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CHAPTER 8
SYSTEMATIC TREATMENT OF THE
ROTHMANNIA COMPLEX IN MALESIA
Based on evidence discussed in the preceding chapters and the cladistic
analyses attempted (Chapter 7), the Rothmannia complex in Malesia cannot at present
be resolved as distinct groups, in any conclusive way. At the same time, both species
of Kochummenia consistently associated as a subclade that was distinct from other taxa
included in the analysis. Hence, as there is also no evidence for refuting its status as a
distinct genus (the analyses in Chapter 7, not including a large array of other genera in
the tribe, did not permit this kind of inference), and because it is a validly published
genus, Kochummenia is here maintained for taxonomic expediency.
Rothmannia is here accepted as being strictly defined only by the African
species with right-contorted corolla lobes (i.e., the type alliance). All Malesian taxa,
except for Kochummenia as explained above, are here treated under “Rothmannia s.l.”.
A key to Kochummenia and Rothmannia s.l. is provided, as well as “generic” and
species descriptions. For Malesia, the taxa here accepted and delimited mostly need a
combination under the appropriate generic name, Rothmannia or a genus to be
separated from it, if the preliminary results of the phylogenetic analysis (section 7.1)
are corroborated. Rothmannia s.l. includes 18 species, involving the need for
recognition of 11 new combinations, five new species and one new variety.

8.1 Key to Malesian taxa of the Rothmannia complex
1a. Flowers subhypocrateriform, with linear corolla lobes, the length more than
10 times the width. Corolla lobes distinctly longer than tube. Anthers fully
included within the tube (inserted at the lower ⅓ of the tube)
(Kochummenia)
2a. Stipules large, 6–8 × 7–9 mm. Mature leaves broad, typically 8–12
cm wide. Flower calyx tube long, 8–11 mm long; corolla lobes long,
6–11 cm long..............K. stenopetala (King & Gamble) K.M. Wong
2b. Stipules smaller, 3–6 × 3.5–6 mm. Mature leaves narrower, 4–7 cm
wide. Flower calyx tube shorter, 3–6 mm long; corolla lobes shorter,
1.5–2 cm long.
3a. Lower surface of mature leaves, stipules and inflorescences
pubescent. Lamina variously shaped, elliptic, obovate or
oblanceolate; drying thin-coriaceous. Inflorescence rachis long
1–2 cm long…………………………...K. parviflora K.M.Wong
3b. Lower surface of mature leaves, stipules and inflorescences
hispid. Lamina narrowly elliptic; drying chartaceous.
Inflorescence rachis shorter, less than 1 cm long..............K. sp. 1
1b. Flowers varying from infundibular, campanulate to hypocrateriform, with
corolla lobes variously shaped, ovate, ovate-elliptic, ovate-oblong, lanceolate
to elliptic, the length not more than 5 times the width. Corolla lobes more-
or-less of same length as or shorter than the tube. Anthers fully included
within the corolla tube, or partly exserted (inserted above the basal ⅓ of the
tube) (Rothmannia s.l.)
4a. Primary branch on erect shoots with only 2 nodes, terminated by an
inflorescence, subsequent higher-order branching typically absent but
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arely develop up to 2 orders. Branch leaves in pairs at the proximal
node and completely vestigial at the distal node (often microscopic in
size). Flowers with infundibular corollas. Anthers long, 20–25 mm
long [Malaya & Sumatra]………...…. “R.” macrophylla R.Br. ex
Hook.f.
4b. Primary branch and successive higher-order modular branch units with
only 2 nodes and a terminal inflorescence each, new modular units
developing from either or both leaf axils at the proximal node of the
previous module. Branch leaves typically in groups of 3’s, each
trifoliate group based on a distal node with only one leaf normally
developed (the other vestigial or very much reduced in size), and the
proximal node with a pair of normal leaves. Flowers with
hypocrateriform or campanulate corollas. Anthers shorter, typically
less than 20 mm long.
5a. Leaves (including young leaves of shoot tips) completely glabrous
on both sides. Corolla tube glabrous both inside and outside
[Papua New Guinea]…………................................ “R. papuana”
5b. Leaves in early stages always hairy (at least on the veins on lower
side, or both sides). Corolla tube glabrous only on inside or
outside, never both.
6a. Stipules on leafy branches long, 10–23 mm long, apex (10–)
15−20 mm long; inner surface of stipules with colleters in a
central triangular area [Papua New Guinea]………. “R.
uvarioides”
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6b. Stipules on leafy branches shorter, less than 10 mm long, apex
shorter, less than 10 mm long; inner surface of stipules with
colleters all over except at the distalmost portion.
7a. Calyx glabrous to scantily puberulous, the hairs if
present, very minute and scattered (detectable only with
× 10 magnification) on the surface.
8a. Corolla throat glabrous inside; lobes narrow, 0.25–
0.3 cm wide [Philippines]………. “R. lagunensis”
8b. Corolla throat hairy inside; lobes wider, 0.5–1.7 cm
wide.
9a. Inflorescence a much-branched cyme, typically
branching 2–3 orders, typically more than 5
flowers per cyme. Corolla tube short, 2–3.5
cm long, mouth narrow, 3.5–4 mm wide, base
1–1.5 mm wide [Sulawesi & Lesser Sunda
Islands].............................… “R. sundaensis”
9b. Inflorescence unbranched or branched to at most
2 orders, typically up to 5 flowers per cyme.
Corolla tube longer, 4–14.5 cm long, mouth
wider, 5–20 mm wide, base wider, 2–5 mm
wide.
10a. Leaf apex short-to-long-acuminate.
246
Corolla lobes 5. Calyx lobes 5.
Anthers 5. Mature fruits large, 6–11.5
× 5–11 cm, pendulous; fruit wall 3–5

247
mm thick, very grainy [Borneo]….. “R.
ridsdalei”
10b. Leaf apex acute to obtuse. Corolla lobes
typically 6–8, rarely 5. Calyx lobes
typically 6–8, sometimes reaching 10,
rarely 5. Anthers 6–10. Mature fruits
smaller, 2.5–4.5(–5) × 2.5–4.5(–5) cm,
erect; fruit wall thinner, 2–3 mm thick,
not grainy [Borneo &
Philippines]…………….. “R. merrillii”
7b. Calyx conspicuously (sparsely to densely pubescent)
hairy on the surface, hairs c. 0.5 mm long, visible to the
unaided eye.
11a. Flower pedicels long, (3–)5–15 mm long. Corolla
campanulate with non-speckled throat.
[Sulawesi]…………….………. “R. forsteniana”
11b. Flower pedicels shorter, typically 0.5–2 mm long,
rarely 3 mm long. Corolla hypocrateriform to
campanulate; if campanulate, with speckled
throat.
12a. Corolla campanulate. Style sparsely hairy
[Myanmar, Thailand, Malay Peninsula,
Sumatra, and Java]…... “R.” schoemannii
(Teijsm. & Binn.) Tirveng.
12b. Corolla hypocrateriform. Style glabrous.

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13a. Corolla tube short, (0.6–)2–3(–3.5) cm
long.
14a. Stipules short, 3–4.5 mm long,
apex 0.5–1.3 mm long. Leaves
thin- to thick-coriaceous.
Corolla tube not distended at the
upper half. Corolla throat with
hairy zone covering part or all
of the tube from the corolla
mouth downwards [Papua New
Guinea]……... “R. nigrescens”
14b. Stipules longer, 5–8 mm long,
apex 2–6 mm long. Leaves
chartaceous. Corolla tube
distended at the upper half.
Corolla throat glabrous
[Philippines]... “R.graciliflora”
15a. Cymes much reduced or with
only 1 order of distinct
branching, 1–3 flowers per
cyme. Calyx tube long, 9–
11 mm long, outer surface
hirsute with suberect hairs.
Corolla lobes wide, 0.4–1
mm. Corolla mouth, 6–7.5

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mm wide [Papua New
Guinea] “R. kassamensis”
15b. Cymes distinctly branched to
2–5 orders, 11–130 flowers
per cyme. Calyx tube
shorter, 3–8 mm long, outer
surface velvety-hairy.
Corolla lobes narrower, 0.1–
0.4 mm. Corolla mouth
only 2.5–3.5 mm wide.
16a. Inflorescence a very
compact cyme with
condensed branches,
rachis 0.5–1 cm long.
Peduncle short, 0.1–0.2
cm long, 4–5 mm thick.
Calyx lobes (4–)6–10
mm long [Papua New
Guinea]…………… “R.”
macromera (Lauterb. &
K. Schum.) Fagerl.
16b. Inflorescence a more laxly
branched cyme, rachis 1–
2.5 cm long. Peduncle
longer, 0.5–2 cm long,
2.5–3.5 mm thick. Calyx

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lobes shorter, 0.5–1 mm
long [New Guinea]... “R.
anisophylloides”
13b. Corolla tube longer, (3.5–)4–14.5 cm long.
17a. Corolla throat with hairy zone covering
part or all of the tube from the corolla
mouth downwards.
18a. Leaf apex cuspidate to caudate, to
1.5 cm long. Stipule apex short, to
c. 0.5 mm long. Number of
flowers 3–9. Calyx lobes typically
3–25 mm long, rarely 1 mm.
Corolla lobes narrow, 0.6–1.4 cm
wide [Sumatra, Peninsular
Malaysia & Borneo]..“R. grandis”
18b.Leaf apex acute to short-
acuminate, to 0.3 cm long. Stipule
apex longer, 1–2.5 mm long.
Number of flowers fewer, 2–3.
Calyx lobes shorter, 0.5–1 mm.
Corolla lobes wider, (1.5–)1.8–2.5
cm wide [Philippines]….... “R.
negrosensis”
17b. Corolla throat glabrous.
19a. Peduncle 0.2–0.4 cm long. Style
and stigma 60–85 mm long, the

8.2 Species enumeration
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stigma included but reaching just
below or slightly above the corolla
mouth and near or above the
anthers.[Philippines].“R.leytensis”
19b. Peduncle longer, 0.5–6.5 cm long.
Style and stigma shorter, 15–50
mm long, the stigma included but
reaching less than ⅔ way along the
corolla tube and well below the
anthers [Borneo]…………….. “R.
pseudoternifolia”
Species are enumerated here in alphabetical order under Kochummenia and
Rothmannia s.l., and specimens cited and examined are incorporated at the end of each
species account. In the specimen listing, two sets of information are provided within
round brackets, viz., one set of brackets containing the collection date and type of
material (abbreviations used are FLO for open flowers or buds; FRU for post-flowering
material, including just calyces or fruits; and ST for reproductive material lacking); and
a second set of brackets containing acronyms of herbaria where material has been
documented (with an exclamation mark (!), indicating which have been studied or
seen). Conversions from measurements in imperial units (e.g., feet or miles) are given
to the nearest unit (m or km) within square brackets. Within each locality, specimens
are listed by author names (surnames) in alphabetical order and ascending numbers for
collecting numbers. For Rothmannia s.l., every species account is accompanied by a

otanical line drawing or a photograph of a representative specimen (frequently the
holotype specimen); for Kochummenia, a photograph of the type specimen is provided
only for K. stenopetala.
Kochummenia K.M.Wong
Malay. Nat. J. 38 (1984) 31; Wong, Tree Fl. Malaya (1989) 366.
TYPE SPECIES: Kochummenia stenopetala (King & Gamble) K.M.Wong
Randia sect. Grandiflorae Ridl., Fl. Malay Penins. 2 (1923) 71, pro parte quoad R.
stenopetala Ridl.
Randia sensu King & Gamble, J. As. Soc. Beng. 72 (1903) 218, pro parte quoad
Gardenia stenopetala King & Gamble.
Shrubs or treelets with typically solitary trunk. Trunk orthotropic with
continuous production of branches. Branches opposite and decussate on the trunk,
plagiotropic by substitution, sympodial in development; primary branch and successive
higher-order modular branch units each with only 2 nodes and a terminal inflorescence
each, new modular units developing from either or both leaf axils at the proximal node
of the previous module. Stipules interpetiolar, free, persistent, inner surface hairy with
2–3 rows of colleters at the basal part. Leaves typically in groups of 3’s, each trifoliate
group based on a distal node with only one leaf normally developed (the other vestigial
or very much reduced in size), and the proximal node with a pair of normal leaves;
petiolate to subsessile, axils of the midrib and lateral veins frequently with domatia on
the lower leaf surface, margin entire. Inflorescences terminal in origin but later
appearing lateral because of displacement to one side during sympodial branch
development; distinctly pedunculate to subsessile, distinctly branched cymes. Flowers
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isexual; small or medium, corolla tube typically not more than 3 cm long; typically
pedicellate, sometimes subsessile. Calyx 5-lobed, colleters present in small groups on
the inner surface, hairy on the outside. Corolla 5-merous; subhypocrateriform; the tube
creamy-white to greenish, the lobes creamy-white to greenish; the outer surface
glabrous or hairy, the inner surface glabrous; lobes contorted to the left in the flower
bud, distinctly longer than the tube; stamens dorsifixed, attached at the lower part of the
corolla tube, fully included within the tube, anthers sessile or on short filaments, linear;
style shorter than the corolla tube, glabrous; stigma included, narrowly subovoid, 2-
lobed, smooth when dried; ovary 2-celled; ovules many, placentation axile. Mature
fruits berry-like, medium, to 3.5 cm across; ellipsoid, crowned with persistent calyx,
outer surface glabrescent to sparsely hairy, drying smooth or wrinkled; 2-loculate.
Seeds many, immersed in a pulp-like placental tissue.
NOTES. The genus was first described by Wong in 1984 after K.M.
Kochummen, a Malaysian botanist. The type species, Randia stenopetala Ridl., was
placed under Randia sect. Grandiflorae by Ridley (1923), who considered it closely-
related to Randia macrophylla (= “Rothmannia macrophylla”) in the present account.
The genus has very similar plant architecture and branching to Rothmannia but differs
significantly in the characters outlined in the key to the Malesian taxa of the
Rothmannia complex.
DISTRIBUTION. Three species, endemic to Peninsular Malaysia.
1. Kochummenia parviflora K.M.Wong, Malay Nat. J. 38 (1984) 32; Wong, Tree
Fl. Malaya (1989) 366.
TYPE: Sinclair & Kiah SFN 39935, Peninsular Malaysia, Terengganu, 36th mile
Terengganu-Besut road (13 July 1953 FRU) (holotype SING; isotype L!).
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Treelet, to 1 m high. Twigs densely pubescent. Stipules triangular, 3−5 mm
long, 4−6 mm wide, sparsely pubescent outside; apex long-acuminate, 2−3 mm long.
Young leaves on shoot tips pubescent on both sides. Mature leaves with petiole
subobsolete to 0.5(−1.3) cm long,1.5–2 mm thick, pubescent; lamina elliptic, obovate
or oblanceolate, 11−21 cm long, 4−7 cm wide, upper and lower surfaces sparsely
pubescent, leaf base symmetric, cuneate to sometimes unequal; leaf apex acute to
acuminate, acumen to 1 cm long; thin-coriaceous when dry; midrib on upper side
slightly raised, sparsely pubescent, on lower side distinctly raised, pubescent to scabrid;
secondary veins 9−12 pairs, on upper side flat to slightly raised, sparsely pubescent, on
lower side raised, pubescent; tertiary venation visible as a much-branched network
between pairs of secondary veins. Inflorescence: peduncle subobsolete to 0.1 cm long,
c. 1 mm thick, pubescent; habit compact, with 1−2 distinct branching orders, rachis 1−2
cm long, primary branches along rachis 1 pair, subobsolete to 0.1 cm long; bracts
narrowly ovate, 3−5 mm long, 1−1.5 mm wide, pubescent. Flowers 3−5 per cyme;
pedicels very short, to 1 mm long, c. 1 mm thick; calyx tube 3−5 mm long, 3−4 mm
wide, pubescent, hairs suberect, surface visible; calyx limb cup-shaped; calyx lobes 5,
linear, 8−12 mm long, sparsely pubescent; corolla subhypocrateriform, white, the tube
1−1.2 cm long, 3−4 mm wide at the throat, c. 2 mm wide at the base; non-speckled
inside, outer surface pubescent, inner surface glabrous; corolla lobes 5, linear, 1.5−2 cm
long, c. 0.1 cm wide, outside pubescent on the half uncovered in the bud, inside
glabrous; anthers 5, fully included within the tube (inserted at the lower ⅓), c. 11 mm
long; style and stigma 10−16 mm long, included but reaching just below the corolla
mouth and the level of the anthers, style smooth, glabrous. Fruits ellipsoid, 3−4 cm
long, 2.5−3 cm wide, sparsely pubescent, drying smooth or sometimes wrinkled; fruit
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wall c. 0.5 mm thick; peduncle 0.1−0.2 cm long, 2−3 mm thick. Seeds ellipsoid, 5–6 ×
5–6 mm.
DISTRIBUTION. Thus far endemic to Terengganu state, Peninsular Malaysia.
Only known from a few collections.
HABITAT. Recorded from primary and secondary lowland forests.
SPECIMENS EXAMINED.
PENINSULAR MALAYSIA. TERENGGANU: Dungun, Bukit Bauk F.R., Compt.
20 B, Chelliah KEP 104371 (19 June 1967 FRU) (KEP!, L!), Dungun, Km 62, road
from Bukit Besi to Kuala Terengganu, Kg Jarangau, Zainudin AZ 3339 (31 January
1991 FLO/FRU) (KEP!, UKMB!); 36th mile Terengganu-Besut road, Sinclair & Kiah
SFN 39935 (13 July 1953 FRU) (holotype SING; isotype L!); North to Kg Buloh,
Samy SA 663 (30 August 1986 FLO) (KEP!).
2. Kochummenia stenopetala (King & Gamble) K.M.Wong, Malay. Nat. J. 38
(1984) 32; Wong, Tree Fl. Malaya (1989) 366.
Basionym: Gardenia stenopetala King & Gamble, J. As. Soc. Beng. 72 (1903) 218.
LECTOTYPE (designated by Wong, 1984): Curtis 3144, Perak, Bujong Malacca (sine
date FLO) (SING!; isolectotype CAL).
Randia stenopetala (King & Gamble) Ridl., Fl. Malay Penins. 2 (1923) 79; Corner,
Wayside Trees of Malaya 1 (1952) 557.
Rothmannia stenopetala (Ridl.) Bremek., Kon. Ned. Akad. van Wet.–P 60, 1 (1957) 6.
(Fig. 53)
Treelet, to 4 m high, to 10 cm diameter. Twigs densely velvety hairy to hispid.
Stipules broadly triangular, 6−8 mm long, 7−9 mm wide, velvety hairy to sparsely
hispid outside; apex long-acuminate, 3−4 mm long. Young leaves on shoot tips hispid
255

to velvety hairy on both sides. Mature leaves with petiole subobsolete to 0.7−1.5(–2)
cm long, 2.5−4 mm thick, velvety hairy to hispid; lamina typically broadly elliptic to
broadly oblanceolate, 20−35 cm long, (6−)8−12 cm wide, upper surface sparsely hispid
to glabrescent, lower surface scabrid, velvety hairy to hispid; leaf base symmetric,
cuneate to sometimes unequal; leaf apex acute to acuminate, acumen to 1.5 cm long;
thin-coriaceous to chartaceous when dry; midrib on upper side slightly raised, glabrous
to sparsely pubescent, on lower side distinctly raised, velvety hairy to hispid; secondary
veins 10−18 pairs, on upper side flat, glabrous to sparsely pubescent, on lower side
raised, velvety hairy; tertiary venation visible as a much-branched network between
pairs of secondary veins. Inflorescence: peduncle 0.1−0.3 cm long, 2−3 mm thick,
velvety hairy; habit laxly arranged, with 1−2 distinct branching orders, rachis 3−7 cm
long, primary branches along rachis 1−2 pairs, 0.3−0.5 cm long; bracts narrowly
triangular, 3−5 mm long, 1−2 mm wide, velvety hairy. Flowers 5−9 per cyme; pedicels
very short, to 2 mm long, 1.5−2 mm thick; calyx tube 8−11 mm long, 4−6 mm wide,
densely velvety hairy to hispid, hairs suberect, surface visible; calyx limb cup-shaped to
short-cylindric; calyx lobes 5, linear, 7−12 mm long, pubescent; corolla
subhypocrateriform, white to green-white, the tube 1.5−2 cm long, 7−9 mm wide at the
throat, 2.5−3 mm wide at the base; non-speckled inside, outer surface sparsely
pubescent, inner surface glabrous; corolla lobes 5, linear, 6−11 cm long, 0.1−0.2 cm
wide, outside sparsely pubescent on the half uncovered in the bud, inside glabrous;
anthers 5, fully included within the tube (inserted at the lower ⅓), 10−12 mm long;
style and stigma 14−20 mm long, included but reaching just below the corolla mouth
and the level of the anthers, style smooth, glabrous. Fruits ellipsoid, 4−6 cm long, 2−3
cm wide, sparsely pubescent, drying smooth or wrinkled; fruit wall c. 1 mm thick;
peduncle c. 0.3 cm long, 2−3 mm thick. Seeds ellipsoid, 7–8 × 5–6 mm.
256

DISTRIBUTION. So far endemic to Perak state, Peninsular Malaysia (Fig.
54). Most of the collections were from the early to mid-20th century. Only recently
recollected from Bujang Melaka and Gunung Bubu F.R., Perak.
forests.
HABITAT. Primary and secondary lowland, Hill and Upper Dipterocarp
USES. The stem has been reported to treat headaches.
SPECIMENS EXAMINED.
PENINSULAR MALAYSIA. PERAK: Bujong Malacca, Curtis 3144 (December
1895 FLO) (lectotype SING!, isolectotype CAL), Pereira et al. JTP 792 (August 2000
FRU) (SAN!); Larut Hill, 3000 ft [915 m], =Curtis 3144 (25 December 1901 FLO)
(K!, SING!), Larut, Scortechini 1446 (sine datum FLO) (K!); Kledang Saiong, Tachun
32200 (21 February 1934 FLO) (SING!); Kuala Kangsar, Kg Hitam Jong, Hashim KL
123 (12 January 1957 ST) (KEP!); Gunung Bubu F.R., Chan CML 2 (2003 FLO)
(SAN!).
3. Kochummenia sp. 1
Kochummenia sp.,Wong, Tree Fl. Malaya (1989) 366.
Treelet, to 1.5 m high. Twigs hispid. Stipules triangular, 4−6 mm long, 3.5−4 mm
wide, hispid outside; apex long-acuminate, 2.5−3 mm long. Young leaves on shoot tips
hispid on both sides. Mature leaves with petiole to 1 cm long, 1.5−2 mm thick, hispid;
lamina narrowly elliptic, 20−30 cm long, 4−7 cm wide, upper and lower surfaces
hispid, leaf base cuneate to sometimes unequal; leaf apex acute to acuminate, acumen
to 1 cm long; chartaceous when dry; midrib on upper side flat to slightly raised, hispid,
on lower side distinctly raised, hispid; secondary veins 10−13 pairs, on upper side flat,
257

hispid, on lower side raised, hispid; tertiary venation visible as a much-branched
network between pairs of secondary veins. Inflorescence (bud): peduncle to 0.2 cm
long, c. 3 mm thick, hispid; habit compact, with c. 2 distinct branching orders, rachis
0.2 cm long, primary branches along rachis 1−2 pairs, c. 0.3 cm long; bracts narrowly
ovate, c. 3 mm long, 2.5−3 mm wide, hispid. Flowers 6 per cyme; pedicels very short,
c. 2 mm long, c. 1 mm thick; calyx tube 4−6 mm long, 2.5−3 mm wide, hispid, hairs
erect, surface visible; calyx limb cup-shaped; calyx lobes 5, long-linear, 9−10 mm long,
hispid; corolla subhypocrateriform, greenish white, the tube c. 0.6 cm long; non-
speckled inside, outer surface pubescent to hispid, inner surface glabrous; corolla lobes
5, linear, 1.8−2.4 cm long, c. 0.1 cm wide, outside hispid on the half uncovered in the
bud, inside glabrous. Fruits not seen.
NOTES. This taxon is closely related to K. parviflora, in having short corolla
lobes but differs in the characters mentioned in the key to the species of Kochummenia
in Malesia.
DISTRIBUTION. Thus far, endemic to Terengganu state, Peninsular Malaysia.
Known only from one collection.
HABITAT. Recorded on logged kapur forest, on shale.
SPECIMENS EXAMINED.
PENINSULAR MALAYSIA. TERENGGANU: Kemaman Hills, 3 miles SW Kg
Ayer Putih, near Sg Buaya, Whitmore FRI 20362 (20 August 1971 FLO) (KEP!, SAR!,
SING!).
258

Fig. 53. The lectotype specimen of Kochummenia stenopetala (King & Gamble) K.M. Wong, Curtis
3144, Perak, Bujong Malacca (SING).
259

260
Kochummenia parvifolia
Kochummenia stenopetala
Kochummenia sp. 1
Fig. 54. Distribution of Kochummenia species in Peninsular Malaysia.

Rothmannia sensu lato
Thunb., Kongl. Vet. Akad. Handl. 37 (1776) 67–68.
Fagerlind, Arkiv Bot. Stockh. 30 A, 7 (1943) 1; Bremekamp, K. Ned. Akad. van Wet–
P 60, 1 (1957) 3; Backer & Bakhuizen, Fl. Java 2 (1965); Yamazaki, J. Bot. 45
(2) (1970) 337; Tirvengadum, Nord. J. Bot. 3 (1983) 466; Wong, Malay. Nat. J.
38 (1984) 46; Wong, Tree Fl. Malaya (1989) 324; Puff & Wong, Sandakania 2
(1993) 13; Beaman & Anderson, Pl. Mt. Kinabalu 5 (2004) 340.
TYPE SPECIES: Rothmannia capensis Thunb.
Gardenia sensu Linn., sensu Miq., Fl. Ind. Bat. 2 (1856) 228, pro parte quoad G.
grandis Korth. & G. schoemannii Teijsm. & Binn.; sensu Merrill, J. Str. Br.
Roy. As. Soc. Spec. No. (1921) 564 & sensu Masamune, En. Phan. Born.
(1942) 682, pro parte quoad G. grandis Korth. & G. merrillii Elmer.
Randia sect. Euclinia DC., sensu Hook.f., in Benth. & Hook.f., Gen. Pl. 2 (1873) 89
pro parte & Fl. Brit. India 3 (1880) 114, pro parte quoad R. macrophylla R.Br.;
sensu K.Schum., in Engl. & Prantl, Nat. Pflanzenf. 4, 4 (1891) 76 pro parte
quoad R. macrophylla (R.Br.) Hook.f.
Randia sect. Gardenioides Hook.f., Fl. Brit. India 3 (1880) 114, pro parte quoad R.
exaltata Griff.; Ridley, Fl. Malay Penins. 2 (1923) 71.
Randia sect. Grandiflorae Ridl., Fl. Malay Penins. 2 (1923) 71, pro parte quoad R.
macrophylla R.Br. ex Hook.f.
Randia sensu Koord. & Valeton, Bijdr. 8 (1902) 88 & sensu King & Gamble, J. As.
Soc. Beng. 72 (1903) 214–215, pro parte quoad R. exaltata & R. macrophylla
Benth. & Hook.f. (sic!); sensu Merrill, J. Str. Br. Roy. As. Soc. Spec. No.
(1921) 562–564, pro parte quoad R. grandis (Korth.) Valeton ex Winkler & R.
kuchingensis W.W. Sm.; sensu Masamune, En. Phan. Born. (1942) 708, pro
261

parte quoad R. grandis (Korth.)Valeton ex Winkler, R. keithii C.E.C. Fischer &
R. kuchingensis W.W.Sm.; sensu Backer & Bakhuizen, Fl. Java 2 (1965) 310,
pro parte quoad R. schoemannii (Teijsm. & Binn.) Bakh.f.
Treelets to small, medium or large trees with typically solitary trunk or (in “R.
macrophylla”) shrublike, unarmed, often encountered with horizontal stem sequences
found above or below ground, these producing erect above-ground shoots. Trunk or
above ground shoot an orthotropic system with continuous or (in some species outside
Malesia) episodic production of branches. Branches opposite and decussate on the
trunk, typically plagiotropic by substitution, sympodial in development; primary branch
(and, in most species, successive higher-order modular branch units) with only 2 nodes
and a terminal inflorescence each; higher-order modular units developing from either
one or both leaf axils at the proximal node of the previous module. Stipules
interpetiolar, free or only very slightly fused at the base, persistent to caducous, inner
surface hairy with colleters restricted to 1–several rows at the basal part or forming a
central triangular area. Leaves typically in groups of 3’s, each trifoliate group based on
a distal node with only one leaf normally developed (the other vestigial or very much
reduced in size) and the proximal node with a pair of normal leaves, or (in “R.
macrophylla”) leaves completely vestigial at the distal node and only the proximal node
with a pair of normal leaves; petiolate to subsessile, axils of the midrib, secondary
veins, and sometimes other vein junctions frequently with domatia on the lower leaf
surface, margin entire. Inflorescences terminal in origin, (in all species except “R.
macrophylla”) later appearing lateral because of displacement to one side during
sympodial branch development; distinctly cymose or sometimes much reduced to a 1-
flowered structure (but always with bracts marking potential branching on the main
inflorescence axis); distinctly pedunculate to subsessile. Flowers bisexual; small or
262

large, corolla tube typically not more than 15 cm long; typically pedicellate, sometimes
subsessile. Calyx 5–8(–10)-lobed, colleters present in small groups on the inner
surface, glabrous or glabrescent to hairy on the outside. Corolla typically 5-merous (in
Malesia, 2 species, “R. merrillii” and “R. negrosensis”, often 6–8); hypocrateriform or
campanulate (in Malesia only “R. forsteniana” and “R. schoemannii” have campanulate
corollas) or infundibular (“R. macrophylla”); the tube creamy-white to white with a
maroon tinge at the base in “R. macrophylla”, the lobes creamy-white, (in “R.”
macrophylla and “R. schoemannii”) with purple or reddish purple speckles or blotches
running down the corolla throat; the outer surface glabrous or hairy, the inner surface
glabrous or with hairs covering part or all of the tube from either the corolla mouth
downwards (“R. grandis”, “R. sundaensis”, “R. merrillii”, “R. negrosensis”, “R.
nigrescens”, and “R. ridsdalei”) or below the level of the anthers (“R. macrophylla”
and “R. schoemannii”); lobes contorted to the left in the flower bud, distinctly shorter
than or more-or-less as long as the tube; stamens (dorsi)medifixed, attached at the
upper part of the corolla tube or sometimes the tips slightly protruding (inserted above
the basal ⅓), anthers sessile or on short filaments, linear; style shorter than, as long as,
or slightly exceeding the corolla tube, or (in a single known species, “R. sundaensis”)
of two lengths in a heterostylous condition, mostly glabrous, but sometimes (e.g., “R.
schoemannii”) with scattered hairs; stigma included or partially exserted (in one
species, “R. sundaensis”), clavate to fusiform and sometimes narrowly oblong, 2-lobed,
smooth to ribbed or grooved when dried; ovary 2-celled; ovules many, placentation
axile. Mature fruits berry-like, indehiscent, medium to large, 2–5 cm or more across;
globose, more-or-less globose to broad ellipsoid, sometimes narrowly ellipsoid or
oblong or spindle-shaped, outer surface glabrous to hairy, drying smooth, wrinkled,
ridged, lenticellate or with a dense covering of minute flakes; 2-loculate. Seeds many,
immersed in a pulp-like placental tissue.
263

DISTRIBUTION. In the Malesian region, 18 species (including 2 varieties of
“Rothmannia pseudoternifolia”) are recognised. Out of this 18 species, 17 are endemic
to Malesia, including 5 endemic to Papua New Guinea; 4 to Philippines, 1 to Celebes
and 1 to Borneo and one species, “R. schoemannii” widespread and distributed in
Myanmar, Thailand, Peninsular Malaysia, Sumatra and Java (A fuller biogeographical
discussion is provided in section 9.4 of Chapter 9).
USES. Documentation on the uses of Rothmannia s.l. species in the Malesian
region is very scant. Some of the species have the potential for cultivation as
ornamental plants due to its showy flowers, e.g., “R. macrophylla” and “R. merrillii”.
In addition, most of the flowers are fragrant and may have potential in the perfumery
business. The timber of “R. schoemannii” has been reported by Burkill (1966) to be
heavy but not durable, however, some species in Papua New Guinea have moderately
hard and durable wood which are used for construction building.
1. “Rothmannia” anisophylloides [combination needed]
Basionym: Randia anisophylloides Wernham, J. Bot. 56 (1918) 74.
TYPE: Forbes 626, Papua New Guinea, Central Prov., Sogere region, Mt Korroro,
(Fig. 55)
2000 ft [609 m] (29 January 1886 FRU) (holotype BM!; isotype P!).
Tree, to 20 m high, to 20 cm diameter. Bark shallowly fissured, dark grey to
black; inner bark light brown to pinkish straw. Sapwood yellowish to orange-straw.
Twigs densely velvety hairy when young, becoming glabrescent when old. Branch
leaves typically in groups of 3’s, each trifoliate group based on a distal node with only
one leaf normally developed (the other vestigial or very much reduced in size), and the
proximal node with a pair of normal leaves. Stipules broadly triangular, 5−8 mm long,
264

6−11 mm wide, velvety hairy outside; apex short-to long-acuminate, 2−5 mm long.
Young leaves on shoot tips velvety hairy on both sides. Mature leaves with petiole
subobsolete to 0.7(−1.0) cm long, (2.5−)3–4 mm thick, velvety hairy; lamina broadly
elliptic to broadly obovate or oblanceolate, 15−30 cm long, (8−)10−18 cm wide, upper
surface sparsely pubescent to glabrescent, lower surface velvety hairy to sparsely
pubescent; leaf base subcordate, sometimes unequal; leaf apex acute to acuminate,
acumen to 0.5 cm long; thin- to thick-coriaceous when dry; midrib on upper side flat,
glabrous to sparsely pubescent, on lower side distinctly raised, densely tomentose to
velvety hairy; secondary veins (8−)10−12 pairs, on upper side flattened to sunken,
glabrous to sparsely pubescent, on lower side raised, pubescent; tertiary venation
visible as a much-branched network between pairs of secondary veins. Inflorescence:
peduncle 0.5−2.0 cm long, 2.5−3.5 mm thick, velvety hairy; habit laxly arranged, with
3−4(−5) distinct branching orders, rachis 3−4 cm long, primary branches along rachis
3−4 pairs, 1−2.5 cm long; bracts ovate, 2−4 mm long, 3−5 mm wide, velvety hairy.
Flowers (20−)40−130 per cyme; pedicels very short, 1–3 mm long, 1.5−2.5 mm thick;
calyx tube 3−8 mm long, 3−5 mm wide, densely velvety hairy, hairs suberect, surface
not visible; calyx limb cup-shaped; calyx lobes 5, short-triangular, 0.5−1 mm long,
densely tomentose or velvety hairy; corolla hypocrateriform, white to cream, the tube
0.6−1.7 cm long, 2.5−3.5 mm wide at the throat, 1−1.5 mm wide at the base; non-
speckled inside, outer surface sparsely to densely pubescent, inner surface glabrous;
corolla lobes 5, narrowly elliptic, 0.7−1.3 cm long, 0.1−0.4 cm wide, outside pubescent
on the uncovered half of the lobes, inside glabrous; anthers 5, fully included within the
tube (inserted at the upper ⅓), 6−15 mm long; style and stigma 5−17 mm long,
included but reaching just below the corolla mouth and the level of the anthers, style
slightly ridged, glabrous. Fruits ellipsoid, 3−4 cm long, 2.5−3 cm wide, pubescent,
265

drying with minute flakes; fruit wall 1−2 mm thick; peduncle 0.5−2 cm long, 2.5−3.5
mm thick. Seeds not seen.
galawon.
VERNACULAR NAMES. Galang-galang-buant (Waskuk); mete (Wagu);
DISTRIBUTION. Thus far known from the Windowoi Mountain in West Irian
Jaya Province, Indonesia, and Papua New Guinea (Central, Madang, Northern, and East
Sepik Provinces) (Fig. 84).
HABITAT. Occurs in lowland primary and disturbed forests, including the
transition zone between hill forest and alluvial plain, to 900 m, sometimes along rivers.
SPECIMENS EXAMINED.
PAPUA NEW GUINEA. MADANG PROV.: Aiome to Ramu river, Dovulat, 200ft
[61 m], Coode & Katik 32777 (11 March 1968 FRU) (L!, SING!); SE of Aiome, about
1 m S of airtrip, 250ft [76 m], Frodin NGF 26947 (22 June 1966 FLO) (L! ), North of
Aiome station, forest on spur, 400ft [122 m], Henty NGF 27438 (29 June 1966
FLO/FRU) (L!, SING!); below Wanuma village on main track to Ramu, 600 m,
Womersley NGF 48678 (31 July 1974 FLO) (BRI!, L!). NORTHERN PROV.:, near
USO resthouse c. 5 km NW of Gwaiari village, 50 m, Hoogland 3654 (16 August
1953 FLO) (L!). EAST SEPIK PROV.: Ambunti subdistrict, along Yapa, Hunstein
river, 400ft [122 m], Hoogland & Craven 10644 (25 July 1966 FLO) (L!); Hunstein
Range, 150 m, Takeuchi 6107 (15 July 1990 FLO) (L!), Waskuk Hills, hill forest, 60
m, Takeuchi & Regalado 10171 (16 June 1995 FLO) (L!). CENTRAL PROV.:
Koitaki, 1500ft [457 m], Carr 12875 (31 July 1935 FLO) (L!, SING!); Sogere
Region, Mt Korroro, 2000 ft [609 m], Forbes 626 (29 January 1886 ST) (holotype
BM!; isotype P!). INDONESIA. WEST IRIAN JAYA PROV.: Teluk Wandama,
Wondiwoi Mountains, 900 m, Schram BW 13448 (13 March 1962 FLO) (L!).
266

Fig. 55. The isotype specimen of “Rothmannia anisophylloides”, Forbes 626, Papua New Guinea,
Sogere region, Mt Korroro (P).
267

2. “Rothmannia” forsteniana [combination needed].
Basionym: Gardenia forsteniana Miq., Ann. Mus. Bot. Lugduno-Batavi 4 (1869) 236;
Koorders, Meded. ’s Lands Plantentuin. 19 (1898) 494; Koords.-Schum., Syst.
Verz. 11 (1914) 121; Koorders, Suppl. Fl. N.O. Celebes Part II (1922) 59, pl.
125.
LECTOTYPE (to be designated): Forsten 386, Celebes, Amurang (Leiden sheet n.
908.216-335) (November 1860 FLO) (L!).
Gardenia pelenkahuana Teijsm. & Binn. ex Koord., Meded. ’s Lands Plantentuin. 19
(1898) 495.
LECTOTYPE (to be designated): Koorders Kds. 19707β, Celebes, Minahasa
(20 March 1895 FRU) (BO!).
Rothmannia forsteniana Miq., nom. invalid., Kessler et al., Checklist of woody plants
(Fig. 56)
of Sulawesi, Indonesia. Blumea Supp. 14 (2002) 121.
Tree, to 15 m high, to 20 cm diameter. Bark greyish; inner bark brownish to
yellowish brown. Sapwood ochre. Twigs sparsely pubescent to hispid. Branch leaves
typically in groups of 3’s, each trifoliate group based on a distal node with only one
leaf normally developed (the other vestigial or very much reduced in size), and the
proximal node with a pair of normal leaves. Stipules broadly triangular, 2−7 mm long,
5−8 mm wide, glabrescent to pubescent outside; apex long-acuminate, 1−3 mm long.
Young leaves on shoot tips sparsely pubescent to hispid on both sides. Mature leaves
with petiole 0.1−0.5 cm long, 1−2.5 mm thick, pubescent to hispid; lamina broadly
elliptic-to-obovate to oblanceolate, (12−)14−26 cm long, (5−)8−15 cm wide, upper
surface glabrescent to sparsely pubescent to hispid, lower surface sparsely pubescent,
velvety hairy to hispid; leaf base cuneate, slightly rounded to unequal; leaf apex
268

acuminate, acumen to 1(−1.5) cm long; chartaceous to thin-coriaceous when dry;
midrib on upper side flat, glabrescent to sparsely pubescent, on pairs, on upper side flat,
lower side raised, sparsely pubescent, velvety hairy to hispid; secondary veins
(8−)10−16, glabrescent, sparsely pubescent to hispid, on lower side raised, pubescent,
velvety hairy to hispid; tertiary venation visible as a much-branched network between
pairs of secondary veins. Inflorescences: peduncle 0.5−1.5 cm long, 1−2 mm thick,
pubescent to hispid; habit laxly arranged, unbranched to with 1(−2) distinct branching
orders, rachis (0.8−)1−3 cm long, primary branches along the rachis 0–2 pairs, 0.2−0.5
cm long; bracts narrowly ovate, 2−2.5 mm long, 1.5−2 mm wide, sparsely pubescent to
hispid. Flowers 1−3(−4) per cyme; pedicels (3−)5−15 mm long, 0.5−1 mm thick; calyx
tube 4−11 mm long, 3−4 mm wide, sparsely pubescent to hispid, hairs appressed to
stiffly erect, surface visible; calyx limb short-cylindric, calyx lobes 5, short-triangular
to narrowly triangular, 1−5 mm long, pubescent to hispid; corolla campanulate, white,
the tube 3.5−5.5 cm long, 15−20 mm wide at the throat, the upper expanded tube length
2.5−5 cm long, the cylindric portion at the base 1−1.6 cm long and 1−3 mm wide; non-
speckled inside, outer surface sparsely pubescent, inner surface glabrous; corolla lobes
5, ovate, 3−4.1 cm long, 1−2 cm wide, outside sparsely pubescent on the uncovered
half of the lobes, inside glabrous; anthers 5, fully included within the tube (inserted at
the upper ⅓), 12−14 mm long; style and stigma 35−55 mm long, included but reaching
just below the corolla mouth and the anthers, style smooth, glabrous. Fruits ellipsoid,
8−14 cm long, 4−6.5 cm wide, glabrous to glabrescent, drying smooth and sometimes
shiny; fruit wall 1.5−2 mm thick; peduncle 0.5−1.5 cm long, 1−1.5(−2) mm thick.
Seeds slightly to much flattened and broadly ovate or suborbicular to narrowly
triangular, 7−9 mm × 9−10 mm.
269

NOTES. In the original publication, Ann. Mus. Bot. Lugduno-Batavi 4 (1869)
236 by Miquel, the type of the basionym, Gardenia forsteniana was not clearly
indicated. It is only indicated that Forsten was the collector where the specimen(s)
were collected from Celebes, around Tondano and Amurang district (“Celebes, in sylvis
circa Tondano et in distr. Amurang: Forsten”). There are four sheets from the L
herbarium, bearing Forsten’s handwriting with two of those sheets collected from
Tondano and the other two from Amurang.
Based on Article 9, paragraph 9.2 of the International Code of Botanical
Nomenclature, Vienna Code (2006), the sheet Herb. Lugd. Bat. 908216-335, Forsten
986, Celebes, Amurang is to be designated as the lectotype for the name.
This species stands out as a very distinct species from the other Malesian
species by having a corolla with campanulate shape, without speckles in the throat,
very long pedicels, (3−)5−15 mm long, and large ellipsoidal fruits, 8–14 × 4–6.5 cm.
Sulawesi).
Provinces).
VERNACULAR NAMES. Jambu hutan (Indonesia); kaju telor (Minahassa,
DISTRIBUTION. Confined to Sulawesi (North, Central, Gorontalo and South
HABITAT. Occurring in primary and disturbed lowland forest, also in hill and
lower montane forest, on limestone, clayey volcanic or alluvial soils, to 1050 m alt.
SPECIMENS EXAMINED.
SULAWESI. No locality. 500 m, Kjlllberg 2484 (10 October 1929 FLO) (BO!).
NORTH SULAWESI PROV.: 600 m, Kaudern 52 (10 April 1917 FLO) (L!),
Koorders Kds 18736 β, (12 March 1895 ST) (L!); Amurang: Forsten, s.n. (sine
datum FLO) (L!); Forsten 386 (Leiden sheet n. 908.216-335) (November 1860 FLO)
(lectotype L!); Tondano, Forsten s.n. (sine datum FLO) (L!); Mt Manimporose, Lam
2433 (15 April 1926 FRU) (L!); Mt Kawawtak, Alston 16241 (6 July 1954 FRU) (L!);
270

Bolaang Mongondow, Dumoga Bone National Park, Edwards camp, 750 m, De Vogel
& Vermeulen 6805 (29 March 1985 FLO) (L!); Dumoga Bone National Park, Tinayola
Camp on Sg Olama, 340 m, Milliken & Poluo WM 884 (10 August 1991 FRU) (L!);
Manado: 600 m, Koorders Kds. 18594β (5 April 1895 FRU) (BO!, L!), 500 m,
Koorders Kds. 18596β (15 January 1895 ST) (L!), Koorders Kds. 18740β (26 January
1895 ST) (BO!), 200 m, Koorders Kds. 19707β (20 March 1895 FRU) (BO!), Koorders
Kds. 24077β (2 June 1896 FRU) (BO!, L!); 340 m, sine coll. bb 14146 (27 November
1929 ST) (BO!), sine coll. 19135 (November 1911 FLO) (BO!); 220 km W of
Manado, c. 50 km inland from Pangi on Sg Ilanga, 350 m, Burley, Tukirin et al. 3681
(6 March 1990 FRU) (L!, SING!); N slope of Mt Kiabat, 1500 m, Forman 258 (27 June
1956 FLO) (BO!, K!); N slope Bukit Timampas near Mt Kiabat, 500 m, De Vogel 2633
(31 October 1973 FRU) (L!). GORONTALO PROV.: 250 km W of Gorontalo, 75
km inland from Papayuto, on tributary of Sg Papayuto, 150 m, Burley, Tukirin et al.
4071 (26 March 1990 FRU) (L!). CENTRAL SULAWESI PROV.: Mt Roroka
Timbu, W slope, c. 80 km S–SE of Palu, 1400 m, De Vogel 5118 (30 April 1979
FLO/FRU) (BRI!, L!); Sopu Valley, c. 60 km S–SE of Palu, c. 40 m high, De Vogel
5616 (29 May 1979 FRU) (BRI!, L!); Donggala Regency, Pangi-Binangga Nature
Reserve, Puncak Beringin Study Area, 590 m, Bynum 8318 (11 February 1991 FRU)
(L!); 6 km W–SW of Pendolo, c. 3 km S–SW of Boe, 850 m, Wieringa 1848 (18
August 1993 ST) (L!); SE of Bampres, NE of Tongoa, 750 m, Johansson, Nybom &
Riebe JNR 396 (16 March ? FRU) (L!); Luwuk area, inland from Batui and Seseba on
Batui River, at Totop camp, 2 hours upriver from Sinsing, 180 m, Coode 6023 (19
October 1989 FLO) (L!, SING!), Coode 6027 (19 October 1989 FRU) (L!); Bukit
Kerinduan, Dako, Mt Toli-Toli, Ramlanto & Zainal Fanani 524 (25 February 1985
FRU) (BO!, L!). SOUTH SULAWESI PROV.: Soroako-Wasuponda Road, Km 19,
500 m, Van Balgooy 3925 (7 July ? FRU) (L!).
271

Fig. 56. The proposed lectotype specimen to be designated for “Rothmannia forsteniana”,
Forsten 386, Celebes, Amurang (Leiden sheet n. 908.216-335) (L).
272

3. “Rothmannia” graciliflora [combination needed].
Basionym: Randia graciliflora Merr., Philipp. J. Sci., C 10 (1915) 109; Merr., Enum.
Philip. Fl. Pl. 3 (1923) 527.
LECTOTYPE (to be designated): Miranda F.B. 17988, Philippines, Mindanao,
Misamis, Mt Malindang (January–February 1913 FLO) (US!) (isolectotypes L!,
P!, SING!, NY!).
Gardenia negrosensis sensu Merr., Philip. Fl. Pl. 3 (1923) 527 pro parte quoad Ramos
(Fig. 57)
& Edaño B.S. 36787; non Merr. (1915).
Tree, to 18 m high. Twigs pubescent when young, becoming glabrescent when
old. Branch leaves typically in groups of 3’s, each trifoliate group based on a distal
node with only one leaf normally developed (the other vestigial or very much reduced
in size), and the proximal node with a pair of normal leaves. Stipules triangular, 3−5
mm long, 4−7 mm wide, sparsely pubescent outside; apex acuminate, 0.5–1 mm long.
Young leaves on shoot tips pubescent on both sides. Mature leaves with petiole
0.1−0.4 cm long, 1−1.5 mm thick, pubescent; lamina elliptic, obovate to oblanceolate,
12−16(−24) cm long, 3.5−7(−9) cm wide, upper surface sparsely pubescent when
young, becoming glabrescent when mature, lower surface sparsely pubescent; leaf base
cuneate, symmetric to unequal; leaf apex acuminate, acumen 0.4−1 cm long;
chartaceous when dry; midrib on upper side flat, glabrous to sparsely pubescent, on
lower side raised, pubescent; secondary veins 7−10(−13) pairs, on upper side flat,
glabrous to glabrescent, on lower side raised, pubescent; tertiary venation visible as a
much-branched network between pairs of secondary veins. Inflorescences: peduncle
0.3−1 cm long, 1−2 mm thick, pubescent; habit laxly arranged, with 2−3 distinct
273

anching orders, rachis 1.2−2.5 cm long, primary branches along the rachis 2–3 pairs,
0.5−1 cm long; bracts ovate, 1.5−2.5 mm long, 1.5−2 mm wide, pubescent. Flowers
11−25 per cyme; pedicels subobsolete to 1 mm long, c. 1 mm thick; calyx tube 3.5−4
mm long, 2−3 mm wide, sparsely pubescent at the limb but densely pubescent at the
hypanthium, hairs suberect, surface visible; calyx limb cup-shaped; calyx lobes 5,
narrowly triangular, 0.5−2 mm long, sparsely pubescent; corolla hypocrateriform, the
tube 1.8−2.4 cm long, bulging at the upper half, 2−2.5 mm wide at the throat, 1−1.5
mm wide at the base; non-speckled inside, outer surface pubescent, inner surface
glabrous; corolla lobes 5, narrowly elliptic, 0.6−0.9 cm long, 0.15−0.2 cm wide, outside
sparsely pubescent on the uncovered half of the lobes, inside glabrous; anthers 5−6,
fully included within the tube (inserted at the upper ⅓), 6−7 mm long; style and stigma
6−10 mm long, included but reaching less than ⅔ along the corolla tube, and well
below the anthers, style smooth, glabrous. Immature fruits globose, 2−2.8 cm long,
2−2.5 cm wide, glabrescent to glabrous, drying smooth; fruit wall c. 1 mm thick;
peduncle 0.5−1.5 cm long, 0.2−0.3 mm thick. Seeds not seen.
NOTES. In the original publication of Merrill in the Philipp. J. Sci., C 10
(1915) 109, there is only one specimen cited which is the type specimen for the species.
However, the holotype specimen which is in PNH (as Merrill had worked there), was
destroyed, hence, one of the isotypes has to be designated as the lectotype.
Based on Article 9, paragraph 9.2 of the International Code of Botanical
Nomenclature, Vienna Code (2006), one of the isotypes is here selected as the lectotype
i.e., from the United States National Herbarium in the Smithsonian Institution (US).
DISTRIBUTION. Thus far only known from 5 collections, recorded from the
southern part of the Philippines (Mindanao and Basilan) (Fig. 68).
274

Fig. 57. The proposed lectotype specimen to be designated for “Rothmannia graciliflora”, Miranda F.B.
17988, Philippines, Mindanao, Misamis, Mt Malindang (US).
275

SPECIMENS EXAMINED.
PHILIPPINES. MINDANAO. Zamboanga, Villamil 21884 A (May–June 1914
FRU) (K!), Babaran 26139 (July 1916 FRU) (K!, P!); Malangas, Ramos & Edano
36787 (October–November 1919 FRU) (L!), Misamis, Mt. Malindang, Miranda 17988
(January–February 1913 FLO) (lectotype US!) (isolectotypes L!, P!, SING!). Basilan,
Tecson & Rouve 25804 (May 1916 FRU) (K!).
4. “Rothmannia” grandis [combination needed].
Basionym: Gardenia grandis Korth., Ned. Kruidk. Arch. 2 (1851) 191; Walp., Ann.
Bot. Syst. 2 (1851) 796; Miq., Fl. Ind. Bat. 2 (1856) 231; Miq., Ann. Mus. Bot.
Lugduno-Batavi 4 (1869) 236; Merrill, Bibl. En. Born. Pl., J. Straits Branch
Roy. Asiat. Soc., Sp. No. (1921) 564; Masamune, En. Phan. Born. (1942) 708.
TYPE: Korthals, s.n., Borneo, Kalimantan, Mt Sakoembang (Leiden sheet no.
908.216-321) (sine datum FLO) (holotype L!)
Randia grandis (Korth.) Valeton ex Winkler, Bot. Jahrb. Syst. 44 (1910) 559; Merrill,
Bibl. En. Born. Pl., J. Straits Branch Roy. Asiat. Soc., Sp. No. (1921) 562;
Masamune, En. Phan. Born. (1942) 708; Coode et al., Checkl. Flow. Pl. &
Gymn. Brunei (1996) 289.
Randia kuchingensis W.W. Smith, Notes Roy. Bot. Gard. Edinb. 8 (1915) 326; Merrill,
Bibl. En. Born. Pl., J. Straits Branch Roy. Asiat. Soc., Sp. No. (1921) 562;
Masamune, En. Phan. Born. (1942) 708.
LECTOTYPE (designated by Wong et al., 1987): Haviland 676, Sarawak,
Kuching (29 December 1892 FLO) (lectotype SING; isolectotypes BM!, K!,
L!).
276

Rothmannia kuchingensis (W.W. Smith) K.M.Wong, Malay. Nat. J. 41 (1987) 270,
Tree Fl. Malaya 4 (1989) 406.
Rothmannia malayana K.M.Wong, Malay. Nat. J. 38 (1984) 51, Tree Fl. Malaya 4
(Fig. 59)
(1989) 406.
TYPE: Loh FRI 19252, Kelantan, proposed Batu Papan F.R. (22 February 1972
FLO) (holotype KEP!; isotype SING!)
Treelet, to 10 m high, to 17 cm diameter. Bark smooth to fissured or finely
cracked, grey-brown; inner bark yellowish green. Sapwood light brown. Twigs
pubescent when young, becoming scantily pubescent or glabrescent when old. Branch
leaves typically in groups of 3’s, each trifoliate group based on a distal node with only
one leaf normally developed (the other vestigial or very much reduced in size), and the
proximal node with a pair of normal leaves. Stipules ovate-triangular, 2−7 mm long,
3−8 mm wide, sparsely to densely pubescent outside; apex acuminate, 1−3 mm long.
Young leaves on shoot tips scabrid to velvety hairy on both sides. Mature leaves with
petiole 0.1−0.6(−1.0) cm long, 2−3 mm thick, pubescent; lamina broadly to narrowly
elliptic, obovate to oblanceolate, (8.5−)10−22(−31) cm long, 2.5−9(−12) cm wide,
upper surface glabrescent to glabrous, lower surface glabrescent, scabrid to velvety
hairy; leaf base cuneate, slightly rounded to unequal; leaf apex short- to long-
acuminate to acute, acumen to 2.5 cm long; thin- to thick-coriaceous when dry; midrib
on upper side flat to slightly raised, glabrescent to scantily pubescent, on lower side
raised, sparsely to densely pubescent or velvety hairy; secondary veins (5−)6−15 pairs,
on upper side flat, glabrous to glabrescent, on lower side slightly raised, glabrescent to
pubescent or velvety hairy; tertiary venation visible as simple connections between
pairs of secondary veins to invisible. Inflorescences: peduncle (0.1−)0.2−0.9 cm long,
277

2.5−4 mm thick, sparsely to densely pubescent or velvety hairy; habit compact to laxly
arranged, with 1−2 distinct branching orders, rachis 0.5−2 cm long, primary branches
along the rachis 1−2 pairs, (0.2−)0.3−0.7(−1.8) cm long; bracts ovate, 2−6 mm long,
2−4 mm wide, sparsely to densely pubescent or velvety hairy. Flowers 3–9 per cyme;
pedicels 1−2 mm long, 1.5−2.5 mm thick; calyx tube 5−13 mm long, 3−6 mm wide,
sparsely pubescent, hairs suberect, surface visible; calyx limb cup-shaped; calyx lobes
5, short-triangular to long-linear, (1−)6−25 mm long, sparsely pubescent; corolla
hypocrateriform, white to creamy white, the tube 4−7.5 cm long, 4−8 mm wide at the
throat, 2−4 mm wide at the base; non-speckled inside, outer surface sparsely
pubescent, inner surface with hairy zone covering part of the tube from the corolla
mouth downwards; corolla lobes 5, elliptic, oblong to ovate, 1.5−4.2 cm long, 0.6−1.4
cm wide, outside sparsely pubescent on the uncovered half of the lobes, inside hairy at
the base; anthers 5, fully included within the tube (inserted at the upper ⅓), 15–20 mm
long; style and stigma length 50−75 mm long, included but reaching just below the
corolla mouth and the level of or above the anthers, style smooth, glabrous. Fruits
globose to ellipsoid, 4–6.5 cm long, (2.5−)4−5.5 cm wide, glabrous, drying smooth;
peduncle 0.2−1.0 cm long, 2.5−6 mm thick. Seeds slightly to much flattened and
broadly ovate or suborbicular to narrowly triangular, 9−10 mm × 5−6 mm.
NOTES. The type specimens of Rothmannia kuchingensis (Basionym: Randia
kuchingensis), Rothmannia malayana and Gardenia grandis were examined.
Vegetatively and reproductively, all of them have similar and overlapping
characteristics. Palynologically, representatives of R. kuchingensis and R. malayana
showed similar sexine ornamentation, i.e., gemmate.
278

Gardenia grandis is an older name, hence, the epithet, grandis takes
precedence. Therefore, Rothmannia kuchingensis and R. malayana are here proposed
as new synonyms of “R. grandis”.
This species is very variable in terms of hairiness on the lower leaf surface
(glabrescent, scabrid to velvety hairy); corolla tube length (5−13 mm long); and calyx
lobe length ((1−)6−25 mm long).
The specimens from Bukit Raya, Kalimantan have extremely long calyx lobes
(1.5−2 cm long); long bracteoles (4−5 mm long); and long anthers (19−20 mm long),
whereas the typical “R. grandis” has shorter calyx lobes (0.1−1.2 cm long); shorter
bracteoles (1−3 mm long) and shorter anthers (15−17 mm long). However, the
vegetative characters are similar to those of “R. grandis”.
VERNACULAR NAMES. Medang suid, Rentap Hitam (Brunei); Berabas
hutan (Sarawak).
DISTRIBUTION. Peninsular Malaysia (Kelantan, Pahang and Johor states),
Sumatera (West, South & Riau Provinces) and Borneo (Kalimantan, Sarawak &
Brunei) (Fig. 58). Frequently found in swamps, including peat swamp forest, and
kerangas forest. Also in lowland Dipterocarp forest, along ridges, to 1200 m alt.
SPECIMENS EXAMINED.
PENINSULAR MALAYSIA. KELANTAN: proposed Batu Papan F.R., Loh FRI
19252 (22 February 1972 FLO) (KEP!, SAN!, SING!). PAHANG: Rompin, summit
of Mt Lesong, 2000 ft [609 m], Mohd Shah & Ahmad MS 3107 (25 November 1973
FRU) (SING!); Tahan Woods, Kuala Teku, narrow ridge crest, 600 ft [183 m],
Whitmore FRI 4749 (20 February 1968 FLO) (KEP!, SING!). JOHORE: Sg Kayu,
Kiah SFN 32153 (22 October 1936 FRU) (L!, SING!), Linggui area, Kochummen
KMK 1 (9 June 1992 FRU) (KEP!), Labis F.R., Mt Janing Plateau, 430 m, Heah FRI
32491 (4 June 1986 FLO) (KEP!), Wong FRI 30918 (23 October 1985 FRU) (KEP!,
279

L!, SAN!), trail to West of Mt Janing, on plateau, Pereira et al. JTP 856 (23 October
2002 ST) (SAN!). SUMATRA. WEST PROV.: Teysmann 917 (1911 FRU) (BO!).
SOUTH PROV.: Sg Lalang-Petaling, Laumonier TFB 356 (11 December 1979 FRU)
(L!); Sg Lalang, Laumonier YL 222 (1 September 1979 FRU) (L!). RIAU PROV.:
Tigapuluh Mountains, 5 km W of Talanglakat on Rengat–Jambi Rd., Bukit Karampal
area, 100 m, Burley, Tukirin et al. 1821 (3 December 1988 FRU) (L!), Mt Lancang
ridge, 600 m, Burley, Turkirin et al. 1913 (5 December 1988 FRU) (L!, SING!).
KALIMANTAN. No locality. Jaheri 1893 (sine datum FLO/FRU) (BO!); Teysmann
8345 (sine datum FRU) (BO!). Mt Sakoembang, Korthals s.n. (Leiden sheet no.
908.216-321) (sine datum FLO) (holotype L!). Bacutok, Winkler 3273 (1910 FRU)
(L!). C KALIMANTAN: Palangkaraya, Arboretum Nyaru Menteng area,
Ambriansyah AA 1543 (6 February 1996 FLO/FRU) (L!), Kuala Kurun, Mt Mas, 100
m, Tukirin 548 (10 March 1979 FLO) (L!); Bukit Raya: 130 m Nooteboom 4028 (26
November 1982 FLO) (BO!, BRI, L!), Nooteboom 4175 (5 December 1982 FRU)
(BRI!, L!), Nooteboom 4293 (14 December 1982 FLO) (L!); Sampit: Nyaru Menteng,
Arboretum, off km 28 Road to Sampit, 50 m, Sidiyasa 1419 (28 January 1995 FLO)
(L!, SAN!); 7 km E of Sampit, near sea level, Alston 13077 (18 January 1954
FLO/FRU) (BM!); Kuala Kuajan, 20 m, Kostermans 8029 (30 July 1953 FRU) (L!).
W KALIMANTAN: Semangit, Selimbau, Afriastini 1008 (5 May 1983 FLO) (BO!);
Sintang, Bukit Baka National Park, SE of camp, along ridge of lower Bukit Simat, 325
m, Church, Mahyar, Ruskandi & Nurdin 471 (1 November 1993 FLO/FRU) (L!,
SING!); Ketapang, Mt Palung National Park, Cabang Panti Research site, 30 m,
Laman, Ismail & Mirmanto TL 1295 (17 October 1997 FRU) (BO!). SARAWAK.
No locality: Native Collector 1240 (1 May 1912 FLO) (PNH!); Native collector 1866
(sine datum ST) (L!); sine coll. S 10372 (sine datum FRU) (SAR!). Kelapaan, Brooke
8835 (19 July 1954 FRU) (L!). KUCHING DIV. Kuching: sine coll. s.n. (27 April
280

1893 FRU) (SING!), Barlett s.n. (7 March 1905 FLO) (BM!), Haviland s.n. (30 June
1892 FLO) (SING!), Haviland 676 (29 December 1892 FLO/FRU) (BM!, K!, L!,
SING!), Haviland = 676 (29 December 1892 FLO) (BM!, SING!), sine coll. Sarawak
Museum Series 601 (13 September 1905 FLO) (SAR!), Haviland & Hose 676 (15
March 1893 FLO) (P!), 6th Mile, Egon S 442 (25 January 1927 FRU) (SAR!);
Santubong, Teluk Bandung, Dayang Awa & Othman Ismawi S 47086 (18 September
1984 FRU) (SAN!, SAR!), Hewitt 602 (January 1907 FLO/FRU) (SAR!), South of Mt
Santubong, 200 ft [61 m], Hj. Bujang S 13143 (17 May 1961 FLO) (L!, SAR!); Camp
Permai, Hj. Othman S 60112 (20 September 1990 FRU) (L!, SAN!, SAR!), Abg.
Mohtar et al. S 65117 (27 June 1992 FLO/FRU) (SAR!); Nyabau, Similajau F.R., 90 ft
[27 m], Brunig S 11949 (18 November 1959 FRU) (SAR!); summit of Mt Bungoh,
3500 ft [1067 m], Brunig S 9509 (9 October 1957 FRU) (SAR!, SING!); Matang,
Hewitt 863 (sine datum FRU) (SAR!), Matang Rd., Native Collector 574 (4 July 1911
FRU & FLO) (PNH!), Native Collector 1253 (23 February 1912 FLO) (K!, PNH!), 9th
Mile, Native Collector S 6226 (20 April 1914 FLO) (SAR!); Stapok F.R., 4 miles S of
Kuching, Dawos & Mamat S 28931 (11 March 1970 ST) (SAR!); Stampin, 4 miles
south east of Kuching, Anderson S 25951 (3 July 1966 FLO) (SAR!); Penrissen Rd.,
2nd Mile, Haviland 560/676 (sine datum FLO) (SAR!); Bako National Park, Tanjung
Limau forest, 0−200 ft [0–61 m], Carrick & Enoch JC 426 (3 May 1959 FRU) (KLU!,
SAR!, SING!); Lintang path, 100 m, Chai S 19723 (3 January 1964 FRU) (SAR!).
Lundu. Mt Pueh, Hj. Othman et al. S 49955 (3 October 1985 FRU) (KEP!, L!, SAR!).
SRI AMAN DIV. Sri Aman: Lingga, Hewitt 385 (June 1906 FLO) (SAR!); path to Kg
Kara, Kg Pungur Tapang, Ilias S 42727 (5 September 1980 FRU) (KEP!, SAN!,
SAR!). SARIKEI DIV.: Binatang, Pulau Bruit, Sanusi S 9288 (19 November 1957
FLO) (SAR!, SING!), 10 ft [3 m], Sanusi S 12952 (20 June 1960 ST) (SAR!), Sg
Kelepu, 10 ft [3 m], Anderson 7904 (9 June 1957 FLO) (L!, SAR!, SING!), Kelepu,
281

Brooke 8801 (11 July 1954 FRU) (L!, SAR!, SING!); Sg Semup, 10 ft [3 m], Anderson
S 5113 (12 April 1958 FRU) (L!, SAR!, SING!); Gelegei, Daro F.R., Hj. Bujang S
19667 (20 May 1964 FLO/FRU) (SAN!, SAR!, SING!), Matu Daro, 12 ft [3.6 m],
Sanusi S 12259 (20 November 1959 FRU) (SAR!, SING!). BINTULU DIV. Bintulu:
Bukit Urang, 120 ft [37 m], Brunig S 11905 (14 August 1959 FRU) (SAR!), 100 ft [30
m], Spawi S 11960 (20 November 1959 FLO/FRU) (SAR!), 80 ft [24 m], Brunig S
12044 (23 May 1960 FLO) (SAR!), 25 m, Brunig S 12085 (21 February 1960 FRU)
(L!, SAR!); Nyabau, Similajau F.R., 30 ft [9 m], Brunig S 11950, 18 November 1959
FRU) (SAR!). Miri: Niah, Haviland & Hose s.n. (May 1892 FLO) (K!). SIBU DIV.
Sibu: Naman F.R., Ahmad S 14327 (18 February 1966 FRU) (K!), S 14334 (18
February 1966 FLO/FRU) (SAR!), 100 ft [30 m], Clarke S 13802 (18 April 1961
FLO/FRU) (SAN!, SAR!, SING!); Sg Sawei, Loba Kabang, 4 ft [1 m], Anderson S
25/1 (25 January 1961 FRU) (L!, SAN!, SAR!, SING!); Sungai Trusan, Rantau
Panjang, Yahud et al. S 82569 (23 February 1999 FRU) (SAN!). BETONG DIV.
Betong: Brunig S 7466 (1 February 1956 FRU) (SAR!). BRUNEI. Belait: Bukit
Sawat, Sg Mau, 15 m, Ariffin et al. BRUN 16452 (4 July 1996 FRU) (BRUN!, SAN!);
Bukit Puan-Labi Rd., mile 1 [Km 1.6], Sinclair 10506 (30 August 1960 FRU) (L!,
SAR!, SING!).
282

MALAY
PENINSULA
SUMATRA
Fig. 58. Distribution of “Rothmannia grandis”.
283
BORNEO

Fig. 59. The proposed holotype specimen of “Rothmannia grandis”, Korthals, s.n.,
Borneo, Kalimantan, Mt Sakoembang (Leiden sheet no. 908.216-321) (L).
284

5. “Rothmannia” kassamensis [new species description needed].
TYPE (to be designated): Brass 32331, Papua New Guinea, Eastern Highlands Prov.,
(Fig. 60)
Kassam Pass (31 October 1959 FLO) (holotype L!).
Small tree, to 7 m high. Bark slightly peeling, blackish grey; inner bark wine
red. Sapwood yellowish straw. Twigs velvety hairy when young, becoming sparsely
pubescent when older. Branch leaves typically in groups of 3’s, each trifoliate group
based on a distal node with only one leaf normally developed (the other vestigial or
very much reduced in size), and the proximal node with a pair of normal leaves.
Stipules ovate-triangular, 4−6 mm long, 4−7 mm wide, sparsely pubescent outside;
apex long-acuminate, 4−6 mm long. Young leaves on shoot tips velvety hairy on the
lower surface, glabrescent on the upper surface. Mature leaves with petiole 0.2−0.3 cm
long, 1.5−2 mm thick, pubescent to velvety hairy; lamina elliptic, lanceolate to
oblanceolate, 14−18 cm long, (5−)6−8 cm wide, upper surface glabrous, lower surface
velvety hairy; leaf base cuneate, symmetric to subrounded; leaf apex long-acuminate,
acumen 1−2 cm long; thin-coriaceous to chartaceous when dry; midrib on upper side
flat, glabrescent, on lower side raised, velvety hairy; secondary veins 6−9 pairs, on
upper side flat to slightly sunken, glabrous, on lower side raised, velvety hairy; tertiary
venation visible as a much-branched network between pairs of secondary veins.
Inflorescences: peduncle 0.1−1 cm long, 1.5−2 mm thick, tomentose; habit compact,
with 1 distinct branching order, rachis c. 0.5 cm long, primary branches along the
rachis 1 pair, 0.15−0.3 cm long; bracts ovate 4–7 mm long, 4–5 mm wide, tomentose
to pubescent. Flowers 1−3 per cyme; pedicels 2−3 mm long, c. 1 mm thick; calyx tube
9−11 mm long, 4−6 mm wide, hirsute, hairs erect, surface visible on the limb; calyx
285

limb cup-shaped; calyx lobes 5, triangular at the base, becoming long-linear at the apex,
4−6 mm long, pubescent; corolla hypocrateriform, white to cream, the tube 1.5−2 cm
long, 6−7.5 mm wide at the throat, 2−3 mm wide at the base; non-speckled inside, outer
surface sparsely pubescent at the upper half, puberulous at the lower half, inner surface
glabrous; corolla lobes 5, ovate-oblong to ovate, 1.4−1.8 cm long, 0.4−1 cm wide,
outside pubescent on the uncovered half of the lobes, inside glabrous; anthers 5, fully
included within the tube (inserted at the upper ⅓), 6−7 mm long; style and stigma
12−16 mm long, included but reaching just below the corolla mouth and the level of or
above the anthers, style smooth, glabrous. Fruits ellipsoid, 4−5.5 cm long, 2.5−3.5 cm
wide, velvety hairy, drying smooth; fruit wall 1−1.5 mm thick; peduncle c. 1 cm long,
c. 1 mm thick. Mature seeds not seen.
DISTRIBUTION. Confined to the Eastern Highlands Prov. of Papua New
Guinea (Fig. 84).
1700 m alt.
HABITAT. Found in Castanopsis-Lithocarpus forest, sometimes on ridges, to
SPECIMENS EXAMINED.
PAPUA NEW GUINEA. EASTERN HIGHLANDS PROV.: Kainantu subdistrict.,
Kassam Pass, Brass 32331 (31 October 1959 FLO) (holotype L!), 1400 m, Forman
LAE 60368 D (25 June 1974 FRU) (BRI!, L!), 4200ft [1280 m], Vandenberg & Henty
NGF 35004 (24 January 1968 FRU) (BRI!, L!, SING!), 5500ft [1676 m], Streimann
NGF 47892 (11 July 1970 FRU) (L!).
6. “Rothmannia” lagunensis [combination needed].
Basionym: Gardenia lagunensis Merr., Philip. J. Sci., C 10 (1915) 110; Enum. Philip.
Fl. Pl. 3 (1923) 530.
286

Fig. 60. “Rothmannia kassamensis”. A, leafy branch with inflorescences. B, close-up of the lower leaf
surface with velvety hairs covering the midrib, secondary veins and entire surface. C, calyx tube with
long-linear lobes. D, longitudinal section of flower. E, outer surface of corolla lobe. F, fruit.
A–E from Brass 32331 (holotype, L), F from Forman LAE 60368 D (BRI).
287

NEOTYPE (to be designated): Ramos Bur. Sci. 14934, Philippines, Luzon, San
(Fig. 61)
Antonio Province, Laguna (June 1912) (L!).
Tree, to c. 25 m high, to 35 cm diameter. Bark smooth, grey-black; inner bark
light brown. Twigs pubescent when young, becoming glabrescent to glabrous when
old. Branch leaves typically in groups of 3’s, each trifoliate group based on a distal
node with only one leaf normally developed (the other vestigial or very much reduced
in size), and the proximal node with a pair of normal leaves. Stipules ovate, 2−3 mm
long, 3−5 mm wide, glabrous outside; apex acute, to 1 mm long. Young leaves on
shoot tips pubescent on both sides. Mature leaves with petiole 0.5−1.1 cm long, 1−1.5
mm thick, glabrous to scantily puberulous; lamina oblanceolate, lanceolate to elliptic,
6−12(−13.5) cm long, 2−6 cm wide, upper surface glabrous, lower surface scantily
pubescent becoming glabrous; leaf base cuneate, attenuate, equal to unequal; leaf apex
acuminate, acumen to 1 cm long; thin-coriaceous when dry; midrib on upper side flat,
scantily puberulous to pubescent, on lower side raised, glabrous; secondary veins
5−6(−7) pairs, on upper side flat, glabrous, on lower side slightly raised, scantily
puberulous to pubescent; tertiary venation visible as a much-branched network between
pairs of secondary veins. Inflorescences (bud): peduncle subsessile to 0.5 cm long,
1−1.5 mm thick, glabrous; habit compact, unbranched or with 1(−2) distinct branching
orders, rachis 0.2−0.5 cm long, primary branches along the rachis 1 pair, 0.2−0.5 cm
long; bracts ovate, 1.5−2 mm long, 1.5−2 mm wide, glabrous, hairy along the margin.
Flowers in buds 1−3(−5) per cyme; pedicels subobsolete to 1 mm long, 1−1.5 mm
thick; calyx tube 4−6 mm long, 3.5−4.5 mm wide, scantily puberulous to pubescent,
hairs appressed, surface visible; calyx limb cup-shaped; calyx lobes 5, triangular, 1−1.5
288

mm long, scantily puberulous; corolla hypocrateriform, the tube 7−8 cm long, c. 3 mm
wide at the throat, c. 1.5 mm wide at the base; non-speckled inside, outer surface
scantily puberulous at the upper half, inner surface glabrous; corolla lobes 5, oblong,
1−1.5 cm long, 0.25−0.3 cm wide, glabrous or with minute hairs sparsely distributed at
the margin or fringes of the lobes outside, glabrous inside; anthers 5, fully included
within the tube or only with tips slightly protruding (inserted at the upper ⅓), 12−14
mm long; style and stigma 75−85 mm long, included but reaching just below the
corolla mouth and the level of or above of the anthers, style smooth, glabrous. Fruits
globose to ellipsoid, 2.5−3.5(−5) cm long, 2.5−4 cm wide, glabrous, drying typically
smooth; fruit wall c. 1 mm thick; peduncle 0.4−0.5 cm long, 2−3 mm thick. Seeds
slightly to much flattened and broadly ovate or suborbicular to narrowly triangular,
8−10 mm × 6–7 mm.
NOTES. Merrill (in the Philipp. J. Sci., C 10 (1915) 110) had designated the
collection C.F. Baker 3176 as the type specimen of the basionym, Gardenia lagunensis.
However, the holotype of this collection which was deposited in PNH had been
destroyed during the Second World War. Attempts were made to locate the isotypes of
this collection from the herbaria at POM and NY, where C.F. Baker’s collections were
normally deposited (Stafleu & Cowan, 1976), but were futile. The search continued to
other herbaria in the United States, e.g., UC and A, but was again futile.
Based on Article 9, paragraph 9.6 of the International Code of Botanical
Nomenclature, Vienna Code (2006), the specimen from the Leiden herbarium of
Ramos’ collection, Bur. Sci 14945 from the Philippines, Luzon, San Antonio Province,
Laguna collected in June 1912 is to be designated as the neotype.
This species is very similar to “R. merrillii” but is readily distinguishable by
having glabrous corolla throats, pubescent at the upper half of the outer surface of the
corolla tube; slender corolla lobes 1−1.5 × 0.25−0.3 cm; style and stigma included but
289

Fig. 61. The proposed neotype specimen to be designated for “Rothmannia lagunensis”,
Ramos, Bur. Sci. 14934, Philippines, Luzon, San Antonio Province, Laguna (L).
290

eaching just below the corolla mouth and the level of or above the anthers whereas “R.
merrillii” has hairy corolla throats, a glabrous outer surface of the corolla tube; larger
corolla lobes, 2−4.5 × 0.5−1.5 cm; style and stigma included but reaching less than ⅔
way along the corolla tube and well below the anthers.
DISTRIBUTION. Known only from Luzon island of the Philippines (Fig. 68).
HABITAT. Recorded in Mixed Dipterocarp forest, to 300 m.
SPECIMENS EXAMINED.
PHILIPPINES. LUZON. Linutan, Loher 1481 A (sine datum FLO) (K!), Montalban,
Loher 1482 A (sine datum FLO) (K!); Camarines Sur, Alambra F.B. 28093 (July 1920
FLO) (K!); Quezon National Park, 25 km E of Lucena, 200–300 m, Soejarto et al.
7828 (12 July 1992 FRU) (L!); Tayabas, Bawan F.B. 24936 (December 1915 FRU)
(P!); Laguna, Conicosa F.B. 29461 (August 1923 FRU) (A!), Ramos 1401 (February
1913 FRU) (L!, SING!); Rizal, Ramos 2635 (May 1907 FRU) (BO!, L!); Mt Susong-
Dalaga, Ramos & Edano Bur. Sci. 29392 (August, 1917 FRU) (K!); San Antonio,
Ramos Bur. Sci. 14934 (June 1912 FRU) (neotype L!), Ramos Bur. Sci. 23845
(October 1915 FRU) (A!); Angat, Ramos Bur. Sci. 22297 (December 1914 FLO) (A!,
L!, PNH!, SING!).
7. “Rothmannia” leytensis [new species description needed].
TYPE (to be designated): Wenzel 641, Philippines, Leyte (8 March 1914 FLO)
(holotype A!; isotype BM!).
Gardenia negrosensis sensu Merr., Enum. Philip. Fl. Pl. 3 (1923) 530, pro parte quoad
(Fig. 62)
Wenzel 741, 641; Sandique 26596; non Merr. (1915).
291

Tree. Twigs velvety hairy when young, becoming sparsely pubescent to
glabrescent when old. Branch leaves typically in groups of 3’s, each trifoliate group
based on a distal node with only one leaf normally developed (the other vestigial or
very much reduced in size), and the proximal node with a pair of normal leaves.
Stipules ovate-triangular, 5−7 mm long, 5−7 mm wide, sparsely pubescent to
tomentose; apex acute to acuminate, c. 1 mm long. Young leaves on shoot tips velvety
hairy on both sides. Mature leaves with petiole 0.1−0.6 cm long, 2−3 mm thick,
pubescent to tomentose outside; lamina elliptic to obovate, 10−18 cm long, 4.5−8 cm
wide, upper surface glabrous to glabrescent, lower surface sparsely pubescent; leaf
base cuneate to symmetric; leaf apex acute to acuminate, acumen to 0.5 cm long; thin-
coriaceous when dry; midrib on upper side flat to slightly raised, glabrous, on lower
side raised, pubescent; secondary veins 7−10 pairs, on upper side flat to slightly
sunken, glabrous, on lower side raised, pubescent; tertiary venation visible as a much-
branched network between pairs of secondary veins. Inflorescences: peduncle 0.2−0.4
cm long, 2−3 mm thick, velvety hairy to tomentose; habit compact, with 1−2 distinct
branching orders, rachis 0.5−1 cm long, primary branches along the rachis 1−2 pairs,
0.4−0.6 cm long; bracts ovate, 2−3 mm long, 2−2.5 mm long, densely pubescent.
Flowers 4−9 per cyme; pedicels to c. 1 mm long, to c. 1 mm thick; calyx tube 5−7 mm
long, 3−4.5 mm wide, densely tomentose, hairs appressed to suberect, surface visible;
calyx limb cup-shaped; calyx lobes 5, triangular, 1−2.5 mm long, tomentose; corolla
hypocrateriform, the tube 6–8.2 cm long, 4−5 mm wide at the throat, 1−2 mm wide at
the base; non-speckled inside, outer surface sparsely pubescent, inner surface glabrous;
corolla lobes 5, elliptic to oblong, 1.2−2.5 cm long, 0.4−0.9 cm wide, outside pubescent
on the uncovered half of the lobes, inside glabrous; anthers 5, fully included within the
tube or only with tips slightly protruding (inserted at the upper ⅓), 11−14 mm long;
292

Fig. 62. “Rothmannia leytensis”. A, leafy branch with inflorescences. B, close-up of the lower leaf
surface. C, outer surface of calyx tube. D, outer structure of flower. E, longitudinal section of flower.
F, outer surface of corolla lobes. G, Fruit. H, seeds.
A–B from Sandique 26596 (K), C–F from Wenzel 641 (BM), G–H from Edano 41727 (A & L).
293

style and stigma 60−85 mm long, included but reaching just below the corolla mouth
and the level of or above the anthers, style smooth, glabrous. Fruits globose, 3−4 cm
long, 3−4 cm wide, sparsely pubescent when young, becoming glabrous when mature,
drying smooth; fruit wall c. 1 mm thick; peduncle 0.3−0.5 cm long, 3−4 mm thick.
Seeds slightly to much flattened and broadly ovate or suborbicular, 9−11 mm × 7−8
mm.
NOTES. Vegetatively, this species is very closely allied to “R.
pseudoternifolia” but is distinguishable by the reproductive characters, i.e., relatively
shorter peduncle and longer style and stigma complex (see Key to Malesian taxa
recognized in the Rothmannia complex in 8.1)
DISTRIBUTION. Endemic to the Philippines (Leyte).
SPECIMENS EXAMINED.
PHILIPPINES. No locality: Wenzel 741 (30 May 1914 FRU) (BM!). LEYTE:
Wenzel 641 (8 March 1914 FLO) (holotype A!; isotype BM!); Mt Abucayan, Edano
41727 (February 1923 FRU) (A!, L!, SING!), Sandique 26596 (March 1917 FLO) (K!).
8. “Rothmannia” macromera (Lauterb. & K.Schum.) Fagerl., Ark. Bot. 30 A (7)
(1943) 39.
Basionym: Randia macromera Lauterb. & K.Schum., Fl. Deutsch. Schutzgeb. Südsee
(1900) 562; Valeton, Bot. Jahrb. Syst. 60 (1926) 90 excl. var. uvarioides
Valeton (= “Rothmannia uvarioides” in the present work); Merrill & Perry, J.
Arnold Arbor. 25 (1944) 202.
NEOTYPE (to be designated): Schlechter 16215, Papua New Guinea, “Kaiser
(Fig. 63)
Wilhelmsland” (Leiden sheet no. Herb. Lugd. Bat. 927.320-165) (29 June 1907
FLO) (L!).
294

Small tree, to 12 m high. Bark grey-brown; inner bark straw. Sapwood straw
to red. Twigs sparsely pubescent to glabrous. Branch leaves typically in groups of 3’s,
each trifoliate group based on a distal node with only one leaf normally developed (the
other vestigial or very much reduced in size), and the proximal node with a pair of
normal leaves. Stipules broadly triangular, 5−8 mm long, 7−10 mm wide, sparsely
pubescent to glabrescent outside; apex cuspidate to acuminate, 2−8 mm long. Young
leaves on shoot tips velvety hairy on both sides. Mature leaves with petiole
subobsolete to 0.2 cm long, 3−4 mm thick, hairy; lamina broadly elliptic, oblanceolate
to obovate, 18−30(−35) cm long, (8−)9−13 cm wide, upper surface glabrous, lower
surface densely velvety hairy; leaf base rounded to subcordate; leaf apex short-
acuminate, acumen c. 0.7 cm long; when dry thick-coriaceous; midrib on upper side
flattened to slightly sunken, glabrous, on lower side distinctly raised, velvety hairy;
secondary veins 6−10 pairs, on upper side flattened to slightly sunken, glabrous, on
lower side raised, velvety hairy; tertiary venation visible as a much-branch network
between pairs of secondary veins. Inflorescences in buds: peduncle 0.1−0.2 cm long,
4−5 mm thick, pubescent; habit compact, with 2−3 distinct branching orders, rachis
0.5−1 cm long, primary branches along the rachis 2−3 pairs, 0.1−0.2(−0.4) cm long;
bracts ovate, 4−6 mm long, 2.5−5 mm wide, sparsely pubescent. Flowers 11−24 per
cyme; pedicels sessile to 1 mm long, 1–1.5 mm thick; calyx tube 4−5 mm long, 3−4
mm wide, velvety hairy to pubescent, hairs appressed to suberect, surface visible; calyx
limb cup-shaped; calyx lobes 5, subulate, (4−)6−10 mm long, pubescent; corolla
hypocrateriform, white, the tube 1–1.5 cm long, 3 mm wide at the throat, 2.5 mm wide
at the base; non-speckled inside, outer surface densely pubescent, inner surface
glabrous; corolla lobes 5–6, narrowly elliptic, 0.8−1.4 cm long, 0.3−0.4 cm wide,
295

outside densely hairy on the uncovered half of the lobes, inside glabrous; anthers 6,
fully included within the tube (inserted at the upper ⅓), 8−10 mm long; style and
stigma 11−20 mm long, included but reaching just below the corolla mouth and near
the anthers, style smooth, glabrous, stigma with ridges or grooves. Fruits spindled-
shaped, 6–8.5 cm × 2.5–3.2 cm wide. Seeds unknown.
NOTES. In the original publication of the basionym, Randia macromera in the
Fl. Deutsch. Schutzgeb. Südsee (1900), Lauterbach and Schumann cited two
specimens, Lauterbach 1286 (flowering) and Lauterbach 1524 (flowering and fruiting),
both from Kaiser Wilhelmsland (formerly the German colony of the northeastern part
of mainland New Guinea), and collected in 1890. Both these specimens (together with
their duplicates) were not located. According to Van Steenis-Kruseman (1950) in
Cyclopaedia of Collectors Vol. 1, collections from Lauterbach’s trip to Kaiser
Wilhelmsland in 1890 were deposited in the Berlin Herbarium. There is a possibility
that these specimens were destroyed during the Second World War. Attempts were
made to locate the syntypes in the Berlin Herbarium and other herbaria where
Lauterbach’s collection were believed to have been deposited, i.e., BO, BM, K, L, and
the US (Van Steenis-Kruseman 1950). However, the search was futile.
A later publication by Valeton in Die Rubiaceae von Papuasien (1926) in
Engler’s Bot. Jahrb. Syst. 60 pg. 90, listed again two of the specimens (i.e., Lauterbach
1286 and Lauterbach 1524) cited by Lauterbach & Schumann (1900) and three other
collections under Randia macromera. One of the specimens cited in this account was
Schlechter 16215, collected on 29 June 1907 from Kaiser Wilhelmsland, which was
located at L. This is the only specimen that was possible to find and also matches the
original description. As such, based on Article 9, paragraph 9.6 of the International
Code of Botanical Nomenclature, Vienna Code (2006), the specimen at Leiden (Herb.
Ludg. Bat. 927.320-165) is to be designated as the neotype for this species.
296

Fig. 63. The proposed neotype specimen to be designated for Rothmannia macromera
(Lauterb. & K. Schum.) Fagerl., Schlechter 16215, Papua New Guinea, Kaiser
Wilhelmsland (L).
297

The fruiting material of this species has not been seen, but according to the
description by Lauterbach & Schumann (1900) and Valeton (1926), the fruits are large,
6–8.5 × 2.5–3.2 cm, and spindle-shaped with a thick beak, c. 1cm long above.
DISTRIBUTION. Found so far only in Papua New Guinea (Morobe Province)
(Fig. 84).
HABITAT. Recorded mostly on Araucaria forest, on slope, to 1060 m alt.
SPECIMENS EXAMINED.
PAPUA NEW GUINEA. Kaiser Wilhelmsland, 200 m, Schlechter 16215 (29 June
1907 FLO) (neotype L!). MOROBE PROV.: Crooked Creek, about 4 miles SW of
Bulolo, Araucaria forest on slope, 2600 ft [790 m], Hartley TGH 10882 (7 November
1962 FLO) (L!), Wau, Crooked Creek, Bulolo, logging area, Araucaria forest, 2450 ft
[745 m], Galore NGF 15772 (October 1962 FLO) (L!); Bulolo Valley, 3500 ft [1060
m], Womersley & Jones NGF 8814 (6 June 1956 FLO) (BRI!, L!, SING!).
9. “Rothmannia” macrophylla (R.Br. ex Hook.f.) Bremek., K. Ned. Akad. van
Wet.–P 60, 1 (1957) 3; Wong, Malay. Nat. J. 38 (1984) 46; Wong, Tree Fl. Malaya
(1989) 324; Kochummen, Tree Fl. Of Pasoh Forest (1997) 385.
Basionym: Randia macrophylla R. Br. ex Hook.f., Fl. Brit. India 3 (1882) 114; King
& Gamble, J. As. Soc. Beng. 72 (1903) 215; Ridley, Fl. Malay Penins. 2 (1923)
78; Corner, Wayside Trees of Malaya 2 (1988) 648.
LECTOTYPE (designated by Wong, 1984): Penang, Wallich, Cat. 8304A (1822
(Fig. 65)
FLO) (K!)
Treelets or shrublike plants, sometimes small trees, frequently with horizontal
stem portions on or below the ground, producing erect above-ground shoots reaching 5
298

m (rarely 8 m) high, typically to 4 cm diameter. Bark brownish black. Twigs scabrid
to hispid. Branch leaves in pairs at the proximal node and completely vestigial at the
distal node (often microscopic in size). Stipules broadly triangular, 15−17 mm long, 7–
8 mm wide, sparsely hispid to pubescent outside; apex long-acuminate, 8–12 mm long.
Mature leaves with petiole subobsolete to c. 1.2 cm long, 2–3 mm thick, hispid to
pubescent; lamina broadly oblanceolate, narrowly elliptic to oblong, (14–)18−35 cm
long, 3.5–13.5 cm wide, upper surface sparsely pubescent, glabrescent to glabrous,
lower surface scabrid to hispid; leaf base symmetric, cuneate to sometimes unequal;
leaf apex acute to acuminate, acumen 0.5–1 cm long; chartaceous to thin-coriaceous
when dry; midrib on upper side slightly raised, sparsely pubescent, on lower side
distinctly raised, pubescent, velvety hairy to hispid; secondary veins 10–21 pairs, on
upper side flat to sunken, glabrous to sparsely pubescent, on lower side raised,
pubescent to hispid; tertiary venation visible as a much-branched network between
pairs of secondary veins. Inflorescence: peduncle subobsolete, to 0.2 cm long, 3−4
mm thick, pubescent to velvety hairy; habit compact, a much reduced cyme, 1–3-
flowered; bracts broadly ovate, 7–8 mm long, 3−5 mm wide, pubescent to hispid hairy.
Flowers bisexual; pedicels very short, to 4 mm long, 3−4 mm thick; calyx tube 6–15
mm long, 5–7 mm wide, pubescent, velvety to hispid, hairs suberect to erect, surface
visible; calyx limb cup-shaped; calyx lobes 5–6, narrowly triangular, 12–30 mm long,
pubescent to hispid hairy; corolla infundibular, the tube white with maroon tinged at the
base on the outside, the tube 9–15 cm long, (20–)30–55 mm wide at the throat, 3–5 mm
wide at the base; speckled inside, outer surface pubescent to hispid hairy, inner surface
with hairy zone covering part of the tube from below the level of the anthers; corolla
lobes 5, ovate with acute apex, 2.5–5.5 cm long, 1.5–2.5 cm wide, outside pubescent on
the uncovered half of the lobes, inside glabrous; anthers 5, fully included within the
tube (inserted at the upper ⅓), 20–25 mm long; stigma fusiform, slightly ribbed when
299

dried, style and stigma 80–140 mm long, included but reaching just below the corolla
mouth and the level of the anthers, style smooth, glabrous. Fruits ellipsoid, 4–7 cm
long, 2−3.5 cm wide, hispid when young becoming pubescent; fruit wall 1−2 mm thick;
peduncle 0.5−3 cm long, 3−5 mm thick. Seeds ellipsoid, 5–8.5 × 6–8 mm.
NOTES. Although Robert Brown's name Rothmannia macrophylla R.Br. was
used in Wallich's Catalogue (no. 8304), this is not valid; Fagerlind (1943) had also
accepted this taxon as a species of Rothmannia prior to Bremekamp's (1957)
combination.
Ridley (1923) had thought that it is closely allied to Randia stenopetala
(=Kochummenia stenopetala) as both shared a similar habit (shrub) and membranous,
more-or-less hairy leaves, as well as conspicuous flowers, and so he placed them in the
same section, Randia sect. Grandiflorae. Wong (1984) pointed out that although the
infundibular corolla shape of “R. macrophylla” is very similar to that of the type
species of Euclinia (E. longiflora), the latter differs from Rothmannia species in having
tetrad pollen grains instead of single grains; chaffy instead of firm stipules; and leaves
clustering at the ends of twigs which are simultaneously deciduous instead of leaves at
the ends of twigs and not deciduous.
It is considered as an aberrant taxon of Rothmannia because of a number of
distinct characteristics that do not conform to the other Rothmannia species. The
primary branches have only 2 nodes each, terminated by an inflorescence, without
subsequent higher-order branching (in the other Rothmannia species, the primary
branch and successive higher-order modular units each have only 2 nodes and a
terminal inflorescence, with new modular units developing from either one or both leaf
axils at the proximal node of the previous module). The branch leaves are never in
groups of 3's, the distal branch node having both vestigial (often microscopic in size),
300

and the proximal node with a pair of normal leaves (in the other Rothmannia species,
the leaves are typically in groups of 3’s, each trifoliate group based on a distal node
with only one leaf normally developed, the other vestigial or reduced, and a proximal
node with a pair of normal leaves). Its anthers are 20–25 mm long (less than 20 mm
long in the other Rothmannia species from Malesia). The pollen grains have a wide
ectoaperture, 7 µm or more (less than 5.5 µm in the other Rothmannia species).
VERNACULAR NAMES. Indonesia: kelalui (Sumatra); Peninsular Malaysia:
kacubong hutan, pecah pinggan, kayu tabat, pokok kumatan (Malay); moh-jelu
(Semelai); hidong babi, terong hantu, kayu mogang (Temuan); segerang (Jakun).
DISTRIBUTION. Endemic to West Malesia. It is common in all states in the
Peninsular Malaysia. Also found in Singapore and Sumatra (Jambi, Central Sumatra,
Riau and Riau Islands Provinces) (Fig. 64). Cultivated at the Bogor Botanic Gardens,
Java, Indonesia.
HABITAT. It is a very common understorey component of the lowland
primary forest, often seen being trampled or snapped by wild pigs for construction of
their breeding nests (Ickes et al. 2003). Common in primary lowland to hill dipterocarp
forest, in disturbed or secondary forests, also recorded in swampy areas, to 1200 m alt.
USES. In Peninsular Malaysia, it has been documented by Burkill (1966) that
the juice was used by the Jakun (a native tribe) for blackening their teeth. As the
species has large showy flowers, it may be a good prospect for ornamental purposes
(Lemmens & Wulijarni-Soetjipto, 1992).
SPECIMENS EXAMINED.
PENINSULAR MALAYSIA. KEDAH: Sik, Bukit Enggang Forest Reserve, Compt.
17, 150 m, Zainudin et al. 4481 (13 June 1993 FLO/FRU) (UKMB!); Mt Bintang F.R.,
500 ft [152m], Sidek S 270 (4 August 1968 FLO) (SING!); Kedah Peak, Ridley s.n.
(June 1893 FLO) (SING!), 2800–4000 ft [854–1219 m], Robinson & Kloss 6179
301

(December 1915 FLO/FRU) (K!, SING!), lower slope of Kedah Peak, sine coll s.n.
(June 1893 FLO) (SING!); Mt Baling, Best SFN 21210 (31 October 1929 FLO)
(SING!). PERAK: No locality. Mantly's collection s.n. (sine datum FLO) (SING!);
500–800 ft [152–244 m], Dr King's Collector 10356 (July 1886 FLO) (L!); Larut,
Wray 1632 (sine datum FLO/FRU) (BM!, U!), Larut, Briah, Wray 4214 (December
1892 FLO) (K!, SING!); Maxwell's Hill, Wray 3232 (October 1888 FLO) (SING!);
Chang Kat Mentri, Sg Perak, Boden Kloss SFN 6524 (September 1918 FLO) (K!);
Tapah, Md. Haniff 14262 (17 September 1924 FLO) (SING!); Trolak F.R., Chelliah
KEP 104693 (23 March 1967 FRU) (KEP!); Lumut, Bryant s.n. (November 1892 FLO)
(SING!); Dindings, Mt Tungal, Ridley s.n. (July 1894 FLO) (SING!). PENANG: No
locality. Wallich 8304 A (1822 FLO/FRU) (lectotype K!). Wallich 8304C (sine datum
FLO) (P!). Teluk Bahang, Curtis 966 (8 March 1905 FLO) (SING!), Curtis 966 (July
1893 FLO) (SING!); Penara Bukit, Curtis 966 (August 1889 FLO) (SING!); Penang
Hill, Ridley s.n. (June 1891 FLO) (SING!), 2500 ft [762 m], Ridley s.n. (June 1898
FLO) (SING!). SELANGOR: Ulu Kuang, Provencher Nr.T. 18 (December 1971
FLO) (KLU!, SING!); Ulu Gombak, Ridley 152 (22 March 1905 FLO) (BM!), Gombak
F.R., Poore 1038 (28 March 1962 FLO) (KLU!); Sg Buloh F.R., Kasim & Rahim 5021
(1 October 1977 FLO) (UKMB!), T. & P. 191 (2 October 1968 FLO) (L!, SING!),
Hardial & Sidek 446 (19 January 1966 FLO) (L!, SING!), 50 ft [15 m], Kochummen
94638 (24 October 1962 FLO/FRU) (KEP!), Sg Buluh F.R., near Kg Cubit,
Kochummen FRI 29141 (23 October 1980 FLO/FRU) (KEP!, L!, SAN!); Bangi,
Universiti Kebangsaan Malaysia campus, Kasim 5021 (19 December 1972 ST)
(UKMB!), Zainudin & Rahim 3184 (15 May 1973 ST) (UKMB!), Razali 3618 (16
November 1973 FRU) (UKMB!), 250 ft [76 m], Zainudin & Rahim 1472 (18 May 1973
ST) (UKMB!), Fernarium, Pereira, Noorsiha & Razali JTP 631 (18 June 1999 FLO)
(SAN!), Bangi F.R., 75–100 m, Balgooy 2089 (10 September 1974 FLO) (L!); Ulu
302

Selangor, 300 ft [91 m], Wyatt-Smith KEP 63729 (27 February 1947 FRU) (KEP!),
Hulu Selangor, Kerling, Gading F.R., Plot 65, Pereira, Noorsiha & Asri JTP 645 (15
July 1999 ST) (SAN!), JTP 646 A (15 July 1999 ST) (SAN!), JTP 646 C (15 July 1999
ST) (SAN!), JTP 648 (15 July 1999 ST) (SAN!), Plot 61, Pereira, Noorsiha & Asri JTP
647 (15 July 1999 FLO) (SAN!); Semenyih, Hume FMS 7833 (9 July 1921 FLO)
(SING!); Semenyih, Sg Tekala, Tan s.n. (23 November 2005 FLO/FRU) (SAN!); Ulu
Selangor, Kanching F.R., Plot 47, Aziz KEP 65548 (29 July 1949 ST) (KEP!); Ulu
Langat, K. Pansom, Gadoh sub Millard KL 869 (9 October 1958 FRU) (KEP!), Mt
Linyu, Gadoh sub Millard KL 1213 (25 November 1958 ST) (KEP!); Batu Tiga, Arcem
s.n. (1 July 1902 FLO) (SING!). KUALA LUMPUR: Ridley 10484 (2 February 1899
FLO) (SING!), Curtis s.n. (February 1890 FLO) (SING!); Kepong, Symington 20527
(22 August 1929 FLO) (SING!), Kepong, FRIM, Compt. 31, Wyatt-Smith KEP 60607
(30 December 1946 FLO) (KEP!, L!), FRIM, Jalan Kapur, Pereira et al. JTP 846 (2
November 2001 FLO) (SAN!), Kepong Plantations, Pawanche 13446 (19 March 1928
FLO) (SING!). NEGERI SEMBILAN: Mt Berumbun, Alvines 2022 (20 June 1885
FLO) (SING!); Tampin, Burkill SFN 3508 (30 July 1918 FLO) (SING!), Burkill 4963
(8 August 1919 FLO) (SING!); Ulu Rembau, Md. Nur SFN 11773 (29 November 1923
FRU) (SING!); Pasoh F.R., Mat Asri FRI 21855 (30 December 1974 FLO) (KEP!);
Port Dickson, Sg Menyala F.R., Sinclair SFN 40163 (13 November 1953 FLO)
(SING!), Vethevelu FRI 32930 (20 December 1983 FLO) (KEP!); Mt Angsi, 1500 ft
[457 m], Loh FRI 17347 (20 February 1971 FRU) (KEP!). MALACCA: No locality.
sine coll. 43 (June 1888 FLO) (SING!), Alvines s.n. (sine datum FRU) (SING!), Alvines
16 (sine datum FLO/FRU) (SING!), Alvines 32 (sine datum FRU) (SING!), Alvines 156
(4 November 1894 FLO) (SING!), Alvines 254 (17 January 1885 ST) (SING!), Alvines
303 (15 December 1894 FLO) (SING!), Alvines 734 (5 November 1894 FLO) (SING!),
Alvines 1053 (sine datum FRU) (SING!), Burkill 440 (13 February 1914 FLO)
303

(SING!), Cuming 2365 (sine datum FLO) (BM!), Dr King's Collector Griffith 2822
(1845 FLO) (K!), Maingay 1301 (6 July 1865 FLO) (L!), Maingay 1301 A (22 June
1868 FLO) (K!). Bukit Bruang, Goodenough 1334 (July 1893 FRU) (SING!); Sg
Udang, Goodenough 1349 (August 1893 FLO) (SING!). PAHANG: Endau-Rompin,
near Sg Selindang, Route 1, Latiff, Nik & Laily ALM 3194 (17 May 1989 FRU)
(UKMB!); Rompin, Mohammad FMS 17180 (17 January 1930 FLO) (SING!); near
Bentong, Sabai Estate, 400 ft [122 m], Mohd. Shah 175 (27 January 1958 FLO)
(SING!); Lesong F.R., 250 ft [76 m], Samsuri & Ahmad SA 428 (17 February 1971
FLO) (SING!); 6th mile S of Kg Aur, 400 ft [122 m], Samsuri & Ahmad SA 486 (18
February 1971 FLO) (SING!); Tasik Bera, Gianno KLU 501 (18 December 1980 FLO)
(L!); Raub, Mohamad 20339 (1 November 1929 FRU) (SING!); near Raub, 400 ft [122
m], Md. Haniff 16907 (11 November 1924 FLO) (SING!); Sg Pahang, Fort Iskandar,
Woods Phytochemical Survey of Malaya Herbarium 1708 (28 February 1959 FLO)
(KEP!, L!, SING!); Jerantut, Remy & Teo KL 3985 (8 January 1991 FLO) (KEP!,
SING!); Kuala Jelai, Ulu Betok, Machado s.n. (13 March 1905 FLO) (SING!); Chini
Lake, forest near the lake, Low et al. LYW 57 (12 April 2005 FRU) (SAN!).
KELANTAN: Sg Keteh, Md. Nur & Foxworthy SFN 12038 (2 October 1924 FRU)
(SING!); Kemahang F.R., Compt. 16, Chelliah FRI 6536 (1 July 1968 FRU) (KEP!);
Sg Tapah, Ulu Sg Aring, Cockburn FRI 7179 (22 September 1967 FRU) (KEP!).
TERENGGANU: No locality. Rostado s.n. (1 February 1904 FLO) (SING!).
Dungun, Soepadmo KLU 9130 (sine datum FRU) (KLU!); Dungun, Hutan Jambu
Bongkok, Kasim s.n. (29 August 1964 FLO) (KLU!), Kasim et al. MK 1612 (29 May
1979 ST) (UKMB!), JOHORE: Mt Pulai, Ridley 3724 (1892 FLO) (BM!, SING!);
9th mile, Mawai-Kuala Sedili new road, Sinclair 10565 (24 January 1961 FLO) (L!,
SING!); Kluang F.R., Compt. 79, 100 ft [31 m], Ng KEP 97951 (30 January 1966 FLO)
(KEP!, L!); Batu Pahat, Banang F.R., Compt. 10, 200 ft [61 m], Chelliah KEP 98238
304

(7 October 1966 FLO) (KEP!, L!); Rawai, Sg Kayu, Corner s.n. (14 April 1935 FRU)
(SING!); Kota Tinggi, Druse s.n. (February 1916 FLO) (SING!), Ridley s.n. (sine
datum FLO) (SING!), Kota Tinggi, Kuala Sedili, Kadim & M. Noor 165 (24 June 1959
FRI) (SING!), Kota Tinggi, Nam Heng Rubber Estate, Teruya 320 (17 December 1926
FLO) (SING!), Kota Tinggi, above waterfall reservoir, Pearce 343 (26 November 1980
FLO) (KLU!); Kahang, Hardial & Samsuri HS 1069 (3 April 1973 FLO) (SING!); Mt
Panti, 800 ft [244 m], Corner s.n. (20 January 1936 FLO) (SING!). SINGAPORE.
No locality. Hullet 117 (June 1884 FLO) (K!), Hullet 339 (6 May 1886 FLO) (SING!),
Gaudichaud 62 (1836–37 FLO) (P!), Mohd Shah & Sidek MS 4068 (5 March 1981
FLO) (SING!), Ridley s.n. (April 1905 FLO) (BM!). Changi, Goodenough 2855a (5
March 1905 FLO) (SING!); Nee Soon, Samsuri, Ganesan, Leong & Gwee NES 250
(26 August 2003 ST) (SING!); Bukit Timah Nature Reserve, Hardial 358 (7 January
1965 FRU) (SING!), Lai LJ 468 (20 June 1995 FLO) (SING!), Who, Phuah & Hardie
NRS 1268 (6 September 1992 FRU) (SING!), steps to summit, Tang & Haji 1125 (30
November 1995 FLO) (SING!), 50 m, Maxwell 77–94 (24 February 1977 FLO) (L!);
MacRitchie Reservoir, Maxwell 78–60 (31 March 1968 FLO) (L!), Maxwell 79–12 (7
November 1979 FLO) (L!), South side of MacRitchie, Sinclair s.n. (8 December 1950
FLO) (L!); Ang Mo Kio, Ridley s.n. (March 1889 FLO) (SING!); Chan Chu Kang,
Ridley s.n. (sine datum FLO) (SING!). INDONESIA. SUMATRA. JAMBI PROV.:
S Kefasung, Sg Tabir, Laumonier YL 791 (1 March 1980 FLO) (L!), Laumonier YL
814 (March 1980 FRU) (L!); Sg Tabir, Alas Kusuma, Camp Lubuk INTAN,
Laumonier YL 851 (February 1980 FLO) (L!). RIAU PROV.: Tigapuluh Mountains,
5 km W of Talanglakat on Rengat-Jambi Rd., Bukit Karampal area, 100 m, Burley,
Tukirin et al. 1430 (13 November 1988 FLO/FRU) (BO!, KEP!, L!, SING!). RIAU
ISLANDS PROV.: N of Dalu Dalu, River Mahato Kiri area, 100 m, Laumonier YL
6246 (24 March 1984 FRU) (L!); Lingga Archipelago, Pulau Singkep, Weq Dabo, 50
305

m, Bunnemeijer 7250 (4 August 1919 FLO) (BO!). C SUMATRA PROV.: Berapit,
Inragiri Uplands, Buwalda 6550 (14 April 1939 FLO) (L!); Ijerman Expedition, 40 m,
Koorders Kds. 22385 (7 March 1891 ST) (BO!).
Cultivated. INDONESIA. JAVA. Bogor Botanic Garden (cultivated in VA.2a),
Kuswata 14 (9 November 1959 FLO) (KEP!).
Fig. 64. Distribution of “Rothmannia macrophylla”.
306
MALAY
PENINSULA
SUMATRA

Fig. 65. The lectotype specimen of Rothmannia macrophylla (R.Br. ex Hook.f.) Bremek., Wallich,
Cat. 8304A, Penang (K).
307

10. “Rothmannia” merrillii [combination needed].
Basionym: Gardenia merrillii Elmer, Leafl. Philipp. Bot. 1 (1906) 5; Leafl. Philipp.
Bot. 4 (1912) 1330; Gibbs, Journ. Linn. Soc. Bot. 42 (1914) 94; Merr., Enum.
Philip. Fl. Pl. 3 (1923) 530; Merr., Philipp. J. Sci. 29 (1926) 421, pro parte excl.
Celebes specimens; Merr., Bibl. En. Born. Pl., J. Straits Branch Roy. Asiat.
Soc., Sp. No. (1921) 564 pro parte quoad Gibbs 4340, Foxworthy 600;
Masamune, En. Phan. Born. (1942) 682.
LECTOTYPE (to be designated): Merrill 669, Philippines, Island of Calomian,
Culion (February 1903 FLO) (lectotype US!) (isolectotypes A!, SING!, NY!).
Randia keithii C.E.C. Fisch., Bull. Misc. Inform. Kew (1932) 180; Masamune, En.
Phan. Born. (1942) 708.
TYPE: Keith 1215, North Borneo, Sandakan, 500 ft [152 m] (20 February 1929
FLO) (holotype K!).
Rothmannia keithii (C.E. Fischer) Bremek., Proc. K. Ned. Akad. van Wet.–P 60 (1957)
6.
(Fig. 67)
Treelet to small tree, to 6(−12) m high, to 10 cm diameter. Bark smooth to
finely fissured, grey-black to dark brown; inner bark brownish to greenish yellow.
Sapwood whitish to white-orange. Twigs glabrous. Branch leaves typically in groups
of 3’s, each trifoliate group based on a distal node with only one leaf normally
developed (the other vestigial or very much reduced in size), and the proximal node
with a pair of normal leaves. Stipules shortly ovate-triangular, 1−3 mm long, 3–5(–6)
mm wide, glabrous to sometimes scantily pubescent outside; apex cuspidate, 0.5−1 mm
long. Young leaves on shoot tips scantily pubescent, especially on the lower surface.
Mature leaves with petiole (0.1−)0.3−1.3 cm long, 1−1.5 mm thick, glabrous to scantily
308

puberulous; lamina narrowly to broadly elliptic, elliptic-oblong, oblanceolate to
obovate, 5−20 cm long, (1.5−)4–8(−10) cm wide, upper surface glabrous, lower surface
typically glabrous, sometimes sparsely puberulous; leaf base attenuate, cuneate to
sometimes unequal; leaf apex acute to obtuse, acumen to 0.3 cm long; thin to thick-
coriaceous when dry; midrib on upper side flat, glabrous, on lower side raised, glabrous
to occasionally scantily puberulous; secondary veins 5−8(−9) pairs, on upper side flat,
glabrous, on lower side raised, glabrous to occasionally puberulous; tertiary venation
visible as simple connections between pairs of secondary veins to almost invisible.
Inflorescences: peduncle 0.15−0.5 cm long, 2−3 mm thick, glabrous to sparsely
puberulous; habit compact, unbranched to with 1(−2) distinct branching orders, rachis
0.1–0.5 cm long, primary branches along the rachis 1(−2) pairs, 0.1–0.5 cm long; bracts
ovate, 1−2.5 mm long, 1−2 mm wide, glabrous to sparsely puberulous. Flowers
1−5(−9) per cyme; pedicels subobsolete to 1 mm long, c. 1 mm thick; calyx tube
4−6(−10) mm long, 3−6 mm wide, glabrous to scantily puberulous, hairs appressed,
surface visible; calyx cup-shaped; calyx lobes (5−)6−8(−10), short-triangular, 0.5−2.5
mm long, glabrous to sparsely puberulous; corolla hypocrateriform, white, the tube
3.5−7.5(−12) cm long, 5−9 mm wide at the throat, 2−3 mm wide at the base; non-
speckled inside, outer surface glabrous, inner surface with hairy zone covering part or
all of the tube from the corolla mouth downwards; corolla lobes (5−)6−8, lanceolate,
oblong-elliptic to obovate, 2−4.5 cm long, 0.5−1(−1.5) cm wide, outside glabrous or
with minute hairs sparsely distributed at the margin or fringes of the lobes, inside
glabrous; anthers 6−10, fully included within the tube (inserted at the upper ⅓), 6−10
mm long; style and stigma 10−34 mm long, included but reaching less than ⅔ way
along the corolla tube and well below the level of the anthers, style smooth, glabrous.
Fruits globose, 2.5−4.5(−5) cm long, 2.5−4.5(−5) cm wide, glabrous, drying smooth to
309

wrinkled; fruit wall 1−2 mm thick; peduncle sessile to 0.3 cm long, 2−4 mm thick.
Seeds slightly to much flattened and broadly ovate or suborbicular to narrowly
triangular, 10−11 mm × 8−10 mm.
NOTES. In 1906, Elmer published the basionym, Gardenia merrillii in Leafl.
Philipp. Bot. 1, pg. 5 and cited two specimens, Merrill 669 from Culion, collected in
February 1903 and Garcia FB 1226 from Mindoro, collected in January 1903. As there
was no clear type designated, one of the two syntypes is to be designated as the
lectotype. Most of Merrill’s specimens were deposited at the Philippine National
Herbarium (PNH) (Van Steenis-Kruseman, 1950). These may have been destroyed
during the Second World War. Attempts to locate the holotype were futile.
Based on Article 9, paragraph 9.2 of the International Code of Botanical
Nomenclature, Vienna Code (2006), one of the duplicates of Merrill 669 is to be
designated as the lectotype, i.e., from the United States National Herbarium,
Smithsonian Institution (US).
This species is very variable in terms of leaf size; corolla tube length; hairiness
on the nerves (glabrous to pubescent), peduncle (glabrous to pubescent), bracteoles
(glabrous to pubescent), domatia (glabrous to pubescent) and number of flowers per
cyme.
The Sabah specimens tend to have fewer flowers, usually 1, rarely 3 per cyme.
However, the Philippine specimens vary from solitary flowered to 3–5-flowered. The
flowers are commonly arranged as 3−5 flowers per cyme, to a maximum of 9 flowers.
Furthermore, the corolla tube length typically varies from 3.5 cm to 7.5 cm long,
however, in one specimen collected by Ridsdale, from Palanan, N.E. Luzon (ISU 93):
the corolla tube is extremely long 10−12 cm long.
310

VERNACULAR NAMES. Sabah: kelimpong (Kudat); sarakat (Dusun-
Rungus); cempaka putih (Malay). Philippines: rosal-gobat (Mindoro); Bagaoi, niog-
niog (Bisaya)
DISTRIBUTION. Endemic to the Philippines and Borneo (Sabah) (Fig. 66).
HABITAT. Primary to secondary forests, including kerangas forest, sandstone
hills, coastal forest and on ultrabasic substrate, and calcareous outcrops, to 850 m alt.
SPECIMENS EXAMINED.
PHILIPPINES. No locality: Marche B 119 (10 January 1885 FLO) (P!); Vidal 1447
(sine datum FRU) (L!). LUZON. Tayabas Lucena, Merrill 2888 (June 1903 FLO)
(BM!, K!); Isabela Prov., Palanan, Diguyo, 50 m, Ridsdale, Dejan & Baquiran ISU 93
(22 April 1991 FLO) (L!, SAN!). CAGAYAN PROV.: Bernardo F.B. 27085 (October
1917 FRU) (A!, K!). MINDORO. No locality: Merritt F.B. 8568 (January 1908
FRU) (SING!); Pinamalayan, Ramos Bur. Sci. 40890 (June 1922 FRU) (L!);
Bongabon & Pinamalayan, Maliwanag 221 (5 February–5 April 1941 FLO) (PNH!);
Baco, Merrill 1195 (January 1903 FLO) (K!), Merrill 1226 (January 1903 FLO) (A!,
K!, SING!, NY!), Baco river, Mcgregor 137 (April to May 1905 FLO) (K!); Paluan,
Ramos Bur. Sci. 39474 (April 1921 FRU) (SING!); Mansalay, Paniza PNH 9426 (18
April 1948 FRU) (A!), Mansalay, Bo, Manaul, Sulit PNH 17140 (30 December 1952
FLO) (L!); N slope of Mt Alinyaban, 5 km SW of Puerto Galera, Burley 128 (24 April
1987 FLO) (K!), Puerto Galera Bay, Santos 5165 (16 April 1952 FLO) (L!), Mt
Yagaw, 350 m, Sulit & Conklin PNH 16774 (28 November 1952 FRU) (L!, PNH!).
Golo Island, Merrill PNH 11540 (August 1922 FLO/FRU) (A!, L!, SING!).
PALAWAN: No locality. Agama PNH 21618 (May 1914 FRU) (P!); Curran 3517
(January 1906 ST) (L!); Escritor Bur. Sci. 21571 (August 1913 ST) (P!); Aborlan,
Sagpangan, along river, Celestino & J. Ramos PNH 23090 (16 May 1955 FLO) (L!,
PNH!); about 4 km off Aborlan-Inagawan main road, 50–100 m, Soejarto et al. 6608
311

(15 April 1989 FLO) (L!); Iraan Mountains, Sulit PNH 12549 (May–June 1950 FLO)
(L!), Km 112, Pinagbatuan, Mendoza & Espiritu PNH 91187 (27 August 1964 FLO)
(PNH!, SAN!); St. Paul's Bay, Cabayungan Village, 40 m, Podzorski SMHI 2081 (10
May 1984 FLO) (KEP!, PNH!, SAN!); Puerto Princesa, c. 36 km north of the city, 40
m, sine coll. SMHI 1063 (11 May 1984 FRU) (L!); Mt Pulgar, Elmer 13126 (April
1911 FRU) (L!); Irawan River valley head, 150 m Ridsdale SMHI 84 (15 March 1984
FLO) (L!), Ridsdale SMHI 98, (16 March 1984 FLO) (KEP!, PNH!, SAN!); Mt
Beaufort, 150–170 m, Ridsdale SMHI 1689 (15 May 1984 FRU) (L!); N of Tagburos,
c. km 17, N of Puerto Princessa, low forest on ultrabasic, Ridsdale 1096 (27 October
1985 FRU) (L!); Vidal 1447 (sine datum FRU) (L!), Babuyan, Edano 266 (16 March
1947 ST) (SING!), Km 36, Puerto Princesa to Roxas, 10 m, Dransfield SMHI 1237 (12
May 1984 FLO/FRU) (KEP!, K!, L!, SAN!); Narra, Victoria Peaks, Trident Mining
Company concession area, partially damaged steep-slope forest on ultrabasic soils,
300–450 m, Ridsdale SMHI 1809 (22 May 1984 FLO) (L!), 520 m, Podzorski SMHI
2091 (18 May 1984 FRU) (L!), 490–590 m, Ridsdale SMHI 1704 (17 May 1984 FRU)
(L!); Mt Victoria, 850 m, sine coll. SMHI 1704A (17 May 1984 FRU) (L!); Bato-Bato,
10 km south of Narra, on side road from the main trunk Narra-Aboabo, 100 m, Soejarto
& Fernando 7384 (26 January 1991 FRU) (L!); Taytay: Lake Manguao, valley stream,
Ridsdale SMHI 378 (9 April 1984 FLO/FRU) (L!); Ibangley, Sagcal, Romero et al.
PPI 364 (31 January 1991 FLO) (L!); Bataraza, Sumbiling, Sitio Gamayon, Bulanjao
Range, 50 m, Soejarto & Madulid 9052 (22 March 1995 FLO) (L!); Quezon Sitio
Bongalon, Cordero & Espiritu PNH 91599 (13 May 1964 FRU) (L!); Busuanga Island,
Lopez Bur. Sci. 41371 (September 1922 FRU) (L!), Ramos Bur. Sci. 41254 (September
1922 FLO) (L!); Culion Island, Merrill 612 (December 1902 FRU) (SING!), Merrill
669 (February 1903 FLO) (lectotype US!; isolectotypes A!, NY!, SING!); Bayas
Island: Copeland 112 (January 1904 FRU) (P!). Sibuyan Island, Romblon,
312

Magdiwang, Silum, Sitio Kulasian, Reynoso, Fuentes & Rivers PPI 14116 (20 February
1994 FRU) (L!). PANAY: McGregor Bur.Sci. 32195 (May–August 1918 FRU) (A!),
McGregor Bur. Sci. 32392 (May–August 1918 ST) (SING!), McGregor Bur. Sci.
32584 (May–August 1918 FLO) (BO!, SING!); Semirara Island, Alcasid PNH 6421
(March 1940 FLO) (A!); Buenavista, Taleon PNH 33906 (7 April 1955 FLO) (SAN!,
PNH!); Southwestern range of Mt Baloy, Laggit, c. 3 km SE of Lublub, 250–300 m,
Varadarajan, Reyoso, Fernando & Mosing 1552 (8 August 1987 FRU) (K!, L!).
NEGROS: Canlaon National Park, Mt Canlaon, Wasay trail, Stone et al. PPI 6307 (23
March 1992 FLO) (L!); Mt Katugasan, Edano PNH 21784 (March 1954 FLO) (K!,
PNH!, SING!). Guimaras Island, Buenavista, Sulit PNH 11787 (3 March 1950
FLO/FRU) (L!, PNH!, SING!). Paragua Island: 1500 ft, San Antonio Bay, dry slopes,
Merrill 839 (February 1903 FLO) (K!). BORNEO. SABAH. No locality: Governor
Creagh s.n. (August 1895 FLO) (K!). Kudat: Boden-Kloss SFN 19296 (16 September
1927 FRU) (SING!); near Kudat, Meijer SAN 19935 (10 September 1959 FLO) (L!,
SAN!), Meijer SAN 19936 (10 September 1959 FRU) (SAN!); behind STM Telecom
Station, Kamaruddin & Sukup KMS 2193 (11 March 1988 FRU) (SAN!); Bak bak, 200
ft [61 m], Brand SAN 30924 (17 July 1962 FRU) (SAN!); Loro F.R., 400 ft [122 m],
Tai SAN 51081 (18 May 1965 FRU) (K!); Balambangan Island, W from base camp,
1.5 km from base camp, Dolois et al. SP 18462 (24 May 2003 FLO) (SNP!); Malawali
Island: 50 ft, Diwol & Awang SAN 89971 (25 March 1979 FRU) (SAN!), 30 ft [9 m],
Fox SAN 69505 (24 February 1970 FRU) (SAN!), Tanjong, Sugau et al. SAN 145703
(8 August 2002 FRU) (SAN!); Labuan F.R.: Austin A 1245 (29 February 1948 FLO)
(KEP!, SING!), 50 m, Diwol SAN 135356 (8 September 1998 FLO/FRU) (K, L, KEP,
SAN!), 50 ft [15 m], Maidin SH 7328 (7 June 1937 FRU) (KEP!, L!, SING!); Lajong
F.R.: Majawat SAN 132063 (28 October 1991 FRU) (L!, SAN!, SAR!), Leopold, Pius,
Tangah & Soinin SAN 145151 (16 October 2002 FLO/FRU) (SAN!). Pitas: Kg Payas,
313

George & Amin SAN 121258 (12 May 1987 FRU) (KEP!, SAN!). Kota Kinabalu:
200 ft [61 m], Saikeh SAN 67153 (10 February 1971 FRU) (SAN!); Bukit Padang,
Meijer SAN 23827 (15 February 1961 FLO) (SAN!); Gaya Island:. Haviland 1438
(August 1892 FLO) (K!), 200 ft [61 m], Aban SAN 42017 (26 July 1966 FRU)
(SAN!), 100 ft [31 m], Ampuria SAN 41307 (4 March 1964 FRU) (SAN!), Berhaman,
Meijer & Madani SAN 134601 (30 June 1992 FRU) (SAN!), 300 ft [91 m], Kanis &
Kuripin SAN 56117 (16 February 1966 FRU) (KEP!, SAN!), south west, 100 ft [31 m],
Stone, Anderson & Leopold SAN 86850 (15 April 1977 FRU) (KEP!, SAN!, SAR!),
150ft [46 m], Wyatt-Smith KEP 80296 (4 August 1954 FLO) (KEP!, L!); Taman
Tunku Abdul Rahman, beach vegetation, Pereira & Postar JTP 698 (4 March 2000
FRU) (SAN!); 250 ft [72 m], Mujin SAN 33575 (8 April 1963 FRU) (SAN!), trail to
highest point, Rimi et al. SP 7288 (23 March 1995 FRU) (SAN!, SNP!), trail to Police
Beach, Dolois et al. SP 9417 (20 May 1999 FRU) (SNP!), Police Beach trail to base
camp, Dolois et al. SP 9508 (21 July 1999 FRU) (SAN!, SNP!); Trail to Padang Point,
Rimi et al. SP 7802 (25 June 1998 FRU) (SNP!); Manukan Island: Rimi et al. SP 7840
(24 June 1998 FLO) (SNP!), Wong & Jamili s.n. (1994 FLO/FRU) (SAN!, SAR!,
SING!). Beaufort: Siaunggau Forest Reserve, Diwol & Karim SAN 78155 (31 May
1974 FRU) (SAN!, SAR!, SING!). Kota Belud: 500 m, Gibbs 4340 (16 May 1913
FLO) (K!), Henderson SFN 38959 (8 April 1950 FLO/FRU) (L!, SING!). Sandakan:
Creagh s.n. (August 1895 FLO) (K!), 100 m, Foxworthy 600 (January–March 1916
FLO) (PNH!), Keith 1215, 500 ft [152 m] (20 February 1929 FLO) (K!); COF's
compound, 300 ft [91 m], cultivated, Keith S.H. 8862 (4 March 1938 FRU) (KEP!, L!,
SING!), near catchment area, Orolfo 831 (3 May 1926 FRU) (SING!); 26 ft [8 m],
Ramos 1195 (24 September 1920 FRU) (PNH!), Wood 741 (February–March 1920
FRU) (P!); behind Cheng Ming School, 300 ft., Pereira SAN 41096 (12 April 1964
FRU) (K!, SAN!); Buli Sim Sim, 800 ft [244 m], Keith A 1506 (21 February 1948
314

FRU) (BKF!, KEP!, P!, SING!); SW of Trig Buli Sim Sim, 500 ft [152 m], Pereira
SAN 41052 (28 November 1963 FRU) (SAN!), 0.5 mile SW of Trig Buli Sim-Sim, 400
ft [122 m], Pereira SAN 41073 (8 December 1963 FRU) (SAN!); Sibuga Road, 200 ft
[61 m], Abu Bakar SAN 27750 (15 November 1961 ST) (SAN!); Kebun China:
Ampon, Talip & Chow SAN 68479 (25 November 1969 FRU) (SAN!), Singh SAN
34719 (25 April 1962 FRU) (SAN!), low undulating country, Singh SAN 39256 (13
August 1963 FRU) (SAN!); Batu Sapi, 90 ft [27 m], Sayu SAN 38822 (30 September
1963 FRU) (SAN!); Leila F.R.: Charington SAN 23664 (3 January 1961 FRU) (K,
KEP!, L, SAN!, SAR!), 300 ft [91 m], James SAN 32552 (10 November 1962 FRU)
(SAN!, SAR!), Kapis SAN 34688 (10 March 1962 FRU) (KEP!, L!, SAN!, SAR!,
SING!), Kumin SAN 58837 (12 November 1968 FRU) (SAN!), Ogata 10646 (4
August 1968 FRU) (L!), 91 ft [28 m], Patrick & Jali SAN 62547 (15 March 1968 FLO)
(SAN!, SING!), 100 ft [31 m], Saikeh SAN 65559 (14 May 1969 FRU) (SAN!), 350 ft
[107 m], Sam SAN 28831 (28 February 1962 FLO) (KEP!, SAN!, SAR!, SING!), 75 ft
[23 m], Sayu SAN 38887 (10 August 1963 FRU) (SAN!, SING!), water catchment
area, Stone 6688 (17 March 1967 FLO) (KLU!, L!); Jalan Kulam, 90 ft [27 m], Kumin
SAN 73587 (12 August 1971 FRU) (SAN!, SAR!), Patrick SAN 70208 (24 August
1970 FRU) (SAN!, SAR!); Taman Datuk Chong Thien Vun, Amin SAN 95147 (22
April 1982 FRU) (KEP!, SAN!, SAR!), Meijer SAN 141282 (10 November 1994 FRU)
(SAN!); Kabili-Sepilok F.R.: 0–70 m, Kessler SAN 118033 (23 February 1989 FLO)
(SAN!, L!), Ridsdale 2029 (21 May 1989 FRU) (L!); RP292–2, Plot 2×1, Pereira &
Diwol JTP 398 (8 January 1997 ST) (SAN!), 30 m, Pereira & Postar JTP 581 (10
February 1999 FRU) (SAN!), Pereira et al. JTP 848 (May 2002 FRU) (SAN!); 150 m,
Wong & Puff WKM 2109 (11 April 1993 FRU) (SAN!); Mile 13, Labuk Rd., 50 ft [15
m], Fox SAN 70682 (29 July 1970 FRU) (SAN!); Seguntor, 300 ft [91 m], Postar et al.
SAN 141146 (9 September 2004 FRU) (SAN!). Labuk Sugut: Bonggaya, Bukit Obar,
315

100–190 m, Sugau et al. JBS 266 (23 September 1997 FRU) (L!, SAN!); Beluran, near
Kg Baba, ridge, 620 m, Postar et al. SAN 145528 (19 August 2004 FRU) (SAN!);
Tawai F.R., Tawai Hill, 150–1310 m, Sukup SA 1946 (24 August–2 September 1991
FRU) (UKMB!), Leopold & Goh et al. SAN 133845 (26 October 1999 FRU) (SAN!).
BORNEO
Palawan
Fig. 66. Distribution of “Rothmannia merrillii”.
316
PHILIPPINES

Fig. 67. Proposed lectotype specimen to be designated for “Rothmannia merrillii”, Merrill 669,
Philippines, Island of Calomian, Culion kept in the United States National Herbarium (US).
317

11. “Rothmannia” negrosensis [combination needed].
Basionym: Gardenia negrosensis Merr., Philipp. J. Sci., C 10 (1915) 111–112, pro
parte quoad Celestino Bur. Sci. 7333, Merr., Enum. Philip. Fl. Pl. 3 (1923) 530,
pro parte quoad Ramos & Edaño Bur. Sci. 31358 & Cardona FB 23868, excl.
Ramos & Edaño Bur. Sci. 36787 (= “Rothmannia graciliflora”), Sandique FB
26596 Wenzel 641 (= “R. leytensis”).
NEOTYPE (to be designated): Celestino Bur. Sci. 7333, Philippines Negros, Cadiz
(Fig. 69)
(February–March 1909 FLO) (L!).
Treelet to small tree, 5−12 m high. Twigs sparsely velvety hairy, becoming
glabrescent when old. Branch leaves typically in groups of 3’s, each trifoliate group
based on a distal node with only one leaf normally developed (the other vestigial or
very much reduced in size), and the proximal node with a pair of normal leaves.
Stipules broadly ovate-triangular, 2−4 mm long, 5−8 mm wide, sparsely to densely
pubescent outside; apex acute to cuspidate, c. 1 mm long. Young leaves on shoot tips
pubescent to velvety hairy on both sides. Mature leaves with petiole 0.5−1.3 cm long,
2−3 mm thick, velvety hairy to glabrescent; lamina broadly elliptic to ovate, to
sometimes lanceolate, (11.5−)14−22 cm long, (4−)6−10 cm wide, upper surface
glabrous, lower surface puberulous to velvety hairy; leaf base cuneate, subrounded to
unequal; leaf apex acute to short-acuminate, acumen to c. 0.3 cm long; thin-coriaceous
to chartaceous when dry; midrib on upper side flat, glabrous, on lower side raised,
velvety hairy; secondary veins 7−13 pairs, on upper side flat, glabrous, on lower side
slightly raised, velvety hairy; tertiary venation visible as simple connections between
pairs of secondary veins. Inflorescences: peduncle 0.2−0.3 cm long, 2.5−4 mm thick,
318

velvety hairy to tomentose; habit compact, with only 1 distinct branching order, rachis
0.2−0.5 cm long, primary branches along the rachis 1 pair, 0.2−0.5 cm long; bracts
broadly ovate, 3−4 mm long, 3−5 mm wide, pubescent. Flowers 2−3 per cyme;
pedicels subobsolete to 1 mm long, c. 1 mm thick; calyx tube 5−7 mm long, 3−6 mm
wide, sparsely pubescent to velvety hairy, hairs appressed, surface visible outside;
calyx limb cup-shaped to conical; calyx lobes 5−6, short-triangular, 0.5−1 mm long,
pubescent to velvety hairy; corolla hypocrateriform, the tube 6−9 cm long, 6−10 mm
wide at the throat, 2.5−3 mm wide at the base; non-speckled inside, outer surface
glabrescent to scantily pubescent, inner surface with hairy zone covering part or all of
the tube from the corolla mouth downwards; corolla lobes (5–)6−8, broadly ovate,
2.7−4.5 cm long, (1.5−)1.8−2.5 cm wide, outside scantily puberulous on the uncovered
half of the lobes, inside glabrous; anthers (5–)6–8, fully included within the tube
(inserted at the upper ⅓), 9−12 mm long; style and stigma c. 60−90 mm long, included
but reaching just below the corolla mouth and the level of or above the anthers, style
smooth, glabrous, stigma 2-lobed. Fruits globose, 5.5−6 cm long, 2.5−6 cm wide,
glabrous, drying smooth; fruit wall c. 1 mm thick; peduncle to 0.2 cm long, c. 4 mm
thick. Seeds slightly to much flattened and broadly ovate or suborbicular to narrowly
triangular, 10−11 mm × 7−9 mm.
NOTES. Merrill (1915) in Philipp. J. Sci., C 10: 111, designated Everett FB
7255, collected from Negros, Philippines as the type of the basionym, Gardenia
negrosensis. However, the holotype of this specimen was not located and is presumed
destroyed in the Philippine National Herbarium during the Second World War. Most of
Merrill’s primary collection (including holotypes) from the Philippines was deposited
in PNH (Van Steenis-Kruseman, 1950). The isotypes, if any, were also not located
319

after search at the various relevant herbaria where Merrill’s collections have been
distributed, i.e., A, K, L, P, NY, SING, U, and US.
Besides the type indicated by Merrill, he had also listed other specimens in the
original description. One of the specimen, Celestino Bur. Sci. 7333 collected from
Cadiz, Negros, around Feb–Mar 1909 and kept in Leiden, was available for study.
Based on Article 9, paragraph 9.6 of the International Code of Botanical
Nomenclature, Vienna Code (2006), Celestina Bur. Sci 7333 from L is to be designated
as the neotype for the species name.
This species is very similar to “R. merrillii”. However, “R. negrosensis” is
consistently hairier than “R. merrillii” in a number of parts: the lower leaf surface
(lamina) and its veins, inflorescence peduncle, bracts and calyx are velvety hairy to
pubescent compared to “R. merrillii”, which is typically glabrous to scantily puberulous
or only sparsely pubescent. The degree of hairiness in the inflorescences, calyces and
leaves compared to “R. merrillii” have been pointed out by Merrill (1915). The outer
surface of the corolla tube is typically glabrescent to scantily puberulous, but in “R.
merrillii” is always glabrous. Another main distinguishing character of “R.
negrosensis” is found in the corolla lobes, which are broader (typically 1.8–2.5 cm
wide, but rarely less than 1.8 cm wide) than those of “R. merrillii” (typically 0.5–1 cm,
rarely to 1.5 cm wide). In the original description of Gardenia negrosensis, the fruits
were described as “subglobose to ovoid, up to 2.5 cm in diameter”. However, it is
uncertain if this fruit measurement referred to a mature fruit. The only fruiting
specimen (with mature fruits) available for study has a larger diameter than was
described in the original publication, i.e., 6 cm diameter.
DISTRIBUTION. Thus far only found in the Philippines (Negros, Mindoro and
Palawan islands) (Fig. 68).
HABITAT. Recorded in lowland to hill forests, to 850 m alt.
320

SPECIMENS EXAMINED.
PHILIPPINES. NEGROS: Cordona 23868 (February–April 1915 FLO) (A!, SING!);
Capiz Prov., Jamindan, Ramos & Edano Ramos 31358 (April–May 1918 FLO) (L!).
Cadiz, Celestino 7333 ( February–March 1909 FLO) (neotype L!). MINDORO:
Calintaaan, Poypoy, Sitio Sapaton, Mt Iglit-Baco National Park, 850 m, Barbon,
Garcia PPI 22603 (26 August 1996 FRU) (L!); Mt Halcon, Edano PNH 3512
(January–February 1948 FLO) (L!, SING!), Mt Halcon, Ramos & Edano Ramos 40730
(March 1922 FLO) (A!); San Teodoro, Sibuang Camp, Ridsdale 857 (12 February 1985
FRU) (L!). Sibuyan Island, Capiz Prov., Mt Guinting-Guinting (Magallanes), Elmer
12246 (April 1910 FLO) (L!), Mt Guinting-Guinting, 300 m, Madulid 6897 (16 June
1987 FRU) (L!). PALAWAN: St. Paul’s Bay, Caboyugan village, W side of the road,
50 m, Ridsdale SMHI 1649 (7 May 1984 FRU) (K!, L!).
Palawan
321
Luzon
Fig. 68. Distribution of “Rothmannia graciliflora”, “R. lagunensis”, “R. leytensis” and “R. negrosensis”.
[Note that the symbol indicated on the map for “R. leytensis” does not reflect the exact localities because
they are either not stated down on the specimen labels or could not be found.]

Fig. 69. Proposed neotype specimen to be designated for “Rothmannia negrosensis”, Celestino
Bur. Sci. 7333, Philippines, Negros, Cadiz (L).
322

12. “Rothmannia” nigrescens [combination needed].
Basionym: Timonius nigrescens Valeton, Bot. Jahrb. Syst. 61 (1927) 46.
LECTOTYPE (to be designated): Ledermann 9560, New Guinea, Sepik (1912–13
FLO) (L!).
Randia bartlettiana Merrill & Perry, J. Arnold Arbor. 29 (1948) 165.
(Fig. 70)
TYPE: Clemens 41856, Papua New Guinea, Morobe district, Boana, 2500–
4500 ft [762–1371 m] (26 September 1940) (holotype A!; isotypes L!, UC!).
Tree, to 20 m high, 10−25(−45) cm diameter. Bark slightly cracked, grey to
brown; inner bark straw. Sapwood cream. Twigs scantily puberulous to glabrescent.
Branch leaves typically in groups of 3’s, each trifoliate group based on a distal node
with only one leaf normally developed (the other vestigial or very much reduced in
size), and the proximal node with a pair of normal leaves. Stipules short-triangular,
3−5 mm long, 4−6 mm wide, glabrescent to sparsely pubescent outside; apex
acuminate, 1−3 mm long. Young leaves on shoot tips appressed pubescent on both
sides. Mature leaves with petiole 0.1−0.5 cm long, 1.5−2 mm thick, glabrescent to
finely appressed pubescent; lamina oblanceolate, narrowly to broadly elliptic,
(10−)14−18(–30) cm long, (2.5−)5−8.5 cm wide, upper surface glabrescent becoming
glabrous, lower surface sparsely pubescent; leaf base cuneate, attenuate to unequal;
leaf apex acute to acuminate, acumen to 0.8 cm long; thin- to thick-coriaceous when
dry; midrib on upper side flat, glabrescent becoming glabrous, on lower side raised,
glabrescent to sparsely puberulous; secondary veins 6−9 pairs, on upper side flat,
glabrescent becoming glabrous, on lower side raised, glabrescent to sparsely
puberulous; tertiary venation typically visible as a much-branched network between
pairs of secondary veins. Inflorescences: peduncle 0.3−0.6(−1.0) cm long, 1.5–3 mm
323

thick, glabrescent to appressed pubescent; habit laxly arranged, with (1–)2–3 distinct
branching orders, rachis 0.3−1(−2.5) cm long, primary branches along the rachis 2–3
pairs, 0.4–0.9 cm long; bracts ovate, 1.5−3 mm long, 1−2 mm wide, glabrescent to
appressed pubescent. Flowers 10−30 per cyme; pedicels 1−2 mm long, 0.5−1.5 mm
thick; calyx tube 3−6 mm long, 2−4 mm wide, scantily puberulous to appressed
pubescent throughout, hairs appressed, surface visible; calyx limb cup-shaped; calyx
lobes 5, short-triangular to sometimes linear, 0.5−1(−8) mm long, scantily puberulous
to appressed pubescent; corolla hypocrateriform, white, the tube (1−)1.8−3(−3.5) cm
long, 2−4.5 mm wide at the throat, 1−2 mm at the base; non-speckled inside, outer
surface sparsely to densely appressed pubescent, inner surface with hairy zone covering
part or all of the tube from the rim of the corolla mouth downwards; corolla lobes 5,
narrowly oblong to elliptic, (1.2−)1.6−2 cm long, 0.2−0.4 cm wide, outside pubescent
on the uncovered half of the corolla lobes, inside hairy at the base; anthers 5, fully
included within the tube (inserted at the upper ⅓), 6−8 mm long; style and stigma
10−30 mm long, included but reaching just below the corolla mouth and the level of the
anthers, style smooth, glabrous. Fruits globose to ellipsoid, 3−5 cm long, 3−4 cm
wide, glabrous, drying smooth; fruit wall (1.5−)2−3 mm thick; peduncle 0.6–1.3 cm
long, 2−4(−5) mm thick. Seeds slightly to much flattened and broadly ovate or
suborbicular to narrowly triangular, 5−6 mm × 6−8 mm.
NOTES. In 1927, Valeton published the basionym, Timonius nigrescens in
Engler’s Bot. Jahrb. Syst. Vol. 61, pg. 46. In this account, the type specimen was
clearly indicated, Ledermann 9560 from Sepik, Papua New Guinea, taken during the
Sepik Expedition in 1912–13. According to Van Steenis-Kruseman (1950),
Ledermann’s holotype collections from this expedition were deposited in the Berlin
324

Herbarium (B) and the duplicates distributed to L, K, SING and US. The holotype of
this collection was destroyed during the Second World War.
Based on Article 9, paragraph 9.2 of the International Code of Botanical
Nomenclature, Vienna Code (2006), the sheet from L, which is the isotype of
Ledermann 9560 is to be designated as the lectotype for this species.
Most of the specimens placed under this species have typically longer corolla
tubes, 1.8−3 cm, sometimes reaching 3.5 cm long. However, the type specimen has
shorter corolla tubes, to c. 1 cm long only.
(Amale).
VERNACULAR NAMES. Garamgudzun (Dumpu); Siton (Faita); Manimah
DISTRIBUTION. Widespread in Papua New Guinea (Madang, Morobe,
Western, Milne Bay, Southern, Northern, Central and East Sepik Provinces) (Fig. 84).
HABITAT. Found in primary and secondary lowland to hill forests, frequently
along streams, also on ultrabasic and volcanic soils, to 730 m alt.
SPECIMENS EXAMINED.
PAPUA NEW GUINEA. MOROBE PROV.: Markham Valley, Patep creek, 2300 ft
[701 m], Millar NGF 11759 (6 October 1959 FLO) (BRI!); Lae subdistrict, Buso, ridge
to Mt Kawea, 200 m, Streimann & Foreman NGF 24454 (26 April 1972 FRU) (BRI!,
K!, SING!); Garaina subdistrict, Saru River, 2400 ft, Gillison, Kairo & Seruvato NGF
25768 (12 December 1968 FLO/ FRU) (BR!, L!, SING!); Kipu-Wakaia trail, ½ mile E
of Luluai's burial place, at edge of Kunai patch, 2,300 ft [701 m], Frodin NGF 26400 (8
January 1966 FRU) (L!, SING!); Golden Pines subdistrict, Palenkwa, Wau, Streimann
& Kairo NGF 39459 (24 April 1969 FRU) (L!); Boana, 2500–4500 ft [762–1371 m],
Clemens 41856 (1940 FLO/FRU) (A!, L!, UC!); Lae, Native Collector 4650 (6
November 1952 FLO/FRU) (BM!, L!); Oomsis Creek, c. 18 miles [Km 29], W of Lae
along Lae-Bulolo Road, 400 ft [122 m], Hartley TGH 9669 (8 January 1962 FRU) (L!),
325

about 18 miles [Km 29], W of Lae, 400 ft [122 m], Hartley TGH 10060 (23 March
1962 FRU) (BRI!, L!); near the Butibum River about 7 miles N of Lae, 100 ft [31 m],
Hartley TGH 9846 (1 February 1962 FRU) (BRI!, L!), hills adjacent to the Busu river,
near Bumayong settlement, 200 m, Takeuchi 7013 (4 November 1990 FLO) (L!); NE
of Lae, Busu River, near suspension bridge over Busu River, Thorne & Henty 27454
(23 May 1960 FLO) (BRI!). MADANG PROV.: Usino subdistrict, 400 ft [122 m],
Henty NGF 28003 (17 August 1966 FLO) (L!, SING!), Amiaba River, 500 ft [152 m],
Forman, Noble & Farley NGF 45927 (11 January 1970 FRU) (L!); Bundi subdistrict,
Braham, 550 ft [166 m], Henty NGF 49253 (17 March 1972 FRU) (L!); Josephstaal,
T.N.G., 250 ft [76 m], White NGF 10311 (1958 FLO) (L!, SING!); Madang subdistrict,
Gogol River, 140 ft [43 m], Katik NGF 46674 (7 May 1979 FRU) (L!, SING!).
WESTERN PROV.: Kiunga subdistrict, 100 ft [31 m], Streimann LAE 51817 (11
August 1971 FRU) (BRI!, L!, SING!); near Ingambit village, 480 ft [146 m], Henty,
Ridsdale & Galore NGF 33014 (13 June 1967 FRU) (L!, SING!); vicinity of the Mipan
track, near mouth of the June river, northern Lake Murray area, 100 ft [31 m], Pullen
7478 (12 October 1967 FRU) (K!); Ramu valley, c. 5 miles SE Faita airstrip, lower
slopes Bismark Range, 600 ft [183 m], Saunders 227 (9 June 1955 FLO) (BM!, L!).
MILNE BAY PROV.: Peria Creek, Kwagira River, 50 m, Brass 24224 (28 August
1953 FLO) (L!), Ma-u River, Rabaraba, 488 m, Katik LAE 56293 (28 June 1972 FLO)
(BRI!, L!). EAST SEPIK PROV.: No locality, Ledermann 9560 (1912–1913 FLO)
(lectotype L!). SOUTHERN PROV.: Mt Bosavi, Northern side, 600−700 m, Jacobs
9326 (19 October 1973 FLO/FRU) (L!). NORTHERN PROV.: Dobodura,
Cavanaugh NGF 2035 (21 March 1945 FRU) (L!). CENTRAL PROV.: Sogeri
Region, Forbes 175 (sine datum FRU) (L!), Forbes 462 (1885–6 FRU) (L!, P!).
326

Fig. 70. Proposed lectotype specimen to be designated for “Rothmannia nigrescens”,
Lederman 9560, New Guinea, Sepik (L).
327

13. “Rothmannia” papuana [new species description needed].
TYPE (to be designated): Carr 11492, Papua New Guinea, Central Prov., Lolorua (24
(Fig. 71)
February 1935 FLO) (holotype A!; isotypes BM!, L!).
Tree, to 25 m high, to 45 cm diameter. Bark grey-brown; inner bark deep red.
Sapwood olive green to light red. Twigs glabrous. Branch leaves typically in groups
of 3’s, each trifoliate group based on a distal node with only one leaf normally
developed (the other vestigial or very much reduced in size), and the proximal node
with a pair of normal leaves. Stipules broadly triangular, 5−9 mm long, 5−7 mm wide,
glabrous outside; apex acute to cuspidate, 1−3 mm long. Young leaves on shoot tips,
glabrous on both sides. Mature leaves with petiole 0.1−0.5 cm long, 1.5−2 mm thick,
glabrous; lamina broadly elliptic, lanceolate to oblanceolate, 10−19 cm long, 5−7.5 cm
wide, upper surface glabrous, lower surface glabrous; leaf base cuneate, sometimes
attenuate; leaf apex acute, acumen 0.2−0.3 cm long; thin-coriaceous to chartaceous
when dry; midrib on upper side flat, glabrous, on lower side raised, glabrous; secondary
veins 6–9 pairs, on upper side flat, glabrous, on lower side raised, glabrous; tertiary
venation visible as simple connections between pairs of secondary veins.
Inflorescences: peduncle 0.1−1 cm long, 1.5−2 mm thick, glabrous; habit laxly
arranged, with 3−4 distinct branching orders, rachis 3−3.5 cm long, primary branches
along the rachis 1−2 pairs, 1–1.5 cm long; bracts short-triangular, 2−2.5 mm long, c. 2
mm wide, glabrous. Flowers 5−10 per cyme; pedicels 2−4 mm long, c. 1 mm thick;
calyx tube 3−5 mm long, 3−5 mm wide, glabrous outside; calyx limb cup-shaped; calyx
lobes 5, broadly short-triangular, 0.5−1 mm long, glabrous outside; corolla
hypocrateriform, white, the tube 1−1.5 cm long, 4−5 mm wide at the throat, 2−2.5 cm
wide at the base; non-speckled inside, outer surface glabrous, inner surface glabrous;
328

corolla lobes 5, broadly elliptic to ovate, 1.2−1.5 cm long, 0.5−0.7 cm wide, outside
and inside glabrous; anthers 5, fully included within the tube (inserted at the upper ⅓),
5–9 mm long; style and stigma 10−13 mm long, included but reaching just below the
corolla mouth and the level of the anthers, style smooth, glabrous. Fruits globose,
3.5−4 cm long, 3−3.5 cm wide, glabrous, drying smooth; fruit wall c. 1 mm thick;
peduncle c. 0.4 cm long, c. 5 mm thick. Seeds slightly to much flattened and broadly
ovate or suborbicular to narrowly triangular, 6−7 mm long × 6−7 mm wide.
NOTES. In the type specimen, Carr 11492, which has flowers, the lateral
branches of the ultimate vegetative node of a branch segment before terminal flowering
is conspicuously elongated, 1–2 cm long. This character has not been observed in the
fruiting specimens of this species or in the other Malesian species. Perhaps it is not a
consistent feature. The species is distinct in having both sides of the leaves completely
glabrous, even in the young shoot tips, and completely glabrous inflorescences.
DISTRIBUTION. Confined to Papua New Guinea (Central Province only).
Only known from 3 collections thus far (Fig. 84).
HABITAT. Recorded from seasonally dry evergreen rain forest, to c. 70 m alt.
SPECIMENS EXAMINED.
PAPUA NEW GUINEA. CENTRAL PROV.: Lolorua, 100ft [30 m], Carr 11492
(24 February 1935 FLO) (holotype A!; isotypes BM!, L!); Port Moresby subdistrict,
Kuriva forestry area, near Veimauri River, 200ft [61 m], Streimann & Kairo LAE
51558 (6 May 1971 FRU) (BRI!, L!, SING!); near Port Moresby, 3 miles past Brown
Bridge, 100ft [30 m], White NGF 8262 (4 May 1960 FRU) (L!, SING!).
329

Fig. 71. “Rothmannia papuana”. A, leafy branch with inflorescences. B, close-up of lower leaf surface
showing domatia at the axils of the midrib and secondary veins. C, calyx tube glabrous. D, longitudinal
section of the flower. E, outer surface of the corolla lobe. F, fruit.
A–E from Carr 11492 (holotype, A), F from Streimann & Kairo LAE 51558 (BRI).
330

14. “Rothmannia” pseudoternifolia [combination needed].
Basionym: Gardenia pseudoternifolia Valeton, Icon. Bogor. 4 (1912) 123, t.
CCCXXXVIII.
TYPE: Teysmann 19411, Insula Liengga, prope Sapientjan (February 1911) (holotype
BO!; isotype L!).
Rothmannia kuchingensis sensu Coode et al., Checkl. Flow. Pl. & Gynm. Brunei (1996)
(Fig. 74)
290, non (W.W.Smith) K.M.Wong (= “Rothmannia grandis”; Rothmannia
schoemannii sensu Kessler et al., Checklist for a tree flora of the Balikpapan-
Samarinda area, East Kalimantan, Indonesia (1992) 67, non (Teijsm. & Binn.)
Tirveng.; Rothmannia sp. nov. in Kessler & Sidiyasa, Trees of the Balikpapan-
Samarinda Area, East Kalimantan, Indonesia (1994) 202, fig. 16; Rothmannia
sp. 2 in Beaman & Anderson, The Plants of Mount Kinabalu (2004) 340.
Medium to large tree, to 45 m high, to 50 cm diameter. Bark smooth to mottled,
fissured, lenticellate and scaly, pale to blackish-grey to reddish brown; inner bark
yellowish to reddish brown. Sapwood pale orange or maroon, grey-brown, pinkish, to
yellowish. Twigs sparsely to densely pubescent, tomentose to velvety hairy to hispid.
Branch leaves typically in groups of 3’s, each trifoliate group based on a distal node
with only one leaf normally developed (the other vestigial or very much reduced in
size), and the proximal node with a pair of normal leaves. Stipules broadly ovate-
triangular, 3−10 mm long, 3−10 mm wide, glabrescent, sparsely to densely pubescent
or hispid outside; apex acuminate to caudate, 2−8 mm long. Young leaves on shoot
tips pubescent, velvety hairy to hispid on both sides. Mature leaves with petiole
subobsolete or to 5 cm long, 1−3 mm thick, sparsely to densely pubescent, tomentose,
331

velvety hairy to hispid; lamina ovate, obovate, elliptic to oblanceolate, 7−27 cm long,
2.5−12 cm wide, upper surface glabrous to sparsely pubescent or hispid, lower surface
sparsely to densely pubescent, velvety hairy to hispid; leaf base cuneate, subcordate,
subrounded to unequal; leaf apex acute, short-to-long acuminate, acumen to 1.5 cm
long; thin-to-thick-coriaceous to chartaceous when dry; midrib on upper side flat,
glabrescent, pubescent to hispid, on lower side distinctly raised, sparsely to densely
pubescent, tomentose, velvety hairy to hispid; secondary veins 7−18 pairs, on upper
side flat, glabrescent to sparsely pubescent to hispid, on lower side raised, sparsely to
densely pubescent, tomentose, velvety hairy to hispid; tertiary venation visible as a
much-branched network between pairs of secondary veins. Inflorescences: peduncle
0.5−6.5 cm long, 1−4 mm thick, pubescent, tomentose, velvety hairy to hispid; habit
compact to laxly arranged, unbranched to with 1−3 distinct branching orders, rachis
(0.2−)0.5−3.5 cm long, primary branches along the rachis 1−2(−3) pairs, 0.5−2 cm
long; bracts broadly to narrowly ovate, 2−9 mm long, 2.5−4 mm wide, pubescent,
velvety hairy to hispid. Flowers (1−)3−20 per cyme; pedicels subobsolete to 2 mm
long, 1−2.5 mm thick; calyx tube 3−10 mm long, 2.5−6 mm wide, pubescent, velvety to
hispid, hairs suberect to erect, surface visible, calyx limb cup-shaped; calyx lobes
5(−6), short-triangular, narrowly triangular to long-linear, 0.5−15 mm long, sparsely to
densely pubescent, velvety to hispid; corolla hypocrateriform, white to pale yellow, the
tube 3.5−9 cm long, 2−6 mm wide at the throat, 1−3 mm wide at the base; non-
speckled inside, outer surface pubescent, tomentose to hispid, inner surface glabrous;
corolla lobes 5(−6), ovate to elliptic, 1.2−3 cm long, 0.2−1 cm wide, outside pubescent
on the uncovered half of the lobes, inside glabrous; anthers 5, fully included within the
tube (inserted at the upper ⅓), 7−13 mm long; style and stigma 11−48 mm long,
included but reaching less than ⅔ way along the corolla tube, and well below the
332

anthers, style smooth, glabrous. Fruits globose to ellipsoid, 3−10 cm long, 3−7.3 cm
wide, glabrous, sparsely to densely pubescent or velvety hairy, drying smooth to
wrinkled; fruit wall 1−3 mm thick; peduncle 0.2−4 cm long, 10 mm thick. Seeds
slightly to much flattened and broadly ovate or suborbicular to narrowly triangular,
9−11 mm × 6−7 mm.
NOTES. Many of the specimens were previously misidentified as Randia
grandis, Gardenia forsterniana, and Porterandia anisophylla.
This species is very variable in vegetative and reproductive characters.
However, two varieties can be discerned based on the type of hairs occurring on the
petiole, lower leaf surface, and inflorescences (see key to varieties).
VERNACULAR NAMES. Sarawak: Mengkudu hutan; engkudu hutan (Iban).
Brunei: badal ayam; bedal manok (Iban); lengang & badal mano’ (Dusun).
Kalimantan: bengkil; teratat (Loa Djanon); kaju pringit.
Kalimantan).
DISTRIBUTION. Endemic to Sumatra and Borneo (Sabah, Sarawak, Brunei &
HABITAT. Widespread and common in primary and secondary forests in the
lowlands and hill forests, including kerangas forests, limestone and seasonal flood-
plain areas (e.g., fresh-water swamp forests), to c. 1640 m alt.
Key to the varieties of “Rothmannia pseudoternifolia”
Petiole, lower leaf surface, and inflorescences velvety hairy or tomentose to pubescent
or very sparsely pubescent to puberulous, the hairs always short (generally not more
than 0.5 mm long) and soft to the touch………………….…. “var. pseudoternifolia”
Petiole, leaf lower surface, and inflorescences sparsely hispid-hairy, the hairs longer (c.
1–1.5 mm long) and bristly, generally stiff or harsh to touch………..... “var. hispida”
333

Within “var. pseudoternifolia”, a total of five subgroupings may be discerned
(Figs. 72a–b) and, which do not segregate sufficiently to merit any formal taxonomic
distinction. Two scatter diagrams in Figs. 72a–b show five subgroups recognizable
within the typical variety, variation of inflorescence peduncle length (cm) with degree
of hairiness (hair density) of the lower leaf surface (Fig. 72a); and variation of leaf
width (cm) with degree of hairiness (Fig. 72b). From these, the following can be
deduced:
(i) Bornean specimens confined to Kinabalu, Sabah that have compact cymes
(branching to only 1 order), short peduncle, narrower leaves and less hairy
leaves (Subgroup 1) are different from the three other subgroups (3, 4 & 5) that
have variable peduncle length and leaf width but considerably more hairy
leaves (Figs. 72a–b). However, they are closely related to Subgroup 2
(Bornean specimens from Sabah generally, East and Central Kalimantan and
the Lawas region of Sarawak) that have rather variable peduncle length and
leaf width and less hairy leaves.
(ii) The type alliance from Sumatra (Subgroup 5) is closer to the other
subgroups than to the Kinabalu taxon (Subgroup 1) but nevertheless, there is
overlap among these different subgroups.
“R. pseudoternifolia var. pseudoternifolia” appears to be a comparatively widespread
taxon showing some geographical variation in different characters, where changes have
not yet progressed to the stage where discrete groups are clearly separable.
DISTRIBUTION. Endemic to Sumatra and Borneo (Sabah, Sarawak, Brunei &
Kalimantan) (Fig. 73).
334

a)
Inflorescence peduncle length (cm)
b)
Maximum leaf breadth width (cm)
(cm)
6
5
4
3
2
1
0
14
12
10
8
6
4
2
Sub-group 1
Sub-group 2
Sub-group 3
Sub-group 4
Sub-group 5
0 1000 2000 3000 4000 5000
0 1000 2000 3000 4000 5000
Hair density (no. of hairs/cm 2 )
Fig. 72. Scatter diagrams showing the distribution of individuals within “Rothmannia pseudoternifolia
var. pseudoternifolia” based on: a, hair density versus inflorescence peduncle length (only mature or
near mature length were taken); b, hair density versus maximum leaf width.
335

SPECIMENS EXAMINED.
SUMATRA. RIAU PROV.: Tigapuluh Mountains, 5 km W of Talanglakat on
Rengat-Jambi Rd., Bukit Karampal area, 100 m, Burley, Tukirin et al. 1323 (9
November 1988 FRU) (KEP!, SING!); NW of Tigapuluh Mountains, S of Bukit
Huluspin, 400 m, Laumonier YL 5740 (1 January 1984 FRU) (L!); Indragiri uplands,
Muara Padjanki, Buwalda 6453 (9 April 1939 FLO/FRU) (K!, SING!). RIAU
ISLANDS PROV.: Lingga Island, prope Sapientjan, Teysmann 19411 (February
1911 FLO) (holotype BO!; isotype L!). BORNEO. SABAH. Tawau: Elmer 21075
(October 1922–March 1923 ST) (K!, L!, SING!, U!); Tajong F.R., Bongsu SAN 63035
(23 June 1968 ST) (SAN!); near Tajong F.R., 120 m, Pereira, Leopold, Pung Ubaldus
& Salamat JTP 579 (27 November 1998 ST) (L!, SAN!), JTP 580 (27 November 1998
ST) (SAN!); Imam Estate, 10 miles [16 km] N of Tawau, 350 ft [107 m], Wood SAN
17248 (9 December 1955 FLO/FRU) (BRI!, KEP!, L!, SING!), Mile 26, Apas Rd., 50
ft [15 m], Meijer SAN 19307 (25 June 1959 FRU) (L!, SAR!, SING!); Brantian, Mile 6
[Km 9.6], Benawood logging area, Fedilis & Sumbing SAN 88420 (19 July 1978 ST)
(L!, SAN!), Mile 3 [Km 1.9], Hap Seng logged area, Fedilis & Sumbing SAN 89114
(24 November 1978 FRU) (KEP!, SAN!, SING!); Kalabakan, Mile 12 [Km 19.3], Hap
Seng logged area, Fedilis SAN 94878 (11 June 1982 FRU) (KEP!, SAN!, SAR!);
Kalabakan, Ulu Sg. Sepaku, Fedilis & Sumbing SAN 95747 (22 March 1983 FLO) (L!
SAN!); Bukit Gemok, Leopold, Zamrie & Goh SAN 106706 (5 August 2002 ST)
(SAN!); Tawau Hills Park: E of Park, Leopold & Sigin SAN 111182 (19 October 1985
FRU) (SAN!), next to the pond at side of Sabah Park’s office, 260 m, Pereira, Leopold,
Pung, Ubaldus & Salamat JTP 547 (24 November 1998 FLO/FRU) (L!, SAN!), near
hanging bridge, Pereira, Leopold, Pung, Ubaldus, Salamat JTP 560 (26 November
1998 FRU) (SAN!), 260 m, Pereira, Ubaldus & Postar JTP 658 (25 November 1999
FLO) (SAN!), 1000 ft [305 m], Pereira, de Wilde & Postar JTP 669 (13 January 2000
336

FRU) (SAN!). Lahad Datu: Danum Valley, Stirling Ecological Plots, Ridsdale CER
2458 (19 June 1997 ST) (L!, SAN!), Ridsdale CER 2462 (22 June 1997 FRU) (L!,
SING!); Mile 4, Segama Rd., Chai SAN 29791 (27 August 1962 FRU) (KEP!, SAN!,
SAR, SING!); Kennedy Bay, V.J.R. No. 14 Block, Pilus SAN 47015 (24 August 1964
ST) (SAN!) Sandakan: Sepilok, behind sawmill, Arshid SAN 85474 (21 May 1977
FLO) (K!, SAN!, SAR!), Kabili-Sepilok F.R., Compt. 6, 15 miles W of Sandakan, 15 ft
[5 m], Charington A 3895 (24 February 1955 FLO) (BRI!, KEP!, L!, SING!), 110 ft
[33 m], James SAN 39049 (4 April 1964 FLO) (K!, L!, SAN!, SING!), Arboretum,
forest adjacent to Kabili-Sepilok F.R., 10 m, Pereira & Postar JTP 582 (10 February
1999 ST) (SAN!). Labuk Sugut: Mile 110 [Km 177], Telulpid-Ranau Road, 400 ft
[122 m], Leopold SAN 89222 (16 August 1978 FRU) (KEP!, SAN!, SING!); Beluran,
near Kg Baba, 680 ft [207 m], Postar, Petrus, Tawadong & Jemson SAN 145527 (19
August 2004 FRU) (SAN!). Kinabatangan: Ulu Sg Pinangah, Sigin & Lideh SAN
107184 (15 October 1984 FLO/FRU) (KEP!, SAN!, SAR!); logging road to Sapulut,
Amin, Mijol & Joseph SAN 107312 (23 October 1984 FRU) (KEP!, SAN!, SING!);
Ulu Sg Pingas-Pingas, Asik SAN 110951 (19 March 1985 FLO) (SAN!); Deramakot
F.R., Diwol SAN 124572 (19 September 1990 FRU) (SAN!); Bukit Tangkunan, sample
plot, 300 ft [91 m], Leopold SAN 61077 (17 June 1967 ST) (SAN!); Tongod, Maliau
Basin, E of Belian camp, 300 m, Pereira, de Wilde, de Wilde-Duyfjes, Postar &
Ubaldus JTP 814 (12 July 2001 ST) (SAN!). Kudat: Matunggong, Bukit Gumantong,
1300 ft [396 m], Sugau, Diwol & Dauni SAN 144303 (21 February 2002 FLO) (SAN!,
SING!); Hutan Simpan Lajong, Majawat SAN 132062 (28 October 1991 FRU) (L!,
SAN!); Loro Kecil F.R., 70 ft [21 m], Corpus & Lajangah SAN 36129 (19 May 1963
FRU) (SAN!, SAR!); Kota Marudu, Torintidon F.R., Berhaman SAN 133163 (12
October 1991 FRU) (SAN!), above Kg Lampada, Aban et al. SAN 99497 (6 August
1983 FRU) (SAN!). Penampang: Kepayan F.R., 45 m, Ag Amin SAN 104065 (17
337

October 1991 FRU) (L!, SAN!), Ag Amin SAN 114895 (25 November 1989 FRU)
(SAN!); Sugud Recreational F.R., 370 m, Pereira, Goh, Chuah & Leopold JTP 544 (8
September 1998 ST) (L!, SAN!), 600 ft [183 m], Pereira, Diwol & Ubaldus JTP 663 (2
December 1999 FRU) (SAN!), 600 ft [183 m], JTP 664 (2 December 1999 FRU)
(SAN!), 300 m, Pereira, Noorsiha, Diwol & Postar JTP 730 (8 March 1999 FLO)
(SAN!). Tuaran: Tamparuli, 150 m, Sato, Andau & Sintiong 1128 (12 July 1981 ST)
(SAN!). Kota Belud: Mt Templer F.R., 1000–1500 ft [305–457 m], Kanis & Kuripin
SAN 54362 (9 February 1966 FLO) (SAN!); Kiau Nuluh, 735 m, Pereira, Noorsiha,
Diwol & Postar JTP 586 (24 February 1999 ST) (SAN!); Kogiitan, Jusimin 211 (22
January 1993 FRU) (K!, SAN!, SNP!). Ranau: Kg Melangkap Tomis, 1 km from
Tomis, Lorence LL 1301 (17 November 1995 FRU) (SAN!, SNP!); along Paramat
River, Lorence LL 141 (12 December 1995 FRU) (SAN!, SNP!); Kepangian, Amin &
Jarius SAN 106774 (15 February 1986 FRU) (SAN!); Tambiu, Amin & Jarius SAN
116151 (22 August 1986 FRU) (SAN!); Poring Hot Springs, trail to canopy walkway,
560 m, Pereira, Noorsiha, Diwol & Postar JTP 590 (27 February 1999 ST) (SAN!);
Nalumad trail, 4 miles NE of Poring, Meijer & Sadau SAN 49786 (12 September 1965
FRU) (SAN!, SNP!); Kinabalu National Park, 2700 ft [823 m], Puasa BNB 3462 (23
September 1931 ST) (K!); Penibukan, 4000 ft [1219 m], Clemens & Clemens 40720 (1
October 1933 FLO) (A!, K!, L!), 4000–5000 ft [1219−1524 m], Clemens & Clemens
30485 (28 December 1932 FRU) (K!, L!), Clemens & Clemens 31549 (7 February
1933 FRU) (BO!, K!, L!), E of camp, 4000–5000 ft [1219−1524 m], Clemens &
Clemens 30883 (10 January 1933 FLO) (K!), 4000 ft [1219 m], Clemens & Clemens
32151 (14 March 1933 FRU) (A!, K!, L!). Tambunan: Trusmadi, 600 ft [183 m],
Francis SAN 50409 (2 August 1966 FLO) (KEP!, SAN!); Km 64, Jalan Kota
Kinabalu-Tambunan, 1200 m, Lily 6832 (10 September 1984 FLO) (SAN!); Rafflesia
Information Centre, Rafflesia F.R., Pereira, Puff & Ahmad JTP 842 (2 September 2001
338

FLO) (SAN!), Rafflesia trail, Postar, Nilus, Hastie & Petrus SAN 144692 (27 June
2003 FLO) (SAN!), 4290 ft [1307 m], Pereira, Goh, Chuah, Hew et al. JTP 499 (14
September 1997 FLO) (SAN!), 4040 ft [1231 m], Pereira, Goh, Chuah, Hew et al. JTP
500 (14 Septemeber 1997 ST) (L!, SAN!); Pegalan, Ignatius & Clement SAN 140139
(2 May 1995 FRU) (SAN!), Jalan Pegalan, 620 m, Pereira, Wong, Sugau, Leopold,
Joseph, Nilus & Puff JTP 156 (10 March 1995 FRU) (L!, SAN!); Kirokot, Ignatius &
Clement SAN 139995 (5 April 1995 FRU) (SAN!), SAN 140048 (19 April 1995 FRU)
(SAN!), above Rampon River, Joseph et al. SAN 123870 (27 April 1988 FRU) (SAN!).
Keningau: Lanas, Shang Liang logging area, Sumbing SAN 118547 (30 October 1986
FLO/FRU) (SAN!), Sumbing SAN 119727 (24 July 1987 FLO) (SAN!); Witte Range,
Shang Liang logging area, Asik SAN 118355 (11 October 1986 FLO) (KEP!, SAN!);
UIu Sg Tinagalan F.R., Fedilis & Asik SAN 113139 (21 November 1985 FRU) (SAN!);
1 mile E of Tambunan, on path to Kaingaran village, 2600 ft [702 m], Wood & Wyatt-
Smith A 4444 (29 August 1954 FLO/FRU) (KEP!, L!, SING!); Crocker Range, Mile 16
[Km 25.7], Kimanis Road, Fedilis SAN 113849 (21 August 1986 FRU) (SAN!).
Pensiangan: Pensiangan Kayu F.R., Asik 113043 (14 November 1985 FRU) (SAN!),
Asik SAN 136016 (22 July 1992 FRU) (SAN!), Fedilis & Sumbing SAN 110346 (16
October 1985 FRU) (KEP!, SAN!, SAR!), Fedilis SAN 110390 (19 October 1985
FRU) (SAN!); Sapulut F.R., Suali & Sigin SAN 101363 (24 October 1983 FRU)
(SAN!); logged area Ponontomon, Asik SAN 128039 (25 September 1989 FLO)
(SAN!). Tenom; Mendalom F.R., Saikeh SAN 72012 (13 January 1972 FRU) (L!,
SAN!); Paling-paling Hills, 700–800 m, Coode & Lamb Coode 7134 (2 January 1992
FLO/FRU) (SAN!); Crocker Range, George SAN 31932 (6 December 1962 FLO)
(SAN!); Tomani, 600 ft [183 m], Pitty SAN 63154 (19 August 1968 FRU) (SAN!).
Papar: Mandahan F.R., Ulu Sg Puntih, Fedilis & Sumbing SAN 127555 (27 February
1989 FLO) (SAN!), Bukit Mandahan F.R., 100ft [31 m], Diwol & Tarmiji SAN 80042
339

(9 June 1975 FLO) (SAN!). Kuala Penyu: Kapayan F.R., 35 m, Ag Amin SAN
127253 (17 August 1993 FRU) (SAN!); along Jalan Berangkok, 5 m, Pereira,
Noorsiha, Diwol & Postar JTP 598 (3 March 1999 ST) (SAN!); Bukit Kapayan,
Othman SAN 27347 (14 January 1962 FRU) (SAN!, SAR!); forest near Kg Memalad,
35 m, Pereira, Diwol & Ubaldus JTP 660 (1 December 1999 FRU) (SAN!), JTP 662 (1
December 1999 FRU) (SAN!), Pereira, Noorsiha, Diwol & Postar JTP 701 (3 March
1999 ST) (SAN!), JTP 702 (3 March 1999 ST) (SAN!). Beaufort: Beaufort Hill, 100
ft [31 m], George SAN 38091 (7 March 1962 FLO) (SAN!, SAR!). Sipitang:
Mendalong, Lumaku F.R., Saikeh SAN 72373 (18 October 1972 FLO) (L!, SAN!);
Melalia F.R., Ag Amin SAN 127018 (12 October 1990 FLO) (SAN!); Klias F.R., Diwol
& Karim SAN 77824 (24 July 1973 FRU) (SAN!). BRUNEI. Belait: Mukim Labi,
Bukit Teraja, Chin, Joffre et al. 4631 (12 November 1995 FRU) (KEP!, SING!); along
the new road from Sg Mau to Merimbun, near Sg Merangking, Wong WKM 377 (24
August 1988 FRU) (KEP!, L!, SAN!, SAR!, SING!); Sg Liang Forest Recreational
Park, Arboretum, 20 m, Ariffin & Ibrahim BRUN 17403 (18 April 1996 FLO)
(BRUN!), Wong 305 (28 July 1988 FRU) (BRUN!, KEP!, SAN!, SING!); Andulau
F.R., 150 ft [46 m], Ashton BRUN 48 (24 May 1957 FRU) (BRI!, BRUN!, KEP!, L!,
SAR, SING!), Compt. 7, near Sg Liang, 50 m, Coode & Wong Coode 6769 (14 April
1990 FLO) (BRUN!), Compt. 6, 150 ft [46 m], Smythies, Wood & Ashton SAN 17483
(25 April 1957 FRU) (BRI!, BRUN!, KEP!, L!, SING!), Compt. 19, Puff Ch. Puff
890725-2/1 (25 July 1989 FLO/FRU) (BRUN!, WU!). Tutong: Tanjung Maya, Bukit
Udal, Bukit Kukub, Hussain, Suhaili, Ibrahim & Clayton BRUN 16470 (22 December
1994 FRU) (BRUN!); Nong Maling, Bukit Sukang, Tasik Merimbun, Bernstein JHB
504 (3 August 1993 FRU) (BRUN!, K!); Andulau F.R., at Bukit Besong, Niga NN 231
(8 August 1991 FRU) (BRUN!, KEP!, SAN!, SAR!, SING!). Brunei-Muara: Brunei
Town, Van Niel 3905 (25 May 1985 FLO/FRU) (L!). Temburong: Ulu Temburong
340

River, at landing point LZ 298, Wong s.n. (19 September 1988 ST) (BRUN!),
Temburong River, just upstream from Wong Nguan rapids, 220 m, Coode & Wong et
al. Coode 6601 (7 April 1990 FLO) (BRUN!, L!, SAN!, SAR!, SING!); Amo, Ulu
Belalong L.P. 382, south facing slope below campside, 500 m, Coode 7894 (23 January
1994 FLO/FRU) (BRUN!, K!, KEP!, SAN!, SAR!, SING!). SARAWAK: No locality,
Beccari 1455 (1865–68 FLO) (K!). KUCHING DIV.: Lundu, Mt Gading, 280 m,
Chai S 18495 (20 July 1963 FRU) (KEP!, SAN!, SAR!, SING!), 750 ft [228 m], Chew,
CWL 610 (20 July 1963 FRU) (K!, L!, SING!), 1700 ft [518 m] , James S 34241 (19
September 1974 FRU) (SAN!, SAR!), Mt Kanji, 600 m, Dayang Awa & Ilias S 44336
(18 April 1984 FLO) (KEP!, L!, SAN!, SAR!); Mt Pueh, Batu River, Hj Othman S
56650 (24 March 1989 FLO) (SAN!, SAR!); Bau, Bengoh range, Pangkalan Tebang,
Mohizah, Yahud et al. S 7386 (7 May 1996 FRU) (SAR!). SRI AMAN DIV.: Lubok
Antu, Batang Ai National Park, Tinting Kemuyang, Nanga Lelayang, 670 m, Rantai et
al. S 87158 (23 July 2002 ST) (SAR!). KAPIT DIV.: Batang Balleh, Bukit Melatai,
Melatai River, 380 m, Yii S 48528 (25 April 1985 FLO) (SAR!), Belaga, Bakun HEP,
Berangan River, Pereira, Saw, Mat Asri, Lai & Rantai JTP 255 (18 August 1995 FRU)
(SAN!, L!). MIRI DIV.: Miri, Lambir National Park, Lambir Hills, Sibat S 24308 (21
May 1966 FLO) (KEP!, SAN!, SAR!, SING!), Rena S 40289 (19 September 1978
FRU) (KEP!, SAN!, SAR!), Lee S 46521 (30 June 1983 FRU) (KEP!, SAN!, SAR!),
Mile 18 [Km 29], 500 ft [152 m], Rena S 39448 (14 May 1978 FRU) (KEP!, SAN!,
SAR!), Ulu Sg. Jangkang, Abg Mohtar et al. S 47135 (25 October 1983 FRU) (KEP!,
SAN!, SAR!), Ecological Research Plot, Rantai, Jugah & Manggi S 66645 (22 July
1993 ST) (SAR!), Ulu Sg Lepoh, 20 th mile Miri-Bintulu Rd., 1100 m, Ilias & Yeo S
38304 (29 October 1976 FRU) (KEP!, SAN!, SAR!), Ulu Luak, S. Setap Rd., 50 m,
Othman S 21335 (16 May 1964 FLO/FRU) (KEP!, SAN!, SAR!, SING!), Baram, Hose
589 (April 1895 FLO) (K!), Niah National Park, Sg Sekaloh, 450 ft [137 m], Yii S
341

40138 (9 April 1979 FLO) (KEP!, SAN!, SAR!); Marudi, outskirts of Marudi, N of
town, Marudi F.R., 30 m, Fuchs 21253 (10 August 1963 FRU) (K!, L!, SAR!), Bok
Tisam, Sg. Peti, Sibat S 22802 (28 April 1965 FLO) (KEP!, SAN!, SAR!, SING!).
LIMBANG DIV.: Limbang, Ulu Sg Limbang, Sg Sipayang, logging area, 610 m,
Julaihi, Vincent et al. S 76667 (7 September 1997 FRU) (SAR!); Lawas, Kota F.R.,
near Sg Telau, 1000 ft [328 m], Chai & Ilias S 30689 (20 October 1971 FRU) (SAN!,
SAR!, SING!). KALIMANTAN. E KALIMANTAN: Kutai, sine coll. bb 13008
(23 November 1928 ST) (BO!), Boschproefstation bb 13009 (23 November 1928 ST)
(BO!), E Kutai, Sg Susuk region, 20 m, Kostermans 5536 (29 June 1951 FLO/FRU)
(K!, L!, SING!), E Kutai, Sangkulirang, 75 m, Boschproefstation bb 11880 (18
September 1927 ST) (BO!), 50 m, Boschproefastation bb 11947 (24 September 1927
FLO) (BO!), Susuh River, 20 m, Kostermans bb 34721 (29 June 1951 ST) (L!); W
Kutai, 5 m, Neth. Ind. For. Service bb 16269 (1 December 1931 FLO) (L!); Berau, 50
m, sine coll. Bb 12091 (23 October 1927 ST) (BO!), Mt Ilas Bungaan, 300 m,
Kostermans 13911 (17 September 1957 FLO/FRU) (K!, L!, SING!); KPC area road
from Camp Bravo Charlie to Sanggatta, 75 m, Ambri et al. AA 1601 (31 March 1996
FLO) (BO!, L!, SAN!); N of Balikpapan, Sg Wain, 60 m, Kostermans 4346 (1 October
1950 FLO) (K!, L!, SING!), 40 m, Kostermans (30 August 1950 FLO/FRU) (L!,
SING!); Samarinda, Loa Lempong Blajan River, Nedi 730 (18 March 1955 FRU) (K!,
L!), Loa Djanan, W Samarinda, 30 m, Kostermans 6614 (26 April 1952 FLO/FRU)
(BRI!, K!, L!, SING!); Wanariset research area, LBN plot, 50 m, Van Balgooy 6094
(19 September 1991 FLO/FRU) (L!), Wanariset location area, 50 m, Ambri & Ariffin W
206 (25 May 1990 FRU) (L!); Tarakan, from old forest garden, Meijer s.n. (1
November 1953 FLO) (BO!). C KALIMANTAN: Sintang, HPH Km 85–87, 110 m,
Church, Mahyar, Indah, Ismail & Hamzah 1260 (1 May 1994 FRU) (L!, SING!),
headwater of Kahayan River, 5 km NE of Haruwu village, ridge W of Nyoohay
342

tributary, 350 m, Burley, Tukirin et al. 370 (23 March 1988 FLO/FRU) (K!, KEP!, L!,
SAR!, SING!); Sampit River, near Kuala Kuajan, 20 m, Kostermans 8062 (2 August
1953 FRU) (L!). W KALIMANTAN: Mt Bentuang area, 5–10 km N of Masa village,
150 km NE of Pontianak, Sengman, 150 m, Burley, Tukirin et al. 2459 (10 June 1989
FRU) (K!, KEP!, SING!), Muara Ilay, Kac. Beduai, Sanggau, Plot M3, 50 m, Suzuki,
Koike & Nome K 3089 (19 September 1991 FLO/FRU) (L!).
MALAY
PENINSULA
SUMATRA
Fig. 73. Distribution of “Rothmannia pseudoternifolia var. pseudoternifolia”.
343
BORNEO

Fig. 74. Proposed isotype specimen of “Rothmannia pseudoternifolia var. pseudoternifolia”,
Teysmann 19411, Insula Liengga, prope Sapientjan (L).
344

“Rothmannia” pseudoternifolia var. hispida [new variety description needed].
Rothmannia sp. 1 sensu Coode et al., Checkl. Flow. Pl. & Gynm. Brunei (1996) 290,
pro parte, quoad Puff 900810-1/1.
TYPE (to be designated): Diwol SAN 110731, Sabah, Sandakan, Segaliud (22 August
(Fig. 76)
2003 FLO/FRU) (holotype SAN!; isotypes K!, L!).
DISTRIBUTION. Endemic to Borneo (Sabah, Sarawak, Brunei and
Kalimantan) (Fig. 75).
SPECIMENS EXAMINED.
BORNEO. SABAH. Tawau: Kalabakan, Gunung Rara F.R., about 20 miles NW of
Luasang camp, 1200 ft [366 m], Shea SAN 75601 (20 April 1972 FLO/FRU) (SAN!,
SAR!, SING!); Tawau Hills Park, near hanging bridge, 260 m, Pereira, Ubaldus &
Postar JTP 659 (25 November 1999 FLO) (SAN!), trail to Hot Springs, c. 2100 m from
Park HQ, 740 ft [225 m], Pereira, de Wilde, de Wilde-Duyfjes & Postar JTP 666 (12
January 2000 FLO/FRU) (SAN!); Madai F.R., Mansus et al. SAN 117723 (20 June
1987 FLO) (L!, SAN!); Merotai Besar, Aban SAN 31328 (21 September 1962 FRU)
(SAN!). Lahad Datu: Tabin Wildlife Sanctuary, along Hatton trail, Pereira et al. JTP
486 (15 June 1997 FLO) (L!, SAN!). Labuk Sugut: Segaliud, KTS Arboretum, 200
m, Diwol SAN 110731 (22 August 2003 FLO/FRU) (holotype SAN!; isotypes K!, L!);
Segaliud Lokan F.R., Mile 41, Labuk Rd., RP 295, Leopold & Patrick SAN 72690 (26
February 1971 FLO) (SAN!); Ulu Sapi F.R., Pereira et al. JTP 851 (16 July 2002 ST)
(SAN!). Kinabatangan: Sukau, KOCP plots, Pereira & Postar JTP 732 (1 March
2000 ST) (SAN!), Pereira, Nilus & Postar JTP 733 (19 May 2000 FLO) (SAN!);
Gomantong logged area, Joseph SAN 122785 (22 January 1988 FRU) (SAN!);
Tangkulap F.R., Leopold SAN 144521 (10 April 2004 FLO) (SAN!); Tongod, Ulu Sg.
Pinangah, Sigin SAN 107421 (22 October 1984 FRU) (SAN!); Tongod, Karamuak,
345

Hap Seng Plantation Road to Sg Tangkulap, 100 m, Diwol SAN 90310 (14 June 1982
FRU) (SAN!). Pitas: Paitan F.R., George SAN 121284 (15 May 1987 FLO) (KEP!,
SAN!). BRUNEI: Belait: Ingei River, upstream from the confluence with Belait
River, Wong 608 (1 January 1989 FRU) (BRUN!), Belait River, around Kuala Ingei, 30
m, Puff et al. Ch. Puff 900810-1/1 (10 September 1990 FRU) (BRUN!, KEP!, L!,
SAN!, SING!). SARAWAK. KAPIT DIV.: Kapit, Bukit Raya, Hj Suib S 22274 (13
November 1964 FRU) (KEP!, SAN!, SAR!, SING!), 950 ft [290 m], Jugah S 23875
(16 October 1965 FRU) (KEP!, SAN!, SAR!, SING!), 600 ft [183 m], Au S 23938 (25
October 1965 FRU) (KEP!, SAN!, SAR!, SING!), 500 ft [152 m], Wright S 23899 (19
October 1965 FRU) (KEP!, SAN!, SAR!, SING!); Pelagus, 80 m, Lee S 40666 (12 July
1979 FRU) (KEP!, SAN!, SAR!), Ulu Pelagus, Sg Iran, 200 ft [61 m], Smythies S 9465
(14 September 1958 FRU) (SAR!, SING!), Balleh, Ulu Sg Entuloh, Nanga Sekawi, Hj
Othman, Rantai, Jugah, Jegong & Johnny S 57763 (20 September 1989 FRU) (SAN!,
SAR!), Batang Balleh, Sg Mengiong, Sg Entulu, 320 m, Lee S 54646 (19 July 1987
FRU) (SAN!, SAR!); Kawasan Perlindungan Maias Lanjak-Entimau, Sg Joh, 215 m,
Kulip & Salleh et al. SAN 133608 (30 June 1993 FRU) (K!, L!). MIRI DIV.: Miri,
Niah F.R., Coupe 6, Anderson S 31662 (23 April 1972 FLO) (SAR!, SING!).
BINTULU DIV.: Bintulu, Nyabau catchment area, 500 ft [ 165 m], Sibat S 24630 (23
June 1966 FRU) (KEP!, SAN!, SAR!, SING!); Dataran Tinggi Merurong, Sg Jejalong,
350 m, Othman, Yii et al. S 48870 (11 October 1984 FRU) (KEP!, SAN!, SAR!),
Tubau, Bukit Sekiwa, 200 m, Abg Mohtar et al. S 52768 (26 August 1986 FRU)
(SAN!, SAR!), Tubau, Bukit Sekiwa, Ulu Jejalong, 300 m, Abg Mohtar S 53956 (2
September 1986 FRU) (SAR!), 300 m, Abg Mohtar S 53949 (2 September 1986 FLO)
(SAN!, SAR!). KALIMANTAN. E KALIMANTAN: Nunukan Island, southern
part of Nunukan Island, 20 m, Kostermans 9224A (3 January 1954 FLO/FRU) (BO!,
K!, L!); N of Tarakan, Meijer 1932 (November 1953 FLO) (BO!, K!, L!), Berau,
346

INHUTANI 1 area, road to Tj. Redeb, c. 8 km from Labana (base camp INHUTANI 1),
logged over in 94/95, 50 m, Kessler et al. Berau 335 (18 February 1997 FLO) (L!,
SAN!), Berau, INHUTANI I area, Labanan, Plot 4, 20 m, Goverse & Adriansyah 495
(18 Marhc 1997 FLO/FRU) (KLU!, L!, SAN!); Jalan Pipa, P.T. Kelian Equatorial
Mining Area, 160 m, Kessler et al. PK 2066 (17 March 1997 FRU) (L!). W
KALIMANTAN: Betung Kerihun National Park, along Mendalom River, 160 m,
Sidiyasa et al. 1957 (2 March 2000 FRU) (BO!).
BORNEO
Fig. 75. Distribution of “Rothmannia pseudoternifolia var. hispida”.
347

Fig. 76. “Rothmannia pseudoternifolia var. hispida”. A, leafy branch with inflorescences. B, close-up
of hairs on the upper leaf surface, i.e., hispid. C, close-up of lower leaf surface. D, calyx limb-tube
hispid. E–F, longitudinal section of flower. G, outer surface of corolla lobes. H, fruit.
All from Diwol SAN 110731 (holotype, SAN).
348

15. “Rothmannia” ridsdalei [new species description needed].
Rothmannia sp. 1, Beaman & Anderson, The Plants of Mount Kinabalu (2004) 340.
TYPE (to be designated): Ridsdale PBU 396, Indonesia, Kalimantan, Central
(Fig. 78)
Kalimantan, P.B.U. base camp and environs, side river of Rekut, riverside forest
(6 June 1990 FLO/FRU (holotype L!; isotypes BO!, K!, KEP!, SAN!).
Medium-sized tree, to 15 m high, to 15 cm diameter. Bark dark brown to dark
grey; inner bark orange. Sapwood yellow. Twigs glabrous. Branch leaves typically in
groups of 3’s, each trifoliate group based on a distal node with only one leaf normally
developed (the other vestigial or very much reduced in size), and the proximal node
with a pair of normal leaves. Stipules shortly ovate-triangular, 1.5−4 mm long, 3−6
mm wide, glabrous outside; apex obtuse, acute to short-acuminate, to 1 mm long.
Young leaves on shoot tips scantily pubescent on lower surface. Mature leaves with
petiole (0.3−)0.5–1.5 cm long, 1−2 mm thick, glabrous; lamina obovate, elliptic to
oblanceolate, 8−21−(26) cm long, 3.5−9(−13) cm wide, upper surface glabrous, lower
surface glabrous to glabrescent; leaf base cuneate, attenuate, rounded to unequal; leaf
apex short-to-long-acuminate, acumen to 1.3 cm long; thin-coriaceous to chartaceous
when dry, bullate on the upper leaf surface when fresh; midrib on upper side slightly
raised, glabrous to glabrescent, on lower side raised, glabrous to sometimes pubescent;
secondary veins 6−12 pairs, on upper side slightly raised, glabrous, on lower side
raised, glabrous to scantily puberulous or pubescent; tertiary venation visible as simple
connections between pairs of secondary veins or invisible. Inflorescences: peduncle
0.1−0.6 cm long, 1.5−3 mm thick, glabrous to glabrescent; habit compact, unbranched
or with 1 distinct branching order, rachis 0.1−0.3 cm long, primary branches along the
rachis 1 pair; 0.1−0.3 cm long; bracts ovate, 1.5−2 mm long, 2−3 mm wide, glabrous,
349

hairy only at the margin. Flowers (1−)2−3 per cyme; pedicels c.1 mm long, 1−1.5 mm
thick; calyx tube 6−12 mm long, 5−8 mm wide, glabrous to scantily puberulous, hairs
appressed, surface visible, calyx limb cup-shaped; calyx lobes 5, short- triangular,
0.5−2 mm long, glabrous but scantily hairy at the margin; calyx limb cup-shaped;
corolla hypocrateriform, white, the tube (8−)10.5−14.5 cm long, 6−9 mm wide at the
throat, 2−3 mm wide at the base, non-speckled inside, outer surface glabrous to
glabrescent, inner surface with hairy zone covering part or all of the tube from the rim
of the corolla mouth downwards; corolla lobes 5, ovate-elliptic to lanceolate, 2.3−4.5
cm long, 0.6−1.7 cm wide, outside scantily puberulous at the margin or fringes of the
lobes, inside glabrous; anthers 5, fully included within the tube (inserted at the upper
⅓), 6−12 mm long; style and stigma 60−80 mm long, included but reaching less than ⅔
way along the corolla tube and well below the level of the anthers, style smooth,
glabrous. Fruits globose, 6−11.5 cm long, 5−11 cm wide, glabrous, drying smooth;
fruit wall 3−5 mm thick, very grainy; peduncle 0.1−0.3 cm long, 5−11 mm thick.
Seeds slightly to much flattened and broadly ovate or suborbicular to narrowly
triangular, 10−14 mm × 8−10 mm.
NOTES. Vegetatively, this species is similar to “R. merrilliii” but differ in the
few characters mentioned in the key to Malesian taxa of the Rothmannia complex.
VERNACULAR NAME. Sarawak: Eloh sa’nieng (Kelabit).
DISTRIBUTION. Endemic to Borneo (Sabah, Sarawak, Brunei and
Kalimantan) (Fig. 77).
HABITAT. From lowland and hill forest to lower montane forest, frequently
near rivers, to 1500 m alt.
SPECIMENS EXAMINED.
350

BORNEO. SABAH. Kota Belud: Kg Kiau Nuluh, Jusimin 369 (22 April 1993 FRU)
(SNP!), 695 m, Pereira, Noorsiha et al. JTP 585 (24 February 1999 ST) (SAN!);
Kandasan, 10 miles W of Ranau, 3500 ft [1067 m], Wood & Kapis SAN 16337 (14
December 1955 FRU) (BRI!, L!, SING!). Ranau: Mile 37.5 [Km 60.3], Ranau Rd.,
Mikil SAN 29250 (23 February 1962 FRU) (L!, SAN!, SAR!), Ranau Rd., 4550 ft
[1387 m], Badak SAN 32341 (10 November 1962 FLO/FRU) (L!, SAN!); Mile 38 [Km
11.6], Ranau Rd., 4500 ft [1371 m], Tai SAN 42535 (14 January 1965 FRU) (K!,
SAN!, SNP!); Sosopodon F.R., 1360 m, Pereira, Noorsiha et al. JTP 587 (25 February
1999 ST) (SAN!); Tenompok: below Tenompok, 4700 ft [1432 m], Carr 27221 (3
May 1933 FRU) (SING!), Clemens s.n. (March 1932 FLO) (BM!), 5000 ft [1524 m],
Clemens & Clemens 28522 (22 February 1932 ST) (L!), Clemens & Clemens 28713
(March 1932 FLO) (BM!, K!, L!), Clemens & Clemens 29661 (10 May 1932 FRU)
(BO!, K!, L!), Clemens & Clemens 29661A (April–May 1932 FLO) (BO!, K!, L!),
Pereira & Sugau JTP 591 (26 February 1999 ST) (SAN!), 800 m, de Wilde et al. SAN
142230 (8 December 2003 ST) (SAN!); Dallas: 3000 ft [914 m], Clemens & Clemens
26365 (12 September 1931 FRU) (K!, L!, SING!), Clemens & Clemens 27351 (2
December 1932 FRU) (K!, L!), Clemens & Clemens 29251 (11 April 1932 ST) (BM!,
K!, L!), Clemens & Clemens 29748 (26 May 1932 ST) (K!, L!), Clemens & Clemens
30345 (1931–32 ST) (K!); Mt Kinabalu: Ulu Langanani, 3800 ft [1158 m], Chew,
Corner & Stainton RSNB 1724 (11 August 1961 FRU) (BO!, K!, L!, SAN!, SING!);
Penibukan, Penibukan ridge, 4000 ft [1219 m], Clemens & Clemens 51698 (September
1933 FRU) (BM!), Dahobong river, 3500 ft [1067 m], Carr SFN 26382 (5 March 1933
FRU) (SING!), 900 m, Nooteboom & Aban 1510 (10 March 1970 FLO) (K!, L!,
SAN!), 4000–5000 ft [1219−1524 m], Clemens & Clemens 31635 (15 February 1935
FLO) (BO!, K!, L!); 4000 ft [1219 m], Clemens & Clemens 32201 (17 March 1933
FLO/FRU) (K!, L!), Penibukan ridge, 4000 ft [1219 m], Clemens & Clemens 40494 (28
351

September 1933 FLO) (K!, L!). Tambunan: Crocker Range, Km 64, Kota Kinabalu–
Tambunan Rd., 1250 m, Beaman et al. 7318 (30 October 1983 FLO) (L!); Trusmadi,
Joseph et al. SAN 109441 (20 May 1986 FRU) (SAN!); Mahua 1340 m, Rimi et al. SP
19265 (28 August 2003 FRU) (SAN!). Sipitang: Long Pasia, Postar & Davidson SAN
145771 (3 September 2002 FLO) (SAN!). BRUNEI. Temburong: Amo, Kuala
Belalong, Sg Belalong, 60 m, Dransfield et al., JD 7077 (15 February 1992 FRU)
(BRUN!, K!, SAN!), Eastern slope of Bukit Bangar, 10–30 m, Hotta 13095 (19 January
1964 FRU) (SAR!). SARAWAK. SRI AMAN DIV. Lubok Antu: Sg Jelok, near
Bukit Sengkajang, Lanjak-Entimau, 2100 ft [640 m], Chai S 34038 (18 March 1974
FRU) (KEP!, SAR!). MIRI DIV. Miri: Sg Medalam, Gunung Mulu National Park,
Chai S 39473 (13 October 1977 FLO/FRU) (KEP!, SAN!, SAR!); Sg Tinjar, Dulit
Range, 860 m, Dayang Awa & Yii S 46788 (16 October 1983 FRU) (KEP!, L!, SAN!,
SAR!). LIMBANG DIV. Limbang: Lawas, Kota F.R., Ulu Lawas, 800 ft [244 m],
Chai & Ilias S 31115 (21 October 1971 FRU) (SAR!). KAPIT DIV. Kapit: Belaga,
Rejang, Haviland 2156 (November 1892 FLO) (K!, SING!), Sg. Entulu, Sg Mengiong,
near stream, Lee S 54630 (18 August 1987 FLO) (SAR!), Mujong, Temiai River, Hose
mountain, submontane forest, 1260 m, Lai, Jugah & Awang S 64099 (6 December
1991 FRU) (L!, SAN!, SAR!); Baram, Kelabit Highland, at the foot of Apo Duat, 1300
m, Sie S 35470 (12 November 1974 FRU) (SAN!, SAR!). KALIMANTAN. E
KALIMANTAN: W Kutai, near Mt Kemoel, 1000 m, Endert 3763 (2 October 1925
FRU) (BO!, L!), Central Kutei, Belajan River, Gunung Kelopok, near Tabang, 50 m,
Kostermans 10496 (16 April 1955 FRU) (A, K!, L!). C KALIMANTAN: P.B.U. base
camp and environs, side river of Rekut, Ridsdale PBU 396 (6 June 1990 FLO/FRU)
(holotype L!; isotypes BO!, K!, KEP!, SAN!).
352

BORNEO
Fig. 77. Distribution of “Rothmannia ridsdalei”.
353

Fig. 78. “Rothmannia ridsdalei”. A, leafy branch with inflorescences. B, calyx tube scantily
puberulous. C, external of flower. D–E, longitudinal section of flower. F, outer surface of corolla
lobe. G, anther. H, a fruit with a mass of seeds.
A from Ridsdale PBU 396 (holotype, L), B–G from Clemens & Clemens 32201 (L), and H–I from
Carr 27221 (SING).
354

16. “Rothmannia” schoemannii (Teijsm. & Binn.) Tirveng., Nord. J. Bot. 3
(1983) 469; Wong, Malay. Nat. J. 38 (1984) 49; Tree Fl. Malaya 4 (1989) 406; Ng,
Manual of forest fruits, seeds and seedlings, Malay. For. Rec. 34 (1992) 491, Figs.
809–810; Lemmens & Wulijarni-Soetjipto, Plant Resources of South-East Asia 3
(1992) 151.
NEOTYPE (to be designated): Koorders Kds. 6859 β, Res. Bantam (Bogor sheet no.
BO-1334900) (16 July 1892 FRU) (neotype BO!) (isoneotype K!).
Basionym: Gardenia schoemanii Teijsm. & Binn., Nat. Tijdschr. Ned. Ind. 3 (1852)
327; Teisjm. & Binn., Catalogus van ’s Lands Plantentuin te Buitenzorg (1854)
98; De Vriese & Dozy, Ned. Kruidk. Arch. 3 (1855) 403; Miq., Fl. Ind. Bat. 2
(1856) 232.
Randia schoemannii (Teijsm. & Binn.) Bakh. f., in Backer, Bekn. Fl. Java 15 (1956)
86; Backer & Bakhuizen, Fl. Java 2 (1965) 310; Corner, Wayside Trees of
Malaya 2 (1988) 648.
Randia exaltata Griff., Notul. Ad Plantas Asiaticas 4 (1854) 262; Kurz, For. Fl. Br.
Burma 2 (1877a) 46, J. Asiat. Soc. Bengal 46 (2) (1877b) 157; Hooker, Fl. Br.
India 3 (1880) 114, pro parte quoad Kew. Dist. 2826, excl. Kurz s.n. (=
Rothmannia pulcherrima (Kurz) Tirveng.); Koord. & Valeton, Bijdr. Booms.
Java 8 (1902) 88 pro parte, excl. Gardenia pulcherrima Kurz in syn.; King &
Gamble, J. Asiat. Soc. Bengal 72 (1903) 214, pro parte, excl. Andaman Islands
(=R. pulcherrima (Kurz) Tirveng.); Brandis, Ind. Trees (1906) 383, pro parte,
excl. G. pulcherrima in syn.; Koord., Exkursionflora Java 3 (1912) 256; Koord.-
Schum., System. Verzeichen. Part I, 1 Fam 270 (1913) 84; Koord. & Valeton,
Atlas der Baumarten van Java 3 (1915) Figs. 527, 528; Pitard, Fl. Gén. Indo-
Chine 3 (1923) 245, pro parte, excl. G. pulcherrima Kurz in syn.; Ridley, Fl.
Malay Penins. 2 (1923) 78; Craib, Fl. Siam. Enum. 2, 1 (1932) 101, pro parte,
355

excl. G. pulcherrima in syn.; Burkill, A Dictionary of the Economic Products of
the Malay Peninsula 2 (1966) 1897.
TYPE: Griffith 962, Myanmar (sine datum FLO/FRU) (holotype K!).
Rothmannia exaltata (Griff.) Bremek., K. Ned. Akad. van Wet.–P 60,1 (1957) 7.
(Fig. 80)
Medium to large trees, to 30 m high, to 45 cm diameter. Bark smooth to
fissured, sometimes flaky, brownish black; inner bark yellowish brown to purple-
brown. Sapwood pale yellow, base of the trunk fluted, up to 3 m high. Twigs
glabrescent to sparsely pubescent. Branch leaves typically in groups of 3’s, each
trifoliate group based on a distal node with only one leaf normally developed (the other
vestigial or very much reduced in size), and the proximal node with a pair of normal
leaves. Stipules triangular, 1.5–3 mm long, 2–4 mm wide, sparsely pubescent outside;
apex acute to short-acuminate, to 0.5 mm long. Young leaves on shoot tips pubescent
on both sides. Mature leaves with petiole 0.1–1 cm long, 0.5–2 mm thick, pubescent,
puberulous to glabrescent; lamina oblanceolate to elliptic, 6.5–15 cm long, 2–6 cm
wide, upper surface glabrous, lower surface scantily puberulous to glabrescent; leaf
base attenuate, cuneate to unequal; leaf apex acute to acuminate, acumen to 0.5 cm
long; thin-coriaceous to chartaceous when dry; midrib on upper side slightly raised to
flat, glabrescent to glabrous, on lower side raised, sparsely pubescent to puberulous;
secondary veins 6–8 pairs, on upper side flat, glabrous, on lower side raised, glabrous
to sparsely puberulous; tertiary venation visible as a much-branched network between
pairs of secondary veins. Inflorescences: peduncle 0.1–0.5 cm long, 2–3 mm thick,
appressed pubescent; habit laxly arranged, with 2–3 distinct branching orders, rachis
0.5–1(–2) cm long, primary branches along the rachis 2–3(–4) pairs, 0.1–0.4 cm long;
bracts ovate, 1–2 mm long, 2–3 mm wide, appressed pubescent. Flowers (5–)6–20 per
356

cyme; pedicels 0.5–2 mm long, 1–1.5 mm thick; calyx tube 3–5 mm long, 3–5 mm
wide, sparsely pubescent on the limb, densely pubescent on the hypanthium, hairs
appressed, surface visible on the limb; calyx limb cup-shaped; calyx lobes 5, short-
triangular, 0.5–1 mm long, appressed sparsely pubescent; corolla campanulate, white,
the tube 2.6–4.5 cm long, 9–20 mm wide at the throat, the upper expanded part of the
tube 2–4 cm long, the cylindric portion at the base 0.6–1.2 cm long and 2–2.5 cm wide
at the base; speckled with purplish spots inside, outer surface sparsely to densely
pubescent, inner surface with hairy zone covering part or all of the tube from below the
level of the anthers downwards; corolla lobes 5, ovate to oblong, 0.7–1.3 cm long, 0.5–
1.2 cm wide, outside sparsely pubescent on the uncovered half of the corolla lobes,
inside glabrous; anthers 5, fully included within the tube or only with tips slightly
protruding (inserted at the upper ⅓), 10–15 mm long; style and stigma 35–41 mm long,
included but reaching just below the corolla mouth and the level of the anthers, style
smooth, sparsely hairy. Fruits globose, 5–7.5 cm long, 3–7 cm wide, glabrous, drying
smooth; fruit wall 3–4 mm thick; peduncle 0.4–0.7 cm long, 5–8 mm thick. Seeds
slightly to much flattened and broadly ovate or suborbicular to narrowly triangular, 10–
13 mm × 8–10 mm.
NOTES. The type specimen of the basionym, Gardenia schoemannii published
by Teijsm. & Binn. in Nat. Tijdschr. Ned. Ind. 3, pg. 327 in 1852 was collected from
Java, Residentie Bantam and named after Dr. C. Schöman. Tirvengadum (1982,
unpublished thesis) indicated the type in a very general way and Wong (1984) assumed
the type was from Bantam. However, no specimens with the Bantam locality collected
before 1852 have been located in BO, K or L, where most of Teysmann’s collections
were deposited. Therefore, the type is either not clearly designated in the original
publication or is missing. Hence, a neotype is to be designated.
357

In Teijsm. & Binn.’s Catalogus van ’s Lands Plantentuin te Buitenzorg (1854),
“Hsskl. e Bantam” (=Hasskarl & Bantam) was indicated under the entry of Gardenia
schomannii. At Leiden, several sheets of Hasskarl’s collections were all bearing the
locality Java (not Bantam). One of these sheets, (Rijskherbarium No. L4505/63; Herb.
Lugd. Bat. 908.217-445) was the presumed type of Wong (1984). Based on Article 9,
paragraph 9.6 of the International Code of Botanical Nomenclature, Vienna Code
(2006), the neotype is selected from one of Koorders’s collections from Bantam which
has “Res. Bantam Afd. Tjiringin bij. Pgr. Tjanmara” stamped on the sheet, although the
collection date (16 July 1892) is clearly much later than the original publication of the
species name.
Vegetatively, this species resembles an Indo-Chinese species, Rothmannia
eucodon. Both species have campanulate corollas. However, “R. schoemannii” is
distinguishable from R. eucodon by the following floral characters: shorter flower
pedicels, 0.5–2 mm long (3–8 mm long in R. eucodon); narrower corolla mouth, 9–20
mm wide (25–35 mm wide in R. eucodon); and shorter calyx tube 3–5 mm long (6–7
mm long in R. eucodon).
VERNACULAR NAMES. Duk kop (Siamese, Thailand); Kelompang gajah,
Poko laga (Malay Peninsula); joyjow (Temuan, Peninsular Malaysia).
(Fig. 79).
DISTRIBUTION. Myanmar, Thailand, Peninsular Malaysia, Sumatra, and Java
HABITAT. Primary to secondary forests in evergreen, lowland and hill forests,
sometimes along streams, near the coast, or in mangroves, to 700 m alt.
USES. Burkill (1966) reported that the timber is heavy but not durable; from
the fruits are derived a black dye.
SPECIMENS EXAMINED.
358

MALESIA. JAVA. No locality: sine coll. s.n. (sine datum ST) (L!); H.L.B s.n. (sine
datum ST) (L!), sine coll. E 23 (September 1890 FLO) (BO!). WEST JAVA: Res.
Preanger, Afd. Soekeboemi, Plaboecan ratoe, Koorders Kds. 6856 β (27 May 1890 ST)
(BO!), Koorders Kds. 6857 β (6 June 1890 FLO) (BO!, SING!), Koorders 6858 β (23
June 1890 FRU) (BO!), Koorders Kds. 12556 β (27 August 1891 ST) (BO!), Koorders
Kds. 15735 β (20 October 1894 ST) (BO!), Koorders Kds. 33095 β (15 April 1899 ST)
(BO!); Res. Bantam, Afd. Tjaringan, Koorders Kds. 6859β (16 July 1892 FRU)
(neotype BO!) (isoneotype K!); Pgr. Palaboean, Koorders Kds. 42702 β (30 April
1893 ST) (BO!); Tanjung Lajar, near the light house, Udjungkulom Reserve, Mt
Pajung, 100 m, Kostermans 122 (13 November 1960 FLO) (L!), near lighthouse, 10 m,
Kostermans 21825 (9 December 1964 FRU) (L!); 50 m, Kostermans & Kuswata 51 (17
August 1960 FLO/FRU) (KEP!, L!, SAN!, SING!); Res. Batavia, Depok, Beumee 6099
(31 January 1924 ST) (BO!), Koorders Kds. 31049 (26 August 1898 FRU) (BO!);
Bogor: cultivated in Hort. Bog., sine coll. C.H.B V.D. 51 (sine datum FLO) (BO!, L!),
sine coll. 13381 (sine datum ST) (BO!); Res. Batavia, Afd. Buitenzorg, Koorders Kds.
24053 β (27 August 1896 FLO) (BO!, K!). EAST JAVA: No locality, Teysmann s.n.
(10 February 1905 ST) (P!). SUMATRA: LAMPUNG PROV.: NW of Kota Agung,
350–400 m, Jacobs 8288 (9 May 1968 FRU) (KEP!, L!, SING!). NORTH SUMATRA
PROV.: Sekundur Forest Reserve, E side of Gunung Leuser National Park, Langkat,
base camp at Aras Napal, upper Besitang River area, 100 m, de Wilde 21276 (5 August
1991 FRU) (L!). PENINSULAR MALAYSIA. KEDAH: Rawei Island, Ridley
15867 (1911 FLO) (K!, SING!); Koh Mai F.R., Kiah SFN 35199 (6 April 1938 FRU)
(BO!, SING!); Sik, Enggang F.R., Kochummen KEP 78903 (7 December 1956 FLO)
(L!); North Kedah, Compt. 42, Ulu Muda F.R., Chan FRI 6736 (18 January 1969
FLO/FRU) (L!, SAR!, SING!). PERAK: junction of Sg Kewa and Sg Perak,
Whitmore FRI 0563 (10 July 1966 FRU) (KEP!, L!, SING!); Tapah Hills, hillside, 3000
359

ft [914 m], Ng FRI 1306 (10 July 1966 FRU) (L!, SING!). PULAU LANGKAWI:
Pulau Bumbun Besar, Van Balgooy 2299 (19 February 1975 FRU) (UKMB!); Mat
Chinchang F.R., Khairuddin FRI 31820 (24 February 1983 FRU) (KEP!, L!, SAN!,
SING!). PENANG (PULAU PINANG): Penara Bukit, 1000ft [305 m], Curtis 793
(April 1886 FLO/FRU) (K!, SING!); waterfall garden, Burkill s.n. (February 1916
FLO/FRU) (BO!, SING!), Curtis s.n. (March 1890 FLO) (SING!); Pulau Buetong,
Curtis =793 (March 1892 FLO) (SING!); Penang Botanic Gardens: Abdul Kadir s.n.
(17 March 1936 ST) (SING!), Henderson 460 (1 March 1938 ST) (SING!), behind
carpenter shed, Md. Noor s.n. (28 September 1918 ST) (SING!). MALACCA: Alvins
739 (sine datum ST) (SING!); Bukit Klanara, Alvines 1147 (13 March 1985 FLO)
(SING!). SELANGOR: Ulu Langat, Ulu Lepok, Gadoh sub Millard 883 (14
September 1958 ST) (L!, SING!); Semenyih, Hume 8403 (8 January 1921 ST)
(SING!); Ulu Gombak F.R., 800 ft [244 m], Ahmad FRI 94460 (18 September 1960
FRU) (KEP!, L!, SING!), Ulu Gombak, 17 miles, 1000ft [305 m], Nur SFN 34229 (25
October 1937 FRU) (SING!); Janda Baik, Pereira et al. JTP 794 (2 September 2000
ST) (SAN!), JTP 795 (2 September 2000 ST) (SAN!). PAHANG: Peromat KL 4367
(6 May 1994 FRU) (KEP!); Taman Negara, Kuala Kenyam, 100 m, Van Balgooy 2586
(30 April 1975 FLO/FRU) (BRI!, L!), Taman Negara, trail to Bumbun, Trenggan
Lodge, Pereira et al. JTP 853 (12 October 2002 ST) (SAN!), trail to Bumbun Tabing,
Pereira JTP 854 (13 October 2002 ST) (SAN!); Pulau Tioman, Corner 29951 (18
August 1935 FRU) (K!, L!, SING!); Ayer Surin, c. 1000 ft [305 m], Henderson 18406
(19 May 1927 FLO/FRU) (K!, SING!); Sg Mai Estate, Jenderok Halt., Kadim &
Mahmud K 7 (26 March 1959 FLO) (KEP!, L!, SAR!, SING!); Lepar F.R., Suppiah
KEP 104992 (17 June 1967 FRU) (L!, SING!); Kuantan, Panching Forest, c. 15 km
north of Panching, 130 m, Ogata 10548 (10 June 1968 FRU) (KEP!, L!), road to Bukit
Tinggi, 1400 ft [427 m], Wong FRI 32091 (13 March 1981 FRU) (KEP!, L!, SAN!);
360

Bentong, Jalan Bukit Tinggi, Pereira et al. JTP 798 (2 September 2000 FRU) (SAN!);
Lancang, Abdul Latiff ALM 83l2 A41 (24 February 1985 ST) (UKMB!).
KELANTAN: Kuala Krai, near Sg Durian F.R., Rahim FRI 104267 (3 May 1967
FLO/FRU) (KEP!, L!, SING!). NEGERI SEMBILAN: Kuala Pilah, Kochummen FRI
16117 (19 June 1970 FRU) (L!, SING!). THAILAND. NARATHIWAT PROV.:
Khao Tan Yong, Chai-anan 45 (5 June 1973 ST) (BKF!). OUTSIDE MALESIA.
THAILAND. No locality. Hamid F.M.S. 2962 (3 February 1919 FRU) (SING!).
SURAT THANI PROV.: Panom, 50 m, Kerr 12421 (25 March 1927 FLO) (BK!, K!);
Kaw Samui, under 50 m, Kerr 12617 (10 April 1927 FLO) (BK!, K!); Kaw Pa- ngan,
Put 1213 (8 November 1927 FRU) (K!); Kaw Tao, under 10 m, Kerr 11151 (29
December 1926 FRU) (BK!, K!), 1300 m, Kerr 12763 (15 April 1927 FLO) (BK!, K!),
5 m, Kerr 12763 A (15 March 1927 FLO) (BM!), Kerr 12800 (18 April 1927 FRU)
(BK!, K!). Trang Prov.: Trang, 50 m, Keith 413 (20 May 1890 FLO) (SING!),
Vanforuk 646 (March 1915 FLO) (BKF!); Kaw Tibang, c. 100 m, Kerr 19079 (23 April
1930 FLO) (BK!); Khao Chong: 81 m, Bunnab 424 (12 March 1966 FLO) (BKF!),
150 m, Maxwell 75–902 (16 August 1975 FRU) (BK!, L!), 200 m, Maxwell 85–157 (2
February 1985 FRU) (UKMB!, SING!), Saard 3281 (sine date FLO) (K!),
Sangkhachand 1925 (27 June 1969 FLO) (BK!), Sangkhachand 2236 (18 December
1969 FRU) (BK!), sine coll. SP 80 (18 March 1969 FLO) (K!). RANONG PROV.:
Ranong, Pengkhlai CP 1310 (18 September 1968 FRU) (BKF!, L!), Ngao waterfall,
sine coll. WN 689 (8 September 1984 FRU) (BKF!); Ka Per, Klongnaka, 50 m,
Niyomdham et al. 303 (14 July 1979 FRU) (BKF!). KRABI PROV.: Kaw Pu, below
50 m, Kerr 18969 (13 April 1930 FLO) (BK!, K!). PANG-NGA PROV.: Pang-nga,
Similum National Park, Koh 4, 45–75 m, Niyomdham & Puudjaa 3335 (26 November
1992 FRU) (BKF!); Kao Praml, Sutheesonn 2474 (28 April 1967 FLO/FRU) (BK!);
Khong Nang Yon, 100 m, Geesink & Santisuk 5063 (30 April 1973 FLO/FRU) (BKF!,
361

L!, P!). NAKHON SI THAMMARAT PROV.: Lansaka Dist., Khao Luang
National Park, Garom Falls, 300 m, Maxwell 85–885 (15 September 1985 FRU) (L!,
UMKMB!). PRACHUAP KHIRI KHAN PROV.: Huai Yang waterfall, S of
Prachuap, Beusekom & Santisuk 2757 (10 February 1970 FLO/FRU) (BKF!, L!, P!).
CHONBURI PROV.: Kow Kieo, 500 m, Maxwell 76–6 (16 January 1976 FRU)
(BK!). MYANMAR. No locality. Griffith 962 (sine datum FLO/FRU) (holotype
K!), Griffith s.n. (sine datum FLO) (K!, P!). South Tennaserim, Theinkun Chaung
forest, 200 ft [61 m], Parkinson 1915 (15 February 1926 FLO) (K!); Mergui distr.,
Dandawur, Lace 4804 (14 April 1909 FLO/FRU) (K!).
Fig. 79. Distribution of “Rothmannia schoemannii”.
362
MALAY
PENINSULA

Fig. 80. Proposed neotype specimen to be designated for Rothmannia schoemannii (Teijsm. & Binn.)
Tirveng. Koorders Kds 6859 β, Res. Bantam (Bogor sheet no. BO-1334900) kept in Bogor Herbarium
(BO).
363

17. “Rothmannia” sundaensis [new species description needed].
TYPE (to be designated): van Balgooy 3946, Indonesia, Sulawesi, between Soroako
and Nickel Plant site, 500 m (10 July 1979 FRU/FLO) (holotype L!).
Rothmannia merrilli sensu Kessler et al., Checklist of woody plants of Sulawesi,
(Fig. 82)
Indonesia, Blumea Supp. 14 (2002) 121, non (Elmer) Ridsdale & J.T.Pereira.
Small tree, to 15 m high, to 25 cm diameter. Twigs glabrescent when young,
becoming glabrous when old. Branch leaves typically in groups of 3’s, each trifoliate
group based on a distal node with only one leaf normally developed (the other vestigial
or very much reduced in size), and the proximal node with a pair of normal leaves.
Stipules triangular to ovate, 2−3(−5) mm long, 4−5 mm wide, glabrous outside; apex
obtuse, acute to cuspidate, to c. 1 mm long. Young leaves scantily pubescent on lower
surface. Mature leaves with petiole 0.2−0.7(−1) cm long, 1−2 mm thick, glabrous;
lamina elliptic, oblanceolate to obovate, (7−)9−16 cm long, (3−)4−9 cm wide, upper
surface glabrous, sometimes shiny, lower surface glabrous; leaf base cuneate, attenuate
to unequal; leaf apex acute to short-acuminate, acumen to 0.5 cm long; thin-to thick-
coriaceous when dry; midrib on upper side flat, glabrous, on lower side raised, sparsely
puberulous to glabrous; secondary veins 5−7(−8) pairs, on upper side flat, glabrous, on
lower side raised, sparsely puberulous to glabrous; tertiary venation visible as simple
connections between pairs of secondary veins. Inflorescences: peduncle 0.3−1.2 cm
long, 2−2.5(−3) mm thick, glabrous; habit laxly arranged, with (1−)2−3 distinct
branching orders, rachis 0.5−1.5 cm long, primary branches along the rachis (1−)2
pairs, 0.5−1 cm long; bracts narrowly ovate, 1−1.5 mm long, 1−1.5 mm wide, sparsely
puberulous to pubescent at the apex. Flowers (4−)7−14 per cyme; pedicels obsolete to
364

1 mm long, c. 1 mm thick; calyx tube 4−6 mm long, 3.5−4 mm wide, glabrous
throughout or puberulous at the limb and glabrous at the hypanthium, hairs appressed,
surface visible; calyx limb cup-shaped; calyx lobes 5−6, short-triangular, 1−2 mm long,
scantily puberulous to glabrous but sparsely puberulous on the margin; corolla
hypocrateriform, white, the tube 2−3.5 cm long, 3.5−4 mm wide at the throat, 1−1.5
mm wide at the base; non-speckled inside, outer surface scantily puberulous, inner
surface with hairy zone covering part or all of the tube from corolla mouth downwards;
corolla lobes 5, elliptic to elliptic-oblong, 1.6−2(−2.5) cm long, 0.5−1 cm wide, outside
scantily puberulous on the uncovered half of the lobes and near the margin, inside
glabrous; anthers 5, fully included within the tube (inserted at the upper ⅓), 5−12 mm
long; style and stigma 13−14 mm long, when included but reaching less than ⅔ way
along the corolla tube, and well below the anthers but 30–35 mm long when partially
exserted, style smooth, glabrous, stigma 10–17 mm long. Fruits globose to obovoid,
3.5−5 cm long, 2.5−4 cm wide, glabrous, drying smooth and sometimes shiny; fruit
wall 1−1.5 mm thick; peduncle 0.5−1 cm long, c. 3 mm thick. Seeds slightly to much
flattened and broadly ovate or suborbicular to narrowly triangular, 9−10 mm × 7−8
mm.
NOTES. This is a species close to “R. merrillii”, with which it is probably
vicariant. Many of the Celebes specimens were misidentified as “R. merrillii”.
However, this species is distinguishable from “R. merrillii” in having typically more
flowers per cyme (7–14, rarely as few as 4) whereas “R. merrillii” has typically 3–5
flower per cyme, rarely reaching 9 flowers per cyme; in its peduncles that are 0.5–1 cm
long (peduncles to 0.3 cm long in “R. merrillii”); the generally shorter corolla tube, to
only 3.5–4 cm long (3.5–7.5(–12) cm long in “R. merrillii”); narrower corolla mouth,
3.5–4 mm wide (5–9 mm wide in “R. merrillii”); narrower corolla tube base, only 1–1.5
mm wide (2–3 mm wide in “R. merrillii”); and 5 anthers 6–10 in “R. merrillii”).
365

There is one collection, SVD 4121 that has flowers with style and stigma
partially exserted from the corolla mouth. The rest of the specimens have shorter
styles, i.e., included and reaching less than ⅔ way along the corolla tube and well
below the anthers. A heterostylous condition may occur in this species, but more
material and observations are needed to substantiate this.
There is one fruiting collection, Sidiyasa & Didi 1885 which is very close to
this species, but differs in having larger mature leaves, 20−25 × 10−12 cm whereas the
typical “R. sundaensis” has smaller leaves, (7−)9−16 × (3−)4−9 cm.
DISTRIBUTION. Endemic to Sulawesi (Southeast, Central and South
Provinces) and Lesser Sunda Islands (West Flores Island) (Fig. 81).
HABITAT. In primary and secondary forests, including heath forests, on sandy
or ultrabasic soils, to 900 m alt.
SPECIMENS EXAMINED.
SULAWESI. No locality. Bosekwezen s.n. (6 June 1925 FLO) (BO!). SOUTHEAST
SULAWESI PROV.: Tawanggo, Kolaka, Mt Poli-polia, 200 m, Prawiroatmadjo &
Maskuri 1449 (22 October 1978 FRU) (SAR!), around Opa swamp, W of Poli-polia,
primary forest, 200–300 m, Prawiroatmodjo & Soewoko 1919 (23 November 1978
FRU) (L!). SOUTH SULAWESI PROV.: en Onderh. Ond. Afd. Malili Oesoe, 100
m, Boschproefstation CEL II 338 (4 April 1931 FRU) (L!), Malili, Boschwezen bb
1925 (sine datum ST) (BO!), Oesoe, 100 m, Maturandong 113 (2 October 1931 FRU)
(BO!), 100 m, Neth. Ind. For. Service CEL II 306 (10 February 1931 FLO/FRU) (BO!,
L!, SING!), 200 m, Neth. Ind. For. Service CEL II 395 (10 April 1934 ST) (L!);
Soroako road, 150 m, Balgooy 3894 (6 July FLO & FRU) (L!); between Soroako and
Nickel Plant site, 500 m, Balgooy 3946 (10 July 1979 FRU/FLO) (holotype L!); Batu
Besi between Tabarano and Larona river, 300 m, Balgooy 4082 (19 July 1979
FLO/FRU) (L!). CENTRAL SULAWESI PROV.: west part of Batudaka Island,
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Kayome, Sidiyasa et al. 1885 (25 December 1999 FRU) (BO!). LESSER SUNDA
ISLANDS. Flores: Schmutz 3606 (26 May 1905 FRU) (L!); Manggarai, 300 m,
Schmutz SVD 4017 (28 January 1978 FRU) (L!), Naga, 900 m, Schmutz SVD 4121 (6
May 1978 FLO) (L!), 450 m, Schmutz SVD 4781 a (25 February 1981 ST) (L!).
FLORES
SULAWESI
Fig. 81. Distribution of “Rothmannia forsteniana” and “R. sundaensis”.
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“Rothmannia
forsteniana”
“Rothmannia
sundaensis”

Fig. 82. “Rothmannia sundaensis”. A, leafy branch with inflorescences. B, close-up of lower leaf
surface showing glabrous to minute pit-domatia on the axils of the midrib and secondary veins. C, calyx
tube puberulous. D–E, longitudinal section of flower. F, outer surface of corolla lobe. G, anther. H,
infructescence. I, seeds.
A–G from van Balgooy 3946 (holotype, L), H–I from Balgooy 4082 (L).
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18. “Rothmannia” uvarioides [combination needed].
Basionym: Randia macromera var. uvarioides Valeton, Bot. Jahrb. Syst. 60 (1926) 91.
NEOTYPE (to be designated): Ledermann s.n., Papua New Guinea, Sepik (1912–13
(Fig. 83)
FLO) (SING!)
Treelet, to 5 m high. Bark brownish green; inner bark straw. Sapwood white.
Twigs scabrid when young becoming glabrescent when old. Branch leaves typically in
groups of 3’s, each trifoliate group based on a distal node with only one leaf normally
developed (the other vestigial or very much reduced in size), and the proximal node
with a pair of normal leaves. Stipules narrowly ovate, 10−23 mm long, 6−10 mm wide,
pubescent outside; apex long-acuminate, 15−20 mm long. Young leaves on shoot tips
scabrid both sides. Mature leaves with petiole subobsolete to 0.8 cm long, 2.5−4 mm
thick, scabrid; lamina broadly elliptic to oblanceolate, (20−)25−35 cm long, 10−15 cm
wide, upper surface glabrous, lower surface glabrous to scabrid; leaf base rounded to
subcordate, sometimes unequal; leaf apex acute to acuminate with a pointed tip,
acumen 0.5−1 cm long; thin-coriaceous when dry; midrib on upper side slightly sunken
to flat, glabrous, on lower side distinctly raised, scabrid to puberulous; secondary veins
10−15 pairs, on upper side flat, glabrous to glabrescent, on lower side raised, scabrid to
puberulous; tertiary venation visible as a much-branched network between pairs of
secondary veins. Inflorescences: peduncle 0.1−0.2 cm long, 3−3.5 mm thick,
pubescent; habit compact, with 2−3 distinct branching orders, rachis 1−1.5 cm long,
primary branches along the rachis 2 pairs, 0.2−0.25 cm long; bracts ovate with a long-
acuminate apex, 8−10 mm long, 3−4 mm wide, pubescent. Flowers 5–10 per cyme;
pedicels 2−3 mm long, 1.2−1.5 mm thick; calyx tube 3−4 mm long, 3−4 mm wide,
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densely pubescent, hairs appressed to suberect, surface visible; calyx limb cup-shaped;
calyx lobes 5, subulate, (9−)10−12 mm long, pubescent; corolla hypocrateriform,
creamy white, the tube (3.5−)4–5 cm long, (3.5−)4−7 mm wide at the throat, 1.5−2 mm
wide at the base; non-speckled inside, outer surface sparsely pubescent, inner surface
glabrous; corolla lobes 5, ovate to elliptic-oblong, 2.5−5 cm long, 1.2−1.5 cm wide,
outside pubescent on the uncovered half of the lobes, inside glabrous; anthers 5, fully
included within the tube (inserted at the upper ⅓), 8−12 mm long; style and stigma
45−50 mm long, included but reaching just below the corolla mouth and the level of or
above the anthers, style smooth, glabrous. Immature fruits elliptic to subglobose.
Seeds unknown.
NOTES. In Valeton’s account, Die Rubiaceae von Papuasien (1926) in
Engler’s Bot. Jahrb. Syst. 60, pg. 91, he proposed the var. uvarioides, which is here
raised to specific rank because it is sufficiently distinct from the typical var.
macromera. Some of the distinct features that differentiate specimens placed under
var. uvarioides from var. macromera are the glabrous to scabrid lower leaf surface in
var. uvarioides (densely velvety hairy in var. macromera); longer stipules, 10–23 mm
long in var. uvarioides (5–8 mm long in var. macromera); colleters inside the stipules
of var. uvarioides are confined to a triangular area with more than 5 rows at the central
part (colleters are arranged in a narrow zone usually of 1–2 rows at sparse intervals, and
along the bottom edge of the stipule base in var. macromera); 5–10 flowers per cyme
in var. uvarioides (more flowers, 11–24 per cyme in var. macromera); and longer
corolla tube, (3.5–)4–5 cm long in var. uvarioides (1–1.5 cm long in var. macromera).
Attempts have been made to locate the holotype of Ledermann’s collection listed in
Valeton’s account from the Berlin Herbarium (where most of Ledermann’s collection
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were kept) and duplicates in the other herbaria, i.e. L, K and SING (Van Steenis-
Kruseman, 1950). The search was futile.
One unnumbered flowering collection made by Ledermann in the Sepik region
during the Sepik Expedition in 1912–13 (the same collecting time and area as the
specimens listed in Valeton’s account) was discovered and examined at the Singapore
Herbarium.
Based on Article 9, paragraph 9.6 of the International Code of Botanical
Nomenclature, Vienna Code (2006), as this is the only specimen that was available for
study and serves to match the original description, it is to be designated as the neotype
for this species.
None of the fruting specimens in this species listed in Valeton had been located
during this study. However, according to Valeton in Bot. Jahrb. Syst. 60 (1926) 91,
“the immature infructescence has short peduncle, with fruits in groups of 3–5, and the
fruits are elliptic or subglobose, glabrescent and crowned by a calyx which at length
falls off; the pericarp is fleshy-coriaceous, thick”. However, the matching of material
to this species is based mainly on the vegetative and flowering characteristics provided
by Valeton (1926).
DISTRIBUTION. Recorded so far only in Papua New Guinea (East Sepik
Province) (Fig. 84).
HABITAT. Recorded in disturbed forest on hill slopes, to 125 m alt.
SPECIMENS EXAMINED.
PAPUA NEW GUINEA. EAST SEPIK PROV. Ledermann s.n. (1912–13 FLO)
(neotype SING!); Garamambu, J.S.W N.G.F. 3749 (21 August 1949 FLO) (L!); near
Ambunti, 300ft [91 m], Hoogland & Craven 10185 (1 June 1966 FLO) (L!), 400ft [122
m], Hoogland & Craven 10249 (9 June 1966 FLO) (L!); Ambuti subdistr., near Wagu,
200ft [61 m], Hoogland & Craven 10375 (29 June 1966 FLO) (L!).
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Fig. 83. Proposed neotype specimen to be designated for “Rothmannia uvarioides”, Ledermann s.n.,
Papua New Guinea, Sepik (SING) (reproduced from Singapore Herbarium).
372

“ ”
Rothmannia anisophylloides
Rothmannia kassamensis
“ ”
Rothmannia macromera
“ ”
Rothmannia nigrescens
“ ”
NEW GUINEA
373
“ Rothmannia uvarioides”
“ Rothmannia papuana”
Fig. 84. Distribution of “R. anisophylloides”, “R. kassamensis”, “R. macromera”, “R. nigrescens”, “R.
papuana” and “R. uvarioides”.
Rothmannia spp. indet
The following specimens were determined as “Rothmannia” but lacked
sufficient material for precise species determination. However two groups can be
discerned based on a combination of characters. Both these groups lack mature
flowering material which yields the most reliable character in distinguishing species
among the Malesian taxa.
“R. sp. indet 1”
PENINSULAR MALAYSIA: PERAK: IPOH, Kinta Hill F.R., Kuala Liyang, 2200
ft [670 m], Ramli KEP 99779 (15 October 1966 FRU) (KEP!). KEDAH: Mahang, G
Inas, Compt. 22, 220 m, Khairuddin FRI 38325 (14 February 1999 FRU) (K!, SING!).
The leaves and fruit surface texture (epidermis flaking into a dense covering of
minute pieces) appear very similar to that in Rothmannia pulcherrima, from the

Andaman Islands. However, this group has very large mature fruits, c. 9 × 8.5 cm that
are lobed when dried, whereas in R. pulcherrima the fruits are smaller, 3.2–4 × 2.5–5
cm and mature fruits are not lobed. In addition, the fruit wall of this taxon is very thick
and leathery, c. 10 mm thick (in R. pulcherrima it is only 2–3 mm thick). It is also
noted in the field label of Khairuddin FRI 38325 that the “corolla tube is 10 cm long”.
In R. pulcherrima, the corolla tube reaches only 3 cm long and has campanulate corolla
form. However, no flowering material was available for confirming the length of the
corolla tube and corolla form in this taxon. This combination of fruit and flower
characters has not been seen in any of the other Malesian species here recognised.
When mature flowering material becomes available for this taxon, it will likely be
shown to be different from the other Malesian species.
“R. sp. indet 2”
PENINSULAR MALAYSIA: SELANGOR: Ulu Selangor, Gading F.R., 1500 ft [457
m], Loh FRI 13404 (22 July 1969 FRU) (SAN!). SUMATRA: JAMBI PROV.:
Bukit Pening Pening, NW part of Tigapuluh mountain range, S of Bukit Hulusipin, 400
m, Laumonier YL 5642 (18 January 1984 FRU) (L!). CENTRAL SUMATRA
PROV.: W Indragiri, Teluk region near bivacque Dewan I, Meijer 4236 (10 January
1956 FRU) (L!).
This species is closest to R. sootepensis, based on vegetative characters.
However, due to lack of flowering material, the taxonomic status cannot be confirmed.
374