Malesian Euphorbiaceae Descriptions

54. HANCEA (Euphorbiaceae)

J.W.F.Slik, S.E.C. Sierra, K.K.M. Kulju, P.C. van Welzen & M. Aparicio

Slik, J.W.F. & P.C. van Welzen. 2001. A taxanomic revision of Mallotus sections Hancea and Stylanthus (Euphorbiaceae). Bluma 43: 3–66.

Sierra, S.E.C., M. Aparicia, K.K.M. Kulju, Ž. Fišer, P.C. van Welzen & R.W.J.M. van der Ham. 2006. Re-shaping Mallotus [Part 1]: Expanded cicrumscription and revision of the genus Cordemoya (Euphorbiaceae). Blumea 51: 519–540.

Sierra, S.E.C., K.K.M. Kulju, J.F. Veldkamp & P.C. van Welzen. 2007. Resurrection of Hancea and lectotypification of Adisca (Euphorbiaceae). Blumea 52: 361–366.

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Subgenus Hancea

Key to the Sections

Section Hancea

Key to the Malesian species (of subgen. Hancea)

Species descriptions of sect. Hancea

Section Oliganthae

Species desctiption of sect. Oliganthae

Hancea Seem.

    Hancea Seem., Bot. Voy. Herald (1857) 409; S.E.C.Sierra, Kulju, Veldkamp & Welzen, Blumea 52 (2007) 362; G.L.Webster in Kubitzki, Fam. Gen. Vasc. Pl. 11 (2014) 128, Fig. 29. — Hancea Seem. subgen. Hancea: S.E.C.Sierra, Kulju, Veldkamp & Welzen, Blumea 52 (2007) 362. — Mallotus sect. Hancea (Seem.) Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 199; Airy Shaw, Kew Bull. 20 (1966) 39; Kew Bull. 21 (1968) 388; Kew Bull. 23 (1969) 80; Kew Bull. 26 (1972) 293; Kew Bull. 27 (1972) 86; Kew Bull. Add. Ser. 4 (1975) 160; Kew Bull. Add. Ser. 8 (1980) 164; Kew Bull. 36 (1981) 324; Slik & Welzen, Blumea 46 (2001) 4; in Slik, Tropenbos-Kalimantan ser. 4 (2001) 146. — Type: Hancea hookeriana Seem.

    Cordemoya Baill., Adansonia 1 (1861) 255. — Boutonia Bojer, Hort. Maurit. (1837) 282, nom. nud., non DC. (1838); in Bouton, Trav. Soc. Hist. Nat. Maurice (1846) 151. — Mallotus Lour. sect. Cordemoya (Baill.) Müll.Arg., Linnaea 34 (1865) 186. — Echinus Lour. sect. Cordemoya (Baill.) Baill., Bull. Mens. Soc. Linn. Paris 2 (1891) 977. — Cordemoya Baill. subg. Cordemoya: S.E.C.Sierra et al., Blumea 51 (2006) 526. — Hancea Seem. subg. Cordemoya (Baill.) S.E.C.Sierra, Kulju & Welzen in S.E.C.Sierra, Kulju, Veldkamp & Welzen, Blumea 52 (2007) 362. — Type: Boutonia mascareinensis Bojer [= Hancea integrifolia (Willd.) S.E.C.Sierra, Kulju & Welzen.

Shrubs to big trees, monoecious and/or dioecious; branches with slender or conspicuously swollen nodes. Indumentum composed of simple hairs and sometimes capitate glandular hairs with multicellular stalks (subg. Hancea and H. capuronii), or sessile peltate-stellate hairs with a central cell (subg. Cordemoya). Resin sometimes present (H. integrifolia and H. spinulosa). Stipules axillary or interpetiolar. Leaves simple, alternate or opposite (strongly unequal or not), chartaceous to (sub)coriaceous, adaxial extrafloral nectaries absent or when present inconspicuous; venation pinnate or triplinerved (rarely palminerved), midrib, nerves and veins prominent, veins scalariform, veinlets reticulate. Inflorescences ramiflorous, axillary, or terminal, unisexual to bisexual, with one bract per node; staminate ones with 1–3 flowers per bract, bracts persistent, bracteoles absent; pistillate ones with 1 flower per bract, bracts persistent to caducous, bracteoles rarely present. Flowers actinomorphic, not exceeding 1 cm diam.; sepals persistent; petals and disc absent. Staminate flowers: sepals 2–4, valvate, free to basally connate; stamens: thecae 2, parallel, opening extrorsely and lengthwise; pollen ornamentation areolate with scabrae; pistillode absent. Pistillate flowers: sepals (3)4–6(7), valvate or imbricate, free; staminodes rarely present; ovary with 1 ovule per locule; style short, stigmas erect, inner surface densely covered with short papillae, outer surface hairy. Fruits dehiscent capsules, opening loculicidally-septicidally or septicidally-loculicidally, spiny, spines sometimes with terminal glands. Seeds (sub)globose, somewhat trigonous in transverse section; caruncle or aril absent.

    Distribution — Madagascar, Mauritius, Réunion, from South China throughout Southeast Asia and Malesia to New Guinea.

Subgenus Hancea

    Hancea Seem. subgen. Hancea: S.E.C.Sierra, Kulju, Veldkamp & Welzen, Blumea 52 (2007) 362. — Mallotus sect. Hancea (Seem.) Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 199; Airy Shaw, Kew Bull. 20 (1966) 39; Kew Bull. 21 (1968) 388; Kew Bull. 23 (1969) 80; Kew Bull. 26 (1972) 293; Kew Bull. 27 (1972) 86; Kew Bull. Add. Ser. 4 (1975) 160; Kew Bull. Add. Ser. 8 (1980) 164; Kew Bull. 36 (1981) 324; Slik & Welzen, Blumea 46 (2001) 4; in Slik, Tropenbos-Kalimantan ser. 4 (2001) 146. — Type species as genus: Hancea hookeriana Seem.

Branches with slender or conspicuously swollen nodes. Indumentum composed of simple hairs and capitate glandular hairs with multicellular stalks. Stipules interpetiolar or axillary, margin entire or serrate. Leaves opposite (strongly unequal) or alternate; blade peltate or not, chartaceous to coriaceous, margin entire (to rarely serrate), extrafloral nectaries absent; venation pinnate or triplinerved (rarely palminerved), looping close to the margin (rarely ending in the margin). Inflorescences axillary or ramiflorous, with one flower per bract. Staminate inflorescences racemes or panicles. Staminate flowers: sepals (2 or) 3 or 4, free to basally connate; connective sometimes widened. Pistillate inflorescences racemes and/or developing one terminal flower. Pistillate flowers: 3- (or 4)-locular. Fruits opening loculicidally-septicidally or septicidally-loculicidally, inside glabrous to densely hairy.

    Distribution — From Burma to South China throughout Southeast Asia and Maleaia to New Guinea.

    Note — The other subgenus (subgen. Cordemoya (Baill.) S.E.C.Sierra, Kulju & Welzen) occurs in Madagascar, Mauritius and Réunion.

Section Hancea

    Hancea Seem. section Hancea: S.E.C.Sierra, Kulju, Veldkamp & Welzen, Blumea 52 (2007) 363. — Mallotus sect. Hancea (Seem.) Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 199; Airy Shaw, Kew Bull. 20 (1966) 39; Kew Bull. 21 (1968) 388; Kew Bull. 23 (1969) 80; Kew Bull. 26 (1972) 293; Kew Bull. 27 (1972) 86; Kew Bull. Add. Ser. 4 (1975) 160; Kew Bull. Add. Ser. 8 (1980) 164; Kew Bull. 36 (1981) 324; Slik & Welzen, Blumea 46 (2001) 4; in Slik, Tropenbos-Kalimantan ser. 4 (2001) 146. — Type species as genus: Hancea hookeriana Seem.

    Diplochlamys Müll.Arg., Flora 47 (1864) 539. — Mallotus sect. Diplochlamys (Müll.Arg.) Baill. ex Durand, Index Gen. Phan. (1888) 370; Pax in Engl., Prantl, Pflanzenfam. III.5 (1890) 55, isonym; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 177; Airy Shaw, Kew Bull. 21 (1968) 389. — Cordemoya subgen. Diplochlamys (Müll.Arg.) S.E.C.Sierra, Kulju & Welzen, Blumea 51 (2006) 532. — Cordemoya sect. Diplochlamys (Müll.Arg.) S.E.C.Sierra, Kulju & Welzen, Blumea 51 (2006) 533. — Type: Diplochlamys griffithianus Müll.Arg. [ = Hancea griffithiana (Müll.Arg.) S.E.C.Sierra, Kulju & Welzen].

Branches with conspicuously swollen nodes. Stipules interpetiolar, margin entire. Leaves opposite, strongly unequal, one of each pair stipuliform; blade not peltate, margin entire (rarely serrate), venation pinnate (rarely palminerved), looping close to the margin (rarely ending in the margin). Inflorescences axillary; staminate ones racemes or panicles; pistillate ones racemes and/or developing single flowers terminally. Staminate flowers: sepals (2 or) 3 or 4, free to basally connate. Pistillate flowers: sepals (3 or) 4–6; ovary 3- (or 4-)locular. Fruits opening septicidally-loculicidally, inside glabrous to hairy, spines with or without terminal glands.

    Distribution — From Burma to South China throughout Southeast Asia and Maleaia to New Guinea.

    Habitat & Ecology — Usually in the understorey of primary forest, sometimes also present in secondary forest (mainly selectively logged) in shaded places.

Key to the Malesian species (of subgenus Hancea)

1. Hancea cordatifolia (Slik) S.E.C. Sierra, Kulju & Welzen

    Hancea cordatifolia (Slik) S.E.C.Sierra, Kulju & Welzen in S.E.C.Sierra, Kulju, Veldkamp & Welzen, Blumea 52 (2007) 363. — Mallotus cordatifolius Slik, Blumea 43 (1998) 225; Slik & Welzen, Blumea 46 (2001) 7, Fig. 1, Map 1; in Slik, Tropenbos-Kalimantan ser. 4 (2001) 149, Fig. 7.1, Map 7.1. — Type: PNH (Gutierrez et al.) 117545 (L holo), Philippines, Samar Island, Mt Sohoton.

Small tree up to 7 m tall, dbh up to 8 cm. Indumentum dense, yellowish, short to long. Branches smooth, glabrescent. Stipules persistent, narrowly triangular, 7–10 by 1–1.1 mm, margin entire, apex acute, hairy. Non-reduced leaves: petioles 5–7 by c. 1 mm, hairy, leaf blade ovate, 6.5–16.5 by 3.5–8.5 cm, length-width ratio 2–2.5, chartaceous, base deeply cordate, surrounding the petiole, slightly oblique, margin irregularly wavy, apex acute to acuminate, upper surface smooth, evenly hairy, lower surface smooth, evenly hairy, domatia absent; venation palmate, veins 6 at petiole attachment, 9–12 secondary veins per side along midrib, ending parallel to the margin. Stipuliform leaves (narrowly) triangular, 2–2.5 by c. 0.5 mm, hairy, margin entire, apex acute to rounded. Staminate inflorescences up to 3.5 cm long, basally c. 0.8 mm thick, unbranched, hairy, nodes up to 31 per inflorescence; bracts narrowly triangular, at right angles with rachis, 4.3–5 by c. 0.7 mm, margin entire, apex acute, hairy. Buds globose with acuminate apex, hairy. Staminate flowers 4.5–5 mm in diameter, red; bracteoles absent; pedicels 3.5–4.5 mm long, hairy; sepals 3, persistent (to caducous), basally connate, ovate, 3–3.8 by 1.8–2 mm, margin irregular, apex acuminate, hairy abaxially, glabrous adaxially; stamens up to 100, filaments up to 4 mm long, glabrous, anthers c. 0.5 by 0.5 mm, basifixed; connective not widened. Pistillate inflorescences and fruits unknown.

    Distribution — Philippines (Mt Sohoton on Samar Island).

    Habitat & Ecology — Logged-over forest.

2. Hancea eucausta (Airy Shaw) S.E.C. Sierra, Kulju & Welzen

    Hancea eucausta (Airy Shaw) S.E.C.Sierra, Kulju & Welzen in S.E.C.Sierra, Kulju, Veldkamp & Welzen, Blumea 52 (2007) 363. — Mallotus eucaustus Airy Shaw, Kew Bull. 23 (1969) 80; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 161; Slik, Priyono & Welzen, Gard. Bull. Singapore 52 (2000) 56, Fig. 26; Slik & Welzen, Blumea 46 (2001) 9, Fig. 2, Map 1; Slik, Priyono & Welzen in Slik, Tropenbos-Kalimantan ser. 4 (2001) 95, Fig. 5.29; Slik & Welzen in Slik, Tropenbos-Kalimantan ser. 4 (2001) 150, Fig. 7.2, Map 7.1. — Type: S (Sibat ak Luang) 23630 (K holo; L iso), Borneo, Sarawak, Third Division, Bukit Iju, Ulu Arip, Balingian.

    Also check: Slik, Macaranga and Mallotus species of Borneo (under Mallotus eucaustus).

Shrubs to small trees up to 23 m tall, dbh up to 20 cm; bole straight. Outer bark smooth, hoop-marked, rugose, mottled; sapwood hard. Indumentum dense, whitish to yellowish, short. Branches smooth, glabrescent. Stipules caducous to semi-persistent, triangular to narrowly triangular, 4–11 by 1–1.8 mm, margin entire, apex acute, hairy, puncticulate to clavate glands few to absent. Leaves in one plane. Non-reduced leaves: petiole 3–11 by 1–2 mm, hairy, clavate glands few to numerous; blade sub-ovate to elliptic (to obovate), 7–24.5 by 1.5–7 cm, length-width ratio 2.7–5, coriaceous, base cuneate to acute to obtuse, slightly oblique, margin entire to slightly wavy to apically dentate, apex acuminate to aristate, upper surface smooth, hairy on the midrib, glabrescent, puncticulate glands few, inconspicuous, lower surface smooth, hairy, especially on veins, puncticulate to clavate glands numerous, inconspicuous, domatia absent; venation pinnate, 11–17 secondary veins per side, ending parallel to the margin. Stipuliform leaves narrowly triangular, 4–8.3 by 1.5 mm, hairy, margin entire, apex acute. Staminate inflorescences up to 3.5 cm long, basally 0.3–0.8 mm thick, unbranched, hairy, puncticulate to clavate glands present, nodes up to 16 per inflorescence; bracts narrowly triangular, 2.5–5.5 by 0.6–0.8 mm, margin entire, apex acute, hairy abaxially, glabrous (to few hairs) inside, puncticulate glands present. Buds globose with acute apex, hairy. Staminate flowers c. 3.5 mm in diameter, reddish yellow to red; bracteoles absent; pedicels 1–4 mm long, hairy; sepals 3, persistent, basally connate to free, elliptic to orbicular, c. 3 by 2.5 mm, margin entire to irregular, apex acute, hairy abaxially, glabrous adaxially; stamens up to 46, filaments up to 1.5 mm long, glabrous; anthers c. 0.5 by 0.6 mm, basifixed; connective more or less widened. Pistillate inflorescences up to 7 cm long, basally 0.9–1 mm thick, unbranched, hairy, puncticulate glands few, inconspicuous, nodes up to 11 per inflorescence; bracts persistent, narrowly ovate to narrowly triangular, 1.8–4 by 0.6–0.8 mm, margin entire, apex acute, hairy abaxially, glabrous to hairy adaxially, puncticulate glands few, inconspicuous. Pistillate flowers: bracteoles absent; pedicels 2.5–5 mm long, hairy, puncticulate glands few to absent; sepals 5–6, free, narrowly triangular, 2.5–4 by c. 0.6 mm, margin entire, apex acute, hairy abaxially, glabrous to hairy adaxially, puncticulate to clavate glands few, inconspicuous; ovary 3(–4)-locular, echinate, densely woolly hairy, spines 0.5–1 mm long, with long hairs; style 0.5–1 mm long, hairy; stigmas 7–10 mm long, densely covered with plumose to finely granulate papillae on the inside, hairy abaxially, clavate glands absent or present. Fruit a shortly echinate, lobed capsule, 8–15 by 5–8 mm, green to light red, hairy; carpels glabrous to hairy adaxially; column c. 4 by 3.5 mm. Seeds glossy, globose, c. 4 by 4 by 5 mm; hilum elliptic to slightly deltoid, c. 1 by 0.5 mm.

    Distribution — Borneo.

    Habitat & Ecology — Mainly a locally common understorey tree in primary forest, but also in secondary, logged-over forests; mostly on well-drained, sandy to (red) clayey soils. Altitude: up to 340 m.

    Uses — Fire-wood (hence its name eucaustus, which means easy to burn).

3. Hancea grandistipularis (Slik) S.E.C. Sierra, Kulju & Welzen

    Hancea grandistipularis (Slik) S.E.C.Sierra, Kulju & Welzen in S.E.C.Sierra, Kulju, Veldkamp & Welzen, Blumea 52 (2007) 363. — Mallotus grandistipularis Slik, Blumea 43 (1998) 227; in Slik & Welzen, Blumea 46 (2001) 11, Fig. 3, Map 1; in Slik, Tropenbos-Kalimantan ser. 4 (2001) 153, Fig. 7.3, Map 7.1. — Type: Burley, Tukirin et al. 2021 (L holo; CANB, K, US iso), Indonesia, Sumatra, Riau Province, Tigapulu Mts, 5 km West of Talanglakat.

Small trees up to 15 m tall, dbh up to 15 cm. Outer bark smooth, hoop-marked. Indumentum dense, short to long. Branches lenticellate, glabrescent, tubular to clavate glands few to numerous when young. Stipules semi-persistent, ovate to obovate, 8–22 by 2.3–9 mm, margin irregular wavy to entire, apex (rounded to) acute (to acuminate), hairy, especially along margin. Leaves in one plane. Non-reduced leaves: petioles 9–45 by 1.5–2 mm, hairy, tubular to clavate glands few to numerous when young; blade ovate to obovate, 13.5–30.5 by 4–9.5 cm, length-width ratio 3–4, coriaceous, base obtuse to rounded, margin entire to slightly wavy, apically sometimes slightly dentate, apex acuminate to caudate, upper surface smooth, glabrous to few hairs on midrib and veins, lower surface smooth, hairy, tubular to clavate to puncticulate glands numerous when young; venation pinnate, 10–13(–15) secondary veins per side, ending parallel to the margin. Stipuliform leaves obovate, 4–10 by 1.8–3.5 mm, margin entire to wavy, apex acute, hairy, especially along margin, tubular to clavate glands numerous when young. Staminate inflorescences unknown. Pistillate inflorescences up to 10 cm long, basally c. 1.8 mm thick, unbranched, hairy, tubular to clavate glands few to numerous, nodes up to 9 per inflorescence; bracts ovate to obovate, 3–4 by 1.3–1.4 mm, margin entire to irregular, apex acute, hairy, tubular to clavate glands on the outside only. Pistillate flowers: pedicels 5–6 mm long, hairy, tubular to clavate glands numerous; sepals 4–5, persistent, free, ovate to narrowly triangular, 5.5–6 by c. 1.1 mm, margin entire, apex acute, hairy, tubular to clavate glands present on outside only; ovary 3-locular, echinate, densely hairy, glands absent, spines 2–2.5 mm long, ending in an inconspicuous gland, apically few long hairs; styles c. 1.8 mm long, hairy; stigmas up to 9.5 mm long, densely covered with granulate papillae on the inside, hairy abaxially. Fruit an echinate, lobed capsule, c. 18 by 15 mm, greyish-red, hairy; carpels glabrous to sparsely hairy adaxially, puncticulate glands absent to numerous; column c. 7.5 by 3.8 mm. Seeds glossy, globose, 4.5–5 by 3.5–4.5 by 4–5 mm; hilum elliptic, 1–1.2 by 0.5–1 mm.

    Distribution — Central Sumatra.

    Habitat & Ecology — In primary lowland Dipterocarp forest; on loamy soils. Altitude: up to 500 m.

4. Hancea griffithiana (Müll.Arg.) S.E.C. Sierra, Kulju & Welzen

    Hancea griffithiana (Müll.Arg.) S.E.C.Sierra, Kulju & Welzen in S.E.C.Sierra, Kulju, Veldkamp & Welzen, Blumea 52 (2007) 363. — Diplochlamys griffithianus Müll.Arg., Flora 47 (1864) 539; in DC., Prod. 15, 2 (1866) 1024. — Mallotus griffithianus (Müll.Arg.) Hook.f., Fl. Brit. India 5 (1887) 433; Boerl., Handl. Fl. Ned. Ind. 3(1) (1900) 290; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 178; Ridl., Fl. Malay Penins. 3 (1924) 291; Pax & K.Hoffm. in Engl. & Harms, Natürl. Pflanzenfam. ed. 2, 19c (1931) 115; Burkill, Dict. Econ. Prod. Malay Penins. 2 (1935) 1395; Meijer, Bot. News Bull. (1967) 52; Airy Shaw, Kew Bull. 21 (1968) 390; Whitmore, Tree Fl. Malaya 2 (1973) 116; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 163; Slik, Priyono & Welzen, Gard. Bull. Singapore 52 (2000)57, Fig. 28; Slik & Welzen, Blumea 46 (2001) 12, Fig. 4, Map 2; Slik, Priyono & Welzen in Slik, Tropenbos-Kalimantan ser. 4 (2001) 95, Fig. 5.31; Slik & Welzen in Slik, Tropenbos-Kalimantan ser. 4 (2001) 155, Fig. 7.4, Map 7.2. — Type: Griffith KD 4961 (K holo), India orientali Malayensi.

    Buettneria uncinata Mast. in Hook.f., Fl. Brit. India 1 (1884) 377. — Type: Maingay KD 1450 (K holo), Malacca.

    Mallotus impar Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 396; Merr., J. Str. Roy. Asiatic. Soc. (1921) 339; Univ. Calif. Publ. Bot. 15 (1929) 157; Pax & K.Hoffm. in Engl. & Harms, Natürl. Pflanzenfam. ed. 2, 19c (1931) 117. — Type: Hose 281 (B holo, lost; EDI, K, L iso), Südwestmalayische Provinz, Borneo, Sarawak, Baram Distrikt.

    Mallotus woodii Merr., Philipp. J. Sci. (1918) 81; J. Str. Roy. Asiatic Soc. (1921) 340; Univ. Calif. Publ. Bot. 15 (1929) 158; Pax & K.Hoffm. in Engl. & Harms, Natürl. Pflanzenfam. ed. 2, 19c (1931) 117. — Type: Wood 451 (PNH holo, lost; K iso), Sabah, Marutai, near Tawau.

    Also check: Slik, Macaranga and Mallotus species of Borneo under Mallotus griffithianus.

Small trees up to 20 m tall, dbh up to 21 cm; bole straight. Outer bark smooth to slightly fissured, hoop-marked, dippled; sapwood soft to medium hard. Indumentum sparse, short to long. Branches smooth, sparsely (to densely) lenticellate, glabrous to sparsely hairy, glabrescent, clavate glands few to numerous, reddish, on young parts. Stipules early caducous, narrowly triangular (to ovate), 4.5–6.5 by 1.3 mm, margin entire, apex acute, sparsely hairy abaxially, glabrous adaxially, puncticulate glands few to numerous. Non-reduced leaves: petiole 28–90 by 0.9–2.3 mm, apically strongly pulvinate and bending upwards, glabrous to sparsely hairy, glabrescent, clavate glands present when young, puncticulate when older; blade ovate to elliptic (to slightly obovate), 9–30 by 2.5–18 cm, length-width ratio 2–4.3, coriaceous, base obtuse to acute to rounded, sometimes slightly oblique, margin entire to slightly wavy to apically sometimes dentate, apex acuminate to cuspidate, upper surface smooth, glabrous to sparsely hairy on midrib, puncticulate glands few to numerous, lower surface smooth, sparsely hairy on midrib, puncticulate glands few to numerous, domatia absent; venation pinnate, 11–17 secondary veins per side, ending parallel to the margin. Stipuliform leaves narrowly triangular, 2.5–4 by 1–1.5 mm, hairy, clavate glanded, margin entire, apex acute. Staminate inflorescences up to 13 cm long, basally 1–1.5 mm thick, branched, each branch with basal bract, hairy, clavate glands numerous, nodes many; bracts persistent, triangular to ovate, at right angles with the axis, 1.5–1.6 by 0.5–0.6 mm, margin entire, apex acute, sparsely hairy on outside only, clavate glands numerous on outside only. Buds globose with acuminate apex, hairy, clavate glands numerous. Staminate flowers 4.5–7 mm in diameter, white to reddish yellow to reddish; bracteoles absent; pedicels 1.3–5 mm long, hairy, clavate glands numerous; sepals 3–4, persistent, basally connate, ovate, 1.7–3.5 by 1.1–2 mm, margin entire to slightly irregular, apex acute, hairy on outside only, clavate glands numerous, on outside only; stamens up to 86, filaments 1.3–1.7 mm long; anthers 0.3–0.5 mm high, basifixed; connective not to slightly widened, sometimes slightly overtopping anther. Pistillate inflorescenses up to 12 cm long, basally 1–3 mm thick, rarely branched, hairy, glabrescent, clavate glands numerous, with sticky yellow exudate, nodes up to 14 per inflorescence; bracts persistent, ovate to narrowly triangular, 2–4 by c. 1 mm, margin entire, apex acuminate to cuspidate, hairy on outside only, clavate glands numerous. Pistillate flowers: bracteoles early caducous to sometimes subtending the flower, ovate, 1.3–1.8 by 0.4–0.8 mm, margin entire, apex acute, hairy on outside only, clavate to conical glands numerous; pedicels 2.3–27 mm long, hairy, apically with long hairs, clavate to conical glands numerous; sepals 5, persistent, free to basally connate, ovate to narrowly triangular, 4–8 by 0.9–2.3 mm, margin entire, apex acute, hairy on outside only, clavate glands on outside only; ovary 3-locular, echinate, densely hairy, clavate glands numerous, spines 2–4 mm long, apically often with globose gland, long hairs apically, clavate glands present; style 1–2 mm long, hairy, clavate glands present; stigmas 9–15 mm long, densely covered with granulate papillae on the inside, hairy on the outside, clavate glands present on the outside. Fruit an echinate, lobed capsule, 12–18 by 8–12 mm, yellow when young, pink to red to red-brown when mature, (glabrous to) hairy, clavate glands absent to few; carpel inside glabrous to hairy, clavate glands absent to few; column 7–8 by 5–6 mm. Seeds glossy, globose, c. 6 by 5 by 5 mm; hilum elliptic to ovate to triangular, 1–1.5 by c. 1 mm.

    Distribution — From Peninsular Thailand and Malaysia into Borneo.

    Habitat & Ecology — Mainly a locally common understorey tree in primary forest, but also in secondary logged-over forests; on steep slopes, ridges, hill sides, flat land, along paths, river banks, sometimes on periodically inundated terrain; present on most soil types from peat to sandstone to (yellow) sandy loam to ultrabasic soil to yellow sandy clay to rarely on limestone. Altitude: up to 480 m.

    Note — A very constant species with little variation in characters.

5. Hancea hirsuta (Elmer) S.E.C. Sierra, Kulju & Welzen

    Hancea hirsuta (Elmer) S.E.C.Sierra, Kulju & Welzen in S.E.C.Sierra, Kulju, Veldkamp & Welzen, Blumea 52 (2007) 364. — Mallotus hirsutus Elmer, Leafl. Philipp. Bot. (1915) 2648; Airy Shaw, Alph. Enum. Euphorb. Philipp. Isl. (1983) 36; Slik & Welzen, Blumea 46 (2001) 15, Fig. 5, Map 2; in Slik, Tropenbos-Kalimantan ser. 4 (2001) 157, Fig. 7.5, Map 7.2. — Type: Elmer 13480 (PNH holo, lost; K, L, NY iso), The Philippines, Mindanao, Agusan province, Cabadbaran (Mt Urdaneta).

Small trees up to 8 m tall, dbh up to 15 cm. Indumentum dense, short to long. Branches smooth to slightly fissured, glabrescent, clavate glands absent to few. Stipules caducous to semi-persistent, narrowly triangular, 9–14 by 1.5–1.8 mm, margin entire, slightly incurved, apex acute, hairy, clavate glands absent to few outside. Non-reduced leaves: petiole 5–16 by 1–1.2 mm, hairy, clavate glands absent to few; blade ovate to elliptic (to slightly obovate), 8.5–19 by 3.5–6 mm, length-width ratio 2.2–3.3, chartaceous to coriaceous, base (rounded) to obtuse, slightly oblique, margin entire to wavy, apex acuminate to cuspidate, upper surface smooth, hairy (to glabrous), especially on midrib and nerves, lower surface smooth, hairy, domatia absent; venation pinnate, 7–9 secondary veins per side, ending parallel to the margin. Stipuliform leaves narrowly triangular to ovate, 5–8 by 1.5–1.8 mm, margin entire, apex acute, glabrous to hairy abaxially, glabrous adaxially. Staminate inflorescences unknown. Pistillate inflorescences up to 10.5 cm long, basally c. 1 mm thick, unbranched, hairy, clavate glands absent to few, nodes up to 5 per inflorescence, usually only terminal node with fruit; bracts persistent, narrowly triangular, 9–12 by 1–1.3 mm, margin entire, apex acute, hairy, glabrous adaxially, clavate glands absent to few. Pistillate flowers: bracteoles absent; pedicels 5–7 mm long, hairy, clavate glands absent to few; sepals 5, persistent, free, narrowly triangular, 7–12 by 0.9–1.4 mm, margin entire, apex acute, hairy, glabrous adaxially; ovary 3-locular, echinate, densely hairy, spines 2–5 mm long, hairy; style c. 1 mm long, hairy; stigmas 10–16 mm long, densely covered with granulate papillae on the inside, hairy on the outside. Fruit an echinate, lobed capsule, 10–15 by 9–10 mm, hairy in- and outside; column c. 4.5 by 3.5 mm. Seeds glossy, globose, c. 4 by 4.5 by 5 mm; hilum elliptic to deltoid, c. 1.5 by 0.6 mm.

    Distribution — Mindanao in the Philippines.

    Habitat & Ecology — In forest. Altitude: 1333 m.

6. Hancea kingii (Hook.f.) S.E.C. Sierra, Kulju & Welzen

    Hancea kingii (Hook.f.) S.E.C.Sierra, Kulju & Welzen in S.E.C.Sierra, Kulju, Veldkamp & Welzen, Blumea 52 (2007) 364. — Mallotus kingii Hook.f., Fl. Brit. India 5 (1887) 439; Boerl., Handl. Fl. Ned. Ind. 3(1) (1900) 290; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 197; Ridl., Fl. Malay Penins. 3 (1924) 293; Pax & K.Hoffm. in Engl. & Harms, Natürl. Pflanzenfam. ed. 2, 19c (1931) 117; Airy Shaw, Kew Bull. 21 (1968) 389; Kew Bull. 23 (1969) 81; Kew Bull. 26 (1972) 295; Whitmore, Tree Fl. Malaya 2 (1973) 115; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 163; Welzen, Slik & Bollendorff in Welzen et al., Thai For. Bull. (Bot.) 28 (2000) 101; Slik & Welzen, Blumea 46 (2001) 18, Fig. 7, Map 1; in Slik, Tropenbos-Kalimantan ser. 4 (2001) 161, Fig. 7.7, Map 7.1; Welzen, S.E.C.Sierra, Slik & Bollendorff in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 402. — Type: King's Collector 7414 (K holo; G iso), Malaya, Perak.

Small trees up to 25 m tall, dbh up to 21 cm; bole straight. Outer bark smooth, slightly lenticellate, brittle, scaly; inner bark hard, granular. Indumentum dense, short. Branches smooth, glabrescent, clavate to puncticulate glands few to numerous. Stipules caducous to persistent, ovate, 8–14 by 2–3 mm, margin entire, apex acute to aristate, hairy with few long hairs along margin, clavate glands absent or present. Non-reduced leaves: petiole 25–62 by 0.8–2 mm, hairy, puncticulate to clavate glands few to numerous; blade elliptic to obovate, 11–31.5 by 4–11.5 cm, length-width ratio 2.2–3, chartaceous, base rounded to obtuse to cuneate, sometimes slightly oblique, margin slightly wavy, apex acuminate to cuspidate, upper surface smooth, hairy on midrib and base of veins, puncticulate to clavate glands few to numerous, lower surface smooth, hairy, especially on veins, clavate to puncticulate glands usually numerous, domatia absent; venation pinnate, 12–14 secondary veins per side, ending parallel to the margin, apically sometimes ending in margin. Stipuliform leaves ovate, 4–7 by 2 mm, margin entire, apex acute, hairy, clavate glands absent or present. Staminate inflorescences up to 9 cm long, basally c. 1.8 mm thick, branched, hairy, clavate glands numerous, nodes many per inflorescence; bracts persistent, ovate to obovate, 1–1.5 by 0.4–0.5 mm, margin entire to irregular, apex rounded, hairy, clavate glands numerous on outside only. Buds globose, hairy, clavate glands numerous. Staminate flowers 3–5 mm in diameter, greenish white; bracteoles absent; pedicels 1–2 mm long, hairy, clavate glands numerous; sepals 2–3, persistent, basally connate, orbicular to elliptic, c. 3 by 2–3 mm, margin entire, apex acute, hairy abaxially, glabrous adaxially, clavate glands numerous on outside only; stamens up to 98, filaments 3.5–4 mm long, glabrous; anthers c. 0.5 by 0.6 mm, dorsifixed; connective widened or not. Pistillate inflorescences up to 15 cm long, basally c. 2 mm thick, unbranched (to branched), hairy, clavate glands numerous, nodes up to 15 per inflorescence; bracts persistent, ovate to deltoid, 1.7–2.5 by 1.5–2 mm, margin entire, apex acute, hairy with long hairs along margin, clavate glands numerous. Pistillate flowers green; bracteoles usually absent, subtending the calyx, ovate, 1.8–4 by 0.7–1.5 mm, margin entire, apex rounded, glabrous to hairy with long hairs along margin, clavate glands numerous; pedicels 4–19 mm long, hairy, clavate glands numerous; sepals 5, persistent, free, deltoid to ovate, 1.8–4.5 by 1.5 mm, margin entire, apex acute, glabrous to sparsely hairy with long hairs along margin, clavate glands numerous; ovary 3-locular, echinate, densely woolly hairy, clavate glands absent to few, spines 2–7 mm long, usually with apical gland, glabrous to hairy, sometimes long hairs apically, clavate glands absent to few basally; style 1–1.5 mm long, hairy, clavate glands present; stigmas 6–7 mm long, densely covered with granulate papillae on the inside, hairy on the outside, clavate glands numerous on the outside. Fruit an echinate, lobed capsule, 15–16 by 12–13 mm, green, hairy, clavate glands absent to few; carpel inside glabrous to hairy, puncticulate to clavate glands few; column 7–8 by 5 mm. Seeds glossy, globose, 5.5–6 by 4.5–6 by 5.5–6 mm; hilum ellipic to deltoid, 1.5–2 by 1 mm.

    Distribution — Peninsular Thailand and Peninsular Malaysia.

    Habitat & Ecology — Primary and secondary forests; on ridges, hill sides, low undulating terrain, and along streams. Altitude: up to 833 m.

    Note — A fairly constant species with little variation in characters.

7. Hancea longystyla (Merr.) S.E.C. Sierra, Kulju & Welzen

    Hancea longistyla (Merr.) S.E.C.Sierra, Kulju & Welzen in S.E.C.Sierra, Kulju, Veldkamp & Welzen, Blumea 52 (2007) 364. — Mallotus longistylus Merr., Philipp. J. Sci. 13 (1920) 560; Enum. Philipp. Flow. Pl. 2 (1923) 434, Airy Shaw, Alph. Enum. Euphorb. Philipp. Isl. (1983) 36; Slik & Welzen, Blumea 46 (2001) 20, Fig. 8, Map 2; in Slik, Tropenbos-Kalimantan ser. 4 (2001) 163, Fig. 7.8, Map 7.2. — Type: BS (Ramos & Pascasio) 35173 (PNH holo, lost; K iso), Philippines.

Shrubs to small trees up to 10 m tall, dbh up to 12 cm, sometimes monoecious. Indumentum absent to sparse, short. Branches smooth (to slightly fissured), slightly pustulose, glabrous to sparsely hairy. Stipules persistent, ovate to broadly ovate, 5–27 by 4–17 mm, margin entire to wavy, apex acute to rounded, sparsely hairy, venation parallel, clearly visibLecomte Non-reduced leaves: petiole 6–40 by 0.8–2.2 mm, sparsely hairy; blade ovate, 7.5–26 by 3.5–9.5 cm, length-width ratio 2.3–3.3, (chartaceous to) coriaceous, base obtuse to rounded to slightly cordate, slightly oblique, margin entire to wavy, apex acute to acuminate, upper surface smooth, glabrous to sparsely hairy midrib and veins, lower surface smooth, (pustulose), glabrous to sparsely hairy on midrib and veins, domatia absent; venation pinnate, 8–11 secondary veins per side, ending parallel to the margin. Stipuliform leaves ovate, 1.5–3.3 by 1.2–1.5 mm, margin entire, apex rounded, hairy. Staminate inflorescences up to 9 cm long, basally c. 0.7 mm thick, unbranched, hairy, nodes up to 33 per inflorescence; bracts persistent, broadly ovate to ovate, 5–6 by 4 mm, half to completely enfolding the rachis, margin entire to irregular, apex acute, hairy. Buds globose with acuminate apex, hairy. Staminate flowers c. 7 mm in diameter; bracteoles absent; pedicels 2–4 mm long, hairy; sepals 3(–4), persistent, basally connate, elliptic to obovate, c. 5 by 3.5 mm, margin entire to irregular, apex acuminate, hairy, glabrous adaxially; stamens more than 100, filaments up to 2 mm long, glabrous; anthers 0.6–0.7 by 0.5–0.6 mm, basifixed; connective not widened. Pistillate inflorescences up to 10 cm long, basally 0.5–0.7 mm thick, unbranched, sparsely hairy, nodes up to 10 per inflorescence; bracts persistent, broadly ovate to ovate, enfolding the rachis when young, 5–10 by 1–3 mm, margin entire to irregular, apex acute, hairy. Pistillate flowers: bracteoles absent; pedicels 5–9 mm long, hairy, glands absent to puncticulate; sepals (4)5, persistent, free, narrowly ovate to narrowly elliptic, 6–7 by 1–1.6 mm, margin entire, involute, apex acute, hairy; ovary 3-locular, echinate, densely hairy, glands absent to puncticulate, spines up to 2 mm long, sparsely hairy; style 1.5–2 mm long, densely hairy; stigmas 12–17 mm long, densely covered with granulate papillae on the inside, densely hairy abaxially. Fruit an echinate, lobed capsule, c. 10 by 6.5 mm, green to reddish-green, hairy; carpel inside glabrous to sparsely hairy; column 4–5 by 3–3.5 mm. Seeds glossy, globose, c. 4 by 3.5 by 4 mm; hilum deltoid, c. 1.2 by 1 mm.

    Distribution — Samar Island in the Philippines.

    Habitat & Ecology — In primary forest. Altitude: up to 266 m.

8. Hancea papuana (J.J.Sm.) S.E.C. Sierra, Kulju & Welzen

    Hancea papuana (J.J.Sm.) S.E.C.Sierra, Kulju & Welzen in S.E.C.Sierra, Kulju, Veldkamp & Welzen, Blumea 52 (2007) 364. — Mallotus papuanus (J.J.Sm.) Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 202; Airy Shaw, Kew Bull. 21 (1968) 400; Kew Bull. 23 (1969) 81; Kew Bull. Add. Ser. 8 (1980) 164; Slik & Welzen, Blumea 46 (2001) 22, Fig. 9, Map 4; in Slik, Tropenbos-Kalimantan ser. 4 (2001) 165, Fig. 7.9, Map 7.4. — Mallotus hookerianus (Seem.) Müll.Arg. var. papuanus J.J.Sm., Nova Guinea 8 (1912) 787; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. viii (1919) 19; in Engl. & Harms, Natürl. Pflanzenfam. ed. 2, 19c (1931) 117. — Type: Gjellerup 316 (L holo; A, BO, K, U, P iso), Irian Jaya, Biwak Hollandia.

Small trees up to 15 m tall, dbh up to 10 cm. Outer bark smooth. Indumentum dense, short to long. Branches smooth, (pustulose), glabrescent. Stipules early caducous, narrowly triangular to linear, 10–17 by 1–1.2 mm, margin entire, sometimes involute, apex acute, densely hairy. Non-reduced leaves: petiole 5–12 by 1–2 mm, densely hairy; blade ovate to elliptic to obovate, 6–23 by 2.5–8.5 cm, length-width ratio 2.3–4, coriaceous, base rounded to cordate, conspicuously oblique, margin entire to slightly wavy (to apically dentate), apex acuminate to cuspidate, upper surface smooth, glabrous to hairy, lower surface smooth, hairy, domatia inconspicuous, venation pinnate, 9–15 secondary veins per side, ending parallel to the margin. Stipuliform leaves narrowly triangular to linear, 5–14 by 1 mm, margin entire, apex acute, hairy. Staminate inflorescences up to c. 4.5 cm long, basally c. 0.5 mm thick, unbranched, hairy, nodes up to 28 per inflorescence; bracts ovate to narrowly triangular, 1.8–2.3 by 0.5 mm, margin entire, apex acute to aristate, hairy, glabrous to hairy adaxially. Buds globose with acuminate apex, hairy. Staminate flowers c. 4.5 mm in diameter, pale brownish; bracteoles absent; pedicels 2–2.5 mm long, hairy; sepals 3, more or less connate, elliptic, 1.8–2 by 1.3–1.6 mm, margin entire to irregular, apex acuminate, hairy, glabrous adaxially; stamens up to c. 36, filaments 1.8–2 mm long, glabrous; anthers 0.3–0.5 by 0.6 mm, basifixed; connective slightly widened. Pistillate inflorescences up to 9 cm long, basally c. 0.5–0.8 mm thick, usually only terminal fruit developing, unbranched, hairy, nodes up to 6 per inflorescence; bracts narrowly triangular, 4–7 by 0.4 mm, margin entire, involute, apex acute, hairy. Pistillate flowers yellowish; bracteoles absent; pedicels c. 2 mm long, hairy; sepals 5, free, narrowly triangular, c. 6 by 0.7 mm, margin entire, apex acute, hairy; ovary 3-locular, echinate, hairy, spines 3–7 mm long, with long hairs; styles 0.7–1 mm long, hairy; stigmas up to 15 mm long, densely covered with granulate papillae on the inside, hairy abaxially. Fruit an echinate, lobed capsule, c. 11 by 5 mm, green, hairy; carpel inside hairy; column unknown. Seeds glossy, roughly globose, c. 5 by 4.5 by 5 mm, hilum deltoid, c. 1 by 1 mm.

    Distribution — Only known from the area around Jayapura, Irian Jaya, Indonesia.

    Habitat & Ecology — Primary forest understorey; along streams, in river valleys, on steep slopes; on clay, gravel and rich humus soils. Altitude: up to 650 m.

9. Hancea penangensis (Müll.Arg.) S.E.C. Sierra, Kulju & Welzen

    Hancea penangensis (Müll.Arg.) S.E.C.Sierra, Kulju & Welzen in S.E.C.Sierra, Kulju, Veldkamp & Welzen, Blumea 52 (2007) 364. — Mallotus penangensis Müll.Arg., Linnaea 34 (1865) 186; in DC., Prod. 15, 2 (1866) 961; Hook.f., Fl. Brit. India 5 (1887) 440; Boerl., Handl. Fl. Ned. Ind. 3(1) (1900) 288; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 201; Ridl., Fl. Malay Penins. 3 (1924) 293; Pax & K.Hoffm. in Engl. & Harms, Natürl. Pflanzenfam. ed. 2, 19c (1931) 117; Meijer, in Bot. News Bull. (1967) 52; Airy Shaw, Kew Bull. 21 (1968) 399; Whitmore, Tree Fl. Malaya 2 (1973) 116; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 161, 164; Kew Bull. 36 (1981) 324, 328; Keßler & Sidiyasa, Tropenbos Series 7 (1994) 133; Slik, Priyono & Welzen, Gard. Bull. Singapore 52 (2000) 62, Fig. 39, 40; Welzen, Slik & Bollendorff in Welzen et al., Thai For. Bull. (Bot.) 28 (2000) 105; Slik & Welzen, Blumea 46 (2001) 24, Fig. 10, Map 3; Slik, Priyono & Welzen in Slik, Tropenbos-Kalimantan ser. 4 (2001) 99, Fig. 5.42, 5.43; Slik & Welzen in Slik, Tropenbos-Kalimantan ser. 4 (2001) 167, Fig. 7.10, Map 7.3; Welzen, S.E.C.Sierra, Slik & Bollendorff in elzen & Chayam., Fl. Thailand 8, 2 (2007) 420. 2007. — Type: Wallich 8576 (G holo; A, K, P iso), Malaysia, Pulau Penang.

    Mallotus echinatus Elmer, Leafl. Philipp. Bot. 3 (1910) 925; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 204; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 433; Pax & K.Hoffm. in Engl. & Harms, Natürl. Pflanzenfam. ed. 2 (1931) 117; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 164; Kew Bull. Add. Ser. 8 (1980) 166. — Type: Elmer 12253 (PNH holo, lost; EDI, K, L, NY iso), The Philippines, Sibuyan Island, Capiz Province, Magallanes (Mt Giting-giting).

    Mallotus sarawakensis Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 201; Merr., J. Str. Roy. Asiatic Soc. (1921) 340; Pax & K.Hoffm. in Engl. & Harms, Natürl. Pflanzenfam. ed. 2, 19c (1931) 117. — Type: Hose 184 (B holo, lost; A, EDI, K, L iso), Borneo, Sarawak, Baram District.

    Mallotus leptophyllus Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 203; Merr., J. Str. Roy. Asiatic Soc. (1921) 339; Pax & K.Hoffm. in Engl. & Harms, Natürl. Pflanzenfam. ed. 2, 19c (1931) 117. — Type: Hose 26 (B holo, lost; L iso), Borneo, Sarawak, Baram District.

    Mallotus pseudopenangensis Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 203; in Engl. & Harms, Natürl. Pflanzenfam. ed. 2, 19c (1931) 117. — Lectotype, selected here: BS (Ramos) 7396 (US lecto; NY isolecto), Philippines, Luzon, Cagayan.

    Mallotus xylacanthus Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 203, 397; S.Moore, J. Bot. 63 (1925) 103; Pax & K.Hoffm. in Engl. & Harms, Natürl. Pflanzenfam. ed. 2 (1931) 117. — Lectotype (Slik & Welzen, Blumea 46, 2001): Forbes 3237 (L holo), Sumatra.

    Mallotus papuanus (J.J.Sm.) Pax & K.Hoffm. var. glabrescens Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. viii (1919) 19. — Lectotype (Slik & Welzen, Blumea 46, 2001): Ledermann 9803 (L holo), New Guinea, Aprilfluss.

    Mallotus papuanus (J.J.Sm.) Pax & K.Hoffm. var. intermedius Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. viii (1919) 19. — Type: Ledermann 9539 (B holo, lost; K, L iso), New Guinea, Etappenberg.

    Mallotus penangensis Müll.Arg. var. grandifolia Ridl., Fl. Malay Penins. 3 (1924) 293. --- Lectotype (Slik & Welzen, Blumea 46, 2001): Ridley 6481 (SING holo), Singapore, Sungei Morai.

    Mallotus tenuipes Airy Shaw, Kew Bull. Add. Ser. 8 (1980) 393. — Type: S (Wright & Othman Ismawi) 32221 (K holo; L iso), Sarawak.

    Also check: Slik, Macaranga and Mallotus species of Borneo under Mallotus penangensis.

Small trees up to 27 m tall, dbh up to 40 cm; bole straight, fluted. Outer bark smooth, hoop-marked, slightly fissured; sapwood quite hard. Indumentum sparse, short to long. Branches smooth to rarely fissured when older, glabrescent, clavate glands absent to sometimes few at nodes. Stipules caducous (to semi-persistent), narrowly triangular to ovate, 2–10 by 1–3 mm, margin entire, apex acute to sometimes rounded, glabrous to hairy abaxially, glabrous adaxially, clavate to puncticulate glands absent or present. Non-reduced leaves: petiole 4–45 by 0.8–1.5 mm, sparsely hairy, glabrescent, clavate to puncticulate glands absent to sometimes few; blade ovate to obovate, 5–25.5 by 2–11 cm, length-width ratio 1.7–5.1, coriaceous, base acute (to obtuse to rounded), rarely oblique, margin entire to slightly wavy, apex acuminate to cuspidate, upper surface smooth, glabrous to sparsely hairy on veins, clavate to puncticulate glands absent to few, lower surface smooth, glabrous to sparsely hairy, especially on midrib, domatia absent or present, with simple hair tufts; venation pinnate, 7–14 secondary veins per side, ending parallel to the margin. Stipuliform leaves narrowly triangular to ovate, 1–4.7 by 1.3–2 mm, margin entire to irregular, apex acute to rounded, hairy abaxially only, clavate to puncticulate glands absent to few. Staminate inflorescences up to 7.5 cm long, basally 0.4–1 mm thick, unbranched, hairy, clavate to puncticulate glands absent to few, nodes up to 54 per inflorescence; bracts triangular to ovate to deltoid, 0.6–2.5 by 0.5–2.5 mm, margin entire to rarely irregular, apex acute, hairy abaxially only, clavate to puncticulate glands absent. Buds globose with acuminate apex, hairy, clavate to puncticulate glands absent to numerous. Staminate flowers 3.7–5.5 mm in diameter, greenish-white to pinkish-white to red; bracteoles absent to rarely present, narrowly triangular, c. 2 by 0.4 mm, margin entire, apex acute, hairy abaxially only, clavate to puncticulate glands absent to few; pedicels 2–3 mm long, hairy, clavate to puncticulate glands absent or present; sepals 3–4, free to basally connate, broadly ovate to elliptical, often recurved, 2–5 by 2–2.5 mm, margin entire to irregular, apex acute to acuminate, hairy abaxially only, clavate to puncticulate glands absent to numerous; stamens up to c. 98, filaments 3–4 mm long, glabrous to rarely hairy, clavate glands absent; anthers 0.6–0.7 by 0.5–0.7 mm, basifixed to dorsifixed; connective widened, sometimes apically protruding. Pistillate inflorescences up to 12 cm long, basally 1–1.5 mm thick, usually only terminal fruit developing, unbranched, hairy, clavate glands absent to numerous, nodes up to 18 per inflorescence; bracts persistent, broadly ovate to ovate, 0.9–2 by 0.8–1.1 mm, margin entire, apex acute to acuminate, hairy abaxially only, clavate glands absent or present. Pistillate flowers: bracteoles absent; pedicels up to 10 mm long, hairy, clavate to puncticulate glands absent or present; sepals 4–5, usually free to sometimes basally connate, narrowly triangular, often recurved, 2–7 by 0.5–1.2 mm, margin entire, apex acute, hairy abaxially only, clavate glands absent to numerous; ovary 3-locular, echinate, densely woolly hairy, clavate glands absent or present, spines up to 5 mm long, hairy, apically usually few long hairs, clavate glands absent to few; style 0.7–1 mm long, hairy, clavate glands absent; stigmas 5–16 mm long, densely covered with granulate papillae on the inside, hairy abaxially, clavate glands absent to few. Fruit an echinate, lobed capsule, 9–15 by 6–14 mm, green to pinkish to red; carpel inside simple to stellate hairy, puncticulate glands absent or present; column 3.5–7 mm long, 3.4–5 mm wide. Seeds glossy, roughly globose, 4.5–7 by 3–6.5 by 2.5–8 mm; hilum elliptic to orbicular, 0.5–1.7 by 0.5–1.5 mm.

    Distribution — From Southern Thailand and the Andaman Islands to the Philippines and New Guinea.

    Habitat & Ecology — Mainly in the understorey of primary forests or late secondary forests, where it can be very common, sometimes also in young secondary forest or scrub, along creeks, river banks and road sides; usually on dry, well drained terrain, occasional also swampy places; soils variable: from clay to sandy to loamy, also on limestone. Altitude: up to 1666 m.

    Uses — Leaves used against "sakit kepala" (headache) in Sumatra. The wood has a faint sweet aroma and is used for fires and for house construction (Malaysia).

    Notes — A very variable species, hence the large number of synonyms. The petioles range from very short (c. 0.5 cm) to very long (c. 4.5 cm); the stipules can be very small (c. 2 mm long) to quite large (c. 1 cm long); domatia can be present or absent; leaf sizes range from small (c. 5–7 cm long) to very long (up to c. 27 cm); the pistillate inflorecences can be slender with only 1 terminal fruit developing, or quite thick and with many fruits developing over the whole inflorescence; the fruits can have long spines (whole distribution range) to very short to almost verrucose spines (mainly the Moluccas and New Guinea). As a rule (but not always) the plants become more hairy, the stipules become longer, domatia are more often present, and the inflorescence axes become thicker with more fruits with increasing altitude.

10. Hancea stipularis (Meijer ex Airy Shaw) S.E.C. Sierra, Kulju & Welzen

       Hancea stipularis (Meijer ex Airy Shaw) S.E.C.Sierra, Kulju & Welzen in S.E.C.Sierra, Kulju, Veldkamp & Welzen, Blumea 52 (2007) 364. —  Mallotus stipularis Meijer [Bot. News Bull. 7 (1967) 53, anglice] ex Airy Shaw, Kew Bull. 21 (1968) 398; Meijer, Bot. News Bull. 10 (1968) 231; Kew Bull. 26 (1972) 169; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 164; Kew Bull. 36 (1981) 329; Slik, Priyono & Welzen, Gard. Bull. Singapore 52 (2000) 63, Fig. 41; Welzen, Slik & Bollendorff in Welzen et al., Thai For. Bull. (Bot.) 28 (2000) 109; Slik & Welzen, Blumea 46 (2001) 27, Fig. 11, Map 4; Slik, Priyono & Welzen in Slik, Tropenbos-Kalimantan ser. 4 (2001) 99, Fig. 5.45; Slik & Welzen in Slik, Tropenbos-Kalimantan ser. 4 (2001) 170, Fig. 7.11, Map 7.4; Welzen, S.E.C.Sierra, Slik & Bollendorff in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 429, Fig. 36. — Type: Lörzing 14746 (K holo; L iso), Sumatra, N. Sibajak, W. of Bandarbaru.

    Also check: Slik, Macaranga and Mallotus species of Borneo under Mallotus stipularis.

Shrubs to small trees up to 17(–27) m tall, dbh up to 30 cm. Outer bark smooth to (slightly) rough, scaly to flaking, mottled. Indumentum usually dense, short to long. Branches smooth, pustulose when young, glabrescent, clavate glands absent or present. Stipules semi-persistent, narrowly ovate to ovate to linear, 7–13 by 1–3.5 mm, often blackish when dried, margin entire, involute, apex rounded (to acute), hairy, long hairs along margin, clavate glands absent to few basally. Non-reduced leaves: petiole pustulose, 9–30 by 0.8–1.1 mm, hairy, clavate glands absent (to few); blade elliptic to obovate, 8.5–23 by 3–6.5 cm, length-width ratio 2.5–4, chartaceous to coriaceous, base obtuse to truncate to slightly cordate, often oblique, margin entire to slightly wavy to apically dentate to serrate, apex cuspidate, upper surface smooth, glabrous to hairy, glabrescent, puncticulate glands absent to numerous, lower surface smooth, hairy, puncticulate to clavate glands absent or present, domatia usually present, with hair tufts; venation pinnate, 11–15 secondary veins per side, ending parallel to the margin or in the margin. Stipuliform leaves linear to narrowly ovate, 4–10 by 1.5–2.5 mm, margin entire to wavy, apex rounded to acute, hairy, long hairs along margin, clavate glands absent. Staminate inflorescences up to 5 cm long, basally c. 0.6 mm thick, unbranched, hairy, tubular to clavate glands present, nodes up to 18 per inflorescence; bracts ovate, c. 1.1 by 0.6 mm, margin entire, apex acute, hairy abaxially only, clavate glands absent to few. Buds globose, hairy. Staminate flowers c. 4.5 mm in diameter, green-yellow to reddish; bracteoles absent; pedicels 1–2 mm long, hairy, clavate glands present; sepals 3, basally connate, elliptic, recurved, c. 3 by 2 mm, margin entire, apex acute, hairy abaxially only, clavate glands absent; stamens up to c. 56, filaments c. 2 mm long; anthers c. 0.4 by 0.3 mm, basifixed to dorsifixed; connective not to slightly widened. Pistillate inflorescences up to 8 cm long, hairy, clavate glands absent (to present), nodes up to 9 per inflorescence; bracts ovate to narrowly ovate, 1.5–4 by 1 mm, margin entire, apex acute to rounded, hairy, clavate glands absent to few. Pistillate flowers white; bracteoles absent or present, subtending the calyx, ovate, c. 3 by 1 mm, margin entire, apex acute, hairy, long hairs along margin, clavate glands absent; pedicels 3–4.5 mm long, hairy, clavate glands absent to numerous; sepals 3–5, free, ovate to narrowly triangular, 4–7 by 0.9–1.5 mm, margin entire, apex rounded to acute, hairy, long hairs along margin, clavate glands absent; ovary 3-locular, echinate, woolly hairy, clavate glands absent, spines 0.5–1.8 mm long, hairy, long hairs apically, clavate glands absent; style 1–2.5 mm long, hairy, clavate glands absent; stigmas 6–11 mm long, densely covered with granulate papillae on the inside, hairy abaxially, clavate glands absent. Fruit an echinate, lobed capsule, 10–14 by 5–10 mm, greenish; carpel inside hairy, clavate glands absent; column 4.5–5 by 4.5–5.5 mm. Seeds glossy, globose, c. 5 by 5 by 4 mm; hilum elliptical to deltoid, c. 1.5 by 1–1.5 mm.

    Distribution — Southern Thailand, Sumatra and Borneo.

    Habitat & Ecology — Mainly a locally common understorey tree in primary forest but also in secondary forest; soils alluvial to dry, basalt derived to clay rich to sandy. Altitude: up to 1100 m.

    Notes — Thai specimens can be almost glabrous.

11. Hancea wenzeliana (Slik) S.E.C. Sierra, Kulju & Welzen

    Hancea wenzeliana (Slik) S.E.C.Sierra, Kulju & Welzen in S.E.C.Sierra, Kulju, Veldkamp & Welzen, Blumea 52 (2007) 364. — Mallotus wenzelianus Slik, Blumea 43 (1998) 229; Slik & Welzen, Blumea 46 (2001) 29, Fig. 12, Map 4; in Slik, Tropenbos-Kalimantan ser. 4 (2001) 172, Fig. 7.12, Map 7.4. — Type: Wenzel 2697 (G holo; UC, A iso), The Philippines, Mindanao, Surigao Province, Jubud.

Small trees up to 7 m tall, dbh up to 15 cm. Indumentum dense, short. Branches smooth, pustulose, glabrescent, clavate glands absent to few. Stipules semi-persistent, narrowly triangular, 11–20 by 1.4–2.2 mm, margin entire, apex acute, hairy, clavate glands absent to numerous. Non-reduced leaves: petiole 10–22 by 1.5–2 mm, pustulose, hairy, clavate glands absent to numerous; blade (ovate to) elliptic, 15–27 by 4–10 cm, length-width ratio 2.6–3.7, chartaceous, base cuneate, margin slightly serrate, apex cuspidate, upper surface smooth, sparsely hairy, on midrib and veins, puncticulate glands numerous, lower surface smooth, hairy, especially on midrib and veins, puncticulate glands numerous, domatia absent; venation pinnate, 9—12 secondary veins per side, ending in the margin. Stipuliform leaves narrowly triangular, 3.5–12 by c. 1 mm, margin entire, apex acute, hairy, clavate glands present. Staminate inflorescences unknown. Pistillate inflorescences up to 11 cm long, basally c. 1 mm thick, only terminal fruit developing, unbranched, hairy, clavate glands present, nodes up to 6 per inflorescence; bracts narrowly triangular, 5–11 by 0.6–1.3 mm, margin entire, involute, apex acute, hairy, glabrous to hairy adaxially, clavate glands few to numerous. Pistillate flowers: bracteoles rarely present, linear, c. 3.3 by 0.2 mm, margin entire, involute, apex acute, hairy, clavate glands few; pedicels 1.5–6 mm long, hairy, clavate glands few; sepals 4 or 5, free, narrowly triangular, 8–12 by 0.9–1 mm, margin entire, apex acute, hairy abaxially only, clavate glands on outside only; ovary 3-locular, echinate, densely hairy, clavate glands few, spines 2–4 mm long, hairy, long hairs apically, clavate glands absent; style c. 1 mm long, hairy, clavate glands absent; stigmas up to 22 mm long, densely covered with granulate papillae on the inside, hairy abaxially, clavate glands absent. Fruit an echinate, lobed capsule, 10–14 by c. 10 mm, hairy, clavate glands few; carpel inside hairy, clavate glands absent; column c. 5 by 3–4 mm. Seeds unknown.

    Distribution — Mindanao in the Philippines.

    Habitat & Ecology — In forest. Altitude: up to 150 m.

Section Oliganthae (Airy Shaw) S.E.C.Sierra, Kulju & Welzen

    Hancea Seem. sect. Oliganthae (Airy Shaw) S.E.C.Sierra, Kulju & Welzen in S.E.C.Sierra, Kulju, Veldkamp & Welzen, Blumea 52 (2007) 364. — Mallotus Lour. sect. Oliganthae Airy Shaw, Kew Bull. 21 (1968) 389. — Cordemoya Baill. sect. Oliganthae S.E.C.Sierra, Kulju & Welzen in Sierra et al., Blumea 51 (2006) 535. — Type: Mallotus subpeltatus Blume [= Hancea subpeltata (Blume) M.Aparicio].

Branches with slender nodes. Stipules axillary, margin serrate. Leaves alternate; blade peltate, margin entire. Leaves alternate; blade peltate, margin entire; venation triplinerved, looping close to the margin. Inflorescences ramiflorous to axillary; staminate ones racemes; pistillate ones developing a single terminal flower. Staminate flowers: sepals 3 (or 4), basally connate. Pistillate flowers: sepals 5 or 6; ovary 3-locular. Fruits opening loculicidally-septicidally, inside densely hairy; spines without terminal glands.

    Distribution — From Burma to Thailand, Peninsular Malaysia, Sumatra, Borneo and Java.

12. Hancea subpeltata (Blume) M.Aparicio

    Hancea subpeltata (Blume) M.Aparicio, Blumea 52 (2007) 364. — Adisca subpeltata Blume, Bijdr. 11 (1826) 610. — Rottlera subpeltata (Blume) Baill., Étude Euphorb. (1858) 423. — Mallotus subpeltatus (Blume) Müll.Arg., Linnaea 34 (1865) 189; Welzen, Slik & Bollendorff in Welzen et al., Thai For. Bull. (Bot.) 28 (2000) 109; Welzen, S.E.C.Sierra, Slik & Bollendorff in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 431, Fig. 37. — Cordemoya subpeltata (Blume) M.Aparicio in S.E.C.Sierra et al., Blumea 51 (2006) 535. — Lectotype (M.Aparicio, 2006): Blume s.n. '1510' (holo L), Java, Krawong.

    Mappa rhynchophylla Miq., Fl. Ned. Ind. 1 (1859) 403. — Rottlera rhynchophylla (Miq.) Miq., Fl. Ned. Ind., Eerste Bijv. (1861) 181, 454. — Lectotype (M.Aparicio, 2006): Teijsmann HB 3645 (holo U), Sumatra, Tubuan, Ogan Ulu.

    Mappa acuminatissima Zipp. ex Span., Linnaea 15 (1841) 349, nom. nud. — Representative specimen: Zippelius s.n. (L?).

    Also check: Slik, Macaranga and Mallotus species of Borneo under Mallotus subpeltatus.

Shrubs to small trees, up to 15 m high, dbh up to 15 cm, dioecious. Outer bark smooth, grey-brown. Indumentum composed of short (with a pointed tip), long simple and tufted hairs, and capitate glandular hairs with multicellular stalks. Stipules narrowly triangular, 5–7 by 1.8–2 mm, caducous to early caducous, margin serrate, hairy, apex acute, sparsely to densely hairy on both sides. Leaves: petiole 1–14 cm by 1–1.3 mm, not grooved above, glabrescent; blade ovate to elliptic, 7.5–27 by 6–11 cm, length/width ratio 2.5–3, chartaceous, base subpeltate up to 2 mm, rounded to obtuse, margin entire, apex caudate, upper surface sparsely hairy, glabrescent, lower surface sparsely hairy on midrib and nerves, with few glandular hairs, nerves 5–7 per side, with hair tuft domatia. Inflorescences single or two together; peduncle 0.5–1.2 by c. 1 mm; staminate racemes 0.5–0.8 cm long, nodes per branch 3–6; pistillate ones 1.5–3 cm long, bracts 4 or 5; bracts triangular, 1.8–2.5 by 1–1.5 mm, persistent, margin entire, apex acute, upper surface hairy, lower surface densely hairy; bracteoles absent. Flowers: pedicels densely hairy; sepals margin entire, apex acute. Staminate flowers 5–7 mm in diam.; pedicels 5–7 mm long; sepals ovate, 4.5–5 by 2–2.5 mm, densely hairy outside, subglabrous to sparsely hairy inside; stamens 200–250, glabrous, filaments 2–7 mm long, thecae ellipsoid, 0.5–0.7 by 0.4–0.5 mm, connective widened. Pistillate flowers c. 5 mm in diam.; pedicels 0.7–2 mm long; sepals triangular, 4–6 by 1–1.5 mm, valvate and/or imbricate, hairy; staminodes absent; ovary 1–2 by 2–3 mm, densely hairy; style 1–1.5 mm long; stigma 8–12 mm long. Fruits 8–15 by 15–20 mm, pale green, densely hairy outside, spines 140–170, straight, 6–10 by c. 1 mm, densely hairy; wall 1–2 mm thick; column 9.5–15 by 9–14 mm. Seeds 9–12 by 9–11 by 8–10 mm, smooth, shiny, dark brown.

    Distribution — Burma, Thailand, Peninsular Malaysia, Sumatra, Java, Borneo.

    Habitat & Ecology — Locally in understory of evergreen forest. Altitude: sea level up to 1400 m. Flowering: February to May; fruiting: June to January.