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Studies in Malesian Gentianaceae III:
Cyrtophyllum reapplied
to the Fagraea fragrans alliance
K.M. Wong1 and M. Sugumaran2
1Singapore Botanic Gardens,
1 Cluny Road, Singapore 259569
wkm2000@gmail.com
2Rimba Ilmu Botanic Garden,
Institute of Biological Sciences, University of Malaya,
50603 Kuala Lumpur, Malaysia
maran@um.edu.my
ABSTRACT. Cyrtophyllum Reinw., one of several distinct lineages among the Fagraea
complex, is the correct genus to which ve species of Southeast Asian trees should be assigned,
including the widespread F. fragrans. Cyrtophyllum minutiorum K.M.Wong is a new species
described here. Two new combinations are made: C. caudatum (Ridl.) K.M.Wong and C.
wallichianum (Benth.) M.Sugumaran & K.M.Wong.
Keywords. Cyrtophyllum, Fagraea fragrans, Gentianaceae, Malesia, Potalieae, Potaliinae,
Southeast Asia
Introduction
The results of a molecular phylogenetic study of the Fagraea complex (Sugumaran
& Wong 2012) demonstrated the distinctness of a number of generic lineages from
Fagraea Thunb. s.s. (Wong & Sugumaran 2012). Among these, Cyrtophyllum Reinw.
and Picrophloeus Blume were readily distinguished from Fagraea s.s., Limahlania
K.M.Wong & M.Sugumaran and Utania G.Don because the rst two genera have
owers with conspicuously exserted styles (typically more than 40% of their length)
and laments (greater than 70% of their length) (Sugumaran & Wong 2012). Also,
Cyrtophyllum and Picrophloeus frequently have cymes bearing numerous small
owers (corollas narrow, the mouth often not more than 10 mm wide), compared to
the other genera, which typically have larger owers (corollas typically much wider)
in variable numbers. However, Cyrtophyllum has axillary cymes and Aubréville’s tree
architectural model, whereas Picrophloeus and the other three genera all have terminal
cymes and consistently other architectural models (Scarrone’s in Picrophloeus and
Fagraea s.s., Fagerlind’s in Limahlania, Roux’s in Utania) (Sugumaran & Wong
2012; Wong & Sugumaran 2012).
The history of Cyrtophyllum is interesting. Ridley (1893) had used the name
C. fragrans (Roxb.) DC. Later, he described an allied species under Fagraea, but
nevertheless appreciated (Ridley 1918) that this genus name was available for the
alliance of trees that was quite different from Fagraea: “It would probably be best to
497
Gardens’ Bulletin Singapore 64(2): 497–510. 2012
keep up the genus Cyrtophyllum for the Tembusu trees which differ so much from the
epiphytic true Fagraeas with their eshy leaves and owers, and included stamens,
from the tall trees with their leaves and owers and long projecting stamens. The
genus Cyrtophyllum would thus contain C. fragrans..., C. giganteum..., C. wallichii...,
C. caudatum...” In his Flora of the Malay Peninsula (Ridley 1923), he adopted
Cyrtophyllum for this group of species. Burkill (1935), on the other hand, took the
broad view as Blume (1838) did, and, under Fagraea, stated: “Cyrtophyllum... which
some botanists separate, is not really distinct, and is united here.” The uncertainties
have found their way as different taxonomic approaches throughout this period, viz.,
as a section of Fagraea (Blume 1838); as a subgenus of Fagraea (Miquel 1857); and
more recently as Blume’s section again but with more discordant elements thrown in
(Leenhouts 1962; see also Wong & Sugau 1996).
Here we provide a revision of Cyrtophyllum, which includes the well-known
tree species in Southeast Asia, C. fragrans (synonym Fagraea fragrans Roxb.).
Cyrtophyllum Reinw.
in Hornschuch, Syll. Pl. Nov. 2 (1825) 8; Blume, Bijdr. Fl. Ned. Ind. (1826) 1022;
Ridley, Fl. Malay. Pen. 2 (1923) 421; Wong & Sugumaran, Gard. Bull. Sing. 64 (2012)
490. TYPE SPECIES: C. peregrinum Reinw. (= C. fragrans (Roxb.) DC.).
Fagraea sect. Cyrtophyllum (Reinw.) Blume, Rumphia 2 (1838) 34 pro parte excl.
F. speciosum; Leenhouts, Fl. Males. I, 6 (1962) 303 pro parte excl. F. elliptica & F.
umbelliora; Wong & Sugau, Sandakania 8 (1996) 7 pro parte excl. F. elliptica. -
Fagraea sect. Parviorae Benth., J. Linn. Soc. Bot. 1 (1857) 100 pro parte quoad C.
peregrinum. - Fagraea subg. Cyrtophyllum (Reinw.) Miq., Fl. Ind. Bat. 2 (1857) 375
pro parte excl. F. sumatrana.
Medium-sized to large trees usually more than 3 m and some reaching 25-30 m tall.
Trunk monopodial with episodic growth; developing wave-like sympodial branch
systems or ‘terminalian’ branching with successively higher orders of outwardly
directed, then upturned, (indeterminate) branch segments (plagiotropy by apposition)
(Aubréville’s architectural model de Hallé et al. 1978). Trunk bark becoming ssured
in older trees. Vegetative shoot apices with light yellowish resin. Leaf arrangement
on branches decussate; leaf margin entire; petiolar sheaths of a leaf pair fused and
forming a shallow cup-like ochrea that loosely clasps the stem; petiolar base auricles
absent. Inorescence axillary, a few- to many-owered and few- to multi-branched
cyme, basal branches nearly as long as or longer than rachis. Flowers bisexual, small,
up to 10 mm wide at the corolla mouth; calyx lobes 5; corolla white to creamy white,
corolla lobes 5, overlapping to the right; stamens 5, typically two thirds or more
exsert, anthers versatile, sagittate; style typically one third or more exsert; stigma
capitate (knob-like). Fruits small, subglobose to slightly ellipsoid, up to about 10
mm in diameter; colour at maturity yellow-orange to bright red; with small amounts
498 Gard. Bull. Singapore 64(2) 2012
of translucent sticky latex in fruit epidermis and fruit wall; epidermis separating as
a thin translucent lm from pericarp (fruit surface appearing crinkled on herbarium
specimen). Seeds numerous; placentation axile; polygonal; surface areolate.
Distribution and diversity. Bengal, Burma, Thailand, Indo-China, Andaman Islands,
Malay Peninsula, Sumatra, Java, Borneo, Mindoro, Balabac, Palawan, Celebes and
north-west New Guinea. Five species known.
Habitat. Lowland forests.
Key to Cyrtophyllum species
1a. Inorescence branching to just 1, rarely 2, orders. Corolla tubes 10–25 mm long;
stamens with laments 16–38 mm long; styles 14–55 mm long…….............… 2
b. Inorescence branching to 2–4 orders. Corolla tubes not exceeding 10 mm long;
stamens with laments 7–17 mm long; styles 9–22 mm long……….............… 3
2a. Leaf apex caudate. Calyx 2.5–3 mm across, corolla tube subcylindric (not
conspicuously ared in its upper part), 10–12 mm long. (Borneo) ........................
............................................................................................................ C. caudatum
b. Leaf apex acuminate. Calyx 3–4 mm across, corolla tube infundibular (ared in
its upper part), 12–25 mm long. (Malay Peninsula) ................... C. wallichianum
3a. Corolla tube infundibular (ared in the upper half), larger (lower narrowed part of
corolla 1.5–2(–2.5) mm wide); leaf secondary veins at or slightly raised on the
upper leaf surface in dried material .................................................... C. fragrans
b. Corolla tube subcylindric (not noticeably ared in the upper part), smaller (lower
narrowed part of corolla 1–1.5 mm wide); leaf secondary veins at to frequently
sunken on the upper leaf surface in dried material ………....................………. 4
4a. Leaves chartaceous, margin conspicuously wavy (more conspicuously so in
fresh material), secondary veins forking towards the leaf margin but not forming
conspicuous loops; pedicels 3–5 mm long (owers distinctly pedicellate); corolla
tube 7–8 mm long; style 12–22 mm long. (Widespread across Malesia.) ...............
.......................................................................................................... C. giganteum
b. Leaves coriaceous, margin plane, secondary veins forming distinct loops towards
the margin; pedicels 0–1 mm long (owers subsessile); corolla tube 4–5 mm
long; style 9–10 mm long. (Restricted to Borneo.) ...................... C. minutiorum
1. Cyrtophyllum caudatum (Ridl.) K.M.Wong, comb. nov. — Fagraea caudata Ridl.,
J. Str. Br. Roy. As. Soc. 79 (1918) 97; Merr., J. Str. Br. Roy. As. Soc. Sp. No. (1921)
491; Wong & Sugau, Sandakania 8 (1996) 8; Coode et al. Checkl. Fl. Pl. Gymn. Brunei
(1996) 174. TYPE: Lobb s.n., 1853, ‘Borneo’ (Sarawak) (holo K).
499
Cyrtophyllum reapplied to Fagraea fragrans and allies
Fagraea fragrans auctt. non Roxb., Leenhouts, Fl. Males. I, 6 (1962) 304 pro parte,
Ashton, Manual Non-Dipt. Trees Sarawak 2 (1988) 310 pro parte, quoad F. caudata
in syn.
Tree, 6–7 m tall, perhaps bigger; trunk to 2.5 cm diameter or more; bark smooth, dark
brown. Leaves elliptic to slightly oblanceolate; (4–)7–9(–11) cm long, (1.5–)2.5–3.5(–
4) cm wide; base cuneate; apex caudate, 1–1.5 cm long; margin plane when fresh and
in dried specimens; coriaceous; upper and lower surfaces smooth; midrib prominent
below, sunken above; secondary veins 7–9 pairs, upper side faint and immersed in the
blade, lower side faint to very slightly sunken or even prominent; tertiary veins faint
to inconspicuous; petioles 0.8–2.2 cm long, 1.5–2 mm thick, petiolar sheaths of a leaf
pair fused and forming a shallow cup-like ochrea that tightly clasps the stem; petiolar
base auricles absent. Inorescence a few-owered cyme 5–9.5 cm long; peduncle
2.5–6 cm long, 1(–1.5) mm thick; with just a single level (tier) of branching on the
main axis, typically not rebranched. Flower pedicel 5–10 mm long, c. 1 mm thick;
calyx (from the base to the lobe apices) 3.5–4 mm long, glabrous, calyx cup 2.5–3 mm
wide, calyx lobes erect, 1–1.5 mm long, 1–1.5 mm wide, margins glabrous; corolla
tube subcylindric, 10–12 mm long, 1.5–2 mm wide near the base, inside glabrous to
minutely papillate; corolla lobes broad-ovate to subobovate, 5–6 mm long, 4–5 mm
wide; stamens inserted at the uppermost third of the corolla tube; laments 16–18 mm
long, exsert 11–12 mm in the open ower; anthers not seen; style 14–15 mm long,
protruding 11–13 mm from the corolla mouth in the open ower; stigma c. 0.3 mm
across. Infructescence peduncle 4–7.5 cm long, 1–1.5 mm thick. Fruits (immature)
to 5–6 mm across; the base loosely to tightly clasped by the calyx lobes. Seeds not
examined.
Distribution. Endemic to NW Borneo (SW Sabah, Brunei, Sarawak).
Habitat. Hills and ridges in lowland mixed dipterocarp forest.
Specimens examined: BORNEO. Brunei. Belait, Batu Patam, along ridge north of summit,
Wong WKM 1060 (BRUN, K, SING). Sarawak. Beccari 2956 (K); 1st Div. Gunung Santubong
South, Bujang S. 12999 (K, L, SAR, SING); Bako N.P., Lintang path, Chai S. 19707 (A, BO,
K, KEP, L, MEL, MOSC, SAN, SAR, SING), Bukit Gondol, Ilias S. 17908 (A, BO, K, KEP,
L, SAN, SAR, SING), Md. Shah P. 5647 (A, K, L, SAR, SING); Lobb s.n. 1853 (holo K); 1st
Div. Mt. Matang, near Valombrosa, M. & J. Clemens 7783 (K); Mt Mike Low LYW 213 (KLU);
Bintulu, Nyabau F.R., Brunig S. 12050 (SAR); Lambir N.P., Sg. Jangkang, Mokhtar et al. S.
47187 (K, KEP, L, MO, SAN, SAR); Kuching, Ulu Sg. Rayu, Yakup S. 7716 (SAR, SING);
Kelaung F.R., sine coll. S. 7431 (SAR).
Ridley (1918) observed that the species he was describing under Fagraea, when
appropriately transferred to Cyrtophyllum, should be C. caudatum, although that
combination was not effected.
500 Gard. Bull. Singapore 64(2) 2012
2. Cyrtophyllum fragrans (Roxb.) DC., Prod. 9 (1845) 31; Ridley, Trans. Linn. Soc.
Lond. 2nd ser. Bot. 3 (1893) 323. — Fagraea fragrans Roxb. [Hort. Beng. (1814) 84,
nom. nud.], Fl. Ind. ed. Wall., 2 (1824) 32; Don, Gard. Dict. 4 (1837) 68; Blume, Mus.
Bot. 1 (1850) 172; Benth., J. Linn. Soc. Bot. 1 (1857) 100; Miq., Fl. Ind. Bat. 2 (1857)
375; Kurz, For. Fl. Brit. Burm. 2 (1877) 205; Clarke in Hooker f., Fl. Brit. India 4
(1883) 85; Ahern, Compil. Timber Tree Philip. Isl. (1901) 88, pl.; Koord. & Val., Bijdr.
9 (1903) 86; Ridley, J. Str. Br. Roy. As. Soc. 50 (1908) 121; Elm., Lea. Philip. Bot. 2
(1909) 597; Dop, Fl. Gén. I.-C. 4 (1914) 177; Merrill, J. Str. Br. Roy. As. Soc. Sp. No.
(1921) 492; Cammerl., Bull. Jard. Bot. Btzg. III, 5 (1923) 318; Foxw., Mal. For. Rec.
3 (1927) 157; Corner, Wayside Trees Malaya (1940) 424; Kerr in Craib, Fl. Siam. En.
3 (1951) 55; Browne, For. Trees Sar. & Brun. (1955) 244, t. 32; Smythies, Common
Sarawak Trees (1965) 87; Kochummen, Tree Fl. Malaya 2 (1973) 273; Cockburn,
Trees Sabah 1 (1976) 211; Wong & Sugau, Sandakania 8 (1996) 11; Coode et al.
Checkl. Fl. Pl. Gymn. Brunei (1996) 174. — Willughbeia fragrans (Roxb.) Spreng.
Syst. Veg. 4 (1827) Cur. Post. 71. TYPE: Hunter, Wallich Cat. no. 1597E, “Pullo
Penang” (holo K).
Cyrtophyllum peregrinum Reinw. in Hornschuch, Syll. Pl. Nov. 2 (1825) 9; Blume,
Bijdr. Fl. Ned. Ind. (1826) 1022; DC., Prod. 9 (1845) 31; Ridley, Fl. Malay. Pen. 2 (1923)
421; Holttum, Gard. Bull. S.S. 5 (1931) 189. — Fagraea peregrina Blume, Rumphia 2
(1838) 34, t. 80; Mus. Bot. 1 (1850) 172. LECTOTYPE (Wong & Sugumaran 2012):
Reinwardt s.n., Java (L sheet no. 908.127-246 & barcode L0005030).
Fagraea cochinchinensis A. Chev., Cat. Pl. Jard. Bot. Saigon 33 (1919) 66, pro
specim., Merrill, Enum. Philip. Fl. Pl. 3 (1923) 314, pro specim., non basionym Aidia
cochinchinensis Lour., Fl. Cochinch. 1 (1790) 143.
Fagraea fragrans auctt. non Roxb.: King & Gamble, J. As. Soc. Beng. 74 (2) (1908)
611 pro parte, excl. Derry 272; Leenhouts, Fl. Males. I, 6 (1962) 304 pro parte,
Ashton, Manual Non-Dipt. Trees Sarawak 2 (1988) 310 pro parte, excl. F. caudata
Ridl., F. gigantea Ridl., F. speciosa sensu Ridl. non Blume, F. sororia J.J.Sm., F.
wallichiana Benth., F. lanceolata Wall. in syn.; Kochummen, Tree Fl. Malaya 2 (1973)
273 pro parte, excl. Cyrtophyllum lanceolatum, Fagraea wallichiana in syn.; Grifn
& Parnell, Fl. Thailand 6 (3): 198 pro parte, excl. C. giganteum Ridl. in syn.
Tree, rarely to just 3–4 m tall, more often big, to 30 m tall; trunk to over 1 m in
diameter; bark deeply and ruggedly ssured, grey-brown to dark brown. Leaves
elliptic; (5.5–)7.5–11(–13) cm long, (2–)3.5–4.5(–5.3) cm wide; base cuneate; apex
short-cuspidate to caudate; (3–)5–8(–9) mm long; margin plane when fresh (in dried
specimens sometimes slightly wavy); chartaceous to thin-coriaceous; upper and lower
surfaces smooth; midrib prominent below, at to slightly raised above; secondary
veins (7–)9–12 pairs, upper side faint and immersed in the blade, lower side faint to
very slightly prominent; tertiary veins faint to inconspicuous; petioles 1–1.3(–1.6) cm
501
Cyrtophyllum reapplied to Fagraea fragrans and allies
long, 1–1.5 mm thick. Inorescence (3.5–)4–7(–7.7) cm long; peduncle (2.8–)3–3.5
cm long, 1–1.5 mm thick; with (2–)3(–4) levels (tiers) of branching on the main axis,
the branch tiers (2–)3–5(–6) mm apart, lowest tier typically branched to 2(–3) orders.
Flower pedicel (3–)5–8(–11) mm long, 0.5(–1) mm thick; calyx (from the base to
the lobe apices) (1.5–)2–2.5 mm long, glabrous, calyx cup (1.5–)2–2.5(–3) mm wide,
calyx lobes erect, 1–1.5(–2) mm long, 1–1.5(–2) mm wide, margins glabrous; corolla
tube somewhat infundibular, (4–)6–8 mm long, lower narrowed part 1.5–2(–2.5) mm
wide, upper ared part (3–)4–6(–7) mm wide at the top, the lower narrowed tubular
part nearly as long as the upper ared part, inside glabrous to minutely papillate;
corolla lobes broad-ovate to subobovate, (3–)4–6 mm long, 2–3(–4) mm wide;
stamens inserted at the middle of the upper ared part of the corolla tube; laments
(10–)12–16(–17) mm long, exsert (8–)10–12(–13) mm in the open ower; anthers
1–1.5 mm long, 0.5–1 mm wide, each anther sac somewhat narrowly ellipsoid; style
(14–)18–22 mm long, protruding (8–)10–12(–14) mm from the corolla mouth in the
open ower; stigma c. 0.5 mm across, lobes 2, low and rounded, parting to present two
slightly raised, hemispherical, papillate inner surfaces when receptive, not recurving.
Infructescence peduncle (2.5–)3.5–5(–6.2) cm long, 1–2 mm thick. Fruit when
mature to 4–6 mm across; the base loosely to tightly clasped by the calyx lobes. Seeds
0.5–1 mm across.
Distribution. Principally a mainland SE Asia and W Malesian species: Bengal,
Andaman Islands, Burma, Thailand, Indo-China (Laos, Vietnam, Cambodia), Malay
Peninsula, Sumatra, Java, Borneo, SW Philippines (the Palawan chain from Balabac
NW through Palawan, Culion and Busuanga, and continuing into Mindoro); also
Celebes.
Habitat. One of our most commonly encountered trees in West Malesia, Cyrtophyllum
fragrans establishes easily in open areas and secondary forests including along
roadsides. It grows easily on sandy sites, including around sandy tailings of former
tin-mines in the Malay Peninsula. It is less commonly found in lowland high forest,
but easily encountered in kerangas (tropical heath forest) vegetation and coastal or
beach forest.
Specimens examined: BORNEO. Brunei. Belait, Anduki F.R., Anderson S. 4941 (SAR),
Andulau, Taman Rekreasi Hutan Sungai Liang, Sugumaran et al. SM 227 (KLU), Sungai
Liang, Hussain HUS 30 (BRUN, SING). Kalimantan. Bandjermasin, Boschwezen 2176
(BO); Sanggau, Pengoedang, Neth. Ind. For. Service bb 29174 (SING); Sentarum Wildlife
Reserve, far north-west corner of Danau, Sungei Seriang, Giesen 140 (K), western border of
Danau, Nanga Kenelang, Giesen 148 (K); Sungai Kenara, Hallier 1359 (P, SING). Sabah.
“N Borneo”, Vilamil 300 (P); Beaufort, Cuadra NBFD A 1377 (K, SING); Gaya Island F.R.,
Kuripin SAN 28778 (SING), Gaya island (south), Aban SAN 57839 (SAN); Jesselton, Wood
2557 (SING); Kimanis, Bayak NBFD 2114 (K); Kota Belud, Kampung Lantige & Abai R.
F.R., Kandilis 7118 (SING); Kuala Penyu, Road to Pantai Tanjung Aru, Rimi et al. SPN 06605
(KEP); Mempakul, Bangawan, Abubakar NBFD 4111 (K, SING); Papar, Mandahan, Dewol &
Termiji SAN 80011 (SAN, SING), Talip Bidin SAN 80685 (KEP, SAN, SING); Sandakan, below
502 Gard. Bull. Singapore 64(2) 2012
Mr. Fox House, 50 ft, Jali SAN 65627 (SAN, SING); Sandakan, below Sabah Hotel, Sam SAN
25509 (SAN, SING); Sandakan, Chinese Cemetery, Chow & Aban SAN 75925 (SAN, SING);
Sandakan, Elopura, Agama A 2102 (SING), Cuadra A 2218 (SING), Darby road, Cuadra A
3197 (K, SING); Sandakan, Ernestina Road, Meijer SAN 24941 (SAN, SING); Sandakan, Jalan
Kapok, Clemente 4963 (SING); Sandakan, Leila F.R., 300 ft, Leopold & Termiji SAN 76680
(SAN, SING); Sandakan, Mile ¼ Derby road, Aban Gibot SAN 75916 (KEP, SAN); Sandakan,
Taman Rimba Sport Complex, Dewol SAN 105553 (SAN, SING). Sarawak. Beccari 3428
(K); J. & M.S. Clemens 22561 (P: barcode P03976237); Kuching, Museum Garden, Mamit S.
33492 (SAR); Kuching, Taman Budaya, Yahud et al., S. 57555 (K, KEP, L, MO, SAN, SAR);
Simunjan, Balai Ringin P.F., Stipni Bin Dollah S. 303 (SAR, SING), 1705 (SAR); Sadong, Ulu
Gunong, Omar 64 (SING).
BURMA. Martabania. Wallich Cat. no. 1597a (GDC: barcode G00134008), Wallich Cat.
no. 1597b (P: barcode P00349660), Wallich s.n. (P: barcode P00349658). Tenasserim &
Andamans. Helfer 3735 (P: barcode P00349659).
CAMBODIA. Bejaud 740 (P: barcode P00349636); Dam ta trau, Poilane 14071 (P: barcode
P00349651). Kompong Thom. Prey Krabey, reserve Sambo Spieu, Magnen s.n. 1909 (P:
barcode P00349640). Kompong Spen. entre Pum Love & Pum Rong, Poilane 17553 (P:
barcode P00349652). Siem Reap. Phnom Kulen, Martin 221 (P: barcode 00349644).
CELEBES. Boni. Heyne 2863 (BO). Malili. 25 m, Neth. Ind. For. Service Cel/IV-86 (SING),
Cel/IV-136 (SING), Cel/IV-137 (SING), Tabarano, Neth. Ind. For. Service bb 32356 (SING).
Moena. Wapai, Neth. Ind. For. Service bb 21738 (P: barcode P03976230). Manado.
Kolonodale, 400 m, Neth. Ind. For. Service bb 31525 (SING). Moena. Makoemoro?, 25 m,
Neth. Ind. For. Service bb 26302 (SING), bb 26303 (SING).
INDIA. Roxburgh s.n. (BR: barcode BR0000006912710).
JAVA. sine coll. no date (L); Blume s.n. (GDC: barcode G00134012); Korthals s.n. (L: barcode
L0005029); Reinwardt, s.n. (L sheet no. 908.127-246 & barcode L0005030). Batavia. Bodjong,
Bakhuizen v/d Brink 6354 (SING); Bodjong Ejot, Bakhuizen v/d Brink 6354 (P: barcode
P03976232); Buitenzorg, Hallier 202 (BO), Welter s.n. (SING), Moera Enim, Teysmann 4019
(BO). Semarang. Ngarengan kult., Kalshoven s.n. (BO). Soerabaja. Insel Bawean, Tambak,
Teysmann 1767 (BO).
LAOS. Banthat a 12 km de Savannakhet, Poilane 16384 (P: barcode P00330715), mare a 20
km de Savannakhet, Vidal 1752 (P: barcode P00330717).
MOLUCCAS. Smith s.n. 1796-1805 (BM: barcode 001053465).
PENINSULAR MALAYSIA. Johor. Johore Bharu, Carrick 1406 (SING). Kedah. Rahim KEP
12386 (SING). Melaka. Merlimau, Alvins s.n. (SING), Derry 53 (SING). Negeri Sembilan.
Cubitt 706 (SING); Forest Dept. 697 (SING); Angsi Forest Reserve, Othman KEP 23732
(KEP); Bahau Reserve, Mat Deris 654 (SING); Port Dickson, Blue Lagoon, Bremer 1819
(KLU), Cape Rachado F.R., roadside to Light House, Tsou 207 (NY); Seremban, Bain 18856
(KEP). Pahang. Ridley 1028 (SING); Kuantan, Abdul Rahman FMS 4172 (SING), Mahamud
FMS 3729 (SING), Mohd Soh 15735 (SING); Muazam Shah-Menchali main road, Sugumaran
et al. SM 212, leafy branch only (KLU); Pulau Berhala, Burn-Murdoch SFN 303 (SING).
Penang. Askey FMS 2554 (SING); “Pullo Penang”, Hunter, Wallich Cat. no. 1597E, (holo
K), Wallich Cat. no. 1597c (GDC: barcode G00134009); Sungai Pinang, Ogata 10344 (KEP);
Tasek Gelugor, Fyfe 29321 (SING); Tulloh (sic! = Teluk) Bahang, Curtis 314 (SING), Curtis
s.n. 1893 (SING), v 1892 (SING), vii 1892 (SING). Perak. Pangkor Island, Telok Nipah,
Chin 3137 (KLU). Selangor. Gombak, Klang Gates Quartz Ridge, Daniel Lee et al. s.n., leafy
branch only (KLU); Kepong, Forest Research Institute, Vethevelu FRI 29649 (KEP, SING);
Kuala Lumpur, Ramli KEP 94100 (KEP), Government Hill, Kalong 17463 (SING), Circular
503
Cyrtophyllum reapplied to Fagraea fragrans and allies
Road, Omar 7425 (SING), University of Malaya campus, Tan Bee Cheok s.n. (KLU), Zakiah
Hassan s.n. (KLU). Terengganu. Kemaman, Osman FMS 26996, FB, FL, 13 vii 1971 (KEP).
PHILIPPINES. Mindoro. Merritt 8842 (PNH, US?). Palawan. Curran Bur. For. 4501 (PNH,
US?); Aborlan, Iraan Mountains, Sulit 12512 (SING); Palawan, Cenabre 29990 (SING);
Taytay, Merrill 1202 (SING), Roque 30274 (SING).
SINGAPORE. Cantley’s Collector s.n. (SING); Botanic Gardens, Henderson 1329 (SING);
Nature Reserves Sector 17, Turner et al. NRS 269 (SING); Pulau Pawai, Sidek S 99 (SING);
Surrounding reservoir, Cantley’s Collector s.n. (SING!).
SUMATRA. Palembang. Endert 881 (SING), Moesi-Oeloe, Endert 160 (BO), 161 (BO).
Riauw islands. Pulau Kundor, Teruya 1624 (SING).
THAILAND. Bangkok. Marcan 707 (SING). Krabi. Krabi-Trang Road, Weerachai Nanakorn
WN 514 (SING); Jurin?, Kerr 8232 (SING). Surat Thani. Prsundej et al. 249 (KLU); Ko
Pha-ngan, Phromdet 25 (P: barcode P00349653). Narathiwat. Bacho, Sangkhachand 184 (P:
barcode P00349654); Sungai Padi, Bourke s.n. 1924 (P: barcode P00349655). Trat. Taphan
Hin, Geesink 6490 (P: barcode P00349656).
VIETNAM. Harmand 804 (P: barcode P00349666). Minh Hai. Iles de Poulo Condor, Harmand
873 (P: barcode P00349668), Pierre 3699 (P: barcode P00349677). Tourane. Quang-Nam, Da-
Nang, J. & M.S. Clemens 3862 (P: barcode P00349662). Lam Dong. Dalat & environs, Squires
889 (P: barcode P00349680). Long. Quang Tri, Chevalier 41189 (P: barcode P00349663).
A drawing of this species by William Hunter, collector of the type specimen, is on
display in the Penang Museum and a picture of this work is found in Gardner et al.
(2011: 116). In the past, two other species found in the Malay Peninsula, namely,
C. giganteum and C. wallichianum have been confused with C. fragrans. These two
species are not as common as C. fragrans; for differences, see under those species.
Burkill (1935) used the name Fagraea cochinchinensis A.Chev. for this
species, but apparently this had been based on Aidia cochinchinensis Lour. (a name
dating from 1790; Rubiaceae) and was therefore in the rst instance misapplied to
material of F. fragrans (Leenhouts 1962). Arguably, both taxa have some resemblance
in their paired leaves and branched cymes of small, cream-coloured to white owers
with exserted styles and stamens.
Holttum (1935) has given an account of the owering of this species in
Singapore, which is gregarious (many or nearly all trees in a population with
synchronised owering). It has two owering seasons, the main one around mid-year
and another towards year’s end, the owers opening several weeks following the
bud stage. At such times, it may be appreciated how different nomenclatural systems
may converge on an essential trait, for the species epithet in the latinised scientic
name refers to fragrant owers, and its Malay name, tembusu, could well allude to the
conspicuous sweet-and-sour scent that transpires into more than a hint of fermentation
(Malay: busu, unpleasantly odourous). Apparently, the owers last several days
even though the stamens are spent after the rst day of bloom, thus sustaining and
intensifying the perfumed aura around the tree. The fruits, which take a few months
to mature to an attractive yellow to red, are probably mainly dispersed by birds or
bats (Corner 1940). Of this tree, Ridley (1893: 323) observes that “it is difcult to say
whether the tree is more beautiful when covered with owers or fruits.”
504 Gard. Bull. Singapore 64(2) 2012
3. Cyrtophyllum giganteum (Ridl.) Ridl., Fl. Malay. Pen. 2 (1923) 421. — Fagraea
gigantea Ridl., J. Str. Br. R. As. Soc. 79 (1918) 98; Foxw., Mal. For. Rec. 3 (1927)
157; Burkill, Dict. (1935) 995; Corner, Wayside Trees Malaya (1940) 425; Browne,
For. Trees Sar. & Brun. (1955) 245. Kochummen, Tree Fl. Malaya 2 (1973) 273;
Cockburn, Trees Sabah 1 (1976) 211; Wong & Sugau, Sandakania 8 (1996) 11; Coode
et al. Checkl. Fl. Pl. Gymn. Brunei (1996) 174. LECTOTYPE (Wong & Sugau 1996):
Ridley 8921, Singapore, Garden Jungle (SING).
Fagraea sororia J.J.Sm. ex Cammerl., Bull. Jard. Bot. Btzg. III, 5 (1923) 319, pl. 5;
Heyne, Nutt. Pl. (1927) 1270. LECTOTYPE (Wong & Sugau 1996): Endert 44E 1P
515, Sumatra, Palembang (L; isolecto BO, K, SING).
Fagraea fragrans auctt. non Roxb.: King & Gamble, J. As. Soc. Beng. 74 (2) (1908)
611 pro parte, quoad Derry 272; Leenhouts, Fl. Males. I, 6 (1962) 304 pro parte,
Ashton, Manual Non-Dipt. Trees Sarawak 2 (1988) 310 pro parte, quoad Cyrtophyllum
giganteum Ridl., F. gigantea Ridl., F. sororia J.J.Sm., F. speciosa (non Blume) Ridl.
in syn.
Fagraea speciosa auct. non Blume: Ridley, J. Str. Br. R. As. Soc. 50 (1908) 122.
Tree, rarely to just 3–4 m tall, more often big, to 30 m tall or more; trunk to over 1
m in diameter; bark closely and nely ssured, grey-brown to dark brown. Leaves
elliptic; (4.4–)5–7(–7.5) cm long, (1.8–)2.2–3(–3.2) cm wide; base cuneate; apex
short-cuspidate to caudate; 4–6 mm long; margin conspicuously wavy when fresh
and in dried specimens; chartaceous; upper and lower surfaces smooth; midrib
prominent below, at to sunken above; secondary veins 4–6 pairs, upper side faint and
immersed in the blade, lower side faint to very slightly prominent; tertiary veins faint
to inconspicuous; petioles (0.8–)1–1.5(–2.3) cm long, 1–1.5 mm thick. Inorescence
(3.5–)5–6.5(–7.5) cm long; peduncle 3.5–5(–5.5) cm long, 1–1.5 mm thick; with 4–5
levels (tiers) of branching on the main axis, the branch tiers (5–)6–10(–12) mm apart,
lowest tier typically branched to (2–)3 orders. Flower pedicel 3–5 mm long, 0.3–0.5
mm thick; calyx (from the base to the lobe apices) 1.5–2 mm long, glabrous, calyx cup
1.5–2 mm wide, calyx lobes erect, 1–1.5 mm long, 1–1.5 mm wide, margins glabrous;
corolla tube subcylindric, 7–8 mm long, 1–1.5 mm wide near the base, upper part
very gradually and slightly wider, inside glabrous to minutely papillate; corolla lobes
broad-ovate to subobovate, 3–4 mm long, 2–2.5(–3.5) mm wide; stamens inserted at
the uppermost third of the corolla tube; laments 13–15 mm long, exsert 12–13 mm in
the open ower; anthers 1–1.5 mm long, 0.5–0.8 mm wide, each anther sac somewhat
narrowly ellipsoid; style (12–)18–22 mm long, protruding (5–)12–14 mm from the
corolla mouth in the open ower; stigma c. 0.5 mm across, lobes 2, low and rounded,
parting to present two slightly raised, hemispherical, papillate inner surfaces when
receptive, not recurving. Infructescence peduncle (2.5–)3–4(–4.8) cm long, 1–1.5 mm
thick. Fruit when mature to 4–6(–7) mm across; the base tightly clasped by the calyx
lobes. Seeds 1–2 mm across.
505
Cyrtophyllum reapplied to Fagraea fragrans and allies
Distribution. Malay Peninsula, Sumatra, Borneo.
Habitat. Lowland mixed dipterocarp forest.
Specimens examined: BORNEO. Brunei. Belalong, Ulu Ropan, 2000–2500 ft, Ashton BRUN
5275 (BRUN, K, SING); Tutong, Ulu Tutong, 150 ft, Ashton BRUN 908 (BRUN, K, SING).
Kalimantan. East Kutai, Sangkulirang district, Kerajaan River region, Kostermans 34793
(BO, K, L), Sungai Kerajaan, 40 m, Kostermans 5804 (BO, SING), Sungai Susuk, 40 m,
Kostermans 5693 (BO, K, SING); Muara Teweh, Popas?, Neth. Ind. For. Service bb 27769
(SING); Nunukan Island, Kostermans 8612 (BO, SING), 8955 (BO, SING); S.-O. Borneo,
Horyoep?, Winkler 2468 (SING); Sanggau, Bindang, Neth. Ind. For. Service bb 28143 (BO, K,
SING). Sabah. Keningau, Nabawan, Dewol & Karim SAN 78059 (K, L, SAN, SAR, SING);
Sandakan, Patrick SAN 25509 (KLU, SAN), Batu Sapi road, Meijer SAN 24942 (K, KEP,
L, SAN, SAR); Tawau, Apas Road, Brand SAN 21500 (SAN, SING), Kawa Road, Jaswir &
Aban SAN 26277 (BO, K, KEP, L, SAN, SING), Table Estate, 200 ft, Wood SAN 16650 (SAN,
SING). Sarawak. Baram, Melinau Gorge (4º10’N, 114º55’E), 1000 ft, Chew CWL 444 (K,
SING), Ulu Sungai Melinau Paku, Anderson 4085 (K, L, SAN, SAR, SING); Kuching, Bukit
Hujan, Omar 353 (K, SING); Lawas, Mt Bugoh ridge, Smythies BRUN 812 (K); Limbang, Ulu
Medamit, 900 ft, Chai, Wright & Othman S. 32335 (K, L. SAR, SING); Miri, S. Ukong, 10 m,
Othman S. 21394 (SING).
PENINSULAR MALAYSIA. sine coll. 16711 (KEP); Johor. Ulu Endau, Labis F.R., Sungai
Jasin, Ogata KEP 110427 (KEP). Melaka. Air Keroh Botanical Garden, Sugumaran & Lee
SM 193, leaves and stem bark (KLU), SM 194, leaves only (KLU); Bukit Saliokor?, Derry
272 (SING). Negeri Sembilan. Kuala Kelawang, Triang F.R., Lau & Jalil FRI 18249 (KEP).
Pahang. Kuantan, Mohd Soh FMS 15750 (SING), Rahman 15738 (SING). Selangor. Ayer
Hitam F.R., sine coll. KEP 55873, leaves only (KEP); Kajang, Bangi Reserve, Foxworthy
10289 (SING); Sungai Buluh F.R., Walton FMS 30770 (KEP). SINGAPORE. Singapore.
Garden Jungle, Ridley 8921 (lecto SING).
SUMATRA. Palembang. Buurman van Vreeden 77 (BO); Endert 44E 1P 429 (BO, L), 44E
1P 515 (BO, K, L, SING), 44E 2P 673 (BO, L); Banjoeasin en Koeboestreken, Endert 44E 2P
706 (BO, K, L); Lematang Ilir, Semangoes, Neth. Ind. For. Service bb 32270, leafy branch only
(SING); Medan 19, Bengkalis, Houtvester s.n. (BO); Moeara Doea Sh., Grashoff 400 (BO);
Moeara Enim, Teysmann 3796 (BO, L); Rawas, Dumas 1550 (BO, K), Grashoff 1060 (BO,
L). Lampongs. Kebang, Teysmann 4210 (BO, L); Tandj-Penang, Bruinsma 12 (BO). Upper
Riauw. Tenajan, Pakanbaru, Soepadmo 232 (BO, K, SING).
Cyrtophyllum giganteum and C. fragrans have very similar owers that appear only to
have consistently different corolla tube widths. However, several vegetative features
are very distinctive, such as the distinctively wavy margins in fresh leaves of C.
giganteum (C. fragrans have leaf margins which are generally plane). There are more
pairs of secondary veins in C. fragrans (7–9(–11)) that form distinct loops towards
the leaf margin but in C. giganteum there are fewer pairs of secondary veins (4–6)
that fork towards the leaf margin and do not form conspicuous loops. The mature
tree form is also often distinguishable: C. giganteum develops a very straight and
columnar bole like many other lowland forest canopy species, whereas C. fragrans
usually reaches up to only 30 m and typically develops several erect main branches
that are as tall as the main trunk. A well-grown C. fragrans tree develops a coarse,
506 Gard. Bull. Singapore 64(2) 2012
reticulately-sinuously ridged and ssured bark, as pictured in Gardner et al. (2011:
117); but that in a mature C. giganteum tree, as shown in Plate 35 of Ashton (1988:
313), is regularly and shallowly ssured.
4. Cyrtophyllum minutiorum K.M.Wong, sp. nov. C. caudato Ridley similis sed
cymis ramosioribus, oribus minoribus tubis corollae brevioribus (4–5 mm longis) et
stylis brevioribus (9–10 mm longis) differt. TYPE: Burley, Tukirin et al. 3278, 1–6 Jul
1989, Borneo, West Kalimantan province, G. Bentuang area, 5–10 km north of Masa
village, ridge SW of G. Bentuang, 1100 m alt. (holo SING; iso A, BO, K).
Tree, documented as 10 m tall; trunk to 15 cm diameter; bark texture in mature tree
trunk not known. Leaves elliptic to slightly lanceolate or oblanceolate; (5–)8–10(–
11) cm long, (1–)2–3(–3.5) cm wide; base cuneate; apex acuminate, hardly 0.5 cm
long; margin plane in dried specimens; coriaceous; upper and lower surfaces smooth;
midrib prominent below, channelled to raised above; secondary veins (4–)5–7 pairs,
upper side faint and immersed in the blade, lower side faint to immersed; tertiary
veins faint to inconspicuous; petioles (5–)12–18 cm long, 1.5–2 mm thick, petiolar
sheaths of a leaf pair fused and forming a shallow cup-like ochrea that tightly clasps
the stem; petiolar base auricles absent. Inorescence a few-owered cyme 8–10 cm
long; peduncle 4–4.5 cm long, 1(–1.5) mm thick; with 1(–2) levels (tiers) of branching
on the main axis, lowest tier typically branched to 3–4 orders. Flower pedicel 0–1 mm
long (owers subsessile), c. 0.5 mm thick; calyx (from the base to the lobe apices) c.
1.5 mm long, glabrous, calyx cup c. 1.5 mm wide, calyx lobes erect, c. 0.5 mm long, c.
0.5 mm wide, margins glabrous; corolla tube subcylindric, 10–12 mm long, c. 1 mm
wide near the base, inside glabrous to minutely papillate; corolla lobes broad-ovate
to subobovate, 2.5–3 mm long, c. 2–2.5 mm wide; stamens inserted at the uppermost
third of the corolla tube; laments c. 7 mm long, exsert c. 5 mm in the open ower;
anthers not seen; style 9–10 mm long, protruding c. 4 mm from the corolla mouth
in the open ower; stigma c. 0.1 mm across. Infructescence peduncle 4–5 cm long,
1–1.5 mm thick. Fruit (immature) c. 2 mm across; the base loosely to tightly clasped
by the calyx lobes. Seeds not examined.
Distribution. Borneo, W Kalimantan, only known from the type collection.
Habitat. On a ridge at the upper limit of Mixed Dipterocarp Forest or its transition to
lower montane forest.
5. Cyrtophyllum wallichianum (Benth.) M.Sugumaran & K.M.Wong, comb. nov.
— Fagraea wallichiana Benth., J. Linn. Soc. Bot. 1 (1857) 98; Clarke in Hooker
f., Fl. Brit. Ind. 4 (1883) 85; King & Gamble, J. As. Soc. Beng. 74 (2) (1908) 607;
Corner, Ways. Trees Malaya (1940) 426; Wong, Saw & Kochummen, Malayan Nat. J.
41 (1987) 267; Wong & Sugau, Sandakania 8 (1996) 12. — Fagraea lanceolata Wall.
507
Cyrtophyllum reapplied to Fagraea fragrans and allies
[Cat. (1829) no. 1599, nom. nud.] Schnizl., Iconogr. 2 (1851) t. 131, f. 1, nom. illeg.,
non Blume (1826); Miquel, Fl. Ind. Bat. 2 (1857) 376; Burkill, Dict. (1935) 995. —
Cyrtophyllum lanceolatum (Wall.) DC., Prod. 9 (1845) 31, nom. illeg.; Ridley, Fl. Mal.
Pen. 2 (1923) 421. TYPE: Wallich Cat. no. 1599, Penang (holo K; iso BM, GDC).
Fagraea fragrans auctt. non Roxburgh (1824): Kochummen, Tree Fl. Malaya 2 (1973)
273 pro parte, quoad Fagraea wallichiana Benth. in syn.; Leenhouts, Fl. Males. I, 6
(2) (1962) 304, pro parte, Ashton, Manual Non-Dipt. Trees Sarawak 2 (1988) 310
pro parte, quoad Cyrtophyllum lanceolatum DC., F. lanceolata Wall., F. wallichiana
Benth. in syn.
Tree, rarely to just 3–4 m tall, more often bigger, to 25 m tall; trunk to over 1 m in
diameter; bark ssured, grey-brown to dark brown. Leaves narrow to broadly-elliptic
to oblanceolate to obovate; (3.4–)6–10(–13) cm long, (1.4–)2–3(–5) cm wide; base
cuneate; apex acute to short-cuspidate; 2–5(–1.2) mm long; margin plane when fresh
and in dried specimens; thin-coriaceous; upper and lower surfaces smooth; midrib
prominent below, sunken above; secondary veins (4–)5–7 pairs, upper side faint to
immersed in the blade, lower side faint to very slightly prominent; tertiary veins faint
to inconspicuous; petioles (0.3–)0.8–1.2(–1.7) cm long, 1–1.5 mm thick, petiolar
sheaths of a leaf pair fused and forming a shallow cup-like ochrea that tightly clasps
the stem; petiolar base auricles absent. Inorescence a few-owered cyme (2.5–)3–
6(–7.8) cm long; peduncle (1.9–)2.3–3.5(–5.3) cm long, 1(–1.5) mm thick; with 1(–2)
levels (tiers) of branching on the main axis, the branch tiers 10–14 mm apart, lowest
tier typically branched to 1(–2) orders. Flower pedicel (4–)5–8 mm long, 1–1.5 mm
thick; calyx (from the base to the lobe apices) (3–)4–5(–6) mm long, glabrous, calyx
cup 3–4 mm wide, calyx lobes erect, 2–3 mm long, 2–2.5 mm wide, margins glabrous;
corolla tube somewhat infundibular, (12–)20–25 mm long, lower narrowed part 1–2
mm wide, upper ared part 6–8(–10) mm wide at the top, the lower narrowed tubular
part nearly as long as the upper ared part, inside glabrous to minutely papillate; corolla
lobes broad-ovate to subobovate, (5–)7–8 mm long, 3–4.5(–5) mm wide; stamens
inserted at the lower end of the upper ared part of the corolla; laments (27–)30–38
mm long, exsert 20–23(–28) mm in the open ower; anthers 1–1.5 mm long, 0.5–1
mm wide, each anther sac somewhat narrowly ellipsoid; style (34–)42–45(–55) mm
long, protruding 22–25(–30) mm from the corolla mouth in the open ower; stigma
c. 0.5 mm across, lobes 2, low and rounded, parting to present two slightly raised,
hemispherical, papillate inner surfaces when receptive, not recurving. Infructescence
peduncle 2–2.5(–3) cm long, 1–1.5 mm thick. Fruit when mature to 7–9 mm across;
the base loosely to tightly clasped by the calyx lobes. Seeds 1–1.5 mm across.
Distribution. Endemic to the Malay Peninsula.
Habitat. Lowland forest on hills and ridges.
Specimens examined: PENINSULAR MALAYSIA. Johor. Hutan Simpan Labis, Gunung
508 Gard. Bull. Singapore 64(2) 2012
Beremban, Wong FRI 30882, leafy branch only (KEP). Kedah. Gunung Jerai, Kochummen
FRI 18086 (KEP, SING), FRI 29495 (KEP), KEP 85027, leafy branch only (KEP), KEP 94416
(KEP). Kelantan. Relai Forest Reserve, Cockburn FRI 7251 (KEP, SING), FRI 7280 (KEP,
SING), FRI 7411 (KEP, SING). Malacca. Maingay 1029 (BM, mounted on the same sheet,
lower half, as the isotype Wallich Cat. no. 1599). Penang. Wallich Cat. no. 1599 (holo K; iso
BM, GDC); Govt. Hill, Curtis s.n. 1899 (SING), Ridley 7066 (SING); Moniots Road, Burkill
SFN 3330 (SING), Curtis 375 (SING); Penang Hill, Ahmad Shukor AS 91 (SING), Corner SFN
31597, leafy branch only (SING), Low et al. LYW 206, leafy branch only (KLU), Nauen s.n.
1940 (SING), Samsuri Ahmad SA. 999 (KEP, KLU, SING), Sidek S. 226 (SING!), Symington
KEP 28043 (SING); Western Hill, Nauen s.n. 1940 (SING). Terengganu. Ulu Brang, Gunung
Padang, Whitmore FRI 12803 (KEP, SING).
Cyrtophyllum wallichianum is generally a smaller tree (up to 25 m tall) of hills
and ridges, compared to C. fragrans, which can grow taller (up to 30 m tall) but is
typically found on gentler terrain in the lowlands. C. wallichianum is vegetatively
quite similar to C. fragrans but the oral characters are different. The inorescence
of C. wallichianum is usually less branched, 1(–2) orders, with fewer owers; that
of C. fragrans is generally more branched, 2(–3) orders, and so bears more owers.
The corollas of C. wallichianum are much bigger, 12–25 mm long, 6–10 mm wide,
whereas in C. fragrans they are smaller, (4–)6–8 mm long and 1.5–2(–2.5) mm wide.
As noted by Ridley (1918), F. caudata (= C. caudatum) which occurs only
in Borneo (Sarawak and Brunei), closely resembles C. wallichianum. Cyrtophyllum
caudatum differs from C. wallichianum in having more coriaceous, lanceolate-caudate
leaves and extremely slender peduncles and pedicels, cylindric corolla tubes and
shorter stamens.
Indeterminate material
A taxon that resembles C. caudatum and C. minutiorum in leaf characters, but without
owering or fruiting material, occurs in New Guinea. Its habitat, too, is similar: ridge
forest in hilly terrain, only even higher at c. 800 m elevation. This is quite possibly a
distinct species.
Specimens examined: NEW GUINEA. Japen Island, c. 800 m, Neth. Ind. For. Services bb.
30336 (BO, SING), bb. 30352 (BO, SING).
ACKNOWLEDGEMENTS. We thank the Keepers and Curators of the A, BO, K, KEP, KLU,
L, NY, US, SAN, SAR and SING herbaria for permission to consult reference materials and
loans of specimens. J.J. Wood, J. Dranseld, N. Martland (Kew), E.F. de Vogel, C.E. Ridsdale,
J.-F. Veldkamp, L. Willemse (Leiden), C.C. Davis, D. Boufford, W. Zhang, K. Gandhi
(Harvard), L.G. Saw, R.C.K. Chung (Kepong), C.L. Chan (Kota Kinabalu), A.A. Joffre, K.
Arifn (Brunei), R. Kiew (formerly SING), S. Lee, C.J.H. Soh and M.I. Bazilah (SING) are
thanked for assisting with a whole range of research matters. Y.W. Low, Singapore Botanic
Gardens, assisted with research and assessment of information related to types and specialised
collections. We thank C. Puff for help with the Latin diagnosis used.
509
Cyrtophyllum reapplied to Fagraea fragrans and allies
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510 Gard. Bull. Singapore 64(2) 2012
