Malesian Euphorbiaceae Descriptions

21. BRIDELIA

 

S. Dressler

 

Dressler, S. 1996. The genus Bridelia (Euphorbiaceae) in Malesia and Indochina - A regional revision. Blumea 41: 263—331.

 

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Genus description

Key to the species

Subgenus Bridelia

Subgenus Gentilia

 

Bridelia Willd.

 

    Bridelia (`Briedelia') Willd., Sp. Pl. 4 (1806) 978; corr. Spreng., Anleit. Kenntn. Gew. ed. 2, 2 (1818) 887; Mόll.Arg. in DC., Prodr. 15, 2 (1866) 492; Hook.f., Fl. Brit. India 5 (1887) 267; Gehrm., Bot. Jahrb. Syst. 41, Beibl. 95 (1908) 1; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 54; Ridl., Fl. Malay Penins. 3 (1924) 183; Gage, J. Asiat. Soc. Bengal. 75 (1936) 484; Backer & Bakh. f., Fl. Java 1 (1964) 475; Airy Shaw, Kew Bull. 26 (1972) 227; Whitmore, Tree Fl. Malaya 2 (1973) 74; Airy Shaw, Kew Bull. Add. ser. 4 (1975) 63; Kew Bull. Add. ser. 8 (1980) 43; Kew Bull. 36 (1981) 272; Kew Bull. 37 (1982) 10; Enum. Euphorb. Philipp. Isl. (1983) 11; G.L.Webster, Regnum Veg. 129 (1993) 153; Ann. Missouri Bot. Gard. 81 (1994) 39; S.Dressler, Taxon 45 (1996) 337, nom. cons. prop.; Blumea 41 (1996) 273; Radcl.-Sm., Gen. Euphorbiacearum (2001) 19; S.Dressler & Welzen in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 141; G.L.Webster in Kubitzki, Fam. Gen. Vasc. Pl. 11 (2014) 71. — Lectotype (designated by G.L.Webster, 1993): Bridelia scandens (Roxb.) Willd. [= Bridelia stipularis (L.) Blume].

 

Small trees to shrubs, sometimes scramblers, occasionally thorny from horizontal aerial roots or persisting twig bases, usually monoecious; branchlets terete. Bark smooth or tearing off in scales or long stripes. Indumentum consisting of simple, appressed to patulous, short or longer hairs, at least on the young parts, in certain taxa lacking. Leaves alternate, simple, smaller towards the twig tips; stipules caducous; petioles mostly thicker and darker than midvein; blade symmetric, base obtuse to acute, apex obtuse, rounded, acute or acuminate, margin entire to slightly crenate; venation pinnate, craspedodromous but secondaries terminating in a fimbrial vein (subg. Bridelia: sect. Bridelia, Scleroneurae sensu Jabl., 1915) or brochidodromous (secondaries joined together in arches before the margin (subg. Gentilia: sect. Cleistanthoideae), secondary veins mostly uniformly, narrowly to moderately acutely angled, tertiary veins straight percurrent (scalariform) to weakly percurrent and recurved towards midvein, areolation reticulate. Inflorescences scariously bracteate, axillary fascicles, glomerulate, sometimes spicate or terminal on leafless branchlets, diclinous flowers in the same inflorescence with usually the central pistillate ones surrounded by staminate ones, dichogamous. Flowers usually small to minute, hypogynous, rarely perigynous, actinomorphic, whitish to greenish, sessile to shortly pedicelled, pedicel slightly elongating in fruit; sepals 5, valvate, thick, fleshy, persistent in fruit; petals 5, minute, alternate with sepals, very variable in shape, obovate or obcuneate, subentire to lobulate, thin; disc plain, broadly pulvinate or saucer-shaped, fleshy .Staminate flower: disc circular ; stamens 5, episepalous, filaments united below into a short central column, apically free, filiform and spreading, with pistillode terminating the column, anthers ovoid to ellipsoid, basifixed or dorsifixed, introrse, dithecal, tetrasporangiate, longitudinally dehiscent. Pistillate flowers: disc with additional tubular inner disc present, conically truncate, membranaceous, enclosing the ovary and splitting into disc-scales adpressed to the outer portion of the disc during fruit development; ovary 2- or 3-locular, locules each with 2 ovules, anatropous, descending, epitropous, only 1 developing; in subg. Gentilia only one locule developing; styles usually 2, free or united over a variable length, bifid above; stigmas terete to flat. Fruits small drupes, ellipsoid to roundish, sometimes pointed or bilobate at apex, with fleshy mesocarp and hard endocarp which sometimes splits (e.g., B. stipularis, B. ovata), bilocular or 1-locular by abortion; if bilocular endocarps subgloboid with flat ventral and convex dorsal surface, if unilocular endocarp ellipsoid. Seeds 1 (subg. Gentilia) or 2 (rarely 3) per fruit, ovoid and pointed or semi-ovoid with one flat, often sulcate side (if 2-seeded); testa smooth, glabrous, brownish. Embryo straight or slightly curved, cotyledons foliaceous, endosperm fleshy or membranaceous.

    Distribution — About 50 species in the Old World tropics, distributed in Tropical Africa, Madagascar, Yemen and in Asia ranging from India and S China throughout Indochina, Malesia to N Australia and the Solomons and Vanuatu.

    Habitat & Ecology — The species in Southeast Asia are usually part of the primary and secondary forest vegetation either as big trees or as smaller trees or shrubs in the understorey. Some species are scrambling and one is reported to be a climber. Some species seem to be restricted to a certain type of habitat, e.g., B. cinnamomea peat swamp forests, B. parvifolia sand dunes, B. oligantha dry savannahs. The species occur from sea level up to 1800 m, but in sect. Scleroneurae there are several species restricted to lower altitude.

    Note — Closely related to Cleistanthus, with which it forms tribe Bridelieae of subfam. Phyllanthoideae, but easily distinguishable by its drupaceous fruit and mostly 2-locular ovary as well as by the often more chartaceous leaves with a different venation: Cleistanthus always has brochidodromous secondary veins, whereas in Bridelia only sect. Cleistanthoideae of subg. Gentilia shows this feature. The greater part of the genus shows craspedodromous venation with secondary veins joining a fimbrial vein.

 

Key to the species

 

1a.

Leaves with a distinct fimbrial vein with secondary veins joining it (craspedodromous). Drupes bi- or trilocular. Seeds plano-convex (subg. Bridelia).

2

1b.

Leaves without a distinct fimbrial vein; secondary veins arching near the margin (brochidodromous). Drupes unilocular. Seeds ± ellipsoid with a deep longitudinal groove (subg. Gentilia). 

11

2a.

Ovary and drupes regularly trilocular. – Secondary veins (10–14 pairs) con­spicuously steeply ascending. New Guinea. 

9. B. triplocarya

2b.

Ovary and drupes bilocular; rarely some trilocular may be present.

3

3a.

Flowers large, 6–12 mm in diameter (calyx).

4

3b.

Flowers small, up to 6 mm in diameter. (calyx). 

5

4a.

Secondary veins in 9–14 pairs. Leaves tomentose to velvety beneath. Fruits 6–11 mm in diameter, bilobate, often dehiscing at apex. Floral disc 4–6 mm in diameter. Sri Lanka to Lesser Sunda Islands.

7. B. stipularis

4b.

Secondary veins in (12–)17–23 pairs. Leaves glabrous to pilose beneath. Fruits 17–30 mm in diameter, plum-like, rarely trilocular. Floral disc up to 3 mm in diameter. Halmahera, New Guinea.

3. B. macrocarpa

5a.

Secondary veins usually in more than 12 pairs.

6

5b.

Secondary veins usually in less than 12 pairs.

9

6a.

Fruits very big, 1.5–3 cm in diameter, plum-like. 

3. B. macrocarpa

6b.

Fruits smaller, up to 1.5 cm in diameter     

7

7a.

Inflorescences terminal on mostly leafless twigs, spike-like. Secondary veins numerous (> 18 pairs). Leaves coriaceous. 

6. B. retusa

7b.

Inflorescences in the axils of normal leaves. Secondary veins in less than 18 pairs. Leaves chartaceous to subcoriaceous.

8

8a.

Leaves glabrous beneath. Small tree to 8 m high. SE Asia, Java, Lesser Sunda Islands — Leaves drying to olive green to blackish.

5. B. ovata

8b.

Leaves puberulous to pubescent beneath. Large tree, up to 15 m high. New Guinea.

2. B. erapensis

9a.

Leaves thinly chartaceous, glaucous beneath. Apex (bluntly) acute to shortly acuminate. Plant mostly rather delicate, twigs whip-like, slender.

8. B. tomentosa

9b.

Leaves stiffly chartaceous to coriaceous, lighter beneath. Apex obtuse to rounded. ― Twigs more robust, often strongly branched. 

10

10a.

Leaves conspicuously narrow-oblong (leaf index > 2), stiffly (sub)coriaceous. SE Asia, W Malesia.

1. B. curtisii

10b.

Leaves broadly elliptic (leaf index 1–2.1), stiffly chartaceous. New Guinea.

4. B. oligantha

11a.

Big woody climber. Ovary hairy. Leaves drying greenish-brown, somewhat bullate.

15. B. whitmorei

11b

Bushes, scramblers or small trees. Ovary glabrous (tubular disc sometimes with hairs). Leaves ± plain. 

12

12a.

Leaves puberulous to tomentose beneath. 

13

12b.

Leaves glabrous beneath.

16

13a.

Inflorescence cushions strongly protruding, rather large (> 7 mm in diameter). Plant nigrescently blackening when drying. Tertiary veins conspicuously ladder-like. Branchlets robust, rough, strongly pustulate-lenticellate. Pistillate tubular disc sometimes hairy outside.

14. B. pustulata

13b.

Inflorescence cushions smaller (< 7 mm in diameter). Plant not nigrescent. Tertiary venation not conspicuously ladder-like. Branchlets less robust, neither rough nor pustulate. Pistillate tubular disc always glabrous outside.

14

14a.

Secondary veins in 6–9 pairs with a narrow acute angle of divergence (< 45°). Leaf base mostly acute, apex very shortly acuminate (3–7 mm). Stipules very short (1.5–2 mm). Styles not exserted.

11. B. cinnamomea

14b.

Secondary veins in > 9 pairs with a moderately acute angle of divergence (usually > 45°). Leaf base obtuse to truncate, rarely acute, apex longer acuminate. Stipules longer than 2 mm. Styles exserted.

15

15a.

Flowers conspicuously pedicelled (2–6 mm), 3–5 mm in diameter. Inflorescence cushions of rather ‘smooth’ appearance. Secondary veins in 11–18 pairs. 

12. B. glauca

15b.

Flowers (sub)sessile, 2–3 mm in diameter. Inflorescence cushions of rather ‘rugged’ appearance. Secondary veins in 9–13 pairs.

13. B. insulana var. subnuda

16a.

Inflorescence few-flowered (< 14). Leaves stiffly chartaceous to coriaceous, glossy above, margin revolute.

10. B. adusta

16b.

Inflorescence many-flowered (> 14). Leaves chartaceous, rarely subcoriaceous, usually dull above, margin not revolute. 

17

17a.

Flowers conspicuously pedicelled (2–6 mm), 3–5 mm in diameter. Inflorescence cushions of rather ‘smooth’ appearance. Secondary veins in 11–18 pairs.

12. B. glauca var. acuminatissima

17b.

Flowers (sub)sessile, 2–3 mm in diameter. Inflorescence cushions of rather ‘rugged’ appearance. Secondary veins in 9–13 pairs.    

13. B. insulana

 

Subgenus Bridelia

 

    Bridelia subgenus Bridelia: S.Dressler, Blumea 41 (1996) 276. — Bridelia sect. Eubridelia Gehrm., Bot. Jahrb. Syst. 41, Beibl. 95 (1908) 11, nom. inval. — Bridelia subg. Eubridelia (Gehrm.) Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 55, nom. inval.

 

1. Bridelia curtisii Hook.f.

 

    Bridelia curtisii Hook.f., Fl. Brit. India 5 (1887) 273; Boerl., Handl. Fl. Ned. Indiλ 3 (1900) 271; Ridl., J. Straits Branch Roy. Asiat. Soc. 59 (1911) 167; Fl. Malay Penins. 3 (1924) 184; Gage, J. Asiat. Soc. Bengal. 75 (1936) 488 (in obs.); S.Dressler, Blumea 41 (1996) 278, map 1; S.Dressler & Welzen in Welzen et al., Thai For. Bull. 28 (2000) 61; in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 143, plate V: 3. — Bridelia ovata var. curtisii (Hook.f.) Airy Shaw, Kew Bull. 26 (1972) 229. — Type: Curtis 97 (K holo; SING iso), Penang, at Tulloh Bahang.

    Bridelia griffithii auct. non Hook.f.: Gehrm., Bot. Jahrb. Syst. 41, Beibl. 95 (1908) 38.

 

Scrambling shrub to small tree, up to 7 m high; branches spreading, glabrous to puberulous, with scattered lenticels. Stipules early caducous, narrowly triangular, glabrous, up to 4by 0.6–0.7 mm. Leaves: petiole terete, glabrous, rarely puberulous, 3.5–6 by 0.8–1.2 mm; blade (elliptic or lanceolate to) conspicuously oblong with parallel or only slightly curved sides, 30–80 by 10–36 mm, length/width ratio (1.5–)2–2.8, stiffly coriaceous; base (cordate to) rounded or obtuse (to acute); apex rounded (to bluntly acute); margin entire, surfaces subglabrous to slightly puberulous, then also adaxially scattered hairs on the main nerves, green abov, greyish green beneath; venation conspicuously prominent on both sides, secondary veins in 5–10(–12) pairs, arching and joining the marginal vein, tertiary veins irregularly percurrent, but mostly not scalariform, irregular reticulate areolation. Inflorescence: glomerules with up to 12 sessile, green flowers; bracts many, broadly ovate-triangular, up to 1 by 0.8 mm, scattered puberulous and with ciliate margin. Flowers rarely with up to 1 mm long puberulous stalk; sepals triangular-ovate, 1.5–2 by 1–1.5 mm, glabrous, greenish yellow; petals very variable in shape, base cuneate to spathulate, apex roundish, notched or lobulate, up to 1 by 0.7 mm, whitish yellow; disc glabrous. Staminate flowers 3–5 mm in diameter; disc 1.4–1.7 mm in diameter; stamens: free part of filaments up to 0.7 mm long, anthers shortly ellipsoid, 0.6–0.7 by 0.3–0.4 mm; staminal column c. 1 by 0.3 mm, pistillode conical-ovoid with 2-lobed top, up to 0.7 by 0.2 mm. Pistillate flowers 4–5 mm in diameter; outer disc c. 2.5 mm in diameter, inner one tubular, up to 1 mm long, margin sometimes ciliate; ovary globoid, c. 1.5 mm in diameter, styles up to 1.2 mm long, basal half united, apically free, deeply bifid. Infructescences with up to 6 fruits, mostly 2 or 3. Fruits depressed-ovoid, apically emarginate, slightly bilobate, 4.5–6.5 mm in diameter, when fresh fleshy, greenish purple, blackish when dry; endocarps 2, with woody, brown, semigloboid, apically splitting endocarp, c. 4 by 5 by 2.5 mm, Seeds semi-globoid to semi-tear-shaped, with lateral furrow, c. 3 by 4 by 2 mm, brown.

    Distribution — Indochina (Cambodia, Cochinchina), Thailand, Andaman & Nicobar Islands, Malay Peninsula, N Sumatra.

 

Bridcurt-map.gif (94799 bytes)

 

    Habitat & Ecology — Reported from mangroves, tidal riversides, evergreen for­ests and open, disturbed areas (savannah); often on limestone. Alt.: up to 25 m. Ripe berries eaten by Myophoneus (van Balgooy 2379, L).

    Uses — Fruits edible and medicinally used in Cambodia (Martin 155, L).

    Notes — 1. The characteristics of this species are the typical oblong leaf mostly with both ends rounded, the stiffly coriaceous texture, and the prominent venation on both sides of the dry leaf. The plant is a scrambling shrub or small tree obviously confined to lower altitudes.

2. Bridelia curtisii is morphologically to be placed between B. ovata and B. tomentosa. Nevertheless it is clearly distinguished from both so that I consider it to be a distinct species in this closely related group. The main differences to B. ovata are the lower number of secondary veins and the stiffly coriaceus texture of the characteristically narrowly oblong leaves with both ends rounded (leaf index > 2). From B. tomentosa on the other hand it differs by having stiffer leaves, but also an often stronger growth habit with bigger leaves, flowers, and fruits. This species has an acute leaf apex and base.

3. Airy Shaw (1972a) mentioned B. pedicellata Ridl. as a synonym of B. ovata var. curtisii. Ridley himself (1924: 184) already synonymised this taxon with B. ovata but he did this with B. curtisii as well. Three of the four syntypes of B. pedicellata are housed in Kew and two of them have a leaf shape which superficially resembles B. curtisii. To clarify this matter I (Dressler) lectotypify this species with Ridley 8326 (K), a specimen which clearly represents B. ovata.

4. Strangely Gehrmann (1908) placed B. curtisii among the synonyms of B. griffithii but said nothing about his reasons. Jablonszky (1915: 74) only cited Gehrmann with the remark not to have seen the type. There is no justification for a conspecificy of these both species since they belong to the different subgenera.

5. Two collections from N Thailand (Santisuk 1507 and van Beusekom & Phengklai 2530, both L) show a remarkable difference: their leaves are lanceolate with a distinctly acute apex. As there are no other macromorphological features which distinguish them from B. curtisii, to which they are most similar in leaf texture and venation, I did not venture to describe a new species although these collections seem clearly distinguished by their altitudinal occurrence too (1000–1400 m). Further material is needed to clear their taxonomic situation.

 

2. Bridelia erapensis S.Dressler

 

    Bridelia erapensis S.Dressler, Kew Bull. 51 (1996) 601, Fig. 1, Map 1; Blumea 41 (1996) 280, map 2. — Type: Hartley 12224 (L holo; CANB, K, LAE iso), Papua New Guinea, Morobe Prov., Markham River Valley at Erap, W of Lae along the Lae-Goroka road.

    Bridelia ovata auct. non Decne.: Hartley et al., Lloydia 36 (1973) 264.

    Bridelia macrocarpa auct. non Airy Shaw: Airy Shaw, Kew Bull. Add. Ser. 8 (1980) 43, p.p.

 

Briderap-total.gif (83462 bytes)

 

Tree, up to 15 m high, stem 25 cm diameter; flowering branches slender, glabrous to pubescent, scattered lenticellate. Outer bark greyish brown, scaly and fissured, inner bark salmon or pink, blaze straw. Stipules not seen, early caducous. Leaves: petiole subterete, pilose to pubescent, 4–6 by 1.2–2 mm; blade ovate (to elliptic), 3–10 by 2–6 cm, length/width ratio 1.5–1.9, rigidly chartaceous to subcoriaceous; base obtuse, rounded to slightly cordate, margin entire to distantly shallowly crenate where veins reach the margin, apex obtuse to acute, upper surface glabrous, dark green, lower surface puberulous, greyish green, often glaucous; venation prominent on both sides, stronger below, secondary veins in 14–18 pairs, fairly straight, only slightly arching, but often forking before joining the fimbrial vein, secondaries often forming loops near the margin and tertiary veins joining the fimbrial vein, tertiary veins ± patent to secondaries, percurrent, scalariform, reticulate areolation. Inflorescences paucibracteate glomerules of 2–5 sessile, greenish yellow flowers; bracts (broadly) triangular, up to 1.3 by 1.2 mm, pilose to glabrous. Staminate flowers not seen. Pistillate flowers 3.5–5 mm in diameter, up to 0.5 mm long stalked; sepals triangular, 1.6–2.2 by up to 1.7 mm, glabrous; petals variable in shape, base spatulate to cuneate, apex triangular to lobulate with gnawed margin, 1–1.3 by 0.6–1 mm; outer disc c. 2 mm in diameter, inner one up to 1.1 mm long, margin irregular erose to dentate; ovary ellipsoid, conical at apex, 2-locular, c. 2 by 1.5 mm diameter, styles 0.8–1.3 mm long, deeply bifid, hence resembling 4 separate styles, stigmas lobate. Infructescences with up to 3 almost sessile fruits. Fruits globose, 5–6 by 5–6 mm, smooth, green turning to purple when ripe; endocarps 2, semigloboid, with pointed base, c. 4.5 by 4.5 by 2.5–3 mm. Seeds semigloboid with shallow ventral furrow, c. 3 by 3.5 by 1.5 mm; testa reddish brown.

    Distribution — NE New Guinea.

 

Briderap-macr-olig-map.gif (75597 bytes)    B. erapensis (triangle; B. macrocarpa = dot; B. oligantha = square)

 

    Habitat & Ecology — Lowland monsoon or rain forests; reported from c. 45 m altitude.

    Notes — 1. This new species was based on two collections from the Markham River Valley near Erap in the Morobe Province of New Guinea. The epithet refers to the locality, known to have a rather dry climate (Van Welzen, 1995, pers. comm.). A third collection (Clemens 10501, E) was later traced, also from the same area.

2. One of the specimens was mentioned by Airy Shaw (1980) under B. macrocarpa but with reference to its unusual cordate leaf base. As I found another vegetatively similar collection with much smaller bilocular fruits and as the combination of characters does not fit in any other species, the specimens have to be considered as an undescribed taxon. This species is characterised by ovate, basally obtuse to slightly cordate leaves of a (sub)coriaceous texture with a puberulous undersurface. Its venation pattern resembles more that of B. triplocarya. It has less secondary veins which often branch and curve before the margin, hence the secondaries form indistinct loops and join the fimbrial vein only sometimes. Tertiary veins arising from the branching of the secondaries join the fimbrial vein as well. It has only 2–5 flowers per glomerule and the fruits are bilocular and of only 5–6 mm in diameter. So it is clearly distinguished from B. macrocarpa and also from B. triplocarya.

 

3. Bridelia macrocarpa Airy Shaw

 

    Bridelia macrocarpa Airy Shaw, Kew Bull. 25 (1971) 512; Kew Bull., Add. Ser. 8 (1980) 43, t. 1, f. 2; S.Dressler, Kew Bull. 51 (1996) 606, Map 2; Blumea 41 (1996) 283, map 2. — Type: Hoogland 4913 (K holo; CANB, L, LAE iso), New Guinea, Madang Prov., near Mawan village, Gogol Valley.

    Bridelia beguinii Airy Shaw, Kew Bull. 37 (1982) 10 — Type: Beguin 2039 (L holo; BO, L iso), Moluccas, Halmahera, Soa Tobaroe.

 

Tall slender tree (up to 30 m high) with straight clear bole, buttresses, thick, up to 1.5 m; crown long, forked; branches slender, glabrous to scattered pilose, young glaucous. Outer bark dark greyish brown, flaking in square scales, with an acrid smell; underbark pale purple to red; inner bark pink or red, in papery layers, no exudate; sapwood white, cream or pinkish; heartwood brown, very hard. Stipules narrowly triangular, 5–7 by 2.5–4 mm, whitish pubescent, early caducous. Leaves: petiole subterete, sparsely pilose, 7–13 by 1–2.5 mm; blade broadly elliptic (to obovate), 64–220 by 30–116 mm, length/width ratio 1.3–2.6, chartaceous; base (cordate to) rounded to obtuse, margin entire to shallowly crenate where veins reach margin, apex (emarginate to) obtuse to acute, upper surface glabrous except pilose on veins, light to dark green, lower surface inconspicuously whitish or brownish pilose, greyish green; venation prominent on both sides, stronger below, secondary veins in (12–)17–23(–28) pairs, fairly straight, only slightly arching, but often forking before joining the marginal vein, tertiary veins ± patent to secondaries, simple or rarely forked percurrent, scalariform. Inflorescences glomerules with (8–)9–16 sessile, yellowish green to creamy flowers, mostly not terminal and not at special leafless branches, with pistillate flowers surrounded by staminate ones; bracts triangular to semi-ovate, up to 1.5 by 2 mm, whitish pilose. Flowers: pedicel indistinct, up to 0.5 mm long; sepals triangular; petals variable in shape, base spatulate, apex roundish to lobulate with gnawed margin, (1–)1.5–2 by 1–1.5 mm. Staminate flowers 3.5–5 mm in diameter; sepals 1.3–1.5 by c. 0.8 mm; disc 1.8–2.5 mm in diameter; stamens: free part of filaments c. 0.8 by 0.1 mm diam, anthers ovoid, 0.5–0.8 by 0.3–0.4 mm, column 0.7–1.5 by 0.2–0.5 mm, pistillode ovoid, bifid at apex, c. 0.5 by 0.2 mm. Pistillate flowers 6.5–10 mm in diameter; sepals 3–4 by 3–3.5 mm; outer disc 2.5–3 mm in diameter, inner one 1–2 mm long when tubular, scales 1–1.3 mm long; ovary globoid, conical at apex, pointed, 2(3)-locular, c. 2.5 by 2.5–3 mm, styles 0.8–1 mm long, deeply bifid as to 4 separate styles. Infructescences with l almost sessile fruit. Fruits globose, c. 25 by 17–30 mm in diameter, smooth, firm, succulent, red to purple in natural state; concerning the fruit in dried state cf. Airy Shaw (1971: protologue); mature endocarps and endocarp not seen, globular with 3 prominent vanes splitting along the vanes into 3 segments (see NGF (Jackson) 4567). Seeds suborbicular, narrowly concavo-convex, c. 1 cm in diameter; testa reddish brown, cotyledons flat (Airy Shaw, 1971).

    Distribution — Moluccas (Halmahera), New Guinea.

 

Briderap-macr-olig-map.gif (75597 bytes)    B. macrocarpa (dot; B. erapensis = triangle; B. oligantha = square).

 

    Habitat & Ecology — Lowland rain forests, also tall secondary forest; on sandy or clayey soil, reported from flat, poorly drained or temporarily flooded terrain. Alt.: sea level up to 150 m.

    Uses — Reputation for durability of up to 10 years as posts, also used for carved bowls, lime sticks etc. (NGF (Jackson) 4567, K, L).

    Note — This species has some affinities with B. stipularis and B. retusa, both from S Asia and W Malesia, but nevertheless it is easily distinguishable by the large characteristic fruits which may have three seeds and by the lack of the special leafless branches which form the inflorescence. From B. stipularis it is easily distinguishable by having much more lateral veins and 8–16 flowers per glomerule. Bridelia retusa has similar but thicker and more coriaceous leaves.

 

4. Bridelia oligantha Airy Shaw

 

    Bridelia oligantha Airy Shaw, Kew Bull. 32 (1978) 386; Kew Bull. Add. Ser. 8 (1980) 44; Kew Bull. 37 (1982) 11; S.Dressler, Kew Bull. 51 (1996) 607, Map 1 (‘oligandra’); Blumea 41 (1996) 284, Map 2. — Type: Pullen 6822 (K holo; A, BO, BRI, CANB, L, LAE, US iso), Papua New Guinea, Central Prov., Port Moresby Subprov., Tavai Creek area, 72 km SE of Port Moresby.

 

Shrub or small tree, up to 8 m high, dbh 10–20 cm; branches slender, glabrous throughout, rarely pilose above inflorescence axils, scattered lenticellate. Bark grey, longitudinally grooved, wood cream. Stipules rather long persistent, narrowly triangular to linear, sometimes falcate, 1(–2) by 0.5–0.7, medially c. 0.2 mm wide, margin ciliate. Leaves comparatively small; petiole terete, 4–7 by 0.8–1 mm, glabrous; blade (slightly ovate to) broadly elliptic, 2–6 by 1.3–4.3 cm, length/width ratio 1.1–2(–2.1), stiffly chartaceous to subcoriaceous, base obtuse to rounded (to acute), margin entire, rarely faintly crenate where veins reach margin, apex (emarginate to) obtuse to broadly rounded, upper surface glabrous, olive green, lower surface (sub)glabrous, pale glaucous green; venation faintly prominent to obscure on both sides, abaxially often rather inconspicuous, secondary veins in 8–11 pairs, rather irregular, straight to arching and joining the marginal vein, sometimes branched near margin, tertiary veins irregularly reticulate or rarely percurrent, not scalariform, reticulate areolation. Inflorescences glomerules with 1–8 sessile greenish yellow flowers with several glabrous bracts; latter broadly triangular, 0.8–1.2 by c. 0.8 mm, with ciliate margin and thick, dark keel (midrib). Flowers 3–4 mm in diameter; sepals triangular, 1–1.5 by 0.9–1.2 mm, glabrous; petals very variable in shape, base cuneate, spatulate to nailed, apex roundish, rhomboidal or triangular, 0.6–1 by 0.4–1 mm, light with darker central part; disc glabrous;  Staminate flowers greenish cream; disc c. 1.8 mm in diameter; stamens: free part of filaments very short, < 0.5 mm long, anthers shortly ellipsoid, 0.4–0.6 by c. 0.3 mm, column c. 1 by 0.3 mm, rather stout; pistillode shortly ovoid to bluntly conical, c. 0.5 by 0.2 mm. Pistillate flowers: outer disc c. 2 mm in diameter, inner one tubular with erose margin, up to 0.7 mm long; ovary globoid, c. 0.8 mm in diameter; styles 0.5–0.8 mm long, deeply bifid, only united at the base, apically bilobed, not exserted in flower. Infructescences with up to 3 fruits, mostly only 1. Fruits slightly compressed globoid, sometimes subacute at base and apex, bilocular, 4.5–5 by 3.5–4 mm in diameter, yellowish green when fresh, dry blackish; endocarps 2, endocarp brown, semigloboid, subacute at base and apex, apically often splitting, c. 3 by 4 by 2 mm. Seeds semigloboid, shallowly ventrally sulcate, c. 3 by 3 by 1.5 mm, brown, shining, embryo flat.

    Distribution — New Guinea (Central Province: around Port Moresby).

 

Briderap-macr-olig-map.gif (75597 bytes)    B. oligantha (square; B. erapensis = triangle; B. macrocarpa = dot).

 

    Habitat & Ecology — Low monsoon thickets or forests, mixed savannah and grassland. Alt.: up to 50 m.

    Notes — 1. Characteristic for B. oligantha are the rather small and broadly elliptic leaves with a smooth, perfectly glabrous undersurface as well as the few flowers per inflorescence only. The veins are normally not prominent. The very broadly round shape of the leaves and their thicker texture and smoother lower surface distinguish it from broad-leaved forms of B. tomentosa var. glabrescens. Without any doubt the latter species is very closely related to B. oligantha. Morphologically similar is B. parvifolia from Indochina, a rare small shrub from dune thickets. Here the intemodes are often very short and the leaves are mostly conspicuously nigrescent as opposed to B. oligantha where they dry to a greenish brown colour.

2. I agree with Airy Shaw (1978) who stated that this species represents a distinct local endemic but I cannot confirm his reference to Timor (Airy Shaw, 1982: 11: B. cf. oligantha) which could possibly be based on a broad-leaved specimen of B. tomentosa. So far we know this species only from a restricted area near Port Moresby with a dry climate and savannah-like vegetation near the coast.

 

5. Bridelia ovata Decne.

 

    Bridelia ovata Decne., Nouv. Ann. Mus. Hist. Nat. 3 (1834) 484; Span., Linnaea 15 (1841) 347; Baill., Ιtude Euphorb. (1858) 583; Miq., Fl. Ned. Indiλ 1, 2 (1859) 364; Mόll.Arg. in DC., Prodr. 15, 2 (1866) 495; Kurz, Prelim. Rep. For. Pegu, App. A: cix., App. B. (1875) 78, in clavi; Kurz, For. Fl. Burma 2 (1877) 368; Fern.-Vill. in Blanco, Fl. Filipp., ed. 3, Nov. App. 4 (1880) 186 (see below); Hook.f., Fl. Brit. India 5 (1887) 274; Boerl., Handl. Fl. Ned. Indiλ 3 (1900) 271; Gehrm., But. Jahrb. Syst. 41, Beibl. 95 (1908) 33; J.J.Sm., Meded. Depart. Landb. Ned.-Indiλ 10 (1910) 324; Koord., Exkurs.-Fl. Java 2 (1912) 485, in clavi; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 61, 63; Ridl., Fl. Malay Penins. 3 (1924) 184; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1926) 489; Gage, J. Asiat. Soc. Bengal. 75 (1936) 488; Henderson, J. Malayan Branch Roy. Asiat. Soc. 17 (1939) 69; Corner, Wayside Trees Malaya, ed. 2, 1 (1951) 243, ed. 3, 1 (1988) 279; Backer & Bakh.f., Fl. Java 1 (1964) 475, in clavi; Airy Shaw, Kew Bull. 26 (1972) 229; Whitmore, Tree Fl. Malaya 2 (1973) 74, in clavi; Airy Shaw, Kew Bull. 37 (1982) 11; Pham-hoang Ho, Cayco Vietnam 2 (1992) 291; S.Dressler, Blumea 41 (1996) 285, Map 3; S.Dressler & Welzen in Welzen et al., Thai For. Bull. 28 (2000) 64; in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 147. ― Amanoa ovata (Decne.) Baill., Adansonia 6 (1866) 336, p. p. — Bridelia ovata var. genuina Mόll.Arg. in DC., Prodr. 15, 2 (1866) 495, nom. inval. — Type: Riedlι s.n. (P holo; A, E, K, L, NY iso), Timor.

    Bridelia lanceolata Kurz ex Teysm. & Binn., Natuurk. Tijdschr. Ned.-Indiλ 27 (1864) 45. — Cleistanthus lanceolatus (Kurz ex Teijsm. & Binn.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 507; Boerl., Handl. Fl. Ned. lndiλ 3 (1900) 271; Hallier f., Meded. Rijks-Herb. 1 (1910) 6. — Type: Teijsmann s.n. [BO; probable isotype: G-DC (IDC microfiche Candolle Prodromi Herbarium 2511.19 & 20), K], Java, prope Japara.

    Bridelia glauca auct. non Blume: Koord. in Junghuhn-Gedenkboek (1910) 173 (sec. Hallier f., 1910).

    Bridelia ovata var. acutifolia Mόll.Arg. in DC., Prodr. 15, 2 (1866) 495. — Type: Helfer KD 4884 (G-DC, IDC microfiche Candolle Prodromi Herbarium 2508.24), in Tenasserim and Andamans.

    Bridelia burmanica Hook.f., Fl. Brit. India 5 (1887) 269; Gehrm., Bot. Jahrb. Syst. 41, Beibl. 95 (1908) 28; Jabl. in Engl., Pflanzenr. Heft 65 (1915) 69; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1926) 489. — Syntypes: Wallich 7888 (BM, K, K-Wall.), Burma (Hort. Bot. Cale.); Kurz s.n. (BO), Burma, Ava.

    Bridelia amoena auct. non Wall. ex Baill.: Kurz, For. Fl. Burma 2 (1877) 368.

    Bridelia kurzii Hook.f., Fl. Brit. India 5 (1887) 272; Williams, Bull. Herb. Boissier, sιr. 2, 5 (1904) 31; Brandis, Indian Trees (1906) 560; Gehrm., Bot. Jahrb. Syst. 41, Beibl. 95 (1908) 32; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 65. — Type: Kurz s.n. (K holo), Nicobar Islands, Kamorta.

    Bridelia pedicellata Ridl., J. Straits Branch Roy. Asiat. Soc. 59 (1911) 167; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 63; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1926) 490; Pham-boang Ho, Cayco Vietnam 2 (1992) 291. — Lectotype (designated by S.Dressler, 1996): Ridley 8326 (K holo; SING iso), Malaya, Kedah, Langkawi.

    Bridelia kurzii auct. non Hook.f.: Williams, Bull. Herb. Boissier, sιr. 2, 5 (1905) 31.

 

   

 

Scrambling shrub or small tree, up to 8 m high, dbh 10 cm; crown flattened; branche glabrous, with scattered lenticels. Bark fissured, dark greyish brown. Stipules early caducous, narrowly triangular, subulate, up to 7(–10) by 1.2 mm, glabrous. Leaves: petiole terete, (3–)4–6 by 0.8–1.5 mm, glabrous; blade elliptic (to slightly obovate), 5–18 by 2–8(–10) cm, length/width ratio 1.4–2(–2.2), chartaceous, often conspicuously blackish olive when dry; base (slightly cordate to) obtuse, margin entire, apex obtuse to rounded (to bluntly acute), upper surface glabrous, light green; lower surface glabrous, greyish green to glaucous; venation prominent on both sides, secondary veins in 13–17 pairs, arching and joining the marginal vein, sometimes branching, tertiary veins irregularly percurrent, sometimes scalariform, well-developed reticulate areolation. Inflorescences multibracteate glomerules of 1–20 (or more) subsessile to shortly pedicelled yellowish green flowers; bracts ovate-triangular, up to 2 by 1–1.5 mm, with scattered hairs and membranaceous margin. Flowers subsessile to shortly stalked (the staminate flowers mostly), pedicel up to 2(–2.5) mm long, glabrous, pistillate flower base or pedicel often shorter and stouter, up to 1.5 mm in diameter; sepals triangular, up to 2 by 1.5 mm, glabrous, greenish cream tinged with red; petals elliptic, 0.5–1.2 by 0.7–1 mm, apex roundish or notched, whitish yellow; disc glabrous. Staminate flowers 3–5 mm in diameter; disc c. 2 mm in diameter; stamens: free part of the filaments up to 0.8 mm long, anthers shortly ellipsoid, c. 0.5 by 0.3–0.4 mm; staminal column c. 1 by 0.3 mm; pistillode conical-ovoid with 2-lobed top, up to 0.7 by 0.4 mm. Pistillate flowers 4–6 mm in diameter; outer disc c. 2.5 mm in diameter, inner one tubular, up to 1 mm long, fully covering the ovary; latter globoid, 0.6–0.7(–1) mm in diameter; styles up to 1.2 mm long, only the very base united, apically deeply bifid. Infructescences with up to 9 fruits. Fruits depressed-ellipsoid, emarginate at apex, bilobate, sometimes obconical at base, 5–7 by 6–7.5 by 7–8 mm, fleshy, pale greenish purple when fresh, dry blackish; endocarps 2, endocarp woody, brown, semigloboid, tapering at base, apex splitting, c. 6 by 4–5.5 by 3–3.5 mm. Seeds semi-globoid with a lateral furrow, c. 3.5–5 by 4.5–5 by 2–2.5 mm, shining reddish black, rugulate.

    Distribution — Burma, Thailand, Andaman & Nicobar Islands, N Malaya (Perlis, Lankawi, Penang), E Java, Lesser Sunda Islands (Bali, Sumbawa, Sumba, Flores, Alor, Timor).

 

Bridpust-map.gif (96561 bytes)

 

Femandez-Villar (1880) mentions this species as seen in Luzon, Philippines. I cannot confirm its occurrence there and assume that he misinterpreted B. tomentosa. Koorders (1912) reports a record by Backer for W Java ('Batavia') which I cannot confirm either.

    Habitat & Ecology — Reported from sandy beaches, savannahs, dry evergreen or deciduous forests and moist monsoon forests; on sandy or calcareous soil or limestone rocks; locally common. Alt.: Sea level up to 400 m.

    Uses — Medicinal use of the leaves is reported several times, e.g., as a purgative (van Beusekom & Santisuk 2728, L; Thailand), against lues (syphilis) (van Beusekom et al. 3398, L; Thailand), and also for wrapping cigarettes (Kerr 4419A, K, L; Thailand).

    Notes — 1. This species can be recognised by its rather large, broadly elliptic and chartaceous leaves with more than 14 pairs of secondary veins and the often conspicuous olive blackish colour in dry state. The branches and leaves are entirely glabrous and the latter are blunt on both ends. It seems to have a rather wide ecological range as the field notes prove. But the peculiar distributional pattern (from Burma and Thailand to the Lesser Sunda Islands, unknown from Sumatra) reveals it as requiring a strong (to severe) dry season. Bridelia curtisii is morphologically and ecologically similar, but distinguishable by having a narrow-oblong leaf-shape (index > 2), a stiffly coriaceus texture, and less than 10 pairs of secondary veins. Bridelia tomentosa has much more slender and longer branches, and mostly smaller and apically acute leaves.

2. An exceptional trilocular fruit was observed in Santisuk 556 (K) from Thailand whereas the other ones were normally bilocular.

3. The inclusion of Australia (Northern Territory) within the distributional range by Airy Shaw (1972) seems to go back to the unification of B. exaltata with B. ovata. No specimens could be traced and Australia is therefore omitted here. The occurrence in the Philippines stated by Fernandez-Villars (1880) could not be confirmed and probably relies on a misidentification. Merrill [Enum. Philipp. Flow. Pl. 2 (1923) 424] excluded the species as well.

4. In the Index Kewensis B. lanceolata was erroneously reported to occur in Malaya, but in the protologue Java is cited.

5. The syntype of B. burmanica in the Wallich Herbarium represents B. ovata but has exceptionally large leaves with c. 20 pairs of secondary veins. However, the other syntype collected by Kurz clearly represents this taxon.

 

6. Bridelia retusa (L.) A.Juss.

 

    Bridelia retusa (L.) A.Juss., Euphorb. Gen. (1824) 109, t. 7, f. 22; Baill., Ιtude Euphorb. (1858) 584, Atlas t. 25, f. 25–34; S.Dressler, Blumea 41 (1996) 289, Map 4; S.Dressler & Welzen in Welzen et al., Thai For. Bull. 28 (2000) 64; in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 148. — Clutia retusa L., Sp. Pl. (1753) 1042; ed. 2 (1763) 1475; Willd., Sp. Pl. 4, 2 (1806) 883. — Bridelia retusa (L.) Spreng., Syst. Veg. 3 (1826) 48, pro comb. nov.; Thwaites, Enum. Pl. Zeyl. (1861) 279; Mόll.Arg. in DC., Prodr. 15, 2 (1866) 493; Bedd., Fl. Sylv. S. India (1872) t. 260; Brandis, Forest Fl. NW India (1874) 449, t. 55; Kurz, Prelim. Rep. For. Pegu, App. A: cix, App. B (1875) 78, in clavi; For. Fl. Burma 2 (1877) 368; Fern.-Vill. in Blanco, Fl. Filipp., ed. 3, Nov. App. 4 (1880) 186; Gamble, Man. Indian Timb. (1881) 595; Trimen, Syst. Cat. Flow. Pl. Ceylon (1885) 78; Hook.f., Fl. Brit. India 5 (1887) 268; G.Watt, Dict. Econ. Prod. India 1 (1889) 536; Kuntze, Rev. Gen. Pl. (1891) 594; Trimen, Handb. Fl. Ceylon 4 (1898) 10; Woodrow, J. Bomb. Nat. Hist. Soc. 12 (1899) 369; Boerl., Handl. Fl. Ned. Indiλ 3 (1900) 271; Prain, Bengal Pl. (1903) 927, repr. (1963) 694; Brandis, Indian Trees (1906) 560; Bourdillon, For. Trees Travancore (1908) 320; Gehrm., Bot. Jahrb. Syst. 41, Beibl. 95 (1908) 30, 33; Talbot, For. Fl. Bombay 2 (1911) 435, f. 488; Craib, Kew Bull. (1911) 457; Aberdeen Univ. Stud. 57 (1912) 183; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 69; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 424 (as excluded spp.); Ridl., Fl. Malay Penins. 3 (1924) 184; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1926) 487; Osmaston, For. Fl. Kumaon (1927) 473; Burkill, Dict. Econ. Pl. Malay Penins. (1935) 366; Merr., Pap. Michigan Acad. Sci. 24 (1939) 76; Talbot, Syst. List Trees Bombay, ed. 3 (1949) 442; Neal, Bernice P. Bishop Special Publ. 50 (1965) 504; Airy Shaw, Kew Bull. 23 (1969) 67; Kew Bull. 26 (1972) 230; Kew Bull. 36 (1981) 274; Grierson & Long, Fl. Bhutan 1 (3) (1987) 769; Pham-hoang Ho, Cayco Vietnam 2 (1992) 292; Saldanha, Fl. Marnataka 2 (1996) 123. — Bridelia retusa (L.) A.Juss. var. genuina Mόll.Arg. in DC., Prodr. 15, 2 (1866) 493, nom. inval. — Bridelia airy-shawii P.T.Li, Acta Phytotax. Sin. 20 (1982) 117, nom. superfl.; Anon., Wealth of India 213 (1988) 295, f. 62. — Lectoype (S.Dressler, 1996): Herb. Hermann 2:  fol. 71, #367 (BM).

    Clutia squamosa Lam., Encycl. 2 (1786) 54. — Bridelia retusa (L.) A.Juss. var. squamosa (Lam.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 493; Hook.f., Fl. Brit. India 5 (1887) 268. — Bridelia squamosa (Lam.) Gehrm., Bot. Jahrb. Syst. 41, Beibl. 95 (1908) 30; Haines, J. Bot. 59 (1921) 190; Anon., Wealth of India 213 (1988) 297. — Bridelia squamosa (Lam.) Gehrm. var. typica Gehrm., Bet. Jahrb. Syst. 41, Beibl. 95 (1908) 30, nom. inval. — Type: Hb. Sonnerat (P?); Hb. Commerson (P?) (n.v.), India oriental.

    Clutia spinosa Roxb., Pl. Corom. 2 (1802) 38, t. 172. — Bridelia spinosa (Roxb.) Willd., Sp. Pl. 4 (1806) 979 (p.p., excl. specim. in B-W # 18957), non Hort. ex DC. (1833), non Wight (1852); Roxb., Hort. Bengal. (1814) 69; Spreng., Syst. Veg. 3 (1826) 48; Roxb., Fl. Ind., ed. Carey, 3 (1832) 735; J.Graham, Cat. Pl. Bombay (1839) 184; Voigt, Hort. Suburb. Cale. (1845) 155; Wall., Numer. List [1847 (1845?)] nr. 7883; P.T.Li, Acta Phytotax. Sin. 26 (1988) 63; Fl. Reipubl. Pop. Sin. 44, 1 (1994) 32, t. 9, f. 1–3. — Lectotype (designated by S.Dressler, 1996): Roxburgh s.n. (Hb. Martius) (BR; BM iso, # 19312, p.p.), without locality.

    Bridelia amoena Wall. ex Baill., Ιtude Euphorb. (1858) 584; Voigt, Hort. Suburb. Calc. (1845) 156, nomen. — Type: Wallich in Hb. Gaudichaud (P?), Calcutta?

    Bridelia crenulata Roxb., [Hort. Bengal. (1814) 70, nomen;] Fl. Ind., ed. Carey, 3 (1832) 734 (descr.). — Bridelia retusa (L.) A.Juss. var. roxburghiana Mόll.Arg. in DC., Prodr. 15, 2 (1866) 493, nom. superfl. — Bridelia roxburghiana (Mόll.Arg.) Gehrm., Bot. Jahrb. Syst. 4l, Beibl. 95 (1908) 30, nom. superfl.; Voigt, Hort. Suburb. Calc. (1845) 155; Hook.f., Fl. Brit. India 5 (1887) 268; Prain, Bengal Pl. (1903) 927, repr. (1963) 694. — Type: Wallich 7880 [BM, G-DC n.v. (IDC microfiche Candolle Prodromi Herbarium 2508.14), K, K-Wall], Botanic Garden Calcutta, from S. Harris anno 1798.

    Bridelia retusa (L.) A.Juss. var. glauca Hook.f., Fl. Brit. India 5 (1887) 268 — Type not designated.

    Bridelia fordii Hemsl., J. Linn. Soc. 26 (1894) 419; P.T.Li, Fl. Reipubl. Pop. Sin. 44, 1 (1994) 34. — Syntypes: Ford 249, 254 (K), China, Kwangtung.

    Bridelia retusa (L.) A.Juss. var. glabra Gehrm., Bot. Jahrb. Syst. 41, Beibl. 95 (1908) 30. — Type not designated.

    Bridelia retusa (L.) A. Juss. var. pubescens Gehrm., Bot. Jahrb. Syst. 41, Beibl. 95 (1908) 30. — Type not designated.

    Bridelia retusa (L.) A.Juss. var. stipulata Gehrm., Bot. Jahrb. Syst. 41, Beibl. 95 (1908) 30. — Type not designated.

    Bridelia.squamosa (Lam.) Gehrm. var. meeboldii Gehrm., Bot. Jahrb. Syst. 41, Beibl. 95 (1908) 30. ― Type not designated.

    Bridelia cambodiana Gagnep., Bull. Soc. Bot. France 70 (1923) 432; in Lecomte, Fl. Indo-Chine 5 (1926) 494, f. 62.12–16, 63.1. — Lectotype (designated by S.Dressler, 1996): Chatillon s.n., 9 Aug. 1909 (P holo; P iso), Cambodge, Prey-kdey.

    Bridelia pierrei Gagnep., Bull. Soc. Bot. France 70 (1923) 434; in Lecomte, Fl. Indo-Chine 5 (1926) 494; Airy Shaw, Kew Bull. 23 (1969) 67; Kew Bull. 26 (1972) 229; Pham-hoang Ho, Cayco Vietnam 2 (1992) 292; P.T.Li, Fl. Reipubl. Pop. Sin. 44, 1 (1994) 32. — Lectotype (designated by S.Dressler, 1996): Pierre 6287 (US holo; BM, BO, BR, E, K, NY, P iso), Cambodia, Chaudoc, Mt Cam.

    Bridelia montana auct. non Willd.: J. Graham, Cat. Pl. Bomb. (1839) 184; Wall., Numer. List [1847 (1845?)] nr 7879, p.p.; Baill., Ιtude Euphorb. (1858) 583; Dalzell & A.Gibson, Bomb. Fl. (1861) 233.

 

(Big shrub to) medium to large-sized tree, up to 10(–20) m high, dbh up to 30 cm, trunk with scattered, up to 4 cm long, pointed thorns, stems somewhat spiny; branches rusty tomentose to woolly when young, later pilose to glabrous. Bark dark brown to blackish, thick, roughly cracked, flaking in scales. Stipules ovate-triangular, to 2 by 2 mm, whitish woolly, early caducous. Leaves: petiole subterete, 7–12 by 1–2.5 mm, sparsely pilose; blade elliptic (to slightly obovate), 63–253 by 30–115 mm, length/width ratio 1.4–2.7, stiffly (sub)coriaceous, turning pinkish brown before falling, base (acute to) (subcordate to) obtuse to rounded, margin entire to shallowly crenate where veins reach margin, apex (emarginate to) acute to obtuse, upper surface glabrous except midvein rarely pilose, dark glossy green; lower surfacelight brownish pubescent to tomentose, pale dull greenish, often glaucous; venation prominent on both sides, conspicuously stronger below, secondary veins in (16–)19–23(–27) pairs, fairly straight to only slightly arching upwards, but often forking before joining the marginal vein, tertiary veins ± patent to secondaries, simple or rarely forked percurrent, conspicuously scalariform. Inflorescences multibracteate glomerules of (3–)8–15 subsessile flowers on usually leafless branches, rarely small (up to 5 by 1 mm), woolly, thick, lanceolate leaves extant, hence of spike-like appearance, mostly terminal on twigs, sometimes also in axils of persisting normal leaves, ‘spikes’ often branched, up to 20 cm long, glomerules mostly of one sex only, but often both sexes in inflorescence; bracts acuminately triangular, 1.5–2 by 1.3–1.5(–2) mm, with dark and abaxially prominent midvein, pilose to tomentose. Flowers: pedicel slender, up to 2 mm long; sepals ovate-triangular, glabrous to pilose at base, rarely also at the apex; petals variable in shape, base spatulate, apex roundish to lobulate with gnawed margin, whitish, 0.8–1.6 by 0.6–1 mm. Staminate flowers 4–5 mm in diameter, pale yellow green to brown, sepals up to 1.7 by 0.8–1 mm; disc 1.2–1.5 mm in diameter, yellow to pale orange; stamens: free part of filaments c. 0.8–1 by 0.1 mm, white, anthers ovoid, c. 0.5–0.7 by 0.3–0.4 mm, reddish to purplish, staminal column 1–1.3 by 0.2–0.4 mm; pistillode narrow ovoid-conical, c. 0.7 by 0.3–0.4 mm diam. Pistillate flowers 5–5.5 mm in diameter, reddish to brown; sepals up to 2 by 1.2–1.5 mm; outer disc up to 2.2 (in fruit 2.7) mm in diameter, inner one up to 1.6 mm long when conical-tubular; ovary globoid, apically tapering into style, 2-locular, 1–1.8 by 0.7–1.6 mm in diameter; 2 styles 0.5–1 mm long, united below, shortly bifid above, stigmas papillose. Infructescences with 1–3 fruits. Fruits depressed-globose, sometimes apical bluntly pointed (remnant of style), sometimes bilobate, 5–8 by 5–9 mm in diameter, smooth, glossy, light green with minute pale greenish dots, ripening dark reddish when fresh,dark to bluish black when dry; endocarps 2, semispherical, brown, woody to stony, c. 5 by 6 by 5 mm. Seeds semiglobular with ventral furrow, c. 4.5 by 5 by 3 mm; testa smooth, reddish brown.

    Distribution — India, Sikkim, Bhutan, Sri Lanka, Burma, Indochina, S China, Thailand, Malay Peninsula (?, see note 3), Sumatra. Fernandez-Villar's (1880) report for the Philippines was already corrected by Merrill (1923).

 

Bridretu-map.gif (65426 bytes)

 

    Habitat & Ecology – Quite common in forests and open land, reported from dry evergreen or deciduous forests; sandy-loamy soil, granite or basalt derived sandy soil, and limestone. Alt.: 50–600(–1400) m. Reported to be resistant to fire in regularly burnt grassy savannah (de Wilde & de Wilde-Duyfjes 18956, L). Fruits are eaten by birds (pigeons) in India (Koelz 29237, L; Neal, 1965: 504).

    Uses — Cf. Wealth of India 2B (1988) 295. The dull red wood is used for construction, railway ties, fuel (Neal, 1965: 504), rafters, posts, and floor-boards; also used for cart-shafts, wheels and agricultural implements; suitable for tool-handles and helves (Useful Pl. India, 1986: 87). Bark contains tannin (16–40%); in pharmacological trials it exhibited antiviral, hypoglycaemic, hypotensive properties (Useful Pl. India, l.c.), used for tanning and medicinal (Burkill, 1935) and as a poison (Neal, l.c.). Leaves used as fodder (Useful Pl. India, l.c.), eaten by cattle, and said to free them from intestinal worms (Voigt, 1845; J. Graham, 1839: 184). Fruit edible (Useful Pl. India, l.c.), very adstringent (Fernandes 28, L).

    Notes — 1. This species is identifiable by its spicate or sparsely branched, leafless or almost leafless, axillary inflorescences which can reach a length of 30 cm, and by its stiff-coriaceous leaves with 15–25 pairs of prominent secondary veins. Some similarities in leaf morphology exist with B. macrocarpa which, however, has its flower glomerules exclusively in the axils of normal leaves and develops big fruits.

2. Linnaeus described Clutia retusa based on material from Sri Lanka. Sprengel in 1826 was always considered to have made the combination in Bridelia. But Jussieu's older homonym Bridelia retusa is based on Clutia retusa sensu Willdenow (1806: 883) who cites Linnaeus' Sp. Pl. (ed. 2, 1763) and most of the Linnean sources. Hence Willdenow's concept is congruent with that of Linnaeus and both are homotypic. I have no idea why Mόller Argoviensis in DC. (1866) synonymised Jussieu's name with B. stipularis. Hence, the selection of B. airy-shawii P.T.Li as a nomen novum is nomenclaturally superfluous.

3. I have no certainty about the occurrence on the Malay Peninsula as I have seen no recent collections. The records in the literature are based on one specimen cited by Ridley (1924), but Whitmore (1973) suspects this to be B. stipularis. However, Hooker (1887) already gave Malacca as part of the range, probably based on specimens from the Wight Herbarium (# 2602, 2604) which have no clear indication of the place of provenance (Peninsula Indiae Orientalis), but which are often from the Malay Peninsula (Welzen, pers. comm.).

4. I saw three collections from the Hawaiian Islands, where the plant is obviously cultivated (cf. Neal, 1965: 504).

5. Willdenow made the new combination Bridelia spinosa based on Roxburgh (1802) but also cited material extant in B-W which represents B. montana Willd.

6. According to the leaf width and form, the indumentum, the shape of the inflorescences and the stipules several varieties were distinguished, but as these features are variable they are all united under B. retusa. In India plants with smaller, stiffer, more acute leaves are often to be found, which have been referred to as B. squamosa. These often stem from higher altitudes or drier localities. The material from Thailand often shows large leaves.

7. Airy Shaw regarded B. pierrei as closely related to but distinct from B. retusa due to its inflorescence structure (fewer flowers per inflorescence, persistent large stipules, etc.). All specimens determined by him as B. pierrei are in a very early stage of their inflorescence development and none is in anthesis or even fruiting. The stipules are still present and often the inflorescence axis is not yet grown out. A fruiting syntype in US has no persistent stipules.

8. Bridelia cambodiana was said to differ mainly by having the inflorescences on leaved branches which occurs sometimes in B. retusa as well. I subsume it under the latter as it perfectly matches otherwise.

 

7. Bridelia stipularis (L.) Blume

 

    Bridelia stipularis (L.) Blume, Bijdr. (1826) 597; Hassk., Cat. Hort. Bot. Bogor. (1844) 240; Baill., Ιtude Euphorb. (1858) 584; Miq., Fl. Ned. Indiλ l, 2 (1859) 364; Fl. Ned. Indiλ, Suppl. (1861) 445; Mόll.Arg. in DC., Prodr. 15, 2 (1866) 499; Miq., Ann. Mus. Lugd.-Bat. 4 (1869) 120; Bedd., Fl. Sylv. S. India, For. Man. (1873) 201; Brandis, Forest Fl. NW India (1874) 449; Kurz, For. Fl. Burma 2 (1877) 369; Fern.-Vill. in Blanco, Fl. Filipp., ed. 3, Nov. App. 4 (1880) 186; Gamble, Man. Ind. Timb. (1881) 596; S.Vidal, Sin. Gen. Pl. Leρos. Filip. (1883) 38, Atlas t. 82 f. F.; Phan. Cuming. Philipp. (1885) 141; Rev. Pl. Vasc. Filip. (1886) 233; Hook.f., Fl. Brit. India 5 (1887) 270; Collett & Hemsley, J. Linn. Soc. 28 (1889) 122; G.Watt, Dict. Econ. Prod. India 1 (1889) 536; Kuntze, Rev. Gen. Pl. (1891) 594; Woodrow, J. Bomb. Nat. Hist. Soc. 12 (1899) 369; Boerl., Handl. Fl. Ned. Indiλ 3 (1900) 271; Talbot, Syst. List Trees Bombay, ed. 2 (1902) 298 (n.v.), ed. 3 (1949) 443; Prain, Bengal Pl. (1903) 928, repr. (1963) 694; Merr., Philipp. J. Sci. 1, Suppl. (1906) 78; Brandis, Indian Trees (1906) 560; Gehrm., Bot. Jahrb. Syst. 41, Beibl. 95 (1908) 29; J.J.Sm., Meded. Depart. Landb. Ned.-Indiλ 10 (1910) 315; Craib, Kew Bull. (1911) 457; Talbot, For. Fl. Bombay 2 (1911) 437, f. 489; Craib, Aberdeen Univ. Stud. 57 (1912) 183; Koord., Exk.-Fl. Java 2 (1912) 485, in clavi; Merr., Fl. Manila (1912) 284; F.C.Gates, Philipp. J. Sci., Bot. 9 (1914) 426; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 55; Merr., Sp. Blanc. (1918) 220; Bibl. Enum. Born. Pl. (1921) 336; Enum. Philipp. Flow. Pl. 2 (1923) 424; Ridl., Fl. Malay Penins. 3 (1924) 183; S.Moore, J. But. 63, Suppl. (1925) 92; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1926) 492; Osmaston, For. Fl. Kumaon (1927) 474; Hand.-Mazz., Symb. Sin. 7 (1931) 221; Merr. & Chun, Sunyatsenia 2 (1935) 264; Burkill, Dict. Econ. Pl. Malay Penins. (1935) 366; Gage, J. Asiat. Soc. Bengal. 75 (1936) 485; Corner, Wayside Trees Malaya, ed. l (1940) 243, ed. 2, 1 (1951) 243, ed. 3, l (1988) 279; Masamune, Enum. Phan. Born. (1942) 390; Backer & Bakh. f., Fl. Java 1 (1964) 475; Anon., Fl. Hainanica 2 (1965) 142; Meijer, Bot. News Bull. Forest Dept, Sabah 7 (1967) 39, in clavi; Corner & Watanabe, Ill. Guide Trop. Pl. (1969) 339; Airy Shaw, Kew Bull. 26 (1972) 230; Whitmore, Tree Fl. Malaya 2 (1973) 74, in clavi; Airy Shaw, Kew Bull., Add. Ser. 4 (1975) 65; Kew Bull. 36 (1981) 274; Kew Bull. 37 (1982) 11; Enum. Euphorb. Philipp. Isl. (1983) 11; Anon., Wealth of India 2B (1988) 296; Grierson & Long, Fl. Bhutan 1 (3) (1987) 769; Whitmore, Tree Fl. Indon., Checkl. Kalimantan 1 (1990) 121; PROSEA (Lemmens & Wulijarni-Soetjipto, eds.) 3 (1992) 133; Pham-hoang Ho, Cayco Vietnam 2 (1992) 292; P.T.Li, Fl. Reipubl. Pop. Sin. 44, 1 (1994) 38, tab. 8, f. 5–7; S.Dressler, Blumea 41 (1996) 2, Fig. 1, Map 5; S.Dressler & Welzen in Welzen et al., Thai For. Bull. 28 (2000) 65, Fig. 3; in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 149, fig. 32. — Clutia stipularis L., Mant. Pl. (1767) 127; Willd., Sp. Pl., ed. 4, 4 (2) (1806) 883 (as Clutya); Blanco, Fl. Filip. (1837) 818; ed. 2. (1845) 564, ed. 3, 3 (1879) 229 (as Clutya). — Bridelia stipularis (L.) Blume var. typica Gehrm., Bot. Jahrb. Syst. 41, Beibl. 95 (1908) 29, nom. inval. — Lectotype (designated by S.Dressler, 1996): LINN 1206.13 (LINN; IDC microfiche LINN Herbarium 1206.13).

    Clutia scandens Roxb., PI. Corom. 2 (1802) 39, t. 173. — Bridelia scandens (Roxb.) Willd., Sp. Pl. 4 (1806) 979; Roxb., Hort. Bengal. (1814) 70; Spreng., Syst. Veg. 3 (1826) 48; Roxb., Fl. Ind., ed. Carey, 3 (1832) 736; J.Graham, Cat. Pl. Bombay (1839) 184; Voigt, Hort. Suburb. Calc. (1845) 156; Wall., Numer. List [1847 (1845?)] nr 7878 (excl. E); Baill., Ιtude Euphorb. (1858) 584; Dalzell & A.Gibson, Bomb. Fl. (1861) 233; Trimen, Syst. Cat. Flow. Pl. Ceylon (1885) 78; Handb. Fl. Ceylon 4 (1898) 11; Gehrm., Bot. Jahrb. Syst. 41, Beibl. 95 (1908) 29; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 55; Saldanha, Fl. Karnataka 2 (1996) 123. — [Scherunam cottam Rheed., Hort. Malab. 2 (1679) t. 16 (sec. Roxb., 1832; Dillwyn, Rev. Hort. Malab., 1839; Nicolson et al., Regnum Veg. 119 (1988) 107]. — Lectotype (designated by S.Dressler, 1996): Roxburgh s.n., s.d. (BR lecto; BM # 19310, K, K-Wall. # 7878A iso).

    Bridelia zollingeri Miq., Fl. Ned. Indiλ 1, 2 (1859) 364. — Type: Zollinger 878 (erroneously cited as 8782) (U, fragment, holo; P iso), Java, Bandung.

    Bridelia dasycalyx Kurz, J. Asiat. Soc. Beng. 42, ii (1874) 241; For. Fl. Burma 2 (1877) 369; Hook.f., Fl. Brit. India 5 (1887) 271; Brandis, Indian Trees (1906) 560; Gehrm., Bot. Jahrb. Syst. 41, Beibl. 95 (1908) 29. — Bridelia dasycalyx Kurz var. genuina Kurz, J. Asiat. Soc. Beng. 42, ii (1873) 241, nom. inval. — Lectotype (designated by S.Dressler, 1996): Kurz 1605 (K holo; BO iso), Burma, Ava, Prome, Pegu.

    Bridelia dasycalyx Kurz var. aridicola Kurz, J. Asiat. Soc. Beng. 42, ii (1874) 241; For. Fl. Burma 2 (1877) 369; Hook.f., Fl. Brit. India 5 (1887) 271. — Lectotype (designated by S.Dressler, 1996): Kurz 2475 (BO), Burma, Brome, Pegu.

    Bridelia stipularis (L.) Blume var. ciliato Gehrm., Bot. Jahrb. Syst. 41, Beibl. 95 (1908) 29, syn. nov. — Type not designated.

    Bridelia stipularis (L.) Blume subsp. philippinensis Jabl. in Engl., Pflanzenr. IV.147.viii (19i5) 57. — Lectotype (designated by S.Dressler, 1996): FB 1884 (Ahern's coll.) (US # 851432 holo; BO, K, NY iso), Philippines, Luzon, Prov. Rizal, Bosoboso.

 

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Scrambling shrub (up to 16–20 m long) or small tree (up to 8 m high), branching sparingly or laxly and pendulous; branches young tomentose, velvety or woolly, later pilose. Outer bark dark purple to brown, slightly flaking; inner bark pale green, soft; sapwood pale white. Stipules (narrowly) triangular to ovate, 3–10 by 2–4 mm, base (cordate to) rounded to truncate, apex long-acuminate, whitish to reddish brown pilose to velvety, rather late caducous. Leaves: petiole subterete, 3–8(–9) by 1(–1.8) mm, ± densely tomentose to velvety; blade (ovate to) elliptic (to obovate), 32–172 by 13–112 mm, length/width ratio 1.2–2.6, chartaceous, base (emarginate to) obtuse to rounded (to acute), margin subentire to faintly crenate where veins reach margin and slightly undulated when dry, apex (rounded to) obtuse to mostly acute, both surfaces with whitish, greenish, reddish or brownish hairs, upper surface pilose to villose, bright dark green, lower surface densely tomentose to softly velvety, at least on veins, otherwise pilose, lighter green to greyish green to often glaucous; venation prominent below, secondary veins in 9–13(–l5) pairs, relatively straight, only very slightly arching and joining the marginal vein, tertiary veins ± patent to secondaries, simple or rarely forked percurrent, scalariform. Inflorescences glomerules of (1–)3–6 usually big (sub)sessile flowers set along the leafless or smaller-leaved ends of string-like twigs, each cluster with several woolly bracts; bracts very variable in shape, broadly triangular to broadly ovate, 3–4 by 3.5–4 mm, apex apiculate, caducous in fruit. Flowers: pedicel stout but short, c. 1 by 2 mm in diameter, hairy; sepals (acuminately) triangular, 4–5 by 2–3 mm, with variable indumentum from plain over base or apex hairy to densely tomentose, sometimes even adaxially; petals very variable in shape, (1.5–)2–3 by (1–)1.5–2.5 mm, base cuneate to spatulate, apex roundish to lobulate. Staminate flowers cream to greenish or yellowish, 6–10 mm in diameter; disc plain, 5–6 mm in diameter; stamens: free part of filaments c. 1.5 by 0.1 mm, anthers ellipsoid, c. 1 by 0.5 mm, purple, staminal column c. 2 by 0.5 mm; pistillode conico-ovoid with 2- or 4-merous top, c. 1 by 0.5 mm. Pistillate flowers reddish green, up to 12 (in fruit 14) mm in diameter; outer disc 4–6 mm in diameter, inner one c. 1.5 mm long, inside mostly hairy, hence after splitting a ring of hairs or hairy scales around fruit base; ovary ovoid to globoid, 1.5–2 by 1.5–2 mm, styles 1.5–3 mm long, apex bifid (c. 1 mm). Infructescences with mostly 1 or 2, up to 5 fruits. Fruits ellipsoid to subglobose or ovoid, sometimes barrel-shaped, often bilobate, 7–12 by 6–11 mm in diameter, blunt to pointed at apex with remnants of style,glossy glaucous green with red dots, turning dull dark reddish to black, with fleshy mesocarp splitting between 2 endocarps; latter horny, light brown, splitting loculicidally from apical, after dehiscence receptacle with persistent sepals, petals, hairy disc-scales, and columella-like structure. Seeds semi-ovoid, dorsally keeled, ventrally sulcate, subapical hilum, 6–7 by 3–5.5 by 2–3.5 mm, shiny, striate, reddish brown; embryo flat; cotyledons obovate, simply folded; endosperm thin.

    Distribution — Sri Lanka, India, Nepal, SE Asia mainland, Malay Peninsula, Sumatra, Java, N Borneo, Philippines (unknown from Mindanao), Lesser Sunda Islands (Bali, Lombok, Sumba, Flores, Alor, Timor).

 

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    Habitat & Ecology — Primary and secondary forests, often near wet places (rivers, swamps, seashore, mangrove); soil: mostly reported from sandy soil, but also from limestone or loamy soil above igneous rocks. Alt.:sea level up to 400(–1100) m.

    Uses — Cf. Wealth of India 2B (1988) 296. The plant is said to be poisonous (Airy Shaw, 1975, Maidin 235, Borneo). Twigs used for basketry (Useful Pl. India, 1986: 87). Bark used as medicine against intestinal worms (Hohenacker 345, L) and for tanning (PROSEA 3, 1992, 133). Decoction of bark used for cough, fever, and asthma; also shows hypotensive hypoglycaemic action on animals (Useful Pl. India, l.c.). Leaves medicinal (cf. Burkill, 1935), infusion of the leaves against colic on Java in the vicinity of Bogor (K. Heyne, Nutt. Pl. Ned. Ind., ed. 3,1950, 918), used for jaundice (Useful P1. India, 1. c.). Fruits are said to be edible (Airy Shaw, 1975; Useful Pl. India, l. c.; Borneo: Cuadra A264) and to produce a sticky fluid when wounded (van Beusekom et al. 4420, Thailand) and to yield a black colouring matter (PROSEA, l. c.). Seeds possess hemaglutinating properties and yield a fatty oil (Useful Pl. India, l.c.).

    Notes — 1. The leaves are very variable in size and shape; those subtending inflorescence clusters are often smaller and shorter petioled. There are many collections with obovate leaves from the Lesser Sunda Islands and with subrotundate ones from Thailand. Nevertheless, the species is easily recognisable by its abaxially tomentose to velvety leaves having only 9–14 pairs of secondary veins. Other diagnostic characters comprise the relatively large flowers (up to 12 mm diam.) with the hairy inner disc in the pistillate ones and the large, often barrel-shaped, dehiscing fruits (only the fruits of B. macrocarpa are bigger) as well as the scrambling habit.

2. Some collections, mostly from the Asian mainland, lack the hairs inside the inner pistillate disc and hence the hairy ring at the fruit base. Bridelia scandens was based mainly on this feature (beside a generally more delicate appearance, smaller leaves and flowers, and differing indumentum of the sepals, cf. Jablonszky, 1915). Nearly all these collections are very variable and intergrading, and, e.g. in Thailand, both character states of disc hairiness occur together so that I agree with all previous authors who considered both taxa to be conspecific.

3. I place B. stipularis subsp. philippinensis into synonymy under the species as I cannot observe any distinction of the Philippine material, neither in indumentum nor in leaf size, from specimens collected elsewhere. Both these characters are extremely variable.

4. Globose galls on the lower leaf surface are often found; the are resembling valvate flower buds, up to 5 mm diam.; cf. Docters van Leeuwen-Reijnvaan, Marcellia 8 (1909) 89 (sec. Jablonszky, 1915: 57).

5. Pseudococcus virgatus (Cockerell) - Coccidae - a mealy bug was found on this species in Luzon, Manila, Malate, April 1917 (Philipp. J. Sci., D, 13, 1918, 146).

 

8. Bridelia tomentosa Blume

 

    Bridelia tomentosa Blume, Bijdr. (1826) 597; Hassk., Cat. Hort. Bot. Bogor. (1844) 240; Wall., Numer. List [1847 (1845?)] nr 7874; Benth., Hook. J. Bot. Kew Gard. Misc. 6 (1854) 8; Seem., But. Voy. Herald (1857) 410; Baill., Ιtude Euphorb. (1858) 584; Hook.f., Fl. Tasman. (1859) xlvi; Miq., Fl. Ned. lndiλ 1, 2 (1859) 364; Fl. Ned. Indiλ, Suppl. (1861) 445; Benth., Fl. Hongk. (1861) 309; Mόll.Arg. in DC., Prodr. 15, 2 (1866) 501; Benth., Fl. Austral. 6 (1873) 120; Kurz, Prelim. Rep. For. Pegu (1875) App. A: cix., App. B.: 78, in clavi; For. Fl. Burma 2 (1877) 367; Fern.-Vill. in Blanco, Fl. Filipp., ed. 3, Nov. App. 4 (1880) 186; Gamble, Man. Ind. Timb. (1881) 596; S.Vidal, Rev. Pl. Vasc. Filip. (1886) 234; Hook.f., Fl. Brit. India 5 (1887) 271; G.Watt, Dict. Econ. Prod. India I (1889) 537; Kuntze, Rev. Gen. Pl. (1891) 594; Forbes & Hemsl., J. Linn. Soc. 26 (1894) 419; Henry, List Pl. Formos. (1896) 82 (= Trans. Asiat. Soc. Japan 24, Suppl.: 82); Boerl., Handl. Fl. Ned. Indiλ 3 (1900) 271; F.M.Bailey, Queensl. Fl. 5 (1902) 1411; Prain, Bengal Pl. (1903) 928, repr. (1963) 694; Hayata, J. Coll. Sci. Tokyo 20 (1904) 30, t. 3A; Matsum. & Hayata, J. Coll. Sci. Tokyo 22 (1906) 362; Brandis, Indian Trees (1906) 560; Gehrm., Bot. Jahrb. Syst. 41, Beibl. 95 (1908) 32; J.J.Sm., Nova Guinea 8 (1910) 231; Meded. Depart. Landb. Ned.-Indiλ 10 (1910) 320; Ridl., J. Straits Branch Roy. Asiat. Soc. 59 (1911) 167; Craib, Kew Bull. (1911) 457; Aberdeen Univ. Stud. 57 (1912) 183; Koord., Exk.-Fl. Java 2 (1912) 485, in clavi; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 58, f. 11A; W.Fitzg., J. Proc. Roy. Soc. W. Austral. 3 [1918 (1916-1917?)] 163; Merr., Bibl. Enum. Born. Pl. (1921) 336; Ridl., Fl. Malay Penins. 3 (1924) 184; S.Moore, J. Bot. 63, Suppl. (1925) 92; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1926) 488; Gage, J. Asiat. Soc. Bengal. 75 (1936) 487; M.R.Hend., J. Malayan Br. Roy. As. Soc. 17 (1939) 69; Corner, Gard. Bull. Str. Settlem. ser. 3, 10 (1939) 291; Corner, Wayside Trees Malaya, ed. 1 (1940) 243, ed. 2, 1 (1951) 243, f. 72, ed. 3, 1 (1988) 280, f. 78; Masam., Enum. Phan. Born. (1942) 390; F.G.Browne, For. Trees Sarawak (1955) 177; Corner & Watanabe, Ill. Guide Trop. Pl. (1969) 338; Airy Shaw, Kew Bull. 26 (1972) 231; Kew Bull. Add. Ser. 4 (1975) 65; Kew Bull. 31 (1976) 382; Hsieh, Fl. Taiwan 3 (1977) 452; ed. 2, 3 (1993) 430; Airy Shaw, Kew Bull. Add. Ser. 8 (1980) 45; Kew Bull. 35 (1980) 602; Muelleria 4 (1980) 223; Kew Bull. 36 (1981) 274; Kew Bull. 37 (1982) 11; Enum. Euphorb. Philipp. Isl. (1983) 11; Grierson & Long, Fl. Bhutan 1 (3) (1987) 770; P.T.Li, Acta Phytotax. Sin. 26 (1988) 64; Anon., Wealth of India 2B (1988) 297; Hnatiuk, Austral. Fl. Fauna Ser. 11 (1990) 180; Chapman, Austral. Fl. Fauna Ser. 12 (1991) 477; P.T.Li, Fl. Reipubl. Pop. Sin. 44, 1 (1994) 30, 31, t. 8, f.1–4; S.Dressler, Kew Bull. 51 (1996) 607, Map 2; Blumea 41 (1996) 297, Fig. 2, Map 6; P.I.Forst., Austrobaileya 5 (1999) 413. — Lectotype (designated by Airy Shaw, 1980): Blume s.n. (BO holo; BM, BR, L, NY iso), Java, Buitenzorg (Bogor).

    Bridelia louteirii Hook. & Arn., Bot. Beechey Voy. (1837) 211, excl. syn. Lour. (Mόll.Arg. in DC., 1866: 501). — Type: Lay & Collie s.n. (Captain Beechey) (K?).

    ?Bridelia urticoides Griff., Not. Pl. Asiat. 4 (1854) 481 (sec. Index Kewensis). — Type: Griffith s.n. (not traced, CAL?), Burma, Mergue, ad littoram Ins. Madamaca.

    Bridelia monoica sec. Merr., Philipp. J. Sc., Bot. 13 (1918) 142. — vix Clutia monoica Lour. (sec. Corner, 1939: 291 f.); Merr., Enum. Philipp. Flow. Pl. 2 (1923) 423; Lingnan Sci. J. 5 (1927) 109; Trans. Amer. Philos. Soc. II, 24 (2) (1935) 234; Burkill, Dict. Econ. Pl. Malay Penins. (1935) 365; Kaneh., Formosan Trees (1936) 333, f. 287; K.Heyne, Nutt. Pl. Ned. Indiλ ed. 3 (1950) 918; Keng, Taiwania 6 (1955) 38; H.L.Li, Woody Fl. Taiwan (1963) 419; Salvosa, Lex. Philipp. Trees (1963) 89; Backer & Bakh.f., Fl. Java 1 (1964) 475, in clavi; Anon., Fl. Hainanica 2 (1965) 142, f. 365; Airy Shaw, Kew Bull. 26 (1972) 228, 231, in clavi; Whitmore, Tree Fl. Malaya 2 (1973) 74, in clavi; Pham-hoang Ho, Cayco Vietnam 2 (1992) 290.

 

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Erect to scrambling shrub or small tree, up to 13 m high, up to 15 cm dbh; branches very slender, often long and whip-like, scrambling, spreading or drooping, glabrous to tomentose, with scattered lenticels, old branches sometimes thorny. Bark smooth to slightly roughened, inner bark pink. Stipules early caducous, ovate to narrowly triangular, 2–4(–5) by c. 1 mm, sometimes aristate at apex, pubescent to tomentose. Leaves often rather small; petiole terete, glabrous to tomentose, (2–)3–5.5 by 0.8–1.5 mm; blade elliptic to obovate, 25–140 by 10–60 mm, length/width ratio (1.7–)2–3.7, thinly chartaceus, base obtuse (to acute), margin entire, rarely shallowly crenate where lateral veins reach the margin, apex (bluntly) acute (to shortly acuminate, upper surface glabrous to scattered appressed hairs, dull dark green, lower surface glabrous to tomentose, pubescent on main nerves, dull green to whitish green, often conspicuously glaucous; venation (weakly) prominent, secondary veins in 7–12(–15) pairs, continuously arching and joining the marginal vein, tertiary veins percurrent, irregularly scalariform, irregular reticulate areolation. Inflorescences small multibracteate glomerules of up to 10(–20) sessile green flowers; bracts broadly triangular, up to 0.7 by 0.5 mm, puberulous, with ciliate margin. Flowers scented, faint musty smell, sessile, rarely with up to l mm long glabrous stalk (pistillate flowers stouter stalked than staminate ones); sepals triangular, glabrous, greenish yellow; petals very variable in shape, base cuneate to spatulate, apex roundish, notched or lobulate, up to 1 by 0.8 mm, whitish yellow; disc glabrous. Staminate flowers 2–3 mm in diameter; sepals 0.8–1.2 by 0.5–0.8 mm; disc 1–1.2 mm in diameter; stamens: free part of filaments up to 0.5 mm long, anthers shortly ellipsoid, 0.4–0.5 by 0.3–0.4 mm; staminal column c. 0.7 mm long; pistillode conical-ovoid, up to 0.4by 0.2 mm. Pistillate flowers 3–4 mm in diameter; 1.2–1.5 by 0.8–1.2 mm; outer disc c. 2.2 mm in diameter, inner one tubular, up to 0.6 mm long, covering Ύ of the ovary, margin sometimes erose; ovary globoid, c. 1 mm in diameter, style completely divided into 2 arms, very short, to 0.5 mm long, stigma lobed. Infructescences with up to 7 fruits. Fruits (sub)globoid, sometimes depressed and laterally compressed, emarginate at apex, slightly bilobate, 4.5–6.5 mm in diameer, fleshy, greenish to blue, blackish when dry; endocarps 2, semigloboid, c. 4 by 5 by 2.5 mm, woody, brown, apically splitting. Seeds semigloboid to broadly semi-tear-shaped, with deep lateral furrow, 3–3.5 by c. 4 by 2–2.5 mm, brown, rugulate.

    Distribution — India (Madras, Bengal, Assam-Khasi Hills, Bihar, Orissa), E Nepal, Sikkim, S China, Hainan, Taiwan, Myanmar, Indochina, Thailand, Andaman (Middle and South) and Nicobar Islands, Malay Peninsula, Sumatra, Java, Borneo (SE Kalimantan), Philippines (unknown from Mindanao), Celebes, Lesser Sunda Islands (Sumbawa, Sumba, Flores, Alor, Timor), Moluccas (Tanimbar & Kei Is.), New Guinea, N Australia; common throughout its range, but rare in Borneo.

 

Bridtome-map.gif (70139 bytes)

 

    Habitat & Ecology — Deciduous to evergreen forests, primary to secondary vegetation; reported from sandy or loamy soil, limestone, and over granite bedrocks. Alt.: sea level up to 1000 m. Reported to be rather common on waste land and in secondary forests (Browne, 1955: 177, Corner, 1988: 280). Crown full of butterflies (KEP FRI 13000, L, Malacca).

    Uses — Cf. Wealth of India 2B (1988) 297. Timber tree: wood suitable for baskets, carts, wheels, and tool-handles (Heyne, 1950: 918; Useful Pl. India, 1986: 87), for report of wood quality cf. Browne (1955: 177), for chemical compounds of the stem cf. Wealth of India, l.c. Tannin-producing plant: bark astringent, used in colic, also for tanning (Useful PI. India, l.c.), used to colour wood black (Meijer 5363, L, Sumatra). Infusion of leaves against colic (Heyne, l.c.). Fruits eaten (Useful Pl. India, l.c.). Seeds used by children for bullets in bamboo guns (Hu 5013, K). Well known in village medicine, in folk tales (cf. Corner, 1988: 280).

    Notes — 1. The characteristic features of this variable and widespread species comprise the long, slender, often whip-like branches, the elliptic, rather small and narrow, pointed leaf with conspicuous whitish glaucous undersurface, the small (mostly < 10 flowers) inflorescences and the the small flowers (staminate 2–3 mm, pistillate 3–4 mm in diameter).

2. The present species forms, together with B. curtisii and B. harmandii, a species complex, which is often difficult to disentangle. The main differences from B. curtisii are the acute to acuminate leaf tip, the often glaucous undersurface of the leaf, its weaker texture, as well as the smaller flowers. Bridelia curtisii shows a rather distinct leaf shape. Nevertheless intergrading specimens occur (e.g., Lφrzing 16248, L, Sumatra). Bridelia harmandii, which occurs in Thailand and Indochina, differs in its stiffer coriaceous, rather broad leaves, an always strong indumentum in many parts (also outside the calyx), rather persistent subulate stipules, and less flowers per glomerule.

3. Within the area studied a considerable variation could be found in the indumentum of the leaves and to a lesser extent in the leaf shape. Such a diversity is not very surprising within such a widespread and ecologically adaptable species. In the Philippines an accumulation of absolutely glabrous plants could be observed. This glabrous form is referred to as B. tomentosa var. glabrescens Benth. and together with the type variety these are the only infraspecific taxa which could be observed in Indochina and Malesia (see below). The varieties trichadenia Mόll.Arg. and eriantha Airy Shaw are so far only known from Australia (Northern Territory) (Airy Shaw, Kew Bull. 35, 1980).

4. One aberrant trilocular fruit was seen (Mizushima & Liao 10885, L, Taiwan). Such an exception occurs rarely in several species of the genus (cf. however B. triplocarya) and was already reported for B. tomentosa by Hooker & Arnott (1837: 211; under B. loureirii).

5. The fact that the widespread B. tomentosa was not collected in Borneo (except in the very Southeast: Ambriansyah & Arifin AA 1079; Motley 493; Winkler 2635, 2965, 2973, 3023) seems worth mentioning and could be explained by the relatively drier climate there. In New Guinea the species is apparently only found in the south as well. Bridelia tomentosa is obviously a plant showing a seasonal drought pattern of distribution.

 

This species is represented in Southeast Asia by two varieties (a. var. tomentosa and b. var. glabrescens), which are widespread all over the area. The latter variety is characterised by lack of an indumentum (especially on the undersurface of the leaf). Airy Shaw (1976: 383) already pointed out, intermediate forms with minute traces of pubescence do occur.

 

a. var. tomentosa

 

    Bridelia tomentosa Blume var. tomentosa: S.Dressler, Blumea 41 (1996) 302. — Bridelia tomentosa Blume var. genuina Mόll.Arg. in DC., Prodr. 15, 2 (1866) 501, nom. inval. — Amanoa tomentosa (Blume) Baill., Adansonia 6 (1866) 336. — Type as the species.

    Bridelia lancifolia Roxb., [Hort. Bengal. (1814) 70, nom. nud., (‘lanceofolia’)] Fl. Ind. ed. Carey, 3 (1832) 737 (‘lanceaefolia’); Wall., Numer. List [1847 (1845?)] nr. 7884; Airy Shaw, Kew Bull. 26 (1972) 231 (‘lanceifolia’); Kew Bull. Add. Ser. 4 (1975) 65; Kew Bull., Add. Ser. 8 (1980) 45. — Lectotype (designated by S.Dressler, 1996): Roxburgh s.n. (BM holo; K iso).

    Bridelia rhamnoides Griff., Not. Pl. Asiat. 4 (1854) 480. — Type: Griffith s.n. (K, CAL n.v.), Burma, in sylvis Mergue, Oct. 1834.

    Bridelia tomentosa Blume var. rhamnoides Mull. Arg. in DC., Prodr. 15, 2 (1866) 502. — Type: Hoffmannsegg s.n. [B, †, & in hb. Franquev. (P?), fragm. G-DC (IDC microfiche Candolle Prodromi Herbarium 2510.18)], Java.

    Bridelia tomentosa Blume var. chinensis Mόll.Arg. in DC., Prodr. 15, 2 (1866) 501. — Syntypes: Park s.n. (G-DC n.v.; IDC microfiche Candolle Prodromi Herbarium 2510.8), China; Motley 493 (K), Borneo prope Banjarmassing.

    Bridelia tomentosa Blume var. ovoidea Benth., Fl. Austral. 6 (1873) 120. — Type: Gulliver s.n. (K), Australia, Northern Territory, Wood Island.

    Bridelia phyllanthoides W.Fitzg., J. Proc. Roy. Soc. W. Austral. 3 (1918) 163. — Type: Fitzgerald 823 (NSW, n.v.), W Australia, base of Mt Broome.

 

b. var. glabrescens Benth.

 

    Bridelia tomentosa Blume var. glabrescens Benth., Hook. J. Bot. Kew Gard. Misc. 6 (1854) 8; S.Dressler, Blumea 41 (1996) 302. — Type: Champion s.n. (K?), Hongkong, East Point, Hedges.

    Bridelia glabrifolia Merr., Enum. Philipp. Flow. Pl. 2 (1923) 422; Salvosa, Lex. Philipp. Trees (1963) 88. — Bridelia tomentosa Blume var. glabrifolia (Merr.) Airy Shaw, Kew Bull. 31 (1976) 383; Airy Shaw, Kew Bull. 35 (1980) 603. — Bridelia tomentosa Blume var. lanceaefolia sec. Mόll.Arg. in DC., Prodr. 15, 2 (1866) 502 (non Bridelia lanceaefolia Roxb.). — Bridelia lancaefolia sec. Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 60, (non Bridelia lanceaefolia Roxb.). — Lectotype (designated by Airy Shaw, 1980): Gaudichaud s.n. (G-DC; IDC microfiche Candolle Prodromi Herbarium 2510.20), Phlippines, Manila.

    Bridelia ovata auct. non Decne.: Benth., Fl. Austral. 6 (1873) 120; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 61.

    Note — The first description of a glabrous form of B. tomentosa has unfortunately been overlooked, so that the commonly known name (var. glabrifolia) has to be changed to var. glabrescens. The type could not be traced in Kew.

 

9. Bridelia triplocarya Airy Shaw

 

    Bridelia triplocarya Airy Shaw, Kew Bull. 32 (1978) 385; Kew Bull. Add. Ser. 8 (1980) 45; S.Dressler, Kew Bull. 51 (1996) 617, Map 2; Blumea 41 (1996) 303. — Type: Schodde 2768 (K holo; BO, CANB, E, L, SING iso), Papua, Central Distr., 1.5 km N of Rigo.

 

Slender tree, c. 7 m high, dbh c. 8 cm; branches slender, scattered pilose to puberulous. Outer bark dark grey, rough, furrowed; wood pink cream outside, dark brown inside. Stipules early caducous, narrow-triangular, subulate, c. 3 by 0.3 mm, whitish puberulous. Leaves: petiole subterete, 4–8 by 0.7–1.3 mm, scabrous to sparsely puberulous; blade elliptic, 4–11 by 1.5–4.5 cm, length/width ratio 1.9–3, coriaceous, base obtuse to acute, margin entire to rarely distantly shallowly crenate where veins reach margin, apex (obtuse to) acute, upper surface glabrous slightly glossy green; lower surface light puberulous to pubescent, almost glaucous; venation prominent on both sides, stronger below, secondary veins in 10–14 pairs, steeply ascending, fairly straight, only slightly arching, but often forking before joining the fimbrial vein, tertiary veins irregular percurrent, simple or rarely forked, sometimes scalariform, reticulate areolation. Inflorescences paucibracteate glomerules of 2–5 sessile flowers; bracts (depressed) triangular, tiny, up to 0.8 by 1 mm, whitish pilose to puberulous. Staminate flowers not seen. Pistillate flowers 4–5 mm in diameter in fruit; pedicel indistinct, up to 0.5 mm long; sepals triangular, 1.4–1.8 by c. 1.3 mm, glabrous; petals variable in shape, triangular to lobulate with gnawed margin, 0.7–1 by 0.5–0.7 mm, base spatulate, apex roundish; scales of inner disc 1–1.5 mm long; ovary not seen. Infructescences with 1 or 2 almost sessile fruits, up to 0.5 mm stalked. Fruits globose, 8–9 by 6–8 mm in diameter, (2- or) 3-locular, smooth, dull pink maroon, bluish black when dry, glaucous, conspicuously stipitate (1–2 mm) by tapering fruit base and often 6–9-ribbed; endocarps with the shape of a spherical segment with acute base, ventrally carinate, dorsally with costa, margin erosely winged, c. 6.5 by 8 by 3.5 mm. Seeds flat-semigloboid, plano-convex with shallow ventral furrow, base subacutely tapering, 3.5–4 by 4–5 by 1–1.5 mm; testa reddish brown.

    Distribution — New Guinea, so far only known from the type collection.

    Habitat & Ecology — Monsoon hill forest. Alt.: c. 15 m.

    Notes — 1. Bridelia triplocarya usually has trilocular fruits, which is very unusual in the genus. Exceptionally, fruits with three locules can be found in some other species (e.g., B. ovata, B. macrocarpa), but normally they are characteristic for Cleistanthus, where they are developed as fully dehiscing woody capsules. Although B. triplocarya shows three locules it is clearly a Bridelia because of its indehiscent drupes with copious fleshy mesocarp. A fimbrial vein is also not found in Cleistanthus. Apart from the fruit this species is characterised by the stiffly coriaceous leaves which are puberulous beneath and show a prominent venation on both sides. It seems to be a very rare endemic in New Guinea.

2. This taxon seems to have some affinities with B. macrocarpa and B. erapensis, both from New Guinea. In the former species trilocular fruits are occasionally found, but these have plum-like dimensions. The leaves are larger and have more secondary vein pairs. They have normally only a pilose indumentum beneath. Also the inflorescences are much larger, with 8–16 flowers. The latter species, however, has ovate leaves with an often cordate base, and bilocular fruits.

3. In leaf shape and the number of fruits per infructescence there is a superficial resemblance between B. triplocarya and B. exaltata from E Australia as Airy Shaw already noted. But the venation pattern, the leaf texture, and the fruits clearly distin­guish both taxa.

 

Subgenus Gentilia

 

    Bridelia subg. Gentilia (Beille) Jabl. in Engl., Pflanzenr. Heft 65 (1915) 71; S.Dressler, Blumea 41 (1996) 304. — Gentilia Beille, Compt. Rend. Hebd. Seances Acad. Sci. 145 (1907) 1294; G.L.Webster, Ann. Missouri Bot. Gard. 81 (1994) 39 (in synonymy). — Bridelia sect. Monospennae Gehrm, Bot. Jahrb. Syst. 41, Beibl. 95 (1908) 12. — Bridelia sect. Gentilia (Beille) Gehrm., Jahresber. Schles. Ges. Vaterl. Cult. 1908 (1909) 28, nom. superfl. — Lectotype species (designated by G.L.Webster, 1994): Gentilia hygrophila Beille (= Bridelia ndellensis Beille).

 

10. Bridelia adusta Airy Shaw

 

    Bridelia adusta Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 63, 224 (Latin diagnosis); Kew Bull. 32 (1978) 383; Anderson, Checkl. Trees Sarawak (1980) 179; Whitmore, Tree Fl. Indon., Checkl. Kalimantan 1 (1990) 120; S.Dressler, Blumea 41 (1996) 304, Map 7. — Type: S (Anderson) 30854 (K holo; A, E, L, SAN iso), Borneo, Sarawak, Baram Distr., Gunong Api.

 

Tree, up to 18 m high, girth c. 80 cm; branches glabrous, only very young twigs puberulous, prominently lenticellate. Bark smooth, greyish brown to green; inner bark brownish; sapwood pale yellow. Stipules caducous, but often present on younger twigs, narrow ovate-triangular to subulate, 5–8 by c. 1 mm, with scattered rusty hairs to puberulous. Leaves: petiole subterete, (3–)4–7 by 1–1.5 mm, glabrous to scattered hairy; blade slightly ovate to slightly obovate, 40–100 by 15–35 mm, length/width ratio 2.4–3.2, stiffly coriaceous, chartaceous when young, base broadly cuneate to subrotundate, margin entire, conspicuously revolute, apex acuminate, acumen up to 15 mm long, upper surface glabrous, rarely a few scattered hairs along the main veins, shiny olive green; lower surface glabrous, light green, slightly glaucous, often conspicuously cinnamon-brownish when dry; venation: secondary veins in 7–9(–11) pairs, mostly not prominent on adult leaves, rarely slightly prominent if leaf is not fully developed, sometimes darker than lamina, slightly bent and obliquely running towards the margin, distally narrowed and inconspicuously joining the next secondaries, no distinct fimbrial vein, margin cartilaginous, tertiary veins irregularly percurrent, ultimate venation reticulate. Inflorescences multibracteate glomerules of up to 10 (sub)sessile flowers, axil sometimes leafless, but not at special leafless branches; bracts triangular, tiny, up to 1 by 1.5 mm, margin minutely ciliate. Staminate flowers not seen. Pistillate flowers ± perigynous, 2.5–4 mm in diameter, whitish pink; pedicel at most 1 by 1 mm, puberulous like basal part of receptacle; sepals (narrowly) ovate to triangular, 1.5–2.5 by 1–1.5 mm, brownish puberulous outside; petals whitish, obovately spathulate, c. 1–1.3 by 0.8–1 mm, base ± narrowed, apex erose; outer disc 1.5–2 mm in diameter, inner one crateriform, c. 0.5 mm long, completely surrounding the ovary, margin erose; ovary ovoid, apically terminating into a style, up to 1 by 0.8 mm in diameter; styles very short, c. 0.5 mm long, shortly bifid at apex, not exserted. Fruits subgloboid to ovoid, acute or blunt at apex, 7–8 by 5–8 mm in diameter. Seed one per fruit, not seen.

    Distribution — N Borneo (so far known from two localities only).

 

Bridadus-whit-map.gif (95017 bytes)    Distribution of B. adusta (dots; B. whitmorei = squares).

 

    Habitat & Ecology — In primary forests, among limestone boulders. Alt.: 800–1800 m

    Notes — 1. This species was described as a rare endemic from Sarawak and is, so far, only known from two collections, one from the original tree and another one from Sabah.

2. This species clearly belongs to sect. Cleistanthoideae with its one-seeded drupes and absent fimbrial vein, and not to sect. Scleroneurae as was erroneously mentioned by Airy Shaw. Morphologically it is rather close to B. glauca. The distinguishing features are: stiffly coriaceous leaves, revolute margin, less secondary veins (7–9 pairs only), its more acute leaf base, and subsessile flowers and fruits.

3. More material is needed to corroborate the distinction of this taxon which is undoubtedly not that much isolated as Airy Shaw thought, putting it into the wrong section. But admittedly the combination of coriaceous leaves and few secondary veins is unique in sect. Cleistanthoideae as well.

 

11. Bridelia cinnamomea Hook. f.

 

    Bridelia cinnamomea Hook.f., Fl. Brit. India 5 (1887) 273; Boerl., Hand]. Fl. Ned. Indiλ 3 (1900) 271; Gehrm., Bot. Jahrb. Syst. 41, Beibl. 95 (1908) 38; Ridl., Fl. Malay Penins. 3 (1924) 185; Gage, J. Asiat. Soc. Bengal. 75 (1936) 490, 492; Whitmore, Tree Fl. Malaya 2 (1973) 74, in clavi; Tree Fl. Indon., Checkl. Kalimantan 1 (1990) 120; S.Dressler, Blumea 41 (1996) 307, Fig. 3, Map 9; S.Dressler & Welzen in Welzen et al., Thai For. Bull. 28 (2000) 61, Fig. 2; in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 142, fig. 31. — Bridelia griffithii Hook.f. var. cinnamomea (Hook.f.) Gehrm., Bot. Jahrb. Syst. 41, Beibl. 95 (1908) 38; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 64; Anderson, Checkl. Trees Sarawak (1980) 179; Airy Shaw, Kew Bull. 36 (1981) 273. ― Lectotype (designated by Gage, 1936): King's collector 7101 (K holo; SING iso), Malaya, Kinta (erron. Kinla in Hook.f., 1887).

    Bridelia griffithii Hook.f., Fl. Brit. India 5 (1887) 272, incl. var. griffithii; Brandis, Indian Trees (1906) 561; Gehrm., Bot. Jahrb. Syst. 41, Beibl. 95 (1908) 38; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 74. — Lectotype (designated by S.Dressler, 1996): Griffith KD 4883 (K holo; U iso), Malacca.

    Bridelia ovata auct. non Decne.: Kurz, For. Fl. Burma 2 (1877) 368 (sec. Hook.f., 1887).

    Bridelia gehrmannii Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 73; Merr., Bibl. Enum. Born. Pl. (1921) 335; Masam., Enum. Phan. Born. (1942) 390. — Lectotype (designated by S.Dressler, 1996): Haviland & Hose 1858 (L, holo; BM, E, K, L, SING iso), Borneo, Sarawak, Kuching.

    Bridelia moonii auct. non Thwaites: Gehrm., Bot. Jahrb. Syst. 41, Beibl. 95 (1908) 37 as to the Bornean reference (sec. Merrill, 1921).

 

Scrambling shrub, rarely small tree, up to 6 m high; branches often creeping, greyish to brwon, whitish lenticellate, reddish brown pubescent to tomentose, glabrescent. Outer bark thorny, brownish grey; inner bark fibrous, yellow green. Indumentum reddish brown. Stipules early caducous, widely ovate, 1.5–2 by c. 2 mm, brownish pubescent. Leaves: petiole terete, 4–8 by c. 1 mm, whitish to brownish pubescent or tomentose; blade elliptic (to slightly obovate), 40–120 by 18–65 mm, length/width ratio 1.5–2.4, chartaceous, base (bluntly rounded to) acute, margin ± entire, apex rounded to shortly acuminate, acumen 3–7 mm long, upper surface glabrous except pilose on the veins, deep green; lower surface puberulous to pubescent, rarely subglabrous, bluish to geryish green with pale brownish venation; latter visible above, prominent beneath, secondary veins in (5–)6–9(–10) pairs, especially at the base with a rather acute angle of divergence, distally arching towards the margin and joining the next secondaries, not the weak marginal vein, tertiary veins percurrent, scalariform, ± right-angled to midvein. Inflorescences multibracteate glomerules of 10–20 subsessile to pedicelled, brownish red, yellow- to green-centred flowers, normally not terminal and not at special leafless branches; bracts narrowly triangular, up to 4 by 1 mm, brownish tomentose to woolly. Flowers 4–5 mm in diameter, smelling (Lφrzing 16440, L); pedicel up to 3 by 0.5 mm, brownish pubescent; sepals triangular, 1.5–2 by 1.2–1.5 mm, puberulous, at least at base outside; petals variable in shape, c. 1 mm by 0.7–0.8 mm, base spathulate, apex roundish to lobulate, greenish yellow with pink base. Staminate flowers: disc c. 1.8 mm in diameter; stamens: free part of filaments 1–1.2 by c. 0.1 mm, anthers shortly semigloboid, c. 0.4–0.5 mm in diameter, staminal column c. 1 mm long; pistillode conical-ovoid, 0.7–0.8 by 0.3–0.6 mm, blunt at apex, bifid. Pistillate flower: petals outside pubescent to tomentose and somewhat fleshy; disc c. 1.8 mm in diameter, wrinkled, inner one 0.7 mm long when tubular, scales c. 1 mm long; ovary globoid, apex conically pointed, c. 1.2 by 1.2 mm; styles c. 1 mm long, deeply bifid (upper third), not exserting flower, stigma papillose. Infructescences with c. 3 or 4 subsessile fruits. Fruits ovoid to globoid, tapering or only pointed at apex (remnant of styles), 6–7 by 5–5.5 mm in diameter, glossy yellow green, tinged reddish, dark brown when dry; endocarp woody, glossy. Seeds one per fruit, (semi-)ellipsoid with lateral furrow, 4–5 by 2.3–3.5 mm; testa light brown.

    Distribution — Thailand, Malay Peninsula, Sumatra, Borneo (known only from Sabah and Sarawak).

 

Bridcinn-map.gif (89876 bytes)

 

    Habitat & Ecology — Primary and secondary mixed peat swamp forests; occasional. Alt.: Sea level up to 600 m.

    Notes — 1. This species can easily be identified by its rather small leaves with acute base and reddish indumentum below, by the comparatively few (6–9) lateral nerves that have a conspicuously acute angle of divergence in the basal part of the leaf. Especially the staminate flowers are often shortly pedicelled and the styles are not exserting in the pistillate ones.

2. It belongs to a group of closely related species together with B. insulana, B. glauca, and B. pustulata. Morphologically, the latter seems closest related, although it is distinguishable by several features (tree, leaf size and form, number of secondary veins, big protruding inflorescence cushions, pistillate inner disc sometimes hairy outside).

3. Apparently, B. cinnamomea is confined to swampy areas and ranges in lower altitudes.

4. The material distributed by Kew as Griffith (Kew Distr. 4883) is obviously heterogeneous: Specimens labelled Griffith (Kew Distr. 4883) =Maingay 1371 represent the type collection of B. pustulata and the ones solely named Griffith 4883 the type collection of B. griffithii (= B. cinnamomea).

 

12. Bridelia glauca Blume

 

    Bridelia glauca Blume, Bijdr. (1826) 597; Span., Linnaea 15 (1841) 347; Hassk., Cat. Hort. Bot. Bogor. (1844) 240; Miq., Fl. Ned. Indiλ l, 2 (1859) 364; D.Dietrich, Syn. Pl. 5 (1852) 383; Baill., Ιtude Euphorb. (1858) 584; Mόll.Arg. in DC., Prodr. 15, 2 (1866) 497; Boerl., Handl. Fl. Ned. Indiλ 3 (1900) 271; J.J.Sm., Meded. Depart. Landb. Ned.-Indiλ 10 (1910) 307; Hallier f., Meded. Rijks-Herb. 1 (1910) 7; Koord., Exk.-Fl. Java 2 (1912) 485, in clavi; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 74; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 423; Pap. Michigan Acad. Sci. 24 (1939) 76; Holthuis & H.J.Lam, Blumea 5 (1942) 200; K.Heyne, Nutt. Pl. Ned. lndiλ, ed. 3 (1950) 918; Salvosa, Lex. Philipp. Trees (1963) 89; Backer & Bakh.f., Fl. Java 1 (1964) 475, in clavi; Meijer, Bot. News Bull. For. Dept., Sabah 7 (1967) 39, in clavi; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 64; Kew Bull. Add. Ser. 8 (1980) 43; Anderson, Checkl. Trees Sarawak (1980) 179; Airy Shaw, Kew Bull. 36 (1981) 273; Kew Bull. 37 (1982) 10; Enum. Euphorb. Philipp. Isl. (1983) 11; Whitmore, Tree Fl. Indon., Checkl. Kalimantan 1 (1990) 120; S.Dressler, Kew Bull. 51 (1996) 606, Map 3; Blumea 41 (1996) 311, Fig. 4, Map 10; S.Dressler & Welzen in Welzen et al., Thai For. Bull. 28 (2000) 63; in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 145. — Lectotype (designated by Mόller Argoviensis, 1866): Blume s.n. (Lholo; L, NY iso), in insulae Javae in sylvis ad pedem montis Salak.

    Bridelia multiflora Zipp. in Miq., Ann. Mus. Lugd.-Bat. 4 (1869) 119; Koord., Versl. Minahasa (1898) 582; Boerl., Handl. FI. Ned. Indiλ 3 (1900) 271; Hallier f., Meded. Rijks-Herb. 1 (1910) 7. — Type: Zippelius (235c?) (L holo; BO iso), Amboina.

    Bridelia pubescens Kurz, J. Asiat. Soc. Beng. 42, ii (1874) 241; Kurz, Prelim. Rep. For. Pegu (1875) App. A: cix., App. B.: 78, in clavi; For. Fl. Burma 2 (1877) 367; Hook.f., Fl. Brit. India 5 (1887) 270; Prain, Bengal Pl. (1903) 928, repr. (1963) 694; Brandis, Indian Trees (1906) 560; Gehrm., Bot. Jahrb. Syst. 41, Beibl. 95 (1908) 37; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 73; Parija & Misra, J. Indian Bot. Soc. 12 (1933) 227; Merr., Pap. Michigan Acad. Sci. 24 (1939) 76; Hurusawa, J. Fac. Sci. Univ. Tokyo, Sect. 3, Bot. 6 (6) (1954) 322; Airy Shaw, Kew Bull. 26 (1972) 230; Grierson & Long, Fl. Bhutan 1 (3) (1987) 770. — Lectotype (designated by S.Dressler, 1996): Kurz 2474 (K holo; BM, K iso), Burma: Pegu, Yomah.

    Cleistanthus myrianthoides C.B.Rob., Philipp. J. Sci., Bot. 6 (1911) 325. — Type: FB (Curran) 10679 (US # 709742 iso), Philippines, Luzon, Prov. Camarines, Caramoan, San Roque.

    Bridelia lauraefolia Elmer, Leafl. Philipp. Bot. 7 (27 March 1915) 2637. — Lectotype (designated by S.Dressler, 1996): Wenzel 411 (BM holo; GH # 45887 iso), Philippines, Mindanao, Leyte, Dagami.

    Bridelia glauca Blume f. laurifolia Jabl. in Engl., Pflanzenr. IV.147.viii (22 June 1915) 75; Salvosa, Lex. Philipp. Trees (1963) 89. — Lectotype (designated by S.Dressler, 1996): FB (Curran, Merrill & Zschokke) 18195 (L holo; K, NY iso), Philippines, Prov. Benguet, Mt Pulong.

    Bridelia nooteboomii Chakrab., J. Econ. Taxon. Bot. 5 (1984) 949. — Type: Nooteboom 4761A (PBL n.v., holo; L, PBL n.v., iso), Thailand, Northern Region, NW of Chiengmai, Doi Pui.

 

Bridglau-habit.gif (51652 bytes)    Bridglau-fruit.gif (25832 bytes)

 

(Shrub to) tree, up to 30 m high, with a clear bole up to 18 m high, 40 cm dbh, sometimes with a few buttresses, up to 1.5 m high, 3 m out, 10 cm thick, basal trunk sometimes provided with root-thorns; branches reddish brown pubescent to tomentose (especially when young), inconspicuously lenticellate, older branchlets thorny. Outer bark dark greyish brown, to 5 mm thick, smooth, rarely fissured, not peeling off; inner bark yellowish white, up to 4 mm thick; sapwood up to 5 cm thick, yellowish white to orange, no exudate; heartwood ochre to dark brown. Stipules ovately triangular, 8–12 by 3–5(–6) mm, plain green, brownish pubescent, caducous, but often present on younger twigs. Leaves: petiole subterete, 5–11 by 1–2.5 mm, pubescent; blade elliptic, slightly ovate (to elliptic), 40–280 by 20–120 mm, length/width ratio 1.9–3.1, (membranaceous to) chartaceous, base acute, obtuse, or (often) truncate, margin entire, apex acute to acuminate, acumen up to 25 mm long, upper surface glabrous, rarely along the main veins scattered hairy, dark green, sometimes glossy, lower surface light brownish to reddish pubescent, rarely glabrous or tomentose, light green, slightly glaucous beneath, brownish when dry; venation: secondary veins in (7–)11–18(–20) pairs, prominent beneath, continuously bent towards the margin, joining the next secondaries, tertiary veins weakly percurrent, ± right-angled to secondaries, only slightly prominent, ultimate venation reticulate. Inflorescences multibracteate glomerules of many (up to 50) conspicuously pedicelled, rarely only subsessile flowers, not terminal and not at special leafless branches; bracts triangular, < 1 mm long, densely brownish pubescent, very inconspicuous (lens!). Flowers 3–5 mm in diameter, yellowish green; pedicel (1–)2–6 by 0.3–2 mm, densely pubescent, rarely glabrous, pedicel of pistillate flowers sometimes stouter; sepals (narrowly) triangular, 1.5–2.3 by 1–1.5 mm, brownish pubescent or puberulous outside; petals lanceolate, spathulate, cuneiform or rectangular, 0.6–0.8 by 0.2–0.5 mm, base ± narrowed, whitish; disc brownish. Staminate flowers: disc 1.5–2 mm in diameter, yellow whitish; stamens: free part of filaments up to 1 mm long, very slender, whitish; anthers ellipsoid to slightly ovoid, 0.5–0.6 by 0.3–0.4 mm, pale yellow, staminal column 0.8–1.5 mm long; pistillode conical cylindrical, blunt at apex, up to 0.5 by 0.2 mm. Pistillate flowers ± perigynous; outer disc 1.2–1.5 mm in diameter, inner one crateriform, c. 0.5 mm long, completely surrounding the ovary, with a ring of few hairs inside, the pistillate flowers; ovary globose to slightly ovoid, apically terminating into the style, 0.6–0.7 by 0.3–0.4 mm in diameter, styles up to 1.2 mm long, lower half united, apical branches of styles bifid, slightly exserting. lnfructescences with up to 8 fruits. Fruits (conical) ellipsoid, acute or blunt at apex, 5.5–10(–12) by 4.5–7.5(–9) mm in diameter, pale purple, turning black with some light (white or pink) spots, bluish black when dry; mesocarp juicy, sweetish; endocarp woody, ellipsoid, 5–8 by 4–8 mm. Seed one per fruit, ellipsoid with shallow lateral furrow, 4.5–6.5 by 3–4.5 mm; testa rugulate, light greenish brown.

    Distribution — India, Bhutan, Sikkim, Burma, Thailand, Laos, Taiwan, Malay Peninsula, Sumatra, Java, Borneo, Philippines (not found on Palawan), Celebes, Moluccas (Talaud Is., Morotai, Halmahera, Obi, Bacan = Batjan, Taliabu Is., Ambon, Ceram), New Guinea; Bismarck Archipelago (New Britain).

 

Bridglau-map.gif (76040 bytes)

 

    Habitat & Ecology — In (sometimes disturbed) primary and secondary forests, often near rivers; reported from well-drained volcanic as well as from sandy, loamy, or clayey soil; over sandstone, limestone, grey schists, or granite; scattered to common. Alt. up to 1500 m.

    Uses — Timber tree, wood hard and durable, used for house and bridge construction (J.J.Sm., 910; K.Heyne, 1950), good house posts and fuel in Mindanao, Philippines (PNH 97554, L; Zwickey 708, NY); used in the Agri ritual on Mindanao (PHN 36045, L). Fuits said to be sweetish and edible (Prawiroatmodjo 422, Borneo).

    Notes — 1. This species can be easily identified by its mostly conspicuously pedicelled flowers, its often truncate leaf base, by the (in sect. Cleistanthoideae) comparatively large number of lateral veins (11–18), and by the perigynous flowers, where the inner disc has a ring of hairs inside and the styles are only slightly exserted. It resembles B. insulana a little, but is nevertheless distinguishable by its larger and pedicelled flowers and the greater number of secondary veins. Because of the tiny bracts the deflowered glomerules resemble smooth cushions and do not have the rather rugged appearance of those of B. insulana. The endemic B. moonii from Sri Lanka shares the number of secondary vein pairs (12–18) with B. glauca but has sessile flowers and fruits and a mostly acute leaf base.

2. The collections from the Asian mainland were always referred to as B. pubescens. However, I could not trace any feature distinguishing it from B. glauca. Some collections (including the type of B. pubescens) have rather short pedicels only (often due to an early stage of inflorescence development) but others are distinctly pedicelled and no hiatus could be traced. Both taxa are therefore considered to be conspecific.

3. The occurrence of the rather uncommon phenomenon of root-thorns was described for B. pubescens (Parija & Misra, 1933) and probably the trunk spines reported in some other Bridelia spp. are of the same origin.

4. A careful study of the isotype of B. nooteboomii in L revealed that the species does not belong to sect. Scleroneurae at all as stated by Chakrabarty. The fruits are obviously unilocular and the secondary veins do anastomose before the margin. Both features clearly indicate that the species belongs to sect. Cleistanthoideae. I cannot find any character, which distinguishes the taxon from B. pubescens (= B. glauca). A relation to B. tomentosa as stated by the original author is absolutely not evident.

 

Three varieties may be distinguished under Bridelia glauca:

 

― Leaves glabrous beneath   

b. var. acuminatissima

― Leaves pubescent beneath     

a. var. glauca

― Leaves rufous-tomentose beneath

c. var. sosopodonica

 

 

a. var. glauca

 

    Bridelia glauca Blume var. glauca: S.Dressler, Blumea 41 (1996) 314. — See further under the species.

 

b. var. acuminatissima (Merr.) S.Dressler

 

    Bridelia glauca Blume var. acuminatissima (Merr.) S.Dressler, Blumea 41 (1996) 314. — Bridelia acuminatissima Merr., Philipp. J. Sci., Bot. 9 (1914) 473; Enum. Philipp. Flow. PI. 2 (1923) 422; Airy Shaw, Enum. Euphorb. Philipp. Isl. (1983) 11. — Type: M.Ramos 1551 (A, BM, L, NY, SING iso), Philippines, Luzon, Prov. Camarines, Mt Isarog.

 

Bridelia acuminatissima described from the Philippines is based mainly on its rather glabrous and slenderly acuminate leaves. These characters are not confined to this area only and occur in various combinations elsewhere. The leaf shape is often variable and the leaf apex of pubescent forms of B. glauca is sometimes rather acuminate too, but the lack of an indumentum is considered worth taxonomic ranking. Therefore, I reduced this taxon to a variety under B. glauca especially as it shares important other features (e.g. pedicelled flowers) with the latter.

 

c. var. sosopodonica (Airy Shaw) S.Dressler

 

    Bridelia glauca Blume var. sosopodonica (Airy Shaw) S.Dressler, Blumea 41 (1996) 315. — Bridelia sosopodonica Airy Shaw, Kew Bull. 23 (1969) 67; Kew Bull. Add. Ser. 4 (1975) 65; Whitmore, Tree Fl. Indon., Checkl. Kalimantan l (1990) 121. , Blumea 41 (1996) 314. —  Type: SAN (Sinanggul) 38274 (K holo; L iso), Sabah (N Borneo), Ranau Distr., Bukit Sosopodon, Kundasang, near Mile 38.

 

Specimens with a stronger rufous tomentose indumentum originally described as a very local Bornean endemic (B. sosopodonica Airy Shaw) appear to occur also elsewhere (e.g., India, Laos, Java, Philippines, Moluccas) and the other distinguishing features mentioned by Airy Shaw fall within the variability of B. glauca (stipule and fruit size). Therefore, this taxon is reduced to varietal rank with the strongly developed indumentum as distinguishing character, although I have to admit that specimens intergrading with B. glauca var. glauca occur.

 

13. Bridelia insulana Hance

 

    Bridelia insulana Hance, J. Bot. 15 (1877) 337; Gehrm., Bot. Jahrb. Syst. 41, Beibl. 95 (1908) 33; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 63; P.T.Li, Acta Phytotax. Sin. 26 (1988) 63; Fl. Reipubl. Pop. Sin. 44, 1 (1994) 37, p.p.; S.Dressler, Kew Bull. 51 (1996) 605, 606, Map 3; Blumea 41 (1996) 315, Map 11; P.I.Forst., Austrobaileya 5 (1999) 410; S.Dressler & Welzen in Welzen et al., Thai For. Bull. 28 (2000) 63; in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 146. — Type: Pierre 19762 (BM holo; K, P? iso), Cochinchina (= S Vietnam), ins. Phukok.

    Bridelia penangiana Hook.f., Fl. Brit. India 5 (1887) 272; Boerl., Handl. Fl. Ned. Indiλ 3 (1900) 271; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 75; Ridl., Fl. Malay Penins. 3 (1924) 185; Burkill, Dict. Econ. Pl. Malay Penins. (1935) 365; Gage, J. Asiat. Soc. Bengal. 75 (1936) 492; Airy Shaw, Kew Bull. 26 (1972) 229; Whitmore, Tree Fl. Malaya 2 (1973) 75, in clavi; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 64; Kew Bull. 31 (1976) 382; Kew Bull. Add. Ser. 8 (1980) 44; Kew Bull. 35 (1980) 602; Muelleria 4 (1980) 223; Anderson, Checkl. Trees Sarawak (1980) 179; Airy Shaw, Kew Bull. 36 (1981) 273; Kew Bull. 37 (1982) 11; Enum. Euphorb. Philipp. Isl. (1983) 11; Whitmore, Tree Fl. Indon., Checkl. Kalimantan 1 (1990) 120; Hnatiuk, Austral. Fl. Fauna Ser. 11 (1990) 180; Chapman, Austral. Fl. Fauna Ser. 12 (1991) 477. — Bridelia griffitthii Hook.f. var. penangiana (Hook.f.) Gehrm., Bot. Jahrb. Syst. 41, Beibl. 95 (1908) 38. — Type: Curtis 527 (K holo; SING iso), Malaya, Penang, on Government Hill.

    Bridelia minutiflora Hook.f., Fl. Brit. India 5 (1887) 273; Koord., Versl. Minahasa (1898) 581; Boed., Handl. Fl. Ned. Indiλ 3 (1900) 271; Brandis, Indian Trees (1906) 561; J.J.Sm., Nova Guinea 8 (1910) 231; Meded. Depart. Landb. Ned.-Indiλ 10 (1910) 310; Koord., Exk.-Fl. Java 2 (1912) 485, in clavi; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 76; Merr., Bibl. Enum. Born. Pl. (1921) 335; Enum. Philipp. Flow. Pl. 2 (1923) 423; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1926) 493; Merr., Univ. Calif. Publ. Bot. 15 (1929) 155; C.T.White, Proc. Roy. Soc. Queensl. 47 (1936) 80; Holthuis & H.J.Lam, Blumea 5 (1942) 200; Masam., Enum. Phan. Born. (1942) 390; K.Heyne, Nutt. Pl. Ned. Indiλ, ed. 3 (1950) 918; Salvosa, Lex. Philipp. Trees (1963) 89; Backer & Bakh.f., Fl. Java 1 (1964) 475, in clavi; Meijer, Bot. News Bull. For. Dept., Sabah 7 (1967) 39, in clavi; Hyland, Card Key Rain For. Trees North Queensl. (1971) 71; Peekel, Fl. Bismarck Archip. (1984) 297, f. 481; Chapman, Austral. Fl. Fauna Ser. 12 (1991) 477; Pham-hoang Ho, Cayco Vietnam 2 (1992) 289. — Lectotype (designated by Airy Shaw, 1980): Griffith KD 867 (K holo; K iso), Burma, Tenasserim, Mergui.

    Bridelia ovata auct. non Decne.: Merr., Philipp. Bur. For. Bull. 1 (1903) 30 (sec. Merrill, 1923). 

    ?Bridelia griffithii Hook.f. var. glabra Gehrm., Bot. Jahrb. Syst. 41, Beibl. 95 (1908) 38. — No type designated.

    Bridelia minutiflora Hook.f. var. abbreviate J.J.Sm., Meded. Depart. Landb. Ned.-Indiλ 10 (1910) 313. — Lectotype (designated by Dressler, 1996): Koorders 25549 (L holo; BO iso), Java, Prov. Preanger.

    Bridelia platyphylla Merr., Philipp. J. Sci., Bot. 7 (1912) 384. — Type: FB (Aguilar) 11167 (type fragm: US # 706710), Philippines, Luzon, Prov. Bulacan, Angat.

    ?Bridelia palauensis Kaneh. in Kaneh. & Hatus., Bot. Mag. Tokyo 53 (1939) 152; Kaneh., J. Dept. Agric. Kyushu Imp. Univ. 4 (1935) 347. — Type: S. Hatusima 4803 (not traced), Palau, Peliliu.

    Bridelia morotaea Airy Shaw, Kew Bull. 37 (1982) 10. — Type: bb (Tangkilisan 123) 33815 (K holo; A, BO, BR, CALC, L, LAE, M, P, S iso), Moluccas, Morotai, Subdistr. Tobelo, N Totodokoe.

    Bridelia nicobarica Chakrab. & Vasudeva Rao, J. Econ. Taxon. Bot. 5 (1984) 945. — Type: N.G.Nair 3523A (PBL holo; L, PBL (nr. 3523B) iso), Nicobar Islands, Sawai.

 

Medium to large tree, 10–25 m high, dbh 25–60 cm; crown big, diffusely spreading; bole clear or multistemmed, sometimes buttressed; branches slender, glabrous, inconspicuously lenticellate, sometimes with a tendency to climbing, older branchlets thorny. Outer bark grey-brown, cracking, scaly; inner bark reddish to brownish, fibrous with granular wedges; sapwood whitish to yellowish to pale orange; blaze odour cucumber-like (Hyland 3437, L). Stipules early caducous, very narrowly triangular, up to 5 by 0.8–1.3 mm, sparsely brownish puberulous. Leaves when crushed have a distinct strong almond odour (Kajewski 2116, L); ; day-old foliage with foul smell like Hedyotis (Takeuchi et al. 4389, L); petiole subterete, 4–7(–8) by 1–1.8 mm, glabrous; blade (broadly) elliptic (to slightly obovate), 45–210 by 25–88 mm, length/width ratio 1.5–2.7, chartaceous (to subcoriaceus), base obtuse, rounded to acute, margin entire, apex obtuse to shortly acuminate, acumen up to 15 mm long; upper surface glabrous, dark green, sometimes glossy, lower surface glabrous, rarely slightly puberulous, then especially at veins, slightly glaucous above, brownish when dry; venation: secondary veins in 9–13(–14) pairs, prominent beneath, continuously bent towards the margin, joining the next secondaries, tertiary veins weakly percurrent, ± right-angled to secondaries, only slightly prominent, ultimate venation reticulate. Inflorescences multibracteate glomerules of many (15–30) sessile to shortly pedicelled flowers, forming cushion-like clusters, not terminal and not at special leafless branches; bracts triangular, c. 1.2 by 1.2 mm, brownish puberulous with ciliate margin. Flowers faintly scented (Hyland 7801, L; BSIP 12981, L); pedicel at most 1.5 mm long, sparsely puberulous; sepals triangular, c. 1.2 mm by 1 mm, brownish puberulous outside; petals elliptic, trilobed or irregularly lobed, tiny, 0.3–0.5 by 0.3–0.5 mm, base cuneate. Staminate flowers 2–2.5 mm in diameter, creamy yellow; disc brown 1–1.2 mm in diameter; stamens: free part of filaments up to 0.8 mm long, anthers ovoid to semigloboid, c. 0.4 mm in diameter, dark red, staminal column c. 1 mm long; pistillode conical ovoid, blunt at apex, up to 0.5 by 0.2 mm. Pistillate flowers 2–3(–3.5) mm in diameter, whitish cream; disc red, outer one c. 1.5 mm in diameter, inner one 0.5–0.7 mm long, completely surrounding the ovary; ovary ovoid, apically terminating into style, c. 1 by 0.7 mm, two branches of styles 1–1.5 mm long, shortly bilobed at apex, exserting. Infructescences with up to 10 fruits. Fruits ellipsoid to ovoid, pointed at apex, 6–11 by 4–6 mm in diameter, green ripening red, then purple-black, bluish black when dry; mesocarp rather thinl; endocarp woody. Seed one per fruit, ovoid, tapering at apex, 6–7 by 3.5–4.5 mm in diameter; testa greenish brown.

    Distribution — Burma, Thailand, Indochina, Nicobar Islands, Malay Peninsula, Sumatra, Borneo, Philippines, Palau Islands, Java, Lesser Sunda Islands (Bali, Flores, Sumbawa), Moluccas (Morotai), New Guinea, Bougainville, Solomon Islands, Sta. Cruz Group, New Hebrides (Espirito Santo), Australia (Queensland).

 

Bridinsu-map.gif (82967 bytes)

 

This species is reported to occur in China (P.T.Li, 1988, 1994), but no relevant specimens could be traced, these records probably refer to B. balansae.

    Habitat & Ecology — Primary and secondary rain forests, often along rivers, but also on dry land, sandy to loamy or clayey soil, or on limestone. Alt.: Sea level up to 1500 m.

    Uses — Timber sometimes used in house-building, Minahasa, Celebes (Koorders, 1898; J.J.Sm., 1910; Airy Shaw, 1980), as house posts (Kajewski 2116, L, Bougainville), for knife handles; as glossy black (Kostermans 18190, L, Sumbawa); bark used for colouring the saguwer (palm wine) red (Holthuis & H.J.Lam, 1942); fruits reported to be edible in Borneo and Indochina (Tukirin 501, L; Gagnepain, 1926); native medicine for headache in Sabah (Tandom 2817, L; Airy Shaw, 1975), decoction of leaves is applied as a lotion against itch (Burkill, 1935); cover for smokes, New Guinea (NGF 14854, L).

    Notes —1. Characteristics of this species are the (broadly) elliptic leaves with often acute to obtuse base and 9–13 pairs of secondary veins only. The leaves are mostly glabrous beneath except for a few collections from E Malesia. The most outstanding feature, however, is formed by the very small flowers (up to 3 mm diam. only), whereas the calyces of B. glauca and B. balansae show diameters from 4–6 mm (up to 7 mm in fruit). The inflorescences have numerous flowers, and when these are fallen off the inflorescence-cushions are of a rather rugged appearance due to the large bracts. Bridelia balansae is distinguishable by its fewer but larger flowers with the pistillate flowers always shortly and stoutly pedicelled and the mostly stiffer, rather elliptic leaves. This species shares the rather low number of secondary vein pairs (8–13) with B. insulana as opposed to B. glauca. The latter differs by having conspicuously stalked flowers, an often truncate leaf base, and usually pubescent undersurfaces of the leaves.

2. Examining the holotype it proved that there can be no doubts about the conspecifity of B. insulana, a long neglected but older name, with B. penangiana, a commonly accepted name of this widespread species, which unfortunately leads to a name change. P.T.Li (1988) already recognised the necessity of this nomenclatural alteration.

3. Bridelia minutiflora var. abbreviata is conspecific with B. minutiflora (= B. insulana) as the supposedly distinguishing characters (very short style and staminal column, less regular venation, bracts weaker hairy) fall within the normal variability and are even not consistent in the type material.

4. Merrill based his B. platyphylla on its larger leaves with 12–16 pairs of secon­dary veins, although he confirmed the occurrence of leaves half as long with only 8­10 vein pairs. The type in US shows one leaf with 12 and several with 8–9 pairs. Thus this taxon was already synonymised with B. minutiflora Hook. f. by Merrill himself in 1923. Airy Shaw (1972, 1975) therefore cited it as a synonym under B. penangiana. In his Philippines checklist (1983), however, he referred to it as B. glauca Blume. He probably came to this conclusion after examining one paratype of B. platyphylla (BS 14557, K). The isotype from US, however, clearly belongs to B. insulana.

5. Bridelia morotaea is conspecific as the supposedly distinguishing characters (smaller, adaxially shining, subcoriaceous leaves with acuminate apex) fall within the normal variability of B. insulana.

 

This species seems to have a broad ecological amplitude, which is also reflected in its wide range and finds its expression in the morphological variability. Most of the Australian specimens are of a more xerophytic appearance (e.g., more prominent higher venation).

 

Two varieties are recognised under this species:

― Leaves glabrous beneath

a. var. insulana

― Leaves pubescent beneath 

b. var. subnuda

 

a. var. insulana

 

    Bridelia insulana Hance var. insulana: S.Dressler, Blumea 41 (1996) 320. — See further under the species.

 

b. var. subnuda (K.Schum. & Lauterb.) S.Dressler

 

    Bridelia insulana Hance var. subnuda (K.Schum. & Lauterb.) S.Dressler, Blumea 41 (1996) 230. — Bridelia subnuda K.Schum. & Lauterb., Fl. Schutzgeb. Sόdsee (1900) 393. — Bridelia penangiana Hook.f. var. subnuda (K.Schum. & Lauterb.) Airy Shaw, Kew Bull. Add. Ser. 8 (1980) 45. — Neotype (S.Dressler, 1996): Tappenbeck 64 (WRSL hololecto; B? † holo), New Guinea, Kaiser-Wilhelmsland, Ramufluss.

 

This taxon differs from B. insulana var. insulana, which is typically glabrous, only by having an indumentum on the abaxial side of the leaves and ± hairy petioles and young branchlets. It seems to be restricted to New Guinea and neighbouring islands, but intermediate forms could be found in the Philippines too. The indumentum of the lower leaf surface is rather variable so that no sharp limits can be drawn between both varieties.

 

14. Bridelia pustulata Hook.f.

 

    Bridelia pustulata Hook.f., Fl. Brit. India 5 (1887) 271; Boerl., Handl. Fl. Ned. Indiλ 3 (1900) 271; Gehrm., Bot. Jahrb. Syst. 41, Beibl. 95 (1908) 38; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 74; Ridl., Fl. Malay Penins. 3 (1924) 185; Burkill, Dict. Econ. Pl. Malay Penins. (1935) 366; Gage, J. Asiat. Soc. Bengal. 75 (1936) 489, 492; Airy Shaw, Kew Bull. 23 (1969) 67; Whitmore, Tree Fl. Malaya 2 (1973) 74, in clavi; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 64; Kew Bull. 36 (1981) 274; Whitmore, Tree Fl. Indon., Checkl. Kalimantan 1 (1990) 120; S.Dressler, Blumea 41 (1996) 320, Fig. 5, Map 12. — Lectotype (designated by S.Dressler, 1996): Maingay KD 1371 (K holo; BM, GH, K, L iso), Malacca.

 

Bridpust-habit.gif (65236 bytes)    Bridpust-male.gif (42804 bytes)    Bridpust-female.gif (52835 bytes)    Bridpust-fruit.gif (82598 bytes)

 

Tree, up to 20 m high; trunk with stiltroots and sharp thorns (2.5–3 cm long); bole clear; crown dense; branches conspicuously light brownish lenticellate, rusty brown pubescent when young. Outer bark dark brown, smooth, scaly, or fissured, with acrid smell; inner bark reddish to pale brown, fibrous; sapwood white brown, merging into darker brown, hard, heavy heartwood. Stipules early caducous, linear to very narrowly triangular, up to 8 by 1.5 mm, sparsely brownish pubescent. Leaves conspicuously blackish brown when dry; petiole terete, 7–11 by 1.2–2.8 mm, brownish pubescent, rarely nearly glabrous; blade (ovate to) elliptic to obovate, 90–230 by 30–115 mm, length/width ratio 1.6–3, chartaceous when young to (sub)coriaceous, base obtuse to cuneate, margin entire, apex obtuse (to shortly acuminate, acumen to 12 mm long, upper surface glabrous but pilose on veins, lower surface brownish pilose to pubescent; venation darker above, prominent beneath, secondary veins in 11 or 12 pairs, continuously bent towards the margin, joining the next secondaries, tertiary veins percurrent, scalariform, ± patent to secondaries. Inflorescences multibracteate glomerules of many (> 20) sessile flowers, forming big cushion-like clusters with the pistillate flowers surrounded by staminate ones, normally not terminal and not at special leafless branches; bracts ovate-triangular, 1.5–1.8 by up to 2 mm, brownish pubescent to tomentose. Flowers 3–4 mm in diameter, pink tobrownish red; pedicel at most 0.3 mm long; sepals narrowly triangular, 1.5–1.7 by 0.8–1 mm, brownish puberulous, especially at base and apex; petals elliptic, tiny, 0.5–0.8 by 0.3–0.6 mm, apex sometimes irregularly lobed, white with darker centre (vein). Staminate flowers: disc c. 1.2 mm in diamete; free part of filaments up to 0.8 mm long, anthers ovoid to semigloboid, c. 0.4 mm in diameter, dark red, staminal column c. 1.2 mm long; pistillode conical-ovoid, blunt at apex, up to 0.5 by 0.3 mm.Pistillate flowers: outer disc 1.5–1.7 mm in diameter, inner one c. 0.5 mm long and sometimes hairy outside; ovary globoid, c. 0.6 by 0.5 mm in diameter, styles c. 2.3 mm long, united in lower quarter, shortly bilobed at apex, exserting, stigma lobate. Infructescences with up to 10 fruits, sometimes with pedicel up to 1.5 mm long. Fruits ellipsoid to obovoid, blunt at apex, sometimes remnants of styles present, 7–8 by 5–7 mm in diameter, green ripening red then purple-black, bluish black when dry; mesocarp rather copious; endocarp woody, 6–7 by 4.5–5 mm. Seed one per fruit, ellipsoid with deep lateral furrow, 4 by 5 mm; testa light brown, granulate to rugose.

    Distribution — Malay Peninsula, Sumatra, Borneo, Philippines.

 

Bridovat-map.gif (99454 bytes)

 

    Habitat & Ecology — Primary forests. Alt.: Sea level up to 350 m.

    Uses — Timber tree, used for posts in native houses because of its durability (Burkill, 1935; Jansen et al. (eds.), PROSEA, Basic List of Species, 1991, Wageningen).

    Notes — 1. This species is characterised by the following features: big, protruding cushion-like inflorescence clusters with many sessile small flowers (more than 20); conspicuously blackish brown leaves in dry state with dark, prominent, scalariform tertiary venation; pistillate flowers with the inner tubular disc sometimes hairy outside and the styles strikingly exserted. The whole plant blackens conspicuously during drying.

2. The Kew and Leiden type specimens of B. pustulata (Maingay KD 1371) bear the annotation "Griffith (Kew Distrib. 4883)." The specimens labelled solely "Griffith (Kew Distrib. 4883)" in K and U, however, are lectotype sheets of B. griffithii and represent B. cinnamomea. The material distributed as Griffith 4883 is obviously heterogeneous!

3. This species seems to occur infrequently. It closely resembles B. cinnamomea and B. insulana.

 

15. Bridelia whitmorei Airy Shaw

 

    Bridelia whitmorei Airy Shaw, Kew Bull. 27 (1972) 77; Whitmore, Tree Fl. Malaya 2 (1973) 74, in clavi; S.Dressler, Blumea 41 (1996) 322, Map 7. — Type: KEP FRI Whitmore 8586 (K holo; KEP, L iso), Malaya, Pahang, S Tembeling, near K. Keniyum, S. Redab.

 

Big woody climber; branchesconspicuously light brownish lenticellate, glabrous. Stipules not seen, early caducous. Leaves glabrous; petiole terete, 4–6 by c. 1 mm, glabrous, rarely slightly puberulous; blade (slightly ovate t0) elliptic to obovate, 65–140 by 30–65 mm, length/width ratio 1.7–2.4, (sub)coriaceous, base rotundate (to acute, obtuse, or slightly cordate), margin entire, apex obtuse (to shortly acuminate, acumen up to 1 cm long, often slightly curved), upper surface dark olive green when dry, lower surface glaucous; venation slightly prominent above, prominent beneath, secondary veins in 9–12 pairs, sinuously curved and branched towards the margin, joining the next secondaries and forming outer secondary veins, tertiary veins distantly percurrent, irregular and sinuous; intercostal areas somewhat bullate when dry. Inflorescences multibracteate glomerules of up to 8 sessile flowers, not terminal and not at special leafless branches; bracts narrowly triangular, 2–2.5 by up to 1.3 mm, brownish velutinous to tomentose. Staminate flowers not seen. Pistillate flowers 5–7 mm in diameter; pedicel at most 0.3 mm longl; sepals narrowly ovate-triangular, subulate, up to 3.5 by 1.5 mm, sparsely pilose at base; petals and disc not seen; ovary ellipsoid, pubescent, styles c. 2 mm long, united in the lower part, free and bilobed in the apical third, exserting, stigma lobate. Infructescences with up to 5 fruits, sometimes up to 1 mm stalked. Fruits ellipsoid to ovoid, up to 12 ny 8 mm in diameter, apex acute, base rotundate, yellow green, blackish when dry; mesocarp fleshy; endocarp woody. Seed one per fruit, compressed ellipsoid with lateral furrow, c. 8 by 6 by 4–5 mm; testa purple to dark brown, rugulate. – Note: The description is based on a single fruiting collection.

    Distribution — Malay Peninsula, so far only known from the type locality.

 

Bridadus-whit-map.gif (95017 bytes)    Distribution of B. whitmorei (squares; B. adusta = dots).

 

    Habitat & Ecology — Rocky streambed. Alt.: 240 m.

    Note — With the unilocular and one-seeded fruits as well as the lack of the fimbrial vein this taxon clearly belongs to subg. Gentilia. The general appearance (leaf shape, fruits) recalls B. penangiana, but the flowers are much larger and the leaves are stiff and coriaceus. With these features the taxon is quite close to B. balansae, but the leaves are rather different: they are neither lanceolate nor acute at the base, and they are of a conspicuous olive green colour when dry. An interesting feature is the somewhat bullate structure of the intercostal areas in sicco. The habit as a big woody climber seems to be unique among the taxa studied. Merely B. cinnamomea and B. stipularis grow as scrambling shrubs. The roughly lenticellate branches are reminiscent of B. pustulata but the leaves are not blackish when dry, they lack the closely scalariform higher venation, and the plants show only small few-flowered inflorescence cushions which are not strongly protruding.