THE GARDENS’ BULLETIN, SINGAPORE

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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 David J. Middleton Dr Jana Leong-Skomickova

(Editor-in-Chief) (Managing Editor)

Felicia Tay YeeWenEow Christina Soh

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

Editorial Advisory Board

Professor Sir Peter Crane

Yale University

USA.

Dr Rogier P.J. de Kok

Royal Botanic Gardens, Kew

UK.

Dr W. John Kress

National Museum of Natural History

Smithsonian Institution

U.S.A.

Dr Mark Hughes

Royal Botanic Garden Edinburgh

UK.

Dr Kiat W. Tan

Gardens By The Bay

Singapore

Dr Nigel P. Taylor

Singapore Botanic Gardens

National Parks Board, Singapore

Dr Ian M. Turner

Royal Botanic Gardens Kew

UK.

Dr Jan-Frits Veldkamp

National Biodiversity Center

The Netherlands

Dr Jun Wen

National Museum of Natural History

Smithsonian Institution

U.S.A.

Professor Nianhe Xia

South China Institute of Botany

PR. China

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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 inti'on and //77L(UAA)-F(GAA) intergenic spacer. NorrJicJ. 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) Flor-a Viliensis Nova, A New Flora

q/’E’//7 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 GenBanlc should be

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

Style of nomenclatural summaries. The foUowmg style is required:

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

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

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

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

they should also be mentioned in the text briefly, e.g., “Nomenclatural references researched include

Blume (1849) and Roxburgh (1824).”

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Front cover picture: Licuala miriensis (Photo by K.M. Wong)

The Gardens’ Bulletin

Singapore

VOL. 67(2) 2015 ISSN 0374-7859

CONTENTS

R.P.J. de Kok

Cryptocarya nitens (Lauraceae), a new species record for Singapore 253

Y.Y. Sam

Boesenbet'gia basispicata (Zingiberaceae), a new record for

Peninsular Malaysia 261

K.M. Wong

Schizostachyum kuisingii, a new species of bamboo

(Poaceae: Bambusoideae) from Peninsular Malaysia 267

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

Novitates Bruneienses, 4. New records in the Araliaceae, Araucariaceae,

Arecaceae, Fagaceae, Musaceae and Thymelaeaceae 275

T.L. Yao

Three new species of Loxocarpus (Gesneriaceae) from Sarawak, Borneo 289

W.H. Ardi & D.C. Thomas

Studies on Begonia (Begoniaceae) of the Moluccas II:

Anew species from Seram, Indonesia 297

M. Rodda

Anew typification of Hoy a zollingeriana (Apocynaceae, Asclepiadoideae) 305

R.P.J. de Kok

A revision of Cryptocarya (Lauraceae) from Thailand and Indochina 309

H. T. Liru, J. Leong-Skornickova, L.X.B. Nguyen, C.T. Do & T.T. Hoang

Newmania sessilanthera (Zingiberaceae): a new species from Vietnam 351

P.S. Ashton

A few last words on Far Eastern Dipterocarps 357

D.M. Johnson & N.A. Murray

A contribution to the systematics ofXylopia (Annonaceae) in Southeast Asia 361

K.M. Wong

Studies in Southeast Asian Melastoma (Melastomataceae), 1 .

Morphological variation in Melastoma malabathricum and notes

on rheophytic taxa and interspecific hybridisation in the genus 387

I. M. Turner

Clarifying the nomenclature of Crateva trifoliata (Capparaceae) 403

BOOK REVIEWS

A Guide to Begonias of Borneo. Ruth Kiew, Julia Sang, Rimi Repin

& Joffre Ali Ahmad. 2015. (D.C. Thomas) 407

Forest Trees of Southern Thailand, Volume 1 (A-Es). Simon Gardner, Pindar

Sidisunthom & Kongkanda Chayamarit. 2015. (D.J. Middleton) 409

Flora of Cambodia, Laos and Vietnam.

Fascicle 33 Apocynaceae. David J. Middleton. 2014.

Fascicle 34 Polygalaceae. Colin A. Pendry. 2014.

Fascicle 35 Solanaceae. Sovanmoly Hul & Pauline Dy Phon. 2014.

(K.M. Wong & M. Rodda) 411

Date of publication: 23 December 2015

National Parks Board

Singapore Botanic Gardens

1 Cluny Road

Singapore 259569

Printed by Oxford Graphic Printers Pte Ltd

Gardens’ Bulletin Singapore 67(2): 253-259. 2015

doi: 10.3850/S2382581215000204

253

Cryptocarya nitens (Lauraceae),

a new species record for Singapore

R.PJ. de Kok

275 Cricklewood Lane, London, NW2 2JJ, U.K.

dekokrogier@gmai] . com

ABSTRACT. Ciyptocarya nitens (Blume) Koord. & Valeton is newly recorded for Singapore.

It was discovered during surveys in Bukit Tiinah Nature Reserve and the Nee Soon Swamp

Forest. A description is given, together with a key based mostly on vegetative characters for all

Cryptocaiya species occurring in Singapore, A short overview of the Lauraceae of Singapore

shows that, in total, 57 species belonging to 14 genera have been recorded, of which 47 species

in 1 3 genera are native. Ciyptocarya nitens is lectotypified in addition to two of its synonyms.

Keywords. Cryptocarya, Endiandra

Introduction

Species of the Lauraceae (Laurel Family) are of major ecological and economic

importance in Southeast Asia as they comprise a major part of almost any forest in the

region. The family is very complex, posing many challenging taxonomic and systematic

questions, and for various reasons the identification of species is problematic. Some

systematic work has been done, for instance Kostennans (1952, 1957a, 1957b, 1964)

wrote a series of papers on various genera or parts of genera during his career, and

he also dealt with some aspects of the higher taxonomy. The most recent overview

of the family as a whole was by Rohwer (1993). In Eastem Asia and Australasia,

only for Peninsular Malaysia (Kochummen, 1989), China (Li Xiwen et al., 2008)

and Australia (Le Cussan & Hyland, 2007) are recent treatments available, dealing

with all species occumng in their territories. The classification and delimitation of the

genera within the Lauraceae has always been problematic. The general framework of

characters which has been employed by most authors was first established by Nees

von Esenbeck (1836) in his classification of the family. However, the outcome of such

a classification can vary greatly, depending upon the weight given to each particular

character. Recently, with the help of molecular data, some taxonomic relationships are

becoming clearer (Rohwer et ah, 2014).

The checklist for Singapore records 57 species of Lauraceae, of which only

47 are native (Chong et ah, 2009). This is a high percentage (82%) of native taxa for

a plant family as the average for Singapore is 51.3% (Chong et al., 2009). Amongst

the native species of Lauraceae, the vast majority have a widespread distribution,

occurring also in Peninsular Malaysia and at least one other area, such as Borneo and/

or Sumatra. No Lauraceae are restricted to Singapore and only two species found in

254

Gard. Bull. Singapore 67(2) 2015

Singapore are endemie to the Malay Peninsula (Actinodaphne malaccensis Hook.f.

and^. pruinosa Nees). According to Kochummen (1989) and De Kok (in press I & II),

almost all Lauraceae species native to Singapore are found growing in lowland forest

and usually also in hill or montane forest, or rarely also in swamp forest in Peninsular

Malaysia. Two species are restricted to swamp forest (e.g. Litsea gracilipes Hook.f

and Nothaphoebe coriacea Kosterm.).

Most of the species recorded from Singapore are reported as common in

Peninsular Malaysia (Kochummen, 1989; De Kok, in press I, in press II). However, in

Singapore, these taxa have been assigned a wide range of of conservation assessments

(Fig. 1, column 1), from extinct (ex), critically endangered (ce), endangered (en),

vulnerable (vu) to common (co) (Chong et al., 2009). In contrast, those species that are

reported to be rare, uncommon or scattered in Peninsular Malaysia have been assigned

to a more restricted set of conservation assessments, namely extinct, endangered or

critically endangered (Fig. 1, column 2).

Most genera recorded as native in Peninsular Malaysia are also native in

Singapore, with three exceptions. The first two are not problematic: Hexapora Hook.f.

is a monotypic genus endemic to Penang Island; Cinnadenia Kosterm. is a genus of

two species which is represented in Peninsular Malaysia by only one species from the

hill forests of Selangor and Negeri Sembilan. However, the third exception, Endiandra

R.Br., is curious. It does not seem to be native in Singapore as the two recorded

Fig. 1. The conservation status of Lauraceae in Singapore. Column 1 shows the conservation

status in Singapore for species that are common in Peninsular Malaysia; Column 2 shows the

conservation status in Singapore for those that are rare, scattered or uncommon in Peninsular

Malaysia, ex = Extinct, ce = Critically endangered, en = Endangered, vu = Vulnerable, co =

Common in Singapore (based on Chong et al., 2009).

Cryptocarya nitens in Singapore

255

species, Endiandra impressicosta C.K.Allen and E. longipedicellata C.T. White &

W.D. Francis, are only known from cultivation and are originally from Australia and

New Guinea, not from Peninsular Malaysia (Kochummen, 1989). Of the ten species

in this genus that are native to Peninsular Malaysia, most are reported to be rare, with

only two {Endiandra macrophylla (Blume) Boerl. and E. maingayi Hook.f) said to

be common in lowland forest (Kochummen, 1 989). As stated earlier, species that are

considered uncommon in Peninsular Malaysia tend to be (critically) endangered or

extinct in Singapore (see Fig. 1, column 2), so the absence of Endiandra as a native

genus in Singapore is possibly the result of local extinction (before a single specimen

could be collected), rather than a natural phenomenon.

New record for Singapore

During my visit to the Herbarium of Singapore Botanic Gardens in May 2015, 1 was

able to confiim the identification of Ctyptocarya nitens (Blume) Koord. & Valeton,

which is a new record for Singapore. This species was collected twice in 2006 from

Bukit Timah and more recently, but sterile, from a survey of the Nee Soon Swamp

Forest.

Cryptocarya nitens (Blume) Koord. & Valeton, Meded. Lands Plantentuin 68: 220-

223 (1904); Kochummen, Tree FI. Mai. 4: 136 (1989). - Tetranthera nitens Blume,

Mus. Bot. 1 : 375 (1851). -Type: [Indonesia] Sava.Blume s.n. (lectotype L [L0036214],

designated here; isolectotype L [L0036213]). (Fig. 2)

Cryptocarya areolata Gamble, Bull. Misc. Inform. Kew 1910: 144 (1910). - Type:

[Peninsular Malaysia] Perak, May 1 884, King s collector 61 71 (lectotype K, designated

here; isolectotypes BM [BMOO 1124601], L [L0036209, L0036210], P [P02010230],

SING [SING0209177, SING0209I78]).

Cryptocarya buhongana Gamble, Bull. Misc. Inform. Kew 1910: 144 (1910).

- Type: [Peninsular Malaysia] Ulu Bubong, July 1886, Kings collector 10570

(lectotype K [KOO 1084494], designated here; isolectotypes K [KOO 1084495], SING

[SING0046590]).

Cryptocarya nativitatis Rendle ex Baker.f in Andrews, Monogr. Christmas Isl.

187 (1900). Type: [Australia] Christmas Island, Phosphate Hill, 1897, Andrews

158 (holotype BM [BM000799315]; isotypes BO, K [K000768438], SING

[SING0069586].

Trees 8-30 m tall, dbh 15-30 cm. Bark smooth or scaly, grey-brown, inner bark

brownish, wood cream coloured with spicy odour. Twigs strongly angular when

young, velutinous, hairs brown. Leaf lamina leathery, elliptic to lanceolate, ovate to

256

Gard. Bull. Singapore 67(2) 2015

obovate (6-)7.5-23 x (2.2-)3.5-10.5 cm, apex acute to acuminate, base cuneate or

rounded, slightly unequal; pinnately veined, secondary veins 5-10 pairs, tertiary veins

scalarifomi; upper surface glabrous, sometimes with hairs on midrib and secondary

veins, midrib sunken, secondary veins sunken, tertiary veins faint, shiny, (light to

yellow) green, drying greenish brown; lower surface glabrous sometimes with hairs

on midrib and secondary veins, midrib and secondary veins raised, tertiary veins

faint, light blue, pale green, glaucous. Petiole slender, 10-30 mm long, velutinous.

Inflorescences 10-15 mm long, velutinous, greenish white; bracteoles triangular

0.5-0. 6 mm long, caducous. Flowers pale green to yellow or white, hairs yellowish;

perianth tube 1.2-1. 6 mm long, velutinous; perianth lobes elliptic to lanceolate, 1.3-

1.8 X 0.8-1 mm, apex acute, velutinous, greenish white. Stamens 1.2-1. 6 mm long,

hairy, dull greenish yellow, anther bright yellow-orange. Ovary clavate, 1-1.2 mm,

glabrous; style 1.2-1. 6 mm long, linear; stigma inconspicuous. Fruit (dried) globose,

8-16 mm diameter, shallowly ridged, sparsely hairy, black when mature. Stalk slender

when mature.

Distribution. Peninsular Thailand, Malaysia, Singapore, Indonesia (Sumatra and Java),

Australia (Christmas Island).

Ecology. Over its general range it is found on riversides in open, mixed (including

bamboo) hill to lowland forest, growing on limestone, sandstone and granite, at 0-250

m altitude. In Singapore recorded from swamp forest and along water courses.

Provisional regional lUCN conservation assessment. Least Concern.

Phenology. Flowering from March to October; fruiting from January to November.

Notes. In the original description of Tetranthera nitens Blume (1851: 375), only

one collection is cited, of which there are two morphologically virtually identical L

specimens available for lectotypification. The specimen L0036214 is selected here as

lectotype because the label mentions at least some of the additional information cited

in the original description, while the other specimens lacks any label data.

In the original description of Cryptocarya areolata Gamble (1910: 144), four

different collections are cited: Kings collector 6017, 6171, 8630 and Wray 2456.

Gamble’s top set of specimens are housed at K and from these the duplicate of Kings

collector 6171 is selected here as the lectotype. It, uniquely amongst the syntypes, has

dissected dowers glued on a card with notes in Gamble’s handwriting.

In the original description of Cryptocaiya bubongana Gamble (1910: 144),

only one collection is cited: King’s collector 10570. From the two virtually identical

K collections (both morphologically and in label date), one is selected here as the

lectotype [KOO 1084494].

This species is easy to distinguish from the other Singaporean Cryptocarya

species as the under surface of its leaves is glabrous. In all other species have small

appressed hairs can be seen with a x 10 hand lens.

Cryptocarya nitens in Singapore

257

Fig. 2. Cryptocarya nitens (Blume) Koord. & Valeton, twig with fruits photographed from the

Bogor Botanie Gardens, tree no. XX.B.59. (Photo: Jens Rohwer)

Selected specimens examined. SINGAPORE: Bukit Timah Nature Reserve, Publie Utilities

Board Catehment, 27 Jun 2006, Khoo KMS 14 (SING); ibidem, 7 Jul 2006, Chua CSC 1 (SING).

Key to the Cryptocarya species in Singapore based mostly on vegetative characters

1 a. Lower surface of mature leaves glabrous apart from veins C. nitens

lb. Lower surface of mature leaves sparsely hairy to velutinous 2

258

Gard. Bull. Singapore 67(2) 2015

2a. Midrib sunken at base on upper leaf surface 3

2b. Midrib raised at base on upper leaf surface 5

3a. Petiole channelled; bracteoles elliptic, < 1 mm long C.ferrea

3b. Petiole half terete; bracteoles elliptic to lanceolate or linear, > 1 mm long 4

4a. Leaves leathery; bracteoles elliptic to lanceolate, (1 .5-)2-3.5 nmi long

C. rugulosa

4b. Leaves thinly leathery; bracteoles linear, 1 . 1-1 .8 mm long C. kurzii

5a. Leaves bullate; petiole swollen C. griffithiana

5b. Leaves not or only slightly bullate; petiole slender 6

6a. Leaf base symmetrical, hairs msty to dark brown (when dried) C. impressa

6b. Leaf base slightly oblique, hairs light brown (when dried) C. malayana

ACKNOWLEDGEMENTS. This research was supported in 2015 by a Singapore Botanic

Gardens Research Fellowship which is gratefully acknowledged. The author is grateful to

Dr Helen Fortune-Hopkins who corrected my English, to Dr Jens Rohwer who kindly gave

pemiission to use his pictures, and to the curators of the BM, BO, K, KEP, L and SING herbaria

for access to the spechnens used in the present study.

References

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

Brill.

Chong, K.Y., Tan. H.T.W. & Corlett, R.T. (2009). ^1 Checklist of the Total Vascular Plant Flora

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

of Biodiversity Research, National University of Singapore.

De Kok, R.P.J. (in press 1). A revision of Ctyptocaiya R.Br. (Lauraceae) of Peninsular Malaysia.

Kew Bulletin.

De Kok, R.RJ. (in press 11). A revision of Beilschmiedia Nees (Lauraceae) of Peninsular

Malaysia. Blumea.

Gamble, J.S. ( 1910). New Lauraceae from the Malayan region 1. Bull. Misc. Inform. Kew 1910:

142-153.

Kochuimnen, K.M. (1989). Lauraceae. In: Ng, F.S.P. (ed) Tree Flora of Malaya 4: 98-178.

Kuala Lumpur: Longman Sdn. Bhd.

Kostennans, A.J.G.H. (1952). A historical survey of Lauraceae, part 11. J. Sci. Res. (Jakarta)

1: 113-127.

Kostennans, A.J.G.H. (1957a). Lauraceae. Reinwardtia 4: 193-256.

Kostennans, A.J.G.H. (1957b). Lauraceae. Communi. Forest Res. Inst. Bogor 51 \ 1-64.

Kostennans, A.J.G.H. (1964). Bibliographia Lauracearum. Bogor, Indonesia: Departemen

Urusan Research Nasional.

Le Cussan, J. & Hyland, B.P.M. (2007). Lauraceae (excluding Cassytha). In: Wilson, A.J.G.

(ed) Flora of Australia 2: 106^86. Melbourne: CSIRO Publishing.

Cryptocarya nitens in Singapore

259

Li Xiwen, Li Jei, Huang Puhua, Wei Fa’nan, Cui Hongbin & Van der Werff, H. (2008).

Lauraceae. In: Wu Zhengyi & Raven, R (eds) Flora of China 7: 102-254. Beijing:

Science Press and St. Louis: Missouri Botanical Garden.

Nees von Esenbeck, C.G.D. (1831). Lauraceae. In: Wallich, N. Plantae Asiaticae Rariores, vol

2: 58-76. London.

Rohwer, J.G. (1993). Lauraceae. In: Kubitzki, K., Rohwer, J.G. & Bittrich, V. (eds) The

Families and Genera of Vascular Plants, vol. 2: 366-391. Berlin: Springer- Verlag.

Rohwer, J.G., De Moraes, P.L.R., Rudolph, B. & Van der Werff, H. (2014). A phylogenetic

analysis of the Cryptocarya group (Lauraceae), and relationships of Dahlgrenodendron,

Sinopora, Triadodaphne and Yasunia. Phytotaxa 158: 111-132.

Gardens’ Bulletin Singapore 67(2): 261-265. 2015

doi: 10.3850/S2382581215000216

261

Boesenbergia basispicata (Zingiberaceae),

a new record for Peninsular Malaysia

Y.Y. Sam

Forest Research Institute Malaysia,

52109 Kepong, Selangor, Malaysia

samy en@frim . gov.my

ABSTRACT. Boesenbergia basispicata K.Larsen ex Sirirugsa was recently discovered from

a limestone hill in Perak, Peninsular Malaysia. This brings the total number of Boesenbergia

species in Peninsular Malaysia to 12. A description, colour plates and notes on this species are

provided.

Keywords. Limestone, Peninsular Thailand, Perak

Introduction

Boesenbergia Kuntze is a genus of gingers found in tropical Asia. The genus consists

of about 80 species ranging from India to China and Southeast Asia (Larsen et al.,

1998; Saensouk & Larsen, 2002; Mood et al, 2014). Holttum (1950) published the first

comprehensive treatment of the genus in Peninsular Malaysia in which he recognised

eight species, namely Boesenbergia clivalis (Ridl.) Schltr., B, curtisii (Baker) Schltr.,

B. Jiava Holttum, B. longipes (King & Prain ex Ridl.) Schltr., B. plicata (Ridl.)

Holttum, B. prainiana (King ex Baker) Schltr., B. pulcherrima (Wall.) Kuntze and B.

rotunda (L.) Mansf. Among these, Boesenbergia rotunda is not considered to be native

as only cultivated plants were sighted. Since then, there was no taxonomic work on

the genus in Peninsular Malaysia until Lim (2008) named a new species, Boesenbergia

kenali C.K.Lim, from Kelantan. He also provided a checklist for Peninsular Malaysian

Boesenbergia in which 1 1 species were listed. The species newly added to the list

included Boesenbergia lurida (Ridl.) Loes., which was resurrected from synonymy

of B. plicata (Ridl.) Holttum, and the rediscovery of B. minor (Bak.) Schltr. Recently,

a Thai species, Boesenbergia basispicata K.Larsen ex Sirirugsa, was discovered in

northern Peninsular Malaysia, adding another species to the list.

Boesenbergia basispicata K.Larsen ex Sirirugsa, Nordic J. Bot. 7: 423 (1987);

Sirirugsa, Nat. Hist. Bull. Siam Soc. 40: 67 (1992). - TYPE: Thailand, Naklion Si

Thammarat, Kao Luang, in evergreen forest, 300 m, 2 May 1926, Kerr 15575 (holotype

K; isotypes BK, BM). (Fig. 1)

262

Gard. Bull. Singapore 67(2) 2015

Rhizomatous evergreen herb c. 50 cm tall, l-(2) leafy shoots per clump. Rhizome

short, not branched, 8-10 mm in diam., light yellow internally, roots tuberous. Leafy

shoot composed of 3-4 leaves, arranged distichously, with no obvious pseudostem;

bladeless sheath 1-2, to 1 1 cm long, mottled with red, soon decaying; leaf sheath 1 1.5-

15 cm, longitudinally ridged, green mottled sparsely with red, margin thin and broad;

ligule thin, bilobed, lobes triangular, c. 2 mm long; petiole 1 1 .5-1 5 cm long, deeply

channelled; lamina ovate to narrowly ovate, largest 25-28 x 10.3-13.5 cm, adaxially

green, main veins slightly raised (not plicate), abaxially pale green, tinged red on old

leaves, apex acute to attenuate, base oblique and subcordate. Inflorescence radical,

7.5-9 cm long, mottled with red; peduncle 1-3 cm long, covered with sheathing bract;

spike conical, 6.5-8 cm long, composed of 10-11 distichously an*anged overlapping

fertile bracts (lowermost bract fertile bracts 22-28 x c. 10 mm, elliptic, boat-

shaped, glabrous, apex acute, margin incurved, each bract supporting single flower;

bracteole 1 7-20 mm long, shorter than bract, cylindrical, open to the base, glabrous,

apex acute. Flower 4-5 cm long, white except the variegation on labellum; calyx

c. 5 mm long, tubular, glabrous, apex tiTincate, sometimes with a slight incision;

floral tube 30-35 mm long, slender, c. 1 mm in diam. at base, widening to 2 mm

distally, glabrous externally and internally; corolla lobes c. 13 x 4-6 mm, elliptic,

membranous, apex obtuse, margin incurved but not overlapping, dorsal lobe with

hooded apex; androecial tube c. 1 cm long, sparsely hairy internally; labellum c. 18 x

1 7 mm, saccate, suborbicular, abaxially sparsely hairy, white, centrally yellow mottled

with red, the yellow extending as 2 bands to apical, ending c. 5 mm before touching

the apex, apical area chilli-red, slight pinkish tinge on both sides, apex wrinkled and

deflexed, margin entire; lateral staminodes c. 10 x 5 mm, broadly elliptic, sparsely

hairy abaxial ly, apex broadly acute. Stamen c. 7 mm long, with sparse glandular hairs

abaxially; filament c. 2 mm long; anther c. 6 mm long, without anther-crest, anther

thecae dehiscing longitudinally. Ovary obovoid, c. 3 mm long, glabrous, unilocular,

free-basal placentation in mature flowers; style c. 46 mm long, filiform, glabrous,

stigma less than 0.5 mm long, club-shaped, glabrous, ostiole transverse, without cilia,

face upwards; epigynous glands 2, c. 5 mm long, filiform. Fruit c. 1 0 x 5 mm, ellipsoid,

glabrous, wall thin and translucent (dehiscence not observed); seeds ellipsoid, c. 7 x 2

mm, arillate, aril white, laciniate.

Additional specimens examined’. PENINSULAR MALAYSIA: Perak: Hulu Perak, Belukar

Semang Lores! Reserve, Bukit Telor Ayam 1, limestone hill area, 15 Nov 20\\,Mohd. Hairul et

al. FRI 72405 (KEP); Tanah Hitam, limestone hill next to Rahman Plantation, 372 m altitude,

5°35.80’N 101°00.09’E, \1 M 2012, Sam FRI 68937 {KEV).

Habitat and ecology. Herbaceous vegetation on karst limestone substrate, surrounded

by regenerating lowland evergreen rainforest. The plants were found on two small

limestone outcrops covered with thick vegetation. They were seen growing in the

crevices and ledges on the outcrop, about 10 m above the ground. These narrow

pockets contained thick humus supporting the growth of herbaceous plants such as

Boesenbergia basispicata, aroids {Arisaema fimbriatum Mast.), gesneriads {Damrongia

Boesenbergia basispicata, a new record for Peninsular Malaysia

263

Fig. 1. Boesenbergia basispicata K.Larsen ex Sirirugsa. A. The limestone outcrop where the

plants were found, adjacent land has been cleared for planting rubber trees. B. Plant clinging

onto the limestone cliff face. C. Radical inflorescences. D. Flower. E. The dehisced anther

thecae. F. Stigma. G. The cross section of ovary. H. Free-basal placentation. (Photos: Y.Y.

Sam)

lacunosa (Hook.f.) D.J.Middleton & A.Weber, Epithema saxatile Blume, Microchirita

rupestris (Ridl.) A.Weber & Rafidah, Paraboea verticillata (Ridl.) B.L.Burtt), ferns and

lycophytes (Adiantum malesianum Ghatak, Pyrrosia penangiana Holttum, Selaginella

frondosa Warb.). This is the first report of Boesenbergia basispicata growing on a

limestone substrate. In Southern Thailand, the plants are often found on damp slopes

(Sirirugsa, 1992) and also near streams on sandstone with thick leaf litter (Mood, pers.

comm.) in lowland evergreen forest.

264

Gard. Bull. Singapore 67(2) 2015

The population of Boesenbergia basispicata at Belukar Semang Forest

Reserve is potentially threatened by disturbance. Firstly, the Forest Reserve may

suffer from encroaclmient through illegal land clearance, since it is now adjacent to

a rubber plantation and thereby more accessible. Secondly, the land clearance for the

adjoining rubber plantation has adversely affected the environment as the humidity

and temperature have become less favourable to understorey plants like Boesenbergia

basispicata.

Distribution. Peninsular Thailand and Peninsular Malaysia.

Notes. Boesenbergia basispicata was first discovered in Naklion Si Thammarat and

Trang Provinces, Peninsular Thailand (Sirirugsa, 1987) and later in the southernmost

province of Narathiwat (Sirirugsa, 1992). It is one of only around eight species of

Boesenbergia with an inflorescence emerging directly from the rhizome; most species

produce a terminal inflorescence. In Peninsular Malaysia, Boesenbergia basispicata

is the first species recorded with a radical inflorescence and this makes it easy to

distinguish from the other species with a terminal inflorescences, especially from the

otherwise similar B. prainiana.

In general, the plants at Belukar Semang Forest Reserve fit Sirimgsa’s

descriptions but differ in the ovate to narrowly ovate laminas with subcordate bases,

whereas the Thai plants have oblong laminas with obtuse or cuneate to decurrent

bases, the broader laminas (10.3-13.5 vs 4-9.5 cm), longer peduncles (1-3 vs 0.5

cm) and floral tubes (30-35 vs 25 mm) and smaller fruits (c. 10x5 mm vs c. 7 x 2

mm). A reviewer also noted that pseudostems have been observed in Thai populations.

Another marked difference observed in the Peninsular Malaysian plant is its unilocular

ovary with free-basal placentation in contrast to the incompletely trilocular ovary in

Siriruga’s account.

ACKNOWLEDGEMENTS. This study is supported by the ‘Documentation and Inventory of

Flora Malaysia’ Project under the Ministry of Natural Resources and Environment’s Tenth

Malaysian Plan. The author would like to thank Dr Ruth Kiew and Dr R.C.K. Chung, for their

comments and suggestions on the manuscript, and Mohd. Hainil Mohd. Amin and Angan Atan,

for their assistance in the field.

References

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

82-105.

Larsen, K., Lock, J.M., Maas, H. & Maas, P.J. (1998). Zingiberaceae. In: Kubitzki, K. (ed) The

Families and Genera of Vascular Plants 4: 474^95. Germany: Springer- Verlag.

Lim, C.K. (2008). Anew species of Boesenbergia Kuntze from Kelantan, Peninsular Malaysia.

Folia Malaysiana 9(1): 69-76.

Boesenbergia basispicata, a new record for Peninsular Malaysia

265

Mood, J.D., Veldkamp, J.F. & Prince, L.M. (2014). A new species and a new record of

Boesenbergia (Zingiberaceae) for Thailand. Gard. Bull. Singapore 66: 207-214.

Saensouk, S. & Larsen, K. (2002). Boesenbergia baimaii, a new species of Zingiberaceae from

Thailand. Nordic J. Bot. 21: 595-597.

Sirirugsa, P. (1987). Three new species and one new combination in Boesenbergia

(Zingiberaceae) from Thailand. Nordic J. Bot. 1: 421^25.

Sirirugsa, S. (1992). A revision of the genus Boesenbergia Kuntze (Zingiberaeeae) in Thailand.

Nat. Hist. Bull. Siam Soc. 40: 67-90.

Gardens’ Bulletin Singapore 67(2): 267-274. 2015

doi: 10.3850/S2382581215000228

267

Schizostachyum kuisingiU a new species of bamboo

(Poaceae: Bambusoideae) from Peninsular Malaysia

K.M. Wong

Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, Singapore 259569

wkm2000@gmail.com

ABSTRACT. Schizostachyum kuisingii K.M.Wong is a new species of bamboo from Peninsular

Malaysia, distinguished from closely related species from adjacent areas in Indonesia, such as

Schizostachyum castaneum Widjaja, Schizostachyum lutescens Widjaja and Schizostachyum

mampouw Widjaja, by the hairs of the culm intemode, glabrescent foliage leaves, leaf ligule

fonn, a palea longer than the lemma and the presence of two lodicules in the flower.

Keywords. Bambuseae, Malay Peninsula, Malesia, Melocanninae, taxonomy

Introduction

Of some 60 bamboo species in 14 genera documented for Peninsular Malaysia (Wong,

1995), 11 are classified in Schizostachyum Nees. This paper describes a hitherto

unknown Schizostachyum species that Mr Tan Kui Sing brought me to see in Johor

state in Peninsular Malaysia in 2008, during a period when we were gathering bamboos

to represent a special living collection for Peninsular Malaysia. We have subsequently

been able to study the literature and specimens pertaining to the Schizostachyum

diversity in Southeast Asia, with the conclusion that this bamboo is new to science.

Herbarium acronyms used here follow Thiers (continuously updated).

Schizostachyum bamboos

Schizostachyum and allied genera, such as Melocanna Trin., Cephalostachyum

Munro, Pseudostachyum Munro, Teinostachyum Munro, Neohouzeaua A.Camus,

Dendrochloa C.E.Parkinson and Leptocama L.C.Chia & H.L.Fung, belong to the

Melocanninae subtribe of the Bambuseae (BPG, 2012; Kellogg, 2015). Whereas

Melocanna has always been easy to distinguish based on its long-necked sympodial

rhizomes and fleshy fruit pericarp, the relationships of the other genera mentioned here

have been contentious (Holttum, 1946, 1956). Otherwise the genera of Melocanninae

generally share a number of conspicuous characters: a branch complement of many

slender subequal branches developing from a single branch bud, and a glabrous ovary

with a rigid hollow attenuating upward extension of the ovaiy apex (analogous to a

stylar structure and often just simply called the ‘style’) connecting with the stigmas

268

Gard. Bull. Singapore 67(2) 2015

(Holttum, 1956); also there is a eonspicuous white-waxy ring just below each culm

node (Wong, 1 995).

Holttum (1946) had suggested that Schizostachyum might include

Cephalostachyum, Teinostachyum and Pseudostachyum because the number of flowers

per pseudospikelet and fiision of filaments may not be stable distinguishing characters.

Later, Holttum (1956) added Neohouzeaua to his broad concept of Schizostachyum,

Although limited-sampling molecular phylogenetic investigations have yet to clearly

resolve their generic positions, Pseudostachyum seems to be a distinct lineage (Yang

et ah, 2007). Although Xia (1993) recognised Cephalostachyum diwd Pseudostachyum

as distinct genera, he also deemed it is practical to include Dendrochloa, Leptocanna,

Neohouzeaua and Teinostachyum within Schizostachyum. From the molecular analyses

of Yang et al. (2007), it seems likely that Cephalostachyum may be heterogeneous,

with its type species C. capitatum Munro and an alliance including C. latifoHum Munro

(syn. C. fuchsianum Gamble) and C. pallidum Munro fomiing one possible generic

entity, and others like C. pergmcile Munro and C. virgatum (Munro) Kurz being

more closely allied to a group of Schizostachyum species including S. brachycladum

(Kurz) Kurz and S. zollingeri Steud. As it stands, too little is known about comparative

pseudospikelet and floral structure for this group of genera and it is premature still to

speculate if differences such as keeled or convolute palea, 2 versus 3 stigmas, presence

of glumes and number of flowers will con'elate with clades recognised from molecular

analyses, when these eventually benefit from sufficient taxon sampling that includes

nomenclatural types.

The consensus seems to be that Melocanna, Pseudostachyum and

Cephalostachyum could be distinguished from Schizostachyum, and the last could

probably moXn&Q Dendrochloa, Leptocanna, Neohouzeaua and Teinostachyum, as well

as some species currently placed in Cephalostachyum (such as C. pergracile and C.

virgatum). The present new species is consistent with the type of Schizostachyum,

S. blumei Nees, in having slender 1 -flowered pseudospikelets that terminate with a

rachilla extension bearing a temiinal vestigial flower, absence of glumes, convolute

paleas with 2 long-pointed tips and 3 stigmas. It differs from the type and many other

species of the genus in consistently having two lodicules (the type species does not

have lodicules, whereas a number of others have three, although variable numbers are

sometimes encountered).

The new species

Schizostachyum kidsingii K.M.Wong, sp. nov.

Schizostachyum kuisingii is similar to S. castaneum Widjaja, S. lutescens Widjaja and

S. mampouw Widjaja in having an erect clump habit, narrowly triangular to ovate-

lanceolate culm-sheath blades that are at first erect, and generally small culm-sheath

auricles with fine bristles on their margin. It differs from these three species in having

glabrescent leaf blades (the other species have persistently hairy leaf blades), minutely

serrate-ciliate leaf ligules (the other speeies have entire-glabrous leaf ligules), and two

Schizostachyum kuisingii, a new bamboo from Peninsular Malaysia

269

lodiciiles in its flower {S. lutescens and S. mampouw do not have lodicules; the flowers

of S. castaneum are not known). - TYPE: Peninsular Malaysia, Johor, Bekok, old

logging track past Kampung Tomoh and c. 10 km before the Selai gateway to Endau-

Rompin State Park, 26 November 2008, K.M. Wong, YW Low, Zulkapli Ibrahim &

K.S. Tan WKM 2896 (holotype SING; isotypes K, KLU). (Fig. 1-3)

Medium-size clumping bamboo to 18 m high, erect to slightly arching outward.

Culms plain green, 2. 5-5. 5 cm diameter; mid-culm intemodes to 40^5 cm long,

with scattered appressed pale hairs all over and a white-waxy zone below each node;

branches many at each node, slender and subequal, arising from a single branch

bud. Culm sheaths light green, with loose irritant stiff chestnut-brown hairs all over

the back; auricles low rim-like, c. 1 nim high, dark purplish black, with pale brown

bristles 14-22 mm long on the margin; ligule with a 1-1.5 mm high rim-like base with

2-3(-6) mm long bristles on the margin; blade narrowly triangular to ovate-lanceolate,

erect in the more basal sheaths, and erect becoming patent to reflexed at mid-culm or

higher, dark green. Foliage leaves 8-32 cm long, (1.4-)2.3-3.6 cm broad, adaxial

surface glabrous, abaxial surface sparsely pale minute-hairy becoming glabrescent;

auricles rounded to elongate lobes extending free of the sheath margin, 0.5-2 mm long

bearing bristles 2-5 mm long on their margin; ligules 1 mm high, minutely serrate-

ciliate. Pseudospikelets green, very slender, only 1-1.5 mm diameter, 10-14 mm

long, 1 -flowered with a rachilla extension 7-12 mm long bearing a terminal vestigial

flower; basal bracts subtending prophyllate buds 2-3, each 5-veined, the back with

veiy sparse short hairs. Flower with lemma 10-11 mm long, 9-veined, apical cusp 1

mm long, glabrous; palea 12-18 mm long, 9-veined, apex bifid with cusps 2-2.5 mm

long, glabrous; lodicules 2, obovate to oblanceolate, 2. 5-3. 5 mm long, 0.5-1 .5 mm

wide, with ciliate margin; stamens 6, filaments free; anthers 4—5 mm long, apex blunt,

maroon; ovary ovoid, c. 1 mm long, glabrous; style to 13-19 mm long, exserted c. 4

mm in the mature flower, rigid, glabrous, white; stigmas 3, hairy, white.

Etymology^ This new species honours Mr Tan Kui Sing, an experienced horticulturist

with wide interests in exploring the native Malaysian flora for novel candidate species

in tropical landscaping. He brought the existence of this new species to the attention

of the author.

Provisional lUCN conservation assessment. As far as is known, this species exists

only in the village at Bekok, Johor, Peninsular Malaysia. In spite of its novelty, its

status as an indigenous Malaysian bamboo is doubtful, as with a suite of other bamboo

species known only in cultivation or in association with human settlements, which

Holttum (1958) referred to as “village bamboos”. Because only one locality has been

documented, it seems appropriate to consider this species as “Data Deficient” in

conservation terms (lUCN, 2012).

Notes. Besides the key differences from its close congeners, Schizostachyum castaneum,

S. lutescens and S. mampouw (Widjaja, 1997), noted in the diagnosis above, there

270

Gard. Bull. Singapore 67(2) 2015

Fig. 1. Habit of Schizostachyum kuisingii K.M.Wong. (Photo: K.M. Wong)

Schizostachyum kuisingii, a new bamboo from Peninsular Malaysia

271

Fig. 2. Key vegetative eharacteristics of Schizostachyum kuisingii K.M.Wong. A. Detail of

eulm sheath showing loose brown hairs, spreading blade and bristly, dark rim-like auricles;

culm intemode with a covering of appressed white hairs, and a pale waxy zone just below the

node. B. Branch complement with a cluster of many slender subequal branches developing from

a solitary branch bud. C. Detail of leaf sheaths showing pale-bristly auricles. D. Pseudospikelet

clusters. (Photos: K.M. Wong)

272

Gard. Bull. Singapore 67(2) 2015

Fig. 3. Schizostachyum kuisingii K.M.Wong, pseudospikelet and flower structure. A.

Pseudospikelet with (from base) prophyll (pr), three bracts subtending prophyllate buds (br, br

2, br 3), a pseudospikelet branch (psb) developing from the bud subtended by br3, the lemma

(1) and palea (p) of the perfect flower with its protruding stiff style and stigmas. B. Bract. C.

Prophyllate bud. D. Lenmia. E. Apically bicuspidate palea. F. Gynoecium with a stiff style,

with six free stamens and two lodicules. G. Detail of blunt anther apex. H. Perfect flower with

lemma removed, revealing a slender rachilla extension with a terminal vestigial flower. Scale

bars represent 5 mm except for G (1 nun). Drawn by the author from WKI/1 2896 (SING).

Schizostachyum kuisingii, a new bamboo from Peninsular Malaysia

273

Table 1. Schizostachyum kuisingii K.M.Wong compared with closely related species.

A dash indicates feature not known for a particular species. The description of "hairy" ovaries

in various Indonesian species such as S. lutescens Widjaja and S. mampouw Widjaja (Widjaja,

1997) is probably erroneous, as the species known to the present author (including .S. kuisingii)

and described by others have invariably smooth (glabrous) ovaries and fruits.

S. kuisingii

S. castaneum

S. lutescens

S. mampouw

Midcuhn intemode

length (diameter)

(cm)

40-55

(2.5-5.5)

45-70

(4.5-6)

45-60

(2^)

30-60

(2.5-3)

Hair type on

intemode surfaee

White appressed

hairs only

White and brown

hairs

White hairs and

scattered brown

hairs

—

Hair type on back

of culm sheaths

Dense chestnut-

brown hairs

Dense chestnut-

brown hairs

Chestnut-brown

hairs

White to brown

hairs

Culm-sheath

auricles

Low rim only 1

mm high, bristles

14-22 mm long

Low lobes to 2 mm

high, bristles 4-1 1

mm long

—

Eow rim 1-2 mm

high, bristles 14

mm long

Culm-sheath

ligules

1-1.5 mm high

base, bristles 2-3

mm long, at times

to 6 mm long

1 imn high base,

denticulate (small 1

mm teeth)

1 mm high base,

bristles 2-3 mm

long

Culm-sheath blade

Narrowly

triangular to

ovate-lanceolate,

erect then patent to

reflexed

Narrowly triangular

to ovate-lanceolate,

erect

Narrowly

triangular to

ovate-lanceolate,

erect

Lower surface of

foliage leaves

Sparsely pale

minute-hairy,

glabrescent

Hairy

Sparsely hairy

Leaf sheath ligules

Minutely serrate-

ciliate,

1 mm high

Entire, glabrous,

1 mm high

Entire, glabrous,

1 mm high

Entire, glabrous,

1 mm high

Lemma length,

hairiness

10-11 mm

(shorter than

palea),

glabrous

c. 12 mm

{longer than

palea),

glabrous

9-10 mm

(shorter than

palea),

glabrous

Palea length,

hairiness

12-18 mm,

glabrous

c. 9

mm,

glabrous

10-15 mm,

glabrous

Lodicules

—

Anther colour

Maroon

—

Yellow

274

Gard. Bull. Singapore 67(2) 2015

are further distinctions (Table 1). Schizostachyum kuisingU culm intemodes have

only white appressed hairs, whereas those of S. castaneum and S. lutescens have a

mixture of white and brown hairs (those of S. mampouw have not been documented).

Schizostachyum castaneum also differs from S. kuisingU and S. mampouw in having

small lobe-like culm-sheath auricles (instead of rim-like auricles) and denticulate

culm-sheath ligules (instead of bristly ligules). In addition, Schizostachyum lutescens

differs from S. kuisingU and S. mampouw by its lenuuas longer than the paleas (in the

other two species, the lemmas are shorter than the paleas). Schizostachyum kuisingU

has maroon anthers, whereas S. lutescens and S. mampouw have yellow anthers.

ACKNOWLEDGEMENTS. I am grateful to Y.W. Low and Zulkapli Ibrahim for assistance in

the field. Pemiission to examine Schizostachyum material in the BO herbarium is gratefully

acknowledged. Dr Soejatmi Dransfield kindly looked over the material of this species in

providing additional verification of its novelty.

References

BPG (Bamboo Phylogeny Group) (2012). An updated tribal and subtribal classification of the

bamboos (Poaceae: Bambusoideae). In: Gielis, J. & Potters, G. (eds) The 9th World

Bamboo Congress Proceedings. Pp. 3-17. Massachusetts: World Bamboo Organisation.

Holttum, R.E. (1946). The classification of Malayan bamboos. J. Arnold Arbor. 27: 340-346.

Holttum, R.E. (1956). The classification of bamboos. Phytomorpholog\> 6: 73-90.

Holttum, R.E. (1958). The Bamboos of the Malay Peninsula. Gard. Bull. Singapore 16: 1-135.

lUCN (2012). lUCN Red List Categories and Criteria: Version 3. 1. 2nd ed. Gland, Switzerland

and Cambridge, UK: lUCN.

Kellogg, E.A. (2015). Poaceae. In: Kubitzki, K. (ed) The Families and Genera of Vascular

Plants. Vol. XIII. Heidelberg, New York, Dordrecht, London: Springer.

Thiers, B. [continuously updated]. Index Herbariorum: A global directory of public herbaria

and associated .staff. New York Botanical Garden’s Viitual Herbarium. http://sweetgum.

nybg.org/ih/ (accessed on 6 May 2015).

Widjaja, E.A. (1997). New taxa in Indonesian bamboos. Reinwardtia 1 1 : 57-152.

Wong, K.M. (1995). The Bamboos of Peninsular Malaysia. Malayan Forest Records No. 41.

Kuala Lumpur: Forest Research Institute Malaysia.

Xia, N.H. (1993). Studies on the genus Schizostachyum and other bamboos from China. J.

Prop. Subtrop. Bot. 1; 1-10.

Yang, H.-Q., Peng, S. & Li, D.-Z. (2007). Generic delunitations o^ Schizostachyum and its allies

(Gramineae: Bambusoideae) inferred from GBSSI and timL-F sequence phylogenies.

Taxon 56: 45-54.

Gardens’ Bulletin Singapore 67(2): 275-287. 2015

doi: 10.3850/S238258121500023X

275

Novitates Bruneienses, 4. New records in the Araliaceae,

Araucariaceae, Arecaceae, Fagaceae, Musaceae and

Thymelaeaceae

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

'Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, Singapore 259569

wkiTL2000@gmail.com

^Bioinei National Herbarium, Forestry Department,

Ministry of Industry and Primai 7 Resources, Jalan Menteri Besar,

Berakas, BB3910 Brunei Damssalam

ABSTRACT. Aralia merrillii C.B.Shang (Araliaceae), Agathis lenticula de Laub.

(Amicariaceae), Licuala collina Saw and L. iniriensis Saw (Arecaceae), Lithocarpus buUatus

Hatus. ex Soepadmo and L. hallieri (Seemen) A. Camus (Fagaceae), Musa lawitiensis var.

suratii (Argent) Hakkinen (Musaceae) and Aqiiilarla microcarpa Baill. (Thymelaeaceae) are

newly recorded for the Brunei flora.

Keywords. Angiosperms, Borneo, Brunei Darussalam, conservation, gymnosperms

Introduction

The Botanical Sun>ey of Brunei Darussalam is a collaborative programme between

the Government of Brunei Damssalam and the National Parks Board, Singapore, in

which the Ministry of Industry and Primary Resources and the Forestry Department in

Bmnei partner the Singapore Botanic Gardens to continue botanical exploration and

documentation in Bmnei. In this fourth instalment of the series dedicated to reporting

the scientific results of this collaboration, we record eight plant species new for the

Bmnei flora, in six different families. The baseline data, to assess whether a taxon is a

new record or not, comes from Coode et al. (1996) and later taxonomic works in the

particular groups. Conservation assessments for the species presented in this paper are

based on lUCN (2012), adopted with a regional focus on Bmnei. Herbaria acronyms

of herbarium specimens examined follow Thiers (continuously updated).

New records for Brunei

ARALIACEAE

Aralia merrillii C.B.Shang, J. Nanjing Inst. Forest. 1985(2): 28 (1985). - Aralia

scandens (Mem) Ha, Novosti Sist. Vyssh. Rast. 11: 229 (1974), nom. illeg. non A.

scandens Poir, Encycl. Suppl. 1:419(1811). - Acanthophora scandens Mem., Philipp.

276

Gard. Bull. Singapore 67(2) 2015

J. Sci., C. 13: 316 (1918). - TYPE; Philippines, Mindanao, Lake Lanao, Camp

Keithley, 1 October 1906, M.S. Clemens 752 (holotype PNH, destroyed; isotype A

[A00067728]). (Fig. 1)

Specimen examined. BRUNEI: Temburong: Ulu Temburong National Park, steep slope at LP

58, 1000 m asl, 6 Aug 2014, Wong & Jangarun WKM3363 (BRUN, SING).

This thorny scrambling-climbing species is known from the Malay Peninsula to the

Philippines and Sulawesi, and in Borneo has been recorded on Mt Kinabalu (Philipson,

1979; Beaman et al., 2001). The tripimiate leaves, up to 1.5 m long, and large panicle,

with the flowers or fruits in ultimate umbels, are distinctive. Besides that, Aralia

merrillii is also the only Aralia species in Malesia to have a scrambling-climbing

liana habit (Philipson, 1979). This species belongs to the section Aralia L. sect.

Dimorphanthus (Miq.) Miq., for which the presence of prickles is a synapomorphy

(Wen, 2004).

Provisional lUCN consen^ation assessment. Data Deficient (DD) (TUCN, 2012) for

Brunei as this is the only collection known thus far. Its typical occurrence is disturbed

or fringe habitats in montane localities, with the documented occurrence in the inner

mountainous region of the Ulu Temburong National Park. Field observations are badly

needed for a better understanding of its conservation status in Brunei.

ARAUCARIACEAE

Agathis lenticula de Laub., Blumea 25(2): 537 (1979). - TYPE: Sabah, Mt. Kinabalu,

Park Headquarters, 9 August 1978, de Laubenfels P619 (holotype L; isotypes A

[A00022582], K, RSA [RSA0000051], SAN, US [US00345172]). (Fig. 2)

Specimen examined. BRUNEI: Temburong: Ulu Temburong National Park, ridge W of LP 58,

1000 m asl, 6 Aug 2014, Wong & Jangarun WKM 3303 (BRUN, SING).

This stately conifer was previously known only for the Crocker Range, including

Mount Kinabalu, in Sabah, and is by no means rare on the ridges in this part of the

Ulu Temburong National Park. It is not expected to occur in the Tutong and Belait

districts because of the generally lower ten’ain there. The glaucous foliage, including

somewhat asymmetric leaf with acute apex and base, as well as short-pedunculate

male cones, are distinctive (Yii, 1995).

Provisional lUCN consei'vation assessment. Data Deficient (DD) (lUCN, 2012) for

Brunei as this is the first and only collection recorded so far. As the montane ridges of

the Ulu Temburong National Park are little explored, more specialised collections and

detailed field observations of the taxon are badly needed for a better understanding

of its conservation status in Brunei. Besides that, the species occurs within the Ulu

Temburong National Park, and there are no imminent threats known to be present.

New records for Brunei flora

277

Fig. 1. Aralia merrillii C.B.Shang (Araliaceae), Wong & Jangarun WKM 3363. (Photo: K.M.

Wong)

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Gard. Bull. Singapore 67(2) 2015

Fig. 2. Agathis lenticula de Laub. (Araucariaceae), the characteristic columnar bole with

scaly-dippled bark and (inset) glaucous leaves and male cones. Wong & Jangarun WKM 3303.

(Photos: K.M. Wong)

New records for Brunei flora

279

ARECACEAE

Licuala collina Saw, Kew Bull. 67(4): 599 (2012). - TYPE: Malaysia, Sarawak, Tama

Abu Range, Bario, 1250 m, 6 November 1989, Dayang Awa & Lee S 51191 (liolotype

K; isotype SAR). (Fig. 3A)

Specimens examined. BRUNEI: Belait: Sukang, Labi Hills Forest Reserve, east of LPUkur 16,

100 m asl, 24 Aug 1997, Iliga BRUN 18698 (BRUN); Ulu Buau, Kampoug Buau, 8 Mar 1999,

Ariffin BRUN 19100 (BRUN). Temburoiig: Ulu Temburong National Park, ridge E of LP 58,

c. 1000 m asl, 8 Aug 2014, PVong & Jangarun WKM3329 (BRUN, SING).

This was recorded as a lower montane forest palm and also with one occurrence in

kerangas forest at 1 00 m asl (Saw, 2012). It is now also recorded at the lower elevation

in Brunei in riparian Mixed Dipterocarp Forest. It would appear to have a general

occurrence in the highland areas on the Setap Shale formation of Brunei and NE

Sarawak and adjacent lower elevations.

This is a usually solitary palmlet 0.5-1. 5 m high. The inflorescence is typically

short, with a single partial inflorescence (unbranched or with few branches), without

peduncular or rachis bracts. There are 3-5 leaf segments (in Sarawak 4-10 recorded),

with the central segment broader and distinctly bifid, and frequently one of the halves

further bearing a narrow division.

Provisional lUCN conservation assessment. Data Deficient (DD) (lUCN, 2012)

for Brunei as the species is only recorded three times. However, all known Brunei

populations exist within protected areas, namely, the Ulu Temburong National Park

and Forest Reserves where disturbances are now curtailed. Meanwhile in Sarawak,

Licuala collina has been categorised as Vulnerable (VU) by Saw (2012).

Licuala miriensis Saw, Kew Bull. 67(4): 628 (2012). - TYPE: Malaysia, Sarawak,

Miri, Riam Road, 15 m, 3 December 1962, Au S 16794 (holotype SAR; isotypes K,

L). (Fig. 3B-D)

Specimen examined. BRUNEI: Belait: Lumut, Sungai Liang, Bukit Agis-Agis, 24 Dec 2003,

Ariffin BRUN 20467 (BRUN); Andulau Forest Reserve, Compartment 18, 13 Aug 2014, Wong,

Ariffin & Jangarun WKM 3392 (BRUN, SING).

This species was previously recorded only for the Miri-Lambir and Bintulu areas in

Sarawak (Saw, 2012). The two collections here record it for Brunei for the first time.

It is a distinctive, elegant, slender-stemmed understorey palm to 1.5-2 m high

with c. 20 narrow segments per leaf. Although Saw (2012) records 46 species of

Licuala for Bomeo, a number remain known only from scanty material, including

the present species, which was diagnosed as having an inflorescence specimen “with

proximal portion broken off’ and peduncle, prophyll and peduncular bracts unknown.

In the key provided in Saw (2012), Licuala miriensis is diagnosed through the lead

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Fig. 3. New Licuala records. A. Licuala collina Saw. Wong & Jangarun WKM 3329. B-D.

Licuala miriensis Saw. Wong, Ariffin & Jangarun WKM 3392. B. Habit. C. (from left)

Leaf blade, lower portion of petiole, infructescence. D. Detail of partial infructescence.

(Photos: K.M. Wong)

New records for Brunei flora

281

in couplet 2 giving “All partial inflorescences spicate...” and, indeed, adopting this

character helps derive the identity of the Brunei material as this species. However,

as the Brunei material demonstrates, the other lead in couplet 2 (“Proximal partial

inflorescence branched. . .”) should have been the correct grouping because that partial

inflorescence does have a solitary branch to 4 cm long {WKM 3392). This character

is sometimes easily missed, including in BRUN 20467 where that branch has broken

off and the remaining partial inflorescence looks spicate. [Following what should have

been the correct lead for this species in this key, however, one arrives at the cluster of

L. atrovirens Saw, L. mukahensis Saw, L. cordata var. ashtonii Saw and L. maculata

Saw, species that differ in many characteristics from L. miriensis. Incidentally, the

couplets 19 and 20 should have their numbers interchanged so as to match the correct

character-states.]

Slight discrepancies remain between the Brunei material and the description

in Saw (2012). The Brunei material has thin-membraneous to papei*y calyx lobes

and corolla lobes, both in fresh (fruiting) and dried material, whereas the available

description states that the calyx is “membranous... becoming thick” and the corolla

is “fleshy becommg very thick in fruiting specimen”. These are, however, minor

differences that probably point simply to some variation present.

This extremely elegant Licuala deserves introduction into horticulture although,

where we were able to observe this in Bmnei, it is a solitary palm occurring with very

low frequency in very moist shaded understorey conditions, and probably also slow-

growing. It is not known how this species might adapt and respond to an increased

light regime.

Provisional lUCN conservation assessment. Data Deficient (DD) (lUCN, 2012) for

Brunei as the species is only known from two localities so far. However, this status

needs reassessment for a better understanding of its conseiwation status in Brunei, as

more botanical explorations are made. Meanwhile, its probable status in Sarawak is

considered to be Endangered (EN A2c) (Saw, 2012).

FAGACEAE

Lithocarpus bullatus Hatus. ex Soepadmo, Reinwardtia 8: 223 (1970). - TYPE:

Borneo, Sabah, Mt. Kinabalu, 15 April 1933, J. & M.S. Clemens 32715 (holotype

BO; isotypes A [A00033959], BM [BM000951969], K [K000832515], L, NY

[NY00248587], UC). (Fig. 4A)

Specimen examined. BRUNEI: Temburong: Ulu Temburong National Park, ridge E of LP 58,

c. 1000 m asl, 8 Aug 2014, Wong WKM 3323 (BRUN, SING).

The type of this species is J. & M.S. Clemens 32715 from Mount Nungkok, a minor

peak of the Kinabalu massif, at 1200-1500 m elevation in lower montane forest

(Soepadmo, 1 970). It is a Bomean endemic not uncommon in lower to upper montane

forest to 3000 ni on Mount Kinabalu and the Crocker Range in Sabah, and has also

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Fig. 4. New Lithocarpus records. A. Lithocarpus bullatus Hatus. ex Soepadmo. Wong WKM

3323. B. Lithocarpus hallieri (Seemen) A.Camus. Wong & Jangarun WKM 3295. (Photos:

K.M. Wong)

been reeorded for the Mulu National Park (Sarawak), in Kalimantan (Indonesian

Borneo), as well as on ultramafic soil in Beluran (Sabah) (Soepadmo et ah, 2000).

In Brunei, it is locally common in the hills around 1000 m in southern Temburong.

The strongly bullate leaves are highly distinctive and no other Lithocarpus species

approaches in this character.

Provisional LUCN conservation assessment. Least Concern (LC) (lUCN, 2012) for

Brunei as there are good populations well within national parks, including the Ulu

Temburong National Park.

Lithocarpus hallieri (Seemen) A.Camus, Riviera Sci. 18: 40 (1932). - TYPE: Borneo,

Kalimantan, Mt. Lianggagang, Llallier 2655 (holotype BO; isotype L). (Fig. 4B)

Specimen examined. BRUNEI: Temburong: Ulu Temburong National Park, ridge W of LP 58,

1000 m asl, 6 Aug 2014, Wong & Jangarun WKM 3295 (BRUN, SING).

This Bornean endemic tree is generally known for the other Bornean states and can be

common in some localities, such as Ranau in Sabah.

The specimens were collected from a medium-sized tree to 35 m tall and 60 cm

diameter. The acorns have cupules enveloping most of the nut and are large, the size of

New records for Brunei flora

283

a fist (to 12 cm long, 10 cm diameter), obovoid and woody, with 5-7 distinct lamellae.

The nut itself is c. 5 cm diameter.

Provisional lUCN conservation assessment. Data Deficient (DD) (lUCN, 2012) for

Brunei as there is just a single collection, although several trees were seen on the

ridges around the collection site. However, this status requires reassessment for a better

understanding of its conservation status in Brunei, as more botanical explorations are

made, especially to the largely inaccessible hills at the Bmnei-Sarawak (Malaysia)

border.

MUSACEAE

Musa lawitiensis Nasution & Supard. var. suratii (Argent) Hakkinen, Adansonia

28(1): 60 (2006). - Musa suratii Argent, Gard. Bull. Singapore 52: 203 (2000). -

TYPE: Sabah, Tenom, Kallang, 8 September 1989, Surat & Lamb 268/89 (holotype

SAN; isotype E). (Fig. 5)

Specimens examined. BRUNEI: Temburong: Amo, Ulu Temburong National Park,

Temburong river just downstream from Kuala Machang, true left bank, old landslip, 4°32’09”N

1 1 5H 2 T 3”E, 1 00 m asl, 1 7 May 2(dU,Leeet al. SL 1018 (BRUN, SING, IB SC), 1 8 May 2014,

Wong et al. WKM3274 (BRUN, SING).

Borneo, well within the centre of diversity for bananas, has 1 3 known species excluding

the introduced Musa textilis Nee (Hakldnen, 2006). For Brunei, Coode et al. (1996)

recorded only Musa borneensis Becc. {Hotta 13877), M. campestris Becc. {Bernstein

225, Coode 6792, Cowley 14, Hotta 12460, Johns 6800), and M. tuberculata M.Hotta

{Hotta 13878), as well as the widely cultivated M. textilis {Hotta 13017).

Musa lawitiensis Nasution & Supardiyono, endemic to Borneo, has four varieties

(Hakkinen, 200i6).Musa lawitiensis var. lawitiensis is known in W Kalimantan (Bentuan

Karimun National Park) and Sarawak’s Kapit Division; M. lawitiensis var. kapitensis

and M. lawitiensis var. sarawakensis so far only from Sarawak’s Kapit Division; and

M. lawitiensis var. suratii from SW Sabah (Tenom), W Sarawak (Lubok Antu) and

now Brunei (Temburong). Argent (2000), when publishing the original description of

Musa suratii Argent, noted that this taxon was “clump fomfing” with suckers “vertical

in young clumps but becoming angled outwards in the larger older clumps”. He aptly

described the plant looking “from a distance more like a Heliconia than a Musa as, in

addition to the very slender habit, the leaves although actually spirally arranged are

displayed more or less distichously in a single plane”. Where we collected this plant

beside the Temburong River, a small population of plants were seen with pseudostems

typically inclined towards the river, so that the lower leaves were inserted spirally

but more apical leaves assumed a pseudo-distichous aiTangement: the sheaths showed

spiral insertion clearly but the petioles oriented the leaf blades into essentially two

rows, so that the banana had the habit of a large ginger from a distance. This habit

would seem to adapt the plant to a good display of leaf surface on a steep earth bank.

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Gard. Bull. Singapore 67(2) 2015

Fig. 5. Musa lawitiensis var. suratii (Argent) Hakkinen. Wong et al. WKM 3274. A. Habit.

B. Downarched inflorescence with slender fruits. C. Characteristics of the leaf blade and

inflorescence tip. (Photos: K.M. Wong)

New records for Brunei flora

285

111 the field, the leaning habit of this small slender banana (c. 3.5^ m tall), with

the narrow pseudo-distichous leaves that are richly waxy on their lower (abaxial) leaf

surfaces, a horizontal to arching inflorescence, small slender fruits (c. 10 cm long,

c. 1 .2 cm diameter), and very slender, orange-pink male bud with a sharp apex, are

highly distinctive. The seeds are tuberculate, characteristic of Musa section Callimusa,

and as many as 300 were counted for one fruit.

Provisional lUCN consei^ation assessment. Data Deficient (DD) (lUCN, 2012) for

Brunei as the species is only known from the Ulu Temburong National Park, represented

by two collections made on the same botanical fieldwork. Further field observations

will contribute to a better understanding of its conservation status in Brunei.

THYMELAEACEAE

Aquilaria microcarpa Bail!., Adansonia 11: 304 (1875). - TYPE: Borneo, Sarawak,

1872, Beccari PB 2886 (holotype FI [FI008102]; isotypes G [G00190871], L

[L0010148], M [M0145881], NY [NY00386214], P [P00650666], S [S-G-553]). (Fig. 6)

Specimens examined. BRUNEI: Belait: Bukit Sawat, Jalan Meranlcing, 26 Jul 1997, Ogata

Og-B 519 (BRUN); Kampong Merangking, 20 Dec 1994, Suhaili BRUN 16407 (BRUN);

ibidem, 23 Jul 1999, Joffre BRUN 18367 (BRUN); Kampong Singap, Jalan Singap-Bukit, 24

Jun 1997, Said BRUN 18496 (BRUN); Sungai Liang, Arboretum, 7 Aug 1996, Clayton BRUN

7657<5(BRUN); ibidem, 19Aug \991 , Ariffin BRUN 1 8643 (BRUN). Brunei-Muara: Kilanas,

Jalan Dadap waterfall, 16 Jun 1998, Noor Azam BRUN 18854 (BRUrN), Temburong: Batu

Apoi, 23 Apr 1935, Zainal KEP 30376 (BRUN, KEP); Bukit Patoi, 1959, Ashton Voucher

3689 (BRUN); Kuala Belalong, 15 May 2014, Lee et al. SL 974 (BRUN, SING). Tutong:

Rambai, Ulu Sungai Tutong, Ulu Sungai Medit, 14 Aug 2009, Yusop BRUN 22684 (BRUN, K,

KEP, L, SAN, SAR, SING); Tasik Merimbun, Bang Oncom, 19 Nov 1992, Bernstein JHB 358

(BRUN); Bukit Tangan, 16 Sep 2000, Suzuki K 13261 (BRUN, KAG).

An early checklist by Hasan & Ashton (1964) listed Aquilaria malaccensis Lamk.,

which, although recorded for N and E Borneo (Hou, 1960), does not seem to be present

in Brunei. The later checklist by Coode et al. (1996) confinned the presence in Brunei of

Aquilaria beccariana Tiegh. but \Qft Bernstein JHB 358 (leafy twig only) unidentified.

Aquilaria microcarpa is now verified by a number of collections of both flowering and

fruiting specimens: the perianth is bell-shaped with lobes less than or equal the tube

length and the fmit is subcordate and up to c. 1.5 cm long only {A. beccariana has

floral tubes much longer than the lobes, and a longer obovoid fruit). Furtheraiore, its

leaves diy olive to medium brown, whereas those of Aquilaria beccariana dry a pale

golden brown. The latter species is more typical of kerangas, peatswamp and riverine

(alluvial) forests in Mixed Dipterocarp Forest, whereas Aquilaria microcarpa is more

generally recorded from Mixed Dipterocarp Forest on sandy clays.

286

Gard. Bull. Singapore 67(2) 2015

Fig. 6. Aquilaria microcarpa Baill. Flowering leafy branch. Lee et al. SL 974. (Photo: K.M.

Wong)

Provisional lUCN conservation assessment. Least Concern (LC) (lUCN, 2001) for

Brunei as the species is reasonably well collected with populations existing in the Ulu

Temburong National Park, and other Forest Reserves and conservation areas, where

logging is not permitted.

ACKNOWLEDGEMENTS. The Ministry of Industry and Primary Resources and the Forestry

Department, Bmnei Darussalam, and the National Parks Board and Botanic Gardens, Singapore,

supported this work as part of their collaboration for A Botanical Survey of Brunei Darussalam.

Field coordination and logistics were assisted through Noralinda bind Haji Ibrahim, Roslinah

binti Haji Mohsin (Forestry Department), Watu bin Awok, Jangamn anak Eri and Azlan bin

Pandai (BRUN); Serena Lee and Paul Parusuraman Athen (SING). We are grateful to the

keepers and curators of the BO, BRUN, SAN, SAR and SING herbaria for facilitating study of

specimens in their care. Specimen images were also accessed in Global Plants (continuously

updated).

New records for Brunei flora

287

References

Argent, G. (2000). Two interesting wild Musa species (Musaceae) from Sabah, Malaysia.

Card. Bull. Singapore 52: 203-210.

Beaman, J.H., Anderson. C. & Beaman, R.S. (2001). The Plants of Mount Kinabalu 4.

Dicotyledon Families Acanthaceae to Lythraceae. Kota Kinabalu: Natural History

Publications (Borneo). (Araliaceae, Pp. 119-126)

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

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

Ministry of Industry and Primai 7 Resources.

Hakkinen, M. (2006). Musa lawitiensis Nasution & Supard. (Musaceae) and its intraspecific

taxa in Borneo. Adansonia 28(1): 55-65.

Hasan bin Pukol & Ashton, P.S. (1964). A Check List of Brunei Trees. Reprinted 1988. Brunei

Darussalam: Forestry Department.

Hou, D. (1960). Thymelaeaceae. In: Van Steenis, C.G.G.J. (ed) Flora Malesiana, Series I, 6:

1-48.

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

and Cambridge, UK: lUCN.

Global Plants (continuously updated). JSTOR® Global Plants. ITHAKA https ://plants.jstor.

org/ (accessed on 22 May 2015).

Philipson, W.R. (1979). Araliaceae — I. In: Van Steenis, C.G.G.J. (ed) Flora Malesiana, Series

1,9(1): 1-105.

Saw, L.G. (2012). A revision of Licuala (Arecaceae, Coryphoideae) in Borneo. Kew Bull. 67:

577-654.

Soepadmo, E. (1970). Florae Malesianae Praecursores XLIX: Malesian species of Lithocarp us

Bl. (Fagaceae). Reinwardtia 8: 197-308.

Soepadmo, E., Julia, S, & Go, R. (2000). Fagaceae. In: Soepadmo, E. & Saw, L.G. (eds) Tree

Flora of Sabah and Sarawak v 3: 1-117. Kuala Lumpur: Sabah Forestry Department,

Forest Research Institute Malaysia & Sarawak Forestry Department.

Thiers, B. (continuously updated). Index Herbariorum: A global directory of public

herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium.

http://sweetgum.nybg.org/ih/ (accessed on 31 May 2014).

Wen, J. (2004). Systematics and biogeography of Aralia L. sect. Dimorphanthus (Miq.) Miq.

(Araliaceae). Cathaya 15-16: 1-187.

Yii, P.C. (1995). Araucariaceae. In: Soepadmo, E. & Wong, K.M. (eds) Tree Flora of Sabah

and Sarawak 1: 27-32. Kuala Lumpur: Sabah Forestry Department, Forest Research

Institute Malaysia & Sarawak Forestry Department.

Gardens’ Bulletin Singapore 67(2): 289-296. 2015

doi: 10.3850/S2382581215000241

289

Three new species of Loxocarpus (Gesneriaceae)

from Sarawak, Borneo

T.L. Yao

Forest Research Institute Malaysia,

52109 Kepong, Selangor, Malaysia

yaotzeleong@frim.gov.my

ABSTRACT. Three new species, Loxocarpus burttii T.L.Yao, Loxocarpus littoralis T.L.Yao

and Loxocarpus segelamensis T.L.Yao are described from Sarawak, Malaysia.

Keywords. Borneo, Gesneriaceae, Loxocarpus, new species, Sarawak

Introduction

Loxocarpus R.Br. is a small genus consistmg of 20 species distributed in Sumatra,

Peninsular Thailand, Peninsular Malaysia and Borneo. The genus was previously

included in Henckelia Spreng. (Weber & Burtt, 1998; Banka & Kiew, 2009) but was

reinstated by Middleton et al. (2013) based on earlier phylogenetic studies (Weber

et al., 2011). The species of Loxocaij}us from Peninsular Malaysia were revised (as

Henckelia sect. Loxocarpus) by Banka & Kiew (2009), although since then Yao et al.

(2012) have described a new species from Kelantan. Loxocarpus is most diverse in

Borneo, represented by nine species, and here a fiirther three species are described.

All new species described here have a relatively long capsule (10-17.5 mm

long). Loxocarpus was principally characterised by a short plagiocarpic capsule less

than 10 mm long (Banka & Kiew, 2009). However, molecular phylogenetic studies

(Yao, 2012) support the inclusion of several long-capsuled species which had been

placed in Didymocarpus Wall, and Henckelia.

Loxocarpus burttii and L. segelamensis belong to the ‘saintpaulioid’

morphological group (Yao, 2012) which is confined to Borneo. Species in this group

resemble Saintpaulia H.Wendl. (Gesneriaceae) and are characterised by their flat-

faced corolla with a pair of exposed stamens. The short campanulate corolla with

deltoid lobes observed in L. littoralis is shared with L. argenteus B.L.Burtt and L.

pauzii T.L.Yao. All three new species are known only from a single locality each and

L. segelamensis and L. littoralis are represented only by the type specimen.

New species

Loxocarpus burttii T.L.Yao, sp. nov.

This species is similar to Loxocarpus verbeniflos (C.B. Clarke) B.L.Burtt in its

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Gard. Bull. Singapore 67(2) 2015

relatively long capsules (10-12 mm long vs 9-20 mm long) and its usually ovate or

elliptic lamina, but differs in its buff trichomes (not silvery) and adpressed tricbomes

along the petiole (vs spreading in Z, verbenifios). - TYPE: Borneo, Sarawak, Miri

District, Lambir Hills National Park, below Bkt. Lambir, sandstone cliff, c. 1500 feet,

24 September 1978, Burtt B11597 (holotype E; isotypes KEP, SAR). (Fig. 1, 2)

Rosulate herb with numerous crowded leaves. Rootstock woody, 2^(-9.5) cm long,

4-6 mm thick; trichomes buff, adpressed, dense; adventitious roots long, wiry. Petiole

base of withered leaves persistent. Stem lacking or distinctive, 1-2 cm long. Leaves

alternate, crowded at the top; trichomes dense, buff, adpressed on petiole, equally

dense on lamina above and beneath. Petiole relatively thick, (0.8-)2.8-3.8 cm long,

0. 7-1.1 mm diam., grooved above; longest petiole about equal in length to the lamina.

Lamina dark green above, paler beneath, moderately thick, ovate or elliptic or obovate,

(2.5-)3.3-4.1 X (0.8-)l .3-1.5 cm; base cuneate, equal, margin entire or undulate,

apex acute or blunt; midrib and veins obscure above, inconspicuous beneath, lateral

veins 3 pairs. Inflorescence axillary, a reduced cyme with a single flower; trichomes

buff, eglandular, shaggy, dense on peduncle and bracts; mixed with shorter glandular

trichomes on pedicel. Peduncle slender, 4.5-6. 5 cm long; bracts narrowly lanceolate,

c. 1.5 X 0,3 mm, apex blunt; pedicel 0.5-0. 9 mm long. Flower trichomes buff and

eglandular, shaggy and dense on the outer surface of the calyx, less dense, shorter

and erect on the outer surface of the corolla; dense on the ovary. Calyx green, upper

lip 3-lobed, lower lip 2-lobed; lower lobes larger than the upper lobes, tube c. 0.6

mm long, c. 1 mm wide, lobes deltoid, apex acute, c. 1.4 mm long, base c. 0.5 mm

wide. Corolla mauve, flat-faced, tube very short, c. 0.6 mm long, c. 1.1 mm wide;

upper lip with 2 lobes c.. 2.5 x 2.5 mm, lower lip with 2 lateral lobes c. 2.3 x 2 mm

and a median lobe c. 3 x 2.3 mm; lateral and median lobes oblong. Fertile stamens 2,

yellow; anthers projecting beyond corolla tube, exposed; filaments straight, thickened

in distal half, c. 2.2 mm long; anthers coherent only at the ventral tip, kidney-shaped, c.

1.7 X 0,7 mm; staminodes 3, club-shaped. Nectary absent. Ovary narrowly conical or

slightly oblique, c. 1.8 mm long, c. 1 mm wide; style curved upwards near the stigma,

c. 3 mm long; stigma capitate, exposed. Capsule long slender conical, 10-12 mm

long, 1. 1-1.4 mm wide, style c. 1.5 nun long, sub-persistent; valves straight, splitting

dorsally to form a trough; trichomes a mix of eglandular and glandular trichomes,

adpressed to erect. Seeds not seen.

Distribution. Borneo, Sarawak, known only from Lambir FR, Bukit Lambir.

Habitat. Hill forest at c. 460 m asl. On sandstone cliffs or on acidic sandy soil.

Etymology. This species is named after the collector of the type, B .L. Burtt (191 3-2008),

a British botanist who collected extensively and is well known for his contributions to

the taxonomy of the Old World Gesneriaceae.

Three new Loxocarpus from Borneo

291

Fig. 1. Distribution map of Loxocarpus burttii T.L.Yao (•), L. littoralis T.L.Yao (■) and L.

segelamensis T.L.Yao (A).

Additional specimens examined. MALAYSIA: Sarawak: Bukit Lambir, 23 Oct 1993, yt// & La

Frankie AI 191 (SING [SING 106370]); ibidem, 25 Oct 1993, Ali & La Frankie AI 192 (SING

[SING106371]); ibidem, 2 Nov 1976, Ilias & Yeo S.38365 (K, KEP [KEP60951], SAR).

Note. A handwritten annotation by Burtt on the folder of the specimen selected here

as the holotype mentioned that it is possibly a new species and listed S.38365 as

conspecific.

Fig. 2. Loxocarpus burttii T.L.Yao. A. Habit. B. Leaf. C. Corolla opened to show stamens.

D. Bracts, calyx, ovary, style and stigma. E. Stamens. F. Staminode. Drawn by N. Mohamad

Aidil from (A, B) Burtt B11597, and (C-F) Bias & Yeo S. 38365.

Three new Loxocarpus from Borneo

293

Loxocarpus littoralis T.L.Yao, sp. nov.

This species has a relatively long capsule similar to that of Loxocarpus violoides

(C.B. Clarke) T.L.Yao (17 mm long vs 15-30 mm long) but differs in the dense

spreading woolly trichomes on the lamina upper surface (not thin adpressed), grooved

upper surface of the petiole (not terete), deltoid corolla lobes (not oblong) and conical

capsule (not cylindric). - TYPE: Borneo, Sarawak, Kuching District, Tanjung Po,

in deep shade on dripping wet rocks near the sea, 5 October 1955, Brooke 10614

(holotype L [L645488]). (Fig. 1)

Rosulate herb. Rootstock woody, to 1 cm long, c. 3 mm thick or lacking; rootstock with

buff trichomes; adventitious roots long, wiry. Petiole base of withered leaves caducous.

Stem lacking. Leaves alternate, crowded at the top of the rootstock; trichomes buff,

woolly and shaggy, dense on lamina above and less dense beneath. Petiole slender,

c. 2 cm long, c. 1 mm thick, grooved above; longest petiole shorter than the lamina.

Lamina moderately thick, ovate or broadly ovate, c. 2.5 x 1 .7 cm; base cuneate or

rounded, equal, margin shallowly serrate, apex blunt; midrib and veins obscure above,

distinct beneath, lateral veins 3 pairs. Inflorescence axillary, a reduced cyme with a

single flower; trichomes buff, shiny. Peduncle slender, c. 5 cm long; bracts narrowly

lanceolate, c. 3 x 0.5 mm, apex acute; pedicel c. 3 mm long. Flower: corolla violet,

tube short with deeply dissected lobes; upper lip with 2 upper lobes, lower lip with 2

lateral lobes and a median lobe; upper lobes smaller, lateral and median lobes larger,

lobes recurved, deltoid. Capsule long, slender conical, c. 17 mm long, c. 2.5 mm wide;

valves straight, splitting dorsally to form a trough. Seeds not seen.

Distribution. Borneo, Sarawak, known only from the type specimen from Tanjung Po

(Bako NP).

Habitat. On dripping wet rocks near the sea in deep shade.

Etymology. It is named for its seashore habitat.

Note. There is only one fully opened flower on the type specimen which I decided not

to dissect, thus preventing a fuller description of the calyx and corolla lobes. More

material is needed.

Loxocarpus segelamensis T.L.Yao, sp. nov.

This species is similar to Loxocarpus violoides in the relatively long capsule (17.5

mm vs 15-30 mm long) and in the ovate lammas, but differs in its buff trichomes (not

silvery), the longer lamina (c. 7.4 cm long vs up to 5.5 cm), and the presence of c. 10

inflorescences in flower at any one time, with each inflorescence bearing 3-7 flowers

(vs 1-2 inflorescences per plant, bearing 1 or rarely 2 flowers). - TYPE. Borneo,

Sarawak, Manidi District, Baram, Sungai Segelam, around Long Selatong Lepo Ga’,

on rocky ridge, 24 July 1977, Chin 2797 (holotype KLU [KLU29970]). (Fig. 1, 3)

Three new Loxocarpus from Borneo

295

Creeping herb with buff, shiny trichomes, stem decumbent. Rootstock c. 6 cm long,

c. 2 mm thick; trichomes adpressed, dense; adventitious roots wiry. Petiole base of

withered leaves persistent. Erect stem distinct, c. 8 cm long. Leaves alternate, below

well-spaced, crowded at the top; trichomes adpressed, dense on petiole, on lamina

above and beneath dense. Petiole slender, c. 6 cm long, terete; longest petiole about

equal in length to the lamina. Lamina dark green above, paler beneath, thin, ovate, c,

7.4 X 2.9 cm; base cuneate, equal, margin finely semilate, apex acuminate or acute;

midrib and veins obscure above, distinct beneath, lateral veins 4(-5) pairs, lowermost

pair arising Ifom lamina base. Inflorescences axillary, a reduced cyme with a single

flower or once branched, c. 10 per plant, flowers 3-7; trichomes eglandular, adpressed

on peduncle, bracts and pedicel. Peduncle slender, 1 0.5-1 1 .5 cm long; bracts linear, c.

4.4 X 0.4 mm, apex acuminate; pedicel to 1 .2 nun long. Flower trichomes eglandular,

adpressed on calyx, outer surface of the corolla and ovary, dense. Calyx green, upper

lip 3-lobed, lower lip 2-lobed; lower lobes larger than the upper lobes, tube c. 1 mm

long, c. 1.9 mm wide, lobes linear-lanceolate, apex thickened, blunt, c. 3.9 mm long,

base c. 1 mm wide. Corolla pale blue, flat-faced, tube veiy short, c. 1.8 mm long,

c. 2.9 mm wide; upper lip with 2 lobes c. 7 x 3.8 nun, divided to the base, lower

lip with 2 lateral lobes, c. 5.6 x 3.5 mm and a median lobe c. 4.8 x 3 mm, lobes

deeply divided; lateral and median lobes oblong. Fertile stamens 2, yellow; anthers

projecting beyond the corolla tube, exposed; filaments straight, thickened in the distal

half, c. 4.5 nmi long; anthers coherent only at the ventral tip, kidney-shaped, c. 2.5 x

1.4 mm; staminodes 3, the two lateral club-shaped, the central a minute protrusion.

Nectaiy absent. Ovary conical, slightly oblique, c. 2.8 mm long, c. 1 .8 mm wide; style

straight, c. 2.3 mm long; stigma capitate, exposed. Capsule slender cylindric, c. 17.5

mm long, c. 1.9 mm wide, style sub-persistent; valves straight, splitting dorsally to

form a trough; trichomes eglandular, adpressed, dense. Seeds not seen.

Distribution. Borneo, Sarawak, known only from the type specimen Irom Ulu Long

Selatong.

Habitat. On rocky ridge.

Etymology. It is named for its type locality, Sungai Segelam.

ACKNOWLEDGEMENTS. These new species were discovered during the course of iny

studies for a Master of Science degi'ee at the University of Malaya, Kuala Lumpur, Malaysia.

I am grateflil to R. Kiew (KEP) and N.W. Haron (University of Malaya) for their guidance

throughout my study. T thank the Training Committee of the Forest Research Institute Malaysia

for sponsoring my study for the first two years, and the Institute of Research Management

and Monitoring, University of Malaya for the provision of a research grant (PS173-2008B).

I gratefully acloiowledge the curators at E, K, KEP, KLU, L, SAN, SAR, SING and SNP for

granting me permission to examine specimens in their care. I thank R. Kiew, D.J. Middleton

and an anonymous reviewer for their constructive comments on the manuscript.

296

Gard. Bull. Singapore 67(2) 2015

References

Banka, R.A. & Kiew, R. (2009). Henckelia section Loxocarpus (Gesneriaceae) in Peninsular

Malaysia. Edinburgh J. Bot. 66: 239-261.

Middleton, D..T., Weber, A., Yao, T.L., Sontag, S. & Moller, M. (2013). The current status of the

species hitherto assigned to Henckelia (Gesneriaceae). Edinburgh J. Bot. 70: 385^04.

Weber, A. & Burtt, B.L. (1998 [‘1997’]). Remodelling of Didymocarpus and associated genera

(Gesneriaceae). Beitr. Biol. Pflanzen 70: 293-363.

Weber, A., Middleton, D. J., Forrest, A., Kiew, R., Lim, C.L., Rafidah, A.R., Sontag, S., Triboun,

R, Wei, Y.G., Yao, T.L. & Moller, M. (2011). Molecular systematics and remodelling of

Chirita and associated genera (Gesneriaceae). Taxon 60(3): 767-790.

Yao, T.L. (2012). A Taxonomic Revision of Loxocarpus (Gesneriaceae). Unpublished M.Sc.

thesis. Kuala Lumpur: University of Malaya.

Yao, T.L., Kiew, R. & Haron, N.W. (2012). Loxocarpus pauzii (Gesneriaceae), a new species

from Peninsular Malaysia. Blumea 57: 134-135.

Gardens’ Bulletin Singapore 67(2): 297-303. 2015

doi: 10.3850/S2382581215000253

297

Studies on Begonia (Begoniaceae) of the Moluccas II:

a new species from Seram, Indonesia

W.H. Ardi> & D.C. Thomas"

^Bogor Botanic Gardens, Jl. Tr. H. JuandaNo. 13, Bogor, Indonesia

wisn001@lipi.go.id

^Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, Singapore 259569

daniel_thomas@nparks . gov. sg

ABSTRACT. A new species of Begonia L., Begonia galeolepis Ardi & D.C. Thomas, is

described from Seram, Maluku province, Indonesia. The species is endemic to Seram and

belongs to Begonia section Petermannia. An identification key to the seven Begonia species

known from the Moluccas is provided.

Keywords. Begonia galeolepis, Maluku Islands, Manusela National Park

Introduction

Our understanding of the Begonia L. flora of the Moluccas (also laiown as the Maluku

Islands), an archipelago within Indonesia located between the islands of Sulawesi and

New Guinea, is very limited because of the paucity of herbarium collections fi*om the

region and a lack of alpha-taxonomic baseline work on eastern Malesian Begonia.

The Begonia Resource Centre (Hughes et al., 2015) comprises records of only 138

Moluccan Begonia specimens, most of which are not identified to species level, and the

rest belongmg to six species (Ardi et al., 2014; see identification key to the Moluccan

species below).

Recent expeditions by Bali Botanic Garden, and a joint expedition between

Bogor Botanic Gardens and Fairchild Tropical Botanic Gardens to major islands of the

Moluccas (Halmahera, Seram and Temate), have brought to light material of several

Begonia species that do not confomi with any species previously reported from the

archipelago. Three endemic species have recently been described (Wiriadinata, 2012;

Ardi et al., 2014) from material collected on Halmahera, the largest island of the

Moluccas, raising the total number of Moluccan species to six (Ardi et al., 2014). From

the second largest island, Seram, Hughes (2008) reported only a single species, Begonia

brachybotjys Meir. & L.M.Perry. This species belongs to the Begonia rieckei Warb.

species complex, which shows a wide distribution east of Wallace’s Line, However,

given (i) Seram’s substantial area (c. 17,100 kiiF) and topographical heterogeneity,

(ii) the presence of diverse and suitable ecosystems including lowland rain forests and

limestone hills, as well as (iii) the species-richness and prevalence of narrow endemics

in Begonia (Hughes & Hollingsworth, 2008), this is likely to be an underestimation.

298

Gard. Bull. Singapore 67(2) 2015

Hughes (2008) already noted that even the very limited herbarium material indicated

that a number of species remain to be described from the archipelago,

A new species is described below from material collected on Seram and

cultivated in Bali Botanic Garden and Bogor Botanic Gardens. The species is very

distinct because of the presence of unusual appressed, fleshy and branched red scales

on the stems and leaves (Fig. 1, 2). In common with the majority of Moluccan species,

it belongs to Begonia sect. Petemiannia (Klotzsch) A.DC., which is distinguished by

protogynous inflorescences, two-flowered female inflorescences or solitary female

flowers, three-locular ovaries with axile placentation and bilamellate placentae, fruits

with equal or subequal wings, and anthers with unilaterally positioned slits (Doorenbos

et ah, 1998). All available Begonia specimens from BO, E, K, L and SING have been

consulted, and hence it must be assumed, at least until more intensive collecting reveals

otherwise, that this species has a restricted range and is endemic to Seram (Fig. 3).

Begonia galeolepis Ardi & D.C. Thomas § Petermannia, sp. nov.

Species resembling Begonia holosericeoides Ardi & D.C. Thomas in the creeping

habit and leaf shape, but differs consistently by the shorter peduncles of the female

inflorescences (up to 5 mm long), the sparse to moderately dense indumentum of

fleshy, branched, appressed red scales on stems, petioles and abaxial leaf lamina veins,

and the very long pedicels of male flowers (up to 1 1 cm). In conti'ast, in Begonia

holosericeoides the peduncles of the female inflorescences are 1.1 -3. 5 cm long, the

petioles and stems show a dense indumentum of branched pinkish hairs, and the

pedicels of the male flowers are 1.5-4 cm long. - TYPE: Indonesia, Seram, Manusela

National Park, cultivated in Bogor Botanic Gardens, 19 May 2015 (voucher made

from cultivated material), Wisnu Ardi 102 (holotype BO; isotypes Herbarium of Bali

Botanic Garden, KRB, SING) (Fig. 1-3).

Perennial, monoecious herbs, stems initially semi-erect, but prostrate and creeping

in older plants, rooting at the nodes when in contact with the substrate, up to c. 40

cm long; stems, primai*y veins on the abaxial stipule surfaces, petioles, and primary

and secondary veins on the abaxial leaf lamina surfaces with sparse to moderately

dense indumentum of multicellular, fleshy, appressed, red scales up to 6 x 4 mm, the

larger ones branched in the distal part, and all above-ground vegetative parts with

microscopic glandular hairs. Stem branched; intemodes 0.8-5. 5 cm long, up to 1 cm

in diameter, terete, green. Leaves alternate; stipules persistent, 1 .8-2.5 x 1-2 cm, ovate

to triangular, acuminate, setose, seta to 16 mm long, margin entire and sometimes

slightly revolute, green to reddish with small paler spots, translucent at the margins;

petioles c. 7-22 cm long, adaxially deeply channelled, with red scales fonning a

ring at the petiole-lamina transition; lamina basifixed, 16-23.8 x 12.5-17.2 cm, very

asymmetric, broadly ovate to suborbicular, base cordate and lobes not or sometimes

slightly overlapping, apex shortly acuminate, margin distantly dentate, the teeth bristle-

pointed, adaxial surface shiny reddish dark green with bright green veins, glabrous,

abaxial surface pale reddish green, primary veins 7-9, actinodromus, secondary veins

craspedodromus. Inflorescences protogynous; female inflorescences 1-2-flowered,

Begonia galeolepis, a new species from Moluccas

299

one node basal to male inflorescences, peduncles up to 5 mm long; male inflorescences

composed of 2-3 monochasial partial inflorescences, each monochasium with 2-A

flowers; bracts ovate to elliptic, 10-20 x 7-10 mm, green to red, with an abaxially

prominent midrib and a sparse indumentum. Male flowers: pedicels 4-11 cm long,

glabrous or glabrescent; tepals 2, broadly ovate, 9-18 x 11-15 mm, base slightly

cordate, margin entire, apex rounded, white or white tinged with pink at the margin,

abaxially sparsely hairy; androecium of 45-51 stamens, yellow, filaments 1-2 mm

long, slightly fused at the very base, anthers 1-1.5 mm long, obovate, dehiscing

through unilaterally positioned slits c. !/2 as long as the anthers. Female flowers:

pedicels 1.5^ cm long, sparsely hairy, green-reddish; tepals 5(-6), white tinged with

pink, unequal, the outer four larger, 9-15 x 6-11 mm, obovate, the inner one smaller,

6-15 X 4-5 mm, obovate, abaxially sparsely hairy to glabrescent; ovary obovoid, c.

10-15 X 6-8 mm (excluding the wings), green, glabrous or sometimes sparsely hairy,

locules 3, placentation axile, placentae bilamellate, wings 3, red, base slightly rounded

or cuneate, apex truncate, style basally fused, 3 -branched, each stylodium bifurcate in

the stigmatic region, stigmatic surface a spirally twisted papillose band, orange. Fruits

on up to 5 cm long pedicels, seed bearing part obovoid, up to c. 17 x 10 mm (excluding

the wings), sparsely hairy, dehiscent, splitting along the wing attachment, wing shape

as for ovary, up to 11 mm at the widest point (subapically). Seeds unknown.

Fig. 1. Branched trichomes and scales in Begonia sect. Petermannia. A-B. Begonia galeolepis

Ardi & D.C. Thomas, appressed, branched red scales. C-D. Begonia holosericeoides Ardi &

D.C. Thomas, branched trichomes. E-F. Begonia ozotothrix D.C. Thomas, branched trichomes.

Scale bars: A-F = 2 mm. (Photos: Wisnu H. Ardi)

300

Gard. Bull. Singapore 67(2) 2015

Fig. 2. Begonia galeolepis Ardi & D.C. Thomas. A. Growth habit in cultivation. B. Red scales

on stem. C. Red scales on the petiole and abaxial leaf surface. D. Stipules. E. Bracts. F. Male

inflorescence. G. Female inflorescence. H. Fruit. I. Male flowers. J. Female flower. K. Cross-

section of ovary showing placentation. Scale bars: A = 20 cm, B, C, K = 5 mm, D, E, G-J = 1

cm, F = 4 cm. (Photos: Wisnu H. Ardi)

Begonia galeolepis, a new species from Moluccas

301

Fig. 3. Distribution of Begonia galeolepis Ardi & D.C.Thomas. Collection sites are indicated

by a circle (Manusela National Park), a square (Japoetih) and a triangle (Buria). Specimen

location infonnation was georeferenced using the GeoNames geographical database at http://

www.geonames.org/.

Habitat. Primary lowland rainforest, growing on vertical moist limestone walls, at

river embankment, in half-shade to fiilly sun-exposed, at c. 20 m altitude.

Distribution. Endemic to Seram, Maluku Province, Indonesia. Locally common.

Etymology’. The specific ephithet is derived from the classical greek (“ya^^eog”

[galeos^^ dogfish, small shark, and “}i87iLg” [lepis]= scale) and refers to the fleshy red

scales on the petioles and stems, whose shape resembles shark scales (Fig. 1).

Notes. Begonia galeolepis is distinct within section Petennannia by a combination of

several characters such as the creeping stem, the characteristic red scales on stems and

leaves, few-flowered monochasial male inflorescences, and the very long pedicels of

the male flowers (up to 1 1 cm long). Branched hairs are rare in Asian Begonia, and

have been only described from two other species in section Petermannia, Begonia

holosericeoides Ardi & D.C.Thomas from Halmahera (Ardi et al, 2014) and B.

ozotothrix D.C.Thomas from Sulawesi (Thomas et al., 2009). The branched hairs

of Begonia ozotothix are much smaller, white and translucent and rounded in cross-

section, and the hairs of B. holosericeoides are thinner, pinkish and show fllifonn apices

in comparison to the thick, appressed red scales of B. galeolepis (Fig. 1). These scales

rather resemble hairs found at the lamina-petiole transition in the Sumatran species

Begonia sublobata Jack which, however, is otherwise morphologically dissimilar

and belongs to section Reichenheimia (Hughes & Girmansyah, 2011). Similar hairs

302

Gard. Bull. Singapore 67(2) 2015

are also found in some species of the American Begonia section Gireoudia (e.g. B.

heracleifolia Cham. & Schltdh), which is only very distantly related to Asian Begonia

(Moonlight et ah, 2015).

Provisional lUCN conservation assessment. Data deficient (DD). Begonia galeolepis

is known from only five collections from five locations (Batoe Keye, G. Buria,

Japoetih-Pileana, Manusela National Park, and W. Kasoe), two of which (Batoe Keye

and W. Kasoe) could not be georeferenced with certainty (Fig. 3). This indicates that

the species used to have a wide distribution on Seram. However, two of the collection

sites, Buria and Japoetih, have been almost completely converted for human habitation

and agriculture. The third collection site is in a legally protected area, Manusela

National Park, where no signs of major anthropogenic disturbance were noticed.

Further exploration is required to assess the species’ current range on the island.

Additional specimens examined. INDONESIA: Seram: 18 Sep \9\1 ,L.M.R. Ratten 114 (BO);

W. Kasoe, West Ceram, 28 Aug 1918, L.M.R. Ratten 1607 (BO); Japoetih-Pileana, 27 Oct 1 937,

P.J. Eyma 1811 (AA, BO); Batoe Keye, 29 Dec 1937, P.J. Eyma 2474 (BO); G. Buria, Kec.

Taniwel, 8 Dec 1984, Ramlanto 310 (BO); cultivated in Bali Botanic Garden from vegetative

material collected in Manusela National Park, 27 Apr 2011 (voucher made from cultivated

material), W.H. Ardi 61 (BO).

Identification key to the Begonia species of the Moluccas

la. Plants erect 2

lb. Plants creeping 3

2a. Leaves broadly ovate; female flowers with 2-5 tepals; male flowers with 2 tepals,

anther connectives not projecting at apex B. rieckei complex

2b. Leaves oblong, elliptic or broadly elliptic; female flowers with 6 tepals; male

flowers with 4 tepals, anther comiectives projecting at apex B. aptera

3 a. Leaf apex acuminate 4

3b. Leaf apex rounded 6

4a. Adaxial leaf surface densely hirsute with red hairs B. sageaensis

4b. Adaxial leaf surface glabrous 5

5a. Stems and petioles with sparse to moderately dense indumentum of fleshy,

appressed and branched red scales; female inflorescence peduncles up to c. 5 mm

long; male flower pedicels up to 1 1 cm long B. galeolepis

5b. Stems and petioles with dense indumentum of branched pinkish trichomes with

filifonn apices; female inflorescence peduncles up to 3.5 cm long; male flower

pedicels up to c. 4 cm long B. holosericeoides

Begonia galeolepis, a new species from Moluccas

303

6a. Male flowers with two tepals; female inflorescence peduncles 2-3 mm long; ovary

densely hairy B. holosericea

6b. Male flowers with four tepals; female inflorescence peduncles c. 1 mm long; ovai*y

glabrous or glabrescent B. aketajawensis

ACKNOWLEDGEMENTS. The authors would like to express gratitude to the Seram

Expedition team from Kebun Raya Bali (I Made Ardhaka, Kadek Erosi Undaharta, and 1 Cede

Tirta) for giving pennission to use their materials, to the curators of A, B, BM, BO, CEB, E, K,

L and SING for allowing us access to herbarium material, and to two anonymous reviewers and

David Middleton for their constructive comments. WA’s work was supported by a Singapore

Botanic Gardens’ Research Fellowship, which is grateflilly acknowledged.

References

Ardi, W.H., Kusuma, Y.W.C., Lewis, C.E., Risna, R.R., Wiriadinata, H., Abdo, M.E. & Thomas,

D.C. (2014). Studies on Begonia (Begoniaceae) of the Molucca Islands 1: Two new

species from Halmahera, Indonesia, and an updated description of Begonia holosericea.

Reinwardtia 14(1): 19-26.

Doorenbos, J.M., Sosef, S.M. & De Wilde, J.J.F.E. (1998). The sections of Begonia including

descriptions, keys and species lists. Studies in Begoniacoae VI. Wageningen Agr. Univ.

Pap. 98(2): 1-266.

Hughes, M. (2008). An Annotated Checklist of Southeast Asian Begonia. Edinburgh: Royal

Botanic Garden Edinburgh.

Hughes, M. & Girmansyah, D. (2011). Searching for Sumatran Begonia described by William

Jack: following in the footsteps of a 1 9th century Scottish botanist. Card. Bull. Singapore

63(1 & 2): 83-96.

Hughes, M. & Hollingsworth, P.M, (2008). Population genetic divergence corresponds with

species-level biodiversity patterns in the large genus Begonia. Molec. Ecol. 17: 2643-

2651.

Hughes, M., Moonlight, R, Jara, A. & Pullan, M. (2015). Begonia Resource Centre. Royal

Botanic Garden Edinburgh, http://elmer.rbge.org.uk/begonia/ (accessed on 7 Jul. 2015).

Moonlight, P.W, Richardson, J.E., Tebbitt, M.C., Thomas, D.C., Hollands, R., Peng, C.I. &

Hughes, M. (2015). Continental-scale diversification patterns in a megadiverse genus:

the biogeography of Neotropical Begonia. J. Biogeogr. 42(6): 1137-1 149.

Thomas, D.C., Ardi, W.H. & Hughes, M. (2009). Two new species of Begonia (Begoniaceae)

from Central Sulawesi, Indonesia. Edinburgh J. Bot. 66(1): 103-1 14.

Wiriadinata, H. (2012). A new species of Begonia (Begoniaceae) from Sagea Lagoon, Weda

Bay, Halmahera Island, North Moluccas, Indonesia. Reinwardtia 13(3): 263-270.

Gardens’ Bulletin Singapore 67(2): 305-307. 2015

doi: 10.3850/S2382581215000265

305

A new typification of Hoya zollingeriana

(Apocynaceae, Asclepiadoideae)

M. Rodda

Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, Singapore 259569

rodda.michele@gmail.com

ABSTRACT. The typrfication of Hoya zollingeriana Miq. is discussed and a lectotype is

selected from the original material. An earlier typification was based on a specimen belonging

to a different taxon and is rejected.

Keywords. Eriostemma, Hoya diversifolia, Java, lectotype, neotype

Introduction

In the recently established online journal 'Hoya New' [http://www.rare-hoyas.

com/publication.htm], Hoya zollingeriana Miq. was combined in Eriostemma

zollingerianum (Miq.) Kloppenb, and typified (Kloppenburg, 2014). The combination

is unnecessary as Eriostemma (Schltr.) Kloppenb. & Gilding (Kloppenburg &

Gilding, 2001) is a genus that has been shown to be genetically and morphologically

indistinguishable from Hoya R.Br. in all recently published phylogenies of the

genus (Wanntorp et ah, 2006a, 2006b; Wanntorp, 2007; Wanntorp & Forster, 2007;

Waimtorp & Kunze, 2009; Wanntorp et ah, 2011; Rodda & Ercole, 2014; Rodda et al.,

2014; Wanntorp et al., 2014). Eriostemma is currently recognised at sectional level

{Hoya sect. Eriostemma Schltr.). The species included in Hoya sect. Eriostemma are

characterised by a terrestrial habit, large flowers (>3 cm across, often much larger), a

prominently stalked staminal corona, club-shaped or clavate pollinia without pellucid

margins, and large follicles with a thick spongy pericaip. Regardless of the taxonomic

status of Eriostemma, the identity of Hoya zollingeriana needs to be ascertained as,

since its publication, it has rarely appeared in the taxonomic literature. I have been

able to find only two authors who cite the name; Koorders (1912: 100), who accepted

the species even though he didn’t examine any type material, and Backer & Bakhuizen

van den Brink (1965: 269), who considered it to be a synonym of Hoya diversifolia

Blume, Kloppenburg (2014) stated that 'Since no holotype species [sic] has previously

been designated for this species: 1 hereby designate #12615 (BO) as the holotype'. That

specimen is later indicated as Miquel 12615 and a low-resolution photograph of the

sheet, obtained before it was re-mounted in 1999, is also published. If that specimen

is indeed original material for Hoya zollingeriana then Kloppenburg's paper can be

considered to be an effective lectotypification following ICN Art. 9.9 (McNeill et al..

306

Gard. Bull. Singapore 67(2) 2015

2012) due to need to correct holotype to lectotype. In August 2014, 1 examined all

Hoya specimens at BO and noted that the specimen indicated by Kloppenburg (2014)

as Miquel 12615 is instead Teysmann 12615, an undated sterile specimen collected

in Sulawesi that 1 tentatively identify as Hoya excavata Teijsm. & Binn. Teysmann

12615 is not amongst the materials cited by Miquel (1857: 518) in the publication of

Hoya zoUingeriana and, therefore, it cannot be the lectotype of the name. If there is no

extant original material, and again applying Art. 9.9 of the ICN (McNeill et al., 2012),

then the typification by Kloppenburg (2014) could possibly count as an effective

neotypification. Given that Kloppenburg (2014) does not indicate whether he has made

a thorough search for all original material of Hoya zoUingeriana, T have examined

herbarium specimens of Hoya at BO, BM, CGE, G, E, FT, K and P (herbarium codes

from Thiers [continuously updated]) in an attempt to locate possible original material

and verify whether Kloppenburg’s (2014) effective neotypification should stand or not.

Lectotypification of Hoya zoUingeriana

Hoya zoUingeriana Miq., FI. Ned. Ind. 2: 518 (1857) [20 Aug 1857]. - Eriostemma

zollingerianum (Miq.) Kloppenb., Hoya New 3(1): 6 (2014) [27 Sep 2014] [epublished].

-TYPE: Indonesia, Java, bij Lalaei, Mei, Zollinger, H. s.n. (lectotype P [P05029459],

designated here). = Hoya diversifolia Blume

Notes. Hoya zollingeriana was based on a Zollinger collection indicated only as 'Java,

bij Lalaei, Mei' (Miquel, 1857). Zollinger’s duplicates can be found in many herbaria,

with significant sets at BO, CGE, G, K, LE, R However, Zollinger’s private set was lent

to Miquel, who used it for his Flora van Nederlandsch Indie (Miquel, 1 857), and is now

incorporated into P (Van Steenis-Kruseman, 1950). In P I found a specimen (barcode

P05029459, https://science.mnhn.fr/institution/mnhn/collection/p/item/p05029459)

labelled Hoya zoUingeriana in Miquel ’s handwriting. This is the only original material of

the taxon that has been found. It is a well-preserved fertile specimen that can be identified

as Hoya diversifolia Blume, a species also described from Java and widespread in East

and Southeast Asia. Backer & Bakhuizen van den Brink (1965: 269) already previously

synonymised Hoya zoUingeriana under H. diversifolia, a decision I can confirm.

ACKNOWLEDGEMENTS. This research received support from the National Parks Board

(Singapore) that sponsored herbarium visits in Asia and Europe. I thank the curators of BO,

BM, CGE, G, E, FI, K and P herbaria for allowing access and/or for providing high quality

images of herbarium specimens and, in particular. Dr Sovanmoly Hul (P) who sent a high

quality scan of the lectotype of Hoya zollingeriana. An anonymous reviewer and Dr David

Middleton, Editor of Gardens’ Bulletin Singapore, are thanked for suggesting improvements

to this manuscript.

A new t3T3ification of Hoya zollingeriana

307

References

Backer, C.A. & Bakhuizen van den Brink, R.C. (1965). Flora of Java, vol. 2. Groningen:

N.V.P. Noordhoff.

Kloppenburg, R.D. (2014). Eriostemma zollingerianum Kloppenburg. Hoya New 3(1): 6-7.

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Kloppenburg, R.D. & Gilding, E. (2001). Eriostemma (Schlechter) Kloppenburg & Gilding.

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Koorders, S.H. (1912). Exkursionsflora von Java, vol. 3. Jena: G. Fischer.

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

up to the year 1950. Flora Malesiana, Series f 1: 5-605. Jakarta: P. Noordhoff.

Wanntorp, L. (2007). Pollinaria of Hoya (Marsdenieae, Apocynaceae) - Shedding light on

molecular phylogenetics. Taxon 56(2): 465-478.

Wanntorp, L. & Forster, P.L (2007). Phylogenetic relationships between Hoya and the

monotypic genera. Madangia, Absolmsia, andMicholitzia (Apocynaceae, Marsdenieae):

Insights from flower morphology. Ann. Missouri Bot. Card. 94(1): 36-55.

[http://doi.org/10.3417/0026-6493(2007)94[36:prbhat]2.0.co;2]

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

Dischidia — Toward phylogenetic understanding. Int. J. PI. Sci. 170(3): 331-342.

[http://doi.org/10.1086/596329]

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

phylogeny of Hoya (Marsdenieae, Apocynaceae) inferred from nuclear and

chloroplast DNA sequences. Molec. Phylogenet. Evol. 39(3): 722-733.

[http://doi.Org/10.1016/j.ynipev.2006.01.022]

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

Hoya (Mai'sdenieae, Apocynaceae): Inferences from the chloroplast triiL region and the

rbcL-atpB spacer. Syst. Bot. 31(3): 586-596. [http://doi.org/10.2307/25064187]

Wanntorp, L., Gotthardt, K. & Muellner, A.N. (2011). Revisiting the wax plants (Hoya,

Marsdenieae, Apocynaceae): Phylogenetic tree using the matK gene and psbA-trnH

intergenic spacer. Taxon 60(1): 4-14.

Wanntorp, L., Grudinski, M., Forster, PL, Muellner-Riehl, A.N. & Grimm, G. W. (2014). Wax

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Gardens’ Bulletin Singapore 67(2): 309-350. 2015

doi: 10.3850/S2382581215000277

309

A revision of Cryptocarya (Lauraceae)

from Thailand and Indochina

R.PJ. de Kok

275 Cricklewood Lane, London, NW2 2JJ, U.K.

dekokrogier@ginail .com

ABSTRACT. A revision of the species of Ciyptocar\>a R.Br. (Lauraceae) from Thailand

and Indochina (Laos, Cambodia and Vietnam) is presented, comprising a key to species,

full descriptions, distribution maps, provisional lUCN conservation assessments, ecological

information and ethno -botanical notes where appropriate. In this treatment, 16 species

are recognised, one species name is validated (Cryptocatya globularia Kostenn. ex de

Kok), nineteen names are lectotypified and nine names are placed into synonymy for the

first time. Using standard lUCN criteria, most species are considered to be Least Concem,

one is assessed as Data Deficient (C. globularia), and six are Endangered {Cryptocatya

chanthaburiensis Kostenn., Cryptocatya hainanensis Men\, Cryptocatya laotica (Gagnep.)

Kosterm., Cryptocatya pallens Kostenn., Cryptocatya pustulata Kostenn. and Cryptocatya

sublanugmosa Kosterm.).

Keywords. Cambodia, Cryptocatya, Laos, taxonomy, Thailand, Vietnam

Introduction

Cryptocarya R.Br. occurs tliroughout the tropics (with the exception of central Africa)

and has its centre of diversity in Southeast Asia. The total number of species is estimated

to be between 200 and 250. The classification of the genera within the Lauraceae

has always been problematic. The general framework of characters which have been

employed by most authors was first established by Nees von Esenbeck (1836) in his

classification of the family. However, the outcomes of such a classification can vaiy

greatly, depending upon the weightage given to each particular character. Recently,

with the help of molecular data, some taxonomic relationships are becoming clearer

(Rohwer et al., 2014). Cryptocarya is now placed in the Cryptocary’a group together

with Beilschmiedia Nees and Endiandra R.Br. as one of the early divergent clades

within the family (Rohwer, 2000; Rohwer et al., 2014). The genus has never been

revised in full, but a number of important modem regional heatments exist. The

Australian taxa were revised by Hyland (Hyland, 1989; Le Cussan & Hyland, 2007),

who recognised 47 species, of which 38-39 (80-82%) were considered to be endemic

to Australia. The Chinese taxa have been revised by Li Xiwen et al. (2008), who

recognised 21 species, of which 15 (71%) are endemic to China. In Brazil, 13 species

are recognised, of which 11 (85%) are endemic (De Moraes, 2007), and amongst

the 1 7 taxa currently recognised from Peninsular Malaysia, only three (c. 18 %) are

310

Gard. Bull. Singapore 67(2) 2015

endemic (De Kok, in press). The genus has never previously been revised for Thailand

and is not even mentioned for Thailand by Suvatti (1978). In his Flore Generale de

Llndo-Chine, Lecomte (1914) only recognised five species {Ctyptocarya caesia

Blume (now Cryptocarya ferrea Blume), C. lenticellata Lecomte (now C. concinna

Hance), C. ferrea, C. ohlongifolia Blume and C. ochracea Lecomte (both now C.

ferrea)), although 17 were recorded in the most recent checklist for Vietnam (Le,

2003: 82-85). In this account, 1 6 species are recognised from Thailand and Indochina

(Laos, Cambodia and Vietnam), with 13 recorded from Thailand, 10 from Vietnam, 8

from Laos and 5 from Cambodia. The numbers of endemic taxa per country are very

low, only two from Thailand and one from Vietnam. The number of endemic species

for the region as a whole is six, which is c. 37 % of the total.

Distribution within Thailand and Indochina

The species distribution patterns within Thailand and Indochina are interesting. A

distinct set of species {Cryptocaiya kiirzii Hook.f., C. nitens (Blume) Koord. & Valeton

and C. rugulosa Hook.f.) occurs only in Peninsular Thailand but are then also found

in Peninsular Malaysia. Some of them are common in Peninsular Malaysia but are

represented by only a few specimens in Thailand: for instance Cryptocarya rugulosa

is only known from one specimen, while C. nitens is known from just two. A different

set of species occurs in northern Thailand, northern Vietnam and northern Laos and

then usually also in southern China {Cryptocaiya concinna and C. hainanensis Merr.).

In contrast, a few taxa {Cryptocaiya amygdalina Nees, C. densiflora Blume, C.

impressa Miq. and, to some extent, C. diversifolia Blume and C. ferrea) bridge the gap

between, on the one hand, northern Thailand and northern Indochina, and on the other.

Peninsular Thailand and Malaysia. These species tend to be very widespread and in

one case occurs all the way to Australia {Cryptocaiya densiflora).

A small group of species are endemic to the region {Cryptocaiya chanthaburiensis

Kostenu., C. glohularia Kosterm. ex de Kok, C. laotica (Gagnep.) Kosterm., C.

pallens Kosterm., C.pustulata Kosterm. and C. sublanuginosa Kosterm.). Most occur

in eastern Thailand, Cambodia, southern Laos and southern Vietnam, which together

form an area that is considered to be under-collected (Parnell et al., 2003; Marsh et

al., 2009), while the others occur either in northern Vietnam {Cryptocaiya globularia)

or northern and central Thailand (C, pallens). Most of these taxa are represented in

herbaria by very few specimens, which has made revising their taxonomy and the

production of an identifrcation key very difficult. This applies in particular to those

taxa for which flowers {Ciyptocary^a globularia) or fruits (C. pustulata) are lacking

in the herbaria consulted. This small number of collections also means that based on

the area of occupancy alone, all these endemic taxa have been assessed as Endangered

according to lUCN standards (lUCN, 2012; see http://geocat.kew.org).

A distinct group of species, characterised by markedly long bracteoles, is

co mm on in Peninsular Malaysia (see discussion of morphology in De Kok (in press))

but is almost entirely absent from the present study area. The only exception is

Cryptocarya of Thailand and Indochina

311

Cryptocarym nigulosa, which is mainly a species from the Sunda Plateau that only just

gets across the border into Peninsular Thailand.

In conclusion, the Oyptocary^a flora of the region studied here is dominated

in the south by species which are common in Peninsular Malaysia and other areas

of the Sunda Plateau, while in the north it is dominated by species that also occur in

southern China. Very little overlap occurs between these two groups. A significant

number of endemic species is found only in the region comprising southern Laos,

eastern Thailand, Cambodia and southern Vietnam.

Material and Methods

This study is based mainly on observations of specimens from the following herbaria:

A, BISH, BKF, BM, BO, C, CAS, E, GH, IBK, K, KEP, L, MEL, MO, MPU, NY, P,

PE, RUPP and SfNG.

In the descriptions that follow:

i) All characters and measurements are from mature material, unless

indicated otherwise;

ii) All collections of Cryptocarya from Indochina and Thailand seen by the

author are cited;

iii) Selected specimens of Indo-Chinese and Thai taxa from outside the

present area are cited only when this material contributed to the taxonomic

descriptions, except for material cited in De Kok (in press);

iv) All synonyms of Cryptocarya taxa from Indochina and Thailand are

included. Synonyms from outside the area are included only in those

cases where type material has been seen by the author;

v) In the descriptions the indumentum is characterised as: ‘glabrous’ when

the underlying surface is without any hairs or with just a few occasional

hairs; ‘sparsely hairy’ when it is showing more underlying surface than

hairs; ‘densely hairy’ when showing less underlying surface than hairs;

and ‘velutinous’ when the underlying surface is completely covered with

hairs;

vi) Petioles and fruit stalks are sometimes described as swollen. This means

not that they are necessary thicker, but that over the length of the structure

from the base the width increases and then decreases near the apex, giving

the whole structure a swollen appearance;

vii) Scans of type material, where seen, were accessed at http://plants.jstor.

org on 20 October 2014;

viii) For the provisional lUCN conservation assessments, all Extents of

Occurrence (EOO) and Areas of Occupancy (AOO) were calculated

using http://geocat.kew.org on 21 October 2014.

312

Gard. Bull. Singapore 67(2) 2015

Taxonomic treatment

Cryptocarya R.Br., Prod. FI. Holl. 402 (1810), nom. cons.; Gamble, J. Asiatic Soc.

Bengal. 75: 37-51 (1912); Ridl., FI. Malay Pen. 3: 76-82 (1924); H.Liou, Laurac.

Chine & Indo-Chine 95-102 (1932); Kosterm., Meded. Bot. Herb. Utrecht 42: 557

(1937); Kosterm., Reinwardtia 4: 243-244 (1957); Kostenn., Reinwardtia 8: 21-196

(1970); Kochummen, Tree FI. Malaya 4: 132-138 (1989); B.Hyland, Austral, Syst.

Bot. 2: 162-213 (1989); Rohwer, Fam. Gen. Vase. PI. 2: 366-391 (1993); van der

Werff & H.G.Richt., Ann. Miss. Bot. Gard. 83: 409^18 (1996); van der Werff, Blumea

46: 134 (2001); Le Cussan & B.Hyland, FI. Austral. 2: 140-178 (2007); P.L.R.de

Moraes, ABC Taxa 3: 1-191 (2007). - TYPE SPECIES: Cryptocarya glaucescens

R.Br. (lectotypified by Kostermans, Notul. Syst. (Paris) 8: 112 (1939)).

Caryodaphne Blume ex Nees, Syst. Laurac. 225 (1836). - TYPE SPECIES:

Caryodaphne laevigata (Blume) Nees, Syst. Laurac. 227 (1836) (= Cryptocarya

laevigata Blume) (lectotypified by Kostermans, J. Sci. Res. (Jakarta) 1 : 122 (1952)).

Pseudocryptocarya Teschn., Bot. Jahrb. Syst. 58: 411^13 (1923). - TYPE SPECIES:

Pseudocryptocarya pauciflora (K.Schum. & Lauterb.) Teschn., Bot. Jahrb. Syst. 58:

411-413 (1923) (= Cryptocarya laevigata Blume).

Kerrdora Gagnep., Notul. Syst. (Paris) 14: 31 (1950). - TYPE SPECIES: Kerrdora

laotica Gagnep., Notul. Syst. (Paris) 14: 31 (1950). (= Cryptocarya laotica (Gagnep.)

Kostemi.).

Shrubs to treesy evergreen. Bud scales few. Leaves alternate or rarely (sub)opposite

(not in this region), pinnately veined, rarely triplineiwed. Inflorescence type 3 (using

terminology of Van der Werff, 2001), axillary, involucral bracts absent, bracteoles

triangular, linear to elliptic or orbicular, usually caducous during flowerhig. Flowers

bisexual, small; perianth lobes 6, isomorphic, caducous, tube turbinate or ovoid,

persistent, constricted at top after flowering. Stamens 9, inserted on throat of perianth

tube, in 1st and 2nd rows introrse, glandless; in 3rd row extrorse, with glands at base;

anthers 2-locular; staminodes 3, shortly stalked, glandless; ovary sessile, free during

flowering, enveloped by perianth tube; style linear; stigma small or inconspicuous.

Fruit drupe-like, wholly enveloped by fleshy or hard, dilated perianth tube, but leaving

a ring like aperture at apex. Stalk sometimes swollen when mature.

Distribution. About 200-250 species, pan-tropical except for central Africa. In this

revision 16 species are recorded from Thailand and Indochina.

Use. The wood of many species is used as a light hardwood.

Cryptocarya of Thailand and Indochina

313

Key to the species

(This key works best for young flowering material, fruiting material is often difficult

to name with any degree of certainty)

la.

lb.

2a.

2b.

3a.

3b.

4a.

4b.

5a.

5b.

6a.

6b.

7a.

7b.

8a.

8b.

Leaves triplinerved C. densiflora

Leaves pinnately veined 2

Mature lower leaf surface glabrous, except on veins 3

Mature lower leaf surface sparsely hairy to velutinous 8

Young twigs with dark brown hairs; mature fruit globose, smooth or faintly

ridged, sparsely hairy (occurring in Peninsular Thailand only) C. nitens

Young twigs with whitish to yellowish brown hairs; mature fmit narrowly

ellipsoid, ellipsoid or ovoid, smooth to ribbed, glabrous or sometimes with a

few hairs at apex (occurring throughout the region) 4

Leaves papery 5

Leaves leathery 6

Twigs densely tomentose; upper surface of leaves sparsely hairy, midrib more

densely hairy; bracteoles <0.5 mm long, caducous; mature fruit (dried) obovoid,

12-ribbed C. concinna

Twigs sparsely hairy; upper surface of leaves glabrous with a few hairs on the

midrib; bracteoles > 4 mm long, persistent; mature fruit (dried) ellipsoid, smooth

C. laotica

Tertiary veins scalariform; petiole 6-20 mm long, slender to swollen when

mature; mature fruits smooth, stalk strongly swollen (currently only known from

Thailand) C. amygdalina

Tertiary veins reticulate to scalariform; petiole 5-15 mm long, slender or only

slightly swollen when mature; mature fruits smooth or ridged, stalk slender to

only slightly swollen (known from Cambodia, Laos, Vietnam and southeastern

Thailand) 7

Tertiary veins scalarifonu on lower surface; petiole 5-8 mm long; mature fruits

(dried) globose, smooth with some faint ridges C. hainanensis

Tertiary veins reticulate to scalariform on lower surface; petiole 8-15 mm long;

mature fruit (dried) ellipsoid, smooth C. sublanuginosa

Midrib raised or flattened at base on upper leaf surface 9

Midrib sunken at base on upper leaf surface 10

9a.

Upper leaf surface bullate, apex emarginate to acuminate; mature fruit with

pronounced or faint ridges C. diversifolia

314

Gard. Bull. Singapore 67(2) 2015

9b. Upper leaf surface slightly bullate, apex long acuminate; mature fruit smooth ....

C. impressa

10a. Perianth lobes velutinous to densely hairy (hairiness of perianth tube and

petals lobes similar); twigs and inflorescences with yellowish to brown hairs;

bracteoles elliptical, < 1 mm long, seldom persistent; fruit (dried) ovoid to

ellipsoid, sometimes obovoid, stalks swollen when mature C.ferrea

10b. Perianth lobes sparsely hairy (clear difference in hairiness between the perianth

tube and the petals lobes) or specimens in fruit 11

lla. Bracteoles 1-3.5 mm long 12

llb. Bracteoles < 1 mm long 15

12a.

12b.

Mature fruit globose, 17-18 mm diameter, finely and sparsely hairy at apex, stalk

not or hardly swollen when mature C. globularia

Mature Ifuit ellipsoid, <13 mm diameter, glabrous, stalk not swollen to swollen

when mature 13

13a. Leaves leathery; bracteoles elliptic to lanceolate, (1 .5-)2-3.5 mm long; perianth

tube 1 .2-2 mm long; fruit stalk swollen when mature (occurring in Peninsular

Thailand) C. nigulosa

13b. Leaves thinly leathery; bracteoles linear, 1-2.5 mm long; perianth tube 0.8-1. 5

mm long; stalk sometimes swollen when mature 14

14a. Leaf base asymmetric in more than 50% of mature leaves; fruit 11-15 mm long,

stalk not or slightly swollen when mature (occurring in east Thailand and southern

Indochina) C. chanthahuriensis

14b. Leaf base symmetric in most mature leaves (> 75%); fruit 18-2 1 miu long, stalk

swollen when mature (occurring in Peninsular Thailand) C. kurzii

15a. Leaves with tertiary veins iiTegularly reticulate; bracteoles triangular

C. concinna

1 5b. Leaves with tertiary veins scalariform; bracteoles round or elliptic to linear

16a.

16b.

Petiole channelled, velutinous; perianth lobes velutinous outside, lobes 1-1.6

mm wide C.ferrea

Petiole half-terete, glabrous to velutinous; perianth lobes sparsely hairy outside;

lobes 0.6-1. 5 mm wide 17

17a.

17b.

Twigs and petioles with dark brown hairs; bracteoles round, caducous; perianth

lobes 0. 8-0.9 mm wide C. pallens

Twigs and petioles with yellowish hairs; bracteoles elliptic to linear, often

persistent; perianth lobes 0.6-1 .5 mm wide C. pustulata

Cryptocarya of Thailand and Indochina

315

1 . Ciyptocmya amygdalina Nees in Wallich, PL Asiatic Rar. 2: 69 (1831); Kosterm.,

Bibliogr. Laurac. 384 (1964); H.W.Li et al., FL China 7: 250-251 (2008). -

amygdalina Buch.-Ham. ex Wall., Numer. List 2585 (1830), nom. nud. - TYPE:

[India?] Datgong, 26 March 1809, [Herb. Hamilton, Wallich Cat. 2585] (lectotype

K-W [KOOl 116509], designated here; isolectotype E [E00393147]). (Fig. 1)

Trees 4.5-25 m tall, dbh 10-30 cm. Bark smooth to rough, reddish or pale or greyish

brown; wood brownish to yellow, sometimes smelling of lavender. Twigs smooth to

slightly longitudinally ridged, glabrous but sparsely hairy when young, hairs whitish

to reddish or yellowish. Leaf lamina leathery, lanceolate to elliptic, 4-21 x (1.7-

)2.5-ll cm, apex emarginate to acuminate, base cuneate to almost rounded, slightly

asymmetric; pinnately veined, secondary veins 5-10 pairs, straight but curved near

margin, tertiary veins scalarifonn; upper surface glabrous with few hairs on major

veins, midrib sunken, secondary veins sunken, tertiary veins faint to inconspicuous,

bright to deep green, shiny; lower surface glabrous with few hairs on midrib and major

veins, midrib raised, secondary veins raised, tertiary veins distinct, glaucous, grey-

brown. Petiole 6-20 nun long, slender to swollen, half-terete, glabrous to sparsely

hairy, hairs yellowish to reddish. Inflorescences 5-14 cm long, densely to sparsely

hairy, hairs yellowish to reddish; bracteoles triangular to round, 0.5-1. 7 mm long,

caducous. Flowers scented, hairs yellow; perianth tube 1-1.3 mm long, velutinous to

sparsely hairy; perianth lobes narrowly ovate to elliptic, 1.3-2 x 0.8-1. 2 mm, apex

round to acute, sparsely hairy, pale gi'een to light yellow. Stamens 0.4-0. 5 mm long,

hairy, cream, anthers yellow. Ovary 1-1.1 x c. 0.6 mm, glabrous or sparsely hairy at

apex; style c. 1 mm long, glabrous, stigma inconspicuous. Fruit (dried) ellipsoid or

naiTowly ellipsoid, 19^0 x 9-15 nim, glabrous, smooth, yellow-red to black when

mature. Stalk red, strongly swollen when mature.

Distribution. Southern China, northeastern India and the Andaman Islands, Bhutan,

Myanmar, Thailand, Peninsular Malaysia, Indonesia (Sumatra) (see Fig. 1).

Ecology. Growing in primary and secondary evergreen broad-leaved or mixed

deciduous lowland to hill forests, sometimes along rivers, at 50-1525 m altitude.

Phenology. Flowering from Febmaiy to July (October); fruiting from March to

November (December).

Provisional lUCN conservation assessment. Least Concern.

Selected additional specimens examined. THAILAND: Chiang Mai: Doi Siithep, 26 Apr

1958, Sorensen et al. 5048 (K [2 sheets]). Phrae: between Ban Nam Ki'ai and Pha Tuem, 16

Apr 1970, Smitinand & Cheke 10817 [BKF 46511] (K, P). Surat Than!: Phanom district,

Khlong Phanom National Park, Khao Sok River, 23 Mar 2005, Gardner et al. ST 1718 (KEP).

Notes. In the original description of this species only one collection was cited (Nees von

Esenbeck, 1831: 69) of which there are two specimens available for lectotypification.

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Gard. Bull. Singapore 67(2) 2015

Fig. 1. Distribution of Cryptocarya amygdalina Nees (A).

one at E and one in K-W. The specimen in the Wallich herbarium at K-W is selected

here as the lectotype.

2. Cryptocarya chanthaburiensis Kosterm., Nat. Hist. Bull. Siam Soc. 25: 33

(1975[‘1974’]). - TYPE: Thailand, ‘Chanthaburi’ [now in Trat], Khao Kuap, 27

Cryptocarya of Thailand and Indochina

317

December 1929, Kerr 17846 (holotype K [K000734460]; isotypes BK [BK257960],

BM [BMOO 11 24606], C [C10013568]). (Fig. 2)

Cryptocarya ferrea van grandiflora Lecomte, FI. Indo-Chine 5: 148 (1914); Kostenn.,

Bibliogr. Laurac. 399 (1964); T.C.Le, Checklist PI. Sp. Vietnam 85 (2003). - TYPE:

[Laos] Mekong, Paklai, 1866-1868, ThoreJ s.n. (lectotype P [P02010425], designated

here; isolectotypes K [K000734461], P [P02010426, P02010428]).

Cry^ptocarya ohlongifolia auct. non Blume: Lecomte., FI. Indo-Chine 5: 146 (1914);

H. Liou, Laurac. Chine & Indo-Chine 101 (1932); Kostenn., Bibliogr. Laurac. 423

(1964); T.C.Le, Checklist PI. Sp. Vietnam 85 (2003).

Tree or shrub 2-1 3 m tall, dbh up to 15 cm. Twigs velutinous when young, becoming

almost glabrous later, hairs yellow. Leaf lamina (thinly) leathery, elliptic to lanceolate,

5.5-19 X 2. 6^. 6 cm, apex round to acuminate, base cuneate or rounded, symmetric

to asymmetric; pimiately veined, secondary veins 5-9 pairs, tertiaiy veins scalarifomi;

upper surface glabrous to very sparsely hairy, midrib sunken at base, hairy, secondary

veins sunken, tertiaiy veins inconspicuous, dull or shiny; lower surface sparsely hairy,

midrib raised, secondary veins raised, teitiaiy veins distinct, hairs yellow. Petiole 8-1 5

mm long, slightly swollen or not, half-terete, velutinous, hairs yellowish, appressed.

Inflorescences 6.5-9 cm long, velutinous; bracteoles linear 1-2.5 mm long. Flowers

with yellowish hairs; perianth tube 1-1.5 mm long, velutinous; perianth lobes elliptic,

I . 2-1 .8 X 0.6-1 . 1 mm, sparsely hairy, apex rounded to acute. Stamens 1 .5-2 miu long,

densely hairy at base. Ovary clavate, c. 2 mm long; style c. 0.5 mm long, linear; stigma

inconspicuous. Fruit (dried) ovoid to ellipsoid, 11-15 x 8-10 mm, smooth, glabrous,

black when mature. Stalk not or only slightly swollen when mature.

Distribution. Southeastern Thailand, Laos, Cambodia, southern and central Vietnam,

including the island of Phti Quoc (see Fig. 2).

Ecology. Along margins of evergreen forests or in bamboo thickets, 250-600 m

altitude.

Phenology. Flowering from December to February; fruiting from November to

February.

Provisional lUCN conservation assessment. Endangered (EN B2ab(ii,iii)). This

species is known from a small number of collections from Indochina and Thailand.

An analysis of the Extent of Occurrence (EOO) gives a conservation assessment of

Least Concern, but an analysis of the Area of Occupancy (AOO) gives the assessment

of Endangered. Given the small area of occupancy and the intensive logging and

landscape modification that has occurred in the last 50 years, it must be considered to

be endangered.

318

Gard. Bull. Singapore 67(2) 2015

Fig. 2. Distribution of Cryptocarya chanthaburiensis Kosterm. (▼) and Cryptocaiya concinna

Hance (D).

Additional specimens examined. VIETNAM: An Giang: Arboretum de Trambom, 11 Nov

1922, Poilane 164 (P). Da Nang: Ba Na, 3 Dec 1923, Poilane 9034 (K, P [2 sheets]); Ba Na,

21 Jul 1930, Poilane 17927 (P). Kien Giang: Phu Qudc, Feb 1874, Li Pi [Pierre] 1432 (BM

[2 sheets], BO [10 sheets], K [3 sheets], P [2 sheets]). Quang Tn: Lang Khoai, 10 Jan 1932,

Poilane 19930 (P). CAMBODIA: Pursat: Veal Veng District, some 250 m east of military

headquarters, 16 Feb 2000, Eanghourt 46 (K, RUPP). Unknown province: Expedition du

Mekong, Lacoine, 1866-1868, Thorel s.n. (P [2 sheets]); Expedition du Mekong, 1866-1868,

Thorel s.n. (P [2 sheets]). LAOS: Savannakhet: pres de Bauthat, 12 km de Savannakhet, 26

Jun 1929, Poilane 16328 (P). THAILAND: Chanthaburi: Krat district, Khao Kuap, 23 Dec

1929, Kerr 17712 (BM, K, P). Chon Buri: Si Racha, Nong Nam Kliio, 10 Nov 1926, Collins

1421 (K).

Notes. Kostermans wrote the name ‘Cryptocarya petelotii’ on a herbarium sheet

{Poilane 19930) of this species but the name was never published. This invalid name

was taken up in An Illustrated Flora of Vietnam (Ho, 1999: 380) and in the Checklist

of Plant Species of Vietnam (Le, 2003: 85), again without the name being validly

published.

In the original description of Cryptocarya ferrea var. grandiflora, Lecomte

(1914: 148) only cites one collection for which there are three P and one K specimens

available for lectotypification. The specimen P020 10425 is selected here as the

lectotype.

Cryptocarya of Thailand and Indochina

319

This species is morphologically very similar to Ciyptocaiya ferrea, from which

it differs in having longer (> 1 mm long) and linear bracteoles, whereas the bracteoles

of C. ferrea are shorter (< 1 mm) and elliptical In addition, the fruits stalks are not or

only slightly swollen when mature in Cryptocarya chanthaburiensis, whereas they are

swollen when mature in C.ferrea.

3. Cryptocarya concinna Hance, J. Bot. 20: 79 (1882); H.Liou, Laurac. Chine & Indo-

Chine 101 (1932); RH.Ho, 111. FI. Vietnam 1: 376 (1999); T.C.Le, Checklist PI. Sp.

Vietnam 82 (2003); H.W.Li et al, FI. China 7: 252 (2008). - TYPE: [China] Hong-

Kong, Wong-Nei-Chung, 23 August 1 880, Ford 2 1 748 (holotype BM [BM00095088 1];

isotypes A [A00041350], K [K000763886, K000768389]). (Fig. 2)

Cryptocarya lenticellata Lecomte, Notul. Syst. (Paris) 2: 333 (1913); H.Liou, Laurac.

Chine & Indo-Chine 99-100 (1932); T.C.Le, Checklist PI. Sp. Vietnam 83 (2003);

H.W.Li et al, FI. China 7 : 252 (2008). - TYPE: [Vietnam] Tonkin, Forets du Mont Bavi,

August 1887, Balansa 2412 (lectotype P [P020 10201], designated here; isolectotypes

BO [2 sheets], K [K000734468], P [P02010199, P02010200]).

Cryptocaryfa microcarpa F.N.Wei, Guihaia 15: 210 (1995); H.W.Li et al, FI. China 7:

252 (2008). - TYPE: China, Guangxi, Huanjiang County, Mulun Village, Hong Tong

Valley, 14 August 1994, Mulun exped. M0193 (holotype IBK [IBK00190178]).

Tree 4-25 m tall, dbh 10-40 cm. Bark grey or brownish, wood brown. Twigs grey-

brown, finely striate, densely tomentose, hairs yellowish brown. Leaf lamina papery,

elliptic-oblong or oblong, (3-)5-13 x (1.5-)2-3(-6) cm, apex obtuse, acute or shortly

acuminate, base cuneate, often unequal; pinnately veined, secondary veins 4-7 pairs,

tertiary veins iiTegularly reticulate; upper surface sparsely hairy, more densely so on

midrib, midrib sunken at base, secondary veins sunken, tertiary veins conspicuous,

green-yellow in young leaves, shiny; lower surface glabrous to sparsely hairy, midrib

raised, secondary veins raised, tertiary veins conspicuous. Petiole 4-10 mm long,

slender to slightly swollen, half-terete, sparsely hairy, wrinkled. Inflorescences 2-12

cm long, sparsely hairy to velutinous, indumentum often very patchy; bracteoles

triangular, 0.2-0.5 mm long, caducous. Flowers green to yellow or white, fragrant,

hairs yellowish; perianth tube 1-1.5 mm, velutinous; perianth lobes oblong, 1.2-2 x

1-1.5 mm, apex round to acute, sparsely hairy. Stamens 1-1.5 mm long, hairy at base;

anthers oblong, 0.8-1 mm. Ovary clavate; style c. 0.5 mm long; stigma inconspicuous.

Fruit obovoid, 15-22 x 5-11.5 mm, glabrous, 12-ribbed, black or blue-black when

mature. Fruit stalk not swollen when mature.

Distribution. Southern China, eastern Thailand, Laos, Cambodia, Vietnam (see Fig. 2).

Ecology. Evergreen broad-leaved forests, from 550-1200 m altitude.

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Gard. Bull. Singapore 67(2) 2015

Phenology. Flowering from March to August; fruiting from June to December.

Vernacular names. Cay Mot, Re Dap or Co San Coc (Vietnamese); Co puin link

(Muong); Ko mi ham (Laotian).

Provisional lUCN Conservation Assessment. Least Concern.

Use. The wood is finely grained, heavy and durable, and is used for furniture or house

constmction.

Selected additional specimens examined. CHINA; Hainan: 17 Oct 1933, Liang 63456 (K, P);

Ching Mai District, Mei Maan, 25 Sep 1 932, Lei 1 7 (K, P); Hung Mo Shan, 12 Aug 1929, Tsang

& Fung (688) 18222 (P). Guangdong: Tinghushan, 1978, Chow 78133 (K); Tinghushan, 1978,

Chow 78130 (K). Fujian: Yeuping Cave, 12 Aug 1924, Chung 2994 (K). TAIWAN: Locality

unknown: 26 Nov 1918, Wilson 11115 (K).

VIETNAM: Ha Tay: Massif du Tam Dao, Aug 1931, Petelot 5360 (P); Massif du Tam Dao,

Aug \ 93l, Petelot 5361 (P); Bavi, Vallee de Lankok, Massif du Tam Dao, 2 Dec 1888, Balansa

2397 (P). Lao Cai: Cha-pa [Sa Pa], Massif de Nui Bien, 7 Sep 1926, Poilane 13181 (K, P [2

sheets]). Nghe An: De Vinh, Ke Nhe, Sep 1917, Chevalier 38162 (P [2 sheets]). Nha Trang:

Massif du Hon Ba, 22 Sep 1918, Chevalier 38680 (P [5 sheets]). Hoa Binh: Sendee forestier

du Tonkin s.n. (P). Quang Tri: Massif du Doug, 5 Jul 1924, Poilane 11203 (P [2 sheets]).

Locality unknown: Forest de Vien-yen, Dec 1918, Chevalier 39613 (P [2 sheets]); Hau Phuc,

20 Jan 1923, Butreau 53 (P).

LAOS: Savannakhet: 20 km de la route de Savannakhet a Quang Tri, 28 Jan 1925, Poilane

11665 (P). Locality unknown: Fau Meia, Nea Hani Firo, 15 Sep 1920, Poilane 1865 (P [2

sheets]).

CAMBODIA: Kanipot: Mont de I’Elephant, 1 Aug 1919, Chevalier 349 (P [2 sheets]).

THAILAND: Nakhon Ratchasima: National Park, 20 Dec 1962, Phengkhlai 598 (K).

INDOCHINA: Locality unknown: Schmid 1808 (P).

Notes. In the original description of Cryptocarya lenticellata Lecomte (1913: 333),

only one collection is cited for which there are three P, two BO and one K specimen

available for lectotypification. The specimen P020 10201 is selected here as the

lectotype.

4. Cryptocarya densiflora Blume, Bijdr. 556 (1856[‘1852’]); Gamble, J. Asiatic Soc.

Bengal. 75: 39^0 (1912); Ridl., FI. Malay Pen. 3: 77 (1924); FI.Liou, Laiirac. Chine &

Indo-Chine 98 (1932); Kosterm., Bibliogr. Laurac. 395-396 (1964); Kochummen, Tree

FI. Mai. 4: 134 (1989); P.H.Ho, 111. FI. Vietnam 1: 377 (1999); H.W.Li et al., FI. China

7: 248-249 (2008); Le Cussan & B.Hyland, FI. Austral. 2: 157 (2007). - Caryodaphne

densiflora (Blume) Nees, Syst. Laurac. 228-230 (1836). - TYPE: [Indonesia, Java]

Mt. Salak, Blume s.n. (lectotype L [L0036111], designated here; isolectotypes L

[L0036101, L0036102, L0036103, L0036104, L0036105, L0036107, L0036112), U

[U0002692], NY [NY00355062, NY00355063, NY00355064]). (Fig. 3)

Cryptocarya of Thailand and Indochina

321

Cry^ptocaiya flewyi A.Chev. ex H.Liou, Laurac. Chine & Indo-Chine 98 (1932);

Kosterm., Bull. Bot Surv. India 10: 287 (1968); T.C.Le, Checklist PL Sp. Vietnam

83 (2003). - TYPE: [Vietnam] Sud Annam, Nha Trang Prov., Massif du Hon Ba,

28-31 August 1918, Chevalier 38863 (lectotype P [P02009912], designated here;

isolectotypes P [P02009911], MO [MO 1295 102]), synon. nov.

Cryptocarya annamensis C.K.Allen, J. Arnold Arbor. 23: 459-460 (1942); P.H.Ho,

111. FI. Vietnam 1 : 376 (1999). - TYPE: [Vietnam] French Indochina, Annam, Station

Agricole de Blao Prov., de Haut Donai, Poilane 22294 (holotype P [P00745435];

isotypes A [A00041362, A0041363, A00041364], BM [BM000950884], K

[K000734455], L [L0036060], P [P00745436], US [US00099476]), synon. nov.

Cr)^ptocanm oligoneura Kosterm., Nat. Hist. Bull. Siam Soc. 25: 34 (1975[‘1974’]).

- TYPE: Peninsular Thailand, Nakhon Si Thammarat, 4 May 1955, Thaworn 258

(holotype C [Cl 00 11 742]), synon. nov.

Tree, rarely a shrub, 3-20 m tall, dbh 5-35 cm. Bark (dark) grey to (reddish) brown

or black, smooth or dippled, aromatic when crushed, wood (pale) yellow to white.

Twigs sparsely hairy, longitudinally ridged. Leaf lamina leathery, elliptic to oblong,

7-21 X 3.2-14 cm, apex long-acuminate, base rounded to cuneate; triplinerved;

secondary veins 2-3 pairs, pinnate, miming up to y4 of leaf length towards the apex,

tertiary veins scalarifonn; upper surface glabrous, midrib and secondary veins sunken,

tertiary veins faint or inconspicuous, dark to yellowish green, young leaves white-

green, crushed leaves aromatic; lower surface glabrous to sparsely hairy, midrib and

secondary venation raised, reticulations distinctly visible, usually glaucous. Petiole

slender, 10-16 mm long, channelled, glabrous to sparsely hairy. Inflorescences 2.5-8

cm long, densely hairy, hairs brown to pale yellow; bracteoles ti'iangular, 0.5-1 mm

long, caducous. Flowers white to (creamy-greenish) yellow to brownish; perianth tube

1.2-1. 6 mm long, densely hahy; perianth lobes elliptic, 2-3 x 1-1.4 mm, apex acute,

densely hairy. Stamens 1.4-2. 8 mm long. Ovary clavate, c. 2 mm; style c. 1 mm long,

linear; stigma inconspicuous. Fruit (dried) (depressed) globose to ellipsoid, 12-20 x

16-20 mm, glabrous, smooth or with 12 shallow longitudinal ridges and/or warty, dark

purplish or (bluish) black when mature. Stalk slender to only slightly swollen when

mature.

Distribution. Southern China, Laos, Cambodia, Vietnam, Thailand, Malaysia

(including Sarawak and Sabah), Brunei, Indonesia (Kalimantan, Sumatra, Java,

Sulawesi, Lesser Sunda Islands (Flores and Sumbawa), the Moluccas (Am Islands)

and Papua), Philippines, Papua New Guinea, northeastern Australia (see Fig. 3).

Ecology. Common in primary and secondary forests, sometimes along rivers, in peat

swamps or kerangas forest. Soil: limestone, sandstone, granite or ultra-basic rock

derived soils or sandy loam or clay soils, at 0-1600 m altitude.

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Phenology. Flowering in January to July (October); fruiting in April to November.

Vernacular names. Ca duoi hoa (Vietnamese); Cay com (Muong)\ Kokhe (Laotian).

Provisional lUCN conservation assessment. Least Concern.

Use. The wood is used for house construction and furniture making in China and for

boats in Indochina.

Selected additional specimens examined. VIETNAM: Hoa Binh; Lung Van, Tan Lac, 27 Jan

1931, Poilane 18951 (P [2 sheets]). Lao Cai: Cha-pa, Cho-bo, 14 Aug 1926, Poilane 12934

(P [3 sheets]); 31 Jan 1943, Petelot 8579 (P); Feb \93\, Petelot 5358 (P [4 sheets]); Aug 1935,

Petelot 5445 (P [2 sheets]). Lam Do'ng: Bao Loc, Haut Donai, 22 Jan 1933, Poilane 21763 (P,

SING); Haut Donai, 3 Feb 1933, Poilane 21809 (P [2 sheets]); Haut Donai, Station Agricole

de Blao, 1 1 May 1933, Poilane 22483 (P [2 sheets]); Blao, Sam Dong, 5 Apr 1953, Schmid s.n.

(P). Locality unknown; Tonkin et Annam, 26 Jan 1931, Poilane 18917 (K, P).

LAOS; Champassack: Plateau des Bolovens, pres de la Station Agricole, 20 Nov 1938,

Poilane 28489 (P [2 sheets]); Plateau des Bolovens, entre Muong Bok Kao et Phong Thani,

5 Oct 1928, Poilane 15845 (P [3 sheets]). Hiiaphan: Sam Neua, 9 Oct 1920, Poilane 2026

(K, P [2 sheets]). Saravane: Sommet du Pou Set, 24 Oct 1928, Poilane 16128 (P [2 sheets]).

CAMBODIA; Kampong Cham: 23 Jan 1939, Poilane 28804 (P [4 sheets]).

THAILAND: Chiang Mai: Chieng Saen, 25 Mar 1921, Kerr 5133 (BM, K); Fang, 21 Feb

1969, Van Beusekom & Phengklai 2661 (K, P). Narathiwat: Sungei Kolok, Nikom Waeng, 25

Cryptocarya of Thailand and Indochina

323

Feb 1974, Larsen & Larsen 32603 (K). Kanchanaburi: Kao Ri Yai, 2 Feb 1933, Kerr 10409

(BM, K); Thong Pha Phum, Pilok, 25 Jan 2009, Middleton et al. 4785 (K).

INDOCHINA: Locality unknown: north du Tonkin et du Laos, 10 Dec 1925, Poilane 25054

(K, P).

Notes. The oldest name of this species is Laurus triplinervia Reinw. ex Blume, but

a combination in Cryptocarya would be too similar to the well-known Cryptocarya

triplinervis R.Br. In line with the ICN (McNeill et al., 2012), and in particular article

53.3, example 11, 1 am retaining the second oldest name.

The name Ciyptocary^a dens (flora is based on two gatherings, each with several

duplicates (Mt. Salak, Blume s.n. L0036101, L0036102, L0036103, L0036104,

L0036105, L0036107, L00361 1 1, L00361 12, U0002692, NY00355062, NY00355063,

NY00355064) and on the collection: "Ki-tjetja' Reinward s.n, (BO, L0036108,

L0036109, L0036110). The duplicate in L of Mt. Salak, Blume s.n. (L0036111) is

selected here as the lectotype.

In the original description of Cryptocarym fleuryi A.Chev. ex H.Liou (1932: 98)

only one collection is cited for which there are two P and one MO specimens available

for lectotypification. The specimen P0200991 2 is selected here as lectotype.

This is one of a very few species of Ciyptocary^a which can easily be identified

without using a key. The combination of ti*iplinei*ved leaves, with the major veins not

joining at apex and which dry pale yellowish/reddish brown with a clearly paler under

surface, is a unique combination of characters for this species in Thailand and Indochina.

5. Cryptocarya diversifolia Blume, Mus. Bot. 1 (1851). - TYPE: [Indonesia] Insulae

Sumatra, in Provincia Palembang, 1834, Pretorius s.n. (lectotype K [K000768453],

designated here; isolectotypes NY [NY00581223], U [U0002715, U0002716]). (Fig.

Cryptocarya crassinervia Miq., FI. Ned. Ind. 1: 924 (IS5S); Gamble, J. Asiatic Soc.

Bengal. 75: 41-42 (1912); Ridl., FI. Malay Pen. 3: 78 (1924); Kochummen, Tree

FI. Mai. 4: 133-134 (1989). - Cryptocarya griffithiana var. crassinervia (Miq.) Ng,

Gard. Bull. Singapore 57: 67 (2005). - TYPE: [Indonesia] Sumatra, Fort de Kock,

Teysmann %.n. [1007 HB] (lectotype L [L0036097], designated here; isolectotypes BO,

L [L0036098]), synon. nov.

Cryptocarya infectoria (Blume) Miq., FI. Ned. Ind. 1: 924 (^1858); Kochummen, Tree

FI. Mai. 4: 135 (1989); P.H.Ho, 111. FI. Vietnam 1: 378 (1999); T.C.Le, Checklist PI.

Sp. Vietnam 84 (2003). - Cylicodaphne infectoria Blume, Mus. Bot. 2: 11 (1856

[‘1852’]). -TYPE: [Indonesia] Archipelago Indico, Waitzs.w. (lectotype L[L003 6 159],

designated here; isolectotypes L [L0036157, L0036158]).

Cryptocarya obtusifolia Merr., Philipp. J. Sci. 21: 344 (1922), nom. illeg. (non

Cryptocarya obtusifolia Meisn., Prodr. 15: 508 (1864)); Kosterm., Reinwardtia 7:

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Gard. Bull. Singapore 67(2) 2015

312 (1968); P.H.H6, 111. FI. Vietnam 1: 379 (1999); T.C.Le, Checklist PI. Sp. Vietnam

84 (2003); H.W.Li et al., FI. Cliina 7: 253 (2008). - Cryptocaiya impressinervia

H.W.Li, Acta Phytotax. Sin. 17: 70 (1979); H.W.Li et ah, FI. China 7: 253 (2008). -

TYPE: China, Hainan, Ng Chi Leng, 21 December 1921, McClure 8581 (lectotype

CAS [CAS0033144], designated here; isolectotypes A [A00041359, A00041360],

K [K000768393], MO [MO1889406], NY [NY00581220, NY00581221], P

[P020 10336]), synon. nov.

Cryptocarya lecomtei Kosterm., Nat. Hist. Bull. Siam Soc. 25: 34 (1975 [‘1974’]).

- TYPE: Thailand, Chanthaburi [now in Chon Buri], Si Racha, 17 January 1946,

Nakkarn 206 (holotype C [Cl 00 11 743]), synon. nov.

Trees 3-23 (-3 3) m tall, dbh 5-40 cm. Bark scaly to smooth, brownish to black, inner

bark reddish brown; wood pale yellow, aromatic. Twigs stout, longitudinally ridged,

velutinous, hairs yellowish to reddish brown. Leaf lamina thickly leathery, elliptic

to lanceolate, 6-32 x (4.5-)8-15 cm, apex emarginate to rounded or acuminate, base

cuneate to rounded, clearly to slightly asymmetric; pinnately veined, secondai*y veins

5-12 pairs, tertiary veins scalariform; upper surface bullate, glabrous except for velvet

midrib and secondary veins, midrib raised at base, secondary veins sunken, cuiwing

and joining near margin, tertiary veins clear to inconspicuous, dark green, metallically

shiny; lower surface velutinous, midrib raised, secondary veins raised, tertiary veins

distinct, sparsely to densely hairy, bluish green to silvery grey, glaucous. Petiole 7-27

mm long, swollen, velutinous, half-terete or shallowly channelled, longitudinally

ridged. Inflorescences 4-16 cm long, longitudinally ridged, velutinous, hairs yellowish

to reddish; bracteoles triangular to linear, 1-3 mm long, caducous. Flowers greenish

white to (yellowish) brown; perianth tube velutinous, 2-2.4 mm long; perianth lobes

elliptic to lanceolate, 3. 7^.4 x 1.5-1. 8 mm, apex acute, densely hairy, greenish.

Stamens 2, 6-3.2 mm long, hairy. Fruit (dried) globose to ellipsoid, 20-27 x 13-16

mm, distinct to faintly ridged, sparsely hairy to glabrous, black when mature, smelling

of resin. Stalk slender when mature.

Distribution. India (Andaman Islands), China (Hainan), Vietnam, Thailand, Malaysia

(including Sarawak and Sabah), Brunei, Indonesia (Kalimantan and Sumatra) (see

Fig. 4).

Ecology. Primary and secondary forest, sometimes in swamps, at 0-1000 m altitude.

Phenology. Flowering in (May) July to November; fruiting from January to August.

Provisional lUCN conservation assessment. Least Concern.

Selected additional specimens examined. VIETNAM: Bien Hoa: 18 Oct 1931, Poilane 19972

(K). Ha T%: Mont Bavi, 1 888, Balansa 2396 (P [2 sheets]).

THAILAND: Phetchaburi: Kaeng Krachan National Park, near top of Khao Panoen Thung,

Cryptocarya of Thailand and Indochina

325

Fig. 4. Distribution of Cryptocarya diversifolia Blume (A).

26 Jan 2005, Williams 1101 (K, KEP). Surat Thani: Phanom district, Khlong Phanom National

Park, 8 May 2006, Gardner & Sidisunthorn ST 2698 (KEP).

Notes. Four specimens are available for lectotypification of this name (two at L

(fonnerly U), one at K, and one at NY). The specimen at K has flowers and bracteoles

attached to the inflorescence and is therefore selected here as the lectotype.

In the original description of Ciyptocaiya crassiner^ia Miq. (Miquel, 1858:

924), he only cites one collection for which there are two L and one BO specimens

available for lectotypification. The specimen L0036097 is selected here as lectotype.

In the original description of Cylicodaphne infectoria Blume (Blume, 1856:

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Gard. Bull. Singapore 67(2) 2015

11), he only cites one collection for which there are three L specimens available for

lectotypification. The specimen L0036159 is selected here as lectotype.

As the original type material in Manila was destroyed at the end of the Second

World War, a lectotype needs to be selected for Cry'ptocary^a ohtusifolia Merr. In the

original description of Ciyptocary^a obtusifolia, Merrill (1922: 344) cites one collection

for which there are numerous specimens available for lectotypification. The specimen

in CAS0033144 was verified by himself, so it is selected here as the lectotype.

This species is recognised as a variety of Oyptocaiya griffithiana by Ng

(2005: 67-68). In the present work, Cryptocarya diversifolia and C. griffithiana are

recognised as separate species, as they differ with regard to inflorescence bract length

(< 4 mm long as opposed to > 5 mm long), mature fruit surface texture and the amount

of swelling of the fruit stalk.

6. Cryptocarya ferrea Blume, Bijdr. 557 ([‘1852’] 1856); Gamble, J. Asiatic Soc.

Bengal. 75: 45^6 (1912); Ridl., FI. Malay Pen. 3: 79 (1924); Fl.Liou, Laurac. Chine

& Indo-Chine 101 (1932); Kosterm., Bibliogr. Laurac. 399 (1964); Kochummen,

Tree FI. Mai. 4: 134 (1989); P.H.Ho, 111. FI. Vietnam 1: 377 (1999); Ng, Gard. Bull.

Singapore 57: 64-67 (2005). - TYPE: [Indonesia, Java] Cheribon Province, Mt.

Tierimai, October, Blume s.n. (lectotype L [L0036127], designated here; isolectotypes

BO,L[L0036128]). (Fig. 5)

Cryptocaiya oblongifolia Blume, Bijdr. 557 (1856[‘1852’]); Kosterm. Bibliogr.

Laurac. 423 (1964); P.H.Ho, 111. FI. Vietnam 1: 379 (1999). - Cryptocarya ferrea

var. oblongifolia (Blume) Meisn., Prodr. 15: 70 (1864). - TYPE: [Indonesia, Java]

Buitenzorg Prov., Bantam, Tjanjor, October-December, Blume s.n. (lectotype L

[L0036137], designated here; isolectotypes BO, GH [GH00041376], K [K000768429],

L [L0036131, L0036132, L0036133, E0036134, L0036135, L0036136), NY

[NY00355071, NY00355072, NY00355073], U [U0002707]).

Cryptocarya ochracea Lecomte, Notul. Syst. (Paris) 2: 333-334 (1913); Lecomte,

FI. Indo-Chine 5: 145 (1914); Kosterm., Bibliogr. Laurac. 425 (1964); P.H.Ho, 111. FI.

Vietnam 1 : 379 (1 999); TC.Le, ChecklistPl. Sp. Vietnam 85 (2003). -TYPE: [Vietnam]

Cochinchine, Bien Hoa Prov. Song-Lu, February 1877, Pierre 1620 (lectotype P

[P00745428], designated here; isolectotypes A [A0004 13 65], BM [BMOOl 124604], K

[2 sheets], MPU [MPU018667], P [P00745427, P00745429, P00745430]).

Cryptocarya caesia auct. non Blume: Lecomte, FI. Indo-Chine 5: 146 (1914);

Kosterm., Bibliogr. Laurac. 390 (1964); TC.Le, Checklist PI. Sp. Vietnam 82 (2003).

Cryptocarya albiramea Kostenn., Nat. Hist. Bull. Siam Soc. 25: 33 (1975 [‘1974’]). -

TYPE: Peninsular Thailand, Krabi, Tambon Khao Phanom, 2 April 1930, Kerr 18815

(holotype K; isotypes BM, BO), synon. nov.

Cryptocarya of Thailand and Indochina

327

Shrubs or trees (2-)8-22 m tall, dbh 5^0 cm. Bark grey to reddish brown, smooth

to finely scaled, wood white to yellowish. Twigs smooth to angular, velutinous when

young, almost glabrous when mature; hairs yellowish to brown, appressed. Leaf

lamina leathery, narrowly elliptic to oblong or lanceolate, 6-20 x 2-9 cm, apex blunt

to acuminate, base cuneate to rounded, sometimes slightly oblique; pinnately veined,

secondary veins 4-14 pairs, tertiary veins scalarifomi; upper surface glabrous with

sparsely to densely hairy midrib and major veins, midrib and secondary veins sunken,

tertiary veins sunken to inconspicuous, light to dark green, young leaf red, glossy;

lower surface sparsely hairy with denser patches along and on veins, hairs yellowish

to brown, midrib and secondary veins raised, tertiary veins inconspicuous to distinct,

glaucous, grey-green. Petiole 6-20 mm long, slender to slightly swollen, half-terete

to channelled, velutinous; hairs yellowish, appressed. Inflorescences 1.5-20 cm

long, velutinous when flowering to almost glabrous when in fruit, greenish yellow

to cream; bracteoles elliptical, < 1 imn long, seldom persistent. Flowers pale green

to pale yellow, hairs yellowish; perianth tube 0.9-1 .6 mm long, velutinous; perianth

lobes elliptic, 1.5-2. 5 x 1-1.6 mm, apex round to acute, velutinous to sparsely hairy.

Stamens 1.3- 1.7 mm long, densely hairy. Ovary clavate, c. 0.6 imu; style c. 1 mm

long, linear; stigma inconspicuous. Fruit (dried) ovoid to ellipsoid, sometime obovoid,

ll-28(-35) X 7-17.5 mm, glabrous, smooth to obscurely ridged, apex with a narrow

ring like aperture, black when mature. Stalks swollen when mature.

Distribution. Southern Vietnam, Cambodia, Laos, Thailand, Malaysia (including Sabah

but not yet recorded from Sarawak), Singapore, Philippines, Indonesia (Kalimantan,

Sulawesi, Sumatra and Java) (see Fig. 5).

Ecology. Growing in rainforest, sometimes in peat swamps or bamboo forests, from

30-1200 m altitude.

Phenology. Flowering from December to September; fruiting from (December)

January to April.

Vernacular names. Cou Kirp (Vietnamese); Kril dam or Sma Krdbei (Khmer); May ky

guan (Laotian).

Provisional lUCN conservation assessment. Least Concern.

Selected additional specimens examined. VIETNAM; Bien Hoa; Dinh Quan, 47 km de la route

nr 20, 20 Nov 1932, Poilane 21401 (P). Kien Giang: Phu Quoc, 30 Dec 1919, Poilane 878 (P

[3 sheets]).

LAOS: Savannakhet: Bassin du Se-Moun, 1875-1877, Voyage du Harmand, no. 76 (P [3

sheets]).

CAMBODIA: Kampong Cham: 18 Jul 1930, Bejaud 823 (P [2 sheets]). Kampot:

Sihanoukville, 24 Jan 1966, Vidal 5042 (P); Nord du Kampot, 3 Feb 1928, Poilane 14615

(BM, P [2 sheets], SING). Locality unknown: 1877, Harmand (Pierre) 3179 (P [3 sheets]).

THAILAND: Chumphon: Lang Suan District, Pang Wan, 15 Feb 1927, Kerr 11955 (BM,

328

Gard. Bull. Singapore 67(2) 2015

Fig. 5. Distribution of Cryptocarya ferrea Blume (A).

K). Pattani: Bukit Pattani, 25 Jan 1931, Put 3632 (BM, BO, K, P). Prachuap Khiri Khan:

Khao Tao near Hua Him, 18 Sep 1918, Kerr 16020 (BM, K). Surat Thani: Phanom District,

Khao Sok National Park, 2006, Middleton 3993 (K); Phanom District, Khao Sok National

Park, nature trail north of headquarters, 24 Mar 2005, Gardner et at. ST 1 733 (K, KEP).

INDOCHINA: Locality unknown: Voyage du Harmand s.n. (P).

Notes. There are several Blume gatherings available at L and BO for the lectotypification

of Cryptocarya ferrea Blume. The L0036127 specimen, clearly mentioning the original

collection site and including two inflorescences with mature flowers, is selected as a

lectotype here.

Cryptocarya of Thailand and Indochina

329

In the original description of Oyptocarya oblongifolia Blume (Blume, 1856:

557), only one collection is cited for which there are numerous specimens available for

lectotypification. The specimen L0036137 is selected here as lectotype.

In the original description of Cryptocarya ochracea Lecomte (Lecomte, 1913:

333), only one collection is cited. For the lectotypification of this name there are four

specimens available at P, two at K and one each at A, BM and MPU. The specimen

P00745428 is selected here as the lectotype.

This species is often difficult to distinguish from Ciyptocarya kurzii and C.

rugulosa when specimens are in fruit. Table 1 sets out the differences between their

flowers and vegetative characters.

7. Cryptocarya globularia Kosterai. ex de Kok, sp. nov.

Cryptocarya globularia differs from the closely related Cryptocaryra chanthaburiensis

in having mature fruit which are globose rather than ovoid to ellipsoid and which

are also much larger (17-19 x 16.5-18.7 mm, rather than 11-15 x 8-10 mm in C.

chanthaburiensis) and which are sparsely hairy rather than glabrous. - TYPE:

[Vietnam] Tonkin, May 1887, Environs de Tu-Phap, Balansa 2413 (holotype P

[P020 10448]; isotype K (photograph at BO)). (Fig. 6, 7)

Tree 5-6 m tall, dbh 12-20 cm. Twigs in cross-section angular and velutinous when

young, smooth and glabrescent when old; hairs yellow, erect. Leaf lamina leathery,

elliptic to oblong, 7.6-12 x 4—6.7 cm, apex slightly emarginate to short acuminate,

base cuneate to rounded, symmetric to asymmetric; pinnately veined, secondary veins

7-9 pairs, tertiary veins scalarifomi; upper surface glabrous except for the midrib,

secondary veins sunken, tertiary veins and reticulations sunken; lower surface sparsely

hairy, midrib and secondary veins strongly raised, tertiary veins and reticulations

strongly raised, glaucous. Petiole 8-12 mm long, slender, half-terete, glabrous to

velutinous; hairs yellow, short. Inflorescences 6-10 cm long, glabrous to velutinous;

bracteoles linear, 1-2.5 mm long. Flowers unknown. Fruit (dried) globose, 17-19 x

16.5-1 8.7 mm, sparsely hairy, smooth, shiny, black when mature. Stalk not or hardly

swollen when mature.

Distribution. Northern Vietnam (see Fig. 7). Only known from the type.

Ecology. Lowland.

Phenology. Flowering unknown; fhiiting in May.

Provisional lUCN consenration assessment: Data Deficient (DD). This species is only

known Ifom one collection which was collected in 1887 in a part of Vietnam which

has seen significant reductions in forest cover. More fieldwork is needed to establish

the extent of the remaining population(s) and their current threats. This species is

therefore classified as Data Deficient.

330

Gard. Bull. Singapore 67(2) 2015

Table 1. Morphological differences between Cryptocaiya ferrea Blume, C. kurzii Hook.f

and C. rugiilosa Hook.f Fruiting specimens of these species are often impossible to name

satisfactorily.

C. ferrea

C. kurzii

C. rugulosa

Petiole when

Channelled,

Half-terete, glabrous

Half-terete, velutinous

young

velutinous

to velutinous

Bracteoles

Elliptical, < 1 mm

Linear, 1 . 1 - 1 . 8 mm

Elliptic to lanceolate.

long

( 1 . 5-)2-3 . 5 mm long

Outer surface

of flowers

Velutinous

Sparsely hairy

Notes. The name of this species has not previously been validly published. Kostennans

wrote the name on two duplicates, one at P and one at K. The name has not subsequently

been used by other authors. The fruits are reported to be eaten by birds.

8. Cryptocarya hainanensis Merr., Philipp. J. Sci. 2 1 : 343-344 (1922); H.Liou, Laurac.

Chine & Indo-Chine 100-101 (1932); Kosterm., Bibliogr. Laurac. 405^06 (1964);

P.H.Ho, 111. FI. Vietnam 1: 377 (1999); T.C.Le, Checklist PI. Sp. Vietnam 83 (2003);

H.W.Li et al. FI. China 7: 251 (2008). - TYPE: China, Hainan, Five Finger Mountain,

Ng Chi Leng, 20 December 1921, McClure 8707 (lectotype CAS [CAS0033142],

designated here; isolectotypes A [A00041352], BISH [BISH 1006 120], K, MO

[MO1889407], P [P02010435]). (Fig. 7)

Cryptocarya rolletii H.Wang & H.Zhu, Guihaia 19: 197 (1999); H.W.Li et al., FI.

China 7: 251 (2008). - TYPE: China, Yunnan, Jingong, Mengsoon, 16 April 1997,

Wang et al. MS 125 (holotype HITBC (not seen); isotype IBK [IBK00190189]).

Cryptocarya hainanensis forma grandifolia H.Liou, Laurac. Chine & Indo-Chine 100-

101 (1932); T.C.Le, Checklist PI. Sp. Vietnam 83 (2003). - TYPE: [Vietnam] Annam,

Thanh H6a Prov., Phong Y, 15 July 1920, Poilane 1632 (holotype P [P02010434];

isotypes P [P02010436, P02010435], SING [SING0209180]).

Tree 7-20 m tall, dbh 25-35 cm. Bark smooth, brownish, wood yellow. Twigs smooth

to slightly longitudinally ridged, sparsely hairy to glabrous, hairs whitish. Leaf lamina

leathery, lanceolate to oblong-lanceolate, 8.5-18.5 x 2.5-7 cm, apex acuminate

to long acuminate, base cuneate, slightly asymmetric; pinnately veined, secondary

veins 5-7 pairs, tertiary veins reticulate; upper surface glabrous with few hairs on

main secondai 7 veins, midrib sunlcen at base, secondary veins sunken, tertiary veins

inconspicuous, greenish, shiny; lower surface glabrous with few hairs on midrib and

332

Gard. Bull. Singapore 67(2) 2015

Fig. 7. Distribution of Cryptocarya laotica (Gagnep.) Kosterm. (A), C. hainanensis Merr. (♦),

C. pallens Kosterm. (▼) and C. globiilaria Kosterm. ex de Kok (•).

main secondary veins, midrib raised, secondary veins raised, tertiary veins distinct.

Petiole 5-8 mm long, slender to slightly swollen, half-terete, glabrous to very sparsely

hairy, hairs yellowish to reddish. Inflorescences 3-10 cm long, densely hairy, hairs

yellowish brown; bracteoles lanceolate-linear, c. 1 mm long. Flowers with perianth

tube c. 1 mm long, velutinous; perianth lobes broadly ovate, c. 1 mm long, apex acute,

velutinous. Stamens < 1 mm long, hairy at base. Ovaiy ovoid, c. 0.7 mm, style c. 0.7

mm long, stigma capitate. Fn/iV globose, 14-30 x 13-21 mm, smooth with some faint

ridges, glabrous or sometimes with a few hairs at apex, shiny, black or blackish blue

when mature. Stalk slightly swollen when mature.

Distribution. Southern China, Laos, Vietnam (see Fig. 7).

Ecology. Evergreen broad-leaved forests; 470-1600 m altitude, sometimes on

limestone.

Cryptocarya of Thailand and Indochina

333

Phenology. Flowering from January to April; fruiting from April to November.

Vernacular name. Cay gin (Vietnamese).

Provisional lUCN Consei^ation Assessment. Endangered (EN B2ab(ii,iii)). This

species is known from a small number of collections from China and Indochina. An

analysis of the Extent of Occurrence (EOO) gives a conservation assessment of Least

Concern, but an analysis of the Area of Occupancy (AOO) gives an assessment of

Endangered. Given the small area of occupancy and the fact that intensive logging and

landscape modification have occurred since some of these collections were made, this

species must be considered to be endangered.

Selected additional specimens examined. VIETNAM: Lang Son: Entre Dong Mo et Van-Linh,

16 Dec 1940, Petelot 6692 (P [2 sheets]). Ninh Binh: Ciic Phao'ng National Park, 15 Mar

2000, Loc et al. P 10021 (P); Cue Phirong National Park, 20 Nov 2000, Cuong et al. NMC

1267 (P). Quang Binh: Ban Kil, 80 km de la route col 7, Vallee du Soug Ga, 27 Jan 1932,

Poilane 19953 (P). Da Nang: Lien Chieu pres de Tourane, 17 Aug 1923, Poilane 7583 (K, P

[2 sheets]).

LAOS: Champasak: Phou, 3 Mar 1922, Poilane 20310 (P). Vientiane: Pu Tat, Viengchan, 21

Apr 1932, Kerr 21183 (BM, K [2 sheets], P).

INDOCHINA: Locality unknown: Schmid 1821 (P).

Notes. As the original type material in Manila was destroyed at the end of the second

world war, a lectotype needs to be selected. In the original description of Cryptoccuya

hainanensis Merr., Merrill (1922: 343) cites only one collection for which there

are numerous specimens available for lectotypification. The specimen in CAS

(CAS0033 142) was verified by himself, so it is selected here as the lectotype.

This species is very similar to Cry^ptocarya impressa from which it differs mainly

in having a bigger fruit, a lower leaf surface that is glabrous (apart from a few hairs on

the veins), and a midrib that is sunken at base on the upper surface of the leaves. It is

also similar to Cryptoccuya globularia, from which it differs in having leaves that are

glabrous (apart from a few hairs on the veins) and mainly glabrous fruits (see Table 2).

9. Cryptocarya impressa Miq., FI. Ned. Ind. 1: 923-924 (^1858); Gamble, J. Asiatic

Soc. Bengal. 75: 42-43 (1912); Ridl., FI. Malay Pen. 3: 78 (1924); Kochummen, Tree

FI. Mai. 4: 135 (1989). - TYPE: [Indonesia] Sumatra, Payo Kombo, ’Mohdang apie

ape’ Teysmann 1005 (lectotype U [U0002696], designated here; isolectotype BO).

(Fig. 8)

Cryptocarya impressa var. tonkinensis Lecomte ex H.Liou, Laurac. Chine & Indo-

Chine 99 (1932); P.H.Ho, 111. FI. Vietnam 1: 377 (1999); T.C.Le, Checklist PL Sp.

Vietnam 83-84 (2003). - TYPE: [Vietnam] Tonkin, Prov. De Phu Cho, reserve

forestiere de Chan-Mong, 21-22 April 1914, Fleiuy 30111 (lectotype P [P02010233],

designated here; isoleetotype P [P020 10089]), synon. nov.

334

Gard. Bull. Singapore 67(2) 2015

Trees 10-33 m tall, dbh 25^5 cm. Bark smooth to finely fissured, grey to reddish

brown. Twigs velutinous; hairs erect, rusty brown. Leaf lamina leathery, elliptic to

nan*owly obovate, 6-19 x 3-8 cm, apex long-acuminate, base euneate; piimately

veined, secondary veins 5-9 pairs, tertiaiy veins scalarifonn; hairs erect, dark brown;

upper surface slightly bullate, glabrous to sparsely hairy, veins velutinous, midrib

slightly raised at base, secondary veins sunken, tertiary veins sunken, dark green; lower

surface densely hairy, veins velutinous, midrib and secondary veins raised, tertiary

veins and reticulations prominently raised, bluish grey, glaucous. Petiole slender,

5-15 mm long, velutinous, half-terete to shallowly channelled, smooth. Inflorescences

3-13 cm long, velutinous, hairs light rusty brown; bracteoles linear, 2. 5-3. 8 mm long,

velutinous, caducous. Flowers yellow, hairs reddish; perianth tube 0.7-1. 4 mm long,

velutinous to sparsely hairy; perianth lobes elliptic, 1 .2-2 x 0.9-1 .2 mm, apex acute,

velutinous. Stamens 1.1 -1.5 mm long, densely hairy. Ovary clavate, c. 2 mm; style

c. 1 mm long, linear; stigma inconspicuous. Fruit (dried) globose to ovoid, 15-16 x

10-15 mm, glabrous, smooth, blue when mature. Stalk slender when mature.

Distribution. Vietnam, Laos, Peninsular Malaysia and Sarawak, Singapore, Indonesia

(Sumatra) (see Fig. 8).

Ecology. Forest and swamps at 150-900 m altitude. Sometimes over shale.

Phenology. Flowering from May to September; fruiting from April to July.

Vernacular name. Cay hoang mang (Vietnamese).

Provisional lUCN consei'vation assessment. Least Concern.

Use. The timber is used in road construction.

Selected additional specimens examined. VIETNAM: Tay: Col d’Chilao, Massif du Lam

Dao, 8 Aug 1933, Poilane 22834 (P [2 sheets]). Ninh Binh: Cue Phuong National Park, 27 Oct

2000, Cuong et al. NMC 1202 (K, P). Da Nang: Col du Nuages, pres Tourane, 28 Aug 1923,

Poilane 7784 (P).

LAOS: Luangprabang: Phou Phung, 1 Mar 1932, Poilane 20241 (BM, K, P, SING).

INDOCHINA: Locality unknown: Chevalier s.n. (P).

Notes. Two different Teysmann gatherings are mentioned in the original description

of Cryptocarya impressa (Miquel, 1858). Of those, only one has a collection number

and of this collection there are specimens at U, now L, and BO respectively which are

available for lectotypification. As Miquel was based at U, the specimen U0002696 is

selected here as the lectotype.

After studying the type material of Cryptocarya impressa var. tonkinensis

Lecomte ex H.Liou at P, and given the information from the original description (as

the P material no longer has any fruits), it appears that this is a small-leaved version

Cryptocarya of Thailand and Indochina

335

Fig. 8. Distribution of Cryptocarya impressa Miq. (A).

of C. impressa with a young fruit. In the original description of this name, Liou Ho

(1932: 99) cites only one collection. For the lectotypification of this name there are two

specimens available at P. The specimen P020 10233 is selected here as the lectotype.

This species seems to be absent from Thailand, southern Vietnam and Cambodia,

although it is common in Peninsular Malaysia, Laos and northern Vietnam. This may

be a result of the undercollection of this genus in the region.

10 . Cryptocarya kurzii Hook.f., FI. Brit. India 5: 119 (1886); Gamble, J. Asiatic Soc.

Bengal. 75: 48-49 (1912); Ridl., FI. Malay Pen. 3: 80 (1924); Kochummen, Tree FI.

336

Gard. Bull. Singapore 67(2) 2015

Table 2. Differences between Cryptocan^a chanthaburiensis Kosterm., C. hainanensis Merr.,

C. impressa Miq. and C. globiilaria Kosterm. ex de Kok.

C. chanthaburiensis

C. hainanensis

C. impressa

C. globularia

Under

surface of leaf

Sparsely hairy to

almost glabrous

Glabrous

Densely

hairy

Sparsely hairy

Midrib at

base on the

upper surface

Sunken

Raised or

flattened

Sunken

Indumentum

on fruit

Glabrous

Glabrous or

sometimes with

a few hairs at

apex

Glabrous

Sparsely hairy

Fruit shape

and size

(in mm)

Ovoid to ellipsoid,

11-15 X 8-10

Globose,

14-30 X 13-21

Globose to

ovoid,

15-16 X

10-15

Globose,

17-19 X

16.5-18

Mai. 4: 135 - 136 (1989); Ng, Gard. Bull. Singapore 57: 64-67 (2005). - Oyptocaiya

wightiana var. griffithii Meisn., Prodr. 15: 70 (1864). - TYPE: [Myanmar] Tenasserim,

Mergui, Griffith 1142 [Kew Distribution 4274] (lectotype K [K000768400], designated

here). (Fig. 9)

Shrubs or trees 2.5-30 m tall, dbh 10-25 cm. Bark smooth to slightly scaly, grey-

black to reddish brown, wood yellow to pinkish white. Twigs velutinous when young,

becoming more glabrous when older, hairs yellow to brown. Leaf lamina thinly

leathery, elliptic to lanceolate, 5-18 x 2.2-6. 5 cm, apex round to acuminate, base

cuneate or rounded, symmetrical; pinnately veined, secondary veins 4-9 pairs, tertiary

veins scalarifomi; upper surface glabrous, except hairs on midrib, midrib sunken,

secondary veins sunken, tertiary veins inconspicuous, bright to darkish green, young

leaves salmon pink, dull or shiny; lower surface sparsely hairy to almost glabrous,

hairs yellow, midrib and secondary veins raised, tertiary veins faint below, silvery

to greenish grey or bluish. Petiole slender to slightly swollen, 10-13 mm long, half-

terete, glabrous to velutinous, hairs yellowish, appressed. Inflorescences 3-12 cm

long, velutinous to sparsely haiiy, greenish yellow; bracteoles linear 1 .4-1 .8 mm long,

caducous. Flowers pale yellow to yellowish brown or pale gi'eenish yellow, hairs

yellowish; perianth tube 0.8-1. 2 mm long, velutinous to sparsely hairy; perianth lobes

elliptic, 1.6-1 .8 x 0.8-1. 3 mm, apex round to acute, sparsely hairy. Stamens 1.2-1. 4

mm long, densely hairy. Ovary clavate, c. 0.6 mm; style c. 1 mm long, linear; stigma

inconspicuous. Fruit (dried) ovoid to ellipsoid, 18-21 x 10-15 mm, smooth, glabrous

Cryptocarya of Thailand and Indochina

337

Fig. 9. Distribution of Cryptocarya kurzii Hook.f. (A).

when mature, sparsely hairy when young, apex with a wide ring-like aperture, purple

blue or black when mature. Stalk swollen when mature, up to 3.3 mm in diameter.

Distribution. Southern Myanmar, Thailand, Malaysia (including Sabah and Sarawak),

Singapore, Bmnei, Indonesia (Kalimantan and Sumatra), Philippines (see Fig. 9).

Ecology. Growing in primary to secondary forests, often in wet localities, at 0-610 m

altitude. Sometimes over granite.

Phenology. Flowering from August to May; fruiting from July to May.

338

Gard. Bull. Singapore 67(2) 2015

Vernacular name, Hu ban bai leg (Thai).

Provisional lUCN consei^ation assessment. Least Concern.

Selected additional specimens examined. THAILAND: Phangnga: Nai Chong, 11 May 1973,

Geesink & Santisuk 5356 (K). Nan: Pua, Doi Phu Kha National Park, 14 Nov 2001, Srisanga

2245 (SING). Narathiwat: Kok Dan Peat Swamp Forest, 17 Feb 1984, Niyomdham 786 (K).

Satun: La-ngu District, Tarutao National Park, 1 8 Feb 2005, Gardner et al. ST 1556 (K, KEP).

Trang: Khao Chong, 15 Apr 1969, Phusomsaeng 154 (K, P). Yala: Than To, Ban Chulaphon

Phatthana, 13 Feb 2004, Middleton 3046 (K).

Notes. Hooker (1 886: 119) bases this name on the description of Cryptocaiya wightiana

var. griffithii Meisn. (Meissner, 1864: 70). In this earlier description two specimens are

cited: Tenasserim, Mergui, Griffith 1142 [Ke\v Distribution 4274]) and Canara, Stocks

s.n. (v.s. in Hooker herbarium). Meissner (1864: 70) states that he had not seen the

later specimen, and Hooker himself does not mention it in his description. 1 could not

find the Stocks specimen in K under the obvious names. Given that a lectotype should

be chosen, I select the Griffith 1142 [Kew Distribution 4274]) here as the lectotype as

it is the only one that I know still to be extant.

11 . Cryptocarya laotica (Gagnep.) Kosterm., Bull. Bot. Surv. India 10: 287 (1968);

P.H.Ho, Til. FI. Vietnam 1: 378 (1999); T.C.Le, Checklist PI. Sp. Vietnam 84 (2003).

-Kerrdora laotica Gagnep., Notul. Syst. (Paris) 14: 31 (1950). - TYPE: Laos, Muang-

awn, Chieng-kwang, 6 April 1932, Kerr 20939 (lectotype P [P00745432], designated

here; isolectotypes BM [BM000950885], K [2 sheets]). (Fig. 7)

Tree 1.8-9 m tall, dbh up to 30 cm. Twigs angular, finely striate, sparsely hairy, hairs

yellowish brown. Leaf lamina papery, elliptic to (narrowly) lanceolate, 7-18 x 1.2-6

cm, apex obtuse to acuminate, base cuneate, rarely asymmetric; pinnately veined,

secondary veins 6-8 pairs, tertiary veins reticulate; upper surface glabrous except for

a few hairs on the midrib and main secondary veins, midrib sunken at base, secondary

veins sunken or rarely raised, tertiary veins faint; lower surface glabrous, midrib

raised, secondaiy veins raised, tertiary veins conspicuous. Petiole 3-10 mm, wrinkled,

slender to slightly swollen, half-terete, sparsely hairy, hairs yellowish. Inflorescences

1.5-6 cm long, densely hairy to velutinous; bracteoles triangular, 4-7.5 mm long,

persistent. Flowers with yellowish hairs; perianth tube 0.5-1. 5 mm long, velutinous;

perianth lobes elliptic, 1.8-2 x 1-1.1 mm, apex acute, sparsely hairy. Stamens 1-1.5

mm long, densely hairy. Ovary clavate, c. 1.5 mm long; style c. 1 mm long; stigma

inconspicuous. Fruit (dried) ellipsoid, 13-17 x 7-10 mm, smooth, glabrous. Stalk not

or slightly swollen when mature.

Distribution. Laos, central Thailand, northern and central Vietnam (see Fig. 7).

Cryptocarya of Thailand and Indochina

339

Ecology. Evergreen broad-leaved forests over granite-derived soils, at about 900-2000

m altitude.

Phenology. Flowering from November to April; fruiting from December to May.

Provisional lUCN conservation assessment. Endangered (EN B2ab(ii,iii)). This

species is known from a small number of collections from Indochina. An analysis of the

Extent of Occurrence (EOO) give a conservation assessment of Least Concern, but an

analysis of the Area of Occupancy (AOO) gives an assessment of Endangered. Given

its small area of occupancy and that intensive logging and landscape modification has

occurred since most of the collections were made, it must be considered endangered.

Additional specimens studied. VIETNAM: Bac Giang: Route du Hanoi a Lang Son, Mar 1933,

Petelot 5245 (P [2 sheets]); ibidem, 10 Dec 1946, Poilane 32686 (P). Kon Turn: Massif du

Ngok Guga, pres de Dak To, 27 Nov 1946, Poilane 35663 (P).

THAILAND: Tak: Muang sub district, Taksin Maharat National Park,F*ouwa etal. 3941 (KEP).

Notes. This species is distinctive by having reticulate tertiary venation which is

prominent on the glabrous underside of the leaves. In Indochina and Thailand, it shares

this character only with Ciyptocaiya concinna and some specimens of C. sublanuginosa,

although the character is more common in Peninsular Malaysia (De Kok, in press).

12. Cryptocarya nitens (Blume) Koord. & Valeton, Meded. Lands Plantentuin 68:

220-223 (1904); Kochummen, Tree El. Mai. 4: 136 (1989). - Tetranthera nitens

Blume, Mus. Bot. 1: 375 (1851). - TYPE: [Indonesia] Java, Blume s.n. (lectotype L

[L00362I4], designated by De Kok (2015); isolectotype L [L0036213]). (Fig. 10)

Trees 8-30 m tall, dbh 15-30 cm. Bark smooth or scaly, grey-brown, inner bark

brownish, wood cream coloured with spicy odour. Twigs strongly angular when

young, velutinous, hairs brown. Leaf lamina leathery, elliptic to lanceolate, ovate to

obovate (6-)7.5-23 x (2.2-)3.5-10.5 cm, apex acute to acuminate, base cuneate or

rounded, slightly unequal; pimiately veined, secondary veins 5-10 pairs, tertiary veins

scalariform; upper surface glabrous sometimes with hairs on midrib and secondary

veins, midrib sunken, secondary veins sunken, tertiary veins faint, shiny, (light to

yellow) green, drying greenish brown; lower surfaee glabrous sometimes with hairs

on midrib and secondary veins, midrib and secondary veins raised, tertiary veins faint,

pale green, glaucous. Petiole slender, 10-30 mm long, velutinous. Inflorescences

10-15 mm long, velutinous, greenish white; bracteoles triangular 0.5-0.6 mm long,

caducous. Flowers pale green to yellow or white, hairs yellowish; perianth tube 1.2-

1.6 mm long, velutinous; perianth lobes elliptic to lanceolate, 1.3-1 .8 x 0.8-1 mm,

apex acute, velutinous, greenish white. Stamens 1.2-1. 6 mm long, hairy, dull greenish

yellow, anther bright yellow orange. Ovary clavate, 1-1.2 mm, glabrous; style 1.2-1. 6

mm long, linear; stigma inconspicuous. Fruit (dried) globose, 8-16 mm diameter,

shallowly ridged, sparsely hairy, black when mature. Stalk slender when mature.

340

Gard. Bull. Singapore 67(2) 2015

Fig. 10. Distribution of Cryptocarya nitens (Blume) Koord. & Valeton (•) and C. rugulosa

Hook.f. (A).

Distribution. Peninsular Thailand, Malaysia, Singapore, Indonesia (Sumatra and Java),

Australia (Christmas Island) (see Fig. 10).

Cryptocarya of Thailand and Indochina

341

Ecology. Riversides in open mixed (including bamboo) hill to lowland forest, growing

on limestone, sandstone and granite, at 0-250 m altitude.

Phenology. Flowering from March to October; fruiting from January to November.

Provisional lUCN conset^ation assessment. Least Concern.

Selected additional specimens examined. THAILAND: Nakhon Si Thammarat: Tha Sala,

Khao Luang National Park, Gardner & Tippayasri ST 1660 (KEP). Ranong: Muang Len, 13

Jan 1966, Hansen & Smitinand 11941 (K). Yala: Than To, Ban Chulaphon Phatthana, 9 Feb

2004, Middleton 2875 (K).

Notes, See De Kok (2015) for additional information on this species.

13. Cryptocarya pallens Kosterm., Nat. Hist. Bull. Siam Soc. 25: 34-35 (1975

[‘1974’]). - TYPE: Northern Thailand, Chiang Mai, Doi Angka [Doi hithanon], Mae

Ka Pak drainage, 24 April 1931, Put 3767 (holotype K; isotypes BM [BMOOl 1 24603],

BO, P [P00745426]). (Fig. 7)

Cryptocarya shoreifolia Kosterm., Nat. Hist. Bull. Siam Soc. 25: 36 (1975 [‘1974’]),

as ‘shoreaefolia’. - TYPE: Thailand, Phitsanulok, Tung Salaeng Luang, 15 Febmary

1964, Hansen et al. 11178 (holotype BKF (not seen); isotypes C [C10013574,

C 100 13 575], K, L), synon. nov.

Tree 7-20 m tall. Twigs striate, dark brown, covered with short, erect hairs. Leaf

lamina leathery, lanceolate to elliptic, 6-16(-24) x 2.2-5. 8(-8) cm, apex emarginate

to acuminate, base cuneate, symmetric to slightly asymmetric; pinnately veined,

secondary veins 5-8(-14) pairs, tertiary veins scalarifomi; upper surface glabrous

except for a few erect hairs on midrib, midrib sunken at base, secondary veins sunken,

tertiary veins and reticulations sunken or inconspicuous; lower surface sparsely hairy

with hairs erect, midrib raised, secondary veins raised, tertiary veins and reticulations

prominently raised, glaucous. Petiole 9-15 mm long, slender or only slightly swollen,

smooth, half-terete to channelled, sparsely to densely hairy, hairs msty brown to pale

yellowish. Inflorescences 3-8 cm long, velutinous to sparsely hairy, hairs yellow;

bracteoles round, 0. 1-0.2 mm long, caducous. Flowers with perianth tube 0.9-1. 3

mm long, velutinous; perianth lobes lanceolate, 1.1-1. 7 x 0.8-0.9 mm, apex acute,

sparsely hairy, sometimes velutinous. Stamens 1-1.5 mm long, hairy at base. Ovary

clavate, c. 1 mm long; style c. 1 mm long, linear; stigma inconspicuous. Fruit (dried)

ellipsoid, 13.5-22 x 6.7-13 mm, smooth or with faint ridges, glabrous, deep violet

when mature. Stalk not or slightly swollen when mature.

Distribution. Northern to central Thailand (see Fig. 7).

Ecology. Evergreen forest, between 200-1350 m altitude.

342

Gard. Bull. Singapore 67(2) 2015

Phenology. Flowering from January to April; fruiting from January to June.

Vernacular names. Mak Ki Ai or Salawt (Thai).

Provisional lUCN conservation assessment. Endangered (EN B2ab(ii,iii)). This

species is known from a small number of collections from Thailand. An analysis of

the Extent of Occurrence (EOO) give a conservation assessment of Least Concern,

but an analysis of the Area of Occupancy (AOO) gives an assessment of Endangered.

Given the small area of occupancy and that intensive logging and landscape

modification has occurred since the collections were made, it must be considered to

be endangered.

Additional specimens examined. THAILAND: Clianthaburi: Kao Sabap, 5 Jan 1930, Kerr

17963 (BM, K). Chiang Mai: Me Ka Pak drainage, 9 Jun 1939, Garrett 1146 (K [2 sheets], P);

Pang Tawn, 29 Apr 1931, Put 3814 (BM, K [2 sheets], P). Lampang: Che Saun, 1921, Kerr

4747 (BM, K), 31 Jan 1921, Kerr 4746 (BM, K). Nakhon Ratchasima: Pak Thong Chai, 26

Apr 1968, Phengnaren & Smitinand 647 (K). Petchabun: Kao Keo Kang, Dan Sai, 10 Apr

1922, Kerr 5791 (BM, K [2 sheets], P) and 5791A (BM, K).

14. Ciyptocarya pustulata Kosterm., Nat. Hist. Bull. Siam Soc. 25: 35 (1975 [‘1974’]).

- TYPE: Thailand, Clianthaburi, 27 February 1956, Sangkhachand 615 (lectotype C

[C10013570], designated here; isolectotype C [C10013569]). (Fig. 11)

Tree 1.6-15 m tall, dbh 15-20 cm. Twigs smooth to slightly longitudinally ridged,

velutinous when young, sparsely hairy to glabrous when mature, hairs yellowish.

Leaf lamina leathery to membranous, lanceolate, sometimes obovate, 9-23 x 2-7.5

cm, apex round to acuminate, base rounded to slightly cuneate, slightly asymmetric;

pinnately veined, secondaiy veins 6-10 pairs, tertiary veins scalarifonn; upper

surface glabrous with few hairs on midrib and main secondary veins, midrib and main

secondary veins sunken, tertiary veins visible; lower surface sparsely hairy, midrib

raised, secondary veins curving and raised, tertiary veins visible, glaucous. Petiole

6-13 mm long, slightly swollen, half-terete to channelled, velutinous to glabrous

when more mature, hairs yellowish. Inflorescences 1.8-12 cm long, velutinous to

sparsely hairy; bracteoles elliptic to linear, < 1 mm long, often persistent. Flowers

yellow, scented, hairs yellowish; perianth tube 0.3-2 imn long, velutinous; perianth

lobes elliptic, 0.6-1. 3 x 0.6-1. 5 mm, densely to sparsely hairy, apex acute. Stamens

1-1.3 mm long, hairy. Ovary clavate, glabrous; style c. 1 mm long, linear; stigma

inconspicuous. Fruit (dried young fruits only) ellipsoid when very young, becoming

more globose when older, smooth, hairy, glabrescent when older. Stalk in mature state

unknown.

Distribution. Southeastern and northern Peninsular Thailand (see Fig. 11).

Cryptocarya of Thailand and Indochina

343

Fig. 11. Distribution of Cryptocarya pustulata Kosterm. (•) and C. sublanuginosa Kosterm.

(A).

Ecology. Growing in evergreen lowland forest or in bamboo forest, sometimes along

river banks, at 150-1300 m altitude.

Phenology. Flowering from February to March; fruiting starting in April.

Vernacular name. Mang Kat (Thai).

Provisional lUCN conservation assessment: Endangered (EN B2ab(ii,iii)). This

species is only known from seven collections, made between 1926 and 1956. An

analysis of the Extent of Occurrence (EOO) give a conservation assessment of Near

344

Gard. Bull. Singapore 67(2) 2015

Tlireated, but an analysis of the Area of Occupancy (AOO) gives the assessment of

Endangered. As this part of Thailand has seen significant reduction in forest cover, and

given the small area of occupancy, this species must be considered to be endangered.

Additional specimens examined. THAILAND: Surat Thani: Pak Sai, 3 1 Mar 1927, Kerr 12496

(BM, K, P); Kaw Zao, 15 Apr 1927, Kerr 12749 (BM [2 sheets]), Kerr 12744 (BM [2 sheets]);

ibidem, 18 Apr 1927, Kerr s.n. (BM); ibidem, 3 1 Dec 1926, Kerr 11200 (BM [2 sheets], SING).

Chumphon: Pato District, Lars Quan, 27 Feb 1927, Kerr 12144 (BO, BM, K, P).

Notes. The fruits are only known from very immature collections. They are hairy and

ellipsoid at first but very soon become glabrous and globose. They have no ridges

during this process and the stalk is not swollen when young.

In the original description, Kostennans (1975: 35) did not distinguish which of

the two sheets at C is the holotype of this name. I have therefore selected the specimen

Cl 00 135 70 as the lectotype.

15 . Cryptocarya rugulosa Hook.f., FI. Brit. India 5: 118 (1886); Gamble, J. Asiatic

Soc. Bengal. 75: 43-44 (1912); Ridl., FI. Malay Pen. 3: 78-79 (1924); Kochummen,

Tree FI. Mai., 4: 136 (1989). - TYPE: [Malaysia] Malacca, 26 June 1867, Maingay

1262 (lectotype K [K001084564], designated here; isolectotypes GH [GH00041377],

K [KOO 1084565]). (Fig. 10)

Trees (3-) 12-40 m tall, dbh 25-35 cm. Bark smooth to dippled and/or scaly, reddish-

brown; wood (pale) yellow white to brown, aromatic. Twigs round in cross-section

or slightly angular, velutinous when young, flaking off in patches when old, hairs

yellowish. Leaf lamina leathery, elliptic to lanceolate or ovate, 4-14(-20) x 2-5(-6.5)

cm, apex round to (long) acuminate, base cuneate, even to slightly oblique; pimiately

veined, secondary veins 3-13 pairs, tertiaiy veins scalariform; upper surface glabrous

with sparsely to densely hairy midrib, hairs yellowish, midrib sunken, secondary veins

sunken, tertiary veins sunken or inconspicuous, glossy, dark green, young leaves red;

lower surface sparsely hairy, veins densely hairy, hairs yellowish-reddish, secondary

and tertiary veins raised, grey green, faintly glaucous. Petiole slender to slightly

swollen, 7-15 mm long, half-terete, velutinous, hairs yellowish. Inflorescences 1.5-

10 cm long, velutinous to densely hairy, hairs brown to yellowish; bracteoles elliptic to

lanceolate, ( 1 .5-)2-3.5 mm long, persistent. Flowers yellowish brown to pale yellow;

perianth tube 1.2-2 mm long, velutinous; perianth lobes elliptic, 1.2-2 x 1-1,3 mm,

sparsely hairy, apex acute. Stamens 1.2-1. 5 mm long. Ovary clavate, 1-1.2 mm; style

1-1.2 mm long. Fruit (dried) ellipsoid to obovoid or ovoid, 17-26 x 9.5-15.5 mm,

smooth or with faint ridges, glabrous with sometimes some hairs at apex, apex with

a wide ring-like aperture, purple to black when mature. Stalk swollen when mature.

Distribution. Peninsular Thailand, Malaysia (including Sabah and Sarawak), Indonesia

(Sumatra) (see Fig. 10).

Cryptocarya of Thailand and Indochina

345

Ecology. Growing in primary or secondary forests, often in peat swamps or in sandy

soil, sometimes over shale, ultrabasic or granite, at 1 5-650 m altitude.

Phenology. Flowering from January to March (July); fruiting all year around.

Provisional lUCN cons citation assessment. Least Concern.

Selected additional specimens examined. THAILAND: Narathiwat: freshwater swamp-forest

south of Narathiwat, 9 Mar 1974, Larsen & Larsen 33126 (K).

Notes. The type material 'Malacca, 26 June 1867, Maingay 1262" at Kew contains

two sheets. The specimen on the first sheet has both flowers and fmits, whereas the

specimen on the second sheet does not have flowers and only one immature fruit. The

specimen on the first sheet (KOO 1084564) is selected here as the lectotype.

16. Cryptocarya sublaniiginosa Kostenn., Nat. Hist. Bull. Siam Soc. 25 (1 975[‘ 1974’])

36. - TYPE: Laos, Viengchan [Vientiane], Muang Ban, 28 April 1932, Kerr 21285

(holotype K; isotypes BM [BMOO 11 24602], BO, P [P02010455]). (Fig. 11)

Tree 8-15 m tall, dbh 10-20 cm. Bark brown. Twigs striate, densely hairy when

young, glabrous when mature; hairs erect, yellowish. Leaf lamina leathery, lanceolate

to rarely elliptic or ovate, 5-20 x 1.4-6. 6 cm, apex acute to shortly acuminate, base

cuneate, symmetric to asymmetric; pimiately veined, secondary veins 6-10 pairs,

straight but curved near margin, tertiary veins scalariform to reticulate; upper surface

glabrous with a few hairs on midrib, midrib sunken at base, secondary veins sunken or

inconspicuous, tertiary veins faint to inconspicuous; lower surface glabrous except for

occasional hairs along major veins, midrib and secondary veins with a few hairs, midrib

and secondary veins raised, tertiary veins distinct, glaucous. Petiole 8-15 mm long,

slender to slightly swollen, half-terete, densely hairy to sparsely hairy. Inflorescences

6-14 cm long, velutinous to sparsely hairy; bracteoles linear, 1-2.5 mm long. Flowers

with white to yellowish hairs; perianth tube 0.8-1 mm long, velutinous to sparsely

hairy; perianth lobes elliptic, 1.8-2 x 0.8-1 mm, apex acute, velutinous to sparsely

hairy. Stamens 1.2-2 mm long, densely hairy. Ovaty clavate, c. 1.5 mm; style c. 1 mm

long, linear; stigma inconspicuous. Fruit (dried) ellipsoid, 8.5-18 x 6.4-10.5 mm,

smooth, glabrous, black when mature. Stalk not swollen when mature.

Distribution. Southeastern Thailand, Cambodia, Laos, Vietnam (see Fig. 11).

Ecology. Growing in forest on clay soil at 25-200(-1000) m altitude.

Phenology. Flowering from January to April; fruiting from November to January.

Vernacular names. Cay mong gd (Vietnamese); Mbdy Ky Quan (Laotian).

346

Gard. Bull. Singapore 67(2) 2015

Provisional lUCN Conservation Assessment. Endangered (EN B2ab(ii,iii)). This

species is known from a small number of collections from Indochina. An analysis of

the Extent of Occun'ence (EOO) give a conservation assessment of Least Concern,

but an analysis of the Area of Occupancy (AOO) gives an assessment of Endangered.

Given the small area of occupancy and that intensive logging and landscape

modification has occuiTed since the collections were made, it must be considered to

be endangered.

Additional specimens examined. VIETNAM: Ba Ria-Vung Tau: Montibus ad Baria Galicea,

1 867, Pierre 3598 (BO, BM, K, P). An Giang: Arboretum de Trambom, 9 Dec 1 922, Poilane

1 74 (P); ibidem, 18 Mar \9'3\, Poilane 19139 (P [5 sheets]). Quang Tri: Massif de Doug Che,

21 May 1924, Poilane 10503 (P [2 sheets]).

CAMBODIA: Kampot: Mont de TElephant, 16 Aug 1919, Poilane 377 (K, P). Locality

unknown: Mulu Prey, Jan 1 876, Hannand 268 [Pierre 3 1 78] (BM, K).

LAOS: Attapeu: Bassind’Attapeu, 1875 1877, 1287 (BO, P [2 sheets]). Savannakhet:

20 km de la route de Savannakhet a Quaug Tri, 21 Jan 1925, Poilane 11714 (P); 28 Jan

1925, Poilane 11797 (BO, P); Bassin du Se-Moun, 1875-1877, Harmand 268 (P [3 sheets]).

THAILAND: Chon Burl: Sriracha Forest, 6 Apr 1923, Collins 865 (K); ibidem, 23 Nov 1927,

Collins 1975 (BM, K).

Dubious names

Cryptocarya godefroyana

On the sheet of Godefroy-Lebeuf 287 (Cambodia, ‘Sud Quest du Grand Lac’) housed

at Kew, Kostennans attached an identification slip in 1972 with the name ‘Cryptocarya

godefroyana’. As far as I have been able to determine, this name has never been validly

published. This is not unknown for unpublished names to be found on Kostennans ’s

identification slips. The specimen is sterile and does not resemble any known

Cryptocarya from the region. One other suggestion written on the specimen, in an

older hand than Kostennans ’s, is Roydsia Roxb., now a synonym of Stixis Lour. After

checking specimens in Kew’s herbarium, this identification was also rejected. I do not

believe that this specimen represents a Cryptocarya and it most likely belongs to a

different genus m the Lauraceae.

Excluded names

Cryptocarya chinensis (Hance) HemsL, J. Linn. Soc., Bot. 26: 370 (1891); RH.Ho,

111. FI. Vietnam 1: 376 (1999); T.C.Le, Checklist PI. Sp. Vietnam 82 (2003); H.W.Li

et al. FI. China 7: 249 (2008). - Beilschmiedia chinensis Hance, J. Bot. 20: 79-80

(1882). [China] Hong-Kong, Valle Su-kun-pu, May 1881, Ford (Herb. Propr.) 21705

(holotype BM (not seen); isotype MO [MO 188941 2]).

Cryptocarya of Thailand and Indochina

347

This species is endemic to southern China (Guangdong, Guangxi, Hainan, Sichuan and

Taiwan). It was also reported to occur in Vietnam (Ho, 1999: 376; Le, 2003: 82), but

I could not find any specimens to support this claim. The species is morphologically

very close to Cryptocarya densiflora, from which it differs in having a smaller fruit

that is clearly 12-15 ribbed, a leaf blade that is smaller and narrower, and glabrous

twigs and petioles.

Cryptocarya chingii W.C. Cheng, Contr. Biol. Lab. Sci. Soc. China., Bot. Ser. 10:

111-113 (1936); P.H.HO, 111. FI. Vietnam 1: 376 (1999); T.C.Le, Checklist PI. Sp.

Vietnam 82 (2003). - TYPE: [China] Cherriang, Shunshi, south of Ping Yang, 10 July

1924, Ching 2055 (holotype PE; isotypes BO, E [E00386424], P [P02010170], US

[US00099482]).

This species has been recorded for Vietnam (Ho, 1999: 376; Le, 2003: 82). However, I

have not seen any material from Vietnam to support this, although, given its distribution

in southern China, it may occur in northern Vietnam.

Cryptocarya cuneata Blume, Bijdr. 558 (1826); T.C.Le, Checklist PI. Sp. Vietnam 85

(2003). = Dehaasia cuneata (Blume) Blume, Rumphia 1: 164, t. 46 (1837). - TYPE:

[Indonesia, Java] Nusae Kambaugae, November, Blume s.n .(type L (L0036313)).

Cryptocarya maclurei Merr., Philipp. .1. Sci. 21: 344 (1922); H.Liou, Laurac. Chine

& Indo-Chine 99 (1932); RH.Ho, 111. FI. Vietnam 1: 378 (1999); T.C.Le, Checklist PI.

Sp. Vietnam 84 (2003). - TYPE: China, Hainan, Ng Chi Leng, 22 Dec \92\, AdcClure

8508 (holotype PNH (not seen, presumably destroyed); isotypes A [A00041357,

A00041358], BISH [BISH1006122], K [K000768391], MO [MO1889405], NY

[NY00355052], P [P02010366]).

This species has been recorded from Vietnam (Liou Ho, 1932: 99; Ho, 1999: 378; Le,

2003: 84). However, I have compared the Vietnamese material supposedly belonging

to this species with Chinese specimens of Cryptocarya maclurei, including the type

material, and I have come to the conclusion that the Vietnamese specimens are better

placed in Cryptocarya sublanuginosa.

Cryptocarya metcalfiana C.K.Allen, J. Arnold Arbor. 23: 457-458 (1942); RH.Ho 111.

FI. Vietnam 1: 378 (1999); T.C.Le, Checklist PL Sp. Vietnam 84 (2003). - TYPE:

[China] Hainan, Chim Shan, Fan Maan, Ts’uen & vicinity, Ling Shui (ling-tui) District,

3-20 May 1932, Fung 20087 (holotype NY [NY00581230]; isotypes A [A00041356],

E [E00386408], BO, K [K000768392], MO [MO1889403], P [P02010176], US

[US00956269, US00099512]).

348

Gard. Bull. Singapore 67(2) 2015

In the past, this species was considered to be endemic to Hainan Island in southem

China. More recently it has also been reported from Vietnam (Le, 2003: 84). However,

I could not find any specimens supporting this claim. Ciyptocarya metcalfiana

is morphological very close to C. concinna, but differs from it in having a longer

inflorescence (> 1 0 cm long versus < 8 cm long) and glabrous leaves (versus leaves

sparsely hairy in C. concinna).

Cryptocarya merrilliana C.K. Allen, J. Arnold Arbor. 23: 456-457 (1942); T.C.Le,

Checklist PL Sp. Vietnam 82 (2003); H.W.Li et ah, FI. China 7: 253 (2008). - TYPE:

[China] Hainan, Kumyum, Lau 27635 (holotype A (not seen); isotypes IBK, MO

[M0255232]).

This species has been recorded for Vietnam (Le, 2003: 82). However, I have not seen

any material from Vietnam to support this, although, given its distribution in southem

China, it may occur in northern Vietnam.

Cryptocarya obovata R.Br., Prod. FI. Holl. 402 (1810); P.H.H6, 111. FI. Vietnam 1: 379

(1999); T.C.Le, Checklist PI. Sp. Vietnam 85 (2003). - TYPE: [Australia] New South

Wales, Port Jackson-Hunter River, Brown 3016 (holotype BM [BM000838232]).

This species is endemic to Queensland and New South Wales in Australia (Le Cussan

& Hyland, 2007: 1 70). It was reported to occur in Vietnam by Le (2003 : 84) but I could

not find any specimens supporting this claim.

ACKNOWLEDGEMENTS. The author is grateful to the curators of A, BISH, BKF, BM, BO,

C, CAS, E, GH, IBK, K, KEP, L, MEL, MO, MPU, NY, P, PE, RUPP and SING herbaria for

access to and/or loan of specimens used in the present study. The drawing of Ciyptocarya

globularia was made by Juliet Beentje. I am very grateful to Helen Hopkins for checking

my English and for the comments of two anonymous reviewers. This research received

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

European Community Research Infrastructure Action under the FP7 “Capacities Program.”

This work was also supported in 2015 by a Visiting Research Fellowship at FRIM and a

Research Fellowship at the Singapore Botanic Gardens, which are gratefully acknowledged.

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Nees von Esenbeck, C.G.D. (1831). Lauraceae. In: Wallich, N. Plantae Asiaticae Rariores, vol

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Nees von Esenbeck, C.G.D. (1836). Systema Laurinarum. Berolini: Sumptibus Veitii et

Sociomm.

Ng, F.S.R (2005). Taxonomic notes on Bornean Cryptocarya R.Br. (Lauraceae). Card. Bull.

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Parnell, J.A.N., Simpson, D.A., Moat, J., Kirkup, D.W, Chantaranothai, R, Boyce, P.C.,

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Muasya, A.M., Prajaksood, A., Pendry, C.A., Pooma, R., Suddee, S. & Wilkin, P.

(2003). Plant collecting spread and densities: their potential impact on biogeographical

studies in Thailand. J. Biogeograp. 30: 193-209.

Rohwer, J.G. (2000). Toward a phylogenetic classification of the Lauraceae: Evidence from

inatK sequences. Syst. Bot. 25: 60-71.

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Rohwer, J.G., De Moraes, P.L.R., Rudolph, B. & Van der Werff, H. (2014). A phylogenetic

analysis of the Cryptocarya group (Lauraceae), and relationships of Dahlgrenodendron,

Sinopora, Triadodaphne and Yasunia. Phytotaxa 158: 111-132.

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Gardens’ Bulletin Singapore 67(2): 351-355. 2015

doi: 10.3850/S2382581215000289

351

Newmania sessilanthera (Zingiberaceae):

a new species from Vietnam

H.T. Lim^ J. Leong-Skomickova^, L.X.B. Nguyen^, C.T. D 6 ^ & T.T. Hoang^

^Southern Institute of Ecology, Vietnam Academy of Science and Technology,

1 Mac DTnh Chi Street, District 1 , Ho Chi Minh City, Vietnam

lhtruong@sie.vast.vn

“Herbarium, Smgapore Botanic Gardens, National Parks Board,

1 Cluny Road, Singapore 259569

jana_skomickova@nparks.gov. sg

^Agency of Environment Protection, Department of Natural Resource and Environment,

Phu Yen Province, Vietnam

"^Forest Science Institute of Central Highlands and South of Central Vietnam,

9-11 Hung Vu'cmg Street, Da Eat City,

Lam Dong Province, Vietnam

ABSTRACT. Newmania sessilanthera Luu & Skomick., a new species from Phu Yen province

in central Vietnam, is described and illustrated here. Notes on this species, the third described

to date, and a new key to the species of Newmania N.S.Ly & Skornick. are provided.

Keywords. Conservation, Phu Yen province, species key

Introduction

Newmania N.S.Ly & Skornick., containing the two species N. serpens N.S.Ly &

V V

Skornick. and N. orthostachys N.S.Ly & Skornick. from Quang Ngai province in

central Vietnam, was only recently described (Leong-Skomickova et al., 2011). Here

we describe and illustrate a third species from Phu Yen province, central Vietnam.

Newmania sessilanthera Luu & Skornick., sp. nov.

Similar to Newmania orthostachys N.S.Ly & Skornick. in general appearance and the

upright inflorescence with spikes composed of condensed bracts, but differing in the

thin and plicate leaf blades, the deeply bilobed purple labellum with a bright red base

and white margins basally (incision V 2 -V 5 of the total length of the labellum), and the

stamen with no filament (vs. smooth non-plicate leaf blades, purple labellum with

prominent white lines and incision '/4-V3 of the total length, and the stamen with a well-

developed filament). - TYPE: Vietnam, Phu Yen province, Song Hinh district, Buon

Kit village. Song Hinh Forest Enterprise, 269 m asl, 12°49'51.87"N 108°53'56.88"E,

25 June 2014, Lieu Hong Tru&ng, Tran Gidi, Do Cao TriPY29 (holotype SGN; isotypes

SGN, SING, VNMN). (Fig. 1)

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Gard. Bull. Singapore 67(2) 2015

Terrestial herb to 80 cm tall, with 2-3 leafy shoots per rhizome, 3-6 cm apart. Rhizome

subterranean, branched, 8-12 mm in diam., light brown externally, cream internally,

aromatic. Leafy shoot slender, leafless for about 30-55 cm above the base, with 6-13

distichously arranged leaves; leafless sheaths 3-5, tubular in basal V4-V5 (lowermost

1-2 sheaths short and almost open), greenish or with reddish brown tinge, glabrous,

becoming brown and papery with age; ligule 7-10 mm long, obscurely bilobed

(sometimes nearly entire at the top of leafy shoot), green, turning brown and papery

with age, glabrous; petiole thick (pulvinous), c. 2-6 mm long, ?>-A mm broad, light

green, glabrous; lamina elliptic to weakly obovate, 10-20 x 4-6.5 cm, uppeimost two

to three leaves smaller, thin, visibly plicate, green and glossy above, lighter beneath,

glabrous on both sides, base cuneate to attenuate, apex acute, often ending in a small

sharp mucro (c, 1 mm), margin entire. Inflorescence arising from the rhizome at the

base of the leafy shoot, about 2 inflorescences per rhizome, spikes condensed, erect,

narrowly ovate, 7-10 cm long, with 7-10 flowers, opening in succession from base to

top; peduncle 2-3 cm long, 4-5 nun in diam., sheathing bracts 3-20 mm long, 3-12

mm wide, cream white with dark red-purple reticulate pattern, glabrous, apex obtuse,

sometimes ending in a small mucro; fertile bracts subtending a single flower, spirally

arranged, overlapping each other by 2 / 3 - V4 of thek length, ovate, 25-47 x 20-27 mm,

open to base, cream white to pale ochraceous with dark red-purple reticulate pattern,

glabrous, apex obtuse, sometimes ending in a small mucro; bracteoles one per flower,

18-24 X 10-13 mm, open to the base, semi-translucent, tinged with dark red-purple

reticulate pattern, glabrous. Flower 8-9 cm long; calyx tubular, c. 12 mm long, 1.8 mm

in diam., unilaterally slit 9-10 nun, semi-translucent white with slight red-purple tinge,

glabrous, apex obtuse;J?ora/ tube cylindrical, 6. 2-7. 3 cm long and c. 1 .5 in diam., pure

white, glabrous; dorsal corolla lobe elliptic, c. 1 7 x 7 nun, concave, pure white with

translucent lines, glabrous, sharply mucronate at apex (c. 1 .5 mm); lateral corolla lobes

narrowly elliptic, 15-16 x 5 rnm, pure white with translucent lines, glabrous; lateral

staminodes petaloid, obovate, 16-17 x 7-7.5 mm, pure white, glabrous; labellum

broadly ovate, 21-23 x 13-14 mm at broadest point, deeply bilobed, glabrous, slit

between lobes about V2-V5 of the length of the labellum, marginally reflexed, basally

white, with bright red patch in the middle, centrally purple, lighter puiple towards

margins and apex. Stamen 6 mm long, entirely g\eLbvous; filament absent; anther c. 6

X 1.8 mm, connective tissue pure white, glabrous; anther crest absent; thecae 6 mm

long, straw coloured to yellow, dehiscing longitudinally along entire length, glabrous;

pollen cream to pale yellow. Style white, glabrous; stigma naiTowly cup-shaped c.

1 mm long, c. 0.4 mm in diam., semi-translucent white; ostiole facing more or less

upwards, ciliate. Epigynous glands absent. Ovary c. 2.5 x 2 mm, white, glabrous,

trilocular, placentation axile, with ovules along entire axis. Fruit an ovoid to ellipsoid

capsule, c. 19 x 13 mm (not yet fully ripe) with persistent calyx, semi-translucent

cream coloured, glabrous, 7-10 seeds. Seeds obovoid, bluntly angular, 8-11 x 3 ^

mm, pink-red (not fully ripe), shortly haii*y, shiny; aril whitish translucent, irregularly

laciniate, c. 8 mm long, lobes 2.5-5 mm long. — All measurements based on living

flowering material from which the type and additional two collections were made.

Newmania sessilanthera, a new species from Vietnam

353

Fig. 1. Newmania sessilanthera Lim & Skomick. A. Habit. B. Flowers. C. Leaves (abaxially).

D. Flower enclosed in bract. E. Base of leafy shoot with inflorescences. F. Flower dissection,

from left: calyx (slit open); bracteole; bract; corolla lobes, labellum and staminodes; ovary,

floral tube and stamen (on top). G. Detail of anther from front and back. H. Fruit and seeds.

From type Luu Hong Truong, Tran Gidi, Do Cao Tri PY29 (Photos: Luu Hong Truong)

354

Gard. Bull. Singapore 67(2) 2015

Additional specimens examined. VIETNAM: Phii Yen: Tay Hoa district, Hoa Thinh ward,

My Lam village, Suoi Liep Forest Enterprise, 151 m asl, 12°52'24"N 109°14'09"E, 6 Aug

2015, Nguyen LeXiidn Bach, Hoang Thanh Tnrdng PY507 (SGN, SING); ibidem, 12°51'09"N

109°13'03"E, 340 m asl, 9 Aug 2015, Nguyen Le Xudn Bdch, Hoang Thanh Truong PY636

(SGN, SING).

Distribution. So far the species has been collected from three localities in Song Hinh

and Tay Hoa districts, all in Phii Yen province in central Vietnam.

Habitat and phenology’. Understorey of lowland evergreen tropical forests, growing

on moist soils on granite and along streams. Flowering occurs from June to August,

followed by fruiting which likely extends to September or early October.

Etymology. The specific epithet reflects the lack of the filament in this species.

Provisional lUCN conservation assessment. Endangered (EN Blab(iii)). So far three

localities, each with fewer than 100 scattered individuals, have been obseiwed. The

known EOO is currently slightly less than 100 km^ but there is enough suitable habitat

in the vicinity that it is likely the real EOO is more than 1 00 km- but certainly less than

5000 km^. The type locality is adjacent to agricultural land, recently established from

cleared forest. Further deforestation may occur posing a threat to the population. The

other two populations are within large forested areas with no obvious tlireats.

Notes. Newmania, now with three species, remains endemic to central Vietnam,

although preliminary studies from various researchers suggest further species

extending tliroughout the Annamite mountain range to southern Vietnam. Newmania

species seem to be steno-endemic and, therefore, susceptible to any rapid habitat

changes. This has implications for any conservation effort.

The lack of a filament in Newmania sessilanthera differentiates it readily

from the other two currently known species. Additional differences from Newmania

orthostachys are outlined above in the diagnosis. It is somewhat similar to Newmania

serpens by the presence of a striking red patch on the purple and white labellum,

but differs by its erect inflorescences composed of compact spikes (vs. inflorescence

prostrate on the ground with more or less lax spikes).

Key to Newmania

1 a. Inflorescence prostrate on the ground; with more or less lax spikes N. serpens

lb. Inflorescence more or less erect; spikes condensed 2

2a. Lamina thick and smooth; labellum purple with prominent white lines, bilobed

with incision %-V3 of the total labellum length; filament well-developed, 7-1 1 mm

long N. orthostachys

Newmania sessilanthera, a new species from Vietnam

355

2b. Lamina thin, visibly plicate; labellum purple with bright red base and white

margins basally, bilobed with incision Vz-Vs of the total labellum length; filament

absent N. sessilanthera

ACKNOWLEDGEMENTS. The new species was discovered during the implementation of

the project “Databasing the biodiversity in Phu Yen province (Phase 1 - Forest biodiversity

database)” fiinded by Phu Yen Provincial Department of Natural Resource and Environment/

Agency of Environment Protection in 2014-2015. The authors are grateful to the managers and

staff of local forest enterprises and forest protection departments in the province, especially

Mr Nguyen Van Toan, Mr Tran Duy Tan, Mr Pham Ngoc Minh, Mr Phan Duy Linh, Mi' Tran

Ngoc Tien and Mr Nguyen Huu Khanh for their support during the fieldwork. We especially

thank Mr Nguyen Nliir Thirc, Ms Le Dao An Xuan, Mr Tran Trung True, Mr Tran Gioi and Dr

Nguyen Thanh Men for their kind cooperation. The research of the second author is funded

by National Parks Board, Singapore, and the Czech Science Foundation, GACR (grant no.

14-13541S).

References

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

and Cambridge, UK: lUCN.

Leong-Skomickova, J., Ly, N.S., Poulsen, A.D., Tosh, J. & Forrest, A. (2011). Newmania: a

new ginger genus from central Vietnam. Taxon 60(5): 1386-1396.

Gardens’ Bulletin Singapore 67(2): 357-360. 2015

doi: 10.3850/S2382581215000290

357

A few last words on Far Eastern Dipterocarps

RS. Ashton

The Arnold Arboretum of Harvai'd University, 125 Arborway,

Boston, MA 02130-3500, U.S.A.

and

Royal Botanic Gardens, Kew

Richmond, Surrey TW9 3AE, U.K.

pashton@oeb.harvard.edu

ABSTRACT. Three additions to the Dipterocarpaceae are presented. Recent collections from

southern Vietnam have finally revealed the identity of Dipterocarpiis condorensis Pien'e,

necessitating reduction of D. caudatus Foxw. and nomenclatural adjustment of its subspecies;

an unexpected novelty, Dipterocarpus pseudocorniitus P.S. Ashton, is described from Tawi-

Tawi, SW Philippines; while a new lower montane point endemic, Vatica palongensis P.S.

Ashton, is reported from West Kalimantan.

Keywords. Dipterocarpus, Kalimantan, Philippines, Vatica, Vietnam

Introduction

Although my 1 982 Flora Malesiana account has held remarkably well, there has been a

continuing trickle of novelties, as witnessed by the two new Shorea Roxb. ex Gaertn.f.

species and one new Hopea Roxb. species in the Tree Flora of Sabah and Sarawak,

that richest of dipterocarp provinces (Ashton, 2004); and new discoveries in the genus

most easy to miss, Vatica L., from Peninsular Malaysia (Saw, 2002; Symington, 2004;

Chua et ah, submitted). The following further novelties have arisen while curating the

Dipterocarpoideae held at the herbarium of the Royal Botanic Gardens, Kew. Notable

has been the discovery of an isotype of Dipterocarpus condorensis Pierre, the Paris

duplicate of which I have not had the opportunity to examine as it had been misplaced

in the cabinets.

Dipterocarpus condorensis resolved

Dipterocarpus condorensis Pierre, FI. For. Cochinch. 3, 14: P1.214 (1889); Smitinand,

Vidal & Ho, FI. Cambodge, Laos & Vietnam 25: 10 (1990). - TYPE: Vietnam, Pulau

Condor, Pierre s.n, (lectotype K [K000671095], designated here).

Dipterocarpus condorensis subsp. condorensis

Dipterocarpus caudatus Foxw., Philipp. J. Sci., C 13: 177 (1918); Ashton, Gard. Bull.

Singapore 31:8 (1978); Ashton, FI. Males., Ser. 1, Spermat. 9: 305 (1982); Ashton,

358

Gard. Bull. Singapore 67(2) 2015

Tree FI. Sabah & Sarawak 5: 94 (2004). - TYPE: Philippines, Luzon, Camarines Prov.,

Barrio Hibatac, Alvarez FB 21193 (lectotype K, designated by Ashton (1978)).

Dipterocarpus condorensis subsp. penangianus (Foxw.) P.S.Ashton & Luu, comb,

nov. - Dipterocarpus penangianus Foxw., Malayan Forest Rec. 10: 72 (1932);

Symington, Malayan Forest Rec. 16: 185 (1943); Ashton, Gard. Bull. Singapore 31:

8 (1963); Ashton, Man. Dipt. Brunei 43 (1964). - Dipterocarpus caudatus subsp.

penangianus (Foxw.) P.S.Ashton, Gard. Bull. Singapore 31:8 (1978); Ashton, FI.

Males., Ser. 1, Spennat. 9: 305 (1982); Ashton, PROSEA 5, 1: 174 (1993); Coode et

ah, Checkl. FI. PI. Gymnosperms Brunei Darussalam 68 (1996); Newman et al., Man.

Males, Dipts., Philippines 66 (1996); Newman, Borneo Medium-Flea vy Hardwoods

80 (1998); Ashton, Tree FI. Sabah & Sarawak 5: 97 (2004). - TYPE: Penmsular

Malaysia, Penang, Mt. Olivia, HaniffSFN 3484 (holotype SENG; isotype K).

Dipterocarpus caudatus auct. non Foxw.: Symington, Foresters’ Man. Dipt. 364

(2004).

Additional specimens examined. VIETNAM: Ba R|a-Vung Tau: Binh Chau-Phuac Bmi

Nature Reserve, branchlets with flowers and with young fruit, fallen ripe fruit, Lim Hong

Truong 23, 24, 25 (K).

Notes. The identity of Dipterocarpus condorensis Pierre has remained a mystery for

more than a century. It is based solely on fallen fruit, from the isolated Vietnamese

island of Pulau Condor (Con Dao) in the South China Sea, which do not closely match

any Indo-Chinese species. 1 failed to find the presumed type, in the Paris herbarium,

when preparing my account for Flora Malesiana (Ashton, 1982). Smitinand et al.

(1990), in the Flore du Cambodge, du Laos et du Vietnam, provided no description,

merely quoting that of Pierre along with the number 55 1 1 provided in Pien*e’s original

description, implying that they also failed to find the type in Paris where two of the

authors prepared the account. The Paris collection, furthennore, appears not to have

been photographed. The entry under the name Dipterocarpus condorensis on the Paris

herbarium website, barcode P05454280, consists only of notes on scraps of paper

attached to a herbarium sheet, possibly in Pierre’s hand, including a drawing of a

dissected mature embryo which appears to be the draft of that in Pierre’s plate. It

is dated 1876, but unnumbered. There is a collection at Kew which, although not

numbered by Pierre, bears the name and a description in Pierre’s hand matching part

of his protologue description. I fonnerly assumed it to represent abnonual Iruit of the

most similar Indo-Chinese species, Dipterocarpus turbinatus Gaertn.f., and named

it accordingly. The fruit, however, differs from those of that species notably in its

subspherical rather than ellipsoid-fusifonn calyx tube, and shorter, to 6 cm versus to

13 cm long cal>Tc lobes. The conundmm has now been resolved by Liai Hong Truong,

who has collected material in flower, young and ripe fruit from Vung Tau on the

southern Vietnamese coast, of a species not previously known from Indo-China. The

fmit are an exact and diagnostic match for Dipterocarpus condorensis but the material

Far Eastern Dipterocarps

359

as a whole matches the type subspecies of Dipterocarpus caudatus Foxw. from the

Philippines. A further taxon is thus added to those which are confined to the Philippines

and Indochina, including southemmost coastal China. Among dipterocarps, these

include Dipterocarpus alatus Roxb. & G.Don, and Vatica mangachapoi Blanco. In

this case, though, there remains an overland link, Dipterocarpus condorensis subsp.

penangianus, of coastal forests in Peninsular Malaysia and Northwest Borneo, which

penetrates inland only in Ulu Temburong, Brunei, thus fonning a link to those species

which do occur in Indo-Bunna, Peninsular Malaysia and the Philippines, but which are

rare in northern Borneo except in the extreme west and north-east, notably Anisoptera

costata Korth., Dipterocarpus hasseltii Blume and Shorea guiso (Blanco) Blume.

New Malesian Dipterocarpaceae

Dipterocarpus pseudocornutus P.S. Ashton, sp. nov.

Dipterocarpus cornutus superficie similis, alae calycis in fructu congeste contorto

prominente differt. - TYPE: Philippines, Tawi-Tawi, Seratang Languyan, FJ.M.

Gaerlan & E.C. Sagcal PPI 10209 (holotype K).

Tree. Young parts densely shortly evenly buff pubescent, raceme and petiole persistently

so; leaf blade beneath buff lanate. Bud not seen. Twig c. 5 mm diam. apically, stout,

somewhat compressed. Leaf blade coriaceous, corrugate, ovate, 8-15(-21) x 4-8(-

11) cm, apex subacute, base obtuse to narrowly subcordate; petiole 3-5 cm long, c. 3

mm diam. Raceme to 15 cm long, hardly branched. Flower bud to 2 cm long; calyx

tube 12x8 mm, ellipsoid with intricately folded wings to 6 mm wide; stamens c. 30,

anthers 4 mm long at anthesis, lorate, with 4 mm filiform appendages. Fruit unknown.

Notes. This extraordinary collection, from islands populated by the Moro people and

long thought deforested, was apparently found in a patch of degraded forest. Had

it been without flowers, it would have likely been claimed as a major extension of

the range of Dipterocarpus cornutus Dyer, a species of Peninsular Malaysia and

the slightly seasonal northern Sumatra and south-eastern and southern Borneo.

The densely intricately folded wings of the calyx either implies that this character

is evolutionarily plastic, or that this new taxon is allied to others which bound the

South China Sea; Dipterocarpus lamellatus Hook.f., of SW Sabah and Brunei, D.

lowii Hook.f, which follows the expanded Riau Pocket excepting Simeulue (Ashton,

2014), andO. intricatus Dyer, of southernmost Indo-China.

Vatica palungensis P.S. Ashton, sp. nov.

Similar to Vatica oblongifolia Hook.f , especially to V. oblongifolia subsp. selakoensis

P.S. Ashton, and to V. dulitensis Symington, but with terete twigs (vs compressed), a

sparse buff indumentum (vs dense vinous), and 8-10 pairs of secondary veins (versus

> 10). - TYPE: Indonesia, Kalimantan Barat, Ketapang, Gunung Palung National

360

Gard. Bull. Singapore 67(2) 2015

Park, Lower montane kerangas forest over granite, 665 m, T. G. Laman, LA. Rachman,

& E. Mirmanto TL 772 (holotype K; isotype A).

Young parts sparsely rugose grey-brown pubescent. Tnng c. 2 mm diam. at first, terete.

Leaf blade coriaceous, drying pale grey-brown above, ocherous beneath, nan'owly

elliptic-ovate to elliptic-obovate, 7-1 5 x 3-6 cm, acumen to 1 cm long, broad, base

obtuse; veins 8-10 pairs, ascending, raised on both surfaces, more so beneath; petiole

8-15 mm long, 1.5 mm diam., slender. Raceme short. Flowers unknown. Fruit to 10

mm diam., globose, rugulose; sepals to 3 x 2.5 mm, obtuse, becoming reflexed.

Additional specimens examined. INDONESIA: Kalimantan Barat: Ketapang, Gunung Palung

National Park, Lower montane kerangas forest over granite, 705 m, T.G. Laman, I.A. Rachman,

& E. Mirmanto TL 1374 (K, A), ibidem, 1000 m, fallen leaves and fruit, RS. Ashton s.n. (A).

Notes. Superficially a member of the endemic Borneo species group around the

widespread Vatica oblongifolia Hook.f. which share a short powdery vinous

indumentum. The present species differs from others notably in its tomentum. As in so

many groups of sister dipterocarps, these species are particularly well represented in

the north-west. At present a point endemic, this novelty also shares the naiTow range

and high habitat specificity of several of the others.

References

Ashton, RS. (1982). Dipterocarpaceae. In: Van Steenis, C.G.G.J. (ed) Flora Malesiana, Series

I, 9(2): 237-552.

Ashton, RS. (2004). Dipteroearpaceae. In: Soepadmo, E. & Saw, L.G. (eds) Tree Flora of

Sabah and Sarawak 5: 63-388, 485^96.

Ashton, RS. (2014). On the Forests of Tropical Asia: Lest the Memory’ Fade. UK: Royal

Botanic Gardens, Kew and USA: The Arnold Arboretum of Harvard University.

Chua, L.S., Nor-Ezzawanis, A.T., Yong, W.S.Y. & Hamidah, M. (submitted). Two new species

of Vatica from Reninsular Malaysia. Phytotaxa.

Saw, E.G. (2002). Anew species of Vatica (Dipterocarpaceae) from Peninsular Malaysia. Gard.

Bull. Singapore 54: 247-251.

Smitinand, T., Vidal, J.E. & Ho, RH. (1990). DiptNocarpacees. In: Morat, Rh. (ed) Flore

du Cambodge, du Laos et du Vietnam, vol. 25. Raris: Museum National d’Histoire

Naturelle.

Symington, C.F. (2004). Foresters ’ Manual of Dipterocarps. Malayan Forest Records No.

16. Revised by Ashton, RS. & Appanah, S., edited by Barlow, H.S. Kepong: Forest

Research Institute Malaysia and Kuala Eumpur: Malaysian Nature Society.

Gardens’ Bulletin Singapore 67(2): 361-386. 2015

doi: 10.3850/S2382581215000307

361

A contribution to the systematics of Xylopia

(Annonaceae) in Southeast Asia

D.M. Johnson & N.A. Murray

Department of Botany & Microbiology, Ohio Wesleyan University

Delaware, OH 43015 USA

drniohnso@owu.edu

namurray@owu.edu

ABSTRACT. Herbarium and field study ofXylopia L. (Annonaceae) for the Flora of Peninsular

Malaysia and the Flora of Thailand projects has clarified regional diversity patterns within

this ecologically significant lowland rainforest genus. Two species groups represented within

Southeast Asian floras ai'e delineated, one centred on Xylopia ferruginea (Hook.f & Thomson)

Baill. and the other on Xylopia malayana Hook.f & Thomson. In Xho Xylopia ferruginea group,

a new species, Xylopia erythrodactyla D.M. Johnson & N.A.Murray, is distinguished from X.

ferruginea, and a new combination, Xylopia sumatrana (Miq.) D.M. Johnson & N.A.Murray,

is proposed, based on an earlier name for the species currently known as Xylopia stenopetala

Oliv. In the Xylopia malayana group, review of the species Xylopia elliptica Maingay ex

Hook.f. & Thomson resulted in the recognition of tliree additional species: Xylopia platycarpa

D.M.Johnson & N.A.Murray, from southern Thailand and northwestern Peninsular Malaysia,

Xylopia ngii D.M.Johnson & N.A.Murray, fi'om Peninsular Malaysia, Sumatra and Borneo,

and Xylopia heterotricha D.M.Johnson & N.A.Mun*ay, from Sumatra and Borneo. The taxon

Xylopia malayana Hook.f. & Thomson var. obscura Kochummen is placed in synonymy under

Xylopia elliptica sensu stricto. Xylopia fusca Maingay ex Hook.f. & Thomson var. sessiliflora

Kochummen & Whitmore is distinguished from Xylopia fusca, and raised to species status as

Xylopia sessiliflora (Kochummen & Whitmore) D.M.Johnson & N.A.Murray. We recognise 23

Xylopia species in the Sundaic region of Southeast Asia, and provide evidence that additional

collecting and taxonomic analysis in the region is needed.

Keywords. Annonaceae, biogeography, Borneo, Malay Peninsula, Sumatra, Sundaland,

Thailand, Xylopia

Introduction

The Annonaceae, a flowering plant family of 2500 species, including the economically

important soursop, custard apple and ylang-ylang, is widespread across the tropics.

The family is most diverse, and ecologically most significant, in tropical Asia, where

it is represented by c. 40 genera and 800 species. In southeastern Asia it is one of

the dominant families in lowland wet forests. Corlett & Turner (1997) determined

that Annonaceae ranked fourth in species-richness among flowering plant families

in Singapore; Appanah et al. (1993) found that Annonaceae ranked first in species

diversity among lianas of Malaysian forests. In long-term ecological plots in Southeast

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Gard. Bull. Singapore 67(2) 2015

Asia Annonaceae usually rank among the top ten tree families in both number of

individuals and number of species, though not in basal area (see examples in Losos &

Leigh, 2004).

The genus Xylopia L., the only pantropical genus in the family, comprises

180-200 species of trees and shrubs worldwide. The highest concentration of species

in Southeast Asia occurs in the Sundaic region extending from the Kra Isthmus in

southern Thailand to Wallace’s Line, a region of high diversity for many plant taxa.

Xylopias are distinctive among the Annonaceae in their cone-shaped buds, elongate,

aromatic flowers, and dehiscent fruits with seeds bearing arils or fleshy seed coats. In

West Africa the peppery fruits of Xylopia aethiopica (Dunal) A.Rich. have long been

used as a spice (Dunal, 1817; Burkill, 1985) and are sold commercially. Despite its

significance, the genus has never been monographed.

In preparing keys and descriptions of the genus Xylopia (Annonaceae) for the

Flora of Peninsular Malaysia and Flora of Thailand projects we had the opportunity to

study material in herbaria with important holdings for the region as well as to observe

several species in the field. At the beginning of our study 1 3 Xylopia taxa were known

from Peninsular Malaysia, Singapore and Thailand combined (Sinclair, 1953, 1955;

Kochummen et ah, 1970; Kochummen, 1972a, 1972b; Chalennglin, 2001; Gardner et

ah, 2015).

Analysis of morphological and preliminary molecular data (Stull et ah, 2011;

Thomas et ah, 2015; Stull et ah, in prep.) has shown that the Southeast Asian species of

Xylopia fall into two groups, one including Xylopia ferruginea (Hook.f. & Thomson)

Baill. and a second including Xylopia malayana Hook.f & Thomson, each with

their respective allies. Our study revealed a much greater diversity of species than

previously recognised in both groups. To document this diversity, and reconcile it with

the previously existing taxonomy and nomenclature for the genus, the following paper

is presented. A full treatment, including keys and distribution maps, is forthcoming.

Conseiwation assessments using lUCN (2012) criteria are not included in

this account as more data are required for these than we currently have available.

Although historical EOOs and AOOs can be calculated, we are conscious that many of

the collections have been made in lowland forest areas that have suffered from rapid

deforestation. In these cases the Population Reduction (A) criterion would be more

appropriate, as has been used for many dipterocarp species (lUCN, 2014), but again

we would require more on-the-ground knowledge than we currently have.

Xylopia ferruginea group

The Xylopia ferruginea group is characterised by stilt roots, relatively long (5-19

mm) flower pedicels, a flat receptacle lacking a staminal cone (Fig. 2K), flat narrowly

oblong stamens with a tongue-shaped apex to the anther connective (Fig. 2J), stigmas

studded with small papillae (Fig. 21), and rugose seeds (Fig. 2C-D). In addition, most

species of the Xylopia ferruginea group have relatively numerous (up to 20) linear and

somewhat torulose monocarps.

Xylopia in Southeast Asia

363

Study of the group resulted in demarcation of a new species, and recognition of

an earlier name for an existing species.

Xylopia erythrodactyla D.M.Johnson & N.A.Murray, sp. nov.

Species resembling Xylopia ferruginea in the rusty pubescence of the leaves and

flowers and the long narrow monocarps, but differing consistently in the more densely

pubescent and thicker leaves, thicker pedicels (1.3-2. 5 mm), longer sepals (4. 8-7. 9

mm), broader outer petals (3-3.7 rmn wide at the midpoint) and narrowly oblong and

weakly torulose monocarps 7.5-10.7 cm long and 0.6-1. 1 cm wide. In contrast, in

Xylopia ferruginea the pedicels are 1-1.3 mm thick, the sepals are 3-5.5 mm long, the

outer petals are 2-2.2 mm wide at the midpoint, and the monocarps are linear, strongly

tomlose, 6.2-11.6 cm long and 0.4-0.6 cm wide. - TYPE: Malaysia, Sarawak, Teluk

Bandung, Santubong, 1st Division, 18 September 1984 (fr), Awa & Ismawi S.47080

(holotype KEP; isotypes ASU, K, L, SAR). (Fig. 1, 2)

Tree up to 30 m tall, dbh up to 75 cm, bole smooth with stilt roots at the base;

secondary branches drooping. Bark smooth, light brown, brown tinged with red,

brick-red, or orange, very finely fissured. Twigs light grey to brown, eventually

dark grey, densely femiginous-pubescent/velutinous, eventually glabrate. Leaf with

larger blades 12.4-26 cm long, 3.9-7 cm wide, subcoriaceous to coriaceous, strongly

discolorous, oblanceolate, oblong-oblanceolate, or narrowly elliptic, base rounded to

cuneate and short-decurrent, apex short-acuminate, the acumen 2-7 mm long, glabrous

adaxially, densely ferruginous-pubescent, the pubescence especially pronounced

along the midrib, secondary veins, and larger higher-order veins abaxially; midrib

impressed adaxially, raised abaxially; secondary veins 1 1-14 per side, diverging at

50-60° from the midrib, brochidodromous, these and higher-order veins indistinct

adaxially, strongly raised abaxially; petiole 5-12 mm long, deeply canaliculate

(margins nearly meeting), pubescent. Inflorescences axillary or from axils of fallen

leaves, 1-3-flowered, densely ferruginous-pubescent; peduncles 1-2 per axil, 2 mm

long; pedicels 2 per peduncle, 7-14 mm long, 1.6-2. 5 mm thick; bract 1, attached

!/3-!/2 distance from base of pedicel, 2.9-3 mm long, ovate, apex acute to obtuse; buds

linear-lanceolate, sometimes somewhat falciform and slightly twisted, apex obtuse.

Sepals ’/g-’A-connate, 4-7.9 mm long, 4-5.3 mm wide, coriaceous, broadly ovate to

triangular, apex acute to acuminate, occasionally obtuse, pubescent along margins

and at apex adaxially, ferruginous-pubescent abaxially. Petals pale yellow to white

in vivo; outer petals curving outward at anthesis, 38-45 imn long, 4-5.3 mm wide at

base, 3-3.7 mm wide at midpoint, linear-lanceolate, obtuse, densely grey-puberulent

adaxially, densely feiTuginous-pubescent abaxially; inner petals erect at anthesis, 32-

40 mm long, c. 3.5 mm wide at base, c. 1.3 mm wide at midpoint, linear, densely

grey-pubemlent on both surfaces except for glabrous concave base. Stamens up to

77, 2. 7-3. 5 mm long, narrowly oblong, often setose along edges of anther locules,

apex of connective 0.4-0. 6 mm long, oblong, densely long-papillate, filament 0.3-0. 5

mm long, glabrous; staminal cone absent; outer staminodes c. 18, c. 2.7 mm long.

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Gard. Bull. Singapore 67(2) 2015

Fig. 1. Xylopia erythrodactyla D.M.Johnson & N.A.Murray. A. Two leaves, abaxial view. B.

Flower bud and abaxial view of monoearp, the latter showing beginning dehiscence with single

seed visible. C. Single monoearp in lateral view, resting on adaxial surface of a leaf

(Photos: N.A. Murray)

oblanceolate, flat, apex obtuse. Carpels 16-20; ovaries 1.5-2. 5 mm long, narrowly

oblong, densely fermginous-pubeseent with hairs obseuring lower portion of stigmas;

stigmas loosely connivent, e. 2.2 mm long, dark, with a few seattered hairs and

studded with amber-eoloured papillae. Torus flat, c. 3.4 mm in diameter. Pedicel of

fruit 11-20 mm long, 4-7 mm thick, pubescent; torus of fruit c. 8 mm high, 9-14

mm in diam., depressed-globose, sparsely pubescent to glabrate. Monocarps red with

brown tomentum in vivo, up to 20 per fruit, 5.5-10.7 cm long, 0. 6-1.1 cm wide and

thick, linear to narrowly oblong, weakly torulose, terete in cross-section, apex rostrate,

the beak 2.5-9 mm long, base contracted into a stipe 8-10 mm long, 3.5^ mm thick,

longitudinally wrinkled, ferruginous-pubescent to glabrate; pericarp 1.7 mm thick.

Seeds in a single row, parallel to long axis of monoearp, 6-12 per monoearp, 7.2-8. 8

mm long, 5. 5-5. 6 mm wide, 4.7-5. 1 mm thick, ellipsoid, elliptic in cross section, dark

brown, rugose, flattened or a little concave at micropylar end, rounded at chalazal end,

raphe and antiraphe distinctly raised; aril and aril plate absent.

Distribution. Occurs in Terengganu in northeastern Peninsular Malaysia, and on the

northern coast of the island of Borneo in Sarawak (East Malaysia) and in Brunei.

Considering its restricted habitat and the pace of development in Sarawak, this species

Xylopia in Southeast Asia

365

Fig. 2. Xylopia eiythrodactyla D.M. Johnson & N.A.Murray. A. Habit. B. Inflorescence with

flower buds, side view. C. Seed, view of micropylar end. D. Seed, side view. E. Fruit. F.

Outer petal, adaxial view. G. Inner petal, adaxial view. H. Stigma apex. I. Carpel. J. Stamen.

K. Close-up of flower with petals and stamens removed, to show sepals, caipels and torus.

L. Schematic side view of flower at anthesis, one outer petal removed. Drawn by Kate Ball

from (A) Awa S. Ismawi S. 47080, ASU; (B) Sibat ak Luang S. 24502, L; (C-E) Chew & Kiah

SFN. 40982, A; (F, G, K) Zehnder S. 16803, A; (H-J) Rogstad 704, A; and (L) field sketch.

366

Gard. Bull. Singapore 67(2) 2015

may be more vulnerable than it appears from the collecting data.

Ecology. The bulk of the Sarawak and Brunei collections have come from either

lowland peat swamp forest or heath (kerangas) forest (tenninology following Saw,

2010). The collection localities in Terengganu are all from lowland dipterocarp forest,

as are several in Sarawak. In general, however, the ecological range of this species is

much narrower than that of Xylopia fermginea. Elevational range is sea level to 250

m, with one collection from Terengganu from c. 800 m.

Phenology. The species shows two distinct flowering and fruiting periods throughout

its distribution: flowers have been collected in late March and May-July, and then

again in November-December, while fruits have been collected from February to July

and also from September to December.

Local name. Ako (Sarawak: Bojeng bin Sitam 9319, Zainuddin bin Bolhassan S. 11941).

Etymology. The species is named for the red and finger-like monocarps (Fig. IB-C),

which are thicker and less torulose than those of its congonQi Xylopia fermginea.

Additional specimens examined. MALAYSIA: Peninsular: Terengganu: 18th mile, Jalan

Kelantan, 31 Mar 1957 (fr), Chiew & Kiah SEN 40982 (A, K, L [4 sheets], M, SING [2 sheets]);

Pulau Redang, Pasir Mah Kapit, 300 m, 5 Mar 1 989 (fr), Saw FRl 36487 (A, K, KEP, L, SAR,

SING); near Kampong Gong Nangka, Marang, 6 Jul 1 953 (st), Sinclair & Kiah bin Salleh SEN

39837 (L, SING [3 sheets] ); 19th mile Kuala Trengganu Besut road, 15 Nov 1954 (st), Sinclair

& Kiah bin Salleh 40473 (L, SING); 1 8th mile Kuala Trengganu-Besut Road (west side), 7 Sep

1955 (young fr), Sinclair & Kiah bin Salleh SEN 40748 (A, K, L, SING [2 sheets]); Ulu S. Loh

below E face G. Mandi Angin, 2300 ft., 13 Jul 1968 (If), Whitmore FRI 12144 (K, KEP, L);

Borneo: Sarawak: TelokAsam, Bako N. R, 4 Jun 1963 (W), Ashton S. 17913 (A, ASU, K, KEP,

L, SAR, SING); Ulu Kenyana, Mukah, 20 Oct 1963 (young fr), Ashton S. 19488 (A, K, KEP, L,

SAR, SING); Kuching, Setapok F.R., 24 Jul 1957 (fl), Bojeng bin Sitam 9319 (BO, K, L, SAR,

SING); G. Pueh F.R., June 1956 (st), BrunigS.6369 (SING); Selang F.R., Feb 1956 (st), Brunig

S. 7225 (SING); long path from Tg. Po to Telok Kruim, Bako National Park, Kuching, 1st div.,

15 m, 16 May 1980 (fr). Citing S. 42286 (K, KEP, L, SAR); Ulu Sungai Pasir Biawak, Lundu,

Kuching, 8 Apr 1997 (if), Jamree et al. S. 76728 (K, KEP, L, SAR); Kem Pennai, Santubong,

29 Jun 1992 (fr), Othmcin et al. S.65133 (KEP, L); Sampadi F.R. (near road), 25th mile, Bau/

Lundu Road, 1st Division, 750 ft, 28 Jun 1968 (fl). Pale S. 269 15 (K [2 sheets], L, SAR, SING);

Bako National Park, just above mangrove swamp at open area on slope at start of Jalan Lintang,

6 Dec 1981, Rogstad 703 (Afr), 704 (Afl); Nyabau Catchment area, Bintulu, 4th Division, 300

ft, 11 Jun 1966 (fl), Sibat ak Luang S.24502 (BO, KEP, L, SAR, SING); Kuching, 100’ alt.,

Setapok F.R., 15 Nov 1957 (fi-), Yacup 8939 (K, L, SAR, SING); Bintulu, Similajau F.R., 17

Nov 1959 (fl), Zainuddin bin Bolhassan S. 11 94 1 (K, L, SAR, SING); Kuching, Sg. Teruntum,

Sarawak Mangrove Reseiwe, 27 May 1962 (fl), Zehnder S. 16803 (A, K, L, SAR, SING).

BRUNEI: Andulau F.R., 18 Sep 1957 (st), Ashton B RUN 560 (L, SING); Belait District, Ulu

Sungai Badas, 28 Mar 1989 (fl), Nangkat NN105 (A, AAU, K, L, SAR, SING); Belait District,

Lumut Hills, 30 Mar 1968 (st), van Niel 4454 (L); Ulu Badas, Andulau Stateland, Sg. Liang,

Belait (Labi Rd. 10 km), 25 Jun 1996 (fl), Ogata et al. Og-B203 (L).

Xylopia in Southeast Asia

367

Notes. Trees of this species were observed in Bako National Park in June of 2003,

where the plants were frequent along the edge of dipterocarp forest on a slope

bordering mangroves. At this site there were many seedlings and saplings present;

flowers or fruits were only found on individuals exceeding 1 5 m in height. The petals

of fallen flowers were sweetly scented, the scent reminiscent of Gardenia (Rubiaceae).

At anthesis the outer petals were widely spread while the inner petals were more or

less erect (Fig. 2L). Dehiscing monocarps, gathered from the ground at this locality,

were photographed (Fig. IB).

The dried seeds, when soaked, become tan in colour, revealing a sarcotesta c.

0.2 mm thick, which may be somewhat incomplete at the chalazal end. The sarcotesta

breaks away in chunks from the woody layer of the seed coat underneath, often with

patches of the woody coat adhering to it.

The type description of Xylopia altissima BoerL, based on a specimen collected

by Teysmann on the island of Lingga (Boerlage, 1 899), suggests the new species in

its emphasis on the dense tomentum of the abaxial leaf surface, but Boerlage (p. 203)

also emphasised the strongly reticulate leaves (“venis tenuibus in nervos ti ansversis et

reticulatis pertensa”) with acuminate apices. These features are frequent in Sumatran

specimens of Xylopia ferniginea s.s., which in general have larger and broader leaves

than those of Peninsular Malaysian and Bornean plants, as well as a tendency toward

a subcordate rather than truncate leaf base. These leaf features do not occur in Xylopia

erythrodactyla.

Two sterile specimens found in the herbarium at M bear leaves resembling those

of Xylopia e/ythrodactyla. The specimens were collected from a plant grown in the

Bogor Botanic Garden. The provenance of the plants is not certain, but “Borneo” is

written in pencil on one of the sheets. The actual labels give the fol lowing information:

Hort. Bogor. IV B 1 8, 2 May 1 895, Spelta [?] s.n. (M [2 sheets], as “Sapotacea Sambas

V. d. Florsf ’ on one sheet, “Columnifera?” on the other). Duplicates of these specimens

were not seen at L or BO; the catalogue by Dakkus (1957) did not show any listings

under either the names or the number.

Xylopia sumatrana (Miq.) D.M.Jolmson & N.A.MuiTay, comb. nov. - Unona

sumatrana Miq., FI. Ned. Ind., Eerste bijv. 3: 377 (1861). -Xylopia malayana Hook.f.

& Thomson var. macrocarpa BoerL, Icon. Bogor. 1(2): 123 (1 899). -TYPE: Indonesia,

Sumatra occid. in prov. Priaman, Diepenhorst s.n. (holotype U).

Xylopia stenopetala Oliv., Hook. Icon. PL 21: t. 1563 (1887). - TYPE: Malaysia,

Penang, Government Hill, 600 ft, June 1886, Curtis 857 (lectotype K, designated by

Turner (2011); isolectotype SING).

Distribution. Peninsular Malaysia, Sumatra, Borneo, Philippines (Mindanao).

Additional specimens examined (representative specimens). MALAYSIA: Peninsular: Pahang:

Frasers Hill at the lower gate, 3000’ alt, 29 May 1968 (fl, fr), Ng FRl 6172 (A, K, SING); G.

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Gard. Bull. Singapore 67(2) 2015

Benom Game Reserve, Ulu Ki'au, 1800’ alt, 22 Apr 1967 (fl), YusoffKEP 99124 (A, K, SING);

Penang: Tunnel road, Penang Hill, 2200 ft, 28 May 1938 (fr), Henderson SFN 21425 (SING

[2 sheets]); Selangor: Kepong, F.R. Inst., 50th mile. Gap Road, 15 April 1990 (fl), Kochummen

FRl 29090 (A, K, SAR); Ulu Gombak F.R., 1800 ft alt, 18 Jan 1966 (fr), Kochummen FRl

80497 (A, K, SING); Borneo: Sabah: Lamag District, close to exit stream of Gunong Lotung,

lake Inarat, 1200 ft, 21 May 1976 (ft), Cockburn SAN 83321 (K, L, SAR, SING); Tawau Dist.,

Mile 26, Apas Rd, 120 ft, 25 Jun 1959 (If), Meijer SAN 19321 (K, L, SING); Sarawak: Pk in

Kuching, Dec 1892 (fl), Haviland =10[?] (BM, L, MO, SING); Perkulen Ampat, b.p.m.d. (fl,

fir), Haviland 142/10 [A. 13. 9] (SAR, SING); Bkt. Lobang, Punung Lusong, Sg. Linau, Belaga,

7th division, 470 m, 14 Jun 1 979 (fr), Lee S. 39802 (ASU, K, L, SAR); path from Kpg. Seropak

to Bungoh Range, Bau, secondary forest on hillside, c. 1000 ft, 29 Nov 1969 (fl), Paie & Mamit

S.29584 (A, K, L, SAR, SING),

INDONESIA: Borneo: East Kalimantan: Meratus, 5230 area PT. ITCI, Kenangan

Balikpapan, 70 m, 27 Sep 1992 (fr), AmhriansyahA. A:611 (A, K, L); PT. ITCI, road Kenangan

to G. Meratus, than to Basecamp Birawa, km 52, 500 m, 28 Mar 1995 (fl), Kessler et al.

PK.949 (A, K); E Kutei, Sangkulirang subdivision, Sg. Susuk region, 10 m, 26 Jun 1951 (fl),

Kostermans 5456 (A, BM, K, L [2 sheets], SING); West Kalimantan: G. Bentuang area, 5-10

km N of Masa village, 150 km NE of Pontianak, steep ridge above Semawang River, 00°52’N,

110°26’E, 50 m, 23 Jun 1989 (fl, fr), Burley et al 2823 (A [2 sheets], F, K, L, MO, NY [2

sheets], SING); Sumatra: Aceh: Gunung Leuser Nature Resei^ve, Gunung Bandahara c. 6 km

NE of Kampung Seldok (Alas Valley), c. 25 km N of Kutatjane, c. 800-1000 m alt, 20 Mar

1975 (fl, fir), de Wilde & de Wilde-DuyJjes 15599 (K [2 sheets], MO, US); North Sumatra:

East Coast, vicinity of Loemban Ria, Asahan, 5 Feb-12 Apr 1934 (st), Rahmat si Boeea 8096

(A, MICH, US); East Coast, Asahan NE of Tamuan Delok and W of Salabat, 500 m, 15 Jun-9

Jul 1936 (fl), Rahmat si Boeea 9269 (A [2 sheets], K, L, MICH, NY, US, W).

PHILIPPINES: Mindanao: Surigao Sur: Manobo District, PICOP Bislig, Apr 1976 (fl), Rojo

325 (MO).

Notes. The type specimen of Unona sumatrana Miq. from the Utrecht herbarium (now

housed at L) includes two separate branches, one with a fruit attached, and the other

with two small leaves. The two leaves are slightly obovate-oblong, one 6.5 cm long

and 3 cm wide and the other 3.9 cm long and 2.3 cm wide. The leaves are retuse at

the apex and broadly cuneate and decurrent at the base. The fruit shows the distinctive

characteristics of Xylopia stenopetala: there are two monocarps and a portion of a

third on the specimen, none of them fully mature. The intact monocarps are naiTowly

oblong, one 6.8 cm long and 0.8 cm wide and slightly torulose, sparsely pubescent,

with about 7 seeds arranged in a single row. The monocarps are acute at the apex and

taper to a stipe c. 8 mm long. The monocaips are borne on a pedicel 10 mm long and

4.3 mm thick, and a torus 1 1 mm in diameter, 8 mm high, and depressed-globose in

shape.

Although the name Unona sumatrana was reduced to a variety of Xylopia

malayana by Boerlage (1899), and placed in synonymy under A malayana by Turner

(2011), the fmit on the type specimen of Unona sumatrana clearly distinguishes it

from A. malayana: the narrow monocarps have up to 7 seeds (seeds usually 3 or fewer

per monocaip in A malayana), and, at 6.8 cm in length, are already longer than those

found in A malayana (which never exceed 4 cm in length).

Xylopia in Southeast Asia

369

The type of the name Unona sumatrana from 1861 was thus found to represent

the same species as Xylopia stenopetala from 1887 and the earlier name must take

precedence. This name is not to be confused with Xylopicrum siimatranum (Miq.)

Kuntze, Revis. Gen. PI. 8, 1891, which is based on Par artabotrys sumatranus Miq.,

now considered a taxonomic synonym of Xylopia malayana (Sinclair, 1955; Turner,

2011 ).

Xylopia sumatrana has been collected in Peninsular Malaysia in the states of

Pahang, Penang and Selangor, and on the islands of Sumatra, Borneo, and Mindanao

(Philippines), where it is found at elevations of 50-1000 m, the widest elevational

range of any Asian species of Xylopia.

Xylopia malayana group

Members of Xylopia malayana group lack stilt roots (with the exception of Xylopia

sessiliflora treated below), the flower pedicels are 5 mm long or less, the receptacle

bears a distinct but low and irregularly laciniate staminal cone fonned from the connate

bases of the filaments, the stamens are clavate with a transversely flattened apex to the

anther connective, the stigmas lack papillae, and the seeds are smooth. Within this

group the monocarps tend to be relatively few in number (ten or fewer, but sqq Xylopia

heterotricha below) and relatively broad and oblong.

Species of the Xylopia malayana group are distinctive and well circumscribed

with the exception of X. elliptica and X. malayana. We found that the primary source

of taxonomic difficulty with these two species stemmed from the inclusion of multiple

distinct taxa under the name Xylopia elliptica. The simplest route to clarification is to

retrace the taxonomic history of that species.

Hooker & Thomson (1872: 86) originally Xylopia elliptica on a single

Maingay collection from Malacca. The protologue, reproduced verbatim below,

shows that the diagnosis emphasised the glabrous branches, the small elliptic, obtuse,

membranous, glabrous leaves and the solitary flowers:

14. X. elliptica, Maingay niss,\ branches glabrous, leaves small elliptic obtuse mem-

branous glabrous, tip rounded, neives faint reticulate, flowers small solitary erect pubescent,

sepals subacute united to the middle, ovaries 1-3.

Malacca, Maingay.

A lofty tree; trunk thick; branches glabrous, almost black; branchlets pubescent. Leaves

1 1 / 2-2 by 1 %-l 14 in., base obtuse or acute, pale on both surfaces, browner beneath; petiole 14

in., puberulous. Flowers 14 -% in., slender; peduncle half as long or shorter, and calyx rusty-pu-

bescent; bratcs [sic] median, minute. Petals pale brown-tomentose; outer linear-subulate, from

a rather broad base, concave; inner trigonous, base excavated. Stamens minute. Ovaries sunk

in the deeply urceolate torus, hidden amongst long white hairs; ovules 4-6.

The circumscription of the species was enlarged by King ( 1 892), who identified

two additional specimens, Wray 3194 from Perak and Curtis 2482 from Penang, as

belonging to this species. The latter specimen in particular departed from Hooker and

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Gard. Bull. Singapore 67(2) 2015

Thomson’s protologue, however, in having distinctly pubescent leaves and multiple

flowers per inflorescence home on longer pedicels. Ridley (1922) subsequently

maintained King’s concept of the species mtact. Sinclair (1955) identified four additional

specimens as Xylopia elliptica: Awang 42444 from Kedah, Wray 3562 from Perak,

20309, without collector’s name, from Pahang, and Yeob 5037 from Selangor. Sinclair

stressed the significance of the pubescence of twigs and leaves as a characteristic

important in distinguishing this species from the otherwise similar Xylopia malayana,

and apparently on this basis identified a number of specimens of Xylopia Ifom Sarawak

and Sabah in various herbarium collections as belonging to X. elliptica.

It has become clear from re-examination of these specimens, plus study of a

wider range of material than was available to these authors, Xylopia elliptica in the

sense of Sinclair is a mixture of four very distinctive species. The concept adopted here

is one of Xylopia elliptica in a restricted sense, and the recognition of three previously

undescribed species to accommodate the variants.

Xylopia elliptica Maingay ex Hook.f & Thomson, FI. Brit. India 1: 86 (1872). -

Xylopicrum ellipticum (Maingay ex Hook.f. & Thomson) Kuntze, Revis. Gen. PI. 1:8

(1891). - TYPE: Malaysia, Malacca, 9 May 1867, Maingay 2376 [Kew Distribution

no. 82] (lectotype K [2 sheets, barcode nos. K000574709, K000574712], designated

by Turner (201 1); second-step lectotype, designated here: K000574709; isotype CAL).

Xylopia malayana var. obscura Kochummen, Gard. Bull. Singapore 26(1): 49 (1972).

- TYPE: Malaysia, Terengganu, Gunong Padang Expedition, Ulu Brang, camp 1 nr.

K. Lallang, 1000’ alt, 15 September 1969 (fl, fr), Whitmore 12594 (holotype KEP;

isotypes A, K, L, SING).

Distribution. Peninsular Malaysia. Although it has been collected from four states in

Peninsular Malaysia, Xylopia elliptica is known from only eight collections, all from

forests below 300 m in elevation.

Additional specimens examined. MALAYSIA: Peninsular: Johor: Compt 10, Rengam F.R.,

14 Nov 1966 (fl), Kochummen FRI 2188 (A, K, KEP, L, SING); Compt 34, Gunong Arang

F.R., 13 Sep 1969 (fr), Kochummen FRI 2761 (K, KEP, L); Keluang, 20 Nov 1990 (fl, fl-), Teo

& Remy KL 3968 (KLU); Hutan Simpan Endau, 23 Oct 1997 (fl), Teo & Tetu KL 4951 (KEP);

Pahang: Chini Forest Reserve, 11 Dec 2008 (fl), Khairil bin Mahmoud et al, s.n. (UKMB);

without definite locality [Raub, according to Sinclair (1955)], collector unknown, KEP 20309

(KEP).

Notes. Xylopia elliptica in the sense of its type applies to relatively small-leaved

plants with glabrous to sparsely pubescent leaves, inflorescences of a single flower,

and relatively short petals. This same circumscription also includes the taxon Xylopia

malayana var. obscura. Xylopia elliptica in this restricted sense is endemic to Peninsular

Malaysia, where it is infrequent in lowland forests up to an elevation of 300 meters in

Johor, Melaka, Pahang, and Terengganu. The report of Xylopia elliptica for southern

Xylopia in Southeast Asia

371

Thailand in Gardner et al. (2015) is based upon a misidentified specimen oi Xylopia

pierrei Hance; the southern Thailand population oiX. pierrei is disjunct from the main

distribution of the species in eastern Thailand, Cambodia, and Vietnam.

The detached fruit on the sheet of KEP 20509 does not look like a fruit of

Annonaceae, Sinclair’s detennination of the specimen notwithstanding, but the leaves

of the collection possibly represent this species.

Turner (2011) designated the specimens K000574709 and K000574712 at K as

lectotypes of the name Xylopia elliptica. Both sheets include leafy twigs and flowers.

Hand-written descriptive notes that are incorporated into the protologue are present on

K0005 74709, and it is therefore designated as a second-step lectotype as pennitted by

Article 9.17 of the ICN (McNeill et al., 2012). The sheet K000574712 does not bear

these notes.

Xylopia platycarpa D.M.Johnson & N.A.Murray, sp. nov.

Resembling Ay/c»/?/a vielana by its pubescent leaves with a broadly cuneate to rounded

base, but differing in the longer and narrower petals often curled at the apices and

the flat monocarps that are white and marked with fine red veins. - TYPE: Thailand,

Trang Province, Yanta Khao District, Peninsular Botanic Garden (Thung Kliai), near

office, 07°28N 99°38'E, 25 m, 7 July 2005 (fl, fr), Gardner et al ST1882 (holotype L;

isotypes BKF, L) (Fig. 3A-D, Fig. 4A-G)

Tree up to 22 m tall, dbh up to 48 cm. Bark orange to pale brown, flaking. Twigs

brown or grey to blackish brown, eventually lenticellate, pubescent; double-branching

occasional. Leaf with larger blades 6. 5-9.4 cm long, 2.4-4. 1 cm wide, chartaceous

to subcoriaceous, lanceolate to oblong-lanceolate, elliptic-lanceolate, or ovate, base

broadly cuneate to rounded-truncate, apex acute to obtuse, pubescent but becoming

glabrate adaxially, pubescent abaxially; midrib slightly impressed adaxially, raised

abaxially; secondary veins 7-12 per side, diverging at 45-50° fi'om the midrib,

brochidodromous, these and higher-order veins indistinct to slightly raised adaxially,

indistinct or slightly raised abaxially; petiole 6-8 mm long, shallowly canaliculate,

pubescent. Inflorescences axillary, 1-3-flowered, dusty yellow-pubescent; peduncles

1 or sometimes 2 per axil, 1.5-4 mm long; pedicels 1 or 2 per peduncle, 1-4.2 mm

long, bracts 2, the uppeimost just below the calyx, 1.7-2. 3 mm long; buds linear and

sometimes falcate, apex acute. Sepals Y 3 -V 3 comiate, 2-3 mm long, c. 3.1 mm wide,

coriaceous, ovate to broadly triangular, apex acute, pubescent. Petals pale yellow in

vivo; outer petals 20.6-28 mm long, 2-3.1 mm wide at base, 0.8-1. 5 mm wide at

midpoint, linear, apex acute, sometimes curling at the tips, densely puberulent except

for glabrous base; inner petals 16.5-22.3 mm long, 1.5-2 mm wide at base, 0.7-0. 8

mm wide at midpoint, linear, apex acute, flat at base except for two slightly thickened

areas along the margin, densely puberulent except for glabrous base. Stamens c. 70,

1.2-1 .4 mm long, narrowly oblong to clavate, apex of anther connectives 0.2-0. 3 imn

long, bluntly conical to subglobose, puberulent; staminal cone c. 0.6 mm high, 1.4-2

mm in diameter, low, jagged; outer staminodes 1.2-1. 5 mm long, oblong, flat, apex

372

Gard. Bull. Singapore 67(2) 2015

obtuse to acute; inner staminodes c. 0.8 mm long, narrowly oblong. Carpels 3-5;

ovaries 0.6-0. 8 mm long, lanceolate to narrowly oblong, pubescent; stigmas connivent,

1.5-1. 8 mm long, black, shiny, with a few hairs at apex but otherwise glabrous. Torus

flat, 2.2-2. 5 mm in diameter. Pedicel of fruit 3-5.5 mm long, 1-2 mm thick; torus of

fruit 1-1.5 mm high, 3 mm in diameter. Monocarps cream-coloured with fine dull

red veining outside, bright red inside in vivo, up to 4 per fruit, up to 5.2 cm long, c.

0.8 cm wide, c. 0.5 cm thick, oblong and slightly falcate, flattened-ellipsoid in cross

section, apex forming a beak c. 5 mm long, base contracted into a narrow stipe 8-14

mm long, 1-1.5 imn wide, strongly rugose, sparsely pubescent to glabrate; pericarp c.

0.5 mm thick. Seeds arranged in a single row, oblique to long axis of monocarp, up to

3 per monocai*p, grey in vivo, 7. 9-8.3 mm long, 62-6.1 mm wide, 4. 7^. 8 mm thick,

broadly ellipsoid, elliptic to semicircular in cross section, smooth, flat on micropylar

end, rounded on chalazal end, sarcotesta translucent, black layers of seed coat visible

underneath; aril forming a fleshy crown-like ring around the rnicropyle, c. 2.8 mm

high, c. 4 mrn in diameter.

Distribution. Restricted to a small area of southern Thailand and northwestern

Peninsular Malaysia.

Ecology. All localities are in lowland evergreen forest.

Phenology. Flowers in July and August, fruit collected in July.

Local name. Kerangi lotong (Kedah: Awang 42444).

Etymology. The species is named for its unusually flattened and beanlike rnonocarps.

Additional specimens examined. MALAYSIA: Peninsular; Kedah: Perangin Forest Reserve,

30 Jul 1938 (fl), Awang 42444 ([Sinclair cites specimen at K, but this was not found in July

2014], KEP); Penang; Sungei Penang, Aug 1890 (fl), Curtis 2482 (BM, CAL (Sinclair, 1955),

K, SING [2 sheets], US).

Notes. Xylopia platycarpa stands apart from other Asian species of the genus by its

uniquely flattened cream-coloured rnonocarps with red veining. It occupies a very

narrow distribution in northwestern Peninsular Malaysia and southern Thailand. At

present this species is only known from three localities and has not been collected from

Peninsular Malaysia since 1938. Its biogeography is unusual, in that the distribution

of the species crosses the Kangar-Pattani Line, which has been widely recognised as a

prominent botanical transition line on the Malay Peninsula (Woodruff, 2003).

The new species most closely resembles A>7o/7/a vielana of northeastern Thailand,

Laos, Cambodia, Vietnam and southern China by virtue of its persistently pubescent

twigs and chartaceous, pubescent, acute to obtirse leaves. It has been identified as that

species in a recent floristic work (Gardner et al., 2015), but Xylopia platycarpa has,

in addition to the uniquely flattened pallid rnonocarps, longer completely pale yellow

petals (16.5-26 mm long) that tend to curl at the apices. The petals oi Xylopia vielana,

Xylopia in Southeast Asia

373

in contrast, are only 11-14 mm long, do not curl at the apex, and are flushed with red

or purple at the base.

The Curtis specimen, cited here as belonging to Xylopia platycarpa, was

identified by King (1892), Ridley (1922), and Sinclair (1955) as Xylopia elliptica, but

we found that it differs from the type of that name by its distinctly pubescent twigs

and leaves, the leaves lanceolate rather than elliptic in shape, and the tendency to have

multiple flowers per leaf axil, these often borne on a relatively long common peduncle

(Fig. 4B). The Awang specimen, also cited xmdiQv Xylopia elliptica by Sinclair (1955),

shares these same features, and in fact Sinclair used relative lengths of peduncle and

pedicel as a way to distinguish X. malayana and X. elliptica’. pedicel longer than

peduncle in X. malayana, and peduncle longer than pedicel inX. elliptica. This pattern

of long peduncles and relatively short pedicels however, was never seen in specimens

of Xylopia elliptica s.s. in our study.

Xylopia ngii D.M.Johnson & N.A.Murray, sp. nov.

Species differing from Xylopia malayana in the narrowly acuminate to caudate apex

of the leaf, the sepals only 2-2.9 mm long and sparsely pale brown pubescent with

acute to apicLilate apices, the outer petals longer and nan*ower (up to 34 mm long and

0.7-1 .6 mm wide at the midpoint), the larger monocarps 3 . 1-6.4 cm long with pericarp

2-4 mm thick and sessile or broadly short-stipitate, and 8-12 seeds per monocarp,

the seeds 14-18 mm long. - TYPE: Malaysia, Sabah, Tawau, Cpt. A., sub-cpt. 13,

Bombay Bunnah T. C. Licence Area, Kalabakan, 30 mi WNW of Tawau, 350 ft, 30

April 1954 (fr, A & L sheets also have flowers). Wood A 3454 (holotype A; isotypes K,

L, SING). (Fig. 3E-F, Fig. 40-V)

Tree up to 40 m tall, dbh up to 60 cm, with a clear bole up to 30 m and steep buttresses

up to 5 m high and 1 m wide. Bark brown to reddish brown, somewhat scaly, sapwood

yellow to cream-yellow. Twigs dark brown to brownish grey, eventually lenticellate,

glabrous or finely but sparsely pubescent and soon glabrate; double-branching

occasional. Leaf with larger blades 6.5-10.9 cm long, 2.4-4. 8 cm wide, chartaceous

or occasionally subcoriaceous, slightly discolorous, elliptic, ovate, oblong-elliptic, or

elliptic-oblanceolate, base cuneate, sometimes obliquely or broadly so, apex narrowly

acuminate to caudate, the acumen 3.5-14 mm long, often deflexed to one side when

pressed, glabrous adaxially, glabrous or sparsely pubescent abaxially; midrib impressed

adaxially, raised abaxially, secondary veins 7-12 per side, diverging at 60-75° from

the midrib, weakly brochidodromous; secondary and higher-order veins slightly

raised adaxially, slightly raised to raised abaxially; petiole 4-9 mm long, shallowly

canaliculate, transversely wrinkled, glabrous or sparsely pubescent. Inflorescences

axillary or from the axils of fallen leaves, occasionally axillary on expanding axillary

shoots, 1-5-flowered, pale brown pubescent; peduncles 1-2 per axil, 1-2 mm long or

lacking; pedicels 1-3 per peduncle, 1.7-7 mm long, bracts 2, the lower at about the

midpoint of the pedicel and the upper subtending the sepals, upper bract larger than

lower bract, 1. 6-2.4 mm long, 2. 1-2.8 mm wide, clasping, semicircular; buds linear.

374

Gard. Bull. Singapore 67(2) 2015

apex obtuse or acute. Sepals V3-V2-com'iate, 2-2.9 mm long, 2-2.8 mm wide, coriaceous,

ovate-triangular, acute to apiculate, pale brown appressed-pubescent abaxially. Petals

yellow-green to greenish white in vivo; outer petals 13.5-34.2 mm long, 1.9-2. 5 mm

wide at base, 1-1 .6 mm wide at midpoint, linear, flat adaxially, longitudinally ridged

abaxially, densely puberulent on both surfaces except for glabrous base; inner petals

17.2-29 mm long, 1,8-1, 9 mm wide at base, 0.7-0.8 mm wide at midpoint, linear

to filiform, shallowly concave at base, acute at the apex, longitudinally ridged and

densely puberulent on both surfaces. Stamens 26-29 (includmg staminodes), 1.5-1. 7

mm long, clavate to narrowly oblong, apex of anther connectives c. 0.4 mm long,

oblong to conical, obtuse, densely papillate, filament 0.3 mm long; staminal cone c.

0.3 mm high, c. 1 . 1 mm in diam., a low ring surrounding the bases of the ovaries; outer

staminodes 7-8, narrowly oblong to clavate, flat, apex acute, glabrous. Caipels 3-4;

ovaries 1-1.2 mm long, nan'owly oblong, pubescent; stigmas somewhat connivent,

1 .3 mm long, barely exceeding tops of stamens, clavate, glabrous. Toms c. 1.3 nmi in

diameter. Pedicel of fruit 3-20 mm long, 4.5-10 mm thick at midpoint; torus of fruit

6-17 mm in diameter, 4-9 mm high. Monocarps greenish brown to brown, dehiscing

when mature to reveal a bright pink endocarp in vivo, up to 4 per fruit, 3. 1-6.4 cm

long, 2.5-3. 5 cm wide, 2-2.7 cm thick, oblong, ellipsoid, ovoid, or nearly globose,

apex rounded, base sessile or contracted into a stipe 3-5 mm long and c. 7 mm thick,

dark brown to black, blotched with lighter-coloured corky spots, glabrate or with a few

scattered hairs; pericarp 2-A mm thick. Seeds lying in two short partially overlapping

rows, transverse to long axis of monocaip, 8-12 per monocai*p, light green in vivo,

14-18 mm long, 10.3 mm wide, 8-8.1 mm thick, more or less ellipsoid, narrowly

oblong to cuneiform in cross section, smooth, dark grey with white crusty patches, flat

across micropylar end, chalazal end rounded; micropyle encircled by a rough irregular

aril plate.

Distribution. Occurs in Peninsular Malaysia, East Malaysia (Sarawak and Sabah), and

Indonesia (Riau Province of Sumatra and Central Kalimantan, East Kalimantan, and

West Kalimantan Provinces of Borneo).

Ecology. Mixed lowland dipterocarp forest and sometimes secondary forest, from sea

level up to 300 m.

Phenology. This species has been collected in flower in January, March-May,

and November; fruits have been collected Febmary-April, June-September, and

November-December.

Local names. Djangkang, djerendjang (East Kalimantan), medang (name also

commonly used for various Lauraceae), ngkurari (West Kalimantan: de Jong 439),

podjeng (East Kalimantan: Kostermans 10205).

Etymology. With pleasure we name this distinctive species in honour of Dr Francis

Ng, fornier Deputy Director General of the Forest Research Institute of Malaysia

Xylopia in Southeast Asia

375

and recipient of the 2009 David Fairchild Medal for Plant Exploration. Dr Ng has

contributed to knowledge of the flora of the region through a large body of work,

including the Tree Flora of Malaya and his study of the family Ebenaceae. Through his

studies of the seeds and seedlings of Southeast Asian forest trees (Ng, 1991; Ng & Mat

Asri Ngah Sanah, 1991) Dr Ng obtained valuable seedling data for the new species.

Additional specimens examined. MALAYSIA. Peninsular: Negeri Sembilan. PasohF.R., Plot

2 at the slope near the Percolation Pits and Lateral Flow, Pasoh, Kuala Pilah, 1 Mar 1977

[presumably had a fruit, as there is a seedling specimen attached], Mat Asri FRI 25722 (KEP);

Pasoh Forest Research Station, Ecology Plot No. 1, tree no. 1325, 6 Oct 1983 (st), Mat Salleh

& Normalawati KMS 40 (SING); Pasoh Forest Reserve, 50 ha Long-teim Ecological Research

Plot, 2A58N 102°18'E, 20 Feb 1988 (fr), LaFrankie 2758 (KEP); Pahang: Ulu Sg. Kepong,

Gua Peningat, 15 Jul 1970 (fl buds), Burgess FRI 19047 (KEP, L); Perak: “Upper Perak,”

May 1889 (fl), Wray 3562 (K, SING); Selangor: Bemam River State Land, 28 Jul 1920 (fr),

Yeoh 5037 ([Sinclair cites duplicate at K, but not found July 2014], KEP); Terengganu: Mandi

Angin Exped., S. Loh nr. Kuala Datok, 5 Jul 1968 (fl), Whitmore FRI 8965 (K, KEP, L);

Borneo: Sabah: Tawau District, Jalan Kuala Apas, 31 May 1961 (fr), Bakar SAN 1 7302 (L);

Sandakan District, Leila F.R., 18 Apr 1962 (fl), D.B. & J.S SAN 34713 (K, SING); Sipitang

District, Mesapol, 1 Jul 1962 (young fr), Mikil SAN 27180 (L [2 sheets]); Tawau District, ch.

5 Jalan from ML 15 1/2 Quain Hill Road, c. 100 ft, 15 Jun 1964 (ft), Pereira SAN 44208 (K,

SING); Keningau District, Sook-Tulid road mile 7%, 3000 ft, 27 May 1965 (fl), Sadau SAN

49574 (L [2 sheets], SING); Tawau, mi 1 0 1 /2 on road through Apas Forest Reserve ( 1 0 mi E of

Tawau), 300 ft, 7 Nov 1955, Wood SAN 17182 (A+ sep. carpol., L, SING); Sarawak: Without

definite locality, without date, Beccari 1578 (FI-W, K), Beccari 1579 (FI-W, K).

INDONESIA. Borneo: Central Kalimantan. Sintang HPH km 70 W of camp of main

(new) logging road, 0°51'54"S [given as 00°5133.6" S on one sheet] 112°1330"E [given as

112°13'29^9"E on one sheet], 120 m, 17 Apr 1994 (fl, ft), Mahyar et al. 990 (A, K, L, SAR,

SING); East Kalimantan: Wanariset research area, Rd Samboja-Semoi, km 2 Rintis B. Baru,

01°S, 117°E, 50 m, 3 Aug 1991 (ft), Ambri & Arifin W801 (A, BO, K, L); Bukit Soeharto

area off km 65 Balikpapan-Samarinda, 50 m, 20 Nov 1993 (fl), Ambri & Arifin A. A. 891 (A,

BO, K, L); Wanariset, Rintis Baru, Plot Matthijs, 01°S, 117°E, 50 m, 29 Jan 1992 (fl), Ambri

6 Arifin W1007 (A, K, L [2 sheets], MO); Sg. Wain region N of Balikpapan (fl), Kostermans

4488 (L, SING); Mentawir region, N of Balikpapan, 20 m, Sep 1950 (st, fr on L & SING

sheets), Kostermans 4512 (BO, K, L, SING); Loa Djanan, W of Samarinda, 30 m, 15 Apr

1952 (fl), Kostermans 6451 (A, K); island Nunukan (northern part), 100 m, 2 Dec 1953 (fl'),

Kostermans 8895 (A, BO, K, L, SING); Balikpapan distr., nr village Mentawirlow, 3 Mar 1955

(st), Kostermans 10154 (L, NY, SENG); East Kutei, Loa Djanan region along road Balikpapan.

Samarinda, km 25, 15 Mar 1955 (fl), Kostetmans 10205 (L); E Kutei, Sg Tiram, 40 m, 15 Apr

1952 (fl), Kostermans bb.35023 (A, K); East Kutai Reserve, vicinity of Sengata and Mentoko

Rivers, altitude below 300 m, 0°30N, 117°20'E, 15 Apr 1978, Leighton 154 (L); Z.O. Afd.

V. Borneo, Boeloengan, Noekoekaii (Boschtuin), 18 May 1939 (st), Neth. Ind. For. Service

bb. 29395 (A, L); West Kalimantan: Kabupaten Sanggau, 10 Aug 1993 (fr), de Jong 439 (L,

NY); Melawi, Tjatit, B. Gontuk, c. 180 m, 3 Mai' 1939 (fl), Netherlands Indies Forest Service

bb.27013 (A, BISH, K, L, NY, SING); Melawi, B. Ulu, Bomsepan, 275 m, 23 Jul 1939 (young

fr), Neth. Indies For Service bb. 29037 (L, SING). Sumatra: Riau: Indragiri, the uplands,

Danau Mengkuang, 21 Apr 1939 (young fr), Buwalda 6633 (A, K, L, SING); Riouw en Ond.

Indrag. Bovenlanden, Danau Mengkuang, 21 Apr 1939 (young fr), Netherlands Indies Forest

Service bb.27571 (A, K, L, NY).

376

Gard. Bull. Singapore 67(2) 2015

Notes. Material of this species is usually identified in herbarium collections as Xylopia

malayana, but X. ngii differs consistently from that species in its smaller (2-2.9 mm

long) sepals sparsely covered with pale brown pubescence and the longer (13.5-34.2

mm) and narrower (1-1.6 mm wide at the midpoint) outer petals. In Xylopia malayana

the sepals are 2.7^ mm long and densely covered with rusty pubescence, and the

outer petals are 13.5-20 mm long and 1.5-2 mm wide at the midpoint. The leaves of

Xylopia ngii have a characteristic narrow acumen, which is often bent to the side in

pressed specimens, rather than the gradually acuminate tip seen in the leaves of X.

malayana, which tends to press flat; the bending of the leaf apex inX ngii is the result

of the fact that its leaf midrib is usually curved downward towards the apex. The most

striking difference between the two species, however, is in the finits and seeds: the

monocarps oi Xylopia ngii are 3. 1-6.4 cm long, 2. 5-3. 5 cm wide, sessile or broadly

short-stipitate with a stipe 3-5 mm long and c. 7 mm thick and a pericarp 2-A mm

thick, each monocarp containing 8-12 seeds 14-18 mm long. l\\ Xylopia malayana

the monocarps are 1 .9-3.9 cm long, 0.9-1 .7 cm wide, distinctly stipitate with a narrow

stipe 2.5-7 mm long, and with a pericarp 0.3-0. 5 mm thick, each monocarp containing

up to 3 seeds 7.5-8 mm long. In fruit Xylopia ngii can only be confused with X.

dehiscens (Blanco) Men’., a species of the Philippines and northern Borneo, which has

a blunt leaf apex and a fmit drying black and without the corky blotches oiX. ngii.

Part of the difficulty in separating Xylopia ngii from X. malayana and other

congeners has been the fact that the flowers and fruits of the species are rarely collected

together. Thus the type collection and the specimen Mahyar et al. 990 have played a

critical role in our delimitation of the species. While the taxonomic confusion has

been principally with Xylopia malayana, specimens of this species have also been

confused with X. elliptica and even with X. fusca Maingay ex Hook.f. & Thomson:

the specimens Wray 3562 and Yeob 5037 were identified as X. elliptica by Sinclair

(1955), the specimens Amhriansyah & Arifin AA 891, Ambriansyah & Arifin W 801,

and Ambriansyah & Arifin W 1007 were identified as X. malayana in Sidiyasa et al.

(1999). Turner (2014) determined the collections Mikil SAN 27180 and Pereira SAN

44208 as Xylopia malayana, and Sadau SAN 49574 as X. fusca. Xylopia ngii was

illustrated in LaFrankie (2010: 100, reproduced in Fig. 3F here), where it was called

Xylopia fusca.

The descriptions of seedlings of Xylopia malayana" in Ng (1991) and Ng &

Mat Asri Ngah Sanah (1991) are based on this species. A seedling specimen is attached

to the KEP sheet of the collection Mat Asri FRI 25722, the apparent original seed

source for the seedling studies.

In the Pasoh Forest Reserve an individual of Xylopia ngii, tree #414079, was c.

40 m tall with a dbh of 43.9 cm, with steep buttresses at its base (Fig. 3E). The tree was

growing on some of the highest ground in the plot (Manokaran et al., 1992).

Xylopia heterotricha D.M.Jolmson & N.A.Murray, sp. nov.

Species differing from its congeners by the twig pubescence consisting of a mixture of

acicular hyaline spreading hairs 0.5-0. 7 mm long and curling appressed yellow-brown

Xylopia in Southeast Asia

377

Fig. 3. A-D: Xylopia platycarpa D.M.Johnson & N.A.Murray. A. Two open flowers and a

single bud. B. Open flowers and a single dehisced monocarp. C. Monocarps of two fruits, the

uppermost ones in the photograph just beginning to dehisce. D. Single open flower, showing

curled ends of petals. Xylopia ngii D.M.Johnson & N.A.Murray. E. Base of tree #414079,

Pasoh Forest Reserve, showing the prominent buttresses and absence of stilt roots. F. Dehisced

monocarps and leafy branches. (Photos: A-D: S. Gardner; E: N.A. Murray; F: J. LaFrankie)

378

Gard. Bull. Singapore 67(2) 2015

B-D, H-J, 0-R

5 mm

Fig. 4. A-G. Xylopia platycarpa D.M.Johnson & N.A.Murray. A. Habit. B. Inflorescence.

C. Outer petal, adaxial view. D. Inner petal, adaxial view. E. Profile view of twig indument.

F. Monocarp, lateral view. G. Seed, lateral view. H-N. Xylopia heterotricha D.M.Johnson &

N.A.Murray. H. Flower bud, lateral view. I. Base of outer petal, adaxial view. J. Inner petal,

adaxial view. K. Profile view of twig indument. L. Fmit. M. Seed, lateral view. N. Habit. O-V.

Xylopia ngii D.M.Jolinson & N.A.Murray. O, P. Flower buds, lateral view. Q. Outer petal,

oblique adaxial view. R. Inner petal, adaxial view (twisted toward apex). S. Monocarp, lateral

view. T. Profile view of twig indument. U. Habit. V. Seed, lateral view. Drawn by D.M. Johnson

from (A, C, D, F, G) Gardner STI882, L; (B, E) Cwyis 2482, K; (H) Piiasa 4550, KEP; (I,

J, N) Puasa 4550, A; (K) McDonald & Ismael 3526, A; (L, M) Ambri & Arifin W602, A; (O,

S, T) Wood SANA 3454, A; (P) Ambri & Arifin WI007, MO; (Q, R) Ambri & Arifin ±4891,

A; (U) Mahyar et al. 990, SING; (V) spirit collection gathered beneath flee #414079 in Pasoh

Forest Reserve.

Xylopia in Southeast Asia

379

hairs 0. 1-0.2 mm long, the latter often persisting on leafless twigs. It differs from

Xylopia elliptica in addition in having the leaf blades persistently pubescent abaxially,

the petals 28-47 mm long (versus 5.5-10 mm long) and the monocarps up to 17,

stipitate (stipes 7-10 mm long) and distinctly pubescent at maturity (versus monocarps

1-2, short-stipitate (stipes 1.5-3. 5 mm long) and glabrate). - TYPE: Malaysia, Sabah,

Sandakan, Bettotan, 27 March 1 935 (fl), Puasa 4550 (holotype KEP; isotypes A, K, L,

SING, US). (Fig. 4H-N)

Tree up to 36 m tall, dbh up to 48 cm, with a clear bole up to 26 m, occasionally with

buttresses up to 1.5 m high. Bark light grey, rarely white, yellowish green, or dark

brown, smooth. Twigs dark brown, densely pubescent, the pubescence consisting of

a mixture of acicular hyaline spreading hah's 0.5-0. 7 mm long and curling appressed

yellow-brown hairs 0. 1-0.2 mm long, the latter often persisting on leafless twigs;

double-branching occasional. Leaf with larger blades 7.9-10.2 cm long, 2.4-3. 7 cm

wide, chartaceous, often mottled adaxially when dried, nan'owly elliptic to oblong,

base cuneate, apex acuminate, the acumen 8.5-18 mm long, punctate and glabrous

adaxially except for the densely hispid-pubescent midrib, pubescent with loosely

appressed hairs abaxially; midrib plane adaxially, raised abaxially; secondary veins

9-13 per side, diverging at 45-60° from the midrib, weakly brochidodromous,

secondary and higher-order veins slightly raised adaxially, slightly raised to raised

abaxially; petiole 2.5-3 mm long, deeply canaliculate, pubescent. Inflorescences

axillary or from the axils of fallen leaves, 1 -flowered, brownish to yellowish pubescent;

peduncles absent; pedicels 4. 5-5. 5 mm long, bracts 2, the lower bract attached at

or below the midpoint and the upper bract attached to distal half of pedicel, upper

bract 3 mm long, ovate, acute; buds linear, apex acute. Sepals Yi-connate, 3^ mm

long, coriaceous, broadly ovate, apex acute, acuminate, or caudate, brown-pubescent

abaxially. Petals white to yellow in vivo\ outer petals 28-47 mm long, c. 2.5 mm

wide at base, 1-1.2 mm wide at midpoint, linear, flat adaxially, longitudinally ridged

abaxially, apex acute, densely puberulent on both surfaces except for glabrous base;

inner petals 30-36 mm long, 1.6-1. 7 mm wide at base, c. 0.8 mm wide at midpoint,

filiform, apex acute, longitudinally ridged and densely puberulent on both surfaces.

Stamens 65-74, 1.8-1. 9 mm long, clavate to narrowly oblong, apex of anther

connectives 0.4-0. 5 mm long, bluntly conieal, densely papillate, filament 0. 3-0.4 nmi

long; staminal cone barely visible, a low ring surrounding the bases of the ovaries;

staminodes 16-19, c. 2 mm long, narrowly elliptic, narrowly oblong or oblanceolate,

glabrous. Carpels 17-18, ovaries c. 0.8 mm long, oblong, densely pubescent; stigmas

more or less connivent, c. 1.2 imn long, linear, dark, glabrous or occasionally with a

tuft of hairs at the apex. Torus flat or a little concave beneath the carpels, c. 3.2 mm

in diameter. Pedicel of fruit 3-14 mm long, 4.5-10 mm thick at midpoint; torus of

fruit 4-9 mm high, 6-17 mm in diameter. Monocarps light green to glaucous green,

dehiscing when mature to reveal a bright red endocaip in vivo, up to 17 per fruit, 2.8-

3.6 cm long, 1.1-1 .4 cm wide, 1.2-1 .3 cm thick, oblong to ellipsoid, roughly circular

in cross section, apex rounded, base contracted into a stipe 7-10 mm long and 2, 8-3. 5

mm thick, irregularly sunken but raised into a longitudinal ridge along the abaxial

380

Gard. Bull. Singapore 67(2) 2015

midline, pubescent; pericaip 0.3-0. 5 mm thick. Seeds an'anged in two in*egular rows,

perpendicular to long axis of monocarp, 5-6 per monocarp, grey- white to purple

in vivo, 7. 1-7.6 mm long, 5. 3-5. 9 mm wide, 4. 6^. 9 mm thick, broadly ellipsoid,

elliptic in cross section, smooth, obliquely flattened at micropylar end, rounded at

chalazal end, sarcotesta evident after soaking as a thin pale yellow layer over dark

brown seed coat layers; aril plate surrounding micropyle 3. 2-3. 3 mm long, 2.7-3 mm

wide, a flat ring, white.

Distribution. Occurs in East Malaysia (Sarawak and Sabah), and Indonesia (Bangka

Belitung, Bengkulu and South Sumatra Provmces of Sumatra and neighbouring

islands, and Central Kalimantan and East Kalimantan Provinces on Borneo).

Ecology. Lowland primary and mixed dipterocarp forest, often on hilltops or ridges,

occasionally in riparian or logged over and secondaiy forest, in one case in forest

consisting of “pole-sized dipterocarps.” It has been collected on granitic sand,

sandstone, and red clay at 50-500 m. Associates at one site were species of Aporosa

Blume, Dipterocarpus C.F.Gaertn., Eugenia L., Hopea Roxb. and Shorea Roxb. ex

C.F.Gaertn.

Phenology. Flowers collected in March-May and July-October; fruits collected in

January, April-July and September-October.

Local names. Banetan koening (Sumatra: Endert 176 E.l[8 or P]. 886), banetan poetih

(Sumatra: Boschproefstation nr. For T 971), banitan (Billiton: van Possum 46), banitan

gadan g {Smn?iXvdi: Dumas 1527), banitan laki (Bangka: Andong 111), bebanditan (East

Kalimantan: Ambri & Arifin AA7), karai (Sabah: Maidin 3056), karai batu (Sabah:

Puasa 4550), miwi (East Kalimantan: Kostermans 6634), pisang-pisang (Sabah: Ong

SAN 120806, Wood SAN A 4819; NOTE: this is a widely used common name for

Annonaceae generally), sebukau (Sarawak: Jacobs 5409), usaj (Bangka: Kostermans

& Anta 223).

Etymology^. Named for the twig pubescence, which consists of a mixture of two

different types of trichomes: longer erect straight trichomes and shorter curled and

more appressed trichomes.

Additional specimens examined. MALAYSIA: Borneo: Sabah: Kuala Penyu (Beaufort),

Kepayan, 30 May 1968 (fr), Binideh 55770 (K, KEP, L, SAR); Sabah, Nabawan District, Sg.

Pingas-Maitland area, 23 May 1986 (fl), Fidilis Krispinus SAN 115831 (K, KEP); Sandakan

87.5 Hap Seng logging, 24 Jun 1976 (ft), Leopold & Taha SAN 8355 1 (K, L [2 sheets]); Beluran

District, Bidu-Bidu F.R., 2 1 Jul 1990 (ft*), Madani SAN 128873 (K, KEP); Marotai, 8 Apr 1933

(fl), Maidin [B.N.B. 3056] (A, K); Lahad Datu dist., Diwats, Kennedy Bay Co. area, 60’ alt, 9

Jun 1961 (fr). Main Chai SAN 25078 (K, L); Tawau District, Sepulut Reseai'ch Plot, 13 Aug

1987 (ft), Ong SAN 120806 {K, KEP, L); Tiulon, Nabawan, 600 ft, 16 Sep 1976 (ft), Tarmiji &

Dew'ol SAN 84191 (K, L, SAR, SING); mi. 61 Telupid Road, 20 m alt, 4 .lul 1980 (fr), Tenniji

Xylopia in Southeast Asia

381

Arshid SAN 92570 (K, KEP, L, SAR); Lahad Datu District, Cpt. 6, North Borneo Timber Co.,

Concession Area, Kretam, 22 May 1954 (fr), Wood SANA 4819 (A, L, SING); Sandakan Cpt.

16, Sepilok Forest Reserve 15 mi W of Sandakan, 30 Apr 1955 (fl), Wood & Charington 16318

(A, KEP, L, SING); Sarawak: 3rd Division, Kapit District, Belaga subdistrict, left bank of

Rajang River c. 10 km below Belaga, Segaham Range near Belaga airfield, 2°40'N 1 13°50'E,

<500 m, 3 Sep 1958 (ft*), Jacobs 5409 (K, L, SAR, US), 7 Sep 1958 (fl), Jacobs 5453 (K, L,

SAR, SING, US); Ulu Sg. Semawat Belaga, 7th Division, 23 Oct 1981 (young fr), Othman et

al. S.43397{ASU,L, SAR).

INDONESIA: Sumatra: Bangka Belitung: Bangka, Lobok Besar, 5 m, 14 Sep 1949 (st),

Andong 111 (L [2 sheets]); Ond. Luid. Bangka, 20 m, 16 Apr 1927 (st), Boschproefstation

bb.ll299 [Mohamad Oetei 149] (A); Lobok-besar, SE Bangka, 30 Aug 1949 (fl), Kostermans

& Anta 223 (A, K, L, NY, SING); Lobok-besar, G. Lading, 100 m, 30 Sep 1 949 (fl), Kostermans

&Anta 1012 (A, K, L, SING); Billiton, s.d. (f\), van Rossum 46 (L [2 sheets]); Bengkulu: Lais,

Talang Benal, 250 m, 20 Mar 1925 (st), Idris 5/bb.8858 (A); South Sumatra: Res. Palembang

ond. Afd. Lemakang Ilir, 75 m, 8 Sep 1924 (fl), Bo.^chproefstation nr. F or T 971 (L);

Palembang, Rawas, 27 Apr 1917 (st), Dumas 1527 (L); Res. Palembang, ond. Afd. Banjoeasin

en Koeboestreken, 25 m, 6 Feb 1 920 (fl), Endert 1 76 E. 1[8 or P],886 (L [2 sheets]); Banjoeasin

en Koeboestreken, 16 Nov 1915 (fr), Grashqff 828 p.p. (L); Borneo: Central Kalimantan:

Samba, 1994-1995 cutting blocks of PT Handiyani, 0°43T6.7"S 112°50'34.2"E, 340 m, 24

Jan 1995 (ft), Jar\ne & Ruskandi 5229 (A, KEP, L); P. B. U. base camp and environs. Trail

Jalang Nancy, 16 Jun 1990 (fl, fr), Ridsdale PBU55 (L [2 sheets]); East Kalimantan: Sepaku

PT ITCI, Kenangan, Balikpapan, 30 Oct 1990 (fl), Ambri & Arifin AA7 (A, K, L); Wanariset,

Wanariset Research I area, off km 35, Wanariset-semoi Road, 70 m, 16 Jan 1 991 (fr), Ambri &

Arifin W602 (A, K, L); Bulungan, Nunukan, Boschtuin, 8 May 1939 (fl), Netherlands Indies

Forest Seixice bb. 29395 (A, L, SING); Loa Djanan, W of Samarinda, 30 m, 27 Apr 1952 (fir),

Kostermans 6634 (A, BO, K, L, SING); Boeloengan, Kabiran, G. Shnendoeroel, 5 Aug 1927

(st), Zwaan bb. 11722 (A); Pujungan Dist., Kayan-Mentirang Nature Reserve on Bahau River at

or to 3 km above confluence with Gong Biou River towards Long Alango, 02°50'N 115°50'E,

6 Jul 1992 (ft), McDonald & Ismail 3526 (A, F, L, SING); West Kutei, 20 Mar 1932 (st),

Netherlands Indies Forest Service bb. 16521 (A, L); Kutei, 15 Jul 1957, Schut K.22 (BO, K, L,

SING); Sungai Wain Protected Forest of km 15 Balikpapan-Samarinda, 50 m, 11 Jan 1992 (fr),

Sidiyasa et al. 818 (A, K, L).

Notes. Xylopia heterotricha is distinctive because of the unusual character of the twig

pubescence, which consists of a mixture of long light-coloured acicular hairs and

short brown curled hairs. The leaf blades are relatively small and oblong, often with

a blotchy mottled appearance to the adaxial leaf blade surface when dried. The petals

are very long, approaching in length those of Xylopia magna Maingay ex Hook.f. &

Thomson from Peninsular Malaysia, Singapore and Borneo. The number of carpels

and monocarps is exceptional for the Xylopia malayana group, reaching up to 18:

the more typical number for species in this group is fewer than 7, rarely up to ten

(Keng & Heaslett, 1973); The monocarps are superficially similar to those of Xylopia

malayana in being persistently pubescent and distinctly stipitate, but are usually

wrinkled or puckered when dry and never obliquely striate as in the monocarps of

Xylopia malayana.

Previous reports of Xylopia elliptica from Borneo (e.g., Sinclair, in lift.; Turner,

2011, 2014) all refer to X. heterotricha.

382

Gard. Bull. Singapore 67(2) 2015

Xylopia sessiliflora (Kochummen & Whitmore) D.M.Johnson & N.A.MiiiTay, comb,

nov. - Xylopia fusca var. sessiliflora Kochummen & Whitmore, Fed. Mus. J. 13:

135 (1970 [‘1968’]), - TYPE: Malaysia, Negeri Sembilan, Kuala Pilah, Pasoh F.R.,

Compt. No. 14, 19 February 1927, Mohd. Yatim bin Aseh, Forester, for Serting KEP

62984 (holotype KEP).

Distribution. Peninsular Malaysia. Known from only five collections, each from a

different state in Peninsular Malaysia. Last collected in 1972.

Additional specimens examined. MALAYSIA: Peninsular: Johor: E Johore, Sungai Kaya, 15

Mar 1937 (fl), Kiah SFN 32403 (A, BKF, K, KEP, SING); Melaka: Without definite locality,

without date (fl), Herb. Cantley 5842 (SING); Pahang: Tasek Bera FR, 22 Jul 1972 (fl), Chan

FRI 16942 (A, K, KEP, L, SAR, SING); Perak: Without definite locality, without date (bud),

Wray 3194 (CAL (Sinclair, 1955), SING [2 sheets]).

Notes. The combination of dense appressed pubescence of the abaxial leaf surfaces

and twigs and the very short flower pedicels readily distinguishes Xylopia sessiliflora

from all other Peninsular Malaysian Xylopia species. Xylopia sessiliflora differs from

X. fusca of the Xylopia ferniginea group in a number of significant ways. The leaf

indument is silvery grey rather than golden, the leaf blades are lanceolate, ovate, or

elliptic and usually acuminate at the apex, rather than oblong and obtuse, rounded, or

rarely short-acuminate at the apex, the pedicel is only 2-3.4 mm long as opposed to

5-10 mm long, the calyx is only %-connate rather than ’A-Yi-connate, the petals are

up to 44 mm long rather than only up to 26 nmi long, and there are 8-10 rather than

3-5 carpels. The paratype Kiah SFN 32403 had been identified by Sinclair (1955) as

Xylopia malayana. To confuse things further, this specimen was identified in turn as

hot\\ Xylopia elliptica and mcdayana in the same publication by Comer (1978).

The fmit of this species is not yet known, and may provide additional characters

for distinguishing it from other Xylopia species in the Pasoh Reserve and elsewhere.

According to Kochummen (1997), trees of this species in the Pasoh Reserve have stilt

roots, the only similarity with Xylopia fusca and other members of the X. ferruginea

group.

Four specimens were cited in the protologue of Xylopia fusca var. sessiliflora.

In addition to the type and Kiah SFN 32403 already discussed, these two were cited:

Johor: Labis F.R., KEP 105267 and Kedah: Gunong Inas F.R., KEP 104789. The latter

two specimens are sterile; the Kedah specimen definitely does not look like other

material of this species, and the Johor specimen is not deteraiinable.

Conclusion

A clearer understanding of regional patterns of distribution of the 23 Xylopia species

occurring in the Sundaic region of Southeast Asia is now possible. Six species are

widespread across the Malay Peninsula, Sumatra and Borneo: Xylopia caudata Wall.

Xylopia in Southeast Asia

383

ex Hook. f. & Thomson, X ferruginea, X. fusca.X. malayana,X. ngii andX. sumatrana.

Xylopia does not occur on Java. Two species are shared between the Malay Peninsula

and Sumatra: X. dicarpa Hook.f. & Thomson and X. oxyantha (Wall, ex Hook.f. &

Thomson) Hook.f. & Thomson. Two species are shared between the Malay Peninsula

and Borneo: Xylopia eiythrodactyla and X. magna. Xylopia magna was previously

known only from the Malay Peninsula; the following specimen represents the first

record of that species for Borneo: Malaysia, Sarawak, logging area, Bukit Tubeh,

Tebedu, Serian, 23 Jun 1996 (fr), Jamree et al. S. 73290 (KEP). Xylopia heterotricha is

shared between Sumatra and Borneo.

Xylopia elliptica, X. platycarpa, X. sessiliflora and X. subdehiscens (King)

J. Sind, are known only from the Malay Peninsula, and Xylopia conifolia RidL, X.

kuchingensis TM.Tumer & D.M. Johnson, X. mucronata Boerl., X. orestera T.M.Tumer

& D.M.Johnson and X. pulchella Ridl. are unique to Borneo. There is no species

cun*ently known to be endemic to Sumatra.

Sundaic species with distributions extending outside of the region include

Xylopia densifolia Elmer and X. dehiscens (Blanco) Merr., which occur in Borneo

and the Philippines. Xylopia pierrei, occurring in Thailand, Cambodia, and

Vietnam, just reaches the Malay Peninsula. Xylopia vielana is the only species in

Thailand that does not occur in the Sundaic region, its distribution extending instead

from northeastern Thailand into Vietnam and southern China. With the notable

exception of the Malay Peninsula endemics mentioned above, all of which belong

to the Xylopia malayana group, the X. ferruginea gi'oup and the X. malayana group

exhibit strongly similar biogeographic patterns in the region.

Some of the rarer species of Xylopia in the area may be simply under-collected.

Several species are large trees more than 40 meters in height, and this, coupled with

sparseness of individuals in populations, may cause them to be overlooked. Some

lowland evergreen rainforest xylopias may, however, exist only in small fragmented

relict populations of the Sundaland lowland evergreen forests that contracted into

refugia following the last glacial maximum (Cannon et al., 2009). This explanation

accords well with the extraordinary diversity of the genus found at sites such as the

Pasoh Reserve, where in 2014 we verified the occurrence of seven Xylopia species in

or near the Pasoh pennanent 50-hectare research plot (Manokaran et al., 1992). While

the Pasoh plot is known as an area of high diversity, Xylopia species within such

a small area is unequalled in our field experience with the genus.

The intensive inventories of lowland rainforest plots in southeastern Asia (see

Losos & Leigh, 2004) have documented that Annonaceae are a vital component of

these forests and that Xylopia species are invariably a substantial component of that

Annonaceae diversity (Manokaran et al., 1992; Lee et al., 2002). Data from these

research plots can begin to give us a measure of population structure for many species,

including those of Xylopia, but only if species-level identification can be reliably

established. It is hoped that this contribution toward clarification of Xylopia taxonomy

in the region will enable stronger cross-comparisons among these long-term plots.

Our results make clear, however, that further collecting and taxonomic analysis of the

Malay Peninsula flora is still critically needed.

384

Gard. Bull. Singapore 67(2) 2015

ACKNOWLEDGEMENTS. We thank the Singapore Botanic Gardens and the Forest Research

Institute Malaysia (FRTM) for their vital support of our project. The Singapore Botanic Gardens

provided critical financial support in the form of SBG Research Fellowships, and we were

generously supported at FRIM as FRIM Fellows, funded by the Ministry of Science, Technology

and Innovation, Malaysia (MOSTI) under the “Flora of Peninsular Malaysia project” (01-04-

01-000 Khas) and “Documentation & Inventory of Flora of Malaysia project,” based at Forest

Research Institute Malaysia. In Singapore we received kind assistance from Edmund Chia,

Bazilah Ibrahim, Nura Bte Abdul Karim, Joseph Lai, Samantha Lai, Serena Lee, Michael

Leong, Paul Leong, Jana Leong-Skomickova, Low Yee Wen, Shawn Lum, David Middleton,

Michele Rodda, Benito Tan, Nigel Taylor, Elango Velautham, Craig Williams, and Winnie

Wong. We express our appreciation for assistance in Malaysia from Saw Leng Guan, Richard

C.K. Chung, Mohamed A. Latiff, Ruth Kiew, Ummul Nazrah Abdul Rahman, Yao Tze Leong,

Rafidah Abdul Rahman, Avelinah Julius, Norsham Suliaina Yaakob, Ming Yee Chew and Ong

Poh Tek. Sabbatical leave support from Ohio Wesleyan University is gratefully acknowledged,

as is financial support from the TEW Presidential Discretionary fund at Ohio Wesleyan, which

supported our further herbarium work at L and K. Additional funding to support herbarium

work at L was provided by Naturalis Biodiversity Centre in Leiden kindly amanged by Erik

Smets. At Kew we were hosted by David Goyder and the Drylands Africa team and received

assistance from Ian Turner. For providing key photographs of two of the new Xylopia species

to help illustrate this paper we thank Simon Gardner and James LaFrankie. Kate Ball provided

the excellent pen and ink rendering of Xylopia eiythrodactyla. For assistance with preparation

of the figures for publication we thank Sara Stuntz, Doug Thompson and Emily Gattozzi. We

thank the two anonymous reviewers for suggesting improvements to the manuscript. Finally,

we thank curators of the following herbaria for making specimens available to us either by loan

or during our visits: A, AAU, ASU, BISH, BKF, BM, BO, K, KEP, KLU, L, M, MICH, MO,

NY, SAR, SING, U, UKMB, US.

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387

Studies in Southeast Asian Melastoma (Melastomataceae), 1.

Morphological variation in Melastoma malabathricum

and notes on rheophytic taxa and interspecific

hybridisation in the genus

K.M. Wong

Herbarium, Singapore Botanic Gardens, National Parks Board,

1 Cluny Road, Singapore 259569

wkin2000@gmail .com

ABSTRACT. Morphological variation in the widespread Indo-Pacific Melastoma

malabathricum L. is discussed, as well as the possibility of hybridisation or introgression

with other species. Hypanthium indumentum is a reliable character to diagnose the group and

species. Five rheophytic taxa, including four distinctive species and a variant of the widespread

M. malabathricum, which are superficially alike because of their stenophyllous leaf form,

are enumerated for Soutlieast Asia and a key is provided for their identification. Melastoma

kahayanense K.M. Wong from Borneo is described for the first time and the SumaXmn Melastoma

stenophyllum Merr. is considered a distinct species and not synonymous to M. malabathricum.

Various putative hybrids or introgression products involving M. malabathricum are identified.

Keywords. Brunei, identification key, indumentum, introgression, Kalimantan, Malesia,

morphology, Sarawak, Sumatra, taxonomy, variation

Introduction

The most recent revision of Melastoma L. by Meyer (2001) included 22 species,

of which an overwhelming 18 were considered to have ranges within or including

Malesia. Of these, eight species are listed for the Malay Peninsula and nine species

for Borneo. Nevertheless this appears to be very incomplete because collections that

were apparently not consulted include new taxa characterised by distinctive forms

of hypanthium mdumentum (see Wong & Low, 2015). In addition Meyer adopted

wide species concepts for a number of taxa without much clarification and which

sometimes appeared to be inconsistent. This series presents various discussions and

taxonomic changes, including the description of new or overlooked taxa. We begin

with a commentary on the fundamental significance of the hypanthium indumentum

type and variation within Melastoma malabathricum L., the type species. Based

on this and other characters, distinct Southeast Asian rheophytic species are then

distinguished from a rheophytic form of Melastoma malabathricum. Finally, the

evidence for interspecific hybridisation in the genus is reviewed and putative hybrids

between Melastoma malabathricum and other species in Malesia are identified.

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Hypanthium indumentum type a reUable distinguishing character

Species with distinctive types of hypanthium indumentum consistently differ in other

morphological characters, ecology or distribution (Naudin, 1 850; Cogniaux, 1891; Li,

1944). The main hypanthium indumentum types documented for Melastoma include

bristles (filiform or terete, hair-like indumentum, as in M. moUe Wall, ex Ridl. or

M. sanguineum) (Fig. lA, B) or scales (visibly flat even under xlO magnification,

usually triangular to ovate or lanceolate (= narrowly ovate) indumentum, as in M.

imbricatum Wall, ex C.B.Clarke orM. malabathricum) (Fig. IC, D-J), or sometimes

penicillate emergences (outgrowths of hypanthium tissue that bear bristles on their

edges or at their tips, as in the trans-oceanic vicariants M. beccariamim Cogn. andM.

saigonense (Kuntze) Merr.; Cogniaux, 1891; Flansen, 1977; Meyer, 2001) (Fig. IP,

Q). Thus far, the scales documented are mainly triangular to ovate or lanceolate flat

scales with irregularly and shallowly serrate margins (e.g., Melastoma malabathricum,

M. normale, M. stenophyllum MeiT.) (Fig. ID-L), or subulate-subentire (e.g., M.

ariffinii K.M.Wong) (Fig. IN), or deeply divided nearly to base forming several linear

segments (e.g., M. ashtonii K.M.Wong) (Fig. 10) (Wong & Low, 2015).

Variation in Melastoma malabathricum

“On account of the great variation among the plants which must be identified as M.

malabathricum in the wide sense, no satisfactory classification of them has been

proposed.” - E.J.H. Comer, Gardens’ Bulletin Straits Settlements 10: 300 (1939).

Melastoma malabathricum was lectotypified by Bremer in Jarvis et al. (1993: 65).

The lectotype is Herb. Hermann, Vol. 1: 55, No. 171, Ceylon in the Natural History

Museum, London (BM). There are two elements numbered “171”. One is a leafy twig

without flowers (barcode BM000621418) and is found on the same sheet together

with another separate leafy twig bearing an open flower (barcode BM000621419).

Both these elements are ostensibly conspecific. They have twigs covered in appressed

ovate, triangular and broad-lanceolate scales, the margins of which have short irregular

serrations. The flower has a hypanthium clothed and completely hidden in overlapping,

flat, namowly tiiangular to lanceolate scales (length typically three times or more the

width), also with irregular and shallow marginal serrations. All these character-states

are also found together in most of the material attributable to this species from India

through Malesia (Fig. ID-J), into northern Australia and the Pacific region, where

indeed leaf size and shape, number of flowers in an inflorescence, and flower size

appear to vary continuously without any significant morphological gap that could be

used to justify taxonomic division. Hypanthium scale dimensions are less variable

and, although the hypanthium scales of material from Sri Lanka (Ceylon) (Fig. ID,

2A) are merely half the width of the largest hypanthium scales found in Malesian

material (Fig. lE-I, 2C), there are intenuediate sizes between these extremes, and the

smaller scale sizes occur also in Malesia (Fig. IE, F, G, J).

Variation in Melastoma malabathricum

389

Fig. 1. Hypanthium indumentum types found in Melastoma species. Bristles in A. Melastoma

molle Wall, ex Ridl., and B. M. sanguinetim Sims. Scales with serrate margins in C. M,

imbncatum Wall, ex C.B. Clarke, D. M malabathricum L. from Ceylon, E. Malay Peninsula-

Selangor, F. Malay Peninsula-Pahang, G. Sumatra, H. .lava, I. Bomeo-Sarawak, and J. Borneo-

West Kalimantan, K. M. nonnale D.Don, L. M. stenophyllum Merr. and M. M. kahayanense

K.M.Wong. Subulate-subentire scales in N. M. ariffinii K.M.Wong. Scales that are deeply

divided into linear segments in O. M. ashtonii. Penicillate emergences in P. M. saigonense

(Kuntze) Merr. and Q. M. beccarianum Coqn. Drawn by K.M. Wong from (A) Symington KEP

21361\ (B, left) Webb et al. WA 64, (B, middle and right) Sidek SK 409; (C, three on left) Shah

& Sidek MS 1123, (C, right) Everett FRI 1 3992; (D) Gardner 296; (E) Hume 7313; (F) Burkill

& HaniffSFN 16660; (G) Jacobs 8193; (H) Zollinger 1938; (I) Othman S 21107; (J) Church

et al. 557; (K) Ribu (Dr Prain’s collector) s.n. 5 Apr 1902; (L) Rahmat Si Boeea 7000; (M)

Mahyar 890; (N) Chai & Ilias S 31110; (O) Ashton BRUN 5629; (P) Webb et al. WA 81; (Q)

Wong WKM 189; all from SING. 2 mm scale shown applies throughout; measurements were

made with a Micro-Scale with 0. 1 mm divisions from Minitool Inc.

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Fig. 2. Hypanthium scales and upper leaf surfaee scabrid-hairy indumentum. A-D. Melastoma

malabathricum L. A, B. Typical in material from Ceylon {Gardner 296 (= Hooker 8246),

SfNG); C, D. Malay Peninsula {Burkill & Shah HMB 935, SING). E, F. Melastoma normale

D.Don from Sikkim {Hooker 9577, SfNG). (Photomicrographs courtesy of Y.W. Low)

This is, therefore, a highly variable speeies and, in the present assessment, no

eonsistent varieties or forms seem possible to delimit using clear-cut and discontinuous

characters. It is possible that, although apparently extreme environments may be

expected to pose selection pressure that skews some morphological expression,

this has not happened because the different populations still participate in sufficient

Variation in Melastoma malabathricum

391

genetic exchange. It is useful to identify some of this variation, bearing in mind

the characteristics highlighted in the previous paragraph are consistently present

throughout.

(i) Leaves are typically elliptic, with 5 longitudinal veins (on the upper surface distinct

as 5 longitudinal furrows, the outermost two very near to the leaf margins). In the

Malay Peninsula and Borneo there are rheophytic fonns with leaves narrowly

elliptic to almost linear, with 3 longitudinal veins (on the upper surface distinct as

3 longitudinal fnrrows), and sometimes 3-5 longitudinal veins on some or many

leaves.

(ii) Leaf blades are typically (1 .6-)2.5-4.5(-8) cm wide. In the rheophytic forms leaf

blades are (0.3-)0. 7-1.1 (-2.2) cm wide.

(iii) The upper leaf surface typically has short rows of crystalliferous cells immersed in

the leaf tissue (visible as pale spicule-like patterns under magnification) proximal

to appressed-suberect scabrid hairs typically up to 0.5 mm long (Fig. 2B & D).

(iv) Flowers with open corollas typically span 4-5.5 cm in dried material. In the

rheophytes, flowers have open corollas spanning (2.5-)3.5^ cm in dried material.

Thus a rheophytic variant may be encountered that has leaves as small and narrow as 2

cm long and 1 .6 cm wide, with small flowers (spanning only 2.5 cm in dried material).

These are, however, comiected by intennediate states with other variants, all of which

have the hypanthium scales and twig scales described above for the species.

The status of Melastoma normale D.Don, which was reduced to synonymy of

M. malabathricum by Comer (1939) and treated as a subspecies ofM malabathricum

by Meyer (2001), apparently because of similar hypanthium scale types found in both

(Fig. ID-J, K; 2 A, C, E), requires comment. In spite of the type of Melastoma normale

D.Don, which should be a Hamilton collection from Nepal (Don, 1 825), not being

traced (see also Meyer, 2001), there are other Himalayan (including Nepal) collections

available that have been identified with it (Naudin, 1850; Clarke in Hooker, 1879),

such as Hooker s.n. from Sikkim (G-DC, G003 19474), Native Collectors No. 22 from

Khasia [Kliasi] Hills (G-DC, G003 19472), and Wallich 4039 from Nepal (BM, K,

P; also the type of M. wallichii DC., placed in synonymy of M. normale) that amply

display consistent characteristics: a dense twig indumentum of erect-spreading bristles

and leaf upper surfaces with 0.6-1 mm long hairs that are erect and then curved over

(Fig. 2E), visibly longer than those of typical M. malabathricum, Don (1825) himself

noted ''ramis undique setoso-pilosis, foliis... supra hispide pilosis...'' Fig. 2 shows the

similar hypanthium scale character but different leaf upper surface indumentum in

these two species. These moiphological differences, together with the high elevation

distribution of Melastoma normale, and in the absence of any phylogenetic work

to indicate otherwise, suggest that Comer (1939) and Meyer (2001) were hasty in

synonymising or changing the rank of the taxon. As such, Melastoma normale is still

considered a distinct species here.

Additionally, a number of other taxa relegated to the synonymy of Melastoma

malabathricum by Meyer (2001) also seem quite distinct from it, not having the flat

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narrowly triangular-lanceolate scales covering the hypanthium, as inM malabathricum

(Fig. 2A, C). Included are the following

(i) Hypanthium with dense filiform (bristly) scales: Melastoma clarkeanum Cogn. -

TYPE: Bunna, Tenaserrim, 1000 m, Heifer 2241 (K, P); Melastoma homostegium

Naudin - TYPE: Philippines, Cuming 927 (BM, K, P); Melastoma rohustum

Bakh.f. - TYPE: Indonesia, Riau, Lingga, Gunung Walker, Bunnemeijer 6585

(L); Melastoma roemeri Mansf - TYPE: Papua New Guinea, Madang, Hellwig-

Gebirge, 1000-1500 m, Von Roemer 1123 (L). The relationship among these taxa

require study but nonetheless fonn recognisable entities with a very different

hypanthium scale type.

(ii) Hypanthium with narrow-lanceolate scales with extended bristle-like cauda:

Melastoma pelagicum Naudin - TYPE: Solomon Islands, Isle Ysabel, Hombron

s.n. (P).

(hi) Hypanthium with sparse, subulate and subentire scales: Melastoma scabnim Ridl.

- TYPE: Malay Peninsula, Kedah, Langkawi, Telaga Tujuh, Ridley 15813 (K).

There are other taxa placed in the synonymy of Melastoma malabathricum by Meyer

(2001) that, although they have the imbricately airanged serrate hypanthium scales

of M. malabathricum, differ in other characteristics. An example is Melastoma

stenophylliim, which differs by its solitary flowers, a broad-triangular hypanthium

scale type that is less than three times as long as broad (Fig. IL), and near-glabrous

upper leaf surfaces.

Rheophytic taxa and their morphological distinction

Rheophytes are plant taxa adapted to conditions within the flood zone of swift-flowing

streams and rivers, and typically have highly dissected or nan'ow leaf blades (the latter

'stenophylly'), assumed to be adapted to reduce damage by providing less resistance

to swift waters during inundation (Van Steenis, 1981). These plants most often grow

on sandy or stony stream banks or islands, and from rock crevices and at rapids,

developing strong root holdfasts and easily resprouting or branching. Van Steenis

(1981) noted that Northwest Borneo is particularly rich in rheophyte diversity.

A number of Melastoma taxa are also rheophytic in adaptation. Despite their

superficial morphological similarity, typically being low, much-branched shrubby forms

with very naiTowly elliptic to linear leaves, and their shared preference for a similar

ecology within the flood zone of swift-flowing streams, they can be distinguished by

the form of the hypanthium scales, as described in Wong & Low (2015), as well as a

number of other characters. Such rheophytic taxa were either overlooked or have not

been studied for the last revision of the genus by Meyer (2001).

Here we present a key to the Southeast Asian rheophytic species, compared with

M. malabathricum, with which they could be confused because the latter also includes

rheophytic variants. A new rheophytic species, Melastoma kahayanense, is described

Variation in Melastoma malabathricum

393

that has the M. malabathricum hypanthium scale type (flat, triangular to lanceolate,

serrate) (Fig. IM) but with consistently solitary flowers and upper leaf surfaces that

are subglabrous or with very minute hairs, in addition to stenophyllous leaves and a

rheophytic distribution.

Key to Southeast Asian Melastoma rheophytes

la. Hypanthium scales quite flat, margins irregularly serrate to short-ciliate but not

deeply incised towards their base. Calyx without conspicuous intersepalar lobes,

often only a low protuberance (1-1.5 mm high) bearing 1-few prolonged scales

l-2(-3) mm long 2

lb. Hypanthium scales basally thickened or inflated, margins subentire to sparsely

denticulate or the scale deeply incised almost to base to form several linear

segments. Calyx with or without conspicuous, well-fonned intersepalar lobes

2a. Flowers (l-)3-7(-9) in a cyme. Upper leaf surface hairs appressed-suberect and

scabrid, typically up to 0.5 mm long, rarely to 1 mm long, but always conspicuous

with xio magnification (India to SE Asia and the Pacific, including Malesia)

M. malabathricum

2b. Flowers solitary. Upper leaf surface subglabrous to minutely scabrid, hairs very

tiny and barely emergent from leaf surface, requiring x50 magnification to see

clearly (Borneo, Sumatra) 3

3a. Leaf blades (2. 5-) 5. 5-7 (-9) cm long, (0.4-)0.8-l (-1.2) cm wide. Flowers in dried

material with open corolla spanning 4.5-5 cm, hypanthium 0.5-0. 6 cm diameter,

calyx lobes 7-8 mm long; hypanthium scales broadly triangular, their length less

than 3 times the width (Sumatra) M stenophyllum

3b. Leaf blades only up to 4 cm long, 0.6 cm wide. Flowers in dried material with

open corolla spamiing 2.5-3 cm, hypanthium not more than 0.5 cm diameter,

calyx lobes 3.5-6 mm long; hypanthium scales narrowly triangular to lanceolate,

their length 3 times the width or more (E Kalimantan) M. kahayanense

4a. Hypanthium scales subulate, subentire to sparsely denticulate (occasionally with

small teeth near the scale apex). Calyx without distinct intersepalar lobes, often

just low protuberances bearing 1-few prolonged scales to 2 mm long. Upper leaf

surface mostly glabrous or sometimes with minute hairs barely emergent from

leaf surface, requiring x50 magnification to see clearly (NW Bomeo: Brunei,

Sarawak, W Kalimantan) M. ariffinii

4b. Hypanthium scales deeply divided almost to base to fonu several linear segments.

Calyx with conspicuous, well-formed narrowly triangular intersepalar lobes 4-5

mm long. Upper leaf surface totally glabrous (Brunei: Ingei River) M. ashtonii

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Rheophytic Melastoma enumerated

1. Melastoma ariffinii K.M.Wong, Gard. Bull. Singapore 67: 71 (2015). - TYPE:

Bmnei, Tutong, Ramba, Ulu Tutong, down valley to SW of helicopter pad LP 239,

150-200 m asl, 8 May 1992, Johns, Niga, Shanang & Han 7547 (holotype BRUN;

isotype K).

Melastoma polyanthum van linearifoUum Bakh.f., Meded. Bot. Mus. Herb. Rijks Univ.

Utrecht 9 1 : 69 ( 1 943). - TYPE: Borneo [West Kalimantan], Giinong Narik, Mt Kelam,

May 1894, Molengraaff B3460 (lectotype L, designated in Wong & Low, 2015).

Additional specimens examined. BRUNEI: Temburong: First big waterfall on R. Temburong,

c. 500 ft, 6 Nov 1959, Ashton BRUN 759 (BRUN, K, SING). SARAWAK: 5th Division: Ulu

Lawas, Kota Forest Reserve, on bank of Sg. Kota, 21 Oct 1971, Chai & Ilias S 31110 (A, K,

L, SAN, SAR, SING). Kapit Division: Balleh, Ulu Mengiong, Sg. Entejum, 27 Oct 1988,

Othman, Rantai & Jiigah S 56078, (K, KEP, L, SAR, SING); Sg. Belaga at upper rapids, 12

Apr 1963, Ashton S 18242 (K, L, SAR, SING).

This occurs in the Northwest Borneo region, including Brunei, Sarawak and W

Kalimantan.

2. Melastoma aslitonii K.M.Wong, Gard. Bull. Singapore 67: 74 (2015). - TYPE:

Brunei, Belait, Sg. Ingei, rocky river bank, 21 January 1959, Ashton BRUN 5639

(holotype BRUN; isotypes K, L, SING).

Additional specimens examined. BRUNEI: Belait: Falls just upstream from Batu Melintang, 4

Jan 1989, Wong WKM 680 (BRUN, K, SING).

This species seems to be restricted to the Sungai Ingei area in Brunei.

3. Melastoma kahayanense K.M.Wong, sp. nov.

This species resembles M. malabathricum in its flat, irregularly serrate, imbricate

hypanthium scales but differs by its solitary flowers and near-glabrous upper leaf

surfaces, in addition to its stenophyllous leaves and rheophytic habit. - TYPE: Borneo,

E Kalimantan, headwaters of Sungai Kahayan, 0”28’S 1 13®44’E, 200 m asl, 30 March

1988, Mahyar 890 (holotype SING; isotypes A, BO). (Fig. 3A, B)

Rheophytic bush to 2 m high. Twigs with a dense covering of appressed triangular to

ovate-lanceolate flat and finely serrate-edged scales. Leaves with petioles 0.2-0.4 cm

long, c. 0.05-0.1 cm diameter; blades linear, 1.2^ cm long, 0.2-0. 6 cm wide, the 3

longitudinal veins sunken on the upper surface and prominent on the lower surface,

covered by a mixture of larger lanceolate appressed scales (c. 0.5 mm long) and smaller

appressed lanceolate scales (<0.2 mm long), lamina on the upper surface in dried

Variation in Melastoma malabathricum

395

Fig. 3. Hypanthium scale types and upper leaf surface detail. A, B. Melastoma kahayanense

K.M.Wong. A. Hypanthium seale narrowly triangular-lanceolate, the length 3 times or more the

width. B. Upper leaf surfaee with eonspicuous pale elongate spicule-like rows of crystalliferous

cells, most of which are barely emergent as minute hairs. C, D. Melastoma stenophyllum Merr.

C. Hypanthium scale broad-triangular, the length less than 3 times the width. D. Upper leaf

surface with conspicuous pale elongate spicule-like rows of crystalliferous cells, most of which

are barely emergent as minute hairs. A, B from Mahyar 890 (SING); C, D from Rahmat Si

Boeea 7000 (SfNG). (Photomicrographs courtesy of Y.W. Low)

material with abundant conspicuous elongate spicule-like rows of crystalliferous

cells which are typically immersed or proximal to minute hairs barely emergent from

leaf surface and requiring x50 magnification to see clearly, on the lower surface with

minute appressed scabrid hairs up to 1.5 mm long. Flowers solitary, pedicel 2.5-4

mm long; hypanthium 4-6 mm long, 3-4 mm diameter, brownish green, densely

provided with scales, the scales fiat, narrowly triangular to lanceolate, 0.3-0.8(-l)

mm long, the length 3-4 times the width, with finely short-serrate margins (the

serrations less than 0.1 mm deep), lobes 5, triangular, 3.5-6 mm long, 1.5-2. 5 mm

wide, without conspicuous intersepalar lobes; petals obovate, 1.3-1. 5 cm long, 0.8-1

cm wide, noted as purple; stamens 10, 5 longer and 5 shorter. Fruits c. 6-8 mm long,

4-5 mm diameter.

Etymology. The species is named after its type locality, the Kahayan River in East

Kalimantan, Indonesian Borneo.

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Additional specimens examined. BORNEO: East Kalimantan: headwaters of Sungai Kahayan,

5 Ian NE of Haniwu Village, 0°28’S 113°44’E, 200 m asl, 3 Apr 1988, Burley et al. 574 (A,

BO, SING).

Distribution and habitat. Apparently restricted to the Kahayan river basin in E

Kalimantan, on the banks and rocks of swift-flowing streams.

Provisional lUCN conservation assessment. Data Deficient (DD) because it is

documented only from one expedition and its current status is unknown.

4. Melastoma malabathricum L. rheophytic variants

Melastoma polyanthum var. angustifolia Ridl., FI. Malay Pen. 1: 765 (1922). - TYPE:

[Peninsular Malaysia], Pahang, Kuala Tahan, February 1921, Seimund 952 (lectotype

SING, designated here).

Similarly, the names M. longifolium Naud. (TYPE: Singapore, July 1939, Guillou s.n.

(P)) andM obvolutum var. angustifolium Cogn. (TYPE: Sarawak, 1965-68, Beccari

3974 (K)) are also synonyms of M. malabathricum; in fact the leaves of their type

specimens do not depart significantly from the smallish elliptic leaves typical of

M. malabathricum and are not immediately reminiscent of the leaves of rheophytic

taxa. Van Steenis (1981) accepted Jacobs 5580 from Brunei (Temburong river, 30

Sep 1958 (L)) as a true rheophyte because that specimen grew in rocky islets along

the Temburong river, and identified it as M. borneense Bakli.f. That name (TYPE:

Borneo, without date, de Vriese 168 (L: barcode L0008992)) is a synonym of M.

malabathricum, which is also the identity of the Jacobs specimen; these specimens,

too, do not have unusually stenophyllous leaves.

The following additional collections also amply illustrate rheophytic variants of Melastoma

malabathricum. MALAY PENINSULA: Kelantan: Gunong Stong, Lata Jari, 13 Oct 1934,

Symington KEP 37820 (KEP, SING). Pahang: Kuala Tahan, Feb 1921, Seimund 555 (SING),

956 (SING); Kuala Teku, Feb 1921 , Seimund 419 (SING), 426 (SING), 427 (SING); Sungai

Tahan, 7 Sep 1937, Corner s.n. (SING), 2 Mar 1977, Ng FRI 27026 (KEP, SING), rocky

islets in Tahan River, 1891, Ridley 2664 (SING); Sungai Teku, 20 Feb 1968, Shah MS 1333

(SING).

Note. Ridley (1922) merely cited “collected on the Tahan River by Seimund” but there

are a number of such collections in the SING herbarium. One of these is designated as

the lectotype here.

5. Melastoma stenophyllum Merr., Mich. Acad. Sci. Arts Letters 24: 83 (1938). -

TYPE: Sumatra, near Hoeta Bagasan, 7 September 1934^ February 1935, RahmatSi

Boeea 7000 (holotype MICH; isotypes A, L, SING, US).

Variation in Melastoma malabathricum

397

This species is restricted to Sumatra. Van Steenis (1981) wrongly identified S

18242 from Sarawak and Molengraaff B3460 from West Kalimantan as this species

(both specimens are M. ariffinii; see above). The hypanthium scales (Fig. IL) and

features of the leaf upper surface are also shown in Fig. 3C & D.

Possibility of interspecific hybridisation and introgression

There is molecular evidence for natural hybridisation among Melastoma species,

attributed to overlaps in distribution and flowering time, as well as shared pollinators

(Liu et ah, 2014). There are also species of hybrid origin, such as one in China

(probably mistakenly) identified as Melastoma affine D.Don (the type of which has

not been traced but is of a provenance originally stated as “India Orientalis”) that is

intermediate in moi*phological characteristics between M. candidum D.Don and M.

sanguineum Sims (Liu et ah, 2014), and M inteimedlum Dunn that has been shown

to have most nuclear gene haplotypes shared with M. candidum and M. dodecandrum

Lour, and chloroplast spacer sequences identical to either putative parental species

(Dai et ah, 2012). Thus in the sorting of “variable” taxa, the present work adopts

the approach that the character-state combinations found to be common to the most

number of collections is taken to represent a natural species, whereas odd character-

state combinations could be regarded as possible hybrids or introgression products of

the species with the same floral hypanthium scale features. It is, of course, possible

that a number of distinct, established species occur with the same hypanthium

scale type (e.g., Melastoma malabathricum and M. stenophyllum discussed above;

Fig. ID- 1, L), but then these differ in more than one character. In both the Malay

Peninsula and Borneo, the odd possibilities of hybrids or introgression products are

comparatively few and overlaps are within the range of putative parental species.

Notwithstanding, other consistent character-state combinations in Borneo (where

there is greater physiographic and edaphic differentiation, and possibly more effective

reproductive barriers among populations) are considered to represent distinct species,

especially when the ecology or distribution differs from that of similar species. With

this approach, at least any variants that are considered significantly different are then

identified, and the morphological assessment results in a taxonomy that attempts a

hypothesis for fiirther work on the phylogenetic relationships.

Using this approach, a comparatively small number of collections from Malesia

(lowlands to mountains up to only around 1 800 m) were found to have the hypanthium

scale type of Melastoma malabathricum but which differ in small vegetative characters,

including having twigs with spreading scales (otherwise of the same fonu) or upper leaf

surfaces with longer hairs (0.5-1 .2 mm). These are regarded as representing possibly

hybrid material or introgression products. Below, we list the specimens known as such.

Recommendation Ii.2A of the Melbourne Code (MeNeill et ah, 2012) notes

that names or epithets in a hybrid fomiula should preferably be in alphabetical order

or place the female parent first, or indicate female and male parents using symbols,

emphasising that if a non-alphabetical sequence is used, its basis should be clearly

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indicated. However, in the cases we consider below, the direction of the crosses

are not detennined, but it is of relevance to note which species is thought to have

contributed the hypanthium scale characters and which the stem and branch scale type

or upper leaf surface hair type. Thus in our case we do not use hybrid formulas with a

multiplication sign (x) as prescribed but indicate the hybridity differently, using a plus

sign (+) between the putative donor of the hypanthium scale type (named first) and the

donor of the other character(s) considered (named second).

(i) Putative hybrid or introgression products of Melastoma malabathricum + M.

muticum Ridl.

(Twig scales triangular, ovate or lanceolate, serrate, appressed (as in both species);

upper leaf surface with long hairs, 0.5-0.8(-1.2) mm long, intermixed with shorter

minute hairs, as in Melastoma muticum; hypanthium scales as inM malabathricum).

The possibility of Melastoma muticum hybridising has been considered before by

experienced collectors. There is a note on Stone 10828 (KLU), collected at 1036 m

along the then new Genting Highlands road (with its attendant corridor of disturbance)

up the Peninsular Malaysian Main Range, originally identified as Melastoma muticum

but which we regard here as a putative Melastoma muticum + M. velutinosum Ridl.

(see below), that states: "M. malabathricum just reaches this altitude; they are here

found growing together. Do they [M. muticum and M. malabathricum'] hybridize?"

MALAY PENINSULA: Johore: Gunung Blumut, West Ridge, 2900 ft asl, 14 May 1968,

Whitmore FRI 8716 (KEP, SING). Pahang: Cameron Highlands, between Gunung Brinchang

and Green Cow Area, 1600-2000 m asl, 19 Oct 1967, Shimizu et al. M 13479 (KEP, KYO,

SING). Perak: Gunung Korbu FR, 22 Jul 1933, Symington KEP 32240 (KEP, SING); Maxwell

Hill, summit of Gunung Hijau, 4500 ft asl, 4 Dec 1965, Shah & Sidek MS 1105 (SING).

SUMATRA: North Sumatra: Brastagi, Dec 1930, Symington KEP 23985 (SING).

All these collections are from montane areas where the distribution of Melastoma

malabathricum and M, muticum Ridl. (which has a characteristic mixed indumentum

on the upper leaf surface) overlap.

(ii) Putative hybrid or introgression products of Melastoma malabathricum + M.

sanguineum

(Twig scales sparse, small and ovate-appressed and sometimes mixed with larger

or longer suberect-erect bristly scales, as in M. sanguineum; upper leaf surface and

hypanthium scales as inM malabathricum).

MALAY PENINSULA: Kedah: Gunong Jerai, 3200 ft, Burkill HMB 3345 (SING); Langkawi,

Pulau Ular, 22 Nov 1941, Corner s.n. (SING). Penang: Air Itam Dam Reserve, 3 Mar 1965,

Hardial & Samsuri 227 (SING); enroute from Penang Hill to Waterfall Botanic Garden, c. 550

m asl, 14 Oct 1 967, Shimizu et al. M 12990 (KYO, SING); ‘top of Hill’, Dec 1 895, Ridley s.n.

(SING), 1520 ft. Mar 1881, King’s Coll 1540 (SING). Perlis/Thai border: Setul [=Satun],

Mar 1910, Ridley 15056 (SING). Selangor: Ulu Gombak, 18 m.s., 15 Jun 1966, CarrickJC

1486 (SING).

Variation in Melastoma malabathricum

399

(iii) Possible hybrid or introgressioii products with parent species unidentified:

Melastoma malabathricum + unknown taxa

(Twig scales suberect- spreading, not appressed as in Melastoma malabathricum', other

characteristics including hypanthium scale characters as in M. malabathricum).

BORNEO: Sabah: Sipitang, 5 miles from Kampung Mendulong, road to Meligan, 4 Sep 1983,

Lee SAN 96893 (A, SAN, SING). Sarawak: Mulu National Park, along Sg. Melinau, south of

Long Birar, 4‘'00N, 1 100 m asl, 12 Apr 1978, Stone 13681 (KLU, SAR).

MALAY PENINSULA. Kedah: Koh Mai Forest Reserve, 4 Apr 1938, Kiah SFN 37397 (SING);

Kuala Pegang to Kg. Ibul, 6 Feb 1968, Whitmore FRI 4631 (KEP, SING); Langkawi, Guard

Regiment Camp, 10 Dec 1960, Alphonso & Samsuri A 131 (SAN, SAR, SING); Kuah, 4 Aug

1961, Samat 26 (KLU, SING); Langkawi, south coast 4 miles west of Kuah, 10 Dec 1969,

Whitmore FRI 12976 (KEP, SING). Kelantan: Gua Musang, 18 Aug 1971, Boev 302 (KLU);

Kota Bahru, 23 Apr 1937, Corner s.n. (SING). Kuala Lumpur: University of Malaya, around

Jabatan Botani, 11 Sep 1986, Rosmawati 3 (KLU); University of Malaya, wasteland in campus,

6 Sep 1974, Jayamohan s.n. (KLU). Pahang: Tasek Bera, north end, Kuim, 30 m asl, 8 Mar

1969, Flenley 58 (KLU); LIlu Sg. Krau, NE Gunung Benom, 800 ft asl, 1 Mar 1967, Whitmore

FRI 3138 (KEP, SING). Perak: Batu Gajah, 26 Jun 1924, Burkill & HaniffSFN 13362 (SING);

Kuala Kangsar, 14 Jun 1925, HaniffSFN 14919 (SENG); Kuala Kangsar, LubokMerbok, 21 Oct

1924, HanijfSFN 16007 (SING). Selangor: Kepong, Bukit Beruang, 18 Nov 1927, Pawanche

KEP 13422 (SING); Klang Gates quartz ridge, 19 Mar 1960, Carrick 650 (KLU); ibidem, 18

Nov 2006, Lee & Wong s.n. (KLU); Pudu, 16 Aug 1922, Seimimd FMS Museums 10327 (SING);

Ulu Gombak 18 mile stone, 6 Mar 1968, Teo T&P 51 (K, L, SING); Ulu Kuang, May 1972,

Provencher NrT51 (KLU). Trengganu: Kuala Berang, 29 Apr 1937, Corner SFN 33490 (SING).

Singapore: Changi Reserve, 11 Mar 1889, Goodenough s.n. (SENG); Seletar Reservoir, 10 Mar

191 \, Anthony 259 (SING).

A number of such collections have odd characters noted: having witches broom

{Goodenough s.n.) or galls {Anthony 259), chlorotic {Burkill & Haniff SFN 13362),

or with white flowers {Seimund FMS 10327). It is not possible to detemiine without

appropriately designed studies if these could be related to recessive conditions such as

produced in a hybrid swarm.

Possible hybrid or introgression products of Melastoma muticum + M.

malabathricum also exist. This material has hypanthium scales resembling those of

Melastoma muticum (very narrow, near-filiform scales with length 5-8 times the width,

and sparse inconspicuous short cilia on the margin); twig scales triangular-ovate and

appressed as in M. malabathricum', upper leaf surface with long and very minute hairs

intermixed as inM muticum.)

MALAY PENINSULA: Pahang: Cameron Highlands, Gunong Brinchang, 5 Nov 1960, Poore

453 (KLU); ibidem, sunmiit, 6500 ft, 11 May 1965, Stone 5612 (KLU).

Also, there are specimens that are possibly the hybrid or introgression products of

Melastoma muticum + M. velutinosum (which have the hypanthium scale type of M.

muticum but suberect bristles covering the twigs as in M. velutinosum)', both species

are found in the Main Range mountains of the Malay Peninsula.

400

Gard. Bull. Singapore 67(2) 2015

MALAY PENINSULA: Pahang: Genting Highlands, 3400 ft [1036 m] asl, 8 Jul 1972, Stone

10828 (KLU); Ulii Kali, 1 Dec 2012, Sugumaran SM 345 (KLU).

It may seem, from this listing, that a good number of collections are considered

of hybrid or introgression origin. In fact, such variation has not been detected for all

species, and for some species less widespread than Melastoma inalabathricum, the

number of such odd taxa is very small, represented by I-few collections. It would

stand to reason that in a very widespread species, such as Melastoma malabathricum,

distributed from Sri Lanka through Malesia and mainland Southeast Asia to north

Australia and the Pacific region, and from near sea level to around 1 800 m on mountains,

there would be more collections of putative hybrids or introgression products to be

expected.

ACKNOWLEDGEMENTS. I am grateful to the curators of the following herbaria for hosting

these studies: BRUN, KLU, SAN, SAR, SING. Low Yee Wen, Bazilah Ibrahim and Alsa Moo

of SING were especially helpful with specimen imaging and annotations. David Middleton's

suggestions have helped improve the manuscript for publication.

References

Cogniaux, C.A. (1891). Melastomaceae. Monogr. Phan. 7: 314-1183.

Comer, E.J.H. (1939). Notes on the systematy and distribution of Malayan phanerogams III.

Gard. Bull. Straits Settlements 10: 239-329.

Dai, S., Wu, W., Zhang, R., Liu, T., Chen, Y, Shi, S. & Zhou, R. (2012). Molecular evidence for

hybrid origin Melastoma intermedium. Biocliem. Syst. Ecol. 41: 136-141.

Don, D. (1825). Prodromus Florae Nepalensis. London: J. Gale.

Hansen, C. (1977). The Asiatic species oi Osbeckia. Ginkgoana 4: 1-150.

Hooker, J.D. (1879). Flora of British India, vol. 2. London: L, Reeve & Co.

Jarvis, C.E., Barrie, F.R., Allan, D.M. & Reveal, J.L. (1993). A ListofLinnaean Generic Names

and Their Types. Regnum Vegetabile 127. Konigstein: Koeltz Scientific Books.

Li, H.-L. (1944). Studies in the Melastomataceae of China. J. Arnold Arbor. 25: 1-41.

Liu, T., Chen, Y, Chao, L., Wang, S., Wu, W., Dai, S., Wang, F., Fan, Q. & Zhou, R. (2014).

Extensive hybridization and introgression between Melastoma candidum and M.

sanguineum. PLOS One, DOI: 10.1 371 /joumal.pone,0096680.

McNeill, J., Banie, F.R., Buck, W.R., Demoulin, V., Greuter, W., Hawksworth, D.L., Herendeen,

P.S., Knapp, S., Marhold, K., Prado, J., Pmd’homme van Reine, W.F., Smith, G.F.,

Wiersema, J.H. & Turland, NJ. (2012). International Code of Nomenclature for algae,

fungi, and plants (Melbourne Code). Regnum Vegetabile 154. Konigstein: Koeltz

Scientific Books.

Meyer, K. (2001). Revision of the Southeast Asian genus Melastoma (Melastomataceae).

Blumea 46(2): 351—398.

Naudin, C. (1850). Melastomaceamm quae in Musaeo Parisiensi continentur monographicae

descriptionis et secundum affinitates distributionis tentamen. XXVIII. Melastoma. Ann.

Sci. Nat. 13: 273-296.

Variation in Melastoma malabathricum

401

Ridley, H.N. (1922). The Flora of the Malay Peninsula, vol. 1. Polypetalae. London: L. Reeve

& Co. {Melastoma, Pp. 763-766)

Van Steenis, C.G.G.J. (1981). Rheophytes of the World: An Account of the Flood-Resistant

Flowering Plants and Ferns and the Theory of Autonomous Evolution. Alphen aan den

Rijn, The Netherlands: Sijthoff & Noordhoff.

Wong, K.M. & Low, Y.W. (2015). Novitates Bmneienses, 3. Eight new woody plants in the

Brunei flora, including five new species. Card. Bull. Singapore 67: 69-84.

Gardens’ Bulletin Singapore 67(2): 403-406. 2015

doi: 10.3850/S2382581215000320

403

Clarifying the nomenclature

of Crateva trifoliata (Capparaceae)

LM. Turner

Research Associate, Royal Botanic Gardens, Kew

Richmond, Surrey TW9 3AE, U.K.

ABSTRACT. Capparis trifoliata Roxb., the supposed basionym of '‘Crateva adansonii subsp.

trifoliata (Roxb.) Jacobs’ and "Crateva trifoliata (Roxb.) B.S.Sun’, is shown to represent an

illegitimate renaming of Crateva religiosa G.Forst. However, Jacobs effectively excluded the

nomenclatural type of Capparis trifoliata, thus avoiding creating a superfluous name, and

instead described Crateva adansonii subsp. trifoliata Jacobs validated by the citation of the

Latin description of Crateva erythrocarpa Gagnep. Sun also excluded the type of Crateva

religiosa, but his effective transfer of C. adansonii subsp. trifoliata to specific rank does not

have priority over Crateva erythrocarpa Gagnep., which is therefore the couect name for

this taxon when it is considered a distinct species. The name C/^ateva rmrvula Buch.-Ham. is

formally lectotypified.

Keywords. Capparis, China, Crateva, India, nomenclature, Roxburgh, typification

Introduction

William Roxburgh named an Indian species Capparis trifoliata. The name was included

in Roxburgh’s Hortus Bengalensis (Roxburgh, 1814: 41) but as no description was

included the name remained invalid. A brief description was eventually published in

the second edition of Flora Indica (Roxburgh, 1832: vol. 2, p. 571), 17 years after

Roxburgh had died. Jacobs (1964), in a revision of the genus Crateva, included

Capparis trifoliata within the wide-ranging (seasonally dry tropical Africa and Asia)

and variable Crateva adansonii DC. In order to categorise the variation within the

species, Jacobs recognised a series of five subspecies separable on leaf characters.

These consisted of Crateva adansonii subsp, adansonii in Africa, C. adansonii subsp.

odora (Buch.-Ham.) Jacobs in India, C. adansonii subsp. trifoliata (Roxb.) Jacobs

in Indochina, C. adansonii subsp. formosensis Jacobs in China and Taiwan and C.

adansonii subsp. axillaris (C.Presl) Jacobs in the Philippines and Java. While some

recent works have favoured splitting up Crateva adansonii, others have maintained

Jacobs’s system (Chayamarit, 1991; Liu & Liao, 1996; Philcox, 1996). However,

there is a nomenclatural problem with Capparis trifoliata Roxb., upon which Jacobs

evidently based C. adansonii subsp. trifoliata. In the original validation in Flora Indica,

Roxburgh cited "Crateva religiosa Willd. 2. 853’ in synonymy. This is a reference to

Willdenow’s Species Plantarum. In tum, Willdenow referred to Vahl (1794: 62) and

Forster (1786: 203). The latter element (also refen*ed to by Vahl) relates to the original

404

Gard. Bull. Singapore 67(2) 2015

publication of Crateva religiosa G.Forst. Roxburgh, Willdenow and Vahl, in their

somewhat confused view, also all referred to the Hortus Malabaricus element ‘niirvala’

(Van Rlieede tot Draakestein, 1682: 49-50, t. 42) - now considered to represent a

separate species. Roxburgh stated that his species was native to India and certainly

included what Jacobs later referred to as Crateva adansonii subsp. odora as shown by

a Roxburgh specimen from William Hooker’s herbarium now at Kew and Roxburgh’s

leones no. 157 also at Kew. Jacobs (1964; 199) referred to a Roxburgh specimen

in the Herbarium of the East India Company as the holotype of Capparis trifoliata.

This does seem to be a specimen of Crateva adansonii subsp. trifoliata. It is mounted

with another Roxburgh specimen but Jacobs made it clear that he was referring to

the specimen annotated ‘ 1’ on the sheet. The origins of the specimen are unknown.

Nomenclaturally, Capparis trifoliata Roxb. represents a superfluous renaming of

Crateva religosa and, as no type was indicated by Roxburgh, it is typified by the type

of C. religiosa (McNeill et al., 2012: Art. 7.5), Therefore .Jacob’s subspecies’ name

also appears to be based on Crateva religiosa. Similarly so does "Crateva trifoliata

(Roxb.) B.S.Sun’ which has been used in recent Chinese floras (Sun, 1999; Zhang &

Tucker, 2008),

However, apparently unwittingly, Jacobs (1964) effectively excluded the type

of Crateva religiosa from C, adansonii subsp. trifoliata by recognising Crateva

religiosa as a taxon distinct from C. adansonii subsp. trifoliata in the same paper. Such

exclusion by implication is permitted under the code (McNeill et al., 2012: Art. 52.2;

cf Ex. 7). While Jacobs provided a description in English of Crateva adansonii subsp.

trifoliata this is not sufficient to validate a new taxon at this date. However Jacobs also

cited Crateva erythrocarpa Gagnep. in synonymy. This was published by Gagnepain

with a Latin description. Therefore Crateva adansonii subsp. trifoliata is validated by

the Latin description of C. eiythrocarpa.

Sun (1999) also seemed to base Crateva trifoliata on Capparis trifoliata Roxb.

However, Sun excluded Crateva religiosa G.Forst. from the flora of China, noting

that the application of the name to the species used to make fish lures in Taiwan

and the Ryukyus was en'oneous. Therefore, like Jacobs, Sun excluded Roxburgh’s

nomenclatural type and effectively created a new combination at species rank based

on Jacobs’s subspecies. But this combination only has priority at species rank from

the date of Sun’s publication. Therefore the correct name at species rank is Crateva

erythrocarpa Gagnep.

Crateva adansonii subsp. trifoliata Jacobs, Blumea 12: 199 (1964). - Crateva trifoliata

(Jacobs) B.S.Sun, FI. Reipubl. Popularis Sin. 32: 489 (1999). - TYPE: W. Roxburgh

s.n. (ETC 6972C, specimen annot. 1) (holotype K-W [barcode no. KOOl 126439]).

Crateva erythrocarpa Gagnep., Bull. Soc. Bot. France 55: 322 (1908). - TYPE:

Vietnam, montagne de Chaudoc, .July 1 876, F.-J. Harmand 608 (lectotype P [barcode

no. P05427398], designated by Jacobs (1964: 199)),

Nomenclature of Crateva trifoliata

405

Crateva religiosa G.Forst, Diss, PI. Esc. 45^6 (1786). - Capparis trifoliata Roxb.,

FI. Lnd. 2: 571 (1832), nom. illeg., superfl. - TYPE: ?India, G. Forster s.n. (lectotype

K [barcode no. K000651081], designated by Jacobs (1964: 191)).

Roxburgh, Willdenow and Vahl all cited the Van Rlieede element ‘ niirvala ’ in synonymy.

This has also been considered to be a separate species, Crateva nw^ala Buch.-Ham.

Jacobs (1964: 194) effectively lectotypified the name with a Buchanan-Hamilton

specimen in the Wallich Fferbarium (K-W). However, as Nicolson et al. (1988) have

noted, Buchanan-Hamilton did not consider his own specimens to be the tme Crateva

nurvala but a variety of it. Therefore Jacobs’s typification is incorrect. Nicolson et

al. (1988) stated ‘However it appears that Van Rheede’s illustration should be taken

as the type.’ It is questionable whether this is a valid typification - the opening of the

sentence casts doubt making it unclear that the authors are accepting their typification.

I therefore formally typify the name here. Note that after the type of a Loureiro name

was refound Crateva nw^ala was included in synonymy of Crateva magna (Lour.)

DC. (Jacobs, 1976).

Crateva magna (Lour.) DC., Prodr. 1: 243 (1824). - Capparis magna Lour., FI.

Cochinch. 1: 331 (1790). - TYPE: Cochinchina, J. Loureiro s.n. (lectotype BM

[barcode no. BM000629693], designated by Jaeobs (1976: 822)).

Crateva nwyala Buch.-Ham., Trans. Linn. Soc. London 15: 121 (1827), as ‘niirvala’.

- TYPE: Van Rheede, Hort. Malab. 3: t. 42 (1682) (lectotype, designated here).

ACKNOWLEDGEMENTS. I am very grateful to Prof J. McNeill (E) for guidance on

nomenclatural matters and Wong Wei San (SUTD) for Chinese translation. Comments from

two anonymous reviewers helped improve the paper.

References

Chayamarit, K. (1991). Capparaceae. In: Smitinand, T. & Larsen, K. (eds) Flora of Thailand

5(3): 241-271. Bangkok: Royal Forest Department.

Forster, J.R. (1786). Florulae Insularum Australium Prodromus. Goettingen: Joann. Christian

Dieterich.

Jacobs, M. (1964). The genus Crateva. Bluniea 12: 177-208.

Jacobs, M. (1976). Capparaceae. In: Van Steenis, C.G.G.J. (ed) Flora Malesiana 7(4): 822.

Leyden: Noordhoff International Publishing.

Liu, T.-S. & Liao, J.-C. (1996). Capparaceae. In: Huang, T.-C. (ed) Flora of Taiwan, 2nd

edition, 2: 734-744. Taipei: Editorial Committee of the Flora of Taiwan.

406

Gard. Bull. Singapore 67(2) 2015

McNeill, J., Barrie, F.R., Buck, W.R., Demoulin, V., Greater, W., Hawksworth, D.L., Herendeen,

RS., Knapp, S., Marhold, K., Prado, J., Prud’homme van Reine, W.F,, Smith, G.F.,

Wiersema, J.H. & Turland, NJ. (2012). International Code of Nomenclature for algae,

fungi, and plants (Melbourne Code). Regnum Vegetahile 154. Konigstein: Koeltz

Scientific Books.

Nicolson, D.FI., Suresh, C.R. & Manilal, K.S. (1988). An Interpretation of Van Rheede s Hortus

Malabaricus. Regnum Vegetabile 119. Konigstein: Koeltz Scientific Books.

Philcox, D. (1996). Capparaceae. In: Dassanayake, M.D. {qC) A Revised Handbook to the Flora

of Ceylon 10: 23-50. Rotterdam: A.A. Balkema.

Roxburgh, W, (1814). Hortus Bengalensis. Serampore: Mission Press.

Roxburgh, W. (1832). Flora Indica, vol.1-3. Calcutta: W. Thacker & Co.

Sun, B. (1999), Capparaceae (excluding Cleome). In: Wu, C. (ed) Flora Reipublicae Popularis

Sinicae 32: 484-531. Beijing: Science Press.

Vahl, M. (1794). Symbolae Botanicae, vol. 3. Copenhagen: Nicolaus Moller et filius.

Van Rheede tot Draakestein, FI. A. (1682). Horti Indici Malabarica pars Tertia. Amsterdam:

Van Someren & Van Dyck.

Zhang, M. & Tucker, G.C. (2008). Capparaceae. In: Wu, Z., Raven, P.H. & Hong, D. (eds)

Flora of China 7: 433^50. Beijing: Science Press.

Gardens’ Bulletin Singapore 67(2): 407-408. 2015

doi: 10.3850/S2382581215000332

407

BOOK REVIEW: A Guide to Begonias of Borneo. Ruth KieWj Julia Sang, Rimi

Repin and Joffre Ali Ahmad. 2015.

Kota Kinabalu: Natural History Publications (Borneo). 21.5 x 15 cm, soft or hard cover, 304

pp. ISBN 978-983-812-160-6 (softcover), ISBN 978-983-812-161-3 (hardcover). Price RM 80

(softcover), RM 140 (hardcover).

Begonias are well-known ornamentals which are

cultivated for their spectacular blooms and range of leaf

shapes and colouration. The diversity of the common

hybrids pales, however, in comparison to the diversity of

wild begonias. Begonia is estimated to have more than

1 800 species, placing it amongst the ten largest genera of

flowering plants. Some 200 species have been described

from the biodiversity hot-spot of Borneo alone, but no

monograph on Bornean begonias is available. Therefore,

up till now detailed information on Bornean species could

only be accessed by consulting an array of specialist

literature. Photos capturing the stunning variation in leaf

shape, colour and texture, as well as their flowers, have

not been available for most Bornean species.

The authors of A Guide to Begonias of Borneo, together combining decades

of experience of Begonia systematics and conservation, managed to unlock this

information for anybody interested in Begonia and appreciative of botanical diversity

and the rich natural heritage of Borneo. The guide provides a richly illustrated 25-

page introduction giving concise information on previous research, diversity, typical

habitats, dispersal biology, identiflcation and conservation of Bornean Begonia. The

main body of the guide consists of species pages, with usually two or sometimes more

pages dedicated to each of the 134 featured species (c. 70% of the documented species

diversity in Borneo). These species pages include (i) brief descriptions of growth habit,

leaves, flowers and fruits, sometimes elaborating on unique features of the species; (ii)

a full page photograph of the growth habit, usually a large photograph showing an

inflorescence or a fruiting or flowering leafy branch, as well as some smaller inset detail

shots of flowers and fruits; (iii) information on species distributions on Borneo, but no

distribution maps, and usually also some information on habitats where the species can

be found; (iv) notes on the etymology of the species epithet and, sometimes, additional

information on plant usage and ethnobotany.

The descriptions are concise and use only few technical terms, making information

on the species easily accessible, and the figures are of high quality. The guide does not

provide an identification key to the species nor specimen-level information but, given

the current rate of new species discovery, this kind of information in a static book

format would soon be outdated and would be better placed in a more dynamic online

database format (e.g., see the Begonia Resource Centre available at http://padme.rbge.

org.uk/begonia/). The descriptions and figures are detailed enough, however, to aid

with species identification.

408

Card. Bull. Singapore 67(2) 2015

The guide can serve as a beautiful coffee table book, but it is also scientifically

relevant. In the introduction, the authors emphasise that every field trip in Borneo

brings to light further new species of Begonia, and that the approximately 200 species

described from the island probably represent only a fraction, possibly about one

third, of the estimated total Bornean Begonia species diversity. Given the narrow

distributions of the vast majority of Asian Begonia species and the rates of land

conversion and deforestation in Borneo, this means that many species will likely go

extinct before they have been described. In this race against time, the recent precursor

papers in the Journal Sandakania, m which 60 new Bornean species were described,

and A Guide to Begonias of Borneo, are key publications. These will stimulate interest

in this fascinating group of plants and greatly facilitate further species descriptions and

assessments of the conservation status of Bornean Begonia.

Daniel C. Thomas

Singapore Botanic Gardens

Gardens’ Bulletin Singapore 67(2): 409-410. 2015

doi: 10.3850/S2382581215000344

409

BOOK REVIEW: Forest Trees of Southern Thailand, Volume 1 (A-Es). Simon

Gardner, Pindar Sidisunthorn and Kongkanda Chayamarit. 2015.

Bangkok: Kobfai Publishing Project. 24 x 17 cm, hard cover, 768 pp. ISBN 978-616-7150-39-0

(English edition), ISBN 978-616-7150-43-7 (Thai edition). Price 2200 Baht.

In 2000, the remarkable book A Field Guide to the

Forest Trees of Northern Thailand was published in

both English and Thai. That book contained easy-to-

use keys, simple descriptions and numerous paintings

and photographs, thereby making identification of

the trees of the region, and access to information

about them, much simpler. It became a best seller.

The same three authors have now turned their hands

to a sister project, the trees of southern Thailand,

again with English and Thai language editions. The

biggest difference is in the sheer scale of the task:

southern Thailand has a much greater number of

tree species than northern Thailand and no longer

will the information fit into a single volume. Indeed

three volumes are planned of which the first has

just been published, the second is planned for early

2016, and the third for late 2016. The co mm itment to excellence displayed in the

northern Thailand book is again very evident in the first volume of this new work. It is

both an excellent source of knowledge and simply a beautiful book to peruse.

The inside front and back covers provide an illustrated glossary of technical

botanical terms (e.g. leaf type, leaf arrangement, leaf shape, inflorescence type, fruit

type etc.). The opening chapter gives information on how to use the guide, how to go

about identifying trees, the classification of trees into orders and families (using APG

III), and how to collect parts from trees in order to identify them later. The Introduction

is divided into three sections: Geography, Flora and Vegetation. The Geography

section includes information on geology, topography and climate; the Flora section

tells us there are at least 2250 tree species in southern Thailand, comprising about

75% of Thailand’s total, as well as how geographical and historical factors have led to

such species richness in the region; and the Vegetation section guides us through the

major forest types and their major floristic compositions. Black and white photos and

drawings abound to illustrate the text.

The bulk of the book, however, is given over to the systematic presentation

of the trees of southern Thailand. The arrangement is alphabetical by family, then

alphabetical by genus within each family, and then alphabetical by species within each

genus. Keys are provided to the genera within a family and to the species within a

genus for the larger genera. For the larger families spot characters are given for the

genera to aid identification. Families, genera and species are described, with diagnostic

characters highlighted in bold type. But this is no ordinary Flora, as more than half

of almost every page of the volume is devoted to colour photographs, line drawings

410

Gard. Bull. Singapore 67(2) 2015

and paintings of trees. The authors recommend various ways of using the information

contained within the volume to identify trees, including just flicking through and

looking at the pictures. As a professional botanist myself, used to using weighty

academic Floras, 1 have lost count of the number of times that 1 have fruitlessly been

attempting to identify something and wished 1 could just look at pictures. This guide to

southern Thai trees gives us both licence and opportunity to indulge in such activities!

The first volume includes all plant families from Acanthaceae to Escalloniaceae

so includes major families such as Annonaceae, Apocynaceae, Burseraceae,

Combretaceae and Ebenaceae. And, of course, it has the Dipterocarpaceae, the

characteristic family of Asian tropical forests. Eight genera and over 70 species are

covered in the book from this family alone, with photographs and drawings of whole

trees, bark, leaves, flowers and fruits for most of them.

Inevitably in a project of such scale not all identifications are correct. For

example, Johnson & Murray’s paper on Xylopia in this issue of Gardens ' Bulletin

Singapore notes that there are two misidentifications in this genus. At the same time,

however, the Xylopia paper draws our attention to an extremely positive aspect of the

wider project that resulted in this book. A vast amount of field work was necessary

to collect and photograph the species in this volume and those collections are, and

will continue to be, available for taxonomic study. Johnson & Murray’s study of the

collection accompanying the photos of the incon'ectly identified Xylopia vielana

led to the description of a new species, Xylopia platycarpa. Perhaps more such

examples will arise, particularly in those families not yet completed for the Flora of

Thailand. Nevertheless, it is abundantly clear that the authors have striven hard to

produce a work of botanical accuracy as well as aesthetic appeal and a small number

of misidentifications is an inevitable consequence of such an ambitious project. The

authors are to be commended for having that ambition and seeing it to fruition.

David J. Middleton

Singapore Botanic Gardens

Gardens’ Bulletin Singapore 67(2): 411-414. 2015

doi: 10.3850/S2382581215000356

411

BOOK REVIEW: Flora of Cambodia, Laos and Vietnam

Fascicle 33 Apocynaceae by David J. Middleton. 2014.

(288 pages, 48 colour photographs, 48 line drawings; paperback; text in English; 160 x 240

mm. ISBN 978-2-85653-751-0. £35 incl. packing & postage)

Fascicle 34 Polygalaceae by Colin A. Pendry. 2014.

(72 pages, 14 colour photographs, 5 line drawings; paperback; text in English; 160 x 240 mm.

ISBN 978-2-85653-752-7. £15 incl. packing & postage)

Fascicle 35 Solanaceac by Sovanmoly Hul & Pauline Dy Phon. 2014.

(104 pages, 29 colour photographs, 25 line drawings; paperback; text in French; 160 x 240 mm.

ISBN 978-2-85653-750-3. £20 incl. packing & postage)

Paris: Publications scientifiques du Museum national d’Histoire naturelle & Edinburgh: Royal

Botanic Garden Edinburgh.

FLORA of

CAMBODIA, LAOS

and VIETNAM

FLORA of ^

CAMBODIA, LAOS

and VIETNAM

FLORE du

CAMBODGE, du LAOS

et du VIETNAM

"SOLANACEAE

Sovanmoly Hut & l^uline Dy Phon

It has been ten years, a whole decade, after the publication of the last issue of the

Flore du Cambodge, du Laos et du Vietnam (Flora of Cambodia, Laos and Vietnam,

or FCLV), a project begun by Andre Aubreville in 1960 to succeed La Flore generale

de I Lndochine (The General Flora of Indochina) coordinated by H. Lecomte & H.

Humbert during 1907-1934, and H. Humbert until 1951. That last issue was Fascicle

32, Myrsinaceae, by Chi Ming Hu & Jules Vidal, issued in December 2004.

The new issues are Fascicle 33 (Apocynaceae, by David J. Middleton, in

English), Fascicle 34 (Polygalaceae, by Colin A. Pendry, in English), and Fascicle

35 (Solanaceae, by Sovanmoly Hul & Pauline Dy Phon, in French), all issued in

December 2014.

These finely produced issues not only welcome back a key regional project

after a hiatus but also see a redoubled organisation and collaboration to engage more

specialists in what is the longest-running international science endeavour in the

Cambodia, Laos and Vietnam (CLV) region. The added use of English as an alternative

language will certainly make the revisions more amenable to potential contributors as

well as more accessible to users of floras.

It is easy to see why botanists have been attracted to this region and perhaps

also fairly easy to understand why efforts at the documentation of the plant life have

412

Gard. Bull. Singapore 67(2) 2015

not been on a dauntless path since the Portuguese Jesuit missionary Joao de Loureiro

published the Flora Cochinchinensis in 1790. Loureiro's work was an extension of his

interest in local medicinal plants after spending 35 years, mostly as chief mathematician

and naturalist, in the service of the King of Cochinchina, a region in the southern

third of modem Vietnam that included Saigon and Hue, and at some later point a

French colony (Memll, 1935). Loureiro's specimen collections, which would have

been important reference material, were partly destroyed, and his plant descriptions

were very brief and, understandably, often ambiguous or unclear. The CLV region has

been, and is still to a large degree, a landscape difficult to traverse: initially because of

thick vegetation cover, disease and mountainous terrain in many parts, and latterly still

with the risk posed by landmines from the 20th century war years that riddle parts of

the landscape. The relative isolation of mainland SE Asia caused by the Miocene uplift

of the Himalayas, connectivity with the Chinese Henduan Mountains to the northwest

and the main South Chinese karst region to the north, general climatic distinction

from the wetter tropical region of Malesia to the south, as well as emergence through

episodes of climatic oscillations, geological changes and vegetation range expansions

and contractions, have been important biogeographic features that have shaped a very

rich and special biota. Loureiro's work was important but can be difficult to relate

to other pioneering botanical work with floristic overlaps in the region, especially

when specimen material is lacking or poorly referenced. The trauma of the Second

World War and subsequent political realigmnents and regional conflicts would stretch

the pause in scientific research into the 1990s. And, meanwhile, floristic and other

progress has raced on around the CLV region. It is not surprising that the CLV flora

remains one of the poorest explored and documented, complex to tackle even for the

specialist, and for which much painstaking research is now required..

Economic refonns and the opening up of the CLV region have brought its

countries into wider collaboration with outside botanical institutions. The early

French effort, fostered much through the collecting activities of such as Balansa,

Chevalier, Eberhardt, Haiinand, Petelot, Pierre, Schmid, Thorel, Vidal, and Poilane,

brought in some 75,000 specimen collections (Regalado et al., 2005); probably now

over 30,000 collections have been added through collaborative programmes since

the 1990s. Encumbered by so many historical factors, it is still most meaningful to

approach the flora of this block of mainland SE Asia with a single definitive regional

flora — even if each country uses a different language and, as faster progress on the

Vietnamese front demonstrates, there is a local need to produce national floras, such as

the Flora of Vietnam series (in Vietnamese) initiated through the Hanoi-based Institute

of Ecology & Biological Resources (which can emphasize conservation efforts more

directly), and checklists, as in the 3 -volume Checklist of Plant Species of Vietnam

(in Vietnamese) produced through the Vietnam National University in Hanoi. In the

end, all checklists and floras will be subject to revision — some more intensively than

others — as further exploration, collecting and revision will show. In this regard, even a

number of earlier FCLV accounts are ripe for revision, although tackling these should

subsequently be more systematic and easier given its regional scope. Even with a

shift towards online floras that are more easily updated as parts are made available.

413

continuing the printed version of the FCLV is not without merit. It is one thing being

able to access websites with a good satellite cellphone from just about anywhere, and

another to afford sufficient and up-to-date computer equipment and software to carry

out sustained flora work that could make reference to the results of the published

revisions. Moving the taxonomic inventory into many locations on mainland SE Asia,

and encouraging sustained local efforts at continuing documentation (the Flora is not

an end-point), is not devoid of challenges.

The present treatment of Apocynaceae for Fasc. 33 (38 genera, 119 native

spp.) revises only two subfamilies, Rauvolfioideae and Apocynoideae, and does

not cover the other three subfamilies which have been traditionally included in the

Asclepiadaceae. No genus is overly represented, with the largest mostly 6-10 spp., but

more than half of the species are native climbing species, including some big lianas.

Seventeen species are endemic to the region, most of them in Vietnam. Another 12

commonly cultivated species are also identified. In all, this Apocynaceae treatment

includes 1 09 species for Vietnam, 64 species for Laos, and 49 species for Cambodia.

In Fasc. 34, Polygalaceae (5 genera, 43 spp.) includes the cosmopolitan Polygala

(over 500 spp., 22 spp. in CLV), the largest genus in this family of c. 1000 species.

Nine species are endemic to the region, again mostly in Vietnam and especially in

Xanthophyllum. There are 34 species known in Vietnam, 28 species in Laos, and 16

species in Cambodia. The life fomi coverage is interesting; besides herbs, slinibs, trees

and lianas, there is also the diminutive myco-heterotrophic Epirixanthes elongata,

which is an easily overlooked inliabitant of the forest floor but otherwise a widespread

taxon ranging from E India to S China and Moluccas.

Fasc. 35 presents the Solanaceae (8 genera, 48 spp. including Solanum

camranhense Dy Phon & Hul newly published in the revision) and includes Solanum,

the largest genus in the family (c. 1500 spp., 34 spp. in CLV). In CLV all Capsicum,

Cestrum, Datura, Lycium, Nicotiana and Physalis species, along with numerous

Solanum species, are introduced from the New World: they include weeds as well as

major and minor crops. Only five species are apparently endemic to the region. There

are 41 species known in Vietnam, 29 species in Laos, and 23 species in Cambodia.

Standard inclusions for all three revisions are a concise introduction with notes

on taxonomy, diversity, moiphology, palynology, karyology and pertinent biological

aspects as known, uses, keys (to genera, and also to species) and detailed descriptions

of all taxa, with notes on ecology and distribution. Citations of specimens studied are

provided. The descriptions in Polygalaceae and Solanaceae are of about maximum

length compared to past revisions in the FCLV, but the Apocynaceae treatment includes

some lengthy descriptions in a number of cases. For Solanaceae, good notes are given

on the local use of numerous species and at the end of the volume there is an index

of vernacular names in different native languages. There are a good number of rather

exquisite line illustrations in the Apocynaceae (48 by Alain Jouy) and Solanaceae (25

by Dominique Storez and Alain Jouy) issues, but far fewer for Polygalaceae (only five

by Holly Somerville); the inclusion of colour images of species as plates in the final

section of each issue is helpful (8 plates for 38 spp. in Apocynaceae, 4 for 8 spp. in

Polygalaceae, 7 for 16 spp. in Solanaceae). The presentation of genera is alphabetical

414

Card. Bull. Singapore 67(2) 2015

for Apocynaceae but not so for the other two revisions. Printing quality is excellent but

the text font style tends to deliver a somewhat faint print.

Overall, these have been excellently produced. The FCLV is well off on its

second leg.

References

Merrill, E.D. (1935). A commentary on Loureiro's 'Flora Cochinchinensis'. Trans. American

Philos. Soc., n.s., 24(2): 1^45.

Regalado, Jr., J.C., Nguyen Tien Hiep, Phan Ke Loc, Averyanov, L. & Harder, D.K. (2005).

New insights into the diversity of the Flora of Vietnam. In: Friis, 1. & Balslev, H. (eds)

Plant Diversity and Complexity Patterns: Local, Regional, and Global. Proceedings of

an international symposium held at the Royal Danish Academy of Sciences and Letters,

Copenhagen, Demnark, 25-28 May, 2003. Det Kongelige Danske Videnskabernes

Selskab, Denmark.

K.M. Wong & M. Rodda

Singapore Botanic Gardens

The Gardens’ Bulletin

Singapore

VOL. 67 2015

ISSN 0374-7859

VOLUME 67

No. 1

No. 2

pages 1-252 (printed version published 4 June 2015)

pages 253^14 (printed version published 23 December 2015)

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 David J. Middleton Dr Jana Leong-Skomickova

(Editor-in-Chief) (Managing Editor)

Serena Eee & Felicia Tay Yee Wen Eow Cliristina Soh

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

Editorial Advisory Board

Professor Sir Peter Crane

Yale University

U.S.A.

Dr Rogier P.J. de Kok

Royal Botanic Gardens, Kew

U.K.

Dr W. John Kress

National Museum of Natural History

Smithsonian Institution

U.S.A.

Dr Mark Hughes

Royal Botanic Garden Edinburgh

U.K.

Dr Kiat W. Tan

Gardens By The Bay

Singapore

Dr Nigel P. Taylor

Singapore Botanic Gardens

National Parks Board, Singapore

Dr Ian M. Turner

Royal Botanic Gardens Kew

U.K.

Dr Jan-Frits Veldkamp

National Biodiversity Center

The Netherlands

DrJun Wen

National Museum of Natural History

Smithsonian Institution

U.S.A.

Professor Nianhe Xia

South China Institute of Botany

PR. China

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

is Singapore $100,00 including postage. Overseas subscribers should make payment in the form of

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

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

Botanic Gardens, 1 Cluny Road, Singapore 259569”.

National Parks Board

Singapore Botanic Gardens

1 Cluny Road

Singapore 259569

Printed by Oxford Graphic Printers Pte Ltd

CONTENTS OF VOLUME 67 (2015)

VOL. 67(1), June 2015

J. Leong-Skornickova & P.C. Boyce

Hanguana in Singapore demystified: an overview

with descriptions of three new species and a new record 1

M. Rodda, D. Lee & S. Mishak

Dischidia acutifoUa (Apocynaceae, Asclepiadoideae)

a new record for the Singapore flora 29

I.M. Turner

Vxom Ancistwcladus to Tristaniopsis via Tetramerista

- the convoluted history of a Wallich collection and its impact

on the native flora of Singapore 35

S. Lindsay & D.J. Middleton

The clarification and synonymisation of two taxa of Vittaria

from Peninsular Malaysia and a new combination

in Haplopteris (Pteridaceae subfam. Vittarioideae) 39

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

Bothriospermum (Boraginaceae), a new generic record

for the Flora of Peninsular Malaysia 45

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

Novitates Bruneienses, 1. A background to the botanical survey of

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

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

Novitates Bruneienses, 2. A remarkable new species of Begonia

sect. Petermannia (Begoniaceae) from Brunei Darussalam 61

K. M. Wong & Y.W. Low

Novitates Bruneienses, 3. Eight new woody plants

in the Brunei flora, including five new species 69

S. Rahayu, U. Meve & M. Rodda

Hoya imdulata (Apocynaceae, Asclepiadoideae), a new myrmecophytic

species from Borneo, and typification ofH. darwimi 85

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

Begonia bimaensis, a new species of Begonia

from Sumbawa Island, Indonesia 95

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

A new species of Paraboea (Gesneriaceae) from Thailand 101

H. Kurzweil & Saw Lwin

New orchid records for Myanmar, including

the first record of the genus Stereosandra 107

M.F. Newman

Anew species of Zingiber (Zingiberaceae) from Lao RD.R 123

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

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

and the synonymy of Z menghaiense and Z. stipitatum 129

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

Pennilabium labanyaeanum (Orchidaceae),

a new species from Meghalaya, Northeast India 143

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

Tripogon bimiicronatus (Poaceae: Chloridoideae: Tripogoninae),

a new species from India 151

K. Bransgrove & D.J. Middleton

A revision of Epithema (Gesneriaceae) 159

B.R. Maslin

Synoptic overview of Acacia sensu lato (Leguminosae: Mimosoideae)

in East and Southeast Asia 231

BOOK REVIEW

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

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

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

& A. Rasiya Beegam. 2014. (J. Leong-Skornickova) 251

VOL. 67 (2), December 2015

R.P. J. de Kok

Cryptocarya nitens (Lauraceae), a new species record for Singapore 253

Y.Y. Sam

Boesenbergia basispicata (Zingiberaceae), a new record for Peninsular Malaysia 261

K.M. Wong

Schizostachyum kuisingii, a new species of bamboo

(Poaceae: Bambusoideae) from Peninsular Malaysia 267

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

Novitates Bruneienses, 4. New records in the Araliaceae, Araucariaceae,

Arecaceae, Fagaceae, Musaceae and Thymelaeaceae

275

T.L. Yao

Three new species of Loxocarpus (Gesneriaceae) from Sarawak, Borneo

289

W.H. Ardi & D.C. Thomas

Studies on Begonia (Begoniaceae) of the Moluccas II:

A new species from Seram, Indonesia 297

M. Rodda

A new typification of Hoya zollingeriana (Apocynaceae, Asclepiadoideae) 305

R.P.J. de Kok

A revision of Cryptocaiya (Lauraceae) from Thailand and Indochina 309

H. T. Lifu, J. Leong-Skornickova, L.X.B. Nguyin, C.T. DO & T.T. Hoang

Newmania sessilanthera (Zingiberaceae): a new species from Vietnam 351

P.S. Ashton

A few last words on Far Eastern Dipterocarps 357

D.M. Johnson & N.A. Murray

A contribution to the systematics of Xylopia (Annonaceae) in Southeast Asia 361

K.M. Wong

Studies in Southeast Asian Melastoma (Melastomataceae), 1 .

Morphological variation in Melastoma malabathricum and notes

on rheophytic taxa and interspecific hybridisation in the genus 387

I. M. Turner

Clarifying the nomenclature of Crateva trifoliata (Capparaceae) 403

BOOK REVIEWS

A Guide to Begonias of Borneo. Ruth Kiew, Julia Sang, Rimi Repin

& Joffre Ah Ahmad. 2015. (D.C. Thomas) 407

Forest Trees of Southern Thailand, Volume 1 (A-Es). Simon Gardner, Pindar

Sidisunthom & Kongkanda Chayamarit. 2015. (D.J. Middleton) 409

Flora of Cambodia, Laos and Vietnam

Fascicle 33 Apocynaceae. David J. Middleton. 2014.

Fascicle 34 Polygalaceae. Colin A. Pendry. 2014.

Fascicle 35 Solanaceae, Sovanmoly Hul & Pauline Dy Phon. 2014.

(K.M. Wong & M. Rodda) 411