Sandakania August 1998 - Malaysian Agricultural Repository

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SANDAICANIA

A journal of plant systematics, morphology and natural history

published by the Forest Research Centre, Sandakan, Sabah, Malaysia ISSN 0128-5939

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SANDAKANIA

A journal of plant systematics, morphology and natural history

with two issues a year

published by the Forest Research Centre, Sandakan, Sabah, Malaysia with the support of the

Science and Technology Unit, Chief Minister's Department, Sabah, Malaysia

Editorial Committee

I Chairman, Robert C. Ong Editor, K.M. Wong Co-editor, L.G. Saw Assistant editors, Joan T. Pereira

John B. Sugau Richard C.K. Chung Production Manager, C.L. Chan

Advisors

P.S. Ashton {Harvard University, U.S.A.) J. Dransfield {Royal Botanic Gardens, Kew, U.K.) A. Latiff {Universiti Kebangsaan Malaysia, Malaysia)

A.L. Lim {University of Malaya, Malaysia) U. Maschwitz {J.W. Goethe University, Germany) Mohamed Abdul Majid, Haji {University of Malaya, Malaysia)

C. Puff {University of Vienna, Austria)

Communications addresses

Robert C. Ong, Forest Research Centre, Forestry Department, P.O. Box 1407, 90715 Sandakan, Sabah, Malaysia (Fax 6089-531068, email [email protected]).

K.M. Wong, Institute of Biological Sciences, University of Malaya, 50603 Kuala Lumpur, Malaysia (Fax 603-7594178, email [email protected]).

L.G. Saw, Forest Research Institute Malaysia, Kepong, 52109 Kuala Lumpur, Malaysia (Fax 603- 6365531, email [email protected]).

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SANDAKANIA No. 11 August, 1998

Taxonomic notes on the Aquifoliaceae of Sabah and Sarawak

Susyn Andrews

Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, U.K.

Summary. Thirteen new species and two new \arieties of lIex from Sabah and Sarawak are described. lIex nervufosa (Loes.) S. Andre\\" is recognised at a new rank, while six other ]fex species (I. beccarian(J Loes., I. borneellsis Loes., I. CYIIlOS(J Blume. I. gfmnerata King, I.

/ul\'lwndii Loes .. I. lna/accensis Loes.) are lectotypified. The nomenclature and taxonomy of I. havilandii Loes. is also discussed.

In the course of studying the Aquifoliaceac for the Tree Flora of Sabah and Saravvak Project, several new taxa have been recognised. These include three new species and a variety from Sabah, nine new species and a variety from Sarawak and one new species common to both states. Keys to the new taxa will be puhlished in a forthcoming volume of the Tree Flora of Sahah and Sarawak.

THE NEW TAXA

Hex apiensis S. Andrews sp. nov. (J ceteris speciehlls epiphyticis Homeensihlls fotiis sessifihlls. in(ructescentiis pferwnque c(lllfif7oris spicLlTis. fmctihlls lJlI/JCSCClIlihlls dift'r>rl.

TVPllS: /\lIdcrsol1 S.2987R. Sur(/\\'(lk, Gunong Api i1lOfotrjlus SAR.' i.lolrl'i A. E. K'. L'. SING!).

Epiphyte. Twigs peeling, somewhat glossy, reliliish tan to redliish gre) when dried, glabruus.

Leaves alternate, coriaceous, glabrous and dull brown when dried, punctate below, oblong to broadly oblong. 10-15 x 6-8 cm. base rounded or cordate. margin entire, apex acuminate:

midrib glabrous, slightly raised above, prominent below, lateral veins 6-11, subopposite to alternate, fairly widely spaced, more or less paralleL ascending and looping before the leaf margin. suhobscure above. prominent below, intercostal veins hroadly reticulate, ohscure above and helow; sessile. lntlorescences not knO\vn. Infructescences axillary or caulillorous, spicate. 1-2 cm long, glabrous. Fruits (immature) globose, pubescent. 4.5 mm in diameter, green ripening to red/purple, fruiting pedicel verrucose, pubescent. 4 mm long: pyrencs 5, 3-3.5 x 1.3-1.5 mm, glossy. rounded and lineale. (Fig. :2 L~N)

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DISTRIBUTION. Endemic to Borneo. so far only known from the type specimen and one other, both found on Gunong Api, Sarawak.

HABITAT. Primary forest on limestone mountain~ in pockets of organic soil around limestone bouldcrs. 680-900 m altitude. Epiphyte, recorded growing in the crowns of Hopea ilndt'rsonii and Pa/wjllillm sp.

SPECIMENS EXAMINED~BORNEO. SARAWAK: NE. flank of Gunong Api. Ulu Melinau, Anderson S.29878 (A'. E. K!. L!, SAR!, SING!) fruiting specimen; Anderson S.30848 (A!, E!, K!. L!. SAR!. SING!) fruiting specimen.

Named after Gunung Api. this species was previously identified as /lex. sp. lO in many herbaria.

Hex celehensis Capit. var. ranauana S. Andre\\'\ var. nov. a varietate tvpico inj70rescentiis hispidis l)(Ifilis quam pubescentibus, (/((jIlC distri/Jutione geographica (Sumatrwl1 BorneollelllCflle incofit. neque Sulmresi nee Insl/los MO/llccanas nee Sundaims Minores)

d~ffert. TYPIlS: Clemens & Clemens 29245. Sabal/. MOllnt Kinabalu, Kundasang (/lOlotVPllS SING! isotvpi BM", BO!, G!, K', L/, NY!).

lIex sp. an. Ivighfiallo Wall. ex Wight. in Andrew~. Sandakania 5 (1994) 49.

Tree to 18 m tall. 60 cm in diameter. Bark smooth, dark green. Twigs mid-dark grey when dried, glabrous. new growth shortly pubescent. Leaves alternate. semi-chartaceous, glabrous, slightly punctate below. elliptic or lanceolate, 6-9.2 x 2-3.4 cm; base acute. margin entire, apex caudate and obtuse; midrib glabrous and sunken above. prominent with scattered hairs below. lateral veins 6-9. subopposite to alternate. irregularly spaced. more or less parallel, ascending and looping before leaf margin. obscure to subprominent above and below.

interc()stal veins broadly reticulate. obscure ahove and below; petiole )-8 mm long.

puberulous. Intlorescences axillary, in fascicles or few-flowered cymes. hispid or pubescent.

Calyx hispid. 4-6-merous, strongly ciliate. Corolla glabrous. 4-6-merous. ciliate. white.

Stamens (md staminodes 4-6. 2 mm long. Ovary globose. I mm long. pubescent. stigma capitate to columnar. pubescent. Fruits globose, hispid when young. chartaceous. 4--4.5 mm in diameter. brownish-red, fruiting pedicel pubescent. 7-8 mm long; pyrenes 5-7. 2.8-3.5 x

1.5-2 111m, sulcate. (Fig. 3K-M)

DISTRIBUTION. Occurring 111 Sumatra and Borneo, known only from three localities in Sabah.

HABITAT. Found in primary lower montane forest, 1280-1530 m altitude.

SPECIMENS EXAMINED~SUMATRA.East Coast, Yates 1510 (K!) male t1owers; Mount Sago, near Pajakumbuh, Meijer 5116 (L!) female flowers and fruit; loco cit. Meijer 5175 (BM!,

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L!) female flowers; loco cit. Meijer 5891 (L!) flowering specimen; loc. cit. Majadjo 190 (U):

Camp 3, Gn. Bandahara. c. 25 km NNW of Kutatjare. de H'ilde & de Wilde-Du,jies 13020 (BO!, K!, L'J fruiting specimen; Camp I, loc. cit.. de vliilde & de Wilde-Du"}iies 15047 (BO'.

K!, U) male f1owers: Air Sirah on watershed abovc Padang. Barisan Range. de Vogel &

Vermeulen 7609 (BO!. K!. L!) female flowers. BOR"IEO. SABAH: Kundasang. Mount Kinabalu, Clemens & Clemens 29245 (BM!, BO!,G!, K'. L'. NY'. SING') fruiting specimen:

Tenompok, Mount Kinabalu, Clemens & Clemens 28808 (EM!. BO!. G!. K!. L!, NY!) fcmale flowers and young fruit; loe. cit., Clemens & Clemens 28808A (EO!. G!. K!, L!, NY') female flowers and young fruit; Sosopodon below Kinabalu, near Kundasang. Miki! SAN 46711 (K!,

U, SAN!) fruiting specimen; lalan Lering. Sosopodon. Meijer SAN 42476 (A!. K!. SAN!)

immature inflorescences; loco cit.. Meijer SAN 42466 (A'. K!. SAN!) immature inflorescences; loco cit., Gibot SAN 60625 (K'. L!, SAN!, SNP!J fruiting specimen.

This taxon has formerly been confused with I. H'ightiana Wall. ex Wight from S. India and Sri Lanka. but it differs in the glabrous corolla and the presence of a mesocarp in the fruit. Variety celebensis is native to Sulawesi. the Moluccas and the Lesser Sunda Islands, where it occurs on calcareous soils and montane forest. 1900-2400 m altitude.

Hex c1ethritlora S. Andrnl's sp. nov. a ceteris taxis sl'icllli/7oris in{/orescentiis 1i1llSClIlis

longissimis. llsque 12 em IOf/gitudine differt. Tvpu.l: Wood SAN A 4188. Sabah. MoulIl Silalll (hoZotypus KEP' isotvpi A. BO. K, L. SING).

Virgate shrub to 4 m high. Bark greyish-whitc. Twigs silver-grey, smooth. new growth shortly pubescent. Leaves alternate. thick coriaceous, glabrous. punctate below. orbicular to elliptic.

4-10.5 x 3-6.6 cm, base oblique or obtuse. margin entire and translucent when dried. apex mucronate or sharply acute; midrib glabrous. sunken above, prominent below. lateral veins 3-6, subopposite to alternate, fairly widely spaced, more or less parallcl. ascending and looping before the leaf margin, obscure abovc, prominent below, intercostal veins broadly reticulate, obscure above, semi-obscure below; petiole 2-4(-1 ()) mm long. glabrous. Male inflorescences ax i llary, spicate, to 12 cm long. pubescent. female inflorescences Cl\illary.

compressed spicate, to 1 cm long, pubescent; bracts adpressed pubescent. acute. persistent.

Calyx glabrous, tuberculate and glandular, 45-mcrous. ciliate. Corolla papillate inside. 4-5

merous, eciliate, male flowcrs white, females white becoming purplish-white, unscented:

stamens and staminodes 4-5, I mm long. Ovary oblong, 1.4 mm long. purplish. stigma discoid. Fruits (immaturc) globose, glabrous, 3 m111 in diameter, reddish in colour, fruiting pedicel pubescent, 2-3 mm long; pyrenes 4-5. 3 x 1.5 mm, sulcate. (Fig. 3A. B) DISTRIBUTION. Endemic to Bornco, so far only found from one locality in Sabah.

HABITAT. Montane forest on ultramafic ridges. 750-880 m altitude.

SPECIMENS EXAMINED-BORNEO. SABAH: Bukit Silam, Lahad Datu, Wood SAN A 4188 (A, BO, K, KEP!, L, SING) fruiting specimen; Langgi SAN 67507 (L!, SAN!) fruiting

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specimen; Andrews 806 (K!, SAN!) male flowers; Andrel\'s 810 (K!, SAN') male flowers;

Andrews 811 (K!, SAN') female flowers.

The long male inflorescences resemble ClefiJm racemes. which has been the subject of an in

house joke among members of the Kew Southeast Asian team for several years. This taxon was previously identified as !lex sp. 3 in many herbaria.

Hex fruticosa S. Alldrn,'s sp. nov. ({ ceteris !licibus j}(lITif()liis Borneellsifms rwnunculis IClIlIilms fuscis ../i)/iis Imll([tis apice obtusc ({cumillatis difjl'rl. J)jJlIS: Pail' S.2657I. Samwak.

Bukit S({long. Ulu Sampumu. Melinau (holotYPlis SAR! isot.\pi K'. L!).

Shrub to 2.5 m. occasionally epiphytic. Twigs thin. dark coloured. pubescent. Leaves alternate, coriaceous. glabrous. bullate, punctate below. elliptic-mate. 3--1-.2 x 1-1.6 ern. base cuneate.

margin entire. apex blunt acuminate: midrib glabrous. sunken above. prominent below. lateral veins 3-6. subopposite to alternate. widely spaced. ascending and obscure befort' leaf margin, obscure above. subprominent below. intercostal veins obscure above. subobscure below:

petioles 2 111111 long. pubescent. Inflorescences not known. lnfructescences spicate. to 1.2 crn long, pubescent. Fruits (immature) globose. 2-2.2 mm in diameter. somewhat wrinkled when dried, fruiting pedicel 2 mrn long. pubescent: pyrenes 4.2-2'-+ x 1.2-1.4 mm. smooth.

(Fig. 3 E-G) DISTRIBUTION. Endemic to Borneo. known only from two localities in Sarawak.

HABITAT. Found in montane forest and on edge of Kakus sandstone scarp. 950-1380 m altitude.

SPECIMENS EXAMINED-BORNEO. SARAWAK: Bukit Salong. Ulu Sampura.

Melinau. Pail' S.26571 (K!. U. SAR!) fruiting specimen: Bukit Lumat. Ulu Arnau, Mujong.

Ashton S.21258 (K!, L!, SAR'. SING) sterile specimen.

This species was previoLlsly identified as lIcx sp. () in many herharia.

Hex ijuensis S. AfldreH's sp. nov. species (,l1dcll1ica. a ceteris speciehus Sarmmkensihus ramuncutis pal/idis decorticwltihus atque desquul/lIIlltihus,foliis gmndilms 14-15 x 4.5-7 em nitidis o/Jlongis epunctatis. infj'ucteseentiis pu!Jeseentibus differt. TvplIs: Kudi S.23783.

Sarawak. Bukil1ju (holotVjJllS SAR! isotypi A!. BO'. CCE. K.'. L.'. MEL. MOSC. SAN!. SING).

Epiphyte. Twigs flaking and peeling. glabrous. pale coloured when dried. Leaves alternate, coriaeeous, glabroLls. glossy above. epuI1ctate below. drying mid brown. broadly oblong, 14-15 x 4.5-7 Clll. base shortly attenuate. margin entire. apex acuminate: midrib glabrous.

deeply sunken above. prominent below. lateral veins 5-8. subopposite to alternate. widely spaced, more or less parallel, ascending and looping before the leaf margin. obscure above.

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prominent below. intercostal veins broadly reticulate. obscure above and below: petiole 3 mm long. glahrous. Inflorescences not known. Infructescences axillary. ~picate. up to :2 mm long.

pubescent: hracts pubescent. acute. persistant. Fruits (immature) ovoid. 4 mm in diameter.

bright yellow, fruiting pedicel pubescent. 1-2 mm long: pyrenes 4-). :; x 1.3 mm. glossy and

deeply sulcate. (Fig. IE. F)

DISTRIBUTION. Endemic to Borneo. only known from the type locality in Sarawak.

HABITAT. Found in primary dipterocarp forest. at 180 m altitude.

SPECIMENS EXAMINED-BORNEO. SARAWAK: Bukit lju. Ulu Sg. Arip. Balingian.

Kwli S.23783 (A'. BO!, CGE. K!. L!. MEL. MOSC. SAN!. SAR!. SING) fruiting specimen.

Named after Bukit Iju. this taxon was found growing on the bole of a huge tree. where the fleshy leaves and bright yellow fruit were particularly noted. In many herbaria this species was prev iously identified as l/c.\ sp. 9.

Hex jacobsii S. Andre\\'s sp. nov. a ceteris seansorihu.1 Borneensibus rWIIlIllclIlis tellllihus .lil.leis, foWs !) 16.5 x 3.7-7 em subcoriaceis epullc/a!is, prrenis lacrillli!iJ/'/lIihus differt.

Tvplls: Jacohs 5057. SW'!II\'(/k. Mount Penrissol/ (I/Ii!otYjJIIS SAR.' i.I(!/lpi B. G!. K.'. Lt. US).

!lex sp. 8 in Andrews. ;\yuifoliaceae in Checklis! F!O\rerillg P/(/I/!.I & GrlllIlOSperll/\ BI'II/lei f){/rl/ssalam (1996) 31.

Epiphytic climher. rarely a small tree to 7 m. Twigs slender. dark coloured. drying glossy reddish brown to black. lenticellate. glabrous. Leaves alternate. scmi-coriaceous. drying dark brown above. dull mid-brown helow, glabrous. epullctate. elliptic-oblong to broadly elliptic

oblong. 9-16.5 x 3.7-7 cm. base shortly attenuate. margin entire. apex long-acuminate: midrib glabrous, sunken above. prominent below. lateral veins 5-8. subopposite to alternate, widely spaced, more or less parallel, ascending and looping before the leaf margin. semi-prominent above, prominent below. intercostal veins broadly reticulate. obscure above. prominent below:

petiole 4-7 mm long. glabrous. Male inflorescences not known. Female int10resccnces axillary. spicate, 1-2 cm long. glabrous. Calyx glabrous. lobes )-6. eciliate. Corolla )-6

merous. eciliate, greenish-white to white: stamil10des 5-6, 1.4 111111 long. Ovary ublong. 1.8 mm long. yellow. stigma columnar. Fruits globose, ellipsoid when young. glabrous. 5 111111 in diameter, glossy red. fruiting pedicel glabrous. 2 cm long: pyrenes 5-6. 3-4 x 1.2-1.5 mm.

smooth and glossy. tear-like. (Fig. 3N. 0)

DISTRIBUTION. Endemic to Borneo. recorded from se\eral localities in Sarawak and une in Brunei.

HABITAT: Mixed lowland forest on shale in river valleys. riverine forest. mixed dipterocarp forest on sandstone substrate. on very steep slopes in primary forest on basalt hillsides.

100-1100 111 altitude.

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SPECIMENS EXAMINED-BORNEO. SARAWAK: N. slopes of Mount Penrisson. S. of Kuching. Jacobs 5057 (B, GI, K!, L!, SARI. US). fruiting specimen; Ulu Mujong. below N.

Semperaja. Vnyong S.21159 (K!, L!. SAN!. SAR!, SING!). fruiting specimen; Bt. Mersing.

Anap, LuanE? S.22425 (A BO!, K!. KEP!, L!, MEL, SAN!, SARL SING!), fruiting specimen;

Bkt. Balu Semakan Cave, Ulu Sempurau. Melinau. Poie S.25824 (A!, K!, L!. SAR!, SING!), fruiting specimen: Bukit Peninjau, Lanjak-Entimau PF.. Lubok Antu District. Chai S.33909 (K!, KEP!. L!, MO, SAN!, SAR!) female tlowers and fruit; Path from Melinau Gorge to Ulu Sg. Melinau. G. Mulu National Park, Paie S.36523 (K!, KEP!. L!, MO. SAN!, SARI) fruiting specimen: Sg. Abang, Linau Balui. Be1aga, Lee S.40006 (KI. KEP!, L!. SAR!) fruiting specimen; Lambir National Park. Miri. Yii et al. S.43164 (K!, KEP!, L!, MO. SAN!. SAR!) fruiting specimen; across the stream from Camp I, Mount Mulu. Primack S.42433 (A!, K!, KEP!. L!, SAN!, SAR!) fruiting specimen. BRUNEI: Temburong River, just above Wong Nguan rapids, Coode & Wong 6500 (BRUN, K!. III young innorescenccs: loe. cit., just upstream from Wong Nguan rapids. Coode & Wong 6574 (BRUN. K!. L!) flowers female.

This species is named after Marius Jacobs (192l)-1983) to whom Flora Malesiana owes so much: he also collected the earliest known specimen of Ihis species in 1958.

Mikil SAN 44302 a fruiting specimen from Trusmadi, above Kaingaran River in Sabah, could possibly be an extension of this species, but more material needs to be seen. It was collected from a 12 m tree with a clear bole of 8 Ill. Duplicates were distributed to K!. KEP. L. SAR!

from SA:--.I!.

Hex kelabitana S. Andrews sp. nov. a ceteris IlICiblls Borneellsihus foliis 9-14.5 X 5-9 cm coriacei.1 fragilihlls plerumqllc integris )'cflation£' cOf/spieua fJlIllctuatiol1('cjlle eius subtus promineflti differt. TvplIs: Awa et al. S.5U466, Sural\'(/k, roule to Pa' VII/or (holotlpus SARi isonpi K!, KEP!, U, MO. SAN).

Tree to 7 m tall. 6 cm in diameter. Twigs hrownish. slightly puherulous. Leaves alternate, glabrous, coriaceous and brittle. prominently punctate below. broadly elliptic or broadly oblong, 9-14.5 x 5-9 cm. base broadly cuneate. margin entire or undulate with a few minute spines. apex retuse: midrib glabrous. deeply sunken ahove. prominent below. lateral veins 7-10. subopposite to alternate. widely spaced. ascending and obscure hefore leaf margin, subprominent above, prominent below. intercostal veins obscure above. subprominent below:

petiole 4-7 mm long, glabrous. Male inflorescences cymosc. pubescent: female inflorescences not seen. Calyx slightly puhescent, 5-6-merous. ciliate. Corolla 5merous. ciliate, white:

stamens 5. 1.5 mm long. Pisti ]Jode rudimentary. Fruits (immature) elliptic. glahrous, 2 mm in diameter, pale green to red. fruiting pedicel pubescent, 2.5 mm long; pyrenes 5-6. 3 x 1.2 mm,

sulcate. (Fig. 3 S-U)

DISTRIBUTION. Endemic to Borneo. Recorded from two localities in Sarawak. Possibly occurring in Kalimantan.

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HABITAT. Found in submontane mossy forest kerangas forest and secondary semi-kerangas forest: 1000-1330 m altitude.

SPECIMENS EXAMINED-BORNEO. SARAWAK: Pa Ukat 3 miles east of Bario.

Kelabit Highlands. Nooteboom & Chai 02172 (L!. SAR!) fruiting specimen: Kelabit Highlands, Bario, Chai S.35361 (K!. L!, MO, SAR!) male tlowers; Route to Pa'Umor, Bario.

Awa et al. S.50466 (KL KEPL L!. MO. SAN. SAR!) fruiting specimen; Summit of Bukit Pagon KeciL B. Pagan Periuk, Limbang, AW(l & Lee S. 47737 (K, KEP. L. MO, SAN. SAR!) fruiting specimen. KALIMANTAN: Long Api. west of Bawan. Krayan. M. Kuto et al. B

8469 (BO!) fruiting specimen: Between Long Bawan and Panado, R. Geesink 8946 (L!) fruiting specimen.

Named after the Kelabit Highlands in Sarawak. This taxon was previously identified as !lex sp. 20 in many herbaria.

Ilex keranjiensis S. Andrews sp. nov. ab I. hypoglauca (Miq.) Loes. petiolis multo brevioribus tenuioribusque et inflorescentiis magis compactis differt. Ambae species in habitationibus similaribus dispositae sunt. Typus: Anderson S.19123. S{/rawak, Tanjong Keranji (holotypus SARl isotvpi K!, LI, SAN', SiNG').

Small tree to 5 m tall, 20 em in diameter. Bark smooth. light grey. Twigs dark grey. puberulent, new growth dark brown when dried. Leaves alternate. coriaceous, glabrous. dull above, faintly punctate and paler below, obovatc. elliptic or obovate-oblong, 5.5-12 x 2-5 cm. base cuneate, margin entire and revolute, apex obtuse and retuse: midrib with scattered hairs and sunken above, glabrous and prominent below, lateral veins 5-8. subopposile to alternate. widely spaced. more or less paralleL ascending and looping hefore the leaf margin, sunken and subprominenl above. prominent below. intercostal veins broadly reticulate, obscure above, subprominent below; petiole 3-8 mm long, pubcrulous. Inflorescences not known.

Infructescences axillary, in lax compound cymes, to 3.5 cm long, pubescent. Fruits (very immature) narrowly ovoid and ribbed, green; fruiting pedicels pubescent, 4 mm long.

(Fig. 3 H-J) DISTRIBUTION. Endemic to Borneo, known only from the type locality of Tanjong Keranji, Sarawak.

HABITAT. Found in Shorea albida peat swamp forest and peat swamp forest.

SPECIMENS EXAMINED-BORNEO. SARAWAK: Tanjong Keranji, lower Saribas. four miles from banks of river. Anderson S.19123 (K!, L', SAN!, SAR!, SING!) fruiting specimen;

Tg. Keranji, Sarabas. Anderson S.26693 (A!, K!, L!, SAR '. SING!) rruiting specimen; S.P. 47, Tg. Kerangi, Pus a, Hamidi S.14216 (SAR!) sterile.

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This taxon was previously identified as !lex sp. 16 pro parte in many herbari<l. It has been mistakenly called 1. sc/erophYl/oides Lues. which is a synonym of 1. H'ol/ic/7ii Hook. f.. a widespread species in S.E. Asia. The latter has white peeling twigs. epunctate leaves and smooth fruits. IIex keranjiensis has also been confused with 1. hvpoglauca (Mig.) Loes. which occurs in similar habitats in Sumatra. Banka and Borneo. but differs in the thick petioles. 1-2 cm long and in the much longer and larger infructescences.

Hex kinabaluensis S. Andrews sp. nov. Ilh 1. remluta Stapf /i)liis planis. inj70rcscentiis fascicul{l{is glahris potius quam cnnosis IJlil7Cru/i.I, atqlle fructilms maiorilms [lYrenis 4 pot ius quam 3 instructis differt. T"pus: Andrews 890, S(//)(Ih, MOllnt Kinahalu (!zolotypus K' isot.'vpi SAN! SNP.').

flex sp. I in Andrews. Sandakwzi(/ 5 (19lJ4) 49.

Virgate shrub to 2.5 m high. Bark greyish-white. Twigs greenish purple. new growth purple.

glabrous. channelled when dried. Leaves alternate. eoriaceoLis. glabrous. punctate below.

elliptic to obovate. 1.5-3 x 0.8-1 cm. base cuneate. margin serrate in upper half. apex mucronate; midrib glabrous. sunken above, prominent below. lateral veins 3-5. subopposite to alternate. fairly widely spaced, more or less paralleL ascending and looping before the leaf margin. obscure above, promincnt below. intercostal veins broadly reticulate. obscure above.

prominent below; petiole 4-5(-10) mm long. winged when dried. glabrous. Inflorescences glabrous, male in fascicles. female in fascicles or solitary. Calyx glabrous. 4-merous. eciliate.

Corolla glabrous. 4-merous. ciliate. white with pInk or deep red on reverse of lobes. faintly scented; stamens and staminodes 4.3.5--+ 111m long. Ovary oblong. 3.2--3.5 lllm long, grecn or purple. stigma capitate. Pislilllllie oblong. 2-2.2 ¹¹¹¹¹¹ long. Fruib oval. glabrous. I cm in diameter, dark red to purple. wrinkled when dried. fruiting pedicel glabrous. 4.5-7 mm long:

pyrenes 4,8-8.5 x 3-3.5 mm, striate. (Fig. 3 C. D)

DISTRIBUTION. Endemic to Borneo. Known only from the Kinabalu area in Sabah.

HABITAT. Found on sunny slopes on ultramafic mcks in open montane Leptospennwn forest.

2130-2970 m altitude.

SPECIMENS EXAMINED-BORNEO. SABAH: Marai Parai. Clemens 32313 (A!, BM!.

BO!, L!, NY!) flowers in bud; Vicinity of Paka Cave. Bogle. Bogle & Meijer 438 (NY!) fruiting specimen; Summit Trail. abme Carsons Camp. Andrcll's 884 (K'. SNP!) fruiting specimen; loe. cit .. Andrews 886 (K!) 2 seedlings. died 1988; Summit Trail. before Chain 225, Andrews 888 (K!) fasciated branch; loco cit.. Andrews 889 (K!, SAN'. SNP!) male flowers;

loco cit., Andrews 890 (K!. SAN!, SNP!) fruiting specimen; Kinabalu. Wong, ChaN & Nais 93

30 (SAN!. SNP!): Beginning of new Mesilau Trail (Layang-Layang end). Andrews 1579 (K', KEP!, SAN!, SNP!) male flowers; loc. cit.. Andrews 1580 (G!, K!. SAN!, SNP!) fruiting specimen; Summit Trail, Andrc\\'s 1585 (K'. KEP!. SAN!, SNP') male flowers; Panar Laban, Amin et al. SAN 117233 (L') fruiting specimen.

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Collections of this taxon have often been confused in herbaria with l. revv/uta Stapf. another small-leaved holly that occurs in the same area. [n the field when sterile. 1. killiliJalllcnsis appears very similar to Rapanc([ das.\phvl/o (Stapfl Mez as the glossy. dark green leaves are often closely set at the ends of branches. Fasciation appears quitc common.

IIex megaphylla S. Andrews sp. nov. spcciesfoliis Rrandibus 19-35 x 5.5-13 em a/iquatenus falcalis cpllnetatis nervis /{[Icmlihus 18-25 disringllituf: Tvpus: Anderson 5.4750. Sam\\'ak,

Gunollg Afii (holotvpus 5iAR' isolvpi A, K.'. L.'. SING.').

Tree to 9 m taIL 45 cm in diameter. Bark smooth. greyish. Twigs thickish. drying black.

lenticellate, glabrous. Leaves alternatc. coriaceous and brittle, glabrous. drying blackish above. brownish below, epunctate. somewhat falcate. broadly oblong or oblong-Ianceolate, 19-35 x 5.5-13 cm, base cuneate or cordate. margin entire. apex acuminate: midrib glabrous.

sunkcn above, prominent below. lateral veins 18-25, subopposite to alternate, more or less parallel. ascending and looping before the margin. subprominent to obscure above. prominent beneath, intercostal veins broadly reticulate, sunken and subprominent above. prominent to subprominent beloVo': petiole 1-1.5 cm long. thick. channelled above. glabrous. Inflorescences axillary in compound cymes: male cymes to 6 cm long. pubescent. female cymes 3-4 cm long and i I11ll1ature in bud. slightly puberulent on pri mary and \econdary pedicels. tertiary pedicels glabrous. Calyx glabrous. 4-5-merous, eciliate. Male corolla 4-S-merous. ciliate. white:

stamens 4-5. 1.3 mm long. Fruits (very immature) ovoid. (Fig. I A) DISTRIBUTION. Endemic to Borneo, only recorded from two localities in Sarawak.

HABITAT. Found in submontane forest on limestone hilb and cliffs. 910-1000 m altitude.

SPECIMENS EXAMINED-BORNEO. SARAWAK: Gunong Api, Baram District.

Anderson S.4750 (A. K'. L'. SAR!, SINe') immature female tlowers and fruits; C. Api limestone, Ulu Melinau, Chai S. 30364 (A, BO!, K!. L'. SAN, SAR!, SING') male flowers:

G. Buda, G. Mulu N.P.• Chai S.39907 (L!, SAR!) malc flowers.

This is namcd megaphyl/a hecause of its large Icaves, possibly the longest in the gelllls flex.

This taxon was formerly identified as flex sr. II in many herbaria and was also previously mistaken for I. grandij"o/i{[ Merr.. which is endcmic to Sumatra.

IIex mesilauensis S. Andrews sp. nov. ah aliis duahus sl'eeiebus opposifijc)/illS Borneensibu.1 jc)/iis subtlfS punclatis differt. Tvpus: Chew & Corner RSNB 4RR5. Saha/i, Mesi/{lu Ril'er

(hO/OflpIIS SAN! isotypi A!. K.', U, SARi, SlNG!).

flex sp. '2 in Andrews. SWldak{[nia 5 (1994) 49.

Tree to 15 m tall. 120 cm in diamcter. Bark fissurred irregularly, with very large and numerous lenticels. Twigs glabrous, dark brown, lenticcls white. Leaves opposite or slightly alternate,

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coriaceous, bullate. glabrous, punctate below, spathulate or obovate, 6-7,5 x 3-5 cm, base attenuate, margin entire. slightly revolute. apex emarginate: midrib glahrous and sunken above, prominent below, lateral veins 6-8, opposite to subopposite, widely spaced, more or less paralleL ascending and occasionally looping before leaf margin, obscure above, subprominent below, intercostal veins broadly reticulate, sunken and obscure above, subprominent below; petiole 7-8 mm long, glabrous. Inflorescences male, immature, spicate to 2 cm long, puberulous; bracts glabrous. acute. Calyx glahrous, 4-merous. ciliate. Corolla 4

merous, strongly ciliate. Female inflorescences not known. Infructescences not known.

(Fig. 2D, E) DISTRIBUTION. Endemic to Borneo, known only from a single collection by the Mesilau River, Ranau, Sabah.

HABITAT. Montane forest, at 1520 m altitude.

SPECIMENS EXAMINED-BORNEO. SABAH: Mesilau River. Chell' & Corner RSNB 4885 (A!, K!, L!, SAN!, SAR!. SING!) immature male int1orescences.

The only known locality of this taxon has now heen built upon. Despite the lack of mature flowering and fruiting materiaL it is thought that l. mesi/allensis is distinct enough to warrant specific rank. It is to be hoped that more material will he located.

I1ex poiensis S. Andrews sp. nov. ab aliis scwlsorihus Borneensihus in omnibus partihus glabris, foliis integris subtus punctatis petiolo lm'l'i 3-4 mm longo nerl'is lateralihus 5--6 late dispositis dUlert. Tvpus: Clemens & Clemens 225X2. Sa rawak, MOlillt Poi (holotypus SAR!:

isotvpi BO!, K!).

Climber to 15m. 60 cm in diameter. glabrous in all its parts. Leaves alternate. coriaceous.

punctate below. broadly oblong. elliptic or hroadly elliptic. 8.5-11 x 4-6 cm, base broadly cuneate, margin entire, apex acute: midrib sunken above. prominent below, lateral veins 5-6, alternate to opposite, widely spaced, more or les~ paralleL ascending and looping before leaf margin, subobscure above, prominent below, intercostal veins broadly reticulate, obscure above, subobscure below: petiole .'1-4 n1l11 long. Inflorescences not known. lnfructescences axillary, spicate to 3.5 cm long; bracts persistent. acute-acuminate. Fruits (unripe) globose.

4-5 mm in diameter, wrinkled when dried, yellowish-green, fruiting pedicel 6-8 mm long;

pyrenes 8-11, 3 x 1.5 mm, centrally sulcate. (Fig. 2 A-C)

DISTRIBUTION. Endemic to Borneo. known only from two localities in Sarawak,

HABITAT. Occurring in primary forest and near summits on tertiary granodiorite, 460-1370 m altitude.

SPECIMENS EXAMINED-BORNEO. SARAWAK: sine loc., Feb.-June 1914, Native Co/lector 2263 (K!) fruiting specimen: near Upper Cave. Mount Poi, Clemens & Clemens

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20365 (BML BO!. K!, SAR') fruiting specimen: Mount Poi. Clemens & Clemens 22582 (BO!, K'. SAR!) fruiting specimen: Gunong Gading, Lundu District. Chai S.18467 (A. BO!, K!, KEP!, L!, MEL, SAN!, SAR!, SING!) fruiting specimen: loco cit .. Chew CWL 584 (A!, K!, SING!) fruiting specimen.

This is named after Mount Poi in the Ist Division of Sarawak. This species was previously identified as /lex sp. 17 in many herbaria.

I1ex promecophylla S. Andrews sp. nov. species j(lliis oblollRis palliciis 10.5-17.51-24) x 4.5-6.5(--10) em illtegris subtus punctatis longepetiolatis petiolis gracilibus 2-3 em longis distinguitw: Typus: Fidilis & Sumbing SAN 1lO918, Sahah, Ulu Sg. Pingas Pil/RC1.l' (holotypus SAN.' isotypi K!, KEP!, U, SAR!).

/lex sp. 5 in Andrews, Aquifoliaceae in Checklist Flowering Plants & Gymnosperms of Brunei Darussalam (1996) 31.

Tree to 25 m tall, 65 cm in diameter. Bark smooth, dark brown, grey. whitish-cream or yellowish-grey, granular. Twigs glabrous, pale coloured. Leaves alternate, coriaceous.

glabrous, pinkish or yellowish-green above and glaucous below when young, mature leaves mid green above. paler and punctate below. oblong, obovate-oblong or broadly obovate

oblong, 10.5-17(-24) x 4.5-6.5(-10) cm. base cuneate. margin entire. apex mucronate or retuse, rarely obtuse; midrib glabrous, sunken above, prominent below. lateral veins 6-8(-13).

subopposite to alternate. more or less parallel, ascending and looping before the margin, obscure above. prominent beneath. intercostal veins hroadly reticulate. obscure above.

subprominent below; petiole 2-3 cm long. glabrous. Male inllorescences in compound cymes.

glabrous on primary pedicels, pubescent on secondary pedicels. Female inflorescences not known. Calyx glabrous, 5-6-merous, ciliate. Corolla 4-6-merous. ciliate, white to greenish

white. stamens 4-6, 3-3.5 mm long. Ovary rudimentary. Fruits globose. 5-9 mm in diameter, wrinkled when dried, red. fruiting pedicels glabrous. rarely puberulous. 5-10 mm long;

pyrenes 4-6, 6-7.5 x 3-4 mm. striate, sulcate. (Fig.2I-K)

DISTRIBUTION. Endemic to Borneo. found 111 several localities l!1 Sabah and one in Sarawak. It also occurs in Brunei.

HABITAT. Found in primary forest on slopes, on sandy, yellowish soil, on black rocky soil and along ridges. 250-620(-1520) m altitude.

SPECIMENS EXAMINED-BORNEO. SABAH: G. Lumaku FR., west of Sapong Est, Tenom, Rundi SAN 43152 (A!, K!, L!, SAN!) immature fruiting specimen; Trusmadi F Res ..

above Ulu Koingaran River, Tambunan, Mikil SAN 41781 (K!) fruiting specimen; Bt.

Tinuntol, Croker Range, Melalap, Ogata 11476 (L!) male tlowers; Ulu Sg. Pingas Pingas, Keningau, Fidilis & Sumhing SAN 110918 (K!. KEP!, L!, SAN!, SAR!) fruiting specimen;

Syt. Benawood, logged area. Sg. MaaduI1. Nabawan. Fidilis & Asik SAN 104342 (SAN!) male

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flowers: East of Sg. Siburan, RP474, Sepulut, Keningau, Leopold Madani ct ill. SAN 119226 (SAN') fruiting specimen. SARAWAK: Lambir National Park. Miri. George 5.40252 (E'. K', KEP!, L!, MO, SAN~, SARU male flowers. BRUNEI: Bukit Teraja. Ashton BRUN 1(BRUN, L!. SAR!. SING!) fruiting specimen: Temburong. Kuala Belabong, Ashton S.3114 (BRUN!):

loc. cit.. Ashton S. I504 (BRUN').

The name promecophvlla is from the Greek words meaning oblong leaves.

Hex renae S. Andrews sp. nov. speciesf(!liis nitidi.1 ol'lltis intcgris 6-12.5 x 4-6.5(-Y.5) em, hrevipetiolatis petiolis cmssis /-2.5 em long is distinguitlll: Typus: Hm'iland & Hose 2011.

Sarawak, Mount Lamhia (holotYJiUS BO! is(mpi BM!. L.').

Shrub or small tree to 5 m tall. 20 cm in diameter. Bark whitish. Twigs dark grey, glabrous.

Leaves alternate, coriaceous. glahrous and glo~sy abO\e. paler and slightly punctate below.

ovate, broadly ovate or oblong-ovate. 6-12.5 x ..J.-6.5(-9.5) cm. base hroadly cuneate, margin entire. apex obtuse. midrib glabrous. sunken ahO\e. promi nent below. lateral veins 5-8, subopposite to alternate. widely spaced. more or less parallel. ascending and looping hefore the leaf margin, sunken and subobscure above. prominent below. intercostal veins broadly reticulate. obscure above. subobscure below: petiole 1-2.5 cm long. thick, glabrous. Male inflorescences immature, axillary, cymose. puherulous. Female inflorescences not known.

Calyx glabrous . ..J.-5-merous. strongly ciliate. Corolla ..J.-5-merous. ciliate. pale yellow. Fruits (unripe) globose, 3-3.5 mm in diameter, green. fruiting pedicel puberulous, 2-3 mm long:

pyrenes 5. 3 x 1 mm. sulcate. (Fig. 2F-H)

DISTRIBUTION. Endemic to Borneo. round in only two localities within the Lambir Hills National Park. Sarawak.

HABITAT. On the summits of hills on kerangas soil and pnmary heath forest on Kakus sandstone, 150-600 m altitude.

SPECIMENS EXAMINED-BORNEO. SARAWAK: Mount Lambia. Hm'il(//u! & Hose 2011 (EM!, BO'. L') male flowers: Lambir Hills FR .. Miri. DUll hill Hj. Bilker S.3022 (K!,

L!. NB. SAR!. SING!) immature int1orescences: Tatai Memu:-,s. Ulu Tiau. Mujong. Balleh. ak

Unyong S.21229 (K!, L', SAR!, SING!) male f1owers: Lambir National Park. George S.40406 (E!. K!, KEP!, L!, SAN!. SAR) male t10wers and immature fruit (K'): Lambir Hill, Lambir National Park. Miri. Yii et ill. S.431 I I (K'. L'. SAR). fruiting specimen: on summmit of Lambir Hills N.P.. Ulu Sg. Lehau. 20th Mile. Miri/Bintulu Road. Pail' & Yt'(! S.3835..J. (8. K!, KEP!, L!, SAN, SAR) fruiting specimen.

This taxon is named after Rena anak George ( 1955-1994), who was the Officer-in-Charge of the Semengoh Botanical Research Centre, Sarawak. It was previously identified as !lex sp. 18 in many herbaria.

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I1ex spicata Blume vw: harmsiana (Loes. ) S. ^Andrewsvar. nov. pf'tiolo j -2( -4) em longo distil1!!,uitlll: potius qualll 7-/2011 ut in vat: .lpicmLl, atq/le Sar(/\I'(/k Bnmeique if/('oli!, Ileljue Sumutrillll, ]ovam, Sul{l\\'esi 11(,1' Tractus Bonl('ci1sis Sabol! Kalilililll!onamque. Tvp/ls: Bc('cari 3223, SaraH'ak, Mount Tiallg Laju, Provo del Batallg Lupar (hulotypus FL-B.' isot)jJi K').

1. harmsiana Laes. Monogr. Aquif()I. I (! 901) 433; MerrilL Bibl. Enum. Bornean PI. (1921) 352; Masamune. Eflllfll. Phaf/. Born. (1942) 415; Beaman & Anderson. Contrib. Univ. Mich.

Herb. 21 (1997) 112.

1. sp. :'spi('(lta Blume in Andrews. Aquifoliaceae ^In Checklist Flowering Plants &

Gnnnosperms

0/

Brunei Darussalam (1996) 30.

Virgate shrub, rarely a tree to 13.5 m tall. often epiphytic or climbing. Bark smooth, white or silver-grey. Twigs glabrous. Lea\'es alternate. cariaccous. glabrous, densely punctate beneath.

elliptic. ovate or oblong-ovate. 5-8.5(-13) x 2.5-4.5(-6) cm; base rotundate. margin entire.

apex acuminate to long-acuminate; midrib glabrous and deeply sllnken above. prominent and glabrous below; lateral veins 5-7(-8). subopposite to alternate. widely spaced. more or less parallel. ascending and looping hefore leaf margin. obscure above. prominent below;

intercostal veins broadly reticulate. sunken and obscure above. subprominent below; petiole 2--4 111m long. glabrous. Inflorescences axillary. spicate and slightly twisted to 3 Clll long.

glabrous. Calyx verrucose. 4-6-merous, ciliate. Corolla glabrous. 4-6-merous, ciliate. creamy white. Stamens 4-6. 0.9 111m long. Staminodes 4-6. 0.3 mmlong. Ovary globose, 0.5 ¹¹¹¹¹¹long but obovate when young. glabrous; stigma capitate. glabrous. Fruits globose. 2.5-5 mm in diameter, reddish ripening to purplish-black, closely wrinkled when dried. fruiting pedicel glabrous, 3-5 mm long; pyrenes 4-6. 1.7 x I mm. centrally sulcate. (Fig. 3 P-R) DISTRIBUTION. Endemic to Romeo; recorded from several localities in Sarawak and also found ill Brunei.

HABITAT. Found in montane and submontane forest and on basalt and limestone cliffs and ridges. 730-2300 111 altitude.

SPECIMENS EXAMINED-BORNEO. SARAWAK: Mount Tiang: Laju. Provo del Batang Lupar. Ueccari 3223 (E'. K I) fruiting specimen; Gunong Api. Baram District. Allderson S.4734 (SAR!); Bt. Mersing. Anap. Luang S.22363 (K!. L. SAR!. SING!) fruiting specimen;

Bt. Pantu, Melinau. Kapit. Paie S.25739 (A!, E!. K!, L!. SAR!) fruiting specimen; Bt. Salong.

U. Sampurau waterfall, Melinau. Paie S.29284 (E!. K!. LI. SAR!) fruiting specimen; loe. ci! ..

Paie S.29288 (K!. SAR!) fruiting specimen; Gunong Murut. Lawas. Pail' 26453 (K!. L!.

SAR I) fruiting specimen: Mount Murud, Kelabit Highlands. Noo/c/JoOIll & Chai 2020 (K!.

KEPi. LI. SAR!) fruiting specimen; foe. cit.. Nootcboom & Chai 2042 (L!) fruiting: specimen;

Gunong: Api. ulu Melinau. Tutoh, Anderso/l S.30927 (A!. KI. L'. SAR!. SING') fruiting specimen: loc. cit., Chai S.30430 (K!, L!, SAR!) fruiting specimen; Bt. Dulit. Ulu Sg. Tinjar, Baram District. Tong S.34l)()() (K!, KEP!, L'. SAN!. SAR!) fruiting specimen; Aru Tukil, Bario. Baram District. Clwi S.35409 (K!, KEPi. L!. SAN!. SARI) fruiting specimen; Gunong

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Api, Gunong Mulu National Park, Baram District, Argent & Jenny 1031 (E~. K~. SAR~)

fruiting specimen; Bt. Sampadai, Ulu Sampirau. Melinau. Ludong S.41174 (K'. KEP~. L!.

SAN!. SAR!) fruiting specimen; Gunong Murud, Yii S.44489 (K!) male flowers; Ulu Sg.

Kayan, Dulit Range, Belaga. Awa & Yii S.46744 (K!. KEP!, SAR!) immature inflorescences;

Bt. Tebunan, Ulu Trusan, Lawas. Lee S.52485 (K'. L!. SAR!) fruiting specimen; MOllnt Murud, summit ridge, Beaman 11468 (K!, UNIMAS). BRUNEI: North ridge Bt. Retak, Temburong District. Wong 757 (A!. K!. SAN!) female flowers; Bt. Belalong, Temburong District. Dransfield 7153 (K!. L!) female f10wers and fruit; Gunong Pagon near helipad.

Temhurong. Coode 7421 (K!. L!) female tlowers.

I1ex nervulosa (Loes.) S. Andrews stat. nov.

1. venulosa Hook. f. var. nervulosa Loes .. Mongr. AquiJ()I. I (1901) 90; Ridley. Flora Malay Pen. (1922) 442; Kiew, Gard. Bull. Sing. 31 (1978) 82; Kiew. Tree Flora Malara 3 (1978) 8;

Turner, lac. cit. 47. I (1995) 131. Typus: Beccari S.fl. IHerb. Beccari Coli. No. 2935], Singapore. Woodlands (holotypus FL-B!; possible isotypi Beccari s.n. [Herb. Beecari CoIl.

No. 2935AJ and Beccari S.n. [Herb. Beceari ColI. No. 2935B] FL-B!).

I. sclerophylloides Loes. forma - Airy Shaw, Kew Bull. 9 (1939) 509.

lIex sp. 4 in Andrews. Aquifoliaceae in Checklisl Flowering Plants & Gymllosperms Brunei Darussalam (1996) 31.

Tree to 38 m high. 60 em in diameter. Bark white. grey or whitish-green with some mottling.

smooth or slightly fissured. somewhat corky. Twigs peeling. glabrous. greyish to mid-brown or tan. Leaves alternate. coriaceous to thickly coriaceous. glahrous. punctate helow. oblong

ovate, elliptic to broadly elliptic, 7.5-15 x 3.5-6.5 cm. hase cuneate, margin entire. apex acute or obtusely acuminate; midrib glahrous, deeply sunken above, prominent below. lateral veins 6-10, subopposite to alternate, fairly widely spaced. more or less parallel. ascending and looping before the leaf margin, obscure above, prominent below. intercostal veins broadly reticulate, obscure above and below; petiole 1-2 cm long. thick. glabrous. Inflorescences axillary in lax compound cymes, glabrous or rarely pubescent. Calyx glabrous. 4-merous, eciliate or rarely ciliate. Corolla male, 4-merous, female 4-7-merous. eciliate, greenish-white, unscented; stamens 4, 2.5 mm long, staminodes not known. Ovary oblong. 1.2-1.-1- mm long.

stigma capitate. Fruits globose, glabrous, 3-3.5 mm in diameter, red. fruiting pedicel glabrous, rarely puberulent, 4-5 mm long; pyrenes 4.3.5-4 mm x 1.5-2 mm, striate. (Fig. I B-D) DISTRIBUTION. Malay Peninsula. Singapore, Sumatra and Borneo. Recorded from several localities in Sarawak and three areas in Sabah. Also found in Brunei and Kalimantan.

HABITAT. Occurring in open heath forest. seasonal swamps, kerangas forest. suh-montane kerangas, rocky beach forest and acid sands; near sea level to 850 m altitude.

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SPECIMENS EXAMINED-SUMATRA. Saboengan. E. Coast. Si Toroes 3738 (A'. K!) female flowers. SINGAPORE. Woodlands. Beccari s.n.

I

Herb. Beccari ColI. No. 2935 J male flowers; Beccari s.n. [Herb. Beccari ColI. No. 2935A] male flowers: Beccari S.Il. [Herb.

Beccari ColI. No. 2935B] male flowers. PENINSULAR MALAYSIA. 28th Mile, Pontian Rd., Johore, Kosterman & Henderson s.n. (SING!) immature inflorescences; Linggiu-Sindora Forest, Bandar, Tenggara, Johore, Lesmy FRI 35947 (K'. KEP!, SINU!) fruiting specimen;

Gunong Banang, Johor, Ridley 11110 (K!. SING!) male flowers: Bukit Pengaram. Ridlev 11111 (K', SING!) immature inflorescences; Summit of G. Panti. E. Panti FR., S. lohore.

Everett FRI 13834 (K!, KEP', L!) female flowers: G. Panti Timor. KOla Tinggi. Cockhurn FRI 7767 (KEP!, L!) in flower; loc.cit., Jumali & Henslett 4240 (SINU') fruiting specimen; top of Ophir, Hullett 749 (K!) female flowers: Malacca, Ridley 3195 (SING!); Menchali FR., Pekan, Pahang, Ngah KEP 67631 (K!, KEP') immature inflorescences; 16th Mile, Kemaman Road, Pahang, Smith KEP 76588 (A!, K!, KEP!) fruiting specimen; Anak Endau, Ulu Sungai, S.

Pahang, Cockburn FRI 11141 (K!, KEP!, L') female flowers: Menchali forest, Pahang, 1.

Carrick 802 (K!) female flowers; 16th Mile, Kcmaman Road. Pahang KEP 76588 (KEP!) fruiti ng speci men; Kledang Saiong, Perak, SYmington 25895 (KEP I). BORNEO. SABAH:

Lumat, Beaufort, Singh SAN 24321 (BO!, K!, L!, SAR!) male flowers; Weston, Amin SAN 102957 (K!, L, SAN!) female flowers; Forest area Brunei Bay, Weston, Amin & Haya SAN 86354 (K!, KEP!. L!, SAN!, SAR', SING!) fruiting material; Lumat Estate, Beaufort, Lantoh SAN 73239 (K, L!, SAN!) fruiting material; Lumat Beaufort, Sadau SAN 50365 (L!. SAN) flowering material; Hutan Simpan Sianggau. Weston, Sigun SAN 127067 (SAN I) female flowers; H.S. Singgangan, Weston, Amin SAN 105163 (KEP!. L!, SAN!) flowering and fruiting material; VlR Sinnggau, Weston, Sipitang, Leopold Madani SAN 111462 (SAN!) immature flowers. SARAWAK: in Kuehing, Haviland & Hose 3616 (K') male flowers; C.

Santubong East, Kuching, Bujang S.13480 (K!, L!, NB. SAR!, SING!) male flowers; Sagan Rouge. Limbang, Brunig S.I 098 (SAR!) female flowers; Mount Stupong. Jantan S.5193 (A!, NY!) fruiting material; Stapok Road, Ac/zm{[(/ 19 (SAR!) ~terile: SempeJi FR., S.6030 (SAR!) sterile: Telok Pandan Besar. Bako N.P., Chai & Paie S.I7298 (A'. BO'. K!. L!. MEL. SAN!, SAR!, SING!) fruiting material; Bako National Park, Brunig S.7705 (SAR!. SING!) female flowers; loco cit., Carrick & Enoch lCn5 (SAR!, SING!) female flowers: loc. cit., Othman el al. S.37160 (K!, KEP!, L, MO, SAN!, SAR!) fruiting material; loco cit., ,)'tone 13438 (SAR!);

lalan Lintang, foe. cit., Paie S.36409 (K!, KEP'. L!. MO. SAN'. SAR!) male tlowers; Telok Krium, loe. cit., Yii S.42203 (K!, KEP!, L!, MO. SAN!) fruiting material; Padang Baut Plateau, Andrews 1573 (G'. K!, SAR!) fruiting material: loc. cit., Andrews 1574 (K'. SAR!) male flowers; Tk. Tajor Forest, loco cit., Carrick & Enoch lC507 (SAR!, SINGl); foe. cit..

Carrick & Enoch lC4111 (SAR!, SING!); foe. cit., Carrick & Enoch 136 (SAR'. SING!); Ulu Koyan, Mount Dulit, Richards 1820 (A!, K!, SINGl) male flowers; just below summit ridge of Bungoh Range, Anderson S.29047 (A!, E!. K!, KEP', Ll. SAR!) fruiting specimen.

BRUNEI: Badas FR., Brunig S.I 061 (SAR!) inflorescences; B ukit Puan, Belait, A.I/llon .1.825 (BRUN!) sterile; Berakas F.R., Berakas, Allderson S.4930 (BRUN'. SAR!) male flowers.

KALIMANTAN: Bulungan, Sg. Sebakis region, Kostermans 9290 (A!, BO!, G'. K'. L', SING!) fruiting material; loc. cit., Kostermans 9262 (BO!. L!, SING); Mount Palimasan. near Tabang on Belajan River. W. Kutei, Kosternwlls 12790 (K!. L'. SING!) male flowers;

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Goenoeng Klam, Hallier B.2442 (BO!, L!) immature inflorescences; Soengai Smittomw, Hallier B.1270 (BO!, L!).

In March 1866, the Italian botanist Odoardo Beccari returned to Singapore after his joint expedition to Sarawak with the Italian Marquis of Doria. The latter returned to Europe, while Beccari spent a week at Woodlands on the lohore Straits (van Steenis-Kruseman 1950). While there, he collected several specimens including a single, unnumbered collection of !lex.

[Beccari numbered only part of his SE Asian collections; those from Singapore, Sulawesi and the Moluccas were unnumbered, as were a certain proportion of the material from the Malay Peninsula, Sumatra, lava, Borneo, the Lesser Sunda Islands and New Guinea. The entire FI-B herbarium comprising some 15,000 sheets of Beccari numbered and unnumbered specimens was eventually given individual sheet numbers by the FI staff. Many of the Beccari s.n. sheets are only represented at Florence (van Steenis 1955).]

This Beccari s.n. collection consists of 3 sheets labelled Herb. Beccari ColI. No. 2935, 2935A and 2935B. Sheet 2935 was the only sheet determined and annotated by Loesener in 1900 as I. venulosa var. nervulosa var. nov. It is dated March 1866, Singapore at Woodlands and in the lower left hand comer is the word Alheretto. Sheet 2935A is labelled "Singapore Bung.

Lemepon, 3/66" and Alheretto. On sheet 2935B is written "Singapore, Bung Lemelon" and "3.

1866". Both the A and B labels are in different unknown handwriting. To date, I have been unable to find out the meaning of "Bung. Lemepon" or "Bung Lemelon", while alhereto is Italian for a grove of trees. In spite of the different labels, all three sheets are from the same collection and consist of male inflorescences.

T. Loesener visited the Herbarium at Florence in March 1900 and annotated many !lex specimens. In 190 I he described the Woodlands material as var. nervulosa of I. venulosa Hook. f., citing only the Beccari s.n. material he had seen in Florence. lIex venulosa is a tree of the warm, broadleaved forests, woods and open thickets of Burma, Meghalaya and Assam in India, Bhutan and Yunnan in SW China. It occurs from 910-2740 m and is a tree or shrub to 8 m. It has conspicuous lenticels on its branches and the 15-22-lateral-nerved, epunctate leaves have a 2-3 cm long drip tip. The inflorescences are in pseudopaniculate compound cymes, while the fruits contain 5-7 pyrenes. !lex lIervulosa, on the other hand, occurs in a very different habitat from near sea level to 850 m and is a tree up to 38 m high. The branches are peeling not lenticellate, and the leaves which are punctate beneath, have only 6-10 lateral veins, while the apices are acute to obtusely acuminate. The inflorescences are in lax compound cymes and the fruits examined to date contain 4 pyrenes. Even Ridley (1922a) was suspicious as he commented that he had seen no specimens from the Malay Peninsula and so sank var. nervulosa into I. venulosa!

While working on the Aquifoliaceae for the Tree Flora of Malaya (1978b), Kiew realised that I. venulosa did not occur within the region. She had examined photographs of the Beccari s.n.

collection, but equated it with I. macrophylla Wall. ex Hook. f., reducing it to synonymy (Kiew 1978a). However, I. macrophylla, one of the few !lex taxa native to Singapore, differs from var. nervulosa in several ways. It is an evergreen tree but can be deciduous in dry weather

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on the Peninsula (Corner 1952), while I. nervulosa is completely evergreen. The older branches of 1. macrop/zrl1a are heavily lenticellate compared to the peeling ones of I.

nervulosa. The leaves are epunctate beneath as opposed to punctate and the female white flowers are 8-IO-merous, as opposed to the 4-7-merous. greenishwhite flowers of I.

nervulosa. The fruits of I. lIl([crophylla are a glossy scarlet with 0-9 sulcate pyrenes compared to the red fruits with 4 striate pyrenes of I. nervlIfosa. The mainly pubescent I. macrop/n//u occurs in the Malay Peninsula, Singapore and Sarawak in primary lowland forest up to 150 ^III altitude. Collections from Thailand, Sumatra and its islands, and Kalimantan require further study before it is known if they too belong to I. macrophrlla.

/lex nerl'u/osa brings the number of /lex taxa native to Singapore up to four, none of which are endemic. /lex cymosa Blume is the most common, being widely distributed throughout S.E.

Asia. However, it is rated as rare in Singapore (Ng & Wee 1994), as is I. 17wcroph.vlla, which is only known elsewhere with certainity, from Sarawak and the Malay Peninsula. /lex maingavi Hook. f. is considered vulnerable in Singapore and is endemic to the Malay Peninsula. As I. ncrl'ulosa has not been collected on Singapore sinee Beccari's time, it should be regarded as extinct in the wild. According to Ian Turner (pers. comm.) Woodlands is now entirely covered with housing and industrial estates with only a I ittle mangrove left along the Straits. At the time when Beecari was collecting, there would still have been some forest. and as can be seen from the ecological notes above, I. nerniiosa is known from a number of forest habitats as well as from seasonal swamps and acid sands

P.W. Richards, one of the participants of the 1932 Oxford University Expedition to Sarawak, collected an unknown holly on Mount Dulit at Ulu Koyan. This was described by Airy Shaw in 1939 as an unnamed form of I. sclcrophv//oidcs Locs., a species which has since been reduced to synonymy under I. wallichii Hook. f, a widespread taxon from Burma through to Indo-China and West Malesia (Andrews. in press). Richards 1820 was collected in white sand forest at c. 850 m, from a tree 24-27 m in height.

Several of the lowland SE Asian taxa have become totally confused, as inconsistent naming has been applied to the different island groups. The result has been a hotchpotch of epithets which has created total chaos within the herbarium covers. Airy Shaw mentioned not only the above taxon. but also linked it with I. altern!jolill (Zoll. & MOl'.) Loes. and l. pleiob}'({c/Ziil/il Loes., two Javanese species which have yet to be tackled satisfactorily!

I.ECTOTYPIFICATIONS Hex beccariana Loes.

Monogr. Aquifol. 1 (1901) 96; Merrill, Bib!. Enum. Bornean PI. (1921) 352; Masamune, Enum. Phan. Born. (1942) 414. Syntypes: Beccari 14~2, Sarawak. Mount Mat(t)ang (BO, FI

B, K); Bcccari 1879, foe. cit. (FI-B, K); Beccari 2056, foe. cit. (BO, FI-B, K). Lectotype, chosen here: Beccari 1482, Sarawak, Mount Mat(t)ang (FI-B); isolectotypes (BO, K).

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Fig. 1. !lex megaphylla. A. Flowering leafy twig. From Anderson S.4750. !lex nervulosa. B. Fruiting leafy twig. C. Immature fruit. D. Detail of lower leaf punctation. From Yii S.42203. !lex ijuensis. ~.

Fruiting leafy twig. F. Immature fruit. From Jugah S.23783. Drawn by Margaret Tebbs.

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Fig, 2. !lex poiensis. A. Fruiting leafy twig. B. Fruit. C. Detail of lower leaf punctation. From Clemens 22582. !lex mesilauensis. D. Flowering leafy twig. E. Detail of lower leaf punctation. From Chew & Comer RSNB 4885. !lex renae. F. Flowering leafy twig. G. Immature fruit. H. Detail of lower leaf punctation. From Haviland & Hose 2011 except G, from Paie & Yeo S.38354. !lex promecophylla. I. Leafy twig. J. Fruit. K.

Detail of lower leaf punctuation. From George S.40252 ex.cept for 1, from Mikil SAN 41781. !lex apiensis.

L. Fruiting leafy twig. M. Fruit. N. Detail of lower leaf punctation. From Anderson S.30848 except for M, from Anderson S. 29878. Drawn by Margaret Tebbs.

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Fig. 3. !lex clethriflora. A. Flowering leaf twig. B. Detail of lower leaf punctation. From Andrews 806. !lex kinabaluensis. C. Fruiting leafy twig. D. Detail of lower leaf punctation. From Andrews 884. !lexfruticosa.

E. Fruiting leafy twig. F. Fruit. G. Detail of lower leaf punctation. From Paie S.26571 . !lex keranjiensis. H.

Fruiting leafy twig. I. Immature fruit. J. Detail of lower leaf punctation. From Anderson S.19123. !lex celebensis var. ranauana. K. Fruiting leafy twig. L. Fruit. M. Detail of lower leaf punctation. !lex jacobsii.

N. Fruiting leafy twig. O. Fruit. From Yii S.43164. !lex spicata var. harmsiana. P. Fruiting leafy twig. Q.

Fruit. R. Detail of lower leaf punctation. From Lee S.52485.!lex kelabitana. S. Leafy twig. T. Immature fruit.

U. Detail of lower leaf punctation. From Dayang S.50466. Drawn by Margaret Tebbs. •

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DISTRIBUTION. Endemic to Borneo.

HABITAT. Not known.

SPECIMENS EXAMINED-BORNEO. SARAWAK: Mount Matang. Beccari 1482 (FI

B!. lectotype chosen here: BO!. K!. isolectotypes) male flowers; lac. cit. BecCliri 187<) (original syntype. FI-B!. K!) male tlowers; lac. cit. Bec(({ri 2056 (original syntype. BO'. FI

B!, K!) fruiting specimen.

!lex beccariana was described in 1901 hased on three numbers collected by Odoardo Beccari (1843-1920) on Mount Matang, Sarawak in 1866; namely Beccari 1482. Beccari 1879 and Beccari 2056. Loesener named his new species in honour of this celebrated Italian botanist and naturalist and stated that he had seen material from B (now destroyed) and at FI-B. K and M.

Beccari 1482 was collected in April 1866 from a male plant. Both the sheets at FI- Band K have plenty of tlowers and were determined by Loesener when he visited Florence in March 1900 and Kew. An inadequate duplicate was found at BO in 19<)5 but this was not seen by Loesener. Bec('uri 1879 and Beccari 2056 were both collected in July 1866 on a subsequent visit to Mount Matang which is not far from Kuching. The fonner is a male-tlowered specimen and was determined by Loesener at FI-B and K, where there arc t\Vo sheets. The latter number contains good fruit on both the FI-B and K sheets. but ~lI1other duplicate at BO is definately inferior. In this case, only the FI-B sheet has been determined hy Loesener.

I have chosen Beccari 1482 at FI-B as the lectotype. as it has a good number of male inflorescences and plenty of mature foliage.

Hex borneensis Loes.

Monogr. Aqu(f()I. I (1901) 248 + tab. 6, fig. I a-c: Merrill. RiM Enum. Bornean PI. (1921) 352; Masamune. Emull. Phan. Born. (1942) 415: Dudley. Fed Rep. 94, 3--4 (1983) 167.

Syntypes: Beccari 1652. Sarawak, Mount Mat(t)ang (FI -B): Beccari 2577, lac. cit. (FI- B. KJ.

Lectotype, chosen here: Beccari 1652, Sarawak. Mount Mat(t)ang (FI-B).

Diospyros sororia Bakh., Bull. Jard. Bot. Buit. 1512 ( 1(37) 125.

DISTRIBUTION. Endemic to Borneo.

HABITAT. Found in primary lowland dipteroearp forest or on hillsides. rarely in secondary forest: 90-1000 m altitude.

SPECIMENS EXAMINED-BORNEO. SARAWAK: N. slopes of Mount Penrissen.

Jacobs 5103 (K!, L!) fruiting specimen: Serian, Sabal F.R., Paie S.16647 (BOI. K!. L!. SAN!,

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SING!) fruiting specimen; Semengoh Arboretum. Penrissen Road, Chai S.37172 (K!. KEP!, U, SARI) sterile; loe. cit. ISIJumi & Ching S.37193 (K'. KEP!. L!, SAR!) fruiting specimen;

loe. cit., Ghazalli S.13659 (K!, U) fruiting specimen; loc. cit., Paie S.335337 (K!) male flowers; Mount Matang. Beccari 1652 (FI-B!, lectotype chosen here) fruiting specimen; lac.

cit., Beccari 2577 (original syntype, PI-B'. K!) male flowers; Gunong Magili Forest Reserve, Sadang. Mead 00046 (BO!' K') sterile: Bukit Senyandang, Lingga, Lee S.44023 (KI. L!) fruiting specimen; Bukit Alak, Tebakang area. A\\'a & Paie S.45619 (K!) fruiting specimen;

Sg. Pesu, Dataran Tinggi Merurong, Othman, Yii et al. S.48909 (L!, SAR!) fruiting specimen.

KALIMANTAN: Mount Liang Gagang. Hallier 3077 (A!, BO!, K!, L') male flowers;

Gllnllng OaYllh, Bentiang, Pontianak, She{/ 26X34 (80!, KI. L') male flowers; Gunong Bentiang area, Burley, Tukirin et al. 2512 (K I) fruiting specimen; loe. cit., Burley. Tukirin et af. 2475 (K!, L!, SING!) fruiting specimen; loc. cit., Burlev, Tukirin et al. 3252 (K!,

u.

SING!) fruiting spccimen; loe. cit., Burl('\'. Tukirin et al. 2988 (K!, U. SING!) fruiting specimen; loc. cit .. Burley, Turkirin er al. 2893 (K!. L!. SING!) sterile: Gunong Jako, Bentiang, Shea 26880 (BO!, K!) fruiting material: Gunong Pao, Bentiang, Shea 27131 (BO!, K!, U) fruiting specimen; de Vriese s.n. (L!); de Vriese s.n. (L!).

flex homeensis was described by Loesener and illustrated by his sister Elisabeth who showed the exceptionally, large pyrenes of this species (Loesener 190I). It was based on two collections made by Beccari on Mount Matang. where in May 1866 he gathered a fruiting specimen Beccari 1652. This unicate is held at FI-B. where it was annotated by Loesener in March 1<)00. It is chosen here as lectotype.

Jn November 1866, Beccari 2577 was collected consisting of immature inflorescences. Of the two sheets at PI-B, only one has a determination by Loesener on the main label, while an unknown hand has added "gros,\'o frutice" to it. The second sheet (Herb. Beccari ColI. No.

2532A) has a main label only with 2577 handwritten on it. At Kew there are two sheets of Beccari 2577 which were not seen by Loesener and neither has original labels.

Hex cymosa Blume

Bijdragen (1827) 1149; Hook. f., Fl. Brit. India I (1875) 605; King, Jour. Asiatic Soc. Beng.

64, 2 (I X95) 136; Loesener. MOI1()Kr. Aquilal. 1 (\ 9(1) 74: Koorders-Schumacher. S-"st.

Verzeich. 1 (1912) 157; Valeton, BUdrage 13,13 (1914) 7; Merrill, Bibl. Enum. Bornean PI.

( 1921) 352; Ridley, Flora Malay Pen. 1 (1922) 442; Masamune, Emu1/. Phan. Born. (1942) 415; Browne, Forest Trees Samwak & Brunei (1955) 200; Meyer. Bot. News Blill. Herh. For.

Dept. <) (1967) 91; Anderson, Trees Pe(/r Swamp Forests Soul\\'ak (1972) 25, plate 8A;

Anderson, Check List Trees Sarawak (1980) 150; Corner, Wayside Trees Malaya 1 (1988) 163;

Keng, Concise, Fl. SinKapore (1990) 114; Turner. Gard. Bull. SinK. 45. 1 (1993) 37; Kessler

& Sidiyasa, Trees Balikpapan-Samarinda Area. East Kalimantan, Indmzesia (1994) 54;

Andrews, Sandakania 5 (1994) 43; Turner, Gard. Bull. Sing. 47, 1 (1995) 131; Andrews, Aquifoliaceae in Checklist Flowering Plants & Gnnnosperms of Brunei Darussalam (1996) 30. Syntypes: Blullle s.n. Herh. Lugh. Bat. No. 904.134.97, Java (L); Blume s.n. Herb. Lugh.

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Bat. No. 904,134,106, loco cit. (L); Blume s.n. Herb. Lugh. Bat. No. 904,134,98, loc. cit. (L);

Blume s.n., Herb. Lugh. Bat. No. 952,63,761, loco cit. (L): Blume s.n., Herb. Lugh. Bat. No.

904,28,89 (L). Lectotype. chosen here: Blume s.n. Java. Herb. Lugh. Bat. No. 904,134,97 (l).

Prinos cymosa Hassk., Tijdschr. Nat. Gesch. & Phys. 10 (1843) 140; Hasskarl, Cat. PI. Hort.

Bog. (1844) 231: MiqueL Fl. Ned. Ind. 1,2 (1859) 595: MiqueL I.c. Suppl. I (1860) 201, 514:

Mueller, Ann. /Jot. srst. 7. 4 (1869) 572.

Prinos cymosa Hassk. var. minor Miq .. Fl. Ned. Ind. Suppl. 1 (1860) 20 I, 514.

Leucodermisjavanica Planch. ex Hook. f.. Fl. Brit. Ind. I (1875) 605.

/lex philippinensis Rolfe, Jour. Linn. Soc. Bot. 21 (1884) 308.

I. ("umingiana Rolfe, Jour. Linn. Soc. Bot. 21 (1884) 308.

I. cymosa Blume var. cumingiana (Rolfe) loes., Monogl: Aquifol. I (1901) 76; Elmer. Leaf7.

Phil. Bot. 5, 38 (1913) 1964: Merrill, Enum. Phil. Plants 2 ( 1923) 478.

DISTRIBUTION. Malay Peninsula, Singapore, Thailand. Sumatra and neighbouring islands, Java, Sulawesi. Borneo and the Phili