Malesian Euphorbiaceae Descriptions




P.C. van Welzen & L.J. Bulalacao


P.C. van Welzen & L.J. Bulalacao. 2007. The genus Alchornea (Euphorbiaceae) in the Malay Archipelago and Thailand. Syst. Bot. 32: 803—818.


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Genus description

Key to the species

Species descriptions

Excluded species


Alchornea Sw.


    Alchornea Sw., Prodr. 6 (1788) 98; Mόll.Arg. in DC., Prodr. 15, 2 (1866) 899; Hook.f., Fl. Br. India 5 (1887) 420; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 220; Backer & Bakh.f., Fl. Java 1 (1963) 485; Airy Shaw, Kew Bull. 26 (1972) 211; Whitmore, Tree Fl. Malaya 2 (1973) 53; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 28; Kew Bull. 35 (1980) 593; Kew Bull. Add. Ser. 8 (1980) 25; Kew Bull. 36 (1981) 249; Kew Bull. 37 (1982) 4; Alphab. Enum. Euphorb. Philipp. Isl. (1983) 3; G.L.Webster, Ann. Missouri Bot. Gard. 81 (1994) 81; Radcl.-Sm., Gen. Euphorbiacearum (2001) 192; Secco, Fl. Neotr. 93 (2004) 55; Welzen & Bulalacao, Syst. Bot. 32 (2007) 804; G.L.Webster in Kubitzki, Fam. & Gen. Vasc. Pl. 11 (2014) 112, Fig. 24. — Type: Alchornea latifolia Sw.

    Cladodes Lour., Fl. Cochinch. (1790) 574. — Type: Cladodes rugosa Lour. [= Alchornea rugosa (Lour.) Mόll.Arg.]

    Hermesia Humb. & Bonpl. ex Willd., Sp. Pl. 4 (2) (1805) 809. — Type: Hermesia castanifolia Humb. & Bonpl. ex Willd. [= Alchornea castanifolia (Humb. & Bonpl. ex Willd.) A.Juss.]

    Schousboea Schumach., Beskr. Guin. Pl. (1827) 449 (non Willd., Sp. Pl. 2: 579. 1799, Combretaceae). — Type: Schousboea cordifolia Schumach. [= Alchornea cordifolia (Schumach.) Mόll.Arg.]

    Stipellaria Benth., Hooker’s J. Bot. Kew Gard. Misc. 6 (1854) 2. — Lectotype (designated by Thin, Tap Chi Sinh Hoc 6 (1984) 26-29): Stipellaria trewioides Benth. [= Alchornea trewioides (Benth.) Mόll.Arg.]

    Lepidoturus Bojer ex Baill., Ιtude Euphorb. (1858) 448. — Type: Lepidoturus alnifolius Bojer ex Baill. [= Alchornea alnifolia (Bojer ex Baill.) Pax & K.Hoffm.]

    Bleekeria Miq., Fl. Ned. Ind. 1 (2) (1859) 407 (non Hassk., Retzia 1: 38. 1855, Apocynaceae). — Type: Bleekeria zollingeri (Hassk.) Miq. [= Alchornea tiliifolia (Benth.) Mόll.Arg.]


Shrubs to small trees, monoecious or dioecious; branching sympodial. Indumentum in Malesia simple, usually hairs of two lengths. Stipules 2, linear to needle-like, hairy outside, late caducous, usually 1 to several papillae-like glands basally along the margin. Leaves alternate, simple; petiole short to long, round or reniform, somewhat pulvinate basally and even less so apically; stipellae absent (blades obovate) or 2 (blades ovate), needle-like and erect to somewhat patent or leaf-like and pendulous to patent, often with papillae-like glands basally; blade ovate (to elliptic) to obovate, base emarginate to cuneate, margin serrate with upright glandular teeth, apex acute to caudate, upper surface glabrous to pubescent on venation, extrafloral nectaries absent or round ones present; lower surface generally more hairy, basally and usually apically with extrafloral nectaries (usually absent when nectaries present on upper surface), hair-tuft domatia often present; venation pinnate or triplinerved, nerves ending open in the margin or curved and closed, veins laxly scalariform, veinlets reticulate. Inflorescences ramiflorous (staminate) to terminal (pistillate), usually racemes, sometimes branched panicles, usually with one sex (A. liukiuensis excepted), exceptionally mixed sexes; bracts triangular to ovate, caducous, hairy outside, glabrous inside, margin often with papillae-like glands at base, often below bracts or on base of bracts round extrafloral nectaries; bracteoles present, very small on staminate axes. Flowers symmetric, sessile or pedicelled; pedicels with abscission zone and often with raised, round extrafloral nectaries when pistillate; sepals glabrous to hairy outside, glabrous inside; petals and disc absent. Staminate flower in groups of 3—25 per node; calyx valvate, usually 2-4-partite, lobes ovate, apiculate; stamens 4-8, inserted on a ring-like collar of receptacle (not excreting), filaments strap-like, anthers oblong, parallel, dorsifixed, 2-locular, opening latrorsely with longitudinal slits; pistillode absent. Pistillate flowers solitary per node, pedicellate or sessile; sepals usually 5, imbricate, margin often with papillae-like glands, outside hairy, inside glabrous; ovary 3(4)-locular, 1 ovule per locule; style absent to long; stigmas long triangular, margins reflexed, lowly papillate above, slightly hairy beneath, apex not split (to rarely split or somewhat frayed). Fruits slightly or deeply 3-lobed capsules, dehiscing septicidally and/or partly loculicidally, hairy or glabrous outside, glabrescent, glabrous inside, wall smooth or warty, exocarp often dehiscing from mesocarp; columella narrow, apex not or only slightly T-shaped. Seeds ellipsoid, somewhat ridged on the inside, smooth or verrucate, apically usually with a dented hilum, brown or black.

    Distribution — A pantropical genus with about 40 species (Secco, 2004).


Key to the species



Leaves (elliptic to) obovate; venation penninerved; stipellae absent

1. Alchornea rugosa


Leaves ovate; venation triplinerved; stipellae or glands present at insertion of blade



Inflorescences up to 7 cm long. Style absent. Staminate bracts 1.4—2 by 1.3—2.4 mm, pistillate ones 1.3—2.4 mm wide, stiff, midrib usually raised abaxially, often striate because of parallel venation. Leaf blades 1.5—11 by 1.2—7 cm; lower surface with many round extrafloral nectaries

4. Alchornea sicca


Inflorescences up to 21 cm long. Style 0.4—6 mm long. Staminate bracts either 0.5—1.2 by 0.3—1 mm or 0.8—3 by 0.8—2 mm, pistillate ones generally smaller, 0.3—1.3(—3.2) mm wide, papery, venation not visible. Leaf blades 4.5—27 by 2—20 cm; lower surface with apically none to many round extrafloral nectaries



Lower leaf surface with basal extrafloral nectaries only, apical ones absent. Ovaries and fruits smooth (to somewhat muricate). — Malaysia: Penang Island

2. Alchornea rhodophylla


Lower leaf surface with basal and apical extrafloral nectaries. Ovaries and fruits especially on upper part muricate (to almost smooth when ripe). — West Malesia, absent from Penang Island



Stipellae 1.2—2.5 by 0.3—0.4 mm, subulate, upright or patent. Lower leaf surface slightly hairy on venation. Staminate bracts 0.6—0.7 by 0.4—0.7 mm. Pistillate bracts 0.8—4 mm high, bracteoles 0.5—2.3 mm high. Styles 0.5—1.3 mm long

1. Alchornea parviflora


Stipellae 3—18 by 0.6—7 mm, ovate and leaf-like (to subulate or gland-like; sometimes additional subulate or gland-like stipellae present under normal ones), hanging to patent. Lower leaf surface subglabrous to generally hairy all over. Staminate bracts 0.8—3 by 0.8—2 mm. Pistillate bracts 1.5—7.7 mm high, bracteoles 1—5.5 mm high. Styles 0.4—6 mm long

5. Alchornea tiliifolia


1. Alchornea parviflora (Benth.) Mόll.Arg.


    Alchornea parviflora (Benth.) Mόll.Arg., Linnaea 34 (1865) 168; in DC., Prodr. 15, 2 (1866) 902; Merr., Enum. Philipp. Fl. Pl. 2 (1923) 438; Airy Shaw, Alphab. Enum. Euphorb. Philipp. Isl. (1983) 3. 1983; Welzen & Bulalacao, Syst. Bot. 32 (2007) 807, fig. 1b, c, f; 2i-k, 3. — Stipellaria parviflora Benth., Hooker’s J. Bot. Kew Gard. Misc. 6 (1854) 4. — Type: Cuming 1800 (holo K; iso L), Philippines, Island of Negros.

    ? Alchornea adenophylla Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 251; Ridl., Fl. Malay Penins. 3 (1924) 278; Whitmore, Tree Fl. Malaya 2 (1973) 54; Airy Shaw, Kew Bull. 36 (1981) 251. — Type: Scortechini s.n. (n.v., not traceable), Malaysia, Perak. See note 1.

    Alchornea borneensis Pax & K.Hoffm., Mitt. Inst. Allg. Bot. Hamburg 7 (1931) 227; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 28. — Type: Hans Winkler 353 (holo HBG; iso HBG), Indonesia, West Borneo, bei Lebang Hara.

    Acalypha tiliifolia auct. non (Benth.) Vidal: Vidal, Revis. Pl. Vasc. Filip (1886) 244, p.p., pro Vidal y Soler 592 (K) (see note 2); Ridl., Fl. Malay Penins. 3 (1924) 277.

    Alchornea sicca auct. non (Blanco) Merr.: Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 251.

    Alchornea trewioides auct. non (Benth.) Mόll.Arg.: Airy Shaw, Kew Bull. 26 (1972) 212.


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Shrubs to small trees, up to 9 m high, bole up to 4 m high, d.b.h. up to 38 cm; branchlets somewhat pilose when young, glabrescent. Outer bark thin, very finely roughened, with sparse lenticels, whitish to red-brown; inner bark pale greenish to brownish; sapwood whitish to yellowish. Stipules 2.3—5 by 0.3—1 mm. Leaves: petiole 0.8—10.2 cm long, round to reniform in transverse section, basally and less so apically indistinctly (constricted) pulvinate, pilose, brown; stipellae needle-like, 1.2—2.5 by 0.3—0.4 mm, subulate but flattened towards blade, upright or patent, hairy, light green, with sometimes reddish tips, marginal basal gland present; blade ovate, 7.5—18 by 3.1—11 cm, length/width ratio 1.6—3, broadest at lower third, base slightly emarginate to rounded, margin laxly serrate with upwardly bent glandular teeth, apex cuspidate to caudate, often mucronate, upper surface hairy on nerves, extrafloral nectaries absent, dark green; lower surface slightly hairy on venation, more densely so on nerves, glaucous to grey-light green, basally two (outside nerves) or four (between nerves) round slightly raised, light brown extrafloral nectaries, in nerve axil or slightly distant from axil, additional ones apically and usually along the margin, often slightly pocket-like domatia or slight hair tuft domatia in nerve axils; venation triplinerved, flat adaxially, raised abaxially, nerves 5—9. Inflorescences ramiflorous, axillary and terminal racemes (pistillate and staminate) or seldom panicles (staminate), single or several together (staminate), up to 10(—21) cm long, shortly pilose, light green; bracts triangular, hairy outside, staminate ones small, 0.6—0.7 by 0.4—0.7 mm, light brown, pistillate ones 0.8—4 by 0.3—1 mm, papery, venation not visible, basally often with glandular teeth or a large marginal gland, light green; bracteoles several, on node in bract or along pedicel, triangular, 0.5—2.3 by 0.4—1 mm, hairy outside. Staminate flowers only seen in bud, several per node, pedicelled, pedicel with abscission zone, sepals 3 or 4, basally united, ovate, ca 4.5 by 0.8 mm, hairy outside, bright light green; stamens 6 or 8, filaments ca 1 mm long, white, anthers ca 0.6 by 0.4 mm. Pistillate flowers: pedicel 2.2—8 mm long, 1.3—2 mm from top with abscission zone, often gland near this zone or absent but sometimes with thickened glabrous parts (Borneo); sepals 5(6), broad triangular lower part, narrow triangular upper part, of different sizes, 1.5—7 by 0.7—1.5 mm, outside hairy, often with a single to several papillae-like glands along lower part of margins; ovary subglobose, 1.5—3.2 by 1.5—3.2 mm, 3-locular, densely hairy, somewhat muricate, dull light green; style 0.5—1.3 mm long, light green to red; stigmas 4—9.8 mm long, with short papillae adaxially, light green to red. Fruits 7.5—10.5 by 7—10 mm, muricate in upper half, pilose, glabrescent, yellowish green; columella T-shaped, 5—5.5 mm long, slender, only basally some remnants of septae. Seeds 5.5—6 by 4.5—5 by 3.3—4 mm, somewhat warty.

    Distribution — Thailand, Malay Peninsula, Sumatra, Borneo, Philippines.

   Habitat & Ecology —  Primary forest, secondary forest, along road sides; soil: clay-loam, limestone. Altitude: 50—1300 m. Flowering probably whole year round; fruiting: February to July.

    Vernacular names — Philippines: Malacboc (Vidal 1886).

    Notes —  1. This species had different names on the major islands groups in Malesia: Alchornea parviflora on the Philippines (accepted name), A. borneensis on Borneo, A. adenophylla on Sumatra and the Malay Peninsula, and A. trewioides in Thailand (wrong interpretation of this species, see note 3). The synonymy of the name A. adenophylla by Pax & Hoffmann (1914) is problematic, because the type of A. adenophylla could not be located. Most specimens identified as A. adenophylla are A. parviflora, and, therefore, this name is tentatively treated as synonym of A. parviflora. This view is consistent with the species Pax & Hoffmann (1914) recognized for Sumatra and the Malay Peninsula.

2. Vidal (1886) accidentally made the new combination Acalypha tiliifolia instead of using the name Alchornea tiliifolia (Benth.) Mόll.Arg., when he tried to identify his specimen, Vidal 592 (the confusion is easy to explain, both genera are treated on the same page and under the species the genus name is always abbreviated to A.). Unfortunately, Alchornea tiliifolia does not occur in the Philippines, thus Vidal not only made a new combination, he also applied it incorrectly. In this case Murphy’s law applies, the collection is also a mixtum: The Kew (K) duplicate of Vidal 592 is indeed Alchornea parviflora, but the Leiden (L) specimen is Mallotus peltatus (Geiseler) Mόll.Arg.

3. Alchornea parviflora and A. trewioides were confused in Thailand. Alchornea trewioides occurs in S.E. China and N. Vietnam, and the specimens in Thailand belong to A. parviflora. The two species differ in quite a few characters (between brackets the character states in A. trewioides). Alchornea parviflora has slender leaves: length/width ratio of the blade 1.6-3 (1.2-1.6); the stipellae are shorter, 1.2-2.5 mm long (2.3—4.3(-10) mm); the staminate buds and sepals are hairy, though sometimes glabrous in the Philippines, but then the inflorescences are branching (glabrous sepals, unbranched inflorescences); the staminate bracts, 0.6-0.7 mm long, and the pistillate bracts, 0.8-4 mm, are usually shorter (0.6-1.2 and 2.3-5 mm, resp.); the ovary is warty (smooth); and the apex of the columella is T-shaped (not widened apically).


2. Alchornea rhodophylla Pax & K. Hoffm.


    Alchornea rhodophylla Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 249; Whitmore, Tree Fl. Malaya 2 (1973) 54; Welzen & Bulalacao, Syst. Bot. 32 (2007) 809, fig. 3. — Alchornea discolor Hook.f., Fl. Br. India 5 (1887) 421, nom. illig, non Engl. & Poepp., Nov. Gen. 3 (1845) 19. — Type: Wallich 7777 (holo K-W; iso K, 2x!), Malaysia, Penang.

    Alchornea villosa (Benth.) Mόll.Arg. var. glabrata Hook.f., Fl. Br. India 5 (1887) 421; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.viiv(1914) 248; Ridl., Fl. Malay Penins. 3 (1924) 277. — Type: Curtis 214 (holo K!), Malaysia, Penang.


Shrubs to small bushy trees, up to 3.5 m high; flowering branches 2—5 mm in diameter, shortly, not densely sericeous when young, glabrescent. Stipules 4—9 by 0.6—1 mm. Leaves: petiole 1—7 cm long, round in transverse section, basally and less so apically indistinctly (constricted) pulvinate, pilose; stipellae ovate, 2.3—6.5 by 0.5—1.5 mm, hanging, succulent, margin and lower surface hairy, no marginal basal glands; blade ovate, 6—20 by 2—7.5 cm, length/width ratio 1.2—3.8, broadest at lower third, usually drying reddish brown, base narrowly rounded, margin laxly serrate with upwardly bent glandular teeth, apex cuspidate to caudate, often mucronate, upper surface shortly, sparsely hairy on basal part of midrib and nerves, extrafloral nectaries absent; lower surface green to magenta, shortly, sparsely hairy on midrib and nerves, basally usually 2( or 4) round slightly raised extrafloral nectaries, apical additional ones absent, glabrescent hair tuft domatia in nerve axils; venation triplinerved, not well visible above, raised beneath, nerves 8 or 9 up to the apex. Inflorescences racemes, ramiflorous (then often with big brachyblasts when staminate), axillary and terminal when generally pistillate, single (both sexes) or several together (staminate), staminate ones up to 16 cm long, pistillate ones up to 8.5 cm long, shortly pilose, extrafloral nectaries sometimes present beneath nodes or on lower pedicels; bracts triangular, hairy outside, without basal glands, staminate ones small, 0.5—1.2 by 0.3—1 mm, pistillate ones 1.8—4.7 by ca 1 mm, basally succulent, papery, venation not visible; bracteoles several, on node and very small (staminate), or in bract or along pedicel (pistillate), triangular, much smaller than bracts, hairy outside. Staminate flowers in groups of up to 6, ca 3 mm in diameter, white; pedicel ca 0.5 mm long, hairy, probably with abscission zone; sepals 4 (usually 2 together), ovate, 1.2—1.8 by 0.7—0.9 mm, apically acuminate, hairy outside; stamens 7 or 8, filaments 0.5—0.7 mm long, anthers 0.5—0.9 by 0.4—0.6 mm. Pistillate flowers ca 2 mm in diameter (without stigmas); pedicel 3.7—10 mm long, 2—3 mm from top with abscission zone, with raised gland beneath sepals; sepals (5? or) 6, triangular, base broadened, 2.3—3.5 by 0.8—1.2 mm; ovary ca 1.5 by 1.5 mm, 3-locular, smooth (to somewhat muricate); style 1.3—1.5 mm long; stigmas 7.5—17 mm long. Fruits 9—12.5 by 7—7.5 mm, smooth (to somewhat muricate), shortly sericeous, glabrescent; columella T-shaped with arms somewhat upright, 4.5—5 mm long. Seeds 4.8—5.8 by 4—5.3 by 2.7—4 mm, somewhat warty.

    Distribution — Endemic in Malaysia, State of Penang, Penang Island.

    Habitat & Ecology — Primary forest, mountain forest, by roadside. Altitude: 600—900 m. Flowering: May, September, and November; fruiting: March and June.

    Note — This species is endemic on Penang Island on the N.W. coast of Peninsular Malaysia. The species is names after the reddish drying leaves (a typical but not unique character). Typical are the smooth ovaries and fruits and the absence of extrafloral nectaries on the lower leaf blade surface near the apex (only basally present).


3. Alchornea rugosa (Lour.) Mόll.Arg.


    Alchornea rugosa (Lour.) Mόll.Arg., Linnaea 34 (1865) 170; in DC., Prodr. 15, 2 (1866) 905; Kurz, Forest Fl. Burma 2 (1877) 385; Hook.f., Fl. Br. India 5 (1890) 422; J.J.Sm., Meded. Dept. Landb. Ned.-Indiλ 10 (1910) 467; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 243; Merr., Enum. Philipp. Fl. Pl. 2 (1923) 438; Gagnepain in Lecomte, Fl. Indo-Chine 5 (1926) 379; Merr., Trans. Amer. Philos. Soc., New Ser. 24 (1935) 237; Holthuis & Lam, Blumea 5 (1942) 199; Airy Shaw, Kew Bull. 26 (1972) 211; Whitmore, Tree Fl. Malaya 2 (1973) 53; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 28; Kew Bull. 35 (1980) 594; Kew Bull. Add. Ser. 8 (1980) 25; Kew Bull. 36 (1981) 251, Fig. 1: B.1-4; Kew Bull. 37 (1982) 4; Alphab. Enum. Euph. Philipp. Isl. (1983) 4; Welzen & Bulalacao, Syst. Bot. 32 (2007) 809, fig. 1d; 2a-h; 4. — Cladodes rugosa Lour., Fl. Cochinch. (1790) 574. — Type: Loureiro s.n. (holo BM), Cochinchine.

    [Croton apetala Blume, Catalogus (1823) 104, nom. nud. --- vernacular: ki-leat.]

    Conceveiba javanensisConceveibum javanense’) Blume, Bijdr. (1825) 614; Fern.-Vill., Nov. App. (1880) 195. — Aparisthmium javense (‘javensis’) (Blume) Endl. ex Hassk., Cat. Hort. Bot. Bogor (1844) 235; Zoll., Linnaea 28 (1857) 330. — Alchornea javensis (Blume) Mόll.Arg., Linnaea 34 (1865) 170; in DC., Prodr. 15, 2 (1866) 905; Ridl., Fl. Malay Penins. 3 (1924) 278; Backer & Bakh.f., Fl. Java 1 (1963) 485. — Lectotype (Welzen & Bulalacao, 2007): Blume 1334 (hololecto L), Indonesia, Java. See note 1.

    Adelia glandulosa Blanco, Fl. Filip. (1837) 814 (see Merr., Spec. Blanc., 1918, 225, for interpretation). — Neotype (Welzen & Bulalacao, 2007): Merrill Species Blancoanae 725 (holo L), Philippines, Luzon, Rizal Prov., Montalban. See note 2.

    Tragia innocua Blanco, Fl. Filip., 2nd ed. (1845) 479 (non Walter, Fl. Carol.: 229. 1788); 3rd ed. (1879) 94 (see Merr., Spec. Blanc., 1918, 225, for interpretation). — Neotype (Welzen & Bulalacao, 2007): BS (M. Ramos) 7664 (holo L), Philippines, Luzon, Ilocos Norte Prov. See note 2.

    Alchornea hainanensis Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 242. — Alchornea hainanensis Pax & K.Hoffm. var. glabrescens Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 242, nom. illig. — Syntypes: Henry 8119Y (K), China, Hainan; Henry 8536 (K), China, Hainan; Henry 8778 ( K), China, Hainan; Swinhoe s.n. (K), China, Hainan. See note 3.

    Alchornea hainanensis Pax & K.Hoffm. var. pubescens Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 243. — Alchornea rugosa (Lour.) Mόll.Arg. var. pubescens (Pax & K.Hoffm.) H.S.Kiu, Fl. Reipubl. Popul. Sin. 44, 2 (1996) 69. — Type: Henry 8543 (n.v.), China, Hainan.

    Alchornea pubescens Merr. (non (Britton) Secco, 1999), Philipp. J. Sc. 20 (1922) 399; Enum. Philipp. Fl. Pl. 2 (1923) 438; Airy Shaw, Alphab. Enum. Euphorb. Philipp. Isl. (1983) 4. — Type:FB (Castillo) 22724 (holo PNH, destroyed; iso K), Philippines, Luzon, Cagayan Prov., Peρablanca. See note 4.

    Alchornea rugosa (Lour.) Mόll.Arg. var. macrocarpa Airy Shaw, Kew Bull. 26 (1972) 211. — Type:Larsen, Santisuk & Warncke 2398 (holo AAU), Thailand, Northern, 15 km E of Mae Sariang. See note 5.

    Alchornea petalostyla Airy Shaw, Kew Bull. 35 (1980) 395; Alphab. Enum. Euphorb. Philipp. Isl. (1983) 4. — Type: Vidal y Soler 3719 (holotype: K), Philippines, Luzon, Nueva Vizcaya Prov., Bayombong. See note 6.

    Actephila excelsa auct. non Mόll.Arg.: Koord., Gedenkb. Franz Junghuhn (1910) 172.


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Shrubs to trees up to 10 m tall, d.b.h. up to 15 cm, monoecious, sexes usually on different branches; flowering branches glabrous, puberulous to hirsute , lenticellate, 2—6 mm thick. Outer bark pale to dark grey to pale brown, with lichens, smooth to finely fissured, sometimes peeling off in thin, minute flakes; inner bark white or pinky brown, wood creamy white, finely textured, moderately hard and heavy. Stipules 2—12 by 0.2—1.2 mm. Leaves: petioles 0.1—6 cm long, flat to channelled above, basally and apically to completely somewhat pulvinate, subglabrous to pubescent to hispid; stipellae absent; blade (elliptic to) obovate, 4—34.5 by 1.8—13 cm, length/width ratio 1.5—5.3, base narrowly cordate (to cuneate), margin serrate or dentate with 15—20 small teeth, sometimes ending in upwards pointing glands, apex acuminate to caudate, upper surface glabrous to pubescent with scattered short hairs on midrib and veins, dark green, lower surface glabrous except for scattered hairs on midrib and veins, with or without hair tuft domatia, dark to lighter green, extrafloral nectaries seldom absent, either present on the upper surface, or usually on the lower surface or on both surfaces, usually at least 2 basally present near petiole insertion, others parallel to midrib or margin, oval to circular, crater-like with reddish black wall, shiny and striate inside; venation penninerved, up to 12 nerves, distinct below. Staminate inflorescences strongly branched (ca 15—50 times) (sub)terminal panicles or single long thyrsoid racemes, up to 42 cm long, or less branched (5—10 times) axillary panicles up to 9 cm long, green to pale yellow, flowers in clusters of 9-25 per node; bracts rhomboid, 1—1.7 by 0.9—1.6 mm, puberulous outside, glabrous inside, circular glands usually at the base; bracteoles deltoid, 0.4—0.6 by 0.8—1 mm, hirsute outside, glabrous inside; stalked glands cylindrical, ca 0.5 by 0.2 mm, hirsute. Staminate flowers ca 3 mm in diameter, yellow-green to pale green to reddish, buds green to maroon; pedicel 0.25-0.75 mm long; calyx 2—4-partite, 1.8—1.9 by 1—1.7 mm , margin membranous; stamens 4 (or 6 or 8, see note 3), dark pink, filaments up to 1 mm long, whitish to black, glabrous, anthers 0.9—1 by 0.9—1 mm, light yellow or cream. Pistillate inflorescences terminal groups of racemes or panicles, up to 23 cm long or axillary racemes ca 4 cm long, rachis green, ribbed, puberulous; bracts triangular, basally usually with 2 glands or 2 bracteoles, just beneath flower, ca 1.3 by 1.3 mm, outside slightly hairy. Pistillate flowers ca 3 mm in diameter, green; pedicel 1—4 mm long, sometimes with a distinct abscission zone, seldom with one or a few elevated, round glands; sepals 4 or 5, rhomboid to triangular, 1—3.5 by 0.7—1.8 mm, margin denticulate, a few ending in glands, sometimes an elevated round gland present; ovary green or purplish, 3(4)-locular (see note 7); style short, 0.5—1(—2) mm long (see note 8), stigmas trifurcate, narrowly triangular (to broadly obovate, see note 5), up to 5.8 mm long (see notes 5 and 7) and 1.2 mm wide (see note 5), apically seldom split once or several times (note 6), decurrent, often recurved, light green to yellow or reddish. Fruits 3—12 by 4—10 mm, purplish red to finally brown, subglabrous to pubescent outside, glabrous inside; columella 3—5 mm long, apically narrowly obtriangular. Seeds 4.5—7.3 by 4.5—6 by 3.5—4.8 mm, (somewhat) pustular with pustules more or less in rows.

    Distribution — Sri Lanka, India (Assam, Nicobar Islands), Myanmar, Thailand, Vietnam, China (Hainan), Malesia: Peninsular Malaysia, Sumatra, Java, Borneo (Sarawak, Sabah), Philippines, Sulawesi, Lesser Sunda Islands, Moluccas, New Guinea, Australia (Queensland).

    Habitat & Ecology — Usually frequent in understorey in shade or semi-shade of primary (Dipterocarp) rain forest, evergreen forest, deciduous forest, gallery forest, secondary forest to deforested and very degraded seasonal forest with bamboo, scrubs, and even in alang-alang (grass fields), very often in wet places like marshes or along water courses; also in cultivated places, like shifting cultivation, shrubbery of coconut and narra (Pterocarpus indicus) plantations. Soil: various, often on or near limestone, partly even on guano near caves, clay loam, deep clay, ultramafic; bedrock: shale, quartzite, sandstone, granite. Altitude: sea level up to 1200(—2800) m. Flowering and fruiting whole year through.

    Vernacular names — Thailand: Vietnam: Bo met, Cay soi giai, Cay muoi, Cay chua mot, Cay dat mot (Gagnepain 1926). China: Foo tsa. Java: Kibewok, ki-leat, kissengat (Blume, 1823, 1825); singugu; tapen (Sundanes: Jasilin dial.); tjotjok kilik. Borneo: Sabah: Boot (Suluk). Philippines: Aguyui, kanauai, tayan (Tagalog); banauli (S.L. Bis.); gasugasi, halum, malambingan (Sul.); gaugau (C. Bis.); limo, samburagat-bukid (Tagbanua); makabangon (partly after Merrill 1923). Celebes: Umbana. Moluccas: Ngofa dofu (Halmahera). New Guinea: Ara fatem.

    Uses — Wood for fuel (Philippines). Leaves are used as vegetable (Moluccas). Seeds used as purgative (Gagnepain 1926). Leaves used for cigaret paper (Irian Jaya).

    Notes — 1. Blume (1825) described Conceveiba javanensis (a synonym of A. rugosa), which was transferred to the genus Aparisthmium by Hasskarl (1844), but with an incorrectly spelled epithet: ‘javense’. Mόller (1865) transferred the species to the genus Alchornea, but still maintained the incorrect spelling. The spelling was improved by Backer and Bakhuizen van den Brink f. (1963). Airy Shaw (1975) treats the two spellings (‘javanensis’ and ‘javense’) as different species, but all authors clearly refer to Blume and thus to the same type.

2. Merrill (1918) selected Merrill Species Blancoanae 725 as illustrative specimen for two names of Blanco: Adelia glandulosa and Tragia innocua. This specimen cannot be used as neotype for both names, therefore, we selected it as neotype for Adelia glandulosa, the name most discussed by Merrill. For Tragia innocua we chose a specimen collected by Ramos as neotype (BS 7664), because Ramos collected most specimens for the Species Blancoanae series of Merrill (Van Steenis-Kruseman 1950: 427a).

3. Alchornea hainanensis differs in the number of stamens (6 or 8 instead of 4), no other characters are found in which it differs from rugosa. The number of stamens varies between 4 and 8 in Alchornea, therefore, A. hainanensis is synonymised with A. rugosa.

4. Typical for A. pubescens is the very dense indumentum on twigs and lower leaf surface, the large, narrow leaves, and the long cuspidate apex. Also mentioned by Merrill (1922, 1923) was the high(er) number of nerves. The species is known only from the type and that makes its status suspect. The number of nerves does not differ from many more typical A. rugosa specimens, nor is the apex really typical (other specimens of A. rugosa are also caudate). The leaves are indeed long and slender, but again, this is not exceptional (e.g., compare with Cuming 1204 and Vidal y Soler 1766). Most striking is the indumentum of the type specimen, but intermediates with the (sub)glabrous specimens of A. rugosa are present (e.g., the two specimens identified as A. petalostyla, see note 5, Cuming 990 and Vidal y Soler 1766). On the other hand, specimens from other areas can be very hairy, too (Petelot 1175 from Vietnam, Amdjah 168 from Java, S 21688 from Borneo, PNH 10651 from the Philippines). Thus, A. pubescens has to be treated as a synonym of A. rugosa.

5. At first we wanted to maintain variety macrocarpa. The type (N. Thailand) has indeed much larger fruits (12 by 10 mm versus 3—8 by 4—7 mm), large (but not larger) and elliptic leaf blades (14—30 by 6—13 cm versus 4—32 by 1.8—8.5 cm and generally obovate in common rugosa), and an acute blade base (instead of narrowly cordate). Also typical are the stigmas which have multiple frays at the end. Similar specimens, but with normal-sized fruits are present in S. Thailand (J.F. Maxwell 85-454, 87-443, 87-444) and N. Peninsular Malaysia (Stone et al. 15217) and in S. Peninsular Malaysia (Ahmad 278, Loh Hoy Shing KEP FRI 6867). This is a very disjunct distribution. However, the elliptic shape and cuneate leaf base are also found in Indian, Thai, and Vietnamese specimens, which further equal normal rugosa. Moreover, the ‘macrocarpa’ specimens from S. Thailand and Peninsular Malaysia are often (partly) slightly obovate. The multiple frayed stigmas occur very accidently within A. rugosa (see next note). Thus, only the type specimen is really exceptional in its large fruits. As it is the only specimen showing this character we consider it to be an exceptional specimen rather than a good variety.

6. Airy Shaw (1980) described A. petalostyla, and indeed the type specimen shows peculiar, large petal-like stigmas, which are broad and obovate with an erose rounded apex, up to 1.7 by 1.2 mm. In typical A. rugosa the stigmas are narrowly triangular and much longer (but see next note) with a gradually acute tip. Several specimens bridge the gap in stigma sizes between A. petalostyla and A. rugosa: R.C. McGregor BS 11320 (also identified as A. petalostyla by Airy Shaw) to Castro, Barbon & Garcia PPI 22287, Quisumbing PNH 8049 to Edaρo PNH 15443 and all other A. rugosa specimens. Moreover, several specimens have stigmas with a single to multiple split apex, these are in principal also obovate, but these specimens look like normal A. rugosa. No other characters support the separate status of A. petalostyla, therefore, we interpret it as a synonym of A. rugosa. Similar styles, but then triangular and very adnate to the ovary, are also found on Sulawesi (de Vriese & Teijsmann s.n., L  0384092; Sidiyasa & Didi 1866) and the Moluccas (Toxopeus 301).

7. Two specimens, Sidiyasa & Didi 1866 from Sulawesi and Kostermans 775 from the Moluccas (Morotai), are exceptional in a sometimes 4-locular ovary and especially in the very short stigmas, less than 1 mm long.

8. Generally, pistillate flowers hardly possess a style, exceptional is De Vogel 3062 from the Moluccas with ca 2 mm long styles.


4. Alchornea sicca (Blanco) Merr.


    Alchornea sicca (Blanco) Merr., Philipp. J. Sc. 5, Bot. (1910) 192; Philipp. J. Sc. 7, Bot. (1912) 383; Sp. Blancoan. (1918) 224; Enum. Philipp. Fl. Pl. 2 (1923) 439; Airy Shaw, Alphab. Enum. Euphorb. Philipp. Isl. (1983) 4; Welzen & Bulalacao, Syst. Bot. 32 (2007) 813, fig. 3. — Excoecaria sicca Blanco, Fl. Filip. (1837) 787; Fl. Filip., ed. 2 (1845) 542; Fl. Filip., ed. 3, 3 (1879) 193, t. 307. — Neotype (Welzen & Bulalacao, 2007): Merrill Species Blancoanae 642 (holo K), Philippines, Luzon Island, Rizal Prov., Pasig River, opposite Guadalupe.

    Croton drupaceus (‘drupaceum’) Blanco, Fl. Filip., ed. 2 (1845) 519, nom illig, non Roxb., Fl. Ind. 3 (1832) 684. — Neotype (Welzen & Bulalacao, 2007): Elmer 9410 (neotype: K!, here designated), Philippines, Luzon Island, Laguna Prov., Los Baρos (see note 2).

    Alchornea philippinensis Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 249. — Type: Merrill 988 (holo PNH or B, destroyed; iso L), Philippines, Luzon Island, Manila (see note 3).

    Alchornea coudercii Gagnep., Bull. Soc. Bot. France 71 (1924) 138; in Lecomte, Fl. Indo-Chine 5 (1926) 380, fig. 44: 1. — Syntypes: Thorel s.n. (P), Laos, Compong-luong; Thorel s.n. (P), Laos, Oudong; Couderc s.n. ( P, 3 sheets), Cambodia, Wat-preas near Kompur; Godefroy (or Harmand) s.n. (P, 2 sheets), Cambodia; Pierre s.n. (P), Cochinchine, Mt. Lu; Talmy s.n. (P) Cochinchine, Saigon Botanical Garden; Thorel s.n. (P). Cochinchine, Cai-cong (see note 4).


Shrubs to small trees, probably deciduous in part of Indochina (flowering when in leaf), flowering branches 0.8-3 mm in diameter, slightly pilose when young, glabrescent. Indumentum yellowish white, hardly present. Bark thin, smooth, grey. Stipules 3.5—7.5 by 0.3—0.6 mm. Leaves: petiole 0.3—6 cm long, round to reniform in transverse section, very indistinctly basally pulvinate, mainly adaxially pilose, quickly glabrescent; stipellae subulate, 0.2—3.5 mm long, erect to patent; blade ovate, 1.5—11 by 1.2—7 cm, length/width ratio 1.5—3, broadest at lower third, base slightly emarginate to rounded, glabrous, margin laxly serrate with glandular teeth, apex cuneate to cuspidate, mucronulate, upper surface subglabrous, few hairs on (basal part of) nerves, extrafloral nectaries absent, dull dark green; lower surface slightly hairy on nerves, dull light green, basally 2 to several round slightly raised extrafloral nectaries outside the basal nerves, many additional ones along margin or in the blade, often hair tuft domatia in nerve axils and axils of nerves with veins; venation triplinerved, slightly raised adaxially, raised abaxially, nerves 5—8. Inflorescences single, axillary and terminal racemes, up to 7 cm long, (subglabrous to) somewhat pilose, green; bracts basally leaf-like (not measured), upper ones triangular to ovate, 1.4—2 by 1.3—2.4 mm, acuminate, stiff, midrib usually raised abaxially, often striate because of parallel venation, margin hairy, outside few hairs, basally big glands especially in pistillate plants, pale light green; bracteoles distinct, staminate ones ovate, 1.5—1.7 by 0.2—0.3 mm, pistillate ones 1.1—1.9 by 0.5—0.8 mm, stiff, hairy like bracts, pale light green. Staminate flowers only seen in bud or partly destroyed by insects, several per node, 2—3.7 mm in diameter; pedicel ca 0.5 mm, apically with an abscission zone, sepals 2(—4), ovate, 1.2—2 by 0.6—1.6 mm, few hairs outside, pale red; stamens 6 (or 8), filaments 0.5—1 mm long, dull red and cream, anthers ca 0.8 by 0.5—0.6 mm. Pistillate flowers sessile; sepals (3—)5, often still several united, ovate, of different sizes, 0.7—1.1 by 0.6—0.8 mm, margin hairy, apex rounded to emarginate, outside subglabrous, one sepal basally often with a single raised glands, pale light green; ovary 1—1.3 by 1—1.5 mm, 3-locular, smooth; style absent; stigmas 5.2—7.8 mm long, maroon. Fruits 6—7 by 7.5—8 mm, smooth, glabrescent; columella ca 4.8 mm long, very slender, apically narrowly obtriangular. Seeds ca 5 by 4 by 3.5 mm, very warty.

    Distribution — Laos, Cambodia, Vietnam, Philippines (Luzon: Rizal & Laguna Prov.).

    Habitat & Ecology — Riverbanks and open secondary growth thicket. Rhyolite bedrock. Altitude: ca. 75 m. Flowering: March to May.

    Vernacular names — Philippines: Balanti (Tagalog).

    Notes — 1. A very distinct, but poorly collected species. Especially the large, stiff bracts are remarkable.

2. Merrill (1918) refers to one representative specimen for two Blanco names: Excoecaria sicca and Croton drupaceus, of which the original type material is lost. This specimen (Merrill Species Blancoanae 642) is selected as neotype for Excoecaria sicca. Of Alchornea sicca three Philippine specimens are known, two are already used as types, either as neotype for E. sicca, or (Merrill 988) as type for A. philippinensis (see note 3), this leaves the third specimen (Elmer 9410) to be selected as neotype for C. drupaceus.

3. When Pax & Hoffmann (1914) described A. philippinensis they referred to Merrill’s article (1912) in which Merrill stated that in the Philippines not one species (A. sicca with A. parviflora as synonym, see Merrill, 1910), but two species are present, A. sicca and A. parviflora. Merrill separated these two again. Unfortunately, Pax & Hoffmann interpreted Merrill incorrectly, they thought that Merrill still kept both names together under A. sicca (see their treatment on page 251) and that his separate taxon was still undescribed, after which they described it as A. philippinensis.

4. The Indochinese material, formerly known as A. coudercii Gagnep., is in all characters equal to A. sicca of the Philippines. The only difference is in the usually larger size of the leaves and petioles (generally larger in Indochina), but overlap is present, thus A. coudercii cannot be recognized as a separate species.


5. Alchornea tiliifolia (Benth.) Mόll.Arg.


    Alchornea tiliifolia (Benth.) Mόll.Arg., Linnaea 34 (1865) 168; in DC., Prodr. 15, 2 (1866) 903; Kurz, Forest Fl. Burma 2 (1877) 385; Hook.f., Fl. Br. India 5 (1887) 421; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 250; Ridl., Fl. Malay Penins. 3 (1924) 227; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1926) 382; Airy Shaw, Kew Bull. 26 (1972) 212; Whitmore, Tree Fl. Malaya 2 (1973) 54; Welzen & Bulalacao, Syst. Bot. 32 (2007) 814, fig. 1a, e; 5. — Stipellaria tiliaefolia Benth., Hooker’s J. Bot. Kew Gard. Misc. 6 (1854) 4.. — Type: Wallich 7829 (holo K-W; iso K), India, Sillet.

    [Croton chiamala Wall., Numer. List (1847) no. 7775, nom. nud. — Representative specimen: Wallich 7775 (K-W, n.v.), India, Goygalpura]

    Stipellaria villosa Benth., Hooker’s J. Bot. Kew Gard. Misc. 6 (1854) 4. 1 — Alchornea villosa (Benth.) Mόll.Arg., Linnaea 34 (1865) 168; in DC., Prodr. 15, 2 (1866) 902; Hook.f., Fl. Br. India 5 (1887) 421; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 247; Ridl., Fl. Malay Penins. 3 (1924) 277; Whitmore, Tree Fl. Malaya 2 (1973) 54; Airy Shaw, Kew Bull. 36 (1981) 251. — Alchornea villosa (Benth.) Mόll.Arg. var. genuina Mόll.Arg. in DC., Prodr. 15, 2 (1866) 902, nom. illeg; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 248. — Lectotype (Welzen & Bulalacao, 2007): Cuming 2307 (holo K; iso: K), Malaysia. See note 1.

    Alchornea zollingeri Hassk., Retzia (1855) 156. — Bleekeria zollingeri (Hassk.) Miq., Fl. Ned. Ind. 1 (2) (1859) 407. — Stipellaria zollingeri (Hassk.) Miq., Fl. Ned. Ind, eerste bijv. (1861) 182, 460. — Type (Miquel, 1859): Zollinger 3054 (= Teijsmann HB 4189) (holo L; iso K, P), Indonesia, Sumatra, Lampong [= Lampung Prov.], grown in Kebun Raya (Bogor Botanical Garden). See note 2.

    Aparisthmium sumatranum Rchb.f. & Zoll. in Zoll., Linnaea 28 (1856) 331. — Type: Zollinger 3054 (holo P; iso K, L), Sumatra, Lampong [= Lampung Prov.]. See note 2.

    Alchornea villosa (Benth.) Mόll.Arg. var. lanceolata Mόll.Arg. in DC., Prodr. 15, 2 (1866) 902; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 248. — Type: Zollinger 1809 (= Teijsmann HB 4186) (holo L; iso P), Indonesia, Sumatra, Lampong [= Lampung Prov.]. See note 2.

    Alchornea villosa (Benth.) Mόll.Arg. var. ? latisepala Hook.f., Fl. Br. India 5 (1887) 421; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 248. — Typte: Helfer KD 4752 (holo K), Burma, Tenasserim. See note 3.


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Evergreen shrubs to small trees, up to 11 m tall, d.b.h. up to 12 cm, sparsely branched, monoecious, sexes usually on different inflorescences; flowering branches hairy with short and long hairs, glabrescent, 2—7 mm thick. Outer bark thin, smooth, whitish to grey to tan; inner bark thin, white to brown; wood white. Stipules 4—14 by 0.8—2 mm. Leaves: petioles 0.6—18 cm long, round or reniform, basally (and apically) pulvinate, hirsute; stipellae long ovate and leaf-like (to subulate to more or less glandular; sometimes extra subulate or glandular ones beneath normal ones; see note 4), 3—18 by 0.6—7 mm, pendulous when leaf-like (to patent when subulate), margin usually with a single to a few glandular teeth, glabrous above, hairy underneath, reticulate nerves usually distinct above; blade ovate, broadest in lower third, but above insertion, 4.5—27 by 2—20 cm, length/width ratio 1—2.3, base emarginate to widely cuneate, margin laxly serrate with upwardly bent glandular teeth; apex acuminate to caudate, upper surface subglabrous to hairy on (basal part) venation, dark green, extrafloral nectaries absent; lower surface subglabrous (see note 4) to generally hairy all over, also between (visible) veins with very short hairs, lighter green, basally usually 2—4 round, slightly raised extrafloral nectaries, additional apical ones present, hair tuft domatia absent; venation triplinerved, visible above, but especially beneath, nerves 5—8 up to the apex. Inflorescences light green, erect, ramiflorous to axillary racemes (staminate) or terminal racemes (to basally branched) (pistillate), single or several together (mainly staminate), sometimes short pistillate ones among the staminate ones or apical flower(s) in staminate raceme pistillate, staminate inflorescences up to 15 cm long, pistillate ones up to 17 cm long, occasional side branch up to 12 cm long, all shortly pilose, extrafloral nectaries sometimes present beneath nodes or on the base of the bracts; rachis hairy, round; bracts triangular to ovate, hairy outside, with or without basal glands, staminate ones small, 0.8—3 by 0.8—2 mm, pistillate ones 1.5—7.7 by 0.3—1.3(—3.2) mm, papery, venation not visible, caducous; bracteoles several, on node and very small (staminate) or along pedicel (pistillate) and triangular, 1—5.5 by 0.3—1.7(—3) mm, hairy outside. Staminate flowers 2—3 mm in diameter, light greenish except the anthers; pedicel 0.3—1 mm long, abscission zone halfway, often more hairy than rest of pedicel; calyx 2-partite at anthesis, later (3)4(5) lobes, sepals ovoid, 1—1.4 by 0.7—1.3 mm, subhispid (to glabrous, Thailand) outside, glabrous inside; stamens 6(—8), connate at base on a ring-like receptacle, filaments 0.6—0.7 mm long, hairy, anthers oblong, 0.7—0.9 by 0.4—0.6 mm, cream. Pistillate flowers 3—4.5 mm in diameter, light green; pedicel absent to 11.5 mm long in fruit, with abscission zone beneath extrafloral nectaries; sepals 5-7(8), triangular, 2.8—8 by 0.6—3.5 mm, margin without glands to with glandular teeth towards the base, outside hairy, inside glabrous, large extrafloral nectaries often present beneath sepals; ovary 3-locular, 1.3—2.2 by 1—1.5 mm, subglabrous to hairy, muricate in upper part; style 0.4—6 mm long, basally broader than apically, stigmas 6.8—15 mm long, apex seldom somewhat split at the end. Fruits 7—10 by 7—12.5 mm, muricate (especially in upper third) when young to often nearly smooth when mature (see note 4), shortly hairy outside, glabrous inside, greyish brown to greyish black; columella 5—7.5 mm long, 3-angled, T-shaped with arms somewhat diagonal. Seeds 5.2—7.5 by 4.5—6 by 3.2—6 mm, dark brown, warty.

    Distribution — N.W. India, Myanmar, Laos, Thailand, N. Vietnam, Sumatra, Java, Borneo.

    Habitat & Ecology — Primary rain forest, evergreen forest, deciduous forest, secondary forest, montane forest, bamboo scrub, thickets, pioneer along road sides and in open places, often along water; soil: clay, granite, sandstone, shales. Altitude: 30-1300 m. Flowering and fruiting whole year through.

    Uses — The bark is used for string (Ridley 1924).

    Vernacular names — Vietnam: Cay dom dom, cay dong chau, cay nong lua (Gagnepain 1926). Malaysia: Bulan, lisau-lisau, ramin bukit, ramin hitam, sumin jantan (partly after Ridley 1924). Sumatra: Krangkrang, latih poetih, lisau-lisau, poedeng; leloeti/leluti (Malay).

    Notes — 1. At first sight, we wanted to keep Alchornea tiliifolia and A. villosa separate, because they seemingly differed in the type of stipellae, but intermediates were common and both names had to be united. Alchornea tiliifolia, whether in its former narrow sense, or in the wider sense (united with A. villosa) is, like A. rugosa, a rather variable species. Generally, the leaves are broadly ovate, and the bracts and pistillate sepals are large and broad, while the style is usually relatively long.

2. Two specimens (Zollinger 1809 and 3054) collected by Zollinger on Sumatra were used as types for various names. The specimens were seeminly brought at least partly alive to the botanical garden of Bogor (Kebun Raya) and samples of these botanical garden plants were used by Hasskarl (1855) to describe Alchornea zollingeri. Independently, based on (probably) another duplicate (in P), Reichenbach & Zollinger (1856) described Aparisthmium sumatranum based on Zollinger 3054. Miquel (1859) noticed that both names were based on the same collection and he united them under Bleekeria zollingeri (correctly choosing the oldest epithet) and later on he (1861) made a new combination in Stipellaria. Miquel (1859) only mentioned Zollinger 3054 and thus lectotypified Hasskarl’s name with this specimen. Later, Mόller (Argoviensis) (1866) used the other specimen (Zollinger 1809 from Bogor Botanical Garden, Hasskarl’s specimen) to describe a new variety under Alchornea villosa (var. lanceolata).

3. Alchornea villosa var. latisepala is only known from the type (Helfer KD4752) from Myanmar. It seems an exceptional specimen that differs in the width of the bracts, bracteoles and sepals. However, after the union of A. villosa and A. tiliifolia the measurements are not exceptional and, therefore, the name is treated as a synonym.

4. Thai specimens (except for the north and southern peninsula) tend to be rather glabrous, tend to have much smaller, gland-like stipellae and generally have almost smooth fruits (slightly muricate near style).


 Excluded species


Alchornea blumeana Mόll.Arg., Linnaea 34 (1865) 167; in DC., Prodr. 15, 2 (1866) 900. — Lectoype (Welzen et al., Blumea 43, 1998, 157) Blume 1521 (L, barcode L  0027207), Indonesia, Java = Wetria insignis (Steud.) Airy Shaw.


Pseudotrewia ? cuneifolia Miq., Fl. Ned. Ind. Eerste Bijv. (1861) 183, 462. — Alchornea cuneifolia (Miq.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 900; Miq., Ann. Mus. Bot. Lugd.-Bat. 4 (1869) 122cuneata’). — Type: Teijsmann HB 3883 (holotype: U), Indonesia, Sumatra orient. in prov. Palembang, prope Batu Radja. = Trigonostemon longifolius Baill. (see: Airy Shaw, Kew Bull. 36: 355. 1981)