Flora of Thailand
Euphorbiaceae
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Identification key for staminate material
Identification key for pistillate and fruiting material
Burm. ex L., Sp. Pl.: 1027. 1753; Gen. Pl. ed. 5: 451. 1754; G.L.Webster, Ann. Missouri Bot. Gard. 81: 52. 1994; Radcl.-Sm., Gen. Euphorbiacearum: 69. 2001; Petra Hoffm. in Chayam. & Welzen, Fl. Thailand 8, 1: 51. 2005; Antidesma Malesia Thailand: 25. 2006; G.L.Webster in Kubitzki, Fam. Gen. Vasc. Pl. 11: 81. 2014. Bestram Adans., Fam. 2: 354. 1763. Stilago L., Mant. Pl.: 16. 1767. Rhytis Lour., Fl. Cochinch.: 660. 1790.
Dioecious shrubs and trees. Indumentum simple. Petiole terete or channelled adaxially, sometimes basally and distally swollen and geniculate. Stipules caducous or persistent, sometimes foliaceous, entire. Leaves evergreen or deciduous, alternate, simple, eglandular, blades symmetrical, entire, pinnately veined, basally sometimes decurrent, apically usually mucronate, membranaceous to coriaceous, domatia present or absent. Inflorescences raceme-like, axillary or cauline, erect to pendulous, infructescences usually pendulous. Bracts 1 per flower. Pedicel short to absent, inarticulate. Flowers usually light yellowish green, often turning red when mature. Sepals 3-8, imbricate in bud, fused to various degrees. Petals absent. Staminate flowers: disc cushion-shaped (enclosing the bases of the stamens and pistillode), extrastaminal-annular or consisting of free alternistaminal lobes; stamens 2-8, usually antisepalous, filaments free, anthers extrorse in bud, versatile, connective elongated, U-shaped, thecae resembling swollen ends of the U, raised at anthesis, longitudinally dehiscing, pistillode present or absent. Pistillate flowers: disc annular; ovary 1-locular; ovules 2; stigmas 2-6, acute. Fruits drupaceous, globose to ellipsoid or laterally compressed (lens-shaped), base symmetrical to asymmetrical, style terminal to lateral, maturing unevenly depending on position in infructescence and exposure to the sun from green through white and red to black, sometimes glaucous or mottled with grey, smooth, shiny, fleshy, soft, juicy, edible, sour to bitter-sweet; endocarp lignified, foveolate, the sculpture usually obvious through the dried mesocarp. Seeds ecarunculate, 1 per fruit (rarely two, then fruits twice as big); endosperm in mature seeds present, cotyledons thin, flat, several times wider and longer than radicle.
About 100 species in the Old World Tropics, with the highest number in South-East Asia. 18 species in Thailand. Classification: Subfamily Phyllanthoideae, subtribe Antidesminae (Webster, 1994).
Notes: Specimens should either be in fruit or bear mature staminate flowers. Pistillate flowering or sterile material may be difficult and sometimes impossible to determine. Descriptive terms are illustrated in fig. 1 and 2. Many structures can only be seen in sufficient detail under a dissecting microscope; dried flowers may have to be softened. All measurements are given in the dry state; fruit length includes calyx and stigmas. Species in parentheses key out in this place only as an exception.
Key for Staminate Material
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1a. |
Leaf apex rounded or retuse, more rarely obtuse. |
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1b. |
Leaf apex acute, acuminate or caudate. |
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Leaf base acute or obtuse, mostly cuneate. Stamens 2 (or 3). Sepals fused for 2/3rd of their length, glabrous outside. Stipules 0.5-1 mm wide. Petiole 1-2 mm wide. Inflorescences simple to branched twice. Flowers 1-1.5 mm long pedicellate. Disc cushion-shaped (Fig. 4a). Pistillode absent. |
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2b |
Leaves base cordate to rounded, very rarely acute. Stamens 4-6. Sepals free, pubescent outside. Stipules 1-2 mm wide. Petiole 0.7-1 mm wide. Inflorescences consisting of 10-20 racemes. Flowers sessile. Disc consisting of free lobes (Fig. 4c). Pistillode present. |
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Midvein in sicco sharply raised adaxially (Fig. 3c), distinctly perceptible to the touch. |
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3b. |
Midvein in sicco impressed to gently raised adaxially (Fig. 3a, b, d), not distinctly perceptible to the touch. |
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Leaves 6-7.5 cm long, 2.5-3.5 times longer than wide, chartaceous to subcoriaceous. Indumentum of stipules, inflorescence axis and bracts whitish. Stipules 2-5 mm long. |
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4b. |
Leaves 13-27 cm long, 4-5.5 times longer than wide, coriaceous. Indumentum of stipules, inflorescence axis and bracts ferrugineous. Stipules 4-10 mm long. |
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Inflorescences at least partly cauline. |
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5b. |
Inflorescences axillary (sometimes terminally condensed). |
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Petiole (at least of some leaves) longer than 10 mm, 1-1.5 mm wide, basally and distally usually slightly swollen and sometimes geniculate. |
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6b. |
Petiole shorter than 10 mm (if longer, then wider than 1.5 mm), basally and distally not swollen or geniculate. |
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Tertiary veins close together, more than 10 between every two secondary veins (Fig. 1a). Petiole 1.5-4 mm wide. Inflorescences (4-)15-24 cm long. Leaves (7-)13-24(-50) by (2.5-)4-8(-15) cm. |
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7b. |
Tertiary veins widely spaced, less than 9 between every two secondary veins (Fig. 1b). Petiole up to 2 mm wide. Inflorescences 3-8 cm long. Leaves (5-)8-14(-19) by (2-)3-5(-7.5) cm. |
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Stipules, petiole and lower leaf side hairy all over. Indumentum ferrugineous. Inflorescences branched, consisting of 4-11 racemes. Disc extrastaminal-annular (Fig. 4b; but may sometimes appear cushion-shaped, Fig. 4a), glabrous (sometimes hairs originating from the base of the disc; Fig. 5a). Pistillode hairy. |
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8b. |
Stipules, petiole and lower leaf side glabrous. Indumentum whitish or absent. Inflorescences simple or branched once. Disc cushion-shaped (Fig. 4a), hairy (Fig. 5b, c). Pistillode glabrous. |
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Stipules foliaceous, more than 4 mm wide. |
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9b. |
Stipules not foliaceous, up to 4 mm wide. |
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Disc pilose (Fig. 5b, c). Flowers 2-3 by 1.5-2 mm. Stamens 2-2.5 mm long, 1.5-2 mm long exserted from the calyx. Stipules, petiole and leaves glabrous. |
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10b. |
Disc glabrous (Fig. 5a), rarely pilose (Fig. 5b, c). Flowers 1-2.5 by 1-1.5 mm. Stamens 0.8-2 mm long, 0.5-1.5 mm long exserted from the disc. Stipules, petiole and major veins on lower leaf surface pilose. |
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Leaves drying olive-green. Plant with whitish indumentum or glabrous, indumentum never ferrugineous. Inflorescence much-branched, consisting of 10 or more racemes. Flowers 1-2.5 mm long. Stamens 1-2 mm long. Sepals often glandular-fimbriate. |
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11b. |
Leaves drying reddish brown. Plant with ferrugineous indumentum, at least at the tip of the branchlets. Inflorescences simple or branched, consisting of up to 7 racemes. Flowers 1 mm long. Stamens 0.8-1 mm long. Sepals never glandular-fimbriate. |
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Bracts longer than 1.5 mm, conspicuous even in very young inflorescences. Sepals 5-7, free. Disc cushion-shaped, enclosing the bases of the filaments and pistillode (Fig. 4a), glabrous (Fig. 5a). Indumentum especially of young plant parts densely hirsute and never ferrugineous. Leaves abaxially hairy, drying olive-green. |
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12b. |
Bracts up to 1.5 mm long. Leaves and flowers various. |
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Stamens 2, rarely 3. Pistillode absent. Pedicels 1-1.5 mm long. |
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13b. |
Stamens 3-5. Pistillode present or absent. 0-1.5(-2) mm long. |
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Disc glabrous (sometimes hairs at base of calyx; Fig. 5a). |
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14b. |
Disc hairy all over or at least at the margin (Fig. 5b, c). |
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Flowers 3-4 by 3 mm, sessile, smelling strongly of bats droppings (unpleasant cheesy smell), attracting flies. Stamens 1.5-2 mm long exserted from the calyx. Anthers 0.5 by 0.7 mm. Pistillode present. |
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15b. |
Flowers 1-2.5 by 1-2 mm, sessile or pedicellate, without unpleasant cheesy smell. Stamens 0.5-2 mm long exserted from the calyx. Anthers various. Pistillode present or absent. |
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Indumentum lacking or whitish. Abaxial leaf surface glabrous except sometimes along the midvein. Often cultivated. |
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16b. |
Indumentum ferrugineous. Abaxial leaf surface pubescent all over. Rare. |
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Disc extrastaminal-annular (Fig. 4b). |
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17b. |
Disc consisting of free lobes (which are short and may appear to be fused extrastaminally in A. pendulum; Fig. 4c) or disc cushion-shaped (Fig. 4a), enclosing the bases of the filaments and, if present, pistillode (observe carefully, dried flowers may have to be softened). |
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Stipules 1-3 by 0.2-0.5 mm. Petiole 0.7-1 (-1.2) mm wide. Indumentum ochraceous. Leaves abaxially pubescent all over or only along the veins, up to 13 cm long, drying olive-green, greyish or reddish brown. Inflorescences 4-11 cm long, terminal or more rarely axillary. |
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18b. |
Stipules 3-7 by 0.5-1 mm. Petiole 1.2-1.5 mm wide. Indumentum ferrugineous. Leaves abaxially pubescent all over, distinctly velvety to the touch, up to 18 cm long, always drying reddish brown. Inflorescences 4-7 cm long, axillary or more rarely cauline. |
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Disc consisting of free lobes (Fig. 4c). |
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19b. |
Disc cushion-shaped, enclosing the bases of the filaments and pistillode (Fig. 4a). |
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Petiole 1.5-6 mm wide, glabrous. Leaves (13-)25-50 by (4-)7-14 cm, chartaceous to coriaceous, usually drying yellowish-brown. Inflorescences c. 15 cm long, simple, solitary or fascicled in twos, axis glabrous. Pistillode glabrous. Stipules usually persistent, 0.5-3 mm wide. Midvein usually gently raised adaxially (Fig. 3b). |
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20b. |
Petiole 0.5-1.2 mm wide, hairy. Leaves 3-11(-16) by 1-4(-5.5) cm, chartaceous to membranaceous, drying olive-green, reddish or greyish brown. Inflorescences 2-11 cm long, simple or branched, axis hairy. Pistillode pubescent. Stipules early caducous, 0.2-0.5 mm wide. Midvein impressed adaxially (Fig. 3a). |
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Inflorescences 2-4 cm long. Young twigs first dark brown with conspicuous lenticels, then light grey to whitish, glabrous to puberulent. Stamens 0.5 mm long exserted from the calyx. Petiole (4-)6-17 mm long. Leaves glabrous or slightly hairy only along the midvein, higher venation not conspicuously tessellated. |
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21b. |
Inflorescences 4-11 cm long. Young twigs brown, (usually densely) ochraceous-tomentose. Stamens 1-2 mm long exserted from the calyx. Petiole 2-11 mm long. Leaves abaxially hairy at least along the veins, higher venation finely tessellated. |
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Indumentum especially of young plant parts and inflorescences ferrugineous to ochraceous. |
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22b. |
Indumentum whitish or absent. |
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Petiole 1 mm wide. Leaves (6-)9-14(-20) by up to 7 cm, tertiary veins weakly percurrent (Fig. 1b) to reticulate (Fig. 1c). Inflorescences 2-6 cm long, simple or branched, very slender, with flowers more widely spaced, not touching each other. Bracts 0.5 by 0.3 mm. Disc glabrous (Fig. 5a). Pistillode absent or tiny, up to 0.1 mm long, glabrous to sparsely pilose. |
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23b. |
Petiole 2-5 mm wide. Leaves (8-)15-30(-55) by up to 25 cm, tertiary veins percurrent, close together (Fig. 1, see a, d resp.). Inflorescences 6-14 cm long, always simple, mostly thick, with flowers so densely arranged that they touch each other. Bracts 0.7-1.5 by 0.5-1 mm. Disc glabrous (Fig. 5a) or pubescent (Fig. 5b, c). Pistillode present, 0.2-0.4 mm long, pubescent. |
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Plant rheophytic, leaves 4-10 times longer than wide or narrower than 2 cm. |
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24b. |
Plant not rheophytic, leaves (1.2-)2.5-3.5(-6.6) times longer than wide. |
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Average leaf size 5-11 by 1-2 cm. Leaf apex acuminate. |
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25b. |
Average leaf size 3-6 by 0.4-1.2 cm. Leaf apex acute, sometimes rounded at the very tip. |
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Inflorescence, pedicels and flowers slender, usually very regular. Petiole 0.5-1 mm wide. Sepals free. Leaves chartaceous to subcoriaceous, often conspicuously shiny, narrowly elliptic-oblong to ovate or obovate, average leaf size 6-10 by 2-3.5 cm. |
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26b. |
Inflorescence, pedicels and flowers not conspicuously slender. Petiole 0.7-2 mm wide. Sepals free to halfway fused. Leaves variable, not conspicuously shiny. |
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Disc consisting of free lobes (Fig. 4c). Pedicel 0. |
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27b. |
Disc cushion-shaped, enclosing the bases of the filaments and pistillode (Fig. 4a). Pedicel various. |
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Inflorescence c. 15 cm long. Pistillode glabrous. Petiole 1.5-6 mm wide. Stipules 6-25 by 0.5-3 mm, usually persistent. Leaves (13-)25-50 by (4-)7-14 cm. |
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28b. |
Inflorescence 4-12 cm long. Pistillode hairy. Petiole 0.7-1.5 mm wide. Stipules 2-6 (-10) by 0.5-1 mm, usually caducous. Leaves 2-15(-20) by 2-4.5(-9) cm. |
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Sepals free, pubescent outside. Young twigs brown. Leaves 1-2.25 times longer than wide, base rounded to cordate, rarely acute. Inflorescences 4-8 cm long, much-branched, consisting of 10-20 racemes. |
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29b. |
Sepals fused for 1/2 of their length or more, glabrous to sparsely pilose outside. Young twigs whitish. Leaves 2-4.7 times longer than wide, base acute to rounded. Inflorescences 6-12 cm long, simple or branched once. |
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Indumentum especially of young plant parts, inflorescences and disc ferrugineous to ochraceous. Petiole 2-5 mm wide. Leaves 8-55 cm long, drying reddish brown. Flowers sessile. Stamens 0.5 mm long exserted from the calyx. |
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30b. |
Indumentum whitish or absent. Petiole 0.7-2 mm wide. Leaves 1.5-30 cm long, drying various colours. Flowers sessile or pedicellate. Stamens 0.5-2 mm long exserted from the calyx. |
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Young twigs light grey, glabrous or nearly so. Leaves drying greyish green to reddish brown, chartaceous to coriaceous. Anthers 0.5-1 mm long and wide. Sepals never glandular-fimbriate. Flowers 2-3 mm long. |
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31b. |
Young twigs brown, ± densely hairy. Leaves usually drying olive-green, membranaceous to chartaceous, only larger leaves sometimes coriaceous. Anthers 0.2-0.5 mm long and wide. Sepals often glandular-fimbriate. Flowers 1-2.5 mm long. |
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Lower leaf side patulously hirsute all over, especially along the veins. |
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32b. |
Lower leaf side glabrous except along the major veins. |
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Plant not rheophytic, leaves 1-3.5(-6.6) times longer than wide. |
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33b. |
Plant rheophytic, leaves 4-10 times longer than wide or narrower than 2 cm. |
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Average leaf size 5-11 by 1-2 cm. Leaf apex acuminate. |
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34b. |
Average leaf size 3-6 by 0.4-1.2 cm. Leaf apex acute, sometimes rounded at the very tip. |
Key for Pistillate and Fruiting Material
See notes under Key for Staminate Material.
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1a. |
Leaf apex rounded or retuse, more rarely obtuse. |
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1b. |
Leaf apex acute, acuminate or caudate. |
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2a. |
Leaf base acute or obtuse, mostly cuneate. Sepals fused for 2/3rd of their length, glabrous outside. Ovary glabrous. Stipules 0.5-1 mm wide. Petiole 1-2 mm wide. Fruiting pedicel 1.5-3 mm long. Inflorescences simple to branched twice. |
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2b. |
Leaf base cordate to rounded, very rarely acute. Sepals free, pubescent outside. Ovary pubescent. Stipules 1-2 mm wide. Petiole 0.7-1 mm wide. Fruiting pedicel 0-1 mm long. Inflorescences consisting of (1-)10-20 racemes. |
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3a. |
Midvein in sicco sharply raised adaxially (Fig. 3c), distinctly perceptible to the touch. Inflorescences simple. |
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3b. |
Midvein in sicco impressed to gently raised adaxially (Fig. 3a, b, d), not distinctly perceptible to the touch. Inflorescences various. |
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4a. |
Leaves 6-7.5 cm long, 2.5-3.5 times longer than wide, chartaceous to subcoriaceous. Indumentum of stipules, inflorescence axis and bracts whitish. Ovary glabrous. Fruit 5-6(-8) by 4-5(-6) mm, glabrous. Infructescence 4-8 cm long. Stipules 2-5 mm long. |
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4b. |
Leaves 13-27 cm long, 4-5.5 times longer than wide, coriaceous. Indumentum of stipules, inflorescence axis and bracts ferrugineous. Ovary appressed-pubescent. Fruit 8-11 by 5-8 mm, pilose. Infructescence 7-15 cm long. Stipules 4-10 mm long. |
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5a. |
Inflorescences at least partly cauline. |
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5b. |
Inflorescences axillary (sometimes terminally condensed). |
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6a. |
Petiole (at least of some leaves) longer than 10 mm, 1-1.5 mm wide, basally and distally usually slightly swollen and sometimes geniculate. |
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6b. |
Petiole shorter than 10 mm (if longer, then wider than 1.5 mm), basally and distally not swollen or geniculate. |
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7a. |
Tertiary veins close together, more than 10 between every two secondary veins (Fig. 1a). Petiole 1.5-4 mm wide. Leaves (7-)13-24(-50) by (2.5-)4-8(-15) cm. Calyx 0.3-0.5 mm long. Infructescence 10-25(-69) cm long. Fruiting pedicel 0.5-9 mm long. |
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7b. |
Tertiary veins widely spaced, less than 9 between every two secondary veins (Fig. 1b). Petiole up to 2 mm wide. Inflorescences 3-8 cm long. Leaves (5-)8-14(-19) by (2-)3-5(-7.5) cm. Calyx 0.7-1.2 mm long. Infructescences 3-15 cm long. Fruiting pedicel 0.5-2.5 mm long. |
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8a. |
Stipules, petiole and lower leaf side glabrous. Indumentum absent to whitish. Inflorescence 4-10 cm, infructescence 4-15 cm long. Sepals free or nearly so. Disc hairy (Fig. 5b, c). Fruit 6-8 by 4-6 mm, with a lateral to subterminal (Fig. 6c) (rarely terminal; Fig. 6a) style. |
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8b. |
Stipules, petiole and lower leaf side hairy all over. Indumentum ferrugineous. Inflorescence 2-4 cm, infructescence 3-8 cm long. Sepals halfway fused. Disc glabrous (sometimes hairs at base of calyx; Fig. 5a). Fruit 4-5 by 3 mm, with a terminal to slightly subterminal style (Fig. 6a). |
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9a. |
Stipules foliaceous, more than 4 mm wide. |
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9b. |
Stipules not foliaceous, up to 4 mm wide. |
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10a. |
Disc pilose (Fig. 5b, c). Calyx 0.7-1.2 by 1.2-1.5 mm. Stipules, petiole and leaves glabrous. |
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10b. |
Disc glabrous (Fig. 5a), rarely pilose (Fig. 5b, c). Calyx 0.3-0.8 by 0.5-0.8 mm. Stipules, petiole and major veins on lower leaf surface pilose. |
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11a. |
Leaves drying olive-green. Plant with whitish indumentum or glabrous, indumentum never ferrugineous. Inflorescence simple to much-branched, consisting of 1-13 racemes, 4-10 cm long. Sepals often glandular-fimbriate. Fruiting pedicel 1-4 mm long. Fruit 3-6(-8) by 2.5-4(-6) mm. |
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11b. |
Leaves drying reddish brown. Plant with ferrugineous indumentum, at least at the tip of the branchlets. Inflorescence simple, rarely branched once or in fascicles of two, 1.5-6 cm long. Sepals never glandular-fimbriate. Fruiting pedicel 1-20 mm long. Fruit 6-10 by 4-7 mm. |
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12a. |
Bracts longer than 1.5 mm, conspicuous even in very young inflorescences, less so in old infructescences. Fruit lens-shaped, obliquely ellipsoid or bean-shaped, 4-7 by 4-6 mm, with a distinctly asymmetrical base and lateral styles (Fig. 6c), laterally compressed (Fig. 6d). Sepals 5-8, free. Disc glabrous (Fig. 5a). Indumentum especially of young plant parts densely hirsute and never ferrugineous. Leaves abaxially hairy, drying olive-green. |
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12b. |
Bracts up to 1.5 mm long. Leaves, flower and fruit various. |
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13a. |
Sepals fused for more than 1/2 of their length. |
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13b. |
Sepals free to fused for up to 1/2 of their length. |
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14a. |
Fruit roughly bean-shaped, with a lateral style and an asymmetrical base (Fig. 6c), 5-8 by 4-6 mm. Leaves chartaceous to submembranaceous. |
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14b. |
Fruit mostly ellipsoid, with a terminal (Fig. 6a) to subterminal (sometimes lateral in A. leucocladon; Fig. 6c) style and a symmetrical base, size various. Leaves chartaceous to coriaceous. |
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15a. |
Fruit (3-) 4-6 (-8) by 3-4 (-6) mm, laterally compressed (Fig. 6d) or terete (Fig. 6b). Midvein impressed to flat adaxially (Fig. 3a, d). Leaves with or without domatia (Fig. 2), pilose at least along the major veins. |
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15b. |
Fruit 5-13 by 4-9 mm, laterally compressed (Fig. 6d). Midvein gently raised to flat adaxially (Fig. 3b, d). Leaves without domatia, completely glabrous or pubescent. |
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16a. |
Inflorescence 2-3 cm, infructescence 2-5(-8) cm long. Leaves with domatia between midvein and secondary veins abaxially (Fig. 2). Calyx pitcher-shaped, lobed for up to 1/3rd of its length, margin never glandular-fimbriate. |
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16b. |
Inflorescence 3-10 cm, infructescence 6-20 cm long. Leaf domatia present (Fig. 2) or absent. Calyx various. |
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17a. |
Calyx pitcher-shaped, lobed for 1/4th-1/2 of its length, margin never glandular-fimbriate. Leaves often with domatia (Fig. 2), ochraceous-pilose at least along the veins abaxially, higher venation finely tessellated. Fruit without or with inconspicuous white pustules when dry. |
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17b. |
Calyx cup-shaped, lobed for ½ of its length, margin often glandular-fimbriate. Leaves without domatia, usually glabrous except along the major veins, higher venation not conspicuously tessellated. Fruits usually with conspicuous white pustules when dry. |
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18a. |
Plant not rheophytic, leaves 2-3.5(-6.6) times longer than wide. |
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18b. |
Plant rheophytic, leaves 4-10 times longer than wide. |
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19a. |
Sepals fused for up to 1/2 of their length; calyx in flower and fruit bowl-shaped to shallowly cup-shaped. Fruit 10-13 by 7-11 mm. Infructescence 25-75 cm long. Flowers sessile, fruiting pedicel 0-1(-3) mm long. Stipules usually persistent, 6-25 mm long. Leaves (13-)25-30(-50) cm long. Petiole 1.5-6 mm wide. |
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19b. |
Sepals fused for 2/3rd-3/4th of their length; calyx in flower and fruit cup-shaped, pitcher-shaped or cylindrical. Fruit 5-11 by 4-7 mm. Infructescence 7-17 cm long. Flower shortly pedicellate, fruiting pedicel 1-9 mm long. Stipules early caducous, 2-6(-10) mm long. Leaves (5-)10-18(-32) cm long. Petiole 0.7-2 mm wide. |
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20a. |
Young twigs whitish. Infructescence 7-10 cm long. Fruiting pedicel 1-2.5 cm long. Disc hairy (Fig. 5b, c). Fruit with subterminal to lateral style (Fig. 6c, e). Calyx 0.5-0.8 mm long, inside at the base without or with whitish hairs. Petiole 0.7-1.5 mm wide. Stipules 0.5-1 mm wide. Leaves membranaceous to chartaceous. |
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20b. |
Young twigs brown. Infructescence 10-17 cm long. Fruiting pedicel 2-9 cm long. Disc glabrous (Fig. 5a). Fruit with terminal to slightly subterminal style (Fig. 6a). Calyx 1-1.5 mm long, inside at the base with long ferrugineous hairs. Petiole 1.5-2 mm wide. Stipules 1.5-2 mm wide. Leaves coriaceous. |
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21a. |
Indumentum lacking or whitish. Abaxial leaf surface glabrous except sometimes along the midvein. Ovary glabrous. Often cultivated. |
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21b. |
Indumentum ferrugineous. Abaxial leaf surface pubescent all over. Ovary sparsely pilose. Rare. |
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22a. |
Indumentum especially of young plant parts and inflorescences ferrugineous to ochraceous. |
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22b. |
Indumentum whitish or absent. |
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23a. |
Fruit 4-5 by 3 mm. Fruiting pedicel 0-1 mm long. Infructescence 3-8 cm long. Leaves up to 18 cm long. Stipules 3-7 by 0.5-1 mm. |
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23b. |
Fruit 6-20 by 4-9 mm. Fruiting pedicel 1-23 mm long. Infructescence 4-65 cm long. Leaves up to 50 cm long. Stipules 3-35 by 1-18 mm. |
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24a. |
Sepals free. Petiole 0.7-1 mm wide. Disc glabrous (Fig. 5a) to pubescent (Fig. 5b, c). Ovary ovoid to globose. Leaves drying olive-green, 1-2.25 times longer than wide, abaxially pubescent to nearly glabrous. |
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24b. |
Sepals fused for 1/2 of their length. Petiole 1.2-1.5 mm wide. Disc glabrous (Fig. 5a). Ovary almost cylindrical. Leaves drying reddish brown, 2-4 times longer than wide, abaxially ferrugineous-pubescent all over, distinctly velvety to the touch. |
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25a. |
Petiole 1 mm wide. Leaves (6-)9-14(-20) by up to 7 cm, tertiary veins weakly percurrent (Fig. 1b) to reticulate (Fig. 1c). Fruit ellipsoid, globose, ovate or obovate, compressed (Fig. 6d) or terete (Fig. 6b), glabrous, rarely thinly puberulent. Disc glabrous (Fig. 5a). Ovary globose, glabrous, rarely pilose. Inflorescence 1.5-6 cm long, infructescence 4-16 cm long. |
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25b. |
Petiole 2-5 mm wide. Leaves (8-)15-30(-55) by up to 25 cm, tertiary veins percurrent, close together (Fig. 1, see a, d resp.). Fruit obliquely ovate (mango-shaped) to falcate (Fig. 6e), always laterally compressed (Fig. 6d), hairy (old fruits sometimes almost glabrous). Disc glabrous (Fig. 5a) to pubescent (Fig. 5b, c). Ovary ovoid to falcate, densely appressed-pubescent. Inflorescence (5-)10-30 cm, infructescence 10-65 cm long. |
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26a. |
Fruit 10-13 by 7-9 mm, reticulate, laterally compressed (Fig. 6d). Sepals never glandular-fimbriate. Inflorescences simple, 8-35 cm, infructescences 25-75 cm long. Stipules 6-25 mm long. Leaves (13-)25-30(-50) cm long, midvein flat (Fig. 3d) to gently raised (Fig. 3b), rarely slightly impressed adaxially (Fig. 3a). Stipules, petiole and inflorescence axis usually glabrous. Fruiting pedicel 0-1(-3) mm long. |
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26b. |
Fruit 3-8 by 2.5-6 mm, areolate, compressed (Fig. 6d) or terete (Fig. 6b). Sepals often glandular-fimbriate. Inflorescences simple or branched, 2-10 cm long, infructescences 2.5-20 cm long. Stipules 1.5-7(-13) mm long. Leaves (1.5-)3-15(-30) cm long, midvein impressed adaxially (Fig. 3a). Stipules, petiole and inflorescence axis usually hairy. Fruiting pedicel 1-6 mm long. |
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27a. |
Fruit ventrally and dorsally compressed (Fig. 6f), 7-8 by 4-5 mm, pilose. Disc, ovary and abaxial leaf side (at least along the veins) densely hirsute. |
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27b. |
Fruit laterally compressed (Fig. 6d) or terete (Fig. 6b). Fruit size and indumentum various. |
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28a. |
Plant rheophytic, leaves 4-10 times longer than wide or narrower than 2 cm. Fruit terete (Fig. 6b). |
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28b. |
Plant not rheophytic, leaves 1-3.5(-6.6) times longer than wide. Fruit compressed (Fig. 6d) or terete (Fig. 6b). |
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29a. |
Average leaf size 3-6 by 0.4-1.2 cm. Leaf apex acute, sometimes rounded at the very tip. |
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29b. |
Average leaf size 5-11 by 1-2 cm. Leaf apex acuminate. |
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30a. |
Fruit laterally compressed (Fig. 6d). |
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30b. |
Fruit terete or hardly laterally compressed (Fig. 6b). |
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31a. |
Disc glabrous (Fig. 5a). Fruiting pedicel (2-)3-6 mm long. Fruit glabrous. |
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31b. |
Disc hairy (Fig. 5b, c) (if glabrous, fruiting pedicel 0-1 mm long and fruit hairy). Fruiting pedicel 0-2.5 mm long. Fruit pilose or glabrous. |
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32a. |
Fruit 6-8 by 4-6 mm, with a lateral to subterminal (Fig. 6c), rarely terminal style (Fig. 6a). Leaf base acute. Leaves drying greyish green to reddish brown, glabrous. Inflorescence 4-10 cm, infructescence 4-24 cm long, simple or branched once or twice. Sepals rounded to acuminate. Stipules, petiole and leaves glabrous. Young twigs light grey. |
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32b. |
Fruit 3-5 by 2.5-3(-3.5) mm, with a terminal to slightly subterminal style (Fig. 6a). Leaf base rounded to cordate, rarely acute. Leaves drying olive-green, pubescent at least along the major veins abaxially. Inflorescence 2-3 cm, infructescence 3-7 cm long, usually much-branched, consisting of (1-) 10-20 racemes. Sepals acute. Stipules, petiole and leaves usually pubescent. Young twigs brown. |
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33a. |
Calyx pitcher-shaped, lobed for Όth-1/2 of its length, margin never glandular-fimbriate. Leaves often with domatia (Fig. 2), ochraceous-pilose at least along the veins abaxially, higher venation finely tessellated, 3-12.5 by 1-4.2 cm. Petiole 0.7-1(-1.2) mm wide. Fruit without or with inconspicuous white pustules when dry. |
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33b. |
Calyx cup-shaped, lobed for c. ½ of its length, margin often glandular-fimbriate. Leaves without domatia, usually glabrous except along the major veins, higher venation not conspicuously tessellated, 6-30 by 2-12 cm. Petiole 1-2 mm wide. Fruits usually with conspicuous white pustules when dry. |
1. Antidesma acidum Retz., Obs. Bot. 5: 30. 1788, acida; Mόll.Arg. in DC., Prodr. 15, 2: 249. 1866; Airy Shaw, Kew Bull. 26: 352. 1971; Grierson & Long, Fl. Bhutan 786, fig. 48 h-j. 1987; Petra Hoffm. in Chayam. & Welzen, Fl. Thailand 8, 1: 59, fig. 14. 2005; Antidesma Malesia Thailand: 63, Map 3. (2006. Stilago diandra Roxb., Pl. Corom. 2: 35, pl. 166. 1802. Antidesma diandrum Roth, Nov. pl. sp.: 369. 1821. Antidesma lanceolarium (Roxb.) Wall., Cat.: 7284. 1832; Wight, Ic. Pl. Ind. Orient. 3(1): 4, t. 766. prob. 1844. Antidesma wallichianum Presl, Epim.: 235. 1849.
Shrub to tree, up to 10 m, diameter up to 10 cm, usually branched from the base. Young twigs pilose to pubescent, brown. Stipules usually persistent, linear, 3-7 by 1-2 mm, slightly pilose to (especially apically) densely pubescent. Petiole 2-7 by 1-2 mm, pilose to densely pubescent. Leaf blade obovate to elliptic-oblong, (2-)5-10(-18) by (1-)2.5-4 -8) cm, (1.6-)2-2.5(-4.5) times longer than wide, basally acute or obtuse, mostly cuneate, apically long to very shortly acuminate to rounded, chartaceous, glabrous or rarely slightly pilose adaxially, pilose to pubescent at least in the axils between midvein and secondary veins, rarely completely glabrous abaxially, midvein flat to impressed adaxially, tertiary veins reticulate, domatia present. Staminate inflorescences 5-14 cm long, axillary, simple or branched twice at the base, axis glabrous to pubescent. Bracts orbicular to lanceolate, 0.6-1 by 0.6 mm, glabrous. Staminate flowers 2-2.5 by 1-1.5 mm; pedicel 1-1.5 mm long, very thin; calyx 0.5 by 0.8 mm, globose to cup-shaped, sepals 4, fused for c. 2/3rd of their length, irregularly shaped, glabrous outside, pubescent inside with hairs often exceeding the calyx; disc cushion-shaped, enclosing the bases of the filaments, pubescent; stamens 2, rarely 3, 1.5-2 mm long, 1.5-2 mm long exserted from the calyx, anthers 0.3 by 0.6 mm; pistillode absent. Pistillate inflorescences 2-3 cm long, axillary, simple to branched twice at the base, axis glabrous to pubescent. Bracts orbicular to lanceolate, 0.5-0.7 by 0.5 mm, glabrous. Pistillate flowers 2 by 1 mm; pedicel 0.2-1 mm long; calyx 1 by 1 mm, pitcher-shaped, sepals 4 (or 5), fused for c. 2/3rd of their length, apically acute, glabrous outside, pubescent inside with hairs often exceeding the calyx; disc shorter than the sepals, glabrous inside, glabrous to pilose outside; ovary ovoid, glabrous, style usually terminal, stigmas 3 or 4. Infructescences 2-5(-8) cm long, pedicel 1.5-3 mm long; fruits ellipsoid, terete or laterally compressed, basally symmetrical, with a terminal to slightly subterminal style, 4-6 by 3-4 mm, glabrous.
T h a i l a n d. NORTHERN: Mae Hong Son, Chiang Mai, Chiang Rai, Nan, Lamphun, Lampang, Phrae, Tak, Phitsanulok, Kamphaeng Phet; NORTH-EASTERN: Phetchabun, Loei, Nakhon Panom; EASTERN: Buri Ram, Surin, Si Sa Ket, Ubon Ratchasima; SOUTH-WESTERN: Uthai Thani, Kanchanaburi, Ratchaburi; CENTRAL: Saraburi; SOUTH-EASTERN: Chon Buri, Chanthaburi; PENINSULAR: Ranong, Phangnga, Krabi, Satun.
D i s t r i b u t i o n. India (type) including Andaman & Nicobar Islands, Nepal, S China, Bangladesh, Burma, Laos, Vietnam, Cambodia, Thailand, Java). Absent in Malesia except Java; absent in Sri Lanka. Distribution map in Baker & al. (1998: 252, fig. 6D).
E c o l o g y. In dry deciduous, deciduous and evergreen forest, forest edges, open spaces, bamboo thickets; open or half-shady habitats; secondary, often disturbed, much degraded or frequently burnt vegetation. On sandy gravel, red volcanic soil, sand, silt or red lateritic soil, over limestone, granite and shale-granite. 0-1600 m altitude.
U s e s. Fruits eaten. Young leaves used in curry and as vegetable (India and Thailand).
V e r n a c u l a r. Mao ta khwai (เม่าตัาควาย) (Chiang Rai), makmao (มักเม่า) (Lampang); mao (เม่า) (Northern); mao soi (เม่าสร้อย) (Loei); hao cha (Karen).
2. Antidesma bunius (L.) Spreng., Syst. Veg. 1: 826. 1824; Wight, Ic. Pl. Ind. Orient. 3(1): t. 819; Petra Hoffm. in Chayam. & Welzen, Fl. Thailand 8, 1: 61, fig. 15. 2005; Antidesma Malesia Thailand: 74, Fig. 7; Map 6. 2006. Stilago bunius L., Mant. Pl. 1: 122. 1767.
Tree (to shrub), up to 30 m, diameter up to 1 m, often fluted or with buttresses. Young twigs glabrous to densely ferrugineous-pubescent, brown. Stipules early caducous, linear, 4-6 by 1.5-2 mm, pubescent. Petiole 3-10(-17) by 1.5-2 mm, glabrous to densely ferrugineous-pubescent. Leaf blade oblong to elliptic, more rarely obovate, (5-)10-18(-32) by (2-)4-6(-9) cm, (2-)2.5-3(-6) times longer than wide, coriaceous, basally acute to rounded, rarely cordate, apically acuminate, acumen apically often rounded, glabrous, or pilose to ferrugineous-pubescent only along the midvein adaxially, glabrous to ferrugineous-pubescent all over abaxially, midvein flat adaxially, tertiary veins reticulate, drying olive-green adaxially, lighter and more reddish abaxially, domatia absent. Staminate inflorescences (6-)15-25 cm long, axillary, branched, consisting of 3-8(-14) racemes, axis pubescent to glabrous. Bracts deltoid to lanceolate, 0.5 by 0.5 mm, pubescent. Staminate flowers 3-4 by 3 mm, sessile; calyx 1 by 1.5 mm, cup-shaped, sepals 3 or 4(5), fused for 2/3rd-3/4th of their length, apically obtuse, glabrous to ferrugineous-pubescent on both sides; disc variable, extrastaminally-annular to lobed, lobes free or enclosing the bases of the stamens and pistillode, much shorter to extending to the same length as the sepals, glabrous; stamens 3 or 4(5), 2-3 mm long, 1.5-2 mm long exserted from the calyx, anthers 0.5 by 0.7; pistillode clavate to cylindrical, 1 by 0.5 mm. Pistillate inflorescences (4-) 8-17 cm long, axillary, simple or more rarely branched, consisting of up to 4 racemes, axis glabrous to pubescent. Bracts deltoid, 0.5-1.5 by 0.5-1 mm, pubescent to pilose. Pistillate flowers 2.5-3 by 1.5 mm; pedicel 0.5-1(-2) mm long; calyx 1-1.5 by 1.5 mm, cup-shaped, sepals 3, fused for 2/3rd-3/4th of their length, apically obtuse to rounded, glabrous to pilose on both sides; disc shorter than the sepals, glabrous; ovary ellipsoid, glabrous to very sparsely pilose, style terminal to subterminal, stigmas 3 or 4(6). Infructescences 10-17 cm long, robust; pedicel 2-4(-9) mm long; fruits ellipsoid, laterally compressed, 5-11 by 4-7 mm, basally symmetrical, with a terminal to slightly subterminal style, glabrous, rarely pilose.
2a. var. bunius: Petra Hoffm. in Chayam. & Welzen, Fl. Thailand 8, 1: 63. 2005; Antidesma Malesia Thailand: 75, Map 6. 2006. Antidesma sylvestre Lam., Enc. 1: 207. 1783, sylvestris. Antidesma bunius (L.) Spreng. var. wallichii Mόll.Arg. in DC., Prodr. 15, 2: 263. 1866. Antidesma floribundum Tul., Ann. Sci. Nat., Bot., Sιr. 3: 189. 1851. Antidesma ciliatum C.Presl, Epimel.: 234. 1849. Antidesma cordifolium C.Presl, Epim.: 235. 1849. Antidesma andamanicum Hook.f., Fl. Br. India 5: 364. 1887. Sapium crassifolium Elmer, Leafl. Philipp. Bot. 2: 485. 1908. Antidesma collettii Craib, Bull. Misc. Inform. Kew 1911(10): 461. 1911. Antidesma thorelianum Gagnep., Bull. Soc. Bot. France 70: 124. 1923.
Young twigs glabrous to very shortly pubescent. Petiole glabrous to pubescent (especially adaxially). Leaf blade glabrous, or pilose only along the midvein, often minutely white pustulate. Inflorescence axis glabrous to pubescent. Staminate calyx glabrous to pubescent outside, ferrugineous-pubescent at the base inside, hairs often exceeding the calyx. Pistillate calyx glabrous to sparsely pilose outside, ferrugineous-pubescent at the base inside, hairs often exceeding the calyx; ovary glabrous.
T h a i l a n d. NORTHERN: Chiang Mai, Chiang Rai, Lamphun, Lampang, Tak; NORTH-EASTERN: Loei, Nakhon Phanom; SOUTH-WESTERN: Kanchanaburi; SOUTH-EASTERN: Chon Buri, Trat.
D i s t r i b u t i o n. India (type) incl. Andaman & Nicobar Islands, Sri Lanka, S China, Burma, Laos, Vietnam, Thailand, Singapore, Borneo, Java, Philippines, Sulawesi, Lesser Sunda Islands, Moluccas, New Guinea, Christmas Islands (Indian Ocean), Tahiti, Hawaiian Islands. As A. bunius is widely cultivated as a fruit tree, it is impossible to distinguish truly wild occurrences.
E c o l o g y. In wet evergreen forests, open forests, dipterocarp forest, thickets, bamboo thickets, teak forest, river banks, forest edges, by roadsides, semi-cultivated and cultivated areas, gardens, shady or open habitats, usually in secondary but also in primary vegetation. On sand, loam or clay over (coral) limestone or granite bedrock. 0-2100 m altitude.
U s e s. Widely cultivated as a fruit tree. Wood subject to termite attacks and therefore unsuitable for work in contact with the soil (Vietnam, Poilane 10901). Leaf eaten as vegetable (Java).
V e r n a c u l a r. Ba mao ruesi (บ่าเม่าฤาษี), mamao dong (ขะเม่าดง) (Chiang Mai); maeng mao khwai (แมงเม่าควาย), mao chang (เม่าช้าง) (Chanthaburi).
2b. var. pubescens Petra Hoffm., Kew Bull. 54: 350. 1999; in Chayam. & Welzen, Fl. Thailand 8, 1: 64. 2005.
Young twigs, petiole and inflorescence axis densely ferrugineous-pubescent. Leaf blade glabrous, or ferrugineous-pubescent only along the midvein adaxially, ferrugineous-pubescent all over abaxially, especially along the veins. Staminate calyx densely ferrugineous-pubescent on both sides. Pistillate calyx pilose on both sides; ovary very sparsely pilose. Infructescences and fruits not seen.
T h a i l a n d. NORTHERN: Lamphun (type: Me Lee, winit 295, holotype K!); NORTH-EASTERN: Phetchabun (Lom Khao).
D i s t r i b u t i o n. Endemic to Thailand.
E c o l o g y. In evergreen forest by stream. 700-1020 m altitude.
V e r n a c u l a r. Mamao petra (ขะเม่าเปตรา) (General).
N o t e. These specimens have been referred to by Airy Shaw (1971: 359) as A. cf. nienkui Merr. & Chun. Apart from the considerable disjunction, A. bunius var. pubescens differs from the type and only specimen of A. nienkui from Hainan in the shorter petioles and fused calyx lobes. Both taxa, however, have a similar dense ferrugineous indumentum in most parts, which distinguishes them from the nearly glabrous A. bunius var. bunius.
3. Antidesma forbesii Pax & K.Hoffm. in Engl., Pflanzenr. 81: 153. 1922; Petra Hoffm. in Chayam. & Welzen, Fl. Thailand 8, 1: 64. 2005; Antidesma Malesia Thailand: 122, Map 15. 2006.Antidesma plagiorrhynchum Airy Shaw, Kew Bull. 28: 272. 1973; Kew Bull. 36: 356, fig. C1-2, 363. 1981. Antidesma pradoshii Chakrab. & Gangop., J. Econ. Taxon. Bot. 21: 479. 1997.
Treelet, up to 8(-12.5) m, diameter 8(-15.6) cm. Young twigs glabrous to puberulent, first dark brown with conspicuous lenticels, then light to whitish grey. Stipules early caducous, deltoid, 1 by 0.5 mm, glabrous. Petiole basally and distally often slightly swollen and geniculate, (4-)6-17 by 0.5-1 mm, pilose to puberulent. Leaf blade ovate to elliptic, (4.5-)6-11(-16) by (1.5-)2-4(-5.5) cm, 2.3-4.1 times longer than wide, chartaceous to submembranaceous, basally acute to obtuse, apically acuminate, acumen apically rounded, glabrous or pilose only along the midvein adaxially and/or abaxially, midvein impressed adaxially, tertiary veins reticulate, drying olive-green, lighter abaxially. Staminate inflorescences 2-4 cm long, axillary, simple or branched, consisting of up to 4 racemes, slender, axis pubescent. Bracts elliptic, 0.4-0.5 by 0.4 mm, pilose. Staminate flowers 1-1.5 by 1-2 mm, sessile; calyx 0.5 by 0.8-1 mm, bowl-shaped, sepals 5, fused for 2/3rd of their length, apically rounded, shortly pilose outside, glabrous inside but long hairs at the base; disc consisting of 4 or 5 free lobes, lobes ± obconical, with two shallow imprints apically, 0.2 by 0.3 mm, as long as the sepals, glabrous; stamens 4-5, 1-1.5 mm long, 0.5 mm long exserted from the calyx, anthers 0.2 by 0.3 mm; pistillode cylindrical, 0.8 by 0.3 mm, pubescent. Pistillate inflorescences 2-5 cm long, axillary, simple, sometimes in fascicles of 2 racemes, axis shortly pubescent. Bracts elliptic, 0.5 by 0.5 mm, pilose. Pistillate flowers 1.5 by 1.5 mm; pedicel 0.7-1.2 mm long; calyx 0.7 by 1 mm, cup-shaped, sepals 4 or 5, fused for 1/2 or a little more of their length, apically rounded, pilose outside, glabrous inside but with long hairs at the base; disc shorter than the sepals, glabrous; ovary subglobose, laterally compressed, glabrous to sparsely pilose, style lateral, thick, stigmas 3-6. Infructescences 5-11 cm long; pedicel 2-5 mm long; fruits roughly bean-shaped, with a ventral bulge, laterally compressed, 5-8 by 4-6 mm, basally asymmetrical, with a lateral style, glabrous to very sparsely pilose.
T h a i l a n d. PENINSULAR: Pangnga.
D i s t r i b u t i o n. Thailand (one collection), Peninsular Malaysia (Pahang, Cameron Highlands only), Sumatra (type).
E c o l o g y. In montane rain and mossy forest. 1000-1900 m altitude.
V e r n a c u l a r. Mamao fop (ขะเม่าฝอบ) (Phangnga).
4. Antidesma ghaesembilla Gaertn., Fruct. 1: 189, t. 39. 1788; Wight, Ic. Pl. Ind. Orient. 3(1): t. 820. 1844-45; Gagnep. & Beille in Lecomte, Fl. Indo-Chine 5: 505, fig. 64.11-19. 1926; Petra Hoffm. in Chayam. & Welzen, Fl. Thailand 8, 1: 65. 2005; Antidesma Malesia Thailand: 124, Fig. 13e-j, Map 16 (as map 15 on p. 126). 2006. Antidesma pubescens Roxb., Pl. Corom. 2: 35, pl. 167. 1802. Antidesma paniculatum Willd., Sp. pl. 4.2.: 764. 1806, paniculata. Antidesma vestitum Presl, Epim.: 232. 1849.
Shrub, treelet or tree, up to 20 m, diameter up to 32 cm. Twigs greenish, brown, grey, young twigs sometimes pinkish, pubescent, brown. Stipules early caducous, subulate, 3-6 by 0.5-1 mm, pubescent. Petiole 4-10(-17) by 0.7-1 mm, pubescent, glabrescent. Leaf blade oblong, rarely slightly ovate or obovate, (2-)4-7(-16) by (2-)3-4.5(-9) cm, (1-)1.3-1.7(-2.25) times longer than wide, chartaceous to coriaceous, basally rounded to cordate, very rarely acute, apically rounded (to obtuse, very shortly acuminate, truncate or retuse), pubescent to glabrous, often only the major veins of both sides and the margin pubescent, midvein flat adaxially, tertiary veins reticulate to weakly percurrent, drying olive-green. Staminate inflorescences 4-8 cm long, axillary, much-branched, consisting of 10-20 racemes, axis ferrugineous-pubescent. Bracts lanceolate, 0.5-1 by 0.5 mm, pubescent. Staminate flowers 2-3 by 2-3 mm, sessile; calyx 0.5-1 by 1.5 mm, sepals 4-6, free, deltoid to oblong, apically acute to obtuse, pubescent outside, more or less glabrous inside; disc consisting of 4-6 free lobes, lobes ± obconical, well-separated from each other, 0.5 by 0.5 mm, pubescent; stamens 4-6, 2 mm long, 1-1.5 mm long exserted from the calyx, anthers 0.5 by 0.5 mm; pistillode obconical, 0.5 by 0.2-0.3, pubescent. Pistillate inflorescences 2-3 cm long, axillary, usually much-branched, consisting of (1-)10-20 racemes, axis ferrugineous-pubescent. Bracts lanceolate, 0.8 by 0.4 mm, pubescent. Pistillate flowers 1.5-2 by 1-1.5 mm; pedicel 0-1 mm long; calyx 0.7 by 1-1.5 mm, sepals 5 or 6, free, deltoid, apically acute, pubescent outside, glabrous inside; disc shorter than the sepals, glabrous to pubescent especially at the margin; ovary ovoid to globose, pubescent, style usually subterminal, glabrous, stigmas (2)3(5). Infructescences 4-7 cm long; pedicel 0-1 mm long; fruits lens-shaped, more rarely ellipsoid, laterally compressed, 3-4(-5) by 2.5-3(-3.5) mm, basally symmetrical, with a terminal, more rarely subterminal style, pilose.
T h a i l a n d. NORTHERN: Mae Hong Son, Chiang Mai, Chiang Rai, Lamphun, Lampang, Tak, Phitsanulok, Nakhon Sawan; NORTH-EASTERN: Phetchabun, Loei, Khon Kaen; EASTERN: Nakhon Ratchasima, Buri Ram, Si Sa Ket; SOUTH-WESTERN: Kanchanaburi, Ratchaburi, Phetchaburi?, Prachuap Khiri Khan; CENTRAL: Ang Thong, Bangkok; SOUTH-EASTERN: Chon Buri, Chanthaburi; PENINSULAR: Chumphon, Surat Thani, Pangnga, Nakhon Si Thammarat, Trang, Songkhla, Yala, Narathiwat.
D i s t r i b u t i o n. India incl. Nicobar Islands, Sri Lanka, S China, Bangladesh, Burma, Laos, Vietnam, Cambodia, Thailand, Peninsular Malaysia, Sumatra, Borneo (Kalimantan, Sabah), Java, Philippines, Sulawesi, Lesser Sunda Islands, Moluccas, New Guinea, N Australia. Type without locality.
E c o l o g y. In savannah, grassland (often regularly burnt, said to be fire resistant, NGF 9809), open forest, dense scrubby forest, vine thickets, fresh-water swamps, edge of mangrove, coastal fringes, secondary vegetation around towns and villages (Malay: 'belukar'); on roadsides and river banks, on dry to swampy ground; usually in secondary vegetation. On clay, sand, lateritic soil, black peat, limestone, sometime on ultrabasic, over shale and granite bedrock. 0-1250 m altitude.
U s e s. Fruits eaten locally.
V e r n a c u l a r. Khamao pha (ขะเม่าผา), mamao (มะเม่า) (Northeastern); mao khai pla (เม่าไข่ปลา) (Chon Buri); mang mao (มังเม่า) (Chanthaburi); ma mao khao bao (มะเม่าข้าวเบา) (Chumphon), mao thung (เม่าทุ่ง) (Chumphon, Songkhla); ku-chae (กูแจ) (Malay-Narathiwat).
5. Antidesma helferi Hook.f., Fl. Br. India 5: 357. 1887; Petra Hoffm. in Chayam. & Welzen, Fl. Thailand 8, 1: 66. 2005; Antidesma Malesia Thailand: 129, Map 17. 2006. Antidesma pachystemon Airy Shaw, Kew Bull. 23: 279. 1969.
Shrub or tree, up to 20 cm, diameter up to 15 cm. Young twigs glabrous or nearly so, light grey. Stipules caducous or persistent, rarely foliaceous (Geesink & Santisuk 5166), linear to oblanceolate, 3-8(-20) by 0.5-1(-5) mm, apically acute, glabrous. Petiole sometimes sharply keeled abaxially, 3-10 by (0.7-)1-2 mm, glabrous. Leaf blade elliptic, oblong or ovate, (5-)9-14(-19) by (2-)3.5-5.5(-7.5) cm, (1.9-)2.6(-4.3) times longer than wide, chartaceous to coriaceous, basally acute, sometimes decurrent, apically acute or acuminate, glabrous, midvein flat to shallowly impressed adaxially, tertiary veins reticulate, widely spaced (4-7 between every two secondary veins), drying greyish green to reddish brown, domatia absent. Staminate inflorescences 3-8 cm long, axillary to almost cauline, simple or branched once at the base, axis pilose to glabrous. Bracts deltoid to linear, 0.7-1.2 by 0.3-0.5 mm, pilose or glabrous. Staminate flowers 2-3 by 1.5-2 mm; pedicel 0-1 mm long; calyx 0.7-1 by 1.2-1.5 mm, sepals 4, fused for 1/3rd of their length, 0.7-1 mm wide, apically rounded, pilose to glabrous outside, glabrous inside; disc cushion-shaped, enclosing the bases of the filaments and pistillode, pilose; stamens 4, 2-2.5 mm long, 1.5-2 mm long exserted from the calyx, anthers 0.5-0.7 by 0.5-1 mm; pistillode conical or globose, 0.1-0.3 by 0.1-0.3 mm, glabrous. Pistillate inflorescences 4-10 cm long, terminal, axillary or almost cauline, simple, rarely branched once or twice at the base, sometimes in fascicles of 2 racemes, axis patulously pubescent to glabrous. Bracts narrowly deltoid to linear, 0.5-1(-1.5) by 0.3-0.4 mm, pubescent to glabrous. Pistillate flowers 1.5-3 by 1.5-2 mm; pedicel 0-1 mm long; calyx 0.7-1.2 by 1.2-1.5 mm, sepals 4 or 5(6), free or nearly so, 0.5-0.8 mm wide, orbicular to almost linear, apically rounded to acuminate, pilose to glabrous outside, glabrous inside; disc shorter or as long as the sepals, setulous outside and at the margin; ovary ellipsoid to globose, sparsely to densely setulous, style subterminal to lateral, stigmas (2)3 or 4. Infructescences 4-15 cm long; pedicel 0.5-2.5 mm long; fruits ellipsoid to lens-shaped, laterally compressed, 6-8 by 4-6 mm, basally symmetrical or asymmetrical, with a subterminal to lateral, rarely terminal style, glabrous to setulous.
T h a i l a n d. PENINSULAR: Ranong, Pangnga, Krabi, Nakhon Si Thammarat, Trang.
D i s t r i b u t i o n. Peninsular Burma (type), Thailand, Peninsular Malaysia, Sumatra (Aceh and East coast).
E c o l o g y. In primary evergreen or mossy forest, often near water or on marshy ground. On alluvial and greyish clay-mud soil. 100-1300 m altitude.
N o t e s. The specimens from Peninsular Malaysia and from Sumatra have bigger, more elliptic, thinner and dull leaves than those from Burma and Thailand, whereas the generative characters are uniform throughout the distribution area.
The fruiting specimens Kerr 18703 from Krabi, and Larsen & al. 45988 from Khao Luang, Nakhon Si Thammarat, differ by their glabrous discs from the remainder of the collections. In this and all vegetative characters the latter specimen much resembles the unmatched collection van Beusekom & Phengkhlai 967 from the same locality and altitude.
The stipules of Geesink & Santisuk 5166 are exceptional in that they are leaf-like.
6. Antidesma japonicum Siebold & Zucc., Abh. Math.-Phys. Cl. Kφnigl. Bayer. Akad. Wiss. 4(3): 212. 1846 (repr. as Fl. jap. fam. nat. 2: 88. 1846); Petra Hoffm. in Chayam. & Welzen, Fl. Thailand 8, 1: 66. 2005; Antidesma Malesia Thailand: 134, Map 18. 2006.
Shrub to tree, up to 5(-8) m, diameter up to 10 cm. Young twigs pubescent to pilose, brown. Stipules usually caducous, narrowly deltoid to subulate, 2-5 by 0.7-1 mm, puberulent to pilose. Petiole basally and distally sometimes slightly swollen, 2-8 by 0.5-1.5 mm, pubescent to almost glabrous. Leaf blade narrowly elliptic-oblong to ovate or obovate, (3.5-)6-10(-13) by (1-)2-3.5(-4.7) cm, (1.2-)3(-4.4) times longer than wide, subcoriaceous to chartaceous, basally acute to rounded, apically acute to acuminate, sometimes caudate, glabrous, sometimes some hairs only along the midvein, midvein impressed to flat or distinctly raised adaxially, tertiary veins weakly percurrent in wider, reticulate in narrower leaves, widely spaced, drying light olive- to greyish green. Staminate inflorescences 2-6 cm long, axillary, sometimes terminally condensed, simple or branched, consisting of up to 5 racemes, slender, axis pilose. Bracts deltoid to lanceolate, 0.5-0.7(-1) by 0.3-0.5 mm, glabrous to pilose, margin often glandular-fimbriate. Staminate flowers 1-1.5 by 1-1.5 mm; pedicel (0.5-)1-1.5 mm long; calyx 0.4-0.6 mm long, sepals 3-5, free, deltoid to lanceolate, apically acute to rounded, glabrous on both sides; disc cushion-shaped, fully or partially enclosing the bases of the filaments and pistillode, glabrous; stamens 3-5, 1-2 mm long, fully exserted from the spreading sepals, anthers 0.3-0.5 by 0.3-0.5 mm; pistillode absent, flat, clavate, cylindrical or 3-fid, 0-0.5 by 0-0.2 mm, glabrous. Pistillate inflorescences 2-5 cm long, axillary to terminal, simple or branched once, rarely twice, axis nearly glabrous to puberulent. Bracts deltoid to linear, 0.3-1 by 0.2-0.5 mm, pubescent to almost glabrous, margin sometimes glandular-fimbriate. Pistillate flowers 1-1.5 by 1 mm; pedicel (0.5-)1-1.5 mm long; calyx 0.5-0.7 by 0.7 mm, sepals 3-5(7), free, 0.3-0.5 mm wide, deltoid, apically acute to rounded, glabrous but sometimes long hairs at the base inside; disc much shorter to as long as the sepals, glabrous; ovary fusiform to ellipsoid, glabrous, style terminal, stigmas 3-5. Infructescences 4-9 cm long; pedicel (0-)3-6 mm long; fruits ellipsoid to lens-shaped, laterally compressed, 5-6(-8) by 4-5(-6) mm, basally asymmetrical to slightly symmetrical, with a subterminal, rarely nearly terminal style, glabrous, rarely white pustulate.
