KEW BULLETIN VOL. 64: 587–625 (2009)
The genus Teijsmanniodendron Koord. (Lamiaceae)
R. P. J. de Kok1, G. Rusea2 & A. Latiff3
Summary. A revision of the genus Teijsmanniodendron Koord. (Lamiaceae) is presented with a summary of its
taxonomic history, keys, full descriptions, distribution maps, conservation assessments, ecological information and
ethno-botanical notes. In this treatment, 23 species are recognised, one new combination is made: Teijsmanniodendron havilandii (Ridl.) G. Rusea, 14 names are placed into synonymy for the first time, one species is neotypified,
five are lectotypified and six new species are described: T. bullatum G. Rusea, T. latiffii G. Rusea, T. obscurinerve G.
Rusea, T. punctatum G. Rusea, T. renageorgeae G. Rusea and T. zainudinii G. Rusea. One species name is validated, T.
scaberrimum Kosterm. ex G. Rusea and one species is excluded from the genus: T. petelotii Moldenke.
Key Words. Distribution, Lamiaceae, taxonomy, Teijsmanniodendron.
Introduction
Teijsmanniodendron (Lamiaceae) is a genus of trees
native to the rain forests of South East Asia. The genus
was revised as part of a PhD thesis at the Department
of Botany, Faculty of Life Sciences, National University
of Malaysia (Rusea 1998). This work is presented here,
with modifications, with information on its taxonomic
history, species delimitation and morphology, as well
as identification keys, full descriptions with distribution maps, conservation assessments, ecological and
ethno-botanical notes for all 23 recognised species.
This research moves on from Rusea & Latiff (2001)
which examined trichome diversity and its distribution
within the genus.
Following the discussion in Kostermans (1951) and
Moldenke (1980a) concerning the original spelling of
the genus name, we are using the original Teijsmanniodendron version over the later Teysmanniodendron.
Taxonomic History
The genus Teijsmanniodendron, originally proposed by
Koorders (1904), is named after Johannes Elias
Teijsmann (1808 – 1882), Curator of Botanic Gardens,
Buitenzorg (now Bogor), Indonesia. The genus was
based on one species only, T. bogoriense Koord.,
described from specimens gathered from two cultivated trees of unknown origin in the Botanical Garden
at Buitenzorg (Koorders 1904). Koorders distinguished Teijsmanniodendron from closely related Vitex
by its fruit, a one-celled, one-seeded indehiscent
capsule, rather than 1 – 4 one-celled, one-seeded
drupe. He concluded that this genus represented a
new subtribe (Teysmanniodendreae) within the tribe
Viticoideae (Verbenaceae). Lam (1919), basing his
work on specimens preserved at Leiden and Utrecht
herbaria, followed this. However, in addition he
described a new monotypic genus Xerocarpa H. J.
Lam (Lam 1919), now placed in Teijsmanniodendron
(Kostermans 1951). Later, (Lam & van den Brink
Bakhuizen 1921) revised the genus again, based on
more extensive material at Bogor. They transferred
two Vitex species (V. pteropoda Miq. and V. aherniana
Merr.) to Teijsmanniodendron and placed it in the
subtribe Teysmanniodendreae (Viticoideae, Verbenaceae) together with the monotypic genus Xerocarpa
H. J. Lam (Lam 1919). This was later confirmed by the
anatomical studies of Junell (1934). However, at least
Lam seems to think at the time that more species of
Vitex, and probably his own genus Xerocarpa, should be
placed in Teijsmanniodendron (Kostermans 1951).
Kostermans (1951) was the next to revise the genus,
recognising 12 species and one variety based on his
field observations and on specimens deposited both in
BO and SING. He considered Teijsmanniodendron to be
closely related to Vitex and its segregation perhaps
debatable. Provisionally segregating the two genera on
the basis of the swollen articulation of the petioles and
petiolules in Teijsmanniodendron, he also emphasised
that Teijsmanniodendron has an ovary with four ovules, of
which only one develops into a mericap, while in Vitex
the ovary with 4 ovules develops into 1 – 4 mericaps. He
also proposed two practical sections within Teijsmanniodendron, namely “Unifoliolatae” and “Plurifoliolatae”. The
Accepted for publication April 2009.
1
Royal Botanic Gardens Kew, Richmond, Surrey TW9 3AB, UK. e-mail: R.deKok@kew.org
2
Universiti Putra Malaysia, 43400, Serdang, Selangor, Malaysia.
3
Universiti Kebangsaan Malaysia, Bangi, Malaysia.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
588
former consists of species with simple leaves and the
latter of those with palmately compound 3 – 7-foliolate
leaves (Kostermans 1951). Following this delimitation,
he transferred seven species of Vitex to Teijsmanniodendron, notably T. coriaceum (C. B. Clarke) Kosterm., T.
hollrungii (Warb.) Kosterm., T. holophyllum (Baker)
Kosterm., T. novo-guineense (Kaneh. & Hatus.) Kosterm.
(= T. holophyllum), T. sarawakanum (H. Pearson)
Kosterm., T. smilacifolium (H. Pearson) Kosterm. and
T. subspicatum (Hallier f.) Kosterm. He continued
to work on the genus and later described three
more species: T. pendulum Kosterm. (= T. bogoriense),
T. simplicioides Kosterm. and T. sinclairii Kosterm.
(Kostermans 1962b).
Later, Moldenke (1980a & b, 1981) recognised 27
taxa comprising 22 species and five varieties, including
three new species, T. bintuluense Moldenke, T. borneënse
Moldenke (= T. bogoriense), and T. kostermansii Moldenke
(= T. bogoriense), a number of new varieties, and one
new combination: T. unifoliolatum (Merr.) Moldenke.
In the present study, Teijsmanniodendron is distinguished from Vitex in having well-developed and
conspicuously swollen articulations of the petioles
and petiolules. We recognise 23 species, six of which
are new to science. The two species recorded from
Vietnam (T. petelotii Moldenke and T. pierrei Moldenke)
are somewhat anomalous. T. peteloti has not been
accepted in the genus as it does not have a swollen
petiole base and apex, and the whorled leaves are
uncharacteristic for the genus and is most likely a Vitex.
T. pierrei is reduced to a synonym of T. holophyllum.
Taxonomic Position of Teijsmanniodendron
in the Lamiaceae
Traditionally this genus was placed in the Verbenaceae
(Koorders 1904; Lam 1919; Lam & van den Brink
Bakhuizen 1921; Junell 1934; Kostermans 1951;
Moldenke 1980a), although in the Index Kewensis it
was originally erroneously placed in the Araliaceae
(Prain 1908). The tribe Teijsmanniodendreae was
created by Koorders (1904) to accommodate this
genus. Later authors adopted Briquet’s (1897) system
of classification for the family and placed Teijsmanniodendron in the tribe Viticoideae, subtribe Teijsmanniodendreae. However, some authors placed the genus
within its own tribe in the subfamily Caryopterioideae
Briq. (Moldenke 1971; Raj 1983).
Cantino (1992) in his cladistic analysis based
mainly on morphological and anatomical data, placed
Teijsmanniodendron in a clade together with members
of the Viticoideae. This clade was part of a larger
group comprising mainly of taxa of Lamiaceae. In
conclusion, Cantino (1992) suggested that members
of the Viticoideae, including Teijsmanniodendron, and
other non-Viticoideae genera traditionally considered
as Verbenaceae such as Clerodendrum L., be incorpo© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
KEW BULLETIN VOL. 64(4)
rated into the Lamiaceae, leaving the Verbenaceae
restricted to subfamily Verbenoideae. This conclusion
has been supported by several molecular studies on
the families (Wagstaff & Olmstead 1997; Wagstaff et al.
1998 and Olmstead et al. 2001) and the new classification has now been generally accepted (Harley et al.
2004; Heywood et al. 2007).
Morphology
Leaflet shape, size and number of leaflets have been
found to be variable characters within species of
Teijsmanniodendron (e.g. T. pteropodum, T. subspicatum,
T. sarawakanum and T. hollrungii) both here and by
Kostermans (1951), and are therefore not considered
reliable for species delimitation. In species with
compound leaves, the number of leaflets seems to be
consistent in some species (T. renageorgeae and T.
bullatum), while in others (T. pteropodum, T. bogoriense, T.
ahernianum and T. coriaceum) it can vary from 3 – 7 per
leaf. Again this follows the observations of Kostermans
(1951) and the character is not considered in this
treatment for species identification.
Informative characters are the absence or presence
of indumentum pits in the lower leaf surface (Rusea &
Latiff 2001), colour and pattern of venation, the
texture of young leaflets (rough vs. smooth), and the
colour of dried leaves.
Whilst petiole length can be inconsistent in compound-leaved species, it is less so in those with simple
leaves and is hence somewhat reliable when distinguishing between the latter species. More functional is the
shape, absence or presence of indumentum and absence
or presence of a wing or appendages on the petiole.
Inflorescence structure does not vary within the
genus (Kostermans 1951). The shape and lobing of
the calyx can be distinctive, especially the welldeveloped lobes before and during anthesis. However,
these lobes soon become invisible as the calyx expands
during the development of the fruit.
The corolla, stamen and ovary characters are
uniform between species and not of much taxonomic
value (Kostermans 1951). One exception is Teijsmanniodendron ahernianum, which usually has five stamens
rather than four.
Fruit characters like woodiness of pericarp, shape and
size, indumentum and glaucousness appear to be useful
characters in species delimitation, although some may be
difficult to recognise by the general user and are therefore of limited value in species recognition. The fruit wall
can vary from woody (Teijsmanniodendron pteropodum) to
leathery (T. coriaceum) (Ng 1992): when dried it can be
smooth, wrinkled to striate or slightly lobed.
Characters derived from hairs and glands on the
leaf surface (Rusea & Latiff 2001) and petiole anatomy
and pollen morphology (Rusea 1998) are to a certain
extent good secondary characters.
THE GENUS TEIJSMANNIODENDRON KOORD. (LAMIACEAE)
Distribution
The natural distribution of the genus is from the Nicobar
Islands (Great Nicobar and Katchal Islands) in India
(Srivastava 1992), through peninsular Thailand and
South Vietnam (Birar), then across Malesia to New
Guinea and the Solomon Islands, with the exception of
Java and the Lesser Sunda Islands. The centre of
diversity is in Borneo where all 23 currently recognised
species are recorded. One species occurs in the south
east of Vietnam (Teijsmanniodendron holophyllum) and
one species (T. pteropodum) is reported as rare on the
Nicobar Islands, India. Three species occur in peninsular
Thailand (T. bogoriense, T. pteropodum, T. coriaceum).
Eight species are recorded from Peninsular Malaysia
(T. bogoriense, T. coriaceum, T. hollrungii, T. holophyllum,
T. pteropodum, T. simplicifolium, T. simplicoides, T. sinclairii).
In Indonesia, Sumatra has eight species (T. ahernianum,
T. bogoriense, T. coriaceum, T. glabrum, T. holophyllum,
T. pteropodum, T. simplicifolium, T. subspicatum); all
specimens of the three species recorded from Java
(T. ahernianum, T. bogoriense, T. pteropodum) are cultivated
plants in Bogor Botanical Garden. Only six species occur
east of Wallace’s Line (T. ahernianum, T. bogoriense,
T. hollrungii, T. holophyllum, T. pteropodum, T. unifoliolatum).
Materials and Methods
This study is based mainly on observations of natural
populations during fieldwork in Peninsular Malaysia,
Sabah and Sarawak (1996 – 1997) by Rusea, and in
Sabah in 2006 and Sarawak in 2007 by de Kok.
Specimens from the following herbaria have also been
examined: A, BISH, BKF, BM, BO, E, K, KEP, L, LAE,
NSW, NY (including photographs of types), SAR, SAN,
SING and UKMB.
In the following descriptions:
i) all measurements and colour descriptions are from
mature material;
ii) for compound leaves, the measurements given
relate to the central leaflet;
iii) all collections cited have been seen by the authors
unless indicated otherwise;
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) for information on surface glands see Rusea &
Latiff (2001);
vi) in the fruit description a pericarp is considered
thin if it less then 0.1 mm thick.
Taxonomic Treatment
Teijsmanniodendron Koord. (1904: 19 – 31, Figs. 2 & 3);
King & Gamble (1909: 841 – 857); Hallier f. (1918: 47
589
& 50 – 55); H. J. Lam (1919: 98 – 100); H. J. Lam & van
den Brink Bakhuizen (1921: 29 – 31); Merrill (1921:
514; 1923: 394 & 398 – 399; 1929: 262); Ridley (1923:
630 – 633); Kostermans (1951: 75 – 106); Kochummen
(1978: 298); Moldenke (1980a: 460 – 494; 1980b: 18 –
43); Wiselius & Sosef (1998: 548 – 550). Type species:
Teijsmanniodendron bogoriense Koord.
Xerocarpa H. J. Lam (1919: 98). Type species: Xerocarpa
avicenniaefoliola H. J. Lam (1919: 99) = Teijsmanniodendron ahernianum (Merr.) Bakh.
Trees up to 50 m tall, DBH up to 120 cm, with or without
buttresses. Bark smooth to somewhat scaly or flaky.
Twigs usually terete to tetragonal, glabrous to variously
pubescent. Leaves with petioles thickened or swollen at
both ends; decussate, simple or palmately compound
with 3 – 7 leaflets, when compound middle leaflets
largest, others gradually smaller, margin entire. Inflorescences paniculate with trichotomous cymes, terminal or
axillary; bracts leaf-like; bracteoles ovate, lanceolate to
linear, usually < 1 mm; calyx persistent, usually enlarged
in fruit, 5-lobed; corolla (4 –) 5-lobed, 2-lipped, upper
lip (1 –) 2-lobed, lower lip 3-lobed, middle lobe the
largest, the throat villous inside, shortly tubular; stamens
4 (− 5), distinctly didynamous, exserted, inserted in the
middle or lower part of the corolla tube; anthers
biloculed, dorsifixed, dehiscing by long slits; style
terminal, slender, with bifid stigma; ovary imperfectly
2-celled, each cell 2-ovuled; only one ovule developing,
the other 3 are suppressed; ovules hemianatropous,
attached laterally near the apex of the ovary cells. Fruits
drupaceous, dry, the exocarp either thick with scattered
sclerenchymatic cells or thin and very brittle. Seeds one,
pendulous, oblong, with membranous testa. Seedling
with epigeal germination; cotyledons emergent, leafy;
hypocotyl elongated.
HABITAT. Teijsmanniodendron is found growing in
various kinds of vegetation from the landward edge
of mangroves to submontane forest, but most species
are found in primary or secondary lowland rainforest.
In primary lowland rainforest they tend to be more
common in areas with slightly more disturbance such
as on ridges or near streams.
Most species seem not to be very common and
consequently the numbers of collections are low given
their extensive distribution. This contrasts with the
closely related genus Vitex, with which it shares many
ecological characteristics and which is relatively well
represented in collections.
USES. The wood of Teijsmanniodendron is used for
general purposes such as house construction (rafters
and posts), interior work, salt-water piling, telephone
poles, framing, moulding, and for making boxes and
crates. The fruits of T. pteropodum have been used
medicinally, both internally and externally for intestinal complaints.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
590
KEW BULLETIN VOL. 64(4)
Key to Species
1. Leaves 3 – 7-foliolate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Leaves unifoliolate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
2. Petioles winged or with pseudo-stipules at base; fruits woody (0.8 – 4.2 × 0.8 – 5 cm); pericarp woody and thick
(2 – 3 mm) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Petioles not winged nor with pseudo-stipules at base; fruits woody, leathery or fleshy, (1 – 8 × 0.5 – 5 cm);
pericarp thick (2 – 3 mm) or thin (< 0.1 mm) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3. Petioles usually 2 – 7.5 cm long, not winged but with oblong shaped oblique pseudo-stipules at base . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6. T. glabrum
Petioles usually > 5 cm long, winged or with round and straight pseudo-stipules at base . . . . 12. T. pteropodum
4. Petioles < 2 cm long; leaflets narrowly lanceolate (< 3 cm wide), apex narrowly acuminate . . . . 14. T. renageorgeae
Petioles > 2 cm long; leaflets (ob)lanceolate to elliptic (> 2 cm wide), apex round to acuminate . . . . . . . . . . . 5
5. Clear ridge between the petioles (interpetiolar ridge) present; fruits smooth or only slightly striate, large
(mature fruits ≥ 2 cm long and ≥ 2 cm wide); pericarp woody and thick (2 – 3 mm) . . . . . . . . . 3. T. bogoriense
Clear ridge between the petioles (interpetiolar ridge) absent; fruits smooth or striate, leathery and small
(mature fruits ≤ 2 cm long and ≤ 1.5 cm wide); pericarp thin (< 0.1 mm) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6. Leaflets (3 –) 5 – 7, shiny above when dried; stamens (4 –) 5; fruits 1.5 – 2 × 1 – 1.5 mm, globose, apex round,
smooth when dried . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. T. ahernianum
Leaflets 3, not shiny when dried; stamens 4; fruits 1 – 1.5 × 0.5 – 1.1 mm, ellipsoid, apex truncate or depressed,
surface striate when dried . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7. Mature leaves glabrous, rarely with some soft, < 0.1 mm long, hairs . . . . . . . . . . . . . . . . . . . . . . . 5. T. coriaceum
Mature leaves sparsely hairy; hairs hispid and > 2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Leaflets not bullate. West to central Sarawak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. T. bintuluense
Leaflets strongly bullate. East to central Sarawak, Brunei . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. T. bullatum
9. Lower leaf surface punctate with tiny holes (10× lens) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Lower leaf surface not punctate with tiny holes (10× lens) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
10. Young leaves smooth below; lateral veins > 15 pairs; mature fruits with a powdery surface . . . . . . 9. T. hollrungii
Young leaves (slightly) rough below, lateral veins < 15 pairs; mature fruits hispid or with a powdery surface . . . . . 11
11. Leaves oblanceolate, young leaves green; fruits hispid and glaucous but not powdery . . . . . . 11. T. obscurinerve
Leaves ovate to lanceolate, young leaves coppery reddish; fruits with a powdery surface and only a few hairs at
apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13. T. punctatum
12. Young twigs, inflorescences and leaf veins covered with hispid hairs . . . . . . . . . . . . . . . . . . . 16. T. scaberrimum
Young twigs, inflorescences and leaf veins glabrous or covered with soft hairs . . . . . . . . . . . . . . . . . . . . . . . . 13
13. The nodes of young twigs significantly more hairy than young petioles or stems . . . . . . . . . . . . . . . . . . . . . . 14
The nodes of young twigs glabrous or similarly hairy as young petioles or stems . . . . . . . . . . . . . . . . . . . . . . . . 15
14. Leaves lanceolate, < 22 cm long; petioles with short and long (0.5 – 2 mm) yellowish hairs; ovary hairy at apex
only . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18. T. simplicioides
Leaves oblong, > 20 cm long; petiole with short (< 1 mm) white to brownish hairs, whole ovary hairy . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10. T. latiffii
15. Young twigs square in cross-section . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Young twigs round in cross-section . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
16. Young twigs hairy; petioles 2 – 6 cm long; fruits not covered with glaucous layer . . . . . . . . . . . . 8. T. holophyllum
Young twigs glabrous; petioles < 3.5 cm long; fruits sometimes covered with glaucous layer . . . . . . . . . . . . . . . . 17
17. Lateral veins the same colour as the green/brown background in both dried and fresh material; fruits covered
with glaucous layer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22. T. unifoliolatum
Lateral veins usually distinctly white on a green/brown background in both dried and fresh material; fruits
sometimes covered with glaucous layer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15. T. sarawakanum
18. Mature petioles ≤ 1 cm long; leaf blade thick coriaceous. Only found on ultrabasic substrates in North Central
Sabah . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23. T. zainudinii
Mature petioles, ≥ (0.8) – 1 cm long; leaf blade chartaceous to coriaceous. Widespread in many types of
habitat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
19. Leaves with at least one pair of major lateral veins originating at base of leaf, all major lateral veins uniformly
prominent and arching towards to apex (as in the genus Smilax) . . . . . . . . . . . . . . . . . . . . . 20. T. smilacifolium
Leaves without pairs of major lateral veins originating at base, all major lateral veins not uniformly prominent
and not arching towards apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
THE GENUS TEIJSMANNIODENDRON KOORD. (LAMIACEAE)
591
20. Fruit apex depressed and with few hairs; young leaves rough below . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
Fruit apex round and glabrous; young leaves smooth below . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
21. Leaves lanceolate, 8 – 38 cm long, sub-bullate in dried specimens, lateral veins 7 – 10 pairs; petiole pulvinus
sometimes with a distinct black ring; fruit glabrous with few hairs at apex , smooth . . . . . . . . . 21. T. subspicatum
Leaves elliptic, 20 – 27 cm long, strongly bullate in dried specimens, lateral veins (5 –) 8 – 20 pairs; petiole
pulvinus usually with a distinct black ring; fruit glabrous, smooth with faint longitudinal groove . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19. T. sinclairii
22. Leaves elliptic, reddish in dried specimens, lateral veins 4 – 8 pairs; fruits smooth but covered with glands laying
on the surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. T. havilandii
Leaves lanceolate, light-brown in dried specimens, lateral veins ≤ 4 pairs; fruits smooth with fine longitudinal
ridges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17. T. simplicifolium
1. Teijsmanniodendron ahernianum (Merr.) Bakh.
(1935: 74); Kostermans (1951: 84 – 88); Moldenke
(1980a: 467 – 471). Type: Philippines, Luzon, Principe
Prov., Baler, Aug. 1902, Merrill 1007 (holotype PNH;
isotype K!).
Vitex aherniana Merr. (1904: 18); H. J. Lam (1919: 206);
Merrill (1923: 394).
Xerocarpa avicenniaefoliola H. J. Lam (1919: 98 – 100);
Kostermans (1951: 84). Syntypes: Papua New
Guinea, Etappenberg, 25 Oct. 1912, Ledermann
9510 (syntype L!); Papua New Guinea, near April
R., 14 Nov. 1912, Ledermann 9667 (syntype L!);
Papua New Guinea, Camp 18, near April R., 20 Nov.
1912, Ledermann 9789 (syntype L!); Papua New
Guinea, Camp 18, near April R., 21 Nov. 1912,
Ledermann 9792 (syntype L!); Papua New Guinea,
Camp Malu, 4 Jan. 1913, Ledermann 10427 (syntypes
K!, L!); Papua New Guinea, Camp Malu, 4 Feb.
1913, Ledermann 10828 (syntype L!).
Vitex curranii H. J. Lam (1919: 207 – 208); Kostermans
(1951: 84). Type: Philippines, Negros, Oct. 1909,
Curran 17463 (holotype PNH; isotype L!).
Vitex bogoriensis H. J. Lam in H. J. Lam & van den Brink
Bakhuizen (1921: 60) [non Teijsmanniodendron bogoriense Koord.]; Kostermans (1951: 84). Type: Indonesia, Botanic Gardens Buitenzorg [Bogor],
originally from Banka, Anonymous XI. H. 37 (holotype BO!, photo. K), nom. invalid.
Vitex bankae H. J. Lam in H. J. Lam & Bakhuizen van
den Brink (1921: 62 – 63); Kostermans (1951: 84).
Type: Indonesia, Banka, Blinju, 17 Oct. 1914,
Grashoff 36 (syntype BO!, photos, K, L!); Indonesia,
Banka, Djebus, Teysmann s.n. (syntype L!); Indonesia, Banka, Menumbing, Teysmann s.n. (syntype L!);
Indonesia, Banka, Blinju, Teysmann s.n. (syntype
BO!, photos K, L!).
Vitex bulusanensis Elmer (1939: 3798); de Kok (2008:
36). Type: Philippines, Luzon, Sorsogon Prov.,
Irosin (Mt Bulusan), Aug. 1916, Elmer 17004
(holotype PNH; isotypes BM!, NY!).
Tree 9 – 50 m tall, DBH 10 – 60 cm; buttresses < 2 m
high, thin. Bark smooth, fissured or with small flakes,
grey to reddish-black; slash light yellow to pale-brown;
sapwood white to brown; heartwood very hard, blackish-brown. Twigs terete, pubescent when young, glabrescent when mature, without a ridge between the
opposite petioles, greyish. Leaves 3 – 7-foliolate; leaflets obovate, (7 −) 25 – 35 × (2 –) 8 – 13 cm (lateral
leaflets 5 – 15 × 2 – 5 cm), apex acute, base cuneate,
margin flat, coriaceous, glabrescent (especially on
midrib) to glabrous on both surfaces, shining above;
midrib raised on both surfaces; lateral vein pairs 8 –
15, slightly looping near margin; reticulation obscure
or invisible. Petioles terete, 6 – 12 cm long, tomentose,
especially basally and distally; petiolules 0.5 – 5 cm
long, equal in all leaflets, furrowed above, tomentose
at the base. Inflorescences 15 – 30 cm long; peduncles
tomentose, 5 – 12 cm long; bracteoles ovate, < 1 mm
long. persistent. Flowering calyx infundibular, 2.5 – 3 ×
2 – 5 mm, outer surface sericeous when young,
glabrescent when mature; lobes 0 – 5, if present up
to 0.5 mm long, apex acute to round; fruiting calyx
cupuliform, 5 – 8 × 5 mm, margin entire, outer
surface glabrous. Corolla (4 −) 5-lobed, sericeous,
glabrous at the base, white to yellow or red, with a
faint sweet smell; central lobe of lower lip spathulate,
3 – 8 × 5 – 6 mm, apex round to emarginate, margin
serrate and undulate, throat villous; other lobes
isomorphic 3 – 5 × 1.5 – 2 mm, apex round, hairy
outside; tube 2 – 3 mm long, glabrous. Stamens (4 –) 5,
2 – 4 mm long, attached to the upper to lower part of
the tube, white; anthers blue or black. Ovary globose,
1 – 1.5 mm diam., glabrous; style 3 – 4 mm long,
exserted; stigma white or blue. Fruits globose, 1.5 – 2 ×
1 – 1.5 cm, apex round, smooth, glabrous, purple to
(blue) black; pericarp thin.
DISTRIBUTION. Malaysia: Sarawak and probably Sabah.
Indonesia: Sumatra, Moluccas and West New Guinea and
probably Kalimantan and Sulawesi. Brunei, Philippines,
Papua New Guinea and the Solomon Islands. Map 1.
SELECTED SPECIMENS EXAMINED. MALAYSIA. Sarawak:
Bako National Park, 30 Aug. 1977, Bernard Lee S 39411
(K, KEP, L, SAR); Bintulu, Semilajau Forest Reserve,
28 Sept. 1972 Ilias Paie S 32062 (SAR). BRUNEI. Bt.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
592
KEW BULLETIN VOL. 64(4)
Map 1. Distribution of Teijsmanniodendron ahernianum.
Pasir Puteh, 20 Jan. 1959, Ladi ak Bikas BRUN 5110 (K,
KEP, L, SAN). INDONESIA. Sumatra, Bangka, 15 Sept.
1949, Anta (exp. Kostermans) 706 (K, L); Sumatra,
WNW of Medan, Sikundur Forest Reserve, 3 Aug.
1979, de Wilde & de Wilde-Duyfjes 19274 (L); Maluku,
Obi Island, 16 Oct. 1953, de Haan 1829 (BM, K); Irian
Jaya, Sukarnapura, 7 Aug. 1966, Kostermans & Soegeng
234 (K); Biak Island, 12 Sept. 1966, Kostermans &
Soegeng 927 (K). PHILIPPINES. Leyte Naval, 8 Aug. 1992,
Barbon PPI 8490 (K); Luzon, Prov. Zambales, April
1903, Merrill 1762 (K). PAPUA NEW GUINEA. Koneipa
Village, Kipu, Morobe Distr., 7 Jan. 1966, Streimann &
Kairo 26131 (BO, K, L); Gulf Province, Avi Avi R., 30
Oct. 1996, Takeuchi & Kulang 11376 (K); Fly R.,
Palmer R., June 1936, Brass 6984 (K); Bougainville,
Tonolei Harbour, 22 Aug. 1969, Foreman 45678 (K).
SOLOMON ISLANDS. New Georgia Group, Baga Island,
9 Jan. 1963, Whitmore 1311 (K); Wagina Island, 16
March 1964, Whitmore collectors 5431 (K).
HABITAT. Commonly found growing in primary and
secondary (submontane) forest, sometimes in heath
forest or swamps on clay-loam to white sand, sometimes
over granite, (ultra)-mafic or iron-ore rich soils; 0 to
1000 m.
CONSERVATION STATUS. Least concern (LC).
PHENOLOGY. Flowering all year around but mainly from
August to April; fruiting from February to August.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
VERNACULAR NAMES. Malaysia: Sarawak: entabuluh
(Iban language). Philippines: Kulipapan, Salipapa or
Dangula (Tagalog language). Indonesia: Bangka: Kayu
Melak (Malay language); Maluku, Obi island: Tété;
Irian Jaya: Oproere (Biak language), Gerèng (Moejoe
Language). Papua New Guinea: East Sepik Province:
Me Mape; Koropuitasona (Middle Waria Language);
Neviai (Waskuk Language); Miap (Wagu Language);
Bougainville: Kona or Kanara.
USES. The strong timber is used in house construction
and as tool handles in Papua New Guinea.
2. Teijsmanniodendron bintuluense Moldenke (1973:
355 – 356 & 1980a: 471 – 472). Type: Malaysia,
Sarawak, 4th Division, Bintulu, Segan Forest Reserve,
17 Sept. 1972, Chai S 31713, (holotype SAR!; isotypes
K!, L!).
Tree 2.5 – 8.5 m, DBH 10 – 14 cm. Bark smooth, light
to greyish-brown. Twigs tetragonal, densely tomentose
when young, without a ridge between the opposite
petioles. Leaves 3-foliolate; leaflets narrowly elliptic,
12 – 30 × 5 – 8 cm, apex acuminate, base acute,
margin straight, coriaceous, feels rough, not bullate,
glabrous above, stiff-pubescent beneath; midrib prominent and raised beneath; lateral veins 6 – 8 pairs;
THE GENUS TEIJSMANNIODENDRON KOORD. (LAMIACEAE)
593
intercostal veins reticulate, distinct on both surfaces
and slightly raised beneath. Petioles 5 – 7 cm long, halfterete, densely tomentose; petiolules 2 – 3 cm long,
densely tomentose. Inflorescences < 10 cm long, densely
tomentose throughout, (dark) purplish (blue); bracteoles narrowly lanceolate, 0.5 – 0.7 cm, persistent.
Flowering calyx 5 – 5.5 × 1.5 – 2 mm, densely tomentose
and glandular when young; lobes 2 – 3 mm long, apex
acute to acuminate; fruiting calyx 8 – 10 mm diam.,
cupuliform, accrescent; lobes prominently erect, glabrous. Corolla purple to pale blue, densely tomentose;
central lobe of lower lip 3 – 3.5 × 2 – 3 mm,
spathulate, margin round and entire; other lobes
1.5 – 2 × 1.5 – 2 mm, apex acute; tube 3.5 – 4 mm
long, glabrous at base. Stamens 3 – 4 mm long, inserted
in the lower part of the corolla tube, slightly villous.
Ovary c. 1 mm diam., globose, apex glabrous to
sparsely hairy; style 4 – 5.5 mm long, light purple;
c. 0.5 mm long, apex acute. Fruits globose to ellipsoid,
1 – 2 × 0.5 – 1 cm, apex truncated, leathery,
longitudinally striate, glabrous, bloom sometimes
present, orange to dark purple; pericarp thin.
S 47024 (K, L); Kuching, Santubong, 1958, Zehnder S
9571 (K, L, SAR); 7st division, Ulu Sungai Belaga, 31
Oct. 1981, Othman et al. S 43481 (K, L, SAR); Bintulu,
Balingian, Ulu Arip, 31 July 1965, Sibat Luang S 23670
(K, KEP, L, SAR); Ulu Batang Belaga, 29 Oct. 1981,
Hansen 859 (SAR); 3th division, Nanga Tunok, Melinau Community, 3 Aug. 1967, Burt & Martin 4764 (E).
HABITAT. Found growing in primary dipterocap or
heath forest, often on hills or ridges. Soil: yellow sandy
clay or sandstone; c. 10 – 250 m.
CONSERVATION STATUS. Least concern (LC).
PHENOLOGY. Flowering from September to October;
fruiting from May to November.
ETYMOLOGY. Named after the type locality Bintulu in
Sarawak.
VERNACULAR NAME. Sarawak: Entabuluh (Iban language).
NOTE. This species is morphologically closely related
to Teijsmanniodendron bullatum, from which it differs by
not having bullate leaves and a different distribution
(East to Central Sarawak and Brunei rather than West
to Central Sarawak).
DISTRIBUTION. Malaysia: endemic to the western part
of Sarawak. Map 2.
SPECIMENS EXAMINED. MALAYSIA. Sarawak: 1st Div.,
Bako National Park, Path to Teluk Paku, 10 Nov. 1963,
Paul Chai S 18014 (K, KEP, L, SAR); Kapit, Bukit Raya,
22 Oct. 1965, Wright S 23975 (K, L, SAR); Kapit, Bukit.
Raya, 19 Oct. 1965, Wright S 23951 (K, L, SAR); 1st
Division, Bako, 13 Sept. 1984, Dayang Awa & Ilias Paie
3. Teijsmanniodendron bogoriense Koord. (1904: 20 –
21 & Fig. 2); H. J. Lam (1919: 97); Bakhuizen van den
Brink in H. J. Lam & Bakhuizen van den Brink (1921:
29); Merrill (1921: 154 & 512); Kostermans (1951:
88 – 92); Kochummen (1978: 298); Anderson (1980:
344). Type: Indonesia, Bogor Botanic Gardens, section
XI.G.82, Nov. 1949, Kostermans s.n. (neotype L!,
selected here; isoneotypes BO!, K!).
Map 2. Distribution of Teijsmanniodendron bintuluense.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
594
Vitex longifolia Merr. (1910: 227); Teijsmanniodendron
longifolia (Merr.) Merr. (1923: 398); de Kok (2008:
36). Type: Philippines, Mindanao, Surigao Prov., 5
July 1907, Hutchinson 7574 (holotype PNH; isotypes
K!, US!).
Vitex flabelliflora Hallier f. (1918: 50); Kostermans
(1951: 88). Type: Indonesia, East Kalimantan, O.
Borneo, Sungei Talut, Penihier, 1896 – 1897, Jaheri
(in Exp. Nieuwenhuis) 1539 (holotype L!; isotypes
BO!, photo. K!).
Vitex lasiantha Hallier f. (1918: 50); H. J. Lam (1919:
201); de Kok (2007: 600). Type: Papua New Guinea,
SE Neuguinea, im Walde an der Etnabai, 2 Dec.
1904, Koch 45 (holotype BO!; isotype L!), synon. nov.
Vitex merrillii H. J. Lam (1919: 212 – 213); H. J. Lam in
H. J. Lam & Bakhuizen van den Brink (1921: 58);
Vitex euphlebia Merr. ex H. J. Lam (1919: 212), in
syn.; de Kok (2008: 36). Type: Philippines, Mindanao, Butuan Province, Aug. 1912, Fénix 15906
(holotype PNH; isotypes BM!, K!, L!, NSW!).
Teijsmanniodendron bogoriense var. pentaphyllum Moldenke
(1967b: 400). Type: Malaysia, Sabah, Sandakan,
Sepilok Forest Reserve, 5 Aug. 1948, Cuadra A 877
(holotype SAN; isotype K!), synon. nov.
Teijsmanniodendron kostermansii Moldenke (1952: 57).
Type: Indonesia, South Kalimantan, Southern Borneo, Puruktjahü Subdiv., near Matara Djaän, 20
Oct. 1926, Lot Obi 75 [bb. 10495] (holotype BO!,
photo. K!); isotype BO!, photos K, L!), synon. nov.
Teijsmanniodendron pendulum Kosterm. (1960: 352 –
353 & 369); Moldenke (1980b: 19 – 20). Type:
Indonesia, Kalimantan, West Kutei, Palimasan, near
Tabang, 13 Sept. 1956, Kostermans 13007 (holotype
BO!; isotypes BM!, BO!, K!, L!, SING!), synon. nov.
Teijsmanniodendron borneënse Moldenke (1980a: 479 –
480). Type: Indonesia, Kalimantan, Balikpapan
peak (Genung Beratus), 16 July 1953, Kostermans
7555 (holotype NY, scan seen), synon. nov.
Tree 15 – 45 m high, bole 2.5 – 20 m high, DBH 20 –
145 cm; buttresses 0 – 2 m high. Bark smooth to scaly,
lenticellate, greyish-brown to light-brown/whitish;
slash very hard to brittle, (pale) yellow; sapwood white
to yellowish. Twigs glabrous to glabrescent, lenticellate,
with distinct transverse stipular line between leaves.
Leaves (3 –) 5 (– 7)-foliolate; leaflets lanceolate, 9 –
25 × 2 – 11 cm (lateral leaflets 5 – 15 × 1.5 – 7 cm),
apex broadly acuminate, base acute to cuneate,
margin slightly recurved, coriaceous to chartaceous,
glabrous on both surfaces; midrib prominent, raised
on both surfaces; lateral veins 3 – 12 pairs, looping
into an imperfect intramarginal vein near margin;
intercostal veins reticulate, prominent on both surfaces. Petioles subterete, 1 – 6 cm long, glabrous;
petiolule 0.3 – 3 cm long, subterete, sparsely pubescent to glabrescent. Inflorescences < 30 cm long,
pubescent when young, glabrescent when mature;
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
KEW BULLETIN VOL. 64(4)
bracteoles caducous. Flowering calyx irregularly campanulate, 2 – 25 × 25 – 35 mm; lobes 5, pubescent at
base, glabrescent near margin, violet; fruiting calyx
shallowly cupuliform, 3.0 × 1.0 cm, margin entire,
glabrous. Corolla 4 – 5-lobed, white to blue, lobes
pubescent, fragrant; central lobe of lower lip spathulate, 0.3 – 1.0 × 0.3 – 0.5 cm, violet with yellow spot at
base, densely villous; side lobe of lower lip 1.5 – 3.5 ×
3 mm, apex acute; lobes of upper lip 1 – 3.5 × 3 mm,
apex acute; tube 0.5 – 2 mm long, white. Stamens 4,
3.5 – 6 mm long, didynamous, exserted, inserted in
the middle of the corolla-tube, glabrous to hairy at
base, anthers deep purple. Ovary globose, glabrous,
apex densely hairy and glandular; style 4 – 7 mm long,
stigma apex acute. Fruits globose, 2 – 8 × 2 – 5 cm,
apex round, smooth, glabrous, shiny, reddish-purple
to black; pericarp woody, thick (2 – 3 mm).
DISTRIBUTION. Southern Thailand to the Philippines
and south to Papua New Guinea (except Java and
Lesser Sunda Islands and, apparently, Peninsular
Malaysia). Map 3.
SELECTED SPECIMENS EXAMINED. MALAYSIA. Sabah:
Tawau, Impong Road, 13 March 1978, Fedillis &
Sumbing SAN 88324 (K, KEP, SAN, SAR); Tawau, 10
acres plot Lua Song, Kalabalakan, 25 Sept. 1967,
Binideh SAN 59762 (SAN); Kota Belud, Sungai Telupid, 19 Sept. 1972, Shea & Minjulu SAN 76173 (SAN);
Lahad Datu, Forest Reserve, Ulu Segama, 11 Aug.
1986, Donggop SAN 116870 (SAN); Sandakan, Mostyn,
Madai Forest Reserve, 5 Aug. 1966, Sinanggul SAN
57074 (SAN); Sandakan, Mile 60, 23 Aug. 1963,
Sinanggul SAN 39106 (KEP, SAN); Sarawak, Belaga,
Plieran Rapids, 8 May 1994, Lai et al. S 67959 (K, KEP,
L); Mt Kalulong, 13 Feb. 1956, Pickles 3731 (BM).
INDONESIA. Sumatra: Sikundur Forest Reserve, 3 Aug.
1979, de Wilde & de Wilde-Duyfjes 19274 (K); Gunung
Bandahara, 20 March 1975, de Wilde & de Wilde-Duyfjes
15531 (K, L); Kalimantan: East Kalimantan, East Kutei,
Sangkulirang, Sungai Menubar, 12 June 1951, Kostermans 5166 (BO, L); Balikpapan, Peak of G. Beratus, 11
July 1952, Kostermans 7449 (L); West Kalimantan, West
Boneo, Berau, 29 May 1934, Neth. Ind. For. Service bb.
19044 (BO, L); East Kalimantan, Pujungan Distr.,
Kayan-Mentarang Nat. Res., 29 June 1992, MacDonald
& Ismail 3491 (L); Borneo, West koers, Djembajan
(Kelesan), 27 June 1938, Neth. Ind. For. Service bb 25135
(L); Borneo, Sampit, 15 Nov. 1929, bb 13944 (L);
Sulawesi Celebes, Pangkawjene, Teysmann 11785 (K,
L); Sulawesi: Malili, 11 March 1933, Neth. Ind. For.
Service V-244 (L); Sulawesi Utara, Sungai Papayuto, 29
March 1990, Burley et al. 4175 (E); Maluku: Moluccas,
Ambon, 16 Dec. 1929, bb 14274 (L); Irian Jaya [West
Papua]: Hollandia Res., Tami, 21 Jan. 1955, Schram BW
1653 (K); Ransiki, 16 July 1948, Ilham 6 [bb. 33255] (K).
THAILAND. Narathiwat: Kaluwotai, Khao Sannak, 23
April 1986, Niyomdham et al. 1207 (BKF, K); Kaluwotai,
THE GENUS TEIJSMANNIODENDRON KOORD. (LAMIACEAE)
595
Map 3. Distribution of Teijsmanniodendron bogoriense.
Khao Chana, 19 Sept. 1985, Niyomdham et al. 1044
(BKF). PHILIPPINES. Mindanao: Butuan, Sept. – Oct.,
1913, Rafael 20752 (K). PAPUA NEW GUINEA. West
Sepik Distr., Imonda Patrol Post, 25 Nov. 1971,
Streimann & Martin 52871 (K); Western Distr., Kiunga,
6 Aug. 1971, Streimann 51713 (K); Madang Distr.,
Koropa, 8 Aug. 1955, Hoogland 5061 (BM).
HABITAT. Primary and secondary rainforest (occasional in submontane forest), flood plains or swamps
on sand, black or yellow loam, sometimes over basalt,
sandstone or limestone soils; 0 – 700 (– 1200) m.
CONSERVATION STATUS. Least concern (LC).
PHENOLOGY. Flowering and fruiting the whole year
round.
ETYMOLOGY. Named after Hortus Bogoriense in Java,
where the type specimen was grown and collected.
VERNACULAR NAMES. Sabah: Buak-buak (Malay language); Sarawak: Entabuluh or Memaru (Iban language); Pangajen Asai (Kenyah language). Indonesia,
Kalimantan: Gragai or Galigantan (Balikpapan area);
Irian Jaya: Prau (Selogof language); Běsokja (Ransiki
language). Papua New Guinea: Kossijt (Salawati
Island); Gugba (Madang language); Timiri (Bembi
language); Parrapik (Kaigorin Language); Ballebal (Jal
Language); Ludri (Koropa Language).
USES. Its wood is reported to be used for timber,
especially to make planks.
TYPIFICATION. In the original publication by Koorders
(1904), two different trees growing in the Bogor
Botanical Gardens are cited (‘‘colitur in Hortus
Bogoriensi, plot IX.D.78 & IX.D.78a’’). No specimens
with those plot numbers can now be found in BO, L
and K. The illustrations accompanying the original
description unfortunately do not exhibit some of the
key characters. Therefore a new type has to be
selected. A specimen collected from Bogor Botanic
Gardens (section XI.G.82) by Kostermans in Nov.
1949, of which several duplicates exist at BO, K and
L is the best candidate. The one at L with both flowers
and fruits is selected as the new type.
NOTES. Based on numerous observations in the field
by the authors, we are convinced that Teijsmanniodendron borneënse and T. kostermansii are only slight
morphological variants (sometimes easily observed
on a single tree) of T. bogoriense. T. pendulum is a
small-leaved form of T. bogoriense.
4. Teijsmanniodendron bullatum G. Rusea, sp. nov. A
T. bintuluensi foliis valde bullatis nec laevibus et
distributione geographica differt. Typus: Malaysia,
Sarawak, Sibu Division, Mukah, Ulu Kenyana, 21 Oct.
1963, Ashton S 19587 (holotypus SAR!; isotypi K!, L!).
Tree 1 – 12 m tall, DBH 15 – 20 cm. Bark smooth, greybrown. Twigs without a ridge between the opposite
petioles. Leaves 3-foliolate; leaflets elliptic, asymmetric,
8.5 – 24 × 3.5 – 8 cm, lateral leaf base distinctly
oblique, base cuneate, margin recurved, strongly
bullate, shiny, coriaceous, glabrous above, feels rough,
stiff-pubescent beneath, greenish-brown, young leaves
bright yellow-green; midrib prominent, flat to sunken
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
596
above, raised below; lateral veins spaced far apart,
prominent, 5 – 7 pairs, looping near margin; intercostal veins prominent beneath and strongly sunken
above. Petioles 1 – 6 (– 8) cm long, subterete, sparsely
to densely tomentose; petiolule 8 – 13 mm long.
Inflorescences densely tomentose, purple; bracteole
persistent. Flowering calyx 1.5 – 3 × 1.5 – 2 mm,
tomentose, yellow; lobes 0.5 – 1 mm long, apex acute;
fruiting calyx about 6 – 7 mm diam., shallowly
infundibular, glabrous, lobes erect, yellow. Corolla
tomentose, mauve to purple; central lobe of lower lip
oblong, c. 3 × 1 mm, apex round and entire, yellow
blotch in throat, villous inside; other lobes c. 2 ×
1 mm, apex acute; tube c. 2 mm long, glabrous at base.
Stamens 4, 3 – 4 mm long, exserted, inserted in the
lower part of the corolla tube; anthers black. Ovary
globose, c. 1 mm diam., glabrous; styles c. 3 mm long,
stigma c. 0.5 mm long, apex acute. Fruits globose to
ellipsoid, 1 – 1.3 × 0.5 – 1.1 mm, apex truncate,
smooth when fresh, striate when dried, glabrous,
yellow to pink; pericarp thin. Fig. 1.
DISTRIBUTION. Only known from Central and East
Sarawak, Brunei and south-western Sabah. Map 4.
SPECIMENS EXAMINED. MALAYSIA. Sarawak: Miri, NE
Lambir Hills, Tukau formation, 29 Nov. 1962, Au &
Ashton S 16450 (K); Bintulu Division, Simalajau
National Park, Sungai Kebalak, 18 June 1990, Mohtar
& Othman S 59564 (K, SAR); Sabah: Nabawan, Labou
Forest Reserve, 18 May 1992, Fidilis SAN 129935 (K).
BRUNEI. Belait: Andulau Forest Reserve, 22 June 1996,
Ariffin Kalat 17471 (K); Andulau Forest Reserve, 15
July 1957, Ashton 256 (K); Belait, New road to
Merankin from 13 Labi road, 10 Nov. 1990, Kirkup
253 (K, L); Teraja Forest Reserve, 18 Dec 1963, Hotta
12726 (L).Tutong: Bukit Udal, 22 Dec. 1994, Osman
16469 (K); Labi, Bukit Telingan, 2 Nov. 1990, J.
Dransfield et al. 6828 (K, KEP, L); Labi: Sungai
Ramayoh, 12 Jan. 1994, Coode 7827 (K, KEP); Brunei,
19 Nov. 1934, KEP 30468 (K).
HABITAT. Primary and secondary dipterocarp and
heath (kerangas) forest on yellow sandy or sandy clay
soils, sometimes over sandstones; c. 15 – 250 m.
CONSERVATION STATUS. Least concern (LC).
PHENOLOGY. Flowering in June to December; fruiting
in July to January.
VERNACULAR NAMES. Malaysia, Sarawak: Entabuluh
(Iban language); Brunei: Maratubu (Iban language).
NOTES. This species is morphologically closely related
to Teijsmanniodendron bintuluense, from which it differs
by having strongly bullate leaves and a different
distribution (West to Central Sarawak rather than East
to Central Sarawak and Brunei).
5. Teijsmanniodendron coriaceum (C. B. Clarke)
Kosterm. (1951: 80 – 84); Kochummen (1978: 308);
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KEW BULLETIN VOL. 64(4)
Moldenke (1980a: 480 – 484); Anderson (1980: 344).
Basionym: Vitex coriacea C. B. Clarke (1885: 586);
Ridley (1923: 630 – 632). Type: Malay Peninsula,
Malacca, Griffith 6065 (lectotype K!, selected here;
isolectotype SING!).
Vitex venosa H. J. Lam in H. J. Lam & Bakhuizen van
den Brink (1921: 61); Kostermans (1951: 80). Type:
Indonesia, Sumatra, Palembang, Banjuasin and
Kubu countries, 24 Dec. 1915, Grashoff 890 (syntypes BO!, photos K, L!).
Tree 4 – 30 m tall, bole 8 – 20 m high, DBH 6 –
100 cm. Bark smooth, slightly scaly to fissured, grey to
pale-brown or brown; slash dull-blackish or pale
brown; sapwood orange-ochre. Twigs glabrous, without
a ridge between the opposite petioles. Leaves 3 (– 4)foliolate; leaflets oblanceolate, 5 – 18 (– 20) × (2 –)
3 – 5 (– 7) cm, apex acute to obtuse, base acute, base
of lateral leaflets sometimes oblique, margin strongly
recurved, coriaceous, not too strongly bullate, not
shiny above when dried, glabrous on both surfaces,
sometimes hairy beneath in young leaflets; midrib
prominent and raised beneath; lateral veins 5 – 8
pairs, curving and joined near margin to form an
imperfect intramaginal vein, conspicuous above,
prominent and raised beneath; intercostal veins reticulate, distinct and raised beneath. Petioles 2 – 5 cm
long, glabrous to slightly hairy, subterete in crosssection. Inflorescences 4 – 15 cm long, slightly pubescent; bracteoles linear to oblong, 0.2 – 0.4 cm long,
persistent, pubescent. Flowering calyx broadly infundibular, c. 1.5 cm long, sparsely pubescent; lobes 1 –
0.5 mm long, apex acute to obtuse; fruiting calyx
enlarged, shallowly cupuliform, margin entire. Corolla
pubescent, white, violet to dark-violet, slightly fragrant;
central lobe of lower lip spathulate, 4.5 – 5 × 3.5 –
4 mm, apex round, throat villous inside, violet with a
yellow patch at base; side lobes of lower lip c. 2 ×
1.5 mm, apex round; lobes of upper lip c. 1.5 × 2 mm,
apex round to acute; tube 5 – 6 mm long. Stamens 4,
4 – 5 mm long, distinctly didynamous, exserted,
inserted in the lower part of the corolla tube, sparsely
pubescent; anthers globose. Ovary glabrous, creamcoloured; styles purple, long-exserted; stigmas palepurple. Fruits ellipsoid, 1 – 1.5 × 0.5 – 0.8 cm, apex
truncated, leathery, striate, glabrous, deep blue or
black; pericarp thin. Seedlings see Ng (1992).
DISTRIBUTION. From Peninsular Thailand south to
Sumatra and Borneo. Map 5.
SELECTED SPECIMENS EXAMINED. THAILAND. Satun:
Talay Bahn [Thalae Ban] National Park, Kwan Bohn,
2 July 1985, Maxwell 85 – 652 (BKF); Songhkla: Ton
Ngaa Chang, 21 Aug. 1992, Niyomdham & Puudjaa
3084 (BKF); Pattani, Bulait, July 1923, Kerr 7100 (E).
MALAYSIA. Peninsular Malaysia: Maingay 1203 (BM,
K); Pahang, Taman Negara, Bukit Terisik, 31 Aug.
THE GENUS TEIJSMANNIODENDRON KOORD. (LAMIACEAE)
597
Fig. 1. Teijsmanniodendron bullatum. A habit; B petiole detail; C underside leaf detail; D underside leaf closer detail; E calyx, fruit
detail; F fruit, side view. All from Coode 7827. DRAWN BY JULIET BEENTJE.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
598
KEW BULLETIN VOL. 64(4)
Map 4. Distribution of Teijsmanniodendron bullatum (●) and T. obscurinerve (■).
1970, Soepadmo KLU 843 (K, KEP, L, SING); Negeri
Sembilan, Pasoh Forest Reserve, Plot I, Kuala Pilah, 17
July 1976, Mat Asri FRI 25624 (K, KEP, L); Bintang
Hijau Forest Reserve, 27 July 1970, Everett FRI 14490
(K, KEP, L, SING); Sabah: Tawau, Hap Seng logged
Map 5. Distribution of Teijsmanniodendron coriaceum.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
area, mile 3 Brantian, 21 Nov. 1978, Fidilis & Sumbing
SAN 89106 (K); Sandakan, Bidi-Bidi Hills above
Kiabau, 3 June 1964, Meijer SAN 43828 (K, KEP, L,
SAN); Sarawak: Kapit, Bukit Nungsang, Entuloh,
Mengiong, 14 Nov. 1979, Othman S 41397 (K, L,
THE GENUS TEIJSMANNIODENDRON KOORD. (LAMIACEAE)
SAR); 5th Div., Limbang, Ulu Sungei Medamit, 9 Oct.
1972, Othman & Wright S 32258 (K, L, SAR); Bintulu,
Maging, Sungai Banut, 24 Oct. 1985, Ilias Paie S 48610
(K, KEP, SAR); 7th Div., Kapit, Bukit Raya, 23 Oct.
1965, Wright S 23988 (K, L, SAR). BRUNEI. Kuala
Belalong, 21 Sept. 1959, Ashton BRUN 3379 (K).
INDONESIA. Sumatra: Bangka, 27 Oct. 1948, Anta
(exp. Kostermans) 1350 (K); Muara Enim Distr., Tjaban
Forest Reserve, Sept. 1955, Kostermans 12084 (K, L);
Jambi, W of Muarabungo, Jan. 1984, Purnajaya TFB
4866 (L); W Sumatra, Pesisir Selatan Distr., SungailasiSindang, 21 Feb. 1983, Laumonier et al. TFB 4121 (L);
N Sumatra, Central Tapanuli, Bonan Dolok-Barus,
July 1985, Purnadjaja & Setiabudi TFB 5091 (L);
Kalimantan: W of Samarinda, Loa Djanan, 11 April
1952, Kostermans 6388 (BO, K, L); Nunukan island, 3 –
5 Dec. 1953, Kostermans 8924 (BO, K); Kalimantan
Timur, Samboja, 26 Aug. 1992, Ambrisyah & Ariffin AA
564 (K, L); Sampit region, near Kuala Kuajan, 27 July,
1953, Kostermans 7962 (L).
HABITAT. Found growing in primary or secondary
forests on sand, sandy clay or clay, sometimes over
sandstone or granite soils; 100 – 1200 m.
CONSERVATION STATUS. Least concern (LC).
PHENOLOGY. Flowering from July to December; fruiting from March to April.
VERNACULAR NAMES. Peninsular Thailand: Kaza
(Malay language); Peninsular Malaysia: Meroyan Batu
(Malay language). Sarawak: Entabuluh (Iban language)
or Madang. Indonesia, Kalimantan: Mahuwi Kuning or
Ma-uhi Putih (Samarinda), Kaju Gading, Kaju Krasak
(Dayak language).
USE. The heartwood is said to be durable and
comparatively strong.
TYPIFICATION. Two collections were mentioned in the
original description of Vitex coriacea: Griffith 6065 and
Maingay 1203 (Clarke 1885: 586). The Griffith 6065
collection at K is selected here as the lectotype.
6. Teijsmanniodendron glabrum Merr. (1929: 263);
Kostermans (1962a: 166 – 167); Teijsmanniodendron
bogoriensis var. glabrum (Merr.) Bakh. ex Moldenke
(1967a: 23), in synon; Moldenke (1980a: 484 – 486);
Anderson (1980: 344 – 345). Type: Malaysia, Sabah,
Elphinstone Province [Tawau Distr.], near Tawao [Tawau],
Oct. 1922 – March 1923, Elmer 21320 (lectotype K!,
selected here; isolectotypes A!, BO!, BISH!, BM!, L!, NY!).
Teijsmanniodendron pteropodum var. auriculatum Bakh. ex
Kosterm. (1951: 94); Teijsmanniodendron pteropodum
var. auriculata Bakh. ex Moldenke (1959: 354), in
synon.
Teijsmanniodendron pteropodum f. juv. auriculatum (Bakh.
ex Kosterm.) Moldenke (1980b: 27 – 28). Type:
Indonesia, Kalimantan, E. Kutai, 13 June 1928,
Abdulhamid 47 [bb. 12563] (syntype BO!, photo.
K!), synon. nov.
599
Tree, seldom shrub, 4 – 33 m tall, bole < 20 m high, DBH
6 – 16 cm, usually straight; buttresses < 1 m high. Bark
smooth to slightly scaly, whitish, greyish to (greenish)
brown; slash brittle, whitish, yellowish to yellowishbrown; sapwood white to yellow or brownish. Twigs
glabrous, raised ridge between two leaves. Leaves 3foliolate; leaflets oblong or ellipsoid, 9 – 15 × 4 – 7 cm,
apex acuminate with tip slightly to strongly curved, base
sub-equal, margin slightly recurved, slightly shiny above;
midrib prominent, raised on both surfaces; lateral veins
6 – 11 pairs, arching towards the margin, conspicuous
beneath; intercostal veins reticulate. Petioles 2 – 5 (– 7.5)
cm long, terete, glabrous; pseudo-stipular appendages
basal, obtusely oblong; petiolules 1 – 3 cm long.
Inflorescences up to 20 cm long, younger parts sparsely
pubescent, older parts glabrous; bracteoles lanceolate,
< 0.3 cm long, glabrous to glabrescent, caducous.
Flowering calyx 2 – 3 mm long, campanulate, externally
glabrous to glabrescent, whitish-green to purple; lobes 5,
triangular-ovate, c. 0.5 mm long, apex acute to rounded;
fruiting calyx patelliform, lobed, about 1.5 × 0.5 cm,
glabrous. Corolla pubescent, white, fragrant; central lobe
of lower lip 3 – 4.5 × 2 – 3 mm, apex round to truncate;
other lobes 2 – 3 × 1.5 – 2 mm; tube 3 – 5 mm long.
Stamens 4, c. 5 mm long, distinctly didynamous when
mature, exserted, inserted in the lower part of the
corolla tube, villous, white; anthers blue. Ovary glabrous,
slightly villous at apex; styles 3 – 5 mm long, stigmas
blue or violet. Fruits ellipsoid or ovoid-ellipsoid, 0.8 – 3.5
× 0.8 – 4.5 cm, apically obtuse, smooth, glabrous, shiny,
pale-grey to brown; pericarp woody, thick (2 – 3 mm).
DISTRIBUTION. Borneo and north Sumatra. Map 6.
SELECTED SPECIMENS EXAMINED. MALAYSIA. Sabah:
near Tawau, Elmer 21616 (BO, K, L); Sandakan, Mile
57, Telupid Road., Tarmiji & Paul SAN 87629 (SAN,
SING); Sepilok Forest Reserve, Aban Gibot SAN 73773
(K, KEP, L, SAN); Sandakan Bay, Patrick SAN 39713
(KEP, L, SAN); Bintang Baru Logging Area, Tarmiji
SAN 81284 (SAN); Lahad Datu, Mostyn, Sinanggul
SAN 57016 (L, SAN); Sarawak: Miri, Lambir Hills, 9
Nov. 1976, Ilias Paie & Yeo S 38458 (K, SAN, SAR,
SING). BRUNEI. Belait: Ulu Sungei Ingei, 3 Jan. 1989,
Wong s.n. (K). INDONESIA. Sumatra, E side of Gunung
Leuser Natural Park, Sekundur Forest Reserve, 7 Aug.
1991, de Wilde & de Wilde-Duyfjes 21294 (K, L); North
Sumatra, Laras Estate below Pematang Siantar, 7 Feb.
1930, Lörzing 16087 (K, L); Kalimantan: NE Borneo,
Numukan Island, 12 Dec. 1953, Kostermans 9027 (K,
L); E Borneo, Berau, Mt Njapa on Kelai R., 20 Oct.
1963, Kostermans 21397 (K).
HABITAT. Growing in primary and secondary forests
on sandy loam or blackish soil, sometimes over limestone or sandstone; 0 – 120 m.
CONSERVATION STATUS. Least Concern (LC).
PHENOLOGY Flowering from (February) May to June
(November); fruiting from June to October.
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KEW BULLETIN VOL. 64(4)
Map 6. Distribution of Teijsmanniodendron glabrum.
VERNACULAR NAMES. Borneo, Sabah: Salunapid
(Dusun language), Ragas (Orang Sungei language),
Buak-buak (Malay language).
TYPIFICATION. In the original description (Merrill
1929), two different collections are cited (Elmer
21616 (BISH!, BM!, K!, L!, SING!) and Elmer 21320
(BISH!, BM!, BO!, K!, L!, NY!). The specimen Elmer
21320 (K) is at the fruiting stage; since fruits are more
diagnostic in this genus than flowers, the fruiting
specimen is selected here as the lectotype.
7. Teijsmanniodendron havilandii (Ridl.) G. Rusea
comb. nov.
Vitex havilandii Ridl., Bull. Misc. Inform., Kew 1929: 262
(1929). Type: Malaysia, Sarawak, Sadong, Coalmine
Hill, Haviland 861 (lectotype K! K000183029; isolectotypes K! K000183028, SAR!).
Tree 8 – 15 m tall, DBH 10 – 25 cm. Bark smooth, grey;
slash yellowish; sapwood pale yellow. Twigs terete,
glabrous, whitish. Leaves unifoliolate, elliptic, (5 –)
7 – 10 (– 15) × 2 – 5 (– 7) cm, apex acuminate, base
acute to rounded, margin slightly recurved, coriaceous, glabrous, not strongly bullate, mature leaves
rusty-coloured when dry; midrib prominent and raised
beneath; main lateral veins 4 – 8 pairs, not originating
from base, arching towards the margin, prominently
raised beneath, some forming incomplete intramarginal veins; intercostal veins reticulate, obscure beneath.
Petioles 1 – 1.5 cm long, subterete, glabrous, without a
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
distinct black ring. Inflorescences slightly puberulous;
bracteoles narrowly lanceolate, c. 1 mm long, persistent. Flowering calyx 2 – 3 × 1.8 – 3 mm, with a few
hairs, dark reddish; lobes up to 1 mm long, broadly
acute; fruiting calyx cupuliform, 7 – 8 × 7 – 12 mm,
accrescent, glabrous, irregularly lobed. Corolla pubescent, purple to pale-blue; central lobe of lower lip
oblong to orbicular, 4 – 5 × 2.5 – 4 mm, apex round;
side lobes of lower lip 2 – 2.3 mm long, apex acute;
lobes of upper lip 1.5 – 2 mm long, apex round; tube
3.5 – 5 mm long. Stamens 4, 3 – 5 mm long,
didynamous, exserted, inserted in the middle part of
corolla tube. Ovary c. 1 mm diam., apex villous; style
c. 5 mm long, purple; stigma black. Fruit ellipsoid, 12 –
15 × 8 – 9 mm, apex round, smooth, glabrous except
for few hairs at apex, glandular, dark-brown; pericarp
thin.
DISTRIBUTION. Malaysia, Sarawak, Kuching, Sadong
hills and Semengoh Arboretum. Map 7.
SELECTED SPECIMENS EXAMINED. MALAYSIA. Sarawak:
Kuching, Semengoh Arboretum (T.7799 = P.240), 13
Aug. 1976, Ilias Paie S 37038 (K, KEP, L, SAR);
Kuching, Semengoh Arboretum (T.7796 = P.224), 8
Nov. 1974, Ilias Paie S 35766 (K, L); Semengoh
Arboretum, near tree no. 2228, 19 August 1966,
Anderson S 25971 (K, L, SAR); Semengoh Arboretum,
near tree no. 47, 23 June 1960, Anderson S 12574 (L);
Semengoh Arboretum, 21 Aug. 1962, Rosli S 14768 (K,
L); Semengoh Arboretum, Aug. 1961, Rosli S 3358 (L);
Semengoh Arboretum, 12th Mile, Penrissen Rd., 6
THE GENUS TEIJSMANNIODENDRON KOORD. (LAMIACEAE)
601
Map 7. Distribution of Teijsmanniodendron latiffii (●) and T. havilandii (■).
Jan. 1970, Othman Ismawi S 29452 (K, L); Semengoh
Arboretum, 12th Mile, Penrissen Rd., (near tree no.
2241), 9 Nov. 1966, Banyeng ak Nudong & Sibat ak
Luang S 26242 (K, L, SAR); Kuching, 1893, Bartlett s.n.
(BM); Semengoh Arboretum, 12 Sept. 1994, Rantai et
al. S. 67390 (SAR).
HABITAT. Found growing in yellow sandy clay in mixed
dipterocarp forests on ridges; 50 – 100 m.
CONSERVATION STATUS. This species is only known
from a number of collections from two areas, one old
collection from the Sadong hills, and several from the
Semengoh Arboretum (a protected area of approximately 14 ha, surrounded by the larger Semengoh
Nature Reserve). The species is significantly less
common than Teijsmanniondendron simpicifolium, which
forms one of the dominant understorey trees in the
Arboretum. Given the small area of occupancy and
the pressures on the forests in Sarawak and those in
the Kuching area in particular, this species is best
classified as Vulnerable (VU).
PHENOLOGY. Flowering in April to August; fruiting
from August to January.
TYPIFICATION. Two specimens at K are available for
lectotypification (Sadong, Coalmine Hill, Haviland
861, K000183029 and K000183028). As fruits are more
important for species identification in this genus, the
specimen with mature fruits (K000183029) is selected
here as the lectotype.
8. Teijsmanniodendron holophyllum (Baker) Kosterm.
(1951: 97 – 99); Kochummen (1978: 308 – 309);
Moldenke (1980a: 492 – 494); Anderson (1980: 345).
Type: Malaysia, Sabah, Sandakan, April 1895, Governor
Creagh [Hugh Low] s.n. (holotype not located; isotype K!).
Vitex holophylla Baker (1896: 25 – 26); Ridley (1923:
631 – 632).
Teijsmanniodendron novo-guineense (Kaneh. & Hatus.)
Kosterm. (1951: 103); Moldenke (1980b: 18 – 19).
Basionym: Vitex novo-guineënsis Kaneh. & Hatus.
(1942: 116 – 117). Type: Indonesia, Irian Jaya,
Geelvink Bay, Near Nabire, Ayerjat, 9 May 1940,
Kanehira & Hatusima 12578 (holotype not located;
isotype BO!, photo. K), synon. nov.
Teijsmanniodendron pierrei Moldenke (1962: 273 – 274).
Type: Vietnam, Baria, March 1867, Pierre 37 (holotype not located; isotypes A!, K!, NY!), synon. nov.
Teijsmanniodendron subspicatum var. acutifolium Moldenke
(1981: 431). Type: Malaysia, Sabah, Sandakan, Foot
of Mt Mentapok, Kg. Miruru, 4 June 1979, Aban Gibot
& Petrus SAN 90237 (holotype TEX; isotypes K!,
KEP!, L!, SAN!), synon. nov.
Tree 2 – 30 m, bole often straight to 20 m high, DBH
5 – 60 cm; buttresses to 1 m high. Bark smooth to
slightly scaly, whitish, grey to brown; slash greenishyellow to pale brownish. Sapwood white to pale yellow
or yellow. Twigs glabrous to pubescent, square. Leaves
unifoliolate, elliptic-lanceolate or (ob)lanceolate, 12 –
35 × 4 – 7 (– 12) cm, apex acuminate, base acute,
rounded to slightly cordate, margin not to slightly
recurved, slightly bullate, coriaceous, not punctate
beneath, glabrous on both surfaces, but slightly
tomentose when young; midrib prominent and raised
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
602
beneath; lateral veins 5 – 15 pairs, prominent and
raised beneath; intercostal veins reticulate, conspicuous above, distinct and raised beneath. Petioles 1.5 –
6 cm long, (sub)terete, glabrous, distal pulvinus
sometimes with distinct black ring. Inflorescences up to
50 cm long, slightly hirsute; bracteoles lanceolate to
elliptic-lanceolate, caducous, 2 – 5 mm long. Flowering
calyx campanulate, 2 – 2.5 × 1.5 – 2 mm, tomentose and
glandular when young; fruiting calyx campanulate, 12 –
16 mm in diam., glabrous when mature. Corolla deeppurple or white, velvety; central lobe of lower lip 3 – 4 ×
2 – 3 mm, oblong, apex round, throat villous inside;
other lobes 1.5 – 2 × 1.2 – 2 mm, apex acute to round,
pale purple; tube 3 – 4 mm long. Stamens 4, 4 – 5 mm
long, exserted, inserted in the middle or lower part of
the corolla tube; filaments slightly villous; anthers
c. 0.5 mm long, purple. Ovary c. 1 mm diam., villous
at the top; styles 6 – 7 mm long. Fruits subglobose, 2 –
2.2 × 0.9 – 1.2 cm, apex depressed to acute, smooth
with very faint longitudinal stripes, glabrous except few
hairs at apex, dark brown to bluish, shiny; pericarp thin.
DISTRIBUTION. Sumatra, Borneo and Tawi-Tawi Island
in the Philippines and a number of rarely collected
enigmatic populations, one in Johore and one in
Terengganu in Peninsular Malaysia, one each in South
Vietnam and in Irian Jaya, Indonesia. Map 8.
SELECTED SPECIMENS EXAMINED. MALAYSIA. Penisular
Malaysia: Johor, Sungei Kaliang, 1892, Ridley [Lake &
Kelsali] 4031 (BM, K); Terengganu, Dungun, Jerangau
State Land, 5 Sept. 1966, Ismail 98947 (KEP); Sabah:
Map 8. Distribution of Teijsmanniodendron holophyllum.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
KEW BULLETIN VOL. 64(4)
Lahad Datu, Mt Silam, 13 July 1963, Meijer SAN 37924
(K, SAN); Telupid, Hutan Simpan Tangkulap, 23 May
1989, Majawat et al. SAN 118195 (K, SAN); Keningau,
Nabawan, Tiulon, 10 Sept. 1976, Tarmiji & Dewol SAN
83897 (K); Sarawak: Kuching, Padawan, Mini Hydro
Station, Kg. Sadil, 16 Sept. 1987, Yii S 51349 (K, SAR);
Miri, Lambir, Ulu Sungai Lepoh, 19 Oct. 1976, Ilias
Paie & Yeo S 38310 (K, SAR, SING) Bintulu, Nyabau, 7
June 1987, Othman & Munting S 54313 (K, KEP, SAR);
Kuching, Tiang Bekap, Mt Mentawa, 26 July 1963,
Chew Wee Lek 682 (K L, SAR, SING). BRUNEI. Bukit
Puan, 15 May 1957, Ashton 7862 (K); Bank of Sungei
Belait between S. Burau and Sukang, 1 Nov. 1989,
Forman 1129 (K). INDONESIA. Sumatra: West Aceh,
Meulaboh, 27 Feb. 1986, Laumonier YL 6793 (L); Atjeh
Province, Kloet Nat. Res., 12 July 1985, de Wilde & de
Wilde-Duyfjes 20008 (L); Anamba Islands, 15 April 1928,
Henderson SFN 20408 (K, SING); Kalimantan: Kapuas
Ulu, Kp. Embalu, 20 Nov. 1953, Esche bb. 35258 (K); E
Kalimantan, Berau, 19 Sept. 1997, Ambriansyah &
Arbainsyah Berau 587A (K); Central Kalimantan, Sintang,
7 May 1994, Church et al. 1350 (BISH, E, K, SAR); Irian
Jaya [West Papua]: Mamberamo, Nov. 1926, Doct. van
Leeuwen 11260 (K). PHILIPPINES. Tawi-Tawi Island,
Languyan, 27 May 1992, Gaerlan et al. 10122 (K).
HABITAT. Found growing in primary or secondary
forests, on hillsides or ridge-tops, along riverbanks, on
sandy, dark-brown or black soils, sometimes ultra basic;
60 – 265 m.
CONSERVATION STATUS. Least concern in Borneo.
The populations in both Peninsular Malaysia and
THE GENUS TEIJSMANNIODENDRON KOORD. (LAMIACEAE)
Vietnam were only collected once, in the 19th century
and given the pressures on the forests in those
areas; they must be considered at least “Critically
Endangered”. The population in Irian Jaya was only
collected twice in the 19th century and given the
extent of the forests there it must be considered “Data
Deficient”.
PHENOLOGY. Flowering from February to September;
fruiting from June to November.
VERNACULAR NAMES. Malaysia, Sarawak: Entabuluh
(Iban language), Kenjau (Penan language); Sabah:
Kemuning (Lahad Datu area). Brunei: Mertubu or
Martubu (Iban language).
NOTES. Teijsmanniodendron holophyllum is similar to T.
hollrungii in leaf shape and texture, but differs by
having a bullate and non-punctate leaf under-surface.
It is also quite similar to T. subspicatum in its leaf
shape and bullate under-surface but does not have
the reddish-brown drying colour and subspicate
inflorescence.
9. Teijsmanniodendron hollrungii (Warb.) Kosterm.
(1951: 103 – 105); Kochummen (1978: 309); Moldenke
(1980a: 486 – 492); Anderson (1980: 345). Type: Papua
New Guinea, Hatzfeldhafen, 1886, Hollrung 377 (holotype B; isotype BO!, K!).
Vitex hollrungii Warb. (1894: 208).
Vitex clarkeana King & Gamble (1909: 845); Kostermans
(1951: 103). Types: Malaysia, Pangkor Island, Curtis
1611 (syntype K!); Malaysia, Perak, Oct. 1884,
Scortechini 1383 (syntype K!); Malaysia, Telok Sera,
Ridley 7990 (syntype K!); Malaysia, Bernam R.,
King’s Collector 8788 (syntype K!); Malaysia, Malacca,
1845, Griffith 6064 (syntype K!); Malaysia, Johore,
Sembrong R., Lake & Kelsall 4059 (syntype not
located); Borneo, Motley 1269 (syntype not located);
Borneo, Beccari 166 (syntypes K!); Borneo 1853,
Lobb s.n. (syntype K!); Borneo, Haviland 1580
(syntype K!).
Teijsmanniodendron monophyllum Kurata (1947: 203);
Kostermans (1951: 103). Type: Indonesia, Irian
Jaya, Bumi R. near Nabire, 21 April 1939, Inokuma
& Hara 679 (holotype not located).
Vitex simplicifolia C. B. Clarke (1885: 586); Kostermans
(1951: 103). Type: Peninsular Malaysia, Malacca,
Griffith s.n. (holotype not located; isotype K!).
Tree (seldom shrub) 2 – 25 m, bole 2 – 10 m high,
DBH 2 – 45 cm; buttresses up to 0.8 m high. Bark
smooth to slightly flaky, soft, white to whitish-grey;
slash yellowish to pale red or brown; sapwood white or
greyish to yellow; heartwood hard. Twigs stout, terete,
variously pubescent; nodes swollen. Leaves unifoliolate,
lanceolate, 15 – 35 × 5 – 13 cm, apex long acuminate,
base acute to narrowly rounded, margin straight,
coriaceous, above somewhat glossy, feels smooth
603
beneath, glabrous on both surfaces, densely punctuated beneath; midrib slender, flat or slightly impressed
above, slightly rounded and raised beneath; lateral
veins 13 – 20 pairs, arching towards the margin and
sometimes joined; intercostal veins reticulate, distinct
beneath. Petioles slender, up to 0.5 cm long, smooth,
terete. Inflorescences up to 40 cm long; bracteoles linear,
3 – 5 mm long, inconspicuous, puberulent, caducous;
peduncles sharply tetragonal, densely rusty-pubescent.
Flowering calyx campanulate, 3.5 – 4 × 3 – 3.5 mm,
hairy; lobes c. 1 mm long, apex acute; fruiting calyx
much accrescent, 10 – 18 mm diam., shallowly cupuliform, unevenly truncate. Corolla white, cream yellow to
purple, fragrant, velvety; central lobe of lower lip 4 –
5 × 3 – 5 mm, oblong to spathulate, apex rounded,
with few hairs to glabrous above, lilac to purple, yellow
spot at base; side lobes of lower lip 3 – 4 × 2 – 2.5 mm,
apex acute, white; lobes of upper lip 4 – 4.5 × 2 – 2.5,
apex round to acute, white; tube 5 – 7 mm long,
funnel-shaped. Stamens 4, 7 – 10 mm long, distinctly
didynamous, villous at base, pale purple to yellow or
white; anthers c. 1 mm long, white. Ovary globose 1 –
1.5 mm diam.; apex flattened, pubescent; styles to 10 –
15 mm long, purple; stigmas yellowish-white. Fruits
globose to clavate, 1.5 – 2 × 1.5 – 2 cm, apex round,
smooth, glabrous except for few hairs at apex, somewhat fleshy, covered with sooty yellowish or greenishbrown powder which forms a cracked layer when dry,
olive-green to (greyish) black; pericarp thin.
DISTRIBUTION. From Peninsular Malaysia southeast to
Papua New Guinea and the Solomon Islands (except
Sumatra, Java and the Lesser Sunda Islands). Map 9.
SELECTED SPECIMENS EXAMINED. MALAYSIA. Peninsular
Malaysia: Perak, Sungai Krian Estate, 10 July 1938,
Spare 36010 (K); Peninsular Malaysia, Pahang, Endau,
Pianggu, Aug. 1917, Evans s.n. (K); Sabah: Beaufort,
Kampong Bakau, 14 Nov. 1976, Talib Bidin SAN 84578
(K, KEP, SAN); Kinabatangan, Lamag Tenegang
Kechil, 16 June 1965, Eging Banang SAN 51952
(SAN); Papar, Bongawan Forest Reserve, 10 Feb.
1976, Dewol & Talib SAN 80369 (SAN); Lahad Datu,
Paris Camp, Segama, 24 Sept. 1984, Dewol et al. SAN
67532 (KEP, SAN, SAR); Sarawak: Bintulu, Sungei
Segan, 20 Oct. 1963, Anderson S 18582 (K, SAR); Ulu
Lundu, 18 Nov. 1963, Ashton S 18868 (K, SAR).
BRUNEI. Sungai Damit, near Kuala Belait, 6 Nov.
1990, J. Dransfield et al. 6797 (BRUN, K, SAR).
INDONESIA. Kalimantan: near Samarinda, East Kutei,
19 Aug. 1951, Kostermans 6131 (K); Kalimantan
Selatan, Pleihari, 19 Aug. 1965, Sauveni 921 (K);
Bangarmasin, 1857 – 1858, Motley 1269 (K); Pleihari,
19 Aug. 1965, Sauveur 921 (L); Tapin R., 5 km
upstream from Margasari, 12 Dec. 1988, Giesen 55
(L); Celebes [Sulawesi]: Oct. 1913, Rachmat. (Exp. van
Vuuren) 792 (L); Soeda Islands, Mangoli, Atje (Exp. van
Hulstijn) 32 (L); Moluccas [Maluku]: Ambon, July –
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Map 9. Distribution of Teijsmanniodendron hollrungii.
Nov. 1913, Robinson 1867 (BM, K, NSW); Seram Timur,
15 Feb. 1986, Mirmanto & Ruskandi 76 (K); Irian Jaya
[West Papua]: Ransiki, Meos Island, 22 July 1956,
Kalkman BW 3571 (K). PAPUA NEW GUINEA. West
Sepik: Vanimo, Ossima, 27 Jan. 1969, Streimann &
Kairo 39202 (K); New Britain: Cape Hoskins, 6 Aug.
1954, Floyd 6481 (K). SOLOMON ISLANDS. Eastern
Choiseul, 6 March 1964, Whitmore’s Collectors BSIP
5284 (K); Santa Ysabel, 6 Jan. 1964, Whitmore’s Collectors
BSIP 3575 (K).
HABITAT. Found growing in primary or secondary
forests, usually along rivers, tidal creeks, peat swamps
or inland side of mangroves on sand, clay or loam; 0 –
50 (– 100) m.
CONSERVATION STATUS. Least concern (LC).
PHENOLOGY. Flowering all year round; fruiting from
May to March.
VERNACULAR NAMES. Malaysia, Sabah: Renggas
(Kedayan language), Kapur-kapur (Brunei language);
Sarawak: Entabuluh (Iban language), Ladip (Malay
language), Rengas (Kutching), Senumpol (Binatang).
Brunei: Putat (Binuni language). Indonesia: Kalimantan: Lihampit, Kaju Kolok Ampit (Dayak language), Irian
Jaya: Ap (Merauke), Seriga (Salawati Island). Papua
New Guinea: La Pone (W Nakanai, New Britain).
Solomon Islands: Huhuhu, Luluka, Aibu or Aisofolota
(Kwara’ae language).
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
USE. In Sarawak for treating scabies, the ash from the
leaves is added to coconut oil and applied to the head.
NOTE. We have not seen the type of Teijsmanniodendron
monophyllum Kurata, but from the description and the
accompanying plate (Kurata 1947) it is clear that this
name must be placed under T. hollrungii.
10. Teijsmanniodendron latiffii G. Rusea sp. nov.
Teijsmanniodendrono unifoliolato folii forma texturaque
similis, sed ab ambabus inflorescentia densissime
pubescenti, bracteis bracteolisque numerosis persistentibus differt. Typus: Malaysia, Sarawak, 4th Div., Marudi
Bok-Tisam, Sungai Mentagai, 1 May 1965, Sibat ak
Luang S 22819 (holotypus SAR!; isotypi K!, L!, KEP!).
Tree 5 – 15 m tall, DBH 10 – 20 cm. Bark pale brown to
greyish, flaky. Twigs densely pubescent when young,
terete. Leaves unifoliolate, oblong, 20 – 40 × 5 – 10 cm,
apex acuminate, base acute, margin straight to
strongly recurved, glabrous, coriaceous, slightly bullate, shiny above, not punctate beneath, feels slightly
rough beneath, dark to dull green; midrib raised with
a sharp ridge above and beneath; lateral veins 8 – 12
pairs, sunken above, raised beneath; intercostal veins
scalariform. Petioles 1.5 – 4 cm long, subterete, densely
to sparsely hairy. Inflorescences up to 30 cm long,
THE GENUS TEIJSMANNIODENDRON KOORD. (LAMIACEAE)
densely pubescent; bracteoles < 5 mm long, lanceolate, green or violet. Flowering calyx campanulate, 3 –
4 × 1.5 – 2 mm, densely hirsute externally; lobes
c. 1 mm long, apex acute; fruiting calyx infundibular,
much accrescent; lobes 5, broadly connate, apex
acute, densely hirsute externally. Corolla white to violet,
golden pubescent; central lobe of lower lip spathulate,
c. 1.5 × 3 mm, apex round, white to violet with yellow
patch at the base of the lip, throat villous; other lobes
1 – 1.5 × 1 – 1.2 mm, apex acute; tube c. 2.5 mm long.
Stamens 4, 2.5 – 3 mm long, slightly didynamous,
villous at base; anthers black. Ovary globose, c. 1 mm
diam., villous, more densely at apex; styles c. 5 mm
long. Fruits globose, c. 5 × 10 mm, apex round,
sparsely tomentose, more densely at apex, purple.
Fig. 2.
DISTRIBUTION. Malaysia (Sarawak) and Indonesia
(Kalimantan). Map 7.
SPECIMENS EXAMINED. INDONESIA. Kalimantan: Bukit
Raya, Batu Badinging, 7 Feb. 1983, Veldkamp 8584
(KEP, NSW); Kalimantan Tengah, Kotawarigin, 8 Sept.
1996, Ambriansyah & Arbainsyah AA1961 (L); Kotawarigin, Sungai Mentaya, 12 May 1993, Argent et al. 93135
(L).
HABITAT. Growing in primary forest on ridges; 50 –
100 m.
CONSERVATION STATUS. Least concern (LC). A rarely
collected species but with a wide distribution.
PHENOLOGY. Flowering and fruiting from September
to May.
ETYMOLOGY. This species is named after Prof. A.
Latiff, Professor of Botany and Curator of UKMB
Herbarium, Malaysia.
11. Teijsmanniodendron obscurinerve G. Rusea sp.
nov. Teijsmanniodendron hollrungii et T. punctatuto
fructibus hispidis nec glabris neque glaucis differt.
Typus: Malaysia, Sabah, Sandakan, Kuala Sungai Sasau
and Sungai Tongod, Labuk Sugut, 24 Sept. 1984,
Matin Pikkoh SAN 67646 (holotypus SAN!; isotypi K!,
KEP!).
Tree 3 – 15 m tall, bole up to 10 m high, DBH 30 –
40 cm. Bark smooth, whitish; slash yellowish; sapwood
yellow. Twigs smooth, distinctly tetragonal, glabrous.
Leaves unifoliolate; oblanceolate to elliptic, 15 – 28 ×
3 – 10 cm, apex acuminate, base rounded to cordate,
margin thickened and slightly recurved, glossy above,
punctate beneath, coriaceous, glabrous on both surfaces, young leaves feel rough beneath; midrib prominent
and raised on both surfaces, whitish beneath when dry;
lateral veins obscured above, slightly raised beneath, 8 –
14 pairs, arching and looping near the margin forming
an intramarginal vein, intercostal veins reticulate,
obscured on both surfaces. Petioles 1.2 – 2.5 cm long,
605
glabrous, subterete. Inflorescences up to 20 cm long,
sparsely to densely hairy; bracteoles ovate, 2 – 2.2 ×
0.5 – 1 mm, apex acute, caducous. Flowers (purple) blue
to purple; flowering calyx campanulate, 1 – 2 × 1.5 –
2 mm, densely appressed tomentose; lobes deltoid,
c. 0.5 mm long, apex acute. Fruiting calyx accrescent, 5 –
6 mm diam., cupuliform, 5-lobed to truncate. Corolla
funnel-shaped, densely pubescent; central lobe of lower
lip absent, throat villous inside. Stamens 4, distinctly
didynamous, exserted, inserted in the middle part of
the corolla tube; filaments unknown. Ovary apically
obtuse and villous. Fruits ellipsoid, 15 – 20 × 7 – 15 mm,
apex round, smooth, hispid hairs especially at apex,
glaucous, not powdery when dried, dark purple;
pericarp thin (< 0.1 mm). Fig. 3.
DISTRIBUTION. Malaysia, Sabah. Map 4.
SPECIMENS EXAMINED. MALAYSIA. Sabah: Ulu
Nangoh, 20 June 2001, Postar Miun SAN 151108 (K);
Beluran Distr., Meliau, Sungai Sulosong, 13 Aug. 2003,
Diwol & Markus SAN 136546 (KEP, SAN); Ulu Tongod
Forest Reserve, 17 Aug. 2004, Sugau SAN 145509
(KEP); Bukit Masasau, 29 July 1983, Sigin & Rahim
SAN 99689 (K); Nangoh, Bukit Masassau, 29 July 1983,
Sigin et al. SAN 99769 (SAR); Jesselton, Sungai
Sapatali, 12 Dec. 1962, Madani SAN 33190 (K, SAR).
HABITAT. Found growing on river banks; 100 – 150 m.
CONSERVATION STATUS. This species is only known
from five relatively recent collections. As almost
nothing is known about the ecology of this species a
conservation status of “Data Deficient” is proposed.
PHENOLOGY. Flowering from July to September
(December); fruiting from June to August.
VERNACULAR NAMES. Sabah: Buak-buak Pulas (Malay
language).
NOTE. This species is distinctly different from any
known Teijsmanniodendron by its hispid fruits, narrowly
oblong leaves and obscure venation on both surfaces.
We have not seen any mature flowers, but given the
lack of variation in the corolla morphology in the
genus, this is not seen by us as a great problem in
describing this as a new species.
12. Teijsmanniodendron pteropodum (Miq.) Bakh. in
H. J. Lam & Bakhuizen van den Brink (1921: 29);
Anderson (1980: 345); Kostermans (1951: 92 – 95);
Kochummen (1978: 309); Moldenke (1980b: 21 – 27).
Type: Indonesia, Sumatra, Palembang, Dangku Lematang, Teijsmann s.n. (holotype L!; isotypes BO!, K!).
Vitex pteropoda Miq. (1861: 242 & 567); Gamble in King
& Gamble (1909: 851); H. J. Lam (1919: 170);
Ridley (1923: 633).
Vitex philipinensis Merr. (1903: 52); Kostermans (1951:
92). Type: Philippines, Mindanao, Province of
Zamboanga, March 1901, Ahern 387 (holotype
PNH; isotypes BO, US!).
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KEW BULLETIN VOL. 64(4)
Fig. 2. Teijsmanniodendron latiffii. A habit; B leaf underside; C indumentum detail at node; D underside leaf detail; E corolla
opened with ovary and stamens. All from Sibat ak Luang S. 22819. DRAWN BY JULIET BEENTJE.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
THE GENUS TEIJSMANNIODENDRON KOORD. (LAMIACEAE)
607
Fig. 3. Teijsmanniodendron obscurinerve. A habit; B detail of node and petiole base; C underside leaf detail; D inflorescence detail.
All from Matin Pikkoh SAN 67646. DRAWN BY JULIET BEENTJE.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
608
Vitex peralata Miq. (1861: 242 & 567). Type: Indonesia,
Sumatra, Palembang, Dangku Lematang, Teijsmann
s.n. (holotype L; isotypes BO!, K!), synon. nov.
Vitex peralata King in King & Gamble (1908: 112);
Kostermans (1951: 92); Teijsmanniodendron peralata
(King) Balakrishnan (1980: 176). Types: Malaysia,
Perak, at Simpang, Wray 2029 (syntype K); Malaysia
Perak, Wray 2254 (syntype K!); Malaysia, Perak, at
Simpang, July 1888, Wray 2305 (syntype K!); Malaysia, Perak, near Larut, July 1881, King’s Collector
2064 (syntype K!); Malaysia, Perak, near Larut,
1884, Kunstler [King’s Collector] 6187 (syntype K!);
Malaysia, Perak, near Larut, Nov. 1884, King’s
Collector 6874 (syntype K!); Malaysia, Perak, near
Larut, Nov. 1885, King’s Collector 8299 (syntype K!),
nom. invalid.
Vitex koordersii H. J. Lam in H. J. Lam & Bakhuizen van
den Brink (1921: 64); Kostermans (1951: 92).
Types: Indonesia, Sumatra, near Pelembang, Buurman v. Vreeden 158 (syntype BO!, photo. K!);
Indonesia, Central Sumatra, Rawang, Sigati, Koorders 10483 β (syntype BO); Borneo, Jaheri s.n.
(syntype BO!, photo. K).
Teijsmanniodendron pteropodum f. cristatum Moldenke
(1979b: 473). Type: Malaysia, Sabah, Penampang
Distr., Sungai Sosopodon, 16 Dec. 1963, Mikil SAN
37769 (holotype TEX; isotype L!), synon. nov.
Tree 6 – 40 m tall, bole 5 – 20 m high, DBH 12 –
60 cm; buttresses < 4 m high. Bark smooth to flaky,
lenticellate, white to yellow to black; slash yellow to
purplish mottled; sapwood yellow. Twigs glabrous,
pale-brown. Leaves 3 – 7-foliolate; leaflets lanceolate,
obovate to ovate, central leaflets 10 – 55 × 3 – 19 cm,
apex obtuse to acuminate, base attenuate to cuneate,
margin slightly to strongly recurved, coriaceous, shiny
or glossy, bright- or dark-green above, lighter beneath,
glabrous; midrib and lateral veins sunken above,
prominent and raised beneath; lateral veins (5 –)
12 – 16 pairs; intercostal veins reticulate, conspicuous.
Petioles 7 – 30 cm long, triangular in cross section,
glabrous, narrowly to broadly winged, usually throughout the length of the petiole; petiolule 0 – 5 cm long,
broadly grooved above. Inflorescences 20 – 100 cm long,
reddish-brown or pale purplish-red; bracteoles lanceolate to ovate, 2 – 10 × 3 – 7 mm, persistent, apically
abruptly acute, glabrescent on both surfaces, whitish
or pale-greenish to purple. Flowering calyx 2 – 2.5 ×
1.5 – 2 mm, campanulate, sparsely pubescent externally, glabrous internally; lobes 1.5 – 3.5 × 2 – 2.5 mm,
purple to rose-coloured; fruiting calyx hypocrateriform,
1.7 – 5 × 3 – 4.5 cm, greatly spreading, marginally
undulating, glabrous, brownish. Corolla pubescent,
violet to pale-blue violet; central lobe of lower lip
3.5 – 4 × 3.8 – 5 mm, spathulate, apex round, throat
brown-striate; side lobes of lower lip 2 – 2.5 × 2 –
3 mm, apex round, lobes of upper lip 1.5 – 2 × 2 mm,
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
KEW BULLETIN VOL. 64(4)
apex acute, reflexed; tube 4 – 7 mm long. Stamens 4,
distinctly didynamous, exserted, inserted above or at
middle part of the corolla tube; filaments 2 – 6.5 mm
long, white to pale-violet, slightly villous at base;
anthers dark-violet or dark-purple, glabrous. Ovary
globose, apex depressed, villous at apex; style 8 –
10 mm long, exserted, glabrous; stigmas shortly bifid.
Fruits ellipsoid, sometimes lobed when dry, 1.5 – 4.5 ×
2 – 5 cm, smooth, irregularly wrinkled when young,
glabrous, brown; pericarp woody, thick (2 – 3 mm).
Seedlings see Ng (1992).
DISTRIBUTION. India, Nicobar Islands (Srivastava
1992); Southern Thailand to Sumatra, Borneo and
the Philippines. Map 10.
SELECTED SPECIMENS EXAMINED. THAILAND. Narathiwat: Towo, Phlu Kok Gu No, 19 July 1983, Niyomdham
702 (BKF). MALAYSIA. Peninsular Malaysia: Pahang,
Lesong Forest Reserve, 9 June 1979, Kamarudin 28032
(K, KEP, L); Johore, Maokil Forest Reserve, 26 July
1970, Kochummen FRI 16082 (KEP, SAR, SING); Sabah:
Lahad Datu, Silam Old Camp, 21 Dec. 1965, Ahmad
Talip SAN 52935 (SAN); Sepilok Forest Reserve, 11
Aug. 1959, Meijer SAN 19692 (SAN); Gunong Tambuku Heng Hiap, 14 March 1980, Fidilis & Sumbing
SAN 91763 (K, KEP, SAN); Sungai Gum Gum, Mile 17,
10 Dec. 1974, Tarmiji Arshid SAN 79648 (K, KEP, L,
SAN); Tawau, Camp, Luasong, 7 March 1977, Fidilis
Krispinus SAN 82174 (SAN); Beaufort, Lupak Camp,
13 Feb. 1962, Kinted SAN 19080 (K, KEP, SAN);
Sarawak: Kapit, Bukit Raya, 10 Nov. 1966, Suib et al. S
24234 (K, L, SAR); Sungai Perman, Batang Balui, 8
March 1987, Yii S 53426 (K, KEP, SAR); 4th Div.,
Bintulu, Segan Forest Reserve, 19 Sept. 1972, Chai S
32138 (BO, K, L, SAR); Sungai Kelawait, Tatau, 29
Sept. 1963, Ashton S 16477 (K, KEP, SAR, SING); Base
Camp along Sungai Melinau, 1 April 1978, Stone 13593
(K, KEP, SAR, SING); Belaga, Batang Balui, Ulu
Sungai Pemian, 20 March 1987, Yii S 53639 (K, KEP,
L, SAR); 7th Div., Kapit, Baleh, Sungai Sekawi, Ulu
Sungai Entutuh, 23 Sept. 1989, Othman & Rantai S
57825 (K, KEP, L, SAR); Baram Distr., Tutoh, Gunong
Api, Ulu Melinau, 4 Oct. 1971, Anderson S 31766 (K, L,
SAR). BRUNEI. Belait: Bukit Sawat, Jalan Labi, 14 Aug.
1991, Niga 288 (K); Bukit Sawat, Sungai Belait, 23 Oct.
1988, Wong 569 (K, L); Melilas, Sungai Belait, 1 Sept.
1989, Forman 1138 (K, L, SAR, SING). INDONESIA.
Sumatra: Upper Riauw, Pakanbaru, Tenajan R., 12
Aug. 1960, Soepadmo 43 (BISH, BKF, K); Bengkulu, S.
Ketahun, 9 Oct. 1993, Afriastini 2602 (K); Simaloer
[Simalur] Island, Tapah, 8 Dec. 1919, Achmad 1545
(K); Kalimantan, W. Koetai, 30 Oct. 1925, Endert 4717
(BO, K, L); Kalimantan: Sangkulirang Distr., Mt
Medadem, 7 Aug. 1957, Kosterman 13366 (K, L, BO);
Balikpapan-Samarinda road, 18 Aug. 1992, Ambrisyah
& Arifin AA553 (K, L); Central Kalimantan, Upper
Samba R., 28 Nov. 1982, Mogea 3761 (BO, K).
THE GENUS TEIJSMANNIODENDRON KOORD. (LAMIACEAE)
609
Map 10. Distribution of Teijsmanniodendron pteropodum.
PHILIPPINES. Samar, May – April 1914, Ramos 17449
(K); Leyte, Palo, Jan. 1906, Elmer 7096 (K).
HABITAT. Growing in primary and secondary forests,
including heath forest, often along riverbanks or in
freshwater swamps, on clay or sandy soils, sometimes
over basalt, sandstone or limestone; 0 – 960 m.
CONSERVATION STATUS. Least Concern (LC).
PHENOLOGY. Flowering from December to October;
fruiting from February to December.
VERNACULAR NAMES. Malaysia, Sabah: Buak-buak
(Tapak Itik) (Malay language); Sarawak: Entabuluh
(Iban language). Brunei: Mengkulat or Merkulat (Iban
language). Indonesia, Kalimantan: Miang (Dayak
language, ‘miang’ means can cause itchiness and
therefore the wood is not cut); Sumatra: Tjiempana
or Lipanoeh Pajo (Simalur Islands).
NOTES. The distictive winged petiole is a very variable
character, sometimes plants can be found to have
leaves with complete wings, wings only along the basal
part of the petiole or with no wings at all (de Kok,
pers. obs.).
13. Teijsmanniodendron punctatum G. Rusea sp. nov.
A Teijsmanniodendron hollrungii venis lateralibus foliorum paucioribus minus quam 15 numero differt. A
T. obscurinervis supeficie fructum maturorum sanosa
differt. Typus: Malaysia, Sarawak, Kuching, Semengoh
Forest Reserve, 14 July 1979, Othman Ismawi S 40040
(holotypus SAR!; isotypi K!, KEP!, L!).
Tree 4 – 8 m tall, DBH 10 – 30 cm. Bark greyish. Twigs
tetragonal, scabrous. Leaves unifoliolate, ovate to
lanceolate, 11 – 17 × 4 – 6 cm, apex acuminate, base
round to acute, margin flat, coriaceous, glabrous,
sparsely punctate beneath, feel rough beneath;
midrib prominent and raised on both surfaces; lateral
veins 6 – 11 pairs, prominent and raised beneath,
arching towards margin and looping into imperfect
intramarginal vein; intercostal veins obscure above,
visible beneath, subscalariform to reticulate, young
leaves coppery-reddish. Petioles 1 – 2 cm long, terete,
sparsely tomentose. Inflorescences 3 – 4 cm long, densely
tomentose; bracteoles narrowly lanceolate, densely
pubescent. Flowering calyx 2 – 2.5 × 1.5 – 2.2 mm,
campanulate, densely pubescent and whitish at base;
lobes 0.8 – 1 mm long; fruiting calyx shallowly
infundibular, irregularly lobed, densely pubescent.
Corolla purple, central lobe of lower lip spathulate,
2.5 – 3.5 × 1.5 – 2.5 mm, apex round, undulate, throat
pinkish-violet with yellow streak inside; side lobe of
lower lip 2 – 3 × 1 – 1.5 mm, apex acute; lobes of
upper lip 2 – 2.5 × 1.5 – 2 mm, apex acute; tube 3 –
3.5 mm long, purple, densely pubescent. Stamens 4, 3 –
4 mm long, distinctly didynamous, exserted, inserted
in lower part of the corolla tube, white; anthers
ellipsoid, brown. Ovary globose, c. 1 mm diam., apex
slightly villous; styles c. 5.5 mm long, stigmas c. 0.5 mm
long, blue to white. Fruits globose, 1 – 1.5 cm
(immature) diam., apex round, smooth, surface
powdery, apex with a few hairs; pericarp thin. Fig. 4.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
610
KEW BULLETIN VOL. 64(4)
Fig. 4. Teijsmanniodendron punctatum. A habit (from Bujang S 32547); B detail of node and petiole (from Rantai et al. S. 66077);
C underside leaf detail; D underside leaf closer detail; E inflorescence detail; F flower side view; G corolla opened; H fruit, side view
(from Othman Ismawi S 40040). DRAWN BY JULIET BEENTJE.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
THE GENUS TEIJSMANNIODENDRON KOORD. (LAMIACEAE)
DISTRIBUTION. Malaysia, Sarawak, Only known from six
collections, collected between 1957 and 1994, from
the Semengoh Arboretum, Kuching. Map 11.
SPECIMENS EXAMINED. MALAYSIA. Sarawak: Kuching,
Semengoh Arboretum, near tree 537, Aug. 1961,
Galau S.14828 (K, L, SAR); Semengoh Arboretum,
near tree no 4930, 19 Sept. 1972, Bujang & Othman
Ismawi S.32467 (K, L, SAR); Semengoh Arboretum,
tree no 6191, 4 Oct. 1972, Bujang S.32549 (K, L, SAR);
Semengoh Arboretum, 30 Sept. 1957, Bojang b Sitan
S.9388 (K, L, SAR); Semengoh Arboretum, southern
trail, 13 Sept. 1994, Rantai et al. S.66077 (K, KEP, SAR).
HABITAT. Found growing on slope of low hill, as
understorey trees of primary lowland forests on yellow
sandy clay soil; c. 100 m.
CONSERVATION STATUS. This species is only known
from six relatively recent collections from one protected area (Semengoh Arboretum). Given the small
area of occupancy (14 hectares) of the species and the
pressures on the forest in Sarawak and those in the
Kuching area in particular, this species is best classified
as “Vulnerable”.
PHENOLOGY. Flowering from July to October; fruiting
in July.
ETYMOLOGY. The Latin name refers to the many tiny
dots, which are filled with a sandlike substance (Rusea
& Latiff 2001).
VERNACULAR NAME. Sarawak: Entabuluh (Iban language).
14. Teijsmanniodendron renageorgeae G. Rusea sp.
nov. Ab omnibus ceteris speciebus compositifoliis
611
petiolo non alato atque minus quam 2 cm longo et
foliis ad apicem minus quam 2 cm latis differt.
Typus: Malaysia, Sarawak, Limbang, Ulu Medamit,
Sungai Ensungai, Tg. Long Amok, 21 Sept. 1980,
Rena George S 43074 (holotypus SAR!; isotypi K!,
KEP!, L!, SAN!).
Tree 4 – 13 m tall, DBH 15 – 20 cm. Bark smooth,
greyish-brown. Twigs terete, scabridulous when young,
glabrescent when mature, without a ridge between
opposite petioles, greyish-brown. Leaves 3-foliolate;
leaflets narrowly lanceolate, 8 – 10 × 2 – 3 cm, apex
narrowly acuminate, base symmetric (lateral leaflets
have distinctly oblique bases), margin strongly
recurved, glabrous, coriaceous, shining above, glaucous beneath; midrib sunken above, prominent and
slightly raised beneath; lateral veins 5 – 6 pairs,
obscure above, prominent beneath; intercostal veins
obscure on both surfaces. Petioles subterete, grooved
above, < 2 cm long, scabrous; petiolule 1 – 1.3 cm
long. Inflorescences 9 – 14.5 cm long, glabrous to
sparsely hairy; bracteoles narrowly lanceolate, pubescent. Flowering calyx 2 – 2.5 × 1 – 1.5 mm, campanulate, scabridulous, dark green; lobes 1 – 1.5 mm long,
apex acute; fruiting calyx c. 3.5 × 4 mm, fleshier than
flowering calyx. Corolla bilabiate, pubescent, violet with
yellow spot on the lip; central lobe of lower lip
damaged; side lobes of lower lip c. 2 × 1 mm, apex
round; lobes of upper lip damaged; tube c. 3 mm long.
Stamens 4, didynamous; anthers black. Ovary c. 1 mm
diam., glabrous with few hairs at apex. Fruits fresh
unknown, immature dried ones c. 2.5 × 3.5 mm;
Map 11. Distribution of Teijsmanniodendron renageorgeae (●) and T. punctatum (■).
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
612
young fruit globose, apex depressed, smooth, glabrous
with few hairs at apex. Fig. 5.
DISTRIBUTION. Known from two localities: Malaysia,
Sarawak, Ulu Medamit, Limbang and from Brunei,
Batu Melintang Hot Spring. Map 11.
KEW BULLETIN VOL. 64(4)
SPECIMEN EXAMINED. BRUNEI. Batu Melintang, Hot
Spring, 4 Jan. 1989, Kessler et al. 410 (L).
HABITAT. An understory tree in mixed dipterocarp
forest on loamy soil; 0 – 150 m.
CONSERVATION STATUS. This species is only known
from two relatively recent collections. As almost
Fig. 5. Teijsmanniodendron renageorgeae. A habit (from Bujang S 32547); B underside leaf detail; C detail of node and petiole
base; D inflorescence detail; E fruit in calyx. DRAWN BY JULIET BEENTJE.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
THE GENUS TEIJSMANNIODENDRON KOORD. (LAMIACEAE)
nothing is known about its ecology, a status of Data
Deficient (DD) is proposed.
PHENOLOGY. Flowering from September to January;
fruiting (young fruits) in September.
ETYMOLOGY. The species is named after the late Ms
Rena George (1956 – 1994), Assistant Forest Botanist
in the Sarawak Forest Department.
NOTES. Despite the incomplete material (only one
damaged flower seen), we decided to describe this
new species here, because its small narrowly lanceolate
leaflets, distinctly oblique base of the lateral leaflet
and the simple panicle with cluster of subsessile
flowers, make it very different from any other known
compound-leaved Teijsmanniodendron.
15. Teijsmanniodendron sarawakanum (H. Pearson)
Kosterm. (1951: 100 – 102); Moldenke (1980b: 29 –
31); Anderson (1980: 345). Type: Malaysia, Sarawak,
1865 – 1868, Beccari 2280 (lectotype K!, selected here).
Vitex sarawakana H. Pearson (1907: 60); Ridley (1929:
262); Merrill (1921: 514 – 515).
Vitex tetragona Hallier f. (1918: 53); H. J. Lam (1919:
202); H. J. Lam in H. J. Lam & Bakhuizen van den
Brink (1921: 52 – 54); Kostermans (1951: 100).
Type: Indonesia, Kalimantan, Gunung Pembliangan,
Nov. 1912, Amdjah 955 (holotype L!; isotype BO!).
Tree (2 –) 7 – 40 m tall, bole 8 – 16 m high, DBG 10 –
50 (– 165) cm; buttresses c. 0.5 m out. Bark smooth to
slightly scaly to flaky, white, grey to black; sapwood
hard or soft, yellowish or light brown to white. Twigs
glabrous, square in cross-section distally. Leaves unifoliolate; elliptic to oblanceolate or lanceolate, 8.5 –
30 × 2 – 11 cm, apex acuminate, base acute to
rounded, margin strongly recurved to sub-recurved,
chartaceous, glabrous, strongly bullate or not, shiny
above, not punctate beneath; midrib prominent;
lateral veins 5 – 12 pairs, arching and looping near
the margin, prominent and slightly sunken above,
raised beneath, colour distinctly paler than lamina;
intercostal veins scalariform-reticulate, distinct, raised
and whitish beneath. Petioles 1 – 3.5 cm long,
subterete, glabrous. Inflorescences up to 25 cm long,
glabrous, sometimes hairy at base; purplish; bracteoles
persistent, densely appressed-puberulous, caducous.
Flowering calyx campanulate, 1.5 – 2.5 × 1.5 – 2 mm
long, purple, densely minutely pubescent; lobes c.
1 mm long, apex acute; fruiting calyx shallowly cupuliform, 5 – 8 × 10 – 12 mm, irregularly lobed. Corolla
white or whitish-purple to pale pinkish-mauve or blue,
velvet, pubescent; central lobe of lower lip spathulate,
2.5 – 3 × 2.5 – 3 mm, apex round, white to purple with
yellow or purple markings at base, erect hairs at base;
side lobes of lower lip 1.5 – 2 × 1 – 2 mm, apex acute,
whitish-violet to yellow; lobes of upper lip 1.5 – 2 × 1 –
2 mm, apex acute, whitish-violet to yellow; tube 3 –
613
3.5 mm long, violet. Stamens 4, 3 – 4 mm long, slightly
didynamous, exserted, inserted halfway on tube, hairy
at base, white to yellow; anther c. 0.5 mm long, black.
Ovary globose, glabrous with velutinous upper half;
styles 3 – 8 mm long; stigma c. 0.5 mm long, apex
acute. Fruits globose, 10 – 25 × 15 – 25 mm, smooth,
apex round to depressed, smooth, glabrous with
velutinous apex or covered by a dark bluish-green
powder, dark grey to orange, later black; pericarp
thin.
DISTRIBUTION. Borneo. Map 12.
SELECTED SPECIMENS EXAMINED. MALAYSIA. Sarawak:
Simanggang, Bukit Sua, 21st Mile, Kuching-Simanggang Road, 5 Sept. 1966, Anderson S 24794 (K, L, SAR);
Tebedu, Bukit Simurus, 12 Feb. 1985, Mohtar et al. S.
49286 (K, KEP, SAR); Kapit, Bukit Raya Pelagus, 11
Aug. 1961, Anderson S 14717 (K, SAR); Kapit, Rejang,
March 1893, Havilland & Hose 3550 (BM, K, SAR,
SING); Kapit, Rejang R., Havilland 796 (K, L, SAR);
Bukit Sue, Kuching-Serbian Road, Burt & Martin B
4732 (E, SAR); Sabah, Tawau, 20 Oct. 1978, Fidilis &
Sumbing 88993 (K); Bukit Silam, 12 Oct. 1966,
Sinanggul SAN 57264 (K); Labuk Sugut Distr., near
Karamuak R., 24 – 25 June 1984, Beaman 10298 (K).
BRUNEI. Temburong: 21 Aug. 1990, Wong 1981 (K).
INDONESIA. Kalimantan: 150 km NE of Pontianak,
Gunung Bentuang, 28 June – 6 July 1989, Tukirin &
Burley et al. 3187 (E, K, KEP, L); 50 km NW of
Tumbangmiri, Sikatan Wana Raya logging concession,
19 April 1988, Burley et al. 714 (KEP, SAR); Central
Kalimantan, 92 km from Sangai, Kotawaringin, Nov.
1993, Tuke P13 605 (E, K); West Kalimantan, Gunung
Setutup, 11 Feb. 1992, Koyama et al. 3460 (L).
HABITAT. Found growing in primary and secondary,
sometimes heath forests on clays to sandy loams,
sometimes over shale, sandstone, limestone or ultrabasic rock; 35 – 900 m.
CONSERVATION STATUS. Least Concern (LC).
PHENOLOGY. Flowering from February to October;
fruiting from May to November.
VERNACULAR NAMES. Sarawak: Entabuluh (Iban language); Brunei: Mertuboh (Iban language).
16. Teijsmanniodendron scaberrimum Kosterm. ex G.
Rusea sp. nov. A Teijsmanniodendron simplicioidi ramulis
inflorescentiisque atque venulis foliorum pilis hispidis
obtectis differt. Typus: Indonesia, Kalimantan, Sangkulirang Distr., at Karangan R. near Batu Pondang, 1
Sept. 1957, Kostermans 13644 (holotypus K!; isotypi
BO!, KEP!, L!, NY!).
Teijsmanniodendron scaberrimum Kosterm. ex Moldenke
(1971: 911 & 1980b: 31), nom. nud.
Teijsmanniodendron holophyllum var. pubescens Moldenke
(1952: 57). Type: Indonesia, Kalimantan, Berouw
[Berau], 25 Oct. 1927, van der Zwaan 609 (bb.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
614
KEW BULLETIN VOL. 64(4)
Map 12. Distribution of Teijsmanniodendron sarawakanum.
12144) (holotype BO!, photo. K; isotypes L!, NY!),
synon. nov.
Tree 2 – 20 m tall, DBH 15 – 30 cm; buttresses absent
or up to 3 m high. Bark smooth to scaly, grey to light
brown; wood yellowish to pale brown. Twigs terete,
with stiff hairs to glabrescent. Leaves unifoliolate;
elliptic, 15 – 18 × 7 – 9 cm, apex broadly acute, base
broadly acute, margin strongly recurved, strongly
bullate, coriaceous, glabrous and shiny above, densely
hispid hairs, especially on veins beneath; midrib
sunken above, sharply raised beneath; lateral veins
5 – 6 pairs, arching and looping towards margin,
distinct and raised beneath; intercostal veins reticulate, distinct and raised on both surfaces. Petioles 3 –
4 cm long, terete, densely scabrous. Inflorescences up to
25 cm long, covered with hispid hairs, dark brown;
bracteoles linear, < 4 mm long, pubescent, persistent,
red. Flowering calyx 2.5 – 3 × 2 – 3 mm long, densely
hairy; lobes 1 – 1.2 mm long, apex acute; fruiting calyx
shallowly infundibular, 12 – 15 mm diam., slightly
pubescent at base, rim entire to slightly irregularly
lobed. Corolla dark violet to dark reddish-brown;
central lobe of lower lip oblong to spathulate, 3.5 –
4 × 2 – 3 mm, apex round, purple, covered with white
hairs; side lobes of lower lip 2.5 – 3 × 2 – 2.3 mm, apex
acute; lobes of upper lip 2.2 – 3 × 1.8 – 2 mm, apex
acute; tube 5 – 7 mm long; Stamens 4, 5 – 7 mm long,
didynamous, with few white hairs. Ovary globose, c.
1 mm diam., with a dense tuft of hairs at apex; style 8 –
8.5 mm long, glabrous to with few white hairs. Fruits
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
ellipsoid, 1 – 1.6 × 0.9 – 1.3 cm, smooth, impressed
apically, glabrous with some hairs at apex, dark
yellowish-orange; pericarp thin.
DISTRIBUTION. Brunei and Indonesia: East Kaliman-
tan. Map 13.
SPECIMENS EXAMINED. BRUNEI. Belait: Sungai Mutip,
26 July 1993, Atkins 588 (K, KEP); Andulau Forest
Reserve, 31 July 1963, Fuchs et al. 21155 (L). INDONESIA.
Kalimantan: Berouw [Berau], near Tdg. Redeb, 5 Oct.
1963, Kostermans 21113 (K); Berouw [Berau], near Tdg.
Redeb, 31 Oct. 1963, Kostermans 21565 (K); Berouw
[Berau], near Tdg Redeb, 31 Oct. 1963, Kostermans
21545 (L, SAR); Berau, 20 June 1992, Amiril Saridan AS
1 (K, L); N Kutei, Sangata, P.T. Porodisa Area, 21 Nov.
1994, Sidiyasa et al. 1173 (K, L).
HABITAT. Found growing in primary or secondary
forests, near rivers in floodplains, sometimes over
sandstone; 25 – 1000 m.
CONSERVATION STATUS. Least concern (LC). A rarely
collected, widely distributed species, which is reported
to be common when it is found.
PHENOLOGY. Flowering from June to November;
fruiting from July to September.
17. Teijsmanniodendron simplicifolium Merr. (1929:
263 – 264); Kostermans (1951: 96 – 97); Kochummen
(1978: 309); Moldenke (1980b: 32 – 34); Anderson
(1980: 345). Type: Malaysia, Sabah, Elphinstone Province [Tawau Distr.], Tawao [Tawau], Oct. 1922 – March
615
THE GENUS TEIJSMANNIODENDRON KOORD. (LAMIACEAE)
Map 13. Distribution of Teijsmanniodendron scaberrimum (●) and T. zainudinii (■).
1923, Elmer 21837 (holotype PNH; isotypes A!, BISH!,
BM!, BO!, K!, L!, NY!, SING!).
Teijsmanniodendron simplicifolium var. cordifolium Moldenke (1979a: 252). Type: Malaysia, Sabah, Ranau,
Simpang trail, 18 Sept. 1967, Aban Gibot SAN 60725
(holotype SAN!), synon. nov.
Tree 5 – 32 m tall, bole 3 – 14 m high, DBH 12 –
60 cm; buttresses to 2 m high, about 50 cm out. Bark
smooth to slightly scaly or fissured, white to whitishgreen or brown; sapwood hard, heavy, yellow to
brown. Twigs terete, glabrous to hairy. Leaves unifoliolate, lanceolate, 7 – 13 × 2 – 5 cm, apex acuminate,
base acute to rounded (cordate), margin flat, not
bullate, coriaceous, glabrous, light brown when dried;
midrib prominent; lateral veins 3 (– 4) pairs, curving
and ascending, arching towards the margin, very
conspicuous beneath; intercostal veins densely reticulate, distinct beneath. Petioles 0.8 – 1.2 cm long,
subterete, grooved above, glabrous, rarely with few
hairs in particular on the swollen articulations, distal
pulvinus sometimes with distinct black ring. Inflorescences up to 15 cm long, glabrous; bracteoles small,
lanceolate to elliptic-lanceolate, caducous. Flowering
calyx campanulate, 1 – 2 × 1 – 2 mm, glabrous, yellow
or whitish; lobes up to 0.5 mm long, apex acute;
fruiting calyx broadly infundibular, 7 – 10 × 3 – 8 mm,
irregularly lobed, glabrous. Corolla glabrous outside,
pale-yellow or white, fragrant; central lobe of lower lip
spathulate, 2 – 2.5 × 2 – 2.5 mm, apex round, with a
dark-yellow spot at the base; side lobes of lower lip
1.5 – 2.5 × 1 – 1.5 mm, apex acute; lobes of upper lip
1 – 2.5 × 1 – 1.5 mm, apex acute to round; tube 3 –
4 mm long, with blue-purple longitudinal stripes.
Stamens 4, 3 – 4 mm long, didynamous, exserted,
inserted in the lower part of the corolla tube; anthers
dark-purple. Ovary globose, c. 1 mm diam., apex
glabrous, covered in glands; style 3 – 5 mm long,
stigma c. 0.1 mm long. Fruits ovoid, 13 – 20 × 11 –
15 mm, apex notched, smooth with fine longitudinal
ridges, glabrous; pericarp thin.
DISTRIBUTION. Peninsular Malaysia, Sumatra and Bor-
neo. Map 14.
SELECTED SPECIMENS EXAMINED. MALAYSIA. Peninsu-
lar Malaysia: Pahang, Lesong Forest Reserve, 9 June
1979, Chan 25184 (K, KEP); Terengganu, 30.5 mile
Jerangau Road, 5 Sept. 1966, Rahim Ismail 99801 (K);
Sabah: Pandewan, Sungai Mesopo, 21 Feb. 1986, Fidilis
& Asid SAN 114020 (K, KEP, L, SAN); Lamag, Sungai
Lokam, 19 March 1963, How SAN 63305 (SAN);
Kinabatangan, Sungai Lokan, 7 Oct. 1968, Agis Horr.
SAN 64874 (SAN); Beaufort, Beaufort Hill, 19 Aug.
1965, Layangah SAN 44587 (SAN); Sungai Imbak, 2
Sept. 1992, Wong 2348 (E); Sarawak: Kapit, Bukit Raya,
12 Nov. 1964, Suib S 22249 (K); Bukit Mersing, Anap,
Dec. 1964, Sibat ak Luang, S 25045 (K, KEP, L, SAR);
Nanga Pelagos, 24 July 1938, Daud & Tachun, S 35644
(K, KEP, L, SAR); 4th. Div, Miri, Suai, Ulu Sungai
Sibau, 22 July 1977, Ilias Paie S 39165 (K, KEP, L, SAR).
BRUNEI. Andulau Forest Reserve, 20 Sept. 1957, Ashton
611 (K, KEP). INDONESIA. Kalimantan: West Kaliman© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
616
KEW BULLETIN VOL. 64(4)
Map 14. Distribution of Teijsmanniodendron simplicifolium.
tan, Serawai, 2 Feb. 1995, Church et al. 1687 (BISH, E,
K); Central Kalimantan, West Koetai, 27 Sept. 1925,
Endert 3625 (K, L); Sumatra, Riouw en Oud. Inderagirische bovenlanden [Riau and Indragiri], Keritang,
24 June 1939, Neth. Ind. For. Sevice bb 28654 (L).
HABITAT. Found growing in primary and secondary
forests on (sandy) clay or yellow sands, sometimes over
sandstone, basalt, shale or ultrabasic substrates; 0 –
1200 m.
CONSERVATION STATUS. Least Concern (LC).
PHENOLOGY. Flowering from (December) February to
August (September); fruiting from March to October
(December).
VERNACULAR NAMES. Malaysia, Sabah: Buak-buak
(Malay language); Sarawak: Ubah Sireh or Entabuluh
(Iban language), Esak or Punok (Kayan Language);
Indonesia, Kalimantan: Kaju Gadang or Kemuning
(Dayak language).
USES. The wood is used in Sabah and said to be
durable against insect attack; in Sarawak the wood is
used for canoes and the bark for roofing.
NOTES. Teijsmanniodendron simplicifolium var. cordifolium
Moldenke is rejected here as a separate taxon as the
defining character (leaf base cordate) according to
Moldenke (1979a) is not stable. Specimens with leaflets that are both rounded to slightly cordate at base
can be easily found.
18. Teijsmanniodendron simplicioides Kosterm.
(1962b: 303 & 325). Type: Indonesia, SE Kalimantan,
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
SE Borneo, Berouw Betemoean [Berau Betemuan], 28
May 1934, van der Zwaan 1074 (bb 19034) (holotype
BO!; isotypes A!, BO!, L!, NY!).
Teijsmanniodendron simplicifolium var. kostermansii Moldenke (1952: 57 & 1971: 641).
Tree 5 – 60 m tall, bole 3 – 20 m high, DBH 10 –
120 cm, fluted or with steep buttresses. Bark whitish,
pale yellow to greyish-brown, smooth; inner bark
yellowish; sapwood yellowish, hard. Young twigs densely
golden pubescent, terete. Leaves unifoliolate, sometimes
pseudo-whorled, lanceolate, 6 – 10 (– 22) × 3 – 8 cm,
apex acuminate, base rounded to broadly acute,
margin flat, glabrous on both surfaces, coriaceous;
midrib prominent above, prominent and slightly raised
beneath; lateral veins 4 – 10 pairs, sunken above, raised
beneath, arching towards the margin, not looping but
becoming faint near margin towards apex; intercostal
veins prominent. Petioles 0.5 – 3 cm long, densely
pubescent when young, sometimes glabrous when old.
Inflorescences densely pubescent to at least densely hairy
at base; bracteoles 1 – 5 mm long, caducous. Flowering
calyx campanulate, 1.5 – 2 × 2 – 3 mm, margin
undulating and uneven, glabrous; fruiting calyx 7 –
10 mm diam., erect, margin undulating and uneven,
glabrous. Corolla yellowish-white to (bluish) yellow or
purple; central lobe of lower lip 3 – 4 × 3 – 5 mm,
spathulate, apex round, throat villous inside; side lobe
of lower lip 2 – 2.5 × 1.8 – 2 mm, apex round; lobes of
upper lip 2 – 2.5 × 2 – 3 mm, apex round; tube 4.5 –
5 mm long. Stamens 4, 3 – 4 mm long, didynamous,
THE GENUS TEIJSMANNIODENDRON KOORD. (LAMIACEAE)
yellow; anther c. 0.5 mm long, round. Ovary globose, c.
1 mm diam., villous at apex; style 3 – 4 mm long, stigma
c. 0.1 mm long. Fruits ellipsoid, 1 – 2.2 × 0.6 – 1.2 cm,
smooth, apex round, glabrous with a few glands;
pericarp thin.
DISTRIBUTION. Peninsular Malaysia (Terengganu) and
Sabah; Indonesia: Kalimantan Timur. Map 15.
SELECTED SPECIMENS EXAMINED. MALAYSIA. Peninsular
Malaysia: Terengganu, Dungun, Bauk Forest Reserve, 4
Sept. 1966, Ismail 98944 (L, KEP); Sabah: Sandakan,
Hutan Sibuga Forest Reserve, 25 April 1962, Singh SAN
34721 (K, SAN); Sandakan, Leila Forest Reserve, 28
Feb. 1962, Sam SAN 28833 (K, SAN); Lahad Datu,
Forest Reserve, Ulu Segama, 11 Aug. 1986, Maidin SAN
116861 (K, SAN); Beaufort, Beaufort road, 16 Oct.
1962, Lajangah SAN 32196 (K).
HABITAT. In primary and secondary forests often on
ridges, on sandy, clay or clay loam, sometimes over
sandstone or ultrabasic; 60 – 150 m.
CONSERVATION STATUS. Least concern (LC).
PHENOLOGY. Flowering recorded from September to
April; fruiting from March to October.
VERNACULAR NAMES. Sabah: Buak-buak (Malay language); Kilons (Dusun – Papar language).
NOTES. This species is a very close relative of Teijsmanniodendron simplicifolium, and further research may well
prove that it is just a variety of it. T. simplicioides differs
from T. simplicifolium in having densely hairy young
petioles and branches and fruits that are entirely
smooth.
617
19. Teijsmanniodendron sinclairii Kosterm. (1958: 6 –
8); Kochummen (1978: 309); Moldenke (1980b: 36 –
37); Anderson (1980: 345). Type: Peninsular Malaysia,
Terengganu, Kuala Terengganu, Besut Road, Sungai
Nerus riverbank, 18 Sept. 1955, Sinclair & Kiah SFN
40877 (holotype SING!; isotypes BO!, E!, K!, L!).
Tree 3 – 20 m tall, DBH 2 – 45 cm. Bark smooth, grey;
sapwood white to pale yellowish-brown. Twigs terete,
stout; young twigs minutely scabrous, older ones
glabrous. Leaves unifoliolate, elliptic, 17 – 25 × 8 –
13 cm, apex acuminate, base acute, margin strongly
recurved, distinctly bullate, coriaceous, glabrous (hairy
on veins below in Peninsular specimens), shiny; feels
rough beneath; midrib sunken above, raised beneath;
lateral veins (5 –) 8 – 10 (–20) pairs, sunken above,
prominent and raised beneath, arching towards the
margin; intercostal veins densely reticulate, impressed
above, distinct and raised beneath. Petioles 2 – 4 cm
long, terete, glabrous, distal pulvinus with distinct
black ring. Inflorescences up to 30 cm long, sparsely
scabrous; bracteoles c. 3 × 8 mm, persistent, narrowly
ovate to elliptic-lanceolate, apex acute. Flowering calyx
campanulate, 2 – 3 × 2 – 3 mm, with few hairs; lobes
c. 0.5 mm long, apex broadly acute, erect; fruiting calyx
shallowly cupuliform, 10 – 12 × 5 – 7 mm, rim
irregularly lobed, minutely pubescent. Corolla funnelshaped, petals blue, dark-violet or purple, pubescent;
central lobe of lower lip oblong to round, 3.5 – 3.8 ×
2 – 3 mm, apex round, yellow-brown patch at base,
throat villous at base; lower lip side lobes 2 – 2.2 ×
Map 15. Distribution of Teijsmanniodendron simplicioides.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
618
KEW BULLETIN VOL. 64(4)
1.5 – 2 mm, apex round; lobes of upper lip 1.5 – 2 ×
1 – 1.2 mm, apex round; tube 2 – 4 × 2 mm. Stamens 4,
5 – 7 mm long, didynamous, exserted, inserted in the
middle part of the corolla tube, white; anther black.
Ovary globose, c. 1 mm diam., pubescent at apex; style
6.5 – 7 mm long; stigma blue. Fruits ellipsoid, 15 – 17 ×
10 – 12 mm, apex depressed, glabrous, smooth with
faint longitudinal groove, glabrous, dark-green; pericarp thin.
ETYMOLOGY. This species is named after James
Sinclair, 1913 – 1968, botanist at the Singapore
Botanic Gardens.
VERNACULAR NAME. Sarawak: Entabuluh (Iban language).
NOTES. The specimens from Peninsular Malaysia differ
from the Borneo collections in having hairs on the
veins beneath the leaf and a more densely pubescent
inflorescence.
DISTRIBUTION. Peninsular Malaysia: (Terengganu) and
20. Teijsmanniodendron smilacifolium (H. Pearson)
Kosterm. (1951: 95 – 96); Moldenke (1980b: 37 – 39).
Type: Malaysia, Sarawak, 1865 – 68, Beccari 1097
(lectotype K!, selected here; isolectotypes BM!, NY!).
Vitex smilacifolium H. Pearson (1907: 59); H. J. Lam
(1919: 175); H. J. Lam & Bakhuizen van den Brink
(1921: 48 & 51); Merrill (1921: 514).
Borneo. Map 16.
SELECTED SPECIMENS EXAMINED. MALAYSIA. Sabah:
Lahad Datu, Mt Silam, 21 Dec. 1966, Sinanggol SAN
57450 (K, L, SAN); Sarawak, Bintulu, Tatau, Kuala
Sungai Mina, 13 March 1965, Anderson 20939 (K);
Bintulu, Kakus, Ulu Mayang, 19 July 1964, Sibat ak
Luang S 21756 (K, SAR). INDONESIA. Kalimantan: E
Kalimantan, Berau, 16 Dec. 1997, Ambriansyah &
Arifin Berau 1070 (E, K); W Kalimantan, Serawai, Batu
Lintang, 24 Jan. 1995, Church et al. 1433 (BISH, E, K).
HABITAT. Found growing in primary and secondary
forests, often along riverbanks on alluvium, sometimes
over sandstone or basalt; 0 – 500 m.
CONSERVATION STATUS. Least concern (LC). However, the Peninsular Malaysian population has been
collected only once and more work is needed to
determine its conservation and taxonomic status.
PHENOLOGY. Flowering from March to October;
fruiting from September to December.
Map 16. Distribution of Teijsmanniodendron sinclairii.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
Tree 4 – 30 m, bole 10 – 25 m high, DBH 10 – 55 cm;
buttresses present. Bark smooth, powdery to scaly,
whitish-grey to yellow or black; slash yellow, grey to
pale-brown; sapwood white to yellowish-brown, soft.
Twigs terete, glabrous. Leaves unifoliolate, oblanceolate, 12 – 20 × 4 – 8 cm, apex acute or broadly
acuminate, base broadly acute to rounded, margin
slightly recurved, glabrous on both surfaces, coriaceous, shiny above, not punctate beneath; midrib
prominent above and below; lateral veins 3 – 4 pairs,
least one pair of major lateral veins originating at base
of leaf, all major lateral veins uniformly prominent
THE GENUS TEIJSMANNIODENDRON KOORD. (LAMIACEAE)
and arching towards to apex, intercostal veins finely
reticulate, obscure above, prominent beneath. Petioles
1 – 2 cm long, subterete, glabrous. Inflorescences up to
30 cm long, glabrous; bracteoles small or minute,
caducous. Flowering calyx campanulate, 1 – 2 × 1 –
2 mm, sparsely and minutely glandulous; lobes
c. 0.5 mm long, apex acute; fruiting calyx cupuliform,
6 – 9 mm diam., accrescent, irregularly lobed,
glabrous. Corolla dull dark glaucous; pale-purple or
blue to white, pale mauve, white (yellow); central lobe
of lower lip darkish-violet near the villous throat, with
a distal pale-yellow spot, cream; side lobe of lower lip
cream; lobes of upper lip cream; tube funnel-shaped,
upper part marked externally with glands. Stamens 4,
distinctly didynamous, slightly exserted, inserted below
the middle part of the corolla tube; filaments darkpurple to faint-violet, basally dilated and villous;
anthers dark-purple or black. Ovary globose, c. 1 mm
diam., glabrous, densely glandular-dotted; styles
c. 6 mm long, white. Fruits globose, 1.2 – 1.8 × 0.8 –
1.2 cm, apex round to acute, finely ribbed when dry,
glabrous, black; pericarp thin.
DISTRIBUTION. Borneo. Map 17.
SELECTED SPECIMENS EXAMINED. MALAYSIA. Sabah,
Sandakan, Telupid, Mile 80, 27 July 1972, Leopold
Madani et al. SAN 75369 (K); Beaufort Hill, 19 Aug.
1965, Lajangah SAN 44587 (K, L, SAN); Sarawak, 1865 –
1868, Beccari 1137 (K); Kuching, Semengoh Arboretum,
17 March 1962, Galau S 15620 (K, L, SAR); Kapit, Bt.
Raya, 15 Nov. 1964, Suib S 22298 (K, L, SAR). Bintulu,
Nyabau, 8 Dec. 1961, Ilias Paie S 15877 (K, L, SAR);
619
Balingian, Nanga Arip, 17 Oct. 1963, Ashton S 19580 (K,
SAR). INDONESIA. Kalimantan: Central Kalimantan,
Sambas, 31 Jan. 1995, Jarvie & Ruskandi 5803 (K,
SAR); Kalimantan Tengah, P.B.U. Base Camp, 11 July
1990, Ridsdale PBU22 (E, K); W Kalimantan, am Kapûas,
29 Sept. 1893, Hallier B 219 (L); Gunung Palung, 4 Nov.
1996, Laman et al. TL 334 (L).
HABITAT. Found growing in primary or secondary
forests on clay or black stony soil, sometimes over
shale; 0 – 1000 m.
CONSERVATION STATUS. Least concern (LC).
PHENOLOGY. Flowering from November to July; fruiting from October to November.
TYPIFICATION. Two collections were cited in the
original publication (Pearson 1907): Borneo, Sarawak,
Beccari 1097 (syntype, BM!, K!, NY!) and Beccari 1137
(syntype, K!). The Beccari 1097 collection is chosen
here as the lectotype as it has more duplicates.
21. Teijsmanniodendron subspicatum (Hallier f.)
Kosterm. (1951: 99 – 100); Moldenke (1980b: 39 –
42); Anderson (1980: 345). Type: Indonesia, Sumatra,
1880, Forbes 3204 (lectotype L!, selected here; isolectotypes BM!, BO!, L!, SING!).
Vitex subspicata Hallier f. (1918: 52 – 53); H. J. Lam
(1919: 177 – 178); H. J. Lam & Bakhuizen van den
Brink (1921: 52); Merrill (1921: 514).
Tree 3 – 20 m, DBH 6 – 40 cm; buttresses to 1 m high.
Bark smooth to flaky cracked, yellow to grey; slash
white or light-greenish, yellowish, yellow or orange,
Map 17. Distribution of Teijsmanniodendron smilacifolium.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
620
granular, soft; sapwood pale brown to yellowish. Twigs
white or pale ochraceous, stout, terete, glabrous.
Leaves unifoliolate, lanceolate, (8 –) 15 – 30 (– 38) ×
(3 –) 10 – 14 (– 16) cm, apex acuminate, base broadly
acute to rounded, margin recurved, coriaceous, subbullate beneath, shiny and glabrous on both surfaces,
scruffy, not punctate beneath; midrib impressed
above, prominent and raised beneath; lateral veins
7 – 10 pairs, arching towards the margin and sometimes forming an (imperfect) intramarginal vein,
prominent and raised beneath; intercostal veins reticulate, prominulent and slightly raised beneath. Petioles
1 – 3 cm long, subterete, glabrous, distal pulvinus
sometimes with distinct black ring. Inflorescences up to
25 cm long, dark purplish-red, very minutely puberulent; bracteoles minute, linear, persistent. Flowering
calyx campanulate, 1.5 – 2 × 2 – 2.2 mm long,
puberulent, green to dark red-purple; lobes up to
0.3 mm long, apex broadly acute; fruiting calyx
shallowly infundibular, 6 – 8 × 10 – 18 mm, much
accrescent, irregularly dentate, dirty brownish-yellow.
Corolla pubescent, lilac to purple; central lobe of lower
lip round to spathulate, 2 – 4 × 1.5 – 2 mm long, band
of white hairs along length of lip, lilac-blue with yellow
blotch at base; side lobes of lower lip 1.5 – 2.8 × 0.8 –
1 mm, apex round to acute, dark violet; lobes of upper
lip 1 – 1.2 × 0.8 – 1 mm, apex round, dark violet; tube
2.5 – 3.5 mm long, pale yellow or white to pale-dark
purple. Stamens 4, 3 – 5 mm long, distinctly didynamous, exserted, inserted in the lower part of the
corolla tube, white, basally pale-blue, villous; anthers
Map 18. Distribution of Teijsmanniodendron subspicatum.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
KEW BULLETIN VOL. 64(4)
black, c. 0.5 mm long. Ovary globose, c. 1 mm diam.,
villous at apex; style 5 – 7 mm long, exserted, white to
pale blue. Fruits ellipsoid, 9 – 20 × 8 – 15 mm, apex
depressed, smooth, glabrous with few hairs at apex,
black; pericarp thin.
DISTRIBUTION. Sumatra and Borneo. Map 18.
SELECTED SPECIMENS EXAMINED. MALAYSIA. Sabah:
Lahad Datu, Gunung Silam, 21 Dec. 1965, Talip SAN
52933 (K); Beluran, Sungai Meliau, 26 July 1983, Sigin
et al. SAN 99716 (K); Tenom, Crocker Range, above
Amboi, 26 March 1965, Sadau SAN 49460 (K);
Sarawak, 7th Div. Kapit, Bukit Raya, 13 Oct. 1965,
Jugah ak Kudi S 23856 (K, L, SAR, SING). BRUNEI.
Belait: Melilas, along river bank of Ulu Belait, 22 July
1993, Atkins 529 (K). INDONESIA. Sumatra: Medan,
Sikundur Forest Reserve, 3 Aug. 1979, de Wilde & de
Wilde-Duyfjes 19321 (K); Sikundur Forest Reserve, 6
Aug. 1979, de Wilde & de Wilde-Duyfjes 19466 (K);
Kalimantan: Gunung Meranti, 25 Feb. 1994, Argent
94117 (E, KEP); E Kalimantan, W Kutei, Mt Palimasan
on Belajan R., 19 Sept. 1956, Kostermans 13161 (BM, K, L);
Central Kalimantan, Kabupaten Kapuas, Katunjung
village, 15 Oct. 2001, Sidiyasa et al. 2609 (K, L); Central
Kalimantan, Kotawaringin Timur, 92 km from Sangai, 3
July 2002, Tuke P2 2066 (E).
HABITAT. Found growing in primary and secondary
forests, sometimes swamp or kerangas forest or along
riverbanks on white sand to sandy or loamy clays,
sometimes on acid soils or limestone; 10 – 400
(– 1000) m.
THE GENUS TEIJSMANNIODENDRON KOORD. (LAMIACEAE)
CONSERVATION STATUS. Least concern (LC).
PHENOLOGY. Flowering in September to June (August);
fruiting from July to May.
VERNACULAR NAMES. Malaysia, Sabah: Kedaras (Dusun
language); Sarawak: Selumpo, Medang Sisit, Entabuluh
(Iban language), Ubah Putih (Malay language).
Brunei: Mertubu (Iban-Brunei language).
TYPIFICATION. Four different collections were cited in
the original description (Hallier 1918): Sumatra, 1880,
Forbes 3204 (syntypes BM, BO, L, SING); W Kalimantan, grossen Sambasflusse [Sungai Sambas Besar], 4
Nov. 1893, Hallier B 1064 (syntypes BO, L); W
Kalimantan, Terussan, zwischen dem grossen und
dem kleinen Sambas [between the Sungai Sambas
Besar and Sambas Kecil], 5 Nov. 1893, Hallier B 1122
(syntypes BO, L); S.E. Kalimantan, Berg Pamatton [Mt
Pamaton], Korthals s.n. (syntypes L). Forbes 3204, which
has fruits, is chosen here as the lectotype, because fruit
characters are more informative in this genus than
flowers, and there are many duplicates of this collection in various herbaria.
22. Teijsmanniodendron unifoliolatum (Merr.) Moldenke
(1952: 58 & 1980b: 42 – 43); Anderson (1980: 345).
Type: Philippines, Mindanao, Zamboanga Distr., Melangas,
Oct. – Nov. 1919, Ramos & Edano 37048 (holotype PNH;
isotype K!).
Vitex unifoliolata Merr. (1922: 438 – 439).
Tree 1.5 – 26 m tall, DBH 10 – 35 (– 100) cm,
sometimes fluted. Bark smooth to flaky, whitish to
greyish-green or brown; sapwood soft, white to paleyellow; hardwood brown, hard. Twigs tetragonal,
glabrous, smooth. Leaves unifoliolate, narrowly elliptic to lanceolate, 12 – 40 × 4 – 11 cm, apex
acuminate, base acute to rounded, margin flat to
recurved, shiny above, glabrous on both surfaces,
slightly bullate, young leaves feel rough beneath, not
punctate beneath, coriaceous; midrib prominent and
raised on both surfaces; lateral veins 6 – 12 pairs,
arching towards the margin, sometimes forming
intramarginal veins, prominent and raised beneath;
intercostal veins laxly reticulate, distinct and slightly
raised beneath, obscured above, concolorous with
leaf blade. Petioles 1 – 3 cm long, subterete, glabrous.
Inflorescences up to 40 cm long; peduncles about 6 cm
long, glabrescent, violet; bracteoles lanceolate to
elliptic-lanceolate, caducous. Flowering calyx campanulate, 1.8 – 3 × 1 – 2 mm long, green to (dark)
purple, pubescent; lobes c. 0.5 mm long, apex acute
to obtuse; fruiting calyx cupuliform, 0.7 – 1.2 cm
diam., erect, lobed to entire, glabrous to pubescent.
Corolla velvety, scented, blue to violet or (white)
purple; central lobe of lower lip orbicular to spathulate, 2.5 – 4 × 2 – 3 mm, apex round, yellow spot at
base, erect hairs at base; side lobe of lower lip 2.5 –
621
3 mm long, apex acute, blue; lobes of upper lip 1.5 –
3 mm, apex acute, blue; tube 3 – 5 mm long, purple
to violet. Stamens 4, 2 – 5.5 mm long, distinctly
didynamous, greatly to shortly exserted, inserted in
the middle or lower part of the corolla tube, villous at
base, white to faintly violet; anthers black. Ovary
globose, c. 1 mm diam., villous at apex; styles
c. 6 mm, white; stigma c. 0.5 mm. Fruits globose,
0.7 – 2 × 1 – 1.7 cm, smooth, apex round to
depressed, glabrous except for a few hairs at apex,
covered with sooty yellow- or greenish-brown powder
which forms a cracked layer when dry, pale purple to
black; pericarp thin with pithy layer within.
DISTRIBUTION. Borneo and Southern Philippines.
Map 19.
SPECIMENS EXAMINED. MALAYSIA. Sarawak: Sungai
Selangsar, 8 Nov. 1979, Othman et al. S 41315 (K,
KEP, SAR); Ulu Kenawit, Tatau, 29 Sept. 1963, Ashton
S.16476 (K); Bt. Mersing, 25 Sept. 1964, Sibat ak Luang
S 22370 (K); Nanga Pelagos, 23 July 1937, Daud &
Tachun SFN 35638 (BISH, K, SAR); Kapit, Belaga
Subdistr., left bank of Rajang R., near Belaga airfield, 1
Sept. 1958, Jacobs 5391 (K, L); Sungai Sama, 13 Aug.
1938, Daud & Tachun SFN 36053 (BISH, K); Mulu
National Park, Ulu Sungai Berar, 3 Oct. 1977, Chai
39632 (K, L, KEP, SAR); Upper Rejang R., 1929,
Clemens & Clemens 21825 (BM, K); Kuching, Semengoh Forest Reserve, 19 Nov. 1994, Jegong & Shalih S
68689 (E, K, SAR); Ulu Katibas, Bukit Pelandok, 19
Nov. 1997, Pearce et al. ITTO/BB 585 (SAR); Sabah:
Beluran, S of Labuk Bridge, 8 Dec. 1979, Dewol
Sundaling SAN 91068 (K). BRUNEI. Tutong: Ulu Supan,
10 Jan. 1958, Ashton BRUN 863 (K, KEP, L). INDONESIA.
Kalimantan, Kotawaringin, 3 July 1994, Tuke Djuda
94255 (E, K, L); E Kalimantan, Berouw [Berau], Mt
Ilas Bungaan, 12 Sept. 1957, Kostermans 13845 (K);
Berouw [Berau], Mt Ilas, 8 Sept. 1957, Kostermans 13710
(K); E Kalimantan, Sangkulirang Distr., Karangan R.,
near Batu Pondong, 2 Sept. 1957, Kostermans 13649
(KEP, L); E Kalimantan, Gunung Buntung, 4 Jan. 1981,
Kato & Wiriadinata B 5369 (L); Kalimantan, Bukit Raya,
Jan. 1983, Nooteboom 4544 (L); Central Kalimantan,
Kotawaringin Timur, 12 May 1993, Argent et al. 93135
(K).
HABITAT. Growing in primary and secondary, sometimes in heath (kerangas) forests, often along rivers or
in seasonally inundated forest on sand, clay or sandy
loam or shale-derived soil, sometimes over basalt,
sandstone or limestone; 0 – 800 m.
CONSERVATION STATUS. Least concern (LC).
PHENOLOGY. Flowering from (March) August to April;
fruiting from June to January.
VERNACULAR NAMES. Malaysia, Sarawak: Entabuluh
(Iban language). Indonesia, Central Kalimantan: Kayu
Sanaman (Dayak language). Philippines: Babako (Tagalog
language).
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
622
KEW BULLETIN VOL. 64(4)
Map 19. Distribution of Teijsmanniodendron unifoliolatum.
23. Teijsmanniodendron zainudinii G. Rusea sp. nov.
A Teijsmanniodendron smilacifolio petiolis maturis minus
quam 1 cm longis, lamina crasse coriacea differt et
habitat in Sabah septentrionali substrata calcarea
ultra-basica tantum incolens. Typus: Malaysia, Sabah,
Sandakan, Telupid, Bukit Tawai Forest Reserve, 11
April 1994, Zainudin AZ 5009 (holotypus UKMB!;
isotypi KEP!, SAN!).
Teijsmanniodendron subspicatum var. parvifolium
Moldenke (1979a: 252 & 1980b: 42). Type: Malaysia,
Sabah, Sandakan Distr., Tawai Plateau, Ulu Karamuak,
Meijer SAN 39328 (holotype SAN!; isotype K!),
synon. nov.
Tree 2 – 10 (– 25) m tall, DBH 3 – 20 cm. Bark smooth,
pale green to whitish; sapwood yellow to yellowish–
ochre; resin yellowish-ochre turning brown on exposure.
Twigs greyish-brown, glabrous, terete. Leaves unifoliolate,
elliptic to lanceolate, 7 – 9 × 3 – 5 cm, apex acuminate
to acute, base round to slightly cordate, margin flat or
recurved, thick coriaceous, glabrous above, feel smooth
beneath, not punctate beneath; midrib prominent, flat
to slightly raised above, raised beneath; lateral veins 5 – 7
pairs, faint above, rather prominent beneath but not
raised, arching towards margin and looping into a faint
intramarginal vein; intercostal veins invisible above, fairly
faint beneath, finely reticulate. Petioles < 1 cm long,
subterete, scabrous. Inflorescences panicles of subsessile
branched cymes, violet; bracteoles narrowly lanceolate,
< 3 mm long, caducous. Flowering calyx infundibular, 1 –
2 × 1.5 – 2 mm, glabrous, reddish-purple; lobes up to
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
0.5 mm long, apex acute glabrous; fruit calyx cupuliform,
12 – 14 × 4 – 5 mm, lobes irregular, glabrous, dark
greyish-brown. Corolla slightly pubescent on both surfaces; purple to violet or white with yellow hairs; central
lobe of lower lip spathulate, 2 – 2.5 × 3 – 3.5 mm, apex
round or emarginate, reflexed, throat villous inside; side
lobes of lower lip 2 – 3 × 2 – 2.5 mm, apex round; lobes
upper lip 2 – 3 × 1.5 – 2 mm, apex round; tube 4 –
5.5 mm long. Stamens 4, 4 – 6 mm long, didynamous,
exserted, inserted in the lower part of the tube. Ovary
globose, 1 – 2 mm diam., glabrous, apex covered with
glands; style 3 – 5 mm long. Fruits ellipsoid, 14 – 17 ×
10 – 12 mm, apex acute with slight notch at apex,
smooth, glabrous; pericarp thin. Fig. 6.
DISTRIBUTION. Malaysia, Sabah: Only known from a
few collections from the ultrabasic outcrops North
West of Sandakan. Map 13.
SPECIMENS EXAMINED. MALAYSIA. Sabah: Kinabatangan,
Bukit Tawai Forest Reserve, 11 April 1994, Bojo &
Cheksum 36 (KEP); Bukit Tawai Forest Reserve, E of
sawmill near Karamuak R., 11 June 1992, Meijer &
Madani SAN 131966 (E, K, KEP, L, SAR); Bukit Tawai
Forest Reserve, 4 March 1985, Mansus et al. SAN 107787
(K, L); Bukit Tawai Forest Reserve, logging road to first
waterfall, 7 April 1994, Mat-Salleh KMS 3331 (KEP); Bukit
Tawai Forest Reserve, logging road to first waterfall, 7
April 1994, Mat-Salleh KMS 3303 (E, KEP, L); Bukit Tawai
Forest Reserve, Sungai Meliau, 9 April 1994, Soepadmo et
al. FRI 41327 (KEP); Tongod, Sungai Pinangah Forest
Reserve, 3 Nov. 2004, Sugau et al. SAN 142192 (KEP);
THE GENUS TEIJSMANNIODENDRON KOORD. (LAMIACEAE)
623
Fig. 6. Teijsmanniodendron zainudinii. A habit (from Meijer & Madani SAN 131966); B detail of node and petiole base; C underside
leaf detail; D flower, side view (from Meijer & Madani SAN 24184); E corolla opened and style; F young fruit; G mature fruit (from
Manus & Dewol SAN 107787). DRAWN BY JULIET BEENTJE.
Telupid, Bukit Tangakunan, 5 Feb. 1982, Rahim SAN
92990 (L); Ranau, Bukit Hampuan, 28 Feb. 1961, Singh
SAN 24184 (K).
HABITAT. Found in lowland dipterocarp forests and
heath forest dominated by Gymnostoma sumatrana
(Jungh. ex de Vriese) L. A. S. Johnson on clay over
ultrabasic substrates; 150 – 850 m.
CONSERVATION STATUS. This species is only known
from a few collections from the ultrabasic outcrops
North West of Sandakan. These outcrops usually have
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2010
624
some protective status, but they only cover a relatively
small area and are, like most ultrabasic vegetations,
prone to forest fires. Given the small area of occupancy and the higher fire risk, this species is best
classified as “Vulnerable”.
PHENOLOGY. Flowering in February to April; fruiting
from March to November.
ETYMOLOGY. Named after Ahmed Zainudin Ibrahim,
Plant Collector at UKMB and collector of the type
specimen.
NOTES. Teijsmanniodendron subspicatum var. parvifolium
Moldenke has been raised here to species level and
given the new name: Teijsmanniodendron zainudinii.
This became necessary as more specimens showed
that this taxon differs in a series of characters from T.
subspicatum (including smaller leaves 7 – 9 × 3 – 5 cm
rather than 8 – 38 × 3 – 16 cm and fewer lateral veins
5 – 7 pairs rather than 7 – 10, intercostal veins almost
invisible rather than prominent and the petiole < 1 cm
long rather than 1 – 3 cm).
Excluded Species
Teijsmanniodendron petelotii Moldenke (1950: 225 –
226). Type: North Vietnam, Tonkin, province de
Sontây, 16 April 1941, Mont Bavi, Petelot 6801
(holotype NY!; isotype A!).
This species is not a Teijsmanniodendron due to the
lack of a swollen petiole base and apex and the leaves
being arranged in whorls of three, which is not
characteristic for this genus. It is most likely to be
either a Vitex or Premna.
Acknowledgements
The authors are grateful to the Curators of A, BISH, BKF,
BM, BO, E, K, KEP, L, LAE, NSW, NY, SAR, SAN, SING
and UKMB herbaria for access and/or loan of specimens
used in the present study. This study is part of a revision by
G. Rusea of the genus Teijsmanniodendron carried out for
the Tree Flora of Sabah and Sarawak Project and funded the
Malaysian Government through an IRPA grant made
available to Forest Research Institute of Malaysia. It was
also supported through the generosity of the Royal
Society, (travel grant 2005) made available through the
Royal Botanic Gardens, Kew, which supported G. Rusea’s
visit to Kew. We are grateful for this support. Rogier de
Kok was supported during fieldwork by the Kew TOBU
fund. The drawings were made by Juliet Beentje. The
authors are grateful to David Mabberley, Gemma
Bramley and Engkik Soepadmo for useful comments
made on earlier versions of the manuscript.
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