Kew Bulletin

, Volume 67, Issue 3, pp 293–329

A revision of the genus Gmelina (Lamiaceae)

Article

DOI: 10.1007/s12225-012-9382-4

Cite this article as:
de Kok, R. Kew Bull (2012) 67: 293. doi:10.1007/s12225-012-9382-4

Summary

A revision of the genus Gmelina L. (Lamiaceae) is presented with a summary of its taxonomic history, keys, full descriptions, distribution maps, conservation assessments, ecological information and ethno-botanical notes. For the New Caledonia species no descriptions and distribution maps are given as they have been provided in an earlier publication. In this treatment, 31 species are recognised, 18 names are placed into synonymy for the first time, five are lectotypified, one is neotypified (G. racemosa (Lour.) Merr.) and seven names are excluded from the genus. Two varieties (G. palawensis var. palawensis and G. palawensis var. celebica) are raised to subspecies level. Four new species are described: G. australis de Kok, G. basifilum de Kok, G. hollrungii de Kok and G. peltata de Kok.

Key Words

distribution taxonomy 

Introduction

The genus Gmelina was first described by Linnaeus in 1753 on the basis of one species Gmelina asiatica L. The genus is named after Johann Georg Gmelin (1709 – 1755), who was Professor of Medicine, Botany and Chemistry in Tübingen, Germany (1747 – 1755), and Professor of Chemistry and Natural History in St. Petersburg, Russia (1731 – 1747). He was elected to accompany Vitus Bering’s Second Kamchatka Expedition (1731 – 1742), which formed the basis of his Flora Sibirica (1747 – 1769).

The genus occurs naturally from India and South China to North Australia and Fiji. Some species have been introduced to many subtropical and tropical countries, either as a timber tree (Gmelina arborea Roxb. ex Sm.) (Harley et al.2004) or as an ornamental (G. asatica, G. elliptica Sm. and G. philippinensis Cham.). The total number of species has been estimated from about 33 (Munir 1984) to about 40 (Mabberley 2008): in this revision 31 species are accepted, with two subspecies. Most of the genus has been provisionally revised by Moldenke (1984c, d, e, f, g, h) in a series of articles. In this series the taxa were presented in alphabetical order, but strangely he abruptly stopped after G. tonkinensis Moldenke ( = G. elliptica) despite the ‘to be continued’ note at the base of the page. In addition, a substantial number of regional revisions have been published. Most notable are Lam (1919) for SE Asia and the Pacific, and Lam & Bakhuizen van den Brink (1921) and Fletcher (1938a) for Thailand. These revisions are now completely out of date. More recent local revisions are available for Australia (Munir 1984), New Caledonia (Mabberley in Mabberley & de Kok 2004), India (Rajendran & Daniel 2002), Sri Lanka (Moldenke & Moldenke 1983), China (Shou-liang & Gilbert 1994) and Vietnam (Vũ Xuân Phuöng 2007). The tree species of Borneo have been revised in the Tree Flora of Sabah and Sarawak (Bramley et al.2011); a revision of the species of mainland Asia has been the subject of a PhD study (Mohammad Harun-ur Rashid, Trinity College Dublin) and those of Peninsular Malaysia have been revised by Bramley & de Kok (in press).

Unlike most other genera in the Lamiaceae (de Kok 2007, 2008 & in press; de Kok et al.2009), the genus Gmelina seems to be particularly rich in species and morphological variation in New Guinea and its surrounding islands. The number of native species is considerably fewer on the islands of the Sunda-Shelf. A second, but less important centre of speciation is on the SE Asian mainland, in particular Indo-China and Southern China.There have been very few attempts to subdivide Gmelina. Briquet (1895) divided the genus into two sections, Microstromatae and Bracteosae, based on the size, colour and venation of the floral bracteoles, with G. philippinensis as the only species in the section Bracteosae, and G. asiatica as the type of section Microstromatae. This division is no longer used and is not practical given the wide variation in bracteole morphology within this genus.

Rumphius’s species

Rumphius described three species in his genus Tittius in Herbarium amboinense vol. 3 (1750), two of which are now thought to be part of Gmelina. The species Tittius alba Rumph. and T. rubra Rumph. are identified in this study as G. moluccana (Blume) Backer ex K. Heyne. The third species of the genus, T. litoralis Rumph. is placed by Merrill (1917) under the species in Guettarda speciosa L. (Rubiaceae).

Uses

The genus is most known for its use as a timber tree (Gmelina arborea) (Dvorak 2004), but it is also extensively used as a garden plant (mainly G. philippinensis, but also G. asiatica and G. elliptica). The fast growing timber tree G. arborea is increasingly being used in reforestation projects. The cultivated species of the genus also have a wide range of medicinal uses (Burkill 1966).

Morphology

Habit

Most species are tall trees, often with buttresses, but some very common species (Gmelina asiatica, G. elliptica and G. philippinensis) are qualified either as (scrambling) shrubs or as small trees. One species is a liana proper (G. uniflora Stapf). The spines present in some species (G. asiatica, G. elliptica and G. philippinensis) are modified branches and would be better defined as thorns (Beentje 2010). The leaves of the seedlings are 3 – 7-lobed (Ng 1992).

Indumentum

There are two types of indumentum present in Gmelina. The most common are simple multicellular hairs that occur in all species and are common in the family Lamiaceae. In some species these can form dense mats on the underside of the leaves, on the infloresence and/or calyces, and in those cases can have taxonomic significance. The second type is a form of peltate scales that are either present or absent in a species. When they are present there is great variety in form and number. In some species the lower leaf surface is completely covered with these scales resulting in a whitish appearance (G. uniflora and G. asiatica). In other species the number of scales are fewer and they are less conspicuous without a × 10 lens (G. phillipensis and G. smithii Moldenke). The shape of the scales can vary from those with entire margins to those with incised margins that begin to resemble stellate hairs.

Glands

Glands on leaves. There is only one type of gland present on the leaves of Gmelina. Discoid glands are present as either solitary glands or in gland fields at the base of the leaves or in rows along the main vein. These glands are reminiscent of similar discoid glands in other genera (Clerodendrum L. and Oxera Labill.) in the Lamiaceae (Cantino 1990). In a small number of species these glands are partly enveloped in a cup-like structure at the base of the leaves. This cup-like structure seems to be a modification of the base of the leaf blade, where a number of glands, which form a dense gland-field, have been incorporated. These cup-like glandular structures are often used as a useful field character for the genus as they are prominent in the widespread species G. arborea. From notes on herbarium specimens it is clear that ants are attracted to the extrafloral glands on the leaves and calyx.

Glands on calyx. Discoid glands similar to those found on the leaves are present in some species on the outer calyx surface. In his study of extrafloral nectaries, Burck (1891) noted that glands on the calyx of some species (Gmelina asiatica and G. philippinensis [as G. bracteata]) growing in the botanic gardens at Bogor in Java, were visited by ants, which then patrolled the inflorescences. The presence of these glands on the calyx is relatively common in the genus and is often a good taxonomic character for species recognition.

Glands on bracteoles. A small group of species (Gmelina australis de Kok, G. papuana Bakh., G. ledermannii H. J. Lam and G. schlechteri H. J. Lam) from New Guinea and northern Australia is marked by having discoid glands on the bracteoles. These glands are always present and are often very prominent, giving the bracteoles the appearance of having ‘eyes’.

Inflorescences

Almost all species have a terminal inflorescence, very rarely also an axillary one. Only one species (Gmelina uniflora) has consistently an axillary inflorescence only. In the latter case the inflorescence also has a small number of flowers; while most species have many-flowered inflorescences. The inflorescence is of a very common type in the family and is similar to the second type inflorescences in the genus Vitex L. (de Kok 2007). The inflorescence is a panicle with the bracts that are always leaf-like and persistent. The main inflorescence axes are seemingly indeterminate. The main axis and major side branches (when present) bear compact to lax small cymes.

Bracteoles

The variation in bracteoles in Gmelina is remarkable: they can be small and narrow and falling off early in the development of the inflorescence (G. asiatica), while in other species they can be big, colourful and remain on the infloresence even during the fruiting period (G. philippinensis). In his study of extrafloral nectaries Burck (1891) noted that at Bogor the corollas of G. philippinensis [as G. bracteata] were significantly less broken into by insects than those of a species with smaller bracteoles (G. asiatica). His conclusion was that the many, big bracteoles provided room for ants to make nests that, in so doing, would be better able to protect the corollas.

Flowers

Calyx. All species have five calyx lobes. However in many species these lobes are hardly developed and sometimes appear to be practically absent, although it is possible to find relictual or very small lobes on some calyces. The outer surface of the calyx is either glabrous or velutinous when in flower, this character usually persisting when the plant is in fruit, though sometimes the calyx will lose some of its hairs during the fruiting period and then may appear glabrous. In one species (Gmelina racemosa (Lour.) Merr.) the calyx is very big (> 5 mm long) and completely envelopes the fruits, with the calyx lobes growing into wing-like structures. In this case the calyx is persistent with the fruit and calyx dispersed as one unit.

Corolla. There are two types of corolla shape in the genus. The corollas of Gmelina asiatica, G. elliptica and G. philippinensis are always a uniform bright yellow and have four corolla lobes with a prominent anterior lip. The second type has five lobes, very rarely four (G. arborea, which usually has corollas with four and five lobes together on one plant and G. delavayana Dop), due to a split in the posterior lip; the two lips are generally unequal, but the anterior one is much less prominent than in the first corolla type.

The colour is much more difficult to define in these species: some are uniformly white or pink to pale yellow or purple, but more often they have several colours in each flower, and there is a distinct difference between the main corolla parts and the lip, or between the tube and lobes, or between the inside and outside of the corolla.

The inside of the corolla is always glabrous, while the outside surface can be glabrous to velutinous. The corolla lobes can, in some species, be velutinous on the margins. Most corollas seem to continue to grow when they open, such that measurements of corolla parts can vary greatly within species.

Fruits

Unlike some other closely related genera (Teismanniodendron and Premna) in the Lamiaceae (de Kok 2008 and in press; de Kok et al.2009), fruit characters have not traditionally been considered to have taxonomic significance in this genus. Gmelina fruits are two-celled, each cell containing two ovules. It is unusual for both ovules in a cell to develop.

Within the mature fruit there always is a cavity, the origin of which is yet to be determined. It is present in every mature fruit, does not contain any seeds and is clearly not a gall. In young fruits this cavity can be filled with pith. Such cavities can be very small, in particular in young fruits, but in some species a cavity can make up more than 50 % of the total volume of the mature fruit. The shape varies with species and the state of development of the fruit, as well as the number of seeds developing within that particular fruit. Usually the cavity is below the seeds, but it can occupy the whole length of the fruit or sometimes it is only distal to the seeds. Any possible function of this cavity is unclear; there is no indication that species in this genus are distributed via water, which would explain the cavities as floating devices. A more likely explanation could be that it is an efficient way of increasing the size of the fruit, and species of this genus are therefore able to mobilise a different suite of dispersers from their close relatives. More research on fresh material is needed to study the development of these cavities and elucidate their possible function.

Materials and Methods

This study is based on close examination of herbarium specimens from the following herbaria: BM, BO, K, L, LINN, P and SING (abbreviations following Thiers 2011); many (type) specimens were diagnosed using images on the web.

In the following descriptions:
  1. i)

    all measurements and colour descriptions are from mature material;

     
  2. ii)

    all collections cited have been seen by the authors unless indicated otherwise;

     
  3. iii)

    all measurements of the stigma are made from the longer branch;

     
  4. 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;

     
  5. v)

    all conservation status assessments follow the criteria set by (IUCN 2001).

     

Gmelina

GmelinaL. (Linneaus 1753: 626); Schauer (1847: 678); Briquet (1895: 173); Lam (1919: 214 – 228); Lam & Bakhuizen (1921: 64 – 71); Fletcher (1938a: 422 – 424); Moldenke (1984c: 308 – 342; 1984d: 424 – 442; 1984e: 460 – 499; 1984f: 32 – 54; 1984g: 102 – 126; 1984h: 154 – 182); Moldenke & Moldenke (1983: 388 – 401); Munir (1984: 91 – 116); Shou-liang & Gilbert (1994: 32 – 34); Rajendran & Daniel (2002: 153 – 171); Mabberley in Mabberley & de Kok (2004: 22 – 34); Harley et al. (2004: 195 – 196); Vũ Xuân Phuöng (2007: 135 – 146); Bramley et al. (2011); Bramley & de Kok (in press). Type: Gmelina asiatica L.

[Michelia Amman (1739: 218, t.18). Based on Michelia spinosa Amman (1739: 218 – 219, t.18)  =  Gmelina asiatica L or Gmelina elliptica Sm.].

[Tittius Rumph. (Ruphius 1750: 38 – 39); Merril (1917). Based on Tittius alba and Tittius rubra  =  Gmelina moluccana (Blume) Backer ex K. Heyne; Tittius litoralis  =  Guettarda speciosa L. (Rubiaceae)].

Cumbulu Rheede ex Adans. (Adanson 1763: 199). = Gmelina arborea Roxb. ex Sm.

Ephielis Sol. ex Seem. (Seemann 1865: 258), nom. illegit., non Ephielis Schreb. (Schreber 1789: 253). Based on Ephielis simplicifolia Sol. ex Seem. = Gmelina dalrympleana (F. Muell.) H. J. Lam.

Trees, shrubs or rarely lianas, sometimes spiny, sometimes deciduous. Leaves simple, decussate, sometimes lower surface covered with hairs or with peltate scales, often with discoid glands at base, sometimes these glands concentrated in a cup-like structure. Inflorescence terminal, rarely axillary, dense to lax, panicle with cymose side branches; bracts leaf-like, decreasing in size towards apex; bracteoles present. Calyx usually weakly 2-lipped, 5-lobed, sometimes appearing fewer, persistent, usually accrescent, usually with large discoid glands without. Corolla zygomorphic, 2-lipped, 4 – 5-lobed, blue or purple-violet, white, yellow, brownish, mauve or purple pink, inside glabrous, tube widely funnel-shaped, greatly enlarged at throat, posterior lip 1 – 2-lobed, anterior lip 3-lobed, mid-lobe largest. Stamens 4 (− 5), didynamous, inserted usually from halfway to at the apex of the corolla tube, only once at the base (Gmelina basifilum), anthers dorsifixed; anterior pair longer, always fertile, included to exserted; posterior pair rarely sterile. Pollen 3-colpate. Ovary imperfectly 2-celled, each cell 2-ovuled; only two ovules developing, the other two are suppressed. Style filiform, terminal; stigma-lobes 2, unequal. Fruits drupaceous, with very fleshy mesocarp and tough endocarp, cavity present. Seeds 0 – 4 seeded, seeds without endosperm.

distribution. India and South China south to Australia, New Caledonia and Fiji.

habitat. Growing in deciduous or evergreen forest or anthropogenic vegetation; alt. 0 – 1500 (− 3000) m.

uses. Planted as ornamentals (Gmelina asiatica, G. elliptica and G. philippinensis); used as light and medium-weight timbers (G. arborea). G. arborea is used increasingly in SE Asia as a pioneer species in reforestation projects.

Key to Gmelina species

  • 1. Shrubs or small trees, sometimes climbing, twigs usually with spines; corolla 4-lobed, bright yellow........................2

  • Trees or lianas, rarely shrubs, twigs always without spines; corolla 5 (very rarely 4)-lobed, white to yellowish, bluish to purple or red...............................................................................................................................................4

  • 2. Lower leaves surface densely villous…..........................................................................................................8. G. elliptica

  • Lower leaves surface glabrous or with only a few hairs on the veins..............................................................................3

  • 3. Bracteoles linear to elliptic, < 6 mm wide, usually not persistent...............................................................2. G. asiatica

  • Bracteoles ovate-lanceolate, > 10 mm wide, usually persistent......................................................23. G. philippinensis

  • 4. Lower leaf surface covered with whitish peltate scales, sometimes also with simple hairs...........................................5

  • Lower leaf surface without peltate scales........................................................................................................................21

  • 5. Lianas or small trees; inflorescence few flowered, axillary; corolla pale yellow to white; endemic to Borneo...........................................................................................................................................30. G. uniflora

  • Trees, rarely shrubs; inflorescence usually many flowered, terminal, rarely axillary ones also present; corolla bluish to reddish purple to white; widespread.................................................................................................................6

  • 6. Mature lower leaf surface velutinous, usually with yellow hairs.....................................................................................7

  • Mature lower leaf surface glabrous to sparsely hairy with white hairs on veins..........................................................10

  • 7. Leaves with two glands in a cup-like structure at the base; corolla (pale reddish) yellow...................1. G. arborea

  • Leaves without glands or in gland fields at base or along midvein; corolla (cream-) white to pink...........................8

  • 8. Fruit apex truncate, sparsely covered with peltate scales, grey when mature; endemic to Indo-China.....................................................................................................................................................12. G. lecomtei

  • Fruit apex round, glabrous, bluish-purple when mature; endemic to Australia or New Caledonia......................9

  • 9. Inflorescence open; endemic to Australia...........................................................................................14. G. leichhardtii

  • Inflorescence compact; endemic to New Caledonia.....................................................................19. G. neocaledonica

  • 10. Calyx lobes  >  4 mm long when in flower, endemic to China and Indo-China.........................................................11

  • Calyx lobes  <  2.5 mm long when in flower, widespread................................................................................................12

  • 11. Leaf  <  5 cm long; flowering branches slender  <  2 mm diam.; corolla 4-lobed...................................7. G. delavayana

  • Leaf  >  5 cm long; flowering branches  >  2 mm diam.; corolla 5-lobed..................................................24. G. racemosa

  • 12. Calyx glabrous when in flower; sometimes covered with peltate scales.....................................................................13

  • Calyx velutinous when in flower; sometimes covered with peltate scales..................................................................16

  • 13. Leaves with side veins  >  9.................................................................................................................................................14

  • Leaves with side veins  <  7...............................................................................................................................................15

  • 14. Corolla pale pink to purplish brown; frurayishish white when mature; endemic to the Solomon Islands..........................................................................................................................................................22. G. peltata

  • Corolla blue; fruit purple or mauve when mature; endemic to Fiji.....................................................31. G. vitiensis

  • 15. Corolla mid lobe apex acuminate, outer surface glabrous to hairy; fruit apex covered with peltate scales; endemic to (Indo-) China.......................................................................................................................5. G. chinensis

  • Corolla mid lobe apex rounded, outer surface velutinous; fruit without peltate scales; endemic to the Moluccas and New Guinea.......................................................................................................................................15. G. lepidota

  • 16. Shrubs; corolla lobes almost isomorphous, tube elongated (tube  >  24 mm long), not greatly expanding above the calyx, velutinous; endemic to New Caledonia................................................................................29. G. tholicola

  • Trees; corolla lobes clearly different from one other, tube 8 – 14 mm long, greatly expanding above the calyx.................................................................................................................................................................................17

  • 17. Leaves with two glands in a cup-like structure at the base.....................................................................1. G. arborea

  • Leaves without glands, or glands arranged in gland fields at base or along midvein.............................................18

  • 18. Filaments inserted at base of flower tube...............................................................................................4. G. basifilum

  • Filaments inserted at apex of flower tube....................................................................................................................19

  • 19. Calyx lobes  >  1 – 2.5 mm long when in flower; endemic to New Guinea................................................28. G. smithii

  • Calyx lobes  <  1 mm long when in flower; endemic to Australia or the Solomon Islands.........................................20

  • 20. Bracteoles clavoid to rounded, 2.5 – 5 mm long; flowers blue to white with a purple lip; endemic to Australia...................10. G. fasciculiflora

  • Bracteoles lanceolate to spathulate, 5 – 30 mm long; flowers (grey) white, sometimes with a purple base; endemic to the Solomon Islands....................................................................................................25. G. salomonensis

  • 21. Mature lower leaf surface hairy.....................................................................................................................................22

  • Mature lower leaf surface glabrous or with hairs on the veins only..........................................................................23

  • 22. Bracteoles lanceolate, 4 – 10  ×  1 – 5 mm; Moluccas to Solomon Islands.........................................18. G. moluccana

  • Bracteoles rounded, 13 – 16  ×  11 – 13 mm; occurring in Eastern New Guinea..............................27. G. sessilis

  • 23. Corolla lobes almost isomorphous, tube not greatly expanding above the calyx; endemic to New Caledonia................................................................................................................................................17. G. magnifica

  • Corolla lobes greatly different from one other, tube greatly expanding above the calyx; occurring in Australia, Solomon Islands and New Guinea...............................................................................................................................24

  • 24. Calyx glabrous when flowering, lobes 0 – 1 mm long.................................................................................................25

  • Calyx velutinous when flowering, lobes  <  0.5 mm long.............26

  • 25. Leaves with two cup-glands at base; mature fruit with a truncate apex; Australia and New Guinea.........................................................................................................................................6. G. dalrympleana

  • Leaves with several discoid glands at base; mature fruit with a rounded apex; endemic to Palau Islands and occurring on the Caroline Islands......................................................................................................20. G. palawensis

  • 26. Bracteoles with several disc-like glands present..........................................................................................................27

  • Bracteoles without disc-like glands..............................................................................................................................30

  • 27. Bracteoles petiolate, 12 – 23  ×  5 – 9 mm, apex rounded to acute; flowering calyx 4 – 4.5  ×  6 – 7 mm; fruit apex truncated; endemic to Australia, Northern territory.............................................................................3. G. australis

  • Bracteoles sessile, fruit apex rounded; endemic to New Guinea.............................................................................28

  • 28. Inflorescences dense without long side branches; bracteoles rounded on the main axis, 7 – 15  ×  2 – 8 mm, apex acute; flowering calyx 3.5 – 5  ×  2.5 – 4 mm; corolla lip 6 – 10 mm long.........................................21. G. papuana

  • Inflorescences narrow with long side branches; bracteoles apex acuminate to long acuminate.........................29

  • 29. Bracteoles on main inflorescence axis rounded, apex long acuminate; flowering calyx 3.5 – 5  ×  3.5 – 4 mm; corolla lip 7.5 – 10 mm long.............................................................................................................13. G. ledermannii

  • Bracteoles on main inflorescence axis lanceolate, apex acuminate; flowering calyx 2.5 – 3  ×  2.5 – 3 mm; corolla lip 5 – 8 mm long.................................................................................................................................26. G. schlechteri

  • 30. Leaves with cup-like glands at base; Moluccas, New Guinea, Australia and the Solomon Islands..............................................................................................................................................11. G. hollrungii

  • Leaves without cup-like lands, discoid glands sometimes present at base; endemic to New Caledonia...................31

  • 31. Evergreen small tree or shrub, up to 3 m tall; endemic to north-west New Caledonia.....................9. G. evoluta

  • Deciduous tree, up to 20 m tall; endemic to Valée de Thy (New Caledonia).......................16. G. lignum-vitreum

1. Gmelina arboreaRoxb. ex Sm. (in Rees 1810); Lam (1919: 219 – 220); Greaves (1981: 237 – 258); Moldenke (1984c: 337 – 342; 1984d: 424 – 442; 1984e: 460 – 469).

Gmelina arborea var. arborea Roxb. ex Moldenke & Moldenke (1983: 390 – 394). Type: Rheede,. Hort. Malab. 1 (1678) t. 41, selected here.

Gmelina oblongifolia Roxb. (Roxburgh 1814: 95, nom. nud.; 1832: 83); Moldenke (1984h: 156 – 157), synon. nov. Type: Icon. Roxburgh t. 2306 (K!).

Premna tomentosa Miq. ex C. B. Clarke (1885: 581), nom. illegit., non Willd. (Willdenow 1802).

Premna arborea Roth (1821: 287); Rajendran & Daniel (2002: 156). Type: ‘Ind. Orient.’, Heyne s.n. (holotype?).

Gmelina rheedei Hook. (Hooker 1848: t. 4395) as rheedii; Lam (1919: 219). Type: Curtis’s Bot. Mag. 4: t. 4395.

Gmelina arborea var. glaucescens C. B. Clarke (1885: 582); Rajendran & Daniel (2002: 155). Type: [India] Khasia Mts, 24 July 1850, Hooker s.n. (holotype K!).

Gmelina arborea var. canescens Haines (1910: 487); Rajendran & Daniel (2002: 155). Type: [India, Bengal] Santal Parganahs, 2 Feb. 1907, Haines 5769 (holotype CAL (n.v.); isotype K!).

Gmelina arborea forma dentata Moldenke (1961: 14); Moldenke & Moldenke (1983: 390 – 394). Type: India, Siwalik & Jaunsar division, Jhajra, 14 March 1922, AzizullahShah s.n. (holotype CAVA scan seen).

Trees 3 – 30 m high, DBH 10 – 200 (− 450) cm, sometimes deciduous. Bark smooth; whitish to greyish-reddish brown. Twigs densely tomentose, spines absent. Leaves broadly ovate to reniform, 8 – 25 × 4.5 – 20 cm, apex rounded to acuminate, base cuneate to (deeply) cordate, margins entire, sometimes slightly lobed, papery; upper surface with hairs on veins, dark green, shiny; lower surface glabrous to densely hairy, covered with peltate scales, two cup-like glands clustered around the base; veins 3 – 5 pairs, sometimes 3 veined from base. Petiole terete, 3.5 – 13 cm, hairy. Inflorescence terminal or axillary, narrow; peduncle 10 – 30 cm. Bracteoles lanceolate to linear or triangular, 8 – 15 × 1.3 – 2 mm long, deciduous. Calyx 2 – 7 × 4 – 5 mm, 5-lobed, velutinous, not accrescent, green; lobes 0.8 – 1 × 0.7 – 2 mm, acute, outer surface with scattered discoid glands. Corolla (pale reddish) yellow, outer surface hairy, sparsely glandular, 4 – 5-lobed; anterior lip 3-lobed; mid-lobe spathulate 15 – 20 × 15 – 25 mm, (reddish) yellow, apex rounded; side lobes 10 – 15 × 9 – 12 mm long, apex rounded, reddish brown; posterior lip 1 – 2-lobed, lobes 6 – 7 × 6 – 12 mm, apex rounded, reddish brown; tube 15 – 20 mm long. Stamens inserted at apex of tube, glabrous, sometimes with a few glands, yellow; anterior pair 15 – 20 mm long, anthers 2 – 3 mm long, black; posterior pair, fertile; 12 – 14 mm long, anthers c. 1 mm long. Ovary glabrous; style 15 – 23 mm long, stigma c. 1 mm long. Fruit 12 – 20 × 10 – 15 mm, ellipsoid to obovoid-ellipsoid, glabrous, apex rounded, (dark) purple to yellow when mature, recorded as having a bitter sweet taste.

distribution. Natural distribution: from Mirpur in India and Jamu in Pakistan east to South Yunnan in China and northern Vietnam, south to Northern Thailand and Sri Lanka (Map 1). It is introduced in most tropical countries as a timber-, or shade- tree as well as an ornamental.
Fig. 1

Gmelina australis. A habit; B glands on bracteole; C flower side view; D calyx; E corolla; F style, stigma and ovary; G fruit (dried). From Blake 17169, sheets 1 & 2. drawn by juliet beentje.

Fig. 2

Gmelina basifilum. A habit; B detail leaf underside, scales; C flower side view; D calyx; E corolla; F stamen; G style and ovary; H fruit (dried). From Koster BW 11115. drawn by juliet beentje.

Fig. 3

Gmelina hollrungii. A habit; B detail leaf underside, cup-like glands; C flower side view; D calyx; E corolla; F style and ovary; G stigma. From Takeuchi 6373. drawn by juliet beentje.

Fig. 4

Gmelina peltata. A habit; B narrow leaf; C detail leaf underside, gland field; D detail leaf underside, scales; E flower side view; F calyx; G calyx detail, scales; H corolla; J style, stigma and ovary; K fruit. A, CJ from Gafui & collectors 16942, B from Runikera & collectors 12504, K from Runikera & collectors 12504. drawn by juliet beentje.

Map 1

Probable natural distribution of Gmelina arborea (●); distribution of Gmelina moluccana (▲); Gmelina vitiensis (■).

selected specimens examined. bangladesh. [East Pakistan] Chittagong, Chuak R.F., 27 Feb. 1966, Majumder & Islam 38A (K). INDIA. Maharashtra State: Khandala [Khaudala] saddle, 21 April 1943, Santapau 1932 (K); Tamil Nadu State: Mont Nilghiri, Thomson s.n. (K); Hort. Cal., 11 Sept. 1819, Wallich 1832 (K); Meghalaya State: Khasia Mts, 24 July 1850, Hooker s.n. (holotype K); Jharkhand State: [Bengal] Santal Parganahs [Santhal Pargana], 2 Feb. 1907, Haines 5769 (isotype K); Ultarakhand State: Siwalik & Jaunsar division, Jhajra, 14 March 1922, AzizullahShah s.n. (holotype CAVA). LAOS. Khammouan, Nakai Talang, 1 March 2007, Vannachak BT 913 (L). MYANMAR. Pyiamaua, March 1913, Gamble s.n. (K). SRI LANKA, Nalanda, 19 May 1947, Worthington 2787 (K). THAILAND. Rachasima, Pu Kio, 25 Feb. 1931, Kerr 20271 (K); Huai Krasa, 90 km S of Tak, 18 March 1968, Hansen & Smitinand 12952 (K).

habitat. Growing in deciduous or evergreen forest or anthropogenic vegetation; alt. 0 – 1500 m. It is a relatively short-lived tree that colonises openings in the forest, where the ground has been disturbed. Usually it is not common and it does not grow on poorly drained soils. Soil (sandy) loam to clay.

conservation status. Least Concern.

phenology. In the more seasonal areas it is reported to flower between April and May; fruiting occurs between May and July; in wet-tropical areas flowering and fruiting throughout the year. The seeds fail to germinate in the full shade, but germinate well in the open (Mohamad & Ng 1982).

uses. Planted widely as an ornamental, shade and/or timber tree. The wood is used for various purposes including construction, furniture, and musical instruments. The tree also has a wide range of medicinal uses (Burkill 1966).

vernacular names. India, Bengal: Ato Kasmar; Bombay presidency: Sioni (Kanarese language); Sri Lanka: Et demata (Shingalese Language) or Kumil (Tamil Language); Myanmar: Yamanay; China: yun nan shi zi; Laos: Saw or Mai so; Thailand: The rong (Karieng language); Peninsular Malaysia: Bulang (Malay Language) or Jemane; Sabah: Jemane (Malay Language) or Jamaná (Dusun Language); Fiji: Yemane.

notes. Chromosome 2n  =  36 (Mangenot & Mangenot 1962: 411). The type material of Gmelina oblongifolia Roxb. falls within the total morphological variation of G. arborea ex. Sm. and is therefore placed into its synonomy.

2. Gmelina asiaticaL. (Linnaeus 1753: 626); Lam (1919: 221 – 223); Lam & Bakhuizen (1921: 69 – 70). Gmelina asiatica var. typica Bakh. in Lam & Bakhuizen (1921: 69 – 70), nom. illegit., superfl. Type: Herb. Linn. 780–2 (holotype LINN!).

Gmelina parviflora Roxb. (Roxburgh 1802: 32 – 33, t. 162); Gmelina asiatica f. parviflora (Roxb.) Moldenke (1984a: 42). Type: Roxburgh (1799 – 1805), Pl. Coast. Coromand., Pl. 162.

Gmelina parviflora Pers. (Persoon 1806: 142). Based on Roxburgh (1799 – 1805 Pl. Coast. Coromand., Pl. 162, t 32 and Burman (1768) Flora Indica, p. 332.

Gmelina asiatica Wall. (Wallich 1828 n. 1818), nom. nud., nom. illeg., non Gmelina asiatica L. Based on: [India] Botanical Garden, 22 Nov. 1814, [Wallich] 1818 (K-W!).

Gmelina attenuata H. R. Fletcher (1938b: 203 – 204), synon. nov. Type: Thailand, Payap, Chiengmai, 4 July 1922, Kerr 6224 (holotype E scan seen).

Gmelina paniculata H. R. Fletcher (1938b: 204); Moldenke (1984h: 161), synon. nov. Type: Thailand, Prachinburi, Krabin, Aranya, 18 Oct. 1928, Put 2086 (holotype E scan seen; isotype K!)

Gmelina asiatica forma lobata Moldenke (1975: 47), synon. nov. Type: Sri Lanka, Wilpattu National Park, Smithsonian Camp, Marai Villu, 30 June 1969, Wirawan, Cooray & Balakrishnan 899 (holotype NY scan seen; isotypes K!, L!).

Shrubs, sometimes climbing, 2 – 4 m high, rarely deciduous. Twigs minutely hairy when young; spines usually present, up to 25 mm long. Leaves ovate to elliptic or deltoid, 0.5 – 5 (− 13)  ×  0.5 – 3.3 (− 6) cm, apex rounded to acute, papery, base rounded to cuneate, margins entire to 5-lobed; upper surface glabrous, or with a few hairs on veins, dark green, shiny; lower surface glabrous or with a few hairs on veins, whitish, covered with peltate scales; veins 3 – 4 pairs, sometimes 3-veined from base, few discoid glands present. Petiole 5 – 30 mm, glabrous or with a few hairs, peltate, scales absent to many. Inflorescence terminal, sometimes appearing axillary on short side shoots, 1.5 – 10 cm long, pendulous or erect, sparsely hairy, covered with white peltate scales; bracteoles 8 – 10  ×  3.5 – 6 mm, linear to elliptic, cuspidate, caducous. Calyx 4 – 5-lobed, few hairs present, discoid glands sometimes present; flowering calyx 4 – 6  ×  3.5 – 4 mm; lobes acute, 0 – 1 mm long; fruiting calyx 7 – 8 mm diam., erect to patent. Corolla 4-lobed, covered with yellow hairs, sparsely glandular; anterior lip 3-lobed, mid-lobe oblong, 12 – 35  ×  10 – 15 mm long, apex rounded, patent, cuspidate, margins sometimes reflexed; side lobes 4 – 10  ×  5 – 8 mm, sometimes oblique, apex acute, erect to reflexed; posterior lip 1-lobed, 1.5 – 3.5  ×  1 – 6 mm, oblong, apex acute to rounded; tube 10 – 23 mm long. Stamens inserted at apex of tube, glabrous, sometimes with a few glands; anterior pair 10 – 20 mm long, anthers 2 – 3 mm long; posterior pair fertile, 2 (− 3); 8 – 11 mm long, anthers c. 1 mm long. Ovary glabrous with a few hairs at apex; style 20 – 32 mm long. Fruit (dried) 13 – 30  ×  8 – 30 mm, clavoid, glabrous, apex rounded, yellow when mature.

distribution. India and Sri Lanka east to Northern Thailand and North Vietnam (Map 2). Introduced and sometimes cultivated in gardens in Peninsular Malaysia (Kochummen 1978) and/or naturalised along roadsides. Cultivated throughout the tropics.
Map 2

Probable natural distribution of Gmelina asiatica (●); distribution of Gmelina australis (▲); Gmelina basifilum (■); Gmelina hollrungii (♦); and Gmelina leichhardtii (◘).

selected specimens examined. india. Bombay Presidency [Maharashtra State]: North Konkan, Karjat region, 10 Feb. 1949, Fernandes 88 (K); Tamil Nadu State: Attur distr., Salem, 30 March 1979, Perumal 22539 (K). MYANMAR. Upper Burma: 1888, Collet 554 (K). SRI LANKA. Yala: 13 June 1965, Worthington 7103 (K); Wilpattu National Park: Smithsonian Camp, Marai Villu, 30 June 1969, Wirawan, Cooray & Balakrishnan 899 (holotype NY, isotypes K, L). THAILAND. Lampoo [Lamphun Province]: Mae Tah Distr., Doi Kuhn Dahn Nat. Park, summit of Doi Hoa Chang, 3 June 1994, Maxwell 94676 (L); Prachinburi Province: Krabin, Aranya, 18 Oct. 1928, Put 2086 (holotype E, isotype K); Chiang mai Province: Payap, Chiengmai [Chiang Mai], 4 July 1922, Kerr 6224 (holotype E). VIETNAM. Cochinchina: 1862 – 1866, Thorel 60 (K); Tonkin: Bac Ninh, May 1891, Balansa 4972 (K).

habitat. In deciduous or evergreen forest, secondary and coastal vegetations. Soil sandy clay or a coarse gritty substrate often in poor soils, sometimes over granite or sandstone; alt. 0 – 1300 m.

conservation status. Least Concern.

phenology. Flowering and fruiting throughout the year.

vernacular names. Sri Lanka: Demata (Singhalese language), Kumil (Tamil language); Peninsular Malaysia: Bulang or Bulongan (Malay language). Cambodia: Köncaang; Laos: Phoung nou. China: Ya zhou shi zi. Philippines: Banganga (Cebuano language). Indonesia, Wareng (Javanese language), West Timor: Hau bako (Dawan language); Alor: Lombaul.

uses. The species has a wide range of medicinal uses (Burkill 1966).

notes. The type material of Gmelina asiatica forma lobata Moldenke, G. paniculata H. R. Fletcher and G. attenuata H. R. Fletcher fall within the total morphological variation of G. asiatica L. and is therefore placed into its synonomy.

3. Gmelina australisde Koksp. nov. a G. schlechteri bracteolis petiolatis maioribusque atque ad apicem rotundatis acutisve, calyce ad anthesin maiore et fructu ad apicem truncato differt. Typus: Australia, Northern Territory, c. 5 km E Nourlangie Rock, 10 Nov. 1972, Martensz 304 (holotypus K!; isotypi CANB, DNA, NT).

http://www.ipni.org/urn:lsid:ipni.org:names:77120319-1

[Munir (1984: 106 – 109 as Gmelina schlechteri p.p., Australian material only].

Tree 12 – 16 m high. Bark smooth to fissured, grey. Twigs densely hairy when young, interpetiolar ridge present, spines absent. Leaves elliptic to obovate, 18 – 27  ×  9 – 16 cm, apex rounded to acuminate, base rounded to cuneate, margin entire, sometimes slightly lobed; upper surface glabrous, green; lower surface glabrous with a few simple hairs on veins, peltate scales absent, pale green; two cup-like glands at the base; veins 5 – 7 pairs, sometimes 3-veined from base. Petiole 4 – 6.5 cm long, half-terete, a few simple hairs present. Inflorescence terminal, open, up to 20 cm long, with long side branches, velutinous; bracteoles lanceolate, 12 – 23  × 5 – 9 mm, apex rounded to acute, petiolate, with several disc-like glands present. Calyx 4 – 4.5  ×  6 – 7 mm, 5-lobed, sometimes appearing fewer, velutinous with sometimes a glabrous margin, outer surface with discoid glands; lobes up to 0.5 mm long, triangular; fruiting calyx c. 7 mm diam., erect. Corolla 5-lobed, outer surface hairy, sparsely glandular on lobes, pale lilac; anterior lip 3-lobed; mid-lobe oblong, 5 – 9  ×  4 – 5 mm, apex round, lilac, with two yellow marks; side lobes 4.5 – 5  ×  4.5 – 5 mm long, apex round; posterior lip, 2-lobed, 4.5 – 5  ×  6 – 7 mm, oblique, apex round; tube 8 – 11 mm long. Stamens inserted at apex of tube, glabrous, glands present; anterior pair 8 – 15 mm long, anthers c. 2 mm long; posterior pair, fertile, 6 – 10 mm long, anthers c. 1.5 mm long. Ovary glabrous with a few hairs at apex. Style 15 – 20 mm long, stigma c. 2 mm long. Fruit (dried) 14 – 17  ×  8 – 13 mm, clavate, apex truncated, glabrous, glossy, reddish purple when mature. Fig. 1.

distribution. Australia: Northern Territory restricted to the tropical forest of the Darwin region. Map 2.

specimens examined. australia. Northern Territory: c. 5 km E Nourlangie Rock, 10 Nov. 1972, Martensz 304 (holotype K); sandstone tableland between Gerowie Creek and Mary R., 4 Oct. 1946, Blake 17169 (K).

habitat. Along the fringes of rainforests, sometimes on black sandy soil; alt. c. 300 m.

conservation status. Least Concern.

phenology. Flowering Oct. – Nov.; fruiting Nov.

notes.Gmelina australis differences from G. schlechteri in having bracteoles which are petiolate, bigger, rounded to acute at the apex; a bigger calyx when in flower and a fruit with a truncated apex.

4. Gmelina basifilumde Koksp. nov. a Gmelina smithii ramulis lateralibus unflorescentiae longis (nec brevibus), filamentis ad basin (nec ad apicem) tubi corollae insertis et ovario ad apicem piloso (nec omnino glabro) differt. Typus: [Indonesia] Netherlands New Guinea, Geelvinkbaai, Japen Island, Soemberbaba, 2 July 1961, Koster BW 11115 (holotypus K!; isotypi L!, SING!).

http://www.ipni.org/urn:lsid:ipni.org:names:77120320-1

Trees 7 – 10 m high, DBH 7 – 10 cm. Bark whitish brown to grey. Wood (medium) hard, white to cream. Twigs hairy, spines absent. Leaves elliptic to lanceolate, 13 – 17  ×  5 – 8.5 cm, apex acute, base cuneate to rounded, margins entire; upper surface glabrous, dark green; lower surface glabrous with simple hairs on veins of young leaves, upper surface dark green; lower surface glaucous, covered with peltate scales, discoid glands absent; veins 7 – 10 pairs, sometimes 3-veined from base. Petiole half-terete, channelled, 3 – 5 cm, glabrous to sparsely hairy. Inflorescence terminal, 12 – 25 cm long, side branches up to 9 cm long, densely hairy, scales present; bracteoles ovate-lanceolate, 4 – 5  ×  2 – 2.5 mm, sessile, apex acute, caducous. Calyx 3 – 4.5  ×  3 – 4.5 mm, 5-lobed, velutinous when in flower, with a glabrous margin, glabrous when in fruit, surface covered with peltate scales, outer surface without discoid glands; lobes 0.8 – 1  ×  1 – 1.2 mm; apex acute to rounded; fruiting calyx 5 – 12 mm diam., glabrous, patent. Corolla 5-lobed, purple to white, outer surface velutinous, inside velutinous on lobes, glabrous in tube; anterior lip 3-lobed; mid-lobe 4 – 5  ×  4 – 4.5 mm, oblong, apex rounded; side lobes 3 – 4  ×  3 – 4 mm, apex rounded; posterior lip 2-lobed, 3 – 4  ×  3.5 – 4 mm, apex rounded; tube 2 – 5 mm long. Stamens inserted at base of tube, hairy at base; anterior pair 7 – 8 mm long, anthers c. 1.5 mm long; posterior pair, fertile, 6 – 7 mm long, anthers c. 1 mm long; ovary c. 2 mm diam., glabrous with tuft of hairs at apex. Style 6.5 – 7 mm long, stigma 1 – 1.5 mm long. Fruit (dried) 12 – 25  ×  13 – 30 mm, globose, apex rounded, glabrous, purple green to reddish purple when mature. Fig. 2.

distribution. The islands of New Guinea and New Britain. Map 2.

additional specimens examined. indonesia. New Guinea: Geelvinkbaai, Japen Island, Soemberbaba, 2 July 1961, Koster BW 11115 (holotype K, isotypes L, SING). PAPUA NEW GUINEA. Madang Province: Gogol R., 6 May 1970, Katik 46669 (K, SING); West New Britain Province: near Linga-linga, 29 May 1973, Henty & Lelean 49499 (K); Central Province: Sogeri, 6 May 1970, N.G.P. 4167 (SING).

habitat. Growing in primary and secondary forests, sometimes along rivers; alt. 0 – 500 m.

conservation status. This is a rare species with a very wide distribution in undisturbed forest, though it can also survive in secondary forests. Therefore, despite the limited number of collections known, I would provisionally propose a conservation assessment of Least Concern.

phenology. Flowering July; fruiting May – July.

notes. The specific epithet refers to the position at which the stamens are inserted, namely the base of the corolla tube, a character unique in the genus.

5. Gmelina chinensisBenth. (Bentham 1861: 272); Oliver (1889: t. 1974); Moldenke (1984f: 33 – 35); Shou-liang & Gilbert (1994: 33). Type: [China] Hong Kong, 1853 – 1856, Wright 490 (holotype K!; isotype NY scan seen, US scan seen).

Gmelina balansae Dop (1914: 322 – 323). Type: [Vietnam] Indo-Chine, Annam, a Hué, Harmand s.n. (syntype HM (n.v.); isosyntypes K!, P!); Laos, Phron-thane, Spire 233 (syntype B scan seen); [Vietnam] Tonkin, Bon 5413 (syntype ?); [Vietnam] Tonkin, Mont Bavi, Vallé du Lankok, 28 June 1887, Balansa 3806 (syntype HM (n.v.); isosyntype K!, L!, NY scan seen, P! × 2 sheets).

Gmelina lecomtei var. annamitica Dop (1933: 896); Vũ (2007: 143), synon. nov. Type: [Vietnam] Indo-Chine, Annam, a Hué, Harmand s.n. (holotype HM (n.v.); isotypes K!, P!).

Gmelina speciosa Moldenke (1940: 418 – 419), synon. nov. Type: [Vietnam] Tonkin, Mont Bavi, Vallé du Lankok, 28 June 1887, Balansa 3806 (holotype K!; isotypes HM (n.v.), NY scan seen, P!).

Tree or shrubs 1.5 – 20 m high, DBH 15 – 30 cm. Bark grey to grey-brown, finely fissured, wood yellowish. Twigs hairy when young, spines absent. Leaves ovate to elliptic, 5 – 18  ×  3 – 11 cm, apex acute, base cuneate, margins entire; papery; upper surface glabrous; lower surface glabrous to sparsely hairy, covered with peltate scales, a few discoid glands clustered around the base and veins, bluish green; veins 3 – 5 pairs, sometimes 3-veined from base, sometimes these prominent. Petiole terete, slightly channelled, 2 – 5.5 cm, glabrous. Inflorescence terminal, 9 – 20 cm, velutinous; bracteoles rounded to lanceolate, 8 – 15  ×  4 – 6.5 mm, sparsely hairy. Calyx 6 – 8  ×  8 – 8.5 mm, 5-lobed, sometimes appearing fewer, glabrous to sparsely hairy, covered with peltate scales, outer surface with few discoid glands, grey-brown; lobes triangular, 0.5 – 1  ×  2 – 4 mm, apex acute; fruiting calyx c. 14 cm diam. Corolla 5-lobed, yellow outside, white or pink-purple, inside, outer surface glabrous to sparsely hairy, often covered with peltate scales; inside glabrous; anterior lip 3-lobed; mid-lobe 11 – 17  ×  9 – 10 mm, apex acuminate, yellow; side lobes 9 – 16  ×  8 – 10 mm long, apex rounded; posterior lip, 2-lobed, 7 – 12  ×  7 – 10 mm, apex rounded to truncate; tube 17 – 22 mm long. Stamens inserted at apex of tube; anterior pair 13 – 20 mm long, anthers 2 – 2.5 mm long; posterior pair, fertile, 15 – 17 mm long, anthers 2 – 3 mm long. Ovary velutinous at apex. Style 25 – 27 mm long, stigma c. 1 mm long. Fruit (dried) 10 – 30  ×  6 – 15 mm, clavate, apex truncated, glabrous, apex covered with peltate scales, few discoid glands present.

distribution. South China, Laos and Vietnam south to Da Nang (see Map 3).
Map 3

Distribution of Gmelina chinensis (◘); Gmelina fasciculiflora (◊); Gmelina ledermannii (▲); Gmelina papuana (●); Gmelina schlechteri (♦).

specimens examined. china. Hong Kong: 1853 – 1856, Wright 490 (holotype K; isotype NY, US). Laulao Island, 10 Aug. 1888, Native collector 68/88 (K). LAOS. Khammouan: Nakai, 23 May 2006, Nanthavong BT 501 (L); Nakai distr., Ban Sop Phone, 8 Oct. 2006, Nanthavong BT 666 (L). VIETNAM. Mt Bani: 25 km from Tourane, May – July 1927, Clemens 3980 (K, L); Quang-nam: Da Nang, 1923, Poilane 7533 (K); Tonkin: Mont Bavi, Vallé du Lankok, 28 June 1887, Balansa 3806 (isosyntypes K, L, NY, P); Mont Bavi, Vallé du Lankok, 28 June 1887, Balansa 3806 (holotypes HM, K, L, P, NY). Annam, a Hué, Harmand s.n. (isosyntypes K, P); Laos: Phron-thane, Spire 233 (syntype B).

habitat. Growing in mixed forest on mountain slopes; alt. 500 – 1200 m altitude. Sometimes on poor laterite soils.

conservation status. Least Concern.

phenology. Flowering April – July; fruiting June – Oct.

vernacular names. China: Shi zi.

notes. The type material of Gmelina speciosa Moldenke and G. lecomtei var. annamitica Dop fall within the total morphological variation of G. chinensis Benth. and is therefore placed into its synonomy.

6. Gmelina dalrympleana(F. Muell.) H. J. Lam (1919: 223), Moldenke (1984f: 35 – 39); Munir (1984: 109 – 114); Vitex dalrympleana F. Muell. (Mueller 1864: 128). Type: [Australia, Queensland] Rockingham Bay, Dallachy s.n. (lectotype MEL 583504 scan seen; isolectotypes K! × 2 sheets), selected by Munir (1984: 109).

Ephielis simplicifolia Sol. ex Seem. (Seemann 1865: 258); Munir (1984: 110). Vitex macrophylla R. Br. (Brown 1810: 512), Gmelina macrophylla (R. Br.) Benth. (Bentham 1870: 65), nom. illegit., non Gmelina macrophylla Wall. ex Schauer (1847: 680)]; Munir (1984: 110). Type: Australia, Queensland, Cape Grafton, 1768 – 1771, Banks & Solander s.n. (holotype BM!).

Shrub or tree, 1 – 40 m high, DBH 5 – 70 cm. Bark fissured or flaky, greyish to grey-brown. Wood dark straw coloured with yellow streaks; twigs glabrous, spines absent. Leaves broadly ovate to ovate-oblong or elliptic-obovate, 6 – 37  ×  5 – 23 cm, margins entire, apex obtuse to obtuse acuminate, base cordate to cuneate; upper surface glabrous, dull or shiny (dark-olive) green; lower surface glabrous, peltate scales absent, with glands in a cup like structure at base, pale or greyish green; veins 4 – 8 pairs. Petiole 5 – 40 mm long, half-terete, slightly channelled, glabrous to hairy. Inflorescence terminal, 12 – 30 cm long, glabrous to hairy; bracteoles linear, 2 – 3  ×  1 – 1.5 mm, apex acute, caducous. Calyx 5-lobed, sometimes appearing fewer, glabrous, with black discoid glands; flowering calyx 4 – 6  ×  3 – 3.5 mm; lobes 0.5 – 1  ×  1.5 – 4 mm, apex round; fruiting calyx 6 – 7 mm diam., erect to patent. Corolla 5-lobed, velutinous, white, pink-yellow to pink or (pale) purple; anterior lip 3-lobed, mid-lobe oblong, 7 – 10  ×  5 – 6 mm long, apex rounded, dark pink to mauve with yellow spot at base; side lobes oblong-ovate, 5 – 9  ×  3.5 – 6 mm, apex rounded; posterior lip 2-lobed, oblong, 5 – 9  ×  4 – 5 mm, apex rounded; tube 10 – 22 mm long. Stamens inserted at apex of tube, glabrous, sometimes with a few glands, white; anterior pair 9 – 13 mm long, anthers c. 2 mm long; posterior pair, fertile, 7 – 9 mm long, anthers 1.5 – 2 mm long. Ovary 1 – 1.5 mm diam., glabrous, style 12 – 20 mm long, stigma c. 1.5 mm long. Fruit obovoid, 8 – 20  ×  5 – 25 mm, apex truncated, glabrous, pinkish to red when mature.

distribution. Australia: North Queensland and the island of New Guinea. Map 4.
Map 4

Distribution of Gmelina dalrympleana (■); Gmelina delavayana (▲); Gmelina elliptica (●) and Gmelina sessilis (▼).

selected specimens examined. australia. Queensland: Rockingham Bay, Dallachy s.n. (lectotype MEL; isolectotypes K); Cape Grafton, 1768 – 1771, Banks & Solander s.n. (holotype BM); Cairns: Woree, Bruce Highway, 10 March 1988, Gray 4766 (K). INDONESIA. Merauke area: Pr. Kurik, South Polder, 6 June 1961, Hoogerwerf 118 (L.); Koerik Camp, c. 15 km NE of Koembe village, 7 Sept. 1954, van Royen 4891 (L); North West Guinea: Djalan Kp. Keliki, 8 Aug. 1941, Anta (exp. Wentholt) 250 (L); Birds Head Peninsula: surroundings of Ayawasi, 16 March 1996, Ridsdale 2311 (L); surroundings of Ayawasi, 17 Sept. 1996, Wanda Ava 4800 (L); Nova Guinea Neerlandica Meridionalis: 29 Aug. 1907, Branderhorst 23 (K, L). PAPUA NEW GUINEA. Western Province: Daru Island, 21 Sept. 1972, Streimann & Lelean 18459 (K, L, SING); Oriemo R., Jan. 1959, White & Gray 10374 (K); Wassi Kussa R., Tarara, Dec. 1936, Brass 8539 (K, L).

habitat. Growing as a tree in rainforest to open forest, dunes or swamps (in Papua New Guinea sometimes associated with Melaleuca or Eucalyptus-Banksia woodland), sometimes a shrub in savanna woodlands; alt 0 – 450 m. Soil sometimes sandy.

conservation status. Least Concern.

phenology. Flowering Sept. – April; fruiting Feb. – Oct.

vernacular names. Australia: Dalrymple’s White Beech or Queensland Beech or Long-leaved Gmelina. Indonesia: Ara afa (Ayawasi).

uses. The wood is used in Australia for floorboards and planks.

7. Gmelina delavayanaDop (1914: 321); Moldenke (1984f: 41 – 43); Shou-liang & Gilbert (1994: 33). Type: China, Yunnan, Ta-pin-tze, Delavay 3595 (lectotype NY scan seen, designated here; isolectotype P scan seen).

Gmelina montana W. W. Sm. (Smith 1916: 107 – 108); Shou-liang & Gilbert (1994: 33). Type: [China] Yunnan, western flank of the Tali Range, Aug. 1913, Forrest 11662 (holotype E (n.v.); isotype K!).

Shrub 0.3 – 3 m high; twigs hairy when young, spines absent. Leaves ovate to elliptic, 1.5 – 5  ×  1.5 – 3.5 cm, apex acute, base cuneate, margins entire; sometimes slightly lobed, papery; upper surface glabrous; lower surface glabrous, sometimes with hairs on veins, covered with peltate scales; veins 3 – 4 pairs, sometimes 3-veined from base. Petiole 3 – 8 mm, half-terete, channelled, glabrous or with a few hairs, peltate scales absent to many. Inflorescence terminal, lax, 8 – 12 cm long, sparsely hairy, covered with peltate scales; bracteoles elliptic to lanceolate, 6 – 50  ×  1 – 35 mm, apex acute. Calyx 5-lobed, discoid gland present, 6 – 11  ×  6 – 7 mm; lobes 2.5 – 3.5 mm long, apex acute, not accrescent, erect. Corolla 4-lobed, sparsely covered with hairs, bluish to reddish purple, base yellow; anterior lip 3-lobed, mid-lobe rounded to spathulate, 5 – 15  ×  10 – 11 mm long, apex acute to round, patent; side lobes 5.5 – 15  ×  8 – 10 mm, rounded to spathulate, sometimes oblique, apex acute to rounded, erect to reflexed; posterior lip entire to slightly lobed, 8 – 18  ×  10 – 13 mm, oblong, apex rounded; tube 4 – 5 mm long, infundibulate. Stamens inserted at apex of tube, glabrous, sometimes with a few glands; anterior pair 14 – 16 mm long, anthers 2.5 mm long; posterior pair, fertile, 9.5 – 13 mm long, anthers c. 1 mm long. Ovary glabrous, sparsely glandular; style 17 – 20 mm long, glabrous. Fruit (dried) 6 – 7  ×  6 – 7 mm, globose, apex round, glabrous, black when mature.

distribution. China: endemic to South West Sichuan and Yunnan. Map 4.

specimens examined. china. Yunnan: June 1908, d’Alleizette s.n. (L); 1917 – 1919, Forrest 15620 (K); western flank of the Tali Range, Aug. 1913, Forrest 11662 (isotype K); Ta-pin-tze, 17 Aug. 1888, Delavay s.n. (P); Ta-pin-tze, Delavay 3595 (lectotype NY, isolectotype P); Ta-pin-tze, 1888, Delavay s.n. (P × 4); 13 June 1887, Delavay s.n. (P); 10 July 1885, Delavay s.n. (P); July 1746, Schneider 1746 (K); western flank of the Tali Range, Aug. 1913, Forrest 11662 (isotype K); NW Yunnan, N of Yung-peh, Forrest 22081 (K); W Yunnan, 1933, McLaren 250 (K). SW Szechuan: Sept. 1922, Forrest 22499 (K);

habitat. In thickets and along streams on mountain slopes; alt. 1500 – 3000 m.

conservation status. Least Concern.

phenology. Flowering May – July; fruiting July – Sept.

vernacular name. China: Xiao ye shi zi.

notes. Dop (1914: 321) cited a number of specimens in his original description of Gmelina delavayana: China, Yunnan, Ta-pin-tze, Delavay 170; China, Yunnan, Ta- pin-tze, Delavay 3595; China, Yunnan, Pint chouan, Ducloux 4698; China, Yunnan, Pint chouan, Ducloux 4707. From these, I select Delavay 3595 from NY as the lectotype as the specimen has a letter attached from Dop stating that this is G. delavayana.

This species seems to be morphologically close to Gmelina asiatica, G. elliptica and G. philippinensis. It shares with them their typical habit (a shrub) and the number of corolla lobes (four); it can be distinguished from them by the absence of spines, as well as by the different corolla shape and colour.

8. Gmelina ellipticaSm. in Rees (1810); Moldenke (1984f: 43 – 54); Munir (1984: 95–98); Rajendran & Daniel (2002: 161 – 166 as G. asiatica); Zhengyi & Raven (1988: 84 as G. asiatica). Type: East Indies, Anon. in Herb. Linn. 780–1 (holotype LINN!).

[Gmelina vestita Wall. (Wallich 1828: 50), nom. nud., synon. nov. Based on: [Myanmar] Kyouk Talong, Irrawaddy R., 28 Sept. 1826, Wallich 1820 (K-W!).]

Gmelina villosa Roxb. (Roxburgh 1814: 46, nom. nud.; 1832: 86); Lam (1919: 217 – 219); Gmelina asiatica var. villosa (Roxb.) Bakh. in Lam & Bakhuizen (1921: 70). Lectotype: Rumphius (1750: t. 39) selected by Rajendran & Daniel (2002: 166–168).

Gmelina villosa Naves (1877: t 215). Type: unknown.

Gmelina integrifolia Hunter ex Ridl. (Ridley 1909: 101 – 102); Lam & Bakhuizen (1921: 70). Type: [Penisular Maylasia, Penang], Prince of Wales Island, Hunter s.n. (holotype not located).

Gmelina tomentosa H. R. Fletcher (1938b: 204 – 205); Moldenke (1984h: 181 – 182), synon. nov. Type: Thailand, Rachasima, Ban Chum Seng, Korat, 23 May 1929, Nai Noe 211 (holotype E scan seen; isotype K!).

Gmelina tonkinensis Moldenke (1940: 419; 1984h: 182), synon. nov. Type: Vietnam, Tonkin, Fu Chap, to the rocks of Notre Dame, May or June 1887, Balansa 3807 (holotype LE (n.v.); isotypes K! × 2 sheets, L!, NY scan seen, P! × 4 sheets).

Shrubs to small trees, sometimes straggling, 1 – 5 (− 8) m high, DBH 2.5 – 20 (− 70) cm. Bark smooth, (pale) grey; wood hard. Twigs villous; spines usually present, up to 30 mm long. Leaves elliptic to ovate, rarely round, 2 – 9.5 (− 14)  ×  1.5 – 5 (− 8.5) cm, apex rounded to acuminate, base rounded to cuneate, margins entire to rarely 1 – 5-lobed; upper surface glabrous or with a few hairs on veins, green, pubescent when young; lower surface velutinous, hairs usually yellowish, covered with white peltate scale; sometimes with discoid glands at base; veins 3 – 4 pairs, sometimes 3-veined from base. Petiole 6 – 30 mm long, half terete, villous. Inflorescence terminal, 2.5 – 7 cm long, erect, villous; bracteoles lanceolate or ovate-lanceolate, 10 – 20  ×  (1.5–) 4 – 8 mm, apex acuminate, caducous. Calyx 5-lobed, sometimes appearing fewer, densely pubescent, becoming more glabrous over time, discoid glands present, peltate scales present; flowering calyx 3 – 5  ×  3 – 4 mm, lobes 0.5 – 1 mm long, apex rounded to acute; fruiting calyx 4 – 8 cm diam., margin undulated, patent. Corolla 4-lobed, covered with yellow hairs; anterior lip 3-lobed; mid-lobe 4.5 – 13  ×  4.5 – 14 mm, apex acute, patent, sometimes reflexed; side lobes 3.5 – 7.5  ×  3 – 7 mm, apex rounded to acute; posterior lip 5 – 9  ×  2 – 10 mm, apex acute to emarginate, patent to reflexed; tube 10 – 16 mm long, infundibulate. Stamens inserted at apex of tube, yellow; anterior pair 15 – 17 mm long, anthers 1.5 – 2.5 mm long; posterior pair, fertile, 10 – 18 mm long, anthers 1.5 – 2 mm long. Ovary glabrous; style 20 – 25 mm long, stigma c. 2.5 mm long. Fruit (dried) 10 – 20  ×  8 – 20 mm, globose to obovoid, glabrous, yellow when mature.

distribution. India: Andaman and Nicobar Islands; Myanmar to South China and Vietnam; south to Malaysia; Philippines: known from a number of islands: Basilan, Mindoro, Mindanao and Negros; Indonesia: from Sumatra to the Moluccas (Map 4). Cultivated throughout the tropics.

selected specimens examined. [East Indies], anon. in Herb. Linn. 780–1] (holotype LINN). CHINA. Yunnan: Kishuanbanna, 1980, Chow & Wan 80202 (K). INDIA. Nicobars: Feb. 1875, Kurz 26058 (K). INDONESIA. Amamba Islands: Terempak, 31 March 1928, Henderson 20147 (K, SING); Zuid Sumatra: Palembang, 31 Aug. 1947, de Raadt 21 (L); Sumatra: Simaloer, 13 Feb. 1918, Schmael 239 (L); Djambi, Doesoen Baroe, Sept. 1925, Posthumus 862 (L); North Sumatra, Medan, 27 Feb. 1928, Lörzing 12957 (K); East Java: Besuki, Blambangan Peninsula, 28 May 1957, Jacobs 4923 (L); Kalimantan: Barat, Gunung Palung Nat. Park, Dec. 1997, Laman et al. TL529 (K); East Borneo: Berau, Slopes of Mt Njapa on Kelai R., 13 Oct. 1963, Kostermans 21273 (L); SE Borneo, Hajoep, 2 June 1908, Winkler 2270 (L); Celebes: Kendari, Gunung Abesu, 6 Feb. 1986, Moh. Amir 70 (L), NE Celebes, Minahasa, Tasikoki, near Kema, SE of Mount Klabat, 5 July 1956, Forman 407 (L); Central Sulawesi: c. 25 km south of Palu, 21 April 1979, van Balgooy 2955 (L); Pulau Buton: Wakunti, Bau-bau, 26 June 1978, Widjaja 551 (L); East Bali: Kunu, 15 Oct. 1985, van Balgooy 5270 (L); East Sumbawa: Bay of Danggar, 22 Sept. 1984, Snellius-II 11151A (L); Lombok: N- Seite of Rindjani Vulkan gebirge, 30 April 1909, Elbert 750 (L); Flores, 16 Jan. 1973, Verheijen 3224 (L); West Timor: Usapi, 7 June 1981, Kooy 1302 (L); Alor Ketjil: Kabola peninsular, 4 May 1938, Jaag 469 (L); Ambon, 2 April 1918, Kornassi 1079 (L); West Ceram: 1918, Rutten 1667 (L). MALAYSIA. Penang: 1824, Phillips s.n. (K); Perak: Bota Kiri, near Ipoh, 11 March 1958, Shah 322 (K); Sabah: Keningau, Ulu Sungai. Pingas- pingas, 19 March 1988, Fidilis & Asik 122120 (K). MYANMAR. Toungoo Distr.: 11 Aug. 1911, Lace 5397 (K); Kyouk Talong: Irrawaddy R., 28 Sept. 1826, Wallich 1820 (K-W). PHILIPPINES. Basilan: 17 Jan. 1904, Hallier 4295a (L); Manaul: Mansalay, Mindoro, Dec. 1952, Sulit 17009 (K). SINGAPORE. near Changi Road, 13 Jan. 1933, Teruya 2163 (K). THAILAND. Phuket: near airport, 9 May 1968, van Beusekom & Phengkhlai 498a (K); Ratchasima: Pak Thong Chai, 13 May 1969, Phengnaren 639 (K); Siam: Kamboerie, Teysmann 5941 (K); Ban Chum Seng, Korat, 23 May 1929, Nai Noe 211 (holotype E; isotype K). VIETNAM. Tonkin: Fu Chap, to the rocks of Notre Dame, May or June 1887, Balansa 3807 (isotypes K, L, NY, P).

habitat. Growing in primary and secondary vegetations. Soil sandy or sandy loam or sandy clay, often poor, sometimes over basalt or limestone; alt. 0 – 600 m.

conservation status. Least Concern.

phenology. Flowering Jan. – Oct.; fruiting from Feb. – Oct.

vernacular names. Thailand: (Kang- or Nom-) Maow or Tam mio; Peninsular Malaysia: Bulangan, Bulang or Pokok Buah dulang (Malay language): Sabah: Bulangan (Malay Language), Kutang or Belingkot (Murut Language), Tangginang or Kutang (Dusun Language), Taring pelandok (Kadayan Language); Sumatra: Pakok boelang (Riau archipelago), Pokak baelandga (Pulau Lingga), Bariang (Simaloer island), Warengen (Medan); Wareng (Javanese language); Sulawesi: Pundanga (Palopo), Rerubo, Kayu palapi or Tanggalasi (Kendari), Bidara (Pulau Buton); Kayu palapi (Laweha), Wewenganga (Minhassa); Moluccas, Ambon: Karanjam, Alor Ketjil: Lombaul, South West Timor: Hau bako (Dawan language).

uses. In India, an infusion of the leaves is used for eye complaints (Rajendran & Daniel, 2002). In Peninsular Malaysia several medicinal uses are known (Burkill 1966).

notes. This species is recorded from a small area in Queensland, Australia (Munir 1984). The general distribution of the species goes as far south as the Flores and Sumba Islands (Map 4) and was not discovered in Australia until the 1970s. It is therefore very unlikely that this is a native of Australia. The fact that it is cultivated throughout the tropics and that it is becoming a pest problem in Queensland supports this assumption.

The differences between Gmelina elliptica and G. asiatica are usually easily to determine (G. elliptica has leaves which are densely villous beneath, and G. asiatica has either glabrous leaves or leaves with only a few hairs on the veins beneath). However, in Borneo, Sumatra, Java and the Lesser Sunda Islands there are specimens which seem to be morphologically intermediate; these specimens have lobed, somewhat hairy leaves with many peltate scales on the lower surfaces, the typical form having leaves with an entire margin, densely hairy and few peltate scales on the lower surfaces. On the specimen Lörzing 12957, the typical form for G. elliptica and the intermediate form are both present, and in the remark on the label, he states ‘Young shoots differ especially in their leaves’. This is confirmed by some of the other specimens with apparently intermediate leaf morphology: in these the bigger leaves are more hairy and less lobed and are, as such, more typical for G. elliptica than the smaller leaves. Given that typical G. asiatica does not naturally occur on these islands and that G. elliptica is native to these areas, I have concluded that these intermediate specimens should be seen as part of G. elliptica. More research is needed to determine whether these forms represent either hybrids between G. elliptica and the introduced G. asiatica or a natural variation within G. elliptica. This variation is one of the main reasons why G. asiatica and G. elliptica are sometimes seen as two forms of one species.

I have not seen any type material of Gmelina integrifolia Hunter ex Ridl. (Ridley 1909: 101 – 102), but from the description it is clear that it is part of the variation of G. elliptica. The type material of G. tonkinensis Moldenke, G. tomentosa H. R. Fletcher and G. vestita Wall. fall within the total morphological variation of G. elliptica Sm. and is therefore placed into its synonomy.

9. Gmelina evoluta(Däniker) Mabb. (Mabberley 1998: 316). Vitex evoluta Däniker (1933: 408). Type: New Caledonia, Koumac, foot of Mt Kaala, 25 Feb. 1925, Däniker 1228 (holotype Z scan seen; isotype Z × 2 sheets, scan seen).

conservation status. Least Concern.

note. For a description and more information see Mabberley in Mabberley & de Kok (2004).

10. Gmelina fasciculifloraBenth. (Bentham1870: 65); Moldenke 1984g: 106 – 107). Vitex leichhardtii var. glabrifolia F. Muell. (Mueller 1865: 35), Benth. (Bentham 1870: 65 as ‘glabrata’). Type: Australia, Queensland, Rockingham Bay, 16 Oct. 1868, Dallachy s.n. (lectotype MEL 583159 scan seen; isolectotypes K! × 2 sheets) selected by Munir (1984: 102).

Trees (4 –) 10 – 30 m high, DBH 12 – 80 cm. Bark somewhat scaly, grey. Twigs tomentose when young, hairs brown, spines absent. Leaves ovate-elliptic, 3 – 16  ×  2.5 – 9 cm, apex obtuse to obtusely acuminate, base rounded to cuneate, margin entire, coriaceous; upper surface shiny; lower surface covered with peltate scales; veins 4 – 7 pairs, sometimes 3-veined from base. Petiole half-terete, channelled, 5 – 30 mm long, glabrous. Inflorescence terminal, densely tomentose, 10 – 25 cm long; bracteoles clavoid to round, 2.5 – 5  ×  2 – 4 mm, sessile. Calyx 5-lobed, green, tomentose outside, hairs rusty brown; flowering calyx 2.5 – 4  ×  2.5 – 3.5 mm; lobes triangular, 0.4 – 0.6  ×  0.5 – 1 mm, apex acute, erect; fruiting calyx 7 – 15 mm diam. Corolla 5-lobed, tomentose when young, becoming less so with age, white to blue; anterior lip 3-lobed, mid-lobe ovate to broadly elliptic, 5 – 10  ×  5 – 8 mm, apex rounded to acute, purple with a yellow marking at base; side lobes 4 – 8  ×  3 – 6 mm, oblong- ovate long, apex rounded; posterior lip 2-lobed, 3 – 7  ×  3 – 6 mm; tube 8 – 14 mm long. Stamens inserted at apex of tube, few glandular hairs; anterior pair 10 – 11 mm long, anthers c. 1.2 mm long; posterior pair, fertile, 8 – 9 mm long, anthers c. 1 mm long. Ovary 1 – 2 mm diam., glabrous, villous at apex. Style 12 – 15 mm long, stigma c. 3 mm long. Fruit globular, 10 – 20 mm diam., apex rounded glabrous, smooth, shiny, glands absent, violet or blue to bright purple or violet when mature, taste sour.

distribution. Australia: Northern Queensland. Map 3.

selected specimens examined. australia. Queensland: Julatten, 11 Jan. 1977, Gray 222 (K); Boonjee, 13 Dec. 1979, Gray 1591 (K); Barron, 15 Nov. 1972, Hyland 6486 (K); Mission Beach, 58 Miles SE of Atherton, 14 Nov. 1963, Hyland 3720 (K); Rockingham Bay, 16 Oct. 1868, Dallachy s.n. (lectotype MEL; isolectotype K, P).

habitat. Growing in rainforests or along rivers and beaches; alt. 0 – 760 m.

conservation status. Least Concern.

phenology. Flowering Nov. – Dec.; fruiting Nov. – Jan.

vernacular names. North Queensland Beech or Grey Teak.

uses. The wood is used in furniture making and woodcarving.

11. Gmelina hollrungiide Koksp. nov. a G. schlechteri glandulis disciformibus e bracteolis carentibus differt. A G. evoluta glandulis cupuliformibus ad basin foliorum dispositis differt. Typus: [Papua New Guinea] New Guinea, Augusta Station, 1887, Hollrung 651 (holotypus K!; isotypi P!).

http://www.ipni.org/urn:lsid:ipni.org:names:77120321-1

Gmelina macrophylla Schumann in Schumann & Hollrung (1889: 120), nom. illegit., non Gmelina macrophylla Wall. ex Schauer (1847: 680). Type: [Papua New Guinea] New Guinea, Augusta Station, 1887, Hollrung 651 (holotype B (n.v.); isotypes K!, P! × 2 sheets).

Trees, 6 – 40 m high; DBH 30 – 75 cm. Bark smooth to fissured, grey to brown; Twigs velutinous, spines absent. Leaves elliptic-round, oblong or obovate, 10 – 36 × 5 – 25 cm, apex rounded to shortly acuminate, base rounded to cuneate, margins entire, coriaceous; upper surface glabrous, few hairs on veins, (yellow) green, shiny; lower surface glabrous, pale green to yellow, few hairs on veins, peltate scales absent, pale beneath, with cup-like glands at base; veins 5 – 8 pair, sometimes 3-veined from base. Petiole 15 – 80 mm long, half-terete, channelled, sparsely hairy. Inflorescence terminal, 15 – 30 cm long, brown tomentose; bracteoles linear, 2.5 – 4  ×  0.5 – 1 mm long, apex acute, sessile. Calyx 5-lobed, sparsely to densely hairy; with scattered black discoid glands; flowering calyx 3 – 5  ×  3 – 4 mm; lobes 0.8 – 1  ×  1 – 1.2 mm, apex rounded; fruiting calyx up to 8 mm diam., patent. Corolla 5-lobed, velutinous, white, pale lilac to mauve or purple; anterior lip 3-lobed, mid-lobe oblong, 5.5 – 10  ×  3 – 4 mm, apex rounded to acute, lilac with a yellow spot at base; side lobes oblong, 3.5 – 8  ×  2 – 5 mm, apex rounded to acute, oblique; posterior lip 2-lobed, oblong to ovate- oblong, 3 – 7  ×  2 – 5 mm; tube 7 – 12 mm long. Stamens inserted at apex of tube, few glands; anterior pair 6 – 10 mm long, anthers 2 – 2.5 mm long; posterior pair, fertile, 6 – 8 mm long, anthers c. 2 mm long. Ovary 1 – 1.5 mm diam., glabrous with some glands at apex, style 12 – 20 mm long. Fruit (dried) 8 – 25  ×  6 – 16 mm, obovoid, apex truncated, glabrous, shiny, blue or red to purple when mature. Fig. 3.

distribution. Indonesia: Moluccas: Morotai and Aru Islands, West Papua; Papua New Guinea; Northern Australia; the Solomon Islands. Map 2.

selected specimens examined. australia. Port Essington, Armstrong 556 (K); Rockingham Bay, Dallachy s.n. (K). INDONESIA. Moluccas: Morotai, Mt Permatang, 23 May 1949, Main & Aden 941 (K); Aroe Islands, Pulau Trangan, 1 July 1938, Buwalda 5431 (K, L); Aru Archipelago, Kobroor, 25 Oct. 1994, Nooteboom 5880 (K, L); Aru Islands, Pulau Trangan, 23 Oct. 1994, van Balgooy 6610 (K); West Papua: Sorong, 1 Oct. 1948, Pleyte 1094 (K, L). PAPUA NEW GUINEA. Augusta Station, 1887, Hollrung 651 (holotype K; isotypes K, P); Morobe Province: Lae, Saru R., 20 July 1970, Streimann et al. 47995 (SING); Wampit, Bupu Village, 13 July 1967, Millar 22979 (L); Central Province: Brown R., Dec. 1963, Kumul 13071 (K); Western Highlands Province: Hagen distr., Road above Baptist Mission, 9 July 1968, Miller 36668 (L); Baiyer R. Sanctuary, 5 June 1980, Wiakabu 73468 (K, L); East Sepik Province: Hunstein range, Mt Samsai, 24 July 1990, Takeuchi 6373 (K); Northern Province: Isuarava, 24 Feb. 1936, Carr 15748 (L, SING). SOLOMON ISLANDS. Santa Ysabel, 21 Sept. 1965, Hunt 2649 (K).

habitat. Growing in rainforest, sometimes as canopy tree, or in more open types of forest; alt. 30 – 1400 m. Soil sandy or clayey, sometimes over sandstone, limestone or ultra-basic rock.

conservation status. Least Concern.

phenology. Flowering July – Nov.; fruiting July – Feb. Fruit eaten by cassowaries.

vernacular names. Indonesia: Adoen or Adun mangai laplabai (Aru Islands); Anjoes (Manokwari, Warnapi).

uses. On the Aru Islands the species is considered a good tree for timber.

notes.Gmelina hollrungii differs from G. schlechteri in having bracteoles without disc-like glands and differs from G. evoluta in having cup-like glands at the base of the leaves.

12. Gmelina lecomteiDop (1914: 322); Vũ (2007: 141 – 142). Type: [Vietnam] Indo-Chine, Annam, Chapa à Muong-Xen, 31 Oct. 1911, Lecomte & Finet 421 (holotype P scan seen; isotype HM (n.v.), K!).

Shrub to trees 3 – 7 m high, DBH c. 10 cm. Twigs densely hairy when young, spines absent. Leaves ovate to elliptic, 4 – 13  ×  3 – 12 cm, margin enire, sometimes slightly lobed, apex acute to acuminate, base slightly cordate or cuneate to slightly decurrent, papery; upper surface glabrous, with few hairs on veins at base, bluish green; lower surface velutinous, hairs usually yellow, covered with peltate scales, few discoid glands present at base; veins 3 – 6-pairs, sometimes 3-veined from base. Petiole 1.5 – 7 mm, terete to half-terete, channelled, velutinous. Inflorescence terminal, 5 – 12 cm long, velutinous to sparsely hairy, covered with peltate scales; bracteoles 7 – 13  ×  5.5 – 10 mm, elliptic, margin slightly lobed, sparsely hairy, covered with peltate scales, often with discoid glands present. Calyx 12 – 15  ×  11.5 – 15 mm, 5-lobed, covered with peltate scales, sparsely hairy, often with discoid glands present, not accrescent; lobes triangular, 4 – 7  ×  5 – 7 mm, anterior lobes biggest, apex acute, erect. Corolla 5-lobed, covered with peltate scales, white to pink, odourless to fragrant; anterior lip 3-lobed, mid-lobe rounded, 5 – 6  ×  7 – 6 mm long, apex, patent; side lobes oblong, 4 – 5  ×  6 – 7 mm, apex rounded, erect; posterior lip 2-lobed, 4 – 9  ×  7 – 10 mm, oblong, apex rounded; tube 10 – 14 mm long, infundibular. Stamens inserted at apex of tube, glabrous, sometimes with a few glands; anterior pair c. 10 mm long, anthers c. 3 mm long; posterior pair, c. 7 mm long, anthers c. 2 mm long. Ovary pubescent at apex, covered with peltate scales at apex; style c. 10 mm long, glabrous. Fruit (dried) cylindrical, 15 – 40  ×  10 – 12 mm, apex truncate, sparely hairy, covered with peltate scales, discoid glands present, grey.

distribution. China: Yunnan, Kwantung and Hainan; Vietnam and Laos. Map 5.
Map 5

Distribution of Gmelina lecomtei (●); Gmelina lepidota (▲); probable natural distribution of Gmelina philippinensis (■) with grey-shaded square as a questionable data point; distribution of Gmelina peltata (▼).

specimens examined. china. Kwantung: Sin-fung distr., Fuk Lung monastery, Hau T’ong Shan, 1 – 19 June 1938, Taam 808 (K); Sin-fung distr., Ch’a-P’ing village, Ah P’o Kai Shan, 1 – 24 May 1938, Taam 635 (K); Fang Ch’eng distr., T’aan Faan, Kung P’ing Shan and vicinity, 15 – 24 Aug. 1936, Tsang 26664 (K); Fang Ch’eng distr., T’aan Faan, Kung P’ing Shan and vicinity, 15 – 24 Aug. 1936, Tsang 26666 (K); Hainan: Ngai Distr., Yeung Ling, Shan, 13 June 1932, Lau 75 (K); Lingshin, 25 April 1932, Ko 52188 (K); Yaichow, 8 July 1933, Liang 61985 (K). LAOS. Khammouan Nakai Distr.: 19 May 2006, Newman et al. LAO 1154 (L). VIETNAM. Annam [Central region]: Chapa à Muong-Xen, 31 Oct. 1911, Lecomte & Finet 421 (holotype P); Tonkin: Ha-coi, near Chuk-phai, Taai Wong Mo Shan, 3 May – 22 June 1939, Tsang 29235 (K, L); Dam-ha, Lung Wan Village, Sai Wong Mo Shan, 18 May – 5 July 1940, Tsang 29869 (K, SING); Tien-yen, Kau Nga Shan and vicinity, 23 Sept. – 7 Oct. 1940, Tsang 30588 (K, SING); Ha-coi, Tong Fa market, Taai Wong Mo Shan and vicinity, 11 – 23 Sept. 1939, Tsang 29546 (K, SING); Bankeuin, near Quang Yen, Aug. 1885, Balansa 937 (K).

habitat. In open forest and thickets, in dry sandy or clayey soil; alt. 200 – 1000 m.

conservation status. Least Concern.

phenology. Flowering Aug. – Sept.; fruiting June – Aug.

vernacular name. China: Yue nan shi zi.

notes. This species is often confused with Gmelina racemosa and the other Indo-China endemics; the differences are given in Table 1.
Table 1.

Differences between Gmelina chinensis, G. lecomtei and G. racemosa.

 

G. lecomtei

G. lecomtei

G. chinensis

Lower leaf surface

hairy

glabrous

hairy to glabrous

Calyx lobes

present

present

minute

Bracteoles (mm)

elliptic, 7 – 13 × 5.5 – 10

lanceolate, 7.5 – 25 × 2.5 –10

lanceolate, 8 – 15 × 4 – 6.5

Bracteole margin

slightly lobed

entire

entire

Corolla (anterior lip length, mm)

small, (5 – 6)

big, (5 – 10)

big, (11 – 17)

Ovary apex

sparsely hairy

velutinous

glabrous

13. Gmelina ledermanniiH. J. Lam (1919: 226); Moldenke (1984g: 109 – 110); Type: [Papua New Guinea] New Guinea, Malu, near Sepik-river, 3 March 1912, Ledermann 6537 (lectotype K!), selected here.

Tree 10 – 20 m high. Bark dark, rough. Twigs sparsely hairy when young, with interpetiolar ridge present, spines absent. Leaves elliptic to slightly obovate, 8 – 19  ×  5.5 – 9.5 mm, slightly oblique, apex rounded to acute, base rounded, rarely cuneate, margins entire; upper surface glabrous, dull to glossy, dark green; lower surface glabrous with few simple hairs on veins, hairy when young, pale green, peltate scales absent; cup-like glands at base; veins 6 – 8 pairs, sometimes 3-veined from base. Petiole 2 – 4 cm long, half-terete, sparsely hairy. Inflorescence terminal, narrow, up to 20 cm long, with long side branches, velutinous; bracteoles rounded (in particular on main axis) to lanceolate, 4 – 11  ×  4.5 – 6.5 mm, apex long acuminate, sessile, with several disc-like glands present. Calyx 3.5 – 5  ×  3.5 – 4 mm, 5-lobed, sometimes appearing fewer, velutinous sometimes with a glabrous margin, outer surface with discoid glands; lobes up to 0.5 mm long, apex acute to rounded; fruiting calyx 5 – 6 mm diam., erect. Corolla 5-lobed, outer surface hairy, densely glandular on inside of lobes, cream to yellow; anterior lip 3-lobed; mid-lobe oblong, 7.5 – 10  ×  3 – 5.5 mm, apex rounded, mauve with two yellow marks; side lobes 4.5 – 5  ×  4.5 – 5 mm long, apex rounded; posterior lip, 2-lobed, 4.5 – 5  ×  3.5 – 4 mm, apex rounded; tube 6.5 – 7.5 mm long. Stamens inserted at apex of tube, glabrous, glands present; anterior pair 8 – 11 mm long, anthers c. 2 mm long; posterior pair, fertile, 6 – 8 mm long, anthers c. 1.5 mm long. Ovary glabrous with a few hairs at apex. Style c. 10 mm long, stigma c. 1.5 mm long. Fruit (dried) 10 – 11  ×  5 – 6 mm, tubular, apex rounded, glabrous, smooth, glossy, purple or bright blue when mature.

distribution. Indonesian New Guinea and Papua New Guinea: Sandaun and east Sepik Province (see Map 3).

specimens examined. indonesia. New Guinea: Mimika, Najaja (Oeta), 21 June 1941, Neth. Ind. For. Service bb. 32852 (Lundquist 133) (K). PAPUA NEW GUINEA. Morobe Province: Dschischugari, 25 May 1909, Schlechter 19566 (K); East Sepik Province: New-Guinea, Malu, near Sepik-R., Ledermann 10455a (K); Malu, near Sepik-R., 3 March 1912, Ledermann 6537 (lectotype K); along Yapa (Hunstein R.), 1 Aug. 1966, Hoogland & Craven 10799 (K, L); near Ambunti, 31 May 1966, Hoogland & Craven 10167 (K, L); Sandaun Province: Telefomin distr., W of Fiak airstrip, 27 March 1992, Frodin et al. 2586 (K); Papua, Eastern Division, Asia R., 11 May 1926, Brass 1376 (K).

habitat. A tree in stunted or secondary forest, sometimes along rivers, in sandy or poor soil; alt. 90 – 800 m.

conservation status. Least Concern.

phenology. Flowering May – Aug.; fruiting March – Aug.

vernacular names. Papua New Guinea: Yoni (Wagu Language); Bouwen or Boing (Waskuk Language) or As Kobokim (Miyanmin Language). Indonesia: Boetao (Mimika area).

uses. Wood used for construction and canoe building (as it is easily worked) and firewood.

notes. In the originally description three specimens were mentioned: Ledermann 6537 and 10455a and Schlechter 19566. The Ledermann 6537 specimen at K is selected here as the lectotype as it is the only one with mature flowers.

14. Gmelina leichhardtii (F. Muell.) Benth. (Bentham 1870: 66); Moldenke 1984g: 110 – 115); Munir (1984: 99 – 102); Vitex leichhardtii F. Muell. (Mueller 1862: 58). Type: [Australia] New South Wales, Myall Creek, 30 Nov. 1843, Leichhardt s.n., (lectotype MEL scan seen), selected by Munir (1984: 99).

Tectona australis W. Hill (1862: 20), synon. nov. Type: [Australia] Queensland Woods, [London exib. 1862] Hill 30 (holotype K!).

Tree 10 – 40 m high, DBH up to 120 cm, often semi-deciduous. Bark flaky or fissured, (dark) grey. Twigs tomentose, light brown, spines absent. Leaves narrowly elliptic or ovate-elliptic, 6 – 15  ×  4 – 9 cm, apex rounded to acute, papery, base rounded to cuneate, margins entire, upper surface glabrous, sometimes with hairs on veins, dark green; lower surface densely tomentose, peltate scales present, basal glands absent; green; veins 7 – 8 pairs, sometimes 3-veined from base, lower surface with strong ladder-like tertiary venation. Petiole 15 – 55 mm, half-terete, channelled, tomentose. Inflorescence 10 – 25 cm long, tomentose; bracteoles linear, 1.5 – 2  ×  1 – 1.2 mm, apex acute, caducous. Calyx 5-lobed, sometimes appearing fewer, pubescent, black discoid glands present; flowering calyx 4 – 6 × 3 – 6 mm; lobes triangular, up to 0.2 mm long, apex acute; fruiting calyx 10 – 20 mm diam. Corolla 5-lobed, tomentose outside, glandular hair inside, (cream-) white; anterior lip 3-lobed; mid-lobe spathulate to ovate, 8 – 13  ×  7 – 10 mm, apex rounded, violet with two yellow spots at base; side lobes elliptic-oblong, 5 – 10  ×  5 – 8 mm, apex rounded, sometime oblique; posterior lip, 2-lobed, lobes oblong, 6 – 9  ×  5 – 6 mm, apex rounded; tube 4 – 8 mm long, with two bluish stripes ending in yellow spots at the filament insertion. Stamens inserted at apex of tube, covered with glandular hairs; anterior pair 12 – 14 mm long, anthers 1.5 – 2 mm long; posterior pair, fertile, 10 – 11 mm long, anthers 1.2 – 1.5 mm long. Ovary 1 – 2 mm diam., sparsely pubescent. Style 10 – 16 mm long, stigma c. 1.5 mm long. Fruit 11 – 20  ×  13 – 15 mm, almost globular, apex rounded, glabrous, bluish-purple when mature, reported to have a disagreeable taste.

DISTRIBUTION. Australia: from the coast of central Queensland to the south coast of New South Wales. Map 2.

selected specimens examined. australia. Queensland: Queensland Woods, [London exib. 1862] Hill 30 (holotype K); South Kennedy, NE of Eungella, 26 Nov. 1989, McDonald 4471 (K); Brisbane Botanic Gardens, 31 Oct. 1930, Hubbard 4738 (K); Dalrymple Heights and vicinity, July – Nov. 1947, Clemens s.n. (K); New South Wales: Botanic Gardens Sydney, Oct. 1908, Boorman s.n. (L); Sydney Woods, 1854, MacArthur 193 (K); New Holland, Morton Bay, 1845, Leichhardt s.n. (P); Myall Creek, 30 Nov. 1843, Leichhardt s.n., (lectotype MEL).

habitat. Growing in primary and secondary vegetations.

conservation status. Least Concern.

phenology. Flowering Oct. – Jan.; fruiting Jan. – April.

vernacular name. White or Native Beech.

uses. The wood is used in furniture making and wood carvings. The species has been commercially overexploited in the past (Francis 1951).

notes. The type material of Tectona australis W. Hill falls within the total morphological variation of Gmelina leichhardtii (F. Muell.) Benth. and is thefore placed into its synonomy.

15. Gmelina lepidotaScheff. (Scheffer 1876: 41 – 42) Moldenke 1984g: 115 – 116). Type: [Indonesia, New Guinea] L’Ile de Tow, 18 Aug. 1871, Teysmann 6744 (holotype BO!; isotypes L!, NY scan seen).

Gmelina misoolensis Moldenke (1952: 54; 1984g: 118), synon. nov. Type: [Indonesia] New Guinea, Misool Island, Fakal, 30 Sept. 1948, Pleyte 1087 (holotype BO!; isotypes BO! L! NY scan seen).

Gmelina lepidota var. lanceolata Moldenke (1958: 325 – 326; 1984g: 116), synon. nov. Type: Papua New Guinea, New Britain, Open Bay, Nantambu, June 1945, Mair 1894 (holotype LAE (n.v.); isotypes L! × 2 sheets).

Trees 7.5 – 30 m high, DBH 3 – 60 cm. Bark (dark) grey-brown, fissured, wood light straw coloured. Twigs hairy when young, glabrous later, spines absent. Leaves elliptic to lanceolate, rarely ovate, 4 – 18  ×  2.3 – 7 cm, apex rounded to acuminate, base cuneate, margins entire; upper surface glabrous, dark green; lower surface glabrous with a few hairs on the veins, lightly brownish green or dull grey brown, covered with peltate scales, with discoid glands clustered around the base and along the vein; veins 4 – 7 pairs. Petiole half-terete, slightly channelled, 1 – 3 cm, sparsely covered with peltate scales. Inflorescence terminal and axillary, with short side cymes, 4 – 7 cm long, sparsely hairy; bracteoles lanceolate, 0.8 – 2  ×  0.1 – 1.5 mm, apex acute, persistent. Calyx 5-lobed, glabrous, covered with peltate scales, olive, greenish purple or purple-brown, outer surface with discoid glands; flowering calyx 2 – 3.5  ×  2 – 3 mm; lobes triangular, 0.5 – 0.8 mm long, apex acute; fruiting calyx 5 – 6 mm diam., patent. Corolla 5-lobed, outer surface velutinous, inside glabrous, sparsely glandular, cream to yellow or (light) purple; anterior lip 3-lobed; mid-lobe spathulate 4.5 – 5  ×  3 – 4 mm, apex rounded; side lobes 1.5 – 3  ×  2 – 2.5 mm, apex rounded; posterior lip 2-lobed, 2 – 4  ×  1.5 – 2 mm, apex rounded; tube 8 – 9 mm long. Stamens inserted at apex of tube, white, base covered with hairs, mauve; anterior pair 6 – 10 mm long, anthers c. 1 mm long; posterior pair, fertile, 5 – 8 mm long, anthers 0.8 mm long. Ovary c. 1 mm diam., glabrous; style 8 – 11 mm long, stigma c. 1.5 mm long. Fruit (dried) 9 – 20  ×  6 – 25 mm, ellipsoid to clavoid, apex rounded, glabrous, red or pale purple when mature.

distribution. Indonesia: Moluccas: Morotai Island; the islands of New Guinea and New Britain. Map 5.

selected specimens examined. indonesia. Moluccas: Morotai, Gunung Pare Pare, 28 May 1949, Main & Aden (exp. Kostermans) 1292 (K, L, SING); Morotai, Gunung Pare Pare, 28 May 1949, Main & Aden (exp. Kostermans) 1276 (K, L, SING); Obi Island: Jikodolong, 26 Nov. 1974, de Vogel 4342 (L); [Indonesia, New Guinea] L’Ile de Tow, 18 Aug. 1871, Teysmann 6744 (holotype BO!; isotype L); Misool Island, Fakal, 30 Sept. 1948, Pleyte 1087 (holotype BO; isotypes BO, L, NY); West Papua: Nederland’s New Guinea, Radjah Ampat, Weigeo Island, E bank of Majalibit Bay, Waifoi, 18 Jan. 1955, van Royen 5222 (K, L); Vogelkop Peninsula, Ajamaroe, 8 May 1958, Versteegh BW 4998 (L); New Guinea, Hollandia, 21 Dec. 1954, Versteegh BW 658 (L); West Irian, near Sukarnapura [= Djajapura], 4 August 1966, Kostermans & Soegeng 148 (K); West Irian, in town of Sukarnapura [= Djajapura], 7 Aug. 1966, Kostermans & Soegeng 230 (K); West Irian, above Hollandia [ = Djajapura], 15 Aug. 1966, Kostermans & Soegeng 369 (K); Netherlands New Guinea, Hollandia [ = Djajapura], 17 Jan. 1957, Versteegh BW 4696 (SING). PAPUA NEW GUINEA. Western Province: near Lake Daviumbu, 4 Oct. 1967, Pullen 7436 (L); New Britain Province: Mt Tangis, Western Slopes, 29 May 1966, Frodin 26866 (L); New Britain, Open Bay, Nantambu, June 1945, Mair 1894 (isotypes L); Southern Highland Province: Lake Kutubu, near Moro, 4 Oct. 1961, Schodde 2368 (L); Morobe Province: Sikong Range, Poiyu road, 2 Jan. 1989, Takeuchi et al. 4342 (L).

habitat. Growing from rainforest to xerophytic or open vegetation, sometimes dominated by Myrtaceae (notably Eucalyptopsis papuana C. T. White); alt. 10 – 1000 m. Clayey soil, sometimes over limestone, serpentine, or volcanic soils.

conservation status. Least Concern.

phenology. Flowering and fruiting Aug. – May.

vernacular names. Indonesia: Rafah (Maibrat Language). Papua New Guinea, New Britain: Namavue (Mt Tangis); Kanike’hua (Kutubu Language).

uses. The wood is used for dugout canoes.

notes. Type material of Gmelina lepidota var. lanceolata Moldenke and G. misoolensis Moldenke fall within the total morphological variation of G. lepidota Scheff. and is therefore placed into its synonomy.

16. Gmelina lignum-vitreumGuillaumin (1952: 539); Moldenke (1984g: 116 – 118); Mabberley in Mabberley & de Kok (2004). Type: New Caledonia, Forêt de Thy, Aug. 1949, Sarlin 81 (holotype P scan seen; isotypes P × 2 sheets, scans seen).

conservation status. Listed as Critically Endangered (IUCN red list download on 7 Jan. 2011 (www.iucnredlist.org/apps/redlist)).

note. For a description and more information see Mabberley in Mabberley & de Kok (2004).

17. Gmelina magnificaMabb. in Mabberley & de Kok (2004: 23 – 26). Type: New Caledonia, Pouébo, 24 July 1974, MacKee 28975 (holotype P, scan seen; isotype P, scan seen).

conservation status. Least Concern.

note. For a description and more information see Mabberley in Mabberley & de Kok (2004).

18. Gmelina moluccana(Blume) Backer ex K. Heyne (1917: 118); Vitex moluccana Blume (1826: 813 – 814); Moldenke (1984g: 118 – 126); Merrill (1917: 452) Type: Rumph. Herb. Amb. T. 3 t 20, selected here [Tittius alba Rumph. (Rumphius 1750: 38 – 39)] [Tittius rubra Rumph. (1750: 38 – 39)].

Volkameria obovata Roxb. (Roxburgh 1814: 95, nom. nud.; 1832: 62); Clerodendrum bovatum (Roxb.) Walp. (Walpers 1844 – 1848: 112), synon. nov. Type: [Indonesia] Moluccas, n.d., anon. (holotype BR, scan seen).

Gmelina macrophylla Wall. ex Schauer (1847: 680); Lam (1919: 220 – 221), synon. nov. Type: H. B. C. [Hortus Botanicus Calcuta], 1826, ex Amboyna [ = Indonesia, Ambon], Wallich 1819 (holotype G (n.v.); isotype K-W!).

Gmelina glandulosa Hallier f. (1918: 57), Moldenke (1984g: 121); Gmelina moluccana forma glandulosa (Hallier f.) Moldenke (1984g: 121, nom. nud.).

Gmelina moluccana forma glandulosa (Hallier f.) Moldenke (1984h: 155), synon. nov. Type: [Indonesia, Molucas] Banda-gruppe, Ay Insel, June 1821, Reinwardti 1362 (syntypes L!); [Indonesia, Molucas] Insel Neira, May 1859, de Vriese s.n. (syntype L!).

Gmelina sessilis var. ramiflora Moldenke (1953: 178 – 179). Gmelina sessilis forma ramiflora (Moldenke) Moldenke (1984h: 178 – 179), synon. nov. Type: [Indonesia, Papua] New Guinea, West Irian, Hollandia, Bernhard Bivak, 26 July 1938, Meijer Drees 371 (holotype BO (n.v.); isotype NY, scan seen).

Trees 9 – 30 m high, DBH 15 – 100 cm. Bark rough, fissured and peeling off in irregular scales, white to grey or light–dark brown; slash orange-brown to creamy yellow, turning red-brown on exposure. Wood soft to hard, creamy white, with pink tinge. Twigs hairy, spines absent. Leaves elliptic to orbicular, margins entire, often shallowly lobed, 10 – 41  ×  6 – 20 cm, apex rounded to acute, coriaceous, base cordate to cuneate,; upper surface glabrous, with some hairs on veins, light to dark green, sometimes shiny; lower surface covered with rusty brown hairs, often with glands clustered around the base, peltate scales absent, (pale) green; veins 6 – 11 pairs, sometimes 3- veined from base, lower surface with strong ladder-like tertiary venation. Petiole 3 – 7 cm long, half-terete, channelled, hairy. Inflorescence terminal, 8 – 30 cm, velutinous, ochreous-golden; bracteoles lanceolate, 4 – 10  ×  1 – 2.5 (− 5) mm wide, apex acute to acuminate, caducous. Calyx 5-lobed, sometimes appearing fewer, 3 – 5  ×  3 – 5 mm, velutinous, olive green, outer surface with discoid glands; lobes triangular, 0 – 1 mm long, apex acute to rounded; fruiting 4 – 15 mm diam., erect to patent. Corolla 5-lobed, outer surface hairy, inside glabrous, sparsely glandular, white to mauve, sometimes fragrant; anterior lip 3-lobed; mid-lobe spathulate to oblong, 3 – 14  ×  3.2 – 8 mm, apex rounded, blue to dark purple with yellow blotch at base; side lobes oblong 7.5 – 8  ×  4 – 5 mm, apex rounded; posterior lip, 2-lobed, 5 – 7  ×  4 – 5 mm, apex rounded; tube 8 – 10 mm long. Stamens inserted at apex of tube, white; anterior pair 10 – 12 mm long, anthers c. 1.5 mm long; posterior pair, 7 – 9 mm long, anthers 1 – 1.2 mm long. Ovary c. 1 mm diam., glabrous; style 11 – 14 mm long, white, stigma c. 1 mm long. Fruit (dried) 10 – 18  ×  10 – 16 mm, globose, later cylindrical, apex truncate when mature, glabrous, blue or purple to red.

distribution. Indonesia: Moluccas: Ceram, Halmahera, Ternate and Morotai; Papua: Papua New Guinea and Solomon Islands. Map 1.

selected specimens examined. indonesia. Moluccas: n.d., anon. (holotype BR); Amboyna [ = Indonesia, Ambon], Wallich 1819 (isotype K-W); Amboina, July – Nov. 1913, Robinson 296 (BM, L); Banda-gruppe, Ay Insel, June 1821, Reinwardti 1362 (syntypes L); Insel Neira, May 1859, de Vriese s.n. (syntype L); Ile de Banda, 1829, Reinwardt 5158 (L, P); Tanimber-eilanden, Loeroemboen, 20 April 1938, Neth. Ind. For. Service bb. 24431 (L) West Ceram, Kairatu, Gemba, 6 June 1959, Kuswata & Soepadmo 103 (K); Halmahera, North Gaila, 21 Sept. 1921, Beguin 1746 (K, L); Ternate, Amo, 14 April 1950, Bish 4 (K); New Guinea: Japen Island, Soemberbaba, 13 July 1961, Koster BW 11162 (L); [Netherlands New Guinea, Division Hollandia], Sabronsamon, 13 Aug. 1957, Kalkman BW 6209 (K). PAPUA NEW GUINEA. Manus Island: Mt Dremsel, 21 June 1971, Stone & Streimann 53636 (L); West New Britain Province: Mt Bango, 19 May 1968, Henty & Lelean 29453 (K). SOLOMON ISLANDS. North West Santa Ysabel: Binusa, 25 Jan. 1966, Beer’s collectors BSIP 6716 (K); San Cristobal, Wairaha R., 11 May 1964, Whitmore BSIP 4255 (K).

habitat. Growing in primary and secondary, well drained, (rain) forests; alt. 0 – 800 m, sometimes in clayey soil or sandy loam, sometimes over limestone; sometimes associated with Araucaria hunsteinii K. Schum.

conservation status. Least Concern.

phenology. Flowering Jan. – Oct.; fruiting March – Dec.

vernacular names. Indonesia, Moluccas: Ambon: Caju titti; Halmahera: Foehoe or Gal (North Galia Lanaguge). Papau: Nassa-onggoe or Nakaoengga (Kamtoek Language); Anjoes (Warnapi and Manokwari); Ainoes (Biak language); Ankieève or Angki-e-oe (Manikiong Language); Bas (Mooi Language); Kaina (Karoon Language); Sarot (Amberbaken Language); Tanee (Kho Language); Merkemmek (Arfak Sidai language); Ainoe (Ambai language). Papua New Guinea: Morobe Distr.: Aron gazo (Garaina Language); Central Distr.: Waria or Gow (Brown R.); Gulf Division: Omormi (Kikori Language); Jacquinot Bay: Ossogee (Warea language). Solomon Islands: Arokoko (Kwara’ae language).

uses. The wood is used in making small boats and house panelling as it is reported to be soft, light and easy to work, and also durable in seawater, but not in fresh water. The species is reported to be cultivated in the Moluccas for this particular use (Rumphius 1750: 38). The pulverised bark is use to clean wounds and ulcers (Heyne 1950: 1321).

notes. Type materials of Gmelina sessilis var. ramiflora Moldenke, G. moluccana f. glandulosa (Hallier f.) Moldenke, G. macrophylla Wall. ex Schauer and Volkameria obovata Roxb. fall within the total morphological variation of Gmelina moluccana and are therefore place into its synonomy.

19. Gmelina neocaledonicaS. Moore in Rendle et al. (1921: 375); Moldenke (1984h: 155 – 156); Mabberley in Mabberley & de Kok (2004: 30 – 33). Type: New Caledonia, Comboui, 26 Oct. 1914, Compton 2158 (holotype BM (n.v.); isotype NSW (n.v)). For a description and more information see Mabberley in Mabberley & de Kok (2004).

specimens examined. new caledonia. Upper Tontouta valley, 20 Jan. 1955, McKee 3484 (L).

conservation status. Least Concern.

20. Gmelina palawensis
  • 1a. Corolla outside white, leaves elliptic to slightly ovate, blade often unequal...............................................................

  • ...............................................................Gmelina palawensis subsp. palawensis

  • 1b. Corolla outside (pale) pink, leaves elliptic to slightly obovate, blade equal...............................................................

  • ....................................................................Gmelina palawensis subsp. celebica

20.1 Gmelina palawensis subsp. palawensis

Gmelina palawensis H. J. Lam (1919: 224 – 225); Moldenke (1984h: 157 – 159). Type: Palau Islands, Babelthaop, Ngarsul, 21 Feb. 1914, Ledermann 14331 (lectotype B 10 0379005 scan seen), selected here.

Trees 10 – 12 m high. Twigs glabrous, spines absent. Leaves elliptic to slightly ovate, 9 – 15  ×  5 – 9 cm, apex rounded to acute, base cuneate, often unequal, margins entire; upper surface glabrous; lower surface glabrous, with discoid glands clustered around the base, peltate scales absent; veins 4 – 7 pairs, sometimes 3-veined from base. Petiole 4 – 7 cm long, glabrous, half-terete, channelled. Inflorescence terminal, rarely axillary, narrow, up to 15 cm long, glabrous; bracteoles lanceolate, 5 – 7.5  ×  0.8 – 2 mm, apex acute, sessile, caducous. Calyx 5-lobed, sometimes appearing fewer, glabrous, outer surface sometimes with discoid glands; flowering calyx 4 – 5  ×  3 – 4 mm; lobes 0 – 0.1 mm long, apex acute to rounded; fruiting calyx 7 – 9 mm diam., erect to patent. Corolla 5-lobed, white, outer surface hairy; anterior lip 3-lobed; mid-lobe spathulate to oblong, 4 – 5  ×  2.5 – 4 mm, apex rounded, dark or bright blue-purple with yellow spot at base; side lobes oblong, 3 – 7  ×  2 – 5 mm, apex rounded; posterior lip 2-lobed, oblong, 2 – 5  ×  2 – 3 mm, apex rounded; tube 8 – 10 mm long. Stamens inserted at apex of tube; anterior pair 7 – 10 mm long, anthers c. 2 mm long; posterior pair 6 – 8 mm long, anthers c. 2 mm long. Ovary glabrous; style c. 15 mm long, stigma c. 1.5 mm long. Fruit 18 – 23  ×  10 – 13 mm, clavoid, apex rounded, glabrous.

distribution. Endemic to the Palau Islands; introduced to the Caroline Islands. Map 6.
Map 6

Distribution of Gmelina palawensis subsp. palawensis (◊); Gmelina palawensis subsp. celebica (▲); Gmelina racemosa (■); Gmelina smithii (◘); Gmelina uniflora (●); Gmelina salomonensis (Open image in new window).

specimens examined. caroline islands. Pelew Island: July 1933, Kanehira 2280 (K); Truk-Moen Island: 20 Aug. 1980, Fosberg 60359 (K). PALAU ISLANDS. Arumogui- hukin-sinrin, 17 Sept. 1933, Hosakawa 6969 (L); Kaiguru, 15 April 1936; Babelthuap [Babeldaob] Island: Gaspan, 1 Jan. 1963, Stone 4618 (K, L); Ngarsul, 21 Feb. 1914, Ledermann 14331 (lectotype B); 5 – 7 April 1950, Fosberg 32528 (K); Takamatsu 1594 (K); Malakal Island: 25 Aug. 1965, Fosberg 47522 (K).

vernacular name.Blaheos (Ngarsul Island); Blachaiósch (Korror Island); La eos or La eus (Malakal Island).

habitat. Growing in secondary vegetation or primary forest; alt. 25 – 300 m. Soil weathered volcanic material.

conservation status. Least Concern.

phenology. Flowering Feb. – Aug.; fruiting Jan. – April.

uses. Wood used in house construction.

notes. Of the various collections cited in the original description (Palau Islands, Babelthaop, Ngarsul, 24 Feb. 1914, Ledermann 14420; Palau Islands, Babelthaop, Ngarsul, 21 Feb. 1914, Ledermann 14331; Palau Islands, Korror, Raymundus 114; Palau Islands, Korror, Raymundus 310) only one (Ledermann 14331) could be found, and is selected here as the lectotype. Given the morphological and geographical differences between the two parts of this species, it would be more consistent if this variation was recognised at subspecific level (see de Kok 2007).

20.2 Gmelina palawensis subsp. celebica (Moldenke) de Kokcomb. et stat. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77120323-1

Gmelina palawensis var. celebica, Phytologia 3: 417 (1951); 1984h: 159 – 160). Type: [Indonesia, Sulawesi] Celebes, Malili, 2 Aug. 1929, Kjellberg 2001 (holotype BO 21308!; isotype BO 21309!).

Trees 5 – 15 m high, DBH c. 10 cm; bark smooth, grey to brown. Twigs glabrous, spines absent. Leaves elliptic to slightly obovate, 9 – 19  ×  5 – 7.5 cm, apex rounded to acute, base rounded to cuneate, equal, margins entire; upper surface glabrous, shiny; lower surface glabrous, with discoid glands clustered around the base, peltate scales absent; veins 7 – 9 pairs, sometimes 3-veined from base. Petiole 2.5 – 4 cm long, glabrous, half-terete. Inflorescence terminal, narrow, up to 20 cm long, glabrous to sparsely hairy at apex; bracteoles lanceolate to spathulate, 6 – 15  ×  1 – 7 mm, apex rounded to acute, sessile, caducous. Calyx 5-lobed, sometimes appearing fewer, glabrous, outer surface without discoid glands, green, sometimes dark red brown at apex; flowering calyx 3.5 – 5  ×  4 – 4.5 mm; lobes 0 – 0.1  ×  0 – 0.1 mm long, triangular, apex acute; fruiting calyx 6 – 10 mm diam., patent. Corolla 5-lobed, outer surface hairy, (pale) pinkish to lilac or purple outside, white inside, fragrant; anterior lip 3-lobed; mid-lobe oblong, 5 – 7  ×  4 – 5 mm, apex rounded, cream pink with two longitudinal yellow honey marks; side lobes oblong, 4 – 6  ×  3 – 4 mm, apex rounded, oblique; posterior lip 2-lobed, oblong, 5 – 6  ×  3 – 4 mm, apex rounded; tube 10 – 13 mm long. Stamens inserted at apex of tube; anterior pair 8 – 12 mm long, anthers c. 2 mm long, brown; posterior pair, 7 – 10 mm long, anthers c. 2 mm long, brown. Ovary c. 2 mm diam., glabrous; style 13 – 20 mm long, stigma c. 1.5 mm long. Fruit 17 – 22  ×  12 – 15 mm, clavoid, apex rounded, surface with bumps, glabrous, red or purple when mature.

distribution. Endemic to Sulawesi, including the Sangihe islands. Map 6.

specimens examined. indonesia. [Sulawesi] Celebes: Malili, 2 Aug. 1929, Kjellberg 2001 (holotype BO; isotype BO); Malili, 3 Feb. 1934, Neth. Ind. For. Service Cel./IV-131 (K); Malili, 3 Feb. 1934, Neth. Ind. For. Service Cel./IV-129 (SING); Malili, 10 Dec. 1994, Sidiyasa 1326 (K, L); Malili, 18 Oct. 1934, Neth. Ind. For. Service Cel./III-168 (L); Malili, 19 Sept. 1930, Boschproefst 441 (K, L); Malili, 27 Jan. 1925, Boschproefst 8560 (L); surroundings of Matano Lake, near Soroako, NE of Malili, 22 July 1976, Meijer 11514 (L); Selatan, Soroako, S Shore of Lake Matano, 10 June 1979, de Vogel 5707 (K, L); Selatan, Luha, Island in Lake Towuti, 18 July 1979, de Vogel 6367 (K, L); South Sulawesi, Lake Matano, SE of Soroako, 28 June 1979, van Balgooy 3797 (K, L); South Sulawesi, Soroako, 3 July 1979, van Balgooy 3858 (K, L); Sangihe Island, Kalumelahana, 12 Sept. 1998, Hicks 58 (K).

habitat. Growing in secondary vegetation and rainforest or landside of mangrove; alt. 20 – 730 m. Soil red clayey or podsolic, often over ultra-basic, limestone or schists.

conservation status. Least Concern.

phenology. Flowering June – Jan.; fruiting July – Nov.

vernacular name.Mahangparata (Sangihe language); Woeroko or Kajoe poetih (Malili).

21. Gmelina papuanaBakh. (Bakhuizen 1929: 71 – 72 & Pl. 16 & 17). Gmelina sessilis var. papuana (Bakh.) Moldenke (1953: 178; 1984h: 176 – 178). Type: [Papua New Guinea] Papua, Iawarere, 25 Nov. 1925, Brass 695 (holotype BO 1336416!; isotype BO 1336415!).

Gmelina brassii Moldenke (1958: 324 – 325; 1984f: 32 – 33). Type: Papua New Guinea, Milne Bay Distr. [ = Province], Cape Vogel Peninsula, Dabora, 10 April 1953, Brass 21915 (holotype LAE!; isotypes K! L! US scan seen).

Tree 7 – 30 m high, DBH 10 – 40 cm. Bark grey to pale brown, fissured to scaly. Wood pale brown to whitish yellow or white. Twigs hairy, with interpetiolar ridge present, spines absent. Leaves elliptic or lanceolate to ovate, 9 – 29 × 4.5 – 14 cm, slightly oblique, apex rounded to acuminate, base rounded to cordate, margins entire; upper surface glabrous with few simple hairs on veins, dark green, matte to glossy; lower surface glabrous with few simple hairs on veins, peltate scales absent, (pale) green; two cup-like glands clustered at the base; veins 3 – 9 pairs, sometimes 3-veined from base. Petiole 2 – 5 cm long, terete, channelled, hairy. Inflorescence terminal, dense, up to 20 cm long, without long side branches, velutinous, brownish; bracteoles lanceolate, 7 – 15  ×  2 – 8 mm, those on the main axis significantly bigger than those on the side axes; apex long acuminate, with several disc-like glands present. Calyx 5-lobed, 3.5 – 5  ×  2.5 – 4 mm, velutinous, greenish brown, outer surface with discoid glands; lobes up to 0.5 mm long, apex long acuminate; fruiting calyx 5 – 6 mm diam., erect. Corolla 5-lobed, outer surface hairy, sparsely glandular, white or yellowish to pinkish/lilac; anterior lip 3-lobed; mid-lobe oblong, 6 – 10  ×  5 – 6 mm, apex acute, mauve or purple, with two yellow marks; side lobes 3.5 – 5  ×  3 – 5 mm long, apex acute to rounded; posterior lip 2-lobed, 4 – 6  ×  3 – 4 mm, apex rounded; tube 7 – 13 mm long. Stamens inserted at apex of tube, glabrous, glands present; anterior pair 9 – 12 mm long, anthers c. 1.5 mm long; posterior pair, fertile, 7 – 9 mm long, anthers 1.3 – 1.5 mm long. Ovary glabrous with a few hairs at apex. Style 13 – 15 mm long, stigma c. 2 mm long. Fruit (dried) 7 – 10  ×  4 – 6.5 mm, clavate, apex rounded, glabrous, apart from some hairs at apex, black or purple when mature.

distribution. Southern part of Papua New Guinea. Map 3.

specimens examined. papua new guinea. Milne Bay Province: Alotau, 28 March 1970, Henty & Katik 42917 (L); near Mapo, March 1945, Smith 1345 (L); Cape Vogel Peninsula, Dabora, 10 April 1953, Brass 21915 (holotype LAE; isotypes K, L, US); Cape Vogel Peninsula, Menapi, 28 March 1953, Brass 21719 (L); about 0.5 mile N of Waigani Plantation, March 1945, Smith 1298 (K, L); near Kaporika Village, 5 June 1964, Henty 16985 (K, SING); Alotau subprov., 1 mile of Ulabo camp, 6 March 1984, Gideon 77041 (K, L); Normanby Island, Sewa Bay, 21 Oct. 1971, Lelean & Streimann 52522 (K, L); Normanby Island, near Miadeba airstrip, 23 Nov. 1976, Croft et al. 68878 (K, L); Normanby Island, Waikaiuna, 12 April 1956, Brass 25383 (K, L); Normanby Island, Sewa Bay, 20 April 1956, Womersley 8680 (K, L SING); Fergusson Island, E of lake Lavu, 15 Nov. 1976, Croft et al. 68823 (K, L); Fergusson Island, Agamoia, 22 June 1956, Brass 27281 (K, L); Modewa Bay, Modewa, 17 Dec. 1956, Brass 28910 (K, L); Western Highlands Province: near Dagarunga Ridge, 1 Sept. 1968, Henty & Streimann 38948 (K); Central Province: Brown R. Bridge, 10 Nov. 1953, Jackson & McDonald 4580 (K); Brown R. Forest Reserve, Jan. 1960, McDonald 8213 (K); Iawarere, 25 Nov. 1925, Brass 695 (holotype BO, isotype BO), Idenburg R., Bernhard Camp, Feb. 1939, Brass 12751 (K). Morobe Province: Wampit R., Bupu Village, 3 March 1964, Millar 23247 (K).

habitat. A (sub) canopy tree in primary or secondary forest or in transition zone between rainforest and savanna, sometimes over limestone or on beaches; alt. 0 – 1400 m.

conservation status. Least Concern.

phenology. Flowering (Nov.) March – June; fruiting Nov. – June. The fruits are reported to be eaten by ‘Canowangs’ (Bakhuizen van den Brink 1929: 71 – 72), which is possibly the bird genus Strepera, known in Australia as Currawongs.

vernacular names. Papua New Guinea, Milne Bay: Alongaya (Upper Waria Language); Petapeta (Kaporika village); Gow (Brown R. area).

notes. The wood is used for carving drums.

22. Gmelina peltatade Koksp. nov. a G. lepidota venulis lateralibus magis numerosis plus quam 9 (nec minus quam 7) numero utroque latere, bracteolis longioribus plus quam 2 mm (nec minus quam 2 mm) longis atque non persistentibus et fructu maturo griseo-albido (nec rubro nec pallide purpureo) differt. Typus: Solomon Islands, Small Malaita, Pululaha Area, 13 Aug. 1969, Gafui & Collectors 16942 (holotypus K!).

http://www.ipni.org/urn:lsid:ipni.org:names:77120324-1

Trees 7 – 25 m high, DBH 10 – 60 cm. Bark smooth to deeply fissured or scaly, grey to (light to dark) brown. Wood soft to hard, white or whitish brown. Twigs hairy, when young, spines absent. Leaves elliptic to lanceolate, 5 – 15.5  ×  2 – 6 cm, apex acute to acuminate, base rounded to cuneate, margins entire; upper surface glabrous with simple hairs on veins at base, dark green; lower surface glabrous with simple hairs on veins, sometimes with discoid glands clustered around the base, covered with peltate scales, grey; veins 9 – 16 pairs. Petiole half-terete, channelled, 1.2 – 3 cm, hairy. Inflorescence terminal and axillary, 10 – 20 cm, sparsely hairy, peltate scales present; bracteoles lanceolate, 2 – 5  ×  0.8 – 3 mm long, apex acute, caducous. Calyx 5 lobed, 2.5 – 3  ×  2 – 3 mm, glabrous, surface covered with peltate scales, outer surface with discoid glands; lobes 0.5 – 0.8  ×  0.8 – 1 mm; apex acute to rounded; fruiting calyx 9 – 12 mm diam., patent. Corolla 5-lobed, pale pink to purplish brown, outer surface hairy, sparsely glandular, scented; anterior lip 3-lobed; mid-lobe 6 – 25  ×  3.5 – 5 mm, oblong, apex rounded; side lobes 4 – 5  ×  5 – 5.5 mm, apex rounded to acute, sometimes oblique; posterior lip, 2-lobed, 4 – 5  ×  4 – 4.5 mm, apex rounded; tube 8 – 13 mm long. Stamens inserted at apex of tube, glabrous; anterior pair 10 – 12 mm long, anthers c. 1.6 mm long; posterior pair, fertile, 6 – 9 mm long, anthers c. 1.5 mm long; ovary glabrous with a few hairs at apex. Style 10 – 12 mm long, stigma 1.5 – 2 mm long. Fruit (dried) 13 – 20  ×  11 – 16 mm, globose, apex rounded, glabrous, green to greyish white when mature. Fig. 4.

distribution. Solomon Islands. Map 5.

specimens examined. solomon islands. San Cristobal: Oneibia Area, 19 Nov. 1968, Runikera & Collectors 12660 (K); Oneibia Area, 26 Jan. 1969, Mauriasi 14552 (K); Tetere Makira Bay area, 23 Oct. 1968, Runikera & Collectors 12504 (K); Maeaba Ridge – Maru Bay, 3 Dec. 1968, Gafui & Collectors 12904 (K); Small ( = South) Malaita: Pululaha Area, 13 Aug. 1969, Gafui & Collectors 16942 (holotype K); Rokera, 25 July 1968, Runikera & Collectors 10406 (K); N of Palasu’u School, 29 Feb. 1969, Gafui & Collectors 17309 (K); South West Malaita: Are are distr., Kiu, 9 Dec. 1963, Lipaqeto 3511 (K); Fauro Island: NW Fauro, Barahono R. area, 14 April 1969, Mauriasi & Collectors 13921 (K); E Fauro, Haliuna R. area, 18 April 1969, Mauriasi & Collectors 13993 (K); Guadalcanal: Beaufort, Kombau, 21 Feb. 1946, Walker 236 (K).

habitat. Growing in secondary or primary (beach) forests, often on well drained soils; alt. 0 – 210 m.

conservation status. Least Concern.

phenology. Flowering July – Nov.; fruiting Dec. – Oct.

vernacular name. Solomon Islands: Maladala (Kwara’ae Language).

uses. The wood is used in making outriggers for boats.

NOTES. The specific epithet refers to the peltate scales, in particular those on the leaves and calyx.

23. Gmelina philippinensisCham. (Chamisso 1832: 109); Merrill (1912: 405 – 406; 1918: 333 – 334; 1935: 335); Moldenke (1984h: 161 – 172); Vũ (2007: 135 – 137); Gmelina asiatica var. philippinensis Bakh. in Lam & Bakhuizen (1921: 70 – 71). Type: [Philippines] Luçonia, Manilla, Roman-Zoffiana s.n. (holotype LE, picture at K seen). [Gmelina finlaysoniana Wall. (Wallich 1828: 6317), nom. nud. Based on: [Vietnam, Da Nang] Turon Bay, Wallich 6317 (K-W!).]

Gmelina asiatica Blanco (1837: 492), nom. illegit., non Gmelina asiatica L; Merrill (1918: 333 – 334). Type: unknown.

Gmelina inermis Blanco (1837: 493); Merrill (1918: 333 – 334). Type: unknown.

Gmelina hystrix Schult. ex Kurz (1870: 81); Kochummen (1978: 306). Type: [Thailand] Siam, Bangkok, in hortis, Teysmann 5946 (holotype L!).

Gmelina bracteata Burck (1891: 98 – 99). Type: Burck (1891: t. VII, Fig. 5 – 6).

Gmelina finlaysoniana var. silvestris forma colorata Kuntze (1891: 507); Gmelina philippinensis forma colorata (Kuntze) Moldenke (1984b: 234; 1984h: 172). Type: [Vietnam, Vung Tàu, Ba-Ria Vung Tàu Prov.] Cape St. James, n.d., anon. (holotype NY, scan seen).

Gmelina finslaysoniana var. silvestris forma viridibracteata Kuntze (1891: 507); Gmelina philippinensis forma viridibracteata (Kuntze) Moldenke (1984b: 234; 1984h: 173). Type: [Vietnam, Vung Tàu, Ba-Ria Vung Tàu Prov.] Cape St. James, n.d., anon. (holotype NY, scan seen).

Gmelina philippinensis forma transitoria Moldenke (1969: 210; 1984h: 172 – 173). Type: Philippine islands, Guimaras Island, Feb. or March 1950, Sulit 11741 (holotype US scan seen; isotypes L! × 2 sheets).

Gmelina szechwanensis K. Yao (1982: 211 – 212 & 125 as ' szechuanensis' on p. 124, and ‘szechuenensis’ on p. 125), synon. nov. Type: [China] Sichuan, Puge, 8 Aug. 1960, unknown 614137 (holotype NAS (n.v.)).

Gmelina thothathriana A. Rajendran & P. Daniel (2002: 169 – 171), synon. nov. Type: [India] Andaman Islands, Dithaman, 28 Feb. 1959, Thothathri 9243 (holotype CAL (n.v.); isotype MH (n.v.)).

Climbing shrubs or small tree, 2 – 8 m high, DBH 3 – 11 cm. Bark light brown or grey to green, rough. Twigs sparsely hairy; spines usually present, up to 10 mm long. Leaves elliptic to obovate, 2.5 – 11  ×  1.4 – 5 cm, margins entire, sometimes shallowly 3-lobed, apex rounded to acute, base cuneate to decurrent; upper surface glabrous, with sometimes a few hairs on main vein, green; lower surface glabrous with some hairs on the veins, white stellate hairs present, green; sometimes with discoid glands at base; veins 3 – 4 pairs, sometimes 3-veined from base. Petiole 9 – 30 mm long, channelled, sparsely hairy, pink. Inflorescence terminal, 3 – 13 cm long, erect to patent, sparsely hairy; bracteoles ovate-lanceolate, 9 – 30  ×  10 – 50 mm, apex rounded to acute, cuspidate, usually persistent, light brown, pale green or purple or maroon outside, pale green with pink inside, sometimes with discoid glands. Calyx 5-lobed, sometimes appearing fewer, densely to sparsely pubescent, becoming more glabrous over time, pale yellow green to green, discoid glands present, small white glands present; flowering calyx 3 – 4  ×  2.5 – 3 mm, lobes c. 0.2 mm long, apex acute; fruiting calyx 4 – 10 cm diam., margin straight to undulate, patent. Corolla 4-lobed, yellow, velutinous, slightly fragrant; anterior lip 3-lobed, mid-lobe 3 – 9  ×  6.5 – 12 mm, apex acute to rounded, patent; side lobes 3.5 – 7  ×  6 – 8 mm, apex acute to rounded, slightly oblique; posterior lip 1-lobed, 4.5 – 15  ×  6 – 14 mm, apex acute to rounded; tube 10 – 16 mm long. Stamens inserted at apex of tube, yellow; anterior pair 18 – 22 mm long, anthers 1.5 – 3 mm long, yellow; posterior pair, 11 – 13 mm long, anthers c. 2 mm long. Ovary glabrous, sometimes with a few discoid glands; Style 28 – 40 mm long. Fruit (dried) 15 – 20  ×  9 – 12 mm, obovoid, apex rounded, glabrous, yellow when mature.

distribution. This species has an interesting and complex distribution. It is considered to be native in the Philippines, East Cambodia and Central and South Vietnam and possibly east Thailand (Map 5). As it is now also cultivated as a garden plant throughout the tropics and has become naturalised in many areas the exact limits of its natural distribution may be impossible to determine.

selected specimens examined. cambodia. Mondulkiri Province: Pech Chenda distr., O Lorh stream, 30 Jan. 2001, Meng Monyrak et al. 452 (K); Ratanakiri Province: Lum Phat distr., Keng Sann village, 25 March 2001, Meng Monyrak et al. 742 (K). MALAYSIA. Sarawak: Kapit, Upper Rejang R., 1929, Clemens 21089 (K). PHILIPPINES. Mindanao: Butuan subprovince, March – July 1911, Weber 1028 (K); Laguna Province:, Mt Maquiling, June – July 1917, Elmer 18278 (K) Guimaras Island: Feb. or March 1950, Sulit 11741 (holotype US; isotypes L). THAILAND. Siam: Bangkok, in hortis, Teysmann 5946 (holotype L). VIETNAM. Annam: Dalat, March – April 1932, Squires 920 (K); Nha-trang: 11 – 26 March 1911, Robinson 1344 (K); Tourane: 6 Sept. 1920, Poilane 1455 (K); 100 km S of Hue, May – July 1927, Clemens 3152 (K); [Da Nang] Turon Bay: Wallich 6317 (K-W); Vung Tàu, Ba-Ria Vung Tàu Province: Cape St. James, n.d., anon. (holotype NY); Cape St. James, n.d., anon. (holotype NY).

habitat. Growing in secondary vegetation or (mixed) deciduous forests; alt. 60 – 490 m. Soil clay.

conservation status. Least Concern.

phenology. Flowering Jan. – Oct.; fruiting March – Nov.

vernacular names. Thailand: Sam ma nga or Cling Chai. Vietnam: Cây ngăy nui.

uses. The species is used extensively throughout the tropics as a garden plant. In Peninsular Malaysia the fruit is mixed with lime and applied externally to the throat as a remedy for coughs.

notes. In 1912 Merrill stated that the Philippines specimens of Gmelina philippinensis he had seen always have pale green bracteoles, while the specimens he had seen from Indo-China have purple bracteoles. Kuntze (1891: 507) founded his garden-based forms of Gmelina finslaysoniana var. silvestris on this supposed difference in bracteole colour, with forma viridibracteata (green bracteoles) and forma colorata (brown bracteoles). There is very little evidence from herbarium specimens to suggest a clear geographical distribution pattern in bracteole colour. This is mainly due to the fact that very few collectors record the bracteole colour. Specimens from Cambodia are reported to have pale green bracteoles with pink inside (Meng Monyrak 452, 742); I have seen one specimen from Vietnam with maroon bracteoles (Squires 920), while a specimen from Sarawak is reported to have bracteoles with purple abaxial surfaces (Clemens 21089). In his studies of extrafloral nectaries, Burck (1891) notes that the persistent bracteoles in this species reduce the percentage of flowers being robbed of their nectar through perforations at the base; 20 – 40% of total flowers observed in G. asiatica (which has small bracteoles) being robbed, vs 3 % in G. philippinensis (as G. bracteata Burck.). Futhermore, he claims that the space within the inflorescences created by the persistent bracteoles are used by ants for their nests. I have not seen the type materials of G. thothathriana A. Rajendran & P. Daniel and G. szechwanensis K. Yao, but from the descriptions and plates it is clear that they fall within the total morphological variation of G. philippinensis Cham. and are therefore placed into its synonymy.

24. Gmelina racemosa(Lour.) Merr. (Merrill 1935: 336); Moldenke (1984h: 173 – 175); Vũ (2007: 140 – 141). Lantana racemosa Lour. (Loureiro 1790: 376 – 377). Type: [China] Hainan, Fan Ya, 25 April 1922, McClure 9281 (neotype K!), selected here.

Gmelina hainanensis Oliv. (Oliver 1889: 1874), nom. invalid.; Moldenke (1984g: 107 – 109); Shou-liang & Gilbert (1994: 33). Type: [China] Hainan, Henry s.n. (holotype K!; isotype P!).

Gmelina annamensis Dop (1933: 894); Moldenke (1984c: 336); Vũ (2007: 137 –138). Type: Vietnam, Annam, Quantri, entre Laùg-Lut-Ha, and Lang-Pa-Ka, 10 May 1927, Poilane 13301 (holotype P, scan seen; isotype HM (n.v.)).

Tree or shrub 2 – 22 m high, DBH 10 – 18 cm. Bark grey. Twigs hairy when young, spines absent. Leaves ovate to elliptic, 5 – 16  ×  4 – 10 cm, apex acute to acuminate, base cuneate to almost cordate, margins entire, papery; upper surface glabrous with few simple hairs on veins at base, green, shiny; lower surface glabrous to sparsely hairy, green; few discoid glands clustered around the base and veins, covered with peltate scales; veins 3 – 4 pairs, the first pair often starting from the very base. Petiole terete, slightly channelled, 2 – 6.5 cm, glabrous to sparsely hairy. Inflorescence terminal, 5 – 8 cm, velutinous; bracteoles lanceolate, 7.5 – 14 (− 25)  ×  2.5 – 6 (− 10) mm, grey, with few disc-like glands present. Calyx 5-lobed, 5 – 13  ×   7 – 15 mm, sparsely hairy, covered with peltate scales, outer surface with few discoid glands, not accrescent; lobes triangular, 4 – 10  ×  6 – 8 mm, anterior lobes biggest, apex acute to rounded. Corolla white to yellowish or pinkish outside, red to orange or purplish inside; outer surface sparsely hairy, covered with glands; sparsely glandular, ill-smelling or fragrant; anterior lip 3-lobed; mid-lobe spathulate, 5 – 10  ×  10 – 11 mm, apex rounded; side lobes rounded, 8 – 9  ×  8 – 9 mm, apex rounded; posterior lip, 2-lobed, oblong, 10 – 11  ×  7 – 7.5 mm, apex rounded; tube 2.5 – 5 mm long. Stamens inserted at apex of tube; anterior pair 22 – 23 mm long, anthers 2.3 – 3 mm long; posterior pair, fertile, 20 – 22 mm long, anthers 2 – 2.5 mm long. Ovary 2.5 – 3  ×  2.5 – 3 mm, velutinous in upper half. Style 23 – 30 mm long, stigma 1.5 – 2.5 mm long. Fruit (dried) 10 – 12  ×  4 – 7 mm, clavate with a long stalk, apex truncated, glabrous with a few hairs at apex, covered with peltate scales, few discoid glands present, grey.

distribution. China: Hainan and Kwangtung; North Vietnam. Map 6.

specimens examined. china. Hainan: Fan Ya, 25 April 1922, McClure 9281 (neotype K, isotype P); Henry s.n. (holotype K); Non Pong Shan, Taam-chau Distr., 17 Sept. 1927, Tsang 904 (16403) (K); Ching Mai Distr., Ku Tung Village, Lei 450 (K); Seven Finger Mts, Lingmon, 7 May 1932, Liang 61767 (K); Tung Koo Sha, Near Shan Ho Village, Aug. 1932, Fung 20570 (K); Hung Mo Shan, Lai [Loi] area, 6 May 1929, Tsang et al. 97 (K); Nar Tai Lee, May 1893, Chinese collector 454 (K); Ngai Distr., Between T'ang K'iu (Din-Kio) and Po T'eng, April – May 1932, McClure 20049 (K); Shui Mei Shan (Taam Chau Distr.), 25 May 1928, Tsang 500 (LU 17249) (K); Yaichow, March – June 1933, How 70801 (L); Dung Ka to Mo San, 1932 – 1933, Chun & Tso 43542 (L). VIETNAM. Tonkin: Kau Nga Shan, Tien-yen, 23 Sept. – 7 Oct. 1940, Tsang 30588 (L, SING); Taai Wong Mo Shan, Tong Fa Market, Ha-coi, 11 – 23 Sept. 1939, Tsang 29546 (L); Sai Wong Mo Shan (Sai Vong Mo Leng), Lung Wan Village, 18 May – 5 July 1940, Tsang 29869 (L); Annam: Massif de Dong Cho, Quang Tri, 2 June 1924, Poilane 10723 (K, P); Quantri, entre Laùg-Lut-Ha, and Lang-Pa-Ka, 10 May 1927, Poilane 13301 (holotype P).

habitat. Growing in open grassy areas to dense (mixed) monsoon forest; alt. 300 – 500 m. In primary and secondary forest. Soil sandy or clay or loamy. Reported to be a drought tolerant tree and light-demanding with natural regeneration occurring in open places (Chinh et al.1996: 741).

phenology. Flowering April – July; fruiting May – Sept.

conservation status. Least Concern.

vernacular names. China: Ku Zi. Vietnam: Cây tlai or Lõi thọ hải nam.

uses. Reported to be a fine timber, with wood being resistant to rot. Used in ship, house, bridge building and tool making (Chinh et al.1996: 741; Shou-liang & Gilbert (1994: 33).

notes. The name Gmelina hainanensis was proposed at the end of the description of G. chinensis. Oliver (1889) gives a short analysis of how this new species differs from G. chinensis and then states ‘it may be called Gmelina hainanensis’. According to article 34.1 (McNeill et al.2006) in the botanical code it is not allowed to reserve names for taxa you do not yourself recognise at the time, therefore the name G. hainanensis is invalid. No type material of Lantana racemosa Lour. could be found at BM, K or P. I have thererefore selected a neotype from China, with flowers and mature calyxes for this name.

25. Gmelina salomonensisBakh. (Bakhuizen 1935: 72 – 73). Type: Solomon Islands, Ysabel [Isabel] Island, Tiratoña, 8 Dec. 1932, Brass 3309 (holotype BO 1337463!; isotypes A (n.v.), BO 1337464!, L!).

Gmelina salomonensis forma glabrescens Moldenke (1953: 178); Gmelina moluccana var. glabrescens (Moldenke) Moldenke (1970: 435; 1984h: 155). Type: Solomon Islands, Bougainville Island, Karngu, Buin, 10 Oct. 1930, Kajewski 2228 (holotype BO (n.v.); isotype L! NY, scan seen).

Gmelina salomonensis var. elliptica Moldenke (1969: 71); Gmelina moluccana var. elliptica (Moldenke) Moldenke (1970: 435). Type: Solomon Islands, Santa Ysabel, Kolokofa R., 6 April 1966, Teona BSIP 6371 (holotype US, scan seen; isotype L!)

Tree, 6 – 40 m high, DBH 12 – 400 cm, buttresses present. Bark smooth to fissured or scaly, white to (yellow) grey. Wood hard, (brownish) white. Twigs sparsely hairy, spines absent. Leaves elliptic to ovate, 14 – 28  ×  7 – 16 cm, apex acute to acuminate, base cuneate to cordate, margins entire, sometimes partly lobed; upper surface glabrous, with sometimes a few hairs on main vein, bright green, shiny; lower surface glabrous, sparsely hairy, covered with peltate scales, sometimes with discoid glands at base, grey green; veins 9 – 12 pairs, sometimes 3-veined from base, lower surface with weak tertiary venation, yellow. Petiole 30 – 90 mm long, channelled, glabrous to velutinous, peltate scales present, green to red brown. Inflorescence terminal, 12 – 35 cm long, velutinous, covered with peltate scales; bracteoles lanceolate to spathulate, 4.5 – 30  ×  2 – 10 mm, apex acute, caducous, sessile to petiolate. Calyx 5-lobed, glabrous to densely hairy, covered with peltate scales; lobes few, discoid glands present; flowering calyx 1.5 – 3  ×  2 – 4.5 mm, lobes triangular to linear, 0.5 – 1 mm long, apex acute; fruiting calyx 5 – 9 cm diam., irregularly lobed, patent. Corolla 5-lobed, glabrous to sparsely hairy, white outside, purple base, pink inside, scentless or with sweet smell; anterior lip 3-lobed, mid-lobe rounded to oblong, 6 – 9  ×  3 – 7 mm long, apex rounded to acute, white, sometimes with pink stripes and with a yellow patch at base; side lobes oblong, 4 – 9  ×  2 – 10 mm, sometimes oblique, apex rounded; posterior lip 2-lobed, oblong, 5.5 – 7  ×  4 – 6 mm, apex rounded; tube 7 – 10 mm long. Stamens inserted at apex of tube; anterior pair 10 – 12 mm long, anthers 1 – 1.5 mm long; posterior pair, fertile, 9 – 11 mm long, anthers 1.2 – 1.3 mm long. Ovary glabrous. Style 9 – 11 mm long, stigma 1.5 – 2.5 mm long. Fruit (dried) 6.5 – 11  ×  8 – 16 mm, globose to conical, often with depressed centre when fresh, often pyramid-shaped when dried, glabrous, apex round, black, purple or blue when mature.

distribution. Endemic to the Solomon Islands. Map 6.

selected specimens examined. solomon islands. Guadalcanal: E Guadalcanal, Makina area, 24 Sept. 1968, Mauriasi & collectors BSIP 11275 (K); Gold ridge, 28 July 1945, Walker & White BSIP 59 (K); Gold ridge, 16 Oct. 1962, Whitmore BSIP 649 (K); SE Guadalcanal, Betebete (Avu avu), 23 May 1968, Runikera & collectors BSIP 9723 (K); NW Guadalcanal, Lambi Bay area, 16 April 1968, Gafui & collectors BSIP 9232 (K); Santa Isabel Province: Ysabel, Kolokofa R., 6 April 1966, Teona BSIP 6371 (holotype US, isotype L); Santa Isabel Island, Tiratoña, 8 Dec. 1932, Brass 3309 (holotype BO, isotypes BO, L); Barora Fa Island, Isaisao Pt. Area, 4 Sept. 1969, Mauriasi & collectors BSIP 16588 (K); Santa Isabel Island, Allardyce harbour, 14 April 1965, Bengough BSIP 4525 (K); Santa Isabel Island, Maringe lagoon, near Tiratona village, 22 Oct. 1963, Whitmore BSIP 2301 (K); Kolombangara Island: E coast, Bosua Point, 25 Nov. 1962, Whitmore & Womersley BSIP 851 (K); SE Kolombangara, W of Vila R., 11 Dec. 1967, Dennis & collectors BSIP 8522 (K); Bougainville Island: Karngu, Buin, 10 Oct. 1930, Kajewski 2228 (isotypes L, NY).

habitat. Growing in primary and secondary forest, sometimes well drained, often along ridges, sometimes the co-dominant tree with Campnosperma; alt. 0 – 700 m.

conservation status. Least Concern.

phenology. Flowering and fruiting all year round.

vernacular names.Koko (Tiratoña, Ysabel Island); Arokoko or Arakoko (Kwara’ae Language); Golita (Roviana Language); Karto (Bougainville Island, Karngu); Marovo (Qoliti language).

uses. Wood is used for making canoes.

26. Gmelina schlechteriH. J. Lam (1919: 226); Munir (1984: 106 – 109); Gmelina dalrympleana var. schlechteri (H. J. Lam) Moldenke (1953: 178; 1984f: 39 – 41). Type: Papua New Guinea, Central Province, c. 12 km N of Amazon Bay, Nunumai, 21 June 1969, Pullen 7662 (neotype CANB (n.v.); isoneotypes BO (n.v.), K!, L! × 2 sheets), selected by Munir (1984: 106).

Gmelina palawensis var. novoguineënsis Moldenke (1952: 54 – 55; 1984h: 160), synon. nov. Type: [Papua New Guinea] Morobe Province, Morobe, N. G. F 2922 (holotype BO!; isotypes K!, L!).

Tree 6 – 32 m high, DBH 9 – 60 cm. Bark (grey) brown, scaly or longitudinally fissured. Wood (medium) hard, pale brown to (pale) pinkish. Twigs sparsely hairy when young, with interpetiolar ridge, spines absent. Leaves elliptic to ovate, 10 – 23  ×  4.5 – 14.5 cm, slightly oblique, apex rounded to acuminate, base rounded, rarely cordate or cuneate, margins entire; upper surface glabrous, dull dark green; lower surface glabrous with few simple hairs on veins, peltate scales absent, (yellow) green; two cup-like glands at the base; veins 5 – 8 pairs, red brown, sometimes 3-veined from base. Petiole 1 – 7 cm long, terete to half-terete, red brown, hairy. Inflorescence terminal or axillary, narrow, up to 40 cm long, with long side branches, pale yellow, velutinous; bracteoles lanceolate, 2 – 9  ×  0.5 – 3 mm, apex long acuminate, sessile, with several disc-like glands present. Calyx 2.5 – 3  ×  2.5 – 3 mm, 5-lobed, sometimes appearing fewer, dark purple green, velutinous with sometimes a glabrous margin, outer surface with discoid glands; lobes up to 0.5 mm long, apex rounded to acute; fruiting calyx 5 – 6 mm diam., erect. Corolla 5-lobed, outer surface hairy, sparsely glandular, white to whitish pink or pale purple; anterior lip 3-lobed; mid-lobe oblong, 5 – 8  ×  3 – 5 mm, apex rounded, white to pale purple, with two yellow marks; side lobes 3.5 – 7  ×  2 – 4 mm long, apex round; posterior lip, 2-lobed, 2 – 4  ×  1.5 – 3 mm, apex rounded; tube 8 – 9 mm long. Stamens inserted at apex of tube, glabrous, glands present; anterior pair 10 – 12 mm long, anthers c. 2 mm long, brown; posterior pair, fertile, 9 – 10 mm long, anthers c. 1.5 mm long, brown. Ovary glabrous with a few hairs at apex. Style 10 – 12 mm long, stigma c. 1.5 mm long. Fruit (dried) 9 – 11  ×  5 – 6 mm, clavate, apex rounded, glabrous, sometimes with hairs at apex, often with a ring of scars around apex, purplish-blue to purple (pink) when mature.

distribution. Papua New Guinea: Morobe, Oro (Northern) and Central provinces. Map 3.

selected specimens examined. papua new guinea. Northern Province: N of Ioma, 1 June 1967, Coode & Katik 29970 (K, L); between Ambasi and Devatutu villages, 22 July 1953, Hoogland 3405 (BM, K, L); Morobe Province: Morobe, N. G. F 2922 (holotype BO, isotypes K, L,); Paiawa, 12 May 1970, Johns 47311 (K, L); SE of Lae on the coast, opposite Lasanga Island, 10 Nov. 1973, Jacobs 9536 (L); Buso R., 27 June 1980, Rau 609 (K, L); Kui, 3 Nov. 1966, Henty 28077 (K, L); Saru R., SE of Garaina, 20 July 1970, Streimann and students 47995 (K, L); mouth of Mo R., 31 Jan. 1972, Streimann 24350 (K, L); Kamiali Wildlife Management Area, 19 Feb. 2001, Takeuchi et al. 15114 (K); S of Buso Forestry camp, 30 June 1977, Conn et al. 288 (K, L); Operation Drake Phase IV, 13 Oct. 1979, Wint 47 (K); Lasanga Island: 4 Nov. 1969, Streimann 44257 (K, L); near Anna Village, Opo Creek, 4 March 1978, Kerenga 73871 (K); Central Province: near Marshall Lagoon, Maigo, 18 Oct. 1963, Kairo 17260 (K, L); Kupiano area, Marshall Lagoon, Jan. 1981, Vinas & Nagari 4856 (L); c. 12 km N of Amazon Bay, Nunumai, 21 June 1969, Pullen 7662 (isoneotypes K, L); Papua: Vailala R., 13 Feb. 1926, Brass 959 (K).

habitat. A tree in primary and secondary forest, often along rivers or at the coast, usually in flat lands, rarely on ridges, usually in alluvial soils, usually poorly drained; sometimes over ultra-basic rock; sometimes associated with Anisoptera thurifera (Blanco) Blume or Dillenia nalagi Hoogland; alt. 0 – 300 (− 600) m.

conservation status. Least Concern.

phenology. Flowering and fruiting throughout the year.

vernacular names. Papua New Guinea: Pò’a or Bauma (Orokaiva Language); Boa (Yekora language).

notes. The original type (New Guinea, Kaui Mts, 25 Dec. 1908, Schlechter 17042), seems to be lost and Munir (1984: 106) selected a neotype when he revised the Australian species. The type material of Gmelina palawensis var. novoguineënsis Moldenke falls within the total morphological variation of G. schlechteri H. J. Lam and is thefore place into its synonomy.

27. Gmelina sessilisC. T. White & W. D. Francis (1927: 257 – 259); Moldenke (1984h: 175 – 176). Type: [Papua New Guinea] Papua, Baroi, Oct. 1922, Lane-Poole 303 (holotype BRI (n.v.); isotypes BO! K!).

[Gmelina sessilis C. T. White & W. D. Francis ex Lane-Poole (1925: 136 – 137) nom. invalid.]

Trees 6 – 30 m high, DBH 13 – 75 cm; lightly buttressed. Bark fissured, peeling in irregular flakes, grey brown to brown; slash (orange) brown to red brown. Wood soft, creamy white. Twigs hairy, spines absent. Leaves elliptic to orbicular, often ovate, 14 – 24  ×  10 – 19 cm, apex rounded to acute, base rounded to cuneate, margins entire, often slightly lobed; upper surface glabrous with simple hairs covering the veins, glossy, (dark) green; lower surface covered with simple hairs, with glands clustered around the base, peltate scales absent, dull, pale green or glaucous; veins 5 – 7 pairs, yellowish, sometimes 3-veined from base, lower surface with strong tertiary venation. Petiole half-terete, slightly channelled, 4 – 7 cm, hairy. Inflorescence terminal, narrow, 13 – 21 cm, velutinous; bracteoles rounded, 13 – 17  ×  11 – 13 mm wide, apex acuminate to rounded, sessile, with discoid glands. Calyx 4 – 6  ×  3.5 – 5 mm, 5-lobed, sometimes appearing fewer, velutinous, green, outer surface with discoid glands; lobes 1 – 2  ×  1 – 1.8 mm, apex acute; fruiting calyx c. 7 mm diam., erect to patent. Corolla 5-lobed, cream to light purple, with darker markings, outer surface velutinous, glabrous at base, inside glabrous, sparsely glandular; anterior lip 3-lobed; mid-lobe spathulate to oblong, 4 – 8  ×  3 – 4 mm, apex rounded, yellow patch at base; side lobes 4 – 5  ×  2.5 – 3 mm, apex rounded, oblique; posterior lip 3 – 5  ×  2 – 8 mm, apex rounded, erect; tube 6 – 11 mm long. Stamens inserted at apex of tube, white; anterior pair 5 – 14 mm long, anthers c. 1.5 mm long; posterior pair, fertile, 4 – 10 mm long, anthers c. 1.5 mm long. Ovary glabrous; style c. 12 mm long, stigma c. 2 mm long. Fruit (dried) c. 12  ×  10 mm, cylindrical, apex truncate, glabrous, bright blue to glossy violet.

distribution. New Guinea: Northern, Milne Bay, Gulf and New Britain Provinces. Map 4.

specimens examined. papua new guinea. Papua: Baroi, Oct. 1922, Lane-Poole 303 (isotypes BO, K); Northern Province: Samboga R. area, Dododura Plain, 16 Feb. 1945, anonymous 2005 (K, L); Milne Bay Province: W of Opanabu, 24 July 1969, Kanis 1305 (K, L); Gulf Province: Middle Tauri R. 14 March 1966, Craven & Schodde 1021 (K, L); New Britain Province: Talasea subdistr., 1 mile E of Lavege Village, 17 April 1959, White 10883 (K, L, SING); junction of Tauri and Kapau Rs, near Putei, 9 March 1966, Craven & Schodde 974 (L).

habitat. Growing in primary to secondary forest; alt. 30 – 60 m. Sometimes associated with Artocarpus sp., Evodia sp. Maniltoa sp., Planchonia sp., Parinari sp., Pterocymbium sp., Sterculia sp. and Tetrameles nudiflora R. Br. Soil alluvium or grey sandy loam.

conservation status. Least Concern.

phenology. Flowering March – April; fruiting Feb. – May.

notes. Lane-Poole (1925) was the first one to use the name Gmelina sessilis, but from his description, it is clear that he did not intend to formally describe it and he must have known that C. T. White & W. D. Francis were to make a formal description at a later date. He has included no Latin diagnosis, does not select a type, and gives C. T. White & W. D. Francis as the authors of the name.

28. Gmelina smithiiMoldenke (1958: 326 – 327; 1984h: 180 – 181). Type: Papua New Guinea, Central Highlands [Madang Province, Aiyura, Oct. 1944, Smith 1064 (holotype LAE scan seen; isotypes K!, L!).

Trees 10 – 45 m high, DBH 10 – 100 cm. Bark fissured to smooth, pale blue-grey or whitish brown. Wood soft, pale yellow brown. Twigs hairy, interpetiolar ridge present, spines absent. Leaves elliptic to obovate or ovate, 7 – 13.5  ×  4 – 8 cm, apex rounded to acute, base rounded to cuneate, margins entire; upper surface glabrous, (pale olive) green, glossy; lower surface glabrous, pale green to pale bronze, with some discoid glands clustered around the base, covered with peltate scales; veins 6 – 11 pairs. Petiole half-terete, channelled, 1 – 2.5 cm, glabrous with few hairs at base. Inflorescence terminal, 7 – 19 cm, velutinous; bracteoles ovate to linear, 5 – 7  ×  2 – 5 mm, decreasing in size and particularly in width towards base, apex acute, persistent, blackish. Calyx 5-lobed, velutinous; flowering calyx 3.5 – 5  ×  3.5 – 5 mm; lobes 1 – 2.5  ×  1 – 1.2 mm; apex acute to rounded; fruiting calyx 9 – 10 mm diam., patent. Corolla 5-lobed, white to purple or red, outer surface hairy, sparsely glandular; anterior lip 3-lobed; mid-lobe 4 – 6  ×  3.5 – 5 mm, oblong, apex rounded, pale blue with yellow spots and lines, sometimes with a purple apex; side lobes 3 – 5  ×  4 – 5 mm, apex rounded to acute; posterior lip, 2-lobed, 3 – 6  ×  3 – 5 mm, apex rounded; tube 3 – 6 mm long. Stamens inserted at apex of tube; anterior pair 8 – 10 mm long, anthers 1.5 – 2 mm long; posterior pair, fertile, 8 – 10 mm long, anthers 1.5 – 2 mm long. Ovary glabrous; style 7 – 14 mm long, stigma 1.5 – 2 mm long. Fruit (dried) 7 – 12 × 10 – 15 mm, globose, apex rounded, glabrous, red to pink when mature.

distribution. Papua New Guinea: Morobe, Eastern Highlands and Madang Provinces. Map 6.

specimens examined. papua new guinea. Eastern Highlands Province: c. 1 mile S of Akuna, 19 July 1963, Hartley 12065 (K, L); Anumba to Ilafo, near Okasa, 24 May 1967, Coode & Lelean 29938 (K, L, SING); Madang Province: Aiyura, Oct. 1944, Smith 1064 (holotype LAE, isotypes K, L); Morobe Province: Menyi Village, Slate Creek, 3 August 1982, Streimann 8488 (K, L); Bulolo-Watut divide, above Taun creek, 24 Aug. 1964, Kairo 25552 (K, L, SING); Wau sub-distr., Eraulu logging area, 13 June 1974, Katik & Eddowes 62098 (K, L).

habitat. Growing in lower or mid mountain rainforests; alt. 1525 – 1980 m. Sometimes associated with Fagaceae forest or Cinnamomum sp., Elmerrillia sp. and Symplocos sp.

conservation status. Least Concern.

phenology. Flowering and fruiting May – Oct.

vernacular names. Madang area: Yabeina (Anona Language), Kaboya (Aiyura Language); Eastern Highland area: Wakhai (Fore Language).

29. Gmelina tholicolaMabb. in Mabberley & de Kok (2004: 27). Type: New Caledonia, Néhoué, Valléede la Rade, 20 Jan. 1979, MacKee 36450 (holotype P, scan seen).

conservation status. Least Concern.

note. For a description and more information see Mabberley in Mabberley & de Kok (2004).

30. Gmelina unifloraStapf (1895: t. 2391); Lam (1919: 217); Lam & Bakhuizen (1921: 65). Gmelina uniflora var. typica Bakh. in Lam & Bakhuizen (1921: 66), nom. superfl. Type: [Indonesia], South Borneo, Banjermassing [Banjarmasin], 1857 – 1858, Motley 1204 (holotype K!).

Gmelina uniflora var. villosa Bakh. in Lam & Bakhuizen (1921: 66 – 67). Type: [Indonesia] Borneo, Soengei Bloe-oe, Jaheri exp. Nieuwenhuis 1109 (syntype BO (n.s)); [Indonesia] Borneo, Soengei Bloe-oe, Jaheri exp. Nieuwenhuis 1300 (syntype BO (n.v.), isosyntypes L! NY scan seen); [Indonesia] Borneo, Soengei Bloe-oe, 1896 – 1897, Jaheri exp. Nieuwenhuis 1463 (syntype BO (n.v.); isosyntype L!).

Small tree or liana, 4 – 30 m high, DBH 8 – 30 cm. Bark light to reddish brown, smooth; sapwood white to pale orange; twigs hairy, spines absent. Leaves broadly elliptic or obovate-elliptic, 10 – 20  ×  6.5 – 12 cm, apex obtuse or shortly acuminate, papery, base rounded to cuneate, margins entire; upper surface glabrous, green to dark green; lower surface covered with peltate scales, sometimes sparsely hairy, light green to grey-green or silvery, sometimes with glands at base present; veins 4 – 6 pairs, sometimes 3-veined from base. Petiole 1.5 – 4.5 cm long, half terete, glabrous. Inflorescence 2.5 cm long, usually consisting of a single or only a few flowers, axillary, velutinous, covered with peltate scales; bracteoles 12 – 14  ×  3 – 4 mm, lanceolate, apex acuminate. Calyx 4 – 5-lobed, 14 – 20  ×  7 – 10 mm, velutinous, dark brown to pale green, becoming pale yellow when in fruit, sometimes with glands, slightly accrescent; lobes 7 – 10 mm long, apex acuminate. Corolla 5-lobed, white to pale yellow, yellow guides in throat, velutinous; lip 3-lobed, mid-lobe 5 – 13  ×  8 – 12 mm, spathulate, apex rounded, glabrous; side lobes 4 – 10  ×  4 – 8 mm, apex rounded; posterior lip 2-lobed, lobes 5 – 7  ×  6 – 7 mm, apex rounded; tube 10 – 32 mm long. Stamens inserted at apex of tube, white; anterior pair 10 – 18 mm long, anthers 2.5 – 3 mm long; posterior pair, fertile, 11 – 13 mm long, anthers 2 – 2.5 mm long. Ovary glabrous with apex covered with hairs; style 15 – 30 mm, long, stigma c. 3 mm long. Fruit 2.5 – 4  ×  1.5 – 2.5 cm, clavoid, apex rounded, glabrous, covered with scales when young, smooth, (pale) yellow.

distribution. Endemic to Borneo. Map 6.

selected specimens examined. brunei. Temburong, Batu Apoi, 30 Jan. 1994, Dransfield 7488 (K). INDONESIA. Kalimantan: Soengei Bloe-oe, Jaheri exp. Nieuwenhuis 1300 (isosyntypes L, NY); Soengei Bloe-oe, 1896 – 1897, Jaheri exp. Nieuwenhuis 1463 (isosyntype L); Central Kalimantan: Kotawaringin Timur, 62 km from Sangai, 28 Sept. 1996, Argent et al. 9627 (K); Kalimantan: KTC, Camp Buntut Emban, 30 Jan. 1983, Wiriadinata 3484 (L); Kalimantan Timur: Around Jelini, along Sungai Belayan, NW of Tabang, 7 Jan. 1979, Murata et al. 1148 (L); South Kalimantan: Banjermassing [Banjarmasin], 1857 – 1858, Motley 1204 (holotype K). MALAYSIA. Sarawak: Kelabit Highlands, Pa Dalih, 4 Oct. 1993, Christensen 531 (K); Kapit Distr., Balleh R., Balang/Balleh watershed, 7 July 1969, Anderson & Ilias S. 28480 (K); Kuching, 1 Nov. 1894, Haviland & Hose 919 (BM, K); Sabah: Sepulut, Iabang, 14 Oct. 1988, Fidilis 125236 (K); Lahad Datu, Malambabula Nam Hing Co, 28 Nov. 1962, Muin Chai 31738 (K).

habitat. Growing in primary and secondary forest; alt. 0 – 1100 m. Soil (sandy) clay or loam, sometimes over sandstone, shale, limestone or ultramafic.

conservation status. Least Concern.

phenology. Flowering Jan. – Sept.; fruiting Aug. – March.

vernacular names. Sarawak: Susu-susu (Kedayan Language); Akar enkeliae or Akar ankaleh (Iban language); War Imang Bayang (Kelabit Language). Brunei: Akar inklis (Iban language) or Paginggi (Brunei language).

uses. In Sarawak it is used for treating goitres, by first pounding the leaf and rubbing the paste on the affected part. It is also used for soap by crushing the bark and add water. The fruits are reported to be eaten in Brunei.

31. Gmelina vitiensis(Seem.) A. C. Sm. (Smith 1978: 414, 1991: 205 – 206, fig. 23); Munir (1984: 110); Gmelina vitiensis Seem. (Seemann 1862: 440), nom. nud.; Vitex vitiensis Seem. (Seemann 1866: 190 – 191, t 45); Munir (1984: 110 – 113). Type: [Fiji] Viti, Ngau, Oct. 1855, (HMS Herald) Milne 224 (lectotype K!; isolectotype K!), selected here.

Trees 12 – 30 m high, DBH 28 – 75 cm, buttresses sometimes present. Bark smooth to fissured or scaly, brownish white. Wood white to creamy coloured, hard. Twigs brown tomentose to glabrous, spines absent. Leaves ovate to ovate- lanceolate, 3 – 15  ×  1.5 –14 cm, apex rounded to acute, base rounded to cuneate or slightly cordate, margins entire, papery, glabrous or with a few hairs on veins; upper surface green; lower surface discoid glands sometimes present, covered with peltate scales, grey; veins 7 – 12 pairs. Petiole 1 – 4 cm, half-terete, shallowly channelled, sparsely hairy. Inflorescence terminal or axillary, narrow, 5 – 17 cm long, glabrous to densely hairy, covered with peltate scales; bracteoles elliptic, 8 – 15 mm long, petiolate, caducous. Calyx 5-lobed, glabrous, covered with peltate scales, outer surface with scattered discoid black glands; flowering calyx 2.5 – 4  ×  2.5 – 5 mm; lobes 0.3 – 1  ×  0.5 – 1.2 mm, apex acute; fruiting calyx 7 – 8 mm diam., irregularly lobed. Corolla 5-lobed, blue, outer surface velutinous, sparsely glandular; anterior lip 3-lobed; mid-lobe oblong to spathulate 5 – 8.5  ×  3 – 6 mm, apex rounded; side lobes oblong, 4 – 5  ×  3 – 6 mm, apex round; posterior lip, 2-lobed, 3 – 5  ×  2 – 4 mm, apex rounded; tube 2 – 10 mm long. Stamens inserted at apex of tube, white; anterior pair 7.5 – 8.5 mm long, anthers c. 1 mm long; posterior pair, fertile, 6 – 7 mm long, anthers c. 0.6 mm long. Style 7 – 9 mm long, stigma c. 2 mm long. ovary c. 1 mm diam., apex with few hairs. Fruit 9 – 12  ×  10 – 13 mm, globose to ovoid, apex rounded, glabrous, purple or mauve when mature, shiny.

distribution. Fiji. Map 1.

selected specimens examined. fiji. Viti, Ngau, Oct. 1855, (HMS Herald) Milne 224 (lectotype K, isolectotype K); Sasa logging area, 18 Sept. 1968, Howard H 181 (K); Sertua, Korovou hill, 1961, Bola K.V. 1 (K); Sotava labasa, 11 May 1962, Damanu 55 (K); Nadroga & Navosa, Nausori Highland, 25 June 1970, Nasogiri 17354 (K).

habitat. Growing in primary and secondary forests; alt. 0 – 900 m. Soil well-drained.

conservation status. Least Concern.

phenology. Flowering and fruiting throughout the year.

vernacular names. Fiji: Rosawa or Sevua.

uses. The wood is used to make furniture (Smith 1991).

notes. This species was considered conspecific with Gmelina dalrympleana by Munir (1984); he also stated that Gmelina is not native to Fiji and that no specimens from wild populations are known to exist. He postulated that the type of G. vitiensis must have come from a plant that was introduced to Fiji. This was contested by Smith (1991) who claims that the species is abundant on several separate islands in the Fiji archipelago and occurs often in dense and undisturbed forest.

In this study, Gmelina vitiensis was found to differ from G. dalrympleana in having a lower leaf surface covered with peltate scales and by the absence of two cup-shaped glands at the base of the leaves. Furthermore, it is clear that G. vitiensis is not only endemic to Fiji, but that there are many specimens at K that clearly have come from wild sources. I therefore agree with Smith’s (1991) conclusion. There are two specimens labelled ‘Viti, Ngau, October 1855, (HMS Herald) Milne 224’ at K. The specimen with the original label is designated here as the lectotype.

Dubious taxa

Gmelina coromandeliana Burm. f. (Burman 1768: 132) Rajendran & Daniel (2002: 162). Type: Plukenet (1691: 234, t. 97, f 2) and Sloane (1725: 100, t 207, f 1).

This name is based on a plate in Plukenet (1691) and a copy of this figure in Sloane (1725). These plates depict a sterile specimen with opposite simple leaves and long spines, which could be Gmelina asiatica as Rajendran & Daniel (2002) believe, but it could just as easily be a species from one of several other families.

Gmelina inermis Wight ex Wall. (Wallich 1828: 1816d), nom. nud.; Gmelina asiatica L. forma inermis (Wight ex Wall.) Moldenke (1984a: 42). Based on [Indonesia] Madura, Wallich 1816d (holotype not located).

No material on which this name is based could be found. All other material seen with the Wallich number 1816 is Gmelina elliptica, and given the locality cited, this identification is also possible for the 1816d specimen. However, without a specimen no definite identification can be made.

Gmelina lobata Gaertn. (Gaertner 1791: 268); Lam & Bakhuizen (1921: 69); Gmelina elliptica forma lobata (Gaertn.) Moldenke (1953: 178; 1984g: 103 – 106). Type material: Gaertner (1791: Pl. 56, Fig. 2) and Rumphius (1750: 1, p 129, t 40)  =  Gmelina asiatica or G. elliptica.

This name is based on two different elements. The plate in Rumphius is Gmelina elliptica L., while the plate and description in Gaertner (1791: 268, t 56, Fig. 2) depicts a Gmelina, but which of the two possible species, G. asiatica or G. elliptica, is depicted is impossible to determine.

Gmelina palawensis var. dingatensis Moldenke (1951: 418; 1984h: 160). Type: Philippines, Mindanao, Suirago, Dingat, 1902, Ahern 461Q (holotype BO!; isotype BO!).

I have found it impossible to identify the collections on which this name is based. The original description is very short and only refers to characters common to many species. Futhermore, there is a problem with the remaining type material. In the original description only one collection is cited: Philippines, Dingat, 1901 or 1902, Ahern 461Q (BO × 2) in Bogor. In January 1952 on a det. label on a herbarium specimen deposited in NY, Moldenke also cites the collection: Philippines, Mindanao, Suirago, 1902, Ahern 461 (NY 001375580 scan seen) as part of the type material. The collection locality of the latter specimen is problematic and most likely refers to Surigao in the north-east of the Island of Mindanao. This would mean that we are talking about two different collections, from two different islands, and the latter cannot be considered as type material. Taking only the remaining type material into account, this name could refer to a Gmelina (but given the locality this seems unlikely), or to a species of Faradaya F. Muell. or Clerodendrum L.

Gmelina sinuata Link (1822: 128); Moldenke (1984h: 179 – 180). Type: ?

This name is based on a specimen growing in the Berlin Botanic Gardens in the early 1800s. The description is inadequate to determine the identity of the plant, even to family level. No specimen could be found associated with this name and subsequently its identity remains unknown

Excluded species

Gmelina indica Burm. f. (Burman 1768: 132, t. 39, Fig. 5). 1768; Lam (1919: 228). Gmelina javanica Christm. (Christmann 1777: 134 ), nom. superfl. Type: Burman (1768: 132, t. 39, Fig. 5)  =  Flacourtia indica (Burm. f.) Merr. (see Sleumer 1954: 76).

Gmelina siamica Moldenke (1955: 226). Type: north-eastern Thailand, Loei, Phu Krading, Sala Chomphon, 13 Sept. 1954, Smitiand 1920 (holotype not located; isotype BKF (n.v.), L!)  =  Wightia speciosissima (D. Don) Merr. (see Yamazaki 1985: 15).

Gmelina speciosissima D. Don (1825: 104 – 105); Gmelina tacabushia Buch.-Ham. ex D. Don (1825: 105). Type: Nepal, ‘Gmelina Taeabushia’ Buchanan-Hamilton MSS (holotype not located). = Wightia speciosissima (D. Don) Merr. (see Yamazaki 1985: 15).

Acknowledgements

The author is grateful to the curators of B, BM, BO, BR, CAVA, E, L, LINN, MEL, NY, P, SING and US for access to their collections. The drawings were by Juliet Beentje, the Latin diagnoses were written by Melanie Thomas. The author is grateful for the useful comments made by David Mabberley, Ian Turner and Gemma Bramley on an earlier version of this paper.

Copyright information

© The Board of Trustees of the Royal Botanic Gardens, Kew 2012

Authors and Affiliations

  1. 1.Royal Botanic Gardens, KewRichmondUK

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