Malesian Euphorbiaceae Descriptions

67. MALLOTUS (Euphorbiaceae)

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References

Genus description

Descriptions of still recognized sections:

    Mallotus section Mallotus

    Mallotus section Philippinensis (grade to sect. Mallotus)

    Mallotus section Polyadenii

    Mallotus section Rottleropsis

    Mallotus section Stylanthus

Key to the species

Species

Key to the species of East Kalimantan

Notes on Unidentified Specimens

Dubious Species

References

    Bollendorff, S.M., P.C. van Welzen & J.W.F. Slik. 2000. A taxonomic revision of Mallotus Section Polyadenii (Euphorbiaceae). Blumea 45: 319–340.

    Kulju, K.K.M., S.E.C Sierra & P.C. van Welzen. 2007. Re-shaping Mallotus [Part 2]: Inclusion of Neotrewia, Octospermum and Trewia to Mallotus s.s. (Euhporbiaceae s.s.). Blumea 52: 115–136.

    Sierra, S.E.C., M. Aparicio, M.J.H. Gebraad, K.K.M. Kulju & P.C. van Welzen. 2007. The morphological range in Mallotus (Euphorbiaceae) and a taxonomic revision of its section Rottleropsis (including Axenfeldia) in Malesia, Thailand and Africa. Blumea 52: 21–113.

    Sierra, S.E.C. & P.C. van Welzen. 2005. A taxonomic revision of Mallotus section Mallotus (Euphorbiaceae) in Malesia. Blumea 50: 249–274.

    Sierra, S.E.C., P.C. van Welzen & J.W.F. Slik. 2005. A taxonomic revision of Mallotus section Philippinensis (former section Rottlera – Euphorbiaceae) in Malesia and Thailand. Blumea 50: 221–248.

    Slik, J.W.F. website: Macaranga and Mallotus species of Borneo

    Slik, J.W.F., Priyono & P.C. van Welzen. 2000. Key to the Macaranga and Mallotus species (Euphorbiaceae) of East Kalimantan, Indonesia. Gard. Bull. Singapore 52: 11–87.

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

    Welzen, P.C. van & S.E.C. Sierra. 2006. The Mallotus wrayi complex (Euphorbiaceae). Blumea 51: 373–388.

    Welzen, P.C. van, S.E.C. Sierra, M.J.H. Gebraad & K.K.M. Kulju. The distinction of five Mallotus species formerly in section Hancea (Euphorbiaceae). Blumea 51: 367–372.

Mallotus Lour.

    Mallotus Lour., Fl. Cochinch. (1790) 635, nom. cons.; Pax & K.Hoffm. in Engl., Pflanzenr. VI.147.vii (1914) 145; Airy Shaw, Kew Bull. 26 (1972) 292; Whitmore, Tree Fl. Malaya 2 (1974) 113; Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 160; G.L.Webster, Ann. Missouri Bot. Gard. 81 (1994) 89; Bollendorff et al., Blumea 45 (2000) 319; Welzen, Slik & Bollendorff in Welzen et al., Thai For. Bull. (Bot.) 28 (2000) 93; Slik & Welzen, Blumea 46 (2001) 3; Radcl.-Sm., Gen. Euphorbiacearum (2001) 225; Welzen, S.E.C.Sierra, Slik & Bollendorff in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 384; G.L.Webster in Kubitzki, Fam. Gen. Vasc. Pl. 11 (2014) 126, Fig. 28. — Type: Mallotus cochinchinensis Lour. [= Mallotus paniculatus (Lam.) Müll.Arg. var. paniculatus].

    Trewia (‘Trevia’, see notes 1 and 2) L., Sp. Pl. 2 (1753) 1193, nom. rej.; Burm.f., Fl. Indica (1768) 298 (‘Trevia’); Neck., Elem. Bot. 2 (1790) 268; Willd., Sp. Pl. ed 4, 2 (1806) 834; Sm. in Rees, Cyclop. 36 (1819) 7; Blume, Bijdr. (1826) 612; Endl., Gen. Pl., Suppl. 3 (1843) 98; Baill., Étude Gen. (1858) 408; Miq., Fl. Ned. Ind. 1, 2 (1859) 399; Müll.Arg. in DC., Podr. 15, 2 (1866) 953; Bedd., Fl. Sylv. 2 (1872) 281; Kurz, Forest Fl. 2 (1877) 378; Benth. in Benth. & Hook.f., Gen. Pl. 3, 1 (1880) 318; Hook.f., Fl. Br. Ind. 5 (1887) 423; Pax in Engl. & Prantl, Nat. Pflanzenfam. 3, 5 (1896) 53; Brandis, Ind. Trees (1906) 590; T.Cooke, Fl. Presid. Bombay 2, 3 (1906) 613; J.J.Sm., Meded. Dept. Landb. Ned.-Indië 10 (1910) 389; Talbot, Forest Fl. Bombay Pres. Sind 2 (1911) 481; Pax & K. Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 140; Gamble, Fl. Presid. Madras 2, 7 (1925) 1318; Backer & Bakh.f., Fl. Java 1 (1964) 481 (‘Trevia’); Airy Shaw, Kew Bull. 26 (1972) 343; Whitmore, Tree Fl. Malaya 2 (1973) 134; Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 200; Kew Bull. 36 (1981) 350; Alphab. Enum. Euph. Philipp. Isl. (1983) 46; Grierson & D.G.Long, Fl. Bhutan 1, 3 (1987) 798; G.L.Webster, Ann. Missouri Bot. Gard. 81 (1994) 90; H.S.Kiu et al., Fl. Reip. Pop. Sin. 44, 2 (1996) 11; Philcox in Dassan. & W.D.Clayton, Rev. Handb. Fl. Ceylon 11 (1997) 142; Susila & N.P.Balakr., J. Econ. Taxon Bot. 22 (1998) 345; Govaerts et al., World Checkl. Bibliogr. Euphorb. 3 (2000) 1560; Radcl.-Sm., Gen. Euphorb. (2001) 232; ; Kulju, S.E.C.Sierra & Welzen, Blumea 52 (2007) 119, map 1. — Canschi Adans., Fam. Pl. 2 (1763) 443, nom. superfl. — Type: Trewia nudiflora L. [= Mallotus nudiflorus (L.) Kulju & Welzen].

    Neotrewia Pax & K. Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 211; Merr., Enum. Philipp. Fl. Pl. 2 (1923) 437; Pax & K. Hoffm. in Engl. & Harms, Nat. Planzenfam. ed. 2, 19c (1931) 118; Airy Shaw, Kew Bull. 37 (1982) 31; Alphab. Enum. Euph. Philipp. Isl. (1983) 39; G.L.Webster, Ann. Missouri Bot. Gard. 81 (1994) 90; Govaerts et al., World Checkl. Bibliogr. Euphorb. 4 (2000) 1217; Radcl.-Sm., Gen. Euphorb. (2001) 232. — Type: Neotrewia cumingii (Müll.Arg.) Pax & K. Hoffm. [= Mallotus cumingii Müll.Arg.].

    Octospermum Airy Shaw, Kew Bull. 19 (1965) 311; Kew Bull., Addit. Ser. 8 (1980) 176; G.L.Webster, Ann. Missouri Bot. Gard. 81 (1994) 90; Govaerts et al. , World Checkl. Bibliogr. Euphorb. 4 (2000) 1219; Radcl.-Sm., Gen. Euphorb. (2001) 232. — Type: Octospermum pleiogynum (Pax & K.Hoffm.) Airy Shaw, [= Mallotus pleiogynus Pax & K.Hoffm.

Shrubs, trees, to rarely climbers, dioecious (or monoecious), up to 35 m tall, evergreen or sometimes deciduous. Indumentum consisting of simple, tufted, stellately-tufted, or stellate, or rarely peltate hairs; glandular hairs globular to disc-shaped, usually conspicuous, rarely absent. Bark sometimes prominently hooped (rings around stem); wood soft to hard. Stipules axillary, small, often caducous. Leaves alternate to opposite, usually unequal in size when opposite, simple, sometimes lobed; blade: base sometimes peltate, margin entire to crenate to dentate, often with minute glandular teeth, upper surface almost always with elliptic or orbicular, usually conspicuous, extrafloral nectaries, which are basally and/or marginally distributed (or along midrib or all over the lamina), glandular hairs present or absent; lower surface usually with glandular hairs, hairy domatia sometimes present; venation pinnate, triplinerved or palminerved, nerves looping or ending in the margin, veins scalariform, veinlets reticulate. Inflorescences axillary or terminal, rarely ramiflorous or cauliflorous, spikes, racemes, panicles, glomerules or umbel-like, or reduced to one flower, unisexual, rarely bisexual; bracts 1 (several when umbel-like) per node, margin entire, rarely trilobed, bracteoles sometimes present; staminate flowers 1–15 (to rarely more) per bract, pistillate flowers one per bract (to several when umbel-like). Flowers not exceeding 10 mm diam., usually pedicellate, pedicels sometimes with abscission zone; sepals valvate, free or connate, usually persistent; petals absent. Staminate flowers: sepals (2 or) 3–5; interstaminal disc-glands sometimes present; stamens 15–130, filaments free or connate, anthers basifixed, thecae 2, longitudinally dehiscent, connective narrow to distinctly broadened, sometimes irregularly apiculate, papillose; pollen 3- (rarely 4-)colporate, ornamentation perforate/microreticulate with scabrae; pistillode sometimes present. Pistillate flowers: sepals (2 or) 3–6; disc absent; ovary (1- or) 2- or 3- (to rarely 9-)locular, 1 ovule per locule; style present or absent; stigmas simple, (erect to) reflexed, (papillose or) plumose. Fruits dehiscent to tardily dehiscent capsules, or indehiscent, drupaceous and variously fleshy, usually lobed (to globose), rarely with wings or longitudinal ridges, surface smooth to spiny (to somewhat rugose to verrucose), hairy (to glabrous), sometimes with glandular hairs, opening septicidally-loculicidally to loculicidally-septicidally, or rarely only loculicidally; column usually present. Seeds (±) globose, rarely triangular in cross section or lenticular, sometimes with a thin sarcotesta, surface smooth to rugose, caruncle absent.

    Distribution — From West (Senegal to Liberia) to East (Ethiopia to Tanzania) Africa to West Madagascar, Pakistan, Sri Lanka, India, Bangladesh, Myanmar and Nepal, South China, Korea, South Japan, throughout Southeast Asia and Malesia to East Australia and the West Pacific (Solomon Islands, Palau, Federated States of Micronesia, New Caledonia, Fiji). 54 species in Malesia.

    Notes — 1. Trewia L. is the oldest name for the genus, as circumscribed here. However, we have submitted a proposal to conserve Mallotus against Trewia to maintain nomenclatural stability (see text).

2. Originally, Linnaeus (1753) spelled the genus name as ‘Trevia’, but later he inconsistently used both ‘Trevia’ and ‘Trewia’ (see Nicolson et al., Regnum Veg. 119, 1988). Since Linnaeus, almost all authors have adopted the spelling ‘Trewia’. The present code (McNeill et al., Intern. Code Bot. Nom., Regnum Veg. 46, 2006: art. 13.4) gives priority to the original spelling given in Species Plantarum (1753) and therefore ‘Trevia’ should be used. However, since this genus is here present only in the synonymy of Mallotus, we have continued to follow the widely used spelling ‘Trewia’.

Mallotus Lour. section Mallotus

    Mallotus Lour. sect. Mallotus: Pax & K. Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 162; Airy Shaw, Kew Bull. 21 (1968) 386, S.E.C.Sierra & Welzen, Blumea 50 (2005) 250. — Mallotus sect. Eumallotus Müll.Arg., Linnaea 34 (1865) 186, nom. inval. — Type: Mallotus cochinchinensis Lour. [= Mallotus paniculatus (Lam.) Müll.Arg. var. paniculatus].

    Echinus Lour., Fl. Cochinch. 1 (1790) 633. — Mallotus sect. Echinus Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 162, nom. superfl. — Type: Echinus trisulcus Lour. [= Mallotus paniculatus (Lam.) Müll.Arg. var. paniculatus].

    Rottlera Roxb. ex Willd. sect. Trelotra Baill., Étude Euph. (1858) 425.— Type: Rottlera japonica (Thunb.) Spreng. [= M. japonicus Müll.Arg.].

    Mallotus sect. Melanolepis auct. non Müll.Arg.: Müll.Arg., Linnaea 34 (1865) 184, quoad Mallotus barbatus Wall. ex Müll.Arg.

    Rottlera Roxb. ex Willd. sect. Eurottlera auct. non Rchb.f. & Zoll.: Rchb.f. & Zoll., Linnaea 28 (1857) 314, quoad Rottlera zippelii Zoll. & Moritzi, Rottlera zippelii Zoll. & Moritzi var. minor Rchb.f. & Zoll. (nom. nud.), Rottlera paniculata (Lam.) Blume.

Shrubs to small trees, monoecious or dioecious; branches glabrescent. Dark coloured exudate present in fresh twigs. Indumentum composed of stellate hairs, and sessile, globular to disc-shaped, light yellow to orange glandular hairs. Stipules with entire or subentire margin. Leaves alternate to apically subopposite, simple; petiole basally pulvinate; blade subpeltate or peltate, upper surface glabrescent, extrafloral nectaries basally or marginally, orbicular to elliptic, midrib, nerves and veins occasionally with glandular hairs, prominent, 3- or palminerved, nerves looping to ending in the margin, veins scalariform, veinlets reticulate. Inflorescences axillary or terminal, unisexual or bisexual; bracts 1 per node; bracteoles present or absent, both triangular to linear triangular, hairy, margin entire or subentire, apex acuminate. Flowers actinomorphic, not exceeding 1 cm diam.; pedicels hairy; sepals persistent, valvate, densely hairy outside, subglabrous to sparsely hairy inside, with glandular hairs on both sides, margin entire, apex acute; petals and disc absent. Staminate inflorescences panicles, erect, with several flowers per bract. Staminate flowers: sepals 3–5, ovate to elliptic, free, reflexed; stamens 40–90, glabrous, anthers basifixed, thecae 2, separate from each other, ovoid, opening extrorse and lengthwise, sometimes the apex with glandular hairs, connec-tive widened, papillose; pistillode present (or absent), flattened to raised, 0.1–0.3 mm long, consisting of wart-like, glabrous appendices. Pistillate inflorescences racemes, panicles or spikes, becoming pendulous, with one flower per bract. Pistillate flowers: pedicels absent or present; calyx 3–5, lobes ovate, connate (on the base or almost to the apex), erect, persistent in fruits; ovary with visible spines or not, with glandular hairs, locules 3–5, 1 ovule per locule; style up to 1.5 mm long; stigmas narrowly triangular, plumose, densely covered with papillae above, outer surface hairy and with glandular hairs; staminodes present or absent. Fruits dehiscent capsules, surface spiny and densely hairy; wall glabrous inside. Seeds ellipsoid to globose, somewhat trigonous in transverse section; caruncle or aril absent.

    Distribution — From India (W Bengal) to South China and South Japan, throughout Southeast Asia and Malesia to East Australia and West Pacific (Solomon Islands). Four species in Malesia.

Mallotus section Philippinensis Pax & K.Hoffm.

    Mallotus Lour. sect. Philippinenses Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 178. — Type: Mallotus philippensis (Lam.) Müll.Arg.

    Rottlera Roxb. ex Willd., Sp. Pl., ed. 4, 2 (1806) 641, 832, non Willd. (1798). — Rottlera Roxb. ex Willd. sect. Eurottlera Rchb.f. & Zoll., Linnaea 28 (1857) 314, nom. inval. — Mallotus Lour. sect. Eumallotus Müll.Arg., Linnaea 34 (1865) 186, nom. inval.— Mallotus Lour. sect. Rottlera ((Willd.) Rchb.f. & Zoll., corr.) Airy Shaw, Kew Bull. 21 (1968) 392, nom. superfl. — Type: Rottlera tinctoria Roxb. [= Mallotus philippensis (Lam.) Müll.Arg.].

Woody climbers or small trees, dioecious; branches glabrescent. Indumentum composed of stellate, and/or simple hairs, and sessile, globular to disc-shaped, light yellow to red glandular hairs (sometimes drying greenish). Stipules: margin entire, apex acuminate. Leaves alternate to apically subopposite, simple; petiole basally and apically pulvinate or not; blade not peltate, upper surface glabrous except on midrib and nerves (to rarely hairy all over), extrafloral nectaries basally or marginally, orbicular to elliptic, midrib, nerves and veins occasionally with glandular hairs, prominent, 3-nerved, nerves looping to ending in the margin,veins scalariform, veinlets reticulate. Inflorescences paniclesor racemes, the latter axillary (1 or 2 together) or terminally grouped (1 to several), erect, unisexual; bracts 1 per node; bracteoles present or absent, both triangular,margin entire, apex acute to acuminate. Flowers actinomorphic, not exceeding 1 cm diam.; pedicels hairy; sepals persistent, valvate, densely hairy outside, subglabrous to sparsely hairy inside, with glandular hairs on both sides, margin entire, apex acute; petals and disc ab-sent. Staminate inflorescences with several flowers per bract. Staminate flowers: sepals 3–5, ovate to elliptic, free, reflexed; stamens18–75, glabrous to hairy, anthers basifixed, thecae 2, separate from each other, ovoid to ellipsoid, opening extrorse and lengthwise, sometimes the apex with glandular hairs, connective widened or not, papillose; pistillode present or absent. Pistillate inflorescences with one flower per bract. Pistillate flowers: sepals 3–6, ovate or narrowly triangular, free, reflexed or erect, persistent in fruits; ovary with glandular hairs, locules 2 or 3, 1 ovule per locule; stigmas sessile or not, narrowly triangular, plumose, densely covered with papillae above, outer surface hairy and with glandular hairs; staminodes absent. Fruits dehiscent capsules, surface with glandular hairs; wall glabrous to sparsely hairy inside. Seeds globose, somewhat trigonous in transverse section; caruncle or aril absent.

    Distribution — From Pakistan to South China and South Japan, throughout Southeast Asia and Malesia to East Australia and New Caledonia.

Mallotus section Polyadenii Pax & K.Hoffm.

    Mallotus Lour. sect. Polyadenii Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 197; Airy Shaw, Kew Bull. 16 (1963) 350; Kew Bull. 20 (1966) 39, 43; Kew Bull. 21 (1968) 388, 397; Kew Bull. 26 (1972) 292, 299; Kew Bull. 27 (1972) 87; Kew Bull. Add. Ser. 4 (1975) 160, 166; Kew Bull. 31 (1976) 392; Kew Bull. 35 (1980) 162; Kew Bull. 36 (1981) 323; Bollendorff, Welzen & Slik, Blumea 45 (2000), 320; in Slik, Tropenbos-Kalimantan ser. 4 (2001) 122. — Type species: Mallotus polyadenos F.Muell.

    Coccoceras Miq., Fl. Ned. Ind., Eerste bijv. (1860) 455; Müll.Arg. in DC., Prodr. 15, 2 (1866) 949; Benth. in Benth. & Hook.f., Gen. Pl. 3 (1880) 318, 319; Pax in Engl. & Prantl, Pflanzenfam. 3, 5 (1890) 55; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 209; G.L.Webster, Ann. Missouri Bot. Gard. 81 (1994) 90. — Type species: Coccoceras sumatranum Miq. [= Mallotus sumatranus (Miq.) Airy Shaw]

Shrubs to trees, occasionally with buttresses, dioecious, rarely monoecious; branchlets terete to flattened, usually lenticellate. Indumentum consisting of simple and stellate, white hairs, sometimes also villose red hairs; all parts with transparent to yellow to orange-red discoid glands, highest density on young parts and lower leaf surface, fewest on branchlets. Bark smooth, sometimes finely fissured; sapwood often pink turning dark after exposure. Leaves petiolate, simple, opposite to alternate, unequal in size when opposite; stipules small, triangular, margin entire, caducous; blade elliptic to obovate, chartaceous to coriaceous, never entirely symmetric, base acute to slightly cordate, margin entire to slightly crenate with marginal glands in the crenations, glands protruding in young leaves, apex rounded to obtusely cuspidate, often ending in a gland, upper surface dull, usually glabrous, basally (and submarginally and apically) with impressed glands, lower surface dull, pubescent or glabrous except for hair-tuft (or pocket) domatia in axils between midrib and nerves; venation pinnate, basally often triplinerved, nerves brachidodromous to eucamptodromous, clearly raised below, slightly raised above, veins scalariform to sometimes reticulate, veinlets reticulate. Inflorescences catkin-like when young, raceme-like thyrses, axillary though staminate ones often terminal, solitary or in groups of 2 or 3, unbranched (except sometimes in M. polyadenos), straight. Bracts persistent, triangular, margin entire, apex acute to acuminate. Flowers small, actinomorphic, white, yellow to green; calyx lobes unequal in width, margin entire; petals, disc, and pistillode absent. Staminate flowers in clusters of 3–7 per node, shortly pedicellate; calyx lobes 3–5, basally connate, generally ovate, membranous; stamens numerous, inserted on a dome-shaped receptacle, filaments filiform, elongating at anthesis, anthers ovoid to ellipsoid, basifixed to dorsifixed, thecae 2, parallel, opening lengthwise with slits, connective slender to broad, sometimes split. Pistillate flowers usually solitary per node; pedicels elongating in fruit, articulate, abscission zone sometimes geniculate; calyx lobes 3–6, basally to halfway connate, triangular (to ovate), base thickened; ovary ovoid to ellipsoid, (2- or) 3- (or 4-)locular, uniovulate, placentation axillary; ovules anatropous, descending, epitropous; stigmas sessile or not, strongly plumose with long, branched glandular-granular papillae. Fruits mainly tricoccoid, dehiscent woody capsules, first splitting septicidally from base to apex, then loculicidally from base to apex, or indehiscent (or tardily dehiscent) and winged or ridged, smooth; column bell- to anchor-shaped from lateral view. Seeds globose to ovoid, shiny or dull, smooth or sometimes with white protuberances, light to dark brown.

    Distribution—Eight species in Sri Lanka, S India, SE Asia (Myanmar, Laos, Cambodia, Vietnam, Thailand), W Malesia (Peninsular Malaysia, Sumatra, Borneo), New Guinea, Australia (NE Queensland), and the Solomon Islands. Unknown from Central Malesia.

Mallotus section Rottleropsis Müll.Arg.

    Mallotus Lour. sect. Rottleropsis Müll.Arg. in DC., Prodr. 15, 2 (1866) 957; Airy Shaw, Kew Bull. 21 (1968) 379; Kew Bull. 26 (1972) 301; S.E.C.Sierra et al., Blumea 52 (2007) 34. — Type species: Mallotus lappaceus Wall. ex Müll.Arg., see note 1.

    Echinocroton F. Muell., Fragm. Phytogr. Austr. 1 (1858) 31. — Mallotus Lour. sect. Echinocroton (F. Muell.) Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 148; Airy Shaw, Kew Bull. 20 (1966) 41. — Type species: Echinocroton claoxyloides F. Muell. (= Mallotus claoxyloides (F. Muell.) Müll.Arg.).

    Plagianthera Rchb.f. & Zoll., Acta Soc. Regiae Sci. Indo- Neerl. 1 (1856) 19. — Mallotus Lour. sect. Plagianthera (Rchb.f. & Zoll.) Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 156. — Type species: Plagianthera oppositifolia Rchb.f. & Zoll. (= Mallotus blumeanus Müll.Arg., non Mallotus oppositifolius (Geiseler) Müll.Arg.).

    Mallotus Lour. sect. Axenfeldia Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 187; Airy Shaw, Kew Bull. 21 (1968) 393; Kew Bull. 26 (1972) 293. — Type species: Axenfeldia intermedia Baill. (= Mallotus intermedius (Baill.) N.P.Balakr.).

Shrubs to trees, dioecious or rarely monoecious. Indumentum composed of simple, tufted, stellately-tufted (= 5 rays, radially symmetrical), rarely peltate hairs, and usually globular to disc-shaped glandular hairs, rarely capitate glandular hairs. Stipules usually present. Leaves opposite, unequal in size, rarely also alternate; blade base rarely peltate; margin entire to crenate to dentate, often with minute glandular teeth, upper surface usually with generally conspicuous extrafloral nectaries, sometimes with glandular hairs, lower surface usually with glandular hairs, hair tuft domatia sometimes present, venation pinnate, triplinerved or palminerved, nerves looping or ending in margin. Inflorescences axillary or terminal, rarely ramiflorous or cauliflorous, unisexual, rarely bisexual; bracts 1 (to several when inflorescence umbel-like) per node, bracteoles sometimes present. Flowers not exceeding 10 mm diam.; sepals free or connate, usually persistent; petals absent. Staminate inflorescences racemes, rarely glomerules, with 1–15 (to rarely more) flowers per bract/node. Staminate flowers: sepals (2–)3–5(–6); disc-glands rarely present, 3–16; stamens 20–130, filaments free or connate; pistillode often present. Pistillate inflorescences racemes, umbel-like, or sometimes reduced to a terminal flower; with 1 to several (when umbel-like) flowers per bract. Pistillate flowers: sepals (2–)3–6; staminodes rarely present; ovary (1–)2–4(–5)-locular, 1 ovule per locule; style usually present; stigmas (erect to) reflexed, (papillose or) plumose adaxially. Fruits dehiscent to tardily dehiscent capsules, or indehiscent, drupaceous and variously fleshy, oblate (to globose), usually lobed, surface smooth (to somewhat rugose to verrucose) to spiny, glandular and/or hairy (to glabrous), opening septicidally-loculicidally to sometimes loculicidally-septicidally, rarely only loculicidally; wall rarely with a thick, fleshy, whitish endocarp; column usually present. Seeds ± globose, rarely lenticular or triangular in cross section, surface smooth to rugose; rarely covered with a thin sarcotesta.

    Distribution — From SW (Senegal to Liberia) to SE (Ethiopia to Tanzania) Africa to W Madagascar, Sri Lanka, India, Myanmar and Nepal, S China, throughout SE Asia and Malesia to E Australia and the W Pacific (Solomon Islands, Republic of Palau, Federated States of Micronesia and Fiji).

    Notes — 1. Wallich 7738 and 7845 from Myanmar are mentioned in the protologue of M. lappaceus. Although we have only been able to study them from digital pictures sent by K, their appearance and description by Müller Argoviensis (1866) agree with the concept of sect. Rottleropsis s.l. The collections have long stipules and bracts, which are features that are also present in M. dispar and M. leucocalyx.

2. We estimate that c. 30 names have been published for non-Malesian taxa (not treated in this study), which might also belong to sect. Rottleropsis s.l.

Mallotus section Stylanthus (Rchb.f. & Zoll.) Pax & K.Hoffm.

    Mallotus sect. Stylanthus (Rchb.f. & Zoll.) Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 172; Airy Shaw, Kew Bull. 20 (1966) 38; Kew Bull. 21 (1968) 387; Kew Bull. 26 (1972) 293; Kew Bull. Add. Ser. 4 (1975) 160; Kew Bull. Add. Ser. 8 (1980) 163; Muelleria 4 (1980) 233; Kew Bull. 36 (1981) 323; Slik & Welzen, Blumea 46 (2001) 31; in Slik, Tropenbos-Kalimantan ser. 4 (2001) 174. — Adisca Blume, Bijdr. Fl. Ned. Ind. 11 (1825) 609. — Rottlera sect. Stylanthus Rchb.f. & Zoll., Verhand. Natuurk. Vereen. Ned.-Indië 1 (1856) 312. — Type species: Adisca floribunda Blume [ = Mallotus floribundus (Blume) Müll.Arg.].

Shrubs to small trees, dioecious, (strongly) smelling of fenugreek when dried. Indumentum simple, tufted and stellate, most parts covered with glands (gland-dotted) producing a clear yellowish globosely shaped exudate. Branches often conspicuously lenticellate. Stipules usually caducous. Leaves terminally grouped on branches, simple, peltate or not, alternate to apically pseudo-opposite, chartaceous when dried; petioles often with constricted base and / or apex when dried; blade margin with glands, upper surface always with conspicuous basal macular glands, more or less gland-dotted, usually densely covered with white granular dots, lower surface usually conspicuously gland-dotted, domatia absent or present; venation palmate, tripliveined or pinnate, ending in or parallel to the margin. Inflorescences not branched, axillary to terminal, usually solitary; bracts usually (early) caducous. Flowers actinomorphic, small, not exceeding 1 cm in diameter, petals absent, disc absent. Staminate inflorescences raceme-like thyrses, several flowers per bract. Staminate flowers: bracteoles usually absent; sepals 2–5, often recurved; anthers basi- to dorsifixed; thecae 2, parallel, extrorse, opening lengthwise, sometimes connective widened; pistillode absent. Pistillate inflorescences racemes. Pistillate flowers: calyx caducous to (partly) persistent, completely enclosing ovary with only stigmas exerted, when fruits develop the calyx is torn open into 2–5-lobed; ovary usually 3-locular, echinate; ovules 1 per locule, axillary, apotropous, ascending; styles persistent, apically splitting into 3 lineate, plumose stigmas; staminodes absent. Fruits dehiscent, usually 3-lobed, echinate, loculicidal-septicidal capsules; carpels inside glabrous or tufted to stellate, gland-dotted or not. Seeds globose to laterally flattened, usually 5–8 mm in diameter.

    Distribution — From India and Sri Lanka to Southern China to New Guinea, the Solomon Islands and Australia.

    Habitat & Ecology — In primary and secondary forest, often preferring open places (gaps, river and road sides), but sometimes also in the forest understorey.

Key to the species

(only distribution within Malesia provided)

1.6 Mallotus actinoneurus Airy Shaw (sect. Rottleropsis)

    Mallotus actinoneurus Airy Shaw, Kew Bull. 33 (1978) 63; S.E.C.Sierra et al., Blumea 52 (2007) 46, map 1. — Type: KEP FRI (Sohadi) 17911 (holo K; iso L), Peninsular Malaysia, Trengganu, Jerteh, Gunong Tebu Forest Reserve.

Shrubs, 1.5–3 m high, dioecious. Indumentum composed of simple, tufted and stellately-tufted hairs, glandular hairs absent. Stipules narrowly triangular, 10–13 by 2.5–3.5 mm, persistent, margin entire, apex acute, densely to sparsely hairy. Leaves opposite to rarely alternate; petiole 14–61 by 1.5–3 mm, densely hairy; blade narrowly obovate, 25–32 by 7–14 cm, length/width ratio 2.1–3.6, chartaceous, base shallowly emarginate, margin entire to denticulate, with glandular teeth, apex caudate, upper surface sparsely to scatteredly hairy on midrib and nerves, larger extrafloral nectaries marginal in lower half, 3 or 4 per side, 1–4 mm from margin, elliptic to orbicular, 0.7–0.8 by 0.7–0.8 mm, smaller ones all over the blade, 0.2–0.3 by 0.2–0.3 mm, lower surface sparsely hairy, domatia absent, venation pinnate, nerves 9 or 10 per side, looping and closed near margin. Staminate inflorescences and flowers unknown. Infructescences racemes, with a single terminal fruit, c. 1.7 cm long, erect, single; peduncle c. 2 mm long; bracts linear-triangular, 3.5–7 by 0.7–1 mm, margin entire or trilobed, apex acute, densely hairy; bracteoles absent. Pistillate flowers (based on young fruit) c. 10 mm diam.; pedicels c. 2.5 mm long, densely hairy; sepals 3, triangular to narrowly triangular, 8–10 by c. 3 mm, free, margin entire, apex acute; ovary c. 9 by 8 mm, 3-locular, densely hairy; style c. 2.2 mm long; stigmas c. 6 by 1–1.2 mm. Fruits capsules, 15–18 by 23–27 mm, opening septicidally-loculicidally, surface smooth, densely hairy; wall 1.8–2 mm thick, glabrous inside; column c. 11.5 by 11.5 mm. Seeds ± globose, c. 12 by 11.5 by 12 mm, surface rugose, dull, light brown.

    Distribution — Endemic in the border area of Thailand and the Malay Peninsula.

    Habitat & Ecology — In logged area. Altitude c. 80 m. Fruiting: April.

2. Mallotus attenuatus Airy Shaw (sect. Rottleropsis)

    Mallotus attenuatus Airy Shaw, Kew Bull. 33 (1978) 63; S.E.C.Sierra et al., Blumea 52 (2007) 47, map 2. — Type: NGF (Streimann & Kairo) 27781 (holo K; iso A, BO, BRI, CANB, SING), Papua New Guinea, Central Prov., Brown River.

Shrubs, dioecious. Indumentum composed of simple and stellately-tufted hairs, glandular hairs absent to rarely present. Stipules linear-triangular, 2–4 by 0.2–0.3 mm, caducous, margin entire, apex acute, scatteredly hairy to glabrous outside, glabrous inside. Leaves opposite; petiole 2–8 by 0.8–1 mm, densely to sparsely hairy; blade narrowly elliptic, 8.2–16 by 1.7–4.7 cm, length/width ratio 3.2–4.8, membranaceous, base cuneate to attenuate, margin crenulate, without glandular teeth, apex caudate, upper surface glabrous, extrafloral nectaries sometimes present, inconspicuous, marginal in lower half, 1 per side, 0.2–0.5 mm from margin, elliptic to orbicular, 0.2–0.3 by 0.2–0.3 mm, lower surface scatteredly hairy, glabrescent, glandular hairs absent to rarely present (on midrib of young leaves), domatia sometimes present, venation pinnate, nerves 9–12 per side, looping and closed near margin. Staminate inflorescences reduced racemes, c. 0.5 cm long (still young, with buds), single, erect; peduncle 2–4 mm long; axes densely hairy, nodes per branch 2 or 3, with 3–5 flowers per bract; bracts linear-triangular, 2–2.2 by 0.2–0.3 mm, persistent, apex acute, margin entire, scatteredly hairy to glabrous; bracteoles triangular, 0.5–1 by 0.2–0.3, persistent to caducous. Staminate flowers 3–4 mm diam.; pedicels 2.5–3 mm long, densely hairy; sepals 3, elliptic to obovate, 2–2.2 by 1–1.2 mm, free, persistent, margin entire, apex acute, densely hairy outside, glabrous inside; stamens 25–30, glabrous, filaments 1.8–2.5 mm long, free, thecae ellipsoid, 0.2–0.3 by 0.2–0.25 mm, connective widened; pistillode present. Pistillate inflorescences, flowers and fruits unknown.

    Distribution — New Guinea.

    Habitat & Ecology — Altitude c. 90 m. Flowering: February.

    Note — Mallotus attenuatus differs from M. claoxyloides in the absence to rarely presence of glandular hairs (always present in M. claoxyloides), inconspicuous extrafloral nectaries, sometimes absent (conspicuous, always present in M. claoxyloides), 9–12 nerves per side (7 or 8 in M. claoxyloides) and a cuneate leaf base (shallowly emarginate, rounded, obtuse, to cuneate in M. claoxyloides).

3. Mallotus barbatus Wall. ex Müll.Arg. (sect. Mallotus)

    Mallotus barbatus Wall. ex Müll.Arg., Linnaea 34 (1865) 184; Backer & Bakh.f., Fl. Java 1 (1964) 482; Airy Shaw, Kew Bull. 37 (1982) 28; Corner, Ways. Trees Malaysia, ed. 3, 1 (1988) 305; Welzen, Slik & Bollendorff in Welzen et al., Thai Forest bull., Bot. 28 (2000) 97; ; S.E.C.Sierra & Welzen, Blumea 50 (2005) 251, fig. 1, map 1. — [Rottlera barbata Wall., Num. List (1847) no. 7822, nom. nud.; Baill., Étude Euph. (1858) 423, nom. nud. — Lectotype (Sierra & Welzen, 2005): Amherst 1483 = Wallich Numer. List 7822C (holo G no. 1165; iso G no. 1164, K-W (photo in L), LE), Myanmar].

    Mallotus esquirolii H.Lév., Repert. Spec. Nov. Regni Veg. 9 (11 Aug. 1911) 461, non H.Lév., Repert. Spec. Nov. Regni Veg. 9 (10 May 1911), (see Lauener, Notes Roy. Bot. Gard. Edinburgh 49, 1983, 482). — Mallotus leveilleanus Fedde, Repert. Spec. Nov. Regni Veg. 10 (1911) 144 (‘Léveillanus’). — Mallotus leveillei Fedde ex H. Lév., Fl. Kouy-Tchéouchéou (1914) 165, nom. superfl. — Type: Esquirol 120 (holo E; photo in A), China, Kweichow, Ouang-Mou.

    Mallotus barbatus Wall. ex Müll.Arg. var. congestus F.P.Metcalf, Lingnan Sci. J. 10 (1931) 487 (‘congesta’). — Mallotus lotingensis F.P.Metcalf, J. Arnold Arbor. 22 (1941) 206. — Lectotype (Sierra & Welzen, 2005): Tsiang 1131 (holo SYS; iso A, NY), China, Kwangtung, San Kai Tin.

    Mallotus barbatus Wall. ex Müll.Arg. var. pedicellaris Croizat, J. Arnold Arbor. 19 (1938) 135. — Type: Fang 680 (holo A), China, Szechuan, Chung-hsien.

    Mallotus croizatianus F.P.Metcalf, J. Arnold Arbor. 22 (1941) 204. — Mallotus barbatus Wall. ex Müll.Arg. var. croizatianus (F.P.Metcalf) S.M.Hwang, Acta Phytotax. Sin. 23 (1985) 295.— Type: Taam 4 (holo SYS; iso A), China, Kwangsi, Pai-shou, Liang-chiang.

    Mallotus luchenensis F.P.Metcalf, J. Arnold Arbor. 22 (1941) 206. — Type: Ching 5699 (holo SYS; iso A, NY), China, Kwangsi, Shan Fang, N. Luchen.

    Mallotus barbatus Wall. ex Müll.Arg. var. hubeiensis S.M.Hwang, Acta Phytotax. Sin. 23 (1985) 296. — Type: Li 7620 (holo SCIB, n.v.), China, Hubei.

    Mallotus barbatus Wall. ex Müll.Arg. var. wui H.S.Kiu in H.T.Chang & H.S.Kiu, Guihaia 23 (2003) 99. — Type: Exped. Yue 74-4586 (holo SCIB, n.v.), China, Guandong, Fengkai, Qixing.

Shrubs to small trees up to 12 m high, dbh up to 15 cm, monoecious or dioecious. Outer bark finely striate, smooth, up to 6 mm thick, greyish with patches of brown, on cross section reddish brown; sapwood straw coloured with pinkish brown streaking. Indumentum densely hairy, very soft-floccose, flocci up to 6 mm long. Stipules linear-triangular, 10–17 by 0.8–1.2 mm, persistent, margin subentire, apex acuminate. Leaves alternate to apically subopposite; petiole 3–40 by 2–7 mm, blade peltate for 10–80 mm, broadly ovate to ovate, 10.5–58 by 9–45 cm, length/width ratio 1–1.3, base truncate, rounded or obtuse, margin dentate, sometimes 2-lobed at widest part of blade, longest lobes up to 50 mm, upper surface pinkish red when young, dull green when old, basally with 0–4 extrafloral nectaries, 1.2–4.5 by 0.7–2 mm, marginal nectaries 0–9 per side, 2–6(–18) mm from margin, 0.7–1 by 0.5–1 mm, lower surface brownish green, palminerved, nerves 7–11 per side, mostly ending in the margin. Inflorescences terminal, unisexual or bisexual, axes greenish tawny, basally 2–5 mm thick; bracts 5–20 by c. 1 mm, bracteoles linear triangular 2.7–4.3 by 0.4–0.6 mm, persistent, margin subentire. Staminate inflorescences up to 65 cm long, side branches up to 30 cm long, with 2 or 3 flowers per node, nodes per branch up to 130. Staminate flowers 6–9 mm diam.; pedicels 3–5.5 mm long; sepals 4 or 5, 3–5 by 1.3–3 mm, pale light cream to tan cream; stamens 60–85, pale light green to yellow, filaments 1–4 mm long, anthers 0.3–0.4 by 0.2–0.3 mm, pale light yellow; pistillode absent (stamens absent from middle of flower, centre sometimes with reduced wart-like appendices). Pistillate inflorescences racemes, up to 55 cm long, occasionally with side branches, up to 10 cm long, nodes up to 80. Pistillate flowers 4–6 mm diam.; pedicels 1–45 mm long; calyx (3-) or 4- or 5- or (6-)lobed, connate on the base, 2.5–5 mm long, lobes 2–3 by 1.5–2.5 mm; ovary 3(–5)-locular, 2.5–3.5 by 2.2–3 mm, spines not individually visible; style up to 1.5 mm long; stigmas 2–6 mm long, yellow; staminodes sometimes present, also persistent in fruits. Fruit 10–21 by 14–20 mm, with strong smell, yellow with reddish touch, spines numerous, straight, thin, less hairy, up to 7 mm long, hairs forming a continuous layer, rubbing off; column 7–9 by 1–1.5 mm. Seeds ellipsoid, 4.8–5.5 by 3–4.5 by 2–4 mm, surface smooth, black; hilum c. 1 by c. 1.3 mm.

    Distribution — From India (W Bengal) to South China, Southeast Asia mainland, Malay Peninsula, Sumatra, and Java.

    Habitat & Ecology — Locally common in deciduous and evergreen forest, mostly in open, often very disturbed or burned places, along road and river sides; on a large variety of soil types, like shale, limestone, sandstone, and sandy clay. Altitude: sea level up to 1400 m. Flowering and fruiting the whole year through.

    Uses — The roots and fruits are used against muscle stiffness. The seeds yield a fatty oil used for making candles.

    Vernacular names —Malay Peninsula: Baleh anging, Ba-le-a-nging, Balek angin, Balik angin, Tampun (Malay).

4. Mallotus blumeanus Müll.Arg. (sect. Rottleropsis)

    Mallotus blumeanus Müll.Arg., Linnaea 34 (1865) 195; Backer & Bakh.f., Fl. Java 1 (1964) 483; Airy Shaw, Kew Bull. 36 (1981) 325; S.E.C.Sierra et al., Blumea 52 (2007) 48, map 3. — Rottlera oppositifolia Blume, Bijdr. (1826) 608, non Mallotus oppositifolius (Geiseler) Müll.Arg. — Plagianthera oppositifolia (Blume) Rchb.f. & Zoll., Acta Soc. Regiae Sci. Indo- Neerl. 1 (1856) 20. — Rottlera blumeana (Müll.Arg.) Scheff., Ann. Mus. Bot. Lugduno-Batavum 4 (1869) 124. — Lectotype (S.E.C.Sierra et al., 2007): Blume 1366, pro fem. (holo L, barcode L0291898; iso NY), Java, Salak, Parang.

Small trees, 13–23 m high; bole 4–7 m high, dbh 15–60 cm, dioecious, crown 7–14 m high. Outer bark smooth. Indumentum composed of stellately-tufted hairs and yellow to orange glandular hairs. Stipules narrowly triangular, 3.5–4 by 0.8–1.2 mm, early caducous, margin entire, apex acute, densely hairy. Leaves opposite; petiole 12–75 by 1–2.5 mm, sparsely to scatteredly hairy; blade ovate to elliptic, sometimes slightly falcate, 13–33 by 5–15 cm, length/width ratio 2.1–2.8, chartaceous, base rounded, obtuse to cuneate, margin denticulate to crenulate, sometimes with glandular teeth, apex acuminate to caudate, upper surface scatteredly hairy, larger extrafloral nectaries basal near the petiole insertion, 1–3 per side, elliptic, 1.2–2 by 0.5–1 mm, smaller ones marginal in upper half, 5–14 nectaries per side, 0.8–2.8 mm from margin, orbicular, 0.2–0.3 by 0.2–0.3 mm, lower surface sparsely to scatteredly hairy, scatteredly covered with glandular hairs (sometimes only present towards the leaf margin), domatia absent, venation triplinerved, nerves 6–9 per side, looping and closed near margin. Inflorescences racemes, single, erect; peduncle 10–15 by 1–2 mm long; axes densely to sparsely hairy; bracts triangular, 1.5–3 by 0.8–1 mm, persistent to caducous, margin entire, apex caudate, densely to sparsely hairy; bracteoles absent. Flowers: pedicels densely hairy; sepals basally connate, persistent, densely to sparsely hairy outside, glabrous inside, glandular hairs absent, margin entire, apex acute. Staminate inflorescences 7.5–20 cm long, nodes per branch 33–45, with 6–8 flowers per bract. Staminate flowers 3–3.3 mm diam.; pedicels 1.5–7 mm long; calyx (rarely 2- or) 3-lobed, lobes ovate to elliptic, 1.8–2.5 by 1–1.5 mm; stamens 28–40, glabrous, filaments 0.25–1.5 mm long, flattened, free, thecae ovoid, 0.1–0.2 by 0.1–0.2 mm, connective umbrella-like; pistillode absent. Pistillate inflorescences 8.5–16 cm long, nodes per branch 19–70, with 1 flower per bract. Pistillate flowers 3–4 mm diam.; pedicels 1.3–2 mm long; calyx 3–5-lobed, lobes triangular to narrowly triangular, 2.5–3 by 1.2–1.8 mm; ovary 1.8–2 by 1.5–1.8 mm, 3-locular, densely hairy, glandular hairs absent; style 0.2–0.5 mm long; stigmas 2–3 by 1–1.5 mm. Fruits drupes, ellipsoid, 10–12 by 4–4.4 mm, surface smooth, sparsely hairy, glandular hairs rarely present; wall: woody part c. 0.1 mm thick, endocarp forming a thick fleshy layer around the seeds; column absent. Seeds lenticular, 5.5–7 by 5–7 by 3–4 mm, surface rugose, dull, black.

    Distribution — Sumatra, Java.

    Habitat & Ecology — In primary or old secondary and riparian forest; on ferrosol-loam or limestone. Altitude 10–1000 m. Flowering and fruiting throughout the year.

    Vernacular names — Sumatra: Entatau, Kijak tukau, Medang tekuku. Java: Ki tendjo, Ki tengi (Sundanese); Daon kajuole, Hantap batu, Pingku.

    Notes — 1. Mallotus blumeanus can be easily confused with M. sphaerocarpus, but these two species differ in their indument and fruits. Mallotus blumeanus has brown yellow hairs, usually appressed (yellow and erect in M. sphaerocarpus) ellipsoid and densely hairy fruits (spheroid and sparsely hairy in M. sphaerocarpus).

2. Airy Shaw (1982) mentions that M. blumeanus occurs in Sulawesi and the Lesser Sunda Islands. This is probably based on, among other collections, Schmutz 4133 (L), from Flores (Lesser Sunda Islands). However, this is a misidentification, in Kulju et al. (2007) it is determined as M. cf. cumingii Müll.Arg.

5. Mallotus brachythyrsus Merr. (ex sect. Hancea)

   Mallotus brachythyrsus Merr., Sarawak Mus. J. (1928) 526; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 162; Slik & Welzen, Blumea 46 (2001) 49, Fig. 19, Map 8; in Slik, Tropenbos-Kalimantan ser. 4 (2001) 194, Fig. 7.19, Map 7.8; Welzen et al., Blumea 51 (2006) 369. — Type: Mjoberg 147 (PNH holo, lost; BM, UC iso), Borneo, Sarawak, Mt Poi.

   Mallotus beccarii Airy Shaw, Kew Bull. 27 (1972) 86; Kew Bull. Add. Ser. 4 (1975) 162. — Type: Beccari 1800 (K holo), Borneo, Sarawak, First Division, Mt Matang.

Also check: Slik, Macaranga and Mallotus species of Borneo

Shrubs up to 3 m tall, dioecious. Indumentum glabrous to dense, whitish-yellowish, simple to tufted to stellate. Branches lenticellate, glabrescent, gland-dotted or not. Stipules caducous, narrowly triangular to ovate, 7–10 by 1.3–2 mm, margin entire to irregular, involute, apex acute, glabrous to hairy abaxially, glabrous adaxially, gland-dotted or not. Leaves simple, opposite, strongly unequal, one of each pair reduced, stipuliform. Non-reduced leaves: petioles 2–5 by 0.8–1 mm, hairy, glabrescent, gland-dotted or not; blade ovate to obovate, 6–20 by 2–7.5 cm, length-width ratio 2.2–3.4, chartaceous to coriaceous, base acute to cuneate to slightly cordate, usually oblique, margin (slightly) dentate, marginal glands 7–17 per leaf side, apex acuminate to cuspidate, upper surface smooth, red-brown when dried, glabrous, densely gland-dotted (or not), basal macular glands 2–4 on veins, 0.5–3 mm from midrib, apical macular glands 0–9 on veins, 1–7 mm from margin, lower surface smooth, creamish when dried, hairy, especially on midrib and veins, gland-dotted or not, domatia absent or present, with hair tufts; venation pinnate, 9–13 secondary veins per side, ending parallel to or in the margin. Stipuliform leaves ovate to narrowly triangular, 2.5–12 by 1.5–2 mm, margin entire to irregular, apex acute, glabrous to hairy, glabrous adaxially, gland-dotted or not. Inflorescences in axils of stipuliform leaves, unbranched, solitary. Flowers actinomorphic, small. Staminate inflorescences raceme-like thyrses, up to 2–2.6 cm long, basally c. 1 mm thick, hairy, gland-dotted, nodes up to 11 per inflorescence, flowers 1–6 per node; bracts persistent, (broadly) ovate, 1–2 by 0.9–1.5 mm, margin irregular, apex acute, hairy abaxially only, gland-dotted or not; buds globose (to slightly ovate), hairy, gland-dotted. Staminate flowers up to 3 mm in diameter, greenish to cream; bracteoles absent; pedicels c. 1 mm long, hairy, gland-dotted; sepals 3, ovate, 1.5–2 by 0.7–1 mm, margin entire, apex acute, hairy abaxially only, gland-dotted outside only; stamens up to c. 60; filaments up to 1 mm long, glabrous; anthers c. 0.2 by 0.3 mm, basifixed; thecae 2, parallel, extrorse, opening lengthwise, glabrous; connective not (to slightly) widened. Pistillate inflorescences racemes, up to 5 cm long, basally c. 1 mm thick, hairy, gland-dotted, nodes up to 4 per inflorescence; bracts persistent, ovate, clasping the pedicel, 1.8–2 by 1–1.2 mm, margin irregular, apex acute, hairy abaxially only. Pistillate flowers pedicels 5–6 mm long, hairy, gland-dotted or not; sepals 3, ovate, 2.5–5 by 1.3–2.8 mm, margin irregular, apex acute, hairy abaxially only, gland-dotted; ovary 3-locular, echinate, densely hairy, gland-dotted, spines 0.6–0.8 mm long, glabrous to hairy, not gland-dotted; ovules 1 per locule, axillary, apotropous, ascending; styles persistent, up to 1 mm long, hairy, gland-dotted; stigmas 3–4 mm long, densely covered with granulate papillae on the inside, hairy abaxially, gland-dotted outside. Fruit an echinate, lobed capsule, 5–7 by c. 5 mm, loculicidal-septicidal, green, hairy, not gland-dotted; carpel inside glabrous (to hairy), not gland-dotted; column c. 4 by 4–5 mm. Seeds glossy, roughly globose, c. 5 by 4–5 by 4.5–5 mm; hilum elliptic to deltoid, c. 1.2 by 0.5–3 mm.

    Distribution — Borneo (Sarawak, Central Kalimantan).

    Habitat & Ecology — In primary and secondary forest; on yellow sandy clay soils. Altitude: up to 500 m.

    Notes — The plants can range from almost completely glabrous to conspicuously hairy. Airy Shaw (1972) described the species M. beccarii, which differs from M. brachythyrsus in being very hairy as opposed to nearly glabrous. However, all characters, except for the glabrescence, are variable enough to include M. beccarii as a synonym of M. brachythyrsus. The specimen collected in Central Kalimantan differs from the specimens known from Sarawak in not being gland-dotted on most parts.

6. Mallotus brevipetiolatus Gage (sect. Rottleropsis)

    Mallotus brevipetiolatus Gage, Rec. Bot. Surv. India 9 (1922) 242; Ridl., Fl. Malay Penin. 3 (1924) 286; Airy Shaw, Kew Bull. 21 (1968) 393; Welzen et al., Thai Forest Bull., Bot. 28 (2000) 97; S.E.C.Sierra et al., Blumea 52 (2007) 50, map 4. — Lectotype (S.E.C.Sierra et al., 2007): Ridley 15222 (holo K; iso BO, SING), Peninsular Malaysia, Perlis, Bukit Lagi.

Shrubs to small trees, up to 8 m high, dioecious. Outer bark smooth. Indumentum composed of simple, tufted and rarely peltate hairs and yellow to orange glandular hairs. Stipules triangular, 2.5–3.5 by 1.3–2.2 mm, early caducous, margin subentire, apex acute, scatteredly hairy. Leaves opposite; petiole 2–6 by 1–2 mm, scatteredly hairy to glabrous; blade elliptic to obovate, 4–21.5 by 2.2–8.1 cm, length/width ratio 2.5–3.8, chartaceous, base shallowly emarginate to cuneate, margin denticulate to crenulate, with glandular teeth (located in a minute notch), apex acuminate to caudate, upper surface glabrous, sparsely to scatteredly covered with glandular hairs, extrafloral nectaries inconspicuous, touching the midrib to marginal in lower half, 1–3 per side, 2–8.5 mm from margin, elliptic to orbicular, 0.6–1.2 by 0.2–0.4 mm, lower surface glabrous, scatteredly covered with glandular hairs, domatia absent, venation pinnate, nerves 8–16 per side, looping and closed near margin. Inflorescences racemes, single, erect; peduncle 2–12 by 0.9–1.2 mm; axes sparsely hairy; bracts scatteredly hairy to glabrous, deltoid or triangular, 1–1.7 by 0.8–1.4 mm, persistent, margin subentire, apex acute, scatteredly hairy to glabrous; bracteoles absent. Flowers: pedicels scatteredly hairy; sepals free, persistent, margin entire, apex acute, scatteredly hairy to glabrous outside, glabrous inside, with glandular hairs outside. Staminate inflorescences 7–22 cm long, nodes per branch 9–68, with 6–9 flowers per bract. Staminate flowers 3.5–5 mm diam., with sweet scent; pedicels 1.5–7 mm long; sepals 3 or 4, elliptic to obovate, 2–3 by 1.2–2 mm; stamens 20–35, filaments 1–3 mm long, free, glabrous, thecae ellipsoid, 0.3–0.4 by 0.2–0.25 mm, connnective not widened; pistillode absent. Pistillate inflorescences 4.7–10 cm long, nodes per branch 7–11, with 1 flower per bract. Pistillate flowers 2.5–3 mm diam.; pedicels 1–3 mm long; sepals 4–6, linear-triangular, 1.2–4.3 by 0.5–0.8 mm; ovary 1–1.4 by 1.3–1.6 mm, 3-locular, scatteredly hairy, densely covered with glandular hairs; style 0.5–0.7 mm long; stigmas 1.3–3.5 by 0.8–1 mm. Fruits capsules, 5.8–8.9 by 9–13 mm, opening septicidally-loculicidally, surface spiny, scatteredly hairy, sparsely to scatteredly covered with glandular hairs, spines 50–70, curved, 1.5–3 by 0.2–0.3 mm, apex hairy; wall 0.8–1 mm thick, sparsely hairy to glabrous inside; column 4.3–5 by 4.8–7 mm. Seeds ± globose, 4.5–6.8 by 3.8–5.9 by 4.5–6 mm, surface smooth, shiny, dark brown.

    Distribution — Thailand, Peninsular Malaysia.

    Habitat & Ecology — In evergreen or mixed evergreen deciduous forest; on hill sides and shaded areas; on clay or limestone soil. Altitude 10–470 m. Flowering and fruiting throughout the year. Staminate flowers are visited by two genera of bees, Trigona and Apis dorsata.

7. Mallotus caudatus Merr (sect. Rottleropsis)

    Mallotus caudatus Merr., Philipp. J. Sci., Bot. 13 (1918) 83; J. Straits Branch Roy. Asiat. Soc., Special number (1921) 338; Pl. Elmer. Born. (1929) 157; Welzen & S.E.C.Sierra, Blumea 51 (2006) 376, fig. 1b, 2b, 3j, k; map 1. — Type: Villamil 376 (holo PNH, †, photo in A; iso K), British North Borneo (Sabah), Marauti watershed near Tawau.

    Mallotus wrayi auct. non King ex Hook.f.: Airy Shaw, Kew Bull. 16 (1963) 349, p.p. (M. caudatus); Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 52, p.p. (M. caudatus); Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 171, p.p. (M. caudatus).

Shrubs to trees, up to 8(–15) m high, clear bole up to 10 m high, dbh up to 8 cm; flowering branchlets 3–5 mm thick, flattened and somewhat ridged when dry, hairy, glabrescent. Outer bark smooth, (green to) white to white-brown to grey, to spotted red to dark brown, c. 0.5 mm thick; inner bark whitish to yellowish to green to brownish (to black?), c. 2 mm thick; sapwood medium hard to very hard, fairly fine grained, white to yellow to pale orange to brown. Indumentum present on most parts, stiff simple hairs and stellate groups of very soft and thin whitish hairs on the leaf blades, glabrescent; glandular scales yellowish to orangey to dark brown. Stipules and bracts of terminal buds triangular, 4–5.5 by 1.7–2 mm, hairy, caducous. Leaves opposite, similar shape, one usually somewhat smaller; petiole 1.3–8 cm long, round, somewhat hairy, basally and especially apically strongly pulvinate, upper pulvinus oblique; blade partly over upper pulvinus (pseudo-subpeltate), ovate to elliptic, 3.9–44.5 by 1.8–11 cm, length/width ratio 2.2–6, symmetric, membranous to subcoriaceous; base cuneate, very base narrowly cordate, margin entire or with one to few glandular teeth per margin or laxly denticulate, usually somewhat undulate, apex cuspidate to caudate; upper surface with stellate soft hairs when young, basally one pair of extrafloral nectaries on basal nerves, elliptic, c. 1 mm diam., additional extrafloral nectaries often present (up to 9 pairs) along the basal part of the midrib (sometimes only in some leaves) and in the apical third, a few cm from the margin on the nerves and larger veins, circular, c. 0.3 by 0.3 mm; lower surface smooth, not to somewhat lighter green than upper surface, subglabrous to mainly hairy on the midrib and (basal) nerves, hair tuft domatia present, but not very distinct, glandular scales present, orangey yellowish; venation triplinerved, slightly raised above, raised beneath and of lighter colour than lamina, nerves 7–9 per side up to the apex, looped and closed near margin, veins scalariform, veinlets laxly reticulate to almost scalariform. Inflorescences racemes, axillary to terminal, single to seven together when staminate, greenish brown to white, hairs mainly straight, patent, ferruginous, glabrescent; staminate ones up to 19 cm long, 0.5–2.2 mm thick, with cymosely flowering groups of more than 15 flowers per node, pistillate ones up to 26 cm long, 1.7–2.5 mm thick, with few flowers, one per node; bracts ovate to elliptic, (2.3–)3.3–10 by 1.3–3 mm, outside hairy, inside glabrous. Staminate flowers 2–4.5 mm diam., green to white to yellow, fragrant; buds mainly obovoid; pedicel 2.2–3.4 mm long, hairy; sepals 3 or 4, ovate to elliptic, 2.5–3 by 1.5–2 mm, reflexed, hairy, especially outside, glandular scales only outside; stamens (24–)35–60, filaments up to 2.6 mm long, glabrous, anthers c. 0.3 by 0.2 mm, thecae separated from each other by connective, yellow, top of connective with a few large cells (perhaps glandular). Pistillate flowers 7–7.5 mm diam., greenish to white to (greenish) yellow; pedicel up to 3 mm long with an abscission zone subbasally; sepals splitting variously, (2–)4 or 5 (or 6), ovate, 3–5 by 1.3–3 mm, mainly hairy outside, glandular scales only outside; ovary 3-locular, 1.2–2.5 by 1.8–2 mm, spines up to 1 mm long, dense, in 6 groups (2 per locule), tomentose, purple; style 1.3–3 mm long, stigmas up to 9 mm long, 0.6–1 mm wide, densely papillate above with branching papillae more than 1 mm long, abaxially densely (yellowish when dry) stellately tomentose. Fruits lobed capsules, (11–)12.5–17 by 7–10 mm, reddish green to white (to light blue?), spines up to 4 mm long, tomentose, breaking off; wall woody, inner part c. 1.2 mm thick; column T-shaped, up to 4 mm high. Seeds globose, 6–7.5 mm diam., brown to black, shiny.

    Distribution — Borneo. Most specimens either show soft hairs or stiff hairs (but many are more or less in between). The soft-haired ones usually have additional extrafloral nectaries along the basal part of the adaxial midrib. Initially it seemed that these forms were geographically separate, but Map 1 shows this not to be the case.

    Habitat & Ecology — In understorey of primary and dipterocarp forest, riverine forest, kerangas forest, selectively logged forest, mainly on undulating land and ridges of hills; soil yellow to brown sand or red clay, laterite; bedrock: Belait Series, sandstone, Setap Shales. Altitude: sea level up to 500(–1100) m. Flowering and fruiting: throughout the year through. Flowers much visited by bees.

    Vernacular names — Brunei: Bantas (Brunei, Iban); Enserai (Iban). Kalimantan: Kroti. Sabah: Babalitak (Tidong); Balek balek angin bini (Brunei); Bongbong, Mingkig pateh (Dusun); Labah, Lambukan, Limbukan, Minungbong (Dusun Kinabatangan); Kandang jagong (Kadayan); Kemenyan (Suluk); Kolokos (Tengara); Lamba; Romi romi; Singa kapot; Tola tola, Toloo toloo (Sungai); Rendang jagong, sagar-sagar (Malay). Sarawak: Bantas, Retih (Iban); Sekayah (Kenyah).

    Uses — Wood is excellent firewood.

    Note — SAN (Amin et al.) 69332 is exceptional in having pistillate flowers with staminodes.

8. Mallotus cauliflorus Merr. (sect. Rottleropsis)

    Mallotus cauliflorus Merr., Philipp. J. Sci., Bot. 7 (1912) 399; S.E.C.Sierra et al., Blumea 52 (2007) 52, map 5. — Type: BS (M. Ramos) 13909 (holo PNH, †; iso A, CAHUP, K, US), Philippines, Luzon, Cagayan.

Woody, dioecious. Indumentum composed of simple and tufted hairs and yellow to orange glandular hairs. Stipules narrowly triangular, 18–30 by 8–10 mm, persistent, margin entire, apex acute, densely hairy, glabrescent. Leaves opposite; petiole 35–90 by 2–2.5 mm, scatteredly hairy to glabrous; blade elliptic to obovate, 16–31 by 6–12 cm, length/width ratio 2.3–2.5, coriaceous, base rounded, obtuse to cuneate, margin entire to slightly repand, with glandular teeth, apex acuminate to caudate, upper surface glabrous, sparsely covered with glandular hairs, extrafloral nectaries at the axils between midrib and nerves, from base to apex, 1–20 per side, 3–60 mm from margin, elliptic to orbicular, 0.5–1 by 0.5–0.8 mm, lower surface sparsely hairy to subglabrous, densely covered with glandular hairs, domatia sometimes present, venation triplinerved, nerves 10–15 per side, looping and closed near margin. Inflorescences axillary, ramiflorous or cauliflorous racemes, 1–3 together, erect; axes sparsely hairy, with 1 flower per bract; bracts triangular, margin subentire, densely hairy outside, glabrous inside; bracteoles absent. Flowers: pedicels scatteredly hairy; sepals 3, persistent, margin entire, apex acute, sparsely hairy outside, glabrous inside, with glandular hairs. Staminate inflorescences up to c. 10 cm long; peduncle 12–28 mm long; nodes per branch 36–60; bracts 0.6–0.9 by 0.4–0.6 mm, apex acute. Staminate flowers c. 3 mm diam.; pedicels 2.5–5 mm long; sepals 3, elliptic, 2.5–3 by 1.5–2 mm, free; stamens 35–42, glabrous, filaments 0.5–2.8 mm long, free, thecae ellipsoid, 0.2–0.3 by 0.2–0.25 mm, connective umbrella-like; pistillode absent. Pistillate inflorescences 7.5–12.5 cm long; peduncle 28 mm long; nodes per branch c. 13; bracts 3–4 by 1.5–2 mm, apex acute. Pistillate flowers 3–4 mm diam.; pedicels 0.2–0.4 mm long; calyx 3-lobed, lobes triangular, 4–5 by 1–3 mm, basally connate; ovary 2–2.5 by 2–2.5 mm, 3-locular, sparsely hairy, densely covered with glandular hairs; style 0.3–0.5 mm long; stigmas 3–4 by 0.8–1 mm. Fruits capsules, 9–11 by 9–11 mm, opening septicidally-loculicidally, surface spiny, sparsely hairy, densely covered with glandular hairs; spines 90–120, curved, 0.5–1 by 0.1–0.2 mm, sparsely hairy; wall c. 0.8 mm thick, glabrous inside; column 3.5–5 by 1.5–2.5 mm. Seeds globose, 4.5–5 by 4–4.5 by 4–4.5 mm, surface smooth, dull, light brown.

    Distribution — Endemic to the Philippines.

    Habitat & Ecology — Altitude c. 80 m. Flowering and fruiting: January to December.

9. Mallotus chromocarpus Airy Shaw (sect. Rottleropsis)

    Mallotus chromocarpus Airy Shaw, Kew Bull. 32 (1978) 403; Kew Bull. 33 (1978) 64; Kew Bull., Addit. Ser. 8 (1980) 164, 225; S.E.C.Sierra, Welzen & Slik, Blumea 50 (2005) 238, fig. 6, map 4. — Type: NGF (McDonald) 8204 (holo K; iso A, BRI, L), Papua New Guinea, Central Province, Mori River.

    Mallotus discolor auct. non (F. Muell.) F. Muell. ex Benth.: Airy Shaw, Kew Bull. 20 (1966) 44.

Trees up to 30.5 m high, dbh up to 40 cm; bole up to 18 m high; crown up to 9.5 m long; branchlets glabrescent. Outer bark rough, long and transversely fissured to smooth, up to 19 mm thick, greyish with patches of brown, on cross section reddish; sapwood cream; heartwood straw-coloured. Indumentum tomentose, composed of stellate hairs, and sessile, globular to disc-shaped, orange glandular hairs. Stipules absent. Leaves alternate, simple, symmetric; petiole 25–80 by 1–2 mm, glabrescent; blade ovate to elliptic, 6–18 by 5.3–14 cm, length/width ratio 1.3–1.8, coriaceous, base rounded to cuneate, peltate for 3 mm, margin entire, apex caudate, upper surface dull green, glabrous, basally with 2 (or 4) extrafloral nectaries on the nerves near the petiole insertion, elliptic, 2–3 by 0.7–1 mm, marginal nectaries 0–6 per side, elliptic, 0.5–1 by 0.3–0.5 mm, lower surface greenish grey to brownish grey, not glabrescent, domatia present, woolly, venation prominent, 3-nerved, nerves 4–6 per side, looping, veinlets reticulate. Inflorescences racemes, axillary (1–3 together) or terminally grouped, erect, unisexual, axes basally c. 1 mm thick; bracts triangular, persistent, margin entire, airy on both sides; buds light green. Flowers actinomorphic, not exceeding 1 cm diam.; pedicels hairy; sepals persistent, valvate, densely hairy outside, subglabrous to sparsely hairy inside, with glandular hairs on both sides, ovate, margin entire, apex acute; petals and disc absent. Staminate inflorescences up to 15.5 cm long, flowers 3–5 per node, nodes up to 40; bracts 0.6–0.7 by 0.3–0.5 mm, apex acuminate. Staminate flowers 2.8–3.5 mm diam.; pedicels 1.5–2 mm long; sepals (2 or) 3, 1.8–2.2 by 1.7–2 mm, free; stamens 30–36, glabrous, filaments 0.6–1 mm long, free (to basally connate), anthers ovoid, basifixed, 0.5–0.6 by 0.5–0.6 mm, often at the apex with glandular hairs, light yellow, connective broad (umbrella-like); pistillode absent. Pistillate inflorescences up to 10.5 cm long, flowers 1 per node, nodes up to 13; bracts 0.6–0.8 by 0.4–0.6 mm, apex acute. Pistillate flowers 2–2.5 mm diam., pedicels 2–4 mm long, hairy, gland dotted; sepals (3 or) 4 (or 5), 1.5–2 by 1–1.2 mm, slightly connate at base; ovary (3- or) 4- or (5-)locular,1.5–1.8by1.8–2mm,withglandularhairs;stigmassessile,plumose,1–1.2 mm long, densely covered with papillae above, outer surface hairy and with glandular hairs. Fruits indehiscent drupes, 5–8 by 8–10 mm, with (3 or) 4 (or 5) distinctly long ridges, orange, sparsely warted and hairy, densely gland-dotted; wall c. 0.5 mm thick, glabroustosparselyhairyinside;column4–4.2by1.2–1.3mm. Seeds ±globose,4–4.2 by 3.7–4 by 3.6–3.8 mm, glossy, brown; hilum 3.5–3.8 by c. 1 mm.

    Distribution — Endemic in New Guinea.

    Habitat & Ecology — Locally common in the canopy of young to old secondary forest, or on the grassland-edge of swamp-forest. Altitude: sea level up to 150 m. Flowering and fruiting: March to October.

    Vernacular name — New Guinea: Osari (Orokaiva-Mumuni).

10. Mallotus claoxyloides (F.Muell.) Müll.Arg. (sect. Rottleropsis)

    Mallotus claoxyloides (F.Muell.) Müll.Arg., Linnaea 34 (1865) 192; Airy Shaw, Kew Bull. 20 (1966) 42; Kew Bull., Addit. Ser. 8 (1980) 165; P.I.Forst., Austrobaileya 5 (1999) 460, 493; S.E.C.Sierra et al., Blumea 52 (2007) 53, map 2. — Echinocroton claoxyloides F.Muell., Fragm. Phytogr. Austr. 1 (1858) 32. — Echinus claoxyloides (F. Muell.) Baill., Adansonia 6 (1866) 315, incl. var. genuina Baill., nom. inval. — Type: F. von Mueller s.n. (‘Hill & Mueller’) (holo MEL barcode 708383; iso, GH, K), Australia, Queensland, Moreton, Brisbane River.

    Rottlera affinis Baill., Étude Euphorb. (1858) 36, t. 19, f. 29–31, 424, non Hassk. (1842). — Lectotype (S.E.C.Sierra et al., 2007): Verreaux 592 (holo P; iso P), Australia.

    Echinus claoxyloides (F.Muell.) Baill. var. cordata Baill., Adansonia 6 (1866) 315. — Mallotus claoxyloides (F.Muell.) Müll.Arg. var. cordatus (Baill.) Airy Shaw, Muelleria 4 (1980) 232. — Lectotype (P.I. Forster, 1999): Beckler s.n. (holo MEL barcode 515956 n.v.; iso P n.v.), Australia, New South Wales, Richmond River.

    Mallotus claoxyloides (F.Muell.) Müll.Arg. var. glabratus Domin, Biblioth. Bot. Band 22, Heft 89, Abt. 3 (1927) 888. — Type: Dietrich 524 (holo PR; iso AD, CANB, L, MEL), Australia, Queensland, S Kennedy, Port Mackay.

Shrubs to small trees, 0.5–8 m high, dioecious or rarely monoecious. Indumentum composed of simple and stellately-tufted hairs and yellow to orange glandular hairs. Stipules triangular to linear-triangular, 1.8–6 by 0.6–1 mm, persistent to caducous, margin entire, apex acute, scatteredly hairy to glabrous. Leaves opposite to rarely alternate; petiole 2–60 by 0.8–1 mm, glabrescent; blade elliptic to obovate, 4–17 by 2.5–9 cm, length/width ratio 1.6–2.1, (sub)coriaceous, base rounded, obtuse, cuneate to shallowly emarginate, margin entire, (denticulate), dentate to crenate, with glandular teeth, apex rounded to acuminate, upper surface sparsely hairy to glabrous on midrib and nerves, extrafloral nectaries marginal in lower half, 1–3(–5) per side, 1–2 mm from margin, elliptic to orbicular, 0.5–1 by 0.4–0.6 mm, lower surface sparsely hairy, glabrescent, sparsely covered with glandular hairs, domatia sometimes present, venation inconspicuously triplinerved, nerves 7 or 8 per side, indistinctly looping or ending in margin. Inflorescences single, erect; axes densely to sparsely hairy; bracts narrowly triangular, persistent, apex acute, margin entire, densely hairy outside, scatteredly hairy inside; bracteoles narrowly triangular 0.5–1 by 0.2–0.3 mm, early caducous. Flowers: pedicels densely hairy; sepals free, persistent, margin entire, apex acuminate, densely hairy outside, glabrous inside, with glandular hairs outside. Staminate inflorescences reduced racemes, 3.5–4 cm long; peduncle 7–20 mm long; nodes per branch 3–5, with 6–10 flowers per bract; bracts 1.5–4.5 by 0.3–1 mm, apex acuminate. Staminate flowers 5–6.5 mm diam.; pedicels 2–5 mm long; sepals (2 or) 3, elliptic to obovate, 4–5 by 1.8–2.2 mm; stamens 29–45, filaments 1.8–5.5 mm long, free, glabrous, thecae ellipsoid, 0.2–0.3 by 0.2–0.25 mm, hairy or glabrous, connective widened; pistillode present. Pistillate inflorescences umbel-like, 0.8–1.5 cm long, usually with an extra flower near the base of the axes; peduncle 3–6 mm long; nodes per branch 1 or 2, with (1–)3–5 flowers per bract; bracts 1.5–2.8 by 0.2–0.4 mm, apex acute. Pistillate flowers 4–6 mm diam.; pedicels 3–10 mm long; sepals 3 or 4, triangular, 3–5 by 0.8–1.2 mm; ovary 1.8–2 by 1.5–1.8 mm, 3-locular, densely hairy, scatteredly covered with glandular hairs; style 0.5–1 mm, stigmas 3–3.5 by 1.2–1.5 mm. Fruits capsules, 6–8.5 by 11–13 mm, opening septicidally-loculicidally, surface spiny, densely hairy, scatteredly covered with glandular hairs; spines 65–80, curved, 1–1.3 by 0.1–0.2 mm, sparsely hairy; wall 0.8–1 mm thick, glabrous inside; column 4–5 by 3–5 mm. Seeds globose, 5–5.5 by 4–5 by 4–5 mm, surface smooth, dull, light-brown.

    Distribution — New Guinea, Australia.

    Habitat & Ecology — Locally common in edges of evergreen forests, semi-evergreen vine-thickets and mangroves; in araucarian notophyll and microphyll vineforest; along dried gullies and creeks; on sedimentary, coral sand, limestone, rocky, sandy loam or basalt soil. Altitude 20–500 m. Flowering and fruiting throughout the year.

    Notes — 1. See note under M. attenuatus.

2. According to label information, the fresh leaves emit a strong smell of benzaldehyde.

11. Mallotus concinnus Airy Shaw (ex sect. Hancea)

    Mallotus concinnus Airy Shaw, Kew Bull. 32 (1978) 406; Welzen et al., Blumea 51 (2006) 371. — ?Erismanthus sp.: M.R.Hend., J. Malayan Branch Roy. Asiat. Soc. 17 (1939) 70. — Type: SF (Henderson) 19543 (holo K; iso K, SING), Malaya, Kelantan, Gua Panjang, Gua Ninik.

Shrub, up to 3 m tall; branchlets terete, up to 3 mm thick. Indumentum sparse, very short simple or stellately bundled hairs and yellowish glandular scales. Stipules early caducous, triangular, 1.2–1.5 by 0.5–0.8 mm, margin entire, apex acute, outside with simple hairs. Leaves simple, opposite, unequal, one much smaller and often caducous. Non-reduced leaves: petiole 1–4 mm long, with simple hairs; blade elliptic to obovate, 6.8–13.2 by 2.5–5.2 cm, length/width ratio 2.5–2.7, pergamentaceous, symmetric, drying greenish or brownish, base cordate, symmetric, margin coarsely serrate to subserrate, teeth ending in glands, apex cuspidate to caudate, upper surface with basally a single or two pairs of small extrafloral nectaries on the basal nerves, glabrous, lower surface with simple hairs on the midrib and nerves, many glandular hairs and hair tuft domatia present; venation pinnate, nerves 7–9 per side, arching, looped and closed near margin, venation especially raised beneath. Reduced leaves subsessile, obcordate to reniform, 4–11 by 4–12 mm, margin laxly dentate, apex rounded to acute, upper surface without extrafloral nectaries, lower surface with glandular hairs; venation distinct on both sides. Staminate inflorescences and flowers unknown. Pistillate inflorescences single, axillary, up to 1.5 cm long, few-flowered, not densely hairy with simple hairs and glandular hairs; bracts broad and short, broken. Pistillate flowers c. 1.75 mm in diameter, white; pedicel 1–2 mm long, puberulous; sepals 4 (single badly preserved flower seen), ovate, c. 1.8 by 1.2 mm, revolute, glandular hairs inside and outside, outside laxly puberulous; ovary 3-locular, muricate, with a few short hairs and many glandular hairs; stigmas c. 2.5 mm long, plumose-papillate above. Fruits capsular, red (label information), not seen; column 4–5 mm long. Seeds not seen.

    Distribution — Peninsular Malaysia: Kelantan.

    Habitat & Ecology — On dry rocky summit in scrubby vegetation with wet humus and a dense ground cover of mainly orchids and ferns; soil: limestone. At 300–500 m altitude.

12. Mallotus connatus M. Aparicio (sect. Rottleropsis)

    Mallotus connatus M.Aparicio in S.E.C.Sierra et al., Blumea 52 (2007) 54,  fig. 4, map 6. — Type: Mahyar et al. 1197 (holo L; iso A, SING), Borneo, Kalimantan, Sintang, HPH Km 86.

    Mallotus moritzianus auct. non Müll.Arg.: Slik, Priyono & Welzen, Gard. Bull. Singapore 52 (2000) 60, f. 35.

    Mallotus rufidulus auct. non (Miq.) Müll.Arg.: Airy Shaw, Kew Bull. 36 (1981) 328, pro Borneo.

Shrubs to small trees, up to 9 m high; dioecious or rarely monoecious. Indumentum composed of simple, tufted and stellately-tufted hairs and yellow to orange glandular hairs. Stipules linear-triangular, 3–7 by 0.5–1 mm, early caducous, margin entire, apex acute, densely to sparsely hairy. Leaves opposite; petiole 5–70 by 1–1.8 mm, densely to sparsely hairy; blade ovate to elliptic, 7–27 by 5–14 cm, length/width ratio 1.5–3, membranaceous to chartaceous, base rounded, obtuse to cuneate, margin entire, apex caudate, upper surface densely to sparsely hairy on midrib and nerves, extrafloral nectaries marginal throughout blade, up to 20 nectaries per side, 1–8 mm from margin, sometimes also all over the blade, elliptic to orbicular, 0.2–0.5 by 0.2–0.5 mm, lower surface sparsely to scatteredly hairy, sparsely covered with glandular hairs, domatia absent, venation triplinerved, nerves 4–6(–9) per side, looping and closed near margin. Inflorescences racemes, 1 (or 2) together, erect, unisexual or rarely bisexual; peduncle 20–60 by 0.5–1.5 mm; axes sparsely hairy; bracts narrowly triangular or trilobed, 3–5 by 0.8–3 mm, persistent, margin entire, apex acute, densely hairy; bracteoles narrowly to linear-triangular, 2–3.8 by 0.3–0.5 mm, early caducous. Flowers: pedicels sparsely to densely hairy; sepals margin entire, apex acute, densely hairy outside, with scattered glandular hairs, glabrous inside. Staminate inflorescences 8–22 cm long, nodes per branch 11–13, with 5–13 flowers per bract. Staminate flowers 3–4 mm diam.; pedicels 1–4 mm long; calyx 3- or 4-lobed, lobes ovate, 3–5 by 0.4–0.6 mm; stamens 45–60, glabrous, filaments 3.5–5 mm long, connate in lower half, thecae ovoid, 0.2–0.3 by 0.2–0.25 mm, connective widened; pistillode present. Pistillate inflorescences 16–21 cm long, nodes per branch 3–10, with 1 flower per bract. Pistillate flowers 4–4.2 mm diam.; pedicels 3–7 mm long; sepals 4 or 5, triangular, 5–6.5 by 1–2 mm, free; staminodes rarely present; ovary 2.5–3 by 2.5–3 mm, 3-locular, densely hairy, green, scatteredly covered with glandular hairs; style 2–4 mm long; stigmas 3–7 by 0.8–1 mm. Fruits capsules, 10–15 by 10–16 mm, pale green to green, opening septicidally-loculicidally, surface spiny, densely hairy, scatteredly covered with glandular hairs; spines 200–250, ± straight, 1.5–3 by 0.2–0.3 mm, densely hairy; wall 0.8–1.2 mm thick, glabrous inside; column 7–8 by 7–8 mm. Seeds ± globose, 8.2–10 by 8.2–10 by 8.2–10 mm, surface smooth, dull, brown.

    Distribution — Borneo, Philippines.

    Habitat & Ecology — Primary and secondary forest, along river sides; on alluvial to clay-loam soil. Altitude 30–200 m. Flowering and fruiting: March to November.

Vernacular names — Borneo: Kemantah rimba.

    Note — Mallotus connatus can easily be confused with M. rufidulus. The main differences are found in leaf and floral characters: Leaves drying green to green-brown (brown in M. rufidulus); stipules linear-triangular (narrowly triangular in M. rufidulus); pistillate inflorescences with entire or trilobed bracts (trilobed in M. rufidulus), pistillate flower buds ellipsoid (ovoid in M. rufidulus), connate filaments (free to slightly connate at the base in M. rufidulus), styles 2–4 mm long (0–1(–2)) mm in M. rufidulus), stigmas 0.8–1 mm thick (1–1.3 mm in M. rufidulus). Besides differences in morphology they also differ in geographical distribution; M. connatus is found in Borneo and the Philippines, while M. rufidulus is found in Sumatra, Java, Borneo and Lesser Sunda Islands.

13. Mallotus cumingii Müll.Arg. (sect. Rottleropsis)

    Mallotus cumingii Müll.Arg., Linnaea 34 (1865) 195; in DC., Prodr. 15, 2 (1866) 978; S.Vidal, Rev. Pl. Vasc. Filip. (1886) 245; Kulju, S.E.C.Sierra & Welzen, Blumea 52 (2007) 121, fig. 1, map 2. — Neotrewia cumingii (Müll.Arg.) Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 212; Merr., Enum. Philipp. Fl. Pl. 2 (1923) 437; Airy Shaw, Kew Bull. 37 (1982) 31; Alphab. Enum. Euph. Philipp. Isl. (1983) 39; De Guzman et al., Guide Philipp. Fl. Fauna 3 (1986) 292. — Type: Cuming 798 (holo G-DC, microfiche IDC #800; iso BM, CGE, K, L, W), Philippines, Luzon, Tayabas Province.

    Trewia ambigua Merr., Philipp. J. Sci. 1, Suppl. 1 (1906) 79. — Syntypes: FB (Borden) 623 (BM, K, US), Philippines, Luzon, Bataan Prov., Lamao River; FB (Borden) 1251 (BM, K, US), Philippines, Luzon, Bataan Prov., Lamao River, Mt Mariveles; FB (R. Meyer) 2798 (K, NY, US), Philippines, Luzon, Bataan Prov., Lamao River, Mt Mariveles.

Shrubs to small trees up to 18(–25) m high, dbh up to 30(–45) cm, dioecious, evergreen; young branches usually flattened when dry, sparsely to densely hairy, older ones terete, glabrescent; flowering branches 1–5 mm thick. Outer bark smooth to rugose, often peeling in thin plates, grey to (yellowish or reddish) brown, sometimes patchy. Indumentum with simple to (stellately) tufted hairs, whitish to yellowish (to brownish); glandular hairs yellowish to reddish, in most parts absent (to sparsely present). Stipules (narrowly) triangular to obovate, 1.8–6 by 0.7–2 mm, early caducous, margin entire, apex obtuse to acute, densely hairy outside, sparsely (to densely) hairy inside. Leaves opposite; petiole 0.4–11.8 cm long, with or without a basal and apical pulvinus, glabrous to densely hairy; blade ovate to obovate, 4.8–35.5 by 1.5–19.5 cm, length/width ratio 1.3–3.5, papery, base (truncate to) obtuse to cuneate to attenuate, margin subentire to wavy (to subdentate), glandular teeth 2–12(–16) per side, often puberulus or papillose and located in a minute notch, apex (rounded to) acuminate to cuspidate, usually ending in a gland; upper and lower surface glabrous, to sparsely hairy on the venation; upper surface dark green, somewhat shiny, smooth to sparsely granular dotted, basal extrafloral nectaries 2–4(–8), 0.5–18 mm from petiole insertion, marginal extrafloral nectaries 0–5(–12) per side, 1.6–15 mm from margin; lower surface paler green, densely dotted with minute bumps, glandular hairs sometimes present in young leaves, caducous to sparsely present in adult leaves, hair-tuft domatia sometimes present; venation pinnate, nerves 4–11 per side, looped and closed near the margin. Inflorescences axillary racemes, single, or rarely 2 together when staminate, axis sparsely to densely hairy; bracts mostly caducous, ovate to triangular, somewhat cymbiform, margin entire, apex (obtuse to) acute to acuminate, both sides hairy to subglabrous inside when pistillate. Flowers: pedicels sparsely to densely hairy, with basal abscission zone; sepals sparsely hairy outside, (sub)glabrous inside. Staminate inflorescences up to 21 cm long, basally 0.6–1.5 mm thick; bracts 1.2–2.5(–3.7) by 0.6–1.3 mm. Staminate flowers: buds globose, apiculate or not; flowers 3.2–5.2(–6.9) mm diam., white to cream to yellowish, with sweet odour; pedicels 1.8–5.2(–6.8) mm long; sepals (2 or) 3 or 4, free (to basally connate), ovate to elliptic, 1.8–3.2 by 0.8–1.9 mm, often recurved when dry, apex obtuse to acuminate, sometimes with glandular hairs outside; disc-glands often present; stamens 30–60, filaments 0.9–3.1(–4.2) mm long, free, glabrous, thecae 0.2–0.5 by 0.15–0.25 mm, opposite to subalternate, opening latrorsely to irregulary, connective up to twice as wide as the filament, papillose, apex irregulary apiculate or widened. Pistillate inflorescences up to 12.5(–19) cm long, basally 0.7–2.5 mm thick; bracts 1.1–2.6 by 0.5–0.9 mm. Pistillate flowers: buds ovoid, 0.7–1.1 mm diam. just before anthesis; flowers greenish, with a faint, sweet odour; pedicels 0.9–7.4 mm long; calyx 2.9–4.5 mm long, opening via 1 or 2 sutures, soon caducous; ovary ellipsoid, densely hairy, glandular hairs usually present, locules 1 (or 2); style 0.5–3(–4) mm long, sparsely to densely hairy, smooth to verrucose; stigma 4.4–8.4 mm long, reflexed, stigmatic side plumose, non-stigmatic side hairy. Fruits indehiscent, dry drupes, (oblate to) spheroid to broadly ellipsoid, often slightly bilobed when 2-seeded, 9–15 by 9–15 mm, often with style remnants, surface smooth to slightly verrucose, hairy, with or without some glandular hairs, speckled, (yellowish) green to (reddish) brown when fresh, brown to dark grey when dry; pericarp 0.4–2.9 mm thick, dry, hard to somewhat brittle. Seeds oblate to spheroid to broadly ellipsoid, 6–10 by 7–12 mm, surface smooth, white to brown, sarcotesta absent.

    Distribution — Borneo, Philippines, Sulawesi.

    Habitat & Ecology — In primary and secondary forest, frequently in damp places along rivers and streams. Altitude up to 1000(–1660) m. Flowering and fruiting (in Philippines): throughout the year, flowering especially from March to June.

Vernacular names — Philippines: Banilad, bunog (Tagbanua); batobato (Tagalog); wamut (Subanun); apanang, balogo, cabantala, potian, suysuysagpub, tabunghain.

    Notes — 1. This species shows a large variation in leaf size, which is partly caused by the size inequality of each leaf pair. The variation is, however, continuous and does not correlate with other characters. Specimens from Batan Island, Philippines (BS 80605, BS 80686, BS 80720) differ from others by having only ovate leaves with low

length/width ratios and somewhat longer staminate bracts.

2. The distribution of this species, previously only know from the Philippines and Sulawesi, is here extended to Borneo. Only one Bornean specimen, S series 28705 (Kapit Distr., Sarawak), is known so far.

3. Five specimens from the Lesser Sunda Islands (Kostermans 18263, Kostermans 18736, Mulyati et al. 503, Schmutz 4133, Schmutz 4797) resemble M. cumingii closely, but differ from it by the consistent presence of glandular hairs on the lower leaf surface, and short and stout staminate inflorescences covered with hairs longer than in M. cumingii. As pistillate specimens are not known, the status of these collections remains unclear.

14. Mallotus darbyshirei Airy Shaw (sect. Rottleropsis)

    Mallotus darbyshirei Airy Shaw, Kew Bull. 34 (1980) 597; S.E.C.Sierra et al., Blumea 52 (2007) 58, map 7. — Type: Darbyshire & Hoogland 7956 (holo BRI; iso A, L), Papua New Guinea, Aitape, Aiserap.

Shrubs, up to 4 m high, dioecious; branches hollow. Indumentum composed of simple and stellately-tufted hairs and dark red glandular hairs. Stipules triangular, c. 7 by 3 mm, early caducous, margin entire, apex acute, densely hairy outside, glabrous inside. Leaves opposite; petiole 18–54 by 1.8–3 mm, densely to sparsely hairy; blade elliptic to obovate, 17–34.5 by 7–18 cm, length/width ratio 1.9–2.4, chartaceous, base obtuse to cuneate, margin entire, undulate, apex caudate, upper surface sparsely to scatteredly hairy on midrib and nerves, extrafloral nectaries along the midrib, 20–30 per side, also all over the blade, elliptic to orbicular, 0.6–0.8 by 0.5–0.6 mm, lower surface sparsely hairy, scatteredly covered with glandular hairs (sometimes only present near the leaf margin), domatia absent, venation inconspicuously triplinerved, nerves 5 or 6 per side, looping and closed near margin. Staminate inflorescences unknown. Pistillate inflorescences umbels, 1 or 2 together, erect, 3.8–4.5 cm long, axes densely hairy; peduncle 25–27 by 1.2–1.5 mm; nodes per branch 1 or 2, with 3 or 4 flowers per bract; bracts triangular, c. 2 by 1 mm, margin entire, apex acute; bracteoles absent. Pistillate flowers 5–6 mm diam.; pedicels 6–10 mm long, densely hairy; sepals 3, triangular to narrowly triangular, 4–4.5 by c. 1.5 mm, free, early caducous, margin entire, apex acute, densely hairy outside, glabrous inside; ovary 3–4 by 3–4 mm, 3-locular, densely hairy, without glandular hairs; stigmas sessile, 3–4 by 0.8–1 mm. Fruits capsules, c. 9 by 14 mm, green, opening septicidally-loculicidally, surface spiny, densely hairy, without glandular hairs; spines 500–600, ± straight, 2.5–3 by 0.2–0.3 mm, densely hairy; wall 0.8–1 mm thick, densely hairy inside; column c. 6 by 5 mm. Seeds ± globose, c. 6.5 by 4.5 by 4.5 mm, surface smooth, dull, brown.

    Distribution — Endemic to New Guinea.

    Habitat & Ecology — On gentle slope. Altitude c. 15 m. Fruiting: June.

    Vernacular name — Wommollo (Wapi).

15. Mallotus decipiens Müll.Arg. (sect. Rottleropsis)

    Mallotus decipiens Müll.Arg., Linnaea 34 (1865) 194; Hook.f., Fl. Br. Ind. 5 (1887) 434; Airy Shaw, Kew Bull. 26 (1972) 303; S.E.C.Sierra et al., Blumea 52 (2007) 59, map 8, plate 1d. — Lectotype (S.E.C.Sierra et al., 2007): Wallich 8009 (holo G-DC, IDC microfiche 800, the sheet with the field number 2130 and staminate infl.; iso K-W, IDC microfiche 7394), Myanmar, Mon Prov., banks of Ataran River (‘2130, Myanmar, ripae Attran’), see note 1.

    Coelodiscus eriocarpoides Kurz [J. Asiat. Soc. Bengal 42 (1873) 244, nom. nud.] Forest Fl. Burma 2 (1877) 392. — Type: Parish s.n. (CAL n.v., fide Hook.f., 1887: 434), Myanmar, Upper Taninthayi Prov. (‘Myanmar, Upper Tenasserim’), see note 2.

    Mallotus cuneatus Ridl., J. Straits Branch Roy. Asiat. Soc. 59 (1911) 181. — Mallotus resinosus (Blanco) Merr. var. cuneatus (Ridl.) N.P.Balakr. & Chakrab., Rheedea 1 (1991) 39. — Type: Ridley 15190 (holo SING; iso K), Peninsular Malaysia, Perlis, Bukit Lagi, see note 3.

    Coelodiscus muricatus auct. non. (Wight) Gagnep.: Gagnep. in Lecomte, Fl. Indo-Chine 5 (1925) 369, excl. basion.: Eberhardt 3258 (K), Vietnam, Lang Son; Bon 2574 (P), Vietnam, Hanoi; Pételot 1277 (P, UC), Vietnam, Tonkin, Cho Ganh; Pételot 1417 (UC), Vietnam, Tonkin, Cho Ganh; Pételot 1418 (UC), Vietnam, Tonkin, Cho Ganh; Pierre s.n. (P), Cambodia, Mt Schrall; all others not seen.

Shrubs, 0.5–4 m high, dioecious. Outer bark slightly rough. Indumentum composed of simple and tufted hairs and yellow to orange glandular hairs. Stipules triangular, 2–4 by 1–1.8 mm, caducous, margin entire, apex acute, scatteredly hairy outside, glabrous inside. Leaves opposite; petiole 2–24 by 0.9–1.4 mm, sparsely hairy; blade obovate, 2.8–20 by 1.6–12 cm, length/width ratio 1.9–2.4, membranaceous to chartaceous, base obtuse to cuneate, margin denticulate, dentate (specially towards the apex) to crenate, with glandular teeth, apex cuspidate to caudate, upper surface scatteredly hairy to glabrous on midrib and nerves, extrafloral nectaries marginal in lower half, 1 or 2 per side, 1.5–3 mm from margin, elliptic to orbicular, 0.8–2.2 by 0.6–0.8 mm, lower surface sparsely hairy, densely (to sparsely) covered with glandular hairs, domatia sometimes present, venation triplinerved, nerves 6–8 per side, looping and closed near margin or ending in margin. Inflorescences racemes, single, erect; peduncle 8–32 by 0.2–1 mm; axes densely to sparsely hairy; bracts triangular, persistent, margin entire to subentire, apex acute to acuminate, densely to sparsely hairy outside, glabrous inside; bracteoles sometimes present, triangular, 0.2–0.3 by 0.2–0.3 mm, caducous, margin entire. Flowers: pedicels densely to sparsely hairy; sepals persistent, margin entire, apex acute, sparsely hairy and with glandular hairs outside, glabrous inside. Staminate inflorescences 7–16 cm long, nodes per branch 22–41, with 2–7 flowers per bract; bracts 1–2 by 0.9–1 mm. Staminate flowers 2.5–4 mm diam.; pedicels 1.8–2.2 mm long; sepals 3, elliptic to obovate, 1.5–2.2 by 0.7–1.5 mm, free; stamens 22–32, filaments 1.3–3 mm long, free, glabrous, thecae ovoid, 0.2–0.3 by 0.2–0.25 mm, usually glabrous, connective widened; pistillode present. Pistillate inflorescences 6.8–16 cm long, nodes per branch 13–36, with 1 flower per bract. Pistillate flowers 0.8–1 mm diam.; pedicels 0.3–1 mm long; calyx 3-lobed, lobes triangular, 1.5–1.7 by 0.8–1 mm; ovary 0.5–0.6 by 0.5–0.6 mm, 3-locular, sparsely hairy, scatteredly covered with glandular hairs; style 0.2–0.5 mm long; stigmas 0.7–1 by c. 1 mm. Fruits capsules, 3–4 by 4–6 mm, pale light green, opening septicidally-loculicidally, surface spiny, sparsely hairy, sparsely covered with glandular hairs; spines 150–180, curved, 0.5–0.7 by 0.2–0.25 mm, scatteredly hairy; wall c. 0.2 mm thick, glabrous inside; column 2.9–3.1 by 2.9–3.1 mm. Seeds ± globose, 3.6–3.8 by 3.1–3.2 by c. 3.2 mm, surface smooth, shiny, brown-grey.

    Distribution — Bangladesh, Myanmar, Laos, Vietnam, Thailand, Peninsular Malaysia (northern).

    Habitat & Ecology — Locally common in very degraded, seasonal, evergreen hardwood forests with bamboo, deciduous dipterocarp-oak and pine forests, dry secondary scrub; in shaded areas, along roadsides, or on steep slopes; on granite, limestone, calcareous rock, sandstone, or shale bedrock soil. Altitude: sea level up to 800 m. Flowering and fruiting throughout the year.

    Notes — 1. Müller Argoviensis (1865) described Mallotus decipiens on “Wall.! In hb. DC.” However, there are two gatherings in Wallich’s Numerical List: 7725 and 8009, curiously both labelled “Attran, 17 May 1827”, to which 7725 adds ‘ripae’, i.e. riverbanks, and 8009 the field number 2130. In G-DC there is a sheet labelled “2073, Attran R. 17 May 1827” with a pistillate plant. The 7725 sheet in K-W has both staminate and pistillate inflorescences, which indicates a mixed collection, possibly of field numbers 2073 and 2130.

2. The type, place of deposit, and identity of Coelodiscus eriocarpoides Kurz are accepted here on the base of the remarks made by Hooker f. (1887: 434). The brief diagnosis corresponds to our concept of M. decipiens.

3. Balakrishnan & Chakrabarty (1991) recognized M. cuneatus at the varietal level, mentioning that it differs from M. resinosus by the presence of triplinerved leaf venation and hairy capsules (pinnate venation and densely glandular capsules in M. resinosus). However, there are more characters that separate them: Extrafloral nectaries with at least one pair always present on the nerves (below the nerves in M. resinosus), 6–8 nerves (8–12 in M. resinosus), smaller fruits, 3–4 by 4–6 mm (larger fruits, 6–8 by 8–15 in M. resinosus), presence of short spines, 0.5–0.7 by 0.2–0.3 (longer, 1–2.2 by 0.3–0.5 in M. resinosus).

16. Mallotus didymochryseus Airy Shaw (sect. Rottleropsis)

    Mallotus didymochryseus Airy Shaw, Kew Bull. 20 (1966) 40; S.E.C.Sierra et al., Blumea 52 (2007) 61, map 7. — Type: NGF (C.T. White) 8255 (holo K; iso CANB, L), Papua New Guinea, Lawes.

Small to large trees, 8–28 m high, dioecious. Outer bark shallowly longitudinal fissured. Indumentum composed of simple, stellately-tufted and stellate hairs (on petioles), glandular hairs absent. Stipules triangular to narrowly triangular, 1–1.8 by 0.4–0.6 mm, early caducous, margin entire, apex acute, densely hairy. Leaves opposite; petiole 25–80 by 2–3 mm, densely to sparsely hairy; blade ovate, 12–23.5 by 9.5–22 cm, length/width ratio 1.1–1.4, chartaceous, base cordate to rounded, margin entire to shallowly repand, with indistinct glandular teeth, apex rounded, acute to acuminate, upper surface densely hairy on midrib and nerves, extrafloral nectaries marginal from base to apex, 10–35 per side, 1–6 mm from margin, elliptic to orbicular, 0.2–0.7 by 0.2–0.5 mm, lower surface sparsely hairy, domatia absent, venation triplinerved, nerves 6 or 7 per side, indistinctly looping or ending in margin. Inflorescences racemes, single, erect; peduncle 6–45 by 1.5–2 mm; axes densely hairy; bracts triangular to narrowly triangular, margin entire, apex acute, densely hairy, early caducous; bracteoles absent. Flowers: pedicels densely hairy; sepals free, persistent, margin entire, apex acute, densely hairy outside, glabrous inside. Staminate inflorescences 20–22 cm long, nodes per branch 35–47, with 5–7 flowers per bract; bracts 1–2.2 by 0.4–0.6 mm. Staminate flowers 5–7 mm diam.; pedicels 3.5–4 mm long; sepals 3 or 4, ovate to elliptic, 2.5–3 by 1.5–1.8 mm; stamens 60–90, filaments 1–3 mm long, free, glabrous, white, thecae ellipsoid, 0.2–0.3 by 0.2–0.25 mm, hairy, pale light cream, connective widened; pistillode present. Pistillate inflorescences 5–12 cm long, nodes per branch 8–30, with 1 flower per bract; bracts 1–3 by 0.5–0.9 mm. Pistillate flowers 3–5 mm diam.; pedicels 5–8 mm long; sepals 4–6, triangular, 2.5–3 by 0.8–1 mm; ovary 1.6–1.8 by 1.5–1.7 mm, 2-locular, densely hairy; style 0.2–0.4 mm long; stigmas 2.3–2.7 by c. 2 mm. Fruits tardily dehiscent capsules, opening loculicidally, 10–12.5 by 12–17 mm, orange yellow, surface wrinkled, densely hairy; wall c. 0.5 mm thick, glabrous inside; column 8–9 by 3–3.2 mm. Seeds ± globose, 6.5–7.5 by 7–8 by 6–7 mm, surface slightly rugose, dull, whitish cream.

    Distribution — Endemic to New Guinea.

    Habitat & Ecology — Scattered in the understorey of seasonally dry evergreen rain forest or in the upper tier of mixed primary or secondary alluvial lowland or swamp forest (periodically flooded to 1 m deep); on forest edges. Altitude: sea level up to 580 m. Flowering and fruiting: November to July.

    Vernacular names — New Guinea: Indonesia: Kamuretjia (Tarie); Papua New Guinea: Tivamunana (Maya); Okaibu.

    Notes — 1. All the fruits studied were closed (even the ripe ones) and lacked the characteristic sutures present of dehiscent Mallotus species.

2. Mallotus didymochryseus can be distinguished from M. trinervius by the sand-papery texture of the upper surface of the leaves (soft in M. trinervius), 10–35 marginal extrafloral nectaries (5 or 6 in M. trinervius), cordate to rounded leaf base (rounded to cuneate in M. trinervius) and the tardily dehiscent, small, 10–12.5 by 12–17 mm, orange yellow fruits (dehiscent, large, 17–22.5 by 24–29 mm, brown ochre in M. trinervius).

17. Mallotus dispar (Blume) Müll.Arg. (sect. Rottleropsis)

    Mallotus dispar (Blume) Müll.Arg., Linnaea 34 (1865) 191 (excl. var. psiloneurus Müll.Arg. = M. resinosus (Blanco) Merr.); in DC., Prodr. 15, 2 (1866) 971; Airy Shaw, Kew Bull. 21 (1968) 380; Kew Bull. 26 (1972) 303; Whitmore, Tree Fl. Malaya 2 (1973) 115; Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 169; Welzen et al., Thai Forest Bull., Bot. 28 (2000) 99; Slik, Priyono & Welzen, Gard. Bull. Singapore 52 (2000) 56, Fig. 25; in Slik, Tropenbos-Kalimantan ser. 4 (2001) 94, Fig. 5.28; S.E.C.Sierra et al., Blumea 52 (2007) 62, map 9. — Rottlera dispar Blume, Bijdr. (1826) 608. — Mallotus dispar (Blume) Müll.Arg. var. lasioneurus Müll.Arg., Linnaea 34 (1865) 191, nom. inval. — Type: Blume s.n. (L), Java, Kuripan.

    Claoxylon ? stipulosum Rchb.f. ex Zoll., Acta Soc. Regiae Sci. Indo-Néerl. 1 (1856) 20; Miq., Fl. Ned. Ind. 1, 2 (1859) 387. — Type: Zollinger 3681 (?), Java, Bogor Botanical Garden, originally from Nusa Kambangan.

Also check: Slik, Macaranga and Mallotus species of Borneo

Shrubs to small trees, 2–10 m high, dioecious or rarely monoecious. Indumentum composed of simple and tufted hairs and yellow to orange glandular hairs. Stipules narrowly triangular, 5–6 by 1.2–1.5 mm, persistent to caducous, margin entire, apex acute, densely hairy outside, sparsely hairy inside. Leaves opposite; petiole 3–50 by 0.8–1.3 mm, densely hairy; blade ovate to obovate, 5–20 by 4–9 cm, length/width ratio 2.1–2.5, membranaceous to chartaceous, base rounded, obtuse to cuneate, margin denticulate, dentate to crenate, with glandular teeth, apex acuminate to caudate, upper surface glabrous, extrafloral nectaries marginal in lower half, rarely along the midrib, 1 or 2 (or 3) per side, 1.5–7(–12) mm from margin, elliptic to orbicular, 0.5–2 by 0.5–1 mm, lower surface sparsely hairy, sparsely covered with glandular hairs, domatia sometimes present, venation triplinerved, nerves 7–9 per side, looping and closed near margin. Inflorescences racemes, single, erect, unisexual to rarely bisexual; peduncle 15–30 by 0.5–0.8 mm long; axes densely hairy; bracts narrowly to linear-triangular, margin entire, apex acute, densely hairy outside, scatteredly hairy to glabrous inside; bracteoles absent. Flowers: pedicels densely to sparsely hairy; sepals free, persistent, margin entire, apex acute, densely hairy outside, scatteredly hairy to glabrous inside, with glandular hairs outside. Staminate inflorescences 6–12 cm long, nodes per branch 12–18, with 1–4 flowers per bract; bracts 3–5.5 by 0.7–1.3 mm. Staminate flowers 3.5–4 mm diam.; pedicels 2–4 mm long; sepals 3 or 4, elliptic to obovate, 1.8–2.3 by 0.5–1.3 mm, apex acute; stamens 28–50, glabrous, filaments 1.5–2.5 mm long, free, thecae ovoid, 0.2–0.3 by 0.2–0.25 mm, connective widened; pistillode present. Pistillate inflorescences 5–12 cm long, nodes per branch 6–9, with 1 flower per bract; bracts 3.5–6 by 0.8–1.7 mm; bracteoles absent. Pistillate flowers 2.5–4 mm diam.; pedicels 1.5–2 mm long; sepals 3–5, narrowly triangular, 3–4 by 0.8–1.2 mm; stami-nodes rarely present; ovary 0.8–1.5 by 1.8–2 mm, 3-locular, densely hairy, sparsely covered with glandular hairs; style 0.7–1.3 mm long; stigmas 3.5–6 by 1.5–2 mm. Fruits capsules, 8–12 by 9.5–13 mm, light green, opening septicidally-loculicidally, surface spiny, sparsely hairy, sparsely covered with glandular hairs; spines 60–80, curved, 2–4 by 0.1–0.2 mm, sparsely hairy; wall 0.7–1 mm thick, glabrous inside; column 3–5 by 3–5 mm. Seeds ± globose, 4–5 by 4–5 by 4–5 mm, surface smooth, shiny, brown, aril or arillode absent.

    Distribution — From Thailand to Peninsular Malaysia, Sumatra, Java.

    Habitat & Ecology — Locally in evergreen or coastal forest mixed with lowland dipterocarp forest. On clay, limestone or brown soil, on cliffs. Altitude: sea level up to 600 m. Flowering and fruiting throughout the year.

    Note — Mallotus dispar differs from M. minimifructus in the presence of 1 or 2 (or

3) pairs of extrafloral nectaries, with al least one pair always present on the nerves,

(2–7 pairs below the nerves in M. minimifructus).

18. Mallotus eximius Airy Shaw (sect. Rottleropsis)

    Mallotus eximius Airy Shaw, Kew Bull. 32 (1978) 400; S.E.C.Sierra et al., Blumea 52 (2007) 64, map 10. — Type: KEP FRI (Loh) 13517 (holo K; iso A, L, SING), Peninsular Malaysia, Ulu Trengganu.

Small trees, 2–6 m high, dioecious. Indumentum composed of simple, tufted and stellately-tufted hairs and light yellow to orange glandular hairs. Stipules narrowly triangular, 5–12 by 1.5–5 mm, early caducous, margin entire, hairy, apex acute, densely hairy outside, sparsely hairy inside. Leaves opposite; petiole 60–120 by 2–3 mm, densely hairy; blade ovate to elliptic, 12–48 by 7–25 cm, length/width ratio 1.7–2.6, chartaceous, base sometimes peltate up to 5 mm, cordate, rounded to obtuse, margin entire, apex caudate, upper surface densely hairy on midrib and nerves, extrafloral nectaries marginal throughout blade, up to 30 per side, 2–30 mm from margin, sometimes also all over the blade, elliptic to orbicular, 0.2–0.8 by 0.2–0.5 mm, lower surface sparsely hairy, sparsely covered with glandular hairs, domatia absent, venation triplinerved, nerves 7–9 per side, looping and closed near margin. Inflorescences racemes, single, erect; axes densely hairy; bracts persistent, densely hairy, margin entire or trilobed, apex acute; bracteoles sometimes present. Flowers: pedicels densely hairy; sepals connate in lower half, margin entire, apex acute, surfaces densely hairy, without glandular hairs. Staminate inflorescences 10–15 cm long; peduncle 10–20 mm long; nodes per branch 18–20, with 4–6 flowers per bract; bracts triangular, 2.8–3.2 by 0.6–0.8 mm; bracteoles entire or trilobed, 0.8–2 by 0.3–0.4 mm. Staminate flowers 4.8–5.2 mm diam.; pedicels 1–3 mm long; calyx 3-lobed, lobes triangular to narrowly triangular, 3–4.1 by 1.8–2 mm; stamens 30–35, filaments 3–5 mm long, connate in lower half, hairy, thecae ovoid, 0.2–0.3 by 0.2–0.25 mm, subglabrous, connective not widened; pistillode present. Pistillate inflorescences 15–34 cm long, nodes per branch 6–12, with 1 flower per bract; bracts trilobed, 6–9 by 4–6 mm; bracteoles absent. Pistillate flowers 6–8 mm diam.; pedicels 6–11 mm long; calyx 5- or 6-lobed, lobes triangular, 4.5–6 by 3–4 mm; ovary 3.2–4 by 4–5 mm, 3-locular, densely hairy, glandular hairs absent; style 2–3 mm long; stigmas 4–7 by 1.2–1.4 mm long. Fruits capsules, 17–20 by 18–20 mm, green to brown green, opening septicidally-loculicidally, surface spiny, densely hairy, without glandular hairs; spines c. 700, ± straight, 1.5–3 by 0.4–0.6 mm, densely hairy, hairs forming a continuous layer; wall 1.7–2 mm thick, glabrous inside; column 12–13 by 10–11 mm. Seeds ± globose, 10–12 by 9–11 by 9–11 mm, surface smooth, dull, brown.

    Distribution — Peninsular Malaysia, NW Borneo.

    Habitat & Ecology — Locally common in understorey of primary to secondary forests, scrubs, mostly found in disturbed sites, also on hillsides; on granite to limestone soil. Altitude 150–400 m. Flowering and fruiting: May to September.

    Vernacular name — Borneo: Brati manuk (Dayak).

19. Mallotus floribundus (Blume) Müll.Arg. (sect. Stylanthus)

    Mallotus floribundus (Blume) Müll.Arg., Linnaea 34 (1865) 187; in DC., Prod. 15, 2 (1866) 962; Rolfe, J. Bot. 23 (1885) 215; Hook.f., Fl. Brit. India 5 (1887) 432; Boerl., Handl. Fl. Ned. Ind. 3(1) (1900) 288; J.J.Sm., Meded. Dept. Landb. Ned.-Indië 10 (1910) 417; Elmer, Leafl. Philipp. Bot. 4 (1911) 1297; Koord., Exkurs.-fl. Java (1912) 492; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 173; in Engl., Pflanzenr. IV.147. viii (1919) 17; Merr., J. Str. Roy. Asiatic Soc. (1921) 338; Enum. Philipp. Flow. Pl. 2 (1923) 433; Ridl., Fl. Malay Penins. 3 (1924) 290; Pax & K.Hoffm. in Engl. & Harms, Natürl. Pflanzenfam. ed. 2, 19c (1931) 115; Burkill, Dict. Econ. Prod. Malay Penins. 2 (1935) 1395; Corner, Wayside Trees Mal. (1940) 271; Backer & Bakh.f., Fl. Java 1 (1964) 483; Meijer, Bot. News Bull. (1967) 51; Airy Shaw, Kew Bull. 21 (1968) 385; Kew Bull. 26 (1972) 305; Whitmore, Tree Fl. Malaya 2 (1973) 113; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 172; Kew Bull. 31 (1976) 392; Kew Bull. Add. Ser. 8 (1980) 166; Kew Bull. 36 (1981) 323, 326; Alph. Enum. Euphorb. Philipp. Isl. (1983) 36; Corner, Wayside Trees Mal., ed. 2 (1988) 306; Nguyen Nghia Thin, Euph. Vietnam (1995) 11; Slik, Priyono & Welzen, Gard. Bull. Singapore 52 (2000) 57, Fig. 27; Slik & Welzen, Blumea 46 (2001) 33, Fig. 13, Map 5; Slik, Priyono & Welzen in Slik, Tropenbos-Kalimantan ser. 4 (2001) 95, Fig. 5.30; Slik & Welzen in Slik, Tropenbos-Kalimantan ser. 4 (2001) 177, Fig. 7.13, Map 7.5. — Adisca floribunda Blume, Bijdr. Fl. Ned. Ind. 11 (1825) 610. — Rottlera floribunda (Blume) Hassk., Cat. Hort. Bot. Bogor. (1844) 238; Rchb.f. & Zoll., Linnaea 28 (1856): 313; Baill., Etude Euphorb. (1858) 426; Miq., Fl. Ned. Ind. 1 (1859) 393. — Mappa floribunda (Blume) Zoll. & Mor., Syst. Verz. (1855) 17. — Mallotus floribundus (Blume) Müll.Arg. var. genuinus Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 174, nom. inval. — Lectotype (Slik & Welzen, Blumea 46, 2001): Blume s.n. (L sheet. 904 105 – 104) (L holo), Java.

    Ricinus tanarius Lour., Fl. Cochinch. 1 (1790) 584, non L.; Fl. Cochinch. 2 (1793) 717, nom. inval. — Type: Igyn G. 123 (BM holo), Bach dan nam.

    Mallotus amentiformis Müll.Arg., Flora 47 (1864) 468; in DC., Prod. 15, 2 (1866) 962. — Type: Griffith KD 4762 (K iso), Malaya Peninsula.

    Mallotus annamiticus Kuntze, Revis. Gen. Pl. 2 (1891) 608; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 204; Merr., Trans. Amer. Philos. Soc., New Ser. 24 (1935) 236. — Coelodiscus annamiticus (Kuntze) Gagnep. in Lecomte, Fl. Gen. Indo-Chine 5 (1925) 375. — Type: Kuntze 3823 (NY holo; NY iso), Annam, Turong.

    Mallotus floribundus (Blume) Müll.Arg. var. pilosus Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 174; Airy Shaw, Kew Bull. Add. Ser. 8 (1980) 166. — Lectotype (Slik & Welzen, Blumea 46, 2001): Lauterbach 1232 (WRSL holo), Papua New Guinea.

    Also check: Slik, Macaranga and Mallotus species of Borneo

Shrubs to small trees up to 18(–25) m tall, dbh up to 30 cm; bole mostly straight; crown bushy, much branching. Outer bark smooth to fissured, lenticellate; sapwood soft. Indumentum glabrous or simple to tufted, whitish to creamish. Branches lenticellate, glabrescent, gland-dotted on young parts. Stipules early caducous, triangular to ovate, 1.6–3.8 by 0.6–1.3 mm, margin irregular to entire, apex acute (to acuminate), hairy abaxially only, gland-dotted abaxially or not. Leaves alternate to apically subopposite, peltate, petiole insertion 3–27 mm from base of leaf blade; petiole 11–152 by 0.4–2.5 mm, glaucous, basally usually constricted when dried, apically sometimes slightly constricted, glabrous to sparsely hairy, glabrescent, gland-dotted or not; blade (ovate) to broadly ovate to orbicular, 2–20 by 2–17.5 cm, length-width ratio 0.7–1.9, base rounded to truncate to slightly emarginate, margin irregularly wavy (to slightly dentate to crenate), marginal glands 1–12 per leaf side, apex acute to acuminate (to slightly cuspidate), upper surface smooth, glabrous (to sparsely hairy at petiole insertion), sparsely gland-dotted or not, especially at petiole attachment, basal macular glands 0–8 on veins, 3–55 mm from petiole insertion, apical macular glands 0–19 on veins, 1–6(–17) mm from margin, lower surface smooth, usually glaucous, glabrous to hairy, gland-dotted, domatia present, basally often conspicuously large; venation palmate, 7–11 veins originating from petiole, 3–6 secondary veins per side along midrib, ending in the margin. Staminate inflorescences up to 23 cm long, basally up to c. 1 mm thick, solitary to sometimes grouped together, glabrous to hairy, gland-dotted, nodes up to 46 per inflorescence, flowers 1–9 per node; bracts early caducous, triangular to ovate to broadly ovate, cup-shaped around pedicel base, 0.6–2.6 by 0.5–2.8 mm, margin entire to irregular, apex acute (to acuminate to rounded), hairy abaxially, glabrous (to sparsely hairy) inside, gland-dotted or not. Buds globose, glabrous to hairy, gland-dotted or not. Staminate flowers 2–6 mm in diameter, white yellowish, slightly fragrant; pedicels 1.3–5 mm long, glabrous to hairy, gland-dotted or not; sepals 2–5, free to basally connate, elliptic to ovate (narrowly triangular to triangular), often recurved, 0.5–5 by 0.5–2.3 mm, margin entire, apex (rounded to) acute (to acuminate), glabrous to hairy, gland-dotted or not; stamens up to 56, filaments up to 4 mm long, glabrous to rarely hairy, rarely gland-dotted; anthers 0.3–0.5 by 0.3–0.6 mm, basifixed; thecae glabrous to hairy along aperture; connective not (to slightly) widened. Pistillate inflorescences up to 22 cm long, basally 1–2 mm thick, usually solitary, hairy, glabrescent, gland-dotted, nodes up to 14(–19) per inflorescence; bracts caducous, (triangular to) ovate to broadly ovate, 0.9–3.5 by 0.6–2 mm, margin irregular to entire, apex (rounded to) acute (to acuminate), hairy, gland-dotted or not. Pistillate flowers pale green to whitish; bracteoles absent to caducous, (broadly) ovate to elliptic, rarely linear to triangular, 0.5–1.4 by 0.3–0.8 mm, margin entire to irregular, apex (rounded to) acute (to acuminate), glabrous to hairy abaxially only, gland-dotted or not; pedicels 0.7–7 mm long, glabrous to hairy, gland-dotted or not; calyx caducous, 2–5-lobed, free to more or less connate, narrowly triangular to triangular, 1.1–6 by 0.2–1.8 mm, margin entire to irregular, apex acute (to acuminate), hairy abaxially, glabrous to hairy adaxially, gland-dotted or not; ovary (2–)3(–4)-locular, echinate, densely woolly hairy, densely gland-dotted, spines up to 7 mm long, hairy, gland-dotted or not, especially basally; style up to 6 mm long, glabrous to hairy, gland-dotted; stigmas 4–11 mm long, densely covered with granulate papillae adaxially, glabrous to hairy abaxially, gland-dotted abaxially. Fruit an echinate, lobed capsule, 10–20 by 7–12 mm, green, glabrous to hairy, densely gland-dotted; carpel inside glabrous to hairy, gland-dotted or not; column 4–7 by 4–10 mm. Seeds glossy, globose, 4.7–8 by 5 by 3–8 mm; hilum deltoid to cordate, 0.5–2.2 by 0.8–3 mm.

    Distribution — Southeast Asia to New Guinea and the Solomon Islands.

    Habitat & Ecology — Locally common in primary and secondary forests, mainly in open places; often found on river banks, along roads, in gaps, clearings, and open fields; in badly drained swampy areas as well as in dry areas; on most soil types, from sandy to loamy to clayey to limestone. Altitude: up to 500(–933) m.

    Uses — The rather aromatic male flowers are used in Java with rice-flour in making scented medicinally applied powders. In Peninsular Malaysia a decoction of the root is administered against fever, after child birth, and against stomachache and cholera. In Sumatra the tough wood is used to make small objects. Cultivated as an ornamental in Burma.

    Notes — This species, although very wide spread, is remarkably uniform in shape throughout its distribution range. There are only two characters that change conspicuously from west to east. Firstly, east of Borneo the lower leaf surfaces of the plants are often densely velvety hairy as opposed to (almost) glabrous west of Sulawesi. This was reason for Pax and Hoffmann (1914) to describe the variety pilosus. Secondly, in eastern New Guinea and the Solomon Islands the conspicuous hair tufts in the axils of the basal secondary leaf veins are often (Papua New Guinea) or always absent (Solomon Islands).

20. Mallotus havilandii Airy Shaw (ex sect. Hancea)

   Mallotus havilandii Airy Shaw, Kew Bull. 20 (1966) 39; Kew Bull. 23 (1969) 82; Kew Bull. Add. Ser. 4 (1975) 161, 163; Slik & Welzen, Blumea 46 (2001) 52, Fig. 20, Map 8; in Slik, Tropenbos-Kalimantan ser. 4 (2001) 196, Fig. 7.20, Map 7.8; Welzen et al., Blumea 51 (2006) 368. — Type: Kalong for Haviland 1720 (K holo; L iso), Sarawak, Mt Koum.

    Also check: Slik, Macaranga and Mallotus species of Borneo

Small trees up to 8 m tall, dbh up to 15 cm, dioecious. Outer bark smooth. Indumentum dense, simple to tufted to stellate. Branches smooth, lenticellate, glabrescent, gland-dotted, nodes not to only slightly swollen. Stipules semi-persistent, narrowly triangular, 6–11 by 0.6–1.5 mm, margin entire, involute, apex acute, hairy, not gland-dotted. Leaves simple, opposite, unequal, one of each pair much smaller, small leaf sometimes caducous. Non-reduced leaves: petioles 2–5 by 1.5–2.5 mm, hairy, gland-dotted; blade ovate to obovate, 11–24.5 by 3.5–8 cm, length-width ratio 2.8–4.1, chartaceous, base cordate, oblique, margin wavy to apically dentate to serrate, marginal glands 9–15 per leaf side, apex cuspidate to aristate, upper surface smooth, hairy, especially on veins, not gland-dotted, basal macular glands present, similar to apical glands, often glands present on all veins, 2–20 mm from margin, lower surface smooth, hairy, gland-dotted, domatia absent; venation basally palmate to apically pinnate, veins 7 from petiole insertion, 11–16 per side along midrib, ending parallel to or apically in margin. Reduced leaves cordate, 15–30 by 14–22 mm, margin dentate, apex acuminate, indumentum and glands similar to non-reduced leaves, macular glands on upper surface, especially basally along veins. Inflorescences in axils of reduced leaves, unbranched, solitary. Flowers actinomorphic. Staminate inflorescences thyrse-like racemes, up to 23 cm long, basally c. 1 mm thick, hairy, gland-dotted, nodes up to 47 per inflorescence, flowers up to 3 per node; bracts persistent, narrowly triangular, 3–7 by 0.5–1 mm, margin entire, involute, apex acute, hairy, gland-dotted; buds globose with acuminate apex, hairy. Staminate flowers 2.5–3.5 mm in diameter; pedicels 3–4 mm long, hairy, rarely gland-dotted; sepals 4, more or less connate, elliptic to ovate, recurved, 1.8–2.5 by 1–1.8 mm, margin entire to irregular, apex acute, hairy, gland-dotted; stamens up to 44; filaments up to 2 mm long, glabrous; anthers 0.3 by 0.4 mm, basifixed; thecae 2, parallel, extrorse, opening lengthwise, glabrous; connective not widened to widened. Pistillate inflorescences racemes, up to 27 cm long, basally 1–1.5 mm thick, hairy, gland-dotted, nodes up to 12 per inflorescence; bracts persistent, narrowly triangular, 4–10 by 0.8–1 mm, margin entire, involute, apex acute, hairy, gland-dotted. Pistillate flowers pedicels 1.5–5 mm long, hairy, gland-dotted; sepals 2–5, sometimes apically split, more or less connate to free, ovate to narrowly ovate, c. 4 by 1.3 mm, margin entire to irregular, apex acute, hairy abaxially only, gland-dotted; ovary 3-locular, echinate, densely hairy, gland-dotted, spines up to 0.5 mm long, hairy, long simple hairs apically, rarely gland-dotted; ovules 1 per locule, axillary, apotropous, ascending; styles persistent, 1–1.5 mm long, hairy, gland-dotted; stigmas up to 4.5 mm long, densely covered with granulate papillae on the inside, hairy abaxially, gland-dotted outside. Fruit an echinate, lobed capsule, c. 10 by 7 mm, loculicidal-septicidal, light green to green, hairy, gland-dotted; carpel inside hairy, gland-dotted; column unknown. Seeds glossy, roughly globose, c. 6 by 6 by 5 mm; hilum deltoid to cordate, c. 1.5 by 2 mm.

    Distribution — Only known from Sarawak, west of Kuching.

    Habitat & Ecology — Only found on limestone. Altitude: up to 166 m.

21. Mallotus insularum (Airy Shaw) Slik (ex sect. Hancea)

   Mallotus insularum (Airy Shaw) Slik in Slik & Welzen, Blumea 46 (2001) 54, Fig. 21, Map 8; in Slik, Tropenbos-Kalimantan ser. 4 (2001) 198, Fig. 7.21, Map 7.8; Welzen et al., Blumea 51 (2006) 368. — Mallotus miquelianus (Scheff.) Boerl. var. insularum Airy Shaw, Kew Bull. 20 (1966) 40. — Type: Buwalda 4575 (K holo; L iso), Moluccas, Tanimbar Islands (Timor Laut), Pulau Jamdena.

Shrubs up to 3.5 m tall, dioecious. Indumentum dense, simple to tufted to stellate. Branches smooth to slightly fissured, sometimes lenticellate, glabrescent, gland-dotted or not, nodes slightly swollen. Stipules usually caducous, sometimes persistent, triangular to ovate, 1.5–4.5 by 0.4–1 mm, margin irregular to regular, apex acute, hairy abaxially only, gland-dotted or not. Leaves simple, opposite, unequal. Non-reduced leaves: petioles 1–5 by 0.5–1 mm, hairy, gland-dotted or not; blade elliptic to obovate, 5–14 by 2.5–5.5 cm, length-width ratio 1.9–4, chartaceous, base cuneate to truncate to cordate, sometimes slightly oblique, margin wavy to dentate, marginal glands 3–13 per leaf side, apex acute to acuminate, upper surface smooth, glabrous to hairy, especially on midrib and veins, gland-dotted or not, basal macular glands 2–6 on veins, 1–12 mm from midrib, apical macular glands absent, lower surface smooth, hairy, especially on veins, gland-dotted, domatia absent; venation pinnate, veins 7–11 per side, ending parallel to or apically in the margin. Reduced leaves cordate to ovate to obovate, 7–42 by 5–25 mm, margin wavy to dentate, apex acute, upper surface with 2–6 basal macular glands, glabrous, gland-dotted, lower surface hairy, gland-dotted. Inflorescences in axils of reduced leaves, unbranched, solitary. Flowers actinomorphic. Staminate inflorescences thyrse-like racemes, up to 9.5 cm long, basally 0.5–1 mm thick, hairy, gland-dotted or not, nodes up to 30 per inflorescence, flowers up to 3 per node; bracts persistent, ovate, 1.3–4 by 0.5–0.8 mm, margin entire to irregular, apex acute, hairy abaxially, glabrous to hairy adaxially, not gland-dotted; buds (globose to) ovate, hairy, gland-dotted or not. Staminate flowers up to 4 mm in diameter, yellow green; pedicels 1–3 mm long, hairy, gland-dotted or not; sepals 2–3, more or less connate, ovate to elliptic, c. 2.5 by 1.5 mm, margin irregular, apex acute, hairy abaxially only, not gland-dotted; stamens up to 53; filaments 2.3–3.5 mm long, glabrous; anthers 0.2–0.3 by c. 0.4 mm, basifixed; thecae 2, parallel, extrorse, opening lengthwise, glabrous; connective usually widened. Pistillate inflorescences racemes, up to 5 cm long, basally c. 1 mm thick, hairy, gland-dotted sparsely, nodes up to 6 per inflorescence; bracts persistent, narrowly triangular to triangular, 3–5 by 0.8–1 mm, margin entire, involute, apex acute, hairy. Pistillate flowers pedicels 4–5 mm long, hairy; sepals 2–4, more or less connate to free, ovate to narrowly triangular, 2–2.5 by c. 1 mm, margin entire to irregular, apex acute, hairy; ovary 3-locular, echinate, densely hairy, gland-dotted, spines up to 0.5 mm long, hairy, long simple hairs apically, rarely gland-dotted; ovules 1 per locule, axillary, apotropous, ascending; styles persistent, 0.5–1 mm long, hairy, gland-dotted; stigmas up to 2.5 mm long, densely covered with granulate papillae on the inside, hairy abaxially, gland-dotted outside. Fruit an echinate, lobed capsule, 6–7 by c. 5 mm, loculicidal-septicidal, hairy, gland-dotted; carpel inside hairy; column unknown. Seeds unknown.

    Distribution — Lesser Sunda Islands, the Moluccas and Tanimbar Islands.

    Habitat & Ecology — In primary (monsoon) forest; dry places. Altitude: up to 1000 m.

    Notes — Plants from Bali and Lombok have less pronounced dentate leaves in comparison to the plants from the Moluccas and the Eastern Lesser Sunda Islands. From the mountains of Flores an aberrant specimen (Schmutz 2860) was collected which differs from the type by its less conspicuously unequal opposite leaves.

22. Mallotus korthalsii Müll.Arg. (sect. Rottleropsis)

    Mallotus korthalsii Müll.Arg. in DC., Prodr. 15, 2 (1866) 976; Merr., Enum. Philipp. Fl. Pl. 2 (1923) 433; Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 170; Slik, Priyono & Welzen, Gard. Bull. Singapore 52 (2000) 58, Fig. 29; in Slik, Tropenbos-Kalimantan ser. 4 (2001) 96, Fig. 5.32; S.E.C.Sierra et al., Blumea 52 (2007) 72, map 13. — Rottlera korthalsii (Müll.Arg.) Scheff., Ann. Mus. Bot. Lugduno-Batavum 4 (1869) 124. — Type: Korthals s.n. (L barcode L0293734), Borneo, Kalimantan Selatan, Gunung Sakumbang.

    Mallotus petanodon Airy Shaw, Kew Bull. 32 (1978) 401. — Type: Whitmore 3071 (holo K), SE Indonesia, Borneo, Kalimantan Timur (see note 1).

    Also check: Slik, Macaranga and Mallotus species of Borneo

Shrubs to large trees, up to 25 m high, dbh 4–69 cm, dioecious. Outer bark smooth. Indumentum composed of simple and tufted hairs and orange to dark red glandular hairs. Stipules triangular, 1.5–3 by 0.8–1 mm, early caducous, margin entire, apex acute to acuminate, densely to scatteredly hairy outside, scatteredly hairy inside. Leaves opposite; petiole 10–90 by 0.5–1.5 mm, densely to sparsely hairy; blade ovate to obovate, 5.5–25 by 3–15 cm, length/width ratio 1.2–1.7, membranaceous to chartaceous, base rounded, obtuse to cuneate, margin entire to dentate, with glandular teeth, apex caudate, upper surface scatteredly hairy on midrib and nerves, extrafloral nectaries marginal from base (to apex), 3–10 per side, 0.5–8 mm from margin, elliptic to orbicular, 0.3–0.8 by 0.3–0.8 mm, lower surface scatteredly hairy, densely covered with glandular hairs, domatia sometimes present, venation triplinerved, nerves 3–5 per side (see note 2), looping and closed near margin or ending in margin. Inflorescences racemes, single, erect; axes densely hairy; peduncle 5–12 mm long; bracts deltoid, c. 1 by 0.8–1 mm, persistent to caducous, margin entire, apex acute, densely hairy outside, inside glabrous; bracteoles absent. Flowers: pedicels densely hairy; sepals persistent, margin entire, outside sparsely to scatteredly hairy with simple and glandular hairs, glabrous inside. Staminate inflorescences 3–22 cm long, nodes per branch 20–25, with 3–15 flowers per bract. Staminate flowers 2–5 mm diam.; pedicels 0.5–3 mm long; sepals 4 or 5, ovate to elliptic, 1.8–2.5 by 0.5–1.5 mm, free; stamens 23–33, filaments 2.3–3 mm long, free, hairy to glabrous, thecae ovoid, 0.1–0.2 by 0.2–0.3 mm, hairy, connective not widened; pistillode absent. Pistillate inflorescences 4–24 cm long, nodes per branch 15–80, with 1 flower per bract. Pistillate flowers 3–3.5 mm diam.; pedicels 1.5–3 mm long; calyx 4–6-lobed, lobes triangular to narrowly triangular, 1.5–3 by 0.7–1.2 mm, margin subentire; ovary 1.5–2.1 by 1.9–2.1 mm, 3-locular, densely hairy, scatteredly covered with glandular hairs; style 0.2–0.5 mm long; stigmas 2–3 by 1.5–2 mm. Fruits capsules, 5–11 by 10–14 mm, green, opening septicidally-loculicidally, surface spiny, densely to sparsely hairy, scatteredly covered with glandular hairs; spines (20–)100–200, ± straight, 1–1.7 by (0.1–)0.3–0.8 mm, hairy; wall 1–1.5 mm thick, densely hairy to glabrous inside; column 5–6 by c. 4 mm. Seeds ± globose, 3–5 by 4–5 by 4–5 mm, surface smooth, shiny, dark brown.

    Distribution — Peninsular Malaysia, Sumatra, Java, Borneo, Philippines.

    Habitat & Ecology — Primary and secondary dipterocarp forest, marshy and savannah areas; on ridges, along forest edges, road and river sides, on steep slopes and in logged areas; on limestone. Altitude: sea level up to 400 m. Flowering: April to September; fruiting: March to November.

    Vernacular name — Sumatra: Kayu jobar.

    Notes — 1. Airy Shaw (1978) reported M. petanodon to be different from M. korthalsii because of the presence of “long spreading repand teeth in the upper half of the leaves, and the almost smooth cocci of the capsules”. However, these features are also present in M. korthalsii over its entire distribution and are part of the variation of the species. Furthermore, Airy Shaw had only seen one sample from Kalimantan Timur, Borneo (Whitmore 3071) and based on this single collection he described M. petanodon. Until now, we have seen three collections from Kalimantan Timur (Kato & Wiriadinata B5524; Keßler & Arbainsyah B1361 and Wiriadinata 811) and they all have the typical characters of M. korthalsii. Therefore, M. petanodon is synonymized with M. korthalsii.

2. The first pair of nerves end in the upper half of the leaf blade.

23. Mallotus lackeyi Elmer (sect. Stylanthus)

    Mallotus lackeyi Elmer, Leafl. Philipp. Bot. 4 (1911) 1298; Pax & K.Hoffm. in Engl., Pflanz

enr. IV.147. vii (1914) 176; Merr., Enum. Philipp. Fl. Pl. 2 (1923) 433; Philipp. J. Sci. 29 (1926) 382; Univ. Calif. Publ. Bot. 15 (1929) 158; Meijer, Bot. News Bull. (1967) 51; Airy Shaw, Kew Bull. 26 (1972) 308; Whitmore, Tree Fl. Malaya 2 (1973) 114; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 172; Alph. Enum. Euphorb. Philipp. Isl. (1983) 36; Keßler & Sidiyasa, Tropenbos Series 7 (1994) 133; Slik, Priyono & Welzen, Gard. Bull. Singapore 52 (2000) 58, Fig. 30; Slik & Welzen, Blumea 46 (2001) 38, Fig. 15, Map 6; Slik, Priyono & Welzen in Slik, Tropenbos-Kalimantan ser. 4 (2001) 96, Fig. 5.33; Slik & Welzen in Slik, Tropenbos-Kalimantan ser. 4 (2001) 182, Fig. 7.15, Map 7.6. — Type: Elmer 13017 (PNH holo, lost; EDI, L, NY iso), The Philippines, Palawan, Puerto Princesa (Mt Pulgar).

    Mallotus sanchezii Merr., Philipp. J. Sci. 7 (1912) 402. — Type: Quadras (PNH holo, not traced), Philippines, Mindanao, Misamis province, Talisayan.

    Macaranga leightonii Whitmore, Kew Bull. 39 (1984) 607. — Type: Not yet indicated (see: Whitmore, Gen. Macaranga Prodr., 2008: 283).

    Also check: Slik, Macaranga and Mallotus species of Borneo

Shrub to small tree up to 17 m tall, dbh up to 20 cm; bole straight to crooked; crown spreading. Outer bark smooth to slightly fissured, knobby; sapwood soft. Indumentum dense. Branches lenticellate, glabrescent, densely gland-dotted. Stipules usually persistent, inserted up to c. 3 mm above petiole insertion, narrowly triangular to triangular, 5–9 by 2–3.5 mm, margin entire to irregular, involute, apex acute (to acuminate), densely hairy, especially along midrib, gland-dotted abaxially or not. Leaves alternate to apically subopposite, peltate, distance between leaf blade base and petiole insertion 7–38 mm; petiole 25–180 by 0.6–2.3 mm, basally thickened, densely hairy, glabrescent, gland-dotted sparsely to densely; blade ovate (to orbicular), 5–33.5 by 3–21 cm, length-width ratio 1.2–2.3, base rounded (to truncate), margin (slightly) dentate to irregularly wavy, marginal glands 18–33 per leaf side, apex aristate, upper surface smooth, sparsely hairy, especially on veins and along margin, gland-dotted sparsely to densely, basal macular glands 0–5 on veins, 4–18 mm from petiole insertion, apical macular glands 0–7 on veins, 1–4 mm from margin, lower surface smooth, densely hairy, gland-dotted sparsely to densely, domatia absent; venation palmate, 7—10 veins from petiole insertion, 5–9 secondary veins per side along midrib, ending parallel to or in the margin. Staminate inflorescences up to 32 cm long, basally 1–4 mm thick, usually solitary, densely hairy, gland-dotted, nodes up to 64 per inflorescence, flowers 1–15 per node; bracts caducous, triangular to narrowly triangular, 1.5–6.5 by 0.7–1.3 mm, margin entire, apex rounded to acute, densely hairy, usually glabrous adaxially, gland-dotted; buds ovate with acute apex, densely hairy. Staminate flowers 3–3.5 mm in diameter, white to yellow, not to slightly fragrant; pedicels 0.8–3 mm long, hairy, not gland-dotted; sepals 3–5, free to basally connate, narrowly elliptic to elliptic to ovate, sometimes recurved, 1.8–2.5 by 0.7–1 mm, margin entire, apex acute to slightly acuminate to cuspidate, hairy abaxially, especially basally, glabrous adaxially, gland-dotted or not; stamens up to 34, filaments up to 3.3 mm long, glabrous; anthers 0.2–0.3 by 0.4–0.5 mm, basifixed; connective widened. Pistillate inflorescences up to 37 cm long, basally 2–4 mm thick, densely hairy, gland-dotted, nodes up to 54 per inflorescence; bracts caducous, linear to triangular, 3.5–7 by 0.4–1.5 mm, margin entire, involute, apex acute, hairy, glabrous to hairy adaxially, gland-dotted outside only. Pistillate flowers white to cream; bracteoles caducous, linear, 1.3–3.5 by 0.2–0.7 mm, margin entire, involute, apex acute, hairy, gland-dotted; pedicels up to 7 mm long, often recurved at fruit maturity, hairy, gland-dotted; calyx persistent, 3–6-lobed, basally connate, narrowly triangular to triangular, 3.5–5 by 0.6–1.8 mm, margin entire, apex acute, hairy, glabrous adaxially, gland-dotted; ovary 3-locular, densely echinate, densely hairy, gland-dotted, spines up to 2 mm long, densely hairy, gland-dotted or not; style persistent, up to 1 mm long, hairy, gland-dotted; stigmas 4–5 mm long, densely covered with granulate papillae on inside, hairy abaxially, gland-dotted outside. Fruit an echinate, lobed capsule, 10–13 by 3.5–7 mm, light green to yellow-green (to reddish), hairy, gland-dotted; carpel inside hairy, not gland-dotted; column 3.5–5 by 5–5.5 mm. Seeds glossy, globose, c. 5 by 4.5 mm; hilum deltoid to cordate, c. 1.5 by 1.5–2 mm.

    Distribution — Borneo and the Philippines.

  dots = M. lackeyi; squares = M. thorellii.

    Habitat & Ecology — Locally common in primary and secondary forests where it prefers open places; often along streams and rivers, in gaps, along roads, but also growing in the forest understorey; on dry and periodically flooded areas; on most soil types, from yellow to brown sand to clay to sandstone to limestone. Altitude: up to 800(–2693) m.

    Note — This species is very uniform in shape throughout its distribution range. In Sarawak and Peninsular Malaysia specimens of M. peltatus can look very similar to M. lackeyi. These M. peltatus specimens can be recognised by their less dense and more tufted indumentum, their upper leaf surface that is not gland dotted, their small and early caducous stipules, and their fruits with long styles.

24. Mallotus lancifolius Hook.f.  (sect. Rottleropsis)

    Mallotus lancifolius Hook.f., Fl. Br. India 5 (1887) 434; Welzen & S.E.C.Sierra, Blumea 51 (2006) 376, fig. 2a, 3i; map 2. — Lectotype (Welzen & Sierra, 2006): Maingay KD 1451 (holo K; iso K), Malaya.

    Mallotus moritzianus Müll.Arg. var. laevigatus Müll.Arg., Linnaea 34 (1865) 191. — Mallotus laevigatus (Müll.Arg.) Airy Shaw, Kew Bull. 19 (1965) 328; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 52; Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 160; Kew Bull. 36 (1981) 326; Chakrab., J. Econ. Taxon. Bot. 6 (1985) 496. — Macaranga laevigata (Müll.Arg.) Backer & Bakh.f., Fl. Java 3 (1968) 649, nom. inval. (‘laevigatus’; see note 3) — Lectotype (Welzen & Sierra, 2006e): Zollinger 1554 (holo L; iso BM, FI, G (2x), M, S, ZT), Java.

    Rottlera glaberrima auct. non Hassk.: Rchb.f. & Zoll., Linnaea 28 (1856) 313; Miq., Fl. Ned. Ind. 1, 2 (1859) 393 (all p.p., excl. type). — Mallotus glaberrimus auct. non (Hassk.) Müll.Arg.: J.J.Sm., Meded. Dept. Landb. Ned.-Indië 10 (1910) 435; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 192; Merr., J. Straits Branch Roy. Asiat. Soc., Special number (1921) 338; Backer & Bakh.f., Fl. Java 1 (1963) 484 (all p.p., excl. type). See note 2.

Mallotus wrayi auct. non King ex Hook.f.: Whitmore, Tree Fl. Malaya 2 (1973) 115, p.p. (‘M. lanceifolius’).

Shrubs to trees up to 13 m high, dbh up to 20 cm, crown up to 5 m high; flowering branchlets 2–3 mm thick, flattened and somewhat ridged when dry; apical parts stellately hairy, glabrescent, mainly remaining hairy on nodes. Outer bark smooth to shallowly, finely fissured, whitish to yellow-brown to pale brown to reddish grey-brown to brown-black, 0.5 mm thick, fibrous; inner bark c. 4 mm thick, hard, pale greenish to pinkish to ochre to pale brown to outside brown-red, striped; sapwood white to straw, hard. Indumentum usually not dense, long simple hairs and stiff, short stellate hairs, glandular scales orange or yellowish. Stipules and bracts of the terminal buds triangular, 2–10 by 2–3 mm, usually long, appressed hairy outside, especially on the central part, margins becoming glabrous and more or less translucent cartilaginous, stipules very early caducous. Leaves opposite (alternate in young stages), similar in shape, one somewhat smaller; petiole 0.8–5(–8.5, see note 5) cm long, round, basally and apically pulvinate, upper pulvinus relatively thin to oblique with blade base partly overlaying, slightly hairy; blade ovate to elliptic (to obovate), 5–28 by 2.3–10 cm, length/width ratio 1.9–4.3, symmetric, membranous to subcoriaceous; base (emarginate, see note 5, to) rounded to attenuate to cuneate, very base usually narrowly cordate, margin (entire to) serrulate to serrate in upper two-third with usually coarse teeth, apex (acute to) caudate; upper surface basally with one pair of extrafloral nectaries, up to 1.3 by 0.5 mm, in upper third usually several, usually deep, extrafloral nectaries (mainly 0.3 by 0.3 mm) on and in between nerves (S 63606 exceptional in having additional small extrafloral nectaries along basal part of midrib), at most slightly hairy (to hairy) on midrib and basal nerves; lower surface sparsely hairy on venation, glabrescent, glandular scales very few, almost absent, hair tuft domatia present; venation triplinerved, flat to slightly raised above, raised below, 8–12 nerves per side, veins scalariform, veinlets reticulate. Inflorescences racemes, axillary to (pseudo)terminal, mainly single, reddish, slightly to densely covered with stiff hairs; staminate ones short, up to 10 cm long, or long, up to 25 cm long, diameter 0.8–1.4 mm, with cymosely groups of more than 10 flowers; pistillate ones up to 14 cm long, 1.2–2.1 mm thick, a single flower per node; bracts ovate to triangular, 2–8 by 1–2.5 mm, often stiff and patent, later recurved, densely hairy on back with usually glabrous to glabrescent margins. Staminate flowers 3–4 mm diam., greenish to white (to pale yellow), buds ovoid; pedicel 3–6 mm long, with abscission zone in basal two-third, hairy, less so above abscission zone; sepals 3(–5), ovate to elliptic, 2.3–4.3 by 0.8–2.2 mm, reflexed, hairs and glandular scales outside; stamens 30–65, filaments up to 1.7–4 mm long, glabrous, anthers 0.5–0.7 by c. 0.3 mm, thecae distinctly separated from each other by connective, latter with apically glandular cells. Pistillate flowers 2.5–3 mm diam., quickly increasing in size, yellowish; pedicel 1.8–4.5(–7 in fruit) mm long, abscission zone ± halfway, densely hairy; sepals breaking up to about 4 or 5, triangular, 2.5–6 by 0.4–1.7 mm, few to many stellate hairs outside, reflexed; ovary 3-locular, c. 2.8 by 1.7 mm, hairy outside, covered with not very dense spines up to 2 mm long, latter dark red, pointing in all directions, with few hairs, especially basally; style 0.5–1.5 mm long, bright red, stigmas 2.7–8.3 mm long, 0.2–0.7 mm wide, bright red, with usually branching papillae above, less than 1 mm long, somewhat sericeous beneath. Fruits lobed capsules, 8–16 by 4–9 mm, yellow-green, spines up to 2.5 mm long, breaking off; wall woody, 1.2–1.5 mm thick; column T-shaped, up to 4 mm high. Seeds subglobose, 5.8–9.5 by 5.2–9 by 5–5.2 mm, shiny.

    Distribution — Malay Peninsula, Sumatra, Java, Borneo. According to Chakrabarty (1985) also on the Andaman and Nicobar Islands.

    Habitat & Ecology — In understorey of primary dipterocarp, secondary, submontane, and montane forest, selectively logged forest; soil moist, yellow clay, volcanic rocks, sandstone. Altitude 30–1650 m. Flowering: January to April, July, August, October to December; fruiting: January, March, June, August to November.

    Vernacular names — Sumatra: Eroepeka (Enggano); Talog (Malay Palembang). Borneo: Sarawak: Bantas.

    Notes — 1. The epithet laevigatus is the older (Müller Argoviensis, 1865) than the epithet lancifolius (Hooker, 1887), but it was first used on an infraspecific level; only much later Airy Shaw (1965) was the first to use it as a species epithet. This makes the epithet lancifolius the oldest epithet on the species level.

2. Airy Shaw (1965) clearly explains the wrong interpretation of the species Rottlera glaberrima as described by Hasskarl (Flora 25, 2, Beibl., 1842, 41). Hasskarl’s description is extremely brief, but shows the leaves to be alternate. Airy Shaw correctly identified it as a Macaranga species (Macaranga glaberrima (Hassk.) Airy Shaw). Reichenbach & Zollinger (1856) incorrectly identified a specimen from the Bogor Botanical Garden with Hasskarl’s name. Their specimen is a true Mallotus with opposite leaves (one smaller than the other). Several authors (e.g., Pax & Hoffmann, 1914; Backer & Bakhuizen van den Brink, 1963) considered the name and description by Reichenbach & Zollinger as a later homonym, but this is not the case, because Reichenbach & Zollinger clearly refer to Hasskarl. Müller Argoviensis (1865) interpreted Reichenbach & Zollinger’s description correctly as a Mallotus and created the new combination Mallotus glaberrimus. He too referred to Hasskarl, which means that this name is also a synonym of Macaranga glaberrima (Hassk.) Airy Shaw. Luckily, Müller Argoviensis (1865) also described a new variety (laevigatus) under Mallotus moritzianus Müll.Arg., which is the same taxon. Therefore, the combination Mallotus laevigatus (Müll.Arg.) Airy Shaw is the correct name.

3. Unfortunately, Backer & Bakhuizen van den Brink, in their addenda to the Flora of Java (1968) accidentally reversed the solution of Airy Shaw and they unintentionally created a new combination: Macaranga laevigatus. Luckily, an invalid name because references are missing.

4. The leaves on Java and partly on Borneo are narrower and ovate, on Sumatra more elliptic and larger, while many specimens on Borneo have relatively broad leaves. The fruits on Java are smallest, those on Sumatra and Borneo are usually larger.

5. Two specimens from the Malay Peninsula have extra large leaves and long petioles (up to 8.5 cm long): SF 30028 and SF 30147. The base of the leaves is rounded to even emarginate. The two specimens are sterile and might be suckers.

25. Mallotus lauterbachianus (Pax & K.Hoffm.) Pax & K.Hoffm. (sect. Rottleropsis)

    Mallotus lauterbachianus (Pax & K.Hoffm.) Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 157; S.E.C.Sierra et al., Blumea 52 (2007) 74, map 7. — Coelodiscus lauterbachianus Pax & K.Hoffm. in Pax, Feddes Repert. Spec. Nov. Regni Veg. 8 (1910) 481. — Type: Weinland 257 (ULM?, n.v.), Papua New Guinea, Bumi.

    Mallotus lullulae Airy Shaw, Kew Bull. 21 (1968) 384. — Type: Brass 28701 (holo K; iso L, S), Papua New Guinea, Woodlark Island, Kulumadau.

Shrubs to small trees, 2–9 m high; branches hollow, dioecious. Indumentum composed of simple and stellately-tufted hairs, glandular hairs absent. Stipules narrowly triangular, c. 4.5 by 1 mm, early caducous, margin entire, apex acute, sparsely hairy outside, glabrous inside. Leaves opposite (to subopposite); petiole 10–103 by 1–2 mm, scatteredly hairy; blade ovate to elliptic, 10.5–30 by 6.5–14 cm, length/width ratio 1.6–2.1, chartaceous, base rounded, obtuse to cuneate, margin crenulate, with glandular teeth, apex cuspidate, upper surface glabrous, extrafloral nectaries along the midrib, 5–18 per side, elliptic to orbicular, 0.8–1 by 0.7–1 mm, lower surface scatteredly hairy, glabrescent, domatia present, venation triplinerved, nerves 5–9 per side, indistinctly looping and closed near margin. Inflorescences single, erect; axes sparsely hairy; bracts triangular to narrowly triangular, 2.5–5 by 0.8–1.8 mm, persistent, margin entire, apex acute, sparsely hairy outside, glabrous inside; bracteoles narrowly triangular, 1–3 by 0.3–1 mm, persistent. Flowers: pedicels densely hairy; sepals free, margin entire, apex acute, persistent, densely to sparsely hairy outside, scatteredly hairy to glabrous inside. Staminate inflorescences racemes, 5–14 cm long; peduncle 5–60 mm long; nodes per branch 5–8, with 8–10 flowers per bract. Staminate flowers 5–8 mm diam.; pedicels 6–10 mm long; sepals 3, ovate to elliptic, 4–6 by 2–3 mm; stamens 30–65, filaments 1.8–5 mm long, free, hairy, thecae ovoid, 0.3–0.4 by 0.2–0.25 mm, glabrous, connective widened; pistillode present. Pistillate inflorescences racemes or umbel-like, 2.7–4 cm long; peduncle 20–27 mm long; nodes per branch 1–4, with 1–3 flowers per bract. Pistillate flowers 3–4 mm diam.; pedicels 4–6 mm long; sepals 3, narrowly to linear-triangular, 6–7.1 by 0.8–1 mm; ovary 2.5–2.8 by 2–2.3 mm, 3-locular, densely hairy; style c. 2 mm long; stigmas 3–4.5 by 0.8–1 mm. Fruits unknown.

    Distribution — Endemic to New Guinea.

    Habitat & Ecology — Frequent in secondary forest. Altitude 10–500 m. Flowering and fruiting: July to November.

26. Mallotus leucocalyx Müll.Arg. (sect. Rottleropsis)

    Mallotus leucocalyx Müll.Arg. in DC., Prodr. 15, 2 (1866) 970; S.E.C.Sierra et al., Blumea 52 (2007) 75, map 5, plate 1a-c. — Type: Unit. Stat. Explor. Exped. s.n. (holo G-DC, IDC microfiche 800; iso GH, US), Philippines, Mindanao, Caldera.

Shrubs to small trees, up to 8 m high, dioecious. Outer bark smooth. Indumentum composed of simple, tufted and stellately-tufted hairs and yellow to orange glandular hairs. Stipules triangular to narrowly triangular, 4–10 by 2–4 mm, persistent to caducous, margin entire to subentire, apex cuspidate, scatteredly hairy to glabrous outside, densely hairy inside. Leaves opposite; petiole 20–100 by 1.3–1.8 mm, densely to sparsely hairy; blade ovate to obovate, 10.5–26 by 5.8–15 cm, length/width ratio 1.5–1.9, membranaceous, base shallowly emarginate, rounded to cuneate, margin denticulate to dentate, with glandular teeth, apex acute to acuminate, upper surface scatteredly hairy to glabrous on midrib, extrafloral nectaries marginal throughout the blade, up to 12 per side, 1–7 mm from margin, sometimes also all over the blade, elliptic to orbicular, 0.3–0.5 by 0.3–0.5 mm, lower surface sparsely hairy, sparsely covered with glandular hairs, domatia absent, venation triplinerved, nerves 8 or 9 per side, looping and closed near margin or ending in margin. Inflorescences racemes, single, erect; peduncle 15–28 by c. 1 mm; axes densely hairy; bracts trilobed, persistent, margin entire, apex acuminate to cuspidate, densely hairy outside, glabrous inside; bracteoles absent. Flowers: pedicels densely hairy; sepals free, persistent, margin entire, outside densely to sparsely hairy and scatteredly covered with glandular hairs, glabrous inside. Staminate inflorescences 3.5–12 cm long, nodes per branch 8–23, with 2–7 flowers per bract; bracts 5–8 by 3–4 mm. Staminate flowers 3–4.2 mm diam.; pedicels 2–4 mm long; sepals 3 or 4, elliptic to obovate, 2.5–2.8 by 0.7–1.2 mm, apex acuminate; stamens 20–40, glabrous, filaments 1–3 mm long, free, pale light yellow, thecae ovoid, 0.2–0.3 by 0.2–0.25 mm, connective widened; pistillode present. Pistillate inflorescences 9.4–16 cm long; nodes per branch 7–15, with 1 flower per bract; bracts 5–10 by 3–5 mm. Pistillate flowers 2.8–3 mm diam.; pedicels 1–2 mm long; sepals 3–6, narrowly triangular, 3.8–4.1 by 0.8–1 mm, apex acute; ovary 1.8–2 by 1.8–2 mm, 3-locular, densely hairy, pale light green, scatteredly covered with glandular hairs; style 1.5–2 mm long; stigmas 4–5 by 1.3–1.5 mm. Fruits capsules, 7–7.5 by 9–11 mm, green, opening septicidally-loculicidally, surface spiny, densely hairy, scatteredly covered with glandular hairs; spines 200–230, curved, 3–4.2 by 0.2–0.3 mm, densely hairy; wall 0.7–1 mm thick, densely to sparsely hairy inside; column 3.4–5 by 3–3.5 mm. Seeds ± globose, 3.2–4.2 by 3–4 by 3–3.8 mm, surface smooth, shiny, light brown.

    Distribution — Myanmar, Thailand, Peninsular Malaysia, Sumatra, Java, Borneo (Sabah), Philippines, Sulawesi.

    Habitat & Ecology — In understorey of primary or secondary forests, in open or shaded areas; on clay or limestone. Altitude: sea level up to 650 m. Flowering and fruiting throughout the year.

    Vernacular names — Borneo (Sabah): Dulang jantan (Bajau), Mandahasi (Dusun Kinabatangan), Sempurna.

27. Mallotus leucodermis Hook f. (sect. Polyadenii)

    Mallotus leucodermis Hook.f., Fl. Brit. India 5 (1887) 441; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 180; Ridl., Fl. Malay Penins. 3 (1924) 291; Airy Shaw, Kew Bull. 16 (1963) 350, 352; Kew Bull. 20 (1966) 39; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 53; Airy Shaw, Kew Bull. 21 (1968) 397; Whitmore, Tree Fl. Malaya 2 (1973) 116, Fig. 10; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 167; Kew Bull. 36 (1981) 326; Bollendorff, Welzen & Slik, Blumea 45 (2000) 326, fig. 1b, 2a, 3; map 2; Slik, Priyono & Welzen, Gard. Bull. Singapore 52 (2000) 58, Fig. 31; in Slik, Tropenbos-Kalimantan ser. 4 (2001) 96, Fig. 5.34; Bollendorff, Welzen & Slik in Slik, Tropenbos-Kalimantan ser. 4 (2001) 129, Fig. 6.1b, 6.2a, 6.5; Map 6.2. — Mallotus leucodermis Hook.f. var. leucodermis: Airy Shaw, Kew Bull. 21 (1968) 397. — Lectotype (Bollendorff et al., 2000): Maingay KD 1433 (K, holo; iso in K), Malacca.

    Coccoceras muticum Müll. Arg. var. pedicellatum Hook.f., Fl. Brit. India 5 (1887) 424 ('pedicellata'); Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 210; Ridl., Fl. Malay Penins. 3 (1924) 291. — Lectotype (Bollendorff et al., 2000): Griffith KD 4789 (K, holo; iso in K, P), Malacca, Alor Gajah.

    Also check: Slik, Macaranga and Mallotus species of Borneo

Shrub to tree, up to 35(–54) m high, dioecious, more or less glabrous; branchlets not to finely ribbed, rarely lenticellate, white to grey with conspicuous discoid glands to brown. Stipules triangular, rarely falcate, 1.5–2 by 0.8–2 mm, early caducous, sometimes ridged, glabrous, margin entire or rarely slightly dentate, apex acute. Leaves subopposite to alternate and each pair unequal in size; petiole terete to angular, sometimes adaxially channelled, 10–25 by 0.5–3 mm, glabrous or rarely with stellate hairs basally and distally, base and apex more or less pulvinate; blade elliptic to obovate, 5–22 by 2.3–11.5 cm, index 1.6–2.7, chartaceous to coriaceous; base obtuse to acute and sometimes slightly emarginate; margin entire to slightly irregularly crenate with 0–16 glands per leaf side in sinuses; apex acuminate to obtusely cuspidate, sometimes ending in a marginal-type of gland; upper surface glabrous, grey-green to reddish brown to dark brown, 0–4 basally, impressed glands on each leaf-side, elliptic, 1–5 by 0.5–0.8 mm, brown to black, 1–5 mm from petiole attachment, apically none to numerous circular impressed glands among the margin; lower surface olive green to pale- to red- to dark brown, rarely villose hair tuft domatia, sometimes in excavations; venation basally triplinerved, basal nerves mainly ending in margin above middle of blade, 2–6 nerves per leaf side, mainly eucamptodromous, veins scalariform. Staminate inflorescences axillary and terminal, solitary or often 2–4 together, rachis terete to angular, 4–22 cm by 0.4–1.3 mm, 22–79 nodes per rachis with 3–6 flowers per node; bracts triangular to rhomboid, 0.5–0.8 by 0.5–1.5 mm, margin glabrous to ciliate, apex acute to acuminate; pedicels terete to flattened, 2–4 by 0.2–0.3 mm, stellate hairs or glabrous, abscission zone at very base; flowers 2–5 by 3–5 mm in diam., calyx lobes 2–4, ovate to obovate, 2–3 by 1–2 mm, villose to stellate, white to yellow to pinkish to red-brown, recurved, apex acute, stamens 17–41, filaments 1–4 mm long, anthers reniform to oblong, 0.5–0.8 by 0.2–0.3 mm, dorsifixed to rarely basifixed, glabrous, connective slender, rarely split basally, receptacle c. 0.3 mm high by 0.5 mm diam. Pistillate inflorescences axillary to rarely terminal, solitary or 2–4 together, rachis angular, 3–32.5(–60) cm by 0.8–2.5 mm, 14–53 nodes per rachis, 1 flower per node; bracts triangular to rhomboid, 0.5–1.5 by 0.5–1.6 mm, basal part sometimes thickened, blade glabrous to ciliate, apex acute to acuminate; pedicels terete to angular, sometimes pubescent, 6–55 by 0.3–1.3 mm, abscission zone 0.5–4 mm from apex; flowers 2.5–6 by 2–5 mm in diam., calyx lobes 4–6, up to halfway connate, triangular to narrowly triangular, 2–2.6 by 0.5–1.5 mm, persistent, yellow to reddish brown, basally thickened, apically recurved, some villose hairs inside, ovary globose, 2–4-locular, style 0.5–1.5 by 0.5–0.6 mm, stigmas 2–4 by 0.3–1 mm, basally erect, apically recurving. Fruits woody, dehiscent capsules, 8–15 by 10–25 mm in diam., locules globose to ovoid, without wings, 8–9 mm wide; column bell-shaped in lateral view, 6–8 mm long by 5–7 mm wide basally and 1–3 mm wide apically. Seeds globose to ovoid, 5–8 mm in diam., shiny to dull, surface smooth or sometimes with white protuberances, light to dark brown.

    Distribution — Disjunct in W Malesia (Peninsular Malaysia, Borneo, Sumatra) and Irian Jaya.

    Habitat & Ecology — Primary mixed Dipterocarp forest, secondary forest, forest edges, logged areas, often riparian, in swampy areas. Alt.: sea level to 1440 m. Flowering: Mar.–Nov.; fruiting: Mar.–Nov.

    Vernacular names — Peninsular Malaysia: Teropok (Temuan). Sumatra: Toebo-lat-oeding, toebobalie pajo, toebo-lala, suruchon pajo. Borneo: Galungan.

    Notes — 1. Whitmore's (1973) fruit description does not correspond with the fruits of M. leucodermis. He probably described the winged, beaked fruits of M. sumatranus. The corresponding figure, however, shows a coccus without wings which could be M. leucodermis.

    2. The three specimens of Irian Jaya differ from the W Malesian specimens in their relatively thick petioles, very rare marginal glands, their more conspicuous veins and their (fuchsia) red discoid glands on the inflorescences.

    3. Mallotus leucodermis and M. muticus are sympatric and are impossible to identify with certainty without pistillate flowers or fruits. The following list of vegetative characters may be helpful for identification:

28. Mallotus longinervis M. Aparicio (sect. Rottleropsis)

    Mallotus longinervis M.Aparacio in S.E.C.Sierra et al., Blumea 52 (2007) 76, fig. 6, map 10. — Typus: S (Banyeng ak Nudong & Sibat ak Luang) 26206 (holo L; iso A, BO, K, SAN, SING), Borneo, Sarawak, Kuching, Semengoh Arboretum, 12th mile Penrissen Road.

Small trees, 2–8 m high, dioecious. Indumentum composed of simple and stellately-tufted hairs and yellow to orange glandular hairs. Stipules deltoid, 0.4–0.5 by 0.4–0.5 mm, early caducous, margin entire, hairy, apex acute, densely hairy. Leaves opposite; petiole 10–50 by c. 1 mm, sparsely to scatteredly hairy; blade elliptic, 10–25 by 3.5–8 cm, length/width ratio 2–3, chartaceous to subcoriaceous, base obtuse to cuneate, margin entire to denticulate, with glandular teeth, apex caudate, upper surface glabrous, extrafloral nectaries marginal in lower half, 4–6 per side, 1–5 mm from margin, elliptic to orbicular, 0.5–0.8 by 0.5–0.8 mm, lower surface scatteredly hairy to glabrous, densely covered with glandular hairs, domatia sometimes present, venation triplinerved, nerves 2 or 3 per side, looping and closed near margin. Inflorescences racemes, single, erect; peduncle 2–9 by 0.5–1 mm; axes densely to sparsely hairy; bracts deltoid, 0.6–0.9 by 0.5–0.9 mm, persistent to caducous, margin entire, apex acute, densely hairy; bracteoles absent. Flowers: pedicels densely hairy; sepals free, persistent, margin entire, apex acute, densely hairy, also with glandular hairs outside, scatteredly hairy to glabrous inside. Staminate inflorescences 8–12 cm long, nodes per branch 10–18, with 3–5 flowers per bract. Staminate flowers 1.8–2 mm diam.; pedicels 0.5–0.8 mm long; sepals 3, obovate, 1.8–2.5 by 1.3–1.5 mm; stamens 18–25, filaments 0.5–1.5 mm long, free, glabrous, thecae ellipsoid, 0.2–0.3 by 0.1–0.2 mm, hairy, connective not widened; pistillode absent. Pistillate inflorescences 6–17 cm long, nodes per branch 6–15, with 1 flower per bract. Pistillate flowers 2–2.5 mm diam.; pedicels 2.5–3 mm long; sepals 5 or 6, narrowly triangular, 1.5–2 by 0.3–0.5 mm; ovary 1–1.3 by 1–1.2 mm, 3-locular, densely hairy, sparsely covered with glandular hairs; style 0.5–1 mm long; stigmas 1.8–2.3 by c. 0.8 mm. Fruits capsules, 5–5.5 by 5.5–6 mm, green, opening septicidally-loculicidally, surface smooth, densely hairy, sparsely covered with glandular hairs; wall 0.1–0.2 mm thick, densely hairy inside; column 2–2.2 by 1.5–2 mm. Seeds globose, 2–2.2 by 1.7–1.8 by 2–2.5 mm, surface rugose, dull, brown.

    Distribution — Endemic to Borneo (Arboretum Semengoh F.R.).

    Habitat & Ecology — Road and river sides. Altitude c. 90 m. Flowering: August; fruiting: November.

    Vernacular name — Borneo: Entaempulor (Iban).

    Note — The collection S 3389 from Sarawak was identified by Airy Shaw as M. milaciformis (see Dubious species, probably M. korthalsii), but we think that it represents M. longinervis. According to the protologue (syntypes not seen) M. smilaciformis should have 3–5 nerves and minute processes (spines), while S 3389 has 2 or 3 nerves and no spines on the fruits. Additionally, M. smilaciformis is reported for Peninsular Malaysia, while S 3389 is found in Borneo. So far, all collections seen of M. longinervis were collected in Sarawak in the Semengoh Arboretum, therefore, it is possibly endemic to that place.

29. Mallotus macrostachyus (Miq.) Müll.Arg. (sect. Mallotus)

    Mallotus macrostachyus (Miq.) Müll.Arg. in DC., Prodr. 15, 2 (1866) 963; Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 165; Corner, Ways. Trees Malaysia, ed. 3, 1 (1988) 306; Welzen, Slik & Bollendorff in Welzen et al., Thai Forest bull., Bot. 28 (2000) 101; Slik, Priyono & Welzen, Gard. Bull. Singapore 52 (2000) 59, Fig. 32; in Slik, Tropenbos-Kalimantan ser. 4 (2001) 97, Fig. 5.35; S.E.C.Sierra & Welzen, Blumea 50 (2005) 254, fig. 2, map 2.— Rottlera macrostachya Miq., Fl. Ned. Ind., Eerste bijv. (1860) 454. — Lectotype (Sierra & Welzen, 2005): Teijsmann HB 3245 (U), Indonesia, Sumatra.

    Mallotus insignis Müll.Arg., Linnaea 34 (1865) 193; in DC., Prodr. 15, 2 (1866) 975. — Type: Schomburgk 52 (holo K), Singapore.

    Mallotus albus auct. non (Roxb.) Müll.Arg.: Müll.Arg., Linnaea 34 (1865) 188, quoad Motley 530 (CGE, K), Indonesia, Borneo; Wallich Numer. List 7820 (K-W mixed), Malay Peninsula, Penang.

    Mallotus barbatus auct. non Wall. ex Müll.Arg.: Müll.Arg., Linnaea 34 (1865) 184, quoad Wallich Numer. List 7820 (K-W mixed), Malay Peninsula, Penang.

    Also check: Slik, Macaranga and Mallotus species of Borneo

Shrubs to small trees up to 16 m high, dbh up to 25 cm, dioecious, occasionally monoecious; crown spreading, flat. Outer bark finely fissured, soft, brownish grey, inner bark fibrous inwards, yellowish green, cambium yellow, sapwood white. Indumentum tomentose to puberulent. Stipules triangular, 2.8–7 by 1–1.8 mm, persistent, margin entire, apex acute. Leaves alternate to apically subopposite; petiole 2–36 by 1–4 mm; blade subpeltate for 1.5 mm, peltation near petiole insertion often thick and frequently curved upwards, broadly ovate to ovate, 9.2–31 by 7.8–25 cm, length/width ratio1–1.3, base truncate, rounded, to slightly cuneate, margin dentate, or serrate, rarely shallowly 2-lobed at widest part of blade, upper surface dark green, basally with 0–2 extrafloral nectaries, 1.3–3 by 0.5–2.5 mm, marginal nectaries 0–9 per side, 1.5–10 mm from margin, 0.6–1 by 0.6–8 mm, lower surface brownish grey, domatia with a dense tuft of woolly hairs, 3-nerved, nerves 7 or 8 per side, mostly ending in the margin. Inflorescences axillary or terminal, unisexual or bisexual, axes light brownish, basally 2–4 mm thick; bracts triangular,1–2.5 by 0.6–10 mm, persistent, margin entire, bracteoles absent. Staminate inflorescences up to 112 cm long, side branches up to 65 cm long, with 1 or 2 flowers per node, nodes per branch up to 120. Staminate flowers 4–4.5 mm diam., fragrant; pedicels 3.2–4.2 mm long; sepals 3 or 4, 2–2.7 by 1.2–1.7 mm; stamens 60–90, filaments 1–3 mm long, white, anthers 0.3–0.35 by 0.2–0.25 mm, light yellow; pistillode present. Pistillate inflorescences spikes, up to 80 cm long, seldomly with side branches (up to 14 cm long), nodes up to 150 (see note 1). Pistillate flowers 3–7 mm diam., sessile; calyx 4- or 5-lobed, connate almost to the apex, 3–5 mm long, lobes 0.5–1.7by0.8–1.5 mm,light brownish;ovary 3-locular, 3.2–4 by1.8–2.2mm, spines individually not visible, style up to 1.2 mm long; stigmas 4–5 mm long, dark brown; staminodes absent. Fruits 11–16 by 10–14 mm, light green with brown tomentum, spines numerous, straight, thin, less hairy, up to 4 mm long, hairs forming a continuous layer, rubbing off; column 5–6 by 1.2–2 mm. Seeds ellipsoid, 4–5 by 4–4.5 by 3.5–4 mm, surface irregular, dark brown; hilum 2.3–2.5 by 1.8–2 mm.

    Distribution — Thailand to Singapore, Sumatra, and Borneo.

    Habitat & Ecology — Locally common in deciduous and evergreen forest, mostly in open, often very disturbed or burned places, along road and river sides, swampy areas; on a large variety of soil types, like shale, limestone, sand, and basalt. Altitude: sea level up to 800 m. Flowering and fruiting the whole year through. The male flowers are visited by Trigona bees.

    Uses — The leaves are used as an antidote against snake-poison. The wood is used as firewood.

    Vernacular names —Malay Peninsula: Selau, Balek angin. Sumatra: Baliek anggien, Batang balik; Si tarak balanggingan. Indonesia (Borneo): Kapit mawat (Kenyah), Kayu balikangin, Kayu kapit, Buah beliangin, Kajoe si tarak balanggingan. Sabah: Dahau (Kadazan Kinabatangan); Dahu (Dusun/Bundu Tuhan); Dauah (Dusun Putatan); Mangabong, Ngoludon, Padauh-padau, Pedau-pedau (Dusun); Tondon tondon (Suluk). Sarawak: Balek angin; Entupak, Kelempah pinggai, Kelipak pinggai, Kerimpah pinggai (Iban). Brunei: Lekon abai, Lokon (Dusun); Entupak, Nentupak (Iban).

    Note — BS 2576 has aberrant female inflorescences, probably infected by a virus.

30. Mallotus macularis Airy Shaw (sect. Rottleropsis)

    Mallotus macularis Airy Shaw, Kew Bull. 25 (1971) 526; Kew Bull., Addit. Ser. 8 (1980) 168; S.E.C.Sierra et al., Blumea 52 (2007) 78, map 7. — Type: BW (Schram) 9349 (holo L), New Guinea, Bakaro, Mankowari.

Shrubs, 2–5 m high, dioecious. Indumentum composed of simple and stellately-tufted hairs, glandular hairs absent. Stipules narrowly triangular, 3–7 by 0.7–1.5 mm, early caducous, margin entire, apex acute, densely hairy outside, sparsely hairy inside. Leaves opposite; petiole 10–84 by 1.8–2 mm, densely hairy; blade ovate to elliptic, 15–31 by 5.5–15 cm, length/width ratio 1.7–2.6, chartaceous, base shallowly emarginate, rounded, obtuse to cuneate, margin entire, undulate, without glandular teeth, apex acute to caudate, upper surface glabrous, extrafloral nectaries along the midrib, sometimes also all over the blade, elliptic to orbicular, 0.6–0.8 by 0.6–0.8 mm, lower surface scatteredly hairy, domatia present, venation inconspicuously triplinerved, nerves 6–10 per side, indistinctly looping and closed near margin or ending in margin. Inflorescences single, erect; axes densely hairy; bracts linear-triangular, 2.9–3.1 by 0.5–0.7 mm, persistent, margin entire, apex acute, glabrous; bracteoles triangular 0.9–1.1 by 0.8–1 mm, caducous, margin entire, apex acute. Flowers: pedicels densely hairy; sepals free, persistent, margin entire, apex acute. Staminate inflorescences racemes, 4.5–6 cm long; peduncle 6–19 mm long; nodes per branch 5–7, with 8–11 flowers per bract. Staminate flowers 5–5.5 mm diam.; pedicels 3–4.5 mm long; sepals 3, ovate to elliptic, 3–5.5 by 2–2.5 mm; stamens 55–65, glabrous, filaments 2.5–4 mm long, free, thecae ovoid, 0.2–0.3 by 0.2–0.25 mm, connective widened; pistillode present. Pistillate inflorescences umbel-like, 12–17 cm long, nodes per branch 1, with 2 or 3 flowers per bract. Pistillate flowers 3–3.5 mm diam.; pedicels 6–7 mm long; sepals 3, triangular, 2.9–3.1 by 1.4–1.6 mm, densely hairy outside, glabrous inside; ovary 2.8–3 by 2.8 mm, 3-locular, densely hairy, scatteredly covered with glandular hairs; style 0.6–0.8 mm long; stigmas c. 2 by 0.8 mm. Fruits capsules, 7.5–9 by 14–17 mm, opening septicidally-loculicidally, surface spiny, densely hairy, scatteredly covered with glandular hairs; spines 70–90, straight, 1–1.2 by c. 0.8 mm, densely hairy; wall 1 mm thick, sparsely hairy inside; column 5.5–7 by 4–6.8 mm. Seeds ± globose, 7–7.5 by 6–7 by 6–7 mm, surface smooth, shiny, dark brown.

    Distribution — Endemic to New Guinea.

    Habitat & Ecology —In primary or disturbed lowland forest; on limestone or sandy peaty clay soil. Altitude: sea level to 130 m. Flowering and fruiting throughout the year.

    Note — Plants have a strong smell when dried.

31. Mallotus minimifructus S.E.C. Sierra (sect. Rottleropsis)

    Mallotus minimifructus S.E.C.Sierra in S.E.C.Sierra et al., Blumea 52 (2007) 79, fig. 7, map 5. — Typus: S (Y.P. Ching) 62305 (holo L; iso CGE, K, KEP, SAN), Borneo, Sarawak, Ulu Sungai Kebhor.

Shrubs to small trees, 3–10 m high; bole up to 5 m high, dioecious or rarely monoecious, crown up to 5 m long. Outer bark smooth. Indumentum composed of simple and tufted hairs and yellow to orange glandular hairs. Stipules triangular to narrowly triangular, 4–10 by 1.4–1.6 mm, persistent to caducous, margin entire, apex acute to caudate, sparsely hairy outside, scatteredly hairy to glabrous inside. Leaves opposite, petiole 1–50 by 0.6–1 mm, densely hairy to scatteredly hairy; blade ovate to obovate (the smaller leaf obcordate), 14–22 by 5–10 cm, length/width ratio 1.8–3.3, membranaceous (to chartaceous), base shallowly emarginate, obtuse to cuneate, margin denticulate, dentate to crenate, with glandular teeth, apex caudate, upper surface densely to scatteredly hairy on midrib and nerves, extrafloral nectaries marginal in lower half, 2–7 per side, 1–2.5 mm from margin, elliptic to orbicular, 0.2–0.3 by 0.2–0.3 mm, lower surface sparsely to scatteredly hairy, sparsely covered with glandular hairs, domatia rarely present, venation triplinerved, nerves 6–9 per side, indistinctly looping and closed near margin or ending in margin. Inflorescences racemes (see note 1), single, erect, unisexual or rarely bisexual; axes densely to sparsely hairy; peduncle 9–24 by 0.8–1 mm; bracts triangular, persistent, margin entire, apex acute, sparsely hairy outside, glabrous inside; bracteoles sometimes present, triangular, 1–1.5 by 0.2–0.3 mm, early caducous. Flowers: pedicels densely to sparsely hairy; sepals 3 or 4, free, persistent, margin entire, apex acute, densely hairy and with glandular hairs outside, glabrous inside. Staminate inflorescences 6–10 cm long, nodes per branch 12–27, with 1–5 flowers per bract; bracts 1.2–1.4 by 0.5–0.6 mm. Staminate flowers 3.3–4 mm diam.; pedicels 1.7–3 mm long; sepals elliptic to obovate, 2.8–3.2 by 1–1.2 mm; disc-glands present; stamens 40–90, glabrous, filaments 1–2.5 mm long, free, thecae ovoid, 0.2–0.3 by 0.2–0.25 mm, connective widened; pistillode absent. Pistillate inflorescences 3.4–12 cm long, nodes per branch 10–20, with 1 flower per bract; bracts 1.5–1.7 by 0.6–0.7 mm; bracteoles absent. Pistillate flowers 2.8–3.8 mm diam.; pedicels 1–3 mm long; sepals narrowly triangular, 2–2.4 by 0.8–1 mm; staminodes rarely present; ovary 1.3–1.5 by 1.3–1.6 mm, 3-locular, sparsely (to densely, see note 2) hairy, sparsely covered with glandular hairs; style 0.2–0.6 mm long; stigmas 2.2–5.5 by 0.8–1.2 mm. Fruits capsules, 5–10 by 8–11 mm, green, opening septicidally-loculicidally, surface spiny, sparsely (to densely) hairy, sparsely covered with glandular hairs; spines 100–200 (rarely more), curved, 1.5–3.5 by 0.1–0.2 mm, sparsely hairy, red; wall 0.7–0.9 mm thick, glabrous inside; column 3.8–4 by 3.5–4 mm. Seeds ± globose, 4–5.1 by 3.5–5 by 3.5–4.5 mm, surface smooth, shiny, dark brown.

    Distribution — Borneo, Philippines, Sulawesi.

    Habitat & Ecology — Locally common in understorey of primary forest or in disturbed areas; on basalt, clay-loam, limestone, sandy soil or ultramafic soil. Altitude: sea level up to 800 m. Flowering and fruiting throughout the year.

    Uses — Roots are used against fever.

    Vernacular names — Borneo (Sarawak): Entak empuluh. Philippines: Tiolu gletin.

    Notes — 1. Angian 10494 has inflorescences with very short internodes at the apex.

2. SAN 42227 differs from the other material studied in having fruits that are densely

covered with spines and hairs resembling those of M. leucocalyx.

3. See note under M. dispar.

32. Mallotus miquelianus (Scheff.) Boerl. (ex sect. Hancea)

   Mallotus miquelianus (Scheff.) Boerl., Handl. Fl. Ned. Ind. 3(1) (1900) 290; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 200; Merr., J. Str. Roy. Asiatic Soc. (1921) 339; Enum. Philipp. Flow. Pl. 2 (1923) 434; S.Moore, J. Bot. 63 (1925) 103; Merr., Philipp. J. Sci. 29 (1926) 382; Univ. Calif. Publ. Bot. 15 (1929) 158; Pax & K.Hoffm. in Engl. & Harms, Natürl. Pflanzenfam. ed. 2 (1931) 117; Burkill, Dict. Econ. Prod. Malay Penins. 2 (1935) 1395; Airy Shaw, Kew Bull. 20 (1966) 40; Meijer, Bot. News Bull. (1967) 53; Airy Shaw, Kew Bull. 26 (1972) 295; Whitmore, Tree Fl. Malaya 2 (1973) 114; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 163; Kew Bull. 36 (1981) 326; Alph. Enum. Euphorb. Philipp. Isl. (1983) 36; Keßler & Sidiyasa, Tropenbos Series 7 (1994) 133; Slik, Priyono & Welzen, Gard. Bull. Singapore 52 (2000) 59, Fig. 33; Slik & Welzen, Blumea 46 (2001) 55, Fig. 22, Map 9; Slik, Priyono & Welzen in Slik, Tropenbos-Kalimantan ser. 4 (2001) 97, Fig. 5.36; Slik & Welzej in Slik, Tropenbos-Kalimantan ser. 4 (2001) 200, Fig. 7.22, Map 7.9; Welzen et al., Blumea 51 (2006) 368. — Rottlera miqueliana Scheff., Ann. Mus. Bot. Lugd.-Bat. 4 (1868-69) 124. — Lectotype (Slik & Welzen, Blumea 46, 2001): Korthals s.n. (L sheet nr. 904105-154) (L holo), Borneo, Sakoembang, regione fl. Doesson.

   Mallotus anisophyllus Hook.f., Fl. Brit. India 5 (1887) 436; Boerl., Handl. Fl. Ned. Ind. 3(1) (1900) 290; Merr. & Rolfe, Philipp. J. Sci 3 (1908) 107; Elmer, Leafl. Philipp. Bot. 4 (1911) 1286; Ridl., Fl. Malay Penins. 3 (1924) 293. — Type: Maingay 1413 (K holo; L iso), Malaccaa.

    Also check: Slik, Macaranga and Mallotus species of Borneo

Shrubs up to 3(–10) m tall, dbh up to 10 cm, dioecious; bole crooked, vegetative reproduction via root shoots. Outer bark smooth, scaly; sapwood hard, tough, durable. Indumentum sparse to dense, simple to tufted to stellate. Branches smooth to slightly fissured, lenticellate, hairy, not gland-dotted, nodes only slightly swollen. Stipules early caducous, ovate, 4–13 by 1.8–3 mm, margin entire to irregular, apex acute, hairy abaxially only, not gland-dotted. Leaves simple, opposite, unequal, one of each pair much smaller. Non-reduced leaves: petioles 2–7 mm long, basally 2–3 mm thick, hairy, not gland-dotted; blade elliptic to obovate, 7–32 by 2.5–13 cm, length-width ratio 2–4.7, chartaceous, base cuneate (cordate), oblique, margin wavy to dentate, marginal glands 10–23 per leaf side, apex cuspidate to aristate, upper surface smooth, glabrous (to hairy), not gland-dotted, basal macular glands 2–6 on veins, 1–6 mm from midrib, apical glands 2–13 per side between the veins, 4–10 mm from margin, lower surface smooth, hairy, especially on veins, gland-dotted or not, domatia absent or present, with hair tufts; venation pinnate, 11–18 secondary veins per side, ending in marginal glands especially apically. Reduced leaves (semi-)persistent, cordate, 8–55 by 8–48 mm, margin entire to wavy, marginal glands sometimes present, apex rounded, upper surface with (0–)2 (to several) basal macular glands on veins, rarely also a few apical glands on veins, glabrous to hairy, not gland-dotted, lower surface sometimes with domatia, hairy, gland-dotted. Inflorescences in axils of reduced leaves, unbranched, solitary. Flowers actinomorphic. Staminate inflorescences thyrse-like racemes, up to 13.5 cm long, basally c. 1 mm thick, hairy, not gland-dotted, nodes up to 18 per inflorescence, flowers up to 6 per node; bracts persistent, ovate, 1.5–3 by 0.9–1.7 mm, margin entire to irregular, apex acute, hairy abaxially only, gland-dotted or not; buds ovate (to globose), apex acute to acuminate, hairy, gland-dotted. Staminate flowers 2.5–6 mm in diameter, yellow-green to white-green; pedicels 2.5–5.5(–10) mm long, glabrous to hairy, gland-dotted or not; sepals (2–)3, free, ovate to elliptic, 1.8–5 by 1–1.8 mm, margin entire to irregular, apex acute, glabrous to hairy, gland-dotted; stamens up to 55; filaments 2.3–4.5 mm long, glabrous; anthers c. 0.3 by 0.5 mm, basifixed; thecae 2, parallel, extrorse, opening lengthwise, glabrous; connective widened. Pistillate inflorescences racemes to rarely thyrse-like racemes, up to 10 cm long, basally c. 2.3 mm thick, hairy, gland-dotted or not, nodes up to 13 per inflorescence, flowers 1(–3) per node; bracts persistent, ovate, 3–4.5 by 1–2 mm, margin entire to irregular, apex acute, hairy abaxially only, not gland-dotted. Pistillate flowers dirty green to yellow-white to purplish-brown; bracteoles sometimes present, ovate, c. 1.3 by 1 mm, margin entire to irregular, apex acuminate, hairy, not gland-dotted; pedicels 1–6 mm long, hairy, gland-dotted or not; sepals 3, free, ovate, 4–8 by 2–3.3 mm, margin entire to irregular, apex acute, hairy abaxially only, gland-dotted; ovary 3(–4)-locular, echinate, hairy, gland-dotted, spines 0.8–1.5 mm long, hairy, few long hairs apically, gland-dotted; ovules 1 per locule, axillary, apotropous, ascending; styles persistent, up to 1 mm long, hairy, not gland-dotted; stigmas up to 5(–9) mm long, densely covered with granulate papillae on the inside, hairy abaxially, not gland-dotted. Fruit an echinate, lobed capsule, 8–12 by 5–7 mm, loculicidal-septicidal, green to brown-red; carpel inside glabrous, not gland-dotted; column 3.3–4 by 4–4.6 mm. Seeds glossy, roughly globose, c. 4.5 by 4–4.5 by 4.5 mm; hilum elliptic to deltoid, 0.5–1.3 by 0.5 mm.

    Distribution — Peninsular Thailand, Peninsular Malaysia, Sumatra, Borneo and the Philippines.

    Habitat & Ecology — Locally common in primary and secondary forests (scrub), mostly found on disturbed sites but also in the forest understorey; on undulating terrain, ridges, marshy areas, near streamlets, forest edges, and roadsides; on alluvial (swampy) to well drained terrain; on a large variety of soil types, from limestone to sandy soils to clayey loam. Altitude: up to 700 m.

    Uses — Used for making walking sticks. Poison.

    Notes — Very hairy specimens can be found in Sabah (and to a lesser extend also in Sarawak and the Philippines) as opposed to the nearly glabrous specimens from other localities.

33. Mallotus mollissimus (Geiseler) Airy Shaw (sect. Mallotus)

    Mallotus mollissimus (Geiseler) Airy Shaw, Kew Bull. 26 (1972) 297; Backer & Bakh.f., Fl. Java 1 (1964) 482; Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 165; Kew Bull., Addit. Ser. 8 (1980) 168; Kew Bull. 36 (1981) 326; Kew Bull. 37 (1982) 28; Alphab. Enum. Euph. Philipp. Isl. (1983) 37; P.I.Forst., Austrobaileya 5 (1999) 473, 494; Keßler, Blumea, Suppl. 14 (2002) 49; Welzen, Slik & Bollendorff in Welzen et al., Thai Forest bull., Bot. 28 (2000) 102; Slik, Priyono & Welzen, Gard. Bull. Singapore 52 (2000) 60, Fig. 34; in Slik, Tropenbos-Kalimantan ser. 4 (2001) 97, Fig. 5.37; S.E.C.Sierra & Welzen, Blumea 50 (2005) 257, fig. 3, map 3. — Croton mollissimus Geiseler, Croton. Monogr. (March 1807) 73. — Chrozophora mollissima (Geiseler) A.Juss. ex Spreng., Syst. Veg. 2 (1826) 851. — Echinus mollissimus (Geiseler) Baill., Adansonia 6 (1866) 316. —Type: Herb. Vahl s.n. (holo C; photo at A), China, see note 1.

Croton ricinoides Pers., Synop. Pl. 2, 2 (Sept. 1807) 586. — Rottlera ricinoides (Pers.) A.Juss., Euphorb. Gen. (1824) 111, t. 9, f. 29A. — Mallotus ricinoides (Pers.) Müll.Arg., Linnaea 34 (1865) 187, nom. superfl. — Lectotype (Forster, 1999): Lahaye s.n. (holo P, Herb. A. Juss. 16578, [IDC microfiche no. 6206]), India, see note 2.

    Adisca zippelii Blume, Bijdr. (1826) 611. — Adisa zippelii (Blume) Steud., Nom. Bot. ed. 2, 2 (1840) 28, sphalm. — Rottlera zippelii (Blume) Hassk., Cat. (1844) 238. — Mappa zippelii (Blume) Zoll. & Moritzi ex Moritzi, Syst. Verz. (1846) 17. — Mallotus zippelii (Blume) F.Muell., Fragm. Phytogr. Austr. 4 (1864) 139. — Lectotype (Sierra & Welzen, 2005): Blume s.n. (holo L barcode 0294214; iso L barcode 0294172), Java, Mt Salak.

    Adelia bernardia auct. non L.: Blanco, Fl. Filip. (1837) 814; Merr., Sp. Blanc. (1918) 221. — Adelia barbata Blanco, Fl. Filip. ed. 2 (1845) 561. — Neotype (Sierra & Welzen, 2005): Merrill Species Blancoanae 474 (holo US; iso K, L), Philippines, Luzon, Rizal.

    Mallotus pycnostachys F.Muell., Fragm. Phytogr. Austr. 4 (1864) 138. — Lectotype (Forster, 1999): MEL 232430, 32-34, Australia, Queensland, North Kennedy Distr., Mt Elliott.

    Mallotus barbatus auct. non Wall. ex Müll.Arg.: Merr., Philipp. J. Sci., Bot. 7 (1912) 397. — Mallotus confusus Merr., Philipp. J. Sci. 16 (1920) 559. — Type: BS (Ramos & Edaño) 29153 (holo PNH†; iso A, P, US), Philippines, Luzon, Tabayas Province, Mt Tulaog.

    [Mappa sp.: Moritzi, Syst. Verz. (1846) 17].— Mappa zippelii (Blume) Zoll. & Moritzi var. minor Rchb.f. & Zoll., Linnaea 28 (1857) 315, nom. nud. — Rottlera peltata auct. non Roxb.: Miq., Fl. Ned. Ind. 1, 2 (1859) 395. — Voucher: Zollinger 319 (G, L), Indonesia, Java.

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

Shrubs to small trees up to 12 m high, dbh up to 15 cm, dioecious, occasionally monoecious; bole up to 5 m high; crown up to 6 m long. Outer bark rough to finely fissured or pustular lenticellate, up to 2 mm thick, yellowish green, under surface light reddish

brown; inner bark up to 6 mm thick, fibrous, light greenish; sapwood cream with watery sap; wood medium hard, cream. Indumentum dense, sometimes soft-floccose, flocci up to 4 mm long, rarely tomentose. Stipules narrowly triangular, 0.7–1.5 by 0.1–0.3 mm, caducous to persistent, margin subentire, apex acuminate. Leaves alternate to apically subopposite; petiole 30–200 by 1.5–4 mm; blade subpeltate or peltate for 1.5–40 mm, ovate, 10–35 by 8–30 cm, length/width ratio 1.2–1.4, reddish brown when young, base truncate to cuneate, margin entire to dentate, seldom 2-lobed at the widest part of the blade, longest lobes up to 15 mm long, upper surface green, basally with 0–2(–4) extrafloral nectaries, 0.8–4 by 0.4–1.8 mm, marginal nectaries 0–15 per side, 2–10 mm from margin,0.5–1 by 0.5–1 mm, lower surface greenish grey to brownish, with domatia or not; 3- or palminerved, nerves 8–11 per side, mostly ending in the margin. Inflorescences axillary or terminal, unisexual or bisexual, axes greenish tawny, basally 1–4 mm thick; bracts narrowly triangular, 1.8–3 by 0.2–0.5 mm, caducous to persistent, margin subentire; bracteoles absent. Staminate inflorescences up to 43 cm long, side branches up to 20 cm long, with 3–5 flowers per node, nodes per branch up to 100. Staminate flowers 3–5 mm diam.; pedicels 3–3.5 mm long; sepals 3 or 4, elliptic, 2–3 by 1.5–2 mm, greyish; stamens 50–80,yellowish, filaments 1.5–3 mm long, anthers 0.2–0.3 by 0.1–0.2 mm; pistillode present. Pistillate inflorescences racemes, up to 35cm long, occasionally side branches up to 13 cm long, nodes up to 130; bracts caducous to persistent. Pistillate flowers 3–4 mm diam.; pedicels up to 0.5 mm long; calyx 3–5-lobed, connate on the base, 2–3 mm long, green with brown indumentum, lobes ovate, 1.5–2 by 1–1.7 mm; ovary 3-locular, 2.2–3 by 2–3 mm, spines individually visible; style up to 1.2 mm long; stigmas 2.2–2.8 mm long, greenish yellow; staminodes absent. Fruits 10–16 by 11–17 mm, greenish brown, smelling strong when dried, see note 3, spines numerous, curly, thin, less hairy, up to 7 mm long, hairs not forming a continuous layer, easily rubbing off; column 5–6 by 1–1.8 mm. Seeds ellipsoid, 4–5 by 3.8–4 by 3–3.2 mm, smooth, black, shiny; hilum c. 1 by 0.6–0.7 mm.

    Distribution — From Sumatra to East Australia and West Pacific (Solomon Islands).

    Habitat & Ecology — Locally common in deciduous and evergreen forest, mostly in open, often very disturbed or burned places, gallery forest, mangrove edges, swamp forest, riparian vegetation, notophyll vineforest, on mountain slopes, open grasslands and along hill and road sides; on a large variety of soil types, like alluvial, basalt rocks, clay, coral limestone, loam, rocky, sandy, ultrabasic, and volcanic. Altitude: sea level up to 1700 m. Flowering and fruiting the whole year through. Observed to be visited by small brown ants.

    Uses — The roots are used against headaches and malaria. The inner bark is used as a strap for carrying and for sing sing decorations. The leaves are used as food covering, against stomach cramp, and together with the bark used to cure the spleen. The wood is used for making scabbards, ritual spears, building construction, and as firewood.

    Vernacular names — Sumatra: Kajoe si balihangin. Java: Djalikangin, Toetoep beling. Sabah: Dapulan, Dullong perampuang (Bajau); Patau jantan (Bajec); Kasobong (Daho); Andalamit, Angkutangkut, Dahu, Dau, Kedayan, Lukabang, Padau-padau, Padahu, Pandau pandu (Dusun); Laba-laba (Kinabatangan); Bajau papar, Murut tenom (Kwijau); Ambutan, Dawoh, Lapakon (Murut); Dapulon-dapulon, Pais tahun (Suluk); Apat retu, Apatmaratu, Pit dau (Sungei); Andalasan, Kehat, Labah-labah, Labah lahat, Ludai, Padauk, Porak sembelik, Purak sambalik. Sarawak: Labah-labah (Kedayan); Lalapakon (Murut). Brunei: Jabai, Laba-laba, Markubong, Prestaang, Tokatoka, Tuka, Sawarbubu. Philippines: Hanonomo (Bisáya); Indang (Lánao); Bampalan (Súlu); Hinlaumo (Tagálog); Ipalan (Yakán); Aab-ben, Anumoh, Anumútagabas, Atay-atay, Baray, Cahoy-dalaga, Gapasgapas, Malang-malang, Maracapau, Tagaan. Sulawesi: Kayou wollo sola, Laransiana oeroene, Lemoro, Ota-ota. Papua (Irian Jaya): Bekoeaap, Bekwaap, Bie grai, Majetrip (Hattam); Mangafafin (Biak). Papua New Guinea: Atatiza (Garaina, Waria); Majetrip (Hattam); Wapi (Marok); Siripupo (Orakiva); Wasittatt (Wein); Ah papper, Goreh-gobu, Hukoitiiti, Hukopu, Kikikanauant, La putu la basi, Moraka, Ongohn, Tiwotiwo, Ukoiitiiti.

    Notes — 1. Forster (1999) states that Müller Argoviensis (1866; 964 sub. M. ricinoides) mentioned a Staunton collection from China as the type of M. mollissimus. But actually Müller Argoviensis indicated two specimens from China: a Staunton collection in herbarium G and a Vahl collection in herbarium C. Part of the confusion has arisen, because, when reading Müller Argoviensis, one can think that there is one single collection (Staunton s.n.) with the holotype in G and the isotype in C. However, Müller Argoviensis added ‘i.e.’, meaning a single form! (id est), this is the original specimen to the C sheet.

Moreover the Staunton collection cannot be the type of M. mollisimus, because there is no evidence that Geiseler ever saw it. It was given by Lambert to De Candolle in 1816, as was written on the label (for additional information see also Miller, Taxon 19, 1970: 543).

Since the Crotonis Monographiam from Geiseler (1807) is based on material from Vahl at the Copenhagen Herbarium, by circumstantial evidence, the most likely specimen to be the type is the Vahl specimen found in C as was actually said by Müller Argoviensis. Like Croizat (1938: 141) we doubt the Chinese origin of this specimen as no other one has come to our attention. The nearest occurrence could be Tu Phap in North Vietnam, but the provenances given by the collector d’Aleizette cannot always be trusted. The specimens cited by Gagnepain (in Lecomte, Fl. Indo-Chine 4, 1925: 357) and Poilane 1685 cited by Croizat as well, as far as seen belong to M. metcalfianus.

A letter by Croizat on July 14, 1938, that is after the publication of his paper (April 27, 1938) suggests a possible Indonesian origin and this seems a good suggestion.

2. Croton ricinoides was described from India. The lectotype was collected by Lahaye, who never was in India (cf. Van Steenis-Kruseman, Fl. Males. ser. 1, 1, 1950: 307) but did collect in Malesia.

34. Mallotus monanthos Airy Shaw (sect. Rottleropsis)

    Mallotus monanthos Airy Shaw, Kew Bull. 32 (1978) 402; S.E.C.Sierra et al., Blumea 52 (2007) 83, map 1. — Type: KEP FRI (Whitmore) 3544 (holo K; iso A, L), Peninsular Malaysia, Pahang, Krau Game Reserve, Kuala Lumpat.

Small trees, 2.4–5 m high, monoecious (probably also dioecious, see note). Indumentum composed of simple and tufted hairs, glandular hairs absent. Stipules linear-triangular, 9–11 by 1.5–2 mm, caducous, margin entire, apex acute, scatteredly hairy outside, densely hairy inside. Leaves opposite; petiole 5–15 by 2.5–4 mm, densely hairy; blade narrowly obovate, 25–42 by 7–11.5 cm, length/width ratio 3.4–4, coriaceous, base attenuate, margin entire, undulate, apex caudate, upper surface glabrous, larger extrafloral nectaries marginal, 2 or 3 per side, 0.5–2.5 mm from margin, elliptic to orbicular, 0.8–1 by 0.7–0.8 mm, smaller ones from base to apex, 9–13 per side, 4–7 mm from margin, orbicular, 0.2–0.3 by 0.2–0.3 mm, lower surface glabrous, domatia absent, venation pinnate, nerves 15–18 per side, looping and closed near margin. Inflorescences erect, several together; peduncle absent; bracts linear-triangular, persistent, margin entire, apex acute, hairy, sparsely hairy to glabrous outside, sparsely hairy inside; bracteoles absent. Flowers: pedicels densely hairy; sepals free, persistent, margin entire, apex acute, densely hairy outside, glabrous inside. Staminate inflorescences (only bracts and flowers seen): bracts numerous, c. 40, 3–4 by 0.4–0.5 mm. Staminate flowers 4–5 mm diam.; pedicels 1–1.7 mm long; sepals 3, ovate, 5–8.5 by 1.5–3 mm; stamens 20–23, glabrous, filaments 2–5 mm long, basally connate, thecae ellipsoid, 0.6–0.7 by 0.2–0.25 mm, connective widened; pistillode present. Infructescences reduced to 1 terminal fruit, 1 cm long; peduncle reduced; bracts c. 15, 8–10 by 1.3–1.5 mm. Pistillate flowers (based on young fruit): pedicels c. 3 mm long; sepals 3, narrowly triangular, c. 7 by 1.8 mm; ovary 3-locular, densely hairy; style c. 3 mm long; stigmas c. 8 by 1 mm. Fruits capsules, c. 17 by 20 mm, opening septicidally-loculicidally, surface smooth, densely hairy; wall c. 1 mm thick, glabrous inside; column c. 4 by 2 mm. Seeds ± globose, c. 9 by 9.5 by 9.3 mm, surface smooth, dull, light cream brown.

    Distribution — Endemic to Peninsular Malaysia.

    Habitat & Ecology — Dense lowland forest. Altitude 365 m. Flowering and fruiting: March.

    Note — The two collections studied (KEP FRI 3544 and Soepadmo 736) have la-

bels that only refer to the fruits. Soepadmo 736 (A) consists of a loose fruit. KEP FRI 3544 (A, K) also consists of fruits, while the L specimen has two branches, one with

female (longer and few) and the other with male bracts (shorter and numerous), with

the staminate flowers stored in an envelope. Based on KEP FRI 3544 (L) it is likely

that the species is monoecious (if the branches belong to the same plant), while based

on Soepadmo 736 (A) it may be dioecious; it is therefore evident that further material

needs to be studied.

35. Mallotus montanus (Wall. ex Müll.Arg.) Airy Shaw (sect. Rottleropsis)

    Mallotus montanus (Wall. ex Müll.Arg.) Airy Shaw, Kew Bull. 32 (1977) 78; Welzen et al., Thai Forest Bull., Bot. 28 (2000) 102; S.E.C.Sierra et al., Blumea 52 (2007) 83, map 8. — [Croton montanus Wall., Numer. List (1845) 7723B, nom. nud., non Willd.] — Coelodiscus montanus Wall. ex Müll.Arg. in DC., Prodr. 15, 2 (1866) 759. — Lectotype (S.E.C.Sierra et al., 2007): Wallich 7723B (holo G-DC (IDC microfiche 800); iso BM, E, K-W (IDC microfiche 7394), L), Peninsular Malaysia, Penang.

    Mallotus eriocarpus auct. non Müll.Arg.: Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii 160; Whitmore, Tree Fl. Malaya 2 (1973) 115; quoad specimen from Peninsular Malaysia.

Shrubs to small trees, up to 6 m high, dioecious or rarely monoecious. Indumentum composed of simple, tufted, stellately-tufted and stellate hairs (on petiole and midrib of blade) and light yellow-orange glandular hairs. Stipules deltoid to triangular, 0.6–1.3 by 0.5–0.6 mm, early caducous, margin entire, apex rounded, densely hairy. Leaves opposite; petiole 1–7 by 0.5–2 mm, densely hairy; blade ovate to elliptic, 5–24 by 3–12 cm, length/width ratio 1.6–2.5, chartaceous, base cordate to obtuse, margin denticulate, with glandular teeth, apex acute to caudate, upper surface sparsely hairy (densely hairy on midrib and nerves), sometimes with glandular hairs, extrafloral nectaries marginal from base to apex 7–17 per side, 0.4–2 mm from margin, elliptic to orbicular, 0.2–0.4 by 0.2–0.4 mm, lower surface sparsely hairy, sparsely covered with glandular hairs, domatia usually present (towards the petiole insertion), venation triplinerved, nerves 4–7 per side, looping and closed near margin or ending in margin. Inflorescences racemes, single, erect, unisexual or rarely bisexual; peduncle 7–35 by c. 1 mm, densely hairy; bracts margin entire, apex acuminate, densely hairy; bracteoles absent. Flowers: pedicels densely hairy; sepals free, persistent, margin entire, apex acute, densely hairy outside, scatteredly hairy to glabrous inside, with glandular hairs outside. Staminate inflorescences 2.5–4 cm long, nodes per branch 5–9, with 2–4 flowers per bract. Staminate flowers 3–3.5 mm diam.; pedicels 1.2–1.5 mm long; sepals 3 (or 4), ovate, 1.5–2 by 2–2.5 mm; stamens 20–30, filaments 2–3 mm long, free, densely hairy, thecae ovoid to ellipsoid, 0.2–0.3 by 0.2 mm, hairy to glabrous, connective not widened; pistillode present. Pistillate inflorescences 3–7 cm long, nodes per branch 1–6, with 1 flower per bract. Pistillate flowers 4.5–5.5 mm diam.; pedicels 0.8–1 mm long; sepals 2 or 3, triangular, 2.5–3 by 1.3–1.5 mm; ovary 1–1.2 by 1.2–1.5 mm, 3-locular, densely hairy, sparsely covered with glandular hairs; style 0.2–0.4 mm long; stigmas 1–2 by 0.6–0.8 mm. Fruits capsules, 6–13 by 15–19 mm, opening septicidally-loculicidally, surface smooth, densely hairy, sparsely covered with glandular hairs; wall c. 0.5 mm thick, sparsely hairy inside; column 4–10 by 4.5–6 mm. Seeds ± globose, 4.2–6.5 by 4.2–6.5 by 4.2–6.2 mm, surface smooth, shiny, light brown.

    Distribution — Thailand, Peninsular Malaysia.

    Habitat & Ecology — Locally common in primary, secondary and evergreen forest; along edges of rubber plantations; on calcareous rock. Altitude 100–400 m. Flowering and fruiting: March to November.

    Note — See note under M. subcuneatus.

36. Mallotus muticus (Müll.Arg.) Airy Shaw (sect. Polyadenii)

    Mallotus muticus (Müll.Arg.) Airy Shaw, Kew Bull. 16 (1963) 351; Kew Bull. 20 (1966) 39; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 53; Whitmore, Tree Fl. Malaya 2 (1973) 116, 117, fig. 10; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 167; Kew Bull. 36 (1981) 327; Bollendorff, Welzen & Slik, Blumea 45 (2000) 329, fig. 2b, 4; map 3; Slik, Priyono & Welzen, Gard. Bull. Singapore 52 (2000) 61, Fig. 36; in Slik, Tropenbos-Kalimantan ser. 4 (2001) 98, Fig. 5.39; Bollendorff, Welzen & Slik in Slik, Tropenbos-Kalimantan ser. 4 (2001) 132, Fig. 6.2b, 6.6.; Map 6.3. — Coccoceras muticum Müll.Arg., Flora 47 (1864) 470; in DC., Prodr. 15, 2 (1866) 950; Hook.f., Fl. Brit. India 5 (1887) 424; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 210; Ridl., Fl. Malay Penins. 3 (1924) 294. — Coccoceras muticum Müll.Arg. var. muticum: Müll.Arg., Flora 47 (1864) 470; in DC., Prodr. 15, 2 (1866) 950; Hook. f., Fl. Brit. India 5 (1887) 424; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 210; Ridl., Fl. Malay Penins. 3 (1924) 294. — Type: Griffith KD 4770 (K), Malacca.

    Mallotus borneensis Müll.Arg. in DC., Prodr. 15, 2 (1866) 980; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 198. — Rottlera borneensis (Müll.Arg.) Scheff., Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 125. — Lectotype (Bollendorff et al., 2000): Korthals s.n., L. sheet no. 905.105-21 (L, holo), Indonesia, Borneo.

    Coccoceras borneense J.J.Sm., Bull. Jard. Bot. Buitenz., ser. 3, 6 (1924) 94 (pro spec. nov.). — Lectotype (Bollendorff et al., 2000): Teysmann s.n., L. sheet no. 914.150-496 (L, holo), Indonesia, Borneo, Soengai Landak.

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

Tree up to 35 m high, dioecious, mainly glabrous; branchlets finely ribbed, not to rarely lenticellate, glabrous, with white to greyish bark with conspicuous discoid glands. Stipules early caducous. Leaves subopposite to alternate, pairs unequal in size; petiole terete to angular, adaxially channelled, 18–83 by 0.7–3 mm, glabrous to sometimes puberulous at ends, base and apex usually not pulvinate; blade elliptic to obovate, 8.2–24.8 by 3.7–14 cm, index 1.3–2.4, chartaceous to coriaceous, base rounded to obtuse and often minutely auriculate, margin entire to slightly irregularly crenate, 6–16 marginal glands per leaf side, usually in sinuses, triangular to globose, thickened, c. 0.7 by 0.3 mm, protruding especially in young leaves, apex acuminate to obtusely cuspidate, sometimes ending in a margin-type of gland, upper surface green to grey to brown, 0–3(–8) basal glands on each side, elliptic, impressed, c. 1.5 by 0.8 mm, grey or brown, 0–8 mm from petiole attachment, apically none to 5 circular, impressed glands on last pair of nerves, lower surface pale green to brown, domatia with villose to long stellate white hairs in nerve axils; venation basally triplinerved, basal nerves ending in margin below and above middle, further pinnate, with 3–5(–7) nerves per side, eucamptodromous, veins conspicuously scalariform. Staminate inflorescences and flowers as those of Mallotus leucodermis, see there. Pistillate inflorescences axillary to rarely terminal, solitary or 2 or 3 together, rachis angular, 3.5–23.5 cm by 0.5–2 mm, 10–35 nodes per rachis, 1 flower per node; bracts 1–1.5 by c. 1 mm, margin glabrous to sometimes ciliate, apex acuminate to cuspidate, patent to reflexed; pedicels terete and often angular near abscission zone, 1–7(–12) by 0.5–1 mm, often geniculate, abscission zone 1–2 mm from apex; flowers 3–4 mm in diam., calyx lobes (4 or) 5, connate to up 0.5 mm from base, triangular to narrowly triangular, 2–2.5 by 0.5–1.2 mm, long white villose hairs on the inside, reddish brown, margin sometimes villosely ciliate, recurved when old, apically acuminate to cuspidate, basally thickened, ovary ovate in lateral view, 2- or 3-locular, covered with discoid glands, style 1.5–2 by c. 0.5 mm, stigmas 2–2.5 by 0.5–1 mm, basally erect, apically recurving. Fruits indehiscent, woody capsules, broadly angular-ovate to obcordate in lateral view, 12–25 by 11–22 mm in diam., apically style remnant often bent; locules 0.7–1 cm wide, slightly to clearly ridged over valve joints; column narrowly rhomboid with basal part slightly longer than apical one, c. 15 mm long by 6 mm wide. Seeds ovoid, 6–7 by 4–5 mm in diam., shiny to dull, smooth, brown.

    Distribution — W Malesia: Peninsular Malaysia, Sumatra, and Borneo.

    Habitat & Ecology — Primary and secondary forest, in periodically inundated forest, forest edges, logged plains, often in riparian forests, and swampy areas. Alt.: sea level to 500 m. Flowering: Mar.–Oct.; fruiting all year round.

    Vernacular names — Sumatra: Senggawan. Borneo: Kalimantan: Kelepuek, perupuk (Modang Dayak); Sabah: Randang jangun, salungapit or suko lapit (Kedayan); salung api, sandanaap; tapai longan (Murut).

    Note — Whitmore's (1973) fruit description does not correspond with the fruits of M. muticus. He described the fruits as spiny, which they are not. He correctly described them as indehiscent, however, the corresponding figure shows a split-open capsule.

37. Mallotus nudiflorus (L.) Kulju & Welzen (sect. Rottleropsis)

    Mallotus nudiflorus (L.) Kulju & Welzen in Kulju, S.E.C.Sierra & Welzen, Blumea 52 (2007) 124. — Trewia (‘Trevia’) nudiflora L., Sp. Pl. (1753) 1193; Willd., Sp. Pl. ed 4, 2  (1806) 834; Sm. in Rees, Cyclop. 36 (1819) 8; Wight, Icon. Pl. Ind. Or. (1852) 21, t. 1870, 1871; Baill., Étude Euph. (1858) 409; Miq., Fl. Ned. Ind. 1, 2 (1859) 400; Müll.Arg. in DC., Prodr. 15, 2 (1866) 953; Bedd., Fl. Sylv. 2 (1872) 281; Kurz, Forest Fl. 2 (1877) 379; Benth. in Benth. & Hook.f., Gen. Pl. 3, 1 (1880) 318; Hook.f., Fl. Br. Ind. 5 (1887) 423; Pax in Engl. & Prantl, Nat. Pflanzenfam. 3, 5 (1896) 53; Brandis, Ind. Trees (1906) 590; T.Cooke, Fl. Presid. Bombay 2, 3 (1906) 614; J.J. Sm., Meded. Dept. Landb. Ned.-Indië 10 (1910) 390; Craib, Bull. Misc. Inform. Kew (1911) 466; Talbot, Forest Fl. Bombay Pres. Sind 2 (1911) 481; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 140; Troup, Silv. Ind. Trees 3 (1921) 841; Merr. , Enum. Philipp. Fl. Pl. 2 (1923) 431; Gamble, Fl. Presid. Madras 2, 7 (1925) 1319; Gagnep. & Beille in Lecomte, Fl. Indo-Chine 5 (1925) 343; Backer & Bakh.f., Fl. Java 1 (1964) 481 (‘Trevia’); Airy Shaw, Kew Bull. 20 (1966) 405; Kew Bull. 23 (1969) 79; Kew Bull. 26 (1972) 343; Whitmore, Tree Fl. Malaya 2 (1973) 134; Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 200; Kew Bull. 36 (1981) 350; Alphab. Enum. Euphorb. Philipp. Isl. (1983) 46; Grierson & D.G.Long, FL. Bhutan 1, 3 (1987) 799; Nicolson et al., Regnum Veg. 119 (1988) 119; H.S.Kiu et al., Fl. Reip. Pop. Sin. 44, 2 (1996) 11; Vu Van Dung, Vietnam For. Trees (1996) 249; Philcox in Dassan. & W.D.Clayton, Rev. Handb. Fl. Ceylon 11 (1997) 143; Susila & N.P.Balakr., J. Econ. Taxon Bot. 22 (1998) 345, p.p.; S. Gardner et al., Field Guide For. Trees N. Thailand (2000) 322; Govaerts et al., World Checkl. Bibliogr. Euphorb. 3 (2000) 1560. — Trewia integerrima Stokes, Bot. Mat. Med. 4 (1812) 570 (‘Treuia’), nom. superfl. — Trewia nudiflora var. nudiflora: Susila & N.P.Balakr., J. Econ. Taxon Bot. 22 (1998) 347, t. 1. — Type: Rheede, Hort. Malab. 1: 76, t. 42 (‘Canschi’) 1678.

    Trewia macrophylla Roth, Nov. Pl. Spec. Ind. Or. (1821) 373 (non Blume, 1825). — Type: Heyne s.n. (B†), India orientalis.

    Trewia macrostachya Klotzsch in Klotzsch & Garcke, Bot. Ergebn. Reise Königl. Hoh. Priz. Waldemar Preussen (1862) 117, t. 23. — Type: Hoffmeister s.n. (n.v., B, †), Himalaya.

    Mallotus cardiophyllus Merr., Philipp. J. Sci., Bot. 7 (1912) 398. — Type: Cuming 1267 (holo PNH, †; iso A, BM, CGE, K, L, W), Philippines, Luzon, Cagayan Province.

Trewia nudiflora L. var. tomentosa Susila & N.P.Balakr., J. Econ. Taxon Bot. 22 (1998) 351, t. 3. — Type: G.V. Subba Rao 62480A/B (holo MH n.v.; iso MH n.v.), India, Andra Pradesh, Srikakulam Distr., near Regadi.

    Rottlera indica auct. non Willd.: Willd., Gött. J. Natürwiss. 1 (1797) 8, quoad. descr., voucher Klein s.n. (B-Willd, microfiche IDC #7440 index nr. 18504), India orientalis.

(Large shrubs to) trees up to 30 m high, dbh up to 50(–80) cm, dioecious, deciduous, flowering before or at the time of leaf appearance; branches ribbed when dry, young ones glabrous to hairy, sometimes with glandular hairs, older ones glabrescent; flowering branches 1.6–8.5 mm thick. Outer bark thin, smooth (to slightly rough), grey to brown (to tan), sometimes patchy. Indumentum of simple to tufted hairs, short to long and soft, whitish to yellowish to brownish, density in most parts varying greatly; glandular hairs yellowish, often (almost) absent. Stipules (narrowly) triangular, 1.8–5.2 by 0.5–1.5 mm, early caducous, margin entire, apex acute, (subglabrous to) hairy outside, (sub)glabrous inside. Leaves opposite to subopposite; petiole 1.8–12 cm long, often with a basal constriction when dry, glabrous to hairy, sometimes with glandular hairs; blade ovate, 6–21 by 6.3–16.5 cm, length/width ratio 1–2.3(–4.8 in immature leaves), papery, base cordate to obtuse (to acute to cuneate especially in immature leaves), not peltate to subpeltate, very base often attenuate, margin subentire (to rarely basally serrate with 1–7 minute teeth per side), apex (acute to) acuminate; upper surface glabrous to sparsely hairy (to densely hairy at venation), dark green, usually granular dotted, basal extrafloral nectaries 2–5, 1–16 mm from petiole insertion, marginal extrafloral nectaries 0–8(–20) per side, 3–26 mm from margin; lower surface glabrous to densely hairy, lighter green, glandular hairs absent to sparse, hair-tuft domatia sometimes present; venation triplinerved (to 5-nerved), nerves 3–7 per side, looped and closed near the margin. Inflorescences axillary racemes, pendulous when staminate, single, or often 2 or 3 together when staminate, axis subglabrous to hairy; staminate flowers many, (2 or) 3 (or 4) per node, pistillate flowers 1–5 at apical part, single per node; bracts early caducous to somewhat persistent when staminate, triangular, staminate ones cymbiform to recurved, margin entire to ciliate, apex acute, hairy to apically glabrous outside, (sub)glabrous inside. Staminate inflorescences up to 30 cm long, basally 0.6–2.1 mm thick; bracts 1.5–3.9 by 0.8–1.7 mm. Staminate flowers: buds ovoid to ellipsoid, hairy-apiculate; flowers 4.7–9 mm diam.; pedicels 3.9–10 mm long, hairy, with widened and more hairy abscission zone at basal part; sepals (2 or) 3 or 4, free, (ovate to) elliptic, (3.2–)3.7–6 by (1.8–)2.1–3.7 mm, often recurved, apex acute to acuminate, subglabrous to hairy outside, sometimes with scattered glandular hairs, (sub)glabrous inside, light yellowish to light greenish; receptacle slightly swollen; disc-glands absent; stamens (25–)45–75(–130), filaments 1.1–6 mm long, free, glabrous, whitish to light greenish, thecae 0.8–1.7 by 0.3–0.6 by 0.2–0.4 mm, opposite to subalternate, opening extrorsely to latrorsely, light yellowish to light greenish, connective widened or not. Pistillate inflorescences 1.5–10.5 cm long; bracts 1.2–3.2 by 0.5–2.1 mm. Pistillate flowers: pedicel 1.1–9 mm long, usually more hairy than inflorescence axis; calyx 4.7–6.4 mm long, connate with 3–5 lobes, opening further with 1 or 2 sutures, soon caducous, hairy outside, (sub)glabrous to sparsely hairy inside; ovary ovoid to ellipsoid, densely shortly hairy, glandular hairs present to absent, locules 3–5; style (1.7–)2.4–4.5(–5.9) mm long, hairy, glandular hairs (present to) absent; stigmas 12–24 mm long, plumose inside, hairy outside. Fruits indehiscent, drupaceous, oblate (to spheroid), 18–29 by 21–35 mm when dry, up to 35 by 45 mm when fresh, surface rugose when dry, subglabrous to hairy, light green to light brown to greyish, speckled; pericarp 2–8 mm thick when dry, mesocarp somewhat hard and fleshy, endocarp crustaceous. Seeds in cross section somewhat triangular to almost elliptic, 8–12 by 7–10 by 6–8 mm, surface smooth, brown to black, with thin sarcotesta.

    Distribution — From India and Nepal throughout mainland Southeast Asia to southern China (Hainan, Yunnan) and West Malesia (Malay Peninsula, Sumatra, Borneo, Java, Philippines).

    Habitat & Ecology — Deciduous to evergreen forest, usually disturbed and (partly) open sites, usually near streams and rivers, often on alluvial, but also on dry ground; bedrock varying (granite to limestone to shale to quartzite). In lowland Nepal M. nudiflorus is a common tree in riverine forests and dispersed by rhinoceros and domestic cattle (Dinerstein & Wemmer, Ecology 29, 1988, 1768-1774). Altitude up to 1200 m. Flowering in mainland Asia: mainly from December to April; fruiting: May to September.

    Vernacular names —Java: Binung putjang, Gemblok. Borneo: Pridja buaja.

    Uses — Wood is used for implements; leaves for fodder (Nepal). Medicinal uses: to remove swelling and bile, to relieve flatulence, for gout and rheumatic afflictions (India, Nepal).

38. Mallotus paniculatus (Lam.) Müll.Arg. var. paniculatus (sect. Mallotus)

    Mallotus paniculatus (Lam.) Müll.Arg. var. paniculatus: S.E.C.Sierra & Welzen, Blumea 50 (2005) 263, fig. 4, map 4. — Mallotus paniculatus (Lam.) Müll.Arg., Linnaea 34 (1865) 189; Backer & Bakh.f., Fl. Java 1 (1964) 483; Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 166; Kew Bull., Addit. Ser. 8 (1980) 169; Kew Bull. 36 (1981) 327; Kew Bull. 37 (1982) 29; Alphab. Enum. Euph. Philipp. Isl. (1983) 37; Corner, Ways. Trees Malaysia, ed. 3, 1 (1988) 307; P.I. Forst., Austrobaileya 5 (1999) 478, 496; Keßler, Blumea, Suppl. 14 (2002) 50; Welzen, Slik & Bollendorff in Welzen et al., Thai Forest bull., Bot. 28 (2000) 103; Slik, Priyono & Welzen, Gard. Bull. Singapore 52 (2000) 61, Fig. 37; in Slik, Tropenbos-Kalimantan ser. 4 (2001) 98, Fig. 5.40; W.J.Kress et al., Checkl. Myanmar, rev. ed. 4 (2003) 232; S.E.C.Sierra & Welzen, Blumea 50 (2005) 261. — Croton paniculatus Lam., Encycl. 2 (1786) 207; A.Juss., Euphorb. Gen. (1824) 33, comb. in Rottlera not made!. — Rottlera paniculata (Lam.) Blume, Bijdr. (24 Jan. 1826) 604; Spreng., Syst. Veg. 2 (Jan.–Mar.1826) 877 (isonym). — Lectotype (Forster, 1999): Commerson s.n. (holo P, Herb. A. Juss. 16579 [IDC microfiche no. 6206]), Indonesia, Java.

    Echinus trisulcus Lour., Fl. Cochinch. 1 (1790) 633. — Mappa cochinchinensis Spreng., Syst. Veg. 2 (1826) 878, nom. superfl. — Lasipania tricuspis Raf., Sylv. Tell. (1838) 22, nom. superfl. — Type: Loureiro s.n. (holo BM? n.v.), Vietnam, Annam, Cây hón.

    Mallotus cochinchinensis Lour., Fl. Cochinch. 1 (1790) 635. — Trewia tricuspidata Willd., Sp. Pl. ed. 4, 2 (1806) 835, nom. superfl. —[Mallotus chinensis Lour. ex Müll.Arg. in DC., Prodr. 15, 2 (1866) 965, in syn. sphalm. pro cochinchinensis]. — Lectotype (Sierra & Welzen, 2005): Loureiro s.n. (BM [photo at BRI]), Vietnam, Annam.

    Rottlera alba Roxb. [Hort. Beng. (1814) 73, nom. nud.] ex Jack, Descr. Malay. Pl. 1 (1820) 26; Roxb., Fl. Ind. 3 (1832) 829, isonym. — Mallotus albus (Roxb. ex Jack) Müll.Arg., Linnaea 34 (1865) 188, quoad basionym et Wall. 7818. — Lectotype (Sierra & Welzen, 2005): William Roxburgh fil. s.n. ≡ Wallich Numer. List 7818A (Hb. Roxb.!) (holo BR sh 849430 in fr., curiously the label says male, P, fide leaf fragment in A, Icon Ined. 1712; iso BR, CAL, K-W), Malaysia, P. Penang.

    Croton appendiculatus Elmer, Leafl. Philipp. Bot. 1 (1908) 312. — Type: Elmer 9215 (holo PNH†; iso L, NY, US), Philippines, Luzon.

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

Shrubs to small trees up to 15 m high, dbh up to 24 cm, dioecious, rarely monoecious. Bole up to 4 m high; crown up to 4 m long; bark with much tannic acid, outer bark rough to finely fissured or pustular lenticellate, reddish brown with yellowish brown lenticels, up to 10 mm thick; inner bark and wood white. Indumentum tomentose to puberulent. Stipules triangular, 0.8–1.5 by 0.4–0.7 mm, caducous, margin entire, apex acute. Leaves alternate to apically subopposite; petiole 30–180 by 0.4–0.7 mm, greenish tawny; blade subpeltate for 1.5 mm, broadly ovate to ovate, 4–23.5 by 3–15 cm, length/width ratio 1.2–2, reddish brown when young, base rounded to cuneate, margin entire to (irregularly) dentate, sometimes 2-lobed at widest part of blade, never hastate, longest lobes up to 30 mm, upper surface dull dark green, basally with 0–2 extrafloral nectaries, 1.8–5 by 1–2.5 mm, yellowish green, marginal nectaries 0–11 per side, 2–4 mm from margin, 0.7–0.8 by 0.7–0.8 mm, lower surface light brownish grey to coppery, domatia absent, 3-nerved, nerves 6–8 per side, looping or ending in the margin. Inflorescences axillary or terminal, unisexual to rarely bisexual, axes greenish tawny, basally 2–4 mm thick; bracts triangular to narrowly triangular, 1.9–2.2 by 0.3–0.4 mm, bracteoles 0.4–1 by 0.3–0.5 mm, persistent, margin entire. Staminate inflorescences up to 45 cm long, side branches up to 28 cm long, with 3–7 flowers per node, nodes per branch up to 70, hairy. Staminate flowers 3–5 mm diam.; pedicels 3–4 mm long; sepals 3 or 4, elliptic, reflexed, 2.2–3 by 1.2–1.8 mm, pale yellow; stamens 40–65, filaments 1.5–2 mm long, white, anthers 0.2–0.3 by 0.1–0.2 mm, orange-yellow; pistillode present. Pistillate inflorescences panicles, occasionally racemes, up to 30 cm long, nodes up to 75 (per branch). Pistillate flowers 3–4 mm diam.; pedicels up to 1 mm long; calyx 3–5-lobed, connate on the base, 2–2.6 mm long, cream brown, lobes ovate, 1.5–2.2 by 0.7–1 mm; ovary 3-locular, 1.3–1.5 by 1.7–2 mm, cream brown, spines individually visible; style up to 1 mm long, yellow; stigmas 1.2–2 mm long, yellow; staminodes absent. Fruits 5–12 by 6–14 mm, greenish tawny, spines few, straight to curved, thick, hairy, up to 7 mm long, hairs not forming a continuous layer, not rubbing off; column 3–3.3 by 2–2.2 mm. Seeds globose, 2.5–3 by 2.5–3 by 2–2.5 mm, smooth, black, shiny; hilum c. 1 by c. 1 mm.

    Distribution — From India to Taiwan, throughout Southeast Asia and Malesia to East Australia and New Guinea.

    Habitat & Ecology — Locally common in deciduous and evergreen forest, mostly in open, often very disturbed or burned places, in ravines and on dry sites like plateaus, ridges and slopes, in thickets, and along rivers and roadsides; on a large variety of soil types, like basalt, clay, granite, laterite, limestone, loam, rocky, sandy, ultrabasic, and volcanic. Altitude: sea level up to 1800 m. Flowering and fruiting the whole year through. It grows fast and was observed to be visited by ants and birds eating the fruits.

    Uses — The roots are boiled and drunk after child birth. The bark is used for construction, making strings, and it is used by the Loi (in China) to make a coarse cloth from which men’s jackets were made. The leaves are used against fever. The indumentum of the young leaves is applied on the penis after circumcision. The wood is used for paper pulp, wallboard, light construction, and as firewood.

    Vernacular names —Malaya Peninsula: Balek angin, Balek angin kechil (Semelai); Buerakaeputeh (Malay); Balek (Sakai); Balik angina, Musapal itam, Poko balek angina. Sumatra: Balik angin (Alas); Saringkut (Karo); Bali-bali angina, Balik angina, Balik-balik Angin silai, Boedi, Kajoe lappisi, Kajoe si balik angina, Kajoe si djoekkat, Kajoe si tarak balanggingngan, Lilouw. Java: Tjalik angin (Sundanese), Ki tjoelih angin perak, Ki-manggong, Ki-talikangin, Walung. Borneo: Sabah: Balabakan (Bandukam, Bajau); Parak balik (Dusun Kwijau); Dauh (Dusun Putatan); Entupak (Iban); Sirindien (Land Dayak); Balek angin. Sarawak: Dagoh (Dusun); Berlekut (Kelabit); Sirindienjamis (Land Dayak); Balek angin (Malay); Balikangina. Indonesia (Kalimantan): Butaq (Tunjung Benua); Bayur, Keminting boeroeng, Waring kangin laki. Philippines: Hinlaumong-puti (Bisáya); Anaplán (Bukídnon); Lamai. Sulawesi: Kajukuo (Uma), Kapilah. Papua (Irian Jaya): Kanibor (Kutubu); Lengle (Miwaute); Gayanbangemba (Waskuk); Bangiam, Kolamalama. Papua New Guinea: Poekwa (Hattam).

    Note — Next to M. paniculatus var. paniculatus the var. formosanus (Hayata) Hurus. is recognized. Both varieties occur in Taiwan and differ in partly overlapping characters (see Sierra & Welzen, 2005).

39. Mallotus peltatus (Geiseler) Müll.Arg. (sect. Stylanthus)

    Mallotus peltatus (Geiseler) Müll.Arg., Linnaea 34 (1865) 187; in DC., Prod. 15, 2 (1866) 967; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 174; in Engl. & Harms, Natürl. Pflanzenfam. ed. 2, 19c (1931) 115; Corner, Wayside Trees Mal. (1940) 272; Backer & Bakh.f., Fl. Java 1 (1964) 483; Airy Shaw, Kew Bull. 26 (1972) 307; Whitmore, Tree Fl. Malaya 2 (1973) 113; Airy Shaw, Kew Bull. 36 (1981) 327; Corner, Wayside Trees Mal., ed. 2 (1988) 307; Slik, Priyono & Welzen, Gard. Bull. Singapore 52 (2000) 62, Fig. 38; Slik & Welzen, Blumea 46 (2001) 41, Fig. 16, Map 7; Slik, Priyono & Welzen in Slik, Tropenbos-Kalimantan ser. 4 (2001) 99, Fig. 5.41; Slik & Welzen in Slik, Tropenbos-Kalimantan ser. 4 (2001) 184, Fig. 7.16, Map 7.7. — Aleurites peltatus Geiseler, Croton. Monogr. (1807) 81. — Type: Rottler s.n (C holo), in herbariumVahl as a Croton species from India orientalis.

    Adisca acuminata Blume, Bijdr. Fl. Ned. Ind. 11 (1825) 610. — Rottlera acutifolia Hassk., Cat. Hort. Bot. Bogor. (1844) 238; Miq., Fl. Ned. Ind. 1(2) (1859) 393. — Mappa acutifolia (Hassk.) Zoll. & Mor., Syst. Verz. (1855) 17. — Rottlera acuminata (Blume) Baill., Etude Euphorb. (1858) 426 (non A. Juss., 1824), nom inval. — Mallotus acuminatus (Blume) Müll.Arg., Linnaea 34 (1865) 187; in DC., Prod. 15, 2 (1866) 966; Kurz, Forest Fl. 2 (1877) 383; Hook.f., Fl. Brit. India 5 (1887) 431; J.J.Sm., Meded. Dept. Landb. Ned.-Indië 10 (1910) 422; Koord., Exkurs.-Fl. Java (1912) 492; Ridl., Fl. Malay Penins. 3 (1924) 290; S.Moore, J. Bot. 63 (1925) 103: Backer & Bakh.f., Fl. Java 1 (1964) 483. — Lectotype (Slik & Welzen, Blumea 46, 2001): Blume s.n. (L sheet 904 75 - 689) (L holo), Java.

    Rottlera longifolia Rchb.f. & Zoll., Verhand. Natuurk. Vereen. Ned.-Indië 1 (1856): 31; Miq., Fl. Ned. Ind. 1(2) (1859) 394; Fl. Ned. Ind. Eerste Bijv. (1862) 181; Scheff., Ann. Mus. Bot. Lugd.-Bat. 4 (1868-69) 123. — Mallotus longifolius (Rchb.f. & Zoll.) Müll.Arg., in DC., Prod. 15, 2 (1866) 967. — Mallotus longifolius (Rchb.f. & Zoll.) Müll.Arg. var. genuinus Müll.Arg., in DC., Prod. 15, 2 (1866) 967, nom. inval. — Lectotype (Slik & Welzen, Blumea 46, 2001): Teysmann s.n. (L sheet 904 105 - 380) (L holo), Sumatra.

    Rottlera oblongifolia Miq., Fl. Ned. Ind. 1(2) (1859) 396. — Mallotus oblongifolius (Miq.) Müll.Arg., Linnaea 34 (1865) 192; in DC., Prod. 15, 2 (1866) 973; J.J.Sm., Meded. Dept. Landb. Ned.-Indië 10 (1910) 425; Koord., Exkurs.-Fl. Java (1912) 491; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 193; Merr., J. Str. Roy. Asiatic Soc. (1921) 339; Ridl., Fl. Malay Penins. 3 (1924) 293; S.Moore, J. Bot. 63 (1925) 103; Pax & K.Hoffm. in Engl. & Harms, Natürl. Pflanzenfam. ed. 2, 19c (1931) 117; Backer & Bakh.f., Fl. Java 1 (1964) 484; Meijer, Bot. News Bull. (1967) 52; Airy Shaw, Kew Bull. 21 (1968) 388; Kew Bull. 26 (1972) 306; Whitmore, Tree Fl. Malaya 2 (1973) 116; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 173; Kew Bull. Add. Ser. 8 (1980) 168; Muelleria 4 (1980) 233; Kew Bull. 36 (1981) 327; Alph. Enum. Euphorb. Philipp. Isl. (1983) 37; Corner, Wayside Trees Mal., ed. 2 (1988) 306; Kessler & Sidiyasa, Tropenbos Series 7 (1994) 133. — Mallotus oblongifolius (Miq.) Müll.Arg. var. genuinus Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 193 nom. inval. — Type: Zollinger 245 (U holo; A, BM, G, L, MEL, P iso), Java, Tjikoja.

    Hancea muricata Benth., Fl. Hongk. (1861) 306. — Type: Hance 7298, Hongkong. (K holo; P, MEL iso).

    Rottlera flavigutta Miq., Fl. Ned. Ind., Eerste Bijv. (1862) 453. — Type: Teysmann HB 3667 (U holo), Sumatra orient. in prov. Palembang, prope Muara-enim.

    Mallotus porterianus Müll.Arg., Linnaea 34 (1865) 185; in DC., Prod. 15, 2 (1866) 960; Hook.f., Fl. Brit. India 5 (1887) 432; Boerl., Handl. Fl. Ned. Ind. 3 (1900) 288; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 192; Ridl., Fl. Malay Penins. 3 (1924) 292; S.Moore, J. Bot. 63 (1925) 103; Pax & K.Hoffm. in Engl. & Harms, Natürl. Pflanzenfam. ed. 2 (1931) 117; Burkill, Dict. Econ. Prod. Malay Penins. 2 (1935) 1398; Corner, Wayside Trees Mal. (1940) 273. — Lectotype (Slik & Welzen, Blumea 46, 2001): Porter 9094 (K holo), Peninsular Malaysia, Penang.

    Mallotus furetianus Müll.Arg., Linnaea 34 (1865) 190; in DC., Prod. 15, 2 (1866) 968; F.B.Forbes & Hemsl., J. Linn. Soc. Bot. 26 (1894) 439; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 194; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1925) 352; Pax & K.Hoffm. in Engl. & Harms, Natürl. Pflanzenfam. ed. 2, 19c (1931) 117. — Type: Furet 126 (P holo), China, Honkong.

    Mallotus lambertianus Müll.Arg., Linnaea 34 (1865) 190; in DC., Prod. 15, 2 (1866) 968; Boerl., Handl. Fl. Ned. Ind. 3(1) (1900) 289; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 194; in Engl. & Harms, Natürl. Pflanzenfam. ed. 2 (1931) 117. — Rottlera lambertiana (Müll.Arg) Scheff., Ann. Mus. Bot. Lugd.-Bat. 4 (1868-69) 125. — Type: Lambert s.n. (G holo, n.v.), Moluccas.

    Mallotus helferi Müll.Arg., Linnaea 34 (1865) 190; in DC., Prod. 15, 2 (1866) 968; Kurz, Forest Fl. 2 (1877) 384; Hook.f., Fl. Brit. India 5 (1887) 431; Boerl., Handl. Fl. Ned. Ind. 3(1) (1900) 289. — Mallotus oblongifolius (Miq.) Müll.Arg. var. helferi (Müll.Arg.) Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 194. — Type: Helfer 138 (G holo, n.v.), India orientalis, insula Kolonkhin fluminis Saluin.

    Mallotus longifolius (Rchb.f. & Zoll.) Müll.Arg. var. pubescens Müll.Arg., in DC., Prod. 15, 2 (1866) 967 — Lectotype (Slik & Welzen, Blumea 46, 2001): Zollinger 1812 (L holo), Sumatra.

    Mallotus stylaris Müll.Arg., in DC., Prod. 15, 2 (1866) 973; Boerl., Handl. Fl. Ned. Ind. 3(1) (1900) 290. — Rottlera stylaris (Müll.Arg.) Scheff., Ann. Mus. Bot. Lugd.-Bat. 4 (1868-69) 125. — Lectotype (Slik & Welzen, Blumea 46, 2001): Korthals s.n. (L sheet 904105-332) (L holo), Borneo.

    Mallotus puberulus Hook.f., Fl. Brit. India 5 (1887) 435; Boerl., Handl. Fl. Ned. Ind. 3(1) (1900) 290; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 172; Ridl., Fl. Malay Penins. 3 (1924) 290. — Type: Scortechini s.n. (K holo), Perak.

    Mallotus columnaris Warb., Bot. Jahrb. 13 (1891) 349; Boerl., Handl. Fl. Ned. Ind. 3(1) (1900) 289; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 176; Merr., Philipp. J. Sci. 11(5) (1916) 283. — Type: Warburg 20515 (B holo, lost; A, EDI iso), Kl. Key und Aru Inseln.

    Mallotus odoratus Elmer, Leafl. Philipp. Bot. 4 (1911) 1299; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 176; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 434; Pax & K.Hoffm. in Engl. & Harms, Natürl. Pflanzenfam. ed. 2, 19c (1931) 115. — Type: Elmer 12584 (PNH holo, lost; EDI, L, NY iso), Philippines, Palawan, Puerto Princesa (Mt Pulgar).

    Mallotus alternifolius Merr., Philipp. J. Sci. 7 (1912) 395. — Type: F.B. Curran 4124 (PNH holo, lost), Philippines, Palawan, near Puerto Princesa.

    Mallotus camiguinensis Merr., Philipp. J. Sci. 7 (1912) 397. — Type: B.S. Fénix 4047 (PNH holo, lost), Philippines, Babuyanes Islands, Camiguin.

    Mallotus oblongifolius (Miq.) Müll.Arg. var. siamensis Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 194. — Lectotype (Slik & Welzen, Blumea 46, 2001): J. Schmidt 691 (C holo, UC iso), Südwestmalayische Provinz, Siam.

    Mallotus oblongifolius (Miq.) Müll.Arg. var. villosulus Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 194; Airy Shaw, Kew Bull. 20 (1966) 42; Kew Bull. Add. Ser. 8 (1980): 169; Muelleria 4 (1980) 234. — Type: Lauterbach 1054 (B holo, lost), Papuasische Provinz, Neu-Guinea, Kaiser Wilhelmsland, am Gogolflusse.

    Mallotus tenuispicus Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. vii (1914) 201; Airy Shaw, Kew Bull. 21 (1968) 400. — Type: Schlechter 14467 (B holo, lost; K, WRSL iso), Papuasische Provinz, Neu-Guinea, Torricelli-Gebirge.

    Mallotus maclurei Merr., Philipp. J. Sci. 21 (1922) 347; Lingnaam Agric. Rev. 2 (1924) 29; Lingnan Sci. J. 5 (1927) 110. — Type: McClure 8558 (PNH holo, lost; A, NY iso), Ng Chi Leng.

    Mallotus peekelii Pax & K.Hoffm., Notizbl. Bot. Gart. Berlin-Dahlem 10 (1928) 384. — Type: Peekel 921 (B holo, lost; BRI iso), Papuasische Provinz, Neu-Mecklenburg, Lamekot.

    Mallotus oblongifolius (Miq.) Müll.Arg. var. rubriflorus Chakrab., J. Econ. Taxon. Bot. 6 (1985) 496. — Type: Chakrabarty 10174 (PBL holo, PBL iso), Great Nicobar Is., hill above Baludera.

    Mallotus peltatus (Geiseler) Müll.Arg. var. rubriflorus Chakrab., J. Econ. Taxon. Bot. 6 (1985) 497. — Type: Nair 3694 (PBL holo, PBL iso), South Andaman Is., Obragunj.

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

Shrubs to small trees up to 15(–21) m tall, dbh up to 30 cm; bole fluted, twisted to slanting; crown branching freely, wide spreading to rounded pyramidal. Outer bark smooth to fissured, lenticellate, mottled. Indumentum sparse to dense, whitish to cream. Branches lenticellate, glabrescent, gland-dotted on young parts. Stipules early caducous, triangular to ovate, 1.8–5 by 0.7–1.5 mm, margin entire to irregular, apex acute, hairy abaxially only, gland-dotted or not. Leaves alternate to apically subopposite, not peltate to peltate (sometimes both in one individual), distance between leaf blade base and petiole insertion 2–17 mm; petiole 4–110 by 0.5–1.5 mm, constricted to thickened basally and apically, hairy, especially basally and apically, glabrescent, gland-dotted or not, especially basally and apically; blade ovate to obovate, 3.5–21(–29) by 1–10(–13) cm, length-width ratio (1.3–)2–4, base attenuate to cordate, margin (slightly) dentate to irregularly wavy (to slightly crenate), marginal glands 10–28 per leaf side, apex acuminate to cuspidate to aristate, upper surface smooth (to pustulose), glabrous to hairy, especially on veins, sparsely gland-dotted or not, especially along veins, basal macular glands 2–7(–13) on veins, 1.5–14(–21) mm from petiole or midrib, apical macular glands 0–8 on veins, 1–9 mm from margin, lower surface smooth, hairy, especially on veins, glabrescent, gland-dotted or not, domatia present usually with hair tufts; venation pinnate to triplinerved to palmate, when peltate 8–10 veins originating from petiole insertion, 5–13 secondary veins per side along midrib, ending parallel to the margin. Staminate inflorescences up to 24 cm long, basally 0.5–1.5 mm thick, rarely grouped, hairy, gland-dotted, nodes up to 38(–70) per inflorescence, flowers 1–12 per node; bracts early caducous, deltoid to ovate, 0.9–2.3 by 0.4–1.2 mm, margin entire to irregular, apex rounded to acute, hairy abaxially only, gland-dotted outside only, especially basally; buds globose to ovate with acute to acuminate apex, glabrous to hairy, gland-dotted or not. Staminate flowers 3–6 mm in diameter, whitish green to cream to reddish, fragrant; pedicels 0.6–3 mm long, glabrous to hairy, gland-dotted or not; sepals 3–5, free to basally connate, (triangular to) elliptic (to obovate), often recurved, 1.5–2.5 by 0.5–1.3 mm, margin entire to irregular, apex acute to acuminate, glabrous to hairy abaxially only, gland-dotted or not; stamens up to 50; filaments 1–3.5 mm long, glabrous, sometimes gland-dotted; anthers 0.2–0.3 by 0.3–0.5 mm; connective widened, sometimes with protruding apex. Pistillate inflorescences up to 28 cm long, basally 1–2.8 mm thick, hairy, glabrescent, gland-dotted, nodes up to 29 per inflorescence; bracts caducous (to persistent), ovate, cup-shaped, half surrounding the pedicel, 1–2.8 by 0.7–1.5 mm, margin entire to irregular, apex acute, hairy abaxially only, gland-dotted or not. Pistillate flowers green to (light) yellow to yellow-red, fragrant; bracteoles absent to caducous, triangular to ovate, 0.8–1.3 by 0.3–0.6 mm, margin entire to irregular, apex acute, hairy abaxially only, gland-dotted outside only; pedicels 1–7 mm long, glabrous to hairy, gland-dotted or not; calyx caducous, lobes (1–)2–3(–4), free to more or less connate, triangular to ovate, 0.5–4.5 by 0.2–2.3 mm, margin entire to irregular, apex acute to acuminate, hairy abaxially only, gland-dotted or not; ovary (2–)3(–4)-locular, echinate, glabrous to hairy, densely gland-dotted, spines up to 5 mm long, hairy, usually ending in a few long hairs, rarely gland-dotted basally; style 2–7 mm long, glabrous to hairy, gland-dotted or not; stigmas 3–7 mm long, densely covered with granulate papillae on the inside, glabrous to hairy abaxially, gland-dotted outside. Fruit an echinate, lobed capsule, 7–15 by 5.5–8 mm, purple tinged to dark red, glabrous to hairy, densely gland-dotted; carpel inside hairy, gland-dotted or not; column 4–6 by 4–6.5 mm. Seeds glossy, roughly globose, 4–6 by 3–5 by 3–5.5 mm; hilum deltoid, 1–2.5 by 1–2.5 mm.

    Distribution — From India, the Andaman Islands and Southern China to New Guinea.

    Habitat & Ecology — Locally common in primary to secondary forests, preferring open places like river banks, forest edges, road sides, cleared areas, but sometimes also found in the forest understorey; both in wet (riverine, swampy) areas and on well drained soils; soils include thick humus, limestone, sandstone, sandy clay, sandy loam soil, volcanic rock, gravel, red shale. Altitude: up to 1000(–1800) m.

    Notes — This species is very variable in leaf shape and indumentum throughout its distribution range. This variability is the main reason for the large number of synonyms for this species. Airy Shaw had already reduced this abundance of names to two species, M. oblongifolius (without peltate leaves) and M. peltatus (with peltate leaves). He suggested that these two species should be united into one species, but did not do so himself. Considering the large variablity in leaf shape, the fact that some individuals have both peltate and non-peltate leaves on the same branch and the lack of other differentiating characters, we have to conclude that M. peltatus and M. oblongifolius are in fact one and the same species. In general there are three main leaf types: 1) the obovate-penniveined type which usually occurs in India, Southern China, Thailand, Malaysia, Borneo, Sumatra and Java. 2) the cordate-tripliveined type which usually occurs on the Andamans, Sumatra, Malaysia, the Philippines, Sulawesi, the Moluccas and New Guinea. 3) the peltate-palmativeined type which is usually found on the Andamans, Malaysia, Sumatra, Northern Borneo, Java and the Lesser Sunda Islands. The indumentum of the plants ranges from almost completely glabrous to densely hairy on most parts. In general the plants found in Sumatra, Malaysia, Northern Borneo, Java, and New Guinea can be very hairy. Some specimens collected in Sumatra, Malaysia, Vietnam and Northern Borneo are very similar to M. lackeyi, but differ from that species in their smaller, early caducous stipules, not densely gland-dotted upper leaf surface, longer styles, and longer spines of the fruits.

40. Mallotus philippensis (Lam.) Müll.Arg. (sect. Philippinensis)

    Mallotus philippensis (Lam.) Müll.Arg., Linnaea (1865) 196 (‘philippinensis’); Backer & Bakh.f., Fl. Java 1 (1964) 483; Airy Shaw, Kew Bull. 26 (1972) 300; Whitmore, Tree Fl. Malaya 2 (1973) 115; Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 168; Kew Bull., Addit. Ser. 8 (1980) 170; Kew Bull. 36 (1981) 328; Kew Bull. 37 (1982) 29; Alphab. Enum. Euph. Philipp. Isl. (1983) 37; Philcox in Dassan. & W.D.Clayton, Rev. Handb. Fl. Ceylon 11 (1997) 158; P.I.Forst., Austrobaileya 5 (1999) 480; Welzen et al., Thai Forest Bull., Bot. 28 (2000) 105; Keßler, Blumea, Suppl. 14 (2002) 50; W.J.Kress et al., Checkl. Myanmar, rev. ed. 4 (2003) 232; S.E.C.Sierra, Welzen & Slik, Blumea 50 (2005) 230, fig. 4, map 1. — Croton philippense Lam., Encycl. 2 (1786) 206. — Rottlera philippensis (Lam.) A.Juss. ex Spreng., Syst. Veg. 2 (1826) 877; Scheff., Ann. Mus. Bot. Lugd.-Bat. 4 (1869) 124 (‘philippinensis’: isonym). — Echinus philippensis (Lam.) Baill., Adansonia 6 (1866) 314 (‘philippinensis’). — Type: Sonnerat s.n. (holo P, barcode P00279571), Philippines, see note 1.

    Croton punctatus Retz., Observ. Bot. 5 (1789) 30 (‘punctatum’), non Jacq. (1787). —Type: J. König s.n. (holo LD; iso C), Sri Lanka, see note 2.

    Croton coccineus Vahl, Symb. Bot. 2 (1791) 97. — Lectotype (Sierra et al., 2005): J. König s.n. (holo C; iso C, LD), Sri Lanka, see note 2.

    Rottlera tinctoria Roxb., Pl. Corommandel 2, 3 (1802) 36, t. 168. — Lectotype (Sierra et al., 2005): Wallich Numer. List 7832A (holo K, photo in L; iso K (photo in L), Icon Ined. 106 (CAL, K)), India.

    Croton montanus Willd., Sp. Pl. ed. 4, 1 (1805) 547. — Type: Klein s.n. (holo Herb. Willd. 17874, (IDC microfiche no. 1261), B), India.

    Rottlera aurantiaca Hook.&Arn., Bot. Capt. Beechey Voy. 6 (1838) 270.—Type: Unknown s.n. (holo K, barcode K000185510), China, Fujian, Loo Choo.

    Rottlera tinctoria Roxb. var. monstruosa Ham. ex Dillwyn, Rev. Ref. Hortus Malab. 5 (1839) 22; Mabb., Taxon 26 (1977) 532; Nicolson, Suresh & Manilal, Interpr. Van Rheede Hort. Malab. (1988) 112; Manilal, Van Rheede Hort. Malab. 5 (2003) 87. — Type: Van Rheede tot Draakestein & Munnicks, Hort. Malab. 5 (1685), t. 24 (holo), India.

    Rottlera affinis Hassk., Flora 25 (1842) 41. — Type: Unknown s.n. (holo BO?, n.v.), Indonesia, W Java.

    Mappa stricta Rchb.f. & Zoll., Acta Soc. Regiae Sci. Indo-Neerl. 1 (1856) 31; Linnaea 28 (1857) 310; Miq., Fl. Ned. Ind. 1, 2 (1859) 402.— Macaranga stricta (Rchb.f. & Zoll.) Müll.Arg. in DC., Prodr. 15, 2 (1866) 1004.—Type: Korthals s.n. (holo BO?, n.v.), Indonesia, Sumatra, Prov. Padang, Prope Lubu alang (= Lubukalung), Mantjiro inc. 24 April 1834.

    Aconceveibum trinerve Miq., Fl. Ned. Ind. 1, 2 (1859) 389. — Type: Zollinger 1101 (holo U), Myanmar.

    Mallotus reticulatus Dunn, J. Linn. Soc., Bot. 38 (1908) 365. — Mallotus philippensis (Lam.) Müll.Arg. var. reticulatus (Dunn) F.P.Metcalf, J. Arnold Arbor. 22 (1941) 207. — Type: Hongkong Herb. 3429 (holo HK; iso K), China.

    Euonymus hypoleucus H. Lév., Repert. Spec. Nov. Regni Veg. 13 (1914) 260. — Type: Cavalerie 2733 (holo E; iso E, K), China, Kweichow, Lo-fou, 4 April 1906.

    Mallotus philippensis (Lam.) Müll.Arg. var. tomentosa Gamble, Fl. Presid. Madras 2, 7 (1925) 1322. — Lectotype (Sierra et al., 2005): Gamble 14368 (K), India, Connoon Ghat.

    Mallotus bicarpellatus T.Kuros., Edinburgh J. Bot. 61 (2004) 31. — Type: Nicolson 2282 (holo TI; iso BM, KATH, L), Nepal, Bagmati Zone, Bhaktapur District.

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

Shrubs to small trees up to 25 m high, dbh up to 48 cm; bole up to 15 m high; crown up to 18 m long. Outer bark smooth to shallowly fissured, up to 8 mm thick, grey with patches of brown, on cross section reddish brown; sapwood cream, heartwood pink to brown-coloured. Indumentum tomentose, often villous, composed of stellate and/or simple hairs, and sessile, globular to disc-shaped, red (sometimes dull yellow to orange, rarely black) glandular hairs. Stipules triangular, 1–1.5 by 0.4–0.7 mm, caducous to persistent. Leaves alternate to apically subopposite; petiole 15–130 by 0.8–2 mm, basally and apically pulvinate; blade ovate to elliptic, sometimes obovate, 4–25 by 2–13 cm, length/width ratio 1.2–5.2, coriaceous, base truncate to cuneate, margin entire, rarely dentate to serrate (with glandular teeth), slightly sinuate, apex obtuse to acuminate, upper surface dull green, basally with 2 (or 4) extrafloral nectaries on the blade (0–3 mm from petiole inser-tion), orbicular to elliptic, (0.4–)1–3 by (0.3–)0.8–1.2 mm, lower surface greenish grey to brown, not glabrescent, domatia rarely present, nerves 6–11 per side, looping. Inflorescences racemes, axes brownish, basally 1–2 mm thick; bracts persistent, hairy outside, glabrous to subglabrous inside. Staminate inflorescences up to 17 cm long, flowers 1–3(–4) per node, nodes up to 45; bracts 0.5–0.8 by 0.6–1.2 mm. Staminate flowers 3–5 mm diam.; pedicels 1.8–3.5 mm long; sepals 3 or 4, ovate to elliptic, 2–3 by0.7–2.5mm; stamens 18–33,filaments0.5–4mm long,glabrous,anthers ellipsoid, 0.5–0.8 by 0.3–0.4 mm, glabrous to sparsely hairy,lightyellow, connectivenotbroad; pistillode absent. Pistillate inflorescences up to 21 cm long, nodes up to 40; bracts 0.7–1.5 by 0.6–1.2 mm. Pistillate flowers 4–7 mm diam., pedicels 0.5–2 mm long; sepals (3 or) 4 or 5, narrowly triangular, 1–2 by 0.7–1.8 mm, reflexed; ovary (2- or) 3-locular, 1–1.5 by 1–1.8 mm; style up to 1 mm long; stigmas 2–7 mm long. Fruits 4–12 by 4.5–11 mm, opening loculicidally-septicidally, red; wall 0.3–0.5 mm thick; column 3–7 by 2–4 mm. Seeds ± globose, 3–5.5 by 3–5.5 by 2.5–5.2 mm, surface smooth, glossy, black; hilum 1–2.5 by 1–2 mm.

    Distribution — From Pakistan to South China and South Japan, throughout Southeast Asia and Malesia to East Australia and West Pacific (Solomon Islands).

    Habitat & Ecology — Locally common in understorey of primary to secondary forests, scrub, mostly found on disturbed sites; on ridges, forest edges, road and river sides, steep slopes, marshy and savannah areas; in wet (riverine, swampy) to well-drained terrains; on a large variety of soil types, like granite, limestone, sandstone, sandy clay, sandy loam soil, volcanic rock, gravel, quartz, shale, and rock. Also cultivated (Hawaii and Miami, USA). Altitude: sea level up to 1600 m. Flowering and fruiting the whole year through.

    Uses — Ornamental (red fruits). The wood is used for rafters, tool handles, matchboxes, and house-posts. The fruits and bark are used as an antihelminthic, to relieve constipation, and against cutaneous affections. The fruits and leaves are used against colds and to cure stings and bites of snakes and other poisonous animals. The leaves are used as a fodder. The glands of the fruits are used as a red dye. The roots are used for dissolving coagulated blood and contusions. The oil of the seeds is used as a substitute for tung oil (Vernicia Lour., Euphorbiaceae) in the formulation of rapid-drying paints, varnishes, hair fixers and ointments. Partly after Ambasta (1986) and Manilal (2003).

    Vernacular names —Malay Peninsula:Min-ya-ma-ya, Mue-ra-kae-pu-te (Malay); Balikangin, Mingak madia, Kasiran. Sumatra: Kajoe poetat, Madang mansiro, Masiho, Moacho, Toeba sira, Tumbasira. Java: Kangke djuan (Madurese); Kaju tike, Kapasan, Kemesoe, Palan, Papasan, Tal tidang, Talaman, Tapèn, Tekeg-tekegan. Borneo: Sabah: Ayagkun (Bajau Tuaran); Asin asin, Mata kunau (Dusun), Bai bai (Dusun Tampoluri); Magundasing (Murut Tenom); Balinasi (Kwijau). Philippines: Dalunis (Ibanág); Banáto (Ibanág, Igorot, Tagálog); Anangkuli, Buas. Sulawesi: Paedje-paedje. Lesser Sunda Islands: Hotel ewi (Bunag); Biofluke, Bnafo (Dawan); Hadju puser, Kaika, Kajoe tapis, Poeser, Pudjar, Pure, Puser. Moluccas: Galoega foeroe (Ternate), Intiboro. New Guinea: Papua (former Irian Jaya): Kikindam (Kotte); Maile (Kulumo); Poiro (Matapaili), Nakokupote (Minu); Ugo (Musa, Safia); Gamete (Naukwate, Onjob); Matsiuna (Roro); Iruda, Mamada, Nakunmenia. Papua New Guinea: Damon (Andjai), Rah (Gab Gab); Dammoen, Dammonnie, Domoni (Ke-bar); Sies (Manikiong); Andeh, Haha, Sebijreraka, Nokloe.

    Notes — 1. Forster (1999) mentioned as the holotype of M. philippensis the sheet P 16581, but this number refers to a Commerson s.n. collection of M. papillaris.

2. Croton punctatus and C. coccineus were based on different specimens in different herbaria, both with the same annotations of J. König. The name C. coccineus, being based on a different specimen, is not superfluous.

3. Under the number Wallich Numer. List 7772A there are two very distinct species as can be seen on the IDC microfiche nr. 7394 of the Wallich herbarium in K. The specimen found on the left hand base corner is M. philippensis. The one found on the right hand base corner is M. distans as it is generally known based on Müller Argoviense’s description of the fruits (densely hairy mixed with punctuate glandular hairs).

4. The collections Ludwigs 247 and 465 (M) have on the label as locality ‘Kamerun: Victoria’, but it is likely that this is a mistake.

5. The sheets Dickason 6641, Unknown s.n. (L 0436439) and Zollinger 3873, have aberrant staminate and pistillate inflorescences, respectively; and Kukkonen 6791 has aberrant stems; all of them are probably infected by a virus (these sheets look the same as Rottlera tinctoria Roxb. var. monstruosa Ham. ex Dillwyn).

41. Mallotus pleiogynus Pax & K. Hoffm. (sect. Rottleropsis)

    Mallotus pleiogynus Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 187; Kulju, S.E.C.Sierra & Welzen, Blumea 52 (2007) 128, fig. 3, map 4. — Octospermum pleiogynum (Pax & K.Hoffm.) Airy Shaw, Kew Bull. 19 (1965) 312; Hooker’s Icon. Pl. 38 (1974) t. 3716; Kew Bull., Addit. Ser. 8 (1980) 176, t. 8. — Type: Hollrung 782 (holo B, †; iso K), New Guinea, Kaiser-Wilhelmsland, Augusta Station.

Trees, up to 43 m high, dbh up to 70 cm, dioecious, evergreen; buttresses absent to up to 2 by 1.2 m; older branches more or less glabrescent, flowering branches 2–4 mm thick. Outer bark smooth, brown to grey; inner bark white to yellow within, cross section yellowish brown to brown. Indumentum dense in most parts, usually consisting of both short, (stellately) tufted and longer pilose, simple to (stellately) tufted hairs, whitish to brownish, sometimes one hair type rare or absent; glandular hairs present on some parts, yellowish to brownish to red. Stipules absent. Leaves alternate, subpeltate; petiole 3–13.5 cm long, usually with a basal constriction when dry, sparsely to densely hairy, often partly glabrescent; axillary buds often 2 above each other; blade ovate (to elliptic), 8.8–22.5 by 7–16 cm, length/width ratio 1.2–1.7, papery, base rounded to obtuse (to acute), margin (sub)entire, apex acuminate to cuspidate; upper surface subglabrous (to sparsely hairy), glabrescent, usually granular dotted, basal extrafloral nectaries 2(–4), conspicuous, dark brown, raised, close together at the petiole insertion, marginal extrafloral nectaries 9–20 per side, smaller, on venation, 0.5–2 mm from the margin; lower surface densely hairy, glandular hairs present, yellowish (to ± colourless), hair-tuft domatia sometimes present; venation triplinerved, nerves 5–8 per side, looped and closed near or at the margin. Inflorescences axillary racemes, single, or 2 together when staminate; bracts persistent to caducous, triangular, margin entire, apex acute to acuminate, hairy on both sides. Flowers: pedicels with a basal abscission zone; buds apiculate; sepals triangular to ovate, hairy outside. Staminate inflorescences up to 17 cm long, basally 1.1–1.8 mm thick; bracts 0.5–1.8 by 0.2–0.4 mm. Staminate flowers: buds globose; flowers 1.8–3.5 mm diam.; pedicel 0.5–2.6 mm long; sepals 3 or 4, 1–2.4 by 0.7–2.1 mm, usually unequal in width, apex acute to acuminate, glabrous inside; receptacle swollen into a torus; disc-glands absent; stamens 15–50, filament 0.3–0.8 mm long, free, glabrous, thecae 0.3–0.4 by 0.2–0.3 by 0.1–0.2 mm, ellipsoid, opening latrorsely, connective papillose, darker coloured, conspicuously peltate, covering the thecae like an umbrella, thecae and connective usually with some yellowish glandular hairs; pistillode absent. Pistillate inflorescences up to 7.8 cm long, basally 1.5–2.4 mm thick; flowers 4–8; bracts present on pedicel below abscission zone and often somewhat adnate to it, 1.2–2.4 by 1.2–1.5 mm, bracteoles sometimes present, near the abscission zone, smaller than the bracts. Pistillate flowers: buds ellipsoid, flowers 3.5–7 mm diam.; pedicel 2.4–6 mm long; sepals (4 or) 5 (or 6), 2.7–4.5 by 1.1–2.5 mm, recurving and persistent in fruit, apex acute to acuminate, subglabrous (to sparsely hairy) inside; ovary (sub)glabrous, covered with glandular hairs, locules (7 or) 8 or 9; stigmas 2.2–4.7 mm long, sessile, ± erect, slightly connate at base, narrowly triangular, plumose, outer surface with a ± hairy groove with glandular hairs. Fruits indehiscent drupes, 2–4 per infructescence, transversely ellipsoid, 9–12 by 18–22 mm, with 7–9 distinct longitudinal ridges when dry, surface somewhat irregulary wrinkled, covered with glandular hairs, yellow to orange when fresh, yellowish to reddish brown with dark grey ground colour when dry; pericarp toughly fleshy, hard when dry. Seeds radially arranged; obloid, somewhat flattened, 5.2–5.6 by 6.2–6.8 mm, surface slightly rugose, brown to light brown, shiny, sarcotesta absent.

    Distribution — New Guinea.

    Habitat & Ecology — Locally rather common in primary or sometimes secondary forest; terrain usually flat; soil sandy to clayey, rarely loamy or limestone. Altitude up to 660 m. Flowering and fruiting: throughout the year.

    Vernacular names — Irian Jaya: Aromaraipi (Roberbai, Japen dialect); Bangembo

(Waskuk); Manggafafin (Biak); Naroe, Sinaroe, Tinaro (Mooi); Sebijreraka, Seborreraka

(Manikiong); Barorong, Boear, M’brèrehie, Njakwe, Pembrièn. Papua New Guinea:

Fofoto (Onjob, Koreaf dialect); Osari (Orokaiva, Mumuni dialect); Sikusikura (Mana-

galase); Hopkos mange.

42. Mallotus polyadenos F.Muell. (sect. Polyadenii)

    Mallotus polyadenos F.Muell., Fragm. 6 (1868) 184; Benth., Fl. Austr. 6 (1873) 142; Bailey, Syn. Queensl. Fl. (1883) 479; Queensl. Fl. 5 (1902) 1448; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 198; Airy Shaw, Kew Bull. 20 (1966) 43; Kew Bull. Add. Ser. 8 (1980) 170; Kew Bull. 35 (1980) 655; Bollendorff, Welzen & Slik, Blumea 45 (2000) 329, fig. 1c; map 4; in Slik, Tropenbos-Kalimantan ser. 4 (2001) 136, Fig. 6.1c, 6.8; Map 6.4. — Lectotype (Bollendorff et al., 2000): Dallachy s.n., MEL sh. no. 708736 (MEL), Australia, Queensland, Rockingham Bay.

Shrub to tree, up to 28 m high, dioecious or rarely monoecious, mainly glabrous; branchlets sometimes lenticellate, ribbed, whitish grey and red-brown mottled to dark brown. Stipules triangular, half conical, 0.8–1.5 by 0.4–1 mm, (late) caducous, dark red to brown, apex acute. Leaves opposite to subopposite to rarely alternate, pairs unequal in size; petiole terete and deeply channelled to reniform in transverse section, 0.4–25 by 0.4–2 mm, glabrous to glabrescent, base and apex slightly to clearly pulvinate with transversal cracks; blade ovate to obovate, 4–17.8 by 1.4–7.8 cm, index 1.5–4.3, coriaceous to rarely chartaceous, base acute to rounded and slightly emarginate, margin entire, rarely dentate, 0–5 marginal glands per side, protruding in young leaves, intruding in old ones, triangular to round, apex obtusely acute to acuminate, rarely ending in a margin-type gland, upper surface green to silvery grey to brown, basally 0–10 impressed elliptic glands per side, c. 1 by 0.5 mm, brown or black, 0–45 mm from petiole attachment, on first pair of nerves and veins, lower surface green to olive-green to brown, domatia sometimes pocket-like with simple villose to velutinous white hairs; venation pinnate to often basally triplinerved, basal nerves ending in margin below middle, rare above, 4–9 nerves per side, brachidodromous, veins scalariform close to midrib, gradually becoming reticulate towards margin. Inflorescences axillary to terminal, solitary or 2 or 3 together, straight to rarely sinuous; bracts triangular to rhomboid, margin ciliate towards base, apex acute to acuminate, sometimes patent. Staminate inflorescences 2.8–20.7 cm by 0.3–0.8 mm, 20–47 nodes per rachis, 3–5 flowers per node; bracts 0.5–2 by 0.5–1.2 mm, basally often thickened; pedicels terete to angular, 0.8–4 by 0.3–0.5 mm, abscission zone at very base, densely covered with stellate hairs; flowers 4–7 mm in diam., calyx lobes 3–5, ovate to elliptic, 2.5–4.5 by 0.8–2.8 mm, margin rarely ciliate, apex occasionally recurved with few villose hairs, yellow to red-brown, stamens 20 to over 100, filaments 1.5–3 mm long, anthers ellipsoid to reniform, 0.3–0.5 by 0.2–0.8 mm, basifixed to dorsifixed, margin occasionally brown mottled, connective basally broad, deeply to entirely split, receptacle c. 0.3 mm high by 0.5 mm in diam. Pistillate inflorescences 3–21 cm by 0.5–2 mm, 12–45 nodes per rachis, 1–3 flowers per node; bracts 0.5–1.2 by 0.8–1.3 mm, base decurrent, margin concave; pedicels terete to flattened, 1–28 by 0.1–0.6 mm, abscission zone 0–1.2 mm from apex; flowers 1–4 mm in diam., calyx persistent, inside basally ring of villose hairs, lobes 3–5, ovate to triangular, 1.5–4 by 0.5–3 mm, yellow to red-brown, basally thickened, margin ciliate, apex acute with few villose hairs, ovary ovoid (2)3(4)-locular, covered with discoid glands, inside basally often with some villose hairs, stigmas subsessile, persistent, 0.8–4 by 0.4–1.2 mm, decurrent. Fruits subglobose, dehiscent, woody capsules, deeply trigonous, 4–6 by 7–10 mm diam., surface rarely with some stellate hairs, cocci twisting apically after dehiscence, each locule 3–6 by 2–5 mm wide; column more or less anchor-shaped in lateral view, c. 3 by 0.5–1 mm wide at base and 3–4.5 mm wide at apex. Seeds globose to subglobose, 3–4 mm in diam., with a few granular dots, light to dark brown.

    Distribution — New Guinea, Australia (NE Queensland).

dots = M. polyadenos; stars = M. puber.

    Habitat & Ecology — Dry, open or closed, often riparian rainforest, in Melaleuca-Eucalyptus-Tristania forest as well as in secondary swamps or in cleared dune woodland, on moderate slopes, on coral limestone. Alt.: sea level to 800 m. Flowering: May–Dec.; fruiting: Aug.–Dec.

    Note — One specimen only had branched staminate inflorescences which are terminal and sinuous (Clemens s.n., Australia, Queensland, Mount Fox; GH).

43. Mallotus puber Bollendorff  (sect. Polyadenii)

    Mallotus puber Bollendorff in Bollendorff, Welzen & Slik, Blumea 45 (2000) 334, fig. 1d, 5; map 4; in Slik, Tropenbos-Kalimantan ser. 4 (2001) 138, Fig. 6.1d, 6.9; Map 6.4. — Mallotus leucodermis Hook.f. var. puberulus Airy Shaw, Kew Bull. 21 (1968) 396. — Type: BSIP (Whitmore) 2998 (K, holo; iso in L), Solomon Islands, Choiseul.

Small tree, up to 15 m high, dioecious, pubescent to glabrous; branchlets sometimes lenticellate, glabrous to puberulous to hirsute, ribbed, reddish to greyish to dark brown. Stipules ridged, 2–2.5 by c. 1 mm, mainly early caducous, glabrous, apex acute to acuminate. Leaves alternate to rarely opposite, pairs unequal in size; petiole terete to angular and often narrowly channelled adaxially, 14–90 by 1–3 mm, glabrous to hirsute, base and apex usually not to slightly pulvinate, sometimes twisted; blade elliptic to ovate, 12–27 by 7.2–14.3 cm, index (1.1–)1.5–2.1, chartaceous to rarely subcoriaceous, base broadly rounded to cordate, margin entire, sometimes with a few irregular teeth, (0–)3–11 inconspicuous marginal glands per side in sinuses, apex acute to obtusely acuminate, ending in a margin-type of gland (sometimes giving a mucronate aspect), upper surface glabrous to hirsute, simple and stellate hairs on venation, dark green to brown, basally 0–13 circular, randomly distributed glands, c. 0.5 mm in diam., brown or black, lower surface olive green to brown, domatia with long simple and short stellate hairs, venation basally triplinerved to almost palmate with basal nerves ending in margin at or above middle, further pinnate, with 3–5 nerves per side, veins scalariform with 4–6 mm wide intervals, veinlets reticulate to scalariform. Inflorescences axillary, solitary or 2 or 3 together, rachis angular; bracts basally decurrent, margin ciliate, apex acute, patent. Staminate inflorescences 3–8.5 cm by 0.5–1 mm, 39–46 nodes per rachis, mainly 3 flowers per node; bracts c. 1 by 1 mm; pedicels terete to flattened, 1.5–2 by 0.4–0.5 mm, abscission zone 1–1.5 mm from apex, indumentum dense; flowers only seen in bud, calyx lobes 4, elliptic, c. 2 by 1.5 mm, discoid glands inside, apex acute, yellowish brown, stamens c. 50, anthers ellipsoid, dorsifixed, connective splitting. Pistillate inflorescences 8–38.5 cm by 0.8–3 mm, 13–36 nodes per rachis, 1 flower per node; bracts 0.8–1.2 by c. 1 mm; pedicels terete or angular, 7–48 by 0.5–0.7 mm, generally geniculate at abscission zone, latter 0.5– 7 mm from apex; flowers 3–4 mm in diam., calyx lobes 6, triangular to narrowly triangular, 1.5–1.8 by 0.3–0.5 mm, subcoriaceous, apex acute, few to densely hirsute hairs on the outside, dark red brown, ovary reniform to ellipsoid, c. 2.5 by 2 mm, 2(3)-locular, covered with discoid glands and rarely with hairs, stigmas sessile, 2–4 by 0.5–1 mm, decurrent, few hairs. Fruits subglobose, woody, dehiscent capsules, 5–8 by 7–10 mm in diam., without wings; locules globose to ovoid, c. 5 mm wide, inside basally with stellate and simple hairs; column anchor- to bell-shaped in lateral view, 4–5 by 1–2 mm wide basally and 3.2–5 mm wide apically. Seeds globose to ovoid, 4–5 mm in diam., dull, sometimes with a few granular dots, light to dark brown with darker venation; micropyle protruding conspicuously.

    Distribution — Solomon Islands (Choiseul, Kolombangara, New Georgia, San Isabel).

dots = M. polyadenos; stars = M. puber.

    Habitat & Ecology — Lowland primary forest, mostly well drained hills, rare in swamp forest. Alt.: up to 130 m. Flowering: July–Nov.; fruiting: Dec.–Feb. and Aug.–Sep.

    Vernacular names — Ketonwane, Faiketo, Raumomote, Malaketo (Kwara'ae).

    Note — The epithet puber is used, because puberulus already exists on the species level (Mallotus puberulus Hook. f.).

44. Mallotus repandus (Rottler) Müll.Arg. (sect. Philippinensis)

    Mallotus repandus (Rottler) Müll.Arg., Linnaea (1865) 197; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1925) 365; Backer & Bakh.f., Fl. Java 1 (1964) 483; Airy Shaw, Kew Bull. 26 (1972) 301; Whitmore, Tree Fl. Malaya 2 (1973) 114; Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 170; Kew Bull. 36 (1981) 328; Kew Bull. 37 (1982) 29; Alphab. Enum. Euph. Philipp. Isl. (1983) 37; McPherson & Tirel, Fl. Nouvelle-Calédonie 14 (1987) 106, pl. 20; Philcox in Dassan. & W.D.Clayton, Rev. Handb. Fl. Ceylon 11 (1997) 165; P.I.Forst., Austrobaileya 5 (1999) 484; Welzen et al., Thai Forest Bull., Bot. 28 (2000) 107; Slik, Priyono & Welzen, Gard. Bull. Singapore 52 (2000) 63; in Slik, Tropenbos-Kalimantan ser. 4 (2001) 99, Fig. 5.44; Keßler, Blumea, Suppl. 14 (2002) 50; W.J. Kress et al., Checkl. Myanmar, rev. ed. 4 (2003) 232; ; S.E.C.Sierra, Welzen & Slik, Blumea 50 (2005) 234, fig. 5, map 3. — Croton repandus Rottler, Ges. Naturf. Freunde Berlin Neue Schriften 4 (1803) 206 (‘repandum’). — Mallotus repandus (Rottler) Müll.Arg. var. genuinus Müll.Arg., Linnaea (1865) 197, nom. inval. — Rottlera repanda (Rottler) Scheff., Ann. Mus. Bot. Lugd.-Bat. 4 (1869) 124. — Lectotype (Sierra et al., 2005): Rottler s.n. = Wall. Numer. List 7774, fide specim. in Herb. Wall. (IDC microfiche no. 7394) (holo LIV; iso K-W, L s.n. barcode L0436491 ex. CAL), India, Marmelon.

    Croton rhombifolius Willd., Sp. Pl. ed. 4, 1 (1805) 555.— Rottlera rhombifolia (Willd.)Thwaites, Enum. Pl. Zeyl. (1861) 272.— Type: Roloff s.n. (holo B, Herb. Willd. 17900, (IDC microfiche no. 1261)), Sri Lanka.

    Helwingia populifolia Spreng., Pl. Cogn. Pug. (1815) 89; Airy Shaw, Kew Bull. 15 (1962) 419. — Type: Unknown s.n. (holo B, †), India, Bengalia.

    Rottlera scabrifolia A.Juss., Euphorb. Gen. (1824) 111, t. 9, f. 29B. — Mallotus repandus (Rottler) Müll.Arg. var. scabrifolia (A.Juss.) Müll.Arg., Linnaea (1865) 197. — Type: Unknown s.n. (Herb. A. Juss. 16583, (IDC microfiche no. 6206), P), Indonesia, Timor.

    Rottlera viscida Blume, Bijdr. (1826) 608. — Lectotype (Sierra et al. 2005): Blume 1716 (holo L, barcode L0436521; iso L barcodes L0436522, L0436536), Indonesia, Java, Nusa Kambanga.

    Rottlera scandens Span., Linnaea 15 (1841) 348. — Mallotus scandens (Span.) Müll.Arg. in DC., Prodr. 15, 2 (1866) 982. — Type: Spanoghe s.n. (holo L, barcode L 0293546), Indonesia, Lesser Sunda Islands, Timor.

    Rottlera dicocca auct non. Roxb.: Roxb. [Hort. Beng. (1814) 73, nom. nud.] Fl. Ind. 3 (1832) 829, pro specim.; Baill., Étude Gen. (1858) 423 (‘dioica’ Roxb.).—Vouchers: Roxburgh s.n. (BM barcode BM00813956; BR barcode 849411; Icon Ined. 480 (K, CAL)), India.

    Croton volubilis Llanos, Mem. Real Acad. Ci. Exact. Madrid 4 (1856) 503. — Neotype (Sierra et al., 2005): Merrill Species blancoanae 842 (holo US; iso L, P), Philippines, Luzon, Rizal.

    Rottlera? cordifolia Benth., Fl. Hongkong. (1861) 307. — Type: Wright s.n. (holo K barcode K000185495; iso NY barcode NY00273207, US barcode 00096200), Hongkong.

    Trewia nudiflora auct non. Willd: Hance, J. Bot. 16 (1878) 14. — Voucher: Sampson 9420 (K), China, Guan-dong.

    Mallotus contubernalis Hance, J. Bot. 20 (1882) 293. — Lectotype (Sierra et al., 2005): Sampson & Hance 17694 (K), China, Guandong, West River, Ting-ü-shan.

    Mallotus chrysocarpus Pamp., Nuovo Giorn. Bot. Ital. 17 (1910) 413. — Mallotus contubernalis Hance var. chrysocarpus (Pamp.) Hand.-Mazz., Symb. Sin. (1931) 214. — Mallotus repandus (Rottler) Müll.Arg. var. chrysocarpus (Pamp.) S.M.Hwang, Acta Phytotax. Sin. 23 (1985) 297. — Type: Silvestri 1290 (holo FI), China, Hubei, ‘Monte Triora’.

    Mallotus contubernalis auct. non Hance: Pax & K. Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 180. — Mallotus illudens Croizat, J. Arnold Arbor. 19 (1938) 146. — Lectotype (Sierra et al., 2005): Wang 239 (holo A; iso NY), China, Jiangxi.

    Mallotus repandus (Rottler) Müll.Arg. var. megaphyllus Croizat, J. Arnold Arbor. 19 (1938) 146.—Type: Kingdon Ward 8922 (holo A; iso NY), Laos, Mong Hsing.

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

Woody climber up to 10 m high. Outer bark dark brownish grey. Indumentum sparse, composed of stellate and/or simple hairs, and sessile, globular to disc-shaped, light yellow to orange glandular hairs. Stipules triangular, 0.5–0.6 by 0.3–0.6 mm, per-sistent to caducous. Leaves alternate to apically subopposite; petiole (10–)24–64 by 0.8–2 mm, basally and apically not pulvinate; blade broadly ovate to elliptic, 4–19 by 3–13.5 cm, length/width ratio 1.1–2.2, chartaceous, base cordate to cuneate, margin with glandular teeth, apex acute to caudate, upper surface dull dark green, glabrescent, rarely densely hairy, marginally with 2–4(–10) extrafloral nectaries per side, orbicular to elliptic, 0.3–1 by 0.2–0.4 mm, lower surface light green, glabrescent, domatia present or absent, nerves 4–6 per side, ending in the margin. Inflorescences racemes or panicles, axes basally 1–1.5 mm thick; bracts persistent, hairy outside, glabrous to subglabrous inside. Staminate inflorescences up to 15 cm long, flowers 1–3 per node, nodes up to 28; bracts 0.8–1.5 by 0.4–1 mm. Staminate flowers 4–5 mm diam.; pedicels 1.8–5.2 mm long; sepals 3 or 4, ovate to elliptic, 2.5–3.5 by 1.2–2.3 mm; stamens 30–75, glabrous to sparsely hairy, filaments 1–2 mm long, anthers ellipsoid, 0.5–0.8 by 0.3–0.4 mm, light yellow, connective not broad; pistillode absent. Pistillate inflorescences up to 13 cm long, nodes up to 20; bracts 1–2 by 0.5–0.8 mm. Pistillate flowers 2.5–3 mm diam., pedicels 1–4 mm long; sepals 4 or 5, narrowly triangular, 1.8–2.2 by 0.6–0.8 mm, reflexed; ovary (1-or)2-or 3-locular, 1–1.2 by 1.2–1.3 mm; style up to 1.5 mm long; stigmas 1.5–2.5 mm long. Fruits 5–11 by 7–13 mm, opening loculicidally-septicidally, yellowish brown; wall 0.5–1 mm thick, outside grooved or not, shallowly depressed above the septa or not, hairs smooth to slightly rough; column 3–7.5 by 1–2.5 mm. Seeds ± globose, 4–6 by 4–6.2 by 3.5–4.5 mm, surface smooth, glossy, black; hilum 1.5–2 by 1–1.5 mm.

    Distribution — From Nepal to South China and Taiwan, throughout Southeast Asia and Malesia to East Australia and New Caledonia.

    Habitat & Ecology — Locally scattered in the understorey of primary to secondary forests, scrub and disturbed sites; on ridges, forest edges, mangrove swamp edges, road and river sides, steep slopes and dry ground; on various soil types: limestone, granite, sandy loam, and rock. Altitude: sea level up to 1500 m. Flowering and fruiting the whole year through.

    Vernacular names — Malay Peninsula: Ku-ko-mu-ya (Malay). Java: Katjoe-kilang, Merangan, Sindukan, Toekal takal. Philippines: Panuá-lan (Tagálog); Adgao, Ambao, Tapin. Lesser Sunda Islands: Nono nuifmetan (Dawan); Keterakaba, Ikur wase, Rowe. New Guinea: Papua (former Irian Jaya): Toho (Wagu); Wananugapok (Waskuk); Ngontoen.

45. Mallotus resinosus (Blanco) Merr. (sect. Rottleropsis)

    Mallotus resinosus (Blanco) Merr., Spec. Blanc. (1918) 222; Enum. Philipp. Fl. Pl. 2 (1923) 436; Airy Shaw, Kew Bull. 26 (1972) 294; Kew Bull., Addit. Ser. 4 (1975) 161; Kew Bull 35 (1980) 656; Kew Bull. 37 (1982) 29; S.E.C.Sierra et al., Blumea 52 (2007) 88, fig. 9, map 15. — Adelia resinosa Blanco, Fl. Filip., ed. 2 (1845) 562. — Neotype (Balakrishnan & Chakrabarty, 1991: 37): Merrill Species Blancoanae 485 (‘483’) (holo US; iso A, L, NY, P), Philippines, Luzon, Batangas, Mount Batulao.

    [Croton muricatus (‘muricatum’) Wall., Numer. List (1845), nom. nud.]. — Claoxylon muricatum Wall. ex Wight, Icon. Pl. Ind. Or. 5 (1852) 24, t. 1886. — Rottlera muricata (Wight) Thwaites, Enum. Pl. Zeyl. (1861) 273. — Mallotus muricatus (Wight) Müll.Arg., Linnaea 34 (1865) 191. — Mallotus muricatus (Wight) Bedd., Fl. Sylv. 2 (1873) ccviii, pro nov., isonym. — Mallotus muricatus (Wight) Müll.Arg. var. genuinus Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 190, nom. inval. — Coelodiscus muricatus (Wight) Gagnep. in Lecomte, Fl. Indo-Chine 5 (1925) 369, pro comb. — Lectotype (S.E.C.Sierra et al., 2007): Wight KD 2672 (holo K, marked ‘A’; iso A, C n.v., L), India, ‘Courtallum, July 1835, in Herb. Ind. Or. Hook.f.’.

    Mallotus dispar (Blume) Müll.Arg. var. psiloneurus Müll.Arg., Linnaea 34 (1865) 191. — Type: Zollinger 3804 (holo G barcode G00011915; iso G barcode G00011934, K, L), Java.

    Mallotus stenanthus Müll.Arg., Linnaea 34 (1865) 191. — Lectotype (S.E.C.Sierra et al., 2007): Wallich 7751B (holo G-DC n.v., fide IDC microfiche 800; iso K-W n.v., photo seen, IDC microfiche 7394), India, ‘Courtallum, Herb. Madras’.

    Mallotus walkerae Hook.f., Fl. Br. Ind. 5 (1887) 437. — Mallotus muricatus (Wight) Müll.Arg. var. walkerae (Hook.f.) Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 190. — Lectotype (Philcox in Dassan. & W.D.Clayton, Rev. Handb. Fl. Ceylon 11, 1997: 150): Walker s.n. (holo K), Sri Lanka.

    Mallotus walkerae Hook.f. var. laxiflora Hook.f., Fl. Br. Ind. 5 (1887) 437. — Type: Walker 1217 (holo K), Sri Lanka.

    Mallotus spec. nov. 2: Welzen et al., Thai Forest Bull., Bot. 28 (2000) 110. — Mallotus viridis Welzen & Chayam. in Smitinand, Thai Pl. Names, rev. ed. (2001) 343, nomen; Welzen & Chayam., Kew Bull. 56 (2001) 652, f. 2. — Type: Santisuk 6684 (holo BKF), Thailand, South-eastern, Chanthaburi, Soi Dao Nua.

    Mallotus muricatus auct. non Müll.Arg.: Kurz, Forest Fl. Burma 2 (1877) 384, ‘excl. synon.’, fide Hooker f., see next. — Mallotus andamanicus Hook.f., Fl. Br. Ind. 5 (1887) 439. — Lectotype (S.E.C.Sierra et al., 2007): Kurz s.n. (holo CAL n.v.; iso CAL, L, see note 1), S Andaman Islands.

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

Shrubs to small trees, up to 8 m high, dioecious. Indumentum composed of simple and tufted hairs and light yellow to yellow-orange glandular hairs. Stipules triangular to narrowly triangular, 2.3–4.3 by 1–1.7 mm, early caducous, margin subentire, apex acute to rounded, scatteredly hairy outside, glabrous inside. Leaves opposite; petiole 2.5–20(–35) by 0.8–1.3 mm, sparsely to scatteredly hairy; blade elliptic to obovate, 3.5–25 by 1.7–10.7 cm, length/width ratio 1.7–4.3, chartaceous, base obtuse, cuneate to shallowly attenuate, margin denticulate, dentate to crenate, with glandular teeth, apex acute to caudate, upper surface glabrous, usually scatteredly covered with glandular hairs, extrafloral nectaries touching the midrib to marginal in lower half, 1 or 3(–5) per side, 1–3 mm from margin, elliptic, 0.5–3.5 by 0.3–1 mm, lower surface densely (to sparsely) covered with glandular hairs, domatia absent, venation pinnate, nerves 8–12 per side, looping and closed near margin or ending in margin. Inflorescences racemes, single, erect; peduncle 3–7 by 1–1.2 mm; axes densely hairy; bracts ovate or deltoid to triangular, 1.2–2.5 by 1–1.6 mm, persistent or caducous, margin subentire, apex acute to acuminate, sparsely to scatteredly hairy outside, glabrous inside; bracteoles absent. Flowers: pedicels densely to sparsely hairy; sepals persistent, margin entire, apex acute, light brown with glandular hairs outside. Staminate inflorescences 1.7–17.3 cm long, nodes per branch 7–39, with 3–7 flowers per bract. Staminate flowers 2.5–4 mm diam.; pedicels 2–5 mm long; sepals 3 or 4, elliptic to obovate, 1.5–2.3 by 1.2–1.3 mm, free, sparsely hairy outside, glabrous inside; stamens 29–40, filaments 1.5–2.5 mm long, free, glabrous, thecae ellipsoid, 0.2–0.3 by 0.2–0.25 mm, hairy, connnective widened; pistillode present. Pistillate inflorescences 2.1–7.2 cm long, nodes per branch 6–10, with 1 flower per bract. Pistillate flowers 1.5–2 mm diam.; pedicels 0.5–1 mm long; calyx 4–6-lobed, lobes narrowly triangular, 2.7–3 by 0.7–0.8 mm, basally connate, scatteredly hairy to glabrous outside, glabrous inside; ovary 1.3–1.7 by 1.4–1.9 mm, 3-locular, sparsely covered with glandular hairs; style 0.2–0.5 mm long; stigmas 2–3.5 by 1 mm. Fruits capsules, 6–8 by 8–15 mm, opening septicidally-loculicidally, surface spiny, scatteredly hairy to glabrescent, densely to sparsely covered with glandular hairs; spines 40–80, straight to curved (see note), 1–2.2 by 0.3–0.5 mm, scatteredly hairy to glabrous; wall 0.7–1 mm thick, sparsely hairy to glabrous inside; column 4–6 by 4–5 mm. Seeds ± globose, 5–8 by 4–6.5 by 4–6 mm, surface smooth, shiny, brown to brown-grey.

    Distribution — S India, Sri Lanka, Andaman & Nicobar Islands, Thailand, Laos, Vietnam, Philippines, Borneo, Java, Lesser Sunda Islands, Sulawesi, Moluccas, Papua New Guinea, Australia.

    Habitat & Ecology — Locally in understorey of mixed evergreen deciduous, secondary and notophyll vine forest and grassland; near roads, near streams and sea; on clay, granite, loam, limestone, sandstone or volcanic soil. Altitude: sea level up to 1000 m. Flowering and fruiting throughout the year.

    Vernacular names — Philippines: Agudín (Dum), Hangbi táwad (Mango), Tibalbagyon (Tagbanua). Moluccas: O hararoko (Tobelorese). Lesser Sunda Islands: Anak wati, Mboréng.

    Notes — 1. Balakrishnan & Chakrabarty (1991) report the presence of four Kurz collections at CAL, unfortunately without date and sex or fruiting stage. A stamiante and fruiting specimen collected by Kurz is at L and has been appointed as lectotype. Because the material from CAL has not been studied, we do not know exactly how many sheets bearing male specimens there might be. In K there is no accepted specimen labelled as M. andamanicus.

2. Mallotus resinosus has a pronounced midrib on the upper surface of the leaves.

3. See note 3 under M. decipiens.

46. Mallotus rufidulus (Miq.) Müll.Arg. (sect. Rottleropsis)

    Mallotus rufidulus Miq. ex Müll.Arg. in DC., Prodr. 15, 2 (1866) 970; S.E.C.Sierra et al., Blumea 52 (2007) 91, map 6. — [Rottlera rufidula Miq., Fl. Ned. Ind., Eerste bijv. (1861) 181, nom. nud.; 453]. — Type: Diepenhorst HB 2510 (holo U), Sumatra, Priaman.

    Mallotus zollingeri Müll.Arg., Linnaea 34 (1865) 193. — Type: Zollinger 1098* (holo G, barcode G00016017; iso BM, G, P), Java.

    Rottlera dispar auct. non Blume (1826): Moritzi, Syst. Verz. (1846) 17 (nomen); Rchb.f. & Zoll. , Acta Soc. Regiae Sci. Indo- Neerl. 1 (1856) 17 (descr.!). — Mallotus moritzianus Müll.Arg. , Linnaea 34 (1865) 190; Slik, Priyono & Welzen, Gard. Bull. Singapore 52 (2000) 60, Fig. 35; in Slik, Tropenbos-Kalimantan ser. 4 (2001) 98, Fig. 5.38. — Mallotus moritzianus Müll.Arg. var. scaber Müll.Arg., Linnaea 34 (1865) 190, nom. inval. — Rottlera moritziana (Müll.Arg.) Scheff., Ann. Mus. Bot. Lugduno-Batavum 4 (1869) 125. — Lectotype (S.E.C.Sierra et al., 2007): Zollinger 1616 (holo G, barcode G00011931; iso BM, FI, G, P, ZT), Java.

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

Shrubs to small trees, up to 6 m high, dioecious or rarely monoecious. Outer bark smooth. Indumentum composed of simple and stellately-tufted hairs and yellow to orange glandular hairs. Stipules narrowly triangular, 2.8–7 by 0.7–1 mm, early caducous, margin entire, apex acute, sparsely hairy. Leaves opposite; petiole 8–70 by 0.7–1.2 mm, densely to sparsely hairy; blade ovate to elliptic, 8–24 by 3.8–17 cm, length/width ratio 1.6–3, membranaceous to chartaceous, base rounded, obtuse to cuneate, margin entire, rarely dentate, sometimes with glandular teeth, apex caudate, upper surface densely to sparsely hairy on midrib and nerves, extrafloral nectaries marginal throughout blade, up to 22 per side, 2–10 mm from margin, sometimes also all over the blade, elliptic to orbicular, 0.2–0.5 by 0.2–0.5 mm, lower surface sparsely to scatteredly hairy, sparsely covered with glandular hairs, domatia absent, venation triplinerved, nerves 4–6 per side, looping and closed near margin. Inflorescences racemes, single, erect, unisexual or rarely bisexual; axes sparsely hairy; peduncle 20–80 by 1–1.2 mm; bracts triangular, 1–4.2 by 0.7–1.3 mm, persistent to caducous, margin entire, apex acute, densely to sparsely hairy; bracteoles triangular, 0.5–0.7 by 0.2–0.3 mm, early caducous. Flowers: pedicels densely hairy; sepals persistent, margin entire, apex acute, densely hairy and with few glandular hairs outside, glabrous inside. Staminate inflorescences 9–20 cm long, nodes per branch 6–13, with 5–10 flowers per bract. Staminate flowers 3–4.5 mm diam.; pedicels 2.5–5 mm long; sepals 3 (or 4), triangular, 2.9–3.1 by 1.8–2, free; stamens 50–60, glabrous, filaments 1.5–3.2 mm long, free (to slightly connate at the base), thecae ovoid, 0.2–0.3 by 0.2–0.25 mm, connective not widened; pistillode present. Pistillate inflorescences 16–20 cm long, nodes per branch 2–8, with 1 flower per bract. Pistillate flowers 4–5 mm diam.; pedicels 3–8 mm long; calyx 3-lobed, lobes narrowly triangular, 5–6 by 1–1.4 mm, basally connate; ovary 2.5–3 by 2.5–3 mm, 3-locular, densely hairy, glandular hairs scattered; style 0–1(–2) mm long; stigmas 3–4 by 1–1.3 mm. Fruits capsules, 8–20 by 7–19 mm, green, opening septicidally-loculicidally, surface spiny, densely hairy, scatteredly covered with glandular hairs; spines 200–250, ± straight, 2.8–4 by 0.2–0.3 mm, densely hairy; wall 0.8–1.1 mm thick, glabrous inside; column 7–8 by 7–8 mm. Seeds ± globose, 6.5–9 by 6–9 by 6–9 mm, surface smooth, dull, light cream-brown.

    Distribution — Sumatra, Java, Borneo, Lesser Sunda Islands.

    Habitat & Ecology — Riverine forest; on limestone. Altitude 30–750 m. Flowering and fruiting: June to November.

    Vernacular names — Sumatra: Kalikih utan. Java: Kaijae tekeh, Waru rot. Lesser Sunda Islands: Tjempa, Pampa.

    Notes — 1. See note under M. connatus.

2. Kostermans s.n. (L barcode L0294643) has two labels. One is a field label with

the locality ‘Ujung Kulon’ (Java). The other one is a typed label with the locality ‘Berau

near Tgd. Redeb, E Indonesia, Borneo’. We think that the collecting locality is more

likely to be in Java.

47. Mallotus sphaerocarpus (Miq.) Müll.Arg. (sect. Rottleropsis)

    Mallotus sphaerocarpus (Miq.) Müll.Arg. in DC., Prodr. 15, 2 (1866) 976; S.E.C.Sierra et al., Blumea 52 (2007) 93, map 3. — Rottlera sphaerocarpa Miq., Fl. Ned. Ind., Eerste bijv. (1861) 454. — Type: Diepenhorst HB 2119 (holo U; iso G-DC, fragm., fide microfiche IDC 800; BO (‘Teysmann’)), Indonesia, Sumatra, Priaman.

Small to large trees, 8–30 m high, dbh 10–35 cm, dioecious. Indumentum composed of stellately-tufted hairs and yellow to orange glandular hairs. Stipules narrowly triangular, 4.8–7 by 0.8–1.2 mm, early caducous, margin entire, apex acute, densely hairy. Leaves opposite to rarely alternate; petiole 56–80(–130) by 1.8–3 mm, densely to sparsely hairy; blade ovate to elliptic, sometimes slightly falcate, 12–32 by 10–18 cm, length/width ratio 1.4–2, membranaceous to chartaceous, base obtuse to cuneate, margin denticulate to crenulate (rarely dentate), with glandular teeth, apex cuspidate to caudate, upper surface scatteredly hairy to glabrous, larger extrafloral nectaries basal near the petiole insertion, 1–3 per side, elliptic, 0.5–1.5 by 0.2–0.7 mm, smaller ones marginal in upper half, 8–13 nectaries per side, 1–3.5 mm from margin, orbicular, 0.2–0.3 by 0.2–0.3 mm, lower surface sparsely hairy, scatteredly covered with glandular hairs (sometimes only present towards the leaf margin), domatia absent, venation triplinerved, nerves 6–10 per side, looping and closed near margin. Inflorescences racemes, single, erect; axes densely hairy; peduncle 5–10 by 1–2 mm; bracts triangular, 1.5–3 by 0.5–1 mm, persistent to caducous, margin entire, apex caudate, densely hairy; bracteoles absent. Flowers: pedicels densely hairy; sepals persistent, margin entire, apex acute, densely hairy outside, glabrous inside, glandular hairs absent. Staminate inflorescences 16–21 cm long; nodes per branch 63–100, with 6–13 flowers per bract. Staminate flowers 3–3.4 mm diam.; pedicels 2–7 mm long; sepals 3, ovate to elliptic, 2–3 by 1.7–2.1 mm, free; stamens 30–38, glabrous, filaments 0.3–1.8 mm long, flattened, free, thecae ovoid, 0.1–0.2 by 0.1–0.2 mm, connective umbrella-like; pistillode absent. Pistillate inflorescences 9–14 cm long, nodes per branch 12–60, with 1 flower per bract. Pistillate flowers 4–5 mm diam.; pedicels 0.8–1.2 mm long; calyx 3–5-lobed, lobes triangular, 2.5–3 by 1.2–2 mm, basally connate; ovary 2–2.2 by 2–2.2 mm, 3-locular, densely hairy, glandular hairs absent; style 0.4–0.7 mm long; stigmas 1.8–3 by 1–1.4 mm. Fruits indehiscent, spheroid, 15–19 by 15–19 mm, with 3 longitudinal ridges when dry, surface slightly verrucose, warts 20–100, 0.5–1 by 0.2–0.3 mm, densely hairy, glandular hairs rarely present; wall: woody part c. 0.1 mm thick, endocarp fleshy; column absent. Seeds lenticular, 7–10.5 by 6–7.5 by 4–5 mm, surface rugose, dull, black.

    Distribution — Endemic to Sumatra.

    Habitat & Ecology — Locally common in disturbed secondary forest, also found in primary forest; in open areas; on sandy clay soil. Altitude 50–500 m. Flowering: February to March; fruiting: April to October. The fruits are eaten by orang-utans.

    Vernacular names — Sumatra: Pepuah (Gajo); Rumpi rawan.

    Notes — 1. Koop 130 and 200 are atypical in having membranaceous, dentate and

opposite to alternate leaves.

2. See note 1 under M. blumeanus.

48. Mallotus spinifructus Welzen & S.E.C. Sierra (sect. Rottleropsis)

    Mallotus spinifructus Welzen & S.E.C.Sierra, Blumea 51 (2006) 382, fig. 3a–h; map 3. — Type: Leighton 92 (L), Borneo, East Kalimantan, East Kutai Reserve, vicinity of Sengata and Mentoko Rivers, 0°30' N, 117°20' E.

Trees, up to 10 m high, dbh up to 12 cm; flowering branchlets 3–4 mm thick, flattened and somewhat ridged when dry, apical parts stellately hairy, glabrescent; bracts of the terminal buds triangular, c. 1.5 by 0.7 mm. Outer bark grey to greyish pale brown, firm, smooth; inner bark dull brown, coarsely fibrous. Indumentum of simple and stiff, stellately bundled hairs, glandular scales yellowish. Stipules triangular, basally usually quickly decreasing in width, 7–10 by 1–1.7 mm, rather late caducous, outside hairy. Leaves opposite, similar in shape, one somewhat smaller; petiole 1–6 cm long, round, hairy, basally and apically pulvinate, especially upper pulvinus well-developed, oblique with blade base partly overlaying; blade ovate to elliptic, 14.5–24.3 by 5–9.5 cm, length/width ratio 2.6–2.9, symmetric, membranous, base cuneate, very base narrowly cordate, margin entire, apex caudate, upper surface basally with one pair of somewhat elliptic extrafloral nectaries, up to 1.1 by 0.9 mm, in upper third usually several small extrafloral nectaries (mainly 0.3 by 0.3 mm) on the nerves, at most slightly hairy on the midrib; lower surface hairy on venation, glandular scales distinct, hair tuft domatia present; venation triplinerved, flat and somewhat indistinct above, raised and distinct beneath, yellowish, 8 or 9 nerves per side, veins and veinlets scalariform. Inflorescences racemes, axillary to (pseudo)terminal, single, densely hairy; staminate ones only seen in bud, up to 6 cm long, c. 0.8 mm wide, with groups of up to c. 10 buds, pistillate inflorescences up to 7.5 cm long, a single flower per node; bract triangular, 5.5–6 by 1.3–1.8 mm, densely hairy outside, almost floccose, recurved. Staminate flowers only seen in bud, buds globose, pedicelled; pedicel densely hairy; calyx apparently splitting into three sepals, densely hairy outside; stamens perhaps c. 30, anthers separated by connective. Pistillate flowers c. 7.5 mm diam., subsessile when young, in fruit pedicel up to 8? mm long, hairy; sepals 5, triangular to ovate, 4.5–5 by 1.3–2 mm, hairy outside, reflexed; ovary 3- (or 4-)locular, densely long spiny, spines not in groups but pointing all directions, with loose long patent white hairs, glandular scales present; style c. 1.5 mm long, stigmas c. 7 mm long, c. 1 mm wide, with long, branching papillae above, (sub)glabrous beneath. Fruits lobed capsules, 1.8–1.9 by 1–1.1 cm, green when immature, spines up to 4.5 mm long, very slender; wall woody, basally c. 0.7 mm thick, apically c. 1.7 mm thick; column broadly T-shaped to obtriangular, c. 5.5 mm long. Seeds subglobose 6–8 by 5–7 mm, shiny.

    Distribution — Malay Peninsula and Borneo (Central & East Kalimantan).

    Habitat & Ecology — (Logged) riverine forest, alluvial soil, well-drained. Altitude 200–300 m. Flowering: September; fruiting: November to January.

    Vernacular names — Borneo: Kreh merah (Kutai National Park).

49. Mallotus subcuneatus (Gage) Airy Shaw (sect. Rottleropsis)

    Mallotus subcuneatus (Gage) Airy Shaw, Kew Bull. 26 (1972) 304; Whitmore, Tree Fl. Malaya 2 (1973) 115; Welzen et al., Thai Forest Bull., Bot. 28 (2000) 109; S.E.C.Sierra et al., Blumea 52 (2007) 94, map 12. — Coelodiscus subcuneatus Gage, Rec. Bot. Surv. India 9 (1922) 240. — Lectotype (S.E.C.Sierra et al., 2007): Robinson & Kloss 6173 (holo K; iso SING), Peninsular Malaysia, Kedah, Langkawi.

    Mallotus dispar auct. non. (Blume) Müll.Arg.: Airy Shaw, Kew Bull. 21 (1968) 380.

Shrubs to small trees, up to 8 m high, dioecious. Indumentum composed of simple, tufted, stellately-tufted and rarely peltate hairs and yellow-orange glandular hairs. Stipules linear-triangular, 3–4 by 0.4–0.6 mm, early caducous, margin entire, apex acute, densely hairy. Leaves opposite; petiole 30–62 by 1–1.5 mm, densely hairy; blade ovate to elliptic, 7–20 by 4.5–8 cm, length/width ratio 1.5–2.5, membranaceous to chartaceous, base rounded, obtuse to cuneate, margin denticulate, with distinct glandular teeth, apex (acute to) cuspidate, upper surface densely hairy to glabrous on midrib and nerves, extrafloral nectaries marginal in lower half, 1–10 per side, 1–10 mm from margin, elliptic to orbicular, 0.2–0.7 by 0.2–0.3 mm, lower surface sparsely hairy, sparsely covered with glandular hairs, domatia absent, venation triplinerved, nerves 4–6 per side, looping and closed near margin or ending in margin. Inflorescences racemes single, erect; axes densely hairy; bracts narrowly to linear-triangular, 2.5–3 by 0.7–1 mm, persistent, margin entire, apex acute, densely hairy; bracteoles absent. Flowers: pedicels densely hairy; sepals free, persistent, margin entire, apex acute, densely hairy outside, glabrous inside, glandular hairs absent. Staminate inflorescences c. 3.5 cm long, nodes per branch 5–8, with 3–6 flowers per bract. Staminate flowers 4–5 mm diam.; pedicels 1–2 mm long; sepals 3, ovoid, 3.2–3.3 by 1.5–1.7 mm; stamens 30–40, filaments 2.5–4 mm long, basally connate, densely to sparsely hairy, thecae ovoid to ellipsoid, 0.1–0.2 by 0.1–0.15 mm, glabrous, connective widened; pistillode present. Pistillate inflorescences 4–10 cm long, nodes per branch 2–5, with 1 flower per bract. Pistillate flowers 2.4–2.7 mm diam.; pedicels 3–5 mm long; sepals 4 or 5, narrowly triangular, 4.5–5 by 1.8–2 mm; ovary 1.8–2.3 by 1.8–2 mm, 3-locular, densely hairy, glandular hairs absent; style 1–2 mm long; stigmas 1.5–2 by 0.5 mm. Fruits capsules, 8–10 by 13–16 mm, pale green, opening septicidally-loculicidally, surface spiny, densely hairy, scatteredly covered with glandular hairs; spines 100–140, curved, 0.8–1 by c. 0.6 mm; wall c. 0.5 mm thick, sparsely hairy to glabrous inside; column 5–5.5 by 6–8 mm. Seeds ± globose, 5–7 by 6–7 by 6–7 mm, surface smooth, shiny, light brown.

    Distribution — Thailand, Peninsular Malaysia.

    Habitat & Ecology — Locally in understorey of evergreen forest, near waterfalls and streams; on granite soil. Altitude: sea level up to 150 m. Flowering and fruiting: February to May.

    Vernacular names —  Peninsular Malaysia: Tampay, Kayu toon tiew (Temuan).

    Uses — Poisonous fruits.

    Note — Mallotus subcuneatus can easily be confused with M. montanus because of the hairs on the lower leaf surface, hairy filaments, seed shape and colour and their distribution in Thailand and Peninsular Malaysia. The main differences are found in domatia, leaf blade upper surface indument, bracts and fruits: domatia absent (present in M. montanus), upper surface of leaves glabrous (sparsely hairy in M. montanus), glandular teeth inconspicuous (conspicuous in M. montanus), bracts large, c. 3 by 0.7–1 mm (small, 0.6–0.8 by 0.4–0.6 mm in M. montanus); filaments basally connate (free in M. montanus), fruits with spines (absent in M. montanus), seeds large, 5–7 by 6–7 by 6–7 mm, dark brown (small, 4.2–6.5 by 4.2–6.5 by 4.2–6.2 mm, light brown in M. montanus).

50. Mallotus sumatranus (Miq.) Airy Shaw (sect. Polyadenii)

    Mallotus sumatranus (Miq.) Airy Shaw, Kew Bull. 16 (1963) 351; Kew Bull. Add. Ser. 4 (1975) 167; Kew Bull. 36 (1981) 329; Bollendorff, Welzen & Slik, Blumea 45 (2000) 329, fig. 2d, 6; map 1; Slik, Priyono & Welzen, Gard. Bull. Singapore 52 (2000) 63, Fig. 42; in Slik, Tropenbos-Kalimantan ser. 4 (2001) 100, Fig. 5.46; Bollendorff, Welzen & Slik in Slik, Tropenbos-Kalimantan ser. 4 (2001) 140, Fig. 6.2d, 6.10, Map 6.1. — Coccoceras sumatranum Miq., Fl. Ned. Ind., Eerste bijv. (1860) 456; Müll.Arg. in DC., Prodr. 15, 2 (1866) 950; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 209; in Engl. & Harms, Pflanzenfam., ed. 2, 19c (1931) 118. — Type: Teysmann s.n. (U, holo; iso in L), Sumatra, Prov. Lampong, prope Siringkebau.

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

Small tree, up to 12 m high, dioecious; branchlets sometimes lenticellate, glabrescent, finely ribbed, light brown to grey, sometimes mottled with black. Stipules triangular, rarely falcate, 1.5–2.5 by 0.5–1 mm, ridged, caducous, apex acute to acuminate, with some simple hairs. Leaves opposite to rarely alternate, pairs more or less equal in size; petiole terete to flattened, 5–62 by 0.8–2.1 mm, sometimes with transversal cracks, hirsute, sometimes with (stalked) stellate hairs, base and apex not pulvinate, sometimes wrinkled; blade ovate to obovate, rarely narrowly elliptical, 5.5–18.5 by 2–8 cm, index 1.6–4.3, chartaceous to rarely subcoriaceous, base emarginate to cordate, margin slightly to clearly crenate, with 7–15 marginal glands per side in sinuses, rarely pubescent, apex acute to bluntly acuminate, rarely ending in a margin-type of gland, upper surface glabrous or with many stellate hirsute hairs on petiole attachment, pale- to red- to dark brown, basally 1–6 impressed elliptic glands per side, up to 1.5 by 0.8 mm, brown to grey, lowermost pair 0.5–3 mm from petiole attachment, apically 2–4 more circular glands per side, submarginal on veinlets, lower surface red-brown to brown, domatia with long stellate hairs in axils, venation basally triplinerved, basal pair ending in margin below to above middle, 4–11 nerves per leaf side, eucamptodromous, apically occasionally brachidodromous, raised below, veins conspicuously scalariform with 2–3 mm intervals. Inflorescences axillary, staminate inflorescences also terminal; bracts triangular to rhomboid, 1–1.3 by 0.8–2 mm, base sometimes thickened and decurrent, margin not to slightly ciliate, apex acute. Staminate inflorescences 2 or 3 together, 6–16.2 cm by 0.8–1.5 mm, 32–37 nodes per rachis, 3–6 flowers per node; pedicels terete to flattened, 2–5 by 0.1–0.2 mm, abscission zone at the very base, stellate hairs rare, red; flowers 3.5–4 mm in diam., calyx lobes (2 or) 3 or 4, ovate, 1.6–3 by 1.2–1.5 mm, recurving, apex acute, with few villose hairs, reddish brown, stamens 30–42, filaments 1.5–3 mm long, anthers oblong, 0.5–0.8 by c. 0.2 mm, basifixed, reddish brown, connective slender, not split, receptacle 0.2 by 0.5 mm in diam. Pistillate inflorescences solitary, 4.2–19.5 cm by 0.8–1.5 mm, 8–24 nodes per rachis 1 or 2 flowers per node; pedicels terete to flattened, 2–42 by 0.5–0.8 mm, abscission zone 1–2 mm from apex; flowers c. 4 mm in diam., calyx lobes 4–6, connate at base, 2–3 by 0.6–1 mm, basally thickened, reddish brown, apex acute to acuminate, with few villose or long, multicellular, uniseriate hairs, ovary winged, elongated ovoid in lateral view, (2- or) 3- (or 4-)locular, covered with discoid glands, ovules taking up only a third of the wing, style 0.8–2 by c. 0.8 mm, stigmas persistent 2–3.5 by 0.4–1 mm, spreading horizontally, recurved at the apex. Fruits woody, winged, indehiscent to tardily dehiscent capsules, fruit body roughly equilateral triangular from above, each side measuring 6–10 mm, smooth or with some hook-like protuberances, lower fruit body hirsute, wings spreading horizontally, 1.5–3 cm long, basally 2–5 mm wide, often slightly twisted, ribbed lengthwise; locules 3–6 by 3–5 mm wide; column bell-shaped, c. 4 by 3 mm wide at base and 2 mm wide at apex. Seeds subglobose, c. 3 mm in diam., shiny, brown.

    Distribution — Indonesia (Sumatra and Borneo).

    Habitat & Ecology — Primary rain forests on river banks, swamp forests. Alt.: 10–30 m. Flowering: June, Oct.; fruiting: Feb.–Nov.

    Vernacular names — Kalimantan: Belanti, Peroepoek.

51. Mallotus thorelii Gagnep. (sect. Stylanthus)

    Mallotus thorelii Gagnep., Notul. Syst. 4 (1923) 53; in Lecomte Fl. Indo-Chine 5 (1925) 358; Airy Shaw, Kew Bull. 26 (1972) 307; Slik & Welzen, Blumea 46 (2001) 47, Fig. 18, Map 6; in Slik, Tropenbos-Kalimantan ser. 4 (2001) 191, Fig. 7.18, Map 7.6. — Lectotype (Slik & Welzen, Blumea 46, 2001): Thorel s.n. (P holo; P, G iso), Oudan, Me Kong.

Shrubs to small trees up to 10 m tall. Indumentum dense, whitish to yellowish. Branches lenticellate, hairy, glabrescent, gland-dotted. Stipules early caducous (to persistent), (narrowly) triangular to ovate, 2.5–4.5 by 0.9–1.4 mm, margin irregular to entire, apex acute to acuminate, hairy abaxially, glabrous (to sparsely hairy) inside, gland-dotted. Leaves alternate to apically subopposite, peltate, petiole insertion 3–22 mm from base; petiole 15–130 by 0.7–1.7 mm, basally (sometimes also apically) usually slightly thickened to constricted when dried, hairy, glabrescent, gland-dotted; blade ovate to triangular (to orbicular), 3–16 by 2–15 cm, length-width ratio 1–2, base rounded to truncate to slightly emarginate, margin irregularly wavy to slightly dentate, marginal glands 10–20 per leaf side, apex acuminate to cuspidate to aristate, upper surface smooth, sparsely hairy, especially on veins and along margin, glabrescent, densely gland-dotted, basal macular glands 2–6 on veins, 3–18 mm from petiole insertion, apical macular glands 0–3 on veins, 1–3 mm from margin, lower surface smooth, hairy, especially on veins, densely gland-dotted, domatia present often with hair tufts, venation palmate, 8—11 veins originating from petiole insertion, 4–7 secondary veins per side along midrib, ending in the margin. Staminate inflorescences up to 11 cm long, 1–1.3 mm thick at base, usually solitary, hairy, gland-dotted, nodes up to 38 per inflorescence, flowers up to 6 per node; bracts caducous, triangular to ovate, 1.5–2.8 by 0.5–1.2 mm, margin entire to irregular, apex acute to acuminate, hairy abaxially, glabrous (to long simple hairy at base) adaxially, gland-dotted abaxially, rarely adaxially; buds globose with acuminate apex, hairy, gland-dotted. Staminate flowers 3.5 mm in diameter; bracteoles rarely present, ovate to obovate, 0.5–1.2 by 0.2–0.4 mm, margin irregular to entire, apex acute, glabrous to hairy, glabrous adaxially, not gland-dotted; pedicels 1.3–2.3 mm long, hairy, gland-dotted; sepals 4, basally connate, elliptic, recurved, 2–2.5 by 0.8–1 mm, margin entire, apex acuminate, glabrous to hairy, glabrous adaxially, gland-dotted; stamens up to 40; filaments up to 2 mm long, glabrous, gland-dotted or not; anthers 0.3–0.5 by 0.5 mm, basifixed; thecae rarely gland-dotted; connective not widened to widened. Pistillate inflorescences up to 14.5 cm long, basally 0.7–1.3 mm thick, hairy, glabrescent, gland-dotted, nodes up to 18 per inflorescence; bracts early caducous, triangular, 3.5–3.8 by 1.3 mm, margin entire, apex acute, hairy abaxially only, gland-dotted outside only. Pistillate flowers: pedicels 1–1.5 mm long, hairy, gland-dotted; calyx persistent, lobes 2–5, free to more or less connate, ovate to triangular, 1.3–3.5 by 0.7–2.8 mm, margin entire, apex acute (to cuspidate), hairy abaxially only, gland-dotted; ovary 3(–4)-locular, verrucate to shortly echinate, densely woolly hairy, glabrescent, densely gland-dotted, spines up to 1 mm long, densely hairy, gland-dotted, especially basally; style persistent, 0.6–1.5 mm long, hairy, gland-dotted; stigmas 2–4 mm long, densely covered with granulate papillae on the inside, hairy abaxially, gland-dotted outside. Fruit an echinate to verrucose, lobed capsule, 6–8 by 5–6 mm; carpel inside hairy, not gland-dotted; column c. 4 by 2.5 mm. Seeds glossy, roughly globose, 2.5 by 1.8 by 2 mm; hilum elliptic, 0.8 by 0.4 mm.

    Distribution — Southeast Asian mainland (Thailand, Peninsular Malaysia, Cambodia).

  squares = M. thorellii; dots = M. lackeyi.

    Habitat & Ecology — In primary and secondary forest; in scrub, along forest edges and river sides. Altitude: up to 50 m.

52. Mallotus tiliifolius (Blume) Müll.Arg. (sect. Rottleropsis)

    Mallotus tiliifolius (Blume) Müll.Arg., Linnaea 34 (1865) 190; in DC., Prodr. 15, 2 (1866) 969; S.E.C.Sierra et al., Blumea 52 (2007) 96, map 17, plate 2. — Rottlera tiliifolia Blume, Bijdr. (1826) 607 (‘tiliaefolia’); Backer & Bakh.f., Fl. Java 1 (1964) 484; Airy Shaw, Kew Bull. 26 (1972) 305; Kew Bull., Addit. Ser. 4 (1975) 170; Kew Bull., Addit. Ser. 8 (1980) 171;  Kew Bull. 36 (1981) 329; Kew Bull. 37 (1982) 30; Alphab. Enum. Euph. Philipp. Isl. (1983) 38; P.I.Forst., Austrobaileya 5 (1999) 489; Welzen et al., Thai Forest Bull., Bot. 28 (2000) 110. — Rottlera blumei Decne., Nouv. Ann. Mus. Hist. Nat. 3, 3 (1834) 486, nom. superfl. — Lectotype (S.E.C.Sierra et al., 2007): Blume ‘1676’ (holo L barcode L0294967; iso L barcode L0292539, BO), Java.

    Halecus litorea Rumph., Herb. Amboin. 3 (1743) 196, t. 126, nom. inval.; Merr. (1917) 318. — Croton tiliifolius Lam. var. aromaticus auct. non (L.) Lam.: Lam., Encyl. Meth. Bot. 2 (1786) 206, quoad syn. Rumph.; Müll.Arg. in DC., Prodr. 15, 2 (1866) 969 (see note). — Voucher: Rumphius plate 126. — Representative specimen: Robinson 367 (holo US; iso BM, L, NY), Moluccas, Ambon, Amboina.

    Croton acuminatus Lam., Encyl. Meth. Bot. 2 (1786) 207 (‘acuminatum’), non M. acuminatus Müll.Arg. (1865). — Rottlera acuminata (Lam.) A.Juss., Euphorb. Gen. (1824) 33. — Type: Commerson s.n. (holo P-JU # 16580 (IDC microfiche 6206)), Papua New Guinea, New Britain, Port Praslin.

    Adelia papillaris Blanco, Fl. Filip., ed. 2 (1845) 562. — Mallotus papillaris (Blanco) Merr., Philipp. J. Sci., Bot. 7 (1912) 238; Enum. Philipp. Fl. Pl. 2 (1923) 434. — Mallotus zollingeri auct. non Müll.Arg.: Fern.-Vill. in Blanco, Fl. Filip., ed. 3 (1880) 195. — Neotype (selected here): Merrill Species Blancoanae 50 (holo US; iso A, BM, F, GH, L, NY, P), Philippines, Batangas, San Pedro.

    Mallotus playfairii Hemsl. in F.B. Forbes & Hemsl. J. Linn. Soc., Bot. 26 (1894) 441. — Type: Playfair 324 (holo K; photo in A), Taiwan, Ape Hill.

    Croton enantiophyllus K.Schum. in K.Schum. & Lauterb., Nachtr. Fl. Deutsch. Schutzgeb. Südsee (1905) 296. — Type: Parkinson s.n. (holo B, †; iso NSW barcode 604690, male specimen), Papua New Guinea, New Britain.

    Mallotus palauensis Hosok., Trans. Nat. Hist. Soc. Formosa 25 (1935) 25. — Type: Hosokawa 7284 (holo TAI; iso L), Republic of Palau, Babeldaob Island.

    Mallotus ponapensis Hosok., Trans. Nat. Hist. Soc. Formosa 25 (1935) 26. — Type: Hosokawa 6124 (holo TAI, iso TAI), Federated States of Micronesia, Pohnpei, Saba-tic.

    Mallotus repandus auct. non (Rottler) Müll.Arg., Linnaea 34 (1865): K.Schum. & Lauterb., Fl. Deutsch. Schutzgeb. Südsee (1901) 395. — Voucher: Naumann s.n. (G?, K?, LE? n.v.), Papua New Guinea, Neu-Hannover.

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

Shrubs to small trees, up to 15 m high, dioecious or rarely monoecious. Outer bark smooth and shallow longitudinal fissured. Indumentum tomentose, composed of simple, stellately-tufted and stellate hairs (on petiole and midrib of blade) and yellow to orange hairs (capitate glandular hairs are also present but not easy to detect). Stipules absent. Leaves opposite to rarely alternate, smelling of skunk when fresh; petiole 20–65 by 1–2 mm, densely hairy; blade broadly ovate to ovate, 4.5–21 by 3.8–17.5 cm, length/width ratio 1–1.3, chartaceous, base cordate, truncate, rounded to cuneate, margin entire to denticulate, sometimes with glandular teeth, apex acute to caudate, upper surface sparsely hairy to scatteredly hairy, sometimes scatteredly covered with glandular hairs, extrafloral nectaries marginal from base (to apex), 2–8 per side, 0.3–2 mm from margin, elliptic to orbicular, 0.3–0.8 by 0.3–0.5 mm, lower surface densely hairy, densely (to scatteredly) covered with glandular hairs, domatia absent, venation triplinerved, nerves 7–11 per side, looping and closed near margin or ending in margin. Inflorescences racemes, single, erect, unisexual or rarely bisexual; axes densely hairy; bracts triangular to narrowly triangular, 0.8–3 by 0.5–1 mm, margin entire, apex acute, densely hairy; bracteoles absent. Flowers: pedicels densely hairy; sepals margin entire, apex acute, densely hairy and with glandular hairs outside, sparsely hairy to glabrous inside. Staminate inflorescences 7–22 cm long; peduncle 30–40 mm long; nodes per branch 12–25, with 5–12 flowers per bract. Staminate flowers 2.8–5.5 mm diam.; pedicels 2–4 mm long; sepals 4 or 5, ellipsoid to ovoid, 2.5–3 by 0.9–1.5 mm; stamens 45–70, glabrous, filaments 0.5–2.3 mm long, free (rarely aggregated into a cluster), thecae ellipsoid, 0.2–0.3 by 0.2–0.25 mm, connective widened; pistillode present. Pistillate inflorescences 5–15 cm long; peduncle 10–40 mm long; nodes per branch 9–16, with 1 flower per bract. Pistillate flowers 2–4 mm diam.; pedicels 1.7–2.5 mm long; calyx 4–6-lobed, lobes narrowly triangular, 2–2.3 by 0.5–0.7 mm; staminodes rarely present (usually persistent in fruits); ovary 1.3–1.5 by 1.2–1.4 mm, 3- (or 4-)locular, densely hairy, sparsely covered with glandular hairs; style 0.4–0.7 mm long; stigmas 2.5–4.3 by 0.8–1 mm. Fruits capsules, 5–15 by 7–15 mm, opening septicidally-loculicidally, surface spiny, densely hairy, sparsely covered with glandular hairs; spines 15–60, straight, 0.5–2 by 0.2–0.3 mm, densely hairy; wall 0.8–1 mm thick, sparsely hairy inside; column 4–6 by 3–4 mm. Seeds globose, 5–7 by 5–7 by 4.7–5 mm, surface smooth, dull, light brown-orange.

    Distribution — From Thailand to Taiwan, throughout Peninsular Malaysia, Malesia, Republic of Palau, and Federated States of Micronesia to the W Pacific (Solomon Islands, Fiji).

    Habitat & Ecology — Primary forest, understorey of evergreen to deciduous forest, coastal or disturbed riparian vegetations; on edges of mangroves, swampy areas, sandy areas behind the beach, along forest margins, trails, steep valley slopes, lake and sea shores, the end of tidal marsh; in wet terrains. Altitude: sea level up to 350 m. Flowering and fruiting throughout the year.

    Vernacular names — Java: Djasilin, Ring rin, Waru lot. Brunei: Tapian batu. Malaysia/Sabah: Tuka tuak laut. Philippines: Anamú saláurid, Gapas gapas. Moluccas: War-war. Sulawesi: Lendé. New Guinea: Maimeireh (Gabobora); Idirur (Onjob).

    Note — Lamarck (1786) described Croton tiliifolius with a variety ß aromaticus. As it turns out Croton tiliifolius consists of three taxa. The typical variety is based on a Commerson collection from Mauritius, which is a true Croton (see Coode, 1982: 44). The variety ß consists of two parts, based on references to Linnaeus’ Fl. Zeylanica (1747) and Rumphius’ Herb. Amboin. 3 (1743). The first part, “Croton foliis cordatis scabris subserratis petiolatis caule arboreo” from Sri Lanka, was used by Linnaeus in 1753 for his Croton aromaticus, which is also still regarded as a Croton (Philcox, 1997: 101). The second part, which has caused the most confusion, is the Halecus litorea of Rumphius from Ambon. It is a Mallotus as plate 126 in Rumphius’ book shows. This was also observed by Merrill (1917: 318), who gave a brief survey on the identification of the Rumphian species.

Another cause of the confusion is that Blume described a Rottlera tiliifolia (1826) which has been equated with Rumphius’ species and probably because of the similarity of the epithet with Lamarck’s species. An example of this is found in Müller Argoviensis (1865) where he made the combination Mallotus tiliifolius. The first reference given is to Lamarck’s name, the second to the one of Blume and the third to Rottlera acuminata A.Juss. (from Timor and a synonym of Blume’s combination). It would therefore appear that the combination is based on the Mauritian name. However, this is not so, as is evident from the treatment in 1866, where Müller clearly distinguished between Croton tiliifolius Lam. (p. 587), Croton aromaticum L. (p. 588) and Mallotus tiliifolius Müll.Arg. (p. 969). In the publications of 1865 and 1866 the last taxon has exclusively Southeast Asian and Chinese collections and what obviously happened was that the letter ß after Croton tiliifolius Lam. was accidentally omitted in 1865. The distinction between Croton tiliifolius and Rottlera tiliifolius was also made by Decaisne (1834) who erroneously renamed Blume’s taxon to Rottlera blumei. Blume’s epithet tiliifolius is in fact younger (1826) than the epithet acuminatus (De Jussieu’s name is based on Croton acuminatum Lam., 1786). However, the combination Mallotus acuminatus was already in use (also by Müller Argoviensis) and, therefore, the epithet tiliifolius is then the oldest available. The combination proposed is based on that of Blume and the correct name for the taxon therefore must be Mallotus tiliifolius (Blume, and not Lam.!) Müll.Arg.

53. Mallotus trinervius (K. Schum. & Lauterb.) Pax & K.Hoffm. (sect. Rottleropsis)

    Mallotus trinervius (K. Schum. & Lauterb.) Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 156; S.E.C.Sierra et al., Blumea 52 (2007) 99, map 2. — Syndyophyllum trinervium K. Schum. & Lauterb., Fl. Deutsch. Schutzgeb. Südsee (1901) 405. — Coelodiscus trinervius (K. Schum. & Lauterb.) Pax, Feddes Repert. Spec. Nov. Regni Veg. 8 (1910) 481; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iii (1911) 105. — Type: Tappenbeck 84 (holo B, †), New Guinea, Kaiser-Wilhelms-Land, Ramufluss.

Small trees, 6–19 m high; bole up to 11 m high, dbh 18–40 cm, dioecious, crown up to 8 m high. Outer bark shallowly longitudinally fissured. Indumentum composed of simple, stellately-tufted and stellate hairs (on petiole and midrib of blade) and yellow glandular hairs. Stipules triangular to narrowly triangular, 1.8–2 by 0.4–0.6 mm, early caducous, margin entire, apex acute, densely hairy outside, sparsely hairy inside. Leaves opposite; petiole 30–90 by 1.8–2.5 mm, sparsely hairy; blade ovate to elliptic, 9–26 by 6–15.5 cm, length/width ratio 1.4–1.8, chartaceous, base rounded, obtuse to shortly attenuate, margin entire, apex obtuse, acute to acuminate, upper surface sparsely to scatteredly hairy on midrib, extrafloral nectaries marginal from base to apex, 7–25 per side, 0.5–6 mm from margin, elliptic to orbicular, 0.2–0.6 by 0.3–0.6 mm, lower surface densely (see note) to sparsely hairy, glabrescent, sometimes scatteredly covered with glandular hairs, domatia absent, venation triplinerved, nerves 5–7 per side, looping and closed near margin. Staminate inflorescences racemes, 9–17 cm long, single, erect; axes densely hairy; peduncle 8–29 by 1–1.5 mm; nodes per branch 16–20, with 5–7 flowers per bract; bracts triangular, 0.9–1.1 by 0.7–0.9 mm, margin entire, apex acute, densely hairy outside, glabrous inside, early caducous; bracteoles absent. Staminate flowers 5–6 mm diam.; pedicels 2–3.5 mm, densely hairy; calyx 3–5-lobed, lobes ovate to elliptic, 3.8–4.8 by 1.5–2.5 mm, persistent, margin entire, apex acute, densely hairy outside, glabrous inside, without glandular hairs; stamens 100–120, glabrous, filaments 2.5–5 mm long, free, thecae ellipsoid, 0.2–0.3 by 0.2–0.25 mm, connective widened; pistillode present. Pistillate inflorescences and pistillate flowers unknown; infructescences 4–6 cm long; peduncle 3–55 by 1–2.2 mm long; nodes per branch 7–19, with 1 flower per bract; sepals 4–6, narrowly triangular, 2.4–3 by 0.8–1.1 mm; ovary 2- or 3- (or 4-)locular, densely hairy. Fruits tardily dehiscent capsules, 17–22.5 by 24–29 mm, opening loculicidally, surface smooth, densely hairy, glandular hairs absent; wall 3–8 mm thick, sparsely hairy to glabrous inside; column 10–15 by 2.8–3 mm. Seeds ± globose, 9–10 by 8.5–9.5 by 7–8.5 mm, surface rugose, dull, whitish cream.

    Distribution — Endemic to New Guinea.

    Habitat & Ecology — In primary or secondary forest, occasionally in flooded rainforests of river plains, in swamp-forests or swamp-grasslands; on alluvial or peaty soil. Altitude: sea level up to 1400 m (Airy Shaw 1980c: 172, mentions 810–3450 m).

    Note — See note 2 under M. didymochryseus.

54. Mallotus wrayi King ex Hook.f.  (sect. Rottleropsis)

    Mallotus wrayi King ex Hook.f., Fl. Br. India 5 (1887) 433; Ridl., Fl. Malay Penins. 3 (1924) 287; M.R.Hend., J. Malayan Branch Roy. Asiat. Soc. 17 (1939) 71; Airy Shaw, Kew Bull. 14 (1960) 361; Kew Bull. 16 (1963) 349, p.p. excl. M. caudatus; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 52, p.p. excl. M. caudatus; Whitmore, Tree Fl. Malaya 2 (1973) 115, p.p. excl. M. lancifolius; Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 171, p.p. excl. M. caudatus; Kew Bull. 36 (1981) 330; Slik, Priyono & Welzen, Gard. Bull. Singapore 52 (2000) 64, f. 43; Tropenbos-Kalimantan Ser. 4 (2001) 100, f. 5.47; Welzen & S.E.C.Sierrra, Blumea 51 (2006) 384, fig. 1a; map 4. — Lectotype (Welzen & Sierra, 2006): King’s collector 3456 (holo K; iso BM, G, SING), Malaya, Perak, at Larut.

    Kunstlerodendron cuspidata Ridl., Fl. Malay Penins. 3 (1924) 283. — Type: Ridley 7911 (holo K? n.v.; iso BM, SING), Malaya, Telok Sera, Dindings, February 1896.

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

Shrubs to trees, up to 8(–23) m high, clear bole up to 15 m high, dbh up to 10(–40) cm; flowering branchlets 2–4 mm thick, flattened and somewhat ridged when dry, (sub)glabrous; bracts of the terminal buds up to 4.5 mm long. Outer bark whitish to grey to green to brownish, smooth; inner bark whitish to yellowish to yellowish brown to pink to brownish red or orange-brown; sapwood white to yellowish to pale orange to pale brown. Indumentum mainly absent, lower leaf blade surface with very thin, translucent white stellate hairs with only horizontal branches; inflorescences with curly brownish simple to stellately grouped hairs; glandular scales few, yellowish to orangey to dark red. Stipules triangular, c. 8 by 2 mm, very early caducous. Leaves opposite, similar shape, one usually somewhat smaller; petiole 0.7–10(–32) cm long, round to adaxially slightly flattened in transverse section, (sub)glabrous, basally and especially apically strongly pulvinate, upper pulvinus oblique; blade partly over upper pulvinus (pseudo-subpeltate), ovate to elliptic (to slightly obovate), 4.5–27 by 1.8–11 cm, length/width ratio 2.3–5, symmetric, membranous to subcoriaceous; base mainly broadly cuneate, very base narrowly cordate, margin entire, usually somewhat undulate, apex cuspidate to caudate; upper surface glabrous, dark green, basally one pair of extrafloral nectaries on basal nerves, elliptic, 0.3–1.3 by 0.2–0.8 mm, sometimes an additional, smaller one along the midrib in basal third, in upper third small extrafloral nectaries on the nerves, circular, c. 0.3 by 0.3 mm; lower surface smooth, somewhat bluish, venation glabrous, in between veins subglabrous to densely hairy (then with silvery sheen), hairs flat against surface, hair tuft domatia present, few glandular scales; venation triplinerved, slightly raised above, raised beneath, nerves 6–9 per side up to the apex, looped and closed near margin, veins scalariform, veinlets laxly reticulate to almost scalariform. Inflorescences racemes, axillary to terminal, single to seven together when staminate, greenish brown, hairs curly, patent, ferruginous, glabrescent; staminate ones up to 5(–11) cm long, 0.4–1.2 mm thick, with cymosely flowering groups of up to 5 flowers per node, pistillate ones up to 4(–11) cm long, 0.9–2 mm thick, with few flowers, one per node; bracts ovate, 0.7–2.8(–3.3) by 0.7–1 mm, outside hairy, inside glabrous. Staminate flowers 3–4.7 mm diam., whitish to yellowish; buds round to ovoid (see note); pedicel 2–4(–7.8) mm long, hairy; sepals 3 or 4, ovate to elliptic, 2.3–3.2 by 1.2–2 mm, reflexed, hairy, especially outside, glandular scales only outside; stamens 25–40 (high numbers in Malay Peninsula), filaments 0.8–2.2 mm long, glabrous, anthers 0.3–0.4(–0.7) by c. 0.3(–0.7) mm (see note), thecae slightly to very distinctly separated from each other (sometimes in same flower). Pistillate flowers c. 3.7 mm diam., green; pedicel up to 4 mm long with an abscission zone in basal half; sepals 3, finally splitting into 5 parts, ovate, 3.5–4 by 0.8–2 mm, yellowish green, mainly hairy outside, glandular scales only outside; ovary (2- or) 3-locular, c. 2 by 2 mm, spines up to 1 mm long, pointing in all directions (not grouped in rows), moderate to dense, purple, mainly glabrous (base excepted); style c. 1.2 mm long, stigmas up to 7 mm long, 0.2–0.7 mm wide, densely papillate above with branching papillae of usually less than 1 mm long, narrow strip of hairs on abaxial side (indistinct by the naked eye). Fruits lobed capsules, 13–23 by 6–12 mm, yellow to light orange to purple-orange turning brown to brownish grey, spines up to 5.5 mm long, breaking off; wall woody, inner part 1–1.5 mm thick; column T-shaped, up to 6 mm high. Seeds subglobose, 7–9 by 6.5–9 by 6–8.5 mm, brown, shiny.

    Distribution — Malay Peninsula, Sumatra (Riau), Borneo.

    Habitat & Ecology — Primary (dipterocarp) forest, pole dipterocarp forest, swamp forest, felled forest, secondary forest, hill side, valley side, along rivers; soil brown, yellow (sandy) or red clay, yellow sandy loam; sedimentary rock, basalt, granite derived soil. Altitude 10–1000 m. Flowering: February to April, June to October; fruiting: February, April to July, September to December.

    Vernacular names — Malay Peninsula: Balek angin. Borneo: Brunei: Luta. West Kalimantan: Krotis, Mpore, Sampa. Sabah: Mangki. Sarawak: Bantas.

    Notes — 1. BRUN (Ahmad et al.) 15713 from Brunei is not only exceptional in its extremely long leaves, but more so in its large ovoid staminate buds with very big anthers (c. 0.7 by 0.7 mm instead of c. 0.3 by 0.3 mm).

2. The leaf size and shape is rather variable. Usually the leaves are large, but especially in Sarawak and West Kalimantan many specimens with small, more slender, almost rheophytic leaves are found. These specimens also tend to have somewhat smaller fruits. Specimens show a gradual transition from small and narrow to large and relatively broad. Moreover, there is also no geographical cline as small-leaved specimens are also present in Sabah and large-leaved ones in Sarawak. Therefore, no taxonomic significance is given to the smaller-leaved form.

3. Fruits on Borneo are usually less hairy than those on the Malay Peninsula.

4. The leaves usually only have one basal pair of extrafloral nectaries, sometimes an additional smaller pair is found slightly above the base along the midrib. Ambriansyah AA 774 (East Kalimantan, Long Sungai Barang) is exceptional in having leaves with often many smaller extrafloral nectaries, a situation usually found in Mallotus caudatus.

Notes on Unidentified Specimens

1. Brass 1333, New Guinea, Murua River.

The specimen looks like M. attenuatus in the presence of inconspicuous extrafloral

nectaries and absence of glandular hairs. However, the leaves are triplinerved, while

in M. attenuatus they are pinnate.