Flueggea (Euphorbiaceae s. l. or Phyllanthaceae) in Malesia

Abstract

Flueggea (including Richeriella) is revised for the Flora Malesiana area. Three species are recognized. Two of them are generally small trees and they are allopatric, F. gracilis (ex-Richeriella) being present in West Malesia and F. flexuosa in East Malesia and the west Pacific. The two subspecies of the generally shrub-like F. virosa are also allopatric, with subsp. virosa having a palaeotropical distribution ending just west of New Guinea, and subsp. melanthesoides being present in New Guinea and Australia.

Full text

Systematic Botany (2010), 35(3): pp. 541–551
© Copyright 2010 by the American Society of Plant Taxonomists
541
Flueggea Willd. was established by Willdenow (1806) based
on F. leucopyrus Willd. from S. India and Sri Lanka. The
generic name appeared as both Flüggea (p. 637) and Fluggea
(p. 757) in Willdenow’s original article, but Article 60.6 of the
International Code of Botanical Nomenclature ( McNeill et al.
2006 : 106) indicates that Flueggea is the correct spelling of the
name. De Jussieu (1824 : 16, 17) accepted Flueggea in his treat-
ment of Euphorbiaceae, observing that the fruits could be
fleshy or dry and two- or three-locular. Baillon (1858 : 590–594)
transferred Phyllanthus L. species with five stamens and a pis-
tillode (including Phyllanthus virosus Wall. [= P. virens Wall. ex
B. D. Jacks., nom. nud.]) into Flueggea and divided the genus
into five sections. He distinguished Flueggea from Securinega
Comm. ex Juss. by the extrorse anthers of the former and the
introrse anthers of the latter. Müller (1866 : 448) considered
Flueggea to be a section of Securinega , but Bentham (1880 : 245)
separated the two genera again on account of differences in
their fruit and seed. Pax and Hoffmann (1922) did not include
Flueggea as part of their revision of Phyllantheae because they
omitted treatment of two of the largest subtribes, Glochidiinae
and Phyllanthinae, Flueggea being included in the latter.
Bentham’s interpretation continued to be followed until Pax
and Hoffmann (1931) resurrected Müller’s interpretation and
placed Flueggea back into Securinega. Webster (1984) provided
evidence that the inclusion of Flueggea in Securinega was erro-
neous. Baillon (1858 : 590–592) had already pointed out a basic
difference between the anthers of the two genera but a palyn-
ological study ( Webster 1984 : 260, 263, 264) showed funda-
mental differences in the morphology of the pollen grains
of the two genera. Pollen of Flueggea has short colpi and a
spinulose exine, whereas that of Securinega has long colpi and
a thin, finely reticulate exine. Furthermore, ovules of Flueggea
are hemitropous whereas those of Securinega are anatropous.
Also, the wood anatomy shows both genera to be distinct, hav-
ing different wood anatomical types ( Mennega 1987 : 123). On
this evidence, Securinega was assigned to a monogeneric sub-
tribe within Phyllantheae by Webster (1994 : 42). It comprises
a small group of species from Madagascar and the Mascarene
Islands. Flueggea was placed in tribe Phyllantheae subtribe
Flueggeinae along with eight other genera (see Webster 1984 :
259–261, 1994; Radcliffe-Smith 2001 ).
Hoffmann et al. (2006) used phylogenetic results to reclas-
sify the infrafamilial taxa within the Phyllanthaceae (formerly
Euphorbiaceae subfamily Phyllanthoideae; family concept
introduced by Savolainen et al. 2000 ). They classified Flueggea
in subfamily Phyllanthoideae tribe Phyllantheae. The phy-
logenetic analyses (e.g. Kathriarachchi et al. 2005 ) on which
they base their conclusions also showed Securinega to be dis-
tinct from Flueggea .
Molecular phylogenetic analyses ( Kathriarachchi et al.
2005 ) placed Richeriella Pax & K. Hoffm. in clade F1 as part of
a polytomy with two species of Flueggea . Webster (1984) was
the first to suggest merging the two genera for which con-
vincing evidence exists. Leaf anatomically ( Levin 1986 ) and
palynologically ( Punt 1962 ; Köhler 1965 ; Sagun and van der
Ham 2003 ) Richeriella is like Flueggea and Margaritaria L. f.;
wood anatomy ( Mennega 1987 ) also places the three genera
together. However, there are morphological differences: long
inflorescences in Richeriella and fasciculate flowers in Flueggea
and Margaritaria , subsessile staminate flowers in Richeriella ,
and seeds with a one-layered exotegmen with ribbon-like
cells in Richeriella ( Stuppy 1995 ; Tokuoka and Tobe 2001 ) and
multilayered with oblong cells in Flueggea and Margaritaria .
Hoffmann et al. (2006) united both genera and extended the
circumscription of Flueggea . Their view is followed here.
Webster (1984) , in his revision of Flueggea , presented the
latest infrageneric classification. He divided the genus into
two sections, sect. Flueggea and sect. Pleiostemon (Sonder)
G. L. Webster. Flueggea sect. Flueggea is further subdivided
into three subsections, subsects. Flueggea , Geblera (Fischer &
Meyer) G. L. Webster, and Acidothamnus G. L. Webster. The
three Malesian species all belong to Flueggea sect. Flueggea
subsect. Flueggea .
This revision encompasses all species found in Malesia
(Malay Archipelago), the triangular area ranging from
Malaysia in the west to the Philippines in the north and Papua
New Guinea in the east.
Taxonomic Treatment
Flueggea Willd., Sp. Pl. 4, 2: 637. 1806; A. Juss., Euphorb. Gen.:
16. 1824; Blume, Bijdr.: 580. 1825; Baill., Étude Euphorb.:
590. 1858; Benth. & Hook. f., Gen. Pl. 3: 276. 1880; Hook.f.,
Fl. Brit. India 5: 328. 1887; Beille in Lecomte, Fl. Indo-
Chine 5: 528. 1927; G. L. Webster, Allertonia 3: 273. 1984;
Ann. Missouri Bot. Gard. 81: 43. 1994; Welzen, Thai Forest
Bull. (Bot.) 28: 56. 2000; Radcl.-Sm., Gen. Euphorb.: 35.
2001; C. Barker & Welzen in Chayamarit & Welzen, Fl.
Thailand 8, 1: 299. 2005. Securinega Comm. ex Juss. sect.
Flueggea (Willd.) Müll. Arg. in DC., Prodr. 15, 2: 448. 1866;
Pax & K. Hoffm. in Engl. and Harms, Nat. Pflanzenfam.
ed. 2, 19c: 60. 1931.—TYPE: Flueggea leucopyrus Willd.
Flueggea (Euphorbiaceae s. l. or Phyllanthaceae) in Malesia
Christine Barker 1 , 3 and Peter C. van Welzen 2
1 Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, U. K.
2 National Herbarium of the Netherlands, Leiden University Branch, P.O. Box 9514, 2300 RA Leiden, The Netherlands
3 Author for correspondence (c.barker@kew.org)
Communicating Editor: Daniel Potter
Abstract—Flueggea (including Richeriella ) is revised for the Flora Malesiana area. Three species are recognized. Two of them are generally
small trees and they are allopatric, F. gracilis (ex- Richeriella ) being present in West Malesia and F. flexuosa in East Malesia and the west Pacific.
The two subspecies of the generally shrub-like F. virosa are also allopatric, with subsp. virosa having a palaeotropical distribution ending just
west of New Guinea, and subsp. melanthesoides being present in New Guinea and Australia.
Keywords— Euphorbiaceae s. l. , Flueggea , Malesia , Phyllanthaceae , Richeriella , Securinega .

542 SYSTEMATIC BOTANY [Volume 35
Acidoton P. Browne, Civ. Nat. Hist. Jamaica: 355. 1756, 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: 90. 1815.—TYPE:
Bessera inermis Spreng. [= Flueggea virosa (Roxb. ex Willd.)
Royle subsp. virosa ].
Geblera Fisch. & C. A. Mey., Index Sem. Hort. Petrop. 1: 28.
1835.—TYPE: Geblera suffruticosa (Pall.) Fisch. & C. A.
Mey. [= Flueggea suffruticosa (Pall.) Baill.].
Colmeiroa Reut., Biblioth. Universelle Genève 38: 215. 1842.—
TYPE: Colmeiroa buxifolia Reut. [= Flueggea tinctoria (L.)
G. L. Webster].
Pleiostemon Sond., Linnaea 23: 135. 1850.—TYPE: Pleiostemon
verrucosum (Thunb.) Sond. [= Flueggea verrucosa (Thunb.)
G. L. Webster.]
Neowawraea Rock, Indig. Trees Haw. Isl.: 243. 1913.—TYPE:
Neowawraea phyllanthoides Rock [= Flueggea neowawraea
W. J. Hayden].
Richeriella Pax & K. Hoffm. in Engl., Pflanzenr. IV.147.xv: 30.
1922; Merr., Enum. Philipp. Fl. Pl.: 411. 1923; Airy Shaw,
Kew Bull. 25: 489, 491, 492. 1971; Kew Bull. 26: 328. 1972;
Whitmore, Tree Fl. Malaya 2: 128. 1973; Airy Shaw, Kew
Bull., Addit. Ser. 4: 190. 1975; Alphab. Enum. Euphorb.
Philipp. Isl.: 43. 1983; G. A. Levin, Ann. Missouri Bot. Gard.
73: 75. 1986; Mennega, Bot. J. Linn. Soc. 94: 123, Table 2.
1987; G. L. Webster, Ann. Missouri Bot. Gard. 81: 44. 1994;
Stuppy, Syst. Morph. Anat. Samen biovul. Euphorb.: 157,
162. 1995; Radcl.-Sm., Gen. Euphorb.: 36. 2001; T. Tokuoka &
Tobe, J. Plant Res. 144: 75. 2001; Sagun & R. W. Ham,
Grana 42: 205, Fig. 7 e–j, Table 2. 2003; Kathriarachchi
et al. Mol. Phylog. Evol. 36: 127, Figs. 1 , 3 , 4 . 2005;
Welzen in Welzen & Chayamarit, Fl. Thailand 8, 2: 515.
2007.—TYPE: Richeriella gracilis (Merr.) Pax & K. Hoffm.
[= Flueggea gracilis (Merr.) Petra Hoffm.]
Securinega auct. non Comm. ex Juss.: Backer & Bakh.f., Fl. Java
1: 465. 1963; Airy Shaw, Kew Bull. 26: 340. 1972; Whitmore,
Tree Fl. Malaya 2: 131. 1973; Airy Shaw, Kew Bull., Addit.
Ser. 4: 196. 1975; Kew Bull. 35: 687. 1980; Kew Bull., Addit.
Ser. 8: 200. 1980; Kew Bull. 36: 345. 1981; Alphab. Enum.
Euph. Philip. Isl.: 45. 1983.
Shrubs or trees, usually dioecious, branching monopodial
plagiotropic. Indumentum generally absent. Stipules triangu-
lar, 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 actino-
morphic, 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, dehisc-
ing extrorsely with longitudinal slits; pistillode consisting of
2 or 3 large strap-like filaments, apically bent. Pistillate flow-
ers: pedicels long, slender, enlarging in fruit; disc flat, annu-
lar, 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 reticu-
late 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 and 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.
Key to the Species of Malesian F LUEGGEA
1. 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 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. F. virosa
1. 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 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Inflorescences fascicles. Fruits 3.5–4 mm in diameter—Philippines (not Palawan) to New Guinea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. F. flexuosa
2. 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. F. gracilis
1. Flueggea flexuosa Müll. Arg., Linnaea 34: 76. 1865;
C. B. Rob., Philipp. J. Sci. 4, Bot.: 74. 1909; G. L. Webster,
Allertonia 3: 284. 1984. Securinega flexuosa (Müll. Arg.)
Müll. Arg. in DC., Prodr. 15, 2: 450. 1866; Merr., Philipp.
J. Sci. 9, Bot.: 491. 1914; Enum. Philipp. Fl. Pl. 2: 290.
1923; Whitmore, Guide For. Brit. Solomon Is.: 151, 201.
1966; Airy Shaw, Kew Bull., Addit. Ser. 8: 201. 1980;
Alphab. Enum. Euph. Philip. Isl.: 45. 1983. Acidoton
flexuosus Kuntze, Rev. Gen. Pl. 2: 592. 1891.—TYPE:
PHILIPPINES. Bohol, s. d., Cuming 1855 (holotype:
G-DC; isotype: K!).
Phyllanthus acuminatissimus C. B. Rob., Philipp. J. Sci. 3, Bot.:
200. 1908. Securinega acuminatissima (C. B. Rob.) C. B. Rob.,
Philipp. J. Sci. 4, Bot.: 73. 1909.—TYPE: PHILIPPINES.
Mindanao, Dist. of Davao, Santa Cruz, s. d., R. S. Williams
2807 (holotype: NY; isotypes: K!, NY).
Securinega samoana Croizat, Bull. Bernice P. Bishop Mus. 184: 45.
1945.—TYPE: SAMOA. Savaii, Tufutafoe-Falelima, 30 Sep
1931, E. Christophersen 2765 (holotype: NY; isotypes: K!, P).
Shrub-like trees to trees, evergreen, up to 17 m high, bole
10 m, dbh up to 33 cm; buttresses occasionally present, up to

2010] BARKER AND VAN WELZEN: FLUEGGEA IN MALESIA 543
F ig . 1. Distribution of Flueggea flexuosa Müll. Arg. (circle) and F. gracilis (Merr.) Petra Hoffm. (star).
1 m high, thin, equal; twigs terete to angled, smooth to stri-
ate, flexuous, often weakly so, occasionally lenticellate; flow-
ering 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; sap-
wood white to pale brown, soft to usually hard, heartwood
hard, red to pale brown. Stipules 1–1.3 × 1–1.7 mm, acumi-
nate, caducous. Leaves: petiole 5–11 mm long, channelled
above; blade elliptic to oblong, 6–18.5 × 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, vena-
tion 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 × 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 × 0.7–0.9 mm, inner 3 larger, obovate-oblong, 1.3–1.7 ×
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 ellip-
soid, 0.4–0.6 × 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; pedi-
cel 3.5–7.7 mm long; sepals 5, apex broadly rounded, outer 2
oblong, 1.2–1.5 × 0.8–1.1 mm, inner 3 larger, broadly ovate to
orbicular, 1.2–1.6 × 1.0–1.5 mm; disc flat, thick, 1–1.6 mm in
diameter; ovary 3(4)-locular, 1–1.5 × 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 per-
sistent, dehiscing irregularly; columella early caducous. Seeds
obovoid-triangular, 2–2.5 × c. 1.5 × 1.3 mm, with a broad, shal-
low 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). Figure 1 .
Habitat and Ecology— The species is present in lowland
primary and secondary rainforests, kaingin (shifting cultiva-
tion) fields, often along rivers; soil: loam, clay, limestone. Alt.:
up to 500 m.
Phenology— 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 con-
struction, fencing and sometimes for heavy construction and
bridging. In the Reef Islands it has been used for buildings

544 SYSTEMATIC BOTANY [Volume 35
that stand in the sea as the wood is said to be durable in salt
water ( Henderson and Hancock 1988 : 190–192). On Vanuatu
the bark and leaves are used medicinally, the latter are
boiled and rubbed over the body for curing scabies, espe-
cially 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. Solomon Islands: Mamufu’a or Mamufua
(Kwara’ae). Vanuatu: Mamau (Butu); Nemameiu (Hiu). Fiji:
Pou. Samoa: Poumuli ( Croizat 1945 ). Wallis and Futuna
(Horne Island): Poutea.
Notes—Flueggea flexuosa is clearly related to the wide-
spread 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.
Representative Specimens Examined— FIJI. Rotuma: Itutiu Distr.,
Kilinga, 12 Aug 1938, St. John 19468 (L). INDONESIA. Maluku: Morotai,
Totodoku, 28 Apr 1949, Kostermans 7815 (L). Papua: Vogelkop, Warnapi, N
of Ransiki, 80 km S of Manokwari, Aug 1948, Kostermans 4758 (L). NEW
ZEALAND. Niue: Horn Island, Alofi, 26 Oct 1968, McKee 19784 (L). PAPUA
NEW GUINEA. Bougainville: near Aku village, 10 miles W of Buin, 24 Sep
1964, Craven & Schodde 530 (L). PHILIPPINES. Catanduanes: Brgy. Si-ay,
San Miguel, 26 Nov 1991, Barbon, Alvarez & Garcia PPI 5526 (L). Cebu:
Gahong (Puti), Ipil, Carmen, 10 Jun 1997, Gaerlan & Reynoso PPI 20232 (L).
Gamaguin Island: Mambajao, Nov 1912, Elmer 14244 (L). Leyte: Palo, Jan
1906, Elmer 7280 (L). Luzon: Albay Prov., Guinobatan, Banao, Jun 1953,
Mendoza PNH 18588 (L); Sorsogon Prov., Irosin (Mt. Bulusan), Nov 1915,
Elmer 15218 (L). Mindanao: Agusan Prov., Cabadbaran, Aug 1912, Elmer
13455 (L). SAMOA. Upolu: Lefaga, Safa’atoa, 14 May 1968, Bristol 2100 (L).
SOLOMON ISLANDS. Big Nggela: N of Haleta village, 13 Jun 1969, Gafui
et al. BSIP 15040 (L). Choiseul: Dalepima Area, Mbirambira, 15 Jan 1970,
Gafui et al. BSIP 18820 (L). Fauro: 29 Apr 1964, Whitmore BSIP 4196 (L). Gizo
(Giso): 8 Jul 1968, Mauriasi et al. BSIP 11707 (L). Guadalcanal: Nalimbu
River, 3 miles inland, 6 Feb 1964, Whitmore BSIP 2550 (L); Forest adjacent
to Tina River, 12 miles inland from the coast, 15 Nov 1962, Womersley &
Whitmore BSIP 1154 (L). Kolombangara: Sandfly Harbour, 20 Feb 1963,
Whitmore BSIP 1502 (L). Malaita: near Bitima Anchorage, 27 Aug 1945,
C. T. White BSIP 104 (L). Malauppaina: Three Sisters, 9 Dec 1969, Mauriasi
et al. BSIP 17998 (L). Mbagha: Baga, 4 Feb 1964, Whitmore’s Collectors BSIP
2936 (L). Mono: Palusua, 1 May 1969, Mauriasi et al. BSIP 14179 (L). New
Georgia Islands: Guruva Island, 22 May 1964, Whitmore’s Collectors BSIP
3742 (L); Hamarai Island, 22 Oct 1964, Cowmeadow’s collectors BSIP 3202
(L); New Georgia, Mango River, 21 Jun 1965, Maenu’u BSIP 6049 (L).
Ranongga: Viri upper area, 1 Jul 1969, Mauriasi et al. BSIP 15731 (L). Reef
Islands: Nanienubuli, 5 Feb 1965, Inimua BSIP 6531 (L). Rob Roy: 9 Mar
1964, Whitmore’s Collectors BSIP 5331 (L). San Cristobal: Hotabila River,
Mara Bay, 27 Nov 1968, Gafui et al. BSIP 12831 (L). Santa Cruz: Mbania, 9
Oct 1969, Mauriasi et al. BSIP 17625 (L). Santa Isabel: Gorona, 21 Dec 1932,
Brass 3372 (L). Shortland: SE of Koang Kopi River, 24 Feb 1969, Runikera et al.
BSIP 13067 (L). Small Malaita: N of Palasu’u school, 29 Sep 1969, Gafui
et al. BSIP 17304 (L). Small Nggela: W of Kombe village, 3 Jul 1969, Gafui
et al. BSIP 15282 (L). Treasury Island: Maloaini, 3 Jun 1969, Mauriasi et al.
BSIP 14268 (L). Ulawa: Suumoli Plantation, 4 Feb 1965, Teona BSIP 6206
(L). Vaghena: Wagina, 20 Mar 1964, Whitmore’s Collectors BSIP 5531 (L).
VANUATU. Hiu: Yageugemona, garden area above village, 22 Oct 1993,
Kalomor 63 (L). Iles Torres, Tegua, 27 Nov 2007, Pillon 1132 (L).
2. Flueggea gracilis (Merr.) Petra Hoffm. in Petra Hoffm.,
Kathriar. & Wurdack, Kew Bull. 61: 44. 2006. Baccaurea
gracilis Merr., Philipp. J. Sci. 1, Suppl.: 203. 1906. Richeriella
gracilis (Merr.) Pax & K. Hoffm. in Engl., Pflanzenr.
IV.147.xv: 30. 1922; Merr., Enum. Philipp. Fl. Pl.: 411.
1923; Lingnan Sci. J. 14: 22. 1935; Punt, Wentia 7: 26. 1962;
Eg. Köhler, Grana Palyn. 6: 45, 72, 100, t. 4: fig. 5–8. 1965;
Airy Shaw, Kew Bull. 25: 489. 1971; Kew Bull. 26: 328.
1972; Alphab. Enum. Euphorb. Philipp. Isl.: 43. 1983;
Welzen in Welzen & Chayamarit, Fl. Thailand 8, 2: 515,
Fig. 68. 2007.—TYPE: PHILIPPINES. Palawan, Dec 1905,
Bermejos BS 218 (lectotype, here designated: K!; isolecto-
type: US). PHILIPPINES, Palawan, Dec 1905, Bermejos
BS 258 (syntype: K); PHILIPPINES, Palawan, Mar 1906,
Curran FB 3828 (syntypes: K!, US).
Richeriella malayana M. R. Hend., Gard. Bull. Straits Settlem.
7: 122, Pl. 32. 1933; J. Malayan Branch Roy. As. Soc. 17:
71. 1939; Airy Shaw, Kew Bull. 25: 489, 491, 492. 1971;
Whitmore, Tree Fl. Malaya 2: 128. 1973; Airy Shaw,
Hooker’s Icon. Pl. 38: t. 3703. 1974; Kew Bull., Addit.
Ser. 4: 190. 1975.—TYPE: MALAYSIA. Perak, Gunong
Pondok, 7 Jun 1930, M. R. Henderson SFN 23790 (holo-
type: SING; isotypes: K!, 2 sheets, SING).
Richeriella malayana M. R. Hend. var. macrocarpa Airy Shaw,
Kew Bull. 25: 490. 1971; Kew Bull., Addit. Ser. 4: 190.
1975.—TYPE: MALAYSIA. Sarawak, Kuching dist., Tiang
Bekap, 5 Jun 1960, J. A. R. Anderson S 12542 (holotype: K!;
isotypes: L!, SAR).
Shrubs to trees, evergreen, up to 20 m high, dbh up to 15 cm;
bole sometimes fluted; flowering branchlets 2–3 mm in diam-
eter, 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 × 1–2.2 mm,
stiff, caducous. Leaves: petiole 3–9 mm long, flattened above;
blade elliptic-oblong to slightly obovate, 6.7–23 × 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 acumi-
nate (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 ×
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 × 1 mm, 2 or 3, inner ones larger, 1.2–1.3 × 1.2–
1.3 mm; disc lobes 5, c. 0.2 × 0.2 mm; stamens 5, filaments 1.7–3
mm long, white, anthers ellipsoid, 0.5–0.8 × 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 × 1.1–1.4
mm, 3 inner ones c. 1.5 × 1.5 mm; ovary 3(4)-locular. Fruits
lobed, dehiscent, 8–12 × 4.5–7 mm, black, slightly succulent,
somewhat wrinkled when dry; columella 2.4–4 mm long.
Seeds obovoid, 3–5.5 × 2.4–4.7 × 1.8–4.5 mm, with a broad,
shallow hilar invagination. Figure 2 .
Distribution— China (Hainan), Peninsular Thailand, Malay
Peninsula, Borneo, Philippines (Palawan). Figure 1 .
Habitat and Ecology— Primary and secondary rainfor-
est, open shrubbery; soil: sandy clay, sandy beach, limestone
rocks with loam cover. Elevation: Sea level up to 333 m.
Phenology— Flowering and fruiting probably all year round.
Vernacular names— Borneo, Sarawak: Mabon (Land
Dayak). Philippines: Marabakawan (Palawan).
Notes— The long inflorescences instead of fascicles are a
unique character for F. gracilis . The names Richeriella graci-
lis , 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

2010] BARKER AND VAN WELZEN: FLUEGGEA IN MALESIA 545
Fig. 2.Flueggea gracilis (Merr.) Petra Hoffm. a. Habit. b. Inflorescence with fruit remnants (columellas and sepals) and buds. c. Staminate flower.
d. Pistillate flower. e. Fruit. f. Seeds. [a: H. Y. Liang s. n., L 0137873; b, f: Edaño PNH 239; c: Curran FB 4123; d, e: Anderson S 20979; all L]. Drawing by
Hanneke Jelles (L), 2006.

546 SYSTEMATIC BOTANY [Volume 35
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) hav-
ing smaller inflorescences with fewer branches. Uniting both
species creates a continuous distribution. Specimens iden-
tified as Richeriella malayana var. macrocarpa , known from
Sarawak and part of Indonesian Kalimantan, have larger
fruits than the typical F. gracilis specimens (11–12 × 6–7 mm
versus 8–9.5 × 4.5–6 mm), but the difference is not impres-
sive. 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. spe-
cies show a comparable variation ( Alchornea rugosa (Lour.)
Müll. Arg., Austrobuxus nitidus Miq., Cheilosa montana Blume).
Therefore, the name is also synonymised with F. gracilis .
Representative Specimens Examined— CHINA. Hainan: 24 Feb 1934,
Liang H.Y. s. n. (L). INDONESIA. Kalimantan Tengah: Tumbang Tapi,
40 km N of Tumbang Samba, on the Samba River near base camp of the
Handyani Timber Comp., 18 Nov 1982, Mogea 3535 (L). Kalimantan Timur:
W Koetai, Kombeng, 22 Nov 1925, Endert 5139 (L). MALAYSIA. Sabah,
Tongod, Sungai Enodol, 23 Jul 1983, Sundaling SAN 99460 (L). Sarawak:
First Division, Kuching Distr., Tiang Bekap, 5 Jun 1960, J. A. R. Anderson S
12542 (L); Second Division, 21st mile Simanggang Road, 5 Sep 1966, J. A.
R. Anderson S 24795 (L). Terengganu: Batu Biwa, 10 Jun 1968, Cockburn FRI
10561 (L). PHILIPPINES. Palawan: Lipuun Island, 28 Apr 1964, Mendoza &
Cordero Jr. PNH 91341 (L); Quezon, Tawa-Tawa, 1 Feb 1994, Gaerlan, Romero &
Sagcal PPI 13434 (L); Tagduan, S of Puerto Princesa on E coast, 26 Jan 1991,
Reynoso, Romero & Sagcal PPI 240 (L).
3. Flueggea virosa (Roxb. ex Willd.) Royle, Ill. Bot. Himal.
Mts.: 328. 1836; Voigt, Hort. Suburb. Calcut.: 152. 1845;
Baill., Étude Euphorb.: 593. 1858; C. B. Rob., Philipp. J.
Sci. 4, Bot.: 74. 1909; J. J. Sm., Meded. Depart. Landbouw
10: 51. 1910, Merr., Enum. Philipp. Fl. Pl.: 390. 1923; G. L.
Webster, Allertonia 3: 287. 1984; C. Barker & Welzen in
Chayamarit & Welzen, Fl. Thailand 8, 1: 301, Fig. 75, Plate
XVI: 4. 2005. Phyllanthus virosus Roxb. ex Willd., Spec.
Pl. 4, 1: 578. 1805; Roxb., Fl. Ind. 3: 659. 1832; Wall., Numer.
List: 7928 A, B, C (Goyalpara), E. 1847. Securinega virosa
(Roxb. ex Willd.) Baill., Adansonia 6: 334. 1866; Backer &
Bakh.f., Fl. Java 1: 466. 1963; Airy Shaw, Kew Bull. 26:
340. 1972; Whitmore, Tree Fl. Malaya 2: 131. 1973; Airy
Shaw, Kew Bull., Addit. Ser. 4: 196 (excl. New Guinea and
Australia). 1975; Kew Bull. 36: 345. 1981; Alphab. Enum.
Euph. Philip. Isl.: 45. 1983. Acidoton virosus (Roxb. ex
Willd.) Kuntze, Rev. Gen. Pl. 2: 592. 1891.—TYPE: INDIA.
Nandaradah, Klein 748 (holotype: B-W!).
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 × 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 ×
0.6–6.2 cm, length/width ratio 1.6–2.3, base cuneate to
rounded and decurrent with the petiole, sometimes asym-
metric, 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 tri-
angular, small. Staminate flowers 20–40 per axillary cluster,
1.7–2.2 mm in diameter, white to pale green to yellow to suf-
fused 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 × 0.5–0.7 mm, the inner 3
broadly ovate, 1.6–2 × 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 ×
0.3–0.4 mm, cream to light yellow, connective maroon; pistil-
lode (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 clus-
ter, 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 mar-
gins to yellowish; disc 0.7–1 mm in diameter, thin, entire, yel-
low; 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 sub-
globose, shallowly 3-lobed, 4–5 × 4–5 mm in diameter, baccate,
fleshy, sweet and tasty, white (to brown); columella 1.3–4 mm
long. Seeds 2.2–2.5 × 1.7–1.8 × 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— 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 iden-
tified as such. It might be that F. leucopyrus and F. virosa are
identical, but determining this is beyond the scope of this
study. The fruit sometimes develops into a gall, with the pedi-
cel 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 ).
Key to the Subspecies of F LUEGGEA VIROSA
1. 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 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3a. F. virosa subsp. melanthesoides
1. Leaf blades usually less than 5 cm long, apex rounded to obtuse; veins not coarse or prominently
raised below. —Africa, Malesia excluding New Guinea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3b. F. virosa subsp. virosa
3a. Flueggea virosa (Roxb. ex Willd.) Royle subsp. melan-
thesoides (F. Muell.) G. L. Webster, Allertonia 3: 294.
1984. Leptonema melanthesoides F. Muell., Hooker’s J.
Bot. Kew Gard. Misc. 9: 17 (as ‘ melanthesioides ’). 1857.
Flueggea melanthesoides (F. Muell.) F. Muell., Trans. Bot.
Soc. Edinburgh 7: 490. 1863. Securinega melanthesoides
(F. Muell.) Airy Shaw, Kew Bull. 31: 352. 1976; Kew Bull.
35: 687. 1980; Kew Bull., Addit. Ser. 8: 213. 1980.—TYPE:
AUSTRALIA. Northern Territory, Victoria River, s. d.
Mueller s. n. (holotype: K!).

2010] BARKER AND VAN WELZEN: FLUEGGEA IN MALESIA 547
Flueggea novoguineensis Valeton [Bull. Dépt. Agric. Indes Néerl.
10: 26 (as ‘novaguineensis ’), nom. nud . 1907] ex Hallier f.,
Meded. Rijks-Herb. 36: 5 (as ‘ nova-guineensis ’). 1918.—
TYPE: INDONESIA. Irian Jaya (Papua), Merauke, 1904,
Koch 456 (lectotype, selected by Webster 1984 : 295: L!).
Flueggea virosa (Roxb. ex Willd.) Royle f. reticulata Domin,
Biblioth. Bot. 22 (Heft 89 iv): 878. 1927.—TYPE:
AUSTRALIA. Queensland, Cloncurry, s. d., Domin 5951
(lectotype selected by Webster (1984) : PR, n. v., photo-
graph K!).
Flueggea virosa (Roxb. ex Willd.) Royle var. aridicola Domin,
Biblioth. Bot. 22 (Heft 89 iv): 878. 1927. Securinega melan-
thesoides (F. Muell.) Airy Shaw var. aridicola (Domin) Airy
Shaw, Muelleria 4: 213. 1980; Kew Bull. 35: 688. 1980.—
TYPE: AUSTRALIA, Queensland, Cloncurry, s. d., Domin
5954 (lectotype selected here from Webster’s (1984) selec-
tion: PR, n. v., photograph K!).
Shrubs to small trees, up to 6.5 m high with spreading
branches. Outer bark smooth, light grey. Leaves: blades 2–10.5 ×
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. Figure 3 .
Habitat and Ecology— Lowland rain forest, tropical savan-
nas, 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.
Phenology— 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— 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 syn-
type. His choice probably had to do with a later annotation by
Hallier on the sheet. Fluggea virosa is the most widespread spe-
cies 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.
Representative Specimens Examined— AUSTRALIA. Northern
Territory: 4 miles NNW El Sharana, UDP Falls Road, 18 Jan 1973, Martensz &
Schodde AE 425 (L); 13 miles NE of Katherine, 7 Mar 1965, I. B. Wilson 380 (L).
Queensland: Cook Dist., Barkly Highway, 4 km N of Spear Creek,
F ig . 3. Distribution of Flueggea virosa (Roxb. ex Willd.) Royle subsp. melanthesoides (F. Muell.) G. L. Webster (square) and Flueggea virosa (Roxb. ex
Willd.) Royle subsp. virosa (circle).

548 SYSTEMATIC BOTANY [Volume 35
12 km N of Mt. Isa (by air), 31 Jan 1995, A. Fraser 91 (L). INDONESIA. Irian
Jaya/Papua: Merauke, 14 Aug 1904, Koch 457 (L). PAPUA NEW GUINEA.
Central Prov.: Port Moresby Subdist., Waigani, Jul 2000, Pulsford UPNG
296 (L). Gulf Prov.: Yule, 16 Jan 1955, Womersley & Simmonds NGF 5096 (L).
Northern: Tufi subprov., ca. 3 miles from Aku on Kuruaku track, 9 Aug
1954, Saunders 29 (L). Western Prov.: Daru Island, 20 Aug 1967, Ridsdale
NGF 33751 (L).
3 b . Flueggea virosa (Roxb. ex Willd.) Royle subsp. virosa (See
under the species for type and homotypic synonyms).
Xylophylla obovata Willd., Enum. Hort. Berol.: 329. 1809.
Phyllanthus lucidus Hort. ex Steud., Nomencl. Bot., ed. 2,
2: 327. 1841; non Müll. Arg. (1865: 370). Securinega obo-
vata (Willd.) Müll. Arg. in DC., Prodr. 15, 2: 449. 1866.
Cicca obovata (Willd.) Kurz, For. Fl. Brit. Burma 2: 354.
1877. Flueggea obovata (Willd.) Wall. ex Fern.-Vill., Nov.
App.: 189, nom. superfl., non Baill. (1861). 1880. Acidoton
obovatus Kuntze, Rev. Gen. Pl. 2: 592. 1891.—TYPE:
GERMANY. Berlin, Botanical Garden, Cultivated (holo-
type: B-W, n. v.).
Bessera inermis Spreng., Pl. Min. Cogn. Pug. 2: 90. 1815. Drypetes
bengalensis Spreng., Syst. Veg., ed. 16, 3: 902, nom. superfl.
1826.—TYPE: INDIA. Bengal, collector not indicated.
Flueggea microcarpa Blume, Bijdr.: 580. 1825. Securinega micro-
carpa (Blume) Müll. Arg. in DC., Prodr. 15, 2: 434. 1866.—
TYPE: INDONESIA. Java, Blume s. n. (lectotype, selected
here: L!, photo K; isolectotype: L!).
Phyllanthus angulatus Schumach. & Thonn. in Schumach.,
Beskr. Guin. Pl.: 415. 1827. Flueggea angulata (Schumach. &
Thonn.) Baill., Adansonia 1: 80 [the reference to Schrank,
Syll. Pl. Nov. 2, 1828, 65 is incorrect as it only concerns
Phyllanthus obtusus Schrank]. 1860.—TYPE: GUINEA.
Thonning s. n. (syntypes: C!, 3 sheets).
Phyllanthus dioicus Schumach. & Thonn. in Schumach., Beskr.
Guin. Pl.: 416. 1827.—TYPE: GUINEA, Thonning s. n.
(syntypes: C!, 8 sheets).
Phyllanthus obtusus Schrank, Syll. Pl. Nov. 2: 65. 1828.—TYPE:
GUINEA, collected/grown in Regensburg Botanical
Garden, collector not indicated.
[ Flueggea leucophylla Wall., Numer. List: no. 7916, nom. nud.
1829. Phyllanthus leucophyllus Strachey & Winterb. ex
Baill., Étude Euphorb.: 593 (name on plant sheet, as syn-
onym, nom. nud.). 1858.—Representative specimen:
Herb. Wallich 7916 (K!)].
Phyllanthus retusus Roxb., [Hort. Bengal.: 69, nom. nud. 1814]
Fl. Ind., ed. 2, 3: 657, non Dennst. (1818, nom. nud.).
1832.—TYPE: INDIA. Bengal, Hb. Roxburgh (n. v.).
[ Flueggea arborescens Bojer, Hortus Maurit.: 278, nom. nud.
1837.]
Cicca pentandra Blanco, Fl. Filip.: 701. 1837; Fl. Filip., ed. 2: 486.
1845; Fl. Filip., ed. 3: 105. 1879.—TYPE: PHILIPPINES.
Luzon, Rizal Province, near Mandaloyon, 18 Apr 1914,
Merrill Species Blancoanae 242 (neotype, selected here: K!;
isoneotype: L!). See note.
[ Phyllanthus griseus Wall., Numer. List: no. 7814 A (p.p.), nom.
nud. 1847.—Representative specimen: NEPAL. Herb.
Wallich 7918 A (p.p.) (K!)]
Phyllanthus glaucus Wall. [Numer. List: no. 7927 B, nom. nud.
1847] ex Müll. Arg., Linnaea 32: 14. 1863 (non P. glaucus
(Müll. Arg.) Müll. Arg. (1865a: 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: 349, nom. superfl. (as ‘flüggei-
formis ’). 1866. (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: BURMA. Herb. Wallich 7927 B (holotype: K!).
[ Phyllanthus rotundatus ( ‘rotundata ’) Wall., Numer. List: no.
7928 D, nom. nud., non Poir. (1804). 1847.—Representative
specimen: Herb. Wallich (Herb. Wight) 7928 D (K!).]
Securinega abyssinica A. Rich., Tent. Fl. Abyss. 2: 256 (incl.
the nomen Phyllanthus polygamus Hochst. in synon-
ymy). 1850. Flueggea abyssinica (A. Rich.) Baill., Étude
Euphorb.: 593. 1858.—TYPE: ETHIOPIA (Abyssinica).
Dscheladscheranne versus fluvium Tacaze, 4 Jun 1840,
Schimper 877 (syntypes: K!, L!, 3 sheets, P); ETHIOPIA
(Abyssinica). Prope Djeladjeranne, 31 Jul 1840, Schimper
1698 (syntypes: K!, L!, 2 sheets, P).
Flueggea ovalis Baill., Étude Euphorb.: 593. 1858.—TYPE: Riedlé
s. n. (n. v.).
[ Phyllanthus reichenbachianus Sieber ex Baill., Adansonia 1:
80, nom. nud. in synonymy of Flueggea angulata Baill.
1860.—Representative specimen: SENEGAL. Sieber Fl.
Seneg. 49 (n. v.)].
Flueggea senensis Klotzsch in Peters, Naturwissensch. Reise
Mossambique 6, 1: 106. 1861. —TYPE: MOZAMBIQUE.
Sena (named after place of collecting, not to be confused
with F. sinensis Baill.), collector not indicated (holotype:
B, destroyed).
Diasperus portoricensis Kuntze, Rev. Gen. Pl. 2: 602. 1891.
Phyllanthus portoricensis (Kuntze) Urban, Symb. Antill.
4: 338. 1905. Conami portoricensis Britton in Britton &
P. Wilson, Sci. Surv. Porto Rico & Virgin Islands 5: 475.
1924.—TYPE: PUERTO RICO. Cayey – Guayama (culti-
vated), 14 Mar 1874, Kuntze 489 (holotype: NY!).
[ Phyllanthus virens Wall. ex B. D. Jacks., Index Kew. 2: 512.
1895, 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: NIGER. Tiédiana, 18 Jun 1899, A. Chevalier 997
(holotype: P).
Shrubs to treelets, sometimes somewhat climbing, up to
8 m high, basally up to 9 cm thick, usually deciduous; branch-
lets 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 × 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 apicu-
late at tip; secondary veins not coarse or prominently raised
below. Figure 4 .
Distribution— Throughout tropical Africa ( Fig. 3 ) 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).

2010] BARKER AND VAN WELZEN: FLUEGGEA IN MALESIA 549
F ig . 4. Flueggea virosa (Roxb. ex Willd.) Royle subsp. virosa . a. Habit. b. Staminate flower. c. Pistillate flower. d. Pistillate flower with part of sepals
removed. e. Fruit. f. Seed, ventral view. g. Seed, lateral view. [a, b: Barnes 1109 ; c, d: Maxwell 87–668 ; e: Murata, Fukuoka & Phengkhlai T-16569 ; f, g: Kostermans
Khwae 1257 ; all L]. Drawing by Jan van Os (L), 2003.

550 SYSTEMATIC BOTANY [Volume 35
Habitat and Ecology— Evergreen and deciduous forests,
thickets, waste lands, open swamps, near fields or in aban-
doned fields, savannah, near streams, along roadsides, along
the beach; soil: clay, loam, sand, alluvial; bedrock granite, lime-
stone, or shale. Elevation: Sea level up to 1,000(–2,300) m.
Phenology— 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 ). 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 (includ-
ing turmeric, Blumea balsamifera , Mimosa pudica var. unijuga ,
Scoparia dulcis ).
Vernacular Names— Ethiopia: Harmaso. Comores:
Nhambah. Burma: Pakat. Thailand: Mi thong pla (Karen).
Indochina: Cày cang pa; Co cang pa; Co mai ten; Xeur
nam ( Beille 1927 ). Cambodia: Thmenh trei (Khmer). Malay
Peninsula: Berbeti; Beti; Beti ayer; Mumbuti (Batek). Sumatra:
Mata-mata-ponei; Tjoemata or Tjoemateh (Malay) ( Miquel
1860 ). 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 spec-
imens were cited in his description of Cicca pentandra . Webster
(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) .
Representative Specimens Examined— (limited to a single specimen
per country)—BANGLADESH. Chittagong: Teknaf Game Reserve, 80 km
from Cox’s bazaar, 15 Apr 1994, Huq & Mia 10339 (L). BENIN. Atlantique:
Kpomassè, Aïdjèdo, 31 Aug 1999, Essou et al. 1795 (BENIN). Mono:
Aplahoué, SW of Adogonou, near Adjarala waterfalls of Mono River, 8
Nov 1997, Van der Maesen et al. 6240 (WAG). BURKINA FASO. Boulgou:
some km SE of Tenkodogo, 4 Sep 1996, Madsen 5526 (WAG). BURMA
(MYANMAR). Tenasserim: Tawer dist., E of Kanbauk village, Dawei River,
Yebyu township, J. F. Maxwell 98–1147 (L). CAMBODIA. Stung Treng:
Thala Barevath Dist., Preuh Rum Keil Village, Ramsar Site, Monyrak &
Meng 160 (L). CAMEROON. Central Province: left bank Sanaga River,
near Ferry Nachtigal, about 20 km N of Obala, 19 Nov 1965, Leeuwenberg
7037 (L). CHINA. Guangdong: Jen-hwa, Shek Pik Ha village, 21–30
Apr 1936, W. T. Tsang 26246 (L). COMOROS. Mohéli: N. side, 31 Dec
1938, Lam & Meeuse 6157 (L). ETHIOPIA. Kefa: c. 200 km SW of Addis
Abeba, near the bridge over the Omo-Bottega River in the road Addis
Abeba to Jimma, 23 Aug 1965, W. J. J. O. de Wilde & De Wilde-Duyfjes 7876
(K, WAG). GHANA. Volta Region: km 27 Hohoe - Peki Road, 2 km S of
Ve Agombe, 9 Mar 1977, Leeuwenberg 11178 (WAG). INDIA. Andaman and
Nicobar Islands: Car Nicobar, Kimus, 10 Jun 1974, N. G. Nair 1583 (L).
INDONESIA. Jawa Barat: Preanger, Cicoeroeg, Cisokan, Bakhuizen van den
Brink Sr 2813 (L). IVORY COAST. Abidjan: Adiopodoume, 17 km W of
Abidjan, 18 May 1962, Leeuwenberg 4211 (L). JAPAN. Kyushu: Oita, Kuju
Mountain, 30 Oct 1926, S. Saito 3011 (L). LAOS. Sayaboury: Phiang Dist.,
Nampouy village area, along Nam Pouy River, 29 Aug 1999, J.F. Maxwell
99–238 (L). MADAGASCAR. Toliara (Tulear): Tôlanaro, Reserve Naturelle
11, Andohahela, Parcelle 2, vicinity of Gite d’Etape, NE of Amboasary,
near Hazofotsy, 30 Nov 1991, Leeuwenberg 1134 (L). MALAYSIA. Kelantan:
Sungei Lebir at Kuala Relai at Jentah, 26 Apr 1976, B. C. Stone & Sidek
12453 (L). MALI. Kayes: Parc National de la Boucle du Baloué. Missira,
on area Benbugu, 13 Jun 1979, Dekker 404 (WAG). NAMIBIA. Karibib:
Otjosondu, 13 Feb 1963, Seydel 3327 (L). NEPAL. s. d., Wallich s. n. (L).
NIGERIA. Bauchi State: Duguri Dist., Yankari Game Reserve, Wikki-
Gaji confluence, 19 Aug 1971, Geerling 4038 (WAG). PHILIPPINES:
Luzon: Albay Prov., Taqui River, San Bernardo, Tiwi Side, SE slopes of
Mt. Malinao, Reynoso et al. PPI 21258 (L). SENEGAL. Sénégal-Oriental:
Gouloumbou, Sankange, 27 Aug 1991, Van der Steur 85 (WAG). SOUTH
AFRICA. Transvaal: Barberton, Highlandscreek Barberton, 15 Oct 1938,
Kleinhoonte 836 (L). TAIWAN. Pingtung Hsien: Kenting – Hengchun,
1 May 1928, S. Saito 7729 (L). THAILAND. Central: Saraburi Prov., between
Saraburi and Pok Chong, 18 Jul 1973, Murata, Fukuoka & Phengkhlai T-16519
(L). TOGO. Maritime: Forêt classée de la Lili, Lillikopé, 22 Jul 1971, Breteler
7015 (WAG). UGANDA. Kashari: E Ankole U2 dist., Bisya near Rubindi,
14 Jun 1997, Rwaburindore 4136 (L). VIETNAM. Ninh Binh: Cuc Phuong
National Park, CP54 site, Cuong 207 (L). ZIMBABWE. Mashonaland West:
Kariba District, Kessesse River, Nov 1959, Goldsmith 59/66 (L).
Excluded Names— Only names of Malesian taxa not or
partly treated by Webster (1984) are mentioned here:
Flueggea javanica Blume, Bijdr.: 580. 1825.—TYPE: INDONESIA.
Java, Gunung Parang, s. d., Blume s. n. , barcode L 0813547
(holotype: L) = 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: 334. 1866.—TYPE: AUSTRALIA,
Queensland, Gilbert River, s. d., F. Mueller s. n. (lectotype,
selected here: K ‘Securinega leucopyra ’). = Flueggea leu-
copyrus Willd. Webster (1984) reduced this variety to
Flueggea virosa (Roxb. ex Willd.) Royle subsp. melanthe-
soides (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.
Acknowledgments. We are grateful to the directors and curators of
the following herbaria for the loan of material: BO, GH, HBG, L, LAE,
PNH, SAN, SING and WAG. We would also like to thank P. Hoffmann (K)
and C.M. Wilmot-Dear (K) for their assistance and encouragement and the
reviewers and Daniel Potter for their constructive remarks. The beautiful
drawings were made by Jan van Os (L) and Hanneke Jelles (L).
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