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); © The Board of Trustees of the Royal Botanic Gardens, Kew, 2010 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. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2010 600 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 – © The Board of Trustees of the Royal Botanic Gardens, Kew, 2010 604 KEW BULLETIN VOL. 64(4) 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!). © The Board of Trustees of the Royal Botanic Gardens, Kew, 2010 606 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. 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