Malesian Euphorbiaceae Descriptions

43. ENDOSPERMUM (Euphorbiaceae)

Arias Guerrero & van Welzen, P.C. 2011. Revision of Malesian Endospermum (Euphorbiaceae) with notes on phylogeny and historical biogeography. Edinburgh J. Bot. 68: 443–482.

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Genus description

Species descriptions

Excluded names

Endospermum Benth.

    Endospermum Benth., Fl. Hongkong. (1861) 304, nom. cons., non Blume (‘Endespermum’, 1823 = Dalbergia, Papilionaceae), nom. rej.; Müll.Arg. in A.DC., Prodr. 15, 2 (1866) 1131; Benth. in Benth. & Hook.f., Gen. Pl. 3 (1880) 322; Hook.f., Fl. Brit. India 5 (1887) 458; Pax in Engl. & Prantl, Pflanzenfam. 3, 5 (1890) 90; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iv (1912) 33; Ridl., Fl. Malay Penins. 3 (1924) 305; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1926) 452; L.S.Sm., Proc. Roy. Soc. Queensl. 58 (1947) 52; J.Schaeff., Blumea 19 (1971):177; Airy Shaw, Kew Bull. 26 (1972) 258; Whitmore, Tree Fl. Malaya 2 (1973) 93; Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 109; Kew Bull., Addit. Ser. 8 (1980) 78; Kew Bull. 36 (1981) 293; Kew Bull. 37 (1982) 18; Alphab. Enum. Euphorb. Philipp. Isl. (1983) 24; G.L.Webster, Ann. Missouri Bot. Garden 81 (1994) 101; Radcl.-Sm., Gen. Euphorbiacearum (2001) 281; Chantharaprasong in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 255; S.Arias Guerrero & Welzen, Edinburgh J. Bot. 68 (2011) 460; G.L.Webster in Kubitzki, Fam. Gen. Vasc. Pl. 11 (2014) 160. — Endospermum Subgen. Euendospermum Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iv (1912) 34, nom. inval.; L.S.Sm., Proc. Roy. Soc. Queensl. 58 (1947) 53. — Type: Endospermum chinense Benth.

    Capellenia Teijsm. & Binn., Natuurk. Tijdschr. Ned.-Indië 29 (1867) 238, Fig. — Endospermum Benth. Subgen. Capellenia (Teijsm. & Binn.) Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iv (1912) 36; L.S.Sm., Proc. Roy. Soc. Queensl. 58 (1947) 52. — Type: Capellenia moluccanaTeijsm. & Binn. [= E. moluccanum (Teijsm. & Binn.) Kurz].

Trees, dioecious, rarely monoecious; branches with pith or hollow; when hollow then with pores and inhabited by ants. Indumentum consisting of simple hairs, solitary and/or stellately bundled. Stipules triangular, caducous. Leaves spirally arranged, present at the top of the branches, simple; petiole long, in transverse section channelled above at the base and/or the apex, otherwise round; blade (orbicular to) ovate to obovate, symmetric, usually coriaceous, rarely papyraceous, base peltate or not, very often with 1 or 2 large extrafloral nectaries near the insertion, margin entire or somewhat undulate, often with minute glands, lower surface often with glands or hair tuft domatia in the first and second bifurcations of the nerves; venation indistinct above, distinct underneath, basally triplinerved or palmate, upper part pinnate, veins scalariform, veinlets reticulate. Inflorescences axillary (thyrsoid?) panicles or racemes; bracts triangular, apex acute to acuminate. Flowers regular, unisexual, exceptionally bisexual (only some specimens in one species); sessile or with a short pedicel; calyx lobes basally or completely connate; petals absent; disc annular, indisctinct but perhaps toothed and up the androphore in E. diadenum. Staminate flowers in groups of up to 3 flowers per node; calyx cupular, indistinctly lobed or 4-lobed, rarely 4-toothed; stamens 5–12, androphore cylindrical or obovate, filaments short, anthers dorsifixed, 3- or 4-thecate, thecae separate from each other, reniform, opening extrorse with lengthwise slits; pistillode on top of androphore, knob-like or cylindrical. Pistillate flowers single per node, rarely in groups of up to 3; calyx indistinct or 4- or 5-lobed or toothed; ovary globose, 1-6- (or 7)-locular, densely covered with minute hairs; ovules 1 per locule; style absent, stigma discoid or lobed. Fruits indehiscent, baccate, rugose, stigma persistent, columella absent. Seeds ellipsoid or globose, ecarunculate or with a slender caruncule, ribbed.

    Distribution — The genus comprises ten species with a palaeotropical distribution, mainly ranging from Southeast Asia main land throughout the Malesian region (except for Java and the Lesser Sunda Islands), North Queensland (Australia) to the west Pacific (Solomon Islands, Vanuatu and Fiji Islands). In the Malesian region eight species are recognized. 

    Note — The genus description is based on the eight species that occur in the Malesian region.

Key to the Malesian species

1. Endospermum diadenum (Miq.) Airy Shaw

   Endospermum diadenum (Miq.) Airy Shaw, Kew Bull. 14 (1960) 395; J.Schaeff., Blumea 19 (1971) 186, map 2; Airy Shaw, Kew Bull. 26 (1972) 258; Kew Bull., Addit. Ser. 4 (1975) 109; Kew Bull. 36 (1981) 293; Chantharaprasong in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 256, plate XV: 3; S.Arias Guerrero & Welzen, Edinburgh J. Bot. 68 (2011) 464, fig. 5, 6. — Melanolepis ? diadena Miq., Fl. Ned. Ind. Eerste Bijv. (1860) 455. — Mallotus diadenus (Miq.) Müll.Arg. in A.DC., Prodr. 15, 2 (1866) 959. — Rottlera diadena (Miq.) Scheff., Ann. Mus. Bot. Lugd.-Bat. 4 (1869) 125. — Type: Teijsmann HB 3807 (holo L), Indonesia, Sumatra, Lubu-alang.

   Endospermum borneense Benth. ex Müll.Arg., Flora 47 (1864) 469; Müll.Arg. in A.DC., Prodr. 15, 2 (1866) 1132; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iv (1912) 35; Merr., J. Straits Branch Roy. Asiat. Soc., Special no. (1921) 346; Corner, Gard. Bull. Straits Settlem. 10 (1939) 298. — Type: J. Motley 1126 (holo K), Indonesia, Borneo, prope Banjarmassing.

   Endospermum malaccense Benth. ex Müll.Arg., Flora 47 (1864) 469; in A.DC., Prodr. 15, 2 (1866) 1132; Hook.f., Fl. Brit. India 5 (1887) 458; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iv (1912) 34; Ridl., Fl. Malay Penins. 3 (1924) 305; K.Heyne, Nutt. Pl. Ned.-Ind. 2^nd ed., 2 (1927) 956; Corner, Gard. Bull. Straits Settlem. 10 (1939) 296; Whitmore, Tree Fl. Malaya 2: 93 (1973) Fig. 7. — Type: Griffith KD 4721 (holo K; iso P), Peninsular Malaysia.

   Endospermum ovalifolium Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iv (1912) 34; Ridl., Fl. Malay Penin. 3 (1924) 305; K.Heyne, Nutt. Pl. Ned.-Ind. 2^nd ed., 2 (1927) 956; Corner, Gard. Bull. Straits Settlem. 10 (1939) 297. — Type: Unknown s.n. (holo B, lost), Singapore, Bukit Timah. Synonymy follows Schaeffer (1971).

   Endospermum borneense auct. non Benth. ex Müll.Arg.: Becc., Malesia 2 (1884) 45. — Endospermum beccarianum Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iv (1912) 35; Merr., J. Straits Branch Roy. Asiat. Soc., Special no. (1921) 346; Corner, Gard. Bull. Straits Settlem. 10 (1939) 298. — Endospermum beccarianum Pax & K.Hoffm. var. crassirameum Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iv (1912) 35, nom. inval. (should have been autonym). — Type: Beccari PB 3137 (holo B, lost; iso Fi, n.v., P), Malaysia, Sarawak. See note 1.

   Endospermum beccarianum Pax & K.Hoffm. var. tenuirameum Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iv (1912) 35. — Type: Beccari PB 819 (holo B, lost; iso FI, n.v., NY, P), Malaysia, Sarawak.

   Endospermum chinense Benth. var. malayanum Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iv (1912) 36. — Endospermum malayanum (Pax & K.Hoffm.) Chatterjee, Kew Bull. 1949 (1950) 564. — Type: Forbes 2779 (holo B, lost; iso BM, L, P), Indonesia, Sumatra, Palembang.

Trees, dioecious, up to 36 m high, bole straight up to 25 m high, diameter 13–60 cm; without or with buttresses up to 2 m high; branches with a hard pith, flowering ones 6–8 mm thick, pilose, containing white to yellow latex. Outer bark grey to green to red to brown, smooth, hooped and/or with white lenticels; inner bark pale yellow to orange to brown to grey, sap clear; sapwood white to yellow; heartwood yellow to light brown. Indumentum consisting of stellately bundled and simple hairs. Stipules 3.1–5 by 1.2–2 mm, densely hairy on both sides. Leaves: petiole 5.5–15 cm long, channelled above at the base, otherwise round; blade ovate to obovate, 10.5–23.5 by 7.7–19 cm, length/width ratio 1.2–1.4, coriaceous, rarely papyraceous, base never peltate, (cordate to) truncate to cuneate, abaxially with or without 1 or 2 knob-like, green extrafloral nectaries near the insertion, 0.4–0.5 by 1.5–2.2 mm, basally without or rarely with a rim around the glandular part, margin entire or somewhat undulate, with numerous minute glands, apex round to acuminate, upper surface glabrous to sparsely covered with minute stellately bundled hairs and/or simple hairs, dark green, glossy, lower surface sparsely to densely covered with minute stellately bundled hairs and/or simple hairs (see notes 2 and 3), pale green to yellow and often with glands at the more marginal bifurcations of the nerves, hair tuft domatia lacking; venation basally triplinerved or palmate with 7 nerves of which the 2 outer ones less distinct, upper part pinnate with 4 or 5 pairs of nerves. Inflorescences racemes, pilose, up to 20 cm long; bracts triangular, 1.4–2.5 by 0.8–2.2 mm, apex acute, densely hairy on both sides, glandless. Staminate flowers white to yellow to green, sessile or with a pedicel up to 0.6 mm long; calyx indistinctly lobed or 4-toothed, 0.7–1.4 by 1.2–1.6 mm, pilose outside, glabrous inside, pale green; stamens c. 9, yellow to ochre, dull; androphore cylindrical, 1.4–2.4 by 0.4–0.8 mm, free part of filaments 0.4–0.7 mm long; anthers 4-thecate, 0.4–0.6 by 0.5–0.6 mm; pistillode knob-like or split in 2 parts, c. 0.2 by 0.2 mm, pilose or sparsely hairy on top. Pistillate flowers single or in groups of up to 3, greenish to yellow, with a pedicel up to 3.1 mm long; calyx indistinctly lobed or 5-toothed, 1.2–2.3 by 1.8–2.6 mm, densely hairy outside, pilose inside; ovary cylindrical, more often 2- than 3-locular, 1.4–2.4 by 1.1–1.7 mm; stigma indistinctly lobed, flattened, 0.2–0.6 by 1.2 —1.8, glabrous, stigma discoid, 0.9–2 mm wide. Fruits blue to green to yellow, globose, 4.8–7 by 5.2–9 mm, pilose. Seeds globose, 3.8–4.2 by 3.1–4.1 mm, white, ecarunculate.

   Distribution — Thailand, Peninsular Malaysia, Sumatra and Borneo.

   Habitat & Ecology — In primary, secondary, swampy, mossy, lowland mixed dipterocarp forests, kerangas forests and disturbed (logged) areas; along rivers and roads, open spaces, on hills, ridges and steep slopes. An opportunist species which grows well and is characteristic of gaps in the high forest, hence common in areas of old shifting cultivation (Whitmore, 1973). Soil: granitic sand, sandy soil, sandy loam, sandy clay, yellow and red clay soil and ultramafic soil. Altitude: up to 1375 m. Flowering and fruiting throughout the year; ripe fruits eaten by birds.

   Uses — The wood used for construction (Kalimantan, Borneo), though the timber is in general soft, light, and suitable for match boxes, splints, drawing boards, blackboards, toys (Whitmore, 1973), planks and clogs (Burkill, 1935).

   Vernacular names — Malay peninsula: Bebaru bukit, Kuyu sendok, Membulan, Poko susundo, Sendok sendok, Sindor sindor. Sumatra: Alifambang, Alifambang bungo, Alifambang uding, Djelanding, Kaju labuh, Kundui, Labu, Libut, Madag tapak kudu, Mara bulan, Modang kombiri, Njelanding, Ogan ulu, Simar antipa, Sonduk, Terbulan. Borneo. Sarawak & Brunei: Entabulan, Sendok sendok. Sabah: Dalam mata, Katimahar, Melokan. Kalimatan: Garung, Lempaung, Nangair, Pajaug gunung, Parupaek, Pempiring (Schaeffer, 1971).

   Notes — 1. Endospermum beccarianum var. crassirameum was selected by us as the typical variety (containing the type specimen) of E. beccarianum, therefore the name is invalid as the autonym rule applies.

2. This species is very variable in leaf shape and indumentum throughout its distribution range, making especially the identification of sterile specimens difficult. The leaf shape varies from ovate with a cordate to truncate base to obovate with a cuneate base. This variation in leaf shape can even be observed on the same branch. The indumentum on the lower leaf surface also varies considerably. Corner (1939) studied dry specimens and many living trees and he concluded that this variability depended on the age of the trees. Sapling leaves have only long simple hairs, while in mature trees minute, stellately bundled hairs are formed between the simple hairs, covering all of the lamina. When the leaves wither, the long simple hairs disappear, and only the flattened, minute, stellately bundled hairs remain visible.

3. The majority of specimens studied have a lower undersurface that is densely covered by minute, stellately bundled hairs and the presence of simple hairs is less common. Specimens of E. diadenum with a combination of stellately bundled and simple hairs can easily be confused with E. peltatum. It is for this reason that reproductive characters are necessary for identification. The following list of characters may be helpful with identification:

4. Endospermum diadenum also closely resembles E. quadriloculare (see note 2 under E. quadriloculare).

2. Endospermum domatiophorum J.Schaeff.

   Endospermum domatiophorum J.Schaeff., Blumea 19 (1971) 190, map 4 (‘domatiphorum’); Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 79; S.Arias Guerrero & Welzen, Edinburgh J. Bot. 68 (2011) 467, fig. 1a, 7. — Type: Pullen 5922 (holo CANB), Papua New Guinea, Northern Division, Bariji-Managalase area, S. of Toma.

Trees, dioecious, up to 40 m high, bole straight, up to 30 m high, diameter 10–60 cm; branches with hard pith, flowering ones 3–6 mm thick, glabrous to pilose, with a reddish watery exudate. Outer bark grey to brown; inner bark yellow to orange; sapwood cream; heartwood deeper cream. Indumentum consisting of minute stellately bundled hairs. Stipules 3–3.4 by 1–1.3 mm, pilose outside and inside. Leaves: petiole 2–9 cm long, channelled above basally and apically, pilose; blade ovate to obovate, 5–20 by 3.9–16.6 cm, length/width ratio 1.2–1.3, coriaceous, base never peltate, truncate to cuneate, abaxially with or without 1 or 2 round large, yellow extrafloral nectaries near the insertion, 1.3–2.5 by 1–2.2 mm, basally with a rim around the glandular part, margin entire, with minute glands, apex round to acute, drying brown, upper surface glabrous or sparsely hairy, dark green, glossy, lower surface glabrous to sparsely hairy, duller and paler than above, hair tuft domatia in the first and second order bifurcations of the nerves, rarely with glands in the more marginal bifurcations; venation triplinerved or palmate with 5 or 7 nerves of which the 2 outer ones less distinct, upper part pinnate with 4–6 pairs of nerves. Inflorescences panicles, up to 13 cm long, pilose, first branches up to 2 cm long; bracts triangular, 1.5–3 by 1–3 mm, apex acute, pilose outside and inside. Staminate flowers white to yellow, sessile or with a pedicel up to 1 mm long, pilose; calyx completely or basally connate, indistinctly lobed or 4-lobed, 1.1–1.9 by 2.1–2.3 mm, pilose outside, glabrous inside; stamens 6–8; androphore cylindrical, 2.2–2.3 by 0.6–0.8 mm, free part of filaments 0.4–0.6 mm long, anthers, 3- or 4-thecate, 0.6–1 by 0.5–0.8 mm; pistillode somewhat pyramidal, 0.5–0.8 mm long, glabrous. Pistillate flowers single per node, white to cream, sessile or with a pedicel up to 1 mm long, pilose; calyx completely connate, indistinctly lobed, 1.6–2.1 by 1.9–2.7 mm, pilose outside, glabrous inside, green; ovary ovoid to globose, 2.3–2.5 by 1.1–1.5 mm, 1-locular, green; stigma somewhat conical to pyramidal, 0.3–1 by 1.1–1.3 mm, glabrous, light yellow. Fruits ovoid, 6–8 by 3–5 mm, sparsely hairy to pilose, yellow to green, dull. Seeds ellipsoid, somewhat flattened, 4.3–4.5 by 2.2–2.9 mm, ecarunculate, black.

   Distribution — Papua New Guinea.

   Habitat & Ecology — In primary, montane and disturbed forest, associated with Araucaria, Anisoptera and Nothofagus forests; on hills, ridges, in gullies, margins of forest and open areas. Soil: old well-drained volcanic and ultra-basic soil. Altitude: up to 2620 m. Flowering throughout the year; fruiting: from April to December.

   Vernacular names — Boroboro (Schaeffer, 1971), Devarip (Daga), Mayak (Bariji-Managalese).

   Note — Endospermum domatiophorum resembles E. medullosum and can be distinguished from the latter by the presence of hair tuft domatia on the lower leaf surface in the first and second bifurcations of the nerves. Moreover, in E. domatiophorum the lower leaf surface is usually glabrous to sparsely covered with flattened, minute, stellately bundled hairs (only visible with a hand lens), while in E. medullosum the indumentum is tomentose and visible to the naked eye. A diagnostic character for the identification of the species is the presence of a rim around the basal part of the round extrafloral nectaries near the leaf insertion.

3. Endospermum medullosum L.S.Sm.

   Endospermum medullosum L.S.Sm., Proc. Roy. Soc. Queensl. 58 (1947) 53, Plate 1; J.Schaeff., Blumea 19 (1971) 181, map 5; Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 79; S.Arias Guerrero & Welzen, Edinburgh J. Bot. 68 (2011) 468, fig. 1e, 8. — Type: NGF (C.T. White, Dadswell & L.S. Smith) 1738 (holo BRI, n.v.; iso L), Papua New Guinea, Lae.

   Macaranga sp.: Lane-Poole, For. Resources Terr. Papua & New Guinea (1925) 105.

Trees, dioecious, up to 54.8 m high, bole straight, up to 36.6 m high, diameter 30–120 cm; without or with buttresses up to c. 4 m high and 2 m long; branches with a hard pith, flowering ones 7–12.5 mm thick, pilose to densely hairy. Outer bark yellow to pale brown to grey, 5–12.7 mm thick, smooth or with longitudinal fissures and transverse cracks; inner bark cream to yellow to brown, with fragrance of roasted nuts; sapwood undefined or white to cream; heartwood white to straw, soft. Indumentum consisting of stellately bundled and simple hairs. Stipules 5–10 by 1.8–3 mm, densely hairy on both sides. Leaves: petiole 6.4–14.3 cm long, channelled above at the base, pilose to densely hairy, with orange white sap; blade ovate to elliptic, 10.1–27.5 by 8.7–17.5 cm, length/width ratio 1.2–1.6, coriaceous, base not or rarely peltate, emarginate to rounded, abaxially with 2 short, cylindrical, extrafloral nectaries near the insertion, 1.5–1.8 by 1.2–1.6 mm, rimless, at first green and smooth, then orange and finally red with cracked or warty surface, margin entire or somewhat undulate, with numerous minute glands, apex acute, upper surface glabrous or sparsely covered with stellately bundled hairs and/or hirsute simple hairs, dark green, glossy, lower surface densely hairy with minute stellately bundled hairs with many (>10) short flat branches and one longer erect branch and/or simple hairs, lamina completely covered, white to grey to pale green, glands in the more marginal bifurcations of the nerves, hair tuft domatia generally lacking; venation yellowish, basally triplinerved or palmate with 5–8 nerves of which the 2 outer ones less distinct, upper part pinnate with 5–7 pairs of nerves. Inflorescences panicles, pilose or densely hairy, up to 16.7 cm long, with the first branches up to 4.5 cm long; bracts triangular, 3.5–4.2 by 1.9–2.8 mm, apex acuminate, densely hairy on both sides, often with 2 round glands near the base, c.0.8 mm in diam. Staminate flowers whitish to greenish yellow to yellowish, fragrant, sessile or with a pedicel up to 0.3 mm long, pilose; calyx indistinctly lobed or 4-lobed, 1.8–2.1 by 1.4–2 mm, basally connate, densely hairy outside, glabrous inside; lobes triangular, c. 0.3 by 0.6 mm, apex acute; stamens 5–8, yellow, androphore obovate, 1.7–2.2 by 0.6–1.2 mm, free part of filaments c. 0.4 mm long, anthers 3- or 4-thecate, 0.7–0.9 by 0.6–1 mm; pistillode cylindrical, c. 0.4 mm long, sparsely hairy beneath. Pistillate flowers single per node, sessile or with a pedicel up to 0.3 mm long; calyx completely connate, indistinctly lobed, 2.1–2.4 by 1.7–1.9 mm, densely hairy outside, glabrous inside; ovary obovoid, 1.5–1.9 by 1.2–1.4 mm, green, 1-locular; stigma somewhat pyramidal, 0.5–0.9 by 0.9–1.4 mm, glabrous. Fruits pale green to yellow, dull, ovoid to globose, 5–7 by 5–5.4 mm, sparsely hairy. Seeds globose, 4.6–4.8 by 3.5–3.7 mm, ecarunculate.

   Distribution — The Moluccas, New Guinea, Solomon Islands and Vanuatu.

   Habitat & Ecology — In primary or late secondary forest, alluvial forest, on ridges, along creeks, on hills, steep slopes and flood plains; associated with Pometia (Sapindaceae), Celtis (Ulmaceae), Fagaceae, Myrtaceae, Annesijoa (Euphorbiaceae) and Pimelodendron (Euphorbiaceae). Soil: clay, sandy clay, gravel and deep well-drained volcanic scoria. Altitude: up to 1070 m. Flowering and fruiting throughout the year.

   Uses — The wood is used for making canoes. Young leaves are sometimes eaten as a vegetable (Schaeffer, 1971).

   Vernacular names — New Guinea: Adokko (Manikiong), A’ugo (Koropa; Onjob), Joemkejoe (Kemtoek), Joerasan (Asmat), Karubu (Bembi), Kirikendita (Madang), Koindza (Jal), Moenabore (Biak), Nakau (Talasea), Rikwa (Nemo), Sajoemena, Saimena (Mooi), Sisibkoiru (Wanigela), Teppateh (Rawa). Solomon Islands (Schaeffer 1971): A’sa, Kandiki, Manogo.

   Notes — 1. The specific epithet is derived from the Latin word ‘medulla’ meaning pith and refers to the branchlets containing pith in this species as opposed to other New Guinean species (E. moluccanum and E. myrmecophilum) that are hollow (Smith, 1947).

2. Endospermum medullosum resembles E. domatiophorum (see note under latter) and E. myrmecophilum. Endospermum medullosum can be distinguished from E. myrmecophilum by the presence of pith in the branches, the leaf base mostly not peltate, and a tomentose indumentum visible to the naked eye.

3. The short, cylindrical extrafloral nectaries near the leaf insertion have a lateral position, and they are sometimes found at the petiole apex and can be seen from above.

4. The specimens NGF 1783, NGF 25932, and NGF 29307 differ from other material studied by having hair tuft domatia in the first and second bifurcation of the nerves and the extrafloral nectaries near the insertion are round and have a rim around the basal part.

4. Endospermum moluccanum (Teijsm. & Binn.) Kurz

   Endospermum moluccanum (Teijsm. & Binn.) Kurz, J. Bot. 5 (1867) 23; Becc., Malesia 2 (1884) 38; Merr., Interpr. Herb. Amboin. (1917) 326; K.Heyne, Nutt. Pl. Ned.-Ind. 2^nd ed., 2 (1927) 956; Rant, Natuurk. Tijdschr. Ned.-Indië 94 (1934) 113; Ann. Jard. Bot. Buitenzorg 48 (1938) 123; Schodde, Blumea 15 (1967) 401, in obs.; J.Schaeff., Blumea 19 (1971) 182, map 2; Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 80; Kew Bull. 37 (1982) 18. — Capellenia moluccana Teijsm. & Binn., Natuurk. Tijdschr. Ned.- Indië 29 (1867) 239, Fig.; S.Arias Guerrero & Welzen, Edinburgh J. Bot. 68 (2011) 470, fig. 9, 10. — Type: Teijsmann & Binnendijk s.n. (holo BO), Indonesia, Moluccas. See note 1.

    [Arbor regis Rumph., Herb. Ambon. 2 (1741) 257, t. 85, nom. inval. — Hernandia sonora auct. non L.: Stickman in L., Amoen. Acad. 4 (1759) 117, 122, 141.]

   Endospermum formicarum Becc., Malesia 2 (1884) 44; Schum. & Hollr., Fl. Kais. Wilh. Land (1889) 804; Warb., Bot. Jarb. 13 (1891) 348; Schum. & Lauterb., Fl. Schutzgeb. Südsee (1900) 406; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iv (1912) 36, Fig. 11; Docters vanLeeuwen, Treubia 10 (1929) 431; L.S.Sm., Proc. Roy. Soc. Queensl. 58 (1947) 58; Schodde, Blumea 15 (1967) 401, in obs.. — Type: Beccari PP 648 (holo FI, n.v.), Nuova Guinea [= Indonesian Papua], Andai. Synonymy after Schaeffer (1971).

   Endospermum labios Schodde, Blumea 15 (1967) 397; Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 79. — Endospermum labios Schodde subsp. labios: Schodde, Blumea 15 (1967) 399, Fig. 1. — Type: Brass 32607 (holo CANB; iso US), Papua New Guinea, Markham Valley, Umi River.

   Endospermum labios Schodde subsp. gracilipes Schodde, Blumea 15 (1967) 400. — Type: BSIP (Whitmore) 3979 (holo LAE; iso US), Solomon Islands, SE Choiseul Island, ridge NE from harbour at Ruruvai.

Trees, dioecious, up to 33.4 m high, bole up to 18.4 m high, diameter 12–40 cm; without or with buttresses up to 1.4 m high, extending 0.6 m over the ground; branches hollow, flowering ones 10–16.2 mm thick, conical at the apex, glabrous to sparsely hairy, with exit pores for nesting ants; latter round, 1.8–3 mm in diam.; apex of shoots with triangular bract-like structures (or their scars), 1.5–1.6 by 1.4–1.9 mm (see note 1). Outer bark greyish, mottled with white, pink or green patches, smooth except for some shallow longitudinal and transverse cracks; inner bark green, turning light orange to white, sap whitish to yellow, milky or watery; sapwood pale yellow; heartwood white to pale yellow. Indumentum consisting of stellately bundled and simple hairs. Stipules unknown. Leaves: petiole 8.4–23.5 cm long, basally and apically channelled above, otherwise terete, sparsely hairy to pilose; blade ovate to orbicular, 15.7–35 by 12.3–25.5 cm, length/width ratio 1.2–1.4, papyraceous or rarely coriaceous, base peltate, truncate, abaxially with 2 extrafloral nectaries near the insertion, cylindrical, 0.9–2.6 by 1.5–2.1 mm, depressed in the middle, margin entire or somewhat undulate, with minute glands, apex acuminate to cuspidate, upper surface glabrous to sparsely covered with stellately bundled hairs and/or simple hairs, dark green, shiny, lower surface glabrous to pilose covered with hirsute hairs, duller and paler than above; venation palmate with (7)8-10 nerves, upper part pinnate with 5 or 6 pairs of nerves. Staminate inflorescences panicles, pistillate ones racemes, pilose to densely hairy, up to 33.7 cm long, first branches up to 12.5 cm long; bracts either leaf-like in basal parts of inflorescence, ovate, c. 13.5 by 10.2 mm, sessile or with a petiole up to 3 mm, with 2 round glands at the base, c. 1.1 by 1 mm, base rounded, apex acuminate, pilose outside and inside or bracts triangular in higher parts of inflorescence, 6–8 by 2.2–4.2 mm, apex acuminate to cuspidate, curved, pilose outside, glabrous to sparsely hairy inside, often with 2 glands underneath at the base, c 1.2 mm in diameter. Staminate flowers pale yellow, fragrant, sessile; calyx 4-lobed, 2.3–4 by 2.2–3.2 mm; lobes triangular, 0.9–2 by 1.2–1.7 mm, basally connate, apex acute, sparsely hairy outside, glabrous inside; stamens 9–12, androphore cylindrical, 4.6–5.5 by 1.4–2.8 mm, free part of filaments 0.4–0.8 mm long, anthers yellow, 4-thecate, 0.7–1 by 0.7–1.2 mm; pistillode cylindrical, 0.7–1 by 0.4–0.5 mm, glabrous. Pistillate flowers single per node, light green to whitish, sessile; calyx indistinctly lobed, 3.8–6.2 by 4.8–11.3 mm, green, sparsely hairy outside, glabrous inside; ovary cylindrical to obovoid, 2.1–7.4 by 1.8–7.8 mm, (3-)4-6-locular; stigma yellow, indistinctly lobed, concave, 0.5–1.5 by 3–6.6 mm, glabrous. Fruits light green, globose, 5–23.8 by 5.5–18 mm, sparsely covered with minute hairs. Seeds ellipsoid or globose, 4.8–6.3 by 3–4.1 mm, ecarunculate.

   Distribution — Sulawesi, the Moluccas, New Guinea and the Solomon Islands.

   Habitat & Ecology — In primary and secondary forest, swampy and alluvial forest, disturbed (logged) areas, along rivers and roads. Soil: clay, sandy clay, limestone, gravel, loose volcanic soil. Altitude: up to 900 m. Flowering and fruiting throughout the year.

   Uses — In the Moluccas the tree is used for constructing semi-permanent houses. On New Guinea the bark is considered to have anti-fertility properties, and the leaves are used as treatment for infections and aches.

   Vernacular names — Moluccas: Kaju radja, Pea-pea, Ofo. New Guinea: Bondoa (Waskuk), Kidikendi (Madang), Koin (Jal), Kobaguno (Kutubu), Kornu, Kungum ( Orme), Lalaul, Luki, Mangoeriw ( Biak), Mawr ( Mawan), Munampun (Kaigorin), Nauwauwa, Nakok malu (Nakanai), Ngebbie, Njebbie (Manikiong), Pehpi (Bembi), Poteh (Rawa), Sajon (Tehid), Sejoem (Mooi), Seraida ( Tehid), Taingino (Wapi), Taraparo (Wandammen), Wakopak (Amberbaken), Wakpak ( Andjai), Wapaap (Kebar).

   Notes — 1. Teijsmann & Binnendijk published Capellenia moluccana in the same year as Kurz published Endospermum moluccanum (1867). We could not trace the exact dates of publication of the journals, but from Kurz’s short article it is obvious that he used Teijsmann & Binnendijk’s publication and that he interpreted the figure correctly as an Endospermum. This means that Teijsmann & Binnendijk published their paper before Kurz published his.

2. The triangular bract-like structures found near the apex of the branchlets may serve as food bodies for the ants. They are only present near the branch apex, but many scars are present on lower parts of the branches as if the ‘bracts’ were eaten away and had served as food bodies for the ants living in the hollow pith.

3. Specimens from the Moluccas have smaller fruits in comparison with specimens from New Guinea.

4. Airy Shaw (1980) recognised two different species, E. moluccanum (Teijsm. & Binn.) Kurz and E. labios Schodde. The characters that Airy Shaw used to distinguish the species, such as the indumentum on the lower leaf surface and inflorescence and the shape of the stigma and bracts, are too variable to be used in species delimitation. However, there is a considerable difference in fruit size, whereby E. moluccanum (in the strict sense) has smaller fruits than E. labios. This difference may be explained by differences in climatological conditions. Endospermum moluccanum occurs in Sulawesi and Moluccas, areas with a drier climate than where the E. labios form is found. The latter occurs in the wetter parts of New Guinea and the Solomon Islands. Both forms have hollowed branches, triangular bract-like structures near the apex of the branchlets (an autapomorphy), two cylindrical extrafloral nectaries near the leaf insertion and the leaf base always peltate. For these reasons we follow Schaeffer (1971) rather than Airy Shaw (1980) and consider the names to be synonyms with Endospermum moluccanum having priority.

5. Endospermum myrmecophilum L.S.Sm.

   Endospermum myrmecophilum L.S.Sm., Proc. Roy. Soc. Queensland. 58 (1947) 56, Plate II; J.Schaeff., Blumea 19 (1971) 181, map 3; Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 80; S.Arias Guerrero & Welzen, Edinburgh J. Bot. 68 (2011) 472, fig. 1b, 11. — Type: NGF (C.T. White, Dadswell & L.S. Smith) 1640 (holo BRI, n.v.; iso L), Papua New Guinea, Morobe Prov., Yalu.

Trees, dioecious, up to 36.5 m high, bole up to 23 m high, diameter 20–90 cm; without or with buttresses up to c. 1 m high; branches hollow, flowering ones 10–17 mm thick, conical at the apex, glabrous to sparsely hairy, with a watery whitish sap, with exit pores for nesting ants; latter round, 2.4–2.8 mm wide. Outer bark white to grey to light brown, fairly smooth, longitudinally fissured and covered with insignificant lenticels, c. 4 mm thick; inner bark white to orange to light brown; sapwood straw to yellow to light brown, soft; heartwood cream. Indumentum consisting of stellately bundled and simple hairs. Stipules 2–8.7 by 1.4–2.2 mm, densely hairy on both sides. Leaves: petiole 5.2–16.3 cm long, channelled above at the base, sparsely hairy; blade ovate to orbicular, 8.5–27.4 by 9.7–22 cm, length/width ratio 0.9–1.2, coriaceous, rarely papyraceous, base peltate or not, emarginate to truncate (to rounded), abaxially with 2 flat, elliptic, yellowish to green extrafloral nectaries near the insertion, 1–2.4 by 1.3–4 mm, margin somewhat undulate, often with minute glands, apex round to acute, upper surface glabrous, dark green, glossy, lower surface densely covered with minute stellately bundled hairs and/or simple hairs, rarely with glands in the more marginal bifurcations of the nerves, paler and duller than above; venation amber to yellow, palmate with 7–9 nerves, upper part pinnate with 4 or 5 pairs of nerves. Inflorescences panicles, up to 23.3 cm long, pilose to densely hairy, first branches up to 4.5 cm long; bracts triangular, 1.9–3.2 by 1.1–1.5 mm, apex acuminate, densely hairy on both sides. Staminate flowers cream to yellow to pale green, sessile or with a pedicel up to 1.2 mm, pilose; calyx indistinctly lobed or 4-lobed, 1–1.3 by 0.8–1.8 mm, basally connate, hairy outside, glabrous inside; lobes triangular, c. 0.3 by 0.4 mm, apex acute; stamens c. 5, androphore obovate, 2–2.5 by 1.2–1.5 mm, free part of filaments c. 0.3 mm long, anthers 4-thecate, 0.7–0.8 by 0.8–0.9 mm; pistillode cylindrical, c. 0.4 by 0.2 mm, glabrous. Pistillate flowers single or in groups of up to 3 flowers per node, green, sessile or with a pedicel up to 3 mm long, pilose; calyx indistinctly lobed or 4-toothed, 1.1–1.4 by 1.3–1.9 mm, densely hairy outside, glabrous inside; ovary obovate, 1.2–1.8 by 1.1–1.4 mm, 1-locular; stigma flattened or somewhat pyramidal, indistinctly lobed, c. 0.4 by 0.9–1.4 mm, glabrous. Fruits pale green, globose, 5–6.5 by 4.8–6.1 mm, pilose to densely hairy. Seeds globose, 3.7–4.6 by 4–4.5 mm, ecarunculate.

   Distribution — New Guinea (Papua) and Australia (North Queensland).

   Habitat & Ecology — In lowland primary rain forest, swamps and alluvial forests, and logged areas; along streams, on ridges, hills, margins of forests and floodplains. Soil: brown sandy loam. Altitude: up to 457 m. Flowering: from February until July; fruiting: from February to November.

   Vernacular names — New Guinea: Eladina (Koriari), Sisib-koiru (Wanigela).

   Notes — 1. Endospermum moluccanum and E. myrmecophilum are the only two myrmecophilic species within Endospermum and are, therefore, probably closely related. Both have hollow branches with exit pores for the ants. Endospermum myrmecophilum differs from E. moluccanum by the absence of the triangular bract-like structures at the apex of the branches and the extrafloral nectaries near the leaf insertion are flat and elliptic and not cylindrical. Moreover, the fruits are always 1-locular in comparison with the (3)4–6-locular fruits of E. moluccanum.

2. See also note 2 under E. medullosum.

3. The specimens LAE 68860 and NGF 19645 differ from other material studied in having branches with pith.

6. Endospermum ovatum Merr.

   Endospermum ovatum Merr., Philipp. J. Sci. 9, Bot. (1914) 481; Enum. Philipp. Fl. Pl. 2 (1923) 457; J.Schaeff., Blumea 19 (1971) 190, map 4; Airy Shaw, Alphab. Enum. Euphorb. Philippine Isl. (1983) 24; S.Arias Guerrero & Welzen, Edinburgh J. Bot. 68 (2011) 474, fig. 12. — Type: BS (Fénix) 15921 (holo PNH, lost; iso BM, US), Philippines, Mindanao, Butuan Subprov., near Sumulao.

Trees, dioecious, up to 30 m high, diameter c.30 cm; branches with a hard pith, flowering ones 5–5.2 mm thick, glabrous to sparsely hairy. Outer bark white, smooth; sapwood yellow; heartwood yellow. Indumentum consisting of minute stellately bundled hairs. Stipules c. 1.5 by 0.5 mm, densely hairy outside, glabrous inside. Leaves: petiole 4–9.3 mm long, channelled above basally and apically, otherwise round, sparsely hairy to pilose; blade ovate to elliptic, 8.2–13.2 by 5.4–10.2 cm, length/width ratio 1.3–1.5, coriaceous, base not peltate, round to acute, abaxially with or without 1 or 2 short cylindrical extrafloral nectaries near the insertion, 1.2–1.6 by 0.8–0.9 mm, rimless, margin entire, rarely with minute glands, flat, apex acuminate, upper surface glabrous, lower surface sparsely hairy to pilose with minute stellately bundled hairs, domatia absent; venation triplinerved, upper part pinnate with 4 or 5 pairs of nerves. Inflorescences panicles, up to 17.5 cm long, pilose to densely hairy, first branches up to 3.5 cm long; bracts triangular, c. 2.3 by 1.5 mm, apex acute, densely hairy on both sides. Staminate flowers cream, with a pedicel up to 1 mm long; calyx indistinctly lobed, 1.1–1.4 by 1.7–1.9 mm, completely connate, densely hairy outside, sparsely hairy inside; stamens 8 or 9, androphore cylindrical, 1.8–1.9 by c. 0.9 mm, free part of filaments 0.3–1 mm long, anthers 3-thecate, 0.4–0.5 by 0.5–0.7 mm; pistillode cylindrical, c. 0.4 by 0.2 mm, sparsely hairy. Pistillate flowers unknown. Fruits disintegrated, unknown. Seeds globose, c. 3.1 by 3.2 mm, ecarunculate.

   Distribution — Philippines (Mindanao).

squares: E. ovatum; dots: E. quadriloculare

   Habitat & Ecology —In logged areas. Flowering: April; fruiting: August.

   Uses — For making toothpicks and matchsticks.

   Vernacular name — Gubas.

   Note — This description is based on two specimens only. More material is needed, especially specimens with pistillate flowers and/or fruits.

7. Endospermum peltatum Merr.

   Endospermum peltatum Merr., Publ. Bur. Sci. Gov. Lab. 35 (1906) 35; Philipp. J. Sci. 1, Suppl. 1 (1906) 82; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iv (1912) 37; Chatterjee, Kew Bull. 1949 (1950) 564; Merr., Enum. Philipp. Fl. Pl. 2 (1923) 457; J.Schaeff., Blumea 19 (1971) 188, map 3; Airy Shaw, Kew Bull. 26 (1972) 259; Kew Bull., Addit. Ser. 5 (1975) 109; Kew Bull. 37 (1982) 18; Alphab. Enum. Euphorb. Philippine Isl. (1983) 24; Chantharaprasong in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 257; S.Arias Guerrero & Welzen, Edinburgh J. Bot. 68 (2011) 475, fig. 1f, 13. — Lectotype (designated by Schaeffer, 1971): Merrill 2603 (holo K), Philippines, Luzon, Tayabas Prov., Pagbilao.

    [Endospermum philippinense Elmer ex Merr., Enum. Philipp. Fl. Pl. 2 (1923) 457, nom. nud., in synon.].

Trees, dioecious, up to 50 m high, bole up to 29 m high, diameter 16–90 cm, without or with buttresses up to 3 m high, extending c. 2 m over the ground; branches with a hard pith, flowering ones 8–11 mm thick, pilose. Outer bark white to yellow to green to brown to grey, 0.5-13 mm thick, smooth; inner bark pale yellow to orange to brown; sapwood white to pale yellow to straw-coloured; heartwood straw-coloured, soft. Indumentum consisting of stellately bundled and simple hairs. Stipules 4.8–5.8 by 1.7–2.5 mm, densely hairy on both sides. Leaves: petiole 7.5–26 cm long, channelled above at the base, densely hairy to pilose; blade narrowly to broadly ovate, 11–29.8 by 8.7–23.5 cm, length/width ratio 1.2–1.3, coriaceous, base peltate or not, cordate to rounded (to acute), abaxially with 2 cylindrical large extrafloral nectaries near the insertion, 1.3–1.9 by 1.1–1.8 mm, rimless, margin entire with numerous minute glands, apex acute to acuminate, upper surface glabrous or sparsely covered with strigose simple hairs and/or stellately bundled hairs, lower surface densely hairy to pilose with stellately bundled hairs, with at most 5 equal hairs in a bundle, and/or simple hairs, lamina visible, greyish green to pale green, dull and with glands in the more marginal bifurcations of the nerves, domatia absent; venation yellow beneath, basally triplinerved or palmate with 5–9 basal nerves, upper part pinnate with 5–8 pairs of nerves. Inflorescences panicles, densely hairy, up to 42 cm long with the first branches up to 13 cm long; bracts triangular, 1.8–3.2 by 1.8–1.9 mm, apex acuminate, pilose both sides, often with 2 flat round glands near the base, c. 0.5 mm wide. Staminate flowers white to yellow to pale green, sessile or with a pedicel up to 1.5 mm long, pilose; calyx 4-lobed, 1.5–1.7 by 1.7–2.1 mm, basally connate, densely hairy outside, pilose inside; stamens 9 or 10, androphore obovate, 1.8–2.2 by 0.9–1.1 mm, free part of filaments 0.8–1.5 mm long, anthers 3-thecate, 0.4–0.5 by 0.8–1 mm; pistillode cylindrical, splitting into 2 or 3 parts, 0.5–0.6 by 0.4–0.8 mm, sparsely hairy on the top. Pistillate flowers single per node, green to yellow, sessile or with a pedicel up to 3.2 mm long; calyx 4-lobed or toothed, 1–1.4 by 2–2.6 mm, basally connate, densely hairy outside, pilose inside; ovary globose, 2- or 3(-4)-locular, 1.5–2.8 by 1–2 mm; stigma flattened, 1.5–1.9 by 0.3–0.6 mm, glabrous. Fruits brown to green to yellow, globose, 5–8.2 by 6–2.7 mm with or without 3 triangular lobes on top, 0.4–1 mm long, lobes on inside often with a groove. Seeds globose, 3.4–5 by 3.2–3.8 mm, ecarunculate.

   Distribution — Peninsular Thailand, Andaman Islands (Schaeffer, 1971), Malay Peninsula, Borneo, and the Philippines.

   Habitat & Ecology — In primary (mixed dipterocarp) forest, secondary forest, and disturbed areas; along streams and roads, on hills, ridges and steep valley slopes. Soil: sandy, sandstone, loam soil on limestone, silty clay loam, alluvium and periodically inundated soil. Altitude: up to 640 m. Flowering: from April to November; fruiting: from February to November.

   Uses — Seeds contain edible oil. In the Philippines the wood is used for making wooden shoes and matches.

   Vernacular names — Borneo: Buah icras, kemiri (Malay), Kedjó luk long, Marampangi (Dusun), Peridja buaja, Sedok-Sedok, Seduk- seduk (Malay). Philippines: Gubas.

   Notes — 1. The epithet ‘peltatum’ indicates that the leaf base is peltate. However, this character is variable and non-peltate leaves, even together with peltate ones, are found.

2. The species from the Philippines more often have cordate leaves than specimens from Borneo and the Malay Peninsula.

3. SMHI 479, SMHI 881, SMHI 963, and Soejarto 7682 which were collected on Palawan Island differ from the other specimens studied by having a less dense indumentum on the lower leaf surface

8. Endospermum quadriloculare Pax & K.Hoffm.

   Endospermum quadriloculare Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iv (1912) 36, Fig. 10; S.Moore, J. Bot. 63, Suppl. (1926) 104; J.Schaeff., Blumea 19 (1971) 190, map 3; Airy Shaw, Kew Bull. 36 (1981) 294; S.Arias Guerrero & Welzen, Edinburgh J. Bot. 68 (2011) 476, fig. 1d, 12. — Type: Forbes 2751 (holo B, probably lost; iso BM, n.v., GH, L, P), Indonesia, Sumatra, Palembang.

   Endospermum banghamii Merr., Contr. Arnold Arbor. 8 (1934) 89; J.Schaeff., Blumea 19 (1971) 185, map 5; Airy Shaw, Kew Bull. 36 (1981) 293. — Type: W.N. & C.M. Bangham 739 (holo CGE; iso A), Indonesia, Sumatra, Atjeh, Takengon.

   Endospermum ronaldii J.Schaeff., Blumea 19 (1971) 185, map 4. — Type: SF (M.R. Henderson) 18396 (holo SING, n.v.; iso K, n.v., L), Malaysia, Pahang, Pulau Tioman, Sedagong.

Trees, probably dioecious, but rarely monoecious because of bisexual flowers, up to 40 m high, bole clear, diameter c.60 cm; without or with buttresses up to 2.45 m; branches with a hard pith, flowering ones 5.2–7 mm thick, glabrous to sparsely hairy. Outer bark whitish, smooth, with circular rings; inner bark yellow; wood white to yellow. Indumentum consisting of minute, stellately bundled hairs. Stipules 4.2–4.7 by 1.4–1.5 mm, pilose. Leaves: petiole 4.9–15.3 cm long, channelled above at the base, glabrous to sparsely hairy; blade elliptic to ovate, 9.2–25 by 6.3–14.1 cm, length/width ratio 1.5–1.8, papyraceous to rarely coriaceous, base peltate or not, emarginate to acute, abaxially with or without 1 or 2 lateral cylindrical, large extrafloral nectaries near the insertion, 1.1–3 by 1–1.4 mm, rimless, margin somewhat undulate, rarely with minute glands, apex acute to cuspidate, upper surface glabrous, dark green, glossy, lower surface glabrous to densely covered with minute stellately bundled hairs, duller and paler than above, without domatia; venation basally triplinerved or palmate with 5–7 basal nerves of which the 2 outer ones less distinct, upper part pinnate with 4–6 pairs of nerves. Inflorescences panicles, densely hairy, up to 18 cm long with the first branches up to 6 cm long; bracts triangular, 1.8–2.4 by 1.5–1.6 mm, apex acuminate, densely hairy both sides, often with 2 round glands near the base, c. 1 mm wide. Staminate flowers pale yellow, with a pedicel up to 1.2 mm long; calyx indistinctly lobed, 1.1–1.2 by c. 1.8 mm, pilose outside, sparsely hairy inside; stamens c. 12, yellow to brown, androphore obovate, 2–2.1 by 1.2–1.3 mm, free part of filaments c.0.5 mm long, anthers 3-thecate, 0.6–0.8 by 0.8–0.9 mm; pistillode 3- or 4-lobed, 0.4–0.5 by c. 0.6 mm, sparsely hairy beneath. Pistillate or bisexual flowers single per node, with a pedicel up to 4 mm long, pilose; calyx indistinctly lobed, 1.3–2.4 by 1.8–3.8 mm, pilose outside and inside; bisexual ones with or without a cylindrical androgynophore of c. 2.2 by 0.8 mm, with 8–10 stamens at base of ovary, comparable to those of staminate flowers, persistent in fruit; ovary globose (unisexual) to cylindrical (bisexual), (3)4 or 5-locular, 1.3–3 by 0.8–2.4 mm; stigma lobed or not, 2–2.2 by 0.9–1.4 mm, glabrous, with or without raised triangular lobes on top, c 1 mm long, up to 2 mm in fruit, lobes on the inside with a groove. Fruits pale blue to green to brown, globose or ellipsoid, 7.8–13 by 6–11 mm, pilose; stigma, somewhat enlarged, 1.8–2.1 by 2.5–3.2 mm. Seeds ellipsoid, 5.9–5.6 by 2.5 –3.2 mm, with a slender caruncle, 1–1.5 mm long.

   Distribution — Thailand, Peninsular Malaysia, Sumatra.

dots: E. quadriloculare; squares: E. ovatum. 

   Habitat & Ecology — In primary and evergreen forests, along roads, on hills and ridges. Altitude: up to 1220 m. Flowering: from April to May; fruiting: from May to August.

   Vernacular name — Peninsular Malaysia: Sesendok.

   Notes — 1. Bisexual flowers are only known from the type specimens of E. banghamii and E. ronaldii. The flowers of E. banghamii have dehisced anthers that contain pollen that may be fertile. The anthers are located at the base of the ovary. Endospermum ronaldii has branched inflorescences with solitary flowers and the anthers are arranged on an androgynophore, basal to the ovary. Both names are placed in synonymy because the types do not differ in any other characters from E. quadriloculare and both specimens are unique (though collected in well sampled areas), which indicates that they are more likely to be exceptional forms than good species. Bisexual flowers are rare in Euphorbiaceae and seldom occur in Malesia, e.g., seemingly bisexual flowers with staminodes are found in Agrostistachys staminodiatus Sevilla (Sevilla & Welzen, 2001) and real bisexual flowers occur in several New Guinean species of Aporosa Blume (Schot, 2004).

2. Endospermum quadriloculare and E. diadenum are sympatric species and some sterile specimens may easily be confused. In general, E. quadriloculare differs from E. diadenum in having a sparser indumentum (most specimens that we studied have a glabrous to sparsely lower leaf surface), leaf base peltate or not, papyraceous leaves rarely with glands around the margin, and the extrafloral nectaries near leaf insertion are longer (1.1–3 by 1–1.4 mm) than those of E. diadenum (0.4–0.5 by 1.5–2.2 mm).

Excluded names

Endospermum eglandulosum Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii: 418 (1914). — Type: Beccari PB 1347 (isotype: BM000645755), Borneo, Sarawak. = Sterculia macrophylla Vent. (Sterculiaceae), e.g., see Govaerts et al. (2000), World Checkl. Bibliogr. Euphorb. 2.