Malesian Euphorbiaceae Descriptions

3. AGROSTISTACHYS (Euphorbiaceae)

S. Sevilla & P.C. van Welzen

Sevilla, S. & P.C. van Welzen. 2001. Revision and Phylogeny of Agrostistachys and Chondrostylis (Euphorbiaceae). Blumea 46: 71–97.

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Genus description

Key to the species

Species descriptions

Excluded species

Agrostistachys Dalzell

    Agrostistachys Dalzell, Hooker’s J. Bot. Kew Gard. Misc. 2 (1850) 41; Airy Shaw, Kew Bull. 14 (1960) 472; Kew Bull. 26 (1972) 210; Whitmore, Tree Fl. Malaya 2 (1973 ) 52; Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 26; Kew Bull., Addit. Ser. 8 (1980) 23; Kew Bull. 36 (1981) 248; Alph. Enum. Euphorb. Philipp. Is. (1983) 3; G.L.Webster, Ann. Missouri Bot. Gard. 81 (1994) 69; Sevilla & Welzen, Blumea 46 (2001) 77; Radcl.-Sm., Gen. Euphorbiacearum (2001) 132; Welzen & Sevilla in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 41; G.L.Webster in Kubitzki, Fam. Gen. Vasc. Pl. 11 (2014) 110. — Type: Acalypha indica Dalzell

    Sarcoclinium Wight, Icon. Pl. Ind. Orient. 5 (1852) 24. — Type: Sarcoclinium longifolium Wight [= Agrostistachys borneensis Becc.]

    Heterocalyx Gagnep., Notul. Syst. 14 (1950) 33. — Type: Heterocalyx laoticus Gagnep. [= Agrostistachys indica Dalzell]

Shrubs or trees, dioecious (monoecious A. hookeri and A. staminodiatus); flowering twigs glabrous. Indumentum of simple hairs only. Stipules ovate to narrowly triangular, margin entire or ciliate with trichomes, apex rounded to aristate, caducous to late caducous or persistent, scars up to 5 cm below the leaves. Leaves alternate, distichous to spiral, simple; petiole reniform in transverse section, basally pulvinate; blade elliptic to obovate, symmetric, papery to coriaceous, base attenuate (emarginate to rounded in A. sessilifolia; sometimes obtuse to rounded in A. borneensis), margin entire and without glands to dentate and teeth with glands, apex rounded to caudate, upper and lower surface smooth, glabrous (sometimes the young leaves pubescent); venation raised on both sides, especially beneath, nerves 10–40 per side, at the margin looped and joined, veins and veinlets scalariform to reticulate. Inflorescences axillary and sometimes cauliflorous, racemes, single or up to 15 superposed above each other per axil (or growing from the same point), unbranched, glabrous to sericeous. Bracts alternate to spiral along rachis (overlapping each other and hiding rachis in A. indica and sometimes in other species when flowers still in bud), margin ciliate, sometimes with trichomes, glabrous to sericeous, sometimes covered with papillae outside, inside sometimes with glands. Flowers single or up to 10 per bract, actinomorphic; pedicels glabrous to sericeous; sepals 2–5, valvate, basally connate; petals 5 (absent in A. hookeri), free (to connate). Staminate flowers: disc lobes 4–7 (annular in A. hookeri); stamens 8–16, glabrous, filaments thread-like, anthers 2-locular, deltoid, basally deeply cleft, basifixed, opening introrse with longitudinal slits, connective apically elongated (in A.hookeri spade-like with the anthers in the basal lobes); pistillode absent or present, 2– or 3–partite. Pistillate flowers: sepal margin ciliate, sometimes with trichomes; petal margin entire; disc lobes 5 (annular in A. gaudichaudii and A. hookeri); pistil (2- or) 3-locular, 1 ovule per locule, ovary glabrous to sericeous, style subsericeous or absent, stigmas 3, split in upper 0.5–1 mm (not split in A. hookeri), upper surface with broad papillae, lower surface covered with smaller papillae. Fruits rhegmas, lobed, septicidal and partly apically locucidal, smooth, glabrous (to subsericeous), wall thin, woody; column after dehiscence apically not broadened, with narrow, quite entire septal margins; sepals and styles persistent. Seeds obovoid, base very acute, apically with a caruncle.

    Distribution — South India, Sri Lanka, Thailand, Malay Peninsula, Sumatra, Borneo, Philippines, New Guinea.

Key to the species

1. Agrostistachys borneensis Becc.

    Agrostistachys borneensis Becc., For. Borneo (1902) 331; Airy Shaw, Alph. Enum. Euphorb. Philipp. Is. (1983) 3; Sevilla & Welzen, Blumea 46 (2001) 78, fig. 2, 3e, f, h, map 1; Welzen & Sevilla in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 41, fig. 8. — Lectotype (designated by Sevilla & Welzen, 2001): Beccari PB 3117 (lecto FI, photo L; isolecto K, P), Sarawak, Kuching.

    Sarcoclinium longifolia Wight, Icon. Pl. Ind. Orient. 5 (1852) 24, pl. 1887, 1888; Müll.Arg. in DC., Prodr. 15, 2 (1866) 727. — Agrostistachys longifolia (Wight) Trimen, Syst. Cat. Fl. Pl. Ceylon (1885) 81, nom. illeg., later homonym, non Kurz; Benth. ex Hook.f., Fl. Brit. India 5 (1887) 407; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vi (1912) 100, fig. 49 E; Whitmore, Tree Fl. Malaya 2 (1973) 53; Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 27; Kew Bull., Addit. Ser. 8 (1980) 24. — Agrostistachys coriacea Trimen, Handb. Fl. Ceylon 4, Suppl. (1931) 265, nom. superfl. — Type: Wight s.n. (n.v.), India, Western slopes of the Neilgherries (see note 1).

    Agrostistachys longifolia (Wight) Trimen var. latifolia Hook.f., Fl. Brit. India 5 (1887) 407. — Agrostistachys latifolia (Hook.f.) Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vi (1912) 100; Merr., J. Straits Branch Roy. Asiat. Soc., special number (1921) 337. — Syntypes: King’s collector 3459 (K, photo L), Malaysia, Perak, Larut; King’s collector 3523 (K, photo L), Malaysia, Perak, Larut; Scortechini 1985. (K, photo L), Malaysia, Perak.

    Agrostistachys meeboldii Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vi (1912) 100.—Syntypes: Meebold 13048 (B, †), India, Malabar coast, Undanbanthola; Meebold 13079 (B, †), India, Malabar coast, Udanbanthola; Wight KD 2612 (K, photo L; A, L), India, Trabancore, Courtallam.

    Agrostistachys leptostachya Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vi (1912) 102; Merr., Philipp. J. Sci. 16 (1920) 553; J. Straits Branch Roy. Asiat. Soc., special number (1921) 337; Enum. Philipp. Fl. Pl. 2 (1923) 428. — Agrostistachys longifolia (Wight) Trimen var. leptostachya (Pax & K.Hoffm.) Whitmore, Gard. Bull. Straits Settlem. 26 (1972) 52; Tree Fl. Malaya 2 (1973) 53. — Lectotype (designated by Sevilla & Welzen, 2001): Hose 162 (lecto K, photo L; isolecto L), Borneo, Sarawak (see note 2).

    Agrostistachys intramarginalis Philcox, Kew Bull. 50 (1995) 119. —Type: Waas 1556 (holo K, photo L; iso NY, US), Sri Lanka, Sabaragamuwa Prov., Ratnapura Distr., Walankanda.

Shrubs, to 7 m high; d.b.h. to 1.9 m; flowering twigs to 1.5 cm thick, rather smooth to ribbed to rough. Outer bark green to brown to grey to black, smooth to narrowly fissured; inner bark yellowish to dark red to brown; sapwood cream to pale yellow to brown. Stipules ovate to triangular, 0.5–4 by 1–8 mm, late caducous to persistent, scars up to 1.5 cm below leaves, sometimes thick, margin entire, ciliate with trichomes, apex rounded to acute, glabrous to subsericeous abaxially, with trichomes at base adaxially, venation distinctly parallel. Leaves: petiole 0–2 cm long; blade elliptic to obovate, 6–70 by 2–18 cm, index 2–5.7, subcoriaceous to coriaceous, base attenuate, sometimes obtuse to rounded, margin entire, apex rounded to cuspidate; veins scalariform, veinlets scalariform to reticulate. Inflorescences axillary and cauliflorous, single or up to 4 together, up to 35 cm long, glabrous to sericeous; bracts overlapping each other and hiding rachis when in bud, ovate (to obovate), 1–4 by 1–4 mm, margin sometimes with trichomes, apex rounded to acuminate, outside sometimes covered with papillae, inside with glands that exude white latex, glabrous to subsericeous, venation usually distinctly parallel. Flowers single or up to 5 together; pedicel glabrous to sericeous; disc lobes orbicular to broadly ovate, glabrous. Staminate flowers 2–4 by 1.5–4 mm; pedicel 1–3 mm long; sepals 2–5, lobes  1.5–3 by 1–2 mm, apex mucronate to acuminate, venation usually distinctly parallel; petals 5, 1–2.5 by 1–2.5 mm, glabrous, margin entire, apex irregular to obcordate; disc lobes 4–6, 0.3–1 mm in diam.; stamens 8–12, in two whorls, the outer whorl inserted between the disc lobes, second whorl sometimes tube-like around the pistillode, filaments free (first whorl) or shortly connate (second whorl), 2–3 mm long, connective sometimes covered with papillae; pistillode 2- or 3-partite, glabrous to subsericeous. Pistillate flowers 2–5 by 2–5 mm, pedicel 2–5 mm long; sericeous; sepals 5, lobes 2–3 by 1–2 mm, margin with trichomes, apex acute; petals 5, 1–2 by 1.5–2 mm, margin entire, apex irregular; disc lobes 5, 0.5–1 mm in diam., sometimes incompletely divided; ovary 1–3 by 1.5–4 mm, subsericeous to sericeous, stigmas sessile, 1– 2.5 mm long, split in upper 0.5–1 mm, glabrous. Fruits 0.5–1 by 0.5–1 cm high. Seeds 4–6 by 3–5 mm.

    Distribution — South India, Sri Lanka, Peninsular Thailand, Malay Peninsula, Sumatra, Borneo, Philippines, and New Guinea.

    Habitat & Ecology — Primary and secondary (logged) forests, wet evergreen forests, dipterocarp forests, swamp forests, kerangas forests, montane forests, mossy forests, along forest roads, ridges, streams, on gentle to steep slopes, on rock in dry exposed areas, common in undergrowth and abundant in open areas. Soil: red, yellow sandy or light brown clay, limestone, acid sand. Alt: 50–980 m. Flowering and fruiting throughout the year.

    Uses — Cosmetic use as tooth paste to dye the teeth black: burned wood exudes a black latex, which is smeared on the teeth. Repeated treatments are needed after 3 days otherwise the colour disappears.

    Vernacular names — Peninsular Malaysia: Perak: Jenjulong. Borneo: Sarawak: Gelimtum, malo, malau pucuk, malau puchok, maha bu/punubéru; Brunei: Julong-julong, malau puchok. Papua New Guinea: Wagu.

    Notes — 1. The combination Agrostistachys longifolia was already made by Kurz in 1875 (a synonym of A. indica). Therefore, the well-known name A. longifolia may not be used and the second oldest epithet on the species level, borneensis, has to be preferred.

2. Beccari PB 1381 has been used as a syntype of A. borneensis by Beccari (1902) and as a syntype of A. leptostachya by Pax & Hoffmann (1912). Therefore, to prevent further confusion, A. leptostachya has to be lectotypified with Hose 162.

3. This species is very variable in the shape and size of the leaves, length of the petioles, and size of the inflorescences, not only between the different islands, but even on a single island.

2. Agrostistachys gaudichaudii Baill. ex Müll.Arg. 

    [Sarcoclinium gaudichaudii Baill., Étude Euphorb. (1858) 310, nom. nud.] — Agrostistachys gaudichaudii Baill. ex Müll.Arg., Linnaea 34 (1865) 144; in DC., Prodr. 15, 2 (1866) 725; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vi (1912) 103; Airy Shaw, Kew Bull. 26 (1972) 210; Whitmore, Tree Fl. Malaya 2 (1973) 52; Sevilla & Welzen, Blumea 46 (2001) 81, map 2; Welzen & Sevilla in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 42, plate II: 2. — Lectotype (designated by Sevilla & Welzen, 2001): Gaudichaud 66 (lecto P; isolecto A, P), Malaysia, Penang Island (see note under description).

    Agrostistachys maingayi Hook.f., Fl. Brit. India 5 (1887) 406; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vi (1912) 105. — Type: Maingay KD 1425 (1571) (holo K, photo L; iso L), Malaysia.

    Agrostistachys filipendula Hook.f., Fl. Brit. India 5 (1887) 407; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vi (1912) 99. — Syntypes: Hullett s.n. (K, photo L), Singapore; Hullett 852 (K, photo in L), Singapore; King’s collector 4669 (K, photo L), Malaysia, Perak, Larut; King’s collector 5736 (K, photo L, L), Malaysia, Perak, Larut.

Shrubs or trees, to 15 m high, d.b.h. to 33 cm; flowering twigs to 1 cm thick,  smooth. Outer bark red-brown to grey; inner bark pale brown to yellow; sapwood white to cream to orange. Stipules narrowly triangular, 2–20 by 2–7 mm, sericeous, caducous, scars up to 4.5 cm below leaves. Leaves alternate; petiole 0.5–4.5 cm long; blade (narrowly) elliptic to obovate, 14–54.5 by 4.5–22 cm, index 1.7–5.2, papery to subcoriaceous, base attenuate, margin entire to crenulate, apex acuminate to cuspidate, glossy on both sides, darker above, nerves 11–24 per side, veins scalariform, veinlets scalariform to reticulate. Inflorescences single or up to 15 above each other, up to 20 cm long, glabrous to subsericeous; bracts alternate along rachis, overlapping each other and hiding rachis when flowers still in bud, broadly ovate, 2.5–5 by 2.5–6 mm, apex forming obtuse hood, sericeous, outside glabrous to subsericeous. Flowers single or up to 3(–10) toegether; pedicel glabrous to subsericeous; sepals longer than petals, lobes elliptic to ovate, nerves distinctly parallel; petals 5, free, obcordate, glabrous, margin entire, apex emarginate; disc glabrous, yellow-red. Staminate flowers 2–3 by 2.5–3.5 mm; pedicel 2–3 mm long; sepals 3, c. 2 by 1.3–2 mm, glabrous, margin entire, apex acute; petals 1.3–1.5 by 1.2–1.5 mm, white to yellow to pale green; disc lobes 5, orbicular to reniform, 0.4–0.5 by 0.4–1 mm; stamens 10, free, 1.5–3 mm long, anthers pale yellow; pistillode absent. Pistillate flowers 2–3 by 2.5–3.5 mm; pedicel 0.2–1 cm long; sepals 5, 2–3 by c. 1 mm, apex rounded to acute; petals 1.5–3 by 1.5–3 mm, red-brown; disc annular; ovary 1–2.5 by 1.5–3 mm, sericeous; stigma sessile 3, 1.5–2 mm long, split in c. upper 0.5 mm part, glabrous. Fruits: 0.5–1.2 by 0.8–1.5 cm high, grey to green. Seeds  7–9 by 5–8 mm.

    Distribution — Peninsular Thailand and Peninsular Malaysia (Kedah, Kelantan, Trengganu, Pahang, Malacca, Johore).

    Habitat & Ecology — Lowland primary forests, evergreen forests, dipterocarp forests, margin of primary evergreen forests, secondary forests, bamboo forests, swampy forests, logged forests, on hills, in shades of taller trees, along streams, in cleared areas. Soil: limestone, alluvium. Alt.: sea level to 50 m. Flowering: mainly Februaty to April, less so in June to September; fruiting: March to June, October to December.

    Note —. Helfer 4921 was used by Müller as one of the syntypes of A. gaudichaudii, but he also indicated it to be the type of A. indica Dalzell subsp. longifolia Müll.Arg. (a synonym of A. indica ). In order to solve this problem, the other syntype of A. gaudichadii, Gaudichaud 66, has been selected as the lectotype, which makes Helfer 4921 the only type of subsp. longifolia (and var. integra Pax & K. Hoffm., an illegitimate name which should have repeated the subspecific name).

3. Agrostistachys indica Dalzell 

    Agrostistachys indica Dalzell, Hooker’s J. Bot. Kew Gard. Misc. 2 (1850) 41; Müll.Arg. in DC., Prodr. 15, 2 (1866) 726; Hook.f., Fl. Brit. India 5 (1887) 406; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vi (1912) 103, fig. 20 C, D; Airy Shaw, Kew Bull. 14 (1960) 472; Kew Bull. 26 (1972) 210; Kew Bull. 29 (1974) 312; Kew Bull., Addit. Ser. 4 (1975) 26; Kew Bull., Addit. Ser. 8 (1980) 24; Sevilla & Welzen, Blumea 46 (2001) 83, map 4; Welzen & Sevilla in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 43. — Agrostistachys indica Dalzell subsp. genuina Müll.Arg. in DC., Prodr. 15, 2 (1866) 726, nom. illeg.; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vi (1912) 104. — Type: Hb. Dalzell s.n. (holo K, photo L), India, Bombay.

    Agrostistachys indica Dalzell var. longifolia Müll.Arg. in DC., Prodr. 15, 2 (1866) 726; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vi (1912) 105. — Agrostistachys longifolia (Müll.Arg.) Kurz, Prelim. Rep. Forest Pegu (1875) App. A cxi, App. B 79; Forest Fl. Burma 2 (1877) 377. — Agrostistachys indica Dalzell var. subintegra Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vi (1912) 105, nom. illeg. — Type: Helfer 4921 (holo K, photo L; iso in A, C), Myanmar, Tenasserim (see note 1 under A. borneensis and under A. gaudichaudii).

    Agrostistachys maesoana Vidal, Revis. Pl. Vasc. Filip. (1886) 242; Merr., Enum. Philipp. Fl. Pl. 2 (1923) 428. — Agrostistachys indica Dalzell subsp. longifolia Müll.Arg. var. maesoana Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vi (1912) 105; Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 24; Alph. Enum. Euphorb. Philipp. Isl. (1983)3.— Type: Vidal 897 (holo PNH, †; iso in L, A), Philippines, Luzon, Tayabas Prov., Guinayangan.

    Heterocalyx laoticus Gagnep., Notul. Syst. 14 (1950) 33. — Syntypes: Kerr 20895 (L, P), Kerr 20895A (L, P), Laos, Tawieng, Chieng-kwang.

Shrubs or trees, to 0.5 m high, d.b.h.  to 3 cm; flowering twigs  to 3 mm thick, rather smooth, quite slender and flexible. Outer bark brown to yellowish green, grey mottled, smooth; inner bark yellow; sapwood white. Stipules narrowly triangular, 4–17 by 2–4 mm, caducous, scars up to 5 cm below leaves;. Leaves alternate; petiole 0.2–7 cm long; blade (narrowly) elliptic to obovate, 8–40 by 1.5–13 cm, index 2.2–5.7, papery to subcoriaceous, base attenuate, margin dentate, teeth ending in a gland, apex acute to caudate, shining on the upper surface, slightly paler beneath; venation yellowish beneath, nerves 10–27 per side, veins scalariform, veinlets scalariform to reticulate. Inflorescences single or up to c. 6 above each other, up to 3 cm long, subsericeous, white to dark green; bracts overlapping each other and hiding rachis, also when flowers not in bud, ovate, 2–6 by 2–5 mm, yellowish green, apex acuminate, outside sericeous, venation distinctly parallel. Flowers single; pedicel sericeous; sepals 2–5, lobes ovate, margin ciliate, subsericeous to sericeous outside, nerves distinctly parallel; petals 5 (9), longer than sepals, imbricate, glabrous, margin entire, apex obtuse to acuminate, white to brown-red; disc lobes orbicular to broadly obovate, glabrous, dark brown when dry. Staminate flowers 3.5–5 by 3–4 mm; pedicel 4–6 mm long; sepal lobes 3–4 by 2–3.5 mm, apex mucronate; petals ovate to narrowly obovate, 2–5 by 1–2.5 mm, cream; disc lobes 5–7(–10), 0.4–0.5 mm in diam.; stamens 13–16, filaments basally united, 2.5–4 mm long; pistillode absent. Pistillate flowers 5–7 by c. 3 mm; pedicel c. 4 mm long; sepal lobes 3–5 by 1–2 mm, subapically mucronate, apex acute; petals orbicular to obovate, 2.5–3.5 by 1–1.5 mm, pinnate venation distinct; disc lobes 5 (or 10), 0.5–1 mm in diam.; ovary 0.5–1.5 by 1–1.7 mm, sericeous; style c. 1 mm long, subsericeous; stigmas 3, 1–1.5 mm long, split in upper 0.5 mm, subsericeous below. Fruits: 12–13 by 8–10 mm high. Seeds 5–8 by 5–7 mm.

    Distribution — South India, Sri Lanka, Myanmar, Laos, Vietnam, Thailand, Borneo, Philippines, and Papua New Guinea.

    Habitat & Ecology — Primary forests, secondary wet evergreen forests, dipterocarp forests; on low hills or ridges, frequent along dry water courses, rivers, and cascades, abundant in undergrowth. Soil: compact, clay, sand, among boulders. Alt: sea level to 400 m. Flowering: January to February, April to July; fruiting: March to July, September to October.

    Vernacular names — Philippines: Haginis (Panay Bisaya); malagasaha (Kagascas, Tayabas Prov.); puso-puso (Quezon Prov., Tayabas Prov.); cacao-cacao/kakao-kakao (Bisaya); malacapé (Palawan I.); ulam-ulam (Tagbanua); gaginis (Negros Occidental); (partly after Merrill, 1923).

    Note—The specimens of A. indica from India and Ceylon, (usually) have (linear) elliptic bladess and long petioles; the specimens from Thailand and Vietnam obovate blades and long petioles; and the specimens from Borneo and Philippines (A. maesoana) obovate blades and short petioles (sometimes even indistinct).

4. Agrostistachys sessilifolia (Kurz) Pax & K.Hoffm.

    Agrostistachys sessilifolia (Kurz) Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vi (1912) 102; Airy Shaw, Kew Bull. 20 (1966) 26; Kew Bull., Addit. Ser. 4 (1975) 27; Kew Bull. 36 (1981) 249; Sevilla & Welzen, Blumea 46 (2001) 85, map 5. — Sarcoclinium sessilifolium Kurz, Flora 33 (1875) 31. — Type: Kurz s.n. (holo K, photo L), Singapore.

    Agrostistachys longifolia (Wight) Trimen var. malayana Hook.f., Fl. Brit. India 5 (1887) 407. — Syntypes: Griffith KD 4739 (C; K, photo L; P L), Malacca;  Maingay KD 1407 (K, photo L; L), Malacca; Wallich 7717 (K, photo L; L), Malaysia, Penang.

    Agrostistachys sessilifolia (Kurz) Pax & K.Hoffm. var. graciliflora Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 27. — Type: SAN (Talip) 65844 (K; photo L), Sabah, Tawau Distr., Mt. Wullersdorf.

Shrubs, to 6 m high, d.b.h. to c. 1.9 m; flowering twigs to 1.5 cm thick, rather smooth. Outer bark grey; inner bark light brownish; sapwood yellowish. Stipules triangular, 5–12 by 3–6 mm, late caducous, scars up to 1.5 cm below leaves, margin entire, apex aristate, venation distinctly parallel. Leaves alternate to distichous to spiral; petiole 0.2–0.5 cm long; blade obovate to linear obovate, 25–55 by 7–9.5 cm, index 3.5–5.6, coriaceous, base emarginate to rounded, margin entire to crenulate, apex acuminate to cuspidate, dark green above, light green below; nerves 15–30 per side, veins scalariform to reticulate, veinlets reticulate. Inflorescences axillary, single, up to 60 cm long, subsericeous; bracts alternate to spiral, ovate to triangular, 2–5 by 2–3.5 mm, apex cuspidate to caudate, subsericeous outside, venation distinct parallel. Flowers up to 10 together; pedicel sericeous; disc lobes orbicular to broadly ovate. Staminate flowers c. 4 by 2 mm; pedicel c. 3 mm long; sepals 2 or 3, 2–2.5 by c. 2 mm, margin entire, apex mucronate, glabrous; petals 5, obcordate, c. 1.5 by 1.5 mm, white; disc lobes 5, c. 0.8 mm in diam.; stamens 8–10, in two whorls, filaments basally free (outer whorl) or shortly connate (inner whorl), c. 3 mm long; pistillode 2- or 3-partite. Pistillate flowers c. 3 by 3 mm; pedicel c. 3 mm long; sepals 5, 2–4 by 1–1.5 mm, margin with trichomes, apex acute; petal number unknown, lobes obcordate, c. 2–2.5 mm long; ovary c. 1.5 by 2.5–3 mm, sericeous; stigmas sessile, 3, 2–2.5 mm long, split in upper c. 0.5 mm part. Fruits: 0.5–1 by 0.5–1 cm high. Seeds  5–6 by 4–5 mm.

    Distribution — Singapore; North Borneo (Sabah).

    Habitat & Ecology — Primary forests, evergreen forests, swampy forests, logged and disturbed forests; in low undulating country, on lower slopes or lower hillsides, on flat land, near the beach. In shaded area. Birds eat the fruit. Soil: acid sandy soils, and sedimentary rock. Alt.: 30–150 m. Flowering and fruiting: throughout the year.

    Note — The specimens from North Borneo (formerly known as A. sessilifolia var. graciliflora) have smaller stipules, shorter and thinner inflorescences, and the leaves are less wide than the specimens from Singapore. We also found specimens with the two kinds of inflorescences together, therefore, the variety graciliflora cannot be accepted as a separate taxon.

5. Agrostistachys staminodiatus Sevilla

    Agrostistachys staminodiatus Sevilla in Sevilla & Welzen, Blumea 46 (2001) 86, fig. 3a-d, g, map 6.—Type: W.J.J.O. de Wilde & B.E.E. de Wilde-Duyfjes 19505 (holo L; iso L), North Sumatra, Aceh Prov., Gunung Leuser Nature Reserves, Sikundur Forest Reserve.

Shrubs, to c. 5 m high; monoecious; flowering twigs  to 6 mm thick, glabrous, smooth. Stipules broadly obovate to orbicular, c. 5 by 5 mm, caducous to late caducous, scars up to 1.5 cm below leaves, margin ciliate, apex rounded to emarginate, trichomes adaxially, venation distinctly parallel. Leaves alternate; petiole 0.2–0.3 cm long; blade narrowly obovate, 32–45 by 8–11 cm, index 3–4.5, coriaceous, base attenuate, margin entire, apex acuminate to cuspidate; nerves 25–30 per side, veins scalariform, veinlets scalariform to reticulate. Inflorescences axillary and cauliflorous, single or up to 3 above each other, up to 25 cm long, subsericeous to sericeous; bracts alternate along rachis, broadly obovate to orbicular, 1–4 by 1–8 mm, margin with trichomes, apex rounded to emarginate, outside glabrous with papillae, inside with glands exuding white latex, nervation distinctly parallel. Flowers single or up to 8 together, staminate and pistillate flowers on same inflorescence, basally staminate, apically pistillate. Staminate flowers c. 3 by 3 mm; pedicel c. 1.5 mm long; sepals 2 or 3, lobes ovate, c. 1.5 by 2 mm, apex acuminate, outside glabrous, venation distinctly parallel; petals 5, c. 1.5 by 1.5–2 mm, shorter than sepals, glabrous, margin entire, apex irregular to obcordate; disc lobes 5, c. 5 mm in diam.; stamens c.10, filaments free, 2–3 mm long; pistillode present. Pistillate flowers 3.5–4 by 2.5–3 mm; pedicel c. 2 mm long, sericeous; sepals 2–5, lobes ovate, 1.5–2 by 1.5–2 mm, margin with trichomes; petals 5, 2–2.5 by c. 1.5 mm, margin entire, apex irregular; disc 5 lobes (sometimes incompletely divided), c. 0.5 mm high; staminodes c. 10, inserted between the disc lobes, c. 1.3 mm long; ovary c. 2 by 1.5–2.5 mm, glabrous to subsericeos; stigmas sessile, 3, 1–2 mm long, split in upper c. 0.5 mm part, glabrous. Fruits: c. 1.2 by 0.8 cm high. Seeds c. 0.5 by 0.4 mm.

    Distribution — Peninsular Malaysia and Sumatra. 

    Ecology & Habitat — Shaded places in undisturbed dense forests; on c. 15 years old logged over forests and recently logged over forests. Soil: yellow sand-loam. Alt: 50–100 m. Flowering: July and February; fruiting: May and October.

    Vernacular names — Peninsular Malaysia: Jenjulong (Pahang).

 Excluded species

Agrostistachys africana Müll.Arg., Flora 47 (1864) 534. — Syntypes: Mann 582 (B, †), Fernando Po; Mann 1079 (B, †), St. Thomas. = Pseudoagrostistachys africana (Müll.Arg.) Pax & K. Hoffm.

Agrostistachys comorensis Pax, Bot. Jahrb. Syst. 23 (1897) 523. — Syntypes: Humblot 382 (B, †), Comores; Humblot 390 (B, †), Comores; Schmidt 191 (B, †), Comores. = Tannodia cordifolia (Baill.) Baill.

Agrostistachys ugandensis Hutch., Kew Bull. (1917) 233. — Syntypes: Fyffe 19 (K, photo L), Uganda, Namalala Forest; Fyffe 64 (K, photo L), Uganda, Namalala Forest. = Pseudoagrostistachys ugandensis (Hutch.) Pax & K. Hoffm.