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A revision of the genus Gmelina (Lamiaceae)

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A revision of the genus Gmelina L. (Lamiaceae) is presented with a summary of its taxonomic history, keys, full descriptions, distribution maps, conservation assessments, ecological information and ethno-botanical notes. For the New Caledonia species no descriptions and distribution maps are given as they have been provided in an earlier publication. In this treatment, 31 species are recognised, 18 names are placed into synonymy for the first time, five are lectotypified, one is neotypified (G. racemosa (Lour.) Merr.) and seven names are excluded from the genus. Two varieties (G. palawensis var. palawensis and G. palawensis var. celebica) are raised to subspecies level. Four new species are described: G. australis de Kok, G. basifilum de Kok, G. hollrungii de Kok and G. peltata de Kok.
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A revision of the genus Gmelina (Lamiaceae)
Rogier de Kok
1
Summary. A revision of the genus Gmelina L. (Lamiaceae) is presented with a summary of its taxonomic history,
keys, full descriptions, distribution maps, conservation assessments, ecological information and ethno-botanical
notes. For the New Caledonia species no descriptions and distribution maps are given as they have been provided
in an earlier publication. In this treatment, 31 species are recognised, 18 names are placed into synonymy for the
rst time, ve are lectotypied, one is neotypied (G. racemosa (Lour.) Merr.) and seven names are excluded from
the genus. Two varieties (G. palawensis var. palawensis and G. palawensis var. celebica) are raised to subspecies level.
Four new species are described: G. australis de Kok, G. basilum de Kok, G. hollrungii de Kok and G. peltata de Kok.
Key Words. distribution, taxonomy.
Introduction
The genus Gmelina was rst described by Linnaeus in
1753 on the basis of one species Gmelina asiatica L. The
genus is named after Johann Georg Gmelin (1709
1755), who was Professor of Medicine, Botany and
Chemistry in Tübingen, Germany (1747 1755), and
Professor of Chemistry and Natural History in St.
Petersburg, Russia (1731 1747). He was elected to
accompany Vitus Berings Second Kamchatka Expedi-
tion (1731 1742), which formed the basis of his Flora
Sibirica (1747 1769).
The genus occurs naturally from India and South
China to North Australia and Fiji. Some species have
been introduced to many subtropical and tropical
countries, either as a timber tree (Gmelina arborea
Roxb. ex Sm.) (Harley et al. 2004) or as an ornamental
(G. asatica,G. elliptica Sm. and G. philippinensis Cham.).
The total number of species has been estimated from
about 33 (Munir 1984) to about 40 (Mabberley 2008):
in this revision 31 species are accepted, with two
subspecies. Most of the genus has been provisionally
revised by Moldenke (1984c,d,e,f,g,h) in a series of
articles. In this series the taxa were presented in
alphabetical order, but strangely he abruptly stopped
after G. tonkinensis Moldenke ( = G. elliptica) despite
the to be continuednote at the base of the page. In
addition, a substantial number of regional revisions
have been published. Most notable are Lam (1919) for
SE Asia and the Pacic, and Lam & Bakhuizen van
den Brink (1921) and Fletcher (1938a) for Thailand.
These revisions are now completely out of date. More
recent local revisions are available for Australia (Munir
1984), New Caledonia (Mabberley in Mabberley & de
Kok 2004), India (Rajendran & Daniel 2002), Sri Lanka
(Moldenke & Moldenke 1983), China (Shou-liang &
Gilbert 1994) and Vietnam (VũXuân Phuöng 2007).
The tree species of Borneo have been revised in the
Tree Flora of Sabah and Sarawak (Bramley et al. 2011); a
revision of the species of mainland Asia has been the
subject of a PhD study (Mohammad Harun-ur Rashid,
Trinity College Dublin) and those of Peninsular Malaysia
have been revised by Bramley & de Kok (in press).
Unlike most other genera in the Lamiaceae (de
Kok 2007,2008 &in press; de Kok et al. 2009), the
genus Gmelina seems to be particularly rich in species
and morphological variation in New Guinea and its
surrounding islands. The number of native species is
considerably fewer on the islands of the Sunda-Shelf.
A second, but less important centre of speciation is on
the SE Asian mainland, in particular Indo-China and
Southern China.There have been very few attempts to
subdivide Gmelina. Briquet (1895) divided the genus
into two sections, Microstromatae and Bracteosae, based
on the size, colour and venation of the oral
bracteoles, with G. philippinensis as the only species in
the section Bracteosae, and G. asiatica as the type of
section Microstromatae. This division is no longer used
and is not practical given the wide variation in
bracteole morphology within this genus.
Rumphiuss species
Rumphius described three species in his genus Tittius
in Herbarium amboinense vol. 3 (1750), two of which are
now thought to be part of Gmelina. The species Tittius
alba Rumph. and T. rubra Rumph. are identied in
this study as G. moluccana (Blume) Backer ex K.
Heyne. The third species of the genus, T. litoralis
Rumph. is placed by Merrill (1917) under the species
in Guettarda speciosa L. (Rubiaceae).
Accepted for publication April 2012. Published online 13 July 2012
1
Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, UK. e-mail: r.dekok@rbgkew.org.uk
KEW BULLETIN VOL. 67: 293 Y329 (2012) ISSN: 0075-5974 (print)
ISSN: 1874-933X (electronic)
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2012
Uses
The genus is most known for its use as a timber tree
(Gmelina arborea) (Dvorak 2004), but it is also extensively
used as a garden plant (mainly G. philippinensis, but also
G. asiatica and G. elliptica). The fast growing timber tree
G. arborea is increasingly being used in reforestation
projects. The cultivated species of the genus also have a
wide range of medicinal uses (Burkill 1966).
Morphology
Habit
Most species are tall trees, often with buttresses, but
some very common species (Gmelina asiatica,G.
elliptica and G. philippinensis) are qualied either as
(scrambling) shrubs or as small trees. One species is a
liana proper (G. uniora Stapf). The spines present in
some species (G. asiatica,G. elliptica and G. philippi-
nensis) are modied branches and would be better
dened as thorns (Beentje 2010). The leaves of the
seedlings are 3 7-lobed (Ng 1992).
Indumentum
There are two types of indumentum present in
Gmelina. The most common are simple multicellular
hairs that occur in all species and are common in the
family Lamiaceae. In some species these can form
dense mats on the underside of the leaves, on the
inoresence and/or calyces, and in those cases can
have taxonomic signicance. The second type is a form
of peltate scales that are either present or absent in a
species. When they are present there is great variety in
form and number. In some species the lower leaf surface
is completely covered with these scales resulting in a
whitish appearance (G. uniora and G. asiatica). In other
species the number of scales are fewer and they are less
conspicuous without a ×10 lens (G. phillipensis and G.
smithii Moldenke). The shape of the scales can vary
from those with entire margins to those with incised
margins that begin to resemble stellate hairs.
Glands
Glands on leaves. There is only one type of gland
present on the leaves of Gmelina. Discoid glands are
present as either solitary glands or in gland elds at
the base of the leaves or in rows along the main vein.
These glands are reminiscent of similar discoid glands
in other genera (Clerodendrum L. and Oxera Labill.) in
the Lamiaceae (Cantino 1990). In a small number of
species these glands are partly enveloped in a cup-like
structure at the base of the leaves. This cup-like
structure seems to be a modication of the base of
the leaf blade, where a number of glands, which form
a dense gland-eld, have been incorporated. These
cup-like glandular structures are often used as a useful
eld character for the genus as they are prominent in
the widespread species G. arborea. From notes on
herbarium specimens it is clear that ants are attracted
to the extraoral glands on the leaves and calyx.
Glands on calyx. Discoid glands similar to those found
on the leaves are present in some species on the outer
calyx surface. In his study of extraoral nectaries,
Burck (1891) noted that glands on the calyx of some
species (Gmelina asiatica and G. philippinensis [as G.
bracteata]) growing in the botanic gardens at Bogor in
Java, were visited by ants, which then patrolled the
inorescences. The presence of these glands on the
calyx is relatively common in the genus and is often a
good taxonomic character for species recognition.
Glands on bracteoles. Asmallgroupofspecies
(Gmelina australis de Kok, G. papuana Bakh., G.
ledermannii H. J. Lam and G. schlechteri H. J. Lam) from
New Guinea and northern Australia is marked by
having discoid glands on the bracteoles. These glands
are always present and are often very prominent,
giving the bracteoles the appearance of having eyes.
Inorescences
Almost all species have a terminal inorescence, very
rarely also an axillary one. Only one species (Gmelina
uniora) has consistently an axillary inorescence only.
In the latter case the inorescence also has a small
number of owers; while most species have many-
owered inorescences. The inorescence is of a very
common type in the family and is similar to the second
type inorescences in the genus Vitex L. (de Kok
2007). The inorescence is a panicle with the bracts
that are always leaf-like and persistent. The main
inorescence axes are seemingly indeterminate. The
main axis and major side branches (when present)
bear compact to lax small cymes.
Bracteoles
The variation in bracteoles in Gmelina is remarkable: they
can be small and narrow and falling off early in the
development of the inorescence (G. asiatica), while in
other species they can be big, colourful and remain on
the inoresence even during the fruiting period (G.
philippinensis). In his study of extraoral nectaries Burck
(1891) noted that at Bogor the corollas of G. philippinensis
[as G. bracteata]weresignicantly less broken into by
insects than those of a species with smaller bracteoles (G.
asiatica). His conclusion was that the many, big bracteoles
provided room for ants to make nests that, in so doing,
would be better able to protect the corollas.
Flowers
Calyx. All species have ve calyx lobes. However in many
species these lobes are hardly developed and sometimes
appear to be practically absent, although it is possible to
nd relictual or very small lobes on some calyces. The
outer surface of the calyx is either glabrous or velutinous
294 KEW BULLETIN VOL. 67(3)
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2012
when in ower, this character usually persisting when the
plant is in fruit, though sometimes the calyx will lose
some of its hairs during the fruiting period and then may
appear glabrous. In one species (Gmelina racemosa
(Lour.)Merr.)thecalyxisverybig(>5mmlong)and
completely envelopes the fruits, with the calyx lobes
growing into wing-like structures. In this case the calyx is
persistent with the fruit and calyx dispersed as one unit.
Corolla. There are two types of corolla shape in the
genus. The corollas of Gmelina asiatica,G. elliptica and
G. philippinensis are always a uniform bright yellow and
have four corolla lobes with a prominent anterior lip.
The second type has ve lobes, very rarely four (G.
arborea, which usually has corollas with four and ve
lobes together on one plant and G. delavayana Dop),
due to a split in the posterior lip; the two lips are
generally unequal, but the anterior one is much less
prominent than in the rst corolla type.
The colour is much more difcult to dene in these
species: some are uniformly white or pink to pale yellow
or purple, but more often they have several colours in
each ower, and there is a distinct difference between the
main corolla parts and the lip, or between the tube and
lobes, or between the inside and outside of the corolla.
The inside of the corolla is always glabrous, while
the outside surface can be glabrous to velutinous. The
corolla lobes can, in some species, be velutinous on
the margins. Most corollas seem to continue to grow
when they open, such that measurements of corolla
parts can vary greatly within species.
Fruits
Unlike some other closely related genera (Teismannio-
dendron and Premna) in the Lamiaceae (de Kok 2008
and in press; de Kok et al. 2009), fruit characters have
not traditionally been considered to have taxonomic
signicance in this genus. Gmelina fruits are two-celled,
each cell containing two ovules. It is unusual for both
ovules in a cell to develop.
Within the mature fruit there always is a cavity, the
origin of which is yet to be determined. It is present in
every mature fruit, does not contain any seeds and is
clearly not a gall. In young fruits this cavity can be
lled with pith. Such cavities can be very small, in
particular in young fruits, but in some species a cavity
can make up more than 50 % of the total volume of
the mature fruit. The shape varies with species and the
state of development of the fruit, as well as the
number of seeds developing within that particular
fruit. Usually the cavity is below the seeds, but it can
occupy the whole length of the fruit or sometimes it is
only distal to the seeds. Any possible function of this
cavity is unclear; there is no indication that species in
this genus are distributed via water, which would
explain the cavities as oating devices. A more likely
explanation could be that it is an efcient way of
increasing the size of the fruit, and species of this
genus are therefore able to mobilise a different
suite of dispersers from their close relatives. More
research on fresh material is needed to study the
development of these cavities and elucidate their
possible function.
Materials and Methods
This study is based on close examination of herbarium
specimens from the following herbaria: BM, BO, K, L,
LINN, P and SING (abbreviations following Thiers
2011); many (type) specimens were diagnosed using
images on the web.
In the following descriptions:
i) all measurements and colour descriptions are from
mature material;
ii) all collections cited have been seen by the authors
unless indicated otherwise;
iii) all measurements of the stigma are made from the
longer branch;
iv) cited specimens represent not only the typical form of
the species, but also more extreme forms (sometimes
formally described), and the intermediates between them;
v) all conservation status assessments follow the criteria
set by (IUCN 2001).
Gmelina
Gmelina L. (Linneaus 1753: 626); Schauer (1847: 678);
Briquet (1895: 173); Lam (1919: 214 228); Lam &
Bakhuizen (1921:6471); Fletcher (1938a: 422 424);
Moldenke (1984c: 308 342; 1984d: 424 442; 1984e:
460 499; 1984f:3254; 1984g: 102 126; 1984h: 154
182); Moldenke & Moldenke (1983: 388 401); Munir
(1984:91116); Shou-liang & Gilbert (1994:3234);
Rajendran & Daniel (2002: 153 171); Mabberley in
Mabberley & de Kok (2004:2234); Harley et al. (2004:
195 196); VũXuân Phuöng (2007:135146);
Bramley et al. (2011); Bramley & de Kok (in press).
Type: Gmelina asiatica L.
[Michelia Amman (1739: 218, t.18). Based on Michelia
spinosa Amman (1739: 218 219, t.18) = Gmelina
asiatica LorGmelina elliptica Sm.].
[Tittius Rumph. (Ruphius 1750: 38 39); Merril
(1917). Based on Tittius alba and Tittius rubra =
Gmelina moluccana (Blume) Backer ex K. Heyne;
Tittius litoralis =Guettarda speciosa L. (Rubiaceae)].
Cumbulu Rheede ex Adans. (Adanson 1763: 199). =
Gmelina arborea Roxb. ex Sm.
Ephielis Sol. ex Seem. (Seemann 1865: 258), nom.
illegit., non Ephielis Schreb. (Schreber 1789: 253).
Based on Ephielis simplicifolia Sol. ex Seem. =
Gmelina dalrympleana (F. Muell.) H. J. Lam.
Trees, shrubs or rarely lianas, sometimes spiny,
sometimes deciduous. Leaves simple, decussate, some-
295A REVISION OF THE GENUS GMELINA (LAMIACEAE)
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2012
times lower surface covered with hairs or with peltate
scales, often with discoid glands at base, sometimes
these glands concentrated in a cup-like structure.
Inorescence terminal, rarely axillary, dense to lax,
panicle with cymose side branches; bracts leaf-like,
decreasing in size towards apex; bracteoles present.
Calyx usually weakly 2-lipped, 5-lobed, sometimes
appearing fewer, persistent, usually accrescent, usually
with large discoid glands without. Corolla zygomor-
phic, 2-lipped, 4 5-lobed, blue or purple-violet, white,
yellow, brownish, mauve or purple pink, inside gla-
brous, tube widely funnel-shaped, greatly enlarged at
throat, posterior lip 1 2-lobed, anterior lip 3-lobed,
mid-lobe largest. Stamens 4 (5), didynamous,
inserted usually from halfway to at the apex of the
corolla tube, only once at the base (Gmelina basilum),
anthers dorsixed; anterior pair longer, always fertile,
included to exserted; posterior pair rarely sterile.
Pollen 3-colpate. Ovary imperfectly 2-celled, each cell
2-ovuled; only two ovules developing, the other two
are suppressed. Style liform, terminal; stigma-lobes 2,
unequal. Fruits drupaceous, with very eshy mesocarp
and tough endocarp, cavity present. Seeds 0 4
seeded, seeds without endosperm.
DISTRIBUTION. India and South China south to Aus-
tralia, New Caledonia and Fiji.
HABITAT. Growing in deciduous or evergreen forest or
anthropogenic vegetation; alt. 0 1500 (3000) m.
USES. Planted as ornamentals (Gmelina asiatica,G.
elliptica and G. philippinensis); used as light and
medium-weight timbers (G. arborea). G. arborea is used
increasingly in SE Asia as a pioneer species in
reforestation projects.
Key to Gmelina species
1. Shrubs or small trees, sometimes climbing, twigs usually with spines; corolla 4-lobed, bright yellow. . . .... 2
Trees or lianas, rarely shrubs, twigs always without spines; corolla 5 (very rarely 4)-lobed, white to yellowish,
bluish to purple or red . . ............................................................4
2. Lower leaves surface densely villous.............................................8. G. elliptica
Lower leaves surface glabrous or with only a few hairs on the veins . .............................3
3. Bracteoles linear to elliptic, < 6 mm wide, usually not persistent . . ......................2. G. asiatica
Bracteoles ovate-lanceolate, > 10 mm wide, usually persistent . ....................23. G. philippinensis
4. Lower leaf surface covered with whitish peltate scales, sometimes also with simple hairs. .............. 5
Lower leaf surface without peltate scales ................................................. 21
5. Lianas or small trees; inorescence few owered, axillary; corolla pale yellow to white; endemic to
Borneo ..................................................30. G. uniora
Trees, rarely shrubs; inorescence usually many owered, terminal, rarely axillary ones also present; corolla
bluish to reddish purple to white; widespread . ............................................6
6. Mature lower leaf surface velutinous, usually with yellow hairs .................................. 7
Mature lower leaf surface glabrous to sparsely hairy with white hairs on veins...................... 10
7. Leaves with two glands in a cup-like structure at the base; corolla (pale reddish) yellow . .....1. G. arborea
Leaves without glands or in gland elds at base or along midvein; corolla (cream-) white to pink........8
8. Fruit apex truncate, sparsely covered with peltate scales, grey when mature; endemic to Indo-China. . 12. G. lecomtei
Fruit apex round, glabrous, bluish-purple when mature; endemic to Australia or New Caledonia ........9
9. Inorescence open; endemic to Australia .....................................14. G. leichhardtii
Inorescence compact; endemic to New Caledonia . . . ........................19. G. neocaledonica
10. Calyx lobes > 4 mm long when in ower, endemic to China and Indo-China . . . ...................11
Calyx lobes < 2.5 mm long when in ower, widespread ...................................... 12
11. Leaf < 5 cm long; owering branches slender < 2 mm diam.; corolla 4-lobed . . . ........7. G. delavayana
Leaf > 5 cm long; owering branches > 2 mm diam.; corolla 5-lobed .................24. G. racemosa
12. Calyx glabrous when in ower; sometimes covered with peltate scales . . . .........................13
Calyx velutinous when in ower; sometimes covered with peltate scales . . . ....................... 16
13. Leaves with side veins > 9 . ............................................................14
Leaves with side veins < 7 . ........................................................... 15
14. Corolla pale pink to purplish brown; frurayishish white when mature; endemic to the Solomon Islands . . . 22.G.peltata
Corolla blue; fruit purple or mauve when mature; endemic to Fiji . . ...................31. G. vitiensis
15. Corolla mid lobe apex acuminate, outer surface glabrous to hairy; fruit apex covered with peltate scales;
endemic to (Indo-) China..................................................5. G. chinensis
Corolla mid lobe apex rounded, outer surface velutinous; fruit without peltate scales; endemic to the Moluccas
and New Guinea. . .......................................................15. G. lepidota
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16. Shrubs; corolla lobes almost isomorphous, tube elongated (tube > 24 mm long), not greatly expanding above
the calyx, velutinous; endemic to New Caledonia. . . ..............................29. G. tholicola
Trees; corolla lobes clearly different from one other, tube 8 14 mm long, greatly expanding above
the calyx ..................................................................................17
17. Leaves with two glands in a cup-like structure at the base . . ..........................1. G. arborea
Leaves without glands, or glands arranged in gland elds at base or along midvein . ...............18
18. Filaments inserted at base of ower tube........................................4. G. basilum
Filaments inserted at apex of ower tube . . .............................................19
19. Calyx lobes > 1 2.5 mm long when in ower; endemic to New Guinea . ................28. G. smithii
Calyx lobes < 1 mm long when in ower; endemic to Australia or the Solomon Islands .............20
20. Bracteoles clavoid to rounded, 2.5 5 mm long; owers blue to white with a purple lip; endemic to
Australia ............................................... 10. G. fasciculiora
Bracteoles lanceolate to spathulate, 5 30 mm long; owers (grey) white, sometimes with a purple base;
endemic to the Solomon Islands . . . .....................................25. G. salomonensis
21. Mature lower leaf surface hairy . . .......................................................22
Mature lower leaf surface glabrous or with hairs on the veins only .............................23
22. Bracteoles lanceolate, 4 10 ×15 mm; Moluccas to Solomon Islands . .............18. G. moluccana
Bracteoles rounded, 13 16 ×11 13 mm; occurring in Eastern New Guinea............27. G. sessilis
23. Corolla lobes almost isomorphous, tube not greatly expanding above the calyx; endemic to New
Caledonia................................................. 17. G. magnica
Corolla lobes greatly different from one other, tube greatly expanding above the calyx; occurring in
Australia, Solomon Islands and New Guinea ............................................24
24. Calyx glabrous when owering, lobes 0 1 mm long. ........................................25
Calyx velutinous when owering, lobes < 0.5 mm long . . ...................................26
25. Leaves with two cup-glands at base; mature fruit with a truncate apex; Australia and New
Guinea..........................................................6. G. dalrympleana
Leaves with several discoid glands at base; mature fruit with a rounded apex; endemic to Palau Islands and
occurring on the Caroline Islands ..........................................20. G. palawensis
26. Bracteoles with several disc-like glands present .............................................27
Bracteoles without disc-like glands . . . ..................................................30
27. Bracteoles petiolate, 12 23 ×59 mm, apex rounded to acute; owering calyx 4 4.5 ×67 mm; fruit apex
truncated; endemic to Australia, Northern territory . ...............................3. G. australis
Bracteoles sessile, fruit apex rounded; endemic to New Guinea ................................ 28
28. Inorescences dense without long side branches; bracteoles rounded on the main axis, 7 15 ×28 mm,
apex acute; owering calyx 3.5 5×2.5 4 mm; corolla lip 6 10 mm long . . .........21. G. papuana
Inorescences narrow with long side branches; bracteoles apex acuminate to long acuminate . . . ...... 29
29. Bracteoles on main inorescence axis rounded, apex long acuminate; owering calyx 3.5 5×3.5 4 mm;
corolla lip 7.5 10 mm long. . . ..........................................13. G. ledermannii
Bracteoles on main inorescence axis lanceolate, apex acuminate; owering calyx 2.5 3×2.5 3 mm;
corolla lip 5 8 mm long . . .............................................26. G. schlechteri
30. Leaves with cup-like glands at base; Moluccas, New Guinea, Australia and the Solomon Islands. 11. G. hollrungii
Leaves without cup-like lands, discoid glands sometimes present at base; endemic to New Caledonia . . . . . . . 31
31. Evergreen small tree or shrub, up to 3 m tall; endemic to north-west New Caledonia. . . ......9. G. evoluta
Deciduous tree, up to 20 m tall; endemic to Valée de Thy (New Caledonia) ........16. G. lignum-vitreum
1. Gmelina arborea Roxb. ex Sm. (in Rees 1810); Lam
(1919: 219 220); Greaves (1981:237258);
Moldenke (1984c: 337 342; 1984d: 424 442;
1984e: 460 469).
Gmelina arborea var. arborea Roxb. ex Moldenke &
Moldenke (1983: 390 394). Type: Rheede,. Hort.
Malab. 1 (1678) t. 41, selected here.
Gmelina oblongifolia Roxb. (Roxburgh 1814: 95, nom.
nud.;1832: 83); Moldenke (1984h: 156 157),
synon. nov. Type: Icon. Roxburgh t. 2306 (K!).
Premna tomentosa Miq. ex C. B. Clarke (1885: 581),
nom. illegit., non Willd. (Willdenow 1802).
Premna arborea Roth (1821: 287); Rajendran & Daniel
(2002: 156). Type: Ind. Orient.,Heyne s.n.
(holotype?).
Gmelina rheedei Hook. (Hooker 1848: t. 4395) as rheedii;
Lam (1919: 219). Type: Curtiss Bot. Mag. 4: t. 4395.
Gmelina arborea var. glaucescens C. B. Clarke (1885: 582);
Rajendran & Daniel (2002: 155). Type: [India]
Khasia Mts, 24 July 1850, Hooker s.n. (holotype K!).
297A REVISION OF THE GENUS GMELINA (LAMIACEAE)
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2012
Gmelina arborea var. canescens Haines (1910: 487);
Rajendran & Daniel (2002: 155). Type: [India,
Bengal] Santal Parganahs, 2 Feb. 1907, Haines
5769 (holotype CAL (n.v.); isotype K!).
Gmelina arborea forma dentata Moldenke (1961: 14);
Moldenke & Moldenke (1983: 390 394). Type:
India, Siwalik & Jaunsar division, Jhajra, 14 March
1922, AzizullahShah s.n. (holotype CAVA scan seen).
Trees 3 30 m high, DBH 10 200 (450) cm,
sometimes deciduous. Bark smooth; whitish to greyish-
reddish brown. Twigs densely tomentose, spines
absent. Leaves broadly ovate to reniform, 8 25 ×4.5
20 cm, apex rounded to acuminate, base cuneate to
(deeply) cordate, margins entire, sometimes slightly
lobed, papery; upper surface with hairs on veins, dark
green, shiny; lower surface glabrous to densely hairy,
covered with peltate scales, two cup-like glands clustered
around the base; veins 3 5 pairs, sometimes 3 veined
from base. Petiole terete, 3.5 13 cm, hairy. Inorescence
terminal or axillary, narrow; peduncle 10 30 cm.
Bracteoles lanceolate to linear or triangular, 8 15 ×1.3
2 mm long, deciduous. Calyx 2 7×45 mm, 5-lobed,
velutinous, not accrescent, green; lobes 0.8 1×0.7
2 mm, acute, outer surface with scattered discoid glands.
Corolla (pale reddish) yellow, outer surface hairy,
sparsely glandular, 4 5-lobed; anterior lip 3-lobed; mid-
lobe spathulate 15 20 ×15 25 mm, (reddish) yellow,
apex rounded; side lobes 10 15 ×912 mm long, apex
rounded, reddish brown; posterior lip 1 2-lobed, lobes 6
7×612 mm, apex rounded, reddish brown; tube 15
20 mm long. Stamens inserted at apex of tube, glabrous,
sometimes with a few glands, yellow; anterior pair 15
20 mm long, anthers 2 3 mm long, black; posterior pair,
fertile; 12 14 mm long, anthers c. 1 mm long. Ovary
glabrous; style 15 23 mm long, stigma c. 1 mm long. Fruit
12 20 ×10 15 mm, ellipsoid to obovoid-ellipsoid,
glabrous, apex rounded, (dark) purple to yellow when
mature, recorded as having a bitter sweet taste.
DISTRIBUTION. Natural distribution: from Mirpur in
India and Jamu in Pakistan east to South Yunnan in
China and northern Vietnam, south to Northern
Thailand and Sri Lanka (Map 1). It is introduced in
most tropical countries as a timber-, or shade- tree as
well as an ornamental.
SELECTED SPECIMENS EXAMINED. BANGLADESH. [East
Pakistan] Chittagong, Chuak R.F., 27 Feb. 1966,
Majumder & Islam 38A (K). INDIA. Maharashtra State:
Khandala [Khaudala] saddle, 21 April 1943, Santapau
1932 (K); Tamil Nadu State: Mont Nilghiri, Thomson
s.n. (K); Hort. Cal., 11 Sept. 1819, Wallich 1832 (K);
Meghalaya State: Khasia Mts, 24 July 1850, Hooker s.n.
(holotype K); Jharkhand State: [Bengal] Santal Parga-
nahs [Santhal Pargana], 2 Feb. 1907, Haines 5769
(isotype K); Ultarakhand State: Siwalik & Jaunsar
division, Jhajra, 14 March 1922, AzizullahShah s.n.
(holotype CAVA). LAOS. Khammouan, Nakai Talang,
1 March 2007, Vannachak BT 913 (L). MYANMAR.
Pyiamaua, March 1913, Gamble s.n. (K). SRI LANKA,
Nalanda, 19 May 1947, Worthington 2787 (K). THAI-
LAND. Rachasima, Pu Kio, 25 Feb. 1931, Kerr 20271 (K);
Huai Krasa, 90 km S of Tak, 18 March 1968, Hansen &
Smitinand 12952 (K).
HABITAT. Growing in deciduous or evergreen forest or
anthropogenic vegetation; alt. 0 1500 m. It is a
relatively short-lived tree that colonises openings in
the forest, where the ground has been disturbed.
Usually it is not common and it does not grow on
poorly drained soils. Soil (sandy) loam to clay.
CONSERVATION STATUS. Least Concern.
PHENOLOGY. In the more seasonal areas it is reported
to ower between April and May; fruiting occurs
between May and July; in wet-tropical areas owering
and fruiting throughout the year. The seeds fail to
germinate in the full shade, but germinate well in the
open (Mohamad & Ng 1982).
USES. Planted widely as an ornamental, shade and/or
timber tree. The wood is used for various purposes
including construction, furniture, and musical instru-
ments. The tree also has a wide range of medicinal
uses (Burkill 1966).
VERNACULAR NAMES. India, Bengal: Ato Kasmar;
Bombay presidency: Sioni (Kanarese language); Sri
Lanka: Et demata (Shingalese Language) or Kumil
(Tamil Language); Myanmar: Yamanay;China:yun
nan shi zi;Laos:Saw or Mai so;Thailand:The rong
(Karieng language); Peninsular Malaysia: Bulang
(Malay Language) or Jemane;Sabah:Jemane (Malay
Language) or Jamaná (Dusun Language); Fiji:
Yemane.
NOTES. Chromosome 2n = 36 (Mangenot & Mangenot
1962: 411). The type material of Gmelina oblongifolia
Roxb. falls within the total morphological variation of
G. arborea ex. Sm. and is therefore placed into its
synonomy.
2. Gmelina asiatica L. (Linnaeus 1753: 626); Lam
(1919: 221 223); Lam & Bakhuizen (1921:6970).
Gmelina asiatica var. typica Bakh. in Lam & Bakhuizen
(1921:6970), nom. illegit., super. Type: Herb.
Linn. 7802 (holotype LINN!).
Gmelina parviora Roxb. (Roxburgh 1802: 32 33, t.
162); Gmelina asiatica f. parviora (Roxb.) Moldenke
(1984a: 42). Type: Roxburgh (1799 1805), Pl.
Coast. Coromand., Pl. 162.
Gmelina parviora Pers. (Persoon 1806: 142). Based on
Roxburgh (1799 1805 Pl. Coast. Coromand., Pl.
162, t 32 and Burman (1768)Flora Indica, p. 332.
Gmelina asiatica Wall. (Wallich 1828 n. 1818), nom.
nud., nom. illeg., non Gmelina asiatica L. Based on:
[India] Botanical Garden, 22 Nov. 1814, [Wallich]
1818 (K-W!).
298 KEW BULLETIN VOL. 67(3)
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2012
Gmelina attenuata H. R. Fletcher (1938b: 203 204),
synon. nov. Type: Thailand, Payap, Chiengmai, 4
July 1922, Kerr 6224 (holotype E scan seen).
Gmelina paniculata H. R. Fletcher (1938b: 204); Mol-
denke (1984h: 161), synon. nov. Type: Thailand,
Prachinburi, Krabin, Aranya, 18 Oct. 1928, Put 2086
(holotype E scan seen; isotype K!)
Gmelina asiatica forma lobata Moldenke (1975: 47),
synon. nov. Type: Sri Lanka, Wilpattu National
Park, Smithsonian Camp, Marai Villu, 30 June
1969, Wirawan, Cooray & Balakrishnan 899 (holo-
type NY scan seen; isotypes K!, L!).
Shrubs, sometimes climbing, 2 4 m high, rarely
deciduous. Twigs minutely hairy when young; spines
usually present, up to 25 mm long. Leaves ovate to
elliptic or deltoid, 0.5 5(13) ×0.5 3.3 (6) cm,
apex rounded to acute, papery, base rounded to
cuneate, margins entire to 5-lobed; upper surface
glabrous, or with a few hairs on veins, dark green,
shiny; lower surface glabrous or with a few hairs on
veins, whitish, covered with peltate scales; veins 3 4
pairs, sometimes 3-veined from base, few discoid
glands present. Petiole 5 30 mm, glabrous or with a
few hairs, peltate, scales absent to many. Inorescence
terminal, sometimes appearing axillary on short side
shoots, 1.5 10 cm long, pendulous or erect, sparsely
hairy, covered with white peltate scales; bracteoles 8
10 ×3.5 6 mm, linear to elliptic, cuspidate,
caducous. Calyx 4 5-lobed, few hairs present, discoid
glands sometimes present; owering calyx 4 6×3.5
4 mm; lobes acute, 0 1 mm long; fruiting calyx 7
8 mm diam., erect to patent. Corolla 4-lobed, covered
with yellow hairs, sparsely glandular; anterior lip 3-
lobed, mid-lobe oblong, 12 35 ×10 15 mm long,
apex rounded, patent, cuspidate, margins sometimes
reexed; side lobes 4 10 ×58 mm, sometimes
oblique, apex acute, erect to reexed; posterior lip 1-
lobed, 1.5 3.5 ×16 mm, oblong, apex acute to
rounded; tube 10 23 mm long. Stamens inserted at
apex of tube, glabrous, sometimes with a few glands;
anterior pair 10 20 mm long, anthers 2 3 mm long;
posterior pair fertile, 2 (3); 8 11 mm long, anthers c.
1 mm long. Ovary glabrouswith a few hairs at apex; style
20 32 mm long. Fruit (dried) 13 30 ×830 mm,
clavoid, glabrous, apex rounded, yellow when mature.
DISTRIBUTION. India and Sri Lanka east to Northern
Thailand and North Vietnam (Map 2). Introduced
and sometimes cultivated in gardens in Peninsular
Malaysia (Kochummen 1978) and/or naturalised
along roadsides. Cultivated throughout the tropics.
SELECTED SPECIMENS EXAMINED. INDIA. Bombay Presi-
dency [Maharashtra State]: North Konkan, Karjat
region, 10 Feb. 1949, Fernandes 88 (K); Tamil Nadu
State: Attur distr., Salem, 30 March 1979, Perumal
22539 (K). MYANMAR. Upper Burma: 1888, Collet 554
(K). SRI LANKA. Yala: 13 June 1965, Worthington 7103
(K); Wilpattu National Park: Smithsonian Camp, Marai
Villu, 30 June 1969, Wirawan, Cooray & Balakrishnan
899 (holotype NY, isotypes K, L). THAILAND. Lampoo
[Lamphun Province]: Mae Tah Distr., Doi Kuhn Dahn
Nat. Park, summit of Doi Hoa Chang, 3 June 1994,
Maxwell 94676 (L); Prachinburi Province: Krabin,
Aranya, 18 Oct. 1928, Put 2086 (holotype E, isotype K);
Chiang mai Province: Payap, Chiengmai [Chiang
Map 1. Probable natural distribution of Gmelina arborea (); distribution of Gmelina moluccana (); Gmelina vitiensis ().
299A REVISION OF THE GENUS GMELINA (LAMIACEAE)
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2012
Mai], 4 July 1922, Kerr 6224 (holotype E). VIETNAM.
Cochinchina: 1862 1866, Thorel 60 (K); Tonkin: Bac
Ninh, May 1891, Balansa 4972 (K).
HABITAT. In deciduous or evergreen forest, secondary
and coastal vegetations. Soil sandy clay or a coarse
gritty substrate often in poor soils, sometimes over
granite or sandstone; alt. 0 1300 m.
CONSERVATION STATUS. Least Concern.
PHENOLOGY. Flowering and fruiting throughout the year.
VERNACULAR NAMES. Sri Lanka: Demata (Singhalese
language), Kumil (Tamil language); Peninsular Malay-
sia: Bulang or Bulongan (Malay language). Cambodia:
Köncaang; Laos: Phoung nou. China: Ya zhou shi zi.
Philippines: Banganga (Cebuano language). Indone-
sia, Wareng (Javanese language), West Timor: Hau bako
(Dawan language); Alor: Lombaul.
USES. The species has a wide range of medicinal uses
(Burkill 1966).
NOTES. The type material of Gmelina asiatica forma
lobata Moldenke, G. paniculata H. R. Fletcher and G.
attenuata H. R. Fletcher fall within the total morpho-
logical variation of G. asiatica L. and is therefore
placed into its synonomy.
3. Gmelina australis de Kok sp. nov. aG. schlechteri
bracteolis petiolatis maioribusque atque ad apicem
rotundatis acutisve, calyce ad anthesin maiore et
fructu ad apicem truncato differt. Typus: Australia,
Northern Territory, c. 5 km E Nourlangie Rock, 10
Nov. 1972, Martensz 304 (holotypus K!; isotypi CANB,
DNA, NT).
http://www.ipni.org/urn:lsid:ipni.org:names:77120319-1
[Munir (1984: 106 109 as Gmelina schlechteri p.p.,
Australian material only].
Tree 12 16 m high. Bark smooth to ssured, grey. Twigs
densely hairy when young, interpetiolar ridge present,
spines absent. Leaves elliptic to obovate, 18 27 ×9
16 cm, apex rounded to acuminate, base rounded to
cuneate, margin entire, sometimes slightly lobed; upper
surface glabrous, green; lower surface glabrous with a
few simple hairs on veins, peltate scales absent, pale
green; two cup-like glands at the base; veins 5 7pairs,
sometimes 3-veined from base. Petiole 4 6.5 cm long,
half-terete, a few simple hairs present. Inorescence
terminal, open, up to 20 cm long, with long side
branches, velutinous; bracteoles lanceolate, 12 23 ×5
9 mm, apex rounded to acute, petiolate, with several
disc-like glands present. Calyx 4 4.5 ×67mm,5-
lobed, sometimes appearing fewer, velutinous with
sometimes a glabrous margin, outer surface with discoid
glands; lobes up to 0.5 mm long, triangular; fruiting calyx
c. 7 mm diam., erect. Corolla 5-lobed, outer surface
hairy, sparsely glandular on lobes, pale lilac; anterior lip
3-lobed; mid-lobe oblong, 5 9×45 mm, apex round,
lilac, with two yellow marks; side lobes 4.5 5×4.5
5 mm long, apex round; posterior lip, 2-lobed, 4.5 5×
67 mm, oblique, apex round; tube 8 11 mm long.
Stamens inserted at apex of tube, glabrous, glands
present; anterior pair 8 15 mm long, anthers c. 2 mm
long; posterior pair, fertile, 6 10 mm long, anthers c.
1.5 mm long. Ovary glabrous with a few hairs at apex.
Map 2. Probable natural distribution of Gmelina asiatica (); distribution of Gmelina australis (); Gmelina basilum (); Gmelina
hollrungii (); and Gmelina leichhardtii ().
300 KEW BULLETIN VOL. 67(3)
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2012
Style 15 20mmlong,stigmac.2mmlong.Fruit (dried)
14 17 ×813mm,clavate,apextruncated,glabrous,
glossy, reddish purplewhenmature.Fig.1.
DISTRIBUTION. Australia: Northern Territory restricted
to the tropical forest of the Darwin region. Map 2.
SPECIMENS EXAMINED. AUSTRALIA. Northern Territory:
c. 5 km E Nourlangie Rock, 10 Nov. 1972, Martensz 304
(holotype K); sandstone tableland between Gerowie
Creek and Mary R., 4 Oct. 1946, Blake 17169 (K).
HABITAT. Along the fringes of rainforests, sometimes
on black sandy soil; alt. c. 300 m.
CONSERVATION STATUS. Least Concern.
PHENOLOGY. Flowering Oct. Nov.; fruiting Nov.
NOTES. Gmelina australis differences from G. schlechteri
in having bracteoles which are petiolate, bigger,
rounded to acute at the apex; a bigger calyx when in
ower and a fruit with a truncated apex.
4. Gmelina basilum de Kok sp. nov. aGmelina smithii
ramulis lateralibus unorescentiae longis (nec brevi-
bus), lamentis ad basin (nec ad apicem) tubi corollae
insertis et ovario ad apicem piloso (nec omnino
glabro) differt. Typus: [Indonesia] Netherlands New
Guinea, Geelvinkbaai, Japen Island, Soemberbaba, 2
July 1961, Koster BW 11115 (holotypus K!; isotypi L!,
SING!).
http://www.ipni.org/urn:lsid:ipni.org:names:77120320-1
Trees 7 10 m high, DBH 7 10 cm. Bark whitish brown
to grey. Wood (medium) hard, white to cream. Twigs
hairy, spines absent. Leaves elliptic to lanceolate, 13 17
×58.5 cm, apex acute, base cuneate to rounded,
margins entire; upper surface glabrous, dark green;
lower surface glabrous with simple hairs on veins of
young leaves, upper surface dark green; lower surface
glaucous, covered with peltate scales, discoid glands
absent; veins 7 10 pairs, sometimes 3-veined from base.
Petiole half-terete, channelled, 3 5cm,glabrousto
sparsely hairy. Inorescence terminal, 12 25 cm long, side
branches up to 9 cm long, densely hairy, scales present;
bracteoles ovate-lanceolate, 4 5×22.5 mm, sessile,
apex acute, caducous. Calyx 3 4.5 ×34.5 mm, 5-
lobed, velutinous when in ower, with a glabrous
margin, glabrous when in fruit, surface covered with
peltate scales, outer surface without discoid glands; lobes
0.8 1×11.2 mm; apex acute to rounded; fruiting
calyx 5 12 mm diam., glabrous, patent. Corolla 5-lobed,
purple to white, outer surface velutinous, inside veluti-
nous on lobes, glabrous in tube; anterior lip 3-lobed;
mid-lobe 4 5×44.5 mm, oblong, apex rounded; side
lobes 3 4×34 mm, apex rounded; posterior lip 2-
lobed, 3 4×3.5 4 mm, apex rounded; tube 2 5mm
long. Stamens inserted at base of tube, hairy at base;
anterior pair 7 8mmlong,anthersc.1.5mmlong;
posterior pair, fertile, 6 7mmlong,anthersc.1mm
long; ovary c. 2 mm diam., glabrous with tuft of hairs at
apex. Style 6.5 7mmlong,stigma11.5 mm long.
Fruit (dried) 12 25 ×13 30 mm, globose, apex
rounded, glabrous, purple green to reddish purple when
mature. Fig. 2.
DISTRIBUTION. The islands of New Guinea and New
Britain. Map 2.
ADDITIONAL SPECIMENS EXAMINED. INDONESIA. New
Guinea: Geelvinkbaai, Japen Island, Soemberbaba, 2
July 1961, Koster BW 11115 (holotype K, isotypes L,
SING). PAPUA NEW GUINEA. Madang Province: Gogol
R., 6 May 1970, Katik 46669 (K, SING); West New
Britain Province: near Linga-linga, 29 May 1973, Henty
& Lelean 49499 (K); Central Province: Sogeri, 6 May
1970, N.G.P. 4167 (SING).
HABITAT. Growing in primary and secondary forests,
sometimes along rivers; alt. 0 500 m.
CONSERVATION STATUS. This is a rare species with a
very wide distribution in undisturbed forest, though it
can also survive in secondary forests. Therefore,
despite the limited number of collections known, I
would provisionally propose a conservation assessment
of Least Concern.
PHENOLOGY. Flowering July; fruiting May July.
NOTES. The specic epithet refers to the position at
which the stamens are inserted, namely the base of the
corolla tube, a character unique in the genus.
5. Gmelina chinensis Benth. (Bentham 1861: 272);
Oliver (1889: t. 1974); Moldenke (1984f:3335);
Shou-liang & Gilbert (1994: 33). Type: [China] Hong
Kong, 1853 1856, Wright 490 (holotype K!; isotype NY
scan seen, US scan seen).
Gmelina balansae Dop (1914: 322 323). Type:
[Vietnam] Indo-Chine, Annam, a Hué, Harmand
s.n. (syntype HM (n.v.); isosyntypes K!, P!); Laos,
Phron-thane, Spire 233 (syntype B scan seen);
[Vietnam] Tonkin, Bon 5413 (syntype ?); [Vietnam]
Tonkin, Mont Bavi, Vallé du Lankok, 28 June 1887,
Balansa 3806 (syntype HM (n.v.); isosyntype K!, L!,
NY scan seen, P! ×2 sheets).
Gmelina lecomtei var. annamitica Dop (1933: 896); Vũ
(2007: 143), synon. nov. Type: [Vietnam] Indo-
Chine, Annam, a Hué, Harmand s.n. (holotype HM
(n.v.); isotypes K!, P!).
Gmelina speciosa Moldenke (1940: 418 419), synon.
nov. Type: [Vietnam] Tonkin, Mont Bavi, Vallé du
Lankok, 28 June 1887, Balansa 3806 (holotype K!;
isotypes HM (n.v.), NY scan seen, P!).
Tree or shrubs 1.5 20mhigh,DBH1530 cm. Bark grey
to grey-brown, nely ssured, wood yellowish. Twigs hairy
when young, spines absent. Leaves ovate to elliptic, 5 18 ×
311 cm, apex acute, base cuneate, margins entire;
301A REVISION OF THE GENUS GMELINA (LAMIACEAE)
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2012
papery; upper surface glabrous; lower surface glabrous to
sparsely hairy, covered with peltate scales, a few discoid
glands clustered around the base and veins, bluish green;
veins 3 5 pairs, sometimes 3-veined from base,
sometimes these prominent. Petiole terete, slightly chan-
nelled, 2 5.5 cm, glabrous. Inorescence terminal, 9
Fig. 1. Gmelina australis.Ahabit; Bglands on bracteole; Cower side view; Dcalyx; Ecorolla; Fstyle, stigma and ovary; Gfruit
(dried). From Blake 17169, sheets 1 & 2. DRAWN BY JULIET BEENTJE.
302 KEW BULLETIN VOL. 67(3)
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2012
20 cm, velutinous; bracteoles rounded to lanceolate, 8
15 ×46.5 mm, sparsely hairy. Calyx 6 8×88.5 mm,
5-lobed, sometimes appearing fewer, glabrous to sparsely
hairy, covered with peltate scales, outer surface with few
discoid glands, grey-brown; lobes triangular, 0.5 1×2
4 mm, apex acute; fruiting calyx c. 14 cm diam. Corolla 5-
lobed, yellow outside, white or pink-purple, inside, outer
surface glabrous to sparsely hairy, often covered with
peltate scales; inside glabrous; anterior lip 3-lobed; mid-
lobe 11 17 ×910 mm, apex acuminate, yellow; side
Fig. 2. Gmelina basilum.Ahabit; Bdetail leaf underside, scales; Cower side view; Dcalyx; Ecorolla; Fstamen; Gstyle and ovary;
Hfruit (dried). From Koster BW 11115. DRAWN BY JULIET BEENTJE.
303A REVISION OF THE GENUS GMELINA (LAMIACEAE)
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2012
lobes 9 16 ×810 mm long, apex rounded; posterior
lip, 2-lobed, 7 12 ×710 mm, apex rounded to
truncate; tube 17 22 mm long. Stamens inserted at
apex of tube; anterior pair 13 20 mm long,
anthers 2 2.5 mm long; posterior pair, fertile, 15
17 mm long, anthers 2 3 mm long. Ovary velutinous at
apex. Style 25 27 mm long, stigma c. 1 mm long. Fruit
(dried) 10 30 ×615 mm, clavate, apex truncated,
glabrous, apex covered with peltate scales, few discoid
glands present.
DISTRIBUTION. South China, Laos and Vietnam south to
Da Nang (see Map 3).
SPECIMENS EXAMINED. CHINA. Hong Kong: 1853 1856,
Wright 490 (holotype K; isotype NY, US). Laulao Island,
10 Aug. 1888, Native collector 68/88 (K). LAOS. Kham-
mouan: Nakai, 23 May 2006, Nanthavong BT 501 (L);
Nakai distr., Ban Sop Phone, 8 Oct. 2006, Nanthavong
BT 666 (L). VIETNAM. Mt Bani: 25 km from Tourane,
May July 1927, Clemens 3980 (K, L); Quang-nam: Da
Nang, 1923, Poilane 7533 (K); Tonkin: Mont Bavi, Vallé
du Lankok, 28 June 1887, Balansa 3806 (isosyntypes K,
L, NY, P); Mont Bavi, Vallé du Lankok, 28 June 1887,
Balansa 3806 (holotypes HM, K, L, P, NY). Annam, a
Hué, Harmand s.n. (isosyntypes K, P); Laos: Phron-
thane, Spire 233 (syntype B).
HABITAT. Growing in mixed forest on mountain slopes;
alt. 500 1200 m altitude. Sometimes on poor laterite soils.
CONSERVATION STATUS. Least Concern.
PHENOLOGY. Flowering April July; fruiting June Oct.
VERNACULAR NAMES. China: Shi zi.
NOTES. The type material of Gmelina speciosa Moldenke
and G. lecomtei var. annamitica Dop fall within the total
morphological variation of G. chinensis Benth. and is
therefore placed into its synonomy.
6. Gmelina dalrympleana (F. Muell.) H. J. Lam (1919:
223), Moldenke (1984f:3539); Munir (1984: 109
114); Vitex dalrympleana F. Muell. (Mueller 1864: 128).
Type: [Australia, Queensland] Rockingham Bay,
Dallachy s.n. (lectotype MEL 583504 scan seen;
isolectotypes K! ×2 sheets), selected by Munir
(1984: 109).
Ephielis simplicifolia Sol. ex Seem. (Seemann 1865: 258);
Munir (1984: 110). Vitex macrophylla R. Br. (Brown
1810:512),Gmelina macrophylla (R. Br.) Benth.
(Bentham 1870: 65), nom. illegit., non Gmelina
macrophylla Wall. ex Schauer (1847: 680)]; Munir
(1984: 110). Type: Australia, Queensland, Cape
Grafton, 1768 1771, Banks & Solander s.n.
(holotype BM!).
Shrub or tree, 1 40 m high, DBH 5 70 cm. Bark
ssured or aky, greyish to grey-brown. Wood dark
straw coloured with yellow streaks; twigs glabrous,
spines absent. Leaves broadly ovate to ovate-oblong or
elliptic-obovate, 6 37 ×523 cm, margins entire,
Map 3. Distribution of Gmelina chinensis (); Gmelina fasciculiora (); Gmelina ledermannii (); Gmelina papuana (); Gmelina
schlechteri ().
304 KEW BULLETIN VOL. 67(3)
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2012
apex obtuse to obtuse acuminate, base cordate to
cuneate; upper surface glabrous, dull or shiny (dark-
olive) green; lower surface glabrous, peltate scales
absent, with glands in a cup like structure at base, pale
or greyish green; veins 4 8 pairs. Petiole 5 40 mm
long, half-terete, slightly channelled, glabrous to hairy.
Inorescence terminal, 12 30 cm long, glabrous to
hairy; bracteoles linear, 2 3×11.5 mm, apex
acute, caducous. Calyx 5-lobed, sometimes appearing
fewer, glabrous, with black discoid glands; owering
calyx 4 6×33.5 mm; lobes 0.5 1×1.5 4 mm,
apex round; fruiting calyx 6 7 mm diam., erect to
patent. Corolla 5-lobed, velutinous, white, pink-yellow
to pink or (pale) purple; anterior lip 3-lobed, mid-lobe
oblong, 7 10 ×56 mm long, apex rounded, dark
pink to mauve with yellow spot at base; side lobes
oblong-ovate, 5 9×3.5 6 mm, apex rounded;
posterior lip 2-lobed, oblong, 5 9×45 mm, apex
rounded; tube 10 22 mm long. Stamens inserted at
apex of tube, glabrous, sometimes with a few glands,
white; anterior pair 9 13 mm long, anthers c. 2 mm
long; posterior pair, fertile, 7 9 mm long, anthers
1.5 2mmlong.Ovary11.5 mm diam.,
glabrous, style 12 20 mm long, stigma c. 1.5 mm long.
Fruit obovoid, 8 20 ×525 mm, apex truncated,
glabrous, pinkish to red when mature.
DISTRIBUTION. Australia: North Queensland and the
island of New Guinea. Map 4.
SELECTED SPECIMENS EXAMINED. AUSTRALIA. Queens-
land: Rockingham Bay, Dallachy s.n. (lectotype MEL;
isolectotypes K); Cape Grafton, 1768 1771, Banks
& Solander s.n. (holotype BM); Cairns: Woree, Bruce
Highway, 10 March 1988, Gray 4766 (K). INDONESIA.
Merauke area: Pr. Kurik, South Polder, 6 June 1961,
Hoogerwerf 118 (L.); Koerik Camp, c. 15 km NE of
Koembe village, 7 Sept. 1954, van Royen 4891 (L);
North West Guinea: Djalan Kp. Keliki, 8 Aug. 1941,
Anta (exp. Wentholt) 250 (L); Birds Head Peninsula:
surroundings of Ayawasi, 16 March 1996, Ridsdale
2311 (L); surroundings of Ayawasi, 17 Sept. 1996,
Wanda Ava 4800 (L); Nova Guinea Neerlandica
Meridionalis: 29 Aug. 1907, Branderhorst 23 (K, L).
PAPUA NEW GUINEA. Western Province: Daru Island,
21 Sept. 1972, Streimann & Lelean 18459 (K, L,
SING); Oriemo R., Jan. 1959, White & Gray 10374
(K); Wassi Kussa R., Tarara, Dec. 1936, Brass 8539
(K, L).
HABITAT. Growing as a tree in rainforest to open forest,
dunes or swamps (in Papua New Guinea sometimes
associated with Melaleuca or Eucalyptus-Banksia wood-
land), sometimes a shrub in savanna woodlands; alt 0
450 m. Soil sometimes sandy.
CONSERVATION STATUS. Least Concern.
PHENOLOGY. Flowering Sept. April; fruiting Feb. Oct.
VERNACULAR NAMES. Australia: DalrymplesWhite
Beech or Queensland Beech or Long-leaved Gmelina.
Indonesia: Ara afa (Ayawasi).
Map 4. Distribution of Gmelina dalrympleana (); Gmelina delavayana (); Gmelina elliptica () and Gmelina sessilis ().
305A REVISION OF THE GENUS GMELINA (LAMIACEAE)
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2012
USES. The wood is used in Australia for oorboards
and planks.
7. Gmelina delavayana Dop (1914: 321); Moldenke
(1984f:4143); Shou-liang & Gilbert (1994: 33).
Type: China, Yunnan, Ta-pin-tze, Delavay 3595 (lecto-
type NY scan seen, designated here; isolectotype P
scan seen).
Gmelina montana W. W. Sm. (Smith 1916: 107 108);
Shou-liang & Gilbert (1994: 33). Type: [China]
Yunnan, western ank of the Tali Range, Aug.
1913, Forrest 11662 (holotype E (n.v.); isotype K!).
Shrub 0.3 3 m high; twigs hairy when young, spines
absent. Leaves ovate to elliptic, 1.5 5×1.5 3.5 cm,
apex acute, base cuneate, margins entire; sometimes
slightly lobed, papery; upper surface glabrous; lower
surface glabrous, sometimes with hairs on veins, covered
with peltate scales; veins 3 4 pairs, sometimes 3-veined
from base. Petiole 3 8 mm, half-terete, channelled,
glabrous or with a few hairs, peltate scales absent
to many. Inorescence terminal, lax, 8 12 cm long,
sparsely hairy, covered with peltate scales; bracteoles
elliptic to lanceolate, 6 50 ×135 mm, apex acute.
Calyx 5-lobed, discoid gland present, 6 11 ×67mm;
lobes 2.5 3.5 mm long, apex acute, not accrescent,
erect. Corolla 4-lobed, sparsely covered with hairs, bluish
to reddish purple, base yellow; anterior lip 3-lobed, mid-
lobe rounded to spathulate, 5 15 ×10 11 mm long,
apex acute to round, patent; side lobes 5.5 15 ×8
10 mm, rounded to spathulate, sometimes oblique, apex
acute to rounded, erect to reexed; posterior lip entire
to slightly lobed, 8 18 ×10 13 mm, oblong, apex
rounded; tube 4 5 mm long, infundibulate. Stamens
inserted at apex of tube, glabrous, sometimes with a
few glands; anterior pair 14 16 mm long, anthers
2.5 mm long; posterior pair, fertile, 9.5 13 mm long,
anthers c. 1 mm long. Ovary glabrous, sparsely glandu-
lar; style 17 20 mmlong, glabrous. Fruit (dried) 6 7×
67 mm, globose, apex round, glabrous, black when
mature.
DISTRIBUTION. China: endemic to South West Sichuan
and Yunnan. Map 4.
SPECIMENS EXAMINED. CHINA. Yunnan: June 1908,
dAlleizette s.n. (L); 1917 1919, Forrest 15620 (K);
western ank of the Tali Range, Aug. 1913, Forrest
11662 (isotype K); Ta-pin-tze, 17 Aug. 1888, Delavay
s.n. (P); Ta-pin-tze, Delavay 3595 (lectotype NY,
isolectotype P); Ta-pin-tze, 1888, Delavay s.n. (P ×4);
13 June 1887, Delavay s.n. (P); 10 July 1885, Delavay s.n.
(P); July 1746, Schneider 1746 (K); western ank of the
Tali Range, Aug. 1913, Forrest 11662 (isotype K); NW
Yunnan, N of Yung-peh, Forrest 22081 (K); W Yunnan,
1933, McLaren 250 (K). SW Szechuan: Sept. 1922,
Forrest 22499 (K);
HABITAT. In thickets and along streams on mountain
slopes; alt. 1500 3000 m.
CONSERVATION STATUS. Least Concern.
PHENOLOGY. Flowering May July; fruiting July Sept.
VERNACULAR NAME. China: Xiao ye shi zi.
NOTES. Dop (1914: 321) cited a number of specimens
in his original description of Gmelina delavayana:
China, Yunnan, Ta-pin-tze, Delavay 170; China,
Yunnan, Ta- pin-tze, Delavay 3595; China, Yunnan,
Pint chouan, Ducloux 4698; China, Yunnan, Pint
chouan, Ducloux 4707. From these, I select Delavay
3595 from NY as the lectotype as the specimen has a
letter attached from Dop stating that this is G.
delavayana.
This species seems to be morphologically close to
Gmelina asiatica,G. elliptica and G. philippinensis.It
shares with them their typical habit (a shrub) and the
number of corolla lobes (four); it can be distinguished
from them by the absence of spines, as well as by the
different corolla shape and colour.
8. Gmelina elliptica Sm. in Rees (1810); Moldenke
(1984f:4354); Munir (1984:9598); Rajendran &
Daniel (2002: 161 166 as G. asiatica); Zhengyi &
Raven (1988:84asG. asiatica). Type: East Indies, Anon.
in Herb. Linn. 7801 (holotype LINN!).
[Gmelina vestita Wall. (Wallich 1828: 50), nom. nud.,
synon. nov. Based on: [Myanmar] Kyouk Talong,
Irrawaddy R., 28 Sept. 1826, Wallich 1820 (K-W!).]
Gmelina villosa Roxb. (Roxburgh 1814: 46, nom. nud.;
1832: 86); Lam (1919: 217 219); Gmelina asiatica
var. villosa (Roxb.) Bakh. in Lam & Bakhuizen
(1921: 70). Lectotype: Rumphius (1750:t.39)
selected by Rajendran & Daniel (2002: 166168).
Gmelina villosa Naves (1877: t 215). Type: unknown.
Gmelina integrifolia Hunter ex Ridl. (Ridley 1909: 101
102); Lam & Bakhuizen (1921: 70). Type: [Penisular
Maylasia, Penang], Prince of Wales Island, Hunter
s.n. (holotype not located).
Gmelina tomentosa H. R. Fletcher (1938b: 204 205);
Moldenke (1984h: 181 182), synon. nov. Type:
Thailand, Rachasima, Ban Chum Seng, Korat, 23
May 1929, Nai Noe 211 (holotype E scan seen;
isotype K!).
Gmelina tonkinensis Moldenke (1940: 419; 1984h: 182),
synon. nov. Type: Vietnam, Tonkin, Fu Chap, to
the rocks of Notre Dame, May or June 1887,
Balansa 3807 (holotype LE (n.v.); isotypes K! ×2
sheets, L!, NY scan seen, P! ×4 sheets).
Shrubs to small trees, sometimes straggling, 1 5
(8) m high, DBH 2.5 20 (70) cm. Bark smooth,
(pale) grey; wood hard. Twigs villous; spines usually
present, up to 30 mm long. Leaves elliptic to ovate,
rarely round, 2 9.5 (14) ×1.5 5(8.5) cm, apex
rounded to acuminate, base rounded to cuneate,
306 KEW BULLETIN VOL. 67(3)
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2012
margins entire to rarely 1 5-lobed; upper surface
glabrous or with a few hairs on veins, green, pubescent
when young; lower surface velutinous, hairs usually
yellowish, covered with white peltate scale; sometimes
with discoid glands at base; veins 3 4pairs,
sometimes 3-veined from base. Petiole 6 30 mm
long, half terete, villous. Inorescence terminal, 2.5
7 cm long, erect, villous; bracteoles lanceolate or
ovate-lanceolate, 10 20 ×(1.5)48 mm, apex
acuminate, caducous. Calyx 5-lobed, sometimes
appearing fewer, densely pubescent, becoming more
glabrous over time, discoid glands present, peltate
scales present; owering calyx 3 5×34 mm, lobes
0.5 1 mm long, apex rounded to acute; fruiting calyx
48 cm diam., margin undulated, patent. Corolla
4-lobed, covered with yellow hairs; anterior lip 3-
lobed; mid-lobe 4.5 13 ×4.5 14 mm, apex
acute, patent, sometimes reexed; side lobes 3.5
7.5 ×37 mm, apex rounded to acute; posterior
lip 5 9×210 mm, apex acute to emarginate, patent
to reexed; tube 10 16 mm long, infundibulate.
Stamens inserted at apex of tube, yellow; anterior pair
15 17 mm long, anthers 1.5 2.5 mm long; posterior
pair, fertile, 10 18 mm long, anthers 1.5 2mmlong.
Ovary glabrous; style 20 25 mm long, stigma c. 2.5 mm
long. Fruit (dried) 10 20 ×820 mm, globose to
obovoid, glabrous, yellow when mature.
DISTRIBUTION. India: Andaman and Nicobar Islands;
Myanmar to South China and Vietnam; south to
Malaysia; Philippines: known from a number of
islands: Basilan, Mindoro, Mindanao and Negros;
Indonesia: from Sumatra to the Moluccas (Map 4).
Cultivated throughout the tropics.
SELECTED SPECIMENS EXAMINED. [East Indies], anon. in
Herb. Linn. 7801] (holotype LINN). CHINA. Yunnan:
Kishuanbanna, 1980, Chow & Wan 80202 (K). INDIA.
Nicobars: Feb. 1875, Kurz 26058 (K). INDONESIA.
Amamba Islands: Terempak, 31 March 1928, Henderson
20147 (K, SING); Zuid Sumatra: Palembang, 31 Aug.
1947, de Raadt 21 (L); Sumatra: Simaloer, 13 Feb. 1918,
Schmael 239 (L); Djambi, Doesoen Baroe, Sept. 1925,
Posthumus 862 (L); North Sumatra, Medan, 27 Feb. 1928,
Lörzing 12957 (K); East Java: Besuki, Blambangan Penin-
sula, 28 May 1957, Jacobs 4923 (L); Kalimantan: Barat,
Gunung Palung Nat. Park, Dec. 1997, Laman et al. TL529
(K);EastBorneo:Berau,SlopesofMtNjapaonKelaiR.,
13 Oct. 1963, Kostermans 21273 (L); SE Borneo, Hajoep, 2
June 1908, Winkler 2270 (L); Celebes: Kendari, Gunung
Abesu, 6 Feb. 1986, Moh. Amir 70 (L), NE Celebes,
Minahasa, Tasikoki, near Kema, SE of Mount Klabat, 5
July 1956, Forman 407(L);CentralSulawesi:c.25km
south of Palu, 21 April 1979, van Balgooy 2955 (L); Pulau
Buton: Wakunti, Bau-bau, 26 June 1978, Widjaja 551 (L);
East Bali: Kunu, 15 Oct. 1985, van Balgooy 5270 (L); East
Sumbawa: Bay of Danggar, 22 Sept. 1984, Snellius-II
11151A (L); Lombok: N- Seite of Rindjani Vulkan
gebirge, 30 April 1909, Elbert 750 (L); Flores, 16
Jan. 1973, Verheijen 3224 (L); West Timor: Usapi, 7
June 1981, Kooy 1302 (L); Alor Ketjil: Kabola penin-
sular, 4 May 1938, Jaag 469 (L); Ambon, 2 April 1918,
Kornassi 1079 (L); West Ceram: 1918, Rutten 1667 (L).
MALAYSIA. Penang: 1824, Phillips s.n. (K); Perak: Bota
Kiri, near Ipoh, 11 March 1958, Shah 322 (K); Sabah:
Keningau, Ulu Sungai. Pingas- pingas, 19 March
1988, Fidilis & Asik 122120 (K). MYANMAR. Toungoo
Distr.: 11 Aug. 1911, Lace 5397 (K); Kyouk Talong:
Irrawaddy R., 28 Sept. 1826, Wallich 1820 (K-W).
PHILIPPINES. Basilan: 17 Jan. 1904, Hallier 4295a (L);
Manaul: Mansalay, Mindoro, Dec. 1952, Sulit 17009
(K). SINGAPORE. near Changi Road, 13 Jan. 1933,
Teruya 2163 (K). THAILAND. Phuket: near airport, 9
May 1968, van Beusekom & Phengkhlai 498a (K);
Ratchasima: Pak Thong Chai, 13 May 1969, Phengna-
ren 639 (K); Siam: Kamboerie, Teysmann 5941 (K);
Ban Chum Seng, Korat, 23 May 1929, Nai Noe 211
(holotype E; isotype K). VIETNAM. Tonkin: Fu Chap,
to the rocks of Notre Dame, May or June 1887,
Balansa 3807 (isotypes K, L, NY, P).
HABITAT. Growing in primary and secondary vegetations.
Soil sandy or sandy loam or sandy clay, often poor,
sometimes over basalt or limestone; alt. 0 600 m.
CONSERVATION STATUS. Least Concern.
PHENOLOGY. Flowering Jan. Oct.; fruiting from
Feb. Oct.
VERNACULAR NAMES. Thailand: (Kang- or Nom-) Maow
or Tam mio; Peninsular Malaysia: Bulangan,Bulang or
Pokok Buah dulang (Malay language): Sabah: Bulangan
(Malay Language), Kutang or Belingkot (Murut Lan-
guage), Tangginang or Kutang (Dusun Language), Taring
pelandok (Kadayan Language); Sumatra: Pakok boelang
(Riau archipelago), Pokak baelandga (Pulau Lingga),
Bariang (Simaloer island), Warengen (Medan); Wareng
(Javanese language); Sulawesi: Pundanga (Palopo),
Rerubo,Kayu palapi or Tanggalasi (Kendari), Bidara
(Pulau Buton); Kayu palapi (Laweha), Wewenganga (Min-
hassa); Moluccas, Ambon: Karanjam, Alor Ketjil: Lombaul,
South West Timor: Hau bako (Dawan language).
USES. In India, an infusion of the leaves is used for eye
complaints (Rajendran & Daniel, 2002). In Peninsular
Malaysia several medicinal uses are known (Burkill 1966).
NOTES. This species is recorded from a small area in
Queensland, Australia (Munir 1984). The general distri-
bution of the species goes as far south as the Flores and
Sumba Islands (Map 4)andwasnotdiscoveredin
Australia until the 1970s. It is therefore very unlikely
that this is a native of Australia. The fact that it is
cultivated throughout the tropics and that it is becoming
a pest problem in Queensland supports this assumption.
The differences between Gmelina elliptica and G.
asiatica are usually easily to determine (G. elliptica has
leaves which are densely villous beneath, and G. asiatica
haseitherglabrousleavesorleaveswithonlyafewhairs
on the veins beneath). However, in Borneo, Sumatra,
307A REVISION OF THE GENUS GMELINA (LAMIACEAE)
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2012
Java and the Lesser Sunda Islands there are specimens
which seem to be morphologically intermediate; these
specimens have lobed, somewhat hairy leaves with many
peltate scales on the lower surfaces, the typical form
having leaves with an entire margin, densely hairy and
few peltate scales on the lower surfaces. On the specimen
Lörzing 12957, the typical form for G. elliptica and the
intermediate form are both present, and in the remark
on the label, he states Young shoots differ especially in
their leaves.Thisisconrmed by some of the other
specimens with apparently intermediate leaf morpholo-
gy: in these the bigger leaves are more hairy and less
lobed and are, as such, more typical for G. elliptica than
the smaller leaves. Given that typical G. asiatica does not
naturally occur on these islands and that G. elliptica is
native to these areas, I have concluded that these
intermediate specimens should be seen as part of G.
elliptica. More research is needed to determine whether
these forms represent either hybrids between G. elliptica
and the introduced G. asiatica or a natural variation
within G. elliptica. This variation is one of the main
reasons why G. asiatica and G. elliptica are sometimes
seen as two forms of one species.
I have not seen any type material of Gmelina
integrifolia Hunter ex Ridl. (Ridley 1909: 101 102),
but from the description it is clear that it is part of the
variation of G. elliptica.ThetypematerialofG.
tonkinensis Moldenke, G. tomentosa H. R. Fletcher and
G. vestita Wall. fall within the total morphological
variation of G. elliptica Sm. and is therefore placed into
its synonomy.
9. Gmelina evoluta (Däniker) Mabb. (Mabberley 1998:
316). Vitex evoluta Däniker (1933: 408). Type: New
Caledonia, Koumac, foot of Mt Kaala, 25 Feb. 1925,
Däniker 1228 (holotype Z scan seen; isotype Z ×2
sheets, scan seen).
CONSERVATION STATUS. Least Concern.
NOTE. For a description and more information see
Mabberley in Mabberley & de Kok (2004).
10. Gmelina fasciculiora Benth. (Bentham 1870: 65);
Moldenke 1984g: 106 107). Vitex leichhardtii var.
glabrifolia F. Muell. (Mueller 1865: 35), Benth. (Bentham
1870:65asglabrata). Type: Australia, Queensland,
Rockingham Bay, 16 Oct. 1868, Dallachy s.n. (lectotype
MEL 583159 scan seen; isolectotypes K! ×2 sheets)
selected by Munir (1984:102).
Trees (4 )1030 m high, DBH 12 80 cm. Bark
somewhat scaly, grey. Twigs tomentose when young,
hairs brown, spines absent. Leaves ovate-elliptic, 3
16 ×2.5 9 cm, apex obtuse to obtusely acuminate,
base rounded to cuneate, margin entire, coriaceous;
upper surface shiny; lower surface covered with
peltate scales; veins 4 7 pairs, sometimes 3-veined
from base. Petiole half-terete, channelled, 5 30 mm
long, glabrous. Inorescence terminal, densely tomentose,
10 25 cm long; bracteoles clavoid to round, 2.5
5×24 mm, sessile. Calyx 5-lobed, green,
tomentose outside, hairs rusty brown; owering calyx
2.5 4×2.5 3.5 mm; lobes triangular, 0.4 0.6 ×0.5
1 mm, apex acute, erect; fruiting calyx 7 15 mm diam.
Corolla 5-lobed, tomentose when young, becoming less
so with age, white to blue; anterior lip 3-lobed, mid-lobe
ovate to broadly elliptic, 5 10 ×58 mm, apex
rounded to acute, purple with a yellow marking at base;
side lobes 4 8×36 mm, oblong- ovate long, apex
rounded; posterior lip 2-lobed, 3 7×36 mm; tube 8
14 mm long. Stamens inserted at apex of tube, few
glandular hairs; anterior pair 10 11 mm long,
anthers c. 1.2 mm long; posterior pair, fertile, 8
9mmlong,anthersc.1mmlong.Ovary12mm
diam., glabrous, villous at apex. Style 12 15 mm
long, stigma c. 3 mm long. Fruit globular, 10
20 mm diam., apex rounded glabrous, smooth, shiny,
glands absent, violet or blue to bright purple or violet
when mature, taste sour.
DISTRIBUTION. Australia: Northern Queensland. Map 3.
SELECTED SPECIMENS EXAMINED. AUSTRALIA. Queensland:
Julatten, 11 Jan. 1977, Gray 222 (K); Boonjee, 13 Dec.
1979, Gray 1591 (K); Barron, 15 Nov. 1972, Hyland 6486
(K); Mission Beach, 58 Miles SE of Atherton, 14 Nov.
1963, Hyland 3720 (K); Rockingham Bay, 16 Oct. 1868,
Dallachy s.n. (lectotype MEL; isolectotype K, P).
HABITAT. Growing in rainforests or along rivers and
beaches; alt. 0 760 m.
CONSERVATION STATUS. Least Concern.
PHENOLOGY. Flowering Nov. Dec.; fruiting Nov. Jan.
VERNACULAR NAMES. North Queensland Beech or
Grey Teak.
USES. The wood is used in furniture making and
woodcarving.
11. Gmelina hollrungii de Kok sp. nov. aG. schlechteri
glandulis disciformibus e bracteolis carentibus differt.
AG. evoluta glandulis cupuliformibus ad basin folio-
rum dispositis differt. Typus: [Papua New Guinea]
New Guinea, Augusta Station, 1887, Hollrung 651
(holotypus K!; isotypi P!).
http://www.ipni.org/urn:lsid:ipni.org:names:77120321-1
Gmelina macrophylla Schumann in Schumann &
Hollrung (1889: 120), nom. illegit., non Gmelina
macrophylla Wall. ex Schauer (1847: 680). Type:
[Papua New Guinea] New Guinea, Augusta Station,
1887, Hollrung 651 (holotype B (n.v.); isotypes K!,
P! ×2 sheets).
308 KEW BULLETIN VOL. 67(3)
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2012
Trees, 6 40 m high; DBH 30 75 cm. Bark smooth to
ssured,greytobrown;Twigsvelutinous,spines
absent. Leaves elliptic-round, oblong or obovate, 10
36 ×525 cm, apex rounded to shortly acuminate,
base rounded to cuneate, margins entire, coriaceous;
upper surface glabrous, few hairs on veins, (yellow)
green, shiny; lower surface glabrous, pale green to
yellow, few hairs on veins, peltate scales absent, pale
beneath, with cup-like glands at base; veins 5 8 pair,
sometimes 3-veined from base. Petiole 15 80 mm
long, half-terete, channelled, sparsely hairy. Inores-
cence terminal, 15 30 cm long, brown tomentose;
bracteoles linear, 2.5 4×0.5 1 mm long, apex
acute, sessile. Calyx 5-lobed, sparsely to densely hairy;
with scattered black discoid glands; owering calyx
35×34 mm; lobes 0.8 1×11.2 mm, apex
rounded; fruiting calyx up to 8 mm diam., patent.
Corolla 5-lobed, velutinous, white, pale lilac to mauve
or purple; anterior lip 3-lobed, mid-lobe oblong, 5.5
10 ×34 mm, apex rounded to acute, lilac with a
yellow spot at base; side lobes oblong, 3.5 8×25mm,
apex rounded to acute, oblique; posterior lip 2-lobed,
oblong to ovate- oblong, 3 7×25 mm; tube 7
12 mm long. Stamens inserted at apex of tube, few
glands; anterior pair 6 10 mm long, anthers 2 2.5 mm
long; posterior pair, fertile, 6 8 mm long, anthers c.
2 mm long. Ovary 1 1.5 mm diam., glabrous with some
glands at apex, style 12 20 mm long. Fruit (dried) 8
25 ×616 mm, obovoid, apex truncated, glabrous,
shiny, blue or red to purple when mature. Fig. 3.
DISTRIBUTION. Indonesia: Moluccas: Morotai and Aru
Islands, West Papua; Papua New Guinea; Northern
Australia; the Solomon Islands. Map 2.
SELECTED SPECIMENS EXAMINED. AUSTRALIA.Port Essing-
ton, Armstrong 556 (K); Rockingham Bay, Dallachy s.n.
(K). INDONESIA. Moluccas: Morotai, Mt Permatang, 23
May 1949, Main & Aden 941 (K); Aroe Islands, Pulau
Trangan, 1 July 1938, Buwalda 5431 (K, L); Aru
Archipelago, Kobroor, 25 Oct. 1994, Nooteboom 5880
(K, L); Aru Islands, Pulau Trangan, 23 Oct. 1994, van
Balgooy 6610 (K); West Papua: Sorong, 1 Oct. 1948,
Pleyte 1094 (K, L). PAPUA NEW GUINEA. Augusta Station,
1887, Hollrung 651 (holotype K; isotypes K, P); Morobe
Province: Lae, Saru R., 20 July 1970, Streimann et al.
47995 (SING); Wampit, Bupu Village, 13 July 1967,
Millar 22979 (L); Central Province: Brown R., Dec.
1963, Kumul 13071 (K); Western Highlands Province:
Hagen distr., Road above Baptist Mission, 9 July 1968,
Miller 36668 (L); Baiyer R. Sanctuary, 5 June 1980,
Wiakabu 73468 (K, L); East Sepik Province: Hunstein
range, Mt Samsai, 24 July 1990, Takeuchi 6373 (K);
Northern Province: Isuarava, 24 Feb. 1936, Carr
15748 (L, SING). SOLOMON ISLANDS. Santa Ysabel,
21 Sept. 1965, Hunt 2649 (K).
HABITAT. Growing in rainforest, sometimes as canopy
tree, or in more open types of forest; alt. 30 1400 m.
Soil sandy or clayey, sometimes over sandstone,
limestone or ultra-basic rock.
CONSERVATION STATUS. Least Concern.
PHENOLOGY. Flowering July Nov.; fruiting July Feb.
Fruit eaten by cassowaries.
VERNACULAR NAMES. Indonesia: Adoen or Adun mangai
laplabai (Aru Islands); Anjoes (Manokwari, Warnapi).
USES. On the Aru Islands the species is considered a
good tree for timber.
NOTES. Gmelina hollrungii differs from G. schlechteri in
having bracteoles without disc-like glands and differs
from G. evoluta in having cup-like glands at the base of
the leaves.
12. Gmelina lecomtei Dop (1914: 322); Vũ(2007: 141
142). Type: [Vietnam] Indo-Chine, Annam, Chapa à
Muong-Xen, 31 Oct. 1911, Lecomte & Finet 421
(holotype P scan seen; isotype HM (n.v.), K!).
Shrub to trees 3 7 m high, DBH c. 10 cm. Twigs
densely hairy when young, spines absent. Leaves ovate
to elliptic, 4 13 ×312 cm, margin enire, sometimes
slightly lobed, apex acute to acuminate, base slightly
cordate or cuneate to slightly decurrent, papery;
upper surface glabrous, with few hairs on veins at
base, bluish green; lower surface velutinous, hairs
usually yellow, covered with peltate scales, few discoid
glands present at base; veins 3 6-pairs, sometimes 3-
veined from base. Petiole 1.5 7 mm, terete to half-
terete, channelled, velutinous. Inorescence terminal,
512 cm long, velutinous to sparsely hairy, covered
with peltate scales; bracteoles 7 13 ×5.5 10 mm,
elliptic, margin slightly lobed, sparsely hairy, covered
with peltate scales, often with discoid glands present.
Calyx 12 15 ×11.5 15 mm, 5-lobed, covered with
peltate scales, sparsely hairy, often with discoid glands
present, not accrescent; lobes triangular, 4 7×5
7 mm, anterior lobes biggest, apex acute, erect.
Corolla 5-lobed, covered with peltate scales, white
to pink, odourless to fragrant; anterior lip 3-lobed,
mid-lobe rounded, 5 6×76mmlong,apex,
patent; side lobes oblong, 4 5×67 mm, apex
rounded, erect; posterior lip 2-lobed, 4 9×7
10 mm, oblong, apex rounded; tube 10 14 mm
long, infundibular. Stamens inserted at apex of tube,
glabrous, sometimes with a few glands; anterior pair
c. 10 mm long, anthers c. 3 mm long; posterior pair,
c. 7 mm long, anthers c. 2 mm long. Ovary pubescent
at apex, covered with peltate scales at apex; style c.
10 mm long, glabrous. Fruit (dried) cylindrical, 15
40 ×10 12 mm, apex truncate, sparely hairy,
covered with peltate scales, discoid glands present,
grey.
DISTRIBUTION. China: Yunnan, Kwantung and Hainan;
Vietnam and Laos. Map 5.
309A REVISION OF THE GENUS GMELINA (LAMIACEAE)
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2012
SPECIMENS EXAMINED. CHINA. Kwantung: Sin-fung distr.,
Fuk Lung monastery, Hau Tong Shan, 1 19 June
1938, Taam 808 (K); Sin-fung distr., Cha-Ping village,
Ah Po Kai Shan, 1 24 May 1938, Taam 635 (K); Fang
Cheng distr., Taan Faan, Kung Ping Shan and
vicinity, 15 24 Aug. 1936, Tsang 26664 (K); Fang
Cheng distr., Taan Faan, Kung Ping Shan and
vicinity, 15 24 Aug. 1936, Tsang 26666 (K); Hainan:
Ngai Distr., Yeung Ling, Shan, 13 June 1932, Lau 75
(K); Lingshin, 25 April 1932, Ko 52188 (K); Yaichow,
8 July 1933, Liang 61985 (K). LAOS. Khammouan Nakai
Distr.: 19 May 2006, Newman et al. LAO 1154 (L).
Fig. 3. Gmelina hollrungii.Ahabit; Bdetail leaf underside, cup-like glands; Cower side view; Dcalyx; Ecorolla; Fstyle and ovary;
Gstigma. From Takeuchi 6373. DRAWN BY JULIET BEENTJE.
310 KEW BULLETIN VOL. 67(3)
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2012
VIETNAM. Annam [Central region]: Chapa à Muong-
Xen, 31 Oct. 1911, Lecomte & Finet 421 (holotype P);
Tonkin: Ha-coi, near Chuk-phai, Taai Wong Mo
Shan, 3 May 22 June 1939, Tsang 29235 (K, L);
Dam-ha, Lung Wan Village, Sai Wong Mo Shan, 18
May 5 July 1940, Tsang 29869 (K, SING); Tien-yen,
Kau Nga Shan and vicinity, 23 Sept. 7Oct.1940,
Tsang 30588 (K, SING); Ha-coi, Tong Fa market, Taai
Wong Mo Shan and vicinity, 11 23 Sept. 1939, Tsang
29546 (K, SING); Bankeuin, near Quang Yen, Aug.
1885, Balansa 937 (K).
HABITAT. In open forest and thickets, in dry sandy or
clayey soil; alt. 200 1000 m.
CONSERVATION STATUS. Least Concern.
PHENOLOGY. Flowering Aug. Sept.; fruiting June Aug.
VERNACULAR NAME. China: Yue nan shi zi.
NOTES. This species is often confused with Gmelina
racemosa and the other Indo-China endemics; the
differences are given in Table 1.
13. Gmelina ledermannii H. J. Lam (1919: 226);
Moldenke (1984g: 109 110); Type: [Papua New
Guinea] New Guinea, Malu, near Sepik-river, 3 March
1912, Ledermann 6537 (lectotype K!), selected here.
Tree 10 20 m high. Bark dark, rough. Twigs sparsely
hairy when young, with interpetiolar ridge present, spines
absent. Leaves elliptic to slightly obovate, 8 19 ×5.5
9.5 mm, slightly oblique, apex rounded to acute, base
rounded, rarely cuneate, margins entire; upper surface
glabrous, dull to glossy, dark green; lower surface
glabrous with few simple hairs on veins, hairy when
young, pale green, peltate scales absent; cup-like glands
at base; veins 6 8 pairs, sometimes 3-veined from base.
Petiole 2 4 cm long, half-terete, sparsely hairy.
Inorescence terminal, narrow, up to 20 cm long, with
long side branches, velutinous; bracteoles rounded (in
particular on main axis) to lanceolate, 4 11 ×4.5
6.5 mm, apex long acuminate, sessile, with several disc-
Map 5. Distribution of Gmelina lecomtei (); Gmelina lepidota (); probable natural distribution of Gmelina philippinensis () with
grey-shaded square as a questionable data point; distribution of Gmelina peltata ().
Table 1. Differences between Gmelina chinensis,G. lecomtei and G. racemosa.
G. lecomtei G. lecomtei G. chinensis
Lower leaf surface hairy glabrous hairy to glabrous
Calyx lobes present present minute
Bracteoles (mm) elliptic, 7 13 × 5.5 10 lanceolate, 7.5 25 × 2.5 10 lanceolate, 8 15 × 4 6.5
Bracteole margin slightly lobed entire entire
Corolla (anterior lip length, mm) small, (5 6) big, (5 10) big, (11 17)
Ovary apex sparsely hairy velutinous glabrous
311A REVISION OF THE GENUS GMELINA (LAMIACEAE)
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2012
like glands present. Calyx 3.5 5×3.5 4 mm, 5-lobed,
sometimes appearing fewer, velutinous sometimes with a
glabrous margin, outer surface with discoid glands; lobes
up to 0.5 mm long, apex acute to rounded; fruiting calyx
56 mm diam., erect. Corolla 5-lobed, outer surface
hairy, densely glandular on inside of lobes, cream to
yellow; anterior lip 3-lobed; mid-lobe oblong, 7.510 ×3
5.5 mm, apex rounded, mauve with two yellow marks;
side lobes 4.5 5×4.5 5 mm long, apex rounded;
posterior lip, 2-lobed, 4.5 5×3.5 4 mm, apex
rounded; tube 6.5 7.5 mm long. Stamens inserted at
apex of tube, glabrous, glands present; anterior pair 8
11 mm long, anthers c. 2 mmlong; posterior pair, fertile,
68 mm long, anthers c. 1.5 mm long. Ovary glabrous
with a few hairs at apex. Style c. 10 mm long, stigma c.
1.5 mm long. Fruit (dried) 10 11 ×56mm,tubular,
apex rounded, glabrous, smooth, glossy, purple or
bright blue when mature.
DISTRIBUTION. Indonesian New Guinea and Papua New
Guinea: Sandaun and east Sepik Province (see Map 3).
SPECIMENS EXAMINED. INDONESIA. New Guinea: Mim-
ika, Najaja (Oeta), 21 June 1941, Neth. Ind. For. Service
bb. 32852 (Lundquist 133) (K). PAPUA NEW GUINEA.
Morobe Province: Dschischugari, 25 May 1909,
Schlechter 19566 (K); East Sepik Province: New-Guinea,
Malu, near Sepik-R., Ledermann 10455a (K); Malu, near
Sepik-R., 3 March 1912, Ledermann 6537 (lectotype K);
along Yapa (Hunstein R.), 1 Aug. 1966, Hoogland &
Craven 10799 (K, L); near Ambunti, 31 May 1966,
Hoogland & Craven 10167 (K, L); Sandaun Province:
Telefomin distr., W of Fiak airstrip, 27 March 1992, Frodin
et al. 2586 (K); Papua, Eastern Division, Asia R., 11 May
1926, Brass 1376 (K).
HABITAT. A tree in stunted or secondary forest, some-
times along rivers, in sandy or poor soil; alt. 90 800 m.
CONSERVATION STATUS. Least Concern.
PHENOLOGY. Flowering May Aug.; fruiting March Aug.
VERNACULAR NAMES. Papua New Guinea: Yoni (Wagu
Language); Bouwen or Boing (Waskuk Language) or As
Kobokim (Miyanmin Language). Indonesia: Boetao
(Mimika area).
USES. Wood used for construction and canoe building
(as it is easily worked) and rewood.
NOTES. In the originally description three specimens
were mentioned: Ledermann 6537 and 10455a and
Schlechter 19566. The Ledermann 6537 specimen at K is
selected here as the lectotype as it is the only one with
mature owers.
14. Gmelina leichhardtii (F. Muell.)Benth. (Bentham
1870: 66); Moldenke 1984g: 110 115); Munir (1984:
99 102); Vitex leichhardtii F. Muell. (Mueller 1862: 58).
Type: [Australia] New South Wales, Myall Creek, 30
Nov. 1843, Leichhardt s.n., (lectotype MEL scan seen),
selected by Munir (1984: 99).
Tectona australis W. Hill (1862: 20), synon. nov. Type:
[Australia] Queensland Woods, [London exib.
1862] Hill 30 (holotype K!).
Tree 10 40 m high, DBH up to 120 cm, often semi-
deciduous. Bark aky or ssured, (dark) grey. Twigs
tomentose, light brown, spines absent. Leaves narrowly
elliptic or ovate-elliptic, 6 15 ×49 cm, apex
rounded to acute, papery, base rounded to cuneate,
margins entire, upper surface glabrous, sometimes
with hairs on veins, dark green; lower surface densely
tomentose, peltate scales present, basal glands absent;
green; veins 7 8 pairs, sometimes 3-veined from base,
lower surface with strong ladder-like tertiary venation.
Petiole 15 55 mm, half-terete, channelled, tomen-
tose. Inorescence 10 25 cm long, tomentose; brac-
teoles linear, 1.5 2×11.2 mm, apex acute,
caducous. Calyx 5-lobed, sometimes appearing fewer,
pubescent, black discoid glands present; owering
calyx 4 6×36 mm; lobes triangular, up to
0.2 mm long, apex acute; fruiting calyx 10 20 mm
diam. Corolla 5-lobed, tomentose outside, glandular
hair inside, (cream-) white; anterior lip 3-lobed; mid-
lobe spathulate to ovate, 8 13 ×710 mm, apex
rounded, violet with two yellow spots at base; side
lobes elliptic-oblong, 5 10 ×58mm,apex
rounded, sometime oblique; posterior lip, 2-lobed,
lobes oblong, 6 9×56 mm, apex rounded;
tube 4 8 mm long, with two bluish stripes ending
in yellow spots at the lament insertion. Stamens
inserted at apex of tube, covered with glandular
hairs; anterior pair 12 14 mm long, anthers 1.5 2mm
long; posterior pair, fertile, 10 11 mm long,
anthers 1.2 1.5 mm long. Ovary 1 2mmdiam.,
sparsely pubescent. Style 10 16 mm long, stigma
c. 1.5 mm long. Fruit 11 20 ×13 15 mm, almost
globular, apex rounded, glabrous, bluish-purple
when mature, reported to have a disagreeable taste.
DISTRIBUTION. Australia: from the coast of central
Queensland to the south coast of New South Wales.
Map 2.
SELECTED SPECIMENS EXAMINED. AUSTRALIA. Queens-
land: Queensland Woods, [London exib. 1862] Hill
30 (holotype K); South Kennedy, NE of Eungella, 26
Nov. 1989, McDonald 4471 (K); Brisbane Botanic
Gardens, 31 Oct. 1930, Hubbard 4738 (K); Dalrymple
Heights and vicinity, July Nov. 1947, Clemens s.n. (K);
New South Wales: Botanic Gardens Sydney, Oct. 1908,
Boorman s.n. (L); Sydney Woods, 1854, MacArthur 193
(K); New Holland, Morton Bay, 1845, Leichhardt s.n.
(P); Myall Creek, 30 Nov. 1843, Leichhardt s.n.,
(lectotype MEL).
HABITAT. Growing in primary and secondary vegetations.
CONSERVATION STATUS. Least Concern.
PHENOLOGY. Flowering Oct. Jan.; fruiting Jan. April.
VERNACULAR NAME. White or Native Beech.
312 KEW BULLETIN VOL. 67(3)
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2012
USES. The wood is used in furniture making and wood
carvings. The species has been commercially over-
exploited in the past (Francis 1951).
NOTES. The type material of Tectona australis W. Hill
falls within the total morphological variation of
Gmelina leichhardtii (F. Muell.) Benth. and is thefore
placed into its synonomy.
15. Gmelina lepidota Scheff. (Scheffer 1876:4142)
Moldenke 1984g: 115 116). Type: [Indonesia, New
Guinea] LIle de Tow, 18 Aug. 1871, Teysmann 6744
(holotype BO!; isotypes L!, NY scan seen).
Gmelina misoolensis Moldenke (1952: 54; 1984g: 118),
synon. nov. Type: [Indonesia] New Guinea, Misool
Island, Fakal, 30 Sept. 1948, Pleyte 1087 (holotype
BO!; isotypes BO! L! NY scan seen).
Gmelina lepidota var. lanceolata Moldenke (1958: 325
326; 1984g: 116), synon. nov. Type: Papua New
Guinea, New Britain, Open Bay, Nantambu, June
1945, Mair 1894 (holotype LAE (n.v.); isotypes L! ×
2 sheets).
Trees 7.5 30 m high, DBH 3 60 cm. Bark (dark) grey-
brown, ssured, wood light straw coloured. Twigs hairy
when young, glabrous later, spines absent. Leaves elliptic to
lanceolate, rarely ovate, 4 18 ×2.3 7cm,apex
rounded to acuminate, base cuneate, margins entire;
upper surface glabrous, dark green; lower surface
glabrous with a few hairs on the veins, lightly brownish
green or dull grey brown, covered with peltate scales, with
discoidglandsclusteredaroundthebaseandalongthe
vein; veins 4 7 pairs. Petiole half-terete, slightly
channelled, 1 3 cm, sparsely covered with peltate scales.
Inorescence terminal and axillary, with short side cymes,
47 cm long, sparsely hairy; bracteoles lanceolate, 0.8
2×0.1 1.5 mm, apex acute, persistent. Calyx 5-lobed,
glabrous, covered with peltate scales, olive, greenish
purple or purple-brown, outer surface with discoid
glands; owering calyx 2 3.5 ×23 mm; lobes
triangular, 0.5 0.8 mm long, apex acute; fruiting calyx
56 mm diam., patent. Corolla 5-lobed, outer surface
velutinous, inside glabrous, sparsely glandular, cream to
yellow or (light) purple; anterior lip 3-lobed; mid-lobe
spathulate 4.5 5×34 mm, apex rounded; side lobes
1.5 3×22.5 mm, apex rounded; posterior lip 2-lobed,
24×1.5 2 mm, apex rounded; tube 8 9mmlong.
Stamens inserted at apex of tube, white, base covered
with hairs, mauve; anterior pair 6 10 mm long, anthers
c. 1 mm long; posterior pair, fertile, 5 8mmlong,
anthers 0.8 mm long. Ovary c. 1 mm diam., glabrous;
style 8 11mmlong,stigmac.1.5mmlong.Fruit (dried)
920 ×625 mm, ellipsoid to clavoid, apex rounded,
glabrous, red or pale purple when mature.
DISTRIBUTION. Indonesia: Moluccas: Morotai Island;
the islands of New Guinea and New Britain. Map 5.
SELECTED SPECIMENS EXAMINED. INDONESIA. Moluccas:
Morotai, Gunung Pare Pare, 28 May 1949, Main & Aden
(exp.Kostermans)1292(K,L,SING);Morotai,Gunung
Pare Pare, 28 May 1949, Main & Aden (exp. Kostermans)
1276 (K, L, SING); Obi Island: Jikodolong, 26 Nov. 1974,
de Vogel 4342 (L); [Indonesia, New Guinea] LIledeTow,
18 Aug. 1871, Teysmann 6744 (holotype BO!; isotype L);
Misool Island, Fakal, 30 Sept. 1948, Pleyte 1087 (holotype
BO; isotypes BO, L, NY); West Papua: NederlandsNew
Guinea, Radjah Ampat, Weigeo Island, E bank of
Majalibit Bay, Waifoi, 18 Jan. 1955, van Royen 5222 (K,
L); Vogelkop Peninsula, Ajamaroe, 8 May 1958, Versteegh
BW 4998 (L); New Guinea, Hollandia, 21 Dec. 1954,
Versteegh BW 658(L);WestIrian,nearSukarnapura[=
Djajapura], 4 August 1966, Kostermans & Soegeng 148 (K);
West Irian, in town of Sukarnapura [= Djajapura], 7 Aug.
1966, Kostermans & Soegeng 230 (K); West Irian, above
Hollandia [ = Djajapura], 15 Aug. 1966, Kostermans &
Soegeng 369 (K); Netherlands New Guinea, Hollandia
[ = Djajapura], 17 Jan. 1957, Versteegh BW 4696 (SING).
PAPUA NEW GUINEA. Western Province: near Lake
Daviumbu, 4 Oct. 1967, Pullen 7436 (L); New Britain
Province: Mt Tangis, Western Slopes, 29 May 1966,
Frodin 26866 (L); New Britain, Open Bay, Nan-
tambu, June 1945, Mair 1894 (isotypes L); Southern
Highland Province: Lake Kutubu, near Moro, 4
Oct. 1961, Schodde 2368 (L); Morobe Province:
Sikong Range, Poiyu road, 2 Jan. 1989, Takeuchi et
al. 4342 (L).
HABITAT. Growing from rainforest to xerophytic or
open vegetation, sometimes dominated by Myrtaceae
(notably Eucalyptopsis papuana C. T. White); alt. 10
1000 m. Clayey soil, sometimes over limestone, ser-
pentine, or volcanic soils.
CONSERVATION STATUS. Least Concern.
PHENOLOGY. Flowering and fruiting Aug. May.
VERNACULAR NAMES. Indonesia: Rafah (Maibrat
Language). Papua New Guinea, New Britain: Namavue
(Mt Tangis); Kanikehua (Kutubu Language).
USES. The wood is used for dugout canoes.
NOTES. Type material of Gmelina lepidota var. lanceolata
Moldenke and G. misoolensis Moldenke fall within the
total morphological variation of G. lepidota Scheff. and
is therefore placed into its synonomy.
16. Gmelina lignum-vitreum Guillaumin (1952: 539);
Moldenke (1984g: 116 118); Mabberley in Mabberley
& de Kok (2004). Type: New Caledonia, Forêt de Thy,
Aug. 1949, Sarlin 81 (holotype P scan seen; isotypes P ×
2 sheets, scans seen).
CONSERVATION STATUS. Listed as Critically Endangered
(IUCN red list download on 7 Jan. 2011 (www.iucnredlist.
org/apps/redlist)).
NOTE. For a description and more information see
Mabberley in Mabberley & de Kok (2004).
313A REVISION OF THE GENUS GMELINA (LAMIACEAE)
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2012