Malesian Euphorbiaceae Descriptions

20. BREYNIA (Phyllanthaceae)

P.C. van Welzen, K. Pruesapan, I.R.H. Telford, H.-J. Esser & J.J. Bruhl

    Van Welzen, P.C. 2003. A revision of the Malesian and Thai species of Sauropus (Euphorbiaceae: Phyllanthoideae). Blumea 48: 319–391.

    Welzen, P.C. van & Pruesapan, K. 2010. Four new species of Breynia (Phyllanthaceae / Euphorbiaceae sensu lato) and one new combination from Thailand and Malaysia. Thai Forest Bulletin (Botany) 38: 111–119.

    Welzen, P.C. van, Pruesapan, K., Telford, I.R.H., Esser, H.-J. & Bruhl, J.J. 2014. Phylogenetic reconstruction promps taxonomic changes in Sauropus, Synostemon and Breynia (Phyllanthaceae tribe Phyllanteae). Blumea 59: 77–94.

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Genus description

Key to the Malesian species (section Cryptogynium and subgenus Sauropus)

Key to the Philippine and Lesser Sunda species (subgenus Sauropus)

Species descriptions

Breynia J.R.Forst. & G.Forst.

    Breynia J.R.Forst. & G.Forst., Char. Gen. Pl. (1775) 73, nom. cons. (non Breynia L., Sp. Pl., 1753: 503, nom. rej., Capparaceae); Müll.Arg. in DC., Prodr. 15, 2 (1866) 438; Airy Shaw, Kew Bull. 26 (1972) 224; Whitmore, Tree Fl. Malaya 2 (1973) 73; G.L. Webster, Ann. Missouri Bot. Gard. 81 (1994) 46; Chakrab. & Gangop., J. Econ. Tax. Bot. 20 (1996) 501; Welzen, Thai For. Bull. 28 (2000) 55, Fig. 1; Welzen & Esser in Welzen et al., Thai For. Bull. 28 (2000) 74; Radcl.-Sm., Gen. Euphorbiacearum (20010 46; Welzen & Esser in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 132; G.L.Webster in Kubitzki, Fam. Gen. Vasc. Pl. 11 (2014) 80; Welzen et al., Blumea 59 (2014) 89. — Type: Breynia disticha J.R.Forst. & G.Forst.

    Melanthesa Blume, Bijdr. 12 (1826) 590. — Lectotype (designated by Webster, Ann. Missouri Bot. Gard. 81, 1994: 46): Melanthesa racemosa Blume (= Breynia racemosa (Blume) Müll.Arg.).

    Melanthesopsis Müll.Arg., Linnaea 32 (1863) 74; in DC., Prodr. 15, 2 (1866) 436. — Lectoype (designated by Wheeler, Taxon 24, 1975: 537): Melanthesopsis lucens (Poir.) Müll.Arg. (= Breynia fruticosa (L.) Hook.f.).

Herbs to trees, monoecious; stem often with 4 raised ribs. Indumentum of simple hairs or absent, often asperities (stiff papillae) present on stem, petioles or leaf margins. Stipules triangular, (early to) late caducous. Leaves distichous, on side branches, simple; petiole usually short; blade symmetric, basally attached, margin entire, surfaces without glands, drying greenish to brownish to blackish above, greenish to dirty brownish and sometimes glaucous-papillate beneath; venation usually indistinct, pinnate, nerves looped and closed near margin, veins reticulate. Inflorescences ramiflorous or cauliflorous racemes to usually axillary fascicles of single to several bracteate flowers, all pendent, pistillate at the top of branches, staminate at lower nodes, the latter often in short bracteate racemes with a few flowers apically and often appearing before the pistillate flowers. Flowers actinomorphic, pedicellate, hanging down; calyx disk-shaped to urceolate, 6-lobed, imbricate; petals, disc, and pistillode absent. Staminate flowers: calyx thin to very thick, lobes minute to distinct, with internal scale (transformed disk gland?) at lobe insertion, closing flowers when pollen not ripe, in some species 3 lobes apically inflexed and grown with adaxial midrib, then scales lacking; stamens 3, united into central vertical androphore, which either splits 3 ways, horizontal or ascending (group without scales) and anthers underneath or androphore not split and anthers longitudinally along it, anthers 2-thecate. Pistillate flowers: calyx persistent, lobes apically mucronate; ovary bell-shaped, 3-locular, apically flat to emarginate, outer margin with or without lobes, inside with 3 simple or split stigmas, in one part of genus these horizontal and crescent moon-like; ovules 2 per locule. Fruits rhegmas/capsules, ovoid, not lobed, tardily dehiscent, smooth, glabrous but sometimes papillate at apex, usually thin-walled and woody when dry to fleshy and inflated in a few species, yellow to red. Seeds sharply trigonous in transverse section, smooth, without or with yellow to reddish sarcotesta.

    Distribution — Genus of 50 or more species, ranging from India and Sri Lanka to S. China, S.E. Asia (main diversity), Malesia, to Australia. 

    Note— The term asperities is used for a kind of stiff papillae, present on the leaves or stems in some species.

Key to the Malesian species of former Sauropus (excl. Breynia s.s.)

(For the Philippines and the Lesser Sunda Islands see also next key)

 (Mal. = Peninsular Malaysia, Sum. = Sumatra, Jav. = Java, Bor. = Borneo, Phil. = Philippines, Sul. = Sulawesi, LSI. = Lesser Sunda Islands, Mol. = Moluccas, NG. = New Guinea.)

Key to the species of the  Philippines and the Lesser Sunda Islands

(subgenus Sauropus)

Breynia J.R.Forst. & G.Forst. Sugenus Breynia

    Breynia J.R.Forst. & G.Forst. Subgen. Breynia: Literature and type as under the genus.

Contains Section Breynia and Section Cryptogynium.

Breynia J.R.Forst. & G.Forst. Section Breynia

    Breynia J.R.Forst. & G.Forst. Section Breynia: Literature and type as under the genus.

    Breynia J.R.Forst. & G.Forst. Section Breyniastrum Baill., Adansonia 6 (1866) 344. — Lectotype (designated by Esser in Welzen et al., Blumea 59, 2014: 89): Breynia stipitata Müll.Arg.

All species of Breynia s.str. (Breynia in the old sense) belong in this type section. These need no new combinations, and the Malesian species are still under revision by Esser & Stuppy. A checklist of accepted names under Breynia in the old sense can be found in Govaerts et al. (World Checkl. Bibliogr. Euphorb. 1, 2000: 278–284).

Breynia coronata Hook.f.

Breynia discigera Müll.Arg.

Breynia glauca Craib

Breynia novoguineensis Esser & Stuppy

Breynia orbicularis Esser & Stuppy

Breynia racemosa (Blume) Müll.Arg.

    Breynia racemosa (Blume) Müll.Arg. in DC., Prodr. 15, 2 (1866) 441. — Melanthesa racemosa Blume, Bijdr. 12 (1826) 492. — Phyllanthus racemifer Steud., Nomencl. Bot. ed. 2, 2 (1841) 327, nom. nov. — Phyllanthus racemosus (Blume) Chakrab. & N.P.Balakr., Bangladesh J. Pl. Taxon. 19 (2009) 712. — Lectotype (designated by Esser in Welzen et al., Blumea 59, 2014: 89): Blume 1036 (lectotype L, sheet no. 903.155-99; isolectotypesA, L), (Indonesia), Java, ‘prope Parong provinciae Buitenzorg’.

Breynia retusa (Dennst.) Alston

    Breynia retusa (Dennst.) Alston

    Sauropus elegantissimus Ridl., Bull. Misc. Inform. (1926) 476; Whitmore, Tree Fl. Malaya 2 (1973) 130. — Type: Hume 9366 (SING), Malaysia, Selangor, Ulu Gombak.

Breynia stipitata Müll.Arg.

Breynia vitis-idaea (Burm.f.) C.E.C.Fisch.

    Breynia vitis-idaea (Burm.f.) C.E.C.Fisch., Bull. Misc. Inform. (1932) 65. — Rhamnus vitis-idaea Burm.f., Fl. Ind. (1768) 61, p.p., only lectotype. — Phyllanthus rhamnoides Retz., Observ. Bot. 5 (1788) 30, nom. illeg., nom. superfl. — Phyllanthus vitis-idaea (Burm.f.) Chakrab. & N.P.Balakr., Bangladesh J. Pl. Taxon. 19 (2009) 712. — Lectotype (designated by Radcliffe-Smith in Nasir & Ali, FL. Pakistan 172, 1986: 14): Breyne, Exot. Pl. Cent.: 8, t. 4. 1678.

Breynia J.R.Forst. & G.Forst. Section Cryptogynium (Müll.Arg.) Welzen & Pruesapan

    Breynia J.R.Forst. & G.Forst. Section Cryptogynium (Müll.Arg.) Welzen & Pruesapan in Welzen et al., Blumea 59 (2014) 89. — Sauropus Blume Section Cryptogynium Müll.Arg., Linnaea 32 (1863) 73;  in DC., Prodr. 15, 2 (1866) 243, as ‘Ceratogynum’; Hook.f., Fl. Brit. India 5 (1887) 334, as ‘Ceratogynum’; Pax&K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 222, as ‘Ceratogynum’; Airy Shaw, Kew Bull. 23 (1969) 43. — Type: Sauropus rigidus Thwaites (= Breynia quadrangularis (Willd.) Chakrab. & N.P.Balakr.) (referred originally to Wight’s genus name Cryptogynum, but as the oldest available name in its rank is - erroneously Cryptogynium, it must be maintained; see Airy Shaw 1969).

    Ceratogynum Wight, Icon. Pl. Ind. Orient. 5 (1852) 26. — Type: Ceratogynum rhamnoides Wight (= Breynia quadrangularis (Willd.) Chakrab. & N.P.Balakr.).

    Sauropus Blume Section Hemisauropus Müll.Arg. in DC., Prodr. 15, 2 (1866) 243; Airy Shaw, Kew Bull. 23 (1969) 55. — Sauropus Blume Subgenus Hemisauropus (Müll.Arg.) Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 225. — Type: Sauropus rostratus Miq. (= Breynia temii (Welzen & Chayam.) Welzen & Pruesapan).

    Breyniopsis Beille, Bull. Soc. Bot. France 72 (1925) 157; in Lecomte, Fl. Indo-Chine 5 (1927) 630. — Type: Breyniopsis pierrei Beille (= Breynia pierrei (Beille) Welzen & Pruesapan).

Staminate flowers disc-like; calyx thin to very thick, lobes minute to distinct, with internal scale at lobe insertion, in some species 3 lobes apically inflexed and grown with adaxial midrib, then scales lacking; stamens 3, united into central vertical androphore, apically split 3 ways, horizontal or ascending (group without scales) and anthers underneath. Pistillate flowers: calyx persistent, lobes apically mucronate; ovary bell-shaped, 3-locular, apically flat with rim along outer margin, inside with 3 split stigmas, these horizontal or upright (group without scales); ovules 2 per locule. 

xx. Breynia brevipes (Müll.Arg.) Chakrab. & N.P.Balakr.

    Breynia brevipes (Müll.Arg.) Chakrab. & N.P.Balakr., Bangladesh J. Pl. Taxon. 19 (2012) 121; Welzen et al., Blumea 59 (2014) 89. — Sauropus brevipes Müll.Arg., Linnaea 32 (1863) 73; in DC., Prodr. 15, 2 (1866) 242; Hook.f., Fl. Br. India 5 (1887) 335; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 222; Beille in Lecomte, Fl. Indo-Chine 5 (1927) 653; Airy Shaw, Kew Bull. 23 (1969) 44; Kew Bull. 26 (1972) 334; Whitmore, Tree Fl. Malaya 2 (1973) 130; Chakrab. & M.G.Gangop., J. Econ. Tax. Bot. 20 (1996) 524, Fig. 3; Smitinand, Thai Pl. Names, rev. ed. (2001) 467; Welzen, Blumea 48 (2003) 348, map 7; in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 532. —Aalius brevipes (Müll.Arg.) Kuntze, Revis. Gen. Pl. (1891) 591. — Phyllanthus myanmarensis Chakrab. & N.P.Balakr., J. Econ. Taxon. Bot. 33 (2009) 715. — Type: Wallich 23? (holo G-DC, n.v., 2 sheets, IDC microfiche DC herbarium 2461/12, 13), India, Prome.

    Sauropus parvifolius Ridl., J. Straits Branch Roy. Asiat. Soc. 49 (1911) 175; Fl. Malay Penins. 3 (1924) 221; M.R.Hend., J. Malayan Branch Roy. Asiat. Soc. 17 (1939) 72. — Type: Ridley 15187 (holo K; iso BM), Malaysia, Kedah, Alor Sta.

Woody herbs to shrubs, up to 1.5 m tall, often smelling of fenugreek when dry; branches with 2 relatively broad ribs, often with asperities between the ribs, glabrescent; flowering branches c. 0.5 mm thick. Indumentum absent. Stipules triangular to basally eared and falcate, 0.5–3.1 by 0.4–1.5 mm, membraneous, persistent. Leaves: petiole 1.3–2.5 mm long, flattened above to squarish in transverse section; blade (ovate to elliptic to) obovate, 0.9–3.3 by 0.6–2.3 cm, length/width ratio 1.1–2.9, papery to chartaceous, base (emarginate to truncate to) cuneate, margin lat (to revolute), apex rounded to acuminate, mucronulate, upper surface dark green, lower surface light dull green, papillate; venation mainly well visible beneath, nerves 6–8 per side, not raised, veins and veinlets reticulate. Inflorescences axillary fascicles, usually single or few staminate ones together, pale green to bright red. Staminate flowers 2–5 mm in diameter; pedicel 2.5–6.2 mm long; calyx 1–2.7 mm deep, lobes 0.5–1.2 by 0.7–2.5 mm, apex slightly emarginate to rounded; stamens: androphore 0.2–0.3 mm long, anthers 0.2–0.3 by 0.2–0.3 mm. Pistillate flowers 5–7(–9 in fruit) mm in diameter; pedicel 1.8–2.7 mm long; calyx lobes obovate, not spade-like to spade-like, 2.2–3.2 by 1.3–2.5 mm, 3 slightly smaller; ovary 0.6–1 by 1.5–1.8 mm; stigmas horizontal, up to 1.3 mm long, split for 0.6 mm, usually curled for less than a full circle. Fruits ovoid, c. 5 by 4 mm; column c. 4 mm long, tapering towards apex. Seeds triangular in transverse section, c. 3.7 by 2.3 by 2.3 mm.

    Distribution — Myanmar, Thailand, Cambodia, N. Peninsular Malaysia (Perlis, Kedah, Penang).

    Habitat & Ecology — Mixed dry dipterocarp forest, decidous forest, bamoo-evergreen forest, evergreen thickets, by streams, forest edges, sandy shores; soil: limestone, sand, red clay, granite. Altitude: 0–700 m. Flowering: February, May to September, November; fruiting: February, May.

    Vernacular name — Thailand: Kraduk kai dam (official Thai name); kham kao.

    Notes — Extremely similar to B. quadrangularis, but differing in smell of fenugreek (though this smell disappears with age of the specimen), the apically hardly to non-lobed staminate sepals, and the horizontal stigmas. The latter is the most typical difference, especially in the north of Thailand. Also, B. brevipes often has asperities along the stem between the ribs, these are absent in the glabrous specimens of B. quadrangularis (check youngest branchlets).

xx. Breynia shawii (Welzen) Welzen & Pruesapan

    Breynia shawii (Welzen) Welzen & Pruesapan in Welzen et al., Blumea 59 (2014) 90. — Sauropus shawii Welzen, Blumea 48 (2003) 372. — Type: P.F. Stevens et al. 513 (holo L; iso A, KEP), Malaysia, Sabah, Lahad Datu, Ulu Sungei Segamat.

Shrubs up to 2.1 m high, bole up to 1.2 m high, dbh up to 2.5 cm; branches horizontal, flowering branches 2.5–3 mm thick, youngest branches with 2 ridges. Outer bark smooth, grey, soft, c. 1 mm thick; inner bark grey and yellow, soft, fibrous. Indumentum absent. Stipules triangular, basally strongly narrowing into a spike-like blade, 1.4–2 by 1–1.5 mm, rather persistent. Leaves: petiole c. 3 mm long, flattened above; blade ovate, 6.8–11.2 by 3.8–4.9 cm, length/width ratio 1.8–2.3, papery, base cuneate, strongly asymmetric, margin flat, apex acuminate to cuspidate, mucronulate, upper surface dark shiny green, lower surface paler, shortly and densely papillate; venation hardly distinct on both sides, nerves 6 or 7, veins + scalariform, veinlets indistinct. Inflorescences axillary fascicles, staminate flowers sometimes in short, up to 2 mm long, racemes. Staminate flowers c. 2.5 mm in diameter; pedicel c. 3.5 mm long; calyx lobed till halfway, obovate, 3 smaller, outer lobes c. 1 by 1–1.2 mm, apex incurved, rounded; 3 inner, larger lobes 0.7–0.8 by 1.5–1.7 mm, apex incurved and grown with blade; stamens erect, androphore c. 0.2 mm long, anthers c. 1.3 by 1 mm. Pistillate flowers unknown; pedicel in fruit 8–13 mm long; calyx cupular, 9–11 mm deep in fruit, lobes obovate, 5–6 by 5–6 mm., apex spade-like; stigmas free from each other, c. 1.2 mm long, erect, split in upper half, apices recurved for less than a full circle. Fruits ovoid, wider than high, 15–16 by 17--19 mm; wall thick, up to 2.5 mm; apically hole between stigmas. Seeds triangular in transverse section, half-moon-shaped, c. 5 by 3.5 by 3.3 mm.

    Distribution — Borneo (Sabah: Tawau & Lahad Datu Distr.).

(B. shawii: triangles; B. asymmetrica: dots; B. micrasterias: stars)

    Habitat & Ecology — Primary forest, once recorded near river. Altitude: 170–500 m. Flowering: January; fruiting: February to April.

    Notes — 1. This entity resembles B. lanceolata in leaf shape and size, but the leaves are basally asymmetric, the venation is indistinct and the lower surface is very glaucous. S. shawii clearly belongs to the Hemisauropus group, but it is unlike the other species in this group in several respects. The strongest resemblance is with S. pierrei (different in leaf size, but just as asymmetric), S. subterblancus (different in the more cuspidate apex, asymmetry, and leaf size), and B. temii (larger leaves, lack of asperities).

2. The name is in honour of H.K. Airy Shaw, who described and revised most Malesian Euphorbiaceae and who was the first to note that two SAN collections from Borneo represented something new (Airy Shaw, 1975, under S. pierrei).

3. See note under B. asymmetrica.

xx. Breynia temii (Welzen & Chayam.) Welzen & Pruesapan

    Breynia temii (Welzen & Chayam.) Welzen & Pruesapan in Welzen et al., Blumea 59 (2014) 90. — Sauropus temii Welzen & Chayam. [ex Smitinand, Thai Pl. Names, rev. ed. (July 2001) 468, nom. nud.] Kew Bull. 56 (Oct. 2001) 654. — Type: Smitinand 2877 (BKF), Thailand, Peninsular, Surat Thani, Bang Bao. See note 2.

    Sauropus rostratus Miq., Fl. Ned. Ind., eerste bijv. (1861) 179, 447, non Breynia rostrata Merr.; Müll.Arg. in DC., Prodr. 15, 2 (1866) 243; Pax & K.Hoffm. in Engl., Pflanzenr. VI.147.xv (1922) 225; Airy Shaw, Kew Bull. 23 (1969) 55; Kew Bull. 36 (1981) 343; Welzen, Blumea 48 (2003) 370, map 17; in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 546. — Aalius rostratus (Miq.) Kuntze, Revis. Gen. Pl. 2 (1891) 591 (‘rostrata’). — Lectotype (designated by Welzen, 2003): Teysmann HB 3678 (holo U), Sumatra, Palembang Prov., River Lamatang near Koeripan (Kuripan).

Shrubs, up to at least 60 cm tall; young branches with 2 broad rims and sometimes 2 additional vague ones, many asperities between and on rims, flowering branches 0.8–1 mm thick. Indumentum absent. Stipules triangular, 1–2.2 by 0.7–1.2 mm, often basally eared, stiff, rather persistent. Leaves: petiole 1.3–2 mm long, with asperities, somewhat flattened above; blade ovate (to elliptic), 2.9–7 by 0.9–2.6 cm, length/width ratio 1.9–3.2, chartaceous, base broadly cuneate, margin flat to revolute, apex gradually acute, mucronulate, drying greyish above, brownish or greenish below, not distinctly papillate, venation indistinct, midrib sharply raised above, nerves 7–9, basal one ending far below middle, veins and veinlets indistinct, reticulate. Inflorescences axillary fascicles or in short staminate racemes, less than 5 mm long; flowers single or in small groups, both sexes at same time, green. Staminate flowers 2–2.5 mm in diameter; pedicel 1.2–2 mm long; calyx lobes without scales, rather small, cupular, 3 lobes folded inwards and apex grown together with raised midrib, 0.5–0.7 by 1.2–1.5 mm, other 3 lobes with incurved apex, 0.5–0.7 by 0.3–0.7 mm; stamens: androphore c. 0.7 mm long, anthers erect, c 1 by 0.8 mm. Pistillate flowers 2.5–5(–8 in fruit) mm in diameter; pedicel 1–2 mm long; calyx cupular, hardly lobed, smaller three lobes 0.5–2 by 1–3 mm, larger three lobes 0.8–2.7 by 1.5–5.5 mm, apex spade-like; ovary bell-shaped with flat apex on which 3 rims between stigmas, 1–1.5 by 1.2–2 mm; stigmas 1–1.4 mm long, split in upper 0.5–0.6 mm, recurved. Fruit ellipsoid, wider than high, c. 6 by 3.7 mm; column broadly triangular, c. 0.5 mm high. Seeds triangular in transverse section, half-moon-shaped, c. 3.3 by 2.4 by 2.8 mm.

    Distribution — Thailand (Peninsular floristic district), Sumatra, Borneo.

    Ecology & Habitat — Scattered in evergreen forest. Altitude under 100 m. Flowering: Aug.

    Vernacular name — Thailand: Mak bang bao.

    Notes — 1. The specimens from the Lesser Sunda Islands, as mentioned by Airy Shaw (1982), are certainly not B. temii, but an atypical form of B. lanceolata. They have a very different type of staminate flower (the normal 'Sauropus' stamens: low androphore, 3 horizontally spreading branches with anthers underneath) and much larger fruits (up to 2 cm diameter).

2. Smitinand 2877 (BKF) from Thailand was originally selected as a new species. It is a very scrappy specimen but easily recognisable as a distinct entity. However, extension of the revisions of former Sauropus to the complete Malesian area showed it to be similar to S. rostratus (now a synonym of B. temii).

Breynia J.R.Forst. & G.Forst. Subgenus Sauropus (Blume) Welzen & Pruesapan

    Breynia J.R.Forst. & G.Forst. Subgenus Sauropus (Blume) Welzen & Pruesapan in Welzen et al., Blumea 59 (2014) 91. — Sauropus Blume, Bijdr. 12 (1826) 595; Müll.Arg. Linnaea 32 (1863) 72; in DC., Prodr. 15, 2 (1866) 239; Hook.f., Fl. Brit. India 5 (1887) 332; Benth. & Hook.f., Gen. Pl. 3 (1880) 271; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 215; Backer & Bakh.f., Fl. Java 1 (1963) 471; Airy Shaw, Kew Bull. 23 (1969) 42; Kew Bull. 26 (1972) 330; Whitmore, Tree Fl. Malaya 2 (1973) 130; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 190; Kew Bull. Add. Ser. 8 (1980a) 199, 221; Kew Bull. 35 (1980b) 669; Kew Bull. 36 (1981) 342; Kew Bull. 37 (1982) 34; Alphab. Enum. Euph. Philip. Is. (1983) 44; G.L.Webster, Ann. Missouri Bot. Gard. 81 (1994) 46; Welzen, Thai For. Bull. 28 (2000) 57, Fig. 10; Radcl.-Sm., Gen. Euphorbiacearum (2001) 30; Welzen, Blumea 48 (2003) 331; in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 521; G.L.Webster in Kubitzki, Fam. Gen. Vasc. Pl. 11 (2014) 79. — Aalius Rumph. [Herb. Amboin. (1743) 207; Lam., Encycl. Méth. Bot. 1, 1 (1783) 1, nom. inval., Art. 32.1(d).] ex Kuntze, Rev. Gen. Pl. 2 (1891) 590, nom. illeg., nom. superfl. — Sauropus Blume Section Eusauropus Müll.Arg., Linnaea 32 (1863) 72; in DC., Prodr. 15, 2 (1866) 240; Hook.f., Fl. Br. India 5 (1887) 332, nom. inval., Art. 21.3. — Sauropus Blume Subgenus Holosauropus Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 216, nom. inval., Art. 22.2. — Lectotype (designated by G.L.Webster, 1994): Sauropus albicans Blume [= Breynia androgyna (L.) Chakrab. & N.P.Balakr.]

    Sauropus Blume Section Sphaeranthi Pax & K. Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 220. — Type species: Sauropus stipitatus Hook.f. [= Breynia gynophora Welzen & Pruesapan].

    Sauropus Blume Section Retroversi Pax & K. Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 221. — Type: Sauropus retroversus Wight [= Breynia androgyna (L.) Chakrab. & N.P.Balakr.].

    Sauropus Blume Section Schizanthi Pax & K. Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 221. — Lectotype (designated by Welzen, 2003): Sauropus trinervius Wall. ex Müll.Arg. [= Breynia trinervia (Wall. ex Müll.Arg.) Welzen & Pruesapan].

    Sauropus Blume Section Glochidioidei Airy Shaw, Kew Bull. 23 (1969) 51. — Type: Sauropus villosus (Blanco) Merr. [= Breynia villosa (Blanco) Welzen & Pruesapan].

Staminate flowers disc-like; calyx thin to very thick, lobes minute to distinct, with internal scale at lobe insertion; stamens 3, united into central vertical androphore, apically split 3 ways, horizontal with anthers underneath. Pistillate flowers: calyx persistent, lobes apically mucronate; ovary bell-shaped, 3-locular, apically flat without rim along outer margin, inside with 3 split stigmas, these horizontal; ovules 2 per locule. 

xx. Breynia androgyna (L.) Merr.

    Breynia androgyna (L.) Chakrab. & N.P.Balakr., Bangladesh J. Pl. Taxon. 19 (2012) 120; Welzen et al., Blumea 59 (2014) 91. — Clutia androgyna L., Mant. Pl. 1 (1767) 128. — Aalius androgynus (L.) Kuntze, Revis. Gen. Pl. (1891) 591 (‘androgyna’). — Sauropus androgynus (L.) Merr., Bull. Bur. Forest. Philipp. Is. 1 (1903) 30; J. Straits Branch Roy. Asiat. Soc. 76 (1917b) 92; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 217; Merr., Enum. Philipp. Fl. Pl. 2 (1923) 405; Gamble, Fl. Madras 2 (1925) 1303; S.Moore, J. Bot. 63, Suppl. (1925) 93; Beille in Lecomte, Fl. Indo-Chine 5 (1927) 645; Backer & Bakh.f., Fl. Java 1 (1963) 471; Airy Shaw, Kew Bull. 26 (1972) 333; Whitmore, Tree Fl. Malaya 2 (1973) 130; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 193; Kew Bull. Add. Ser. 8 (1980a) 199; Kew Bull. 36 (1981) 342; Kew Bull. 37 (1982) 34; Alph. Enum. Euph. Philipp. Isl. (1983) 44; Polunin, Pl. Fl. Singapore (1987) 150, Fig. 153; Pl. Fl. Malaysia (1988) 138, Fig. 138; Levang & Foresta, Econ. Pl. Indonesia (1991) 66; Du Puy & Telford, Fl. Australia 50 (1993) 263; I.M.Turner, Gard. Bull. Singapore 45 (1993) 87; Kapil & Bhatnagar, Ann. Missouri Bot. Gard. 81: 145; M.H.Bergh in Siemonsma & Piluek, PROSEA 8, vegetables (1994) 244; Wightman, Astuti & Munawaroh, Northern Territory Bot. Bull. 19 (1994) 15; Chakrab. & M.G.Gangop., J. Econ. Tax. Bot. 20 (1996) 519, Fig. 1: g–j; C.Q.Lin et al., J. Trop. Subtrop. Bot. 7 (1999) 255, 256; Philcox, Rev. Handb. Fl. Ceylon 13 (1999) 102; Hung et al., Ann. Nucl. Med. Sc. 13: 197; Smitinand, Thai Pl. Names, rev. ed. (2001) 466; Welzen, Blumea 48 (2003) 340, fig. 1a, map 3; in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 527, Fig. 72a. — Phyllanthus androgynus (L.) Chakrab. & N.P.Balakr., J. Econ. Taxon. Bot. 33 (2009) 714. — Lectotype (designated by Welzen, 2003): Hb. Linnaeus 1206.14 (holo LINN).

    Sauropus albicans Blume, Bijdr. (1825) 596; Hassk., Retzia 1 (1855) 162; Müll.Arg. in DC., Prodr. 15, 2 (1866) 240; Kurz, Forest Fl. Burma 2 (1877) 349; Hook.f., Fl. Br. India 5 (1887) 332; Ridl., J. Straits Branch Roy. Asiat. Soc. 59 (1911) 176; Craib, Bull. Misc. Inform. (Dec. 1911) 457; Ridl., Fl. Malay Penins. 3 (1924) 220. — Sauropus albicans Blume var. genuinus Müll.Arg. in DC., Prodr. 15, 2 (1866) 241, nom. inval. — Lectotype (designated by Welzen, 2003): Blume s.n. (holo L, barcode L  0138130), Indonesia, Java, Bantam.

    Phyllanthus strictus Roxb., Fl. Ind. 3, ed. 1832 (1832) 670. — Syntypes: Roxburgh Icones pl. 1685 (K); Roxburgh s.n. (Hb. Forsyth) (K), India; Wallich 7933A (K-W), Wallich 7933B (G-DC, K-W, K); Wallich 7933E (K-W, K), India, Sillet.

    Sauropus retroversus Wight, Icon. Pl. Ind. Orient. 6 (1853) t. 1951 (left); Thwaites, Enum. Pl. Zeyl. (1864) 284; Müll.Arg. in DC., Prodr. 15, 2 (1866) 241; Hook.f., Fl. Br. India 5 (1887) 333; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 221; Beille in Lecomte, Fl. Indo-Chine 5 (1927) 651; Chakrab. & M.G.Gangop., J. Econ. Tax. Bot. 20 (1996) 537, Fig. 9: e–h; Philcox, Rev. Handb. Fl. Ceylon 13 (1999) 104. — Aalius retroversus (Wight) Kuntze, Revis. Gen. Pl. (1891) 591 (‘retroversa’). — Phyllanthus retroversus (Wight) Chakrab. & N.P.Balakr., J. Econ. Taxon. Bot. 33 (2009) 715. — Breynia retroversa (Wight) Chakrab. & N.P.Balakr., Bangladesh J. Pl. Taxon. 19 (2012) 121. — Syntypes: Anonymous s.n. (K), Sri Lanka; Thwaites CP 3134 (G-DC, K), Sri Lanka, Central Prov., Oodoopussalawa (Thwaites, 1864); Walker 274 (K), Sri Lanka. 

    Sauropus gardnerianus Wight, Icon. Pl. Ind. Orient. 6 (1853) t. 1951 (right); Thwaites, Enum. Pl. Zeyl. (1864) 284. — Sauropus albicans Blume var. gardnerianus (Wight) Müll.Arg., Linnaea 32 (1863) 72; in DC., Prodr. 15, 2 (1866) 241. — Type: Gardner 472 (n.v.), Sri Lanka (Ceylon), Hantane.

    Sauropus zeylanicus Wight, Icon. Pl. Ind. Orient. 6 (1853) t. 1952 (left). — Sauropus albicans Blume var. zeylanicus (Wight) Müll.Arg. in DC., Prodr. 15, 2 (1866) 241. — Type: R. Wight s.n. (n.v.), Sri Lanka (Ceylon). (A possible syntype, Gardner s.n. in K).

    Sauropus indicus Wight, Icon. Pl. Ind. Orient. 6 (1853) t. 1952 (right); Hassk., Retzia 1 (1855) 164; Miq., Fl. Ned. Ind. I, 2 (1859) 366. — Type: Wight s.n. (n.v.), India, Courtallum and Shevagherry Hills.

    Agyneia ovata Miq., Fl. Ned. Ind. I, 2 (1859) 367. — Type: Zollinger s.n. (U, holo, barcode U  0002081), Indonesia, Java.

    Sauropus assimilis Thwaites, Enum. Pl. Zeyl. 4 (1861) 284. — Phyllanthus assimilis (Thwaites) Chakrab. & N.P.Balakr., J. Econ. Taxon. Bot. 33 (2009) 715. — Breynia assimilis (Thawits) Chakrab. & N.P.Balakr., Bangladesh J. Pl. Taxon. 19 (2012) 120. — Type: Thwaites CP 2855 (holotype K), Sri Lanka (Ceylon), Central Prov., Allagalla. 

    Sauropus sumatranus Miq., Fl. Ned. Ind., eerste bijv. (1861) 179, 446; Ridl., Fl. Malay Penins. 3 (1924) 220. — Aalius sumatranus (Miq.) Kuntze, Revis. Gen. Pl. (1891) 591 (‘sumatrana’). — Type: Teysmann s.n. (holo U, barcode U  0002082; iso A, G-DC), Indonesia, Sumatra, Rau Distr. (= Riau Prov.).

    Sauropus albicans Blume var. intermedius Müll.Arg., Linnaea 32 (1863) 72; in DC., Prodr. 15, 2 (1866) 241. — Type: Wallich 273? (holo G-DC, n.v.; IDC microfiche DC herbarium 2461-5), India, Prome.

    Sauropus oblongifolius Hook.f., Fl. Br. India 5 (1887) 333. — Syntypes: Griffith KD 4824 (K), India, East Bengal, Mishmee; Masters s.n. (n.v.), India, Upper Assam, Dailoon in the Mishmi Hills. 

    Andrachne sp.: Merr., Philipp. J. Sci. 1, Suppl. (1906) 74. — Sauropus scandens C.B.Rob., Philipp. J. Sci. 4 (1909) 72; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 224; Merr., Enum. Philipp. Fl. Pl. 2 (1923) 405; Airy Shaw, Alph. Enum. Euph. Philipp. Isl. (1983) 44. — Type: FB (Borden) 1934 (holo PNH, †; iso K, NY, US), Philippines, Luzon, Bataan Prov., Mt. Mariveles, Lamao River.

    Sauropus parviflorus Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 218; Beille in Lecomte, Fl. Indo-Chine 5 (1927) 646. — Type: Warburg 5057 (holo B, †), Thailand (Siam), Bangkok.

    Sauropus convexus J.J.Sm., Bull. Jard. Bot. Buitenzorg ser. 3, 6: (1924) 82. — Neotype (designated by Welzen & Pruesapan, 2014): Hortus Bogoriensis s.n., cultivated (L, barcode L0138208), Indonesia, Java, Bogor. Former syntypes were based on living collections of which there are no vouchers: Indonesia, Java, Bogor, Hortus Bogoriensis XV.J.B.IV.1 (died 1950) and XV.J.B.V.5 (died 1945), originally from Leiden Botanical Garden. Smith indicates that there are minor differences in the staminate and pistillate flowers with B. androgyna, which fall well within the variation of this species.

    Sauropus macranthus auct. non Hassk.: Fern.-Vill., Nov. App. (1883) 187 (pro Vidal s.n., Philippines, Luzon, Tayabas).

    Sauropus sp., ? aff. S. stipitato Hook.f.: Airy Shaw, Kew Bull. 26 (1972) 339, p.p. [pro Kerr 8522 (BK, K), Thailand (Siam), Nong Kai (Nawng Kai), Chaiyaburi]. See note 3.

Herbs to shrubs (to treelets), up to 4 m high, dbh up to 3 cm; young branches with 2 or 4 ribs, green, flowering branches 1–2 mm thick. Indumentum absent. Bark light brown to grey. Stipules triangular, 1.8–3.2 by 0.8–1.3 mm, basally somewhat eared, usually thin, usually caducous. Leaves: petiole 2–3 mm long, flattened above; blade ovate (to elliptic), 1.8–9.5 by 0.6–4 cm, length/width ratio 1.5–3.8, papery (to pergamentaceous), base rounded to truncate, margin flat to recurved, apex (round to bluntly) acute, at most with slight sinuses below the apex, often mucronulate, dark green above, (grey-)light green and often somewhat papillate underneath; nerves usually distinct on both sides, nerves 6–10, veins somewhat scalariform, veinlets reticulate. Inflorescences axillary fascicles (or flowers in up to 8 mm long racemes), flowers single or few together, usually all of same sex. Staminate flowers 2.5–6(–20) mm in diameter; pedicel 4.5–13 mm long, green; calyx flat, (small and somewhat lobed to) wide, reflexed and hardly lobed, light yellowish green to maroon outside, maroon inside, lobes indistinct to ovate, 0.7–3 by 1.3–5 mm; stamens: androphore 0.1–0.3 mm long, cream, stamens 0.4–0.6 by 0.4–0.6 mm. Pistillate flowers 4.5–10 mm in diameter; pedicel up to 3.2–11(–14 in fruit) mm long, apically up to 1 mm thick, green; calyx lobes usually obovate, green with maroon to maroon outside, dark maroon inside, smaller ones 1.8–3.5(–4 in fruit) by 1.8–3.8(–5.5 in fruit) mm, larger ones 2.2–5.5 by 2–7 mm; ovary 1.1–1.5 by 1–2 mm; stigmas up to 1.2 mm long, light yellow (with maroon), horizontal, split till halfway, bend, forming less than a circle. Fruits globose (to obovoid), inflated, somewhat fleshy but pergamentaceous when dry, 12–17 by 9–15 mm, white to finally maroon; column 8–10 mm long with apically heart-shaped remnants of the septae. Seeds triangular in transverse section, hollow adaxially, 7–10 by 4.5–6.5 by 3–4.5 mm, white to black.

    Distribution — India, Sri Lanka, Bangladesh?, Myanmar, Laos, Cambodia, Viet Nam, Thailand, Peninsular Malaysia, Singapore, Sumatra, Java, Cocos Islands, Christmas Island (Indian Ocean), Borneo, Philippines, Sulawesi, Lesser Sunda Islands, Moluccas, New Guinea (Irian Jaya).

    Habitat & Ecology — Monsoon (deciduous) forest, secondary forest, swamp forest, open areas and clearings, in scrubs and thickets, waste grounds, hedges, fruit gardens, gardens and house yards, along forest edges, rivers, and roads, near the beach, often a weed or cultivated; in dry to wet soil: shale, (coral) limestone, sand, weathered phyllite, andesite, sandy loam, granite. Altitude: 0–1500 m. Flowering and fruiting whole year through, though less fruiting in January to March.

    Uses — Young shoots, leaves, but also flowers and fruits are eaten, raw, cooked or in the soup, as a vegetable throughout the region; they taste sweet and have a nice or very typical odour. The plant is often cultivated in small fields, as hedges, or allowed as weed near the house. Stripped leaves and young shoots are sold in the market. Dried and crushed root is used medicinally in Chiang Mai (Thailand) against head ache, but seemingly also acts against fever or urinary problems; the leaves are thought to stimulate milk production and recover the womb after child birth. In Java the leaves are also used to treat wounds and against colds (Harini 35). A green dye, used in foo are used as a green dye in food products. (Partly after van den Bergh, 1994). According to Levang & Foresta (1991) the plant is used as a vegetable, fruit, medicinally, as dye, and ornamentally.

    Chemical compounds — Lin et al. (1999) report that the major chemical essential oil constituents of the leaves are: Carvacrol methyl ether (49.35 %), Thymol (14.67 %) and Butylated hydroxytoluene (10.5 %); all other oil compounds are usually less than 2 %. The plant is exceptionally rich in several vitamins and contains the alkaloid Papaverine in harmless quantities (Polunin, 1987, 1988). Hung et al. (2000) relate that in Taiwan B. androgyna is used to slim down and then the amounts of Papaverine become too high and cause Bronchiolitis Obliterans (a fibrosing process of the lungs).

    Vernacular names — Myanmar: Yaung-mapywet; yo-ma-hin-yo (Kurz, 1877). Laos: Pahk wan. Thailand: Cha phak wan, kan tong, ma yom pa, phak kan thong, phak wan, phak wan ban (official Thai name), phak wan tai bai; tho-lui-ka-ne-doh (Karen); na-na siam (Malay). Vietnam: Bu lot, cây bu ngot, cây chum ngot, cây cu de, cây dau ngot, cu ve, dom nghob, ran n’got (Beille, 1927). Peninsular Malaysia: Assin-assin, chekop, chekop manis, chekor manis, kasim, kencur manis; thavasi murunggai (Tamil) (partly after Ridley, 1924). Indonesia: cekop manis, katu, katuk, memata (Levang & Foresta, 1991). Sumatra: Bait memata oetan, daun katu, lakioi, nasi-nasi, simanie, si topoe manoek; katoe (Akar, Malay). Java: Boemo, kera kuhr, katoek/katuk; babing, katukan, katu (Javanese; Levang & Foresta, 1991); daun katuk, katuk (Sundanese; Levang & Foresta, 1991), katuk utan (Sundanese, Djasilin dialect). Borneo: Kalimantan Timur: Daun katuk; Sabah: Chang kok manis (Malay); sayor manis (Dusun). Philippines: Tawitawi (Basilan Island). Moluccas: Katoek.

    Notes — 1. Breynia androgyna is a widespread, often cultivated species, and, probably therefore, very variable. Usually the leaves are rather small, less than 5 cm long, but many exceptions exist; the apex usually gradually narrows into an acute acumen, the staminate flowers are usually hardly lobed (though often somewhat lobed in N. Thailand and Laos, and exceptionally in other areas), the pistillate sepals are somewhat dimorph, and the fruits are inflated and somewhat fleshy.

2. This species is difficult to distinguish from B. lanceolata and B. macrantha. These two species usually have acuminate leaves with very distinct sinuses near the apex, while S. androgynus has acute leaves, gradually narrowing towards the apex without distinct sinuses. However, S. androgynus may have slight sinuses and Philippine B. lanceolata may almost lack the sinuses. Breynia macrantha has larger leaves, longer and more persistent stipules, and different staminate flowers. Compared with S. androgynus, B. lanceolata usually has (much) larger leaves (big overlap), wider staminate flowers (overlap, especially due to some exceptional specimens of B. androgyna in Thailand and Borneo), pistillate sepals usually larger and distinctly dimorph, and larger fruits.

3. S. spec. (Airy Shaw, 1972) is referred to B. androgyna, because staminate flowers with distinct, incurved lobes are often encountered.

xx. Breynia asymmetrica (Welzen) Welzen & Pruesapan

    Breynia asymmetrica (Welzen) Welzen & Pruesapan in Welzen et al., Blumea 59 (2014) 91. — Sauropus asymmetricus Welzen, Blumea 48 (2003) 344, map 4. — Type: Yates 1241 (holo BM, barcode BM000606476), Indonesia, Sumatra.

Small shrub; young branches with 2 faint ribs, flowering branches 1.5–2.5 mm thick. Indumentum absent. Stipules triangular, 3–6 by 1–2.2 mm, basally eared, rather stiff and persistent. Leaves: petiole c. 4 mm long, above channeled; blade ovate, 5.1–13.5 by 3.6–5.2 cm, length/width ratio 1.4–2.6, papery, base assymetric, rounded to very broadly cuneate, margin flat, apex acuminate, mucronulate, upper surface drying dark green, lower surface drying greyish green, densely papillate; venation indistinct on both sides, hardly raised beneath, nerves 8–10 per side, veins and veinlets indistinct. Inflorescences axillary fascicles (to short racemes of less than 1 cm long), flowers single or few staminate ones together. Staminate flowers c. 1.5 mm in diameter; pedicel c. 2.5 mm long; calyx c. 1.2 mm deep, smaller lobes c. 0.8 by 0.6 mm, apex incurved, acute, larger lobes c. 0.8 by 0.7 mm, apex further incurved, rounded; stamens: androphore 0.3–0.4 mm long, anthers c. 0.3 by 0.5 mm. Pistillate flowers unknown; fruiting pedicel c. 10 mm long; calyx lobes c. 1.5 by 1.2 mm. Fruits subglobose, 16–19 by 16–17 mm, wall rather thin, somewhat corky, apically with small hole between stigmas; column not seen. Seed triangular in cross section, c. 13 by 6 by 6 mm.

    Distribution — Sumatra.

(B. asymmetrica: dots; B. micrasterias: stars; B. shawii: triangles)

    Habitat & Ecology — Primary hill forest and secondary forest; soil: limestone, ‘red’ soil. Altitude: 500 m. Flowering: December; fruiting: October to May.

    Vernacular names — Sumatra: Kajoe ira.

    Notes — This species resembles B. shawii from Borneo and at first sight seems to be part also of the former Hemisauropus group, because the staminate calyx lobes are inflexed. However, the apices of three staminate calyx lobes are not grown with the blade as in the former Hemisauropus group, nor are the stamens large and more or less erect as in the former Hemisauropus group, the stamens show the usual 'Sauropus' type with horizontal branches with underneath the anthers. The leaves also have a somewhat different shape like those of B. shawii, both are basally asymmetric, but the leaf base of B. asymmetrica is much wider and the leaves are generally somewhat larger.

xx Breynia calcarea (M.R.Hend.) Welzen & Pruesapan

    Breynia calcarea (M.R.Hend.) Welzen & Pruesapan, Thai For. Bull., Bot. 38 (2010) 113. — Sauropus calcareus M.R.Hend., Gard. Bull. Straits Settlem.7 (1933) 121; J. Malayan Branch Roy. Asiat. Soc. 17 (1939) 72; Whitmore, Tree Fl. Malaya 2 (1973) 130; Welzen, Blumea 48 (2003) 350, map 8. — Type: SF (Henderson) 22316 (holo SING; iso K (4 sheets), KEP, NY), Malaysia, Pahang, Gunung Senyum limestone hill.

Subshrub; branches round, glabrous, outer bark of branchlets blackish, fissuring and flaking; flowering branches 0.7–0.9 mm thick. Indumentum absent. Stipules triangular, 1.5–2.3 by 0.8–1.2 mm, stiff, persistent. Leaves simple, alternate; petiole c. 1.5 mm long, round; blade ovate to round, 1.5–2.8 by 1–2.3 cm, length/width ratio 1.2–1.4, very thin, almost translucent, symmetric, base slightly peltate, emarginate to truncate, margin entire, flat, apex emarginate to subacute, upper surface drying greyish green, lower surface drying more olive green; nerves 6–8 per side, looped and closed near the margin; venation very distinct, reticulate, raised on both sides. Inflorescences axillary fascicles to very short racemes (up to 5 mm long); flowers single or a few together, red. Staminate flowers (partly after Henderson, 1933): 1.6–2 mm in diam.; pedicel up to 8 mm long, round, slightly thickening upwards; calyx consisting of 6 almost completely connate lobes, disc-like, lobes c. 0.5 by 1 mm, apex rounded; scales present; staminal column very short, anthers small. Pistillate flowers c. 3.5 mm in diam.; pedicel 2–3 m long; calyx consisting of 6 basally connate lobes, lobes c. 1.9 by 1.5 mm; ovary c. 1 mm high, top flat, without marginal rims, stigmas 3, flat on top of ovary, apex split and circling for c. 180°. Fruits depressed globose, c. 4.5 mm in diam., black when dry. Seeds straw-coloured. 

    Distribution — Endemic to the Malay Peninsula (Pahang). 

(B. calcarea: dots; B. discocalyx: stars)

   Habitat & Ecology — On limestone; ca 35 m altitude. Flowering and fruiting: July. 

    Note — This description is an amendment to that published in Van Welzen (2003). Diagnostic characters include the absence of hairs and the slightly peltate leaf blades.

xx. Breynia discocalyx (Welzen) Welzen & Pruesapan

    Breynia discoclayx (Welzen) Welzen & Pruesapan in Welzen et al., Blumea 59 (2014) 91. — Sauropus discocalyx Welzen, Blumea 46 (2001) 501, fig. 1;  Blumea 48 (2003) 351, fig. 6, map 8; in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 533, Fig. 74. — Type: van Beusekom & Phengkhlai 566 (holo L; iso AAU, BKF, C, E, K, P), Thailand, (Peninsular,) Khao Saideng, near Ranong.

    Sauropus bonii auct. non Beille: Smitinand, Thai Pl. Names, rev. ed. (2001) 467.

Shrublet with ascending habit, up to 0.5 m high; young branches with 4 distinct ribs. Indumentum absent. Stipules triangular, basally eared, 2.5–3 by 2–2.5 mm, stiff, rather persistent. Leaves: petiole 2.5–4 mm long, flattened above with 3 longitudinal (indistinct) ridges; blade (ovate to) elliptic, 11.5–17.5 by 5.8–7.2 cm, length/width ratio 2–2.3, papery, base attenuate, descending into petiole, margin flat, apex cuspidate, usually mucronulate, lower surface not papillate; venation flat to somewhat sunken above, distinct beneath, nerves 12–15, veins and veinlets reticulate. Inflorescences cauliflorous at ground level to axillary, racemes (to panicles), up to 7.5 cm long, with per node groups of staminate flowers and a single pistillate flower; flowers sometimes on short, up to 2 mm long, branches with (sub)apically the pistillate flower. Flowers reddish green. Staminate flowers 12–13 mm in diameter; pedicel 13.5–15 mm long; calyx hardly lobed, disc-like, translucent when mature; lobes very broadly triangular, 1–1.5 by c. 4.5 mm, patent, apex rounded, scales small; stamens: androphore c. 0,3 mm high, anthers c. 0.3 by 0.7 mm. Pistillate flowers c. 11 mm in diameter; pedicel c. 13 mm long; calyx almost completely split, lobes only basally attached, obovate, c. 5 by 3 mm, more or less with claw, apex rounded; ovary obtriangular, c. 2 by 2 mm, basally very narrow, green; stigmas horizontal, c. 2 mm long, split till halfway, forming more than a circle, green. Fruits red, not seen. Seeds unknown.

    Distribution — Thailand (Northern part Peninsular floristic district: Chumphon, Ranong Prov.). Perhaps also in Malaysia (Perak).

(B. discocalyx: stars; B. calcarea: dots)

    Habitat & Ecology — Wet evergreen forest. Altitude: 400–1500 m. Flowering: February, May.

    Vernacular name — Thailand: Mayom bon.

    Notes — 1. The two known specimens resemble B. beillei Welzen & Pruesapan (formerly Sauropus racemosus Beille). The type of that species (Balansa 3202, P) is quite unlike B. discocalyx in the leaves (smaller and narrower: up to 10.5 by 3.5, length/width ratio c. 3) and the staminate flowers (also broad, c. 15 mm in diameter, but very distinctly lobed, with lobes till about halfway the calyx). A second specimen from Paris, doubtfully identified as B. beillei (leaves basally rounded instead of narrowly cuneate), has pistillate flowers with short, up to 2.2 mm long, pedicels and an ovary which is basally rounded and not narrow, also the diameter of the flower is much smaller, c. 7 mm in diameter and the calyx lobes are c. 3.5 by 2 mm.

2. A third specimen from Malaysia (Goping/Gopeng in Perak, King’s collector 477) bears a strong resemblance to the Thai specimens in the leaves. The staminate flowers are still too young. The pistillate flowers are also somewhat younger.

xx. Breynia lanceolata (Hook.f.) Welzen & Pruesapan

    Breynia lanceolata (Hook.f.) Welzen & Pruesapan in Welzen et al., Blumea 59 (2014) 92. — Sauropus lanceolatus Hook.f., Fl. Br. India 5 (1887) 333. — Type: Griffith KD 4825 (K; ‘4828’ on sheet), India, East Bengal, Mishmee. (N.B. there are more sheets of Griffith KD 4828 in K, one is paratype of Sauropus macrophyllus Hook.f.).

    Sauropus rhamnoides Blume, Bijdr. (1825) 596 (non Breynia rhamnoides (Willd.) Müll.Arg.); Müll.Arg. in DC., Prodr. 15, 2 (1866) 240; J.J.Sm., Meded. Depart. Landb. Ned.-Indië 10 (1910) 191; Merr., J. Straits Branch Roy. Asiat. Soc., special numb. (1921) 329; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 219; S.Moore, J. Bot. 63, Suppl. (1925) 93; Merr., Pl. Elmer. Born. (1929) 139; Backer & Bakh., Fl. Java 1 (1963) 471; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 194; Kew Bull. 32 (1977) 81; Chakrab. & M.G.Gangop., J. Econ. Tax. Bot. 20 (1996) 539, Fig. 11; Smitinand, Thai Pl. Names, rev. ed. (2001) 468; Welzen, Blumea 48 (2003) 367, fig. 1b, c, 3a, 8, map 16; in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 546, Fig. 72b, 75a, 79. — Aalius rhamnoides (Blume) Kuntze, Revis. Gen. Pl. 2 (1891) 591 (‘rhamnodes’). — Phyllanthus rhamnifolius Chakrab. & N.P.Balakr., J. Econ. Taxon. Bot. 33 (2009) 715, non P. rhamnoides Retz. — Breynia macrocalyx Chakrab. & N.P.Balakr., Bangladesh J. Pl. Taxon. 19 (2012) 122, nom. illeg., nom. superfl. — Lectotype (designated by Welzen, 2003): Blume s.n. (L, barcode L  0138511), Indonesia, Java, Montis Salak.

    Sauropus hayatae Beille in Lecomte, Fl. Indo-Chine 5 (1927) 650. — Type: Hayata s.n. (holo P), Viet Nam, Annam, Bum-mo. 

    Sauropus rostratus auct. non Miq.: Airy Shaw, Kew Bull. 37 (1982) 35.

Herbs to (scandent) shrubs to treelets to small vines, up to 6(–10) m high, dbh up to 4(–12) cm; young branches with 2 or 4 ribs, flowering branches 0.8–2 mm thick. Indumentum absent. Stipules triangular, 2–3.5 by 0.8–1.5 mm, basally usually eared, usually thin with membranous margin, usually caducous, sometimes torn-like. Leaves: petiole 3–4 mm long, flattened and grooved along midrib above; blade ovate (to elliptic), 2.2–17.2 by 1–8.4 cm, length/width ratio 1.2–3.2, papery, base cuneate to somewhat oblique, margin flat to recurved, apex (gradually to) suddenly smaller with 2 sinuses acuminate to cuspidate (see note 2 and 3), often mucronulate, dark green above, pale light green, usually not papillate underneath; venation well-visible on both sides, nerves 7–10, veins somewhat scalariform, veinlets reticulate. Inflorescences axillary fascicles, sometimes short staminate racemes of up to 5 mm long, flowers usually single or few together, usually all of same sex. Staminate flowers 4.5–19(–25, Philippines) mm in diameter; pedicel 6.2–12 mm long, green to white; calyx not or hardly lobed and circular (to lobed and more star-shaped in SE Thailand and the Lesser Sunda Islands, or sometimes deeply lobed in the Philippines; see note 2 and 3), white to striped pinkish or reddish to reddish, lobes 1–1.6 by 1.5–2.5 mm, somewhat leathery, apex entire; stamens: androphore 0.1–0.4 mm long, anthers 0.3–0.5 by 0.3–0.4 mm, blackish. Pistillate flowers 6.5–27 mm in diameter; pedicel 7.5–20 mm long, apically 1–1.5 mm thick, very pale greenish; calyx lobes thick, only at very base attached, obovate, yellowish or light green to red, smaller inner ones (4–)6–10 by (3.3–)6–8 mm, larger outer ones (4–)7–16 by (4–)7–11 mm; pistil light yellow to whitish, ovary bell-shaped, 1.8–3.6 by 1.5–2.7 mm; stigmas 1.3–2.3 mm long, horizontal, thick, upper half curved, forming more than a complete circle. Fruits ellipsoid to obovoid, inflated, somewhat fleshy but pergamentaceous when dry, 13–31 by 13–23 mm, white to pale yellow to finally red, somewhat higher than broad, wall thin except near sutures; column 17–19 mm long, in upper ²/₃^rd with heart-shaped remnants of septae. Seeds triangular in section, hollow adaxially, c. 11 by 6–7 by 4–5.5 mm, blackish.

    Distribution: India, Myanmar, Thailand (Southeastern and Peninsular floristic districts), Cambodia, S Viet Nam, Malay Peninsula, Sumatra, Java, Borneo, Philippines, Lesser Sunda Islands (Sumbawa, Flores).

    Habitat & Ecology — Evergreen forest, mixed Dipterocarp forest, lower montane forest, secondary forest to open vegetations, along rivers or forest edges, swamp forest edges, ladang edges, also as village hedge, terrain level to hilly to mountainous; soil: rich in clay, sometimes black in colour, sometimes rocky, sometimes covered with guano, andesite; bedrock: basalt, granite, limestone. Altitude: 5–1750 m. Flowering and fruiting whole year through.

    Uses — The fruits are edible and the leaves are sometimes used as vegetable.

    Vernacular — Thailand: Mayom liam. Malay Peninsula: Asing asing hutan. Sumatra: Alor anteu anteu, katoei. Java: Djiendjieng (Javanese); katoek-badak, katoek endog, telor kotok (Sundanese) (Smith, 1910; Moore, 1925). Borneo: Kalimantan: Kabo, oroh mi(h), rangkok rimba; Sabah: Obah (Malay); Sarawak: Changkok manis (hutan) (Malay). Lesser Sunda Islands: Mintje, tago potjo (Flores).

    Notes — 1. Airy Shaw (1977) tentatively added Thailand to the distribution of B. lanceolata (formerly Sauropus rhamnoides), which was correct. All specimens of SE and Peninsular Thailand, formerly referred to B. garrettii, all appeared to be B. lanceolata. The Malay Peninsular appeared to be a gap in the distribution of B. lanceolata, but here also, the species was not recognised properly, it was mixed with B. androgyna (for differences see note under B. androgyna).

2. The specimens from SE Thailand are somewhat different in that the staminate flowers have a distinctly lobed, hardly spreading calyx. The same syndrome can be found on the Lesser Sunda Islands and more extremely so in the Philippines (see next note).

3. The Philippine specimens of B. lanceolata are very similar to those of B. androgyna (much smaller leaves), because B. lanceolata lacks its distinct subapical, laminar sinuses (apex gradually cuspidate). The same can be observed on the Lesser Sunda Islands. However, the very apex remains much shaper than in B. androgyna. The staminate flowers of B. lanceolata in the Philippines can become very large, up to 25 mm in diameter, which is distinctly different from those of B. androgyna (up to 6 mm in diameter), or they become deeply divided. The specimens with the deeply divided staminate flowers are very exceptional, they are found on Samar (BS 24500), Leyte (Wenzel 370, Wenzel 1583), and in the Camarines Sur Prov. On Luzon (Ahern 288). One Philippine specimen of B. androgyna (BS 26491) shows the same phenomenon of a deeply divided staminate calyx (like the Sri Lankan S. assimilis Thwaites = B. androgyna, the Indian B. trinervia a (Hook.f. & Thomson ex Müll.Arg.) Chakrab. & N.P.Balakr., and Bornean B. micrasterias).

4. For differences between B. lanceolata and B. macrantha see note under latter.

xx. Breynia macrantha (Hassk.) Chakrab. & N.P.Balakr.

    Breynia macrantha (Hassk.) Chakrab. & N.P.Balakr., Bangladesh J. Pl. Taxon. 19 (2012)  121; Welzen et al., Blumea 59 (2014) 92. — Sauropus macranthus Hassk., Retzia 1 (1855) 166; Hort. Bogor. Descr. (1858) 52; Müll.Arg. in DC., Prodr. 15, 2 (1866) 240; Backer & Bakh.f., Fl. Java 1 (1963) 471; Airy Shaw & Radcl.-Sm., Dansk Bot. Ark. 25 (1967) 34, fig. 13; Airy Shaw, Kew Bull. 26 (1972) 336; Whitmore, Tree Fl. Malaya 2 (1973) 131; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 193; Kew Bull. Add. Ser. 8 (1980a) 200; Kew Bull. 35 (1980b) 681; Kew Bull. 36 (1981) 343; Alph. Enum. Euph. Philipp. Isl. (1983) 44; Chakrab. & M.G.Gangop., J. Econ. Tax. Bot. 20 (1996) 529, Fig. 6; Smitinand, Thai Pl. Names, rev. ed. (2001) 467; Welzen, Blumea 48 (2003) 358, fig. 3b, map 12; in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 540, Fig. 75b. — Aalius macranthus (Hassk.) Kuntze, Revis. Gen. Pl. 2 (1891) 591 (‘macrantha’). — Type: Teysmann s.n. (L, barcode L  0138428), Indonesia, Hortus Bogoriensis.

    Sauropus spectabilis Miq., Fl. Ned. Ind., eerste bijv. (1861) 446; Müll.Arg. in DC., Prodr. 15, 2 (1866) 240; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 219; Ridl., Fl. Malay Penins. 3 (1924) 220; Beille in Lecomte, Fl. Indo-Chine 5 (1927) 647; M.R.Hend., J. Malayan Branch Roy. Asiat. Soc. 17 (1939) 30, 72. — Aalius spectabilis (Miq.) Kuntze, Revis. Gen. Pl. (1891) 591. — Type: Teysmann HB 4481 (holo U), Indonesia, Sumatra, Lampung Prov. (Lampong), Radja-bassa.

    Sauropus macrophyllus Hook.f., Fl. Br. India 5 (1887) 333, pro parte; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 226. — Aalius macrophyllus (Hook.f.) Kuntze, Revis. Gen. Pl. (1891) 591 (‘macrophylla’). — Lectotype (designated by Welzen, 2003): Griffith KD 4834 (K), India, Upper Assam, Mishmi Hills at Laee Pane and Yen. (Other, former, syntype, Griffith KD 4828 = S. rhamnoides).

    Sauropus forcipatus Hook.f., Fl. Br. india 5 (1887) 334; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 218; Ridl., Fl. Malay Penins. 3 (1924) 220. — Aalius forcipatus (Hook.f.) Kuntze, Revis. Gen. Pl. (1891) 591 (‘forcipata’). — Phyllanthus forcipatus (Hook.f.) Chakrab. & N.P.Balakr., J. Econ. Taxon. Bot. 33 (2009) 715, non P. macranthus Pax, nor P. spectabilis Miq. — Type: Scortechini 1254 (holo K), Malaysia, Perak.

    Glochidion umbratile Maiden & Betche, Proc. Linn. Soc. New South Wales 30 (1905) 370. — Type: E. Betche s.n., Aug. 1901, barcode NSW247350 (holo NSW), Australia, North Queensland, Atherton.

    Sauropus robinsonii Merr., Philipp. J. Sci. 7 (1912) 407; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 220; Merr., Enum. Philipp. Fl. Pl. 2 (1923) 405. — Type: Philippines: Elmer 6441 (holo PNH, †; iso BM, K (2sheets), L, P), Luzon, Benguet Subprov., Kias.

    Sauropus wichurae Müll.Arg. ex Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 220. — Lectotype (designated by Welzen, 2003): Wichura 2104 (holo G), Indonesia, Java, Tjisurupan.

    Sauropus grandifolius Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 222; Beille in Lecomte, Fl. Indo-Chine 5 (1927) 648. — Henry 11765A (holo K; iso K, 2 sheets), China, Yunnan, Szemao.

    Sauropus grandifolius Pax & K.Hoffm. var. tonkinensis Beille in Lecomte, Fl. Indo-Chine 5 (1927) 648. — Type: Balansa 3842 (holo P; iso K), Viet Nam, Tonkin, vallée de Lankok, mont Bavi.

    Sauropus longipedicellatus Merr. & Chun, Sunyatsenia 2 (1934) 34. — H.Y. Liang 61709 (holo NY; iso A), China, Hainan, Seven Finger Mountain.

Shrubs totreelets, up to 5 m high, up to 22 mm dbh; branches spreading horizontally, with 2, often indistinct ribs, flowering branches 2–4 mm thick. Indumentum absent. Bark thin, smooth, tan to greyish brown. Stipules triangular, 2.3–8.5 by 1.3–2.8 mm, basally eared, dark brown when dry with yellowish margin, stiff, breaking off easily, not caducous. Leaves: petiole 3–5 mm long, cracked when dry, flattened above but with marginal and midrib ridges; blade ovate (to elliptic), 3.8–20 by 1.9–8.2 cm, length/width ratio 1.6–3.5, papery, base broadly cuneate, margin flat, apex (gradually) cuspidate, usually mucronulate, upper surface glossy dark green above, lower surface lighter bright green, never papillate; venation distinct on both sides but hardly raised, nerves 10–17, veins more or less scalariform, veinlets reticulate. Inflorescences axillary fascicles, flowers pendent, usually single or few together of same or mixed sexes. Staminate flowers 2.5–4.5 mm in diameter, red; pedicel 7.5–15 mm long; calyx 1.5–2.8 mm deep, 6-lobed, lobes triangular, 0.5–1.2 by 1–2 mm, leathery, apex rounded to emarginate, scales low; stamens: androphore c. 0.2 mm high, anthers c. 0.3 by 0.3–0.6 mm. Pistillate flowers 3.5–12 mm in diameter, greenish red to red to purplish; pedicel up to 7.5 cm long in fruit, apically more than 1 mm thick; calyx lobes 6, only at very base attached, thick, 3 smaller, c. square to ovate, 2.2–3 by 2.3–3 mm; larger ones elliptic, 3–5.3 by 2.2–3.1 mm; ovary bell-shaped to ellipsoid, 2–2.5 by 2.2–3 mm, stigmas horizontal, c. 2 mm long, upper ²/₃^rd split, thick, curled in more than a complete circle, free from ovary. Fruits flattened globose to somewhat lobed, 17–22 by 10–13 mm, broader than high, (pink to) red to purplish, wall thin, mericarp white and edible; column c. 9 mm long, tapering towards the apex. Seeds triangular in transverse section, hollow adaxially, 10–10.5 by 6.5–8 by 5–6 mm, black or brown.

    Distribution — NE India to S China, SE Asia, throughout Malesia to N Australia (Queensland).

    Habitat & Ecology — (Dry) evergreen forest, primary forest, secondary forest (with much bamboo), scrub; very dry to swampy, along rivers, ridges, on steep slopes; soil: deep fertile soil, humus rich to swampy boulder strewn soil; limestone bedrock. Altitude: 200–2000 m. Flowering and fruiting whole year through, though somewhat less in December—February.

    Uses — On Java planted as ornamental. Mericarp edible. Leaves sometimes used as vegetable, but less popular than B. androgyna.

    Vernacular names — Thailand: Mayom khao. Sumatra: Gening-gening (Karo). Java: Manis manisan. Irian Jaya: Nibasam (Andjai).

    Note — This species has – within Breynia – large leaves. However, especially in Australia the leaves are usually much smaller (less than 10 cm). On Java the pistillate pedicels remain shorter, within the range of B. lanceolata. The latter species can relatively easily be distinguished from B. macrantha, even when in pistillate flower only, as it has caducous stipules and a very different kind of staminate flower (calyx enlarged, circular, hardly lobed).

XX. Breynia micrasterias (Airy Shaw) Welzen & Pruesapan

    Breynia micrasterias (Airy Shaw) Welzen & Pruesapan in Welzen et al., Blumea 59 (2014) 92. — Sauropus micrasterias Airy Shaw, Kew Bull. 14 (1960) 354; Kew Bull. 16 (1963) 344; Kew Bull. Add. Ser. 4 (1975) 194; Welzen, Blumea 48 (2003) 360, fig. 2b, map 4. — Type: Jacobs 5179 (holo K; iso L), Sarawak, 1^st Division, rock formation (Bau series) W. and E. of passage of Sungei Serin (30 miles S of Kuching).

Shrub (with creeping habit), up to 1.5 m high, sprouting from long roots; young branches with 2 faint ribs, flowering branches at least 4 mm thick. Indumentum absent. Stipules triangular, basally eared, 2.5–3 by 1.1–1.3 mm, stiff, rather persistent. Leaves: petiole 3–4 mm long, flattened above; blade elliptic (to somewhat obovate), 14–20.5 by 6.3–8.5 cm, length/width ratio 2.2–2.4, papery, base cuneate, descending into petiole, margin flat, apex cuspidate, not mucronulate, lower surface not papillate; venation rather distinct on both sides, especially beneath, nerves 12–15, veins more or less scalariform, veinlets reticulate. Inflorescences cauliflorous near ground to axillary, racemes (to panicles), up to 14 cm long, red, often with some asperities, with per node groups of staminate flowers and a single pistillate flower; staminate flowers sometimes on short, up to 5 mm long, branches. Flowers dull raspberry red to dark red. Staminate flowers 5.5–8 mm in diameter; pedicel 5–6 mm long; calyx c. 4 mm deep; lobes narrowly triangular, c. 3 by 0.9–1 mm, erect, apex rounded, scales slightly broader than lobes; stamens: androphore c. 0,3 mm high, anthers c. 0.5 by 0.7 mm, cream. Pistillate flowers c. 4 mm in diameter; pedicel c. 2.2 mm long; calyx almost completely split, lobes only basally attached, obovate, 3 smaller, c. 2 by 1.3 mm, 3 larger, c. 2.5 by 1.3 mm, apex rounded; ovary c. 1 by 1.3 mm, pinkish; stigmas horizontal, c. 1 mm long, split till halfway, forming less than a circle. Fruits and seeds unknown.

    Distribution — Endemic in Sarawak, near Kuching.

(B. micrasterias: stars; B. asymmetrica: dots; B. shawii: triangles)

    Habitat & Ecology — Primary and secondary forest at foot of limestone hills, land wet (recorded once) with scattered limestone boulders; soil: yellow clay loam, limestone. Altitude 70–400 m. Flowering: March, August, October.

xx Breynia obscura Welzen & Pruesapan

    Breynia obscura Welzen & Pruesapan, Thai For. Bull., Bot. 38 (2010) 115, fig. 4. — Type: Molesworth-Allen s.n, 31 Jan. 1959. (holo SING!), Malaysia, Perak, Ipoh, Gunong Rapat. 

Subshrub; branches round, hairy, drying light greenish, bark not fissuring; flowering branches c. 0.7 mm thick. Indumentum of whitish patent hairs, most parts hirsute. Stipules triangular, c. 0.8 by 0.5 mm, hairy, not very persistent. Leaves simple, alternate; petiole c. 0.8 mm long, channelled above, hirsute; blade ovate to round, 1.3–1.6 by 1.3–1.7 cm, length/width ratio 0.9–1.1, very thin, almost translucent, symmetric, base emarginate to truncate, margin entire, flat, apex rounded, upper and lower surface only hairy on the midrib, upper surface darker, lower surface glaucous, venation very distinct, reticulate, raised on both sides, nerves 7 or 8 per side, looped and closed near the margin. Staminate flowers single per node, c. 2 mm in diam., glabrous; pedicel up to 8 mm long, round, slightly thickening upwards; calyx consisting of 6 almost free lobes; scales broader than lobes; staminal column short, anthers small. Pistillate flowers, fruits and seeds unknown. 

    Distribution — Endemic to the Malay Peninsula. 

    Habitat & Ecology — Limestone ledge, usually in holes on cliffs. Flowering: January. 

    Notes — Known from the type collection only, one poor quality specimen (few loose leaves and a small staminate fl ower on a short piece of branchlet) with distinct characters like the almost completely free staminate lobes, presence of hairs and basally attached leaves.

xx. Breynia suberosa (Airy Shaw) Welzen & Pruesapan

    Breynia suberosa (Airy Shaw) Welzen & Pruesapan in Welzen et al., Blumea 59 (2014) 92. — Sauropus suberosus Airy Shaw, Kew Bull. 23 (1969) 42; Kew Bull. 25 (1971) 500; Kew Bull. 26 (1972) 338; Whitmore, Tree Fl. Malaya 2 (1973) 130; Smitinand, Thai Pl. Names, rev. ed. (2001) 468; Welzen, Blumea 48 (2003) 375, map 20; in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 549. — Type: Hansen & Smitinand 12030 (holo K; iso L, SING), Thailand, Peninsular, Phuket, Khao Thong Lang, NW of Nai Chong.

Shrubs, up to 4 m high, simply branched; stems at most with indististinct ribs when very young, sometimes with small asperities, older branches with thick, longitudinally fissured, soft, brown cork. Indumentum simple hairs only, glabrous except for inflorescences. Stipules triangular, 3.5–6 by 1.2–4, basally eared, very thick, light margin, stiff, easily breaking off, otherwise persistent. Leaves: petiole 2–4 mm long, flattened dorsoventrally, especially above, sometimes minute asperities; blade elliptic to obovate, 7.7–25 by 2.6–7.3 cm, length/width ratio 3–3.6, papery, base cuneate, margin flat to revolute, apex acute to cuspidate, often mucronulate, venation relatively distinct on both sides, nerves 12 or 13, veins and veinlets reticulate. Inflorescences racemes, cauliflorous in basal part of stem, sometimes over ground, up to 41 cm long, slightly hairy, 4-ribbed or somewhat flattened, rachis dark reddish, with groups of flowers of which one or two pistillate; flowers pink, dark reddish, dark purplish or yellowish. Staminate flowers 3.7–4 mm in diameter; pedicel up to 5.5 mm long; calyx lobes 0.8–1 by 1–1.6 mm, up to one third of calyx height, rounded, often folded inward; stamens: androphore c. 0.2 mm long, anthers 0.4–0.5 by 0.4–0.5 mm. Pistillate flowers 4.3–7.7 mm in diameter; pedicel 2.5–4.5 mm long, glabrous to hairy; calyx lobes thick, mostly obovate, three smaller outer ones 1.2–2.5 by 2.1–4 mm, inner larger ones 1.5–3 by 2–4.2 mm; ovary 1.1–1.8 by 1.1–2 mm; stigmas 1–1.3 mm long, horizontal, split in upper half, forming more than a complete circle. Fruits unknown.

    Distribution — Thailand (Peninsular floristic district), Peninsular Malaysia (Perak).

    Habitat & Ecology — Thicket in moist, evergreen forest on mountain ridge; soil: limestone, rocky with a fair amount of accumulated soil. Altitude: 80–800 m. Flowering: November to January.

    Vernacular name — Thailand: Mayom yak.

xx. Breynia villosa (Blanco) Welzen & Pruesapan.

    Breynia villosa (Blanco) Welzen & Pruesapan in Welzen et al., Blumea 59 (2014) 93. — Kirganelia villosa Blanco, Fl. Filip. (1837) 712; ed. 2 (1845) 493; ed. 3, 3 (1879) 116, t. 399; Merr., Sp. Blancoan. (1918) 217. — Sauropus villosus (Blanco) Merr., Contrib. Arn. Arb. 8 (1934) 86; Airy Shaw, Kew Bull. 23 (1969) 49; Kew Bull. 26 (1972) 339; Whitmore, Tree Fl. Malaya 2 (1973) 130; Airy Shaw, Kew Bull. 36 (1981) 343, Fig. 12A1–4; Alph. Enum. Euph. Philipp. Isl. (1983) 44; Smitinand, Thai Pl. Names, rev. ed. (2001) 468; Welzen, Blumea 48 (2003) 380, map 21; in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 552. — Neotype (Welzen, 2003): Merrill Species Blancoanae 931 (holo L; iso A, BM, K, NSW, NY, P, US), Philippines, Luzon, Rizal Prov.

    [Phyllanthus pubescens Klotzsch (in Meyen, Observ. Bot.), Nova Acta Phys.-Med. Caes. Leop.-Carol. Nat. Cur. 19, Suppl. 1 (1843) 420 [non Moon, Cat. Pl. Ceylon (1824) 65; non Wall., Numer. List (1847) 7917 A, B], nom. illig.] — Glochidion llanosii Müll.Arg., Linnaea 32 (1863) 68; Ridl., J. Straits Branch Roy. Asiat. Soc. 59 (1911) 173; Merr., Enum. Philipp. Fl. Pl. 2 (1923) 400. — Phyllanthus llanosii (Müll.Arg.) Müll.Arg., Flora 48 (1865) 387; in DC., Prodr. 15, 2 (1866) 308. — Sauropus llanosii (Müll.Arg.) Gage, Rec. Bot. Surv. India 9 (1922) 223; Ridl., Fl. Malay Penins. 3 (1924) 221; M.R.Hend., J. Malayan Branch Roy. Asiat. Soc. 17 (1939) 30, 72. — Syntypes: Cuming 593 (n.v.), Philippines; Cuming 595 (G-DC, K, TCD), Philippines; Llanos s.n. (G-DC, n.v., IDC microfiche DC herbarium 2473/18), Philippines, 1858; Meyen s.n. (K (ex-B), 2 sheets), Philippines, Manila; (the types of Klotzsch’s Phyllanthus pubescens were not indicated and probably lost in B during the second world war except for the Meyen specimens in K).

Woody herbs to shrubs, up to 2 m tall, d.b.h. up to 2 cm; branches hirsute, light green, round, many, often longer than the stem; flowering branches 1–2 mm thick. Indumentum of simple hairs, hirsute, sparse to dense on most parts. Stipules triangular, 1.3–4 by 0.2–0.5 mm, subhirsute outside, glabrous inside. Leaves basally smaller than apically on branches; petiole 1.2–2 mm, dorsoventrally flattened, hairy; blade ovate, margins gradually tapering towards each other towards the apex, 1–6.5 by 0.6–1.9 cm, length/width ratio (1.5–)3.4, papery, not translucent, base emarginate to cuneate, sometimes somewhat oblique, margin often revolute, apex acute, often mucronulate, upper surface glabrous to subhirsute, especially when young, dark green, lower surface papillate, pale light green, sparsely to densely hairy; venation flat above, slightly raised below, nerves 9–13, rather distinct, veins and veinlets rather indistinct, reticulate. Inflorescences axillary fascicles, flowers usually few (to many) together of same or mixed sexes. Staminate flowers 1–3 mm in diameter, yellow, glabrous except for a few hairs sometimes on pedicel and calyx outside; pedicel 1.2–4 mm long, glabrous; calyx flat, slightly 6-lobed, lobes up to 1.2 by 0.4 mm, hairy outside, glabrous inside; scales distinct or indistinct and united with lobes, providing a thick apex, apex usually erose; stamens erect, united along long side, androphore c. 0.2 mm long, pinkish, anthers 0.6–0.7 by c. 0.4 mm, connective sometimes slightly elongated. Pistillate flowers 1.2–2 mm in diameter, yellow-green (to white); pedicel 0.7–1(–2.5 in fruit) mm long, hairy; calyx lobes 0.4–0.7 by 0.5–0.7 (in fruit up to 1.3 by 1.3) mm, yellow-green, apex acute, hairy outside, glabrous inside; ovary 0.7–1 by 0.8–0.9 mm, light green, glabrous; stigmas 0.6–1.2 mm long, horizontal, upper 0.5–0.7 mm free and in more than a complete circle, yellow green. Fruits flattened ellipsoid, 8–11 by 4–6 mm, broader than high, yellow to orangish red when mature, apically without a ring; column 1.2–2 mm long, tapering towards apex. Seeds triangular in transverse section, 4.5–5.5 by 3–4 by 2.8–3.2 mm.

    Distribution: Thailand (Southeastern and Peninsular Dist.), Viet Nam (Annam), N Malay Peninsula, Sumatra, Philippines.

    Habitat & Ecology — Locally abundant in especially disturbed sites like secondary growths, meadows, open thickets, scrubs, savannah, coconut plantations, open forest, but also in semi-deciduous forest; often near the sea shore; soil: sand. Altitude: 0–600 m. Flowering and fruiting: September to June.

    Uses — Philippines: Trunk makes good construction material for, for instance, flooring.

    Vernacular names — Thailand: Ngap yai, tan ngan khao (official Thai name). Sumatra: Bebaho (Alas). Philippines: Lanitaga (Bisaya).

    Note — The staminate scales are separate from the calyx lobes in Thailand, but in other areas they are grown together, giving a thick lobe apex with in the middle a constriction. The plants of Sumatra usually have a denser indumentum than those of other areas.