Malesian Euphorbiaceae Descriptions
Barker, C. & P.C. van Welzen. 2010. Flueggea (Euphorbiaceae s.l. or Phyllanthaceae) in Malesia. Syst. Bot. 35: 541–551.
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Key to the species
Flueggea Willd., Sp. Pl. 4, 2 (1806) 637; A. Juss., Euphorb. Gen. (1824) 16; Blume, Bijdr. (1825) 580; Baill., Étude Euphorb. (1858) 590; Benth. & Hook. f., Gen. Pl. 3 (1880) 276; Hook.f., Fl. Brit. India 5 (1887) 328; Beille in Lecomte, Fl. Indo-Chine 5 (1927) 528; G. L. Webster, Allertonia 3 (1984) 273; Ann. Missouri Bot. Gard. 81 (1994) 43; Welzen, Thai Forest Bull. (Bot.) 28 (2000) 56; Radcl.-Sm., Gen. Euphorbiacearum (2001) 35; C.Barker & Welzen in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 299; Petra Hoffm., Kathriar. & Wurdack, Kew Bull. 61 (2006) 44; C. Barker & Welzen, Syst. Bot. 35 (2010) 541; G.L.Webster in Kubitzki, Fam. Gen. Vasc. Pl. 11 (2014) 77. — Securinega Comm. ex Juss. sect. Flueggea (Willd.) Müll. Arg. in DC., Prodr. 15, 2 (1866) 448; Pax & K. Hoffm. in Engl. and Harms, Nat. Pflanzenfam. ed. 2, 19c (1931) 60. — Type: Flueggea leucopyrus Willd.
Acidoton P.Browne, Civ. Nat. Hist. Jamaica (1756) 355., nom. rej., non Sw. (1788). — Type: Acidoton frutescens aculeatum et diffusum P. Browne, nom. rej. [= Flueggea acidoton (L.) G. L. Webster].
Bessera Spreng., Pl. Min. Cogn. Pug. 2 (1815) 90. — Type: Bessera inermis Spreng. [= Flueggea virosa (Roxb. ex Willd.) Royle subsp. virosa].
Geblera Fisch. & C.A.Mey., Index Sem. Hort. Petrop. 1 (1835) 28. — Type: Geblera suffruticosa (Pall.) Fisch. & C. A.Mey. [= Flueggea suffruticosa (Pall.) Baill.].
Colmeiroa Reut., Biblioth. Universelle Genève 38 (1842) 215. — Type: Colmeiroa buxifolia Reut. [= Flueggea tinctoria (L.) G. L. Webster].
Pleiostemon Sond., Linnaea 23 (185) 135. — Type: Pleiostemon verrucosum (Thunb.) Sond. [= Flueggea verrucosa (Thunb.) G. L. Webster.]
Neowawraea Rock, Indig. Trees Haw. Isl. (1913) 243. — Type: Neowawraea phyllanthoides Rock [= Flueggea neowawraea W. J. Hayden].
Richeriella Pax & K. Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 30; Merr., Enum. Philipp. Fl. Pl. (1923) 411; Airy Shaw, Kew Bull. 25 (1971) 489, 491, 492; Kew Bull. 26 (1972) 328; Whitmore, Tree Fl. Malaya 2 (1973) 128; Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 190; Alphab. Enum. Euphorb. Philipp. Isl. (1983) 43; G. A. Levin, Ann. Missouri Bot. Gard. 73 (1986) 75; Mennega, Bot. J. Linn. Soc. 94 (1987) 123, Table 2; G. L. Webster, Ann. Missouri Bot. Gard. 81 (1994) 44; Stuppy, Syst. Morph. Anat. Samen biovul. Euphorb. (1995) 157, 162; Radcl.-Sm., Gen. Euphorb. (2001) 36; T. Tokuoka & Tobe, J. Plant Res. 144 (2001) 75; Sagun & R. W. Ham, Grana 42 (2003) 205, Fig. 7 e–j, Table 2. 2003; Kathriarachchi et al. Mol. Phylog. Evol. 36 (2005) 127, Figs. 1, 3, 4; Welzen in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 515; G.L.Webster in Kubitzki, Fam. & Gen. Vasc. Pl. 11 (2014) 77. — Type: Richeriella gracilis (Merr.) Pax & K. Hoffm. [= Flueggea gracilis (Merr.) Petra Hoffm.]
Securinega auct. non Comm. ex Juss.: Backer & Bakh.f., Fl. Java 1 (1963) 465; Airy Shaw, Kew Bull. 26 (1972) 340; Whitmore, Tree Fl. Malaya 2 (1973) 131; Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 196; Kew Bull. 35 (1980) 687; Kew Bull., Addit. Ser. 8 (1980) 200; Kew Bull. 36 (1981) 345; Alphab. Enum. Euph. Philip. Isl. (1983) 45.
Shrubs or trees, usually dioecious, branching monopodial plagiotropic. Indumentum generally absent. Stipules triangular, small, apex acute. Leaves alternate, simple; petiole short; blade symmetric, papery, margin entire, surfaces smooth, venation pinnate, nerves looped and closed near margin, veins and veinlets reticulate. Inflorescences usually axillary (to ramiflorous), flowers usually in fascicles, sometimes racemes to panicles, bracteate, nodes usually with groups of staminate flowers or a single to several pistillate ones. Flowers actinomorphic, small; pedicels slender; sepals (4 or)5(–7), imbricate, subpetaloid, unequal, margins entire to denticulate; petals absent; disc or disc lobes present. Staminate flowers: pedicel short; disc-glands (4 or)5(–7), interstaminal, alternisepalous; stamens (4 or)5(–7), filaments free, anthers 2-locular, dehiscing extrorsely with longitudinal slits; pistillode consisting of 2 or 3 large strap-like filaments, apically bent. Pistillate flowers: pedicels long, slender, enlarging in fruit; disc flat, annular, sometimes thickened, entire or shallowly 5- or 6-lobed; ovary (2)3(4)-locular, ovules 2 per locule; style absent to short, stigmas free, recurved or spreading, apical part bifid, persistent in fruit. Fruits capsular, dehiscing loculicidally into 3 merocarps or subbaccate, indehiscent, columella usually persistent; endocarp crustaceous. Seeds 2 per locule, hilum sometimes invaginated, testa smooth or occasionally reticulate or verruculose, crustaceous; endosperm copious. Embryo straight to distinctly curved, lacking chlorophyll; cotyledons flat, broad, longer than the radicle.
Distribution — Fifteen species are found throughout the tropics and subtropics, extending into temperate eastern Asia. Three species occur in Malesia.
Habitat & Ecology — Seasonal vegetation, rainforest, montane forest (not in Malesia), tropical savannas, deciduous forests and scrub; occasionally on limestone. Elevation: Sea level up to 3,000 m (1,000 m in Malesia).
Note — For a long time Flueggea was included within Securinega, but as detailed above, the two genera should be separated (Webster 1984: 260). Securinega is restricted to a small group of species from Madagascar and the Mascarene Islands.
Leaf blades small, 1–10.5 cm long, obovate to elliptic or suborbicular, apex rounded or obtuse. Inflorescences fascicles. Shrubs to trees up to 3(–6) m high
Leaf blades large, 6–23 cm long, elliptic to oblong (to obovate), apex acuminate. Inflorescences fascicles or racemes to panicles. Shrubs to trees up to 20 m high
Inflorescences fascicles. Fruits 3.5–4 mm in diameter — Philippines (not Palawan) to New Guinea
|1. Flueggea flexuosa|
Inflorescences racemes to panicles, staminate ones up to 17 cm long, pistillate ones up to 5 cm long. Fruits 8–12 mm in diameter — SE Asia, Malay Peninsula, Borneo, Philippines (Palawan)
|2. Flueggea gracilis|
Flueggea flexuosa Müll.Arg., Linnaea 34 (1865) 76. 1865; C.B.Rob., Philipp. J. Sci. 4, Bot.(1909) 74; G.L.Webster, Allertonia 3 (1984) 284; C.Barker & Welzen, Syst. Bot. 35 (2010) 542, Fig. 1. — Securinega flexuosa (Müll. Arg.) Müll.Arg. in DC., Prodr. 15, 2 (1866) 450; Merr., Philipp. J. Sci. 9, Bot. (1914) 491; Enum. Philipp. Fl. Pl. 2 (1923) 290; Whitmore, Guide For. Brit. Solomon Is.: 151 (1966) 201; Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 201. 1980; Alphab. Enum. Euph. Philip. Isl. (1983) 45. — Acidoton flexuosus (Müll. Arg.) Kuntze, Rev. Gen. Pl. 2 (1891) 592. — Type: Cuming 1855 (holotype: G-DC; isotype: K!), Philippines, Bohol.
Phyllanthus acuminatissimus C.B.Rob., Philipp. J. Sci. 3, Bot. (1908) 200. — Securinega acuminatissima (C.B.Rob.) C.B.Rob., Philipp. J. Sci. 4, Bot. (1909) 73. — Type: R.S. Williams 2807(holotype: NY; isotypes: K, NY), Philippines, Mindanao, Dist. of Davao, Santa Cruz.
Securinega samoana Croizat, Bull. Bernice P. Bishop Mus. 184 91945) 45.—Type: E. Christophersen 2765 (holotype: NY; isotypes: K, P), Samoa, Savaii, Tufutafoe-Falelima.
Shrub-like trees to trees, evergreen, up to 17 m high, bole 10 m, dbh up to 33 cm; buttresses occasionally present, up to 1 m high, thin, equal; twigs terete to angled, smooth to striate, flexuous, often weakly so, occasionally lenticellate; flowering branchlets 1.6–2 mm in diameter. Outer bark pale grey to grey-brown to light or sandy brown, smooth to fissured to scaly; inner bark pink, fibrous, soft, with clear sap; sapwood white to pale brown, soft to usually hard, heartwood hard, red to pale brown. Stipules 1–1.3 by 1–1.7 mm, acuminate, caducous. Leaves: petiole 5–11 mm long, channelled above; blade elliptic to oblong, 6–18.5 by 3–8 cm, length/width ratio 2.3–3.1, base cuneate to rounded and decurrent with the petiole, margin slightly incurved on drying, apex acuminate, the acumen up to 2 cm long, often apiculate, upper surface glossy green, lower surface light green to subglaucous, venation especially distinct underneath, midrib impressed above, with 6–9 pairs of nerves per side, midrib impressed above; secondary veins 6–9 pairs. Flowers in axillary fascicles; bracts triangular to ovate, c. 0.8 by 0.8 mm. Staminate flowers at least 20 per axillary cluster, 3–4.3 mm in diameter, white; pedicel 3–7 mm long; sepals 5, apex broadly rounded, outer 2 oblong, 1.1–1.3 by 0.7–0.9 mm, inner 3 larger, obovate-oblong, 1.3–1.7 by 1.3–1.5 mm; disc segments 5, fleshy, 0.4–0.6 mm diameter; stamens 5, filaments 1–1.4 mm long, white, anthers ellipsoid, 0.4–0.6 by 0.4–0.6 mm, yellow; pistillode 0.7–1.2 mm long, divided into 2 or 3 branches to half to a third of its length, bent part 0.2–0.3 mm long. Pistillate flowers 3–7 per axillary cluster, c. 4 mm in diameter, green to yellowish green; pedicel 3.5–7.7 mm long; sepals 5, apex broadly rounded, outer 2 oblong, 1.2–1.5 by 0.8–1.1 mm, inner 3 larger, broadly ovate to orbicular, 1.2–1.6 by 1.0–1.5 mm; disc flat, thick, 1–1.6 mm in diameter; ovary 3(4)-locular, 1–1.5 by 0.8–1.2 mm; styles almost absent, stigmas 0.6–1 mm long, 2-lobed to half to a quarter of their length. Fruits globose to shallowly 3-lobed, 3.5–4 mm in diameter, red and fleshy when mature, sepals and styles persistent, dehiscing irregularly; columella early caducous. Seeds obovoid-triangular, 2–2.5 by c. 1.5 by 1.3 mm, with a broad, shallow hilar invagination.
Distribution — Malesia: Philippines (Luzon, Catanduanes, Samar, Leyte, Bohol, Negros, Mindanao), Moluccas (Morotai; Sula Islands: Mangoli), New Guinea (Irian Jaya: Vogelkop). Bougainville, Solomon Islands, Vanuatu, New Caledonia, Fiji, Wallis and Futuna (Horne Island), Samoa. Cultivated in Papua New Guinea (Lae Botanic Garden), Vanuatu, Samoa (Upolu), Cook Islands (Rarotonga).
l = Flueggea flexuosa; n = Flueggea gracilis.
Habitat & Ecology — The species is present in lowland primary and secondary rainforests, kaingin (shifting cultivation) fields, often along rivers; soil: loam, clay, limestone. Alt.: up to 500 m. Flowering throughout the year with maxima in March to April and August to November; fruiting is most commonly in June to November, but also in February.
Uses — The heavy, hard, straight timber is much used in the Solomon Islands, Vanuatu and Fiji for general house construction, fencing and sometimes for heavy construction and bridging. In the Reef Islands it has been used for buildings that stand in the sea as the wood is said to be durable in salt water (Henderson and Hancock, 1988, Guide Usef. Pl. Solomon Isl.: 190–192). On Vanuatu the bark and leaves are used medicinally, the latter are boiled and rubbed over the body for curing scabies, especially in small children; the bark is probably applied in the same way.
Vernacular Names — Philippines: Anislag or Anislog (Bikol, Cebu Bisáya, Samar-Leyte Bisáya, Manóbo, Mandáya); Anisláng, Hamislág (Bikol); Katamañgan, Malañgau (Manóbo); Tras (Magindanáo) ( Merrill, 1923 ). Bougainville: Mewana.
Notes — Flueggea flexuosa is clearly related to the widespread F. virosa, having similar more or less baccate fruits that dehisce irregularly and seeds with a hilar invagination. It differs in its larger, acuminate leaves and larger floral parts. Flueggea flexuosa and F. gracilis are both trees and apparently evergreen, which is unusual in the genus. Other species are usually deciduous shrubs found in seasonal habitats.
Flueggea gracilis (Merr.) Petra Hoffm. in Petra Hoffm., Kathriar. & Wurdack, Kew Bull. 61 (2006) 44; C.Barker & Welzen, Syst. Bot. 35 (2010) 544, Fig. 1, 2. — Baccaurea gracilis Merr., Philipp. J. Sci. 1, Suppl. (1906) 203. — Richeriella gracilis (Merr.) Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 30; Merr., Enum. Philipp. Fl. Pl. (1923) 411; Lingnan Sci. J. 14 (1935) 22; Punt, Wentia 7 (1962) 26; Eg. Köhler, Grana Palyn. 6 (1965) 45, 72, 100, t. 4: fig. 5–8; Airy Shaw, Kew Bull. 25 (1971) 489; Kew Bull. 26 (1972) 328; Alphab. Enum. Euphorb. Philipp. Isl. (1983) 43; Welzen in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 515, Fig. 68. — Lectotype (designated by C.Barker & Welzen, 2010): Bermejos BS 218 (lectotype: K!; isolectotype: US), Philippines, Palawan. Former syntypes: Bermejos BS 258 (syntype: K), Philippines, Palawan; Curran FB 3828 (syntypes: K!, US), Philippines, Palawan.
Richeriella malayana M.R.Hend., Gard. Bull. Straits Settlem. 7 (1933) 122, Pl. 32; J. Malayan Branch Roy. As. Soc. 17 (1939) 71. 1939; Airy Shaw, Kew Bull. 25 (1971) 489, 491, 492; Whitmore, Tree Fl. Malaya 2 (1973) 128; Airy Shaw, Hooker’s Icon. Pl. 38 (1974) t. 3703; Kew Bull., Addit. Ser. 4 (1975) 190. — Type: M.R. Henderson SFN 23790 (holotype: SING; isotypes: K!, 2 sheets, SING), Malaysia, Perak, Gunong Pondok.
Richeriella malayana M.R.Hend. var. macrocarpa Airy Shaw, Kew Bull. 25 (1971) 490; Kew Bull., Addit. Ser. 4 (1975) 190. — Type: J.A.R. Anderson S 12542 (holotype: K!; isotypes: L!, SAR), Malaysia, Sarawak, Kuching dist., Tiang Bekap.
Shrubs to trees, evergreen, up to 20 m high, dbh up to 15 cm; bole sometimes fluted; flowering branchlets 2–3 mm in diameter, drying greyish. Outer bark greyish to grey brown to light brown to fawn to dull dark brown, smooth to somewhat flaky, fibrous; inner bark pale yellow green to pale pink, granular, fibrous; sapwood cream to yellow. Stipules 1.7–4.3 by 1–2.2 mm, stiff, caducous. Leaves: petiole 3–9 mm long, flattened above; blade elliptic-oblong to slightly obovate, 6.7–23 by 2.6–9 cm, length/width ratio 2.2–2.7, base cuneate to attenuate and then decurrent with the petiole, margin flat, apex bluntly acuminate (generally damaged), upper surface glossy, dark green above, lower surface paler, midrib yellowish brown; venation especially distinct and raised underneath, with 9–14 nerves per side. Inflorescences axillary to ramiflorous racemes to panicles, pale light green, pendent, staminate ones up to 17 cm long, pistillate ones up to 5 cm long; bracts triangular, 0.5–1.3 by 0.5–1.2 mm. Staminate flowers 1.5–2.5 mm in diameter, cream to white; pedicel 0.3–0.7 mm long; sepals 5, 2 or 3, outer ones smaller, c. 1.2 by 1 mm, 2 or 3, inner ones larger, 1.2–1.3 by 1.2–1.3 mm; disc lobes 5, c. 0.2 by 0.2 mm; stamens 5, filaments 1.7–3 mm long, white, anthers ellipsoid, 0.5–0.8 by 0.4–0.6 mm, yellow; pistillode 2 hooks, 1.2–1.6 mm high, bent part 0.5–0.6 mm long. Pistillate flowers unknown, data from fruits: pedicel 11–20 mm long in fruit; sepals 5, 2 outer ones 1.1–1.4 by 1.1–1.4 mm, 3 inner ones c. 1.5 by 1.5 mm; ovary 3(4)-locular. Fruits lobed, dehiscent, 8–12 by 4.5–7 mm, black, slightly succulent, somewhat wrinkled when dry; columella 2.4–4 mm long. Seeds obovoid, 3–5.5 by 2.4–4.7 by 1.8–4.5 mm, with a broad, shallow hilar invagination.
Distribution — China (Hainan), Peninsular Thailand, Malay Peninsula, Borneo, Philippines (Palawan).
l = Flueggea flexuosa; n = Flueggea gracilis.
Habitat & Ecology — Primary and secondary rainforest, open shrubbery; soil: sandy clay, sandy beach, limestone rocks with loam cover. Elevation: Sea level up to 333 m. Flowering and fruiting probably throughout the year.
Vernacular names — Borneo, Sarawak: Mabon (Land Dayak). Philippines: Marabakawan (Palawan).
Notes — 1. The long inflorescences instead of fascicles are a unique character for F. gracilis. The names Richeriella gracilis, R. malayana, and R. malayana var. macrocarpa are all united under F. gracilis. Airy Shaw (1971) mentioned as differences between these species that R. gracilis has smaller leaves (up to 16 cm), whereas they are up to 28 cm in R. malayana. He also stated that the latter generally has larger inflorescences with more branches than R. gracilis. However, the overlap in leaf size is too large to merit distinction and the size and degree of branching of the inflorescences seems to be a geocline, the more temperate specimens (Hainan, Palawan, Thailand) having smaller inflorescences with fewer branches. Uniting both species creates a continuous distribution.
2. Specimens identified as Richeriella malayana var. macrocarpa, known from Sarawak and part of Indonesian Kalimantan, have larger fruits than the typical F. gracilis specimens (11–12 by 6–7 mm versus 8–9.5 by 4.5–6 mm), but the difference is not impressive. The specimen SAN 99460 from Sabah is more or less intermediate, smaller sized fruits also occur in Sarawak and Kalimantan, and a number of other Euphorbiaceae s. l. species show a comparable variation (Alchornea rugosa (Lour.) Müll. Arg., Austrobuxus nitidus Miq., Cheilosa montana Blume). Therefore, the name is also synonymised with F. gracilis.
Flueggea virosa (Roxb. ex Willd.) Royle, Ill. Bot. Himal. Mts. (1836) 328; Voigt, Hort. Suburb. Calcut. (1845) 152; Baill., Étude Euphorb. (1858) 593; C.B.Rob., Philipp. J. Sci. 4, Bot. (1909) 74; J.J.Sm., Meded. Depart. Landbouw 10 (1910) 51, Merr., Enum. Philipp. Fl. Pl. (1923) 390; G.L.Webster, Allertonia 3 (1984) 287; C.Barker & Welzen in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 301, Fig. 75, Plate XVI: 4; Syst. Bot. 35 (2010) 546, Fig. 3. — Phyllanthus virosus Roxb. ex Willd., Spec. Pl. 4, 1 (1805) 578; Roxb., Fl. Ind. 3 (1832) 659; Wall., Numer. List (1847) 7928 A, B, C (Goyalpara), E. — Securinega virosa (Roxb. ex Willd.) Baill., Adansonia 6 (1866) 334; Backer & Bakh.f., Fl. Java 1 (1963) 466; Airy Shaw, Kew Bull. 26 (1972) 340; Whitmore, Tree Fl. Malaya 2 (1973) 131; Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 196 (excl. New Guinea and Australia); Kew Bull. 36 (1981) 345; Alphab. Enum. Euph. Philip. Isl. (1983) 45. — Acidoton virosus (Roxb. ex Willd.) Kuntze, Rev. Gen. Pl. 2 (1891) 592. — Type: Klein 748 (holotype: B-W!), India, INandaradah.
Shrubs or small trees, deciduous, up to 3(–6) m high; twigs angled, smooth, becoming lenticellate; flowering branches 0.7–2.4 mm in diameter. Stipules 1–3 by 0.7–1 mm, stiff, often caducous. Leaves: petiole 3–8 mm long, channelled above; blade elliptic to obovate, occasionally suborbicular, 1.3–10.5 by 0.6–6.2 cm, length/width ratio 1.6–2.3, base cuneate to rounded and decurrent with the petiole, sometimes asymmetric, margin slightly incurved on drying, apex rounded or obtuse, extreme tip acute or apiculate, surfaces dull, dark olive green above, paler to blue-green underneath; venation promimulous to prominently raised especially beneath, with 5–8 nerves per side. Flowers in axillary fascicles; bracts triangular, small. Staminate flowers 20–40 per axillary cluster, 1.7–2.2 mm in diameter, white to pale green to yellow to suffused red in center, aromatic; pedicel 3–6.5 mm long, green; sepals 5, margin subentire, apex broadly rounded to acute, the outer 2 ovate to elliptic, 0.8–1.2 by 0.5–0.7 mm, the inner 3 broadly ovate, 1.6–2 by 0.8 –1.3 mm; disc segments 5, fleshy, up to 0.2 mm long and wide, light yellow; stamens 5, filaments 1–2 mm long, light green to white, anthers ellipsoid, 0.3–0.5 by 0.3–0.4 mm, cream to light yellow, connective maroon; pistillode (2- or) 3-lobed, apically hooked, divided to two-thirds of its length, lobes 0.7–1.8 mm long, bent part 0.3–0.7 mm, cream to light yellow. Pistillate flowers 3–12 per axillary cluster, 1.5–2 mm in diameter, green; pedicels 1.5–5 mm long, up to 12 mm long in fruit, green; sepals as in staminate flower but slightly smaller, persistent in fruit, green with reddish margins to yellowish; disc 0.7–1 mm in diameter, thin, entire, yellow; ovary 3-locular, ca.1 mm in diameter, green; styles 0.3–0.6 mm long, stigmas 0.5–1 mm long, apically bifid. Fruits subglobose, shallowly 3-lobed, 4–5 by 4–5 mm in diameter, baccate, fleshy, sweet and tasty, white (to brown); columella 1.3–4 mm long. Seeds 2.2–2.5 by 1.7–1.8 by 1.5–2 mm, smooth, shiny, with a deep hilar invagination.
Distribution — Old World Tropics ( Fig. 3 ) from West Africa to Asia and Australia.
Notes — 1. Some specimens of F. virosa (e.g. NIFS 62, BS (Escritor) 21102, Kjellberg 3072), especially those from drier habitats, exhibit shorter lateral branches than is typical. Specimens with spiny branch ends also occur within F. virosa. Flueggea leucopyrus is another species with spiny branch ends. This species occurs in southern India and Sri Lanka and the small leaved Flueggea specimens in Australia are also identified as such. It might be that F. leucopyrus and F. virosa are identical, but determining this is beyond the scope of this study.
2. The fruit sometimes develops into a gall, with the pedicel and fruit increasing in size and becoming united, and the surface being a bit uneven (var. melanthesoides: Heyligers 1160, Pullen 6817, L.S. Smith 4293, Streimann and Kairo LAE 51645, Widjaja 665, Santos 5717; var. virosa: Koelz 24632, PPI 10285, PPI 22363; PPI 25285, PPI 29152, PPI 29288, PPI 29469, PPI 37488, PPI 38626).
Leaf blades usually more than 5 cm long, apex acute to apiculate, occasionally obtuse; veins coarse, promimulous above, prominently raised below. — New Guinea and Australia
Leaf blades usually less than 5 cm long, apex rounded to obtuse; veins not coarse or prominently raised below. — Africa, Malesia excluding New Guinea
a. subsp. melanthesoides (F.Muell.) G.L.Webster
Flueggea virosa (Roxb. ex Willd.) Royle subsp. melanthesoides (F.Muell.) G.L.Webster, Allertonia 3 (1984) 294; C.Barker & Welzen, Syst. Bot. 35 (2010) 546, Fig. 3. — Leptonema melanthesoides F.Muell., Hooker’s J. Bot. Kew Gard. Misc. 9 (1857) 17 (as ‘melanthesioides’). — Flueggea melanthesoides (F.Muell.) F.Muell., Trans. Bot. Soc. Edinburgh 7 (1863) 490. — Securinega melanthesoides (F.Muell.) Airy Shaw, Kew Bull. 31 (1976) 352; Kew Bull. 35 (1980) 687; Kew Bull., Addit. Ser. 8 (1980) 213. — Type: Mueller s. n.(holotype: K!), Australia, Northern Territory, Victoria River.
Flueggea novoguineensis Valeton [Bull. Dépt. Agric. Indes Néerl. 10 (1907) 26 (as ‘novaguineensis’), nom. nud.] ex Hallier f., Meded. Rijks-Herb. 36 (1918) 5 (as ‘novaguineensis ’). — Lectotype (designated by Webster, 1984: 295): Koch 456 (lectotype: L!), Indonesia, Irian Jaya (Papua), Merauke.
Flueggea virosa (Roxb. ex Willd.) Royle f. reticulata Domin, Biblioth. Bot. 22 (Heft 89 iv) (1927) 878. — Lectotype (designated by Webster, 1984): Domin 5951 (lectotype: PR, n.v., photograph K!), Australia, Queensland, Cloncurry.
Flueggea virosa (Roxb. ex Willd.) Royle var. aridicola Domin, Biblioth. Bot. 22 (Heft 89 iv) (1927) 878. — Securinega melanthesoides (F. Muell.) Airy Shaw var. aridicola (Domin) Airy Shaw, Muelleria 4 (1980) 213; Kew Bull. 35 (1980) 688. — Lectotype (designated by C.Barker & Welzen, 2010, from Webster’s (1984) selection: Domin 5954 (lectotype: PR, n. v., photograph K!), Australia, Queensland, Cloncurry.
Shrubs to small trees, up to 6.5 m high with spreading branches. Outer bark smooth, light grey. Leaves: blades 2–10.5 by 1.2–6.2 cm, generally more than 5 cm long, base attenuate to rounded, apex acute to apiculate, occasionally obtuse; secondary veins coarse, promimulous above, prominently raised below.
Distribution — Australia; Malesia: New Guinea.
l = F. virosa subsp. virosa; ¬ = F. virosa subsp. melanthesoides.
Habitat & Ecology — Lowland rain forest, tropical savannas, edge of monsoon forest, monsoon scrub/deciduous vine thickets, edge of mangrove, and in secondary regrowths; soil: silicious rocks, dry stony soil, clayey loam, and silty sand. Elevation: sea level up to 840 m. Flowering: January to May, August to October; fruiting: January to June.
Vernacular names — Papua (Irian Jaya): Salisari; Papua New Guinea: Djora (Minujfia, Kabubu dialect); Kinuru (Onjob, Naukwate dialect).
Notes — 1. Valeton (1907) only cited the collecting locality of the type of Flueggea novoguineensis Valeton ex Hallier f. Hallier (1918) did not discuss any specimens when he validated Valeton’s name. Valeton’s introduction shows that the type should have been collected by J.W.R. Koch. Two collections in Leiden match the date and locality. Of these, Webster (1984) selected Koch 456 as lectotype. Airy Shaw’s 1976 citation of Versteeg 1884 as the type of Flueggea novoguineensis is incorrect as it was collected in 1934 and could never have been a syntype. His choice probably had to do with a later annotation by Hallier on the sheet.
2. Fluggea virosa is the most widespread species in the genus, with subsp. melanthesoides replacing subsp. virosa in Australasia. However, if F. leucopyrus of Sri Lanka / S. India and Australia is conspecific with F. virosa, then the other subspecies would also occur in Australia.
b . subsp. virosa
Flueggea virosa (Roxb. ex Willd.) Royle subsp. virosa: C.Barker & Welzen, Syst. Bot. 35 (2010) 548, Fig. 3, 4 (see otherwise under the species for type and homotypic synonyms).
Xylophylla obovata Willd., Enum. Hort. Berol. (1809) 329. — Phyllanthus lucidus Hort. ex Steud., Nomencl. Bot., ed. 2, 2 (1841) 327 (non Müll.Arg., 1865: 370). — Securinega obovata (Willd.) Müll. Arg. in DC., Prodr. 15, 2 (1866) 449. — Cicca obovata (Willd.) Kurz, For. Fl. Brit. Burma 2 (1877) 354. — Flueggea obovata (Willd.) Wall. ex Fern.-Vill., Nov. App. (1880) 189, nom. superfl., non Baill. (1861). — Acidoton obovatus Kuntze, Rev. Gen. Pl. 2 (1891) 592. — Type: Berlin Botanical Garden, Cultivated (holotype: B-W, n. v.), Germany, Berlin.
Bessera inermis Spreng., Pl. Min. Cogn. Pug. 2 (1815) 90. — Drypetes bengalensis Spreng., Syst. Veg., ed. 16, 3 (1826) 902, nom. superfl. — Type: Collector not indicated, India, Bengal.
Flueggea microcarpa Blume, Bijdr. (1825) 580. — Securinega microcarpa (Blume) Müll. Arg. in DC., Prodr. 15, 2 (1866) 434. 1— Lectoype (designated by C.Barker & Welzen, 2010): Blume s. n. (holo: L!, photo K; iso: L!), Indonesia, Java.
Phyllanthus angulatus Schumach. & Thonn. in Schumach., Beskr. Guin. Pl. (1827) 415. — Flueggea angulata (Schumach. & Thonn.) Baill., Adansonia 1 (1828) 80 [the reference to Schrank, Syll. Pl. Nov. 2, 1828, 65 is incorrect as it only concerns Phyllanthus obtusus Schrank]. — Type: Thonning s. n. (syntypes: C, 3 sheets), Guinea.
Phyllanthus dioicus Schumach. & Thonn. in Schumach., Beskr. Guin. Pl. (1827) 416. 1827.—Type: Thonning s. n. (syntypes: C, 8 sheets), Guinea.
Phyllanthus obtusus Schrank, Syll. Pl. Nov. 2 (1828) 65. — Type: Collected/grown in Regensburg Botanical Garden, collector not indicated, originally from Guinea.
[Flueggea leucophylla Wall., Numer. List (1829) no. 7916, nom. nud. — Phyllanthus leucophyllus Strachey & Winterb. ex Baill., Étude Euphorb. (1858) 593 (name on plant sheet, as synonym, nom. nud.). — Representative specimen: Herb. Wallich 7916 (K)].
Phyllanthus retusus Roxb., [Hort. Bengal. (1814) 69, nom. nud.] Fl. Ind., ed. 2, 3 (1832) 657 (non Dennst., 1818, nom. nud.). — Type: Hb. Roxburgh (n. v.), India, Bengal.
[Flueggea arborescens Bojer, Hortus Maurit. (1837) 278, nom. nud.]
Cicca pentandra Blanco, Fl. Filip. (1837) 701; Fl. Filip., ed. 2 (1845) 486; Fl. Filip., ed. 3 (1879) 105. — Neotype (C.Barker & Welzen, 2010): Merrill Species Blancoanae 242 (holo: K; iso: L), Philippines, Luzon, Rizal Province, near Mandaloyon. See note.
[Phyllanthus griseus Wall., Numer. List (1847) no. 7814 A (p.p.), nom. nud. —Representative specimen: Herb. Wallich 7918 A (p.p.) (K), Nepal].
Phyllanthus glaucus Wall. [Numer. List (1847) no. 7927 B, nom. nud.] ex Müll.Arg., Linnaea 32 (1863) 14 (non P. glaucus (Müll.Arg.) Müll.Arg. (1865: 386) = Glochidion glaucum Müll. Arg.; nor P. glaucus (F.Muell.) Baill. (1866: 343) = Synostemon glaucus F.Muell.). — Phyllanthus flueggeiformis Müll.Arg. in DC., Prodr. 15, 2 (1866) 349, nom. superfl. (as ‘flüggeiformis’; combination made because Müller thought that the epithet glaucus based on Synostemon glaucus should have precedence, but P. glaucus Wall. ex Müll. Arg. is the oldest use of the epithet glaucus within Phyllanthus).— Type: Herb. Wallich 7927 B (holo: K), Burma.
[Phyllanthus rotundatus (‘rotundata’) Wall., Numer. List (1847) no. 7928 D, nom. nud. (non Poir., 1804). —Representative specimen: Herb. Wallich (Herb. Wight) 7928 D (K).]
Securinega abyssinica A.Rich., Tent. Fl. Abyss. 2 (1850) 256 (incl. the nomen Phyllanthus polygamus Hochst. in synonymy). — Flueggea abyssinica (A.Rich.) Baill., Étude Euphorb. (1858) 593. — Syntypes: Schimper 877 (K, L, 3 sheets, P), Ethiopia (Abyssinica), Dscheladscheranne versus fluvium Tacaze; Schimper 1698 (K, L, 2 sheets, P), Ethiopia (Abyssinica), prope Djeladjeranne.
Flueggea ovalis Baill., Étude Euphorb. (1858) 593. — Type: Riedlé s. n. (n. v.).
[Phyllanthus reichenbachianus Sieber ex Baill., Adansonia 1 (1860) 80, nom. nud. in synonymy of Flueggea angulata Baill. — Representative specimen: Sieber Fl. Seneg. 49 (n. v.), Senegal].
Flueggea senensis Klotzsch in Peters, Naturwissensch. Reise Mossambique 6, 1 (1861) 106. — Type: Collector not indicated (holo: B, destroyed), Mozambique, Sena (named after place of collecting, not to be confused with F. sinensis Baill.).
Diasperus portoricensis Kuntze, Rev. Gen. Pl. 2 (1891) 602. — Phyllanthus portoricensis (Kuntze) Urban, Symb. Antill. 4 (1905) 338. — Conami portoricensis Britton in Britton & P. Wilson, Sci. Surv. Porto Rico & Virgin Islands 5 (1924) 475. — Type: Kuntze 489 (holo: NY), Puerto Rico, Cayey – Guayama (cultivated).
[Phyllanthus virens Wall. ex B. D. Jacks., Index Kew. 2 (1895) 512, a sphalm., misreading of P. virosus Wall., non Müll.Arg. (1866: 415)].
Flueggea obovata (Willd.) Wall. ex Fern.-Vill. var. luxurians Chev. ex Beille, Bull. Soc. Bot. France, Mém. 8 (1908) 55. — Type: A. Chevalier 997 (holo: P), Niger, Tiédiana.
Shrubs to treelets, sometimes somewhat climbing, up to 8 m high, basally up to 9 cm thick, usually deciduous; branchlets laxly spreading, older ones often spiny, brown, often scandent, striated lengthwise. Outer bark (glaucous to) light brown, many lenticels, thin, smooth to slightly cracked to flaky. Leaves: blades 1.3–9 by 0.6–5.6 cm, generally less than 5 cm long; obovate to elliptic, occasionally suborbicular, base cuneate to rounded, apex rounded to obtuse, acute or apiculate at tip; secondary veins not coarse or prominently raised below.
Distribution — Throughout tropical Africa to south central, southeast tropical and subtropical Asia; Malesia: Sumatra, Peninsula Malaysia (Perlis to Pahang), Borneo (Kalimantan), Java, Philippines (Babuyanes to Mindanao ), Celebes, Lesser Sunda Islands (Timor, Flores, Sumbawa), Moluccas (Sula Island).
l = F. virosa subsp. virosa; ¬ = F. virosa subsp. melanthesoides.
Habitat & Ecology — Evergreen and deciduous forests, thickets, waste lands, open swamps, near fields or in abandoned fields, savannah, near streams, along roadsides, along the beach; soil: clay, loam, sand, alluvial; bedrock granite, limestone, or shale. Elevation: Sea level up to 1,000(–2,300) m. Flowering and fruiting throughout the year, most commonly from December to July.
Uses — The bark is used for tanning and to produce a black dye for matting (Burkill 1935, Dict. Econ. Prod. Malay Penin.). It has been described as being strongly astringent and to intoxicate fish when thrown into water, rendering them easier to catch (Roxburgh, 1832). In Thailand, the Karen hill tribe use the roots and leaves against diarrhoea, and for women three days after having given birth; the roots are boiled and drunk like tea, the leaves are boiled and used to bathe, mixed with seven other plants (including turmeric, Blumea balsamifera, Mimosa pudica var. unijuga, Scoparia dulcis).
Vernacular Names — Malay Peninsula: Berbeti; Beti; Beti ayer; Mumbuti (Batek). Sumatra: Mata-mata-ponei; Tjoemata or Tjoemateh (Malay) (Miquel, 1860, Eerste Bijv. ). Java: Chong be. lut; Ime. r; Latian; Pre. bilutan; Sigar jalak; Simpereum (Sundanese); Tre. mbilutan. Philippines: Baohit; Botolan; Baiset; Bis Dlongo, Deongo (Magaspang); Granatang gubat; Sula dugat (Luzon Island); Tulitangalung; Tulugtulug. Celebes: Tangkalasi (Moronene).
Note — Blanco’s specimens were not preserved and no specimens were cited in his description of Cicca pentandra. Webster (Allertonia 3, 1984) did not typify this name in his revision, therefore we have selected Merrill 242 as a neotype. This collection was selected as an illustrative specimen by Merrill (1918) .
Only names of Malesian taxa not or partly treated by Webster (Allertonia 3, 1984) are mentioned here:
Flueggea javanica Blume, Bijdr. (1825) 580. —Type: Blume s. n. (holo: L, barcode L 0813547), Indonesia, Java, Gunung Parang = Streblus taxoides (B. Heyne ex Roth) Kurz (Moraceae). The type specimen was recently detected and reindentified as Streblus taxoides.
Securinega virosa (Roxb. ex Willd.) Baill. var. australiana Baill., Adansonia 6 (1866) 334. — Lectotype (designated by C.Barker & Welzen, 2010): F. Mueller s. n. (K ‘Securinega leucopyra’), Australia, Queensland, Gilbert River. = Flueggea leucopyrus Willd.
Webster (Ann. Missouri Bot. Gard. 81, 1984) reduced this variety to Flueggea virosa (Roxb. ex Willd.) Royle subsp. melanthesoides (F.Muell.) G.L.Webster. Unfortunately, Webster did not annotate the chosen lectotype (Mueller s.n., Australia, Queensland, Gilbert River) at K. There are two specimens at K collected by Mueller at Gilbert River. One, annotated “Leptonema ?”, is of insufficient quality to be determined with certainty, whereas the other, selected as lectotype by us, annotated “Securinega leucopyra”, is clearly different from F. virosa in that it has spiny branches and obovate leaves. Securinega virosa var. australiana is therefore here excluded from synonomy under Flueggea virosa subsp. melanthesoides and forms a synonym of what is known as Flueggea leucopyrus in Australia.