TYPE:

Java, Sumatra, Borneo

?Uvaria ovalifolia var. racemiflora Mig., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 7. Ridley,

Sarawak Mus. J. 1(3) (1913) 75. TYPE: Borneo australis in regione fluminis Doessan,

Korthals.

Magnolia fasciculata P. Parm., Bull. Sci. France Belg. 27 (1896) 204, 265. TYPE: India

[probably wrongly localised], s.dat., T.S. Ralph s.n. (lectotype, designated by Turner

(2011b), P (barcode no. P00260050)).

Large woody climber. Twigs with very short brown hairs when young, becoming

glabrous and darkening with age. Leaves chartaceous to subcoriaceous, stellate hairy

along veins and lamina beneath when young, becoming glabrous except along midrib

above, ovate to ovate lanceolate, 13-31 x 7-13 cm, base truncate to rounded, apex acute

-to acuminate. Petiole 3-13 mm long, 1-4 mm thick. Inflorescence terminal or supra-

axillary, 3—4-flowered cymes. Pedicel 1.5—4 cm long, densely brown hairy, basal and

medial bract 3 mm long, 5 mm wide, calyx connate forming a triangular cup, sepals c.

5 mm long, petals red or red-yellow, oblong to obovate, 10 mm long by 7 mm wide,

inner whorl often slightly narrower, stamens many, often reduced to staminodes or even

petalloid structures. Monocarps many, ripening orange, globose or ellipsoidal, c. 1 cm

diameter, drying black or brown, glabrous, sometimes minutely beaked, stipe to 3 cm

long. Seeds 1 to several, , flattened on one or both faces, 7-8 « 5—6 x 4 mm, smooth,

light brown, shiny.

Distribution. From Sri Lanka and India to New Guinea. In Borneo collected from Brunei,

Kalimantan, Sabah and Sarawak. Much collected in Sabah.

Ecology. Lowland forest.

15. Uvaria lobbiana Hook.f. & Thomson

(Thomas Lobb, 1820-1894, British plant collector who collected widely in tropical Asia,

including Borneo over the period 1845-1856)

Fl. Ind. 1 (1855) 100. Sinclair, Gard. Bull. Singapore 14 (1955) 208. Kessler & van

Heusden, Rheedea 3 (1993) 86. — Uva lobbiana (Hook.f. & Thomson) Kuntze, Rev.

Gen. Pl. | (1891) 7. TYPE: Peninsular Malaysia, Malacca, 1845, W. Griffith s.n.

(lectotype, designated by Turner (2011b), K (barcode no. KO00380689)).

Uvaria subrepanda Wall. [Numer. List. (1832) no. 6483, nom. nud.] ex Hook.f. &

Thomson, Fl. Ind. 1 (1855) 101. — Uva subrepanda (Wall. ex Hook.f. & Thomson)

Kuntze, Rev. Gen. Pl. 1 (1891) 8. TYPE: Singapore, 1822, N. Wallich s.n. [EIC 6483]

(lectotype, designated by Turner (2011b), K-W (IDC tufiche WA667/19) (barcode no.

K00044825); isolectotype: CAL).

Climbing Annonaceae of Borneo 455

Uvaria ptychocalyx Miq.. Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 4. Ridley, Sarawak

Mus. J. 1(3) (1913) 74. Mermill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921)

254. Masamune, Enum. Phan. Born. (1942) 298. — Uva ptychocalyx (Miq.) Kuntze,

Rev. Gen. Pl. 1 (1891) 8. TYPE: Borneo, Poeloe Lampei, PW. Korthals s.n. (lectotype.

designated by Turner (2011b), L (barcode no. L0195619)).

Large woody climber. Twigs with tiny adpressed stellate hairs, becoming glabrous with

age. Leaves generally glabrous, some adpressed stellate hairs along veins and midrib

above, ovate, obovate to oblanceolate, 7-24 x 3—9 cm. Inflorescence subopposite

leaves, one or a few flowered. Petiole 3-7 mm long. 1—3 mm thick. Pedicel 3-10

mm long. densely brown hairy, sepals broadly orbicular, —6 mm long, 7-10 mm

wide, densely tomentose, petals red, broadly ovate to oblong. 10 <x 9 mm, verruculose,

densely pale tomentose, stamens many, carpels many. Monocarps very numerous (60

or more). globose or ellipsoidal, to 3 x 2 cm, drying muricate, stipe to 7 cm long. Seeds

c. 5, generally with two flattened faces and one convex, c. 10 x 8 x 3—5 mm, drying

brown, smooth.

Distribution. Burma, Thailand, Malay Peninsula. Sumatra and Borneo. In Borneo

recorded from Brunei, Kalimantan, Sabah and Sarawak.

Ecology. Lowland forest.

16. Uvaria micrantha (A.DC.) Hook.f. & Thomson

(Greek, micros = small, anthos = flower)

Fl. Ind. (1855) 103. Merrill, Enum. Philipp. Fl. Pl. 2 (1923) 155. Masamune, Enum.

Phan. Born. (1942) 298. — Guatteria micrantha A. DC., Mem. Soc. Phys. Genev.

5 (1832) 218. — Uva micrantha (A. DC.) Kuntze, Rev. Gen. Pl. 1 (1891) 8. —

Cyathostemma micranthum (A. DC.) J. Sinclair, Gard. Bull. Singapore, 14 (1955) 225.

Utteridge, Blumea 45 (2000) 388. TYPE: Burma. Martaban, Amherst, 17 June 1827,

W. Gomez 15 [N. Wallich 1287, EIC 6449] (lectotype. designated by Utteridge (2000),

K-W; isolectotypes: BM, G).

Polyalthia fruticans A. DC., Mém. Soc. Phys. Genéve 5 (1832) 216. TYPE: Burma,

Tavoy, 7 August 1827. W. Gomez 49 [N. Wallich 1288, EIC 6430] (lectotype. designated

by Turner (201 1a), K-W; isolectotype: G (barcode no. G00237293)).

Anaxagorea sumatrana Miq.. Fl. Ned. Ind., Eerste Bijv. 3 (1861) 382. — Uvaria

sumatrana (Miq.) Kurz. Rep. Veg. Andaman Isl., App. A (1870) 8. — Uva sumatrana

(Miq.) Kuntze. Rev. Gen. Pl. 1 (1891) 8. — Cyathostemma sumatranum (Miq.) Boerl.,

Icon. Bogor. 1 (1899) 126, t. 58. TYPE: Sumatra, Lampongs, near Tegineneng, J.-E.

Teijsmann s.n. (Herb. Bogor. no. 4383] (holotype: L: isotypes: GH, K, U).

456 Gard. Bull. Singapore 64(2) 2012

Popowia nitida King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 61(2) (1892) 92. TYPE:

Andaman Islands, South Andaman, Hobdaypur, 4 July 1891, G. King s.n. (lectotype,

designated by Utteridge (2000), K (barcode no. K000739196); isolectotype: K

(barcode no. K000739197)).

Large woody climber. Young twigs hairy, becoming glabrous with age. Leaves

chartaceous to subcoriaceous, glabrous above except on midrib, sparsely hairy beneath,

oblong elliptic, 5-14 x 24.5 cm, base cuneate to rounded, apex acute to acuminate.

Petiole 2—3 mm long, c. | mm thick. Inflorescence subopposite leaves, 2-flowered.

Pedicel 2-7 mm, pubescent, sepals broadly ovate 2.5 mm long, petals ovate, 4-5 x

3 mm, stamens many, carpels many. Monocarps to 25 or more, irregularly globose,

12-20 x 10 mm, glabrous, stipe 3—10 mm long, drying very slender c. 0.3 mm thick.

Seeds 1-2, rather irregular in shape with | or 2 flat faces, c. 5 x 4 x3 mm, drying

brown, smooth.

‘Distribution. Widespread from Burma and the Andaman Islands to the north coast of

Australia. In Borneo recorded from Sabah and Kalimantan.

Ecology. Lowland forest.

Notes. In fruit this species can be confused with Sphaerocoryne affinis, but the presence

of stellate hairs will confirm whether a specimen is Uvaria

17. Uvaria monticola Mig.

(Latin, mountaineer)

Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 5. Ridley, Sarawak Mus. J. 1(3) (1913) 74. Merrill,

J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921) 253. Masamune, Enum. Phan.

Born. (1942) 298. TYPE: Borneo, Gunung Balaran, PW. Korthals s.n. (holotype: L

(barcode no. L 0038219)).

Uvaria scandens C.B. Rob., Bull. Torrey Bot. Club 35 (1908) 69. Merrill, Enum.

Philipp. Fl. Pl. 2 (1923) 157. TYPE: Philippines, Mindanao, Prov. Davao, Santa Cruz,

5 May 1905, R.S. Williams 2764 (holotype: NY (barcode no. 00059993); isotypes: NY,

US).

Uvaria cauliflora Ridl., Bull. Misc. Inform. Kew 1912 (1912) 382. Ridley, Sarawak

Mus. J. 1(3) (1913) 74. Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921) 253.

Masamune, Enum. Phan. Born. (1942) 297. Beaman et al., Pl. Mt. Kinabalu 4 (2001)

98. TYPE: Borneo, Sarawak, Tegora, s.dat., G.D. Haviland 417 [c.k.q.a.] [Garai leg. ]

(lectotype, designated by Turner (2012), K (barcode no. K000380300); isolectotype:

K (barcode no. K000380298)).

Climbing Annonaceae of Borneo 457

Uvaria elmeri Merr., Univ. Calif. Publ. Bot. 15 (1929) 61. Masamune, Enum. Phan.

Born. (1942) 297. Kessler & van Heusden, Rheedea 3 (1993) 85. TYPE: Borneo.

Sabah, Tawao, October 1922—March 1923, 4.D.E. Elmer 20870 (lectotype, designated

by Turner (2011b), UC (sheet no. 289944): isolectotypes: A, BISH. BM. BO. C. CM.

DS, GH, L, M, MICH, MO, NY, P, PH, S, SING, U, UC, US, Z).

Uvaria sp.: Merr., Univ. Calif. Publ. Bot. 15 (1929) 63.

Large woody climber. Twigs brown with tightly adpressed stellate hairs when young.

Leaves with hairs along veins and midrib above, becoming glabrous with age, ovate to

obovate or oblanceolate, 10—30 x 6-12 cm, base rounded to cuneate, apex acuminate.

Petiole 2—11 mm long, 1—3 mm thick. Inflorescences mostly cauliflorous, multi-flowered

fascicles or branching leafless twigs from near stem base. Pedicel to 6 cm long, 1 mm

thick when dry, densely brown stellate hairy, medial bract c. 10 x 5 mm, fiower buds

c. | cm diameter, entirely enclosed within calyx, splitting irregularly during anthesis,

petals yellow-green, membranous, oblong-ovate, 15 x 10 mm, densely brown hairy,

reflexing with age, stamens many, carpels many. Monocarps to 25 or more, irregularly

cylindrical, not strongly ridged, to 6 x 2 cm, apex rounded, drying brown becoming

glabrous with age, rough to the touch, stipe to 2 cm. Seeds to 20 or more in 2 rows,

roughly flattened semicircles, c. 13—15 x 7-10 x 3-5 mm, drying brown.

Distribution. Borneo and the Philippines. In Borneo widely collected from Brunei,

Kalimantan, Sabah and Sarawak.

Ecology. Lowland forest.

Notes. In probably the only published study of pollination in an annonaceous climber

from Borneo, Nagamitsu & Inoue (1997) discovered that this species (as Uvaria

elmeri) is mainly pollinated by cockroaches that visit the flowers to feed on stigmatic

exudate and pollen.

18. Uvaria schefferi L.L.Zhou & al.

(R.H.C.C. Scheffer (1844-1880), Dutch botanist, Director of Buitenzorg Botanic

Gardens)

Syst. Biodivers. 7 (2009) 255. — Ellipeia coriacea Scheff., Ann. Jard. Bot. Buitenzorg

2 (1885) 4. Merrill, J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921) 254. Masamune,

Enum. Phan. Born. (1942) 283. TYPE: Borneo, pris de Montrado, J.E. Teijsmann s.n.

[Herb. Bogor. 10842] (holotype: BO (sheet no. BO-1350684)).

Large woody climber. Twigs drying dark grey or black with very tightly adpressed

brown stellate hairs (like tiny mites). Leaves chartaceous to coriaceous, drying

458 Gard. Bull. Singapore 64(2) 2012

typically dark brown above, mid-brown below with darker midrib and veins, lamina

below with uniform distribution of dark dots which are clumps of long stellate hairs set

in dense covering of stellate scales, giving a rough feel to the leaf, some collections are

glabrescent beneath, young leaves have a uniform but sparse scattering of very fine,

pale hairs above, soon dropping, oblong elliptic to obovate, 4.5-30 x 1.7—-10 cm, base

cuneate to obtuse, apex acuminate, midrib sunken above, prominent beneath, lateral

veins 10—20 pairs, looping distinctly within the margin. Petioles 6-16 =< 1-3 mm.

Inflorescences terminal or subopposite leaves, 1—several-flowered. Flower pedicels

5—8 mm (possibly longer), densely brown stellate hairy, sepals broadly ovate, 5 x 7

mm, densely brown adpressed stellate hairy outside, inside tomentum paler brown,

outer petals, ovate, c. 3.5 x 2 cm, densely short brown stellate hairy except for glabrous

minutely verrucose patch near base on inner surface, inner petals lanceolate c. 2.5 =

1 cm, densely short brown stellate hairy except near base adaxially, stamens many, c.

1.5 mm long, apex of connective truncate, flat, polygonous, carpels many, c. | mm

long, drying black with brown stellate hairs. Fruiting pedicels 15—30 mm long, to 3

- mm thick, monocarps 3-5, ovoid, 2.54 x 2—3.5 = 2 cm, ripening orange, drying red-

brown, scurfy, apiculus lateral, stipe to 3 cm long. Seeds 1, black shiny, shaped like an

orange segment with a blunt end, 3 = 2 x 1.2 cm, pale hilum at blunt end.

Distribution. Endemic to Borneo where it has been collected from Brunei, Kalimantan,

Sabah and Sarawak.

Ecology. Lowland and hill forest to 1800 m.

19. Uvaria verrucosa Scheff.

(Latin, warty)

Ann. Jard. Bot. Buitenzorg 2 (1885) 3. Ridley, Sarawak Mus. J. 1(3) (1913) 74. Merrill,

J. Straits Branch Roy. Asiat. Soc. Spec. No. (1921) 254. Masamune, Enum. Phan.

Born. (1942) 298. TYPE: Borneo, Soengei-aja (Kapoeas), J.E. Teijsmann s.n. [Herb.

Bog. no. 8192] (holotype: BO (sheet no. BO-1455790)).

Large woody climber. Twigs drying dark brown or dark grey-brown, striate or latticed,

younger parts with small adpressed brown stellate hairs (like small mites with many

legs). Leaves chartaceous to subcoriaceous, drying dark brown to pale grey-brown

above, brown to dark brown below, midrib slightly sunken above in dry leaves,

prominent below, lateral veins more or less flush above, slightly raised below, laminas

elliptic to obovate, 6—12.5 x 2-4 cm, base acute, obtuse or rounded, apex obtuse to

acuminate, glabrous above except for hairs on midrib, below with a scattering of tiny

stellate hairs, densest on midrib base and petiole, lateral veins c. 14 pairs, arching

forward and looping within margin, tertiary venation reticulat, generally indistinct.

Petiole 2-5 mm long, c. 1 mm thick. Inflorescence subopposite leaves, sometimes

appearing terminal when located above the last mature leaf on a twig, 1—2-flowered.

Climbing Annonaceae of Borneo 459

Flowering pedicel 3-8 mm long, c. | mm thick, densely brown stellate hairy, basal

bract ovate, c. 7 x 4 mm, apex acute, distinct central nerve, hairy on both surfaces,

medial bract (1-2), membranous, ovate, c. 15 x 15 mm, apex obtuse, hairy on both

surfaces, sepals thin, ovate, c. 10 x 8 mm, apex obtuse, hairy on both surfaces, outer

petals lanceolate, c. 40 x 11 mm, apex blunt, densely brown hairy on both surfaces

except for a rounded glabrous patch near the base internally, inner petals slightly

narrower (c. 7 mm), margins reflex but the petals appear to remain erect at anthesis,

stamens many, c. 1.5 mm long, apex domed, villose, papillate, carpels many. Fruiting

pedicel 6-10 mm long, 2—3 mm thick, monocarps to 10 or more, sessile to subsessile

(stipe 1-2 mm long), globose to ellipsoidal, c. 10-15 x 10-15 mm, distinct lateral

apiculus, drying dark brown, muricate, tuberculate, densely covered with dark brown

stellate hairs. Seeds 2.

Distribution. Endemic to Borneo. Recorded from Brunei, Kalimantan, Sabah and

Sarawak.

Ecology. Lowland forest.

ACKNOWLEDGEMENTS. Support for my research on the Annonaceae from the Arnold

Arboretum, Singapore Botanic Gardens, Forest Research Institute Malaysia and Royal Botanic

Gardens Kew is gratefully acknowledged. Thanks are due to the following for their assistance:

J. Gregson, J. Hunnex, J. Wajer (BM); A. Kartonegoro (BO); E. Luccioli, C. Nepi (FI): L.

Gautier (G); A.M.A.S. Attanayake, R.M.K. Saunders, (HKU); E. Wood (HUH): P.S. Ashton,

R.K. Brummitt, M. Cheek, R. de Kok, T. Harris, Y. Harvey, L. Pearce, T. Utteridge, X. van der

Burgt (K); N. Idris, E. Soepadmo, H.S. Tan (KEP): T. Chaowasku, J.F. Veldkamp, L. Willemse

(L); H.-J. Esser (M), P. Milne (MEL). A. Freire-Fierro (PH): O. Sida (PR); J. Pereira, S. Sabran,

J. Sugau (SAN); A. Chung, J. Pao, J. Sang (SAR), S.K. Ganesan, S. Lee, P. Leong, J. Leong-

Skorniékova, B.C. Tan (SING); C. Meade (TCD); C. Dilger-Endrulat (TUB).

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Appendix. List of specimens examined.

Material was seen from the following herbaria: A, BM, BO, GH, K, KEP, L, SAR, SAN, SING.

Abbe, E.C.; 10267: U. beccarii (K); 12074: P. prismatica (A, K). Abdul Rahim; A343: U.

littoralis (K, SING); A446: A. suaveolens (K, L, SING); A493: Fiss. fulgens (A, K,

SING). Abu Bakar; 4241: 4. swaveolens (K, SING). Afriastani, J.J.; 1200: U. littoralis

(K). Alston, A.H.G.; 13100: Fr excisa (BM). Ambriansyah; AA 647: 4. macropodus

(K, L). Ambriansyah & Arbainsyah; AA 145: M. kentii (K); AA 223: Fiss. latifolium

(K); AA 279: A. ochropetalus (K, L); AA 284: U. borneensis (K); AA 286: U. lobbiana

(K); AA 336: M. clementis (K); AA 486: U. javana (K); AA 648: D. chinensis (K); AA

724: M. clementis (K); AA 987: U. argentea (K); AA 1176: D. chinensis (K); AA 1272:

A. ochropetalus (K, SAN); AA 1646: D. dumosus (K, L); AA 1660: U. excelsa (K); AA

1675: Fiss. elmeri (K, SAN); AA 1703: M. clementis (K); AA 1822: D. dumosus (K,

SAN); AA 2132: Fr. borneensis (K). Ambri[ansyah] & Arifin; AA 337: Fr. biglandulosa

(K, L); AA 338: M. clementis (K); AA 345: U. lobbiana (K); AA 349: A. ochropetalus

(K, L); AA 474: U. monticola (K); AA 934: U. littoralis (K); W 344: Fr. glauca (L); W

705: Fiss. manubriatum (K); W 811: U. littoralis (K); W 815: A. suaveolens (K); W 871:

Fiss. manubriatum (K); W 876 (K): Fiss. manubriatum (K); W 995: U. monticola (K).

Ambri et al.; AA 1219: A. suaveolens (SAN); AA 1482: U. littoralis (K); AA 1498: A.

suaveolens (SAN); AA 1590: A. suaveolens (L, SAN). Amdjah; 005: A. suaveolens (K);

50: A. suaveolens (K); 805: A. suaveolens (K); 865: U. lobbiana (K); 878: D. acutus (A, K,

SING); 883: Fr. korthalsiana (L); 885: A. suaveolens (L); 914: U. littoralis (K). Andau,

D.; 242: U. grandiflora (K); 270: U. monticola (K); 303: A. suaveolens (K, SAN); 590: A.

suaveolens (K); 817: U. littoralis (K); 888: U. littoralis (K). Anderson, J.A.R.; 4024: D.

dumosus (SAR); 4201: Fiss. montanum (K, L, SAN, SAR, SING); 8396: D. chinensis (K,

SAR, SING); 8562: M. dielsii (K, SING); 9289: P. prismatica (K, SING); 9708: M. dielsii

(K); 9730: P. prismatica (K, SING); 12532: U. hirsuta (K); 13285: A. sarawakensis (K,

L). Andrews, S.; 737: U. monticola (K); 778: A. costatus (K). Angian; 7757: D. chinensis

(K, SING); 10481: U. littoralis (L, SING). Apostol; 6743: U. littoralis (K, SING); 7688:

D. chinensis (K, SING). Arbainsyah; AA 1851: A. gracilis (K); 1945: U. excelsa (K).

Argent, G.; 93132: P. prismatica (K); 93163: P. prismatica (K); 94120: U. schefferi (K).

Argent, G. & Ruskani; 5013: 4. ochropetalus (A, K). Argent, G. & Saridan, A.; 9326:

Fr. affinis (K, L); 9376: A. ochropetalus (K, L SAN); 9390: U. verrucosa (K). Argent, G.

et al.; 93123: A. ochropetalus (K, L, SAN). Arifin, Z. & Ambriansyah; B 1539: Fiss.

elmeri (K, SAN). Arifin, Z. & Arbainsyah; B 1344: 4. polygynus (L). Arifin & Insyah;

AA 1049: A. suaveolens (L, SAN). Arfin et al.; AA 1106: A. suaveolens (SAN); AA 1697:

U. grandiflora (K). Arsat; 1064: U. littoralis (A); 1336: U. littoralis (K). Azmi, R.; RA

389: S. affinis (L).

462 Gard. Bull. Singapore 64(2) 2012

Bakia, K.; 299: U. littoralis (K). Balajadia, D.; 3810: U. excelsa (K); 3749: U. littoralis

(K). Barber, E.; 127: U. borneensis (K); 376: D. dunalii (K). Bartlett, E.; sn(1893):

A. maingayi (BM); sn: Fiss. manubriatum (BM); sn: A. gracilis (BM). Bayak; 2117:

U. littoralis (K). Beaman, J.H.; 520: U. excelsa (K); 7103: A. suaveolens (K); 7178: A.

suaveolens (K); 7831: U. littoralis (K); 9306: U. littoralis (K); 11017: A. roseus (K); 11834:

Fiss. crassicaule (K); 11983: A. suaveolens (K); 12406: U. borneensis (K). Beccari, O.;

P.B. 232: Fr. affinis (K); P.B. 318: U. cuneifolia (BO, K); P.B. 381: A. maingayi (K); P.B.

393: P. prismatica (K); P.B. 554: A. roseus (K); P.B. 713: A. maingayi (K); P.B. 802:

Fr. grandifolia (K); P.B. 1041: Fiss. rugosum (K); P.B. 1107: Fr. grandifolia (K); P.B.

1112: Fr. biglandulosa (BM, K); P.B. 1120: U. monticola (K); P.B. 1410: Fiss. rugosum

(K); P.B. 1648: U. beccarii (K); P.B. 1760: U. littoralis (K); P.B. 1811: U. clementis

(K); P.B. 2268: M. kentii (K); P.B. 2911: Fr. glauca (K); P.B. 3190: A. gracilis (K); P.B.

3420: D. acutus (K); P.B. 3571: Fr affinis (K); P.B. 3740: M. kentii (K); P.B. 3742:

Fr. glauca? (K); P.B. 3748: A. suaveolens (K); P.B. 3789: Fiss. multivenium (K); P.B.

3899: M. dielsii (K); P.B. 3945: D. acutus (K); P.B. 3976: U. cuneifolia (K). Bernstein,

J.H.; JHB 520: U. excelsa (K). Brooke, W.M.A.; 8740: Fiss. latifolium (K); 10678: F:

rugosum (BM, L). BRUN Series; BRUN 1: Fr. glauca (K); BRUN 546: Fr borneensis

(K); BRUN 569: D. dunalii (K, SAR); BRUN 606: U. excelsa (K); BRUN 2343: Fiss.

montanum (K, L); BRUN 5209: A. hirtipes (K, L); BRUN 5517: Fr. biglandulosa (K,

L); BRUN 5902: Fr. biglandulosa (K); BRUN 15028: U. excelsa (SAR); BRUN 15251:

A. suaveolens (L, SAN); BRUN 15253: A. maingayi (K, L, SAN); BRUN 15350: U.

lobbiana (K, SAR); BRUN 15462: U. excelsa (K); BRUN 15501: U. monticola (K, SAN,

SAR); BRUN 15504: Fr. glauca (K, SAN); BRUN 15578: Fiss. fulgens (K, SAN, SAR);

BRUN 16219: D. dumosus (K); BRUN 16270: U. littoralis (K, SAN); BRUN 16422: U.

excelsa (SAN); BRUN 16423: Fr: excelsa (K, SAN); BRUN 16430: A. polygynus (L,

SAN); BRUN 16431: A. suaveolens (SAN); BRUN 16521: U. littoralis (SAN); BRUN

16633: P. prismatica (K SAN); BRUN 16644: M. clementis (K, SAN); BRUN 16710: A.

sarawakensis (K, L); BRUN 16924: A. suaveolens (L, SAN); BRUN 16988: M. kentii (K,

SAN); BRUN 17078: M. kentii (K, SAN); BRUN 17310: U. excelsa (K); BRUN 17542:

Fiss. rugosum (SAN); BRUN 17526: U. monticola (K); BRUN 17762: A. ochropetalus

(K); BRUN 115410: P. prismatica? (K). Burley, J.S. & Lee, B.; 272: A. costatus (A,

K, L, SAN, SING); 347: Fr glauca (K, SING). Burley, J.S. et al.; 731: Fiss. elmeri

(K, L, SAR, SING); 830: Fiss. kingii (K, SAR, SING); 832: Fr formosa (K, L, SAR,

SING); 2713: A. hirtipes (K, L); 3288: A. hirtipes (K, L); 3321: Fiss. rugosum (K, SING).

Buwalda, P.; 7839: A. roseus (A, K, L).

Campbell, E.J.F.; EG152: 4. ochropetalus (L); EG228: Fr. biglandulosa (SAN). Castillo, M.;

635: U. monticola (K). Chew, W.L.; CWL 125: U. monticola (K, SAN, SING); CWL 297:

Fr. excisa (K, LSAR, SING); CWL 608: U. lobbiana (K, SING); CWL 684: U. lobbiana

(SAR): CWL 1366: M. kentii (K, SING). Chew, W.L. et al.; 1846: Fiss. montanum (K,

SAN, SING). Church, A.C. et al.; 482: 4. gracilis (L, SING); 483: A. gracilis (L SING);

612: D. dumosus (K, SAR, SING); 661: U. monticola (K, SAR, SING); 774: Fr. affinis

(A, K, SING); 818: Fr. cf. ovalifolia (A, BO); 821: U. clementis (A, SING); 860: Fr.

formosa (A, BO, SING); 969: Fr. formosa (A, BO, K, SING); 983: Fiss. kingii (K, SAR,

SING); 1062: Fr, formosa (A, BO, SING); 1067: Fr formosa (A, BO, K, SING); 1210:

A. suaveolens (SING); 1244: A. hirtipes (A, L); 1275: A. gracilis (L, SING); 1740: A.

hirtipes (A, K, L, SAN); 2671: Fiss. manubriatum (L); 2701: Fiss. elmeri (A, K, L).

Clemens, J. & Clemens, M.S.; 3766: Fr. korthalsiana (BM, K); 20150: Fiss. elmeri

Climbing Annonaceae of Borneo 463

(A, BM, K, SAR); 20347: U. lobbiana (SAR); 20925: A. suaveolens (A, L); 21113: Fr

biglandulosa (A, K); 21270: U. sp. (A); 21276: Fr. glauca (A, SAR); 21341: U. argentea

(A, K, SING); 26008: U. monticola (K); 26205: Fr. korthalsiana (K, L); 26286: Fiss.

kinabaluense (A, K, L); 26377: Fr. biglandulosa (BM, K); 26407: Fiss. kinabaluense (K);

26424: A. polvgynus (K, L); 26431: U. schefferi (K); 26541: U. clementis (K); 26620: A.

costatus (A, K, L); 26701: A. costatus (A, K, L); 26811: Fiss. kinabaluense (K); 26829:

Fr. biglandulosa (BM, K, L); 26831: Fr. biglandulosa (BM, L); 26877: D. acutus (K);

2689 bis: Fiss. kinabaluense (K); 27436: Fr. biglandulosa (BM); 27439: Fiss. latifolium

(K); 27447: Fiss. kinabaluense (K); 28127: Fiss. montanum (K); 28328: Fiss. latifolium

(K); 28435: A. polygynus (SING); 28489: Fiss. latifolium (K); 28674: A. kinabaluensis

(SING); 28697: A. kinabaluensis (A, BM, K, L, SING); 28764: A. kinabaluensis (A, BM,

BO, K, SING); 29306: Fiss. latifolium (BM); 29341: A. kinabaluensis (BM); 29342: A.

costatus (K, L); 30351: A. kinabaluensis (A, K, L); 30367: Fiss. kinabaluense (K); 30368:

Fiss. kinabaluense (K); 30534: Fr. korthalsiana (BM); 30651: Fr korthalsiana (A):

30866: Fr. korthalsiana (A, K); 31616: U. sp. (A, BM, K); 31652: Fiss. latifolium? (A, L);

32448: Fiss. montanum (K); 33978: Fiss. montanum (A, BM); 34022: Fr. korthalsiana

(A, K, L); 40326: A. polygynus (K); 40422: Fiss. carrii (A, K, L); 40491: U. schefferi

(K); 40584: Fiss. kingii (K); 40731: Fr. biglandulosa (BM, K); 40781: Fiss. latifolium

(Kinabalu form) (L); 40874: A. kinabaluensis (BM); 50348: A. kinabaluensis (BM);

50373: A. polygynus (A, L); 50379: A. veldkampii (A, K); 50469: A. polygynus (K, L);

51035: A. polygynus (K); 51310: Fiss. elmeri (A, BM, K); 51624: Fiss. elmeri (A, BM,

K). Clemens, M.S.; 9670: U. clementis (A); 9776: M. clementis (A, K); 9783: U. littoralis

(A); 11158: A. suaveolens (K); 11097: Fr. glauca (A). Coode, M.J.E.; 6353: A. gracilis

(K); 6444: U. littoralis (K, SAN); 6477: Fiss. fulgens (K, SAR, SAN); 6758: U. excelsa

(K); 6770: A. maingayi (K, L, SAN); 6981: U. littoralis (SAN); 7119: U. lanuginosa (K,

SAN); 7353: M. dielsii (K); 7661: U. excelsa (K).

Creagh, C.V.; sn (4/1895): U. grandiflora (K); sn(4/1894): U. littoralis (K); sn(19/4/1895): D.

chinensis (K); sn(4/1895): D. dumosus (K); sn(4/1895): Fiss. fulgens (K). Cuadra, A.; A

1151: A. suaveolens (K).

Darnton, S.; 100: U. littoralis (A). de Jong, W.; 723: P. prismatica (L). de Vriese, W.H.; s.n.

(L0484299): U. cuneifolia (L); s.n. (L0484297): U. cuneifolia (L). Dilmy, A.; 1034: U.

griffithii (L). Dransfield, J.; 6284: U. littoralis (K); 7154: Fiss. paniculatum (K, SAR);

7284: M. kentii (K, SAN); 7320: D. dumosus (K, SAN, SAR); 7346: A. suaveolens (SAN):

7383: Fiss. kingii (K, SAR); 7439: A. suaveolens (SAN); 7447: A. suaveolens (SAN).

Elmer, A.D.E.; 20117: U. littoralis (K, SING); 20118: A. hirtipes (K, SING); 20138: A.

suaveolens (K, SING); 20196: U. littoralis (K, SING); 20197: Fr: grandifolia (BM, K,

SING); 20318: Fiss. fulgens (K, SING); 20338: M. clementis (K); 20407: Fr. borneensis

(BM, K, SING); 20418: Fr. grandifolia (BM, K, SING); 20489: 4. hirtipes (K, SING);

20516: Fiss. latifolium (K, SING); 20525: D. dumosus (K); 20604: U. littoralis (K,

SING); 20732: U. javana (K, SING); 20802: Fr. borneensis (BM, K, SING); 20814: Fiss.

latifolium (K, SING); 20857: U. javana (SING); 20870: U. monticola (K, SING); 20879:

Fr. grandifolia (BM, K, SING); 20881: Fiss. e/meri (K); 21081: U. excelsa (K, SING);

21090: U. concava (K, SING); 21103: Fr glauca? (K, SING); 21167: Fr borneensis

(BM, K, SING); 21181: Fr grandifolia (BM, SING); 21197: A. ochropetalus (K, L,

SING); 21211: U. borneensis (K, SING); 21300: Fiss. paniculatum (A, BM, K, SING);

464 Gard. Bull. Singapore 64(2) 2012

21663: A. ochropetalus (K, L, SING); 21689: Fr. biglandulosa (BM, K, SING); 21721:

U. monticola (K, SING). Endert, F.; 1551: U. excelsa (K); 1610: Fiss. fulgens (K); 1935:

Fiss. fulgens (K); 2234: U. griffithii (K); 2244: M. kentii (K); 2333: U. lobbiana (A,

BO, K); 2423: Fiss. kingii (K); 2511: A. polygynus (A); 2513: U. littoralis (K); 2715:

Fr. korthalsiana (A); 3299: Fiss. kingii (A, K); 3312: M. kentii (K, SING); 3430: A.

ochropetalus (K, L); 3904: Fiss. sp. (L); 4471: A. hirtipes (K); 4793: A. veldkampii (K,

L); 4836: M. kentii (K); 5035: Fiss. multivenium (K); 5169: U. grandiflora (K). Enggoh;

10205: A. suaveolens (SING).

Fabia, M.P.; A3060: Fr: glauca (K, SING). Forman, L.L.; 479: M. kentii (K, SING); 997: U.

littoralis (K, SAN); 1149: Fr. glauca (K); 1151: U. littoralis (SAN). Fraser, M.; 168: U.

littoralis (K); 185: U. littoralis (K). Frodin, D. & Ismawi; 2049: A. suaveolens (K).

Geesink, R.; 9220: A. atractocarpus (L). Geh, S.Y.; GSY 309: Fiss. latifolium (SING). Gibbs,

L.S.; 2580: A. suaveolens (K). Giesen, W.; 137: A. suaveolens (L). Goklin, T.; 2091: D.

chinensis (K); 2094: Fiss. fulgens (K); 2341: D. chinensis (K); 2544: Fiss. fulgens (K);

2727: U. littoralis (K). Goverse, E. & Adriansyah; Berau 446: M. kentii (K). Gregson,

J.; 72: A. ochropetalus (SAN); 100: U. littoralis (SAN); 137: U. littoralis (SAN); Gregson,

J. & Bernardus Bala Ola; 72: A. ochropetalus (K, SAN).

Haegens, R.M.A.P. & Klazenga, N.; 438: A. atractocarpus (K, L, SAR). Haegens, R.M.A.P.

et al.; 445: Fiss. kingii (K). Hallier, H.; 815: M. kentii (K); 1018: U. littoralis (SING);

1080: A. suaveolens (K); 3023: A. hirtipes (L). Haviland, G.D.; 3: A. gracilis (K, SING);

403: U. monticola (K); 411: Fiss. manubriatum (K); 416: Fiss. fulgens (K, SING); 417: U.

monticola (K); 421: P. prismatica (K); 422: U. borneensis (K); 540: U. monticola (SAR);

849: Fiss. latifolium (K, SAR, SING); 877: Fiss. rugosum (K); 1021: Fiss. manubriatum

(SAR); 1310: Fiss. kinabaluense (K, SING); 1504: U. cuneifolia (BO, K, SAR, SING);

1518: A. suaveolens in part (K); 1540: A. suaveolens (K); 1750: A. costatus (K, SING);

1775: A. pandanicarpus (K); 1831: Fr grandifolia (K, SING); 1845: Fiss. paniculatum

(K, SING); 1845bis: Fiss. paniculatum (K); 1968: U. excelsa (K, SING); 2102: Fiss.

longipetalum (BM, K); 2103: Fr excisa (K); 2106: A. hirtipes (K, SING); 2212: Fiss.

manubriatum (SAR); 2213: Fr, excisa (BM, SING); 2216: Fr. excisa (BM, K, L, SING);

2250: U. clementis (A, K, SING); 2315: U. littoralis (K); 2325: U. littoralis (K); 2326:

A. hirtipes (K); 2333: Fr. glauca (BM, K); 3152: Fr. glauca (BM, K, SAR); 3334: U.

lanuginosa (K); 3336: Fiss. kingii (BM, K, SAR, SING); b.y.l.a.: Fiss. rugosum (K);

b.z.d.d.: A. gracilis (K); sn(14/3/1893): A. suaveolens (K); sn(3/1893): Fiss. fulgens

(K); sn(9/1892): A. suaveolens (K); sn(9/1892): U. cuneifolia (K); sn(12/10/1894): A.

roseus (BO); sn: P. prismatica (BM, K). Haviland, G.D. & Hose, C.; 3: 4. gracilis (K);

1629 (26/10/1894): A. maingayi (BM, L); 1629 (13/11/1894): 4. maingayi (K); 1629E

(L0180470): A. polygynus (L); 2106: A. hirtipes (K); 3141: Fr ovalifolia (K, SAR,

SING); 3151: Fr. ovalifolia (K); 3160: A. sarawakensis (K); 3334: U. lanuginosa (SAR,

SING); 3335: Fr, glauca (BM, K, SAR); 3339: A. suaveolens (K. SAR); 3340: A. roseus

(K, SAR); sn (4/12/1894): A. gracilis (K). Hewitt, J.; 40: Fiss. fulgens (SAR); 164: Fr

ovalifolia (SAR); 200: P. prismatica (K); 209: Fr. borneensis (SAR); 347: Fr. borneensis

(SAR); 348: Fiss. kingii (SAR); 538: Fiss. kingii (SAR); 912: Fr. borneensis (SAR);

A.7.13: Fr. ovalifolia (SAR); A.12.16: Fiss. fulgens (SAR). Hose, C.; 83: U. littoralis

(K); 112: U. littoralis (K); 160: A. sarawakensis (BM, K); 180: U. littoralis (K); 302: A.

sarawakensis (BM, K); 397: Fiss. kingii (A, BM, K); 601: U. borneensis (K); 635: Fiss.

fulgens (K). Hotta, M.; 12677: A. roseus (L); 12737: A. roseus (L).

Climbing Annonaceae of Borneo 465

ITTO/BB 0148: U. monticola (SAR).

Jacobs, M.; 5138: U. monticola (K, SAR); 5234: Fiss. kingii (K, SAR, SING); 5362: Fr

biglandulosa (K, L, SAR); 5407: A. polygynus (L, SING); 5481: A. suaveolens (K).

Jaheri; 320: A. Januginosus (BO). Jarvie, J.K. & Ruskandi, A.; 5013: A. ochropetalus

(A, L, SAN); 5293: A. roseus (A, K, L); 5312: A. veldkampii (A, L); 5314: U. excelsa (A);

5319: M. kentii (A K); 5322: P. prismatica (A, K, L); 5730: A. polygynus (A, L); 6008: A.

veldkampii (A, L).

Kadir, A.; 2073: Fiss. kingii (K, SING). Kadir, A. & Enggoh; 10348: Fr. korthalsiana (K,

SING). Kalat, A.; ARK 26: U. lobbiana (K); ARK 104: U. excelsa (K, SAN, SAR).

Kandilis; 6245: 4. suaveolens (A, K, L, SING). Kartawinata, K.; 919: Fr grandifolia(K).

Kato, M. et al.; 20643: M. clementis (A); 20731: U. lobbiana (A); 23423: A. suaveolens

(A); 23425: D. acutus (BM). KEP series; KEP 80018: Fiss. latifolium (K, SING). Kessler,

P.J.A.; PK 287: A. roseus (L); 548: A. ochropetalus (L); PK 598: A. ochropetalus (K, L).

Kessler, P.J.A. et al.; PK 353: U. littoralis (K); PK 617: Fr. biglandulosa (L); PK 625: Fr.

borneensis (L); PK 648: U. littoralis (K); PK 819: U. javana (K); PK 833: U. javana (K);

PK 868: U. javana (K); PK 890: A. suaveolens (K); PK 904: A. suaveolens (SAN); PK

908: M. clementis (K); PK 914: U. borneensis (K); PK 946: U. borneensis (K); PK 955:

Fiss. manubriatum (L); PK 958: M. clementis (SAN); PK 993: A. ochropetalus (K, L,

SAN); PK 997: A. ochropetalus (K, L, SAN); PK 1009: M. clementis (K, SAN); PK 1040:

M. clementis (L); PK 1044: D. chinensis (K); PK 1046: Fr. borneensis (K, L, SAN); PK

1047: Fr. glauca (K); PK 1058: A. ochropetalus (K, L, SAN); PK 1063: Fiss. kingii (K);

PK 1074: Fiss. manubriatum (K); PK 1103: U. littoralis (SAN); PK 1141: U. grandiflora

(SAN): PK 1157: M. clementis (K, SAN); PK 1160: U. littoralis (SAN); PK 1170: U.

grandiflora (K, SAN); PK 1176: D. chinensis (K, SAN); PK 1180: U. micrantha(K, SAN);

PK 1188: U. javana (K); PK 1200: Fiss. multivenium (K, SAN); PK 1211: 4. suaveolens

(SAN); PK 1213: U. grandiflora (K); PK 1224: U. lobbiana (K); PK 1228: M. clementis

(K, SAN); PK 1229: A. suaveolens (L, SAN); PK 1342: A. suaveolens (SAN); PK 1358:

U. grandiflora (K); PK 1362: M. clementis (K, SAN); PK 1364: A. suaveolens (SAN):

PK 1369: U. grandiflora (K); PK 1374: A. suaveolens (SAN); PK 1410: P. prismatica (K,

SAN); PK 1434: A. suaveolens (L, SAN); PK 1441: Fr borneensis (K, SAN); PK 1454:

Fiss. manubriatum (SAN); PK 1457: D. dumosus (K, SAN); PK 1745: A. suaveolens (L):;

PK 1746: U. littoralis (K, SAN); PK 1757: D. chinensis (K); PK 1855: M. clementis (K);

PK 1867: U. littoralis (K); PK 1922: A. suaveolens (L); PK 1928: A. suaveolens (L); PK

2018: Fr. borneensis (K, SAN); PK 2099: Fiss. manubriatum (L); PK 2293: Fr. excisa (K,

SAN); PK 2312: M. clementis (K); PK 2314: U. excelsa (SAN); PK 2361: M. clementis

(K); PK 2362: A. suaveolens (SAN); PK 2384: D. chinensis (K, SAN, SAR); PK 2425:

A. suaveolens (L); PK 2621: Fr. affinis (BO, K, L, SAN, SAR); PK 2728: A. gracilis

(K); PK 2740: M. clementis (K); PK 2799: M. clementis (K); Berau 20: M. clementis (K,

SAN); Berau 133: A. Januginosus (K, SAN); Berau 241: U. littoralis (K); Berau 254: A.

suaveolens (L); Berau 276 (K): U. javana (L); Berau 446: M. kentii (SAN); Berau 867:

A. polygynus (BO, K, L, SAN); Berau 1384: Fiss. manubriatum (SAN); Berau 1436:

U. littoralis (K, SAN); W 898: Fiss. kingii (K). Keith, H.G.; 3102: U. monticola (K).

Kirkup, D.W.; 330: 4. lanuginosus (K, L, SAN); 674: A. hirtipes (K); 825: U. littoralis

(K). KL series; KL 3598: U. monticola (K); KL 3600: U. monticola (SING); KL 3602:

Fiss. kingii (K); KL 3616: U. lobbiana (SING). Kokawa, S.; 6336: U. littoralis (SAN);

6345: A. suaveolens (L, SAN); 6347: U. littoralis (SAN). Kokawa, S. & Hotta, M.;

466

Gard. Bull. Singapore 64(2) 2012

435: U. curvistipitata (SAN); 2605: D. chinensis (SAN); 2751: A. suaveolens (L, SAN).

Korthals, P.W.; sn: D. chinensis (K); sn(9/1864): D. dumosus (K); sn (L0182331): Fr

korthalsiana (L); sn (L0182332): Fr. korthalsiana (L); sn (L0037935): Fr. borneensis (L);

sn (L0037936): Fr. borneensis (L); sn (L0037937): Fr. borneensis (L); sn (L0037938):

Fr. borneensis (L); sn (L0187212): Fr. borneensis (L); sn (LO187213): Fr, borneensis

(L); sn (L0182284): Fr glauca (L); sn (L0182285): Fr glauca (L); sn (L0182286): Fr.

glauca (L); sn: U. argentea (L); sn (L0180796): A. sumatranus (L); sn (LO180797): A.

sumatranus (L); sn (L0180798): A. swmatranus (L); sn (L0187136): Fr. biglandulosa (L);

sn (L0187137): Fr biglandulosa (L); sn (L0182243): Fr. biglandulosa (L); sn (L0182244):

Fr. biglandulosa (L); sn (L0187138): Fr. biglandulosa (L); sn (LO182262): Fr: ovalifolia?

(L); sn (L0182131): Fiss. manubriatum (L); sn (L0182132): Fiss. manubriatum (L);

sn (L0182133): Fiss. manubriatum (L); sn (LO182134): Fiss. manubriatum (L); sn

(L0187006): Fiss. manubriatum (L); sn (LO187007): Fiss. manubriatum (L); sn

(L0187009): Fiss. manubriatum (L); sn (L0186729): Fiss. latifolium (L); sn (L0186730):

Fiss. latifolium (L); sn (L0186731): Fiss. latifolium (L); sn (L0187083): P. prismatica (L).

Kramadibrata, K.; 142: M. clementis (K). Kostermans, A.J.G.H.; 4266: A. ochropetalus

(K, L); 4272: A. suaveolens (SING); 4354: A. ochropetalus (K, L); 4375: A. ochropetalus

(K, L, SING); 4452: M. kentii (K); 4508: U. borneensis (K); 4665: D. acutus (K, SING);

4666: U. concava (A, SING); 4733: U. excelsa (K, SING); 4919: U. lobbiana (A, K,

SING); 5001: A. suaveolens (SING); 5176: M. kentii (IK); 5497: Fr. affinis (K, L, SING);

5710: Fiss. multivenium (L); 5793: U. monticola (A, K, SING); 5924: Fr. formosa (BO,

K); 5975: Fr formosa (BO, K, SING); 6412: P. prismatica (K); 6820: A. sumatranus

(BM, K); 6909: A. suaveolens (SING); 6961: U. grandiflora (A, K, SING); 6963: U.

monticola (K); 7107: Fiss. manubriatum (K); 7965: Fiss. manubriatum (K); 8017: M.

dielsii (IK); 8034: M. kentii (K); 8648: M. clementis (K, SING); 8788: Fiss. kingii (K,

SING); 8801: U. monticola (L); 9180: P. prismatica (K, L, SING); 9577: A. ochropetalus

(K, L, SING); 9675: Fr. excisa (K, SING); 9800: P. prismatica (K); 9921: Fiss. kingii (K);

10165: A. suaveolens (SING); 10172A: M. kentii (K); 10495: U. monticola (K, SING);

10562: A. gracilis (L, SING); 10594: Fiss. kingii (K); 10764: M. kentii (L); 12620: Fiss.

kingii (L); 13473: A. veldkampii (K, L); 13732: A. ochropetalus (K, L, SING); 13905:

Fiss. elmeri (K, SING); 21245: U. monticola (K); 21257: D. chinensis (K, SING); 21280:

D. chinensis (K, SING); 21657: M. kentii (K); 21682: U. lobbiana (K); 21683: M. kentii

(K); 82017: P. prismatica? (K).

Laman, T.; TL 225: U. monticola (A); TL 263: M. kentii (K); TL 274: M. kentii (K); TL 877:

U. monticola (A, K); TL 951: Fiss. fulgens (A); TL 971: Fr. biglandulosa (K); TL 996: U.

monticola (A); TL 1223: Fr. borneensis (A, K, L); TL 1237: U. monticola (A); TL 1246:

A. ochropetalus (K, L); TL 1299: Fr: excisa (A, K); TL 1413: Fiss. fulgens (L); TL 1763:

Fiss. multivenium (K). Lamb, A.; 305: U. littoralis (SAN); ALFB 111/87: U. grandiflora

(K). Latupeirissa, E.R.; 95012: Fiss. latifolium (L, SAN). Leeuwenberg, A.J.M. &

Rudjiman; 13048: Fiss. longipetalum (L); 13418: Fiss. manubriatum (L); 13428: Fiss.

paniculatum (L); 13431: A. suaveolens (L); 13433: A. suaveolens (L). Leeuwenberg,

A.J.M. et al.; 14504: 4. swaveolens (L, SAN). Leighton, M.; 159: Fr: affinis? (L); 214:

mixed coll. A. polygynus & A. ochropetalus (L); 242: Fr. biglandulosa (L); 259: Fr. affinis

(L); 353: Fr. affinis (K, L); 420: A. roseus (L); 1068: Fr. grandifolia (L). Lugas, L.; 73:

U. littoralis (K); 131: U. littoralis (K); 179: A. suaveolens (K); 310: U. littoralis (K); 457:

A. costatus (K); 713: M. clementis (K); 1481: Fr grandifolia (K); 1708: A. costatus (K,

SAN); 1742: U. littoralis (K, SAN); 1761: A. costatus (K, SAN); 1763: Fiss. multivenium

(SAN); 2295: Fr. excisa (K); 2612: A. costatus (K).

Climbing Annonaceae of Borneo 467

Mahyar, U.W.; 982: U. lobbiana (A, SING). Maidim, S. Md.; 1529: U. littoralis (K); 1530:

A. suaveolens (K); 2339: A. suaveolens (K). Mail, A.: 3972: A. ochropetalus (K, SING);

7754: Fiss. fulgens (K, SING). Mat Salleh, K.; 1740: Fr: glauca (SAN); 1955: U.

cuneifolia (K, SAN); 2119: U. grandiflora (SAN); 2414: M. kentii (K); 3333: U. excelsa

(A, SAN). Md. Shah & Kadim; MS 1022: U. micrantha (K, SING). Md. Tahir: 715: S.

affinis (A). Meijer, W.; 2023: Fr. borneensis (K, L); 2235: Fiss. kingii (K). Melegrito;

3353: U. littoralis (K). MB 449: U. grandiflora (K). Mogea, J.; 3517: Fr. glauca (K);

4016: A. macropodus (L); 4023: U. monticola (K). Motley, J.; 15: D. chinensis (K); 39:

Fiss. manubriatum (K); 76: U. littoralis (K); 127: U. borneensis (K); 173: U. littoralis

(K); 326: D. dunalii (K). Museum Dyaks; 245: Fiss. latifolium (SAR); 260: Fiss. fulgens

(SAR); 381: U. cuneifolia (SAR). Museum Saifatt; 669: U. cuneifolia (SAR).

Naiang; 2488: U. hirsuta (SAR). Nais, J.; 3676: U. littoralis (SAN). Nangkat, N.: NN 64:

U. littoralis (K, SAN); NN 239: A. suaveolens (L, SAN).Native collector; 159: Fiss.

manubriatum (K); 226: Fiss. latifolium (A, K); 852: Fiss. latifolium (A); 1794: A.

suaveolens (A). Nooteboom, H.P.; 1207: Fiss. kingii (K, L, SAN): 1314: D. dumosus (L).

Nooteboom, H.P. & Chai, P.; 1823: A. macropodus (L, SAR).

Ogata, K.; Og-B 407: A. suaveolens (L); 11392: Fiss. kingii (L).

Paie, I.; 8464: Fiss. fulgens (SAR, SING). Pannell, C.M.; 2567: A. suaveolens (K). Pascual,

G.; 1048: U. monticola (A). Pereira, J.T.; JTP 213: U. concava (A, SAN); JTP 396: U.

monticola (K, SAN); JTP 509: A. suaveolens (L, SAN); JTP 514: U. littoralis (SAN).

Poklin; 2727: U. littoralis (K). Poore, D.; H66: Fiss. montanum (K). Prance, G.T.;

30612: U. excelsa (K); 30698: A. hirtipes (K, L, SAN); 30714: Fr biglandulosa (L,

SAN). Puasa; 1356: U. littoralis (K); 2746: D. chinensis (K). Puasa & Angian; 3828:

Fiss. kinabaluense (K). Purseglove, J.W.; P 3190: A. gracilis (K); P 4419: A. suaveolens

(K, SING); P 4679: Fiss. paniculatum (A, K, L, SAR, SING); P 4965: U. excelsa (K,

SAR, SING); P 5133: U. monticola (K, SAR, SING); P 5205: Fr: glauca (SAR, SING); P

5402: A. gracilis (L, SING); P 5459: U. littoralis (SING).

Rahayu, M.; 652: Fr. biglandulosa (K); 692: Fr. borneensis (K). Ramos, M.; 1143: Fiss.

fulgens (K); 1170: Fr. grandifolia (K); 1171: Fr. korthalsiana (K); 1178: A. suaveolens

(A, BM, K, L); 1281: Fiss. elmeri (A); 1366: A. polygynus (A, BM, BO, K, L); 1380: A.

gracilis (L); 1389: A. ochropetalus (K, L); 1465: A. hirtipes (BM, K); 1474: M. clementis

(A, K, L); 1480: U. clementis (K); 1499: Fiss. latifolium (A, K L); 1667: U. clementis

(K); 1734: A. gracilis (K); 1793: Fiss. fulgens (K); 1910: Fr. grandifolia (BO); 1911:

Fr. grandifolia (BM, K); sn: Fr. grandifolia (K). Reksodihardjo, S.; 17: U. littoralis

(K); 90: Fiss. manubriatum (K); 97: P. prismatica (K); 103: M. clementis (K); 731: Fr.

biglandulosa (L). Richards, P.W.; 1192: Fiss. kingii (K, SING); 1217: Fr glauca (K,

SING); 1258: A. suaveolens (K, SING). Ridley, H.N.; 12464: D. dumosus (BM, K, SING);

sn(_/1/1915): U. borneensis (K); sn(_/1/1915): D. chinensis (K). Ridsdale, C.E.; 1980:

Fiss. bygravei (A, K, L, SAN); 2047: U. excelsa (A, K, SAN); Cerol/27: A. macropodus

(L). RSNB series; 4080: Fiss. montanum (K, SAN, SING); 4285: Fiss. montanum (K,

SAN, SING); 4431: 4. kinabaluensis (K, L, SAN, SING); 4881: 4. costatus (K); 8382: A.

kinabaluensis (K, SING). Rumutom, M.; 163: A. suaveolens (K); 244: U. concava (K);

279: Fr. korthalsiana (K); 530: U. concava (SAN).

468 Gard. Bull. Singapore 64(2) 2012

S series; S 0436: Fiss. paniculatum (SING); S 4024: D. dumosus (SAR); S 5277: Fiss. rugosum

(SAR); S 5387: Fr. borneensis (K, SAR); S 5902: Fr. biglandulosa (SAR); S 7835: Fiss.

latifolium (K, L, SAR); S 8464: Fiss. fulgens (K); S 9114: A. suaveolens (K, SING); S

11021: M. kentii (K); S 11999: Fr: biglandulosa (K, L, SAR, SING); S 12090: U. littoralis

(K); S 12319: M. dielsii (K, SING); S 12403: Fr: affinis (SAR, SING); S 12428: M. dielsii

(K, SING); S 12519: U. lobbiana (SAR); S 12532: U. hirsuta (SING); S 13188: U.

lobbiana (SAR); S 13261: A. suaveolens (K, SAN, SING); S 13262: Fiss. latifolium (K,

SAN, SAR, SING); S 13285: A. sarawakensis (SING); S 13286: U. lanuginosa (K,

SING); S 13288: Fiss. kingii (K, L, SAR); S 13728: U. monticola (K, SAR); S 13755: Fr.

glauca (K); S 13756: U. monticola (SAR); S 13775: Fr borneensis (SAR, SING); S

14366: U. monticola (SAR, SING); S 14780: U. beccariti (K, SAR); S 15965: Fr.

biglandulosa (K, L, SAN, SAR, SING); S 15966: Fiss. latifolium (K, SAN, SAR, SING);

S 16030: U. grandiflora (SAR); S 16279: U. littoralis (K, SAN, SING); S 16300: U.

littoralis (K); S 16323: Fr. biglandulosa (A, K, L, SAN, SAR, SING); S 16410: U.

beccarii (K, SAR, SING); S 16417: U. excelsa (K, SAR, SING); S 16666: Fiss. kingii (A,

K, L, SAN, SAR, SING); S 16969: A. roseus (L); S 18364: M. kentii (K, SING); S 18434:

Fr, excisa (K, SAN, SAR, SING): S 18493: U. lobbiana (A, K, SAR, SING); S 19129: Fr

excisa (K, SAR); S 19270: A. hirtipes (K); S 19561: D. acutus (K, SAN, SING); S 19647:

Fr. biglandulosa (K, SAR, SING); S 20272: U. lobbiana (SAR); S 20889: A. hirtipes (A,

K, L, SING); S 20923: A. venustus (L); S 20991: P. prismatica (K); S 21376: U. excelsa

(K, SAN, SAR, SING); S 21469: U. monticola (A, K, SAR, SING); S 21549: Fiss.

manubriatum (K); S 21634: U. littoralis (SAR); S 22561: M. dielsii (SING); S 22619: U.

excelsa (A, K, SING); S 22770: Fiss. crassicaule (K, L, SAR); S 23003: U. monticola (A,

K, SAR, SING); S 24371: Fiss. kingii (K, SAN, SAR, SING); S 24726: M. kentii (A, K,

SAN, SING); S 25265: Fiss. kingii (K, L, SAN, SAR, SING); S 25281: M. kentii (K,

SING); S 25409: A. suaveolens (K, SING); S 25414: Fiss. rugosum (SAR); S 25433: A.

suaveolens (K, SING); S 25436: Fiss. paniculatum (K, SAR, SING); S 25440: Fiss.

paniculatum (SAR); S 25448: Fr. borneensis (K, SAR, SING); S 25560: Fr. excisa (K,

SAR, SING); S 25629: D. acutus (K, SAR, SING); S 26041: U. lanuginosa (SAR); S

26258: Fr. ovalifolia (SAN, SAR, SING); S 26861: P. prismatica (K, SING); S 26894: P.

prismatica (K); S 26958: A. venustus (L); S 27003: Fiss. rugosum (K, SAR); S 27068: A.

hirtipes (K, L); S 27180: Fiss. latifolium (K, SAN, SAR, SING); S 27220: Fiss. kingii (K,

SAR); S 27247: Fiss. kingii (K, SAR, SING); S 27553: U. concava (SAR); S 27588: Fiss.

fulgens (A, SAR, SING); S 27599: U. littoralis (A, K, SAN, SING); S 27674: U. clementis

(SAR); S 27677: Fiss. paniculatum (K, SAR, SING); S 27722: U. monticola (K, SAR);

27749: A. veldkampii (K, L); S 27795: Fr. ovalifolia (K, SAR); S 27914: Fr borneensis

(SAR); S 27935: Fr. glauca (K, SAN, SAR, SING); S 28010: A. maingayi (K, L, SAN);

S 28025: A. pandanicarpus (K, L, SAN); S 28103: Fr. formosa (K, SAR, SAN, SING); S

28185: Fr. affinis (K, SING); S 28488: A. suaveolens (K, SING); S 28541: A. suaveolens

(K, SING); S 28701: U. monticola (SAR); S 29261: U. monticola (SAR); S 29583: Fiss.

longipetalum (K, L, SAR); S 29684: A. maingayi (A, K, L, SAN); S 29863: U. lobbiana

(SAR); S 30047: Fiss. latifolium (K, SAR, SING); S 31124: U. monticola (K, SAR,

SING); S 31543: Fiss. kingii (K, SAN, SAR, SING); S 31576: U. lobbiana (SAR); S

31679: U. grandiflora (SAR); S 31743: U. verrucosa (K, SAN, SAR, SING); S 31744:

Fiss. kingii (K, SAR, SING); S 32005: M. kentii (K, SAN); S 32106: Fiss. latifolium

(SAR); S 32139: A. sarawakensis (L, SING); S 32176: M. kentii (K, SAN, SING); S

32369: U. borneensis (K, SING); S 33129: U. cuneifolia (A, K, SAR); S 33297: A.

maingayi (A, K, L); S 33431: Fr glauca (K, SAR); S 33571: Fiss. rugosum (K, SAR); S

Climbing Annonaceae of Borneo 469

33748: U. lobbiana (A, SAR); S 34274: Fr. borneensis (K, SAR); S 34349: U. monticola

(K, SAR); S 34995: U. monticola (SAR); S 35028: Fr. formosa (K, SAR); S 35073: Fiss.

elmeri (A, K, SAR); S 35143: Fiss. paniculatum (SAR); S 35288: U. cuneifolia (A, K,

SAR); S 35345: Fiss. montanum (K, L, SAR); S 35632: Fiss. paniculatum (K, SAR); S

35681: U. monticola (K, L, SAR); S 35743: Fiss. paniculatum (K, SAR); S 35788: U.

monticola (K, SAR); S 36639: Fiss. rugosum (K, SAN, SAR); S 36712: M. kentii (K); S

36716: Fiss. manubriatum (K, SAR); S 36727: U. monticola (SAR): S 36826: U. lobbiana

(K, SAR); S 36863: U. monticola (SAR); S 36885: Fiss. paniculatum (K, SAR): S 36894:

A. veldkampii (L); S 37247: Fr. biglandulosa (K, L, SAN, SAR); S 37307: U. cuneifolia

(L, SAR); S 37445: U. clementis (K); S 37958: U. beccarii (K, SAN, SAR); S 38404: M.

kentii (K, SAN); S 38446: A. suaveolens (K, L, SAN); S 39344: U. schefferi (K, SAR); S

39566: U. monticola (SAN, SAR); S 39705: M. kentii (K); S 39840: Fiss. brevistipitatum

(K, SAR); S 40078: U. lobbiana (SAR): S 40121: D. acutus (K, SAN, SAR); S 40123: U.

lobbiana (K, SAN, SAR); S 40553: U. lobbiana (SAR): S 40656: A. costatus (K, L,

SAN); S 40742: U. schefferi (K, SAR); S 40773: Fiss. rugosum (K, SAN, SAR); S 41241:

Fr. glauca (K, SAR); S 41259: U. borneensis (K, SAN SAR); S 41890: U. littoralis (K,

SAN, SAR): S 41931: Fiss. manubriatum (SAR); S 42357: Fr. formosa (K, SAN, SAR):

S 42563: Fr. biglandulosa (K, L, SAR); S 42624: Fiss. manubriatum (K, SAN, SAR): S

43014: A. suaveolens (K, SAN); S 43022: M. kentii (K, SAN); S 43387: Fiss. longipetalum

(K, SAN, SAR): S 43494: Fr. borneensis (K, SAN, SAR): S 43619: Fr borneensis (K, L,

SAR); S 43645: U. monticola (SAR); S 43716: Fr. borneensis (K, L, SAR); S 43726: U.

sp. (SAN); S 43894: A. suaveolens (K, SAN); S 43907: A. roseus (L, SAN); S 43926: U.

verrucosa (SAR); S 44015: U. monticola in part (L, SAR); S 44372: Fiss. fulgens (K, L,

SAR); S 45260: U. lobbiana (SAR); S 45276: A. veldkampii (L, SAN); S 45350: A.

suaveolens (K, LSAN); S 45583: Fr. biglandulosa (K, L, SAN, SAR); S 45667: A. roseus

(K, L, SAN); S 45671: Fr. borneensis (K, SAN); S 46308: Fiss. latifolium (K, L, SAN,

SAR): S 46801: P. prismatica (K); S 46893: Fr. biglandulosa (K, L, SAR); S 47417: Fiss.

manubriatum (K, SAR); S 47651: Fiss. montanum (K, SAR); S 48014: A. atractocarpus

(K, L, SAN); S 48037: Fiss. multivenium (K, SAR): S 48054: A. suaveolens (L); S 48169:

U. cuneifolia (K, SAN, SAR); S 48343: Fr. biglandulosa (K, SAN, SAR); S 48393: U.

monticola (SAR); S 49037: Fr glauca (K, SAR); S 49373: U. lobbiana (K, SAN, SAR);

S 49458: U. monticola (K, SAR); S 49548: Fiss. elmeri (K, L); S 50454: Fr. biglandulosa

(K, SAR); S 51016: A. veldkampii (K, L, SAN); S 51299: U. monticola (SAR); S 51463:

Fiss. kingii (K, SAN, SAR); S 51561: A. suaveolens (K, L, SAN); S 52314: U. monticola

(K, SAN, SAR); S 52340: U. borneensis (K, SAN); S 52509: M. clementis (K, SAN,

SAR); S 52659: U. littoralis (SAR); S 52668: U. cuneifolia (SAN, SAR); S 53355: Fr

biglandulosa (K, SAN, SAR); S 53500: A. atractocarpus (L); S 53516: Fr. biglandulosa

(K, SAR); S 53725: Fr. biglandulosa (K, SAN, SAR); S 53908: Fiss. kingii (K, SAR); S

54000: U. schefferi (SAR); S 54564: A. maingayi (K, L, SAN); S 54662: U. excelsa (K,

SAR); S 56453: U. monticola (SAR); S 57104: Fiss. manubriatum (K, L); S 57164: Fiss.

kingii (K, SAN, SAR); S 57672: Fiss. kingii (SAN, SAR); S 57834: Fiss. bygravei (K, L,

SAN, SAR); S 58599: U. lobbiana (SAR); S 58897: Fiss. kingii (SAR); S 59637: Fiss.

fulgens (SAR): S 59936: M. kentii (K, SAN); S 60572: Fiss. kingii (SAR); S 60776: A. cf.

lanuginosus (L); S 60849: Fiss. montanum (SAR); S 60921: U. schefferi (SAR); S 61120:

Fiss. kingii (K, L, SAN, SAR); S 61465: Fiss. multivenium (K, SAR); S 62313: A.

atractocarpus (L, SAN); S 62346: Fr. glauca (K, SAR); S 63087: Fr: glauca (SAR); S

63251: U. monticola (SAR); S 63627: Fiss. bygravei (K, L, SAN, SAR); S 63673: Fr.

biglandulosa (SAN, SAR); S 63698: Fr. glauca (K, SAR); S 64063: A. ochropetalus (K,

470

Gard. Bull. Singapore 64(2) 2012

L, SAN); S 64120: U. schefferi (SAR); S 64121: U. schefferi (SAR); S 64648: U.

monticola (K, SAR); S 64929: Fr: glauca (K, SAR); S 64963: Fiss. latifolium (SAR); S

65020: Fr. ovalifolia (SAR); S 65060: Fiss. kingii (K, L, SAN, SAR); S 65838: Fiss.

kingii (SAR); S 65840: Fiss. kingii (SAR); S 65898: Fr. borneensis (SAR); S 65972: Fr.

borneensis (SAR); S 65995: U. cuneifolia (SAR); S 66213: U. javana (K, SAN, SAR); S

66268: Fr. borneensis (SAR); S 66917: Fiss. longipetalum (SAR); S 67395: U. clementis

(SAR); S 67557: A. sarawakensis (K, L, SAN); S 68733: Fiss. rugosum (K, SAR), S

68936: U. excelsa (K, SAN, SAR): S 69529: D. dumosus (SAR); S 69940: U. monticola

(SAR); S 70965: Fr. biglandulosa (SAR); S 70982: U. ?borneensis (SAR); S 71221: U.

cuneifolia (SAN, SAR); S 71304: Fr. biglandulosa (SAR); S 71502: U. cuneifolia (SAR);

S 71542: Fiss. kingii (SAR); S 71629: U. schefferi (SAN, SAR); S 71664: A. hirtipes

(SAR); S 71748: Fiss. latifolium (SAR); S 71954: Fiss. montanum (SAR); S 72129: A.

costatus (SAR); S 72196: Fiss. latifolium (SAR); S 72432: Fiss. cf. kingii (L, SAN, SAR,

SING); S 72541: U. beccarii (SAN, SAR); S 72588: Fiss. rugosum (SAR); S 72718: Fr.

ovalifolia (SAR); S 72967: U. lobbiana (SAR); S 73364: Fiss. manubriatum (K, SAR); S

73705: A. suaveolens (K); S 73763: A. gracilis (SAN); S 74389: U. monticola (SAR); S

74644: U. monticola(SAR); 8 74973: Fiss. manubriatum (SAR); S 75099: A. atractocarpus

(K, SAN, SAR); S 76671: Fr borneensis (SAR); S 77208: Fiss. kingii (SAN, SAR,

SING); S 78109: Fiss. rugosum (SAR); S 78263: Fiss. rugosum (SAR); S 78272: Fr

biglandulosa (SAR); S 78316: U. monticola (SAR); S 78747: Fr. borneensis (SAR); S

79056: A. sarawakensis (SAR); S 79113: Fr borneensis (SAR); S 79426: Fiss. kingii

(SAR): S 79464: U. lobbiana (SAR); S 79995: D. acutus (SAN, SAR); S 80674: U.

excelsa (SAN, SAR); S 80953: A. suaveolens (L, SAR); S 81252: D. dumosus (SAR); S

81280: A. costatus (L, SAN, SAR); S 81317: U. monticola (SAR); S 81603: U. cuneifolia

(SAR); S 81631: Fiss. manubriatum (SAR); S 82117: U. lobbiana (SAR); S 82539: Fr

excisa (SAR); S 82825: Fiss. fulgens (SAR); S 83219: U. concava (SAR); S 83237: U.

lobbiana (SAR); S 83331: Fiss. rugosum (L, SAN, SAR); S 83363: Fiss. longipetalum

(SAR); S 83388: U. lobbiana (SAR); S 83406: U. monticola (SAR); S 83484: A.

suaveolens (L); S 83496: Fiss. longipetalum (L, SAR); S 83550: U. lobbiana (SAR); S

83584: U. schefferi (SAR); S 84231: U. cuneifolia (SAR); S 84434: Fiss. rugosum (SAR);

S 84552: Fiss. rugosum (SAR); S 85820: U. excelsa (SAR); S 85883: U. cuneifolia

(SAR); S 86189: Fr. biglandulosa (SAR); S 87259: U. excelsa (SAR); S 87265: Fr.

ovalifolia (SAR); S 87300: D. dumosus (SAR); S 87644: Fiss. manubriatum (SAR); S

87683: Fiss. kingii (SAR); S 88105: A. atractocarpus (SAR); S 89284: U. lobbiana

(SAR); S 89404: U. grandiflora (SAR); S 89703: Fr. ovalifolia (SAR); S 90803: U.

monticola (SAR); S 90815: U. monticola (SAR); S 91077: U. monticola (SAR): S 91589:

Fiss. paniculatum (SAR, SING); S 91755: U. monticola (SAR); S 93100: U. excelsa

(SAR); S 96230: U. lobbiana (SAR). SAN series; SAN A 2920: A. ochropetalus (K);

SAN A 4226: A. ochropetalus (K); SAN A 4291: U. borneensis (A); SAN A 4380: A.

veldkampii (A, L); SAN 2699: U. littoralis (SAN); SAN 3382: U. littoralis (SAN); SAN

7374: Fiss. elmeri; SAN 11508: Fr. affinis (SAN); SAN 16390: A. suaveolens (A, L,

SING); SAN 16546: Fr: glauca (K, L, SING); SAN 17053: Fr: biglandulosa (K, L); SAN

17070: A. suaveolens (L); SAN 17079: U. verrucosa (K); SAN 17103: P. prismatica (A,

SAN, SING); SAN 17134: U. verrucosa (K, SAN); SAN 17316: A. ochropetalus (K,

SAN); SAN 17403: U. monticola (A, SING); SAN 17404: A. hirtipes (A, K, L, SING);

SAN 17507: P. prismatica (A, K); SAN 17597: Fr. biglandulosa (A, L, SING); SAN

18566: M. clementis (K, SAN); SAN 19255: Fr glauca (K, SING); SAN 19617: U.

borneensis (SAN, SING); SAN 19794: P. prismatica (K); SAN 19795: A. suaveolens (K,

Climbing Annonaceae of Borneo 471

L); SAN 20053: U. littoralis (K); SAN 20330: U. littoralis (K, SING); SAN 20451: Fr:

glauca (SAN); SAN 20882: U. littoralis (SAN); SAN 20926: A. suaveolens (K, L, SAN,

SING); SAN 21060: Fiss. brevistipitatum (K, SAN); SAN 22793: U. monticola (SAN,

SING); SAN 23074: A. suaveolens (K, L, SAN); SAN 24589: D. dumosus (K, SAN,

SING); SAN 24592: 4. suaveolens (L, SAN); SAN 24978: Fr. borneensis (K, SAN, SAR,

SING); SAN 25044: U. excelsa (K, SAN); SAN 25150: U. concava (K, SAN); SAN

25608: U. littoralis (SAN); SAN 26014: A. suaveolens (K, SAN); SAN 26325: Fiss.

multivenium (K, L, SAN); SAN 26676: A. costatus (SAN); SAN 26678: D. dumosus (K,

SAN, SING); SAN 26679: U. javana (SAN); SAN 26755: D. chinensis (SAN); SAN

27117: Fiss. fulgens (K, L, SAN, SAR, SING); SAN 27124: U. littoralis (K SAN); SAN

27266: U. littoralis (K, SAN); SAN 27452: U. grandiflora (K, SAN); SAN 27733: Fr.

glauca (K, SAN); SAN 27739: U. littoralis (K, SAN); SAN 27869: Fr. grandifolia (K

SAN); SAN 28158: A. suaveolens (K, L, SAN, SING): SAN 28161: A. suaveolens (SAN);

SAN 28339: D. acutus (K, SAN, SING); SAN 28381: U. littoralis (SAN); SAN 28526:

Fiss. kinabaluense (K, L, SAN); SAN 29372: U. littoralis (K, SAN, SING); SAN 29601:

Fr. borneensis (K, SAN); SAN 29711: U. concava (SAN, SING); SAN 30002: U.

grandiflora (K, SAN); SAN 30010: Fr borneensis (K, SAN, SAR); SAN 30015: U.

excelsa (K, SAN, SING); SAN 30151: A. suaveolens (SAN); SAN 30152: P. prismatica

(K, SAN); SAN 30295: U. littoralis (K, SAN); SAN 30327: P. prismatica (SAN); SAN

30561: Fiss. bygravei (K, L, SAN, SAR, SING); SAN 30855: U. excelsa (K, SAN); SAN

31082: U. monticola (SAN); SAN 31090: A. roseus (K, SAN); SAN 31107: A. suaveolens

(K, SAN); SAN 31320: Fiss. kingii (K, L, SAN, SAR); SAN 31322: Fr: formosa (SAN);

SAN 31663: D. dumosus (K, SAN); SAN 31746: U. littoralis (SAN); SAN 31988: P.

prismatica (K, SAN); SAN 32519: U. cuneifolia (K, SAN); SAN 32686: Fr. korthalsiana

(K, SAN, SAR, SING); SAN 32690: A. suaveolens (K, SAN, SING); SAN 32739: Fr

borneensis (K, SAN); SAN 32752: U. littoralis (K, SAN); SAN 32834: U. borneensis (K,

SAN); SAN 32837: P. prismatica (SAN); SAN 33045: S. affinis (SAN, SAR); SAN

33192: A. suaveolens (K, L, SAN, SING); SAN 33257: Fr borneensis (SAN); SAN

33456: U. littoralis (K, SAN, SING); SAN 33659: S. affinis (SAN); SAN 33826: U.

borneensis (K, SAN); SAN 33960: U. littoralis (SAN); SAN 33999: U. littoralis (K,

SAN, SING): SAN 34084: Fr. borneensis (K, SAN, SAR, SING); SAN 34135: Fiss.

manubriatum (K, SAN); SAN 34960: Fiss. kingii (K, SAN, SAR, SING); SAN 35396: Fr:

excisa (K, SAN); SAN 35416: A. suaveolens (SAN, SING); SAN 35616: Fr: biglandulosa

(K, SAN, SING); SAN 35802: Fiss. latifolium (Philippines form) (SAN); SAN 35869: U.

monticola (SAN); SAN 35871: U. schefferi (K, SAN, SING); SAN 35914: Fiss.

manubriatum (K, SAN, SAR, SING); SAN 35917: M. kentii (K, SAN, SING); SAN

36859: P. prismatica (K); SAN 36863: M. clementis (SAN, SING); SAN 37034: M.

clementis (K, SAN); SAN 37089: U. verrucosa (K, SAN, SING); SAN 37741: A.

suaveolens (K, SAN, SING); SAN 38102: D. chinensis (K, SAN); SAN 38390: D.

chinensis (K, SAN, SING); SAN 39641: P. prismatica (SAN); SAN 40263: A. costatus

(SAN); SAN 40335: Fiss. elmeri (K, L, SAN, SAR); SAN 40798: U. concava (K, SAN);

SAN 40802: Fiss. fulgens (SAN, SING); SAN 40837: U. concava (SAN, SING); SAN

40846: D. chinensis (K, SAN); SAN 41074: A. veldkampii? (SAN); SAN 41257: U.

littoralis (K, SAN); SAN 41357: U. littoralis (K, SAN, SING); SAN 41432: P. prismatica

(SAN); SAN 41655: U. micrantha (K, SAN); SAN 41799: U. schefferi (K, SAN); SAN

41801: M. clementis (K, SAN); SAN 42578: U. excelsa (SAN); SAN 42662: Fr. glauca

(K, SAN); SAN 43015: S. affinis (SAN); SAN 43192: U. littoralis (K, SAN); SAN 43443:

A. macropodus (SAN); SAN 44321: U. excelsa (K, SAN); SAN 45063: U. sp. (SAN);

472 Gard. Bull. Singapore 64(2) 2012

SAN 46087: D. dumosus (K, SAN); SAN 46339: Fr. korthalsiana (K, SAN); SAN 47261:

U. grandiflora (K, SAN); SAN 47422: U. lobbiana (K, L, SAN); SAN 47489: P

prismatica (SAN); SAN 47864: A. suaveolens (K, L, SAN); SAN 48163: U. excelsa (K,

SAN); SAN 48258: Fr, glauca (SAN); SAN 48259: Fr glauca (SAN); SAN 48430: A.

gracilis (K, SAN); SAN 48566: A. suaveolens (SAN); SAN 48591: A. suaveolens (K,

SAN); SAN 48980: 4. ochropetalus (SAN); SAN 49173: U. micrantha (K, SAN); SAN

49298: Fiss. kingii (K, L, SAN); SAN 49359: A. suaveolens (K, SAN); SAN 49364: S.

affinis (K SAN); SAN 49558: U. sp. (K, SAN); SAN 49589: M. kentii (K, SAN); SAN

49666: A. roseus (SAN); SAN 49673: A. suaveolens (SAN); SAN 49852: A. suaveolens

(SAN); SAN 49888: Fiss. fulgens (SAN); SAN 49941: Fr: glauca (SAN); SAN 49942: U.

littoralis (SAN); SAN 50360: P. prismatica (SAN); SAN 50416: A. gracilis (K, SAN);

SAN 50461: U. lobbiana (SAN); SAN 50492: U. excelsa? (SAN); SAN 50555: S. affinis

(SAN); SAN 50557: D. chinensis (K, SAN, SING); SAN 50560: S. affinis (SAN); SAN

50976: A. suaveolens (K, SAN); SAN 51509: U. monticola (SAN); SAN 51630: A.

suaveolens (SAN); SAN 51652: D. chinensis (K, SAN, SAR, SING); SAN 51849: U.

lobbiana (SAN); SAN 52758: Fiss. paniculatum (K, L, SAN, SAR); SAN 52848: M.

kentii (K, SAN); SAN 53056: U. littoralis (K, SAN); SAN 53063: U. littoralis (K, SAN);

SAN 53422: Fiss. kingii (K, SAN, SAR); SAN 53450: U. verrucosa (SAN); SAN 54333:

Fr. glauca (SAN); SAN 54502: Fiss. kingii (K, SAN); SAN 54507: A. ochropetalus

(SAN): SAN 54532: U. beccarii (K, SAN); SAN 54634: U. excelsa (SAN); SAN 54856:

A. lanuginosus (SAN); SAN 54917: U. borneensis (SAN); SAN 54958: U. littoralis

(SAN); SAN 55367: A. suaveolens (K, L, SAN); SAN 55727: D. acutus (SAN); SAN

55773: U. lobbiana (K, SAN, SAR, SING); SAN 56191: Fiss. elmeri (K, SAN); SAN

56382: Fiss. carrii (SAN); SAN 56772: U. curvistipitata (SAN); SAN 56821: Fiss.

manubriatum (K, SAN); SAN 56951: U. borneensis (K, SAN); SAN 56972: U. schefferi

(K, SAN); SAN 57200: U. littoralis (SAN); SAN 57256: U. schefferi (K, SAN); SAN

57259: Fiss. elmeri (K, L, SAN, SAR, SING); SAN 57343: Fr. korthalsiana (K, L, SAN,

SING); SAN 57374: Fiss. elmeri (K, L, SAN, SAR); SAN 57383: Fiss. manubriatum (K,

SAN); SAN 57391: U. littoralis (SAN, SING); SAN 58052: U. schefferi (K, SAN); SAN

59263: Fiss. latifolium (Philippines form) (SAN); SAN 59589: U. excelsa (SAN); SAN

59830: U. littoralis (SAN); SAN 60004: A. suaveolens (K, SAN, SING); SAN 60026: U.

concava (SAN); SAN 60039: U. concava (K, SAN): SAN 60242: Fiss. multivenium (K,

L, SAN); SAN 60274: Fr glauca (K, SAN); SAN 60280: U. concava (SAN); SAN 60404:

U. concava (SAN); SAN 60661: Fiss. carrii (K, SAN); SAN 60896: U. borneensis

(SAN); SAN 60940: U. excelsa (SAN); SAN 61894: D. chinensis (SAN); SAN 62880: A.

suaveolens (SAN); SAN 64442: D. chinensis (SAN); SAN 64464: D. chinensis (SAN);

SAN 64738: A. suaveolens (K, SAN); SAN 64780: U. cuneifolia (SAN, SING); SAN

65449: U. borneensis (K, SAN); SAN 65480: Fiss. kingii (K, L, SAN); SAN 65500: Fiss.

bygravei (K, L, SAN, SAR, SING); SAN 65925: U. littoralis (K, SAN); SAN 66329: A.

macropodus (SAN); SAN 66697: U. concava (SAN); SAN 66814: U. grandiflora (SAN);

SAN 67224: U. excelsa(K, SAN); SAN 67271: A. suaveolens (SAN, SING); SAN 67272:

U. concava (SAN); SAN 67313: U. littoralis (SAN); SAN 67318: A. suaveolens (K,

SAN); SAN 67368: U. schefferi (K, SAN); SAN 67436: U. concava (SAN); SAN 67456:

Fr. glauca (SAN); SAN 67468: A. ochropetalus (SAN); SAN 67555: U. concava (SAN);

SAN 67683: A. ochropetalus (SAN); SAN 67960: Fr. borneensis (SAN); SAN 68191: A.

hirtipes (SAN); SAN 68499: Fiss. multivenium (K, SAN, SING); SAN 68720: U.

monticola (K, SAN); SAN 68811: 4. macropodus (SAN); SAN 69118: U. borneensis

(SAN); SAN 69200: U. sp. (K, SAN); SAN 69461: Fiss. fulgens (SAN); SAN 69465: A.

Climbing Annonaceae of Borneo 473

ochropetalus (SAN); SAN 69532: U. littoralis (SAN); SAN 70051: U. excelsa (K, SAN):

SAN 70583: 4. ochropetalus (K, L, SAN); SAN 70609: A. suaveolens (K, SAN); SAN

70686: U. verrucosa (K, SAN); SAN 71177: A. sp. (SAN); SAN 71181: A. suaveolens (K,

SAN, SING): SAN 71272: A. macropodus (K, SAN); SAN 71283: Fiss. kingii (K, SAN,

SAR, SING); SAN 72409: Fr. glauca (K, SAN, SAR, SING); SAN 73349: Fiss. fulgens

(K, SAN, SING); SAN 73676: U. monticola (K, SAN); SAN 73821: Fiss. kingii (K, SAN,

SAR, SING); SAN 74364: A. roseus (L, SAN); SAN 74866: U. littoralis (K, SAN); SAN

74975: Fr. affinis (K, SAN, SING); SAN 75338: A. ochropetalus (K, L, SAN); SAN

75645: A. ochropetalus (SAN); SAN 75662: U. littoralis (K, SAN, SING); SAN 76069:

U. littoralis (K, SAN); SAN 76088: U. excelsa (K, SAN); SAN 76878: A. macropodus

(K, L, SAN, SING); SAN 76919: A. hirtipes (SAN); SAN 77134: U. grandiflora (SAN);

SAN 77206: U. grandiflora (K, SAN, SING); SAN 77652: A. suaveolens (K, SAN,

SING); SAN 77819: M. kentii (K, SAN, SING); SAN 77851: A. gracilis (K, SAN); SAN

77873: U. borneensis (K, SAN, SING); SAN 77946: D. chinensis (K, SAN, SAR, SING);

SAN 78027: P. prismatica (SAN); SAN 78073: M. clementis (K, SAN); SAN 78075: Fr.

excisa (K, SAN, SING); SAN 78486: U. littoralis (SAN); SAN 78517: A. gracilis (SAN);

SAN 79055: U. littoralis (SAN); SAN 79125: U. littoralis (SAN); SAN 79170: Fiss.

kinabaluense (SAN); SAN 79171: Fr glauca (K, SAN, SAR, SING); SAN 79183: Fr:

glauca (SAN, SAR, SING); SAN 79195: U. littoralis (SAN); SAN 79233: Fr: ?borneensis

(K, SAN, SAR); SAN 79615: A. ochropetalus (L, SAN, SING); SAN 79666: M. clementis

(SAN); SAN 79705: A. ochropetalus (SAN); SAN 79796: M. clementis (K, SAN); SAN

80014: P. prismatica (SAN); SAN 80089: D. acutus (SAN); SAN 80214: A. gracilis (K,

L, SAN); SAN 80237: U. concava (SAN); SAN 80259: Fiss. fulgens (SAN); SAN 80265:

A. hirtipes (SAN); SAN 80277: A. ochropetalus (SAN, SING); SAN 80316: P. prismatica

(SAN); SAN 80354: Fiss. fulgens (SAN); SAN 80355: A. suaveolens (SAN); SAN 80396:

A. ochropetalus (SAN); SAN 80632: P. prismatica (SAN); SAN 80781: S. affinis (K,

SAN); SAN 80810: U. excelsa (K, SAN); SAN 80980: Fiss. kingii (K, L, SAN); SAN

81362: Fiss. bygravei (K, L, SAN); SAN 81393: A. ochropetalus (K, L, SAN); SAN

81444: Fiss. bygravei (SAN); SAN 81456: Fiss. kingii (K, SAN); SAN 81504: U.

curvistipitata (K, SAN, SAR); SAN 81923: A. suaveolens (K, SAN, SING); SAN 81977:

U. beccarii (K, SAN); SAN 81997: U. cuneifolia (K, SAN); SAN 82035: Fiss. fulgens

(K, SAN, SING); SAN 82176: U. concava (SAN, SING); SAN 82231: A. veldkampii (K,

L, SAN); SAN 82277: U. micrantha (K, SAN); SAN 82415: Fiss. fulgens (K, SAN,

SAR); SAN 82456: Fiss. fulgens (SAN, SAR, SING); SAN 82574: A. ochropetalus (K,

L, SAN, SING); SAN 82635: Fr: grandifolia (SAN); SAN 82657: A. suaveolens (SAN,

SING); SAN 83513: A. suaveolens (K, L, SAN); SAN 83616: M. kentii (SAN); SAN

83764: U. lobbiana (K, SAN); SAN 84152: Fr. glauca (K, SAN, SAR); SAN 84428: U.

concava (SAN); SAN 84436: A. suaveolens (K, SAN); SAN 84477: U. concava (SAN,

SING); SAN 84583: A. suaveolens (SAN, SING); SAN 84597: U. littoralis (K, SAN);

SAN 84697: U. clementis (SAN, SAR, SING); SAN 84744: A. suaveolens (SAN, SING);

SAN 84795: Fr. affinis (K, L, SAN, SAR); SAN 84859: Fr: glauca (K, SAN, SAR, SING);

SAN 85055: A. suaveolens (K, SAN, SING); SAN 85119: U. monticola (SAN); SAN

85299: U.? javana (SAN); SAN 85325: U. littoralis (SAN); SAN 85365: Fiss. latifolium

(K, SAN); SAN 85995: A. suaveolens (SAN, SING); SAN 86075: Fiss. latifolium

(Kinabalu form) (L, SAN); SAN 86173: U. littoralis (K, SAN); SAN 86195: A. roseus (K,

SAN): SAN 86201: A. swaveolens (K, L, SAN); SAN 86244: Fr. biglandulosa (SAN);

SAN 86341: Fiss. fulgens (SAN); SAN 86404: Fiss. fulgens (SAN); SAN 86405: A.

suaveolens (SAN); SAN 86468: Fiss. fulgens (SAN); SAN 86619: Fiss. manubriatum (K,

474

Gard. Bull. Singapore 64(2) 2012

SAN, SING); SAN 86641: 4. suaveolens (SAN, SING); SAN 86645: Fr. borneensis (K,

SAN); SAN 86651: Fr grandifolia (K, SAN, SAR); SAN 86687: U. borneensis (K,

SAN); SAN 86795: U. littoralis (SAN); SAN 87341: U. concava (SAN); SAN 87377: Fr.

glauca (SAN); SAN 87379: U. concava (SAN); SAN 87761: 4. ochropetalus (SAN);

SAN 87770: M. clementis (SAN, SAR); SAN 87837: A. ochropetalus (K, L, SAN, SING);

SAN 88055: Fr: biglandulosa (SAN, SAR); SAN 88117: U. littoralis (SAN); SAN 88124:

Fr. glauca (A, SAN, SAR, SING); SAN 88135: U. verrucosa (SAN, SING); SAN 88138:

A. suaveolens (SAN); SAN 88176: Fiss. manubriatum (SAN); SAN 88263: Fr glauca

(SAN, SAR); SAN 88271: A. polygynus (SAN); SAN 88357: A. polygynus (SAN, SING);

SAN 88411: A. suaveolens (SAN); SAN 88432: Fr borneensis (K, SAN, SAR); SAN

88484: D. chinensis (SAN, SAR); SAN 88910: U. grandiflora (SAN); SAN 88965: Fr.

glauca (SAN, SAR, SING); SAN 89042: U. littoralis (SAN); SAN 89118: D. chinensis

(SAN, SAR, SING); SAN 89139: A. suaveolens (SAN); SAN 89141: M. clementis (SAN,

SAR); SAN 89176: Fr. borneensis (K, SAN, SAR); SAN 89317: A. ochropetalus (SAN);

SAN 89516: A. suaveolens (SAN, SING); SAN 89745: P. prismatica (SAN); SAN 89799:

U. borneensis (SAN); SAN 89818: Fr. borneensis (SAN); SAN 89867: Fr. glauca (SAN);

SAN 89920: A. ochropetalus (SAN, SING); SAN 90152: A. polygynus (L, SAN); SAN

90305: Fr glauca (SAN); SAN 90443: A. polygynus (SAN); SAN 90669: Fr. glauca

(SAN); SAN 91078: D. dumosus (SAN, SAR); SAN 91080: U. littoralis (SAN); SAN

91106: U. concava (K, SAN); SAN 91116: A. gracilis (SAN); SAN 91358: D. dumosus

(SAN, SAR); SAN 91359: Fr borneensis (K, SAN, SAR); SAN 91373: M. clementis

(SAN, SAR); SAN 91389: U. concava (SAN); SAN 91401: U. concava (SAN); SAN

91420: A. suaveolens (K, SAN); SAN 91422: A. ochropetalus (SAN); SAN 91433: A.

suaveolens (K, SAN, SING); SAN 91465: U. concava (SAN); SAN 91488: Fr glauca

(SAN, SAR, SING); SAN 91745: A. ochropetalus (SAN); SAN 91782: A. ochropetalus

(SAN); SAN 91801: U. javana (SAN); SAN 91823: S. affinis (SAN); SAN 91828: U.

concava (K, SAN); SAN 91913: D. dumosus (K, SAN, SAR, SING); SAN 91948: A.

lanuginosus (K, SAN); SAN 92017: U. excelsa (K SAN); SAN 92041: A. suaveolens

(SAN, SING); SAN 92475: A. ochropetalus (L, SAN); SAN 92595: A. suaveolens (SAN,

SING); SAN 92972: Fiss. fulgens (K, SAN, SAR, SING); SAN 92973: A. polygynus (L,

SAN, SING); SAN 93088: Fr: grandifolia (K, L, SING); SAN 93096: Fiss. brevistipitatum

(SAN, SAR, SING); SAN 93552: Fiss. cf. latifolium (K, SAN, SAR); SAN 93980: U.

excelsa (K); SAN 94050: A. hirtipes (K, SAN, SAR, SING); SAN 94457: U. concava

(SAN); SAN 94477: A. suaveolens (SAN); SAN 94664: Fiss. kingii (K, SAN, SAR,

SING); SAN 94721: U. borneensis (SAN); SAN 94762: S. affinis (SAN, SAR); SAN

94763: U. javana (K, SAN, SAR); SAN 94786: A. suaveolens (K, SAN); SAN 94791: U.

littoralis (SAN); SAN 94799: Fr glauca (SAN, SAR); SAN 94815: U. lobbiana (K,

SAN, SAR, SING); SAN 94826: A. suaveolens (K, SAN); SAN 94835: Fiss. manubriatum

(K, SAN); SAN 94876: A. suaveolens (K, SAN); SAN 94966: Fr: affinis (A, K, SAN,

SING); SAN 94983: Fiss. brevistipitatum (SAN, SAR); SAN 95021: U. littoralis (SAN);

SAN 95105: A. ochropetalus (SAN); SAN 95178: M. kentii (SAN); SAN 95310: U.

excelsa (K, SAN); SAN 95434: A. ochropetalus (K, L, SAN); SAN 95552: Fr glauca

(SAN, SAR); SAN 95633: A. polygynus (L, SAN); SAN 95637: A. ochropetalus (L, SAN,

SING); SAN 95689: A. ochropetalus (L, SAN); SAN 95725: A. suaveolens (K, SAN);

SAN 95743: D. dunalii (K, SAN); SAN 95962: A. ochropetalus (SAN); SAN 96010: A.

macropodus (K, SAN); SAN 96052: A. ochropetalus (L, SAN); SAN 96132: Fr: glauca

(A, K, SAN, SAR); SAN 96158: Fr. borneensis (SAN, SAR); SAN 96160: Fr glauca

(SAN); SAN 96169: A. ochropetalus (L, SAN); SAN 96173: U. excelsa (SAN); SAN

Climbing Annonaceae of Borneo 475

96177: A. macropodus (K, SAN); SAN 96289: U. grandiflora (A, K, SAN, SING); SAN

96353: A. suaveolens (K, SAN): SAN 96363: U. grandiflora (SAN); SAN 96368: M.

clementis (SAN); SAN 96389: Fiss. bygravei (SAN): SAN 96465: U. borneensis (K,

SAN); SAN 96492: 4. ochropetalus (SAN); SAN 96546: A. gracilis (K, SAN); SAN

96565: U. borneensis (SAN); SAN 96649: A. ochropetalus (K, L, SAN); SAN 96870:

Fiss. kingii (SAN); SAN 96982: A. ochropetalus (K, L, SAN); SAN 97014: M. clementis

(K, SAN, SAR); SAN 97017: U. grandiflora (K, SAN); SAN 97313: U. littoralis (SAN);

SAN 97658: U. excelsa (SAN); SAN 97684: Fr. glauca (K, SAN, SAR, SING); SAN

99225: U. littoralis (SAN); SAN 99417: U. excelsa (K, SAN, SAR); SAN 99424: A.

veldkampii (K, L, SAN); SAN 99452: U. concava (SAN); SAN 99639: U. borneensis

(SAN); SAN 99642: U. littoralis (A, K, SAN); SAN 99648: U. grandiflora (A, SAN);

SAN 99753: M. clementis (K SAN); SAN 99970: Fiss. fulgens (SAN); SAN 99995: U.

borneensis (SAN); SAN 100267: A. suaveolens (SAN, SING); SAN 100312: M. clementis

(SAN): SAN 100331: U. littoralis (K, SAN); SAN 100394: U. concava (A, K, SAN);

SAN 101266: U. littoralis (SAN); SAN 101323: Fiss. latifolium (SAN); SAN 101458: A.

ochropetalus (SAN); SAN 101474: Fr. biglandulosa (SAN, SING); SAN 101773: D.

chinensis (SAN); SAN 101797: U. concava (SAN); SAN 102214: U. concava (SAN):

SAN 102372: M. kentii (SAN); SAN 102663: A. suaveolens (K, SAN); SAN 102711: U.

cuneifolia (K, SAN); SAN 102743: U. littoralis (SAN); SAN 102777: A. suaveolens

(SAN); SAN 102792: P. prismatica (SAN); SAN 103013: A. suaveolens (SAN); SAN

103037: Fiss. fulgens (SAN); SAN 103087: P. prismatica (SAN); SAN 103098: P.

prismatica (SAN); SAN 103232: P. prismatica (SAN); SAN 103273: A. suaveolens

(SAN); SAN 103285: U. littoralis (SAN); SAN 103290: P. prismatica (SAN); SAN

103565: U. monticola (K, SAN); SAN 103632: A. suaveolens (K, SAN); SAN 103825: U.

littoralis (K); SAN 104062: U. concava (SAN); SAN 104063: A. ochropetalus (L, SAN,

SING); SAN 104313: Fiss. bygravei (SAN); SAN 104316: Fr. biglandulosa (SAN); SAN

104321: Fiss. manubriatum (SAN); SAN 105255: Fiss. manubriatum (SAN); SAN

105281: Fr: biglandulosa (K, SAN); SAN 105313: Fr. biglandulosa (SAN); SAN 105350:

Fiss. manubriatum (SAN); SAN 105610: A. suaveolens (K, L, SAN); SAN 105648: U.

littoralis (SAN); SAN 105998: U. clementis (K, SAN); SAN 106121: A. suaveolens (K,

SAN); SAN 106704: U. littoralis (SAN); SAN 106738: A. suaveolens (SAN); SAN

106808: U. littoralis (SAN); SAN 106864: A. polygynus (SAN); SAN 106934: U.

clementis (SAN); SAN 106984: Fr. glauca (SAN); SAN 107131: U. littoralis (K, SAN):

SAN 107163: U. littoralis (SAN); SAN 107220: Fr. borneensis (SAN); SAN 107247: Fr

glauca (K, SAN, SAR, SING); SAN 107334: Fr formosa (SAN); SAN 107610: A.

suaveolens (K, SAN); SAN 107617: M. clementis (K, SAN); SAN 107714: Fr. borneensis

(SAN); SAN 107737: U. littoralis (SAN); SAN 108087: U. littoralis (SAN); SAN

108114: A. macropodus (SAN); SAN 108162: U. littoralis (SAN); SAN 108284: A.

ochropetalus (K, L, SAN); SAN 108469: Fiss. fulgens (K, SAN, SAR); SAN 108555: U.

grandiflora (SAN); SAN 108595: U. littoralis (SAN); SAN 108635: Fr. excisa (SAN);

SAN 108643: A. ochropetalus (K, L, SAN); SAN 108711: Fr glauca (SAN); SAN

109284: U. littoralis (SAN); SAN 109353: A. suaveolens (SAN); SAN 109422: A.

suaveolens (SAN); SAN 109597: Fiss. kingii (SAN); SAN 109643: U. excelsa (K, SAN);

SAN 109908: 4. suaveolens (SAN); SAN 109937: Fiss. brevistipitatum (SAN); SAN

109980: D. dumosus (K, SAN); SAN 1099835: Fiss. multivenium (SAN); SAN 110000: U.

littoralis (K, SAN); SAN 110093: U. argentea (K, SAN); SAN 110109: Fr: biglandulosa

(SAN); SAN 110136: Fiss. kingii (SAN); SAN 110143: A. ochropetalus (SAN); SAN

110145: D. dumosus (SAN); SAN 110271: U. sp. (SAN); SAN 110272: A. costatus (K, L,

476

Gard. Bull. Singapore 64(2) 2012

SAN); SAN 110334: Fr glauca (K, SAN); SAN 110391: Fr. affinis (SAN); SAN 110454:

U. clementis (SAN); SAN 110856: Fr glauca (K, SAN); SAN 110904: Fiss. kingii (K,

SAN, SAR, SING); SAN 111013: 4. ochropetalus (SAN); SAN 111028: A. suaveolens

(K, SAN); SAN 111037: D. acutus (K, SAN); SAN 111109: A. ochropetalus (SAN); SAN

111112: U. littoralis (SAN); SAN 111248: A. suaveolens (SAN); SAN 111420: P

prismatica (SAN); SAN 111446: Fiss. fulgens (SAN); SAN 111508: Fr: affinis (K); SAN

111537: U. littoralis (SAN); SAN 111657: U. verrucosa (SAN); SAN 112135: U.

borneensis (SAN); SAN 112195: U. littoralis (SAN); SAN 112310: U. grandiflora

(SAN); SAN 112938: A. ochropetalus (K, L, SAN); SAN 113087: A. suaveolens (SAN);

SAN 113183: A. suaveolens (SAN); SAN 113188: Fr glauca (SAN); SAN 113233: U.

verrucosa (SAN); SAN 113265: A. hirtipes (SAN); SAN 113300: U. schefferi (K, SAN);

SAN 113314: M. clementis (SAN): SAN 113330: A. suaveolens (K, SAN); SAN 113343:

U. cuneifolia (SAN); SAN 113482: U. littoralis (SAN); SAN 113576: A. costatus (SAN);

SAN 113741: Fiss. kinabaluense (SAN); SAN 113868: Fr: affinis (SAN); SAN 113934:

U. monticola (K, SAN); SAN 113950: Fiss. kingii (SAN); SAN 113988: U. excelsa (K,

SAN): SAN 113994: A. suaveolens (K, SAN); SAN 114010: U. borneensis (SAN); SAN

114024: M. clementis (K, SAN); SAN 114030: Fr glauca (K, SAN); SAN 114042: Fiss.

bygravei (SAN); SAN 114050: Fiss. bygravei (SAN); SAN 114057: A. suaveolens (SAN);

SAN 114066: A. roseus (SAN); SAN 114275: U. excelsa (SAN); SAN 114305: U.

littoralis (SAN); SAN 114313: U. grandiflora (SAN); SAN 114875: Fiss. fulgens (SAN);

SAN 114905: A. suaveolens (L, SAN); SAN 115294: S. affinis (K, SAN); SAN 115307:

A. gracilis (SAN); SAN 115392: P. prismatica (SAN); SAN 115410: P. prismatica (SAN);

SAN 115418: U. littoralis (SAN); SAN 115576: A. suaveolens (SAN); SAN 115616: P

prismatica (SAN); SAN 115672: A. suaveolens (SAN); SAN 115823: Fiss. bygravei

(SAN); SAN 115835: M. clementis (K, SAN); SAN 115877: Fiss. brevistipitatum (K,

SAN): SAN 116060: A. suaveolens (K, SAN); SAN 116294: A. suaveolens (SAN); SAN

116333: A. suaveolens (K, SAN); SAN 116369: A. suaveolens (SAN); SAN 116414: A.

suaveolens (SAN); SAN 116433: U. clementis (SAN); SAN 116446: U. littoralis (SAN);

SAN 116450: A. suaveolens (SAN); SAN 116618: A. costatus (K, SAN); SAN 116646: A.

gracilis (SAN); SAN 116702: Fiss. brevistipitatum (K, SAN); SAN 116719: A. gracilis

(K, SAN); SAN 116737: Fr biglandulosa (K, SAN); SAN 116866: A. ochropetalus

(SAN); SAN 116946: Fr. biglandulosa (K, SAN); SAN 116964: U. littoralis (K, SAN);

SAN 117020: U. littoralis (SAN); SAN 117033: Fiss. kingii (SAN); SAN 117035: A.

suaveolens (K, SAN); SAN 117128: A. suaveolens (K, SAN); SAN 117158: M. clementis

(K, SAN, SAR); SAN 117355: A. suaveolens (K, SAN); SAN 117534: U. littoralis (K,

SAN); SAN 117542: A. gracilis (K, SAN); SAN 117675: A. macropodus (SAN); SAN

117822: U. grandiflora (K, SAN); SAN 117834: A. macropodus (SAN); SAN 117901: A.

suaveolens (SAN); SAN 117910: A. suaveolens (K, SAN); SAN 118225: A. suaveolens

(K, SAN); SAN 118257: Fiss. kingii (K, SAN); SAN 118278: A. suaveolens (K, SAN);

SAN 118305: Fr. biglandulosa (SAN); SAN 118308: A. suaveolens (K, SAN); SAN

118411: D. dumosus (K, SAN); SAN 118466: A. swaveolens (SAN); SAN 118571: Fiss.

brevistipitatum (A, K, L, SAN); SAN 118574: Fr glauca (SAN); SAN 118612: Fr: glauca

(K, SAN): SAN 118670: U. excelsa (K, SAN); SAN 118730: A. suaveolens (K, SAN);

SAN 118815: Fr glauca (SAN); SAN 118829: U. lobbiana (SAN); SAN 118908: U.

grandiflora (K, SAN); SAN 118944: U. littoralis (K, SAN); SAN 118972: U. concava (K,

SAN); SAN 119020: Fiss. fulgens (K, SAN); SAN 119024: A. polygynus (SAN); SAN

119025: A. suaveolens (K, SAN); SAN 119371: Fr glauca (K, SAN); SAN 119374: A.

polygynus (SAN); SAN 119398: U. sp. (K, SAN); SAN 119423: U. excelsa (K, SAN);

Climbing Annonaceae of Borneo 477

SAN 119442: Fr. glauca (SAN); SAN 119493: U. borneensis (K, SAN); SAN 119497: A.

suaveolens (SAN); SAN 119514: Fr biglandulosa (SAN); SAN 119557: Fiss.

brevistipitatum (SAN); SAN 119613: A. suaveolens (SAN); SAN 119715: A. suaveolens

(K, SAN); SAN 119735: Fiss. brevistipitatum (SAN); SAN 119825: A. suaveolens (SAN);

SAN 119936: A. gracilis (SAN); SAN 119976: Fr. glauca (K, SAN); SAN 120041: A.

gracilis (K, SAN); SAN 120069: Fr: glauca (K, SAN); SAN 120344: M. clementis (K,

SAN); SAN 120385: A. roseus (K, L); SAN 120444: U. grandiflora (SAN); SAN 120667:

D. chinensis (K, SAN); SAN 120940: A. suaveolens (K, SAN); SAN 120990: A.

ochropetalus (SAN); SAN 121129: Fr glauca (SAN); SAN 121194: A. suaveolens (K,

SAN); SAN 121491: A. suaveolens (SAN); SAN 121609: A. suaveolens (K, SAN); SAN

121643: A. suaveolens (K, SAN); SAN 121781: M. clementis (K, SAN); SAN 121900: A.

suaveolens (K, SAN); SAN 122034: A. ochropetalus (K, L, SAN); SAN 122048: U.

littoralis (SAN); SAN 122066: U. excelsa (SAN); SAN 122084: Fr. glauca (K, SAN);

SAN 122215: U. excelsa(K, SAN); SAN 122217: A. suaveolens (K, SAN); SAN 122358:

D. acutus (SAN); SAN 122390: M. clementis (SAN); SAN 122417: Fr. biglandulosa (K,

SAN); SAN 122483: U. littoralis (SAN); SAN 122647: A. suaveolens (SAN); SAN

122702: U. grandiflora (SAN); SAN 122722: Fiss. fulgens (SAN); SAN 122751: A.

suaveolens (K, SAN); SAN 122760: U. littoralis (K, SAN); SAN 122884: A. suaveolens

(K, SAN); SAN 123129: 4. suaveolens (K, SAN): SAN 123242: A. suaveolens (K, SAN);

SAN 123244: Fr. biglandulosa (K, SAN); SAN 123368: A. suaveolens (K, SAN); SAN

123449: Fr biglandulosa (SAN); SAN 123452: Fiss. brevistipitatum (SAN); SAN

123491: U. grandiflora (K, SAN); SAN 123595: A. suaveolens (K, SAN); SAN 124067:

Fr. glauca (SAN); SAN 124107: A. lanuginosa (SAN); SAN 124313: U. concava (SAN);

SAN 124315: A. ochropetalus (SAN); SAN 124435: A. suaveolens (SAN); SAN 124588:

Fr. korthalsiana (K, SAN); SAN 124640: U. excelsa (SAN); SAN 124681: Fr, borneensis

(SAN); SAN 125345: M. clementis (SAN); SAN 125435: Fiss. bygravei (SAN); SAN

125626: A. suaveolens (SAN): SAN 125629: Fiss. multivenium (K, SAN); SAN 125670:

Fiss. kingii (SAN); SAN 125675: U. littoralis (K, SAN); SAN 125676: A. suaveolens (K,

SAN); SAN 125693: Fr glauca (K, SAN); SAN 125700: Fiss. bygravei (K, SAN); SAN

125782: Fr. grandifolia (SAN); SAN 125787: A. gracilis (SAN); SAN 126103: U.

littoralis (SAN); SAN 126174: P. prismatica (SAN); SAN 126305: S. affinis (SAN,

SAR); SAN 126386: A. suaveolens (K, SAN); SAN 126420: Fr glauca (SAN); SAN

126487: A. suaveolens (K, SAN); SAN 126546: A. swaveolens (K, SAN); SAN 126716:

Fr. borneensis (SAN); SAN 126731: Fiss. fulgens (SAN); SAN 126796: Fiss. fulgens

(K); SAN 126847: S. affinis (SAN); SAN 126913: S. affinis (SAN); SAN 126915: U.

grandifolia (K, SAN); SAN 127022: A. suaveolens (SAN); SAN 127122: U. littoralis

(SAN); SAN 127364: P. prismatica (SAN); SAN 127597: A. suaveolens (SAN); SAN

127952: A. suaveolens (SAN); SAN 128075: A. gracilis (K, SAN); SAN 128125: A.

gracilis (K, SAN); SAN 128155: Fr. glauca (SAN); SAN 128245: M. clementis (SAN);

SAN 128311: Fr. biglandulosa (K, SAN); SAN 128460: Fr. biglandulosa (SAN); SAN

128859: A. polygynus (SAN); SAN 129337: A. suaveolens (K, SAN); SAN 129344: A.

costatus (K, L, SAN); SAN 129351: Fiss. carrii (L, SAN); SAN 129431: A. suaveolens

(SAN); SAN 129510: D. acutus (K, SAN); SAN 129559: A. ochropetalus (K, SAN);

SAN 129595: U. littoralis (SAN); SAN 129933: A. suaveolens (SAN); SAN 130000: A.

costatus (L, SAN); SAN 130103: U. cuneifolia (SAN); SAN 130134: M. clementis (K,

SAN); SAN 130144: Fiss. kingii (SAN); SAN 130289: U. grandiflora (SAN); SAN

130604: U. verrucosa (SAN); SAN 130859: U. excelsa (SAN); SAN 130884: Fiss.

manubriatum (K, SAN); SAN 131106: U. excelsa (SAN); SAN 131493: Fr. biglandulosa

478

Gard. Bull. Singapore 64(2) 2012

(L, SAN, SAR); SAN 131515: Fr biglandulosa (K, SAN); SAN 131931: Fiss.

brevistipitatum (SAN); SAN 132017: U. littoralis (SAN); SAN 132255: Fr glauca

(SAN); SAN 132360: U. grandiflora (SAN); SAN 132626: A. suaveolens (SAN); SAN

132687: Fr. biglandulosa (SAN, SAR); SAN 132860: Fiss. kingii (SAN); SAN 132936:

Fr. glauca (K, SAN, SAR); SAN 132949: A. ochropetalus (K, L, SAN); SAN 133079: U.

excelsa (SAN); SAN 133165: U. littoralis (SAN); SAN 133299: Fiss. brevistipitatum

(SAN, SING); SAN 133301: A. macropodus (SAN); SAN 133340: Fiss. kingii (A, K,

SAN); SAN 133407: A. suaveolens (SAN); SAN 133454: A. polygynus (L, SAN); SAN

133461: U. littoralis (SAN); SAN 133971: Fr. excisa (K, SAN, SAR); SAN 134272: A.

polygynus (L, SAN); SAN 134410: Fr biglandulosa (A, SAN, SAR); SAN 134484: U.

monticola (SAN); SAN 134563: D. chinensis (K, SAN, SAR); SAN 134815: U. monticola

(SAN); SAN 135137: A. costatus (SAN); SAN 135255: U. excelsa (SAN); SAN 135723:

U. borneensis (SAN); SAN 135822: Fr glauca (SAN); SAN 135848: M. clementis

(SAN); SAN 135917: M. clementis (SAN); SAN 135928: A. ochropetalus (L, SAN);

SAN 135966: D. dumosus (SAN, SAR); SAN 135978: Fiss. bygravei (SAN); SAN

136077: Fiss. bygravei (L, SAN); SAN 136574: U. grandiflora (SAN); SAN 136711: A.

suaveolens (L, SAN); SAN 136733: Fr biglandulosa (K, L, SAN); SAN 136795: A.

suaveolens (SAN); SAN 136853: A. veldkampii (SAN); SAN 136950: U. excelsa (K,

SAN); SAN 136998: Fiss. kingii (K, SAN); SAN 137121: M. clementis (SAN); SAN

138346: U. littoralis (SAN); SAN 138410: A. suaveolens (SAN); SAN 139193: A.

suaveolens (SAN); SAN 139291: Fr. glauca (SAN); SAN 139350: A. suaveolens (SAN);

SAN 139526: A. polygynus (SAN); SAN 139559: A. suaveolens (SAN); SAN 139610: A.

suaveolens (SAN); SAN 141048: A. ochropetalus (SAN); SAN 141487: A. suaveolens

(SAN); SAN 141639: U. borneensis (SAN); SAN 141985: Fiss. multivenium (SAN);

SAN 142979: A. hirtipes (K, L, SAN); SAN 143182: U. monticola (SAN); SAN 143349:

Fiss. kingii (K, L, SAN, SING); SAN 143368: U. javana (SAN); SAN 143374: A.

ochropetalus (SAN); SAN 143377: D. chinensis (K, SAN); SAN 143390: U. littoralis

(SAN): SAN 143460: Fiss. montanum (K, L,SAN, SING); SAN 143523: A. costatus (K,

SAN); SAN 143950: U. monticola (SAN); SAN 144055: U. littoralis (SAN); SAN

144147: U. littoralis (SAN); SAN 144148: A. suaveolens (SAN, SING); SAN 144330: U.

littoralis (K, SAN); SAN 144516: 4. ochropetalus (SAN); SAN 144753: U. concava

(SAN); SAN 144767: U. borneensis (SAN); SAN 146089: A. gracilis (SAN); SAN

146096: M. clementis (SAN); SAN 146945: A. suaveolens (SAN); SAN 147972: A.

?veldkampii (SAN, SAR); SAN 148887: A. suaveolens (SAN); SAN 168499: Fiss.

multivenium (SAN). SBC series; SBC 252: U. lobbiana (SAR); SBC 606: U. lobbiana

(SAR): SBC 1479: U. lobbiana (SAR). SFN series; SFN 10303: Fr: affinis (K, SING);

SFN 10487: U. lanuginosa (K); SFN 10524: M. kentii (K); SFN 10532: Fiss. fulgens (K,

SAR); SFN 25128: Fiss. latifolium (K); SFN 25286: Fr. biglandulosa (K, SING); SFN

27006: Fiss. carrii (SING); SFN 27100: Fiss. montanum (SING); SFN 38982: Fiss.

fulgens (K, SING). Sibil, J.; 136: U. littoralis; 306: Fiss. kingii (K, SAN). Sidiyasa, K.;

490: U. curvistipitata(L); 600: M. kentii (L); 668: A. lanuginosus (L); 775: Fr. biglandulosa

(L); 1188: U. javana (K, SAN); 1270: A. macropodus (K, SAN); 1285: U. monticola (K);

1440: Fiss. manubriatum (K); 1602: Fiss. kingii (K, L); PBU 333: A. gracilis (L); PBU

342: D. dumosus (K); PBU 344: M. kentii (K); PBU 410: A. gracilis (L); PBU 422: U.

excelsa (K); PBU 673: A. macropodus (K, L). Sidiyasa, K. & Arifin, Z.; 1602: Fiss.

kingii (BO); 2003: M. clementis (BO). Sidiyasa, K. & Kochummen, K.M.; 565: U.

excelsa (BO). Sidiyasa, K. et al.; Berau 1218: A. suaveolens (L); Berau 1235: A.

suaveolens (L); Berau 1254: A. macropodus (L); Berau 1257: A. macropodus (K, L).

Climbing Annonaceae of Borneo 479

Sidkan, A.; 1101: U. littoralis (SAN). Simpson, D.A.; 2144: U. littoralis (K, SAN):

2617: U. littoralis (SAN). Sinelair, J.; 10490: 4. polygynus (L). Soetisna, U.; 56: M.

clementis (K). Stevens, P.F. et al.; 6: D. dumosus (SAN); 157: Fiss. rugosum (A, SAR,

SAN); 365: Fiss. fulgens; 385: Fiss. fulgens (A, K, L, SAN); 400: A. macropodus (A, L,

SAN); 410: A. macropodus (A, L, SAN); 419: A. hirtipes (A, L, SAN): 687: U. littoralis

(A, SAN). Subok; 1064: U. /ittoralis (A). Sugau, J.; 288: A. suaveolens (K, SAN); 320:

U. littoralis (SAN).

Teo, L.E. & Pachiappan, G.; T & P 1088(KL 3588): Fr: korthalsiana (K); T & P 1098 (KL

3598): U. monticola(K): T & P 1100 (KL 3600): U. monticola (K); T & P 1102 (KL 3602):

Fiss. kingii (K); T & P 1116 (KL 3616): U. lobbiana (K). Tadong, D.; 13: Fr. grandifolia

(K); 150: D. chinensis (K); 255: U. concava (K); 620: U. excelsa (K). Tadong, L.; 210:

U. littoralis; 475: U. littoralis (K); 541: U. littoralis (K). Takashi, S.; 1405: U. littoralis

(SAN). Tandom; 2955: Fiss. fulgens (K); 3317: Fiss. fulgens (K); 4212: D. chinensis (K,

SING); 4791: A. suaveolens (K). Teijsmann, J.E.; 10838: Fiss. rugosum (L). Thomas,

S.A.; 93: A. suaveolens (SAN); 232: A. suaveolens (SAN). Tukirin; 392: U. schefferi (K).

USEP: 56: M. clementis (K).

Valera, J.; 3819: S. affinis (K); 3823: D. chinensis (K). van Balgooy, M.M.J.; 5837: A. gracilis

(L); 6083: Fr. borneensis (K, L); 6091: Fiss. manubriatum (L). van Balgooy, M.M.J. &

Kessler, P.J.A.; 5923: M. kentii (K). van Balgooy, M.M.J. & van Setten, A.K.; 5499: Fr:

affinis (BO, K, L); 5553: Fr. excisa (L); 5559: Fr. excisa (BO, L). van Niel, J.P.; 3906: A.

gracilis (L); 3913: Fiss. fulgens (L); 4069: Fiss. fulgens (L); 4072: Fiss. fulgens (L); 4259:

M. kentii; 4278: Fiss. fulgens (L); 4361: Fr. biglandulosa (L); 4465: A. roseus (L). van

Valkenburg, J.L.C.H.; 1295: P. prismatica (K). van Welzen, P.C.; 866: 4. ochropetalus

(L). Veldkamp, J.F.; 8567: A. veldkampii (L); 8577: A. gracilis (L); 8574: Fiss. bygravei

(L, SAR).

Winkler, H.; 2641: Fiss. manubriatum (K); 3000: D. dumosus (K). Wiriadinata, H.; 256:

clementis (A, K); 3540: Fr. glauca (A); 3547: A. ochropetalus (L); 3550: U. schefferi (A,

K); 3571: A. veldkampii (A, L). Wong, K.M.; WKM 109: Fiss. fulgens (K, SAN, SAR);

WKM 308: U. excelsa (K); WKM 570: U. beccarii (K); WKM 1554: A. suaveolens (L,

SAN); WKM 1851: Fiss. montanum (K); WKM 2233: A. suaveolens (K, SAN, SING);

WKM 2319: U. borneensis (K, SAN, SING); WKM 2680: A. ochropetalus (K, L, SAN);

sn: U. littoralis (SAN). Wood, D.D.; 444: Fr. ?borneensis (K); 1231: Fr. biglandulosa

(A); 1303: A. ochropetalus (A); 2202: Fiss. fulgens (K). Wood, G.H.S.; A2920: A.

ochropetalus (A, L, SING); A3686: Fr: korthalsiana? (K, SING); A4014: U. grandiflora

(K, SING); A4040: A. suaveolens (K, SING); A4291: U. borneensis (A, SING); A4380:

A. veldkampii (A, L, SING). Wood, G.H.S. & Wyatt-Smith, J.; A4226: 4. ochropetalus

(L, SING).

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Gardens’ Bulletin Singapore 64(2): 481-495. 2012 48]

Studies in Malesian Gentianaceae II:

A taxonomic framework for the Fagraea complex,

including the new genus Limahlania

K.M. Wong! and M. Sugumaran?

‘Singapore Botanic Gardens,

1 Cluny Road, Singapore 259569

wkm2000@gmail.com

*Rimba IImu Botanic Garden,

Institute of Biological Sciences, University of Malaya,

50603 Kuala Lumpur, Malaysia

maran@um.edu.my

ABSTRACT. A molecular phylogenetic study that provided good resolution of the Fagraea s.1.

complex is the basis for constructing a new taxonomic framework in this group. The lineages

identified showed good correspondence with other clades that represent well-established,

recently revised genera in the tribe (Potalieae) and subtribe (Potaliinae) in terms of structure and

statistical support (monophyly), and possessed recognisable morphological characteristics that

were potentially synapomorphic for each monophyletic group. Generic identities are therefore

adopted for the clades within this complex, as well as a somewhat isolated lineage, resulting in

the definition of Fagraea Thunb. s.s.; the reapplication of Cyrtophyllum Reinw., Picrophloeus

Blume, and Utania G.Don; and the circumscription of the new genus Limahlania K.M.Wong &

Sugumaran. A key to the genera of the Fagraea complex is presented and nomenclatural notes

are provided for each genus, in order to facilitate subsequent revisions. Limahlania crenulata

(Maingay ex Clarke) K.M.Wong & M.Sugumaran and U. volubilis (Wall.) M.Sugumaran are

new combinations. Lectotypes are selected for C. peregrinum Reinw., F. ceilanica Thunb.,

Kuhlia morindifolia Blume, and P. javanensis Blume.

Keywords. Cyrtophyllum, Fagraea, Gentianaceae, Limahlania, Malesia, Picrophloeus,

phylogenetic classification, Potalieae, Potaliinae, Racemosae, Utania

From clades to genera: taxonomic concepts in the Fagraea complex

A new classification for the Fagraea Thunb. complex is proposed, based on the results

of molecular phylogenetic analyses reported by Sugumaran & Wong (2012). In that

study, five distinct lineages were clearly identified within Fagraea s.1. that were shown

to correspond to other well established genera in the same tribe (Potalieae) (as treated

by Struwe & Albert 1997, 2004; Struwe et al. 1994; Struwe et al. 2002), in terms of

their monophyly in the molecular phylogenetic analyses as well as circumscription by

potentially synapomorphic morphological characters.

The Fagraea clade in that study, which includes the type species for the

genus, F. ceilanica Thunb., is the basis of the ‘true’ Fagraea (Fagraea s.s.). Fagraea

482 Gard. Bull. Singapore 64(2) 2012

crenulata Maingay ex Clarke is considered representative of a distinct lineage because

in the molecular phylogenetic study, it was only found to associate basally with the

Fagraea clade by chloroplast gene characteristics, but not in the ITS analysis. It is also

morphologically quite well distinguished from Fagraea s.s. No special generic name

has been associated with this taxon and it is the basis of the new genus Limahlania

K.M.Wong & M.Sugumaran.

The other clades correspond to genera resurrected from the synonymy

of Fagraea s.|., and now reapplied to phylogenetically circumscribed entities in

the complex. An appropriate name for the Racemosa clade of Sugumaran & Wong

(2012) appears to be Utania G.Don, which was given to replace the illegitimate name

Kuhlia Reinw. Don (1838) recognised one species, U. morindifolia which was later

recombined as Fagraea morindifolia by Blume (1838). However F: morindifolia is

a synonym of F) volubilis Wall. (Wong & Sugau 1996), which Sugumaran & Wong

(2012: Fig. 5) demonstrated as belonging to their Racemosa clade. This group is

consistently monophyletic in all the analyses with different gene regions.

Another interesting result of the study by Sugumaran & Wong (2012) is the

paraphyly of Fagraea section Cyrtophyllum (Reinw.) Blume as defined by Leenhouts

(1962a), which was consistently resolved as two distinct clades. Wong & Sugau

(1996) recognised two complexes within this “section” which they differentiated by

the exclusively axillary or terminal flowering habit. Indeed, these two complexes are

equivalent to the distinct Gigantea and Elliptica clades, respectively, in the molecular

analyses carried out. In the past, Ridley (1918) had regarded this “section” as a separate

genus, i.e., Cyrtophyllum Reinw., as the species are so different, especially in their

habit, in comparison to other epiphytic members of Fagraea s.l. For Cyrtophyllum,

Blume (1826) had included two species, C. peregrinum Reinw. and C. speciosum

Blume. Subsequent classification reduced C. peregrinum to a synonym of F: fragrans

Roxb., and C. speciosum to synonymy under F: elliptica Roxb. (= Picrophloeus Blume

now). The molecular analyses of Sugumaran & Wong (2012) show that F fragrans

falls within or is closely associated with the Gigantea clade, so that the generic name

Cyrtophyllum is most applicable to the clade.

In the sectional classification of Fagraea s.l. (Leenhouts 1962a), both

Picrophloeus and Cyrtophyllum were reduced to synonymy under Fagraea section

Cyrtophyllum. Picrophloeus was established with only one described species, P

javanensis Blume that was considered synonymous with F: e//iptica. Thus, the generic

name Picrophloeus is applicable to the Elliptica clade.

Characters for clades: circumscription of genera

Sugumaran & Wong (2012) have surveyed the expression of a number of characters in

the Fagraea complex, in comparison to other members of the Potaliinae (Anthocleista

and Potalia). Fagraea s.s. (i.e., the Fagraea clade, and excluding F) crenulata) (Fig.

1) is distinguished by fruits that produce copious creamy pale yellowish latex in the

epidermis and fruit wall. The rest of Fagraea s.|. have fruits that have no latex at all

Taxonomic framework for the Fagraea complex & new genus Limahlania 483

Fig. 1. Fagraea auriculata. A. Tree with orthotropic complexes of branches. B. Each branch

segment is potentially terminated by a cyme, here with fruits formed. C. A fruit longitudinally

cut, revealing how easily the fruit epidermis detaches as a thin peel from the fruit wall, and

latex oozing from the cut surface. D. Close-up of creamy pale yellowish latex oozing from

wound following removal of a fruit calyx lobe. (Photos: K.M. Wong)

484 Gard. Bull. Singapore 64(2) 2012

(Utania or the Racemosa clade, equivalent to Fagraea section Racemosae Benth., and

also Anthocleista and Potalia (Struwe, pers. comm.)), or produce just a translucent

gummy sap from the epidermis or wall when bruised (all others in Fagraea s.1.). Like

Anthocleista and Potalia, Fagraea s.s. has ellipsoid-rounded seeds, contrasting with

polygonal seed shapes found in Cyrtophyllum, Limahlania, Picrophloeus and Utania.

Fagraea s.s. includes not only trees, but also species that are epiphytic, hemi-epiphytic

and scramblers (life forms that are otherwise not found in the other groups of Fagraea

s.1. and which are rare in the Potaliinae generally).

Limahlania (our new genus represented by Fagraea crenulata) has a number

of unique characters in the subtribe, such as the development of prickles on the stem or

trunk and branches (Fig. 2), and serrulate-crenulate leaf margins. It is also distinguished

from all other taxa in the Fagraea complex, and also likely in the subtribe, by having

Fagerlind’s growth architectural model (Hallé et al. 1978; Sugumaran & Wong 2012).

This architecture (Fig. 2) is characterised by episodic stem or trunk growth, non-

equivalent vegetative axes (1.e., clear differentiation into trunk and branches), and a

modular branching system that develops plagiotropically by substitution. In Malesia,

the other lineages or clades in the Fagraea complex characterised by episodic growth

and non-equivalent vegetative axes either develop according to Scarrone’s model with

orthotropic complexes of branches (Fagraea s.s.; the Elliptica clade or Picrophloeus)

(Fig. 1 and 5), or Aubréville’s model with branching that is plagiotropic by apposition

(Fig. 4) (Hallé et al. 1978; Sugumaran & Wong 2012). These taxa all have a monopodial

stem or trunk. Other (non-Malesian) taxa in the subtribe are so far documented with

modular or sympodial stem or trunk systems (Hallé et al. 1978); they include the

African Anthocleista (most species have Leeuwenberg’s model) and the American

Potalia (one species specially analysed corresponds to Chamberlain’s model which

produces a linear sympodium giving a monocaulous habit; others, including a suite

of poorly known species are generally observed to be monocaulous: Struwe & Albert

2004). The apparent exception is a species of Anthocleista documented with Scarrone’s

model (Hallé et al. 1978). Also, Prevost (1978) identifies Koriba’s model for the New

Caledonian F) schlechteri.

On the other hand, Limahlania 1s related to Fagraea s.s. in having petiolar

sheaths that do not or only slightly fuse at the edges, and a peltate stigma structure.

It also resembles Fagraea s.s., Cyrtophyllum and Picrophloeus in having a fruit

epidermis that detaches from the fruit wall easily as a peel, unlike in Urania, where the

epidermis does not easily detach.

Utania (represented in the phylogenetic analyses of Sugumaran and Wong

(2012) by the Racemosa clade) has the most potentially synapomorphic morphological

characters among the Fagraea complex or the Potaliinae. These include a plant

architecture (Roux’s model; Hallé et al. 1978) with continuous stem or trunk growth, and

plagiotropic branches with distichous leaf arrangement (Fig. 3); non-resinous terminal

vegetative buds, and pendulous inflorescences. In comparison, all other members of

the Fagraea complex or the subtribe have episodic stem or trunk growth, with either

orthotropic complexes of branches, or branching that is plagiotropic by apposition or

substitution. Other genera in the Potaliinae have resinous terminal vegetative buds

Taxonomic framework for the Fagraea complex & new genus Limahlania 485

Fig. 2. Limahlania crenulata, showing Fagerlind’s tree architectural model with episodic trunk

development (producing branches in distinct tiers) and modular branch development producing

a plagiotropic system by substitution (each segment potentially terminated by a cyme) (Photo:

K.M. Wong). /nset: Trunk with prickles and developing a lightly fissured bark (Photo: M.

Sugumaran).

486 Gard. Bull. Singapore 64(2) 2012

Fig. 3. Growth architecture and flowering in Urania. Arrow indicating continuous orthotropic

stem growth in Urania volubilis following Roux’s model, where the shoot has no marked resting

period and stem internode lengths are generally similar; and branches are plagiotropic (Photo:

K.M. Wong). /nset: The pendulous, elongate terminal inflorescences with very condensed side-

branches characteristic of the genus, in another Utania species (Photo: M. Sugumaran).

: . Py 7 lar |

Taxonomic framework for the Fagraea complex & new genus Limahlania 48

fens = —_—______1— 5s 2 UMS

a .

wns F

1S)

fee

Sree

aK.

six

Fig. 4. Aubréville’s architectural model in Cyrtophyllum fragrans. A series of vertical brackets

shows distinct growth episodes on the monopodial trunk along which branching intervals

generally alternate as shorter and longer episodes. The brackets along the branch system

show repeated branch segments that develop plagiotropically by apposition (Photo: K.M.

Wong). /nsets: (top) The inflorescences are axillary and do not terminate the growth of branch

segments. (bottom) Deeply fissured bark of adult. (Photos: M. Sugumaran).

488 Gard. Bull. Singapore 64(2) 2012

Fig. 5. Picrophloeus javanensis. A. The tree architectural model of Scarrone found in this

genus is typified by orthotropic branch complexes. B. Terminal cymes, with long-exserted

stamens and styles in the flowers. (Photos: K.M. Wong)

Taxonomic framework for the Fagraea complex & new genus Limahlania 489

(Struwe, pers. comm.). In structure, the Urania inflorescence is an elongate panicle

with cymose branching (branches in several pairs, condensed, and distinctly shorter

than the rachis). In other members of the Fagraea complex and Potaliinae, solitary

flowers, 1—few-flowered cymes and branched cymes in which the basal branches are

longest (nearly as long as the rachis, and mostly rebranched) occur. In addition, Utania

fruits (like those of Anthocleista or Potalia; Struwe, pers. comm.) do not produce

any latex, whereas those of other members of the Fagraea complex do. As explained

above, Utania also has polygonal seeds, as in other members of the Fagraea complex

except for Fagraea s.s. itself (which, like other Potaliinae, have ellipsoid-rounded

seeds).

Cyrtophyllum (the Gigantea clade in the phylogenetic resolution of the

Fagraea complex) and Picrophloeus (Elliptica clade in the same resolution) have

superficial resemblance because of their small flowers and much-protruding stamens

and styles (Fig. 5), compared to other groups, which generally have bigger flowers

as well as less exserted stamens and styles. The only clear morphological distinction

between these two genera is an important one: axillary inflorescences in Cyrtophyllum

and terminal inflorescences in Picrophloeus (Fig. 4 and 5). Wong & Sugau (1996) had

recognised these two groups as distinct complexes within the section Cyrtophyllum

but even earlier, Ridley (1923) had accepted the group with axillary inflorescences (the

Gigantea clade) as the distinct genus, Cyrtophyllum.

Key to genera formerly placed in Fagraea s.1.

la. Inflorescences axillary. Trees with episodic stem / trunk growth, developing a

wave-like sympodial branch system with successively higher orders of outwardly

directed, then upturned, (indeterminate) branch segments (plagiotropy by

Saupe es AMER VT ON A ee eos eB daca vecteecnstadhtdesncsasdoessases-2 Cyrtophyllum

b. Inflorescences terminal. Trees, shrubs, scramblers or hemi-epiphytes with

continuous or episodic stem / trunk growth; branches plagiotropic or orthotropic,

if developing a wave-like sympodial branch systems then the branch segments

2) ERD ETE 0 GE S/P 121) | ane eee ee eee 2

2a. Stem / trunk growth continuous. Leaf arrangement on branches distichous.

Vegetative terminal buds not resinous. Inflorescence a pendulous flowering cyme

with all branches condensed along the rachis. Surface of dried fruits firm and

smooth, the epidermis not detaching from the pericarp ...............:::::eseeeee Utania

b. Stem / trunk growth episodic. Leaf arrangement on branches decussate. Vegetative

terminal buds covered with creamy to yellowish resin. Inflorescence without any

branching (a solitary flower) or an erect cyme with well-developed branches.

Surface of dried fruits wrinkled, the epidermis detaching from the pericarp ........ 3

490 Gard. Bull. Singapore 64(2) 2012

3a. Petiolar sheaths at the base of a leaf pair fully fused to form a cup-like ochrea.

Stigma base not expanding conspicuously, the stigma capitate. Stamens and style

exserted 40% or more from the corolla mouth eeeeeeee te ete Picrophloeus

b. Petiolar sheaths at the base of a leaf pair not fused to slightly fused at the extreme

edges and not forming an ochrea. Stigma base expanding into a circular plate-like

rim, the stigma appearing peltate. Stamen and style not or only slightly exserted

fromthe corolla south. ic: vias sche See eee sec cceesse aces eee

4a. Stem/trunk bark becoming fissured, densely set with prickles. Leaf margin serrulate-

crenulate. Branch system modular, developing plagiotropically by substitution to

form a wave-like sympodial system. Seeds polygonal .................0006 Limahlania

b. Stem / trunk with smooth bark, sometimes becoming cracking-scaly, but never

prickly. Leaf margin entire. Branch system non-modular, developing as orthotropic

complexes and not wave-like in structure. Seeds ellipsoid-rounded ........... Fagraea

Conspectus of genera in the complex

1. Cyrtophyllum Reinw.

in Hornschuch, Syll. Pl. Nov. 2 (1825) 8; Blume, Bidr. Fl. Ned. Ind. (1826) 1022;

Ridley, Fl. Malay. Pen. 2 (1923) 421. TYPE SPECIES: C. peregrinum Reinw. (= C.

fragrans (Roxb.) DC.).

Distribution. Bengal, Indo-China, Malay Peninsula, Sumatra, Java, Borneo, Mindoro,

Balabac, Palawan, Celebes and north-west New Guinea. Five species known.

NOTES ON TYPIFICATION:

Cyrtophyllum fragrans (Roxb.) DC., Prod. 9 (1845) 31.

Fagraea fragrans Roxb. [Hort. Beng. (1814) 84, nom. nud.], Fl. Ind. ed. Wall., 2

(1824) 32: Cammerl., Bull. Jard. Bot. Btzg. HI, 5 (1923) 318. TYPE: Hunter, Wallich

Cat. no. 1597E, “Pullo Penang” (holo K).

Cyrtophyllum peregrinum Reinw. in Hornschuch, Syll. Pl. Nov. 2 (1825) 9; Blume,

Bijdr. Fl. Ned. Ind. (1826) 1022. LECTOTYPE (here chosen): Reinwaradt s.n., Java (L

sheet no. 908.127-246 & barcode L0005030).

Cyrtophyllum was established with a single species, C. peregrinum, thus clearly

the type. The second species in Blume (1826), C. speciosum, is synonymous with

Picrophloeus javanensis Blume (1826).

The Wallich edition of the Flora Indica clearly mentions Hunter’s collection

from “Pullo Penang” under F- fragrans, so that the no. 1597E at Kew is the actual type.

There is also material at De Candolle’s Herbarium (GDC) numbered 1597A (from

Martabania, Burma, not taken as type material) and 1597C (from Penang, Malay

Peninsula, a possible isotype). There is a specimen at the Brussels Herbarium (BR,

barcode 0000006912710) from the Martius Herbarium, which bears an annotation

Taxonomic framework for the Fagraea complex & new genus Limahlania 49]

stating that this was from Roxburgh’s Herbarium and was obtained from an auction at

the Linnean Society of London; this is not directly attributable to Hunter as collector

and its status as type material is uncertain.

For lectotypification of C. peregrinum, it is to be noted that although Reinwardt

(1825) in Hornschuch (1825-1828) did not explicitly indicate a specimen together with

the species, the article in which his taxa were enumerated, Nova plantarum indicarum

genera, concerned the plants of Java. Thus, we have selected Reinwardt’s specimen

for this purpose.

2. Fagraea Thunb.

Kongl. Vetensk. Acad. Handl. 3 (1782) 132, t. 4. TYPE SPECIES: F. ceilanica Thunb.

Distribution. Sri Lanka, India, across Indo-China, S China and Malesia, to northern

Australia and Polynesia. Estimated 55 species.

NOTES ON TYPIFICATION:

Fagraea ceilanica Thunb., Kongl. Vetensk. Acad. Handl. 3 (1782) 132, t. 4.

LECTOTYPE (here chosen): Thunberg s.n., “e Ceilona” (UPS no. 004308; iso UPS

no. 004309, iso S).

The lectotype (UPS 004308) and isolectotype at S have flowering material, the

isolectotype at UPS has fruiting material only. The Stockholm duplicate was

annotated “Habitat in Ceylona” in 1778, the year Thunberg completed his Ceylon

visit. A specimen at the Linnean Society of London Herbarium (LINN barcode no.

LINN-HS292-1) without a clear indication of collector but with Thunberg’s species

name and publication reference for this name written on it, is a possible isotype.

3. Limahlania K.M.Wong & M.Sugumaran, gen. nov.

Fagraeae Thunb. sensu stricto similis sed architectura arboris, caulibus aculeis multis

brevibus, foliis serrulatis-crenulatis et seminibus polygonalis differt. TY PE SPECIES:

L. crenulata (Maingay ex Clarke) K.M.Wong & M.Sugumaran.

Limahlania is distinguished from allied genera in the Fagraea complex by its

development according to Fagerlind’s architectural model (featuring episodic trunk

growth and a modular branch system producing a wave-like sympodial series of axes),

stem bark bearing prickles that are shed with maturity, serrulate-crenulate leaf margin,

and polygonal seed shape. Limahlania crenulata (the Malayan malabera), like C.

fragrans (the tembusu padang tree), has become a tree of some horticultural note in

Southeast Asia, largely due to its ornamental form, beautiful blooms and adaptability

to different site conditions. Both are botanically fascinating trees (Corner 1940; Halle

et al. 1978; Gardner et al. 2011, swb Fagraea).

This new genus honours Professor Lim Ah Lan, who has taught and supervised

numerous students of plant biology at the University of Malaya, and who has been a

most encouraging colleague throughout her long career there, completing in 2012.

492 Gard. Bull. Singapore 64(2) 2012

Distribution. Malay Peninsula, Sumatra, Borneo (southwest and south Kalimantan),

south Indo-China (Cambodia, S Vietnam). A single species.

NOTES ON TYPIFICATION:

Limahlania crenulata (Maingay ex Clarke) K.M.Wong & M.Sugumaran, comb. nov.

Fagraea crenulata Maingay ex Clarke in Hooker f., Fl. Brit. Ind. 4 (1883) 83; King &

Gamble, J. As. Soc. Beng. 74 (2) (1908) 610; Cammerloher, Bull. Jard. Bot. Btzg. III, 5

(1923) 321; Ridley, Fl. Malay. Pen. 2 (1923) 420; Leenhouts, Fl. Males. I, 6 (2) (1962)

324; Kochummen, Tree Fl. Malaya 2 (1973) 271; Wong & Sugau, Sandakania 8 (1996)

59. TYPE: Maingay s.n., Malacca (K).

Fagraea fastigiata auct. non Blume (1838): Ridley, J. Str. Br. R. As. Soc. 30 (1897) 167

pro parte, quoad Ridley 7552.

The original description in Hooker’s publication only identifies the geographical

_ provenance as “Malacca” for a Maingay collection without indicating number. The

unnumbered sheet at K which was accessible to Clarke and Hooker is taken as type.

At least two numbered Maingay specimens (1032, 2976) of this species exist at other

herbaria (Leenhouts 1962b), so these are not considered type material, although

conceivably the notes in Hooker could have meant a representation of this species

via a number of Maingay’s collections from Malacca, including both numbered and

unnumbered material.

4. Picrophloeus Blume

Bidr. Fl. Ned. Ind. (1826) 1019. TYPE SPECIES: P. javanensis Blume.

Distribution. Sumatra, Java, Malay Peninsula, Borneo, Celebes, Maluku, New Guinea.

Four species.

NOTES ON TYPIFICATION:

Picrophloeus javanensis Blume, Bijdr. Fl. Ned. Ind. (1826) 1020. LECTOTYPE (here

chosen): Blume s.n., Java (L: 908.127-201, barcode LOO0S006).

Blume (1826) provided a brief description in Latin for the genus Picrophloeus but

not the species P. javanensis, which was the sole species listed. A single description,

or descriptio generico-specifica, for valid publication of both a genus and species, is

accepted according to Article 42.1 of the Vienna Code (McNeill et al. 2006), when the

genus has but a single species, at least at the time of description. Under the species,

Blume stated “in sylvis altioribus montis Salak’, which 1s attributable to the specimen

Blume, Java at the Leiden Herbarium (L: sheet no. 908.127-201). Therefore the genus

name Picrophloeus and the species P. javanensis are to be regarded as validly published.

5. Utania G.Don

Gard. Dict. 4 (1838) 663. TYPE SPECIES: U. morindifolia (Blume) G.Don (= U.

volubilis (Wall.) M.Sugumaran).

Taxonomic framework for the Fagraea complex & new genus Limahlania 493

Kuhlia Reinw. ex Blume, Bijdr. Fl. Ned. Ind. (1826) 777, nom. illeg., non Kunth

(1825). Kentia Steud., Nomencl. ed. 2 (1840) 845, nom. illeg., non Blume (1838).

Distribution. Indo-China (Cambodia, S Vietnam), the Andaman and Nicobar Islands,

Malay Peninsula, Sumatra, Java, Borneo, Celebes, Maluku, the Philippines and New

Guinea. Estimated 15 species.

NOTES ON TYPIFICATION:

Utania volubilis (Wall.) M.Sugumaran, comb. nov.

Fagraea volubilis Wall. in Roxb., Fl. Ind. ed. Wall. 2 (1824) 36; Wong & Sugau,

Sandakania 8 (1996) 40. TYPE: Jack, E. Bencoolen (Herb. Wallich, sheet marked

“1600. E. Bencoolen” on bottom left) (holo K).

Kuhlia morindifolia Blume, Bijdr. Fl. Ned. Ind. (1826) 777; F. morindifolia (Blume)

Blume, Rumphia 2 (1838) 32, t. 73; U. morindifolia (Blume) G.Don, Gard. Dict. 4

(1838) 663. LECTOTYPE (here chosen): Anon. (probably Blume), Java (L: sheet no.

908.127-731, barcode L0005046; isolecto L: sheet no. 908.127-721; isolecto K).

Utania was a new name for the illegitimate Kuhlia and was accompanied by a genus

description (in English, acceptable under Article 36.1 of the Vienna Code: McNeill et

al. 2006). Don listed a single species under his Utania as “1. K. MORINDAEFOLIA”

(in error for what he should have indicated as “1. U. MORINDAEFOLIA” consistent

with the general format of his enumeration of taxa). Under this species, he referred

to the locality “Native of Java, at the foot of Mount Salak” and also cited “Kuhlia

morindaefolia Blum. bijdr. p. 777” (1826), in which there is a description. Thus Urania

is a validly published genus name and the only one available for use in reference to

Fagraea section Racemosae.

ACKNOWLEDGEMENTS. The Keepers and Curators of the A, BO, K, KEP, KLU, L, NY, US,

SAN, SAR and SING herbaria are thanked for loans of, and permission to study, specimens.

J.J. Wood, J. Dransfield, N. Martland (Kew), E.F. de Vogel, C.E. Ridsdale, J.-F. Veldkamp,

L. Willemse (Leiden), C.C. Davis, D. Boufford, W. Zhang, K. Gandhi (Harvard), L.G. Saw,

R.C.K. Chung (Kepong), J.B. Sugau, J.T. Pereira, M. Postar (Sandakan), C.L. Chan (Kota

Kinabalu), A.A. Joffre, K. Ariffin (Brunei), R. Kiew (formerly SING), S. Lee, C.J.H. Soh and

M.I. Bazilah (SING) are thanked for assisting with a whole range of research matters. Y.W.

Low (SING) assisted in fieldwork and research, procurement and assessment of information

related to types and specialised collections. Research for, and preparation of, this paper have

also benefitted from assistance and comments from L. Struwe (Rutgers University); N. Xia

(South China Institute of Botany); C. Puff (University of Vienna), and A.L. Lim (University

of Malaya). We thank C. Puff for help with the Latin diagnosis used here. The following are

also thanked for their support during the course of the research: M. Sofian-Azirun, Dean of

Science, and R. Hashim, Head of the Institute of Biological Sciences, University of Malaya, as

well as the previous Deans, A.M. Mohamed and A. Nasrulhaq-Boyce. A University of Malaya

Fundamental Research Grant FS 264/2007C helped finance a baseline phylogenetic study, on

which the present contribution is partly based.

494 Gard. Bull. Singapore 64(2) 2012

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new tribal and subtribal classification. In: Struwe, L. & Albert, V.A. (eds.), Gentianaceae—

systematics and natural history. Pp. 21-309. U.K., Cambridge: Cambridge University

Press.

Struwe, L. & Albert, V. A. (2004) A monograph of neotropical Potalia (Gentinaceae: Potalieae).

Syst. Bot. 29: 670-701.

Sugumaran, M. & Wong, K.M. (2012) Studies in Malesian Gentianaceae I: Fagraea sensu

lato—complex genus or several genera? A molecular phylogenetic study. Gard. Bull.

Sing. 64(2): 301-332.

Thunberg, C.P. (1782) Beskrifning pa et nytt och vackert orte Genus, Kalladt Fagraea ceilanica.

Kongl. Vetensk. Acad. Handl. 3: 132—134, tab. IV.

Wong, K.M. & Sugau, J.B. (1996) A revision of Fagraea (Loganiaceae) in Borneo, with notes

on related Malesian species and 21 new species. Sandakania 8: 1-93.

Gardens’ Bulletin Singapore 64(2): 497-510. 2012 497

Studies in Malesian Gentianaceae III:

Cyrtophyllum reapplied

to the Fagraea fragrans alliance

K.M. Wong! and M. Sugumaran?

'Singapore Botanic Gardens,

1 Cluny Road, Singapore 259569

wkm2000@gmail.com

*Rimba IImu Botanic Garden,

Institute of Biological Sciences, University of Malaya,

50603 Kuala Lumpur, Malaysia

maran(@um.edu.my

ABSTRACT. Cyrtophyllum Reinw., one of several distinct lineages among the Fagraea

complex, is the correct genus to which five species of Southeast Asian trees should be assigned,

including the widespread F. fragrans. Cyrtophyllum minutiflorum K.M.Wong is a new species

described here. Two new combinations are made: C. caudatum (Ridl.) K.M.Wong and C.

wallichianum (Benth.) M.Sugumaran & K.M.Wong.

Keywords. Cyrtophyllum, Fagraea fragrans, Gentianaceae, Malesia, Potalieae, Potaliinae,

Southeast Asia

Introduction

The results of a molecular phylogenetic study of the Fagraea complex (Sugumaran

& Wong 2012) demonstrated the distinctness of a number of generic lineages from

Fagraea Thunb. s.s. (Wong & Sugumaran 2012). Among these, Cyrtophyllum Reinw.

and Picrophloeus Blume were readily distinguished from Fagraea s.s., Limahlania

K.M.Wong & M.Sugumaran and Urania G.Don because the first two genera have

flowers with conspicuously exserted styles (typically more than 40% of their length)

and filaments (greater than 70% of their length) (Sugumaran & Wong 2012). Also,

Cyrtophyllum and Picrophloeus frequently have cymes bearing numerous small

flowers (corollas narrow, the mouth often not more than 10 mm wide), compared to

the other genera, which typically have larger flowers (corollas typically much wider)

in variable numbers. However, Cyrtophyllum has axillary cymes and Aubréville’s tree

architectural model, whereas Picrophloeus and the other three genera all have terminal

cymes and consistently other architectural models (Scarrone’s in Picrophloeus and

Fagraea s.s., Fagerlind’s in Limahlania, Roux’s in Utania) (Sugumaran & Wong

2012; Wong & Sugumaran 2012).

The history of Cyrtophyllum is interesting. Ridley (1893) had used the name

C. fragrans (Roxb.) DC. Later, he described an allied species under Fagraea, but

nevertheless appreciated (Ridley 1918) that this genus name was available for the

alliance of trees that was quite different from Fagraea: “It would probably be best to

498 Gard. Bull. Singapore 64(2) 2012

keep up the genus Cyrtophyllum for the Tembusu trees which differ so much from the

epiphytic true Fagraeas with their fleshy leaves and flowers, and included stamens,

from the tall trees with their leaves and flowers and long projecting stamens. The

genus Cyrtophyllum would thus contain C. fragrans..., C. giganteum..., C. wallichii...,

C. caudatum...” In his Flora of the Malay Peninsula (Ridley 1923), he adopted

Cyrtophyllum for this group of species. Burkill (1935), on the other hand, took the

broad view as Blume (1838) did, and, under Fagraea, stated: “Cyrtophyllum... which

some botanists separate, is not really distinct, and is united here.” The uncertainties

have found their way as different taxonomic approaches throughout this period, viz.,

as a section of Fagraea (Blume 1838); as a subgenus of Fagraea (Miquel 1857); and

more recently as Blume’s section again but with more discordant elements thrown in

(Leenhouts 1962; see also Wong & Sugau 1996).

Here we provide a revision of Cyrtophyllum, which includes the well-known

tree species in Southeast Asia, C. fragrans (synonym Fagraea fragrans Roxb.).

Cyrtophyllum Reinw.

in Hornschuch, Syll. Pl. Nov. 2 (1825) 8; Blume, Bidr. Fl. Ned. Ind. (1826) 1022;

Ridley, Fl. Malay. Pen. 2 (1923) 421; Wong & Sugumaran, Gard. Bull. Sing. 64 (2012)

490. TYPE SPECIES: C. peregrinum Reinw. (= C. fragrans (Roxb.) DC.).

Fagraea sect. Cyrtophyllum (Reinw.) Blume, Rumphia 2 (1838) 34 pro parte excl.

F. speciosum; Leenhouts, Fl. Males. I, 6 (1962) 303 pro parte excl. F. elliptica & F-

umbelliflora, Wong & Sugau, Sandakania 8 (1996) 7 pro parte excl. F- elliptica. —

Fagraea sect. Parviflorae Benth., J. Linn. Soc. Bot. 1 (1857) 100 pro parte quoad C.

peregrinum. — Fagraea subg. Cyrtophyllum (Reinw.) Miq., Fl. Ind. Bat. 2 (1857) 375

pro parte excl. F. sumatrana.

Medium-sized to large trees usually more than 3 m and some reaching 25—30 m tall.

Trunk monopodial with episodic growth; developing wave-like sympodial branch

systems or ‘terminalian’ branching with successively higher orders of outwardly

directed, then upturned, (indeterminate) branch segments (plagiotropy by apposition)

(Aubréville’s architectural model fide Hallé et al. 1978). Trunk bark becoming fissured

in older trees. Vegetative shoot apices with light yellowish resin. Leaf arrangement

on branches decussate; leaf margin entire; petiolar sheaths of a leaf pair fused and

forming a shallow cup-like ochrea that loosely clasps the stem; petiolar base auricles

absent. Inflorescence axillary, a few- to many-flowered and few- to multi-branched

cyme, basal branches nearly as long as or longer than rachis. Flowers bisexual, small,

up to 10 mm wide at the corolla mouth; calyx lobes 5; corolla white to creamy white,

corolla lobes 5, overlapping to the right; stamens 5, typically two thirds or more

exsert, anthers versatile, sagittate; style typically one third or more exsert; stigma

capitate (knob-like). Fruits small, subglobose to slightly ellipsoid, up to about 10

mm in diameter; colour at maturity yellow-orange to bright red; with small amounts

Cyrtophylium reapplied to Fagraea fragrans and allies 499

of translucent sticky latex in fruit epidermis and fruit wall: epidermis separating as

a thin translucent film from pericarp (fruit surface appearing crinkled on herbarium

specimen). Seeds numerous: placentation axile: polygonal: surface areolate.

Distribution and diversity. Bengal, Burma, Thailand, Indo-China, Andaman Islands,

Malay Peninsula, Sumatra, Java, Borneo, Mindoro, Balabac, Palawan, Celebes and

north-west New Guinea. Five species known.

Habitat. Lowland forests.

Key to Cyrtophyllum species

la. Inflorescence branching to just 1, rarely 2, orders. Corolla tubes 10-25 mm ane

stamens with filaments 16-38 mm long; styles 14-55 mm long.. rat a2

b. Inflorescence branching to 24 orders. Corolla tubes not ae 10 mm aie ng;

stamens with filaments 7-17 mm long: styles 9-22 mm long..............0.......... 3

2a. Leaf apex caudate. Calyx 2.5-3 mm across, corolla tube subcylindric (not

conspicuously flared in its upper part), 10-12 mm long. (Borneo) .......................-

= pesgcnsSheeee neces Looe eee ee C. caudatum

b. Leaf apex acuminate. Calyx 34 mm across, corolla tube infundibular (flared in

its upper part), 12-25 mm long. (Malay Peninsula) ................... C. wallichianum

3a. Corolla tube infundibular (flared in the upper half), larger (lower narrowed part of

corolla 1.5—2(—2.5) mm wide); leaf secondary veins flat or slightly raised on the

CE CEP IER PST Teer rh Ure rel rey 6 | ne C. fragrans

b. Corolla tube subcylindric (not noticeably flared in the upper part), smaller (lower

narrowed part of corolla 1-1.5 mm wide): leaf secondary veins flat to frequently

sunken on the upper leaf surface in dried material ........ 2.0... eee eee 4

4a. Leaves chartaceous, margin conspicuously wavy (more conspicuously so in

fresh material), secondary veins forking towards the leaf margin but not forming

conspicuous loops; pedicels 3-5 mm long (flowers distinctly pedicellate): corolla

tube 7-8 mm long: style 12-22 mm long. (Widespread across Malesia.) .............-.

Sey cise eR Sag ee C. giganteum

b. Leaves coriaceous, margin plane, secondary veins forming distinct loops towards

the margin: pedicels 0-1 mm long (flowers subsessile); corolla tube 4-5 mm

long: style 9-10 mm long. (Restricted to Borneo.) ..................--- C. minutifiorum

1. Cyrtophyllum caudatum (Ridl.) K.M.Wong. comb. nov. — Fagraea caudata Ridl.,

J. Str. Br. Roy. As. Soc. 79 (1918) 97; Merr., J. Str. Br. Roy. As. Soc. Sp. No. (1921)

491: Wong & Sugau, Sandakania 8 (1996) 8: Coode et al. Checkl. FI. Pl. Gymn. Brunei

(1996) 174. TYPE: Lobb s.n., 1853, ‘Borneo’ (Sarawak) (holo K).

500 Gard. Bull. Singapore 64(2) 2012

Fagraea fragrans auctt. non Roxb., Leenhouts, Fl. Males. I, 6 (1962) 304 pro parte,

Ashton, Manual Non-Dipt. Trees Sarawak 2 (1988) 310 pro parte, quoad F. caudata

in syn.

Tree, 6—7 m tall, perhaps bigger; trunk to 2.5 cm diameter or more; bark smooth, dark

brown. Leaves elliptic to slightly oblanceolate; (4—)7—9(—11) cm long, (1.5—)2.5—3.5(—

4) cm wide; base cuneate; apex caudate, 1—1.5 cm long; margin plane when fresh and

in dried specimens; coriaceous; upper and lower surfaces smooth; midrib prominent

below, sunken above; secondary veins 7—9 pairs, upper side faint and immersed in the

blade, lower side faint to very slightly sunken or even prominent; tertiary veins faint

to inconspicuous; petioles 0.8—2.2 cm long, 1.5—2 mm thick, petiolar sheaths of a leaf

pair fused and forming a shallow cup-like ochrea that tightly clasps the stem; petiolar

base auricles absent. Inflorescence a few-flowered cyme 5—9.5 cm long; peduncle

2.5—6 cm long, 1(—1.5) mm thick; with just a single level (tier) of branching on the

main axis, typically not rebranched. Flower pedicel 5-10 mm long, c. | mm thick;

calyx (from the base to the lobe apices) 3.5—4 mm long, glabrous, calyx cup 2.5—3 mm

wide, calyx lobes erect, 1—1.5 mm long, 1—1.5 mm wide, margins glabrous; corolla

tube subcylindric, 10-12 mm long, 1.5—2 mm wide near the base, inside glabrous to

minutely papillate; corolla lobes broad-ovate to subobovate, 5-6 mm long, 4-5 mm

wide; stamens inserted at the uppermost third of the corolla tube; filaments 16-18 mm

long, exsert 11-12 mm in the open flower; anthers not seen; style 14-15 mm !ong,

protruding 11-13 mm from the corolla mouth in the open flower; stigma c. 0.3 mm

across. Infructescence peduncle 4—7.5 cm long, 1—1.5 mm thick. Fruits (immature)

to 5-6 mm across; the base loosely to tightly clasped by the calyx lobes. Seeds not

examined.

Distribution. Endemic to NW Borneo (SW Sabah, Brunei, Sarawak).

Habitat. Hills and ridges in lowland mixed dipterocarp forest.

Specimens examined: BORNEO. Brunei. Belait, Batu Patam, along ridge north of summit,

Wong WKM 1060 (BRUN, K, SING). Sarawak. Beccari 2956 (K); 1st Div. Gunung Santubong

South, Bujang S. 12999 (K, L, SAR, SING); Bako N.P., Lintang path, Chai S. 19707 (A, BO,

K, KEP, L, MEL, MOSC, SAN, SAR, SING), Bukit Gondol, //ias S. 17908 (A, BO, K, KEP,

L, SAN, SAR, SING), Md. Shah P. 5647 (A, K, L, SAR, SING); Lobb s.n. 1853 (holo K); Ist

Div. Mt. Matang, near Valombrosa, M. & J. Clemens 7783 (K); Mt Mike Low LYW 213 (KLU);

Bintulu, Nyabau F.R., Brunig S. 12050 (SAR); Lambir N.P., Sg. Jangkang, Mokhtar et al. S.

47187 (K, KEP, L, MO, SAN, SAR); Kuching, Ulu Sg. Rayu, Yakup S. 7716 (SAR, SING);

Kelaung F.R., sine coll. S. 7431 (SAR).

Ridley (1918) observed that the species he was describing under Fagraea, when

appropriately transferred to Cyrtophyllum, should be C. caudatum, although that

combination was not effected.

Cyrtophyllum reapplied to Fagraea fragrans and allies S01

2. Cyrtophyllum fragrans (Roxb.) DC., Prod. 9 (1845) 31; Ridley, Trans. Linn. Soc.

Lond. 2nd ser. Bot. 3 (1893) 323. — Fagraea fragrans Roxb. [Hort. Beng. (1814) 84,

nom. nud.|, Fl. Ind. ed. Wall., 2 (1824) 32; Don, Gard. Dict. 4 (1837) 68: Blume, Mus.

Bot. 1 (1850) 172; Benth., J. Linn. Soc. Bot. 1 (1857) 100; Mig., Fl. Ind. Bat. 2 (1857)

375; Kurz, For. Fl. Brit. Burm. 2 (1877) 205; Clarke in Hooker f., Fl. Brit. India 4

(1883) 85; Ahern, Compil. Timber Tree Philip. Isl. (1901) 88, pl.; Koord. & Val., Bijdr.

9 (1903) 86; Ridley, J. Str. Br. Roy. As. Soc. 50 (1908) 121; Elm., Leafl. Philip. Bot. 2

(1909) 597; Dop, Fl. Gén. I.-C. 4 (1914) 177; Merrill, J. Str. Br. Roy. As. Soc. Sp. No.

(1921) 492; Cammerl., Bull. Jard. Bot. Btzg. III, 5 (1923) 318; Foxw., Mal. For. Rec.

3 (1927) 157; Corner, Wayside Trees Malaya (1940) 424: Kerr in Craib, Fl. Siam. En.

3 (1951) 55; Browne, For. Trees Sar. & Brun. (1955) 244, t. 32; Smythies, Common

Sarawak Trees (1965) 87; Kochummen, Tree Fl. Malaya 2 (1973) 273; Cockburn,

Trees Sabah | (1976) 211; Wong & Sugau, Sandakania 8 (1996) 11: Coode et al.

Checkl. Fl. Pl. Gymn. Brunei (1996) 174. Willughbeia fragrans (Roxb.) Spreng.

Syst. Veg. 4 (1827) Cur. Post. 71. TYPE: Hunter, Wallich Cat. no. 1597E, “Pullo

Penang” (holo K).

Cyrtophyllum peregrinum Reinw. in Hornschuch, Syll. Pl. Nov. 2 (1825) 9; Blume,

Bijdr. Fl. Ned. Ind. (1826) 1022; DC., Prod. 9 (1845) 31; Ridley, Fl. Malay. Pen. 2 (1923)

421; Holttum, Gard. Bull. S.S. 5 (1931) 189. — Fagraea peregrina Blume, Rumphia 2

(1838) 34, t. 80; Mus. Bot. 1 (1850) 172. LECTOTYPE (Wong & Sugumaran 2012):

Reinwardt s.n., Java (L sheet no. 908.127-246 & barcode L0005030).

Fagraea cochinchinensis A. Chev., Cat. Pl. Jard. Bot. Saigon 33 (1919) 66, pro

specim., Merrill, Enum. Philip. Fl. Pl. 3 (1923) 314, pro specim., non basionym Aidia

cochinchinensis Lour., Fl. Cochinch. | (1790) 143.

Fagraea fragrans auctt. non Roxb.: King & Gamble, J. As. Soc. Beng. 74 (2) (1908)

611 pro parte, excl. Derry 272; Leenhouts, Fl. Males. I, 6 (1962) 304 pro parte,

Ashton, Manual Non-Dipt. Trees Sarawak 2 (1988) 310 pro parte, excl. F: caudata

Ridl., F gigantea Ridl., F. speciosa sensu Ridl. non Blume, F: sororia J.J.Sm., F:

wallichiana Benth., F: lanceolata Wall. in syn.; Kochummen, Tree Fl. Malaya 2 (1973)

273 pro parte, excl. Cyrtophyllum lanceolatum, Fagraea wallichiana in syn.; Griffin

& Parnell, Fl. Thailand 6 (3): 198 pro parte, excl. C. giganteum Ridl. in syn.

Tree, rarely to just 3-4 m tall, more often big, to 30 m tall; trunk to over 1 m in

diameter; bark deeply and ruggedly fissured, grey-brown to dark brown. Leaves

elliptic; (5.5—)7.5—11(—13) cm long, (2—)3.5—4.5(—5.3) cm wide; base cuneate; apex

short-cuspidate to caudate; (3—)5—8(—9) mm long; margin plane when fresh (in dried

specimens sometimes slightly wavy); chartaceous to thin-coriaceous; upper and lower

surfaces smooth; midrib prominent below, flat to slightly raised above; secondary

veins (7—)9—12 pairs, upper side faint and immersed in the blade, lower side faint to

very slightly prominent; tertiary veins faint to inconspicuous; petioles 1—1.3(—1.6) cm

502 Gard. Bull. Singapore 64(2) 2012

long, 1—1.5 mm thick. Inflorescence (3.5—)4—7(—7.7) cm long; peduncle (2.8—)3-3.5

cm long, I—1.5 mm thick; with (2—)3(-4) levels (tiers) of branching on the main axis,

the branch tiers (2—)3—5(—6) mm apart, lowest tier typically branched to 2(—3) orders.

Flower pedicel (3—)5—8(—11) mm long, 0.5(—1) mm thick; calyx (from the base to

the lobe apices) (1.5—)2—2.5 mm long, glabrous, calyx cup (1.5—)2—2.5(—3) mm wide,

calyx lobes erect, |—1.5(—2) mm long, 1—1.5(—2) mm wide, margins glabrous; corolla

tube somewhat infundibular, (4—)6—8 mm long, lower narrowed part 1.5—2(—2.5) mm

wide, upper flared part (3—)4—6(—7) mm wide at the top, the lower narrowed tubular

part nearly as long as the upper flared part, inside glabrous to minutely papillate;

corolla lobes broad-ovate to subobovate, (3—)4-6 mm long, 2—3(-4) mm wide;

stamens inserted at the middle of the upper flared part of the corolla tube; filaments

(10—)12—16(—17) mm long, exsert (8—)10—12(—13) mm in the open flower; anthers

1—1.5 mm long, 0.5—1 mm wide, each anther sac somewhat narrowly ellipsoid; style

(14—)18—22 mm long, protruding (8—)10—12(—14) mm from the corolla mouth in the

open flower; stigma c. 0.5 mm across, lobes 2, low and rounded, parting to present two

slightly raised, hemispherical, papillate inner surfaces when receptive, not recurving.

Infructescence peduncle (2.5—)3.5—5(—6.2) cm long, 1-2 mm thick. Fruit when

mature to 4-6 mm across; the base loosely to tightly clasped by the calyx lobes. Seeds

0.5—1 mm across.

Distribution. Principally a mainland SE Asia and W Malesian species: Bengal,

Andaman Islands, Burma, Thailand, Indo-China (Laos, Vietnam, Cambodia), Malay

Peninsula, Sumatra, Java, Borneo, SW Philippines (the Palawan chain from Balabac

NW through Palawan, Culion and Busuanga, and continuing into Mindoro); also

Celebes.

Habitat. One of our most commonly encountered trees in West Malesia, Cyrtophyllum

fragrans establishes easily in open areas and secondary forests including along

roadsides. It grows easily on sandy sites, including around sandy tailings of former

tin-mines in the Malay Peninsula. It is less commonly found in lowland high forest,

but easily encountered in kerangas (tropical heath forest) vegetation and coastal or

beach forest.

Specimens examined: BORNEO. Brunei. Belait, Anduki F.R., Anderson S. 4941 (SAR),

Andulau, Taman Rekreasi Hutan Sungai Liang, Sugumaran et al. SM 227 (KLU), Sungai

Liang, Hussain HUS 30 (BRUN, SING). Kalimantan. Bandjermasin, Boschwezen 2176

(BO); Sanggau, Pengoedang, Neth. Ind. For. Service bb 29174 (SING); Sentarum Wildlife

Reserve, far north-west corner of Danau, Sungei Seriang, Giesen 140 (K), western border of

Danau, Nanga Kenelang, Giesen 148 (K); Sungai Kenara, Hallier 1359 (P, SING). Sabah.

“N Borneo”, Vilamil 300 (P); Beaufort, Cuadra NBFD A 1377 (K, SING); Gaya Island F.R.,

Kuripin SAN 28778 (SING), Gaya island (south), Aban SAN 57839 (SAN); Jesselton, Wood

2557 (SING); Kimanis, Bayvak NBFD 2114 (K); Kota Belud, Kampung Lantige & Abai R.

F.R., Kandilis 7118 (SING); Kuala Penyu, Road to Pantai Tanjung Aru, Rimi et al. SPN 06605

(KEP); Mempakul, Bangawan, Abubakar NBFD 4111 (K, SING); Papar, Mandahan, Dewol &

Termiji SAN 80011 (SAN, SING), Talip Bidin SAN 80685 (KEP, SAN, SING); Sandakan, below

Cyrtophylium reapplied to Fagraea fragrans and allies 503

Mr. Fox House, 50 ft, Jali SAN 65627 (SAN, SING); Sandakan, below Sabah Hotel, Sam SAN

25509 (SAN, SING); Sandakan, Chinese Cemetery, Chow & Aban SAN 75925 (SAN, SING):

Sandakan, Elopura, Agama A 2102 (SING), Cuadra A 2218 (SING), Darby road, Cuadra A

3197 (K, SING); Sandakan, Ernestina Road, Meijer SAN 24941 (SAN, SING); Sandakan, Jalan

Kapok, Clemente 4963 (SING): Sandakan, Leila F.R., 300 ft, Leopold & Termiji SAN 76680

(SAN, SING): Sandakan, Mile “% Derby road, Aban Gibot SAN 75916 (KEP, SAN); Sandakan,

Taman Rimba Sport Complex, Dewol SAN 105553 (SAN, SING). Sarawak. Beccari 3428

(K); J. & M.S. Clemens 22561] (P: barcode P03976237); Kuching, Museum Garden, Mamit S.

33492 (SAR): Kuching, Taman Budaya, Yahud et al., S. 57555 (K, KEP, L, MO, SAN, SAR):

Simunjan, Balai Ringin P-F., Stipni Bin Dollah S. 303 (SAR, SING), 1705 (SAR): Sadong, Ulu

Gunong, Omar 64 (SING).

BURMA. Martabania. Wallich Cat. no. 1597a (GDC: barcode G00134008), Wallich Cat.

no. 1597b (P: barcode P00349660). Wallich s.n. (P: barcode P00349658). Tenasserim &

Andamans. Helfer 3735 (P: barcode P00349659).

CAMBODIA. Bejaud 740 (P: barcode P00349636); Dam ta trau, Poilane 1407] (P: barcode

P00349651). Kompong Thom. Prey Krabey, reserve Sambo Spieu, Magnen s.n. 1909 (P:

barcode P00349640). Kompong Spen. entre Pum Love & Pum Rong, Poilane 17553 (P:

barcode P00349652). Siem Reap. Phnom Kulen, Martin 22] (P: barcode 00349644).

CELEBES. Boni. Heyne 2863 (BO). Malili. 25 m, Neth. Ind. For. Service Cel/IV-86 (SING).

Cel/IV-136 (SING), Cel/IV-137 (SING), Tabarano, Neth. Ind. For. Service bb 32356 (SING).

Moena. Wapai, Neth. Ind. For. Service bb 21738 (P: barcode P03976230). Manado.

Kolonodale, 400 m, Neth. Ind. For. Service bb 31525 (SING). Moena. Makoemoro?, 25 m,

Neth. Ind. For. Service bb 26302 (SING), bb 26303 (SING).

INDIA. Roxburgh s.n. (BR: barcode BRO000006912710).

JAVA. sine coll. no date (L); Blume s.n. (GDC: barcode G00134012): Korthals s.n. (L: barcode

L0005029); Reinwardt, s.n. (L sheet no. 908.127-246 & barcode L0005030). Batavia. Bodjong,

Bakhuizen v/d Brink 6354 (SING); Bodjong Ejot, Bakhuizen v/d Brink 6354 (P: barcode

P03976232); Buitenzorg, Hallier 202 (BO), Welter s.n. (SING), Moera Enim, Teysmann 4019

(BO). Semarang. Ngarengan kult., Kalshoven s.n. (BO). Soerabaja. Insel Bawean, Tambak,

Teysmann 1767 (BO).

LAOS. Banthat a 12 km de Savannakhet, Poilane 16384 (P: barcode P00330715), mare a 20

km de Savannakhet, Vidal 1752 (P: barcode P00330717).

MOLUCCAS. Smith s.n. 1796-1805 (BM: barcode 001053465).

PENINSULAR MALAYSIA. Johor. Johore Bharu, Carrick 1406 (SING). Kedah. Rahim KEP

12386 (SING). Melaka. Merlimau, A/vins s.n. (SING), Derry 53 (SING). Negeri Sembilan.

Cubitt 706 (SING); Forest Dept. 697 (SING); Angsi Forest Reserve, Othman KEP 23732

(KEP): Bahau Reserve, Mat Deris 654 (SING); Port Dickson, Blue Lagoon, Bremer 1819

(KLU), Cape Rachado F.R., roadside to Light House, Tsou 207 (NY); Seremban, Bain 18856

(KEP). Pahang. Ridley 1028 (SING); Kuantan, Abdul Rahman FMS 4172 (SING), Mahamud

FMS 3729 (SING), Mohd Soh 15735 (SING); Muazam Shah-Menchali main road, Sugumaran

et al. SM 212, leafy branch only (KLU):; Pulau Berhala, Burn-Murdoch SFN 303 (SING).

Penang. Askey FMS 2554 (SING); “Pullo Penang”, Hunter, Wallich Cat. no. 1597E, (holo

K), Wallich Cat. no. 1597c (GDC: barcode G00134009); Sungai Pinang, Ogata 10344 (KEP):

Tasek Gelugor, Fyfe 29321 (SING): Tulloh (sic! = Teluk) Bahang, Curtis 314 (SING), Curtis

s.n. 1893 (SING), v 1892 (SING), vii 1892 (SING). Perak. Pangkor Island, Telok Nipah,

Chin 3137 (KLU). Selangor. Gombak, Klang Gates Quartz Ridge, Daniel Lee et al. s.n., leafy

branch only (KLU); Kepong, Forest Research Institute, Vethevelu FRI 29649 (KEP, SING);

Kuala Lumpur, Ramli KEP 94100 (KEP), Government Hill, Kalong 17463 (SING), Circular

504 Gard. Bull. Singapore 64(2) 2012

Road, Omar 7425 (SING), University of Malaya campus, Zan Bee Cheok s.n. (KLU), Zakiah

Hassan s.n. (KLU). Terengganu. Kemaman, Osman FMS 26996, FB, FL, 13 vii 1971 (KEP).

PHILIPPINES. Mindoro. Merritt 8S42 (PNH, US?). Palawan. Curran Bur. For. 4501 (PNH,

US?); Aborlan, Iraan Mountains, Sulit 12512 (SING); Palawan, Cenabre 29990 (SING);

Taytay, Merrill 1202 (SING), Roque 30274 (SING).

SINGAPORE. Cantley’ Collector s.n. (SING); Botanic Gardens, Henderson 1329 (SING);

Nature Reserves Sector 17, Turner et al. NRS 269 (SING); Pulau Pawai, Sidek S 99 (SING);

Surrounding reservoir, Cantley s Collector s.n. (SING!).

SUMATRA. Palembang. Endert 881 (SING), Moesi-Oeloe, Endert 160 (BO), 161 (BO).

Riauw islands. Pulau Kundor, Jerwyva 1624 (SING).

THAILAND. Bangkok. Marcan 707 (SING). Krabi. Krabi-Trang Road, Weerachai Nanakorn

WN 514 (SING); Jurin?, Kerr 8232 (SING). Surat Thani. Prsundej et al. 249 (KLU); Ko

Pha-ngan, Phromdet 25 (P: barcode P00349653). Narathiwat. Bacho, Sangkhachand 184 (P:

barcode P00349654); Sungai Padi, Bourke s.n. 1924 (P: barcode P00349655). Trat. Taphan

Hin, Geesink 6490 (P: barcode P00349656).

VIETNAM. Harmand 804 (P: barcode P00349666). Minh Hai. Iles de Poulo Condor, Harmand

~ 873 (P: barcode P00349668), Pierre 3699 (P: barcode P00349677). Tourane. Quang-Nam, Da-

Nang, J. & M.S. Clemens 3862 (P: barcode P00349662). Lam Dong. Dalat & environs, Squires

889 (P: barcode P00349680). Long. Quang Tri, Chevalier 41189 (P: barcode P00349663).

A drawing of this species by William Hunter, collector of the type specimen, is on

display in the Penang Museum and a picture of this work 1s found in Gardner et al.

(2011: 116). In the past, two other species found in the Malay Peninsula, namely,

C. giganteum and C. wallichianum have been confused with C. fragrans. These two

species are not as common as C. fragrans; for differences, see under those species.

Burkill (1935) used the name Fagraea cochinchinensis A.Chev. for this

species, but apparently this had been based on Aidia cochinchinensis Lour. (a name

dating from 1790; Rubiaceae) and was therefore in the first instance misapplied to

material of F) fragrans (Leenhouts 1962). Arguably, both taxa have some resemblance

in their paired leaves and branched cymes of small, cream-coloured to white flowers

with exserted styles and stamens.

Holttum (1935) has given an account of the flowering of this species in

Singapore, which is gregarious (many or nearly all trees in a population with

synchronised flowering). It has two flowering seasons, the main one around mid-year

and another towards year’s end, the flowers opening several weeks following the

bud stage. At such times, it may be appreciated how different nomenclatural systems

may converge on an essential trait, for the species epithet in the latinised scientific

name refers to fragrant flowers, and its Malay name, tembusu, could well allude to the

conspicuous sweet-and-sour scent that transpires into more than a hint of fermentation

(Malay: busw, unpleasantly odourous). Apparently, the flowers last several days

even though the stamens are spent after the first day of bloom, thus sustaining and

intensifying the perfumed aura around the tree. The fruits, which take a few months

to mature to an attractive yellow to red, are probably mainly dispersed by birds or

bats (Corner 1940). Of this tree, Ridley (1893: 323) observes that “it is difficult to say

whether the tree is more beautiful when covered with flowers or fruits.”

Cyrtophyllum reapplied to Fagraea fragrans and allies 50

3. Cyrtophyllum giganteum (Ridl.) Ridl., Fl. Malay. Pen. 2 (1923) 421. — Fagraea

gigantea Ridl., J. Str. Br. R. As. Soc. 79 (1918) 98; Foxw., Mal. For. Rec. 3 (1927)

157; Burkill, Dict. (1935) 995; Corner, Wayside Trees Malaya (1940) 425; Browne,

For. Trees Sar. & Brun. (1955) 245. Kochummen, Tree Fl. Malaya 2 (1973) 273:

Cockburn, Trees Sabah | (1976) 211; Wong & Sugau, Sandakania 8 (1996) 11; Coode

et al. Checkl. Fl. Pl. Gymn. Brunei (1996) 174. LECTOTYPE (Wong & Sugau 1996):

Ridley 8921, Singapore, Garden Jungle (SING).

Fagraea sororia J.J.Sm. ex Cammerl., Bull. Jard. Bot. Btzg. IH, 5 (1923) 319, pl. 5;

Heyne, Nutt. Pl. (1927) 1270. LECTOTYPE (Wong & Sugau 1996): Endert 44E 1P

515, Sumatra, Palembang (L; isolecto BO, K, SING).

Fagraea fragrans auctt. non Roxb.: King & Gamble, J. As. Soc. Beng. 74 (2) (1908)

611 pro parte, quoad Derry 272; Leenhouts, Fl. Males. I, 6 (1962) 304 pro parte,

Ashton, Manual Non-Dipt. Trees Sarawak 2 (1988) 310 pro parte, quoad Cyrtophyllum

giganieum Ridl., F- gigantea Ridl., F- sororia J.J.Sm., F. speciosa (non Blume) Ridl.

in syn.

Fagraea speciosa auct. non Blume: Ridley, J. Str. Br. R. As. Soc. 50 (1908) 122.

Tree, rarely to just 3-4 m tall, more often big, to 30 m tall or more; trunk to over |

m in diameter; bark closely and finely fissured, grey-brown to dark brown. Leaves

elliptic; (4.4-)5—7(—7.5) cm long, (1.8—)2.2—3(—3.2) cm wide; base cuneate; apex

short-cuspidate to caudate; 4-6 mm long; margin conspicuously wavy when fresh

and in dried specimens; chartaceous; upper and lower surfaces smooth; midrib

prominent below, flat to sunken above; secondary veins 4—6 pairs, upper side faint and

immersed in the blade, lower side faint to very slightly prominent; tertiary veins faint

to inconspicuous; petioles (0.8—)1—1.5(—2.3) cm long, 1—1.5 mm thick. Inflorescence

(3.5—)S—6.5(—7.5) cm long; peduncle 3.5—5(—5.5) cm long, 1—1.5 mm thick; with 4-5

levels (tiers) of branching on the main axis, the branch tiers (S—)6—10(—12) mm apart,

lowest tier typically branched to (2—)3 orders. Flower pedicel 3—5 mm long, 0.3—0.5

mm thick; calyx (from the base to the lobe apices) 1.5—2 mm long, glabrous, calyx cup

1.5—2 mm wide, calyx lobes erect, |—1.5 mm long, 1—1.5 mm wide, margins glabrous;

corolla tube subcylindric, 7-8 mm long, 1—1.5 mm wide near the base, upper part

very gradually and slightly wider, inside glabrous to minutely papillate: corolla lobes

broad-ovate to subobovate, 3-4 mm long, 2—2.5(—3.5) mm wide; stamens inserted at

the uppermost third of the corolla tube; filaments 13—15 mm long, exsert 12—13 mm in

the open flower; anthers 1—-1.5 mm long, 0.5—0.8 mm wide, each anther sac somewhat

narrowly ellipsoid; style (12—)18-22 mm long, protruding (5—)12—14 mm from the

corolla mouth in the open flower: stigma c. 0.5 mm across, lobes 2, low and rounded,

parting to present two slightly raised, hemispherical, papillate inner surfaces when

receptive, not recurving. Infructescence peduncle (2.5—)3-4(-4.8) cm long, 1—1.5 mm

thick. Fruit when mature to 4-6(—7) mm across; the base tightly clasped by the calyx

lobes. Seeds 1-2 mm across.

506 Gard. Bull. Singapore 64(2) 2012

Distribution. Malay Peninsula, Sumatra, Borneo.

Habitat. Lowland mixed dipterocarp forest.

Specimens examined: BORNEO. Brunei. Belalong, Ulu Ropan, 2000-2500 ft, Ashton BRUN

5275 (BRUN, K, SING); Tutong, Ulu Tutong, 150 ft, Ashton BRUN 908 (BRUN, K, SING).

Kalimantan. East Kutai, Sangkulirang district, Kerajaan River region, Kostermans 34793

(BO, K, L), Sungai Kerajaan, 40 m, Kostermans 5804 (BO, SING), Sungai Susuk, 40 m,

Kostermans 5693 (BO, K, SING); Muara Teweh, Popas?, Neth. Ind. For. Service bb 27769

(SING); Nunukan Island, Kostermans 8612 (BO, SING), 8955 (BO, SING); S.-O. Borneo,

Horyoep?, Winkler 2468 (SING); Sanggau, Bindang, Neth. Ind. For. Service bb 28143 (BO, K,

SING). Sabah. Keningau, Nabawan, Dewol & Karim SAN 78059 (K, L, SAN, SAR, SING);

Sandakan, Patrick SAN 25509 (KLU, SAN), Batu Sapi road, Meijer SAN 24942 (K, KEP,

L, SAN, SAR); Tawau, Apas Road, Brand SAN 21500 (SAN, SING), Kawa Road, Jaswir &

Aban SAN 26277 (BO, K, KEP, L, SAN, SING), Table Estate, 200 ft, Wood SAN 16650 (SAN,

_ SING). Sarawak. Baram, Melinau Gorge (4°10°N, 114°55’E), 1000 ft, Chew CWL 444 (K,

SING), Ulu Sungai Melinau Paku, Anderson 4085 (K, L, SAN, SAR, SING); Kuching, Bukit

Hujan, Omar 353 (K, SING); Lawas, Mt Bugoh ridge, Smythies BRUN 812 (K); Limbang, Ulu

Medamit, 900 ft, Chai, Wright & Othman S. 32335 (K, L. SAR, SING); Miri, S. Ukong, 10 m,

Othman S. 21394 (SING).

PENINSULAR MALAYSIA. sine coll. 16711 (KEP); Johor. Ulu Endau, Labis F.R., Sungai

Jasin, Ogata KEP 110427 (KEP). Melaka. Air Keroh Botanical Garden, Sugumaran & Lee

SM 193, leaves and stem bark (KLU), SM 194, leaves only (KLU); Bukit Saliokor?, Derry

272 (SING). Negeri Sembilan. Kuala Kelawang, Triang F.R., Lau & Jalil FRI 18249 (KEP).

Pahang. Kuantan, Mohd Soh FMS 15750 (SING), Rahman 15738 (SING). Selangor. Ayer

Hitam F.R., sine coll. KEP 55873, leaves only (KEP); Kajang, Bangi Reserve, Foxworthy

10289 (SING); Sungai Buluh F.R., Walton FMS 30770 (KEP). SINGAPORE. Singapore.

Garden Jungle, Ridley 8921 (lecto SING).

SUMATRA. Palembang. Buurman van Vreeden 77 (BO); Endert 44E IP 429 (BO, L), 44E

IP 515 (BO, K, L, SING), 44E 2P 673 (BO, L); Banjoeasin en Koeboestreken, Endert 44E 2P

706 (BO, K, L); Lematang Ilir, Semangoes, Neth. Ind. For. Service bb 32270, leafy branch only

(SING); Medan 19, Bengkalis, Houtvester s.n. (BO); Moeara Doea Sh., Grashoff 400 (BO);

Moeara Enim, Zeysmann 3796 (BO, L); Rawas, Dumas 1550 (BO, K), Grashoff 1060 (BO,

L). Lampongs. Kebang, Zeysmann 4210 (BO, L); Tandj-Penang, Bruinsma 12 (BO). Upper

Riauw. Tenajan, Pakanbaru, Soepadmo 232 (BO, K, SING).

Cyrtophyllum giganteum and C. fragrans have very similar flowers that appear only to

have consistently different corolla tube widths. However, several vegetative features

are very distinctive, such as the distinctively wavy margins in fresh leaves of C.

giganteum (C. fragrans have leaf margins which are generally plane). There are more

pairs of secondary veins in C. fragrans (7—9(-11)) that form distinct loops towards

the leaf margin but in C. giganteum there are fewer pairs of secondary veins (4—6)

that fork towards the leaf margin and do not form conspicuous loops. The mature

tree form is also often distinguishable: C. giganteum develops a very straight and

columnar bole like many other lowland forest canopy species, whereas C. fragrans

usually reaches up to only 30 m and typically develops several erect main branches

that are as tall as the main trunk. A well-grown C. fragrans tree develops a coarse,

Cyrtophyllum reapplied to Fagraea fragrans and allies 507

reticulately-sinuously ridged and fissured bark, as pictured in Gardner et al. (2011:

117); but that in a mature C. giganteum tree, as shown in Plate 35 of Ashton (1988:

313), is regularly and shallowly fissured.

4. Cyrtophyllum minutiflorum K.M.Wong, sp. nov. C. caudato Ridley similis sed

cymis ramosioribus, floribus minoribus tubis corollae brevioribus (4—5 mm longis) et

stylis brevioribus (9-10 mm longis) differt. TYPE: Burley, Tukirin et al. 3278, 1-6 Jul

1989, Borneo, West Kalimantan province, G. Bentuang area, 5-10 km north of Masa

village, ridge SW of G. Bentuang, 1100 m alt. (holo SING; iso A, BO, K).

Tree, documented as 10 m tall; trunk to 15 cm diameter; bark texture in mature tree

trunk not known. Leaves elliptic to slightly lanceolate or oblanceolate; (5—)8—10(—

11) cm long, (1—)2—3(—3.5) cm wide; base cuneate; apex acuminate, hardly 0.5 cm

long; margin plane in dried specimens; coriaceous; upper and lower surfaces smooth;

midrib prominent below, channelled to raised above; secondary veins (4—)5—7 pairs,

upper side faint and immersed in the blade, lower side faint to immersed; tertiary

veins faint to inconspicuous; petioles (S—)12—18 cm long, 1.5—2 mm thick, petiolar

sheaths of a leaf pair fused and forming a shallow cup-like ochrea that tightly clasps

the stem; petiolar base auricles absent. Inflorescence a few-flowered cyme 8—10 cm

long; peduncle 44.5 cm long, 1(—1.5) mm thick; with 1(—2) levels (tiers) of branching

on the main axis, lowest tier typically branched to 3-4 orders. Flower pedicel 0—1 mm

long (flowers subsessile), c. 0.5 mm thick; calyx (from the base to the lobe apices) c.

1.5 mm long, glabrous, calyx cup c. 1.5 mm wide, calyx lobes erect, c. 0.5 mm long, c.

0.5 mm wide, margins glabrous; corolla tube subcylindric, 10-12 mm long, c. | mm

wide near the base, inside glabrous to minutely papillate; corolla lobes broad-ovate

to subobovate, 2.5—3 mm long, c. 2—2.5 mm wide; stamens inserted at the uppermost

third of the corolla tube; filaments c. 7 mm long, exsert c. 5 mm in the open flower;

anthers not seen; style 9-10 mm long, protruding c. 4 mm from the corolla mouth

in the open flower; stigma c. 0.1 mm across. Infructescence peduncle 4-5 cm long,

1—1.5 mm thick. Fruit (immature) c. 2 mm across; the base loosely to tightly clasped

by the calyx lobes. Seeds not examined.

Distribution. Borneo, W Kalimantan, only known from the type collection.

Habitat. On a ridge at the upper limit of Mixed Dipterocarp Forest or its transition to

lower montane forest.

5. Cyrtophyllum waliichianum (Benth.) M.Sugumaran & K.M.Wong, comb. nov.

— Fagraea wallichiana Benth., J. Linn. Soc. Bot. 1 (1857) 98; Clarke in Hooker

f., Fl. Brit. Ind. 4 (1883) 85; King & Gamble, J. As. Soc. Beng. 74 (2) (1908) 607;

Corner, Ways. Trees Malaya (1940) 426; Wong, Saw & Kochummen, Malayan Nat. J.

41 (1987) 267; Wong & Sugau, Sandakania 8 (1996) 12. — Fagraea lanceolata Wall.

508 Gard. Bull. Singapore 64(2) 2012

[Cat. (1829) no. 1599, nom. nud.] Schnizl., Iconogr. 2 (1851) t. 131, f. 1, nom. illeg.,

non Blume (1826); Miquel, Fl. Ind. Bat. 2 (1857) 376; Burkill, Dict. (1935) 995.

Cyrtophyllum lanceolatum (Wall.) DC., Prod. 9 (1845) 31, nom. illeg.; Ridley, Fl. Mal.

Pen. 2 (1923) 421. TYPE: Wallich Cat. no. 1599, Penang (holo K; iso BM, GDC).

Fagraea fragrans auctt. non Roxburgh (1824): Kochummen, Tree Fl. Malaya 2 (1973)

273 pro parte, quoad Fagraea wallichiana Benth. in syn.; Leenhouts, Fl. Males. I, 6

(2) (1962) 304, pro parte, Ashton, Manual Non-Dipt. Trees Sarawak 2 (1988) 310

pro parte, quoad Cyrtophyllum lanceolatum DC., F- lanceolata Wall., F- wallichiana

Benth. in syn.

Tree, rarely to just 3-4 m tall, more often bigger, to 25 m tall; trunk to over 1 m in

diameter; bark fissured, grey-brown to dark brown. Leaves narrow to broadly-elliptic

to oblanceolate to obovate; (3.4—)6—10(—13) cm long, (1.4-)2—3(—5) cm wide; base

- cuneate; apex acute to short-cuspidate; 2—5(—1.2) mm long; margin plane when fresh

and in dried specimens; thin-coriaceous; upper and lower surfaces smooth; midrib

prominent below, sunken above; secondary veins (4—)5—7 pairs, upper side faint to

immersed in the blade, lower side faint to very slightly prominent; tertiary veins faint

to inconspicuous; petioles (0.3—)0.8—1.2(—1.7) cm long, 1—1.5 mm thick, petiolar

sheaths of a leaf pair fused and forming a shallow cup-like ochrea that tightly clasps

the stem; petiolar base auricles absent. Inflorescence a few-flowered cyme (2.5—)3—

6(—7.8) cm long; peduncle (1.9—)2.3—3.5(—5.3) cm long, 1(—1.5) mm thick; with 1(—2)

levels (tiers) of branching on the main axis, the branch tiers 10-14 mm apart, lowest

tier typically branched to 1(—2) orders. Flower pedicel (4—)5—8 mm long, 1—1.5 mm

thick; calyx (from the base to the lobe apices) (3—)4—S(—6) mm long, glabrous, calyx

cup 3-4 mm wide, calyx lobes erect, 2—3 mm long, 2—2.5 mm wide, margins glabrous;

corolla tube somewhat infundibular, (12—)20—25 mm long, lower narrowed part 1—2

mm wide, upper flared part 6—8(—10) mm wide at the top, the lower narrowed tubular

part nearly as long as the upper flared part, inside glabrous to minutely papillate; corolla

lobes broad-ovate to subobovate, (S—)7—8 mm long, 3—4.5(—5) mm wide; stamens

inserted at the lower end of the upper flared part of the corolla; filaments (27—)30-38

mm long, exsert 20—23(—28) mm in the open flower; anthers 1—1.5 mm long, 0.5—1

mm wide, each anther sac somewhat narrowly ellipsoid; style (34—)42-45(-S5) mm

long, protruding 22—25(—30) mm from the corolla mouth in the open flower; stigma

c. 0.5 mm across, lobes 2, low and rounded, parting to present two slightly raised,

hemispherical, papillate inner surfaces when receptive, not recurving. Infructescence

peduncle 2—2.5(—3) cm long, 1—1.5 mm thick. Fruit when mature to 7—9 mm across;

the base loosely to tightly clasped by the calyx lobes. Seeds 1—1.5 mm across.

Distribution. Endemic to the Malay Peninsula.

Habitat. Lowland forest on hills and ridges.

Specimens examined: PENINSULAR MALAYSIA. Johor. Hutan Simpan Labis, Gunung

Cyrtophyllum reapplied to Fagraea fragrans and allies 509

Beremban, Wong FRI 30882, leafy branch only (KEP). Kedah. Gunung Jerai, Kochummen

FRI 18086 (KEP, SING), FRI 29495 (KEP), KEP 85027, leafy branch only (KEP), KEP 94416

(KEP). Kelantan. Relai Forest Reserve, Cockburn FRI 725] (KEP, SING), FRI 7280 (KEP,

SING), FRI 7411 (KEP, SING). Malacca. Maingay 1029 (BM, mounted on the same sheet.

lower half, as the isotype Wallich Cat. no. 1599). Penang. Wallich Cat. no. 1599 (holo K; iso

BM, GDC); Govt. Hill, Curtis s.n. 1899 (SING), Ridley 7066 (SING): Moniots Road, Burkill

SFN 3330 (SING), Curtis 375 (SING); Penang Hill, Ahmad Shukor AS 91 (SING), Corner SFN

31597, leafy branch only (SING), Low et al. LYW 206, leafy branch only (KLU), Nauen s.n.

1940 (SING), Samsuri Ahmad SA. 999 (KEP, KLU, SING), Sidek S. 226 (SING!), Symington

KEP 28043 (SING); Western Hill, Nauwen s.n. 1940 (SING). Terengganu. Ulu Brang, Gunung

Padang, Whitmore FRI 12803 (KEP, SING).

Cyrtophyllum wallichianum is generally a smaller tree (up to 25 m tall) of hills

and ridges, compared to C. fragrans, which can grow taller (up to 30 m tall) but is

typically found on gentler terrain in the lowlands. C. wallichianum is vegetatively

quite similar to C. fragrans but the floral characters are different. The inflorescence

of C. wallichianum is usually less branched, 1(—2) orders, with fewer flowers; that

of C. fragrans is generally more branched, 2(—3) orders, and so bears more flowers.

The corollas of C. wallichianum are much bigger, 12—25 mm long, 6-10 mm wide,

whereas in C. fragrans they are smaller, (4—)6—8 mm long and 1.5—2(—2.5) mm wide.

As noted by Ridley (1918), FE caudata (= C. caudatum) which occurs only

in Borneo (Sarawak and Brunei), closely resembles C. wallichianum. Cyrtophyllum

caudatum differs from C. wallichianum in having more coriaceous, lanceolate-caudate

leaves and extremely slender peduncles and pedicels, cylindric corolla tubes and

shorter stamens.

Indeterminate material

A taxon that resembles C. caudatum and C. minutiflorum in leaf characters, but without

flowering or fruiting material, occurs in New Guinea. Its habitat, too, is similar: ridge

forest in hilly terrain, only even higher at c. 800 m elevation. This is quite possibly a

distinct species.

Specimens examined: NEW GUINEA. Japen Island, c. 800 m, Neth. Ind. For. Services bb.

30336 (BO, SING), bb. 30352 (BO, SING).

ACKNOWLEDGEMENTS. We thank the Keepers and Curators of the A, BO, K, KEP, KLU,

L, NY, US, SAN, SAR and SING herbaria for permission to consult reference materials and

loans of specimens. J.J. Wood, J. Dransfield, N. Martland (Kew), E.F. de Vogel. C.E. Ridsdale,

J.-F. Veldkamp, L. Willemse (Leiden), C.C. Davis, D. Boufford, W. Zhang, K. Gandhi

(Harvard), L.G. Saw, R.C.K. Chung (Kepong), C.L. Chan (Kota Kinabalu), A.A. Joffre, K.

Ariffin (Brunei), R. Kiew (formerly SING), S. Lee, C.J.H. Soh and M.I. Bazilah (SING) are

thanked for assisting with a whole range of research matters. Y.W. Low, Singapore Botanic

Gardens, assisted with research and assessment of information related to types and specialised

collections. We thank C. Puff for help with the Latin diagnosis used.

510 Gard. Bull. Singapore 64(2) 2012

REFERENCES

Ashton, P.S. (1988) Manual of the Non-Dipterocarp Trees of Sarawak. Vol. 2. Kuching,

Sarawak: Dewan Bahasa dan Pustaka.

Blume, C.L. (1838) Rumphia. Vol. 2. Leiden.

Burkill I.H. (1935) A Dictionary of the Economic Products of the Malay Peninsula. Fourth print

(2002). Vol. 1: 1009-1013. Kuala Lumpur: Ministry of Agriculture and Co-Operatives.

Corner, E.J.H. (1940) Wayside Trees of Malaya. 2 vols. Singapore: Government Printing Office.

Gardner, S., Sidisunthorn, P. & Lai, E.M. (2011) Heritage Trees of Penang. Penang: Areca

Books.

Holttum, R.E. (1935) The flowering of tembusu trees (Fagraea fragrans Roxb.) in Singapore,

1928-1935. Gard. Bull. Straits Settlem. 9: 73-78.

Leenhouts, P.W. (1962) Loganiaceae. In: Steenis, C.G.G.J. van (ed.), Flora Malesiana. Series

1 Vol. 6. Pp. 293-336. Groningen: Wolters-Noordhoff Publishing.

Miquel, F.A.W. (1857) Flora van Nederlandsch-Indié. Vol. 2. Amsterdam.

_ Ridley, H.N. (1893) On the flora of the eastern coast of the Malay Peninsula. Zrans. Linn. Soc.

London, Bot. 3: 267-408 & Pl. 61-66.

Ridley, H.N. (1918) New and rare Malayan plants. J. Straits Branch Roy. Asiat. Soc. 79: 97-98.

Ridley, H.N. (1923) The Flora of the Malay Peninsula. Vol. 2. London: L. Reeve & Co.

Sugumaran, M. & Wong, K.M. (2012) Studies in Malesian Gentianaceae I: Fagraea sensu

lato—complex genus or several genera? A molecular phylogenetic study. Gard. Bull.

Singapore 64(2): 301-332.

Wong, K.M. & Sugau, J.B. (1996) A revision of Fagraea (Loganiaceae) in Borneo, with notes

on related Malesian species and 21 new species. Sandakania 8: 1-93.

Wong, K.M. & Sugumaran, M. (2012) Studies in Malesian Gentianaceae II: A taxonomic

framework for the Fagraea complex, including the new genus Limahlania. Gard. Bull.

Singapore 64(2): 481—495.

Gardens’ Bulletin Singapore 64(2): 511-522. 2012 511

Studies in Malesian Gentianaceae IV:

A revision of Picrophloeus

K.M. Wong

Singapore Botanic Gardens,

1 Cluny Road, Singapore 259569

wkm2000@gmail.com

ABSTRACT. Picrophloeus Blume is the correct generic assignment for four species forming

one of several distinct lineages in the Fagraea complex. They include the widespread Southeast

Asian P. javanensis Blume, otherwise commonly known by the dubious name F: e//iptica Roxb.

Three new combinations are made for species known only in Borneo: P. belukar (K.M.Wong &

Sugau) K.M.Wong, P. collinus (K.M.Wong & Sugau) K.M.Wong, and P. rugulosus (K.M.Wong

& Sugau) K.M.Wong.

Keywords. Borneo, Fagraea elliptica, Gentianaceae, Malesia, Picrophloeus, Potalieae,

Potaliinae, Southeast Asia

Introduction

A molecular phylogenetic analysis of the Fagraea complex (Sugumaran & Wong

2012) revealed the distinctness of a number of generic lineages from Fagraea Thunb.

s.s. (Wong & Sugumaran 2012). These included two genera, Cyrtophyllum Reinw. and

Picrophloeus Blume, that are distinguished from the other lineages by their flowers

with conspicuously exserted styles (typically more than 40% of their length) and

filaments (greater than 70% of their length) (Sugumaran & Wong 2012). Picrophloeus

is readily distinguished from Cyrtophyllum by its terminal cymes (Cyrtophyllum has

axillary cymes) and Scarrone’s tree architectural model (Cyrtophyllum is characterised

by Aubréville’s model) (Sugumaran & Wong 2012; Wong & Sugumaran 2012).

Picrophloeus is presently revised. It includes the widespread P. javanensis

Blume, which has been much identified with the name Fagraea elliptica Roxb., a

problematic name here shown to be dubious.

Picrophloeus Blume

Bijdr. Fl. Ned. Ind. (1826) 1019; Wong & Sugumaran, Gard. Bull. Sing. 64 (2012)

492. TYPE SPECIES: P. javanensis Blume.

Fagraea sect. Cyrtophyllum (Reinw.) Blume, Rumphia 2 (1838) 34 pro parte excl.

FE. peregrina; Leenhouts, Fl. Males. I, 6 (1962) 303 pro parte excl. F. fragrans & F-

umbelliflora, Wong & Sugau, Sandakania 8 (1996) 7 pro parte excl. F. fragrans. —

ay Gard. Bull. Singapore 64(2) 2012

Fagraea sect. Eufragraea Blume, Mus. Bot. | (1850) 172 pro parte quoad F- kimangu,

F. picrophloea & F; speciosa. — Fagraea sect. Parviflorae Benth., J. Linn. Soc. Bot.

1 (1857) 100 pro parte excl. C. peregrinum. — Fagraea subg. Cyrtophyllum (Reinw.)

Migq., Fl. Ind. Bat. 2 (1857) 375 pro parte excl. F: lanceolata.

Medium-sized to large trees to about 30 m tall. Trunk monopodial with episodic

growth, developing orthotropic complexes of branches (Scarrone’s architectural

model fide Hallé et al. 1978). Trunk bark becoming fissured in older trees. Vegetative

shoot apices with light yellowish resin. Leaf arrangement on branches decussate;

leaf margin entire; petiolar sheaths of a leaf pair fused to form a shallow cup-like

ochrea. Inflorescence terminal, a many-flowered and branched cyme, basal branches

nearly as long as or longer than rachis. Flowers bisexual, small, up to 10 mm wide at

the corolla mouth; calyx lobes 5; corolla lobes 5, overlapping to the night, narrowly

elliptic to lanceolate; stamens 5, inserted at the corolla mouth, typically two thirds

or more exsert, anthers versatile; style typically one third or more exsert; stigma

capitate (knob-like). Fruits small, subglobose, up to about 10 mm in diameter; colour

at maturity yellow-orange to bright red; with small amounts of translucent sticky latex

in fruit epidermis and fruit wall; epidermis separating as a thin translucent film from

pericarp (fruit surface appearing crinkled on herbarium specimens). Seeds numerous,

placentation axile; polygonal; surface areolate.

Distribution and diversity. Sumatra, Java, Malay Peninsula, Borneo, Maluku, New

Guinea. Four species known.

Habitat. Lowland to lower montane forests.

Key to Picrophloeus species

la. Leaf surface drying very coarsely wrinkled (resembling a coarsely pitted surface

to the touch). Cymes distinctly pedunculate, the peduncle to I—1.5 cm long

beyond the ultimate leaf pair on the flowering shoot. Flowers sessile (the ultimate

floral bracteole immediately below the calyx cup). Corolla tubes 8—11 mm long.

Filaments in open flowers 15—18 mm long. Fruits larger, 6—9(—10) mm across ....

ee ee eA ce arene ccdo ae ada crsnoaaacesdecumbucr Gdbsconck P. rugulosus

b. Leaf surface drying smooth cr only finely shagreen. Cymes sessile, the basal

branches not clearly elevated from the distalmost leaf pair on the flowering shoot

and appearing as if they originate from the ultimate leaf axils. Flowers pedicellate

(the pedicel at least a short but distinct axis between the ultimate bracteole and

the calyx base). Corolla tubes shorter, 3.5—6 mm long. Filaments in open flowers

(4—)6—11 mm long. Fruits smaller, 3.5—5(—6) mm ACTOS .............eeeeeeeeeseeeeeees 2

Picrophloeus revised ais)

2a. Leaf apex typically obtuse-rounded to emarginate. Inflorescence typically

branched to 5—6 orders. Corolla tube 3.54 mm long .................0::e P. belukar

b. Leaf apex acuminate to cuspidate-caudate. Inflorescence branched to 3—4(—5)

DSSS Gore LA inves: 2 08100 (00 | 3

3a. Leaves thick-coriaceous, secondary veins obscure on lower side. Pedicels 0.5—1

mm long. (Restricted to Borneo, submontane to lower montane forest) .......

ee Pe Stee se See A ake eer aoe Stok suas as cele cebeeawoncalees P. collinus

b. Leaves thin-coriaceous, secondary veins distinct and prominent on the lower side.

Pedicels 1-2 mm long. (Widespread: Sumatra, Java, Malay Peninsula, Borneo,

Moluccas to New Guinea; lower montane forest generally but in the Wallacea

region also in the lowlands, including coastal forest) ...............0 P. javanensis

1. Picrophloeus belukar (K.M.Wong & Sugau) K.M.Wong, comb. nov. — Fagraea

belukar K.M.Wong & Sugau, Sandakania 8 (1996) 15; Coode et al. Checkl. Fl. PI.

Gymn. Brunei (1996) 174. TYPE: Saikeh SAN 72151, Sabah, Beaufort, Beaufort Hill

(holo SAN).

F. elliptica auctt.: Leenhouts, Fl. Males. I, 6 (1962) 303 pro parte; Cockburn, Trees

of Sabah 1 (1976) 210 pro parte; Ashton, Manual Non-Dipt Trees Sarawak 2 (1988)

314 pro parte; Anderson, Check List Trees Sarawak (1980) 239 pro parte. [Fagraea

elliptica Roxb. sensu stricto, F\. Ind. ed. Wall. 2 (1824) 32, nom. dub. |

Tree, to around 30 m tall; trunk to over 1 m in diameter; bark fissured, dark grey-

brown. Leaves obovate to broad-elliptic; 7-22 cm long, 4-13 cm wide; base cuneate;

apex obtuse-rounded to emarginate; margin plane to recurved; coriaceous; upper

and lower surfaces smooth to finely shagreen; midrib prominent below, rounded to

ridged; secondary veins 10—12 pairs, upper side faint and flat or immersed in the blade,

lower side faint to immersed, not prominent; tertiary veins inconspicuous; petioles

34.5 cm long, stout. Inflorescence sessile, the basal branches not clearly elevated

from the distalmost leaf pair on the flowering shoot and appearing as if they originate

from the ultimate leaf axils, to c. 13 cm long, 15—20 cm across, main axis branched

to 5—6 orders, with 5—6 tiers of branches; lowest branches typically 3—5.5 cm long.

Flower pedicel (above the ultimate bracteole) 1-2 mm long; calyx (from the base to

the lobe apices) 1-2 mm long, calyx cup 1-2 mm wide; corolla salverform; cream to

white; corolla tube 3.54 mm long, not conspicuously flared, |-1.5 mm wide, inside

glabrous to minutely papillate: corolla lobes 2-3 mm long, c. 1.5 mm wide; stamens

with filaments 6-8 mm long in the open flower; anthers oblong, c. | mm long; style

11-15 mm long in the open flower; stigma knob-like, c. 0.5 mm across, lobes 2, low

and rounded, not recurving. Fruit when mature to 3.5—5 mm across; fruit calyx lobes

to 1 mm long and wide. Seeds c. | mm across.

514 Gard. Bull. Singapore 64(2) 2012

Distribution. Borneo, all districts; Banka (Bangka) Island.

Habitat. Lowland secondary forest, forest gaps, open sites. This species is a conspicuous

member of the coastal tropical heath (kerangas) forest community of Borneo, easily

seen, for example, in the Sandakan, SW Sabah, Brunei and Sarawak Ist Division

areas. It is also common in secondary forest and some open degraded sites in Borneo

(Wong & Sugau 1996).

Specimens examined: BORNEO. Brunei. Belait, Andulau F.R., /dris et al. BRUN 15655

(BRUN, SING); Berakas, Berakas FR, Ashton S. 7816 (BRUN, SING); Tutong, Smythies

BRUN 0836 (SING), Bukit Beruang, Forman & Blewett 810 (BRUN, K, SING). Kalimantan.

Bangarmassing, Motley s.n. 1857 (K); East Kutai, Sungai Bambangan, Kostermans 6092

(SING); Long Bagun, Wiriadinata 602 (BO, K); Tarakan, Kostermans s.n. December 1963

(SING). Labuan. Botanic Garden, Natural arboretum, Talip SAN 55629 (SAN). Sabah.

Beaufort, Beaufort Hill, Madani SAN 36770 (K, L, SAN, SAR), Meijer NT 172 (SAN), Saikeh

SAN 72151 (holo SAN), SAN 72256 (SAN, SING), 72259 (K, L, SAN, SAR, SING), Kg.

Inuman, Mikil SAN 30215 (SAN), Mantanior, Jalib & Marsal SAN 84772 (SAN, SING),

mile 52 Mesapul road, Mikil SAN 34569 (SAN); Jesselton, Gaya Island F.R., Lajangah SAN

28776 (BO, K, KEP, L, SAN, SAR, SING), Wyatt-Smith KEP 80298 (K, SING), (Pulau)

Gaya, Sinanggul SAN 40136 (KEP), near Ulu Soyong river, Aban SAN 57828 (KEP, L, SAN);

Jesselton, near hospital, Zandom NBFD 2810 (SAN); Kota Kinabalu, Bukit Padang, Beaman

10822 (K, MSC); Sandakan, Postar & Ahmad SAN 147987 (SAN), junction of Ernestina Road,

Brand SAN 34698 (BO, K, KEP, L, SAN, SAR, SING), Leila F.R., Ahwing SAN 39033 (K,

L), Ampuria SAN 32601 (K, L, SAN, SAR, SING), Jawanting & Ampuria SAN 32601 (SAN,

SING), Kanis & Ding Hou SAN 57451 (K, L, SAN), Kumin SAN 74160 (K, L, SAN, SAR,

SING), Wong SAN 22901 (K, KEP, L, SAN, SAR); mile 3/4 North Road below Sabah Hotel,

Sam & Ernesto SAN 26522 (SAN), mile 4 Leila F.R., Dewol & Chow SAN 74258 (K, KEP, L,

SAN, SING), Pum station mainline, Ampon & Madani SAN 47347 (SAN), town, Meijer SAN

20945 (SAN, SING), Sepilok Laut, Jamin SAN 131250 (SAN), Termiji & Paul SAN 85500

(SAN); Sipitang, Caudra A 3293 (K), Bukit Ulu Sipitang, Dewol & Termiji SAN 78358 (K,

KEP, L, SAN, SAR, SING), SFI area, Amin SAN 126518 (SAN), forest behind SFI Qrs., Madani

& Amin SAN 86236 (SAN), Sri Beaufort Area, Dewol SAN SO1I1 (SAN, SING), Merintaman

F.R., Saikeh SAN 72327 (SAN, SING), Seungau, Cuadra A 3293 (SING); Tawau, Padang

Golf, Ismail SAN 107761 (SAN); Weston, Bukit Siungau, Talip SAN 80618 (K, KEP, L, SAN,

SAR, SING), Hutan Simpan Sianggau, Amin SAN 126720 (SAN, SING), SAN 132146 (SAN),

Lubok Darat, Dewol & Karim SAN 78153 (SING), SAN 78358 (SAN). Sarawak. Moulton s

Native Collectors s.n. 1909 (SING); Bukit Lan, Wright 637 (SAR); Kuching, Anderson S.

4921 (SAN), Corner & Brunig 10444 (SAR), Dickson 3 (SING), Kuching Reservoir, Anderson

& Chai S. 28943 (SAR, SING), Selang, Sungai Pasir, Rehal S. 2251 (SING); Mukah, Ulu

Kenyana, Ashton S. 19490 (SING); Serian, Sungai Sabal Tapang, Sinclair SFN 10266 (E, K, L,

SAR, SING); Tarum, Rehal 1750 (SAR).

SUMATRA. Bangka. Jacobs IV-A-127 (SING).

2. Picrophloeus collinus (K.M.Wong & Sugau) K.M.Wong, comb. nov. — Fagraea

collina K.M.Wong & Sugau, Sandakania 8 (1996) 19. TYPE: Aban SAN 50747, Sabah,

Ranau, copper mining area (holo SAN; iso A).

Picrophloeus revised a5

F. elliptica auctt.: Leenhouts, Fl. Males. I, 6 (1962) 303 pro parte; Cockburn, Trees

of Sabah | (1976) 210 pro parte; Ashton, Manual Non-Dipt Trees Sarawak 2 (1988)

314 pro parte, Anderson, Check List Trees Sarawak (1980) 239 pro parte. [Fagraea

elliptica Roxb. sensu stricto, Fl. Ind. ed. Wall. 2 (1824) 32, nom. dub.]

Tree, to around 5 m tall: trunk to c. 25 cm diameter; bark fissured, dark brown. Leaves

elliptic, oblanceolate-obovate; 4-15 cm long, 2—7 cm wide; base cuneate; apex acute;

margin plane to slightly recurved; thick-coriaceous; upper and lower surfaces smooth

to finely shagreen; midrib prominent below, ridged; secondary veins 7—9 pairs, upper

side distinct, lower side obscure; tertiary veins obscure; petioles 1.5—2 cm long, stout.

Inflorescence sessile, the basal branches not clearly elevated from the distalmost leaf

pair on the flowering shoot and appearing as if they originate from the ultimate leaf

axils, 6-9 cm long, 11—18 cm across, main axis branched to 4-5 orders, with 5—6

tiers of branches; lowest branches typically 2.5—7 cm long. Flower pedicel (above

the ultimate bracteole) 0.5—1 mm long; calyx (from the base to the lobe apices) 2—3

mm long, calyx cup 1.5—2 mm wide; corolla salverform; cream to white; corolla tube

4-6 mm long, not conspicuously flared, 1—1.5 mm wide, inside glabrous to minutely

papillate; corolla lobes ovate to lanceolate, 2.5—3 mm long, 1—1.5 mm wide; stamens

with filaments 8—11 mm long in the open flower; anthers oblong, c. | mm long; style

9.5—12 mm long in the open flower; stigma knob-like, c. 0.5 mm across, lobes 2,

inconspicuous. Fruit when mature to 3.5—5 mm across; fruit calyx lobes c. 1 mm long

and wide. Seeds c. | mm across.

Distribution. So far known only in the NW Borneo area (Sabah and Sarawak).

Habitat. In sub-montane to lower montane forest.

Specimens examined: BORNEO. Sabah. Ranau, Mamut, copper mine, Beaman 9962 (K,

MSC), copper mining area, Aban SAN 50747 (holo SAN; iso A), Kinabalu N.P., Bundu Tuhan

view trail, Aban SAN 49430 (SAN), Mt Kinabalu, Puasa 3506 (K). Sarawak. Bt. Sadok, base

camp, Banyeng & Ilias S. 45043 (K, KEP, L. MO, SAN, SAR), summit, Banyeng & Ilias S.

45059 (K, KEP, L, MO, SAN, SAR): Anap, (Kana) trig point, Bt. Naoung, Banyeng S. 19401

(A, BO, K, KEP, L, MEL, SAN, SAR, SING); Bintulu, Merurong plateau, Brunig S. 8791 (K,

SAR); Kuching, Matang, Kubah N.P., Gunung Serapi, Lee S. 54146 (K, SAR), 755 m, Rantai

S. 74271 (SAR, SING), Mt Mike, Low LYW 260 (KLU); Mt. Dulit, Richards 1735 (K), Ulu

Koyan, Synge 1874 (K).

3. Picrophloeus javanensis Blume, Bijdr. Fl. Ned. Ind. (1826) 1020; Don, Gard. Dict.

4 (1837) 66; DC., Prod. 9 (1845) 32. — Fagraea picrophloea Blume, Rumphia 2

(1838) 36, nom. illeg., Mus. Bot. 1 (1850) 173; Migq., Fl. Ind. Bat. 2 (1857) 377; Merr.,

J. Str. Br. Roy. As. Soc. Sp. No. (1921) 492. LECTOTYPE (Wong & Sugumaran

2012): Blume, s.n., Java (L: 908.127-201, barcode L0005006).

516 Gard. Bull. Singapore 64(2) 2012

Cyrtophyllum speciosum Bl., Bijdr. (1826) 1022; DC., Prod. 9 (1845) 31; Ridl., FI.

Malay. Pen. 5 (1925) 322. — Fagraea speciosa (Bl.) Bl., Rumphia 2 (1838) 35, t. 81;

Mus. Bot. 1 (1850) 172; Bentham, J. Linn. Soc. Bot. 1 (1857) 100; Merr., J. Str. Br.

Roy. As. Soc. Sp. No. (1921) 493; non sensu Ridl., J. Str. Br. Roy. As. Soc. 50 (1908)

122 (= C. fragrans). TYPE: Blume 1867, Java (iso L: sheet no. 908.127-210).

Cyrtophyllum speciosum Ridl. var montanum Ridl., Fl. Malay. Pen. 5 (1925) 322.

TYPE: Henderson SFN 10992, Pahang, Cameron Highlands, Gunung Terbakar, 45000

ft. alt., 9 vi 1923 (holo K; iso SING).

Fagraea kimangu Blume, Mus. Bot. 1 (1850) 173. TYPE : Blume, Java, kimangle,

kimangu (holo L: sheet no. 908.127-507).

F: valida Migq., F\. Ind. Bat. 2 (1857) 376. TYPE: Junghunh, Sumatra, “Panoadjih et

- Koeta tinggi: (holo L: sheet no. 908.127-625).

F. sumatrana Miq., Fl. Ind. Bat. 2 (1857) 377. TYPE: Teysmann HB 992, Sumatra,

Payakombo, “kajoe sobo” (holo U: barcode U0003707; iso L: sheet no. 908.127-628).

F. aurantiodora S. Moore, J. Bot. 66 (1928) 105. TYPE: Brass 642, Papua New

Guinea, Sogere (holo BM, iso K).

F-, pseudoelliptica Kanehiro & Hatusima, Bot. Mag. Tokyo 56 (1942) 161, f. 5. TYPE:

Kanehira-Hatusima 12577, New Guinea, Ayerjat, along Boemi River, about 40 km

inland from Geelvink (holo TKU; iso A).

F- pusilliflora Bakh.f. in Backer, Bekn. Fl. Java 7 (1948) fam. 170, p. 13, nom. nud.,

Blumea 6 (1950) 383. TYPE: Endert 77E.1P.746, Sumatra, Palembang (holo L: sheet

no. 923.157-1391).

Fagraea elliptica auctt.: Miquel, FI. Ind. Bat. 2 (1856) 376; Koorders & Valeton, Bijdr.

9 (1903) 84; Cammerloher, Bull. Jard. Bot. Btzg. III, 5 (1923) 316; Leenhouts, FI.

Males. I, 6 (1962) 303; Back. & Bakh. f., Fl. Java 2 (1965) 211; Kochummen, Tree FI.

Malaya 2 (1973) 272; Ashton, Manual Non-Dipt. Trees Sarawak 2 (1988) 314; Wong

& Sugau, Sandakania 8 (1996) 21; guoad F. speciosa Blume, Cyrtophyllum speciosum

Blume, F) picrophloeus Blume & Picrophloeus javanensis Blume in syn.; Anderson,

Check List Trees Sarawak (1980) 239 pro parte.

[Fagraea elliptica Roxb. sensu stricto, Fl. Ind. ed. Wall. 2 (1824) 32, FI. Ind. ed.

Carey 1 (1832) 462, DC., Prodr. 9 (1845) 30, nom. dub.; Willughbeia elliptica (Roxb.)

Spreng., Syst. Veg. 4 (1827) Cur. Post. 71, nom. dub. |

Tree, sometimes to 3-4 m tall but more often bigger, to 10-20 m tall; trunk to about

1 m diameter; bark fissured in younger trees, less conspicuously so and dippled-scaly

Picrophloeus revised s17,

in older trees, grey-brown to dark brown. Leaves elliptic: (8—)11—17(-21) em long,

(3—)4-7(-9.5) cm wide; base cuneate; apex acuminate to short-caudate, 1-3 mm

long; margin plane when fresh (in dried specimens sometimes slightly wavy); thin-

coriaceous; upper and lower surfaces smooth; midrib prominent below: secondary

veins (5—)7—12 pairs, upper side faint and immersed in the blade, lower side faint to

very slightly prominent; tertiary veins inconspicuous; petioles (0.5—)1—1.5(—2.2) em

long, stout. Inflorescence sessile, the basal branches not clearly elevated from the

distalmost leaf pair on the flowering shoot and appearing as if they originate from the

ultimate leaf axils, (6—)7—10(—12.5) long, 10-20 cm across, main axis branched to

4-5 orders, with 4—5 tiers of branches: lowest branches (3.5—)4-10 cm long. Flower

pedicel (above the ultimate bracteole) 1-2 mm long: calyx (from the base to the lobe

apices) 1.5—2 mm long, calyx cup 1.5—2 mm wide; corolla salverform; cream to white;

corolla tube 4—6 mm long, not conspicuously flared, 0.8—1 mm wide, inside glabrous

to minutely papillate; corolla lobes 2.5—3.5(-4) mm long, 0.8-1 mm wide: stamens

with filaments 7-8 mm long in the open flower; anthers oblong, 1—1.2 mm long; style

7—14 mm long in the open flower; stigma knob-like, c. 0.5 mm across, lobes 2, low and

rounded, parting to present two slightly raised, hemispherical, papillate inner surfaces

when receptive, not recurving. Fruit when mature to 4-5 mm across; fruit calyx lobes

up to c. | mm long and wide. Seeds 0.5—1 mm across.

Distribution. Sumatra, Java, Malay Peninsula, Borneo, Moluccas, New Guinea.

Habitat. Lower montane forest, but in the Wallacea region also in the lowlands,

including coastal forest.

Specimens examined: BORNEO. Kalimantan. Sampit, Kostermans 4663 (SING). Sabah.

British North Borneo, Villamil 251 (SING); Mamut Coppermine area, Postar et al. SAN 147993

(SAN): Mt. Kinabalu. Eastern Shoulder, Chew, Corner & Stainton 3 (SING).

JAVA. Blume s.n. (lecto L: 908.127-201, barcode L0005006), Blume s.n. (L: 908.127-507),

Blume 1867 (L: 908.127-210, barcode L0005005). G. Salak. 600-1000 m, Koorders 242476

(BO), 244616 (BO), 366898 (BO). Preanger. Tjibodas, 1400 m, Koorders 428256 (BO),

401546 (BO).

MOLUCCAS. Amboina. Robinson 2037 (BO, SING); Boeroe, 300 m, Neth. Ind. For Service

bb 22839 (SING). Ceram. 40 km E of Masohi in Wae Ruatan / Wae Ruwata (Ruwa) catchment

areas, Burley, Tukirin & Ismail 4335 (A, BO, K, SING); Kp. Kiandarat, G. Kilia, Buwalda

5592 (SING). Halmahera. Tasoa-Gunung Sembilan, 300 m, Pleyte 264 (SING). Morotai.

Mt. Songawo, Main et Aden 1025 (SING). Sulabes (Sanana). Kabauw, 150 m, Neth. Ind. For

Service bb 28875 (SING).

NEW GUINEA. Ayerjat. Along Boemi River, about 40 km inland from Geelvink, Kanehira-

Hatusima 12577 (A, TKU). Mimika Regency. PT-Freeport Indonesia Concession Area,

Utteridge et. al. 458 (SING). Morobe. Markham Point, Womersley & Henty NGF 11681

(SING); Patep, 2300 ft, Millar NGF 9970 (SING). Milne Bay. Rabaraba, junction of Mayu &

Ugat River, Katik, LAE 56309 (SING). Sepik. Aitape Subdistrict, along Bliri River, Darbyshire

& Hoogland 8338 (SING); West Sepik District, Telefomin Subdistrict, Prospect Creek near

Freida River, Henty & Foreman NGF 42535 (SING), NGF 42580 (SING). Sogere. Brass 642

(BM, K). Wissel Lake Region. Eyma 4425 (SING).

518 Gard. Bull. Singapore 64(2) 2012

PENINSULAR MALAYSIA. Kedah. Kuala Muda, Gunung Jerai, Whitmore FRI 20443

(SING); Jerai F.R., Abdullah & Motan 73515 (KEP), Gurun 66405, leafy branch only (KEP),

plantation Jerai, Gurun 59647 (KEP, SING); Kedah Peak, Kochummen KEP 94404 (KEP).

Kelantan. Sungai Keleh, Henderson 19658 (KEP, SING). Pahang. Cameron Highlands, Boh

Plantations, Mohd Nur SFN 32626 (KEP, SING), Gunung Brinchang, Jeo et al., KL 4694

(KEP), Gunung Terbakar, Henderson SFN 10992 (K, SING), SFN 31385 (KEP), Holttum SFN

31385 (SING), Symington KEP 36219 (KEP, SING); Fraser’s Hill, Kiew & Anthony RK 3483

(SING), Kenari Trail, Kiew 3335 (KEP); Gunung Tapis, Chan YC FRI 19906 (KEP, SING):

Ulu Kah, Low LYW 358 (KLU), Stone 15420, leafy branch only (KLU).

SUMATRA. Achmad 58 (SING), 1515 (SING), 16/76 (SING), Batten s.n. 1940 (SING), Forbes

3142 (SING); Yates 2201 (SING); Habinsaran oso. vom Tobasee 1200-1300 m, Lérzing

6568 (BO). Asahan. Loemban Ria, Rahmat 7352 (SING), 7813 (SING), 7952 (SING), 7962

(SING). Palembang. Endert 77E.1P.746 (L); Lematang Ilir, Semangoes, Neth. Ind. For Service

bb. 31728 (SING), bb. 31747 (SING), bb. 32122 (SING); Tjaban F.R., near Muara Enim,

Kostermans 12049 (SING). Panoadjih & Koeta Tinggi. Junghunh s.n. (L). Payakombo.

Teysmann HB 992 (L, U). Si borong borong. Stein Ba/9/73/7 (SING).

Fagraea elliptica Roxb. 1s a dubious name that has often been applied to the taxon

referred to here. This was a name first used in the Hortus Bengalensis of Roxburgh

(1814) without any description (nomen nudum), and later only briefly described in

Roxburgh (1824) (Flora Indica ed. Wall. 2: 32), highlighting the “terminal corymbs”

in this species, in contrast to much more detailed descriptions for other species he

described in Fagraea in the same work. He also did not definitely state a locality,

merely mentioning “native of the Moluccas”.

Roxburgh’s unusually brief description for his F. e/liptica may imply that

there were no specimens immediately available to him and that he was awaiting

material from the Moluccas collected either by his son (also William Roxburgh) or by

Christopher Smith, a less-than-fully compliant nurseryman under Roxburgh Senior’s

direction but who seemed interested in succeeding him at the Calcutta Botanic Garden

(Royal Botanic Gardens Kew 2006, Steenis-Kruseman 1958), or even someone else.

The sheets in Kew, where much of Roxburgh’s collection is kept, do not include F-

elliptica collections made from the Moluccas earlier than 1824 which Roxburgh

(who died in 1815) or Carey (who edited the first publication of the Flora Indica,

with two volumes published in 1820 and 1824) or Wallich (whose taxa included in

these first volumes were clearly marked as his additions) could have seen. Neither

are there any found in the Edinburgh, Geneva, Linnean Society of London, Natural

History Museum (London), and Paris herbaria where also Roxburgh material may be

found (Forman 1997). Smith collected in the Moluccas between 1796 and 1805 (John

Hunnex, Natural History Museum (London), pers. comm.) but we have managed to

determine that these collections do not include F’ e/liptica.

There is also no drawing of this taxon among Roxburgh’s Flora Indica

illustrations at Kew (Sealy 1957). The British Museum has a number of drawings

that had belonged to Patrick Russell (Roxburgh’s friend), and John Fleming (one

of Roxburgh’s collaborating botanists), but none of these are marked by Roxburgh

numbers (Royal Botanic Gardens Kew 2006). At the Library of the Natural History

Picrophloeus revised 519

Museum (London), John Hunnex has managed to locate a painting (in a series of some

180 drawings of plants from the region made by Smith before his death in 1807) that

greatly resembled specimens which had been determined as F) e/liptica. Although this

plant is depicted with the same leafy branch habit, terminal inflorescence and floral and

other dimensions that suitably represented such F e/liptica specimens, the included

drawing of a dissected mature corolla tube clearly displays tiny stamens with short

anthers inserted on the inner surface of the tube and completely included within. This

totally contrasts with F. e/liptica specimens available generally that had conspicuously

long-exserted stamens far exceeding the corolla tube. Was this perhaps a preliminary

representation of what Roxburgh had wanted to include but did not do so satisfactorily

because details such as the nature of the stamens were still to be confirmed? The

rather bright red colour used for the anthers in this painting is unrepresentative for this

alliance of plants (in which anthers were typically yellow turning brown, without a red

phase), suggesting it may have needed to be specially highlighted.

The description of F: el/liptica in Roxburgh (1824) is conspicuous by its

brevity, hardly three lines in all, omitting mention of the stamens, and not discussing

any collection, merely mentioning the Moluccas as a provenance. In contrast, the

description of the preceding taxon in the same work, F. fragrans, is detailed, including

mentioning the exserted stamens, and gives the provenance and a collection number

clearly. The later Carey edition of the Flora Indica (Roxburgh 1832, vol. 1: 462),

repeats the same brevity for F. e/liptica. In his Prodromus, De Candolle (1845: 30

& 32) enumerates both F: elliptica Roxb. as well as Picrophloeus javanensis Blume,

which he was evidently unaware could be synonymous. Candolle notes thus on his

page 30:

“13. F. elliptica (Roxb. fl. ind. ed. 1824. v.2 p.32), caule...., foliis breviter petiolatis

lato-ellipticis laevibus firmis, corymbis terminalibus supra decompositis, corollae tubo

cylindrico. in Moluccis. Caet. ign.”

The final notation (Latin: Caetera ignota = all the rest unknown) is a telling sign that,

decades afterwards, the details of F. elliptica had remained enigmatic. By the time of

the Flora of British India (Clarke in Hooker 1883), this taxon was not enumerated

anymore. However, Miquel (1856) still adopted the name F: e/liptica, with F. speciosa

Blume (published in 1838) as a synonym, and also enumerated the now-illegitimate F:

picrophloeus Blume (based on Picrophloeus javanensis), providing these names that

Malesian botanists then attempted to account for in their region. These names were

later accepted by Leenhouts (1962) as synonymous, with F: e/liptica Roxb. the earliest

correct name.

The unsuccessful searches among the various herbaria for specimen material

that could be specially related to the brief and uninformative description of F: elliptica

in Roxburgh (1924), the continuing inability among interested scholars to provide a

better description in the decades following this, and the existence of a Smith painting

wherein an important discrepancy exists for the very feature that was omitted in the

Roxburgh descriptions, must, unfortunately, imply that Roxburgh and his editors were

greatly hampered by availability of material and lack of familiarity with this taxon.

Here we take the view that F. elliptica Roxb. represents a dubious name that cannot be

520 Gard. Bull. Singapore 64(2) 2012

reasonably ascribed a type specimen from possibly original material. Instead, there is

much advantage in employing P. javanensis Blume as the correct name for the taxon

we intend here, because it is also assigned to the genus name most appropriate to its

taxonomy and was validly published in 1826 (just two years following the appearance

of F. elliptica in Roxburgh’s Flora Indica), when for years later even Roxburghian

scholarly circles could not resolve the enigmatic aspects of F elliptica. Even if

familiarity could be claimed for using F- e/liptica Roxb. in the sense of P. javanensis,

the former name would still need to be neotypified and a new name combination made

in Picrophloeus, both of which could be highly contentious propositions.

4. Picrophloeus rugulosus (K.M.Wong & Sugau) K.M.Wong, comb. nov. — Fagraea

rugulosa K.M.Wong & Sugau, Sandakania 8 (1996) 22. TYPE: Chai & Ilias §. 27929,

Sarawak, 5th Division, Ulu Lawas, Telau, Kota F.R. near Sg. Telau (holo SAN; iso A,

BO, K, KEP, L, SAR, SING).

F. elliptica auctt.: Leenhouts, Fl. Males. I, 6 (1962) 303 pro parte; Cockburn, Trees

of Sabah 1 (1976) 210 pro parte; Ashton, Manual Non-Dipt Trees Sarawak 2 (1988)

314 pro parte; Anderson, Check List Trees Sarawak (1980) 239 pro parte. [Fagraea

elliptica Roxb. sensu stricto, Fl. Ind. ed. Wall. 2 (1824) 32, nom. dub.]

Tree, to around 15 m tall; trunk to c. 25 cm diameter; bark lightly fissured, dark brown.

Leaves elliptic-obovate; 11—21 cm long, 5—9 cm wide; base cuneate; apex acute; margin

recurved when dry; coriaceous; upper and lower surfaces coarsely shagreen; midrib

prominent below, rounded or ridged; secondary veins 9—12 pairs, on both sides faint to

obscure; tertiary veins obscure; petioles 2—3.5 cm long, stout. Inflorescence distinctly

pedunculate (its basal branches distinctly elevated from the ultimate leaf pair), to c.

6 cm long, c. 15 cm across, main axis branched to 3-4 orders, with (3—)4—S tiers of

branches; peduncle 1—1.5 cm long; lowest branches typically 2.5—-6 cm long. Flowers

sessile (the ultimate floral bracteole immediately below the calyx cup); calyx (from the

base to the lobe apices) 3-4 mm long, calyx cup 2-3 mm wide; corolla salverform;

cream to white; corolla tube 8—10(—11) mm long, not conspicuously flared, 1—1.5 mm

wide, inside glabrous to minutely papillate; corolla lobes 4.5—-6 mm long, 2—2.5 mm

wide; stamens with filaments 15—18 mm long in the open flower; anthers oblong, c. 1

mm long; style (18—)22—25 mm long in the open flower; stigma knob-like, c. 0.5 mm

across, lobes 2, inconspicuous. Fruit when mature to 6—9(—10) mm across; fruit calyx

lobes 2—2.5 mm long and wide. Seeds c. | mm across.

Distribution. So far known only in, and probably endemic to, the Brunei region (SW

Sabah, Brunei and NE Sarawak).

Habitat. Primary kerangas and lowland mixed dipterocarp forest up to c. 450 m.

Specimens examined: BORNEO. Brunei. Belait, Andulau F.R., Ashton BRUN 628 (BRUN, K,

—

Picrophloeus revised 5

SING), Andulau Compartment 8, Wood, Smythies & Ashton SAN 17521 (BRUN, K, SING);

Belait, Bukit Puan, Ashton BRUN 637 (BRUN, K, SING), Sukang, Sungei Kuat, Ulu Belait,

Salleh et al. BRUN 15338 (SING), Sungai Liang, at Ulu Sungai Lumut, Niga et al. BRUN 15216

(SING); Belait, Teraja, Wasai Wong Kadir, Ariffin et al. BRUN 15161 (SING); Belait, source of

the Sg. Ingei north of Batu Patam, Wong WKM 1/104 (BRUN, K, SING); Ulu Belait, R. Topi,

Ashton BRUN 214 (BRUN, K, SING); Temburong, Temburong River, just upstream from

Wong Nguan gorge, Wong WKM 1702 (BRUN, K, SING), just upstream from Wong Nguan

rapids, Coode 6539 (BRUN, K); Tutong, Bt. Bahak, near LP 338A, Coode 7029 (BRUN, K):

Tutong, Kampong Bukit Beruang, Niga & Clayton BRUN 15450 (SING); Tutong, Ladan Hill

F.R., Ariffin et al. BRUN 18078 (SING). Sabah. Beaufort, Papar-Beaufort road, mile 16.5,

Lajangah SAN 32270 (SAN); Sipitang, Sri Beaufort Area, Dewol SAN 80111 (KEP, SAN, SAR,

SING). Sarawak. Baram, G. Mulu N.P., Ulu Sg. Berar, Chai S. 39595 (A, KEP, L, MO, SAN,

SAR), Melinau Gorge (4°10°N, 114°55’E), 1000 ft, Chew CWL 442 (K, SING), Ilias S. 16603

(A, BO, K, L, SAN, SAR, SING), near Gunong Api (4°7°N, 114°55’E), 1000 ft, Chew CWL

1172 (K, SING); Miri, Lambir, proposed N.P., Awang Morshidi S. 24094 (K, L, SAR, SING),

Lambir Hills, //ias S. 16603 (SAR, SING); Ulu Lawas, Kota F.R., near Sungai Telau, Chai &

Ilias S. 27929 (holo SAN; iso A, BO, K, KEP, L, SAR, SING).

ACKNOWLEDGEMENTS. The present revision is based on further work subsequent to the

collaboration with M. Sugumaran in unravelling the Fagraea complex. | thank the Keepers

and Curators of the A, BO, K, KEP, KLU, L, NY, US, SAN, SAR and SING herbaria for

permission to consult reference materials and loans of specimens. J.J. Wood, J. Dransfield,

N. Martland (Kew), E.F. de Vogel, C.E. Ridsdale, J.-F. Veldkamp, L. Willemse (Leiden), H.

Noltie (Edinburgh), C.C. Davis, D. Boufford, W. Zhang, K. Gandhi (Harvard), L.G. Saw,

R.C.K. Chung (Kepong), J.B. Sugau, J.T. Pereira, M. Postar (Sandakan), C.L. Chan (Kota

Kinabalu), A.A. Joffre, K. Ariffin (Brunei), R. Kiew (formerly SING), S. Lee, C.J.H. Soh and

M.I. Bazilah (SING) are thanked for assisting with a whole range of research matters. An

introduction to Roxburgh’s importance by the late L.L. Forman (Kew) has enlightened. John

Hunnex, Natural History Museum (London), gave much assistance with the search for material

related to Roxburgh and Smith. Y.W. Low (Singapore) kindly assisted with location of type

images and information.

REFERENCES

Blume, C.L. (1838) Rumphia. Vol. 2. Leiden.

Candolle, A.P. de (1845) Prodromus systematis naturalis regni vegetabilis, sive enumeratio

contracta ordinum generum specierumque plantarum huc usque cognitarium, Juxta

methodi naturalis, normas digesta. Vol. 9. Paris: Sumptibus Sociorum Treuttel et Wirtz.

Forman, L.L. (1997) Notes concerning the typification of names of William Roxburgh

phanerogams. Kew Bull. 52: 523-534.

Hooker, J.D. (1883) The Flora of British India. Vol. 4. Pp. 82-85. London: L. Reeve & Co.

Leenhouts, P.W. (1962) Loganiaceae. In: Steenis, C.G.G.J. van (ed.), Flora Malesiana. Series

1 Vol. 6. Pp. 293-336. Groningen: Wolters-Noordhoff Publishing.

Miquel, F.A.W. (1856) Flora Indiae Batavae. Vol. 2. Lipsiae: Fried. Fleischer.

Ridley, H.N. (1893) On the flora of the eastern coast of the Malay Peninsula. Zrans. Linn. Soc.

London, Bot. 3: 267-408 & PI. 61-66.

522 Gard. Bull. Singapore 64(2) 2012

Ridley, H.N. (1918) New and rare Malayan plants. J. Straits Branch Roy. Asiat. Soc. 79: 97-98.

Ridley, H.N. (1923) The Flora of the Malay Peninsula. Vol. 2. London: L. Reeve & Co.

Roxburgh, W. (1814) Hortus Bengalensis, or a Catalogue of the Plants Growing in the

Honourable East India Companys Botanical Garden at Calcutta. Serampore, Calcutta.

Roxburgh, W. (1824) Flora indica, or, Descriptions of Indian plants, by the late William

Roxburgh. Edited by Dr. William Carey; to which are added descriptions of plants more

recently discovered by Dr. Nathaniel Wallich. Vol. 2. Serampore: Mission Press.

Roxburgh, W. (1932) Flora indica, or, Descriptions of Indian plants /by the late William

Roxburgh. Ed. W. Carey. Vol. 1. Serampore: W. Thacker.

Royal Botanic Gardens, Kew (2006) Roxburgh’s Flora Indica. http://www.kew.org/floralndica/

(accessed on 13 Jul 2012).

Sealy, J.R. (1957) The Roxburgh Flora Indica drawings at Kew. Kew Bull. 11: 297-399.

Steenis-Kruseman, M.J. van (1958) Malaysian plant collectors and collections. Supplement I.

Flora Malesiana |, 5 (4): CCXXXV—CCCXLII (superposed pagination 1—108).

Sugumaran, M. & Wong, K.M. (2012) Studies in Malesian Gentianaceae I: Fagraea sensu

lato—complex genus or several genera? A molecular phylogenetic study. Gard. Bull.

Singapore 64(2): 301-332.

Wong, K.M. & Sugau, J.B. (1996) A revision of Fagraea (Loganiaceae) in Borneo, with notes

on related Malesian species and 21 new species. Sandakania 8: 1-93.

Wong, K.M. & Sugumaran, M. (2012) Studies in Malesian Gentianaceae II: A taxonomic

framework for the Fagraea complex, including the new genus Limahlania. Gard. Bull.

Singapore 64(2): 481—495.

Gardens’ Bulletin Singapore 64(2): 523-527. 2012

im)

1oS)

Studies on Homalomeneae (Araceae) of Peninsular

Malaysia IV:

Homalomena stongensis, a remarkable new species

endemic to Gunung Stong, Kelantan

Zulhazman Hamzah', Peter C. Boyce’ and Mashhor Mansor’

'Faculty of Earth Science, Universiti Malaysia Kelantan,

Jeli Campus, Locked Bag 100,

17600 Jeli, Kelantan, Malaysia

*School of Biological Sciences, Universiti Sains Malaysia,

11800 USM, Pulau Pinang, Malaysia

*‘phymatarum@googlemail.com

ABSTRACT. Homalomena stongensis is described from Gunung Stong, Kelantan, where it is

very locally endemic to steep forested slopes. An updated key to Peninsular Malaysian species

of Homalomena Supergroup Homalomena is provided, and the new species is illustrated.

Keywords. Araceae, Homalomena, Homalomena Supergroup, Peninsular Malaysia, Kelantan,

limestone

Introduction

Fieldwork focussing on Araceae in Peninsular Malaysia continues to reveal novel

species, many seemingly very locally endemic, maybe explaining why they have been

overlooked to date. In particular, Homalomena appears to be represented by a great

many novel taxa in the Peninsula (Ng et al. 201 1a; Zulhazman etal. 2011). Frustratingly,

most are encountered sterile during fieldwork, necessitating samples being bought

into cultivation to flower and enable assessment of their novelty. Recently, one such

taxon gathered during an initial survey at Gunong Stong, Kelantan, has flowered in

the research collection of the first author, and has proved to be both novel and rather

remarkable in its morphology. It is here described.

Homalomena stongensis H.Zulhazman, P.C.Boyce & M.Mashhor, sp. nov.

Species statim distinctissima pedunculo patente declinato spatha ascendenti florens.

Praeterea spatha concolor ad anthesin externe medie virida interne clare luteole alba,

staminodia interpistillaria filamentose clavata, spadix longe stipitatus, longitudo

zonarum pistillatarum cum stipite circa dimidium spadicis longitudinem distinguenda.

TYPE: Malaysia, Kelantan, Jeli, Gunong Stong State Forest Park, forested flanks of

524 Gard. Bull. Singapore 64(2) 2012

Jelawang Falls, 18 April 2010, 5°20°29.0”°N; 101°58710.1°E, Zu/hazman UMK7, (holo,

Herbarium Universiti Malaysia Kelantan, Faculty of Earth Science, UMK). (Fig. 1)

Medium, solitary, evergreen, glabrous, bruised tissue strongly aromatic (reminiscent

of young mango), herbs to 52 cm tall, shoots pleionanthic. Stem up to 26.5 cm tall,

c. 4 cm diam., erect, rooting from nodes and at length declinate, older parts clothed

with brownish-black papery to fibrous petiole base remains, with the roots penetrating

this fibrous layer, active tip erect with oldest leaves spreading. Leaves 6—7 together,

spirally arranged, and clustered towards shoot tips, older leaves by bending of the

petiole with the leaf blade tip touching the ground; petio/es exceeding the blade, 34-42

cm long, non-sheathing portion D-shaped in cross section, 0.9-1.1 cm diam., above

petiolar sheath, base clasping the stem and expanding into an open persistent petiolar

sheath, medium green, petiolar sheath |1—18 cm, '/;—'/ petiole length, persistent, deep

spreading and open, margins hyaline, petiolar sheath for the older leaves, externally

shallowly grooved, the groove extending to the insertion of the blade; leaf blade ovate-

cordate to ovate-lanceolate, 25-30 cm x 18—22 cm, base sagittate to cordate, apex

acute with an apiculate tip 6-7 mm long, semi-glossy, older leaves irregularly quilted

between the primary lateral veins and these leaves, through recurving of the petioles

often with the leaf blade tip touching soil, margins entire and minutely hyaline,

adaxially semi-glossy dark green, adaxially paler green; mid-rib less conspicuous

adaxially, prominent abaxially, wedged shape at mid-vein cross section and primary

lateral veins conspicuously raised abaxially, deeply impressed adaxially; primary

lateral veins 11—1I5 per side, interprimary veins alternating, much less prominent;

secondary and tertiary venation invisible. Inflorescences 3—S together, maturing

sequentially in a simple synflorescence, initially upright, at anthesis with peduncle

spreading-declinate and spathe ascending, fully all declinate after anthesis; peduncle

9-12 cm long, 8.3—9.5 mm diam., at spathe insertion 9-10 mm, dark, somewhat glossy

green. Spathe unconstricted, ellipsoid-cylindrical, 10.4—10.7 cm long, 0.83—0.95 cm

diam. (before anthesis), gaping c. 1.4 cm at anthesis to c. 1.8 cm wide with the rostrum

remaining furled and retaining the tip of spadix, with a conspicuous terminal curved

rostrum to c. 1.5 cm. Spadix c. 7 x 1 cm, obliquely stipitate; stipe 0.9 cm, greenish

white. Pistillate flower zone cylindrical, c. 3 x 1 cm, pistils somewhat laxly arranged,

2.5 x 1.5-2 mm: ovary globose-cylindric, the lowermost ones somewhat obliquely

gibbose, ascending, glossy medium green; stigma sessile, discoid, flat, translucent

white, c. 2 mm diam.; interpistillar staminodes c. 2.8 mm long, exceeding pistils,

stipe very slender, flexuous, tip tear-drop shaped, white, staminodes associated with

distal-most pistils weakly trapezodial in plan view. Staminate flower zone c. 3.9 *

1.1 cm, cylindrical, the tip bluntly weakly tapering, very pale whitish yellow, almost

contiguous with the pistillate zone, the interface marked by a few somewhat laxly

arranged trapezoidal staminodes; staminate flowers weakly and irregularly rhombo-

hexagonal, each with 3—5 stamens, 1.8 * | x 1.5—2 mm, anthers with 4 distinct thecae,

each with a terminal crenate lobe, and overtopped by a large synconnective. Fruiting

spathe declinate. Fruits & seeds not observed.

A new Homalomena from Peninsular Malaysia 525

Fig. 1. Homalomena stongensis H.Zulhazman, P.C.Boyce & M.Mashhor. A. & D. Flowering

plant. Left hand inflorescence at female anthesis; note the spreading-declinate peduncle and

erect spathe; right hand inflorescences at early stages of fruiting, note the declinate peduncle

and spathe beginning to decline. B. Detail of the open petiolar sheath. C. Whole plant. E.

Leaf blade, adaxial view. Note the quilted nature of the blade. F. Inflorescence at pistillate

anthesis. Note the yellowish tinge to the spathe interior, and the manner in which the rostrum

remains closed and retains the tip of the spadix. G. Spadix, with spathe removed. H. Pistillate

flower zone. Note the flexuous interpistillar staminodes. I. Staminate flower zone. (Photos:

Zulhazman H.)

526 Gard. Bull. Singapore 64(2) 2012

Distribution. Peninsular Malaysia, Kelantan, Gunong Stong State Forest Park. Locally

endemic.

Ecology. Perhumid hill dipterocarp forest along flanks of waterfalls on steep (c. 30°)

granite slopes, at 300 m asl.

Notes. Homalomena_ stongensis belongs to the Homalomena Supergroup sensu

Wong & Boyce (2008) & Ng et al. (2011b). Homalomena stongensis is immediately

distinctive from all other Homalomena so far described from Peninsular Malaysia

by flowering with the peduncle spreading-declinate and the spathe ascending. It is

further distinguished by the strongly discolorous, spathe - is externally medium green,

internally bright yellowish white at anthesis, the filamentous-clavate interpistillar

staminodes, long-stipitate spadix, and the combined length of pistillate zone plus stipe

accounting for almost half the spadix length.

Key to Peninsular Malaysian Homalomena

(Supergroup Homalomena)

la. Peduncle at anthesis spreading-declinate with the spathe erect. Interpistillar

staminodes with stipe very slender, filamentous, somewhat sinuous, the head

teardrop-Shape \..zcesszict.soacesatacsceaattnes ctocs oasonseene ccna eee eee ae eee H. stongensis

b. Peduncle at anthesis erect. Interpistillar staminodes stout, straight, the head +

red (0) 010) ne oe rE ANE Acid ocoponcedonat SdsecRecinc.ss sone: 2

2a. Flowering plants large, often exceeding | m tall. Spathe 8-14 cm long; spadix

T—-13 eta LOM greece hc coos eee oe eee H. pontederiifolia

b. Flowering plants medium, seldom exceeding 40 cm tall. Spathe 4-6 cm long;

Spadix 5=6.5:crin LONG: sfc. See creer olessnsie ca me eee eRe eee eee)

3a. Leaf blade abaxially with conspicuous striate pellucid secretory canals running

parallel to the primary lateral veins. (S Peninsular Malaysia as far north as S

Pahang). .assnsnaccedes sce sere eerie eee meat ek ereieite eer enon ene ae eee H. curvata

b. Leaf blade abaxially without striate pellucid secretory canals. (Kedah, Perlis) .....

Savoia baa case sses’eaateo¥sacesonie debe dati Salar Came a RRDRae dee Ore hues eee ace cet a fee tte sae ese ame eset H. truncata

ACKNOWLEDGEMENTS. The authors would like to acknowledge the Kelantan Forestry

Department for allowing them to conduct the study in the Gunung Stong State Forest Park.

Special thanks go to Mr. Nik Yuszrin Yusof, Ms. Naziah Zaid, and Ms. Norzielawati Salleh

for their kind assistance in our work. This project was funded by Universiti Malaysia Kelantan

through the first author’s short-term research grant R/SGJP/A03.00/00279A/001/2009/00002 1

via the Faculty of Agro Industry and Natural Resources. Thanks are due to J.F. Veldkamp (L)

for providing the Latin diagnosis.

A new Homalomena from Peninsular Malaysia

References

Boyce, P.C. & Wong, S.Y. (2008) Studies on Homalomeneae (Araceae) of Borneo 1. Four new

species and preliminary thoughts on informal species groups in Sarawak. Gard. Bull.

Singapore. 60: 1-29.

Ng, K.K., Boyce, P.-C. & Sofiman, O. (201la) Studies on Homalomeneae (Araceae) of

Peninsular Malaysia II: An historical and taxonomic review of the genus Homalomena

(excluding Chamaecladon). Gard. Bull. Singapore 62: 277-289.

Ng, K.K., Sofiman O., Boyce, P.C. & Wong S.Y. (2011b) Studies on Homalomeneae (Araceae)

of Borneo VIII: Delimitation of additional informal suprageneric taxa for Sundaic

Homalomena. Webbia 66(1): 21-28.

Zulhazman, H., Mashhor, M. & Boyce, P.C. (2011) Notes on Araceae of Kuala Koh, Kelantan,

Peninsular Malaysia. Gard. Bull. Singapore 63: 213-218.

Gardens’ Bulletin Singapore 64(2): 529-531. 2012 529

BOOK REVIEW: Etlingera of Sulawesi. 4.D. Poulsen. 2012.

Kota Kinabalu: Natural History Publications (Borneo) Sdn. Bhd. in association with Royal

Botanic Garden Edinburgh and Natural History Museum, University of Oslo. 26 cm = 19 cm,

hard cover, vii + 278 pp. ISBN 978-983-812-138-5. Price RM 250.

The ginger genus Et/ingera is well known through the magnificent Torch Ginger

cultivated throughout the tropics and used in SE Asian cooking. But there is more to

Etlingera. The genus is both large and morphologically diverse with the total number of

species still pending, but estimated by the author to be 150-200 species. All Etlingera

species are evergreen and are found mostly in equatorial evergreen forests, growing

from lowlands to high altitudes of 2700 m. The distribution of this genus spans over

14,000 km, with its western limit in Northeast India, richly represented across SE Asia

and reaching its eastern limit in the Pacific island of Tahiti.

It has been 6 years since Axel Poulsen’s previous book Etlingera of Borneo

(2006, same publisher) and it is obvious that the author has been rather busy getting

yet another stunning book out. It is dedicated to the memory of British naturalist

Alfred Russel Wallace (1823-1913), an avid explorer, geographer, anthropologist and

biologist, who is well known for identifying the Wallace line, dividing the Indonesian

archipelago into the ecozones of Asia and Wallacea—a transitional zone between Asia

and Australia.

Sulawesi is located in the western part of Wallacea and therefore it is no big

surprise that its flora is strikingly different from neighbouring Borneo. The flora is

also far less known and this book reflects perfectly the uniqueness and richness of the

Sulawesi flora as well as the poor state of our knowledge. Only 4 Eflingera species

have been known to occur in Sulawesi in 2008, when the author started his work,

but the current revision presents 48 taxa—an increase of more than 10 fold! It is

noteworthy that Borneo which has only 42 taxa is about 4 times bigger than Sulawesi,

and as the author pointed out, he spent far less time in the field in Sulawesi than in

Borneo, hinting that more species are yet to be found in Sulawesi. The only species

common between the two islands is the ubiquitous Torch Ginger (E¢/ingera elatior).

The readers familiar with usually red, orange and yellow Bornean species will be

surprised with an entirely different colour palette of cream, pink and yellow-green

Sulawesi species instead.

The book is divided into three parts, which are clearly marked by coloured

headers. The first part of the book is divided into 5 major chapters. The brief

introduction recapitulates basic facts about the genus E¢/ingera, while the second

chapter, accompanied by 8 dated maps, explains Sulawesi’s complicated geological

history and touches on its climate and geology. The third chapter deals with the

morphology of an Etlingera plant. It describes in detail the characters of vegetative

and flowering shoots, flowers, infructescences, fruits and seeds. The five very detailed

SEM photographs of stigmas could present somewhat amusing and scary images to

the imaginative viewer, while the line-drawing of various fruit types drawn to scale

and spreading over two pages is helpful for identification.

530 Gard. Bull. Singapore 64(2) 2012

The fourth chapter starts with the history of ginger research on Sulawesi,

recapitulating the importance of botanists collecting there, for example, C.G.C.

Reinwardt, O. Beccari, O. Warburg, the brothers P.B. and K.F. Sarasin, but also

mentioning others like F. Gagnepain and K. Schumman, who described species

based on others’ collections. Informal infrageneric groupings, origin and evolution,

the significance of Wallace’s Line and affinities to neighbouring islands, as well as

ecology, pollination, flexistyly, ethnobotany and conservation issues are also discussed

in this chapter.

The last fifth chapter of the introductory part describes material and methods.

The author shares the dire situation of starting a revision, with most of the types in

the Berlin Herbarium destroyed during the Second World War and how the extensive

fieldwork targeting all type localities proved to be a successful strategy, as fertile

material was found for all but one species. The comparison between Sulawesi, where

the majority of the species are known only from single or very few collections, and

Borneo, where multiple collections exists for most of the species, is nicely summarised

in Figure 37. Beware of the printing hobgoblin, who switched the signs Borneo and

Sulawesi in the chart itself, though the careful reader should not be misled as the

legend is well composed. This chapter also discusses the methodology of collecting

notoriously difficult gingers with specific notes on Et/ingera, and explains terminology

used in the descriptions, how the collections were measured, as well as the species

concept used in this revision. The key to the species is the last item of the introductory

part before the real E¢/ingera show starts.

The main bulk of the book is dedicated to the 48 taxa, which are arranged

alphabetically. This includes 36 new species, two subspecies, one new name and six

new combinations for species, which have been previously recognised as members

of other ginger genera, e.g., Amomum. Six new synonyms are also recognised. Each

taxon has been provided with a detailed description, followed by local names and

uses, etymology, ecology and habitat, distribution, conservation status, other material

examined, and finally, other notes. This is supplemented by a nice distribution map,

drawings of the floral parts and numerous colour photographs showing in great detail

the inflorescences, flowers, ligules, fruits and other important characters. In species

with complex nomenclature or taxonomy, such as Eflingera alba, images of the

original herbarium material are also provided with detailed notes on new synonymy

and selection of lectotype. The account closes with a list of specimens, some of which

may still turn out to be new taxa, but at present there is no adequate material for their

description; there is also an accompanying taxonomic note regarding Amomum names

that have been investigated and proven not to be Ef/ingera species (as many elsewhere

have turned out to be the latter genus).

The third and last part of the book is dedicated to acknowledgements and the

useful necessities—teferences, identification list, indexes to vernacular names, uses

and scientific names.

Etlingera of Sulawesi is a modern, critical revision based on the author’s solid

study of herbarium material as well as fieldwork. There is no doubt that the author

loves his work, loves his plants and enjoys writing books. Kudos are also due to the

Book Review 531

publisher for a beautiful layout, which blends well with the scientific text and ink

illustrations, with colour photographs and other pictures of habitats, old maps and

some hitherto unpublished historical photographs. I find this book supremely well

done, enjoyable throughout and easy to use. Considering the minimal overlap with the

previous Ezlingera of Borneo, the two books together cover some 88 species, which is

more than half of the currently known species. And the Etlingeras? Some are simply

stunning, while others are enjoyably strange as demonstrated right on the cover of the

book, ensuring some ‘oohs’ and ‘aahs’ for anyone who picks up this book for serious

professional interest, or just out of curiosity.

Jana Leong-Skorni¢kova

Singapore Botanic Gardens

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Gardens’ Bulletin Singapore 64(2): 533—534. 2012 533

BOOK REVIEW: Flora of Hong Kong. Volume 4. Hong Kong Herbarium & South

China Botanical Garden (General editors: Hu Qi-ming, Wu De-lin; Associate editor:

Xia Nian-he; Volume editors: Xia Nian-he, Cheung Kwok-wai, Pang Kuen-shum &

Yip Kwok-leung) . 2011.

Hong Kong: Hong Kong Herbarium, Agriculture, Fisheries and Conservation Department. 30.6

em * 21.7 cm, hard cover. xv+379 pp. (main text) +65 pp. (Photos 1—673, illustr. glossary).

ISBN 978—988—98253-—9-3. Price HKD 150.

Volume 4 completes the Flora of Hong Kong project, which succeeds two previous floras

from long ago, Bentham’s Flora Hongkongensis (1861) and Dunn & Tutcher’s Flora

of Kwangtung and Hongkong (1912). With around 2100 native species (out of some

3330 total species) in an area of just over 1000 km’, Hong Kong’s flora is justifiably

rich; the territory encompasses elements of both tropical and subtropical regions. China

has about 31,500 plant species (treated in the near-completing Flora of China project

that will be published as 25 volumes of text and 25 volumes of illustrations, which the

present F/ora complements), nearly 12% of the world total, and many plants generally

distributed in South China were first described from Hong Kong.

Volume | included 10 gymnosperm families sensu Kubitzki (1990) and 75 dicot

families; Volume 2, another 56 families; Volume 3, the remaining 29 families; and

Volume 4, 40 monocot families. A Master Index in a separate volume was published

in 2012. The format is highly accessible, with brief family and genus summaries

(contrast these with some frequently lengthy beginning descriptions in the Flora of

China versions), simple indented (typically artificial) keys (the taxa numbered and

presented following the order they are keyed out), species references citing the major

relevant accounts, taxon descriptions of up to 200-300 words, and short notes on

known localities in Hong Kong, distribution elsewhere, ecology and (where relevant)

uses and taxonomy. The Flora of Hong Kong represents a generally up-to-date

documentation of the plant diversity, with few omissions (e.g., Vanilla, Orchidaceae)

that have resulted from somewhat contemporaneous research publications.

Volume 4 itself has 673 colour photos spread over 59 pages following the main

account. The “big ones” in Volume 4 include Cyperaceae (27 genera, 143 spp..

Deng Yun-fei, Carex by Ng Sai-chit); Poaceae (95 genera, 227 spp., Xia, Yu Hui, Li

Jing, Zhao Nan-xian, Peng Hua); and Orchidaceae (53 genera, 114 spp., Hu Shiu-

ying, Corsica Kong). Other contributors include Guo Li-xiu (Arecaceae), Patrick Lai

(Halophila), Richard Saunders (Burmmaniaceae, Philydraceae), Wang Fa-guo, Xing

Fu-wu and Zheng Xi-long (Dioscoreaceae), Yip Kwok-leung (Hydrocharitaceae,

Limnocharitaceae, Najadaceae), Yu Hui (Araceae), and Zhang Dian-xiang

(Burmmaniaceae), and a great assortment of the other families were covered by the

editors Hu, Wu and Xia, individually.

This is the only modern regional South Chinese flora written in English. It is a

major project of the Hong Kong Herbarium, established in 1878 and China’s first public

herbarium. There are triumphs in different senses. Professor Hu Shiu-ying, Honorary

Editor of the Flora of Hong Kong, provided the preface, in which she celebrated the

completion of this flora, one she began studying in 1968. Professor Hu, whose own

534 Gard. Bull. Singapore 64(2) 2012

contributions appear in the “modern flora specially prepared for Hong Kong” that she

had dreamt of for so long, passed away on 22 May 2012, just a year after Volume 4

appeared.

Orders by mail can be made to the Publications Sales Unit, Information Services

Department, Room 402, Murray Building, Garden Rd., Central, Hong Kong, China or

through the puborder@isd.gov.hk email address.

K.M. Wong

Singapore Botanic Gardens

Gardens’ Bulletin Singapore 64(2): 535-536. 2012 535

BOOK REVIEW: Private Lives. An Exposé of Singapore’s Rainforests. Wang

Luan Keng, Darren C.J. Yeo, Kelvin K.P. Lim & Shawn K.¥. Lum (eds). 2012.

Singapore: Raffles Museum of Biodiversity Research. 21.1 cm < 14.9 cm. 298 pp. Hardcover:

ISBN 978-98 1-07-2472-6, price SGD 35. Paperback: ISBN 978-98 1-07-2473-3, price SGD 24.

This book is well worth noticing and owning because it celebrates the riches of the rain

forest by providing, simply, a well-coordinated string of perspectives and anecdotes

that introduce what rain forests are, nicely educating on key aspects of this vegetation

formation and its plant and animal life. Its context is, firstly, to draw attention to

Singapore's last-remaining natural treasures found in this amazingly complex

vegetation type, but it really is a useful book for all of Southeast Asia generally because

the material is applicable across the region.

There have been other attempts to present the rain forest to a general audience,

but this book, crafted by natural history scientists and conservationists working

together, achieves an excellent introduction for both the student as well as the informed

general reader. Its size, too, allows for easy carrying about and storage, a veritable

mini-compendium of tropical rain forests, their ecological significance, incredible

plant and animal diversity, and the importance of further discovery and protection.

The book cleverly infuses people-oriented elements to make its point. For example,

the introduction does not fail to point out how communities are entrenched in botanical

tradition, with roads named after plants, and so on. It is richly illustrated in colour

throughout, which is quite necessary in a book with such a broad mission. There are

just a few corrections, which the authors themselves take the trouble to point out (in

their website http://exxonmobil.rafflesmuseum.net/) but this in no way diminishes the

value of the book.

Logically structured, the Introduction lays out the importance of rain forests in

maintaining genetic and species diversity, and as a key carbon sink, without hesitation.

Its physical structure contributing to 3-dimensional complexity, to which transitional

developmental phases of individual organisms, cohorts and communities give a

temporal variability, is nicely presented. “Biotic Features” points out the dominance

of tree forms and gives a sketch of the scientific ideas around how such levels of

biodiversity are derived, emphasises the dynamic nature of forests through discussing

the forest growth cycle and succession, and leads the reader comfortably through a

slew of anecdotes about rain forest adaptations among plants. There is a highlight

on mass flowering, a key feature of SE Asia’s climax forests. “Moving Up” relates

about climbing plants and epiphytes. Invertebrates get their own chapter. “Winging It”

discusses creatures that fly or glide, as well as the curious adaptations among plants

for wind dispersal; “Forest Buffet” tells of feeding strategies among animals; “Sylvan

Symphony” recollects the chirps, songs, croaks and choruses contributed by a huge

diversity of organisms that make the forest traveller’s experience so fascinating.

The night brings a different world, as if the organisms have worked out a time-

sharing schedule among themselves. The wonder of nocturnally active animals,

the miracle of specialised night vision, and things that glow in the dark capture our

attention. “Poisons, Fangs & Claws” bring on the realism of food capture as well as

536 Gard. Bull. Singapore 64(2) 2012

chemical defences 1n both plants and animals (here the stinkbugs hold their own), and

the weird and wonderful in both plant and animal armature. “Looks Can Be Deceiving”

describes camouflage and mimicry, including the cross-kingdom mimicry among

plants and animals. The story of decomposition is told via focussing on the agents of

decay, and social organisms and ecological interdependence among plants and animals

are dealt with using similarly short “stories”. Among the last chapters are “Jungle

Grocery” that highlights our continued dependence on forest-derived materials and

foods, and “Taking Stock, Taking Care”, before the final discussion on conserving and

managing forests sensibly is presented.

“Eco-Warrior” seems a nice way to complete such an account. It is a salute,

presented as a finale to this book, to Dr. Wee Yeow Chin, retired academic but tireless

teacher and conservationist, whose own accounts of natural history have sustained

interest around nature in Singapore and the region. It reminds us not to be complacent,

because, like all things good and wonderful, rain forests can well disappear if we do

not take the trouble to understand and protect them.

K.M. Wong

Singapore Botanic Gardens

The Gardens’ Bulletin

Singapore

VOL. 64 2012 ISSN 0374-7859

Centenary Issue 2012

VOLUME 64

No.1 pages 1-280 (published 15 July 2012)

No.2 pages 281—533 (published 12 December 2012)

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.

Dr. K.M. Wong

(Editor-in-Chief)

Salece

Dr. Jana Leong-Skorniékova

(Managing Editor)

Christina Soh

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

Editorial Advisory Board

Professor Sir Peter Crane

Yale University

(OLS

Dr. Rogier P.J. de Kok

Royal Botanic Gardens, Kew

U.K.

Dr. W.J. Kress

National Museum of Natural History

Smithsonian Institution

(GESTAL

Dr. David J. Middleton

Royal Botanic Garden Edinburgh

(ULIK.

Dr. Kiat W. Tan

Gardens By The Bay

National Parks Board, Singapore

Dr. Nigel P. Taylor

Singapore Botanic Gardens

National Parks Board, Singapore

Dr. I.M. Turner

Singapore Botanic Gardens

National Parks Board, Singapore

Dr. J.F. Veldkamp

Nationaal Herbarium Nederland

The Netherlands

Dr. Jun Wen

National Museum of Natural History

Smithsonian Institution

USSVA:

Professor Nianhe Xia

South China Institute of Botany

PR. China

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

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

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

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

Botanic Gardens, 1 Cluny Road, Singapore 259569”.

National Parks Board

Singapore Botanic Gardens

1 Cluny Road

Singapore 259569

Printed by Oxford Graphic Printers Pte Ltd

CONTENTS OF VOLUME 64 (2012)

VOL. 64 (1), July 2012

K.M. Wong

Anondted years of the Gardens’ Bulletin, Singapore \.2.2.z......:...ccsso0..2ccscesecceceecoeeseeeentoees

Charles Clarke and Ch’ien C. Lee

A revision of Nepenthes (Nepenthaceae) from Gunung Tahan, Peninsular Malaysia .......

Heiena Duistermaat

A taxonomic revision of Amischotolype (Commelinaceae) in ASIa ............::csceeceeeseeeeeees

H. Kurzweil and S. Lwin

First record of Taeniophyllum (Orchidaceae) in Myammat ...............:cccssccessccesceesscesseeeeee

C.Y. Ling and S. Julia

Diversity of the tree flora in Semenggoh Arboretum, Sarawak, Borneo ...............:::000000

R.R. Mill and M. Whiting

Podocarpus orarius (Podocarpaceae), a new species from the Solomon Islands and a

taxonomic clarification of Podocarpus spathoides from Malaysia ............::cccccccseeeseeeeneees

Q.B. Nguyen and Jana Leong-Skorni¢kova

Distichochlamys benenica (Zingiberaceae), a new species from Vietnam .................00664

Hans-Juergen Tillich and Leonid V. Averyanoy

Four new species of Aspidistra Ker-Gawl. (Asparagaceae) from China and Vietnam

with a comment on A. longifolia Hook.f. and A. hainanensis W.Y.Chun & F.C.How ........

A.P.J. Ting, S.Y. Wong, J. Jamliah and P.C. Boyce

Phylogenetic study of the Schismatoglottis Nervosa Complex

I.M. Turner

The plant taxa of H.N. Ridley, 4. The primitive angiosperms (Austrobaileyales,

Canellales, Chloranthales, Laurales, Magnoliales, Nymphaeales and Piperales) .............

S.Y. Wong, P.C. Boyce and S.L. Low

Studies on Schismatoglottideae (Araceae) of Borneo XVII: The Schismatoglottis

Hottae Complex, a new informal taxon, and three new species from Sarawak,

SALA) SSRI BOTA (SO a a Reece eee Oe ee eo ene

BOOK REVIEWS

Heritage Trees of Penang. S. Gardner, P. Sidisunthorn & E.M. Lai. 2011.

(SIRS (GRIER) 38.558 8 chee cb cdocconooec BRB RERDAEARCOECRCORT OSC ERR ERE cS BREE spc cc sae ee eee nen

The Wild Orchids of Hong Kong. G. Baretto, PJ. Cribb & S. Gale. 2011.

Utsler yesh SUN Trav ell) Mee eee MD ane eS een Naa 8 ats owhcana fads vacua ceeadeanssstsabewsess savesgeosees

Flora of Peninsular Malaysia. Series II: Seed Plants, Volume 2. R. Kiew,

R.C.K.Chung, L.G. Saw, E. Soepadmo & P.C. Boyce (eds). 2011. (Y.W. Low) ............255

Wild Orchids of Peninsular Malaysia. PT: Ong, P. O'Byrne, W.S.Y. Yong

ill

201

211

221

e250

273

GE INC yer 2A DIP SCI ACG 00 a ec a eee eee 279

VOL. 64 (2), December 2012

P. C. Boyce, Zulhazman Hamzah and Sofiman Othman

Studies on Monstereae (Araceae) of Peninsular Malaysia IV: The enigmatic

Rhaphidophora corneri tetound:alieta/ Si ycatSie te: 2 eee eee ee ee 281

C. Pramod, A.K. Pradeep and J.F. Veldkamp

Coelachne madayensis (Poaceae: Pooideae: Isachneae), a new species from

Kerala, Tindal isc:.c.iecesssesedicezs acne tens oo eee te cee ees omer teece coe enone es concen ee ee 289

M. Rodda, H.D Tran, P.T. Chiew, D. Liew and J. Leong-Skorniékova

The rediscovery of Dischidia hirsuta (Apocynaceae, Asciepiadoideae)

16 Whey 1012421 00) 1 oe 8250 sc 2nccacacn asa Cees cel ea oc ooae sect oe: vaccucekece soosge oountceebate: 293

M. Sugumaran and K.M. Wong

Studies in Malesian Gentianaceae I: Fagraea sensu lato—complex genus

or several genera? A moleculanphylogemetie stir yeessncee erences senescence 301

Pramote Triboun and David J. Middleton

Twenty New species of Paraboecal(Gesneriaceae) kom) Mhailand ees2-- ee eee 333

I.M. Turner

Annonaceae of Bormeo: areview of the climbing species -c.ts--7--s5-22- see ee 371

K.M. Wong and M. Sugumaran

Studies in Malesian Gentianaceae II: A taxonomic framework

for the Fagraea complex, including the new genus Limahlania ............cccccccessceesseessseenseeeees 481

K.M. Wong and M. Sugumaran

Studies in Malesian Gentianaceae III: Cyrtophyllum reapplied

to the Fagraea fragranis Alliance axeeceere- cass = tenn toec ee eee eee eee ee 497

K.M. Wong

Studies in Malesian Gentianaceae IV: A revision Of Picrophloeus .......c.sccecceeseeseseeeeeeeseees Sil

Zulhazman Hamzah, P. C. Boyce and Mashhor Mansor

Studies on Homalomeneae (Araceae) of Peninsular Malaysia IV:

Homalomena stongensis, a remarkable new species endemic

to Gunung Stong, Kelamtatt .....2.5.sics,cusscss-seaccodts-nceesdedecee stone ccencrastesesosevestcundunans (ox career 523

BOOK REVIEWS 2

Etlingera of Sulawesi. 4.D.Poulsen. 2012 (J. Leong-Skorni€kova) ..............-.sscceseeseseereseees 529

Flora of Hong Kong. Volume 4. N.H. Xia, K.W. Cheung, K.S. Pang

& K.L. Yip: (eds). 201) (KM. Wott) acsectecec cere at eee scrote tee voted incre eee tee ee 533

Private Lives. L.K. Wang, D.C.J. Yeo, K.K.P. Lim & S.K.¥. Lum (eds). 2012

00.1) URC 092) epee eee eRe rye eocc sob on ceed adsoor ou snocaocdp asocobenciin-oabonatvodcuadooouasinanonsded S85)

Instructions for contributing authors (continued)

Title and authorship. The title should concisely describe the contents. If a scientific name is used, its

authority is normally excluded, but the family name would be provided. Authors’ names, affiliations

and postal/ e-mail addresses are stated below the title. If more than one author, indicate “corresponding

author”. Avoid footnotes. A short running title (up to six words) should also be provided.

Abstract. The abstract is at most 100-300 words. It should concisely indicate the article’s contents without

summarising it; mentioning novelties and name changes. Keywords: Suggest at most eight keywords, in

alphabetical order.

Scientific names and author abbreviations. Genus and species names of organisms must be italicised and

followed by the authority (with family name in parentheses) when first mentioned in the text or diagnoses.

Standards for author abbreviations include:

Brummitt, R.K. & Powell, C.E. (1992) Authors of Plant Names. Kew: Royal Botanic Gardens, Kew.

Kirk, P.M. & Ansell, A-E. (1992) Authors of Fungal Names [Index to Fungi Supplement]. Wallingford:

CAB International.

Herbarium abbreviations (http://sciweb.nybg.org/science2/IndexHerbariorum.asp) follow

Holmgren, P.K., Holmgren, N.H. & Barnett, L.C. (eds) (1990) Index Herbariorum. Part 1: The Herbaria

of the World [Regnum Veg. vol. 120]. New York: New York Botanical Garden.

Journal and book title abbreviations. For journals:

Bridson, G.D.R., Townsend, S.T., Polen, E.A. & Smith, E.R. (eds) (2004) BPH-2: periodicals with

botanical content: comprising a second edition of Botanico-Periodicum-Hunlianum. Pittsburg: Hunt

Institute for Botanical Documentation.

For books:

Stafleu, FA. & Cowan, R. S. (eds) (1976-88) Taxonomic Literature. 2nd edition. 7 vols [Regnum Veg.

vols 94, 98, 105, 110, 112, 115, 116].

Stafleu, F.A. & Mennega, E.A. (eds) (1992—) Taxonomic Literature. Supplements [Regnum Veg. vols

152130; 132]:

(A useful source of verify ing names of publications is

<http://asaweb. huh. harvard.edu:8080/databases/publication index.html>. If in doubt, list full titles.

Other abbreviations and units of measurement. \f using standard abbreviations and acronyms, give the

full term on first mention. Dates are cited as: | Jan 2000. SI (metric) units of measurement are used and

spelled out except when preceded by a numeral; they are abbreviated in standard form: g, ml, km, etc.

Tables. Yables are numbered in arabic numerals in the order they are first mentioned in the text and carry

an indicative legend at the head. Tables are given at the end of the manuscript.

Illustrations. All drawings, maps, graphs and photographic images (individually or collected in a plate)

are to be numbered in arabic numerals in the order they are first mentioned in the text, as Fig. 1, Fig. 2,

etc. (plate components would be referred to in the text as Fig. 1A, 1B, Fig. 1A—D, etc.). If relevant, scale

bars should be used to indicate magnification.

When grouping photographs, the maximum page area 19.5 x 13 cm must be heeded. High resolution digital

images may be submitted as separate files (line drawings in black and white at 600 dpi, photographs at

300 dpi) sent electronically or in a CD. Do not embed images into the main text file.

References in the text. Citation in the text should take the form: King & Gamble (1886) or (King &

Gamble 1886), or King et al. (1886) if more than three authors to a work. Use 2000a, 2000b, etc. if several

papers by the same author(s) in one year are cited.

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 DELTA 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 trnL(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. & Darwin, S.P. (1988) Rubiaceae. In: Smith, A.C. (ed) Flora Viliensis Nova, A New Flora of

Fiji 4: 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:

Gardenia anisophylla Jack ex Roxb., Fl. Ind. ed. Carey & Wall. 2: 561 (1824).

Medinilla alternifolia Blume, Mus. Bot. 1: 19 (1849).

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

they should also be mentioned in the text briefly, e.g., “Nomenclatural references researched include

Blume (1849) and Roxburgh (1824).”

Homotypic synonyms should be provided in a block, stating the type at the end.

Front cover picture: Cyrtophyllum fragrans in bloom (Photo by Ang Wee Foong)

Back cover: Gardens’ Bulletin covers. 1972-2011 (Montace by Bazilah M. Tbrahim & S Tee)