Malesian Euphorbiaceae Descriptions

17. BLUMEODENDRON (Euphorbiaceae)

P.C. van Welzen & M.A.D. Ottens-Treurniet

van Welzen, P.C. & Ottens-Treurniet, M.A.D. 2015. A revision of the Malesian genus Blumeodendron (Euphorbiaceae). Blumea 61 (2016) 64–82.

Genus description

Key to the Malesian species

Species description

Excluded taxon

Blumeodendron (Müll.Arg.) Kurz

    Blumeodendron (Müll.Arg.) Kurz, J. Asiat. Soc. Bengal 42 (1873) 245; Forest Fl. Burma 2 (1877) 391; J.J.Sm., Meded. Dept. Landb. Ned.-Indië 10 (1910) 458; Koord., Exkurs.-Fl. Java 2 (1912) 493; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 47; Merr., Philipp. J. Sci. 16 (1920) 554; Ridl., Fl. Malay Penins. 3 (1924) 281; Pax & K.Hoffm. in Engl. & Harms, Nat. Pflanzenfam. ed. 2, 19C (1931) 107; Backer & Bakh.f., Fl. Java 1 (1963) 479; Airy Shaw, Kew Bull. 16 (1963) 348; Kew Bull. 26 (1972b) 224; Whitmore, Tree Fl. Malaya 2 (1973) 68; Wheeler, Taxon 24 (1975) 535; Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 57; Kew Bull., Addit. Ser. 8 (1980) 37. 1980; Kew Bull. 36 (1981) 267; Kew Bull. 37 (1982) 9; Alphabet. Enum. Euph. Philipp. Is. (1983) 10; G.L.Webster, Ann. Missouri Bot. Gard. 81 (1994) 77; Radcl.-Sm., Gen. Euphorbiacearum (2001) 170; Chayam. in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 130; G.L.Webster in Kubitzki, Fam. Gen. Vasc. Pl. 11 (2014) 125; Ottens & Welzen, Blumea 61 (2016) 66. — Mallotus Lour. sect. Blumeodendron Müll.Arg. in DC., Prodr. 15, 2 (1866) 956; Benth. in Benth. & Hook.f., Gen. Pl. 3 (1880) 319; Hook.f., Fl. Brit. India 5 (1887) 427; Pax in Engl., Pflanzenfam. 3(5) (1890) 53. — Lectotype (designated by Wheeler 1975): Elateriospermum tokbrai Blume (= Blumeodendron tokbrai (Blume) Kurz).

Trees, dioecious; branchlets generally round in section, nodes thickened. Indumentum consisting of small, orange, lepidote hairs, glabrescent, few species locally with additional simple hairs. Stipules absent. Leaves alternate, subopposite or pseudo-whorled, simple; petiole apically but more so basally pulvinate, in section round but grooved transversely when dry; blade elliptic to obovate, margin entire, revolute, extrafloral nectaries 2, adaxially near base and additional smaller ones along midrib and margin; venation pinnate, marginal vein indistinct, primary and secondary nerves slightly raised above, secondary nerves looped and closed near the margin, tertiary nerves scalariform, higher order nerves reticulate or partly sclariform. Inflorescences axillary or terminally thyrses, one or more together, erect. Flowers: pedicel with abscission zone; flowers actinomorphic, 5-merous, buds globose; sepals elliptic, valvate, margin entire; petals absent; disc present. Staminate flowers: sepals 3 or 4; disc glands on convex receptacle, providing a ruminate surface with the stamens in between the glands; stamens 31–40, anthers basifixed, 2-thecate, opening latrorse via lengthwise slits; pistillode absent. Pistillate flowers: sepals 5 or 6; disc annular, broad; ovary 2- or 3-locular, placenta basal, ovulum single per locule, hemitropous; styles very short; stigmas 2 or 3, entire, often recurved, above papillate. Fruits capsular, globular or ovoid, tardily completely septicidally and partly loculicidally dehiscent into bivalved cocci; pedicels thickened; wall woody, surface somewhat knobbly. Seeds ovoid to subglobular, with sometimes flattened sides, more or less bean-shaped; sarcotesta present.

    Distribution — Nine species ranging from Burma and the Andamans via Thailand through Malesia to New Guinea and the Bismarck Archipelago. 

Key to the species

1. Blumeodendron borneense Pax & K.Hoffm.

    Blumeodendron borneense Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xiv (1919) 14; Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 58; Ottens & Welzen, Blumea 61 (2016) 66, Fig. 1, Map 1. — Blumeodendron tokbrai (Blume) Kurz var. borneense (Pax & K.Hoffm.) J.J.Sm. ex Airy Shaw, Kew Bull. 36 (1981) 269. — Type: Beccari PB 2976 (FI, not seen; iso K), Borneo.

    Blumeodendron concolor Gage, Rec. Bot. Surv. India 9 (1922) 244; Ridl., Fl. Malay Penins. 3 (1924) 281; Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 59. — Type: Curtis KD 1368 (K), [Malay Peninsula,] Pangkor.

    Blumeodendron sp.: Merr., Pl. Elmer. Born. (1929) 157. — pro Elmer 21129 (L), British North Borneo [= Sabah], Tawao.

    Blumeodendron tokbrai auct. non (Blume) Kurz: Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 60, p.p., ‘form with oblong leaves’.

Trees, up to 35 m high, bole to 15 m high, d.b.h. to 30 cm(–2 m); buttresses sometimes present, few, c. 1 m high, c. 0.3 m out, c. 20 cm thick; stem generally round, nodes notably thickened; lenticels indistinct, round; flowering branches 5–28 mm thick, terminal ones often triangular of flattened in section; distance between internodes c. 18 cm; sympodial growth via axillary buds obvious. Outer bark tan to reddish brown, to dark brown, whitish grey, (pale) grey to grey-brown dippled, smooth to rough to fissured and peeling off, brittle, c. 1 mm thick; inner bark white to pale yellow, orange, pale greenish, pale brown outside to yellow inside, c. 4 mm thick; sapwood white to pale yellow; heartwood reddish. Indumentum: simple hairs absent. Leaves in (pseudo-)whorls of 3 leaves to alternate (see notes 1 & 2); petiole 0.7–10 cm long, diam. of thinnest part 0.9–3 mm, shiny, basal pulvinus 1.1–3.4 mm diam.; lepidote hairs present, older parts glabrous; blade ovate to elliptic (see note 3), 6–40 by 2.7–21 cm, length/width ratio 1.7–3.2, pergamentaceous to subcoriaceous, symmetric, glabrous, smooth dark green, drying light green, base rounded to obtuse, margin recurved, apex acuminate to cuspidate, tip to 1 cm long, extrafloral nectaries c. 6–18 along midrib, c. 6–26 along margin, both sides smooth; venation slightly and partly raised above, secondary nerves 4–7 pairs, at c. 46˚ angle with midrib, tertiary nerves perpendicular to midrib and secondary nerves, not raised above, higher order nerves reticulate, not raised above. Inflorescences ramiflorous to axillary and terminal, flowers almost fasciculate when staminate buds young, pistillate inflorescences single, length unknown. Staminate flowers only known from one specimen, either as bud or withered: pedicel c. 19 mm long, round; sepals 3, ovate, c. 4 by 3 mm, completely reflexed; stamens many, anthers c. 0.9 by 0.9 mm. Pistillate flowers unknown. Fruits c. 3 per inflorescence, subglobose, basally slightly sulcate, capsular, 3.5–4.1 by 3.1–4.3 cm, yellow to brown, 2- or 3-locular, ripening from green to yellow, dry brown, surface somewhat knobbly, vein ridges c. 3; pedicels c. 2 cm long, up to c. 0.4 cm diam., abscission zone indistinct; margin slightly thickened; wall 1–2 mm thick; endocarp enclosing two or more seeds; stigmas usually persistent. Seeds bean-shaped, 2.1–2.3 by 1.4–2 by 1–1.1 cm, attached in middle of bean, covered by a thin sarcotesta, dirty white to yellowish, sweet, edible.

    Distribution — Malay Peninsula (very rare, only type of B. concolor) and Borneo.

triangle = B. borneensis; square = B. bullatum

    Habitat & Ecology — In mixed Dipterocarp lowland forest, secondary forest, alluvial forest, mossy forest, primary upper montane forest, often along water and on very wet (but not inundated) soil; soil on (sandy) clay, igneous derived sandy clay. Altitude: 25–1700 m. Flowering: January; fruiting: February-May, July, September, October, December.

    Vernacular names — Kalimantan: Kapol utan; Simpul (Bassap Dyak); Sabah: Indalus (Dusun Kinabatangan), Tampoi (Malay).

    Notes — 1. Quite typical for B. borneense are the light coloured twigs, of which the upper ones are flattened or triangular, the leaves that dry light green, especially the lower surface, the almost fascicled flowers (one specimen in bud seen), and the thin-walled fruits (wall 2 mm at most). Blumeodendron borneense is only known from Borneo. In Borneo a group of specimens dries dark green and resemble B. borneense, but the leaves are alternate to (sub)opposite and seldom in whorls of 3 (end of branches), the flowers are along rachises and the fruits are very thick-walled (c. 5 mm).

    2. The type of B. concolor, Curtis KD 1368 (K) from the Malay Peninsula, strongly resembles B. borneense, as the leaves are of the same size, elliptic and also dry green, the flowers are in fascicles. However, the leaves are alternate. Another example of a specimen with leaves alternate (but close together) or in pseudo-whorls is a cultivated specimen from Borneo in the Bogor Botanical Garden, Gravendeel, de Wilde & Hovenkamp 521 (Kebun Raya IX.C.144). As the leaves in Bornean B. borneense can also be in pseudo-whorls B. concolor is regarded as a synonym of B. borneense.

    3. Specimens from Sarawak, including the type, and Brunei have a tendency to show ovate leaves, those of Sabah and Kalimantan are elliptic.

2. Blumeodendron bullatum Airy Shaw

    Blumeodendron (?) bullatum Airy Shaw, Kew Bull. 19 (1965) 310; Kew Bull. 27 (1972a) 86; Kew Bull., Addit. Ser. 4 (1975) 58; Ottens & Welzen, Blumea 61 (2016) 67, Fig. 2, Map 1. — Type: Haviland & Hose 3658 (holo K; iso BM, L), Sarawak, near Kuching.

Probably trees; branchlets generally round, nodes notably thickened to side of leaf; lenticels absent or indistinct; flowering branches c. 5 mm in diam. Indumentum of simple, white hairs and occasionally lepidote hairs on nodes. Leaves alternate to subopposite; petiole 0.5–1.6 cm long, diam. of thinnest part 0.2–0.5 cm, round, dull, basal pulvinus 0.4–0.5 cm diam., simple hairs present; blade elliptic, 6.1–22.3 by 3.8–10.3 cm, length/width ratio 1.6–2.2, coriaceous, symmetric, strongly bullate, drying brown, base rounded, margin strongly revolute, apex rounded, extrafloral nectaries along margin c. 12, basal ones usually not obvious; venation distinct, sunken above, strongly raised below, covered with simple hairs underneath and basal part of midrib above, marginal vein distinct, nerves 5–8 pairs, at 34–41˚ angle with midrib, tertiary nerves perpendicular to midrib and secondary nerves, distance between tertiary nerves 0.3–0.6 cm; higher order nerves scalariform to reticulate. Inflorescences axillary, very short thyrses, single, c. 0.8 cm long, flowers seemingly fasciculate, but peduncle c. 0.2 cm long, erect; bracts not well visible, c. 0.5 cm long. Staminate flowers c. 8.7 mm diam.; pedicel 3.5–5.5 mm long; buds c. 3.5 mm diam., covered with simple and lepidote hairs, sepals 2 or 3, elliptic, margin entire, c. 3.9–4.4 by 2.9–3.2 mm, valvate, free; stamens c. 33, filaments c. 4.7 mm long, anthers c. 0.6 mm long. Pistillate flowers, fruits and seeds unknown.

    Distribution — Borneo (Sarawak; only known from the type).

triangle = B. borneensis; square = B. bullatum

    Habitat & Ecology — Flowering: November.

    Note — Airy Shaw (1965) was not certain if this species belonged to Blumeodendron, but the absence of stipules, the ruminate staminate disc glands and the presence of orange lepidote hairs on the buds are all indicative of only Blumeodendron.

3. Blumeodendron endocarpum Ottens & Welzen

    Blumeodendron endocarpum Ottens & Welzen; Ottens & Welzen, Blumea 61 (2016) 70, Fig. 3, Map 2. — Type: BW (Kalkman) 6282 (holo L), [Indonesia, Papua,] Div. W. New Guinea, Beriat, c. 12 km S of Teminaboean.

    Blumeodendron kurzii auct. non (Hook.f.) J.J.Sm.: Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 37.

(Shrubs to) trees, to 40 m high, bole to 18 m high, d.b.h. to 40 cm diam.; buttresses sometimes present, to 1.5 m high, to 1.5 m wide, 4–10 cm thick; flowering branches 0.2–0.3 cm in diam., lenticellate, generally round, flat near petioles, distance between internodes usually c. 6 cm. Indumentum absent. Outer bark (light) (greyish) brown to dark brown to dark grey, smooth to shallowly fissured, strongly peeling with large scales, 0.25–0.5 mm thick; inner bark white, creamy, yellowish brown, orange to (light) brown, 4–5 mm thick; exudate sometimes present, light brown; sapwood white, pink or light brown; heartwood orange-brown to (light) brown, 5–12 cm diam. Leaves alternate to subopposite; petiole 2.3–9 cm long, diam. of thinnest part 1–2 mm, basal pulvinus 1.5–4.1 mm diam.; blade elliptic, 11.2–31 by 4.6–13.9 cm, length/width ratio 1.8–3, blade length/petiole length ratio 4.4–4.8, pergamentaceous (to coriaceous), often basally slightly asymmetric, glabrous, base broadly cuneate to attenuate, margin entire, revolute, apex acuminate (to cuspidate), tip round, both surfaces smooth, glabrous, green above, glossy light green to greyish green underneath; above drying greenish brown and brownish green underneath (greener than abaxially), extrafloral nectaries 2 adaxially near base, along midrib 6–15, along margin 8–36; venation: marginal nerve indistinct, secondary nerves (5 or) 6(–8) pairs, at c. 51.1˚ angle with midrib, tertiary nerves perpendicular to midrib and secondary nerves, not raised on both sides, c. 0.8 cm apart, higher order nerves reticulate, not raised on both sides. Inflorescences axillary and terminal, staminate ones mostly 2 per axil, thyrsoid, erect, to 2 cm long, with lepidote hairs. Staminate flowers 8–9 mm diam., white to (pale) yellow(-green); buds 14–30 per inflorescence; pedicel 8.5–17 mm long; sepals 3, ovate, c. 6 by 3 mm; stamens 35–43, filaments 4–5.5 mm long, anthers 0.5–1.1 mm long. Pistillate flowers only seen as young fruits; almost sessile, pedicel c. 2 mm long; ovary 2-locular; style c. 1 mm long; stigmas c. 5 mm long. Fruits flattened ellipsoid, slightly emarginate in the middle; 3.5 (1-seeded)–6 (2-seeded) cm wide by 3.6 (1-seeded)–3.9 (2-seeded) cm high, dry dull brown to whitish, vein ridges absent or indistinct; pedicel thickened up to 0.6 cm diam.; exocarp woody, c. 1 mm thick; surface smooth; endocarp around every seed, thickened to c. 4 mm, not dehiscing, probably rotting away on forest soil. Seeds flattened ovoid, backside more flattened than front side, top emarginate, 2.1–2.3 by 2.1–2.2 by 1.6–1.8 cm; sarcotesta thin, with distinct veins, testa woody, hard.

    Distribution —Sulawesi (Sulawesi Utara), Moluccas (Buru) and New Guinea (Indonesian Papua).

    Habitat & Ecology — Primary forest, seldom young secondary forest, along rivers and on slopes; soil mainly clay to sandy clay to sandy loam, never inundated. Altitude: 5–730 m. Flowering: February, March, September, October, December; fruiting: April, March, June–August, October–December.

    Vernacular names — New Guinea (Papua): Bobrijka, Sagogwo, Sagowgwo, Wobbrijka (Manikiong); Embaam, Embaan, M’baan (Itam); Josevakan (Tehid); Saba, Sawa (Mooi); Tiek (Kebar); Wotiet (Wandammen).

    Notes — Most typical for this species is a thick, woody endocarp around every seed instead around all seeds.

4. Blumeodendron gesinus Ottens

    Blumeodendron gesinus Ottens in Ottens & Welzen, Blumea 61 (2016) 71, Fig. 4, Map 3. — Type: Pereira, Wong, Sugau, Madani, Tangah, Nilus & Puff 158 (holo L; iso SAN), Malaysia, Borneo, Sabah, Tambunan, Jalan Pegalan.

Trees, to 65 m high, bole to 25 m high, d.b.h. to 70 cm; sometimes small buttresses present, c. 23 cm high, c. 30 cm out, c. 3.3 cm thick or fluted up to 1 m; crown spreading; flowering branches 2–3 mm diam., generally round, distance between internodes 2.5–5 cm, generally reddish brown and glabrous when dry, nodes slightly thickened. Outer bark smooth, (reddish) brown to brownish white to grey-brown to pink-brown, sometimes slightly hoop-marked, thin; inner bark chocolate brown to brownish to yellow(-green) to red(-brown) to orange-brown, c. 2.5 mm thick; sapwood white(-red) or pale yellow. Leaves always some alternate, but others subopposite to sometimes 3 in a pseudo-whorl; petiole 0.4–4.2 cm long, diam. of thinnest part 1–1.5 mm, basal pulvinus 1.1–3.3 mm diam.; blade elliptic, 4.6–15.3 by 2–7.4 cm, length/width ratio (1.3–)1.9–2.4(–3.3), coriaceous, symmetric, glabrous, base attenuate to cuneate, margin revolute, apex acuminate with rounded tip, extrafloral nectaries at base sometimes glandless to 1 visible to 2 present, 2-11 along midrib, 2–8 along margin, both surfaces smooth, dark green above when fresh, bright green underneath, drying greyish brown adaxially, abaxial surface orangish brown when dry; venation: marginal vein indistinct, secondary nerves (4–)6–8(–9) pairs, at c. 43.2˚ angle with midrib, tertiary nerves perpendicular to midrib, 1–3 mm apart, fourth order nerves often scalariform, higher order reticulate. Inflorescences axillary and terminal, thyrsoid, erect, staminate ones often 2 together, up to 11.5 cm long, flowers usually in 3-flowered cymes per node, pistillate inflorescences single, up to 3.5 cm long; bracts usually caducous, triangular, c. 0.8 by 0.2 mm, margin entire, with lepidote hairs. Staminate flowers 4.3–6.4 mm diam., yellow-green to yellow; buds 1.6–2.8 mm diam., often sticky with secretion; pedicel above abscission zone 3.1–3.3 mm long, c. 0.4 mm diam.; sepals 3(4), round, 4.5–5.3 by 2.5–3.8 mm long, completely recurved; stamens c. 27, filaments 2–4.5 mm long, anthers 0.4–0.7 mm long. Pistillate flowers 2.3–3 mm diam., pale green to yellow; sepals 5, ovate to triangular, very small, c. 0.5 by 0.5 mm, margin with few hairs; disc a c. 0.4 mm thick ring; ovary obovoid, c. 1.5–3.3 mm high by 2–3 mm wide, sutures thickened, 2(3)-locular, style 0.3–1 mm long, stigmas 1–2.8 mm long, curved horizontally, papillate above. Fruits ellipsoid to obovoid, c. 2.5–3.8 by 3–3.5 cm, probably indehiscent because loculicidal sutures with a rounded thickened rim, green when immature, orangish brown when dry; pedicel c. 4 mm long, c. 4.5 mm diam., abscission zone in the middle, upwards strongly widening; sepal remnants sometimes persistent; disc distinct, drying brownish; wall c. 2 mm thick, surface somewhat knobbly; endocarp enclosing two or more seeds; stigma usually persistent. Seeds ovoid but flattened on one side, not symmetric, attached subapically, 1.9–2.5 by 1.3–2.3 by 1–1.2 cm, sarcotesta thin, veins visible. 

    Distribution — Borneo.

    Habitat & Ecology — Lowland mixed dipterocarp forest to submontane, mossy forest, sometimes along roads in primary forest, seldom in logged over forest. Soil: yellow sandy, clay rich to ultrabasic; bedrock often sandstone. Altitude:25-1375 m. Flowering: February-April, June, August,. September; fruiting: February, May–August, October-December.

    Vernacular names — Borneo: Sabah: Tampoi (Melayu); Sarawak: Bantas (Iban), Bantas ketupong, Empungan (Iban).

    Notes — 1. Gesinus is the first name of the (first) author’s husband. The name is a personal name and not a latinisation, therefore ICN rule 60C.4 applies (http://www.iapt-taxon.org/nomen/main.php?page=art60), the name should not be changed into gesinum.

    2. This new species was generally confused with B. tokbrai, because of the long inflorescences. Very typical are the rims on the fruits and their orangish colour when dry. The same colour can be found on the lower surface of the dried leaves.

    3. The variation in inflorescence lengths seems to be large, but inflorescences with buds are not yet fully grown and will largely extend during maturation of the flowers. 

5. Blumeodendron kurzii (Hook.f.) J.J.Sm.

    Blumeodendron kurzii (Hook.f.) J.J.Sm., Meded. Depart. Landbouw 10 (1910) 463; Koord., Exkurs.-Fl. Java 2 (1912) 493; Pax & Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 48; Ridl., Fl. Malay Penins. 3 (1924) 281; M.R.Hend., J. Malayan Branch Roy. Asiat. Soc. 17 (1939) 69; Backer & Bakh.f., Fl. Java 1 (1963) 480; Airy Shaw, Kew Bull. 16 (1963) 348; Kew Bull. 26 (1972b) 224; Whitmore, Tree Fl. Malaya 2 (1973) 70, Fig. 2; Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 59; Kew Bull. 36 (1981) 269, Fig. 3B; Alphabet. Enum. Euph. Philipp. Is. (1983) 10; Chayam. in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 130; in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 611, Fig. 2; Ottens & Welzen, Blumea 61 (2016) 72, Fig. 5, Map 4. — Mallotus kurzii Hook.f., Fl. Brit. India 5 (1887) 427. — Lectotype (designated by Ottens & Welzen 2016): King’s collector 7114 (K), [Malaysia,] Perak, Larut. (Other syntypes: Helfer KD 5010 (K), Andaman Islands; Anonymous s.n., s.d. (K), [Malaysia,] Perak.)

    Blumeodendron verticillatum Merr., Philipp. J. Sci. 16 (1920) 557; Enum. Philipp. Fl. Pl. 2 (1923) 429; Pl. Elmer. Born. (1929) 157. — Type: FB (Meyer) 2603 (PNH, lost; iso L), Philippines, Luzon, Bataan Prov., Mount Mariveles. (NY noted Elmer 20815 as type, but this is incorrect, the specimen is not cited by Merrill, 1920).

    Blumeodendron sumatranum S.Moore, J. Bot. 63, Suppl. (1925) 102. — Syntypes: Forbes 1522 (BM, GH, 2 sheets, L, 3 sheets), Sumatra, Lampongs, Goenoeng Trang; Forbes 1563 (BM?, GH, L, 2 sheets), Sumatra, Lampongs, Hills NE of Goenoeng Trang; Forbes 1650a (BM, GH, L), Sumatra, Lampongs, Penanggoengan.

    Blumeodendron cuneatum S.Moore, J. Bot. 63, Suppl. (1925) 103. — Type: Forbes 2874 (holo BM; iso A, GH, L, 4 sheets), Sumatra, Palembang, Ayer Angat, foot of Kabo volcano.

Trees, to 35 m high, bole to 30 m high, d.b.h. to 60 cm; sometimes slightly fluted at base, flutes c. 1.5 m high, out 50 cm to sometimes a short buttress; flowering branches 4–22 mm in diam., round to sometimes triangular in section below the nodes, internodes up to 18 cm long; terminal bud surrounded by round or triangular axillary buds. Outer bark dark brown to brown-grey to greyish (black), smooth to cracked in irregular pieces to (powdery) scaly, soft to hard, 0.5-6 mm thick; inner bark beefy red outside to (pale) reddish to brown inside, 3-6 mm thick, sap absent to clear; sapwood white, yellow, reddish or brown; heartwood yellowish red to pinkish brown (to rays brown). Leaves always in whorls of 3–5 per node, young ones yellow-green to light green; petiole 1.3–12 cm long, diam. of thinnest part 1–4 mm, sordid green to brown, (green-brown to) dark brown when dry, generally darker than stem, basal pulvinus c. 4 mm diam., upper pulvinus larger and mainly developed abaxially; blade (ovate to) elliptic to oblong to obovate, 8.8–42(–51, see uses) by 4.4–23.7(–26.5) cm, length/width ratio 1.3–2.5, coriaceous, symmetric, glabrous, dried (dark green to) light brown on both sides, base obtuse to cuneate, margin usually light brown or yellow when dry, flat to revolute, apex acuminate (to cuspidate), extrafloral nectaries often 2, adaxially near base, at both sides along midrib 0–19 and along margin 6–36; venation: marginal vein distinct, secondary nerves (5 or) 6–12 pairs, well visible, 2/3 of length of nerves parallel with others, tertiary nerves perpendicular to midrib, hardly visible above, raised beneath, 0.3–0.4 cm apart, higher order nerves reticulate, indistinct. Inflorescences cauliflorous, ramiflorous, axillary and terminal, thyrsoid, almost fasciculate; staminate ones more than 8 together, up to 1.7 cm long, pistillate ones c. 4 together, up to 3 cm long; bracts absent. Flowers yellowish, yellow-green or light green; pedicel 0.4–1 cm long; staminate buds globose, c. 35 per inflorescence, c. 3.9 mm diam. Staminate flowers 7–7.5 mm in diam.; pedicel c. 10 mm long; sepals 3, 4–5 by 2.2–3.5 mm; disk lobes yellow; stamens 20–25, filaments 2–10 mm long, yellow, anthers 0.75–1 mm long, yellow to later fulvous. Pistillate flowers not seen; ovary 2- or 3-locular. Fruits capsular, subglobose,3.3–5.8 cm wide by 2.8–4.6 cm high, dry dark brown; pedicel thickened, c. 1 cm long, c. 6 mm diam., abscission zone in the middle to subapically; sepals usually persistent; disc distinct, drying dark brown; wall 2–4 mm thick, surface knobbly; margin not or slightly thickened as very low ridges; endocarp enclosing two or more seeds; style very sturdy, at most 1 mm long; stigmas usually persistent, up to 5 mm long, spreading. Seeds bean-shaped, but one end smaller than other, 2.1–2.2 by 1.3–1.6 by c. 1.2 cm, dark brown, attached in middle; sarcotesta yellow.

    Distribution — Peninsular Thailand, Malay Peninsula, Sumatra, Java, Borneo, Philippines (Luzon, Samar).

    Habitat & Ecology — Ranging from primary and evergreen forest to logged over and secondary forest (with bamboo); soil: often rich, varying from igneous derived sandy soil to sandy clay to loamy soil to limestone; bedrock once reported as basalt. Altitude: 5–600(–1800) m. Flowering: March–August, November–January; fruiting: March–June, August–December.

    Uses — Clemens 51511 is tentatively identified as B. kurzii. It is a single, enormous leaf, c. 51 by 26.5 cm, much larger than all other material of B. kurzii. The label indicates that large leaves are used by the Dusun in N Borneo as rain shelter and to repair leaks in roofs. The seeds are eaten in the Philippines.

    Vernacular names — Sumatra: Madanggadjah, Safanggeu bala, Tafanggeu, Tapanggeu delok, Tafanggeu toengo, Tampang. Java: Huru batu. Borneo: Kalimantan: Pelai, Tawiloeng; Sabah: Kulobon (Murut), Medang; Sarawak: Bantas (Iban), Ukut. Philippines: Kabarawang (Samar-Leyte Bisaya).

    Note — Typical are the brown drying leaves present in pseudo-whorls: leaves in a whorl but originating at slightly different levels.

6. Blumeodendron papuanum Pax & K.Hoffm.

    Blumeodendron papuanum Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xiv (6 June 1919) 14; Airy Shaw, Kew Bull. 16 (1963) 349. — Lectotype (designated by Ottens & Welzen 2016): Ledermann 9517 (K), New Guinea, Kaiser Wilhelmsland, Etappenberg. Other syntypes: Ledermann 8898, 9012, 9096 (K), New Guinea, Kaiser Wilhelmsland, Etappenberg.

    Bennettia papuana Gilg, Bot. Jahrb. Syst. 55 (13 Dec. 1918) 283, Fig. 6 (Flacourtiaceae); in Engl. & Harms, Pflanzenr. ed. 2, 21 (1925) 443, fig. 205. — Bennettiodendron papuanum (Gilg) Merr., J. Arnold Arbor. 8 (1927) 10. — Pimeleodendron papuanum auct. non Warb.: Sleumer, Fl. Males. ser. 1, 5 (1954) 65; Djarwaningsih, Blumea 49 (2004) 413, p.p.: Bennetia papuana. — Blumeodendron novoguineense Ottens & Welzen, Blumea 61 (2016) 74, Fig. 6a, b; Map 5, nom. illeg. — Type: Ledermann 8945 (holo B?, lost; iso K, L), [Papaua] New Guinea, Etappenberg. See note 1.

Trees, to 40 m high, bole to 33 m high, d.b.h. to 53 cm; bole sometimes fluted or with low buttresses up to 1.5 m high, out 0.6–2 m, 3–10 cm thick; flowering branches 2–3 mm diam., generally round, with orange lepidote hairs, early glabrescent, long internodes up to 8.5 cm. Outer bark red-brown to grey-brown to dark brown to brownish black, smooth to pustular lenticellate, not fissured nor peeling to little peeling with small to large scales, 0.25-0.5 mm thick; under bark wine-red; inner bark yellow to yellowish brown to red to light to dark brown, 3-12 mm thick; sapwood white to orange-brown to reddish brown to light brown; heartwood light brown to black. Leaves on hardly widened nodes at end of short nodes, alternate to subopposite to in pseudo-whorls of 3 leaves; petiole 1–4.1 cm long, diam. of thinnest part 0.8–1.2 mm, round, basal pulvinus 1.3–2 mm diam., fast fading orange lepidote hairs; blade (ovate to) elliptic, 5.3–17.5 by 2.5–9 cm, length/width ratio 1.5–2.6, pergamentaceous to coriaceous, symmetric, glabrous, base (broadly) cuneate, margin recurved, apex acuminate, tip rounded, extrafloral nectaries on both surfaces along midrib, 2 to many, along margin several, both surfaces smooth, mid to dark green when fresh, dull and lighter beneath, drying brownish green, slightly darker brown underneath; venation slightly raised on both sides, marginal vein indistinct, secondary nerves pinnate, 5–8 pairs, sometimes very parallel, at c. 52˚ angle with midrib, tertiary nerves more or less scalariform, perpendicular to midrib, higher order nerves indistinct, reticulate. Inflorescences axillary, mostly single, staminate rachises up to 9.5 cm long, 1–1.3 mm diam., pistillate ones up to 3 cm long in flower, up to 9 cm when in fruit, 1–1.5 mm diam., thickening during fruit set to c. 2 mm; bracts vestigial; flowers single per node (young additional buds can be present when staminate). Staminate flowers c. 7.5 mm diam., white, sweet scented; pedicel 2–5 mm long, 0.8–1 mm diam.; sepals 3 or 4, ovate, 3.8–5 by 2.5–3.5 mm, green to yellow; disc lobes yellow; stamens c. 40, filaments 3–8.3 mm long, white, anthers 0.4–0.6 mm long. Pistillate flowers 3.8–6 mm diam., green; pedicel c. 3.3 mm long; sepals 3–5, triangular, 1.6–3.1 by 1.4–2 mm, recurved; ovary 2- (or 3-)locular, ellipsoid, c. 3 by 3 mm; style c. 0.8 mm long, sturdy, stigmas 2–5 mm long, recurved. Fruits capsular, flattened obovoid, angular (perhaps not ripe yet) with often slightly raised suture, 2–2.9 cm wide by 2–2.9 cm high; pedicel c. 3 mm long, abscission zone subbasally; wall 1–1.8 mm thick, brown when dry, surface somewhat rugose; endocarp enclosing all seeds; stigma mostly persistent. Seeds bean-like to flattened at one-side, c. 1.9 by 1.3 by 1 cm, attached in middle, black.

    Distribution — New Guinea.

    Habitat & Ecology — Lowland rain forest to Araucaria-Anisoptera forest at higher altitudes; soil: clayey, sandy clay, loam, broken lava, can be inundated in the wet season. Alt.: 8–800 m. Flowering: March–June, August, September, November; fruiting: January, March, July–September, November, December.

    Vernacular names —New Guinea: Papua: New Guinea: Arom (Hattam); Kem (Mooi); Lowkwa (Manikiong); Manaper/Manapper/Manapir (Biak); Moe-e (Tor); Moentawiempi, Moentawinakpopi (Roberbai, Japen dialect); Moëre (Wain); Mwer (Berik); Sehoi/Sohoi (Manikiong); Tabet (Moejoe); Tajapmoetop (Mandobo); Wobbrijka (Manikiong); Wonsoka (Arfak, Sidai dialect); Papua New Guinea: Akop; He-arahai (Mangalese, Bariji dialect). 

    Wood — Sapwood not defined from heartwood, white to straw-coloured, close grained, medium hard to hard, medium heavy to heavy. Pores few, small, visible to naked eye, short radial chains. Rays fine, barely visible to the naked eye. Parenchyma in numerous fine bands. (NGF (Mair) 547; NGF (Havel & Kairo) 17205).

    Notes — 1. The epithet papuana by Gilg is the oldest on species level, however, within Blumeodendron the combination already existed (B. papuanum Pax & K.Hoffm; Pax & Hoffmann 1919). Then, if no other epithet is available, Gilg’s name has to receive a new name within Blumeodendron, as we did (Ottens & Welzen 2015) incorrectly, because papuanum by Pax & Hoffmann (1919) is available.

    2. This species closely resemblances B. tokbrai, but differs in the size and form of the fruits, smaller (2–2.9 by 2–2.9 cm versus 2.3–4.8 by 2.3–4.1 cm) and often angular and with slightly thickened sutures (versus round, without thickened sutures), the thickness of the fruit wall is thinner (1–1.8 mm versus 4–7 mm) and the presence of broader sepals in the pistillate (1.4–2 mm versus 0.5–1.1 mm) and staminate flowers (2.5–3.5 by 1–2 mm). Moreover, the leaves of B. papuanum usually dry brownish green, which also occurs in B. tokbrai, but there most dry leaves are dark brown. 

7. Blumeodendron philippinense Merr. & Rolfe

    Blumeodendron philippinense Merr. & Rolfe in Merr., Philipp. J. Sci. 16 (1920) 555; Merr., Enum. Philipp. Fl. Pl. 2 (1923) 429; Ottens & Welzen, Blumea 61 (2016) 76, Fig. 7, Map 6. — Type: FB (Topacio) 20054 (holo PNH, lost; iso L), Philippines, Luzon, Bataan Prov., Mount Mariveles.

Trees, to 15 m high, d.b.h. to 45 cm; flowering branches c. 5 mm in diam., generally round, with fast fading orange lepidote hairs, internodes c. 4 cm long. Leaves alternate; petiole 3.9–8.9 cm long, diam. of thinnest part1.5–2 mm, round with groove above, basal pulvinus c. 3 mm diam., with lepidote hairs, orange, early caducous; blade ovate, 7.8–21 by 3.7–10.2 cm, length/width ratio 2–2.3, coriaceous, symmetric, glabrous, base broadly cuneate to attenuate, margin recurved, apex acuminate, both surfaces smooth, glossy, upper dark green, lower pale green, adaxial surface browner than abaxial surface when dry, extrafloral nectaries along margin 6–7; venation: marginal vein indistinct, secondary nerves 5 or 6 pairs, usually parallel, at c. 33˚ angle with midrib, tertiary nerves perpendicular to midrib and/or only to secondary nerves, higher order nerves indistinct. Inflorescences terminal and axillary, only consisting of a up to 1.3 cm long rachis, staminate ones 3 or 4 together, pistillate ones single or 2 per node; latter inflorescences thickening after fertilization, to 2.5 mm diam.; lepidote hairs present, orange. Staminate flowers seen in bud: pedicel c. 3 mm long; buds c. 4 mm diam., c. 6 per inflorescence; rest unknown. Pistillate flowers seen in fruiting stage; upper part of pedicel above abscission zone c. 3 mm long; sepals 5, ovate, c. 2 by 2 mm; disc very c. 1 mm high; ovary 2- or 3-locular, style c. 0.8 mm long, stigmas c. 3 mm long. Fruits subglobular, c. 2 per inflorescence, up to 3 cm in width when immature, green (probably unripe), surface knobbly; pedicel c. 6 mm long , to c. 3.5 mm diam., abscission zone basally; wall c. 1 mm thick, drying brown; endocarp enclosing two or more seeds; stigma mostly persistent. Seeds not seen mature. 

    Distribution — Philippines, endemic on Luzon in Bataan Prov.

triangle = B. philippinense; dot = B. subrotudnifolium

    Habitat & Ecology — Secondary forest on a slope with brown soil. Altitude: c. 650 m. Flowering: January, October; fruiting: May, December.

    Note — Distinctive for this species are the alternate, coriaceous leaves and the very short inflorescence. The short inflorescence are reminiscent of B. kurzii (leaves in whorls) and alternate leaves are found in various species, but all with longer inflorescences except for B. borneense from the Malay Peninsula, but the latter form has light green dried leaves instead of brown dry leaves.

8. Blumeodendron subrotundifolium (Elmer) Merr.

    Blumeodendron subrotundifolium (Elmer) Merr., Philipp. J. Sci. 7 (1912) 384; Philipp. J. Sci. 16 (1920) 558; Enum. Philipp. Fl. Pl. 2 (1923) 429; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 49; Whitmore, Tree Fl. Malaya 2 (1973) 70, Fig. 2; Ottens & Welzen, Blumea 61 (2016) 77, Fig. 8, Map 6. — Sapium subrotundifolium Elmer, Leafl. Philipp. Bot. 3 (1910) 930 (‘subrotundifolia’). —Type: Elmer 12349 (holo PNH, lost; iso A, BISH, G, GH, HBG, K, L, NY, US), Philippines, Sibuyan, Capiz Prov., Magallanes (Mt. Giting-Giting).

    Blumeodendron calophyllum Airy Shaw, Kew Bull. 19 (1965) 309; Kew Bull. 25 (1971) 518; Whitmore, Tree Fl. Malaya 2 (1973) 70, fig. 2; Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 58. — Type: S (Brunig) 8867 (holo K; iso L), Sarawak, Bintulu Dist., Niah-Jelalong primary forest.

    Blumeodendron subcaudatum Merr., Philipp. J. Sci. 16 (1920) 557; Enum. Philipp. Fl. Pl. 2 (1923) 429. — Lectotype (designated by Ottens & Welzen 2016): FB (Sherfesee, Cenabre & Cortes) 21075 (holo K; iso A, US ), Samar.

Trees, to 50 m high, bole to 25 m high, d.b.h. to 91 cm; bole sometimes fluted or with low buttresses up to 1.5 m, out c. 1.5 m, c. 2.5 cm thick; flowering branches 3 (staminate)–28 (pistillate) mm diam., generally round, with orange lepidote hairs, early glabrescent, internodes up to c. 5 cm. Outer bark brown to yellow-brown to grey-brown to yellow-grey to grey (to greyish green), fissured to scaly to flaky, soft, lenticellate, 1–3 mm thick; inner bark red, light or reddish brown or dark brown with yellow and light brown spots (laminated), hard; 8–10 mm thick; sometimes exudate reported, red, watery; sapwood cream to white (with pinkish tinge radially), yellow or light reddish brown, very hard; heartwood brown. Leaves alternate to subopposite to in pseudo-whorls; petiole 2.4–18.5 cm long, diam. of thinnest part 1–15 mm, round, basal pulvinus 2–20 mm diam., fast fading orange lepidote hairs; blade elliptic, 6.2–46 by 3–22 cm, length/width ratio 1.2–3.1, coriaceous (slightly bendable) to very coriaceous (breaking), symmetric, glabrous, base emarginate to rounded to cuneate, margin recurved, apex acuminate (to cuspidate), both surfaces smooth, extrafloral nectaries on both surfaces along midrib c. 2–26, along margin c. 8–20, adaxial surface usually drying shiny brown, abaxial surface lighter dull brown; venation: marginal vein indistinct, secondary nerves pinnate, 3–4(–11) pairs, sometimes very parallel, at c. 52˚ angle with midrib, tertiary nerves raised below, perpendicular to midrib and/or only to secondary nerves, higher order nerves indistinct. Inflorescences cauliflorous, ramiflorous to axillary, c. 0.1 cm diam., staminate ones often paired, up to 4.5 cm long, pistillate ones single, up to 2.3 cm long, rachis thickening during fruit set to c. 4 mm, peduncle c. 1.3 cm long, brown; bracteoles triangular, c. 0.5 by 0.2 cm, margin undulate; lepidote hairs orange. Flowers pale yellow to yellow green to yellowish tinged pink to (greenish) red. Staminate flowers 5–6 mm diam.; pedicel 3–25 mm long, brown; buds 1.5–5 mm diam.; sepals 2 or 3, ovate, 4–5.5 by 3–5 mm; disc lobes yellow; stamens 25–40, yellow, filaments 1.5–3 mm long, anthers 0.4–0.5 mm long. Pistillate flowers not seen; sepals 4 or 5, c. 1.5 by 2.2 mm; ovary 2- or 3-locular, dull sordidly purple; style c. 1 mm long, sturdy, stigmas c. 2.5 mm long, recurved, greenish. Fruits capsular, subglobular (smaller) to ovoid (larger), 3.5–6 cm wide by 2.9–4.5 cm high, green (unripe) to yellow to orange-yellow (or red); pedicel c. 1.5 cm long, to c. 0.8 cm diam., abscission zone in the middle; wall 2–4 mm thick, dark brown when dry, margin slightly thickened, but without ridges, surface knobbly; endocarp enclosing two or more seeds; stigma mostly persistent. Seeds bean-like to flattened at one-side, 2.3–3.4 by 1.6–2.4 by 1.1–1.3 cm, attached in middle; sarcotesta cream to purple.

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

triangle = B. philippinense; dot = B. subrotudnifolium

    Habitat & Ecology — Mixed lowland dipterocarp forest, evergreen forest, gallery (Emperan) forest, peat swamp forest, to mossy submontane forest, along logging roads; soil sandy loam, sandy clay, clayey loam, alluvial soil, bedrock: sandstone. Altitude: sea level to 1200 m. Flowering: March–November; fruiting: January, April–November. Fallen seeds / fruits are eaten by birds and animals (e.g., pigs).

    Vernacular names — Malay Peninsula: Gaham badak; Kaum Bada. Sumatra: Babak; Basi; Madang soenting; Makoera; Mamboeloeh; Medang koenik; Oendal; Sikoe kaloeang; Tendal (Malay); Toetoen sijeureuh, Toetoen sijeureuh etem, Toetoen sijeureuh pajo. Borneo: Anambas & Natuna Islands: Medang keladi; Kalimantan: Duhat (Malay); Kahingai; Sibau; Sarawak: Bantas, Bantas belulang (Iban); Belulang; Berti-an (Kenyah); Empungan (Malay Sarikei); Marahbulan; Ngisigi (Land Dayak); Teku (Malay). Philippines: Halilimokon (Samar); Mangamit.

    Notes — 1. Blumeodendron subrotundifolium resembles B. kurzii in leaf-shape, colour of dried leaves and leaf texture. However, the leaves do not arise in whorls from thickened nodes. Also, the inflorescences are generally longer than the almost fasciculate ones of B. kurzii.

    2. Blumeodendron calophyllum is added here as a synonym of B. subrotundifolium. Most specimens can easily be divided over both species as they look spectacularly different. Small, coriaceous leaves with slender petioles and more axillary smaller fruits are present in typical B. subrotundifolium, while much larger, very coriaceous (not bendable) leaves with thick petioles and cauliflorous large fruits are found in B. calophyllum. However, quite a number of specimens bridge the gap between both typical forms (see table 1). Both forms are generally high trees with (very) coriaceous leaves on relatively long petioles, which dry dark shiny brown above and dull brown underneath. Leaf sizes also vary strongly in B. kurzii, a species with which B. subrotundifolium was often confused.

9. Blumeodendron tokbrai (Blume) Kurz

    Blumeodendron tokbrai (Blume) Kurz, J. Asiat. Soc. Bengal 42 (1873) 245; Forest Fl. Burma 2 (1877) 391 (pro Mallotus tokbrai (Blume) Müll.Arg.); emend. J.J.Sm., Meded. Depart. Landbouw 10 (1910) 460; Koord., Exkurs.-Fl. Java 2 (1912) 493; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 48; Pflanzenr. IV.147.xiv (1919) 14; Merr., J. Straits Branch Roy. Asiat. Soc., Spec. No. (1921) 340; Backer & Bakh.f., Fl. Java 1 (1963) 479; Whitmore, Tree Fl. Malaya 2 (1973) 71, Fig. 3; Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 60; Kew Bull., Addit. Ser. 8 (1980) 38; Kew Bull. 36 (1981) 269, Fig. 3A; Kew Bull. 37 (1982) 9; Alphabet. Enum. Euph. Philipp. Is. (1983) 10; Chayam. in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 131; in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 612, Fig. 3; Ottens & Welzen, Blumea 61 (2016) 78, Figs. 6c, 9, Map 7. — Elateriospermum tokbrai Blume, Bijdr. Fl. Ned. Ind. 12 (1826) 621; Hassk., Pl. Jav. Rar. (1848) 251; Miq., Fl. Ned. Ind. 1, 2 (1859) 412. — Mallotus tokbrai (Blume) Müll.Arg. in DC., Prodr. 15, 2 (1866) 956; Pax in Engl., Pflanzenr. IV.147 (1910) 18. — Rottlera tokbrai (Blume) Scheff., Ann. Mus. Bot. Lugduno-Batavi 4 (1869) 122. — Blumeodendron elateriospermum J.J.Sm., Bull. Jard. Bot. Buitenzorg, sér. 2, 8 (1912) 56, nom. illeg., superfl. — Lectotype (designated by Ottens & Welzen 2016): Blume 1531 (L), Java, Mt. Salak.

    Mallotus ? vernicosus Hook.f., Fl. Brit. India 5 (1887) 443. — Blumeodendron vernicosum (Hook.f.) Gage, Rec. Bot. Surv. India 9 (1922) 244; Ridl., Fl. Malay Penins. 3 (1924) 282. — Type: Cantley 9 (K), Singapore, Botanical Garden.

    ? Elateriospermum paucinervia Elmer, Leafl. Philipp. Bot. 2 (1908) 484. — Blumeodendron paucinervium (Elmer) Merr., Philipp. J. Sci. 16 (1920) 555; Enum. Philipp. Fl. Pl. 2 (1923) 428. — Type: Elmer 7416 (holo PNH, lost; iso L), Philippines, Luzon, Tayabas Prov., Lucban.

rees, to 40 m high, bole to 25 m high, d.b.h. to 100 cm; stilt roots or buttresses up to 3 m high, out to 3 m; flowering branches 2‒5.5 mm diam., often somewhat angular, with internodes 3‒7 cm long; nodes hardly thickened. Outer bark red-, greenish- or light brown to (dark) grey to red to orange(-brown), smooth, pustular or fissured, lenticels round, c. 0.5 mm thick; inner bark wine- or orange red to yellow brown, brown, (ochre-)orange, white, purple, yellow-pink and sometimes mottled, 5-10 mm thick; exudate indistinct, but plants becoming sticky; sapwood straw to pale yellow to cream to greyish white; heartwood straw. Leaves usually always a few alternate, but also subopposite to 3 leaves whorled; petiole 1.2–9.4 cm long, diam. of thinnest part 1–2 mm, basal pulvinus 1–3.6 mm diam., hairs present (see note 2); blade (ovate to) elliptic (to obovate), 5.3–31 by 3.1–17.3 cm, length/width ratio 1.4‒2.4(‒3.3), ratio leaf length/petiole length 4.4–4.8, pergamentaceous to coriaceous, asymmetric, with simple and lepidote hairs when young, base attenuate to cuneate, margin slightly recurved, apex acuminate to cuspidate, tip rounded to mucronulate, extrafloral nectaries sometimes adaxially along midrib, c. 10, young leaves resinous, covered with orange lepidote hairs, surfaces drying brown to brown-green to dark green, smooth, abaxial surface browner than adaxial surface when dry; venation slightly raised above, marginal vein indistinct, secondary nerves 5‒9 pairs, at c. 45.6˚ angle with midrib, tertiary nerves perpendicular to midrib, c. 2 mm apart, distinct, raised beneath, higher order nerves reticulate, indistinct. Inflorescences axillary and terminal, staminate ones 1–3 together, to 20 cm long, c. 1.4 mm diam., pistillate ones single, to 10 cm long, c. 1 mm diam.; lepidote hairs orange, simple hairs white. Buds c. 3 mm diam. Staminate flowers 4–7.7 mm diam., white to pale green, sweet scented; pedicel 5.8‒8.4 mm long, 0.3‒0.7 mm diam.; sepals 3 or 4, ovate to elliptic, 2.5‒4 by 1‒2 mm, inside red; stamens 31–36, filaments c. 4 mm long, white, anthers c. 0.5 mm long, yellow. Pistillate flowers 0.9–2.3 mm diam., light green; pedicel c. 2.5 mm long, c. 0.8 mm diam.; sepals 5, triangular to ovate, 1.5‒3 by 0.5‒1.1 mm, inside red; ovary 2- or 3(4)-locular, ellipsoid, 1.7–2.3 mm high, 1.4–2.3 mm diam.; style indistinct, 0.3–0.5 mm long, stigma 1.3–4 mm long, recurved. Fruits ellipsoid to flattened-globular, 3.3–4.8 cm broad by 2.3–4.1 cm high, green to red-brown when dry; pedicel to 1 cm long, up to 3 mm diam., abscission zone basally; sepals not persistent; disc distinct; wall 4–7 mm thick, meso- and endocarp thickened, mesocarp sometimes with cavities when dry (see note 3), surface knobbly; margin mostly pitted when dry; endocarp enclosing 2 or more seeds; stigma usually long persistent. Seeds bean-shaped, 2–3.8 by 1.1–2.2 by 0.9–1.3 cm; sarcotesta yellow.

    Distribution — Thailand, Malay Peninsula, Sumatra, Java, Borneo, Philippines, Sulawesi, Moluccas.

    Habitat & Ecology — Primary dipterocarp lowland forest, peat swamp forest, mangrove, kerangas forest, riverine forest, secondary forests; soils: white sand, sandy clay, clay, sandstone. Altitude: sea level to 1400 m. Flowering and fruiting throughout the year.

    Uses — Sarawak (Borneo): Wood used for canoes and planks; fruits edible.

   Vernacular names — Malay Peninsula: Gaham badak; Kaum bada; Marahbulan. Sumatra: Awa sijeureuh; Batin-batin oeding; Belanti; Beroewa babi; Kalek kasih; Kemili oétan (Malay-Palembang) ; Keteroeng; Lala lalar oeding; Matakoeroeng; Niho (Enggano); Lala lalar oeding; Matakoeroeng; Oekih datan; Panai-panai; Sijeureuh etem; Sijeureuh-pajo; Sijeureuh silai; Sijoeroeh alafai; Tekoeroeng; Tekoeroeng keteroeng; Tijeureuh silai; Toetoen ramboetan dotan. Java: Boerahol; Kendoeng leuweung; Ki tokbraay; Tokbray. Sangi and Talaud Isles: Aluwatu. Borneo: Brunei: Antangon (Iban); Kalimantan: Sibau; Sabah: Gangulang; Gulang gulang; Tombuakat; Sarawak: Bantas (Iban); Buan (Kayan); Empungan (Milanau); Marahbulan; Merbulan; Merahbulan; Oendal; Pelapi (Kayan); Selunsor merah; Takok (Melanau); Teku. Sulawesi: Talaud: Aluwatu.

    Notes — 1. Blumeodendron novoguineense and B. tokbrai resemble each other closely, see note 2 under former.

    2. Simple hairs are usually visible in Sumatran collections.

    3. Some specimens in Borneo dry with green leaves and the fruits are very thick, not only the endocarp is thick also the mesocarp. The mesocarp then contains cavities of which is unclear if these were formed during drying of the fruits. Examples are: A (Wood) 4815, S (Paie) 16992, S (Au) 23937, SAN (Gansau) 47750. Probably S (Anderson & Paie) 28338, with staminate buds, and Ambrianyah & Arifin W 807, with young fruits, also belong to this form. Endert 4029, from SE Kalimantan, dried brownish green and also has fruits – still young – with a thickened mesoderm, but without cavities.

    4. Formerly, no distinction was made between B. tokbrai and B. papuanum. The geographical border between both species is between the the Moluccas (B. tokbrai; fruits larger and thick-walled) and New Guinea (B. papuanum; fruits smaller and thin-walled). The lack of delimitation may be due to the fruits of B. tokbrai, which show a geocline, they are larger in the west (Malay Peninsula and Sumatra) and smaller towards the east (Philippines, Moluccas); it seems that this trend is continued in New Guinea.

EXCLUDED TAXON

    Blumeodendron muelleri Kurz, J. Asiat. Soc. Bengal 42 (1873) 245, nom. superfl. (presented as new name for Paracroton pendulus) = Paracroton pendulus (Hassk.) Miq.