Malesian Euphorbiaceae Descriptions

12. BACCAUREA (Phyllanthaceae)

R.M.A.P. Haegens

Haegens, R.M.A.P. 2000. Taxonomy, Phylogeny, and Biogeography of Baccaurea, Distichirhops, and Nothobaccaurea (Euphorbiaceae). Blumea Suppl. 12: 1–216.

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Genus description

Key to the Malesian Species

Key to the Species of the Philippines (excluding Palawan)

Key to the Species of Sulawesi and the Moluccas

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Species descriptions

Baccaurea Lour.

Baccaurea Lour., Fl. Cochinch. (1790) 661; Müll.Arg. in DC., Prodr. 15, 2 (1866) 456; Bedd., Fl. Sylv. S. India (1869) 280; Kurz, Forest Fl. Burma 2 (1877) 356; Benth. & Hook.f., Gen. Pl. 3 (1883) 283; Hook.f., Fl. Brit. India 5 (1887) 367; Pax in Engl. & Prantl, Nat. Pflanzenfam. 3, 5 (1891) 30; Brandis, Indian Trees (1906) 562; J.J.Sm., Meded. Dept. Landb. Ned.-Indië 10 (1910) 245; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 45; Ridl., Fl. Malay Penins. 3 (1924) 242; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1927) 547; Merr., Trans. Amer. Philos. Soc. New Ser. 24, 2 (1935) 29, 47; Corner, Wayside Trees Mal. 1 (1940) 238; Backer & Bakh.f., Fl. Java 1 (1964) 453; Whitmore, Tree Fl. Malaya 2 (1973) 63; A.C.Sm., Allertonia 1 (1978) 377; Fernando, Philipp. J. Biol. 8 (1979) 301; Airy Shaw, Kew Bull. Add. Ser. 8 (1980) 34; Kew Bull. 36 (1981) 258; G.L. Webster, Ann. Missouri Bot. Gard. 81 (1994) 51; Haegens & Welzen in Welzen et al., Thai For. Bull. 28 (2000) 68; Haegens, Blumea Suppl. 12 (2000) 81; Radcl.-Sm., Gen. Euphorbiacearum (2001) 62; Haegens & Welzen in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 107; G.L.Webster in Kubitzki, Fam. Gen. Vasc. Pl. 11 (2014) 86. — Lectotype (designated by Merrill, 1935): Baccaurea ramiflora Lour.

Pierardia Roxb. ex Jack, Trans. Linn. Soc. London, Bot. 14 (1823) 119; Blume, Bijdr. Fl. Ned. Ind. (1825) 578; Roxb., Fl. Ind. 2 (1832) 254; Endl., Gen. Pl. 2 (1839) 1124; Walp., Repert. Bot. Syst. 5 (1846) 366; Griff., Ic. Pl. Asiat. (1854) 738; Wight, Icon. Pl. Ind. Orient. (1852) 30; Miq., Fl. Ned. Ind. 1, 2 (1859) 358; Fl. Ned. Ind., Eerste bijv. (1861) 441; Baill., Adansonia 3 (1863) 139. — Type: Pierardia dulcis Jack [= Baccaurea dulcis (Jack) Müll.Arg.].

Adenocrepis Blume, Bijdr. Fl. Ned. Ind. (1825) 579; Endl., Gen. Pl. 2 (1839) 1123; Baill., Adansonia 3 (1863) 139; Müll.Arg. in DC., Prodr. 15, 2 (1866) 465. — Type: Adenocrepis javanica Blume [= Baccaurea javanica (Blume) Müll.Arg.].

Microsepala Miq., Fl. Ned. Ind., Eerste bijv. (1861) 444. — Type: Microsepala accuminata Miq. [= Baccaurea javanica (Blume) Müll.Arg.].

Calyptroon Miq., Fl. Ned. Ind., Eerste bijv. (1861) 471; Müll.Arg. in DC., Prodr. 15, 2 (1866) 466. — Type: Calyptroon sumatranum Miq. [= Baccaurea sumatrana (Miq.) Müll.Arg.].

Everettiodendron Merr., Philipp. J. Sci., Bot. 4 (1909) 279. — Type: Everettiodendron philippinense Merr. [= Baccaurea philippinensis (Merr.) Merr.].

Gatnaia Gagnep., Bull. Soc. Bot. France 71 (1924) 870; Airy Shaw, Kew Bull. 14 (1960) 353. — Type: Gatnaia annamica Gagnep. [= Baccaurea ramiflora Lour.].

Shrub to tree, dioecious, with Terminalia branching pattern. Indumentum of simple and often also of stellate hairs. Bark ﬁnely ﬁssured. Leaves simple, alternate, spirally arranged; petiole apically and rarely basally pulvinate, transverse cracks often present; stipules triangular or rarely foliaceous, early to late caducous, often ciliate; blade basally cordate to attenuate; margin entire, marginal glands present; apex usually rounded to cuspidate to rarely retuse; upper surface glabrous, sometimes densely hairy on the veins, raised glands often present, sometimes granulate; lower surface glabrous to densely hairy, raised glands often present, discoid glands sometimes present; nervation raised, secondary veins curved and sometimes closed at margin. Inflorescences axillary to cauline, reduced thyrses, solitary to many clustered together; flowers hypogynous, actinomorphic; pedicel with abscission zone. Staminate inflorescences not to distinctly branched, 10–many-flowered, flowers scattered along inflorescence to clustered at the tip; bracts triangular to rarely (broadly) ovate; branchlets 1–20 (in B. bracteata to 40)-flowered. Staminate flowers: sepals (3 or) 4 or 5(–8), free to slightly fused at base; petals absent; disc absent; stamens 3–10, shorter than sepals, glabrous; anthers introrse, basiﬁxed to dorsiﬁxed, opening with a longitudinal apical slit; staminodes (or disc glands) sometimes present, 3–8; pistillode usually present, small. Pistillate inflorescences not branched, one- to many-flowered; bracts inserted on rachis to usually along pedicel, 1 or 3 per flower, triangular to ovate. Pistillate flowers: sepals 4–6 (or in B. papuana up to 9), caducous to persistent; petals absent; staminodes absent; disc absent; ovary cylindrical to globose, 2–4-locular, ovules 2 per locule, attached at apex of column, wings usually absent; stigmas not cleft to completely divided into 2 lobes; protuberances (see Fig. 1.9) often present. Fruits berries to late dehiscing fleshy capsules; raised glands often present; seeds 0–8, 0–2 per locule, ellipsoid but laterally flattened; arillode enclosing seed totally; cotyledons flat, rarely folded.

Distribution — Forty-three species in India, SE Asia and the Paciﬁc.

Habitat & Ecology — Primary to secondary rain forest or freshwater swamp forest. Soil: sand to clay to loam. Altitude: sea level up to 1600(–1800) m, usually in lowland. Flowering and fruiting throughout the year.

Uses — Arillode of all and pericarp of some species edible, sour to sweet.

1. Baccaurea angulata Merr.

Baccaurea angulata Merr., Univ. Calif. Publ. Bot. 15 (1929) 148; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 35; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 46; Haegens, Blumea Suppl. (2000) 96, Fig. 3.5, Map 3.1, Photo 1. — Type: Elmer 21842 (holo A; iso B, DS, G, L, U), Borneo, Sabah, Tawau.

Tree 6–21 m high, dbh 9.6–40 cm, buttresses absent; branchlets glabrous; young shoots pale straw when fresh, green-grey to blackish when dry; Terminalia branching pattern weakly developed. Indumentum of simple hairs. Bark grey-brown to red-brown to green when fresh, grey when dry, smooth to rough, flaky, soft to hard, corky; inner bark yellow to reddish to green, 0.6–4 mm thick. Heartwood brown. Leaves: petiole 20–125 mm long, glabrous, brown to grey to greenish when dry, raised glands usually present; stipules 4–11 by 2–5 mm, glabrous to densely hairy outside, glabrous inside, margin ciliate, not hyaline; lamina elliptic to obovate, 12–39 by 4–13.6 cm, l/w ratio 2–4.4, thick leathery; base cuneate to attenuate (rounded); apex acuminate to cuspidate to rarely obtuse, (0)3–22 mm long; upper surface glabrous, raised glands absent, usually not granulate, dull to glossy dark green when fresh, grey to green to rarely brown when dry; lower surface glabrous, sometimes subglabrous at midrib, discoid glands absent, pale green with whitish midrib below when fresh, grey to green to rarely brown when dry; secondary veins 9–16 per side, closed at margin; nervation reticulate, not clearly visible. Staminate inflorescences cauline, few clustered together, 0.5–23 cm long, c. 1 mm thick, sparsely hairy, branching minute, up to 50-flowered, flowers scattered along inflorescence, yellow; bracts 1 per branchlet, 1–1.5 mm long, densely hairy outside, glabrous inside, margin ciliate, not hyaline; bracteoles c. 0.5 mm long, 2 per cymule; cymules cylindrical, c. 0.5 mm long, densely hairy, 3-flowered. Staminate flowers 2–2.6 mm diam.; pedicel 2–4 mm long, upper part 1.2–2.5 mm long, densely hairy; sepals 4 or 5, obovate, 1.1–1.4 by 0.6–0.7 mm, apex slightly recurved, outside and inside densely hairy; staminodes 6; stamens 6, 0.6–0.8 mm long, glabrous; ﬁlaments 0.5–0.7 mm long, straight; anthers 0.1–0.2 by 0.2–0.25 by c. 0.1 mm; disc absent; pistillode obtriangular, c. 0.3 mm high, densely hairy, hollow. Pistillate inflorescences cauline to ramiflorous, solitary or up to 7 clustered together, 4–25 cm long, 1–3 mm thick, subglabrous to densely hairy, 8–many-flowered, red; pedicel 2.5–5.5 mm long, upper part thickened, 0.5–1 mm long, sparsely to densely hairy; bracts 3 per flower, sparsely to densely hairy outside, glabrous inside, margin ciliate. Pistillate flowers 4–10 mm diam., pale yellow to cream yellow to greenish; sepals 5 (or 6), elliptic, (3.5–)8–11 by 1–2 mm, outside and inside densely hairy, persistent to caducous; ovary urn-shaped, 3–6 by 1.5–3 mm, 3-locular, densely hairy, wings 6; style absent; stigmas 0.7–1.5 mm long, cleft for upper 0–20(–50)%, persistent to rarely caducous; lobes 0.7–1.5 by 0.7–1.5 mm, glabrous to subglabrous above, glabrous to densely hairy below, protuberances above and below. Fruits obovoid, star-shaped in cross section, 1–3-seeded berries, 30–52 by 10–26 by 10–26 mm when dry, c. 50 by 26 mm when fresh, raised glands present, sparsely hairy outside, glabrous to subglabrous inside, red to purple to pink to red-brown; pericarp 1–2 mm thick, ruminate when dry; column 22–25 mm long, straight; pedicel 4–8 mm long, upper part 0.5–1 mm long. Seeds globose to ellipsoid, laterally flattened, 16–23 by 7–16 by 4–9.5 mm; arillode white; testa cream to greenish; cotyledons often folded, 8–12.5 by 6–11 by c. 0.1 mm; radicle 1.2–3 mm long; endosperm c. 1.5 mm thick.

Distribution — Borneo.

Habitat & Ecology — Primary and secondary rain forest, riverine and non-riverine forest. Soil: sandstone or laterite. Altitude: 0–800 m. Flowering: May, June, August, October, November; fruiting: throughout the year.

Uses — Pericarp and arillode edible, sour to sweet.

Vernacular names — Borneo (Kalimantan): Asem ketiak, pidau, umbing, umbung. Sarawak: Uchong, ujung (Iban). Brunei: Embaling bobou (Dusun); belimbing hutan. Sabah: Embaling (Dusun), belimbing uchong, pelawak, popotong, tampoi hutan.

2. Baccaurea bracteata Müll.Arg.

Baccaurea bracteata Müll.Arg. in DC., Prodr. 15, 2 (1866) 466; Hook.f., Fl. Brit. India 5 (1887) 372; Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 281; Merr., J. Straits Branch Roy. Asiat. Soc. (1921) 330; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 65; Ridl., Fl. Malay Penins. 3 (1924) 246; Merr., Univ. Calif. Publ. Bot. 15 (1929) 145; Pax & K.Hoffm., Mitt. Staatsinst. Allg. Bot. Hamburg 7 (1931) 233; Corner, Wayside Trees Mal. 1 (1940) 239; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 36; Airy Shaw, Kew Bull. 26 (1972) 219; Whitmore, Tree Fl. Malaya 2 (1973) 76; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 46; Kew Bull. 36 (1981) 260; Haegens & Welzen in Welzen et al., Thai For. Bull. 28 (2000) 69; Haegens, Blumea Suppl. (2000) 99, Fig. 3.6, Map 3.3; in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 109; plate IV: 1. — Type: Wallich 7834 (holo K; iso G), Singapore and Penang (both are mentioned on specimen label).

Baccaurea crassifolia J.J.Sm., Bull. Jard. Bot. Buitenzorg 3, 1 (1920) 394; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 66; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 46; Kew Bull. 36 (1981) 260. — Type: Hallier 2158 (holo L; iso BO, K), Borneo, Soengei Kenepai.

Treelet or tree 2–25 m high, dbh 4–40 cm, buttresses sometimes present, up to 1 m high, c. 13 cm thick, stilt roots rarely present, bole rarely fluted; branchlets glabrous to densely hairy, Terminalia branching pattern weakly developed. Indumentum of simple and stellate hairs. Bark brown to grey to pink to red to yellow when fresh, (dark to red-)brown to blackish when dry, 0.25–1 mm thick, usually smooth, hard, flaking in rectangular flakes of c. 10 by 2 mm; inner bark red to pale yellow to (red)-brown to green to pink, 0.25–8 mm thick, hard to soft. Heartwood pink to brown. Leaves: petiole 8–65(–87) mm long, subglabrous to densely hairy, simple hairs longer than those of the stellate tufts, transverse cracks often present, (pink)brown when fresh, black to red brown when dry, raised glands usually absent; stipules triangular, 2–6 by 1–2.5 mm, subglabrous to velutinous outside, glabrous to subglabrous inside, rarely densely hairy, margin ciliate, not hyaline; lamina ovate to elliptic (to obovate), 3.5–21 by 1.7–9.8 cm, l/w ratio 1.4–3.4, papery to subleathery; base truncate to attenuate; marginal glands sometimes slightly visible as small indentations; apex obtuse to cuspidate, up to 20 mm long; upper surface glabrous to subglabrous, small raised glands present, granulate, white green to dark green when fresh, pale to dark green-brown when dry; lower surface sparsely hairy to rarely densely hairy, sparsely hairy to velutinous on the midrib (to subglabrous), stellate hair-tufts at base with a conspicuous black gland, discoid glands usually present, placed in rows to scattered, raised glands sometimes present, bigger, white to pale green below when fresh, (pale) green-brown below when dry; nervation whitish to black-brown above when dry, blackish brown below, often sunken above; secondary veins 3–8 per side, not to almost closed at margin; nervation reticulate to weakly scalariform. Staminate inflorescences axillary to just below the leaves, solitary to 3 clustered together, 0.5–16 cm long, 0.5–1.2 mm thick, sparsely hairy to velutinous, branched, c. 10–100-flowered, flowers scattered along inflorescence, red; sometimes leaf-like bracts present; bracts 1 per branchlet, (broadly) ovate to triangular, conspicuous, 2–6 mm long, caducous to persistent, margin ciliate, sometimes hyaline, densely hairy to rarely subglabrous outside, glabrous inside, red to pale green; bracteoles c. 6, 1–5 mm long; branchlets cylindrical, 2–15 mm long, densely hairy, (3- or) 4–17(–40-)flowered. Staminate flowers 1.5–2.9 mm diam., caducous at the moment of opening; pedicel 0.3–1.2 mm long, upper part c. 0.1–1.1 mm long, densely hairy; sepals 4 or 5, ovate, 1–2.2 by 0.6–1.5 mm, apex recurved, densely hairy outside and inside, white to yellow to green to pink to red to brown when fresh; stamens 4 or 5, 0.25–0.5 mm long, glabrous, yellow; ﬁlaments 0.05–0.3 mm long, straight; anthers 0.1–0.25 by 0.15–0.2 by c. 0.1 mm, light yellow to reddish; disc absent; pistillode usually present, cylindrical to globose, up to 1 mm high, densely hairy, solid. Pistillate inflorescences axillary to just below the leaves, solitary, 1–6.5 cm long, in Borneo up to 16 cm long, c. 1.5 mm thick, sometimes branched, densely hairy, 5–30-flowered; pedicel 1–5 mm long, upper part 0.5–2 mm long, densely hairy, raised glands often present; bracts 1–6 per branchlet, (broadly) ovate, velutinous outside, glabrous inside, margin ciliate, persistent to caducous; rachis pink to brown-red to yellow-brown-green. Pistillate flowers 4–12 mm diam., yellowish to greenish to (red)brown; sepals 4 or 5 (or 6), ovate to elliptic, 3.5–11 by 0.9–3.5 mm, outside and inside densely hairy, caducous to persistent; staminodes rarely present; ovary cylindrical to globose, 2–4 by 1.5–3 mm, 2- or 3-locular, velutinous, wings absent (to 6), brown-yellow; style 0–1.6 by 0.5–1 mm, velutinous, pale yellow green; stigmas 0.9–2.5 mm long, cleft for upper 70–90%, persistent to caducous; lobes 0.8–1.8 by 0.2–0.5 mm, glabrous to subglabrous, protuberances small above, below densely hairy, protuberances strong. Fruits globose to triangular, 3–6-seeded, fleshy capsules, 19–25 mm diam. when fresh, 10.5–23 by 14–25 by 14–25 mm when dry, only partly loculicidally dehiscent, raised glands usually present, subglabrous to densely hairy outside, glabrous to sparsely hairy inside, septa subglabrous to sparsely hairy, red to orange-brown-green; pericarp 1–4.5 mm thick, often ruminate when dry; column 8–19.5 mm long, straight, persistent; pedicel 3–9 mm long, upper part 1.5–4.5 mm long, sometimes thickened. Seeds ovoid to obovoid, (6.2–)7–10 by (4–)4.5–6.5 by 3–4.8 mm; arillode (orange) yellow; testa white; cotyledons sometimes folded, 3–5 by (3–)4–5.5 by 0.05–0.1 mm; radicle 0.5–1.5 mm long; endosperm 0.5–1 mm thick.

Distribution — Thailand, Peninsular Malaysia, Sumatra, Borneo.

Habitat & Ecology — Primary or secondary rain forest, freshwater swamp, peat swamp, heath swamp, kerangas, rarely in mangrove forest. Altitude: sea level up to 390(–900) m. Soil: usually white sand. Flowering and fruiting: throughout the year. Fruits eaten by birds.

Uses — Sarawak: Shoots are used in making laminaang (= Dayak Kenya longhouse). Sabah: used for construction; arillode and pericarp edible, sour.

Vernacular names — Thailand: Khao-rang. Sumatra: Berat mata, tampui kaka. Bangka: Kelempa. Borneo: Kapul pugi nentalon, pugi ranau (Dusun); tampoi paya (Brunei, Kedayan); puak, puak burong, pugi barong (Iban); depot kayu masam (Mili); kelibon (Murut); jemating, selantikan, tampoi hutan, tampoi-tampoi hutan, tampoi munyit, terai rapak.

Note — See note B. simaloerensis.

3. Baccaurea brevipes Hook.f.

Baccaurea brevipes Hook.f., Fl. Brit. India 5 (1887) 372; Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 281; K.Heyne, Nutt. Pl. Ned. Ind. 3 (1917) 73; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 58; Ridl., Fl. Malay Penins. 3 (1924) 250; Corner, Wayside Trees Mal. 1 (1940) 239; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 36; Whitmore, Tree Fl. Malaya 2 (1973) 66; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 47; Kew Bull. 36 (1981) 260; Haegens & Welzen in Welzen et al., Thai For. Bull. 28 (2000) 69; Haegens, Blumea Suppl. (2000) 102, Map 3.4; in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 109, plate IV: 2. — Lectotype (designated by Haegens, 2000): Maingay KD 1366 (lecto K; isolecto L), Peninsular Malaysia.

Tree 3–10 m high, dbh 4–15 cm, buttresses absent; branchlets glabrous, densely hairy when young, Terminalia branching pattern weakly developed. Indumentum of simple hairs. Bark brown to grey to cream to white to green when fresh, brown when dry; inner bark (orange-)brown to white. Leaves: petiole 11–50 mm long, sparsely to densely hairy, usually not apically pulvinate, raised glands often absent; stipules 8–16 by 2–5 mm, glabrous on both sides, midrib velutinous outside, margin ciliate, not hyaline, caducous to late caducous; lamina elliptic to obovate, 15–35 by 6.2–14.4 cm, l/w ratio 1.9–3, papery; base cuneate to rounded; apex cuspidate to acute to rarely rounded, (0–)6–20 mm long; upper surface glabrous, raised glands absent, not granulate, brown when dry; lower surface densely to sparsely hairy, discoid glands absent, brown when dry; secondary veins 11–15 per side, sometimes closed at margin; nervation scalariform to somewhat reticulate. Staminate inflorescences ramiflorous to just below the leaves (to axillary), solitary to 8 clustered together, 8–17.5 cm long, up to 1.5 mm thick, velutinous, many-flowered, flowers scattered along inflorescence; bracts 1–3 per branchlet, 1.5–6 mm long, densely hairy outside, glabrous inside, margin ciliate, not hyaline, persistent; branchlets present to absent, cylindrical, 0–1 mm long, velutinous, 3-flowered. Staminate flowers 1.5–3 mm diam., yellowish green; pedicel 1.7–2.2 mm long, upper part 0.7–1.1 mm long, densely hairy; sepals 3–6, obovate or different in shape, 0.5–2 by 0.5–1.4 mm, apex recurved, densely hairy outside and inside; stamens 5, 0.5–1 mm long, glabrous; ﬁlaments 0.3–0.9 mm long, geniculate; anthers 0.1–0.2 by 0.1–0.2 by 0.1–0.3 mm; disc present to absent; pistillode obtriangular to globose, 0.4–1.1 mm high, densely hairy to velutinous, solid. Pistillate inflorescences cauline (to ramiflorous to axillary), solitary to 3-clustered together, 12–22.5 cm long, 0.7–3 mm thick, densely hairy, many-flowered; rachis pink; pedicel 0–4.5 mm long, upper part 0–1.2 mm long, velutinous; bracts 3 per branchlet, densely hairy outside, glabrous inside, ciliate, persistent. Pistillate flowers 5–7 mm diam.; sepals 4, obovate to elliptic, 5.6–6.5 by 1.5–3.5 mm, densely hairy, caducous to persistent; ovary globose, 2.2–2.6 by 2.2–3 mm, 2- or 3-locular, velutinous; style c. 0.2 by 1 mm wide, velutinous; stigmas c. 1 mm long, persistent. Fruits globose, 1–3-seeded, fleshy capsules to berries, 14–20 by 10–18.5 by 10–17 mm, not dehiscing to rarely splitting, raised glands present, glabrous to densely hairy outside, glabrous inside, white to red to pink; pericarp 0.5–1.5 mm thick; column 15–17 mm long, straight, persistent; pedicel 1–5 mm long, upper part 0.1–1 mm long. Seeds ellipsoid, laterally flattened, 8–13 by 6–8 by 3.1–5 mm; arillode blue to violet to rarely red; testa brown; cotyledons 5–7 by 6–7 by < 0.1 mm; radicle c. 1 mm long.

Distribution — Thailand, Peninsular Malaysia, Sumatra, NW Borneo.

Habitat & Ecology — Primary and secondary rain forest, and seasonal swamp forest. Altitude: 20–700 m. Soil: sand, clay or granite. Flowering: December to April; fruiting: May, July to November.

Uses — Locally grown for fruits.

Vernacular names — Peninsular Malaysia: Rambai, rambai hutan.

4. Baccaurea carinata Haegens

Baccaurea carinata Haegens, Blumea Suppl. (2000) 103, Map 3. — Typus: Hoogland & Craven 10350 (holo L), Papua New Guinea, East Sepik Province, Ambunti subdistrict, near Melawei.

Shrub to tree 7.5–10.5 m high; branchlets glabrous, densely hairy when young, Terminalia branching pattern weakly developed. Indumentum of simple hairs. Bark grey-brown when fresh, flaky; inner bark white, c. 2.5 mm thick. Leaves: petiole 21–59 mm long, glabrous to sparsely (to densely) hairy, transverse cracks usually absent, (dark to red-)brown when dry, raised glands absent to present; stipules 5–11.5 by 2–4.5 mm, glabrous outside, midrib glabrous to densely hairy, margin ciliate, hyaline or not; lamina ovate to obovate, 15–29 by 9.3–15.3 cm, l/w ratio 1.8–2.0, papery to slightly leathery; base attenuate to cuneate; apex rounded to acute, up to 10 mm long; upper surface glabrous, raised glands sometimes present, dark brown to greyish when dry; lower surface glabrous to subglabrous, midrib and secondary veins glabrous to densely hairy, raised glands present, discoid glands absent, (dark) brown when dry; nervation whitish to (light to dark) brown above when dry, (light to red) brown below when dry; secondary veins 10–15 per side, not completely parallel, closed at margin; nervation reticulate to weakly scalariform. Staminate inflorescences axillary to cauline, few clustered together, (4–)7–17 cm long, c. 1 mm thick, densely hairy, subglabrous at base, many-flowered, flowers scattered along inflorescence; bracts 3 per branchlet, 0.5(–2) mm long, glabrous to densely hairy outside, glabrous inside, margin ciliate, not hyaline; branchlets minute, 3-flowered. Staminate flowers 1.5–1.8 mm diam., yellow; pedicel 1.5–1.8 mm long, upper part 0.8–1 mm long, densely hairy; sepals 4, ovate, c. 1 mm long, apex straight, densely hairy outside and inside; stamens 6–9, c. 0.5 mm long, glabrous; ﬁlaments c. 0.3 mm long, straight; anthers c. 0.2 by 0.25 by 0.1 mm; disc absent; pistillode obtriangular, c. 0.5 mm high, densely hairy, solid. Pistillate inflorescences cauline, many clustered together; bracts 3 per branchlet. Pistillate flowers: sepals caducous; ovary 3-locular; stigmas persistent. Infructescences 8–13 cm long, 1.5–2.5 mm thick. Fruits ellipsoid, usually 5-seeded berries, c. 21 by 12 by 12 mm, slightly ruminate when dry, raised glands present, densely hairy outside, glabrous inside, green with pink tinges; pericarp c. 1 mm thick; column c. 17 mm long, straight; pedicel 3–5 mm long, upper part 1.5–3.5 mm long.

Distribution — Papua New Guinea (Prov.: E and W Sepik, Madang).

Habitat & Ecology — Secondary rain forest, swamp forest. Altitude: 50–100 m. Fruiting: August to September.

Uses — Arillode edible.

Vernacular names — New Guinea: Mapiok (Waskuk).

Note — The species is named for its strongly and sharply raised midvein at the lower side of the lamina, which resembles a keel.

5. Baccaurea costulata (Miq.) Müll.Arg.

Baccaurea costulata (Miq.) Müll.Arg. in DC., Prodr. 15, 2 (1866) 464; Boerl., Handl. Fl. Ned. Ind. 3 (1900) 281; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 68; S.Moore, J. Bot. Br. 63 (1925) 98; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 47; Kew Bull. 36 (1981) 261; Haegens, Blumea Suppl. (2000) 105, Map 3.6. — Mappa costulata Miq., Fl. Ned. Ind., Eerste bijv. (1861) 459. — Pierardia costulata (Miq.) Müll.Arg., Flora 47 (1864) 469. — Type: Teijsmann HB 3655 (holo U), Sumatra, Palembang, near Tubuan, Oga-ulu.

Tree up to 10–30 m high, dbh up to 28 cm, buttresses absent; branchlets glabrous to sparsely hairy, densely hairy when young, Terminalia branching pattern weakly developed. Indumentum of simple and stellate hairs. Bark grey, flaky. Leaves: petiole 9–47 mm long, glabrous to densely hairy, transversal stripes sometimes present, brown when dry, raised glands absent; stipules 2.5–5.5 by 1–2.5 mm, densely hairy outside, glabrous inside, base inside densely hairy, margin ciliate, not hyaline; lamina ovate to obovate, 6.1–19.6 by 2.3–7 cm, l/w ratio 2–2.7, papery to slightly leathery; base rounded to cuneate; apex obtuse to cuspidate, up to 21 mm long; upper surface glabrous, raised glands absent, sometimes punctuate, (dark) brown to grey when dry; lower surface (sub)glabrous, densely hairy at secondary veins, discoid glands absent or present, if present in a row between two secondary veins, (light) brown when dry; secondary veins 5–8 per side, closed at margin or almost so; nervation reticulate to weakly scalariform. Staminate inflorescences axillary to somewhat ramiflorous, solitary, 1.5–4 cm long, c. 1 mm thick, densely hairy, many-flowered, flowers scattered along inflorescence; bracts 1–1.5 mm long, 1 per branchlet, margin ciliate, not hyaline, densely hairy outside, sparsely hairy inside, persistent; branchlets cylindrical, 1.5–3 mm long, densely hairy, 3–6-flowered. Staminate flowers 1–2.5 mm diam.; pedicel 1–1.5 mm long, upper part 0.5–1 mm, densely hairy; sepals 4 or 5, ovate, 1–1.7 by 0.6–1 mm, outside and inside densely hairy; disc absent; stamens 5, c. 0.3 mm long, glabrous; ﬁlaments c. 0.15 mm long, straight; anthers c. 0.15 by c. 0.3 by c. 0.15 mm; pistillode, cylindrical to globose, 1.1–1.2 mm high, densely hairy, solid. Pistillate inflorescences axillary to ramiflorous, solitary. Pistillate flowers: sepals 4 or 5, persistent; ovary 3-locular; stigmas sessile, biﬁd, persistent. Infructescences 0.8–3.7(–13) cm long, 1.5–4 mm thick. Fruits 3- (or 4-)seeded, globose berries, 19–30 by 23–30 by 23–30 mm, 50 mm diam. when fresh, glabrous to sparsely hairy outside, subglabrous to sparsely hairy inside, raised glands usually absent; pericarp 1–4.5 mm thick; column 19–25 mm long, straight, persistent; pedicel 2–8 mm long, upper part 0.5–2 mm long. Seeds subglobose, laterally flattened, 12–19 by 11–16 by 3–8 mm; cotyledons c. 8.5–12.3 by 9.5–10.8 by 0.1 mm; radicle c. 1.5 mm long.

Distribution — Sumatra, Borneo.

Habitat & Ecology — Altitude: up to 450 m. Flowering: unknown; fruiting: February, April, July, August.

Uses — Arillode edible, with a bitter aftertaste.

Vernacular names — Borneo: Kampul (Dayak).

Note — In Baccaurea the vegetative parts of male and female plants usually closely resemble each other. This species, however, causes some interpretation problems, because the similarity between male and female is not 100%. The male type specimen is clearly different from all other material. Three fruiting specimens resemble this male specimen most (they do not ﬁt into other species). The female flowering and fruiting specimens, that traditionally have been included here, are hesitantly accepted by me.

6. Baccaurea deflexa Müll.Arg.

Baccaurea deflexa Müll.Arg. in DC., Prodr. 15, 2 (1866) 462; Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 281; J.J.Sm., Meded. Dept. Landb. Ned.-Indië 10 (1910) 248; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 69; Merr., Univ. Calif. Publ. Bot. 15 (1929) 145; Backer & Bakh.f., Fl. Java 1 (1963) 454; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 37; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 47; Haegens, Blumea Suppl. (2000) 108, Fig. 3.7, Map 3.8. — Lectotype (designated by Haegens, 2000): Zippelius s.n. (lecto L, sheet 903.154-568, barcode L 0050885), probably Java.

Treelet or tree 6–25 m high, dbh 10–70 cm, buttresses rarely present, up to 1 m high; branchlets glabrous, densely hairy when young, Terminalia branching pattern weakly developed. Indumentum of simple and stellate hairs. Bark grey to brown reddish when fresh, brown when dry, up to 2 mm thick, smooth to rough, peeling off in strips of 5–10 mm long; inner bark red to brownish to yellowish. Heartwood reddish. Leaves: petiole 17–50 mm long, subglabrous to densely hairy, raised glands absent; stipules 3–6 by 1–2.5 mm, densely hairy outside, glabrous inside, margin ciliate, not hyaline; lamina ovate to obovate, 7.5–19.5 by 3.8–8.9 cm, l/w ratio 1.6–2.5, papery to slightly leathery; base acute to attenuate; marginal glands rarely as small indentations; apex acuminate to cuspidate, 4–14 mm long; upper surface glabrous, rarely granulate, brown to grey when dry; lower surface subglabrous to densely hairy, raised glands absent, few discoid glands sometimes present, (greyish) brown when dry; secondary veins (5 or) 6–8 (or 9) per side, closed at margin or almost so; nervation (weakly) scalariform, brown to white-brown when dry. Staminate inflorescences axillary to cauline, solitary to 4(–many) clustered together, 1–5 cm long, c. 1 mm thick, branched, densely hairy, many-flowered, flowers scattered along inflorescence; bracts 0.5–1 mm long, 1 or 3 (then 2 rudimentary) per branchlet, densely hairy outside, glabrous (to densely hairy?) inside, margin ciliate, not hyaline; branchlets cylindrical, 0.5–2.2 mm long, densely hairy, 3-flowered. Staminate flowers 1–2.5 mm diam., sessile, green; sepals 3 or 4, ovate, 0.5–1.5 by 0.5–1.5 mm, apex slightly recurved, outside and inside densely hairy; stamens (4–)6, 0.3–0.6 mm long, glabrous; ﬁlaments 0.3–0.5 mm long, straight; anthers 0.1–0.15 by 0.1–0.3 by 0.1–0.15 mm; disc absent; pistillode absent to present, globose, up to 0.3 mm high, velutinous, solid. Pistillate inflorescences axillary to ramiflorous, 1–3 clustered together, 3.5–9 cm long, 0.4–1 mm thick, densely hairy, many-flowered, yellow; pedicel 1–2.5 mm long, upper part 0.5–2 mm long, geniculate (90º bent), densely hairy; bracts 1 or 3 (2 very small) per branchlet, densely to sparsely hairy outside, glabrous inside, margin ciliate. Pistillate flowers 1.6–3.1 mm diam., yellow-green; sepals (4 or) 5 or 6, ovate to triangular, 1–2 by 0.4–1.5 mm, outside and inside densely hairy, caducous to rarely persistent; ovary globose to cylindrical, 1.2–2 by 1–1.5 mm, 2-locular, densely hairy; style absent; stigmas 0.3–0.6 mm long, cleft, caducous; lobes 0.3–0.6 by c. 0.2 mm, glabrous above and below. Infructescences 5–17 cm long, 1–2.5 mm thick. Fruits subglobose to ellipsoid, but laterally flattened, usually 1- (or 2-)seeded fleshy capsules, 9–12 by 6–15 by 3–14 mm, loculicidally and often also septicidally dehiscent, raised glands absent, subglabrous to densely hairy outside, velutinous inside; pericarp 0.5–2 mm thick; column 7–12 mm long, curved to straight, persistent; pedicel 2–6 mm long, upper part 1–3 mm long. Seeds subglobose, laterally flattened, 5.8–12 by 4.5–8 by 2.5–4 mm; cotyledons 2.5–4 by 3–5 by c. 0.1 mm; radicle c. 1 mm long; endosperm up to 1 mm thick.

Distribution — Peninsular Malaysia, Sumatra, Borneo (the type is the only specimen from Java).

Habitat & Ecology — Primary rain forest. Altitude: sea level up to 450 m. Soil: sand, clay. Flowering: May, July to November; fruiting: May, July to September, November.

Vernacular names — Sumatra: Bergang gaya, birah mato, djatikau, djentikan betima, djentikan merah, merah matu, tampoei. Simeuluë Island: Bolawah ontur, bolawah silai. Bangka: Kepris, ketjepat, masput, rankop, rankup. Borneo: Jelentikan (Kutai); keliwat’n (Benuag/Tunjun); lubi, tampoi.

Notes — Müller Argoviensis (1866) already noted that this species is very similar to B. costulata. Airy Shaw (1975) regarded the two as conspeciﬁc, with no further explanation. In my opinion, however, the two species are rather different, and the confusion must have been due to incomplete data. The leaves of B. deflexa tend to be larger than those of B. costulata; the midrib (lower surface) usually densely hairy, compared with usually sparsely hairy; the staminate inflorescences less lax, always densely hairy (often sparsely hairy in B. costulata), bracts 0.5–1 mm long, compared with 1–2.5 mm long; stamens (4–)6, compared with 4 or 5, 0.3–0.6 mm long, in contrary to 0.2–0.3 mm long. These differences together with the different appearance of both species in the herbarium material (B. deflexa dark brown, B. costulata more glossy, greyish brown) have me led to conclude that B. deflexa and B. costulata are not conspeciﬁc. I omitted the pistillate inflorescences and infructescences from the discussion because of insufﬁcient knowledge of these characters in B. costulata (see note under B. costulata).

7. Baccaurea dolichobotrys Merr.

Baccaurea dolichobotrys Merr., Univ. Calif. Publ. Bot. 15 (1929) 147; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 47; Haegens, Blumea Suppl. (2000) 110, Map 3.9. — Lectotype (designated by Haegens, 2000): Elmer 21676 (lecto L; isolecto DS, GH, K, US), Borneo, Sabah, Tawau.

Tree 12–24 m high, dbh c. 29 cm, buttresses small; branchlets subglabrous to sparsely hairy; young shoots blackish when dry, Terminalia branching pattern weakly developed. Indumentum of simple and stellate hairs. Bark red-brown to dark reddish when fresh, brown when dry, scaly with cracks of 2 mm long; inner bark red-brown, c. 6 mm thick. Leaves: petiole 35–131 mm long, (sub)glabrous, dark brown to blackish when dry, raised glands usually absent; stipules 4–6 by c. 3 mm, densely hairy to velutinous outside, subglabrous to sparsely hairy inside, usually margin not ciliate, not hyaline; lamina ovate, (8.8–)10–30.3 by (5.7–)6.2–16.5 cm, l/w ratio 1.4–2.2, papery; base rounded to cordate; apex obtuse to acute, 2–15 mm long; upper surface glabrous, secondary veins glabrous to sparsely hairy, sometimes granulate; lower surface subglabrous to sparsely hairy, tuft of hairs present in nerve-axils, raised glands absent or present, discoid glands absent or present; secondary veins 10–12 per side, closed at margin; nervation scalariform. Staminate inflorescences unknown. Pistillate inflorescences ramiflorous to cauline. Pistillate flowers mainly unknown; sepals persistent to rarely caducous; ovary 3-locular, stigmas persistent. Infructescences 14–40 cm long, 1.7–2.5 mm thick. Fruits globose, usually 1- (or 2-)seeded, fleshy capsules, (9.5–) 10–13 by 10–10.5 by 10–13.5 mm, loculicidally dehiscent, raised glands often present, subglabrous to sparsely hairy outside, sparsely to densely hairy inside, yellow to orange-brown; pericarp 0.1–0.3 mm thick; column 9–12 mm long, curved to straight, persistent; pedicel 4.5–9 mm long, upper part 2–5 mm long. Seeds globose, laterally flattened, 5.3–7.3 by 5–7.2 by 2–3 mm; arillode red to orange; testa pale brown; cotyledons 4.2–6.3 by 5–6.5 by c. 0.1 mm; radicle 0.3–1 mm long; endosperm 0.3–1.5 mm thick.

Distribution — Borneo.

Habitat & Ecology — Riverine rain forest. Altitude: 0–700 m. Soil: yellow sand, loam. Fruiting: February, July, October.

Uses — Arillode edible.

Vernacular names — Sarawak: Buah jelentik (Iban).

Note — Airy Shaw synonymised B. dolichobotrys with B. hookeri (= B. polyneura). Baccaurea dolichobotrys is in my opinion clearly different from B. polyneura. The indument on the branches is sparsely hairy in B. dolichobotrys, whereas B. polyneura is densely hairy (B. dolichobotrys is in general less hairy than B. polyneura); the petioles and leaves are on average longer in B. dolichobotrys; the fruits are 10–10.5 mm diam., compared with 12–26 in B. polyneura. Other characters (such as colour of the leaves when dry) differ in range but usually overlap.

8. Baccaurea dulcis (Jack) Müll.Arg.

Baccaurea dulcis (Jack) Müll.Arg. in DC., Prodr. 15, 2 (1866) 460; Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 281; Koord., Exkurs. Fl. Java 2 (1912) 481; K.Heyne, Nutt. Pl. Ned. Ind. 3 (1917) 73; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 51; S.Moore, J. Bot. Br. 63 (1925) 98; Backer & Bakh.f., Fl. Java 1 (1963) 454; Soejarto, Bot. Mus. Leafl. (1965) 82; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 37; Airy Shaw, Kew Bull. 36 (1981) 261; Haegens, Blumea Suppl. (2000) 112, Map 3.10. — Pierardia dulcis Jack, Trans. Linn. Soc. 14 (1823) 120; Walp., Repert. Bot. Syst. 5 (1846) 367; Miq., Fl. Ned. Ind. 2, 1 (1859) 358; Fl. Ned. Ind., Eerste bijv. (1861) 441; Baill., Adansonia 3 (1863) 140. — Type: Jack s.n. (holo K), Sumatra, male flowering specimen.

Baccaurea suvrae Chakrab. & M.Gangop., J. Econ. Taxon. Bot. 18 (1994) 423. — Type: Forbes 3017 (holo CAL; iso L), Sumatra, Palembang, R. Rawas, Soeroelangoen.

Tree 3–30 m high, dbh 4–30 cm, buttresses absent; branchlets (sub)glabrous, hollow; young shoots (pale) red-brown to (pale) brown to grey when dry, Terminalia branching pattern weakly developed. Indumentum of simple hairs. Bark whitish-brown when fresh, grey when dry, c. 1 mm thick, rough; inner bark white, 2–3 mm thick. Leaves: petiole 8–68 mm long, glabrous to subglabrous, brown to red-brown when dry, raised glands usually present; stipules 6–15 by 2.5–6 mm, glabrous outside, but often densely hairy at base, glabrous to densely hairy at midrib, inside glabrous, margin not ciliate, not hyaline; lamina obovate, 7.8–30 by 3.7–11.6 cm, l/w ratio 1.5–3.3, papery to slightly leathery; base acute to attenuate; apex obtuse to acuminate, up to 11 mm long; upper surface glabrous, raised glands usually present, usually granulate, brown to reddish to grey to green when dry; lower surface glabrous, often subglabrous at midrib, discoid glands absent, brown to reddish to grey to green when dry; secondary veins 5–12 per side, closed at margin; nervation reticulate. Staminate inflorescences axillary to just below the leaves, solitary to few clustered together, 0.3–10 cm long, 0.5–1 mm thick, (glabrous to) subglabrous to densely hairy, many-flowered, flowers scattered along inflorescence; bracts 1 per branchlet, 1–2 mm long, (sub)glabrous outside, glabrous inside, margin ciliate, (not) hyaline; bracteoles 0–2, up to 0.3 mm long, sparsely hairy outside, glabrous inside; branchlets absent to cylindrical, 0–3 mm long, densely hairy, (2- or) 3-flowered. Staminate flowers 2–4.5 mm diam.; pedicel 1–3.8 mm long, upper part 0.5–1.5 mm long, densely hairy; sepals 4, ovate, 1.2–3.5 by 0.5–1.6 mm, apex (slightly) recurved, outside and inside densely hairy; staminodes 6–8; stamens 6–8, 0.4–0.9 mm long, glabrous; ﬁlaments 0.4–0.7 mm long, usually geniculate; anthers 0.1–0.15 by 0.15–0.2 by c. 0.1 mm; disc absent; pistillode absent to obtriangular, up to 1 mm high, densely hairy, hollow. Pistillate inflorescences axillary to ramiflorous (to cauline), solitary to few-clustered together, 2.5–12 cm long, 1–2.5 mm thick, (sub)glabrous, 8–30-flowered; pedicel, 2.5–10 mm long, upper part 0.3–4.2 mm, subglabrous to densely hairy; bracts inserted on rachis and pedicel, 3 per branchlet, (sub)glabrous outside, glabrous inside, ciliate. Pistillate flowers 5–10 mm diam., yellow; sepals 4 or 5, elliptic, 3–11 by 1.7–4.5 mm, sparsely to densely hairy outside, densely hairy inside, caducous, whitish when dry; ovary globose, 1.5–2.7 by 1.5–2.7 mm, 3-locular, (sub)glabrous; style absent; stigmas 0.4–0.8 mm long, not cleft to 90%, usually persistent; lobes 0.2–0.8 by 0.3–1 mm, glabrous and ruminate above and below. Fruits globose, 1–3-seeded berries, 24–35 by 11–26 by 11–26 mm when dry, raised glands present, ruminate when dry, (sub)glabrous outside, glabrous inside, yellow; pericarp 1.5–11 mm thick; column 21–24 mm long, straight; pedicel 8–17.5 mm long, upper part c. 1 mm long. Seeds globose to ellipsoid, laterally flattened, (10.6–)12.5–18 by (9–)10–12.5 by 3–5 mm; arillode purple; cotyledons 7–10.5 by 5.5–11.5 by c. 0.1 mm; radicle 1.5–2 mm long; endosperm c. 1 mm thick.

Distribution — Sumatra, W Java.

Habitat & Ecology — Primary rain forest, and cultivated. Altitude: 90–700 m. Flowering: June, August, October; fruiting: May, June, September, November, December.

Uses — Fruits sweet to sour.

Vernacular names — Sumatra: Kapunduang (Minang); cupak, kaloe, ketoepa, menteng negri, toepa. Simeuluë Island: Pranggo.

9. Baccaurea edulis Merr.

Baccaurea edulis Merr., Univ. Calif. Publ. Bot. 15 (1929) 149; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 36; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 47; Haegens, Blumea Suppl. (2000) 113, Fig. 3.8; Map 3.11. — Type: Elmer 21783 (holo A; iso DS, L), Borneo.

Tree 8–33 m high, dbh 5–60 cm, buttresses absent to up to 4 m high by 1 m wide, thin; branchlets glabrous, densely hairy when young, Terminalia branching pattern weakly developed. Indumentum of simple and stellate hairs. Bark pale to red-brown when fresh, brown when dry, c. 0.5 mm thick, smooth, hard to soft; inner bark light red to red-brown to yellow to pale brownish, 2–10 mm thick, soft, sometimes brittle. Heartwood red-brown. Leaves: petiole 12–77 mm long, densely hairy, raised glands absent; stipules triangular to elliptic, 6–11 by 2.5–4.5 mm, sparsely to densely hairy outside, (sub)glabrous inside, margin ciliate, not hyaline; lamina usually elliptic, 8.3–29 by 3.2–11.8 cm, l/w ratio 2.3–3.8, papery; base rounded to cuneate; marginal glands often visible as small indentations; apex acute to cuspidate, 3–20 mm long; upper surface glabrous, midvein usually densely hairy, dark green when fresh, dark brown when dry; lower surface subglabrous to densely hairy, raised glands usually absent, discoid glands present, yellowish green when fresh, light brown (to green-brown) when dry; young leaves with yellowish brown indumentum; secondary veins (7 or) 8–10 per side, closed at margin; nervation reticulate, white (to light brown) above when dry. Staminate inflorescences axillary to just below the leaves, solitary to few clustered together, 1.5–12.5 cm long, c. 1 mm thick, branched, densely hairy, many-flowered, flowers scattered along inflorescence; bracts usually 1 per branchlet, spatulate, 1–4 mm long, densely hairy outside, (sub)glabrous inside, margin ciliate, not hyaline; bracteoles absent; cymules variable in shape, 3–18-flowered, densely hairy. Staminate flowers 1–2.5 mm diam., yellowish green to yellow to white; pedicel 0.5–1.7 mm long, upper part 0.3–1.5 mm long, densely hairy; sepals 4 or 5, ovate to elliptic, 0.7–1.5 by 0.5–1 mm, apex recurved, outside and inside densely hairy; stamens (3 or) 4 or 5, 0.2–0.4 mm long, glabrous; ﬁlaments 0.1–0.3 mm long, straight; anthers 0.1–0.2 by 0.15–0.2 by c. 0.1 mm, pale yellow; pistillode absent to present, globose, up to 0.2 mm high, densely hairy, solid. Pistillate inflorescences ramiflorous to cauline, 2–6 clustered together, 2–18 cm long, 1–1.5 mm thick, densely hairy, 10–25-flowered, green; pedicel 3–7.8 mm long, upper part 0.5–1.5 mm long, densely hairy; bracts persistent, 1(–3) per branchlet, subglabrous to sparsely hairy outside, glabrous inside, margin ciliate. Pistillate flowers 3–7 mm diam., greenish yellow; sepals 4 or 5, ovate, 2–3.5 by 1–2 mm, outside and inside densely hairy, persistent; ovary globose, 2–4 by 1.5–4 mm, 2-locular, densely hairy; style 1–2 by c. 1 mm, densely hairy; stigmas 1–2 mm long, cleft for upper 90%, caducous (to persistent); lobes 1–2 by c. 1 mm, glabrous above, densely hairy below. Infructescences 2–4 mm thick. Fruits globose, 2–4-seeded berries, 5–6 cm diam. when fresh, 23–60 by 23–60 by 23–60 mm when dry, raised glands absent, glabrous outside, densely hairy when young, subglabrous to densely hairy inside, (orange-)brown to whitish to yellow; pericarp 4–10 mm thick; column 19–25 mm long, straight; pedicel 5–14 mm long, upper part 1–6 mm long. Seeds globose to ellipsoid, laterally flattened, 13–20.5 by 8–21 by 2–8 mm; arillode white to yellow; cotyledons 11–12 by 10.5–16 by 0.1–0.4 mm; radicle 1–1.5 mm long; endosperm 1–2 mm thick.

Distribution — (Peninsular Malaysia), Borneo.

Habitat & Ecology — Primary and secondary rain forest, and swamp forest. Soil: sand. Altitude: up to 700 m. Flowering: March, June to August, October; fruiting: February, March to June, August, September.

Uses — Arillode and seed-coat edible, sweet to sour, sold in local markets.

Vernacular names — Borneo: Apor-apor (Bassap); pendal nyumbo, tampoi paya (Iban); pas, tampoi paya (Malay); boenjan, kapul, kapul putih, kelawat’n petik, kepsoet awoet, kulibon, puak burong.

10. Baccaurea javanica (Blume) Müll.Arg.

Baccaurea javanica (Blume) Müll.Arg. in DC., Prodr. 15, 2 (1866) 456; Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 281; J.J.Sm., Meded. Dept. Landb. Ned.-Indië 10 (1910) 253; Koord., Exkurs. Fl. Java 2 (1912) 482; K.Heyne, Nutt. Pl. Ned. Ind. 3 (1917) 73; Merr., J. Straits Branch Roy. Asiat. Soc. (1921) 331; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 50; S.Moore, J. Bot. Br. 63 (1925) 98; Holth. & H.J. Lam, Blumea 5 (1942) 200; Backer & Bakh.f., Fl. Java 1 (1963) 254; Soejarto, Bot. Mus. Leafl. 21 (1965) 96; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 37; Whitmore, Tree Fl. Malaya 2 (1973) 65; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 48; Kew Bull. 36 (1981) 263; 37 (1982) 8; Chakrab. & Rao, J. Econ. Taxon. Bot. 5 (1984) 957; G.L. Webster, Ann. Missouri Bot. Gard. 81 (1994) 51; Haegens, Blumea Suppl. (2000) 116, Fig. 3.9, Map 3.12. — Adenocrepis javanica Blume, Bijdr. (1825) 579; Baill., Étude Euphorb. (1858) 601; Adansonia 3 (1863) 134. — Lectotype (designated by Haegens, 2000): Blume s.n. (lecto L, sheet 903.154-589), Java, Tjanjor, Parang.

Hedycarpus javanicus Miq., Fl. Ned. Ind. 1, 2 (1859) 359. — Lectotype (designated by Haegens, 2000): Hasskarl s.n. (lecto L, sheet 909.25-49, barcode L 0059164), Java.

Microsepala acuminata Miq., Fl. Ned. Ind., Eerste bijv. (1861) 444. — Baccaurea acuminata (Miq.) Müll.Arg. in DC., Prodr. 15, 2 (1866) 463; Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 281. — Lectotype (designated by Haegens, 2000): Diepenhuis s.n. (lecto U, sheet -95/007-45), Sumatra, Priaman, Lubualang.

Baccaurea minutiflora Müll.Arg. in DC., Prodr. 15, 2 (1866) 463; Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 281; J.J.Sm., Meded. Dept. Landb. Ned.-Indië 10 (1910) 257; Koord., Exkurs. Fl. Java 2 (1912) 482; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 58; Backer & Bakh.f., Fl. Java 1 (1963) 455; Airy Shaw, Kew Bull. 36 (1981) 260. — Type: Zollinger 3048 (holo G), Java.

Baccaurea minahassae Koord., Meded. Lands Plantentuin 19 (1898) 625; Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 281; Koord., Syst. Verz. 3 (1914) 67; K.Heyne, Nutt. Pl. Ned. Ind. 3 (1917) 74. — Type: Riedel s.n. (holo K), North Celebes, Gorontalo.

Baccaurea sanguinea J.J.Sm., Icon. Bogor. 4 (1910) 25, t. 319. — Lectotype (designated by Haegens, 2000): Bogor Botanical Garden: VIII.F.3a (lecto BO).

Aporosa dolichocarpa Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 98; S.Moore, J. Bot. Br. 63 (1925) 97; Airy Shaw, Kew Bull. 36 (1981) 264. — Lectotype (designated by Haegens, 2000): Forbes 3159 (lecto L), Sumatra.

Baccaurea leucodermis Hook.f. ex Ridl., Fl. Malay Penins. 3 (1924) 244; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 49. — Syntypes: Kunstler s.n. (K, n.v.), Malakka, Perak, Kinta; Ridley s.n. (K, n.v.), Johore, Castlewood.

Shrub to tree 3–12 m high, dbh up to 30 cm, buttresses absent to up to 30 cm high, 10 cm long, c. 5 cm thick; branchlets glabrous, densely hairy when young, Terminalia branching pattern well-developed. Indumentum of simple hairs. Bark (light) brown to pale grey to yellowish to white to silvery green, 1–2 mm thick, smooth, ﬁnely ﬁssured; inner bark brown to yellow-brown, 2–4 mm thick, soft. Heartwood brown. Leaves: petiole 2–45 mm long, (sub)glabrous, sometimes with a tuft of short hairs at base, brown when dry; stipules 1–6 by 1–2 mm, densely hairy on both sides, margin ciliate, hyaline; lamina obovate (to ovate), 2.9–20 by 1.2–9.5 cm, l/w ratio 1.9–3.9, base cuneate; apex acuminate, up to 15 mm long; upper surface glabrous, sometimes granulate; raised glands sometimes present, (dark) green when fresh, yellowish green to brown when dry; lower surface glabrous, discoid glands rarely present, (dark) green when fresh, yellowish green to brown when dry; secondary veins 4–8 per side, ending open; nervation reticulate, red to brown when dry, slightly raised above; young leaves reddish brown when dry. Staminate inflorescences axillary to just below the leaves, solitary to 4 clustered together, 1.5–14 cm long, 0.5–1 mm thick, subglabrous to sparsely hairy, many-flowered, flowers scattered along inflorescence; bracts 1 per branchlet, 0.5–1.1 mm long, persistent, glabrous on both sides, densely hairy at base outside, margin ciliate, hyaline; bracteoles 0 or 2, up to 0.5 mm long; branchlets cylindrical, 0–2.1 mm long, glabrous to densely hairy, (1–)3-flowered. Staminate flowers 1.7–3.7(–5.5) mm diam., green to white to light yellow to brownish; pedicel 1.2–4.2 mm long, upper part 0.9–2.2 mm long, glabrous to densely hairy; sepals 4 (or 5), ovate, 1.1–4.2 by 0.5–2.3 mm, apex recurved, outside glabrous except one side at margin densely hairy, densely hairy inside, brown outside when dry, whitish inside when dry; staminodes absent or 4–7; stamens 4–7, 0.4–1 mm long, glabrous; ﬁlaments 0.25–1 mm long, apically geniculate; anthers 0.1–0.4 by 0.1–0.2 by 0.1–0.2 mm; disc present, less than 0.1 mm thick; pistillode cylindrical, 0.3–1.1 mm long, densely hairy to velutinous. Pistillate inflorescences axillary to just below the leaves (to ramiflorous), solitary, up to 8.5 cm long, 1.2–1.4 mm thick, glabrous to sparsely hairy (to densely hairy), 1–17-flowered, brown to reddish brown when dry; pedicel 1.6–4 mm long, upper part 0.2–1.1 mm long, glabrous (to densely hairy); bracts 1 or 3 per branchlet, 0.7–0.8 by 0.7–0.8 mm, persistent, glabrous, margin ciliate; flowers up to 1 cm diam., green; sepals 4, ovate to lanceolate, 3–7.5 by 1.2–2.5 mm, outside glabrous, inside densely hairy, outside brown when dry, inside whitish when dry, caducous; ovary globose to cylindrical, 1.2–2.5 by 1–1.9 mm, 2-locular (2- or 3-locular in Borneo), pilose to velutinous, greenish-white; style absent or up to 0.1 mm; stigmas whitish, c. 0.5 mm diam., persistent, not lobed. Fruits globose to cylindrical, bivalved, 1- or 2(–4-)seeded, fleshy capsules, 10–17 by 7–11 by 7–11 mm, not to loculicidally dehiscing, glabrous outside and inside, pale to dark orange to yellow; pedicel 1.5–5 mm long, upper part 0.5–2.5 mm long; pericarp 0.1–1.5 mm thick; column 7–11 mm long, curved when 1-seeded, caducous. Seeds ellipsoid to slightly globose, but laterally flattened, 5–9 by 5.5–7.4 by 1–5 mm; arillode purplish blue to dark purplish to violet; testa red to purplish red; cotyledons 5–6 by 6–6.5 by 0.1–2.5 mm thick.

Distribution — Andaman and Nicobar Islands, Peninsular Malaysia, Sumatra, Java, Borneo, Sulawesi (Talaud Island).

Habitat & Ecology — Primary or secondary rain forest, or freshwater swamps. Altitude: sea level up to 1800 m. Flowering and fruiting: throughout the year.

Uses — Arillode edible, sweet to sour.

Vernacular names — Sumatra: Bosi, toepak rawang. Simeuluë Island: Ahanang bala, ahalang bahok, ahanang bahok pajo, toeanang, toepa rawang. Java: Houtjit, hutjiet, huetjip, heuncit, heuntit. Borneo: Mata plantok (Tidong); sarakat. Bali: Kepinding.

11. Baccaurea lanceolata (Miq.) Müll.Arg

Baccaurea lanceolata (Miq.) Müll.Arg. in DC., Prodr. 15, 2 (1866) 457; Hook.f., Fl. Brit. India 5 (1887) 368; Stapf, Trans. Linn. Soc. London, Bot. 4 (1894) 224; Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 280; Merr., Philipp. J. Sci., Bot. 4 (1909) 277; J.J.Sm., Meded. Dept. Landb. Ned.-Indië 10 (1910) 247; Koord., Exkurs. Fl. Java 2 (1912) 481; K.Heyne, Nutt. Pl. Ned. Ind. 3 (1917) 73; Merr., J. Straits Branch Roy. Asiat. Soc. (1921) 331; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 60; Merr., Enum. Philipp. Fl. Pl. 2 (1923) 411; Ridl., Fl. Malay Penins. 3 (1924) 248; S.Moore, J. Bot. Br. 63 (1925) 98; Merr., Univ. Calif. Publ. Bot. 15 (1929) 150; Pax & K.Hoffm., Mitt. Staatsinst. Allg. Bot. Hamburg 7 (1931) 233; M.R. Hend., J. Malayan Branch Roy. Asiat. Soc. 17 (1939) 69; Corner, Wayside Trees Mal. 1 (1940) 240; Backer & Bakh.f., Fl. Java 1 (1963) 454; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 35; Airy Shaw, Kew. Bull. 26 (1972) 220; Whitmore, Tree Fl. Malaya 2 (1973) 65; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 49; Fernando, Philipp. J. Biol. 8 (1979) 302; Airy Shaw, Kew Bull. 36 (1981) 263; Enum. Euphorb. Philipp. (1983) 9; Haegens & Welzen in Welzen et al., Thai For. Bull. 28 (2000) 70; Haegens, Blumea Suppl. (2000) 119, Fig. 3.10, Map 3.13, Photo 2, 3; in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 110. — Hedycarpus lanceolatus Miq., Fl. Ned. Ind. 1, 2 (1859) 359. — Adenocrepis lanceolatus (Miq.) Müll.Arg., Linnaea 32 (1863) 82. — Type: Zollinger 3265 (holo U; iso K), Java.

Pierardia pyrrhodasya Miq., Fl. Ned. Ind., Eerste Bijv. (1861) 441. — Baccaurea pyrrhodasya (Miq.) Müll.Arg. in DC., Prodr. 15, 2 (1866) 462; Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 281; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 65; Airy Shaw, Kew Bull. 36 (1981) 265. — Lectotype (designated by Haegens, 2000): Junghuhn s.n. (lecto U, sheet 35998, the flowers only), Sumatra, Ankola.

Baccaurea glabriflora Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 59; Merr., Enum. Philipp. Fl. Pl. 2 (1923) 411; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 50. — Type: Merril 853 (holo US; iso K), Philippines, Island of Paragua, San Antonio Bay.

Tree 3–30 m high, dbh 5–60 cm, buttresses absent; trunk gnarled; branchlets (sub)glabrous; young shoots green to grey-green to brownish when dry, Terminalia branching pattern usually weak. Indumentum of simple hairs. Bark grey to pale brown to yellowish to whitish to greenish when fresh, grey when dry, 0.2–3 mm thick, smooth, scaly, sometimes peeling off, soft; inner bark pale brown to white to grey to yellowish, 2–3.5 mm thick. Heartwood yellowish to brownish. Leaves: petiole 16–184 mm long, glabrous, apically and slightly basally pulvinate, green to grey to brown-green when dry, raised glands usually present; stipules (3–)5–16 by 2–6 mm, caducous (to late caducous), glabrous outside and inside, midrib often densely hairy above, margin (not) ciliate, usually hyaline (in Peninsular Malaysia not hyaline); lamina ovate to obovate, 9.2–45 by 3.7–26.5 cm, l/w ratio 1.7–3.4, papery; base attenuate to rounded; apex acute to cuspidate, (0–)3–20 mm long; upper surface glabrous, raised glands present, sometimes granulate, glossy dark green when fresh, (grey to brown to) green when dry; lower surface glabrous, sometimes sparsely hairy at midrib, raised glands present, discoid glands absent, (dull pale) green when fresh, (grey to brown to) green when dry, nervation paler; secondary veins 6–13 per side, closed at margin to almost so; nervation reticulate. Staminate inflorescences cauline, many clustered together, 3.5–18 cm long, 0.1–1 mm thick, glabrous to sparsely hairy, branching minute, many-flowered, flowers scattered along inflorescence, yellow to pink to cream-white; bracts 1 per branchlet, 0.4–1.5 mm long, densely hairy outside, (sub)glabrous inside, margin ciliate, hyaline; bracteoles 0.1–0.5 mm long, 2 per branchlet, glabrous outside, (sub)glabrous inside; branchlets absent, 3-flowered. Staminate flowers 2–7 mm diam., yellow to pink to purple to cream-white; pedicel 1.8–7 mm long, upper part 1.7–7 mm long, (sub)glabrous, densely hairy at base; sepals (3 or ) 4 or 5, obovate to spatulate, often variable in size and shape, 1.1–5.5 by 0.6–2.7 mm, apex slightly recurved or straight, glabrous outside (densely hairy in Peninsular Malaysia), sparsely to densely hairy inside; petals often present, 3–5, rudimentary; staminodes 3–5; stamens (3 or) 4 or 5, 0.4–0.6 mm long, glabrous; ﬁlaments 0.1–0.3 mm long, straight; anthers 0.2–0.35 by 0.2–0.4 by 0.15–0.2 mm; disc absent; pistillode globose to cylindrical, 0.2–0.7 mm high, velutinous, solid. Pistillate inflorescences cauline, few clustered together, 8–33(–50) cm long, 1–5 mm thick, subglabrous to sparsely hairy (glabrous in Peninsular Malaysia), 20–25-flowered, yellowish brown to reddish; pedicel 0.8–4.5 mm long, thickened at abscission zone, upper part 0.1–1.5 mm long, subglabrous to densely hairy (glabrous in Peninsular Malaysia), pink; bracts 3 per branchlet, (sub)glabrous outside, glabrous inside, margin ciliate. Pistillate flowers 3.5–10 mm diam., yellow to orange to purple to reddish cream to whitish; sepals 4 or 5, ovate to obovate, 1–6 by 0.9–3.2 mm, densely hairy outside, (sub)glabrous inside, caducous; petals 2–8, reduced; ovary globose, 1.5–3.2 by 1.5–3.2 mm, 3- or 4-locular, sparsely to densely hairy, wings usually absent, rarely 3 or 4 poorly developed, purple to green; style absent; stigmas 0.7–1 mm long, cleft for upper 10–90%, reddish to ochre, persistent to caducous; lobes 0.7–1 by 0.2–0.6 mm, glabrous and ruminate above, glabrous to densely hairy below. Fruits globose to ellipsoid, 1–4-seeded berries, 24–54 by 16.5–41 by 16.5–41 mm when dry, 38 by 60 mm when fresh, raised glands present, ruminate when dry, glabrous outside and inside, purple or green when young, green to yellow to whitish to greyish to brownish when mature; pericarp 1–10 mm thick; column c. 32 mm long; pedicel (1–)2.5–17 mm long, upper part 0.5–11 mm long. Seeds ellipsoid, laterally flattened, 12.2–26 by 8–15 by 4.8–9 mm; arillode white to grey, translucent; testa yellow to whitish; cotyledons 10–11.5 by 8.9–13 by c. 0.1 mm; radicle 1–2.2 mm long; endosperm 1–2 mm thick.

Distribution — Thailand, Peninsular Malaysia, Sumatra, N Java, Borneo, Palawan.

Habitat & Ecology — Primary and secondary rain forest, on slopes and in riverine forest. Soil: sandy clay, loam. Altitude: sea level up to 1300 m, common in lowland. Flowering: March to December; fruiting: throughout the year.

Uses — Against stomachache: the leaves are pounded in bamboo and mixed with water. Pericarp and arillode edible but sour and therefore eaten with sugar or salt. Rarely in cultivation in fruit gardens. Eaten with chicken rice.

Vernacular names — Peninsular Malaysia: Asam pahong, asam pahung, asam paung, limpanong, pahu asam, pahu temuangi. Sumatra: Tegeiluk (Mentawai); kaloe goegoer, langsat hutan, lempaong, lempaoe-oeng, peng. Java: Lingsoe. Borneo: Limpasu (Banjarese; Bundu tuhan); ampusu’ (Bidayuh); asam pauh, empaong, lampaong, lampawong, lampong (Iban); buah lepasu, lipasu, nipassu (Dusun); kalampesu, lempahong (in Kalimantan); buah lipauh (Kelabit); kelepesoh (Kenyah); tampoy (Malay); buah lepesuh (Punan); empawang, lapahung, lempawong, paong.

Note — The type specimen is a mixtum, the leaves are Terminalia sp.

12. Baccaurea macrocarpa (Miq.) Müll.Arg.

Baccaurea macrocarpa (Miq.) Müll.Arg. in DC., Prodr. 15, 2 (1866) 457; Hook.f., Fl. Brit. India 5 (1887) 375; Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 280; K.Heyne, Nutt. Pl. Ned. Ind. 3 (1917) 73; Pax & K.Hoffm., Engl. Pflanzenr. IV.147.xv (1922) 49; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 35; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 50; Kew Bull. 36 (1981) 263; Haegens, Blumea Suppl. (2000) 121, Fig. 3.11, Map 3.14, Photo 4; Haegens & Welzen in Welzen et al., Thai For. Bull. 28 (2000) 70; in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 111. — Pierardia macrocarpa Miq., Fl. Ind. Bat., Eerste bijv. (1861) 441. — Lectotype (designated by Haegens, 2000): Teijsmann s.n. (lecto U, sheet -95/007-35||36001), Sumatra, Dapoei, Mangala, Lampongs.

Mappa borneensis Müll.Arg., Flora 47 (1864) 468. — Baccaurea borneensis (Müll.Arg.) Müll.Arg. in DC., Prodr. 15, 2 (1866) 460; Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 280; Merr., J. Straits Branch Roy. Asiat. Soc. (1921) 330; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 62; Airy Shaw, Kew Bull. Add. .Ser. 4 (1975) 50. — Type: Motley 199 (holo K), Borneo, near Bandjarmassing.

Baccaurea griffithii Hook.f., Fl. Brit. India 5 (1887) 371; Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 281; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 66; Ridl., Fl. Malay Penins. 3 (1924) 248; Merr., Pap. Michingan Acad. Sci. 20 (1935) 100; Corner, Wayside Trees Mal. 1 (1940) 240; Whitmore, Tree Fl. Malaya 2 (1973) 66; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 50. — Lectotype (designated by Haegens, 2000): King’s Collector 3266 (lecto K), Malakka, Perak, Larut.

Treelet or tree 5–27 m high, dbh 6–64 cm; bole sometimes fluted, up to 5 m high; buttresses absent to low to small; branchlets glabrous; young shoots blackish, Terminalia branching pattern weakly developed. Indumentum of simple hairs and small stellate. Bark dark to pale (red) brown to grey to orange when fresh, greyish-brown when dry, hard, c. 0.1 mm thick, smooth; inner bark red-brown to pale brown to orange to red to pinkish white, soft, 2.5–3 mm thick. Leaves: petiole (10–)22–145 mm long, glabrous to sparsely hairy, brown to blackish when dry, raised glands usually present; stipules 2–9 by 1–5 mm, glabrous to tomentose outside, glabrous inside, margin ciliate, not hyaline; lamina (ovate to) elliptic (to obovate), (7.2–)9–37 by 3.1–17.5 cm, l/w ratio (1.1–)1.6–2.8(–3.5), leathery to papery; base attenuate to cuneate (to rounded); apex (obtuse to) acuminate to cuspidate, up to 20 mm long; upper surface glabrous, sometimes granulate, brown to greenish when dry, nervation brown to rarely whitish when dry; lower surface (sub)glabrous, mostly sparsely hairy on midrib and secondary veins, raised glands absent or present, discoid glands in a row between secondary veins, green with brown nervation when fresh, brown to greenish when dry; secondary veins 6–10(–13) per side, not completely parallel, ending open at margin; nervation reticulate to weakly scalariform; young leaves blackish brown when dry. Staminate inflorescences (axillary to) ramiflorous to cauline, solitary to few clustered together, 0.5–13 cm long, c. 1 mm thick, densely to sometimes sparsely hairy, usually branched, many-flowered, flowers scattered along inflorescence; bracts 1 (or 3) per branchlet, 1–3 mm long, subglabrous to densely hairy outside, glabrous to densely hairy inside, margin ciliate, not hyaline; branchlets cylindrical, (0–)1–2 mm long, densely hairy, 3–12-flowered. Staminate flowers 0.7–2 mm diam., green to yellow to white; pedicel 1–2 mm, upper part 0.1–1.5 mm long, densely hairy; sepals 5, elliptic, 0.7–1.5 by 0.4–0.7 mm, apex straight, outside and inside densely hairy; stamens 5 (or 6), 0.1–0.25 mm long, glabrous, yellowish; ﬁlaments 0.1–0.2 mm long, straight; anthers c. 0.1 by 0.15 by 0.1 mm; disc absent; pistillode globose to obtriangular, c. 0.5 mm high, densely hairy, sometimes hollow. Pistillate inflorescences ramiflorous to cauline, solitary to 3 clustered together, 3.5–18 cm by 2–3 mm thick, subglabrous to densely hairy, 8–many-flowered; pedicel 3–7.5 mm long, upper part 0.4–0.5 mm long, geniculate or not, densely hairy, green; bracts 1 or 3 (then 1 big) per branchlet, sparsely to densely hairy outside, subglabrous to densely hairy inside, margin ciliate. Pistillate flowers 2–4.5 mm diam.; sepals 4–6, ovate, 1.5–2.8 by 1–1.3 mm, outside and inside densely hairy, persistent; ovary globose to cylindrical, c. 2 by 1.2–2 mm, 3- or 4-locular, tomentose; style 0.5–1.5 mm long, c. 1 mm wide, sparsely hairy; stigmas 0.5–1 mm long, cleft, persistent to caducous; lobes c. 1 by 0.4 mm, (sub)glabrous. Infructescences up to 15 cm long, 4–6 mm thick. Fruits (sub)globose, (2- or) 3–6-seeded, fleshy capsules, 30–65 by 34–75 by 34–75 mm, up to 80 mm diam. when fresh, loculicidally (and also septicidally) dehiscent, (sub)glabrous outside, subglabrous to densely hairy inside, raised glands present, brown to yellow to orange to dull red to dark green; pericarp 4–11 mm thick; column 16–32 mm long, straight, caducous; pedicel 7–30 mm long, upper part 5–20 mm long, thickened at abscission zone. Seeds globose to ellipsoid, laterally flattened, (to triangular), 13–23 by 11–18.5 by 4–7 mm; arillode white to yellow to sometimes orange; testa yellow to brown; cotyledons 9–15 by 9–19 by up to 1 mm; radicle 1–2.3 mm long; endosperm c. 1 mm thick.

Distribution — Peninsular Malaysia, Sumatra, Borneo, (Ambon, Irian Jaya).

Habitat & Ecology — Primary rain forest, riverine rain forest and swamp forest, rarely also in secondary forest. Soil: granitic sand, sandy clay or red clay. Altitude: sea level up to 1600 m. Flowering and fruiting: throughout the year.

Uses — Arillode edible, sweet to sour. Sold in markets and cultivated in gardens.

Vernacular names — Peninsular Malaysia: Merkeh (Kelantan); ngeke (Malay); lara (Temuan); rambai, tampoi batang, tampoi, tampui. Sumatra: Tampoei daoen, tampoei benez. Bangka: Medang, tampui. Borneo: Pasin (Bassap Dyab); pegak (Dayak Tunjung); puak, tampoi (Iban); setai (Kenya); djentikan (Malay, Kutei); tampoi (Malay, Kedayan); buah setei, empak kapur, kapul, terai.

13. Baccaurea macrophylla (Müll.Arg.) Müll.Arg.

Baccaurea macrophylla (Müll.Arg.) Müll.Arg. in DC., Prodr. 15, 2 (1866) 460; Hook.f., Fl. Brit. India 5 (1887) 369; Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 281; K.Heyne, Nutt. Pl. Ned. Ind. 3 (1917) 74; Merr., J. Straits Branch Roy. Asiat. Soc. (1921) 331; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 62; Ridl., Fl. Malay Penins. 3 (1924) 247; S.Moore, J. Bot. Br. 63 (1925) 98; Airy Shaw, Kew Bull. 26 (1972) 220; Whitmore, Tree Fl. Malaya 2 (1973) 66; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 51; Kew Bull. 36 (1981) 263; Haegens, Blumea Suppl. (2000) 124, Map 3.15; Haegens & Welzen in Welzen et al., Thai For. Bull. 28 (2000) 71; in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 112. — Pierardia macrophylla Müll.Arg., Flora 47 (1864) 516. — Type: Anonymous (Hb Hook.f.; holo K), Malakka, Puolo-Penang.

Baccaurea beccariana Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 62; Airy Shaw, Kew Bull. 26 (1972) 220. — Type: Beccari PB 3249 [holo FI (?, n.v.); iso G, K], Borneo. The type is to be expected in FI but could not be found.

Tree 8–27 m high, dbh 14–63 cm; buttresses absent or small; bole sometimes fluted; branchlets glabrous, sparsely hairy when young; young shoots brown when fresh, brown to blackish when dry, Terminalia branching pattern weakly developed. Indumentum of stellate hairs. Bark grey to dark red to red-brown when fresh, grey when dry, smooth, flaky; inner bark pale red to pinkish, thick, brittle, soft with sticky sap. Heartwood dark red-brown. Leaves: petiole 30–88 mm long, subglabrous to sparsely hairy, raised glands absent or present; stipules 2–11 by 3–11.5 mm, caducous (to late caducous), (glabrous to) subglabrous to sparsely (to densely) hairy outside, midrib sparsely to densely hairy, (sub)glabrous inside, margin ciliate, not to slightly hyaline; lamina (ovate to) elliptic (to obovate), 9.6–33 by 4.5–14.4 cm, l/w ratio 1.9–2.3(–2.9), papery; base rounded to cuneate; marginal glands not to slightly visible as indentation marks; apex acute, 2–10 mm long; upper surface glabrous, not granulate, dark brown when dry; lower surface glabrous to sparsely hairy, raised glands absent or present, discoid glands present below in a row between two secondary veins; brown when dry; secondary veins 6–13 per side, usually ending open at margin; nervation reticulate; young leaves yellowish green. Staminate inflorescences axillary to ramiflorous, solitary to few clustered together, 2.5–8.6 cm long, 0.5–1 mm thick, densely hairy, many-flowered, flowers clustered at the tip of inflorescence; bracts 1 per branchlet, 1.5–2.5 mm long, sparsely (to densely) hairy outside, (sub)glabrous inside, margin ciliate, usually not hyaline; bracteoles 2 per branchlet, 0.2–0.7 mm long; branchlets cylindrical, 0.7–3 mm long, densely hairy, 3-flowered. Staminate flowers 0.7–1.5 mm diam., green to yellow; pedicel 0–0.7 mm long, densely hairy; sepals 4 or 5, ovate, 0.4–0.9 by 0.4–0.6 mm, outside and inside densely hairy; staminodes absent to 4; stamens 4 or 5, 0.2–0.7 mm long, glabrous; ﬁlaments 0.1–0.7 mm long, straight; anthers 0.1–0.2 by 0.2–0.3 by c. 0.1 mm, yellow; disc absent; pistillode globose to cylindrical, 0.3–0.6 mm high, densely hairy, solid. Pistillate inflorescences ramiflorous, few clustered together, (4–)12–25 cm long, 1–1.5 mm thick, densely hairy to velutinous, few to many-flowered, green; pedicel 2–5 mm long, upper part c. 0.2 mm long, geniculate (90° bent), densely hairy; bracts 3 per branchlet, sparsely to densely hairy outside. Pistillate flowers 2–4 mm diam., green when fresh; sepals 4, lanceolate, c. 1.5 by 1–1.5 mm, outside and inside densely hairy, caducous (to persistent); ovary globose, 1.5–2 by 1–1.5 mm, (2- or) 3-locular, velutinous; stigmas sessile, c. 0.7 mm long, apically not lobed, glabrous, persistent. Infructescences up to 25 cm long. Fruits globose, 2- or 3-seeded, fleshy capsules, 10–16 by 9.5–15 by 9.5–15 mm, loculicidally dehiscent to irregularly splitting, ruminate when dry, raised glands present, (sub)glabrous outside, septa glabrous to densely hairy, usually green to sometimes light yellow to whitish violet to reddish to brown; pericarp 0.5–1.5 mm thick; column 10–14.5 mm long, straight; pedicel 3.5–14 mm long, upper part 0.3–7 mm long. Seeds globose, laterally flattened, 8–12.5 by 7.5–11 by 2–4 mm; arillode translucent; cotyledons 5–8.4 by 7–10 by 0.1–2 mm; radicle 0–1.3 mm long; endosperm cream.

Distribution — Peninsular Malaysia, Sumatra, Borneo, (Moluccas).

Habitat & Ecology — Primary and secondary rain forest, and peat swamp forest. Altitude: sea level up to 650 m. Flowering: February to July, October, November; fruiting: March, June to October.

Uses — Arillode edible, sweet to sour.

Vernacular names — Peninsular Malaysia: Ampai ka, tampoi, tik. Sumatra: Rambai, rambaiklih lanang. Borneo: Jelenteh (Iban); keliw.

14. Baccaurea maingayi Hook.f.

Baccaurea maingayi Hook.f., Fl. Brit. India 5 (1887) 370; Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 281; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 63; Ridl., Fl. Malay Penins. 3 (1924) 246; Whitmore, Tree Fl. Malaya 2 (1973) 65; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 51; Haegens, Blumea Suppl. (2000) 126, Map 3.16. — Lectotype (designated by Haegens, 2000): Maingay KD 1415 (lecto K; isolecto L), Peninsular Malaysia.

Tree 10–22 m high, dbh 16–32 cm, buttresses absent; branchlets densely hairy, Terminalia branching pattern weakly developed. Indumentum of simple and stellate hairs. Bark red to brown when fresh, thin; inner bark red to pink. Leaves: petiole 31–120 mm long, glabrous to densely hairy, brown to black when dry, raised glands absent; stipules elliptic to ovate, 3–7.5 by 1.5–2.8 mm, velutinous outside, glabrous to rarely sparsely hairy inside, margin ciliate, not hyaline; lamina elliptic to ovate, 8–19.5 by 3.8–10.2 cm, l/w ratio 1.6–2.9, papery; base rounded to cordate to attenuate; apex cuspidate, 0–17 mm long; upper surface glabrous, raised glands normally present, often granulate, green to brown when dry; lower surface densely hairy, raised glands, normally present, absent, discoid glands absent to present, brown to green-brown when dry; secondary veins 8–11 per side, closed at margin; nervation scalariform; young leaves black when dry. Staminate inflorescences axillary to just below the leaves (to ramiflorous), solitary to 3 clustered together, 0.7–5 cm long, c. 1 mm thick, densely hairy to velutinous, 14–many-flowered, flowers clustered at the tip of inflorescence rarely somewhat scattered; branchlets cylindrical, 1.5–3 mm long, densely hairy to velutinous, 2-flowered; bracts absent. Staminate flowers 1–1.3 mm diam.; pedicel 0–1 mm long; sepals 4, ovate or variable in shape, 0.5–0.9 by 0.3–0.8 mm, straight, velutinous outside and inside; stamens 4, 0.2–0.5 mm long, glabrous; ﬁlaments 0.1–0.4 mm long, geniculate to straight; anthers c. 0.1 by 0.2 by 0.1 mm; pistillode absent. Pistillate inflorescences ramiflorous, solitary, c. 8.5 cm long, c. 1 mm thick, densely hairy to velutinous, 9–20-flowered; pedicel 3–3.5 mm long, upper part c. 0.5 mm long, geniculate (90° bent), densely hairy; bracts 3 per branchlet, densely hairy outside and inside, margin ciliate. Pistillate flowers 2–3 mm diam.; sepals 6, ovate, 1.8–3 by 0.9–1.4 mm, velutinous outside and inside, persistent; ovary globose, c. 1.3 by 1.3 mm, 3-locular, velutinous; style absent; stigmas small, persistent. Infructescences 6–19(–45) cm long, 1–3 mm thick. Fruits globose, 0–2-seeded, fleshy capsules, 10–20 by 14–22 by 13–20 mm, often loculicidally dehiscent, raised glands sometimes present, velutinous outside, densely hairy to velutinous inside, green to orange; pericarp c. 1 mm thick; column 9–15 mm long, curved to straight, caducous to persistent; pedicel 7.5–13 mm long, upper part 2.5–5 mm long. Seeds ellipsoid, laterally flattened, 8–16 by 7–14 by 2–4 mm; arillode amber; cotyledons 6.5–7.2 by 6–7 by < 0.1 mm.

Distribution — Peninsular Malaysia, Simeuluë Island, Borneo.

Habitat & Ecology — Primary rain forest and peat swamp forest. Altitude: 80–500 m. Flowering: April, May, October; fruiting: March to August, December.

Uses — Arillode edible, sweet.

15. Baccaurea microcarpa (Airy Shaw) Haegens

Baccaurea microcarpa (Airy Shaw) Haegens, Blumea Suppl. (2000) 127, Fig. 3.12, Map 3.17. — Baccaurea nesophila Airy Shaw var. microcarpa Airy Shaw, Kew Bull. 33 (1979) 530. — Type: Brass 27290 (holo K; iso L, US), Papua New Guinea, Fergusson Island, Agamoia.

Tree c. 10 m high, dbh 25–30 cm; branchlets densely hairy when young, blackish when dry, Terminalia branching pattern weakly developed. Indumentum of simple hairs. Leaves: petiole 11–26 mm long, densely hairy when young, sometimes longitudinal cracks present, raised glands absent; stipules c. 2.8 by 1.4 mm, densely hairy outside, margin ciliate, not hyaline; lamina obovate, 5.2–11.8 by c. 2–4.9 cm, l/w ratio 2.4–2.6, papery; base attenuate to cuneate; apex obtuse to acute, up to 5 mm long; upper surface glabrous to subglabrous, raised glands absent, granulate, greenish brown when dry, nervation reddish brown, sometimes slightly sunken; lower surface subglabrous, sparsely hairy at veins, small raised glands present, discoid glands absent to present, reddish brown when dry; secondary veins 5–7 per side, closed at margin; nervation reticulate, brown. Staminate inflorescences unknown. Pistillate inflorescences axillary to just below the leaves, solitary, densely hairy, few to c. 20-flowered. Pistillate flowers: sepals caducous; ovary 2-locular; stigmas persistent. Infructescences 3–7 cm long, c. 1 mm thick. Fruits globose, 1- or 2-seeded, fleshy capsules, 9–10.5 by c. 10 by 10 mm, dehiscing irregularly, raised glands present, sparsely hairy outside, glabrous inside, green; pericarp c. 0.5 mm thick; pedicel 2.2–3.5 mm long, upper part 1–2 mm long. Seeds globose, laterally flattened, c. 6.5 by 7.5 by 4 mm; cotyledons c. 4.5 by 6 by 0.1 mm; radicle c. 1 mm long; endosperm c. 1 mm thick.

Distribution — Papua New Guinea: Fergusson Island.

Habitat & Ecology — Altitude c. 200 m. Flowering: June.

Note — Only known from the type specimen.

16. Baccaurea minor Hook.f.

Baccaurea minor Hook.f., Fl. Brit. India 5 (1887) 370; Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 281; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 64; Ridl., Fl. Malay Penins. 3 (1924) 245; Airy Shaw, Kew Bull. 16 (1963) 342; Soejarto, Bot. Mus. Leafl. 21 (1965) 71; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 37; Whitmore, Tree Fl. Malaya 2 (1973) 66; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 51; Kew Bull. 36 (1981) 264; Haegens, Blumea Suppl. (2000) 129, Fig. 3.13, Map 3.18; Haegens & Welzen in Welzen et al., Thai For. Bull. 28 (2000) 71; in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 112.— Lectotype (designated by Haegens, 2000): King’s Collector 10287 (lecto L; isolecto K), Malakka, Perak.

Aporosa billitonensis Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 97; Airy Shaw, Kew Bull. 16 (1963) 342. — Lectotype (designated by Haegens, 2000): Van Rossum 28 (holo B†; lecto L; isolecto K), Billiton Isl.

Baccaurea pendula Merr., Univ. Calif. Publ. Bot. 15 (1929) 152; Airy Shaw, Kew Bull. Add. Ser. (1975) 51. — Lectotype (designated by Haegens, 2000): Elmer 21508 (holo PNH†; lecto L; isolecto DS, GH, K, US), Borneo, Tawau.

Tree 5–35 m high, dbh up to 8–70 cm; buttresses absent or present, up to 1 m high, 1 m long; branchlets subglabrous, Terminalia branching pattern well-developed. Indumentum of simple hairs. Bark red-brown to brown or black, rarely greyish when fresh, brown to greyish brown when dry, up to 8 mm thick, smooth to rough, soft, flaking with small rectangular ﬁbrous flakes, c. 1 cm wide, 6–7 mm thick; inner bark red to brown, up to 2.5 mm thick, ﬁbrous. Leaves: petiole 7–35 mm long, glabrous, densely hairy when young, transverse cracks present; stipules lanceolate, 1–4 by 0.7–1.5 mm, densely hairy outside and inside, margin not ciliate, not hyaline; lamina ovate to obovate, 3.5–16.3 by 1.6–8.5 cm, l/w ratio 1.5–3.2(–4.2), papery; base (rounded to) cuneate to attenuate; apex acuminate to cuspidate, 1–20 mm long; upper surface glabrous, granulate, brown to greenish brown to white when dry, nervation brown; lower surface glabrous, midrib rarely densely hairy, discoid glands 0–13, often between two secondary veins or in a row at 0–5 mm distance of the midrib, (whitish to greenish) brown when dry; secondary veins 4–8 per side, closed at c. 3 mm from the margin; nervation reticulate to weakly scalariform, brown. Staminate inflorescences axillary to just below the leaves, solitary to 3 clustered together, 1–5 cm long, c. 1 mm thick, densely hairy, 25–60-flowered, flowers clustered at the tip of inflorescence; bracts 1 per branchlet, c. 4.5 mm long, caducous, sparsely to (densely) hairy outside, margin not ciliate; bracteoles 3 per branchlet, small; branchlets cylindrical to spatulate, 1.5–2 mm long, densely hairy, 1- or 2-flowered. Staminate flowers 0.8–2 mm diam., clustered at the tip of the branchlets, greenish yellow to yellow, rarely flushed pink; pedicel absent; sepals 4 (or 5), not uniform in shape, 0.3–1 by 0.3–1 mm, apex straight, densely hairy outside and inside; stamens 4 (or 5), 0.2–1 mm long, glabrous; ﬁlaments 0.2–1 mm long, laterally flattened, straight; anthers 0.1–0.2 by 0.1–0.2 by c. 0.1 mm when dehisced; disc absent; pistillode sometimes present, globose, up to 0.5 mm high, densely hairy, solid. Pistillate inflorescences ramiflorous, solitary to 4 clustered together, 4.5–28 cm long, 1–2 mm thick, densely hairy except at base, 3–20-flowered; pedicel 2–2.5 mm long, abscission zone present, upper part 1–1.5 mm long, densely hairy, pale green; bracts 3 per branchlet, persistent, densely hairy outside and inside, margin ciliate. Pistillate flowers 3–8 mm diam.; sepals 4, ovate to obovate, each with a different size and shape, 2.2–4.5 by 1–2.2 mm, densely hairy at both sides, green when fresh, brown when dry, persistent; staminodes rarely present; ovary cylindrical, 1.5–3 by 1.2–2.6 mm, 3-locular, densely hairy; style 0–3 mm long, tomentose; stigmas 0.8–1.4 mm long, apically cleft for upper 90%, persistent; lobes 0.7–1.3 by 0.3–0.6 mm, glabrous above, hirsute below, with rather long whitish hairs. Fruits globose, 3-seeded, fleshy capsules, 8–14 by 10–15 by 10–15 mm, dehiscing septicidally and loculicidally, raised glands absent, densely hairy outside, velutinous inside, orange to ochre; pericarp 0.8–1.7 mm thick; column 9–13 mm long, usually persistent, straight; pedicel 3–7 mm long, upper part c. 1 mm long. Seeds almost globose but strongly laterally flattened, 5.9–9.5 by 5.4–8.3 by 0.1–0.3 mm, flat; seed-coat red; arillode dull, yellow to orange; cotyledons 4–6 by 5–8 by c. 0.1 mm; radicle up to 2.2 by 1 mm.

Distribution — Peninsular Malaysia, Sumatra, Borneo.

Habitat & Ecology — Primary, old secondary, or bamboo forest, on ridges or slopes. Soil: granite, yellow sand or yellow sandy loam. Altitude: sea level up to 1200 m. Flowering: March to September; fruiting: June to January.

Uses — Used as timber in house construction. Arillode edible, sour-bitter.

Vernacular names — Peninsular Malaysia: Jentek-jentek, kaum tampoi, tampoi, rambai, tinlek-tinlek. Sumatra: Petik. Borneo: Sintak-nyabor (Iban); ubah merah (Malay); bua sarotic, djentikan, obar nasi.

17. Baccaurea mollis Haegens

Baccaurea mollis Haegens, Blumea Suppl. (2000) 131, Fig. 3.14, Map 3. — Type: S (Blicher, Rantai et al.) 60306 (holo L), Malaysia, Sarawak, Nanga Sebatu, Memgiong, Baleh.

Tree 9–16.5 m high, dbh 14–19 cm, buttresses low if present; branchlets sparsely hairy to velutinous, Terminalia branching pattern weakly developed. Indumentum of stellate and simple hairs. Bark red-brown when fresh; inner bark red. Leaves: petiole (34–)38–90 mm long, velutinous, raised glands absent; stipules 10.5–17 by 2.5–4 mm, late caducous, velutinous outside and inside, margin ciliate, not hyaline; lamina ovate to elliptic (to obovate), (8.8–)11.7–27.6 by (4.6–)6.3–12.1 cm, l/w ratio 1.6–2.9, papery; base truncate to acute; marginal glands as identiﬁcation marks; apex acuminate to cuspidate, 4–16 mm long; upper surface (sub)glabrous, not granulate, nervation sunken; lower surface densely hairy, midrib and secondary veins velutinous, raised glands few to absent, discoid glands often present; secondary veins (6–)9–12 per side, closed at margin; nervation reticulate. Staminate inflorescences ramiflorous, solitary to few clustered together, 3–4 cm long, 0.6–1.5 mm thick, velutinous, branching minute, many-flowered, flowers scattered along inflorescence; bracts 1 per branchlet, 1.2–2 mm long, densely hairy outside and inside, margin ciliate, not hyaline; bracteoles absent; branchlets cylindrical, 0.5–1 mm long, densely hairy, 3-flowered. Staminate flowers 1–1.5 mm diam., yellow-green; pedicel c. 0.1 mm long; sepals (3 or) 4 or 5, ovate, 0.6–1 by 0.6–1.2 mm, apex recurved, outside and inside densely hairy; staminodes absent or 4 or 5; stamens 4, c. 0.3 mm long, glabrous; ﬁlaments c. 0.15 mm long, straight; anthers c. 0.15 by 0.2 by 0.1 mm; disc absent; pistillode cylindrical, c. 0.4 mm high, densely hairy, solid. Pistillate inflorescences ramiflorous to cauline, solitary to 2 clustered together, 1.5–3 cm long, 1.5–3 mm thick, velutinous, more than 20-flowered; pedicel c. 2 mm long, upper part c. 1 mm, velutinous; bracts 3 per branchlet, velutinous outside and inside, margin ciliate. Pistillate flowers c. 5 mm diam.; sepals 4 of which 2 adnate, ovate, c. 2 by 1.5 mm, outside and inside densely hairy, persistent; disc absent; ovary globose, c. 3 by 3.5 mm, 2-locular, velutinous; style 0–0.1 mm long, c. 1 mm wide, densely hairy; stigmas 1.2–1.5 mm long, cleft for upper c. 80%, usually persistent; lobes c. 1 by 1 mm, subglabrous above, densely hairy below, protuberances above and below. Infructescences 1–4.5 cm long, 3–6 mm thick. Fruits globose, 1–4-seeded berries, 27–36 by 26–38 by 26–38 mm, raised glands usually absent, not ruminate, glabrous to sparsely hairy outside, subglabrous to sparsely hairy inside, light (yellow-)brown; pericarp (1–)2–6 mm thick; column 20–26.5 mm long, straight, persistent; pedicel 2–7 mm long, upper part 1–2 mm long. Seeds globose to ellipsoid, laterally flattened, 12–15 by 8.2–15 by 3.5–7 mm; arillode deep red; cotyledons 8.5–9.2 by 6–13 by 0.1–0.3 mm; radicle 1–1.5 mm long; endosperm 0.1–0.3 mm thick.

Distribution — Peninsular Malaysia, Sumatra, Borneo.

Habitat & Ecology — Rain forest. Altitude: 60–550(–1800) m. Flowering: March, April, July; fruiting: February, April, June, July, November.

Uses — Arillode and pericarp edible, sweet-sour.

Vernacular names — Borneo (Sarawak): buah telok kejira, buah pekan (Iban); lebek (Kayan); tampoi (Malay).

Note — The species is named for its dense soft indumentum.

18. Baccaurea motleyana (Müll.Arg.) Müll.Arg.

Baccaurea motleyana (Müll.Arg.) Müll.Arg. in DC., Prodr. 15, 2 (1866) 461; Hook.f., Fl. Brit. India 5 (1887) 371; Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 281; K.Heyne, Nutt. Pl. Ned. Ind. 3 (1917) 74; Merr., J. Straits Branch Roy. Asiat. Soc. (1921) 331; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 53; Ridl., Fl. Malay Penins. 3 (1924) 250; Corner, Wayside Trees Mal. 1 (1940) 240; Backer & Bakh.f., Fl. Java 1 (1964) 454; Soejarto, Bot. Mus. Leafl. 21 (1965) 9, 10, 12, 73; Airy Shaw, Kew Bull. 26 (1972) 220; Whitmore, Tree Fl. Malaya 2 (1973) 66; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 52; 8 (1980) 35; Kew Bull. 36 (1981) 264; Haegens, Blumea Suppl. (2000) 134, Fig. 3.15, Map 3.20, Photo 5, 6; Haegens & Welzen in Welzen et al., Thai For. Bull. 28 (2000) 71; in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 113. — Pierardia motleyana Müll.Arg., Flora 47 (1864) 516. — Type: Motley 773 (holo K), Borneo, near Banjarmasin.

Baccaurea pubescens Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 61; Ridl., Fl. Malay Penins. 3 (1924) 250. — Type: Wawra 297a (holo W), Singapore.

Tree 8–20 m high, dbh 13–50 cm, buttresses short; branchlets densely hairy, Terminalia branching pattern weakly developed. Indumentum of simple hairs. Bark brown to red-brown to yellowish when fresh, thin, soft, scaly to flaky; inner bark cream, turning slightly reddish, hard. Leaves: petiole 23–84 mm long, densely hairy to velutinous, brown-grey when dry, raised glands absent; stipules 5–10 by 1.5–3.5 mm, glabrous to sparsely hairy outside, glabrous inside, midrib velutinous outside, margin ciliate, hyaline or not; lamina elliptic to obovate, 13.5–37 by 5.2–15 cm, l/w ratio 1.9–2.5, papery: base rounded to usually narrowly cordate; apex acute cuspidate, (0–)7–22 mm long; upper surface glabrous, not granulate, (dark) green when fresh, brown to green to grey when dry; lower surface densely hairy to sparsely hairy, midrib and secondary veins velutinous, raised glands absent, discoid glands absent, (dark) green when fresh, brown to green when dry; secondary veins 10–17 per side, open at margin; nervation scalariform; young leaves black when dry. Staminate inflorescences cauline to axillary, solitary to 5 clustered together, 2.5–24 cm long, up to 1.5 mm thick, velutinous, many-flowered, flowers scattered along inflorescence, greenish yellow; bracts persistent, elliptic, 1–2.5 mm long, 3 per branchlet, glabrous outside and inside, densely hairy at base inside, margin ciliate, not hyaline; bracteoles absent; branchlets absent to present, cylindrical, 0–3 mm long, densely hairy, 3-flowered. Staminate flowers 2–3 mm diam., green to yellow; pedicel 1.6–2.3 mm long, upper part 0.6–1.6 mm long, densely hairy; sepals 4 or 5, ovate, 0.7–1.4 by 0.5–1.4 mm, usually recurved, densely hairy outside and inside; stamens 5–10, 0.4–0.6 mm long, glabrous; ﬁlaments 0.4–0.6 mm long, geniculate to straight; anthers c. 0.1 by 0.2 by 0.1 mm; disc present, c. 0.1 mm thick; pistillode obtriangular, 0.4–0.5 mm high, velutinous, hollow. Pistillate inflorescences ramiflorous to cauline, solitary to 5 clustered together, 13–42 cm long, 1–2.5 mm thick, velutinous, more than 30-flowered, greenish yellow; pedicel 2–5.5 mm long, upper part 0.2–1 mm long, densely hairy to velutinous; bracts 3 per branchlet, persistent, velutinous outside, glabrous to densely hairy inside, margin ciliate. Pistillate flowers 4–10 mm diam., green to yellow; sepals 4 or 5, ovate, 4–8 by 1.5–3.5 mm, densely hairy outside and inside, persistent; ovary globose, 1.6–2.6 by 1.7–2.9 mm, 3-locular, velutinous; style 0–0.4 by 0.8–1.2 mm wide, velutinous; stigmas 0.5–1 mm long, glabrous above and below, persistent. Fruits globose to ellipsoid, 3-seeded berries, 22–45 by 15–25 by c. 25 mm, raised glands present, glabrous to densely hairy outside, glabrous inside, greenish yellow to white; pericarp 0.5–1.5 mm thick; column 18–23 mm long, straight, caducous; pedicel 4–10 mm long, upper part 1–2 mm long. Seeds ellipsoid, laterally flattened, 13–20 by 9–14.5 by 2.5–4 mm; arillode translucent white to rarely purple; cotyledons 5–9 by 5–9 by c. 0.1 mm; radicle 2–3 mm long.

Distribution — Peninsular Malaysia, Sumatra, Borneo, Halmahera.

Habitat & Ecology — Primary and secondary rain forest, and rarely riparian forest, often cultivated. Soil: yellow clay, sand or limestone. Altitude: 15–500 m. Flowering: January to May, August, October, November; fruiting: January, May, July to September, November, December.

Uses — Locally grown for fruits: arillode edible, sour to sweet. Squeezed cambium and inner bark used as a remedy for sore eyes.

Vernacular names — Peninsular Malaysia: Rambai. Sumatra: Rambai. Borneo: Pekan (Iban); rambai (Iban, Malay); ulup-lavai (Punan).

19. Baccaurea multiflora Burck ex J.J.Sm.

Baccaurea multiflora Burck ex J.J.Sm., Icon. Bog. 4 (1910) 37, t. 312; K.Heyne, Nutt. Pl. Ned. Ind. 3 (1917) 74; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 67; Airy Shaw, Kew Bull. 36 (1981) 264; Haegens, Blumea Suppl. (2000) 136, Fig. 3.16, Map 3.21. — Lectotype (designated by Haegens, 2000): Teijsmann HB 3504 (lecto L), Sumatra, Bangka.

Tree c. 8 m high, dbh 10–20 cm, buttresses absent; branchlets glabrous to sparsely hairy, densely hairy when young; young shoots dark brown to blackish when dry, Terminalia branching pattern weakly developed. Indumentum of stellate and simple hairs. Bark grey-brown to dark brown when fresh, c. 2 mm thick; outer bark ﬁnely ﬁssured; inner bark pale brown, c. 0.5 mm thick. Leaves: petiole 8–45 mm long, glabrous to sparsely hairy, transverse cracks present, dark brown to blackish when dry, raised glands often present; stipules 2.5–5 by 2–3 mm, densely hairy to velutinous outside, subglabrous inside, margin ciliate, not hyaline; lamina obovate, 6.7–13.2 by 2.5–5.9 cm, l/w ratio (1.3–)1.9–2.7(–3.1), (slightly) leathery; base cuneate to attenuate; apex obtuse to acute, up to 11 mm long; upper surface glabrous, raised glands usually present, green when fresh, dark (green-, or grey-)brown when dry; lower surface glabrous, nervation sometimes subglabrous, raised glands usually present, discoid glands absent, pale yellow-green when fresh, (pale) (yellow-)brown with darker nervation below when dry; secondary veins 4–7 per side, often closed at margin; nervation reticulate, indistinct. Staminate inflorescences axillary to ramiflorous, solitary to few clustered together, 1–9.5 cm long, 0.5–1 mm thick, branched, subglabrous to densely hairy, many-flowered, flowers usually scattered along inflorescence, cream-yellow; bracts usually present, leaf-like; bracts 1 per branchlet, 1–3.5 mm long, sparsely to densely hairy outside, subglabrous to densely hairy inside, margin ciliate, not hyaline; bracteoles c. 0.5 mm long, subglabrous outside; branching cylindrical, 1.5–12 mm long, densely hairy, 3–10-flowered. Staminate flowers 1–2 mm diam., cream-yellow; pedicel 1–3.3 mm long, upper part 0.6–1.8 mm long, densely hairy; sepals 4 or 5, ovate to obovate, 0.5–1.8 by 0.4–0.9 mm, apex recurved, outside and inside densely hairy, cream-yellow; stamens 3–6, 0.2–0.5 mm long, glabrous; ﬁlaments 0.1–0.4 mm long, straight; anthers 0.1–0.15 by 0.2–0.3 by c. 0.1 mm; disc absent; pistillode globose, up to 0.2 mm high, velutinous, solid. Pistillate inflorescences probably ramiflorous, few clustered together, c. 3.3 cm long, 0.8–1 mm thick, densely hairy to velutinous, many-flowered; pedicel 1–4 mm long, upper part 0.7–3 mm, velutinous; bracts 3 per branchlet, 2 small, densely hairy outside, (sub)glabrous inside, margin ciliate. Pistillate flowers 2.5–6.8 mm diam.; sepals (4 or) 5, ovate to obovate, 2–3.5 by 1–1.9 mm, outside and inside velutinous, caducous; ovary fusiform to cylindrical, 1.8–4.5 by 1.5–3.6 mm, 2-locular, velutinous; style 0.3–0.9 by 0.9–1.2, densely hairy; stigmas 0.5–1.5 mm long, cleft for upper 0–50%, caducous; lobes 0.5–0.7 by 0.2–0.5 mm, glabrous above, densely hairy below, protuberances above and below. Fruits globose, berries, 24–28 by 22–23 by 22–23 mm when dry, raised glands present, velutinous outside and inside, peel brownish; pericarp 4–6 mm thick; column c. 20 mm long, straight; pedicel 8–9 mm long. Seeds only seen immature; arillode whitish.

Distribution — Sumatra.

Habitat & Ecology — Primary rain forest. Soil: yellow-red loam. Altitude: 50–900 m. Flowering: August (and July, November in cultivation); fruiting: December.

Vernacular names — Sumatra: Rambay hutan, roesip, temoeleëng. Simeuluë Island: Bolawah anteu.

20. Baccaurea nanihua Merr.

Baccaurea nanihua Merr., Interpr. Rumph. Herb. Amboin. (1917) 315; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 67; J.J.Sm., Bull. Jard. Bot. Buitenzorg 3 (1924) 94; K.Heyne, Nutt. Pl. Ned. Ind., 3rd ed., 1 (1950) 914; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 52; Kew Bull 37 (1982) 8; Haegens, Blumea Suppl. (2000) 138, Map 3.22. — Type: Robinson 331 (holo A; iso G, SING, US), Moluccas, Ambon, Koesoekoesoe and Mahija.

Baccaurea tristis Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 69; Airy Shaw, Kew Bull. Add. Ser. 8 (1980) 37. — Lectotype (designated by Haegens, 2000): Ledermann 9811 (lecto L), Papua New Guinea.

Treelet or tree 5–25 m high, dbh 8–40 cm; bole sometimes fluted up to 1 m; mouldings 20 cm out, 10 cm thick; branchlets subglabrous, densely hairy when young; young shoots brown to grey when dry, Terminalia branching pattern weakly developed. Indumentum of stellate and simple hairs. Bark red to red-brown to grey-brown to dark brown when fresh, 0.2–2 mm thick, sometimes slightly peeling off; inner bark red to red-brown to yellowish red, 0.5–3 mm thick. Heartwood pale red to reddish yellow. Leaves: petiole 16–64 mm long, subglabrous to densely hairy, transverse cracks present, apically and sometimes basally pulvinate, raised glands rarely present; stipules 3.5–7.5 by 1–3 mm, velutinous outside, glabrous to densely hairy inside, margin ciliate, not hyaline; lamina ovate to elliptic (to obovate), 6.5–21(–25) by 2.5–10.8 cm, l/w ratio 1.4–2.9, papery to leathery; base attenuate to rounded; marginal glands sometimes visible as small indentations; apex (broadly) acuminate, up to 11 mm long; upper surface glabrous (to densely hairy at midrib), raised glands often present, sometimes granulate, dark brown to grey-brown when dry, nervation dark brown to grey-brown; lower surface glabrous to sparsely hairy (to densely hairy at midrib and veins), raised glands often present, discoid glands if present in a row between secondary veins, light brown when dry; secondary veins 6–10 per side, open at margin; nervation reticulate to weakly scalariform, (dark) brown; young leaves green. Staminate inflorescences axillary (to ramiflorous), solitary to few clustered together, 1–13.5 cm long, up to 1 mm thick, branching present, densely hairy, many-flowered, flowers scattered along inflorescence, rachis pale yellow, bracts leaf-like (up to 2 cm long) usually present; bracts 1 per branchlet, 1.5–3 by 1–1.2 mm, densely hairy outside, subglabrous to densely hairy inside, margin ciliate, not to slightly hyaline; bracteoles 0.5–1.5 mm long, outside and inside densely hairy; branchlets cylindrical, 2–35 mm long, densely hairy, 3–many-flowered. Staminate flowers 1.5–2.5 mm diam., pale yellow to whitish; pedicel 1–2 mm long, upper part 0.6–1.2 mm long, densely hairy, reddish-brown; sepals 4–6, ovate, 0.9–1.5 by 0.3–1 mm, sometimes apically recurved, outside and inside densely hairy; stamens 4–6, 0.3–0.6 mm long, glabrous, brownish; ﬁlaments 0.15–0.5 mm long, straight; anthers 0.1–0.2 by 0.15–0.25 by up to 0.1 mm; disc absent; pistillode globose, 0.1–0.4 mm high, velutinous, solid. Pistillate inflorescences ramiflorous to axillary, solitary, 0.5–4 cm long, 1–2 mm thick, velutinous, few-flowered; pedicel c. 4.5 mm long, upper part 1–1.5 mm long, densely hairy; bracts 1 per branchlet, ovate, velutinous outside, densely hairy inside, margin ciliate. Pistillate flowers 3–7 mm diam.; sepals 5 (or 6), ovate, 1.5–3 by 1–1.5 mm, velutinous outside and inside, usually persistent; ovary globose to cylindrical, c. 2.5 by 2.5 mm, (2- or) 3- (or 4-)locular, velutinous; style c. 1 by c. 1.5 mm, velutinous; stigmas c. 0.1 mm long, persistent to caducous. Fruits globose to suppressed globose, 0–6-seeded, fleshy capsules, 14–25 by 25–29 by 25–29 mm, loculicidally dehiscent, raised glands usually present, densely short hairy outside, sparsely hairy to hirsute inside, septa densely long hairy, (yellowish to reddish) brown; pericarp 3–6.5 mm thick; column 11–13.5 mm long, straight; pedicel 8.5–15 mm long, upper part 3.5–8 mm long. Seeds ellipsoid, laterally flattened, to long triangular, 7.5–11.5 by 3.8–7 by 3–4.7 mm; cotyledons 5–5.5 by c. 5 by 0.1 mm; radicle c. 0.7 mm long; endosperm 2–2.5 mm thick.

Distribution — Borneo, Sulawesi, Moluccas, New Guinea.

Habitat & Ecology — Primary rain forest, riverine rain forest, and edges of mangrove. Soil: sandy loam. Altitude: sea level up to 600 m. Flowering: June to January; fruiting: January, April to July, September, November, December.

Vernacular names — Moluccas: Maroontaboea. New Guinea: Garang kafak (Tehid); mabab (Biak); senesasau (Mekeo); pei (Biak).

21. Baccaurea nesophila Airy Shaw

Baccaurea nesophila Airy Shaw, Kew Bull. 33 (1979) 529; Kew Bull. Add. Ser. 8 (1980) 35; Haegens, Blumea Suppl. (2000) 140, Map 3.23. — Type: Brass 27665 (holo K; iso L, US), Papua New Guinea, Mimisa Island, Quark Mt.

Tree 8–20 m high, dbh 10–30 cm, slightly buttressed; branchlets densely hairy when young, brown to blackish when dry, Terminalia branching pattern weakly developed to sometimes strong. Indumentum of simple and rarely a few stellate hairs. Bark brown to reddish when fresh. Heartwood pinkish. Leaves: petiole 8–31 mm long, glabrous to sparsely hairy, densely hairy when young, raised glands absent; stipules 2.5–6 by 1.5–2 mm, glabrous on both sides, densely hairy when young, margin ciliate when young, (not) hyaline; lamina elliptic to obovate, 3.5–12.9 by 1.9–6.2 cm, l/w ratio 1.6–2.4, papery to slightly leathery, base attenuate to cuneate; apex (retuse to) obtuse to slightly acute, 0–3 mm long; upper surface (sub)glabrous, granulate above, nervation not to slightly sunken, glossy dark green when fresh, green to brown to reddish when dry; lower surface subglabrous, raised glands absent, discoid glands many, dark to pale green below when fresh, (light) green to brown below when dry; secondary veins 4–9 per side, closed at margin; nervation reticulate, brown to red when dry. Staminate inflorescences unknown. Pistillate inflorescences ramiflorous to axillary, solitary to rarely 5 clustered together, few to c. 15-flowered. Pistillate flowers: sepals caducous; ovary 2-locular; stigmas persistent. Infructescences 2.6–7 cm long, 0.5–1.5 mm thick. Fruits globose to ellipsoid to pear-shaped, 2–4-seeded, fleshy capsules, 16–21 by 12–20 by 12–20 mm, dehiscing loculicidally, (sometimes septicidally also), raised glands absent to present, glabrous to sparsely hairy outside, glabrous to subglabrous inside, green to yellow; pericarp 1–1.8 mm thick; column 12–17 mm long, straight to slightly curved; pedicel 2.1–3.6 mm long, upper part 1–2.5 mm long. Seeds globose to ellipsoid, laterally flattened, 8.8–12.5 by 5.7–12 by 3–5 mm; arillode red to orange; testa cream to whitish; cotyledons 5.5–9 by 6–11 by c. 0.1 mm; radicle 1–2 mm long; endosperm c. 1.5 mm thick.

Distribution — Papua New Guinea: Louisiade Archipelago.

Habitat & Ecology — Swamp forest to ridge tops. Altitude: 0–510 m. Fruiting: April, June to November.

Vernacular names — Louisiade Archipelago: Dumi, tum.

22. Baccaurea odoratissima Elmer

Baccaurea odoratissima Elmer, Leafl. Philipp. Bot. 4 (1911) 1276; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 59; Merr., Enum. Philipp. Fl. Pl. 2 (1923) 411; Fernando, Philipp. J. Biol. 8 (1979) 307; Haegens, Blumea Suppl. (2000) 141, Fig. 3.17, Map 3.24, Photo 7. — Lectotype (designated by Haegens, 2000): Elmer 13160 (lecto L; isolecto GH, K, US), Philippines, Palawan, Puerto Princesa.

Baccaurea membranacea Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 49; Airy Shaw, Kew Bull. Add. Ser. 4 1975) 54. — Lectotype (designated by Haegens, 2000): Hose 600 (lecto L, selected here; isolecto K), Borneo, Sarawak, Baram district, Miri River.

Baccaurea trunciflora Merr., Univ. Calif. Publ. Bot. 15 (1929) 151; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 37; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 56. — Lectotype (designated by Haegens, 2000): Elmer 21550 (lecto L; isolecto GH, US), Borneo, Sabah, Tawau.

Treelet to tree 2–17 m high, dbh 4.4–20 cm, buttresses absent; branchlets sparsely hairy, female plants rarely glabrous, Terminalia branching pattern well-developed. Indumentum of simple hairs, usually in combination with stellate hairs. Bark yellow to grey-brown when fresh, whitish grey when dry, 0.1–2.5 mm thick; inner bark yellow to orange-brown, 0.1–0.4 mm thick. Leaves: petiole 9–82 mm long, sparsely hairy, female plants rarely glabrous, (green- or red-)brown when dry, raised glands present; stipules 1.7–8 by 0.5–2 mm, glabrous to sparsely hairy outside, but densely hairy when young, glabrous inside, midrib densely hairy outside, margin (not) ciliate; lamina obovate to elliptic (to ovate), 6.4–19 by 2–9.9 cm, l/w ratio 1.5–3.3, papery; base acute to attenuate, apex rounded to cuspidate, up to 22 mm long; upper surface glabrous, raised glands usually present, green-brown when dry; lower surface glabrous, sparsely hairy at veins, raised glands usually present, discoid glands absent, brown when dry; secondary veins 3–8 per side, usually closed at margin; nervation reticulate to weakly scalariform; young leaves red-brown. Staminate inflorescences axillary to ramiflorous, solitary to few clustered together, 1.4–8 cm long, up to 1 mm thick, densely hairy, more than 30-flowered, flowers scattered along inflorescence; branchlets absent; bracts 3 per branchlet, c. 0.6 mm long, persistent, glabrous to densely hairy outside, glabrous inside, margin ciliate, sometimes hyaline. Staminate flowers 3 per branchlet, 1–2 mm diam., pale yellow; pedicel 0.8–2.5 mm long, upper part 0.6–2.1 mm long, densely hairy; sepals 4(–6), ovate, 0.9–1.2 by 0.7–0.9 mm, apex slightly recurved, densely hairy outside, glabrous inside; stamens 4–7, (0.2–)0.8–0.9 mm long, glabrous; ﬁlaments (0.2–)0.8 mm long, straight; anthers 0.1–0.3 by c. 0.1 by 0.1 mm; disc absent; pistillode subglobose to cylindrical, (0.2–)0.9 mm long, densely hairy, solid. Pistillate inflorescences cauline (to ramiflorous), solitary to 3 clustered together, 2–26.5 cm long, 0.5–1.5 mm thick, sparsely to densely hairy, 9–many-flowered, red; pedicel 1.2–3.5 mm long, upper part 0.5–1 mm long, densely hairy, light green to red; bracts 3 per flower, persistent, sparsely to densely hairy outside, glabrous inside, margin ciliate. Pistillate flowers 5–11 mm diam.; sepals 4, obovate, 4–6 by 2–2.7 mm, sparsely hairy outside, densely hairy inside, greenish yellow when fresh, brown outside when dry, whitish inside when dry, caducous; ovary globose, 1.2–2.5 by 1.7–2.6 mm, 2-locular, densely hairy to velutinous, light green; style 0–0.1 mm long, c. 1 mm broad; stigmas white, 0.5–0.9 mm long, not lobed, glabrous on both sides, persistent, rarely caducous. Fruits subglobose to double globose, 0–2-seeded, fleshy capsules, 7.2–12 by 8–13.5 by (4–)6.8–11, irregularly splitting, glabrous on both sides, densely hairy when young outside, raised glands usually absent, red to shiny brown; pericarp up to 0.5 mm thick; column 6–10 mm long, caducous, straight; pedicel 2.5–9.5 mm long, upper part 0.5–3 mm. Seeds subglobose to ellipsoid, laterally flattened, 6.5–10 by 2.5–8 by 2.5–6 mm; arillode blue; cotyledons 5.5–7 by 6.8–7.2 by 0.1–0.3 mm; endosperm 1.5 mm thick; radicle 1.3–1.5 mm long.

Distribution — Borneo.

Habitat & Ecology — Primary or secondary forest. Soil: basalt, yellow sandy clay, blackish sandstone. Altitude: sea level up to 1250(–1500) m. Flowering: February, April, July to October, December; fruiting: January to March, May to August, October, November.

Uses — Fruit edible.

Vernacular names — Borneo: Enkuni, kune madamayang (Iban); kukunal (Dusun Kinabatangan); kunau (Brunei, Dusun); kunau kunau (Malay); botung kratek, enkunig, enkuri, mata kunau, mata pelanduk, rambai tikus, sarotik, tampoi keli.

Note — Airy Shaw (1975) mentioned this species as a synonym of B. parviflora with a question mark. However, after studying the type material and the original descriptions, I am convinced that this species is conspeciﬁc with B. membranacea and B. trunciflora, and distinct from B. parviflora. Especially the fruit is utterly different, in the number of locules (B. odoratissima 2-locular, B. parviflora 3-locular) as well as in the fruit shape (globose in B. odoratissima, fusiform in B. parviflora).

23. Baccaurea papuana F.M.Bailey

Baccaurea papuana F.M.Bailey, Proc. Roy. Soc. Queensland 18 (1904) 3: Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 57; Airy Shaw, Kew Bull. Add. Ser. 8 (1980) 36; Haegens, Blumea Suppl. (2000) 148, Map 3.25. — Type: F.M.Barton s.n. (holo BRI), Papua New Guinea, Mekeo District.

Baccaurea plurilocularis J.J.Sm., Nova Guinea 8 (1910) 228; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 60; S.Moore, J. Bot. Br. 61 (1923) 46; Airy Shaw, Kew Bull. Add. Ser. 8 (1980) 36. — Lectotype (designated by Haegens, 2000): Versteeg 1263 (lecto L; isolecto K, U), New Guinea, Irian Jaya, base of the Nepenthes Hills.

Baccaurea montana Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 55. — Lectotype (designated by Haegens, 2000): Schlechter 18537 (lecto K; isolecto A), Papua New Guinea, Kaiser Wilhelmsland, Bismarckgebiet.

Shrub to tree 4–20 m high, dbh 6–30 cm, buttresses absent; branchlets subglabrous, Terminalia branching pattern usually strong. Indumentum of simple hairs. Bark (light) brown to (light) grey when fresh, grey when dry, thin, papery; inner bark pale brown to pale yellow, thin, medium hard. Heartwood pinkish brown. Leaves: petiole 7–60 mm long, subglabrous to densely hairy, transverse cracks absent to some present, raised glands absent to present; stipules 3–6.5 by 1–2.3 mm, glabrous, densely hairy when young outside, glabrous inside, margin ciliate, (not) hyaline; lamina (ovate to) elliptic to obovate, 8–20.5 by 2.8–10.4 cm, l/w ratio 1.6–3.3, papery; base attenuate to cuneate to rarely rounded; marginal glands not to slightly visible as identiﬁcation marks; apex (retuse to) obtuse to acute (to cuspidate), up to 13 mm long; upper surface glabrous, midrib glabrous to densely hairy, granulate, dull to shiny, pale to dark green when fresh, (greyish to greenish) brown when dry, nervation whitish to (dark) brown; lower surface subglabrous to sparsely hairy, subglabrous to densely hairy at veins, raised glands usually absent, discoid glands sometimes few present; dull to shiny, pale to dark green when fresh, (greyish to greenish) brown when dry; secondary veins 7–15 per side, closed at margin; nervation reticulate to weakly scalariform, grey-brown; young leaves reddish. Staminate inflorescences axillary to cauline, solitary to few clustered together, 0.5–23 cm long, c. 1 mm thick, sparsely to densely hairy, many-flowered, flowers scattered along inflorescence; bracts 3 (or 4) per branchlet, 0.3–1 mm long, subglabrous to densely hairy outside, glabrous inside, margin ciliate, hyaline; bracteoles absent; branchlets 0–0.4 mm long, 3-flowered. Staminate flowers 1.8–3.5 mm diam., cream to yellow, sweet scented; pedicel 1.2–4.2 mm long, upper part 0.6–3 mm, densely hairy, yellow; sepals 4 or 5, ovate to elliptic, 1–1.8 by 0.5–1.3 mm, apex straight, densely hairy outside and inside; staminodes 4–8; stamens 4–8, 0.3–0.8 mm long, glabrous; ﬁlaments 0.2–0.7 mm long, slightly geniculate to straight; anthers 0.1–0.2 by 0.2–0.3 by 0.1–0.15 mm; disc absent; pistillode globose to obtriangular, 0.3–0.6 mm high, densely hairy, hollow. Pistillate inflorescences cauline, solitary to few clustered together, 7.5–35 cm long, 0.5–2 mm thick, sparsely hairy to velutinous, 5–40-flowered, yellow; pedicel 2.5–10 mm long, upper part 0.5–2(–4) mm, densely hairy to velutinous; bracts 3 per branchlet, sparsely to densely hairy outside, glabrous inside, margin ciliate. Pistillate flowers 4–18 mm diam., greenish to yellow to creamy yellow, fragrant; sepals 5–9, lanceolate to rarely obovate, 3–11 by 0.7–4.8 mm, densely hairy outside and inside, persistent; ovary globose, 1.2–3 by 1.5–3.3 mm, 3- (or 4-) locular, woolly; style absent to 0.01 mm long, c. 1 mm wide, glabrous; stigmas 0.7–1.2 mm long, cleft for upper 0–30%, persistent; lobes c. 0.7 by 0.4 mm, glabrous above and below, ruminate above and below. Fruits pear-shaped to globose, (3–)6-seeded berries, 27.5–43 by 24–40 by 24–40 mm, raised glands absent to present, not to slightly ruminate, glabrous to densely hairy outside, glabrous inside, brown to green to red to pink to purple; pericarp 1–7 mm thick; pedicel 5–8(–23) mm long, upper part 1–3(–4) mm long. Seeds ellipsoid to globose, laterally flattened, 9–22 by 9–12 by 3.5–7 mm; arillode yellow to white; cotyledons 5.5–8 by 5–8 by < 0.1–0.4 mm.

Distribution — New Guinea and (Sulawesi).

Habitat & Ecology — Primary and secondary rain forest, and swamp forest. Soil: laterite to clay. Altitude: sea level up to 1300 m. Flowering: throughout the year; fruiting: January to October.

Uses — Arillode edible, harsh but refreshing.

Vernacular names — New Guinea: Ngesby (Hattam); apo (Maipa; Mekeo); kniebieé (Manikiong); esino (Minuﬁa); ani-abi (Noemfoer); kwame (Onjob); be(h)ga, tomarah (Orokaiva); cobo, gomo (Totubu); mapiok (Waskuk); motroka, nunuhabu, sakowgwo, yanak.

24. Baccaurea parviflora (Müll.Arg.) Müll.Arg.

Baccaurea parviflora (Müll.Arg.) Müll.Arg. in DC., Prodr. 15, 2 (1866) 462; Kurz, Forest Fl. Burma 2 (1877) 357; Hook.f., Fl. Brit. India 5 (1887) 368; Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 281; Brandis, Indian Trees (1906) 562; K.Heyne, Nutt. Pl. Ned. Ind. 3 (1917) 74; Merr., J. Straits Branch Roy. Asiat. Soc. (1921) 331; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 59; Ridl., Fl. Malay Penins. 3 (1924) 243; S.Moore, J. Bot. Br. 63 (1925) 98; Corner, Wayside Trees Mal. 1 (1940) 241; Soejarto, Bot. Mus. Leafl. 21 (1965) 89; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 37; Airy Shaw, Kew Bull. 26 (1971) 220; Whitmore, Tree Fl. Malaya 2 (1973) 64; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 52; Kew Bull. 36 (1981) 265; Enum. Euphorb. Philipp. (1983) 9; Haegens, Blumea Suppl. (2000) 152, Fig. 3.18, Map 3.26; Haegens & Welzen in Welzen et al., Thai For. Bull. 28 (2000) 72; in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 113, fig. 24a-f. — Pierardia parviflora Müll.Arg., Linnaea 32 (1863) 82; Baill., Adansonia 3 (1863) 140. — Lectotype (designated by Haegens, 2000): Wallich 1849 (lecto K), Tavoy, India Orientalis (see note 2).

Pierardia dasystachya Miq., Fl. Ned. Ind. 1, 2 (1859) 358. — Baccaurea dasystachya (Miq.) Müll.Arg. in DC., Prodr. 15, 2 (1866) 458; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 52; K.Heyne, Nutt. Pl. Ned. Ind., 3rd. ed., 1 (1950) 912; Airy Shaw, Kew Bull. 36 (1981) 260. — Lectotype (designated by Haegens, 2000): Zollinger 757z (lecto U; isolecto A, BO, G), Sumatra.

Baccaurea affinis Müll.Arg. in DC., Prodr. 15, 2 (1866) 459. — Type: Wallich 7759 B (holo G-DC; iso ?K), Tavoy, India Orientalis (see note 2).

Baccaurea scortechinii Hook.f., Fl. Brit. India 5 (1887) 368; Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 281; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 56; Ridl., Fl. Malay Penins. 3 (1924) 244; Corner, Wayside Trees Mal. 1 (1940) 242; Smitinand, Thai Forest Bull., Bot. 2 (1955) 25; Nat. Hist. Bull. Siam Soc. 20 (1961) 53; Airy Shaw, Kew Bull. 26 (1972) 221; Kiew, Gard. Bull. Sing. 49 (1997) 37. — Lectotype (designated by Kiew, 1997): Scortechini s.n. (lecto K), Malaysia, Perak. (The type bears ridged fruits.)

Baccaurea singaporica Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 54. — Type: Ridley (K?, n.v.), Singapore.

Baccaurea rostrata Merr., Univ. Calif. Publ. Bot. 15 (1929) 150; Airy Shaw, Kew Bull. 21 (1962) 355; Meijer, Bot. News Bull. Forest. Dept., Sabah 7 (1967) 37; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 52. — Lectotype (designated by Haegens, 2000): Elmer 21554 (lecto L; isolecto A, GH, K, US), Borneo, Sabah, Tawau.

Shrub to tree 3–15 m high, dbh 4–21 cm, buttresses absent, rarely fluted at base; branchlets glabrous, slightly thickened at tip, Terminalia branching pattern well-developed. Indumentum of simple hairs. Bark greenish-grey to grey-black to light yellow-brown to brown, 1.5–3.5 mm thick, rough to smooth, minutely ﬁssured in strips 5 mm wide, papery; inner bark yellow to light brown, 1.5–2 mm thick. Leaves: petiole glabrous, densely hairy when young, 4–55 mm long, brown to reddish when dry, raised glands usually present; stipules 2.5–7 by 1.5–2.5 mm, glabrous to rarely densely hairy, margin ciliate, one side with long hairs, hyaline, midrib hirsute above; lamina elliptic to obovate, 5–23 by 2–9.5 cm, l/w ratio 1.6–3.7, papery; base attenuate to cuneate; apex acuminate, up to 22 mm long; upper surface glabrous, rarely hair-domatia present at base, rarely granulate, (dark) green when fresh, red to brown to green when dry; lower surface (sub)glabrous, raised glands absent, discoid glands sometimes present, (greyish to light-)green when fresh, red to brown to green when dry; secondary veins (3–)5–10 per side, closed at 2–3 mm from margin; nervation reticulate to weakly scalariform, green to yellowish to red when dry; young leaves whitish to pinkish. Staminate inflorescences cauline, at base of trunk, many clustered together, 4–28 cm long, rachis up to 1.2 mm thick, densely hairy, many-flowered, flowers scattered along inflorescence, red to maroon to light yellow; bracts 1 per branchlet, 0.3–1.6 by 0.15–1 mm, subglabrous (to densely hairy) outside, glabrous inside, margin ciliate, sometimes hyaline; bracteoles 2, 0.2–0.4 mm long; branchlets cylindrical, 0.1–2.8 mm long, densely hairy, (1–)3-flowered. Staminate flowers 1–5 mm diam.; pedicel 0.4–6 mm long, upper part 0.2–3 mm, sparsely to densely hairy; sepals 4 or 5, obovate, each with a different size and shape, 0.5–3 by 0.5–1.2 mm, densely hairy outside, glabrous to densely hairy inside, maroon to yellow outside, cream to yellow inside when fresh, whitish outside, brownish white inside when dry; stamens 5–7, 0.5–0.9 mm long, glabrous, cream; ﬁlaments 0.4–0.8 mm long, straight; anthers 0.2–0.3 by 0.1–0.2 by 0.1–0.2 mm; rudimentary disc sometimes present; pistillode obtriangular, 0.3–1.2 mm long, hirsute, hollow inside, cream. Pistillate inflorescences cauline, at base of trunk, many clustered together, up to 28 cm long, 1–4 mm thick, densely hairy (to glabrous), red when dry, fragrant; pedicel 2–6(–9) mm long, often without a clear abscission zone, upper part 0.2–3(–7.5) mm long, densely hairy, bright red; bracts 3, up to 1.2(–1.6) by 1(–1.3) mm, persistent, sparsely to densely hairy outside, glabrous inside, margin ciliate. Pistillate flowers 5–7.5 mm diam.; sepals 4 or 5, narrowly elliptic to obovate, 2.7–8 by 0.9–2 mm, densely hairy outside, glabrous to densely hairy inside, white to yellow to usually red when fresh, caducous to persistent; ovary almost globose, with or without 6 wings, 1.3–2.2 by 1–2.3 mm, (2- or) 3- (or 4-)locular, velutinous, yellow; style 0–2 by c. 1 mm, densely hairy, yellow; stigmas 0.1–1.5 mm long, cleft apically only or up to base, caducous to persistent; lobes 0.2–1.4 mm long, glabrous with protuberances above, sparsely to densely hairy below. Fruits fusiform, 1–3-seeded berries, with or without 4–6 narrow wings, 16–32 by 7–15 mm, glabrous to sparsely hairy outside, raised glands present, dull red to maroon turning purple to black when fresh; pericarp c. 1–2.5 mm thick, glabrous on both sides; column 10–18 mm long, straight. Seeds ellipsoid, laterally flattened, 9–16 by 4–8 by 1–5 mm; arillode red to purple to white; cotyledons orbicular to square, membranous, 5.5–11 by 4–6 by 0.5–3.5 mm, radicle up to 1.5 mm long.

Distribution — Thailand, Peninsular Malaysia, Sumatra, Borneo.

Habitat & Ecology — Primary and secondary evergreen forest, steep hillsides to dry parts of swamps to alluvial forest. Soil: sandstone, white sandy soil. Altitude: sea level up to 1250 m. Flowering: November to June, September; fruiting: March to December.

Uses — Wood used as boxwood. Pericarp and arillode edible, sour-sweet.

Vernacular names — Thailand: Somfai Pah. Peninsular Malaysia: Tamun (Batec); kentamun ayer (Temuan); asam tambun, belembik, dedali, engkuni, setambun, tambun. Borneo: Bamotong-belaboh (Bassap-Mapulu); belembik plandoek, engkuni, konkuni, mata pelanduk.

Notes — 1. Baccaurea parviflora is rather heterogeneous in some characters, especially in the fruits. Kiew (1997) re-established the name B. scortechinii, based purely on material from the Peninsular Malaysia. She listed 19 differential characters. I checked all these characters, except for the fruit colour because this character is impossible to study when only herbarium material is available. She mentioned this as one of the problems of species delimitation in this complex (B. parviflora and B. scortechinii). All 19 characters listed overlap. The values given by Kiew are mean values only. Collections from Sumatra made by De Wilde & De Wilde-Duyfjes in Gunung Leuser Park overlap in all characters. According to Kiew, the most important difference is the position of the inflorescences in combination with fruit shape. Baccaurea scortechinii flowers at the trunk and branches, and has ridged fruits, while B. parviflora flowers at the base of the trunk, and has no ridges at the fruits. However, contrary to these observations, M. van Balgooy photographed a tree, which flowered at the base of the trunk, and had ridged fruits. In July 1998 I observed the same phenomenon in the FRIM campus in Kepong (Photo 8). My conclusion is that these two taxa are conspeciﬁc.

2. In 1997 Kiew erroneously typiﬁed B. parviflora. She mentioned Wallich 7759B (K) as the type specimen. This, however, is an isotype of B. affinis. In the original description of Pierardia parviflora Müller Argoviensis (1863) quoted: “Wall.! absque no. in hb. DC”. I have checked G-DC on microﬁche, but could not recover this specimen. The lectotype Wallich 1849 (K, Lectotype, designated by Haegens, 2000) has no original number and has been studied by Müller Argoviensis as notes in his handwriting are found on the sheet. Later the numbers 1849 and 7759B were added, of which number 7759B is crossed out again.

3. Another extremely closely related Burmese and Thai taxon is B. ptychopyxis. This species, however, always has axillary staminate inflorescences, and is ramiflorous to cauline only as an exception. Baccaurea parviflora is always strictly cauliflorous. The staminate inflorescences in B. ptychopyxis are solitary, whereas B. parviflora has dense clusters. Besides these differences there are some minor characters, such as the fruit length is in B. ptychopyxis 17–20.3 mm, and in B. parviflora 16–32 mm. The material of B. ptychopyxis is, however, very incomplete. Therefore B. ptychopyxis is here interpreted as a species separate from B. parviflora.

4. See note under B. odoratissima.

25. Baccaurea philippinensis (Merr.) Merr.

Baccaurea philippinensis (Merr.) Merr., Philipp. J. Sci., Bot. 10 (1915) 275; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 68; Merr., Enum. Philipp. Fl. Pl. 2 (1923) 411; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 53; Fernando, Philipp. J. Biol. 8 (1979) 303; Airy Shaw, Kew Bull. Add. Ser. 8 (1980) 36; Enum. Euphorb. Philipp. (1983) 9; Haegens, Blumea Suppl. (2000) 157, Fig. 3.19, Map 3.27. — Everettiodendron philippinense Merr., Philipp. J. Sci., Bot. 4 (1909) 279; Elmer, Leafl. Philipp. Bot. 3 (1910) 916. — Lectotype (designated by Haegens, 2000): FB (Everett) 7282 (lecto K; isolecto US), Philippines, Negros, Himugaan River.

Treelet to tree 4.5–12 m high, dbh 8–25 cm; branchlets subglabrous, velutinous when young, Terminalia branching pattern weakly developed. Indumentum of short stellate and usually longer simple hairs. Bark grey-brown when fresh; inner bark pale red-brown. Leaves: petiole 17–75 mm long, subglabrous to densely hairy, transverse cracks usually present, apically and sometimes basally pulvinate, raised glands rarely present; stipules 3–5.5 by 1.5–3 mm, velutinous outside, sparsely to densely hairy inside, margin ciliate, not hyaline; lamina ovate to elliptic (to obovate), (5–)7.2–18 by (2.4–)3.5–9 cm, l/w ratio 1.7–2.3, leathery to papery; base acute to cuneate (to rounded to attenuate); marginal glands sometimes visible as small indentations; apex acuminate to cuspidate, 2–18 mm long; upper surface glabrous, small raised glands sometimes present, rarely granulate, dark green when fresh, dark to grey-brown when dry, nervation slightly sunken; lower surface (sub)glabrous, subglabrous to densely hairy at midrib and veins, small raised glands sometimes present, discoid glands present, sometimes in a row between secondary veins, pale green when fresh, light to brown-green when dry, (dark) brown above and below when dry; secondary veins 4–7 per side, open at margin; nervation reticulate to weakly scalariform. Staminate inflorescences axillary, solitary to 3 clustered together, 1–6.5 cm long, c. 0.1 mm thick, branched, many-flowered, densely hairy to velutinous, flowers scattered along inflorescence; bracts 1 per branchlet, 1.2–3.5 mm long, velutinous outside, glabrous inside, margin ciliate, not to slightly hyaline; bracteoles (7–)9–12 per branchlet, 0.3–2 mm long; branchlets cylindrical, 1.2–8 mm long, velutinous, 9–12-flowered. Staminate flowers 1.2–2 mm diam., brownish; pedicel 0.2–1.3 mm long, upper part 0.1–0.6 mm long, velutinous; sepals 3 or 4, ovate to elliptic, 0.5–1.6 by 0.2–1.1 mm, apex slightly recurved, outside and velutinous inside; stamens (3 or) 4, 0.25–0.4 mm long, glabrous; ﬁlaments 0.1–0.2 mm long, straight; anthers 0.15–0.2 by 0.15–0.2 by 0.1–0.2 mm; disc absent; pistillode globose, absent to 0.2 mm high, velutinous. Pistillate inflorescences axillary; sepals usually caducous; stigmas persistent to caducous. Pistillate flowers: ovary 3-locular. Infructescences 1.9–2 cm long, 1.3–2 mm thick. Fruits suppressed globose, (0–)3–6-seeded, fleshy capsules, 21–34 by 26–38 by 26–38 mm, apparently loculicidally dehiscent, but normally not dehiscing, hard, raised glands usually present, subglabrous to densely hairy outside, sparsely to densely hairy inside, septa densely hairy, (yellowish) brown; pericarp 4–11 mm thick; column 16–23 mm long; pedicel 4.5–9 mm long, upper part 2–6 mm long. Seeds ellipsoid to triangular, laterally flattened, 9.5–11.5 by 5–7 by 3.5–5 mm; cotyledons 6–7 by 5–5.5 by c. 0.1 mm; radicle 1–1.5 mm long; endosperm 0.5–1 mm thick.

Distribution — Philippines, Moluccas (Halmahera).

Habitat & Ecology — Primary and secondary rain forest. Altitude: 80–450 m. Flowering: April to August; fruiting: January to June, August, October, December.

26. Baccaurea polyneura Hook.f.

Baccaurea polyneura Hook.f., Fl. Brit. India 5 (1887) 369; Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 281; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 65; Ridl., Fl. Malay Penins. 3 (1924) 248; S.Moore, J. Bot. Br. 63 (1925) 98; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 37; Whitmore, Tree Fl. Malaya 2 (1973) 67; Airy Shaw, Kew Bull. 36 (1981) 265; Haegens, Blumea Suppl. (2000) 159, Map 3.28; Haegens & Welzen in Welzen et al., Thai For. Bull. 28 (2000) 72; in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 114. — Lectotype (designated by Haegens, 2000): Griffith KD 4910 (lecto K), Peninsular Malaysia.

Baccaurea kunstleri King ex Gage, Rec. Bot. Surv. India 9, 2 (1922) 230; Ridl., Fl. Malay Penins. 3 (1924) 248; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 36; Airy Shaw, Kew Bull. 26 (1972) 219; Whitmore, Tree Fl. Malaya 2 (1973) 65; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 49; Kew Bull. 36 (1981) 263; Chakrab. & Gang., J. Econ. Taxon. Bot. 18 (1994) 425. — Lectotype (designated by Chakrab. & Gang., 1994): King’s Collector 8139 (lecto CAL), Malakka, Perak (see note), n.v.

Baccaurea hookeri Gage, Rec. Bot. Surv. India 9 (1922) 232; Ridl., Fl. Malay Penins. 3 (1924) 247; Whitmore, Tree Fl. Malaya 2 (1973) 67; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 48; Kew Bull 37 (1982) 8. — Lectotype (designated by Haegens, 2000): King’s Collector 6240 (lecto K), Malaysia, Perak, Larut.

Baccaurea cordata Merr., Univ. Calif. Publ. Bot. 15 (1929) 147; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 36; Airy Shaw, Kew Bull. 26 (1971) 219; Kew Bull. Add. Ser. 4 (1975) 47. — Lectotype (designated by Haegens, 2000): Elmer 21513 (lecto A; isolecto BO, DS, K, L, US), Borneo, Sabah, Tawau.

Tree (5–)11–30 m high, dbh 10–32 cm, buttresses small; branchlets velutinous, Terminalia branching pattern weakly developed. Indumentum of simple and stellate hairs. Bark brown to white-brown to red-brown when fresh, brown when dry, 3–5 mm thick, smooth; inner bark red to brown, 2–5 mm thick. Heartwood red to brown. Leaves: petiole 7–78 mm long, velutinous, raised glands usually absent; stipules elliptic to ovate, 2.2–10 by 0.5–2 mm, caducous (to late caducous), velutinous outside, subglabrous to velutinous inside, margin ciliate, not hyaline; lamina ovate to elliptic, 5.5–21.5 by 3.1–12.5 cm, l/w ratio 1.4–2.7, papery to slightly leathery; base rounded to cordate (to attenuate); apex (retuse to) acute, up to 11 mm long; upper surface glabrous to densely hairy, midrib and veins sparsely hairy to velutinous, sometimes granulate; lower surface sparsely hairy to velutinous, raised glands absent to present, discoid glands absent; dark green when fresh, green to brown when dry, black when young when dry; secondary veins 7–14 per side, closed at margin or almost so; nervation scalariform, brown to white-brown when dry. Staminate inflorescences axillary to just below the leaves (to ramiflorous), solitary to 5 clustered together, 1–7 cm long, c. 1 mm thick, velutinous, many-flowered, flowers clustered at the tip of inflorescence, rarely somewhat scattered; bracts 0 or 3 per branchlet, 0.2–1.3 mm long, persistent, velutinous outside, glabrous to rarely velutinous inside, margin ciliate, not hyaline; branchlets cylindrical to spatulate, 1–4 mm long, velutinous, 3-flowered. Staminate flowers 1.1–3.3 mm diam., green to yellow to greenish-white; pedicel 0–0.5 mm long, velutinous; sepals 4–6, ovate, 0.7–1.5 by 0.5–1.5 mm, apex straight, outside and inside velutinous; stamens 4–8, 0.3–0.7 mm long, glabrous, cream to brown; ﬁlaments 0.2–0.6 mm long, geniculate to straight; anthers c. 0.1 by 0.2–0.3 by 0.1 mm; pistillode absent to present, globose, 0–0.3 mm high, velutinous, solid. Pistillate inflorescences axillary to ramiflorous, solitary to 3 clustered together, 1–34.5 cm long, 0.5–4 mm thick, velutinous, 5–50-flowered, yellow; pedicel 2–5.8 mm long, upper part 0.5–2 mm long, geniculate or not, velutinous to sparsely hairy; bracts 0–3 per branchlet, persistent to caducous, velutinous outside, glabrous to velutinous inside, margin ciliate. Pistillate flowers 2.5–6.5 mm diam.; sepals 4 or 5, ovate, 0.5–5.9 by 0.3–3.5 mm, outside and inside velutinous, persistent (or caducous), ochre; ovary subglobose to pyriform, 2.5–4.5 by 2.2–4 mm, 3-locular, velutinous; style 0.1–0.2 mm by 0.5–0.8 mm, velutinous; stigmas 0.6–0.9 mm long, cleft for upper 60–90%, persistent or caducous; lobes 0.5–0.9 by c. 0.2 mm, glabrous above, glabrous to sparsely hairy below. Fruits globose to subglobose, 1–3-seeded, fleshy capsules, 10–26 by 12–26 by 12–26 mm, loculicidally dehiscent, raised glands often present, sparsely hairy to velutinous outside and inside, yellow to orange; pericarp 0.1–4 mm thick; column 9.5–19 mm long, almost straight, persistent; pedicel 5–11(–35) mm long, upper part 1–7 mm long, 80% bent present at abscission zone. Seeds ellipsoid, laterally flattened, 7.8–11 by 5–9.8 by 2–5 mm; arillode red; testa orange; cotyledons 5–9 by 5–7.5 by c. 0.1 mm; radicle 0.5–2 mm long; endosperm c. 0.5 mm thick.

Distribution — Thailand, Peninsular Malaysia, Sumatra, Borneo.

Habitat & Ecology — Primary, secondary and swamp rain forest. Soil: sand, clay, loam, granite. Altitude: 20–600 m. Flowering and fruiting throughout the year.

Uses — Arillode edible, sweet to sour.

Vernacular names — Thailand: Chum rai. Peninsular Malaysia: Jentik-jentik, rambai, rambai hutan. Simeuluë Island: Sirawi, sirawi uding, sirawi pajo. Sumatra: Djetikan betina, djentikan, djenti ompang, méràh mata, pétèkan. Borneo: Jelintik, kayu masam, simpak nyabor (Iban); enyak beruk, kapul burung, tampoi.

Note — Although I had no opportunity to study the lectotype, I am convinced that B. kunstleri is conspeciﬁc with B. polyneura. The following paratypes of B. kunstleri were studied by me: Ridley 6670, 6163, and Goodenough 5084. I was unable to separate the numerous other specimens I have studied into two taxonomic entities.

27. Baccaurea pubera (Miq.) Müll.Arg.

Baccaurea pubera (Miq.) Müll.Arg. in DC., Prodr. 15, 2 (1866) 448; Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 280; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 70; K.Heyne, Nutt. Pl. Ned. Ind., 3rd ed., 1 (1950) 914; Airy Shaw, Kew Bull. 36 (1981) 265; Haegens, Blumea Suppl. (2000) 162, Map 3.30. — Pierardia pubera Miq., Fl. Ned. Ind., Eerste bijv. (1861) 442. — Lectotype (designated by Haegens, 2000): Teijsmann HB 3503 (lecto U), Indonesia, Sumatra, Bangka.

Baccaurea latifolia King ex Hook.f., Fl. Brit. India 5 (1887) 373; Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 281; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 65; Ridl., Fl. Malay Penins. 3 (1924) 249; Whitmore, Tree Fl. Malaya 2 (1973) 67; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 50; Kew Bull. 36 (1981) 265; Chakrab. & Gang., J. Econ. Taxon. Bot. 18 (1994) 425. — Lectotype (designated by Chakrab. & Gang., 1994): King’s Collector 3559 (lecto CAL; iso K), Malakka, Perak.

Baccaurea puberula Merr., Univ. Calif. Publ. Bot. 15 (1929) 145; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 36; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 50. — Type: Elmer 21284 (holo GH; iso DS, K, L), Borneo, Sabah, Tawau.

Baccaurea elmeri Merr., Univ. Calif. Publ. Bot. 15 (1929) 146; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 48. — Lectotype (designated by Haegens, 2000): Elmer 21509 (lecto US; isolecto A, DS, K, L), Borneo, Sabah, Tawau.

Tree 5.7–25 m high, dbh 5–40 cm; bole sometimes fluted, rarely with buttresses up to 30 cm out, up to 6 cm thick; branchlets densely hairy to sometimes sparsely hairy, Terminalia branching pattern weakly developed. Indumentum of stellate and simple hairs. Bark pale brown to greyish when fresh, brown to brown-red when dry, smooth; inner bark red to brown, c. 2 mm thick. Leaves: petiole 34–130 mm long, usually densely hairy, raised glands rarely present; stipules 5.5–11 by 4–8.5 mm, densely hairy outside and inside, margin ciliate, not hyaline; lamina (ovate to) elliptic (to obovate), 13–36 by 5.4–14.9 cm, l/w ratio 1.7–2.4, papery to somewhat leathery; base rounded to attenuate; apex acuminate, 4–10 mm long; upper surface (sub)glabrous, raised glands absent (to present), usually granulate, dark (grey-)brown to blackish when dry, nervation light (to dark) brown to whitish, (slightly) sunken; lower surface densely hairy, raised glands absent (to present), discoid glands (hairy) in a row between secondary veins, yellowish-brown when dry; secondary veins (5–)8–13 per side, usually closed at margin, not completely parallel; nervation reticulate to weakly scalariform, (yellowish) brown. Staminate inflorescences axillary to just below the leaves, solitary to 3 clustered together, 0.5–16.5 cm long, c. 1 mm thick, branched, densely hairy, many-flowered, flowers scattered along inflorescence; bracts 1 per branchlet, triangular to lanceolate, 1.5–4 mm long, densely hairy outside and inside, margin ciliate, not hyaline; bracteoles usually absent, or 1–few; branchlets cylindrical, 4–11 mm long, densely hairy, 5–16-flowered. Staminate flowers 1.3–2.5 mm diam., lime green to whitish yellow to pale green; pedicel 0.5–2.1 mm long, upper part 0.1–0.9 mm long, densely hairy; sepals 4 or 5, ovate to obovate, 0.8–1.3 by 0.3–0.7 mm, apex slightly recurved to straight, densely hairy outside and inside; staminodes 4 or 5, < 1 mm long; stamens 4 or 5, 0.1–0.6 mm long, glabrous; ﬁlaments 0.15–0.5 mm long, straight; anthers 0.1–0.3 by 0.2–0.25 by c. 0.1 mm; disc absent; pistillode flat, 0.1–0.3 mm high, densely hairy. Pistillate inflorescences ramiflorous to cauline; sepals persistent; stigmas caducous. Pistillate flowers: sepals 5, yellow; ovary 3-locular. Infructescences 4–16 cm long, 1–2.5 mm thick. Fruits pear-shaped to (sub)globose, 5- or 6-seeded, fleshy capsules, 25–34 by 29–35 by 29–34 mm, c. 4 cm diam. when fresh, usually not dehiscent; raised glands present to absent, ruminate when dry, densely hairy to sparsely hairy outside, subglabrous inside, brown to yellow ochre; pericarp 2–5 mm thick; column 19–24 mm long, straight, persistent; pedicel 11–14 mm long, upper part 2.5–4 mm long. Seeds globose to ellipsoid, laterally flattened, 13–17.5 by 8.3–14.3 by 5.5–8 mm; arillode yellowish to orange; testa brown; cotyledons 10–12.2 by 9.5–11.2 by 0.1–2 mm; radicle 0–1.5 mm long.

Distribution — Peninsular Malaysia, Sumatra, Borneo.

Habitat & Ecology — Primary, rarely secondary, rain forest. Soil: yellow sand, clay. Altitude: sea level up to 660(–1650) m. Flowering: January to March, May, September to December; fruiting: March to December.

Uses — Fruit edible, sweet.

Vernacular names — Sumatra: Sebenggang, sibongang. Borneo: Puak (Dusun); tampui (Kedayan); pugi (Murut); silu buang (Sebob); kunkurad, monyak buruk, paih, tampoi.

Note — I have identiﬁed Elmer 21306 (A, DS, K, L, US), one of the syntypes of B. elmeri as B. nanihua. The remaining syntype, Elmer 21509, is chosen as lectotype of B. elmeri.

28. Baccaurea purpurea Haegens

Baccaurea purpurea Haegens, Blumea Suppl. (2000) 164, Fig 3.20, Map 3. — Type: NGF (Henty & Lelean) 41916 (holo L; iso A), New Guinea, Central Province, Abau Subdistrict, Cape Rodney.

Tree 6–8 m high, dbh 6.3–10 cm, buttresses absent; branchlets glabrous, grey to blackish when dry, Terminalia branching pattern well-developed to weak. Indumentum of simple hairs. Bark grey to brown when fresh, corky; inner bark cream to light brown; sapwood cream. Leaves: petiole 17–65 mm long, (sub)glabrous, apically and basally pulvinate, light to black-brown when dry, raised glands present; stipules 5–9 by 1.5–2 mm, glabrous outside and inside, midrib subglabrous to sparsely hairy outside, margin (slightly) ciliate, often hyaline; lamina elliptic (to ovate), 7.4–17.5 by 3.2–7 cm, l/w ratio 2.2–3.2, papery to slightly leathery; base cuneate to rounded; apex (rounded to) obtuse to acuminate, up to 15 mm long, dark green when fresh, (grey) green to dark brown when dry; upper surface glabrous, usually not granulate; lower surface (sub)glabrous, raised glands usually present, discoid glands sometimes present, dark green when fresh, greenish to brown when dry; secondary veins 6–11 per side, sometimes closed at margin; nervation reticulate. Staminate inflorescences cauline, many clustered together, 2–24 cm long, up to 1 mm thick, branches minute, subglabrous to sparsely hairy, many-flowered, flowers scattered along inflorescence; bracts 3 per branchlet, 1 large, 0.2–1 mm long, subglabrous outside, glabrous inside, margin ciliate, hyaline; bracteoles absent. Staminate flowers 3 per branchlet, c. 1.5 mm diam., (purplish) pink; pedicel 1–2 mm long, upper part 0.2–0.5 mm long, sparsely hairy to densely hairy; sepals 4 (or 5), ovate to triangular, 0.8–1.5 by 0.5–1 mm, sparsely to densely hairy outside, densely hairy inside, apex straight; staminodes 5 or 6; stamens 5, 0.4–0.6 mm long, glabrous; ﬁlaments 0.2–0.4 mm long, straight; anthers c. 0.2 by 0.25 by 0.1–0.2 mm; disc absent; pistillode globose, c. 0.3 mm long, densely hairy, solid. Pistillate inflorescences probably cauline to ramiflorous. Pistillate flowers: sepals caducous; ovary 3-locular; stigmas persistent. Fruits globose, 3-seeded, fleshy capsules, 16–19 by 14–19 by 14–19 mm, loculicidally dehiscent, subglabrous to sparsely hairy outside, glabrous inside, few raised glands present, ochre; pericarp 1.5–2 mm thick, slightly ruminate when dry, column 13–14 mm long, straight pedicel 7–9 mm long. Seeds globose, laterally flattened, 9.3–10.3 by 9.2–10.3 by 3.5–5 mm; cotyledons c. 5.5 by 7–7.7 by 1 mm; radicle c. 1 mm long; endosperm c. 1 mm thick.

Distribution — Papua New Guinea: Prov.: Central, Northern, Milne Bay.

Habitat & Ecology — Soil: thin soil over limestone. Altitude: 240–400 m. Flowering: February, November; fruiting: March.

Note — The species is named for its purple (to pink) colour of the male flowers.

29. Baccaurea pyriformis Gage

Baccaurea pyriformis Gage, Rec. Bot. Surv. India 9 (1922) 233; Ridl., Fl. Malay Penins. 3 (1924) 249; S.Moore, J. Bot. Br. 63 (1925) 98; Corner, Wayside Trees Mal. 1 (1940) 241; Whitmore, Tree Fl. Malaya 2 (1973) 67; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 53; Kew Bull. 36 (1981) 265; Chakrab. & Gang., J. Econ. Taxon. Bot. 18 (1994) 426; Haegens, Blumea Suppl. (2000) 166, Fig. 3.21, Map 3.32. — Lectotype (designated by Chakrab. & Gang., 1994): Goodenough 1992 (CAL, n.v), Malakka, Ayer Panas (see note).

Baccaurea platyphylla Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 67; Whitmore, Tree Fl. Malaya 2 (1973) 67; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 53. — Lectotype (designated by Haegens, 2000): Ridley 6491 (lecto K), Singapore, Krangi and Bot. Garden Jungle (both localities are mentioned on the specimen label).

Baccaurea platyphylloïdes Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 68. — Type: Ridley 6492 (holo K), Singapore, Tanglin.

Baccaurea pubera (Miq.) Müll.Arg. var. tomentella Miq., Fl. Ned. Ind., Eerste bijv. (1861) 442; Müll.Arg. in DC., Prodr. 15, 2 (1866) 458. — Type: Teijsmann s.n. (n.v.), Indonesia, Sumatra, Bangka.

Tree 7–30 m high, dbh 8–50 cm, buttresses small if present; branchlets velutinous when young, Terminalia branching pattern weakly developed. Indumentum of stellate and simple hairs. Bark brown to grey-brown to red-brown when fresh, brown when dry, 2–5 mm thick, smooth to rough, soft, flaky; inner bark brown to red-brown or pink, c. 2 mm thick. Heartwood red-brown. Leaves: petiole 14–72 mm long, densely hairy, apically and often basally slightly pulvinate, brown hairy when fresh, raised glands absent; stipules lanceolate to elliptic to triangular, 2–8 by 1.5–2 mm, velutinous outside, glabrous to sparsely hairy (to velutinous) inside, margin ciliate, not hyaline; lamina elliptic to obovate, 6.8–18 by 2.7–8.4 cm, l/w ratio 1.9–3.8, papery to leathery; base (rounded to) acute to cuneate; marginal glands visible as identiﬁcation marks; apex acuminate to cuspidate, (0–)1–17 mm long; upper surface glabrous, not granulate, dark to yellowish green when fresh, dark brown when dry, nervation slightly sunken; lower surface (subglabrous to) densely hairy, midrib and veins (sparsely hairy to) velutinous, raised glands absent, discoid glands present, dark to yellowish green when fresh, light brown when dry; secondary veins 6–9 per side, closed at margin; nervation reticulate. Staminate inflorescences (axillary to) just below the leaves to ramiflorous, solitary to few clustered together, 1–7.5 cm long, up to 1.5 mm thick, velutinous, many-flowered, flowers scattered along inflorescence; bracts absent or 1 per branchlet, 0.3–1.5 mm long, densely hairy outside, glabrous inside, margin ciliate, hyaline or not; bracteoles, 1–1.5 mm long, c. 1 per branchlet, velutinous; branchlets cylindrical to spatulate, 0.4–4 mm long, densely hairy, 3–12-flowered. Staminate flowers 1–2 mm diam., yellowish to pale brown; pedicel 0–0.8 mm long; sepals 4 or 5, ovate, 0.4–0.9 by 0.3–0.8 mm, apex recurved, densely hairy outside and inside, yellowish to pale brown; stamens 4 or 5, 0.3–0.6 mm long, glabrous; ﬁlaments 0.2–0.5 mm long, geniculate to straight; anthers c. 0.1 by 0.1–0.2 by 0.1 mm; disc absent; pistillode cylindrical, 0.4–0.5 mm high, velutinous, solid. Pistillate inflorescences ramiflorous (to axillary), solitary, 1.5–3 cm long, c. 1.2 mm thick, velutinous, c. 10-flowered; pedicel c. 1 mm long, abscission zone not clear, velutinous; bracts 3 per branchlet, velutinous outside, sparsely hairy inside, margin ciliate. Pistillate flowers 2.5–3.5 mm diam., dirty brown; sepals 4 or 5, ovate, 1.3–2 by 1–1.4 mm, velutinous outside and inside, persistent; ovary cylindrical, 2–2.5 by 1.5–2 mm, (2- or) 3-locular, velutinous; style c. 0.2 by c. 1 mm, velutinous; stigmas c. 1.3 mm long, cleft for upper c. 70%, persistent; lobes c. 1 by 0.5 mm, glabrous above, densely hairy below, protuberances present above. Infructescences 1.5–9.8 cm long, 1.5–3.5 mm thick. Fruits pear-shaped (to globose), (2- or) 3-seeded berries, c. 38 by 34 by 34 mm when fresh, 18–24 by 14–22 by 14–22 mm when dry, rarely irregularly splitting, raised glands absent, not ruminate, densely hairy outside, subglabrous to densely hairy inside, (yellowish to orange-)brown; pericarp 1.5–2.5 mm thick; column 13–19 mm long, straight; pedicel 3–9 mm long, upper part 0.5–4.5 mm long. Seeds ellipsoid, laterally flattened, 8–17.5 by 5–12 by 2–7 mm; testa red; arillode cream to orange; cotyledons 9–12 by 6.2–11.5 by 1–2 mm (endosperm included); radicle up to 1 mm long.

Distribution — Peninsular Malaysia, Sumatra, Borneo.

Habitat & Ecology — Primary rain forest. Soil: sandstone, sandy loam. Altitude: sea level up to 1000 m. Flowering: March to October; fruiting: January, February, June to November.

Uses — Arillode edible.

Vernacular names — Simeuluë Island: Bolawah iteung falah. Borneo: Papar pingan (Bussup); tampoi hutan (Dusun Ramu); tampoi (Iban); tampoi merah (Malay); bukut.

Note — The following former syntypes have been studied: Derry 66; Griffith 4901; Maingay 1361; Ridley 6491, 6492, 6669; SF 28, 2766, 2949.

30. Baccaurea racemosa (Reinw. ex Blume) Müll.Arg.

Baccaurea racemosa (Reinw. ex Blume) Müll.Arg. in DC., Prodr. 15, 2 (1866) 461; Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 281; J.J.Sm., Meded. Dept. Landb. Ned.-Indië 10 (1910) 249; Koord., Exkurs. Fl. Java 2 (1912) 481; K.Heyne, Nutt. Pl. Ned. Ind. 3 (1917) 75; Merr., J. Straits Branch Roy. Asiat. Soc. (1921) 331; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 51; Merr., Univ. Calif. Publ. Bot. 15 (1929) 150; Backer & Bakh.f., Fl. Java 1 (1964) 454; Soejarto, Bot. Mus. Leafl. (1965) 74; Whitmore, Tree Fl. Malaya 2 (1973) 64; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 53; Kew Bull. 36 (1981) 266; Lemmens et al., Prosea 3 (1991) 16; Haegens, Blumea Suppl. (2000) 169, Fig. 3.22, Map 3.33; Haegens & Welzen in Welzen et al., Thai For. Bull. 28 (2000) 73; in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 117. — Coccomelia racemosa Reinw. ex Blume, Catal. Gew. Buitenzorg (1823) 110; Reinw., Flora 8, 1 (1825) 103; Syll. Ratisb. 2 (1828) 5. — Pierandia racemosa (Reinw. ex Blume) Blume, Bijdr. (1825) 579. — Pierardia racemosa (Reinw. ex Blume) Miq., Fl. Ned. Ind. 1. 2 (1859) 358; Fl. Ned. Ind., Eerste bijv. (1861) 442; Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 281. — Lectotype (designated by Haegens, 2000): Blume s.n. (lecto L, sheet 903.154-627, barcode L 0059160), Java.

Baccaurea wallichii Hook.f., Fl. Brit. India 5 (1887) 375; K.Heyne, Nutt. Pl. Ned. Ind. 3 (1917) 95; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 53; Ridl., Fl. Malay Penins. 3 (1924) 245; Soejarto, Bot. Mus. Leafl. (1965) 99. — Lectotype (designated by Haegens, 2000): King’s Collector 7634 (lecto G), Peninsular Malaysia, Perak.

Baccaurea bhaswatii Chakrab. & M.Gangop., J. Econ. Taxon. Bot. 18 (1994) 420. — Lectotype (designated by Chakrab. & Gang., 1994): Forbes 1653 (lecto CAL; isolecto L), Sumatra.

Treelet to tree 3–15(–20) m high, dbh 5–24 cm, buttresses absent; branchlets glabrous to sparsely hairy; young shoots reddish-black to reddish-brown when dry, Terminalia branching pattern well-developed. Indumentum of simple hairs and rarely stellate hairs. Bark whitish to grey to pale brown to fawn when fresh, greyish to brown when dry, up to 3 mm thick, smooth, papery, flaky, soft; inner bark white to yellow to creamy to ochre to brown, 1–2 mm thick, soft. Heartwood yellowish-brown to yellowish-ochre to brown, hard. Leaves: petiole 12–77 mm long, glabrous to sparsely hairy, red to brown when dry; stipules elliptic to triangular, 3–7.5 by 1–1.5 mm, glabrous to sparsely hairy, densely hairy when young outside, glabrous inside, margin ciliate, usually hyaline; lamina (ovate to) elliptic to obovate, 5.8–22 by 2.3–18.8 cm, l/w ratio 2.2–4.6, papery; base cuneate; apex acuminate to cuspidate, up to 25 mm long; upper surface glabrous, sometimes granulate, raised glands usually present, dark green when fresh, (white to green)-brown when dry; lower surface glabrous, rarely densely hairy at veins, discoid glands rarely present, light green when fresh, brown when dry, yellowish green to pink when young: nervation reddish to brown to black when dry; secondary veins 4–10 per side, almost closed at margin; nervation reticulate to weakly scalariform. Staminate inflorescences cauline to just below the leaves, solitary to 5 clustered together, 3–10 cm long, up to 1.3 mm thick, densely hairy, many-flowered, flowers scattered along inflorescence; bracts 3 per branchlet, of which two very small, 0.2–1.1 mm long, persistent, glabrous to densely hairy outside, glabrous inside, margin ciliate, rarely hyaline; branchlets triangular, 0.1–2.5 mm long, densely hairy, (2- or) 3-flowered. Staminate flowers 1–3 mm diam., creamy yellow to white; pedicel 0.4–2.5 mm long, upper part 0.4–1.9 mm long, densely hairy; sepals 4 (or 5), ovate to obovate, 0.9–1.8 by 0.7–1.5 mm, apex usually recurved, densely hairy on both sides; stamens 5–7, 0.5–0.8 mm long, glabrous; ﬁlaments 0.4–0.6 mm long, straight; anthers white, c. 0.15 by 0.2–0.25 by 0.1 mm; disc present or absent; pistillode cylindrical, 0.3–0.9 mm long, densely hairy, hollow. Pistillate inflorescences cauline to axillary, solitary (to few clustered together), 3–28 cm long, c. 1 mm thick, densely hairy, 4–many-flowered, reddish-green when fresh, reddish to brown when dry; pedicel 1–2.9(–6.7) mm long, upper part up to 1.9(–2.5) mm long, densely hairy, green to brown to pinkish; bracts 3 per branchlet, persistent, densely hairy outside, glabrous inside, margin ciliate. Pistillate flowers 3–9.2 mm diam., greenish yellow when fresh; sepals 4 (or 5), obovate to lanceolate, 2–6 by 1–2.7 mm, densely hairy on both sides, indumentum of silvery hairs when dry, caducous; ovary subglobose, 1.5–1.9 by 1.4–1.7 mm, (2- or) 3- (or 4-)locular, velutinous; style 0.2–0.4 by c. 1 mm, velutinous; stigmas c. 0.3 mm long, not lobed, persistent. Fruits (sub)globose, 1–3-seeded, berries or fleshy capsules, 14–30 by 16–25 by 16–25 mm, outer pericarp loculicidally dehiscing, inner pericarp not dehiscent, glabrous to sparsely hairy outside, glabrous inside, raised glands present, yellow to orange (to pink to red); pericarp 0.8–4.5 mm thick; column 12–15 mm long, caducous, straight; pedicel reddish when dry, 3–9 mm long, upper part 1–4.5 mm long. Seeds obovoid to ellipsoid, laterally flattened, 9–11.5 by 6.5–9.2 by 2.5–6 mm; testa reddish-purple to blue to black; arillode blue to purple to violet; cotyledons 5–7 by 6–7 by 0.1–1 mm, radicle 0–0.3 mm long.

Distribution — Thailand, Peninsular Malaysia, Sumatra, Java, Bali, Borneo, Sulawesi and (Halmahera).

Habitat & Ecology — Primary and secondary rain forest, riverine forest, or fresh water swamp forest. Soil: granite to yellow or red sandy clay. Altitude: 50–1550 m. Flowering: March and May to October; fruiting: throughout the year.

Uses — Arillode edible, sour to sweet.

Vernacular names — Peninsular Malaysia: Asam tambun, rambi, tamut, tampoi. Sumatra: Roesip, kisip. Java: Menteng. Borneo: Kokonau (Dusun); engkuni, enkunik, kayu masam, longkuno, moho liox.

31. Baccaurea ramiflora Lour.

Baccaurea ramiflora Lour., Fl. Cochinch. (1790) 661; Müll.Arg. in DC., Prodr. 15, 2 (1866) 458; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 71; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1927) 551; Merr., Trans. Amer. Philos. Soc. 24, 2 (1935) 232; Airy Shaw, Kew Bull. 26 (1971) 221; Whitmore, Tree Fl. Malaya 2 (1973) 65; Airy Shaw, Kew Bull. 36 (1981) 266; Haegens, Blumea Suppl. (2000) 172, Map 3.34, Photo 9; Haegens & Welzen in Welzen et al., Thai For. Bull. 28 (2000) 73; in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 117, plate IV: 3. — Lectotype (designated by Haegens, 2000): Loureiro s.n. (lecto BM, sheet 31252), Hortis Cochinchinae (see note 1).

Baccaurea cauliflora Lour., Fl. Cochinch. (1790) 661; Müll.Arg. in DC., Prodr. 15, 2 (1866) 458; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 70; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1927) 551; Merr., Trans. Amer. Philos. Soc. 24 (1935) 233. — Neotype (designated by Haegens, 2000): Fleury 32043 (neo P), Vietnam, Cochinchina, Bien Hoa Province, Trañg Bône Forest Reserve (see note 2).

Pierardia sapida Roxb., Fl. Ind. 2 (1832) 254; Hortus Bengalensis (1814) 28. — Baccaurea sapida (Roxb.) Müll.Arg. in DC., Prodr. 15, 2 (1866) 459; Bedd., Fl. Sylv. S. India (1869) 280; Kurz, Forest Fl. Burma 2 (1877) 356; Hook.f., Fl. Brit. India 5 (1887) 371; Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 281; Williams, Bull. Herb. Boissier. 2 (1905) 30; Brandis, Indian Trees (1906) 562; Craib, Aberdeen Univ. Stud. 57 (1911) 461; Koord., Syst. Verz. (1914) 67; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 52; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1927) 548; Merr., Trans. Amer. Philos. Soc. 24 (1935) 233; Corner, Gard. Bull. Straits Settlem. 10 (1939) 290; Wayside Trees Mal. 1 (1940) 241. — Lectotype (designated by Haegens, 2000): Wallich 8072 (lecto K), Burma?

Baccaurea flaccida Müll.Arg. in DC., Prodr. 15, 2 (1866) 459; Hook.f., Fl. Brit. India 5 (1887) 370; Brandis, Indian Trees (1906) 563; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 50. — Lectotype (designated by Haegens, 2000): Wallich 8074 (lecto K), Burma.

Baccaurea propinqua Müll.Arg. in DC., Prodr. 15, 2 (1866) 463. — Lectotype (designated by Haegens, 2000): Wallich 8071 (lecto G) India Orientali, Silhet.

Baccaurea wrayi King ex Hook.f., Fl. Brit. India 5 (1887) 374; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 53; Ridl., Fl. Malay Penins. 3 (1924) 244; S.Moore, J. Bot. Br. 63 (1925) 98; Airy Shaw, Kew Bull. 14 (1960) 354; Kew Bull. 26 (1971) 221. — Lectotype (designated by Haegens, 2000): King’s Collector s.n. (lecto K), Peninsular Malaysia, Perak.

Baccaurea oxycarpa Gagnep., Bull. Soc. Bot. France 23 (1923) 431; in Lecomte, Fl. Indo-Chine 5 (1927) 549, f. 69; Airy Shaw, Kew Bull. 14 (1960) 354; Kew Bull. 26 (1971) 221; Vu Van Dung, Viet-nam Forest Trees (1996) 191. — Lectotype (designated by Haegens, 2000): Pierre 614 (lecto K; isolecto P), Cambodia, Mt Knang-krepeuh.

Gatnaia annamica Gagnep., Bull. Soc. Bot. France 24 (1924) 870; in Lecomte, Fl. Indo-Chine 5 (1927) 540; Pax & K.Hoffm. in Engl. & Harms, Nat. Pflanzenfam. ed. 2, 19c (1931) 50. — Lectotype (designated by Haegens, 2000): Eberhardt 3042 (lecto P; isolecto K, L), Annam, Thua-thien Prov.

Baccaurea sylvestris auct. non Lour.: Lour., Fl. Cochinch. (1790) 662, p.p., inflorescences; Müll.Arg. in DC., Prodr. 15, 2 (1866) 457; Pellegr., Fl. Indo-Chine (1911) 766; A. Chev., Cat. Pl. Jard. Bot. Saigon (1919) 64; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 61; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1927) 551; Merr., Trans. Amer. Philos. Soc. 24, 2 (1935) 233; Mabb. et al., Fl. Males. ser. 1, 12 (1995) 316; Vu Van Dung, Vietnam Forest Trees (1996) 192 (see note 3).

Tree 5–13 m high, dbh 5–33 cm, buttresses absent; branchlets glabrous to sparsely hairy; young shoots greenish white when fresh, brown to greenish white when dry, Terminalia branching pattern well-developed. Indumentum of simple hairs. Bark greyish-brown to grey when fresh, 1–2 mm thick, smooth; inner bark white. Leaves: petiole 10–63 mm long, glabrous to sparsely hairy, raised glands present; stipules 2.5–6 by 1–2.5 mm, caducous (to late caducous), glabrous to sparsely hairy outside, midrib densely hairy to velutinous, glabrous inside, margin ciliate, usually hyaline; lamina elliptic to obovate, 7.1–25.5 by 2.9–8.8 cm, l/w ratio 1.8–4, papery; base attenuate to cuneate; apex acute to cuspidate, 2–12 mm long; upper surface: glabrous, raised glands present, not granulate, glossy dark green when fresh, greenish brown to greyish brown when dry; lower surface glabrous to sparsely hairy, raised glands present, discoid glands absent, green when fresh, greenish brown when dry; secondary veins 4–9 per side, closed at margin or almost so; nervation reticulate to weakly scalariform. Staminate inflorescences axillary to cauline, solitary to many clustered together, 3–8.5(–15.5) cm long, up to 1 mm thick, densely hairy, many-flowered, flowers clustered at the tip of inflorescence, greenish; bracts conspicuous, 1 (or 3) per branchlet, (1.5–) 3–4 mm long, persistent, sparsely to densely hairy outside, glabrous inside, but densely hairy at base, margin ciliate, sometimes hyaline; bracteoles absent; branchlets cylindrical, absent to 1.8 mm long, densely hairy, 3-flowered. Staminate flowers 1.1–4 mm diam., yellow; pedicel 0.8–2.6 mm long, upper part 0.7–2.4 mm long, densely hairy; sepals 4 or 5, different in shape, 1–2.4 by 0.5–1.4 mm, sometimes apically recurved, densely hairy outside and inside; stamens 5–8, 0.7–1.1 mm long, glabrous; ﬁlaments 0.5–0.9 mm long, straight; anthers 0.2–0.3 by 0.2–0.4 by c. 0.2 mm, yellow; pistillode present, obtriangular, 0.9–1.1 mm high, densely hairy, solid. Pistillate inflorescences just below the leaves to cauline, solitary to many clustered together, c. 10 cm long, c. 1 mm thick, densely hairy, 10–15-flowered, red; pedicel 1.5–3 mm long, upper part 0.1–1 mm long, densely hairy; bracts 1 per branchlet, persistent (to caducous), sparsely hairy outside, densely hairy at midrib, glabrous inside, margin ciliate. Pistillate flowers 3–8 mm diam., yellow; sepals 4 or 5, elliptic, 3.5–5 by 1–2 mm, densely hairy outside and inside, caducous; ovary globose, 2–2.3 by 2–2.3 mm, (2- or) 3- (or 4-)locular, woolly, green; style absent; stigmas c. 0.5 mm long, not lobed, glabrous, cream, caducous (to persistent). Infructescences 12–20 cm long, 1–2.5 mm thick. Fruits globose to ovoid to pear-shaped, 2–4-seeded berries, 19–37 by 14–28 by 14–28 mm, raised glands absent, glabrous to sparsely hairy outside, glabrous inside, red to orange to pink to purplish outside, cream inside; pericarp 1–3 mm thick; column fragile; pedicel 1–10(–15) mm long, upper part 0.5–3 mm long. Seeds globose to ellipsoid, but laterally flattened, 9–15 by 9–11 by 1.5–6 mm; arillode white; testa purplish red; cotyledons 5.5–9 by 5.2–8.5 by c. 0.1 mm; radicle 0.5–1.8 mm long; endosperm 1.5–7 mm thick.

Distribution — India (Assam), Burma, China (Yunnan, Hainan), Vietnam, Laos, Thailand, Andaman and Nicobar Islands, Peninsular Malaysia.

Habitat & Ecology — Primary rain forest and cultivated. Soil: sand, granite. Altitude 50–1700 m. Flowering: December to June, September; fruiting: throughout the year.

Uses — Arillode edible, sweet to sour.

Vernacular names — Burma: Kanaso. Thailand: Mahphie. Peninsular Malaysia: Tampoi.

Notes — 1. The front side of the type sheet (BM: 31252) of B. ramiflora refers to page 662 of Fl. Cochinch. and B. sylvestris. This is certainly wrong, because the specimen does not correspond with the description of B. sylvestris. The back side of this sheet refers to page 661 of Fl. Cochinch., where B. ramiflora is described.

2. The type of B. cauliflora is probably lost. There is no original illustration. It is therefore not possible to lectotypify this name. From the description it is however obvious that this name is conspeciﬁc with B. ramiflora. It is here therefore neotypiﬁed. Fleury 32043 is chosen as neotype, because it is a collection from Cochinchine, and it is a fruiting specimen like the original specimen described by Loureiro.

3. The type sheet of B. sylvestris is a mixture. The leaves on this sheet are Lancium domesticum Corrêa, see Mabberley et al. (1995). The description of Loureiro is also congruent with Lancium domesticum. The fruits, for example, contain 5 locules, and each locule has one ovule only (both features not present in Baccaurea). The inflorescences on the type sheet, on the other hand, belongs to B. ramiflora. The fact that the type specimen is a mixture has probably caused the confusion about the taxonomical status of B. sylvestris.

32. Baccaurea reticulata Hook.f.

Baccaurea reticulata Hook.f., Fl. Brit. India 5 (1887) 373; Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 281; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 65; Ridl., Fl. Malay Penins. 3 (1924) 246; Corner, Wayside Trees Mal. 1 (1940) 241; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 36; Whitmore, Tree Fl. Malaya 2 (1973) 67; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 54; Kew Bull. 36 (1981) 266; Chakrab. & Gang., J. Econ. Taxon. Bot. 18 (1994) 424; Haegens, Blumea Suppl. (2000) 174, Fig. 3.23; Map 3.35. — Lectotype (designated by Haegens, 2000): Scortechini s.n. (lecto L, sheet 903.154-630, barcode L 0039846), Peninsular Malaysia, Perak, Goping.

Tree or treelet 6–28 m high, dbh 21–60 cm, bole sometimes fluted to up to 2 m high; branchlets subglabrous, densely hairy when young, Terminalia branching pattern weakly developed. Indumentum of stellate and simple hairs. Bark red to red-brown to orange-brown when fresh, brown when dry, thin, rugose, ﬁbrous to flaky; inner bark red to pink to light brown, 2–5 mm thick. Heartwood red-brown to yellow-brown. Leaves: petiole 18–87 mm long, subglabrous to velutinous with short stellate hairs and long simple hairs, transverse cracks usually, apically and slightly basally pulvinate, raised glands sometimes present; stipules 4–7.5 by 1.5–3.5 mm, velutinous outside and inside, margin ciliate, not hyaline; lamina (ovate to) elliptic to obovate, 5.4–14.5 by 2.8–6.6 cm, l/w ratio 1.6–2.9, papery to slightly leathery; base acute to cuneate (to attenuate); marginal glands sometimes visible as small indentations; apex (obtuse to) acuminate to cuspidate, up to 16 mm long; upper surface glabrous, raised glands often present, sometimes granulate, grey-green to green-white to brown-white, nervation whitish to grey-green, slightly sunken; lower surface subglabrous, densely hairy at midrib and veins, raised glands often present, discoid glands if present in a row between secondary veins, light (yellowish) brown when dry; secondary veins 4–8 per side, closed at margin; nervation (weakly) scalariform, (dark) brown. Staminate inflorescences axillary (to ramiflorous), solitary to 3 clustered together, 1–7 cm long, c. 0.1 mm thick, branched, velutinous, many-flowered, flowers scattered along inflorescence; bracts 1 per branchlet, 1–5 mm long inserted, densely hairy outside and inside, margin ciliate, not to slightly hyaline; bracteoles 0.5–2.5 mm long, 1–3 (or 4) per branchlet; branchlets cylindrical, 1–4 mm long, densely hairy, (3–)9–15-flowered. Staminate flowers 0.8–2.5 mm diam., pale yellow to greenish white to white; pedicel 0.6–1.5 mm long, upper part 0.3–1.2 mm long, densely hairy; sepals 4 (or 5), ovate to elliptic, 0.7–1.8 by 0.2–0.9 mm, apex recurved, densely hairy outside and probably also inside; stamens 4 (or 5), 0.2–0.3 mm long, glabrous; ﬁlaments up to 0.2 mm long, straight; anthers 0.1–0.2 by 0.15–0.2 by c. 0.1 mm; disc absent; pistillode absent to small, up to 0.15 mm high, velutinous. Pistillate inflorescences ramiflorous to just below the leaves. Pistillate flowers: ovary 3- (or 4-)locular. Infructescences 2–5 cm long, 2–3 mm thick. Fruits globose, 3–6-seeded, fleshy capsules, 25–38 by 29–51 by 29–51 mm, loculicidally dehiscent, raised glands present, subglabrous to tomentose outside, subglabrous inside, septa sparsely hairy, green to dirty orange to brown; pericarp 3–6.5 mm thick; column 19–25 mm long, straight; sepals usually persistent; stigmas usually caducous; pedicel 12–21 mm long, upper part 8–15 mm long. Seeds globose to ellipsoid, laterally flattened, 10–13.5 by 8–14.5 by 3–4 mm; arillode whitish yellow, translucent; cotyledons 8–11 by 6.5–12 by up to 0.1 mm; radicle up to 0.1 mm long.

Distribution — Peninsular Malaysia, Sumatra, Borneo.

Habitat & Ecology — Secondary rain forest. Soil: yellow sandy loam. Altitude: sea level up to 330 m. Flowering: January, March, April, August, September; fruiting: June to September, November.

Uses — Arillode edible, sweet.

Vernacular names — Peninsular Malaysia: Tampoi, tampoy, tampui. Sumatra: Kaloe merah, tampoei simba.

33. Baccaurea sarawakensis Pax & K.Hoffm.

Baccaurea sarawakensis Pax & K.Hoffm. in Engl., Pflanzenr. VI.147.xv (1922) 53; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 54; Haegens, Blumea Suppl. (2000) 177, Map 3.36, Photo 10, 11. — Lectotype (designated by Haegens, 2000): Hose 43 (lecto K; isolecto L), Borneo, Sarawak, Baram.

Tree 3–18(–21) m high, dbh 4–21 cm, buttresses absent; branchlets glabrous; young shoots greyish-white when dry, slightly thickened at tip, Terminalia branching pattern well-developed. Indumentum of simple hairs. Bark whitish to green to grey to brown when fresh, greyish-white to brown when dry, c. 1–2.5 mm thick; inner bark yellowish to pale brown or pale red, up to 2 mm thick, soft. Leaves: petiole brown when dry, 10–56 mm long, sparsely hairy, raised glands present; stipules caducous to late caducous, lanceolate, 2.5–9 by 1–2.8 mm, sparsely to densely hairy outside, glabrous inside, margin ciliate, sometimes hyaline; lamina obovate to elliptic, 7–20 by 3–8 cm, l/w ratio 2.3–3, ﬁrm papery to leathery; base cuneate to acute; apex (rounded to) acuminate, up to 17 mm long; upper surface glabrous, sometimes granulate, glossy green when fresh, whitish brown when dry; lower surface glabrous, rarely sparsely hairy at midrib and veins, raised glands usually present, discoid glands sparse or absent, shiny pale green when fresh, whitish green to brown when dry; secondary veins 5–8 per side, closed at margin; nervation reticulate to weakly scalariform, (whitish) brown when dry. Staminate inflorescences axillary to just below the leaves, solitary to 5 clustered together, 3–12 cm long, less than 1 mm thick, densely hairy, more than 60-flowered, flowers scattered along inflorescence, bright yellow; branchlets 0–0.2 mm long; bracts 3 per branchlet, 0.4–1.5 mm long, glabrous to densely hairy outside, (sub)glabrous inside, margin ciliate. Staminate flowers 3 per branchlet, 1–1.4 mm diam.; pedicel 1.9–4.5 mm long, upper part 1–2.8 mm long, densely hairy; sepals ovate to obovate, 4 or 5, apically recurved, 0.8–1.2 by 0.4–0.8 mm, glabrous to densely hairy inside and outside; stamens 4 or 5, 0.15–0.2 mm long, glabrous; ﬁlaments c. 0.1 mm long, straight; anthers c. 0.2 by 0.2–0.3 by 0.1 mm; disc present, 0.1–0.15 mm thick; pistillode globose, 0.25–0.3 mm high, densely hairy, solid. Pistillate inflorescences axillary to ramiflorous, solitary to few clustered together, 5–10 cm long, c. 1 mm thick, densely hairy, 9–16-flowered; bracts 3, usually persistent, densely hairy outside, glabrous inside, margin ciliate; pedicel 2–4.1 mm long, upper part c. 0.2 mm long, densely hairy. Pistillate flowers 2–9 mm diam.; sepals 4 or 5, lanceolate, 1.5–9 mm by 1–2.6 mm, glabrous to densely hairy on both sides, caducous; ovary (sub)globose, 1.5–3.5 by 1.5–3.2 mm, 3-locular, woolly; style (0–)0.5–1 mm by 1–2 mm, glabrous to densely hairy; stigmas c. 1 mm long, slightly lobed apically, caducous; lobes above glabrous with protuberances, below densely hairy. Fruits subglobose to triangular, 0–3-seeded, fleshy capsules, 18–28(–30) by 16–30 by 13–30 mm, dehiscing loculicidally, outside sparsely to densely hairy, inside glabrous, raised glands present, red to yellowish red-pink; pericarp 0.5–3.5 mm thick, column 13–16 mm long, caducous; pedicel 2–11 mm long, upper part 1–2 mm long. Seeds obovoid, laterally flattened, 10–14 by 7–11.5 by 1–5.7 mm; arillode blue to pinkish. Seeds coat blue to purple; cotyledons 6–8 by c. 6 by 0.1 mm.

Distribution — Borneo.

Habitat & Ecology — Primary rain forest. Altitude: up to 800(–1600) m. Soil: loam. Flowering: March, April, June to September; fruiting: January, February, May to December.

Vernacular names — Borneo: Ankuni (Brunei, Dayak); engkuni, gune (Iban); kunau-kunau (Malay); mata kunau (Brunei, Dusun); Ng’kokonau (Dusun Rongos); asam kuning, enkunyit, kayu masam, peladuk.

34. Baccaurea simaloerensis Haegens

Baccaurea simaloerensis Haegens, Blumea Suppl. (2000) 179, Fig. 3.24, Map 3. — Type: Achmad 728 (holo L), Sumatra, Simeuluë Island.

Tree; branchlets glabrous to sparsely hairy, densely hairy when young, Terminalia branching pattern weakly developed. Indumentum of simple and usually short stellate hairs. Bark dark brown when dry, thin. Leaves: petiole 12–40 mm long, subglabrous to sparsely hairy, rarely densely hairy, apically and basally pulvinate, blackish to dark reddish brown when dry, raised glands usually absent; stipules 2–4 by 1–1.5 mm, sparsely to densely hairy outside, midrib densely hairy, glabrous inside, margin ciliate, not hyaline; lamina ovate to elliptic, 6.2–18 by 3.4–9.1 cm, l/w ratio 1.5–2.1, papery; base truncate to acute (to attenuate); apex obtuse to acuminate, up to 13 mm long; upper surface glabrous, small raised glands present, usually granulate, dark brown, nervation sunken; lower surface (sub)glabrous, subglabrous to sparsely hairy at midrib, raised glands present, discoid glands usually present, scattered, pale brown when dry; secondary veins 5 or 6 (or 7) per side, open at margin; nervation reticulate. Staminate inflorescences axillary to just below the leaves, solitary, 3–10 cm long, 0.6–1 mm thick, branched, sparsely to densely hairy, many-flowered, flowers scattered along inflorescence; bracts 1 per branchlet, (broadly) ovate, 2.2–3.5 mm long, usually caducous, densely hairy outside, glabrous inside, margin ciliate, not hyaline; bracteoles usually 4, 1.5–2.5 mm long; branchlets cylindrical, 2–3.5 mm long, 6–20-flowered, densely hairy. Staminate flowers 1.5–2 mm diam., caducous before dehiscence; pedicel 0.3–1 mm long, densely hairy; sepals 4 or 5, ovate, 1–1.3 by 0.7–1.2 mm, apically slightly recurved, velutinous outside and inside; stamens 4–7, 0.3–0.4 mm long, glabrous; ﬁlaments 0.1–0.2 mm long, straight; anthers 0.15–0.1 by 0.15–0.2 by c. 0.1 mm; disc absent; pistillode present, cylindrical to obtriangular, c. 0.5 mm high, velutinous, solid. Pistillate inflorescences axillary, solitary, 2–4.2 cm long, 1–2 mm thick, densely hairy, few-flowered; pedicel 2–8.5 mm long, upper part 1–4.5 mm long, densely hairy; bracts probably 3 per branchlet, spatulate, persistent, subglabrous outside, sparsely hairy at base, glabrous inside, margin ciliate. Pistillate flowers 8–14 mm diam.; sepals 5, lanceolate to elliptic, 5–7.5 by 1.2–1.5 mm, densely hairy outside and inside, caducous to persistent; ovary globose, 2–4 by 2–5 mm, (2-? or) 3-locular, velutinous; style c. 0.5 by 1.5 mm, velutinous; stigmas c. 2.8 mm long, cleft for upper 60–70%, persistent; lobes c. 2 by 0.5 mm, glabrous above, protuberances small, below densely hairy, protuberances strong. Fruits globose, usually 2–4-seeded, berries or fleshy capsules, 15.5–18 by 15.8–23.5 by 15.8–23.5 mm, only outer layer loculicidally dehiscent, raised glands absent, sparsely to densely hairy outside, (sub)glabrous inside, septa subglabrous; pericarp 1.5–6.5 mm thick; column 9–12 mm long, straight. Seeds ellipsoid, laterally flattened, 7–8.5 by 3.5–4.5 by 1.5–3.5 mm; cotyledons 2.8–3 by 2–3 by c. 0.1 mm; radicle 0.6–2 mm long; endosperm 0.2–0.5 mm thick.

Distribution — Sumatra: Simeuluë Island.

Habitat & Ecology — Flowering: June, September, November, December; fruiting: February, May.

Vernacular names — Simeuluë Island: Awa-batoe-fatoeh, bolahna itung, bolawah-iteung-delok, bolawah ontur, bolawah palo, bolawah pajo-oeding, bolawah silai, soerai-datan, toetoen bolafah.

Notes — 1. In this species all staminate flowers in herbarium material fall off before anthesis. In B. bracteata, a close relative, a part of the staminate flowers show the same phenomenon. However, despite the fact that the staminate flowers remain closed, pollination takes place because in the fruit bearing specimen seeds are produced. A possible explanation for the successful pollination could be that the breaking off of the unopened flowers is an artifact, and that the flowers do open in the wild. Another explanation could be that fertilisation is performed by thrips, who are able to enter the closed flowers.

2. This species is named for its restricted distribution on Simeuluë Island, off the coast of Sumatra. All studied collections of this species were made by Achmad, and he spelled the island as Simaloer.

35. Baccaurea sumatrana (Miq.) Müll.Arg.

Baccaurea sumatrana (Miq.) Müll.Arg. in DC., Prodr. 15, 2 (1866) 466; Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 281; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 63; S.Moore, J. Bot. Br. 63 (1925) 98; Merr., Univ. Calif. Publ. Bot. 15 (1929) 151; K.Heyne, Nutt. Pl. Ned. Ind. 1, 3rd ed. (1950) 915; Soejarto, Bot. Mus. Leafl. 21 (1965) 71, 99; Meijer, Bot. News Bull. Forest Dept., Sabah (1967) 36; Whitmore, Tree Fl. Malaya 2 (1973) 65; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 55; Kew Bull. 36 (1981) 267; Chakrab. & Rao, J. Econ. Taxon. Bot. 5 (1984) 957; G.L. Webster, Ann. Missouri Bot. Gard. 81 (1994) 51; Haegens, Blumea Suppl. (2000) 182, Fig. 3.25, Map 3.39; Haegens & Welzen in Welzen et al., Thai For. Bull. 28 (2000) 74; in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 118, fig. 24g-i. — Calyptroon sumatranum Miq., Fl. Ned. Ind., Eerste bijv. (1861) 472. — Lectotype (designated by Haegens, 2000): Teijsmann s.n. (lecto U, sheet 35994) Sumatra.

Baccaurea kingii Gage, Rec. Bot. Surv. India 9 (1922) 231; Hook.f., Fl. Brit. India 5 (1887) 373; Ridl., Fl. Malay Penins. 3 (1924) 245; Whitmore, Tree Fl. Malaya 2 (1973) 65; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 54. — Lectotype (designated by Haegens, 2000): Wray 1364 (lecto SING), Malakka, Perak, Tapa.

Baccaurea bivalvis Merr., Univ. Calif. Publ. Bot. 15 (1929) 148; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 56. — Type: Elmer 21510 (holo US; iso DS, GH, K, L), Borneo, Sabah, Tawau.

Tree (4–)15–43 m high, dbh (5–)15–35 cm, buttresses if present up to 1 m high, c. 6 cm thick; branches glabrous, young shoots dark brown to black, Terminalia branching pattern well-developed. Indumentum of simple hairs. Bark greyish-brown to brown to black, 2–16 mm thick, smooth to rather rough, hard, ﬁbrous, peeling off with longitudinal flakes of c. 1 cm long, raised glands present, (exudate watery, red); inner bark white to red brown to pink, 1.5–2 mm thick, ﬁbrous, hard, scaly. Heartwood reddish to dark brown. Leaves: petiole 5–48 mm long, glabrous; stipules 1–7 by c. 1 mm, sparsely hairy on both sides, densely hairy at midrib outside, margin not ciliate; lamina ovate to sometimes obovate, 3–19 by 1.2–7.5 cm, l/w ratio 1.1–2.9(–3.75), papery; base attenuate to cuneate (to rounded); apex obtuse to acuminate, up to 18 mm long; upper surface glabrous, rarely granulate, a few discoid glands rarely present; lower surface glabrous, a few discoid glands present; green to red when dry, red when young; secondary veins, 5–8(–10) per side, sometimes closed at margin; nervation reticulate. Staminate inflorescences axillary to somewhat ramiflorous, solitary (to 3 clustered together), up to 2.3(–5.4) cm long, densely hairy, 30–75-flowered, flowers grouped at upper part of inflorescence; branchlets spatulate, 2–3.8 mm long, densely hairy, with (2 or) 3 (or 4) flowers; bracts 1 per branchlet, c. 1 mm long, glabrous outside. Staminate flowers sessile, 0.9–1.8 mm diam., yellow-green; sepals 3–5, each with a different size and shape, 0.3–1.2 by 0.3–1 mm, glabrous; stamens 4–6, 0.4–1 mm long, glabrous; ﬁlaments 0.4–0.9 mm long; anthers 0.1–0.15 mm; pistillode ligulate, up to 0.5 mm high, densely hairy. Pistillate inflorescences axillary to ramiflorous, solitary (to 4 clustered together), 0.7–6.5(–9) cm long, 1–2 mm thick, densely hairy except at base, 2–20-flowered; pedicel geniculate (90° bent) at the abscission zone, 2.2–2.5 mm long, upper part c. 1.2 mm, densely hairy; bracts absent. Pistillate flowers 1.5–2 mm; sepals 4 or 5, ovate to obovate, each with a different size and shape, 0.5–1.3 by 0.5–1.4 mm, densely hairy on both sides; ovary cylindrical, 1.8–2.5 by 1–1.4 mm, 2-locular, sparsely hairy to velutinous; style absent or up to 0.1 mm long; stigmas not lobed to apically lobed up to the base, lobes up to 0.5 mm long, glabrous. Fruits ellipsoid, laterally flattened, usually 1-seeded, fleshy capsules, 9–17 by 4–10 mm, dehiscing loculicidally, outside subglabrous, inside glabrous, septa rarely sparsely hairy, yellow to orange to reddish; pedicel straight, 2–8 mm long, upper part 0.5–2 mm; pericarp c. 1 mm thick; column often persistent, 6–12 mm long, curved, flattened; sepals persistent, small. Seeds ellipsoid, laterally flattened, 6–9 by 4–7.5 by 2–4 mm; arillode orange; testa red; cotyledons 3.5–6 by 4–6.5 mm, thin; radicle up to 0.8 mm long.

Distribution — Thailand, Peninsular Malaysia, Sumatra, Borneo and (Java).

Habitat & Ecology — Primary and secondary rain forest. Soil: sand or sandy clay. Altitude: sea level up to 1550 m. Flowering: March, May to November; fruiting: throughout the year.

Uses — Used as timber.

Vernacular names — Peninsular Malaysia: Tampoi. Sumatra: Ketjipot, semasam, semasam pris. Sumatra (Banka): Masput (Malay). Sumatra (Simeuluë Island): Toetoen bolawak iteung pajo, toetoen bolawah kèhè-pajo. Borneo: Perepat (Malay); kayu massam, sangkurat.

36. Baccaurea tetrandra (Baill.) Müll.Arg.

Baccaurea tetrandra (Baill.) Müll.Arg. in DC., Prodr. 15, 2 (1866) 465; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 49; Merr., Enum. Philipp. Fl. Pl. 2 (1923) 411; Fernando, Philipp. J. Biol. 8 (1979) 307; Airy Shaw, Enum. Euphorb. Philipp. (1983) 10; Haegens, Blumea Suppl. (2000) 185, Fig. 3.26, Map 3.41, Photo 12. — Adenocrepis tetrandra Baill., Étude Euphorb. (1858) 601; Adansonia 4 (1863) 135. — Lectotype (designated by Haegens, 2000): Cuming 982 (lecto L; isolecto P, TCD, US), Philippines.

Baccaurea stipulata J.J.Sm., Icon. Bog. 4 (1910) 32, t. 311; Merr., J. Straits Branch Roy. Asiat. Soc. (1921) 331; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 56; Merr., Univ. Calif. Publ. Bot. 15 (1929) 151; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 55; Kew Bull. 36 (1981) 266. — Lectotype (designated by Haegens, 2000): Hallier 3445 (lecto L), Borneo, Amai Ambit.

Baccaurea terminalifolia Elmer, Leafl. Philipp. Bot. 4 (1911) 1277. — Lectotype (designated by Haegens, 2000): Elmer 13089 (lecto DS; isolecto GH, K, L, US), Philippines, Palawan, Puerta Princesa, Mt Pulgar.

Baccaurea brevipedicellata Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 55; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 54. — Lectotype (designated by Haegens, 2000): Hose 48 (lecto L; isolecto K), Borneo, Sarawak, Baram District.

Shrub or tree 2–20 m high, dbh 3–29 cm, sometimes fluted, buttresses absent; branchlets (sub)glabrous, often thickened at tip, Terminalia branching pattern well-developed. Indumentum of simple hairs. Bark brown to grey to whitish green when fresh, brown to grey when dry, 0.1–2 mm thick, smooth to slightly rough, corky, peeling off in 5–8 long cm strips, soft; inner bark red to purple to brown to yellow to white, c. 1.5 mm thick, soft. Heartwood brown. Leaves: petiole 4–75 mm long, (sub)glabrous, transverse cracks at base and tip, brown to grey when dry, raised glands present; stipules 3–11 by 1.2–4.5 mm, caducous (to late caducous), glabrous to densely hairy outside, glabrous inside, margin often ciliate, sometimes hyaline; lamina obovate to elliptic, (3.2–)5–21 by 1.5–12.3 cm, l/w ratio 1.4–3.4, papery; base rounded to cuneate; apex (retuse to) obtuse to acute, up to 20 mm long; upper surface glabrous, raised glands usually present, often granulate, (light yellow-)green when fresh, (greenish to brownish) white, nervation slightly raised; lower surface glabrous, nervation subglabrous, raised glands usually present, discoid glands absent, light green when fresh, (pale greenish to greyish) brown when dry; secondary veins (2–)4–9 per side, often closed at margin; nervation reticulate to weakly scalariform, white to brown when dry; young leaves red. Staminate inflorescences axillary to just below the leaves, solitary to 3(–many) clustered together, 1.5–17 cm long, 0.5–1.5 mm thick, sparsely to densely hairy, many-flowered, flowers scattered along inflorescence, rachis white to red to purple; bracts 1 per branchlet, 0.3–2 mm long, persistent to caducous, glabrous to densely hairy outside, (sub)glabrous inside, margin ciliate, not hyaline; bracteoles (0–)2, up to 1 mm long, subglabrous to densely hairy outside, glabrous inside; branchlets cylindrical, 0–2.5 mm long, densely hairy, 3-flowered. Staminate flowers 1.6–4.5 mm diam., yellow to white; pedicel 0.8–3 mm long, upper part 0.3–2 mm long, densely hairy; sepals 4 or 5, spatulate to sometimes ovate, 0.9–3 by 0.6–1.3 mm, sometimes apically recurved, sparsely to densely hairy outside, densely hairy inside; staminodes absent or 4 or 5; stamens (3 or) 4 or 5, 0.6–1.4 mm long, (sub)glabrous; ﬁlaments glabrous to subglabrous, 0.5–1.3 mm long, often apically geniculate; anthers 0.15–0.25 by 0.15–0.3 by c. 0.1 mm; pistillode obtriangular to cylindrical, 0.6–1.3 mm high, densely hairy, usually hollow. Pistillate inflorescences axillary to just below the leaves (to ramiflorous), solitary to 3 clustered together, 2.5–30 cm long, 0.1–2 mm thick, sparsely to densely hairy, 5–50-flowered, rachis red; pedicel 0.5–7.5 mm long, upper part 0–1.3 mm long, sparsely hairy to densely hairy; bracts (1 or) 3 per branchlet, glabrous to densely hairy outside, glabrous inside, margin ciliate. Pistillate flowers 3–12 mm diam., yellow to green (to red); sepals 4–6, obovate to elliptic, 1.5–5.9 by 0.2–2.4 mm, sparsely to densely hairy outside, densely hairy inside, caducous; ovary globose, 1.2–2.4 by 1–2.1 mm, 2-locular, sparsely to densely hairy, pale green; style absent to 0.4 by 0.7–1 mm, densely hairy; stigmas 0.4–1 mm long, cleft for upper up to 90%, white, persistent to caducous; lobes 0.4–0.7 by 0.2–0.7 mm, glabrous with protuberances above, below glabrous to densely hairy with protuberances. Infructescences to 45 cm long, purple. Fruits globose to ellipsoid, laterally flattened, 1- or 2- or 4-seeded, fleshy capsules, 7–17 by 6.5–15 by 6.5–15 mm, loculicidally dehiscent to irregularly splitting, raised glands sometimes present, glabrous to sparsely hairy outside, glabrous inside, red (to yellow to pink to greenish white); pericarp 0.1–1.5 mm thick; column 8–14 mm long, straight, caducous; pedicel 0.5–8 mm long, upper part 0.2–5.5 mm long, red. Seeds globose to obovoid, laterally flattened, 7–13 by 5–9.5 by 3–6.5 mm; arillode blue to purple; testa red; cotyledons often folded, (3–)3.5–7.5 by 4–8.7 by c. 0.1 mm; radicle 0.9–1.8 mm long; endosperm 0.2–1 mm thick.

Distribution — Borneo, Philippines, Sulawesi, Moluccas (Halmahera only).

Habitat & Ecology — Primary and secondary rain forest, and swamp forest. Soil: sand, clay, loam. Altitude: up to 1500 m. Flowering and fruiting: throughout the year. Flowers odourless.

Uses — Arillode edible, sweet to sour.

Vernacular names — Borneo (Brunei): Kunau; mata kunau (Malay). Borneo (Sabah): Kunau-kunau, mata-pelandok. Borneo (Sarawak): Enkuni(k), kunau (Iban); sekunau (Dusun); kuni, mata pelandok. Borneo (Kalimantan): Pundung undang (Dayak); konkuning, mata pelandoeh. Philippines: Katag, saket.

Note — In the type of B. stipulata the leaf blades are 13–19 cm long and whitish when dry, whereas in the type of B. tetrandra the leaf blades are 6–14 cm long and brown when dry. However, intermediates between these characters are found frequently. The same variability is found in the leaf anatomical characters (Bodegom et al., in prep.). Baccaurea stipulata is therefore considered to be conspeciﬁc with B. tetrandra.

37. Baccaurea trigonocarpa Merr.

Baccaurea trigonocarpa Merr., Univ. Calif. Publ. Bot 15 (1929) 152; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 38; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 54; Haegens, Blumea Suppl. (2000) 188, Fig. 3.27, Map 3.42. — Type: Elmer 21525 (holo A; iso DS, K, L, US), Borneo, Sabah, Tawau.

Tree or shrub 4–15 m high, dbh 5–38 cm, buttresses absent; branchlets glabrous to rarely densely hairy, Terminalia branching pattern usually weak. Indumentum of simple and rarely stellate hairs. Bark white-brown to brown to blackish-brown when fresh, brown to grey when dry, thin, smooth; inner bark yellow to rarely yellowish-red to grey. Leaves: petiole 10–57 mm long, subglabrous to densely hairy, slightly apically and sometimes slightly basally pulvinate, raised glands sometimes present; stipules lanceolate to triangular, 4–9 by 1–3 mm, glabrous to densely hairy outside, (sub)glabrous inside, margin usually not ciliate, usually not hyaline; lamina obovate to elliptic, 7–16.5 by 3.9–8 cm, l/w ratio 1.7–2.9, papery; base cuneate to attenuate; marginal glands usually visible as small indentations; apex acuminate to cuspidate, 3–18 mm long; upper surface glabrous, rarely granulate, dark green when fresh, dark (green)-brown when dry; lower surface (sub)glabrous, glabrous to densely hairy at veins, raised glands usually absent, discoid glands present, sparse to in a row between the secondary veins, dark green when fresh, green to brown when dry; secondary veins 6–10 per side, closed at margin; nervation reticulate to weakly scalariform. Staminate inflorescences axillary to just below the leaves, solitary, 1–3 cm long, c. 1 mm thick, densely hairy, many-flowered, flowers clustered at the tip of inflorescence; bracts absent; bracteoles usually 1 per branchlet, c. 1 mm long, densely hairy outside, subglabrous inside, margin ciliate, not hyaline; branchlets cylindrical, c. 1 mm long, densely hairy, 3-flowered. Staminate flowers 1–2.2 mm diam., pale yellow, sessile; sepals (5 or) 6, obovate, 0.5–1 by 0.4–0.8 mm, apically straight, densely hairy outside and inside; stamens 6, 0.9–1 mm long, glabrous; ﬁlaments c. 0.9 mm long, straight; anthers c. 0.1 by 0.2 by 0.1 mm; pistillode obtriangular, c. 0.5 mm high, velutinous, solid. Pistillate inflorescences usually ramiflorous (to axillary or cauline), solitary (to 4 clustered together), 1.5–3 cm long, c. 1 mm thick, densely hairy, 7–many-flowered; pedicel 1–1.5 mm long, upper part 0.2–0.5 mm long, geniculate (90° bent), densely hairy; bracts (1–)3 per branchlet, persistent, glabrous to densely hairy outside, densely hairy inside, margin ciliate. Pistillate flowers 1–2 mm diam., yellow; sepals (5 or) 6, 0.5–1.2 by 0.2–0.9 mm, ovate, densely hairy outside and inside, persistent; ovary globose, 1–1.4 by 0.7–0.9 mm, (2- or) 3- or 4-locular, velutinous, wings usually present, 6 or 8, small; style absent; stigmas c. 0.5 mm long, glabrous, persistent. Infructescences 4–50 cm long, 1–2 mm thick. Fruits 3- or 4-angular (to globose), 1- or 2-seeded, fleshy capsules, 8–15 by 9–14.5 by 9–14.5 mm, dehiscing loculicidally, raised glands usually absent, (sub)glabrous outside, glabrous inside, subglabrous at column, yellow to orange to purple; pericarp 0.5–1.5(–2) mm thick; column 7–11 mm long, straight, sometimes caducous; pedicel 4–10(–17) mm long, upper part 0.5–3 mm long. Seeds globose to ellipsoid, laterally flattened, 6.2–9 by 6–8 by 3.5–5 mm; arillode orange to purple to reddish; testa white; cotyledons 5–7 by 5.5–7 by 0.1–1; radicle 0.1–0.2 mm long; endosperm thin.

Distribution — Borneo, Singapore but probably only in cultivation.

Habitat & Ecology — Primary rain forest and kerangas forest. Soil: sand, clay, loam. Altitude: 30–1200 m. Flowering: June, August, September; fruiting: January, April to November.

Uses — Firewood.

Vernacular names — Borneo: Engkuhi (Iban); tikak cabo (Murut); butoh pranuk, ngiboruok, ombak pahung, pugé laboh, rambai hutan.

38. Baccaurea velutina (Ridl.) Ridl.

Baccaurea velutina (Ridl.) Ridl., J. Bot. 62 (1924) 300; Fl. Malay Penins. 5 (1925) 331; Whitmore, Tree Fl. Malaya 2 (1973) 67; Haegens, Blumea Suppl. (2000) 190, Map 3.43. — Baccaurea reticulata Hook.f. var. velutina Ridl., Fl. Malay Penins. 3 (1924) 246; Smitinand, Nat. Hist. Bull. Siam Soc. 22 (1967) 170. — Lectotype (designated by Haegens, 2000): SF (Nur) 11319 (lecto K; isolecto A, SING).

Shrub or tree (2.1–)9–15(–18) m high, dbh 6.4–24 cm, buttresses absent; branchlets sparsely hairy to velutinous; young shoots pale to red-brown when dry, Terminalia branching pattern well-developed. Indumentum of stellate and simple hairs. Bark brown to grey-brown to orange-brown when fresh, smooth; inner bark red to brown, thin, ﬁbrous. Leaves: petiole 18–80 mm long, velutinous with long and short hairs, often with longitudinal cracks when old, light to blackish brown when dry, raised glands or lenticels often present; stipules 3–7 by 1.5–3 mm, velutinous outside, glabrous inside, margin ciliate, not hyaline; lamina ovate to obovate, 5.2–19.3 by 3.5–11.2 cm, l/w ratio 1.4–2.8, papery to somewhat leathery; base cuneate to attenuate (to rounded); apex rounded to acuminate, up to 11 mm long; upper surface glabrous, raised glands present, sometimes granulate, (dark to grey)-brown when dry; lower surface subglabrous to sparsely hairy, nervation sparsely hairy to velutinous, raised glands present, discoid glands present, hairy, if many then in a row between secondary veins, brown when dry; secondary veins 5–7 per side, usually open at margin; nervation reticulate; young leaves yellowish brown when fresh. Staminate inflorescences axillary, solitary or in pairs, 2.5–9 cm long, rachis 1–1.8 mm thick, branches minute, velutinous, many-flowered, flowers scattered along inflorescence; bracts 1 per branchlet, 4–7.3 mm long, densely hairy outside, (sub)glabrous inside, margin ciliate, not hyaline; bracteoles 4(–8), small; branchlets, cylindrical, 0.5–1 mm long, velutinous, 4–12-flowered. Staminate flowers 2.5–5 mm diam., yellow to brown; pedicel 0.5–1.3 mm long, upper part 0.2–0.9 mm long, velutinous; sepals 4 (or 5), ovate, 1.8–4 by 1–1.8 mm, recurved to straight at apex, velutinous outside and inside; stamens 5 or 6, 0.3–0.9 mm long, glabrous; ﬁlaments 0.1–0.7 mm long, straight; anthers 0.15–0.2 by 0.2–0.25 by 0.1–1.15 mm; disc absent; pistillode cylindrical to obtriangular, 0.6–1.3 mm high, velutinous. Pistillate inflorescences axillary, solitary, 2.5–5.5 cm long, 1.2–1.5 mm thick, velutinous, few-flowered; pedicel 1.5–3.5 mm long, upper part 0.5–2 mm long, velutinous; bracts 2 per branchlet, ovate, velutinous outside, glabrous inside, margin ciliate. Pistillate flowers 5–10 mm diam., yellow; sepals 4 (and a small 5th), ovate, (3–)6–10 by (1–)2–3 mm, velutinous outside and inside, persistent to caducous; ovary cylindrical, 2.5–4 by c. 2.5 mm, 3-locular, velutinous, dark brown; style 0.5–1 by 1–1.5 mm, white velvety; stigmas 1.2–2.2 mm long, cleft for upper c. 50%, persistent and up to 3 mm long; lobes 0.6–1.1 by 0.2–0.4 mm, glabrous above, velutinous below. Fruits globose, 2–6-seeded, berries to fleshy capsules, 23–28 by 25–31 by 25–31 mm, 2.5 cm diam. when fresh, usually only 1 layer dehiscing loculicidally when dry, raised glands present, (slightly) ruminate when dry, velutinous outside, (sub)glabrous inside, septa subglabrous, (red-)brown; pericarp 3–7.5 mm thick; column 15–17 mm long, straight; pedicel 4–5 mm long, upper part 2–3 mm long. Seeds ellipsoid, laterally flattened, 7.5–10 by 4.6–8 by 3–5 mm; arillode yellow to orange; cotyledons 5.2–6 by 5–6.9 by 0.1–0.5 mm; radicle 1–1.5 mm long.

Distribution — Peninsular Malaysia.

Habitat & Ecology — Rain forest, on ridges, rarely in swamps. Altitude: 330–1450 m. Flowering: April, August, September; fruiting: February to May, September.

Uses — Arillode edible, sour.

Vernacular names — Peninsular Malaysia: Tampoi, tampoy tungau.

Species dubiae

No original material of the following taxa could be traced or obtained so that their taxonomic status could not be evaluated.

Baccaurea airyshawii Chakrab. & M.Gangop., J. Econ. Taxon. Bot. 18 (1994) 419. — Type: Kunstler 1408 (holo CAL, n.v.).

Baccaurea celebica Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 69; Airy Shaw, Kew Bull. 37 (1982) 8. — Type: Warburg 15565 (n.v.).

Baccaurea henii Thin, J. Biol. (Vietnam) 9, 2 (1987) 37. — Type: Duong van Hen s.n. (holo HM, n.v.).

Pierardia macrostachys Wight & Arn., Icon. Pl. Ind. Orient. (1852) 30; Baill., Adansonia 3 (1863) 140. — Lectotype (designated by Haegens, 2000): Wight (lecto plate 1912 in Wight & Arn., Icon. Pl. Ind. Orient. (1852) 28).

Hooker (1885) has considered this name a synonym of B. sapida (= B. ramiflora). The original description and plate of B. macrostachys are, however, not congruent with Hooker’s description of B. sapida. The original description and plate of B. macrostachys are also incongruent with the modern description of B. ramiflora (see no. 34). In B. macrostachys (5–)8–10 stamina are present, whereas B. ramiflora has 5–8 stamina. In my opinion, the plant with staminate inflorescences depicted in Wight & Arn. (1852: plate 1912) is a true Baccaurea, but is maybe not conspeciﬁc with B. ramiflora. The drawing is too crude to establish the exact identity of the depicted plant. Two oddities appear in the description and on plate 1913 (Wight & Arn., 1852) of the plant with pistillate inflorescences and infructescence: the seed has no arillode; the leaves are opposite (but Wight & Arn. were not sure if the leaves belonged to the fruits). For these reasons it is extremely questionable if the female plant is a Baccaurea. The plate with the staminate flowers is therefore chosen as lectotype.

Baccaurea reniformis Chakrab. & M.Gangop., J. Econ. Taxon. Bot. 19 (1995) 449. — Type: King’s Collector 10196 (holo CAL, n.v.).

Excluded names

Baccaurea cavaleriei H.Lév., Fl. Kouy-Tchéou (1914) 158; Rehder, J. Arnold Arbor. 18 (1937) 281; Airy Shaw, Kew Bull. 19 (1965) 314. — Type: Cavalerie 3299 (holo E), China, Kweichow, Lo Fou. = Cleidiocarpon cavaleriei (H. Lév.) Airy Shaw.

Baccaurea esquirolii H.Lév., Fl. Kouy-Tchéou (1914) 158; Rehder, J. Arnold Arbor. 18 (1937) 281. — Type: Cavalerie 3458 (lecto A, selected here), China, Nom-You-Se-Ki-Fou. = Sapium rotundifolium Hemsl.