Malesian Euphorbiaceae Descriptions

51. GALEARIA (Pandaceae)

P.C. van Welzen

Van Welzen, P.C. 2011. Pandaceae (formerly Euphorbiaceae s.l. subfam. Acalyphoideae tribe Galearieae). In: Nooteboom, H.N. (ed.), Flora Malesiana ser. 1, 20: 15–43.

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Genus description

Key to the species

Subgenus Galearia

Subgenus Orthopetalum

Species descriptions

Galearia Zoll. & Moritzi

    Galearia Zoll. & Moritzi in Moritzi, Syst. Verz. (1846) 19, nom. cons., non C.Presl, 1830, nom. rej.; Miq., Fl. Ind. Bat. 1, 2 (1859) 429; Fl. Ned. Ind., Eerste Bijv. (1860) 469; Kurz, Forest Fl. Burma 2 (1877) 407; Benth. in Benth. & Hook.f., Gen. Pl. 3 (1880) 287; Hook.f., Fl. Brit. India 5 (1887) 377; Pax in Engl. & Prantl, Planzenfam. 3, 5 (1890) 81; Pax & K.Hoffm. in Engl., Pflanzenfam. IV.147.iii (1911) 97; Ridl., Fl. Malay Penins. 3 (1924) 254; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1926) 456; Pax & K.Hoffm. in Engl. & Harms, Pflanzenfam. ed. 2, 19c (1931) 171; Forman, Kew Bull. 14 (1960) 311; Kew Bull. 20 (1966) 309; Backer & Bakh.f., Fl. Java 1 (1963) 495; Fl. Java 3 (1968) 649; Meijer, Bot. Bull. Herb. Forest Dept. Sabah 10 (1968) 234; Forman, Kew Bull. 26 (1971) 155; Airy Shaw, Kew Bull. 26 (1972) 362; Whitmore, Tree Fl. Malaya 2 (1973) 97; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 220; Kew Bull. Add. Ser. 8 (1980) 220; Kew Bull. 36 (1981) 365; Kew Bull. 37 (1982) 36; G.L.Webster, Ann. Missouri Bot. Garden 81 (1994) 68; Radcl.-Sm., Gen. Euphorbiacearum (2001) 127; Pooma in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 306; Welzen in Noot., Fl. Males. ser. 1, 20 (2011) 18; Kubitzki in Kubitzki, Fam. Gen. Vasc. Pl. 11 (2014) 271, Fig. 61b. — Galearia Subgen. Galearia: Forman, Kew Bull. 14 (1960) 316; Kew Bull. 20 (1966) 318. — Lectotype (designated by Forman, 1960): Galearia pedicellata Zoll. & Moritzi [= Galearia filiformis (Blume) Boerl. See note].

    Cremostachys Tul., Ann. Sci. Nat., sér. 3 Bot., 15 (1851) 259. — Lectotype (designated by G.L.Webster, 1994): Cremostachys filiformis (Blume) Tul. [= Galearia filiformis (Blume) Boerl.].

    Bennettia R.Br. in Benn. & R.Br., Pl. Jav. Rar. (1852) 249, non S.F. Gray, 1821); Baillon, Étude Gén. Euphorb. (1858) 311; Müll.Arg. in DC., Prodr. 15, 2 (1866) 1036. — Type: Bennettia javanica R.Br. [= Galearia filiformis (Blume) Boerl.].

Shrubs to small (to big in G. maingayi) trees, dioecious, bole sometimes fluted or with buttresses; buds in shoot axils. Indumentum of simple short and/or long hairs, often discolorous. Stipules triangular, often thick, hairy outside, glabrous inside (except G. aristifera), entire or with side-lobes in G. aristifera, caducaous to late caducous. Leaves simple, alternate, distichous; petiole short, round, flat or channelled above, subglabrous to hairy, slightly pulvinate; blades asymmetric, one side broader, pergamentaceous to coriaceous, drying green to brown, often somewhat bullate; base asymmetric, margin entire (to sometimes slightly crenate in G. celebica), flat to recurved, without glands, apex bluntly acuminate to caudate, upper surface smooth, glabrous to hairy on midrib, lower surface smooth, generally more hairy, venation pinnate, slightly raised or sunken above, raised beneath, nerves looped and closed towares margin, veins and veinlets coarsely reticulate. Inflorescences terminal (to cauliflorous in G. celebica), single or several together when seldom subterminal to many when cauliflorous, pendulous, spike-like thyrses, seldom paniculate (generally after damage), subglabrous to hairy, up to 1 m long, staminate flowers in groups, pistillate flowers single (or sometimes in threes in G. fulva); bracts triangular, completely hairy or glabrous inside; bracteoles similar to bracts but smaller. Flowers (4)5-merous; calyx 5-lobed, cupular, lobes basally attached, valvate, triangular, horizontally spreading, hairy outside, glabrous inside except for (partly) midrib in G. fulva, midrib inside thickened; petals (4)5, free, valvate, fleshy, two types, type 1 convex or cucullate with thickened midrib inside, glabrous or apex hairy, latter often hook-like, type 2 more or less straight, tomentellous on both sides, apex acute; disc absent. Staminate flowers pedicellate; pedicel hairy, round, basally with an abscission zone; petals late caducous; stamens (8 or) 10 (or 15) in a single or two whorls, often of different lengths, short ones epipetalous, long ones episepalous, filaments thick, free or adnate to each other and pistillode, glabrous or hairy, straight or S-shaped, anthers 2-thecate, thecae parallel to separated and almost horizontal, opening latrors with lengthwise split; pistillode columnar to obovoid, glabrous to hairy, sealing bud together with petals in subgenus Galearia (in subgenus Orthopetalum the petals close the bud). Pistillate flowers generally on a short pedicel, elongating in fruit, round, hairy, usually with an abscission zone beneath the flower; calyx persistent; petals early caducous, straight, slightly cucullate or cucullate, glabrous to hairy; ovary ovoid, 2–6-locular, hirsute; ovules single per locule, pendulous, anatropous; style absent, stigmas completely split once and in several species several times partly, in G. maingayi stigmas inside crater-like apex of fruit. Fruits latterally flattened to ovoid to subglobular drupes, often ridged or angular, 4- or 5-cornered, hairy, glabrescent, wall fleshy or bony, either thin and entire or thick with cavities, woody when dry; columella absent. Seeds flattened, triangular, ecarunculate.

    Distribution — An Asian genus of five species, ranging from Burma, Thailand and Vietnam throughout Malesia (unknown from the Lesser Sunda Islands and the Moluccas) to the Solomon Islands.

    Note — Webster (1994) proposed Galearia sessilis Zoll. & Moritzi as lectotype, becauce G. pedicellata (synonym of G. filiformis, lectotype by Forman, 1960) can be confused with Bennettia pedicellata R.Br., another synonym of G. filiformis. However, this is not a valid reason and thus the lectotype remains G. pedicellata (see alsoMcNeill et al., 2006, International Code of Botanical Nomenclature, Vienna Code).

Key to the Subgenera and Species

Subgenus Galearia

    Galearia Zoll. & Moritzi subgenus Galearia: Forman, Kew Bull. 14 (1960) 316; Kew Bull. 20 (1966) 318; Kew Bull. 26 (1971) 157; Welzen in Noot., Fl. Males. ser. 1, 20 (2011) 20. — Galearia Zoll. & Moritzi section Eugalearia Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iii (1911) 98, nom. inval. — Type as genus.

    Galearia Sect. Longibracteatae Pax & K.Hoffm.in Engl., Pflanzenr. IV.147.iii (1911) 102. — Lectotype (selected here): Galearia aristifera Miq.

Leaves: petioles flat above to round. Flowers: convex or cucullate, glabrous except often for a few apical hairs. Staminate flowers: filaments and connectives glabrous. Pistillate flowers: ovaries 2-locular. Fruits laterally flattened, wall thin, c. 0.4 mm thick, entire.

    Distribution — Three species in Southeast Asia main land and west Malesia.

1. Galearia aristifera Miq.

    Galearia aristifera Miq., Fl. Ned. Ind., Eerste Bijv. (1860) 471; Boerl., Handl. Fl. Ned. Ind. 3 (1900) 282; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iii (1911) 102, fig. 32; Merr., J. Straits Branch Roy. Asiat. Soc., Spec. No. (1921) 344; Pax & K.Hoffm., Mitt. Inst. Allg. Bot. Hamburg 7 (1931) 230; Forman, Kew Bull. 26 (1971) 159; Whitmore, Tree Fl. Malaya 2 (1973) 98; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 220; Kew Bull. 36 (1981) 366; Welzen in Noot., Fl. Males. ser. 1, 20 (2011) 20, fig.1a–g; 2b; map 1. — Bennettia aristifera (Miq.) Müll.Arg. in DC., Prod. 15, 2 (1866) 1039. — Type: Teijsmann HB 749 (holo U),W. Sumatra, near Lubu(k)alang.

    Galearia leptostachya Pax in Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iii (1911) 102; Merr., J. Straits Branch Roy. Asiat. Soc., Spec. No. (1921) 345 (‘leptostachys’). — Type: Haviland & Hose 979 (iso K, L), Sarawak, near Kuching, Garai.

Shrubs to small trees, up to 5(–12) m, dbh up to 30 cm; flowering branches 2–3.5 mm thick, densely hairy (to almost glabrous), glabrescent. Indumentum simple hairs, hirsute, up to 1 mm long, usually short (yellow) and long hairs (yellow or dark brown) present. Outer bark smooth, chocolate brown and grey mottled. Stipules triangular, 6–15 by 0.5–1 mm, hairy, about halfway usually triangular side lobes. Leaves: petiole 4–7 mm long, round, hairy; blades elliptic (to somewhat obovate), 6.7–26.5 by 4.1–11.1 cm, length/width ratio 1.8–3.6, leathery, , often somewhat bullate, drying light green to somewhat brownish light green, base emarginate to rounded, margin entire, flat, apex acuminate to cuspidate (to caudate), sometimes mucronulate, upper surface generally hairy on midrib and nerves, lower surface hairy allover with short and long hairs, venation slightly sunken above, nerves 9–11 per side. Inflorescences terminal, up to 100 cm long, pale green to greenish brown, hairy, seldom subglabrous, pistillate flowers single; bracts (1.8–)4.5–31 by 0.4–1 mm, hairy, seldom glabrous. Flowers 5-merous; calyx with 5 lobes, up to 0.7 mm deep, lobes triangular, 0.4–0.6 by 0.3–0.5 mm, inside glabrous; petals 5, cucullate, apex rounded, bent inwards, midrib very thick. Staminate flowers 3.5–5.3 mm in diam., (pale) green; pedicel 3.5–5.3 mm long; petals elliptic, c. 1.3 by 1 mm, few hairs basally; stamens 10 in two whorls, one shorter, other longer, filaments glabrous, c. 0.3 or c. 0.6 mm long, anthers 0.25–0.4 by c. 0.25 mm, thecae parallel, basifixed; pistillode 0.7–0.8 mm high, long hairy, especially on the top. Pistillate flowers c. 1.4 mm in diam.; pedicel c. 0.5 mm high, elongating in fruit up to 2.5 mm; petals 5, elliptic, caducous, 2–3.5 by 0.7–1 mm; ovary subglobular, c. 1.3 by 1.3 mm, 2-locular, hirsute; ovules single per locule, pendulous, anatropous; style c. 0.5 mm high, stigmas 0.6–1.4 mm long, deeply bifid, with long papillae above. Fruits flattened ovoid drupes, c. 11 by 8.5 by 6 mm, densely hirsute, wall thin, woody. Seeds not seen.

    Distribution — Peninsular Malaysia, Sumatra, Borneo.

    Habitat & Ecology — (Edge of) primary forest, secondary forest, kerangas forest to common along logging roads in open places; soil: deep clay to dry sandy soil, level to steep terrain. Altitude: 20–770 m. Flowering: February to December; fruiting: June to September.

    Vernacular names — Sumatra: Ekor ekor, Kajoe ekor badak, Kajoe ihoer badak, Kajoe ihoe here, Madang tjoemara djatie. Kalimantan: Perikas (Dayak), Tankai ajok.

    Note — Very typical for this species are the long bracts in the inflorescences (4 mm and longer). However, on the Malay Peninsula these can be much shorter, c. 2 mm long, and then the species resembles G. fulva. Differences can still be found in the heavier indumentum of G. aristifera (generally far less dense in G. fulva but hairy exceptions occur), the pistillode (long hairs in G. aristifera and generally – not always - short ones in G. fulva) and the stipules ((1.8–)4.5–31 mm long and about halfway usually with triangular side lobes in G. aristifera, 1–3 mm long and without side lobes in G. fulva).

2. Galearia filiformis (Blume) Boerl.

    Galearia filiformis (Blume) Boerl., Handl. Fl. Ned. Ind. 3 (1900) 282 (see note); J.J.Sm., Meded. Dept. Landb. Ned.-Indië 10 (1910) 550 (= Koord. & Val., Booms. Java 12); Pax & K.Hofm. in Engl., Pflanzenr. IV.147.iii (1911) 98, fig. 34; Backer & Bakh.f., Fl. Java 1 (1963) 495; Forman, Kew Bull. 26 (1971) 160, fig. 1d, 2c; Airy Shaw, Kew Bull. 36 (1981) 366; Welzen in Noot., Fl. Males. ser. 1, 20 (2011) 23; fig. 1h–n; map 2. — Antidesma filiformis Blume, Bijdr. 17 (1827) 1124. — Cremostachys filiformis (Blume) Tul., Ann. Sci. Nat., sér. 3 Bot., 15 (1851) 260. — Bennettia filiformis (Blume) Müll.Arg., Linnaea 34 (1865) 204; in DC., Prodr. 15, 2 (1866) 1038. — Lectotype (L sheets selected here, K sheet by Forman, 1971): Blume s.n. (holo L (barcode L  0157229); iso K (barcode K000253975), L (barcode L  015720)), Java, Mt. Salak). See note 2.

    Galearia pedicellata Zoll. & Moritzi in Moritzi, Syst. Verz. (1846) 19. — Type: Zollinger 1410 (iso, perhaps holo P, separate inflorescence removed = Ostodes paniculata Blume), Java, between Sudimanik and Tjiburial.

    Galearia sessilis Zoll. & Moritzi in Moritzi, Syts. Verz. (1846) 19; Miq., Fl. Ind. Bat. 1(2) (1859) 429. — Type: Zollinger 567 (iso, perhaps holo P), Java, Tjidurian R.

    Bennettia javanica R.Br., Pl. Jav. Rar. (1852) 249, t. 50. — Type: Horsfield s.n. (holo BM), Java.

    [Ryparia (Ryparosa) dubia Blume ex Baill., Étude Euphorb. (1858) 339, nom. nud. — Representative specimen: Blume s.n. (L, barcodes L  0157215, L  0157215).]

Shrubs to small trees up to 10 m high, bole up to 3 m high, dbh up to 15 cm; flowering branches 1–3 mm thick, sericeous, glabrescent. Indumentum simple hairs, short (< 0.5 mm long), mainly sericeous, dark brown and yellow or yellow only. Outer bark smooth to fissured, very hard. Stipules triangular, 0.8–2 by 0.3–0.6 mm, outside hairy, inside glabrous, usually late caducous. Leaves: petiole 2–3.5 mm long, flattened above, hairy; blades ovate to elliptic, 3–25 by 1.7–8 cm, length/width ratio 2.2–3.9, pergamentaceous to coriaceous, brownish green when young, generally drying light green, base shortly attenuate to cuneat, margin entire, flat, apex long acuminate, often mucronulate, upper surface glabrous except for some hairs on basal part midrib (and nerves), lower surface somewhat hairy, mainly on venation, venation generally slightly raised above but midrib flat, raised beneath, nerves 5–7 per side. Inflorescences terminal (or subterminal when several, probably due to sympodial length growth), up to 52 cm long, green, hairy, pistillate flowers single; bracts 0.8–1.5 by 0.25–0.5 mm, glabrous inside. Buds green. Flowers 5-merous, green to white; calyx with 5 lobes, 0.8–1.3 mm deep, lobes triangular, 0.4–0.6 by 0.4–0.7 mm, glabrous inside; petals 5, elliptic, cucullate, as deep as high, 1–1.2 by 0.8–1.3 mm, 1–1.5 mm deep, glabrous, midrib strongly thickened inside, often furrowed outside. Staminate flowers 3.6–4 mm in diam.; pedicel 6.3–9 mm long; stamens 10 in two whorls, of different lenghts, filaments glabrous, c. 0.5 or 0.5–0.7 mm long, often band-like in lateral view, often S-shaped; anthers 0.4–0.5 by 0.4–0.5 mm, thecae separated, ± parallel, horizontal because of S-shape filaments, cream to light yellow; pistillode 0.7–0.8 by 0.5–1 mm in diameter, glabrous. Pistillate flowers 2.5–3 mm in diam.; pedicel c. 0.4 mm long, elongating in fruit up to 2.5(–4) mm; ovary c. 1.2 by 1.3 mm, very shallowly 5-lobed with central furrow due to imprints of petals, 2-locular, stigmas c. 0.5 mm long, completely split several times. Fruits laterally flattened ovoid drupes, often with 2 ridges along edges, 9–9.5 long by 6.2–7.2 high by 5.5–7.1 mm thick, somewhat hairy, wall thin, woody, c. 0.4 mm thick, very light green to pale whitish green when immature. Seeds seen immature, c. 8.5 mm long by 5.5 mm high by 1 mm thick.

    Distribution — Sumatra, Java.

    Habitat & Ecology — Primary rainforest rich in palms, secondary forest (with much bamboo), riparian forest, secondary montane forest, along sea coast; soil: on rocks, coral, ferrosal loam. Altitude: sea level up to 600(–1500) m. Flowering: March to May, July to September, November; fruiting: May, August, November.

    Vernacular names — Java: Balung kaja (Blume, 1827); Ki keretjek (Sundanese, Djasilin dialect).

    Notes — 1. This species is easy to separate from G. fulva when flowering and the petals are still present (usually early caducous in pistillate flowers): G. filiformis has glabrous petals as deep as high and a glabrous pistillode, while G. fulva has petals with apical hairs, higher than deep and a hairy pistillode. When fruits or petalless pistillate flowers are present then the calyx in G. filiformis is glabrous inside, hairy in G. fulva. Other, overlapping differences are the generally light green colour of dried leaves of G. filiformis versus light green to brownish in G. fulva and the pedicels of the fruits, up to 4 mm in G. filiformis and up to 12 mmin G. fulva. Geographically the species only overlap on Sumatra.

2. The combination Galearia filiformis was first published by Pax (1890) but with as authors (Blume) Benth. References to the basionym and the other combinations are missing, thus this new combination was not published validly. The first to make a valid new combination is Boerlage (1900).

3. Galearia fulva (Tul.) Miq.

    Galearia fulva (Tul.) Miq., Fl. Ind. Bat. 1, 2 (1850) 430; Hook.f., Fl. Brit. India 5 (1887) 378; Boerl., Handl. Fl. Ned. Ind. 3 (1900) 282; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iii (1911) 101; Ridl., Fl. Malay Pen. 3 (1924) 257; Forman, Kew Bull. 26 (1971) 160, fig. 1c, 2d; Airy Shaw, Kew Bull. 26 (1972) 362; Whitmore, Tree Fl. Malaya 2 (1973) 98; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 221; Kew Bull. 36 (1981) 366; Pooma in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 306, fig. 1, plate XVII: 1.1—1.3; Welzen in Noot., Fl. Males. ser. 1, 20 (2011) 23; fig. 1o–u; 2a; 3a–d; 4–6; map 3. — Cremostachys fulva Tul., Ann. Sci. Nat., sér. 3, 15 (1851) 261. — Bennettia fulva (Tul.) Müll.Arg., Linnaea 34 (1865) 205; in DC., Prodr. 15, 2 (1866) 1037. — Type: Wallich 8585C parte ex Herb. Lindley (holo CGE; iso L), Penang.

    Cremostachys lindlaeana Tul., Ann. Sci. Nat., sér. 3, 15 (1851) 262. — Bennettia pedicellata R.Br., Pl. Jav. Rar. (1852) 251, nom. inval., based on same type; Müll.Arg. in DC., Prodr. 15, 2 (1866) 1039. — Galearia pedicellata (R.Br.) Miq., Fl. Ind. Bat. 1, 2 (1859) 430, non Zoll. & Moritzi (1846); Hook.f., Fl. Brit. India 5 (1887) 378; Boerl., Handl. Fl. Ned. Ind. 3 (1900) 282; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iii (1911) 101; Ridl., Fl. Malay Penins. 3 (1924) 257; Gagnep. in Lecomte, Fl., Indo-Chine 5 (1926) 457, nom. inval. — Bennettia lindlaeana (‘lindleyana’) (Tul.) Müll.Arg., Linnaea 34 (1865) 205; in DC., Prodr. 15, 2 (1866) 1037. — Galearia lindlaeana (‘lindleyana’) (Tul.) Hook.f., Fl. Brit. India 5 (1887) 379; Boerl., Handl. Fl. Ned. Ind. 3 (1900) 282; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iii (1911) 98; Ridl., Fl. Malay Penins. 3 (1924) 255. — Galearia caudata Forman, Kew Bull. 14 (1960) 316, nom. inval. — Type: Herb. Wallich 8484C parte ex Herb. Lindley (holo CGE; iso K), Penang.

    Bennettia affinis R.Br., Pl. Jav. Rar. (1852) 251; Müll.Arg. in DC., Prodr. 15, 2 (1866) 1037. — Galearia affinis (R.Br.) Miq., Fl. Ind. Bat. 1, 2 (1859) 430; Hook.f., Fl. Brit. India 5 (1887) 379; Boerl., Handl. Fl. Ned. Ind. 3 (1900) 282; Pax & Hoffm. in Engl., Pflanzenr. IV.147.iii (1911) 99; Ridl., Fl. Malay Penins. 3 (1924) 255; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1926) 458. — Type: Finlayson in Herb. Wallich 8585B pro parte, Herb. Finlay (holo BM), Penang.

    Bennettia finlaysonii R.Br., Pl. Jav. Rar. (1852) 251; Müll.Arg. in DC., Prodr. 15, 2 (1866) 1039; Hook.f., Fl. Brit. India 5 (1887) 380. — Galearia finlaysonii (R.Br.) Miq., Fl. Ind. Bat. 1, 2 (1859) 429; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iii (1911) 100; Ridl., Fl. Malay Penins. 3 (1924) 257. — Type: Finlayson in Herb. Wallich 8585 (no. 28) (holo BM; iso BM), Penang.

    Bennettia jackiana R.Br., Pl. Jav. Rar. (1852) 251; Müll.Arg. in DC., Prodr. 15, 2 (1866) 1039; Boerl., Handl. Fl. Ned. Ind. 3 (1900) 282. — Galearia jackiana (R.Br.) Miq., Fl. Ind. Bat. 1, 2 (1859) 430; Hook.f., Fl. Brit. India 5 (1887) 378; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iii (1911) 101. — Type: Jack in Herb. Wallich 8585A (holo BM), Penang.

    Bennettia phlebocarpa R.Br., Pl. Jav. Rar. (1852) 251; Müll.Arg. in DC., Prodr. 15, 2 (1866) 1038. — Galearia phlebocarpa (R.Br.) Miq., Fl. Ind. Bat. 1, 2 (1859) 429; Hook.f., Fl. Brit. India 5 (1887) 380; Boerl., Handl. Fl. Ned. Ind. 3 (1900) 282; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iii (1911) 100; Ridl., Fl. Malay Penins. 3 (1924) 255; Merr., J. Straits Branch Roy. Asiat. Soc., Spec. No. (1921) 345; Pax & K.Hoffm., Mitt. Inst. Allg. Bot. Hamburg 7 (1931) 229; Meijer, Bot. Bull. Herb. Forest Dept. Sabah 7 (1967) 45. — Type: Herb. Wallich 8585D (holo BM; iso K, 2 sheets), Singapore.

    Bennettia wallichii R.Br., Pl. Jav. Rar. (1852) 251; Müll.Arg. in DC., Prodr. 15, 2 (1866) 1038. — Galearia wallichii (R.Br.) Kurz, Prelim. Rep. Forest Pegu, App. A (1875) cxiii; Forest Fl. Burma 2 (1877) 407; Hook.f., Fl. Brit. India 5 (1887) 379; Boerl., Handl. Fl. Ned. Ind. 3 (1900) 282; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iii (1911) 100; Ridl., Fl. Malay Penins. 3 (1924) 256.— Type: Herb. Wallich 8585E (holo BM; iso K), Burma, Tavoy.

    Galearia angustifolia Miq., Fl. Ned. Ind., Eerste Bijv. (1860) 470. — Type: Teijsmann HB 3925 (holo U, iso K as s.n.), Sumatra, Palembang, near Muara-enim.

    Galearia elliptica Miq., Fl. Ned. Ind., Eerste Bijv. (1860) 469. —Syntypes: Teijsmann H.B. 551 (K as s.n., U), Sumatra, Poeloe Pisang prope Padang; Teijsmann HB 3825 (U, 2 sheets), Sumatra, Palembang, Muara-enim.

    Galearia splendens Miq., Fl. Ned. Ind., Eerste Bijv. (1860) 469; Boerl., Handl. Fl. Ned. Ind. 3 (1900) 282; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iii (1911) 101 — Bennettia splendens (Miq.) Müll.Arg., Linnaea 34 (1865) 205; in DC., Prodr. 15, 2 (1866) 1038. — Type: Teijsmann HB 550 (holo U; iso K, s.n.), Sumatra, Poeloe Pisang, prope Padang.

    Galearia sumatrana Miq., Fl. Ned. Ind., Eerste Bijv. (1860) 469. — Type: Teijsmann 4307 (holo U; iso K, s.n.), Sumatra, prov. Lampong, near Kebang.

    Bennettia subulata Müll.Arg. in DC., Prodr. 15, 2 (1866) 1039. — Galearia subulata (Müll.Arg.) Hook.f., Fl. Brit. India 5 (1887) 379; Boerl., Handl. Fl. Ned. Ind. 3 (1900) 282; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iii (1911) 102; Ridl., Fl. Malay Penins. 3 (1924) 258. — Type: ’Ex Herb. Lindley in Herb. Sonder’(probably Herb. Wallich 8585C p.p., the type of Cremostachys fulva) (holo MEL), Penang (Prince of Wales Island).

    Galearia helferi Hook.f., Fl. Brit. India 5 (1887) 378; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iii (1911) 101. — Syntypes: Helfer KD 4968 (K, NY), Burma, Tenasserim; Kurz s.n. (n.v.), Tenasserim.

    Galearia filiformis auct. non (Blume) Boerl.: Merr., Philipp. J. Sci. Bot. 4 (1909) 280.

    Galearia philippinensis Merr., Philipp. J. Sci., Bot. 9 (1914) 482; Enum. Philipp. Fl. Pl. 2 (1923) 450. — Type: BS (Reillo) 16319 (holo PNH, †; iso K, L), Philippines, Basilan.

    Galearia minor Gage, Rec. Bot. Surv. India 9 (1922) 234; Ridl., Fl. Malay Penins. 3 (1924) 256. — Lectotype (selected here): Ridley s.n. (holo SING), Pahang, Pulau Jellum. See note 1.

    Galearia ridleyi Gage, Rec. Bot. Surv. India 9 (1922) 235; Ridl., Fl. Malay Penins. 3 (1924) 256. — Syntypes: Lake & Kelsall s.n. (SING, barcode 0097602), Johore, Gunung Janeng; Lake & Kelsall s.n. (SING, barcode 0097601), Johore, Simpai; Ridley s.n. (SING, barcode 0097603), Malay, Selangor, Dusun Tua.

    Galearia sessiliflora Merr., J. Straits Branch Roy. Asiat. Soc. 86 (1922) 320; Pl. Elmer. Born. (1929) 161. — M. Ramos 1312 (iso US), Sabah, Batu Lima near Sandakan.

    Galearia stenophylla Merr., J. Straits Branch Roy. Asiat. Soc. 86 (1922) 320; Meijer, Bot. Bull. Herb. Forest Dept. Sabah 7 (1967) 45; Forman, Kew Bull. 26 (1971) 159; Whitmore, Tree Fl. Malaya 2 (1973) 98; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 221. — Type: Ramos 1542 (holo PNH, †; iso K, L), Sabah, Batu Lima near Sandakan.

    Galearia dongnaiensis Pierre ex Gagnep., Bull. Soc. Bot. France 71 (1925) 1025; in Lecomte, Fl. Indo-Chine 5 (1926) 457, fig. 56: 1—6. — Type: Pierre 1969 (holo P, iso K, P, 2 sheets), Vietnam, Prov. Bien-hoa, Mt. Pong-lu.

    Galearia dolichobotrys Merr., Philipp. J. Sci. 29 (1926) 384; Meijer, Bot. Bull. Herb. Forest Dept. Sabah 7 (1967) 45. — Type: Castro & Melegrito 1377 (holo PNH, †; iso A, K), Sabah, Banguey (Banggi) Island.

    Galearia lancifolia Ridl., Bull. Misc. Inform. Kew (1926) 476. — Type: Hume 9931 (holo K), Malaya, Selangor, Ulu Gombok.

Shrubs to small trees, up to 20 m, bole up to 11 m high, dbh up to 25 cm; flowering branches 1–2 mm thick, glabrous to sericeous or hirsute, glabrescent. Indumentum simple hairs, short (< 0.5 mm long) to long (up to 1 mm long), sparsely to densely hirsute or sericeous, dark brown and yellow or yellow only. Outer bark smooth to slightly roughened (to slightly fissured to scaly), green to light brown to reddish brown to white to light to dark grey, often patches of different colours; inner bark pale greenish to whitish to pale yellow to pink to red to brown, soft, granular, c. 3 mm thick; sapwood white to yellowish to pale orange to red, hard. Stipules triangular, 1–3 by 0.3–0.5 mm, hairy. Leaves: petiole 1.3–8 mm long, flattened above, glabrous to hairy; blades ovate to elliptic to obovate, 4.1–32 by 1.2–13.8 cm, length/width ratio 1.9–14.7, pergamentaceous, flat to bullate, drying light green to somewhat brownish light green, base emarginate to attenuate, margin entire, flat to recurved, apex long acute to cuspidate, often mucronulate, upper surface glabrous to hairy on midrib and nerves, dark green, lower surface glabrous to hairy allover, pale glaucous to lighter green, venation slightly sunken to slightly raised above, nerves 7–11(–22) per side. Inflorescences terminal, up to 110 cm long racemes (to seldom paniculate), subglabrous to hairy, dark green to light brown, pistillate flowers single or three per node; bracts 0.7–2.7 by 0.25–0.5 mm, glabrous inside. Flowers (4)5-merous, green (to white); buds yellow-green; calyx with (4)5 lobes, 0.8–1.3 mm deep, lobes triangular to ovate, 0.5–1 by 0.4–0.6 mm, midrib inside generally (partly) hairy; petals (4)5. Staminate flowers 2–3.2 mm in diam.; pedicel 2.5–10 mm long; petals elliptic, cucullate with apex at right angles with rest, 1–1.8 by 0.7–1 mm, higher than deep, apex with hairs on abaxial surface; stamens (8)10 in two whorls, of different lengths, filaments glabrous, 0.3–0.5 or 0.5–0.75 mm long, anthers c. 0.5 by 0.3 mm, thecae parallel; pistillode 0.5–0.7 by 0.3–0.5 mm in diameter, hairy with short to long hairs. Pistillate flowers 1.6–4 mm in diam.; pedicel 0–3.5 mm long, elongating in fruit up to 12 mm; petals 5, elliptic, usually hairs on the apex, 2 types, one slightly cucullate, the other type cucullate with a long, inward bent hook-like apex and a stipe-like base (up to c. 0.7 mm long), 1.7–2 by c. 1 mm; ovary 0.8–1.2 by 0.8–1.2 mm, 2-locular, without petaloid imprints; stigmas 0.5–0.7 mm long, deeply bifid, lobes also split, with long papillae above. Fruits laterally flattened ovoid drupes with ridges along edges, 10.5–12 long by 7.8–8.5 high by 6–6.3 mm thick, somewhat hairy, glabrescent, yellowish to pale green to white, wall thin, woody, c. 0.4 mm thick. Seeds seen immature.

    Distribution — Burma, Thailand, Vietnam, Malay Peninsula, Sumatra, Borneo, Philippines.

    Habitat & Ecology — Primary rain forest, evergreen forest, kerangas forest, fresh water swamp forest, secondary forest, logged forest, open areas, ladangs; along streams, trails, roads, and on ridges; soil: Variable, on alluvial, clay, limestone, loam, sand, sandy clay, shale, ultrabasic (serpentinite); bedrock basalt, granitic, granodiorite, sandstone. Altitude: 8–1300 m. Flowering and fruiting whole year through. Animals eat the fruits.

    Vernacular names — Thailand: Ai-bao. Malaysia: Kelandes (Batek); Pnoron tbas (Semelai). Sumatra: Akar kuning; Kajoe ihoer badak. Borneo: Kayu busih; Kayu warna; Malatidong (Kedayan); Rukam gunung; Tis, Titidong (Dusun).

    Uses — The stem is used to make spring and spear traps (Malay Peninsula, Semelai); the leaves are used as vegetable Dusun (Borneo) or in the sayur (Kalimantan Barat). In Thailand the root is used locally for fever.

    Notes — 1. A lectotype is selected for G. minor, because one of the syntypes, H.O. Forbes 3096 (L) is G. filiformis and not G. fulva.

2. This species complex is very variable. Galearia aristifera almost merges into this species (see note under latter) and in this treatment G. stenophylla is not recognised anymore. In the following paragraphs several series are mentioned showing that all differences in shapes and sizes merge into each other. The only difference that still exist is the presence of two petal types in the pistillate flowers. One type is generally found in G. fulva (rather straight, slightly cucullate petals), while the other type (cucullate with hook-like apex) is mainly present in the G. stenophylla form (however, no differences in fruits and staminate flowers with more typical G. fulva, and the leaf shapes gradually merge with G. fulva).

Series from G. stenophylla with long and narrow leaves (rheophytic type) to G. fulva with shorter, broader leaves, also a series from glabrous to hairy. All ovate: S (Yii) 40686 (glabrous) and M. Ramos 1542 (type G. stenophylla; hairy), both ovate, long and slender (19 x 1.7 cm), to SAN (Bidin) 84815 (shorter, ovate, hairy; 12.3 x 11.8), to S (Paie) 26018 (shorter, relatively broader, 11.2 x 2.6 cm) to very small (SAN (Joseph B et al.) 113516, 4.7 x 1.3 cm). The apex in this series changes from gradually decurrent into an acute apex (narrow leaves) to acuminate/cuspidate (and occasionally caudate) (broader leaves).

From small to larger and bullate leaves (all ovate): S (Paie) 26018 (shorter, relatively broader, 11.2 x 2.6 cm) to broader and larger, and from a flat surface to more bullate: S (Sibat ak Luang) 24625 (12.3 x 3.1 cm; flat) to Castro & Melegrito 1377 (type G. dolichobotrys; 16.5 x 5.3 cm, flat) to Shea 27028 (20.5 x 7.3 cm, bullate).

From ovate to elliptic/oblong (long and slender): M. Ramos 1542 (type G. stenophylla) to Veldkamp 8461 to SAN (Amin et al.) 116397 to SAN (Krispinus) 130005.

From small leaves to large ones: SAN (Joseph B et al.) 113516 (4.7 x 1.3 cm) to S (Paie) 26018 (11.2 x 2.6 cm) to Afriastini 86 (16.3 x 6.3 cm) to R.J. Johns et al. 7564 (28 x 8.4 cm) to Korthals s.n., L  0157356 (29 x 14 cm), SAN (Campbell et al.) 11873 (31 x 11 cm).

From ovate to obovate: S (Paie) 26018 (ovate) to Slik LUPRI-7617 (elliptic) to Suzuki K 1222 (obovate, small leaves) and Tuke P2 1013 (obovate, large leaves).

From green drying leaves to brown drying leaves: SAN (Ampuria) 34161 (light green) to Kessler et al. PK 1665 (greenish brown) to Endert 3207 (brown).

From an attenuate leaf base to an emarginate base: Suzuki K 1222 (attanuate) to San (Dewol et al.) 109582 (rounded) to Kostermans 10003 (emarginate). Puasa 4647 has rounded and attenuate leaves.

Subgenus Orthopetalum

    Galearia Subgen. Orthopetalum Forman, Kew Bull. 14 (1960) 316; Kew Bull. 20 (1966) 318; Welzen in Noot., Fl. Males. ser. 1, 20 (2011) 33. — Type: Galearia celebica Koord.

Leaves: petioles channelled above. Flowers: petals ± straight and flat, completely hairy. Staminate flowers: filaments and connectives hairy. Pistillate flowers: ovaries 2–6-locular. Fruits laterally flattened to 5-cornered to subglobular, wall 6–17 mm thick, with cavities.

    Distribution — Two species, one in west Malesia (Malay Peninsula, Sumatra, Borneo), the other in east Malesia (Sulawesi, New Guinea) and the Solomon Islands.

4. Galearia celebica Koord.

    Galearia celebica Koord., Meded. Lands Plantentuin 19 (1898) 626; Boerl., Handl. Fl. Ned. Ind. 3 (1900) 282; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iii (1911) 103; Forman, Kew Bull. 14 (1960) 313; Kew Bull. 20 (1966), 313, 318, 319, t5:3, 4; Kew Bull. 26 (1971) 157, fig. 1b, 2b1-2; Airy Shaw, Kew Bull. Add. Ser. 8 (1980) 220; Kew Bull. 37 (1982) 36; Welzen in Noot., Fl. Males. ser. 1, 20 (2011) 34; fig. 3e–m; 7a–d; map 4. — Galearia celebica Koord. var. celebica: Forman, Kew Bull. 14 (1960) 315; Kew Bull. 26 (1971) 158; Airy Shaw, Kew Bull. Add. Ser. 8 (1980) 220; Kew Bull. 37 (1982) 36. — Lectotype (Forman, 1960): Koorders 16975 (L), Celebes, Prov. Minahassa.

    Galearia fusca Ridl., Fl. Malay Penins. 3 (1924) 257. — Type: Burkill 6375 (holo K), Malaya, Negri Sembilan, Gemas.

    Galearia celebica Koord. var. pubescens Forman, Kew Bull. 14 (1960) 315; Kew Bull. 26 (1971) 158; Airy Shaw, Kew Bull. Add. Ser. 8 (1980) 220. — Type: NGF (A.G. Floyd) 7018 (holo K; iso L), Papua New Guinea, New Britain, Keravat, Keravat River.

(Shrubs to ) small trees up to 28 m high, bole up to 10 m high; dbh up to 40 cm; buttresses seldom present (Solomon Islands sometimes), thin and plank-like, steep to up 1.5 m high, (seldom fluted to first branch); flowering branches 2–40 mm thick, longitudinally grooved when dry, only young parts hairy (to completely). Indumentum simple hairs, either ver short, whitish hairs (less than 0.5 mm long) and sometimes also long, yellowish, hirsute hairs (up to 1 mm long), see note. Outer bark light brown to greenish grey to grey, smooth (to fissured to scaly), c. 0.2 mm thick; under bark pale greenish cream to yellow to reddish to brown; inner bark pink to pinkish grey to dark red to brownish outside, coarsely fibrous to non fibrous, soft, 3–6 mm thick; wood pale cream to straw to orange to orange-brown, firm, clean cutting, finely radially striate. Stipules triangular, 1.5–2 by 0.5–1 mm, outside subsericeous, inside glabrous, caducous. Leaves: petiole 3–8 mm long, channelled above, subglabrous to hirsute; blades (ovate to) elliptic, 7–32 by 3.3–12 cm, length/width ratio 2.1–3.6, papery to chartaceous, drying light green, base broadly attenuate, margin entire (to slightly crenate), apex usually damaged, acuminate, sometims mucronulate, upper surface shiny (to dull) dark green above, glabrous to hairy on basal part midrib, lower surface (sub)glabrous to softly hirsute, dull, pale to mid green, venation slightly raised above, nerves 6–10 per side. Inflorescences cauliflorous and up to 30 together, or terminal and single, up to 75 cm long, damaged cauliflorous ones branching, side branches racemes up to 100 cm long, subglabrous to hirsute, pistillate flowers single per node; bracts up to 0.5 by 0.5 mm, inside glabrous. Flowers 5-merous; calyx with 5 lobes, 1.5–3 mm deep, glabrous inside, green, lobes shallow, 0.4–1 by 2–2.7 mm, apex rounded; petals 5, ovate, 3.5–7 by 2–3.8 mm, concave, fleshy, c. 0.7 mm thick, apex acute, puberulous on both sides, white to cream to pale pink. Staminate flowers 5–15 mm in diam., (faintly) scented; pedicel 5.7–20 mm long; stamens 10 (or 15), in a single whorl, adnate to each other and to pistillode, filaments very broad, strap-like, with thickened midrib, 1.5–2.3 mm high, hairy, white to pale pink, connective triangular, hairy, seperating thecae, anthers c. 0.7–0.8 by 0.6–0.7 mm, connective without appendix; pistillode 2.8–3.7 by 2.5–3 mm, puberulous, very pale green to light yellow. Pistillate flowers 3.7–4 mm in diam., pedicel 3.7–10.5 mm long; ovary 2–4(–6)-locular, ovoid to obovoid, 2–3.2 by 1.8–2 mm, stigmas on top of ovary, 0.6–1 mm long, with a few papillae above and apical, completely split. Fruits laterally flattened to triangular to square to roundish in transverse section, broadest basally or apically, usually angular, 16–30 long by 11–30 wide by 9–18 mm high, subglabrous, whitish; wall very thick, 6–8 mm. Seeds c. 15 by 10 by 2.5 mm.

    Distribution — Sulawesi, New Guinea, Solomon Islands (unknown for Moluccas).

    squares = Galearia celebica Koord.; dots = Galearia maingayi Hook.f.

    Habitat & Ecology — (Disturbed) primary forest, alluvial forest, beach forest, Castanopsis forest, swamp forest, regrowth; river sides, along path, behind seashore; soil often periodically flooded, sand, sandy clay, reddish clay, coral, ultrabasics. Altitude: sea level up to 1100 m. Flowering and fruiting: whole year through.

    Vernacular names — Irian Jaya: Ammiorapi (Roberbai, Japen Dialect); Barboeroeroego; Kembien (Tehid); Wok (Mooi). Papua New Guinea: Manimk (Saki); Mapiok (Waskuk); Pahapi (Wagu); Papui (Dumpu); Ugan (Amele); Uriri (Faita). Solomon Islands: Bainonoo, O’oa (Kwara’ae).

    Note — Forman (1960) recognized two varieties, base on the absence or presence of a dense indumentum of long, hirsute hairs. The two varieties are synonymised, because there is a complete overlap in distribution and the density of the long hairs is rather variable, varying from subglabrous to absent in the inflorescences to present on leaves and inflorescences.

5. Galearia maingayi Hook.f.

    Galearia maingayi Hook.f., Fl. Brit. India 5 (1887) 377; Boerl., Handl. Fl. Ned. Ind. 3 (1900) 281; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iii (1911) 103; Ridl., Fl. Malay Penins. 3 (1924) 255; Forman, Kew Bull. 14 (1960) 311; Kew Bull. 20 (1966) 312, fig. 1.2; Kew Bull. 26 (1971) 158, fig. 1a, 2a1-2; Meijer, Bot. Bull. Herb. Forest Dept. Sabah 10 (1968) 233; Whitmore, Tree Fl. Malaya 2 (1973) 98; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 221; Kew Bull. 36 (1981) 367; Welzen in Noot., Fl. Males. ser. 1, 20 (2011) 37; fig. 3n–p; 7e–k; map 4. — Syntypes: Maingay 2733 (KD 1412) (K, 2 sheets), Malacca; Maingay 3129 (KD 1412/2) (K), Malacca.

(Big) trees, up to 35 m high, bole up to 20 m high, dbh up to 80 cm; sometimes fluted or buttresses present, up to 3 m tall; flowering branchlets 2.5–3 mm thick, sericeous-tomentose, white to whitish green. Indumentum simple short (less than 0.5 mm long), mainly sericeous hairs, generally most parts rather glabrous except young branchlets. Outer bark white to grey to yellowish grey to yellowish to yellowish brown to light brown to greenish or dark brown, smooth to fissured to slightly vertically ridged-corky, not peeling to peeling, yellow and grey-brown on back, 0.2–1 mm thick; under bark pinkish straw; inner bark pinkish straw to light red to dark red to red-brown, 6–15 mm thick, fibrous, rather brittle to very hard; wood yellowish to orange brown, hard. Stipules triangular, 0.8–1 by 0.6–0.7 mm, thick, outside subsericeous, inside glabrous, very early caducous. Leaves: petiole 5.5–13 mm long, channelled above, (sub)glabrous; blades ovate to elliptic, 5.2–20.5 by 2.1–6.7 cm, length/width ratio 2.5–3.1, coriaceous, drying brownish, base shortly attenuate to acute, margin entire, flat, apex bluntly acuminate, upper surface glabrous (to few hairs when young) , glossy, green above, lower surface glabrous to subsericeous when young, paler green, young leaves light green, venation slightly raised above except for the slightly sunken midrib, nerves 9–11 per side. Inflorescences terminal, up to 18 cm long, sericeous; bracts up to 0.8 by 0.3 mm, thick, inside glabrous. Flowers 5-merous; calyx c. 1.5 mm deep, glabrous inside, lobes 5, triangular, c. 0.2 by 0.2 mm, apex acute; petals 5, elliptic, 3.1–3.5 by 0.8–1 mm, apex acute, hook-like, bend inwards, tomentellous on both sides, greenish white to white. Staminate flowers c. 4 mm in diam. (young flower); pedicel up to 12 mm long; petals probably spreading horizontally; stamens 10, in two whorls, free, filaments round, hirsute, of two lengths, up to 1.7 mm long, connective with hairy apical appendix up to 0.3 mm long, anthers c. 0.4 by 0.6 mm; pistillode c. 2 by 0.5 mm, hairy. Pistillate flowers c. 2 mm in diam., not fully open, pedicel c. 1. 5 mm long, elongating up to 4.5 mm in fruit; ovary 4- or 5(6)-locular, c. 1.2 by 1.2 mm, ribbed, apically narrowing and with crater-like opening, stigmas only seen as split bands on inside of apical opening. c. 0.4 mm long. Fruits subglobular to ovoid to angular-subglobular, 1.5–3.8 cm high by 2.1–4.5 cm diam., green when unripe, fleshy, woody when dry; wall very thick, 8–17 mm. Seeds flattened, boomerang-like in transverse section with hollow side extrors, c. 7.5 mm long.

    Distribution — Thailand (Peninsular: Narathiwat), Malay Peninsula, Sumatra, Borneo.

    dots = Galearia maingayi Hook.f.; squares = Galearia celebica Koord.

Habitat & Ecology — Primary forest, pole dipterocarp forest, disturbed forest, alluvial zone, sandy peat swamp, belukar; soil: sandy loam, sandstone derived soil, shales, clayey soil; bedrock basalt. Altitude: 20–800 m. Flowering: April, May, July; fruiting: January to November, eaten by animals.

    Vernacular names — Malaysia: Minyak berok. Sumatra: Katjang kajoe, Medang keli.